THE U NI V E R S I T Y O F C H I CA G O PR ES S

C H I CA G O , I L L I N O I S

T HE BA K ER 8: T A Y LO R CO M PA NY
N E W YO R K

T H E CA M B RI DG E U NI V E R S I T Y P R E S S
L ND NO O

THE M A R U Z E N - K A B U S H I K I - K A I S HA
T O KY O , O S A A
K ,
KYOT O , F U K U O K A ,
S E NDA I

T H E C O M M E RCI A L P R E S S , L I M I TE D
SH AN GHA I
 A L A B O R A T O RY MA N U A L
FOR

C O M PA R A T IVE V E R T E B R A T E
A N A T O MY

By

L I B B I E HE NR IE TTA M AN P H D
, . .

THE UNIVE RS ITY OF CHI CAG O PRE SS

CH I CAG O, I LL INO I S
X)
CO PY R I G H T 1 9 2 2 B Y H E U N I V E R S I T Y O F C H I CA G O
T

A L L RI G HT S RE SE V DR EP U B LI S H E D FE B RUA Y 1 2 2
.
9 R

N I N E T E E N T H I M P E S I O N F E B R U A Y 1 93 8
R S R

C O MP O S E D A ND P RI NT E D B Y T H E U NI VE R S ITY O F C H I C A G O P R E S S
 P RE FACE
0

S everal years ago the m e thod of p ro c edu re in the labora tory work in ver te
bra te zo Olo gy in this Universi ty was chan ged from the typ e p lan then in common ,

use ,
to the c o mp ara tive p lan NO do u b t a similar cha n ge has be en m ade in
.

m an y o ther in s ti tu tions . A s u i table m anu al for the co mp ara tive m e thod has ,

however hi ther to been la cki n g ; the p resen t pu bli ca tion a ttemp ts to su p ply this
,

n eed There can s car cely be an y qu es tion of the sup eriori ty of the comp ara tive
.

m e thod of s tu dy of ver tebra te ana tom y for by this m e thod the s tu den t n o t
,

o n ly learn s all of the an a tomi cal fa cts brou gh t ou t by the typ e m e thod b u t he
also a c qu ires an u nders tandin g of ver tebra te and hu m an s tru c tu re whi ch he is
q u i te u n able to a ttain by the older m e thod of s tu dy In view of the fa c t tha t .

the m ajori ty of the s tu de n ts taki n g co u rses in ver tebra te ana to m y a t the p resen t
ti m e are p rep ari n g for m edi ci n e it seem s obli ga tory tha t they be ta u gh t the
,

his tory of the hu m an body as revealed by the s tu dy of the a n a to my of ver te

bra tes On the o ther hand the comp ara tive m e thod is p erhap s fau l ty in tha t
.
,

it m ay n o t give the s tu den t a clear cu t p i c tu re of the chara c teris ti cs of the

-

variou s classes of ver tebra tes Thu s while the s tu den t readily learns the his tory
.
,

of the aor ti c ar ches for ex am p le he do es n o t readily asso cia te any p ar ticu lar
, ,

g ro u p of ver t ebra t es wi t h a p ar ti c u lar c o n di tio n of the ar c hes when

,
t he co m
p a r a tiv e m e t hod is followed . This defe c t sho u ld p robably be re m edi ed in the

le c tu re p ar t of the c ou rse I have a ttemp ted to rem edy it to a sli gh t ex ten t by

.

in trodu ci n g the se c tion on the general fea tu res of typi cal chorda tes .

I n this m a n u al I have a t tem p ted n o t o n ly to give the labora tory dire c tions
for the disse c tion of the variou s sys tem s b u t I h ave also p resen ted in c onne c tio n
,

wi th ea ch sys tem a very brief ge n erali z ed and simp lied a cco u n t of the develop
, ,

men t and evo l u tion of tha t sys te m I t has seem ed to m e essen tial tha t s u c h an
.

a cco u n t p re c ede or a cco mp an y the labora tory dire c tio n s in order to give a sign i
ca n c e to the fa c ts revealed by the disse c tio n a t the ti m e when the s tu den t

be com es aware of those fa c ts As the c ons u l ta tion of o th er tex ts d u rin g a dissec

.

tion is i n co n ve n ien t a n d ti m e c o n s u mi n g I have tho u gh t it the m os t p ra c ti cal

-
,

p lan to i n c l u de s u c h e x p la n a tory m a t ter in the labora tory m anu al S m

i p le
.
_

illu s tra tions h ave been added to c larify fu r ther the exp la n a tio n s I have n o t .
,

however in the leas t i n ten ded tha t th ese exp lan a tio n s shou ld take the p la ce of
,

readi n g in the s tan dard tex ts of comp ara tive an a to my The s tu den ts sho u ld .

u n d ers ta n d tha t addi tional o u tside readi n g is exp e c ted of the m .

In m akin g su c h brief an d gen eraliz ed exp la n a tio n s as are given here it is

impossible to take i n to a cco u n t nu m ero u s ex c ep tion s an d varia tions I mu s t .

therefore ask the i n d u l gen c e of the exp er t in ver tebra te a n a to m y for the o missio n
of qu alifying clau ses in the exp lana tory a ccou n ts of the variou s sys tem s ; in
V

3 7 54 6
Vi P RE FACE

som e cases no do u b t I have been u n aw are of the ex cep tio n s ; in o thers I have
, ,

kn owi n gly om i tted them on the grou n ds tha t s ta tem m ts of ex c ep tion s are more
c o n fu si ng tha n i n form a tive to the b egin n er .

To avoid con fu sion the exp lan a tory m a tter is p ri n ted in sligh tl y sm aller
typ e tha n the dire c tio n s for the disse c tio n s .

I have in clu ded in the m an u al su ch m a terials as seem ed to m e to bear mos t

directly o n the s tory of the evolu tion of the variou s sys tems I have trea ted .

the skele ton an d the coelo m a t grea t er len g th than is c o mm o n ly the c ase The .

p revaili n g n egle c t of the s tu dy of the skele ton in c o u rses in ver tebra t e a n a to m y

s eem s to me u n j u s tiable in view of the im p or tan c e of this sys tem in the s tu dy of

p hylo g e n e ti c a n d evol u tio n ary qu es tio n s
. I t is t r u e tha t skele tal m a terial is

som ewha t exp en sive to pu r chase an d m ain tain b u t a good man y of the more,

im p or tan t m a terials can be p rep ared by the in s tru c tor or s tu den ts In p resen tin g .

the ver tebral c olu m n I have adop ted Ga do w s con c ep tion of the develop m en t

of the ver tebrae from sep ara te ar cu alia an idea also adop ted by Schau in slan d in
,

his a cco u n t of the developmen t of the ver tebrae in Her twig s Han db u ch The
.

c o n c ep tio n wo u ld app ear to be c orre c t in the m ain and a t leas t fu rnishes a simp le
,

exp lan a tio n for the varie ty of ver tebral c olu mns encou n tered among ver tebra tes .

T he difcu l ties a ttendin g the s tu dy of the c oelo m and m esen teries do n o t seem
to m e to j u s tify one in disregardin g them . I hop e the simp lied a cc ou n t I have
p rese n t ed af t er lo n g s tu dy an d tho u gh t o n the m a tter will ai d in the u n ders ta n d

in g of this co mp lex su bje c t . As to the an i mals to be disse c ted the elas mobran ch ,

mu s t n a tu rally form the po in t of dep ar tu re in the s tu dy of co mp ara tive an a to m y .

I have des c ribed the ska te in addi tion to the do g sh si n c e so m e tea chers p refer ,

it ; it is c er tain ly more favorable than the do gsh for the s tu dy of the nervo u s
a n d u ro g en i tal sys tem s b u t less favorabl e in my op i n io n
, ,
for the s tu dy of the ,

cir cu la tio n
. The n o t in frequ en t s car ci ty of dog sh in r ecen t years m akes it
desirable tha t an al tern a tive form be des c ribed The bony sh is omi tted be ca u se
.

its sp ecializ ed s t ru c tu re wo u ld co n fu se ra th er tha n aid the s tu den t in follo win g

o u t the evol u tio n of the sys tem s . The fro g is so of ten u sed in gen eral and be gi n
ni n g cou rses tha t it seem s su p eru ou s to co n sider it here F u r ther the u rodeles .
,

serve o u r pu rp ose mu ch be tter I sho u ld like to have in clu ded Cryptob r a nchu s
.

as al tern a tive to Nectu m s b u t the limi ts of sp a c e forbade The r ep tile is imp or

, .

ta n t for the p u rp oses of a c o m p ara tive c o u rse a n d the tu r tle is p erhap s the m os t
,

readily ob tain able form of su f cien t siz e The bird has b een i n clu ded sin c e it
.

seem ed i n advisable to omi t al to ge ther an en tire ver tebra te class I have de .

s cribed bo th the cat an d rabbi t as the form er an im al thou gh p erhap s p referabl e
, ,

in so m e resp e c ts is n o t always readily ob tai n ab le

, .

At the Universi ty of Chi ca g o the c o u rse c o n sis ts of a brief glan ce a t the

ex ter n al f ea tu res of the lower c horda tes and O fr e r es en t a tive ver t ebra tes of all of
p ;
the work given in the m a n u al on the ex oskele ton endoskele ton an d m u s c les , , ,

wi th the ex c ep tio n of the mu s cles of Nec tu ru s ; an d of the co m p le te diss ec tion

 PRE FACE Vii

of the dogsh tu r tle an d m amm al ex cep t the perip h eral dis trib u tion of the
, , ,

c ranial nerves in the m am m al The elas m obran ch u rodele an d mammal wou ld

.
, ,

mak e a c om bi n a tion n early as desirable .

The gen eral s ta t ement s and exp lanat ions given in the man u al have been
t aken from s t andard works and ori ginal p ap ers on c omp ara t ive anat om y vert e ,

brat e z o ology and emb ryology and hum an anat omy I have adop t ed for the
, .
,

mos t p art t he u s u al a cc ou n t s of t he evolu t ion of the variou s sys t ems and p art s

, ,

n o t re gardin g myself as c omp e t en t t o c ri t i c iz e t hem I n a few c ases I have pr e

.

s en t ed som e r e cen t views app earin g in the lit eratu re The laborat ory dire ctions .

an d des c rip t io n s of t he anat om y of t he several anim als have however been , ,

wri tt en ent irely from t he s p e cim ens P ra ct i c ally t he whole of the disse ction
.

has b een p erformed twi c e some of it more t han t wi c e The disse ctions have been
,
.

c arried on simu l t aneou sly wi t h the wri t in g of t he dire c t ions In lo ca tin g and .

nam in g the s t r uctu res I have been assis t ed by va ri ou s t ex t s man u als and ori gin al , ,

a
p p ers . In a n u m ber of c ases I have fo un d it desirable t o devise ad ditional
n am es or t o c han g e old na mes I ha Ve emp loy ed t he t erm s dorsal ven t ral
.
, ,

an t erior an d p os t erior as t hey are u sed in vert ebra t e an at om y abandon in g the

, ,

u sa g e c o mmon in h u man an a t om y This has in volved a chan g e in some fam iliar

.

n am es s u c h as t ha t of t he
,
an t erior abdominal vein
.

I have made a p arti cu lar e ort t o u s e all t e chn i cal words in a very p re cise
an d ex a c t manner and t o den e ea c h s u c h word where it rs t o c cu rs On e is .

c on t in u a l ly s u rp ris e d an d ann oyed in a s tu dy of ver t ebra t e s t ru ctu re t o no t e

the loose and ine x a ct way in whi ch many t erm s are em p loyed I t is in fa c t .
, ,

p ra c t i c ally im p ossible t o nd any p re c ise u sa g e for m any t e rm s s u c h as u r et hra , ,

b u lb u s art erios u s p e ri t on eum olfa ct ory lobe p erin eum and o t hers
, , ,
In s u c h
,
.

c as e s I have been c omp elled t o adop t s u c h a de n i t ion as seems c onsis t en t wi th

t he majo ri t y of t he ana t omi c al fa ct s .

I have a tt emp t ed no t hin g in t his manu al b u t a p lain a c c ou n t of the ana t om y

of the several an imals whic h a c c oun t the s tu dent follows This verica tion
,
.

typ e of labora t o r y man u al has b e en re c en t ly s u bj e ct ed to mu ch c ri ti cis m mu ch ,

of it in my op inion u ndeserved The c riti c s seem t o forget that the s tu den t is

, ,
.

n o t in real it y en ga ge d in verifyin g the s t at ement in the m an u al ; he is en gaged

s

in learnin g t he an at omy of an anim al by the short es t an d easies t rou t e a rou te

whi ch the c riti c s thems elves woul d foll ow if c onfront ed wi t h an anim al with
which t h ey were n ot familiar I t is my op in ion that hum an beings in gen eral
.

see chiey t hat whi ch is p oin t ed ou t t o them ; t his has been p roved over and over
again in the his t ory of biology The large number and c omp lexi ty of t he
.

anat omi c al fa ct s t o b e a c qu ired the limit ed time all owed for th eir a cqu isition
, ,

the larg e siz e of the c lasses and t he l imi t ed n u mber of labora t ory assis t an t s
,

available seem to me to ne cessita t e that detailed and s pe cic laborat ory dir ec
tion s be p rovided .I fthe dire ctions are n ot given in t he man u al t hen the labora ,

t ory ins t ru ct ors are c om p el led to p rovide them v e rball y P erson ally I am unabl e
.
 P RE FACE

to s ee any p edagogi cal differen c e bet w een di re ction s and exp lan a tion s writ t en
in the m an u al and t hose giv en ou t v e rbally by the ins t u ct or b u t I do see a grea t ,

deal of dieren ce as c on c erns the time p at i en c e and en ergy of in s t ru ct ors and, ,

s tu den t s Ou r exp erien c e with labora t ory m anu als of the t yp e in whi ch t he
.

b u rden of dis cover y is left t o the s tu den t is that t he s tu dent be c omes hi ghl y
dissatis ed and that the ins t ru ct ors are b rou ght int o a s ta t e of irri tat ion and
fatigu e by the c on tinu ou s d em an ds for assis ta n c e wi t h whi c h they are bombarded .

F rankly I believe in the c onserva t ion of ins t ru ct ors and have wri t t en t his man u al
, ,

wi th that en d in view In p la ce of insert ing qu es tion s in the labora t ory man u al

.

o u r m et hod is t o hold t horo u g h oral qui zz es on the labora t o r y an d t ex t book work

a t fre qu en t in t ervals .

Al thou gh a number of drawin gs are c all ed for in the m a n u al it is p robable ,

tha t ea ch ins t ru ct or will p refer t o de c ide for himself wha t drawin g s a r e t o be

made Drawin gs mi ght p rot ably be omi t t ed al t o g ether t heir p la c e b ein g
.
,

ta ken by ri gorou s p ra cti c al individu al qu izz es on the di sse c t ed s p e c im en .

I am indeb t ed t o a n umber of au t hors and p u blishers for p er mission t o

rep rodu c e gu res from their pu bli cations Du e a cknowledgm en t is made in .

the le g ends t o these gu res I have n o t lis t ed t he n um ero u s o ri gin al p ap ers t o

.

whi ch I have referred sin c e mos t of t hem are given in the bibliograp hies app ended
,

to Kin gsley s Compar a tive An a tomy of Ver teb r a tes and G oodri c h s a cc ou n t of the

shes in P art IXrs t fas ci cle of L ankest er s A Tr eatise on Z oal ogy I am in '

, , .

deb t ed t o D r C R M oore D r B H Will ier an d D r J W B u chan an for c allin g

. . .
,
. . .
,
. . .

my a tt en tion t o errors and omission s in the rs t draf t of the man u al whi c h has
been u sed in the labora t ory u nd er t heir dire ct ion d u rin g the p as t two years .

My t han ks are also d u e t o M r K en ji Toda for his p a tie n ce and skill in drawin g
.

the ill u s tra tions F in ally I wish t o a ckn owled ge the fa c t tha t the p revio u s
.
,

organ iz ation of the laborat ory work alon g c omp ara tive lin es chi ey t hrou gh ,

the eo rts of Dr J W M a cAr thu r an d M r J G S in c lair has fa c ili ta t ed the

. . . . . .
,

task of p rep arin g this m anu al .

S O labo riou s has been this ta sk an d so g rea t is the number of fa cts t o be con
s id er ed t ha t I c an s c ar c ely hop e t o have avoided errors o m issions and s ta t e , ,

men ts la ckin g in c lari t y I shall be m ore than g ra t ef ul t o hav e my a tt en tio n

.

c alled t o t h em .

L . H H YMA N
.

UNI VE R S ITY O F CHI CA GO

No v em b e r , 1 9 2 1
 G E NE RAL D IRE CTI O NS
S u ppl ie s

1 . D is s ec tin g in s tru men ts n eces sar y for the cou rs e a re :

Mediu m siz ed -
s cal p el .

Fin e s c isso r s .

Co ars e sc is s o r s .

S tou t p obe for dissec ti g

r n .

Lon g sl en der p rob e fo r p rob in g .

Medium sized fo rcep s with s t aight p oin ts

-
r .

A towel an d a l abo a to ry co at o r go wn to p ro tect the cl o thes ar e des i abl e

r r .

B on e s cis so rs o r fo rcep s will b e p o vided in the l abo a to ry r r .

2 Drawin g ma terial s n ecessary for the cou r s e ar e :

.

Drawin g p ap er No 6 This p ap er mu st b e s ti an d hard an d have a smoo th su rfac

, . . e .

Hard d rawi g p en cil 6 o r 8 H n , .

E r aser .

Ru l er .

Red , yellow an d bl u e p e cil s ,

n .

P ad o f emery p ap er to sha rp en the hard p en cil .

3 Ob tain the s u pp l ies amed ab ove an d p r esen t you r sel f with a co mp l ete ou t t a t th
. n e

rs t l abora to ry p erio d Do n o t han dicap you rs el f at the s tart by n egl ectin g to p rovide
.

you rsel f with the n ecess ry material s a .

D rawin g s
1 All d awi g s mu s t b e m ade with a ha d p e cil o n g ood qu al ity d awi g p ap er u nl es s
. r n r n r n ,

o therwis e s p eci d Col o rs a e to b e u sed o ly whe s p eci ed in the dir ec tion s

e . r Shadin g n n .
,

c o s sha tchi g e t c
r a e u d es i abl e a d a re to b e avo ided
n ,
.
,
r n D awin g s made o therwis e than
r n . r

a here s p eci ed w il l n o t b e accep ted

s .

2 Drawi gs a r e to b e lin e d rawi g s on ly that is o ly the ou tl in es of the s tru ctu es a e

. n n , ,
n r r

to b e d rawn E very lin e mu s t rep rese t a s tru c tu re a c tu all y p res en t on the s p ecimen
. n Lin es .

mu s t b e s moo th a d cl e Co rr ec t p ro po r tio n s a re o f the u tmos t imp o tan ce an d a e to b e

n an . r r

ob ta in ed by u s e o f a ru l er I n mak i g a d rawi g it is b es t to ou tl in e the d rawin g rs t with

. n n

very light l i es co ec tin g thes e u n til ac cu ra t e a pp ea an ce an d p o p o tion s a e ob ta i ed

n ,
rr r r r r n .

The erase the l ight l i es u til they are barely visibl e an d go over them with a well sharp e ed
n n n -
n

p e cil mak i g the al lin es m an d cl ear

n ,
n n r .

3 D ra w i
.
g s ar e n o t to bn e d ia g a mm a t i ed u l es s s o di r e c t ed in t h e m a u
r a l M a n y z n n .

s t d e ts do
u n o t s eem to un d e s ta d the dif
n f e ce betw ee a diag am a d a d awin g Ar n er n n r n r . s

tion F igu re 9 p ag e 1 5 in K in g s l ey s Compa a tive A atomy of Ver t br a tes is a d ia g am

an ill u s t ra , , ,
r n e ,
r .

a d Fig u e 4 5 p ag e 5
n r
is a d awin g o f the u pp er hal f o f the s ame s tru c tu re r ep resen ted in the
,
1 ,
r s

di g am
a r T he l tte sho w s wha t the o bj ec t a c tu all y l oo k s l ik e ; the fo mer is fo r p u rp o s e s o f
. a r r

e pl
x a tio
an on l y n .

4 A ll d r.a w i g s m us t b e m a d e d ir
n c tl y f o m th e b j e ct w ith t h e b j ec t b e f o r e th ee s t u d e t
r o o n

a d co mp l eted i
n the l abora tory The mak in g of ro u gh s k etches in the l abo a to y to b
n .
r r e

imp ro ved el sewher e is un s cie ti c in accu rate an d abso l u tely n o t p ermitted

n , ,
.

5 R e m e.m b e r t h a t t he p r im e r eq u is it e o f a d r aw in g is a ccu r a cy A d a w in g i s fo r the .

r

in s tr uc to r a reco d f what yo u have a c tu al ly seen Up on you r Sp ec im en If yo u hav e o t

r o . n

ix
 G E NE RAL DIRE CTI ONS

d is sec ted the s tru c tu res ca ll ed fo r , then it is obvious tha t you can n o t d ra w them accura tel y.

P oo r l a bo ra tory wo k in variably re ects its el f in

r the qu al ity of the d rawin gs .

6 . Drawin g s mu s t co n tain al l o f the d eta il s m en tion ed irP the man u al . If, a fter hon es t
eo rt an d with the aid o fthe l a bora to ry a ssis ta n ts , you a re u n ab l e to id en ti fy cer tain r
s t u c tu r es ,

omit them from the d rawin g an d mak e a n o te to hat you we e u abl e to n d them the eec t t r n .

An u n ea so abl amo t o f time s hou ld o t b e s p e t in l o ca ti g s m all o r u imp o tan t details

r n e un n n n n r .

7 A ll d a
. w in g s m s t b er th o ou g h l y l a b e l ed
u E v e ry d a w i gr m s t b e c o mp l et el y l ab el ed . r n u

rega dl ess f whether the s ame s tru ctu es have al r eady b een l abel ed in some p ecedin g draw
r o r r

in g L abel s a e to b e writ ten o r p rin t ed in har d p e cil p a all el to the to p an d bo ttom edges o f
. r n r

the p ag e a d l in es d awn with a rul er f r om the l ab el s to the p ar ts in dica ted

n r .

8 Draw on the ig ht ha d s u r f ce o f the p ag e o n ly

. r - n a .

9 R e m em
. b t h t t h e l a b o a t o
erry i s t a c t o r s ar e f a m il ia r writh a l l o f t he g
n u r es in
ru the
v rio s t e tb oo k s a d that u d u e es embl a ce b etween you r d awin g s an d s u ch g u es will
a u x n n r n r r

re c t u p o
e y o h es t
ny a d uri e a s s p
onic i o t h a t y ou
n h av e
ra s n o t b ee e er ti g yo u r sel f in the
u n n x n

l bo rato ry
a .

0 T he d awi g s will b e c ll ed in a t in terval s

1 . r n T he da tes on which they are du e wil l b e
a .

an n ou ced in a d va ce b y the l a b o
n to ry in s tru c tors
n ra .

No te s an d Qu i zz e s
1 . N0 qu ir ed in this cou rs e T he n o teb oo k s will con s is t o f d rawin g s onl y
n o tes a r e r e . .

2 O ral a n d w ritten qu izz es u p on the s u bj ec t m a tt er o f the l abo r a to ry wo r k a re to b e

.
-

ex p ec t ed a t an y tim e These qu iz zes will d eal with the an atomy o fthe an im al s you are dis sect
.

in g an d with comp a ra tive an a tomy Y ou will b e ex p ec ted to kn ow tho rou g hly the an ima l s .

an dma terials which you dis sec t an d s tu dy in the l abo ra to ry , an d to b e abl e to co m p ar e them
with on e an o ther , s ta tin g r es embl an c es an d differ en ces You wil l b e r equ ir ed to exhib it you r .

dis s ec t ion s an d to be a bl e to ide tify the s tn ru c tu r es p rese n t on the diss ec tion s .

3 An im o r tan t qu iz will
. p foll ow the co m pl etion o f ea ch s ec tion o f the l a bo ra to r y wo k r

an d will d eal wit tha t sec tio n h .

4 . R eadin g in the tex tb ooks of co m pa ra tive an a to my is ex p ec ted as a pa rt of the l a o r a b

to r y work .
Quiz zes will in cl u de ma terial in su c h te tbook s
x .

D is se c tio n
1 . Disse c t ion do es n ot co n s is t in cu t t in g an an i mal to pieces Dissection co s ists
. n in
se p ar a tin g the p a r ts o f an an i ma l so tha t t hey are mor e cl ea r ly vi ibl e l e vi g the p ar ts
s ,
a n as

in ta ct as p r ac tica b l e .

2. I n d is s ec tin g a n an i m al v ery lit tl e cu ttin g is re qu ired . Cl ean in g aw ay the c on n ec tive

tissu e w hich bi ds tog ther a n e nd c on ceal s s t ru c tu res is the c hief p roc ess in d issection . In
do in g this u se bl n t in s t m e
,
u ru n ts , as p robe fo cep s or n g ers Avoid the u se of s ca lp el
the ,
r , .

an d s ciss o rs . Y ou will p rob a b l y cu t so m ethin g you will n eed l a ter o n

'

I n sho r t do n o t cu t ; .
,

s e par at e the p art s .

3 . Have the an i m al rmly fasten ed . Small er an i mal s ar e g en era ll y pi n n ed to wax

bo ttomed dis s ec tin g pa n s. L arg er an i mals , su c h as are us ed i n the g r ea ter p a rt o f h
t is r
cou s e .

ar e tied to s cr ew e ye s in the co rn ers o f the diss ec tin g p an . P u t the pa r t icu l a r pa rt you a r e

diss ectin g on a s tr et ch.

4 you r own dis secti g Do n o t watch somebody el se do it B egin a t the mo t

. Do n . . s

ea s il y access ibl e po in t o f the s ys tem you a e s tu dyin g a d fo ll ow o u t your s t ru c tu e cl ean i g r n r ,

n

away the tis su es that c o ceal it n .

5 E ercise pa tien ce an d ca re
. x Cl ean the s tru ctu es b y small p or tion s . r .
 G E NE RAL DIRE CT I ONS Xi

6 . Fo ll o w the dir ec tion s precis ely . Do not cu t a n ythin g or re mo v e an ythin g u n l es s

s p eci cally di ec ted to do so r .

7 Y .o u r l abo a to ry g a d e is p a rt
rly d et e m inred by the kin d o f diss r ec tio n s you ma k e .

Material s
1 . Bu t on e s pecime of each a imal is all owed to each s tu de t
n n n . E ach s tu den t wil l b e
g iv en the n eces sa ry s p ec im en s an d w il l etain them to the e d o f the c o
r n u r s e. Do n ot dis ca rd
an y an im al u n t il man u al s o direc ts
the .

2 T he s mall er m a terials which are p ro vided fo r the cl ass as a whol e sho ul d b e r etu rn ed
.

to the b o t tl es o r j ar s f ro m which th ey c ame as s oo n as you ha v e n ished s tu dyin g them .

3 T h
. e l a rg er s p ec im en s w ill b e k e p t in l a rg e ca n s E ach tab l e w ill b e al l o tt ed th
.e n ec e s sa ry
n u mb er o f can s S tu d en ts will a tta ch tag s b ea rin g their n a mes to their s p ecim en s an d
.

k eep them in the can s when they are n o t in u se .

4 S p.ec im en s m u s t al w ay s b e k ep t m o is t an d mu s t n ev e r b e al l o wed to d ry u p , as this

ru in s them for d is s ec tion Do n o t go away an d l ea ve you r s p ecimen s ou t o n the tab l e When
. .

rea dy to l ea v e the l ab o rato ry , wra p the an imal s in mo is t en ed chees ecl o th p r o vid ed in the l a b o r a

to ry an d p u t them in to the ca n s S ee that the can s are al ways cov er ed

. .

5 S tu den ts who , throu g h their o wn ca r el essn es s , ren der their sp ecimen s u n t fo r fu r ther
.

dis s ec tion will hav e to p ay fo r n ew s pecimen s .

6 T he s k el etal ma terial p r o vided in this cou r s e is exp en s ive

. Han dl e it with ca re B e . .

p a r tic u l a rl y ca efu l with s k el eto

r ns p reserved in u id .

R efe ren ce s

Th o ghou t this man u al referen ce is made to a nu mber of te ts a d m al s which may

r u x n an u

p o tabl y b e co s l ted by the s t den t They are ob tai ab l e in the l ib ary Th e ef e e ces
r n u u . n r . es r r n

ar e in dica ted as follows :

B B en sl ey ,
P ractical An atomy of the Ra b bit .

CN H The Ca mbr idge Natu ral His tor y: Vol u me VI I , F ishes As cidian s , etc ; Vo l u me VIII , .
,

Amp hibia an d R ep til es

.

D Dav is on , M ammal ia n An a tomy with S pecial Refer en ce to the Ca t .

H Her twig Man u a l of Z ool ogy t ran sl a ted by ed 191 2 . .
, , ,

K K in g s l ey , Compa r ative An a tomy of Ver tebr ates , 3 d cd , 1 9 2 6
X
. .

L L an k es ter , A Tr ea tise on Z ool ogy P a r t I , rs t fasc icl e

Cyc l o s to m es an d Fis hes

.

.
,

N N wm e Ver tebr a te Zool ogy

an , .

P an d H P a r k er an d Ha swell A Tex tb ook of Z ool ogy, Vol u me I I , 3 d cd

,
.
, 1 921 .

R R eyn o ld s , The Ver tebr a te S kel eton .

R an d J R eigha rd an d J en n in g s , An a tomy of the Ca t .

W Wild er The His tor y of the Hu man B ody
,
.

Wd Compara tive An atomy of Ver tebr ates .

 TABLE O F CO NTE NT S
P AG E

I . G E NE RAL CONS I D E RATI ONS ON ANI MA L F OR M 1

A Descr ip tive
. Te m r s

B P l an es an d Axes
C Symm etry
.

D Metamer is m
. or Seg men ta tio n
E . Cephal iz ation 3
F . Homology an d An al o gy 3
II . T HE P HY L UM CH ORD ATA 5

A T he C
. h r cte is tics of the Cho r d tes
a a r a 5
B The Charac te is tics o f the Ve teb a t s
. r r r e 5
C T he Cl ass i cation o f the Cho d tes
. r a 5
III . G E NE RA L STUD Y OF TYP I CAL CH ORDATE S
A A mphioxu s
.

B . A Tu n ica te
O B al an ogl oss us
U An a to my of a L am p rey

mE y of the Dogs h
x te rn a l An atom
mE tern l An atomy o f t he S k a te
x a

o E te n al An atomy o f a Tel eos t

x r

m So me Gan oid Fishes

H E t n l An ato my o f N ctu u s
x er a e r

H E x t e r n al An atomy of a L izar d
W E x tern al An a to my o f the Tur tl e
E y o f the Pigeon
E x te rn al An a to m
E E tern al An ato my o f a M ammal
x

Z S u mmary

IV . G E NE RAL FE AT URE S OF CH ORD ATE DE VE L OP ME NT

> The C ho r d a te E gg
w vage of the E gg an d the Fo
T he Cl ea r ma tion o f the B l as tu l a
o Forma tion o f the G as tru l a
c Fo ma tion of the Third G m L ayer
r er ,
the Neu r a l Tube ,
an d the No to chord
m Fu rther His tory of the Meso derm
w The Fa te o f t he E c toderm
o T he Fa te o f the E n toderm
m The Fate of the Mes o derm an d the Formation of Mesen chyme
V THE COMPARATI VE ANAT O MY
. OF THE INTE GU ME NT AND THE EXOS KE L E T ON

A . G en eral Cons ide ation s on the S k el eto n

r

B . T he S tru ctu r e o f the S k in

C . The E os k el eton in G n e al
x e r

D E xos keleton
. o f Fis hes

E . E xos kel eto n of A m phib ia

F E xo s k eleton of Re p til es

xiii
xiv T AB LE O F CONT E N S T
P A GE

G . E xo s k el eto n o f B ird s 52
H E x o s k el e to n
. of M a mm a ls 54
1 S u m ma ry
. 55
Q

VI THE E ND OS KE L E TON :
. T HE CO MP AR ATI VE ANAT O MY O F T HE VE RTE B R AL COL U MN
AND RI B S
A . Ge n er a l Co n s idera tio n s on the E n do s k el eton
B T he E m b ryo n ic Or ig in o f the Ver te r ae b an d Ri bs
C S om e P r imiti e Ver te r al Co l u mn s
. v b
D The Ver teb ral Co l u mn o f t he D og s h
.

E . Ver teb r a l Co l umn of Teleo s ts

F . Ver teb r al Co l u mn of Am phibia

G . Ver teb ral Co l u mn of R e p til es

H . Ver teb ral Co l umn o f B ir ds

I . Ver teb ral Co l umn of M ammal s

J . S um mary o f the Ver tebra l Co l u mn an d R ib s

VI I THE E ND OS KE L E T ON :
. TH E COMP AR ATI VE AN AT O M Y OF TH E G I RD LE S , THE

S TE RNU M ,
AN D THE P AI R E D AP P E ND AGE S

A . G en er al Co ns ider a tio n s
B . T he P el vicG ir dl e d the P os te ior P air ed App en d ges
an r a

C . T he P c to al G i dl e the S tern um an d the An te ior P ai ed App e d ag es

e r r , ,
r r n

D . G e al Su m mary o f the G i dles the S t n um an d the P i ed App e dag

en r r ,
er ,
a r n es

VI I I T HE E ND O S K E L E T ON :
. THE CO MP ARATI VE ANATO M Y OF THE S K ULL AND THE

VI S CE RA L S K E L E T O N
> T he Car til a g e S t age o f the S k ull
H The Vis cera l S k el eton
O T he Fo r m a tio n o f the Memb r an e B on es o f the S k u ll
P T he Fo rma t io n of the Car til age B o n es of the S k ull an d the Com p ositio n of

the Co m lete S p k ull 103

e T he S kul l of Nectur us , a y Oss ied Sk ull

P ar tiall 10 7
w T he S kull o f t he Al l ig a tor , a Typ ic l Mo d e n S k ull
a r 1 12

G Ma mmal ian S k ul l
i

. T he B o n es o f the 1 16

H . Ge n er a l S u mma ry o f the S kull an d Vis c er a l S k el eton 1 26

I X TH . E CO MP ARATI VE ANAT O MY OF THE M U S CU L A R S Y S TE M 1 28

A . Ge n er al Con s ider a tion s 1 28

B . Muscles
The o f the Do gs h 1 28

C T he Mus cl es
. of Nectu rus 1 30

D T he Mu s cl es
. o f the Ca t an d R a b b it I33

E . S umma r y I S7

X . THE CO MPARATIVE ANATOM Y O F THE COE L OM , DI GE S TI VE , AND R E S P I R ATO R Y

S Y S TE M S 1 58

A T he Or ig in an d P ar ts o f the Co el o m an d the Mes en teries 1 58

B T he Diges tive Tr ac t an d I ts Der iva tives
.
159
C The Co el o m , Diges tive , a n d R es p ir a to ry
. S ys tems o f E l as mo b r an c hs 163

D The Coel om Dig es tive ,

.
, an d p
R es irato r y Sys tems o f Nect us ur 168

E T he Co el om, Diges tive ,

. an d R es p ira to ry Sys tems of t h T tl e ur e 1 72
F The Co el o m D ig es tive an d Res p ir to ry S ys tems o f t he P i eo
a
.
, , g n
 TABL E OF CONT E N S T X
V

P AGE

p
G The Co el o m , Diges tive , an d R es ir a to r Sys tems o f a Ma mmal
. y 1 83

. p
H T he Com ar a tive An ato my o f the Co el o m an d the Mes en ter ies I O4
I . Summa ry 1 98

XI . THE CO MPAR ATI VE ANAT O MY OF TH E CI R CUL AT OR Y S Y S TE M

20 0

A . G en eral Cons ider a t io n s 20 0

B . ry Sys tem o f E l as mob a n chs

T he Cir cu l a to r 20 7
C T he Cir cu l a to ry Sys tem of Nectu s
. ru 222

D T he Cir cul a tory Sys tem o f the Tu r tl e

.
23 0

E T he Circ ul a to ry Sys tem o f the P ig eon

. 24 1

F T he Ci cul to ry Sys tem o f the M mm al

. r a a 2 48

G S u mmary o f the Cir cul ato ry Sys tem

.

XII THE COMPARATI VE ANATOM Y OF TH E UROGE NI TAL SY STE M

. 2 73

A E mbryo ic Origin an d E vo l u tion o f the U og ital Sy t m

. n r en s e 2 73

B T he U roge ital Sys tem o f E l as mob a c hs

. n r n 2 80

C T he U ogen ital Sys tem o f Nect us

. r ur 283

D T he U ro gen ital Sys tem of the Tu tl e

. r 2 84

E T he U ro gen ital Sys tem o f the Pigeon

. 286

F T he Ur ogen ital Sys tem of the M amm al

. 2 88

G The E mbryon ic Membra e

. n s 29 2

H Summary o f the Ur ogen ital Sys tem

. 294

X III THE COMP ARATI VE ANATOMY OF THE NE RVOUS SY S TE M AND THE

. SE NS E OR GAN S 29 6

A Gen er al Con s ider a tion s

. 296

B . ys tem an d Sen se Orga s o f E l smob

T he Nervo u s S h n a ran c s 30 1
C T he Nervo us Sys tem an d S en s e O g s of Necturu s
. r an 3 18
D T he Nervo s Sys tem an d Sen s e O g s of the Tur tl e
. u r an 3 21
E The Nervous Sys tem an d Sen se O g a s of the P ig eon
. r n 3 28
F The Nervous Sys tem an d S ens e O g n s of the Mamm al
. r a 3 33
G Su mm ary o f the Nervous Sy tem a d the Sens e O g ns
. s n r a 3 59

APPE NDI XA P R ONUNCI ATI ON AND DE RI VATI ON OF TE CHNI CAL W OR D S

.
36 2

APPE ND IXB . P RE PARATI ON OF MATE RI ALS 3 7

3 73
 LAB ORA OR T Y MANUAL FOR VE RTE B RA E ANA OM T T Y
symm e trical the o the typ es o f symm etry will n o t b e co
,
r n s id er ed her e . Fo r them co n s u l t H,
pages 3 5 1 2 -
2 .

Bil teral s ym m t y a The p a ts o f a bil ter al ly e r . r a syn me tr ical a n im al ar e arra n ged

sy mm etr icall y with f

r e er en c e to thr ee ax es , the l on gitu din al , tran sver se , an d s ag it tal axes ;

the two e n d s o f the sag it ta l axis in an y give c

n r oss -
s ec tion a r e u nl i e k . Th er e is b u t on e pl a ne

o f symm etry i n s u ch a n an i ma l tha t pl a ne whic h p asses th o gh the l o g it di al a d s agitt l

r u n u n n a

n m ely median divid es the an im al in to a pp o im t l y id e tical

a esx a , the s a gittal pl an e . It r x a e n

right an d l ef t hal ves , w ich h ar e m irr o r imag es o f ea ch o ther . T he s tru c tu r es o f ve r teb ra t es

a r e either cu t in by the media sagittal pl e in which ca se they are sp ok en o f s p i d

hal f n an ,
a un a re

s t ru c t e o they a e pl aced symm etrica ll y o

ur s, r ea ch s id e o f this p l an e e qu idis ta t f o m it
r n ,
n r ,

i which cas e they a e p i d s tru c t es

n The d ig es tive tr ac t is the o nl y sys t em which do es n o t
r a re ur .

exhib it a symm et ica l l tio to the m ed ia p l n e i the adu l t al thou gh it too is bil a ter all y
r re a n n a n , , ,

symmetrical in ea ly emb ryo ic s t g es r n a .

D . ME TAME RI S M OR SE GM E NTATI ON

S egmen tation metamer is m is tha t s tru c tu r al con dition o ccu r rin g in cer ta in g ro u p s o f
or

an i mal s in which all o r mos t o f the pair ed p a r t s o r s tru c tu r es a r e rep ea t ed a t r egu l ar in ter val s

al on g the an tero post erio r ax is . The bo dy o f s u ch an im al s c on s e u en tl yq is co m p osed of a

l on g itu din al s er ies o f d ivis ion s o r el em en ts , in ea ch o f w hich all or mos t of the s ys tems o f the
body a e ep esen ted r r by en ti e p ai ed o g s or s t c tu es o r by a po tio of the medi
r , either r r r an ru r r n an

un p ai ed s tru c tu r es
r E a ch su ch d ivisio o el em en t o f the b o dy is call ed a met mer e somit
. n r a ,
e,

o s egmen t
r T he a t erio r an d p os t io r bo daries o f ea ch s egme t m ay o m ay no t b
. n er un n r e

ma k d e tern all y by a co s tric tio o f the b o dy wall

r e x I n the fo rm er ca se the a imal is s aid
n n . n

to exhib it bo th xt al a d i ter al m e ta me is m ; i the l a t t e cas e in tern al metameris m al o e

e er n n n n r n r n

is p es t r en .

I n a ideal s eg men ted an imal all of the s eg men ts a re id en tical with ea ch o the in all o f
n r

the detail s o f s t c t e No s ch a imal e is t s becau se b o th the head a d the termin al

ru ur . u n x n

seg men ts mu s t o f n ec ess ity di if o nl y slightly f om the o ther s eg men ts b u t the ri g eder , ,
r ,
n

worm s su ch as N r is a d the ea thwo m cl os ely app o a ch the ideal

,
e e n S ch s eg m e ted a imal s
r r ,
r . u n n

in which th va io s s eg m e t s a e ea ly al ik e a re s aid to p o ss es s homon mous seg me ta tio

e r u n r n r o n n.

T he maj o rity o f s eg me ted a im al s ho wever dis p l ay het r on omou s s eg men ta tio

n in which the
n , ,
e n ,

va io u s s gm en ts h v e b ecom e u nl ike ea ch o ther in ma y esp ec ts

r e a n r .

The eg m n ted g o p s o f an im al s a e the a elids the a r th o po ds a d the ve teb a tes

s e r u r nn ,
r ,
n r r

a d th i
n l a tives I the evol u tion o f s egmen ted an imal s th e e has b e
e r re . n a co n tin ou s p o r en u r

g r e s s io n f ro m the ho mon omou s to the ex tr em e hetero n o mo u s co n dition . Ho mon o mou s

s egmen ta tion r ep res en ts a p rimitiv e an d g en er aliz ed h h the va io yp e
t o f s t ru c tu re in w ic r us

seg m en t s a r e m o r e o r l ess in d ep en d en t o f each o ther

a d e ch is ca p ab l e o f p e fo rm in g all n a r

of the ec s s a y f c tio s
n B t w ith the ev l u tio of hete o omy the s eg me ts becom e
e r un n . u o n r n ,
n

un lik e d ther e is a div i io o f l bo r a mo g them so me po tio s of the bo dy el bo ra ti g

an s n a n , r n a n

c r ta in f
e c tio s d o the s o ther fu c tion s
un nE a ch s eg me t is the
an o l ger c p abl e o f
r n . n n n on a

p erfo min g all o f the f c tio s b t is dep e d n t u pon it f ll ow eg m ts with a es l ti g

r un n ,
u n e s e -
s en r u n

un ica tion an d o rg an iza tion which is l a ckin g in the homon o mou s l y s eg men ted fo rm s .

T he he teron omou s co n dition is derived f rom the ho mo n o m ou s throu gh a n u mb er of

d i er en t p ro ces s es , su ch as l oss o f s egmen ts , fu s ion of a dj a cen t s eg m en ts en l a rg em en t , or

men t s lo s s of o rg an s o r p ar ts f rom s ome s eg men ts w ith their r et en tio n in o ther

red u c tion o f s eg ,

segm en ts s tru c tu r a l chan g es amon g the r ep ea ted o r an s o r p a r t s so tha t tho s e o fd iff er en t s e

, g g
men ts b eco me unl ik e etc , .

ver tebrates a e hete o omo ly segmen ted an imal s with in tern al segmen ta tion on l y
T he r r n us .

T he embryos o f ver t eb t m ch more n ea ly app roach the ho mo o mou s co ditio

ra es an u r n n n ,
 GE NE RAL CONSI DE R ATI ONS ON ANI MAL F ORM

in dica tio n tha t the v er te b

f rom ho mon o mou sl y s egmen ted forms I n the embryon ic
r a t es ar o s e .

devel op men t o f ver teb ra t es the chan g e f ro m a s o mewha t homon o mou s con d itio n to an ex tr eme
hetero n o my can b e direc tly fo ll o wed .

E . CE P HA LIZATI ON

In the v
e o l u tion o f an i mal s ther e is a
p ron oun ced te den cy for the an terio e d of the n r n

body to b ecome more an d mo re d is tin c tly

s ep ara ted an d d ie en tia ted f o m the r es t o f the r r

body as a head This di eren tia tion of the head con s is ts chiey of the local i a tion within
. z

the he d oa f the m a in p a t o f th e erv s sys em

ou t the b rain
r an d o f the mos t im p o rtan t
n

se s e o g an s
n r S in ce the b a in an d the s e s e o g an s co n trol to a very l arg e d eg ree the activi
. r n r , ,

ties a d r es po ses o f the res t o f the bo dy the head thu s b ecomes the d omin an t p ar t of the
n n ,

o rg a is m
n This ce t al ization o r loca lization of n ervou s s tru c tu res an d fu n ction s in the head
. n r

with accompa n yi g domi an ce of the head is call ed c phal iz tion

n n Cephalization is mo re e a .

an d m o r e m ar k ed the higher on e as ce d s in the an imal k i g dom an d is p ar ticul arly pro min en t n n ,

as a s tru ctu al a d fu n c tion al fea tu re of the ver teb a tes

r n r .

I n seg men ted an imal s the advan ce in c ephali a tion is co rr el a ted with the prog r ess ion z

o f the hetero o mou s co n ditio n Heteron omy in fact app ear s rs t in the head r egion a d
n. , , n

g ad u ll y p rog es es po s terio ly
r a rThe an terior e d thu s r etai s the l eas t an d the p o s t erio r
s r . n n

e d the mo s t r es embl a ce to the o rig in al homb n omou s con ditio n

n n This res u l ts i an ill . n u

s ion o f a r e t ea t o f c e ta in sys tems to ward the p o s terio r reg ion s o f the bo dy wher ea s the
r r ,

s it a tio
u in r eal ity a rises f rom the fact that thes e sys tems have dis app ear ed from the an terio r
n

s g m e ts an d a e r etain ed in the po s ter io r s eg men ts

e n r I n the case of cer ta in ver teb ra te o rg an s .
,

as the hear t a r ea l p o s terio r descen t occu rs du rin g the evol u tion of the verteb a tes
, In r .

the ve teb ra tes as in o ther h eteron omou s ly seg me ted an im al s the head is p ro d ced throu g h
r ,
n ,
u

th fu s ion o f a c er tain n u mb er of the mo s t an terio r s eg men ts with a l oss of s o me s eg men ts

e

o r o f p a ts o f s eg men ts an d the d is app ear an c e f rom these head seg me ts of n ea ly all sys tems
r n r

e c ep t the n ervou s sys t em

x As cephal iza tion p rog ress es the head app op ia tes mo e a d
. r r r n

mo e o f the adj a cen t s egme ts in corp o r ati g them in to its s tru c tu e so tha t in g e eral it
r n ,
n r ,
n

may b e sa id tha t the higher the degr ee of cephaliz ation the g rea ter is the n u mb e o fs eg men ts ,
r

co mp o s i g the h ea d
n I n a dvan ced cephal iza tion su ch a s is po ss es sed by ve teb a t es it is
.
,
r r ,

very d if cu l t i deed a l mo s t impo s ibl e to decipher the n um b er an d bou da ies o fthe egmen ts
,
n ,
s ,
n r s

which o ig i all y wen t in to the co mpo ition o f the head ; in fac t the p robl em o f the s eg men ta
r n s ,

tion o f the ve teb ra te hea d has n o t b een comp l etely s ol ved al tho gh it has r eceived the a t te
r ,
u n

tion o f the fo emo s t ve teb a te a a tomis ts

r r r n .

T he ve t eb a tes a e then a imal s chara cte i ed by the po s s es s ion o f bil a tera l symmetry
r r r , ,
n r z ,

in tern al an d ma k edly hete o omou s s eg m e ta tion a d a high deg ree o f cephali a tio
r r n The n ,
n z n.

d etail s o f thei s tru c tu re a e u d e s ta dabl e o n ly in the light o f thes e thr ee b o ad an a tomical

r r n r n r

c on ditio n s .

F . H OMOL O GY AND ANAL O GY

Homol ogous h however un l ike in fun ction or s uper cial a pp ear
s tr u c tures are thos e whic ,

an ce , have the s ame origin as d mo s tra ted by the s tu dy o ftheir emb ryonic o rigi an d develop
,
e n n

men t a d thei p al eon to l ogica l his to ry

n r Thus the wing ofbirds the ipp er o fthe s eal s and the .
, ,

fo re l g o f the ca t ar e homol og ou s s tru c tur es becau se they are a ll mo di ca tio n s o fthe o riginal
e

for e l imb an d devel op i the s ame way u p to a cer tain p oin t n The whol im of co mpar tive . e a a

an a t o m y is to dis cover what s tru ctur e s ar e hom o l og ou s an d to tra ce the m o di ca tio n s o f s u ch

ho mo lo g o us s tr u c tu r es in the cou r s e o f evol u tion .

An al ogou s s t ru c tu r es a re tho s e w hich r es emb l e ea ch o t h er either a s to su pe ci

r al a pp ea r

an c e o r in fu n c tio n b u t which ha ve ha d dier en t o rig in s . Thu s , b o th s h an d sn a k es a re

4 L AB OR A OR T Y MANUAL FOR VE R E B RA E ANA T T
c o vered with sca l es fo r p ro tec tive p rpo ses
u hese scal es are o t the s
,
b ut t n

b ecau s e they o rig i a te f om the s k in i d ieren t ways

n r n Su ch fu c ti al . n on

s all y th
u u
e r es u l t o fe vir o mne t al nc o ndi t io ns i e n th ey a e a,
p
. t t
.io ns
,
r

a .

a imal s b u t whol e a im al s m ay c o me to res embl e each o ther th ou g h th

n n r

en vir on men t as whal es a d shes

,
n S u ch r es embl a ce
. o t b as ed on a n ,
n

call ed con ve genc e

r .O the o ther han d animal s cl os el y r el ted to each o t
n ,
a

in app ea an ce o win g to the differen t en vir on men ts in which they live as

r , ,

is k own as diver gence

n ( S ee.f u rth er N pp 1,5 2 0 ;
,
W pp. 1 6 -
) ,
. .
 11 . T H E P H L UM CH O R DATA Y
A . TH E CH ARA CTE RI S TI CS OF THE CH OR DATE S

Whil e p ris e the g rea ter pa t o f the phylu m Ch rd ta th ee s mall

the v er teb r a tes co m r o a ,
r

gro u ps o f a imal s are u n it d with them i this phylu m becau se they po s ses s ce tain cha
n e n r r

a c t e is tics in co mm on with the v e teb a t es

r These cha acte is tics are : r r . r r

I T he wall o f the pha ryn x o f the emb ryo o r adu l t is p ie ced by o p e i g s the gil l s lits
. r n n , ,
.

o rig i ally fo r es p i a to ry pu rp o s es
n r r .

2 A n ot chord is p es e t in embryo o r adu l t

. o T he n o to cho rd is a ro d lyi g do s al to the
r n . n r

in tes tin e e ten din g fr om an te ior to po s terio r e d a d s ervin g as a s k el etal s u ppo t

,
x r I n ,
n r . n

ve tebra tes the n o to cho rd is p ar tiall y o r wholly r ep l aced by the s ku ll a d v e teb al co l u mn

r n r r .

3 T h e .c en t al e rv o u s s ys tem is
r n early al ways h ll w ( i
n the tu n ica tes in th embryo o o n e

o ly )
n co tain i g u su all y a s i g l e co n ti u ou s cavity b u t in s om e cases a n u mb e o f is o l ted
,
n n n n r a

S pa c es an d is s it a ted en ti ely o n the dors l s id e o f the bo dy ( e c ep t i

,
u r the Hemi h d ta a x n c or a

wher e ther e ar e b o th do s al an d v e t al p o tion s) : I n the in ve teb a tes the cen t al n er vo u

r n r r r r ,
r s

sys tem is al ways solid an d lies m ain l y v en tral in the bo dy .

F o r fu r ther d is cu ss ion o f thes e chara c te s see P a d H p ag es 1 2 ; N pages 3 3 ; r n , ,

1- 2

K pages 2
,
1- .

B . THE CHAR A CTE R I S TI CS O F THE VE R TE B RATE S

T he morphol og ical ch racters of the ver tebra tes are the foll owi g : a imal s with bila te al
a n n r

s ymmet y i te al heteron o mou s s eg men ta tio

r ,
n rn an d c ephali a tion ; with g en er ally two p airs o f n, z

paired j oi ted locomo to r app e dages in the form ofn s or l imbs an d sometimes with u pai ed
n n , ,
n r

app e dag es in ad ditio ; S k i s ep a abl e f om the es t of the b ody wall an d co mmo nl y p ro du c i g

n n n r r r n

p o tec tive s tru c t es s u ch as s cal es feathers hai etc cell u l ar in n atu re ; mu scl e l aye o f the
r ur , , ,
r, .
,
r

b o dy w ll d ecidedl y metame ic in arra g eme t ; with a in tern al S k el eto

a o f ca til ag e o
r n n n n, r r

b on e co s is ti g of a s k ll an d g ill su pp o ts in the head ve tebral col u mn ibs an d bre t

,
n n u r ,
r ,
r , as

bo e in the b ody a d su ppo ts fo the app e dages ; ver teb al colu mn highly metame ic
n ,
n r r n r r ,

co m p o s ed of s u cc ess iv e ri g s a rou d the n o to cho d ; cen t al n ervou s sys tem c o s i ti g o f a

n n r r n s n

b ai mu ch e l a g ed within the s ku l l an d a s p in al co d within the v e teb al col mn ; erves

r n ,
n r , ,
r r r u n

highly metameric i a a g em e t ; head with th ee p i s of se se o g a s eyes ea s an d n os e ;

n rr n n r a r n r n , ,
r ,

dig es tive t ac t g ivi g is e b y ou tg o w th to two dig es tive g l a ds the l iver a d the p an cr eas ;
r n r r n ,
n

pharyn x in timately co ected with the resp i a tory sys tem eithe op i g to the e te io
nn r ,
r en n x r r

by o pe i gs the gil l slits in the w ll s o f which the g ill s ar bo e o g ivi g ise b y ou tg owth
n n , ,
a e rn ,
r n r r

to the l u g s ; hear t al ways ve t a l i the b ody ; ci cul a to ry sys tem cl os ed al ways with a median
n n r n r ,

d orsal a r tery the ao ta an d with o e o r two po tal sys tems ; g e ital a d e c eto ry sys tems
,
r ,
n r n n x r

cl osely r el a ted the e c eto ry d cts g en e ally servin g as g e ital du c ts ; e c eto y a d g e ital
,
x r u r n x r r n n

du cts o p e i g in co mmo with the in tes ti e i to a cl oaca o r o p e i g s ep a t ly n ea r the a s ;

n n n n n ,
n n ra e nu

with a well dev l p ed co el o m -

ever s eg me t ed an d divided in the ad l t in to two o r fou r
e o ,
n n ,
u

co mp ar tmen t s ; viscera s u ppo r ted by mes e teries ( S e f he K pp

e u r t r 2 4 ; P an d H n .
, ,
.
,

p 1 1 9 ; H pp 57576 ; W d pp 1 1 1 3 ; a d s tu dy the diag rams in P a d H p

.
, .
,
.
-
n n ,
.

C . THE CL A S S IFI CATI ON O F THE CH ORDATE S

Sin c e in this m an u al f
r e er en c e by t heir scien ti c na mes to g r o u p s of cho r da tes is un a vo id

ab l e . it is ess en tial tha t the s tu d en t l ear n at o n ce the foll owin g s che me o f cl ass i ca tion . On l y
6 L AB ORA OR T Y M ANUAL FOR VE R E B RA E ANA OM T T T Y
thos e g rou p s a e i cl u d d which are ac t all y m et with or r eferr d to in the ma u a l Fo r
r n e u e n .

c omp l et e cl ass i ca tion see the v ariou s t e tb oo k s as N P a d H CNH e tc S p ec i x , ,

n , , .

m e s of these differ e t g rou p s wil l b e seen in the l abo rato ry in the n ext sec tion o f the
n n

man u al .

P hyl u m Chor d ata

Su bphylum I . Cephal ochordata (Acran ia) , A mphioxus an d its allies .

S u bphyl m II u . Urochor data or Tu n icata , the tu n ica tes or s ea s qu ir ts .

S bphylu m I II
u . Hemichordata (E n ter opn eu s ta) , B al an ogl ossus an d its a ll ies .

S u bphyl um I V . Ver tebrata , all an i mal s with

ver teb al col u mn a r .

Cl ass 1 . Cycl os to mata , the cyc l ostomes o r roun d m ou thed shes -

,
shl ike an i mal s w itho u t
l owe r j aws or p ai ed s r n .

Cl as s 2 P is ces the tru e shes with j aws an d p air ed n s

.
, ,
.

S b cl as s
u E l asmobr a chii shes with a ca til gi o u s k l to a d e p os ed g il l
1 . n ,
r a n s e e n n x

s lits in cl u di g the d og sh s k a tes sha k s

,
n tc , ,
r ,
e .

Su b cl ass 2 T l e s tomi shes with a mo e o l es b o y s k el eton a d with g il l s l its

. e o ,
r r s n n

c o ceal ed u n der a n o p er c l u m
n u .

Or der I Cr oss opter ygii p air ed n s with a b as al s talk ; P l ypt r u s

.
( S ee Fig ,
o e . .

FI G . I
A
. a n t erio r en d o f a cr o s so pt erygia n sh ( P ol ypter u s ) t o Sho w t he l ob e o r s t al k on whi ch
,

t he n is b o r n e . B, a n terio r en d o f a co m m o n t el eos t sh ( t ro u t ) t o Sho w the ab sen ce o f su ch a s tal k .

T his differen ce is n ot, ho wev er , an im p

dis tin gu ishin g c ha r ac ter b etween g r o u s o f shes
o r tan t p .
( F rom
B ridg e an d B ou l en g er in t he Ca mbr idge Natu r al Histor y, c ou r tesy o f the M a cmill a n Com an y p .
)

Or d er 2. p air ed n s withou t a s tal k ( Fig I ) s k l et n l g ly c ti

Chondr os tei, .
,
e o ar e ar

l agin ou s with a s p i al v l ve in the i tes tin e hea t with a co u s a t i

,
r ; a n ,
r n r er os u s

the s tu rg eo n ( Acipen s er ) a d the s poo n b il l ( P l yodon ) n o .

O der 3 r Hol ostei lik e the p r ecedin g b u t s k el eton w el l o s si ed ; the g a p ik e

.
, ,
r

( L epidos teu s) a d the river dog sh or b own (Amia) n .

These th ee o rde s a e commonl y refer ed to as the gan oid shes o wi g to

r r r r n

the shin y s cal es ( g an oid scal es) with which m os t o f them a r e cove ed r .

O der 4 r T l e s tei p air ed n s withou t a s tal k s k el eton w ell oss i ed witho t

. e o , , ,
u

s p i al val ve o r c on u s a terio s s ; a ll o f the c o mm on shes

r This o d er is
r u . r

c o mm o ly r ef e red to a s the t l eo s t s o r b o y shes

n r T he th ee o de s Chon e n . r r r

dr os tei Hol s t i an d T l eos tei a r e o f te g ou p ed tog ether as the A ti pt yg ii

, o e ,
e n r c no er

o r r ay n n ed shes in which the n

-
ays s p in g di ec tly f o m the b o dy in r r r r

co t as t to the Cr os
n r
pter ygii o r fri g e n ed sh s in which the so ay s n -
n e ,
n r

S p rin g f o m a s tal k a d f o m a so t o f f rin g e o n the e d o f the s talk

r n r r n .
 T HE P H L Y UM CHOR DA A T
Class 3 Amphibia , am i ians , lowest fou l egged er te
. ph b r -
v b rat es , s kins na k ed an d s lim y ,

o r w ith o n y b p
l at es ( ex tin c t) , livin g in or n ear wa te r .

Su bcl ass S tegoc phal

I . e tin ct amphib ia e a, x ,
with tails ,
an d c o v er ed wit h an ar mo r
o f bo y p l a t es
n .

Su b cl ass L is s amphibia p es e t day amphib ia

2. ,
r n -
, wit h n ak ed slimy skin s ( a few wit h
min u te , co n c eal ed scal es ) .

Ord er 1 . Ur odel a , with tail s ; the s al aman ders an d n ewts , Nectu r us Ambl ystoma , ,

Cr yptobr an chu s , etc .

Or der 2. An u r a , ta ill es s ; frogs ,

to ad s .

Cl as s 4 . Reptilia , .r e p til es ,
air - b
hin g ve teb a tes cove ed with horn y scal es r ea t r r r .

O d er 1
r Cotyl osau r ia
. Mo s t p rimitive g rou p of e tin c t ep til es r es embl i
. x r ,
ng

S tegocephal a in S k el etal chara c ters S eymour ia . .

Order 2 Chel on ia the tu r tl es body i cl osed in a hard c as e

.
, ,
n .

Ord er 3 . Rhyn cocephalia gin cl u din g b u t o n e an imal , the S phen odon (Ha tter ia)
or tu atara of New Zeal an d , a liz ard-like an i mal with p rimitive S k el etal cha r

ac ters .

Order 4 . S qua mata , u su all y of s mall or mo dera te s ize an d co vered wit h hor ny

scal es .

Su bor der 1 . L a cer til ia , the liza rds , n ea rl y al wa ys wit h l im bs , e yel ids
movabl e .

S u bor der 2. Ophidia , the sn a k es ,

d evoid o f l im s , eyelids b immo va l e b .

Ord er 5 Cr ocodil ia , the

. c r o codil es , al l ig ato rs , g avial s , an d cai man s ,
l arg e
rep til es with bo th horn y s ca l es an d bo y pl a tes
n in the s k in .

Cl ass 5 . birds vertebrates with fea thers

Aves , , .

Cl ass 6 . Mamma lia mammal s ver tebr ates with hair

, , an d mil k gl an ds .

Su b cl ass P r tother ia mammal s l ayin g egg s

I . o , .

Order Mon otr emata the mon o tremes o r egg l ayin g mammal s in cl u d in g on ly
I. , ,
-
,

the du ck b ill ( Or n ithor hyn chu s ) an d the sp in y an tea ters (E chidn a a n d P r o "

echidn a) o f the Au s tr alian reg io n .

Su bcl ass 2. E u ther ia , mamm al s bea r in g the you n g al iv e .

Divis ion I . Didel phia or Metather ia .

Or d er I . Marsu pialia ,
the m ar s u p ial s , mammal s be a r in g the yo u n g in a very

imm a tu r e s tate, hem in a pou ch fo rmed by a fol d of s k i o th

an d carr yi ng t n n e

abdomen pl a ce ta ab s e t o r primitive ; k an g a o o s O p ossu ms e tc

,
n n r , ,
.

Divis ion 2 Mon odel phia or P l acen tal ia the pl a cen tal mammal s withou t a
.
, ,

po ch you n g n ou rished in the u teru s by a pl ace ta which is produ ced

u ,
n ,

by a fu sion b etween certain pa ts of the embryo a d ce tain par ts o f the r n r

ma tern al u teru s This divis ion in cl u d es s ix tee o ders which a e des crib d
. n r r e

in d etail i N p ag es 3 46 4 4 I this man u al w e meet with thr ee o de s

n ,
-
0 . n ,
r r :

Order Car n ivor a the car ivorou s mammal s with cl aws a d Sharp cu tti g teeth;
,
n ,
n ,
n

the b ea s a ccoon s r minks mar te s weasel s o tters dog s foxes wo l ves

,
r , ,
n , , , , , ,

ca ts lion s tig er s hyaen as s eal s wal rus es

, , , , ,
.

O der Roden tia the rode ts with chisel lik e fro t teeth an d bac k teeth w ith
r ,
n ,
-
n ,

a t g rin din g
,
rf aces ; the ha es rabbits q i el s a ts mice po cu pi
su r , ,
s u rr ,
r , ,
r n es ,

gu in ea p ig s .

O de E den tata teeth l ackin g o d eg e era te ; the an t bears sl o ths an d ar madill s

r r ,
r n , . o .
8 L AB ORA OR T Y MANUAL FOR VE R TE B RATE ANATOM Y

Add ition a l
sc ien t i c n am es u s ed fo r c on v en ien c e p
in g rou in g the cl as s es o f ver t e b r a t es

ar e :

I chuzyops id a . This t erm , mean in g shlike an im al s , in cIu d es the two c l a ss es , P is ces an d

Amphibia .

S au ropsida . This term , mean in g re p til e l ik e

-
an im al s , in clu des the two cl a s s es , Reptil ia
an d Aves .

An a mn iota or An amn ia . This term in cl u d es the cl a sses P is ces an d Amphib ia an d r e er s f

to the fa ct that their mbryos
e are n a k ed .

Amn iota This emb aces the th e cl ass es R ptil i Aves d M mmal i an d refe s to
. r re , e a, ,
an a a, r

th fac t tha t the emb ryos o f thes e g o p

e e co v e ed b y a memb e the a m ion r u s ar r ran ,
n .

Ac iata o Acran ia
ran This n ame is sy o ymo s with Cephal ch d t d is deriv ed f o m
r . n n u o or a a an r

the abs ce o f the s k ull in Amphioxu s

en .

C an ia ta
r This te m is syn o ymou s with Vert br ta a d ef s to the f c t th t all ve te
. r n e a n r er a a r

b ates po ssess a S ku l l
r .

Agn th s t ma t
a o This term mea i g witho t j aws is syn o ymo s with Cycl stoma ta
o a. ,
n n u ,
n u o .

Gn ath toma ta os This ame mea in g j awed i l des all o f the cl as es of ver teb a tes
. n ,
n ,
nc u s r ,

e cep t Cycl
x t m ta os o a .

T t ap da
e r This term mea in g fou r foo ted i cl des all o f the la d v e teb tes i e
o .
,
n ,
n u n r ra , . .
,

the c l a sses Amphibia , Reptil ia , Aves , an d Mammal ia .

Io LAB ORA OR T Y MANUAL FOR VE R E B RA E ANA OM T T T Y

the ex terior throu gh the a triop ore Amphioxu s m ain tains a con tinu o u s cir cu la .

tion of wa t er thro u gh its di ges tive tra c t ; wa t er e n ter s the mo u th p ass es thro u gh ,

the wa ll s of the p harynx in to the a tri u m a n d o u t of the a trio p ore Make a

'

, .

drawin g of the an im al from the side .

2 In tern al an ato my
.
The in tern al ana to m y is m os t easily s tu died o n s mall
.

m o u n ted sp e cim en s E x am ine wi th the low po wer of the mi cros cop e

. I de n tify : .

the vario u s p ar ts of the n c on tai nin g re c ta n g u lar bodies the n r a ys servin g

, , ,

as the skele tal s upp or t of the n s ; the m u s cle segmen ts ; an d the di ges tive tra c t ,

o ccu p ying the ven tral half of the body The p ar ts of the di ges tive tra c t may .

be s tu died in so me de tail No te again the oral hood wi th its cirri The ves ti
. .

b u le is narrowed p os teriorly by a membran e the vel u m pier ced by a small op en , ,

in g the mou th The m o u th op ens in to a wide c avi ty the pharyn x ex ten din g
, .
, ,

half the len gth of the body I ts walls are p erfora ted by n u m erou s obliqu e.

sli ts the gill s lits or phar yn geal cl efts The solid p or tio n s of the p haryn geal wall
,
.

be tween the gill clef ts are called the b ran chial b ar s an d ea ch is su pp or ted by an
in tern al skele tal b r on chial r od Su rrou nding the p haryn x is a large cavi ty the
.
,

a trium the ven tral bou n dary of which is visible as a line below the p haryn x
,
.

This lin e m ay be tra c ed to the a triop ore The p os terior en d of the p harynx .

op en s in to a tu b u lar in tes tin e whi ch ex tends s trai gh t to the an u s The rs t p ar t .

of this in tes tine for a S hor t dis tan c e po s terior to the p haryn x is n arrow an d dor
sally lo ca ted ; the in tes tine then wide n s su ddenl y an d from this wide n ed p ar t
a blind sac the liver ex ten ds forward ben ea th the n arrowed por tion
, ,
.

Imm edia tely dorsal to the di ges tive tra c t and n early as wide as the p harynx
is a rod the n otochor d ex ten ding the len g th of the body I t will be seen by
, ,
.

fo cu sin g down thr o u gh the m yo to m es an d is m os t dis tin c t in the head where it ,

ru n s forward n early to the ex treme tip J u s t above the n o to chord is si tu a ted .

the m u ch s m aller n eu r al tu b e bes t re c o gn iz ed by the row of bla ck p i gm e n t sp o ts

,

whi ch it bears These p igmen t spo ts have been S hown to be sen si tive to li gh t
. .

D raw the whol e m o u n t showi n g its s tru c tu r e ,

.

3 Cross s e ction thro u g h the pharyn g eal re gion E x a m in e the c ross
-
. .

se c tio n wi th the low p ower an d iden tify the followi n g : (a) The ep idermis the ,

ou ter coverin g of the body comp osed of a single layer of columnar epi thelial cells .

(b ) The dorsal m e di an p roje c tio n the dorsal n co n tain in g an oval m ass the , , ,

n ray whi ch s u pp or ts it
, ()c The tw o ven t rola.t eral p roje c t io n s t he m e t a ,

p le u ral folds There are . a n u m ber of s m aller folds in th e ve n t ral wal l be t wee n

the two m e ta p le u ral folds ( )

d The m yo t o m es .a series of c ir c u lar m asses l ling ,

the dorsal and la teral p or tio n s of the body wall and sep ara ted fro m ea c h o ther ,

by con n e c tive tissu e p ar ti tio n s The m yo to m es are thi ck dorsally an d thin ou t

.

ven trally Tran sverse mu s cles are p resen t in the ven tral body wall j u s t above
.
,

the s m all folds of the ep id e rm is ()

e The n e u ral t u be a m e dia. n dorsal m ass , ,

oval or trap ez oidal in se c tio n lying be tween the dorsal p or tions of the m yo to mes
, ,

below the n ray Observe tha t it con tains a cen tral can al the n eu r ocoel
.
,
.
 G E NE RAL STUD Y OF TYPI CAL CHORDA E S T I I

B la ck sp o ts in the neu ral tu be are the p i gmen t cup s of simp le eyes or li gh t

p e r c eivi n g or g a n s These
. are dis t rib u t ed alo n g t he a n t erior p ar t of the n e u ral
tu be of Amphioxu s an d are si m ilar in s t ru c tu re to the eyes of p la n aria n s co n sis t ,

in g of a p igm en t cu p lled wi th n erve c ells E x p erim en t has S hown tha t these

.

op ti c cup s are sen si tive to li gh t while the eyesp o t on the an terior end of the
brain is insensi tive to ligh t (f The n o to chord an oval mass larger than the
.
, ,

n e u ral tu be a n d di re c tly ven tral to it (g) The a triu m the l arge cavi ty o ccu p ying
.
,

the ven tral half of the se c tion (h) The p haryn x the s tru c tu re o c cu p yi n g the
.
,

c en ter of the a tri u m I t is elo n ga ted in so m e regio n s h ear t shap ed in o thers

. It , .

c onsis ts of sep ara te p ie ces the bra n chi al bars ea ch i n c losi n g in its o u t er ex trem i ty
, ,

a s ti su pp or t the bran chial rod The sp a ces be tween adja cen t gill bars are the
, .

g ill sli t s or p hary n g eal c lef t s by m eans ,of whi c h t he c avi t y of t he p hary n x c o m
mu n ica tes wi th the a tri u m I n the m e dian dorsal li n e of the phary n x is a de ep
.

g roove t
,
he ep i b r a n ch ia l g r oov e; in t h e m edian ven tral li n e a broad S li gh tly ,

dep ress ed s tr u c tu re the hypob ran chial gr oove or en dos tyl e These grooves
, , .

se cre te m u cu s in whi ch the m inu te food p ar ti cles are cau gh t ( i) The liver an .
,

oval hollow s tru c tu re p resen t in some se c tio n s to the righ t of the p harynx I t .

is comp osed of tall ep i thelial cells whi ch p robably p rodu ce a di ges tive se cre tion
, .

The gonads In c ross se c tion s of larger sp e ci mens these are fo u n d as

.
-

m asses o n e o n ea c h side of and p roje c tin g i n t o the a tri u m

,
D raw the c ross .

s ec tion .

Fo r fu r ther de tails of the a n a to m y of Amphioxu s c o n s u l t P a n d H p ages ,

44 5 ;
-
6 C N H V ,o l V I I a
p g
. es,
1 1 2
3 0 ; N a
p g e s 30 4 2 , .

B A T UNI CATE
1 . E x ternalanato my .
Ob tain
a sp e cim en an d p lac e in a dish of wa ter .

I t is an oval sa c like c rea tu re s c ar c ely r e co gn iz able as a n a n i m al

,
The an i mal .

is in lif e p erm an en tly a tta ched to ro cks or o ther obj ec ts The en d whi ch was .

a tta ched is re cogn iz able by its rou gh an d irregu lar form and by the fragmen ts
of wood or o ther ma terials whi ch adhere to it The opp osi te en d which in the .
,

livi n g s ta te ex ten ds free in to the wa ter bears two op en i n gs the s iphon s Wh en , ,

.

the a ni m al is g en tly squ eez ed je ts of wa ter are observed to sq u ir t fro m the

,

sip ho n s ; herice the n am e sea sq u ir t p opu larly app lied to these a nimals The

u p p er of the two sip ho n s is the or a l or in cu r r en t si p ho n an d is the d eg e n era t e

an terior en d of the an i m al ; the lower sip hon is the atr ial or excu r ren t sip hon
a n d rep resen t s the p os terior en d T he a n tero p os terior axis of the tu n i ca tes is
.

ben t in to a U shap e The region of the body be tween the two sipho n s is there
-
.

fore dorsal an d the rem ain in g an d mu ch larger p ar t of the su rfa c e is ven tral
,
.

When the a n im al is u n dis tu rbed the sip ho n s are widely exp an ded an d a cu rren t
, ,

of wa ter p asses i n to the oral sip ho n a n d ou t of the a trial sip ho n Wh en dis tu rbed .

a n d also g en erally in the p r eserved s ta te the si p ho n s are c losed a n d re tra c ted ,

.

C l os u re a n d r ea c tio n is a ccom p lishe d by cir cular an d lon gi tu din al m u s cl es lo ca ted

I 2 LAB OR A OR T Y MANUAL FOR VE R E B RA E ANATO M
T T Y

in the body wall I n c losin g the rim of the sip ho n s is thro wn i n to folds whos e
.
, ,

n u m ber a n d arra n g em en t varies wi th dif fere n t sp ecies .

Th e o u ter layer of the body wall c o n sis ts of a thi ck to u gh so me times so m e , ,

wha t gela ti n ou s of ten war ty c overin g the tu n ic or tes t whi ch is se cre ted by
, , , ,

the e c toderm be n ea th it I n so m e tu n i ca tes ( Cion a Corell a) the tes t is so tran s

.
,

p are n t t ha t p ra c ti c ally all o f the i n tern al org ans are visible thro u gh it The .

tes t is a tta c hed to the u n derlyin g body o n ly a t the si p hons .

Draw the an imal .

2 .

D isse ctio n of a tu ni c ate (A dem o n s tra tion disse c tion m ay be su bs ti tu ted
.

for in divid u al disse c tion by the s tu den t ) As dier en t tu n i ca tes vary in the .

de tails of their i n tern al s tru c tu re the followi n g a ccou n t is of ne cessi ty somewha t

,

ge n erali z ed
. I t is based c hiey o n the ana to m y of Cion a M ake a cu t t hro u g h .

the base of the tu n i c avoidi n g i n j u ry to the u n derlyi n g wall a n d p eel of

, f the ,

tu n i c . No te its a t ta ch m en ts a t the si p hons and cu t thro u gh th ese a tta ch men ts , ,

dis cardin g the tu nic The sof t b ody wall or man tl e is th u s revealed the tu n i c
.
,

being as already exp lained a se cre tion of the ou ter layer of the m a n tle The
, , .

mu s cle bers whi ch op era te the sip hons may be visible as s trip es in the m an tle ;
they are co n sp i cu o u s in Cion a .

F as te n the a nim al in a wax bo tt o m ed disse c ti n g p a n by p i n s thro ugh the

-

rim s of the sip ho n s an d thro u gh the ex trem e basal en d F ill the p a n wi th wa ter .

to c over the a nim al .

The man tle in closes the vis cera whi ch tend to adhere to it so tha t its removal ,

wi thou t i n j u ry to the vis c era is dii cu l t I n tran sp aren t form s the vis cera m ay
.

be s tu died throu gh the m an tle I n case it is n e cessary to remove the m an tle

.
,

m ake a lon gi tu di n al sli t in it fro m the a trial sip ho n to the basal en d a n d p u ll it

off in s m all s tri p s sep ara ti n g ea ch s tri p g e n tly fro m the u n derlyi n g p ar ts
,
The .

p ri n c i p al i n t er n al p ar t s are t he followi n g The oral si p ho n leads in

. t o a lar g e
thin walled b a g whi ch in so m e form s as Cion a ex te n ds the le n g th of the body
-
, , , .

This b ag is the pharyn x I ts wall ap p ears to the naked eye like m osqu i to ne ttin g
.
,

be cau se it is pier ced by nu m ero u s mi n u te op en i n gs the gil l slits or vis cer al cl efts ,
.

The c avi ty ou tside of the phary n x is the a tr iu m; it c ommu n i ca tes wi th the

ex terior by way of the a trial sip ho n The lower en d of the p hary n x n arrows .

abrup tly in to a shor t esophagu s whi ch op en s in to a widen ed s tomach si tu a ted ,

below or to on e side of the basal en d of the p harynx The s toma ch is cu rved .

so tha t its lon g axis lies a t ri gh t an gles to the lon g ax is of the p hary nx I n so me .

c ases (M ol gu l a ) brown diges tive gl a n ds c a n be see n c overin g the s to m a c h The .

s tom a ch leads in to an in tes tin e whi ch immedia tely m akes a loop dou bli n g ,

ba ck so as to lie p arallel to the s tom a ch I t th en ben ds a n d ex ten ds s traigh t .

up ward toward the a trial sip ho n term i n a tin g by an a n u s si tu a ted wi thi n the
,

a trial c avi ty .

The rep rodu c tive organ s c o n sis t of a sin gle or p aired m ass ea ch bein g her ,

m ap hr o d itic tha t is c o mp os ed in p ar t of a n ovary and in p ar t of a tes tis

, , This .

her ma phroditic gon ad is lo ca t ed ei ther in the sp a ce be tween the in t es tin al loo p

 G E NE RAL STUDY OF TYP I CAL CHORDA E S T I 3

an d the s tom a ch (Cion a Corel l a Ascidia ) in whi ch case it is co mmonly S in gle or

, , ,

may be a t ta ched to the inside of the m a n tle (Cyn thia M olgu l a ) in whi ch case ,

a p air of s u c h glands is of ten p resen t F ro m the ovarian p ar t arises an ovidu ct

.

a n d fro m the t es ti cu lar p ar t a vas de er en s The d s in lose o a

f t w o u c t ru
. n c c n t ct

wi th each o ther ( and in som e cases fu se to one d u c t) and op en in to the a trial

c avi ty near the a trial sip hon I n Cion a the two gen i tal d u c ts are fou nd c losely
.

a tta ched to tha t side of the in tes tine whi ch fa ces the p haryn x an d in this p osi ,

tion r u n far forward o p enin g in to the a tri u m so m e dis tan ce above the anal o en
,
p
mg . The gen i tal op en i n g in Cion a is colored red du e to the p resen c e in its walls ,

of red vesi cl e s s u pp osed to have an excre tory fu n c tion I n M ol gu l a an d o ther

, .

mem bers of the fa mily Mol gu l ida e there is a larg e sa c like organ on the ri gh t side ,

believed to be ex c re tory in n a tu re b u t the p resen ce of ex c re tory org ans in the

,

tu n i c a tes is m ore or less u n c er tai n .

I n the dorsal p ar t of t he m a n tle be twee n the two sip ho n s will be fo u n d an

, ,

elon ga ted m ass the gan glion whi ch co n s ti tu tes the en tire ce n tral nervo u s sys tem
, ,

of the ad u l t tu n i c a te Nerves may be seen ex ten din g fro m its en ds to the sip hons
.
.

Draw the disse c tion .

3 The s tr u ctu re of the pharynx

.
M ake a lo n gi tu dinal sli t in the side of
.

the p harynx forward thro u gh the oral si p ho n and sp read o u t its walls I n the .

median ven tral line of the p harynx there is p resen t a co n sp i cu ou s whi te cord
ex ten di n g the len gth of the p harynx This is the en dos tyl e or hypobr an chial .

g ove whose walls are co mp osed of gla n d u lar and ci l ia ted c ells the form er
r o , ,

p rod u c i n g m u cu s D ire
. c t ly o pp osi t e th e e n dos tyle in wha t may be design a ted
the m edia n dorsal l i n e of the p hary n x is lo c a ted the dor s al l a min a This in .

som e tu ni ca tes (As cidia Cyn thia M olgu l a ) co n sis ts of a deli ca te m em bra n o u s
, ,

fold of the p haryn geal wall while in o thers (Cion a Corell a) it consis ts of a row
, ,

of p ro cesses called l an gu ets an d resembles a frin ge At the an terior en d of the .

p haryn x a t th e bo tt o m of th e oral si p hon t her e g e n erally o ccu rs a c ir c le t of

te n ta c les or irregu lar p ro c esses of the wall an d p os terior to this is a grooved ,

ridge the per ipharyn geal b an d in whi ch the a n terior en ds of en dos tyle an d
, ,

dorsal lam i n a term in a te The la t eral walls of the p haryn x in som e tu n i ca tes
.

( M ol gu l a Cyn thia) ex hibi t a d e n i te n u mber of lo n gi tu din al folds

, .

Cu t ou t as mall p ie c e of the p hary n g eal wall m o u n t in wa ter sp r ead o u t , ,

a t an d ex a m in e u n d er the low p ower of the m i cros c op e or ex ami n e p rep ared

, ,

slides The wall is fou nd divided i n to squ ares by m eans of lo n gi tu din al an d

.

cross bars .Wi thin ea ch squ are so formed are s een s everal of the elonga ted
g ill sli t s or vis c eral c lef t s s ep ara
, t ed fro m ea c h o th e r by s m all e r bars The .

g ill sli t s are c o m m o n ly p arallel t o t h e lo n g i t u di n al bars b u t m a y be cu rved

( M ol gu l a ) O
. n the lo n g i tu di n al bars t here are of t e n p r e se n t n g e r like p ro c esses -

or papill ae whi ch have been S hown to aid in pu shi n g the food con tai n in g m u cou s -

s trands toward the esop ha gu s D raw a p or tion of the p haryn geal wall
. .

F o r fu r ther de tails of the a n a to m y of tu n i c a t es c o n s u l t P an d H p ages 1 4 2 1 ;

,

CNH Vo l VII p a g es 3 5 6 2 ; N p ag es 50 52 H p ag es 50 5 6
, .
,
- , , ,
-
.
I 4 LAB ORA OR T Y MANUAL F OR VE RTE B RA E ANA OM T T Y

C . B AL ANO G L OS S US
1 . Ext ern al a n a to my
a s pec im en p lat e in a dish of wa ter an d
O b tain
.
, ,

ex ami n e (As di fferen t sp e ci es d i er so mewha t in app eara n ce the followin g

.
,

des c rip tion app lies o nl y to B al an ogl oss u s "Dol ichogl os su s " kovval evs kii ) I t is a n .

elo n ga ted worm l ike a n im al abo u t ve or s ix i n ch es in l en g th The body is .

divided i n to three p ar ts the pr ob os cis the col l ar and the tru n k The p robos cis , , .

is the slen der co n i c al p oi n ted s tr u c tu r e a t the an terior en d u sed by the a n i m al

, , ,

in b u rrowin g in to the san d The c ollar is the ba n d en cir clin g the body j u s t
.

p os t erior to t he p robos c is The m o u t h is c o n c.ealed wi t hi n t h e c ollar The .

tr u n k co mp rises the g rea ter p ar t of the a n im al and p res en t s a m ore or less

ru fed app earan ce The m edian ven tral lin e is m arked by a dis tin c t l on gitu di
.

'
n al ban d ; the m edia n dorsal lin e in a si milar b u t less dis ti n c t m a n n er The .

sides of the tru n k for abou t an in ch a n d a ha l f p os t erior to the c ollar p roje c t

som e wha t an d are sligh tly differen t in app earan c e fro m the res t of the t ru n k ;
th ese p roje c tio n s c o n tain the rep rod u c tive org a n s a n d hen c e are n a m ed the

g e n i ta l r id ges B e t wee .n t h e a n t erior p or t io n s of t h e g e ni t al rid g es on t he dorsal

side will b e fou n d two lon gi tu di n al rows of s m all p arallel sli ts the gill pores , .

( The tru e gill sli ts leadin g from the p harynx are U shap ed an d ea ch op ens in to
, ,
-

a chamber the gill p ou ch whi ch in tu rn c omm unica tes wi th the ex terior by m ea n s

, ,

of the gi ll p ore whi ch is so m ewha t sm aller than the gill sli t I n som e balan o
,
.
.

g l o s s id s as P t
,ycho d er a th e U sha p ed g ill sli
,
ts o p e n di re
-
c t ly t o t he e x terior a s ,

is the c ase in the larva of Amphioxu s The s tru c tu re a n d dev elop m en t of the .

bran chi al r egion of b al an ogl o ssids is s trikin gly similar to tha t of Amphioxu s ) .

The rem ain der of the t ru n k is o ccu p ied by the di ges tive tra c t an d p oss esses a
t er m i n al a n u s The sp e cies u n der c o n sidera tion la cks the hep a ti c ca eca a
.
,

series of ou tgrow ths of the i n tes tin al wall gu r ed in P a n d H M ak e a drawin g ,

.

of the a n im al fro m the dorsal side .

F o r fu r ther de tails c o n s u l t P an d H p ages 2 7 ; CNH Vo l VII p a ges 3 2 1 ; -

, , .
,

H p a ges 51 2 1 4 ; N p ages 6 0 6 5
.
.
,

D . ANATO M Y OF A L AMP R E Y
The la mp reys belo n g to the lo w es t class of tr u e ver tebra tes the c lass Cycl o ,

s toma ta or ro u n d m o u thed shes

,
-
.

1 E xt e rnal anato my of a l amp rey

.
P la c e the s p ecI m en in a diss ectin g .

tray T he body c o n sis ts of a s to u t c yli n dri c al h ead an d tru nk a n d a a t ten ed

.

t ail .F ro m the m edia n dorsal li n e in the p os t erior p or tion of the body ar ise
two dor s al ns while the tail is p rovided wi th a cau da l n c o n tin u o u s with
, ,

the se co n d dorsal n The n s are s u pp or ted by n u merou s n r ays slen der

.
,

p arall el c ar t ila g e s u s u ally visibl e t hro u g h th e S ki n There are n o p aired .

ven tral n s s u ch as a r e fo u n d in the tru e shes The skin is sof t sli m y an d . , ,

n aked .
 G E NE RAL STUDY OF TYP I CAL CHORDA E S T 1 5

The a n t erior en d of the head p rese n ts a p e cu liar app earan ce o wing to the ,

app are n t abse n c e of a lower jaw ( the jaw is p robably p r esen t in a de en era te
g
c o n dition ) and the p rese n c e on the ve n t ral S ide of the head of a larg e bowl

shap ed dep ressio n the b u ccal fu n n el The ed ges of the b u ccal fu n n el are p ro
, .

vid ed wi th sof t papill a e and it s in terior is s tu dded wi th brow n hor n y teeth

, , ,

den i tely arra n ged At the bo ttom of the fu n n el is a p rojec tio n called the
.
,

ton gu e; also c overed wi th t ee th The m o u th is j u s t dorsal to the tongu e L amp

. .

reys a tta ch t hem selves to the bodies of shes by m ea n s of the b u ccal fu nnel and
rasp o ff their esh by li n g m ove m en ts of the to n gu e .

On the dorsal su rfa c e of the head is an o p e n i n g the n as al a per tu r e or n os tril , ,

whi ch leads in to the olfa c tory sac Cy clos to m es di ff er fro m o ther ver tebra tes
.

in possessi n g b u t one nos tril al tho u gh thi s is p robably a se c o n dary co n di tio n

, .

On ea ch side of the head on a level wi th the olfa c tory organ is an eye n o t p rovided ,

wi th ey elids P os terior to the eye on ea ch side of the head is a row of s eve n

.

oval op en i n gs the gill s l its

,
.

Alo n g the sides of the body the bo u ndaries of the myo to mes are gen erally
n o ti c eable thro u gh the skin I n the m edian ve n tral line a t the j u n c tion of tr u n k
.

and tail will be fou n d a p it In the an terior end of this pit is the an al op en in g
.

an d i m m edia t ely beh i nd the anu s a p roje c tin g p ap illa the u r ogen ital papil l a , ,

whi ch r eceives the p rodu c ts of the kidneys an d rep rodu c tive organ s and dis
char ge s the m to the ex terior thro u g h an o p ening a t its tip the u r ogen ital ,

a er tu r e
p .

2. S a gittale ction of the anterior

s M ake
a median sagi ttal se c tion of
en d .

you r sp e cim en to a p oin t abou t an in c h p os terior to the las t gill S li t or s tu dy a

se c tion so p rep ared E x amin e the cu t s u rfa c e and iden tify the followi n g :
.

a ) Diges tive tra ct: Observe a gain the b u cc al fu nel wi th its t ee th a n d to n gu e

n .

No te the large mu s cle m asses ex tending p os teriorly fro m the to n gu e by mea n s ,

of whi ch the rasp in g m ovem en ts of the to n gu e are bro u gh t abou t Fin d the .

mo u th op en i n g above the to n gu e and follow it in to a p a ssage the b u ccal cavity ,

whi ch slop es ve n trally The b u ccal cavi ty op e n s a t its pos terior en d i n to two
.

tu bes an u pp er s m aller one the es ophagu s a n d a larg er ven t ral o n e the pha ryn x
, , , , ,

the wall of whi c h is p ier c ed by seven oval o p en in g s A fold the vel u m is p rese n t .
, ,

a t the en tran ce of the b u c c al c avi t y in to the p hary n x The esop hagu s leads .

in to the rem ain der of the diges tive tra c t b u t the p hary nx whi ch in the , ,

embryo co n s ti tu ted the a n terior p ar t of the diges tive t rac t en ds bli n dly in ,

the ad u l t .

b ) Res pir a tor y s ys tem: T he seven Op en i n gs in the wall of the p haryn x are
the i n t ern al gil l s l its They op en in to mu ch en larged gill pouches whi ch bear
.

the gills o n their walls an d whi ch c o m m u n i c a t e wi th the e x t erior thro u gh the

ex tern al gill sli ts alr eady no ted P robe in to o n e of the gill p o u c h es and n o te the
.

leaves or gill l a mell ae born e o n its walls The dorsal por tio n s of the gill p ou ch es
.

in close the n arrow eso p hagu s be tween them .

 1 6 L AB ORA OR T Y MANUAL F OR VE RT E B RA E ANA OM T T Y

)
c Notochor d : The is the broad brown rod si tu a ted j u s t dorsal to
n otochor d ,

the esop ha gu s I t is the chi ef ax ial skele t o n of tho a nim al the ver tebral c ol u mn
.
,

b ein g ver y em bryoni c .

d) Nervou s sys tem: Above the n o to c hord is a n arrow c anal the n eu r al can a l , ,

in whi ch the s p i n al cord and brai n are lo ca ted The spin al cor d is a slen der
.

c ord o c cu p yi n g the c a n al ; the b ra in is an en lar g ed lobed s t ru c tu re si tu a ted j u s t

dorsal to the an terior ex tremi ty of the n o to chord .

e) Olf a ctor y a ppa r a tu s : The se c t ion sho u ld bise c t the olfa c t ory a er tu re
p .

No te tha t a c a n al ex te n ds fro m this ap er tu re an d op en s i n to a s ac the olfa ctory ,

s ac S i tu a t ed j u s t a n t erior to the tip of the brai n

, The folds in the wall of the.

s a c bear the olfa c t ory m u c o u s m e m brane F ro m the a n terior m ar gi n of the

.

o lfa ctory sac a tu b e arises whi ch ben ds ven trally and p os teriorly an d widen s
in to an elo n ga ted sac the pitu itary pou ch lyi n g ven tral to the an terior en d of
, ,

the no to chord The p e cu liar dorsal p osi tio n of the olfa c tory sac as well as its
.

rela tio n to the p i tu i tary s ac (whi ch in o ther ver tebra tes has n o con ne c tion wi th
the olfa c tory c avi ties ) are d u e to a shif ti n g of these org a n s d u ri n g develop m en t
fro m their origin al ven tral p osi tion ( see P and H F ig 79 9 n The ,
.
, .

p i tu i t ary s a c c orres p o n ds t o t he a n t erior lobe of t h e p i tu i t ary body or hy o

p p hysis
of o ther ver tebra tes .

f) P er icar dial ca vity: P os terior to the las t gill p o u c h is a so m ewha t coni c al

c avi ty the per icardial cavity wi thin whi c h the hear t is si tu a ted
, ,
.

M ake a de tailed drawing of the sa gi ttal se c tion .

F o r a c o mp le te a cco u n t of the a n a to m y of c y c los to m es c o n s u l t P a n d H ,

a
p g es 1 1
9 39 ; C N H V o l V I I,
a
p g es 1.
5 5
0
2,
2 1 6 2 2 2

47 4 8,
2 79
8 2 1

3 7 73 , , , ,

3 5 9 5; N p a g es 8 7 9 4
8 - -
.

E . E X TE RNAL ANAT OM Y OF TH E DO G FI S H

The dog sh or dog shark is a tru e sh belo n gi n g to the lowes t group of ,

shes the s u b class E l a smobr an chii I t is a very ge n erali z ed ver tebra te an d

, .
,

hen ce a k n owled ge of its s tru c tu re is in disp ensabl e for an u n ders tandi n g of

ver tebra te an a to my There are two co mmon do gshes u su ally ob tai n able for
.

labora tories the smoo th d ogsh M u stel u s can is a n d the sp i n y dogsh Acan thias
, , , ,

vu l ga r is These two sp e cies are S ligh tly di er en t in several resp e c ts ; a tten tio n
.

will be called t o thes e di er en ces where n e c essary .

1 .

B o dy an d s kim The body of the d og sh has the shap e a n d p ro p or tio n s
whi ch we re cogn i z e as mos t adva n ta geo u s for free swimm in g an im als fu siform -

s
( p i n dle sha ed
p ) and p oin t ed a t ea c h e n d t h u s o fferi n g,
li t t le resis t a n c e to the

wa ter The body is divided in to head t ru n k an d tail whi ch are n o t how ever
.
, , , , ,

dis tin c tly bo u n ded from ea ch o ther Tru n k an d ta il are p rovided wi th n s for
.

p p
u r oses of lo c o m o tion The body is .c lo t hed wi t h m in u t e s ca l es ea ch of whi c h .

bears a tin y spin e P ass the han d over the S kin of the do g sh an d n o te the
.
1 8 LAB ORATOR Y MANUAL FOR VE R TE B RATE ANATO M Y

of tru n k an d tail the pelvic ns In m ale d og shes the m edial side of ea ch p elvi c
,
.
,

n is m odied in to a s to u t p ro cess dire c ted p os teriorly These p ro cesses are .

ca l le cl a s pers and are u sed by the m ales in m a n g wi th the fem al es of

d A ll .

the ns are s u pp or t ed by slender ex ible rays the der mal n r ays i m bedded in , ,

the skin of the ns .

5 An u s B e tween the two p elvi c ns is a large op enin g , the an u s or cl oac al

-
. .

a per tu r e Thi s is the op enin g of a cham ber the cl oaca whi ch re ceives the end
.
, ,

of the in tes tin e an d the termi n a tions of the du c ts of the kidn eys an d r ep rodu c
tive org a n s The a n u s of the d ogsh and of the m ajori ty of ver tebra tes does n o t
.

q ui te c orresp o n d to the an u s of m an and o ther mamm als which re ceives the ,

in tes tine only b u t the term is c omm onl y u sed to desi gn a te bo th c lasses of
,

Op eni n g .

F . E XTE RNAL ANAT OM Y OF THE S KATE

The ska tes or rays are like the do gsh elasmobra n ch shes b u t highly
, ,

modied as regards ex t er n al form and p rop or tions Ob tain a sp e ci men and .

ex ami n e .

1 .

B o dy an d s kim The b ody is di vid ed in to a g rea tly a tten ed an terior

p or tio n c o m p risin g head and t ru n k and a S lender p os t erior p or tion the t ail , .

The broad at form is chara c teris ti c of bo ttom feeding shes ( see N pp 9 7 9 9)

,

-
, .

and resu l ts from a shor tening of the dorsoven tral axis an d an elo n ga tion of the
t ra n sv erse ax is The tou gh skin c on tains s c a ttered s cales of whi ch the spi n es
.

p roje c t c ons p i cu o u sly These s c ales are.of th e sa m e typ e alr e ady n o t ed in t he

dog sh c onsis ting of a b as al pl a te imbedded in the S ki n an d a p roje c ti n g spin e

, ,

dire c ted pos teriorly The s cales of the ska te are mu ch larger an d fewer in n u mber
.

tha n those of the dogs h They are deni tely arran ged in lines an d grou p s the
.
,

arrangemen t dierin g som ewha t in the two sex es In fem ales there are s cales .

over the la teral exp an sions of the tru nk and several rows of s cales on the m edian
dorsal p ar t of the tru nk an d dorsal s u rfa c e of the tail while in m ales the g rea ter ,

p ar t of t h e la t eral e xp a n sions is devoid of s c ales there are fewer rows alo n g the ,

m iddle of the ba ck and tail and the s cales o n the m argins of the head are e n larged
,
.

I n m ales fu r ther m ore there are two rows of cu rio u s ere c tile sp i n es o n the la teral
, ,

exp an sion s of the tru n k abou t an in ch in from the m argi n ; these can be ere c ted
and lowered in to dep ression s in the S kin Are there any s cales on the ve n tral .

s u rfa ce ? No te the m arked differen ces in c olor be tween the dorsal and ven tral
s u rfa ces .

2 Fin s
.
L ike the dogsh the Ska te is p rovided wi th m edian an d p aired
.

ns b u t the former are mu ch redu ced They co n sis t of two s mall dorsal n s
, .

on the dorsal side of the end of the tail The pectoral ns are en orm ou sly .

e nl arged formin g the la teral exp ansions of the tru n k already m en tio n ed several
,

tim es ; they are con u en t an teriorly wi th the m argi n s of the h ead The pel vic .

n s a r e s m aller and imm edia t ely p os terior to the p e c toral n s wi th whi ch they
 G E NE RAL STUDY OF TYP I CAL CHORDA E S T 1 9

are co n ti n u ou s in so m e sp e cies They consis t of two l obes an d in the males

.
,

bear large s tou t cl aspers de ep ly grooved along their p os terior la teral margins
, .

The clasp ers are emp loyed in m a ting wi th the females .

3 H ea
. d The head like the t ru nk is grea tly a ttened its margins c on
.

tin uou s wi th the p e c toral ns I t t ermi n a tes in a p oin ted r os tr u m

. I t bears .

dorsally a p air of larg e p roje c tin g eyes wi thou t lids B ehin d ea ch eye is the , .

large spir acl e or rs t gill s lit bearin g a valve on its an terior fa ce marked by
, ,

p arallel rid g es whi c h re p resen t a ru di m en t ary g ill O n th e ven t ral side of the
.

head is the mou th bou nded by too thbearin g jaws The jaws and tee th are -
.

c o mm o n ly larg er in the m ales than in the fe m ales In fron t of the mo u th are .

the two nos trils ea ch p rovided wi th a fringed ear shap ed ap

, E x tendin g -
.

p os t eriorly fro m ea c h nos t ril t o t h e an g le of t he m o u th is a ap wi th a frin g ed

p os t erior m ar g i n This . is th e n a s of r on t a l p r ocess O n lif tin g pu t his p ro cess
. a
deep groove the oron as al gr oove will be fou nd extendi n g from the nos t ril in to
, ,

the m o u th c avi ty This arran gem en t foreshadows the app eara n ce of a closed
.

p assa g e fro m th e nos t rils in t o t h e m o u t h s y c h as is p resen t in hi g her ver t ebra t es .

P os terior to the m o u th are ve p airs of gill sli ts I n ska tes the p e c toral ns have
.

g rown forward above th e g i l l sli t s and f u sed wi t h th e sides of t h e head th u s ,

shovin g the gill sli ts to the ven tral su rfa c e .

/ a 4 A n.u s The . an us or c l o a ca l a p e r tu r e is a lar g e o p enin g be tween t h e

bases of the p elvi c ns .

G . E X TE R NAL ANAT OM Y OF A TE LE OS T

F o r this p u rp oseany c ommon bony sh can be u sed b u t the followi n g ,

des crip tion is based up on the p er ch The p er ch like o ther comm on sh es is a .

m em ber of the g rea t order Tel eos tei Ob tain a sp e ci men an d no te the followin g
.

p oin t s .

1 .

B o dy an d s kim The body has the S hap e typ i c al of aq u a ti c ani m als ,

thi ckes t in the m iddle a n d tap eri n g to ea ch en d I t is so m ewha t co mp ressed .

la terally I t is in dis ti n c tly divided i n to head tru nk an d tail Tru n k and tail
.
, ,
.

are clo thed wi th s cales arra n ged in diago n al rows overlappi n g ea ch o ther ,
.

These s cales are set in p o cke ts in the deep er par t of the ski n (dermis) as may be ,

de termin ed by rem ovin g on e of them and the s u p er cial layer of the skin (ep i
dermis ) form
,

s a thin hn over their s u rfa ces The head is c overed by the sof t .

ep iderm is an d in som e regions bears small s cales like those on the remain der of
the body B enea th the s c aleless p or tions of the ep iderm is of the head will b e
.

no ted large thi n a t bo n es These bones whi ch are the o u ter bon es of the
, ,
.
,

sku ll are in reali ty no thin g b u t en larged s cales whi ch have su n k from their
,

original su p er cial p osi tion to a deep er lo ca tion A la teral li n e is p resen t alo n g .

ea ch S ide of the body .

2 .

H ea d The head bears a term inal mou th bou nded by well d eveloped
.
-

jaws The termin al p osi tio n of the m ou th is p robably m ore p rimi tive than the
.
20 LAB ORA OR T Y MANUAL FOR VE R E B RA E ANA OMT T T Y

e ral p osi tion fo u n d in the elasmobra n chs On the dors al side of the an terior
v nt .

en d of the head are t wo p airs of n os t rils a p air to each olfa c tory sac This , .

arrang em en t p ermi ts a cu rren t of wa t er to cir cu lhte throu gh the olfa ctory sac .

The large ey es are wi thou t lids The ears si tu a ted behi n d the eyes are invisible
.
, ,

ex tern ally The p os terior an d la teral m argins of the head are form ed by a large
.

ap the gil l cover or oper cu l u m whi ch is su pp or t ed by sev eral oper cu l ar b on es

, , ,

large at s calelike bo n es already no ted I t c overs a wide sli t in the body wall
, , ,
.

kn own as the gil l open in g Atta ched to the ven tral margin of the o p er cu lu m
.

is a m embran e the b r an chios tegal memb r an e s upp or ted by s even b o n y rays the
, , ,

b r an chios tegal rays G rasp the m em bra n e wi th a for c ep s a n d s p read it ou t to

.

see the rays L if t u p the op er cu lu m an d look wi thin the c avi ty which it covers
. .

Fou r cu rved s t ru c tu res the gil l a r ches whi ch sho u ld be sep ara t ed fro m ea ch
, ,

o ther wi th a for cep s will be seen E a ch bears on its ou ter s u rfa ce a gill co n
,
.
,

sis ti n g of a dou ble row of sof t l amen ts an d o n its i n n er margin a series of shor t ,

too thlike p ro c esses the gil l r a ker s ,

Thru s t a p robe inward be tween two gil l
.

ar ches op en the m ou th of the sh and observe tha t the en d of the p robe has
, ,

en tered the m ou th cavi ty The cavi ty of the p harynx is th u s in commun i ca tion

.

wi th the ex terior throu gh the sp a c es be tween the gill ar ches These sp a ces are .

g ill sli t s c orres p ondin g t o t hose w hi c h we saw in t h e d o g s h and ska t e b u t here ,

the p or tio n s of the body wall be tween s u cc essive gill sli t s have disapp eared an d ,

all o p en i n t o a c o mm o n c avi ty c overed by the o p er cu l um Thi s c ondi tion is .

chara c t eris ti c of all shes ex c ep t elas m obra n chs When the sh resp ires the .
,

m ou th Op e n s the op er cu la m ove o u tward the bra n c hios te gal m em bra n e u n folds

, ,

an d c loses the gill o p eni n g ; wa t er is th u s draw n in to the m o u th a n d ba th es

the gills The m ou th then closes the op er cu la move in ward the b ran chios te
.
, ,

g a l m e m bra n e folds u p a n d the wa t er p asses

,
o u t t hro u g h th e g ill sli t s a n d
g ill o p e n i n g .

3 F in s The body is p rovided wi th m edian an d p aired n s Of the former

-
. . .

there are an a n terior an d a p os terior dorsal n a ca u dal n an d a ven tral or , ,

anal n ( The n u m ber an d p osi tion of the m edian n s are very variable in differ
.

en t shes ) The cau dal n is app aren tly symme tri cal wi th the en d of the tail
.
,

formi n g a homocer cal tail The p a ired ns are the sam e as in the d ogsh The
. .

p e c t oral n s are lo ca t ed j u s t behind t he o p er cu l u m b u t t he p elvi c n s have ,

m oved forward fro m their norm al p osi tion a t the level of the a n u s to a p osi tio n
n early level wi th the p e c t oral ns Su ch a forward mi gra tio n of the p elvi c n s
.

is very common in the teleos t shes and is of ten asso cia ted wi th dimi n u tion and
degen era tion of these ns The an terior dorsal n is s upp or ted by sharp hard
.

sp i n es an d these are also p resen t in o n e border of so m e of the o ther n s b u t the

, ,

la tter are s u pp or ted chi ey by ex ibl e bon y n r ays .

4 O p e n in g s
.
I n the m edi an ve n t ral line j u s t in fro n t of the ve n tral n
is a large op en i n g the anu s B ehi n d this is a d ep r ession in to whi ch p roje c ts a
,
.

small eleva tion the u r ogen ita l papil l a I n the p er ch th er e is n o c loa ca b u t the
, . ,
 GE NE RAL S UD T Y OF TYPI CAL CHORDA E S T 21

in tes tine and u rog eni tal sys tems have sep ara te op enings ; this is one of the m arked
di fferen c es be tween elasmobran ch and teleos t shes .

H . S OME G ANOI D F I S HE S
o

As er tain ganoid shes are freq u en tly referred to in the s tu dy of the skele ton
c
,

it is advisable tha t the s tu den t be co m e fa miliar wi th their ap p earan c e The .

g a n oid shes belong to the lower and more p rimi tive orders of the Tel eostomi
a n d hen c e are p la c ed in the s c hem e of c lassi c a tion below the t eleos ts b u t the
,

la t ter have been con sidered rs t to in trodu ce the terminology We S hall ex amine .

briey fou r comm on gan oids .

1
. The g ar p e ( L epid os teu s )
ik The body is c lo thed in a co mp le te armor
.

of rhomboid s cales set in diagonal rows Su ch s cales are c alled gan oid s cales .

and consis t of bo n e wi th an o u ter coa t of a shiny su bs tan c e named gan oin .

These s cales p ass o n to the head as enlarged p la tes whi ch cons ti tu te the bones of ,

the roof of the sku ll The jaws are very elon ga ted p rodu c in g a s n ou t on the
.
,

tip of whi ch are the s m all nos t rils The jaw s bear n u m erou s S harp tee th There
. .

is an op er cu lu m c overing the gills A sp irac le is absen t No te m edian and

. .

p aired ns wi th ,
t heir s t o u t joi n t ed n rays
,
The t ail is he t ero
. c er c al b ut
app roa ches the homo c er c al typ e .

2
. The s tu rg eon ( Acipen s er) There is a c overing of ganoid s c ales arran ged
.

in ve rows on the tru nk wi th areas of app aren tly naked skin (whi ch really bear
small s ca les) be tween the rows The s cales la ck the ou ter coa t of ganoin Mos t
. .

of them bear sharp p os teriorly dire c ted sp ines They pass on to the head .
,

transform in g in to sku ll bones The head is broad and t ermin a tes in a larg e
.

ros tru m bearin g man y s m all s cales on its dorsal su rfa ce The n os tril is divided
, .

by a p ar ti tion in to two op en ings The Op er cu l u nkcon tain s a sin gle bon e Th ere
. .

is a sli tlike sp ira cle above ea ch eye On the ven tral side of the ros tru m are fou r
.

bran ched p ro cesses the b ar b el s u sed as sense organs for the de te c tion of food
, , .

The m ou th is cu riou s in form dis tensible and wi th sensory p ap illae on its

, ,

borders The jaws are degenera te an d there are n o tee th No te p aired and
.
,
.

m edian ns ; the t ail is he tero c er c al .

oo b his rio s sh ros r m is e and d

3
. T h e s p n ill ( y
P o l o d on ) I n t cu
. u th e t u x p e

in to a b r oad fthin sp a tu la te s t ru c tu re p rovided wi th sen se organ s for the detec

, ,

tio n of food At the base of the ros t ru m j u s t in fro n t of the eyes are the n os trils
.
,

ea ch wi th two op en in gs B ehin d the eyes is the sm all sp ira cle The op er cu lu m

. .

c o n tai n s no o p er cu lar bones an d is ex tended i n t o a p oin ted p ro c ess ; the bra n c hio

s tega l m em brane c o n ti n u o u s wi th the o p er cu la

,
has no bra n chios tegal rays
,
.

L if t u p the op er cu lu m an d n o te the n u m ero u s lo n g an d n e gill rakers on the gill

ar ches u sed by the sh in sep ara ti n g food p ar ti cl es from mu d The jaws op en
,
.

widely an d are p rovided wi th minu te tee th The body is n aked No te the ns . . .

The tail is typ i cally he tero cer c al The sh is sin gu larly la ckin g in ex tern al
.

hard p ar ts .
22 LAB ORA OR T Y MANUAL T
FOR VE R E B RAT E ANA OM T Y

4 . or rive r
The b own body is c lo thed in exible
d o g s h ( Amia) The .

s cales similar to those of the p er ch There are lagge s cales on the head The
,
. .

op er culu m an d bran chios tegal m embranes con tain bon es The n os trils have two .

op en in gs the an terior one born e on the tip of the p ro c ess lo ca t ed on the an terior
,

en d of the head the p os t erior o p en i n g si tu a ted in fro n t of the eye

,
There is no .

sp ira cle There is a very lon g dorsal n and a n early ho mo cer cal tail The
. .

sh s tro n gly resem bles the ordinary teleos t typ e .

I . E XTE RNAL ANAT OM Y OF NE CT UR U S

Nectu r us is a salam an der ; it belon gs to the class Amphibia order Ur odel a , .

I t is an ex a mp le of the lowes t lan d ver t ebra tes those whi ch lef t the wa t er ,

a cq u ir ed lu n gs and the air brea thin g habi t and develop ed legs in p la c e of the
-
,

p aired n s O b t a in a
. s p e c i m en and no t e th e fo l lowin g p oin t s .

1 .

B o dy an d s kim The body is t yp i cally ver tebra te in form I t is divisible .

in to head tr u n k and tail as in shes A ne ck is n o t p resen t The S kin is

, , , . .

n aked a n d very slim y wi tho u t s cales or o ther harden ed p ar ts s u c h as are co m

,

m o u ly fou n d in ver tebra t es .

2 .

H ea d The head is broad and a t and has a termi n al m ou th p rovid ed
.

wi th l ips I t bears the u su al three p airs of sen se organs The n os tril s or exter n al
. .

n a r es are a p air of widely sep ara t ed o p en ing s j u s t ba ck of the m ar gin of the u pp er

lip . B y p robing in to the n os t rils de termine tha t they c ommu n i ca te wi th the

m ou th c avi ty by means of Op eni n g s known as the in ter n al n ar es This arran ge .

m en t p erm i ts air to en ter the m ou th cavi ty thro u gh the nos trils and differs ,

g rea t ly fro m th e c o n di t io n fo u n d in the m ajori t y of shes where th e olfa c tory

sa c s en d blin dly and have n o conne c tion wi th the mou th c avi ty I n som e shes .
,

howev er as the ska te there is an ex ternal groove the oronasal groove ex ten ding
, , , ,

from ea ch olfa ctory sac to the m ou th cavi ty ; and by the fu sion of the borders
of thi s groove a clos ed p assage fro m the nos trils to the m ou th c avi ty 15 p rodu ced
( see K pp ,
2 0 6 The
. s m all eyes wi t ho u t eyelids are si tu a t ed on th e sides of
the head The ears as in shes are in tern al only
.
, ,
.

F ro m ea ch side of the p os terior m ar gin of the head sp ri n g three gills ea ch con ,

sis tin g of a frin ge of lam e n ts dep en den t from a dorsal p ro cess They are .

ex ter n al gil l s a n d do n o t c orres p o n d to the gills of shes whi ch are in ter n al .

B e tw een the rs t an d se cond an d the se c o n d and thi rd gills are the gill s l its
,

whi ch op en as in shes in to the cavi ty of the p haryn x The an im al however does

, , .
, ,

n o t p ass wa ter thro u gh the gi l l sli ts b u t res ires by m ea n s of the ex tern al gills
p
whi ch are kep t in con s tan t mo tion throu gh the g en eral s u rfa ce of the body an d , ,

to so m e e x te n t by me ans of its l u n gs .

3 App e n d a g e s
.
The t ru n k bears the two p airs of app en dages These
. .

corresp ond to the p aired n s of shes b u t have evolved i n to typ i c al walkin g

l imb s . E a c h c o n sis ts of the followi n g p ar ts : u pper a r m forear m wr is t an d han d , , , ,

 G E NE RAL STUDY OF T YP I CAL CHORDATE S 23

in the ase of the fore lim b ; thigh s han k an kl e an d foot in the hi n d lim b B o th
c , , , , .

hand an d foo t bear fou r digits ( n gers toes ) al tho u gh ve is the typ i cal ver te , ,

bra te n u mber ; the rs t di gi t is the on e whi ch is missin g The p osi tion of the .

lim bs wi th refer en ce to the body is very p ri mi tive esp ecially in the case of the ,

hind limb an d shou ld be carefu lly s tu died No te tha t the hind limb p roje c ts
, .

ou t a t ri gh t a n gles to the body all of its p ar ts on a p lane p arallel to the g ro u nd

, .

In this p ri mi tive p osi tion the li mb has an an terior or pr ea xial border a p os terior ,

or postaxial border and dorsal an d ven tral su rfa ces In the fore limb however
, .
, ,

the forearm is ben t downward and the hand is dire c ted sli gh tly forward
, This .

al tera tio n of p osi tion is brou gh t abou t chiey by a torsion of the upp er arm ,

whose form er p reaxial s u rfa c e now looks dorsally The p reaxial border of the .

forearm is tu rned m edial ly its p os taxial border la terally As a resu l t of these

, .

cha n ges the anim al is able to lif t i tself to a sli gh t ex t en t above t he gro u nd .

The a t ten ed tail is bordered by a tail n whi ch differs from the n s of shes
in tha t it co n tai n s n o n rays The tail is diphycercal tha t is tru ly symm e tri cal
.
, ,

wi th resp e c t to the ver t ebral c olumn .

4 An u s
. . At the j u n ction of the t ru nk and tail in the m edian ven tral lin e
is the large a n u s or cloa cal ap er tu re wi th mb ria ted borders Amphibia like , .
,

the elas m ob r an chs have a cloa ca

,
.

J . E XTE RNAL ANAT OM Y O F A L IZARD

The liz ards are ty pic al rep tiles class Reptil ia order S qu amata , , .

1
.

B o dy an d s kim The body is c hara c teris ti c ally ver tebra t e in form ,

thi ckes t in the middle ta p ering to ea ch end

,
I t consis ts of h ead n e ck tr u n k .
, , ,

and a long ta il I t shou ld be re called tha t a ne ck is absen t in shes and

.

Amphib ia; its app earan c e is correla ted wi th the assu mp tion of the lan d habi ta t .

The body is comp le tely c lo thed in horny scal es which are thi ckenin gs of the o u ter ,

layer of the ski n ( epiderm is ) and are n o t ho mologou s wi th the s cales of shes .

The s c ales are larger on the head where they are kn own as head s hiel ds .

2.

H e ad The head bears the u s u al three p airs of sen se organ s The
. .

ex tern al nares or nos trils are lo ca ted at the tip of the head and lead as in the ,

Amphib ia in to the m ou th cavi ty where they op en by the in ternal n ar es

,
P os .

terior to the nares are the eyes ea ch p rovided wi th an u pp er and lower m ovable
,

eyelid an d wi th a thi rd eyelid the n ictitatin g membr an e a thin transp aren t

, ,

membrane c on cealed fro m view when n o t in u se in the an terior c or n er of the

eye where it shou ld be so u gh t wi th a for c ep s Halfway be tween the eye an d .

the base of the fore lim b is a sli gh t dep ression in the skin a cross the bo t to m of
whi ch s tre tches a thin m embrane This membrane is the tympan ic memb r a n e .

or dru m membr an e and it covers a ca vi ty the tympan ic cavity or middl e ear

, ,
.

The dep ression may be regarded as the begi nn in g of an exter n al ear The liz ard .

ther efore p ossesses a middle an d a n ex t er n al ear in addi tio n to the i n tern al ear

p res e n t in shes a n d Nectu r u s .

24 LABORA OR T Y MANUAL T
FOR VE R E B RA E ANA OM T T Y

3 Appen d a g e s
.
The t ru n k b ears two p airs of lim bs whi c h hav e the sam e ,

ar t s as those of N t F ive di i t s th t y i al ver t ebra t e m ber are

p
p ec u r us g e c . n u , ,

p rese nt on ea c h li m b ; they t ermi n a te in horn y c laws

The s tu den t sho u ld .

n o t e tha t the li m bs dep ar t s till fu r ther fro m the p ri m i tive p osi tio n des c ribed in

c onne c tio n wi th Nectu r u s th u s lif tin g the anim al above the gro u nd to a grea ter
,

ex ten t I n the hi n d li m b the thi gh s till ex ten ds ou t at righ t angl es to the body
.
,

b u t is sli gh t ly twis ted on its lo n gi tu di n al axis so tha t the original dorsal s u rfa ce
is be co mi n g an terior The shank is dire c ted ven trally and p os teriorly an d has
.
,

u nderg o n e the sam e sor t of t orsion as the thi gh The foo t re tains the p rim i tive .

p osi tio n I.n th e fore li m b the u pp er a rm is ro t a t ed in the o pp osi t e d ire c tion

fro m tha t observed in the thigh so tha t the original p reax ial s u rfa c e is now,

dorsal ; the u pp er arm is dire cted p os teriorly The forearm an d han d are .

di re c ted downward and forward and are so ro ta ted tha t their ori gi n al p reaxial
,

borders look inward in s tead of forward .

4 An u s
. On the ven tral side of the t ru nk behi n d the bases of the hind
.

limbs is the anu s or cloa cal ap er tu re I ts shap e a transverse sli t is chara c ter .
, ,

is tic of li z ards an d s n akes .

K . E XTE RNAL ANAT OM Y OF THE T URTLE

The tu r tle is a rep tile belongin g to the order Chel on ia Ob tain a sp e ci m en .

a n d ex a m i n e .

1 . B o dy an d s kin The form of the body is c on siderably m odi ed from

.

the ty p i cal ver tebra te shap e I t is divided in t o head ne ck t ru nk and tail

.
, , ,
.

The head is S imilar in form to tha t of o ther rep tiles b u t the ne ck is u nu su ally ,

lon g an d ex ible ; the tru nk is rem arkably a t an d broad and the tail diminished ,

in diam e ter and length The tu r tle is thu s one of those form s in whi ch as in
.
,

the S ka te the transverse axis is elo n ga t ed while the a n tero p os terior and dorso
,

ven tral ax es are shor ten ed The skin of the legs tail and o ther exp osed p ar ts
.
, ,

of the body is p rovided wi th s mall horny s cales or horn y thi cken in gs as is chara c
teris tic of rep tiles The tru nk is in closed in a hard shell the o u ter su rfa ce of
.
,

whi ch consis ts of grea tly enlarged horny s cales deni tely arran ged The sk in ,
.

of the head in m os t of ou r c o mm on tu r tles is bare b u t in som e tu r tles is m arked ,

of f i n to larg e head shi elds .

2 .

H ea d The a n terior en d of the head is p oin ted an d eleva ted an d bears
.

a t its tip the two ex tern al nares c lose to g e ther ; this p osi tion of the n os t rils
,

en ables the anim al to brea the air wi th on ly a sligh t exp osu re of the head
above wa ter The jaws are c lo thed wi th hard horny b eaks tee th bein g absen t
.
, ,
.

The large eyes are p rovided as in the l iz ard wi th upp er and lower eyelids an d a
n i c ti ta ti n g m em brane lo c a ted in the an t erior c orn er of the eye
,
Ju s t p os terior .

to the a n gles of the m o u th can be observed a c ir cu lar area c overed by skin ; this ,

is the tymp ani c mem bran e ben ea th whi ch is lo ca ted the cavi ty of the m iddle
ear This m embran e is n o t however su nk b elow the su rfa c e as in the liz ard
.
, . ,
.
26 LAB ORA OR T Y MANUAL FOR VE R E B RA E ANA OM T T T Y

su ch an ex ten t tha t the tym p anic m em bran e is n o lo n ger visible The skin .

arou n d the e n tran c e to the m ea tu s tends to el evatp as a fold ; this fold toge ther
wi th the m ea tu s con s ti tu tes the extern al ear .

3 Tru n k
. The t ru nk is very rm an d inexible owi n g to a fu sio n of the ,

bon es of the ba ck P ass you r n gers alon g the ba ck of the p lu cked bird an d
.

feel the skele to n F eel also in the m edian ven tral lin e of the tru nk the p ro
.

je c t in g keel of th e breas t bo n e t o w hi c,
h are a t t a c hed th e g r e a t wi n g m u s c l e s .

I t is the p resen c e of these mu s c les the breas t of the bird whi ch p rod u c es the

, ,

p l u m p c o n t o u r of th e t ru nk The t ru
. n k bears t h e tw o p airs of li m bs of whi c h ,

the a n t erior p air is re m arkably m odied in to win gs or org a n s of i gh t The .

hin d li m bs have also u n dergone co n siderable modica tio n as a res u l t of the

bip ed m ode of walkin g u sed by birds .

The p ar ts of the wi n g whi ch are homolo gou s to those of the fore lim b of
,

o ther ver tebra tes sho u ld be s tu died o n the p lu cked bird In the folded co n di
,
.

ti o n the se c tio n s of the wi n g m ake angles wi th ea ch o ther l ike the le t t er Z .

The u pp er ar m is shor t dire c ted p os teriorly and sligh tly twis ted on its axis so
, ,

as to bri n g the p reax ial m argin o n the dorsal side The forearm is lo n ger an d .

dire cted forward The wris t an d han d are fu sed to ge ther an d the whole is con
.
,

sid er ab l y elo n ga t ed a n d di re c t ed ca u dad There are b u t three digi ts whi ch

.
,

are re garded as the se co n d third an d fou r th The se co n d di gi t is the p roje c tio n

, ,
.

fo u n d j u s t below the join t be tween forearm an d wris t ; the third di gi t form s the
term in al p oi n t of the wi n g ; the fo u r th canno t be see n ex ter n ally When the .

win g is ex ten ded its p ar ts have n early the p rimi tive p osi tio n des cribed u n d er
,

Nectu ru s The g rea t q u ills of the wi n gs are known as r emiges : those of the
.

han d are c alled pr imar ies ; of the forearm secon daries ; and of the u pp er arm , ,

hu meral s . The p ri m ari es di ffer fro m the o thers in tha t the sof t p ar t of the fea ther
is wider on the p os terior side of the c en tral axis than on the an terior side The .

remi ges are born e o n the p os ta xial m argin of the win g an d the d eep large fea ther ,

folli cles exp os ed by their rem oval shou ld be no ted on the p lu cked bird .

The hin d lim b is p ar tially c lo thed wi th fea thers an d p ar tially wi th horn y

s c ales id en ti cal wi th those fou n d in rep tiles The dig i ts of whi ch there are b u t
,
.
,

fo u r the fth bein g absen t term in a te in claws The p osi tion of the hin d .

li mb wi th referen ce to the body is grea tly al tered The whole li m b in s tead of .

,

ex t en di n g s trai g h t o u t la terally fro m the body is dire c t ed ve n trally th u s raisi n g, ,

the a n im al high abov e the gro u n d In order to a chieve thi s resu l t it is obvio u s
.

tha t the l im b m u s t have been ro ta ted 90 fro m the p rim i tive p osi tion so tha t the

ori gi n al dorsal su rfa c e n ow fa c es an teriorly tha t is has be com e p reaxial The ,
.

toes are c o n seq u e n tly dir e c ted forward wi th the ex cep tio n of the rs t whi ch
, ,

thro u gh a s ec o n dary m odi ca tion c o n n e c t ed wi th the p er chi n g habi t p oi n ts ,

p os t eriorly .

T il The t ail s t m bears a half c i r l of lar e q u ills k n ow n as r ectr ic es

4 a u p c e g
-
. . .
,

Un der the base of the tail is the anu s or c loa cal ap er tu re a trans v erse op en in g ,
 G E NE RAL STUD Y OF TYP I CAL CHORDA E S T 27

wi th p ro t ru din g
lip s On the dorsal Side j u s t in fron t of the base of the tail
.

will be seen by lif ti n g up the tail cover ts or on the p lu cked bird a p ap illa the , , ,

op en ing of the u ropygial gl an d fro m whic h the b rd ob tai n s oil for preeni n g
i ,

its fea thers .

M . E XTE RNAL ANAT OM Y OF A MAMMA L

Fo r this pu rp ose ei ther the ca t or the rabbi t may be u sed . B o th are

ma mm als belo n gi n g to the c lass M ammal ia . The rabbi t is a rod en t , the ca t a

c arn ivore .

1 .

B o dy for m an d s kim The body is divisible i n to head n e ck t ru n k an d , , ,

tail . I ts p ro p or tions dep ar t so mewha t fro m the typ i cal ver tebra te form in the
larger S iz e of the head and the red u ctio n of the tail the la tter fea tu re p ar ti cu larly ,

no ticeable in the rabbi t The body is c lo thed wi th closely s et hairs formi n g a

.
,

c overin g of fu r c hara c teris ti c of m a mm als

, Up on the head are a nu mber of .

esp e cially lo n g an d s tou t hairs the whis kers or vibriss ae whi ch serve as imp or tan t
, ,

organs of tou ch .

2 .He a d T he larg e siz e of the head is du e to the g rea t develop m en t of

.

the brain wi t hi n it The head may c o n se qu en tly be divided i n to an an terior

.

fa c ia l re g io n in fron
,
t of t h e eyes and a n e n lar g ed p os t erior ,cra n ia l re gion .

The mo u th is p rovided wi th well develop ed l ips The u pp er lip is clef t in the -

.

c e n ter deep ly so in the rabbi t exp osi n g the in cis or t ee th

, ,
The extern al n ares .

are large an d elon ga ted overh u n g b y the mobile nose The eyes have u pp er
,
.

an d lower lids an d a ni c ti t a t in g m emb rane The la tter shou ld be so u gh t wi th .

a for c ep s in the an terior corner of the eye and drawn over the ey eball The .

ears are p rovided with a lo n g and ex ible ex tern al fold the pin n a whi ch sp rin gs , ,

from the rim of an op ening the exter n al au ditory mea tu s whi ch des cen ds in to
, ,

the in terior of the sk u ll P inn a a n d m ea tu s con Stitu te the ex tern al ear

. .

3 .T r u n k The t ru nk is divisible i n t o an an t erior c hes t or thor a x s u pp or ted ,

by the ribs and a pos terior a bdomen On the ven tral su rfa ce of the trun k are
,
.

fo u r or ve p airs of tea ts or n ippl es in fem al e sp e ci men s ; they are the Op enin gs

of the milk glan ds or mammar y glan ds The tru nk bears two pairs of lim bs .
,

which co n sis t of the sam e p ar ts as o ther ver tebra te limbs The u pp er se c tio n

.

of ea ch lim b tha t is thigh or u pp er arm is m ore or less in clu d ed in the t run k .
,

The lim bs termina te in c lawed digi ts ve in fro n t fou r behin d the rs t hind , , ,

to e bei n g absen t The claws of the cat and its rela tives are re tra c tile
. .

The limbs have u ndergon e a marked chan ge fro m the p rimi tive posi tio n .

In s tead of ex tendin g la terally fro m the body they p roje c t ven trally an d are
elon ga ted so tha t the body of the a n i mal is c arried hi gh above the g ro u n d
,
This .

c ha n ge has involved a ro ta tion of 9 0 in ea ch limb The hi n d limb is ro ta ted

.

forward so tha t the ori gi n al dorsal su rfa ce is a n terior and the toes p oin t fo r ward .

The knee or join t be tween the thigh an d shank is likewis e dire c ted a n teriorly .

The fore lim b on the co n trary has ro ta t ed pos teriorly so tha t the original
, , ,
28 L AB ORA OR T Y MANUAL F OR VE R TE B RATE ANATOM Y

dorsal s u rfa c e fa ces p os teriorly and t he p reaxial s u rfa c e fa ces la terally Con se .

qu en tly the elbow or joi n t be tween u pp er a r m an d forearm is dire c ted c au dad

, .

AS a r es u l t of the ro ta tion of the u pp er a rm the to eSwou l d p oin t p os teriorly b u t ,

by an addi tional torsion they have been brou gh t aro und to the fron t a gai n .

This torsion i n volves a c rossin g of the two bon es of the forearm the dis tal en d ,

of the p reaxial bon e bein g brou gh t i n tern al to the dis tal en d of the p os tax ial
bon e ( see F ig 2 an d read R p . This p osi tion of the forearm 1 5 known as
, .

BI ND -LI ME

Dia p f po t f
FI G . 2 .
gram s t o ill u s tra te the t o rsio n o f the l imb ; reaxial s ur ace whi te , s axial sur ac e

b l ack . A, p rimi tiv e p

o sition o f the limb s , s een from ab ove ,
the limb s ext en din g a t right a n gl es t o the
b o dy, the p f
reaxia l su r a ce f acin g a n t erio rl y, the p o s taxia l p os terio rl y . B, p o sition of the limb s in
m amm als , from the sid e ; the limb s e ten d verticall y b el o w the b o dy ; the upp er a rm ha s rota ted
s een x

o u t war d a n d b ac kwa r d s o tha t the p rea xial su r face n o w faces l a t erall y ; the fo r ea rm ha s r o ta t ed fo rwar d

a g ain r esu l tin g in a c r o ss in g o f its b o n es ; the hin d limb ha s r o ta ted ou twar d a n d forwar d s o tha t the
,

pos taxial su rface faces l a terall y ( From Fl ower s Osteol ogy of the Mammal ia )
.

.

the p p osi
ron e tion a n d is i m i t a t,
ed in t h e h u m an a r m when th e a r m han g s by
t he side wi th the ba ck of the ha n d dire c ted forward I n this p osi tion the c rossi n g .

of the two lo n g bon es of the forearm can b e fel t If n ow the a rm is raised side .

wise shou lder hi gh wi th the p alm fa ci n g forw ard the two bon es re tu rn to the
, ,

p r im i tive p arallel p osi tio n k n ow n as the s u pi

,
n e p osi tion Th u s m a n c a n c han g e .

hi s forearm fro m the p ro n e to the s up in e p osi tio n b u t in m os t m a mm al s the fore

-
,

a rm is x ed in the p ro n e p osi tio n .

The p osi tion of the p ar ts of the foo t in walki n g is di fferen t in di eren t

m am mals The rabbi t a n d the c a t walk on the di gi ts wi th the rem ai n d er of
.
,

the ha n d and foo t eleva t ed This t yp e of g ai t is k n own as digitigrade M an

. .

walks on the whole sole of the foo t the p ri m i tive m e thod k n own as pl an tigrade
, ,
.

H orses a n d c a ttle a n d o ther u n gu la t es walk on their nails whic h are broaden ed ,

in to hoofs a m ode of walking c all ed u n gu l igr ade

, .
 G E NE RAL STUDY OF TY P I CAL CHORDA J
P S
'

29

ven tral m edian lin e j u s t in fron t of the base of the tail On ea ch S ide of the anu s .

in the rabbi t is a deep hairless dep ression the in gu in al or perin eal s pace on to
, , ,

whi ch Op en the in gu in al gl a n ds (n o t visible ex tern ally) whose se cre tion p rodu c es

the odor c hara c teris ti c of the anim al In the fem ale the u ro geni tal op en in g is .
,

S i tu a t ed imm e di a t ely an t erior and ve n t ral to the an u s it is si milar in a earan e

; pp c

to the a n u s I t is in closed by a fold of skin ex tending aro u nd the r im ; this fold

.

is n am ed the greater l ips or l abia majora The labia toge ther wi th the u rogeni tal .

ap er tu re cons ti tu te the vu lva In the m ale rab bi t there is a hillo ck in fron t of .

the an u s I n the c en ter of this is u su ally visible a p oin ted p roje c tion the end
.
,

of the pen is or organ of copu la tion ; the p enis bears a t its tip an op eni n g the ,

u r ogen ital aper tu r e The hillo ck of skin whi ch folds u p abou t the p enis is n amed
.

the prepu ce or fores kin To each side of the p repu c e and ex tendi n g forward is
.

an oval swellin g c au sed by the m ale gonad or tes tis whi c h is i n c losed wi th
, in it , , .

The dou ble p ou ch of the body wall which in c loses the two tes tes is n amed the
s cr otu m or s cr otal s ac In the m ale c at there is a p air of ro u n ded emi n en ces
.

an terior to the anu s ; ea ch eminen ce c on tains a tes tis or male go n ad an d the ,

do u ble p o u ch of the body wall inc losing the tes tes is nam ed the s cr otu m or s cr otal
sac . An terior to the s c ro tu m is a hillo ck of S kin the pr epu ce or for es kin In , .

the c en ter of the p rep u c e is an op ening whi c h is n o t however as m i gh t be su p , , ,

p osed the u ro
, g eni t al o p enin g I t is m erely the dep ressio n lef t by the wi thdrawal
.

in to the p repu c e of the pen is ; the p enis is generall y so far wi thdrawn in m ale
ca ts as to be invisible ex ternally .

N . S UMMAR Y
Ou r s tu dy f eatu res of rep resen tative ver tebra tes may b e u tiliz ed to direc t
o f the ex tern al

the s tu d en t s a t ten tion to the foll owin g p oin ts

.

I T he ver tebra te bo dy is typ icall y fu s ifo rm tha t is modera tely thick throu gh the tru n k
.
, , ,

tap erin g towar d ea ch en d In ev ery g rou p o f ver teb rates fo rms may b e fou n d which deviate
.

from this typ ical shap e b u t su ch deviation s b ear n o r el ation to the position of the an imal in
,

the v er teb ra te scal e b ein g r a ther adap ta tion s to p ar ticu l ar modes o f lif e
,
.

2 T he bo dy is divided in to head trun k an d tail in the l owes t vertebra tes

. A n eck is
, ,
.

a dd ed in l an d ver te r atesb .

3 . hea d ten ds to in crease rel atively in S ize an d the tail to decrease as on e ascen ds the
The
v e t eb a te s e ies
r r T he fo rm er chan g e is a sso ciated with in c eas e in the s iz e of the b ai ; th
r . r r n e

l a t ter with g r ea ter sp eed a d ag ility o f movemen t n .

4 The s k in of ver teb a tes is comm onl y cl o thed with p ro tec tive s t ru c tu r es s ch as s cal
. r ,
u es ,

fea thers o r hai s These are more compl ex in s tru c tu e in the higher ver teb ates an d be tt
,
r . r r er

tted fo r k eep i g the body warm n .

5 T he
. h ea d th o u gho u t b ea s th ee p ai s o
r f s e se o g an s O f th e
rse the eyres u n de g o r n r . r

littl e cha g throu ghou t the s eries

n e T he ol fa c to ry s a cs a e blin d in he an d op en to
. r s s

the e te io r on ly by mea s of the ex tern al n ar es

x r ,
As s oo ho wever as ve t eb a tes l ef t the
n . n, ,
r r

a q atic en vi o m en t i tern al
u r n a es w e e d evel op ed co
,
n ec tin g the o l fa c t ry
n r cs with the
r ,
nn o sa
3 0 L AB ORe
s RY MANUAL F OR VE R E B RA E ANAT OM T T Y
mou th c avity . This a rr an g e men t p ermits the ani m al to b eathe withou t op in g the mou th
r en .

a d ecid ed ad v an t ag e in air - b r ea t hin g an imal s ,

s in c e ther e by the d yin g of the mou th cavity is
r

avo id ed . I n the l ower b

v er t e r a t es an in t ern al ear onl y is p r es en t. T o thi s is a dd ed , b egin
n in g with the an u r an middl e ea r cl o sed ex tern ally by the tymp an ic memb ran e
Amphibi a , a , .

In An u r a an d man y r ep til es the tymp an ic membr an e is l ev el with the su rfa ce o f the hea d b u t ,

in so me r ep t il es it b eg in s to s in k b el ow the s u rfa ce I n b ird s an d mammals it has descen ded .

d eep ly in to the head , for m in g a n arr ow p ass ag e , the ex tern al au dito ry mea tu s Ar ou n d the .

ex tern al rim o f the m ea tu s in mammal s the s kin el eva t es to f orm a so u n d ca tchin g device , the -

pin n a P in n a an d meatu s con s titu te the extern al ear

. .

6 T he gil l slits an d gill s p res en t in the shes an d l ower Amphibia dis app ea r in the adu l ts
.

o f the higher Amphibia an d a ll fo r ms abo ve them This is du e to the assu mp tion of the air .

brea thin g habit .

7 T he tr un k
. bears two pairs of appen dages Thes e are n s in shes b u t b ecome limbs .
,

in all verteb ra tes above shes S ta g es in this tran s fo rma tion are very imp erf ectly kn own
. .

The p a ts o f the limb s ar e the same throu gh all of the v e t eb a tes al thou gh they a e s u bj ect
r r r ,
r

to con s ide abl e modi ca tion

r T he mos t primitive limb s in s tru c tu e fo m a d po s itio n with
.
,
r ,
r ,
n

ref ere ce to the b ody oc cur in the u rod el e Amphibia

n I n higher verteb ra tes the p osition of the .

l imbs is al ter ed by b en din g an d to rs ion r es u l tin g in an el eva tion of the body ab ove the g ou n d
,
r

with a corr es pon din gly mo r e r ap id p og ess io As a s till fu r ther aid to r ap id moveme t the
r r n. n

dig itig rade o r un gulig rade mod e o f wal king has b een adop ted in man y cases L oss o f digits .

is qu ite co mm on amon g ver teb ra tes ; the mis s in g dig its ar e n ea ly al ways the r s t o r l as t o es r n ,

rarely the middl e on es .

8 In n ea rl y all ver tebrat es e cep t m amm al s the in tes tin e an d the u r og en ital du cts op en
. x

in to a common chamb er the cl oaca which commu n ica tes with the e terior by a s in g l e op e in g
, ,
x n ,

the an u s o r cl o acal ap e tu r e I n all pl ac e tal mamm al s the in tes ti e an d the u og e ital sys tem
r . n n r n

o p en by s ep a at e ap er tu r es the u og e ital o p e in g b ein g s itu ated a l ways an te io r to the a u s

r ,
r n n r n .

The term an u s therefore does n o t have the same s ig n i can ce in mammal s as in o ther
, ,

vert ebra tes .

 IV . G E NE RAL FE ATUR E S O F CH O R D ATE D E VE L O P M E NT
is imp ossible to u nders tand the comp ara tive ana tomy of ver tebra tes
Sin c e it ,

whi ch form s the m ain obje c t of s tu dy of this co u rse wi thou t som e knowled ge ,

of the way in which a ver tebra t e develop s it is necessary tha t the s tu den t learn ,

som e of the elem en tary fa cts abou t ver tebra te developm en t E very s tu den t .

mu s t read ei ther K p a ges 7 1 6

or W p ages 6 0 71 He will be exp ec ted to
, , , .

know the c on ten ts of these p ages thorou ghl y and to be able to answer a t all
tim es q u es tions rela ti n g to the m .

A . THE CH ORDA TE E GG

The m anner of developm en t of the Chorda te egg is dep enden t u p on the

amo u n t of food m a terial or yol k whi c h it c o n tain s On this basis Chorda t e
, ,
.

eggs are classied as follows : ( I ) I s ol ecithal eggs wi th li ttle yol k ; Amphioxus ,

and m amm als ( 2 ) Tel ol ecithal eggs wi th total c leavage There is a modera te
. .

FI G 3 T hr ee ty es o f Chord a te eggs
. . pT he b l ac col orin g r e . k p resen ts the yo lk . A , isol ecithal
eg g with s mall a n d ev en l y dis trib u ted yol a rt icl es kp
B , t el ol ecithal . eg g with to tal u n eq u al cl ea va g e,

k
with the yol mor e a b u n d an t in the veg etal than in the a n imal hal C, tel ol ecithal egg with m ero
f .

p p
b l a s tic cl eava ge, con sis tin g c om l etely o f yol k exce t for the small g ermin al disk o f roto l asm a t o n e p p
p l
o e . T h f h
e siz e o t e g er m in al dis k in the gur e is g rea tl y ex agg er a t ed with re s ec t to the S iz e o f the p
yol k .

amoun t of yOlk whi c h a ccu mu la tes in o n e half of the egg and re tards its develop -

en
m t ; Amphibia c c y los t o m es a,
n d g anoid shes , (3 ) T el ol ec i tha l e g g s . wi th
mer ob l as tic c leava e There is an enor m o u s a m o u n t of y olk and t he p ro t o p las m
g . ,

is redu c ed to a sm all disk whic h oa ts on the s u rfa c e of the yolk T hi s spo t is .

c alled the ger min al dis k Su c h e gg s are c hara

. c t eris ti c of birds re p til es an d , ,

t eleos t s The r
t u
. e e gg in t he c ase of birds is th e y olk th e w h i t e b e in g m erel y ,

a nu tri tive en velop e .

These three t yp es of e gg s are illu s tra t ed diagrammatically in Figure 3 .

( S ee f u r ther H pp 1 4 1 , , .

31
3 2 LAB ORAT OR Y MANUAL T
FOR VE R E B R A E ANA OM T T Y

B . THE CLE AVA E G OF TH E E GG AND THE F ORMA TI O N OF THE B LAS T ULA

Develop m en t begin s by the division of the egg in to two fo u r etc c ells u n til , , .
,

a l arge n umb er of c ells has been p rodu c ed This p ro c ess of divisio n of the egg

.

is call ed cl eavage and the way in whic h it o ccu rs dep en ds o n the amo u n t of yolk
,

which the egg con tai n s I t shou ld be u n ders tood tha t the yolk is i n er t m a terial
.

an d t ha t the p ro c ess of develo pm e n t is c arried o u t o nl y by the livin g p ro to

p las m i c p or tions of the e gg .

1 Hol ob a s tic e qu al cl eava g e

.
I n the c ase of isole ci thal e gg s the en tire
.

egg di vides a n d p rod u c es a n u m ber of app ro x im a t ely eq u al c ells S u c h c leava ge .

is said to be hol obl as tic and equ al The c ells as they i n crease in nu mber gradu ally
.
, ,

wi thdraw fro m the c en ter a n d arra n ge them selves in a sin g le layer on the su rfa c e ,

th us p rodu ci n g a hollow ba l l of c ells Thi s ball is c alled the b l as tu l a; its cavi ty

.

is kn own as the s egmen ta tion cavity or b l as tocoel Su ch a blas tu la is p rod u c ed .

in the developm en t of Amphioxu s Cleavage an d form a tion of the blas tu la

.

in Amphioxu s are ill u s tra ted in F i gu re 4A Simi lar gu res will also be fo u nd in
.

the s tandard tex tbooks as P an d H K W and H ,

S tu dy also the m odel s of
, , ,
.

the cleava g e of Amphioxu s p rovided in the labora tory .

2 Hol ob as tic u ne qu al cl eava g e

.
Thi s typ e o c cu rs in those telole ci thal
.

eggs whi ch con tain a modera te amo u n t of yolk The half of the egg which con .

ta ins the m ajori ty of the yo l k is c alled the vegetal hemis phere; tha t whi ch
c o n tai n s the m ajori t y of the p ro to p las m is the an imal hemis pher e The early .

c leavage p la n es are s hif t ed toward the anim al hem isp here and fu r ther the , ,

c leava g e p ro cesses are delayed in t he ve ge tal hemis p here owi n g to the p rese n c e

of the in er t yolk I n co n sequ en ce of these two fa c tors the c ells p rodu ced in the
.
,

anim al hemisp here are sm aller an d more n u m ero u s than those of the vege tal
hemisp here al thou gh the en tire egg c leaves Su ch cleavage is holoblas ti c b u t
, .

u n equ a l The c ells wi thdraw fro m the c en ter p rod u ci n g a blas tu la wi th a

.
,

som ewha t redu ced b l as tocoel an d a wall sev eral layers of c ells thi ck The c ells .

of the blas tu la are of u n equ al siz es g rading fro m the sm alles t a t the animal pole
to the larges t a t the ve ge tal p ole .

This typ e of developm en t is chara c teris ti c of Amphib ia I t is illu s tra ted in .

F igu re 43 ; sim ilar gu res are p resen ted in vario u s t ex t books as P an d H K , , ,

and N S tu dy al so the m odels of the cleavage of the amp hibian egg p rovided
.
,

in the labora tory Then ob tain a se c tion thro u gh an am p hibian egg in the
.

blas tu la s tag e and ex ami n e u nder the low p ower of the mi cros cop e The blas tu la .

is a hollow sp here whose wall is comp osed of two or three layers of cells The .

wal l of the anim al hemisp here is thin and c o n sis ts of s m all c ells ; it is the fu tu re
dorsal sid e of the embryo The wall of the vege tal hem isp h er e is mu ch thi ck er
.

than tha t of the anim al hemisp here an d is c om p osed of lar g e c e lls lade n wi th yolk ,

and wi th in di s tin c t bou ndaries ; it is the fu tu re ven tral side The blas to coel is .

sma l ler than in t he blas tu la of Amphioxu s a n d is disp la c ed dorsally owin g to , .

the thi c kn e s s of t he ven tr al wall Draw showin g o u tli n es o n ly of the ce lls

. . .
 34 T
LABORA OR Y MANUAL FOR VE R E B R A E ANA O M T T T Y

M e rob l as ti c c leavag e
e s on t i In
i n lar e q u a ti ties of yolk o ly
3 .
gg c a n
.
g g n n

the s m all germi nal disk u nder goes cleava g e This ki n d of c leavage is c alled .

mer oblas tic As a resu l t a mi n u t e disk of c ells is p fodu c ed on the su rfa ce of the
.
,

rela tively en orm ou s yolk A sli gh t sp li t app ears be tween the disk and the yolk
.
,

a n d this c orresp o n ds to the se gm e n t a tio n c avi ty of o ther develo p i n g e ggs ; this

s tage is co n sequ en tly the blas tu la s tage M eroblas tic c leavage is illu s tra ted in .

F i gu re 4C The disk of cells p rod u ced by meroblas ti c cleavage is cal led the
.

blas toderm ; in the fu r ther develop m en t it exp ands over the s u rfa ce of the yolk
which it even tu ally in closes .

C . F O RMA TI ON OF TH E G A S TR UL A
I . In e gg s of the Amphioxu s typ e In s u ch eggs the vege tal he misp her e .
-

begins to ben d in ward and con tin u es this p ro cess of in vagin ation u n til its wall
c o m es in c o n ta c t wi th the wall of the a nim al hem is p here An em b r yo wi th a .

wall two c ell lay ers thick is th u s p rod u c ed I t is called a gas tr u l a The o u ter
-
. .

layer is nam ed the ectoder m and the in n er layer the en toderm B e cau se of their .

imp or tan ce in the s u bsequ en t developm en t these layers are referred to as the ,

rs t two ger m l ayer s The hollow tu be of en toderm is called the ar chen teron or
.

p r im it ive i n t es tin e; th e c avi t y of th e g as t r u la is t h e c avi t y of the ar chen t ero n or

g as tr o co el ; a n d th e o p e m n g of t he ar c hen t ero n t o th e e x t erior is th e b l a s top o re .

No te tha t the blas to coel is elim in a ted in the p rod u c tio n of the gas tru la The .

form a tion of the gas tru la of Amphioxu s is illu s tra ted in Fi gu re 5A an d also in ,

the variou s t e x t books as P an d H W K an d N

, ,
S tu dy fu r ther the m odels
, ,
.

ex hibi ted in the labora tory .

2. In e gg s of the amp hibi an typ e In t hese eggs g as t ru la t io n is so m ewha t

.

m odied by the p resen c e of i n er t yolk in the vege tal h emisp here I t is a c co m .

p l is h ed p ar t ly by t he i n va gi n a tio n of t h e e n tod e rm p ar t i c u l arly a t t h e dorsal lip

of the blas to por e an d p ar t ly by the exp an sion of the e ctoderm ven t rally p u shi n g
,

the en t oder m in to the i n t erior The res u l t is the sa m e as the for egoi n g a gas t r u la
.
,

b ein g form ed A s m all po r tion of the i n clos ed yolk b earin g cell s co mm o n ly r e

.
-

m ai n s for so m e tim e p ro t ru din g t hro u gh the blas t o por e an d is c al l ed the yol k pl u g ,

.

The form a tion of the amphi bian gas tru la is illu s tra ted in F i gu re 5B and in ,

P an d H N and o ther t ex tbooks

, , S tu dy fu r ther the m odels of amp hibia n
.

developm en t illu s tra tin g thi s s tage no tin g esp e cially the sagi ttal se c tion of the
,

g as t ru la . The n ob tain a slide bearing a sa gi ttal se c tion of the g as tru la and s tu dy

wi th the low p ower of the mi cros cop e The g as tru la is sligh tly elo n ga ted in the
.

a n terop os terior dire c tio n The side wi th the thi n n er wall is the dorsal side ;
.

tha t wi th the thi ck wall the v en tral side ; the en d wi th an o p en i n g is the p os terior
,

en d ; the opp osi te en d is a n t erior The wall c o n sis ts of two lay ers ea ch com p osed
.

of m ore than on e shee t of cells The o u ter an d thinn er lay er is the e c toderm
.
,

u n iform in wid th over the whol e em bryo The in n er layer is the en toderm
.
,

sep ara t ed from the e c toderm b y a sli g h t sp a c e and very thi ck ven trally where , ,
 d
en to er m

in vaginatio n

archen teron

ec to er d m
b las tocoel

b l as toco e

d
e n to er m

ec to erm d
archen teron

gastr ocoel
b las tocoel

C
2

FI G
5 Gastrul a tion p
an d formation o f the en tod erm in the t hr ee ty es o f chorda e eg g All
. .

shown in medi an sa git ta l s ection A , Amphioxus;

. t s .

b egin nin g o f the in vagin ati o n ; 2 , i n va in ati o n

.

g
p p
c om l eted ; 3 , com l eted g as trul a havin g a wal l o f t wo l ayers , e t od erm a n d en tod erm , a n d a n i ter al
c n n
c avity , the g a s trocoel . p p
B , am hib ian ; I , b egin n in g o fthe i n va gin ati on ; 2 , ro r es s o fthe mvagin ati on
g
p p
a ccom anied b y d o wn war d g ro wth o f the ec to d erm ; 3 , com l eted g as tr l a , W I th very thi ck en tod erm
u
o

ven tr all y . p p
C, re til e o r b ird ; I , os terio r margin o f the b la s tod erm b egi n mn g t o t u rn u n der ; 2 , c on
.

p
t in ua tio n o f the in v a gin a tion ; a slight s lit b etween en t o d erm a n d yo l k co n s ti tu tes the g as tr oc o el .

p
I n A3 , B 3 , an d C2 the an t erio r en d o f the emb ryo is t o the l ef t, os t eri o r en d t o the ri ght , d orsal su rface
ab o v e , v en t ral b el o w .

Z
(A from P a rker an d Has well s Textb ook of ool ogy, a f ter Hatschek , cou rt esy of
p
t he M acmill an Com an y ; B from Kellicott s Gener al E mbryol ogy, cou r tesy o f Hen ry Hol t an d Com

Pa n Y)

the blas top ore E c toderm an d en toderm are c on ti n u ou s a t the rim of the blas
.

to p ore A p or tion of the e n tod erm the yol k pl u g p ro tru des thro u g h the
.
, ,
3 6 LABORA OR T Y MANUAL FOR VE R E B RA E ANA DOMYT T '

blas top ore an d nearly o cc lu des the op enin g . M ake a diagra m of the se c tio n ,

col or in g ectoder m bl u e an d en toder m yell ow .

3 .I n m e robla s t
i c e gg s 1 p s u ch eggs the form a tion of the en tod erm will
obviou sly be a tten ded by dii cu l ties I t seem s to be a ccomp lished in two ways :.

( )
a by a p ro c ess of i n vol u tion or i n va gin a tio n i n whi ch the c ells a t the fu tu re ,

p os t erior end of th e blas t oder m t u r n u nder and g row forw ard ; ( b ) by a

p ro c ess
of del amin a tion in whi ch the cells on the u n der S ide of the blas tod erm a rrange
,

the m selves to form a lower layer In differen t form s the p ro p or tio n of en toderm
. .

form ed by ea ch me thod varies E ven tu ally the two kin ds of en toderm u ni te

.

to form a sin gle shee t of cells The c avi ty of the ar chen teron is very sm all in
.

m eroblas ti c egg s consis tin g of a clef t be tween the en toderm an d the yolk and
, ,

thq b l a s top or e is red u ced to a sli t a t the p os terior en d of the blas toderm Af ter .

g as tr u la tion has o c c u rred th e blas toder m c o n sis t s of two layers an o u ter e c toderm ,

an d an in ner en toderm ; it lies o n the s u rfa c e of the yolk and by p rolifera tio n a t

its m argins g rad u a l ly s p reads ou t over the yolk even tu ally in clos in g it , .

The gas tru la tion of meroblas ti c eggs by the invagi n a tio n p ro cess is ill u s tra t ed
diagra mm a ti c ally in F i gu re 5C .

D . F ORMATI ON OF THE TH IR D G E RM L A YE R ,
THE NE UR A L T UB E ,

AND THE N OT O CH ORD

These p ro cesses o ccu r p ra c tically sim u l tan eo u sly b u t wil l be des cribed sep a
r a tel y .

I . In Amphioxu s Af ter the

emb ryo has a ttai n ed the g as tru la s t age it
elo n ga t es and p resen ts a a tten ed dorsal s u rfa ce a ro u n ded ven tral su rfa ce an d , ,

re co gn iz able an terior and pos terior end s ( see Fig 5A F rom the dorsola teral .

regio n s of the en toderm whi ch it will be rem embered form s the in ner tu be
,

of the gas tru la holl ow po u ches begin to g row ou t in p airs These p ou ches
,
.

are called the coel omic s acs or mes odermal pouches The walls of the p o u ches .

c o n s ti tu t e the mes oder m or thir d ger m l ayer which u n like the e c toderm an d
, , ,

en toderm consis ts of two wall s The p ou ches grow la terad an d ven trad l li n g
, .
,

the s p a c e be tween e c toderm an d en toderm The ou ter wall of the p ou ch es in .

co n t a c t wi th the e c toderm is called the s oma tic or par ietal m esoderm ; the inn er

wa ll in co n ta c t wi th the en toderm is the s pl an chn ic m esoderm The cavi ty of .

the p o uches is the b ody cavity or coel om E ven tu ally the a n terior an d p os terior
.

walls of the p ou ches break down so tha t those of ea ch side u ni te to form a tu be .

Thu s the coelom ori ginally segmen ted com es to consis t of a p ai r of c on tin u ou s
, , ,

cavi ties o n e on ea c h S ide of the em bryo

, .

M ean tim e the e ctoderm rises u p o n ei ther side of the m edian dorsal line
,

as a fold or ridg e The two folds m ee t ab ove the m edian dorsal e c toderm w hi ch
.
,

is thu s inclosed an d be co m es the n eu r al tu b e the p rimordi u m of the brai n an d ,

spin al cord .
 GE NE R AL F E ATU RE S OF CHOR DATE D E VE L O P M E NT
37

F ro m the m edian dorsal wall of the en toderm a solid rod of cells is eleva ted
and sep ara ted o ff This is the n otochord or p ri mi tive skele to n
.
.

These p ro cesses are illu s tra ted in F i gu re 6 al so in P a n d H p age 56 ; N , , ,

a
p g e 4 5 ; K a
p g e 1 1,; a n d p ar ti cu larly well in W p la tes be tween pag es 6 2 and ,

63 . S tu dy also the m odels of the develop m en t of Amphiox u s .

2 In ver tebrate s
.
I n ver t ebra t es ( wi th the ex c ep tion of c y clos to mes and

p ossibly u rod e les ) t h e m esoder m does n o t arise as p o u ches fro m the e n toderm .

I n s tead it g rows ou t as a solid sh ee t of c ells from the m edian dorsal regi on an d

FI G . 6 Fo rmation o f the
.
'
n eu ral t u b e, n o tocho r d , m eso derm, an d c oel om in Amphioxus . A D,-

c ro s s sec tio n s ; E, fron tal section
A , d ifferen tia tion
. o f the m ed u ll a ry p
l a te a , t he n o to cho r d al
l a te d , p
the n eu ra l f ol d s b , a n d t he m eso dermal ou ches e p
B , t he n eu ral folds have cl o sed a cro s s ab o ve the
.

p p
m edull a ry l a t e ; the m eso d erm al ou ches are f art her evagin a ted C, the medull ary a n d n o to cho rda l .

p p
l a t es are b eg in n in g to cl ose ; t he mesodermal ou ches f a re com l et el y s e a ra ted f rom the en to derm p p .

p
D, the n eu ral tu b e j an d the n o tocho r d c are com l et ed ; the m esod ermal ou ches a re in creasin g in s iz e p .

E, f r o n t al s ec tio n to S ho w t he m es oderma l p o u ches f o ri gin a tin g fr om the en tod erm s egmen tally . a,

med ull ary pl a te; b, n eu ra l fold ; c , n o to cho r d ; d, n o toc ho r dal pl a te ; e, meso derm ; f, m eso dermal
p ou ches ; g, m ; h, en to d erm o r a rc hen teron ; i , co el om I n all g u res the m eso derm is s ti pl ed
ec to d er . p .

( From P arker

Z
a n d Has well s Textb ook of ool ogy, a ft er Hat schek , cou r tes y o f the M a cmill a n Com an y ) p .

arou nd the blas top ore and g radu ally sp reads la terally an d ven trally be tween
e c toderm an d en toderm I ts origin canno t b e den i tely as c ribed to ei ther e c to
.

derm or en toderm sin c e a t the b las top ore these two g erm layers are c on ti n u ou s
,
.

As the m esoderm al shee ts s p read a c en tral sp li t app ears in them di vi din g them
, ,

in to s oma tic a n d s pl an chn ic walls ; the sp li t i tself is the b ody cavity or coel om The .

c oelo m of ver t ebra tes thu s c o n sis ts fro m the rs t of a si n gle p air of c avi ties
38 L AB ORATOR Y MANUAL T
F OR VE R E B RA E ANA OM T T Y

ex ten ding alo n g ea ch side of the body ax is fro m an terior to po s terior en d and is ,

never c omp os ed of a series of c avi ties as in Amphip xu s The en d resu l t is the .

sam e as in Amphioxu s al thou g h arrived a t in a differen t way

, .

The n eu r al tu be is form ed fro m a p air of fold s the n eu r al fol ds whic h app ear , ,

in the m edian dorsal regio n an d fu se tog e ther to p rodu c e a tu be The n otochord

, .

is sep ara ted off as a sol id c ord of c el ls from the roof of the ar chen t eron (Amphib ia ,

rep tiles ) or else arises from the som ewha t in deni te region where the m esoderm al
shee ts ori gina t e ( birds) .

The p ro c esses will b e g rasp ed m ore cl early by refer en c e to F i gu re 7 also to ,

the gu res in K p a g e 1 1 ; P an d H p a g e 1 1 7 ; W p ag e 6 9 3 Ob tain a m o u n ted

, , , .

A
Dia formation
FI G . 7
.
g r am s t o S ho w t he o f the n eu r al tu b e , n o t ocho r d , m es od erm , a n d co el o m
in v er teb ra tes , Amphibi a
b as ed Cro ss
on s ec tion s A , d i er en tia tio n o f the n o to cho rd f in the roo f
. .

o f the en t od erm ; m eso d erm al p

la t es b s rea din g v en trall y p
B , n eu r al fol d s h risin g a t t he sid es o f the
.

p p
medu ll ary l a t e g; n ot ocho r d f s e arated from the en toderm ; meso d ermal l a tes b ex ten d ed far ther p
v en t r all y a n d p
d ev el o in g a c en t r al c avity , the coel om i C, n eu ra l f old s h n earl y c l os ed to
. fo rm t he
n eu r al tu b e p
j; meso derm al l a tes have r eached the midv en tral lin e ; coel omic split i ha s ex ten d ed

v en t rall y. m ; b , m esod erm ; c, en toderm o r a rchen t er on ; d , s oma tic meso derm ; e, sp l an ch

a , ec tod er

n ic mes o d erm ; f, n o tocho r d ; g, med ulla ry p

l a t e ; h, n eu ral f old ; i , co el om ; j , n eu ral t u b e (A from .

Her twig M ark s Textbook ofthe E mbr yol ogy ofM an an d Mammal s , c o u r tesy o fthe M acmill an Com an y )
-

p .

cross se c tion thro u gh an amphibian em bryo a t the s tag e of the form a tion of
-

the m esoderm an d ex am ine u n der the low p ower

, The se c tion is oval in form ; .

it is in m os t c ases s till s u rro u n ded by the deli c a t e egg membr a n e The o u ter .

layer of the embryo is the e c toderm rela tively thin a n d of the sam e wid th over ,

the whole s u rfa c e I n the m edian dorsal line the e c toderm is p rod u cin g or has
.

already p rodu ced the neu ral tu b e I n the form er case the e c toderm exhi bi ts a
.

p air of n e u r a l f ol ds i n c losi n g a thi ck p la t e of e c t oder m be tween t he m I n the .

la tter case the folds have fu sed a cross in the m edian li n e formin g a tu be the , ,

n eur a l tu b e whi c h is the oval hollow m ass in the m edian dorsal li n e j u s t benea th
, ,

the e c toderm The g rea ter p ar t of the se c tion is o ccu p ied by the archen ter on or
.

p r imi tive i n te s tin e c o m p osed of en,

t oder m The ar c he n t eron has a th
. i n dorsal
wall a thi ck ven tral wall whose c ells c on tain yolk an d i n c loses the rela tively
, , ,

sm all gas trocoe which o ccu p ies its dorsal p ar t I n the m edian dorsal region o f
l
. .
 GE NE RAL FE ATURE S OF CHORDATE DE VE LOP M E NT
39

the ar chen tero n a m ass of c ells will be seen p ro tru din g dorsally ; or in so m e
,

slides this m ass of c ells may have sep ara ted from the ar chen teron and lies be tw een
the la tt er an d the n e u ral tu be Thi s m ass is the n otochord B e tween the e c to
. .

derm an d the ar chen teron on ea ch side is a n arrow sheet of cells ex ten di n g I rom
the sid es of the n eu ral tu be ve n trally I n so m e slides these shee ts will ex tend .

only a shor t dis ta n ce while in o thers they rea ch n early to the median ven tral
,

line Th es e she ets are the m eso derm M ake a diagr am of the se c tion col or in g
. .
,

ectoder m bl u e ,
en toder m yell ow ,
an d mes oder m red .

E . F UR TH E R HI S T OR Y OF TH E LI E S ODE RM

The his tory of the m esoderm is of the u tm os t i mp or tan c e for the u n ders tan d
in g of ver tebra t e s tr u c tu re We have already Iio ted tha t the mesod erm sp li ts
.

i n to two layers an o u ter or som a ti c layer an d an inner or sp la n chni c layer an d

, , ,

tha t the sp a c e be tween the two layers is the body c avi ty or coelom The meso .

derm grows from ea ch side of the embryo nic axis ven trally to the m edian ven tral
li n e or in m eroblas ti c egg s grows ou t over the yolk p u shi n g ou t be tween e c to
, ,

derm an d en toderm .

The mes oder m n ex t b ecomes differen tia ted in to thr ee r egion s : a dorsal region ,

c alled the epimer e wh i ch lies to ea ch side of the neu ral tu be ; a middle regio n
, ,

c alled t he mes omer e or n ephr otome si tu a ted la t eral and v en tral to the ep i m ere ;
,

a n d a larg e v en tral region on ea c h side of the ar c hen t eron c alled the h m

yp o er e ,

or l a ter al pl a te E a ch of thes e region s has of cou rse bo th soma ti c an d sp l an ch

.

ni c walls ( s ee F ig 8 A also K p . The ep imere imm edia tely be com es s eg

, , .

men ted tha t is to say dorsoven tral clef ts app ear i n it a t regu lar i n tervals the
, , ,

p ro c ess be g in ni n g a t th e an t erior en d of the e m bryo and p ro ce edi n g p os teriorly

'

Co n seq u en tly the ep im ere be c om es divided u p ing) a longi tu di n al row of blo cks
, ,

a row o n ea ch side of the n eu ral tu be These blo cks are ep im eres or mesobl as tic .
,

s omites ( ori gi n ally c alled p r o v er teb r ae as it was erro n eo u sly s u pp osed tha t they

were p ri mi tive ver tebrae) At rs t the ep imeres are s till c on tin u o u s ven trall y
.

an d la t erally wi th the m eso m ere b u t even tu ally they are co mp le tely cu t o

,

from the res t of the m esoderm ( see Fig 8B an d the gu re in K p I The .

,
.

meso m ere and the hyp o m ere do n o t be com e segm en ted and rem ain p erm an en tly
in c lose rela ti o n to ea c h o ther Wi thi n the m esom ere li ttl e tu b u les app ear which
.
,

op en in to the cavi ty of the hyp omere ; they are the tu b u les of the kidn ey ( see
Fig SB )
. . The hyp o m er es of ea ch side fold arou nd the ar chen teron their in n er ,

walls co min g in co n ta c t above an d below the ar chen teron to form dou ble walled -

m embran es the dor s al an d ven tr al mesen ter ies ( see Fig 8B )

,
The cavi ties of . .

t he two hyp o mer es b ec o m e the coelo m of the ad u l t ; the c avi ty in ea ch epi mere
disapp ears ; an d tha t of the m esom ere rem ains as the c avi ties of the tu b u les of
the kidney .

In em bryos of the amp hibian typ e the ar chen teron is a c losed tu be and the
two hyp o m er es are c losed c avi ti es whi ch m ee t below the archen tero n In .
 4o L AB ORA OR T Y MANUAL T
F OR VE R E B RA E ANA OM T T Y

embryos resu l tin g from m eroblas ti c eggs however the ar chen teron is op en below , ,

an d sp read o u t on the yo l k an d the hyp o m er es ex tend o u t over the yolk

, The .

d iffer en b es be tween the two t yp es of em bryos are ill u s tra ted in F i gu res 8 A and C .

The em bryo in the case of m eroblas ti c developmen t is la ter cons tri c ted from the
n eural

notochor d neural tub e

meso mere
d o rsal mesen tery

matic
so
meso erm d
in tes tine

n eura l

e imere
p

meso mer e

d
en to er m

d
ec to erm

yol k

FI G . 8 .

Diagrams to Show mesod erm in hol ob l as tic an d m er o b l as tic
the di eren tia tion of the
\

p
ty es o fd ev el o men t p p
A an d B , hol ob l as tic ty e ; C an d D m ero b l as tic typ e A , di er en tia tio n o fthe
.
, .

p p
mes od erm in to e im ere, m esomer e, an d hy o m er e B , s epara tion o f the e im ere from the mesom er e ,
. p
app p
earan c e o f kid n ey t u b u l es in the mesom ere , an d cl osu r e o f hy o m er e a r ou n d t he in tes tin e t o form

d orsal an d v en t ral m esen teries C, simil ar to A b u t in the m er ob l as tic ty e sho win g en tod erm a n d
. p ,

p
mesod erm growin g arou n d the yol k ; n o te how emb ryo is s r ea d o u t o nthe s u rfa ce o f the yolk ; x , in di
cat es lin es wher e emb ryo is cu t o f f from yol k in makin g s ec tion s for mi cro s c0 p ic s tu dy D, simila r t o B .

p
b u t in the merob l as tic typ e ; the en t od erm has c om l et el y su rro un d ed the yol k ; the mes od erm ha s
n ea rl y d on e so ; the yol k s a c is s een t o b e a p ar o f the in tes tin e ; t he emb ryo is
t a r tl y c on s tric ted fr om p
the yol k sac , the co n s t ric tion b ein g the yol k stal k .
(C from W il der s
His tor y of the Human B ody,
c our t esy of Hen ry Hol t an d Com p an y .
)

yolk b y the form a tion of deep grooves on all sides The yolk then han gs from .

the ven tral s u rfa c e of the em bryo in closed in a s a c of blas toderm the yol k s a c .
, ,

whi ch is c onne c ted wi th the embryo by a s talk the yol k s tal k as shown in , ,
 42 L AB ORA OR T Y MANUAL F OR VE R TE B RATE ANATOMY

li n i n g of the i n tes tine an d the ep i thelial li ning and ep i thelial c ells of all of the
o u tgrow ths of the in tes tin e whic h in clu de the gill p o u ches and g ills the larynx
, , ,

win dp ip e and lu ngs the tonsils the thyroid and thymu s g lands the liver the
, , , , ,

g all bladder and bil e d u c t t he p an c reas and the u rinary bladder and adja c en t
, ,

p ar ts of the ur o g eni t al sys t e m The s tu den t sho u ld

. no te t ha t o nl y the e p i thelial
c ells of thes e s tru ctu res arise fro m the en toderm .

H . THE FATE OF TH E ME S ODE RM AND THE F ORMA TI ON OF ME S E NCHYMIE

1 . M e s en chyme .
I n the fu r ther develop m en t of the m esoderm mesen ,

chy me p lays an imp or tan t r 6 1e . M esen chym e is n ot a g erm layer b u t a p ar ti cu lar

kid ney d uct

mesenchyme of scl erot ome

f f
o rming rom ep mere i d
notochor

FI G 9 . .
E nl ar g ed
'

view o f the e pim ere o f a c hic k emb ry o o f t wo d ays

in cu b a t ion t o Sho w t he
t r an sformation of a po rtion o f the ep imere in to m esen chyme . T hese mesen chyme c ell s con s titu te the
scl er o tom e f rom which the ver t eb ral c ol umn ar ises .

typ e of tissu e I t is a p rimi tive kind of connec tive tissu e consis ting of b ran ched
.
,

c ells whose bran c hes are m ore or less uni t ed to fo rm a ne twork ( see F ig
, .

Nearly all of the m esen c hym e com es from m esoderm ; however it m ay aris e ,

from the o ther g erm layers also Hen c e tiss u es an d s tru ctu res whi ch arise.

from m esen chym e may owe their ori gin to m ore than on e g erm layer Co n .

sequ en tly to avoid ina c cu ra c y it is u su ally m erely s ta ted tha t they arise from
,

m esen chym e wi tho u t sp e cifyin g the p ar ti cu lar g erm layer or layers involv ed
,
.

When a g erm layer is abo u t to p rodu c e m esen chym e its c ells be com e loose ,

sep ara ting from their fellows lose their ep i thelial form and taking o n a bran ched
, ,
 G E NE R AL FE A TURE S T
OF CHORDA E DE VE L OP M E N T 43

irregu lar shap e wander away by amoeboid movemen ts to m ore or less deni t e
regio n s where they give rise to c er tain tissu es ( see Fig Those p ar ts of the .

m esoderm whi ch do n o t be c om e m esen chym e b u t r etain their ep i thelial chara c

t er is tic s are c alled mes othel iu m .

2 The fat e of the e pim ere s

.
The m edial wall of ea ch ep im ere t ra n sform s
i n to a m ass of m esen chym e c ells whi ch mi gra t e to a p osi tion aro u nd the n o to
c hord a n d there give rise to the ver tebral c ol um n Thi s m ass of m esen chym e .

is k n own as the s cl er otome ( see F ig The o u ter wall of ea ch epim ere trans
.

form s i n to m esen c hym e c ells whi ch mi gra t e to the u nder side of the e c toderm
'

an d th ere give rise to the inner layer ( derm is ) of the ski n This p ar t of the .

ep im ere is c alled the der ma tome ( Fig The rem aind er of the epi m ere p ersis ts
.

neural tub e

mesen tery

FI G . 1 0 Diagram of cross sec tion s o f ver teb rate emb ryos t o Show the dieren tiation o f the e i
p
-
.

mer e in to d erma tome ( skin p rod u cer ) myo tome ( mu scl e p r o du ce r) a n d s cl erotome ( skel eton p rod u c er )
-
,
-
,
-
.

Derm at om e mes en chyme, myo tome of meso thelium I n B a n d C the d erma

an d s cl ero tom e c o n s is t o f .

t om e is s een p
s r ea din g b en ea th the ec tod erm t o f orm the d ermis o f the s kin ; the myo tomes are g r o win g

ven trall y t o f orm the m u s cl e l ay er o f the b ody wall ; the s cl ero t ome is accumul a tin g ar ou n d t he n o to

p
cho r d a t x t o form the v er t eb r a e ; an d the hy om ere in cl o s es the in tes tin e rod u cin g the d o r sal a n dp
ven tr al m es en teries a n d the mesod erm o f the in tes tin e .

in p la c e as m eso theli u m an d is k n own as a myotome or m u s cl e s egmen t E a ch

'
.

m yo tom e b e c om es sep ara ted fro m the adja c en t o n es b y a c o n n e c tive tiss u e

p ar ti tion the m
, y oco mm a or m y os ep tu m The m yo t o m es g ive rise
. to the vo l u n

t ary m u s c les of the body (wi t h c er tain ex c ep tions ) E a ch g rows fro m its ori ginal
.

dorsal p osi tion ven trally b e tween the e c toderm and the hyp om ere to the median
ven tral li n e where it m ee ts its fellow from the opp osi te side There is th u s .

p rod u c ed a c o m p le t e m u s cu lar c oa t for th e body ( see F ig 1 0 ; also K pp 1 5 .

,
.
,

I 7; W ,p
The fa te of theThe m m e s o m ere
eso m ere ives
rise t o the kid n e ys t he
3 .
g . ,

rep rodu c tive org an s an d their d u c ts ( the termin al p or tio n s of the u ro gen i tal
,

du c ts m ay have e c toderm al or en tod erm al linin gs) .

44 T
L ABORA OR Y MANUAL T T
FO R VE R E B RA E ANA OM T Y

4 The fate of the hyp o m ere

. .
The c avi ty of the hyp om ere is the c oelom
of the adu l t The sp lan chni c walls of the hyp qn er es of the two sides fold
.

arou nd the ar chen t eron an d give rise to mesen chym e from which are pro ,

du ced the smoo th mu s cle and c onne c tive tissu e c oa t s of the dig es tive tra c t
-
,

and also the smoo th mu s cle con n e c tive tissu e c ar tilage etc as the c ase may be
, , , .
, ,

of all the deriva tives of the di ges tive tra c t m en tioned above The hyp omer e .

also gives rise to the linin gs of all the c oelomi c c avi ties the serosa of the vis cera
, ,

and all of the m esen teries The sp lan chn i c m esoderm of the hyp om ere p rodu ces
.

the hear t I n the region of the g ill sli ts the hyp o m ere p rod u c es vol u n tary
.

mu s cles .

5. T he p ro d uc ts of the m e s e n c hym
. e The m ese n chym e giv es rise to all

of the conne c tive tissu es of the body in clu din g car tilage an d bo n e ; to all of the
,

in Vo l un tary or sm oo th m u s c les ; to the blood c ells the blood vessels the lymp h
, ,

vessels an d lymp h glan ds ; an d to the volu n tary mu s cles of the app en dag es
, .

I t has already been s ta ted tha t the vas t m ajori ty of the m esen chym e is of m es o
derm al ori gin b u t a sm all p ar t arises from the o ther g erm layers
, .
V . THE CO M P ARATIVE ANAT O M Y OF THE I NTE G UM E NT AND
TH E E X
O S KE LE T O N

A . G E NE RA L CONS IDE RA TI ONS ON THE S KE LE TON

T he ter m s kel eton in cl u des all of the har d e
por tion s o fthe bod ies o fa imal T he skel
ne d n s . e

ton o f the in ve teb ra tes is common ly extern al fo rmi g a hard co veri g i cl o s i g the bod y
r ,
n n n n ,

whil e that ofthe v er tebr ates is bo th e tern al a d in tern al I n in ver teb ra tes fu ther the skel e
x n .
,
r ,

to is a lifel ess secretion con tain i g n o cel l s whil e the ve tebra te S k el eton is al m s t in va iabl y
n ,
n ,
r o r

c ell u l ar either b ein g compo sed en tir ely o fharden ed cell s o r con s is tin g of c ell s a n d in te c ell l a
, , r u r

produ c ts There are two dis tin ct k in ds of s k el eton in v er teb ates differen t i o rigin a d
. r ,
n n

fu n ction : ( ) T he e tern al s k el eton o r exos kel et n d e ived f rom the S k i an d for mi g a cove
I x , o ,
r n , n r

i g a d p ro tective l ayer on the ou tside of the b o dy

n n ( 2 ) T he in tern al s k el eto o r ndos kel ton
. n, e e ,

d rived chie y f rom the i n er wal l of the ep imere an d co s titu ti g a s u ppo r t an d framewo k
e n ,
n n r

fo r the bo dy an d a p l ace ofa ttachmen t o f the vol un tary mu scl es .

We shall s tu dy the ex oskele ton rs t H owever c er tain p ar ts of the exoskel e .

,

ton have be co m e so closely rela ted to the en doskele ton th a t they will be co n sidered
wi th the la tter .

B . THE S TR U CTURE OF THE S KI N

S in c e the
ex oskele ton is derived fro m the skin a thoro u gh u nders ta n di n g of ,

the s tru c tu re of the ski n is p rerequ isi te to a s tu dy of the ex oskele ton The ski n .

or in tegu men t o ccu rs on ly in ver t ebra tes and m ay be dened as those o u ter layers
of the body wall whi ch are easily sep ara ted from the inner layers S tu dy of the .

micro sc Opic app earan c e an d of the develop m en t of the skin reveals tha t it co n sis ts
of two dis tin c t p ar ts : an o u ter layer the epidermis composed of epithel ial cell s , , ,

an d a n in n er layer the der mis or cor iu m c o mp osed of con n ective tiss u e ( Fig
, , .

The epidermis ori gi n a tes dire c tly from the su rfa ce e c toderm of the embryo whi ch ,

by prolifera tion p rodu ces several s tra ta of c ells to form the ep iderm is The .

der mis is form ed by the mesen chym e of the derma tom e I n the p re cedi n g s ec .

tio n we learn ed tha t the derm a t om e ori gi n a tes fro m the o u ter p ar t of the ep i m ere

( g
F i .
43 )
I . M icr o s c 0 pic
r re of the frog s
s t u c tu u n der m od era te

s kim E x am i n e

p o w ers of th e m i c ros c o p e a c ross se c tion thro u g h the fro g s skin-

a n d id en tify

c arefu lly the followi n g p ar ts :

a ) The epider mis : The o u ter p ar t of the skin c o n sis ts of several s tra ta of

epi thelial c ells This is the epidermis The du term os t of the c ell s tra ta co n
. .

sis t of thin a t c ells whi ch have be c om e conver ted i n to a horn y ma terial This
,
.

co r n i ed p ar t of the ep ider m is is desi g n a t ed the s tra tu m corn eu m B en ea th the .

s tra tu m c orn eu m the c ells gradu ally cha n g e from a a tten ed to a rou n ded and
n ally to a colu mn ar shape Th es e l ay ers of rou nded to col u m n ar c ells con s ti tu te
.

the s tr atu m ger min a tivu m ( also c alled s tr atu m mucos u m an d s tr a tu m Ma l pighii) In .

5
46 L AB ORA OR T Y MANUAL T
F OR VE R E B RA E ANA OM T T Y
the fro g s ski n there is n o sharp dem ar ca tion be tween the s tra tum corneu m and

the s tra tu m g erm i n a tivu m The lowerm os t layer of the s tra tu m g erm in a tivu m
.
,

c o n sis tin g of t all c ol u mn ar c ells is the a c tive ti f th e ider mis and is
p o r o n o e p , con

tin u ou sl y p rolifera ti n g c ells whi ch are p u shed o u tward be c o m e a t and hor n y , ,

a n d nally form p ar t of the s tra tu m c o r n eu m The s tra tum c orneum is shed I

.

a t freq u en t in tervals an d is c on ti n u o u sl y ren ewed fro m below .

b ) The der mis or cor iu m: The dermi s is the in n er p ar t of the skin I t c on .

sis t s of c onne c tive tissu e whi ch in the c ase of the fro g is arran ged p ar tly in the
,

form of a loose ne twork lo ca ted j u s t ben ea th the ep iderm is an d p ar tly in the form ,

MM
U
U
T F
A N
R R
T O
S C

M
U
MV '
I
U T
T A
A N
R I
T
S
M
R
E
G

FI G . I I Dia gr amma tic

. cr o ss - s ec tion thr ou gh t he v er teb ra te s kin , b as ed on m am m a l s .

of layers of den se p arallel wavy bers I n addi tion to the c on n e c tive tiss u e
, , .

bers the dermis c o n tains : pigmen t cel ls dark irregul ar bran chin g c ells forming a , , ,

t hin layer j u s t ben ea th the ep idermis ; the cu tan eou s gl an ds ask shap ed bodies -

m
, ,

p rod u c ed by an in fo l di g of th e s t ra tu m g erm ina tivu and Op eni g b y a neck

p
n ,

to the s u rfa c e ; and a t in tervals c ol u mn s c on tainin g sm oo th mu s cle c e lls blood ves ,

sels and nerves these c olu mns c rossing the derm is a t ri gh t an gles to the s u rfa c e
, ,
.

Of these s tru c tu res the cu tan eou s g lands are the m os t c o n sp i cu o u s They are .

really p ar ts of the s tra tu m germiirativum w hi ch have been evagi n a ted in to the ,

loose p or tion of the dermis .

Draw a s mall p or tion of the skin to show the p ar ts n am ed above .

In m t t o l y t h p lif
so e ex s ti g l y
n ll thi k i
e ll d th t t m g min tivu m
ro er a n a er , on e c e c ,
s ca e e s ra u er a ,

th t p t i
a f th pid mi b t w
or on o e ethi d th t t m
er s m b i g th d ig t d th t t m
e een s an e s ra u c o rn eu e n en es na e e s ra u

Mal pighii . We shal l here r egar d the t erms s tr atum g ermin a tivum an d s tra tu m M alp ighii as Syn on y
mou s .
 T HE IN E T GUME NT AND T HE E XO S KE L E TON 47

C . THE E X OS KE LE TON IN G E NE RA L

The ex os f rom the s kin I t is p rodu ced b y harden in g p roces ses in the
k el eto n is derived .

ep id ermis o r the d ermis o r in bo th E xosk el eton d erived fr m the ep idermis is s po k en of as

q
.

e ider mal it is rodu ced b the activit o t

p p ; y y f he s tr atu m ger min ati vu m an d con sis ts ofma n yat hor n y
cel l s pr ess ed r ml y together to ma ke a har d s tr u ctu r e I t will b e seen that ep idermal exosk el e to n .

s tru c tu r es ar e on l y s p ec ial po rtion s of the s tratum co rn eu m E xos k el eton d erived from the .

dermis is sp ok en o f a s der mal ; it n ear ly al ways cons is ts ofb on e pr odu ced by the mes en chyme cel l s ,

or igin atin g fr om the der ma tome F rom the emb ryol og ical po in t of view epider mal exos kel eton
.

is ofectoder mal or igin whil e der mal exos kel eton is mes oder mal
, E p idermal an d dermal exo skel e .

ton a r e d iffer en t b o th mo rp hol og icall y an d embryol og icall y Fr om ou r den ition o fho mol ogy .

it foll o ws tha t exo s k el etal s tru c tu r es o f ep idermal origin are homol og ou s in differen t an imal s ,

an d der m al s t ru c tu res ar e lik ewise homol og ou s I t will b e ou r pu rp os e to t race the homol o gy .

of the ex os k el eton in the differ en t b

v er t e ra t e cl asses .

The houl d al so read the chap ters

s tu den t s on the exos k eleton in the s tan dar d texts of

comp arative an a to my as K W an d W d , , .

D . E X O S KE LE TON OF FI S HE S

M os t shes are cov ered wi th s cales w hi ch are of fo u r kinds , .

The pla c oi d s c al e
1. This typ e of s c ale o ccu rs .
elas mobran ch sh es in the .

I t c onsis ts of a b as al pl a te c arryi n g a p roje c ti n g s pin e G ood examp les are ob tain .

able from the ska te Cu t o u t fro m the ska te a sm all p ie ce of skin con taining one
.

sp ine Clean away the skin so as to exp ose the comp le te s c ale I t consis ts of
. .

a too thed basal p la te from whi ch arises a shiny cu rved sp ine The base of the
, .

spine is hollow the cavi ty bein g known as the pu lp cavity This cavi ty can be lo

.
,

ca t ed by p robi n g wi th a needle p oin t in the c en ter of the u nder side of the basal

p la t e The. S hi ny c oa t in g of t h e s p ine is c o m p osed of en a m el ; th e basal p la t e an d

in terior of the sp ine consis t of den tin e a su bs tan ce similar to bone Draw the ,
.

s cale .

T o u n der s tan d the s tru c tu r e of the pl acoid s cal e it is ecess ary to con s ide its mode o f n r

devel o p men t E a ch s cal e o rig i at es f om a reg ion of c ell mu l tipl ica tio n in the d ermis
. n rThe .

c ell s t h s fo m ed heap u p i to a co e the der ma l papill a which p u shes u p ag ain s t the un der S id e
u r n n , ,

o fthe s t ra tu m g ermin at ivum T he ou ter cell s o f the d erm al p ap ill a beg in to s ec ete d en tin e
. r ,

an d p o d c e the b asal p l a te an d the i s ide o f the s p in e

r u T he s tra tu m g e mi a tivu m in con tac t
n . r n

with the d e mal p ap il l a s ec etes o its u der su rfa ce the su b s tan ce e amel which co sequ e tly
r r n n n ,
I
n n

form s a coat o v er the d en tin e o f the s pi e T he spin e whe fo rmed b eak th ou gh the su r n . n r s r

face The i te io c ell s of the d ermal p ap ill a do n o t un d e go a y cha g e b t emai as a so ft

. n r r r n n u r n

pul p o ccu pyi g th pul p cavity i the i te io of the spin e Fro m this accou t it follows that
n e n n r r . n

the p l a co id sca l e is c omp os ed o f b o th ep id erm al a d de m al co n s titu en ts T he d evel op men t n r .

o f the p l ac o id s ca l e is ill u s tra ted in F ig u r e 1 2 ; al so in W p ag e 8 2 ; CNH Vo l VII pa g e , ,

.
,

1 8 5; W d p ag e 40 ; R p ag e 4
, , .

Draw an im aginary longi tu di n al se c tion throu gh a pla coid s cale colori n g e c to ,

d erm al p ar ts bl u e an d m esod erm al p ar ts red .

T ha t t his s ub s ta n ce is rea ll y en a m el . s u ch as o cc u rs o n t eeth, ha s b een q u es t io n ed , so me main

t a in in g t ha t it is a kin d o f d en t in e ( s ee L, p . We s hal l ho wever her e
, ,
ta ke the u su al View o f its

o rig in .
48 LABORATOR Y MANUAL F OR VE R E B RA E ANA OM T T T Y

The d ogs h is c overed wi th p la coid s cales whi ch are mu ch sm aller and m ore ,

c losely set t o ge ther t han in t he c ase of the ska t e The sp in es of the s cales can .

be fel t b y p assin g the han d over the d og sh skin Hy boilin g a p i ec e of the skin .

wi th alkali the s c ales m ay be sep ara ted ou t E x am in e s u c h isola ted s cales

, .

u n der the m i c ros c op e They are similar to those of the ska te wi th rhom boid
.
,

basal p la tes an d a p roje c tin g sp i n e .

2 .

The ho m ology of tee th an d pla c oi d s cale s The m os t i n teres ti n g p oi n t
abo u t the p la c oid s c ale is tha t its s tru c tu re an d m ode of ori gin are ex a c tly the
S tra tu m str atum

E
FI G . 1 2 .
Fiv e su ccess iv e s ta g es p
in the d ev el o m en t o f the pl acoid s c al e o f the dogs h . A, g ather
in g of the c ell s o f fo rm the d erma a a B evagin a tion o f the d ermal p apill a a d
the d ermis t o l p pil l .
,
n

s ec r e tio n o fthe d e tin e ( c ol o r ed b l a ck ) b y the ou t er c ell s o fthe p apill a

n C c on tin u ed s ecret io n o fd e ti e .
,
n n ,

t hinnin g o f the i t erior o f the p pill a t o fo rm the p ul p

n a D b egin n in g s ecr eti n o f the b a sa l pl a t e a n d
.
,
o

fo rmatio n of the en amel (l eft white) b y the u n d er su r face o f the s tra tum g ermin a tivu m E erup tio n
G X
.
,

o f the s pi ne t hrou gh t he epid mis ,

er c om pl et io n o f the b asal pl a te an d p lp ( F
u . ro m oo d rich in P art I
of L an kes ter

s Tr ea tise on Z ool ogy, c ou r t esy o f the Macmill an Co mp y ) an .

sam e as those of the t ee th of all V er tebra t es F ro m thi s it follows tha t t ee t h .

an d p la c oid s c ales are ho m olo g o u s s t ru c tu res and t e eth are m erely sli g h tly m odi ,

ed p la coid s c ales The hom ology is p robably du e to the fa c t tha t the lin in g of
.

the m o u th c avi t y is really skin tu rn ed in an d he n c e m ay be exp e c t ed to give ,

rise to s tru c tu res sim ilar to those fou n d in the ski n .

Ob tain a lo n gi tu di n al half of an ordin ary ver tebra t e t oo th I den tify the .

followi n g p ar ts : the crown or S hi n y u pp er p ar t c orresp on di n g to the sp i n e of the ,

 50 L AB ORA OR T Y MANUAL T
F OR VE R E B RA E ANA OM T T Y
m ay g enerally be no ti c ed tha t the ep idermis form s a thi n coverin g over the free
t oo thed end . Draw .

Q
h cycl oid an d c te oid s cal es are composed f bon e an d are homol ogou s with g a o id
Bot n o n
s cal es Al l th e typ es o fs cal es a r e p o du c ed by hea p s o fmes en chyme c el l s o f the d ermis
. re r
T he .

po te io edge of each s cal e g ows ou tward ca rryin g the ep idermis with it ; the l atter may r e
s r r r ,

main a s a thin co v e i g o v er the p ro j ecti g p a r t o f the s cal e or m a b e ru bb ed o ff T he

r n n
y .

s ca l e o f sh es a e thu s o f d erm al o i i
s r with the e cep tion o f the pl a co id s cal e which c on
r n,
g x
,

t ain s in a ddition to its d erma l p art a smal l ep id e mal c o n trib tio n , r u .

E . E X OS KE LE T ON OF A MP H I B I A

The v ast maj o rity of p r esen t

day Amphib ia have na k ed S kin s that is an ex os k el eton is
, ,

l a c k in g . p rovided with mi u te con ceal ed d e mal scal es The e ti c t Amphibi

A few ar e n , r . x n a

(S t g ph l ) w e e co mm o ly cl o thed in a heavy a mor o f d e mal pl ates ho mol ogou s w ith

e oce a a r n r r

g a o id s cal es an d s imil ar to them in app ea an c e

n r .

st ra tu m m
s tratu stratum
5at u m
c orn eum
germin at vumi

B
Dia kin
FI G . I 3 .
gr am ma tic l o n gitu din al s ec tio n s t hro u gh the s of A, a t el eo s tom e s h a n d B , a

re p til e, to S ho w t he l o ca tio n s o f t he s cal es . I n A, the s ca l es ar e in the d ermis , whil e in B they r ep r e

s en t thic ken ed p o rtio n s o f the s t r atu m c o rn eu m .

( From W ied ers heim

s Compar ative An atomy o f Ver
teb rates , co u r tesy o f t he M a cmil l a n Co m p an y . )

F . E X O S KE LE TON OF RE P TI L E S

The bodi es of rep tiles are c hara c teris ti cally c lo th ed wi th an ex oskele ton com
p osed of hor n y s c ales These s c ales are of ep iderm al ori gi n rep rese n ti n
.
g p ar ticu ,

l arl y den se co rni ed areas of the s tra tu m c orn eu m

,
I n the form a tion of s u ch a .

horn y s c ale a derm al p ap illa rs t app ears whi ch fu rn ishes n u tri m en t for the c ells
e n gaged in p rodu ci n g the s cale The s tra tu m germina tivu m over the derm al p apilla.

begins to p rolifera te rap idly p rod u cing c ells whi ch be com e a t an d horn y and,

c o m p ressed i n t o a s c ale ( see F ig I n addi tion to the horn y s c ales m a n y .

rep tiles p ossess bon y p la tes of derm al origin si tu a ted ben ea th the ep iderm al
s cales These bo n y p la tes ori gina t e in the sam e m an n er as the s c ales of shes
. .

I n order to avoid c o n fu sion the ep iderm al s c ales will be designa t ed as s cu tes and
,

the derm al s t ru c tu res as pl a tes .

1 E x o s kele to n of the 1izar d

.
R e c all the c o n di tion of the liz ard or r e .
,

ex am ine the sp e ci m en The body is c lo thed wi th horn y s cales o f ep iderm al

.

origin overlapp in g like shingles No te arran gem en t an d si z e of the s c ales on

,
.
 T HE IN E T GUME NT AND THE E XO S KE L E TON 51

vario u s p ar ts of the body The head may bear enlarged s cales or head s hields
.
.

There is n o derm al ex oskele ton in liz ards .

2 . E x o s kele ton of the tu rtl e The ex oskele ton of the tu r tle is somewha t
.
-

c o m p li c a ted F o r the p u rp oses of s tu dy c arap a c es and p las trons which have

.
,

been sep ara ted by sawin g thro u gh the brid ges will be p rovided I t is also de .

sirable tha t these p ar ts shall have been cooked to render the su tu res be tween the
p la t es m ore dis tin c t The followin g
. des c ri p tion is based on the e x oskele ton
of ou r comm on po nd tu r tles ; in o ther families of tu r tles the arran gem en t of the
s cu tes and p la tes may be sli gh tly di er en t .

a ) The car apa ce : The dorsal s u rfa c e of the c arap a c e consis ts of lar e thi n
g
horn y s cales or s cu tes whose bou ndaries are m arked by grooves These s cu tes
, .

are of ep iderm al ori gin form ed from the s tra tu m corneu m The s cu tes are ar
, .

ran ged in ve longi tu dinal rows one m edian and two p airs of la teral rows The, , .

m edian row c o n sis ts of ve n eu ral s cu tes ; on ea ch side of this are fou r cos tal s cu tes
and the m argin s of the carap a c e are covered by a nu mber of small margin al s cu tes .

The u n p aired narrow margin al s cu te at them iddl e of the an terior end of the cara
p a c e is t er m ed t h e n u chal s cu t e ; besides t his there are twelve
p airs of m ar ginal
s cu tes of whi ch the median p os terior two behin d the f th neu ral s cu te are of ten , ,

c a l le d pygal s cu tes Observe tha t the margi n al s cu tes are c on tin u ed over the
.

edge of the c arap a ce to cover the m argins of the u nder side Draw the dorsal .

su rfa ce of the carap a c e showing a c cu ra tely the o u tli n es of the s cu tes (only half
,

n eed be lled in ) .

Tu rn the carap a ce over and s tu dy its ven tral su rfa ce I t is co mp osed of .

heavy bony p la tes of derm al origin The ver tebral col umn wi th its ribs o ccu .

p ies th e m edian li n e and bo th are rm ly fu sed to the c arap a c e

,
The bo u ndaries of .

ea ch p la te are m arked by jagged s u tu res whi ch sho u ld be lo ca ted in iden tifying

the p la tes The p la tes like the s cu tes are arranged in ve lo n gi tu di n al rows
. .

The m edian row of p la tes fu sed to the dorsal sides of the ver tebrae c onsis ts
, ,

of a single large an terior n uchal p la te followed by eigh t sm aller ver teb ral or n eu r al
p la t es ,
ea c h a t t a c hed to a ver t ebra followed by t wo pos tn eu ral or preca udal
,

p la t es n
,
o t a tt a c hed to ver t ebrae On ea ch side of the m edian row is a row of
.

cos ta l p la tes ei gh t p airs of elon ga ted p la tes ea ch a tta c hed to a rib

,
The margins ,
.

are form ed of eleven p airs of margin al p la tes and one single u n p aired pygal pla te
in the median p os terior p osi tion Make an a ccu ra te drawing of the u n der S ide .

of the carap a ce showin g bony p la tes and their rela tion to the ver tebrae and ribs
,
.

Do the s cu tes an d p la tes of the carap a c e corresp ond ?

A d ier en ce o fo p in ion regardin g the bo y pl a tes ofthe tu rtl e s carapace Accordin g
exis ts n .

to o e view al l o f the pl tes a e ofd ermal origin an d the e pan d ed ver teb ra e an d rib s are fu sed
n ,
a r x

to the u n der s f a c e of these d e mal pl ates

ur Acco rdin g to the o ther View which is p robably the
r .
,

co r ec t on e the ver teb ral an d c o s ta l p l ates a e fo rmed en tirel y by the e p an s ion of the ver teb ae
r ,
r x r

a d ribs an d a e the efo e n o t d er mal e o s k el eton a t all b t p a r ts of the en dos k el eton

n ,
r r r x This u .

View a l o i cl u d es the c o cep tion tha t tu rtl es o rig in all y po ss es ed a s et o f rea l d erma l p la tes
s n n s
52 LAB ORA OR T Y MANUAL F OR VE R TE B RATE ANATOMY

ex tern al p es e t vert ebral an d costal pl ates an d that thes e pla tes s bs equ en tly dissa p
to the r n ,
u

p ea red . On bo th po in ts ofview the n u chal margin al po s tn e ral an d pyg al pl ates a r e d ermal

, ,
u ,

an d part o f the e xo s k el eton .

b ) The pl a s tr on :The p las tron like the c arap a c e consis ts of a set of horny , ,

ep iderm al s cu tes coveri n g bo n y derm al p la tes S tu dy the ex t ern al (ven tral) .

su rfa ce of the p las tron I t is covered by six p airs of s cu tes n amed from in
.

fron t ba ckward : gu l ar hu mer al pectoral a bdomin al femor al an d an al I rregu lar

, , , , , .

infr amargin al s cu tes cover the brid ges S tu dy the in tern al ( dorsal ) s u rfa ce .
,

n o tin g the larg e bo n y p la t es u ni ted by jagged s u tu res whi ch c o mp os e it The , .

small an terior p air of p la tes are n am ed epipl as tr a B e tween them is a S in gle .

median p la t e wi th a p os teriorly p roje c ti n g p oin t nam ed the en topl as tr on or in ter ,

cl avicl e B ehi n d these are three p airs of large sq u arish p la tes n am ed the hyo
.

p l a itra hyp p
,
o l a s tra and x iph ipl a s,
tra the r s t n a m ed bei n g m os t an t erior D raw
, .

the p las tron showin g o u tli n es of s cu tes and p la tes

,
.

c) Other exos kel eta l s tr u ctu res : The ex oskele ton also i n clu des the claws
a n d the hor n y beaks whi ch in c ase the jaws E x am i n e these beaks in a dem on .

s tra tion sp e cimen in w hi ch they have bee n loose n ed fro m the u n derlyi n g bones .

Tu r tles also p ossess s cales or thi cken ed s calelike areas o n the legs an d ta il an d ,

in som e cases en larged s cu tes over the head All of these s tru c tu res are of epi .

dermal ori gin consis ting o f sp e cial p or tio n s of the s tra tum corn eu m
,
.

G . E X O S KE LE TON OF B I R DS

B irds
are c lo thed in an ex oskele ton co n sis tin g of fea thers on the g rea ter p ar t
of the body s cales an d c laws o n the fee t and horny beaks Al l of th ese s tru c
, ,
.

tu res are of ep iderm al ori gin form ed fro m the s tra tum c or n eu m ,
There are no .

derm al elem en ts of the ex oskele ton in birds The fea thers of birds are their .

chara c teris ti c fea tu res and will be s tu died in de tail There are three kinds of .

fea thers down fea thers con tou r fea thers and ha ir fea th ers or l opl u mes
, , .

1 S tru ctur e of the d own feather

.
D own fea thers or pl u mu l ae c on s ti tu te .

the u ffy coveri n g of yo u n g birds an d also o ccu r in adu l t birds be tween the bases
of the con to u r fea thers Ob tai n a down fea ther or a p rep ared slide of on e
. .

I den tify the shor t s tem or qu ill the sof t rays or b ar b s whi ch sp ring in a cir cle ,

from the top of the q uill an d the mi n u te side rays or b ar bu les on the barbs Draw
,
. .

2. Devel opm en t o f the d own f eather .

A down fea ther
p apill a of the s kin arises f rom a ,

thefeather papil l a con s is ti g o f a d ermal co e the pu l p covered by the ep id ermis

,
n L a ter this
r , , .

papilla sin ks in to a p it in th Skin called the fe ther foll icl e T he s t a tu m ge min ativum of
e , a . r r

the p ap ill a b eg in s to p olif e t e f o min g a n u mb e of l o gitu din al co l u mn s which p o j ect in to

r ra , r r n r

the pu l p ( see F ig Thes e column s even tu all y sep arate each bein g a barb comp osed o f
.
, ,

co rn i ed cell s . T he o rig in al s t r tum co rn eu m cov e s the ba b s l ik e a shea th which is call ed

a r r ,

the per ider m; it s pl its o p en an d is shed r el ea s i g the b arb s Du rin g this p o ces s the who l e
,
n . r

papill a has been el on ga tin g an d comes to p roj ec t above the su rface T he l ower pa t of the . r

papill a does n ot s plit in to ba bs b u t emai s u divided as the qu il l within which the p l p d ries
r r n n ,
u
 THE I NTE GUME NT AND THE E XOS KE L E TON
53
up . ( See fu r ther K , , p 6
3 ;
. P a n d H , 6
3 9 73 ;
pp
W d.
, pp . 25 It shoul d b e n o ted tha t
p
the dev el o m en t o fa f ea ther is simil ar to tha t o fa r e tilian p s cal e , in vol vin
g a d erm al p p
a ill a for
n u tritive p u rpo ses an d an e c to d ermal thic k en in
g .

3 . S tru ctu re of a is th
c ontou r feather
e c o mm on t y e of fea ther
p .

This
whic h c overs the bodies of birds Ob tain on e and iden tify the followi n g p ar ts
.
.

The S hor t bare hollow p or tion the lower end of whi ch is inser ted in to the fea ther
, ,

folli cle of the skin is the cal amu s or qu ill ; the rem ai ning exp anded p or tion con ,

s titu tin g m os t of the fea ther is the van e The qu ill bears two op enin gs the
, .

infer ior u mb il icu s a t its p ro xim al en d whic h was i n ser ted in the skin and thro u h
,

,
g ,

whic h in the early s ta ges ofthe fea ther the derm al p ap illa p asses ; and the s uperior
,

u mb ilicu s a t the j u n c t ion of vane and q u ill on the ven tral s u rfa e of
c the fea ther .

F ro m the s up erior u mbilicu s p ro tru des a m ore or less well develo ed a ccessory -
p

C 0
FI G . I 4
.
Dia gram s o f the d evel opmen t o f the down A, dermal p apill a
an d c o n to u r fea ther s . .

B , c ross sec tio n o f a l a ter s ta g e o f the d ermal p apill a showin g the thi c k eni n g s o f the s tra tu m g erm in a
-

tivu m . C, c ro ss s ec tion o f a l a ter s ta ge ; ea ch thickenin g has s epa rated to form a b a rb ; the d ermis in
-

the cen ter o f the p ap ill a ha s d eg en era ted in to the pith; the s tra tu m c o rn eu m forms t he p erid erm or
s hea th o f the d o wn f ea ther D, s ec tion across a d evel opin g c o n tou r fea ther , sho win g the t wo en l a r g ed
.

k enin gs o r
t hic b a rb s whi ch b ecome the r a chis an d the o b liq ue cou r s e o f the o ther b arb s .
(A C-
f r om
W ied ersheim s Compar ative An atomy of Ver teb r ates cou r tes y o f the M acmill a n Compan y ; D

, a f ter

Kin gsl ey s Compar ative An atomy of Ver tebr ates copyright b y P B lakis ton s S on a n d Compan y )

, .

.

fea ther c alled the afters haft co n sis ti n g of onl y a few tu f ts in some birds b u t in
,

o thers of a c omp le te fea ther nearly or qu i te as larg e as the p rimary fea ther
, .

The van e c on sis ts of a c en tral axis known as the s haft or rachis whi ch is con tin u
o u s wi th the qu ill and of a sor t of web or m em brane sp rin gi n g fro m ea ch side of the

ra chis E x tend ing alon g the ven tral s u rfa c e of the ra chis is a g roove the
.
,

u mb ilica l gr oove The web or m embrane of the fea thers is obviou sly comp osed
.

of a large nu m ber of p arallel obliqu ely p lac ed rays adheri n g to ea c h o ther

, , .

These rays are the b ar bs and ea c h barb b ears side rays called b ar b u l es exa c tly
, ,

as in the down fea ther The barb u les in terlo ck c au sin g the barbs to adhere to
.
,

p rod u c e an u nb roken s u rfa c e To see the m e thod of in terlo cki n g of the barb u les
.

it is ne c essary to ex am i n e a s m all p ie c e of the fea ther u n der the m i cros cop e It .

will then be no ted tha t the barb u les of o n e side of ea ch barb ( side toward the qu ill)
54 LAB ORA OR T Y MANUAL T
FOR VE R E B RATE ANA OM T Y

are p la c ed diagon ally a c ross the b arb u les of the o ther side ( side toward the tip of
the fea ther ) of the adjoin in g barb and tha t the form er are p rovided wi th hooklike,

p roje c tio n s wh i ch t o ver a n d c a t ch a n g elike ex th n s io n s of th e la t t er The .

diagon al arran gem en t p ermits on e hooked barb u le to ca tch a number of an ged

barb u les B y pu llin g on the barbs on e can u nhook the barb u les an d by s troking
.
,

the barbs o n e can hook them u p a gain The la tter p ro cess is the c hief object of
.

the freq u en t p re en i n g of the fea thers habi tu al wi th birds In the rec t ri c es an d .

remiges the hookl e ts are well develop ed throu ghou t so tha t the barbs are in ter
lo cked over the en tire fea ther ; b u t in the cover ts the l ower barb u les la ck the
.

b ookl ets and the lower barbs are loose a n d u ffy

,
In so m e birds as the os tri ch .
, ,

all of the barb u les la ck hooks and the whole fea ther is uffy , .

Draw a co n to u r fea ther showing its p ar ts , .

Dev l pm t of the con tour f e ther Con tou r feathers ar e s imil ar in s t ru c tu e a n d

e o en a . r

d ev el o pmen t to d o wn feathers excep t that in the l atter the barbs s prin g direc tly fro m the q il l
, u

in a cir cl e whil in the fo rmer they s prin g in a row f ro m either s id e o f a c en t al axis

,
e This is r .

du e to the fac t th t in the dev el o pm e t o fthe con tou r fea ther two of the l on g itu din al col u m s
a n n

formed by the s t t m g ermin ativum become mu ch enl a ged an d fu se to p od u ce the rach is

ra u r r ,

the gr oove b et w ee t hem rem a in i g as the u mb il ica l g oo ve

n n This e l argmen t of these two r . n

c ol u mn s imp ar t s an obl iq u e po s ition to the o ther col umn s which become the b a b s s in ce the ,
r ,

r a ch i s g ows fas ter than the barb s

r T he con tou r fea ther is r oll ed u p in s id e the p e id erm the
. r ,

fu tu re dors al or ou t er su rf ace b ein g ou tside n ext to the p eriderm whil e the fu tu e ven tral o r ,
r

in er sur fac e is in s ide n e t to the d ermal p ap ill a

n x When the p eriderm s pl its the f ea ther at te
.
,
ns

ou t . T he quil l is the bas e o f the f ea ther which fail ed to s plit in to barb s ; the white a k y
ma terial n o ticeabl e in side the qu ill is the d ried r emai s of the d ermal p ap il l a ( see Fig 1 4 ; n .

al s o K p 3 7 Fig
, .
, .

5 F eather tr acts B irds a ear

be c omp le tely c overed by c on tou r to
. .
pp
fea thers b u t a ctu ally the fea thers are born e onl y by c er tain areas of the skin
,

c alled fea ther trac ts or pteryl ae with fea therless areas apteryl ae be tween them
, , , .

Observe the fea ther tr ac ts on a dem o n s tra tion sp e cim en of a yo u n g bird an d con
s u l t also K Fi gu re 2 6 p age 3 5 or P and H F igu re 1 0 3 7 p age 3 72
, , , , , ,

6 S tru ctur e of the IOpl ume

.

The l opl u m es are the hairs visible on a .

p l u c ked bird E x a m in e a re
p p
. ared slid e or re m ove a l o p l u m e fro m a p l u cked
bird m ou n t on a slide in a drop of wa ter an d ex amine u n der the low p ower I t
, ,
.

c o n sis ts of a m ai n ax is beari n g a few t er minal barbs I t is a m i n ia tu re d egener .

a ted co n tou r fea th er D raw . .

H . E X OS KE LE T ON OF MAMMA L S

The ex oskele ton of m amm als c on sis ts p ri maril y of hair fou nd in no o ther ,

ver tebra tes S om e m amm als are p rovided wi th s cales in addi tion
. .

1 S tr u ctu re of hair
.
B y ex am inin g yo u r own skin de term in e tha t ea ch
.

hair sp rin gs from a pit in the skin kn own as a ha ir fol l icl e R emove a ne hair ,
.

from the u n der side of the forearm and ex amin e wi th hi gh p owers of the mi cro
s cop e Observe b y fo cu sin g on the s u rfa c e of the hair the irregu lar wavy
.
 THE IN E T GUME NT AND THE E XOS KE L E TON 55

ou tlin es of the c ells of whi ch it is comp osed E x ami n e wi th the mi cros c op e a .

c ross se c tion thro u gh the skin c on tai nin g g rowi n g hairs a n d s tu dy a hair folli cle
-
, .

E a ch is a deep p it in the skin and is li n ed by ep iderm is At the bo tto m of the .

folli cle the dermis form s a sm all b u lb shap ed enlargemen t the hair papill a The -
, .

s tra tu m germina tivu m over this p apilla is seen p rolifera tin g a conical heap of cells
from whi ch the hair arises Thi s c one ex tends u p the folli cle and af ter a shor t
.

dis tan ce a sp li t is seen sep ara tin g a cen tral shaf t the r oot of the hair from the , ,

walls of the folli cle The tra n si tion from the ordin ary ep i thelial cells of the p r o
.

lifera tin g area to the horn y cells of the roo t of the hair is readily observable The .

li n i n g of the folli cle is also somewha t cornied and is k n own as the ou ter root ,

s hea th .I t is the whi te c oa t whi ch clin gs to the roo ts of hairs when they are
p u lled fro m t heir fol l i c les F ro m t his a cc o.u n t it follows t ha t hairs are e p ider m al
s tru c tu res p rodu ced by the a c tivi ty of the s tra tu m germi n a ti vum Draw a hair .

follicle S howin g its s tru c tu re Fu r ther de tails wil l be fou n d in tex tbooks of
,
.

his tology .

2 . S cal e s of m amm al s A nu mbe r of m amm als p ossess hor n y s c ales like

.

those of rep tiles These m ay cover the body as in the s caly an tea ter (Man is) g
.
, ,

in P a n d H p a ge 48 5 and N p a ge 3 75 b u t comm o n ly o ccu r on the tails o n ly

u re
d , , , , ,

as in ra ts beavers and a n u mber of o ther m am mals Su ch s caly p ar ts are also

, ,
.

p rovided wi t h s c an t y hairs E x a mine a ra t s tail a n d observe the s cales an d

.

hairs up on it .

The arm adillos are the only livi n g m ammals whi ch p ossess like the tu r tle an , ,

arm or comp osed of bo th epiderm al s cu tes an d dermal p la tes Ob tain a dried .

arm adillo arm or and ex amine the ex tern al su rfa ce I t is m ade u p of an an terior .

s hiel d of s m all p oly gonal s c ales a middle region co mp osed of n i n e m ovable b an ds

,

wi th bare areas of skin be tween them and a pos terior s hield similar to the an terior ,

shield The ou ter su rfa ce of the arm or con sis ts of thin horny epiderm al s cales or
.

s cu tes p o l ygo n al on the s hields t rian gu lar on the bands

, ,
The trian gu lar s cu tes ar e .

al tern a tely reversed in p osi tion so tha t in half of them the ap ex p oin ts an teriorly
, ,

and in half p os teriorly The form er bear hairs at their p os terior m argins How
.
.

many hairs to ea c h s cu te P Draw som e s cu t es and hairs to S how their rela tion .

Tu rn the arm or over an d s tu dy the in ternal su rfa c e I t is c omp osed of bon y .

p la tes of d er m al ori g in p oly g o n al on ,

t he S hields re c ta n gu lar on the ba n ds ,
.

Wi th a knife p oin t s c rap e offsome of the epiderm al s cu tes ; n o te rela tive thi ck n ess
of s cu tes and p la tes d ifferen ces in the m a terials of whi ch they are com p osed o n e
,

of horn the o ther of bo n e and the imp ressio n s lef t o n the p la tes by the s cu tes .
,

R ead W p g a e s 87 97
,
o n t he rela tion of,
s c ales to hair a n d t o the fri c tio n ridges .

I . S UMMAR Y
1 . T he ver teb r ate k in co s n s is t s of an ou t er e pidermis , ec t o d er m a l in o rig in a n d c o m o s ep
d o f

ep ithelial c el l s an d a n in n er d ermis , meso dermal in o r ig in an d c o mpo s ed o f con n ec tive tis su e .

2 . E x osk el etal s tru c tu res are p rod u ce d by the s kin , either b y t he ep idermis o r the dermi s
or bo th .
56 LABORATOR Y MANUAL T
FOR VE R E B RA E ANAT OM T Y

3 . p
E id ermal ex o s k el eta l s tru c tu r es con s is t o f ho rn p
an d are co m o s ed of n u mer ou s a t
t en ed d ea d p res sed together Thes e cell s aris e by p olife atio ofthe l ower laye s
corn i ed cell s . r r n r

o fthe ep id ermis ( s t atu m g ermin at ivu m) Of su ch n atu re ar the su pe r cial s cal es o f ep til e
r . r s.

bi ds a d mammal s fea thers hair cl aws a il s beaks hoo fs horn s su ch as tho se ofca ttl e
r ,
n , , , ,
n , , , .

4 De mal exosk el etal s truc tu r es con sis t of bo n e secr eted by the mes en chyme c ell s o f the
. r

dermis .

Of su ch n atu re are the s cal es of shes in whol e o r l a ge p a t the bon y pl at es o f r r ,

r ep til es an d m amm al s the horn s ofd eer an d an tel op e the n rays of shes the p l a tes o f ec
, t , , r en

an d ex tin c t Amphibia .

5 Cer tain ex os k el etal s tru ctu r es con tain ot d ermal an d

. b h
pidermal con s titu ten ts in extri e

ca b y l fus ed h p
Su c ar e the l acoid s cal es o f the el as mo ran c
. b h shes an d the teeth ofall ver te
b a tes
r B y the id en tity o f their s tru c tu r e an d mod e ofd evel o p me t thes e two s tru ctu res ar
. n e

s hown to b e ho mo l og ou s .

6 I n the fo rm a tion of e osk el etal p arts a d ermal p ap ill a is in volved which fu rn ishes
. x ,

n u tritio n for the p rolifera tin g o r s ec r etin g cel l s I n the cas e o fdermal s tru ctu r es the cell s o fthe
.

pap ill a give ris e to the s tru ctu re I n the cas e o f epidermal exosk el eton the p ap il l a tak es n o
.

par t in the formation of the stru ctu re b u t S imply bri gs a blood su pply to the d evel opi g p ar t n n

which ar ises sol ely by p rolifera tion an d su b s equ en t co rn i ca tion o fep idermal cell s .

7 From the for egoin g accoun t an d f om the d e n ition o f ho mol o gy it foll ows tha t all exo
. r

s k el t l s tru c tu res o f ep iderm al o rigin ar e homo l og ou s

e a
an d tha t s imil ar l y a ll exosk el etal ,

s t ru c tu r es o f d ermal o rig in ar e ho mol og ou s wha tever their shape o f o m o fu c tio n ,

r r r n .

8 E p idermal s tru ctu r es exhibit mor e modi ca tion s an d a g r ea ter c ompl exity o f s tru c tu r e
.

than d o d ermal p ar ts .

9 Th er e
. h as b een n o d en it e pr og r es s iv e e v o l u tion o f t he ex o s kel e to thr o u gh o t th n u e

ve teb ra te g rou p s b u t ea ch gr ou p is p r ovid ed with ex o s k el etal s tru c t ur es co rr el a t ed with its

r ,

hab its an d mode of l ife There is however a c er tain ten den cy for the higher vertebrates to
.
, ,

d vel o p complica ted epidermal s tru ctu r es

e .
58 L AB ORA OR T Y MANUAL F OR VE R E B RA E ANA OM T T T Y
or l es s re pl a c ed p
du r in g d ev el o m en t b y o n e, s ec r eted on e
formin g c ell s b by b . T he s k el eton
is then s a id to b e os s i ed B o n e r o du c ed in t is m an n e
. p
r e l a c emen t h r by p of p r e-
ex is t in g

c a r til ag e is kn o wn a s ca r til a ge b on e .

I n v eStig a tio n of the in ter n al heir bon es s k el eton o f v er t e b rates shows that n o t all o f t
a i e in this m a
r s er b u t so m e o f th em d v el o p di ec tl y f o m the m esen ch m e with ou t p as
nn e r r
, y s

i g throu gh a c a til ag e s tag e

n S ch bon e s a e c al l ed d r mal membr n e o r i v s tin g bo es
r . u r e ,
a ,
n e n .

Th y ar e r eall y der iv d fr m the de mis f the s kin an d are the efo re dermal pl a tes ho mol ogo u s
e e o r o r

t o g a o id s c al es an d t o the p l tes o f the t r tl e s arm o r They a e c on s equ en tly p ar ts o f the

n a u . r

e o s k el eto
x They hav e s u nk i wa d fro m their o igin l p o sition in the skin a d hav e
n . n r r a n

a tt ached thems el ves to the en d o s k el eton with which they a r e n ow s o c l os el y a ssoc ia t ed tha t

they m u s t b e c on sider ed a p a t o f it r .

T he s tu d en t shou l d p a ticul arl y u n ders tan d that c ar til ag e an d m embran e b on es l oo k

r

e ac tl y al ik e ; they hav e the s a m e his tol og ical s tru c t r e an d ch emic al c o m p os ition ; th ey a r e

x u

b th c o mpl etel y form ed bo e ; it is n o t p o ssibl e to d is ti gu ish th m by e amin in g them

o n n e x .

I t is b l y their m n n er f r igi that is d i r e t I n o r d er to d eterm i e which b on es o f the

'

n a o o n e n . n

FI G Dia m s to sho w t he skel eton formin g sep ta in A the tail r egio n , a n d B , the t ru n k region
5 gra
-
. 1 .
,

o f a v er t eb r a t e . a, s kin ; b, n eu ra l tu b e ; c , n o to chor d ; d , b l oo d v essel ; e, d orsal s kel etogen ou s sept m ; u

f,
v en tral s kel etoge pt
n ou s s em ; g, ho riz on ta l sk el et o gen ou s s e tum ; h, myose tu m ; i , e axia ar o
u p p p lp t f
t he myo tom e ; j , hyp i l p
ax a a rt o f the myo tome ; k, c oel om ; l , in tes tin e ; m p, c artil ag es fr om which
-

t he v er t eb ra e a r e d
form e m, b asi o s ; n , in e e a ; o, b asiven tral ; p, in ter d orsal q, in ter
d r al t r v n t r l
m u sc u l ar rib ; r , su b p erito n eal rib . In B n o te p o sit io n s o f the v er t eb ral c ar tila g es an d rib s with res p ec t

t o the s kel etog en ou s se pt a. (A a ft er Kin g sl ey

s Compar a tive An a tomy of Ver tebr ates co py ight b y
r

XfL
,

P . B l akis ton

s So n an d Co m p an y; B from Good rich in P art I o an kes ter s

Tr eatis e on Z ol gyo o ,

co u r tesy o f the M acmill an Co m p an y .

)

en d o s k el eton bon es an d which are m emb an e bon es it is n ec es sary to s tu dy th eir

a r e c ar t il ag e r

e m b ry o ic d ev el o p m e t
n This has b e n d o e fo r the m a j o ity o f th b o es o f the s k el eto
n . e n r e n n.

I o d er t o t a c e the ho mol ogy o f the p ar t s o f the s k el eto n in d iffe en t ve teb a tes it is a b s o

n r r r r r

l tel y ess e tial to k ow which b on es ar e car til ag e bo n es an d which ar e d e m al

u n n This in for ma r .

t ion is g iv en in the f oll ow i g p ag es F o r the p es e t w e m ay s ta te tha t d ermal bo n es o c cu r

n . r n

in c o ec tion with the s k u l l j aw s an d p ec to al g i dl e ( th ese p ar ts a l s o c o ta in car til ag e b on es

nn , ,
r r n ,

o f c ou r s e) All o ther p ar ts o f the e dos k el eton a r e compo s ed wholly o f ca til ag e b o e

. n r n .

B . THE E MB R Y ONI C OR I GN I OF TH E VE R TE B RA E AND RIB S

I . The d evel opm e n t ver tebr ate s kel eton is the ver teb al
o f the s cl er otom e .
The axis o f the r

c l mo u o r b ac kb n e which i c o m p o sed o f a l o g it d in al s e ies o f b on es ea ch o f whichis ca ll e d

n o ,
s n u r ,

a v t br aer e T he v t eb a e a is e f ro m the s cl e o tom es

. er W e hav e al eady l earn ed tha t a
r r r . r

s c l e o to m e is a m as s o f m es en chym e o rig i a t i g f r om the m edial wall o f ea ch ep imer e ( s ee

r n n
 T HE E NDOS K E L E TON : VE RTE B RAL COL UM N AND RI B S 59

F ig . 10 , p .
43 ) T he s cl ero t om e mig r ates to the median r eg ion o fthe e mb r yo an d s u r r oun ds the
n otoc ho d an d n e al tub e I t then g ives ris e to c a til g e which is l a id down a ou d the
r ur . r a ,
r n

n o tocho d T he v er teb ra e m ay r emain p e ma en tly i this c ar til ag in o s s ta t e a s in el a s mo

r . r n n u ,

b an chs b ei g s tiffe ed b y c al ci cation o r they l a ter tra sfo m i to b on e a s in the maj ority
r ,
n n ,
n r n ,

o f v e teb a t es
r T he s tu d en t sho u l d u de s tan d cl ea ly tha t the vert eb ral co l u mn do es n o t
r . n r r

c o m e f r o m t he n o to cho d b t is f o rm ed ro d it T he n o to cho rd is i cl osed i s id e the

r u a un . n n

ve tebral c ol u mn a d is r eadily lo cat ed in this p os itio in the mo r e p imitive shes ; b t in

r ,
n n r u

m o s t v e t eb a tes it is g radu ally s qu eez ed do wn an d d is app e r s e cep t fo r r em a ts b etween

r r a x n n

t he ver t eb a e r .

A s cl e o to me does n o t a s might b e su ppo s ed p r odu ce a ver t eb ra as o n e p iece

r , T he , .

s cl e o t om es a r e s eg me tall y a
r an g ed ther e b ei g a s cl e o t om e m ed i l t o ea ch myo to me
n rr It ,
n r a .

ha s al ea dy b een s tat ed howev er tha t the ver teb rae al tern a te with the myo to mes a d a ise
r , ,
n r

a t the i t ers ec tion o fmyo s ep ta with the m edian s k el etog en ou s r eg ion

n This r esul t is a chieved .

a s f o ll o ws E ach s cl er o to me soon b ecomes d ivid ed by a ver tical s p l it in to an a te io r o r

. n r

cr a n i l h al f p o s t e ior to the p l an e o f the p ec edi g myo s ep tu m a d a po s t e io o r

a ,
r dal hal f r n ,
n r r cau ,

an ter io r t o the p l a e o f the s cc eedi g myos ep tu m A ve t eb a a rises f om the fu s io o f the

n u n . r r r n

n otochor d
myoto me
o f scl ero to me

FI G . 1 6 .
- p
T wo s tages in the d ev el o men t o f the ver teb ra e rom the s cl ero tomes , o nl y o n e sid e o f f
the b o dy b ein g shown A , divisio n o f the s cl ero tom e in to a n terio r an d o s terior hal ves
.
B , u nion o f p .

f
t he posterio r hal o f o n e s cl er o to m e with t he a n t erio r ha l o f the su cceed in g s cl ero t ome t o o rm a f f
m r ey s Textb ook of E mbr yol ogy, co u r t esy o f the W B S au n der s

ver teb ra ( F r o P re
. n tis s a n d A . .

Co m p an y .
)

p os terior hal f o f o e s cl erotome with the an te ior hal f o f the su c ceedi g s cl ero tome T he
n r n .

a n t e io r h al f o f o e s id e o f the v e teb ra c o mes f o m the p o t e io r h l f o f a s cl e o to me a d

r n r r s r a r n

the p os te io r h al f o f o n e s ide o f the v e teb ra f r om the an t e io r ha l f o f the s cl e o to m e

r r (I t r r .

shou l d b e u d e s to o d tha t the s cl e o tom es o f the t wo s id es o f the b o dy a is c o o p e a t e in the x r

-
n r r

fo mation o f each ve teb a ) I t thu s happ e s that the c en ter o f the ver teb ra in ters ects a
r r r . n

myo sep tu m a d tha t the v e t eb r a e al tern a t e with the myo to mes ( s ee F ig

n r .

2 The ar cu li
.
I n the for ma tion o f a ver tebr a d e nite p ai ed c a til ag es kn own a s
a a. r r

ar cu al ia a ise in the s cl ero to me

r Theoreticall y there are fou r p ai s o fa cu alia to each v e teb
.
r r r ra .

p
T he os te o r i r h a l v e s o f t h e t w o sc l e r o t o m e s o f e ac h s e g m e n t w h ic h f o m t h e a t e io r h a l f o f r n r

a v er te b r a p r o d u c e t w o
,
p a ir s o f a r c u a li a a d o r s a l p a ir a b o v e t h e n o t o ch o d c a ll e d t h e r ,

b as ido rs al s a n d a ve n t
,
r a l p a ir b e l o w t h e n o t o c h o r d c all ed t h e b as i v e t ra l s S im il a
,
l y n . r

the a t e o
n r i r h a l v e s o f t h e t w o s c l er o to m e s o f e a c h se g m e t w h i c h t o g e th e r f o rm t h e n ,

po s terio r hal f of a ver tebra p rodu ce a d o sal p air o f a rcu alia the in t rdors al s a d ,
r ,
e ,
n

a v e t
n ra l p air t h e in te r ve n tr a l s ,( s ee F ig Th e r e m a y b e o th e r e l
.
e m e t s i a d di t ion n n

b
a o v e an d b el ow t ese h u
s c h as t h e s u p r a dor
,
s a l s a d i f r a v e tr l s b u t t h e s e a e n no t n n a ,
r

o f g e eral o c c rre ce
n T he m e m ub e rs n o f t.h e c h ie f p a irs o f a cu l ia t d t o f se t o g e t h e r r a en u

m ediall y p r o du c i g a r c es oh r p i e ce sn w h i c h s t ra d dl e t h e n o t o ch o d a b o v e a d b e l o w T h s r n . u
 60 L AB ORA OR T Y MANUAL T
FOR VE R E B RA E ANA OM T T Y
d
d in terd
b oun
Seg
ary o f
men t no toc hor d
d
b as i o rsal

M i o

a rc h

b asiventral
a n al

in terven tral
b asiventral

FI G . I 7
Dia g ram s t o S ho w the
. a rcu alia f ro m which the v er teb rae ar e fo rm ed A , side view, .

sho win g the f o u r a rcu alia o f o n e s id e t o each segmen t B , c ross s ec tio n throu gh the b a siven tral s a n d
-
.

b asid o r sal s , sho win g their r el a tio n t o the n o tocho r d C, l a t er s ta ge o f the sam e cro ss s ec tio n as in B ,
.
-

sho win g u nio n o f the b asid o r sal s t o fo rm t he n eu r al a rch a n d o f b asiven tr al s to form t he haem a l a r ch .

Dia p f
throu gh d ev el o in g ver teb r ae t o s ho w o rmatio n o f the
FI G . 1 8 .
g ram matic sec tio n s a rche s

an d the cen t ra . A , earl y s tage, k B D,

s ho win g t he s el et o g en o u s r egio n s b a ro u n d t he n o t oc hord
.
-

p pp
formation o f the c ho rd al ty e o f c en trum : B , a earan ce o f the a r cu alia f a n d the shea th o f the n o to
c ho r d e; C, in va sio n o f sk el et o n -f o rm in g c ell s f ro m the a r cu al ia in to the s hea th o f the n o t ocho r d , a s a t j;

p
D, c om l etio n o f the cho r d al cen trum in the shea th o f the n o tocho r d , co m l etio n o f the a r ches b y fu sio np
o f the a r cu alia , an d u n io n o f c en trum a n d ar c hes p
E an d F , fo rm a tio n o f t he ericho rd al o r a r ch t y e
. p
o f cen trum : E , a pp
ea ran c e o f the ar cu alia f a n d the shea th o f the n o to cho r d e; F , formatio n o f t he

c en t ru m a n d the a rches b y t he f u sio n o f the a rcu alia a r ou n d a n d a b o v e a n d b el o w t he n o tocho r d .

a , n o to chor d ; b , sk el et o g en ou s regio n s ; c n eu r al c a n a l ; d , haema l ca n al ; e, shea th o f the n o t oc ho r d ;

,

f, ar cu alia ; g, n eu ral a rch; h, c en trum ; i , haem al a rch ( From W iedersheim s Compar ative Ana tomy

.

of Ver tebr ates , co u r t esy o f the M acmill an Co mp an y )

 T HE E ND OS KE L E TON : VE RTE B RAL COL UM N AND RI B S 6 1

in g the n eu r al tu be ; the in ter do r sa l s si mil arl y f o rm an in ter cal ar y a r ch, the b as iv en tr al s a

haemal a r ch, an d the in t erv en t r a l s ,
( Th u s p r im an
it iv el y eac h
in ter ha emal ar ch s ee Fig .

s egm en t o f the v e t eb a t bo dy is p ovided with two dors al a rches o n e an terior an d o n e

r r e r
,

po s terior a d two s imil arly p l a ced ven tral a ches Su ch a co dition actu all y o ccu rs in the
,
n r . n

a dul t s o f som e o f the l o wer ver teb a tes This te den cy fo r the p od u c tion o f two ver teb r ae
r . n r

t o each s eg m e t is k own a s dipl s p n dyl ( s ee K pp

n n I m v o b o h r
y 5 n o s t e t e ra tes o wev e , . 1 r
, ,
ther e is a s in g l e ve teb a t o ea h s eg m en t ( l ter ati wi h h m h
r
g t r
t e se g e n t);
c
t is c o ditio n a n n n
a is es throu gh the l o s s o f s m e o f the a r c alia o r th eir f s ion
r
T he m an er o f f o rma tio n o f
o u u . n

t he ve teb a e f o m the a cu alia is d iffer en t in dif

r r r fer en t ve teb r a te c l ass es a d is d esc ib ed in
r r n r
co nn ec tion with ea ch g o p r u .

3 Th f.o m a ti n o f th
e e c e tr u m
r T he co mp l e ted ver teb r a c on s is ts n o t o nl y o f the
o n .

a ches ar is in g f o m t he a c al ia b u t a l s o o f a c e tr al mas s the b od o r c v b

r r r u
y n tr m o f t h e e t e ra n
, e u r .

T he c en tru m is p o du c ed in two ways in ve teb ra tes : ( a) by dev el o p i

r
wi th i t h h th of r n
g n e s e a

n eural s l
p ne

ne u ral a na l

FI G . 19. Dia g r ams o f ty pi c al ver t e b ra e . A , t ail v er t eb r a o f a tel eo s t sh . B , tru n k ver teb r a

o f the sa me, S ho win g O p en in g o f the ha emal a rch to fo rm the t ran s verse proces ses C ver teb ra of a .
,

la n d v er teb r a t e , sho w in g p ti l l y
ar c u ar r el a t io n o f r ib s to the c en t ru m ( After K in g sl ey s Compar ative
.

An a tomy of Ver teb r a tes , co py ight b y Pr . B l a kis ton

s S on an d Co m p an y . )
the n o tocho r d an d b eco mi n g s eco n da r il yf u se d to the arcu alia ; ( b) f ro m the a r cu al ia , eithe r

by fu s ion o f the ba s es o f the ar c hes ,

or f ro m en tir e a r cu al ia ( s ee F ig . The rs t typ e o f
c en tru m is call ed a chor dal c en t ru hs ; the s eco d typ e
m an d is c har a c t er is tic o f el a s m o b r an c n

is n am ed a p r ich r dal o r rch c en tru m a d is f ou d in p c ticall y all o ther ve teb ra tes

e o a T he n n ra r .

c e tru m in b o th cas es is f o r m ed ar u n d the n o to cho r d which is i cl osed within it

n o n .

I n the emb yo n ic d ev el o p m e t o f a ver teb r a the d o s al a r cu alia a ris e r t the ven t al

r n r s , r

a c u alia n ex t a d the c e tru m l as t

r ,
T he s ame p rog es s ion is ob s er vabl e in the phyl og e y
n n . r n

o f v e teb a t es
r I n p r imitiv e ver t eb a tes s u ch as the cy cl o s tomes o n l y the d or al a cu alia a r e
r . r s r

p rese t s traddlin g a larg e p ers isten t n o tochord ( s ee Fig 2 I A) I n m a y g an o ids all of the
n , . . n

a r c alia a pp ear s u rr o u d in g the l a g e n o tochor d b u t the c en tru m has n o t yet d evel o p ed

u n r
; ,

s u ch ve t eb a e a r e d es ign a t ed as a c n tr ou s ( Fig
r r I n the higher shes an d all l a d e . n

ve teb ates well d evel o p ed c e tra a e p r esen t a d the n o to cho d is r edu ced o r ab sen t
r r -
n r ,
n r .

4 Th par ts of a typical ver te b a

. e A typ ical comp l eted ver teb a con s is ts o f a cen tr u m r . r

i cl o si g the n o to cho d a d o r s al n eur al ar ch in cl o s in g the n eu ral tu b e a d a ve tr al ha ma l

n n r , ,
n n e

arch i cl o s i g bl o o d ves s el s
n Neu r al a d haemal arches a e common l y p o l o g ed do s ally
n . n r r n r
62 L AB ORA OR T Y MANUAL FOR VE R E B RA E ANA OM T T T Y

an d ven t y
p ectively i to n eu r l a d haemal spin s I n a ddition th c en t m co mm l y
r all ,
r es ,
n a n e . e ru on

p r es en ts va io s p o j ectin g p o ces ses kn own as apophys es servin g for a ticul tio s with adj o i
r u r r , ,
r a n n

in g v er teb r a e o r with ib s o r fo r the a t ta chm en t o f mu s cl es rI n l ower fo m s the e a e n o t r r r

u n co mmo l y t wo c e t o r a ches o e an t e io r a d o e p os t e io r t o each m et a me e b t

n n ra r ,
n r n n r ,
r ,
u

u s all y o l y o e o f these s t ru c tu es is p es e t t o each s eg m n t o wi g t o the l os s o

u n n fu s ion of
r r n e ,
n r

t he o the s F igu r e 9 ill s tr a te a typ ical ve t eb a with its p ar ts

r . 1 u s r r .

T h i i f r ib E ac h v e rt b is theo e tic all y p o v id ed with a p ai o f r ib s which

s. e ra r r
5 e o r
.
g n o r

p o j ect o t f om the c en tru m i to the b ody wall a d s erve to s tr e g then the l atter T he
r u r n n n .

r ib s l ik e o th er p a ts o f the a ia l s k el et on arise in the s k el etog e ou s s ep ta

,
r The e are two
x , n . r

k i ds o f ib s b o th o f which arise in the myosep ta a d c on s equ e tly ar e s egme tal in a ra ge

n r ,
n n n r n

me t n O e typ e o f r ib is fo rm ed a t the i te s ec tion o f each myo sep tu m w ith the ho riz o t l

. n n r n a

s k el et og e ou s s ep tu m nS in c e the ho izo n t al s ep tu m d ivides the myo tom es i to d o s l an d ven

. r n r a

t a l h al v es ( ee F ig 1 5A p 58 ) s ch r ib s l ie b etw ee mu scl es an d h en c e ar e n am ed i ter mu s

r s .
, . u n n

cu l ar rib s They ar e al so kn own as dorsal or tru e ribs I n termu scu l ar ib s a e charac te is tic o f
. . r r r

the m a j o ity o f ve teb a tes T he r r r .

s eco d typ e o f r ib a ris es a t the n

ln eura s pin e
p oin t s o f in ter s ec tion o f t he m yo

s ep ta with t he v en t al skel eto g e r

n ou s sep tu m o it s d e iv a t iv es It r r .

w ill b e r e cal l ed tha t in the t ru k n

r eg ion t he ve t al s ep tu m is s p lit n r

i to two l at eral s ep ta o wi g t o n ,
n

the in terv en tion o f t he c o el o m an d

its c on ten ts o n the v en tral s id e o f

the b o dy ( Fig 1 5B )
s ee . . T he r i bs
u n d er c on s id er a tion a pp ear a t the
poin ts o f in ter section o f the myo
s ep ta with thes e v en t ol a t a l r er

sep ta They l ie j u s t ou tsid e the .

c o el o m ic li i g b etw een the c o e n n ,

F Di g mm t i
IG . 20 ,
ti t h a g h
ra t h t k
a l o m ie w
c c ros s - s ec all a n d t h e
on m u s c l e l a
r ou y e r e ru n .

o f v a t b t h
er e w i g t h l t i
ra e s f t h ib o t nth k l Th
e re a e y a e d e
on o s i gn a t ed as e rs b ps r i o e s e e r u e ~

t g
o gi
en ou s r e d th p i t i f th
on s an t w k ind f ib
e t
os n g a l r ib s ;
on s o a l s o as v tera l o o
r f l e s o r s. o en a s

H r ib s } a r e char
m H T b k f Z l t f , '

( F P
ro k d War t er a n i g y aS y C S ex oo 0 oc o ,
cou r es o .

th M mill
e C mp y )
ac an o an
ac t eris t ic o f the b o y shes B o th n .

typ es o f rib s m ay b e reg a d ed a s r

e ten s io s o f the
x b as iv en t ral ar cu alia th
n e ir p o s i ti o n in t h e m y o s ep t u m b e i g s h i
,
f t ed d o r s ll y n a

o r v en trall y in d if v
fe e t er t e ra tesb I n m a y
r nsh e s b o t h k i d s o f r ib s a e .
s im u l t a e o l y n n r n us

p es en t on each ve teb a T he two k in ds o f r ib s an d their morphol og ical r el tio s to the

r r r .
a n

b ody wall a e ill u s t a ted in F igu e 2 r r r 0 .

b k v
R i s l i e e te ae a b ,
pp e a r s t a s co llre c t io n s r o f m
,
e e c h y m e w h ch r l a t er t a s f o m s s n i r n r

in to ca til ag e b
T he r i s a e
r r p e m a .e n tl y c a r t il a g i n o u s i n so m e s h e s rb u t i n no t h e r v e r t eb r a t es

a e o s s i ed a t l ea s t i
r p ar t ,
n .

For the co mp rative an atomy o f the ver tebr al col umn a d ribs the s tu de t h u l d
a n ,
n s o co n

sul t K W , ,
an d Wd .

T he t erm p is h er e id d
l eur al as a
a v o e ,
o w in g t o ppli d t
t h e f a c t t h
e a t it i
o rib s
s di ff eren tl y em l oy ed p
in d i eren t t ext s K a d W u se
n th e t er m l eu
.
r al a s s yn o n y m ou s w it h in t erm u s c u l a r ,
w hil e L p
u ses it a s

syn o ym o
n u s wit h s u b erito n eal T h e l a t t e r u sa g e a p
ea rs t o b e co rr e c t . pp .
 THE E NDOS KE L E TON : VE RTE B RAL COL UMN AND RI B S 63

a c en trum In o ther c ases d ip losp on dyly is eviden t We m ay glanc e a t a few

.
.

of these .

I . c u m o
The ver ebra ol n f the s tu r g eon I n the s tu rg eon the ver tebral col
t l .

u m n i s Very p ri mi tive a cen trou s an d c onsis tin g of sep ara te ar cu alia E x amin e
, .

FI G . 21 Some p rimi tiv e v er teb ral

. c ol um n s . A , c ycl os tomes ; the vert eb ral col u mn co n sis ts o f a

ro w o f a r ches , t wo arches to ea ch s eg m en t , r es tin g o n the d o r s al f

su r a ce o f the n o to cho rd . B , s tu r geo n ;
the ver teb r al colu mn p
co n s is t s o f s e a r at e a rcu alia p ar t iall y su r rou n d in g the n o tochord . C, ta il r egion

o f Amia , S ho win g t he t wo c en t ra , k a n d l , to ea ch s egm en t , ill us tra tin g dip l ospo n dyly D, vert eb ra e
.

of an ex tin c t am phib i
b el o n gin g to the S tegocephal a ,
an sho win g the a rcu a l ia a r ou n d the n o tochord .

a , s egmen t al b l oo d v essel s ; b , n erv e ; c , n eur a l t u b e ; d , n o tochord ; e, b a sid o rsa l o r n eu ra l arch; f, b asi "

p
ven tral ; g, in t er dors al ; h, in t erv en tr al ; i , n eu ral s in e o r s u rad orsa l ; j , rib ; k, p
o s terio r cen t ru m ; p
l, an terio r c en t ru m , for ea ch s eg men t ; m, ha em al a r ch; n, ha emal s pi ne; 0, b l ood v essel s . (A an d B
after Go odrich in P ar t IXo f L ankes ter s Tr eatis e on Zool ogy cou rt esy o f the Macmill an Compan y

, ,

C fro m Zitt ell D fr om Kin g sl ey s Compar ative Anatomy of Ver tebr ates a ft er Zit tell copyright b y

, , ,

P B l a kis to n
.

s So n an d Co m p an y
.
)

the dem ons tration speciin en s an d c omp are wi th Figu re 2 1 8 The larg e no to .

chor d tr av e rs es the c e n t er of the ver teb ral column an d is covered dorsall y an d

64 L AB ORA OR T Y MANUAL FOR VE R TE B RATE ANATOM Y

ven trally by the car tila ginou s ar cua lia On the dorsal side are the large ba s i .

dor s al s u n i t ed to form a n eu ral ar ch t o pp ed by a n eu ral s pin e ( su pr a dor s al )

,
.
B e tween the bases of the b asido r sa l s are the s m all in ter dors al s The ven tro .

la teral region s of the no to chord are covered on ea ch side by the large b asiven tr al s
ea ch of whi ch bears a p roje cting p ro c ess No te tha t the b a siv en tral s of the two .

sides do n o t m ee t below the no to chord B e tween the ven tral p or tions of the .

b asiven tr a l s are the s m all in terven tr a l s .

2 The tail vert e b rae of the b own

.
E x am ine a dem o n s tra tio n sp e c im en .

of the tai l of the b own Comp are wi th Figu re 2 I C No te tha t the n o to chord
. .

is n o t visible bein g comp le tely in closed by the ver tebrae In the la tter there
, .

is an al tern a tion of a ver tebra bearing dorsal and ven tral ar ches wi th on e devoid ,

of ar ches The form er co n sis ts of the fu s ed b asido rsal s and b asiven tral s form
.
,

in g n eu ral and haemal ar ches an d a c en tral cen tru m The la tter co n sis ts of the
'

fu sed in ter do rsal s and in terv en tral s There are thu s two ver tebrae to ea ch .

segm en t of the body a con di tio n n o t u n comm o n in p rimi tive ver tebra tes
, .

3 Ver
.t e b rae o f S teg oc epha l a S tu dy F i gu re 2 1 D or K F.igu re 4SE p age 53 , , , .

These are rep resen ta tio n s of the ver tebrae of ex ti n c t Amphibia E a ch ver tebrae .

c onsis ts of s ix p ie c es of whi c h fo u r are visible fro m ea c h side There is a dorsal .

arch comp osed of the fu sed b asido r sal s a large ven tral pie c e co mp o ed of the ,
s

fu sed b asiven tral s and two sm all p os t erior p ie ce s o n ea ch sid e the dorsal on e
, .

being the in terdorsal the ven tral one the i n terven tral The large n o to chord
, ,
.

is in closed by these ar cu alia .

D . THE VE RTE B RA L COL UMN OF THE G

DO F I S H

1 . Cros s -
se ction of the tai l O b tain
a c ross se c tio n throu gh the t ail of the
.
-

do g sh an d s tu dy the cu t s u rfa c e B e s u re tha t you have a se c tio n p assi n g

.

throu gh the j u n c tion of s u c c essive ver tebrae an d n o t thro u gh the c en ter of a

ver tebra The c en ter of the se c tion c on tains the ver tebra co mp osed of car ti
.

,

lag e a clear rela tively sof t ma terial Ou tside of the ver tebra is the layer of
, .

volu n tary mu s cl es c on sis ti n g of a nu mber of l eaves ea ch sep ara ted from its
, ,

n ei ghbor by a p la t e of c onne c tive tiss u e the myoseptu m or myocomma E a ch ,

.

leaf is a myotome or mu scl e segmen t The m yo tom es app ear in whorls in the .

se c tion be ca u se they are zi gz a g in form like those of Amphioxu s E a ch cross .

se ction will co n seq u en tly cu t a n u mber of m yo tom es The mu s cle segm en ts ar e .

somewha t in dis tin ctly divided in to dors al an d ven tr al p or tion s by a p ar ti tio n

whi ch ex ten ds from the cen tral regio n of the ver tebra to the ski n wher e it m e e ts
a line the la teral line ru nnin g alon g the ou ter su rfa c e of the ski n This conn ec
, ,
.

tive tiss u e p ar ti tio n is the hor izon ta l s kel etogeno u s s eptu m The mu s cles above .

it are the dor s al or epa xial mu s cles those below it the ven tr al or hypaxial , ,

mu s cles .

Wh th tien p th
e s ecg h t
onh t f t h
a s ses t b f
ro u y f m i g
e c en er ot f w ill e v er e ra , ou r ra s or n a so r o cross

be seen in the ver t eb ra . These a r e ar eas o f cal ci ca tio n .

66 L AB ORA OR T Y MANUAL F OR VE R TE B RATE ANA OM T Y

c oelom in closes the vis c era som e of whi ch will be observed to be s u sp en ded by a
,

deli ca te m em brane the dors al mes en tery from the q ian dorsal line of the co e
, ,

l omic wa ll S tu dy the v er tebra whic h app ears in the middle of the myo to m es
.
, .

I t co n sis ts of c en t ru m and neu ral ar ch simi lar in ap p earan c e to t hose of tail ,

ver tebrae ; b u t the haem al ar c h app ears to be absen t I t is rep resen ted by a .

p air of s m all c ar tila g es a t the sides of the ve n t ral p ar t of the c en t r u m These ,

c ar tila ges known as tr an sver s e pr ocess es rep rese n t the s tu mp s of the hae m al ar c h
, , ,

E
FI G . 22 .
Diagram s to S ho w the co m p o sitio n o f the v er t eb rae in the differ en t vert eb ra te cl a ss es ,

ac co r d in g to G ado w . A , el asmo b ran ch, v er t eb r a c on sis tin g o f chor d al c en trum a , b asido r sal s b , in t er
d o r sal s c, a n d b a siven t ral s d . B , t el eos t , p
v er t eb ra c om o s ed o f all o f the a r cu alia fu sed tog ether .

C, p s eu d ocen tr ou s v er t eb ra o f a u r od el e ; 1 ,
e a rl y s t a g e sho win g the a rr an ge m en t of the ar cu alia ;

2 , a dul t s t ag e, sho n fu sion o f the a r cu alia al on g the dotted lin es ; n o te s plitti ng of the in ter dorsa l s
an d in ter ven tr al s t o form the en d s o f the v ert eb r a . D, n o tocen trou s v er teb ra o f a n u ran s ; I ,
earl y

s t a g e, showin g t he a r cu alia which form t he v ert eb r a ; 2 , a dul t s t ag e, a r cu alia fu s ed al o n g the d o t t ed

lin es ; n o t e en d o f the p rocoel ou s c en tru m form ed b y the in ter d orsal E g as troc en t rou s v ert eb ra .
,
of

re p til es , b ir d s , an d mamm al s , cen t ru m f o rmed o f the in terven t r al e, the b asiv en tr a l d f un c tio n in g as

the in ter ver teb ral ca r til ag e . F, co m p o sitio n o f the a tl a s a n d a xis o f the all ig a t o r , acco r din g to G ad o w ;

the cen tru m o f the a tl a s 2 fun c tio n s as t he od o n toid p r ocess o f the ax s i . a , cho rd al c en tru m ; b , b asi
d o r s al o r n eu r al a r ch; c, in ter dorsal ( in ter cal ary a r ch in A) ; d , b a siv en tr al or ha emal a rch; e, in ter
ven tr al ( cen trum in E an d F) ; t , a tl a s ;
a y, a xis ; 2 , od o n toid p ( in terven tral o f a tl as)
roc es s .
(F a fter
G a dow in the Cambr idge Natu r al His tor y, cou r t es y o f the M a cmill a n Comp a n y ) .

whi ch m ay b e regarded as havi n g op en ed o u t and shif ted to a m ore la teral p osi

t ion the ar c h p or tion disap p earin g
,
E x a mi n e the hori zo n tal skele tog en o u s s ep .

tu m c arefu ll y and n d wi thin it b y p i ckin g away the mu s cles if n e c essary a , ,

sl en der c ar tilage o n ea ch side whi c h ar ti cu la tes wi th the t r an sverse p ro c ess

,
.

These c ar tila ges are r ibs and sin c e they are lo c a ted in the horizon tal skel etoge
,

no u s sep tu m t hey are in termu s cu lar ribs There is of cour se a pair of such ribs .
, ,

to ea c h ver tebra Draw the se c tion . .

 T HE E NDOS KE L E TON : VE RT E B RAL COL U M N AND RIB S 67

E . VE RTE B RA L CO L UMN OF TE L E OS T S

I . ver tebrae
The tail .
E x amine
a s ep ara te dri ed tail ver tebra of an y bony
sh No te tha t the ver tebra is very mu ch harde r an d more op aqu e than the
.

do g sh ver t ebrae owi n g to the fa c t tha t it is c o mp osed of bo n e

, I de n tify the .

sam e p ar ts as already seen in the d o gsh ver tebrae : the bi con cave or amphicoe
lou s cen tru m bearin g a mi n u te canal in its cen ter for the no to chord the neu ral ,

arch termi n a tin g in a very lon g sharp n eu ral sp in e the haemal ar ch termin a ti n g ,

in a si milar hae m al sp ine The n eu ral canal or sp a ce i n closed wi thi n the neu ral
.

arc h is g en erally s maller than the ha emal can al i n closed by the haemal ar ch ; in this
way the dorsal an d ven tral sides of the ver tebrae may be dis ti n gu ished The .

spin es are dire c ted p os teriorly In som e sh there are two n eu ral sp in es to ea ch
.

ver tebra an an terior and a p os terior on e ; the se cond o n e p robably c orresp onds
,

to the i n t er c alary ar ch of the do g sh Draw a ver tebra . .

2 The tr u n k vert e b rae

.
Ob t ain a sep ara t e dried tr u n k ver tebra of a bon y
.

s h I den tify as b efore the c en tru m and the neu ral ar ch an d neu ral sp i n e
.
.

The haem al ar ch and sp ine app ear to b hmissi n g I n s tead there is a p air of .

p roje c tio n s a t the sides of the base of the c e n t r u m to ea c h of whi ch a lo n

g sle n der
r ib is ar ti cu la t ed These s tump s or tr an svers e processes are the Op en ed bases of
.

the hae mal ar ch Draw a ver tebra wi th ribs

. .

T he ver tebrae ar e ar ch cen tra fo med by the fu sio of the bas es of the
cen t ra o f t el eos t r n

b as ido sal s an d b asiv e t ral s chiey al thou gh in te do s al an d in terven t al el eme ts may b e

r n ,
r r r n

ad d ed to the e d s o f the ce tra ( F ig

n T he n e al ar ch is fo rm ed by the b as ido s al s
n . ur r ,

the haemal a r ch by the b a siven tr al s .

3 . S e ction thr o u gh the tr un k a b on y su ch a se ction iden tify

of sh In.

the p ar ts already des c ribed for a si m ilar c ross se c tion of the d o gs h No te the -
.

mu s cle segm en ts the c en tru m a n d n eu ral ar ch an d sp i n e of the ver tebra and

, ,

the c oelo m wi th its li ni n g F ind the ribs lo ca ted j u s t o u tside of the c oelo m i c
.

lin i n g The ribs are in a dieren t p osi tion than those of the dogsh whi ch are
.
,

in the hori zo n tal skele togenou s sep tu m The ribs of teleos ts are si tu a ted j u s t .

ou tside of the c oelo mi c wall s They are false or s u b p eri to n eal ribs Make a
. .

diagram of the se c tio n sho wi n g the p osi tion of the ribs ,

.

4 F u
. r ther s t u d

y of r ib s S o m e shes have t w o ( or m ore) p airs of ribs

sim u l tan eou sly on ea ch ver tebra E x amp les are P olypteru s a crossop terygian .
,

g a n oid fro m A fri c a and m any t eleos

,
t s in c l u di n g m e m bers of th e sa kh o n
,
h erri n g , ,

and pike families E x amine the skele ton of P olypter u s an d n o te two p airs of
.

ribs a tta ched to ea ch ver tebra The dorsal p air a tta ched to the tran sverse .

p ro c esses of th e ver t ebrae are th e t r u e or in t erm u s c u lar ribs ; the ve n t ral p air

loosely a tta ched to the ven tral su rfa ce of the cen tru m are the false or su b per ito
neal ribs Ver tebrae of shes like the salm on may also be ex ami n ed or se ction s
. ,

thro u gh the tru n ks of s u c h shes No te the ven tral or su b p eri ton eal ribs a n d
. ,

the dorsal or in term u s cu lar ribs ( also called ep ip l eu r al s) lo ca ted in the hori z o n t al
68 LABORAT OR Y MANUAL T
FOR VE R E B RA E ANA OM T T Y

skele togen ou s sep tu m Addi tio n al ribs m ay also be p r esen t ar ti cu la ted wi th

.
,

c en tru m or n e u ral ar c h a n d ex t e n di n g o u t in t o the m yosep ta be tw ee n m yo t o m es .

I t appe ars tha t in teleos ts ribs m ay be form ed ht alm os t any level of the
m yosep ta Make a diag ram S howin g the ver tebra and its ribs
. .

5 The vert ebr al c ol u mn a s a whol e

.
S tu dy the e n tire m o u n ted skele ton .

of a bo n y sh No te tha t the axis an d s tron ges t p ar t of the colu mn co n sis ts of

.

the c e n tra whi c h are joi n t ed to g e ther to form a lo n gi tu di n al series

,
The .

ver tebral c olu mn is di visible i n to two regi o n s a tru n k region an d a tail r egio n , .

I n the form er the haem al ar ch es are red u c ed to s tu m p s the t ran sverse p ro c esses , ,

to the e n ds of whi c h the lo n g sle n der ribs are ar ti cu la t ed In the tail re gio n .

ribs are la ckin g an d haem al ar c hes are p resen t Observe the transi tion be twee n .

tru n k and tail re gio n s In the p os terior p ar t of the t ru nk there is a gradu al

.

elon ga tion of the tra n sverse p ro cess and a redu c tion of the ribs At the b eginnin g
'

of the tail region the red u c ed ribs n ally van ish and the elon ga ted tran sverse ,

p ro c esses f u se to for m t h e hae m al ar c h N o t e t ha t th e n e u ral c a n als of s.u c c e ssive

ver tebrae toge ther form a co n ti n u ou s ca n al whi ch in l ife con tain s the spinal
c ord . S im ilarly the haem al canals of the tail regio n form a c o n tinu o u s cavi ty
in closing in life blood v essels At the en d of the tail observe the enl argem en t .

a n d a t t en i n g of the haem al ar ches form in g wha t are c alled h yp u ra l bo n es w hi c h ,

su pp or t the tail n The las t cen tru m (p robably c on sis tin g of several c en tra)
.

form s an elo n ga ted slen der bo n e ( the u r os tyl e) whi ch tu rn s sharp ly up ward ,

p rod u c i n g an asy mm e t ry ; b u t th e sy mm e t ry of th e t ail is res t or e d by the arran g e

m e n t of the hyp u ral bon es Tails of thi s kind are called homocercal
. .

F . VE RTE B RAL COL UMN OF AMP HIB IA

I . The vert ebral o d ele s l mn

S kele t o n s of u rodeles s u c h
co u of ur Whole
as Nectu r u s an d Cryptobran chu s are n eeded for this s tu dy Wi th su ch a skele to n .

before you no te the p ar ts of the ver tebrae of differen t regio n s The tail ver tebrae .

are sim ilar to the tail ver tebrae of shes E a ch p ossesses a low neu ral ar ch a .
,

haem al ar ch an d a cen tru m beari n g tra n sverse p ro cess es I n the tru nk re gio n
,
.

the ha em al ar ch is absen t an d the tran s vers e p ro cesses bear ribs sep ara ted fro m
, ,

them by a s u tu re The ver tebrae are ar ti cu la ted t o ea ch o ther by p ro cesses

.

kn own as zygapophyses These consis t of a p roje ctio n on ea ch side of the n eu ral

.

sp in e whi ch ts over a sim ilar p roje c tion arisi n g from the an terior en d of the
s u cceedin g ver tebra Thu s ea ch ver tebra has a p air of pr ezygapophyses on its
.
,

an terior en d whose ar ticu la tin g s u rfa c es fa c e up ward an d a p air of post ,

zyga pophys es o n its p os t erior end whose ar ti cu lar s u rfa c es fa c e downward .

Th ese zy gap op hys es yoke the ver tebrae to ge ther .

2

I n s hes t he t ran s ver s e p r o c es s es wer e s ta ted t o b e the s t um p s o f the ha emal ar che s . It s ee ms

p ro b a b l e tha t t he t r an svers e p r o ces s es on the vert eb ra e o f l a n d vert eb ra t es ar e n o t o f this kin d b u t a re

la t eral ou t g ro wths o f the c en t ru m .

Zyga pophy ses a r e p o o rl y d evel o p

ed or ab s en t in shes .
 T HE E NDO S KE L E TON : VE R E B RAL COL T U MN AND RI B S 69

The ver tebral column of u rodeles consis ts of fo u r regio n s n ot very sharp ly ,

marked o ff from ea ch o ther The r s t regio n the cervical or n eck region con
.
, ,

S is ts of one ver t ebra the rs t ver t ebra or a tl a s j u s t ehind sk ll servin

, b the u a n d g ,

as a s upp or t for the sku ll The a tlas la cks ribs F ollowin g the c ervi c al regio n
. .

is the lo n g tr u n k region of similar rib beari n g ver tebrae There is n ex t a region -

.

c o n sis ti n g of b u t o n e ver tebra This is the ver tebra to whose ribs the hin d legs
.

are a tta ched I t is called the s acr al ver tebra and its ribs are known as s acr al
.

ribs Thi s region of the ver tebral column is the s acral region or s acr u m P os
. .

terior to the sa c ru m is the cau d al or tail regio n c o mp osed of ver tebrae whi ch ,

la ck ribs and bea r haem al ar ches in mos t cases B y m ovi n g the ver tebrae ap ar t .

n o t e tha t the e n ds of the cen t ra are c on c ave t ha t is the c e n tra are a m hi c oelo u s
p , ,

as in mos t shes .

The a tta ched en ds of the ribs (Nectu ru s) are forked in to two p ro cesses or
heads a dorsal tu b ercul ar head and a ven tral capitu l ar head whi ch ar ti cu la te
,

wi th similar b u t less m arked dorsal and ven tral p roje c tio n s of the tra n sverse
p ro c esses The
. ribs are in t er m u s cu lar ri b s .

T he ver tebrae of the u rodel e Amphibi a differ f rom those of all o ther l an d ver teb rates in
the m an n er in which the c en tra ar e fo rmed I n the tail r eg ion the cen tra a r e p o du ced by
. r

the fu s ion of the b as es o f b as idor al s an d b a siven t ral s the fo mer g ivi g r is e a l s o to the n eu ral
s ,
r n

a ches the l at ter to the ha emal a r ches

r ,
I n the trun k eg ion the cen tra an d n eu r al a r ches
. r

ar e fo rmed from the b asido rsal s the b as iven tr al s h avin g disapp ear ed
, I n t erdo r sal s an d .

in terv en t r al s ar e ab sen t o r r ep r esen ted by the well d evel o p ed in terve teb al car til ag es fo r m -
r r ,

in g p ad s b etween su cces s ive c en tra In s o me u o d el es thes e car til ages s p l it hal f fu s in g to

. r ,

the ver teb r a in f ron t an d h alf to the v er teb ra b ehin d T he ver tebrae o f u ro del es a e d es ig . r

n a ted as pseu d cen tr ou s v er tebra e a s the c en tr u m do es n o t co r es p on d to that of the m aj or ity

o ,
r

of l an d ver teb r at es ( s ee Fig 2 2 C p .

, .

2. Thevert ebr al col u mn of an anu ran amphibi an the ,

frog The ver tebral
9

c ol u m n of the fro g is sp e ciali z ed in tha t the c a u dal ver tebrae are all u ni ted

in to one pie c e the u r ostyl e or coccyx There is as in Nectu r u s o n e c ervi cal

,
.
, ,

ver tebra the a tlas Thi s is followed by s even tru nk ver tebrae These have
,
. .

low neu ral ar ches no haemal ar ches and cen tra beari n g lo n g tra n sverse p ro cesses
, ,
.

R ibs are app aren tly absen t b u t em bryolo gi cal s tu dies Show tha t the dis tal p ar t
,

of the transverse p ro cess is a rib whi ch has be co me indis ti n gu ishably fu sed to

I
,

the p ro c ess There is one sa c ral ver tebra su pp or ting the hind legs and beyo n d
.

this the long u ros tyle already m e n tio n ed The ver tebrae are yoked toge ther .

by zy gap o p hyses as in Nectu ru s In sin gle ver tebrae from the fro g n o te tha t .

the p os terior en d of the c en t ru m is ro u n ded like a ball w hi le the an terior end is ,

t tog e ther in a ball an d so cke t m an n er - -

.

fo rm ed by the u n ion of the b asido s al an d r s

that the i te do rs als a r e fu s ed to either the n r

f rog the in terdors l s a e f sed to the po s te io r a r u r

e d fo rmin g the ball sha p ed p ro j ec tion

n -
I n terve t al s a r e l a ck i g Ve tebra e o f this type
. n r n . r

a e call ed n otocen tr ous ( s ee F ig 2 D p

r . 2 , .
7O LAB ORA O R T Y MANUAL F OR VE R TE B RAT E ANA OM T Y

G . VE RTE B RA L COL U MN OF RE P TI LE S

The ver tebral colu m n of r ep til es rea ches a d egree of d ifferen tia tio n consider
ably in adva n c e of tha t of the Amphib ia and shes .

.I The vert ebral c ol u mn of the all i g ator S tu dy the whole m o u n ted sk ele to n .

of the alliga tor Observe tha t the colu mn is differen tia ted in to ve region s :
.

the cervical or n e ck region mu ch lo n ger than in Amphib ia ; the thor a cic r egio n
,

bearin g lon g ribs ; the l u mb ar regio n wi thou t ribs ; the s acral regio n comp os ed , .

of two fu sed ver tebrae su pp or ti n g the hind lim bs ; an d the cau dal r egion The
, .

'

tail ver tebrae are the m os t p ri mi tive in form an d will be s tu di ed r s t E a ch .

c onsis t s of a larg e c en t ru m a ne u ral ar ch wi th a hi gh neu ral s p ine p ro mi n en t

, ,

transverse p ro c ess es di re c ted s trai gh t ou t fro m the c en tru m a n d a haem al ar c h , ,

mis sin g on the rs t c au dal ver tebra The haemal ar ches are rela tively s mall an d
.

si tu a ted a t the p os terior en d of the c en tr u m They are of ten referred to in .

t ex ts as chevron b on es Toward the en d of the tail the variou s pro cesses p roje c t
.

in g fro m the ver tebrae t en d to be co m e red u c ed a n d n ally vanish al to ge ther so ,

tha t the las t v er tebrae c o n sis t of c en tra o n l y The sa cral regio n co n sis ts of two .

ver tebrae each beari n g a s tou t sa cral rib to whi ch the s u ppor ts of the hin d
, ,

limbs are ar ti cu la t ed The sac ral ver tebrae have hi gh n eu ral sp i n es b u t la ck the
.

haem al ar ches The lu mbar ver tebrae an terior to the sa cr u m are ve in

.
, ,

n u m ber p ossess h
,
i gh n eu ral sp i n es and broad t ransverse p ro cesses There are .

ten thora ci c ver t ebrae similar in form to the lu m bar ver tebrae b u t beari n g
, ,

long ribs which rea ch the m edian ven tr al line There are nin e cervi cal ver tebrae
,
.
,

of which the two rs t known as the a tl as an d the axis are som ewha t differen t
, ,

from the o ther seven These la tter have s tron g n eu ral ar c hes wi th lon g n eu ral
.

sp i n es an d shor t tra n sverse p ro c esses beari n g ribs M os t of these c ervi cal ribs .

are of p e cu liar form bei n g V shap ed and a tta ched to the cervi cal ver tebrae b y
,
-

the two e n ds of the V Cervi c al thora ci c l umbar an d sa c ral ver tebrae la ck

.
, , ,

haemal ar ches b u t the c ervi c al an d a n terior thora ci c ver tebrae p ossess shor t
,

ven tral p roje c tio n s the hyp apo physes on the cen tra All of the ver tebrae are
, , .

p rov i ded wi t h well develo p ed -

p r e and p o s tz h
yg p p y
a o s es B y m ovin g the .

ver tebra e ap ar t n o te tha t the c en tra are p ro coelou s i e co n c ave in fron t con ,
. .
, ,

vex behi n d .

The rs t two cervi cal ver tebrae the a tlas an d the axis are very rem arkable , ,

a n d deserve f u r ther a t t en tion They are rem arkable in tha t the p ar ts which
.

c o mp os e the p ri m i tive ver tebra are re tai n ed in these two ver t ebr a e alm os t
u n c ha n ged ( F ig 2 2 F ) The a tlas or r s t cervi cal ver tebra is co mp osed of fou r
. .

sep ara te p ie ces which toge ther form a ring The ven tral side of the rin g is
,
.

c o mp osed of a m edia n p ie c e the fu sed b as iven tr al s I t bears a p air of lo n g

,
.

m ovable ribs The sides of the rin g are cu rved bo n es the b asido rs al s The
.
,
.

ri n g is co m p le ted abov e by the n eu ral sp in e The a tlas app ar en tly has no .

c en t ru m b u t the c en t ru m is really p resen t an d has b ec o m e a t t a ched to the an t erior

en d of the se c o n d c er vi c al ver t ebra or axis This c en tr u m is c o mp osed of the .

 T HE E NDO S KE L E TON : VE RTE B RAL COL UMN AND RI B S 7I

fu sed in terv en tral s I t p roje c ts forward in to the ri n g form ed by the a tlas as a

.

p oi n t ed p roje c tio n the od on toid pr ocess The , a t las t herefore p ossesses a ll of .

the p ar t s of a p rimitiv e ver teb r a ex c ep t the i n terdorsal The axis ( also called

the epis tr opheu s ) has a lar g e c e n tru m c o mp osed of th e fu sed in terv en tr al s a n d ,

bears a t its an terior en d the odo n toid p ro cess whi ch as already exp lai n ed is , , ,

really the c en tru m of the a tlas The p lac e of union of the odon toid p ro cess wi th .

the c en tr u m of the axis rep resen ts the redu ced basiven tral of the a xis The axis .

has a s tro n g n eu ral ar ch (b asidorsal s) and broad neu ral spine I t bears a p air .

of ribs whi ch cu riou sly en o u gh have moved forward so tha t they are a tta ched
, , ,

to the odo n toid p ro c ess H en ce it m i gh t be said tha t the a tlas has two p airs of
.

ribs and the axis n on e .

T he ver tebrae of the all igator are c ompo sed in g en eral of the foll owin g a rc al ia T he u .

n eur al ar ch co s is ts o f the fu sed b as id o r s al s ; the c en tru m of the fu sed in terven tr al s ; ba s i

n

ven tral s are p esen t onl y in the atl as the axis an d the cau dal ver teb rae wher e they form the
r , , ,

chevr on b on es ; b u t they ar e p ob abl y rep r esen ted th o ghou t by the in t erv er t eb r al c a til ag es
r r u r

( ab sen t o f c
,
ou r se o n d ied s k e
,
l et o n s ) T he b r
as iv e t ra l s ar e p es en t a s d is t in
. c t b o es the n r n ,

i t er cen t a wedg ed in b etween the v e t eb a e in the primitive r ep til e S phe don which is
n r ,
r r no ,

l ivin g fossil o n a ccou n t ofits p rimitive chara c t eris tics in s ome liz a ds

s om etimes c all ed a , r

an d in man y e ti c t r ep til esx nI n t e do s al s ar e ab se t Ver teb ra e lik e tho s e of the all ig ator
. r r n .

in which the c en tru m is fo rmed o f the i t erv e tr al s are n amed gas tr ocen tr ou s ver teb a e n n r .

G as tro cen trou s ve teb ra e a e cha ac te is tic of ep til es bi ds a d mammal s ( Fig 2 2E p

r r r r r ,
r ,
n .
, .

2. S tu d y of the r ib s The ribs m ay be s tu died on the cervi cal or rs t

.

thora c i c ver t ebra e Observe tha t ea ch rib is a tta ched to the ver tebra by two
.

p r o c esses wh i c h,
are c alled hea ds The u pp er head or tu b er cu l u m is a t t a ched .
, ,

to a shor t blu n t tra n sverse p ro c ess arisi n g fro m the ne u ral sp i n e The lower .

head or capitu l u m is a tta ched to the side of the a n terior end of the c en tru m A
, ,
.

large op en i n g is n a tu rally i n closed by the two heads of the ribs The su cc essive .

op en i n gs form a c an al c alled the ver tebr ar ter ial can al in whi ch blood vessels to
the head are lo c a ted The ribs of the third to seven th cervi cal ver tebrae are
.

shor t an d blu n ted those of the eigh t and nin th cer vical s considerably longer
, ,

a n d those of the rs t ei gh t thora c i c ver tebrae very lo n g and cu rved rea chin g .

to the ve n tral side Observe in the series of the thora ci c ver tebrae tha t the
.

c ap i tu lar h ea d of the rib g rad u ally m oves dorsally u n til it nall y c o m es to be

a tta ched to a li ttle p roje c tion on the tran sverse p ro c ess The l i ttle smoo th .

p la c es on the ver t ebrae t o whi c h the heads of the ribs are ar ti cu la ted a r e

kn own as facets and rib beari n g ver tebrae can always be re cogniz ed by su ch
,
-

fa ce ts .

The thora ci c ribs consis t of three p ar ts The upp er third whi ch has the .

heads is c omp osed of bone and is kn own as the ver teb r al rib The middle thi rd .

is par tly car tilagi n o u s and is called the in ter mediate rib The lower third is .

also p ar tly car tilagi n o u s and is the s tern al rib or costal cartil age The las t two .

thora ci c ribs c o n sis t of ver t ebral ribs onl y .

72 LAB ORA ORT Y MANUAL T
FOR VE R E B RA E ANA OM T T Y

On the ven tral side of the lu mbar region will be no ted a series of riblike bones
n o t a t ta ched to a n y o ther p ar t of the skele ton These are the so ca lled a bdomi.
-

n al rib s They are mem bran e bones and therefore belong to the exoskele ton

.
,

bein g in fa c t hom ologou s wi th the derm al p la tes of the tu r tle s p las tron They
.

are likely to o ccu r in lon g bellied ani m als -

.

Draw a c ervi cal or a n t erior thora ci c ver tebra of the alliga tor wi th all of its
p ar t,
s sho w i n g p ar t i cu larly the rela tion of t he ribs .

T h ver t ebra l ol mn f th t rt le I s t dy of th e c arap a c e of

3. e c u o e u n ou r u .

the tu r tle we already n o te d c er tai n p e cu liari ties of the ver tebral c ol u mn of these
anim als These p e cu l iari ties aris e from the cir cu ms tan ce tha t p ar t of the ver te
.

bral colu mn is fu sed to the ex oskele ton The ver tebral colu mn of the tu r tle .

c onsis ts of c ervi cal ,t ru n k sa c ral and ca u dal re gions

, ,
As u s u al the cau dal .
,

region is the m os t p rimi tive The cau dal ver tebrae have neu ral an d haem al
.

ar ches and tran sverse p ro cesses of whi ch the m os t an terior are ribs as S hown by
, ,

the s u tu re a t their bases The rs t cau dal ver t ebra is fu sed to the sa cru m which
.
,

c o n sis t s of two sa c ral ver t ebrae bearin g sa c ral ribs The rs t cau dal the two .
,

sa cral an d the ten tru nk ver tebrae are all f u sed to the c arapac e As already
,
.

e xp lain ed in c o n n e c tion wi th the ex oskele ton the n eu ral ar ches of these ver ,

t eb r a e are broaden ed to form the ver t ebral p la t es of the c arap a c e and the ribs ,

of the tru nk ver tebrae are si milarly exp an ded to form the cos tal p la tes of the
c arap a c e The rs t an d se cond p airs of ribs are fu sed dis tally The ribs of the
. .

tu r tle p ossess a si n gle head the c ap i tu lar head whi ch ar ti cu la tes a t or near the
, ,

bo u n da ry be tween two s u c cessive c en t ra There are ei gh t cervi c al ver tebra whi ch

.

are devoid of ribs and w hi ch are very ex ibly ar ti cu la ted by ball and so cke t - -

join ts The rs t two are differen tia ted as in the alliga tor in to a tlas an d axis
. .

The odon toid p ro cess whi ch is the c en tru m of the a tlas is the large mass
, ,

a tta ched to the an t erior end of the axis .

H . VE R TE B R A L CO L UMN OF B IR DS

The ver tebral c olu mn of birds like their en tire s tru c tu re is hi ghly sp ecial
, ,

iz ed in rela tion to the i gh t habi t Wi th a p rep ared skel eto n of the bird
.

c hi cken or p i g eon before you no te the followin g p oin ts The ver tebral column
,
.

is divided in to c ervi cal thora ci c lu m bar sa cral an d c au dal regio n s b u t all of

, , , , ,

these ex c ep t the rs t are f u sed to ge ther in order to s tre n g the n the ba ck The .

c ervi c al ver tebrae are n u m ero u s ( s ix teen in the c hi cken thir t een or fo u r t een in
'

t he p i geon) and have very exible ar ti cu la tions birds bein g the only ver tebra tes ,

t ha t can tu rn their heads halfway ro u n d This exibili ty is du e to the shap e .

of the cen tra On isola ted ver tebrae or by sligh tly sep ara tin g so m e of the cer
.

vi cal ver tebrae on the m ou n ted skele ton no te tha t the en ds of the cen tra are
saddle shap ed ; this typ e of c en tru m is heter ocoel ous The rs t an d sec ond .

c ervi cal ver tebrae are the a t las an d the a x is ; the form er is a s m all rin g shap ed -

bon e the la tter bears an odon toid p ro c ess as u su al B ehin d the ax is are the
, .
74 LAB ORATOR Y MANUAL FOR VE R E BRA E ANA OM T T T Y

its p os terior en d has likewise two s u rfa c es whi ch ar ti cu la te wi th the ax is As in .

the case of the alli ga tor , the ve n tral side of the tlas ri n g is the basive n tral ;
a
the si de s an d top the b asido r s a l s ; p ar ts of the t ra n sverse p ro c esses are ribs ;
,

a n d the c en t ru m ( i n terven tral ) is a tta c hed to the ax is Draw the a tlas fro m the .

fron t .

The se con d cervi cal ver tebra the ax is or ep is trop heu s has a very long an d
, ,

broad n eu ral ar ch p rovided wi th a neu ral sp i n e whi ch p roje c ts forward over the
,

a tlas I ts cen tru m bears at its an terior end a p oin ted p roje c tio n the odo n toid
.
,

p ro c ess w hi c h
,
ts in t o the rin g of th e a t las allowi n g the tu rnin g of the head , .

The odon toid p ro cess is in reali ty the c en tru m of the a tlas The axis bears .

la terally the so called tra n sverse p ro cesses b u t o nl y the dorsal p ar t of thi s is the
-
,

real transverse p ro cess the lower half bein g a ru dim e n tary rib This rib as is
, .
,

u sii all y the case is un i ted wi th the ver tebra and the transverse p ro c ess in s u ch
,

a mann er as to leave an op en in g the ver tebrar terial canal The an terior en d , .

of the c en tru m of the axis has two ar ti cu la tin g su rfa ces for the a tlas its p os terior ,

end on e for the nex t ver tebra I ts neu ral ar ch has a p air of pos tz ygap ophyses
,
. .

Draw the axis from the side .

The rem aini n g c ervi c al ver tebrae are more or less si milar They have well .

develop ed neu ral ar ches and spi n es the la tt er in creasin g in hei gh t toward the ,

thora ci c region All have p r e an d p os tz ygap o p hyses an d tran sverse p ro cesses

. .

The la tter are in reali ty in p ar t c om p osed of a rib res u l tin g as before in the ,

form a tion of the ver tebrar terial canal Draw a cervi cal ver tebra from the .

fron t .

2 The thora ci c vert ebra e

.
These ver tebrae are rec o gn iz able thro u gh the
.

fa c t tha t they bear lo n g ribs mos t of whi ch ex ten d to the ven tral side In the
,
.

isola ted ver tebrae the s moo th cos tal facets where the ribs were a tta ched always
serve to iden tify them as thora ci c There are thir teen thora ci c ver tebrae in .

the ca t generally twelve in the rabbi t vario u s n u m bers in o ther m amm a l s (m an

, ,

has twelve) The m ajori ty of the thora ci c ver tebrae have very tall neu ral sp ines
.
,

dire cted ca u dad shor t c en tra s m all pr e an d p os tzygap op hyses an d shor t s tou t
, , , ,

tra n sverse p ro cesses hea rin g a t their e n ds a fa c e t for ar ti cu la tion wi th the

,

upp er head of the rib The cen tru m bears a t its an terior an d p os terior en ds
.

half fa ce ts for the lower head of the rib

-
The las t thora ci c ver tebrae differ .

som ewha t fro m the o thers They are lo n ger and s tou ter wi th shor t n e u ral
.
,

sp in es m ore p romi n en t z ygap op hyses an d mu ch redu ced tran sverse p ro cess es

, ,
.

They bear b u t on e fa ce t for the ribs a sm all dep ressio n near the an terior ,

en d of the c e n t ru m B elow the po s tz ygap 0 p hysis in these las t thora ci c ver te

.

bra e is a pos t eriorly dire ct ed p ro c ess the access or y pr ocess co n spi cu ou s in , ,

the cat .

No te tha t the an t erior a n d p o st erio r en ds o f ver t eb ra e can be rec ognized as foll ows : the smoo th
f
a rticu l a tin g su r a c es o r facets o f the p r ezyg a h
p p yses
o face upward or fo rward whil e the facets o f the

p O S tz
y a h
g p p yses
o face down ward o r b ackward .
 T HE E NDOS K E L E TON : VE RTE B RAL COL UM N AND RI B S 75

3 . The rib s
The ribs of m amm als c on sis t of a dorsal bon y p ar t the ver
.
-
, ,

teb r al rib an d a ve n tral c ar tila gi n ou s p or tion the s tern al rib or cos tal car ti
, ,

la ge The ver tebral rib is ar ti cu la ted to the ver tebra by two heads an upp er
.
,
.
tu b er cu l u m whose fa ce t e n ga ges the cos tal fa c e t on the u n der s u rfa ce of the
,

tran sverse p ro c ess ; and the lower capitu lu m a tta ched be twee n two c e n tra to ,

demifacets i e half a fa c e t on the p os terior en d of o n e c en tru m and half a fa ce t

,
. .
,

on the an terior en d of the su cceedin g c en tru m The tu ber cu lu m is di mi n ished .

on the more p os terior ribs an d wan tin g on the las t three ribs whi ch are p rovided
wi th cap i tu lar heads o n ly The n arrow p ar t of a rib be tween tu ber cu lum and
.

c ap i tu l u m is c all ed the n eck; the p oi n t of grea t es t cu rva tu re j u s t a li ttle beyo n d ,

the tu ber cu lu m the an gl e; the re m ai n der of the rib the b ody

,
Those ribs whi ch , .

rea ch to the m edian ven tral line an d are indep enden tly a tta ched to the m edian
ven tral s tru c tu re ( the breas tbone) are known as tr u e ribs ; those whi ch are n o t
so a tta ched to the breas tbon e are c alled fal s e ribs ; an d those false ribs whose
ven tral en ds are free are oatin g ribs The cat has n in e tru e ribs an d fo u r .

false ribs of which the las t o n e is oa ti n g ; the rabbi t has seven t ru e ribs an d
,

ve false ribs of whi ch the las t three are oa tin g ; in man there are seven t ru e
,

ribs and ve false ribs the las t two oa ting The ribs of m ammals are in ter
,
.

m u s cu lar ribs D raw o n e o f the t yp i cal thora ci c ver tebrae wi th its ribs fro m
.
,

the fro n t .

T h e l u m b ar ver t e b rae
There are seve n l u m bar ver t ebrae in th e c a t
A
4 . .

and rabbi t They are large an d s to u t ver tebrae wi th p rominen t neu ral spin es
.
,

c o n sp i cu o u s z y ga p op hyses a n d lo n g t ra n sverse p ro c esses dire c t ed c raniad

,
.

B elow the p os tz ygap 0 physis is a p oin ted p roje c tion dire c ted cau dall y called , ,

,
the a ccessory p ro c ess Draw a l u mbar ver t ebra from the side
. .

5 Th sa r m The sa r m is o m osed of a variable n u m ber of ver t ebrae

. e c u c u c .
p
fu sed together for ar ti cu la tion wi th the hin d limb s There are three sa cral .

ver tebrae in the ca t gen erally fo u r in the rabbi t (of which only the rs t two really
,

c on t rib u te to the a t t a c h m e n t ) ve in m an The bou ndaries be tween the fu sed

,
.

sa cral ver tebrae are readily m ade ou t by m ean s of the op enings be tween them
t hro u gh whi ch the S p i n al n erv e s p ass ou t an d by m eans of the number of n eu ral ,

sp in es z ygap op hyses etc The rs t sa cral ver tebra ass u mes the grea ter p ar t
, ,
.

of the task of tr ansmi ttin g the s u pp or t of the hind limbs to the ver tebral colu mn ;
for this p u rp ose it has larg e la teral exp an sion s bearin g ar ti cu lar su rfa ces for the
in ser tion of the bon y s t ru c tu re whi ch supp or ts the hin d limb These la teral .

exp ansion s c onsis t in p ar t of t ran sverse p ro cesses an d in p ar t of sa cral ribs ,

in dis tingu ishably fu sed to the ver t ebra .

6 The c au d al v e rt ebrae
.
The c au dal ver tebrae are variable in number
.

in m amm als Neu ral ar ches tran sverse p ro cess es an d z ygap ophyses dimi n ish
.
, ,

T his u s e o f the t erm s t ru e an d fal se shou ld n ot be co n u s ed f w ith the u sag e p r evio u sl y g iven in
which the ex p r ess io n t ru e rib f
re ers to in term u scul a r rib s an d fal se rib to su b p erit o n eal rib s . I n this

co mp a ra t ive s en s e all m amm alian rib s are t rue rib s .
76 LAB ORA OR T Y MANUAL F OR VE R TE B RATE ANATOMY

a dally the las t ver t ebrae consis tin g of c en tra only Very sm all haemal
c u ,
.

ar ches ( chevron bo n es) are p resen t in the tail ver teb rae of the cat b u t are missing ,

on p repared skele ton s Man has three to ve cau dal ver tebrae fu sed in to a .

si n gle pie ce the u ros tyl e or coccyx,

.

G e eral re m ar k on th whole ol m The ends of the cen tra of mos t

7 . n s e c u n .

in amm alian ver tebrae as can be seen by i n s p e c tion of isola ted ver tebrae are

, ,

m ore or less a t Su ch c en tra are called a mphipl atyan

. B e tween the ends of .

the ver tebrae are fo u n d in life car tila gi n o u s disks the in terver teb r al car til ages , .

On the mo u n t ed skele to n observe an op enin g on ea ch sid e be tw ee n s u ccessive

c en tra Throu gh these op en i n gs the in terver tebr al for amin a the sp inal nerves
.
, ,

p ass o u t fro m th e s p i n al c ord The s p inal c ord o c cu p i es the c o n tin u o u s canal .

formed by the n eu ral ar ches H aemal ar ches (b asiven tral s ) are absen t ex cep t .

in the a tlas an d a xis and in the tails of so m e m amm als where they form chevro n ,

bo n es The ver tebrae of m amm als are gas tro cen trou s
. .

J . S UMMAR Y OF TH E VE R TE B R AL COL U MN AN D RI B S

1 . T he v er te rae b a r is e at the in t er se c tion of the myo s ep t a with the mes en chyme

s u rr ou n d in g the n o to cho r d an d n eu r al tu be . E ac h ver tebra is p
by the u io of the r o du c ed n n

p o s te ior hal ves

r of the two s cl er o to mes o f o e s eg m e t with the an terio r hal ves o f the t wo
n n

s c l ero to mes o f the s u cc eedin g s eg men t . Owin g to this man n er of o rig i the ve teb a e al t e n r r r

n a te h the myo tomes

wit .

2 . T he ve teb ae an d rib s ar e r s t fo rmed in ca r til ag e p rodu ced by the ac tivity of the

r r ,

m esen chym e I n the el as mo b r an ch shes they r emain p erman e tly in the ca til ag e s tage
. n r .

I n mo t o ther ve t ebr a tes they oss ify du ri g d evel o p men t

s r They b el on g con sequ en tly to
n .
, ,

t he ca t eg o ry o f c til ag e b o n es ar .

3 .E ac h v e t b a b e g i s a s
r e f ou r p
r a ir s o f c a r t
n il ag e s o r a r cu a lia su rr o u n di g the n o to chord : n

d o r s all y a a t e io r p a ir o f b a s ido rsal s an d a po s t erior p air of in ter do sal s ; ven tr ally a

n n r r n

a t er io p air o f b a siven t al s an d a po s te io r p air o f in terv en t a l s

n r r r r .

4 I n p imitive ver t eb ra e ( o c cu r rin g in s o m e s h e ti c t Amphibia an d r ep til es ) thes e

. r ,
x n

p ieces remain mo e or l es s s ep a ate in the a du l t In the maj o ity o f ver teb ra tes however
r r . r , ,

so me o f them ar e l o s t a d the r em ain de fu se in to a s in g l e s tru c tu r e

n r .

5 .T h e p a r t s o f a s i g l e v e t eb a th u s f
n o m e d by t h
r e fu s io n o
r f o rig ally sep ar a te p iec es
in r

ar e : the c e t ru m o r m a i n body o f the ver teb a which in cl oses the n o to chord ; a do sally
n r ,
r

d i ec ted a ch the n e al a ch in cl o s in g the s p in al co r d ; a d a v e tr ally dir ec ted a ch the

r r ,
ur r ,
n n r ,

haemal arch i cl o si g bl ood vess el s The n eu ral a ch co sis ts of the fu sed b asidorsal s the
,
n n . r n ,

haemal a ch o f the fu s ed b asiven t al s ; the ce tru m is o f d iffe en t ig in i differen t g ou p s

r ,
r n r or n r .

6 T he c e tru m in th el a s mob ran ch shes is f or m ed withi the S hea th o f the n o to cho rd ;

. n e n

it is call ed a c ho d al c tru m r T he c en t a in n early al l o th e v erteb r a tes are pro du ced by

en . r r

the fu s io o f c er ta in o f the a c al ia an d a e kn o wn a s p ericho rd al o r a ch c en t ra

n r u r I n d ifferen t r .

g r o u p s o f v e t e b a t esr di ffe e t
r a cu a l ia co n t rib u t
r n e t o t h e c e r t r a n .

7 T he en d s o f the cen tr a ar e v ariou sly s hap ed amphico el ou s or co cav e at ea ch en d

. : ,
n

( shes u ro d el es) ; p ocoel ou s con cav e i f ron t con ve b ehin d (Amphibia r ep til es ) ; op is tho
,
r ,
n ,
x ,

co el ou s co v e ,
i f o t co n cav e b ehi d ( Amphibia rep til es) ; hetero coel ou s s a ddl e S h p ed
n x n r n ,
n , ,
a

( b ir d s ) ; a n d a m p h ip l a ty a a t a t e a c h e d ( m a
n mm
,
l s) n a .

8 In . d d it i a to ce t m a d a ches ve t eb ae c o mm o ly b ear p r o j ec tin g p oc es s es

on n ru n r ,
r r n r

o r a po p hys es T he mo t co mmo o f thes e a e the tr a s ver se p o ces s es an d the yg a po phys es

. s n r n r z .
 T HE E NDOS KE L E TON : VE RTE B RAL COL UMN AND RI B S 77

T he fo r mer serv e fo r the ar ticu l a tion of the r ibs an d are p robably n o t ho mol og ou s in d iffer en t
ver teb r at es T he zyg ap o physes yo k e su c ces s ive ver teb ra e to g ether
. .

9 . T he haemal ar chespp ea r o b ecome r edu c ed in ver teb r a tes

ten d to dis a r They p e sis t . r

chiey in the tail region an d may al so con tribu te to the a is an d the atl as x ,
.

10 T he v er teb ral co l u mn in shes is divid ed i to t ru k an d tail r egion s on l y

. n n In .

A mphibia a very sho rt cervical r eg ion an d a sacr al r eg ion are a dded I n r ep til es bi ds an d .
,
r ,

mamm al s the c erv ical r eg ion is l o g er an d the t run k reg ion is d iv id ed in to an an terio r tho ra cic
,
n ,

reg ion b earin g l o g r ib s an d a p o s terio r l u mb ar reg ion with r edu ced o r n o rib s
,
n , T he differ .

e ces b etween the v er teb r ae o f the d if

n feren t r eg ion s b eco me mo r e an d mor e mark ed the higher
o e as c en ds in the ver t eb ra te s cal e
n .

1 1 Orig i lly each ver teb ra was p rovided with a p air of r ibs b u t in the higher ver te
. na ,

b ra tes thes e ar e r ed c ed o r ab se t exc ep t in the tru n k o r tho r acic r eg ion s

u n R edu ced ribs .

ar e g en erally p es en t on the cerv ic al v er teb ra e a d al ways on the s acral v e teb ra e

r n r .

12 R ib s ar e o f two k in ds : thos e tha t a ris e a t the in ter sec tion o f the myo s ep ta with the
.

hori on tal sk el etog en ou s sep tu m kn own as tru e or in termu scu l ar ribs ; a d tho se that aris e
z ,
n

a t the i t er s ec t io
n o f myo sep ta with the ven tral s k el etog en ou s s ep tu m o r its d erivatives
n ,

k n own as fal se o r su bp eriton eal ribs The l atter are characteris tic of tel eos ts the former o f
.
,

al l o ther v er t ebra tes S o me shes po ss es s bo th k i ds o f rib s s imul tan eou sly an d may al so
. n

devel op ad ditio al r ib s at o ther l ev el s o f the myos ep ta

n .

I3 R ib s typ ically a r tic l a t e w ith the v e teb a e by two heads the s p ac e between the
. u r r ,

heads fo rmin g a ver tebrarterial can al fo r the p as sag e of bl o od vess els .

VII . TH E E ND O S K E LE T O N : T HE C O M P AR ATIVE ANAT O M Y OF
T HE G IR D LE S ,
T H E S T E R NUM , AND T HE P AI RE D
AP P E ND AG E S

A . G E NE R AL CON S IDE RAT I ON S

De ni tion s The
h ped o r arch shap d p or tion s o f the en d r
1 gir dl es a re c es c en t- s a -
. . e o

s k el eton which fun c tion fo the s u pp o t o f the p ai ed a pp en d ag es

r T he c e te o f the ar ch is r r . n r

d i e c ted v en t all y the p o in ts d o ally

r r ,
T he gi dl es a e com po s ed o f c a til ag e in the l owe
rs . r r r r

fo rm s ; this is p a tl y o r comp l etel y o ss ied in the higher o n es T he pect r al gi dl e su ppor ts the

r . o r

a t erio r a pp e dag es ; the p l vic gi dl e the po s terio r a pp e dag es

n n e r n .

T he s t r n m o r br e tb n e is a el o g a ted s tru c tu e lyin g in the median ve t al lin e o f the

e u as o n n r n r

a terior p art o fthe t run k r eg ion

n I t is comm onl y compo sed ofa chain o fca til ag s o r b on es o r
. r e ,

bo th T he ribs an d .

l l a f ld atera n o s
th e p ec to al gi dl e u s r r

n all y a t icul a te with r

the s t e um S u ch rn .

a rran g em en ts s tr e g n

t h e n t he a n t e i o r r

p ar t of the tru nk in
r el a tio n to t he a ir

b hin g hab it an d
r ea t

the p r es e c e o f l u n g s
n .

T he pair ed appen d
a ges c on s is t o i n s in

the shes an d l imbs in

a ll of the l an d ve r te

Diagr am s
b ra tes Th6 an t en o r
FI G . 23 . t o ill u s tra t e the theo ry medi an
o f the o ri g in o f the

p p or p ec to fal app el l d '

an d air ed n s thr o u gh the er s is t en c e o f cer tain r egio n s o f o rigin all y m m

ag es a r t l cu l a t e W l th
tin u o u s med ia n an d l a ter al n folds A , earl y s ta g e sho win g the median
Femo ral
.
_

the r d l e,
d o r s al n fol d an d the t wo l a t eral n fol d s u n itin g a t the an u s . B , l a t er g
s ta g e ill u s tra tin g p er sis ten c e o f c er ta in r egio n s o fthe n fol d s as the m edian an d thei Su r ppo rt is
an d p a ir ed n s , an d disa pp ea r an c e of the main der
re o f t he n fold s a s
,
t ran m i t t e d to the
s

in dica ted b y d o tt ed lin es ( From W ilder .

s His tor y of the Hu ma n B ody, bo dy by m ean s of this
co u r tesy o f Hen ry Hol t an d Com an y ) p .

gir dl e simil arl y the

,

po s terio r o r p e l v i c
a pp en d ag es a e ar ticu l a ted to the p el vic girdl e a n d t ran s mit their s u ppo t throu gh this gir dl e
r r .

As the s u pp o rt o f the po s te io r appen dag es is g en e all y the mo re impo ta t the p el vic

r r r n ,

g i dl e is co m m on ly s t o g er an d m o e m ass iv e tha the p ec to ral gir dl e

r r n This is p articul a ly
r n . r

t e o f b ip ed v e teb a tes
ru T he a ppe d ages po s s ess an in tern al s k el eto
r r . n n.

2 Th o igin o f th p air d app

. e r d g s d the gir d l e s
e e
T he o rigin o f thes e s tru ctur es
en a e an .

i ver teb a tes is ob s c e

n r Acc o di g t o the mos t p ob abl e theo ry t he fol d theo ry the
ur . r n r ,
n- ,

an c es t al v er t eb a t e po s s es s ed a p air o fc on t in u ou s f ol d s o e ru n n in g al o g each s id e o fthe t ru k

r r ,
n n n .

These fu sed b hi d the a u s to a s in gl e median which e te ded a ou d the tail a d al o g

e n n n x n r n n n

the m ed ian d o s al lin e ( s e Fig 2 3 A)r Thi hypo thetical co ditio r sembl es tha t ac tu lly
e . . s n n e a

o cc rr i g in A mphi xu s with its p air ed v en t al m e tap l eu ral f ol ds an d m edian cau d l an d d o sa l

u n o ,
r a r

78
 THE E ND O S KE L E TON : G IR DL E S , T HE S E R N T UM ,
AND AP PE NDA E S G 79

n . These n f o l ds wer e su pp orted by car til agin ou s n ra ys . T he p aired p es e t n s of r n

shes are su ppo s ed to have arisen throu gh the p er s is ten ce of cer tain n r eg ion s o f the p aired

fol ds an d the median u n p aired n s throu gh the p er s is ten ce of p ar ticu l ar region s ofthe median
,

n fold T he r emain in g p or tion s of the n fol d s have van ish

. ed ( s ee F ig 2 3 B ) This theory . .

is su pp o r ted b y the fac ts that the u n p air ed an d p air ed n s o f shes ar e iden tical in s tru c tu re that ,

in the you n g s tag es o f so m e fo rms con tin u ou s p air ed fol ds ar e p res en t f rom which the p aired n s
a ris e an d tha t in the ex tin ct shar k Cl ad os el ach
, e ( s ee K F ig 1 2 1 p 1 1 5) the arran g emen t o f , . , .

the n rays in the p air ed n s is su ch as to sugg es t s tro n g ly the o r ig in o f the p air ed ns fr om

con tin u ou s fo l d s .

T he girdl es a ros e l a ter than the p air ed n s s in ce ex tin ct fo rm s po ss es s the l a tter withou t ,

the fo rmer T he n s a s al r eady s ta ted ar e su pp o r ted by car til agin ou s n rays which may
.
,

occu r in several rows . I t is su g g es ted t ha t the m os t medial an d an ter io r o f t hese n ra ys

of the n s fo rm median pl ates or bar s of ca r til ag e Su ch
fu sed a c r oss the mid ven t ral l in e to .

pl a tes or bars r epr esen t pr imitive girdl es L a ter ossi ca tion o ccurs in the car til age r esul t .
,

in g in the bo n y gir dl es compo s ed of s ever al b on es as fou n d in the higher ver t eb r at es , .

R ead W .

I n the evol u tion of sh in to Amphibia the p air ed n s tran s fo rmed in to limb s S tag es .

in this tran s fo rm ation ar e very imp erfectly kn own an d d icu l t to con ceive I t is su ppo sed .

tha t the n r ays o f the p air ed n s wer e g radu all y r edu ced in n umb er either by l o s s or fu s ion .

A few enl arg ed n ra ys a

b pe sisted Su ch edu c
r ran g ed s i mil arly to the bo n es of the lim s r . r

t ion a d enl a rg em en t o f n rays real ly o ccu rs in the cro ss o p terygia

n g oids l ivin g a d
an n n

e tin c t
x a d the arran g emen t of the n r ays in thes e fo rms is s tro gl y su gg es tive o f the
,
n n

ar an g emen t o f the b o es in the verteb r ate limb

r ( S ee fu r ther N p 1 76 Fig 1 0 2 a d
n .
, , .
, .
,
n

p 1 78 Fig
.
,
.

R ead fu ther on the comp ara tive an atomy of the s te n u m gir dl es a nd p air ed app e d
r r , ,
n

ages ln K W , ,
an d Wd .

B . THE P E L VI C G I R DL E AND THE P O S TE RI OR P AIR E D AP P E NDA G ES

1 The par ts of a typical g ir d l e an d hin d 1imb B efo r e b e in nin the s tu d y of the

. .
g g
co mp ara tive an a to my o f thes e p ar ts it may b e well to d es cri e the g en eral ized g irdl e an d b
l imb The bon y p el v ic g ir dl e
. aris es from the oss i ca tion o f a car til agin ou s ar c o r l ate h p .

Th r ee c en t er s o f o ss i ca tion app ear in each hal f , r esu l tin g in the p r o du c tion o f t hree p a ir s o f

ca r til ag e b Of on es .each s id e a d ors al thes e ther e ar e a te io on

bon e the
il iu m an n r r

ven t al b o e the pu bis an d a p o s terio ven t al bo e the ischiu m ( Fig

r n T he two r r n .

p b es a d ischia commo ly m et in the median ve t al li e T he ilia a e a tic l ated to th

u n n e n r n . r r u e

sacr al ib s r .

T he hi d limb typ ic lly co s is ts o f th ee eg io s : a p o ima l eg me t the thigh con tai in g

n a n r r n r x s n , ,
n

a s i g l e l o g b o e the femu r ; a m iddl e s eg m e t the s ha k co tai i g two l o g bo es : a p e

n n n ,
n ,
n ,
n n n n n r

a i l tibi a a d a p o s ta ia l bu l a ; a d a d i tal s eg me t thef ot o p s i cl u di g the a kl e sol e

x a n x n s n ,
o r e ,
n n n , ,

a d to es
n c o t ai i g a n u mb er o f mall b o es
n T he g en erali ed an kl e is comp os ed f
n n s n z o

.
,

n i e o r ten b o es i
n thr ee o ws a p o ximal o w o f th ee bo n es
n n a p r ea i l tib ial e a m ed ial
r : r r r x a ,

i t medi m a d a p o s t i l bu l
n er u ; a middl r ow o f on e o r two ce tr al es ; a d a d is tal ow o f
,
n ax a ar e e n n r

ve tar s l s The s ol e o f the foo t i c omp o s ed of ve bon es the metata rs als

a e . an g ed in a t an ss , ,
arr r

ve se r ow
r T he to es o d ig its co ta i a s e ies of s mall bon es the phal a ges of which in
. r n n r ,
n ,

p imitive feet the e a e two to the rs t dig it an d three to a ll o f the o the s Figu re 4 ill s
r r r r . 2 u

t a tes the bon es o f the typ ical p el vic g i dl e a d hin d limb

r r n .

The p e lvi c g ir dl e
2. an d p e l vi c n of ela s m obran c hs
,
F o r this p u rp ose
s tu dy p reserved sk ele to n s of the d o g s h or o th er elasm obra n ch sh . The p elvic
 8O L AB ORA OR T Y MANUAL F OR VE R E BRA E ANA OM T T T Y

g irdle is t h e s trai g h t or sli g h t ly c u rved bar of c ar tila g e p la c ed a c ross t he ven t ral

sid e a t the en d of the t ru nk region be tween the an terior ends of the p elvi c n s .

.
This b ar is call ed t he ischiopu b ic bar an d r ep r es en ts a p rimi tive u n o ssi ed p elvi c
g irdle I n m a n y. elas m obra n c hs ( i n c l u di n g the d o g sh) t he tw o ends of the bar

b ear sli gh t p ro c esses p rojec tin g ou tward ; these are the iliac pr oces s es .

At ta c hed to the p elvi c gir dl e ar ti c u la t in g t o it a t the base of the ilia c p r o

,

c esses are the p elvi c n s

,
The skel eton of the basal p ar t of the ns whi c h is
.
,

imb edd ed in the body wall c onsis ts of a n u m

ber of c ar tilaginou s p iec es the car til agin ou s n ,

r ays ; the s kele ton of the free ex t ernal p ar t of t he

n is c omp osed of a nu m b er of slen der p arall el

der ma l n r ays As t he la tte r are p ar t of the .

ex oskel eton th ey wil l n o t be c o n sidered fu r ther , .

pubi s
The c ar tila gi n ou s n rays are arrang ed in t wo
series a m e dial series c o n sis tin g of o n e or tw o , ,

( in so m e for m s t hree or ve ) m u c h e n lar g ed c ar ti

la g es a n d a la t eral or o u ter series c omp osed of
, ,

a n u mber of sm all c ar tila ges disp osed in o n e or

m ore rows The inner series of n rays are c all ed .

b a s a lia or b as al s the o u ter series r adia lia or , ,

r adial s There is u su ally p resen t a single basal .

,

the metapter ygiu m an elo n ga t ed cu rved ca r tila ge ,

form ing the whole m edial border of the n b u t ,

FI G 4 Di g m f th b
. 2 in som e form s there is an ad di tional basal the
a ra 0 e o n es
,
f th typi l p l i gi dl
e d hi d
ca e v c r e an n
o
m a ed h an er or end of
o

h
o

p op te r yg i u S i t u t a t t e t i t e
m b (F m P k
n . d H w u
ro ar
r
er an as e
,
s
,

T tb k f Z l gy
ex oo o t y f th
o oo o m e t,
a p t ery
c ur esg i u m o
The radials
e
u s u ally c onsis t of a .

M mill a Co mp y)
ac n anrow of rod shap ed c ar tila ges their lo n g a x es a t -
,

ri gh t a n gles t o the a xis of the m e tap terygi um .

I n male s p e cim ens the p os terior radial is g rea tly e nl arg ed to form the c ar tila g e
of the c lasp er The b asal s have p robably arisen t hrou gh the fu sion of a number
.

of sm aller c ar tila ges .

D raw the p elvi c girdle and n of an elasm obran c h .

3 The p elvi c gir d le an d hin d l imb o f u ro d ele s
. S tu dy these p ar t s in dried .

or p referably p r es erved sp e cim en s of Nectu ru s or Cryptobr an chu s The u rodele .

g irdle is in a c o n di tion in t erm edia t e be t ween t h e c ar t ila g ino u s g irdle o f e l as m o

bran chs and the bo n y girdle of rep tiles I t is si tu a ted on the ven tral side a t the .

end of the t ru nk b etween the hin d limb s I ts ven tra l p ort ion has the form of a .

a t tened p la t e whi c h m ay b e design a ted the ischiopu b ic pl ate or pel vic pl ate The
, .

an terior p ar t of thi s p la te is c ar tila gi n o u s a n d is called the pu bic car til age I n the .

p os t erior p ar t of the p la t e a p air of Op aq u e b ony ) rou n ded areas m ay be ,

seen E a ch of these area s is a bone the ischiu m whic h as already exp lained is
.
, , , ,

the p os t erior ven tral bone of typ i ca l girdles The is c hia are p rodu c ed b y o ssi .

cat on i ih t he l
 82 LAB ORA OR T Y MANUAL F OR VE R E B RA E ANA OMT T T Y

segm ent is direc t ed ven trally formin g an a n gle wi th thigh a n d foo t a n d there
, ,

has also been som e sligh t torsion S ee fu r th er o n th ese p oin ts in S e c tion I I I

.
.

The thigh c on sis ts of o n e large bo n e the femu r wh i c h ts in to the a c e tab u lu m

, ,

by a p rom i n en t k n ob the head of the femu r The shank is c omp osed of tib ia an d
, .

bu l a the la tter bei n g the sm all er The ankle is m ade u p of ve ( or in som e

, .

tu r tles s ix) bo n es . At the bases of the tibia a n d b u la is a large tra n sversely ,

elo n ga ted bo n e whi ch is in reali ty c omp osed of fou r bo n es fu s ed ( tib iale

, ,

in ter mediu m cen tr a l e b u l a r e)

, ,
I n som e tu r tles the b u l ar e a t the base of the
.
,

b u la is sep ara t e Dis tal to this c om p o u n d bon e is a row of fo u r bon es the

, .
,

fou r tars al es n umbered from the p rea xial ( tibial) side to the p os ta xial (b u lar)
,

side The app aren t fo u r th tarsale the larges t of the fo u r is r eally the fu sed
.
, ,

fou r th and ft h tarsales D is tal to the tarsales are the ve metatars al s and
.
,

beyon d these the digi ts com p osed of bony join ts or phal an ges term i n a ti n g in
, ,

horny claws The n u mber of p halang es in the di gi ts is tha t regarded as p rim i

.

tive for ver tebra t es n am ely two p halan ges to the rs t di gi t a n d three to
, , ,

all of the o thers ( see F ig I t is wor th y of n o te tha t in the tu r tles a n d

.
,

rep tiles in ge n eral the m ovem en t of the foo t u p o n the l eg o c cu rs be tween

,

the two r ows of tarsal bo n es i e it is an in tr a ta r s a l joi n t

,
. .
, F i gu res of the tarsal .

bon es of tu r tles are given in W d K and R , , .

T h e l vi i r d l d h in d l i m b f b i d These s t r t res like t h

5 . e p c g e an o r s u c u e .
,

rem ainder of the skele ton are highly mo died in birds al thou gh c o n sis ti n g of
, ,

the sam e p ar ts as in o ther ver t ebra t es E xamine isola t ed ba ckbo n es w i th the

.

p elvi c g irdles a t t a c hed or s,

tu dy th e whole m o u n t ed s p e c im e n s The p elvi c .

g irdle c o n sis t s of t hree p airs of bones as in re p t iles i e il iu m i s c h iu m a n d p u b is ,

. .
, , ,
.

Al l three are fu sed o n ea ch side to form a con tin u o u s broad bo n e the in n omin a te ,

b on e . The ili u m is the larges t and mos t dorsal p ar t of the in nom in a te bo n e .

I t forms an elo n ga t ed thi n p la t e c o n c ave in fron t c onvex behind ex ten din g fro m
, , ,

t he las t thora c i c ver tebra to the t ail re gion I t is fu sed alo n g its en tire len gth.

wi th the syn sa c ru m the bou n dar y be tween iliu m an d syn sa c rum g en era lly
,

bei n g marked by a su tu re (I n the bird emb ryo the ilium is ar ti cu la t ed to

.

o n ly two ver tebrae w hi ch are the t ru e sa c ral ver tebrae ) The side of ea ch .

in n om in a t e bo n e is comp os ed of the is chiu m the o n ly bou ndary m ark be tw een ,

thi s and the ili u m b ein g a larg e oval Op e n in g the il iois chiac for amen The pu b is
,
.

is the lon g slen der bone alo n g the ven tral border of the is chium fro m whi ch it is ,

separa t ed by a m ore or l ess dis tin c t su tu re and the sli tl ike ob tu r a tor for a men ,

whi c h m ay be divided in to two or m ore op en i n gs I liu m is chiu m an d pu bis .

, ,

c o n trib u t e t o the form a tion of the a c e tab u l um The an terior en d of the pu bis .

is si tu a ted an t erior to the a c e tab u lu m the no rm al p osi tion of the p u bis b u t , ,

du ri n g developm e n t the p u bis tu rn s p os teriorly an d c om es to p rojec t beyond

the p os t erior end of the is chi u m I n the e mbryo n i c develop m en t of birds
.
,

the three c omp o n e n ts of the i n n o mi n a t e bo n e ili u m is chi u m an d p u bis ori g , , , ,

in ate sep ara t ely . Observe tha t n either p u bi c nor is chial sy mp hyses are p resen t
. ,
 T HE E NDO S KE L E TON : G I RDL E S , T HE S E RN T UM , AND AP P E NDAG E S 83

b u t t he two in n omin a t e bo n es are widely sep ara t ed ventra lly This is p robably .

asso cia t ed wi th the habi t of laying large eggs .

I

The hi n d lim b o ffers several p ecu liari ties The femu r has a lar ge head .

ttin g in to the a ce tab u lum an d a p rom inen t p roj ec tion la teral to the head ,

ca lled the grea t tr ochan ter The dis tal end of the femu r is s hap ed like a pulley
.
,

c o n sis t in g of a c en t ral d e r e ssion wi th c u rv ed rid es the con d l eso n ei th e r

p g y
side Over the joi n t be tween thi gh and shank is an ext ra sm all bone the patell a or
.
,

kn eecap n o t fo u n d in the lower ver tebra t es

, The pa tella is a s es amoid bo n e tha t
.
,

is a bo n e develop ed in a t endon S u ch sesamoid bo n es are q ui te common

, .

in the lim bs of higher ver t ebra tes The shank is composed of two bones a
.
,

m edial larg e o n e and a la t eral shor t ru dim en tary bone The large bo n e is the .

tibiotar s u s I t c onsis ts of the tibia fu sed a t its dis tal end wi th the p ro ximal
.

tarsal bones The p ro xim al end of the tibio tarsu s has two con dyles for ar ti cu la
.

tio n wi th the c ondyles of the fem u r and bears in fro n t two divergi n g eleva tions
,

or cres ts for mu s c le a tta chm en t s The small bone of the shank is the bu l a
.
,

whose dis tal p or tion is a trophied The dis tal end of the tibio tarsu s has a .

p u lley l ik
-
e s u rfa c e for ar ti cu la tion wi th the su cceeding bone the raised ar ti cu lar ,

su rfa c es bein g n am ed malleol i B eyo n d the tibio tarsu s is a lo n g s to u t bo n e

.
,

the ta rs ometa ta r s u s evid en tly fo rm ed by the fu sion of three bones as shown

, ,

by the three ridg es o n its dis tal end The three fu sed bones are the m e ta .

tarsals ( se c o n d third and fo u rth) ; in addi tion the tarsom e ta tars u s in cl u des
, ,

in its p ro xim al p or tion the dis tal ankle bo n es I t will thu s be seen tha t the an kle .

bo n es do n o t exis t separa tely in ad u l t birds b u t the p roximal a n kle bo n es are ,

fu sed to the lower end of the tibia while the dis t al on es are fu sed to the upper
,

ends of the m e ta tarsals The a n kle joint is therefore as in rep tiles an in tr atarsal
.
, , ,

join t The three m eta tarsals which are fu sed to form the tarsom e ta tars u s are
.

the se cond t hird a n d fo u r th b u t a rem n a n t of the rs t m e ta tarsal is p rese n t as

, , ,

a small p roje c tio n on the m edi al side of the dis tal end of this co mp o u nd bo n e .

E a ch m e ta tarsal ar ti cu la tes wi th its resp ec tive digi t co m p osed of p hala n ge s and ,

t erm in a tin g in c laws F if th m e ta tarsal a n d di gi t are q u i te wa n ti n g

. The gai t .

of birds is di gi tigrade .

D raw the p elvi c g irdle a n d hind lim b from the S ide .

6 The p elvi c g ir d l e an d hi n d l i mb of mammal s

.
The p elvi c girdle of .

mamm als is v ery sim ilar to tha t of rep tiles in shap e and sim ilar to tha t of birds ,

in tha t the p elvi c bo n es are fu sed in to in n omin ate bones The girdle co n sis ts .

of the u su al three p airs of bones pu bis ischiu m and iliu m the three of ea ch
, , , ,

S ide bein g indis ti n gu ishably fu sed in to an in n omin a te b on e or hip b one The .

iliu m is the m os t dorsal m os t an terior a n d larges t of the three com p onen ts of the
, ,

in n omina t e bo n e I t ar ti cu la t es wi th the sa c ral ver tebrae an d termin a tes an te

.

rio r l y and dorsally in a c u rved bord er known as the cr es t of the ili u m ,

The ili um .

ex ten ds as far p os teriorly as the a c e tab u lu m ; the dorsal p ar t of the gir dl e p os terior
t o the a c etab u l u m is the is chiu m whi ch is c o n ti n u o u s wi th the ili um The .
 84 L AB ORA OR T Y MANUAL F OR VE R E B RA E ANA OMT T T Y

p os t erior e n ds of t he t wo is c hia fo rm p rom in en t p roj ec tio n s in the rabbi t or a

c u rved ro u g h s u rfa c e in the c a t c alled the is chia l tu b er os it d e t end t oward
, y a n x ,

t he median line as t he r ami ( sin gu lar r a mu s ) o f the is c hium m ee tin

, g in the ,

m edian ven tral lin e t o fo rm the is chia l s ymphys is The an terior ven tral p ar t .

of ea ch i n no mina te bone is form ed by the pu bis E a c h p u bis sends a p rojec tion .

,

the r a mu s t oward the m edian ven tr al line the two ram i u n i tin g as the u bic
, , p
s ymphys is B o th is c hial a n d p u bi c s ym p hyses are in life c om p osed of cart ila e
g
.
.

B e tween the ram i of the is c hium and p u bis is the larg e ob tu r a tor fora men I li u m .
,

is chium and pu bis m ee t a t the a c e tab u lu m ,

a n d t ake p ar t in the form a tion of its walls .

D raw the p elvi c girdle from in fro n t .

On a dem ons t ra tion s p e c im en of half of

the girdle of a ki tten no te c omp le t e sep ara
t ion a n d bo u n daries of ili u m is chium a n d , ,

p u bis N o t e f u r t her the p res en c e in the

.

a c e tab u lum of a sm all bon e the acetab u l ar ,

il i m u

b on e w hi ch fo rm s tha t p ar t of the a c etab u

,

l um whi c h wo u ld o therwise be o ccu p ied by

the pu bis I n the yo u n g of all mamm al s .

the t hree bones of t he p elvi c girdle are sep a

ra t e as in the adu l ts of rep tiles (Fig, .

The hind limb is fairly t yp i ca l The .

femu r has a h ead a grea ter t ro chan t er ,

FI G th h m l d
P la teral to the head (which in the rabbi t
e u an e c

g dlir t b th h wm g th t h
e a ir s bo f e ree o n es o
c o n ti nu es p os t eri orly t erm i na tin g i n a s m all
,

whi h it i
c mps d
co S tippl d gi
o se . e re on s

r p
e t
r esen til g
c ar whi h l t
a e i p
c roje c t
a er io n t h
o sse t h i
es , rd t ro c ha n t er )
, a n d a ,

o b lit ti g th b
era n d i e b tw oun th
lesser t ro chan ter si tu a ted below the head
ar es e een e
, .

b o n es (F m rop dm l
3 d b y th
S e en o an e e
Th e se t ro c han t ers serve for m u s c le a tta c h

an a ot my d p tm t ) e ar en .

m en ts The large ar ti cu la tin g s u rfa c es a t

.

the dis tal end of the fem u r are c o n dyles (m edial a n d la t eral ) a n d t hey bear ,

additional eleva tion s or rou ghen ed ar eas the ep i co n dyles At the k n ee join t ,
.

a p a tella is p resen t The shank is c om p osed of a s to u t t ibia a n d slen d er b u la

.
,

the la t t er in the rabbi t f u sed wi t h the t ibia for the grea t er p ar t of its len gth .

The a n terior fa c e of the tibia p rese n t s a c res t ; its p roxim al ar ti c u la tin g s u rfa c es
are known as c ondyles ; its di s tal ones as m alleoli The bon es of the a n kle ar e .

iden ti c al wi th t hose of the hu m an a nkle a n d are designa t ed by the sam e n am es ,

whi ch are u n for tu n a t ely som ewha t fan cifu l a n d n o t based u p on comp ara tive
, ,

ana tom y The nam e derived from comp ara tive an a tom y is given in p aren
.

thesis af t er the n a m e derived fro m hu m an a n a tom y The a nk le c o n sis t s of seven .

bones ( ca t) or six ( rabbi t) The larges t a nd m os t c o n sp i cu o u s of these whi c h

,
.
,

p roje c t s ba c kward as t h e he e l is the c a l can eu s ( b u l a r e)

,
A r ti c u la ti n g wi th t he .

m all eoli of the tibia a n d b u la is the as tr agal u s or tal u s ( tib ia le) Direc tly in .
 86 LAB O RATOR Y MANUAL F OR VERTEBRAT E ANAT O M Y

s ternum The s t ern u m o ccu rs o nl y in l u ng b r ea thin g ver teb ra t es th a t i s t o say it i s ab s ent

.
-
, ,

in s h es Th ere i s some d ou b t as t o the orig in of the s tern um Acc or din g t o the pr eva il ing
. .

Vi ew g ix en in K a nd W the s t e rn u m of the high er v er t eb ra t es i s pro d u c ed by the f u s i o n of the

, g ,

v ent ra l end s of the ri b s i e of the c os t al c ar til ag es B u t s in c e in the l ow er ver t eb ra t es the ri b s

,
. .
,
.

a r e s h or t a nd d o n o t r ea c h t o the v ent ra l s id e thi s th eo ry n ec ess it a t es the in v enti o n of a dif

, ferent
th eory for the orig in of the s t ernu m of Amphibia The s t ernu m of Amphibia i s c on s equ entl y .

su ppos ed t o h ave aris en th rou gh the fu s i on of c er t a in car til ag es present in the vent ra l
m yos ep t a of u ro d el es S ee W pa g es 1 40
.
43 for d e
, t a i l s of thi s id e,
a R e c ent l y h o w e v e r .
, ,

ev id en c e h as b e en b rou g ht for w ar d in favor of the c o n c ep ti o n th a t the s t ern u m h as t he sa m e

orig in in a ll of the v er t eb ra t es and th a t it s orig in i s ind ep end ent of the r ib s Accor din g t o thi s .

th eory the s t ern u m i s the m edi a n vent ra l por ti o n of

the p ec t oral g i r dl e ; it h as s epara t ed fro m the g i r dl e
a nd ex t end ed pos t eri orl y ; it m ay or m ay n o t c o n
n ec t s ec ond aril y w ith the r ib s ( s ee Fi g
I
.

2 The p arts o f a typi c a l g ir dl e an d for e.

1imb T he p ec t oral g ir dl e i s somew h a t mor e com

.

pl i c a t ed in s t ru c tu re th a n the pel vi c g i r dl e I t .

i
p roco raco
ari s es in par t from the oss i c a tio n of a car til agi
m om n o u s b ar or pl a t e I n thi s p l a t e as in th e c as e . of
the pel v i c p l a t e th r ee pa irs o f bo n es t w o vent ra l
n
pa irs and o e d orsal pa ir ari s e T h es e b on es a re .

o n ea ch s id e : a d orsal b o ne the s capu l a a n ant er i or , ,

vent ral b on e the pr ocoracoid or procoracoid and a , ,

pos t eri or v ent ral b on e the cor acoid ( s ee Fig , .

T he s c apul a i s a n al og ou s t o the il i u m the proc ora ,

c o id t o the pu b i s a nd the c ora c o id t o the i s c hiu m , .

Th es e b o n es l ike the p el vi c b on es are c ar til ag e

b on es s in c e th ey ar i se in c ar til ag e I n the c as e of .

the pec t oral g i r dl e u n l ike the p el v i c on l y two of the

FI G 2 7 . D iagram o fthe b ones of the three cartilage bon es persist in the vertebrate s eries
.
, ,

typical p ec toral g irdl e and fore limb The

.

O f th es e t e s c ap u l a i s al w ay s pr es ent O f the t w o
h
.

cartil age b o n es o f the girdl e are s tippl ed .

t he m em b rane b on e ( cl av icl e) l eft b l an k

v ent ra l pai rs of el em ent s onl y o n e pa i r I S pr es ent 1n

All o f t he lim}, b o es a e cartil ag e b ones

n r
th e m a j or it y of l i.
vin g g ro u ps of r ep ti l e s in b i r d s , ,

( F ro m P ar k er and H a s w ell s T x tb ook o f a nd

in m a mm
e a ls I t i s c o mm o n l y s u ppos ed th a t
Z o l ogy cou r t esy o f t he M acmill an Com
o ,
the el ement w hi c h p ers i s t s i s the c ora c o id a nd th a t
pany ) . the pro c ora c o id i s mi ss in g If thi s th eory b e c or .
~

rec t th er e a r e h ardl y an y l ivin g v er t eb ra t es w ith

,

a separa t e ossi ed pro c ora c o id How ever th ere i s c on s id erab l e evid en c e th a t the per
.
,
~

s is t en t el ement i s reall y the pro c oraco id in presen t r ep til es and th a t in thi s g rou p the co racoid ,

18 mi ss in g I n mammal s h ow ever the matt er i s disp u t ed so me regard in g the p ers is t ent

.
, , ,

el ement as t he pro c ora c o id o th er s as the c ora c o id ( Fi g , Thi s p ers i s t ent v ent ral cartil a ge
.

b o n e o f the p ec t ora l gi rd l e of r ep til es b i rd s and ma mmal s i s ca l l ed t he cor a coid a nd thi s n a me

, , ,

w ill b e ret a in ed h ere w ith t he u nd ers t andin g th a t in r ep til es it is al mos t c ert a inl y ho mol o go u s
,

w ith the pro c ora co id I n n earl y all ma mmal s t he cora co id i s rudiment a ry the o nly car til a ge
. .

bo n e o f t he gi rdl e w hi c h r et ain s it s full i mport an ce b ein g the s c ap ul a o r sh oul d er bl ad e Dia .

grams ill u s t ra t in g the ho mol o gy and parts o f the gi rd l es of vario u s forms are given in
Figu re 2 9 .

In a dditi o n t o the c a rti l age b on es pr es ent in the gi r dl e so me of the origina l cartil ag e is

likely t o pers i s t un o ss i ed T he c ar ti l ag es of mos t c omm o n oc c u rr en c e a re : the s u pras capul a
.
,

F B Hanson American J ourn al of An atomy X

.
, XVI 41 , .
 T HE E ND OSKELETON : G IRD LE S THE STERN U M , ,
AND APPE ND AG E S 87

s itu a ted al ong the d orsal b or d er of the s c apul a a nd sometimes par ti all y os sied the epicor a ,
caids c ar til a g es a nt eri or or medi al t o the c ora c o id pro c ora c o id nt of r p ti i a n a nd
, ( ) el em e e l m am
mal i a n p ec t oral g i r dl es ; a nd c ar ti l ag es b et w een the media l end s of the c ora c o id s al so b u t
(
c on fus ingly n amed ep icor acoids in t ex t s and in this ma nu al Ieft u nnamed),
'

The p ec t oral g i r dl e i s s till fu r th er compl icated by the addition o memb ra n e b ones t o the
f
c ar til age b on es a nd c ar til ag es d es c r ib ed ab ove As previ ou sl y s t a t ed su ch memb ran e b on es are
.
,

homol ogou s wi th the der mal pl a tes of the exos kel eton
c c
and h ave s e ond ari l y b e ome asso i a t ed w ith the c
e nd oskel et o n I t w as s t a t ed in the int ro du c ti on
.

t o the end oskel et o n th a t su ch memb ran e b on es cap la s u

o ccu r in r el a ti on t o c er t a in par t s of the end o

skel et on and we meet th em h er e for the rs t
,

time in asso c ia ti o n w ith the p ec t oral g i r dl e I n

, .

t el eos t s th ere a r e a nu mb er of su c h d ermal

el ement s a dd ed t o the p ec t ora l g i r dl e as s h o w n in
,

Figu r e 3 0 ; th ey form a sor t of v ent ral in c as e

m ent of the g i r dl e O f th es e th er e p ers i s t in the
.

mos t primiti ve l a nd v er t eb ra t es ( ex tin c t ) ve

el ement s
s cap la u
a medi an s ingl e in tercl avicl e pa i red
, ,

l a t eral cl avi cl es a nd l a t eral t o the cl av i cl es pa i r ed

,

cl eithr a ( s in gu l ar c l eith ru m ) ( s ee Fi g 2 A
, 9 ) T he . .

int er cl avi cl e ( al so c all ed ep is tem u m) i s s itu a t ed

i mm edi a t el y v ent ral t o the s t ernu m the c l av i cl es ,

inv es t the p roco r aco ids w hi c h ar e of t en c o n c eal ed

w ithin or d orsal t o th em w hil e the cl eithra ar e
pr c aco id
,
o or
s itu a t ed al on g the b or d ers of the s c apul ae T he .

c l eithr a a r e l os t e ar l y in t he evol u ti o n of l a nd W e

ver t eb ra t es a nd d o n o t o ccu r in l i v in g form s b u t c ac id , or o

the int er cl av i cl es a nd c l avi cl es a r e ch ara c t er i s ti c

of m any rep til es of b i rd s and of m on o t remes I n
, ,
FI G 2 8 Dia gm m5 to m
.
uetra t e t wo

th eori es 0 fthe hO I OIOgy 0 fth i c ar ta g e b one?

pl a c ent al m amm al s the int er cl avi cl e is miss in g P f
the p ec toral
,

a nd in so me of th em the cl avi cl es are al so r edu c ed O f A l aw

l ag e b ones O f the p mm l w e p ec mml gu dl e ;

o r a b s ent T h er e h as th u s o ccu rr ed in the evolu
there a re three b o nes l n each hal f o f the
.

t i o n of the p ec t oral g i r dl e a g ra du al r edu c ti on l n

g irdl e a condition fo u nd o n l y in ex tinc t
the num b er of m emb ra n e bo n es as W i ll b e s een by
,

,
fo rms B o n e theory o f the composition o f _

Tef eren ce to Figu re 2 9

,

the p ec toral g i rdl e o fpresent amnio t es sho w ,

T he p ec t oral g i r dl e thu s c o n s i s t s of a compl ex in g disapp earance o f the procoracoid an d

of car til age an d membr a n e b on es T he c ar ti l ag e
. retention o f the coracoid C al ternative .
,

b on es may ar t icu l a t e vent ral l y W ith the s t ern u m theory showing retention o f the procoracoid ,

b u t rar el y h ave a n y ar ti cul a ti o n o r fu s i on with ( caned howeVer o raCOid) and disappearanc e t

f h e Co ra COid
the v er t eb ral c ol u mn ( ex cep ti o n s : s ka t es p t ero 0 t
,

sau rs) I n thi s resp ec t the pec t oral g i rdl e s t and s

.

in m arked c o nt ras t t o the p el vi c g i rd l e w hi c h as we h ave s een al w ay s h as in l and ver t e

, , ,

b ra t es a rm ar ti cu l a ti o n w i th the sa c ru m I n t el eos t s the memb rane b ones of the g i r dl e

.

m ay ar ti cu l a t e w ith the s kull The c ar til ag e b o n es of the g i r dl e al w ay s a nd the memb ra n e

.
,

b on es n ev er t ake par t in the ar ti cul a tio n w ith the for e l i mb Thi s i s on e of the s imp l es t w ay s
,
.

of di s tin gu i shin g the t w o kind s of c ompon ent s in the g i rdl e .

T he s kel e t on of the for e l i m b i s c o mpos ed of par t s s i mi l ar t o th os e al r ea d y d es c r i b ed for t he

.

hind l im b T h ere i s a proximal s egment the u pper arm or br achiu m c on s is tin g of a s ingl e ,
88 LAB O RATOR Y MANUAL FO R VERTEBRAT E N
A ATOMY

b on e a middl e
the hu mer u s ,
or an tibr achiu m c ont ain in g t w o b ones :
s egm ent the for ear m
, ,

a preaxi al r adiu s a nd a pos t axi al ul n a a nd a dis t al s egment the han d or man u s T he h and , .

i s su b divid ed int o th ree par t s : the wr is t the pal m and the fin gers The w r i s t i s primi ti vel y
, , .

c ompos ed of nine or ten b ones in th ree row s : a proxim al row of th ree n amed the r adial e the in ter , ,

mediu m a nd the ul n are; a middl e row of o n e or two cen tr al es ; a nd a dis t al row of ve

,

FI G . 29 P ec tora1 g irdl es
. some verteb rates A side View o f hal f o f the pec toral girdl e o f an
of .
,

e tinc t amphi b ian Cacops b el ong in g t o the S tegoceph

x ,
al a ; no t e pr es enc e o f the cl eith ru m b
,
B vent ral .
,

Vi ew o f b o th hal ves o f the p ec t oral g irdl e o f a v ery anc ient e tinc t r ep til e S eymour ia b el onging t o the x , ,

Cotyl os au ria ill u strating the compl ete g eneral iz ed pectoral girdl e ; note presence o f b o th coracoid and
,

procoracoid C ventral view o f b o th hal ves o f the pec toral g irdl e o f a modern reptil e a l izard ; not e
D ventral view o fb oth halves o f the pec toral g irdle ofa monotreme
.
, ,

l oss o f o n e o f the coracoid b ones .

,

mamm al the du ckb ill Or n ithor hyn chus i ll us t rating the mos t primitive pectoral gi rdl e fou nd among present
, , ,

mammals ; no te p ersis tenc e o f the coracoid and int ercl avic l e sim il ar t o rep til es E vent ral vi ew o f ,
.
,

o n e hal f o f the p ec toral g irdl e o f an ap e ; no t e ab senc e o f the int ercl av icl e and r edu c tion o f the c oracoid
-

t o t he coracoid proc ess f C artil age cl ose stippling ; cartil age b ones open s tippling ; m emb rane b ones
.
, , ,

b l an k a co raco scap u l a ; b cl eithru m ; c cl avi cl e ; d int ercl avicl e ; e s cap u l a ; f coracoid ( coracoid
.
, , , , , ,

proc ess in E ) ; g procoracoid ; h epicoracoid car til age ; i gl enoid fossa ; j spine o f the scapu l a ( A
, , , ,
.

from W ill iston s W ter R ptil es of the P ast a n d P r es en t University of Chicago P ress ; B after W ill i ston ;

a e

C from Reynol ds The Ver tebr ate S kel eton cou rtesy o f the Macmill an Company ; D from W iedersheim s
, ,

,

Comparative Anatomy of Ver tebr at s cou rtesy o f the Macmill an Company ) e ,

.

car pal es pal m i s c ompos ed of ve el ong a t ed metacarpal s a nd the ngers of phal an ges
. T he
wh os e nu mb ers ar e prim iti vel y the sa me as in the cas e of the t o es ( see Fig .

T h e t r l i d l d t l f l m b n h
I t h e s h
3 e .
p c o a g r e an p e c or a n o e a s o r a c s n es .

we nd the p ec tor a l g i r dl e in a p r imi ti v e c o ndi t i o n res emb l in g th a t o f the p el vi c ,

gi rdl e I t is a cu rved car til age almo st compl etely en cir cl ing the an teri or p ar t of
.
9 0 LAB O RATO RY MANUAL FO R VERTEBRAT E ANATOMY

el o n ga ted p r o ces s T he po s t erio r p ar t is the cor acoid r egi o n b u t n o o s sica t ion

.
,

has o c cu r r ed in ei t h er o f the r egi o ns Dor sa ll y ab v e t he j oin t w i th the fo r e

.

l imb is a b o n e the s capu l a the onl y b o n e in the gir 1e

,
Att ach ed to the d or sal.

b or d er o f the sca p u l a is the s u p rasca pu l ar car til age T h er e is n o s t ern um.

(al th ou gh two or th ree p ai rs o f sma l l car til a ges fou nd in the ven tral wa l l ,

l o c at ed in the myo s ep t a ar e r egar d ed as the s ter num b y s o m e au th or s)

, No te .

th a t the p ec tor al gir d l e has n o co n n ec ti o n w i th the v er t eb ral c ol u mn .

T he fo r e limb is s im il ar to the hin d lim b T he u pp er a rm is com p o s ed o f the

.

hu mer u s ; the for ea rm o f the p r eaxi al r adiu s a n d the p o s t axi al u l n a ; the el em en t s

o f the c arp u s o r w r i s t ar e dii cu l t to dis tin gui s h ( th er e a r e six or s ev en ca r ti l ag es

in th ree r ow s) ; th er e are fou r meta ca r p al s in the p alm fol l ow ed b y p hal an ges .

The midd l e segm en t o f the l im b is b en t a n d di r ec t ed ven tr al l y t h u s el eva ting ,

.

the animal sl i gh tl y ab ove the gr ou nd .

Dr aw the p ec t o ral gi rd l e an d o n e fo r e lim b .

5 . T h e p ec to r al g i r dl e an d s ter n u m o f t h e fr o g I n the f ro g we nd th a t
.

the c ar ti l a g es p r esen t in the u r o d el e gi r d l e h ave b een p ar ti all y r ep l a ced b y b o n e .

A s t er nu m is fu r th er p r es en t in the m edi a n v en t r al l in e b etw een the ven t r a l end s

o f t he h al v es o fthe p ec t or a l gi r d l e S tu dy the girdl e an d s ternum The s t ernum
. .

c o n s i s t s o f two b o n es an d two c a r ti l a g es a rr a n g ed in a l o n gi t u din al s eri es a n d

sep ar a t ed in t o two gro u p s b y the p ec t o r a l gi r d l e T he a n teri or gr ou p p ro j ec ting

.
,

c ra nial l y fr om the gi rd l e c o n s i s t s o f a t erm in a l rou nd ed c a r ti l a ge the ep is tern u m

, , ,

p o s teri or to w hi ch is a b on e the omos ter n u m The p o s ter ior gr ou p p roj ec ting

, .
,

c au d ad f rom the gi r d l e is c om p o sed o f a n a n t eri o r b o n e the s tern u m p r op er a n d

, , ,

a t ermin al r ou nd ed c a r ti la ge the xip his tern u m

,
T he ven t ral p ar t o f the p ec t ora l
.

gi r d l e co n si s ts o f two p air s o f b o n es thei r ven tral end s m eetin g in the m edi an

,

ven tral l in e b etw een the two p ar t s o f the s t ernu m The a n terior p a ir o f b on es .

is the cl avicl es .The cl avi cl es ar e m em b r ane b o n es w hi ch co ver a n d c o n c eal

the p r ocor a coid car til ages ; th ese c ar til ages rem ain u n o ssi ed in An u r a T he .

p o s t eri or v en tra l b o n es o f the gir d l e ar e the cor a coids o ssi ed fr om the cora co id ,

ca r t i l a g es . The m edial end s o f the co r aco id s rem ain in the car ti l a gin ou s co ndi
ti o n ( forming car ti l ages w hi ch a re d esi gn at ed in tex ts as the epicora coids b u t ,

this t erm is co n fu sing a s i t is a l so u s ed in ano th er co nn ec ti o n) The d or sa l p ar t o f .

the gir dl e co n si s t s o f two b o n es o n ea ch sid e : o n e n ex t to the ar ti cu l a ti o n o f the

fo re l imb the s ca pu l a an d a l ar ge thin at b o ne m edi al to the s cap u l a the
, , , , ,

su pr a s capu kz h a vin g a c ar til a gino u s b o r d er

,
T he gi r d l e is n o t co nn ec t ed w i th
.

the ver t eb r al c o l u mn . T he d ep r essi o n whi ch rec ei ves the h ea d o f the h u m eru s

is the gl en oid fos s a ; s cap u l a an d c ora co id tak e p ar t in the form a ti o n o f the g l en o id
foss a b u t the c l a vi c l e d o es n o t T hi s indica t es th a t the cl a vi cl e is o f di ffer en t
,
.

o r i gin from the o th er b o n es o f the gi rd l e Dra w the gi r d l e an d s t er n u m o f the

.

f r og .

6 The pector al gir dl e , s tern u m an d for e l i mb of r eptil e s

.
,
Ou the sk el eton .

o f the tu r tl e ob ser v e th a t a p ec t o ra l gi r dl e is p res en t b u t a s ter n um is ab se n t .

 THE E ND OSKELETO N : GIRD LE S THE STERNU M
, , AND AP P E NDA GE S 9 1

E a ch h al f o f the gi rdl e co nsi s ts o f two ven t ra l p ar ts an d on e d ors al p ar t The .

a n t eri o r ven t ra l p a r t is n o t an ind ep end en t b o n e b u t a p r oj ec ti o n f r om the

s c ap u l a a n d is h en c e n am ed the p r os ca p u l a r p r ocess ( err o n eou s l f d t in

, y r e er r e o

s o m e t ex t s as the p r o co r aco id b o n e) The p os t eri o r v en t ral b o n e the l ar ger o f

'
.
,

the t wo is t he cor a coid (w hi ch as al rea d y exp l ain ed is p rob ab l y in r eali ty the

,

p ro c o ra coid ) The d o r sal b o n e an elo n gat ed b o ne r ea chin g to the carap a ce is

.
, ,

the s ca p u l a Al l o f th ese ar e ca r ti l age b o n es

. In a ddi tio n th ere ar e c er ta in .

b o n es o f the p ec toral g i r d l e w hi ch ar e in cl u d ed in the p l a s tro n an d ar e th ere

fore memb r a n e b on es Th es e ar e the p ai r ed cl avicl es ( ep ip l a s tr a) an d the
.

m edi an in tercl avicle ( en to pl as tr on) R efer to yo ur d rawin g s o f the tu r tl e

.

The p ec to ral

gi dl
r i ga to r co n si s ts o f a s tou t d o rs al b one the s capu l a;
e o f the al l ,

a s t o u t ven t ra l b o n e the c o r a co id ; an d a l o n g sl end er d a gg er sh ap ed b o n e in the

, ,
-

m edi an ven t ral line the in tercl avicl e Cl avi cl es ar e l ackin g

, A s t er num is
. .

p resen t I t is comp o s ed o f a p l a t e o f c ar ti l age b etw een the ven tr al end s o f

.

the c o r a c o id s an d j u s t ab o ve the in ter cl ayicl e ; i t is d raw n ou t p o s t eri or l y in to

l on g cu r ved ca r ti l a g es the xiphis tern a l horn s
, Ob ser ve th a t the rib s ar e a tt ach ed .

to the s ter nu m b y m ea n s o f th ei r s t er n al r i b s a condi ti o n rs t m et w i th in ,

r ep ti l es .

T he b on es o f the fo r e l imb s ar e the sam e a s t h o s e o fNectu ru s in g en eral The .

b o n es o f the ca rp u s or w ri s t o f the tu r tl e are r em a rk ab l y p rimi ti ve in numb er

an d p o si ti o n an d d es erv e fu r t h er a tt en ti o n
,
If n o t suf ci en tl y cl ear o n the sp eci.

m en exp o se th em b y cau ti ou sl y p i ckin g aw ay the d ri ed l i g am en t s c ov erin g th em

, .

At the b as e o i the u l n a ar e two b on es an o u t er u l n ar e an d a n inn er in ter mediu m

, , .

The cen t er o f the c arp us is o ccu p i ed b y a l o ng b o n e whi ch is the fu s ed r adiale ( en d

a t the b a s e o f the ra diu s) an d the cen tr ale ( l arg er en d ) Di s t al to t hi s is a row .

o f ve c arp al es o n e fo r ea c h met a ca rp al
,
Thi s arra n gem en t is th erefore very .

m u ch l ik e t h a t o f the id eal c arp u s F i gu r es o f the carp u s o f tu r tl es are gi ven

.

in W d p a ges 1 6 3 1 6 4 Dra w sh owin g the b o nes of the c arp u s

, ,
.
,
.

7 The pectoral g ir dl e , s tern u m , an d f or e l imb of b ir d s

.
The p ec to r al .

girdl e o f b i rd s is a very compl ete an d general iz ed girdl e I t is composed of the .

s ca p u l a a l o n g sw o r d lik e b o n e l yin g ab o v e the ri b s ; the cor a coid a s tou t b o n e

, ,

r ea c hin g thes t er nu m ; an d the wis hb on e or fu r cu l a in fro n t o fthe cora c o id s a n d , ,

a tt a ch ed a t the l o w er en d b y l i gam en ts o nl y The fu r cu l a is mem b ran e b o ne .

an d r eall y co n s i s ts o f two cl a vicles the two f ork s o f the w i shb o n e u ni t ed to a

, ,

rou nd ed p i ec e the in ter cl avicl e

,
Co racoid an d s cap u l a tak e p ar t in the forma
.

ti o n o f the gl eno id fossa The s tern um of b ird s is hi gh ly sp eci ali zed I t is an

. .

el o n g a t ed b o n e b ear in g a s t r o ng ven tral p ro j ec ti o n the heel o r ca r in a The c arin a ,

.

s er v es fo r the a tta c hm en t o f the po w erfu l w in g mu s cl es As in rep t i l es the ri b s .

,

ar e j o in ed to the s t er nu m b y th ei r c o s t al c ar til a g es The fr o n t en d o f the s t ernum .

has sh or t cos tal p r o ces ses each s id e two lo ng xip his tern al p ro cesses
,
Dr aw from .

the s id e sh o win g gi r d l e an d s t er nu m
,
.
9 2 LAB O RATORY MANUAL FOR VERT EBRAT E ANATOMY

On d em o ns tra ti o n sp ecimen o f the s ter num o f an o s tr i ch or o th er ightl ess

a

b ir d n o t e the ab sen ce o f the k eel o r c arina the s tern u n p r esen tin g a co n vex ven
z ,

t r al f ace:

T he for e l imb o f b i rd s is gr ea t l y mo di ed as a w ing p a r ti cu l arl y in its di s ta l ,

p or ti o n s T he humerus is s tou t ttin g in to the gl en o id fo ssa b y its co nvex

.
,

head . O n ei th er sid e an d sli gh tl y di s tal to the h ead ar e p rom in en t p ro j ec ti o ns ,

the gr eater an d less er tu ber os ities co r resp o ndin g to the t ro ch a n t er s o f the f emu r
, .

T he l es ser tu b ero si ty is co n tinu ed di s tal ly in t o a sh arp r id g e the del toid r idge , .

T hi s sid e o f the hu meru s b earin g the d el t oid r id ge is the p reaxi al or ra di al sid e ,

an d i t is r eadi l y o b ser ved th a t a r o ta ti o n has o ccu rr ed s o th a t the p r ea xi al s ur f a c e

fa ces dors all y in the fol d ed p osi ti o n of the w in g The gr ea ter tu b ero si ty is p os t .

axial an d b ea rs o n its u nd er s ur f a ce a l ar g e h o l e the pneu ma tic for amen ,l ea din g ,

.

in to the air sp ace o f the hum eru s T he ra di u s an d u l n a are typ i c al the fo rm er

.
,

the m o r e s l end er o f the two the l a tter exhib i tin g a t its p r oxim al en d a p r o j ec ti o n
, ,

the ol ecra n on pr ocess o r el b ow h ere met fo r the r s t t im e

,
T he w ri s t is g r ea tl y .

al ter ed co n s i s t in g o f b u t tw o s ep a r a t e b on es
the r adial e a t the b ase of the radiu s
,

a n d the u l n a r e a t the b a s e o f the u l n a T he r em aini n g w r i s t b on es a re fu sed to the

m et acarp al s to form the car p ometacar pu s con s i s tin g o f two el on ga t ed b on es
.

, .

T he metacarp al s co n t rib u ting to th es e el emen ts a r e the thi r d an d fo u r th The .

seco nd m e ta car p al is fus ed t o the p r eaxi al s id e o f the p r o xim al en d o f the thi r d

m eta ca rp al w h er e i t f orms a p ro n ou n ced hu mp F rom thi s hump p roj ec ts the .

s ec o nd di gi t The thir d di gi t is the l o n ges t an d co n s i s t s o f two p h al a n ges

.
,

o f w hi ch the p r o x im al on e is mu ch a tt en ed The fo u r t h di gi t is a sm all p rojec

.

tio n fu sed to the p o s taxi al sid e of the p roxim al p hal anx o f the thi rd di gi t F irs t .

a n d f th m eta ca rp al s a n d digi t s ar e w a n tin g .

8 The pectoral gir dl e , s tern um , an d fore l i mb of mammal s

.
The p ec tor al .

gi rd le o f mos t mammal s is redu ced an d s omewh a t mo di ed O nl y the mo no .

tr emes h ave comp l ete p ec toral gir dles con si s tin g o f scap u lae cora coid s cl avi cl es
, , , ,

an d in t er cl a vi cl e ( see F i g 2 9D p.
,
In all of the p l a cen tal m amm al s the co ra
.
,

c oid is r ed u ced to a sm a ll p r o c ess the cora co id p r o ces s the in t er cl a vi cl e is

, ,

m i ss in g an d the cl avi cl es are o f t en red u ced o r w a n tin g ( F ig 2 9 E )

, Con se . .

q u en tl y th e p ec t ora l gi r d l e o f m o s t m amm al s c o n s i s ts o f the s cap u l ae o nl y ,

w hi ch are co rr esp o ndin gl y enl ar ged a n d imp or tan t as p l aces of mu s cl e a tt ach

m en t .

S tudy the pectoral girdl e of the rab b i t or ca t I t consists of two pairs of .

b o n es the clavicles an d the s capul ae

, The cl a vi cl es ar e sma l l sl en d er b on es
.

imb edd ed in the mu s cl es o f the fron t o f the shou l d er an d as th ey ar e n o t ar ti c ,

ul a ted to the r es t o f the g ir dl e th ey gen er all y fal l o ff in p r ep ar ed sk el et o n s I so .

la t ed cl a vi cl es will b e d em o ns tra ted an d th ey w ill b e s een in p l ace l a t er wh en the

d in some tex ts however as the second an d third an d the metacarpal here ca ll ed second
Reg arde
According to L illie Devel opmen t ofthe Chick the emb ryological evidenc e s t rong l y indica tes
, , ,

as the rs t .
, ,

tha t th pe rsis t ent meta ca rpal s are the seco nd third an d fou rth
e
, , .
94 LAB ORATOR Y MANUAL FOR VERTEBRAT E ANATOMY

T he w r i s t is com p o s ed o f a numb er o f sm all b o n es arra n ged in two tra n s ver se ,

row s . T he p ro xim a l r ow co n si s ts o f fou r p i eces in the rab b i t th r ee in the c a t ; ,

.
the di s tal row o f ve in the ra b b i t fo u r in the ca t Ar ti cu l a tin g wi th the di s tal
,
.

p
en d o f the ra diu s is the l a r ge s ca hol u n ar in the c a t s ep a ra t ed in the r a b b i t in t o ,

a m edi a l n a vicu l ar ( r adiale) a n d a l a t era l l u n a te b on e (in ter mediu m) L a ter al .

to the l u n at e p o r ti o n o r b o ne an d ar ti c u l a tin g w i th the u l n a is the tr iqu etra l b on e

p
( u ln are) The is ifor m is the el emen t p r o ec tin g p rom in en tly la t eral to the
. j
tri qu etral b o n e in the ca t o r b ehi nd i t in the rab b i t The dis tal row of p i eces .

g
b e innin g a t the m edi al sid e a n d p r o c eedin g l a teral l y a r e : the grea ter mu l ta n gu l ar
( r s t ca r a l e ) th p
e l es s e
,
r mu l tan gu l ar ( secon d c ar a l e) the cen tra l ( cen tr a l e m i s s in gp , ,

p
in the ca t) the ca ita te (third ca r ale) an d the hama te (fou r th an d fth car ales
,
p ,
p
fus ed) T h es e carp a l es a r e si tu at ed a t o r near the p r oxim al end s o f th eir r esp ec
.

ti ve metacarp al s Th ere are ve metacar al s o f w hi ch the r s t is ver mu ch

. p y
,
g
red u ced an d ve di i t s w h o s e t erm in al p h a l a n g es su pp o r t the h or ny cl a ws .

D . GE NE RL A SU MM ARY OF THE I
G RD E L S ,
TH E S TR M
E NU ,
AND TH E

PA R
I E D AP P E NDA GE S
r. pa i red app end ages prob a bl y r epr esent r emn ant s of a pa i r of c o ntinu ou s l a t eral n
T he
fol d s s u ppor t ed by c ar til ag in ou s n ray s
,
.

2. The two g ir dl es pro b a b l y aros e th rou gh the fu s i o n in the medi a n l in e o f some of th es e

n ray s .

3. T h e pr im iti v e g i r dl es a r e b ars o r p l a t es o f c ar til a g e in w hi c h s u b s e q ent l y oss i c a ti on

u

o ccu rr ed w ith the forma tio n of car til age b on es .

4 I n ea c h gi rdl e th r ee pa i rs of c ar til ag e b on es ari s e

. T h es e are il iu m pu b i s and is chi u m
. , ,

in the p el vi c g i rdl e ; s capu l a pro c ora c oid a nd c ora c o id in the sa me r espec tive pos iti on s in the
, ,

pec t oral g i r dl e .

5 T he p el vi c gir dl e af t er the th r ee pa i rs of b on es h ave ari s en u nd er goes l itt l e ch a ng e

. In .

bir ds an d mamm al s the bou nd ari es b et w een the b o n es ar e l os t by fu s ion pro du c in g on ea ch ,

s id e an inn omin a te b one Th ere are n ever a ny memb rane b on es asso c i a ted w ith the pel vi c
.

g i r dl e .

6 T he p el vic gi rdl e is free in s h es b u t in all o th er ver t eb ra t es i s rml y ar ti cul a t ed or

.

ankyl os ed imm ova b ly fu s ed) to the sa c ru m by mea ns of the sa c ral r i b s .

7 The p ec t ora l gi r dl e on the o th er h and exhi b it s man y mo di c a ti o ns among the

.
, ,

ver t eb ra t es and is fu r th er c ompl i ca t ed by the a dditi on of memb ran e b on es .

8 I n the ma j orit y o f the l and ver t eb ra t es o nl y two o f the th ree car til ag e b on es of the
.

primitive gi r dl e pers is t The s capul a i s al w ay s pres ent On e vent ral el ement i s g en erall y
. .

pres ent call ed the c ora c o id a nd b el ieved by some to b e h omol og ou s w i th the c ora co id in the
, ,

primitive gi rdl e by o th ers t o b e h omol og ou s w ith the pro c ora c o id

,
.

9 I n all of the pl a c ent a l m ammal s the c ora c o id el ement is r edu c ed to a ves tig e the
.
,

corac o id p rocess borne on the s capu la , .

I O The m amm al ia n s c apu l a is dis t in gu i s h ed by the poss ess i o n o f a sp in e a nd o f the c ora

.

co id pro c ess menti on ed in para g rap h 9 .

n , T he m emb ra n e b o n es a dd ed t o the p ec t ora l g i r dl e vary in d i er en t form s b u t in l iv in g ,

l an d ve r t eb ra t es u su ally c o n s is t of pa ir ed cl avicl es and a medi an u n pa i red int er cl avi cl e .

1 2 The p ec t oral g i rdl e is v ery rarel y co n n n ec ted w ith the ver t eb ra l c o l u

. mn by an ar ti cul a
tion or an k yl os is .
 TH E E ND OSKELETON : G IRD LE S THE ST ERNU M , , AND APPE ND AG E S 95

13. s t ernu m occu rs rs t in the Amphibia and i s fou nd in mos t ver teb ra t es ab ov e
T he
Amphibia I t i s prob abl y the medi an vent ral por ti on of the pec to ral g i rdl e I n many rep tiles
. .
,

and in b i rd s and mammal s the ri b s ar ti cul a t e w ith the s t ernu m

, .

1 4 T he b on es of the l im b s ar e pro b a bl y d eri v ed fro m the c a r til a g in ou s n ray s o f the l o w er

.

s h es Th ey are very s imil ar in all l and ver teb ra t es the dis t al po r ti on s b ein g s u b j ec t to the
. ,

mos t mo di c a tio n s .

I 5 T he l im b s a r e di vid ed int o th ree s eg ment s

. T he b on es of th es e s eg ment s are as foll o w s
.
,

th os e o f the hind l imb b ein g n amed rs t : proximal s egment on e b one femu r o r hu meru s ; , ,

middl e s eg ment two para ll el b on es tib i a and b ul a ra diu s a nd ul n a ; dis t al s eg ment co m

, , , ,

pos ed ofth ree par t s ankl e or w r is t sol e o r palm and dig it s Ankl e or w ri s t c on s i s t s pri mitively
, , , .

of n in e or ten b on es in th ree ro w s : a proximal row of th ree n amed tib ial e o r radi al e int er
, ,

mediu m and b u l are o r ul n ar e ; a middl e row of on e or two c ent ral es ; and a dis t al row o f
,

ve t arsal es o r c arpal es Loss o r fu s ion of any of th es e el ement s is qu it e c o mmo n The

. .

pal m or sol e i s c ompos ed o f ve parall el b on es the met a t arsal s o r met a c arpal s The digit s
, .

c on s is t of p h al anges of w hi ch the pr imitive nu mb ers are pro ceedin g from the rs t to the f th
,

dig it 2 3 3 3 3 Loss of dig it s i s al so c ommon ; n earl y al w ays the inn er and ou t er dig it s
, , , , ,
.

di sapp ear w ith r et enti on of middl e o n es ; very rarely are middl e dig it s l os t Ex t ra digit s .

may al so o cc u r u s u all y on a qu a ti c forms

,
.
 E ND O S KE LET O N: CO M P ARATIVE ANAT O M Y o r

VIII . THE T HE
T HE S KULL AND T HE VI CERAL S S KELET O N
T he or cran iu m i s th a t par t of the end oskelet on w hi c h i s fou nd w ithin the h ea d
s ku l l

w h er e it c overs a nd pro t ec t s the b ra in T he s ku ll i s the m os t c ompl ex of all the par t s of the

.

end os kel e t o n o w in g in par t t o the fa c t th a t it i s d er i ve d from s ev eral dif

,
ferent sou r c es T o .

d et erm in e the h o mol og y of the b o n es of the sku ll of differ ent v er t eb ra t es i s a t r emend ou s

t ask w hi c h i s n o t y et c om pl et ed The earl i er in ves tig a t ors n amed b o n es in d ifferent s kul l s
.

by the sa me n ames w ith ou t a n y a d equ a t e ev id en c e th a t th ey r eall y w ere h om ol og ou s w ith ,

r al
in te n ear
i t r al
n e n ear

b asal pla t e

o ti c caps l u e

scl rotom
e e

ve rt b ral
e

FI G 3 I . .
D ia rams
ill ustrate the d evel opment o f the chondrocrani um from the p rec hordal s
g to
sense capsul es and scl ero tomes o f the verteb rae o f the occipital region A earl y s tage
p ar a chor dal s , , .
,

with the variou s cartilag es s eparate B l ater s tage showing u n ion o f the p rechor d l s with each o th er
.
, ,
a

and with the ol fac tory capsul es t o form the ethmoid plate and u nion o f the p a r acho d l s with each
,
r a

o ther and with the o tic capsul es and occipital ver teb rae to form the b asal pl a te ( Comb ined from .

g ures b y Good rich and W ilder ) .

r esu l tin g c o n fu s ion I n the foll o w in g pa ges a n a tt emp t i s m a d e t o fa c il it a t e the s tu d y of

.

the s kull by a n al yz in g the s ku ll int o it s var i o u s c o mpo n ent s a nd t ra c in g in the p h yl o g en eti c

s cal e the u ni on of th es e c om po n ent s in t o the c o mpl et e skull .

A . THE CAR TI LA GE S TAGE OF THE S KU LL

I . O r i g in o f the c h o n dr o c rani u m

ar is es in the mes en c hyme of the hea d T he
T he s ku ll
. .

rs t s t ep in the form a ti o n of the s ku ll i s the pro d u c ti o n of car til a ge by the mes en chyme Fi ve .

pa ir s ofcartil a ges appear On e pa ir forms al o n g the s id es of the a nt eri or en d of the n o t o c h or d

,

96
9 8 LAB O RATORY M A NU AL FO R VERTEB RATE ANATOMY

the c ho n d ro cra nium p resen ts a l ar ge d ep ress io n , the or b it, whi ch in l i fe hol d s

the eye .

a ) Dors al f
s u r a ce f
o the chon drocr a n iu m:
dy the d orsal su r fa ce T he St u .

ro s tru m o p en s d o r s a l l y b y a l a r g e egg s h ap ed ca v i ty the a n ter ior fon tan elle

,
-
, .

At ei th er s id e o f the b a se Of the ro s t ru m is an olfa ctor y ca p s u le a p r oj ec t in g ,

s t ruc t u r e w i t h thin w a ll s P o s t eri or t o t he ca p su l e an d c o n tin u o u s w i th its

.

d o r s al wa ll is a thi ck p r oj ec tin g sh el f the s u p r aor bital cr es t w hi ch form s the

, ,

d ors al wall Of the or b i t The p os t eri o r en d Of the cr es t con tinu es in to a p ro

.

j e c ti on ,
th e p os tor bi ta l p r oce ss A l o n g the m e di a l
. b or d er s O f th e s u p r aor b i t al

cr es t s ru n s a row O f o p enin g s whic h a r e n er ve foramin a i e , Op enin gs th r o u gh ,

. .
,

whi ch n erves p a ss to an d from the b ra in I n the m edi an l in e j us t b ack O f the .

a n t eri o r fo n t an ell e is an op enin g the epip h yse a l fo r a,

m en t h ro u g h w h ic h a por ,

ti o n Of the b r ain (ep ip hys is o r p in eal b o dy) ex t end s I n the medi a n l ine Of the .

p o s teri o r p art O fthe roo f is a r ou nd ed d ep ressi on the end olymphatic foss a in whi ch , ,

ar e s i t u a t ed two p a i r s O f o p enin g s the t erm in a ti o n s Of the en dolymp h ( sm al l er

,

o u t er h o l es) a n d p er il ymp h (l a r g er m edi a l h o l es) ducts O f the in t er na l ea r Th ese .

d u c ts are ca nal s which co nn ec t the u id l l ed chann el s Ofthe ear w i th the sur fac e
-

O f the s k u ll O n ea ch s id e O f the end ol ym p h a ti c fo ssa is a m assi ve regi o n w hi ch

.

is the au ditory or otic ca ps u l e fu sed to the c ra ni u m ,

T he p o s t eri o r en d Of the .

c r a niu m b ea r s a n o p enin g the for a men magn u m v i si b l e d o rs all y j u s t b ack Of

, ,

the end ol ym p h a ti c f o ss a T h r ou gh thi s o p enin g the b ra in is c on tin u ou s w i th

.

the sp in al cor d Dra w the ch o nd r o cra niu m fr o m the do rs a l sid e

. .

b) Ven tr al a n d l ateral s u rfa ces of the chon dr ocr an iu m: The ven t ra l su r fa ce O f

the ro s t ru m b ea rs a p ro j ec tin g c ar in a o r k eel O n ei t h er s id e O f this m edi al k eel
.

is an o val Op enin g w hi ch m ay b e d esi gn a t ed the r os tr al fen es tr a

,
Th rou gh the .

r o s tr a l f en es t ra e o n e c a n l ook in to the ca v i ty O f the cr a ni u m w hi c h in l i f e is ,

o cc u p i ed b y t he b ra in L a t er al to each r o s t ral f en es tr a is the thin wa lled

.
-
,

olfactory cap s u le I n mo s t sp ec imen s p a r t Of the w al l s o f thi s w il l p rob ab l y

.

b e b rok en aw ay W h en com p l ete the n a sal cap su l e is a n earl y s p h eri c al s tru c

.
,

tu r e w i th a r el a ti vel y sm a ll o p eni n g to the ex t eri o r At the b o ttom O f the n a sal .

c ap su l e is a l ar g e O p enin g w h i ch l ead s in to the c avi ty O f the sk u l l P o s t er i or .

to ea ch n a s al cap su l e an d c o n tin u ou s w i th its p o s t er ior w a l l is the p reor b ital

pr ocess whi ch form s the an t eri or wal l Of the orb i t The wall s Of the orb i t are
,
.

pi er ced b y nerve fo ram in a Of whi c h the l ar ges t l o ca ted in the a n t ero ven tr al ,

r egi o n O f the o r b i t is the op tic for amen t hrou gh w hi ch the op ti c n er ves p as s

, ,

from the eye The v en t ra l w a ll o f the sk u l l b etw een the two o rb i t s is ra t h er

.
1

narrow it s narro w es t p o r ti on b ein g the p l a ce w h er e a p ro c es s Of the u pp er jaw

,

ar ti cu l a t es .The p o s t erior p a r t O f the ven t ra l su r fa ce fo rm s a b roa d b as al pl ate ,

wh o se sid es ar e c omp o sed O f the o ti c c ap sul es I n the m edi an l in e o f the b as al .

p l a te w ill b e seen a s tr eak Of sl i gh t ly di ffer en t c o l o r from the ch ond ro cranium .

P rojec tin g into the or b it may b e present a mu sh room sha ped st ructure -
,
the o ptic pe
dicel , which
in li fe su ppor ts the eyeb all .
 THE E ND OSKELETON : S KULL AND VI S CERAL S KE L E T ON 99

Thi s s t r eak is the n otochord I ts a n terior en d tur n s d or sally in to the ca r ti l ag e

.

an d t erm in a t es a t a b o u t the l ev el O f the sm all m edi a n fo r am en l i n g n ear the

y
a n t er i o r en d O f the b as al p l a t e th r ou g h w hi c h the in t er n al c a ro tid a r t er p
y as s es .

I t w il l b e u nd er s to o d th a t tho s e p or ti o n s Of the cho ridro craniu m l yin g t o ei th er

sid e Of the n o t o c h or d are the p ro d u c ts O f p ar a chor d al s an d th a t the p a r t s an t er ior

to thi s u p to the Ol fa c tory cap su l es ar e the p ro d u c t s O f the pr echord al s The .

p a ra c h o r d a l s a r e f u s ed l a t era ll y w i th the o ti c cap su l es an d the p r echo rdal s w i th

the O l fa c t ory c ap s u l es T he l a t er al an d d o rsal w all s Of the b rain ar e p ro d u ced
.

b y ex t en si o n Of th es e o r i gin al p ar t s The p o s t eri o r en d O f the v en t ral s u r f ace

.

O f the sk u ll is p r o d u ced a t ei th er sid e Of the n o t o ch o r d in to a s l i gh t p ro c ess t he ,

occip ital con dyl e w hi ch a r ti cu l a tes w i th the rs t ver teb ra Dra w a v en tral
, .

view Of the cho nd ro c ra niu m .

B . THE VI S CE RAL S KE LT
E O N
T he vis cer al s kel eton or spl h an c n ocran iu m is th a t p ar t
ndo sk el eton Of the e

whi ch su pp or ts the gi l l s . eTh gi ll

s ar e l o ca t ed in t he ven t ral an d p o s t er i o r p a r t

O f the h ea d TO su pp or
. t th gil l
e s an d en ab l e th em to b e m o v ed for resp i ra t o ry

p u rpo s es a sp eci al sk el e to n is p r esen t w hi ch is c al l ed the vi s cera l s kel eto n ,

b ecau se gill s ar e p ar t O f the w all s O f the di ges ti ve t r a c t a s w il l b e d em o ns tra t ed ,

l a ter . The vi s ceral sk el eto n co n si s t s O f a l o n gi t u din al s eri es Of c res cen t sh ap ed -

c a r ti l a g es ( o r b o n es) si tu a t ed b et w een the gil l s l i t s in the p h aryn g eal w all E ach .

su ch el em en t is d esi g n a t ed a gil l a r ch T h er e are t yp i call y seven gill arch es in

.

ver teb ra tes a l thou gh som e el asmob ra n c hs have nin e ( F i g 3

,
T he gi l l ar ch es
.

in the l ow er ver teb r a tes are cl o sel y as so ci a ted wi th the cho ndr o cra ni um an d in ,

the cou rs e Of evo l u ti o n s om e O f t h em t ak e p ar t in the p r od u c ti o n Of the sk u ll .

I . The vi s c er al s kel e to n of the d og s h O b tain a sp ec im en in which the

.

Visceral sk el eto n has b een l ef t a tta ch ed to the ch o nd ro cra nium an d s tu d y i t

ca r efu l l y The sev en gill ar ch es form a s eri es Of cu rv ed ca r til ag es v en t r a l to

.

the po s t er i or p ar t O fthe ch o nd r ocr a ni um an d ex t endin g p o s t eri or l y to the p ec t ora l

gir d l e The rs t gi l l ar ch the man dib u l ar ar ch is the l arges t an d m o s t m odi ed
.
, ,

Of the s eri es I t is s een w h en vi ew ed from b el ow to c o n sis t o f d o rs al an d ven t r al

.

hal ves E a ch s id e Ofthe d o rsa l h al f is c all ed the pal atoqu adr ate or p terygoqu adr a te
.

c ar ti l a g e ; in p r o l e v i ew i t w i ll b e s een t h a t thi s c ar t i l a g e is c l o sel y a pp l i ed to the

ven tr al su r fa ce Of the ch o nd ro cra nium to w hi ch in l i f e i t is a tta ch ed b y a l i ga

,

men t . I t send s u p a w el l d evel o p ed p al a tal p ro cess in to the orb i t The p terygo

- .

d t t i l a g s b ea r t ee th an d in f a c t con s titu te th e u pp er jaw of th e an ima l

q u a ra e c a r e , ,
.

T he ven t ral h al f Of the mandib u l ar ar ch co n s i s t s Of two h al ves ea ch O f which is ,

kn own as Meckel s car til age T h es e b ear teeth an d to geth er con s titu te the lower

.

j a w o f th e d o g s h T h.e w id e g pa b e t w ee n t he tw o j a w s is the m ou th Op enin g .

At th i
e r p t
os er o i r e nd s th e p t er g
y qo u a d ra t e an d M e ck el s c a r ti l a g es j o in a

t a n

a cu t e a n g l e ca l l ed the a n gl e o f the jaw f o rmi n g a hin g e j oin t p erm i ttin g o p enin g

, ,

an d cl o s in g O f the m ou th .
I OO LAB ORATO R Y MANUAL F OR VERT EBRAT E AN ATOMY

The s ec o nd or hyoid ar c nd er th a n the m a ndib u l ar ar ch to the

h is m o r e sl e

p o s t eri or f a c e O f w hi ch i t is c l osel y a pp l ied I t cgn sis ts o f a v entr al m edi an .

p i ec e the b a s ihya l ; a sl end er b ar the cer a tohyal o n ea ch sid e O f the b a sihyal ;

, , ,

an d a s t o u t p i ece the h y om a,
n dib u l a r d o r s al t o ea c h c era t o hya l,
T he hy m an
o .

dib u l a r a r t i cu l a t es t o the o t i c regi o n O f the sk ul l an d t hu s a c t s a s a su sp en s or Of

the l o w er jaw T he h yoma ndi b u l a r b ear s o n its p o s t er io r m ar gin s om e s l end er
.

p ro j ec ti o n s the gil l r ays w hi ch in l i fe s u pp or t the gi ll s

, , .

T he r em a inin g ar ch es kno w n sim p l y a s gil l or b r on chial ar ch es ar e simil ar

, ,

t o ea ch o t h er E a c h co n si s t s t yp i call y O f ve p i eces n am ed f r om the d or sa l

.
,

sid e v en t r a l l y : p har yn gob r a n chial t he m o s t d or sa l p i ece el o n ga t ed an d di r ec t ed

, ,

p o s t eri orl y ; epib r an chial the su cceedin g mu ch s h or ter p i ec e ; cer atobr an chial
, ,

FI G 3 3. .
D ia ram Of
g the chondrocrani
verteb ral col umn and gill arches of an el asmob ranch
u m, ,

t o sho w par ticu l arl y the part s and r el a tion s O f the s even g ill arches ( Sl ightl y modied from Viall et on s

.

El emen ts de M or phol ogie d es Ver tbr s .

)

a n o th er n ga t ed p i ece ; hyp obr an chial cu rv ed ven tra l p i eces O f whic h t h er e

el o , ,

ar e b u t t h r ee p a i r s to the v e b r a n chia l a r c h es ; a n d the b as ib r a n chia l s tw o in ,

nu mb er an a n t eri o r sm al l o n e s i tu a ted b etw een the m edi al end s o f the r s t

an d seco nd p ai r s O f hyp o b r a n chial s a n d a l ar g e p os t er i o r p i ec e b etw een the b ases

O f the f t h c er a to b ran chial s a n d t ermin a tin g in a c au d al l y di r ec t ed p o in t .

E pi an d cera tob ran chi al s b ea r gil l rays No te th a t the gil l a r ch es a re n o t .

a tt a c h ed t o the v er t eb ral c o l u m n A di agra m O f the gi l l a rch es is gi v en in

.

F i gu r e 3 3 .

Dra w f rom the s id e s h o win g ch o nd ro cra ni u m an d v i scer al sk el eto n

,
.

C . T HE FO RM T I
A O N OF TH E ME M B RA E N BO NE S OF TH E S KU LL
I . O ri g in o f the m em b ran e b o n e s
s a dditi on t o the c h ond ro c ranium a nd
o f the ku ll In
.

g ill ar c h es s till a n o th er c ompon ent ent ers int o the forma ti o n O f the s ku ll T hi s c ompo n ent
, .

c o n s i s t s O f the der mal or membr an e bo n es of the s kull T h es e el em ent s rs t app ear in the .

g an o id s h es I n th es e forms it ca n b e O b s erv ed th a t the g an o id s c al es h ave b ec ome fu s ed

.

o n the h ea d t o form l arg e b on y pla t es in cas in g the h ea d T h es e s c al es w hi c h as we h av e .

,

al r ea d y l earn ed ar e d erm al in ori g in s ink int o the h ea d a nd appl y th ems el v es cl os el y t o the

, ,

c h ond ro c raniu m and m andibu l ar ar ch I t h as al rea dy b een po int ed ou t th a t in the maj ori ty
.

10 2 LAB ORATO RY M AN UAL F OR VERT EBRAT E N
A ATOMY

b i tal

p tt mporal
os e

s tal plat s
ro e

po t fro tal
s n

Oc c1p ita ls

pos ttempo

FI G 34
D rmal sca
e l es ( superc ial b ones Of Of
h ead Of some ganoid shes A
the s kull ) the
C s turg eon ( Acipen s er ) D gar pike ( Lepidosteu s) T he arrangement Of
. .
.
,

Amia B P ol ypter u s
.
,
.
,
.
,
.

some Of these scal es which in t el eosts b ecome the b ones Of the dorsal su rface Of the skull greatly resemb l es
, ,

that Of the dorsal sku ll b ones Of the l and vert eb rates and these scal es have rec eived the same nam es
as the skul l b ones al thou gh the s tric t homol ogy is dou b tful (After Good rich in P art IXOf L ankester s
,

.
,

Tr atis e on Z ool ogy c u r t esy Of the M acm ill an C ompa ny )

e ,
o .
 THE E ND OSKELE TON : SK ULL AND VI S CERAL SKELETON 103

c ) M em b r a n e b o nes of th e gar p ik e: ( F i g 3 4D ) T he h ea d i s p ro l
. o n g e d in t o

a l o n g s n o u t h a vin g the n a s al o p enin gs a t its ex t r em i t Th m l l l b n

y e s a n a s a o es .

su r rou nd the n asa l o p enin g s P o s t eri or to t h ese o ccu p yin g the cen t er O f the
.

dorsal su r fa ce Of the s nou t ar e the elo n ga t ed ethmonas al b o n es P o s ter i or to .

th em ar e the fr on tal s w hos e an terior end s in close the po s t erior end s o f the
,

ethmo n a s al s P o s t er i o r to the fro n tal s ar e the l ar ge p ar ietal s an d b ehind th em

.

a n um b er O ftemp or a l s The ed ges O fthe u pp er jaw a re comp o sed O fthe ma xil l ae

.
,

b earin g t eeth an d each co n s i s tin g o f a s eri es O f s qu ari sh b ones T he l ow er j aw s .

co n s i s t c hi e y o f the l o ng den tar y b o n es b ear in g t eet h T he s id es Of the h ead

, .

b ehi nd the o r b i t ar e co ver ed b y a l ar ge numb er O fsm all cheek p l a tes an d p o s ter i o r ,

to th em b y the l a r g er opercu l a r s
, .

d) Memb r an e b on es of the b own : The s ku l l OfAmia is p erh ap s l ess sp eci al

ized t han in a n y o th er l i vin g t el eo s to m e a c cor din g to L ( F i g 3 4A) I d en ti fy

. .
,

o n the sk u ll a n t eri o r s t a lk ed a n d p os t er i o r n o s tr i l s a n d the o r b i t s B etw een the .

a n t eri or n o s tr i l s p ro j ec t s the sm a l l t r i a n gu l a r der mal mes ethmoid a mem b r ane ,

b o ne . An t eri o r to thi s are the p remax il l ae b ear in g t eeth Co verin g the sp a ce

, .

b etween the f ou r n o s tri l s ar e the n as al s p o s t er i or to t h em in the media n l in e the

,

l ar ge fr on tal s p o s t eri o r to th em the p ar ietal s a n d at the p o s t e ri or en d Of the

, ,

s k u l l the f o u r temp or al s B el ow an d an t eri or to the o r b i t is the l arg e l acr imal

.
,

b el o w the o r b i t s the two s u b or b ital s an d p o s t eri or to the o r b i t two lar ge p os t

,

or b ita l s . F o rm in g the s id es Of the u pp er jaw are the to o t h b ea rin g max ill ae -

.

B e tw een the d o r s al p o s tor b i t al an d the p ar i etal is the squ amos al

, ,
The .

o p er cu l um is su pp or ted b y s ev era l op er cu l ar b on es .

e) Demon s tr a tion of th e sep ar a te or igin of chon dr ocr an iu m an d memb r an e

b on es : E x am in e the d em o n s tr a ti o n sp ecim en Ofthe h ea d O f a ga n o id sh in w hi ch

the in ca s in g mem b ra n e b o n es h a ve b een loo sen ed from the u nd erl yin g ch o nd r o
c ra ni u m . R em o ve the s h ea th O f m em b ra n e b o n es no tin g th a t th ey a re s i tua t ed ,

in the skin an d n o t e the cho nd r o cra ni um sim i la r to th a t Of the dog sh lyin g

, , ,

wi thin the sh ea th S u ch a sp ec im en s ho ws cl ea rl y the o r igin Of the sku l l from

.

two sep ara t e s o u r ces the ca r t i l a gin o us c h o nd ro cra ni um an d the d erm al sca l es
,
.

D TH E FO RM TI
A ON O F CARTI LA GE B ONE S OF THE S KU LL AND
THE THE

C OMP O S I TI ON OF THE C O MP LE TE S KU LL
.

T he n ex t a nd l as t s t ep in the form a ti o n O f the s ku ll i s the pro du c ti on O f c ar til ag e b on es

in the c h o nd ro c raniu m in cl u ding the s en s e c apsu l es and in the mandi b u l ar and hy o id ar c h es

.
, ,

I n d enit e r eg i o n s O f th es e s t r c t res e
u u c n ter s of os s i ca tion th a t i s c ent ers O f b o n e for m a ,

tionari s e Ea ch Of th ese c ent ers t ran sforms a c er t a in area of c ar ti l ag e int o b on e a nd

. ,

ea ch su ch b o n e i s prim iti v el y a c ar til a g e b o n e O f the s ku ll I t h as al r ea dy b een s t a t ed th a t

.

in the maj ority O f ver t eb ra t es the ch ond ro c ra niu m pr es ent in the emb ry oni c s t ag e o nl y , ,

c on s i s t s O f a vent rall y s itu a t ed pl a t e w ith neith er l a t eral n or d orsal w al l s exc ep t a t the ext reme ,

po s t eri or en d ( Fig 3 . Co n s equ ent l y the c ar til a ge b o n es O f the s ku ll formed in the c h o nd ro

, ,

c ranium ar e l ikew i s e l imit ed to the vent ral and ex t reme po s t eri or par t s o f the sku ll wh er e
, ,

th ey form a oor fo r the b ra in T he roof a nd s id es O f the skull a re c ompl et ed by memb ran e

.
 104 L AB OR A OR T Y MANUAL F OR VE R E BRA E ANA OM T T T Y
bon es ,
d eriv ed , as we h a v e a l r eady seen , f r om the d erm al s ca l es o f the g an o id shes . In a

si mil ar way c er ta in c ar til a g e bo n es a r is e in the m an dib u l ar an d hyoid arc hes an d beco me

s hea thedin m emb a e b o es The compl ete v e teb a te s kell the efo e con sis ts of c a til ag e
r n n . r r ,
r r , r

bon es d e ived f o m the cho d oc an i m an d its s en s e c ap sul es a d the s t an d s eco d g il l

r r n r r u ,
n r n

a r ches an d o f memb a e b o es c o v erin g the ca til ag e bo es ev erywher e e c ep t o n the v en tr al

,
r n n r n x

s u rf a ce an d p o s t e io r e d o f the s k u ll W e m ay n ow s ta te in d etail the bo es derived in thes e

r n . n

ways to f o m the s k u l l r .

I Car til g b
. s d r iv d fr om the p e
a e an d p cho d al s
on e Ther e a e four grou p s o f
e e r ar a r . r

su ch car til ag e b on es : on e occipital t w o s phen oid a d o e ethm id g r ou p ( s e F ig 3 , ,

n n o e .

a ) Oc ipit l gr u p: This c o s is ts o f f ou r o c cip ital c a o n

b o s s itu ted a t the po s te ior en d o f the s ku ll ne a r

,

fo rmin g a rin g a ou n d the fo amen magn u m They r r .

ar e on e su pr a occipital ,
t wo ex occi pital s , an d on e

b as ioccipital They a rise f o m the p a acho r dal car

. r r

til ag s t og ether with their d o r sal e ten sio the

e , x n

sy o tic t ec tu m
n an d the ve t eb al el emen ts which r r

a e fu sed t o the po s terio r en d o f the p ar acho r d al s

r

( Fig .

b ) P os ter ior s phen oid gr ou p: This co m pris es

t hree s phen oid bo n es , on the v en tr al s id e o f the
s kull , in fron t of the basio ccip ital . They a e r the
median un p a ir e d b as isphen oid an d the p aired al i

s phen oids ,
on e on ea c h s id e ex t en din g u p into the
back wall of the o bit Thes e b o es come from the r . n

po s te ior p a t o f the p echord al s r r r .

c) A ter ior s phen id g ou p: This in cl u des three n o r

bo n es on the ve tral s ide an te io r to the p ec edin g n r r

an d al s o d erived f r o m the p r echo d a l c ar til ag es r .

Th ey are a m edian u n p air ed pr s phen id an d p aired e o

or b itos pheu oids o n e o n each s id e f o rmin g p ar t o f

F G 3 5 Di g m t
I . h w th
.
_

a ra o S o e ar _ ,

t h W all s O f the o rb it
g m t f th til g b e
ran e en o f th e c ar a e o n es o e

s k ll
u ,
f m b
s een Th p t
ro d) E thm id gr ou p: This g r ou p c on s is ts o f three
a o v e. e s e ara e o

o ti b oc whi h m y b
n es , m y c bo es at the a te ior e d o f the ve tr al s ide o f the
a e as an as ve n n r n n

in mb
nu t p
er , t d Th
are s k ul l j u s t b ehi d a n d f u s ed with the o l f a c to ry ca p
no re resen e . e ,
n

S UP iPit 1 f m
I aOCC $ 1 b V th
al s ul es
or T he b o es a e a m edian mes ethmoid a n d
s 3 11 0 1 a O e e . n r

b i iPit l ( M d i d f m Ki g I y p aired ectethmoids o e o n each s ide o f the p rec edin g

aS OCC a O E ro n S e S
,
n .

C mp ti A t my f v t b t )
T he m esethmo id f o rm s a m edian p ar tition o r s ep
o ar a ve na o o er e r a es

tu m b etween the two ol f a c to ry c a p s l es whil e the u ,

e c te thm o id s c o n tribu te to the p o s terio r wall s o f thes e c a p su l es T he ethm o id s aris e in the .

ethmo id p l a te which r ep r ese t s the fu s ed a t e io r en d s o f the p r echo r dal s

,
n n r .

2. Car til a g e b on e s d erive d fr om the s en s e cap s u l e s .

a ) F r om the otic or au ditor y ca p su l es : A nu mb er o f o tic bon es a r is e in t he wal l s of the

o tic ca p su l es r are fu sed with the p arachordal r eg ion of the sk ull the o tic
. S in c e the l a t te ,

bon es ar e n atu all y cl o sely a ssociated with the o c cipital g ou p of bo es an d o f ten fu sed with
r r n

them The e m ay b e a s ma y as ve o tic bo es ( in t el eo s ts) the pr otic epiotic opisth tic

. r n n ,
o
'

, ,
o ,

p ter ti c aon d s phen,

otic b u t th e y a e co mm o nl y f u s ed t og,
eth er o r f u s ed with n ea rby bo n es
r .

T he thr ee s t n am ed a e the o n es m o s t c on s tan t in the higher v er tebra tes

r r When fu sed .

in to o n e b on e th ey a r e d es ign a ted a s the per i tic o r petr om s toid b o n e o a .

6 ) F r om the optic cap l es : As al ea dy exp l ain ed the o p tic cap su l es d o n o t fu se with the
su r ,

skul l o win g t o the n ec ess ity fo r r etain in g f r ee m ovem en t in the eyes

,
T he o p tic ca p s u l e .
 106 L AB ORA OR T Y MANUAL F OR VE R TE B RATE ANAT OM Y

dermos up raoccipital

pre f
ro n tal

s up ra te mporal C 0
.

FI G .
36 .
-
Dorsal view o f the s kull s of four r epr esen tative vert eb ra tes t o sh w t he r eduction o f the o

memb r an e b o n es o f the roo f in the c ou rs e o f ev ol u tio n A skull o fa n e tin c t mphib ian Capitos au r u s
.
,
x a , ,

b el on gin g t o the S tegocephal a; n o te the l ar g e n um b e r o f m em b ra n e b o n es c o mpl tel y r oo n g the sk ulle .

B , sku ll o f o n e o f the mo s t a n cien t ptil

re S eymou r ia , b el o n gin g t o t he Cotyl os aur ia; the mem b ran e
es ,

b o n es a re n ea rl y a s n u m er o u s p
a s in t he am hib ia n , a r e s im il arl y a rran g ed , a n d co m l et el y r oo f the p
s kull . C, s kul l o f a mo der n r ep p
til e , t he allig a tor ; s ev er a l o f the m emb r an e b o n es r es en t in the ext in ct
fo rms have b een l o s t , an d the r oo f b ea rs s ev eral o p en in g s . D, s ku ll of a mo d ern m ammal , the d o g ,
sho win g g r ea t er l o s s o f m em b ra n e b o n es
s t ill M emb ran e b o n es b l an ;
. k ca r til ag e b o n es s ti ppl ed .

(A f r om R eyn ol d s Th

e Ver tebr a te S kel eto n , co u r t esy o f the M acmil l an Com p an y ; B from Will is ton
s

Water Reptil es f the P as t an d P res en t

o ,
U
niversity o f Chica g o P r es s .
)
 T HE E NDOS KE L E T ON S K ULL AND VISCE RAL S KELE T ON 10 7

5 M e mb r an e
. b on e s a dd e d to the ven tr a l s ur ac e f of the s ku ll . Thes e a re from the
a n ter io r en d ,p os terio ly chiey : c mers pal tin es pteryg ids ( Fig
r , I t shou l d b e s ta ted
o , a , o .

ag a in tha t p a l a t in es an d p t eryg o id s a e m emb ra e b o es in som e an imal s a d ca r til a e b on es

r n n n
g
i o the s
n T he par s phe id is a m emb an e bon e which in Amphibia typ icall y fo rms the
r . a no r

chie f b o e on the v en t ral s u rf a c e o f the s k u ll b t it d o es n o t p e s i t in hi h er f o r ms

n Th u r s
g es e , .

m emb ra e bo es are s itu a ted ven tr al to the ethmo id a d p ar t o f the s phen o id bo es which
n n n n ,

they c on c eal f om su r fac e view b u t the p o s te io r p ar t o f the ven tr al s id e o f the s k u ll is c o m

r , r

p os ed o f car til ag e b on es s phen oid s an d o ccip ital s , .

6 . M em b r an e b on e s a dd e d to the j aw s , an d o ther g il l ar che s .

a ) Upper jaw: T he u pp er jaw (p tery g o qu ad r a te c a r til ag es ) eco mes in cased in mem b b ran e
bo n es . T he hief
c o n es fro m the tip of the jaw p os te io ly r r ar e , o n eac h s ide : pr emaxill a ,
maxill a , ju gal ( mal ar ) , qu adr a toju gal , an d
s qu a mos al ( s ee F ig T he u pp er jaw in .

el a s mo b r an chs is s ep ar a t e f r o m the chon

m
d ro cran iu m , t o which it is g en erally a t
terna l ares

t ached by l ig a m en ts , b u t in a l l o f the l an d
b
v er te r a tes the u pp er j aw is in sep ar ably
fu sed to the v en t r a l s id e o f t he c hon d r oc ra inc

n ium an d b ecom es an in teg ra l p ar t of the

0

p terygoid
s ku H .

b ) L ower jaw: T he l ow e r jaw ( Meckel s

c ar til ag es ) is s imil a r l y shea th ed in mem

lugal
b r an ebo n es M eck el s car til ag e o f ten p er
,

s is tin g within th em The chief m emb r an e .

b on es o f the l ower jaw a re the den tary the ,

spl en ial , the an gu l ar , the s u ran gu l ar ( al so quadratojugal

g iven as s u ra
p a n gu l ar ) , the cor on oid, the
gon ial , an d two or thr ee others oc cu rr i g n

o nl y in ex tin c t forms . In the evol u tion o f FI G 3 7 . .

Diagram of ca rt il a g e a n d m em
t he l owe r jaw ther e ha s b een a c o n tin u al b ra n e b o n es of the u pp er ja w an d oor of the
d ecr eas e in the nu mb er of memb r an e bo n es skull . M emb ran e b o n es b l ank ; c ar til ag e b o n es

( s ee Fig .
s ti ppl ed . T he ca r til ag e b on es me from the p t ery
co

Other gill ar ches : No m emb ran e g o qu a d r ate car til a g e (M odied from Kin g sl ey s

c ) .

Compar ative An a tomy f Ver teb r ates )

bo n es o c c u r in co n n ec tio n with the hyo id
o .

o r o the r g ill a ches r .

With this ac cou n t o f the co mp osition o f th s ku ll in e min d we m ay n ow tu rn to the s tu d y

o f s o m e vert eb r a te s k u ll s .

E . S KU LL OF N E CTUR U S A ,
P AR TIALLY O SS IF I E D S KU LL
sku ll of Amphib ia the ossi c a tion of the c hondro cranium has p ro
I n t he
ceed ed to b u t a limi t ed ex t en t so tha t a p ar tially c ar tila gino u s c hondro c rani um

is p resen t from whic h t he in casing m em bran e bon es can be readily sep ara ted as
in shes The p terygoqu adra te car tilages are i n separably fu sed to the ven tral
.

a n d la t eral sides of t he sk u ll p ro p er a n d are p ar tiall y o ssi ed I n the lower .

jaws M eckel s c ar tilages are p ersis ten t as c ores i n closed by m em bran e bon es

.

The num ber of m em bra n e bo n es in the sku lls of p r esen t day Amphib ia is con -

siderably less than tha t o i ex tin c t form s su ch as shown in F igu re 3 6 A ,

.
108 LAB O RAT O RY M ANUAL FOR VE RT E B RAT E ANAT OMY

of the sku ll of Nectu ru s c om p le te sku lls p r eferably p res er ved

F o r the s tu dy , ,

in uid an d c hon dro c ra nia from whi ch the m em b rane bon es have been rem oved
, , ,

shou l d be a t han d .

I .

G eneral re gion s of the s ku ll The sku ll is p ar tly bony p ar tly car til agi .
,

no u s The bon y p ar t ex is ts in the form of dis tin c t areas or bo n es sep ara t ed from
.
,

FI G L o wer j aws f
o f ou r v e rteb ra t es show t he redu c tio n in t he mb er of b o n es in the
.
38 . to nu

cou r s e o f evol u tio n . A , l ower jaw o f a n ext in c t a p

m hib ian , Tr imer or hachis , b el on gin g t o the S tego
phal a in n er su r fa ce a b o v e, o u t er f b el o w ; n o te l ar g e n u mb er o f m emb ran e b o n es B , l o wer
:
ce , s u r ace .

jaw ofan ex tin ct r e ptil e, La bi dosau r us , b el on gin g t o the Cotyl os aur ia; in n er su r a ce ab ove, o u t er su r ac e f f
b el o w ; n o t e redu c t io n in the n u mb er o f m emb ran e b o n es an d in cr ea sed s ize o f the d en tary h an d the
spl en ial i . C, l ower jawo f a mod ern re p til e, a liz a r d , Varan u s , showin g s till far ther r ed u ctio n in the
nu mb er o f b o n es ; o u ter su rface ab o v e, inn er sur fa ce b el ow D, hal f o f the l o wer jaw o f man , seen .

from the ou ter su rface ; it con sis ts o fb u t on e b o n e, the d en ta ry, all o ther b on es havin g van ished Mem .

b ran e b o n es b l ank ; car til a ge b o n es s tip l ed p a , p r eco r on oid ; b , in t er coron oid ; c , o s t s l enial ; d , coro
. p p
n oid ; e, articu l ar ; f ,
p p
a n gul a r ; g , p rear t icu l a r ; h, d en tary ; i , s l eni al ; j , su ra an gu l a r o r su ran gul ar ; -

l, c o r on oid p r ocess ; m, c on dyl o id p roces s ; n ,

b o dy ; p, m en ta l fo ram en
mu s ;
ra 0, (A an d B from .

W ill is ton W ater of the P as t a nd P r es en t, Un iv ers ity o f Ch i ca go P ress ; C from R eyn ol ds The

R eptil es
'

Ver tebr ate S kel eton , c ou r t esy of the M a cmill an Com p an y ; D from a s p ecimen l oan ed b y the ana t o my
d epartm en t ) .

ea ch o ther alo n g wavy or jagged lines the s u tu res The m embran e bon es are , .

som ewha t dis tin gui shable from t he c ar tila g e bones by their m ore s up er cial
p osi tions The sk ul l is di visible
. i n t o a m edi a n
p or tio n t he s k u ll ro
p p er a nd , ,
I 10 LAB ORA OR T Y M ANUAL F OR VE R E B RA E ANA OM T T T Y

wi th the p arie tal I n fron t of the dorsal p or tio n of the opis tho ti c is an area
.

of car tilage an d in fro n t of thi s is the p ro oti c bo n e

,
x t en din g fr o m the o is th
p .

o ti c obliqu ely fo r ward an d la terally is a slen der mem rane bo n e the s qu amos al , .

The p os t erior p ar t of the squ am osal c overs a p ar t of the op is tho ti c an d lies

above the la teral border of the o ti c cap su le The p ro oti c bon e is wed ged in .

b etw een the squ am osal and the p arie tal On tu rnin g the sku ll la terally so as .

to ob tain a la t era l view of the o ti c c ap s u le a sm all ro u n ded bo n e w ill be fo u n d

si tu a t ed in the la teral wall of the o ti c cap su le I t arti cu la tes behind wi th the .

opis tho ti c an d b ea rs a dorsally p ro jec tin g p ro c ess w hich m ee ts a ven t rally direc ted
p ro c ess fro m th e m idd l e of the sq u a m osal T h i s ro u n ded bo n e is .c alled t h e

col u mell a and is believed to b e eq u ivalen t to the hyo m a n dib u lar bone of shes ,

tha t is the dorsal p or tion of the hyoid gill ar c h

.
,
The in ternal ear is lo ca t ed .

i n side of the p ro oti c bo n e an d the colum ella ts in to an o p eni n g in the p ro oti c

,

bo n e called the oval win dow or fen es tr a oval is The colum ella when the m iddle
,
.
,

ear app ears ( rs t in An u r a) beco m es a bone of the mi ddle ear and it or a t

, , ,

leas t tha t p or tion of it whi ch ts in to the fen es tra ovalis is p robably homologou s ,

wi th the s tapes of the m amm alian middl e ear The hom ologies of the middle .

ea r b on es are however in so m e do u b t ( see K pp 8 0

, , , .

3 B one s of the u pp er ja w
.
The u pp er jaw co mp osed ori gi n ally of the two
.
,

p t e r y g oq u adra t e c ar t i la g es is f u sed to ,
th e sk ul l ro
p p er and fo r m s it s la t e ral
p or t io n s I t.c o n sis t s in N ectu r u s
,
in p ar t of the p ersis t en t p or tio n s of the
,

p t ery g oq u adra t e c ar t ila g es in p ar t of t

,
h e c ar t ila g e bo n.es for m ed i n th e se
c ar tila g es a n d in , p ar t of m e m bra n e bo n es e n shea thin g and rep la cin g the
c ar tila g e .

a ) M emb r a n e b on es of th e u pper j a w: The an t erior tip of the jaw is form ed

of the pr emaxil l ae V shap ed bo n es the an gle of the V dire cted forward an d

,
-
,

c o n s ti tu ti n g the tip of the s n o u t On e lim b of the V is a slen der p m c es s si tu a ted

.

dorsally on top of the a n terior en d of the frontal bone The o ther lim b of the .

V is a t oo th beari n g p ro c e ss fo rm ing the m ar gin of the a n t erior en d of the sk u ll

-
.

P os terior to the p rem a xilla is the t oo th bea ring vomer already m en tion ed and
n o t regarded as a bo n e of the jaw P os t erior to the vom er is the pal a topterygoid
.

bon e the an te rior p or tion of w hi ch bears t ee th F rom the p ro jec ti n g a n gle to

,
.

whi ch the lower jaw ar ti cu la t es to the p os terior angle of the o ti c c aps u le ex ten ds
the sl en der s qu a mosal bo n e .

b ) Ca r til age b on es of the u pper jaw: The qu adr ate bo n e fo rm s the fossa in to
which the en d of the lower jaw ar ti cu la t es and ex ten ds p os teriorly from the fossa
for a shor t dis tan c e in co n ta c t wi th the in n er side of the squ am osal M edial to .

the q u adra t e bone is a c o n siderable area of c ar tila g e the qu a dr a te ca r til age , ,

rep resen tin g u n o ssi ed p or tio n s of the p t erygoqu adra te c ar tilage B esid es the .

qu adra te bon e a p or tio n of the p ala top terygoid bon e is likewise a c ar tilage bon e
,

o s si ed in the p t ery goq u adra t e c ar tila ge .

D raw dorsal an d ven tral views of the sk u ll ou tlin in g the bo n es a c c u ra tely ,

.
 T HE E NDO S KELE TON : S K ULL AND VIS CE R AL S KELE T ON I I "

4 . The chon d r oc r an iu m When . sk u ll of Nectur u s is so aked in warm

the
soap solu tion the m em bra n e bones ca n easily be lif ted off reveali n g the chondro
, ,

c ranium ben ea th The chondro c ra n i um ( in clu din g the olfa c tory and o ti c cap
.

su les) consis ts in larg e p ar t of car tilage wi th a few car tilage bo n es whi ch have ,

alrea dy been iden ti ed .

S tu dy p rep ared c hondro c rania The form of the chondro cranium is si milar
.

to tha t illu s t ra ted in F i gu re 3 2 p 9 7 The p os terior p ar t of the chondro cranium

, . .

c onsis ts c hiey of the ro u nded hollow o ti c c ap s u les ,

These are connec ted dorsally.

by a narrow ar ch of c ar tilage the syn otic tectu m and ven tra lly by the broader
, ,

b as al pl a te form ed by the fu sion of the two para chor dal s

,
The two exo ccip i tal .

bones are os si ed in the basal p la te F rom the an terior end of ea ch o ti c cap .

su le a slender cu rved p re chordal c ar tila ge ex tends forward B e tween these is .

a larg e sp a c e the b as icr an ial fen es tra whi ch in the c omp le te sku ll is covered
, ,

above by fron tals and p arie tals and below by the p arasp henoid Near the .

an terior end in fron t of the p oin t where in the comp le te sk u ll the p arasp h en oid
ends the pr echor dal s are fu sed a c ross to form the ethmoid pl ate already no ted
, , .

F rom the e thm oid p la te a slender p ro c ess c on ti n u es forward on ea ch side the ,

cor n u a or horn s of the p r echor d al s I n fron t of the o ti c cap su le is the q u adra te

.

bone and qu adra te car tilag e The la tter sends p ro c esses to the o ti c c ap su le an d
.

the p re c hordal I n the walls of the o ti c c ap su le o p is tho ti c p ro oti c and colu

.
, , ,

m ella may be iden tied an d their bou ndaries more clearly dis tin gu ished than
in the s tu dy of the en tire sku ll The olfa ctory cap su les are so deli ca te as to
.

b e los t in p rep ari n g the chondro c raniu m .

5 T. h e l ower ja w The . lower jaw c onsis t s of a p air of M e c kel s c ar tila g e s

u ni ted in fron t a n d shea thed for the g rea ter p ar t of their c o u rse in m em bra n e

bones The ou ter s u rfa c e of ea ch half of the lower jaw c onsis ts of the den tary
.
,

a memb rane bo n e beari n g tee th The inner su rfa c e is formed of two m em brane
.

bones the splen ial and the an gu l ar The fo rm er is a small bone si tu a ted a t
,
.

abou t the middle of the inner su rfa c e beari n g the las t grou p of t ee th c on sis ting , ,

of ve or six tee th The a n gu lar c overs the rem ain der of the in n er su rfa c e an d
.

p asses on t o t h e o u t er s u rfa c e a t t h e e x trem e p os t erior end of the jaw below the ,

p os t erior end of t h e den t ary The . ar ti cu la ti n g s u rfa c es of the lower jaw are

c omp osed of car tila ge w h i c h is the p os t erior end of M ec kel s car tila ge This
.
,

ar ti cu la tes wi th the qu adra te of the upp er jaw M e ckel s car tila ge ru ns almos t .

the en tire le n g th of the jaw c o n c ealed be twee n the den tary an d the a n gu lar It .

c an be revealed by rem ovin g these m em bra n e bones .

6 The re m ainin g g ill a r che s

.
The hyoid an d three su cce eding gill ar ches
.

are p resen t in Nectu ru s an d are alm os t comp le tely car tila gin o u s P reserved .

ma terial is n ec essary for their s tu dy The hyoid ar ch is a broad som ewha t

.

V sha ed ar t ila e si t a t ed j u s t p os t erior t o t h e lower jaw in th e oor of t h e

g u
-
p c

m o u th c avi ty On ea ch sid e it is divisible in to two car tila ges a sm all an terior

.

hyp yo h a l a n d a m u c h lar g e r p os t e rior ce r a to h y a l The th ird g ill ar c h
. ( rs t t ru e
 1 1 2 LAB ORA OR T Y MANUAL T
FOR VE R E B RAT E ANA OM T Y

ill be ri g ar ch ) is m ore elonga ted and slen der tha n the hyoid ar c h an d is like
-
g a n

wise divisible in to two p ie c es an a n t erior cer a tobr a n chial and a p os terior

, ,

sligh tly l onger epibr an chial B e tween the m edian ven tral p or tio n s of the
, .

hyoid and third ar ch is a triangu lar copu l a rep resen ting a b asibr an chial piec e .

The fo ur th and f th gill ar ches are shor t cu rved rods of car tila ge on ea ch side ,

c omp osed of epib ranc hia l s At the an t erior end of the ep ibran chial of the fo u r th
.

arch is a small cer atobran chial I n the m edi an ven tral line is a slend er bone.
,

the se c o n d b a s ib r an chial I t will be see n t ha t the gill ar ches of Nectu r u s are

.

red u c ed bo th in nu m ber an d as regards the p ie ces of whi ch they are composed

as comp ared wi th the gil l ar ches of the elasmobran chs F or the typ i cal condi .

t ion of the gill ar ches in elasm obran c hs see F i gu re 3 3 p a ge 1 0 0 a n d c omp are , ,

wi th the co n di tio n in Nectu ru s .

F . THE S KU LL OF THE AL L I G Y
AT OR , A T P I CAL MOD E N
R S KU L L

The skull of the alli ga tor is a n ea rly c omp le tely o ssi ed skull in whi ch the
variou s comp onen ts of which the sk u ll is comp osed are so closely kni t in to a
si n gle s tru ctu re as to b e in sep arable The sku ll of the alli ga tor is p robably as .

t yp i c al and g eneraliz ed a sk u ll as is to be fo u nd am on g livin g land ver tebra tes

a n d has fu r ther the advan t a g e of larg e siz e The s tu den t mu s t learn the bones .

of the skul l and jaw of the alli ga tor an d be able to dis ti n gu ish the c ar tilage from
the m em brane bo n es F i gu res of the alli ga tor sk u ll will be fo u n d in R p ages
.
,

2 52 2 54 2 56 2 6 1 ; P an d H p a ge 3 4 2 ; CNA Vol VIII a e 8 K a e

, , , , p g 45 ; p g 1 04 ,
.
, , .

1 .

G eneral r e gion s an d cavi tie s of the s ku ll Ob tai n a sku ll and s tu dy its .

s truc tu re I t is comp osed of a number of sep ara t e b on es c lo sely join ted to ea ch

.

o ther alon g som ewha t jagg ed li n es k n own as s u tu res The vario u s c omp o n en ts .

of which we learn ed the skul l is c o n s tru c ted are here m orp hologi cally in dis tin
g u is h a b l e fro m ea c h o ther b u t will be p oin t ed ou t la t er The bones of th e roof .

of the sk u ll are p i tted a n d rou ghen ed At the an terior end of the dorsal su rfa ce .

are the two nasal Op enings or an terior n ar es P os terior to the mi ddle of the .

roof are the two large or b its the larges t op enin gs in the roof of the sku ll P o s
,
.

ter io r to ea ch orbi t is the tempor al region in whi ch are several holes k n own as ,

fossae or vacu ities The developm en t of these temp oral fossae is chara c teris ti c
.

of rep tiles and has o ccu rred du ring their evolu tion sin c e the early r ep tiles as , ,

in Fi gu re 3 6 B p ossessed comp le tely roofed sk u lls ( S ee fu r ther on this p oin t

,
.
,

R pp 2 8 5 8 7 an d F ig 44 p
, .
-
,
The alli ga tor sku ll has two p airs of these
.
,
.

,

op enin gs a dorsal p air the su pratemporal fossae on the dorsal side of the po s ,

terior en d of the sk u ll a n d a la tera l p air the infr atempor a l fossae j u s t p os terior ,

to the orbi ts from which ea ch is sep ara t ed by a rod of bo n e the pos tor bital b ar ,
.

L a teral to ea ch su p ra t em p oral fossa is a p roje c ti n g led ge of bo n e the su pr a ,

tempora l a rcade Un dern ea th this overhan gin g led g e is an op en in g the ex ter n a l

.
,

au ditor y mea tu s whi c h leads i n to the c avi ty of the m iddle ear or tympan ic ca vity
,
.

Num ero u s c a n als a n d p assages will be fo u n d en t ering the tymp a ni c c a vi ty an d ,

I I 4 LAB ORA OR T Y MANUAL T
F OR VE R E BRA E ANA OM T T Y

e h maxill ae the large bones a t the sides of the n asal bon es and also beari n g
t et ; , ,

t ee th ; ju ga l or ma l ar the elo n g a ted bo n es fo rmin the lower bo u n dari es of the

,
g
orbi t an d m ee ti n g the p os tfron tal s at the middle of the p os torbi tal bar ; an d
d a toju gal a slen der obliq u ely la ed bo e form i th os t erior bo daries
q ua r , p ,c n n g e p u n

of the la teral t emp oral fossae The qu adr a te is a s tou t bone obliqu ely p la c ed
.

be tween the qu adra toj u gal and the ex o ccipi tal The p os terior en d of the q u ad .

ra t e has a con c ave s u rfa c e for ar ti cu la tion wi th the lower jaw .

Tu rn the sk ul l over a n d id en tify the sam e bones of the m axi l lary ar ch from
below P remaxillae and m axillae fo rm the an t erior half of the ven tral su rfa c e
.
,

mee tin g in the m edian li n e B e twee n the two p rem axillae is the sm all an terior
.

p ala t i n e va c u i ty P os t erior t o
. th e m a xillae in the m edia n re gio n a n d formi n
g
th i n ner bo u n daries of the p os t erior p ala tine va c u i ti es are the pa l a tin es P
e os .

terior to the pala ti n es are the broad pter ygoids whi ch in close the p os t erior nares .

E x ten di n g fro m the sides of the p t eryg oids to the p os terior end of the maxilla e
are the ectopterygoids or tran s pal atines .

All of these bo n es of the u pp er jaw ex c ep t the q u adra te are m em bra n e bon es .

The q u adra te rep resen ts the o ssi ed p os t erior end of the p terygoqu adra te c ar ti
lage The qu adra te ar ti cu la tes wi th the lower jaw as in the m ajori ty of
.
,

ver tebra tes .

4 T.he o cc i pital r e gio n Th i s re g io n for m

. s t he p os t erior e n d of th e sk u ll
an d c o n sis t s of fo u r c ar tilag e bo n es derived fro m the p a r a chordal c ar tila g es .

Tu rn the sk u ll so tha t its p os terior end fa c es you The foram en m a gn u m is .

bou nded by three bon es one on ea ch side the exoccipital s an d on e below the
, , , ,

b a s ioccipital The basio c cip i tal bears m os t of the large rou n ded o ccipi tal c o n dyle
.

b u t bo th exo ccipital s send down p ro c esses whi ch t ake p ar t in the c ond yle Above .

a n d be tween the two exo cc ip ita l s is the t rian gu lar s u pr aoccipita l whi c h ar ti cu la tes ,

wi th the p os terior end of the p arie tal .

5 The b one s of the oti c caps ul e

.
There are three of these su rrou n din g the
.

ex tern al au di tory m ea tu s an d inc losing the ear They are nam ed the epiotic .
,

the opis thotic and the pr ootic The rs t of these is fu sed wi th the su p rao ccipi tal ;
'

.
,

the se c o n d wi th the e x o c ci p i tal ; the third alo n e rem ai n s di s ti n c t in the ad u l t .

They are dif cu l t or imp ossible to see in ex ternal view To nd the p ro oti c .
,

tu rn the sk u ll sidewise so as to ob tai n a p role view and ide n tify a larg e foram en , ,

the for a men ova l e j u s t below the su p ra temp oral fossa

,
The p os terior wall of .

thi s fora m e n is form ed by the p ro oti c bo n e whi ch ar ti cu la t es wi th the q u adra t e

,

b o n e b ehi n d the s u tu re be tween the two o ccu rri n g at the e x t ernal rim of the
,

foram en The o ti c bo n es are of c ou rse c ar tilage bon es fo rm ed in the o ti c c ap s u le

.
, .

6 The p o s terior s phenoi d r e gio n

.
This region is an terior to the o ccip i tal
.

regio n an d form s the oor of the c avi ty o ccu p ied by the brai n Tu rn the p o s .

terior end of the sk ul l t oward you an d ide n tify the V shap ed en d of the -

b as isphen oid b e tw een the basio cci p i tal and the p t ery goids
,
Mos t of the basi .

sphe n oid is c o n c eal ed from view by the p terygoids an d the q u adra tes b u t its ,
 T HE E NDO S KELE T ON : S KULL AND VI S CE RAL S KE LE TON 1 1 5

an terior en dp roj ec ts as a thin blu n t p ro c ess , the r os tr u m, in t o the sp a c e be twee n

the two orbi t s F i n d it in side view of the sku ll Above the ros trum on ea ch
. .

side is an al isphen oid bone ex ten ding u pward to the roof of the skul l and formin g
,

p ar t of t he walls of the fora m e n ovale and the supra temp oral fossa I n the .

in terior of the sku ll is a c avit y ex ten din g fro m the foram en magnum to the ali
sp hen oid bones ; this cavi ty con tai n s the brain in life and the ven tral su rfa ce of ,

the brai n therefore res ts u p on the o cc ipi tal and sp henoid bones derived from the ,

p a r a c hor d a l s a n d p r e c hor d al s .

7 The
. an t erio r an d ethm oid r eg i on s
s phe n oi d These region s remain in the.

car tila ge s tag e in the alliga tor fo rmin g a car tilaginou s p ar ti tion be tween the
,

two nasal p assa g es an d be tween t he orbi ts The c ar tilage is of cou rse wan tin g
.

in the dried sk u ll so tha t the nasal p assa ges and orbi ts app ear to be conn e ct ed .

Draw a ven tral view of the sku ll showing the su tu res a ccu ra tely , .

8 The l ower jaw

. The lower jaw or m andible is c omp osed of two halves o r
.
-

r a mi u ni t ed in fro n t by a symph ys is E a ch ramu s c onsis ts of six sep ara t e bones

. .

Near the p os t erior en d of the jaw is a large va cu i t y the extern al man dibu l ar ,
.

f or a m en ,
a n d in fro n t of t hi s on the inner side a s m aller in tern a l man dib u l ar
f or a m en . The bo n es of ea ch ram u s are : the den ta ry bearin g t ee th and form in
g ,

the o u t er s u rfa c e of the a n t erior two thirds of the ramu s ; the s pl en ial of abo u t
-
,

the sam e shap e a n d siz e as the de n tary and in the sam e p osi tion on the inner
su rfa ce ; the an gu l ar below the ex ternal ma n dib u lar foram en an d sep ara ted
, ,

from the sp len ial b y the i n t ern al mandib u lar foram en ; the su pr a an gu l ar or -

su r an gu l ar above the ex t ern al m a n dib u lar fora m en ; the coron oid a sm all bone
, ,

o n the in n er s u rfa c e be tw een the an t erior e n ds of the an gu lar an d the su p ra

,

an gu lar ; an d the ar ticu l ar above the pos terior en d of the a n gu lar and bearing
,

a con cavi ty for ar ti cu la tion wi th the qu adra te A ca vi ty exis t s in the in terior .

of the ramu s be twe en the den tary an d the sp lenial This c avi ty is o ccu pied in .

life by M e ckel s car tilag e w hi ch it may be recalled is the ori gi n al lower half of

,

the r s t gill ar c h The p os terior en d of this car tilage has ossied i n to the ar ti cu lar
.
,

whi ch is therefore the o n ly c ar tila ge bo n e in the lower jaw The arti cu la tion .

be tween u pp er a n d lower jaw is by way of the ar ti cu lar an d the qu adra te a ,

co n di tion fo u n d in t he m ajori t y of land ver t ebra t es D raw the lower jaw from .

bo th in n er an d ou ter vi ews .

Th h oi d ara tu s
A s has already been e xp lained t his is derived
9. e y app .
,

from the hyoid ar ch a n d rem ainin g gill ar ches The hyoid app ara tu s is gen erally .

missi n g o n dried skele to n s and for its s tu dy p reserved s p e c im en s are ne c essary

,
.

I t c o n sis t s of a broad c ar tila gino u s p la te the b ody of the hyoid and a p air of
, ,

ro esses or ho r n s ( c or n u a ) ex t e n din g p os teriorly an d dorsally from the body

p c ,

o n e o n ei th er side T he cor n u a are p ar tially o ss ied

. The body of the hyoid .

app ara tu s is derived from the bases of the se c on d and o ther ar ches while the
horns are rem n an ts of the thi rd ar ch I n lieu of alli ga tor ma terial the hyoid
. ,

app a ra tu s m ay also b e s tu died o n tu r tle skele ton s I n t he tu r tles there is a .

 1 1 6 L AB ORA OR T Y MANUAL T
F OR VE R E B R A E AN AT OMY T

m edian ven tral p la te the b ody of the hyoid lo ca ted in the oor of the m o u th
, ,

it r ep res en t s t he v en t ral e n ds of t he hyoid an d o ther g ill ar c h es F ro m it p r o .

j p
e c t os t eri o rly t w o p airs of sl en der p ro c esses a wn as t h e a n ter ior a n d, p ter ior
os

hor n: of t he hyoid Th ese are p or tion s of the thi rd and fou r th gill ar c hes
. .

I O The t ee th
.
The t ee th of the alli ga t or o ccu r in a si n gle row on the
.

p re m a xillae m a x,
illae a n d de n tary bo n es ,
Their s t ru c tu re is t he sa
. m e as t ha t
already d es c ribed u n der the exoskele ton The p u lp c avi ty of these t ee th is .

widely op en a t the base and the t ee th c a n t h erefore be r ep la ced an ind en i te

,

nu m ber of tim es ea ch su cc essive t oot h havi n g the sam e p u lp cavi ty as it s

,

alisp hen md

s u p ra oc c ip ital

orb itosphen oi d
ib u lar )

b as xsphen ond

pal atine

FI G 3 9
. Diagram o f the b o n es o f the mammalian
.
-
s ku ll . Memb ran e b on es b lank ; cartil a g e

pp
b o n es s ti l ed (After Kin g sl ey s Comparative Anatomy
.

o f Ver teb r ates ,
py ight
co r by P . B l akis to n

s

S on an d Com p an y . )

p rede c essor The t

. ee th are set in t o s ockets or a l veol i in t he jaw ; t ee t h so p la c e d
are sa id t o be thecodon t The tee th are all alike in siz e and form ; henc e they ar e
.

said to be homodon t .

G . TH E B ONE S OF TH E MAW AL IAN S KU LL

The m amm alian skull is c omp le tely o s si ed wi th the ex c ep tion of a sm all
p ar t of t he e t hm oid re gion The n u m ber of bo n es whi.c h it c o n t ai n s is however , ,

c o n siderably less t han tha t fo u n d in rep tile sk u lls This is du e in p ar t t o a loss .

of bon es an d in p ar t to a f u sio n of p ersis tin g bon es This fu sio n is m os t m arked .

in the hu m an sku ll Thu s as Willis ton says The mos t p rimi tive rep tiles had
.
, ,

no less than seven ty two sep arat e bo n es in the sku ll ; the hu m an skul l has b u t
-

t wen ty ei gh t i n c l u sive of the ear bon es The a cc omp a n yi n g dia gram of

-

.

the m am m alian sk u ll F i gu re 3 9 sho u ld b e s tu died carefu lly a n d will aid in

, ,

a c qu iri n g an u n ders tan di n g of the c o n s t ru c tion of t he m amm alian sku ll The .

 1 1 8 LABORA OR T Y MANUAL FOR VE R TE B RA E ANA OM T T Y

tu be The m ea tu s leads in to the cavi ty Of the b u lla which is the tympa n ic cavity
.
,

or cavi ty O f the m iddl e ea r F rom the dorsal side of the b u lla a ridge b e gin s
.

whi ch p ro c eeds a c ross the p os terior p ar t Of the sku ll to the o th er b u lla ; this
rid ge is the su perior n u cha l l in e ( or l a mbdoidal r idge) an d is the m os t an terior
p oi n t O f a tt a c h m e n t of the m u s c les O f the ver t ebral c ol u m n F ro m t he m iddl e
.

of this ridge there p roje cts p os teriorly the extern al occipital pr otu b eran ce sligh t ,

in the c a t b u t form i n g a re c ta n gu lar p ro m i n en c e in the rabbi t .

On the ven t ral su rfa c e the a n t erior p ar t O f the sku ll is o c cu p ied by the hard
p a l a te .Dorsal t o th i s are t h e nasal p assa g es whi c h o p e n a t the p os t erior en d Of

the hard p ala te by the pos ter ior n a res or choan a e The hard p ala te c on tain s a
.

p air O f O p e n i n g s th e,
i n cis iv e fora m ina or a n ter ior pa l a tin e foramina ; these are

sm all an d a t the an terior ex tremi ty Of the p ala t e in the ca t b u t mu ch longer an d

m ore p rom in en t in the rabbi t They lead i n to the nasal c avi ties At the
. .

p os t erior e n d of th e z y go m a ti c ar c h o n its v en t ral side is a dep ressio n the m a n ,

dibu l ar fos s a for the re c ep tion Of the lower jaw

,
M edial to this is the pterygoid
.

fo s s a for th e a tt a c h m en t O f c er t ain m u s c les The p.t ery g oid fossa in the cat is

c on ti n u ou s wi th t he t emp oral and orbi tal fossae while in the rabbi t it is i n c lu ded
,

be tween two p roje c ti n g p la tes of bon e whi ch p oin t toward the tym p an i c b u lla .

2 M e mbrane bo n e s of the r OOf Of the s ku 11 B e gi n nin g j u s t behind the

. .
-

an terior n ares are the p aired n as al s roong the nasal c avi ties ; ne x t pos terior
, ,

the p air ed fr on ta l s ; and las t the p aired pa r ieta l s term ina ti n g a t the s u p erior
, ,

nu chal lin e B e tween the p os terior ends of the p arie tals is gen erally a sm all
.

t ria n gu lar in terpar ietal bo n e the bo u n daries O f whi c h are n o t always di s tin c t
, .

E a ch fro n tal bo n e form s the dorsal p ar t O f the orbi t and p roje c ts ou t above the
orbi t as the su p raorbi tal ar ch which term ina t es p os teriorly in the z ygom a ti c
p ro c ess of th e fron t al bone ,
already no t ed I n th
. e an t erior wall O f th e orbi t is
the sm all l a cr imal bo n e (p robably hom olo go u s wi th the p refro n t al bone of
rep tiles) At the an terior end of the la c rim al bo n e is an op en in g the p os terior
.
,

en d Of the n as ol a cr imal du ct b y m ea n s o f whi c h the t ears drain in to the nas a l

,

c avi ty .

3 The b one s of the u pp er jaw

.
These as in the all iga tor c omp rise a la teral
.

m ax illary ar ch o n ea ch side a n d a m edia n series Of bones The m axillary ar ch

.

c o n sis ts o n ea c h side Of the followin g elem en t s : prema x il l a in fron t of the ,

an terior n ares and beari n g t ee th ; maxill a formi n g the side Of the fa cial regio n
,

O f the sk u ll a n d also beari n g t ee th ; mo l ar or j u gal fo rm i n g m os t O f the z y go m a ti c

,

ar ch ; and te mpor al comp le tin g the zygoma ti c ar ch and coverin g the side O f the
,

c ra n ial p art Of the sk u ll in c l u di n g the t ym p a n i c b u lla Fu r th er de tails co n c ern

.

in g these bo n es may be no t ed The p rem axillae sen d fr on tal pr ocesses dorsally

.

alo n gside the nasal bones ; these are very p rono u n c ed in the rabbi t The pr e .

m ax illae also form the an t erior p ar t O f the hard p a l a t e by m ea n s Of the ir pa l atin e

p r o ces s es whi c,h m e e t in t h e m edia n ven t ra l li n e and in clu de the i n cisive foram in a .

The m axilla is the m ai n bo n e of the fa cial regio n ; in the ra b bi t it is mu ch fen es

 T HE E NDOS KE L E TON : S K ULL AND VI S CE RAL S K E L E TON 1 1 9

tra ted . It form s p ar t Of the an terior wall of the orbi t ven tral to the la c rim al
,

bon e by its or b ital pr ocess ; it ex t en ds to the fron tal bone above b y its fr on ta l
,

p r oce ss ; it s p a l a ti n e p r o c es s m ee ts its fellow in the m edian ven tral line c o n tin u in g

the hard p ala t e ; its a l veol a r pr ocess bears t ee th ; a nd its zygoma tic proces s con
s titu tes the be gin ni n g of the z y go m a ti c ar c h . The malar or j u gal bon e is dis tin ct
in the ca t b u t in the ad u l t rabbi t is fu s ed to the z ygom a ti c p ro c ess O f the m axilla .

The t emp o r al bo n e is a c omp ou nd bone chara c teris ti c Of m ammals I t con sis ts .

O f a s qu amou s por tion which by its z y gom a ti c p ro c ess c o mp le t es t he z ygo m a tic

ar ch an d whi ch also con t rib u tes to the c ranial wall ven tral to the parie tal ; ,

O f the tympan ic b u l l a c o mp osed of a tym p a ni c bone Of u n c er tain hom olo y ; and

g
of the periotic or petr omas toid bone o n ly sligh tly visible on the su rfa ce an d con
sis tin g of the fu sed o ti c bon es O f lower ver tebra tes The s qu amou s p ar t of the
.

t em p oral bo n e is ho m olo go u s wi th the sq u am osal bo n e O f the lower ver tebra t es

a n d is the m os t p os t erior bo n e Of the m a xilla ry ar ch . Al l O f the bones O f the
m axilla r y ar ch are m em bra n e bones .

The m edi an p or tion of the upp er jaw forms p ar t Of the ven tral su rfa ce Of
the sk u ll . We have already no ted tha t the an t erior p or tion of the hard p ala te
is c omp osed Of the p ala tine p ro c esses Of the p rem axillae and m axillae P os terior .

t o the la tter are the pa l a tin es m em bra n e bo n es w h

,
i ch in c lu de the p os t erior nares .

D orsal to the p ala ti n es the roof of the nasal p assages is co mp le ted by the vomer
Of wh i ch o n ly the ex trem e p os t erior tip is visible in the roof Of the choa n ae .

The vom er form s the oor Ofthe n asal c avi ties an d is bes t seen in a sa gi ttal se c tion
of the sku ll The pos terior p art s Of the p ala ti n es ex t end u p in to the orbi t
.
,

there bei n g in the rabbi t a v e ry deep c lef t be tween them in the midven tral li n e .

Obs erve tha t the q u adra t e is wan ti n g . The lower jaw con sequ en tly ar ticu
la tes wi th the squ am osal ( tempo ral) by m ean s Of a dep ression the man dib u lar ,

fossa on the u n der su rfa c e of the z ygom a ti c p ro c es s of the t emp oral bone This
,
.

fea tu re dis ti n guishes m amm alian sku lls from those Of all o ther ver tebra tes .

Owin g to the absen c e O f the q u adra te all of the bo n es O f the u pp er jaw are m em
b ran e bo n es .

T h o i i tal re ion This re ion s u rro u n ds the fora m e n m a g n u m a n d

4
. e cc p g.
g
c o n sis t s O f a si n gle occipita l bo n e e x t e n di n g fro m the su p erior n u chal line to a

oi t be twe t h a t erior e ds f th t m a i b u llae I t is r eally c o mp osed

p n e n e n n O e y p n c .

O f the fo u r o cc i i tal bo n es p rese n t in rep tiles ; t hese are dis ti n c t in em bryo n i c

p
and you n g m am m als as shown in F i gu re 40 A p age 1 2 7 b u t are fu sed in the adu l ts
, , ,
.

The o ccip i tal bo n e bears the two o ccip i tal c o n dyles whose n um ber ser ves to dis
,

tin gu ish m amm alia n sk u lls from those O f all o ther ver tebra tes ex c ep t Amphib ia ,

wh ere th ere are also two c ondyles The dorsal p ar t Of the o ccip i tal m ay app ear
.

to ex t en d a n teriorly be twee n the p os terior e n ds O f the two p ari e tals ; this is

du e to the fa c t tha t in terp arie tal bo n e is c o mm o n ly fu sed to the o c cip i tal . This
is the c ase in m an . The o ccip i tal bon e bears the su p erior n u chal li n e an d the
j u gu lar p ro cess res ti n g o n the b ul la .
 1 20 LAB ORAT O RY MAN UAL FOR VE RTE B RA E ANA OM T T Y

5 The
. oti c cap s ul e s .
As already exp lained the bon es Of the o ti c c ap su les
,

are all fu sed to gether an d fu sed wi th the squ am osal an d the tym p ani c b ul la to
form the tem p oral bo n e a bon e very chara c teris ti c ofm amm als (Fig 40 C p 1
,
.
, .

T ha t par t of the t em p oral bo n es whi c h is c o mp ose d O f the o ti c bo n es is n am ed the

p e r io ti c or p etr o m a s to id bo n e The
. se p ara t e o t i c bo n es O f wh i c h th e p e t ro m as

toid is c om p os ed c an be se en o n ly in e arly em bryo n i c s ta g es The m as t oid p or tion .

O f this bo n e is visible o n the ex ternal su rfa c e O f the sk u ll be tw een the b u l la and

the o ccip i tal ; it p roje c t s over the b u lla as the m as toid p ro c ess v e ry p rom i n en t in ,

man as the b u m p behi n d the p in n a The p e trou s p or tion O fthe p etro m as toid bo n e
.

i n closes the i n tern al ear an d is visible on ly from the i n side O f the sku ll The .

t ymp a n i c b u lla c o n sis ts O f one or two m em bra n e bones of u n c er tain hom olo gy ;
it in c loses the middle ear an d the three m iddle ear bo n es These can be seen in .

well c lean ed c at sku lls as a deli ca t e chain Of bo n es ex ten di n g a c ross the an terior
-

p ar t O f th e t y m p a n i c c avi t y I f. sep ara t e s p e c im e n s are available id e n tify ,

t h em as follows : the mall eu s or hammer a slender bo n e t erm in a tin g in a k n ob ; ,

the in cu s or an vil a sm aller bone wi th two p oin ted p ro c esses ; and the s ta pes or
,

s tir r u p shap ed like a s tirru p

,
There is good reason to believe tha t the m alleu s
.

rep res en ts a very mu ch red u c ed art i cu lar ; tha t the in cu s is the red u c ed qu adra t e ;
an d the s t a p es is the hyom an dib u lar .

6 The pos terior s phenoi d r e g ion

.
On the ve n tral s u rfa c e of the sku ll in
.

fro n t Of the ventral p ar t O f the o cc ip i tal bon e is the b a s isphen oid The basi .

sp hen oid ex tends la t erally in fron t O f the tymp a n i c b u llae as p ro cesses the win gs ,

O f the basisp henoid . These win gs are in r eali ty the alis phen oids and are sep ara te
in yo u n g s tages ( see F ig 40 B ) . The alisphen oids m ee t the squ am ou s p ar t Of
.

the temp oral bo n es dorsally . The alisp hen oid s ex ten d forward as the pterygoid
p r oc e s s es whi c h m ee t the p os terior ends of the p ala ti n es I n the rabbi t the .

p t ery g oid p ro c ess p resen t s t w o ba ckwardly p roje c t i n g t hin p la t es O f bo n e t he ,

l a ter al an d media l l amel l ae . These i n c lose be tween them the p terygoid fossa .

The m edial lam ella in the rabbi t an d the p terygoid p ro cess i tself in the ca t bear
a p oin ted p ro cess the ha mu lu s The p tery goid p ro cesses c orresp o n d to the
, .

p t ery g oid bo n es O f lower for m s .

7 The an terior s phenoi d re g ion
. I n the m edian ven tral line in fron t of
.

the basisp he n oid is the pr esphen oid a sle n der bone ,

I n the rabbi t thi s is a t the
.

bo t tom of the deep clef t be tween the p ala tines The p resp hen oid sen ds up .

win gs i n to the orbi t whi c h m ee t the fro n t al bones above a n d c on t ain the large
Op ti c foram e n for the p assa ge O f the o ti c nerve I th t t his fora m en is
p n e c a .

the m os t a n terior O f a row O f fo u r foramina To see the win gs of the p resp hen oid
.

tu r n the sk u ll to O b tain a la t eral view The wings are in reali ty the or b itosphen oid
.

bo n es of lower ver tebra tes and are sep ara te in you n g s tages (Fig 40 B p 1 2 7) .
, .

Draw a ven tral view of the sku ll showin g the su tu res a ccu ra tely , .

8 The e thm oi d re g ion an d the sa gittal s e c tion Of the s ku 11 The e thm oid
. .

is nearly comp l etely o s s ied in mammals b u t can b e s tu di ed only in sa gi ttal

1 22 LAB OR A OR T Y MANUAL FOR VE R TE B RATE ANAT OM Y

The m os t p os terior h ese bo n es is the ethmotu r b in al or ethmoid l a b yr in th sit

Of t
u a t ed j u s t in fro n t O f the c ribriform p la t e I t is a gr ea tly folded s tru c tu re the
.
,

fo l ds jn c l o s in g sp a c es k n ow n as the ethmoid cel l s

I n fro n t of the e thm o tu rbin al .

is a n o th er b u t sm aller m ass the maxill otu r bin al born e o n the i n n er su rfa c e Of

, ,

the m a xilla .E a c h n asal bears on its i n n er su rfa c e a si n gle elo n ga t ed led ge Of

bo n e whi c h co n s ti tu tes the n as otu r b in al an d whi ch lies above the u pp erm os t
, ,

s crolls Of the e thm o tu rbi n als The fu n c tio n Of the tu rbin als is t o i n cr ease the
.

resp ira tor y a n d O lfa c tory s u rfa c e of the n ose The e thm o tu rbi n als are c overed .

by the Olfa c tory ep i theli u m while the m axill o tu rb in al s serve to s trai n an d m ois ten
,

the air The la tt er will be s een to p roje c t in to the p assage from the a n terior to
.

the p os t erior n ar es .

F rom the c om p ara tive s t a n dp oi n t the sep tu m Of the n ose is the m ese thm oid ,

the p erp e n di c u lar p la t e of the e thm oid bei n g its os si ed p or tio n whil e the e thm oid ,

labyrin ths are the ec tethm oid s The c ribriform p la te is p rodu c ed by the ex ten
.

sio n O f ossi c a tion p ro c esses be tween the m ese thm oid an d ec t ethm o id s The .

p erfora tio n s in th e p la t e are for th e p assa g e of th e O lfa c t ory n erve Th u s th e .

,

thre e e thm oid bo n es O f lower ver tebra tes are f u sed in t o the si n gle e thm oid bo n e
O f m am m als .

D raw the sagi t tal se c tion .

9 The foramina of the s ku ll
. The sku ll is p ier c ed by n u m erou s Op en l n gs for
.

the p assa ge O f n erves a n d blood vess els a n d so m e tim es o th e r s t ru c tu res These .

are lis t ed below for c onven ien t referen ce .

Cat :
a) fora
I n cis ive m ina A n t erior
. end O f ven tral side Of m axillae ; c onn e ct roof
of m ou th wi th n asal c avi ties .

b ) I nfr a or bita l foram en L arg e o p en i n g in the m axilla a t the be gi n n i n g of

.

the z y gom a ti c ar ch ; for the p assa g e of c er tai n bra n c hes O f the f th c ra ni al n erve
an d blood v essels .

c) Na s ol acr ima l c a n al
.

An t erior en d O i the la c rim al bo n e an d p assi n g

.

thro u gh the m a xilla in t o the n asal c avi ties ; for the drai n i n g O f the tears .

d) S phen opal a tin e foram en I n tha t p ar t O f the p ala ti n e bo n e whi ch ex te n ds

.

in to the orbi t p os terior t o the la c rim al ; for the p assage of bra n ch es of the f th
,

n erve i n t o the n asal c avi t y .

e) P os ter ior pa l a tin e c a n al The p os t erior en d Of this is imm edia t ely ven tral
.

t o the sp hen op ala ti n e foram e n ; its a n t erior en d is in abo u t the m iddl e O f the

p ala tine p ro c ess O f th e m a x illa ; for t he p assa g e of a bra n c h O f the f th n erve

to the p ala t e .

f) p O tic fora m en I n the orbi tosp hen oid p ar t of the p r esp h en oid in the
.
,

p os t erior p ar t Of the orbi t m os t a n t e rior O f a ro w O f fou r foram i n a : for the

,

p assa g e O f t h e o p ti c n e rve .

g ) O r bi ta l ss u re S e c o n
. d a n d larges t O f the row O f fo u r ; t hro u gh it p ass the
thi rd fo u r th a n d si x th n erves t o the m u s c les of the eyeball a n d a p ar t of the f th
, , , .
 T HE E NDOS KE L E TON : S K ULL AND VIS CE RAL S KE L E TON 1 2
3

h) F or amen r otu n du m . Third Of the row ; in the alisp hen oid bone ; trans
mi ts p ar t Of the f th nerve .

i) F or amen oval e . L as t Of the fo u r ; in the alisp hen oid ; t ra n sm i ts p ar t O f the

f th nerve .

j) C a n al for t h e E u s t a c hi an tu be I n the an terior wall of the b u lla its roof.

,

fo rmed by the alisp henoid ; for the p assage Of the E u s ta chian tu be from the
p hary n x i n to the b u lla .

k) P terygoid c anal E a ch b u lla term in a tes a n teriorly in a p oi n t ( s tylifo rrn

.

p ro c ess ) lyi n g on t h e basis p henoid ; t his p oi n t is dire c t ed t o a m in u t e o p enin g ,

the p teryg oid c a n al lyi n g in the s u tu re be twe en the basisp he n oid an d the p tery
,

g o id p ro c ess ; for t h e p assa g e of a bra n c h O f th e f th nerve in t o t h e b u lla .

l) J u gu l a r foram e n L arge foram en on the m edial side Of the p os terior en d

.

of the b u lla for the p assage Of the nin th ten th an d eleven th nerves
, , , .

m) Hypogl oss al foram en I n the m edial side Of the p re ce di n g foram en for

.

the p assage Of the twelf th n e rve .

n ) S tyl oma s toid foram e n At the ve ntral tip of the m as toid p ro c ess for the
.

p assa g e O f t h e seven t h nerve .

Rabbi t :
a ) I n cis ive forami na As in the c a t b u t larger
. .

b ) I nfr a b or b ital foram en As in the ca t b u t more sli tlike and elo n ga ted
.
,

formi n g an infr a or bital can al op en ing i n to the orbi t above the exp an ded p ar t
,

O f the m ax illa .

c) Nasol a cr imal c a n al As in the c at si tu a ted u n der the p oin ted an terior

.
,

en d of the s u p raorbi t al ar ch .

d) P os ter ior pal a tin e foram en On the ve n tral side in the s u tu re be tween
.
,

the p ala tine p ro c ess Of the m ax illa an d the p ala ti n es ; form s the a n terior o p en
in g of the pal atin e can al The p os terior op eni ng is a t the p os terior en d Of the
.

exp anded m ass Of the m axil la lo c a ted in the orbi t This c an al is for the p assage .

O f a bran c h of the f th n erve .

e) S ph en opa l a tin e fora m en In c omm on wi th the p os t erior en d of the

.

p ala ti n e c a n al j u s t des c ribed ; for the p assa ge Of bran c hes O f the f th nerve .

f) A n te r ior a n d p o s te r io r s u p r a or bi ta l fora m i n a The p roje c ti n g a n t erior

. a n d
p os t e rior e n d s of t h e s u p raorbi t al ar c h are c o n t i n u ed in li f e by li g a m e n t s th u s ,

fo rmi n g forami n a throu gh whi ch bran ches Of the f th nerve p ass

,
.

g ) O p tic fora m e n L ar
.
g e Op enin g in th e c e n t er O f th e orbi t for th e p assa g e ,

O f the o p ti c ne r ve .

h) Or b ital ss u r e P os t erior an d ven tral to the p re c edi n g an d in c lu din g

.
,

the foram en ro tu n d u m of o ther m amm als F o r the third fo u r th an d si x th .

, ,

n erves to the eyeball an d the g rea t er p ar t O f the f th

,
.

i) An ter ior middl e an d pos ter ior s phen oidal foram ina
, ,
Three forami n a in a .

row in the la teral lam ella of the p ter ygoid p ro c ess a t the p la c e wh ere this is
c o n ti n u o u s wi th the a li sp henoid F o r the p assa ge O f p ar t of the f th n e r v e
.
.
1 24 LAB ORA OR T Y MANUAL F OR VE RTE B RA E ANA OM T T Y

j) F or amen l acer u m fron t

. O f t he tym p ani c b u lla on the ve n t ral s ur fa ce
In .

a n d in cl u din g the foram en ovale of o ther m amm als ; for the p assa ge of p ar t O f

the f th nerve an d an ar te r y .

k) Exter n al car otid foram en A sm all foram e n in the middle of the m edial
.

su rfa ce of the b u lla ; for the p assage Of the in tern al caro tid ar tery the o ther en d ,

O f the c a n al lyi n g in the foram e n la c eru m .

l) J u gu l ar foramen Ju s t p os terior to the p re cedi n g in the dep ressio n

.
,

be tween o ccipi tal con dyle an d b u lla ; for the n i n th t en th an d eleven th c ranial , ,

n er ves an d a vein
,
.

m) Hypogl oss al canal I n c l u di n g sm all ap er tu res p os t erior to the p re c edi n g

.

for the p assa ge of the twelf th (hyp oglossal) n erve .

n ) S tyl oma s toid foram en I n fron t of the middle O f the m as toid p ro c ess for
.
,

the p assa ge of the seven th n erve .

10 .The lower jaw The lower jaw or man dibl e c on sis ts of a si n gle p air of
.

bon es the den tar ies fu sed in fron t by a symphysis All o ther bo n es seen in the
, ,
.

lower jaw Of the alli ga tor have van ished ( ex c ep t the ar ti cu lar whi ch is s u pp osed
to be the m alleu s of the m iddl e ear) The horiz o n tal p art Of the m an dible is
.

n am ed the b ody the ver ti c al p ar t the r amu s

, ( I n the lower
, ver t ebra t es
. ea c h
half Of the lower jaw is also nam ed ram u s ) The p os terior end Of the m an dible .

( )
ca t e x t ends dorsally in t o a s t ro n g c or on oid p ro c ess whi c h in t he n a tu ral p osi tio n ,

p roje c t s i n to the tem p oral fossa I n the rabbi t this is red u c ed to a sli gh t p r o jec
.

tio n whi c h form s the la t eral bo u ndary Of a deep g roove The ar ti cu la ting .

su rfa ce Of the m an dible is born e on the con dyl oid p ro c ess The dep ressed areas .

in the p os terior par t Of the m an dible are for the inser tion of the mu s cles O f
mas ti ca tion . Near the an terior tip Of the m an dible on the o u ter s u rfa c e is the
men tal foram e n ( or two in the ca t) thro u gh whi ch the nerve Of the lower jaw
exi t s Near the cau dal end of the inner s u rfa c e is the man dib u l ar fo ram en
.

thro u gh whi ch the n erve en ters and p u rs u es a c o u rse in the i n terior of the man
dible to the men tal foram en In the rabbi t there is an addi tion al fo ram en
.

j u s t above the m an dib u lar foram en for the p assa ge O f a vei n ,

.

Owin g to the abse n c e of the q u adra t e an d of all of the bo n es of the lower

jaw ex c ep t the den tary the ar ti cu la tion Of the low er jaw to the sk u ll is between
,

the den ta ry a n d the s qu a mos al This fea tu re dis ti n guishes m amm als from all
.

o ther ver tebra tes for in the la tter the ar ti cu la tion is be tween the ar ti cu lar and
,

the qu adra te . The co n di tion fou n d in m amm als is however app roa c hed by , ,

those rep tiles dire c tly a n ces tral to m a mm als .

1 1 .The tee th The tee th of m ammals p ossess several chara c teris ti c s :

.

they are s et in so cke ts or a l veol i in the jaws a c o n di tio n know n as thecodon t; ,

they are heter odon t tha t is di f ,

fere n tia ted in to several differe n t ki n ds ; they can
,

g e n erally be rep la c ed o n ly o n c e ; and so m e of the m are c o m li c a t ed in for m

p .

The tee th Of m amm als fu rn ish imp or tan t tax o n om i c c hara c t ers Those of the .

ca t and rabbi t ar e very differen t the fo rm er havi n g t ee th chara c te ris ti c O f

,
1 26 LAB ORAT ORY M ANUAL FOR VE RT E B RATE ANAT O M Y

12 . The hyoi d appar a tu s .

This is as already ex p lain ed the rem n an t O f the , ,

hyoid and o ther gill ar ches I t is ge n erally absen t o n p rep ared skele to n s an d
.

isola ted sp e cim ens will b e p rovided for its s tu dy .

The hyoid O f the cat c onsis t s of a bony bar p la c ed in the roo t Of the tongu e
j u s t in fro n t Of the lar yn x ; this bar is c alled the b ody O f the hyoid F rom it .

ex tend two p airs Of p ro c ess es or horn s an an te rior p air and a p os t erior p air , .

The an terior horn s c onsis t of fou r sep ara t e pie ces Of whi ch the term inal piece ,

is a tta ched to the tymp anic b u lla j u s t ven t ral to the s tylom as toid foram en
of the sku ll The groove whi ch it o c cup ies can g en erally be seen on the side of
.

the b u lla The p os terior ho rn s consis t ea c h Of a si n gle p ie c e whi ch is u ni ted to

.

the larynx The body and an terior horns belo n g to the hyoid arch ; the p os terior
.

horns to the third gill ar ch The rem ainin g gill arches are rep resen ted in the .

larynx whi ch will be s tu died la ter .

The hyoid app ara tu s of the r ab b it c onsis t s of a s to u t bon e the b ody Of the ,

hyoid si tu a ted a t the base Of the tongu e in fron t Of the laryn x I t bears two
,
.

p airs O f p ro c esses or h o r n s The an t erior horn is a shor

. t p ie c e c o n n e c t ed by a
mu s c le wi th the j u g ul ar p ro c ess of the sku ll The p os terior horn is a longer .

p ie c e c o n ne c t ed by li g a m en t wi t h th e la r yn x and by m u s c le wi t h th e j u g u lar
p ro c ess The body
. and an t erior horn are re m nan t s O ft he hyoid ar c h th e
p sp t er ior ,

horn of the thi rd gill ar ch .

H . G EN ER AL S UMMAR Y OF THE S KUL L AND VI SCE RA L S KE LE TON

1 . T he s kull b egin s as a car til ag in o u s c as e, the cho n d r o c r an iu m , hose m ethod O f
w
form ation has al r ead y been d es c ir bed . This ca se in cl u des the Ol f a c to ry an d o tic

ca p su l es .

2. T he g ill ar c es eco me cl os elh b

a ssoc ia t ed wit y
the chon droc r an iu m h . The e a er r

u s u all y h
seven o f t em , the r s t fo rmin g the u er an d l ower j aws , the s ec o n d the pp hyoid arch .

T he a r ches ar e r edu c ed in nu mb er an d al ter ed in fun c tio n in l an d ver tebr a tes .

3 . B on es d e ived r fro m the chon d ro cr an iu m an d g ill ar c hes ar e the car til ag e bon es of

the s ku ll .

4 . In addit ion to these the e r are memb ran e bon es a d d ed . They co me f ro m the d ermis
o f the s k in re o rig in ally d ermal s cal es
an d we .

T h ca r til ag e b on es o f the s kull an d se se ca p s u l es ar e the oc cip itals s phe o ids

5 e
. n ,
n ,

ethmo id s o tics an d tu rb in al s
, There ar e fou r o ccip ital s ( su p ra oc cip ital two e o ccip ital s
, .
,
x ,

basioccip ital ) ; a po s te ior g rou p o f three s phen o ids (basisphe o id two alisphe oids) ; an
r n ,
n

a terio gr ou p Of thr ee s p hen oid s ( p r es ph en o id two o b ito sphe o ids ) ; an d a g ro u p o f thr ee

n r ,
r n

ethmo id s ( mesethmoid two ec tethmo id s) T he o tic b on es o ss ied in the ear c ap su l e a e

, . r

gen er ally thr ee in n u mb er b u t m ay b e a s man y as ve ; in m ammal s they are fu sed t o fo m r

a p erio tic o r p etromas to id b on e T he n as al ca p su l es may fu rn ish additio n al c a til g e bon es

. r a .

The occ ip ital s an d s phen o id s fo rm the p o s terio r en d an d oo r o f the bra in ca vity whil e the ,

eth mo id s in clo se the n as al ca vities .

6 . The y g ives rise to two c a til ag e b o n es a qu ad ate derived from

rs t gill ar ch g en er a ll r ,
r

the po s terior en d of the u pp er jaw car til ag es o p teryg o qu ad a te c a til ag es ; an d an ar ticu l a rr r r

d erived from the p os terio r en d o f the l ower jaw ca r til ag es o r M eckel s ca r til ag es I n so me
.

fo rms the p terygo qu adrate car til ages may g ive ris e to p al a tin es an d p teryg o id s in addition .
 TH E E NDOS KE LE TON : S K ULL AND VIS CE RAL S KELE T ON 1 2 7

. y
7 T he s ec o n d o r h o id g ill arc g ives r ise to a c ar til ag e h bo e the hyoman dibul ar
n , , an d

the r r
ema in de O f it t ak es p ar t in the f o rm a tio n o f the hy
o id a pp ar a tu s .

8 All o ther bon es of the s k u ll an d j aws a e memb r an e bo n es

. r .

9 T he
. n u m be r O f m e m b r an e bo es O f the s k u ll is
g ea tes t in the l ower ver tebr a tes a d
n r
,
n

the m emb an e b on es p r es en t the mo s t typ ical a an g emen t in e tin c t an d p imitive g ou p s

r rr x r r

o f Amphibia an d r ep til es T he n u mber Of memb r an e bo es dimin ishes in the higher v er te

. n

b a tes chiey by l o ss b u t al so by fu sion Thu s we may n ote tha t several m emb a e b on es

r , .
, r n

p resen t in the all iga to r ar e missin g in the ca t an d rabbit This l oss o f membran e bo es is . n

ill u s t a ted in F igu r e 3 6 p age 1 0 6

r , .

IO T he n u mb er O f ca r til ag e bon es is likewise g r eatl y red u ced in the higher fo rms by a

.

p roc es s Of f u sion Of o ig i ally s ep ara te bon es Thu s the o ccip ital bo e Of mammal s con sis ts
r n .
,
n

o f f o u r o igi all y s ep ar a t e b o n es ; t he s phen o id s Of m ammal s a r e mo r e o r l es s f u sed an d in

r n

man u n ite to a sin gl e s phen o id bon e which con s is ts o f s ix o rigin all y s ep ar a te car til ag e bon es

FI G .
4C
Some b on es o fthe human
. s kull a t an earl y a ge , showin g their co m p oun d na ture. A, the
oc ci pit l b o
a n e at b ir th, sho win g the ve el emen ts O f which it is co m p o sed ; a, in t er p a rieta l ; b, su p ra

o cc ipit l ; c
a ,
ex occi pit al ; d , is the fo ramen ma gn um ; e, b a sio cci it al p . B , the ph
s en oid b on e in an

e mb ryo o f fou r mon ths , sho win g its co mpon en ts ; f, c en t er fo r the p ph

r es en oid ; g, cen ter fo r the o rb ito
Sph id ; en o p p
h, alis hen oid ; i , c en ter fo r the b asis hen oid ; j , c en ter fo r the lin gul a ; k, cen ter fo r t he
pt yg id
er o p
C, the t em oral b o n e a t b ir th, showin g its thr ee com on en ts ; l , squ amosal ; m, t ym anic ;
. p p
n, pt m
e ro as toid o r erio tic p
M emb ran e b o n es b l an k ; c a rtil a ge b on es O en s ti lin g ; c a rt il a ge, cl os e
. p pp
stippli gn T he su b s equ en t o ssi ca tion o f t he car til a g e o b li t era t es t he b o u n da ries b etween the co m
.

p o n en t s ( A a
. n d C f r o m s ec im e p
n s l o an ed b y the ana to my d e ar tm en t ; B a f ter M cMu rrich s Devel op

p '

p
men t of the H u ma n B ody, Co y right b y P B l akis ton s S on an d Com an y ) . p .

an d in cl u des so me memb r an e bon es in ad d ition

; the eth moid Of mamm al s is c om p osed Of

thr ee hmo id bon es ; the p

et bo es etc ( Fig etr o m as to id Of s ev er al o tic n ,
. .

I T he l o w er jaw u n d erg o es redu c tion to the m amm alian con ditio

I . wher e it co s is ts n, n

o fb u t a s in g l e p air o f bon es I n mos t ver tebra tes the a r ticu l a tio n O f the l o wer jaw is b etw n
. ee

the q a d a t e a d the ar ticu l ar bo n es whil e in mamm al s it is b etween the d en tary a d the

u r n ,
n

s qu am os al bo n es .

1 2 T he qu ad r at e the ar ticu l ar an d the hyo man dibul ar ar e b elieved to b e rep resen ted
.
, ,

in m ammal s in the tin y b o es f ou n d in the m iddl e e ar T he r emn an t o f the qu adr a te fo rm s

n .

the in cu s O f the a r ticul ar the m al l eu s a d O f the hyo ma dib u l a r the s ta p es

, , ,
n n ,
.

13 T he hyo id an d o ther g ill ar ches a r e gr d all y r ed c ed i l an d Vertebrates b u t per s is t

. a u u n

in p a t in the hyo id a pp a r atus an d the c ar til ag es Of the l ryn x

r a ,
 Q

I X . TH E CO M P ARATIVE ANAT O M Y OF T HE M US CUL AR S Y S TE M

A . G E NE R AL CON S I DE R A TI ON S

1 . The kin d s an d origin of m u s cl e

may b e divided The . m u scl es o f t he v er te b r a te body
in to two g en er al cl as ses , the in vol u n tar y an d the vol u n tar y T he in vol u n tar y o r s mooth mu s cl es .

o ccu r in the w all s o f the dig es t ive tr a c t an d o ther v iscer a , an d in the s k in an d c er tain d eriv a

t iv es ther eo f . They o r ig in a t e t h
h may
r ou g h the tran s forma tion of mesen chym e c el l s , whic
b e o f variou s o igi s T he maj o ity o f the s moo th mu scu l a tu r e howev er is p ro d ced by the
r n . r , , u

mes e chyme o f the hypome e s in ce in devel o p men t t he hyp o mere cl o ses arou n d t he a che
n r ,
r n

t e o an d its d eriva tives ( s ee Fig

r n 0 p T he vol tary o r s tr ia ted mu s cl es on the o the . 1 ,
. un ,
r

han d with cer tain e cep tio s sp ec i ed b el ow a rise f rom the myo to mes
,
x nT he myo tom es it , . .

w ill b e r ecall ed a e tho s p o tio s o f the ep im e es r emai i g af ter the p im e e s hav e g ive
,
r e r n r n n e r n

i e to the scl ero to mes a d d erm a to mes

r s F ro m their o rigi al do rs al p os itio s the myo to me
n . n n s

g r
o w d o wn b et w ee n th e h yp o m e e a n d the s k in a d tho s e o f O ppo sit e s id es m eet in the med ia
r n , n

v en t al li e r S ee F igu r e 0 a d t e r ea d S ec tio I V Of the ma u l

n . 1 I n this way the e is
n - n n a . r

p od ced a compl ete coa t Of vol u tary mu scl es lyi g ben eath the s kin This mu scle coat
r u n ,
n .

is d ivid ed i to do s al a d v e tral p a ts by the ho izon tal s k el etog en o s p a tition which

n r n n r r u r

in te s ec ts the s kin a t the l at e al l i e

r T he mu s cl es d o r s al to this s ep tu m ar e call d the
r n . e

epaxial m s cl es thos e b el o w the sep tu m the hypaxial mu sc l es ( Fig s 5A p 58 0 p

u , , . I , .
,
2 , .

All o fthe mu s cl s o ig i a ti g f o m the myo to mes ar e vol un ta y m scl s a d are d e ig a t ed

e r n n r r u e n s n

as p ri t l o a m tic mu s cl es
e a r so No t al l Of the vol u n ta ry mu s cl es a e h w ve of this k i d
a . r ,
o e r, n .

I n t he g ill eg io n o f ve teb a tes a sys tem of vo lu n tary mus cl es is d evelo p ed fo r mo vin g the
r r r

ill a ches S i cre th e g ill s a d el a t ed p a r ts a e O f en t o d erm a l o rig i

n th e m
n u s cl es i th
r r n, n e
g .

wall in th g ill egio

s ar e ho mo l og ou s w ith the mu s cl es o f the r es t o f the d ig s tive t rac t
e r n e

a d n in f c t d iv d fr m the mes en hyme f the hypomer s

a re , a , er e Thes e g ill ar ch mu scl es a e
o c o e . r

co equ e tly d es ign a t ed as vi cer al mu s cl es al tho u gh un lik e the mu s c l es o f the vis c e a they
ns n s , ,
r ,

a e s t i ted
r n d vol
r a ta ry The e a e con s equ e tly two k i ds of volu tary mu scl es id e
a un . r r n n n ,
n

tical i s t ru c t e b t di f e e t i o ig in
u n f n n r the p arietal o r s o ma tic m s cl es d e ived f ro m the
u r r u r

myo tomes a d dis t ibu ted widely o ve the b ody a d the vis c ral m s cl es d e ived f rom
n r r n e u r

the hyp om er es a d fou d o l y in the g il l r eg ion n n n .

T he s tu d e t shou ld n o te tha t the te m s mu s cl e a d mu s cu l a sys tem whe u s ed withou t

n r n r ,
n

f ther qu ali ca tio r ef e o ly to the vol u tary mu scl es I n d issec ti g a ve t eb a te onl y the
ur n ,
r n n . n r r

vo l tary mu s cl es a e s t died as the s tu dy o f the i vo l n ta ry m s cl es p p e l y b el o g s to

un r u ,
n u u ro r n

his tol ogy I t shou ld fur th r b e al ways born e in min d that whe the e p essio n visce al mu s cl es
. e n x r r

is em p l oyed this efe s n ot to the i vo l u ta ry mu s cl es o f the vis cer a b u t to the vol

r r t ry n n un a

m s cl es Of the g ill a ches

u I t is a su med tha t the s tu d e t u n der s ta ds the his to l o g ica l differ
r . s n n

en c e b etween sm o oth an d s tr ia t ed mu s cl e .

Con s ul t al so the a ccou n ts of the com parative ana tom y of the mus cul ar s s te y m in K ,

W , an d Wd .

B . TH E MU S CL E S OF THE G
DO F I S H

I . The p arie tal or

skin from the d og sh a t s m t
o a i c mu s c le s S trip . of
f the
the base of the t ail in the n ei ghborhood of the elvi c n s I doi t his m ake
p n , n g .

a cu t thro u gh the skin ras th t ed e wi t h th ers d s ri f

f a ie e of
g p e cu g e n g a n t p o p
, c ,

1 28
l 3 o LAB O RATO RY M A NUAL FOR VE R TE B RATE ANATOM Y

the ven t ral su rfa ce exten din g forward to the m ou th wi th bers dire c t ed back
, ,

ward ; a n d the addu ctor ma n dib u l ar is the thi ck m u s c le a t the a n gl e O f the jaws
, ,

u sed t c lose the lower jaw T he c o n s tri c tors op en re gill sli t s by dimin ishing the
p .

dis tan c e be tw een the gill ar ch es by their c o n tr a c tio n Th ese mu s cles O f the .

d o g sh p ersis t in hi gher vert eb ra t es af t er the gills have been los t in asso cia tion
wi th those s t ru c tu r es which are derived from the gill ar ches tha t is the jaws the , , ,

hyoid an d the car tilag es of the laryn x

,
.

C . TH E MU S CLE S OF NE CT URUS

Anim als tha t have been p reserved for som e tim e in fo rma li n are p referable for
the s tu dy of the mu s c les M ake a m edia n dorsal i n cisi o n exte n di n g the len gth
.

of the head a n d t run k Loosen the cu t ed ges of ski n wi th the n gers n o tin g in
.
,

the c u t s u rfa c es the a sk shap ed cu taneo u s g la n ds whi c h se c r e t e slim e Then .

wi th the n gers sep ar a te the ski n from the m u s c les in a ven tral dire c tion u n til yo u
have rem oved the skin from the head t ru nk and app endages The gills are to be , ,
.

lef t in p la c e The white brou s m a terial be twee n the skin and mu s cles is the
.

s u b cu ta n eou s c o n n e c tive t iss u e or s u per cia l fa s cia .

1 T he mu s cl e s of the tru n k an d tai1

. Th ese mu s cl es p reserve the g eneraliz ed
.
-

arran gem en t typ i cal O f p rimi tive a n d embryonic ver tebra tes They consis t as .
,

in the do g sh of a series of m yo tom es sep ara t ed by m yosep t a

, ,
The myb tomes .

are lon g n early re c tangular blo cks ex t en din g from the mid dorsal to the mid
,
-

ven tra l line Their n arrowed dorsal en ds slan t fo r ward No t e their division
. .

i n to epaxial an d hypaxial p or tio n s by the horizon tal sep tu m Al thou gh the .

t run k mu s c les app ear to be u nm odied they are in reali ty already sep ara ti n g ,

i n to layers On cu tting i n to the ep axial m u s c les they will be seen to co n s ti tu te

.

a m ass whose bers are all di rec t ed forward ; thi s m ass c orresp onds to the l on
g iss im u s d or s i m u s c le of h i g her fo r m s O n c u t tin g i n t o the .h yp a xial m u s c l es o n ,

the o ther ha n d they will be fo u nd divisible in to three di s tin c t layers

,
The .

ou ter layer or exter n al obl ique m u s c le is c om p osed of bers dire ct ed obli qu ely
, ,

ven trad an d c au dad The middle layer or in ter n al obliqu e m u s cle is comp os ed
.
, ,

O f bers dire c t ed c ra ni ad and v en t rad On c u t tin g thro u gh the i n t ern al obli qu e

.

a third layer the tr an s vers e nex t to the body cavi ty will be fou n d On ei ther .

side of the m edi an ven tral lin e the bers of the m yo tom es are di re c t ed p arallel
to the lon gi tu dinal a xis of the body and form a n a r row ba n d the r ec tu s a bdomin is ,

mu s c le whi ch is ra ther dis tin c t j u s t a n t erior to the p elvi c girdle

,
.

2 The m u s c le s Of the p e l vi c gir d le an d b in d 1im b

.
I n this r egl on O f the .

body as in the case of the do gsh the series O f m yo tom es is i n terrup ted by
, ,

the p res en c e O f lim b m u s c les ex t e n di n g fro m the t ru n k in t o the lim b

, E x am in e .

the ven t ral fa c e of the p elvi c girdl e I t is c overed by m u s c les whi ch m a y be sep a
.

ra t ed in to two m u s cl es The a n t erior mu s c le c overi n g the pu bi c car tilag e ex ten ds

.

from the m edian line Ofthe girdle to the femu r ; it m ay be nam ed the pu b ofemor a
lis ex ter n u s The pos terior m u s c le ex t en ds from the ischial regio n of the girdle
.
 COMPARA I VE ANAT OMT Y O F T HE M U S CUL AR SYS TE M 1 3 1

to the t ibia and is c alled the ischiotib ial is F rom the a n terola t era l m argi n of .

the girdle a m u s c le p ar tly co n c ea led u n der the p u b o fem ora l is ex t ern u s p roje c ts
, ,

forward and its bers be com e con tin u ou s wi th the ven t ral p or tio n s Of the
, .

m yo t om es . This is the p os terior end of the re c tu s abdom inis no ted above .

Arou n d the anu s is a large m ass the an al gl an d , On rem ovi n g this gland two .

m u s cles are reveal ed on ea c h side The m edial one is the is chiocau dalis ext en d
.
,

in g from the p os terior m ar gin of the is chiu m to the tail a n d the la teral o n e is ,

the pyr ifor m its a n te rior end fas tened to the ischio tib ia l is its p os terior en d p r o
, ,

c eedin g i n to the t ail F ollow these two m u s c les in t o the tail

. Tu rn the a nimal .

sidewise ben d the l eg forward and n o te tha t the p yrifo rrn sp li ts n ear its anterior
, ,

en d and gives o f f ano t her m u s c le whic h p asses to t he fe m u r This mu s cle .

is the femor ocau dalis The three mu s cles j u s t men tio n ed are tail or cau dal
.

m u s cles .

Cer tain deni tions m ay now be m ade E a c h mu s c le is a tt a ched a t its two

.

en ds a n d more or less free in t he middle The a tta ched en d whi c h is x ed an d.

imm ovable is nam ed the or igin ; the a tta ched en d whi c h m oves when the mu s cle
c o n tr a c t s is the in s er tion The bes t m e thod of n am ing mu s cles is to combine the
.

origi n a n d i n ser tion in to a co mp o u n d word the ori gin p rec eding The n ames of ,
.

the m ajori ty O f the mu s c les given above are of this ki n d b u t m u s cles are also ,

frequ en tly nam ed from their shap es p osi tions etc In the c ase of m any mu s cles
, ,
.

ei ther end ma y serve as the ori gin or inser tion dep endin g on w h i ch en d is held
,

x ed ; thu s the c au dal mu s cles of Nectu r u s will m ove the tail if the p elvic girdle
,

is x ed or will move the girdle if the t ail remain s s ta tionary The mu s cles of .

the girdle already nam ed serve chi ey to be n d the limb t oward the m edian
ven tral line a movem en t known as addu ction
,
.

On the ven tral su rfa c e Of the thi gh fo u r mu s c les ma y be ide n tied These .

are : the pu b ofemoralis in tern u s the m os t a n t erior On e ori gin a ti n g o n the an terior
, ,

rim Of the a c e t ab u l u m an d in ser t ed o n the di s tal en d Of the fem u r ; the pu b otib ia l is ,

nex t p os terior to the p rec eding ex tendi n g from the a ce tab u l um to the p roxim al
,

en d of the tibia ; nex t the dis tal p or tion Of the is chio tib ialis this p ar t bei n g so me
, ,

tim es desi gn a t ed the gr acil is m u s c le ; and m os t p os teriorly a slender m u s c le the , ,

fe m or o b u l a r is ori g i n a
,
ti n g on the p re c edi n g m u s c le a n d i n s e r t ed on th e b u la .

S o m e of these mu s c les ac t to bend the shank towar d the thigh an a c t known as ,

fl ex ion .

On the dorsal side Of the thigh in addi tion to so m e of the mu s cles already
men tio n ed whi c h app ear also in dorsal view there is fo u n d c hi ey the ilioexten
, ,

s or iu s a broad ba n d ori gi n a tin g on the ili um (whi ch m ay readily be fo u n d as a

,

cu rved bone be tween t he m yo t o m es) and i n ser t ed o n the dis t al en d of the fem u r

by a tendon ( shee t Of c on n e c t ive tissu e) whi ch also p asses o n to the shan k This .

mu s c le a b du cts the thigh tha t is draws it dorsally and s traigh t en s or ex ten ds the
, , ,

shank B e tween the ilio ex ten soriu s and the p u b o femora l is in tern u s p ar tly
. ,

c o n c eal ed by the la t ter is the il iofemor a l is e x t en din g from the ili u m to the fem u r
, ,
.
 1 3 2 LABORATOR Y MANUAL FOR VE R T E B RA E ANA OM T T Y

The homolo gy of these mu s c les wi th those of m amm als is so m ewha t dou b t fu l .

S om e su gg es tio n s are given by W p a g e 2 2 9 ,

A gu r e whi ch m ay aid in id en tify
.

in g the mu s c les will be fo u n d in W p a g e 2 2 8 ,

.

3 The mu s c le s Of the p e ctor al g ir d le an d fore 1im b

.
S tu dy the ve n tral .

su rfa c e of the p e ctoral girdle Coverin g the c ora coid car tila ge is a fan shap ed
.
-

mu s cle the s u pr a cor a coideu s inser ted on the humeru s

, ,
On ei ther side ove rly
.
,

in g the elon ga t ed p ro cora c oid car tilag e is the procor a cohu meral is i n ser t ed on the
, ,

hu m eru s j u s t an t erior to the p rec edi n g B ehind the su p ra cora coid eu s an d

.

p ra c t i c ally c o n t i n u o u s wi t h it is t h e lar g e fa n sha p ed p e-

c tor a l m u s c le w hi ch ,

ori gin at es on the linea alba an d som e of the m yosep ta Coverin g the ven t ral .

s u rfa c e b etween the two p ro cor acohu m eral es is the broad s ter n ohyoid mu s cle , ,

differen ti a ted ou t Of the ven t ral p or tio n s of the m os t an terior myo tom es At its .

an terior en d the two halves of the s tern ohyoid mu s cle fork ano ther p air of ,

mu s c les (gen iohyoids ) bein g inser ted b etw een the tw o p ar ts of the fork The .

s ternohyoid shou ld be tra ced p os t eriorly ; it has som e a tta chm en ts to the
p e c t oral g irdle p asses
,
dorsal to th e p e c t oral m u s c les w hi c h sho u ld be lif t e d t o

see it and be com es c on tinu ou s wi th the re c tu s abdomi ni s The s ternohyoid

,
.

and re c tu s abdom i n is p reserve alm os t p erfe c tly the segm en ted condi tion .

On the dorsal side of the p e c t o r al girdle ide n tify the l a tis s imu s dors i the m os t ,

p os t erior and lar g es t of t h e dorsal g irdle m u s c les I t ori g i n

. a t es by abo u t ve
sep ara te slip s from the m yosep ta Of adja cen t myo tom es an d these sli p s c onverge ,

to the hu m eru s This mu s cle illu s t ra tes very well the c omp o u n d origin of
.

limb mu s cles by slip s or b u ds from several myo tom es An terior t o the la tissim u s .

dorsi and coveri n g the su rfa c e of the s capu la is the dors alis scapu l ae also inser t ed ,

o n the hu meru s An terior to this is the tr a peziu s ( or cu cu l l ar is ) ori gin a tin g by

.

two slip s or heads and inser t ed on the s c a p u la B e tween the trap ezi u s an d the
.

p r o c o r a c o h u m e ra li s is th e om o h y oid m u s c le ru nn i,
n g p arallel t o t h e la tt er fro m ,

the hyoid t o t he girdle Cu t thro u gh the m iddle O f the la tissim u s dorsi an d

.

nd benea th it the typ i c al m yo tom es These be com e in hi gher form s the .

s err a tu s mu s c le .

I n the upp er arm ide n tify on the ven tra l side two mu s c les : an an terior o n e ,

the b iceps e x t e n din g from the hu m eru s to radi u s ; a n d a p os t erior cor a cobr a chial is
,

ori gi n ati n g from the cora coid c ar tila ge an d p assing to the dis tal en d of the
hum er u s The dorsal side of the u pp er a rm is o ccup i ed by the large tr iceps
.

mu s cle wi th ori gi n s on c ora c oid s ca pu la and hum eru s an d i n ser tion o n the
, , , ,

ulna . The mu s cles of the forearm are in two chi ef masses a dorsal exten s or ,

which ex t en ds the ha n d an d a ven tral exor whi ch ben ds the han d .

Th vi er l d h ea d m s le s The vis eral m s les or m s les of t h

4 . e s c a an u c . c u c u c e

g ill ar c hes are c onsiderably m od ied in rela tion t o th e g rea t er se p ara tio n of h ead
from t ru nk an d the degen era tion Of the gill ar ches S kin the head c omp l etely . .

Across the ven t ral s u rfa c e fro m o n e half of the low er jaw t o the o ther p ass es
the myl ohyoid mu s cle I t origi n a t es o n the jaws an d is in ser ted on the m edian
.
1 34 L AB OR A OR T Y MANUAL FOR VE R TE B RATE ANATO M Y

app li c able to ei th er an im al differen c es be tw een them bein g sp ec ically no ted

,

wherever n e cessary .

I The d erm al or in te gu m e n tal m u s cl e s

.
S li n the a n i m al a n d in doi n g so

n o t e the s t ru c tu res n a m ed in the followin g p ara g ra p h ( I f t he a nim al has .

already been skin ned t his p ar t Of the work will have to b e o mi t t ed ) I n ski n
,
.

nin g m ake a m edi an dorsal in c ision fro m the b as e Of the t ail t o the ba ck of the
,

head B e su re that this an d following in cisio n s cu t throu gh the skin o nl y

. .

M ake an in cision thro u gh the skin aro u n d the t hroa t arou n d ankles an d ,

wris ts a n d i n cisio n s alo n g the o u ter su rfa c e Of the lim bs Co n n e ct these wi th

,
.

t he m edia n in cisio n L oosen the ski n alo n g the i n cisio n s and gr ad u ally work
.

the skin loos e fro m the mu s c les u si n g n ge rs a n d ba c k of the s calp el

, Avoid .

as far as p ossible the me thod of cu t ti n g the skin from the body as o n e is liable ,

to cu t in t o the mu s cles by this p ro c ed u re Work from the dorsal side toward the
.

ven t ral sid e L eave the skin o n the head and o n the p erineal region for the p resen t
. .

T he followin g p oi n ts sho u ld be n o t ed d u ri n g the skin nin g The skin is .

c o n ne c t ed wi th the u n derlyi n g mu s c les by a loose weblik e m a t erial t he s u b cu ta ,

n eou s con nective tis s ue or su per cia l fa s cia of t en im p regna ted wi th fa t B elow
, .

this is the m u ch rm er and t o u gher c o n ne c tive tiss u e on the s u rfa c e of the m u s c les ,

formin g the deep fascia P assin g from among the mu s c les in to the skin will b e
.

seen at regu lar in tervals whi ch rep resen t the segm e n t s of the body a slen der
, ,

c ord c omp osed of an ar t ery a vein a n d a se n sory n e r ve

, , ,
Th ese m ay be severed . .

Other blood ves sels n o t segm en tally arran g ed will also be seen p assi n g o n to the
, ,

u n d e r s u rfa c e O f the skin from a n t erior a n d p os terior re gio n s t o w ard the middle
, .

T he ar teries are colored by an i n je c tio n m ass and are rea dily re c o gni z ed The .

vein s are u su ally of a very dark reddish brown c olor All vessels to the ski n -
.

shou ld be severed .

When the ski n has been loosened to the sides of the anim al there will be n o ted ,

a thin layer of mu s c le bers on its u n der su rfa ce app eari n g like a n e s tripi n g , .

Toward the ches t and shoulder regio n thi s ass u m es the fo rm of a thin sh ee t .

This mu s c le is a d erm al or skin mu s cle the pan n icu lu s carn osu s or cu ta n eou s
,

maximu s I t c ov ers the en tir e la teral s u rfa c e of the thorax a n d abdo m en bein g
.
,

m ore p rom in en t a n teriorly On c o n tin u in g to skin forward and v en t rally

.

t he m u s c le will be fo u n d to t ake its ori gin from the o u ter s u rfa c e Of a mu s c le

( la ti ssi m u s dorsi ) si tu a t ed p os t erior to th e sho u lder and fro m t he axilla in the ,

c a t the m e di al side O f the h um eru s ( rabbi t) and from the linea alba and vario u s
, ,

p oi n t s on t he ve n t ral side of t h e t hora x in bo th a n im als These p oi n t s of ori gi n .

shou ld be cu t thro u gh an d the cu tan eou s maxim u s removed wi th the skin to

whi c h it gen erally adheres T he m u s c le is in ser t ed on the skin a n d s e rves to
.

shake the skin I n m an this mu s cle is degen era t e I t is an ou tgrow th O f the

. .

la tissim u s dorsi mu s cle .

Th ere is on e o ther d erm al mu s cle the pla tys ma This will be fo u n d o n the
, .

u n der s u rfa ce of the skin of the n e ck a n d head a n d c o n sis t s Of m a n y di ffere n t ,

 COM P ARA I VE ANATOM T Y OF THE M U SCUL AR S Y STE M 1 35

p ar t s wh i,
c h have re c eived se p ara t e nam es S ome of these will be seen la ter .
,

b u t the s tu dy of the p ar ts of the p la tysm a is alm os t im p ossible in an y b u t freshly

killed sp e ci m en s The p la tysm a m u s c le is i n s er ted o nthe skin of the ears eyelids
.
, ,

lip s et c an d s erves to m ove th em I n ma n it c o n s ti tu t es the mu s c les of fa cial

,
.
, .

exp ressio n The p la tysm a is a vis c eral mu s c le d erived from the mu s cles of the
.
,

hyoid ar ch by ex ten sio n .

In f em ales o n the u n der su rfa c e of the skin of the ve n t ral side the mom
mary or mil k gl a n ds will be n o ted sp read o u t as a thin irre gu lar layer .

The ski n havi n g been rem oved and dis carded c lean aw ay fas cia an d fat ,

from the s u rfa c e of the mu s c les There is gen erally a larg e mass of fat at the .

base of the hi n d l egs I t will n ow be seen tha t the exposed su rfa c e in par t
.

c o n sis t s of mu s c les p i n kish m asses c o m p osed of p arallel bers a n d in p ar t of the

, ,

deep fas cia which form s very s t ron g whi te shee ts The p os terior half of the ba ck .

is c over ed by su c h a sheet known as the l u mb odors al fascia I n the m edia n

,
.

ven tral li n e is the lin ea alba The an gl e be tween the base of the thigh an d
.

the abdom i n al wall is k n own as the in gu in al regio n At the bo tto m Of this will .

be fou n d in the ra bbi t a s to u t whit e shinin g cord the in gu in al ligamen t whi ch , ,

s tre t ches from the p u bi c sym p hysis to the cres t of the I t is absen t in the
ca t. The angle between the u pp er arm an d ches t is c alled the axill a or axil l ar y ,

fossa .

In s tu dyi n g the mu s c les it is n e c essary to sep ara te ea c h mu s cle from its

n ei ghbors This is do n e by sear chin g c arefu lly for the white li n es of c o n n e c tive
.

tiss u e whi c h m ark the bo u n daries of mu s c les an d sli tti n g alo n g these li n es wi th
the p oi n t of the s c al p el Observin g the dire c tio n in whi ch the bers ru n wil l
.

also aid in sep ara ti n g m u s cles sin c e the bers in o n e mu s c le gen erally ru n in
,

the sam e dire c tio n whi c h is u s u ally di f feren t from tha t of the nei ghbori n g m u s cl es
,
.

Af t er fre ei n g the m argi n s of a m u s cle the n gers sho u ld be worked u n der the mu s cl e
u n til it is sep ara t ed fro m its fellows As ea ch mu s c le is i n c losed in a c o n n e c tive
.

tissu e shea th it will sep ara t e sm oo t hly from its n ei ghbors ; the p resen c e of ro u gh
,

ed ges indi c a t es tha t the m u s c le i tself has been c u t Avoid the u se of sharp .

in s t ru m en t s in freei n g the mu s c les I n case it is n e cessary to cu t thro u gh a .

mu s cle in order to reveal ano ther mu s c le ben ea th it a lways cu t thr ou gh the cen ter ,

f the mu s cl e
o .

2. a mu s c le W e shall rs t s tu dy the p ar ts of som e c o n ven ien t

The par ts of .
-

mu s c le in o rder to be com e a c q u ai n t ed wi th the t erm i n olo gy app lied to mu s cles .

F or thi s p u rp ose we shall sel ec t the ex ter n al obliqu e The ex tern al obliqu e is
'

t he larg e m u s c le c overin g the sides of the abdom en The s up ercial fas cia sho u l d .

b e c lean ed away fro m its s u rfa c e On c losely in sp e c ti n g the c lean ed su rfa c e of

.

the ex tern al obli qu e the m u s cl e wil l be O bserved to c o n sis t of n u m ero u s p i n kish

,

s trip es dir ec ted obliq u ely ven trad an d ca u dad .

The p in kish s trip es are b u n dles of mu s cle bers c alled fa s cicu li The whi te ,
.

m a te rial b e tw een the fas c i cu li whi ch bi n ds them toge ther in to a mu s cle an d

 1 3 6 LAB ORA OR T Y MANUAL T
F OR VE R E B RA E ANA OM T T Y

whi ch di p s down be tw een them is p art of the deep fas cia I t form s a shea th the .
,

p e r im y s iu m for ,
ea c h fas c i cu l u s The p ar t of
. th m u s c le w h i c h is c o m p osed of
fas ci cu li wi th their p erim ysia is kn own as the fles y p ar t of the mu s c le or the
, ,

bel ly . The fas ci cu li do n o t ex t end c om p l etely to the en ds of a mu s cle b u t the ,

deep fas cia does Co n s equ en tly the en ds of mu s cles are n on eshy comp osed
.
-
,

of conn e ctive tissu e on ly These co n n e c tive tissu e en ds of mu s c les comm on ly

.

form tou gh shinin g bands or cords kn own as ten don s When the ten don is very ,
.

broad an d at it is of ten called an apon eu r osis or the n am e fascia m ay be re tained

, ,

for su ch broad t en don s The a tta chm en ts of mu s cles ar e always by m ea n s of

.

t endons a p o n e u roses or fas ciae

,
n ever by the m u s c l e bers
,
Mu s c les are .

a tta ched t o bon es or to ten don s ap on eu roses or fas ciae whi ch them selves , , ,

are a tta ch ed to bo n es The pu rp ose Of the volu n ta ry mu s cl es is to move the

.

b bn es of the sk el eton .

The an terior p ar t Of the ex tern al obliq u e is c on cealed u n der a large a t

mu s c le the l a tiss imu s dor s i whi ch c overs the an terior p ar t of the ba ck an d slop es
, ,

toward the u p p er a rm The p os terior bou n dary O f thi s m u s c le shou ld be lo c a ted

.

and s li t an d it shou ld th en be lif ted from the s u rfa c e of the ex ternal Obliqu e by
,

thru s ti n g the n gers be tw een I f fa t is p resen t be twee n these two m u s cles it

.
,

mu s t be clea n ed away The m os t p os terior c hes t mu s cles also cover the an t erior
.

p ar t of th e e x t ern al O bliq u e and sho u ld be lif t ed o f f in a si m ilar m anner The .

ex tern al Obliqu e will then be fou n d to be a tta ch ed to the p os terior ribs by sep ara t e
slip s The x ed p oi n ts of a t ta chm en t of a mu s cle are called its origin When
. .

there is m ore than one ori gi n ea c h o n e is k n ow n as a head ,

When there are .

a n u m ber of poin ts of a tt a chm e n ts segm en tally arra n g ed they are generally ,

d esign a ted as s lips The ori gin of the ex tern al obliqu e is from the p os terior
.

ribs by sep ara te slip s an d from the lu m bodorsal fas cia I ts bers ( fas ci cu li ) .

p ass obli q u ely dow n ward a n d ba c kward a n d in t h

,
e rabbi t the m ore dor s al
on es p ass n early s trai gh t cau dad The movable p oi n ts of a t ta chm en t of a.

mu s c le o n whi ch it ex er ts its effe c t are c alled its in s er tion The i n ser tion Of the .

ex tern al obliqu e is by way of an ex ten sive ap oneu rosis whi ch p asses to the
m ed ian ven tral line T he i n ser tio n is : rabbi t o n the li n ea alba by its ap o n eu
.
,

rosis an d on the i n guin al li gam en t whi ch is in tu rn a tta ched to the ilium and the
,

p u bi c s m
y p hysis ; c a t o n t h,e li n ea alba a n d t h e p u bis by it s a p one u roses and ,

o n the m edia n ven t ral lin e ( ra he) of the t hora x The f u n c tion of a m u s c le
p .

is called its action The a c tion Of the ex tern al Obliq u e is c o n s t ri c tor of the
.

abdom en .

I n the followin g di ss ec tion of the m u s cles the disse c tio n is t o be c o n n ed

s tr ictl y to the left s ide leavin t h ri h t sid in t a t for th e disse c tion of o ther
, g e g e c

3 . The m u s c le s of the
ab d o min al wall The
abdomin al wall is c omp osed .

of three layers of m u s cles wi th their ap o n eu roses The ap o n eu roses of these .

mu s cles ar e O u ite ex t en sive The three layers are : an ex tern al layer the
.
,
 1 3 8 LAB ORATOR Y MANUAL F OR VE R TE B RATE ANATOM Y

They are a tta ched to the sa c ru m the iliu m the ver tebrae an d ribs a t variou s , ,

o i t s a n d t o t he ba c k O f t he sk u ll a n d serve t o m ove t he ver tebral c ol um n

p n , ,

as a .whol e or in p ar t raise the head etc Al l {hrieta l mu s c les n o t i n c lu d ed in

, , .

thi s m ass a r e hyp a x ial m u s c les w hi c h have in som e c ases sp read dorsally so as
,

t o c over t he ep a xial mu s c les .

cut edge o f
cl avo trapezius

l ong head of
i
the tr ceps

trans vers us cos tar um l ong hfa d 0

the tn oeps
serratus ven tral s

FI G 4 1
. .
Ven tral view o f the an t erio r p ar t o f the c a t to sho w t he mu scl es Al l d erm al muscl es
.

hav e b een r emov ed . Su p

mu sc l es o n the right s id e, d eep er
er cial l ayer o f m u scl es o n the l ef t sid e ,
a ft er r em ov a l o f the p ec t o r al mu s cl es , s t ern om as to id , myl ohyoid , an d dig a s t ric . T he n erv es an d

b l oo d v es s el s which c r oss t he axill a hav e b een o m itted . T he view o f the a xill a is d ieren t fr om t ha t
rev eal ed in the dissec tio n given in the text . T he epitr o chl ea ris is ca ll ed ex t en sor an tib ra chii in t he t ex t .

5 T h e.m u s c le s of the c he s t ( g
F i T u r n t he a n i m
. al
o n it s ba c k.a n d
exp ose the c hes t by sp rea ding a n d fas tenin g t he fore lim bs The grea t mu s c les .

c overin g t he c hes t or ve n t ral side of t he t hora x are t he pector a l m u s c les They .

are divisible in to several p or tions whi ch are n ot very deni tely sep ara ble from
 COM P ARA I VE ANATOM
T Y O F THE MU SCUL AR S Y ST E M 1 39

ea ch o ther Accordin g to B there are ve p ar ts in the rabbi t an d a c cording to

.
,

R a n d J fo u r p ar ts in the c a t
, Of these we m ay co n sider the follo w in g . .

R abbi t :
'
a) P ector a l is maj or This is a large mu s cle o ri gi n a ti n g from the whole length
.

of the s t ern um and inser ted on the h um eru s I t covers m os t of the su rfa ce of .

the c hes t b u t the i n ser tion is c o n c ealed by a m u s c le c o m i n g dow n fro m the head
,

( c lavodel t oid ) Ac tion d r aws th

. e a rm toward the c h es t
, .

b ) P ector alis pr imu s A sle n der mu s c le a t the a n terior en d Of the pr e

.

c edi n g a n d c overin g its a n t erior bers Ori gin m a n u bri um of the s t ern u m ; .
,

in s er tio n hu m eru s ; a ction like the p re c edi n g I ts a n terior border is in co n ta ct

, , .

wi th the c lavodel toid whi ch also p ar tly c overs its in ser tion
, .

c) P ector a lis min or Cu t throu gh the m iddle of t he b elly of the p e c t oralis

.

major and up on dee c ti n g the cu t ed g es n o t e i n tern al to it a sim ilar mu s c l e

, ,

the pector al is min or Ori gin m anu bri u m ; i n ser tio n c lavi c le a n d sp i n e Of the
.
, , ,

s c apu la To n d the i n ser tion loosen u p the c lavod el toid an d lo ca t e in its

.

bers a t the sho u lder a sm all slender bo n e the clavi cle The c lavi cle is on the ,
.

in n er su rfa c e of the mu s c le Then loosen the mu s cl e n ext lat eral to the clavo del
.

t oid a lo n g mu s cle c om i n g down from the ba ck of the n e ck ( an terior t rap ez i u s )

,
.

The p e ctoralis min or is i n ser ted by som e bers on the c lavi cle b u t m os t of its ,

bers sweep over the shou lder in ternal to the clavodel toid a n d an terior trap eziu s
an d are i n ser t ed on the sp ine of the s ca p u la Ac tion draws arm an d sho u lder .
,

t oward the ches t

;

Cat :
a ) P ectoan tib r a chia l is . An terior an d mos t sup er cial of the ch es t mu s cles .

O ri gi n , m an u briu m ; inser tion by a a t ,

te n do n o n the fas cia O f the forearm ;
a ction draws the arm toward the ches t
,
.

b) P ector a l is major Nex t p os terior to the p rec edi n g a n d ex ten din g an te

.

rio r l y dorsal to the p re c edi n g w h i ch sho u ld be cu t a c ross ; ori gi n a ti n g on the

s t ern um an d m edian ven tr al rap he and i n ser ted on the h um er u s Ac tion like .
,

the p re cedi n g .

6 ) P ector a lis min or Nex t p os terior to the p re c edin g and covered in larg e
.

p ar t by t h e p e c t oralis m ajor The la tt er sho u ld be cu t t hro u gh an d the ex ten t

.

of the p e ct oralis mi n or n o t ed The p e c toralis m inor is divisible in to severa l .

p ar t s . O ri g ,
i n s t er n u m ; i n ser tion h u m er u s ; a c tion like t h
,
e p re c edi ng The ,
.

in ser tion ca n n o t be fu lly t ra ced a t the p r esen t s tage Of the disse c tio n .

d) X iphihu mer alis The las t Of the ches t mu s cles a t hi n at lon g mu s cle
. , ,

p assin g fro m t h e x i p hoid p ro c ess of t he s t er n u m it s a n terior p ar t p assi n g ,

dor sal t o the p os t erior p ar t of the p e ct ora l is m in or an d in s er ted on the hum eru s ,
.

The inser tio n is c overed by a m ass of fat in the axilla R em ain s of the cu tan eou s .

m axim u s are p robably p resen t o n its su rfa c e Ac tio n like the p re c edi n g .
,
.

6 The mu c s o
. s le f th e ne c k an d th roa t i
( g 4
F s 1 or S li t th e ski n u p .

the c e n t er of t he throa t to the tip of the lower jaw a n d loose n it so as t o exp ose
 1 40 L AB ORAT OR Y MANUAL FOR VE R E B R A E ANAT OM T T Y

fu lly the lower jaw No te in doin g so p art s of the pl a tys ma mu s cle on the u n der
.

sid e of the skin I t sweep s from the m edian dorsal lin e of the n eck arou n d the
.

sid es of the head to fa c e an d ears an d p or tion s Of it gen era lly are a tta ched n ear
,

the a n terior en d of the s t ern u m I n disse c ti n g the thr oa t m u s cles work o n

.

o n e side o n ly leavin g the o ther in ta c t for the s tu dy Of o ther p art s Avoid

,
.

c u tt in g a n y blood vessels A larg e vein the ex tern al ju gu l a r vei n ru ns in

.
, ,

the su p er cial m u s c les of the throa t At the a n g le of the jaw is a rou nded
.

p in kish body th e s,
u b ma x ill ary gla n d o n e of the salivary gla n ds
, O ther s m all .

bodies are lymph gl an ds .

) pS aecia l por tion of the l

p ya t s ma A broad thi n she e t of de rm al.m u s c le
ex ten ds from the m an u brium of the s tern u m forward , forkin g like the le tter V,
ea ch half in ser tin g a t the base of the ear This is the depr essor con chae pos ter ior
'
.

a n d is the m os t su p er c ial mu s c l e o n the v en t ral su rfa c e of the n e ck The .

ex t ern al j u g ular vei n ru n s in it I t is a p ar t of the p la t ys ma

. I t sho u ld be .

well sep ara t ed from the u n derlyi n g mu s cles an d the p os terior end severed an d
tu rn ed forward wi tho u t however inj u ri n g the vein
, , ,
.

b ) S ter n ohyoid This is the lon g mu s cle in the m edian li n e of the neck
.
,

the tw o m em bers of the p air bei n g c losely f u sed in the m edia n ven tral lin e .

Ori gi n m a n u bri um Ofthe s tern um ; in ser tio n a n t erior horn of the hyoid
, , F ollow .

the m u s c l e u p to the t hroa t a n d feel wi th the n gers the bony hyoid o n whi ch
the m u s c le is i n ser t ed Ac tion draws the hyoid p os teriorly or raises the s t ern u m
.
, .

c) S tern omas toid The lo n g mu s c le on ea ch side of the p re c edi n g the two

.
,

m em bers O f the p air c o n v er gi n g toward the ma n u brium Of the s ter n um from

whi c h t hey ori gi n a t e ve n t ral t o t he ori gi n Of the p re c edin g I n ser tio n m as toid .
,

p ro c e ss of t h e sk u ll ; a c t io n si n gly tu r n t he head t o g e t her de r ess the head o n

, p ,

the n e ck .

d) Cl eidoma s toid an d b as iocl a vicu l a r is These two lo n g s trap shap ed m u s cl es

.
-

are next la teral to the p re c edin g an d u n i t e a t the c lavi c le wi th the c lavodel toid .

The Cleidom as toid is the m ore m edial o n e an d lies la t eral an d som ewha t dorsal
to the s t em om as to id Ori gi n m as toid re gio n of the sk u ll ; i n ser tio n clavi c l e ;
.
, ,

a ction eleva t es clavi cle or tu rn s the h ead The b asio clavicu l ar is is sli gh tly
, .

lat eral to the p re c edin g at its c ra n ial en d b u t crosses ven tral to it c au dally so
tha t its i n ser tio n on the c lavi c le is m edial to tha t of the Cleido m as toid Ori gi n .
,

o ccip i tal bo n e ; in ser tio n c lavi cle ; fu n c tio n like p re c edi n g The ori gin s of th ese
, ,
.

mu s c les c an n o t be followed ou t c o n veni en tly .

e) Cl a vodel toid Co n ti n u a tio n of t he two p re c edi n g mu s c les

. Ori gin .
,

c lavi c le ; i n ser tio n h u m eru s ; a c tio n

, rais es the h u m eru s
,
.

f ) M asse ter Th .e g r ea t m ass of m u s c le c overi n g the a n g le of t he jaws i t s ,

o u t er su rfa c e wi th a very tou gh shi n in g fas cia Ori gin z y gom a ti c ar ch ; in s er .

,

tio n o u t er s u r fa ce of the p os t erior en d O f the m a n dible ; a ctio n closes the lower

, ,

jaw ( el eva t or of the j aw ) .

1 42 LAB O RA TO RY MANUAL F O R VE R TE B RATE ANATO MY

s t ern om as t oid Ac tion .

, p u lls clavi cle c ran iad or tu rns h ead a c tin g sin gly or
, ,

lowers h ead on ne ck .

d) Cl a votr a peziu s The

an d cl a vob r a chl
ial
on g m u s cl e o n the side Of the ne ck
.

and p assin g ov er the ven t ral su rfa c e of t he shoul der t o the forearm is som e tim es
c onsidered as o n e mu s c l e the cephal ob r a chial or as t wo ,
I n the la t t er c as e the
,
.

u pp er p art is known as t he cl a votr apeziu s a n d ext en ds from t he sk u ll t o t he c lavi cle .

I t will be c on sid e red la t er The lower p art from t he c lavi c le t o the fore
.

a rm is the cl a vob r achial is Orig in c lavi c le and bers of the c l av o trap eziu s ;
.
,

in sert ion u lna ; a ct io n exor of t he forearm The clavi c l e will be fou nd on

, ,
.

t he inner s u rfa c e of the mu s c le in t he shou lder r egion im bedded in the ,

m u s c le .

e) M a sseter T he g rea t thi c k mu s c le c overin g the a n gle of t he jaws si tu a t ed

.
,

in fron t Of the su b maxillary and p aro t id glands I t is c overed by a very t ou gh .

shin in g fas cia O ri gin zygomat i c arch ; in sert ion p os t erior half of the lat eral
.
, ,

su rfa c e of the mandible ; a ction el evat or of the lower jaw , .

f) T e m p o r a l R em.ove th e skin fro m t h e side of t he head u p t o t he m e d ian

dorsal line A g r ea t mass of mu s cle c overed by a s t ron g shin in g fas cia will be
.

seen o ccu p yin g t he t emp oral fossa of the sku ll dorsal t o the ear Origin from the , .
,

side of t he sku ll from the su p erior nu chal line t o the z ygom ati c p ro c ess of the
fronta l bon e an d from p art Of the zygom at i c ar ch ; in sert ion c oronoid p rocess
, ,

of the man dible ; a c tion eleva t or of the jaw in c omm on wi th the masse t er
, , .

g) D ig a s tr ic The .m u s c le lyi n g alon g t he m edial s u rfa c e of ea ch half of the

m andible . I t ext ends p os t e riorly in t ern al to the s u b maxillary gland O ri gi n .
,

j u gu lar an d mas t oid p ro c ess es of the sku ll ; insert ion ma n dible ; a ctio n d ep res , ,

sor Of t he low er jaw .

h) Myl ohyoid The thin t ransverse sheet p assin g a c ross bet w een the two
.

digast ri c s from one half of the mandible t o the o th er Ori gin m an dibl e the .
, ,

ori gin c on c eal ed by the digas t ri c s ; inser t ion media n rap he ; a ction rais es oor , ,

of t he m ou th an d b rin gs hyoid forward .

i) Gen iohyoid Cu t t hro u gh t he median rap he of t he mylohyoid

. This .

exp oses a p air of lon g sl en der m u s cles the gen iohyoid s lyin g in the median lin e , ,
.

Origi n m andible n ear the sym physis ; i n ser tion body of the hyoid ; a c tio n
, , ,

draws the hyoid forward .

j ) S te r n o t h y r oid S ep ara. t e t he tw o s t e m o hy o id s in th e m edian lin e T h i s .

exp oses the tr achea or win dpipe a tu be s ti ffen ed by r in gs of c art il age At the
,
.

top of t he tr a chea is a cham ber wi th c art il a gino u s wall s the l aryn x The c hi ef ,
.

c art ila g e of the larynx is t he larg e s h i eld shap ed thyr oid c art ilag e form in g t he
-
,

ven t ral walls of the la r ynx Ju s t in fron t of the thyroid c art ilage the b ody of the
.

hyoid is f el t as a bon y bar The s t erno t hyroid mu s cles are lo c a t ed on e o n ea c h

.
,

side of the t ra chea dorsal t o the s t em ohyoid s O ri gin s t ern um in c omm on

, .
,

wi th the s t ern ohyoid ; insert ion t hyroid ca rt ila g e of the laryn x ; a c t ion p ull s
,

the laryn x p os t e riorly .

 COM P ARA I VE T ANA TO M Y O F T HE M U S CUL AR SYS TE M 14
3

k) Thyr ohyoid S hort narrow mu s c le on ea c h sid e of the t hyroid c artila e

g
.

from whi ch it ta kes it s ori gin ; insert ion , p os t e rior horn of t he hyoid ; a c tion ,
raises the laryn x .

.
The majori t y of the mu s cles men ti oned in thi s se cti on are the vis c eral
O 0
O O 0 0

mu s c les They c orr esp on d t o the mu s cles fou nd in t he d o g s h asso cia t ed wi th t he

.

f
o f ac ial nerve

d iga stric pm id glan d

ex tern al
jugu la r vein
vator scap ulae ven tral is

toral is m i no r

secon dd d el toi

pec toralis

serrat us ven tralis

external Ob li ue
q

FI G 4 2. L a teral view o f the a n t erior

.
-
p ar t o f the rab b it to s ho w the m u scl es . T he head is tu rn ed
slightl y to g iv e a v en t r al view o f the thro a t . All d ermal mu sc l es hav e b een re moved .

g ill ar c hes They .are t he m u s c les of t he jaws t h e hyoid an d t he c ar

, t ila g es of t he ,

la r ynx all of whic h s t r uctu res are as learned in the s tu dy of t he sku ll hom olo
, , ,

g o u s wi t h t he gi ll ar c hes of shes The m asse t e r t he t emp oral t he m ylohyoid

.
, , ,

and p art of the di gas t ri c are mu s cles of the mandib ul ar arc h ; the p lat ysm a and
the res t of the digas t ri c are mu s c les of the hyoid ar ch ; while t he mu s cles of t he
rem aining arc h es be c om e mu s cles Of t he p harynx and la r ynx whi ch w ere n o t ,

s tu died .
I 44 LAB ORA TO RY MANUAL FOR VE RTE B RA E ANA OM T T Y

7 . The m u s c le s Of the u pper b ac k an d o

s h ul d er an d back Of the n e ck .

( Fig .

R ab b it .

a ) L a tiss imu s Tu rn the a n im al on on e sid e so that t he sid e on whi ch

dors i .
,

t he mu s c les hav e already been di ss ec t ed will be u pp e rm os t T he larg e a t .

mu s c le ext en di n g obliqu ely from the middle of t he ba c k t o the fore l im b is t he

latissimu s dorsi Origin l u m bodorsal fas cia and p os t erior ribs ; insertion on
.
, ,

t he c res t on the medial side of the hu m e ru s t he in s er t ion c overed by the c hes t ,

mu s cles ; a ct ion draws the a rm ca u dad an d dorsad

,
.

b ) An ter ior and pos ter ior trapeziu s These two mu s c les are the at thin .
,

m u s cles c overin g the u pp er ba c k and ba ck of the n e ck an t erior t o t he la t issim u s

dorsi The p os t e rior t rap ezi u s ori ginat es from the l um bodorsal fas cia an d the
.

n eu ral s p ines of the t hora c i c vert ebrae and is in sert ed on t he s p in e of t he s c a u la

, p .

Action draws t he s c ap u la dorsally

,
The an t erior t rap ez iu s ori gin a t es on the
.

ext ernal o cc ip ital p ro tu beran c e of t he sku ll and l i gam en t in the mid dorsal lin e -

and is in ser t ed on the m eta c romion p ro c ess (whi ch is very lon g in the rabbit ) an d
n earby mu s c l e s an d fas c ia ; a c tio n draws the s c a p u la a n d li m b u p ward and for
,

ward The sp a c e between the two trap ez iu s mu s c les is l l ed by a st ou t

.

fas cia .

c) L eva tor s capu l ae ven tr al is This long slender mu s c le run s along the ven tral
.

border of the an terior trap ezi u s near its i n ser tio n then diver ges to its ori gi n ,

from t he vent ral s u rfa ce of t he sku ll at the s u tu re be tw een o cc ip i tal an d b a sisp he

n o id ; i n sert ion m e t a c ro m io n p ro c ess in c omm o n wi t h t he an t erior t ra p ez iu s ;
,

a ct ion pul ls the s c apu la an t eriorly

, .

d) Rhomb oideu s Cu t a c ross the m iddl e of t he b elli es of the two trap ezi us
.

mu s c les an d the la tissim u s dorsi The large thi ck mu s cle ex t en ding from the
.

vert ebral border of the s capu la t o the mid dorsal lin e is the rhom boideu s -
.

Ori gi n m id dorsal li gam en t Of the n e ck and s u c c eedi n g n e u ral sp ines ; inser tio n
,
-
,

ver tebral border of the s capu la ; a c tion draws s c apu la t oward vert ebral c ol umn ,
.

e) S pl en iu s A fairly broad b u t thi n mu s cle on the ba c k of the ant erior p art

.

of the ne ck u n der the ant erior t rap e zi u s ( Ru n n in g alon g it s ext e rnal s u rfa ce .

is a n arrow s t rap like mu s c le the r homb oideu s capitis see b elow ) Origin of
, ,
.

u id
sp leni s m dorsal line of n e ck and adja c ent fas cia ; in sert ion o ccip ita l region
, ,

of the sku ll an d a tlas ; a ction sin gly tu rns the head t o ge ther raise the head
, ,
.

Un der the sp leni us are the ep a xial mu s cles c on t in u a tion s of those alr eady ,

not ed in the lum bar region .

f) p pS u ra s in a tu s The s u p er c ial
. m u s c u lar layer of t ha t p ar t of t he s c a p la
u

an t e rior to the sp ine c onsis ts of the p e ct oralis m in or whi ch swe ep s over the ,

s capu la to be insert ed on the s p ine and vert ebral bord er L ift u p its ant erior .

border an d sep ara te it from the mu s cle be n ea th it This m u s cle is t he su pr a .

s pin atu s llin g the s u p ras pi n ou s fossa of the s c ap u la Ori gi n s u p rasp inou s
, .
,

fossa ; in sert ion g r ea t er tu berosi ty O fthe hu m e ru s ; a ction ext en ds the hu m eru s

, ,
.
 1 46 LAB O RA TO RY MANUAL FOR VE RT E B RATE ANA OM T Y

s errat u s ven tralis They lie in t e rnal t o the s te rnom as t oid Cl eidom as t oid et c
.
, , .
,

p revio u sly ide n t ied and fart her p os t eriorly t hey fo rm t he layer n e x t i n t e rn al
to the p ect oralis mu s cles Ac tion raise the lm and ben d t he n e c k
.
, .

o) S er ra tu s dor s al is (pos ter ior ) The dorsal half Of the thorax un d ern ea th
.

the la t issim u s dorsi t ra p ezi u s a n d rho m boide u s mu s cles is c ov e r ed by a s t ron g

, ,

ap on eu rosis (p art of the l um bodorsal fas cia) in the ven t ral p art of whi ch mu s c le ,

bers are p rese n t whi ch are inser ted o n the ribs by slip s The for em os t slip s .

are qu i te eshy an d take their ori gin by a ten do n from the m edian dorsal
l ine Of the n ec k . This m u s c le is the serra tu s dorsalis Action rais es the ribs .
,

c raniad .

p) I n ter c os t a l s . O n th e sides of t h e c h e s t a s e ries of m u s c les will be s e e n

r unni n g fro m o n e rib t o the ne x t one They are the ex t ern al in t er c os tals They
. .

ex tend on the ches t wall ven t ral to the inser tion of the s erra tu s dorsalis whi ch ,

mu s c le in fa ct c overs their m os t dorsal p or t io n s Origin p os t erior m argins O f .

,

the ver t ebral ribs ; i n ser tion an t erior margi n s of the s u cc eedin g ver tebral ribs ;
,

a ctio n p ull the ribs forward Observe tha t the bers Of the ex t er n al in ter cos ta ls
,
.

are dire ct ed obliqu ely ba ckward On c arefu lly cu tt in g thro u gh an y Of the

.

ex tern al in t er c os t als a layer of in t ernal i n t erc os t als will be fou n d insid e of them
, ,

t heir bers bein g dire c t ed obliq u ely forward The i n tern al in t erc os t al s a r e
.

bes t seen in the ven t ral thora ci c wall in t ern al to the s cal en es whi ch may be c u t
, ,

thro u gh . Here be twee n the c os t al car tila ges the i n t ernal in t er c os t als are n ot
c overed by the ex t er n al i n ter c os t als O rig in a n d i n s er tion m argin s of the
.
,

ver tebral an d s t ern al r ibs ; a ction lower the ribs The in terc os tals a r e the chief
, .

mu s c les c on c ern ed in the resp ira tory m ovem en t s of the t horax The s cal en es .
,

s erra tu s a n d o ther mu s c les assis t

, .

q) E p a x ia l m u s cl es of the th or ax The m ass of e p a x

. ial m u s c l e s is c o n s p i c u o u s
ru nnin g alon g the dorsal p ar t of the thorax This m ass p asses in tern al to the .

serra tu s dorsal is which shou ld be cu t thro u gh ; it li es u p on the dorsal p or tio n s of the

ribs an d th u s c on c eals the dorsal p or tions of the in ter c os tal mu s c les I t is c overed .

by the t ou gh shinin g lu m bodorsal fas cia whi ch sho u ld be rem oved The ep axial ,
.

m ass is easily divisibl e in t o a n arrow m edian p or tion n ex t to t he m edia n dorsal

lin e the s emis pin al is dor s i and a v e ry thi ck la teral p ort ion the l on gis s imu s
, , ,
.

The latter giv es off on its ve n t ral m argi n the narrow il iocos talis lyin g on the rib s
t o whi ch it is in s er t ed The longissimu s is the c on ti n u ation of the sa c rosp in ali s
.

and is ins e rt ed on the rib s The ep axial m ass may be follow ed alon g the n e ck
.

by cu ttin g the sp len iu s .

The a tt en tion of the s tu dent is di re ct ed t o the fa ct t ha t the in t er c os t al mu s cl es

rep resent the original layer of myot om es all of the m u s cl es ext ern al t o them ,

having been derived by p ro c esses of b u ddin g an d spli tt in g The serratu s .

m u s c les beau t ifu lly illu s t rat e the m e t hod of ori gin of a mu s c le
n u mber O f myo t om es sin c e the slip s by wh
, i ch t hey arose from
ar e s t ill p r ese n t .
 COM P ARA I VE T ANATO M Y OF T HE MU S CUL AR SYST E M 1 47

Cat :
a ) L a tiss imu s dor s i T u r n the animal on one side so tha t the side on whi ch
.

the mu s cles have already been disse ct ed wil l be upp ermos t The large at .

mu s cle ext en din g obliqu ely fo rward from t he m iddle Ofthe ba ck t o the u pp er a rm
is the latissimu s dorsi Ori gin from the ne u ral spines of the las t thora ci c and
.
,

mos t of the l umbar vert ebrae and from the lumbodorsal fas cia ; in sertion by a ,

ten don on the m edial s u rfa c e Of the hum eru s ; a ction p u lls the fore l imb dor ,

sally and c au dal ly .

b ) Tr apeziu s mu scl es There are three trap eziu s mu s cles in the cat ; they
.

are the thin at mu s cles c overing the ba ck and ne ck ant erior to the p re c edin g .

The p os t erior t rap ez iu s or spin otr apeziu s takes o ri gin from the sp in es of the
thora ci c vert ebrae and p asses obliqu ely forward c overin g p art of the la tissimu s ,

to be insert ed on the fas cia of the s capu la ; a ction draws the s cap u la dorsad ,

and c au dad In fron t of this is the middl e tr ap eziu s or acr omiotrapeziu s

. .

Ori gin neu ral sp ines of c e r vi c al and rs t t hora ci c vert ebrae ; insertion me ta
, ,

c r om ion p ro c ess and sp ine of the s c apu l a and fas cia Of the p re c e din g m u s cle ;

a ction draws s capu la dorsad and holds the two s capu lae together The ant erior
, .

trap e ziu s or cl avotr apeziu s is the an t er ior p art of the long mu s cle already des c ribed
as cephal ob rachial Ori gin s up erior nu chal line and median dorsal l ine of neck ;
.
,

p asses obliq u ely ven t rally t o be insert ed on the c lavi c le whi ch is imbedded
o n its inside s u rfa c e ; it is c on t in u o u s wi t h t he cl av ob r achial m u s cle Action .
,

dr aws the clavi cle dorsad an d c ran iad .

c) Leva tor s capu l ae ven tr al is Carefu lly free the three t rap ezi u s mu s cles
. .

Alon g the vent ral border of the acr omiot rap eziu s and app aren tl y c ont inu ou s
with it is seen a at b an dlike mu s cle whi ch p asses ant eriorly divergin g from the
a cr omio tr ap ez iu s and p assin g in t e rnal t o the cl a v o tr ap eziu s whi ch sho u ld be

c ut a c ross Ori gin t ransverse p ro c ess of the at las and o ccipi ta l bone ; in s er
.
,

tion m e t a c rom ion p ro c ess and n ei ghborin g fas cia ; a c tio n draws the s cap ula
, ,

c raniad .

d) Rhomb oideu s Cu t a c ross the middle of t he bell ies of the spin o and
.

a c r om io t r ap ez iu s m u s c les A thi ck mu s c le will be seen be n ea th them ext endin g

.

from the vert eb ral border Of the s capu la to the mid dorsal line ; this is the -

rhomboideu s Origin neu ral sp ines of the vert ebrae and adja c ent ligaments ;
.
,

in sert ion vert ebral border of the s capu la ; a ction draws s capu la dorsad The
, ,
.

m os t ven t ral p or tion of thi s mu s cle is a p ra c ti c all y sep arat e mu s c le the r hom ,

b oideu s capitis which ex t ends as a slender band forward to ori gina t e from the
,

su p erior nu chal line ; inser tion s capu la ; a ction draws the s capu la c ran iad an d
, ,

rota tes it .

e) S pl en iu s This is the large shee t of mu s cle c overin g the ba ck of the ne ck

.

in fron t of the rhom boideu s in t ernal t o the t rap ezii and c rossed by the rhom
,

hoiden s c ap i tis Ori gi n from the mid dorsal line an d fas cia ; i n s ert ion s u p erior
.
,
-
,

nu chal lin e ; a c tion raises or tu rns t he head

,
.
I 4S L AB ORA OR T Y MANUAL FOR VE RT E B R A E ANATO T MY

f) S u pr as pin a tu s .ba On tu rnin g

c k t he ven t ral half of t he c u t a c rom io

t rap e z i u s a s t o u t m u s c le is seen o ccu p yin g t he s u p rasp ino u s fossa of t he s c ap u l a .

O ri gi n whole s u rfa c e of t he s u p rasp in o u s foss a ; i n sert io n g rea t er tu b e rosi t y

, ,

of the hu m eru s next t o the insert ion Of p e ct oralis m in or ; a ction ex t ends the
, ,

hum eru s .

g) D el to ids There
. are t w o del t oids in t h e ca t The c l a v o b r ac hialis already
.

des c ribed is sometim es also c on sidered as a del t oid and is c al l ed the cl avodel toid
by some a u t hori ties The acr omiodel toid is a shor t t hi ck mu s c le p assin g ven
.

trall y from the a c rom ion p ro c ess ; it is ins er t ed on t he s u rfa c e of o t her m u s c les
of the shou lder I t lies lat eral t o the cl avob rachial Action wi t h the next
. .
, .

Tu rn the ap of the a c romiot rap eziu s fo rward a gain P os t e rior t o t he line .

m arki n g t he inser tion of the a c r omio t r ap eziu s and the leva t or s c ap u lae ven t ralis
is a mu s cle whi ch p asses to the upp er arm a c ross the upp er en ds of the m u s c les ,

Of the u pp er arm This is the sp in odel t oid Ori gin sp ine of t he s capu la ;
. .
,

insert ion rid ge ( delt oid rid ge) of t he h um eru s the inser tion bein g c on c eal ed
, ,

by the a c rom iodel t oid ; a ct ion the del t oids rais e and ro t at e t he hu me ru s
, .

h) I nfras pin atu s Cu t a c ross the belly of the la tissim u s dorsi

. B rin g the .

an t erior p art s of the latissimu s and the spin o t rap eziu s forward so as t o exp os e
the p os t erior p art O f the s capu la Two large mu s cles are here seen The an te
. .

rior on e lls the infrasp inou s fossa of the s c apu la from whose su rfa c it t akes

it s ori gin and is insert ed on the grea t er tu berosi t y O f the h um eru s t he in ser tio n ,

b ein g c on c ealed by the del t oids whi c h m ay be cu t a c ross t o see it This m u s cl e .

is the infrasp ina tu s Action ro ta t es the h um eru s

.
,
.

i) Ter es major The s t ou t mu s cle imm edia t ely behind the p re c edin g its
.
,

bers ru nning in the sam e dire ction Origin axill ary border of the s c ap u la an d.
,

fas cia of neighborin g mu s cles ; inser tion in c omm on wi t h la tissimu s dorsi on ,

the m edial s u rfa c e of the h u m eru s ; a c tion ro t a t es the h u m eru s an d lowers it , ,

.

j) T er es m in o r C ar e.f u lly se p ara t e t he in fras p ina t u s fro m t eres m ajor a n d

sep ara t e the form er from t he del t oids and the mu s cles of the u pp er arm On .

the p os t erior border of the infrasp ina tu s and som ewha t c ov e red by it is t he s m all
t eres m i n or O ri gi n ax illary border of t he s c ap u la ; in ser tion grea t er t u b erosi t y ;
.
, ,

a ction assis t s the in fraspina tu s

,
.

k) S u b s capu l ar is P la c e the n ger u n der the an t erior border of the s c ap u la

.

a n d c lear away c o n n e c t ive t iss u e fro m the u nd e r s u rfa c e of the s c a p u la The .

su bs c ap u lar fossa is seen to b e o c cu p ied by a mu s cle the s u bs c ap u laris whi ch , ,

c overs t he inner or m edial s u rfa c e O f t he s c ap u la Ori gin s u bs c ap u lar fossa ; .

,

in sertion lesser tu berosi ty of the h um eru s ; a ction p u lls t he hu m eru s m edially

, ,
.

P os t erior t o the s u bs c ap u laris will be f ou n d p art of t eres m ajor whi ch ex t ends

t o the m edial s u rfa c e of the s c a p u la .

l ) S err a tu s ven tr al is ( an ter ior ) Cu t thro u gh the rhom boideu s c los e t o the
.

vert ebral c olu mn The s capu la then swin gs loose On r aisin g the s c ap u la a
. .
,

large fan shap ed mu s cle is seen extendin g from the v ert ebral border of the
-
1 50 LAB O RAT ORY MA NUAL F OR VE RTE B RATE ANAT OMY

ins ert ion on the t ran sverse p ro c esses Of the v e rt ebra e in c o n ta ct wi th the o ri gi n
of the serra tu s ven t ralis The m ass in the n e ck is divisibl e in t o a m edial
.

biventer cervicis and a la t eral compl exu s All of the ep axial m u s cles ar e very .

c om p lex c onsis tin g of n um ero u s b un dl e s They are a t t a ch ed t o variou s p arts .

of the vert ebra e ribs and h ead an d have the gen eral fu n ct ion Of m ovin g the
, , ,

v e rt ebral c olu mn and relat ed p art s .

8 The mu s cle s of the u ppe r arm

.
S ep ara te th ese m u s c les fro m ea ch o ther
.

a n d id en tify the m .

R abbi t :
a ) Tr iceps br a chii The t ri c ep s is the grea t m u s cl e on the ba ck of the upp er
.
_

a rm . I t c onsis t s of t hree heads whi ch are p ra c ti c a l ly dis t in ct mu s c les F ree .

t he heads The l on g head is the large m ass on t he ba ck of the u pp er arm ; ori gin
.
,

s c ap ul a from the a xillary border ; insert ion ole c ranon The l ater al head is
, , .

on the lat eral s u rfa c e of the u pp er a rm ven t ral t o the p re c edin g ; ori gin grea t er , ,

tu berosi t y of t he h u m eru s The medial head is in c ont a ct wi th the hum e ru s To

. .

see it sp read the o th er two heads ap art an d look de ep be twe en them or cu t throu gh
the middl e of the b elly of the la t eral head ; ori gin alo n g the dorsal s u rfa c e of ,

the h u m eru s All t hree hea ds in sert on the ole c ranon a n d a r e the g rea t ex ten sors
.

O f t he for ea rm .

b ) E xten s or a n tib r achii On the m edial s u rfa c e of the lo n g head Of the t ri

.

c ep s is a fas c ia fro m t he lower p art of whi ch t his m u s c le ori gi n a t es I n s er t ion .

,

ole c ranon ; a ct ion in c omm on wi th the t ri c ep s , .

c) B iceps b r achii On the a n terior s u rfa c e O f the u pp er arm fo u nd b y rem o v

.

in g the insert ions of the p e ct oral m u s c les is a s p in dl e shap ed m u s cle the -

,

bi c ep s Ori gin glen oid fossa ; insert ion ul na and radiu s ; a ction exor of the
.
, , ,

forearm .

d) B r a chia l is This mu s c le is n ext lat eral t o the bi cep s an d in co n t a ct wi t h

.

the h u m e ru s O ri gin ven t ral and la t eral s u rfa c e of the hum eru s ; in sert io n
.
,

a n d a c t ion in c o mm o n wi t h the bi c ep s
, .

Cat :

) a above
Tr iceps b r a chii u nder rabbi t wi t h t
. hree heads
As .

b) E xten s or an tib r achii On the medial side of the lon g head a t hin sh ee t
.
,

O fm u s c l e t akin g ori gin from the la tissim u s dorsi and insert ed on the ole c ran o n ;
,

a c tion in c omm on wi th the t ri c ep s t en din g al so t o ro t a t e the ul na

, ,
.

c) B iceps b r a chii As in the rabbi t ; visible o n ly aft er cu tt in g the i n s e rt io n s

.

of the p ect oral m u s cles Ori gin glenoid fossa ; in sertion radiu s ; a ct ion exor
.
, , ,

O f the forearm .

d) B r achia l is L a t eral to the bi c ep s in c on t a ct wi t h the la t eral head of

.
,

t he t ri c ep s O ri gin la t eral s u rfa c e of the hu m e ru s ; in sert io n u hr a ; a c tio n wi th

.
, , ,

the bi c ep s .

e) An con eu s A sm al l t rian gu lar mu s c le at the elbow jo in t c over ed by

.
,

the dis t al end of the la t eral head of the t ri c ep s whi ch sho u ld be dee ct ed ,
.
 COM P ARA I VE ANA OM T T Y OF T HE M U S CUL AR SYS T E M 1 5
1

Ori gin , dis tal end of hu m eru s ; inser tion la t eral s u rfa c e of the u l n a ; a ction , ,

s t ren gt hens t he elbow join t whi ch it c overs and p ossibly rota t es t he u lna
, , .

m le i
9 Th
. e u s c s O f the th gh ( g
F i . .

R ab b it :
a ) Ten s or fas cia e l a tae The ant erior half of the lat eral s u rfa c e of the
.

thigh is c overed by a t o u gh fas cia or ap one u rosis c all e d the fa s cia l a ta I n the .

dorsal p ar t of t his will be fo und a short mu s cle the t ensor fas c iae lat ae whi ch , ,

is m ore or less c on t inu o u s wi t h adja c en t mu s cles Ori gin ilium ; in s ert ion in .
, ,

t he fas c ia ; a c t ion t i gh t ens t he fas cia, .

sar t orius

AN TERI OR

vastu s

L L
A A
I
R
E D
T E
A M
L

l emo n s

grac il i s

emb ra n o su s

POS TE RI OR
a S US

FI G 43 . .
Cross -
s ec tio n s throu gh the thigh o f A , r ab b it , an d B, ca t to sho w the l oca tio n of the
mus cl es B l a ck
. s p o t s a r e n erv es , s m all c ir cl es , b l oo d vess el s . a, g r ea ter saph en o u s n erv e , a r t ery ,

a n d v ein ; b, p ero n eal n erv e ; c , t ib ial n erv e ; d, s cia tic v ein ; e, femo r al n er ve , ar t ery , a n d vein ; f ,
scia tic

n erve . (A f r om B en sl ey

s P r actical An atomy of the Rab bit, Un iv er sity of T o ron to P r ess .
)

b) B iceps fm e or is S li t op en the fas cia la t a

. fas cia o ccup y . P os t erior t o t he
in g the m iddle p or t ion of t he a eral s rfa e
of the t high is the larg e bi c ep s
l t u c

femoris S ep arat e it from t he mu s cle p os t erior t o it whi c h form s t he p os t erior

.

m argin of t he t hi gh ( sem im em branos u s ) On removin g t he s u rfa c e fas c ia fro m .

the bi c ep s it will be fo u n d di vi sible in t o t wo heads F ollow t hese heads dorsally .

t o t heir ori gins c l earin g away t he t o u gh s u rfa c e fas c ia

, O ri gin of t he smaller .

ant erior head ne u ral sp ines of adja c ent vert ebrae of t he larger p os t erior head
, , ,

is chial tu berosit y ; insert ion by a t en don on the knee cap an d on the fas cia of
,

t he sha nk ; a ction abd u c t or of t he t hi gh and ex o r O f t he shank

, .

c) Gl u teu s maximu s R em ove t he fas cia over t he sa c ral region

. The .

g l u t e u s m a xi m u s is u nder t he fas c ia a t hin mu s c le whose p os t erior p ar,

t is p art ly
c over e d by t he bi c ep s f emoris The mu s cle also has an ant erior p art whi ch is
.

c o n t in u o u s wit h t he t ensor fas c iae la t ae The two p art s of t he gl u t eu s m aximu s .

 I 52 LAB ORATO RY MANUAL FOR VE RT E B RA E ANA OM T T Y

are u ni t ed by an ap oneu rosis Ori gin fas cia of the sa c rum and p art of the ilium ; .
,

insert ion third t ro chant er the sm all p roje c tion below the great t ro chant er ;
, ,

a c tion abdu ct s the thigh

, .

d) Gl u teu s mediu s The larger thi cker mu s cle ant e rior to the p re ceding and
.
,

p ar t ly c overed by i t O ri g in c .res t and an t e r ior p ar,

t of th e ili u m ; inser t ion ,

g rea t er t ro c han t er ; a c t ion like t h e p re c edin g,

When t he t w o g l u t e u s m u.s c les
have been well sep ara t ed the ori gin of the t ensor fas ciae lat ae will b e fou nd on
,
o

the ven t ral ed ge Of the gl u t e u s m edi u s di pp in g deep down t o the iliu m .

e) Va s tu s l a ter al is Cu t t hrou gh the fas c ia la ta and its t ensor by a lon gi

.

tu din al sli t ext e n din g t o the knee Under the fas cia will now be seen the s t ou t
.

vas t u s la t e ralis Origin great er t ro chant er dorsal to the insertion Of t he glu t eu s

.
,

m ediu s .

f ) V a s tu s in te r m e di us This is t h e na m
. e g iven t o wha t a pp ears t o be t h e

p os t erior p ar t of t h e p re c edin g m u s c le I t is p ar t ly se p arable fro .m t h e vas t u s

lat eralis and lies between t his and the an t erior head of the bi c ep s fem oris Origin .
,

g rea t er t ro c han t er and s u rfa c e of t h e fem u r .

g) R e c tu s f em o r is Th i s is t h.e t hin m u s c le whi c h is fold e d over t h e an t e rior

m argin of the t hi gh ; it ext ends on bo t h la t eral an d m edial s u rfa c es O f the thigh .

It ori ginat es on the ilium and t he fas c ia la t a and is fu sed wi t h the bers of
t he t ensor fas ciae lata e The re ct u s fem oris c onsis t s of two p art s the p art j u s t
.
,

des cribed and a se c ond p art whi ch m ay be lo ca t ed as follows S ep arat e the ,

.

re c tu s femo ris from the vas tu s lat eralis to whi c h it will be fou nd s l i gh tly
fu sed c on t in u in g this sep aration t o the m edial side Of the t high sp readin g the
, ,

legs ap art The rs t p art of the re ctu s fem oris m ay be cu t t hrou gh at the
.

middle On the medial side of the t hi gh in ter n al t o the rs t p art Of t he re ctu s

.

femoris and vent ral to the vas tu s lat eralis is the se c ond p art Oft he re ctu s fem oris ,

a cylin dri c al mu s cle Ori gin ili um in front Of the a c e t ab u lu m

.
,
.

h) Va s tu s medial is On t he m edial side of the t hi gh p os t erior t o the rs t

.

p ar t of th e re c t u s fe m oris and n o t very well se p ar a ble fro m i t O ri g in .

,

f em u r .

The re ctu s fem oris and the three vas tu s mu s c les t oge t her c ons t itu t e the
g rea t qu a d r i ce p s f e m o r is m u s c le The o r i g ins O f it s several.
p ar t s have already
been given Insertion on the tibia and the p at ella and the t endon whi c h ext en ds
.
,

over the p at ella ; a ction p owerful ext ensor of the shank ,

.

i) S ar tor iu s The very lon g narrow m u s c le ext endin g like a band alon g
.

the middle of the m edial s u rfa c e of the t hi gh Ori gi n in gu inal l i gam en t ; in s er .

,

t io n t ibia ; a ct ion add u c t or Of the thigh ro t a t or of the t hi g h ex t en sor of the

, , , ,

shank T he sart ori u s lies be tween t he vas tu s medialis and the next mu s c le to
.

be des c ribed and c overs som e large blood vess els

,
.

j) G r ac il is A lar
.
g e t hin m u s c le over th e p os t e rior half of t h e m edial s u r

fa c e of the t high Ori gin pu bi c sym p hysis ; insert ion fas c ia of the dis t al p ort ion
.
, ,

O f the thi gh an d p ro xim al p ort io n of the sha n k : a c t io n add u c t or of t he l eg . .

1 54 LAB ORAT OR Y MANUAL FOR VE RTE B RA E ANAT OM T Y

in sert ion by a s tro n g te n don on the grea ter t ro chan ter O f the fem u r ; a c tio n
, ,

abd u ct or of the t high Alon g the an t erior border Of t his mu s c le the origin of
.

the t e n sor fas c iae lat ae p ass es in t ernally t oward t he ili u m .

f) S a r to r iu s .This m u s c le for m s t h e an t erior m ar g in fro m t he la t eral view

of the t hi gh I t is folded over the m argin and on followin g it t o t he m e di al
.

or inner su rfa c e of t he t hi gh wil l be fou nd t o c over t he an t erior half of t he m edi al

s u rfa c e Ori gin c rest and vent ral border of the iliu m ; in s ert ion p roxim al
.
, , ,

end of the tibia and the p at ella and the fas cia and l igam en t s b etween ; a ction ,

addu ct or and rot at or of the t hi gh and ext ensor of the shank .

g) V as tu s l a ter a l is C u t. t hro u g h t h e fas c ia la t a by a longitu din al slit

ext endin g t o t he p at ella S ep arat e well the sart oriu s from underlying p a rt s
. .

The t ensor fas ciae lat ae is now w ell exp os ed The large s t ou t m u s cle whi ch was .

c overed by the fas cia la t a is t he vas t u s lat e ralis O ri g in great er t ro chan t er and
.
,

su rfa c e Of the femu r .

h) Rectus femor is At it s an t erior m argin the vas tu s la t eralis will be fo u nd

.

p ar t ly se p arable fro m a s t o u t m u s c le lyin g o n it s m edial side and c overed e xt er

nally by the sa rt oriu s The sart oriu s may be cu t a c ross the m iddle The mu s cle
. .

in qu es tion is the re ctu s fem oris Orig in ili um near the a c e t ab u lu m .

, .

i) Vas tu s medial is This is on the m edial side Of the t hi gh p os t erior to

.

the re ct u s femo ri s whi ch its an t erior m argin p art ly c overs I t also is c O

A

v er ed .

ext ernally by the sart oriu s Ori gin femu r .

,
.

j) V a s t u s i n te r m e d iu s On widely sep ara t in g the re ctu s fem oris from the

.

vas tu s lat eralis a mu s cle wil l be se en deep down next t o the shaft Of the fem u r ;
Ori gin s u rfa c e of the femu r
,
.

The re ctu s f em oris and the t hree vas tu s mu s cles are more or less u nit ed to
ea ch ot her and c ons t i tu t e the g reat qu adr iceps femor is mu s cle It s ori gin s have .

been d es c ribed ; all it s p art s are insert ed on t he p a t ella and adja c ent ligaments .

a ction ext ensor of the shank

,
.

k) Gr a cil is . Thi s is the large at mu s cle formin g t he p os t erior half of the

medial s u rfa c e of the t hi gh Ori gin is chial and p u bi c symp hyses ; in ser t ion by
.
,

an ap oneu rosis whi ch p asses t o the t ibia ; a ct io n add uct or of t he l eg ,

.

I) Addu ctor l on gu s a n d addu ctor femor is Cu t t hro u gh the m iddl e of the

.

g ra c ilis and se p ara t e ea c h half fro m t h e u nderlyin g m u s c les The la tt er c on sis t .

of t hree m u s cles p assin g from the median vent ral line t o the f em u r T he m os t .

an t e rior of the three is qu it e small ; thi s is the addu ct or lon gu s Origin p u bis ; .
,

in s ert ion f emu r ; a ct ion wi t h the next The m iddl e mu s c u lar m ass is t he large
, ,
.

addu ct or fem oris ( c orresp on din g t o addu ct or magn u s an d brevis of o t h er mam

mals ) . O ri gi n p u bis ; in sert ion fem u r ; a ct ion addu c t or of t he thi gh
, , ,
.

m) S emimembr a n osu s The large p os t erior p art of the mass whi ch was
.

c overed by the g ra c ilis Origin is c hium ; ins er tio n m edial ep i c on dyle of the
.
, ,

fem u r an d p roxim al en d of the tibia ; a ction ext en sor Of the t hi gh The mu s cle ,
.

is m ore or less divisible in t o two p art s .

 COMPARATI VE ANA OM T Y OF THE MU S CULAR S YSTE M 1 55

n ) S emiten din os us The m ost p os t erior mu s cle Ofthe thi gh pos t erio r to the
.
,

p re c e di n g O ri g in . is c hi al t u berosi t
, y ; insert ion tibia ; a ction exor of the shank , , .

o) Ten u iss imu s Tu rn to the lat eral s u rfa ce of the thigh Cu t throu gh
. .

the middle of the bi c ep s femo ris B enea t h it will b e no t ed a ve r y narrow lo n g

.

mu s cle the t en uis simu s

, O ri gin t ra n sve rse p ro cess of the se c ond c au dal v er t e
.
,

brae in c omm on wi th the cau d ofemoralis ; insert ion on the sam e fas cia as the
, ,

in sertion of the bi c ep s .

On sep ara tin g the bi c ep s from the u nderlyin g m u s cles t hey wil l be revealed
as ext ensions of mu s cles already identied on the m edial s u rfa c e The addu ct or .

fem o ris is seen in c onta ct with the f emu r p ost erior to the vas tu s lat eralis ; the
s emim embranos u s c omes next and the s emit endin osu s is again the mos t c au dal ,

of the thigh mu s cles .

10 The mu s c le s of the shan k

. .

R ab b it :
a ) Tib ial is an ter ior The lateral ( ou t er do rsal ) s ur fac e Of the S hank is
.
,

c overed by the dis t al end of the bi cep s fem oris and fas cia These should be .

removed The most ant erior of the mu s cles Of the lat eral s u rfa c e is the tibialis
.

ant erior Ori gin lat eral c ondyle and tu berosit y Of the tibia ; insert ion se cond
.
, ,

m et a t ars al ; a ction exor of the foot , .

b ) P er on eu s Next dorsal t o the p re cedin g on the lat eral s u r fa c e c onsis tin g

.
,

of a group of several more or less fu sed mu s cles Origin tibia and b u la ; in s er .

,

tion m e t a t arsals ; a ct ion ex or Of the foo t

, ,
.

c) Gas tr ocn emiu s T hi s is the thin b u t broad mu s cle fo rming the cau dal
.

s u rfa ce Of the shan k di visible in t o two nearly sep arat e p ort ions one of whi ch is
, ,

on the lat eral one on the medial su rfa c e of t he shank Ori gin lat eral and m edial
,
.
,

c o n dyles of the femu r and t ibia ; insert io n by a s t ron g t endon the t endon of , ,

Achilles whic h p asses over the heel ( c al c a n eu s) on w hi c h it is in sert ed ; a ct ion

, ,

ext ensor of the foot .

d) S ol eu s Thi s is the mu s cle j u s t int ernal t o t hat p art Of the gas t ro cnemiu s
.

which is on the lat eral su rfa c e Of t he t high O rigin head of the b u la ; insert ion .
,

and a ct ion wit h t he p re ce ding .

e) P l a n tar is Thi s is situ at ed in t ernal to that part of the gas t ro cnemiu s

.

whic h is medial Origin lat eral c ondyle of the femu r ; insertion and a ction
.
,

wi th the p re c edin g .

f) O th er m us c l es of th e s ha n k o
( p t ional ) There are t hree m ore m u s c les of the .

shank ; they lie in c ont a ct with the t ibia They are : the exten s or hall u cis l on gu s .

exp osed on the m edial s u rfa c e of the tibia ; exten s or digitoru m l on gu s c overed by ,

the t ibialis an t erior ; and flex or digitor u m l on gu s be t we en the t ibia an d the sole u s ,

and plan t aris As their n am es imp ly these mu s cles are in s ert ed by lon g
.
,

S len der t endons on t he di gi t s and act t o ex and ext e n d the di gi t s .

I t sho u ld be no t ed t ha t the mu s cles nam ed ext ensor in the shank and foo t
a r e real ly ex o r in t heir a ct io n and t hose n am ed ex o r are ext en sor in t heir
1 56 LAB ORAT ORY MANUAL FOR VE R E B RA E T T ANAT OMY

a ction followin g a cu s t om borrowed from human an at om y The cu s t om arises

, .

from a desire t o retain the nam es app lied to the mu s c les of the fore lim b for the
mu s cles in the c orresp ondin g p osi t ions in the hind lim b Thu s in the su p ine .

p osi t ion of t he forea rm t he e x t e n sors fa c

,
e an t eriorly and t he ex or s p os t eriorly .

S im ilarly the mu s c les on the an t erior side of t he l eg are design at ed ext ensors ,

alt hou gh t hey reall y ex the foo t an d those on the p os t erior side exo rs al thou gh
, , ,

they ext en d t he foo t I n des c ribing the a ct io n the t erm s ex o r an d ext ensor
.
,

are u sed wi th referen c e t o the movem ent p rodu ced and n ot with referenc e to
the p osi t ion O f the mu s cle on the lim b .

Cat :

)
a Tib ial is an ter ior away
. Clean
t he t o u gh fas cia of the S han k and also

the inser t ions of the bi c ep s and the g ra c ilis E x am in e the la t eral ( o u t er) s u rfa c e
.

of the shank The m os t vent ral mu s cle whose ven t ral border is in c ont a ct wi th
.

the t ibia is t he t ibialis an t erior Ori gin p ro x im al p art s Of t ibia an d b u la ;

.
,

inser tion by a s t rong t endon whi ch shou ld be t ra c ed in t o the foo t wh e re it will

be fou nd t o pass obliqu ely to the medial side Of the foot t o be insert ed on the
rst m eta t arsal ; a ct ion exor of the foot ,
.

b ) E xten s or digitor u m l on gu s This is the mu s cle nex t dorsal to t he p re c edin g

.

on the lat eral s u rfa c e of the shank I t is so c losely p la c ed t o the p re c edin g as t o

.

app ear as p art of it b u t the l ine of sep ara tion will be fou nd by a li t tle searc hin g
, .

Ori gin lat eral ep i c ondyle of the femu r ; inser t ion by a s t o u t t endon whi c h if
, ,

followed int o the foot is fou nd t o diverge in t o fou r t endons one Of whi ch is ,

insert ed on ea ch di git ; a ction ext ensors of the digi ts ,

.

c) P er on eu s m u s c les These are next dorsal to the p re c edin g ori gin at in g

.
,

on the b u la There are three of them m ore or less fu sed t o ea ch o ther The
. .

t hree end ea ch in a t endon ; t he t hree t endons p ass over the la t eral s u rfa c e of
the la t eral m alleol u s of the tibia and over the c al c ane u s a n d are in sert ed o n ,

the me t a t arsals and digi ts Act ion ext ensors and ex o r s of t he foo t
.
,
.

d) Gas tr ocn emiu s This is the large mu s cle form in g the p os t erior or c au dal
.

s u rfa c e of the shan k I t is divisible in t o two large p ort ions one on the m edial
.
,

s u rfa c e the o ther on the la t eral s u rfa c e Of the shank The lat eral head is su b
,
.

divisible in t o fou r heads Ori gins from the su rfa ce fas cia the fem u r and the
.
, , ,

t endon and fas c ia of the p lan t aris mu s c le ( see below ) ; insert ion by a s t ro n g ,

t en don whi c h p asses t o the heel bone ( c al c ane u s ) on whi c h it is inser t ed

,
Ac tion .
,

ext ensor of the foot .

e) S ol eu s On c arefu lly sep ara tin g the la t eral head of the gas t ro cnemi u s
.
,

a m u s cle the soleu s will b e fou nd in t e rnal t o it I t is a at mu s c le in c on t a ct wi th

, ,
.

the p ero n eu s mu s c les ven t rally ; it t ap ers ab ru p tly t o a t endon whi c h joins t he
t en don Of the gas t ro cnem i u s Ori gin b u la ; inser t ion c al c an eu s ; a c tion wi th
.
, , ,

the g as t ro cn em i u s of whi c h it is som e t im es c onsidered a p ar t

,
.

aref lly se ara i m dial h ad as ro n em i u s

f) P l an tar is O n
. c u p t n g t he e e O ft h e g t c

a large mu s cle will be fou nd int ernal t o it lyin g bet we en the two heads Of the ,

g a s t ro c ne m i u s w hi c h p ra c t i ca ll y in c lose it I t is f u sed t o a c o n siderable

. e xt e n t
 X THE CO M P ARATIVE ANATO M Y O F THE COE LO M
.

, T HE D I G E S TIVE ,

AND R E S P I RAT O R Y S Y S TE M S

A . THE OR I GN I AND P AR T S OF TH E C OE L OM AND THE ME S E NTE RI E S

1 . O r igin .
The coel o m or b ody cavity o f v er te bra tes is the cavit y o f the hyp omere . I t is
n eve r a t an y s t ag e s eg men ted . T he o u ter wall of t he hyp ome e comes r in co n t ac t wit h the
in n er su rface o f the l ayer of vol un tary mus cl es an d forms the lin in g o f the bo dy wall . This
linin g is kn o wn as the p a r ietal per iton eu m . T he in n e wall s r o f the hyp omer es o f the t wo S id es

co me in c on ta c t in t he m edia n pla ne fol din g ar ou n d the in tes tin e an d in cl osin g the in tes tin e
between their two wall s T he i n er wall s o f the hyp ome e thus b eco me the coverin g l ay r f
. n r e o

the in tes tin e an d o the vis c e a a d a e then n amed the vis cer al per iton eu m o s eros a
r r ,
n Ab o ve
r r .

FI G 44 Diagram s t o sho w the r el a tion s o f cer tain visc era t o the mes en teries A , showin g in tes
. . .

tin e d , s u pp
o r ted b y the d o rsa l m es en t ery c , an d t he hear t g , in cl os ed in t he v en t r al m es en t ery e a n d h .

B , Sho win g the liv er n , in cl o sed in the v en tra l m es en t ery , ar t o f whi ch, t he l ess er o m en tu m l , ex t en ds p
p
b etween the in tes tin e k a n d the liv er , an d art O f which, the fal cifo rm li gam en t p, ex t en d s b et ween the
liv er an d the ven tral b o dy wall C, sho win g rel a tio n O f the in t es tin e q t o the do r sa l m es en t ery c
. .

a , n eu r al tu b e ; b , n o to cho r d ; c , d orsal m esen tery o f the digest ive t r ac t ; d , es o ph ga us ; e, d o rsal m es en

t ery o f the heart or d o r sal m esoca rd ium ; ,
p
g, hear t ; h, v en t r al m esen tery O f t he
f eric ar dia l c avity ;

p
hear t o r v en tr a l m esocar dium ; i , d orsal a o r ta ; j , os t erio r car din al v ein ; k, d u od en um ; l , l ess er om en
p
t um o r he at o duo dena l lig am en t ; m, s erosa o r Vis c eral erito n eum ; n , liv er ; 0 , a rieta l erito n eu m ; p p p
p, fal ciform ligam en t o f the liver ; q sm all in testin e ; r eriton eal cavity ; s b il e d u ct I n A e an d h
, ,
p ,
.
, ,

a n d in B l a n d p form the v en t al m es en tery o f t he dig es tive t rac t whic h in cl o ses t he hea r t a d liv er ;
, ,
r n

in C the m es en t ery is ab s en t
v en tral .
( Fr om P r en tis s an d Arey

s Textb ook f E mbr yol ogy
o ,
c ou rt esy

o f the W B S au n d er s Co m a n y )
. . p .

b
an d el ow the in tes tin e the two walls hyp ome e are in con tact an d f m b
o f the r or a dou l e-wall ed
membr a e which is d es ign ated as a mesen te y
n ,
That p o tion of the mes en tery b tween the r . r e

dors al wall o f the coel o m an d the in tes tin e is call ed the dors al m es en tery ; that b etwee the n

ven t al wall an d the in testin e is the ven tr al mes en tery Diffe en t p or tion s of thes e mes en teries
r . r

receive S p ecial n am es which will b e given in the co ur s e Of the diss ec tion T he do sal mesen tery
, . r

is in tac t fo r its en tir e l en g th in mo s t ve teb a tes b u t the v e t al mes tery very ear ly r r ,
n r en

disapp ear s e cep t in ce tain r egio ns which will b e n o ted l a ter

x r Thes e matters have alr eady been .

1 58
 T HE COE LOM , DI G E STI VE AND RE S P I RATOR Y SYS T EM S
, 1 59

des cri bed in co nn ec tion wit h the s ec tio n o n cho rdate develop men t Sectio IV This '

n s ho ul d

.
,

be r e r ea d an d Figu res 7, 8 , a n d 1 0 s tu died .

( See also Fig 44C; K Figs 8 an d 9 pp
.
, .
,
. 1 4 an d

1 5; W. Fig . :5. P

2. Divis ion s of the co el o m At rs t

. the co el o m con s is ts Of a co n tin u o u s c avity ex ten din g

the en ti re l en gth of the trunk reg ion divided in to two longitudin al halves by the dors al a d , n

ven tral mes en teries With the p ar tial dis app earan ce f the ven tral mes en tery the two
. o ,

halves o f the coelom ar e co n ected ven tral to the in tes tin e ( Fig 44C) In the adul ts of all
n . .

vertebrates the coelom is d ivided in to a t leas t two mp a tmen ts by the formati n o f a p Co r o ar

titio n This p ar tition call ed the tran sverse septum develo ps a t the p os te ior en d of the hear t
.
, , r

an d cu ts o f f the hear t fr om all of the o ther vis cera Th t an sverse s ep tum thus divides th . e r e

coel om in to a s mall an terior co mp a tmen t the p ic dial cavity which co n t ins o n ly the r , er ar a
,

heart an d a very l arge p os terior comp ar tmen t the pl eur oper iton al cavity which con t in s all
, , e , a

o f the o ther vis c era ( s ee Fig 4 5A) The p erica dial cavity in s hes a n d ur odel es is an ter ior
. . r

to the p leurop e ito eal c avity an d the tran svers e s ep tu m in those g ou p s p ass es tran s ve s el y
r n , r r

c oss the bo dy ( Fig 4 5A an d B )

a r I n the An u a an d all ver tebr a tes above An ur a the p e i
. . r r

ca dial c vity has des c en ded p os ter io rly so that it co mes to lie ven tr al to the an te io r p ar t o f
r a r

the p leur Op eriton eal c avity ; the tr ns vers sep tu m then ass umes an obl ique p os ition ( Fig 4 5C) a e . .

In that p or tio n o f the p l eu Op ito n eal cavity which in co ns equ ence of the des c e t Of the
r er n

pe ic di l cavity lies dors l to the p erica dial cavity the l un gs ar e situated This co dition
r ar a a r , . n

o f the coel o m as in F ig u e 4 5C is f o u d in An ura an d rep til es

,
r In birds an d mammals the ,
n .
,

pleurop erito eal cavity is divid d i to an terior an d p os terior p ar ts by the fo matio n f a p r ti

n e n r o a

tio n which d s cen ds f o m the do sal bo dy wal l an d un ites with the tran s v s e s ep tu m ( Fig 4 5D
e r r er .

a nd E ) This p a tition is k own as the bl ique s eptu m in birds an d as the diaphr gm in mamm ls
. r n o a a .

In b ir ds it is non mus cu l ar whil e in mammal s it is in l tra t d with s triated mus cl es de ived f o m

-
e r r

adj c en t myo t omes o f t he bo dy wall

a .

The o b liq u e sep tu m o r d i ph ag m fo rms immedia tely po s terio r to t he l u gs Tha ta r n .

po r tio n of the p l o p e ito eal cavity which is cu t Of

eu r f an ter ior to the o b liq e s p tu m o
r n u e r

d i phr agm cons equ e tly co tains the l u n gs

a I t co n s is ts Of the two pl eu l c vities o r pl e al
n n . ra a ur

s cs e ch in cl os i g a l u n g
a ,
a The two p l eur al cavities ar e comp le tely s ep a ated f o m a h
n . r r e c

o ther t he p e ic a dial c vity con tain in g the hear t b ein g s itu ated in the median lin e b t w e
,
r r a e e n

their ven tral p or tion s That p ar t of the p leuI Op erito eal cavity cu t OfI p os te i r t the
. n r o o

Obl iqu e s ep tunr o di phrag m is ca ll ed the per itoneal o r a bdomin al c avity ; it i c lo

r a the n ses

greate p ar t o f the dig es tive trac t an d th ur ogen ital sys tem

r I t wil l b e s n f o m this e . ee r

acco un t that in b i ds an d mammals the co el om is divided in to fou r co mp

r tm ts the ar en

p ericardi al cavity the two p leur al c vities an d the p eriton eal cavity
,
a ,
.

I t is con ve ien t to s p eak of the vis cera as bein g in cl osed in or con tain ed i the co elo mic
n n

cavities This is n ot howev r a correct exp ess io n s in ce owin g to the f ct th t the vis cera
.
,
e ,
r , ,
a a

ar e cover ed by the vis c er al p e ito n eu m o r s erosa they ar e n o t r ea ll y ins ide o f the c el o m i the
r ,
o n

s ame sense that a chair co ul d b e s id to b e ins ide o f a ro o m They a e ou tside o f it n d h ve a . r a a

the s ame r el at ions to it as if they we e p ushed in to the co el o m car ryi g the co el o mic w ll b efo e r n a r

them To ill s t ate fa rt he o n e can n o t g et in to the in side o f a ten t or a b all oo n by p us hing

. u r r,

ag ains t the w ll ; o e c a ies the ten t o r b all oo n wall befor e him an d always r emains in rea l ity
a n rr

on the o u t id e o f the te t o b a ll o o n
s S imil ly the vis cera ar e ou ts ide Of the co l o m al thou gh
n r . ar ,
e

t hey app ear to b e c o t in ed within its c a vity n a .

B . THE DI G E S TI VE TRA CT AND I TS DE RIVATI VE S

1 . The o rigin o f the d ig e s tiv e trac t .

T he primitive in tes tin e or a r chen ter on , as we
learn ed in the s ec tio n o n de velo p me n t, is p rod uced p rocess o f invagin ation or o ther
b y the
pro cess es in the gas t ru la s tag e O f the e mb ryo . I t is a t r t a s im p l e tube o f en toderm with on e
s
 1 60 LAB ORAT ORY MANUAL F OR VE R E B RA E ANA OM T T T Y
O p enin g to the e terior the bl as top ore which is situ ated at the fu tur e p oste ior e d Of the
x , ,
r n

emb ryo This en to dermal tube p e sis ts as the lin in g o f the adul t digestive tr act an d o f al l f
. r o

its der iva tives ; to it the e a e a dd d o ther l aye s ( con n ec tive t iss e an d muscu l ar l ayer )
r r e r u s

d e iyed f om the s p l c hn ic m esoderm Of the hyp om e evhi h it will b e recall ed fo l ds

r r an ou d r c , ,
ar n

the ar chen tero T he dul t diges tive trac t thus cons is ts o f a thic k wall ed tube comp o sed o f
n. a -
,

a
C
i f
Dia
FI G 45 . .
gr ams t o ill us tra te the divisio n s Of the co el o m in the va rio u s v er t eb ra t e c l a ss es .

The t ran sver s e sept u m an d it s d eriva tives a r e in d ic a ted b y thi ck lin es . A, shes ,
sho win g the div isio n
o fthe c o el om in to p erica r dial c avity a a n d g b y m pl
ea n
eu ro
s o f t hept r a n s ver s e s e tu m d
erit o n eal cavi ty p .

B , u ro del es , simil ar t o shes with t he a dditio n o f the l u n g h whi ch ro j ec ts in to the l eu ro eriton ea l p p p

ca vity g p p
C, t u r tl e ; the er icar dial c avity a ha s d es c en d ed os t eriorl y u n til it lies v en tral t o the an t erio r
.

p a r t O f the pl eu r op eriton eal cavity f

g; t he a n terior ace o f the t ran sv er s e s e tum d has n ow b ec ome ar t p p
O f the wall o f the p
ericar d ia l s a c ; the l u n g his r etr o eriton eal p
D, ea rl y s ta ge O fmamm al s , sho win g the .

f p p
b eg in n in g o f the coel omi c old ( l eu ro erito n ea l m emb ra n e) j d escen din g r om the d o rsa l b ody wall , a n d f
the liver f in cl os ed within t he t ran sv erse s ep tu m d E l a t er s ta ge O f mamm al s sho win g u nion o f the
.
, ,

c o el omic fold j with the t ran sv er se s ep tum d the t wo t o gether fo rmin g t he diaphr agm which s ep a ra tes
,

the pl eu ral cavity k f rom the p erito n ea l ca vity m; the liv er ha s co n s tric ted from the main pa r t o f the
t ran sv er se se pt
m , the c o ns tric tio n b ecomin g the co r on a ry ligam en t i a , ericardial c avity ; b , hear t ;
u . p
c, p
arieta l p
ericardiu m or p
ericar dial sa c ; d , t ra n sv erse s e tu m ; e serosa o f t he liver , th i s b ein g a ar t p ,
p
p
O f the t ran sv erse s e tum o rig in all y ; f, liv er ; g p p
l eu ro eriton eal ca vity; h, l u n g ; i , co ro n ary lig amen t
,

o f the liv er ; j , c oel o mic fold which fo rm s ar t o f the dia hrag m ; k, p p

l eu r al cavity ; l , l eu ro erica r dial p p p
memb ran e or a n terio r c o n tin u a tio n O f t he t ra n sv erse s e tum ; m eriton eal cavity p ,
p .

bo th en todermal an d mesodermal el eme ts the l atter p edomin ating T he an terior an d n ,

r .

p osterior en ds of the d iges tive tube a e fo m d by the in vagin ation of the surface ectod e m
r r e r ,

tha t a t the an te io r en d which b ecom es the lin in g o f the mou th ca vity b ein g cal l ed the s tomo
r

daeu m that a t the p os terior en d linin g the an us the pr octodaeu m

, , ,
.
 I 62 L AB ORA OR T Y MANUAL F OR VE R E B RA E ANA OM T T T Y
c art il agin o us or bon y gil l ar ch al re dy described in conn ec tio with the sk el eton Ce tain b l oo d
,
a n . r

v e el s the
ss o tic
, che l s o t ave s e the vis ceral a ches
a r ar s, a T he viscer al mus cl s al r eady
r r r . e ,

d s ib ed dev el o p f o m the tis s

e cr ,
o f the vis ceral arches
r
D ur in g the ch g
uee f o m t he a q a . an r u

t ic t o the t e
.
s trial hab it o f lif e the vis c eral cl ef ts p o u ches ar ches an d their d eriva tives
rr e , , , ,

u n de g o p r foun d al te
r o tion s ra .

) The tymp n ic cavity and the e ter nal a uditory me tu : B eg in nin g with Amphibia the
e a x a s

r s t vis c e al p o ch fo rms a p ou chl ik e o u tgro wth in the dir tio o f the i tern al ear The e d
r u ec n n . n

o f this o u tgr o wth xp an ds in to a chamb e e the tympa ic c vity o c avity o f the middl e ear T he r, n a r .

s tal k o f the o u tg o wth forms the audit r y or E us tachian t b

r c o n ec tin g the pharyn g eal c avity o u e, n

with the tymp ic c avity I s o me r ep til es i bir ds a n d m mm al s an in v g in a tion o c cur s

an . n ,
n ,
a a

co es p on din g t o the p os ition o f the r s t e ter n al g il l s lit

rr T he b o tt m f this i v agin ation x . o o n

meet the wall o f the tymp nic c avity f ormin g a t the p l ace o f con t c t a membran e of doubl e
s a a

Dia form a tion o f the visc eral p ouches fu rro ws arches an d cl ef ts

FI G 4 7 . .
gr ams t o ill u s tr a te the , , ,
.

A, earl y st a g e, a fr om the w all o f the p ha ry n x b

sho win g the eva gin atio n s which fo rm the visc eral ,

pou ches B l ater s t age ill u stra tin g fo rma tion o f the visceral fu rro ws c b y invagin a tion o f the surface
.
, ,

ec to d erm C l a t er s t ag e showin g form tio n o f the e t ern al g ill slits g b y t he u n io n o f the visc er l
.
,
a x a

pou ches an d fu rows a d b reakin g th ou gh o f the po in ts o f u n ion a viscer l po u ch; b pharyn gea l
r n r .
,
a ,

c vity ; c vis c ral fu rro w ; d

a ,
tymp an ic c a vity ; e thyroid gl a n d ; f in tern al gill slit ; g e t ern al gil l
e , , , ,
x

sl it ; h, v is ceral arch; i, ec t od erm ; j ,

en to d er m .

o rigin , the t ymp an ic membr an e ( ear dru m) . T he p ass age fo rmed b y the in vag in ation is the
ex ter n a l a uditor y mea tu s .

f) Gl andl ike der iva tives o f the vis cer al pouches : I n the a dul ts of l an d v er teb r a t es the
vis c er al p ou ches an d c l e f ts disapp ear , l eaving ,
however c e t in bo di s ar e p ro d u c ed ,
r a e which
by prol iferation o f t he lin in g epithelium o f the vis ceral p ou ches Thes e gl a dlike b dies . n o ,

remn a ts of the vis cer al p u hes ar e very va iab l e in nu mb er a d mo de of

n o c , igin i differen t r n or n

ve t b
r e tes ra Am o g th m m y b e me ti
. n d th true
p l tia t il f m th s dn on e : e e or a a ne on s s. ro e econ

vis cer al p ou ch a d the thymus pa athy ids an d the postbr anchial or pithel ia l bo di s fr om
,
n ,
r ro , e e ,

a var iab l e n u mb er o fvis c e al p ou ches T he th ee l as t amed a g l an dl ik b o di es whi c h p e s is t

r . r n re e r

in the n ec k g ion an d pp
reear t o b el o g to the c a teg o ry o f the g l
a ds o f i tern al s ecr ti n an n e on ,

al tho gh their fu c tio n is un c er tai n

u n Thes e gl an ds ar e p res en t al thou gh imp e fec tly devel op ed
. r

in s hes .

g) T h e tr ac hea an d l u g s : W h en the verteb a tes ado p ted the l an d hab it at the g ill s lit s
n r ,

an d gill s dis app e ed in the a dul t an d their phys iol o g ical r ol e w as t ak en o v er by a n ew o u t

ar ,

g ro w th f o m t he rph a ryn x T his o t g o wth is a tub e which evagin a tes from the midve tr al
. u r n
 THE COE L OM , DIGE S I VE , AND RE S P IR A OR T T Y S YSTE M S 1 63

lin e of the pharyn x . At its dis tal en d it divides in to two s acs . T he tub e is the trachea or win d
pipe, an d the two s acs ath trachea an d lun gs subsequ en tly en l a g e
its en d ar e the lu n gs . Bot r

an d beco me s u b divid ed an d thei s tru ctu re is fu ther comp lic ted by the addition of mes ode
,
r r a r

mal tis su es ( con n ective tiss u e smoo th mu s cl e car til age) to the origi al s imp le en to dermal , , n

l ayer .

h) The swim bl adder : This is an ou tgr owth fro m the diges tive tract o ccu rrin g o n ly in the ,

teleos tome shes I t is a t r s t con n ected with the dig es tive tr ac t g n erall y with the eso phagu s
.
,
e ,

on either do s al or ven tr l s ide by a du ct which p ers is ts t hr ou gho u t l ife in g n o ids

r a T he s wim , a .

b l dder l ies j us t in te al to the dors al bo dy wall o f tel eo s to mes an d is s u pp osed to h ve hydr o

a rn a

s t atic fun c tion s I t is n o t imp obabl e that the s wim bl adder is the fo er u er o fl gs
. r r nn un .

i) The l iver : The liver is a very l arg e g l a d which ar is es by o u tg o wth f ro m the in tes tin e n r .

The s t l k of the ou tg o wth beco mes the b il duct which en l a g es o n the s rf ace o f the l iver to
a r e ,
r u

form the gall bl adde The l iver is s itu ted b etwee the two l aye s o f the ven t al me en t ry
r. a n r r s e

an d is al s o l o ca ted i the t an sv erse s ep tum b u t has g row so l a g e that it p r oj ec ts e te sivel y

n r n r x n

o t o f thes e s tr c tu res ( s ee Fig s 448 an d

u u .

j ) Th e p an cr a s : T h is g l d a ri
e s es by o n e to fo u r g e e all y thr ee o u tg o wths f ro m the
an ,
n r ,
r ,

i tes tin e s l ightl y po s terio to the o ig i o f the l iver

n T he s tal k s o f the ou tg rowths become the
r r n .

p an c r
e a tic duc ts T h e ou t g ro.w ths co m b i e t o fo rm o n e g l an d o f v ar iou s f or m in d iff e e t n r n

ver teb a tes r T he p a c eas may lie b etween the t wo l ayers o f the do s al mes n tery the
. n r r e ,

ven tral m se tery o r b o the n ,

.

k) The yolk s ac: I n the emb ryos o f mos t v er teb ates a yol k ll ed s ac p roj ects fro m the r -

in tes tin e T he wal l o f this s a c is p a t o f the in tes ti al wall which may b e r eg ded a hav i g
. r n ,
ar s n

been st et ched ou t i to a s ac The yolk in the s ac is u sed for the g rowth o f the emb ryo a d
r n . ,
n

at the c o mp l etion o f the emb ryo n ic s tag e the yol k s ac is g r ea tly r edu ced in s i e a d is d rawn z n

in to the in tes tin al wall .

I) The u r in ar y bl adder : This is a lar g e sac g rowin g ou t from the ven tr al wal l n ea r the
posterior te min ation of the in tes tin e I n the embryos of r ep til es bi ds an d ma mmal s it is
r .
,
r ,

rep res en t ed by a l arg e s ac the all an tois which p roj ec ts b eyon d the limits of the bo dy
,
The ,
.

urin ary bl adder is in cl o s ed b etween the two l ayers of the ven tral mes e tery n .

The s tructu r es discussed in this in t odu c tion will b e better un ders too d af ter they hav r e

been stu died in the dissections .

Read als o the accoun ts of the comp arative an atomy of the coel em diges tive an d res p ira , ,

tory sys tems in K W or W d , , .

C . THE COE L O M , DI G E S TIVE ,

AND R E S P I R AT OR Y S Y S TE M S OF E L A S MO B RANCHS

The followi n g dire c tion s app ly chiey to the s pi n y d og sh b u t m ay also be

u sed for the sm oo th d og sh and ska t e the mi n or dif feren ces be tw een these ,

a n im als b ein g sp e ci ed where ne c essary .

I The b o dy wall an d the pl eu r Operiton e al c avity

. M ake a n in cisio n fro m .
-

the lef t side of the c loa c a forward thro u gh the p elvi c girdl e sli gh tly to the lef t
of the midven tral li n e u p to the p e c to ral girdle The in cision will p robably .

cu t thro u gh the ski n rs t a n d sho u ld th en b e e x te n ded thro u gh the mu s c l e lay er .

To assis t in exp osin g the i n terior a tra n svers e i n cision m ay be m ade in the
middle of the la teral body wall on ea ch side I n the ska t e cu t alo n g the l ef t side .

of the cloa ca an d the n alo n g bo th la teral borders of the body cavi ty b u t n o t

anteriorly leavin g the ap of body wal l adhering to the p e ctoral girdle The
,
.

large in ternal cavi ty is the pleu r oper iton eal cavity an d c o n s ti tu tes the gr ea ter
I 64 LAB ORA OR T Y MANUAL FOR VE R TE B RA E ANAT OM T Y

p ar t of oelom I t is lin ed b y a sm oo th shinin g m em bran e the par ietal peri

the c .
,

ton eu m w hi ch adheres c losely to the i n side of the body wall

,
The body w all .

is seen to be c omp osed of three layers : ski n mm cl es and p arie tal p eri to n eum , , .

.

5 The vi s ce ra of the pl e u ro periton e al c avity W ithin the c avi ty are a .
I

n um ber of organ s or viscer a m os t of whi c h belo n g to the di g es tive t ra c t

, At .

the an terior end of the c avi ty is the larg e brow ni sh or g ra yish l iver This con .

sis ts in the sp i n y dogsh of long left and r ight l obes an d a small median lobe in
whi ch is lo c a ted the lon g green ish gal l b l adder I n the smoo th d ogsh the liver .
,

is su bdivided i n to right and l eft l obes the m edian lobe bein g absen t ; the gall ,

b l add er is im bedded in the a n terior p ar t of the lef t lobe a n d is visible as a thin

p la c e in th e liver I n t h
.e ska t e th e liver is c o m p osed of r igh t m e di a n and l ef t , ,

l obes of equ al len g th a n d si z e and the ga l l b l adder is si tu a ted in t he an gles be tween

,

the ri gh t a n d m edia n lobes D orsal to the liver o n the lef t side is the large
.

J shap ed s tomach of ten dis ten ded wi th food

-
, ( I n som e sp e cim en s the s tom a ch
.

is ever ted in to the mou th c avi ty an d shou ld be pu lled ba ck i n to the p l eu rop erito
n eal c avi t y by ex er ti n g a g en tle t ra c tion o n it ) The grea ter p ar t of the s tom a ch
.

c o n sis ts of a lar ge s trai gh t tu be e x t endi n g fro m the an t erior en d of the p le u ro

p eri t o n eal c avi t y t o a p oi n t so m ewha t p os t erior t o t he ends of t h e liver lobes .

I t the n m akes a sharp bend de c reasi n g in diam e ter c o n siderably an d ex te n ds

, ,

an teriorly term i n a ti n g in a c o n s tric tio n the pyl or u s Alo n g the p os terior m argin
, , .

of the ben d of the s t om a c h ( or in the ska te on the dorsal side of the be n d ) is a

dark c olored orga n the s pl een a p ar t of the lymp ha ti c sys tem F ro m the
-
, ,
.

p ylor u s t he shor t in tes tin e e x t ends t o t he a n u s The rs t p ar t of t he i n t es

. ti n e
beyo n d the p ylor u s is c alled the du oden u m I t ex tends for a shor t dis tan c e to .

the ri gh t and then c u rves p os teriorly The b ile du ct a lo n g s tou t du c t is

.
, ,

e asily see n des c e n di n g fro m the gall bladder to en t er the d u ode n u m shor tly

c a u dad of the ben d The bile du ct a c c om pan ied by som e blood vessels ru n s
.

in a s trip of m esen tery I t passes t o the dorsal side of the d u oden al wall an d
.

ru n s for a shor t dis tanc e c au d ad imbedded in the wall before it p en e tra tes in to
the c avi ty of t he d u oden u m I n the cu rve of the d u od en u m rep oses the ven tr al
.

lobe of a whi te gland the pan creas The dors al lobe of the p an c reas whi ch is
, .
,

lo n g an d slen der in the spin y d og sh reac hin g to the sp le en shou ld be lo ca ted by

, ,

raisi n g the s tom a ch and d u oden u m an d looki n g dorsal to them The du ct of the .

p an c reas is so m ewha t di f c u l t t o n d in t he d o g s hes less di

f cu l t in t h e ska t e ; ,

it lies im bedded in the tiss u e of the p an c reas n ear the p os terior m argi n of the
ven t ral lobe an d m ay be exp osed by p i cki n g away the p a n c reas tissu e in this
region B eyo n d the du oden um the i n tes ti n e wid en s considerably an d its
.
,

su rfa ce is m arked by parallel ri n gs Th ese rin gs are the li n es of a t ta chm en t of

.

a sp iral fold the s pir a l val ve whi c h o ccu p ies the in terior of the i n tes tine
, , (A .

p or t ion of th e i n t es ti n e of t en p ro t ru des thro u gh the a n u s a n d sho u ld be p u lled

b a ck in to the coelom by g rasp in g the p or tion in the c avi ty an d ex er ti n g a g en tle

I t has al rea d y b een exp l a in ed that the o rgan s are n ot in rea lity in side o f the coel om .
1'
 I66 LAB ORA OR T Y MANUAL FOR VE RT E B RATE ANA O M T Y

also called the l es ser o men tu m be rou ghly divided in to two por tio n s the
. I t m ay ,

hepa todu oden a l li gam en t ex t e n din g from the liver to the d u odenu m an d c on tain
in g the bile d u c t an d blood vessels an d the gas tr ohepa tic li ga m en t ex ten di n g
fromthe s tom a ch to the liver an d du od en u m an d in the do gshes o ccu p yin g also
,

the a n gle fo rm ed by the be n d of the s tom a c h An o ther r em n a n t of the ven tral

.

m ese n t ery is the s u spen s ory or fa l ciform li gam e n t of the liver This will be .

fou n d a t the a n terior en d of the liver ex ten din g from the midven tral su rface ,

of the liver to the midven tral line of the body wall I n ma tu r e f em ales the .

mou th of the ovidu ct will be n o ti c ed in the fal ciform li gam en t as a fu n n el shap ed -

a p er tu re Af ter seein g the falc iform ligam en t the ap of body wall lef t in the
.

ska te m ay b e cu t off .

E a ch go n ad has a m esen tery whi c h is a sp e cial fold arisi n g from the dorsal
wall very near the ori gin of the dorsal m esen tery The m esen ter y of the ovary .

is ca lled the mes ovar iu m of the tes tis the mes or chiu m I n the c ase of ma tu re
, , .

fem ales ea ch ovidu c t has also a m esen tery the mes otu b ar iu m
, , .

The ant erior end of t he p leu rop eri t oneal c avi t y wil l be fou nd c losed by a
p ar t i t ion t h e
,
tr a n s ver se s ep tu m t h e p os t erior fa c e,
of whi c h is c lo t hed by t h e

p arie t al p eri t one u m The l iver

. is a tt a c hed t o th e se p t u m by t he s t ro n g c or on a r y
li gam ent whi ch is in fa ct a p ort ion of the septum I n its early developm en t the
, , .

liver is in closed in t he t ransverse sep tum and su bsequ en tly be cau se of in c reased , ,

siz e p roje ct s p os t eriorly from the septum whi ch then narrows arou nd the an t erior
,

end of the liver and form s the c oronary ligament ( see Fig 48 p .
, .

The p leu rop eri t oneal c avi t y c omm u ni ca t es wi th the ext erior by m eans of
t he a b domin al por es These will be fou n d one on ea c h side of the an al op en in g
.

( in t h e ska t e p os t erior t o th e an u s ) so m e wha t c on c ealed by a fold of ski n P robe .

in t o t hem and no t e that they lead in t o the p leu rop eri t on eal c avi t y Their .

p pu r ose is obs c u re .

The pl eu op er iton eum is the lin in g o f the hypo mer e an d the p l eu op erit on eal cavity is
r r

the cavity o f the hypomere T he pl eu rop eriton eal membr an e is d ivid ed in to three r eg io s
. n ,

a cco di g to its r el a tio s to o ther s tru c tu es : rs t the par ietal per ito
r n n r m th at p ortio n o f the
,
n eu ,

membran e l i i g the in n er su r face o f the body wall ; secon d the vis er al p r it n u m formin g
n n ,
c e o e ,

the thin ou ter cov erin g o f all o f the vis c e a ; an d third the mes en ter ies o l igamen ts po tio s
r ,
r ,
r n

o f the memb a e e te di g f o m the body wall to the vis c era or fro m o n e vis cu s to an o the
r n x n n r r.

D raw the c on t en ts of the p leu rop eri t oneal c avi t y . M ake a dia gram of an
imaginary se ction t hrou gh the an t erior en d of the p leu rop e ri t on eal cavi t y show ,

in g go n ads s t om a c h and liver an d the rela tio n s of the p leu ro p eri t o n eum t o
, , ,

th em and t o t he body wal l .

Th eri a d ial avi ty M ake i ision t hro h th skin i th m edian

4 .e p c r c an n c u g e . n e

vent ral line from the p ect oral girdle u p t o the low er jaw L eave the girdle .

in t a ct G en t ly s t rip o ff wi th a forc ep s the lay ers of vis ceral mu s cl e u n til you

.

have exp osed a m em brane Thi s mem brane is t he par ietal per icar diu m Slit
. .

op en this mem brane and see t h at it in closes a c avi t y the per icar dial cavity in , ,
 T HE COE L OM , DI G E STI VE , AND RE S P IRATOR Y S Y STE M S 1 67

whi ch t he hea rt is situ a t ed To r eveal this c avit y m ore fully cu t la terally alon g
.

the an t erior fa c e of t he girdl e ke ep in g you r in s t r u m en t in c on ta ct wi t h the irdle

g .

P ort ions of the girdle m ay b e sli c ed away b u t the heart mu s t n ot be in j u red

, .

The p eri cardial c avi t y is t hu s revealed as a c on i cal Cavit y lin ed by the p arietal
p eri c ardi u m and c on t ai n in g t h e heart B y gen t ly lif tin g up t he heart no t e
.

th at it is a tt a c hed only a t it s an t erior and p os t erior ends . At t hese p la c es

the p eri cardial lin in g is dee c t ed from the walls of the e ri c ardial cavi t y and
p
p asses over t h e s u rfa c e of t he h eart as a c ove r in
g layer t
,
he v is ce r a l p car diu m
e r i ,

whi ch is in dis t in g uishably fu sed wi th the heart wall With the hea rt lift ed .

n o t e t h a t the os t erior end of the heart is a fan shap ed chamb er the s in u s

p -
,

ven os u s whose walls are c on t i n u o u s wi t h t he art i t ion t ha t fo m s t h os t ior

, p r e p e r

wal l of the p e ri c ardial cavi t y This p art it ion is the tr an sver se s eptu m whose
.
,

p os t erior fa c e we have alr e ady seen The s eptu m is thu s s een t o be a p art ition
.

whose ant erior wal l is c omp osed of the p ari eta l p eri cardium and whose p os terior
wall of the p ariet al p eri t on eu m The win gs of the s u ms venosu s ar e b u ri ed in
, .

t he t ransverse sep tum ; they c o n s t i tu t e lar e ven o u s c hann els t hro u gh whi ch the
g
venou s blood is retu rned t o the heart .

We m ay now exp lain the form a tion of the t ransv erse septum S in c e the .

heart is si tu at ed in the ven t ral p art of the body it is n ec essa ry in order that the
blood from the dorsal body wall may rea ch t he heart t ha t a b ridge be form ed
p assi n g fro m t he dorsal t o t h e v en t ral side I n
. early e m b ry oni c s t a g es a brid g e
or cyli n d er of mesoderm develop s on ea ch side of the p os t erior en d of the heart
c onne ct in g the s p la n c h n i c m esoderm s u rro u ndin g t he sin u s ven os u s wi t h t he

som ati c m esoderm of the dorsal body wall I n th ese brid ges the main ven ou s .

c ha n n els p ass from t he dorsal body wa l l in t o the sin u s venos u s . L a t er the ,

brid ges en large and n ally fu se wi th ea ch ot h er an d wi th the body wall la t erally ,

formin g a p art i tion the t ransverse sep tu m whi ch thu s cu t s t he heart off from
, ,

the r em ai n der of t he c o elom .In elasm obran chs the f u sio n is n ot q u i t e c omp l et e ,

leavin g an op en in g the per icar dio per iton eal can al dorsal to the sin u s venosu s
,
-
, ,

ext en din g from t he p eri c ardial c avi t y in t o t he p le u rop eri t o n eal c avi t y Thi s
.

op en in g will be s een a t a la t er tim e .

o har n a avi i s r s ira or sys m I s r

5 .T h e m u th an d p y g e l c t e an d t h e e p t y t e . n e t

on e blade of a s c issors in t o t he l ef t c orn er of the m o u t h a n d m ake a c u t t hro u gh

the an gle of t h e jaws ba c k a c ross the ven t ral p ar t s of the gi l l sli t s t hro u gh t he

e t oral ird l e so t ha t m er t o t h e lef t side of t h e s t om a c h A a p is

p c g y ou e g e .

thu s form ed whi c h sho ul d be tu rn ed over to the ri gh t . A large c avi t y is revealed

whi ch at it s p os t erior en d c onverges in t o t he ext remely short es ophagus whi ch
asses at o e i t o t h s t o m a h The eso ha
p g u s m a y be sli g h t ly s l i t t o aid in
p nc n e c .

op enin g the ap .

The n ta erior a r t of t h e c avi t y in c los e d by t h e jaws a n d g i ll ar c hes is t h e

p
mou th or or al cavity I t is bou nded in front by t he u pp er and low er jaws p r o
. ,

vided wi th t e e th .The u pp er an d lower jaws are the two halv es of the rs t or

I 6S LAB ORA OR T Y MANUAL F OR VE RT E B RA E ANAT OM T Y

mandib u lar gil l arch ; the se ct ion of the arch should be iden tied in the cu t
s u rfa c e On the oor of the mou th ba c k of t he t eet h is t he ton gu e; it fo rms a
.

a t sli gh t p roje c t ion whi ch is p ra c t i c al ly im m w a b l e

, ,
I t is s u pp ort ed by the .

s eco n d or hyoid gill arch whi ch shou ld be felt wit hin it and ident ied in se ction

a t t he cu t s u rfa c e ( The t o n gu.e is absen t in t he s ka t e owi n g t o t he r ed u c t ion ,

of the hyoid arch ) .

The p os t e rior and g rea t er p art of the c avit y u nder c onsidera tion is the pharyn x .

I ts wall is p ier c ed by s ix in ter n al gill s l its The rs t of these the s pir acl e is a
.
, ,

rou n ded op en in g in the roof of t he m ou t h imm ediat ely p os t e rior t o the m andib u lar
arch The remain i n g ve gill sli t s are elon ga t ed The int ernal gill slit s c ommu ni
. .

c a t e wi t h larg e c avi t ies t he visceral pou ches whi ch in t u rn op en t o the ext erior
by way of the extern al gil l sl its The tissu e between su c c essive vis c eral p ou ch es is
.

a viscer al ar ch The p ar ts of a vis c eral ar ch shou ld be ex am ined in se ct ion on the

.

left sid e where the arches have been cu t t hro u gh The c en t er of the se ction of .

ea ch vis c eral arc h is form ed by the in ter br an chial s eptu m whi ch ext ends t o the ou t er
su rfa c e of the body where the sp a c es between s u cc essive septa fo rm the ext ernal
,

g ill sli t s O n ea
. c h fa c e of t h e in t erbran c hial se p t um is borne a series of low t hi n ,

folds or p lat es t he br on chial or gill l amen ts The set of l am en t s on on e fa ce

,
.

of the s ep tu m c on s t itu t es a half gill or demibr an ch and the demib ran chs o n the
-

two sides of a sep tu m t o ge t her c ons ti tu t e a whole b ran chia or gil l The gills .
~

are ou tgrow ths of the walls of the vis ceral p ou ches and are c overed wit h en t oderm .

B y ex am inin g all of the sept a de t ermine how many d emib ra n chs are p resen t
an d where t hey are missin g In the inner end of the se ct ion of ea ch vis c eral
.

arch lo cat e the c ross se ction of the c art ilagin ou s gill arch and ext ernal to t his ,

lying in t he sep tum the c art ilages of t he gill r ays Ju s t ext ernal to the se ction
,
.

of the gill arch is the se ction of a blood vessel t he a eren t br an chial vess el
whi ch brin gs ven ou s blood t o the gills At ea ch side of t he gill arch is a se c tio n
'

of ano ther vessel whi c h is inje ct ed wi th a c olored solu tion ; t hese are the eeren t
,

b r an chial vess el s whi ch c arry the aera t ed blood away from the g ills
,
No t e the .

n e bran ches of t hese vessels in t he gill lam en t s The gills are the resp ira t ory .

m e chanism of the animal in whi c h t he blood ob t ains o x yg en an d gives o ff c arbon

diox ide Wa t er is kep t owin g over t he gills by movem en t s of the gill arches
. .

D raw the m o u th cavi ty and p haryn x D raw one vis c eral arc h an d all of .

it s p art s in c ross se ction -

.

D . THE COE L OM, DI G E S TIVE ,

AND RE S P I R AT OR Y S Y S TE MS OF NE CT UR US
Ob t ain a sp e cim en an d p la c e in a wax bo t t om ed disse ct in g pan fas t en in g -
,

it ven t ral side u p by p ins t hrou gh the l egs .

I The vi s c er a of the pl eu r Op er iton eal c avi ty

.
M ake a lon gi tu di n al in c ision .

t hrou gh the body wall a l i t tle t o the lef t of the m edian ven t ral li n e from the lef t
side of the a n u s t hrou gh the p elvi c girdle t o the p e ct oral girdl e Sp read ap art .

the two a p s of the body wall and look wi thin The large c avi t y is as in shes .
, ,
I 70 T
LAB ORA OR Y MAN UAL FOR VE R TE B RATE ANATOM Y

being d esi gn a t ed the gas tr os plen ic ligam en t The lu n g is also a t ta ched to the
.

m eso gas t er by a shor t m esen tery Tha t p ar t of the dorsal mes en t ery s u pp or tin g
.

the sm all i n tes ti n e is the mesen ter y in the lim i te d s en se ; t h a t p ar t s u pp or ti n g

,

the l arge i n t es ti n e is the mes orectu m .

The ven tral m esen t ery is p resen t on ly in the region of the liver an d u rin ary
bladder On e p ar t of it form s the lon g m esen tery ex ten din g be tw een the m edian
.

ven tral li n e of the body wall a n d the m edian lin e of the ven t ral fa c e of the liver .

This is the fal cifor m li gam en t of the liver I t co n tain s a num b er of blood vessels
.

whi c h p ass from the ven tral body wall in to the s u bs tan c e of the liver (wh ere
th ey joi n the hep a ti c p or tal vei n ) I n the free p os terior m argi n of the fal cifo rm
.

ligam e n t the v en tral abdom in al vei n c rosses from the body wall t o the liver .

On raisi n g the liver the ga s tr ohepa tic li gam en t will be see n ex t en di n g from the
a n terior p ar t of the s tom a c h to the dorsal fa c e of the liver I n the region of the .

p a n c reas t he he p a todu o den a l li g a m e n t joi n s the d u od e n u m an d liver a n d in c loses

the gr ea te r p ar t of the p a n c r eas The t ails of the p a n c reas however are sit
.
, ,

u a t ed in the m es en tery of the sm all i n t es ti n e B o th of the li gam e n ts j u s t

.

m en tio n ed ar e p ar ts of the ve n t ral m esen tery The las t p ar t of this m esen tery
.

is fo u n d ex ten di n g from the u rin ary bladder to the m idven t ra l l in e of the body
wall ; thi s is the median ligam en t of the bladder .

E a ch gonad has a m es en tery : mes ova r iu m in the fem ale mes or chiu m in the ,

m ale . The m esen tery of the ovid u c t is the mes otu b ar iu m These sho u ld be .

lo ca ted b y lif ti n g u p the s tr u c tu res in q u es tion .

The fal ciform ligam en t shou ld n o w be severed wi thou t however cu tt in g , ,

thro u gh the v en t ral abdo m i n al v ein The n u m ero u s lobes of the liver app eari n g
.

as s c allop s of the m argin m ay n ow be seen m ore clearly On raisi n g the ri gh t .

side of the liv er the ri gh t lu n g m ay be iden tied dorsal to it I s it of the sam e

, .

len gth as the lef t lu n g ? The sm all ga ll bl adder will b e s een o n the dorsal su rfa c e
of the ri gh t side of the liver I ts d u c t s u rrou n ded b y p a n c reas tissu e m ay b e
.

readily tra c ed to the du od en u m The p an c r eas is said to op en i n to the du od e

.

n u m by a n u m ber of n e d u c t s .

The a n terior en d of the p l eu rop eri to n eal c avi ty is clos ed by a m em bra n e ,

the tr an s ver s e s eptu m The liver is a tta ched to this by the cor on a r y li gam en t
.
,

which is c o n ti n u o u s p os teriorly wi th the fal cifo rm ligam en t T he m ode o f form a .

tio n of the sep tu m a n d the c oro n ary li ga m e n t was des c rib ed in c o n n e c tio n wi th
the do g s h .

3 The p eri c ar d ial c avi ty

.
M ake a m edian ven tral
. i n cision throu gh the
skin from the level of the fore lim bs forward to the lev el of the gills R em ove .

the u n derlyi n g m u s c l e s b it by b it u n til yo u have exp osed a m em bra n e the ,

pa r ieta l per ic a r diu m C

. u t t hro u g h thi s m e m bra n e The p e r ica r d ia l ca vity in
.

whi c h the h ear t is si tu a ted is thu s exp osed Wid en the op eni n g in to the cavi ty
.

by cu tti n g la terally alo n g the an terior m argi n of the p ec toral girdle The .

mu s c les be tw ee n the p e ri cardial c avi ty an d the fore lim bs m ay also be s p li t .

 T HE COE L OM , DI G E STI VE AND RE S P I RATOR Y S Y STE M S
, 1 7 1

The p eri cardial cavi ty is a conical c avi ty li n ed by the p arie tal p eri c ardi um .

On g en tly raisi n g the hear t the p os terior wall of the c avi ty is seen to be form ed
by the tra n sverse sep tu m The t ransverse sep tu m is p ier c ed by two vein s
.

( p
he a t i c sin u ses ) wh i c h e x te n d forward an d e n ter the si n u s ve n os u s the m os t ,

dor sal cham ber of the hear t .

4 The or al c avity an d the pha rynx

.
Op en the m ou th an d cu t thro u gh the
.

an gle of the jaws on each side so that the jaws ca n be s p read wid ely Carry you r .

cu ts ba ck to the gill ar ches The c avi ty th u s e xposed c o n sis ts of a n a n terior

.

or a l cavity a n d a p os t erior pha r yn x .

The oral cavi ty is bou nded ex tern ally by the well develop ed lip s I n tern al .

to the lip s are the sm all c oni c al t ee th There are two rows of t ee th on the roof
.

of the m ou th the p os terior row being the lo n ger E x ternal to the las t t ee th of
, .

the p os t erior row on ea ch side is a sli t o n e of the pos ter ior n a r es or i n t ern al ,

Op en i n g s of the n asal p assa ges P robe i n to o n e of the an terior n ar es an d n o te

.

em ergen c e of the p robe throu gh the p os terior naris The oor of the m o u th .

c avi ty bears a si n gle row of t ee th w h i ch o nc losi n g the m o u th will be fo u n d to t

be tw een the two rows o n the roof P os terior to the tee th is the ton gu e su pp or ted
.

by the s tro n gly d evelop ed hyoid ar ch whi ch shou ld be fel t wi thi n the to n gu e .

The walls of the p har yn x are pier ced by tw o p airs of gill s lits P robe throu gh .

them a n d no t e em er gen c e of the p robe be tween the e x ter n al gills No te the .

c ar tila gi n o u s gill a r ch es s u pp or ti n g the bars (viscer al a rches ) be tween and on

ea ch side of the gill sli ts The walls of the gill sli ts are the viscer al pou ches
. .

Un like the d o g sh they bear no gills the gills bei n g ex ter n al ,

The p haryn geal .

cavi ty n arro w s p os t eriorly i n t o a tu be the es ophagu s ,

B y p assi n g a probe i n to .

the esop hagu s de termi n e tha t it e x t en ds dorsal to the p eri c ardial c avi ty and is
c on tin u o u s wi th the s t om a ch .

5 The laryn x an d the l u ng s

.
I n the oor of the p hary n x m idway be tw een
the se c o n d gills sli t s will be fo u n d a shor t sli t the gl ottis The walls of the glo ttis
, .
,

as shou ld be de term ined by feelin g them wi th a n e for cep s are s tiffen ed by a ,

p air of deli c a t e c ar tila g es t h e a

,
r y ten oid c ar tila g e s These p robably re.
p rese n t
redu ced gill ar ches They are the rs t of the laryn geal car til ages to app ear in
.

the p hylo ge n e ti c series . The s mall cavi ty in to whi ch the glo ttis leads an d whi ch
is i n closed be tween the two aryten oid car tilages is the l aryn x Cu t a c ross the .

g ill sl i t s of th e lef t side so t h a t t h e p hary n g eal c avi t y c a n be o p e n ed m or e wid e ly .

S li t the glo t tis p os teriorly The laryn x is th u s s een to lead i n to a n arrow a t ten ed
.

p assa g e ,
t h e tr ac h e a . The p os t erior end of t his is wide n ed and re c eives t w o

op en in gs P robe in t o ea ch wi th a slen der p robe and n o te em ergen ce of the p robe

.

in t o a l un g The t ra ch ea is t hu s seen to c ommun i ca t e wi th the lu n gs T he air

. .

p assa g e in p ri m i t ive air brea t hi

-
n g v e rt ebra t e s ta kes t he followi n g c o u rse :

an t erior n ares n asal c avi ties p os t e rior n ares oral c avi t y pharyn geal cavi ty
, , , , ,

g lo t t is ,
laryn x t ra,
c hea l u,
n g s The l u n g.s have already bee n n o t ed S li t o p en .

one of th em a n d n o t e the sm oo th in t e rior n ot p rovid ed wi th air sa c s ,

.
 I 72 LAB ORA OR T Y MANUAL FOR VE R E B RA E ANA OM T T T Y

E . THE COE L OM, DI G E S TI VE ,

AND RE S P IR AT OR Y S Y S TE MS OF THE TUR TLE

Ob ta in sp e c im en and p la c e in a disse ctin g p an Sp e cim ens whi c h have

a .

n o t b een in je ct ed sho u ld be em loyed

. p R e m ov e t h e p las t ron .This is do n e by .

sawin g t hrou gh the b ridges on ea ch side lift in g up t he p las t ron and sep arati n g ,

it wi t h a s c alp el from t he su rrou nd in g skin and u nderlyin g membrane .

I The divis ion s an d r e l ation s of the c o el o m

.
The rem oval of the p las tron .

exp oses a membran e t he par ietal per iton eu m whi ch c overs and c on ceals the
, ,

vis cera No t e t ha t the mu s cle layer whi ch is norm ally p resent between the skin
.

and the p eri t oneum is c omp let ely la c kin g in the ven t ral body wa l l of the tu rt le ,

owin g t o the p resen c e of the p las t ron The ven t ral body wall in tu rt les there .

fore c on sis t s of b u t two layers t he skin with its c on tained ex oskelet on and the, ,

p eri t o n e u m O win g . t o t h i s c ir c u m s ta n c e t h e p arie t al p eri t on eu m can be easily

s ep ara t ed from the inside of the body wall a p ro cedu re whi ch is dif cu l t or
'

imp ossible in ot her vert ebra t es No t e however the u su al mu s cles in c onne c .

, ,

tion wi t h t he girdles and lim bs .

In t he m edian line in t he ant erior p art of t he p arie tal p eri t one um short ly
p os t erior t o t h e p e c t oral g irdle is si t u a t ed a t rian g u lar m e m bra n o u s s a c t h e ,

p e r ica r dia l s a c wh i c h in c loses

,
t h e h e ar t I t will be no t i c ed t ha t t h e. hear t is
mu ch more p os t erior in p osi t ion t ha n is the c ase in the shes and Nectu ru s ; in
fa ct t here has o ccu rred a p os t erior des cent of the heart ( and of o ther vis cera as
,

well ) The mem branou s s ac c ov ering the heart is as in the d og sh the parietal
.
, ,

p e r ic a r diu m H ere .it t akes t h e for m of an isola t ed s a c t h e p e r ic a r di al s ac , ,

while in shes a n d Nectu r u s it form ed the linin g of a chamber s u rrou nded by

the body wall The sp a c e between the p eri cardial s a c and the heart is the
.

i di l a vity a p or t ion of the c oelom The ven t ral fa e of t h e i ardial

p e r c ar a c ,
c e p r . c

s a c res t s in t he na tu ral p osi t ion a g ai n s t t he in t ernal s u rfa c e of the p l as t ron ,

while it s dorsal fa ce is fu sed as we shall see t o the p arie t al p erit on eu m Cu t

, ,
.

in t o the ven t ral wall of the p eri cardial sac thu s exp osing the p eri cardial c avi ty ,

and the c on t ain ed h eart .

Two c on spi cu ou s vein s the ven tral a b domin al vein s run longi tu dinally in the
, ,

p arie t al p eri t o n e u m be t ween t h e p eri c ardi a l s a c a n d t h e p elvi c g irdle C u t .

t hro u gh t he p eri t oneum halfway b etween t he hear t and p elvi c gi rdle by a t ran s
verse cu t whi ch severs bo th of the abdomin al veins The large c avi t y t hu s .

exp osed is the pl eu r oper iton eal c avi t y whose walls are lin ed by the p ari eta l ,

p eri t o n eum .

The co el o m o f the t tl e l ik that o f the shes a d Ne t

ur ,
e s co s is t s o f t wo p ar ts a s mall n c ur u ,
n ,

p erica dial cavity an d a mu ch l a g e pl eu op e ito eal cavity W e n o te however that whe eas
r r r r r n .
, ,
r

in the l ow er fo rm s the p e ica d ial c avity is a t e io to the pl e o p e ito ea l cavity a d s epara ted
r r n r r ur r n n

from the l a tte by the tra s ve s e s ep t m in th t tl the p e ica dial ca vity is ve t al to the
r n r u ,
e ur e r r n r

pl eu rop e ito eal cavity a d the t an svers e s ep t m s ems to have disappeared W e may
r n ,
n r u e .

exp l ai thi cha g e as foll o ws

n s n ( See al so . I its po s terio r d es ce t the hear t mu s t n n

n eces sa ily ca ry with it the t an s ve s e sep tum an d the parietal p ericard iu m

r r r r T he l a t ter in .

o rd r to move po s terio l y mu s t s ep a a te f o m the body wall to wh

e r r ich it is a ttached in lower
r
I 74 L AB ORA OR T Y MANUAL FOR VE RTE B RAT E ANATOM Y

P ressbot h lob es of the liver forward a gainst the p ect oral girdl e and look on
the dorsal s u rfa c e of t he l iver The elon gat ed s toma ch will be fou nd cu rvi n g
.

dorsal t o the lat eral border of t he lef t liv er lob e On followin g the s t oma ch .

an t eii orl y t he narrow es ophagu s will be fo u n d en t erin g the s t o m a c h

, The .

st oma ch p asses alon g t he dorsal su rfa c e of the left lobe of the liv er t o t he middle
of whi ch it is atta ched along its en tire lengt h by t he short gas tr ohepatic li gam en t .

Abou t opp osit e t he b rid ge c onne ctin g the two lobes of the liver the s t om a c h ,

p asses insensibly in t o t h e s m all in te s tin e th e rs t p ar t,

of whi c h is t he d u o d en u m .

The du odenum is u n it ed t o the middle of the dorsal s u rfa c e of t he ri gh t lob e

of the liver by the hepatodu oden al ligam en t I n this l i gam en t is sit u a t ed a lon g .

whi t e glan d the pan cr eas Abou t on e qu art er of an in c h ba c k of the ri ght en d

,
.
-

of the p an c reas a pan cr ea tic du ct p asses from the pan c reas in t o the d u od en u m
,

an d m ay be revealed by p i c kin g away the s u bs t an c e of the p an c r eas at

t his p oin t On t he dorsal su rfa ce of the ri gh t lobe of the liver n ear it s l a t
.

eral border is the large gall bl adder which is c onn e ct ed to the du oden um by a
,

short b u t s t ou t bil e du ct B eyond the en t ran ce of the bile d u ct the small

.

in t est ine tu rns sharp ly p os t eriorly and is then thrown in t o a nu mber of c oils .

I n t he c ase of fem ale sp e cim ens it will generally be ne c essary t o rem ove on e of
t he lar ge egg bearin g ovaries at t his p oin t before t he in t es t ine can be c onven ien t ly
-

t ra c ed fart her B y l if t in g t he c oils of the s mall in t es t ine no t e the dor s al mesen tery
.

whi c h a tt a ches it t o the m e dian dorsal line of the c oelom ; this p ar t of the dorsal
mes ent ery is the mesen tery p rop er F ollow the dorsal mesen t ery forward an d
.

no t e the p ortion s of it whi ch su pp ort t he du odenum and t he s t om a c h n amed ,

mes odu oden u m and mes ogas ter resp e ct ively ,

The m esodu odenu m is fu sed t o t he
.

h ep at odu odenal ligamen t so t ha t the two app ear as one b u t the mesogas ter is ,

dis t in ct from the gas t rohep ati c li gamen t Tra c e the sm all in t es t in e p ost eriorly
.
,

n o t in g the c oilin g of t he m esen t ery c orres p ondin g t o t he c oils of the in t es t i n e .

F i n d on t he ri gh t side the e n t ra n c e of the smal l in t es t ine in t o t he l ar ge in tes tin e

or col on At t he j u n c tion of the sm all and large int es t in e is a sli gh t p roje c tion
.
,

t he caecu m The c olon gen erally c rosses the p leu rop eri t on eal cavit y transvers ely
.

an d t hen t u rns p os t e riorly an d ru ns s t rai gh t c au dad t o the c lo ac a No t e t he .

mes ocol on su p p or t in g t he c olon In t he t ransverse p ar t of t he c olon it is fu sed

.

to the m eso gas t er In t he m eso c olo n on t he dorsal side of the c olo n sho r tly
.

beyond the c ae cu m is a ro u n d ed red body the s pl een Tra c e t he c olon t o the ,

.

p la c e wh e re it disa pp ears dorsal t o t h e p elvi c g irdle At t his p oin t .ven t ral t o

t he c olo n will be fo u n d the larg e t hin walled bilobed u r in a ry b l adder

,
-
,
I t is .

g enerally g r ea t ly di s t e n d e d wi t h u rine b u t in so m e s p e c im ens m a y be c o n t ra c t e d

to a sm all m ass The bladder has n o ligame n t s the p eri t on eu m leavin g the
.
,

body wall arou n d t he s t alk of the bladder and p assin g over its su rfa c e t o form
its vis c eral inves tm en t .

Cu t away the p elvi c girdl e by m akin g a cu t t hro u gh ea c h sid e of it wi ththe

bo n e s cissors an d rem ovin g a m edian p ie c e The large in t es tin e wil l b e t ra c ed in t o
.
 T HE COE L OM , DI G E ST I VE AND RE S P I RATORY S YST E M S
, I 75

a tu be , the cl oaca , whi ch p ro c eeds dorsal to the girdle t o the anu s At the p oin t .

of en t ran c e of the large in t es tine int o the c loa c a the u rin ary bladder will be
fou nd att a ched to it s vent ral s u rfa c e by a s t al k On ea ch side of the s talk of .

the bladder in fem ales a large whi t e ovidu ct will also be seen en t erin
, ,
g the .

cloa c a .

The fem ale rep rod u ct ive sys t em may be no t ed a t t his t im e ; tha t of the m ale
is so in c onspi cu ou s t hat it will n o t b e d es c ribed a t this p oint The ovaries ar e .

a p air of larg e sa c like bodies c on t ain i n g in t h eir walls e s of vario s si s

gg u z e .

E a ch ovary is a tt a c hed by it s m esen t ery t he mes ovar iu m t o the dorsal body , ,

wall L at eral t o ea ch ovary ru ns the ovidu ct a large coiled whi t e tu be I t is

.
, .

su pp ort ed by the mes otu b ariu m .

3 .T h e ir t o
re s p a ry s ys te m P ry op en the jaws of the tu r tle an d cu t thro u gh
.

the an gles of the jaws cu t t in g nearer the lower t han the u pp er jaw
, The an t erior .

p or t ion of t he c avi t y thu s revealed is the or al or mou th cavity; the p os t erior

p or t ion t h e p,
h ar yn x The. oral c avi t y is bo u nded by t he jaws whi c h have no

tee th b u t are c lo t hed wi th horny b ea ks of ep iderm al ori gin These beaks ext en d .

as p lat es in t o the m ou th cavi t y In the roof of the mou th c avi t y p ost erior t o
.

the p la t e is a p air of elon g a t ed op eni n gs the pos ter ior n ares ,

P robe t h em an d .

det erm in e that they c onne ct wi th the ant erior nares by p assages whi ch ru n
throu gh the nasal c avi t ies The oor of the mou th c avi t y is o ccup ied by the
.

eshy p oin t ed ton gu e .

In the pha r ynx no t e that nei ther gills nor gill sli t s are p resen t al tho u gh as , ,

we sh al l see shortly the gill arches are rep resen ted B ehind the base of the
, .

t on gu e is an eleva t ion the l ar yn geal pr omin enc e in the c en t er of whi ch is an

, ,

elonga t ed sli t the gl ottis F eel the p air of sm all aryten oid c art il ages on e on ea ch
,
.
,

side of the glottis ; they are de rived from on e of the gill ar ches On ea ch sid e .

of the roof of the pharynx p os t erior t o the mu s c les whi ch c onn e ct the sku ll an d
lower jaw is the op enin g of the au ditory or E u s tachian tu b e a c anal whi ch leads ,

from the p harynx t o the cavi t y of the middl e ear (The op ening may have b een .

des t royed in cu ttin g the jaws ap art ) The au di t ory tu be and also the c avit y of
.

the m id dl e ear are o u t g row ths from the rs t vis c eral p o u ch P os t eriorly t he .

p haryn x narrows in t o t he es o h
p g a u s .

Cu t thro u gh the skin in the m edian ven t ral line of the n e ck and p eel away
the skin fro m n e c k and t hroat S ep ara t e the mu s c les in the m edian line of t he
.

ne ck and n d a t u be s t iffened by rin gs of c art ilage This is the trachea or win d .

pp i e. Tra c e i t forward u n t il it disa pp e ars in t o t he p haryn x In fron t of t his .

la e h hard b d of h h id and by c l eanin g away m u s c l es n d also

'

p c n o t e t e o y t e y o ,

tw o p airs of hor n s of the hyoid ex ten din g p os teriorly The hyoid an d its horn s .

a r e d eriva tives of the s ec o n d third and fo u r th gill ar ches

, , Op en the m o u th an d .

make a cu t aro u nd the la r yn geal p rominen c e freeing it from its p osition on ,

the dorsal s u rfa c e of t he body of the hyoid The s t ru ctu re th u s freed is the
.

la ryn x a n exp an d ed ch amb er at the an t erior end of the t ra c hea

,
F in d in the .
 I 76 LAB ORA OR T Y MANUAL FOR VE R T E B RA E ANA OM T T Y

lat eral wal ls of the la r ynx the two aryt en oid c art ila ges sm all cart ilages s u pp or t ,

in g the two t rian gu lar ap s whi ch in c lose t he glo t t is bet ween t hem P os t erior .

t o t e g lo tt is is a rin g shap ed c art ila g e the cr icoid whi ch is m u c h wid e r on t he

h
-
, ,

vent ral t han on the dorsal side The aryt enoids p robably are rem nan t s of the
.

fth gill arches while the cri c oid is the enl arged rs t c art ilage of t he series of
rin gs in the t ra c hea .

Now t ra c e the t ra chea p os t eriorly No t e the esop hagu s a sof t tu be lying .

, ,

dorsal t o or t o one side of the t ra c hea Find j u st an t e rior to the h eart the p oin t
.

where the t ra c hea bifu r cat es in t o the two b r on chi whi ch p ro ceed to the lu ngs .

R aise t he ri gh t and left lobes of the liver and the s t om a ch and nd dorsal t o
t hem a gains t the c arap a c e a larg e s p on gy organ t he l u n g on ea c h side ,
Tra ce , .

a bron c hu s in t o ea c h l u n g ; it is a c c o m p anied by a pu lm on a ry a rt ery and a

- u lm onary vein S t dy th rela t ion of t h l n t t h le ro eri t o al avi t y

p u . e e u g o e p u p n e c .

No t e tha t the lu ng is in c on ta ct wi t h the inner s u rfa c e of t he c arap a c e and that ,

the p arie tal p eri tone u m p asses over the ven tral s u rfa c e of t he l u n g l eav
in g the l u n g o u t side of t he mem br ane S u c h a rela tion to t he p e ri t oneu m
.

is sp ok en of as retr oper iton eal The p os t erior end of the lu ng however

.
, ,

p roje c t s in t o t he p le u ro p eri t oneal c avi t y and is c lo t hed wi t h t he p eri t one u m .

Cu t op en the l un g and observe it s ext rem ely s p on gy t ext u re ; c ords of c onn e c tive
t iss u e divide the in t erior in t o air s p a c es or a l veol i .

T he p at h followed by the air in resp ira t ion is : ex t ernal nares nasal c avi t ies , ,

in t ernal nares mo u t h c avit y p haryn geal c avi t y glo t tis larynx t ra c hea bro n chi
, , , , , , ,

a n d l u n gs I n the pharyn geal c avi t y the p a t hs of food and air c ross

. .

M ake drawings t o show t he p art s of the di ges tive and resp irat o ry sys t em s .

M ake a dia g ram of a c ross se ction t hro u gh the body a t the level of the heart to
-

show the p eri c ardial and p leu rop eri t oneal cavi ties and m em branes and their
relat ion to the vis c era .

F . THE CO E L O M, DI G E S TI VE , AND R E S P IRA T O R Y S Y S TE MS OF T HE PI G E ON

Ob t ain a sp e cim en and p la c e in a diss ectin g p an The fea thers m u s t be .

rem oved . I t is desirable t ha t t he air sa c s sho u ld have been i n a t ed thro u gh t he

t ra c hea .

r The ora l c avi ty an d the pha ryn x

.
Op en the m o u th widely by c u tti n g .

thro u gh the a n gles of t he jaws An ant erior oral ca vity an d a p os t erior phar yn x
.

are th u s rev eal ed .

a ) Or al ca vity: R oof and oor of t he oral c avi t y are bo u nded la t erally by

horny b ea ks of ep ide rm al ori gin whi ch inc ase the jaws Teet h ar e absen t as in .
,

all livin g birds The roof of the mou th c avi t y bears a p air of elon ga ted pal a tal
.

fol ds wi t h free m b ria t ed margins These p ala tal folds c orresp ond to t he hard
.

ala t e of m a m m als b u t di ffer in t ha t t hey do n o t m ee t in the m edian lin e

p , ,

leavin g here a deep pal atal ssu r e The hard p alat e of m an y birds is th erefore .

a s pl it p ala t e and is n orm ally in the c ondi tion w h i c h in m amm als is the res u l t
I 7S LAB ORATOR Y MANUAL FOR VE R TE B RATE ANATO M Y

an terior p or tio n s fu sed to form the en to glo s sal c ar tila ge P os terior to the en togl o s .

sal car tilage is a median bony p ie ce the bas ihyal P os terior to this is the b asi , .

b r an chial of the third gill ar c h F ro m the p oki t O f j u n c tion of basihyal a n d

.

basibran chi al p roje c ts on ea ch side the lo n g pos ter ior horn O f the hyoid c o n sis tin g ,

of p or tio n s O f the third g ill ar c h On followi n g the p os t erior horn s th ey will

.

be fo u n d to ex t end toward the ears and to be divided in to a p ro xim al longer

p or t io n , th e ce r a to b r a n ch ia l ,
a n d a dis t al shor ter rod the epib r an chia l , .

The c ar tilages of the laryn x m ay nex t be iden tied Cu t aro u nd the laryn geal .

p ro m i n e n c e freei n g i t so tha t i t c a n be drawn ve n t rally A lso free. th e hyoid

app ara tu s from the ven tral su rfa c e O f the larynx The larynx is the exp anded .

c ham ber t h u s revealed a t the t op of the t ra c hea and Op en in g in t o the p hary n geal

c avi ty by way O f the glo t tis B y disse c ti n g in the m ar gi n s of the glo ttis o n
.

ea c h side exp ose a slen der cu rved p ar tially o s si ed a ryten oid car tila ge On
, , , .

the ve n tral side of the laryn x no te the e n larged t ria n gu lar cr icoid car tila g e

F ollow t his aro u nd to the dorsal side where it term i n a tes by m u ch n arrowed
en ds B e tween the two dorsal en ds of the c ri c oid car tila ge is a n o ther m edian
.

c ar tilag e the pr ocr icoid wh

, ,
i ch is in c on ta c t wi th the p os t erior en ds of the a ryte
n o id s . The ary tenoids are derived from c er tai n O f the gill ar ches while the
c ri c oid and p ro cr ic o id car tila ges are the m os t a n terior m odied ri n gs Of
the t ra chea Al tho u gh the lary n x of birds is m orp holo gi c ally the sam e as the
.

laryn x Of o ther ver tebra t es from whi c h so u nds issu e in birds the voi ce is n o t ,

p ro d u c ed in t he laryn x b u t in ano t her p ar t of th e t ra c hea whi c h will be

s een la t er .

E x a m i n e the car tilages of the t ra chea They are broad hard an d bony ven
.
, ,

t r a l l y b u t narrower sof ter a n d c ar tila gino u s in c omp osi tion dorsally

, , ,
There .

is co n sequ e n tly a som ewha t sof t s trip alon g the dorsal side Of the t ra ch ea whi ch
lies a gai n s t the cer vi cal ver tebrae .

Tra c e the eso p ha gu s p os teriorly Shor tly in fron t O f the s tern u m it widen s
.

i n to an e n orm ou s bilobed sac the cr op B irds swallow th eir food whole ; the
,
.

food colle c ts in the crop whi c h is c ap able of grea t dis ten sion an d is p assed on in to
the s t om a ch in s m all q u a n ti ties The c rop shou ld be carefu lly loosen ed on all
.

sides No t all birds possess a crop

. .

T h n t erior i a d th e tor l m l The res ira t ory sys t e m

3 . e a a r s c s an e p c a u s c e s p .

of birds is the m os t rem arkable amo n g ver tebra tes I t c on sis ts n o t o n ly of .

,

t h e l u n g s b u,
t also O f a n u m ber of air sa c s lo c a te d a m o n g th e v is c era a n d O f air
sp a c es in the bon es Th ese air sa c s an d air sp a c es co mmu n i ca t e wi th the lu n gs
.

b y m ea n s of bra n ches O f the bro n c hi This sys tem n o t on ly aids in de crea sing
.

the sp e ci c gravi ty of the bird b u t also in s u res a m ore c o mp l et e exp os u re of the

lu n g tissu e to the air ; for the residu al air is re tai n ed in the air sa c s an d n o t in the
lu n gs as in o th er ver tebra tes an d the air in the l u n gs is c on s equ en tly comp le tel y
,

renewed a t ea ch in sp ir a tion Owing to the deli ca cy Of the air sa cs the s tu den t

.
 THE COE L OM , DI G E ST IVE AND R ES P I RATO R Y S Y STE M S
, 1 79

may n ot be able to lo ca te all Of those m en tioned below b u t som e Of them will ,

be seen They are bes t s tu died in freshly killed sp e cim en s in whi ch th ey have
.

been ina ted throu gh the tra chea .

Dorsal to the c rop in the a n gl e formed by the tyvo halves of the fu r cu la or

wishbon e is si tu a ted the in tercl avicu l ar air sac I ts deli ca te ven tral wall is in
.

c o n ta c t wi th the dorsal wall O f the c rop I t consis ts of two lobes one o n ea ch

.
,

side of the m edian lin e ; in the emb ryo these lobes are sep ara te P u n c tu re the .

in ter clavi cu lar air sa c and nd dorsal to it on ea ch side ano ther s a c the cervica l
, , ,

air sac .

E x tend the m edian ven tral in cision in the skin to the an u s S ep ara te the .

skin from the u n derlying mu s cles on ea ch side Of ches t a n d abdom en The .

g r ea t p e c toral m u s c les are revealed im m edia t ely in t ernal t o the skin a n d o c cu p y

in g the a n gle be tween the keel and the body of the s tern um The pectoral is .

major is the grea t m u s c le covering the en tire s tern um and ex ten di n g to the
hum eru s I t takes origin from the keel of the s ternum the su rfa c e of the body
.
,

of the s ternum and the fu r cu la whi ch will be fo und imbed ded in its an terior
,

border ; its bers co n verge toward the hum eru s and p assing over the shou lder ,

are inser ted on the ou t er and dorsal su rfa c e of the hum eru s The mu s cle sho uld .

be followed to its inser tion Ac tion dep resses the wing Now carefu lly cu t
.
, - .

thro u gh the p e c toralis m ajor sli gh tly to the r ight of the keel Of the s tern um
and alo n g the p os terior margin Of the fu r cu la The mu scle can then be dee c ted
.

and sep ara tes easily from the u n derlying pectoral is min or The large pectoral .

a r ter ies and vein s will p robably be no ti c ed em er gi n g be tween the p e c toral m u s c les

whi ch they supp ly The p e c toralis m inor ori gin a tes from the body O fthe s ternum
.

an d c o n ve r ges t oward the h um eru s On followin g the mu s cle la terally there

.

will be fo u n d be tween it and the p e ct oralis m ajor ano ther air sac the axil l a ry ,

s ac. Cu t i n to the a xilla r y sa c The a n terior wall of this sac is in con ta c t wi th

.

the c ora c oid bo n e a n d la terally on looki n g i n to the sa c the tu berosi ties O f the
,

humeru s will be seen A large op en in g in to the h u m eru s the pn eu ma tic fora men
.
, ,

is readily n o ti c ed ; o n p robin g thi s it will be fou n d to lead in to the in terior Of the

hum eru s I t is the en tran c e to the air spa ce O f the hu m er u s whi c h com mu n i
.

c a tes wi th the a xillary air s a c The axillary air sa c comm u n i ca tes in fron t wi th
.

the in ter clavi cu lar air s a c The p e ctoralis min or m ay now be followed to its
.

in s er tion I t c on verges to a ten don whi ch p asses ven tral to the p os terior end
.

of the cervi cal air s a c an d benea th the shou lder to the dorsal side of the h um eru s
o n whi ch it is i n s er ted To see the in ser tio n tu rn the bird dorsal side u p an d
.

disse ct away the s u p er cial mu scles O f the dorsal side O f the sho u lder The ten .

don O f the p e c to ralis mu s cle owing to its m ode O f inser tion has a pu lley like
, ,
-

a c tion whi ch en abl es the mu s c le to raise the win g Whereas in m amm als all .

of the p e c toral m u s c les ac t toge ther to add u c t the fore lim b in birds the a c tion s ,

of the p e c toralis m ajor and mi n or are Opp osed to ea ch o ther the o n e d ep ressing , ,
1 80 LAB ORA OR T Y MANUAL FOR VE R E B RA E ANA OM T T T Y

the o ther raisi n g the lim b This arran gemen t elimina t es all p ow erfu l mu s c les
.

from the ba ck an d en ables all of the win g m u s cles to take their ori gin from the
rm a n d s t ro n g s tern um .

4

T h e. d ivi ion s o th c oe o a
s f e l m n d the p o s t erior air s a c s
Cu t thr o u gh the .

ven tral abdom in al wall to the r ight Of the m edian li n e B en ea th the skin are .

the thin layers O f abdom i n al m u s c les c orresp o n din g to those of m amm als and ,

in ternal to thi s the p ari etal p eri ton eu m generally im p regn a ted w i th s treaks
of fat Cu t thro u gh this an d ex ten d the in cisio n a n teriorly cu ttin g throu gh
.

the s t er n u m sligh tly to the ri gh t of the k eel keep i n g the s c issors in c o n ta c t wi th ,

the bo n e so as t o avoid i n j u ri n g i n t ern al p ar t s Sp r ead ap ar t the c u t ed ges a n d .

look wi thin .

The sm all cavi t y p os terior to the s tern u m is the per iton eal c avi ty No te .

in it the liver dorsal to the p os te rior en d of the s tern u m the c losely c oiled in tes ,

tin e an d to the lef t the larg e rm gizza r d

,
F ro m the giz z ard a m esen tery ex te n ds
.

to the ve n t ral body wall to the lef t O f the m edia n li n e This may b e desi gna ted .

the ven tr al l iga men t of the giz z ard I t is c o n ti n u o u s a n t eriorly wi th the fal ci
.

for m ligamen t Oi the liver whi ch extends from the median ven tral region Of the
l iver to the m idven tral lin e of the body wall and inn er s u rfa c e of the s t ern u m .

The fal ciform ligam en t an d ven tral li gam en t of the giz z ard to ge ther c ons ti tu te
a p ar ti tio n whi ch divides the p eri to n eal c avi ty i n t o a lar ge ri gh t p or tion a n d

sm aller lef t p or tio n This division is n o t fo u nd in o ther ver tebra tes I n the
. .

p ar ti tio n c o u rses a sm all vein ex ten din g fro m the m ese n t eries in q u es tio n to the
2

liver .

Deec t the p e c toralis m ajor m u s c le on the lef t side of the s ternu m and m ake
a cu t thro u gh the lef t side of the s t ern u m sli gh tly to the lef t of the k e el R emove .

a n d dis card the m edian p ie c e of s t er n um c on tai n in g the keel .

Imm edia tely dorsal to the s tern u m is si tu a ted the deli ca t e per icar dial s ac
c o n tai n i n g the hear t The ven tral wall O f the p eri car dial s ac will p robably have
.

bee n op en ed in cu t ti n g throu gh the s t er n u m The hear t as in the tu r tle has .

, ,

des c ended p os teriorly and a p eri cardial s a c has b een fo rm ed of the a n terior
,

fa ce of the tra n sverse sep tu m an d the pari etal p eri cardi u m as des c ribed in co n
n ec tio n wi th the tu r tle The sp a c e be tw een the p eri cardial s a c an d the hear t
.

is as before the perica r dia l ca vity a p or tion of the c oelom The p eri cardial
, , ,
.

s a c is in c o n ta c t on its ve n t ral s u rfa c e wi th the i n n er s u rfa c e of the s t er n u m an d ,

a n teriorly an d la terally is also in con ta c t wi th the i n n er s u rfa c e Of the body wall .

H en c e o nl y the p os terior p ar t O f the p eri cardial sa c is freed from the body wall
,
.

T his is c o mm on l y c all ed m en t um in tex ts a n d ma n u al s b u t s in ce it is n o t a t a l l

the g rea t er o ,

ho m o l o go u s with the s t ru c tu r e so n am ed in mammal s , it is d es ir a b l e tha t t he n a me b e d r o

'

ed The pp .

p
l igam en t O f the g iz za rd is a m es en tery ecu liar t o b ird s , a n d arises a s a s eco n da ry o u tg r o wth fro m the
s er o sa o f the giz z ard t o the ven tral b o d y wall . I t is p ro b a b ly d u e t o the n eed fo r a ddit io n a l su p p o rt fo r
the heavy gizzard .

7
m ed in m a n u al s the ven t ral a b d o m in al
T his v ein is na vein b u t it d oes n o t a p p ea r to b e ho m o lo go u s

wit h the vein o f t ha t n am e in o ther ver teb ra t es .

 1 82 LABORA ORT Y MANUAL FOR VE RTE BRATE ANAT OM Y

ligam en t B e tw een the two sides Of the du odenal loop s tre tches the mesodu o
.

den u m a p or tion O f the m esen tery Of the in t es tin e

,
I n this is si tu a t ed the pan
.

creas l yi n g be tween the two lim bs O f t he loo p o F ro m a deep dep r ession in the
, .

dor al s u rfa c e of the ri gh t lobe of the liver the two bil e du cts ( there is no gall ,

blad der) em erge and p ass i n to the du oden um The lef t bile du c t is the shor ter .

a n d s to u ter of the two an d en ters the lef t limb of the d u odenu m abo u t half an

in ch beyo n d the gizz ard The m ore slen der righ t bile du ct p asses to the ri gh t
.

lim b of the du oden al loop There are three pan cr ea tic du cts all of whi ch p ass
.
,

from the righ t side of the p an creas in to the ri gh t lim b of the du odenal loop On e .

O fthese arises fro m the a n t erior p ar t of the p an c reas and p asses obliq u ely fo r ward ,

en teri n g the du oden um near the an terior termina tion of the ri gh t limb Of the
loop The o ther two d u cts em erge from the middle of the p an creas and p ass
.

a c ross to the ri gh t lim b of the d u oden u m The d u c ts are generally easily seen
.

by sp readin g o u t the mesen tery .

Tra ce the sm all in tes ti n e p os teriorly from the d u odenu m I t is mu ch c oiled .

a n d su p p or t ed by the mesen ter y which owin g to the sm all sp a c e in t o whi ch the

, ,

i n tes tin e is p a cked is fu sed in many p la ces Near its t ermi na tion the sm all
,
.

in tes ti n e tu rns toward the median lin e widens sli gh tly and t hen r un s str aight
, ,

c au dad in the m edian line At abo u t the middle of the p eri toneal ca vi ty it
.

p asses wi t ho u t enlar g e m en t in t o t h e l a r g e i n te s ti n e T h e p oin t .O f j u n c tion of

large and sm all in tes tine is marked by a p air Of small la teral diver ti cu la the ,

ca eca The large i n tes tine is so shor t as to c ons ti tu te li ttle m ore than a re c tum
.

whi ch soon p asses in to the cl oaca Ow in g to the absen c e of pu bi c and is chial

.

sym p hyses in birds the c loa ca in birds does n o t p ass throu gh the ri n g of the
,

p elvi c g irdle b u t m a y be t ra c ed dire c t ly t o t h e an u s There is n o

. u rinar y blad
der In fem ale sp e cimens the si n gle lef t ovidu ct will p robably be no ted en terin g
.

the lef t side of the cloa ca The si n gle ovary (lef t o n e) is si tu a ted in the an terior
.

p ar t O f th e p eri t oneal c avi t y dorsal , t o t h e g i z z ard .

M ake a drawin g of the di ges tive t ra c t .

The gi zz ard an d p roven t ri cu lu s m ay now be freed from the adja cen t air sa c s
a n d the m ese n t eries On tu rni n g the giz z ard far forward there will be fo u n d
.

be tween the p roven tri cu l u s and the an terior en d Of the righ t lim b Of the d u oden al
loop a ro u n ded red body the s pl een The gi zz ard m ay now be cu t op en alon g
,
.

its p os terior m ar gin The in terior con tains small s ton es and p robably par tially
.

diges ted food No te the ex t rem ely thi ck mu s cu lar wal l s an d the hard horn y
. ,

linin g of the gizz ard Cu t from the giz z ard in to the proven tri cu lu s and n o te the
.

sof t glan du lar walls of the la tt er The gizz ard grin ds u p the food in to sm all
.

p ie c e s t h u,
s t aki n g th e p la c e O f t ee t h and t h e, p rov en t ri c u l u s di g es t s t h e foo d by
means of the di ge s tive u id se c re t ed by the g lan ds in it s walls .

6 The pl e ur al cavitie s an d the ir c o n te n ts

.
The p os terior i n ter media te air
s ac si tu a t ed in the obliqu e se p tu m m ay n o w be pu n c tu red if this has n o t already

been don e The two walls of the sep tum are n ow more clearly observable The
. .
 THE COE L OM , DI G E STIVE , AND R E S P I RAT OR Y SY STE MS 1 83

an terior i n term edia te air sac m ay also be pu n c t u red Ag ains t the dorsal wall .

O f the p leu ral cavi ty on ea c h side will be fo u nd a reddish s o n y a t te n ed or a n

p g g , ,

the l u n g The op en in gs O f som e Of the air sa c s in to the lu ngs will probably b e

.

n o t ed o n so m e s p e c im e n s On cu ttin g in to the lu n g the organ will be fo u nd to

. "

be so lid n o t hollow as in the p re ceding an imals

, , .

The cavi ty in whi ch ea ch lu ng is con tai n ed is as already exp lain ed a pleu ral , ,

c avi ty I t is lin ed by a co l eom ic m em brane the pl eu r a

. As the l u n gs are a t , .

te n ed a gai n s t the dorsal wall of the p leu ral c avi t y the p le u ra p asses over their ,

ven tral fa c es leaving them ou tside so to sp eak The p leu ra fu r therm or e

, , .
, ,

p asses over t he s u rfa c e O f the p eri c ardial sa c a n d li n es the i n n er s u rfa c e of the

body wall .

7 The s yr in x
.
E x a mi n e the p os terior p ar t O f the t ra c hea
. Two slender .

mu s cles the s tern otr a cheal mu s cl es diver g e fro m their i n s er tio n on the ven tral
, ,

su rfa ce Of the tra chea to their origin on the s ternu m These mu s cles shou ld be .

severed The t ra chea disapp ears dorsal to the hear t an d the grea t blood vessels
.

whi ch en t er and leave the hear t These blood vessels mu s t n o t be inj u red
. .

L oosen the t ra c hea a n d p u ll it fo rward The bif u r ca tio n of the tra chea in to the
.

two b r on chi can then be seen dorsal to the hear t Cu t a c ross the bron chi wi th .

a n e s c issors an d draw the t ra chea forward At the p oin t where the tra chea .

forks in to the two bronchi an exp anded cham ber the syrin x is p resen t The , , .

voi ce Of birds iss u es from the syrinx n o t from the la r yn x Alo n g ea c h side of the
, .

tra chea e x te n di n g from the p oin t of inser tion of the s terno t ra cheal mu s cles to
the la teral walls of the syrinx is a mu s c le the in tr in s ic syr in geal mu s cle ,
The .

walls of the syrin x are suppor ted by the las t t ra chea l ri n gs an d the rs t bro n chial
half rin gs The las t two t ra ch eal rings are widely sep ara ted from ea ch o th er
-
.

b u t are co n n e c ted in the m edian ven tral line by m edian p ro cesses M ake a sli t .

in the ven tral wall O f the syrinx and sp read ap ar t the cu t ed ges The cavi ty of .

the syr inx is n am ed the tympan u m I n the dorsal wall O f the tym p a n u m a
.

sli gh t ver ti c al fold is p resen t in the m edian dorsal line ex ten din g forward from ,

the level O f the bifu r c a tion of the tra chea This fold is c alled the semilu n ar .

memb r an e an d its vibra tions are said to p rod u c e the voi ce

,
There are also large .

thi cken in g s in the la teral walls O f the tymp an um whi ch may have som e fu n c tion
in the p rodu c tion O f the voi c e The s terno tra cheal and syri n geal mu s cles do u b t
.

less aid by chan gin g the siz e an d shap e Of the tym p anu m .

G . THE COE L OM, DI G E S TI VE , AND RE S P I R AT OR Y S Y S TE MS OF A MA MMAL

The followi n g dire c tio n s app ly to bo th the rabbi t and the ca t When ever .

the di ffere n c es be twee n the two a n i m als warra n t a sep ara te des c rip tion O f ea ch ,

will be given ; o ther wise they will be des cribed toge ther .

1 The m o u th c avi ty an d the phar ynx

. .

a ) Th e s alivar y gl an ds : The s a liva ry gl an ds are m asses O f g land tis s u e which

are ou tgrow ths of the li n i n g of the mou th cavitv : the s tal k Of the ou tgrow th
1 84 LAB ORA OR T Y MANUAL FOR VE R TE B RAT E ANAT OM Y

rem ain s as the s al ivary du ct The glan ds are si tu a ted am on g the mu s c les O f the
.

h ead an d throa t Th ey shou ld b e lo ca ted a ccordin g to the followin g des c rip tion s
.

a n d their d u c ts follo w ed as far as p ra c ti c abl e . Th ere a r e fo u r airs f alivary

.
p O s

g l ah d s in the rabbi t ve in the c a t ,

The diss ec tio n shou ld b e carried ou t on
.

the same side of the head as tha t o n whic h the m u s c l es were disse c t ed .

The par otid glan d is lo c a ted ven trad a n d c ra n iad of the base Of the p inna
O f the ea r j u s t u n der the skin
,
R em ove the skin from thi s regio n a n d n d the
.

p i n kish g la n d s p read ou t u n der t h e s k i n a n t erior a n d ve n t ral t o th e ea r I ts .

d u c t p asses a c ross the ex tern al su rfa c e O f the m ass eter mu s cle an d p en etra tes
the u pp er lip . The s u b maxil l a ry g lan d has already b een n o ted as a rou ndish m ass
a t the a n gle of the jaw near the p os t erior m argin O f the m asse t er L oosen it .

a n d n d the d u c t sp rin gin g from the in t er n a l s u rfa ce I n the cat the b egi n n i n g
.

o f this d u c t is s u rro u n ded by the elo n ga ted s u b lin gu a l gla n d Tra ce the su b.

m a xillary d u c t fo r ward ; it is a c com p a ni ed in the ca t by the s u blin gu al d u c t .

The d u c t w ill be fou n d to p ass i n tern al to the digas tri c mu s cle This m u s cle .

shou ld be severed The du ct ( or two du cts in the ca t) will then be seen to p ass
.

in tern al to the m ylohyoid mu s c le This in tu rn shou ld be cu t an d the d u c t

.

t ra ced fo r ward . I n the rabbit the sm all a t ten ed su b lin gu al gla n d will soo n be
n o t ed lyin g in the p a th O f the s u b m a x illary d u c t The su b m axillary d u c t (a ccom
.

p a n ie d by th e s u bli n g u al d u c t in t he c a t) si tu a t ed j u s
,
t e x t er n al t o t h e lin ing

of the m o u th c avi ty ru n s forward nearly to the symphysis of the ma n dible and

,

then p en e tra te s the lin in g I n the rabbit the su blin gu al gla n d Op en s in to the
.

mou th c avi ty by several shor t d u c t s w hi ch are imp ra c ti cal to n d The mol ar .

g la n d
, p resen t in th e c a t only is si tu a t
,
ed be tween th e ski n a n d th e e x t ern al s u r

fa c e o i the man dible j u s t in fron t of the masse ter mu s cle I t will be fou nd by
,
.

dee c ting the skin a t this p la c e I t op en s o n to the in side of the cheek by

.

several sm all du c ts im p ra c ti cal to lo ca te The infr a or bital gland in bo th ca t

,
.

a n d rabbi t lies in the oor O f the orbi t a n d will be see n la t er when the eye is

disse c ted .

b ) The mou th ca vity: Cu t thro u gh the skin a t the c or n ers O f the m o u th and
see th a t t he skin is well c lear ed away ov er the a n gles of the jaws Cu t thro u gh.

the m asse te r a n d o th er m u s c l e s a t ta c h ed to the low e r jaw a t the a n gl e of the jaws .

I t sho u ld th en be p o ssibl e to pu ll the lower jaw do wn P r y Op en the m o u th

.
,

g ras p t h e lower jaw a n d

,
ex er t a s t ro n g t ra c t ion T h e jaw wi l
. l g e n erally yi e ld ,

b u t if it does n o t the ram u s O f the m a n dibl e ma y be c u t thro u gh wi th the bo n e

,

s cissors The a n t erior p ar t of the c avi ty thu s reveal ed is the mou th or or al c avi ty
. .

I t is bo u n ded by the lips an d cheeks Tha t p ar t of the ora l c avi ty lyin g be tween
.

the t ee th an d li p s is called the ves tib u l e O f the m o u th The tee th were d es c ribed
.

in c o n n ec tio n wi th the sku ll .

The a n t erior p or tio n of the roof of the oral cavi ty is o ccu p ied by the ha r d
p a l a te
, t h e p os t e rior p ar t by t h e s oft pa l a te w hi c h is ve ry lo n g in the rabbi t .

The differen c e b e twe en the hard an d sof t p ala te sho u ld be de term in ed by feeling .
1 86 T
LAB ORA OR Y MANUAL FOR VE RT E B RAT E ANATOM Y

margin may be con c ealed by a leaf shap ed s t ru c tu re the ep i glo ttis whi ch pr oje c ts
-
, ,

from the base of the to n gu e I fso , the ep iglo ttis shou ld be p ressed ou t Of the
.

way ) The Op en in g form ed by the free bord er of the p ala t e is known as the
.

f
is thmu s of the au ces Thi s op en in g leads in to the c avi ty of the p har ynx S hor tly
. .

an terior to the free border O fthe sof t p ala t e on ea ch sid e is a pit the ton s il l ar foss a , ,

whi ch co n tain s a small m ass O f lym p hoid tissu e the pal a tin e ton s il The t on sillar
, .

fossa is bou n ded in fron t and behin d by low folds an a n terior gl oss opal atin e ,

ar ch a n d a p os t erior pha r yn gopal a tin e a r ch No w sli t the sof t p ala te forward

.

along its m edian lin e A cavi ty the n as opharyn x a p ar t O f the p haryn x is

.
, , ,

revea led dorsal to the so f t palat e At the a n terior en d O f the nasop haryn x
.

are the two pos ter ior n a r es or choan ae the in t ern al en ds Of the n asa l p assa ges
, .

P os terior to them o n the la t eral wall of the naso p hary n x will be no t ed a p air of
O bliq u e sli ts ; they are the Op e n in g s O f the au ditor y or E u s ta chian tu b es c anals ,

w hi c h c onne c t the p harynx wi th the cavi ty O f the m iddle ear .

The p harynx narrows p os teriorly in to the es ophagu s An teriorly to the .

e n t ra n c e i n to the esop hagu s is si tu a t ed the en tran c e in to the resp ira tory tra c t .

This en tran c e is g uarded by a p roje ctin g p ro c ess the epigl ottis whi ch if n o t , ,

already iden tied will be seen on pu lling the tongu e well for ward In the .

p hary n x th e p a ths for food and air are c rossed ( as is the ca se in all O f t he air
brea thing ver tebra tes) I t will be no ted however that owin g to the fo rm a tio n
~

.
, ,

O f the p ala t e and the c onseq u e n t p os t erior m i g ra t ion of the p os t erior n ar es ,

the a ir no lo n ger e n ters the oral cavi ty as is the c ase in Amphib ia and m os t r ep
tiles b u t p ro c eeds dire c tly in to the p haryn x .

2 The hyoi d apparatu s the l a ryn x the tr ache a an d the e s o pha g u s

.
, , , .

P ress the to n gu e dorsally a gains t the lower jaw and n d o n its ex tern al s u rfa c e
a t its base a bone the body of the hyoid
,
This is a s tou t bone in the rabbi t a
.
,

n arrow bar in the ca t . Clear away m u s c les from its su rfa c e so as to r eveal it
and the two horn s or corn u a whi ch ex ten d from its sides I n the rabb it the .

horns are shor t p ro c esses whi ch are c onne c ted by slen der t en din o u s m u s cl es
wi th the j u gu lar p ro c ess of the o ccip i tal bone I n the c at the a n terior hor n is
.

lo n g an d slen der an d consis ts Of a chai n of fo u r bony p ie c es the las t Of w hi c h ,

ar ti cu la tes wi th the tymp a n i c b u lla ; the p os terior horn is shor t an d is u n i ted

to the laryn x . The hyoid and its horns are derived in m amm als fro m the
se co n d an d thir d gill ar ches The hyoid su pp or ts the b ase O f the ton gu e an d
.

s erves for the ori gin a n d inser tion of m u s c les .

In the m edia n ven tral line p os t erior to the body of the hyoid is a c ham ber
wi th ca r tila gin ou s walls the l aryn x or voi c e b ox whi ch cons ti tu tes the p rojec
, ,

tion in the throa t p op u larly k n own as Adam s app le B y m akin g a c u t thro u gh

.

the bas e of the ton gu e an d ge n tly severin g the mu s cle a t ta chm e n t s the larynx ,

may be freed and lif t ed forward At the top of the la r ynx is a large Op eni n g
. ,

the gl ottis from whose ven t ra l m ar gin the e p i glo ttis p roje c t s
,
Dorsal to the .

g lo tt is a n d bo u nd wi th it by m u s c les is ano ther Op enin g g enerally c olla p sed a n d ,

 T HE COE L OM , DI G E STIVE , AND RE S PI RATORY S Y STE MS 1 87

c on c eal ed from view by por tions of the la r ynx This op en in g shou ld be lo ca t ed.

by p robi n g ; the p robe will be fo und to en ter a sof t tu be whi ch p ro ceeds pos t e
rior l y dorsal to the larynx This tu be is the es ophagu s
.
.

The s t ru c tu re of the larynx shou ld now be examin ed in de tail The ven tral .

wall O f the laryn x is su pp or ted by a large shield shap ed car tila ge the thyr oid -
,

car tila g e A shor t dis tan c e p os terior to t his is the c ri coid car tilage which
.
,

form s a ri n g arou nd the larynx The dorsal rim Of the glo ttis be tween the glo ttis
.

an d the o p en in g to the eso p ha gu s is s u pp or t ed by a air of roj tin ar t ila es

p p e c g c g ,

the ar yten oids On look ing in to the glo t tis a p air of folds the vocal cor ds will
.
, ,

be seen ex tendi n g from the ary ten oid car tilages to the thyr oid car tilage They .

n early o c c l u de t he o p e n i n g I n the ca t in addi tio n to these tru e vo cal c ords

.
, ,

there is a p air of fal s e vo cal cords si tu a ted la teral to the fo r m er an d ex tendin g

,

from the tip s Of the ar y tenoid ca r tila ges to the base of the ep iglo ttis I t will .

b e n o t ed th a t the vo cal cords are n o t c ords b u t folds of the la t eral wall O f the
laryn x On disse c ting away the esophagu s from the dorsal side of the laryn x
.

the dorsal side O f the c ri coid car tila ge will be exp osed I t is mu ch broader than
.

the ven tral side B y c leanin g away the mu c o u s m em bra n e c overin g it the two
.
,

aryten oid car tila ges whi ch res t on the an terior ex tremi ty of the dorsal p ar t of
the c ri coid will be exp osed .

F rom the laryn x the tra chea or win dp ip e p ro ceeds p os te riorly I ts walls .

are s tiffen ed by car tilaginou s rin gs whi ch are in comp l ete dorsally leavi n g a
, ,

sof t s trip in the dorsal wall O f the t ra c hea in to whi ch the esop ha gu s ts On .

ea ch side O f the t ra ch ea lyin g a gains t the tra chea and in ternal to the mu s cles is
a a t t e n ed elo n ga t ed body o n e of the lobes O f the thyr oid gla n d
,
The an terior .

en d of ea ch lobe is a t a level wi th t he c ri coid c ar tila ge The c au dal en ds of the

.

two lobes are c o n n e c t ed by a m edia n p or tio n the is thmu s whi ch c ross es the , ,

ven tral side O f the t rachea The t ra chea is n o t to be tra c ed far ther p os teriorly
.

a t this tim e .

The tru nk of mamm als is divided

3 The ple u r al an d p eri c ar d i al cavi tie s
. .

in to an an terior thora c i c r egion and a p os terior abdom inal region E a ch of these .

regions con tain s ca vi ties whi ch are p or tions O f the c oelom The thora ci c region .

has three coelom i c c avi ties the two pl eu r al cavi ties la terally lo ca ted an d the
, , ,

m edian per ica r dia l c avi ty si tu a t ed be tween the two p leu ral ca vi ties
,
.

Wi th the bon e s cissors make a cu t thro u gh the ribs on e half in ch to the lef t -

of the s ternum ex tendin g the cu t the len g th of the s ternum At ea ch en d of

,
.

t his cu t la t erally and dorsally be tween two adja c en t ribs a t ri gh t a n gles to the
,

rs t cu t I n this way a ap is fo rm ed in the ches t wall Op en the ap an d b end

. .

it dorsally so tha t yo u can look wi thin

i The cavi ty thu s revealed is the left
.

p l eu ra l ca vi ty or pl eu r a l s a c as it is O
,
f t en c alled ; a sim ilar s a c e xi s t s on t h e ri g h t
side The p leu ral sa c co n ta ins the sof t sp ongy l u n g In the median region
. .

u nder the s t ern um lies the lar g e hear t No te the deli ca t e p ar ti tio n whi ch
.

s tre tches from the hear t to the ven t ral m edia n line This p ar ti tion is called the .
1 88 LAB ORAT OR Y MANUAL F OR VE RT E B RATE ANA OM T Y

media s tin a l se ptu m o sis t s of the tw o m edial walls of the ri gh t an d lef t

. It c n

p le u ral sa c s in c o n t a c t wi t h ea c h o t h e r A t t he level of the h ear t the two walls

.

sep ara t e so tha t the hea r t and its p eri c ardial sa c are in clos ed b etwee n them .

Thissp a c e be tween the two walls of the m edias tinal sep tu m is called the media s
tin u m .The p os terior wall of the p l eu ral sa c is form ed by a m u s cu lar dom e
shap ed p art i tion the diaphr agm The p l eu ral sa c is lin ed by a sm oo th m ois t
,
.

membra n e the pleu r a ,

The p leu ra is divided in to par ietal and viscer al p ar ts
. .

The p arie tal p leu ra lin es the i n side of the p leu ral cavi ty c overs the an terior fa ce ,

of the diap hra gm and to gether w i th the m edial wall Of the o ther p leu ral sa c forms
,

the m edias ti n al s ep tum The vis ceral p leu ra is tha t p art Of the p leu ra whi ch
.

p asses over th e s u rfa c e of the lu n g to whi c h it is indis tin gu ishably fu sed .

E x am in e the lef t lu n g I t is a s of t sp o n gy or ga n divid ed in to three lobes a

.
,

sm aller an ter ior an d larger middle and pos terior l obes The an terior lobe is
, , .

qu i te sm all in the rabbi t The large p os terior lobe ts ver y nea tly on the convex
.

su rfa c e of the diap hra gm Cu t in to the lu n g ; it appears solid b u t is really com

.

p osed O f inn u m erable m in u te air c ells -

.

Now carefu lly cu t throu gh the m edias tin al sep tum ven tral to the hear t and
look in to the ri gh t p leu ral cavi ty The diap hra gm may be sli t alo n g its l ef t side
.

so as to fa c ili ta t e the sp rea din g ap ar t O f the thora ci c walls The ri gh t p leu ral .

cavi ty is sim ilar to the lef t c avi ty I t co n tain s the ri gh t l u n g

. The ri gh t lu n g .

is som ewha t larger than the lef t lu n g I t is divided in to a n terior m iddle an d

.
, ,

p os t erior lobes The lar

.
g e p os t erior lobe is s u bdivid e d i n t o tw o l ob u l es a m edia l ,

and a l a ter al The m edial lob u le p roje c ts in to a p o cke t form ed by a sp e cial

.

dorsally dire c t ed fold of the m edias tinal sep tum This fold the caval fold has .
, ,

the fu n c tio n O f s u pp or tin g a lar ge vein the pos tcaval vein whi ch as c en ds fro m
, ,

the liv er to the hear t a n d will be fo u n d in c losed in the free dorsal m argin O f the
c aval fold .

E x am i n e the hear t an d the p eri cardial s a c The p eri c ardial sa c or p arie tal
.

p e ri c ardi u m is a s a c of t hi n tiss u e in c losin g the hear t b u t n o t a t ta c hed to it ex c ep t

a t the a n t erior en d where the grea t v essels en t er and leave the h ear t The hear t .

is fre ely m ovable in side of the p eri c ardial s a c The narrow sp a ce be twe en the
.

p eri c ardial s a c a n d the hear t is the per ica r dial ca vity a p or tion of the c oelo m , .

Cu t thro u gh the p eri c ardial s a c so as to exp ose the hear t The s u rfa ce of the .

hear t is in ves ted by a thin m em brane the viscer al per icar diu m in sep arably , ,

adheren t to the hear t wall The vis c eral p eri cardium is con tinu ou s wi th the
.

p eri c ardial sa c a t t h e a n t erior en d where the blood vessels e n t er a n d leave the

h ear t As the hea r t wi th its p eri cardial sa c is situ a t ed in the m edias tinum it is
.
,

eviden t tha t there are thre e coelomi c layers su rrou n din g the hea r t : the vis c eral
p e ri c ardi u m c losely adhere n t t o the hear t wall th e p arie tal p e,
ri c ardi u m or
p e ri c ardial sa c s ep ara t ed from the hea r t by the p eri c ardial c avi ty a n d the p arie tal ,

p l eu ra of t h e m e d i as t inal se p t u m whi c h is ,
c losely f u sed to t he p eri cardial sac

( F ig .
49 3 , p .
1 90 LABORA OR T Y MANUAL T
FOR VE R E BR A E ANATO M T Y

ca vi ty, is expo sed I ts ant erior wall is form ed by the c on cavely ar ched diap hra gm
.

whi c h comp le tely sep ara tes the p eri ton eal from the p leu ral cavi ties P os terior .

to the diap hra gm and shap ed so as to t the co rkav e s u rfa c e of the diap hra gm

is the large lobed l iver gen erally grayish brown in p reserved sp e cim en s P os
, , .

terior to the liver the p eri to n eal cavi ty is ll ed by the coils of the in tes ti n e In .

the ca t the i n t es ti n e is covered ve n t rally by a t hin m em brane imp regn a t ed wi th

s treaks of fa t the grea ter omen tu m This m em brane is p res en t also in the rab b it
,
.
,

b u t is very m u ch sm aller a n d less c onsp i cu o u s At the p os terior end of the p er

.

ito n ea l c avi t y m ay be n o ted the p ea r shap ed u r in ary bl adder g en erally dis ten ded
-
,

wi th u id On raisin g the liver and looking dorsally and to the lef t of it will be
.

fou n d the s tomach wi th the s pleen a tta ch ed to its lef t border On the dorsal wall .

of the p eri to n eal cavi ty a t abou t the level of the p os t erior en ds of the liver lobes
are the kidn eys rou n d orga n s ; to see them gen tly lif t the coils O f the in tes tin e
, , .

I n fem ale sp e c im ens esp e c ially those whi ch are p re gn an t the horn s O f the u ter u s
, ,

will be n o ted as a tu be on ea ch side in the p os terior p ar t of the p eri to n ea l cavi ty .

The p eri to n eal cavi ty is lin ed by a membran e the per iton eu m As in all , .

c oelom a t e a n im als tha t po r tion of the m em brane on the inside of the body wall
,

is the par ietal p eri toneum I n bo th dorsal and ven tral regions the p eri toneu m
.

is de ec ted from the body wall and p asses over the su rfa c e of the vis c era form ing ,

a coverin g layer the vis cer al p eri toneu m or ser osa for all of the vis cera x I n
, ,
r

p assi n g to a n d fro m t he body wall t o the vis c era the p eri to n eum form s do u ble ,

walled m em bran es the mes en ter ies or ligamen ts The dors al m esen tery is p resen t
, .

in ta c t in mamm als and the ven tr al m esen tery p ersis ts in the region of the liver
,

a n d u ri n ary bladder as in o ther ver t ebra t es .

E x am i n e the s tom a ch r s t by raisin g the liver and p ressin g it cra n iad

,
The .

ex p os u re of the s to m a c h is fa cili ta t ed by sli ttin g the diap hra gm o n the lef t side .

The s tom a ch is a large an d rou nded or gan in the rabbi t sm aller and m ore elon ga ted ,

in the ca t . Find where the esop ha gu s em erges from the diap hra gm and en ters
the a n terior s u rfa c e O f the s to m a ch The area O f j u n c tion Of the s tom a ch an d
.

esop ha gu s is called the car dia; and the region O fthe s tom a ch adja cen t to the j u n c
t io n the car diac en d of the s to m a ch
,
T he shor ter sli gh tly c o n cave a n t erior
.
,

su rfa c e O f the s tom a ch from the cardia to the p yloru s is the lesser cu rva ture; the
larger c o n vex p os terior su rfa c e the greater cu rvatu re The sa c like b u lge of
,
.

the s to m a c h to the lef t Of the c ard ia is known as the fu n du s ; the rem ai n der
of the s tom a ch the b ody At the righ t the s tom a ch p asses in to the sm all
,
.

in tes ti n e the p oin t of j u n c tion k n own as the pyl oru s bei n g m arked by a c o n
, , ,

s tri c tion b eyo n d whi ch the sm all in tes tin e m akes an abru p t bend Alo n g the lef t
,
.

side of the s tom a ch lies the s pleen a ra ther large organ in the ca t b u t smaller in
, ,

the rabbi t .

The rela tion s of the s tom a ch to the p e ri to n eu m are som ewha t comp li ca t ed .

R aise the fu n d u s an d no te the mes ogas ter ex t e n din g fro m the dorsal wall t o the
s tom a ch Onl y a sm all p or tio n of the m esogas ter p asses dire c tly to the s toma ch;
.
 T HE COE LOM , DI G E STI VE AND RE S P IRATOR Y S Y STE M S
, 1 9 1

t he g rea ter p ar t Ofit rs t des cends p os teriorly formin g a b ag the gr eater omen tu m , , .

This is a ve ry lar ge an d ex ten sive shee t in the cat c overi n g the in tes tin e ven trally ,

as no ted above I n the rab b it it is a shor t m embran e dep en den t from the grea ter
.

cu rva tu re of the s t o m a c h The grea ter omen tu m is to be thou gh t Of as form ed

.
.

in the followin g way Supp ose one sho u ld gras p the m eso gas t er and p u ll it o s
p
.

ter io r l y drawin g it in to a sa c
, Su ch a s a c wo u ld have two walls ea ch do u ble
.
, ,

i e comp osed of the two layers Of the meso gas ter ; the sa c wo u ld c o n ta in a
. .
,

c avi ty whi c h is know n as the les s er per iton eal s a c a n d wo u ld o en a n t eriorly t his
p (
Op e n in g wil l be seen la t er) (see F ig 50 p B y manipu la tin g the o m en tum
.
,

de termin e that it c onsis ts of two sep ara te walls Havin g form ed the grea ter .

omen tum the m eso gas ter re tu rns to the s tom a ch wall and p asses o n to the s toma ch
along the gr ea ter cu rva tu re The sp leen is in closed in the ven tral wall Of the
.

g rea t o m e n tu m j u s t before the la tter p asses to the s toma ch The p or tion of the .

g rea t o m e n t u m be t ween the sp leen and the s tom a ch is called the gas tr ospl en ic
( g
or a s tr ol ie n a l ) g en t P os terior to the sp leen near the lef t kidney a se cond
li a m .
, ,

ary fu sion the gas tr ocol ic ligam en t has form ed be tween the m eso ga s ter and the
, ,

mesen tery O f the in t es tin e ( see F ig . o

The grea ter om en tum owes its origin in p ar t to the ro ta tion O f the s toma ch .

The line of a tta c hm ent O f the om en tum to the grea ter cu rva tu re is the ori gina l
dorsal su rfa c e of the s tom a ch while the les ser curva tu re is the ori gin al ven tral
,

su rfa ce The m eso gas ter passes over the s tom a ch fo rm in g the vis ceral p eri
.
,

to n eu m of the s tom a c h and i n c losi n g the s tom a c h be tween its walls and a t
, ,

the lesser c u rva tu re is c on tinu ed by a s tro n g ligam en t the les ser omen tu m whi ch , ,

p asses t o t h e liver .

The liver may be s tu died nex t I t p resen ts a convex an terior su rfa c e tti n g
.
,

agains t the p os t erior su rfa ce of the diap hragm and a con cave pos terior s u rfa c e , ,

t tin g over the s tom a ch and rs t p ar t Of the small in tes tin e The liver is divid ed .

in to r ight an d l eft lobes ea ch O fwhi ch is su bdivid ed in to two lobes a median and a

, ,

l a ter al The lef t la teral and ri gh t m edian lobes are larger than the o thers I n
. .

the ca t the ri gh t la t eral lobe is deep ly c lef t in to two lob u les The large elo n ga ted .

g ll b l adder is im bedded in the ri gh t m edian lobe on its dorsal s u rfa c e in the

a ,

rabbi t in a c lef t in this lobe in the ca t On raisin g the liver an d lookin g be twe en
, .

the liver and the s tom a ch ano ther sm all lobe the ca u d ate lobe will be seen ,
I t is , .

si tu a ted be tween the two layers Of the lesser om en tu m The lesser omen tu m .
,

or ga s tr o hepa to du oden al l igamen t is the ligam en t p assin g from the lesser cu rva
- -
,

tu re O f the s tom a ch to the p os t erior s u rfa c e of the liver I t is divisibl e i n to tw o .

p or t io n s : t h e g a s tr o hepa tic li gam en t fro m the lesser cu r va tu re t o t he liver a n d ,

the hepa todu oden a l li gam e n t from the liver to the rs t p ar t O f the sm all i n tes ti n e .

Tha t po r tion of the gas trohepati c ligam en t whi ch con tain s the cau da te lobe of
the liver form s a s a c whi c h c on tinu es a n t eriorly the cavi ty of the grea t er om en
tum I n the hep a tod u oden al li gam en t ru n s the b ile du ct which sho u ld be t ra c ed
.

from the gall bladd er by gen tly dissec ting in the li gamen t No te the cys tic du ct .
1 9 2 L AB ORATOR Y MANUAL F OR VE R TE B RAT E ANATOM Y

from the gall bladder an d the hepa tic d u ct s from the lobes of the liver tha t fro m ,

the ri gh t la tera l lobe bein g esp e cially lar ge The cys ti c an d h ep a ti c d u cts u n i te
.

to form t he common b il e du ct whi ch p ass es to the int es ti n e in the hep a tod u ode n al
li gamen t I t shou ld be t ra c ed to the du oden u m by cleanin g away the c o n ne ctive
.

t issu e a r o u n d it No te to the ri gh t a n d dorsal to the bile d u c t lyin g al so in

.
,

the hep a tod u od en al li gam e n t the larg e hepa tic por ta l vei n
,
This mu s t n o t be
.

inj u red . Imm edia t ely dorsal to th is vein p os t erior to its bra n c h i n to the ri gh t
,

la t eral lobe O f the liver the hep a todu oden al ligam en t has a free border whi ch
,

form s the ven tr al rim of an Op e n in g or sli t of som e siz e the for amen epipl oicu m,

or en t ra n c e in to the cavi ty of the om en tum I t can be iden ti ed wi th c er tain ty

.

by m akin g a sli t in to the cavi ty of the om en tum an d p robin g thro u gh the sli t
t oward the ri gh t t oward the sp o t j u s t des c ribed
, .

The lesser om en tum ex t en ds to the middle of the p os terior fa ce Of the liver ,

where it be com es the serosa of the liver ; here its two walls p ar t an d in cl o sin g the
liver be tween them p ass t o the an terior face of the liver where they a gai n u n i te ,

toform li gam en t s The fal cifor m ligam en t ex ten ds from be twee n the two m edian
.

lobes of the liver to the m edian ven tral l in e ; it is a thin shee t wi th a c on cave
p os t e rior border A n t
. eriorly and dorsally it is c o n t in u o u s wi th the cor on ar y

ligam en t a s to u t ligam en t whi ch a tta ches the liver to the c en tral ten do n of the
,

diap hra gm The coro n ary ligam en t is c ir cu lar in form a n d its rin g of a tta c hmen t
.
,

to the live r bo u n ds a sm all sp a c e o n the a n te rior fa c e O f the liver whi ch is fr e e fro m

s erosa .

No w tra c e the in tes tin e from the p yloru s I ts rs t p or tion the du oden u m
.
, ,

is bou n d to the liver by the hep a todu oden al li gam en t T he d u od en um cu rves .

abru p tly c au dad I n the rabbit it is very lo n g an d fo rm s a loop The p ar t O f

. .

this loop whi c h des c en ds p os teriorly is nam ed the des cen din g l imb ; the shor t
tu rn a t the m os t p os te rior p ar t of the loo p is the tr an s ver s e l imb ; a n d the p ar t
whi ch as c en ds a n teriorly toward the s tom a ch is called the a s cen din g l imb T he .

d u oden um O f the c at d es cen ds c au dad for abou t two in ches an d then tu rn s to the
lef t The du oden um is s u pp or ted by a p ar t O fthe dorsal m es en tery the mes odu o
.
,

den u m . I t is also a tta ched to the ri gh t kid n ey by a m es en t erial fold the du ode ,

n or en al li gam e n t L o ca t ed in the m esod u od en u m is the pan cr ea s

. I t will b e .

seen by sp readin g the m esen tery In the rabbit the p a n c r eas co n sis ts Of s t r eaks
.

of glan d tissu e s ca ttered in the m esen tery and si tu a t ed chiey alo n g the c o u rses
of blood vessels In the cat the p anc reas is a den i te com p a c t whi te glan d which
.

ex tends to the lef t in to the dorsal wall of the grea ter om en tum dorsal to the ,

g rea t er cu rva tu re O fth e s t o m a c h I.n the rabb it t he p a n cr ea tic du c t e n t ers the d u o

den um abo u t an in ch or an in ch and o n e half an terior to the begi n n in g of the

-

as c en din g lim b of the du oden um Thi s lo ca tion of the p an crea ti c d u c t is u n u su al

.

in m am m als In the c at th ere are two pan cr ea tic du cts The p rin cip al on e
. .

join s the comm on bile d uc t a t the p oin t where the la tter en ters the du odenum .

On p i ckin g away the s u bs ta n c e of the p an c reas a t t his p oin t the du ct is readily

1 94 L AB ORA OR T Y MANUAL T
FOR VE R E B RA E ANA OM T T Y

t ra n sverse c olon tu r ns ca u dad the m eso colon is fu sed s ec ondarily to the

m eso gas ter .

The u rin ary bl adder is a s ac o ccu p yin g the p os terior en d Of the p eri tonea l
ca vit y imm edia t ely i n t ern al to the body wall an d ven t ral to the lar ge in tes ti n e
, .

F rom the ve n t ral su rfa c e of the bladder a m esen tery the median l igamen t O f the ,

bladder (median u mbilical fold of hum an ana tom y) ex ten ds to the median ven tral
lin e an d here con tinu es fo rward for some dis tanc e Near the exrt of the bladder .

from the p eri to n eal c avi ty there is on ea ch side a sli gh tly develop ed ligamen t ,

the l a tera l l igamen t O f the bladder .

The term in al p or tio n O f the des c en din g c olon is the r ectu m B o th the rec tum .

a n d t he d u c t of the bladder p ass to the e x t erior thro u gh the rin g form ed by the

p elvi c g irdle a n d ver t ebral c ol u mn They w ill be followed a t a

. la t er t im e in
c o n n e c tion wi th the u ro g eni tal sys t em F o r the p resen t it may be s ta ted tha t
.

the re c tum and u ro g e n i tal d u c t s are c om p le tely se p ara t ed from ea ch o ther A .

c loa ca is therefore abse n t .

The sm all bodies whi ch may have b een no t ed in the m esen tery u su ally b u ried ,

in fat are lymph gl and s p ar ts O f the lymp ha ti c sys t em

, ,
Sm all por tio n s Of .

lym pha ti c tis su e called lymph n odu les are also ab u ndan tly p resen t in the wall
of the in tes tine Aggrega tions of lymp h nodu les are kn own as P eyer s patches
.

.

P eyer s p a t ches are thi cken ed oval sp o t s o n the s u rfa c e of the i n tes tine Of

,

sligh tly differen t color from the res t Of the in tes tinal wall L ook for them . .

They o ccu r in the rabbi t alo n g the en tire sm all in tes tine on the S ide Op posi te
th a t a tta ched to the m esen t ery ; there is a larg er p a t ch a t the p la c e of j u n c tion
of sa ccu lu s ro tu n du s an d c ae cum The walls of the sa ccu lu s r o tu ndu s ca ecum
.
, ,

a n d verm ifo rm app e n dix are c o mp osed alm os t en tirely O f l ym p h nod u les In .

the ca t P eyer s p a tches are readily fo u nd as oval li gh t colored s p o ts along the

-
,

c olon .

S li t o p en vario u s p ar t s O f the di ges tive t ra c t alon g the side opp osi t e the a t ta ch
m en t Of the m esen tery Wash ou t the in terior I n the ca t no te the mark ed
. .

ridges or ru gae in the wall of the s tom a ch ; these are very S ligh t in the rabbi t .

Cu t thro u gh the p yloru s and n o t e the thi ckened rid ge or pyl or ic val ve a t
this p la c e I n the wall of the sm all in t es tin e observe the velve ty app eara n c e
.

d u e to the vil li (n ger like p roje c tions of the m u co u s m em brane)

-
F ind also .

the dep r essio n s m arkin g the p osi tions of the lymp h n od u les I n the rabbi t sli t .

op en the sac cu lu s ro tu ndu s cae cum an d app endix and no te the sp o tted app ear
, , ,

an ce of the i n terior owin g to the lymp h nodu les comp osi n g the walls I n the .

in terior of the cae cum is a sp iral ridge Cu t throu gh the j u n c tio n O f large an d .

sm all in tes tin e and n o te in bo th an im als an eleva tion the il eocolic valve p roje c tin g , ,

in to the ileum .

H . THE COMP ARA TIVE ANATOM Y OF THE COE L OM AND THE ME S E NTE RIE S

S in ce the d evel o men t p an d co m p a tive an a to my o f the coelom an d

ar mes en teries ar e

ex tr e mely dif cu l t an d co m plica ted s u b j ec ts it seems advis abl e at the ris k

. p
o f re et ition to
 THE COE L OM , DI GE STI VE , AND RE SP I RATOR Y S Y STE M S
1 95

mak e some co n cl u din g g en eral re mar k s con c er n in g hem

t . On l y a si mp li ed a ccou n t c an be
g iven her e .

The coel o m of shes

is d ivid ed in to two comp ar tmen ts , a s mall an terior
an d u rod el es

pericar dial cavity an d a mu ch l arg er po s terio r pl eu ro periton eal cavity T he fo rmer is s itu ated .

en tirely an terio r t o the l a t ter a n d far an terior in the bo dy a t the l evel O f th ph

, e aryn x T he .

par tition between the two co el omic cavities in thes e gr ou p s is the tran sverse s ep tu m This .

sep tu m is fo rm ed , as p r eviou s l y exp l a in ed fo r the p u r po se o f con veyin h v f

, g t e r ea t ein s ro m g
the do rs al body wal l i to the pos terior en d o f the hear t I t a is es rs t as a col u mn o f mes
n . r en
chym e on each S id e c on veyi g a vein i to the h ea t
, ; l a t er thes e col u mn s en l a g e an d fus e to
n n r r

fo m a p a tition T he tra s ve s e s ep tu m is a rather thick m emb an e with an terior an d

r r . n r r

po ste ior wal l s which in cl os e between them the g eat vein s n d the p os terior en d Of the hea t
r , r a r .

T he sep tu m in sh s an d u ro del es l ies as the ame imp l ies in a pl an e t an sv e s e to the b o dy

e n r r

axis ( F ig 4 5A a d B p
. n , .

T he lin i g o f the p e ic ar d ial cavity is n amed the p arietal p ericar diu m

n r I n shes a d . n
u ro del es it al s o co s titu tes the l ini n
g O f the body wall in the eg ion u n der con s ideratio
n
T he r n.

se rosa o f liver

t rans verse
sep t u m
rans verse

liver sept u m

trans verse
sep tum

FI G 48 .
p
Dia g rams t o sho w the se aration o f the liver from the tran sverse se tu m an d fo rm a tio n
. p
o f the co ro n a ry l ig am en t A , l iver in cl osed in the t ran svers e s e tu m
. B , b egin n in g o f c on s tric tion o f p .

the liv er f rom the s e tu m a t x p p

C, c om l etion O f the con s tric tion , l eavin g the liver s u s en ded fro m
. p
p
the s e tum b y the c o ro n a ry lig amen t T he f al ciform ligam en t is al so formed at the s ame time b y t he
.

s am e co n s tric tio n (B an d C af ter McM urrich s Devel opmen t ofthe Human B ody, co yright b y P B lakis
.

p .

t on s So n an d Co m a n y )

p .

parietal p ericardiu m is con tin u ou s with the an terior face o f the tran sve se s ep tum which in r , ,

fac t fo rms the po s terio r p art o f the p arietal p erica diu m a d the pos terior wall O fthe p ericardial
,
r n

c avity T he l in in g o f the pl eur o p er ito eal cavity is the p l eu ro periton eu m

. This is con tin uou s
n .

an terio rly with the po s terio r f ac e O f t he tran svers e s e p tu m which c o n s titu tes the a t erior p ar t ,
n

O f the p l eu o per ito eu m a d forms the an terior wal l o f the pl eur o periton eal cavity ( s ee Fig
r n n .

4 5A an d B) .

rel ation s o f the tran sverse sep tum to the liver ar e s omewhat complicated an d requ ire
T he
exp l an a tion T he liver is a div er ticul u m f o m the s mall i tes tin e
. I t happ e s that a t the r n . n

poi t wher e the liver diver ticul u m gr ows ou t ven trally fro m the in testi e the tra svers e
n n ,
n

s ep tum is s itu a ted Co s equ en tl y the liver is co mpell ed to g row ou t i to the s ep tum
. n ,
It n .

lies at r s t within the mes en chym e o f the s ep tu m betwee n the two wall s o f the s ep tu m Owin g , .

to the f act as al eady e pl ain ed th a t the g ea t vei s e ter the hear t by way o f the s ep tu m the
r x r n n ,

l iver al s o ac qu ires impo tan t rel a tio n s to these vein s as will b e dis cu s sed in the section o n the
r ,

cir cu l ato ry sys tem The l iver r ap idl y in c eases in s iz e s o tha t it ca n n o l on g er b e con tai ed
. r n

within the l imits o f the s ep tu m I t c on s equ en tly bu l g es pos teriorly carryi g with it the
.
,
n
1 9 6 LAB ORA OR T Y MANUAL F OR VE R TE B RATE ANATOM Y

po sterior wall O f the s ep tu m which th b ecome the pe ito eal co ve i g o s os a o f the liver
,
us s r n r n r er .

L ater the r egion whe e the liver bu lg e f o m the s ep tu m a ows do wn o n d o s al an d l a tera l

,
r s r n rr r

s ides l eavi g a t erio r an d v e t a l c o ec tio s be tw ee a d the s ep tu m ( Fig

,
n n e liv n r nn n n r er n .

T he an terior con n ec tion med the co ron ary l ig amen t an

is na is , as the na me in dica tes , a c ircu l ar

lig amen t , by mean s o f which the liver is p er man en tly in

al l v e teb a t es s s p e d ed f o m the r r u n r

tran s v ers e s ep tu m o r its d eriva tive The v n tral p ar titio n fo med by the co s t ic tio n o f the
. e r n r

liver fr om the s ep tum is the fal ciform lig amen t O f the l ive which is al s b y vi tu e O f its l oca r, o r

tion a p a t O f the ven tral m es e tery ( Fig 44B p

r W e th s s ee tha t the t
n v s s ep t m .
, . u r an s er e u

has the fol l o wi g p ar ts : the po s terior wall o f the perica di al cavity the a t rio r wall o f the
n r ,
n e

pl eu rop erito eal cavity the fal ciform an d co on ary ligamen ts Of the l ive the sero a Of the
n ,
r r, s

liver a d the mesod erm tiss e o f the liv er

,
n u .

In A u r a an d r ep til es the co el om is divid ed in the s ame way as in shes a n d u ro d l es in to

n e

two comp a tmen ts the p erica r dial an d pl eu ro p erito eal cavities

r ,
I n these g o p s however n . r u , ,

the p ericar dial cavity is n o l on g er s itu a ted an terio r to the pl eu o perito eal c avity b u t has r n

d es cen ded po s teriorly u n til it lies ven tral to the an terio p a t O f the l a t ter cav ity This descen t r r .

o ccu rs du rin g embryon ic d evel op men t I n this des ce t the p e icardial cavity n ec es sa il y . n r r

carr ies with it the tran sverse s ep tum s in ce this sep tum form ed the po s terior wall o f the c av ity , .

Fu rthermor e s in ce the p ar ietal p erica rdiu m o wall O f the p erica dial cavity in the lo we
,
r r r

ver teb ra t es lin ed the i s ide o f the bo dy wall it is ec es sary tha t this m emb an e b ec o m e s p l it
n ,
n r ,

a t l eas t in p a r t fro m the bo dy wall be fo e the d es c en t can ta k e pl ac e

,
This happens mai ly r . n

by the in vas ion Of the pl eu rop erito eal cavity fo wa d As a resul t O f these p oc es s the n r r . r se ,

p arietal pericardium is sep a a ted from the body wall a t l eas t in p a t an d forms a d el ica te
r ,
r ,

s ac abo u t the he t the pericardial s ac ar T he s ep ar atio o f the pericar dial s c f o m the b ody
,
. n a r

wall is compl ete in the An u a b u t in the tu tl e is i comp l ete v e tr all y so tha t the v e t al w ll
r r n n n r a

o f the p ericar d ial cavity is s till in con ta c t v en tr all y with the body w al l ( s ee F ig 4 5C p .
, .

I the g rou p s u n der con s ider a tio n the r el atio n s o f the t an s ve s e s ep tu m em ain the same
n r r r

as b efo e b u t its po s ition is n o w obliq u e in d eed al mos t fron tal whil e fo rm e ly it was t ra
r , , ,
r ns

vers e ( Fig 4 5C) I t s til l p es en ts an terior an d pos terior wall s ( which are n ow n early ven tr al
. . r

an d d o rs al in po s ition ) an d s till in cl o ses the gr ea t vein s an d the po s t erior e d O f the hea t n r

between its walls I ts an terior (ven tral ) face forms par t Of the p erica dial sac whil e its
. r ,

po sterior (do sal ) fac e for ms p art Of the lin in g Of the pl eu o periton eal cavity as p evio u sly To
r r r .

this l a t ter po r tio the liver is a ttached as befo e by the co r o a ry a d fal cifo m l ig am e ts
n r n n r n .

T he pos te ior des cen t o f the p erica dial ca vity bri gs a po tion o f the pl eu op e ito n eal
r r n r r r

c avity dor s al t o the perica dial cavity I n this po r tion Of the pl eu o p erito eal cavity which
r . r n

lies above the hea t the l u g s g row ou t r ,

n .

I n birds the el atio s O f perica dial cavity a n d s ac a d o f the t a svers e sep tu m a e t h

r n r n r n r e

s ame as j us t des crib ed T he perica r dial s ac is ho weve r o ly pa tially s ep a t ed fr om the

.
, ,
n r ar

body wall The r emain der Of the co el o m co r s po di g to the pl e op e ito eal cavity o f
.
,
r e n n ur r n

shes amphib ia n s an d rep til es is in bir ds fu the s ubdivided This is accompl is hed by mea s
, ,
r r . n

O fa p air O f m es e t erial fol ds which g o w v e tr ally fr om the s id es O f the eso ph gu s a d fu s e w ith

n r n a n

the tran sverse s ep tu m ( Fig 4 SD an d E p T he pl eu ro p erito eal cav ity is thu s divided
.
, . n

in to an terio r a d pos terio r co mp a tm e ts

n T he an terior co m p tmen t co s i ts o f the tw o
r n . ar n s

pl eu ral cavities on e on each S ide O f the p erica dial cavity an d each con tai i g a l u g T he
,
r n n n .

po s terior comp a tmen t is n am ed the perito eal cavity The li i g O f the pl eu ral cavities is
r n . n n

kn own as the pl e a ; that O f the p erito eal c vity as the p eriton e m The pa titio b etwee
ur n a u . r n n

pl eu al an d p erito eal cavities is call ed the Obliqu e sep tum I ts media po tio is composed
r n . n r n

o f the t an s ve se s ep tu m ; its l a t eral po r tio n s O f the two fo l ds a l ea dy me t io ed which a r e

r r ,
r n n

k n own as the pl e o pe ito eal memb an es ur r n r .

In mamm al s the co el om is divided as in bi ds in to fou r comp ar tme t : the peric ar dial r n s ,

the tw o O ien t a an d the p er iton eal ca viti s T he p erica r dial cavity has d es ce ded pos te io ly
l
r e . n r r
,
 1 9 8 LAB ORA OR T Y MANUAL F OR VE R E B RA E ANA OMT T T Y
T he rel a tion s o f the m es en teries a r e ver y Si mil ar in al l Of the v er te brates . T he d or s al
mesen tery ex ten ds , u s u all y un b o k en f
r ,
ro m the d o r s al median lin e to the dig es tive t r a c t .

E a ch portion bea
pecial n ame accordi g to the p t O f the d iges tive t ac t whi h it
o f it rs a s ,
n ar r c

s u p po ts I n mamm al s the mes e tery o f the s to mach is d awn ou t p os terio ly in to a b ag th

r . n r r e
,

g ea ter o me tu m which a pp a en tly s erves to p r o tec t the ab domin al vis c er a ( F ig

r n ,
r The .

v e t al m es en tery e te ds fr om the dig es tive t ac t to the ven tral b od y wal l b u t is r ed u c ed to

n r x n r

r emn a ts in the ad ul ts o f all ve teb r a tes

n These r emn an ts are foun d in the r egion O f the
r .

liver a d the bl adder wher e they fo m the g as trohep a tic a d hep a tod od e al ligamen ts the
n r n u n
,

yers o f a m es e t ry T he l iver the hea t an d the u rin ary bl adder are s itu ated in
the two l a n e .
,
r ,

the ve tral m es e t ry ; the p an c eas is g en erall y in the do r s al m es en te ry b t m ay b e in the

n n e r u

ve tral mese te y as w ll ; the diges tive tu b e may b e r egard ed a s s itu ated in the dorsal
n n r e

mes en tery .

Dia f f
FI G .
so gr am s t o Show the ormation Of the gr ea t er om en tum in m ammal s an d the usion
.

o f the m eso g as t er an d t he m esocol o n A, ea rl y s tage, in whic h t he m eso g ast er is b egin nin g t o o rm a

. f
b ag a t g . p
B , the m eso g as ter is d rawn o s terio rl y in t o a l o n g b a g g , whi ch is the g rea ter omen tum ;
the meso g as t er an d mesoc ol on a r e u sin g a t i f
C , com l etion O fthe u sion o fmeso g a s t er an d m es o col o n
. p f
at i . a , liv er ; b , s er osa o fthe liv er ; c , l ess er omen tu m o r g as tr o hep a tO d u od en al lig am en t ; d , s t o ma ch; - -

e, p
l esser eriton eal sa c o r cavity o f the g r eat er om en tu m ; f, meso col on ; g , o r tion O f the m eso g a s ter p
which forms the g rea t er o m en tum ; h, in tes tin e ; i , f u sion O f the m eso gas ter an d meso col on ( F ro m .

P r en tiss an d Arey s Textb ook of E mb r yology, a fter Her twig , c ou rt esy o f the W B Saun ders Com an y )

. . p .

I . S UMMAR Y
I . parts Of the diges tive tract are mu ch the s ame in all O f the ver tebrates Ther e has
T he .

been l ittl e evolu tion Of this sys tem In gen eral the in tes tin e is l on ger an d mo re coil ed in the
.
,

forms .

2 T he mou th cavity is provided with gl an ds in l an d vertebr ates

. .

3 T h
. e n as a l ca vit ies a r e b li d s acs in sh es b u t b e
ng inn in g with Amphibia they c om ,

mu n ica te with the m ou th an d pharyn g eal cavities for res p irato ry pu rpo s es At rs t th es e .

respir a tory p as sages through the n ose o pen in to the an terior en d o f the m ou th cavity b u t l a ter
a ha d p al a te a s hel f o f bo e is d evel o ped ven tral to the n as al p a ss ag es
r ,
n ,
B y this mean s the .

in tern a l n ares a r e m oved po s terio ly I n b i ds an d mammal s the n as al p as sag es ar e exten ded

r . r

s till f ar ther po s terio rl y by the d evel op men t o f the s o f t p al ate ; t he air then p asses dir ec tly

in to the phar yn g eal c avity withou t rs t p as sin g the mou th cavity .

4 T h
. e w all O f th e ph a ry n x i s p i er ce d by g ill s lit s p r ov id e d w it h g ill s in s h es an d witho u t
g il l s in ur od el es A b o ve t he uro
. d el es t h e g ill s lit s v a n is h w it h t he e cep tion O f t he r s t which ,
x

c on tribu tes to the ex tern al au dito ry mea tus The rs t visceral p ou ch evagin a tes to form t he
.
 THE COE LOM , DI G E STI VE AND RE SP I RATOR Y S YSTE MS
, 1 99

au dito r y tu be an d ympan ic cavity The e tod ermal l in in g o f the visceral pou ches pe sis ts
t . n r
as cer tain gl an dl ik e b odies s u ch a s the ton s il s the thymu s an d the ep ithelial bodies
, T he , , .

th yroid gl an d is an evag in a tion from the oo r o f the pharyn x between the s eco d vis ceral n

pouches .

5 The
. pharyn x in shes l eads in to a s in gl e tu be , the e Sophagus . Abo ve s hes it leads
in to two tu bes the eso phagus an d
, the tr a chea . At the cau dal en d O f the trac hea are the
paired l u n gs.

6 . pp er en d O f the trachea is expa ded in to a chamber the l aryn x the wall s O fwhich
T he u n
, ,

are su pp o r ted by the mo di ed g ill arches .

7 B o th eso phagu s an d t achea become mo re a d more el o g ted in higher forms

. r n n a
,

owin g to a d es c en t o f the v isc e a pos terio ly This des cen t l eaves a s pace i the an terior
r r . n

do rs al p art O f the b o dy cavity in to which the l u n g s g o w ou t from the l ower e d O f the t achea r n r .

8 Lun g s are p res en t in all ver teb rates above shes ( they have s econ daril y dis app ea ed
.
r

in some u rod el es) They are p robably homol og ou s with the s wim bladder Of shes Thei
.
. r

wall s beco me mo r e an d mo r e comp l ica ted an d divided in to air s p aces the higher on e as ce ds , n

in the phyl og en tic series I n bird s the l u n g s con n ec t with air s acs i the vis cera a d ai s paces
. n n r

in the bon es .

9 T he i tes tin e of vert ebra tes is p rovided with two gl an ds the l iver an d the p an c as
. n
, re ,

which ar e a ttached by du c ts to the du oden um ( the bil e du c t is g en e al ly an d the p an c e tic r r a

d c t sometimes l ac k in g in cycl os tomes)

u .

I O T he in tes ti e term in a t es in all ver teb rates excep t pl acen tal ma mm al s ( cycl os to m
. n nd es a

tel eo s to mes ) in a chamb er the cl oaca in to which the u in ary an d g e ital du c ts al o o pen
, , r n s .

I n mamm al s ( a d cy cl os tomes an d t el eo s tomes) the cl oaca is l ac kin g an d the u rin ary an d g e ital
n n

du cts o p en sep ara tely fr om the in tes tin e .

II In mos t ve teb rates there is a s aclik e ou tg rowth the u rin ary bl adder fro m the
. r , ,

ven tr al wall O f the cl oaca .

1 2. ver tebr a tes e cep t bi ds an d mammal s the coel om is divided in to two comp t
In all x r ar

me ts the pericardial cavity co tain i g onl y the hear t an d the pl eu ro p erito eal cavity co tain
n ,
n n n n

i g the o ther viscera

n T he two cavi ties ar e s ep ara ted by a p ar tition the tran sverse s ep t m
.
, u .

13 In cycl os tomes sh es an d u r odel es the p ericardial cavity is an terior to the pl eu ro

.
, ,

p rito eal cav ity F om An u ra on the p er icardial cavity is ven tral to the an terior part o f
e n . r

the pl eu rop erito eal cavity This chan ge is du e to a p os terior descen t of the hea t an d peri
n . r

cardial cavity carry i g the tran s ver s e s ep tu m with them

n AS a con s equ en ce O f the des cen t the .

wall o f the p ericardial cavity tog ether with the tran svers e s ep tu m forms a s ac the perica rdial ,

sac a ou n d the hear t

,
r That po r tion Of the pl eu roperiton eal cavity dors al to the heart is
.

des tin ed to fo rm the p l eu ral cavities .

I4 I bi ds an d mamm l s the pl eu rop e ito eal cav ity is divided in to a terior an d pos
. n r a r n n

terior po tion s by the fo rma tio n o f a pa tition which d es cen ds from the d ors al body wall a d
r r n

u n ites with the t an s vers e sep tu m This p ar tition is call ed the obliqu e sep tu m in bi ds an d
r . r

the dia phragm in mamm al s T he d ia phragm is mo e compl ex in o rig in than the Obl iqu e s ep tu m
. r

an d unl ik e the l a tter co tai s a c o s id erabl e amoun t o f s tria ted mu s cl e

n n An terior to the
n .

Obliqu e sep tu m o diaph ag m are the two pl eu r al cavities each o f which con ta in s a l u g ;
r r ,
n

posterior to these partitio s is the periton eal ca vity co tain in g the digestive an d rep odu c tive
n n r

sys tems Thu s in b i ds a d mammal s the coel om is divided in to fou r co p

.
,
r n m a r tme ts the n

pericardial the two pleu ral an d the periton eal cavities

, ,
.

1 5 A do rs al mesen tery s u ppo r ts the dig es tive trac t in all ver tebrates an d remain s p rac ti
.

cally compl ete throu ghou t The ven tral mesen tery o f the dig es tive trac t is absen t in the adul t
.

excep t in the reg io s O f the liver an d the bl adder

n Ther e are special mes e teries for the gonads .
n

an d their du c ts I n mamm al s the mes en tery o f the s tomach devel o p s a s p ec ial po s terior p ro
.

lon ga tion ca ll ed the g r ea ter o men tu m .

 X
I . T HE CO M P ARATIVE ANAT O M Y O F THE CIR CUL AT O R Y
S Y S TE M
A . G E NE RA L CON S IDE RATI ON S

r . The p art s of the c ir cu l a to ry s ys tem . T he circu l a to r y sys te m o f v er teb r a tes com

pris es two s ys tem s of b ra n c hi ng tu bes in cl o s in g cir cu l a tin g u ids , the bl ood-vas cu l ar s ys te m

an d the l ymphatic ys tem The former is th larger an d mo e co s p icu o s o f the two a d i
s . e r n u n s

the o e ref n ed to when the e p res s io

err c ircul to ry s ys tem is em pl oyed witho u t q al i c a t i
x n a u on .

T he bl oo d vas cul a sys tem is a cl os ed sys t em that is it c on s is ts o f a s et o f bra chin g tu b e

-
r , , n s,

he bl d vessel s which a e co tin u ou s with ea ch o ther un co n ec t ed with o ther s ys t ms ( e cep t

t oo ,
r n ,
n e x

the ly mpha tic sys tem) a d in which the in cl o s ed u id t ravel s in a c ir cu it

,
n T he p ts o f th . ar e

bl o od vasc l ar sys tem are the h ar t the a ter ies the vei s a d the capil l a i s T he hea t i
-
u e ,
r ,
n ,
n r e . r s a

c o t ac til e mu s cu l a r o rg a
n r s itu ated in the median ven tral r eg io n in the a te io r p t O f the n n r ar

bo dy I t has es sen tially the fo rm O f an S shap ed tu be su bdivided i to chamb rs which wil l

.
-
n e

b e n am ed in co ec tion with the dis s ec tion nn T he a r teries ar e the vessel s which l v the hea t . ea e r

an d in which the c o tai ed u id ows away fr m the hea t n n T he v ei s a re v es s el s i w hi h th o r . n n c e

co ta in ed u id o ws tow rd the hea rt

n The s tu den t s ho ul d p a rt icul a ly n o te tha t a te i s
a . d r r r e an

ve i s can ot b e d e ed on the ba s is o f the ki d o f bl ood which they co ta i

n n n T h vei s f n n n. e n o

the ve t eb a te body fall i to th ee cl asses : the sys temic v ein s which ow d i ec tly in to th
r r n r ,
r e

right s ide of the hea t ; the pul mo ry v eins which ow from the lun g s in to th l eft S ide f
r na ,
e O

the h ea r t ; a d the po tal vei s o r po r tal s ys tems in which the bl o od d o es o t r etu

n r n d i e ctl y ,
n rn r

to the heart b u t p a ss es i to a sys tem O f cap ill a ries from which it is re co ll ec ted in to s y t m ic
n -
s e

vein s The ca p ill aries a re the min u te micro scop ic v es s el s which con n ect the e d s O f the a te ies
. n r r

with the beg in i g s o f the vein s an d throu gh which the cir cu l a tion is co mp l eted
n n All the .

tis su es O f the body a r e p ermea ted with n etwo ks o f ca p ill aries th o gh the wall s o f which the r r u

g as eo u s a d o the e chan g e b etween the bl oo d an d the body c ell s ta k es pl ce

n r x a .

T he bl ood va s cu l ar sys tem in cl oses a u id the bl o d which i v e teb a t s is c o l o ed ed

-
,
o ,
n r r e r r .

The bl ood con s is ts o f a co l orl es s u id the pl ma in which oa t m ic os c p ic cell the or

,
as ,
r o s, c

pus cl es T he la tter a e o f two g en eral kin ds ed a d white the f mer g iv i g the red co l o to
. r ,
r n ,
or n r

the bl oo d S tu dy O f the bl ood lies ou ts ide o f th limits o f this co s e

. e ur .

T he lympha tic sys tem is an open sys tem tha t is it con s is ts o t o n ly O f br chi g tu b s , ,
n an n e ,

the l ymph ves sel s b u t o f la g e s p a ces the l ymph s i us es ; an d it is f u th

,
r i co mm ica tion
,
n r er n un

with the co el o m ic p c es o f the b ody Lymph s i u ses occur ben ea th the s k i ( th s t d e t may
s a . n n e u n

r ecall the l a g e s b c tan eo s lymph s in s es in the frog ) b etw e

r u u uthe mu s cl es i th m ese u ,
e n ,
n e n

te ies in the wall s O f the dig es tive tr ac t a rou d the c e t al er vo s s ys t m etc

r ,
F rom th s ,
n n r n u e ,
. e e

s in u s es the u id p ass es in t o mor e o r l es s de ite lymph ves s l s which ve tu lly o p e i to n e ,

e n a n n

the vei s O f the bl oo d vas cul a r s ys tem

n I n the l ower ve t b a tes c o trac til e l ymph he r ts a e
-
. r e r n a r

pla ced in the cou rs e Of the lym pha tic ves s el s to aid the o w b u t these a e ab se t i mammals , ,
r n n .

T he lympha tic sys tem fu r ther diers fro m the bl o od va s c la s ys t m in th t o d l s o f tis su - u r e a n u e e,

the l ymph gl ds ar e pl aced in the p a th O f the lymph ves s el s

an ,
T he lym p h gl a d s co s is t o f . n n

a n two k o f co
e ec tive t issu e in which a r e imb edd ed ma s s es o f c ll s k
r nn w a s lymp ho cytes e , no n .

Lym pho cy tes a e a variety o f white b l o od co rp s cl es

r T he fu c tio O f the lym ph gl a ds u . n n n

app ea s to b e to d s troy fo eign p a ticl es b ac t ria etc

r e a d to a dd whit e bl ood co rp u s cl es to
r r ,
e ,
.
,
n

the circul a tion T he to n s il s the thymu s a d the s pl een b el o g to the ca teg ory o f lymph
.
, ,
n n
 20 2 L AB ORA OR T Y MANUAL F OR VE R E B RA E ANA OM T T T Y
hyp om ere the e aris es on each s ide a tubul ar cavity o f the s ame n atu e as the bl oo d ves sel s
r ,
r .

As the hypomere cl os es bel ow in the median v tral li e the two ca v ities a e b ou ght tog ether en n r r

a d fu s e t o fo m the hear t ( Fig

n r

n r
B y either m th
n
O f fo ma tio n the h ea t
. eces s a il y

lies in the ve t al m es e tery O f the g ut a d is the efor e p ovid ed with d o s al an d ve t al n

e

r
r
m r
r

r
r n r

n r

mes e teries kn o wn as the dor sal an d ven tral mes ocar di ( s ee F ig 44A)
n ,
These how ver very a . .
,
e ,

ra p idl y dis app ear the ve tral m esoca rdiu m v an ishin g as soo n a s formed
,
n T he hear t the . n

s wi g s f ee in the co el om a ttached o nl y a t its t wo e d s

n r ,
T he po tion o f the coel om in which n . r

the h ear t lies is a t r s t co ti u ou s with the g en e al cavity O f th hyp o m e s b u t is s oo n cu t

n n r e re ,

Of f f o m the r e t o f the c oel om by the forma tion o f the t ran sv e se s ep tu m a d is the eu p o

r s r n r n

n amed the p er icardial cavity .

id
in ternal
caro t

vi t ell rne
art ery

F I G 52 . .
Diagram s O f earl y v er teb ra t e e mb ryos sho w the d ev el o m en t p Of the
v ess el s A , ea rlies t
. in which the c ir cul a to ry s y s tem
s ta g e , co n sis t s O f the vitellin e vein s , hea r t , ven tral

ao r ta , rs t a or tic a r ch, d o r sa l ao r ta e, an d vitellin e ar teries B , l a t er s tage sho win g t he d evel o men t O f

. p
su ccess ive a or tic a r ches foll owin g the rs t on e . f
T he m ethod Of o rm a tion O f the a o rt ic a r ches b y b u d s
from the ven tral an d d orsa l aor tae is ill u s trated b y the l as t aortic arch in B ( Fr om Wil der s His tor y of

.

the Hu man B ody, c ou r t esy Of Hen ry Hol t an d Co m p an y .

)

4 . The chi e f e m b ryo n ic b l oo d ve s s el s . The ear lies t vessel s to for m in the ve tebra te
r

em bryo a re the vitell in e v ein s . Thes e veiyolk s a c (or i the

ns dev el o p on the f
s u r a ce O f the n

ab s n ce o f a yol k s ac al on g the in t es tin e) in the s pl an chni c m es od e m O f the hypo mer e

e ,
They r .

p ass to the emb ryo in the mes en tery O fthe gu t a d e ter the po s te ior en d o f the hea t At the n n r r .

a terior en d o f the hear t a bl o o d v es s el the ven tr al a r ta a ris es a n d c o

n ec ts with the h ear t
,
o ,
nn .

T he ven t al aor ta e te ds fo rwa d to the an te io r e d o f the pharynx

r x n r H e e it d ivides in t wo r n . r ,

a n d the two b an ches turn d o s all y on e on either S id e o f the pha ryn x

r r This p air o f b an ch es
,
. r

en c ir clin g the pharyn x is the r s t p air o f aor tic ar ch s E ach v es sel lies in the c e t er O f t he e . n

s t o r ma dibul ar vis ceral arch On reachin g the dors al s ide o f the pharyn x the two ves s el s
r n .

tu rn po s teriorly an d a s the dor s al a or tae p r oceed bac k war d s itu t ed in the m edian d o s al r eg ion ,
a r

o f the b o dy wall E ch d ors al ao ta on r eachin g the r eg ion o f the yol k s ac s en d s o u t a vitell in e

. a r

a tery o ver the s u r face O f the yolk s ac

r The ea rly embryo ic cir cul ation is thus comp l eted
. n

( F ig .
52 A) .
 COM P ARA I VE ANA OM T T Y OF THE CIR CUL ATOR Y S Y STE M 20 3

There s u bs equ en tly devel op aroun d the pharyn x additional aortic arches conn ecti g the n

ven t al an d d o rs al a o tae I n typica l ver tebrates s ix su ch p airs O f aor tic arches a pp ea on e

r r .
r,

pair to ea ch p air O f vis cer al a ch s ( Fig 52B ) The aortic arch ru n s in the c e ter O f the vis
r e . .
n

c e al arch
r The aor tic a r ches d ev el op in o rder the s econ d a fter the s t then the third an d
.
r
, , ,

so o P os terio r to the pharyn g ea l r egion the two do rs al aof

n. ta e soo n fu s e t o form a s in gl e ,

dorsal aor ta which con tin u es in to the tail as the caud al a r tery Fro m the aor t a a s d ev el op men t .
,

p ro ceeds b an ches aris e for the variou s p a r ts o f the body I n hes the middl e p art o f each
,
r . s

ao r tic a ch l a ter b r ea k s u p in to a s ys tem o f c ap ill aries which o ccu p ies the

g ill l amen ts b u t
r
,

in on a qu atic forms the a r ches r em ain u n b ok en

n -
r .

I t will n ow b e p e ceiv ed th at p r ovis ion mu s t b e made for the r etu rn o f the ven ou s bl ood
r

Of the emb ryo to the hear t Fo r this p rp o se two p airs o f v ein s a ise in the s o ma tic mes ode m
. u r r

O f the b o dy wall Thes e are a p air O f an ter ior cardin l vein s r etu rn in g bl o d from the an te ior
. a o r

pa t Of the body an d a p air Of p sterior cardin al vein s retu rn i g blood from the p os te io r p ar t
r o n r

F I G 53 L a ter st a ge o f the d ev el o m en t p of the circul a to ry sy s tem, showin g the f

. .
chie v ein s .

Vein s b l an , k a r teries b l a ck . a, in t ern al caro tid ; b , d ors al aor ta ; c , an terio r ca r din al vein ; d , t he s ix
ao rtic ar ches ; e, the s ix gill slits ; f ,
the co n u s ar t erio su s O f t he hea rt c on tin uin g in to the ven tr al a o r ta ;

g, main p ar t o f the hear t ; h, sin u s v en o su s O f the hear t ; i , d uct O f Cu vier o r common c ar d in al vein ;
j , po s terior car din al vein ; l , yol k s ac ; m, su b in tes tinal vein ; n , vitell in e ar tery ;
k, vit ell in e vein ;

0, l a teral ab d omin al o r u mb ilical v ein ; p, all an tois ( u rin a ry b l a dd er) ; q, all an to ic o r u mb il ical ar tery ;
cau dal a r tery ; s , c au d al v ein ( Slightly modied from Viall eton s El emen ts de Morphol ogie des

r, .

Ver tbr s ) .

o f the body I . for t hese vei

h h a re s itu a ted in the dorsal body wal l to reach the
n o r der n s, w ic ,

heart ,
a bridg e
o f m es o derm is formed o n e ch s ide e ten din g be tween the s oma tic a n d a x

s pl a chn ic m es od erm
n W e have al ready l earn ed that this p air o f bridg es is the beg in n i g O f
. n

the t an svers e s ep tu m ; an d by the u n ion a n d exten s ion O f the bridg es the s ep tu m is comp l eted
r .

At the l evel o f the p os ter io r en d O f the hea t the an terior an d po s terior cardin al vein s on each r

s ide u n ite to a c o mmon ves s el the du ct of Cu vier o r common cardin al vein ,

T he c ommon .

cardin al v ein then p asses to the hear t f r om each s id e by way o f the tran svers e sep tum b etween

the two wall s O f which this vein is in cl osed I n addition to the car din al vein s there app ears .

in v er tebra te emb ryos a p air o f vein s in the l ateral o r ven tr al ab domin al wall s kn own as the ,

l t r al o r ven tr al a bdomin al vein s whi ch en ter the heart al on g with the common cardin al vein s
a e , .

They are al so call ed the u mbilical or al l an toic vein s in the embryos of amniotes The main .

veins O f the embryo at this time are ill u s trated in Figu re 53 .

The two vitellin e vein s a r e soon ex ten ded p os teriorly in the embryo by mean s O f a tribu
tary the su bin testin al vein whi ch cou r s es in the mesen tery o f the gu t a n d co n s titu tes the c hief
, ,

vein o f the diges tive t ac t I t is shown in Figu r e 53 I t con ti u es in to the tail as the caudal
r . . n

vein maki g a l oop a rou n d the an u s I n embryos withou t a yol k s ac the s ubin tes tin al a d
, n .
n

vitellin e vein s app ear as o e c on tin u ou s vein n .

20 4 L AB ORATOR Y MANUAL F OR VE R E B RA E ANA OM T T T Y
The '

hu s far d es cribed a e the chief l on gitu di al tru k s o f t he embryo

ar teries an d v ein s t r n n .

F rom them b an ches u su all y s egme tall y arran g ed e te d to v ariou s p a ts O f the bo d y

,
r ,
n ,
x n r .

The bran ch s O f the m ain l o gitu din al v ess l s a c l a i

e i to thr ee kin ds : the medi n vis cer al
n e re ss n a

o r spl a chn ic b a ches u p ir ed b

n n ches t o a d f o m t
r n d ig es tiv e t ac t ; th l ter l vi
,
n a r al ra n r e r e a a s ce

b a ches paired b
r n ,
ches to a d fro m the ur og en ital o g ns ;
r an n d the parietal or s oma tic n r a a

b a ches p ai ed v essel s to a d f om the bo dy wall T his a an g eme t is m o t Obviou s in

r n ,
r n r . rr n s

the a teries ( F ig
r I t is mo e o r l es s p ers is ten t in the a d l t chiey in the p o s terior p ar t
. r u ,

o f the b o dy T he ves s el s to the p air ed app e dages a e s imp ly enl a g ed s o matic b a ches
. n r r r n .

I t has alr eady been exp l ain d th a t a p or ta l sys tem

5 The or igi of the p ortal s ys t m s
. n e . e

is a p or tion o f the ve ou s s ys tem the con s titu en t v ein s o f whi ch i s tead O f en terin g the hea t
n , n r

p as s in to a n etwo k o f c apill aries from which the bl ood is re coll ec ted by a sys temic vein I n
r .

o th er wo d s in a po t al s ys tem a n e twor k o f c p ill ari s is in ter po s ed in the p a th o f a vein or

r ,
r ,
a e

v ein s .Th e a e two po t l sys tems the h patic por t l s ys tem in which the in terpo sed
er r r a ,
e a
,

c a p ill a ies e l o ca ted in the l iv e

r ar an d the r en l por tal s ys tem in which th ey a r e in the kid r, a ne
y , .

T he o rigin o f each o f thes e s ys

tem s m ay b e g iven briey .

a ) The hepatic por tal s ys tem:

This r a ises as foll ows When .

the tran s ve s e s ep tum devel op s r

a t the po s terio r e d o f the n

heart the two vitell in e vein s

do rsal aorta
,

mu s t n a tu ally p ierce the s ep

r

tum on their way to the hea t r .

At this r egion the liver bu ds

fr o m S mall
FI G .
54
Dia ram
g. of a c r o s s s ec tio n
-
thro u gh a v ert eb ra t e
ou t

a n d ex ten d
h t e I n tes un e

the t r an s v er se
( Af ter
s
em b ry o , to s ho w the s eg men tal b ra n ches Of the a o r ta .

M cM u r r ich s

Devel opmen t of the Hu man B ody, co py ight
r by Sep tu m AS 1 t g rows ; the liV er
P B l akis t on
.

s So n an d Com p an y .
) gr adu al ly u s es u p the s u b
s tan ce o f the t r an sver s e s ep ~

turn an d ll s the a vail a bl e s pace in the se p tu m . T he liv e r su b s tan ce thu s c o m es to

s u rr oun d p roximal e ds o f the two vit lli e v in s ( Fig 55A an d B ) At s t the vit llin e
the n e n e . . r e

v ein s p a s s th ou gh the liv er i to the h ea t b u t s o on th ey b eg in t o b eak u p in to s mall er a n d

r n r ,
r

s m all er v es sel s ( Fig 55C) in the liv er u til the liv er is o c cu p ied by a n etwo k o f ca p ill a ies ( o f
. n r r

the kin d kn own as s in s oid s) which p e m e te the liv er s u b s tan ce u Th s the ci cu l a tio in r a . u , r n

the vit llin e vein s p a s s es f om the y olk s c a n d d ig es tiv e t a c t t o the liv er p asses th o gh a
e r a r , r u

c a p ill ary n etw o r k in the l iv er a d fr om the l iv e t o the h ea t in the r em ain in g p oximal p o tio s
,
n r r r r n

O f the vitell in e v ein s ( F ig 55D) T he l a tter a e n o w call ed the hep tic v ein s
. P o s terio r t o. r a .

the liver by ri glik e u n ion s b etween the two vitell in e vein s ( Fig 55C a d D) a s in gl e ves s el the
n . n , ,

h p tic por tal v i

e a i fo m ed an d ca u dad o f th se
e n, io ns the right vein dis app ears l eavi g
s r ,
e un ,
n

f
the l e t vein w ith it s t r ib u ta ry , the s u b in tes tin al vein , n ow n a med the mesen ter ic vein , to form
the c ie h f vei n of the d ig es tive t r ac t . This a r ran g e men t in s u r es tha t a ll O f the ven ou s bl oo d
fro m the d ig es tiv e t r ac t c au d a d O f the car dia mu s t p as s throu g h a ca p ill ary n etwo rk in the
liver b efore it c an r eac h the hea rt.

b ) The This d evel op s a s foll ows At rs t the cau dal vein Op e s in to

r en al p or tal s ys te m: . n

the s u b in t es tin al vei formin g a l oo p a ou d the a u s a s in F igu r e 55C an d D L a ter the

n , r n n .

po ste ior c a din al v ei s g row po s terio ly a n d co ect with the cau dal vein The u n ion o f
r r n r nn .

the c au dal v ein w ith the s u b in tes tin al vein is the b r ok en a s in F igu e 55E the l a t ter vein then n r ,

b ecom in g a t ibu tary o f the hep atic po tal sys tem The e n e t dev el op s between the kid eys
r r . r x n

a v i a t s t s i gl e l
e n t e p a ed
r ir
r
the s b r din al vein ( Fig 55F )
n a This con n ec ts with the u ca .
, , .

c a d l v ein an d p o s terior en ds O f the po s t erior ca din al veins

u a T he bl oo d ows fr om the ta il r .
20 6 L AB ORA OR T Y MANU AL F OR VE R E B RA E ANA OM T T T Y

in to the bca din al v ein s a d th o gh th k id eys in to the p o te i r c din l vein s The e

su r n r u e n s r o ar a . r

n ext o cc s a b ea k b etw een the a t ior a d po s te io r p ar ts o f the p o t io r cardin al vein s

ur r n er n r s er .

T he a terio p ar ts form a co
n rc tio n with the s b ca dirn l v ein s T he p o s ter ior p a ts r etain
nne u r . r

ec tion with the cau dal vein Ther e is then a reve s l o f ow th o gh the kid eys

th ir c o
e nn . r a r u n ,

i
s n cethe bl oo d n ow p ass es f om the ca d a l vei in to the po s terio p a ts O f the p o s t e ior ca r din l
r u n r r r a

v ei s n o w c all ed the r e
n ,
l por tal vein s thr ou gh the kid eys i to the su b c d in l v ein s an d
na ,
n n ar a ,

f o m them i to the an terior po tion s o f the po s terio c a din al vein s ( Fig 5

r n r A t r s t the r r .

cha el s th o gh t he kid eys a r e d ir ec t c o n ec tio s b etween the r e al po t l a d su b ca din al

nn r u n n n n r a n r

v ei s b u t l at er they b ea k u p in to a c ap il l ary n etwo r k

n ,
r I n this .

t em is es tablis hed T he bl oo d f om the t ail mu s t p as s th o gh

. r r u

kidn eys .

We hav e n ow ca rr ie d the c ir cul a to r y sys tem u p to the s tage

o c cu r s in mob r an chs The u r
a d u l t el a s . f ther evolu tion o f the c ircul ato ry sys tem
foll owed on the s p ecim en s .

6 The s e gm en tal char acte r Of the cir cu l atory s ys te m

.
L ike mo s t sys tems o f .

the c ir cul atory s ys tem exhib its m ar k ed t ra c es O f an o r ig in all y highly s egmen ted

dow
d
iges tive tub e m e 10 09

FI G .
56 Dia g
. r am O f t he hy po th ti e cal p imiti
r ve v er t eb r a t e cir cul a t io n from which th a t Of p res en t

v er t eb ra t es wa s p ro b ab l y d erive d m ph
,
e a s iz in gmarkedl y s egmen tal a rr an g em en t o f the rimitiv e
the p
v essel s .
( Af ter Kin gsl ey s
Compar ative An atomy of Ver teb r ates , co yright b y P B lakis ton s S on a n d p .

Com p an y .
)

However , even in ver teb ra t e embryo s the s egmen tal a rr an gemen t o f the bl ood v ess el s is so me
wha t imper fect I t is hel p ful to an u n d er s tan din g O f the cir ul a tory s ys tem t o imag in e tha t it
.

is d eriv ed from a highl y s egmen ted c on dition s u ch as th a t p r es en t in an n elids I n this hyp o .

thetical an ces tr al s ta t e as ill u s tr a ted in Figu r e 56 th ere a r e thr ee m a in l on g itu din al vessel s :
, ,

a do r sal s om a tic v es sel , corr es po n din g to the d o r s al a o r ta ; a v en tr al s o ma tic v ess el corr es p on d ,

in g t o the abdomin al vein ; an d a s pl an chn ic vess el , co rr es po n din g t o the s u bin tes tin al an d
vitellin e vein s an d the ven tral aorta I n ea ch s egmen t thes e l on gitu din al vess els are con n ec ted
,
.

by l oo p s , which ar e O f two k in ds s omatic an d s pl an chn ic I n the an ter ior p ar t o f the body

,
.

on l y the s pl an chn ic con n ec tin g l oop s a e pr ese t r ep res en ted by the aor tic arches con n ec tin g r n ,

the d o r s al a d ven tral aor taen P o s terior t o the hear t ea ch s egmen t is p ro vided with a S oma tic
.

an d a s p l an chn ic l oo p the f o rmer p as s in g in the b o dy wall f o m the ao t a to the ve tr al a b d o mi

,
r r n

n al vei an d the l a tt er p a s s in g in the in tes tin al wall f om the ao ta to the s ub in tes ti al v in

n ,
r r n e .

These tran sver se l oop s p ictu r ed as co ti u o s a e in reality in te p ted by capill aries in the n n u r rru

b ody a d i testin al wall s I n v er teb a tes all O f the co ec ti g l oop s e l o s t e cep t the o tic
n n . r nn n ar x a r

a r ches ; b u t s eg me tally a rr an g ed t r a s v e s e v ess el s c o

n es p o di g to the l p s are pl ai ly
n r rr n n oo n

evid en t in ver t b a te emb ry o s a d p e s is t in c er tain r eg io s o f the

e r d l t as in Figu re 54
,
n r n a u ,
.

Fo r mor e e tens iv e accou n ts of the comp ara tive an atomy Of the cir cu l atory sys tem; K or
x

W s ho ul d b e cons ul ted .
 COMP ARAT I VE ANA OM T Y OF THE CIR C L A OR U T Y S YSTE M 20
.

B . TH E CIR C U LAT OR Y S Y S TE M OF E LA S MO B RANCH S

The followi n g a ccou n t app lies to bo th S p e ci es of dogs h and to the skat e .

.r The chamb er s Of the he art The hear t when . rs t formed is a si mp le

s trai gh t tu be b u t it soon bec om es ben t u pon i t self in the shap e Of the le tter S
, ,

and it s wall be c om es differen tiat ed int o a number of chambers The h ear t Of .

the elas m obran chs is in thi s c ondi t ion c onsis t in g of fo u r chambers The p eri
,

c ardial c avi t y has already been e xp osed in t he p re c edin g disse c t ion Sp read its .

walls ap ar t I den t ify the c ham bers of the hear t as follows On raising the
. .

hear t a t riangu lar chamber will be seen extending from the hear t to the transvers e
s ep tum its two c orners b u ried in the septum This is the s in u s ven os u s the
, .
,

m os t p os t erior c ham ber O f the hear t E a ch c orner of the sin u s venos u s is co n

.

tin u ou s wi th a lar ge vei n the du ct of Cu vier or common cardin al vein whi c h is

, ,

inc losed in the t ran sverse sep tum an d will be seen la ter An terior to the si n u s .

venos u s is the atriu m or au r icl e a large t h in walled chamber exp anded on ea ch

,
-

side of the hear t and app earing as if p aired B etween the two sides of the .

au ri cle res t s the ven tricl e a thi c k walled he art shap ed c hamber the m os t c o n
,
-
,
-
,

sp icuou s p or tion of the hear t from ven t ral view The p oin ted p os terior en d of .

the ve n t ri c le is known as t he apex the broad an t erior end the b ase ,

F ro m the , .

base of the ven t ri c le a t hi c k walled tu be runs forward and p enet rat es the an t erior
-

wall Of the p eri cardial c avi t y This is the con u s ar teriosu s the fou r th and m os t
.
,

an t erior cham ber O f the heart The blood c ir cu la tes t hrou gh the c hambers of
.

the hear t in t he follo wing order : sinu s venos u s a u ri c le ven t ri cle c on u s ar terios u s , , , .

.2 The s ys te mi c ve in s S ys temi c veins have already been dened as

.

those veins whi c h en t er t he heart All sys tem i c vein s in ver tebra tes op en in t o
.

the s in u s venos u s or its eq u ivalen t t ha t is t hey en t er the p hylo gene ti c ally p os t e

, ,

rior end of the heart Owing to differenc es be tween them the d ogshes an d
.
,

ska t e will be des c ribed sep ara tely .

I n disse ct ing the veins they are followed away from the hear t and it is of t en
, ,

c o n venien t to s p eak of the m as if t hey p ro c eeded fro m the hear t t o body

s tru ctu res The s tu den t mu s t however always hear in mind the fa c t tha t
.
, ,

they c onvey t he blood fr om the p ar ts Of the body to the hear t .

D og s h: I nser t one blade of a ne s cissors in the sinu s wall and sli t the
ven tral wal l of the sinu s venos us op en in a c rosswise dire c tion The c avi ty of .

the sinu s is th u s exp osed and sho u ld be washed o u t thoro u ghl y u nder a s t ream
of ru nn ing wat er All Of the sys t emi c veins op en in t o the cavi t y of the sinu s
.

and the op enings may now be iden tied with the cu t ed ges Of the S in u s wall ,

s pread well ap ar t E a chla t eral wing Of the sinu s whi ch lies b u ried in the t rans
.

verse s ep tu m re c eives a very large op en ing the en t ran c e of t he du ct of Cu vier ,

or common car din al vein The natu ral rela tions of this en t ran ce are bes t Obs erved
.

on the in t a c t righ t side On the lef t side c arry the sli t in the sinu s lat era lly to
.
 20 8 LAB ORA OR T Y MANUAL FOR VE RTE B RATE ANATOM Y

m ee t the in c ision p reviou sly m ade across the gill sli ts The en tran c e Of the .

c o mm on c ardinal vein in t o the sin u s is t h u s sli t o p e n The followin g m ay then .

be n ot ed In the sp iny d o gs h j u st m edial to the m ain op enin g of the comm on

.

c ardin al v ein in t o the sin u s are several s m all ap er tu r es m os t O f whi c h app ear t o ,

be su bdivisions Of the chief op enin g The m os t an t erior of thes e sm all ap er tu res

.

is however the op enin g of the infer ior ju gul ar vein In the sm oo th d ogs h the
, ,
.

a cc essory op enin gs are lac kin g an d the ent ran c e of the inferior j u gu lar vein is
,

situ at ed j u s t an t erior to the op enin g of the c omm on c ardinal v ein P rob e i n to .

t he op enin g and n o t e t ha t t he vein c om es from the oor of the m o u th and p ha

r yn geal c avi t ies wher e it ru n s alo n gside t he ven t ral ends of t he g il l ar c hes By .

tu rnin g ba c k t he ap p revio u sly form ed of t he oor of the m o u t h and p haryn geal

, ,

c avi t ies the c o u rse of the vein will be m ore readily followed
,
Now look into .

the p os t erior p art O f t he wa l l of the sin u s pu t t in g t his on a s t ret ch

'
In t he , .

m edian lin e of the p os t erior wall wil l be n o t ed a whi t e fold and o n ea ch side of ,

t his is an o p enin g P robe in t o bo th op enin gs and no t e t ha t yo u r p robe p asses

.

in t ernal t o the c oronary li gam en t and int o th

the l iver by sli tt in g the s u bs t an c e O f the liver and no t e t he ,

and left lobes of the liver t hu s revealed These c avi ties whic .

en t ire le n gt h Of the liver lobes are t he hepa tic s in u ses The .

are the p ersis t ent p roxim al p art s of t he vi t elline veins of the e

Now p robe p os t eriorly in t o one O f the c omm on c ardinal
vis cera in the ant erior p art of the p leu rop eri t on eal c avi ty and
p robe has en t ered a lar g e bl u ish s ac lo c a t ed in t he dorsola t eral
p e ri t oneal c avi t y F ollow .t his s a c and its fellow Of t he o pp osi

E a ch
the vis c era t o one side eac ,

Imm edia tely t o ea c h side

m edian dorsal lin e These t wo vess el s are

.

are the c hief som a t i c veins of the t ru nk .

the liver the two p os t erior cardin al sinu ses

broad c onn e ction whi ch m ay be fou nd by p robin g in t o the left v ein
the p robe t oward t he ri gh t one In t his sam e re gio n ea c h vei n has
.

s u rfa c e an ex tensive c omm u ni c at ion wi th a large blood S in u s the gen ital sin u s
'

, ,

s u rrou n din g ea ch gonad E ac h p os t erior c ard inal sin u s also r eceives n u m ero u s
.

se gmen tal ly arran ged bran ches from the body wal l (par ietal veins) an d from the
kidneys (r en al veins) The kidneys are the lon g slender at organ s brownish
.
, , ,

in c olor whi c h lie imme diat ely la t eral t o the p os t erior cardinal sin u s es ext en d
, ,

in g the en t ire len gt h of the p leu rop eri t o n eal c avi t y The p ari etal an d ren al vein s .

are readily ident iabl e in those sp e cim ens in whi c h they happ en t o b e lled wi th
blood b u t are imp ossibl e t o see wh en emp ty
,
.

O wi g t th f t th t th
n o e ac i f l m b
a h
e ve n s O t d it e asl b t po i th ti
ra n c s a re n o e n e v ess e s u s a ces n e ss u es

withou t d e nite wall s ,

they a re mo r e c o rrec tl y d es ign a t ed s in u s es .
 210 LAB ORA OR T Y MANUAL T
FOR VE R E B RA E ANA OM T T Y

l a t eral m arginof the cloa cal ap er tu re and near the ant erior p art of the ,

the p elvi c n re c eives the il iac vein from t he n Tra c e t his a cc ordin g .

dire ctions for the sp iny sp e cies .

In the m edian ven t ral line are t ra c es O f a ven tr al cu tan eou s vein whi c h
o

however to dwindl e away ant eriorly

,
.

Draw t he sys t emi c veins .

S kate : The sin u s ven osu s c o n sis ts of a tu be on ea ch side the c en tral ,

being redu ced in S iz e and a t t a ched t o the t ransv erse sep tum by a she et of c o n nec
t ive t iss u e whi c h m ay be broken E a c h sid e of the S in u s is b u ried in the tra n s
.

verse sep tum Fol low ou t the righ t side t o the p oin t where it disap p ears dorsal
.

to the c art ila ge of the p e ct oral girdle Caref u lly shave away t he c ar tila g e a n d
.

s u rrou nding tiss u es u nt il the sinu s venos u s can be followed laterally Inser t on e .

blade of a n e s cissors in t he vent ral wal l Of the sinu s and sli t it op en in a c ross
.
wise dire ction The sin u s is seen to be c ontin u ou s on ea ch side wi t h a tu be or
.

c ham ber the common ca r din al vein or duct of Cu vier whi c h tu rns dorsally
,
All , .

of the sys t emi c veins op en in t o the c omm on cardinal vein and their op en ings ,

m ay now be iden tied The j un ction of c omm on c ardinal vein wi th the sin u s
.

venosu s is m arked by a S ligh t fold In the ant e rior wall O f the c omm on c ardinal
.

vein c on cealed by this fold is the sm all op enin g of the infer ior ju gu l ar vein This .

op enin g is so sm all th at the p robe can p robably n ot be p assed in t o T he

i n ferior j u gu la r vein drains the walls of the p eri cardial c avit y and the oor O f
the m o u t h and p haryn geal c avi ties I n t he p os t erior wall of the c omm on c ardinal
.

vein at its j u n ction wi th the sinu s venosu s is the op enin g Of the hepatic sin u s
, ,
.

P robe p os t eriorly in t o this I t leads in t o the hep ati c sin u s a lar ge sp a c e si tu at ed

.
,

between the ant erior margin of the l iver and the transverse
by the c oronary ligamen t In .

Of the ovid u c t whi c h is in c losed in the fal ciform ligam en t I n lo ca tin g the hep a ti c .

sinu s p ress the l iver c au dad away from the t ran sverse septum ; the sin u s forms a
b ag be tween liver and sep tum ven t ral to the esop ha gus Cu t in t o the hep a ti c . .

sinu s A tu be the per icar dio per iton eal ca n al ext ends throu gh the c en t er of its
.
,
-
,

c av it y L ook in the p os t erior wall of the sin u s for the sm all op enin gs of the
.

hepa tic veins whi c h drain the liver The s inu s op ens on ea ch side int o the
.

c omm o n c ardinal v ein .

Now p rob e i n t o the m ain c avity of the c omm on c ardinal vein in a dorsal
and p os terior dire c tion The p robe en ters the pos ter ior car din al sin u s whi c h is
.

a broad thin walled tu be lyin g a gains t the dorsal wall of the p leu rop eri ton eal
,
-

c avi t y I n fem ales it is on the dorsal side of the ovid u c t ; in males dorsal t o
.
,

t he t es tis F o l low the p o s t eror ca rdinal sin u s p os t eriorly

. I t sw e rves t oward .

the m edian li n e where it soon m ee ts it s fe llow of the opp osi t e side to form a sin gle
large median sin u s This sinu s c ommu ni ca t es on its vent ral su rfa c e wi th the
.

l arge gen ital sin u s s u rrou nd in g each gonad M ore p os t eriorly the p os t erior c ardi
.

nal sin u s s ep ara t es a gain int o two vein s which p ro c eed ca u dad on the m ed ial
 COMPARA I VE ANA OM T T Y O F THE CI R CUL ATOR Y S Y STE M 21 1

side Of the kidneys The kidneys are rounded lobes at the p os t erior end Of the
.

p le u ro p eri t oneal c avi t y a g ains t t he dorsal wall T O see t h em rem ove the .
,

p le ur o p e ri t o n e u m fro m th e do rs al wall on ea c h side Of t he c loa c a Do n o t .

inj u re t he d u ct s from the kidneys Af t er e xp os u re of the kidneys the p os t erior

.

cardinal veins will be fou nd on the m edial side of the kidne s They onne t
y c c .

wi th ea ch o t her between the kidneys .

R e tu rn to the c omm on cardinal vein and p rob e in t o it in a dorsal dire ction .

Tu rn the animal dorsal side up and lo ca t e the end Of yo u r p robe M ake an .

in cision in t o the sp o t indi cat ed by the p robe and ext end the in cision longi tu dinal ly
forward to the eye On c areful ly deep ening the in cision an elonga ted cavi ty with
.

smooth wal ls the an terior cardin al sinu s is exp osed It is situ at ed j u s t medial
, , .

t o the dorsal ends of the gill ar ches and vis c eral p ou ches It may be followed .

forward wi th t he aid Of the p robe I t tu rns lat eral ly in front Of the rs t vis ceral
.

p o uc h follows
, alo n g t h e an t erior border O f t his p o uc h and t hen t u rns an t eriorly ,

again .

Wit h the dorsal side Of the anim al s til l fa ci n g you lo ca t e by feel ing the ,

ch i ef an t erior c ar tila ge (p rop te r y giu m ) O f the p e ct oral n I t form s a c res cen ti c

'

rid ge lat eral t o the gill r egion nearly halfway from the mid dorsal line to the -

margin Make an in cision alon g the medial fa ce Of t his c art il age on the left
.

side A vein will be exp osed ru nning along the c art ilage here I t is one of the
. . .

br achial v eins F ollow it p os t eriorly

. I t will be fo u nd t o en t er the c omm on
.

Tu rn the an im al vent ral side u p again Along the lat eral wall Of the pleu ro .

p eri t oneal c avi t y ru ns t he l a ter al a b domin al vein N o t e t he p a r ieta l bran c

.hes
whic h it re ceives from the body wal l at ea ch myosep t um Trac e the lat eral .

abdominal vein an t eriorly It p asses along the int ernal su rfa c e Of the car tilages
.

of the p e ct oral n and p e ct oral girdle and en t ers the c omm on cardinal vein
, .

Cu t in t o the vein where it p asses the c ar t ila ges Im media t ely on the p os t erior
.

side of the c artilage of the p e ctoral girdle a b rachial vein will be fou nd en tering
the la teral vein Imm edia t ely p os t erior to this is ano ther c ar tila ge an d on the
.
,

c au dal side Of t ha t ano t her br a chial vein joins the la t eral vein A third bra chial .

vein was m en tioned in the p re ceding p aragraph Tra c e the la t eral vein p os t e .

rio r l y. I t ori gina t es in a ne twork of s m all vessels on the sides Ofthe large in t es tine
and c loa c a I t p asses on the in n er su rfa c e of the c ar tila ges Of the p elvi c girdle
.

an d p elvi c n S li t the ve in op en alon g the s u rfa c e Of the c art ila ges

. I l iac veins .

wil l be fou nd em ergin g from be tween the c art ila ges and ent ering the la teral vein .

The larges t of the ilia c veins is lo ca t ed alo n g the p os t erior side of the car tilage
of the p elvic girdle P robe in to the il ia c veins and no te their dis t rib u tion in the
.

D raw sys tem i c vein s

t he .

T h he a ti or t l ys t e m A or t al sys t e m is a sys t e m O f veins whi ch

3 . e p c p a s p .

ows in t o a n etwork o f c ap illari es In som e organ The hep a tic p or tal sys t em .
21 2 L AB ORA OR T Y MANUAL T
F OR VE R T E B RA E ANATO M Y

c onsis t s of veins whi c h c olle ct the venou s blood from the diges tive t ra ct and
spleen and p ou r it in to a network Of c ap illaries in the liver L o c at e the bile
, .

d uct L yin g in the hep at od u odenal l igamen t al ongside the bile du c t is a large
.

vein the hepatic por tal vein Tra c e it p os teriorly and identify the branch es
,
.

which it re c eives from the diges tive t ra ct These branc hes differ s l ightly in the .

t hree fo rms u nder c onsidera t ion .

S pin y d og s h: On t ra cing the hep a ti c p o r tal vein p os teriorly it will be ,

fou nd to re c eive rs t a very s m all bran ch whi c h m ay be c alled the du oden al ,

branch whi c h runs along the bile d uct and c ollec ts from the d u odenum and
,

an terior p ort ion Of the sp iral valve P os t e rior to this the hep ati c p ortal vein is .

seen to be formed by the u nion Of three large branc hes The lef t bran ch the .
,

g as tr ic vein p asses,
a t on c e t o the s t om a c h where it is for m ed by the u nion , Of

the dor s al and ven tr al g as t ri c veins whi c h bran ch on the dorsal and ven t ral su r
,

fac es of the s toma ch The middle Of the three main branches Of the hep a ti c
.

p or t al is th e lie n omesen ter ic vein I t p asses p os t eriorly do

. r sal t o t he d u oden um

and is im bedded in the s u bs tanc e Of the dorsal lobe Of the p anc reas from whi ch ,

it re c eives bran ches At the p os terior end Of the p an c reas the vein is seen to be
.

form ed by the u nion of two branches : one the poster ior splen ic vein from the , ,

sp leen ; and the o ther the pos ter ior in testin al vein whi ch c om es from the left ,

side of the sm al l int es tine No t e its num erou s branches from the in t e stinal wa ll
.

along the lines of a tta chm en t of the tu rns Of the sp iral valve The ri gh t branch .

of the three th at fo rm the hep ati c p orta l vein is the pan cr eatico mesen ter ic I t -
.

p asses dorsal t o the p yloru s and is im bedded in the s u bs t an c e Of t he ven t ral lobe

of the p an c reas Here it re ceives the an terior spl en ic vein from the ant erior
.

p ar t of t he s p leen and several veins fro m t h e p an c reas and d u oden u m I t s m ain .

t ru nk the an ter ior mesen ter ic vein is si tu at ed along the ri gh t side of the in t es tin e
, ,

from which it re ceives branches in the sam e m anner as the p os t erior int es tinal
vein .

S m oo th d og s h: The bran ches are si m ilar to those Of the s piny sp e c ies .

The hep ati c p ortal rs t re c eives the ve ry sm all du oden al bran ch from along the
bil e du ct and S hor tly b eyon d this p oin t is formed by the u n ion of three large veins :
the gas tr ic vein on the left the l ien omes en ter ic in the middl e and the pan crea tic o
, ,

mesen ter ic t o the ri gh t The gas t ri c vein p asses t oward the s t om a ch an d re c eives
.

an an terior ven tr al gas t ri c vein It is situ at ed in the m esen t ery between the two
.

limbs of the s t om a c h and in its co u rse re c eives bran c hes from bo t h lim bs The .

lien omes en t eri c vein p asses dorsal to the p yloru s lyin g im bedded in the dorsal
lobe Of the p an c reas At the an terior m argin Of this lob e of the p an c reas it
.

re c eives the an ter ior dors al gas t ri c vein from the c orr esp ondin g p ar t of the
s t om a ch It re c eives num erou s pan creatic veins from the p anc reas At the
. .

T hese b ran ches a re f

g en erall y ll ed with b l oo d a n d there o r e eas il y t ra c ed If they a r e em ty,. p
they ma y b e rea dil y in j p p
ec ted thro u gh the he a tic po rt al vein , even in s ec im en s whic h ha v e b een

p res e rved fo r a l o n g time .

214 LABORA O R T Y MANUAL FOR VE RTE B R ATE ANA OM T Y

vessels in closed in the haem al arch The cau dal art ery is dorsal the cau dal
, .
,

vein imm ediat ely vent ral t o the a rt er y .

D o g s h: P robe in t o the cau dal vein Ob s ewe that the p robe c an be p assed
.

eith er t o the ri gh t or left showin g that the vein forks at the anu s The two
, .

forks are the r en al por tal veins L eavin g you r p robe in one of the renal p or t al
.

vein s lo cat e the kidney in the p leu rop erit oneal c avi ty I t is a long brown
, .

organ si tu at ed agains t the do rsal b ody wall on e on ea ch side of the mid dorsal
,
-

lin e ext ernal t o t he p leu rop erit oneum Sl it the p leu rop erit oneum along the
.

lat eral border near the p os t erior end of the ki dn ey on the side where you r p robe
is ins ert ed and gentl y lift the kidney away from the body wal l A sp a c e will be .

fou nd between the kidney and the body wal l ; in t o this sp a c e you r p robe has
p assed . T his s p a c e is th e renal p or t al vein or sin u s I t . bran c hes in t o the
kidney and also re ceives t rib u t aries from the body wall .

S kate : The kid n ey has already been exp osed L ook along the m edial side
.

Of t he p os t erior p art Of the kidney for a vein c om in g fro m the vert ebral c ol umn .

Do n ot in j u re any du cts on the ven t ral su rfa c e of the kidney In males the vein .

in qu es tion l ies imm ediat ely t o the dorsal side of the m ale du ct whi ch will be ,

seen p a ssin g along the vent ral s u rfa c e Of the kidney t o the c loa c a The du ct .

m ay be l ift ed from the kidney s u rfa c e and ben t t o one side The vein in qu es tion
.

is the r en al por tal vein At rs t it li es alon g the m edial border of the kidney
.
,

b ut soon tu rns on t o the ven t ral s u rfa c e of t hat organ givin g Off bran c hes in t o ,

its s u bs t an c e and re c eivin g t rib u taries from t he body wall lat eral t o the ki dney .

The renal p ortal veins are c ont inu ations of the c a u dal vein ; the la tt er forks at
the anu s givin g rise t o the two re n al p ort al vei n s The forking is however
.
, ,

dif cu lt t o t ra c e in the S kat e .

R eferen c e to F i gu re 55 p age 2 0 5 will show t hat the renal p ort al veins are the
, ,

p os t erior p ar t s of t h e p os t erior c ardinal veins and t h a t t h e a pp aren t p os t erior

p ar t s of t h e p os t erior c ardin a l veins of th e ad u l t are in reali t y t he s u b c ard i nal
veins Whereas in t he embryo the blood ows from the su b c ardin als int o the
.

p os t erior c ardi n als ,

in t h e ad u l t t he dire c t ion O f ow is reversed The renal.
p o r

t al sys t em p rovides t ha t the blood from the t ail mu s t p ass in t o a c ap ill a r y

sys t em in the kidn eys from whi ch the blood is re c olle ct ed int o the s u b c ardinal
veins The pu rp ose Of this arran gem ent is obs cu re ; it seem s t o have been disad
.

van t ageou s for the vert ebra t es la t er sh un t ed p art of this blood in t o ano t her
,

sys t em a n d nally abandoned the renal p or tal sys t em al t oge ther .

D raw the renal p ort al sys t em .

T h ve t l aor t d th t b r n hi l ve el T rn on e m ore
5 . e n r a a an e a e re n a c a s s s u . c

to the p eri c ardial c avi t y of the s p e c im en The c on u s art eriosu s ru ns forward and
.

p ene t ra t es t h e an t e rior wall of t he p eri c ardi al c avi t y C ar

. efu lly p i c k away

m u s cles an d c onne ct ive t issu e from the r egion ex t en din g from the an terior end
of t he p eri c ardial c avi ty to the low er jaw In the m edian vent ral l in e wi ll be
.
 COM P ARA I VE ANATOM
T Y OF THE CIR CUL AT OR Y SY STE M 21 5

r evealed a large vessel the ven tr al a or ta whi ch c on t in u es fo rward from the c onu s
, ,

art eriosu s B y disse cting c aref ully to the left side n d the bran ches Of the
.

vent ral aort a They are as fol lows :

.

Dog s he s : There are three m ain p airs of bra n c he s two Of whi ch s u b di vide ,

in t o two The mos t p os t eri or p ai r Of branches a rises j u s t at the p oin t where

.

the c onu s art erios u s p asses in t o the ven t ral aorta (At this p oint not e the
.

cor on ary ar t e ry p aired in the sp iny d o gs h sin gle in the sm oo t h sp e c ies p assin
, , g ,

alon g the c onu s art erios u s ont o the s u rfa ce Ofthe ven t ri cle and to the walls o f the
p e r i c ardial c avi t y This
. ar t ery sho u ld be p reserved as far as p ossible ) F ollow .

the m os t p os t erior bran c h Of the ven t ral aor t a I t ve ry s hort ly divides in two
.
,

the p os t erior bran c h p ene t ra tin g the in t erbran c hial sep tum Of the S ix th vis c eral
arch the anterior branch the sep tum Of the f th The m iddle bran c h of the
,
.

ven t ral aor ta arises short ly in front of the t hird bran ch and p asses wi thou t
division in t o t he in t erbran chial sep tum Of t he fou r th vis c eral ar ch Af t er givin g .

of f this bran c h the ven t ral aor ta p ro ceeds forward wi t ho u t bran c hin g t o a p oin t
j u s t p os t erior to the lower jaw Here it forks to form its ant erior p air of bran ches
. .

Trac e the lef t bran ch lat erally Af t er som e dis t an ce it forks su pp lyin g the
.
,

se cond and third vis c eral ar ches Tra c e any one of the bran ches of the ven t ral
.

aort a ou t int o the in t erbranchi al sep tum sli ttin g the sep tum No t e the small
,
.

bran ches from the art ery int o the gil l l amen t s The ve p airs Of bran ches of .

the ven t ral aort a are nam ed the a er en t b r an chial art eries H ow m any gills .

does ea ch su pp ly ?
S kate : There are two m ain p airs of bran ches fro m the ven tral aor ta The .

p os t e r ior p air arises where t h e c on u s ar t e r io u s p asses in t o the ven t ral aor t a ( At .

this p o in t n o t e the cor on ary ar t ery p aired p assin g t o t he c onu s ar terios u s and
, ,

to the vis c eral mu s cles P reserve it as w ell as p ossible ) F ollow ou t the p os terior
. .

bran ch Ofthe vent ral aor ta Aft er some dis tan ce it s u bdivides int o three bran ches
.

whi ch p ass t o the fou rth fth and S ixth vis ceral ar ches p ene t ra tin g the in ter
, , ,

bran chial sep t a Tra ce them in t o the sep tum by slitting the s ep tum and no t e the
.
,

bran ches from ea ch to the gil l l am ents of bo th d em ib ran chs of t he s ep tum F 0 1 .

low the ven t ral aort a forward beyond the p air of p os t erior bran ch es I t p asses .

wi thou t branchin g for som e dis t an c e and t hen forks in t o ri gh t an d l eft bran c hes .

F ol low the lef t bran ch Af t er a c onsiderable dis t an ce it forks in t o two vessels

.
,

which p enet rat e the int erbranchial sep ta of the se con d and third vis c eral arches .

The ve p airs of bran ches of the vent ral aort a are n amed the a er en t br an chial
art eries H ow m an y d em ib r an chs does ea ch su pp ly ?
.

In fron t Of the an t erior fork O f the ven t ral aor t a will be fou nd som e soft
brownish d iffu se ma t erial c omp osin g t he thyr oid gland
,
.

The aff eren t branchial art eries are the vent ral halves of the aorti c arch es
whi ch w ere p reviou sly des c ribed as p resen t in the emb ryo We n ot ed that .

there are s ix su c h ar ches in vert ebra te embryos S in c e the ad ul t elasm obra n ch

.
216 LAB ORATOR Y MANUAL T
F OR VE R E B RA E ANA OMY T T
o
has b u t ve p ai rs one p air mu s t be m issin g The missin g p air is the rs t whi ch
,
.
,

shoul d su pp ly the rs t vis c eral a r ch ; it has di sapp eared du r ing d ev elopm ent ,

p robably be c a u se t his ar c h bears no g ills .

D raw the ven tral aor ta and the afferen t bran chial art eries showin g t heir
relation t o the vis ceral arches .

6 The e fferen t b r an chial arte rie s an d the d or s al aorta

.
O p e n the ap
already formed of the oor Of the m ou th and p haryn geal c avities ; tu rn it ou t
ward an d fas t en it in tha t p o si tion The esop hagu s may b e cu t to fa cili tate this
.

p ro c ed u re .Wi t h a f o r c e p s s t rip O f
f t h e m u c o u s m e m brane from t h e roof of t h e

mou th and p haryngeal c avi ties I n the roof there will now be seen fo u r p airs
.

(dogshes) or t hree p airs ( ska t e) Of large blood vessel s ext ending from the an gles
of the gill s l i t s ob l iqu ely cau dad They are the e er en t b r an c hial art eries and
.

rep resen t the dorsal halves of the aor ti c arches Clean away t he c onne ct ive .

tiss u e from these art eries so th a t they are c learly exp osed Tra c e t hem to the
.

left and no t e tha t they disapp ear dorsal to the c ar tilages of the gill arches .

R emove t hese c ar tilag es c arefu lly This is bes t done by cu tting a c ross t h em in
.

the vis c eral ar c h g ras p in g the cu t end and loosening the c art ila ge t oward the
, ,

median dors al line Af t e r the c ar t ilages are rem oved t ra c e ea c h of the efferent
.

bran chial art e ries t oward the gil ls No t e that each is form ed at the dorsal an gle
.

O f the gill s li t by the u nion of two vessels a sm aller pretr ema tic bran c h whi ch ,

c om es from the d em ib r a n ch on the an t erior fa c e Of t he vis c eral p o u ch and a

mu c h larger pos t tr ema tic bran c h whi c h c om es from the d emib r an ch on the
-

p os t erior wal l of t h e vis c er al p o u c h N o t.e t h e s m all vessel whi c h ru ns fro m ea c h

g ill la m en t in t o t h e p r e and p os t t re m a t
-
i c bran c hes I n t he ska t e
. th e rs t
two ef feren t bran c hial art eries u ni t e t o one so t ha t there are b u t t hree p airs O f
main vessels in the roof of the p haryn geal c avi t y .

Next disse ct on t he ri ght side whi ch has been kep t in t a c t in order t o s ee

, , ,

the fu ll c ou rse of the ef feren t bran chial ar t eries R em ov e t he mu c o u s m em bra n e

.

from the oor Of the m ou th and p ha ryn geal c avi ties th u s exp osin g the ven t ral ,

p or t ions of t h e g ill ar c hes R . em ov e t hese c ar t ila g e s c ar e f u lly wi t ho u t dis tu rbin g

any of the art eries and also rem ove t he c artila ge from the ful l len gth Of the
,

vis c eral arch es I t wi l l now be seen tha t the p r e and p os t t r ema ti c bran ches
.
-

are u ni ted at t heir ven t ral ends S O that they form a c omp let e loop a rou nd ea ch
g il l c l e f
. t N o t e f u r t her t h a t th e p os t t re m a t
-
i c bran c h on the an t erior wall O f
each vis c eral ar ch is c onne ct ed wi t h the p ret rem a ti c bran ch on the p os t e rior
fa c e Of the sam e arch by m eans of c ross bran ches abou t t hree in number in the ,

sp iny d ogsh one in the sm oo th d ogsh and skat e Th u s ea ch efferent bran chial
,
.

ar t ery colle c ts from several d emib ran chs .

F rom the vent ral ends of som e of the loop s formed by the p re and p os t
t r em a t i c bra n c hes v essels arise on ea c h side and p ro c eed p os t e riorly in the oo r
of the p haryngeal c avi t y ven tral to the car til ages of the gill arches to supp ly
, ,

t he p eri c ardial c avi t y wa l l Of the heart an d vis ceral mu s c l e s

, , These vess els .
21 8 L AB ORA OR T Y MANUAL FOR VE R TE B RATE ANATOMY
s
and oril y ve rysm all bran ches t o the wall s Of the sp ira c le ) Now t u rn the anim al .

vent ral side up rem ove the mu c ou s m embran e between the sp ira cle and the
,

u pp e r jaw and shave away the c art il a ge abo u t halfway b et ween the u er jaw
, pp
and the c omm on c arot id ar te ry An art ery of m oderat e S iz e will be revealed .

Trac e it t oward t he s p ira c le and no t e t ha t it is the c o n tin u a tion Of the hyoid ean
art ery already seen form ed by the reu nion Of the branches in the sp ira cu lar
,

g ill .I t p asses in t o t h e sk u ll ven t ral t o t he e x t ernal c aro t id ar tery and is t he n ,

kn own as the ven tr al car otid art er y On t rac in g it in t o the sku ll by s craping .

away the c art ilage the vent ral c arotid ar tery will be found t o join the int ernal
,

c aro tid .

The efferent bran chial art eries p ass t o the m edian dorsal line of the roof of
the p haryn g eal c avi t y In the s kate the r s t and se c o n d join t o one and short ly
.
,

p os t erior t o t hi s j u n c t ion a ver teb r al art ery arises on ea ch S ide and p asses in t o the

c art ila g e O f the sku ll wher e it is d is t rib u t ed t o t he brain and sp inal c ord
,
The .

efferent bran chial art e ries join in p airs in the median do rsal line and fo rm a
large tr un k the dors al a orta whi ch p asses int o the p leu rop eri t oneal cavit y
, , .

Draw showing efferent branchial art eries their bran ches and their dis tri
, , ,

b u t ion t o the gills .

We are n o w in a p os itio n to co m pa re the head ar ter ies Of the el as mo r an ch b with the

p mitive pl an pl ain ed a t begin br a ries ar e
'

ri ex the n in g O f this s ec tion . T he a er en t n chial a r te

the v en tr al hal ves ,

the ries the dors al hal ves Of the aortic a ches
f
ef er en t b r an chial a r te r . These
a ches co
r ec t the v e t al a o ta s p i g in g f o m the hear t with the do rs al ao ta an d run in
nn n r r r n r r ,

the wall s o f the pharyn x o n e to ea ch v is c al a rch I n el as mob an ch an d o ther shes the

,
er . r

cen t al p a rt o f ea ch ao tic a r ch b r eaks u p i to a n u mb er O f s mall er ves s el s an d ca p ill aries

r r n

ru n n in g in the g il l s Theo etically an d in ver teb ate embryos the e a e s ix aor tic a ches I
. r r r r r . n

el as m o b ran chs the e a e v e v e tr all y a n d fo r d o s ally

r r The rs t is l ackin g on the ve tr al
n u r . n

s id e an d al s o ab se t o the d o sal S id e ; b u t the s econ d aor tic arch is imp er fec tly rep r esen ted
n n r

b y the hyoidean ar te y which s u ppl ie the S p ira cl e T he fou r effe en t b ran chial ar teries p r es
r s . r

en t in th ir fu ll d evel o p men t in el as mobr an chs a re then the thir d the fou th fth a d s ix th
e , , ,
r , ,
n

a o tic ar ches
r I t s hou l d b e n o ted tha t the ca r o tid a tery S p r in gs fro m the thir d aor tic a ch a
. r r ,

co n d ition un ive sal a mo g v er tebra tes

r T he s t ao tic ar ch is miss i g i a ll a du l t ve teb a tes
n . r r n n r r

( e cep t cycl os tom es) a d above s hes the seco d has al so va ished
x ,
n n n .

The d or s al
aor t d it s bran c he s ara t e th eso ha s fro m th body
Se
7 . a an p e p g u e .

wall on the left side and follow the dorsal aor ta p os t eriorly F rom the do rsal
'

aort a be tween the p oin ts where the third and fou rt h p airs of efferent bran chial
art eries u n it e wi t h it a s u bcl avian art ery is given Off on ea c h side Tra c e the left .

one in to t he p e c toral n I t p ro ceeds obliqu ely c au dad and la t erad p assing on

.

the dorsal wal l of the lar ge b ag fo rm ed by the p os t e r ior c ardinal sin u s ( In the .

ska t e it gives Offthe p os t erior c oron ary art e ry and p asses in t ernal to a large whi t e
ban d the nerve of the p e ct oral n Thi s m ay be cu t throu gh ) At the la teral
,
. .

boundary Of the p os t erior c ardin al sin u s the s u b clavian art ery gives rise to the ,

sm al l l ater al art ery whi ch bran ches in t o the body wall and u su ally p ro ceeds pos
terior l y alon g t he body wall in a p osi t ion on a level wi th t he la t eral line F arther .
 COMPARA I VE ANA OM
T T Y OF THE CIR CUL ATOR Y S Y STE M 219

la terally at the p oint where the lat eral vein ent ers the c omm on c ardi n al vein ,

the s u b clavian art ery gives rise t o t he ven tr al a b ddmin al ar t e ry whi c h p ro c eeds

p os t eriorly halfway be t ween th e la t eral vein and t h e m idven t ral line This .

art e r y is somewhat irregu lar in the sm oo th d ogs h b u t c onspi cu ou s in the s p iny

d ogs h and ska t e I t gives Off bran ches se gmen t ally in to the body wall an d a t
.

the p os t erior end of the p leu ro p e ri t oneal c avi t y anas t om oses wi th the vess el s
supp lying the p elvi c n s Af t er givin g off these bran ches in to the body wall
.

the s u b c lavian art ery now nam ed the br achial ar t ery p ro c eeds in t o the p ee
, ,

t oral n .

The dors al aor ta is a very large vessel situ at ed in the mid dorsal line of the -

p le u ro p eri t oneal c avi t y I t has m. edian u n p aired vis c eral or s p lan c hni c bran c hes
t o the vis cera la t eral vis c eral bran ches t o the u rog eni tal sys t em and som a ti c
, ,

branches to the body wall The median vis ceral bran ches are as follows Work
. .

on the left side tu rnin g the vis cera to the righ t .

S piny d og s h: Ju s t af ter it has p ene tra ted the p leu ro p eri toneal c avi t y the ,

dorsal aor ta gives rise to the large coeliac art ery whi ch dis t rib u tes blood t o the
g onads ,
s t om a c h and ,
liver N ear it.s ori g in th e c oe l ia c ar t ery g ives Of f s m all
branches in t o the adja cent gonads esop hagu s and c ardia c end Of the s tomach
, , .

I t then ru ns p os t eriorly for a c onsiderable di s t an ce wi thou t bran chin g ; it en t ers

the gas t rohep a ti c li gam en t and gives rise t o three bran c hes : the gas tr ic the ,

hepatic and the pan cr eatico mesen ter ic art ery

,
-
The gast ri c art ery p asses to the
.

s t om a ch and divides int o dorsal and ven tral gas t ri c art eries whi ch bran ch on
the s u rfa c e of the s t om a ch and p ene t ra t e it s wall s The hep ati c art ery tu rns
.

ant eriorly ru ns alongside t he bile du ct and ent ers the s u bst an c e of the liver
, ,
.

The p anc reati co mesent eri c art ery p asses dorsal to the p yloru s gives Off sm al l
-
,

bran ches in t o the p ylori c p ortion of the s t oma ch and the ven t ral lobe of the
p a n c reas a m odera
,
t ely lar g e du o d en a l ar t ery in t o th e d u oden u m an d a lar g e ,

an ter ior mes en ter ic art e r y alon g the ri gh t side of the s m al l in t es t ine t o whi ch it

g ives O ff bran c hes a t t h e rin gs of a t t a ch m en t of t he s p iral valve The dorsal .

aorta af ter grvm g rise to the c oel ia c ar ter y ru ns wi thou t fu r ther vis c eral bran ch es
to the free ed ge O f the dorsal mesen t ery Here it gives Off two ar teries whi c h
.

c ou rse in t he border Of the m esen t ery On e of these the gas tr ospl en ic art e ry
.
, ,

p asses t o t h e s p leen and bend O f th e s t o m a c h The o t her

. vesse l is t he s u p er ior

mesen ter ic art ery I t p asses t o the sm all in t es t ine and as the pos ter ior mes en ter ic
.
,

art ery ru ns p os t eriorly on the left side Of the in t es tine wi th bran ches c orresp ond
,

in g t o t hose Of the an t erior mesen t eri c B eyond the gap in the dorsal mesen t ery
.
,

the dorsal aort a gives Of f the inferior mesen ter ic art ery whic h p asses along the
free ant e rior border of the m esore ctum int o the re ctal gland .

S moo th d og sh: S hor tly af ter en te ring the p leu rop eri toneal c avi ty the ,

dorsal aort a gives rise to the l arge coel iac ar tery This has sm all bran ches in to .

the adja c en t gonads and soon divides in t o ga s tr ic and pan cr eatico mesen ter ic -

art eries The gast ri c art ery immediat ely forks in t o a smaller an ter ior gas tr ic
.
 2 20 LAB ORATOR Y MANUAL FOR VE RTE B RATE ANATOMY
.
a rt ery
'
whi ch p asses to the wal ls Of the ante rior p a rt of the s t om a ch and a pos
ter ior gas tr ic art e r y whi c h runs p os t eriorly in the gas t ro hep a t i c li gam en t wh e r e
it bran ches t o bo th lim bs of the s t oma c h . The p os t erior gas t ri c art ery near its
origin from the c oelia c sends abou t two hepatic art eries to the live r The .

p an c rea t i c o
- m esen t e r i c ar t ery sends p a nc r ea tic bran c hes in t o t h e p an c reas , a

la rge du oden al branch in t o the du odenum a poster ior gas tr ospl en ic art ery whi ch
,

ru ns p ost eriorly along the narrowed p art of the sp leen and adjac ent s t om a ch
wall and a large an ter ior mesen ter ic art er y which p asses along the righ t side Of
,

the sm al l in tes t ine and bran ches at the lines of a tta chm en t O f the sp iral valve .

S hort ly p os t erior t o the p oin t Of ori gin of the c oelia c ar tery two art eries a rise
from the dors al aort a They are the gas tr ospl en ic and the s u per ior mesen ter ic
. .

The gas t rosp leni c p ro ceeds to the thi ck p or tion of the sp leen and the adjac ent
s tom a ch wall The s u p erior mesent eri c su pp lies bran ches to the gonads and
.

then as the pos ter ior mes en ter ic art ery p asses alon g t he lef t side of the sm all
, ,

int es t ine bran chin g at the lines of a tta c hm en t of the c oils Of the sp iral valve
, .

The dorsal aort a p ro ceeds u nbran ched for s ome dis t an c e and t hen gives off the
infer ior mesen ter ic art e ry . This p asses in t o the adja c ent gonads t o whi c h it
s upp lies som e branches and then em erging from the g onad p ro ceeds to the re ct al
g lan d where it fo rm s a ne t work O f bran c hes .

S kate : S hor tly af ter en t erin g the p leu ro p eri t oneal c avi ty the dorsal aor ta ,

g ives O ff t h e c oeli a c ar t ery

,
wh i c h s u pp lies a n um ber of or gans
. I t has : a h ep ic
a t
bran ch to the liver ; an an ter ior gas tr ic bran ch whi ch divides in t o dor s al and
ven tr al ga s tr ic art eries t o the s t om a c h wall ; spl en ic b ran c hes t o t he s p leen ;

and a gas tr odu oden al b ran ch from whi ch arise a pos teri or gas tr ic ar ter y t o the
,

p os t e r ior p art of t h e s t o m a c,
h p an c rea ti c br a n c hes t o t he p an c reas ,
an d a du od en a l
bran ch t o the pylo ru s and du odenum S hort ly p os t erior to the o rigin Of the
.

c oelia c art ery the dorsal aort a gives rise t o t he s u per ior mesen ter ic art ery whi c h
, ,

aft er som e sm all bran ches to the p an c reas and sp leen p ro ceeds p os t eriorly along
the sm all in tes tine t o whi ch it s u pp lies bran ches a t the lines of a tt a c hm en t of
the tu rns of the sp iral valve . S hort ly c au dad of the o ri gin of t he su p erior
mesen t eri c art ery the infer ior mesen ter ic ar t ery bran ches from the dorsal aort a
,
.

It sends gen ital art eries to the gonads an d t h eir d u ct s and t hen p asses in the
m esen t e ry to the re ctal gland .

The lat eral vis c eral and som ati c branches of the dorsal aor ta are sim ilar in
the three fo rm s u nder c onsidera tion . The fo rmer c onsis t of the gen ital art eries
al ready not ed (b u t c omp le t ely develop ed onl y in m atu re sp e c imens ) and the
r en al art eries in t o the ki dneys . The latt er are s een by loosening the kidn ey from
the dorsal body wall and lookin g on the dorsal s u rfa c e of the org an . T he som a ti c
bran ches consis t Of p aired par ietal ar te ries to t he body wall p assing ou t along ,

the m yosep ta .The ar teries to the p aired n s are m erely enl arged p arie tal
vessels The s u b c lavian to the p e c toral n was al ready seen The p aired
. .

il iac a rt eries t o the p elvic n s arise from the dorsal aort a short ly in fron t of
222 LAB ORATOR Y MANUAL T
FOR VE R E B R ATE ANA OM T Y
b ran chal i pas sin g throu gh the gill l amen ts re bloo d is ae ated an d b ecomes
a r ter ies . On r

a terial bl oo d
r I t is then coll ec ted by the effe en t bra chial a t er ies an d p as s ed i to the
. r n r n

dor s al ao r ta which dis tribu tes it to all p ar ts Of the body The ven ou s bl ood is r etu rn ed to .

the hea r t by mea s o f the sys temic v ein s an d the two p o tal sys tems
n There is thu s in the r .

el as mob ran chs ( an d all shes ) a s in gl e circul ation thr ou h the hear t Sin ce the hear t co n tai
g . ns

onl y v en ou s bl o od the hear t mu s t Ob tain ar terial bl oo d fro m s o me o ts id e s ou ce

, This is u r .

a ccomp lis hed by mean s o f the co ron a ry ar tery which ar is es f om the effer e t b ran chial ves s el s r n

an d retu rn s to the hea t As we shall s ee the co o ary a t ery o rig i a tes in a dif
r . fe e t man e
r n r n r n n r

in s h than in o ther ver t eb ra tes .

9 The p eri c ar d io p e r i to n e al can al s I n sp e c t the p os terior wall Of the p eri
.
-

c ard ial c avi t y af ter the rem oval of the hear t I n t he d og s he s , a large op ening .

will be fo und in the p os terior wal l dorsal to the p r evio u s a tta chm en t of the
sinu s ven osu s On p robin g in to it it will b e fou nd to l ead in to a ca nal the
.
, ,

p er ic ar di o p er iton ea
-
l c anal si tu a t ed alon g th, e ven t ral wall O f th e eso ha
p g u s in t er
nal to the vis c eral p eri tone u m Of the eso pha gu s E ven tu ally the canal op ens .

in t o the p l eu rop eri t oneal c avi ty by a s mall sl i t I n the s kate there is an Open .

in g O f m odera te siz e in the c en t er Of t he p os t erior wall Of the sinu s venos u s .

On p robin g in t o t his it wil l be fo u nd t o lead in t o a c anal the per icar dio per iton ea l ,
-

c anal w hi c h p asses t hro u gh t he c en t er of the hep a t i c sin u s

,
On p rob in g int o this .

it will be fou nd t o fork in t o two c anals lyin g on the ven t ral wall of the es opha
g u s in t e rnal t o it s serosa They o p en in t.o th e p le u ro p eri t oneal c avi ty by m in u t e
sli ts The p eri cardio p eri toneal c anals serve to c onne ct the p eri c ardial and
.
-

p le u ro p eri t o n eal c avi t i e s and arise thro u,g h t he fa il u re O f th e t ransverse se p tu m

t o close c om ple t ely a c ross the c oelom .

C . THE ULAT OR Y
CIR C S Y S TE M OF NE CT UR U S
amb er s Of the
The ch er ardial avi t y he a rt
has The
already been
I .
p i c c

exp osed ; if a new sp ec im en is p rovided the p eri c ardial and p leu rop eri toneal ,

c avi ties are t o be o p ened as b efore The p ar ts of the hear t visible in vent ral .

view are the ven tr icl e the au r icl es and the con us a r ter iosu s The ven tri cle is
, ,
.

t hi c k walled and c oni cal in form An t erior to the ven tri cle on ei ther side is a
.

thin walled au ri cle

-
Sp rin gin g from the base of the ven tri c le and p assin g for
.

ward between the two a u ri cles is the tu b u lar c onu s ar t erios u s An t eriorly the .

c on u s p asses in t o the enlar ged be ginnin g of t he ven t ral aor ta ; thi s m u s cul ar

exp ansion Of the base Of the vent ral aorta is nam ed the bu l b us ar ter iosu s L if t .

the ap ex of the ven t ri c le and no t e the s in us ven osus S i tu a t ed dorsad t o the

vent ri cle The sinu s venos u s re c eives two large venou s channel s lyin g in the
.

dorsal wal l Of the p eri c ardial c avi t y Thes e are formed by the u nion Of the two .

hepa tic s in uses wi th the two common car din al veins The hep ati c sinu ses are the .

two large veins whi ch em erge from the t ransverse sep tum and p ass in t o the
sinu s The c omm on c ardin al veins join the hep ati c sin uses on t heir la teral
.

s u rfa c es .
 COM P ARA I VE ANA OM T T Y OF T HE CIR CUL ATOR Y S YSTE M 2 23

The he patic portal s ys tem an d the ven tr al

2. abv i
d o min al
e n I n the
.

median ven t ral line Of the b ody wall , p os t erior to the liver in closed in the fal ci
,

form ligamen t is situ at ed the ven tral a bdomin al vein I t is hom olo gou s wi th .

the la t eral abdominal veins of t he elas m obran c hs I t re c eives par ietal bran c hes
.

from the body wall At the level O f the p os t erior end of the liver it leaves the
.

body w all and p asses in t o the liver where it is S itu a ted on the dorsal s u rfac e .

Af t er a shor t c o u rse it joins the hep a ti c p ort al vein at the p lac e of a t t a chm en t
of the hep at odu odenal ligam ent .

The hepatic por tal vein is the vein whi ch c ol le ct s t he blood from the di ges tive
t ra ct . I t is form ed by the uni on of bran ches from the in t es tin e p an creas sp leen , , ,

and s t om a ch S t re tch ou t the dorsal m esent ery of the sm all in t es ti n e In

. .

thi s mesen t ery abo u t halfway be tween the body wall and the in t es tin e r uns a
c onsp i c u o u s vein t he mesen ter ic vein
, Tra c e it p os t eriorly and not e it s b egin
.

ning in the wal l of the larg e int es tine As it p asses forward in t he m esent ery it.

re ceives n umerou s in tes tin al v eins from the sm all in t es t ine I t t hen p asses int o .

the s u bs t an c e of t he p an c reas re c eivin g s m al l pan cr ea tic veins from t ha t organ .

At the level of t he p an c reas the large gas tr ospl en ic vein al so joins t he m esen t eri c
from the lef t On fol lowin g the gas t rosp leni c to the sp leen it wil l be fou nd to
.

be formed of spl en ic bran c hes from the sp leen and gas tric veins from the s t om ach .

The un ion of the mesent eri c gas t rosp len i c and p an c rea ti c veins p rodu ces a
, ,

large vessel the hepatic por tal vein whi c h lies alo n g the c en t er Of the dorsal
, ,

face Of the liv er I t also re ceives the ven t ral abdominal vein as already no ted
. .

F oll ow it alon g the s u rfa c e of the l iver I t bran c hes in t o the l iver s u bs t an c e
.

an d in it s c o u rs e also re c eives addi tional gas t ri c veins from the s t o m a c h and also

veins from the vent ral body wall whi ch p ass in tOthe liver by way of the fal ci
~

fo rm l igam en t (These have p robably been des t royed in the s tu dy of the diges
.

tive t ra ct ) . The hep ati c p ort al vein su bdivides in the su bs tan c e Of the l iver and
even tu ally p ass e s in t o c ap illaries .

Draw the hep a ti c p ortal sys t em .

3 T h re l or t al t e m Tra e t h ven t ral abdo m i n al v e i n p os t eriorly

. e n a p s ys c . e .

I t soon re c eiv es som e ves ical veins from the bladd er S hor tly an t erior to t he
.

h ind l im bs the abdom inal vein is seen to be form ed by the union of the two
p e l vic ve ins whi c h ru n alo n g th e in ner s u rfa c e of t he la t eral body wa l l j u s t in

front of the p elvi c girdle F ollow one of the p elvi c veins I t is joined by the
. .

fe m or a l vein fro m t he h in d l im b The vein for

. m ed by t his u nion is t h e r en a l

p or ta l vein . I t p asses a t on c e t o t he dorsal s u r fa c e of th e adja c en t kidney I n .

m ale sp e cim ens the kidney is a brownish organ of c onsiderable siz e situ a ted at
the side Of the in t es tine In fem ale sp e cim ens the ki dney is mu ch sm al ler and
.

m ore slender and is si tu a t ed a t the c omm on p oin t of a tta chm en t of the mes ova
,

r ium and m es o t u b a r ium

'
I t will be lo ca t ed by layi n g the ovary t o one side and
.

the ovid u c t t o the o ther side At the p os t erior end of the p leu rop eri t on eal c avi ty
.

the ki d ney in the f em al e li e s be tween t he in t es t ine and the ovidu ct H aving

.
2 24 L AB ORA OR T Y MANUAL F OR VE R E B RA E ANA OM T T T Y

lo c at ed the kidney iden tify the renal p or tal vein hear t he lat eral m argin of its
,

dorsal s u rfa c e At the p la c e where t he r en al p or tal vein p asses from t he body

.

wall to the su rfa c e of the kidney the vein re ce ives the caud al vein whi ch as c en ds
, ,

fro m the tail forks and p asses to the su rfa c e O f the p os terior end of ea ch kidn ey
, , .

FI G .
57
Dia ram s t o
.
g s ho w t he o rig in o f the p o s tc ava l v ein an d the cha n g es in the ab do mi n al
vein in a p
m hib ia n s a n d re p til es . A , el a sm o b ra n ch s ta g e , sa m e a s
in Fig u re 556 ; t he l a teral a b d omin a l
v ein s i en t er t he c o mm o n c a r d in al v ein s c a n d a r e n o t co n n ec t ed with the r en al or t al v ein s p p
B , the .

l a t eral a b do min al s i hav e j oin ed the ren al p o r ta l s a t t p os terio rl y , a n d a n t erio rl y p ass in to the liv er f,
where they u nite with the hepa tic po r tal vein h; a n ew vein the pos tcaval vein g is seen gr owin g cau dad , ,

from the liver f where it a ri es from the hep atic v ein s 0 C con dition in the adul ts O f u rodel e amphib i
s .
,

a n s ; the po s t cav al v ein g ha s r ea ched a n d f used with t he p o s terio r car din al s e a n d the su b ca r din al s j

a t the p o in t r ; t he t wo l a t er al ab d o min al v ein s hav e u nited t o form the ven tral ab d omin al vein i which

em p ti es in to the he p a t ic p o r ta l h . D, co n ditio n in a d ul t re p til es ; t he a n terior po r tio n s O f the po s terio r

ca r d in al v ein s n a r e o b literat ed , l eavin g the p o s tcaval v ein g as the sol e d rain a g e for the s u b c ard in al s j
an d the kidn eys min al vein s r emain s e ara te as in el asmob ran chs
k; t he t wo l a teral ab d o a , an terio r p .

ca r d in al v ein ; b , sin u s v en osu s ; c , c o mm on c ar di n al v ein ; d , s u b c l avian v ein ; e, os t erior c a r din al p

vein ; f, liver ; g , p
o s t cav al v ein ; h, he at ic p p
o rt al v ein ; i , l a teral ( or in C ven tr al ) a b d omin a l v ein ;

j, su b c ardin al v ein ; k, kidn ey ; l , iliac or femo ral v ein ; m, cau dal v ein ; n , o b litera ted ar t O f the p o s p
t erio r car din al s ; o , he p at ic v ein s ; p, r en al p o r t al v ein s ; q, p el vic vein s ; r , u n io n O f p o s tca val , p o s terio r

ca r d in al s , an d s u b car din al s ; s , u nio n o f p o s tca va l a n d su b c a rd in al s ; t, u nio n O f ab d o min al v ein with

ren al p o r tal s ys t em .

The renal p or tal vein r u n s forward alo n g the s u rfa c e of the kidn ey in to the s u b ,

s tan ce O i whic h it sends nu m erou s bran c hes I t also re ceives bran ches from the .

body wa l l .

Draw the ren al p or tal sys tem .

I t will be seen tha t the ven tral abdo minal vein form s a c o n n e c tion be tween
the r enal p or tal and h ep a ti c p or tal sys tem s B lood fro m the hi n d legs and tail .
2 26 LAB ORA OR T Y MANUAL F OR VE R E BRA E ANA OM T T T Y
.
s to m ach to the lef t an d the liver to the ri gh t I t p asses forward im bedded in .

the liver s u bs tan c e a n d sho u ld be followed by p i cki n g away the liver tiss u e It .

r ec eives several hepatic vein s fro m the liver ; o n e of the larger Of these lies alo n g
the midven tral line of the liver a n d joins the p os t c aval near the an t erior end of
the liver . At the an terior en d of the liver the p os tc aval vein em erg es as a very
large vessel si tu a ted in the c oron ary ligamen t I t pier c es the transverse sep tu m .

a n d forks in to the two he p a ti c si n u ses whic h af ter bei n g joined by the c o mm on ,

c ardinal vein s en ter the sin u s venosu s The ori gin Of the p os tc aval vein is
,
.

dis cu ssed below .

c) The pos ter ior ca r din al vein s : At the p la c e where the p os t c aval vein tu rns

ve n trally toward the liver it is conne c ted wi th a p air Of vei n s the pos ter ior ,

ca r din al veins Tra c e these an teriorly They lie very near the mid dorsal li n e
. .
-

of the an terior half of the p leu rop eri toneal c avi ty o n e to ei ther side of the dorsal ,

aor ta I n fem ales they are si tu a ted in the meso tu b ariu m along the line wher e
.
,

this u ni tes wi th the dorsal w al l Tra ce the p os terior c ardi n als p os teriorly an d
.

no te co n ne c tions be tween the m and the renal p or tal vein s ( whi ch are of cou rse
the ori ginal p os t erior ends O f the p os terior c ardin als as shown in Fig No t e , .

also the parietal vei n s whi ch en ter the pos terior c ardinals in their co u rse along
the body wall The pos terior cardinals may be tra ced to the t ransverse sep tu m
. .

S hor tly before rea chin g this they diverge fro m ea ch o ther and p en e tra ti n g the
la teral p or tio n s of the sep tu m en ter the c o mmon c ardinal vein pra c ti cally a t the
sam e p oin t as the en tran c e of the j u gu lar an d the s u b clavian .

5 Th
. e p u lm on a ry vein s The pu l mo n a r y vei n is a lar g e vessel si tu a t ed
along the ven tral side of ea ch lu ng i e the side Opp osi te tha t whi ch is a tta ched
,
. .
,

to the dorsal wall The two pu lmonary veins ru n forward in the walls of the
.

lu n gs an d shor tly c au dad Of the transverse sep tu m co n verg e and a t the s ep tu m

u ni te to o n e vessel Th is passes throu gh the tra n sverse s ep tu m and ru n n in g
.

forward in the dorsal wall Of the lef t hep a ti c sinu s en ters the lef t au ri cle .

6 The ven tr al aor ta an d the ao r ti c arche s

.
The c on u s ar teriosu s p ass es .

an teriorly in to the ven tral aor ta The grea ter p ar t of the ven tral aor ta li es
.

wi thin the p eri c ardial cavi ty and owing to the fa c t tha t this por tion of the aor ta
is exp anded an d p ossesses thi cken ed mu s cu lar walls it is n amed the b u l b u s ,

ar ter ios u s Tra c e the ve n tral aor ta forward ou t Of the p eri cardial cham b er
.

by disse c ting away the an terior wall of the chamber The ven tral aor ta very .

soon forks in to two vessels whi ch p ass to the righ t an d lef t Tra ce the righ t one .
,

sin ce the vis ceral ar ches have been lef t in ta c t on tha t side F ollow it to ward .

the gill ar ches I t soo n divides in to two vessels an d s u bs equ en tly the p os terior
.

The t erm b ul b u s a rt eriosu s is very a m b ig u ou sl y us ed in m a n y t exts O f v ert eb ra t e a n a t o my a n d

e mb ryo l o gy . T he t erm shou ld be a ppl i d e o nl y to the ex p

m u s c u l a r b a s e o f the v en tral a o r t a
a n d ed .

p
T he b u l b u s ar terio su s do es n o t ta k e ar t in the hea r t b ea t a n d is n o t a cha mb er O ft he hea r t b u t a o r tio n p
o f the v en t r al ao r ta T his d enit io n d oes n o t c o rr es o n d with the o n e g iven in K
. p Very few g ro u p s .

o fver t eb r a t es ha v e a b u l b us ar t erio su s ; the c hief g ro u p o ss e s s in g it is th p

e T el eos tei T he t erm tru n cu s .

a rt erio su s is a n o t her am b igu o u s n am e p

I t sho u l d ro b a b l y b e u s ed as s yn o n ym ou s with ven tr al ao rt a
. .
 COM P ARA I VE ANAT OM T Y OF T HE CI R CUL AT OR Y S Y STE M 227

one a gain divides in two making a to tal Of three affer en t b r an chial ar ter ies one
, ,

to ea c h of the gills Tra c e ea ch one in to the gill i emovin g the skin fro m the
.
,
'

g ill . A t th e e n t ra n c e in t o th e g ill t h e rs t a ff eren t bran c hi al ar tery g ives O ff

an extern al carotid ar tery which tu r n s medially ru n nin g beside the bra n chial ,

ar tery an d then bran ches in to the oor of the mou th Wi thin the gill ea ch
, .

bran chial ar tery sen ds u p a loop whi ch bran ches am o n g the gi l l l amen ts from ,

whi ch o ther bra n ches colle c t in to a loop on the o ther side of the gill this loop ,

joi n ing the bran c hial ar tery again In addi tion to the two loop s a shor t con .
,

me eti n g bra n ch ru n s thro u gh the base O f ea ch gill .

Nex t tu rn ba ck the ap p revio u sly form ed of the oor O f the mo u th and

, ,

p hary n g eal c avi t ies E x t e n d th e. in c ision p os t eriorly alon g th e lef t side O f the
esop ha gu s S t ri p Off the mu co u s membrane fro m the roof of the mou th and
.

p hary n g eal c avi t ies I n th e roof . w i l l be see n a p a i r of lar g e vessels the r oots ,

or r adices of the aor ta whi ch p ass obliqu ely p os teriorly and u ni te Tra ce the
, .

righ t roo t la terally i n to the vis ceral ar c hes disse c tin g Off the mu c ou s mem brane
from the la tter L o ca te from the inside the branchial ar teries exp osed in the
.

p re c edin g p ara g ra p h and no t e t heir e m er g en c e fro m t h e dorsal ends O f t he vis c eral

ar ches as the eeren t bran chial ar teries The se cond and third e fferen t bran chial .

ar teries u ni te as they e m erge from the vis ceral ar ches th u s forming two efferen t ,

bran chial ar teries o n ea ch side F ro m the co mmon vessel formed by the u nion .

of the se co n d and third efferen t bran chial ar teries an ar tery arises whi ch p asses
p os t eriorly Tra c e. it in t o th e p le u ro p eri t oneal c avi t y and no t e th a t it c o u rses
alo n g the dorsal or a tta ched side of the lu n g I t is the pu l mon ary ar tery F ro m . .

the rs t ef fere n t bran c hial ar tery very near the p la c e where it join s the se c ond
a n d third arises the in ter n al ca r otid ar t ery whic hp a ss es forward on the roof O f

the m o u th Ac c o mp a n yi n g the in tern al caro tid ar tery is the in tern al ju gu l ar

.

( a n terior cardin al ) vein The efferen t bran chial ar teries medial to the ori gi n
.

of the pu lm onary an d in ternal caro tid ar teries u n i te to form the roo t Of the aor ta
on ea ch side F ro m this sp ri n g s the ver teb ral ar tery whi ch p asses a t on ce dorsally
.

in to the S k u ll The two roo ts of the aor ta then pass obliqu ely cau dad an d
.

u ni te in the m id dorsal line to form the larg e dor s al a or ta

-
.

Draw the a fferen t an d efferen t branchial ar teries and their bran ches .

T he aeren t an d ef
feren t bra chial
n a r te ries are as in the el as mobran chs ,
the ven tral an d

d o rs alhal ves es pec tively o f the a or tic arches B es ides the n etwork in the g ills each arch
,
r ,
.
,

forms a compl ete s emici cl e by mean s o f the direct co n ec tin g bran ch in the bas e Of the g il l
r n .

T he thr ee ao tic ches p r esen t in Nectu r u s corres pon d to the thi d fou r th an d s ix th a ches
r ar r , ,
r

o f the o ig i al s i
r nNo te the o rig in of the extern al c a o tid fro m the ve tral p a r t o f the in tern a l
x. r n ,

caro tid fro m the d o s al p ar t O f the third ao r tic ar ch an d o f a n ew ves sel the p ul mon a ry ar tery
r , , ,

f om the l as t or s i th arch ( see Fig 58 C p

r x .
,
.

7 . The b r an che s of the d or sal a rta . o Tra c e

dorsal aor ta p os teriorly the
in to the p l eu rOp eriton eal c avi ty I t ru n s in the m edian dorsal lin e
.
.
I mme di .

a tely beyo n d its origi n it g ives O f

f a s u b cl a vi an ar t ery o n ea c h side Tra c e one .
2 28 LABORATOR Y MANUAL FOR VE RTE B RATE ANA OM T Y

Of t herh I t p asses la terally I t gives Off a co n sp i cu ou s cu tan eou s ar tery which

. .

lies on the inner su rfa ce of the p ec toral girdle I t bran ches to nearby mu s cles .

and the skin The su b clavian then gives Off an ar tery to the shou lder and as
.

the br achial ar tery p asses in to the fore lim b where it bran ches ex te n sively .

I n its co u rse alo n g the p leu rop eri toneal cavi ty the dorsal aor ta giv es off bo th
vis ceral and soma tic branches The median vis ceral branches will be des cribed
.

rs t .

The r s t vis ceral bran ch O f the dorsal aor ta is the gastr ic ar tery I t p asses .

to the s to m a ch and forks i n to the dor s al an d ven tr al gas tr ic ar teries s u pp lyin g the
c or resp ondin g walls of the s to m a c h The ven tral gas tri c ar tery also fu rn ishes
.

a few sm all bran ches to the sp leen S om e dis tan ce p os terior to the origin of
.

the g as t ri c ar tery the coel iaca mes en ter ic ar tery sp ri n gs fro m the aor ta
,
-
It .

p asses ven t rally in t he m esen t ery givin g rise to so m e m es en ter ic bran ches to the
beginn ing of the sm all in tes tine ; it then p ro ceeds to the regio n of the hep a to
du oden al li gamen t where it bran ches in to a s pl en ic ar tery to the sp leen a ,

p an cr ea tico d u-
o d en al ar t ery t o t he p an c reas d u oden u m and , p ylori c re gion of
the s to m a ch and a hepa tic ar tery whi c h ru ns al o n g the dorsal s u rfa c e Of the
, ,

liver in c on ta ct wi th the hep a ti c p or tal vein and s upp lies n u mero u s bran ches
to the liver s u bs tan ce P os terior to the p oin t Of ori gin of the co eliaco
.

m esen teri c vessel the dorsal aor t a gives Off a num ber of mesen ter ic ar teries in to
,

the i n tes ti n e .

The la teral vis ceral bran ches Of the dorsal aor ta c o n sis t of n u m ero u s gen ital
ar teries to the tes tes in the male and ovaries and ovidu c ts in the female and of
, ,

ren al ar t eries to the kidneys The so ma ti c bran ches O f the aor ta co n sis t O f the
.

p ar ie ta l or i n ter cos tal ar t eries These arise

. fro m the dorsal side O f th e aor t a a t
se gm en tal i n tervals ; they p ass dorsally and di vide in two one bran ch goin g to ,

ea ch side of the body These bran ches p ass la terally alo n g the in tern al s u rfa c e
.

of the body wall an d su pp ly the body mu s cu la tu re .

Near the p os terior end of the p leu rop eri toneal c avi ty the aor ta gives Off ,

on ea ch S ide an il iac ar tery which p asses la terally alongside the fem oral vein
,

toward the hi n d lim b I t gives Off an epigas tr ic ar tery whi ch ru ns an teriorly

.

along the body wall a hypogas tr ic bran ch to the u rinary bladder an d cloa ca an d
, ,

as the femor al en ters the hind l imb in to which it shou ld be tra ced I t ru n s
,
.

along the m edial side Of the l eg and a t the kn ee gives rise to a n u mber of bran ches .

The dorsal aor ta p ro ceeds in to the tail as the cau dal ar tery giving Off a p air of ,

cl oacal ar teries in to the cloa ca as it p asses tha t re gion .

Draw the bran ches of the dorsal aor ta .

8 The chamb er s of the hea rt

.
R em ove the hear t fro m the p eri c ardial
c avi ty by cu t ti n g a c ross bo th ends The chambers of hear t were p reviou sly
.

nam ed The sin u s ven osu s is a chamber wi th very thin deli ca te walls I t
.
,
.

re c eives fro m behind the two large tru n ks form ed by the u n ion O f the co mm on
c ardinal vein a n d hep a ti c sin u s on ea c h side An teriorly the si n u s p asses in t o
.
2 30 L AB ORA OR T Y MANU AL FOR VE R E B RA E ANA OM T T T Y
0
has Shifted its an terio r c on n ec tion s . Whereas '

in s hes it

o p ens in to the co mmo n ca r din al

vein , in Amphibia it p as ses i n to hep atic po tal sys tem The b domin al vein th s b comes
the r . a u e

a co n n ec tio n b etwee n the e al an d hep a tic po r tal s ys tems an d p ovid es two ou tl ets fo r the
r n r

bl ood f ro m l egs an d ta il . The pu lm o ary vein s ar e n ew forma tion s

n .

D. THE CIR C ULAT OR Y S Y S TE M OF TH E T UR TL E

Sp e ci m en s for the s tu dy Of this sys tem S hou ld have been dou bly in je c ted ,

tha t is i n to the ar terial sys te m an d in bo th dire c tio n s i n to one ve n tral abdo minal
, ,

vein R emove the p las tro n Wi th the bone s cissors cu t away the sides Of
. .

the c arap a c e be twee n fore an d hi n d li m bs .

I .The chamb er s Of the he art The hear t of the tu r tle p ossesses b u t three
differen t chambers in c on tras t to the fou r chambers p resen t in the form s already
,

c onsidered ; one of the c ha mbers is divided in t o two c o mp le tely se p ara te halves .

E x a mine the hear t in the p eri c ardial c avi ty rem ovi n g the ven tral wall Of the ,

p eri c ard i al sa c if t his has n o t already been do n e F r om th e ven t ral view the .

visible p ar ts of the hear t are the ven tr icl e the thi ck walled c oni cal p os terior -
,

ar t d th i l t hin walled c ham bers one on ea c h side an t erior to the

p a n e a u r c es -
,

ven tri cle The two au ri cles are en tirely sep ara te fro m ea ch o ther as will b e
.
,

seen la ter The ven tri cle is a tta ched to the p os t erior p eri cardial wall by a
.

ligam en t whi ch is app aren tly a remn an t Of the ven t ral m esen tery or ven tral
,

m eso car diu m Of the hear t a s t ru c tu re w hi ch was c o n sidered in the g eneral dis
,

cu ss io n Of the c oelo m Cu t thro u gh this li gamen t raise the ven tri cle an d p ress
.
, ,

it forward A larg e chamber the s in us ven os u s is revealed dorsal to the a u ri c les

.
, ,

a n d a tta ched to the ri g h t a u ri c le The large bases Of the sys temi c vei n s will b e
.

seen en terin g the sinu s P u t the ven tri c le ba ck in p la ce Observe the lar ge
. .

vessels ar teries which sp rin g dire c tl y from the base of the ven tri cle wi thou t
, ,

the in terve n tion Of a c onu s ar terios u s The la t ter is la ckin g or to be very

.
,

a ccu ra te is so shor tened down a s to be invisible The bases Of the ar teries which
,
.

sp rin g from the ven tri cle c orresp ond to the ve n tral aor ta of elas m obran chs The .

ven tral aor ta is thu s seen to have sp li t in to several sep ara te tru nks The ar teries .

will be inves tiga ted la ter and shou ld be lef t u n dis tu rbed a t p resen t .

2 .The ve n tr al ab d o min al ve in s an d the r e n al portal s ys te m Ru n ni n g in .

the ven t ral p le u rop eri to n eu m fro m the p elvi c girdle u p to the hear t are two
large veins the ven tral a bdomin al veins They are ho m ologou s wi th the la teral
,
.

abdo min al vein s of elas m obran chs The two vei n s are generally con n e c ted j u s t
.

an terior to the p elvi c girdle by a c ross bran ch Tra c e the veins forward They -
. .

re ceive per icardial vein s from the p eri cardial sac an d then ea ch tu rns dorsally
to en ter the liver Ju s t a t this tu rn each vein re c ei ves a pector al bran ch from
.

the p e c toral mu s c les O f tha t side Tra c e the p e c toral vein in to the mu s cle
. .

S li t the p l eu rop eri to n e u m alo n g side ea ch abdo m in al vei n and by lif tin g the cu t
edges n d the p la c es where the vein O f ea c h side p en e tra tes the lobe of the liver .

T ra c e the ve n tral abdo min al veins p os teriorly Make a lo n gi tu din al sli t .

in the p leu rop e ri to n eum midwa be twee n the two vein s an d sep ara tin g the cu t
.
,
 COM P ARA I VE ANA OMT T Y OF T HE CIR CUL ATOR Y S Y STE M 23 1

ed ges look wi thin an d lo ca te the u rinary bladder No te the small vesical vein
, .

p assi n g fro m th e bladder in to ea c h abdo m inal vei n . C on ti n u e t o t ra c e the

abdomin al vein s p os teriorly E a ch p asses to one side of the p oin ted an terior
.

ex tremi ty of the p elvi c girdle and a t the sam e time g radu ally tu rn s la terally .

As it tu rn s it re c eives a pel vic vein whi ch r u n s over the ven tral s u rfa c e Of the
mu s cles O f the p elvi c girdle . The lef t p elvi c vein seems to be u su ally larg er than
the ri gh t on e .

I n their c o u rse be twe en the hear t and p elvi c girdle ea ch vein gives Off la terally
o n e or m ore s m all bran c hes whi c h p ass to the borders of the c ara p a c e wh e re t h ey

join the margin o cos tal ve in to be des cribed la ter

-
.

Draw the two abdomin al veins wi th their bran ches th u s far no ted .

Con tinu e to tra c e the abdo minal vei n s in the p os terior di re c tio n . As bo th
have iden ti cal bran ches it is n e c essary to follow o n ly one sele c ti n g the o n e which ,

has bee n m os t su ccessfu lly inje c ted I t p asses alon g the dorsal su rfa ce O f the
.

p elvi c g irdle n ear th e an t erior m ar g in of t h e la t t er t h e g irdle sho u ld be p u lled

toward the s tu den t in order to follow the v ein . G rasp the hi n d l eg on the S ide
on whi ch you are disse c ti n g an d work it ba ck an d for th u n til it is freely movabl e .

P ress the l eg away fro m the carap a c e O ftha t side and cu t thro u gh the ski n be twe en
the leg and the c arap a c e ba ck to the en d of the tail .R em ov e the skin fro m l eg
an d tail . No w tra c e the abdo mi n al vein la t erally alo n g the base O f the l eg .

J u s t beyond the p elvi c v ein a sm all cru ral from thigh m u s cles an d a larger vein
fro m fat en t er the abdo m in al Abo u t an in ch an d o n e half la t eral to this
.
-

t he l arge femor a l v ei n em erg es fro m the l eg an d joi n s the abdo m i n al vein ,

n o w desi gn a t ed the iliac v ein .The f emoral vein sho u ld be followed in to the
l eg by sep ara tin g the m u s cles . The ilia c vein is n ow si tu a ted alon gside a c o n spic
uo u s ar tery the ep i gas tri c ar tery bo th bei n g i m bedded in the abdo mi n al wall fro m
, ,

which sm all vein s p ass in to the ilia c vein Af ter a shor t dis tan c e the ilia c vei n
.

re ceives the epigas tr ic vein whi ch a cc omp an ies the ar tery of the sam e n am e
an teriorly alo n g the cu rve of the c arap a c e The ilia c vein now tu rns abru p tly
.

p os t eriorly and r u n s be twee n the base of the l eg a n d the c arap a c e dee , p ly

imbedded in som e loose tissu e This tissu e S hou ld be cleared away and the
/ .

vein followed I t re ceives bran ches from the carap a ce an d n ear the pos terior
. -

ar t f t h thi h a well m
-
arked s cia tic vein fro m the thigh P os t erior to this
p O e g .

p oi n t it re c eives several s m all bran c hes fro m the l eg and as the cau dal vei n

p ass e s alo n g t h e side of t he t ail re c,

eivin g a t th e base O f th e t ail a cl oaca l bra n ch

from the an al region .

R e tu rn to the p oin t where the ep i gas tri c vein en ters the ilia c vei n At this
.

la e a lar e vein c on tin u es forward fro m th e a n t erior and dorsal s u rfa c e of the
p c g
iliac This vein the r en al por tal vein ru n s forward and dorsally p en e tra ting the
.
, , ,

p le u ro p e ri t o n e u m . C u t th e p le u ro p eri t one u m t ra n sversely halfway be twee n

the h ear t a n d p elvi c girdle cu t ti n g a c ross bo th abdo mi n al vein s

,
Cu t also in to
.

the p u ple ro eri t one um a t the p la c e where th e re n al p or t al vein p asses t hro u g h it .

A lay er of m u s c le will be fo u n d o u tside the p eri t o n e u m a t this p la ce B o th

.
23 2 LAB ORATOR Y MANUAL T
F OR VE R E B RA E ANA OM T T Y
mu s cle s and shou ld be sli t ven trally t6 m ee t the tran sverse incision
m embran e
a cross the p leu rop eri toneu m I n thi s way free a ccess is gained to the p leu ro
.

p eri t o n eal c avi t y Wi t

. h th e lef t ha n d c arefu lly p ress all Of the vis cera forward .

I t is u s u ally n ecessary to de ta c h the lu n g fro m the dorsal wall an d p u sh it for

ward also Wi th the ri gh t han d p ress the p elvi c girdle cau dad A sp a c e cleared
. .

O f vis c era is th u s lef t dorsal to and in fron t O f the p elvi c girdle L ook in to this
.

p la c e n ear the m edian dorsal li n e for a so m ewha t a tten ed org an the kidn ey , ,

si tu a ted agains t the m edi an dorsal wall The kidney is re trop eri toneal tha t is
.
, ,

dorsal to the pleu rop eri toneu m Thi s la tter membrane sho u ld be s trip p ed off
.

fr p m the ven tral fa c e of the kidney ( In male sp e ci m en s the ro u nd ed yellow

.

t es tis a n d bla ck c oiled ep ididymis will be no ted a tta c hed to the ve n tral s u rfa c e
O f the kidney ) The ren al p or tal vein may now be followed from the p oin t wh ere
.

it leaves the ilia c thro u gh the p leu rop eri tone u m t oward the kidney B efore .

rea chin g the kidn ey it re ceives a vein fro m the c arap a ce At abou t the middle .

O f the la t eral border of the kidney is a ss u re ; the re n al p or tal vei n e n ters t his

ssu re and p asses on to the ven tral fa c e O f the kid n ey where it i mm edia tely
forks On e O f its bran ches the ver teb ral vein ru ns forw ard an d may be tra c ed
.
, ,

in well inje c ted sp e cim en s by sep ara ting the lu n g fro m the carap a c e and raisin g
-

the l u n g and also s t ri p p in g Of f the p le u rop eri toneu m fro m the dorsal wall The .

ver tebral vein p asses an teriorly dorsal to the ar ches of the ribs an d re ceives
la terally an in tercos tal bran ch at ea ch s u tu re be tween the c os tal p la tes Of the
c arap a c e The in ter c os tal veins a n as tom ose wi th ea ch o ther in the cu rve O f
.

the c arap a c e by m ean s O f a lon gi tu din al vessel the margin o cos tal vei n whi ch ,
-
,

is the an terior c on ti n u a tion Of the epi gas tri c vein p reviou sly no ted The .

m argi n o cos tal vein also has c onne c ti ons wi th the abdomi n al veins
-
The .

p os t erior bra n c h of t h e re n al p or tal vein p asses p os t eriorly over the ve n tra l

fa c e Of the kid n ey and as the in ter n al ilia c or hypogastric vei n re c eives branches
from the rep rodu c tive organs ( male) bladder c loa ca etc The ren al p or tal
, , ,
.

vein in its p assa ge alon g the ven tral fa c e O f the kidn ey gives off bran ches in to
tha t orga n The renal p or tal vei n is the p os terior p ar t of the p os terior cardi n al
.

vei n (F i gs 55 an d . The ver t ebral vein is form ed by the lo n gi tu di n al fu sio n

of segm en tal bran ches Of the p os terior c ardi n al vein of the em bryo .

D raw these vein s as far as you have seen the m addi n g the m to the drawin g ,

of the abdo mi n al veins already m ade .

The s tu den t shou ld c on sider a t this p oin t the differen c es be tween the con
n ec tion s O f the ven t ral abdo m in al veins of the tu r tle a n d their ho m olo gu es the ,

la teral abdo min al veins O f elasm obran chs In the tu r tle these vein s have form ed
,
.

a c onne c tion with the ren al p or tal sys tem p os teriorly while an teriorly they en ter
the l iver in s tead of the c ardi n al sys tem .

3 T h e h e ti or t l y t e m L if t t h lobes of th e liver se p ara t i n g the m

.
p a c p a s s . u p e

g e n t l y fro m t h e s t o m a c h and d u ode n u m and nd,

on t heir dorsal s u rfa c es a t the

p la c e w her e t h e g as t ro h ep
-
a t o d u ode
-
n al li g a m e n t is a t t a c hed to t h e liver a lar g e
234 LAB ORA OR T Y MANUAL T
FOR VE R E B RATE ANA OM T Y

it may be followed forward As bo th have iden tical bran ches it is ne cessary

.
,

to follow onl y one The one whose bran ches app ear to be l led wi th blood
.

shou ld b e sele c ted ; the lef t one is u su ally easier to follow In sp e cim en s whi ch.

have been p reserved for a long ti me the disse c tion Of the branc hes Of the p re caval
veins is generally u nsa tisfa c tory be cau se the bran ches are Of ten emp ty b u t of , ,

those nam ed below as m any as the c ondi tion O f the s p e cim e n p ermi ts sho u ld
,

be iden tied B e very carefu l n o t to i n j u re the adja cen t ar teries sp rin gin g from
.

the ven tri c le Tra c e the p re caval forward ou t Of the p eri cardial s ac Shor tly
. .

a n terior to the p la c e where the p re caval p ene tra tes the p eri cardial sac the v ein
rec eives pra c ti c ally sim u l taneo u sly fo u r t rib u taries three sm all and one large
, ,

Of ie
. The mos t m edial bran ch is the s mall thyr eo scapu l ar vein whi ch c ollec ts
-

a bran ch from the thyroid gland ( the gland si tu a ted in the fork Of the large
ar teries ) and then p asses to the inn er su rfa c e of the shou lder where it c olle c ts
fro m several mu s cles L a t eral to this vein is the sligh tl y larger in tern al ju gu l ar
.

vein This ru ns an teriorly along the side O f the n e ck in c on ta c t wi th a white

.

n erve ( va gu s or t en th c ra ni al nerve ) I t re c eives m e di ally an ex te n sive n et

.

work Of bra n ches fro m the esop hagu s I t may be t ra ced a n teriorly to the base
.

O f the sk u ll fro m w h i ch it iss u es m akin g an anas to m osis wi th the ex tern al j u gu lar

,

vein soon to be des cribed The third trib u tary Of the p re caval vein is the larg e
,
.

s u b cl avian vein by far the lar ges t Of the fo u r bra n c hes whi c h en t er the p re caval
, .

I t p asses alo n g the side O f the ne ck a n d as the a xill a r y vein tu rn s toward the
sho u lder Here it is seen to be form ed by the u nion of two large bran ches
.
,

the ex ter n al ju gu l ar vei n fro m the n e c k a n d the b r ac hia l fro m the fore li m b The .

ex tern al j u gu lar lies alo n g the side of the n e ck la teral and dorsal to the i n t ernal
,

j u gu lar I t coll ec ts from the head and in its p assage p os teriorly alon g the ne ck
.
,

has a t regu lar i n tervals ver teb ral veins p assi n g in to it fro m be twee n the ver tebrae .

Near its j u n c tio n wi th the bra chial it re ceives the las t Of the ver tebral vein s
whi ch des cends fro m the j u n c tion be tween las t c ervi c al a n d rs t tru n k ver tebra ,

where it c o n n e c ts wi th the an terior end of the ver tebral vein des cribed wi th the
ren al p or tal sys tem The ex t ern al j u gu lar vein also re ceives bran ches from the
.

S kin a n d m u s cles O f the sho u lder re gio n The fou r th and m os t la t eral an d dorsa l
.

of the trib u taries Of the p re caval is a s m all s capu l ar vei n which c o m es fro m the
mu s c les c overin g the s c ap u la .

Draw the bran ches of the p re caval as far as you have fo u n d the m .

b ) The l eft hepatic vein : The lef t hep a ti c vein sho u ld be t ra c ed i n to the lef t
lobe of the liver fro m whi ch it colle c ts venou s blood To do this clear away the
.
,

in terven ing p os terior wall of the p eri c ardial sa c an d p leu rop eri to n eu m .

c) The pos tc a val vein : Tra ce the p os tca val vei n p os t eriorly i n to the ri g h t lobe

of the liver I ts co u rse m ay be followed by m aki n g a sli gh t hole in the vein

.

where it en t ers the si n u s an d p robin g p os teriorly i n to the hole disse c ti n g away ,

the liver s u bs ta n c e alo n g the p robe No te the n u m erou s hep a ti c vei n s which
.

e n ter the p os tcaval d u ring its p assage throu gh the liver Find where the p os t .
 COM P ARA IVE ANA OM T T Y OF T HE CI RCUL ATOR Y S Y STE M 23 5

c aval en ters the liver from behind to the ri gh t of the hep a tic p or tal vein At thi s .

p oin t th e serosa O f the liver is f u sed to t he p l eu r op etito n ea l m e m bra n e over the

ven tral fa ce Of the lu n g This fu sion shou ld be broken and the p os tcaval vein
.

freed On following it p os teriorly it wi l l be fo u nd to swerve toward the median

.

l ine where it r u ns along side a lar g e ar tery ( dorsal aor ta ) The p os tcaval may .

be tra ced to the p os terior end of the pleu rop eri to n eal c avi ty I ts rela tions there .

wil l be des cribed la ter .

Add the lef t hep a ti c and p os tc aval vei n s to you r drawin g Of the p re caval vein .

5 The pul mon ar y vein s

.
A pu l mon ary vein p asses fro m ea c h lu ng to the
.

lef t au ri cle of the hear t I t is si tu a ted p os terior to the bro n chu s where it shou ld
.

be iden tied Follow it toward the hear t I t p asses dorsal to the p re caval
. .

vein The ri gh t p u lmonary ru ns in the dorsal wall of the p eri cardial sac an terior
.

to the sinu s venos u s and joins the lef t vein a t the en tran c e O f bo th in t o the lef t
au ri cle The p oin t Of en tran ce is near the lef t p re caval vein
. .

6 The a orti c ar che s an d their b r an che s

.
F ro m the ven t ri cle three large
ar terial t ru nks ex tend forward To ge ther they cons titu te the ven tral aor ta whi ch
.

m u s t be c o n ceived O fas havi n g sp li t in to t hree t r u nk s Clean away the co n n e c .

tive tissu e fro m these ar teries and sep ara t e the m fro m ea ch o ther The tru nk .

far thes t to the lef t is the pu l mon ary ar tery ; the vessel nex t to it is the
l eft aor ta; the thi rd an d ri gh t hand tru nk is the r ight aor ta b u t it is c on cealed
-
,

from View by the large bran ch the br achiocephalic (i n nomina te) ar tery whi ch
,

it gives Off i mm edia tely on leavi n g the hear t No te the s m all cor on ary ar teries
.

sp rin gin g from the base of the bra chio cep hali c ar tery and bran chi n g over the
su rfa ce of the hear t The bra chio cep hali c ar tery lies in the m edian line an d
.

forks a t on c e in to large bran ches I n the angle Of the fork lies a reddish body
.
,

the thyr oid gl an d .

a ) The b r an ches f
o the b ra chiocephWe shall follow this vessel rs t ;
a l ic a r tery:

it divides a t on c e i n to fo u r t ru n ks ; the larg e m edial o n es are the ri gh t an d lef t

su b cl avian ar t eries the m uch s maller la t eral ones are the ri gh t and lef t ca r otid
,

ar teries Clean away conne c tive tiss u e from these vessels and follow their
.

c o u rses T he two su b clavians em bra c e the thyroid gland be tween t heir bases

and su pp ly sm all thyroid ar teries in to this gland E a ch s u b clavian nex t gives .

Off bra n ches to the ven tral S ide of the ne ck and to the tra chea Of which the chief ,

one is the ven tr al cervical ar tery a vessel arisi n g from the s u b cl avian abou t on e
,

half in ch beyo n d the thyroid gland and bran chi n g p rofu sely in to the esop hagu s ,

tra chea m u s c les of the ne ck an d thy m u s gland

, ,
The thymu s gland is a yellowish .

mass la teral to the ven tral c ervi cal ar tery and re c eivi n g bra n ches fro m it The .

su b clavian ar tery now nam ed the axill ary tu rn s la terally and p asses to the i n ner
, ,

su rfa c e Ofthe p e ctoral girdle where a larg e bran ch aris es an d bran ches ex tensively
,

in to the p e ctoral an d sho u lder mu s cles The axillary t hen tu rn s abru p tly pos teri
.

orly an d abo u t an in ch beyon d the t u rn gives O the sm all dor s al cervical in to the
n eck,
the r s t in ter cos ta l la terally an d the ver teb r al c a u dally The rs t in ter .
,
23 6 L AB ORA OR T Y MANUAL F OR VE R TE B RATE ANATOMY

c os tal r u n s la t erally an d t h en tu rn s pos t eriorly jo ihin g the margin o cos tal art ery
,
-

whi ch co u rs es alon g the cu rve of the c arap a ce The ver t ebral p ass es bac kward .

alon g the ver t ebral c ol u mn dorsal to the ribs alon gside the ver tebral vein an d
g ives O ff a t the s u t u res O f t he c os t al
p la t es t h e in te rco s ta l ar t eries w hi c h run ,

la terally in to the m argi n o cos t al ar tery At the poi n t where the rs t in t er cos tal
-
.

an d ver t ebral aris e the a xillary b en ds shar ly la t erally an d as the b rac ar ery
, p h ial t
p asses i n t o t h e for e limb alo n gside the bra chial vein .

E a c h c aro tid ar tery p asses forward along the ven tral side of the ne ck soon ,

c rossi n g dorsal to the s u b clavian and then c o min g to lie m edial to the su b c lavian .

In specime n s in whi ch the ne ck is drawn in to the S hell the c aro tids u su ally m ake
l do p s in the ne ck region As the caro tid ar tery p asses the th ymu s glan d it gives
.

bra n c hes in to the gland I t then p ro ceeds wi thou t bran ching the en tire leng th
.
, ,

O f the n e ck in con ta c t wi th the in ternal j u gu lar vein an d the va gu s nerve and ,

en ters the sku ll b y a foram en in fron t Of the au di tory re gion .

Draw the b rachio cephc ar tery and its bra n ches .

b ) The pu l mon ary ar ter ies : The p u lrn on ary ar t ery is the one far thes t t o the
lef t O f the three ar terial tru n ks whi ch sp rin g from the ven tri c le I t divides .

imm edia tely i n to r ight and l eft pu lm on ary ar teries TO see this division lif t the .

p u l m o n ary t ru nk and look on it s dorsal side Tra c e th e lef

. t p u l m o n ary r s t I t .

p ro c eeds la t erally p os t erior t o th e lef t aor ta t o w hi c h it is m ore or less bo u n d

by c o n ne c tive tissu e formin g the ar ter ial ligamen t or l igamen t of B otal l u s ( The
,
.

signi can c e of thi s l igam en t will be exp lained la ter ) The p u lmonary p ro ceeds .

dire c tl y to the lef t lu ng in co mp an y wi th the lef t bro n chu s an d lef t pu lm onary

vein Tra ce the ri gh t p u lm o n ary in the sam e way I t is bo u nd to the ri gh t
. .

aor ta by the righ t ar terial li gam en t .

Add these to the p re c edi n g drawin g .

c) The r ight an d l eft a or ta e: Tra c e bo th O f these ar t eries away fro m the hear t .

E a ch m akes a cu rve as it leaves the hear t an d tu rns p os teriorly p assi n g dorsal ,

to the p re caval vein the bro n chi and the p u l m o n ary vessels and disapp earin g
, , ,

dorsal to the lobes of the liver Vessels already s tu died m ay be cu t to follow

.

the aor tae p os teriorly . Tra ce the lef t aor ta rs t G rasp the s tom a ch and lef t .

lobe of the liver an d p ress them to the righ t sep ara ti n g the c ardia c en d O f the ,

s to m a ch from the lu n g The lef t aor ta will be fo u n d p assi n g to the lef t Of the
.

esop hagu s and dorsal to the s tom a ch I t gives o ff sim u l tan eo u sly three large
.

bran ches On e O f these is the gas tr ic ar tery whi ch p asses to the s tom a ch in the
.

cardia c re gio n and follows the cu rve of the s t o m a c h alo n g the le n g th of this organ .

Af ter a shor t dis ta n c e it forks in t o an ter ior an d pos ter ior g as tri c ar teries whi ch
s u pp ly the lesser and grea ter cu rva tures Of the s tom a ch resp e c tively Ano ther .

bran ch fro m the lef t aor ta is the coel iac ar tery I t soon forks i n to an ter ior .

and pos ter ior pan cr eatico du oden al ar teries The a n terior p an crea ti c o d u oden al
-
.
-

ar tery p ass es to the lef t en d of the pa n creas gives Off there bra n ches in t o the ,

p y lori c en d of the s to m a ch and to the liver then tu r n s to the ri gh t and ru ns ,

23 8 L AB ORA OR T Y MANUAL FOR VE RTE B R ATE ANATOMY

inj u red in the di sse c tio n Of the renal p or tal sys tem try the o ther side ) I t ru ns , .

la terally to the p oin t where the renal p or tal vein en ters the p leu rop eri ton eal
cavi ty At this p oin t it divides
. The an terior bran ch con tinu es to the carap a ce
.

an d ru n s forward along the cu rve of the carap a c e s u p p lyin g the fa t bodies and ,

be co min g con tinu ou s wi th the m argi n o cos tal ar tery des cribed above The
-
.

p os t erior bran c h tu rns and p asses m edially p arallel to t he ven t ral abdo m i n al
vein I t s u pp lies the base Of the leg and the p elvi c m u s c les and termina tes on
.

the ven tral s u rfa c e O f the p elvis .

Nex t follow the co mmon ilia c ar tery of the sam e side I t divides a t o n c e .

before it has em erged fro m above the kidney in to an in ter n al il iac and an exter n al
il iac ar tery The ex ternal ilia c forks af ter a shor t di s ta n ce The m edial an d larger
. .

bran ch s upp lies the mu s cles of the p elvis and as the femoral ar tery en ters the
thigh . The s maller and la teral b ran ch p asses deep dorsally to the p oin t where
the iliu m is ar ti cu la ted to the sa c ral ribs ; here it p asses dorsal to a nerve an d
tu rns ven trally as the s ciatic ar tery in t o the hi nd l eg ru nni n g alo n g the m edial ,

su rfa ce Of the ili u m The in ternal il ia c is bes t followed by rep la cin g the kid n ey
.

agains t the dorsal wall p u lli n g the large in tes tine ba ckward and lo ca ti n g the
,

p oin t O f ori gin of th e in t ernal i l ia c s fro m t h e c o mm on ilia c The c hi ef bra n c

. h
of the i n tern al ilia c is the hemorr hoidal ar tery whi ch p asses forward alo n g the
side Of the large in tes ti n e ; in addi tion there are branches to the bladder the ,

rep rodu c tive organ s and the p elvi c region in general

, .

Draw the dorsal aor ta and its bra n ches .

8 The s tru ctu re Of the he art

.
S ep ara t e the hear t of the tu r tle b y cu t ting
a cross the grea t vessels and remove it from the body The p os terior cham ber .

of the hear t is the s in u s ven os u s whi ch re ceives the fo u r g rea t sys temi c veins .

Clea n ou t the blood fro m the sin u s I t is a thin walled cha m ber a t ta ched to the
.
-

ri gh t au ri cle in to whi ch it o p ens by the s in au r icu l ar open in g gu arded by a p air

,
-

O f thin valves Op e n ea ch a u ri cle by m akin g a sli t in the m argin a n d washi n g

.

ou t the blood c lo ts The walls of the au ri cle sare som ewha t sp on gy L ook in to
. .

the lef t a u ri cle an d n o te the thin in ter a u r icu l ar s eptu m whi ch c o mp le tely sep ara tes
the cavi ty Of the lef t a u ri c le fro m t ha t of the ri g h t o n e F ind the o p en i n g of the
.

p u l m o n ary veins in t o t h e dorsal wall of th e lef t a u ri c le n ear the se p tu m F i nd .

on ea ch side the large au r icu l o ven tr icu l ar open in g be tween ea ch au ri cle an d the
-

ven tri cle M ake a cu t all Of the way arou nd the margi n of the ven tri cle so as
.

to m ake dorsal and ven tral a p s of the ven tri cle Sp read ap ar t the two ap s
.

cau tio u sly e x te n din g yo u r c u t inward u n til the two ap s are a t t a c hed o n ly alon g

the base of the ven tri cle No te the ex ceedingly thi ck wa l ls of the ven tri cle and
.

the mu s cu lar c olu mn s p roje c ting in to the in terior The ca vi ty of the ven tri c le
.

is a broad b u t a ttened cavi ty u s u al ly con tainin g a sp ongy n e twork whi ch may

be clean ed ou t Sp reading the two ap s widely no te in the base of the ven t ri cle
.
,

a ba n d p assi n g a c ross fro m one side to the o ther On ea ch side of this ba n d is

.

an a u ri cu lo ve n t ri cu lar o p e n i n g
-
The ban d is a co n ti n u a tion Ofthe i n tera u ri cu lar
.
 COM P ARATI VE ANATOMY OF THE CI R CULATOR Y S Y STE M 239

s ep tum and form s a fold or valve on ea ch S ide which p ar tially o cc lu des the ,

au ri cu lo ven tri cu lar op ening The ri gh t valve con tinu es ven trally i n to a rid ge
-
.

whi ch is on the ven tral ap of the sp e ci m en This rid ge is the in comp le te .

in terven tr icu l ar s eptu m On bri n g in g the two ap s of the sp e cimen toge ther
.
,

it wil l be seen tha t the in terven tri cu lar sep tu m was conne c ted wi th the mu s cu lar
wall Of the dorsal ap and tha t a sp a ce is lef t dorsal to the sep tu m by whi ch the
righ t and lef t ven t ri cles commu ni ca te wi th ea ch o ther The ri gh t ven tri cle to .

the ri gh t O f the sep tu m is very s m all whi le the lef t ven tri c le is mu ch larg er and
,

co m mu ni ca t es wi th the c avi ty of bo th a u ri cles owin g to the in co mp le te chara c ter

Of the in terven tri cu lar sep tu m Sp read the ap s Of the sp e cimen a gain and
.

p ass a p robe ven t ral to the in t erven t ri c u lar se p tu m The p robe e m er g es in .

the p u l m onary ar tery P robe in to the o ther ar terial tr u n ks and nd their Op en

.

in gs in to the ven tri cle The op ening Of the lef t aor ta is to the ri gh t Of the in ter
.

ven tri cu lar sep tu m in to the small righ t ven tri cle whi le tha t of the righ t aor ta
, ,

is to the lef t O f the sep tum in to the lef t ven tri c le ; however owin g to the gap
, ,

dorsal to the sep tu m the lef t aor ta can also Ob tain blood from the lef t ven t ri cle
, .

B y sli t ting o p e n the ar terial tr u n ks nd the li ttle p o cke t like s emil u n ar valves -

which gu ard their exi ts from the ven tri cle They rep resen t the rem ai n s of the .

c on u s ar t erios u s .

W e may mp t to exp l ain the

n ow a tte cou rse Of the r
c i cu l a tion hrough the tu rtl e s hea t
t

r .

The ma tter is som ewhat co mplica ted an d fu r ther detail s will b e foun d in P an d H p ag e 3 59 , .

W e have n o ted tha t a ll bl ood retu rn s to the S in u s ven osu s an d that this in tu rn
O f the ven ou s

c onn ec ts with the right au ricl e which p ass es it on in to the right s ide o f the ven tricl e Al tho gh . u

this is imp er f ec tly s ep arated fro m the l ef t s ide o f the ven tr icl e the v en ou s bl oo d is well r etain ed ,

in the right s ide o win g to the s p o gy n atu re o f the ven t icu l ar wall s
n Mean time the two r .
,

pu hn on ary vein s have retu rn ed the bl ood from the l un gs to the l eft au ricl e S in c e the fun c tion .

o f the l un gs is to a era te the bl ood this bl ood is a terial ,

From the s tru c tu re an d r el ation s o f
r .

the hear t the right au ricl e al ways c o tain s ven ou s bl o od an d the l ef t au ricl e a rterial bl ood
,
n ,
.

T he l eft au ricl e p ass es the ar terial bl o od in to the l ef t Side Of the ven tricl e Ther e is s ome .

s light m ix t r e O f ven ou s an d ar terial bl ood in the ven tricl e

u AS the ven t icl e con tr ac ts bo th . r

k in ds of b l ood are moved toward the arterial trunks W e have n o t ed that the pul mo ary . n

a tery s p i g s fr om the s mall right ven tricl e an d that the op en in g in to this ar tery is to the
r r n

righ t O f an d s omewha t con ceal ed by the i t erven tricu l ar s ep tu m When the ve t icl e con
n . n r

t ac ts the pres su re p rac ticall y cl oses the sep tum so that mos t o f the ve ou s bl ood p a s es ou t
r ,
n s

i to the pul mon ary ar tery

n Simu l ta eou s ly the n ea rly p u e a te ial bl ood in the l eft ve
. n ,
r r r n

tricl e p asses in to the b as e O f the rig h t aor ta s i ce that is co ec ted with the l ef t ven tricl e
,
n nn

an d s in ce the comm u nica tion with the l ef t ao r ta is cl o s ed tem p or a il y by the i terv e tricul ar r n n

s ep tu m . Towa d the e d Of the co t action the remai i g bl oo d in both ve t icl es p as ses

r n n r n n n r

in to the l ef t ao ta as the dimi ished p r es s u re ag ain op e s u p the g ap in the s p t m

r ,
n It n e u .

thus ha pp e s tha t the brachiocepha lic ar tery pass i g to the an terio r p ar t o f the bo d y c o
n n n

tain s n early pu r e a terial bl oo d whil e the l ef t ao r ta co ta in s mixed bl oo d

r ,
On accou t O f the
n . n

fun ction o f right a d l eft ao tae the dors al aor ta al so carries mi ed bl ood I t is u n ivers all y
n r ,
x .

tru e amo g ve t eb ra tes tha t the a r an g eme t o f the ci cul a to y sys tem is s uch tha t the
n r r n r r

pu res t bl oo d is received by the head ; an d this is n o doubt du e to the g r eater oxygen require
men ts o f the n ervou s p a ts O f the head r .
2 4o LAB ORATORY MANUAL FOR VE RTE B RATE ANATOMY

We h at wh ereas in s h es th ere is a s in g le cih

n o w see t cu l a tion th r ou gh the h ea rt w h ich ,

carries onl y ven ou s b l oo d th er e is in the tu r tl e a dou bl e cir cu l a tion o n e h al f o f the h ea r t

, ,

( al ways the l ef t) con veyin g ar terial bl ood an d on e h al f ( al ways the rig h t) ven ou s bl ood Th is .

ch an g e is du e to the d evel opmen t o f the l u n g me th o d o f b r ea th in g w ith the con s equ en t s h if t

in g o f the s in u s ven o su s to the righ t s id e an d the d ivis ion o f the au ricl e in to two s ep ar a te
chamb ers The b l o od co u l d n o t go th rou g h the l u n g s an d th en to the b od y b eca u se all o f the
.

force of the h ear t b ea t woul d b e los t in the p ass ag e o f the b lood th rou gh the capil l aries o f the
lun g s an d the circu l ation woul d s tagn a te in the lun g s The aerated blood r etu rn s to the h ea r t
, .

to take advan tag e o f the d rivin g ac tion o f the h ea r t Thu s the d ou bl e circu l a tion aro se
.
, .

W e n o te h ow ever th a t the meth od emp l oy ed by the tu r tl e ( an d o th er r ep til es) is imp erf ec t

, ,

in th at the a r terial an d the ven ou s b l oo d are mix ed in the h ear t A l ittl e con s id era tion w ill
.

s h o w th at this s itu a tion can n o t b e r emed ied mer el y b e co mp l etin g the in t erven tricu l ar s ep tu m

(wh ich has in d eed h app en ed in the cro cod il es an d a ll ig ato rs) b ecau s e the l eft aor ta op en s in to
the righ t ven tr icl e an d wo u l d s till co n tin u e to r ec eive ven ou s b l ood The d if cul ty is in real ity
.

du e to the p res en ce o f the th ree ar terial tru n ks fo rmed by the s pl ittin g o f the ven tral ao r t a .

W e can s ee th at if the ven tral ao r ta wou ld s p lit in to b u t two trun ks on e o f wh ich ( pul mo n ary) ,

is con n ec ted w ith the rig h t s id e o f the h ear t an d the o th er ( aor ta) w ith the l ef t s id e an d if ,

fu r th er the in terven tricu l ar s ep tu m woul d b e compl eted the dif cu l ty wou l d b e overco me
, ,

an d n o ven ou s b l ood cou l d g et in to the a r terial s y s t em Th is is p recis ely wh a t has h app en ed

.

in b ir d s an d mammal s in w h ich the d ou b l e cir cu l a tion is co mpl ete an d p er fec t I t fo ll ow s

.

from this th a t b ir d s an d mamm al s cou l d n o t h ave evolved fro m an y l ivin g g ro u p s o f rep til es
b u t mu s t h ave aris en fa r b ack in the r ep til ian lin e b efo re the s p l ittin g o f the ven tral ao rta
occu rr ed .

9 Compa i on of the cir cu l atory s ys tem of the tu r tl e with pr ec e din g f orm

. r s s T he.

s tu d en t can h a r d l y fa il to h ave n o ted marked d if feren ces b etw een the circu l a to ry s ys tem o f the
tu r tl e an d o f the el as mo b r a n ch s an d Amphibia Th ese ch an g es a re assoc ia ted with the ado p
.
~

tion o f the air b rea th in g h a b it

-
I n the a r terial s ys tem p ro fou n d ch an g es h ave o ccu rr ed in the
.

a or tic arch es ( F ig 58 p .
, . We h ave l earn ed tha t in ver teb rate emb ry os th er e are s ix aor tic
arch es ; in el as mo b ran ch s the rs t is mis s in g the s econ d in co mpl ete an d fo u r c o mp l ete o n es
, ,

p er s is t ; in N e ct u r u s t h e r s t seco n d a,
n d f t,
h h a v e v a n is h e d o n,l y th e th i r d four th ,
a n d s i
,x th
r emain in g I n the tu r tl e we n o te th a t b u t o n e p air o f ao r tic a r ch es has p ers is ted ; th es e u n ite
.

dorsal ly to form the d o rs al ao r ta ( Fig 5SE ) Th is su rvivin g pair o far ch es r ep resen ts the fou r th
. .

p air an d is u su a l ly r efer red to as the aor tic arch s in ce th er e is b u t o n e

, ,
The thi rd p air o f arch es
.

is qu ite d is con n ected fro m the d o rs al ao r ta an d is rep res en ted by the b as es o f the ca ro tid s T he.

f th pair is ab s en t The b as es o f the s ix th p air p ers is t as the b as es o f the pu lmon ary arteries
.

( Fig 5SE ) T he con n ection o f the s ix th arch es w ith the d ors al aor ta is p res en t in ver teb ra te
. .

emb ry os as the d u c t o f B o ta ll u s or ar terial d uc t b u t a f ter b ir th o r h at ch in g th is c l oses u p an d

,

d egen era tes in to a b an d o f con n ec tive tis su e the ar terial l igamen t n o ted in the d is s ec tio n
,

( g
F i . T h e b r a n c h e s o f th e d o rs a l a or t a ar e s im il a r t o t h ose o f th e a n im al s a l r e a d y c o n

s id er ed The s p littin g o f the ven tral ao r ta in to th ree trun ks has b een emp h as ized d u rin g the
.

d is s ec tion an d the r el a tion o f th is to the dou b l e circul a tio n th rou gh the h ear t expl ain ed ab ove
,
.

I n the ven ou s s y s tem l ikew ise ch an g es h ave o ccu rred

, ,
T he p recaval vein is the an terio r
.

ca rd in al vein o f lo w er fo rms ; its b as e con n ec tin g w ith the s in u s ven osu s is the common card in al

vein The ren al po r tal vein s are the pers is ten t pos terior p arts o f the pos terio r cardin al vein s
.

( F ig s .
55 p 2 0
, 5 a n
. d 57 p . T h e a n t er io r p a r t o f th e p os ter io r ca r d in al v e in i s m is s in g ( F ig .

57 )D a n d is r e p l ac e d f un c t io n a ll y by t h e v e r t e b r al v e in w h
,
ic h is f or m ed by a l o n g itu d in al an as

tomo s is b etw een the s eg men tal b r an ch es o f the em b ry o n ic po s terio r c ard in al I t is imp or ta n t to
.

n o te th a t the po s terior c ar d in al ( r en al po r tal ) ha s ex t en d ed its po s terior con n ec tio n s as in A m ,

h ib i W h i h l m b h th n a l po r tal vein co ll ec ts onl y fr om the tail in Amphib ia

p a . e r ea s n t e e a s o r an c e r e ,

an d re p til es it co ll ec ts fro m the hin d l imb a s w ell ( Fig . Th is is du e to a u n ion b etween

 242 LAB ORATORY MANUAL FOR VERT E B RATE A NAT O MY

V
. ein s e i g dir e c tly in to
op n n gh t au ri cle Th er e ar e th r ee o f the s e vein s the
the ri .
,

two p r eca vais an d the p os tcaval The p o s tcav al en te r s the righ t au ricle fr om
.

b ehin d em er gin g fr o m the liver The p r ecav al s co m e fr o m the a n terio r p a r t

,
.

o f the b o d y on e o n e ac h s ide an d cu r vin g to w ar d the he a r t a t the level o f the

, ,

a u r i c le s en t er the r igh t au r i c le The p u l mon ary vein s m ay b e n o ti ced op enin g

.

in to the lef t au ricle .

2 .The he patic portal sys te m T u rn to the p e ri to n e al c avi ty Cu t a cr o ss

. .

the f a l cif o r m ligam e n t o f the live r n e ar the gi zz ar d n o ti n g r s t the sm all vein p a ss ,

in g fr om the ven tr a l lig am en t o f the gi zz a r d in the fa l cifo r m lig am en t to the

liver The lob es of the liver m ay n o w b e tu rn ed fo rwar d Ru nn i n g alon g the
. .

dor s al su r fa c e o f the liver an d b r an c hin g i n to its su b s tan c e is the l a rge hep a tic
p or tal vein T he m ain p ar t of the vein en ter s the righ t l o b e of the liver cou r s in g
.
,

b e tw een the two b ile du cts The r em ain de r o f it lie s a l on g the d o rs a l su r fac e
.

o f the lef t l o b e o f the live r s en d in g b r an c he s i n to the live r an d a t the lef t r e ceive s

, ,

the l eft an d median gas tr ic vein s f r o m the m ar gi n a n d lef t s ide o f the giz z ar d a n d
fr om the pro ven tri cu lu s Fo ll ow p os teri or l y th a t p ar t o f the hep a ti c p or tal
.

w hi ch lies b e tw een the two b ile d u c ts I t is soon s een to b e form ed by the u n i on .

o f th r ee vein s a s u p er ior mes en ter ic a ga s tr odu oden al an d an infer ior mesen ter ic
, , , .

The su p er i o r m e s en ter i c co lle c ts fr om the gr e at e r p a r t o f the sm a l l in te s ti n e The .

gas tr o du o den a l r e ceive s the r ight gas tr ic vein fr om the r igh t side o f the gizz ar d ;
the p an crea tico du oden al vei n whic h r u n s a l o n g the d u o de n a l l oop c o lle c tin g f r o m
-
,

duo den u m an d p an cr eas ; an d the mesen teric vei n fr om the l as t l oop o f the small
in te s tin e T he in feri or me s en ter i c vein r u n s a lo n g the l arge i n te s tin e fr om which
.

it c o lle c ts m an y b r an c he s At its p os t e ri o r en d it tu rn s d o r sally a n d j oin s the

.

r e n a l p o r ta l s ys te m w he r e it will b e f o ll o w ed l a t e r
,
.

D raw the b r an che s o f the hep ati c p or ta l sys tem .

3 The sy s te mic ve in s
. As a l r ea d y s ta ted the s e con si s t o f two pr ecaval s
.
,

an d o n e p os tca val .

a ) The b ra n ches p recaval vein s : As b o th vei n s h ave iden ti ca l b ran ches

f
o the ,

o n l y o n e n eed b e f o ll o w ed F in d the vei n o n e a ch side l a ter al to the au ri cle an d

.

tr a c e e a ch i n to the r igh t au ri cle lif ti n g the hea r t The lef t p r e caval p asse s
,
.

a r ou n d the lef t a u r i c le to en te r the r igh t a u ri c le T he r igh t p r e cav al is mu ch .

sh o r te r a n d e n t e r s the r igh t au r i c le di r e c tl y .

F o ll ow o n e p r e ca va l fo r war d I t lie s j u s t p o s ter i o r to a l a r ge ar ter y an d is

.

the r e s ee n to b e f o r m ed by the u n i on o f th r ee l a r ge vei n s l a te r al l y the p ector al ,

vein s ligh tly an ter i or an d d o r sal to this the su b cl avian vein an d an t eri o r ly the
, ,

j g
u u l a r ve in E ac
. h o f t he s e ve in s s h ou ld b e f o ll o w ed T he p e c t o r a l ve in a t it s .

u ni on wi th the o the r s r e c eive s the in ter n al mamma ry vein as c e n din g f r o m the ,

in n e r su r fa c e o f the r i b s an d has a l s o a tr i b u ta r y f r om the s te r n u m an d c o r acoid

,
.

The m ain vei n is fo rm ed l a te r a ll y by the u ni o n o f two vein s em er gin g fr o m the

p e c t o r a l m u sc le s T he s e.m a y b e f o ll o w ed i n t o t h e m u sc le s f r o m w hi c h t he y a r e

s ee n to co lle c t m a n y b ran c he s The su b c l a vi an vein p a s s e s deep d o r sall y ven tra l

.
 OM PARATIVE ANATO MY OF
C THE C IR CULATORY SYSTE M 2 43

to a g rou p of n e r ve s ( b r achial plexu s) an d s om ewh a t con cealed by ar teries which

s h ou ld n o t b e i n j u r ed As the br achial vei n it e me r ges f rom the wi n g an d the n
.

r e c eive s a b r an c h f r o m the s h o u lde r m u sc le s . The j u gu lar vein p asses an te ri o r ly

on the d o r s a l side o f the l a r ge a r te rie s . On tra ci n g the j u gu l ar forwar d it will
b e fou n d to r e c eive the fo llo w i n g vei n s n am ed in o r der from the hear t fo rwar d :
,

o n the m edia l side so m e sm a ll a n d t he n a l ar ge b r an ch fr o m the ) (a t the p oi n t

\ fi g
o f e n tr a n c e o f the s e in to the j u g u l ar is si tu a ted a sma ll r eddi s h b o d y the cer vical ,

lymp h gl an d) ; o n the l a t era l side a vei n fro m the s hou lde r an d a t the s ame ,

level the ver tebr al vei n fr om the ver teb ral co lu mn ; medi all y an o ther b ran ch
W
, ,

from the c r a ll y a l a r ge vein f r o m a

p le x u s o f b l o o d ve s s el s in t h e s ki n o f
the n eg l
i m en s m a ll vei n s f ro m the e s op h ag us an d t
g g h ea . O n f r eei n g the

a n t e r i or en d o f the e sop h a gu s ( a l s o tr a chea 3 1d cu ttin g a cr oss it the j u gu l ar ,

vein can b e follow ed to the so f t palate wher e it j oin s its fello w o f the opp o si te
,

s ide . P os te r i or to thi s u n i on e a ch r e c eive s a p lexu s o f vei n s fr om the skin o f

the f ac e . On di ss e c tin g away the s o f t p a l a te fr om the an as tom osi s o f the two
j u gu l ar vei n s b ran che s from the sku ll will b e fou n d passi n g i n to the an as tomo si s
, .

D raw the b r an che s o f the p re ca val vei n as far as fou n d .

b ) The p os tcaval vein : R ai s e the ven tr i cle s o f the he ar t an d n o te on c e m o r e

the l a r ge p o s tca va l vei n e m e r gi n g f rom the live r an d en t e ri n g the righ t au r i cle
b e tw een the two p r e ca va l vei n s No te the l arge hep a tic vei n s whi ch it r e c eives
.

from the liver The lef t on e of the s e hep ati c vei n s r e c eive s the small vei n o f
.

the f a l cif o r m ligam e n t m e n ti on ed p r evi ous l y F oll o w the p os tc ava l vei n in to

.

the p e r i to n e a l c a vi ty tu r nin g a ll o f the vi sc e r a to the lef t

, . The p os tcaval will
b e p i cked up ag ain a t the p os te ri o r mar gi n of the righ t l o b e o f the live r in con tac t
wi th the d o r sa l b o d y w a ll . T he po s tcava l is he r e seen to b e formed by the u ni o n
o f two vei n s the il ia c vei n s
,
In m a le s the two o va l te s te s wi ll b e n o ted a t this
.

p o i n t o f j u n c ti o n I
. n fe m a le s t h e s i n gle o v a r y a n d o vid u c t w ill b e n o t ed t o t h e

lef t con cealin g the lef t ili a c vei n E ach ili ac ru n s a lon g the ven tral fac e o f a
,
.

th r ee l o b ed o r gan the kidn ey w hi ch is se t cl os e again s t the d o r s al b o d y w all

-
, ,
.

Fo ll o w the r igh t ili a c vei n . F r om b e tw een the r s t an d s e co n d lob es o f the

kidn ey it r e ceive s the l arge femor al vein emer gin g fr om the leg The femoral
,
.

vein r e c eives a sm a ll b ran ch from the b o d y wall P o s te ri or to the en tr an c e o f .

the fe m o r a l vei n the ili a c vei n co r r e sp o n d s to the r en a l p or tal vei n o f r ep tile s an d

Amp hib ia an d m ay b e so n am ed F r om b e tw een the s e con d an d thi r d l ob es o f
.

the k id n ey it r e ceive s the s cia tic vei n whic h a l so com e s f r om the thigh At the
.

p o s t e r i o r en d o f t he kid n e y s th e t w o r e n a l p or t a l vei n s h a ve a n a n as to m o s i s w i th
each o the r F r o m this an as tomo si s ri s es the infer ior mesen teric vein a l r ea dy
.
,

n o ted a s a b r an ch o f the hep a ti c p o r ta l s ys t em . I t r u n s in the m e sor e c tu m The

.

an a s to m os i s o f the r e n a l o ta l s l o e c eiv e s i th e m edi a n lin e a s m a ll ca u dal

p r a s r n

vein from the tail an d on eac h s ide an in tern al iliac vei n f rom the ro of of the
p elvi c r eg io n . T he lef t ili ac a n d r e n a l p o r t a l ve in s a r e th e sa m e a s t h e r igh t ,

ex cep t th at in the fem ale the lef t vei n s r e c eive gen ital vei n from the o vary an d s
 2 44 LAB ORATORY MANUAL FO R VERTE B RATE ANAT OMY

o viduc t Thes e may b e s een by tu rn i n g the o vidu ct to the righ t I n thei r co u r s e

. .

o ve r t he kid n e y s the r en a l p o rt a l vein s give o b r an c he s i n t o t he kidn e y a s

in low e r f o rm s . T he r e is p ro b ab l y s om e p ort a l c i r cu l a t i on in the kidn eys b u t ,

mo s t o f the b l oo d f r om the r e n a l p o r t a l s p as s e s di r e c t ly i n to the ili ac vein s .

T he ilia c vein r e c eive s r en al vei n s f rom the kidn ey of w hi ch the r e is o n e c hief

r e n a l vein whi c h ru n s a lon g the m edi a l s ide o f the kid n e y b u t is s o imb edded
,

in the kidn e y s u b s ta n c e a s t o b e di f cu l t to iden t if y B e tw een the two r e n a l

.

p o r t a l a n d ili ac vei n s ru n s t he d or s a l a o r ta .

Tu rn the an ima l d o r s a l s ide u p an d r emo ve the s kin o ve r the thigh B y .

s ep a ra t in g t he m u s c le s p i c k u p the fem o r a l a n d s c i a t i c vein s an d t r ac e t hem in t o

the leg . T he s c i at i c vein accomp an ie s the l a rge s c i a t i c n e rve an d s o on tu rn s

forward to ru n p ar a llel to the fem o ra l vein B o th r ein s a r e accomp an ied by
.

a rt e rie s o f t he s am e n am e .

D raw the b ran che s o fthe p ost cava l vein an d the r en al p orta l s yst em .

4. T h e p u l m on a r y ve in s
. T he p u l m on a r y vei n s e m e r ge o n e ac h s ide f r om

the lun g an d p as s t ow a r d the he a rt immedi at el y p os t e ri o r t o the p r e cav a l vein s .

T he r e is u su a ll y on e pu lm on ary vei n f rom each l un g b u t t he r e m ay b e two N o t e .

the b r an che s c o lle ct ed by e ac h vein f rom the lun g T he vein s p as s to the d ors a l
.

s ide o f the b as e s o f the p r ecav a l s an d en t e r t he lef t au ri c le T hei r en t ran c e in to

.

the au ri c le is b e s t s een l at e r w he n the he a rt is dis s e ct ed .

5 The arterial s ys t e m
. .
I t has alr e ad y b ee n n o ted th at the g r e at ar te r ie s
sp ri n g di r e ct l y f r om t he ven t ri c le . T he y a r e s i tu at ed b e tw een the two au ri c le s .

S ep arat e t hei r b as e s f r om the au ri c le s I t w ill b e t hen b e foun d t h a t t he r e a r e

.

two a r t e ri a l t ru n k s . T he l arge r m edi a ll y l o ca t ed o n e is the aor ta

, T he sm a lle r
.

o n e p a s s i n g t o the lef t an d d o rs al t o the aor ta is the p u l mon ar y ar t e r y

, ,
T he .

ven t ra l aort a in b i r d s is sp li t in to the s e two ve s s e ls .

a) The an ter ior b r an ches of th e a or ta : F o ll ow the a o r ta aw a y f rom the he a r t .

T he ao rta immedi at el y give s ri s e in the m edi an l in e t o two l arg e art erie s the ,

br achiocep halic (in n omin a te) a rt e rie s . T he ao rt a then tu rn s t o the righ t an d

di s app ear s d or s a ll y I t W ill b e f o llow ed at a lat e r t im e I den tify the b r an che s
. .

of the b ra c hi o c ep h a li c a rt e r ie s ; a s b ot h h a ve ide n ti ca l b r a n c he s f o ll o w on l y

on e . E a ch p r o c eed s l at e ra ll y an d s ligh t l y an t e ri o r l y a n d f o rk s in t o two b ran c he s ,

an an t e r i o r common car otid a rt e r y an d a la t e ra l s u b cl a vian a rt e r y T he su b .

c l avian a rt e r y s o on give s r i s e t o a n um b e r o f b r an c he s : the sma ll in ter n a l

mammary art e ry p a s s i n g p o s t e ri o r l y a lon g the i nn e r su r f ac e o f the r i b s the two ,

pector al art e rie s to the p e c t o r a l mu s cle s a lo n g w i th the vein s o f the s am e n am e ,

an d t he axill a r y a rt e r y t o t he w i n g . T he ax i l lary a rt e ry ru n s an t e r i or ly a n d
a f t e r givin g offa b r a n ch i n t o the s h ou lde r e n t e r s t he w i n g a s the b r a chial a r t e r y .

T he two common car otid a r t e rie s p a s s fo rw ar d an d at the level o f the c e r vi cal

lym p h glan d e ac h give s ri s e to a ver tebr al a rt ery w hi ch p a s s e s d o r s a lly i n t o
,

the ve rt e b r ar t e r i a l can a l o f the ve rt e b r a l co lum n T he comm on c a r o tid a r t e rie s

.

then app r oa ch t he m edi an lin e a n d p en e t r at e the m u s c le s o n t he ve n t r a l s u r

2 46 L AB ORATORY MANUAL FOR VERTEB RATE ANATOM Y

e fr om the d o r s a l aor ta ; it su pp lies the m iddl e an d p o s terio r lob e s of the

n xt

kid n ey then p ro ceed s to the late ra l b o dy wa ll an d as the femoral ar t ery su pp lies

, ,

the leg . I t m ay b e f o ll o wed by tu rn i n g the a nim a l d o r s a l side up an d l oo kin g

b e tw een the m u s cle s o f the thigh alon g the co u r s e o f the fem o r a l vei n p r eviou sly
iden tied The fem or a l ar te r y ac com p an ie s the l ar ge s ci a ti c n erve an d b ran ches
.

i n to the leg m u scle s R e tu rnin g to the p er i ton e al cavi ty tr ac e the d orsa l ao r ta

.
,

fu r the r As it p ass e s b e tw een the kidn eys it give s o l u mb ar ar te r ie s in to the

.

d o rsal b o dy wall At the p os te r i or en d o f the kidn eys it fork s At the p oi n t

. .

o f f o r ki n g a r i s e the infer ior mes en ter ic a r t e r y whi ch run s an te ri o r l y in the

,

m so r e c tu m an d an as to m o s e s wi th a b r an c h o f the su p e r i or m e s en te r i c a r ter y
e ,

a n d the ca u dal a r te r y w hi ch p r o c eed s s tr aigh t p os te r i o r l y in the m edi an li n e to

,

the ta il T he two fo rk s o f the d or s al a o r ta a r e n am ed the in tern al iliac ar ter ies

. .

They p as s pos ter i o r l y al on g the r oo f o f the p elvi c r egi on The lef t o n e give s .

of f b ran che s in to the o vidu c t .

D raw the b ran ches o f the d o rs al aor ta .

6 The s tru ctu r e of the he art

.
Fr ee the he ar t by cu ttin g ac ro s s the g rea t
ve ss el s No te th a t al l vei n s en ter the app ar en tl y an teri o r en d o f the hear t hav
.
,

in g s hif ted forw ar d o u t o f the tr an s ve rs e s ep tu m As a l r e a d y s ta t ed ther e is

.
,

n ei the r s in u s ve n o su s n o r co n u s a r te r i o su s in the b i r d s he a r t O n l y two o f the

.

o r igi n a l fo u r c h am b e r s o f the he a r t h a ve p er s i s t ed n a m el y a u ri c le a n d ven tri c le

, , ,

b u t e a ch o f the s e is su b divided i n to two comp le tel y s ep ar a t e h a lve s the righ t ,

a n d lef t a u ri c le s a n d ve n t r i cle s T he au ri cle s a r e sm all thi n wa lled ch amb er s

.
-

a n te r i o r to the ve n tr i c le s I n the wa l l s o f the righ t a u r i cle iden tif y the op en i n gs

.

o f the s ys t emi c vei n s a n d in the lef t au r i c le o f the p u l m on a r y vei n s S li t op en .

the righ t au ri c le an d l o oki n g wi thi n n o te the thi n in ter au r icu l a r s ep tu m s ep a r a ti n g

it fr om the lef t au ri cle A fo ld ex ten d s fr om thi s s ep tu m to the en tran ce o f the
.

p o s tca v a l vei n p a r t l y co n c e a l in g th e e n t r an c e N o t e t h e dee .

p c lef t t he a u r ic u l o ,

ven tr icu l ar op en i n g th rou gh w h ,

i c h the r igh t au r i cle Op en s i n to the righ t ven tr i cle .

S im il a r ly op en the lef t a u ri c le an d n d the lef t au ri cu l o ven t r i cu la r op en i n g -

.

Cu t a c r o ss t he ap ex o f the ve n t ri c le s O b s e rve the c r e s c en t i c f o rm an d r el at ively

.

t h in w a ll s o f the r igh t ve n t ri c le an d c i r cu l a r s e ct i on a n d e n o rm ou s l y t h ic ken ed

w a ll o f the lef t ve n tric le Wh a t app e a r s to b e the in t e rn a l w a ll o f t he ven t r i c le s
.

is the in terven tr icu l ar s ep tu m com p le t ely s ep ar at in g the c avi t ie s o f t he two v en

,

t ric l es . Op en the righ t ven t ri c le by a s li t ex t en din g f r om the p r evi ou s ly cu t

ap ex to t he b a s e N o t e t he s in gle val ve whi c h gu ar d s the righ t a u ri cu lo ven t r i cu lar
.
-

op e n in g ; it is a m u s c u l a r b an d ex t en d in g f rom t he ven t ri cu l a r w a l l t o the

au ri cu l o ve n t ri cu l a r op e n i n g
-
I n the lef t s ide o f the an t e ri o r en d o f the righ t
.

ven t ri c le n d the op en i n g o f the pu lm o n a ry a r t e ry o r p rob e in t o t he b as e o f ,

the p u lm o n ary a rt e ry an d n o t e em e rge n c e o f the p r o b e in t o t he r igh t ven t ri c le .

At the b as e o fthe pu lm on ar y a rt e ry a r e th r ee p o cke t like s emil u n a r v a lve s -

Cu t .

in t o the left ven t ri c le an d n o t e the two t hin memb r an ou s va lve s whi ch gu ar d

the au r i cu lo ven t ri cu la r op en in g
-
T hey a r e ca lled the mitr al va lve E a ch is
. .
 C OM PARATIVE ANATO MY OF THE CIR CULATORY SYS T E M 24 7

a t ta c hed by deli cat e co r d s the chor dae ten din ae to the wa ll o f the ven t ri c le
, , .

T he w a ll o f the ven t ri c le has s eve r a l mu s cu l ar ridge s w hi c h p ro je ct i n to the

ca vi t y ; t he y a r e ca lled t he col u mn a e car n ac F in d to the m edi a l s ide o f the .
,

m i t r a l v a lve t he op en in g o f the lef t ve n t r i c le i n t o the ao r t a

,
P r o b e in t o thi s .

an d s a ti s f y yo u r s elf t h a t it le a d s i n t o t he a o r ta N ot e the th r ee semil u n ar va lve s .

a t t he b egi n n i n g o f t he aor t a .

T he r em o va l o f the he ar t p erm it s the t ra c i n g o f the e s op h ag us in t o the p ro

ven t ri c u lu s a n d o f t he b r on chi i n t o t he lu n g s T he f orm an d ex t en t o fthe lu n gs .

ca n a l s o b e O b s e r ved t o a dv a n t a ge a t t hi s t im e .

7 . The l
cir cu ation g
thr o u h the h ear t an d the c o m par is o n of the cir cu ator l y s y s te m o f
b ir d an d r e pti e . l In
b ird s the hear t is comp l etely d iv ided in to righ t an d lef t au ricl es an d
ven tricl es The ven ou s bl ood en ters the rig h t au ricl e fro m the s ys temic vein s p asses in to the
.
,

r ig h t ven tr icl e an d ou t in to the p u l mon ary ar te r ies w h ich con vey it to the l un g s
,
Af ter .

a er a tion in the l u n g s the b l oo d r etu rn s by way o f the pu l mon ary vein s t o the l ef t au ricl e fr o m ,

wh ich it ows in to the l eft ven tricl e an d ou t o f the ao r ta Thu s the righ t s id e o f the h ear t .
,

co n tain s onl y ven ou s b l ood an d the l ef t s id e on l y a r ter ial b l oo d Th ere is a perfect dou bl e .

cir cu l ation b o th kin d s of b lood owin g s imu l tan eou s ly th rou gh the h ear t the two s treams
, ,

co mp l etel y s ep ar a t ed fr om each o th er S in ce the ao r ta is con n ec ted on ly with l eft ven tricl e

.
,

the ar terial s y s tem r eceives p u r e ar ter ial b l o od T he p er fec tion o f the d ou b l e cir cu l a tion is
.

ach ieved by the s p l itti g o f the ven n t r al ao tar in to two t ru n

ks ao a a d p l o ary w h ich
r t n u m n
a r e con n ec ted w ith the l ef t an d rig h t s id es o f the h ea r t res p ec tivel y T he s in u s ven osu s is
, .

app ar en tl y ab s en t b u t in r eality is r ed u ced an d in co rp or ated in to the r ig h t au r icl e

,
T he .

co n u s ar terio su s is r ep r es en ted by the s emilu n ar val ves o f the ao r ta an d pu l mon ary ar ter y .

The aor tic ar ch es ar e fu r th er mod i ed fro m the con d itio n s een in r ep til es As in the .

tu r tl e the b as es o f the co mm o n car o tid ar ter ies r ep r es en t the t h ir d aor tic arc h es
,
The u n ion .

o f the s u b cl avian s w ith th ese is seco n d ary T he arch o f the ao r ta is the r ig h t fo u r th aor tic
.

ar ch ,
the l ef t fou r th arch h avin g va n is h ed d u r in g embr y on ic d evel op men t Th er e is con .

s equ en tl y in b ir d s n o c o mp l e te ao r tic arc h as in the p r e ced in g fo r ms b u t on l y s o to s p eak , ,

on e h al f an ar ch the p er s is ten t h al f b ein g the rig h t o n e

-
,
The p u l mon ary a r teries r ep resen t
.

the s ix th aor tic a r ch sep ar a ted a s in rep til es fro m the a o r ta ( Fig 58 F p
,
.
, .

The ven ou s s y s tem is r ep t il ian in ch arac ter T he two p recaval vein s a re s imil ar to th o s e
.

o f the tur tl e an d ar e h om o l o g ou s w ith the an ter io r ca r d in al vein s o f l o w er fo r ms ( the in tern al

j u gul ar b r an ch b ein g the o rig in al an t erior car din al ) T he b ases of the p r ecavals en terin g the.

s in u s ven osu s ar e the c o mm o n car d in al vein s T he p o s ter io r car d in al vein s a r e as in the

.

tu r tl e r ep r esen t ed onl y by th eir p os t er ior p or tio n s w h ich a r e n amed in the a d ul t the r en al

,

p o r ta l v e in s T h. es e ve in s h a v.
e a s in re,p t il es a n d A m p h ib ia a b s o r b e d th e v e,
in s o f t he l e g s

an d tail . I n b ir d s it is very in ter es tin g to n o te the u n ion w h ich is in p rog r es s b etw een the
r en al po r tal s y s tem ( p os ter io r ca r d in al s ) an d the pos tcaval vein As we s h all see th is u n io n
.
,

is comp l ete in mamm al s I n b ir d s the r en al p o r tal s y s tem is p r ob ab ly to a s l ig h t d eg ree

.

fu n ction al as a por tal sys tem th rou gh the kidn eys bl oo d p as sin g from the ren al p o r tal vein ,

in to the kid n eys an d re co ll ec tin g in to the r en al vein s trib u ta ry to the p o s tcaval vein M os t
- .

o f the b l oo d h ow ever p as s es dir ec tl y fro m the ren al p o r tal vein s in to the p o s tcaval
, ,
.

T he o rig in o f the p os tcaval vein in b ir d s is the same as th a t g iven fo r r ep til es Th at .

p ar t o f it w h ic h is s itu a te d b e t w ee n t h e ki d n e y s i s f o r m e d o f th e s u b ca r d in a l v e in s c h ie y th e ,

r igh t on e T he p ar t th rou g h an d an terio r to the l iver co mes from the vitellin e vein s
. The .

middl e reg io n o f the vein is a n ew fo r m a tion T he p o s tcaval is s een to b e u su rp in g the r en al

.

p o r t al s y s t e m a n d th u s ex te n d in g i t se l f p o s t er io r l y T h is p r.oc es s is c o m p l e t ed in m a mm a l s .
 2 48 LAB ORATORY MAN UAL FOR VERTEB RATE ANATOMY
The h epa tic ys tem is similar to th at in the fohns already discuss ed The ren al
por tal s .

po r tal s y s tem is id en tical w ith tha t o f r ep til es The in ferior mes en teric vein w hich con n ec ts
.

the two po r tal sy s tems is p r o b ab l y h omol og ou s w ith the ven tral a bd o min al vein s o f rep til es .

I t is h ow ever o f d ecr eas ed impo rtan ce as a ch ann el b etw een the two po r tal sys tems owin g
, , ,

to the j un c tion o f the ren al portal s y s tem w ith the po s tca val vein .

F . THE CI R CUL AT OR Y SYS TE M OF THE MAMMAL

The sp e c im en s h ou ld h a ve b een in je ct ed in the art e ri a l s yst em .

I The c hamb e r s of the he art

.
T he he ar t is r el ativel y l arge an d co mp a c t ,

the ch am b e rs c lo s ely u n i t ed wi th each othe r T he p e ri car di al s ac sh ou ld b e .

r emo ved if t hi s ha s n ot b een d o n e p r evi o u s l y In cas e the t h ym u s gl an d is

.

w ell develop ed it will b e n e c e s s ary t o dis s e ct thi s aw a y f r om the a n t e ri or p a rt

,

o f the he a rt T he gr e at e r p ort i on o f the he art con s i sts o f the two ven tr icl es
. .

,
T he s e co n st i tu t e a rm thi ck w alled c on e h avin g a p o s t eri o r p o in t ed ap ex an d a
-
,

b r oad an t e ri o r b as e This c on e con s i s ts of two com p le t ely s ep ara t ed ven t ri c les

.
,

the righ t an d left ven t ri c le s ; the divi s i on b e t w ee n t hem is m ar ked ext e rn a lly
by an in di st in ct lin e o r gr oove ext en din g f rom the lef t s ide o f the b as e ob liqu ely
t o the r igh t an d t e rm in at in g t o the righ t o f t he ap ex
,
T he gr o o ve con ta in s
.

b r an che s o f the co r on ary art e ry an d vein w hi ch will b e foun d r am ifyin g o ve r the

s u r f ac e o f t he ve n t ri c le s T he left ven t ri c le is mu c h the la rge r o f t he two an d
.

in c lu de s the ap ex An t e ri o r t o the b a s e of each ven t ri c le is a mu ch sm a lle r

.
,

t hin w a lled
-
gen e ra lly d ar k c o lor ed ch amb er t he au r icl e o r a tr iu m E a ch
,
-
, .

a u r i c le in t he co n t r ac t ed s t at e p r e s e n ts a l o b e t he a u r icu l ar app en dage p r o je c t

, ,

in g medi a ll y an d s ligh t ly p o s t e ri or l y o ve r t he ve n t r i c le ; in the c at ( an d man )

this l ob e has a s ca ll op ed m ar gin an d is s h ap ed s om e t hin g like t he hu m an ea r

( he n c e t he n am e a u r i c le m e a n i n g li t
,
t le e a r ) E x t.e n
t
di n g a n t e r i o r l y f r o m t h e

m iddle o f t he b as e o f the ve n t ri c le s f orw a r d b et w een t he t wo au r i c le s is a l a r ge

a rt e r y t he pu l mon ary ar ter y
,
T his make s an ar ch to the left an d di sapp ea r s
. .

D ors al to the pu lm on ary is an othe r art er i a l t run k the aor ta T he s e two t run k s ,
.

a r e ge n e ra ll y imb edded in fat w hi ch s h o u ld b e r em o ved T hey r ep r e s en t the

.

sp li t ven t r a l ao rt a A con u s art e r i os u s is lac kin g the art e rie s sp rin gin g di re ct ly
.
,

f rom the ven t ri c le s Grasp the ap ex o f the he art an d tu rn the heart f orwar d
.

an d n ot e the b as e s o f the gr e at vei n s (p u lm o n ar y an d s y s t emi c vein s) e n t e r in g

t he au ri c le s T he r e is n o s in u s ven o s u s p r e s en t a s a dist in ct c h amb er ; it is

.

g r e a t ly r edu c ed an d ab s orb ed in to the righ t au ri c le w her e it m ay b e l ocat ed by

p h y s i o l o gi ca l e xp e r im e n t s a s a s m a ll s p o t a t t he p o in t w he r e t h e s y s t e m i c vei n s

en t e r t he righ t au ri c le F rom t he mo rp h o l o gi cal p oin t ofview w e may s ay t h at

.
,

the m amm a li an he art con s i s ts o f b u t t wo dif fe r en t c h amb e r s ( e a ch o f whi ch is ,

h ow eve r d ou b le) in c on t rast wi th the sh an d amp hib i an hea rt in whi ch ther e

, ,

In human a n a to my o nl y the ea r shap ed l obe is n amed the au ricl e, the whol e

- c ham ber bein g
t erm ed at r ium . In c o m p a ra tiv e a n a tomy , ho wever , the t erms at riu m a n d au ricl e a re reg a rd ed as

syn o n ym o us .
2 5o LAB ORATORY MANUAL FOR VERTEB RATE ANATOMY

trac n i g its b ran che s t ear t he m e s en t erie s whi ch b i n d t oge the r the co il s o f the
,

in t e st in e as far as n e c e s sa ry an d a ls o s t rip off fat an d lym p h glan d s The .

in tes tin al vei n is the l a r ge ve s s el c o lle ct i n g f r om the gr e at e r p art of t he sm a ll

in t e st in e I t run s in t he middle o f the m e s en t e r y r e c eivin g m an y t ri b u t arie s
.
,

in its cou rs e T he b ran c he s f rom t he jej un u m imm edi at ely b eyon d the
.

du oden u m h ow ever en t e r t he p os t e ri o r p an cr eat i co du o den al vein T he ve ry

, ,
-
.

l arge il eocaecocolic vein co lle c ts f r om the ileum app en dix c a e cu m an d a s c en d , , ,

in g an d t r an s ve r s e c o l o n s Chief am on g it s t ri b u t a rie s a r e : t he a ppeml icu l a r

.

vein f rom the app en dix ; the an ter ior il eocaecal vein f r om t he s accu lu s r otun d u s ,

p ro x im a l p a r t o f t he c a e c u m a dj ac en t ile um an d as c en d in g co l on ; an d t he
, ,

p os ter ior il eoca ecal vein f r om the di s ta l p a r t o f t he ca e c u m a dj ac en t ile um a n d , ,

a s c en di n g c o l on .
0

D raw s h ow i n g the p o r ta l s ys t em
, .

Cat : T he r s t t r i b u ta r y o f the p o r ta l o n fo ll o wi n g the p o r ta l a w ay f r o m

,

t he live r is t he cor on a ry ve in f rom the s t om ac h

, T hi s lie s in t he cu rve b etw een .

p y l oru s a n d s t o m ac h an d a t t he le s s e r cu r v a tu r e is f o rm ed by t he u n i o n o f m a n y

b r an c he s f r om b o t h s ide s o fthe s t om ac h At a b ou t the s am e level as the e n t ra n c e

.

o f t he c o r on a ry vein the a n ter ior p an cr ea tico du oden al vein en t e r s t he t l it

p o r a ;
-
,

c o lle c ts f r om the p a n c r e as an d d u o den um At the s am e level a s the two p r e c ed

.

in g the r ight gas tr o ep ip l oic vein en te r s com in g fro m the p y lo r i c r egion an d g r e a ter
,
-
,

c u r v atu r e o f t he s t om ac h a n d a dj a c e n t gr e a t e r om en tu m T he t h r ee vein s j u s t .

de s cri b ed may en t e r t he p o r ta l s ep ar at ely o r m ay u n i t e wi t h each o t he r in an y

c om b in a t i on b ef o r e e n t e r i n g t he p o r t a l B e yon d t he en t r an c e o ft he s e s m a ll vein s
.

the hep a t i c p o r t a l r e c eive s a l a r ge t r i b u t a ry the ga s tr os p l en ic ve in T hi s p as s e s

,
.

t o t he lef t in t he s u b s t an c e o f t he p an c r e a s r e c eivin g on e o r m o r e sm a ll middl e

g a s tr o e-
p ip l o ic vei n s f ro m t h e s t om ac h w a ll a n d om en tu m an d a p an cr ea tic vein

f rom t he p an cr e as B eyon d t he s e t r i b u ta rie s the gas t r o sp len i c is f o rm ed by t he

.

u n i o n o f t wo m a in b r a n c he s t he r ight a n d l eft s p l en ic ve in s
,
T he lef t s p len i c .

vein p a s s e s in the ga s t r o sp len i c ligam en t a lon g t he s p leen r e ceivin g b ran che s f rom ,

t he s p leen t he g r e a t e r om e n tu m an d s eve r a l l eft gas tr o

, ,
epip l o ic vein s f r o m the

o m e n t u m an d s t o m a c h T he r igh t sp len i c vein c om e s f r om t he righ t en d o f

.

t he s p leen r e c e iv in g a ls o t rib u t a r ie s f r o m t he om en tu m an d s t o m ac h w a ll
, .

B e yon d the e n t r an c e Of t he g a s t r o sp len i c vein the p o r t a l is kn own a s t he s u p er ior

mes en ter ic vein Thi s s oon r e c eive s a sm a ll p os ter ior p a n crea tico du oden al vein
.
-

fr om the p an c r e a s an d di s t a l p art o f t he d u o den u m ; n e x t the infer ior mes en ter ic ,

vein f rom t he de s c en din g co lon a n d r e ctu m ; an d then is s ee n t o b e f orm ed by

n u m e r o u s c on ve r gi n g in tes tin al b r a n c he s f r om t he sm a ll in t e s t in e ca e c u m a n d , ,

a s c en din g c o l on T he lymp h glan d s lyin g a l on g the s u p e r i o r me s en t e r i c ve in

.

as w e l l a s fa t s h o u ld b e r em o ved in t r a c in g t he b ran c he s
,
.

D raw the hep at i c p o rt a l s ys t em .

It will be seen h a t the

t l
r e a tion s o f the h ep atic po rt al s y s tem are the s a me in mammal s
as in all o t h er vertebra tes . The s ys tem co n vey s all o f the v
en ou s b l ood fr o m the d ig es tive
 C OMPARATIVE ANATOMY OF T HE C IR CUL ATORY SYST E M 2 51

t rac t in to the l iver T he pu rpo s e o f th is a rran gemen t is th at the liver cell s

ll
c a p i a ries o f the .

may r emove from the b l ood the dig es ted foo d ma terials There is n o ren al p o r tal s ys tem in .

mamm al s it h avin g b een comp l etel y u su rp ed by the p os tcaval vein

, .

3 . The sy s t e mic ve in s .
T he e r ar e t h ee
r s y s t e mi c vei ns in the r abb i ttwo

p r e ca v a l s a n d o n e p o s t c a v a l an d t wo in t he cat on e p r e caval an d o n e p st
o

ca v a l . T he con di ti on in the c at is du e s imp ly t o t he i

u n on o f t he two p r ec ava l s
an t erior he art ( F ig 59 p
t o the . Alt h ou gh the b ran c he s ar e s imilar in
,
.

t he two an im a l s t he y w ill b e de s c r i b ed s ep ara t el y

,
.

a ) The b r an ches of the p reca val vein : T hi s vein is a l s o c a lled t he an te r i o r

ve n a c ava an d de s cen din g ven a c ava .

R ab b it : T u r n the ap ex o f the he ar t f o rw ar d an d exa m i n e the g r e a t vei n s

w hi c h en t er t he righ t au r i c le T he l eft p recaval vein come s f rom the left an d
.
,

p a s s in g a r o u n d th e lef t a u r i c le e n t e r s the lef t s ide o ft he r igh t au ri c le I t r e c eive s .

s m a ll cor on a ry vein s f r om the he a r t wa ll T he r ight p r eca va l p as s e s di r e c t ly in t o

the righ t a n t e ri o r p ar t o f the r igh t au ri c le N o t e a ddi ti on a l co ron ary vein s I
.

e n t e r in g the r igh t au ri c le di r e ct ly .

Ca r ef u lly t ra c e the r igh t p r e ca v a l f orwa r d c le a rin g aw ay conn e ct ive ti ssu e ,

an d mu s c le f r om a b o u t it s c o u r s e an d f o ll ow it aw a y f r om t he he a r t
,
At the p o in t .

o f e n t r a n c e i n t o t he r igh t au ri c le it r e c eive s f rom b ehi n d the azygos ve in P res s .

the l un g s t o t he lef t an d f o ll ow the a zy gos p o s t e r i o r l y a lon g the d o r sa l t h ora c i c

w a ll n e ar the m edi an li n e N ot e the in tercos tal vein s whi c h en t e r it at s egm en ta l
.

in t e rv a ls ; t hey c ou r s e a lon g the p ost e ri or mar gin of e ac h rib E n t er in g the .

p r e ca v a l im m edi a t el y a n t e r i o r t o t he e n t r an c e o f t he a z y go s is the s u p r eme

in tercos tal vei n the r s t o f the s e r ie s o f in t e r co s t a l vein s

,
S h o rt ly an t e r i or t o .

thi s the in tern al mamma ry vei n e n t e rs the p r e caval T his vein a s c en d s on the

in t e rn a l su rf ac e of the c he st ve ry n e ar the m idven tra l li n e T r ac e it p o st e r i o r ly .

,

n o ti n g b r an c he s f r om the in t e rc o s t a l m u s c le s I t co n t in u e s p o s t e r i o r ly on t he
.

a b d o m i n a l w a ll a s the s u p er ior ep iga s tr ic vei n T he n ex t t r i b u ta r y o f the p r e .

ca v a l is t he ver teb r al vei n I t en te r s t he m edi a l s ide o f t he p r e c av a l at ab ou t the

.

s am e level a s t he i n t e rn a l m amm a ry j o i n s the l a t e r a l s ide I t m ay b e tr ac ed .

deep d o r s a lly t o the c e rvi cal ve rt eb ra e f rom w hi ch it em e r ge s r e c eivin g a cos tocer

vical t r i b u t a ry f r om the n e c k B eyon d thi s p o i n t t he p r e c a va l r e c eive s the l arge
.

s u b c l a vi an vei n f r om t he f o r e lim b Fo llow thi s l a t e r a ll y I t p a s s e s b etw een

. .

the r s t an d s e con d ri b s in t o the axill a an d is t he n kn o wn a s the a xil l ary vei n .

E xp o s e the ax ill a by cu t t in g d ow n t h r ou gh the p e c to ra l m u s c les n e a r the mid

ven t r a l li n e an d a t thei r in s er ti on on the h u m e ru s T he p e ct ora l m u s c le s sh o u ld .

the n b e s ep a r a t ed f r om the u n de r l y i n g s e rra tu s ve n t r a lis b u t s h ou ld n o t b e

r em o ved . T he l arge s t ou t w hi te c o rd s s een c ro s s in g the ax ill a a r e the n e rve s
,

o f the b r a c hi a l p lexu s a n d a r e n o t t o b e i n j u r ed

L ym p h gl a n d s sm a ll r o u n ded .

m a s s e s w ill a l s o b e n o ted in the axill a .

The t erm co ro n ar y ( me n i g literally a crown o r wre ath) is in m amm al s u n fo rtun a tely app lied to
a n

t wo ves s l s tho s e o f the h ea r t wa ll a n d a v ess el o f t he s to mach wal l ha vin g r adia tin g bran ch es
e ,
To .

a o id c o n fu s io
v the l a t ter is refer red to as t he co ro n a ry vess el o f the s to ma h ( co ro n ria en t ric u li)
n c a v .
 2 52 LAB ORATORY MANUAL FOR VERTE B RATE ANATOMY

I n t he

ax illa the ax
.
illary vein r e ceives the f ll i g b r an che s : the l on g
o ow n

thoracic vei n the s u b sca pu l ar vein a n d the cep hal ic vei n

, , T he lon g th o r a ci c
.

vein ru n s cau d a d on the th or aci c w a ll in the s e r ratu s mu scle ; it then p as s es to

the in n e r s u r f a c e o f t he s kin an d ext en d s the e n ti r e len gt h o f the ab d omin al w a ll ,

b ein g e sp e ci a ll y p r om in en t in fem a le s w her e as the extern al mamma ry it c o lle c ts

,

f rom the m ammary gl an d s ( T he gre at e r p art ofthis vein was p rob ab ly r emoved
.

wi th the s ki n ) T he s u b s cap u l ar vein en t e rs the ax ill ary d orsa l t o the p r e c edin g

. .

I t c o lle ct s a con sp i cu ou s b r an c h ( thor acodors al vein ) f r om the l a tis s im u s d ors i an d

c utan e ou s m ax imu s mu sc le s ; it t hen p ass e s th r ou g h t he t e r e s m aj o r m u s c le to

the ext e rn a l s u r f a c e o f the s h ou lde r w he r e it co l le cts fr om v ari ou s m u s c le s . T he

c ep h alic vein is the c hief s u p e r ci a l vein o f the arm I t can b e s t b e pi cked u p

.

o n the ou t e r s u r f ac e o f the u pp e r a rm ; n e ar t he di s tal en d o f the up p e r ar m it

p e n e t r a t e s dee p b e t w ee n m u s c le s a n d p a s s in g b e t w ee n t h e t e r e s m aj o r an d s u b

s c ap u l a ri s m u s c le s em e r ge s o n t he in t e rn a l su r f a c e o f the s h ou lde r an d en t e rs the

ax il l ary vein at the s ame p la c e as o r in c omm o n w i t h the su b s cap u l ar vein .

Immediately b eyon d the en tran ce of thes e trib u taries the axillary vein b ecomes ,

t he b r achial vein o ft he a rm T hi s p ro ceed s a l on g the inn e r s u rf ac e of the upp e r

.

ar m in comp an y w i t h an a rt e r y an d a n e r ve .

R e tu rn t o t he p r e cav a l vein At the p o in t of en tr an c e of the su b c la vi an

.

vein the p re cava l vein r e c eive s fr om the n e ck the exter n al an d in ter n al ju gu l ar

vein s T he ext e rn a l j u gu l ar vein is the lar ge vein whi ch ext en d s forwar d in
.

the dep r ess o r con c h a e p o s t e ri o r m u s c le (m o s t s u p e r c i al mu s c le of the ven tr a l

s u r f a c e o f the n e c k) . I t app e ar s a s the a n t e r i or p ort i o n o f the p r e ca va l . T he
i n t e rn a l j u gul ar vei n is a ve ry sm a ll vein whi ch ru ns a lon gs ide the tr ac he a p as s ,

in g the th yr oid gl an d an d a c comp an yin g the car o tid a r t e r y an d the v a gu s n e r ve

, .

T he p l ac e o f en tr an c e o f the in t e rn al j u g ul ar a s w ell a s its gen e ra l r el a ti on s a r e

highly vari ab le ; it m ay en t e r the p r e cava l a f t e r t he latt e r has r e ceived the s u b
c l av ian b u t it u su a ll y e n t e r s w i t h the ext e rn a l j u gu l ar
,
T he pr e ca val vein m ay
.

t h u s b e s aid t o b e f orm ed by the un i o n o f the s u b c l a vi an ext e rn a l j u gu l ar a n d , ,

in t ern a l j u g ul ar vei n s F o ll o w the ext e rn al j u gu lar S hort l y an t eri or to its

. .

u n i on w i t h the s u b c l avi an it r e c eive s the tr an s ver s e s ca p ul ar ve in f r om the ven t r a l

en d o f the s h o u lde r an d n e ar the s am e level has a c r o ss co nn e ct i on ( tran sver se -

j g
u u l a r vei n ) w i t h it s fell o w o f th e opp o s i t e s ide ( t hi s un i o n w a s p r o b ab l y
de s tr oyed in the pr evi ou s di s s e ction ) Alon g the n e ck it r e c eive s vari ous small
.

t r i b u tarie s f r om mu s cle s an d a b ou t on e in c h p o s t e ri o r t o the an gle o f the j aw s is

s ee n t o b e f o rm ed by the un i on o f two vein s the an ter ior a n d p os ter ior fa cia l vein s
,
.

The an te ri o r fa ci a l vein p ro c eed s t o the an gle o f the j aw s wher e it is s een to b e

formed by the un i on ofvein s f r om the an t eri or p ar t oft he f a ce an d j aws I ts ma in .

t ri b u t a rie s a r e the an gu l ar vei n w hi c h p as s e s o ve r t he ve n t ra l p a r t o fthe m as s e t e r

,

mu s c le an d t he n t u rn s t o the r egi on in f r on t of the e y e an d the deep facial vein ,

whi ch em e rges b etw een the mas s e t e r an d di ga s tri c m u s c les an d p as s e s a l on g the

So na me d becaus e d
the co rre s po n in g vein in man was formerly thought to co nn ec t with the hea d .
2 54 LAB ORATORY MANUAL FOR VERTEB RATE ANATOM Y

p i gastr i c vei n s I n t he1r cou rs e the two mt ern al m amm ary v em s r e cei ve
O O O O O O O

e .

b ran c he s from the diap h ragm c he s t wa ll p er i car di um et c T he p r e cav a l vein

, , , .

n ex t r e c eive s sm a ll b r a n c he s f r om the th ym u s gl an d s an d a dj a c en t m u s c le s a n d
,

at a level b e tw een t he r s t a n d s e c on d r i b s is s een t o b e fo rm ed by the u n i o n o f

two l a r ge vein s T he s e a r e the b r achiocep hal ic o r in n omin a te vein s T he y ar e

. .

the two p r e ca v a l vein s o f e m b ry o n i c s ta ge s w hi c h l at e r u n i t e to f or m the s i n gle

p r e ca v a l vei n o f a d u l t a n a t o m y by th e c r o s s in g o ve r o f t he lef t vei n t o j o i n the

righ t o n e ( Fig 59 p .
,
.T he b ran che s of the two b ra chio c ep hc vein s ar e
iden tic a l an d o n ly on e n eed b e f o ll ow ed p r efe rab ly the right on e s in c e the righ t
, , ,

s i e has n o t b een t ou c hed in the p r evi o u s di s s e c t i on T he l e e e

d p a.c s o f n t r a n c

o f t he v a ri o u s t r i b u ta rie s a r e h o w eve r s om e wh a t v a r i a b le
, , .

Immedi ately an t erior to the j u n ction o f the two b rachiocephalics opp osi te ,

the r s t rib e ac h of t hem r e c eive s o n t he d ors a l s ide a l a r ge t r i b u t ar y

,
T hi s is .

lo cat ed by di ss e ct in g on the d o r s a l s ide ofthe vein an d lif t in g the vein T he main .

t f t h e t r i b u t a r y can b e t r ac ed in t o the c e r vi ca l ve r t e b r a e ; it is t he ver teb r a l

p a r o

vein an d cou rs e s in the ve r te b rart e ri a l can al c o lle ct i n g fr om the b r ai n an d sp in a l

,

co r d . B ef or e it e n t e rs t he b r ac hi oc ep h a li c the ve rt e b r a l is j o in ed by the cos to cer

vical vein wh ,
i c h com e s f r om t he m u s c le s o f the b a c k an d r e c eive s b r an che s a l s o
,

f r om the che st wall on the in n e r su rf ac e o f the r s t two ri b s T he co s to ce r vi ca l

. .

vein may b e p i cked u p by tu rn in g the an im a l d or s al s ide u p an d o n the s ide ,

whe r e the mu s c le s w e r e di ss e ct ed di s s e c t in g in t he s e rratu s ven t ra li s an d the

,

ep axi al mu s c le s T he c ommun i cati on of the ve rt eb ra l an d cos to c e rvi ca l vein s

.

w i th the b rachi ocep h a li c an d wi th e ach o the r is vari ab le an d m ay n o t b e a s

de s cri b ed he r e .

T he b r achi ocep h a li c at the s am e p l ac e as the en tr an c e o f the vein s j u s t

de s cri b ed is s een to b e f o rm ed by the un i on oftwo l arge vein s a lat era l s u b cl avia n ,

an d an an t e ri or ex ter n al ju gu l ar T he su b c l avi an w ill b e f o ll ow ed r s t I t

. .

p a s s e s l a t e r a ll y in f r on t o f t he r s t r ib i n t o t h e a xill a w he r e it is kn o wn a s t he
,

axil l ar y vein E xp os e t he axill a by cu t t in g t h r ou gh the p e c t ora l mu s c le s n e a r

.

the midve n t r a l lin e an d a t t he ir in s e rt i o n o n t he hu m e ru s T he p ec tor a l mu s c les

.

s h ou ld t he n b e s ep a r a t ed f r om the un de r l y in g s e r ra t u s ve n t ra l i s b u t s h o u ld n o t

b e r em oved T he s t ou t w hit e cor d s c r o s sin g t he axill a a r e t he n e rve s o f t he

.

b ra chi al p lexu s an d a r e n o t t o b e i n j u r ed Lym p h gl an d s w ill a l s o b e n o t ed in

.

the axill a . T he m o st m edi al tr i b u t ary o f t he axill a ry vein is t he la rge s u b

s ca p u l ar vein w hi ch p ass es th rou gh the p r oxim a l p a r t o f the u pp er a rm to the

d orsal side of the hu meru s an d colle cts fr om vari ou s mu s cle s of the upp er arm
an d s h ou lde r r e c eivi n g al s o the p os ter ior circu mex vei n f rom the ex t e rn al su r
,

face of the upp er arm T he b egin n in g s o f the su b scapu l ar vein will b e fo un d in

.

the tr ap ezi us mu s cles T he axill ary vein l a teral to the en tran c e o fthe su b s cap u
.

l ar r e ceives the small ven tr al thor acic vein f r o m the m e di al p o r ti on s of the p ec tor
al m us cles L ateral to thi s it r e c eives the l on g thor acic vei n w hi ch ru n s cau d ad
.
,

a l on g the i n n e r su r f a ce o f the p e c to r al m us cles ; an d the thor a codor s al vein whi ch ,

 C OM PARATIVE ANATOM Y OF THE CI R CUL AT OR Y S Y STE M 2 55

cou r s s e p ara llel to the p r e c edin g b u t d ors al to it an d co lle cts c hiey f r om the
l at is s imu s d ors i mu s c le T he r e is a b r oad con n e ct i on b etw een the t ho racod or s al
.

a n d s u b s c ap u l a r vei n s . L at e r a l t o t he s e b r an che s t he ax ill ar y vein is kn own a s

t he b r a chia l vei n . I t ru n s a l on g t he in n e r s u r fa c e o f t he u p p e r a rm in comp an y
wit h n e rve s an d the b r ac hi a l ar t e ry T he s e s tru ct u r e s will b e fou n d by s ep ar a t
.

in g the mu s c le s o n t hi s s u r f ac e o f t he u p p e r arm .

R e tu rn n o w to t he ex t e r n al j u gu l a r vei n . I t s oon s r e c eive s on it s medi a "

s ide t he ve ry sm a ll in ter n al j u gu l ar vei n w hi c h p as s e s f o rw ar d in t he n e c k a l on g

s ide t he t r a c he a in c omp an y w i th t he c a r o t id a r t e ry an d v a g u s n e r ve . T he
m u c h l ar ge r ext e rn a l j u gu l ar vein as sume s a mor e su p e r ci a l p o s i t i on an d in
a ddi t i on t o sm a ll b r a n c he s f r om a dj ac e n t m u s c le s r e c eive s t he l a r ge tr a n s ver s e

sca p u l a r vei n f r om t he s h ou lde r This p as s e s l at e r a lly in f r on t o f t he s h ou lde r

.

a n d an a s t o m o s e s w i t h the cep hal ic vein o f t he arm . T he c ep h a li c vein is t he

su p e r ci a l vein o f t he f o r e l im b an d wi ll b e f ou n d o n the ex t ern a l o r l a t e r a l

s u r f ac e o f the u p p e r arm I t a l s o c on n e c t s w i t h the p o s t e r i o r c i r cu m ex vei n

.

de s cr i b ed ab ove T he ex t e rn a l j u gu l ar an t er i or t o the en t ran c e of the tran sve r s e

.

s capu l ar vein is s i tu a t ed in t he s t e rn o ma s t o id m u s c le . On f o ll o wi n g it f o rw ar d
it is s een to b e f o rm ed a t t he an gle o f the jaw by the u n i on o f t he a n ter ior an d
p os ter ior fa cial vems At t hei r p o in t o f u n i on t he y are c onn e ct ed a cr o ss the
.

ven tra l s ide o f t he t h r oat by the tr an s vers e vein w hi ch has p ro b ab ly b een

de st roy ed T he an t e ri or f ac i a l vein co lle ct s f rom the fac e an d j aw s an d su b
.

m axill ary an d lym p h gl an d s it s m ai n t r i b u t a ry b ein g t he an gu l a r vei n f rom t he

,

regi o n o f t he e y e . T he p o st e r i o r f aci a l vein em e r ge s f r om the p ar o t id gl an d an d

a t the p l ac e o f em e r ge n c e r e c eive s t he p os ter ior au r icu l a r vein f r om t he p i nn a

a n d b ac k o fthe he a d . T he m ain vein t hen l ie s imb edded in the p a r ot id gl an d an d

m ay b e f o ll ow ed by di s s e ct i n g aw ay t he gl an d . I t is t hen s een t o b e f orm ed by
the u n i on o f vei n s f r om the te mp o ra l r egi o n an d r egi o n an t e ri o r t o t he ea r .

D raw t he b ran che s of t he p re caval as far as fou n d .

b ) The b r an ches of the p os tcaval : T he f o ll ow i n g de s c r ip t i on app lie s t o b o th

t he ra bb i t an d t he cat . Tu rn the ap ex o f the he ar t forw a r d an d n o t e t he la r ge
.

vein which en t ers t he r ight au ri c le f rom b ehin d T hi s is the p os tcaval vein

/ .

( a l s o c a lled ve n a c a v a p o s t e r i o r o r i n fe r i o r a n d a s c e n d in g ve n a c a v a.
) I t p ass e s

p o s t e r i o r l y in t he t h o r a x l y i
,
n g s ligh t l y t o t h e r igh t o f t h e m edi a n li n e i n c l o s ed
,

in the f r ee d o r s al b o r de r o f t he ca v a l f o ld o f the p le u r a F o ll ow it cau d a d I t

. .

p a ss e s t h r ou gh t h e d i a p h r a g m f r o m w hi c h it r e c eive s s eve ra l p h r e n ic vei n s . I n

the r ab b it it t hen lie s a ga i n s t the d or s a l w a ll o f t he p e r i ton e a l ca vi ty s ligh t ly

t o t he righ t o f the m edi an lin e d o rs al t o the r igh t m edi an l ob e o f the live r an d
,

in con ta c t w i th the hep a t i c p o r ta l vein . I t then p a ss e s i n to the righ t l a te r al

lob e o f the live r f rom w hi ch it em er ge s n e ar the righ t kidn ey I n the cat the .

p o s t ca v a l ve in p a ss e s i n t o t h e r igh t m edi a n l o b e o f t h e live r a n d i n c l o s ed in t h e

l ive r s u b s tan c e t r a ve r s e s the len g th o f the live r em e r gin g f r om the p o s t e ri o r

lob u le o f the r i gh t l at er a l lo b e N ot e the lar ge hepatic Veins Whic h ow from the
.
2 56 LAB ORA OR T Y MANUAL T T
FOR VE R E B RA E ANA OMY T

live r in to the po st caval vein T hes e ar e b e s t s een by di s s e c tin g in the su b s tan c e

.

o f the liver F o ll ow the p os t c av a l p o s t e ri o r l y car efu lly c le an in g aw ay con n e c

.

t ive t i s su e an d fa t f r om it an d it s t ri b u t ar ie s I t ru n s s ligh t l y t o the r igh t o f

.

the mid d o r s a l lin e o f the p e r i t on e a l c avi ty a l on gs ide the d o r s a l ao rta whi ch

-

mu s t n o t b e in j u r ed T he r s t t ri b u t a ry Of the p o st c av a l is the r ight adr en o

.

l u mb ar vein T hi s p ass e s a l on g the p o s t e ri o r s u r fac e o fa sm a ll gl an d the adr en al

.
,

gl an d w hi ch lie s an t e ri or t o the kidn ey in con t act wi t h the p o s t cava l vein an d

,

w ill b e foun d by di ss e c t in g in the fat in t hi s l o cati on T he adr en o lu mb ar vein

.

r e c eive s b r an c he s fo r the a d r en a l gl an d ( w hi c h is o n e o f the gl an d s Of i n t e rn a l

s e c r e t i o n ) a n d a ls o co ll e c t s f r om the a dj a c e n t b o d y w a ll

. Imm edi at ely p osterior
t o this vein the l arge r ight r en al vein p as s e s f r om the kidn e y in t o the p o s t cav a l .

N ex t by t u rn in g the vi s c er a t o the righ t locat e the lef t a d r e n a l gl an d an d kidn ey

,

an d n d t he l eft a d r en o l um b ar an d r en a l vein s T hey a r e s itu at ed p o s t e ri or t o

.

the righ t o n e s T he left ad ren o lum b ar an d r en a l vein s ge n e ra lly un i t e to a

.

c omm o n s t e m b ef or e t he y en t e r the p o s t c av a l In to the lef t r en al vein open s

.

the vein o f t he lef t g on a d In m a le sp e c im en s this is t he l eft in ter n al s p er ma tic

.

vein ; it m ay b e tr ac ed p ost e ri or ly (in c on tact wi t h the p os t cav a l in the rabb i t)

t o t he s cr o tu m I n fem a le sp e c im en s it is the l eft ovar ian vein w hi c h com e s f r om
.

the ov a ry a sm all o v a l b o d y ly in g ab ou t the m iddle o f the l a t e ra l w a ll o f the

,

p e r i t o n e a l c a vi t y T he r ig.h t in t e rn a l s p e rm a t i c o r o v a r i an vei n e n t e r s t he p o s t
c a v a l di r e c t l y in t he c a t s h o rt l y p o s t e ri o r t o the righ t ki dn e y in the r a bb i t m u c h
, ,

f art he r cau d ad T he p os tcava l vein in it s c o u rs e a lon g t he b od y wa ll r e c eives

a t r e g u l a r in t e r v a l s t he p a i r ed l u mb ar vein s f r om the w a ll ; t he s e a r e s ee n by

l oo s en i n g the vein rai s in g it s ligh t ly an d loo kin g on its d o r s a l su r fa ce T he

, ,
.

lum b ar vein s are then s een p as s in g ven tra lly in the m edi an groove b e tw een mu s c le
m ass e s . N e ar the p os t e ri or en d o f the p er i ton e a l cavi ty the p os t ca v a l r e c eive s
a p ai r o f iliol u mb ar vei n s E ac h o f t he s e in c omp an y w i th an ar ter y ex t en d s
.

lat era lly a lon g the b od y wall an d r e c eive s an an t e r i o r b ran ch fr om the n eigh b o r
h ood of the kidn ey S om e time s the lef t ov ar i an vei n en t e r s the lef t ilio lum b ar
. .

P osterior to this poin t the d o rsal aorta com e s to lie ven tral to the postcaval ,

con c e a lin g the l a tt e r T he di ss e c tion of the r em a in der o f the p o s t c av a l will

.

t he r ef o r e b e defe rr ed u n t il the aorta is s tu died .

I t is ve ry c ommon fo r the p os tcav a l an d its b r an c he s t o v a ry c on s ide r a b l y

f r om the for eg oin g accou n t An app ar en t sp li ttin g o f the p o s t cav a l in to two
.

m ai n t ru n ks p os t e ri o r to the kid n ey s is a comm on v ari ati on ; o the rs ar e m en ti o n ed

in R an d J p age ,

D raw the p ost cava l a n d its b ran c he s as far a s fo ll ow ed .

4. T h e p u l m on ar y ve in s E x a m in e th.e r oo ts o f th e l u n g s a n d n o t e n u m e rou s

vein s s eve r al on each s ide en t er in g the left au ri c le f r om t he lu n gs T hes e are

, ,
.

the p u lm o n a ry vein s T hey lie to either s ide of the p o st ca va l vein th o s e o f the

.
,

Fo r mo re d eta il ed a ccou n t o f the occ u rr en c e a n d o rigin o f th es e va r ia t io n s of the pos tca val in

a

the ca t an d m a n s ee Hu n tin g ton an d M c c l u r e Anatomical Recor d , D ec emb er , 1 9 20

. .
2 58 LAB ORATORY MANUAL FOR VERTE B RATE ANATOMY

o
c he st Wa ll an d co n ti n u e s on the ab d om en a s the su per ior epigas tr ic ar t er y At .

the s am e level f r om t he p o s t e ri o r s u rf ac e of the s u b c l a vi an p r ac t i ca ll y in c omm o n

,

wi t h t he p r e c ed in g t he s u pr eme in ter cos tal a rt e ry a ri s e s I t ru n s p o st eri or ly o n

, .

t he d o r s a l w a ll o f t he t h o r ax an d r e c eive s the r s t in ter cos ta l art e rie s On its

.

a n t e ri o r s u r f ac e a t a b ou t t he s am e level a s t he s e t he s u b c l a vi an a rt e r y give s ri s e

to t he ver teb r al a rt e ry w hi c h p as s e s imm edi at ely d o rs a d t ow a r d t he c e r vi ca l v er

t eb r a e w he r e it e n t e r s t he ve r t e b r a rt e ri a l ca n a l ; an d t o the s u p ercial cervical
a rt e r y w hi c h a s c e n d s in t he l a t e r a l p a r t o f t he n e c k s u pp l yi n g v a r i ou s mu s c le s
, ,

its m ain b r an c h ( a s cen din g cervical ) a ccom p an yi n g the e xt e rn a l j u gu l a r vei n .

T he tr an sver s e a r ter y of the n eck le ave s t he s u b c l a vi an a t the s am e p l a c e o r in

com m on w i t h t he s u p r em e in t e r c o s t a l a r t e r y I t p as s e s d o r s a ll y in f r on t o f t he
.

r s t rib t h rou gh a l oop f o rm ed by two n e rve s an d em e rge s on the m edi a l s ide

,

o f the s e rr at u s ve nt r a li s m u s c le . I t is b e s t f ou n d by l oo ki n g o n t hi s m u s c le an d
the n t r a cin g t he a rt e r y t ow a r d t he s u b c l avi an Af te r givi n g off the f o r ego i n g
.

b r an c he s the s u b c l avi a n p as s e s in f ron t o f the r st rib in t o the axill a w he r e it

is n am ed the axil l ary ar t e ry . T hi s lie s b e tw ee n two o f t he s t ou t n e r ve s b el on g
in g t o t he b ra c hi a l p lexu s . I t s b r an c he s a r e s im il a r t o t h o s e o f t he a xill a ry vei n
a n d ac com p an y t he vei n s . Aft e r givi n g r is e t o t he sm a ll thor acoacr omia l a rt e ry
to the p e ct o r a l an d del t o id m u s c le s t he a x ill ar y give s o f
,
f the l on g thora cic an d
s u b sca p u l a r a r t e r ie s a ccom p an y i n g t he vei n s p r evi o u s l y de s c r i b ed
,
T he fo rm er
.

ru n s p o s t e r i o r l y a l o n g t he s e rr a t u s m u s c le an d t hen as t he exter n al ma mmar y

a r t e r y p a ss e s t o t he u n de r s u r f ac e o f t he s ki n o f t he l a t e r a l a b d om i n a l w a ll b e in g ,

e sp e ci a lly con sp i cu ou s in fem a le s ( M o st of t hi s ve s s el was de s t r oyed in r em o v

.

in g t he s ki n ) T he s u b s cap u l a r has a c o n sp i cu o u s b ran c h ( thor a codor s al a r tery)

.

p ass i n g ca u d a d t o t he l a t i s s i m u s d o r s i a n d cu t an e ou s m axim u s m u s c le s ; it

t he n tu rn s d o r s a ll y an d p e r f o r a t i n g the t e r e s m a j o r em e r ge s on t he o u te r s u r f ac e
o f the s h ou lde r ,
s u p p l yi n g v a r i ou s m u s c le s N e ar the p o in t Of o rigi n o f t he
.

s u b s ca p u l a r the deep a r ter y o f the a rm a r i s e s a n d a f t e r givi n g o f

,
f b r an c he s i n to
the su b s cap u l a r m u s c le p a ss e s b e tw ee n t his m u s c le an d t he t e r e s m a j o r t o the
d or s al p art o f the a rm w he r e it ru n s in com p an y wi t h on e b r an c h o f the c ep ha li c
vei n an d a n er ve all th r ee s itu at ed in t e rn a l to t he l a t e ra l he a d o ft he t r i c ep s w hich
,

s h ou ld b e dee c t ed . T he axill a ry a r te ry n ow p a s s e s t o t he u pp e r a rm w he r e as ,

the b r a chial a rt e ry it cou r s e s a l on g t he i n n e r s u r fa c e o f the lim b in comp an y

w ith the b r a c hi a l vei n an d n e rve s .

D raw t he b r an che s o f t he s u b c lavian .

Cat : At t he level o f the r s t r ib the su b cl a vi an has f ou r b r an che s : in ter n a l

ma mma ry ver teb r a l costocervical axis an d thyr ocervical a xis
, , , The in tern al ma m
.

ma ry sp ri n gs fr o m the ven tr al s u rf a c e o f the su b cl avi an a cco m p an ie s the cor ,

r es p on din g vei n a l o n g t he c he s t w all an d p as s e s o n t o the ab d o m in al wa ll a s

,

t he su p er ior epigas tr ic a r t e r y . T he ver tebr al a r t ery ari s es fr om the d or sal su r

fac e o i the su b c l avian an d pas s es d o r sa ll y in to the v er t eb r a terial can al givi n g ,

of f sma ll b r an c he s in to t he n e c k mu scle s T he costocer vical a xis divides in two

.

a lm o s t a t on c e . On e b r an c h t he su pr eme in ter cos tal a r t erv as e os t rio r l

 COM PARATIVE ANATOMY OF THE CIR CULATORY SYSTE M 2 59

n ear t he d orsal lin e o f the th o rax givi n g off.in tercos tal b ran che s an d then
mid -
,

s u p pl y in g t he deep mu s cle s o f the b a ck T he o ther b ran ch o f the co s to cer vi ca l

.

ax i s le aves the t h o rac i c c a vi ty p ass i n g deep d o r s a ll y in f r on t o f t he r s t r ib

, .

an d div ide s i n t o the tran sver se ar ter y of the n eck su pp lyin g the s e r ra tu s ven tra l is
,

an d r h omb o ideu s m u s c le s a n d the deep cer vical ar t e r y to the ep ax i al mu s c le s

,

o f t he n e ck T he s e b ran c hes a r e b es t foun d by l o okin g amo n g the mu s cle s in

.

q u e s ti o n an d tra cin g the ve ss el s towar d the su b cl avi an T he thyr ocer vical axis .

gen e rally ar i s es an t er i or to the o ther b ran che s I t passe s fo rwar d n ear the .

ca r o tid a r te ry an d a f t e r b ran c hin g to the mu s cles o f the d o r sa l side o f the n e c k

, ,

tu rn s l a t e ra ll y in f r o n t o f the s h o u lde r b ei n g t he n n am ed the tr an sver s e sca pu l ar

,

ar t e ry ; it a cc omp a n ie s the ex t e rn al j u gu l a r vein fo r a sh o r t di s t an c e a n d su p p l ie s

man y m u s cles o f the s h o ulder an d n e ck .

T he su b c l a vi an a r t e ry n ow pass e s in fr on t o f the r st rib in to t he axilla ,

whe r e it is n am ed the axil l a ry art e ry Thi s give s o ff the ven tr al thor acic ar te ry
.
,

p a s s in g m edi a ll y t o t he m edi a l e n d s o f t he p e c t o r a l m u s c le s ; t he l on g th or a cic

a rt e r y p as s in g p o s t e ri o r l y a l o n g t he m iddle r egi on o f t he p e c t o ra l mu s c le s an d
,

t hen to t he l at i ss im u s d o r s i ; an d n e ar t he a rm the l a r ge s u b scap u l ar a r t e ry

, ,
.

T hi s give s off t he thor acodors al a rt e ry lyi n g pa ra llel b u t mo r e d or s al to the l on g

,

th o r aci c a r t e ry an d su pp l yin g the l at i s s imu s d or s i ; the s u b s cap u l a r t hen t urn s

d ors a lly p ass es th rou gh the pr oxim a l p art o fthe u pp e r arm d o r s a l to the h u m eru s
, ,

an d b r an c he s t o the m u s c le s o f the u pp e r a rm a n d m u s c le s o f the b a c k an d

s h ou lde r .

T he axill ary ar tery then pr oceed s as the b rachial to the medi al su r face
o f the f o r e lim b w he r e it a c comp an ie s the b r a c hi a l vein an d s om e n e rve s a n d
, ,

b ran che s in to t he limb .

D raw the b ran che s of the su b c lavi an .

b ) Common car otid ar tery: T he two c omm o n c a ro tid a r t e rie s a ri s e in the

c a t f r om the b r a chioc ep h a li c an d immedi at el y dive r ge ; in t he r abb i t t he r igh t

o n e a r i s e s in com m on w i t h t he righ t s u b c l a vi an w hile t he lef t u s u a ll y sp r i n g s

,

in dep en den t l y f rom the a r ch o f the ao rta T r ac e the c omm on caro tid s fo rwa r d
. .

T heir b r an c he s a r e s imil ar in the two an im a l s T hey p as s an t e ri or ly in the .

n e c k on e t o ea chs id e o f the t ra c he a t o w hi c h t he y give s m a ll b r a n c he s

, ,
At the
.

level of the an t e r io r en d of the th yro id glan d e ach su pp lie s a s u p er ior thyr oid
art e ry t o t he gl an d At the level o f the l a ryn x t he r e a r e b r an che s i n to the
.

laryn x an d a dj acen t p a r ts (p rob ab ly de s t r oy ed) an d an occip ital b ran ch in t o the

d o rs a l mu s c le s o f the n e c k T he common ca ro tid at ab ou t this s am e level give s
.

of f the in ter n al ca r otid a r t e ry I n the ra bb i t thi s a r t e ry a ri s e s a t the p l a c e w he r e

.

the ca r o t id p a s s e s to t he d o r s a l s ide o f the s hi n i n g lig am en t o f t he dig as t ri c In.

the c at it is mu c h sm a lle r an d a ri s e s a t t he s am e level as the o c c ip ita l a r t e r y .

I n b o t h an im a l s the in t er n a l ca r o tid p as s e s d o rs a ll y in com p an y wit h n e r ve s an d

en t ers the s ku ll by a fo ram en t h r ou gh the t ymp an i c b u lla I t n eed n o t b e .

fo ll ow ed T he art e ry b eyo n d this p o in t is ca ll ed t he ex ter n a l car otid a r t e ry

. .
At the

an gle j
o f t he aw it b r an c he s to a ll p a rt s o f hieft bh he
r e
a n c he s a d . I ts c

a r e : t he l in gu al a rt e r y in t o the t o n gu e an d the ex tern al max ill ar y run n in g a l on g

t he ven t ra l b or de r o f the m a ss e t e r mu s c le an d b r an c hin g t o the u pp e r an d l ow e r

l ip s an d j aw s T he m ai n art e ry then p a ss e s a l on g t he p o st e ri o r b or de r o f the
.

m as s e t e r mu s c le I t r e c eive s au r icu l ar an d temp or al b ran c he s f rom the p in n a

.

a n d t emp o r a l r egi on s an d t hen a s t he in ter n a l max il l ar y a rt e r y tu rn s in t e rn a l

, ,

t o the m a s s e t e r mu s c le a n d is lo s t t o view I t n eed n o t b e f o ll ow ed fart he r

. .

D raw the b ran che s of the c omm on caro tid ar tery .

c) Th e thor a cic a or ta : Af t e r h avin g given r i s e t o the s u b c l a vi an s an d the

c a ro t id s the a o rta a r c he s t o the lef t

,
N ot e a s it p a ss e s the left p u lmon ary the
.
,

s t r o n g b r ou s b a n d w h ic h co n n e c ts the two ve ss el s This is the ar ter ial l iga .

men t o r l igamen t of B ota ll u s a n d is the r em n an t o f the em b ryon i c conn e cti on

b etw een the aor ta an d the p u lm on a ry ( F ig 58 p . F o llo w the aorta p o s
, .

t erior l y p r e s s in g t he lef t lu n g t o the r igh t

,
I t de s c en d s p o s t e ri o r l y lyin g aga in s t
.

the d or sa l w a ll o f the t h o r ax t o the lef t o f the m edi an lin e I t is s i tu at ed wi thin .

the m edi as tin um ; the m edi as t i n a l w a ll m ay b e c le a r ed a w ay T he ao rt a in its .

cou rs e a l o n g t he t h o r ax is n am ed t he thor acic a or ta I t s c hief b r an c he s a r e the

.

p a i r ed in ter cos ta l a r t e r ie s w hi c h a r i s e f r o m t h e a o r t a a t s eg m e n t a l in t e r v a l s a n d
ru n a l on g the th o ra c i c w a ll a l o n g the p ost e ri o r m a r gin o f t he ri b s T he a orta .

a l s o has s m a ll b r on chial a rt e r ie s t o the b r o n c hi a n d es op hagea l art e rie s t o the

e s op h a gu s Alon g the d ors al s u rf ace o f the aorta on its lef t s ide ru n s a deli cate
.

tu b e r e s em b li n g a s tr e ak o f fat
,
This is the thoracic du ct the ma in lymp hatic
.
,

chan n el fo r t he p o s t e ri o r p a r t o f the b o d y T r a c e it f orwa r d ; its conn e cti on

.

wi t h the j u gu l ar vein gen e r a lly at the p oin t o f u n ion wi th the su b c l avian may
, ,

b e fou n d .

T he ao rt a p en e t ra t e s the di ap h ragm t o w hi c h in the rabb i t it give s s u per ior

p hr en ic a rt e ries an d p as s es in to the p e ri t on e al cavi ty w he r e it is kn own as
the a b d om in a l ao rt a .

d) The a bdomin al a or ta : T u rn the dige s tive t ra ct t o the righ t an d l oca t e

the d o r s al ao r ta a f t e r it ha s p as s ed the di ap h r agm I t will b e f o u n d a ga in s t the
.

d o rs a l wa ll in the m edi an d o r s al li n e Cle ar away the m e s o gas te r an d c lean t he

.

su rf a c e o f the a o rt a F o ll ow it iden t ifyin g its b ran che s T he s e b ran ches co n

. .

s i s t o f un p ai r ed median vis cer al b r an c he s t o the di ge s t ive t r a c t p a i red l a ter al ,

vis cer a l b r an c he s t o the kid n e y s an d r ep r o d u c t ive o r g an s an d p a i r ed s oma tic ,

b ran che s to the b o d y wa ll .

S h ort ly p o s t e ri o r t o the di ap h ra gm the aort a give s ri s e t o two l a rge u n p ai r ed

vis c er al art e rie s the coel iac an d the s u p er ior mesen ter ic a rt e rie s In the ca t the
,
-
.

s e co n d is s h o rt l y p o s t e r i o r t o the r s t w hi le in the r a bb i t the s u p e ri o r m e s en t e r ic

,

a rt e r y lie s o n e h a lf in c h p o s t e ri o r t o t he c o eli ac
-
As the b r an che s o f t he s e tw o
.

ve ss els a r e di ffe r en t in the two an im al s o wi n g t o the diffe ren c e s in thei r dige s ~

tive t racts it wil l b e n e c e s s a ry t o de s c ri b e t hem s ep ar at el y

,
.

R ab b it : T he co eli ac a rt e ry n e a r its o rigi n f rom the ao rt a give s ri s e t o the

sma l l infer ior phren ic a rt e rie s t o the diap h r a gm B e yon d this p o in t the s pl en ic
.
26 2 LAB ORATORY MANUAL FOR VE RTE B RATE ANATOMY
o

Ca t : T he eli ac ar te ry p as s e s tow ar d the s tom ach an d very s oon divide s

co ,

a t a b ou t t he s am e level in t o t h r ee b r a n c he s
, . T he mo st c r an i a l on e is the hep a tic ,

t he n ex t o n e the l eft gas tr ic a n d m o s t cau d a l an d l a r ge s t is t he s p l en ic

, , . T r ace
the sp le n i c a rt e r y I t cou r s e s in t he g r e at om en tum t ow ar d the sp leen an d f o r k s
. .

On e b r an c h g o e s t o the lef t en d of the sp leen an d s en d s a l so b r an che s in t o the

p an cr e a s an d t he s h o r t g a s tr ic a rt e r ie s t o th e s t o m ac h . T he o t he r b r a n c h p a ss e s

to t he righ t en d o f the s p lee n an d s u pp l ie s a l s o b ra n c he s t o the p an c r e as the ,

om e n tu m a n d the l eft gas tr o epip l oic a rt e r ie s t o the gr e at e r c u r va tu r e

,
-
. T he
lef t gas tri c o r cor on ary art e ry p ass e s to the le s s er cu rv atu r e w he r e it sp li ts in t o
m an y b r an c he s su pp lyin g b o t h s ide s o f the s t om ac h
,
. T he hep a ti c a r t e ry p as s e s
a l o n g the b o r de r o f t he lef t en d o f the p a n c r e a s an d to t he d o r s a l s ide o f the le s s e r

cu r va tu r e an d e n t e r s the hep a to d uo de n a l lig am en t . I t is b e s t f ou n d by s ep ar a t

in g the s t omac h a n d live r . I t lie s t o the lef t s ide o f the hep at i c p o r t a l v ein .As
it p as s e s the p y l oru s it give s off t he l a r ge gas tr odu oden al b ran c h . T hi s b ran che s
in t o the an ter ior p an cr ea tico du oden al ar t e ry de s c en din g a lon g the b egi n n in g o f
-

the d uo de n u m an d s u pp ly in g a l s o the p an c r e a s ; the r ight gas tr o epip l oic p as s in g -

fr om the p y loru s a lon g the gr e a ter cu r vatu r e o f the s t om ac h to the lef t ; an d t he

s m a ll pyl or ic a rt e r y t o the p y l o ri c r egi on ( this m ay a l s o a r i s e in dep e n de n t l y f rom

the hep at i c) .T he hep at i c a rt ery p ro ceed s in to the live r s en din g a cys tic art e ry
to t he ga ll b l a dde r .

T he s u p er i or me s en t eri c a rt e ry s upp lie s the g r e at e r p art o f the in t e s tin e I t .

p a ss e s t o w a r d t he in t e s t i n e . F o ll o w it c
,
le an i n g a w a y fa t an d l ym p h gl a n d s f r o m
it s su rf a c e . I t r s t give s r is e t o the middl e col ic a rt e ry w hi c h p a s s e s t o the tr an s
ve r s e an d de s c en din g p art s of the c o lon A li tt le fa r the r on the s u p e ri o r m e s en
.

t eric give s ri s e s imu l tan e ou s ly t o the p os ter ior p an cr ea tico du oden a l a r t e ry w hi c h

-

as c e n d s a l on g the d uo den um s u p p lyin g it a n d the p an c r e a s a n d an a s t om o s in g

,

wit h t he an t er io r p an cr e a ti c o d u o den a l ; a n d t o the il eocolic a r t e ry to the ca e cum

-

an d t e rm i n a l p o r t i on o f the ile u m an d s e n din g a l s o a r ight col ic b r an c h t o the

as c e n din g c o l on ( thi s l as t may a ri s e in dep en den t l y f rom the s u p e r i o r m e s e n t e ri c) .

T he s u p eri or m e s en te ri c t hen divide s in t o n um e r ou s in tes tin al b ran che s t o the

s m a ll i n t e s t i n e .

D raw the b ra n c he s o f the co elia c an d su p er i o r m e s en t e r i c .

R e tu rn n ow t o the d o r s a l ao rta . I ts n ex t b r a n che s a r e the p a i r ed adr en o

l u mb ar an d r en al a rt e rie s .In the rabb it the ad r en o l umb ars a r e b r an che s o f
the ren a l s b u t in the c a t t he y a r i s e in dep en den t l y
,
.T hey p as s c lo s e to the
a d r en a l gl an d t o w hi c h the y give an a d r en a l b r a n c h an d t hen cou r s e a l on g t he

d o rs a l b o dy w a ll In the c at e ach s en d s a phr en ic art e ry an t e r i o r ly t o t he d ia

.

p h r a gm . T he r e n a l a r t e r ie s a r e l a r ge ve s s el s p a s s in g in t o t h e k id n e y s . T he
a o rt a p o s t e r i o r t o the kidn e ys give s r i s e t o the p a i r ed a r t e rie s t o t he g on a d s ( t he s e

m ay h ow eve r b ran c h f r om the r en a l s)

, ,
. T hey a r e t he in ter n al s p er ma tic a r te rie s
in the c as e o f the ma le an d run p o s t e r i o r ly on the d o r s a l w a ll t o the s cro t u m .

I n the fem a le the co r r e sp o n din g ova r ian a r t e rie s a r e l ar ge r a n d in t he c a t c on ,

 C O MPARATIVE ANATO MY OF THE CIR CUL ATORY SYSTE M 26 3

vo lu t ed an d p as s l at e r a lly t o the o varie s In its passage a lon g the mid d or s a l

,
.
-

lin e the ao rt a give s off p ai r ed l u mb ar art e rie s at s egm en ta l in t e rva ls T he s e .

are f oun d by l o o s en in g t he a ort a an d l o o ki n g on it s d o r s a l s u r f a c e P o st e ri o r

.

to the gen i t a l a r t e r ie s the infer ior mesen ter ic a r i s e s a s an u n p a i r ed vi s c e r a l b r an c h

an d p a s s e s t o the de s c en din g c o l o n a n d r e c tu m run n in g in the m e soco l o n
, In .

the me s o co l o n it f o r k s in t o the l eft col ic a r t e r y p a s s i n g c r an i a d a l o n g t he de s c en d

in g co lo n an d the s u p er ior hemorr hoida l art e r y p as s in g cau d ad t o the p o s t e ri or
p a r t o f t he de s c e n di n g c o l o n a n d t he r e c t um .

Add the s e ve s s el s t o t he d r aw in g of the aor ta .

T he dige s t ive tr ac t may n ow b e r em oved an d di s c ar ded le avin g the en d o f ,

the l a r ge i n t e s t in e in p l ac e . H o ld the stump of the colon together with the

u r i n a r y b l a dde r a n d in fema le sp e c im en s t he u t e ru s ( t he f o r ked co iled t u b e a t

the p o s t e r i o r en d o f t he p e r i t on e a l c avi t y ) b ac k a g ai n s t the pu b e s an d f o ll o w the

ao r t a f a r t he r N ear t he en d o fthe p e r i t on e a l cavi ty it fo rk s in to the two common
.

il ia c art e r ie s in the ra bb i t ; in the c a t it give s o ff a p a i r o f exter n al il ia c a r t e rie s

fo llow ed s h or t ly by a pai r o f in tern al il iac ar t e r ie s An t e r i or t o t hi s p lac e in
.

the c a t o r in the r a bb i t a t the level o f the f o r k o r f r om the comm o n ili ac a rt e r ie s

, ,

a p a i r o f il iol u mb ar a r t e r ie s a r i s e s an d p a ss e s l a t e r a ll y a l on g the b o d y w a ll .

T he ilio lu m b ar a r tery divide s in to an an t e ri o r b ran c h w hi ch p as s e s fo r wa r d ,

t o w ar d t he k idn e y an d a p o s t e ri o r b r an c h w hi c h e x t e n d s t o the t high

, , .

T he two commo n ili ac art e rie s in the r abb it s oo n fo rk in to an an t e r i o r exter n al

il iac an d a p o s t e r i o r in ter n al il ia c I n the cat the ex t e rn a l an d in t e rn a l iliac s
.

a r i s e s ep a r a t el y f r om the a o rt a the l a t t e r imm edi at el y p o s t e ri o r t o the f o rm e r

,
.

Af t e r givi n g ri s e to the ili acs the ao r ta c on t in ues in the mid d or s al lin e as the -

s m a ll media n s acra l o r cau da l a r t e ry ly in g h a lfw ay b e tw een the two in t e rn al

,

iliacs In the ca t t hi s ve s s el ari s e s f rom the fork of the in t e rn a l ili acs In the
. .

r a bb i t it sp r i n g s an t e r i o r t o t he f o r ki n g o f the ao r t a f r om the d or s a l s u r f a c e o f
,

the l a tt e r ; it s o r igi n is c on c e a led by t he p o s t c a v a l vei n a n d w ill b e s ee n l a t e r .

T he s ac r a l a r t er y s u pp lie s the s a c ra l r egi o n an d the tail .

F o llo w t he ex t e rn a l ili ac I t p ass e s l a tero cau d ad ou t o f the p e ri to n e al

ca vi t y in t he r a bb i t t o the d o r s a l s ide o f t he i n gu i n a l l ig am e n t
,
As it p as s e s
.

th ro u gh t he a b d om i n a l w a ll o r s h o r t l y b e yo n d t he w a ll it give s ri s e t o the deep

femor al art ery ( c at ) or the inferior ep igas tr ic ( rabb i t) I n the cat thi s ve s s el .

g i v e s o b r a n c he s i n t o t h e t high w hile
,
t he s e a r e l ac ki n g in t he r a bb i t I n b o t h .

a n im a l s t he f o ll o wi n g b r an c he s a r e p r e s en t : b r an c he s i n t o the m a s s offa t b e tw ee n

the t high s an d in t o the ex t e rn a l ge n i ta l o r g an s o f w hi c h on e b r an ch in m ale

, ,

sp e c im en s c o n s t i t u t e s the extern al s p er ma tic a r t e r y ; an d the m ai n ve s s el then

, ,

a s the infer ior ep igas tr ic a r t e r y tu rn s c r an ia d an d a s c en d s in the a b d omin a l

,

wa ll r un n in g a l on g the in n e r s u r f ac e o f the r e ctu s ab d om in i s m u sc le I t an a s

,
.

t omo ses w i t h the s u p e ri o r ep iga s t r i c a rt e r y In the ra bb i t the r e a ri s e s ei t her

.
,

from the in fe rio r ep igas t ri c at the o rigin of t he l att e r fr om the ext ern al ili ac or
from the ex t e rn a l iliac it s elf n ear by t he s u percial epigas tr ic art e ry whi ch ext en d s
,
26 4 LAB ORATORY MANUAL FOR VERTE B RATE ANATOM Y
0

forward on the in n e r su rf ac e of the s kin of the ab dom in a l wall an d an ast omos e s

wi t h t he ext ern a l mamm ary a b r an ch of t he lo n g t h o raci c T he s e ve ss els ar e
, .

p a r t i c u l a r l y p r om i n e n t in fem a le s ,
b u t t he gr e at e r p a rt o f t hei r c ou r s e is

de st r oy ed in r emovin g t he s kin T he ext e rn a l ili ac n ow n am ed t he femor al

.
, ,

p r oc eed s a l on g t h e c e n t e r o f t h e m edi a l s u r f a c e o f t h e t high givi n g

,
b r a n c he s in t o

the leg mu s c le s . F or t he s e b r an c he s c o n su l t R an d J an d B , .

F o ll ow the in t ern al i li a cs b ein g ca r efu l n ot t o i n j u r e the en d o f the p o st

,

c a v a l vein l y in g in c o n ta ct w i t h t hem n o r a n y p art s o f the u r o gen i t a l s y s t em

( in m a le s d o n o t in j u r e t h e m a le d u ct s cu r vi n g a r o un d t he b a s e o f the u ri n ar y

b l adde r) T he in t e rn a l i lia cs lie agai n st the d ors al w a ll At t hei r o rigin f rom

. .

the comm on i l i a c ( r a bb i t) o r p o st e ri o r t o t hei r o r igi n f r om t he d or s a l ao rt a ( c a t)

each give s ri s e to an u mbil ical art e ry whi ch p as s e s t o the b ladde r o r in fem a le
r abb i t s t o t he u t e ru s r s t wi th a b r an c h t o t he b l a dde r T he in t e rn a l i liacs t hen
.

p a s s t o t h e d o r s a l s ide o f t h e p o s t ca v a l vei n. T o f o ll o w t he m di s s e c t a s dee p l y

as p o ss i b le b et w ee n t he r e c t um a n d t he b as e o f t he t high T he in t e rn al ili ac s
.

give s om e b r an che s t o the p elvi s an d then e ach give s off a middl e ha emorr hoidal
a rt e ry t o the r e ct um . T hi s ac c om p an ie s the r e ct um t o the a n u s b u t cann ot b e
f o llow ed at t hi s t ime In fema le cats the u terin e art e ry ar i s e s f rom the mi ddle
.

h aemorr h oid al an d p as s e s an t e ri o r ly again to the u t e ru s T he in t ern al ili acs .

can n o t b e f o ll ow ed fa rt he r co n ven ien t l y . T hey give b ran c hes t o the tail an d

t high .

D raw the b ran che s of the ili ac s a ddin g t hem to the d rawmg o f the d or s al
aort a a lr e a d y ma de .

7 The pos te r ior portio n of the po s tcaval ve in

.
T he p os tcav al m ay n ow b e
.

fo llow ed cau d a d from the p oin t w he r e it was p r evi ou s ly lef t by r emo vin g the
a rt e r ie s w hi ch co ve r it . I ts t ri b u t arie s s h ou ld b e t ra ced as far a s p r acti cab le t o
the p o s t e ri o r en d o f the p e ri t on e a l c a vi t y di ss e ct in g deep l y d o r s a ll y as b ef o r e
,
.

Rab b it : T he p o s tcav a l r e c eive s a t the s am e level as the f o rki n g o f the d o rsal

,

aorta the two la r ge extern al iliac vein s

,
. I t t he n co n t i n u e s in t he mi d d ors a l -

lin e for a s h ort di stan c e c au dad t o thi s p oin t thi s p ort i on o f t en r e c eivin g the
,

n am e o f common in ter n al i l iac vein an d is t hen s ee n t o b e f o rm ed by the u n i o n

,

o f the two in ter n al il ia c or h y gp o a s tr ic vei n s .T r a c e the ext e rn a l ili ac ; its

b ran che s ar e s imil a r to an d a ccomp an y t h os e o f the a rt e ry of t he s am e n ame .

I t soon r e c eive s the ves ical vein f r om the b l a dde r : t hi s vein in fema le s a l s o co lle cts
f rom the u t eru s At the p lac e whe r e it p as s es th rou gh the ab domin al wa ll the
.
,

ext e rn a l iliac r e c eive s the infer ior ep igas tr ic vein T he main p ar t o f t hi s run s .

fo rwar d a l on g the in t ern al su rf ac e o f the r e ctu s ab d omin i s mu s c le an d an asto

m os e s an t e ri o r l y w i th the s u p e r i o r ep iga s t ri c . T he in fe ri o r ep igas tri c n ea r its
l e f e t e i t t h e t e l il i l e eive t r i b u ta r ie s f r om the fa t
p ac o n r a n c n o e x rn a a c a s o r c s

b e tw een the b a s es o f the thigh s an d the ex t ern a l gen i tal r egi on an d s en d s a ,

su p ercial epigas tr ic vein a l on g the in n e r sur f a c e o f the s ki n o fthe la t e r a l ab domi

n a l w a ll . T his la s t n am ed ves s el is par ti cu lar ly con sp i cu ou s in fem ales b u t is

-
,
26 6 LAB ORATORY MANUAL FOR VERTE B R ATE ANATO MY

p r e ca sfal vein Cu t off the ap ex o fthe heart an d n o t; the thick wa lls an d rou n ded
.

fo rm of the lef t ven t ri c le an d sm a ller s iz e thin n er wa lls an d c r e s c en t i c f orm o f

, , ,

the righ t ve n t ri c le Cu t op e n the righ t ven t r i c le by a n ob liqu e cu t b egin n in g

.

at the cu t s u r f a c e a lr e a d y m a de a n d ex t e n din g ou t th r ou gh the p u hn o n a r t e y

y a r r ,

s li tt i n g op en t hi s a rt e ry Wa s h ou t the righ t ven t ri c le I ts cavity is r a the r

. .

sm a ll the w a ll s b ein g deep l y c lef t by mu s cu l ar ridge s the tr a b ecu l a e car n ac

, , .

F r om the w a ll s p roje c t a n umb e r o f p o in t ed n ger like mu s c le s the p a pil l a r y -

,

mu s c le s w hi c h a r e c on n e ct ed by s len de r b e r s the chordae ten din ae t o a thin

, , ,

memb ran e This m emb r an e co n s i s ts o f th r ee ap s an d is c a lled t he tr icu s pid

.

val ve . T wo o f the ap s can b e s t r e tched by p u llin g o n the cu t s u r f ac e s o f the

ven t ri c le whi le the thi r d lie s co ll ap sed a gai n s t the in t e r ven tri cu l ar s ep tu m to ,

w hi c h it is fa s t en ed w i t h ou t t he in t e r ven t i o n o f the p ap ill a ry mu s c le s T he .

t r i cu sp id v a lve g uar d s the righ t au ri cu l o ven t r i cu l a r op en i n g an d p r even ts t he

-

b l oo d f r om ow in g b ack from the ven tri c le in to the au ri c le In the b as e o f .

the p u hn on ary a r t e r y n ot e the t h r ee p o c ke t s h ap ed s emil un ar v a lve s T he pu l

-
.

m o n a ry a rt e r y is the s o le exi t f rom the r igh t ven t ri c le S imil ar l y cu t op en the lef t .

ven t r i c le by a lon gi tu din a l s li t f r om ap ex to b as e Wash ou t the in t e r i or . .

T he c avity o f the lef t ven t ri c le is c on s idera b ly l arger t h an th at o f t he righ t an d ,

its w a ll s thi cke r T he two ven t ri c le s ar e comp le t el y s ep ar a t ed by the in ter

.

ven tr icu l ar s ep tu m w hi ch app e ar s a s the comm on i n t e rn a l w a ll o f b o t h ven t r i c le s

, .

N o t e in the lef t ven t ri cle the t rab e cu la e ca rn ae the p ap illary mu s c le s an d the , ,

c h o r d a e t en d in a e T he l att e r ar e a t t ac hed to the m emb r an ou s b icu s p id o r

.

mitr a l v a lve whi ch c on s i s t o f b u t two ap s

,
T hi s g uar d s the lef t au r i cu l o .

ven t ri cu lar op en in g an d p r even ts the r egu r gi t at i on o f the b l ood f r om the v en

tr i ele b ac k in t o the au r i c le B y p ro b in g n d the s o le ex i t o f t he lef t ven t ri c le
.
, ,

its op en i n g in t o t he aort a F o ll ow t he p r o b e by a cu t an d n o t e the t h r ee s emil u

'

n a r v a lve s a t the b a s e o f the a o rt a .

M ake d r a wi n gs to illu s t r at e the s t ru ctu r e o f the he a rt .

T he r em ova l o f the he a r t p erm i ts a c le a r er view o f s om e o f the s t ru c tu r e s Of

the p le u r a l ca vi t y T he s tu den t s h ou ld ex amin e ca r efu lly the f o r ki n g o f the
.

t r ac he a in to the b ro n c hi the f o rm o f the l u n g s an d thei r r el a tion to the p le u r a l

,

ca vi t y a n d the p u lm o n a r y a r t e rie s an d ve in s
,
.

9 . C om par is on m amm al iof

an hear t the
an d cir cu l a to ry s y s t e m with thos e o f the p r ec ed

in g a nim al s The c h amb ers of the m ammal ian h ear t l ike th ose o f b ir ds are b u t o f two kin d s
.
, ,

n am el y , au ricl es an d ven t ricl es in c on tr as t to the fou r d iffer en t ch amb er s o fthe sh an d amp h ib

,

ian h ear t an d the th ree o f the r ep til ian h ear t E ach is , h ow ever , comp letely d ivid ed in to two
.

co mp ar tmen ts , righ t an d l ef t , by the fo rm a tion o f sep ta in the cen ter o f the o rig in all y s in g l e
ch amb er T he di vis ion of the a u ricl e in to two ch amb er s b egin s with the A mphibia w hil e
. ,

t h a t o f the ven tricl e beg in s in r ep til es an d is comp l eted in the c r o co d il es , b ir d s an d mammal s ,

.

I n ma mm al s , a s in r ep til es an d b ir d s the con u s a r teriosu s ha s van is h ed l ea vin g as remn an ts

, ,

the s emil un a r va lves a t the b as es o f the g rea t ar teries T he s in u s ven osu s s till p er sis ten t in
.

r ep til es has d isa p pear ed in b ir d s an d mam

m al s , o r more cor r ec tl y s peakin g , is r ep res en ted
,

by a s mall s po t in the wall o f the rig h t au ricl e ; this s po t is n ot d etec ta b l e by gr oss exa min a tion .
 C O MPARATIVE ANATOM Y OF THE CIR CULATORY SYSTE M 26 7

The ven tr al a or ta in m amm al s as in b ird s is sp l it

in to two tru n ks , in con tras t w ith the
th ree tru nks common to rep til es Th ese two a re . the p u l mo rfary a r ter y a n d the a o r ta T he .

p u l m o n a r y a r t e r y l ea ve s t he r ig h t v e n tric l e the a or ta the l ef t ven tr icl e , Al l o f th e s y s temic .

vein s o p en in to the righ t au ricl e an d the pu lmon ary vein s in to the l ef t au ricle T he rig h t .

h al f o f the h ear t is ven ou s an d the l ef t ar terial an d o win g to the co mp l etion o f the in terven
, ,
~

t ricu l ar s ep tu m th er e is n o mix in g ofa r ter ia l an d ven o u s b l oo d in the h ea r t b u t a compl ete a n d ,

p erf ec t d o u bl e c ir cu l a tio n is m a in ta in ed t h r ou g h the h ea r t T h e ve n o u s b l oo d p a s s es f ro m t he .

FI G .
58
.D iagrams how the evolution of the aortic a rches A primi tive con dition with six
to s .
,

ao r tic a rch es B s hes the r s t ao r tic a rch m iss in g

.
, ,
C som e u rod el es t he s t s ec o n d a n d f th
.
, ,
r , ,

ar hes missin g
c D a n u ran s with the con n ec tio n k between the p uhn o n ary a rt eries d an d the a orta
.
, ,

o blit ra tede E r ep til es showin g the v en tr al ao r ta s p lit in t o t h ree t ru n ks an d t he four th a or tic arc h
.
, ,

h a d i p er sis t ent o n bo th sid es

n ,
F bird s the v en t ra l ao ta sp lit in to t wo t ru nks an d the fou rth
.
, ,
r
,

ao r tic a r ch h p er sis t en t o n the righ t s id e o nl y G mammal s the ven tra l ao r ta sp lit in to t wo t ru n ks a n d

.
, ,

t he f ou r th ao r tic a rch i p ersis ten t o n the l ef t sid e o n ly a in ter ru p tio n o f the a or tic a r c h es by the g ill
.
,

c ap ill aries in sh es ; b in t ern a l c ar o tid ; c e t er n a l c a r o tid ; d p u hn o ary d evel op ed fr om the s ix th

, ,
x ,
n
,

a r ch ; e v e t ra l a o r ta ; f d orsa l a o rt a ; g a o r tic a rch; h right fou r th a o r tic a r ch c al l ed t he right aorta

,
n , , , ,

a bo v e u r od el es ; i l f t fou r th a or tic a r ch c all ed the l eft ao r t a a b ov e u ro d el es ; j co mmo n c a ro tid ; k

,
e , , ,

a rt erial lig am en t o r o blit er a t ed v essel o rigin al l y co n ec tin g p u l mo n a ry a n d a o r ta ; l n su b cla vian , .

( Slightly modied from W ilder s His tor y of the Hu man B ody cou r tesy of Hen ry Holt a n d Comp an y )

.
,

ri g h t au ricl e in to h ven tricl e an d o u t th r ou g h the p ul mon ary ar tery to the l u n g s wh ere

the r ig t ,

it is aer a ted T he a r terial b l oo d r e tu r n s by way o f the p u l mon ary vein s to the l ef t au ricl e
.
,

p as s es in t o th e l e f t ven t r ic l e a n d ou t o f the ao r ta , .

L ittl e tr ace is l ef t in the adu l t mamm al o f the origin al s ys tem o f aor tic ar ch es p assin g
ar ou n d the p h aryn x As in all o fthe l an d ver teb r a tes the b ases o fthe c o mmo n caro tid a r teries
.
,

r ep r es en t the r em a in s o f the th ir d aor tic ar ch es ; the arc h o f the ao r ta is the l ef t f o u r th ao r tic

26 8 LAB ORATORY MANUAL FOR VERTE B RATE ANATOM Y

'
5

FI G .
59 .
D ia grams to sho w the developmen t of l
the po st cava v ein in the ca t T he .

s ystem o f v ein s is hat ched the sub car din al

c ro ss ,
v ein s l
c os ely ld
s tipp e , t he hepatic v ein s in dica t e d by
c ros s , ver tica l ,
an d obliq u e h at chin g combin ed ,
the su pracar din al vein s by o p en s tip p lin g ,
an d the

"Con tin u ed on oppos ite page"

 2 7o LAB ORATORY MANUAL FOR VERTEB RATE ANATOM Y

l p o r ta l l vei n s an d th at the po s t d
O O

VCI DS ar e t he o f t he
O

the
O O

ren a p o s teri o r p art p os ter i or car i n a ,

cava l vein b et w een t he kid n ey s is fo r m ed by the s u b car d in al vein s c h ie y t he rig h t s u b car d in al , .

Simil arl y in mamm al s the po s t caval v ein is fo rm ed o f the dis tal en d s o f the p o s terior car d in al s ,

o f t he rig h t su b ca r din a l o f the h ep a tic vein s ( vitell in e) in the r eg ion o f an d an t erio r to the
,

liver an d in the r egion b etween the kidn ey s an d the hin d limbs ofo th er su b ordin a te vein s Th is
, .

will b e cl earer by r eferen ce to Figu re 59 T he man n er o fo rig in of the p o s tcaval f rom s o man y
.

dieren t s ou r ces ex p l ain s the n u merou s va ria tio n s common to th is vein s u ch a s the fr equ en t ,

n din g of a dou bl e po s tcaval an d fu r th er expl ain s the as ymmetrical o rigin o f the vein s to the
,

r ep ro d u c tive o r g an s .

I n con cl u sion it may b e s tated th at the emb ryo logy o f the mamm al ian cir cu l ato ry s y s tem
fu rn ish es a b eau tifu l an d s trikin g examp l e o f the r ep etition of evol u tion ary s tag es I n its .

d evel opmen t the mammalian circu l atory s ys tem p as ses su ccess ivel y th rou gh each o f the s tag es
wh ich we h ave fou n d to p ers is t as the adul t con dition in the typ es we h ave s tu d ied an d the ,

evol u tion of th is s y s t em can b e d etermin ed equ al l y w ell eith er by s tu d yin g its d evel op men t in

the m ammal o r by s tu dy in g an d comp arin g its form in the adu l ts o f the d ifferen t cl asses o f
ver teb rates w h ich w er e an ces tral to the mammal
,
.

G . SUMMAR Y OF TH E CI RC ULATOR Y SYSTE M

1 . T he y sys tem is d erived from the meso d erm
en tir e circul a to r .

2 . T he r s t b l oo d ves s el s are the vitell in e ( omp h al omesen teric) vein s Th es e COUTS C al on g

.

the in tes tin e an d ar e c on tin u ed po s terio rl y as the su b in t es tin al vein I n forms w ith yol k
.

s ac s th ey ar e the vein s o f the y ol k s ac .

3 As the w all s o f the h y p omer e fus e on the ven tral s id e o f the emb ry o the two vitell in e
.

vein s u n ite to form the h eart The h ear t lies in the median ven tral p ar t o f the b ody in cl osed
.

in the ven tr al mes en tery .

4 . T h e a n t e r io r e n d o f th e h e a r t co n t i n u es fo r w a r d a s t h e v e n t r a l a or t a .

5 . T h e v en t ra l a o r ta for m s a s e r ies o f l o o p s t h e aor t

,
ic a r ch es a r o u n d the p h aryn x
, T h e s e .

u n ite d o r sall y t o fo rm the d o rs al ao r tae a t rs t d ou b l e b u t s u b s equ en tl y fu s in g to a s in g l e

, ,

ves sel In typ ical ver teb rates th ere are six p airs of aor tic arch es
. .

6 The do rs al ao r ta p roc eed s po s ter io rl y al on g the mid d o rsal l in e o f the co el om s u pp l y

.
-
,

in g b r an ch es to al l p ar ts o f the b o d y b el o w the h ear t .

7 . T h e c h ie f s o m a t ic v e i ns a t r s t ar e t h e a n t er io r a n d p o s te rio r ca r d in a l v e in s u n itin g at

the level o f t he hear t to a co mmon car d in al vein on eac h s id e w h ich en ter s the s in us veno s u s ;
the su b car d in al veins ex ten d in g al on g the kid n ey s ; an d the vein of the l a teral b od y w all the ,

a b d o min a l or u mb ilica l vein o pen in g in to the co mmon car d in al

, .

8 B o th a r teries an d vein s ar e p rovid ed w ith p aired s eg men ta l an d u n p air ed n on s eg men ta l

.
-

b ran ch es The fo rmer are of two kin ds : the somatic vess el s to the p rodu cts o f the ep imer e
.
,

an d the l a t er al vis ceral b ran ch es to the p ro d u c ts o f the mes omer e T he u n p a ir ed b ran ch es

.

con s is t o f vis cer al vessel s to the d ig es tive tu b e T he ves s el s of the app e d ag es s u b cl avian
n

.

to the an ter ior app en d ag es iliac to the p o s terio r ap p e d ag es

,
n c o n s is t s imp l y o f en l ar g ed

so matic ves s els .

9 . I n a ll v e r t e b r a t es t h e v i t ell in e a n d s u b in t e s t in a l v ei n s b eco m e c o n v er t e d in t o t h e h e p a t ic

p o r t al s y s t em as e ,
x p l a in e d in th e in t ro d u c t io n o f t h is se c ti o n T h e p r o.x im a l p o r ti o n s b et w ee n

the l iver an d the h ea r t fo r m the h ep a tic vein s ; w ith in the l iver a n etw o rk o f c ap ill a ries is

p r o d u c e d ; p o s t er i o r t o t h e l iv e r o n e o f th e v ite ll in es w it h th e s u b in tes t in a l b e co m e s th e h ep a t i c

po r tal vein .

10 T he r en al po r tal s ys tem d evel o p s fro m the po s terio r p a r ts o f the po s terio r ca rd in a l

.

vein s w hich detach from the an terior p or tion s an d ar e th en kn o wn as the ren al po r tal vein s .

They co ll ec t fro m the tail an d pou r th eir b loo d in to the kid n eys fro m wh ich the bloo d p ass es in to
 COMP ARAT IVE ANATO M Y OF THE CIR CUL ATORY SYSTE M 2 71

the su b car d in al vein s wh ich es tab l ish con n ec tion s with the an terio r p or tion s o f the po s terio r
ca r d in al s .

1 1 .T he ven ou s s y s tem of sh es is in es sen tiall y the s tag e r ea ch ed to th is p o in t w ith a ,

ren al po r tal an d a h ep a tic p or tal s y s tem an d the ca r d in al vein s as the c h ief so matic vein s
, In .

shes th ere is al so an in ferior j u gul ar vein in the h ead region .

1 2 .T he ar t erial s ys tem o f sh es is al so in a p rimitive con d ition Th ere is a ven tral aor ta .

c o n n ec t e d by ao r tic a r ch es w ith a d o r s al ao r ta w h ich s u pp l ies the b o d y b el o w the h ear t Of .

the o rig in al s ix p a ir s o f aor tic ar ch es the rs t is miss in g the secon d im p er fec t an d on l y the , ,

l ast fo u r are retain ed en tire I n sh es the cen tral p art ofeach aortic arch is secon d aril y b roken
.

u p in to cap ill a ries in the g il l r eg ion ; the ven tr al h alves o f the arch es th en b ecome the a er en t

b ran ch ial ar teries the do rs al h alves the effer en t b ran ch ial ar teries
, .

13 T he p r imitive ver teb ra te h ear t is a b en t tu b e con s is tin g of fou r ch amb er s n amed

.
,

from the p o s terio r en d fo rw ar d : s in u s ven osus au ricl e ven tricl e an d con u s ar teriosu s Th er e
, , , .

is a s in g l e cir cu l a tion co mp osed of ven ou s b l oo d th rou g h the h ear t ; the b l o o d en ter s the
, ,

s in u s a n d p asses th rou g h the ch amb ers o f the h ear t in the o r d er n amed Th is con d ition is .

p erman en t in s h es ( ex cep t Dipn oi) .

1 4 T he an ter io r car d in al vein s p ers is t in all ver teb r a tes as the in tern al j u g ul ar vein s
. .

T h ese u n ite w ith o th er vein s of the an terior p ar t o f the b ody to form l ar g e tru n ks the p recaval ,

vein s T he b ases o f the p recaval s are the co mmon car d in al vein s I n some mammal s the
. .

l ef t pr ecaval j oin s the rig h t p r ecaval in fron t of the h eart formin g a s in gl e tru nk I n su ch , .

c as es the red u c ed b as e o f the l ef t p recaval ( l ef t co mm on car d in al ) r em ain s as the co r on ary

s in u s a vein o f the h ear t w all

, .

1 5 T he an t er io r p o r tion s o f the p o s terior car d in al vein s l ose th eir impo r t an ce in l an d

.

v er teb ra tes an d d is app ea r ab ove u ro d el es ex cep t th at in mammal s the p r ox im al p o r tion o f

,

the rig h t on e co n t rib u t es t o the a zyg os The p o s terio r p o r tion s of the po s terior car d in al s
.

b ecome the ren al p or tal vein s in all ver teb rates In s h es they coll ect onl y from the tail
.
,

w h il e ab ove sh es th ey al so u su rp the vein s from the l eg wh ich at rs t op en in to the a b domin al

vein s T he ren al por tal vein s in sh es Amphibia an d r ep til es ow in to a cap ill ary sys tem in
.
, ,

the kid n ey s from w h ic h the b l oo d is r e coll ec ted in to the su bc ar d in a l vein s wh ich ru n b etw een
,
-

the two kid n ey s .

16 .S imu l tan eou sly w ith th ese ch an g es in the posterior card in al s th ere app ears a n ew
vein the p os tcaval vein I t is fou n d in a few sh es an d is p res en t con tin u ou sly from the
, .

A mphibia on w ard I t is for med by the u n ion of a h ep a tic vein in the l iver ( wh ich is in tu rn
.

d erived from the vitell in es) with the su b car din al vein s ch iey the righ t su b cardin al T he ,
.

su b car d in al s th en b ecom e p ar ts o f the p o s tcava l vein .

1 7 S imu l tan eou s l y w ith th es e c h an g es the ab d o m in al vein s

. w h ich b ecome con n ec ted ,

with the ren al po r tal vein s w h en the l atter u su rp the vein s from the l eg s ch an g e th eir an terior ,

con n ec tion s Wh er eas th ey or ig in all y en tered the common car din al vein s th ey n ow en ter
.
,

the h ep a tic p o r ta l vein The a b d o min al vein s th u s co me to con s titu te a con n ec tion b etw een
.

the ren a l p o r tal a n d h ep a tic p or tal s y s tems .

18 I n A mphibia a n d r ep til es the p os tcaval vein ex ten d s o n l y to the p os terior en d o f the

.

kid n eys .

19 .I n b ir d s the p os tcava l vein es tab l is h es dir ec t co n n ec tio n s w ith the r en a l po r tal vein s ;
t he ren al p o r ta l c ir cu l a tion is th u s g r ea tly r ed u ced I n mammal s the con n ec tio n is co mp l eted
.

so th a t the vein s fro m the l eg s ta il a n d a dj acen t r eg ion s p a ss d ir ec tl y in to the po s tcava l

, ,
.

T he ren al p o r ta l c ir cu l a tion th en van ish es .

20 W ith the ch an g es ou tl in ed in p arag rap h 1 8 the ab d omin al vein l oses its fu n c tion I t
. .

is p r o b a b l y p r es en t in b ir d s b u t has es ta b l is h ed d ifferen t con n ec tion s an d is l a ckin g in mammals

ex c ep t mo n o tr emes I n emb ry on ic s tag es h o w ever th is vein ( o r vein s ) is o f g r ea t imp o r ta n ce
.
, ,

as the vein s o f the a ll an t o is o r em b r y o n ic r esp ira t o ry o r g an .

2 72 LAB ORATORY MANUAL FOR VERTE B RATE ANATOMY
Q
h epatic po rt al sys tem remain s u n ch an ged th rou gh ou t
T he .

2 2 T he ven tr al ao r ta p ers is ts as a s in gl e vess el in s h es an d Amphibia

. I n r ep til es it
.

sp lits in to th r ee tru n ks rig h t an d l ef t aor t ae an d p u l mo n ary ; in b ir d s an d mammal s in to t wo

,

trun ks the aor ta an d the p u lmon ary

,
.

23 . The aor tic arch es are mo r e o r l es s mo d i ed in l an d ver teb r a tes . I n all o f thes e the
rs t two have van ish ed ; an d in all b u t c er tain Amphibia the f th has l ikewise disap p ear ed .

I n th ese Amphibia ( u ro d el es an d Gymn ophion a ) the th ree ( or fou r) r emainin g ar ch es s till

form comp l ete ar ch es con n ec tin g w ith the d orsal aor ta I n An u r a r ep til es b ird s an d
, .
, , ,

mammal s the th ir d an d s ix th ar ch es los e th eir co n n ec tion w ith the d orsal ao r ta o n l y the fou r th
,

ar ch es r etain in g th is con n ec tion . T he th ir d arch es p er s is t as the b as es o f the ca ro tid arteries ,

the fou r th a r ch es form the aor ta an d the b as es o f the s ix th ar ch es b ecome the righ t an d l eft
,

p u l m o n ary a r t er ies I
. n A n u r a an d r ep til es b o th fou r th ar ches p ers is t formin g righ t an d
,

l eft aor tic ar ch es w hich u n ite d ors al l y to p rod u ce the dors al aor ta ; b u t in b ird s the l ef t fou r th
ar c h an d in m ammal s the rig h t fou r th arc h d is app ear . T he righ t fou r th in b ir d s an d the l ef t
fou r th in mammal s th en remain as a s in gl e ar ch con n ec tin g the d orsal aor ta with the l ef t
ven tricl e The dorsal aorta in all ver teb rates is the main ar tery of the b ody pos terior to the
.

h eart .

2 4 T he p u l mon ary vein s ap pear a s n ew s t ru c tu res in the air b r ea th in g ver teb r a tes
.
- Th ey .

en ter the l ef t au ricl e .S imu l tan eou s ly th ere o c cu rs a ch an g e in the h ear t an d the d ou b le circu
l ation is in itiated .

2 5 I n Amphibi a the f ou r ch amb ers o f the h ear t are r etain ed as in s h es b u t the au ricl e
.
,

is p ar tially o r compl etel y divid ed in to rig h t an d l eft au ricl es . T he s in u s ven osu s is th en

a ttach ed to the rig h t au ric l e w h il e the p u l mo n ary ve in s en ter the l e f t au r ic l e
,
T he r ig h t s id e
.

o f the h eart con se qu en tl y c on t ain s ven ous b l oo d an d the l e f t s id e ar ter ial b l o o d Ther e is th en
.

a d ou b l e cir cu l a tion th r ou g h the h ear t b u t the two kin ds o f b l oo d are imp er f ec tl y sep ar a ted
, .

26 . Ab ove Amphibia the con u s a r terio su s is red u ced to val ves ( the s em il u n ar valves ) a t
the b ases o f the g r eat ar teries w h ich th ereu p on s p rin g d irec tl y f r om the ven tr icl es .

2 7 I n r ep til es the s in u s ven o s u s is r etain ed the two a u ricl es a r e co mp l et el y s ep ara t ed

. , ,

the ven tricl e is u su all y in comp l etely sep ar a ted in to rig h t an d l ef t ch am b ers T he d ou b l e
.

cir cu l at ion is p res en t b u t imp er f ec t as in Amphibia .

28 . I n b ir d s an d m ammal s the s in u s ven o sus is r ed u ced to a mer e n od e in the w all o f the

rig h t au ricl e the s y s t emic vein s th en en terin g the rig h t au r icl e d irec tl y
,
T he ven tricl e is
.

comp l et el y d ivid ed by a p ar tition in to rig h t an d l ef t ven t ricl es ; con se qu en tl y the two cir cu l a

tion s th rou g h the h ear t ar e w h o ll y s ep ara ted . T he r ig h t s id e of the h ea r t is ven ou s the l ef t ,

ar ter ial .
 2 74 LAB ORATORY MANUAL FOR VERTEB RATE ANATO M Y

fu n c tion al c as es T he gl o meru li Of the p ron ep hros are ge er all y o f the t yp e kn o wn as exter n a l

.

gl o m eru li t ha t is th ey p ro j ec t in to the co el o m a dj acen t to the n ep hro s to m e an d are n o t dir ec tl y

, ,

as s o cia ted w it h the tu b u l e ( Fig I n s o me c a s es h o w ever in ter n a l g l o m eru l i w hic h p r o j ec t

.
, ,

in to the tu b u l e o ccu r .

D ia dev elopmen t o f the thr ee kidn eys an d th eir ducts a d their

how
FI G 6 0
. .
gram s t o s the n

r el tio n t o the m al e g o n ad
a A ea rly s ta ge sho win g the p ro n eph ros a d ev el op in g fro m the a t erio r en d
.
,
n

o f t he m es om e e c a n d t he p r on ep h ric d u c t b wh
r i ch ha s n o t yet r ea ch ed the cloaca e B n e t s t a g e .
,
x ,

ill u s t ra t in g the d eg e eration o f the p ro n ephr os a t f the d evel op men t o f the m eso n ephros h fr om the
n ,

middl e p o r tion o f the m esom er e the j u n c tion o f the p ro n eph ric d u c t n o w the m eson ep hric d u c t g
, , ,

with the cloaca an d the begin nin g ofthe meta n ephric eva gin ation i from the meson ephric du ct C l a ter .
,

s t ag e sho win g co nn ec tion between c er tain t u bu l es o f the m eso n ep hro s a n d the t es tis j by m ean s o f
,

t u bu l es the vasa e er en tia p whi ch g ro w ou t f ro m t he meson ephro s ; a n d the p en etration o f t he m eta

, ,

n ep h ric ev gin a t io n in to the p o s terior en d o f the m es omer e wh ere it is s u bdividin g t o fo rm the coll ectin g
a

a p p a r atu s l which becom es a s so c ia ted with s ec r eto ry m etan ep hr ic tu bu l es m d evel op ed in the meso
, ,

m er e D n al s ta g e in whi ch the meso n ephro s has d isappear ed excep t for the r emnan t g which co n
.
, ,

n ec ts with the t es t is j by m ean s o f the va sa eer en tia p; the m es o n eph ric d u c t g p ers is ts as the va s

d efer en s ; the t wo p ar ts o f the metan ephros shown in C have united to form a sin gle organ r a p o r

p ro n ephr ic d u c t ; c meso m
.
,

n ep h ro s ; b ,
er e o r n ep hr o tom e ; d in t es tin e ; e c l o aca ; f d eg en era tin g
, , , ,

p r on e p h r o s ; g m es o n ep h ric o r V
, Vol an d u ct ; h m es o n ep h r os o r W o lfan bo dy ; i m
,
eta n ep hr ic ev a g i a ,
n

t io n f ro m the W o lf n d u c t in B u ret er in C an d D ; j t es tis ; k coil ed p o r tio n o f the va s d fer en s fo rm

a
, , ,
e

in g p a rt o f t he ep ididym is ; l coll ec tin g p ar t o f the metan ephr os d eriv ed fro m the W olian d u c t ; m
, ,

ex ecreto ry t u bul es o f t he m et an ep hr o s d erived f rom the m esom er e ; n n eph r o s tom e; o r en al c o rp u s c l e , ,

o r M l p ighian body ; p vasa eer en tia; q r emn an t o f the meso n ephr o s fo rmin g p a rt o f the epididymis ;
a , , ,

r, m etan ephr os .

b) Mes on ephros : T he s ec o n d ver teb r ate kidn eys ar e the p air e d mes on ephr oi ( sin gu l ar ,
meso n ep hr os a so l kn o wn as t he W ol f/
i a n b od ies a ft er t he e mb O l O Q iS t W o l ff w ho r s t
 COMPARATI VE ANATO M Y OF THE URO GENITAL SYSTE M 2 75

d es cribed th em These develop in the meso mere immediatel y p os terior to the p ron ep hr oi even
.
,

o ver l app in g t he p o s ter ior en d s o f t he p r on ep h r o i They ex ten d over a n um ber of s egmen ts .

,

an d ar e mu c h l arg er t h an the p r o n ep hr oi As t h ey d evel op the p ro n ephr oi g en erall y d eg en

.
,

era te . T he meso n ep hr o s cons is ts Of man y tu b u l es a n u mber to eac h segmen t ; t hes e l ike the , ,

p r o n ep h r ic t u b ul es o p en a t o n e en d in to the coel om by n ep hros tomes an d at the o ther en d

,

t ermin a te in the p ron ep hr ic d uc t ( Fig 6 0 B ) Al thou gh the p ron ephr oi vanis h the p ron ep hric
. .
,

d u c ts p ers is t ; t hey gr o w p os teriorl y al on gsid e the region w here the mes on ep hr oi are develop in g ,

a n d b ecome t he d uc ts of the l a tter They are then kn o wn as the mes on ephric or Wolia n du cts
. .

T he meson ep h ric d u cts gro w p os terior l y un til t h ey r each the c au d al en d o f the in tes tin e .

T o t h is t h ey b eco me conn ec ted p ou rin g t h eir con ten ts in to the t er min al c h amb er o f the
,

in tes tin e w hic h con sequ en tl y r eceives t he n ame o f cl oac a (L atin

, a The s tru ctu r e ,

neura lt b
u e

ep im ere

mesonep
tub u e
hl ir c

Bo man sw
caps u e l
in ternal
pronep h ic
r
bl
tu u e

ephric

h
nep rosto me tome

extern a l
gl m l
o eru us

ME SONEP nR os
P RONE PHROS
'

i
intes t n e

FI G 6 1
. ,
D ia gr am
how the struc ture o f the p ron ep hros an d meson ephros P ron ephros o n the
to s .

right m eso n eph r os o n t he l eft

,
T he chi ef differen ce is in the rel a tion o f the glo meru l u s ; in t he p r o
.

n ep h o s it p r o j ec ts in to the co el om ; in the meso n ep hr os it p ro j ec ts in to the t u bul e wh

r ich forms a ,

B owm an s c ap su l e abou t it

( F r om W ie d ers heim s C om p ar a
.tive A n atom y of V e r tebr at es court es y

o f ,

the M acmill an Comp an y ) .

of the meson ephr os differs fr om t h at of the p ron ep hr os Th e tu b u l es are more n um erou s , .

l on ger , an d mor e co nvolu ted The gl omeru li are al ways of the in ter n al kin d ( Fig E ach
'

. .

glomerul u s p r oj ec ts in to the meson ep hr os , carr yin g the w all of the tu b u l e b efor e it to form a
ask s h ap ed c ap s ul e in w hic h t he g l o mer ul u s res ts The cap su l e is n amed B owman s ca ps ul e,

- .

an d th is , t o g eth er wit h the g l o meru l us , f orms a r o u n d ed b o d y kn o wn as the Mal pighian b ody

o r r en al cor puscl e T he meson ep hr ic tu b ul es g en er al l y o p en in to the coel o m by n ep hros to mes

.

whic h d n ear the r en al corp uscl es

are situ ate .

T he m eso n ep hr o i ar e the kid n eys of the ad ul ts of s h es an d Amphibia an d ar e al s o the ,

c hief f un c tio nal kidn eys o f cy cl os to mes al th ou g h the l atter p o s s ess in ad dition p ers is ten t
,

o n e h r o i T he m e s on eph ric o r Woli an d u c ts ar e the ex cr etory d u c ts in t h es e ver teb r a tes

p r p .
,

o p enin g in to the c l oac a The meso n ep hroi ap p ear in t he emb ryos o f rep til es b ir ds a n d
.
, ,

mammals an d are in mos t c as es fun c tion al in the emb ryos o f ten b eco min g o f l ar g e s iz e
, ,
.

They gen erall y however l ack n ep hro stomes Th ey d egen era te b efore h atchin g o r b ir th or
, ,
.
,

very s hor tl y af ter .

2 76 LAB ORATORY MANUAL FOR VERTE B RATE ANATO MY
o
)
c

The metanephroi: T he t h ir d kidn eys of vertebrates ar e the p air e d meta nephr oi T he .

metan ep h ro i from the remain d er o f the meso mer e p os terior

ar is e in p ar t , to the meso n ep hroi ;
an d in p ar t as an o u t gr o w t h fro m the W o lfan d uc ts This ou tgr o w th o ccu rs as an evagin ation .

from the W o li an d u ct n ear its co n n ec tio n with the cl oaca ; it p us hes an teriorl y an d fus es
with th at p or tion of the metan ep hr os arisin g from the meso mer e ( Fig 6 0 B an d C) T he . .

metan ep hro s is thus an o r g an of dou bl e or igin That p ar t w hich comes fro m the mes o mere .

d evel op s in to the tu b u l es of the kid n ey an d the ren al corp uscles ; th at p ar t derived from the
Wo li an d u ct beco mes the coll ec tin g tu b ul es an d the sp aces ( cal ices ) in to w h ich they emp ty .

T he s talk of the ou tgro w th from the W olian d u c t b eco mes the meta n ephr ic du ct o r ur eter ,
x

wh ich conveys the excretion of the metan ep hros to the cl o aca T he s tru ctur e of the meta .

n ep hr os is s imil ar to t h a t o f the meso n ep hr o s excep t that n ep hr os to mes ar e w an tin g , The .

met an ep hric tu b u l es ar e ver y lon g an d comp lica ted with s everal l oo p s an d con vol u tion s The , .

metan ep hr o i ar e the kidn eys o f the a d ul ts of all r ep til es b ir ds an d mammal s , , .

From the foregoin g accoun t it will b e seen th at the thr ee p airs of ver teb rate kidn eys aris e
s u cces s ivel y fro m t he mes om ere eac h s itu ated cau d ad to the p r eced in g on e
, For t his reas o n .

it s eems p ro b ab l e t h at t h ey r ep resen t p ersis ting p ar ts of a kidn ey w hich or ig in all y ex ten d ed

the en tir e l en g th o f the meso mer e The an terior p art of this h yp oth etical kid n ey has gra du all y
.

d egener a ted so th at the kidn ey ap p ears to descen d poster iorly This d isapp earan ce of o rgans .

fro m an terio r segmen ts is co mmon in heteronomousl y s egmen ted animals .

3 The ur in ar y b l a dd er
.
The u r in ary bl adder is in all forms above s hes a s aclike evagin a
.

tion fr o m the ven tral w al l o f the cl o aca In s h es the b l add er when p resen t is formed in p ar t
. ,

by the en lar gemen t of the termin al p or tions ofthe W o lian du cts as well as by a clo acal evagin a
tion . In the emb ryos of rep til es b irds an d mamm als th ere is an en orm ous cl oacal
, , ,

evagin atio n the all an tois w h ich m ay b e reg ar d ed as a gr eatl y exp an d ed u rin ary b l add er
, ,
.

The a ll an to is s erves resp ira tory as w ell as u rin ary f u n ction s The a d ul t b l add er d evel ops a t .

the b as e of the a ll an toic s talk Ad ul t b ird s l ack a urin ary b l add er

. T he ex cr etory d uc ts .

do n ot g en er al l y o p en in to the urin ary b l add er dir ec tl y ex cep t in mammals

.
,

4 The g on d
. a s an d their d u cts The g on ad s or r ep ro d u c tive o r g an s co ns is t o f a p air o f
.

tes tes in the mal e ovar ies in the femal e

,
Th es e org ans aris e as swell in gs the gen ital r idges
.
, ,

o n the ven tral w all o f the meso mer e imm edia tel y to the medial s id e o f the kid neys
,
They .

p r oj ec t in to the co el o m an d in the ma tur e con dition ar e g en erall y p r ovid ed with mesen teries .

a) M al e: T he t es tis co n s is ts of a mas s of tu b u l es the s emin ifer ous tu b ul es whic h emp ty , ,

in to a n etwork o f tu b u l es cal l ed the r ete whi ch is situ ated us u all y al on g the a ttach ed p ar t o f
,

the tes tis Tu b ul ar con n ec tions are es tab l is hed d urin g emb ryonic s tages b etween the rete
.

an d t he B o w man s c ap s u l es o f the an ter io r p ar t of the a dj acen t meson ep hr os ( Fig 6 0 C)

. .

These conn ec tions ar e call ed the vas a eeren tia an d they travers e the mesorchiu m in to t he ,

meson ep hr o s T he vasa eer en tia ar e th en con tin uous within the meso n ep h ro s with c er tain
.

of the meson ep h ric tu b u l es an d thr o u g h t h em n d exit in to the W oli an d u c t I t th us h app en s .

t h at in n ear l y al l ver teb ra tes the W o li an d u ct s erves as the mal e d u ct I n el as mo b r an chs an d .

Amphib ia in whic h the mes on ep hros is the fun c tion al ad ul t kidn ey the W o li an d u c t in males
, ,

serves as b o th ex cretory a n d mal e d u c t ( Fig I n mal e rep til es b ir ds an d mamma ls

.
, , ,

w here the meson ep h ro s has dis ap p eared the W olian du ct remains as the mal e duct an d is , ,

then n am ed the vas d efer ens ( Fig 6 2 C) I n th ese grou p s the meson ep h ros disap p ears excep t
. .
,

th o se meson ep hric tu b ul es whi c h h ave es ta b l is h ed co nn ec t io ns with the rete tes tis Thes e .

tu b ul es co ns titu te the vas a eer en t ia a n d a p ar t o f t he epididymis Ep idid ymis is the n ame .

g iven to th at p ar t o f the mal e d u c t w h ic h is co nvo lu ted .

A gr ea t deal o f confu sion will be a vo i ed d by re se r vin g the term ur eter for the d uct o f the meta
n ep hr o s .
 2 78 LAB ORATORY MANUAL FOR VERTE B R ATE ANATO M Y

b) F emal e: T he o v arl es ar e m asses o f c onn ec ti ve ti ssu e co n tain in g the d evel o p mg ggs

e ,

eac h egg surro un d e ll s f ormin g a f olli cl e The ovar ies unl ike the
d by a c a p s ul e o f n u triti e c e v .
,

t es tes n ever h ave an y co nn ection with t he kid n ey s

,
T he d u cts o f the ovaries ar e n amed the .

oviducts o r M ul l er ia n d u c ts T he or igin o f t he Mull erian d u c ts is s o mewh at p r ob l ema tic al

. .

I n el as mo b ran c hs th ey ar ise by a sp l it tin g O f the p ro n ep hr ic d uc t ; h alf o f t he p ron ep hr ic

du c t th en b ecomes the ovidu ct an d the o th er h alf b ecomes the meson ep h ric d u ct Al , .

t h ou g h t his mo d e o f o rigin is the on e co mm o n l y a cc ep ted it c a nn o t b e d emon s tra ted fo r ,

o t h er ver t eb r a t es ; in t h em t he o vid u c ts a r ise in d ep en d en t l y in t he m es o mer e T he ovid u c ts .

ar e n ever d irec tl y co nn ec ted w ith t he ovar ies Th ey o p en in to the c oelom n ear the ovaries .

u terus

u terin e tu e b

g
u ro eni tal s in u s

M a me tu b
e
i
u ter n e tu eb

h o rn o f u

b dyo of

FI G 6 3 .
D iagr ams
. y to s how lian o vidu cts A co n dition foun d in
the va riou s t p es o f mamm a .
,

the m aj ority o f f em al e v er t eb ra t es ; t he t wo o v id u c t s ar e co mpl etel y sep ar at e a n d o p en in d ep en d en tl y

i to the cl o a ca
n B E variou s c o n dition s f ou n d in m amm al s sho win g differen tia tion o f the o vid u c t s
.
-
, ,

in to u t erin e t u be u teru s a n d v a gina an d p ro gr essive fu sion o fthe l o wer p ar ts o fthe o viducts : B du pl ex

, , , ,

typ e f ou n d in roden ts in which the t wo v a gina e a r e u nited t o o n e ; C b ipa tite t yp e occu rrin g in c a rn i
, ,
r

vo r es ; n ot o n ly a re the va gin ae fu sed b u t the l o wer p a r ts o f the t wo u teri a re fu s ed t o form a s in gl e

b ody divid ed in t wo by a p ar tition which rep resen t s the fu sed walls o f the t wo u t eri; the u pp er p arts o f
,

the t wo u t eri r em ain s ep ara te a s the horn s ; D b icor n ua te typ e fo u n d in m an y u n gul at es simila r t o C , , ,

e c ep t th at the p art itio n ha s d isapp ea r ed ; E

x s implex t yp e oc c u r rin g in ma n an d the ap es in which
, , ,

bo th vagin ae an d u teri ar e fu sed al on g th eir en tir e l en gths l ea vin g o n ly the u terin e t u bes s ep arat e .

No e fu h
t r t er t h a t in B D the u rethr a j o in s the v a gin a t o form the u ro g enital sin u s which o p en s t o
the e t erio r whil e in E the u r ethra an d va gin a ar e wholly s ep ara te a n d o p en in d epen d en tly t o the
x ,

( F rom W ied ersheim s Compar ative An a tomy of Ver tebr ates c ourt esy of the M acmill an

e t erio r
x .
,

Comp an y ) .

by a fun n el s h ap ed op enin g the os tiu m which p rob ab l y rep r esen ts on e o r mor e of the n ep hr o
-
, ,

s t o m es o f the p r o n ep hr os ( Fig O2 B an d D) T he eggs es cap e fr o m the ovar y by ru p tur e o f

. .

the ova rian w a ll p as s in to the o s tiu m o f the o vid u c ts by meth o ds w hi c h ar e n o t a l wa ys un d er

,

s to od an d ar e con veye d d o wn the ovid u c ts

,
.

T he ovid u c ts in the maj or ity o f ver teb r a tes r emain a s two s ep ar ate tu b es o p en in g in to
t he c lo aca ( F ig 6 3 A) I n mamm al s ea ch ovid u c t is d iff eren tia ted in to a n arr o w er an terio r
. .

o r t ion c all e d t he u ter in e o r F al l opi an tu b e w hic h bears the o s tium an d a wid er mor e m us cul ar
p , ,

p o s t e r io r p o r tio n t h e u terus I n
,
t he m on o t r e m es or e gg.l a yin g m amm al s e a c h u t erus o p en s ,
-
,

s ep ara tel y in to the c l oaca I n the m ar su p ial s the termin al p or tio n of the u teru s is dier en tia ted
.
 C OMPARATIVE ANATO M Y OF THE URO G ENITAL SYSTE M 2 79

as a vagina to recei e v the p en is . b ove marsu p ials the two vagin ae fus e to a
I n m ammal s a

gle vagin a ( h en ce
s in the n ame M on odel phia) Ther e is al so gen er all y mor e o r l ess f us ion o f
.
'

the two u ter i ( Fig . Wh en o nl y t he p os ter ior p o r tio n s o f u ter i ar e f u s ed the f u sed
p or tion ,

is call ed the b ody of the u teru s an d the s ep ar ate p o r tion s the hor n s o f the u teru s In man an d .

o t h er p rim ates t he u teri ar e f u s ed a l o n g t h eir en tire l en g t h p ro d u cin g the s in gl e u ter u s o r

womb T he youn g o f the p l acen tal mammal s d evel op o n l y in the u terin e p ar t of the ovid u cts ;
.

in th os e for ms with p ar tial l y f u sed u ter i on l y in the hor n s ,

.

5 The e ol u tion of the c l oa c

. v a The clo aca is f oun d in al l ver teb ra tes ex cep t c ycl os to mes
.
,

t el eos tomes an d the p l ac en ta l m ammal s

, I t r ec eives the ter min ation o f the in t es tin e an d the
.

ur in ar y an d g en ital d u c ts From the p reced in g accoun t it will b e eviden t tha t in the mal es
.

o f el as mo b r an ch s an d Amphibia t he cl o ac a receives o nl y the W o li an d u c t s w h il e in the ,

femal es b o th ovidu cts an d W olian d u c ts en ter it ( Fig 6 2 A an d B ) I t co mmon l y h ap p en s . .

,

ho wever t h at in the males o f th es e group s ves tiges o f the ovid u c ts ar e p r es en t

, I n the mal es .

bl dd
a er

ureter
ov id
u ct

g i
ur o en tal
i
S n us

FI G 6 4 D iagrams to ill ustrate the cha n ges in the cloaca in mammals du rin g d evel opm en t
. . .

A earl y embryon ic sta ge , showin g the clo a c a r eceivin g the u rin ary bladd er , the r ec tu m , a n d the W olf an
duct as in the lower vertebr ates B l ater s tage showin g the begin nin g o f the fold whi ch divides the
,
.
, ,

cl oaca in to a ven t ral u ro g en it al s in u s whi ch r eceives the u rin ary bla dd er W ol fa n d u c ts an d u r et ers , , ,

an d in to a d o rs a l p art which rec eiv es the r ec tu m C fu r th er

w g ress o f t he fol d d ividin g t he cl o aca
.
, ,

in to u rog enital sin u s an d rectum ; the u ret er has s ep a ra ted from the W olian du c t an d is shiftin g
an terio rly D completion of the fold showin g complete s eparation o f the cloaca in to ven tral u ro
.
, ,

gen ital sin u s an d dorsal rectum Not e in D that the ureter has shifted farther so that it op en s in to the
'
.

u rin ary bl add er .

l
o f rep ti es an d b ir ds the cl oaca receives the Wolfan du cts (vas a deferen tia) an d the ureter s ;
in the f emal es the ovidu cts an d the ur eter s ( Fig 6 2 C an d D) I n a d di tion in man y s h es . .
, ,

Amphibia r ep til es an d the emb ryos o f b ir ds a n d mammals the u rin ary b l add er op en s in to
, ,

the ven tr al w all o f the cl o a c a Ad ul t b irds h ave n o urin ary b l add er ; mammal s h ave on e
.
,

b u t it is n o l o n g er attac h ed to the d ig es tive trac t .

I n p l acen ta l mamm al s m ar ked c h an g es occu r in the r el atio ns of the termin al p or tion s o f

the ur o g en ital d u c ts I n the emb ryo the cl o aca b ecomes divid ed by a fo l d w hic h ex ten d s
.

p os t erior l y to t h e b o d y w all an d s ep a r a tes t he c l oa c a in to t w o p ar ts e ac h w it h its o wn o p en in g ,

to the ex terior ( Fig T he d o r s al p ar t in cl u d es the in tes tin e o n l y ; t h is termin al p or tion

.

o f the in tes tin e is c a ll ed the r ec tu m a n d o p ens to t he ex terio r by the an u s T he ven tr al p ar t .

s ep ar a ted f ro m the cl oaca is c all ed the u r ogen ital s in us I t r eceives the s tal k of the b l add er .

an d the ex cr e to r y an d g en ital d u c ts T he excr etory d u cts ( eith er W o lian d uc ts o r ureter s) a t

.

r s t op en in to the uro gen ital sin us b u t su b sequen tly the u reters shif t s o as to op en in to the b l a d
,

der in all o f the p l acen tal mammal s ( Fig

,
Thu s the ur eters p ass in to the bl add er w hil e the
.
,

Wolian du cts (vas a d eferen tia) in males or the vagin a in femal es un ite with the d u c t o f the
b l adder n amed the u r ethr a formin g a co mmon tu b e o r c hamb er the u rog en ital s in u s wh ic h
, , , ,

o p en s ex tern all y in fr o n t o f the an us by a urog en ital ap er ture I n the femal es o n ly o f the .

2 80 LAB ORATORY MANUAL FOR VERTE B RATE ANATOMY
O
highes t p l acen tal

mamm al s t here is n o ur o gen ital sin us , b u t the h
ur et r a an d the vagin a ar e

s ep arate an d op en s ep ar ate ly to the ex ter io r , t he former an ter ior to the l atter . In su c h cases

h er e are thr ee op enin gs in the p erin eu m : the an us the mou th of the vagin a an d the mou th

t , ,

o f the ur e t hr a .

For more comp l e te a ccoun t s Of the comp ar a tive an ato my of the u rog en it al s ys tem the ,

a pp rop riate ch ap ters of K W or W d s h o u l d b e co n su l ted

, ,
.

B . T HE UR O GE NI TA L SYS TE M OF E LA S M OB RANCH S
The fo llo wi n g ou tlin e app lies to the smoo th an d spin y dog sh an d the sk a te ,

the l a tte r b ei n g de s c rib ed s ep a r a tel y The m aj o ri ty o fthe d o gs h u s ed fo r diss ec .

ti on a r e i m m a tu r e an d it is the r ef o r e dii cu l t o r imp ossi b le to l oca te in them a ll

,

o f the p a r ts o f the u r o ge n i ta l s ys t em At le a s t a fe w m a tu r e m a le s an d fem a les .

will b e o n h an d fo r dem on s tr a ti on T he s k a te s a r e s ex u al ly ma tu r e when s till .

r el a tivel y s m a ll an d m o s t o f the sp e cim e n s u s ed h a ve fu ll y devel op ed r ep r o

,

du c tive sy s tem s .

R em ove the dige s tive tr ac t ex cep t cl oaca an d liver .

1 . The fe male ur o ge n ital s ys te m .

Dog s h: T he o v arie s ar e a p a ir of s o f t b odi e s si tu a ted d o r s a ll y In the ,

.

s p i n y d ogsh the y a r e o v a l in f o rm an d l o ca ted d o r sa l to the live r e a ch w i th a ,

m e s en te ry the m e sovar i u m
,
I n the smoo th sp e cie s they a r e l on g an d s len der
.
,

ex ten di n g the who le len g th o f the co elom mo r e o r le ss fu s ed an d a tta ched , ,

p o s t e ri o r l y to the m e s en t e ry o f the r e c ta l gl an d T he ir p os te ri o r e n d s a r e .

to o thed . T he r e is si n gle m e so var iu m fo r the two o var ie s In b o th sp e cie s .

when m a tu r e the l ar ge eggs co n si s ti n g c hiey Of yo lk a r e r e a dily n o ted in the

, , ,

o v a r ie s .

L oca te the kid n eys l yi n g a gai n s t the d o r sa l b o d y wa ll on e to e ac h side o f ,

the d o r sa l ao r ta They ar e l o n g s le n der b ro wn b o dies Thei r p os ter io r po r ti on s

.
, ,
.

a r e b r o a de r a n d thi cke r t h a n the a n t e r i o r p o r ti o n s an d p r o b a b l y p e r f o rm m os t

o f the w o r k o f ex cr e ti o n The kid n eys a r e r e tr op e ri to n e a l F r ee their la ter al

. .

b o r der s by s li tti n g the p leu r op er i to n eu m an d n o te thi ckn e ss o f the o rgan a t

diffe r en t levels T he kidn eys are m eson ep h roi ; the po s ter ior thi cker p ar t may
.

b e n am ed the caud al mes on ep hr os the an te rio r mo r e s len der p ar t the cr an ial ,

mes on ephr os B e tw een the two kid n eys is a tou gh shin in g lig ame n t w hi ch
.
I

s h ou ld n o t b e mi s ta ke n for a d u c t .

T he oviducts in imma tu r e fe male s ar e s len de r tu b es ru n nin g a lo n g the ven tr a l

fac e o f the kidn eys w i th ou t m e s en te r ie s ,
I n ma tu re fem a le s the y a r e ve r y l ar ge .

tu b e s whi c h sp r in g f r ee f r om the kid n e ys by m e an s O f w ell devel op ed m e s e n te rie s -

,

the mes otu b ar ia Tr a c e the o vid u c ts forw ar d T hey p a ss fo r war d a l on g the

. .

do r s a l c o el omi c wa ll cu rve ar ou n d the an te ri o r b o r de r o f the live r an d en ter

, ,

t he fal cif o r m lig ame n t H er e the two ovidu c ts are un i ted to a common op en in g
.
,

1
T he dal meson ephro s is o ft en r egarded as a metan ephro s an d its du ct as a ur eter
ca u . P ro fe s so r

Kin g s l ey has kin dl y exp ress ed his Opin ion that this u sa ge is un j u tiable s .
 28 2 LAB ORATORY MANUAL FOR VERTEB RATE ANATO M Y
o
b egi n nin g o f w hi ch b e ar s a con sp i cu ou s b il ob ed sw ellin g the ovidu cal gl an d , ,

whi ch s e cr e te s the h o rn y c as e in whi ch the egg s a r e l aid ( N p 1 6 4 Fig , .

, .

T he u ter i p ro ceed to the clo aca su pp o r ted by the thi cken ed m eso tu b aria Cu t .

op ne the l in the m edi an ven tr a l lin e s li t ti n g op e n the i n t e s ti n e

c oa c a , No te .

the op en i n g o f the i n te s tin e i n to the ve n tr a l p ar t o f the c lo a ca an d the co n sp i cu ou s

h orizon tal fo ld whi ch s ep ar a te s this fro m the dor sal u r ogen ital p ar t Cu t in to .

the l a tte r by c u t ti n g f o r w a r d th r ou gh this fo ld T he c lo aca is g r e a tly ex ten de d

.

an d t h i cke n ed in the a n t e rio r di r e c ti on F in d the op en in g s o f the o vid u c ts

.
,

o n e to e a c h s ide o f th i s thi cke n ed p a r t o f the clo aca H alfway b e tween the

.

o p e ni n g s o f the o vid u c ts is the u r ogen ital op en in g in the m ed ian d o r s a l wa ll .

The m ai n p ar t o f the kid n e ys in fem ale sk a te s con si s ts Of a thi ck r ou n ded

lo b e lyi n g agai n s t the d o r s al wall a t e a ch side o f the c l oaca The s e l ob es are .

r eve a led by s tr ip p i n g Of f the thick p leu r op e ri to n eu m whi ch cove r s thei r ven tr a l

su r f a c e s .T h es e l ob es m ay b e n am ed the ca udal meson ep hr os The a n te ri o r .

p a r t o f th e m e so n e p h r o s o r c r an ia l mes on ep hr os is n e a r l y dege n e r a t e in fe m a le s

b u t will b e fou n d as diffu s e b r own is h ti ssu e e x ten d in g f or wa r d ven tr a l to the

o

d o r sal a o r ta F r om the m edi an su r fac e O f the cau d al m e so n ep hr os se ve r al d u c ts

.
,

the acces s ory mes on ep hr ic ducts p as s an t e r io r l y an d m edi a ll y in co n ta c t wi th

,

the p o s te r i o r ca r di n al vei n an d Op e n in to a sm a ll ch am b e r the u r in ary s in u s , ,

situ a ted o n the d o r sa l su r f a c e O f the a n te r i o r en d o f the c l oa ca T he two u ri n ary .

s i n u s e s o f the two side s u n i t e i n to a co m mon ch am b e r w h i ch is so m e ti m e s c a lled

,

the u r in a ry b l adder I t d o e s n o t co rr e sp on d to the b l a dder o f h ighe r fo rm s

.
,

sin c e it c o n sis t s o f the e n l a r ged t e rm i n a ti on s o f the m e s on ep h r i c d u c ts Cu t .

in to thi s n o te the en tran c e in to it o f the two u ri n ary s in u s e s an d n d the Op enin g

, ,

in its m id d o r s a l w a ll by whi c h it op en s i n to the cl o a ca

-
T he Wolian ducts or
.

du cts O f the cr ania l m e son ep h ros ar e s len de r tu b e s ex ten di n g an te r io r l y f ro m the

u ri n a r y b l a dde r l yi n g o n the d o r sal s u r f a c e o f the s tr o n g whi te p o r ti on s o f the
,

mes o tu b aria .

D r aw s h owin g kidn eys go n ads an d thei r du c ts an d the Op en ed cl oaca

, , , ,
.

2 . The m al e ur o ge n ital s y s te m T he tes tes a r e a p ai r of sof t b o die s d o r sall y

.

s itu a ted . I n the sp i n y do g sh they a r e l o ca ted in the an te r i o r p ar t o f the

le e i t o e a l c vi ty d l t o th e live r ; e a ch h a s a m e s o r ch i um I n
p u r o p r n a o r sa
,
.

the sm oo th dog s h the y ar e l on g an d s len de r b odi e s ex te n di n g the le n g th o f the

p le u r o p e r i t o n e a l ca vi ty thei ,
r t oo thed p os t e r i o r e n ds a t t ac hed to t h e m e s e n te r y

o f the r e c tal gl an d They are mor e o r le ss fu s ed an d are supp o r t ed by a s in gle

.

m e s o r chiu m .I n the s k a te the te s tes ar e b r o a d a t b odi e s a gain s t the d o r sal

,

wa ll ; ea ch is p r o vided wi th a m eso r chiu m .

The kidn eys ar e iden ti ca l wi th th os e Of the females an d s hou ld b e n ex t

ex ami n ed accor din g to the dir e c ti on s given un der female s In the ma le skate .
,

how eve r the cr an i al p ar t of the kidn ey is very mu ch b e tte r develop ed t h an in

,

the fe m a le an d ex t en d s fo rw ar d as a r m cy lin d r i ca l b o d y o n ei the r side o f the

mid d o r sa l lin e
-
.
 COMPARATIVE ANATO MY OF THE UR O GENITAL SYSTE M 2 83

As exp lai n ed in the in tro du c ti on the mal e ducts the m aj ori ty o f ver teb ra t es
ar e the mes on ep hr ic or Wolian du cts I n m a tu r e ma les the s e d u c ts ar e con
.

s eq u en tl y m u ch l ar ge r th an in fe m ale s T he W oli an du c ts ru n po s teri or ly .

l
a on g the ven tral f a ce o f the kid n eys In i mma tu r e sp e cim en s each is a s len de r
.
,

s tr aigh t tu b e s im il a r to th a t o f the fe m a le b u t in m a tu r e m a le s it is gr e a tl y
, , ,

co iled . The te s ti s is con n e c ted wi th the cran i al me s on ep h ro s by m ean s o f

deli ca te du c ts the vasa eeren tia whi ch ru n in the m e so r chiu m an d can some tim es
, ,

b e s een by ho ldin g the m e so r chiu m u p to the ligh t The va s a effe r en tia con n e c t .

wi th the tu b u le s Of the m e son ep h r o s The gr e a ter p a r t o f the cr an i al m eso

.

n ep h r o s app ar en t l y s e rves in m a le el a sm o b r a n c h s fo r tr an sm i ttin g the s e r m

p ,

a n d is s o m e tim e s ca lled the ep ididymis s i n ce it co rr e sp o n d s to the he a d o f the

,

ep i didymi s o f mamma l s The sp e rm b e arin g tu b u le s of the me son ep h ros then

.
-

con n e c t w i th the W o lian d u c t Tr ace thi s d u c t p os ter i or l y I ts an teri or p ar t

. .

is g r e a tl y coiled in m a tu r e m ale s b u t in dogs hes (n o t in sk a t e) s traigh te n s as

,

it app r oache s the cau d a l m e s on ep h ros an d in a l l th r ee anim a l s en l ar ge s u po n the

su r f ac e o f the l a tte r to f o r m the s emin al ves icl e Trace it by r emovin g the .

p le u r o p e r i to n e u m f r o m th e ve n tr a l f ac e o f the cau d a l m e s o n e p h r os A t its .

p o s te r i o r e n d s o n the s ide s o f the c l o aca the s e min a l ve s i cle te rm i n a te s in a s a c

, ,

the s p erm s ac whi ch p r o je c ts fo r w ar d as a b lin d s a c lyin g a ga in s t the ven tra l

,

su r f a c e o f the s emi n al ve si c le .

Cu t open the c loaca as di r e c ted u n de r the fe m ale an d ide n tify its p ar ts a s

di r e c ted ther e T he r e is n o differ en c e in the c lo aca o f the dogsh b e tween the
.

m ale s an d fe m a le s b u t in the m a le sk a te the cl oa ca is ve r y mu c h smal ler th an

,

in the fe m a le an d is n o t d ivided i n to in te s ti n al an d u r o geni ta l p ar ts In the .

m edi an d o r sal li n e the r e is in the m ale s k a te a u r ogen ita l p ap il l a .

The sp erm sa cs s h ou ld n ow b e cu t op en an d the p a pil l ae w he r e the s emi n al ,

ve si cle s op en in to them iden tied T he two sp erm s acs u nite at their p os teri or
,
.

en ds to fo rm a u rogen ital sin u s which op en s a t the tip o f the u rin ar y p apill a .

T he acce s so r y m e so n ep h r i c d u c ts a r e si mil ar to tho s e o f the fem a le s ; see the

de s crip ti on u n de r fem ale s In m ale do gshes e ach r u n s alon g the m edi a l side
.

o f the s emin a l ve s i c le an d e n t e r s the sp e r m s a c n e a r the op e n in g o f the ve s i cle .

In the m a le s k a te s eve r a l acc e sso r y d u c ts p as s f ro m the cau d a l m e son ep h r os

in to the sp e rm sa c .

D raw s h owin g gon a ds kidn eys an d thei r du c ts an d the Op en ed clo aca

, , , , .

L o o k in m a le do gs hes for r em n an ts o f the o s ti um an d o vid u c ts in the r egion

o f the live r .

C . THE UR OGE NI TAL SYS TE M OF NE CTURU S

1 . fe mal e
The ur o g e n ital s y s t e m
T he
ova r ies h ave a l r e a d y b ee n n o ted a s
.

elo n gated saclike b o die s b e a ri n g eggs of var i ou s siz e s No te the mesovar iu m . .

L a tera l to ea ch o va ry ru n n i n g a l o n g the d orsa l b o d y wa ll is the ovidu ct a thick , ,

white co iled tu b e supp o r ted by the mes o tu b a riu m Fo llo w it an ter io r l y

,
. .
2 84 LAB ORATORY MANUAL FOR VERTEB RATE ANATO MY

At the an ter io r en d o f leu r op e r i ton e al ca vity it b eco me s o fa m o r e deli ca te
the p
tex tu r e a n d is f as te n ed to the l a te r a l w a ll H er e it op en s by a fun n el shaped .
-

op e n i n g the os tiu m; the d o r sa l r im o f thi s is fas te n ed to the b o d y wa ll b u t the

, ,

ven tra l rim is fr ee an d can b e lif ted to exp o s e the op enin g Tr ac e the ovid u c ts .

p o s t e r i o r l y t o the c l o a c a T he y e n t e r th.i s o n e to e a ch s ide o f the l a r ge in t e s tin e .

Cu t the clo aca op en by a l a ter a l s li t ex te n din g u p in to the i n t e s ti n e N o te the .

p ap il l a e by whi ch the o vid uc ts op en in to the cl o aca .

The kid n eys ar e l on g s len der o rgan s ex ten d in g fr om the cl o aca fo rw a r d

a l o n g the d o r s al su r f a ce s o f the o vid u c ts The y a r e r e tr op eri to n e al They a r e
. .

mes on ep hr oi T he d u c t the W oli a n or mes on ep hr ic duct l ie s a l on g the l a ter a l

.
, ,

b or der o f e ach kid n ey I t is ve r y deli c a t e in the female an d dii cu l t to l oca te

. .

I t p ro ceed s to the c loa ca in to whi c h it op e n s to the d o r s al s ide Of the ovid u c t .

I n tr aci n g it m ake a cu t a l on g o n e side o f the c l o aca f r eein g the cl oaca fr om the ,

b o d y w a ll .

No te the u rin ar y bl a dder ex ten din g fr o m the midven tr a l r egi on o f the

c l oa c a . F in d its op enin g in to the clo a ca .

D raw s h o wi n g o varies kidn eys thei r d u c ts an d the op en ed c loaca

, , , , .

2 . The mal e u r og e n ital s ys te m The tes tes ar e a p ai r o f el o n g a ted b odie s

.

s i tu a ted to the side s o f the s m a ll i n t e s ti n e E ach has a m e s en te ry the mesor .

,

chiu m . D orsal an d l a ter a l to e ach te s tis is the lo n g b r o wn kid n ey la r ger th an ,

in the fem a le The kid n e y is a mes on ep hr os Alon g the l a te r a l b or der o f the

. .

kidn ey is a con sp i cu ou s coiled du c t the Wolian o r meson ep hric du ct T hi s a l so

, .
,

as in ve r te b r a te s in ge n e r a l ac ts as the sp e rm d u c t ,
B y h o ldi n g up the m e so r .

chiu m to the ligh t n o t e the deli c a te d u c ts the va s a e er en tia w hi c h c r o s s it in to , ,

the kidn e y The sp er m p a s s th r ou gh the tu b u le s o f the kidn ey an d in to the

.

W olian d u c t Tr a ce the l atte r to the c l o a ca Open the c l o a ca by a s li t to

. .

o n e s ide O f the m edi an lin e ca r r y in g the s li t i n to the l a r ge i n te s tin e The Op en in gs .

o f the W o lian d u c ts in to the c l oaca ar e ge n e r a ll y diicu l t to l oca t e o win g to ,

thei r s m a ll s i z e No te the u r ina ry bl adder an d its op en in g i n to the ven tr al

.

c l o aca l wa ll .

D raw s h owi n g te s te s kidn eys W o li an du c ts an d op en ed c lo aca

, , , ,
.

D . THE UR OGE NI TAL SYS TE M OF THE URTLE

T

R em o ve the dige s tive trac t , if n ot a r a l e dy d on e leavin g ,

the large in te s tin e
in p l ac e .

I . The fe mal e uro g e n ital s ys te m

i T hi s
l of a p ai r of ovaries
. c o n s s ts as u su a

a n d a p ai r o f M til l er ian ducts o r ovidu cts The o var ie s h ave a l r e ad y b ee n n o ted

.

as l a r ge b a glike b o die s in the p os te r i o r p a r t o f the p leu r op e r i to n e a l ca vi ty .

They usu a lly con tain y ell o w eggs in va r iou s s ta te s o f devel op m en t E a ch ova r y .

is s u p po r ted by a m e s en te ry the mes ovar iu m Alon g the p os te r io r b or der o f

,
.

ea ch o vary run s the oviduct a large white co iled tu b e su pp or ted by the m eso tu
, ,

b ariu m Tr ac e the o vid u c t fo r wa r d an d n d the os tiu m; thi s li es in the m e s en

.
286 LAB ORATORY MANUAL FOR VE RTE B RATE ANATOM Y

The kidn eys w er e p r evi o u s ly iden tied a s htten ed lo b ed b o die s tti n g

,

a g ai n s t the p o s t e r i o r w a ll o f the p le u r o p e r i to n e a l ca vi ty E a ch tes tis is a yello w .

s p he r i ca l b o d y a tta ched to t he ve n tr a l f ac e o f the kidn e y by the m e s o r c h iu m .

I a teral an d p os te r i o r to the te s ti s is an elo n ga ted d a r k co l or ed coiled b o dy the ,

-
,

ep ididymis The te s tis is co nn e c ted to the a n ter i o r p ar t o f the ep idi dymi s

.

by the mi n u te va s a e er en tia whic h r u n in the m eso r chi u m T he va s a effer en tia .

a n d th i s p o r ti o n o f the ep idid ym is a r e the r emn an ts of the mes on ep hros The .

r em ain de r o f the ep idi d ym i s co n s ti tu t e s the m a le d u c t o r va s deferen s ( W ol ia n

duct) . R em o ve the p e r i to n e a l co ve rin g o f the ep i di d ym i s an d u n coil the vas
defer en s Tr a c e it to the cloaca I t en ter s an ter i or to an d a t the b as e of the
. .

b ul b o f the p enis .

N ex t cu t op en the cl oa ca in s e r tin g the b l a de O f the s ci sso r s in to on e co rn er

,

of the a n u s a n d c u t tin gde to av o id in j u rin g the p eni s Sp r e ad

fa r to o n e si .

ap ar t the c l o a c a l w a ll s an d s tu d y the p en i s I t co n sis ts o f two sp o n gy r idge s

.
,

the cor p or a caver n os a o r ca ver n ou s b odies in the ven tr a l w a ll O f the cl oa ca , .

B e tw een the s e fo ld s in the midven tr a l lin e is a deep gr o ove the u r ethral ,

gr oo ve w hi
,
c h in th e n a tu r a l co n di tio n is
p r a c ti ca ll y c on ve r ted in to a tu b e by the
a pp r ox i m a ti o n o f the ca ve rn o u s b o die s The u r e th r a l g r oo ve termin a te s cau d ad
.

a t the b as e o f a he a r t s h ap ed p r o je c ti o n
-
the gl a n s o f the p en i s ,
The an teri o r .

en d s Of the cavern ou s b o dies form the b u l bs a lr e a dy n o ted w hi ch p r oje c t fo rw ar d ,

in to the co elom a t the sides o f the s ta lk o f the b la dde r The b u lb s are lled .

wi th b l oo d which the y r e ceive fr om the in te rn al ili ac vein All p ar ts o f the p eni s .

a r e highl y sp o n gy an d v as c u l ar In the s exu a l ac t the b l o o d fr om the b u lb s

.

ru s he s in to the s p o n gy sp ac e s o f the c a ve rn ou s b o die s an d the gl an s e r e c ti n g ,

the m an d cau si n g the ca ve r n ou s b o die s to co m e in con tac t a b o ve the u r e th r a l

,

g roo ve co n ve r tin g the l atter in to a can al fo r the p assage o f the sp erm

, .

The kidn eys ar e metan ep hr oi an d thei r d uc ts the meta n ep hr ic du cts o r

,

u r eter s The u r e te r w ill b e foun d immedia tel y to the d o r sal s ide o fthe ep idid ym i s
.

whi ch s h ou ld b e r em oved T he u r e ter is a s h o r t s tr aigh t tu b e p r oceedin g to

.

the c lo a ca in to w hi c h it o p e n s j us t an te r i o r to the op e nin g o f the vas defe r en s

, .

The two o p enin g s will b e fou n d a t the sides o f the an ter ior b egi n nin g o f the
u r e th r al gr oo ve .

F in d the Op en in g s o f the a c c esso ry b l a dder s the u r in ary b l a dder an d the , ,

re c tu m in to the c l o a ca T he l a tter is d o r s al to the u r o gen i ta l op en in gs

. .

D raw the male u rogen i ta l sys tem wi th op en cloaca .

E . THE UR O GE NI TA L SYS TE M OF THE PI G E ON

1 . fe mal e
The ur og e n ital s ys te m R em o ve the di ge s tive
tr a c t le a vin g the
.
,

l arge in te s tin e in p l a c e In a du l t b i r d s the r e is a sin gle ovary an d ovidu ct o n the

.

l eft s ide The righ t o va r y an d d u c t a r e p r e s en t in the em b ryo b u t a lm o s t

.

en tir ely di s ap p ea r b efor e ha tchin g The o vary is a m ass con tain in g eggs of
.
 C O MPARATIVE ANATOMY OF THE URO G ENITAL SYSTE M 28 7

vari ou s siz e s si tua ted a t the an ter ior en d of the lef t kidn ey I t is a ttached by
, .

a s h o r t m e so v a ri u m P os terior to the ovary the coiled lef t ovidu ct proceeds to

.

the c l oaca b ei n g supp or ted by the m es o tu b ariu m

,
The os tiu m is s i tu ated in
.

the m es o tu b a riu m n e ar the o v ary ; it is a wide o p e n i n g w i th wi n glike b or de r s

fas ten ed to the meso tu b ariu m A sma ll r emn an t o f the righ t o vidu c t is atta ched
.

to the r igh t s ide o f the clo aca .

The kid n eys ar e metan ep hr oi E ach is a a tten ed th r ee l o b ed or gan si tu ated

.
,
-

a ga i n s t the d o r s al wa l l The u r eters o r metan ep hric du cts ar e lo ca ted j u s t dors al

.

to the r en al p o r ta l vei n s w hi ch s h ou ld b e s tr ipp ed f r om the f a c e o f the kid n e y .

The u r e ter b egin s on each side a t the g roo ve b e tw een the an te ri or an d middle
lob e s of the kidn ey an d ex ten d s s traigh t p osteri orly to the c loaca The lef t .

u r e te r is co n c e a led by the o vid u c t .

The cl o aca is an exp an ded ch amb er r e c eivi n g the r e c tu m on its medi an

ven tral su r fac e the lef t ovidu c t to the le ft the ver y sm a ll righ t o vid u c t to the
, ,

r igh t an d th
,
e u r e te r s d o r sal to the o vid u c ts Cu t i n to the c l oaca t o the r igh t
.

o f the r e c tu m No t e t h at the c avi ty o f the clo ac a is su b divided Ther e

. .

is a l a r ge ven tr a l p or ti o n (cop r oda eu m) in to whi c h the r e c tu m op en s Dor .

s al to this an d s ep a r a ted fro m it by a f o ld is the u r odaeu m in to wh i ch op en

the o vidu c ts an d u r e te r s The op en i n g of the lef t o vid u c t is r e a dily fou n d he r e ;
.

the op e n i n gs o f the u r e te r s are mor e m edi a l an d sm a lle r The mo s t d or s a l .

co mp ar tm en t O f the cl oa ca is the p roctodaeu m a s m a ll c h am b e r w i th a r a i s ed

,

rim w hi c h o p e n s to the an u s
,
In the an te r i o r w all o f the p ro c to d a eu m d o r s a l
.

to the r im an o p e n i n g m ay b e n o ted ; it le a d s in to a s ma ll p o u ch w hi ch s ee m s to
b e fun c tion a l in yo u n g b i r d s b u t degen e r a tes wi th ma tu r i ty I t is ca lled the .

a b r iciu s
b urs a f
o F .

D r aw geni ta l sys tem an d c l oaca

uro .

2 .The m al e ur o ge n ital s ys te m T he tes tes are a p ai r o f ov al o rg an s a t the

.

an te r i o r en d s o f the kid n e y s ; thei r s i z e v a r ie s co n s ide r a b l y with the s e a s on .

The kidn eys ar e the sam e a s in the fem ale an d the des crip ti o n given u n der the
,

female shou ld b e r ea d an d the u r e ter s iden tied The m ale d u c ts vas a defer en tia
.
,

( W ol i a n duc ts ) p
s r in g
,
f r o m th e m edi a l b o r de r n e a r the p o s t e r i o r e n d o f th e

te s te s an d p as s p os t e r i o r l y p ar a llel to the u r e te r s The y ar e s len der con vo l u ted

.

tu b e s . Tr a ce b o th u r e ter s an d v as a defer en ti a to the cl o a ca .

The cl o aca is sm a ller th an in the fem ale an d the li p s o f the an u s mor e p r o tr u d

in g. T he r e c tu m e n ter s m e dia ll y an d ven trall y the u r ogen i ta l d u cts l a te r a lly
,
.

Cu t in to the c l o aca as dir e c ted un der fema le an d iden tify its ch am b er s a s the r e
des crib ed T he y are the sam e in the two sex e s ex cep t th a t the u r o d aeu m is
.
,

sm a lle r an d r e c eive s the two v a sa defe r e n tia in s te a d o f the o vid u c t Ur e ter s .

an d v asa defe r en tia op e n o n sm a ll p ap ill a e in the l a t e r a l w a ll s o f the u r o d a eu m .

Ther e is n o p e n i s in m o s t b i r d s .

D r aw
288 LAB ORATORY MANUAL FOR VE RTE B RATE ANATO M Y

F . THE UR O GENI TAL SYS TE M OF THE MAMMAL

R em o ve the diges tive trac t le avin g the r e c tu m in p lac e

, .

1
.

The kidn eys an d u r e te r s T he kid n eys of m amm al s a re metanep hr oi
.

a n d thei r d u c ts the metan ep hr ic duc ts o r u ret e rs T he kid n eys a r e l ar ge o va l .

o r g an s situ a t ed o n the d o r sa l b o d y wa ll o f the p e r i to n e a l cavity As in all .

ve r te b r a te s they ar e r e tr op e r i to n e al The righ t kidn ey is ge n e rally con siderab ly

.

a n te r i or to the lef t o n e Cle ar aw ay fa t an d co n n e c tive tissu e f r om ab o u t the

.

kid n eys T he m edi al face o f e ach kidn ey is con cave ; thi s co n cavity is ca lled
.

the hil u s F r o m the hil u s a whi te tu b e the u reter p ass e s ou t an d tu rn s po s te

.
, ,

r io r l y
. Fo ll ow the u r e ter s cau d a d c le arin g a way the fa t fr o m a b ou t them an d
,

n o te thei r e n tr an c e in to the b l a dde r I n fem ale s the u r e te r s p ass d o rs a l to the

.

h o rn s o f the u ter u s In m ale s e ac h p ass es d or s al to a whi t e co r d the m ale du c t

.
,

o r vas de fer en s whi ch l o op s o ver the u r e ter an d di sapp e a rs do rs al to the b l adder

, .

R em ove by a cu t the ven tr a l h a lf O f a kid n e y A cavi ty the sin u s is .

, ,

r eve a led w i t hin the hil u s ; th i s sin u s is o c cu p ied c hie y by the ren a l p el vis or ,

exp an ded b egin nin g o f the u r e ter an d a ls o by the r en al ar ter y an d vein In to

, .

the p elvi s the su b s ta n c e o f the kid n e y p r oje c ts a s the r en al p a p ill a o n whi ch ar e

situ a t ed the mi c r o s c op i c op e nin g s o f the c o lle c ti n g tu b u le s The kid n ey s u b .

s ta n c e is r e adil y divided i n to two a r e a s a p e ri p he r a l r egi o n the cor tex an d a

, , ,

c en tr a l r egion the medu ll a

,
T he co r tex c on ta i n s the r en a l co rp u sc les an d the
.

con v o l u ted an d l o op ed p o r ti o n s o f the kid n e y tu b u le s The m ed u ll a is m ar ked .

by lin e s which co n ve r ge to the r e n al p ap ill a ; thes e li n e s a r e the co lle c tin g tu b u le s .

I t wi l l b e r e ca lled th a t the co lle c ti n g tu b u le s the p elvi s an d the u re te r a r e , ,

o u tg r o w th s o f the W o lian d u c t The c o lle c ti n g tu b u les an d r en al papill a

.

toge the r f o rm a pyr a mid o f w hi ch the r e is b u t on e in the r a bb i t a n d ca t b u t

,

a b o u t tw elve in m an .

D r aw .

T he u rin ary bl adder is a p ea r s h ap ed sa c a t the p o s te ri o r en d o f the p er i ton eal

-

c a vi ty . I t is ven tr a l to the r e c tu m in the m a le ven tr a l to b o th r e c tu m an d ,

u t e ru s in the fe m a le T he fr ee an teri or en d o f the b l a dder is n am ed the ap ex

.

o r ver tex the p os t e r i o r p or ti on the fun du s

,
T he fu n d us co n ti n u e s p o s teri or ly
.

a s a n a r r o w ed s ta lk the u r ethr a ( a ls o ca lled n eck Of the b l a dde r ) The b l adder

.
,

is co ve r ed by the p e r i ton eu m whi ch is con tin u ou s wi th th a t o f the ab d omin al

,

wa ll by m ean s O f the m edi an an d l a ter al ligam en ts p r evi ou s ly n o ted The .

p ou c h b e tw ee n t he b l a dde r an d r e c tu m ( m a le ) o r b l a dde r a n d u t e r u s ( fe m a le )
is n am ed the rectoves ical o r ves icou ter in e p ouch r e sp e c tively ,
.

D ra w the ex cr e to r y sys t e m .

The t erm ur ethr a is in m u ch co n f u sion in co m p ara tive a n a t o my , o win t o the dier en ces bet w een g
the uro enita l sy s tems o f vario u s m a mmal s
g Al tho u h in the embryo the ur et hr a is the sa me as the
. g
uro en ital s in u s , t his is n o t the c a s e in the ad ul t s o f m o s t ma mma l s , an d con seq u en tl y the u se o f u r ethra
g
as syn on ym ou s with ur o en ital sin u s a pp ea r s t o b e inad visa b l e
g Urethr a is ther efo re here employed .

in t he sa me sen se as in human an ato my that is , as the n ame o f the duc t leadin fro m the bladd er to
, g
the ext erio r .
2 90 LAB ORATORY MANUAL FOR VERTEB RATE ANATO M Y
o

No w h r ou gh the pu b i c an d i s chi al symp hys es an d s p r ea d the leg s w ell

cu t t
ap a r t Tr a c e the u r e th r a the v a gi n a an d the r e c tu m p os ter i o r l y The u r e th r a
.
, , .

lie s a t r s t on the ven tr al fac e of the vagi n a to w hich it is b oun d by ti ssu e ; it

then u ni te s wi th the v agin a to f o rm a comm on tu b e the u r ogen ital can a l o r s in u s , .

Di sse c t th is fr ee an d lif t it ou t an d follow it to the u rogen i tal ap e r tu r e Cu t .

op en the u r o ge n ita l ap e r tu r e an d n o te the f r ee p o s t e r i o r en d o r gl a n s o f the , ,

c lito r i s p r o je c ti n g i n to the ca vi ty in the r a bb i t

, Fr ee the r e c tu m fr om the u r o .

genita l c an al an d fo llo w it to the an u s Alo n g its sides in the r abb it a re a p ai r o f

.

elon ga ted gl an ds the an al gl an ds In the c a t the r ou n ded an al gl a n ds or s acs

, .

a re si tu a t ed o n e a t e a ch s ide o f t he r e c tu m j u s t i n te r n a l to the a n u s , .

D r aw the fem ale u roge n i ta l s ys tem .

Cu t op en the va gi n a I n the r abb it n o te the ex ter n al u ter in e or ice

.

wi th rai s ed frin ged li p s by m e an s o f w hich e a ch u ter u s op en s in to the va gin a .

The r abb i t u te ru s is of the du pl ex typ e ( Fig 6 3 B p I n the c at the b o d y .

, .

is divided i n to l a te r a l h a lve s by a m edi an p a r ti ti on a h orn o p e ni n g t o e a ch s ide ,

o f the p a r ti tion T he ca t u te r u s is o f the b ip a r tite typ e ( Fig 6 3 C) T he l ow e r

. . .

en d o f the u te r u s ca l l ed the cervix p r o je c ts i n to the v a gi n a by a f o ld

, ,
The .

Op enin g o f the b o d y o f the u t e r u s i n to the v a gi n a is the extern al u ter in e or i ce .

3 The m ale r e pr o d u c tive s ys te m

.
The ex te rn a l p ar ts o r ex tern al gen italia
.

w er e de scr i b ed wi th the ex te rn a l an a tom y R evie w thi s (p The two tes te s

. .

a r e l o dged in the s cr o ta l sac w hi c h is di vided in to two co m p a r tm en ts by a n

i n tern a l p ar titi on Cu t th rou gh the s ki n o ver an d in fro n t O f on e te s ti s ex p o si n g

.
,

the t e s ti s a s an o v a l w hi te b o d y Cle ar a way the con n e c tive ti ssu e a n te r io r

.

to the te s tis an d n d a w hi te co r d the s p er ma tic cor d p a ss i n g fo r w a r d a n d e n te r i n g

, ,

the p e r i ton e a l cavi ty th r o u gh a can al ca lled the in gu in al ca n al T he ex te rn al .

en d o f thi s ca n a l is the extern a l in gu in al r in g; the i n te rn a l en d the in ter n a l ,

in gu in al r in g Tr ac e the sp er ma ti c co r d by cu tti n g op en the c an a l

. .

The sp erm a ti c co r d co n ta in s a whi te d u c t the m a le d u c t o r vas defer en s , ,

an d the b l oo d ve ss el s a n d n e r ve s o f the t e s ti s The two v a sa defer en ti a tu rn .

towar d the m edian l in e lo op o ve r the ve n tr a l su r f ac e s o f the u r e te r s a n d dis

, ,

a pp e ar o n the d o r s a l su r f ac e o f the u r e th r a .

I t is y to exp l ain th es e rel ations o f the tes tes an d th eir d u cts I n the ver te
n o w n ec es s a r .

b r ates p reviou s l y s tu d ied the tes tes lie with in the p erito n eal cavity b u t in mos t adul t mamm als ,

t h ey ar e s itu a ted p o s ter ior to th is c avity I n the emb ryos o f mal e mamm al s h ow ever the
.
, ,

tes tes lie with in the p erito n eal c avi ty ; su b s equ en tl y th ey d es c en d c au d ad Th ere is r s t .

formed a s ac o f t he b ody wall the scro t al s ac which con tain s all Ofthe l ayer s O fthe b od y wall

, ,

skin mus cl es an d p er ito n eu m

, ,
an d in cl o s es a p or tion of t he co el om the vagin al s ec T he ,
.

tes tes th en d es cen d in to the scr o tal s ac as s h o wn in Figur e 6 5 c a rr yin g w it h th em t h eir d uc ts

, , ,

the n erves b lo o d vess el s e tc

,
T he can al al on g w hi ch the d es cen t o ccur s l a ter n ar r o ws an d
,
.

even tuall y is co mp l etel y o b l itera ted in the h ig h er m amm als b u t in so m e r em ain s op en ( S ee .

K p
,
. A n ew can al the in guin al can al is l a t er secon d aril y formed aroun d the sp er matic
, ,

co r d . T he d es cen t of the tes tes exp l ain s the p eculiar loo p in g of the vas a d efer en tia over the
ur et ers an d the co u r s e o f the in t ern al s p er ma tic vein T he s cr otum is now b elieved to s erve .

as a t emper a t u re r egu la tin g mech an ism for the t es tes .

 C O MPARATIVE ANATOM Y OF T HE URO GENITAL SYSTE M 29 1

N ow h rou gh the p u b i c an d i s chi a l s ymp h ys es a n d sp r ea d the leg s ap ar t

cu t t .

Tr ac e the v a s a defe r en ti a an d the u r e th r a c au d a d s ep ar a ti n g them f r om the ,

r e c tu m The u r e te r s m ay b e cu t th r o u gh a n d the u ri n a r y b l a dde r held p o s te

.

r io r l y . T he v a s a defe r en ti a p as s a l on g the d o r s a l s u r fac e O f the u r e th r a I n the .

r a bb it t he y e n l a r ge an d e n te r an e x p an ded s ac the s emin a l ves icl e w hi c h he

p o u c , s ,

f orwa r d b e tw een the r e c tu m an d the b l a dder Cu t i n to the s emi n a l ve sicle a n d .

n o t e the Op en i n g s i n to its ve n tr a l w a ll O f the two v as a defe r e n ti a a n d in its ,

p os t e r i o r d o r s a l w a ll the thi c ke n in g c a u s ed by t he p r os ta te gl an d F i n d t he u n i on .

O f the s e m i n a l ve s icle a n d u r e th r a to f o r m a c o m m on tu b e the u rogen ital s in u s , .

I n t he c a t t he t w o v as a defe r e n t i a j o i n t he u r e th r a w i th ou t t he f o rm a ti on O f a
s em i n a l ve s i c le t he p o i n t o f j u n c t i o n b ein g s u rr o u n ded by a s l igh t e n l a r gem e n t
, ,

i
per ton ea l
cavl ty

i
per tonea l vas deferens deferens
pe i
r ton ea l
cav iy
t

va gi l na

id y mi s

es tis

sc ro t u m
tu n i ca
va gi l
n a is

b ern acul um

FI G 6 5 ia ram s t o i u s tra te the

. . D g ll
d escen t o fthe t es tis in the mal e m mmal The t es tis d escen ds a .

in to the s c r o tu m w h
hic is a s a c o f the o b dy wall co taini g a p or tip n o f the p eriton eal c avity c lled the
n n a

va g in a l s ac ; t the g u bern acu l u m o r lig am en t O f the t es tis sho t en s ; t he p as s ag e x al o n g

in the d esce n r

which the d esc en t o c cu r s an d which a t rs t forms a co n n ec tion b etween the vagin al sac a n d the p e i r

t o u c al c avity is l a t er co mp l etely o bl it era t ed in t he hi gh er m amm al s ; it d o es n o t c o rresp o n d t o t he

i gu in al c a n al ( Fr om P r en t iss an d Arey s Textb ook of E mbr yol ogy cour tesy o f the W B S au n der

n .
, . .

Co mp an y ) .

the p r os ta te g l a n d T he co mm o n.t u b e t h u s f o rm ed i s t he u r ogen ita l can a l o r

s in u s . T he b u l b ou r ethr a l gl an ds o r Cowp er s gl an ds a r e sm a ll s w elli n g s s i tu a t ed

on t he u r o ge ni t a l c an a l s h o r tl y p o s t e r io r t o t he p r os t a t e gl an d in t he r a bb i t,
a b ou t ani n c h p o s t er i o r in the c a t T he t e rm in a l in c h o f the u r o gen ita l c an a l is
.

in c lo s ed in t he p e n i s Cu t i n t o t he p r epu c e an d n ot e t he p o in t ed p r oje c ti on
.

w i th in it ca lled the gl an s o f the p en i s N o t e t h at the p r ep u c e is s imp l y a f o ld

,
.

O f s ki n a r o u n d t he gl an s At t he t ip o f t he gl an s is t he u r ogen ital op en in g
. T he .

gl an s in t he cat b e ar s a n u mb er Of m in u t e sp in e s Di ss e c t an t e ri or ly f r om the .

gl an s exp o s in g the r em ain de r Of the p en i s a s a h ar den ed c y li n d ri ca l s t ru ctu r e .

F i n d w he r e t he u r o gen i t a l c an a l e n t e r s it s an te ri o r en d N o te a l s o t he s tr on g .

a tt ac hm e n t s O f t he p e n i s to the p elvi c r egi on Cu t ac r oss t he middle o f the .

p e n is a n d n o t e t h a t it i s co m p o s ed O f t wo c y lin d r i c a l b o die s t he c or p or a ca vern ,

os a o r c a ver n ou s b odies c l o s el y p l a c ed T he u r o gen i ta l c an al he r e c a lled the

.
,

ca ver n ou s u r ethr a lie s o n t he d o r s a l s ide o f t he p e n i s r e s tin g in a dep r e s s i on

,
29 2 LABORATORY MANUAL FOR VERTE B RATE ANATO MY

b etw een the two ca vern ou s b odi e s At the an te ri or en d o f the pen i s the two .

ca ve rn ou s b o die s di ve r ge f o rm in g the cr u r a Of the p eni s w hi c h a r e att a ched to

, ,

the is chi a T he cave rn ou s b o die s a re sp on gy s tru ctu r e s an d in the s exua l a c t

.

b e come di st en ded w ith b loo d s o th at the p en is is cau s ed to p roje ct ou t o f its

s heath the p repu c e
,
.

D raw the p art s o f the m ale gen i ta l s ystem .

T rac e the r e ctum to the an u s foll ow in g di r e c ti on s given fo r the female .

T he s tru c tu r e o f the t e s ti s may n ow b e in ve s t igat ed E ach t e s ti s is i n c los ed .

in a w hi t e b r ou s s ac the p ar ietal p or tion o f the tu n ica vagin al is whi ch is in

, ,

r e a li ty the p e r i t o n e a l p ou c h ma de by the de s c en t o f the t e s ti s ( F i g Cu t .

o p en t hi s s a c exp o s in g t he ca vi t y in w hi ch the t e s t i s lie s t hi s b ein g a p art of the

, ,

p e r i t on e a l c a vi t y T he tu n ica
. va gin a l is w hi c h li n e s t hi s ca vi t y is r ee ct ed ove r
the su rf a c e o f the t e s t is a s the vis cer al p or tion ofthe tu n i ca v a gi n a li s w hi c h f o rms ,

the ou t e r t hin coat o fthe t e s t i s T he p oin t o fdee c t i on lie s a l on g the mid d o rs al

.
-

lin e of the s crota l s a c an d a me s en t e ry is fo rm ed a lon g t hi s lin e b etw een the s ac

,

an d the t e s t i s T hi s m e s en t e ry is the mes orchiu m c o rr e sp on din g to the b road

.
,

lig am en t o fthe u t erus T he p ost eri or en d o fthe t e s t i s is attac hed to the p os t eri or
.

w all o f t he s cro tu m by a s h o r t b u t s tou t ligam en t the gu bern acu l u m con t in u ou s , ,

w i t h the m e s or c hium T he gu b ern acu lum co rr e sp on d s t o the roun d ligam en t

.

o f the u t e ru s T he du c t of the t e s t i s is the vas deferen s o r Wolian du ct I t lie s

. .

a l o n g the d o rs a l su r f ac e o f the t e s ti s mu c h c o iled the co iled p o r ti on b ein g n am ed ,

the epididymis T he ep i di d ymi s b egin s a t the an t eri or en d o f the t e sti s a s a

.

co il the h
,
ead o f t he ep ididym i s I t then p as s e s d own the d o rs a l su r f ac e o f the
.

t e s ti s as a c o iled tu b e the b ody of the ep ididym is ,

At the p os t e ri o r en d o f .

the t e sti s it f orm s an o t he r co iled m as s the tail of the ep i di d ym i s to w hi c h the

, ,

gu b ern acu lu m is atta ched F rom thi s the vas defe r en s pr o ceed s an t e ri or ly
.
,

mu ch c on v o lu t ed an d p as s e s in t o t he in g uin al c an a l w he r e it b e com e s a s t r a igh t

,

tu b e . T he he ad o f the ep ididym i s is de rived from the me s on ep h ro s an d is

co nn e c t ed wi t h the tu b u le s Ofthe t e s ti s by min u t e va s a e er en tia T he r em ain der .

o f the ep ididymis an d the vas defe r en s a r e the W o lian d u c t .

D raw s h owin g con t en ts of the s crotum

, .

G . THE EMB R Y ONI C ME MB RANE S

1 . G en e r a l .
-
There ar e four e mb ryonic memb ran es in ver tebrates . Th ey a re : the yolk
sac , the mn ion , an d the chor ion ( Fig
all an tois , the a .

T he yolk sa c is s imp l y an evagin atio n from the ven tral w all of the in tes tin e I ts co n n ec .

tion with the in tes tin e forms a n arr o w yol k s tal k The y o lk s ac o ccurs in the emb ryos of all
.

vertebrates havin g merobl as tic eggs an d als o in mammals , for they ar e descen d ed from forms
which h ave merob l as tic eggs The yo lk s ac is l l ed with yolk exc ep t in mammal ian emb ryos
.
,

wh er e it is emp ty .

The all an tois is a l arge eva gin ation from the oo r of the cl o aca I t is p rim aril y r esp ira .

tory in fu n c tion , b u t als o p r ob ab ly s erves to h ol d em b ryo nic ex cr etory ma terials The ad ul t .

bl add er d evel op s at the b ase of the all an toic s talk .

2 94 LAB ORATORY MANUAL FOR VERTEB RAT E ANATO MY
O
umb ilical veins ar e the a b d omin al veins of o l wer forms . The amn ion an d the c horion ne ver

con t ain an y b l oo d vessels .

The yolk s a c may o ccur in the emb r yo s o f an y grou p of ver teb rates The all an to is .
,

amnion a n d ch o rion o ccu r o nl y in rep til es b ir ds an d mamm al s w hich are h en ce d es ig n a t ed

, , , ,

a mn io tes T he all an to is sp read s ou t in s id e o f the c hor io n an d b eco mes fu sed to the ch orio n
.

formin g a chor io all an toic memb ran e ( Fig 6 6 D) In the p l acen tal mammals this cho rio
-
. .

all an toic memb ran e co mes in to cl o s e co n t ac t with the in tern al w all o f the u ter us an d in th e

high est mamm als fu ses in s ep arab l y with the u ter in e wall ( Fig 6 6 E ) The comp oun d stru cture . .

thus p r od u c ed by ap p r oximation or fus ion of the chorio all an to ic memb r an e with the u terin e -

wall is des ign ated the pl acen ta I t is ofvarious sh ap es in various mamma ls the n ame p l acen ta
.
,

b ein g d erived from the disk sh ap e of the hu man p l acen ta .

2 An amn iote e mb ryo of the

. d og s h
o p e n the p r egn an t u te ru s o f a
Cu t .

d o gs h an d r emo ve an em b ryo o r ex am in e em b ryo s p r ovided No t e th at the .

emb ryo is n aked F r om the m iddle o f its ven t r a l wa ll h an gs the large yo lk sac
.
,

lled w i th yo lk an d att a ched t o the b o dy by the n ar row ed yo lk sta lk T he yo lk .

s ac is co ve r ed ex t e rn a ll y by a l a y e r o f the b o d y w a ll a n d in t e rn a ll y co n s i s ts o f

the in t e s t i n a l w a ll in c l o s in g the y o lk As the y o lk is u s ed u p the yo l k sac is

.

gr a du a lly d r awn in t o the b o d y D r aw . .

Am i t e e m b y f th t I f eg t fe m le e av a il a b le Op e n o n e o f
3 . n o r o o e c a p r n a n a s a.r ,

the en l ar gem en ts in the h o rn s o f t he u t e ru s T he en l a r gem en t con ta in s an .

emb ryo N ot e th at the em b ryo is in c lo s ed in a thin m em b r an e the amn ion

.
, .

On the in n e r su r f ac e o f the u t e r in e w a ll at the e n l a r gem en t n ot e a thi c ken ed

vas cul ar r in g of ti s su e T hi s is the pl acen ta ; it w ill p r ob ab ly p eel off fr om
.

the u t e rin e w a ll e sp e c i a lly in a dv an c ed s t a ge s o f p r egn an cy

,
Cu t in to the .

am n i on an d n o t e the u mbil ica l c or d e xt en din g f r om the ven t r a l s ide o f the

a b d om en o f the em b r yo t o t he in n e r s u rf ac e o f the am n ion T he u m b ili c a l co r d .

is a conn e cti on b etw een the em b r yo an d it s o wn m em b r an e s a n d n o t a s p op u lar ly ,

s u pp o s ed a c on n e c t i on b e t w een the emb r y o an d the m o t he r

,
T her e is n o .

dir e ct con n e cti on b etw een em b ryo an d m othe r .

H . S UMIKARY OF THE UROGE NI TAL SYSTE M

1 . T he uro g enital s ys tem is d erived from the meso mere of the emb ryo .

2 . T he urin ar y o r ex cre to r y sys tem con s is ts of the p air ed kid n eys an d h eir d u cts T he
t .

kidn eys mp osed of tu b u l es

are co op enin g at on e en d in to the l
co e om by a n ep hr os to me an d a t

the o th er en d in to the coll ectin g d u ct .

3 I n
. t he ev o l u tio n o f the ver teb rates there have b een thr ee s u ccess i e v kidn eys , eac h
s itua te d more p os terior l y decessor h
t a n its p r e .

4 T he
. r s t ki dn ey o f v ert eb r a tes is call ed the p ron ep hros I ts d u c t is t he p ro n ep h ric .

du ct I t app ears in the emb ryos of all ver teb rates b u t fun ctions in the adul ts of on l y cyclo
.

s tomes an d a fe w s h es .

T h d ki d y is th e m eso n ep hr os o r W o li an bo d y I ts d u c t is the meso n ep hric

5 .e seco n n e .

or Woli an du c t ; this is a con tin ua tion o f the p ro n ep h ric d u c t I t en ters the cloaca The . .

meson ep hros is the fun ctiona l kidn ey o f mo s t a du l t s h es an d all Amphibia .

6 T he thir d kidn ey is the metan ep hr os

. I ts d u c t is the metan ep hr ic d u c t o r ur eter
. .

The ur eter an d p art of the metan ep hros aris e by evagin a tion fr om the Wolian d u ct The .

metan ep hros is the fun c tion al kidn ey of the adul ts of all the amn io tes .
 C O MPARATIVE ANATOM Y OF T HE URO G ENITAL SYSTE M 295

7 A u rin ary b ladd er is

. gen erall y p r es en t as an e ag in ation v of the ven t al wall
r of the
cl o aca I t rec ei es the d u c ts
. v o f t he kid n eys in mam mal s o n l y It . o p en s in to the l
c o ac a in al l

fo rms exc ep t a d ul t placen tal mamm al s .

8 . T he va
o r ies an d tes tes are p air ed bo dies d evel op ed fro m the ven tral f
s ur ac e o f the
mes o mer e an d p r o j ec tin g in to the l co e o m.

9 . T he o varies ar e al w ays l ocated within the p eriton ea l or p l eu r op er ito n eal ca vity in all

ver teb ra t es . T he d u c ts o f the o var ies are the Miill erian d u c ts or o vidu cts They a . r e su p p o se d
to b ed erived from the p ron ep h ric du c t by sp littin g b u t arise in this way onl y in el as mob ran chs, .

IO T he ovid u cts n ever h ave an y d ir ect con n ec tion with the ovaries
. They op en in to .

the co el o m n ear the ovar ies by an o s tium w hich is b elieved to r ep r esen t o n e o r more n ep hr m
s to mes .

II . T he o vidu c ts en ter the l

l y excep t in the p l acen tal mammal s In these
c o ac a s ep ar a te .

l atter the ovidu cts ar e more or less u n ited an d d ifferen tiated in to regions Th is u nio n proceeds .

fro m th eir p os ter ior en ds an teriorl y fo rmin g in the l owes t p l acen tal mammal s rs t a common
,

vagin a in higher forms a p ar tiall y fus ed u terus with s ep ar ate horn s an d in the p rimates a
, ,

s in g l e u teru s p r o d uced by the f u s ion o f two o rigina ll y s ep ara te u teri T he u p p er p or tion s o f .

the ovid u c ts f or m n arro w u terin e tu b es w h ich al ways remain sep ar a te .

12 The tes tes ar e l ocated in tern all y excep t in the hig h er mammals w h er e th ey d es cen d
.
,

temp or ar il y or p erm an en tl y in to p ou ch es of the bo d y w all l o cated ex tern all y in the in g u in a l

region .

1 . v
3 T he d u c ts o f the tes tes in all er teb r a tes ( ex cep t cy cl os to mes an d tel eos tomes) ar e the
v
W O lian or meson ep h ric d uc ts , al so cal l ed the as a d eferen tia I n the mal es of g roup s w here .

the meso n ep hr os is f u n c tion al the W olian d u c ts h ave b o t h gen ital an d ex cr eto r y fu n ctio n s .

In grou p s w he r e a metan ep h ros is p r es en t the Wol ian d u cts h ave o n ly gen ital fu n c tion s .

1 4 T he W o ll an d u c t vas d efer en s
. is al ways d ir ectl y conn ec ted to the tes tis by the
, ,

in terven tion o f a p o r tion of the meso n ep h r os This is differen tiated in to the vas a eer en tia
.

an d the ep id id ymis in p ar t T he remain d er of the ep idid ymis is p ar t Of the Wo li an d u ct

. .

I n mal e amn io tes a p or tion of the meson ep hros t h us al w ays p ers is ts as p ar t of the mal e s ys tem ;
in fem al e amn io tes the meson ep hr os takes n o p ar t in the f c Ttion al fem al e ap p ara tu s .

1 5 A cl o aca w hic h receives the in tes tin e an d ur o g enital d u c ts is p r es en t in mos t ver te

.

b rates ( cycl os tomes tel eos tomes an d p l ac en tal mammal s excep ted) I n p lacen tal mammals
,
.

the cl o ac a sp lits in to a d ors al p or tion w h ich r eceives the in tes tin e an d o p en s to the ex ter io r

by the an us an d a ven tral p or tion the uro genita l c an al or s in us wh ich receives the bl ad
,

der an d the u ro g en it al d u cts T he b l a d d er in mammals d evel o p s a s talk the ure thr a w hich
.
, ,

leads to the exter ior The u reter s s hif t so as to op en d irec tly in to the bl a dder In all mal e
. .

pl acen tal mam m als the vas a d ef eren tia ( W o li an d u c ts ) j oin the ur ethr a to f or m a ur o gen ital

c an al ; in mos t fem al e p l acen tal m am m al s the vag in a s imil ar l y un ites with the u ret hr a for m

in g a ur og en ital can al ; b u t in p rimates the ur eth ra an d vagin a op en sep a ratel y to the ex terio r
'

a u ro g en ital c an al b ein g ab s en t The mal e ur ogenital can al is g en er al l y p r ovid ed in mammals

.

with various gl an d s w hos e secretion is b el ieved to b e ofimpo r tan ce for the s p erm ; its termin al
,

por tion travers es the p en is o r or g an of cop ul a tion a n d for ms the gl an s of the p en is The .

p e nis is f orme d by t he un io n o f two s p o n gy b o d ies t he c avern ous b o dies I t b eg i

,
n s in r ep l es
ti .
,

where the cav ern o us bo dies ar e s ep ar ate .

 X
III . THE C O M P ARATIVE ANAT OMY O F THE NE RV US O SYSTE M
AND THE S E NS E O R GA NS
A . GE NE R A L CON I D S ERATI ONS
In th is s ec tio n o f t he ma n u al we s h all in cl u d e bo th the n er vou s s ys tem p ro p e r an d t he
sen se o r former fu n c tions for the con d uc tion co o r d in ation an d corr el atio n o f
gans . T he ,
-
,

st imu l i w h il e the l a tter is d if

,
fer en tia ted for t he r ecep tio n o f s timu l i .

1 The p art s of the n ervou s s ys tem

. The n ervous sys tem co ns is ts o f th ree p ar ts : the
.
-

cen tr al m, the per ipher al ner vou s sys tem, a n d t he s ympathetic s ystem
n er vou s s ys te T he .

c e n tra l n ervo u s s y s t em is c o mp osed o f the br ain , s itu a ted wit h in t he s ku ll an d o f the s pin al ,

cor d, s itu a t ed w it h in the n eur a l c an al f o rm ed by the n eur al ar c h es o f the ver t eb r ae B rain .

a n d sp in a l c o r d are m a d e u p o f b o t h n erve c ell b odies ( gr ay ma tter ) an d o f n erve-c ell pr oces s es

-

( white matter ) T he p erip h er al n ervo u s s ys t em con sis ts of the cr a n ial n er ves s p r in g in g fr o m

.

the b r ain , a n d the s pin al n er ves , s p rin g in g f r o m the sp in al cor d T he n erves ar e mar ked l y .

m etameric in arr a n g emen t , a p air b ein g typ icall y p r esen t fo r ea c h s eg men t Nerves are .

co mp o s ed o f n er ve cell pr oces s es onl y T he s ymp a th etic n ervou s s ys tem co n tr ol s an d r egu l at es

-
.

in g en eral v l un tary activities a n d the organ s w hich s u b serve th ose f u n c tions as the
the in o ,

h ear t the d iges tive tract the s moot h mus cu l atu re in gen eral the sec etin g g l an d s b l ood ves
, , ,
r ,

s el s r es p ir a tor y a n d ep ro d u c tive s y s tems

, I t co n s is ts c h ie y of a p i ed co rd l yin g a g ain s t
r . a r

the dors al w all o f the co el om f rom w h ic h b ran ch es an d n etworks ex ten d in to t he viscera The .

s ym p a t h e tic s ys t em is co nn ec ted wit h t he p er ip h eral n ervo us s y s te m o f w h ic h in d eed it is

a n o u tg r o w t h Ass ociated with b o th o f these sys tems are n u mero us gan gl ia o f ten s egmen
.
,

t a ll y arr an g ed A g an glion is a collec tion of n erve cell b odies s itu ated outs ide o f the cen tral
.
-
,

n ervous s y s tem W ith in the cen tral n ervo u s s ys tem a s imil ar co ll ec tion is call ed a n ucl eu s
.

o r cen ter a l th ou g h a few o f t h ese a re d es ig n a ted g an g l ia as r eten tio ns f ro m an o l d er t er m i

, ,

n o l ogy

2 . The d evel opm en t of the l n ervo us s ys tem is formed a s

n ervou s s ys tem . T he c en tra

p r ev io u s l y l ear n e d by t h e i n fo l d i n g o f t he e c t o d er m in t h e m e d ia n d o r s a l l in e o f t h e e mb r y o .

A tu b e ex ten d in g t he l en g t h of the emb r yo is th u s p r o d u ced Th at p or tio n o f the tu b e .

s itu a ted w it h in t he h ea d d evel o p s in to the b ra in w h il e th a t p o r tio n p os ter io r to the h ead

,

b ecomes the s pin a l cord T he orig in all y s in g l e l ayer o fenro ll ed ec to d er m c ell s p ro l ifer ates to
.

form a th ick zo n e of cell s aroun d the cen tral c avity M o s t o f t h es e b ecome n erve c ell s .
,

w h il e the rem ain d er give ris e to su pp o r tin g cel ls I n the s p in al co r d o f al l ver teb r a tes a n d .

in the b r ain s of the l o w er o n es the zo n e o f n erve cell s re t a in s its p r imitive p o s itio n a r o u n d

the cen tral c avity b u t in the hig h er ver teb ra tes t h er e is c o n s id er a b l e mig ra tio n o f n erve cell s
,

to the p erip h er y o f the b rain Axon es an d d en d rites ar is e fr o m t he n erve c ell s by o u tg ro wth ;

.

p ar t o f t h ese r em a in in t h e c en t r a l n erv o u s s y s t em f or m in g a p er ip h er a l zo n e o f b ers ca ll ed ,

the w hite ma t ter P ar t o f t he p ro cesses o f the n erve c ell s g r o w o u t fro m the cen tral n ervous
.

s ys tem ( o r g r o w in to it fr o m s en se o r g a ns a n d g an g l ia) f or min g the n erves o r p erip h er a l

n ervou s s y s t em T he g an g l ia o f the p erip h eral n ervo u s s y s tem a r ise c h ie y f r o m the n eu ral

.

cr es ts ; t h ese ar e a p air o f l o n g itu din a l c o r d s o f e c to d er m al c ell s w hic h ar e l e f t o u ts id e o f t he

n eur al t u b e a t t he time o f its c l osure T he s ymp a th etic g an g lia aris e b y the migratio n o f
.

c ells f ro m t he n eur a l tu b e o r fro m the g an g l ia o f the p erip h eral n ervous s y s tem .

In the d evel o p men t O f the b rain t he o r ig in al s imp l e tu b e is r s t mar ked 0 3 in to t h r ee

vesicl es the pr imar y br a in ves icl es by two con s trictio n s ( Fig
, ,
These vis ic l es ar e n amed .

:9 6
 298 LAB ORATORY MANUAL FOR VERTE B RATE ANATO M Y

vis cer al co m p on en t s , t he fo r mer d eal i n g w1 th s tru c tur es I n the b o d y w all the l at ter w ith th os e
O O

O

s it u a t e d in t he vis cer a T he vis c er a l im p u l ses b o th mo tor a n d s en s o r y a r e al w ay s tr an s mit t e d

.

by way O f the s ymp a t h etic s ys tem ex cep t in the c as e Of the visc er al mu s cl es Of the g ill r eg ion
, .

T he sp in a l n erves t r a n s mit a l l f ou r c l ass es O f imp ul s es w h il e t he c r an ial n erves ar e ir reg u l ar

,

in t h is r eg ar d T he s o ma tic s en so r y imp ul s es a r is e in t he s en se o r g an s p er ip h er a ll y l o ca te d
.

in t he b o d y w all a n d p as s a l on g n erves w h ose c ell b o d ies a r e n ea r l y a l w ays s itu a t ed in the

g an g l ia O f t he p er ip h eral n ervou s s ys tem ; the vis ceral sen sor y imp u l s es aris e in n erve en d in g s
in t he vis cer a ; b o t h p a ss in to t he c en tra l n ervo u s s ys tem T he s o ma tic m o to r imp u l s es
.

a r is e f r o m m o t or c ell s w it h in t he c en t r a l n ervo u s s y s t em a n d p as s o u t in t he n erves t o t he

vo lu n tary mu s cl es ; the vis cera l m o to r imp u l ses ar ise in t he b r ain an d c o r d n earl y al w ay s ,

g i
of di es t ve tract

FI G 6 8
. . D iagram
-
hrou gh the spin al c ord an d a spinal n erve t o show the
o f a c r oss -sec tio n t
fu n ction al c omp o n en ts O f the spin al n er ve a n d their r ela tion t o the spin al c ord an d symp athetic sys tem .

S o ma t ic mo to r b ers h eavy co n tin u ou s lin es ; vis c eral mo to r b ers light c o n t in u ous lin es ; s o m tic
, ,
a

s en so r y ber s b r o ken l i es ; vis cer al s en so ry be rs d o t ted lin es ( Slightly al ter ed from Herrick s

,
n , .

I n tr oduction to N urol ogy c ou r t esy O f the W B S au n ders Co mp a n y )

e , . . .

make a r el ay in the s ymp a th etic g an gl ia o r a r ise in th os e g an gl ia ,

an d p a s s to the in vo l u n
t a r y m u s cl es , g l an d s , e t c ( F ig
. .

In a ddi tion t o the fo reg oin g it w ill n a tu rall y b e u n d ers too d t h at a c on sid er ab l e p ar t
,

o f t he cen tr a l n erv ou s s y s tem is c on c er n e d w it h t he c o r r el a t ion a n d c o or d in a t io n O f t he -

f oreg oin g f ou r c l ass es o f f u n c tion s .

The sp in al n erves w ill s erve as examp l es as th ey ar e

4 The p eriph era l n ervo u s s ys tem
. .
,

more t yp ic al .E ac h sp in al n erve is c on n ec ted with the sp in al c o r d by t wo r oots o r b u n d l es

O f b ers a d or s a l a n d a ven tr a l
,
T he d o r s al roo t b ears a gan glion t he dor sa l o r s pin al g an g l ion
.
, ,

w h ic h c o n s is ts o f a col l ec tio n of s en so r y n erve cell s T he b er s sp rin gin g f ro m t h es e cells

.

m ake u p m o s t O f t he d o r s a l r oo t Th is ro o t en t er s t he s p in a l cor d an d co n n ec ts wit h t he

.

d or sa l g r ay c o l u mn T he ven tr al r oo t has n o g an gl ion ; its b er s aris e f r o m t he s o ma tic

.

m o tor c el l s o f the v en tia l c o l u mn a n d t he viscer a l mo to r c ell s o f the l a ter al c o l u mn o f the

gr ay m atter Of the sp in al c o r d ( F ig B eyo n d t he g an g l ion b o t h r oo ts u n it e to f or m a
.
 C O MPARATIVE ANATOM Y OF THE NERVOUS SYSTE M 299

s in gl e s pinal w h ich p asses o u t t hr ou g h the in terver teb ral f or amen a n d al mos t immed i
n er ve

a tel y d ivid es in to th r ee b r an ch es or r a mi
the dor s al ven tr ak; an d vis cer a l r am i Al t h ou g h
, .
,

as in tim ated ab ove t he d o r sal r oo t is a l mo s t p u r el y s ens o ry en tir el y so in t he h ig h er ver te

, ,

b ra tes an d the ven tral roo t p ur el y mo tor the r ami are mixed The d ors al an d ven tr al
, ,
.

rami b o th con t ain s o ma tic mo tor an d s en s o ry b er s p as s in g t o an d f ro m the b o d y w all an d

al so in c l u d e a few vis ceral b er s The so matic mo tor b ers o f t he d ors al r ami p as s to t he

.

ep ax ial m u s c l es t h o se o f the ven tr al rami to the h yp axial mu s cl es

, T he visceral r amu s .

( al so call ed r amu s commu n ican s ) con n ec ts w ith a s ymp a th etic g an gl ion I t carr ies vis c eral .

mo to r b er s f r o m the s p in al cor d in to the s ymp a th etic s y s tem t h es e co n s titu tin g the white ,
.

r a mu s ; an d a l s o c arr ies vis cera l s en s o ry b er s f ro m t he g an g l io n in to t he c o r d th ese f or min g ,

the gr ay r a mu s ( Fig T he w h ite an d g r a y r ami to g e th er fo rm the vis cer al o r c o m

.

mu n ic a tin g ramus w h ich s erves t yp ica ll y to co n n ec t each s p in al g an gl ion with the adj acen t
,

g an gl io n of the s ymp ath etic s ys tem .

T he s p in al n erves are p a ir ed an d seg men t all y arr an g ed Th ere ar e in g en er al as man y .

p airs o f s p in a l n erves a s b od y s eg men ts b el ow the h ea d I n c on n ec tio n w ith the p air ed

.

a pp en d ag es the ven tr a l rami o f sever al s u ccess ive s p in al n e rves f o r m a n et w or k o r pl ex us

from w h ic h the n erves to the app en d age aris e Th is in n erva tion s ho ws : th a t the l imb mu s cl es
.

a ris e f r o m the h yp ax ia l p a r t s o f t he myo to mes s in ce the ven tra l rami s u pp l y o n l y h yp a xial

,

mu s cl es ; t h a t the l imb mu s c l es ar e d er ived f rom s ever al myo to mes s in ce t h er e is b u t o n e ,

S p in al n erve t o ea c h s eg men t o f the b o d y ; an d t h at the mu scl es of the app en d a g es h ave un d er

g on e mu c h to r s ion an d c h an ge of p osition resu l tin g in a cris s cr o s s ar ran g emen t o f th eir

,

n erves s in ce e a c h n erve re t a in s its in n erva tion to t he mus cl e w h ic h it o r ig in all y s u p p l ied

, .

T he cr an ial n erves ar e m u c h l ess t yp ical in arran g emen t t h an the s p in al n erves Mo s t .

o f t h em d o n o t co n t ain t he f o u r c l as s es o f b er s They ar e at tach ed to t he b rain by roo ts

.
,

b u t th es e a r e irr eg u l ar l y arr an g ed Th o se cran ial n erves th at con tain somatic s en sory

.

b ers b ear a g an g l ion corresp o n d in g to a sp in al g an gl ion T he comp osition an d fun ction s .

of t he c r an ial n erves w ill b e d es c rib ed in t he d is sec tio n s Th ere ar e ten cr an ial n erves in.

a n amn io tes an d t w elve in amn io t es b u t t he a dd it io n al t wo ar e n o t n ew f or ma tion s

, T he .

c r an ia l n e rves ar e n o t segm en t a ll y a rra n g ed a l th o u g h p r o b ab l y s o arran g ed in the b e

,

g in n in g .

5 The
. se g men tation I t was p reviously s tated th at the ver teb rate h ead was
of the hea d .

o r ig in a ll y s eg m en t ed b u t the s eg men t a tion is n o w mu c h o b s cu re d

,
I n a t temp tin g to w or k .

o u t t he h ead s egm en ta t io n t he b r ain the cr an ial n erves the vis cer a l ar c h es an d the t ru e
, , , ,

h ead mu s cl es ( n o t vis cer al mu s cl es) h ave b een s tu d ied T he b rain is p l ain l y s egmen ted .
,

p ar t ic u l ar l y in it s p o s t er ior p ar t ; t h es e se g m en t s a r e c al l ed n euromer es P r o b ab l y t he p o s t e .

r ior n eu ro mer es a r e tru e s eg men t al d ivis io n s T he c r an ial n erves are n o w mu c h al tered

.

fro m th eir or ig in al con d ition b u t th ey were former l y seg men tal n erves an d a cer tain amou n t ,

o f evid en ce has co n seq uen t l y b een o b ta in ed f r o m a s tu d y o f t h em T he vis cer al ar c h es .

a r e p l a in l y s eg men t all y arr an g ed b u t the r el a tio n o f th eir seg men t at io n t o t h a t o f the h ead
,

as a w h o l e is n o t en tirel y c l ear I t is g en erall y b el ieved t h a t t he g ill s l its ar e in ter s eg men t a l

.

in p os itio n i e occ u r a t the myos ep ta

,
. .
,
T he b es t evid en ce has b een o b tain ed f ro m the s tu d y
.

o f t he my o to m es o f t he h ead I n cy c l o s to mes a l l o f th es e f orm mu scl es o f the a d u l t b u t in

.
,

o th er ver t eb r a t es man y of t h em d is ap p ear in emb r y o n ic s t ag es Th ose w hic h p ers is t for m .

the mu s c l es o f the ey eb all an d the in tr in s ic mu s cu l a tu r e o f the to n g u e ( T he s tu d en t s h ou l d .

r eco ll ec t t h at the app aren t h ea d muscl es ar e vis c er al m u s cl es d er ived f r o m t he hyp omer es , ,

an d n o t fr o m t he myo to mes ) T he var io us lin es Ofevid en ce l ea d to the fo ll o w in g c o n cl u s io n s :

.

th a t t h er e ar e f o u r h ead seg men t s in f ro n t o f the ea r o f w h ic h the r s t ( an t er io r h ea d c avit y )

is evan es cen t w h il e t he n ex t thr ee ( n am ed 1 2 an d 3 ) g ive r is e to the mu s c l es o f the eyeb a ll ;
, , ,

b ehin d the ear the n u mb er Of h ead s egmen ts app ears to b e var iab l e ab o u t 6 8 in fo r ms ab ove ,
-

cycl o s tomes mor e in c y cl o s to mes

,
These p os to tic s eg men ts d evel op n o mus c l es a bo v e
.
3 0 0 LAB ORATORY MANUAL FOR VE RTE B RATE ANATOM Y

c y c lo st o mes ex c ep t th at the in trin sic mu scl es o f the ton gue ap p ear to b e d erived f ro m th em .

Th es e m atters w ill b e cl ear er a f ter a s tu d y o f t he cranial n erves ( S ee a l s o F ig . .

6 The d e ve l o pm en t o f the s en s e or g an s
.
There are thr ee c hief s en se organs to b e con sid
.

e re d h er e
the n os e the eye a n d t he ear
,
The n o se a rises a s a p air o fin va gin a tio ns o fthe ec to
, .

d er m o f the a n ter ior en d o f t he h ead I n s h es th ese p ersis t as a p air o fs imp l e o l f a c tor y s ac s

.

n o t co nn e c te d w it h the m ou th From Amphibia on war d the Ol f actor y s acs b ecome con n ec ted
.

w ith the o r al c avity by the n as al p as s a ges w h ic h ar e t h en b o th Ol f ac tory an d r esp ir a tory .

T he O l f a c to ry f un c tion is g en erall y l im ited t o t he d or s al r eg ion O f the n as al s ac I n mo s t .

ver t eb r a t es t he w all s o f t he n a s al p as s a g es p ro j ec t in to t he p ass a g es a s cur iou s l y ro ll ed o r

f o l d ed s tru c tur es the tu r bin a l s o r con chae w h ic h s erve to in cr eas e the Ol f actor y s u rfhce an d
, ,

t o w ar m s tr a in an d m o is ten t he a ir p as s in g in
, ,
.

0 T he e y es ar is e in p ar t a s eva g in a t ion s O f the d ien c ep h al on T he tip o f the eva gin a tio n .

t h en in va gin a tes p o d u c g
r in a ou e
d b l w a ll ed cu p the optic cup ( S ee K Fig 2 2 8 p , .
,
. .
,

FI G 6 9
. .
D iag ram to i llu st rat e the
gm en ta tion o f t he ver tebrate head a n d the rela tion o f the
se

c r nial n erv es t o the s e gm en ta tio n

a The n u mb ers a bo v e t he gu r e d esign a te the c ra nial n erv es ; the
.

n u mb ers in the gu re a r e situ ated o n the h ea d myo tom es ; t he s en so ry p ar t o f the n erves i rep resen t ed s

by h eavy con tin u ous lin es ; the mo tor p art by b roken lin es T he a n terior head cavity is the rs t .

myo tome an d th er efore the myotome whi ch is n u mbered I is reall y t he s ec on d myo tome an d so o n ;
b u t a s the myot omes were n um bered befo r e t he a n terio r h ea d cavity was disc ov er ed the old n umbers a re ,

gen erall y ret ain ed T he myotomes n u mbered I 2 a n d 3 p rod u ce the eye mu scl es ; those n um ber ed
.
, ,

d d d g n era t e in the maj o rity o f ver tebr a t es ; tho se fr om 7 o n p r o babl y co n tribu t e to the t o n gu e
4, 5 a n
,
e e

mu scu l at u re b u t n ev er fo rm typ ica l p ariet al m u sc l es su ch a s o ccu r in the t ru nk I t is s een f rom the .

gu re that the thir d cranial n erve a n d the d eep oph thal mic b ran ch o fthe f th b elon g to the rst ( rea lly
s eco n d ) h ea d s egm en t ; the fou r th a n d remain d er O f t he f th t o the s ec o n d ( thir d ) s egmen t ; the s i th x

a n d s even th t o the thir d ( fo u r th ) s egmen t ; the nin th t o the four th ( fth ) s egm en t ; a n d the ten th t o the

fth to eighth ( six th to n in th ) s egmen t s T he g ill slits ar e in tersegmen t l in loca tion The rela tion a

(Af ter Good rich in P art IXof Lan kes ter s

. .

o f the c ra nial n erv es t o the gil l s lits shou l d a l so b e n o t ed

.

Tr eatis e of Z ool ogy cou rt esy o f the M acmill an Comp a n y )

,
.

T he in tern al w al l of d evel o p s in to the r etina or lig h t p er ceivin g l ayer o f the

t he o p t ic cu p -

eye .T he ex ter n al w a ll o f t he c u p b eco mes t he p igm en t l ayer o f t he r etin a T he Op tic .

n erve a ris es in the re tin a a n d p as s es d o w n t he s t al k O f the c u p in t o t he b rain T he r e tin a .

is a p ar t o f t he b r ain w all a s s h o wn by its m an n er of f or m ation

,
T he l en s o f the eye ar is es .

fro m an invagin ation oft he ecto d erm over the op tic c u p The su r f ace e ct o d er m t h en remain s .

as t he c on j u n c tiva o f the e y eb all T he o th er p ar ts o r coa ts o f the ey eb all ( s c l er a c o rn ea

.
, ,

c h o rio id a n d ir is ) a r e d evel o p e d f r om the m es en c h ym e s ur r o u n din g t he Op tic c u p

,
.

The ver teb r a te e ar in its c o mp l ete fo r m co n s is ts o f t h r e e c h amb er s t he in ter n al ea r , ,

the middl e ear a n d the ex ter n al e a r

,
T he in ter n al c ar al o n e is p r esen t in s h es
. I t ar is es .

as a s a c l ike in va g in a tion o f t he ec tod er m o f t he h ea d a t t he l evel o f the hin d b r ain This .

3 0 2 LAB ORATORY MANUAL FOR VERTE B RATE ANATOMY
.
i
s p n al c o r d in t o t he n O n ly the r s t f
. he s e s itu at ed on the d ors al
ou r o r v e o ft ,

s ide O f the b a g f o r m ed by t he p o s t e r i o r c a r din a l s in u s a r e u n i t ed by c r o ss

,

b r a n c he s t o f o rm a t ru e p lexu s the p o s t er i or on e s p as s in g di r e ct ly in to the n

, .

S kate : A l ar ge n u m b e r of ven tr a l r ami su pp l y the p e c to r al n the an te ri or ,

o n e s un i t in g t o a p lexu s S t r ip off t he p leu r op e rit on eu m at t he level o f t he su b

.

c l av ian a rt e r y a n d n o t e t he r e the e n o rm ou s n erve tr u n k o f the b r a chial p l exu s .

I t is f o rm ed by the un i on ( w i thin t he n eu r a l c an al ) of a l a r ge n u m b e r o f ve n t r a l
r am i . T hi s w ill b e s een l at er F o ll ow ou t the n e rve t run k t o t he p e c t o r a l n
. .

I t lie s a l on g the p os t e ri o r s ide of the cu rved ca r t il a ge (p r op t e r y gi um ) w h ic h is

sit u at ed in t he p e c t o r a l n a b ou t h a lf w a y f r om the m id d o r s a l lin e t o t he m a r gi n -
.

Cu t t h r o u gh s kin an d mu s c le s on the d o r s a l s ide o f the an ima l a l on g the p o s t e ri or

a n d l a t e r a l s ide Of thi s ca rt il age a n d exp o s e the t r un k I t s u pp l ie s o n ly the
.

an t e r i o r p a rt o f the p e c t o r a l n T he p o s t e ri o r p art a s a l r e a d y n o t ed is su pp lied

.

by di r e c t ven t ra l r ami n o t f o rmin g a p lexu s T he lu m b os a cra l p lexu s fo r the

,
.

p elvi c n i s l o ca t ed a s f o ll o w s R e m o ve
. t he s kin f r om t he b a s e o f t he n o n t he

d or s al s ide T hi s ex.
p o s e s a f a n s h ap ed l a y e-
r o f m u s c le s Cu t t h r ou gh thi s an d
.

j u s t ven t r al t o it w ill b e f oun d a n um b e r o f n erve s w hi ch dive r ge in to the n

m u s c le s .

2. The s e n s e or g an s F o r the r e s t o f the s e c ti on a l a r ge s ep a r a te he a d will

.

gen e r a lly b e p r o vided I n th at c a s e t he sp e c im en s u s ed u p t o thi s p o in t m ay b e

.

di s ca r ded A ve ry car efu l di s s e cti on of thi s he a d on w hi ch the s tu den t will b e

.
,

gr a ded is r equ i r ed
,
.

a ) The a mp u l l a e of L or en zin i: I t has a l r e a d y b een n o t ed t h a t t he s ki n o f

the he a d is p e r f o r a t ed by p o r e s f r om w hi c h mu cu s e xu de s u n de r p r e s s u r e
, .

N o t e the di s t r i b u t i on of the p or e s R em o ve a p ie c e o f s kin fr om a r egi on b e ar

.

in g p o r e s (in the s k a t e f r om the ve n t r a l s ide o f the he ad ) an d n o t e t h a t ea c h

p o r e le a d s in t o a c a n a l o f v a r y i n g len g t h l yi n g b en e at h the s kin E ac h c an a. l ,

n am ed the can al of L or en zin i t e rm in a t e s in a li tt le b u l b the a mpu l l a ofL or en zin i

, , ,

w hi c h is su pp lied by a n e r ve a deli cat e w hi t e b e r e as il y s een at t ac hed t o t he

,

amp u ll a . T he fun ct io n o f thi s s en s o ry app ar at u s app e ar s to b e the p e r c ep t i on

Of vi b r a t i o n an d p r e s s u r e in the s u rro u n din g w a t e r .

b ) The l a ter a l l in e sys tem: In s he s an d Amp hib ia ( l arv a l s ta ge s on l y O f l an d

A mp hib ia) t he r e is p r e s en t a s ys t em o fs en s e o r g an s c a lled the l a tera l lin e s y s t em
, ,

whi c h is r el at ed t o the a qu ati c mo de o f life T hi s s ys t em is comp let ely l o s t .

,

t o ge t he r w i t h the n e r ve s w hi c h s u p p l y it in the l an d ve rt e b r at e s
,
I t c o n s i s ts .

o f the l a t e r a l li n e c an a l s an d the l a t e r a l lin e n e r ve s T he la t er a l lin e can a l s in

.

s he s con s i st o f tu b e s s itu at ed on the in n e r su r f ac e of the s kin on to whi ch they

op en by p o r e s .I n the c an a l s a r e s en s o ry c ell s w hi c h c l o s ely r e s em b le the s en
s o ry c ell s o f t he ea r an d in f ac t the l at e r a l lin e s y s t em app e a r s t o b e r el a t ed
, , ,

t o the i n t e r n a l ear b o th m o rp h o l o gi c a lly an d f un c t i o n al l y T he f u n c ti on o f the

.

la t e r a l lin e sys t em is b elieved t o b e the p er c ep ti on o f wa t e r vi b r ati on s o f l ow

fr equ en cy .
 COMP AR A I VE ANA O M T T Y OF T HE NE R VO US S YSTE M 0
3 3

Dog s h: Al on g the tru n k the sys tem con si s ts of the l a te r al lin e s whi ch ,

mar k the p o s i t i on o f a can a l F in d t he la t e r a l lin e o n the hea d R emove the

. .

s kin a t t hi s p l a c e n o t i n g the u n de r l y in g c an a l an d t he p o r e s co n n e c t i n g t he c an a l
,

with the s u r f ac e T r a c e the l at e r a l lin e f o rw ard r em ovin g t he skin as you p ro

.
,

c eed . At the level o fthe sp i ra c le s the c an a l s Oft he two l a t e r a l lin e s ar e c o n n e ct ed

by the s u p r a temp or al c an a l An t e ri or t o t hi s e ach fo rk s i n to a su pr aor bital
.
,

c an a l p as s i n g f o rw a r d a b o ve the e y e a n d a n infr a or b ital can a l p a s s in g ve n tr a ll y

b etw een the e y e an d the s p i ra c le an d then f o rw ar d b el ow the ey e Tr a c e the .

s u p rao r b i t a l c a n a l t o the en d o f the r o s t ru m ; he r e it tu rn s a n d p r oc eed s

p o s

t erio r ly a ga in p ar a llel t o its f orm e r cou r s e an d b e com e s c on t in uou s wi t h the

i n fr aor b i tal can al T he l at te r give s Off a hyoman dibu l ar b r an ch run n i n g p os te
.

r ior l y al on g the Side s of t he j aw s an d tu rn s to the ve n t r a l s u r f ac e o f the r o s

,

tru m p as s in g rs t p o s t e ri or t o the n os t ril an d then tu rn in g f orw a r d b e tw een

,

the two n o st ril s . T he r e is a l s o a shor man dibu l ar can al un der the s kin j u st
b ehin d the l ow e r jaw ; it is n ot conn e ct ed wi th the othe r can a ls .

S kate : T he l ater al lin e sys tem is mor e comp lex th an in the dogsh an d
mor e dif cu lt to f o ll ow T he l at e ra l lin e can al ru n s on the d ors a l su r fa c e j u st
.

lat era l t o the mid dor s al sp in e s R emo ve the s ki n at t hi s p l ace an d iden tify
-
.

the can a l T ra c e it f o rw ar d r emovin g the s kin as you p ro ceed At the p o s

.
,
.

t erio r en d o f the c art il a ge ( p r op t e ry gi u m) of the an t e ri or p art o f the p e ct o ra l

n it give s Off two can a l s w hi ch p roc eed p os t e ri o r l y ove r the s u rf ac e of the n .

I t t hen p r o ceed s a b ove the e y e a s the s u p r aor bital can a l app ar en t l y conn e c ti n g ,

wi th its fell ow by a cr o s s un i on on the p o s t eri or p ar t o f the s ku ll

-
T he s u p ra .

o r b i ta l can a l p a s s e s in f r on t o f the e y e an d as the infr a or b ital can a l b el o w t he

ey e In the r egi on of the ey e it give s O b ran ch es ove r the r ostrum an d a l on g

.

b r an ch w hi ch p r oceed s p o s t e ri or ly a l on g the l at er a l margin o f the n On the .

ven tra l s ide o f the s kat e the r e is a p romi n en t can al p as s in g j u st l at er a l to the

gill slits T rac e thi s f orw ar d n ot in g b r an che s b ehin d an d in f ron t of the n os t ril
.
,

an d o n the ven t r a l s u r f a c e o f the r o s tru m On the s u rf a c e o f the p e c to r a l n s

.

a f t e r r e m o v a l o f the s kin the n u m e r ou s ve ry l on g ca n a l s o f L o r e n z i n i a r e n o

,

ticeab l e .

D raw s h owi n g di s tri b u ti on o f the c an als

,
.

c) The olfa ctory organ s : T he s e co n s i s t O fa p a i r O fo lf ac t ory s ac s o n the ven t r a l

s ide o fthe r o s t rum o p en i n g e x t e r n a ll y by the n o s t r il o r ex t e rn a l n a r i s w i th whi c h

, ,

a r e a ss oci a t ed v a r i ou s ap s o f s kin Di s s e ct the s kin away f rom on e O lfactory

.

s a c a n d cu t aw a y t he ap s s o t h a t you can l oo k i n to the sac N o t e the n u m e rou s .

p l a t e s o r l am ell a e a rr an ged i n r o w s i n s ide o f the s ac ; t he s e a r e c o ve r ed w i t h

o lf a c t or y ep i theli u m t he s e n s e o f s m ell b ein g q u i t e keen in s he s
,
P r o ve t o .

you r s elf t h a t t he o lf a c t o r y s ac is c l o s ed in t ern a ll y h a vi n g n o c om m u nicat i on

,

wi t h the o r a l c a vi ty .

D r aw sh o win g the lam ella e

,
.
3 4
0 LAB ORATORY MANUAL FOR VERTEB RATE ANATOMY
O

d) The eye mu s cl es : R em ove the t i s su e f r om ab ou t the ey e ide on t he me

sa s

o f the an im a l a s un de r c) an d c omp le t el y exp o s e the ey e b a ll In d o in g thi s .

,

rs t cu t aw ay t heu pp e r ey elid ( o r in the s k a t e the s kin o ve r the ey e) n o t i n g t h at ,

the inn e r lin in g o f t he e y elid is con t in u o u s wi t h a t hin l a y e r (conju n ctiva ) whi ch

a dhe r e s c l o s el y t o the ex t e rn a l su rf a c e o f the e y e b a ll Next cu t aw ay ve ry ca r e
.

fu ll y the ca r til age b e tw een the eye an d the b rain w hi ch is s een as a whi te ,

lob ed s t ructu r e in the m edi an r egi on an d a ls o the c ar t il age in f ron t of the ey e

, .

D o n o t i n j u r e t he b r a i n an d do n ot cu t in t o t he elev a t i on d o rsa l to the sp i ra c le .

T he s to u t w hit e b an d s s een in t hi s di s s e c t i on ar e c ran i a l n e r ve s I n the s ka t e

, .

Ve ry li t tle cu tt in g is r equ i red T he l arge s om ew h at sp he ri ca l b od y exp os ed is

.
,

the eyeb a ll . I n t he d og shes it is im b edded in a gel a t i n o u s ma t e ri a l w hi c h s h ou ld

b e car efu ll y c le an ed o u t .

T he ey eb a ll r ep o s e s in a cavi ty the or b it to the w a lls o f w hi c h it is a tta c hed

, ,

by mu s cu l a r b an d s the eye mu s cl es T hey a r e v o lu n ta ry m u s c le s de rived f r om

,
.

the my ot ome s o f the s e con d t hi r d an d f ou rt h s egm e n t s of t he he a d

, , T he r e a r e .

s ix o f t he s e e y e m u s c le s whi c h s h ou ld b e ide n t ied a s f o ll ow s F rom the d o r sal .

view f ou r of t hem will b e s een T he on e whi ch is atta ched to the an t e ri or wa ll

.

o f the o r b i t is t he s u p er i or ob l iqu e . T he o the r th r ee o rigin at e f rom the p os t e r o

lat e ra l an gle of the o rb i t an d ar e n am ed recti mu s c le s T he mos t an t eri or on e .

is the in ter n al o r medial r ectu s ; its in s e rti on on the ey e b a ll is cove r ed d ors a lly
by the su p e ri o r Ob liqu e T he n ex t r e ctu s mu s c le is t he s u p er ior rectu s m o r e
.
,

d or s a lly s itu at ed th an the othe r s T he thi r d the ex tern al ( or l a ter al ) rectu s is

.
, ,

i n s e r ted on the p o st e ri o r s u rf ac e o f the ey eb all N ex t r a i s e the e y eb a ll d or

.
,

s a ll y a n d n o t e th a t t he co n j u n ct iv a o r m o s t s u p e r c i a l co a t o ve r the ex t e rn a l

su rf a c e o f t he e y e b a ll is co n t in u ou s wi th the lin i n g o f the l ow e r lid Cu t th r ou gh .

thi s an d f r ee the e y e b a ll ve n t r a ll y c le a n i n g ou t the gela t i n ou s an d b r ou s t i s su e

,

whi c h will b e f ou n d he r e On lif tin g the e y eb a ll the r em a in in g two ey e mu s c le s

.

will b e s een T he infer ior ob l iq u e o ri gin at e s f rom the an t e r om ed ia l c orn e r o f

.

the o r b i t the infer ior r ectu s f r om the p o s t e rom edi a l an gle o f the o r b i t ; b o th a r e
,

in s e r ted in c on ta ct wi t h e ach o the r on the m iddle of the ven tra l s u r f ac e o f the

ey eb a ll T he w hi t e c or d s s een am on g the ey e mu s c le s ar e n er ves
. .

D r aw the ey eb a ll an d it s mu s c le s f rom d ors a l view sh ow in g a s man y o f the ,

m u s c le s a s p o ss ib le .

T he ey e m u s c le s o ri gi n at e f rom the o rb i t an d a r e in s e rt ed on the ey e b a ll .

T hei r ac ti on is t o tu rn the ey e b a l l in va ri o u s di r e c t i on s AS a lr e a d y s ta t ed they

.

a r e de rived f r om th r ee he a d s egm en t s an d a r e in m o s t ve rt e b r a t e s p r ac t i ca ll y

the on l y mu s c le s devel op ed f r om the he ad m y o t om e s T he s e th r ee m u s c le

.

fo rmin g myo tom e s ar e de sign at ed the r s t s e con d an d thi rd a l t h ou gh they ar e

, , ,

in r e al i ty the s e con d t hi r d an d f ou r th o f the he a d myo tom e s s in c e the t ru e

, , ,

rst s egm en t wa s n o t n o ti c ed u n til lat e r A cco r din g t o the r e cen t in ve s tiga

.

ti o n s o f Ne a l the t h ir d m yo tom e give s r i s e t o p a r t O f t he ex t e rn a l r e c tu s the

, ,

s e co n d t o the r e s t O f t he e x t e rn a l r e c t u s an d t o the s u e r i o r o b liq u e w h ile the

p ,
3 0 6 LAB ORATORY MANUAL FOR VERTE B RATE ANATOMY
.

the s ki n . emovin g the s kin b earin g the s e h o le s the en d o lym p h at ic fo ss a

Up on r

o f the c h o n d r oc ran i u m w ill b e f ou n d b en e a t h it In thi s fo ssa ar e two d u c t s .

,

the en dolymp ha tic du cts w hi c h o p en on the s kin by the two h o le s j u s t m en t i on ed

, .

T he s e du c t s c onn e c t the ca vi ty o f the in t e rn a l ear w i t h the s u rfa c e Ve ry ca r e .

fu lly sh ave off wi t h a s ca lp el the c art il age o f the elevat i on con ta in in g the ear ,

wo r kin g o n the s am e s ide a s b efo r e T he r e w ill s oon b e n o t i c ed a ca n a l in the.

ca rt il a ge c o n t a in in g a deli ca t e cu r ved tu b e T hi s tu b e is the an ter ior ver tica l

.

s emicir cu l ar du ct Co n t in u e r em o vin g the car t il a ge w i th o u t in j u r in g t hi s d u c t

.
, .

T he mu s c le s p o s t e ri or t o the ear may a ls o b e r emo ved An ot he r tu b e wi ll s oon .

b e un c ove r ed p o s te ri o r to the r s t on e ; t hi s is the p os ter ior ver tical s emicircu l ar

duct Ther e will n ext b e r eve aled a thi n wal led ch am b er the u tr icu l u s f r o m whi ch
.
-
, ,

t he s e two du cts sp r i n g Con ti n u e p i cki n g aw ay the c ar til a ge in sm all p ie c e s le av

.
,

in g all p ar t s in p l ac e A thi r d du ct the hor izon tal s emicircu l ar du ct l yi n g b el ow

.
, ,

an d l at e r al to the o t he r s will n ex t b e exp os ed ,

When the cartil age has b een re .

moved as far as po s si b le the p ar ts of the i n t ern al ear m ay b e iden tied The

,
.

c e n tr al ch am b e r to w hi ch the d u c ts ar e a tt a c hed co n s i s t s o f a s m alle r d o r s o

an t e r i o r p o r ti o n the u tr icu l u s f rom w h o s e d o r sal t ip the a n t e r i o r ver t i ca l a n d

, ,

horizon tal du cts spri n g ; an d a l arger ven tro po steri o r p ar t the s accu l u s from -
, ,

w hi ch the p os terior ver ti ca l du c t take s o ri gi n The s accu l u s ts i n to a r ou n ded .

depr es sion in the car til age The s emi ci r cu l ar du ct s ar e s len der tu b es cu rved in a
.
,

s emi ci r cle an d e a ch t e rmi n ati n g in a r ou n ded sac the a mp u l l a The am pu ll a e ,

.

o f the an t e ri o r ve r ti cal an d h o ri zo n ta l d u c ts ar e in co n t a c t wi th an d comm u n ica t e

wi th e ach o ther an d al so wi th the an ter ior exten s ion of the ven tr al p art o f the
u tr i cu l u s This ex ten sion is called the recess u s u tr icu l i B o th en d s o f the post eri o r
. .

ver ti cal can al j oi n the saccu l u s In e ach ampu ll a will b e s een a whi t e s en sory
.

p a t c h o r cr is ta t o w hi ,
c h a b r a n c h o f t h e au di t o r y n e r v e is a tt ac hed L a r ge r s en .

so ry p at che s call ed macu l a e al s o o ccu r in the r e c e ssu s u tr i cul i an d in the saccu l u s

, ,
.

In side the s accu lu s a white mas s ofs an d grain s or crystallin e material the otolith
, , ,

is vi s i b le T he mo vem e n t s o f the s e gr ai n s m ay b e con ce rn ed in equ ili b r a tio n

. .

The e n do l ym p h ati c du ct o p en s fro m the m edi al side o f the saccu l u s b u t the co n

n ec tio n is dif c u l t to n d .

D raw s ho wi n g p ar ts o f the i n tern al ear Aft er the drawi n g has b een made the
,
.
,

s a ccul u s m ay b e o p en ed an d the o to li th e xami n ed .

The in te rn al ear has two fu n ction s th at o f hear i n g an d th at of equ ili b r ati on ,

.

The wh ole s tru ctu r e is ll ed wi th a u id the en dolymp h while the c h an n el s in , ,

the ca r til age a r e lled wi t h p er ilymp h Ch an ge s in p r e s s u r e in the e n do l ym p h du e.

ei the r to the impi n gi n g of sou n d wave s on the he ad o r to ch an ge s in the posi tio n

O f the he a d app e ar to b e the s t im u li w hi c h ex ci t e the s en s o r y c ell s o f the cri s t a e

an d m acu l a e p r o d u
,
c i n g in t h e r s t c a s e th e s e n s a t i on o f he a r i n g an d in the

s e co n d s e n sa ti o n s o f the an i m a l s po si ti o n in the w at e r e n ab li n g it to keep in the

,

desir ed p o s i ti o n A cco rdin g to the exp erim en ts o f M axw el l bo th (o r ei ther) the

.
,

c ri s t a e an d the m acu l a e con t r o l equ il ib r a ti o n The c ap aci ty to p er c eive s ou n d s is

.

r esum ab l limi t ed to the cri sta e .

 CO M PARATI VE ANATO MY OF THE NERVOUS SYSTE M 3 0 7

3 . The d or s al e
as p ct of the b r ain T he b rai n is n ow
. to b e exp os ed by c ar e
fu lly p i cki n g away the c art il age in sma ll p ie ce s f rom its roo f . The c ran i al n e rve s ,

whi t e s tr an d s p as s in g th ro u gh the ca rt il a ge mu st n ot b e in j u red On e s ide o f

, .

the he a d ha s t h u s fa r b een lef t in ta ct fo r the s tu d y o f the c r an i a l n e r ve s T hi s .

s ide is n ow t o b e e xp o s ed a l on g w i t h t he b r a in as fa r a s n e c e s s ar y R em o ve the .

u pp e r e y elid a s di r e c t ed u n de r the e y e b u t le ave a ll s t ru c tu r e s in ta c t In .

r em o vin g t he ca r t il age b e tw een the b r a in an d the e y e the f o ll ow in g n e rve s

w ill b e n o t ed : the s u percia l ophthal mic n e r ve runn in g in the w a ll o f the o r b i t

n e ar the d o r s a l s u r f ac e ; the s m a ll tr ochl ea r n e r ve p a ss in g t h r ou gh t he b a c k w a ll

o f the o r b i t t o the su p e r i o r o b liq u e e y e mu s c le ; in the s k a t e the l ar ge r ocu l omotor

n e r ve a c com p an y in g the t r o c hle a r Di s s e ct forwar d to the O lf ac tory s acs exp os

.
,

in g them d or s a lly le avin g the oph th al mi c n e r ve in p lac e

,
R emo ve the s ki n
.

b ehin d the sp i ra c le an d n ot e the hyoman dibu l ar n er ve p a ss in g p ost e r i or to the

s p i racle ; t his n e rve is a l so to b e p r e s e rved T o exp o s e the p o s te r i o r p art o f
.

the b ra in the in t e rn a l e ars o f b o t h s ide s m ay b e cu t t h r ou gh an d the ma ss o f

mu s c le s p o s t e ri o r t o the ear r emo ved a s mu ch a s n e c e s s ary N e r ve s will b e
.

s een p a s s in g t h r o u gh the ca r t il a ge in t he ve n t r a l p a r t o f the ea r b u t a r e n o t t o ,

b e di s s e c t ed ou t fo r the p r e s en t In s h o rt the d o r sa l s ide o f t he b r ain is to b e

.
,

fu ll y exp os ed leavin g al l of the mor e su p e r cia l n erve s in ta ct T he d ors al

, .

asp e c t o f the b r a in w ill t hen b e s tu died r s t a n d the cr an i a l n e r ve s a f t e rw a r d

, .

T he b ra in is s itu at ed in a ca vi t y in the c h o n d ro cran ium w hi ch it on ly ,

p ar t i a ll y ll . s I t i s c o ve r ed by a deli ca t e m em b r an e th e p r
,
im iti ve m en in x in ,

w hi ch the b lo o d ve s s el s o f the b ra in a re s i tu at ed T he m en in x is c on n e cted by

.

s t r an d s w i t h the memb r a n e lin i n g the ca r til a gin ciu s w a ll s o f the c r a n i a l c avi t y .

T he sp ac e b etw een b r a in a n d ch on d r oc r an iu m is lled in life by a u id .

T he mos t an t e ri o r p art o ft he b r a in is the l ar ge olfa ctory b u l b a n e rv ou s m ass ,

s i tu a t ed in c o n t a c t w i t h the d o r s a l w a ll s o fthe o lf ac t o r y s ac s F r om the o lfacto ry

.

s a c a n u m b e r o f ve ry s h o rt b e r s whi c h t o ge t he r c o n s t i t u t e t he olfa ctory n er ve

, ,

p as s in t o t h e O lf ac t o r y b u l b . T he o lf a c t o ry b u l b is s p he r i ca l in t h e d o g s h e s ,

elon gat ed in the s ka te T he o lfac t o ry b u lb is con n e ct ed wi t h the n ext p art of

.

t he b ra in by a s t a lk the olfactory tr act whi ch is s h o rt in the sm o o t h d ogs h

, , .

T he o lfa cto ry tra cts p as s to en la rged rou n ded l ob e s the olfa ctory l ob es at the
, , ,

an t e ri o r ( or l a t e r a l in the s k a t e ).e n d o f t h e m a in m a s s o f t h e b r a in P o s t e ri o r .

to ( d o g s h) o r m edi a l t o ( s k at e) the o lf ac t o ry l ob e s is s i tu at ed an o the r p ai r o f

lo b e s the cer ebr al hemis p her es T he c e reb r a l hem i sp he r e s an d the O lf a ct ory
,
.

lo b e s ar e s ep a ra t ed on ly by a f ain t gr oove All of the p art s th u s fa r m en ti on ed

.

b elon g to the tel en cep hal on T he O lfact o ry b u l b s t r acts an d lo b e s to ge ther c on

.
, ,

s t it u t e the r hin en cep ha l on o r s m ell p o rt i o n O f the b r a in In s he s the c e r e b r a l

.

hem isp he r e s a r e a ls o largely con c e rn ed w i th the s en s e o f sm ell .

P o s t e r i o r t o the c e r e b ra l hem i sp her e s is a dep r e s s ed r egi o n the dien c ep ha lon ,

o r t h a l am e n c ep h a l on ,
ve ry n arr ow in the sm ooth d ogsh T he roo f of thi s is .

thi n an d d is c o l o r ed du e t o the f ac t t h a t it co n s i s ts e n tir ely o f a p lexu s o f b l oo d

,
3 0 8 LAB ORATORY MANUAL FOR VERTEB RATE ANATOMY
O

ve ss els ,
ca lled a chor ioid pl exus T hi s p art i cu l ar p lexu s is n am ed
. the chor ioid

pl exu s of the thir d ven tr icl e T he dien c ep h a lon s er ve s chie y a s

. e e a c n t r of

c o o r din a t i o n o f t he p ri n c ip a l s e n s at i on s s u c h as s igh t he a r i n g a n d s kin s ens a

-
, , ,

ti on s it s ve n t r a l p o r t i on s b ein g dev o t ed t o sm ell an d ta s t e

,
T he dien c ep h a lo n .

is in low e r ve rt e b r at e s t he c hief c on t r o llin g r egi on of the b r ain T he op t i c n erve .

p as s e s f r om t h e o r b i t in t o the ven t r a l s u r f ac e o f the die n c e h a l o n

p I t is q u i t e .

c on sp i cuou s in t he s k at e a n d w ill b e s een in the d o gs hes by ge n t l y p r e s s i n g the

dien cep h a lon t o on e s ide P ost e ri or to the dien c ep h alon is t he midbr ain or
.

mesen cep hal on c on s i s t in g d ors a ll y o f two r ou n ded l ob e s the op tic l ob es or cor p or a

, ,

b igemin a
'
T he Op t i c l ob e s a r e c en t e rs fo r the vi su a l a u di t o ry an d gen e r a l s kin
.
, ,

s en s at i on s an d
,
t he s e a r e con n e c t ed wi t h co rr e sp on din g c e n t e r s in the
dien cep h alon A n e rve the tr ochl ear or fou r th cr an ial n e rve ari s e s f rom the
.
, ,

p o s t e r i o r b o r de r s o f t he op t i c l o b e on e a c h s ide an d p a ss e s f o rw a r d t o an e y e

mu s c le . B y gen t l y p r e s s in g the op t i c l o b e s t o o n e s ide a n e rve t he ocu l omotor , ,

o r thir d cr a n ial n e rve w ill b e s ee n em e r gin g f r om t he ve n t r a l s u rf a c e o f the m id

,

b ra in an d p ass in g t o the o r b it .

P o st e ri or to the op tic lobes an d somewhat overh angin g them is the large

cer eb ell u m o r meten cep hal on In t he sp in y do gs h a n d s k at e thi s is s ligh tl y di vided
.

in to fou r qu a d ran t s by fain t lon git u din al an d t ran s ve rs e g roove s ; in the sm ooth
d o gs h it is ma r ked by s eve ra l t ran s ve rs e gr oove s T he c e r eb ellum is a c en t er
.

fo r the co o r din at i on of mot or impu ls e s in c l u din g the ma in t en an c e of equilib

-
,

r iu m . N o n e r ve s a r e at ta c hed to the c e r eb ellum P ost eri or to the c e r eb ellum

.

the el o n ga t ed r emai n in g s e cti on Of the b r a i n is t he med u ll a ob l on ga ta or m yel en

c ep h a l o n .I t is con t in u ou s p o s t e ri o r ly wi th the s p i n a l co r d T he an t e ri o r p ar t .

Of t he ro o f o f the med u ll a is t hin a n d di s c o l o r ed an d co n s i s t s o f a chor ioid p l exu s ,

th a t o f the four th ven tr icl e T he an t e ri or en d o f t he med u ll a ex t en d s forw a rd

.

a t t he s ide s o f a n d b el ow the c e r e b ell um a s two e a r l ike p ro je c ti o n s s om e tim e s ,

Cal led the au r icles R em o ve the ch o r ioid p lexu s fr om the roo f o f the au ri cles
.

an d the m ed u ll a T he l arge cavi ty r evealed is the cavity of the fou r th ven tr icl e
. .

The au ricles to ge ther wi th the p or tion of the d o rsal pa rt of the m ed u ll a j u s t

p o s t e r i o r t o t he m c o n s ti tu t e th e ac us tico l a ter al ar eas o r
-
p rim a r y c e n,
t e r s o f t h e

l ateral lin e sys tem an d of hearin g an d equilib rati on On liftin g the po s teri o r .

en d o f the c e r e b ell u m it w ill b e seen that the a u ri cle s ar e con t in u o u s wi th the

c e r e b ell um an d wi th ea ch o the r b el ow the ove r h an gi n g p o st e ri o r p o r ti on of the

,

ce r e b ell u m B y way o f this con n e cti on wi th the c e r e b ell um the imp u l se s which
.

com e in f rom the l a te r al lin e sy s tem an d in te rn al ear a r e c on vey ed t o the c e r e

b el lum af te r maki n g a r el ay in the a cu s ti co l a te ral are a T he en ti r e d or sa l -

.

r im o f the m ed u l l a in clu din g the acu s ti co lateral ar eas form s an el on gated

,
-
,

s tr ip on e a ch s ide which is kn own as the soma tic s en s ory col u mn T his co lumn .

as its n am e im p lie s con tain s c e n te r s an d t r a c ts a sso ci a ted w i th a ll o f the b od y

s en s e s exc ep t s mell ta s te an d vis io n w hich as w e h ave s ee n a r e di sp os ed of

, , ,
3 10 LAB ORATOR Y MANUAL FOR VERTEB RATE ANATOM Y

F rom the lfacto ry b u lb a ft er a r elay the O lfact ory impu ls es pas s a l o n g the
o , ,

O lf ac t or y t r a cts t o c e rt a in p art s o f the b r a i n de s cri b ed a b o ve T he o lf ac tory

.

n e r ve is a p u r e s en s o r y n e r ve .

b ) The s econ d or op tic n erve: T he op ti c n e r ve a ri s e s in the r e tin a o f the e y e

a n d p a s s e s t h rou gh t he c o a ts o f the e y e em e r gin g ve n t r a l t o the in t e rn a l r e c tu s
,

m u s c le F in d it t he r e on the in t ac t ey e by p r es s in g t hi s mu s c le a ga in st the ey e
.

b a ll I t is a s t ou t w hi t e t run k w hi c h p ie r c e s the ca r t il a ge o fthe o rb it an d p as s e s

.
,

to the ven t r a l s ide O f the di e n c ep h a l on I t may b e s een he r e by gen t l y r ai s in g

.

the di e n c ep h a l on T he op t i c n er ve is n o t r e a ll y a n e r ve fo r the r etin a in whi ch

.
,

it a ri s e s is a p art o f the b r a in w a ll an d in the r e t in a the r e a r e s eve r a l r ela ys o f

,

n e r ve c ell s b e tw een the r o d s an d co n e s an d t he c ells o f o ri gin o f the op t i c n e r ve .

T he op t i c n e r ve is r e a lly a tr act o f the b r ain I t carrie s s en s ory imp u l s e s on l y

.
,

vi su al impu l s e s an d di s ch ar ge s t hem in to c e rta in p arts of the dien c eph al on an d

,

op tic l ob e s .

c) The fou r th or tr ochl ear n erve: T he t r oc hle ar n e r ve a r i s e s in the mid b ra in

an d em e r ge s in t he gr o o ve b e tw een the op ti c l o b e s a n d the c e r e b ell um T rac e

.

it o n the s ide w he r e the e y e is s t ill in ta ct I t p as s e s f orwa r d in the cr an i a l cavi ty

.

t o a b ou t the level O f the c e r e b r a l hemi sp he r e s ; it the n tu rn s a b ru p t l y la t e ra l ly ,

p ie r c e s t h e w a ll o f t he o r b i t a n d i s di s,
t r i b u t ed t o th e s u p e r i o r O b liq u e m u s c le
o f the e y e b a ll I t is the m ot or n e r ve o f thi s mu s c le an d ca r rie s o n l y s om ati c
.

mot o r imp u ls e s Al th ou gh it app e ar s ext e rn a ll y to em er ge f rom the r oof of the

.

mid b r a in the mot o r c ell s f r om whi ch it o rigin ate s a r e in fa c t in the o o r o f the

,

m id b r ai n in a f o rw a r d ext en s i on o f the s omat i c m o t or co lu mn .

d) The thir d or ocu l omotor n erve: T he ocu lomo t o r n e r ve a ri s e s f r om the o o r

o f the m id b r a in I t is r e a dil y n o t i c ed in the s k at e a s c e n din g t o the o r b i t n e a r
.
,

the p r e c edin g n e r ve I n t he d o gs hes it is deep l y s i tu a t ed a n d is s een by p r e s s in g

.

the c e r e b ellu m aw ay f rom the w a ll o f the o rb i t F o llo w it in t o the o r b it on b o th

.

s ide s ge tt in g its gen e r a l r el a t i o ns r s t o n the s ide w he r e the e y e b a ll wa s r em o ved

, .

I t em e r ge s in t o the o r b i t ve ry n e ar the in s e rt i on o f the s u p er i o r r e c tu s mu s c le

an d is s i t u a t ed ven t r a l t o the su p e r c i a l o p h t h a lm i c n e r ve a l r e a d y n o t ed ,
It .

s h ou ld n o t b e c o n f u s ed w i t h the deep op hthal mic n e r ve w hi c h is in c o n ta c t w i th

it as it en t e r s the o r b i t ; the deep op h t h a lm i c n e r ve r un s t h r ou gh the orb i t in

c o n ta c t w i t h the m edi a l s u r f ac e o f the e y e b a ll T hi s n e rve w ill b e b ett e r s een
.

on t he in t ac t s ide O b s erve the b ran che s given by the ocu lomotor n erve imm edi
.

a t el y a f t e r it s en t r an c e in t o the o r b i t t o the i n t e rn a l a n d s u p e r i o r r e c ti mu s c le s .

On the in t ac t s ide n ow l oo s en t he e y eb a ll an d cu t t he in s e r ti on s of the s u p e ri o r

Ob liqu e an d s u p e r i o r r e c tu s c l o s e t o the e y e b a ll I den tify the deep op hth almic
.

n e r ve p as s in g in t he s p in y d o g s h d o r s a l t o the in t e rn a l r e c tu s l yin g a g a in s t the

eyeb a ll an d in the sm oo th d ogsh an d s k at e ven t ral t o the i n t ern al r e ctu s F r ee .

an d p r e s e r ve t hi s n e r ve Cu t th r ou gh t he in s e rt i on o f the i n fer i o r r e c tu s an d
.

the op t i c n e r ve an d p r e s s in g the e y e b a ll ou tw a r d n o t e the b r an c h o f the oc u l o

, ,

m o t o r n e rve whi ch p ass e s a l on g the p os t e ri or s ide o f the in fe ri or r e c tu s mu s c le

, ,
 C OMPARATIVE ANATO M Y OF THE NERVOUS SYSTE M 3 1 1

tu rn s ven t r a l to it an d then run s forwar d in the obr of the o rb i t t o the in feri o r

,

o b l iqu e No t e a l so tha t the br an c h whi ch su pp lie s the i n fe r i o r r e c tu s give s o ff

.

a b ran c h o n e o f the cil iary n e r ve s w hi c h p as s e s in t o the e y e b a ll in comp an y

, ,

w ith an art e ry Alon g t hi s n e rve small b ro wn mas s e s can b e n o t i ced ; they a r e

.

the cil iary gan gl ia b el on gi n g t o the sym p at he t i c s y s t em T he deep op hthal .

m ic n e r ve a ls o su pp lie s c ili ar y b ran c he s i n t o the ey e b a ll T he fu n cti on o f the .

c i l i a r y n e r ve s is t o con t r o l the sm o o t h m u s c le s o f the i ri s r egu l at i n g t he s iz e o f ,

the p u p il ; t he y a r e vi s c e ra l m o t o r n e r ve s ma ki n g a r el ay in s ymp a the ti c

,

g g a
a n .

It will fro m the fo rego in g a ccoun t t h at the o cu l o mo tor n erve s upp l ies fo u r eye
be s ee n

mus cl es n amel y th o s e w h ich d evel o p fr o m the so call ed r s t h ea d myo to me

, ,
-
I t is a s o ma tic .

mo tor n erve o r ig in a tin g f r o m the s o ma tic mo to r col u mn in t he oor o f the mid b ra in It .

al so h o w ever c ar ries wit h it symp ath et ic b er s o f vis c er a l mo to r fu n c t ion

, , .

e) The s ixth or a b du cen s n erve: T he a b duc en s or igin at e s f rom t he so m at i c

m ot o r co lum n on the ven tr a l su r f ac e o f the an t e ri or en d o f the m ed u ll a I ts .

o rigin wil l b e s ee n l a t e r I t p en e tr ate s the o r b it a t the p oin t Of o r igl n o f the

.

'

ex t e rn al r e c tu s mu s c le an d p ass e s a lon g the ven tra l su r f ac e o f thi s m u s c le t o

whi c h its b e r s a r e di s t ri b u t ed I t will b e s een a s a w hi t e ridge on the ven t r al
.

s u rf a c e o f the mu s c le .

I t will n ow be seen h th fou r th an d s ix th cr an ial n erves are s omatic mo to r

a t the t ir d , ,

n e rves t o t he mus c l es o f the eyeb all T he r eas on for the all o tmen t of t h r ee cr an ia l n erves
.

to six s m a ll mus cl es has al r ea d y b een in dic a ted To ever y s e gmen t o f the ver teb r a te b o d y
.

th ere s h o u l d b e t h eor e tic all y a p air o f mo tor a n d s ens or y n erves S in ce as al read y ex .

p l ain ed th e s ix e ye m u s c l es c o m e f r o m t hree h ea d my o t o m e s it is r ea d il y u n d er s too d

, t h a t
a so m atic m o tor n erve s h o u l d b e p resen t to s u p p l y the d er iva tives o f eac h o f t h ese myo

tomes T he s en sory n erves o f t h ese segmen ts will b e in d ica ted l a ter

. .

f) T he fth or tr igem in u s n er ve: T he t r ig e m in u s is a ve r y l a r ge n e r ve w i t h

fou r main b ran che s in el asm o b ran ch s (th ree in lan d ve rt eb rat e s) T he trigemi .

nu s is hed to the m edu ll a n e ar the an t e ri o r en d Of the s omat i c s en s o ry

a tt a c

co lu mn j u s t b eh in d the au r icle s o f the med u ll a

,
I ts roo t s he r e a r e in ex trica bl y
.

min gled w i th the r oots of the s even th an d eigh th n e rve s the th r ee to ge t he r ,

fo rm in g a c on sp i cu ou s mas s at the p lac e s ta t ed T he t r igem in u s p as s e s th r ou gh .

the a dj ac en t w a ll o f the o rb i t an d s h ou ld b e f o ll ow ed in t o the o r b i t by c a r ef u ll y

p i c ki n g a w a y the ca rt il a ge a r oun d it A s s o o n a s it p en e t ra t e s t he o r b it the

.

t rigem in u s divide s in t o f o u r b r an c he s T he r s t Of the s e the s u per cial oph

.
,

thal mic b ran c h is p art o f the s u p e r c i a l op htha hn ic t run k w hi ch ha s a l r e ad y

,

b een m en t io n ed s eve ra l t im e s T hi s lar ge t ru n k p a s s e s fo rw a r d in the d o rs a l p art

.

o f the c a rt il a ge o f the m edi a l w a ll o f the o r b i t T rac e it f orwa r d I t p as s e s o u t

. .

Of the o r b i t t hr o u gh t he Op h t h al mi c fo r am en in the c ho n dr o cr a n i u m an d a b o ve ,

the o lfac to ry b u l b O n l y a s m a ll p ar t O f this tr un k is tr igem i n a l ; t hi s is s en so ry

.

to the s k in do rs a l to the o r b i t The se co n d b r an ch o f the trigem i n u s is the deep

.
3 12 LAB ORATORY MANUAL FOR VERTE B RATE ANATOM Y

Op hthalmic e ve n r . T his p as se
hr ou gh the o rb i t ven tral to the pr e cedin g an d
s t
leaves the or b i t by the o rb i ton asa l can al Upo n tr acin g it forwar d it will b e .

s een to j o i n the su p e r ci al Op h th almi c an d to b e dis tr i b u t ed in co mm on wi th it .

B o th o f t he s e b r an c he s o f the tr igemin u s a r e p ur e so m a ti c s en so ry n er ve s ar i s in g ,

from s en s e or gan s o f the skin .

The two r em ain in g b r an ches o f the trigemin u s lie in the oor of the or b it .

T0 s ee them r emo ve the eye b all o r s tu d y the side w her e the eyeb all was pr evi ously
r emo ved A b road w hi te b an d the infraor b ital trun k is s een in the oo r of
.
, ,

the o r b i t p ass i n g o b l iq u el y l at e r al l y T hi s tru n k in the d o g s he s is compo s ed o f

,
.

the mix ed b e r s o f the maxill ary b ranch o f the tr igemin u s an d the b uccal br an c h
Of t he s even t h n e rve ( s ee b el o w) In the o r b i t the l ar ger an d mo r e m e dial po rti on
.

o f t he tru n k is the maxill ary b r an ch b u t far the r ou t t hi s b e co m e s in extri ca b l y

,

mi n gled wi th the b u ccal n er ve In the s kat e the i n fr ao r b i tal tru n k is di vi si b le

.

in to th r ee tr un ks ofw hi ch the ou ter on e is the maxil l ary b r an ch Ofthe trigemin u s

, ,

the middle o n e the man dibu l ar b r an c h o f the tr igemi n u s an d the in n er o n e the ,

buccal b r an c h o fthe s even th n er ve As b efor e ho w ever it sho u ld b e r em e m b er ed

.
, ,

t h at the r e is an a d mix tu r e o f b er s o f the fth an d s even th n e rve s in t he s e trun k s .

Tr ac e the maxill ary b r an ch o f the trigemin us an d b u ccal b r an c h o f the s even th

o u t fro m the or b i t al on g the ven t r al s u r f a c e o f the r os tr u m
,
T he b r an ches pas s .

to the r egi on b el o w an d in f ron t o f the ey e to the m edial s ide o f the n ostr il (in ,

the sk at e to the l at er al s ide o f the n os tr il al so ) an d to the an gle o f the j a w s In .

the smoo t h do g sh t he r e is a con sp i cu ous b ran ch a l o n g the l o w e r ja w whi ch

ap p e a r s t o b e a p ar t o f the m an di b u l ar b r an ch o f the tr igem i n u s T he m axill ary .

b ran ch is s en so ry to the s ki n of the rostrum w hile the b u ccal n erve s upplies the ,

in fraor b i tal l at eral lin e can al an d n e ar b y amp u ll ae o f Lo r en zin i -

.

The fou r th b r an c h o f the t rigemin u s is the man dib u l ar b ran ch In the do g .

s he s it s ep ar a t e s fro m the in frao r b i tal trun k wher e the l att er en t er s the or b i t

from the b r ain an d p as s es alon g the po s teri or w all o f the or b i t In the smoo th .

do gsh p ar t o f it s eems to a cco mp an y the in fr ao r b ital tr un k fo r war d an d then

cur ve s to the lo w e r jaw as de s c r i b ed ab o ve The m an di b u lar n erve is s een to
.

b ran ch to var io u s mu s cles in the oor o f the o r b i t (thes e are gill ar ch muscle s) -

an d o n fo ll ow i n g it o u t o f the o r b i t will b e s ee n to b e di s t ri b u ted to mu s c le s

,

o f t he lo w e r jaw an d to s en d a s e n so r y b r an c h to the skin o f the lo w er jaw th is ,

b r an ch b ein g si tuated j u st b ehi n d the teet h In the s kate the po si tio n o f the .

m an di b u l ar n erve was de s cr i b ed abo ve as b etw een the m axill ary an d the b uccal
n e rve s .Fo l l ow it for war d I t cur ves aroun d the an gle o f the jaw an d s upp lies
.

mu scles o f the l o w e r jaw an d the adj acen t skin .

It w ill b e o b served th at all o f the b ran ch es of the fth n erve are S oma tic sensory n erves
c o min g fro m va riou s s en so ry o r g a ns o f the S kin ex cep t the m an d ib ul a r n erve w h ich al s o
,

co n ta in s s o me m o to r b ra n ch es t o muscl es As th o se mu s cl es are vis ceral mu s cles this pa r t

.
,

of the f th n erve be l on g s to the vis cera l mo tor s ys tem The d eep o ph thal mic n erve appears .
3 4
1 LAB ORATORY MANUAL FOR VE R E B RATE
'
ANATOMY
Q

h) The eighth or au ditor y n erve: The au di to r y n erve is e somati c s ensory

a p u r

n e rv e ex ten din g f r o m the in t e rn a l ear to the b r ai n I t en t e r s the an te ri or en d

.

o f the m ed u ll a a n d is the r e m in gled wi th the r oo ts o f the fth an d s eve n th n e r ve s .

F o ll o w it i n to the i n tern al ear o n the side opp o s i t e th a t on whi ch the hyo man
dib u l ar was diss e c ted . No te its b r an che s to e ach ampu l l a an d the fanlike arr an ge
m en t o f the b ran chle ts to the cr i s ta o f ea ch amp u ll a T he au di to r y n er ve al so
.

c o lle c ts a n u mb e r o f b r an c he s f r o m the w a ll s o f the s a ccu l u s an d u tr i cu l us It .

ca r r ie s im p u l s e s fo r he ar in g a n d eq u ili b ra ti on in to the a cu s ti c o l a te r a l ar e a o f -

the m ed u ll a to whi ch it will b e fou n d to b e a ttached T her e is n o mo tor n er ve

.
.

c o rr e sp on din g to the a u di to r y n e r ve .

i) The n in th or gl oss op har yn geal n erve: Thi s n e rve p ass e s th r ou gh the oo r

o f the middle o f the ear cap s u le ( w he r e it is likel y to b e m i s take n fo r a t f
p a r o

the au di to r y) p ar a llel to the h yo m an dib u l a r n e r ve

, Par e away as mu ch of the
.

ea r c ap su le as is n e c e ssa r y to r eve al it .F in d its a t ta chm en t to the m edu ll a

p os t e r i o r t o th e a u di t o r y n e r ve . T ra c e it o u t o f th e ea r c a p u le
s J u s t b
. ef o r e it
exi ts from the ear cap su le it b e ars a sw e l l in g the petros al gan glion In s er t a
, .

kn ife b l ade in to the s e con d ( rs t typ i cal) gill s li t (in the sk a te in to the d o r sal
wa ll o f the corr e sp on din g vi s ce r a l p o u ch) an d s li t the gill c lef t op en d o r sally .

The p e tr o s al gan glion will n ow b e s een to b e lo ca ted n ear the upp e r li mi ts o f the
c lef t . D iss e c t the n e rve fr om the gan gli on towar d the gill s li t I t ve r y soon .

divide s in to th r ee b ran che s two smaller an teri or on e s an d a lar ger pos teri or
, ,

on e . The mos t an te r i or b r a n ch is the p r etrema tic b ran ch an d p ass es to the

a n te r i o r w al l o f the vi sc e r a l p ou c h to w hi c h it is a s e n so ry n e rve The s e con d .

b ran ch is p o s teri o r to the p r e tr em a tic b r an ch ; it is n am ed the p haryn geal b r an ch

a n d is a s en s o r y n e rve to the m ou th cavi ty .
( T hi s b r an ch app e a r s t o b e l ac k i n g
in the S ka te ) T he thir d an d l a r ge s t is the p os t tr ema tic b r an ch
.
-
I t p as s e s to
.

the p o s te r io r w a ll o f the vi s ce r a l p ou c h an d is b o th s en so ry an d mo to r its mo to r ,

c o mp o n e n ts su p p l yi n g the m u s c le s o f the thi r d vi s c e r a l a r c h The gl oss op haryn

.

ge al n erve is the n erv e o f the s econ d vis cer al p ouch an d o f the third viscer al ar ch .

j) T he te n th o r va g us n er v e: T he v a gu s n e r ve i s t h e ve r y l a r ge tr u n k p as s i n g
th r ou gh the p o s te r i o r b o r de r o f the ear cap su le I t is a tta ched to the s ide s o f
.

the p os te ri or p ar t of the m ed u ll a . D iss e c t it ou t an d foll ow its co u rs e I t .

p a ss e s t o t h e a n te r i o r ca r din a l S i n u s the w a ll o f w hi ch is fo r m ed in the d o g shes

,

o f a tou gh m e m b r a n e . Op en u p the an t eri o r ca r din al S in u s by a deep cu t th r ou gh

the mu s c le s m edi a l to the vi s c e r al p ou che s Fo ll ow the va gu s n e rve i n to the
.

an te r i o r ca r din al s i n u s . In the dog shes it divide s in to two tru n k s a t the p o in t

wher e it p en e tr a te s the tou gh wa ll o f the s in u s The m edi a lly situ a ted tru n k
, .

is the l a tera l b r a n ch o f the v a gu s an d p as s e s p o s te r io r l y in te rn al to the l a te r a l

lin e wh os e can a l it supp lie s The la te ral trun k is the viscer al b ran ch o f the
.

vagu s which con tin u es alon g the an teri or c ar din al s in u s I n the s kate the
,
.

vagu s run s for a s h or t di s ta n c e in the sin u s b efor e dividin g in to a m o r e d orsal

l ater al b r an ch whi ch p as s e s pos teri o r ly in te r n a l to the l a te ra l lin e ca n a l to w hi ch
,
 COMPARATIVE ANATOMY OF THE NERVOUS SYSTE M 3 1 5
.
it is di s tri b u ted an d a mor e ven tral viscer al b ran ch In all th r ee forms op en
, .
,

u p the r e m ai n i n g vi s c e r a l p ou c he s a s di r e c ted fo r the n in th n e rve a n d de te rm in e

the dis tr i b u ti on o f the vi s c e r a l b r an ch o f the va gu s Wi th the wa ll s o f the si n u s .

w ell sp r e ad Op en n o te the fou r b ran che s cr ossin g the oo r of the s in u s to the .

vi s cera l pou che s Di ss e c t ou t eac h o f the s e an d o b se r ve th a t e ach b e ar s a gan g

.

li on b eyon d whi ch it divide s in to th r ee b ran ches : an an te ri o r p r e tr emati c

,

b ran ch a middle ph ar yn ge al b ran ch an d a pos te ri o r p o s t tr ema ti c b r an ch

, ,
-
.

The p h aryn geal b r an ch s eems to b e m i ssin g in the sk a te As in the cas e of the .

nin th n e r ve ,
the p r e an d p os t tr e m a tic b r an c he s e mb r a c e the Vi s ce r a l p ou ch
-
,

w hi ch lie s b e tw een them ; all th r ee b ra n che s h ave the s am e fu n c tio n s as des crib ed
fo r the n in th n e rve We th u s see th a t the va gu s n e r ve su pp lies the r emainin g
.

vi sc e ral arche s b egi n nin g wi th the fou r th an d the r emainin g vi s c eral p ouches
, , ,

b egi n n i n g wi th the thi r d Af te r supp l yin g the gill app ar a tu s the vis cer al b r an ch
.

o f the v a gu s p ass e s o n i n to the p e r i car di a l an d p le u rop e r i to n e a l cavi tie s su pp l y

'

in g the he ar t dige s tive tr ac t an d o the r vi s c er a

, ,
.

Ou r s t u d y of the cr an i l n erves s h o w s th a t th ey b ea r a d e n ite rel a tio n to the o rig in al

a

segm en t a tio n o f the h ea d a n d to t he g ill app a r at us T he fth s even th n in th an d ten th .

, , ,

c ran ia l n erves ar e g ill a ch n erves -

eac h as s o c ia ted w ith o n e o r mo r e p ar ticu l ar vis c er al
r ,

ar c h es . I n t he evo l u tion o f the ver teb r a tes eac h n erve co n tin u es to s u pp l y its p ar ticu l ar
ar c h o r ar c h es a n d all d er iva tives th ereo f S in c e the mu scl es of the vis cer al ar ch es are
.

visceral mu scles the mo tor comp on en ts of the gil l arch n erves in all ver teb rates ar e vis ceral
,
-

mo to r in n ature T he s o matic mo tor co mp o n en ts for the fth an d s even th n erves as a l r ea d y

.
,

s t a ted ar e s ep ar a ted in to s p ecial n erves

,
the eye mus c l e n erves There are n o s oma tic
,
-
.

mo to r el emen ts for the n in th an d t en th cr an ial n erves s in ce the myo to mes w h ich S h ou l d ,

give r is e to vol un tary mu s cl es for th eir segmen ts ar e d ege er ate in all ver teb ate s ab ove n r

c yclos tomes ( Fig . 69, p .

5 The occipital , hypob r an chia l , an d r s t s pin al n e rve s

. car ef u ll y
Ve r y
exp os e the spi n al cor d po s te r io r to the m edu lla by sh avi n g a way the car til age
o f the n e u r a l a r c he s in thi n s lic e s On the d o r so la te ra l su r fac e o f the co r d n o te
.

the li ttle sw ellin gs ,

the dors al or s pin al d by the g g
a n lia , a t ta c hed to the co r
sen s ory o r dor s al r oot In the sk a te the sp i n al gan gl i a a r e el on ga ted
. B e tw een .

the r s t sp in al gan gli on an d the roo t of the v a gu s n o te two o r th r ee sm a ll r o o ts

sp r in gi n g f r om the s ide o f the m ed u ll a Thes e ar e the occipital n er ves They . .

in n ervate som e mu s cles of thi s r egi on an d als o help to fo rm the hyp ob ran chial
n e r ve de s cr i b ed b el o w a n d con tri b u t e to the c e r vi co b r a c hi al p lexus
, On pr ess .

in g the sp in a l co r d to o n e side the ven tr al o r motor roots o f the sp i n al n erve s will

b e s een a ri si n g from the ven tr o l a te r al r egi on o f the co r d a t the s ame level as ,

the occip ita l n e r ve s T he ven tra l r o o ts a r e form ed by the u nion of s eve ra l sma ll
.

r o o tle ts co mi n g f r o m the c o r d The m os t an teri or ven tr a l r oo ts a r e si tuate d

.

a n te r i o r to the d o r s a l roo ts w hi c h b el o n g to the s am e s egm e n t .

In the d og sh the union o f d o r s al an d ven tra l ro o ts to fo rm a sp i n a l n e rve

is n o t easy to fo llo w I t may u su a l l y b e s een by ca r efu ll y p arin g d o wn the ca r ti
.

lag e a lo n g the side o f the sp in al co r d In the s kate the un ion is e asily fo llowed ;
.
3 1 6 LAB ORATORY MANUAL FOR VERTEBRATE ANATOMY

the r o o ts hr ou gh the car tila ge a t the side o f the cor d an d u n i te as they

p a s s t
exi t fr om the ca r til age A large n u mb e r o f the mo s t an te ri o r sp in al n erves
.

( r e ally the ven tr al r am i o f the sp in al n e rve s the d o r s al r am i b ein g very s len de r

,

in the S k ate) a re then s een to unite to fo rm the ve ry l a rge n e r ve o f the b r a chi a l

p le xu s p r evi ou s l y n o t ed .

The hyp obr anchial n erve is a tru n k fo rm ed by con tri b u tion s fr o m the o c cip i ta l
n e rves an d the r s t sp i n a l n e rve s In the d og s hes it may b e l o ca ted as fo ll o ws
. .

In ser t on e b lade of the s cissors in the an gle of the jaw an d cu t b ack across the
gill s li ts th rou gh to the s ide of the esop ha gu s as was don e in an e ar ly s tage o f
the di ss e c ti on (if the s am e sp e cim e n is s till b ein g u s ed t hi s cu t will a l r e a d y h ave
,

b een m a de) Op en the ap thus form ed an d exp o s e the ro o f o f the m ou th

. .

M ake a l on gi tu di n a l cu t thr ou gh the mu cous m emb ra n e o f the r oo f in the medi an

d orsal lin e S tr ip the m emb ran e l ater a lly car ryin g wi th it the fr ee d o rs al en d s
.

o f the gill ca r t il a ges w hi c h will b e r e a dil y l o ca ted j u s t l a te r a l to the m id d o r s a l -

lin e .The vi s cer a l b r an ch of the v a gu s n erve which was a lr e a d y s ee n f r om the

,

o the r s ide is n ow exp o s ed

,
I t lie s a l on g the thin ve n tral w al l o f the an te ri o r
.

ca r di n a l s in u s E me r gin g f r om the mu scle n ow exp o s ed in the r oo f o f the mo u th

.

will b e s een the ven tr a l r ami of the sp i n a l n er ve s I n the sp in y dog sh two o f .

the s e ( the y app e ar as o n e b u t will b e fou n d to con s i s t of two on di ss e c ti n g them

towa r d the m ed ian lin e ) p ass o b liq u ely to wa r d the vi s ce r a l b ran ch o f the v a gu s
an d e n te r its s he a th th u s app e ar in g to j o i n it
, The tr u n k they fo rm can ho w
.
,

ever b e r eadil y s ep ara ted from the va gu s T hi s tr u n k is the h yp ob ran chi al

,
.

n e r ve In the smo o th dog sh the se ve r al mos t an te ri or ven tr a l r ami u nite to

.

form a large tru nk the h yp ob ran chi al n e rve T hi s n erve lie s j us t an teri or to the
, .

n e rve s o f the c e r vi co b r a chi a l p lexu s to w hi ch it c o n tri b u te s b r a n c he s I t p as s e s .

to the d or s al s ide o f the vi s ce ra l b r an ch o f the v a gu s (whi ch m ay b e n o ted

p r oc eedi n g t o th e e sop h a gu s ) p os t e r i o r to th e l as t vi s c e r a l p ou ch a n d t u rn
,
i n g
ven trally cou rs e s al on g the oor of the mou th supp l yi n g mu s cles ,
.

I n the s kate the h ypo b ran chia l n e rve le ave s the sp i n al co r d in common wi th
the tru n k o f the b ra c hi al p lexu s L o ca te thi s tru n k aga in in the an te r io r wa ll
.

o f the p leu r op e ri ton e a l ca vi ty d o r s al to the p le u r op e ri ton e u m an d j u s t b ehin d

,

the ca r til age o f the p e c to ral gir dle The e n ti r e tru n k wi th the ex cep ti on of
.

on e b r an ch p as ses o u t do rsal to the c a r til a ge to the p e c to r a l n This on e .

b r an ch the h yp ob r an chi a l n erve tu rn s fo rwar d an d is di s trib u ted to the mu s cles

, ,

o f the oo r o f the mo u th .

The oc ci p i tal an d hyp o b ran chi al n e rve s ar e p ro b ab l y the ho m ol o gu e s o f the

tw e l f th o r h ypo gl o ss al n e rv e o f am nio te s h a vin g b ee n take n in to the craniu m
,

in th o s e forms T he mu s cles su pp lied by the hyp o b ran chial n erve are b elieved
.

to b e de r ived f ro m the mos t po s t e ri o r o f the he a d m yo to m e s an d in highe r fo rms ,

b e co m e the in tr in s i c mu s c le s o f the to n gu e .

6 The ven tr al a s pe ct of the b r ain

.
Car efu l l y f r ee the b rain f r om the chon
.

d ro cr an ium F irs t cu t thr ou gh the o l fa c tory tr a cts an d gen tl y lift the an te

.
3 1 8 LAB ORATOR Y MANUAL FOR VE RTEB RAT E ANATOM Y

fro m its shap e the foss a r homb oidea B isect the brain by a median sagi ttal cu t .
.

E x am ine the cu t s u rfa ce bes t u nder wa ter ,

F ro m the fo u r th ven tri c le a narrow.

p assa g e the,
a qu e d uc t of t h e brain e x t ends an t e riorly I t c,o mm u ni c a t es wi th the .

c avi ty of the c erebell u m the cereb el l ar ven tr icl e and the cavi ties of the o p ti c
, ,

lobes the optic ven tricl es The thick oor of the midbrain ven tral to the aqu edu c t
,
.

form s the cer ebral pedun cles The aqu ed u c t op ens in to the cavi ty of the d ien cep ha
.

lon the third ven tri cle The roof of the dien c ep halon is very thin and consis ts of a
,
.

ch o r ioid (vas cu lar) pl exu s whi ch is folded in to the c avi ty The an terior p ar t of this .

roof in the dogs hes ex tends dorsally in to a s a c the par aphysis res ting agains t the , ,

telen c ep halon I t is regarded as p ar t of the la tter

. P os terior to the p arap hysis .
,

a thin transverse p ar ti tio n the vel u m tr an sversu m is seen in the roof of the
, ,

dien cephalon p ar ti cu larly in the sp iny dogs h This m arks the dorsal bou ndary
,
.

be tween dien cep halon and telen c ephalon The small thi ckened region of the .

dien c ep halon j u s t in fron t of the an terior end of the op ti c lobe is the ha ben u l a ,

a sm ell c en ter F rom the habenu la a slen der p ro cess the pin eal b ody may be
.
, ,

seen in favorable sp e ci men s ex tending dorsally j u s t ba ck of the p arap hysis

, , .

The en tire roof of the diencep halon in cl u din g chorioid p lexu s habenu la an d , , ,

ineal body is na m ed t h e epithal a mus an d is c on c ern ed ch i ey wi t h s m ell The

p , .

la teral walls of the dien cep halon cons ti tu te the thal amu s an i mp or tan t correla ,

tion c en ter for vario u s body senses The ven tral p ar t of the die n cep halon is
.

named the hypothal amu s I t co n sis ts of the i n f u ndib u lu m in clu din g the inferior
.
,

lobes the hyp op hysis and the mammillary bodies The former sho u ld b e iden ti
, ,
.

ed on the se ction The mammil lary bodies are the thi ckened p ar t of the ven tral
.

wall above the vas cu lar sa c The hypo thalamu s is con c erned wi th smell and
.

tas te No te tha t the cavi ty of the third ve n tri c le ex tends in to all p ar ts of the
.

hypo thalamu s The third ven tri cle conne c ts by a p assage the for amen ofMon r o
.
, ,

or in terven tricu l ar for amen wi th the cavi ty in ea c h half Of the telen cep halon
, .

( I n th e ska t e th e t ele n c ep halon is solid )

,
These t wo c avi ti es are n am ed the
.

rs t and s econ d or l a ter al ven tri cles They ex ten d o u t in to the olfa c tory b u lbs
.

thro u gh the olfa c tory t ra c ts Cu t in to the telen cep halon to see its ven tri cles
. .

Draw the sa gi tta l se c tion .

c . THE NE R VOUS S Y S TE M AND S E NSE OR ANS OF NE CTUROS G

I . The nerve ss pi al n .
The
spinal nerves are bes t fou n d as follows M ake .

a longi tu din al cu t along the side of the body below the la teral line Cu t thro u gh .

the ex t ern al an d in ternal obliqu e mu s cles a n d sep ara te this m ass of mu s cle fro m
the thi n layer of tran sverse m u s cles lyin g n ex t to the c oelo m The ven tr al r ami .

of the sp in al nerves wil l now be seen ru n nin g be tween the obliqu e an d trans verse
mu s cles alo n g the m yosep ta and s upp lying the hyp ax ial mu s c les
, ,Tra ce o n e .

of these toward the ver tebral c olumn cu tting away mu s cles from its cou rse ,
.

I t lies j u s t behind the rib whi ch m ay be cu t away The nerve m ay be tra c ed .

up to the ver tebra where it is im bedded in an ora n ge c olored m a t erial On -

.
 CO MPARA I VE ANATOM T Y OF THE NE R VOUS S YSTE M 3 9
1

c arefu lly c learin g this away the dors al gan glion of the spinal nerve will be fo und
imbedded in it I t is a rou nded brownish body from which sp ri n g two nerves
.
,

the ve n tral ramu s j u s t followed and the s maller dors al r amu s whi ch s u p p lies the
ep axial mu s cles .

2
. The l imb pl e x i The ven tral rami of the spi n al nerves form a plex u s
.

for ea ch li mb the mo tor nerves for the li mb mu s cles arisin g from this p lexu s
, .

The b rachial p lexu s is lo ca ted as foll ows Make a cu t in the ven tral body wall
.

j u s t m edial to the base of the fore l imb S ep ara t e the p e c toral and shou lder .

mu s c les from the s ternohyoid mu s cle The bra chial p lexu s is then easily seen
.

ru n n i n g p os terior to the s capu la I t consis ts of the ven tral rami of three sp inal
.

nerves (3 4 and 5) whi ch have cross conne c tions wi th ea ch o ther B eyond the
, ,
-
.

p le x u s n
, erves p ro c eed in t o t he fore li m b .

The l u mbos acral p lexu s is lo ca ted by cu tti n g thro ugh the skin and bo th layers
of obliqu e mu s cles lon gi tu dinally j u s t dorsal to the base of the hin d li mb On .

sep ara ting the Obliqu e fro m the tran sverse mu s c le layers the p lexu s is exp osed .

I t c onsis ts of the ven tral ra mi of three ( so m e tim es fo u r) sp i n al nerves The .

an terior one of the three gives o a sli gh t bran ch to the nerve nex t p os terior to '

it so m e ti m es re c eives a c on trib u tion fro m the ramu s an terior to it and as the

, ,

cr u r a l nerve p asses in to the li m b

, The middle of the three nerves is mu ch the
.

s tou tes t an d af ter re c eiving con trib u tions from the n erve nex t p os terior to it
,

en ters the li m b as the ischiadic (s cia ti c) nerve .

3
. Th e s e n s e or g an s of th e he a d O wi n g t o.the s m all S iz e of the eye and its
similari ty to tha t of o ther ver tebra tes it will n ot be inves ti ga ted .

a ) L a ter a l l in e sys tem: L a t eral line sense or gans are p resen t in Nectu r u s b u t

imp ra c ti cal to nd in gross disse c tion They are si tu a ted alon g lin es similar to
.

those of shes .

b ) Nose: P robe in to the ex ternal naris and fo l low yo u r p robe wi th the cu t ,

op ening the en tire nasal p assage to the in ternal naris No te the folds or olfactory .

l a mell ae in the in terior of the p assage Unl ike the co n di tion in shes the
.
,

olfa ctory sac op ens in to the oral c avi ty and has bo th olfa c tory and resp ira tory
fu n c tio n s .

c) E a r : The ear as in shes c onsis ts of the in ternal ear o nl y im bedded in ,

the o ti c re gio n of the sk u ll E xp ose the dorsal s u rfa ce of the sk u ll by cl eanin g

.

away the mu s cles L o ca te the o ti c c ap su les one a t ei ther side of the p os terior
.
,

en d of the sku ll Cau tiou sly shave away the car tilage here an d lo c a te the three
.

semi cir cu lar d u c t s an d ves tib u le as done in the dog sh There are three of the .

,

form er an ter ior ver tical pos ter ior ver tical and horizon tal ea ch wi th an ampu l l a
, ,

all arran ged ex a c tly as in the dogsh The ves tib u le is divided indis tin c tly in to
.

a dorsal u tricu l u s from which the du c ts sp ring an d a ven tral s accu lus The
, ,
.

la tter con tains a c rys talli n e mass the otolith , .

4 The d or s al a s p e ct of the b r ain

.
R em ove the roof of the sku ll
. Thi s is .

b es t done by s tripp in g it off in slivers wi th a knife Af ter the b rain is reveal ed .

3 20 LAB ORATO R Y MANUAL FO R VE R TEB RATE ANA OM T Y

s tu dy its dorsal su rfa c e The brain is covered by a m embran e the pr imitive

.
,

men in x whi c h is m ore or less divisible i n to the pia ma ter a d eli ca t e p i gm en ted
, ,

m em bran e adheri n g to the brain and an o u ter d u ra m a ter whi ch is sep ara ted
, ,

from the sku ll by the peridu r al sp a c e .

The mos t an terior po r tio n of the brain c on sis ts of the two elo n ga ted lobes of
the tele n cep halo n These are very indis tin c tly su bdivided in to an terior olfactor y
.

l ob es and p os terior cer eb r al hemis pheres The olfa c tory lobes in c lu de the b u lbs
.

a n d t ra c ts of the d o g s h B e tween and behin d the p os terior en ds of the tel

.

en cep hali c lobes is a thin roof con tain in g a chorioid pl exu s The an terior p ar t .

Of thi s bears a p ro c ess the pa raphysis p roje c tin g forward in the g roove be tween
, ,

the two c erebral hemisp heres I t is re garded as a p ar t of the telen cep halo n
. .

The very sm all region p os terior to thi s is the roof of the dien cep halon I t bears .

a dorsally p roje cti n g p ro cess the pin al b ody or epiphysis which lies imm edia tely
, ,

p os t erior to t h e p ara p hysis P os terior

. t o t h e dien c e p halon are th e t wo elo n g a ted
optic l ob es rep resen tin g the dorsal p ar t of the m esen c ep halo n
,
B ehin d this is .

an o ther chorioid p lexu s a dark p i gm en ted m em brane Thi s may be re m oved

, . .

There is then revealed the triangu lar cavity of the four th ven tr icl e The an terior .

en d of this cavi t y is overh u n g by a narrow shelf a t the p os t erior m ar gi ns of the

op ti c lobes This shelf is the cer ebell u m The remai n der of the brain con sis ts
. .

of the medu ll a obl on gata whi ch in closes the cavi ty of the fo u r th ven tri cl e The .

m ed u lla ex tends forward at the sides of the op ti c lobes as p roje c tio n s the grea ter ,

p ar t of wh i c h c ons i s t s of th e a cu s tic o l a te ra l ar e-
a The walls .of t h e m ed u lla are
divided i n to dorsal or s en s ory p or tion s and ven tral or motor p or tio n s the la tter ,

formi n g a broad band on ei ther side of the m edian ven tral groove The fu r ther .

s u bdivisions of these are n o t eviden t in Necturus

'

Draw the dorsal View of the brain .

5 The c r ani al n erve s

. .

a ) The rs t or olf a ctor y n erve: Th i s is a s to u t band p assin g fro m the olfa c tory
s a c to the a n terior end of the t elen c ep halon .

b ) The second or optic n erve: This s m all nerve m ay be seen in the oor of
the c ranial c avi ty by p ressi n g the t elen cep halon to o n e side I t p asses obliq u ely .

c a u dad to the ve n t ral s u rfa c e of the die n cep halon .

c) The eye mu s cl e n er ves : The thir d or ocu l omotor fou r th or tr och ,

l ea r an d ,

s ix th or a bducen s nerves are so s m all in Nectu r us as to be s c ar c ely dis c ern ible in

g ross disse c t io n They

. ori g i n a t e fro m t h e sa m e re g ions of t h e brain a n d s u pp ly
the s a m e eye mu s cles as in elas m obran chs .

d) The fth or tr igemin u s n erve: The fth n erve is the large tru nk arisi n g
from the side of the an terior end of the m edu lla Tra ce it ou t thr ou gh the .

skul l I t p asses in fron t of the o ti c cap su le and immedi a tely en ters a large
.

g gan lion t
,
h e s em i l u n ar or G a ss er i an a
g g n lion of t h e t ri g e m i n u s F ro m t h e .

g gan lion t hree lar g e nerves are g ive n o f f th e op h th a l m,

ic m a x il l ar y a n d ,
m an ,

dibu l a r bran ches The form er c orresp ondin g to the deep op h thalm i c of shes
.
, ,

p asses forward th r o u g h the q u adra t e c ar tila g e a n d r u ns forward a n teriorly alo n g

3 2 2 LABORA OR T Y MANUAL FOR VE RT B RA E ANATOM
E T Y

a ) S pin al n erves a n d l imb pl exi: Carefu lly exp ose the sp i n al nerves of the
t ru n k avoi din g inj u ry to
,
the sym p a the ti c sys te m These nerves , c alled the dor
.

s al sp i n al nerves ru n alon g the su tu res be tween the cos tal p la t es o f the cara
,

p a c e. In m os t c ases ea c h c o n sis ts of two bran ches a small er dors al r amu s

,

a n d a larg er ven tr al ra mu s . On tra ci n g these toward the ver tebral c olu mn they
will be fo u n d to co m e fro m a large ganglion the dors al or s pin al gan glion S i tu a ted
, ,

in con ta c t wi th the cen ter of the cen tr u m .

E xp ose the b r achial pl exu s in the dep ression be tween the ne ck an d the dorsal
end of the s capu la I t is general ly form ed by the c ross u nions be tween the
.
-

Ven tral rami of the las t fo u r cervical sp i n al nerves and the rs t dorsal sp in al
n erve . The las t n am ed nerve may be iden tied as the one in fron t of the rs t
-

typ i c al rib ( really the se cond rib ) . The fou r cer vi cal nerves form a co mp lex
ne twork on the su rfa ce of the shou lder mu s cles F ro m thi s ne twork the large
.

median nerve p ro ceeds along the an terior s u rfa c e of these m u s cles and the
smaller u l n ar an d r adial nerves alo n g the p os terior su rfa c e ; the radial is the m os t
dorsally si tu a ted one The r s t dorsal n erve sends a bran c h near its gan glion
.
_

to the bra chial p lexu s and is then dis trib u ted to the c arap a c e j u s t p os terior to
the fore lim b .

The nex t six dorsal nerves are si milar to the r s t des crip tion given The .

ven tral ram i of the eigh th nin th and ten th dorsal nerves toge ther wi th the two
, ,

s acr a l nerves form the l u mb osacr al pl exu s for the h i nd limb This lies on the
.

medial su rfa c e of the mu s cles coveri n g the ili u m and is fo u nd by carefu lly sep ara t
in g the ili u m wi th its mu s cles fro m the median region . The bran ches fro m the
thr ee dorsal n erves u ni te to a sor t of kno t fro m which several nerves p ro ceed to
the an terior p ar t of the l eg . The two sa cral nerves re c eivin g also a con trib u
,

tion fro m the t en th dorsal nerve u ni te to form a larg e t ru nk the sciatic nerve
, , ,

si tu a ted am o n g the mu s cles on the p os terior side of the l eg .

There is a p air of caudal sp inal nerves c orresp on din g to ea ch cau dal ver tebra ,

b u t these n eed n o t be looked for . There are nin e p airs of cer vical nerves whi ch
will be fo u nd by lookin g in the ne ck a t the sa m e level as the level of em ergen ce
of the nerves of the bra chial p lexu s .

b ) The sympa thetic sys tem: L o ca te the vagu s nerve in the ne ck I t is the
.

c o n sp i cu o u s white c ord run nin g alon g t he side of the ne ck . Tra c e it pos teriorly .

The symp a the ti c t ru nk is bou nd wi th it b u t a t abou t the level of the r s t nerve

of the bra chial p lexu s sep ara tes fro m the vagu s and en ters a swellin g or gan glion ,

the middl e cervical gan glion .The symp a the ti c c ord p ro ceeds dorsally fro m this
a
g g n lion a n d lie s on t he ven t ral s u rfa c e of th e bra c h i al p le xu s where it p rese n t s
two s u c cessive swellin gs whi ch t og e ther c on s ti tu t e the infer ior cervical g angl io n
,
.

Observe bra n ches fro m these gan gl ia . The symp a the ti c c ord p asses to the
a
g g n li on of th e r s t dorsal s p i n al n erve t o w h i c h it s o wn an
g g lion is f u sed I t .

then p ro c eeds as a deli ca te whi te c ord a cross the se cond rib and ag ain form s a

g gan lion wh i c h is f u sed to th e g ga n lio n of th e se c o n d dor s al nerve The

. sym p a
the ti c c ord then p ass es m ore ven trally lyin g on the sid e of the lon g n eck mu s c les
,
.
 COM P AR A I VE ANATOM T Y OF THE NE R VO US S YST E M 3 3
2
0

F ollow it here an d n o te the ganglia which it bears a t in tervals and the bran ches
from these ganglia No te p ar ti cu larly the bran ches be tween the symp a the ti c
.

g an g lia and the adja c en t Sp i n al ga n g lia These bran ches co n s ti tu te the r amu s .

commun ican s and c onsis t of the vis c eral m o tor and vis c eral sen sory bers p assin g

be tween the symp a the ti c an d c en tral nervou s sys tem s The gan glia and .

branches of the symp a the ti c are p ar ti cu larly no ti ceable in the u ro gen i tal region .

Draw spinal nerves p lexi and sym p a the ti c sys tem as far as seen
, , .

2. The s en s e or g an s of the he a d .

a ) The n as al cavities : The ex ternal nares lead in to wide cha mbers the nasal ,

c avi ties .Cu t o ff the ex ternal nares and the roof of the sku ll p os terior to them ,

thu s revealing the nasal c avi ties They are sep ara ted by a median s eptu m
.
,

p ar tly bony F.ro m th e ven t ral re g ion of th e se p tu m a c ons p i cu o u s fold p roje c ts

in to the nasal cavi ty On the p os terior wall of the nasal c avi ty is a sli gh t pro
.

je c ti on ,
a con c ha or tu r b in a l P os t erior to t
. his t he n asal c avi t y c o n ne c t s by a
p assa g e wi t h th e roof of th e m o u t h c avi t y th e nasal c avi ti es t h u s serving as ,

respira tory p assages .

b ) The eye: Al tho u gh the eye is s m all it can be disse c ted wi th a li ttle care
'

I ts p ar ts are very si milar to those of the sh eye Make an in cision throu gh the .

skin arou nd one eye and wi th the bone s cissors remove the S ku ll dorsal to and
be tween the eyes The two eyes are seen to be c lose toge ther sep ara ted by a
.
,

m edian m em brano u s in teror bital septu m Near this sep tu m o n ea ch side ru ns .

an ar tery On the an terior dorsal su rfa c e of the eyeball is a gland the Harderian
.
,

g land . O ver th e p os t erior and ven t ral s u rfa c e o f t he eyeball e x t ends th e mu c h

larger l acrimal or tear gland R em ove these glands thu s exp osing the s u rfa ce
.
,

of the eyeball and the eye mu s cles E x tending from the in terorbi tal sep tum to .

the dorsal s u rfa ce of the eyeball is the s u per ior ob l ique mu s c le P os terior t o .

this and i n ser ted in the eyeball near it is the s u per ior r ectu s B e tween and ven .

tral to these two is the in ter n al rectus P assin g above the in tern al re c tu s are.

two nerves the trochlear to the s u p erior obliqu e m u s c le and the ophthal mic
,

branch of the t rigeminu s L oosen the eyeball ven trally and raising it as far as
.

p ossible e x a m ine th e v e n t ral s u rfa c e The an t erior p ar t of t his .s u rfa c e is c overed

by a a t mu s cle the pyr amidalis whi ch o ri gin a tes on the eyeball and p as ses to
, ,

the eyelids and ni c ti ta tin g m embrane R emove this and c lean the ven tral .

su rfa ce of the eyeball The inferior oblique an d infer ior rectu s mu s cles are then
.

seen c onvergin g to their inser tions on the ven tral s u rfa ce of the eyeball The .

ex ter n al r ectu s is p os terior to them .

R emove the eyeball a n d op en it by cu ttin g Off its dorsal side I d en tify the .

c oa ts of the eyeball the lens the c avi ties of the eye an d the two h u m ors as in
, , ,

the elas m obran c h eye as the s t ru c tu re is p ra c ti c ally ide n ti c al wi th the la t ter

, .

No te however the di er en ce in the shap e of the len s of the tu r tle eye

, ,
.

c) The ear : The ear c o n sis ts o f two p ar ts a middl e ear a n d an in tern al ear , .

T he form er is lo c a ted p os terior to the an gle of the jaws in tern al to a c ir cu lar area
3 4
2 LAB ORATOR Y MANUAL FOR VE RTH RATE ANA OM T Y
Q

of skin R emove this p ie ce of skin an d nd benea th it a s maller car tilagi n ou s

.

cir cu lar p la t e the tympan ic memb r an e or eardr u m

,
M ake a cu t arou n d the .

m argin of this and c arefu lly r ai se it Atta ched to its in tern al s u rfa c e p os t erior
.
,

to the c e n ter is a rod S hap ed bone th

,
-
e col u mell a whose inner end is fas tened to
, ,

the wall of a larg e c avi ty Thi s c avi ty is the tympan ic cavity or cavity of the
.

middl e ear . I t is an eva gina tion fro m the rs t vis ceral p o u ch Ven tral to the .

inn er en d of the c olu mella is a sli t bou nded by raised li p s This sli t is the op en .

in g of the au ditor y tu be c o n ne c ting the p haryn geal cavi ty wi th the cavi ty of

h e middle ear an d r ep resen ti n g the s talk of the eva gin a tion by whi ch the la tter
t
was form ed Consid erably i n ternal to the poin t of a tta ch men t of the in ner end
.

o f the col u m ella lies the i n t ern al ear I t will be more deni tely lo ca ted la ter
. .

I t is si milar in s tru c tu re to the in tern al ear of elas mobran chs .

3 D ors al a s p e ct of the brain

. R e move the roof of the sku ll and exp ose the
.
-

brain The brain is covered by a tou gh membra n e the du ra ma ter On cu tti n g

.
, .

carefu lly thro u gh this a m ore d eli c a t e m embrane the pia ma ter will be fo u n d , ,

adhering to the brain I t is mor e or less pi gmen ted and con tains nu merou s blood
.

vessels S trands c ross be tween the two m embran es The two m em bran es are
. .

derived b y the splitting of the ori ginal single m embrane of the brai n the p rimi tive ,

m eninx of shes The s p a c e be tween the du ra ma ter and pia ma ter is the s u b
.

dural sp a ce ; be tween the form er an d the sku ll the per idu r al s p a ce R emove the
, .

d u ra m a ter fro m the dorsal su rfa ce of the brain .

The brain p ossesses the sam e divisions as in the p re ceding form s b u t the ,

rela tive p rop or tion s of the p ar ts are som ewha t al tered The an terior en d co n .

sis ts of the two olfactory l ob es whi ch in clu de the olfa c tory b ulbs and tra c ts seen
,

in the elas m obran ch ; the whole olfa ctory app ara tu s is eviden tly mu ch red u ced
in the tu r tle as comp ared wi th the sh P os terior to the olfa c tory lobes and .

sep ara ted from them by a sli gh t g roove are the two cerebral hemispher es rela tively ,

m u c h larger than in the el as m obran c hs and Nectu ru s B e tween the p os terior .

en ds of the two c erebral hemisp heres is the dien cephal on its roof co n sis tin g of ,

a chorioid pl exu s which p roj ec ts dors ally as a sac tha t adheres to the du ra ma ter
,

and is gen erally torn off in rem oving the la tter This s ac co n sis ts of the .

p a r a h
p y s is and th e e i h
pp y s is When it is re
. m oved the dien c e p halon is see n

as a dep ressed area p os terior an d ven tral to the c erebral hemisp heres P os terior .

to the die n cep halo n are the two ro u nded optic l ob es belo n gi n g to the midbrai n ,
.

B ehin d them is the cer eb ell u m s m all er than in elasm obran chs b u t larger than
,

in Amphib ia P os terior to the c erebell u m and p ar tly overhun g by it is the me

.

du l l a ob l on ga ta I ts thin dorsal roof con tai n s as u su al a chor ioid pl exu s

. On .

removi n g the roof of the m ed u lla the c avi ty of the fou r th ven tri cle is revealed
,

and the c erebell u m is then seen to p roje c t like a roof over the an terior p ar t
of the medu lla The dorsal rim s of the m edu l la consis t as u su al of the s omatic
.

s en s ory c ol u mn s a n d on raisin g the c erebell um the an terior en ds of these co l

,

u m n s whi c h a re au di tor y in na tu re are seen to b e conne c t ed wi th the cereb el

3 2 6 LABO RATOR Y MANUAL FOR VE RTE B RATE ANATOM Y

the sku ll and nally join s the semilu nar gan glion F o llow the max illary nerve
, .

p os t eriorly B .esides th e bra n c h below the eyeball already no ted there is a

bran ch in the oor of the orbi t ru nn ing obliqu ely forward These two bran ches .

u ni te to for m the main t ru n k of the m axillary nerve a t the os t erior end of the
p
orbi t Trace thi s nerve p os teriorly among the mu s cles to where it p ier ces the
.

sku ll At this p oin t it is joined by the man dibu l ar bran ch of the tri geminu s
. .

Tra c e this la terally Af ter bran chi n g in to adja cen t mu s cles the mandib u lar
.
,

nerv e p ro ceeds ven trally and en ters the lower jaw M andib u lar and maxil lary .

bran ches p ass toge ther through a foramen in the sku ll and c onne c t wi th the
semilu nar gan glion .

f) T h e f ac ia l an d a u di to r y n erves : Thes e arise to ge ther fro m the side of the

m ed u lla j u s t behin d the roo t of the t ri gemi n u s and i mm edia tely sep ara t e in to
an an t erior fa cial nerve and a p os terior au di tory The la tter is dis trib u ted to .

the in t ern al ear Thi s is si tu a ted in the sku ll opp osi te the a cu s ti co fa cial roo t
.
-
.

This p ar t of the sku ll may be broken op en wi th the bone for cep s The semi .

cir cu lar d u c ts a mp u llae and ves tib u le of the in t ernal ear will be n o ted
, , The .

au di tory nerve will be seen bran ching a mong these s tru c tu res The facial nerve .

p asses t hro u g h th e an t erior p ar t of th e ear c a p s u le and wi l l b e seen a g ain

la ter .

g) T he gl oss o p ha r y g
n ea l n e r ve : T hi s arises by a s m all roo t fro m t he m ed u lla
i mmedia tely p os terior to the a cu s ti co fa cial roo t and p asses ou t thr o u gh the -

p os t erior p ar t of th e ear c a p s u le .

h) The vagu s the spin al access ory an d the hypogl ossal : The vagu s and sp in al
, ,

a cc essory ( eleven th ) nerves arise toge ther by a nu mber of roo ts fro m the side
of the m edu lla p os terior to the p re c edin g nerve the more an terior roo ts belon g , s

in g to the va gu s and the p os terior o n es to the a c c essory On cu tti n g thr o ugh.

these roo ts the m ore ve n trally si tu a ted roo ts of the hyp o g lossal ( twelf th ) nerve
will be seen The three nerves p ass ou t fro m the S ku ll close toge ther
. .

i) The a bdu cen s n erve: Cu t thro u gh all of the n erve roo ts on one side of the
brain and til t the brain toward the opp osi te side The abdu c en s nerves will be .

seen sp ri n ging from the ven tral su rfa ce of the m edu ll a a t abo u t the sam e level
as the a cu s tico fa cial roo t -
.

The seven th nin th ten th an d twelf th nerves may be tra ced far ther as
, , ,

follows Tu rn the head ven tral side up and rem ove the skin an d s up er cial
.

m u s cles fro m the hyoid app ara tu s L o c a te the an terior and p os terior horns of
.

the hyoid On the side of the ne ck n ear the dorsal end of the an terior horn and
.
,

p os t erior to i t t h e hy
,
o
p g lossal n erve will be see n e m er i
g g n I t bra n c hes in
. t o
the m u s cles over the an terior horn and sends a bran ch forward in to the to n gu e
mu s c les Very near the p oin t of e m erge n c e of the hyp o glossal b u t si tu a t ed m ore
.

deeply and n earer to the car tilage of the an terior horn will be fou n d the glosso
p hary n g eal nerve I t r u ns be. t ween th e tw o hor n s toward the m e dian ve n t ral

lin e an d su pp lies adja c en t mu s c les an d li n i n g of the m o u th c avi ty L a te ral .

 COMPARATI VE ANATOM Y OF T HE NE R VOUS S YSTE M 3 7
2

to t hese nerves a bran ch of the fa cial will be fo u nd crossing the an terior horn
,

near its dorsal end and p assing in to the mu s cles lyin g along the p os terior border
of the m an dible .

M ake a m edian lo n gi tu dinal in cision throu gh the whole oor of the m o u th

and p haryn geal c avi ties and op en the two ap s so tha t the roof of these cavi ties
is revealed L o ca te the vagu s ( really vago symp a the ti c) tru nk in the n e ck and
.
-

tra c e it a n teriorly to its p oin t of exi t from the sk u ll re movin g the m u co u s mem brane ,

from the roof of the p haryngeal cavi ty The vago symp a the ti c tru nk p asses to .
-

the dorsal side of the hyp o g lossal nerve seen above and there en ters a g anglion ,

the s u per ior cervical g anglion of the symp a the tic F o m this gan g lion n u mero u s r
.

bran ches p ass ou t I n tern al to the hyp oglossal lo ca te the glossop haryngeal
.

nerve the caro tid ar tery bein g si tu a ted be tween the two S li gh tly an terior to
,
.

these will be fo u n d the fa cial nerve as it ex i ts fro m the S ku ll I ts bran ches p ass
,
.

to the m u s cles be twee n the an terior horn of the hy oid and the lower jaw one of ,

them cu rvi n g over the ven tral s u rfa c e of the horn as no ted above The vagu s .

nerve p ro ceeds p os teriorly and su pp lies the hear t an d o ther vis cera I t will be .

no ted tha t the fa cial glossop haryn geal an d vagu s nerves are mu ch red u ced
, , ,

owing to the loss of the la teral line sys tem and the gill app ara tu s No te how .
,

ever tha t these n erves co n tinu e to supp ly the rem ains of the Vis ceral ar ches an d
,

their vis c eral m u s c les .

Add the c ranial n erves to yo u r drawin g of the brain .

5 Ventr al a s p e c t of the b r ain

.
R e m ove the brain fro m the sku ll an d
.

ex am ine the ven tral s u rfa c e On the ven tral su rfa c e of the dien cep halon no te
.

the op ti c chi as m a the infu ndib u lu m j u s t behind this wi th the h yp op hysis p r o

, ,

jec t in g ven t rally fro m th e la tt er N o t e th e roo t s of the abd u c ens

. and hyp g
o o ssa l
n erves arisin g fro m the ven tral s u rfa c e of the m ed u lla .

6 M e di an s a gittal s e cti on
.
Make a median sagi ttal se c tion and s tu dy the
.

cu t s u rfa c e I den tify the fou r th ven tricl e in the m ed u lla the a qu edu ct or p assage
.
,

below the c erebellu m the optic ven tr icl e in the Op ti c lobe the third ven tricl e in
, ,

the dien c ep halo n No te the i n creased si z e of the dien cep halon as comp ared
.

wi th the elasm obran ch an d the ba ckward ex ten sio n of the c erebral hem i
,

sp here over the dien cep halon The dien c ep halon is divided in to epithal amus
.
,

thal amu s a n d hypotha l amu s as in the dog sh ea ch in clu di n g the p ar t s p revi

, ,

ou s ly e n u m era ted No te tha t the cerebral hemisp here p resen ts a sol id medial
.

wall called the septu m Cu t in to the roof or p alliu m of the hemisp here No te
,
. .

its cavi ty the l a teral ven tr icl e an d the larg e m ass p ro tru ding from the oor in to
, ,

the ve n tri cle ; this m ass is the corpus s tr iatu m .

The fu n c tion s of the p ar ts of the tu r tl e brain are similar to those s ta ted for
the elas m obran ch brain The c erebral hemisphere is s till largely olfa c tory
.
,

al thou gh its la teral su rfa c e is beginning to ass u m e the fu n c tions chara c teris ti c
of the m amm alian h emisphere .

D raw the se c tio n .

3 2 8 LAB ORATOR Y MANUAL FOR VE RTE R ATE ANATOM Y

E . TH E NE R VOU S S Y S TE M AND S E NS E OR G ANS OF THE PI G E ON

I . The i
S p n alnerve s an d the s ympathe ti c s y s te m Carefu lly rem ove the
.

remain in g vis cera from on e half of the t ru n k No te the ve n tral rami of the
-
.

spinal n erves p assi n g la terally along the dorsal body wall be tween the ribs in
the t ru n k re gio n Tra c e them toward the ver tebral colu mn an d no te a t the
.
,

p oi n t s where t hey e m er g e fro m t h e ver t ebrae the g g

a n lia of t,h e s m
y p a thetic

s ys tem lyi n g on the sp i n al nerves a n d the deli ca t e whi te c ords of the sym p a the ti c

sys tem ex ten di n g be tween the gan glia alo n g the sides of the ver tebral c ol u mn .

a ) S pin a l n erves a nd l imb pl exi: I n the ne ck the cervical s p i n al n erves will

be seen by sep ara ting the ver tebral colu mn from the ski n They p ass ou t a t .

segmen tal in tervals The vagu s nerve is the whi te cord which p asses v en tral
.

to the p ro xim al p or tions of the cervi c al nerves .

The ven tral rami of the las t cervi cal nerves to ge ther wi th tha t of the rs t
of the t ru nk form the br achial p lexu s to the wing This is a ne twork form ed .
,

by the union of branches of fou r s tou t nerves whi ch re c eives a s mall bran ch ,

from the su c ceeding n erve .

The nex t ve ven tral rami p ass ou t be tween the ribs F ollo wi n g them is .

the l u mb os acr al p lexu s divisible in to three p ar ts the l u mb ar the s acr al and the
, , , ,

p u d en da l p le x u s The
. l u m bar p le xu s is for m ed by th ree nerves ; fro m it nerves
p ass in t o th e thi g h The sa
. c ral p le xu s arises fro m t he u nion of ve n erves t h e ,

r s t of whi ch is the sam e as the thi rd nerve con trib u tin g to the l u mbar p le xu s .

These ve u ni te to p rodu c e a large t ru nk the s ciatic nerve whi ch p asses alo n g

, ,

the dorsal S ide of the t hi gh be tween the mu s cles and p ro ceeds down the l eg ,
It .

will be fou nd by sep ara ting the mu s cles along the middle of the dorsal s u rfa ce
of the thi gh I t cou rses alon gside the femoral ar tery and vein
. .

The rem ainin g sp inal nerves p os terior to the sa c ral p lexu s form the p u den dal
p le xu s a n d p ass obliq u ely p os teriorly to the tail a n d cloa c al r egion .

b ) The sympa thetic s ys tem: This has already been iden tied on the sides of
the ver tebral c ol umn I t consis ts on ea ch side of a chain of two c ords a n d
.

segmen tal ga n glia On e of the cords p asses ven tral to the head of the rib the
.
,

o ther dorsal to it A symp a the ti c ganglion lies fu sed to ea ch spin al nerve in

.

the tru n k re gion as the la tter e m erg es fro m the ver tebral c ol u mn On s c rap in g .

of f one of these symp a the ti c g an glia the s pin al gan glion bel o n gin g to the sp in al
,

n erve will be fo u n d dorsal t o it At abo u t the mi ddle of the rib bearin g region
.
-

a p lexu s of nerves an d ganglia will be seen ex ten ding ven trally from the m ain
sym p a the ti c cords and su rrou ndi n g the dorsal aor ta and its m ain bra n ches to
the di ges tive tra c t Thi s is the coeliac p lexu s P os terior to thi s region the
. .

symp a the tic c ords are redu ced an d consis t of a sin gle tru n k on ea ch side A .

symp a the ti c c ord a ccom p anies the pu dendal p lexu s an d has a gan glion in the
middle of this p lexu s An teriorly the symp a the ti c c ords p ass a cross the ve n tral
.

side of the bra chial p lexu s havin g g a n g lioni c e n larg e m e n ts o n the la tt er a n d

, ,

then e n ter the ver t ebrar terial ca n als .

 LAB ORA OR T Y MANUAL FOR VE RTEB RA E ANA OMY T T

and form in g the iris in fron t ; the sof t retin a No te the p ecu liar rid ged s tru c tu re .
,

the pecten p roje c ti n g fro m the c horioid c oa t thro u gh the re tina in the m edial
,

wall of the eyeball and ex ten din g to the lens I ts fu n c tion is u n known L oosen . .

the le n s a n d observe tha t it is e n cir c led by a s t ru c tu re form ed chiey fro m the

c horioid c oa t This s tru c tu re which holds the len s in p la c e is c alled the cil iary
.
, ,

b ody. I t is m arked by radia tin g rid g es the cil iary pr oces s es and c o n tain s , ,

mu s c les the cil iary mu scl es whi ch c han ge the shap e an d p osi tion of the lens
, , .

No te the shap e O f the lens n earlya t ex ternally m ore co n vex i n tern ally The
, , .

c ham bers of the eye and the two h u m ors are as in the d ogs h P eel o f
f the iris .

from the corn ea and no te the s ti bony ri n g en ci c li n g the c ornea ; it is c omp osed
,
r

of a nu mber of s cl er otic bones .

Draw showing s tru c tu re of the eye

,
.

c) Th e ear : The ear of birds c onsis t s of three p ar ts the extern al ear the ,

middl e ear and the in tern al ear

,
The ex tern al ear co mp rises the p assage the
.
,

ex tern al au ditory mea tu s si tu a t ed below and behind the eye Cu t in to this on

, .

the sa m e side of the head on which the eye was disse c ted an d nd a t its in ternal
en d a cir cu lar t ransp aren t m em bran e the tympan ic memb r an e ,
Throu gh the .

m e mbran e the col u mell a can be seen ex tending from its in tern al s u rfa c e inwardl y .

R em ove the t ymp a n i c m em brane no ting the c ol u mella adhering to its in tern al
,

su rfa ce The cavi ty of the middle ear is n ow exp osed ; medially and ven tra lly
.

it is c o nn e c ted to the p harn ygeal cavi ty by the au di tory tu be ; p os terior an d

S li gh tly dorsal to it is si tu a ted the in t er n al ear The i n n er en d of the colu mella .

adjoi n s a ti n y bo n e the s tapes which ts in to an op en i n g the fen es tr a oval is or

, , ,

ves tib u li whi c h leads in to the in tern al ear

,
Lo ok for these a t the i n n er end of
.

the c olu m ella Nex t carefu lly break away in s mall p ie c es the s p ongy bo n e
.
,

behind the middle ear Three bony s emicircul ar can al s are revealed E ac h of
. .

the m c o n tains a m e m bra n o u s s emicir cu l ar duc t as will be seen by breakin g Op en ,

o n e of them The three du c ts are si tu a ted in the sam e p lanes and have the
.

sam e names as in elasmobra n chs The rem aini n g s tru c tu res of the in tern al ear
.
,

c onsis tin g of two s m all cham bers

the u tr icu l u s an d the s accu l u s are too dif cu l t
to disse c t .

D or sal e t f th br in E ose th brai re m ov i t h roof f th

3
. a s p c o e a x p e n . n g e o e ,

sku ll an d the side of the sku ll where the sense orga n s were disse c ted in clu di n g ,

the m edial wall of the orbi t No te : the du r a mater in closi n g the brain ; on
.

rem ovin g this the very deli c a te pia mater n ex t to the brain s u bs tan ce The
,
.

ori gin of these m em branes is the sam e as given for the tu r tle and the sp a c es ,

bo un ding them have the sam e n am es Un like the con di tion in the p re ceding .

form s the brain co mp le tely lls the cranial cavi ty

,
.

The brain is shor t an d broad and s tro n gly cu rved in correla tion wi th the ,

bip ed gai t The cu rva tu re resu l ts fro m exu r es of the brain in three region s
. .

The chi ef or pr imary exure o ccu rs in the region of the m idbrain wi th the resu l t ,

t ha t the p os terior p ar t of the brai n is b e n t n early a t ri gh t a n gles to the an terior

 CO M PARA I VE ANATOM
T Y OF THE NE RVOUS S YSTE M 33 1

ar The se o n d or n uchal exu r e takes p la c e in the med u lla bendin

p . t c g the ,

m edu lla a t an a n gle to the sp in al cord The pon tal exu re in the regio n ven tral
.

to the c erebellu m bends the brain in the o pp osi te dire c tion from the o ther two
exu r es wi th the resu l t tha t this region of the bra in is dep ressed
, .

The an terior en d of the brain consis ts of the two very s mall olfactory l ob es .

P os terior to them are the large cereb r al hemispher es sep ara ted by a deep s agittal
ssure These are so enl arged p os teriorly as to co mp le tely c on ceal the dien
.

c e p halon fro m dorsal View wi th the ex cep tion of the deli c a t e pin eal b ody wh
, i ch
is seen in the p os terior end of the sagi ttal ss u re The large optic l obes of the.

m idbrain are ven tral to the p os terior ends of the c erebral he misp heres P os terior .

to the he misp heres is the cu rved cer eb el l u m m arked by tra n sverse grooves
, P os .

terio r and ven tral to this is the medu l l a obl on ga ta its an terior end dep ressed ,

ben ea th the cerebellu m The roof of the m edu lla is co mposed as u su al of a

.

c horioid p lexu s .

Draw the brain from the side .

4. T h e c r a ni al n erve s These are so m ewha t dif cu l t to follow in de t ail .

Work on the side lef t i n ta c t There are twelve p airs of cranial nerves
. .

a ) The olfactor y n er ves : These are two s to u t and elon ga ted n erves p assi n g

from the nasal sac s along the dorsal margin of the in terorbi tal sep tu m to the
olfa ctory lobes .

b) The optic n erves : On t e side where the wall of the orbi t was re m oved no te
h

the s tou t whi te optic tract in fron t of the op ti c lobe F ollow this toward the.

orbi t and nd the Op ti c n erve conne c ted wi th its an terior end .

c) The tr ochl ear n erve: The c ranial ori gin of this nerve is difcu l t to see a t

the p resen t s ta ge of the disse c tion I t arises in the deep g roove be tween the
.

op tic lobe and the cerebellu m an d p asses ven trally be tween the Op ti c lobe an d
,

m ed u lla I t ru ns forward in the oor of the c ranial c avi ty to the orbi t

. To .

nd it in the orbi t exp ose the in ta ct eye as before Cu t throu gh the su p erior
,
.

ob liqu e a t its inser tion on the eyeball and lif ti n g it no te the tro chlear nerve p ass
in g to it and sp readi n g ou t on its ven tral s u rfa ce The op h thal mi c branch of
.

the tri g e m inu s ru ns c lose to the tro chlear nerve .

d) The ocu l omotor n erve: The cranial ori gin of this nerve will be seen la ter .

I t bran ches to the in ferior obliq u e s up erior inferior and in ternal re ctu s mu s cles
, , ,

R emove the eye which is s ti l l in p la c e cu tting the eye mu s cles as near the eye
,

ball as p ossible and p reserving the Ophthahn ic nerve in ta ct L ook for the .

bran ches of the o cu lomo tor a mon g the eye mu s cles in qu es tion The branch .

to the inferior obliqu e in the oor of the orbi t is the m os t c ons p i cu o u s of them .

e) The a b du cen s n erve: This will be fo u nd on ex a m i n i n g the p os terior s u rfa c e

of the ex tern al re c tu s mu s cle This nerve also su pp li es the p yramid an d q u adra te

.

mu s cles .

f) The tr igemin u s n erve: This has three bran ches the ophthal mic the max , ,

il l a r y a n d the ma n dib u l ar
, . The op h thalmi c has al ready been no ted in the dorsal
33 2 LAB ORA O R T Y MANUAL FOR VE R TEB RATE ANATOMY
Q

p ar t o f t he orbi t . F ollow it forward no t in g it,s dis t rib u tion t o th e walls of the

n asal c avi ties . I n the oor of the orbi t near its o u ter m argin lo ca t e the maxill ar y
, ,

nerve Tra ce it forward no tin g its bran ches to the orbi t and u pp er jaw Tra c e
.
, .

the m axillary nerve p os t eriorly carefu lly cu t tin g away tiss u es in its p a th
, .

Cau dad of the orbi t it is joi n ed by the man dib ul ar nerve Tra ce this n o tin g .
,

bran ches to mu s cles and m ain tru nk p assing in to the lower jaw Tra c e the .

c o mm on t ru nk of the m axillary and m a n dib u lar nerve t oward the sk u ll a n d no t e

tha t they are joined in the sk u ll by the o p h thal mi c nerve At the p oin t of u nion
.

is the s emilu n ar gan gl ion lying in t he sku ll F rom the ganglion the trigeminu s
, .

nerve m ay be tra c ed to its ori gi n fro m the side of the m ed u lla below the op ti c

lobe .

g) T h e f a cia l n er v e : T h is arises fro m t he m ed u ll a j u s t ba ck of the roo t of

the tri gemin u s and p asses throu gh the an terior p ar t of the ear c ap s u le where ,

it will be fou n d by s crapin g away the la tt er .

h) The au ditory n erve: This arises c lose to the fa cial an d p asses ou t wi th it

in to the ear cap su le to the variou s p ar ts of whi c h it is dis trib u ted .

i) The gl os s opha ryn geal a n d the vagu s : These n erves arise close to ge ther j u s t
behind the ear cap su le and will be fou nd there by carefu lly di sse c tin g in the
mu s cles The gIOS S Op haryn geal is the s maller of the two and an terior in p osi tion
. .

I t en ters a g a n glion the petr os al gan glion b eyond whi ch it is dis trib u ted to the
, ,

p ala t e p,
har y nx ,
and laryn x The .va g u s nerve is c onsiderably lar g er t han th e

g losso p haryn g eal . I t p asses la t erally p arallel and p os t erior t o t h e g l osso p h a r yn

g eal and en t ers it s an
g g lio n t h e j g
u u
,
l ar an
g g lion whi c h is u ,
n i t ed wi t h th e

p e t rosal a
g gn lion , t h e tw o for m in g a m ass B eyond . t his t he va gu s t u rns p os

terio rl y and p asses down the ne ck su pp lying res p ira tory sys te m hear t an d
, , ,

o ther vis cera P or tions of the symp a the ti c s ys tem are in termingled wi th the
.

nin th and ten th nerves .

j ) T h e s p i n a l a cc es s or y a n d th e h yp g
o l os s al : The for m er p asses o u t wi t h t he

vagu s and is dis trib u ted to c er tain mu s cles The hyp o glossal is fo u n d j u s t .

p os t erior t o t h e va g u s I.t is dis t rib u ted to c er t ain ne c k m u s c les and sends a

bran ch forward to the ton gu e .

5 Ve n tr al as p e ct of the b r ain
.
R em ove the brain fro m the c ra n ial c avi ty
.
,

p reservi n g t h e roo t s of th e c ranial nerves as far as p ossible Those n o t ke .

p t
a t ta ched to the brain will be fou nd in the cranial cavi ty .

No t e form of the Olfa c tory lobes and cerebral he m isp heres fro m the ven tral
asp ec t B e tween the op tic lobes is the dien cep halon In the c en ter of thi s is
. .

the optic chias ma m arked by c ross lines

,
F ro m the o p ti c c hia sm a the s tro n g
.

whi te optic tr acts p ass la terad an d dorsad to the op ti c lobes and dorsal p ar t of
the dien c ep halon .B ehind the c hiasm a is a dep ressed area the infu n dib u l u m ,

fro m whi ch the hypophysis ex ten ds ven trally The la tter is u su ally lef t behind
.

in rem ovi n g the brai n and will be fou n d in a deep pit the s el l a tu rcica in the
, , ,

oor of the c ran ial cavi ty The in fu n dib u lu m bears a c en tral clef t where the
.
33 4 LAB ORA OR T Y MANUAL FOR VE R TEB RATE ANATOM Y

m ay be rem oved in order to exp ose the la tter In followin g one nerve adja cen t .
,

n erves m u s t n o t be des t royed .

I
. The s pin al nerve s the s ympathe ti c s ys te m an d the vagu s
, , .

a) Cervical por tion of th e sympa thetic a n d the vagu s : L o c a t e the va gu s nerve

a t a p oin t n ear the lary nx I t lies alongside the caro tid ar tery
. The nerve .

c rossin g the va gu s near the larynx and givin g o bran ches in t o the s ternohyoid ,

s terno thyroid and rela t ed mu s cles is the descen din g b ranch of the twelf th or
,

hypogl oss al nerve .

R ab bit : The va gu s n erve an d the c ervi c al p ar t of the symp a the ti c t ru n k lie

to ge ther on the dorsal s u rfa ce of the c aro tid ar tery The vagu s is larger and .

more la teral .To the medial S ide of the symp a the ti c t ru n k p os terior to the
larynx may be sep ara ted a deli ca te nerve the cardiac b ranchof the vagu s (depressor
,

n erve of the hear t) Tra c e the sym p a the ti c p os teriorly Ju s t in fron t of the su b
. .

cl avian ar tery it en ters a g an g lion the infer ior cervical gan glion
, F ro m this .

an
g g lion c ords p ass t o ei ther side of th e s u b c lavian ar t ery for mi n g the a n s a s u b ,

cl avia and u n i t e a g ain to ano ther g an glion the rs t thor acic ga n glion
,
S i tu a ted
, ,

p os t erior and dorsal to t he ar t ery

Cat : The sym p a the ti c t ru n k is insep arably bo u nd wi th the vagu s the two ,

forming a large vago sympa thetic tru nk c ou rsing la teral to the caro tid ar tery and
-

bound wi th it by a c ommon shea th Tra ce it cau dad Ju s t in fron t of the r s t

. .

rib bran ches of symp a the ti c origin arise from the tru nk and p ro ceed toward the
esop hagu s Shor tly p os terior to this p oin t the symp a the ti c sep ara t es from the
.

vagu s and generall y en ters a ganglion the middl e cervi cal gan g lion which lies
, ,

in con ta ct wi th the va gu s F ro m this g anglion cords p ass on ei ther side of the

.

su b clavian ar tery form in g the an sa s u bcl avia and p ro ceedin g dire c tly dorsally
, ,

u n i t e to form a lar g e g an g lion the infer ior cervica l g an glion whi ch l ies a gai n s t
, ,

the n e ck mu s cles be tween the heads of the r s t and se c ond ribs .

F ro m the i n ferior c ervi cal gan g lion in b o th an imal s car diac bra n ches p ass to
the hear t . The c o n spi cu ou s nerve lying la teral to the vagu s is the phr en ic n erve
or nerve of the diap hra gm The ri gh t vagu s j u s t af ter p assin g ven tral to the
.

su b clavian ar tery gives o ff the recu rren t or pos terior l aryn geal n erve whi ch ru n s
a n teriorly alon g the side of the tra chea to the larynx The lef t re cu rren t nerve .

arises mu ch far ther p os teriorly fro m the lef t vagu s .

b ) The an terior cervical s pin al n er ves : The sp i n al nerves em erge fro m the
spinal cord in p airs be tween su ccessive ver tebrae p assi n g ou t throu gh the in ter ,

ver tebral foramina Those of the c ervi cal region are called the cervical nerves ;
.

there are ei gh t p airs of them The ven tral rami of the rs t fou r c ervi c al n erves
.

are loosely u ni ted wi th ea ch o ther to form the cervical p lexu s ; the las t fo u r
to ge ther wi th the r s t thora ci c form the b rachial p lexu s AS t he rs t two are .

sm all an d more or less min gled wi th the p os terior cranial n erves they will n o t
be s tu died a t this s ta ge of the disse c tion .

To exp ose the c ervi cal n erves p u ll the mu s cles whi ch are in ser ted on the
,

a n terior en d of the s ternu m ( s tern om as toid s ternohyoid s tern o thyroid) la teral ly

, ,
 COMPARA I VE ANA OM
T T Y OF T HE NE RVOUS S YSTE M 33 5
O

or cu t a c ross them where ne cessary th u s exp osing the mu s cu la tu re of the ver te

,

bral colu mn L ook along the side of this dorsal to the c aro tid ar tery an d no te
.
, ,

the ve n tral ra mi of the sp inal nerves e m ergi n g a t in t ervals At abo u t the level .

of the p os terior end of the larynx lies the third cervi cal n erve in the rabbi t fo u r th ,

in the ca t .As already s ta ted t he nerves th u s exp osed are the ven tral rami o n ly ;
,

the dorsal ra mi are exp osed only by m ore radi cal di sse c tion whi ch will n o t be ,

a ttemp ted here The dorsal rami supp ly the ep axial mu s cu la tu re No te the
. .

bran ches of the exp osed ven tral ra mi to the mu s cles of the side of the
ne ck .

F ro m the ven tral ra mu s of the fo u r th cervi c al nerve ( rabbi t) and f th

c ervi c al nerve ( ca t) arises the phr en ic nerve I t p asses p os teriorly p arallel to
.

the va gu s in the rabbi t c lose to the ver tebral m u s cu la tu re

,
I t re c eives a bran ch.

from the f th (rabbi t) or sixth ( ca t) cervi cal nerve and then c on tinu es p os teriorly
in to the thorax As it p asses the sympa the ti c gang lia it re ceives c on trib u tion s
.

from them In the thorax it lies a t the side of the p eri cardial s a c j u s t ven tral
.
,

to the roo t of the lu n g Tra c e it p os teriorly and no te how it sp reads on the

.

su rfac e of the diap hragm The p hren i c nerves are the mo tor n erves of the dia
.

p h r a g m ; their ori g i n fro m th e c e r vi c al nerves S hows t ha t t h e m u s c les of the

diap hra gm are derived fro m c ervi cal m yo tom es .

c) Th e b rachial pl exu s : The ve n t ral ram i of the fo u r th to ei gh th ( rabbi t) or

f th to ei gh th (ca t) cervi cal nerves to ge ther wi th the ven tral ramu s of the rs t
thora ci c n erve are u n i ted by i n ter c o mmu ni ca ti n g bran ches c alled an s ae to form , ,

the bra c hial p lex u s whi c h innerva t es the mu s c les of the sho u lder breas t fore
, , ,

limb an d diap hra gm The fou r th cervi cal ( rabbi t) or f th ( cat) take p ar t in
,
.
, ,

p le x u s o n ly t hro u g h t heir c on t r ib ti tio n t o the p hreni c nerve .

To exp ose the bra chial p lexu s cu t throu gh the p e c toral mu s cles near the
midve n tral lin e an d sep ara te the p e c toral mu s cles fro m the u nderlying serra tu s
mu s cle . The p lex u s lies in the ax illa along wi th the axillary ar tery an d vein .

Then cu t throu gh the p e c toral mu s cles as near as p ossible to their in ser tion o n
the h u m eru s a n d sep ara te the m fro m the mu s cles of the u pp er a rm I n this .

way the c ou rse of the nerv es in to the fore lim b is exp osed .

The c on n e c tions Of the nerves of the p lex u s are so in tri ca te tha t it is

imp ossible to des cribe them The following p oin ts m ay be no ted however
.
, .

R abbi t : The f th c ervi c al i mm edia tely se n ds a bran ch to the six th c ervi cal an d
th en p ro c eeds la terally in to the n e ck mu s c les The six th c ervi cal is a broad
.

n erve whi c h af ter c o mmu ni c a ti n g wi th the seve n th n erve p asses to the sho u lder
, ,

mu s cles . The seven th is s maller and af ter c on trib u tin g to the eig h th like wise
i n nerva tes the shou ld er mu s cles The eigh th c ervi cal an d rs t thora ci c u n i te to
.

one tr u nk as they emerge from the ver tebral c olu mn F rom this tru nk .

arise the nerves of the li m b Cat : The S ix th c ervi cal has a broad co n n e c tio n
.

wi th the seven th and then p ro ceeds to the sho u lder The seven th an d eigh th .

c er v ic a l s and the rs t thora c i c are ve r y s to u t t ru n ks whi ch are in tri c a t ely co n

nec ted wi th e a ch o ther and fro m w hi c h p o c ee d the n e r v e s o f the fore lim i

r .
 33 6 LAB ORATO R Y MANUAL FOR VE RTEB RATE ANATOM Y
Q

The chief nerves from the bra chial p lexu s are the following :
I The phr en ic nerve This was des cribed above
. . .

2 The s u prascapu l ar n erve

. This is the mos t an terior n erve arising from
.

the six th c ervi cal . The m ain p ar t of thi s nerve p asses be tween the su p raspina tu s
an d s u bs cap u lar mu s cles to s u p p ly the s u p ras p ina tu s and infras i n a tu s I
p n .

the ca t a bra n c h of this nerve p asses over the sho u lder to m ore s u p er cial p ar ts .

3 The
. ven t r a l th orac ic nerves These nerves supp ly the p e c toral mu s cles
.

and wi ll be fo u nd en teri n g the inner su rfa c e of these mu s cles be tween the two
in cision s m ade above They are the mos t ven tral of the n erves of the p lexu s
. .

'

There are two of these n erves on e arisin g from the seven th cervi cal the o ther ,

from the eigh th cervi cal and r s t thora ci c The form er is s mall in the rabbi t . .

4 The s u b s ca pu l ar nerves
. There are three of these dorsally si tu a t ed and
.
,

p assin g in t o t h e i n n er su rfa ce of the shou lder The rs t arises from the S ix th

.

c ervi cal and p asses to the s u bs ca p u lar mu s cle ; the se cond arises fro m the seven th

c ervi ca l and s upp lies c hiey teres m ajor ; the third c o mes fro m the seve n th an d

ei gh th cervica l s and ru ns p os t eriorly alo n g the in t ernal s u rfa c e of the la tissimu s

dorsi mu s cle .

5 The
. a xil l a r y n erve This
. n erve ori gina t e s ch i ey fro m the S eve n th c ervi c al .

I t p asses thro u gh the upp er p ar t of the upp er arm ven t ral to the tri cep s and ,

em erging on the la teral su rfa ce pi the u pp er arm supp lies chiey the del toid
mu s cles .

6 The dors al or l on g thor acic nerve

. This nerve is bes t lo ca ted by examin
.

in g the o u ter su rfa ce of the serra tu s ven tralis mu s cle On tra ci n g it an teriorly
.

the nerve will b e fou nd to p ass in t ernal to the sca l en es and to sp rin g fro m the
seven th c ervi cal nerve close to the ver tebral colu mn .

7 The
. m u s c u l oc u tan eou s n erve ( ca t o nl y ) This arises fro
. m th e ve n t ral
su rfa ce of the six th and seven th cervical s I t p asses to the bi cep s mu s cle fork
.
,

in g as it app roa ches the mu s c le The p os terior bran ch con tin u es along the
.

su rfa ce of the mu s cle an d at the elbow p asses to the la teral su rfa ce of the a r m
an d s u pp lies the skin Of the forear m .

8 The r adial n erve

. . Thi s is the larges t nerve sp rin gin g fro m the p lexu s .

S even th an d ei gh th cervica l s and rs t thora ci c nerves c o n trib u te to its form a tio n .

I t p asses to the u pp er arm and cou rsin g be tween the hu mer u s an d the t ri c ep s
tu rn s dis tally I t s u pp lies m any mu s cles of the fore l im b
. .

9 The
. m edi a n n erve Th i
. s nerve lies p os t erior t o th e radial I t arises in .

the ca t by bran ches fro m the las t three ne r ves of the p le x u s an d in the rabbi t
chiey fro m the r s t thora c i c I t p asses to the u p p er arm and then tu rns
.
,

dis tally ru nnin g alon g wi th the bra chial ar tery .

IO The u l n ar nerve This lies j u s t p os terior an d p arallel to the median

. .

n erve origina tin g chiey from the r s t thora ci c n erve The u ln ar an d me

,
.

dian n erves su pp ly the li mb dis tal to the elbow al tho u gh in the rabbi t the , ,

median n erve in n erva tes the bi c ep s .

 ep axial mu s cles To see them tu rn the anim al dorsal side up and carefu lly
.
,

cu t down thro u gh the m ass of ep axial mu s c les c lose to the ver tebrae .The
dorsal rami will then be seen e m ergi n g from the ver tebral colu mn and p en etra t
in g the ep ax ial m ass a cc o mp ani ed by blood vessels
, .There are twelve or thi r teen
p airs of t hora ci c nerves .

f) The a bdomin al portion s of the vagu s an d the sympa thetic : Tra ce the vagi
in to the p eri to n eal cavi ty re moving the liver if n o t already do n e I n the rab b it
, .

the lef t va gu s c rosses the ve n t ral s u rfa c e of the es o p hagu s obliqu ely to the ri gh t

a n d is dis t rib u ted to the lesser c u rva tu re and ven tral s u rfa c e of the s t o m a ch .

The ri gh t vagu s crosses the dorsal su rfa ce of the esop hagu s obliqu ely to the lef t
a n d is dis t rib u ted to the dorsal s u rfa c e of the s to m a ch I n the cat the ve n tral
.

divisio n of the vagu s p asses to the lesser cu rva tu re the dorsal division to the ,

g rea t er cu rva tu re . I n bo th c ases th e va g i form p le x i on th e s t o m a c h ca lled t h e

ven tr al and dor s al gas tr ic pl exi whi c h also c o n n e c t wi th the nearby symp a the ti c
,

p lex i des
,
c ribed in th e n e x t p ara ra
g p h .

L o ca te a gain the p os terior p ar t of the thora ci c p or tio n of the symp a the ti c

tru nk E xp ose it and no t e the nerve the gr ea ter s pl an c hn ic n erve whi ch arises
.
, ,

from the symp a the ti c tru nk on ea ch S ide an d p asses obliq u ely ven trally toward
the diap hragm . In the ca t this nerve is a c co mp a ni ed by ad di tional s m aller
nerves the l es ser spl anchn ic n erves arisin g fro m the symp a the tic S hor tly p os terior
, ,

to the ori gin of the grea ter sp lan chnic n erve The sp lan c hn i c nerves p ass to
.

ei t her S ide of the c ru ra of the diap hra g m i n to the p eri t o n eal cavi ty .
( The c r u ra
of the diap hragm are the mu s cu lar cords whi ch fas ten the diap hra gm to the
lumbar ver tebrae ) Tu rn the abdo min al vis cera to the ri gh t an d look on the
.

lef t su rfa c e of the s up erior m esen t eri c ar tery n ear its ori gi n from the aor ta .

Two p ro mi n en t s ymp a the ti c gan glia will be fo u nd lyin g o n the s u p erior m ese n teri c
ar tery These are the coel iac and s u per ior mes en ter ic gan glia ; the form er lies in
.

fron t of or o n the lef t su rfa c e of the ar tery ; the la t ter behind or on the ven tral
su rfa ce of the vessel The two ga n glia are bou nd to ge ther by a s tron g conne c
.

tion . The sp lan chni c n erves of bo th sides m ay be tra ced in to the coelia c gan g
lion F rom this ga n glio n a p rom in en t coel iac pl exu s will be seen ex ten di n g
.

t oward the s to m a ch where it c o n n e c ts wi th the g as t ri c p lexi of the vagi

,
. This
g r ea t sy m p a the ti c p le x u s form ed aro u n d and dorsal to t h e s t o m a ch is of t e n

c alled the s ol ar p lexu s . F ro m the c oelia c and su p erior m ese n teri c ga n glia an d
adja cen t p lexi also arise p lexi for the liver sp leen adre n al glan ds gonads an d
, , , ,

the grea t blood vessels .S o m e of these will p robably be se en . The inferior

mes en ter ic gan glio n of the sym p a the tic sys tem lies in the m eso c olon alo n gsid e
the inferior m esen teri c ar t ery .I t is si tu a ted in the infer ior mes en ter ic pl exu s
from which ne tworks ex ten d to adjac en t s tru c tu res .

The m ain symp a the ti c t ru nk of the abdom i n al region shou ld n ow be tra ced
ca u dad fro m the p la c e of ori g in of the sp la n c h n i c n erves .The two tr u n ks
des c en d deep dorsally lying in the groo ve be tween two mu s cle masses At .

se gm en tal i n te rvals they have g anglioni c enl argemen ts from whi ch n erves p ass
 COM P ARATI VE ANATOM Y OF THE NE R VO U S S YS TE M 339

to the g a n glia an d p lexi already no ted At the p os terior en d of . the p eri toneal
c avi ty the symp a the ti c t ru nks grad u ally di mi n ish and disapp ear .

g) The l u mb ar a n d s acr al s pin a l n erves an d the l u mb os acr a l l

p us :
ex There
are S even p airs of l u mbar nerves and fo u r ( rabbi t) or three ( cat) p airs of s acr al
nerves The ven tral rami of the las t fou r lu mbar nerves form a l u mb ar p lexu s
.
,

those of the sa c ral nerves a s acr al p lex u s ; b u t sin c e the two p lexi are u ni ted wi th
ea c h o ther they m ay be c onsidered to ge ther as the l u mb os acr al p le x u s
, .

R em ove all vis c era from the p eri toneal cavi ty in cl u di n g the p os t caval vei n ,

a n d aor ta .I n the dorsal wall is a mu s cu lar m ass ex ten di n g fro m the ver tebrae
to the p elvi c girdle This consis ts of a la teral larger mu s cle the iliops oas and a
.
, ,

s maller medial o n e the psoas min or In the rab b it the p soas minor is a slen der
, .

mu s cle whi ch o ccup ies o n ly the p os terior p a rt of the mid dorsal region ; its s to u t -

S hi n in g t endon p asses to the dorsal side o f the in gu inal li g a m en t In the cat the .

p soas m inor e x t e n ds n early as far an t eriorly as the ilio p soas ; it narrows p o s

terio r l y to a te n don whi ch p asses obliq u ely la terally on the ven tral s u rfa ce of
,

the iliop soas whi ch is thu s exp osed bo th m edially and la terally to the tendon of
the p soas minor The p soas minor c overs a p ar t of the iliop soas in bo th animals
.
,

a n d the g rea ter p ar t of the l u mbar p lex u s is si tu a ted be tween the two m u s c les .

No te the abdo mi n al p ar ts of the sym p a the tic c ords be tween the p os terior p or tio n s
of these mu s cles .

L o c a te the las t thora ci c sp inal nerve I t lies abo u t o n e half in ch p os t erior

.
-

to the las t rib The r s t nerve p os terior to this on the dorsal wall is the ven tral
.

ramu s of the rs t lum bar nerve S hor tly p os terior to thi s is the se con d lumbar
.

n erve . These two nerves p ass to the mu s cles an d skin of the abdomin al wall ;
in the ca t each divides in to two bran ches The third lu m bar nerve emerges dorsal
.

to the ilio p soas mu s cle and divides i n to a larger la teral bran ch to the abdo mi n al
wall an d a more slen der m edial bran ch whi ch p asses obliqu ely c au dad rea chin g
, ,

and foll owing the c ou rse of the iliolu mbar ar tery and vein The fou r th lu mbar .

n erve is the rs t O f the l u m bar p lex u s I t has two m ain bra n ches the l a ter a l
.
,

cu tan eou s nerve a n d the gen itofemor al nerve The former is the s tou t tru nk
.
.

whi ch em erges be tween the iliop soas and p soas minor mu s cles and a cc omp anies
the c o u rse of the iliol u mbar ar t ery and vein p assin g to the thigh ,
The gen i to .

fem oral n erve is a lon g slender n erve whi ch ru ns along the m edial border of the
p soas m i n or m u s c le la t eral t,o th e sym p a t he ti c c ords I n the p os t erior p ar t. of
its c o u rse it a cc o mp an ies the ex ternal ilia c ar tery I t su pp lies the thigh an d
.

abdo minal wall of an d adja cen t to the in gu inal region Af ter lo ca tin g these .

two bran ches of the fo u r th lu mbar tra ce the m toward the ver tebral c olu m n ,

rem oving the p soas minor as far as ne cessary F in d the p oin t of e m ergen ce of .

the fo u r th lu mbar fro m the ver tebral c olu mn and no te the c onn e c tion very ,

s tou t in the ca t be tween the fou r th lu mbar and the f th

,
.

The f th lu mbar co n trib u tes by m eans of its c onne c tion wi th the four th
lum bar to the la teral cu taneou s bran ch n am ed above and also forms a s trong
u nion wi th the six th lu mbar To exp ose these rem ove the res t of the p soas
.
34o LAB O RA OR T Y MANUAL FOR VE RTE
BRA E ANA OMT T Y

m inor . The comm o n tru n k form ed by the u n ion of bra n ches from the f th
an d six t h l u m bar n erves p asses la terally as the lar g e femor a l n erve Tra c e this .

to the thi gh I t c o u rses alo n g the c e n ter of the m edial s u rfa c e of the thigh in
.

c o mp any wi t h the fe m oral ar tery a n d vein I t i n n erva t es adja cen t mu s cles of

.

the thigh a n d then c on tin u es down the S ha n k an d foo t as the s aphen ou s nerve .

The ob tu rato r nerve arises from the c on n e c ti n g band be tween the s ix th and
seven th lum bar n erves and p asses obliqu ely c au dad dorsal to the pu bis throu gh , ,

the ob tu ra tor foram en and in to the g ra cilis and o ther mu s c les .

The seven th lu m bar to ge ther wi th the rs t sa cral u ni t e to form a very large

trii n k the s ciatic nerve
, The six th lu mbar an d se con d sa cral also c on trib u te
.

s mall bran ches to this n erve F ollow the s cia ti c nerve I t tu rn s dorsally p ass
. .
,

in g be tween the ili u m and the ver tebral col u m n Thru s t an i n s tru m en t throu gh
.

the p la ce where it tu rns and disse c t where the i n s tr u m e n t e m erg es o n the dorsal
side of the ani mal On sep ara tin g the mu s cles a t this p la c e the s cia ti c n erve is
.

exp osed E xp ose it as near to the ver tebral c ol u mn as p ossible The gl u teal
. .

nerv es will be seen sep ara ting from the a n terior side of the main tru n k an d p ass
in g in to the gl u te u s mu s cles ( The n erve o n th
. e p os t erior side of th e s c ia ti c

tru nk is the p os terior cu taneo u s des c ribed below ) ,

F ollow t he s cia ti c n erve
.

down the l eg Af ter givi n g o bran ches in to the thigh mu s cles it divides shor tly
.

above the knee in to a la teral bran ch the per on eal n erve whi ch p asses be tween
, ,

the inser tion of the bi cep s fe m oris and the g as tro cn em i u s an d a m ore m edial ,

bran ch the tibial nerv e p assin g be tween the two heads of the gas tro cn emiu s
, , .

The sa c ral nerves are u ni ted by ansae to form the sa cral p lexu s The rs t .

sa cral also as seen above takes p ar t in the form a tion of the s cia tic n erve The
, ,
.

chi ef nerves arisi n g fro m the sa c ral p lex u s are the pu den dal n erve a n d the infer ior

haemor r hoidal The la tter arises in the ca t from the p oin t of u n ion of the three
.

sa cral nerves an d p asses to the bladder an d re c tu m The pu de n dal n erve aris es .

from the large t ru nk form ed by the u ni on of the se co n d and thi rd sa c ral nerves
and m ay also in the ca t re ceive a branch fro m the s cia ti c Thi s t ru n k p asses .

la terally p arallel and p os terior to the s cia ti c F rom it arises the p u den dal nerve
.

whi ch tu rns toward the re c tu m an d u ro ge ni tal organs an d the pos terior cu tan eou s ,

nerve whi ch c on tinu es la terally in to the bicep s femoris mu s c le I t will be fou n d .

by tu rn ing the animal dorsal S ide up and lookin g where the s cia ti c n erve was
exp osed The nerve in qu es tion lies immedi a tely p os terior to the s cia ti c n erve
.

a n d en ters the bi c ep s fem oris The fou r th sa cral nerve in the rabbi t is of m oder
.

a te siz e ; it p asses la terally and then tu rns to the sides of the re c tu m wh i ch it

in nerva tes in co mm o n wi th the p u den dal nerve .

Draw showin g the lu mbosa c ral p lex u s

, .

The foregoing nerves are all the ven tral rami of the lu m bar and sa cral n erves .

To see the s m all dorsal rami of the lu mbar nerves p ro ceed as dire cted for the
dorsal rami of the thora ci c n erves The caud al spin al n erves will n o t be co n
.

s id er ed .
34 2 LAB ORATOR Y MANUAL FOR VE RTE
B T RA E ANATOM Y

M ake a diagram throu gh the cord showing its funicu li and the roo ts of the
spi n al nerves .

M ake a clean cu t a cross the cord and ex am i n e the cu t su rfa ce The S e c tio n .

is divisible in to a c en tral darker ma terial the gray ma tter shap ed like a b u tter
, ,

y in whi ch the nerve cells of the c ord are lo ca ted ; and a mu ch thi cker whi te
,

m a terial the white ma tter s u rrou n ding the gray m a tter and c omp osed of nerve
, ,

bers The whi te m a tter is su bdi visible in to the fu n i cu li n amed above E a ch

. .

fu n i cu l u s c on sis ts of a n u mber of trac ts or b u ndles of bers whose fu n ctions are

kn own b u t these tra cts are n o t visibly di fferen tia ted from ea ch o ther
, .

3. T he p eri pher al d is rib u ion of the po s terior c r an ial n erve s I n this

t t
se ction wil l be des cribed the p erip heral co u r se of the f th seven th and nin t h to , ,

twelf th c rani al n e r ves Fo r the co mp le te dis se c tion of these it is ne cessary to

.
,

have a sp e cimen of which the head is in ta c t b u t mos t of them can be found in

, ,

p ar t a t leas t on th
, e sa m e s p e ci m e n on wh ic h t he p revio u s disse c tio n s were m ade .

a ) The el even th or s pin al access or y n erve: This n erve s u pp lies the s tem o

m as toid Cleido m as toid leva tor s cap u lae ven t ralis and trap ezi u s mu s cles
, , ,
I t is .

a p u re mo tor nerve and is app aren tly derived from the vagu s .

Ra b b it : S ep ara te the s terno m as t oid and Cleido m as toid on the o n e hand

from the b asiocl avicu l ar is an d leva tor s capu lae ve n tralis on the o ther Ru n nin g .

n ear the dorsal border of t he leva tor s c apu lae ven tra l is and p arallel to it is the

sp inal a ccessory nerve B ran ches of the se con d to fo u r th spi n al nerves p ass
.

ven tral to it and u ni te wi th it by bran ches Tra ce it p os teriorly and n o te its

.

bran ches on the in n er s u rfa ce of the t rap eziu s Tra ce it an teriorly and n o te .

bran ches to the leva tor s capu lae ven tralis s terno mas toid and Cleidom as toid
, ,
.

Cat : Cu t thro u gh the cl avo tr ap ezius near its o ri gin and dee c t it ven trally ,

thu s exp osing the leva tor s cap u lae ven tralis On the inner s u rfa c e of the clavo
.

trap e zi u s alo n g the dorsal border of the leva tor s cap u lae ven tralis ru ns the m ai n

p ar t of the s p inal a c c essory n erve Tra c e it.

p os t eriorly n o ti n g bran ches
, in t o
the tr ap ezi u s mu s cles an d the leva t or s ca pu lae ve n tralis Tra c e it an teriorly
. .

I t p asses dorsal to the se c o n d c ervi cal nerve to which it is conne cted by a n et

work and near thi s region gives bran ches to the s terno mas toid and Cleidom as toid
,

mu s cles I t then p asses thro u gh the Cleido m as toid m u s cle

. .

b ) The vagu s the s ympa thetic an d the hypogl os s al n erves : F ollow the vagu s
, ,

and sym p a the ti c an teriorly S tre tch the head forward by cu tting a cross the
.

la teral mu s cles of the n e ck At abou t the level of the p os terior en d of the larynx
.

the va gu s and c aro tid ar tery are crossed ven t rally by the des cen din g b r an chof the
hypogl os s al or twelf th c ran ial nerve This p asses obliqu ely cau dad toward
.

the median line and s u pp lies the s tern ohyoid s terno thyroid a n d thyrohyoid , ,

mu sc les Con tin u e forward

. At ab ou t the p la ce where the c omm on caro tid
.

ar tery divides in to ex tern al and in t ernal caro tids a c on sp i cu ou s nerve is see n

crossin g the ven tral s u rfa ce of the va gu s an d caro tid ar tery and cu rvin g an te

rior ly . This is the main p art of the hypoglossal nerve F ollow it forward I t . .
 COMPARA I VE ANA OM T T Y OF THE NE R VO U S S YSTE M 3 43
.

p asses to the dorsal side of th e m ylohyoid m u s c le whi c h m a y be cu t and inn

, e r ,

va tes the mu s cles of the tongu e .

Abou t halfway be tween the des c ending bran ch and m ai n p ar t of the hyp o
g lossal n erve,
b u t dee p er dorsally and p assin g t o t he dorsal side of t he c aro t id
ar tery is Si tu a ted the s u per ior l aryn geal bra n ch of the vagu s n erve I t ru n s
, .

obliqu ely cau dad to the laryn x whi ch it p ene tra tes p assin g thro u gh the b ers ,

of the thyrohyoid mu s cle .

F ollow the vagu s and symp a the ti c on ce m ore At the p la c e where the .

des cen din g bran ch of the hyp oglossal crosses them the two sep ara t e in the cat .

S hor tly a n terior to thi s the vagu s in bo th anim als p resen ts an elon ga ted swellin g ,

the n odos al gan g lion At abo u t the sam e level b u t more m edial in p osi tion the
.
, ,

symp a the ti c tru nk en ters the s u per ior cervical gan gl ion of the symp a the ti c an ,

elon ga ted p inkish body The two ganglia lie j u s t p os terior to the hyp oglossal
.

as it cu rves forward in to the to n gu e .

The hyp o glossal the a ccessory the va gu s and the s ymp a the ti c are all
, , ,

involved in a p lexu s in whi ch the rs t c ervi cal nerves also take p ar t .

c) The n in th or gl os s opha ryn geal n erve: Thi s lies very c lose to the m ain

p ar t of th e hy o
p g lossal nerve b u t m ore dee p l y dorsal D isse c t dire c tly in t er n al .

to the hyp o glossal where it cu rves an teriorly to the tongu e The gl oss opharyn geal .

is a sm aller nerve lying dorsal to the hyp oglossal alon g the S ides of the p harynx
an terior to the larynx I t is si tu a ted be tween the two horns of the hyoid I t
. .

divides in to two branches : a s m aller pharyn geal bran ch p assing medially i n to

the p harynx and a main l in gu al bran ch whi ch en t ers the t o n gu e The form er .

is a m o tor n erve to mu s cles of the p harynx while the lin gu al b ran ch is a n erve ,

of tas te .

F ollow the nerves thu s far des cribed toward the p oin t where they em erg e
from the S ku ll They are fou nd to c onver ge to a p oin t to the medial side of
.

the tym p an ic b u lla Here the nin th ten th and eleven th nerves em erge from the
.
, ,

brain throu gh the j u gu lar foramen lo ca ted on the m edial side of the b u lla
,
.

The twelf th n erve emerges near the o thers throu gh the hyp oglossal foram en
( co n sis ti n g of several op enin gs in the rabbi t) .

d) The s even th or facial n erve: The m ain p ar t of this nerve is very su p er

cial in p osi tion I t e m erg es a t the p os terior end of the m asse ter mu s cle a t the
.

base of the ear in a sor t of dep ression On carefu lly sear chi n g in this region
, .

it will be fo u nd as a s to u t whi te band in c on ta c t wi th the m ain p ar t of the ex ternal

,

c aro tid ar tery . At this p la c e the fa cial gives off a bran ch to the p os terior p ar t
of the digas t ri c mu s cle and the pos terior au ricu l ar n erve to the p i n na ( The
,
.

large nerve to the pinna whi ch may be no tic ed dorsal to this bran ch of the fa cial
is the great au r icu l ar nerve ori gi n a tin g in the c ervi cal p lexu s ) The fa cial then .

p ro c e e ds forward bran c hi
,
n g over th e ex t ernal s u rfa c e of t he m asse t er m u s c le ,

and p asses to the lip s and region of the eye I t supp lies the variou s p ar ts of .

the p la tys m a mu s cle whi ch it may b e re called is a derm al mu s cle of the he ad

,
 LAB ORATO R Y MANUAL FOR VE RT T
B RA E ANA OM T Y
3 44 E
Q

and ne ck serving to move the ears lip s eyelids whiskers etc The p la tysma
, , , , , .

mu s cle is a vis c er al m u s cle ori gi n ally b elo n gin g to the hyoid ar ch hen c e its ,

in n erva tion by the fa cial nerve .

e) The fth or tr igemin u s n erve: Th i s n erve has three m ai n bra n ches : the
ophtha l mic the ma xil l a r y a n d the man dib u l ar
, , The former is bes t s tu died wi th .

the eye si n c e it p asses in t o the orbi t .

To lo ca te the m an dib u lar bran ch of the tri gemi n u s p ro ceed as follows free ,

in g one half of the m andible Cu t thro u g h the a tta chm en t of the di gas tri c to
.

the m a n dible a n d d ee c t the di gas tri c ba ckward Cu t thro u gh the a tta c hm en ts .

of all of the mu s cles alo n g the m edial s u rfa ce of the m an dible keepin g the knife ,

again s t the bon e Nex t free the la teral or ou ter su rfa c e of the body of the
.
,

m an dible fro m mu s c le a tta chm en ts chiey the m asse ter Cu t thr o u g h the
, .

sym p hysis of the m andible (p lac e of j u n c tion of the two halves of the m an dible
a t their a n terior tip s ) Carefu lly be n d the half of the mandible thu s freed ou t
.

ward so as to exp ose the S ide of the m u s cu lar m ass whi ch form s the oor of the
,

m o u th an d p haryn geal c avi ties The m ai n p ar t of the m andib u lar bran ch of

.

the tri geminu s the inferior al veol ar n erve will now be seen p assin g in to a fora
, ,

m en the m a n dib u lar foram en si tu a ted o n the m e di al s u rfa c e of the m a n dible

, , .

I n the rabbi t the myl ohyoid nerve a n o ther bran ch of the m a n di b u lar will be
, ,

n o ted to the m edial side of the inferior alveolar a n d p ro c eedi n g ve n trally

to m u s c les of the oor of the m o u th c a v i ty The in ferior alveolar nerve ru n s .

in the in terior of the m andible s u pp lyin g the tee th an d then em erg es thro u gh
the m e n tal fora m en on the la teral s u rfa c e of the m andible a t the level of the
dias tem a There the n erve n ow n am ed the men ta l n erve m ay be fou n d an d
.
, ,

followed in to the lower lip .

Tra ce the in ferior alveolar n erve p os teriorly I t co n verg es toward ano ther .

bran ch of the ma n dib u lar n erve the lin gu al n erve which sho u ld then be followed
, ,

forward I t p asses i n to the to n gu e lyi n g close to the hyp o glossal The lin gu al
.
,
.

bran ch of the trigemin u s in n erva tes the mu cou s m embran e of the ton gu e b u t ,

is n o t a nerve of tas te .

F ollow bo th li n gu al an d inferior alveolar n erves c e n trally ag ain I n fro n t .

of the tym p anic b u lla behi n d the p oi n t where the body of the man dible bends
dorsally in to the ra mu s of the m an di ble will be seen the au r icu l otempor al bran ch
of the man dib u lar nerve join i n g the o ther two On tra ci n g it p erip herally it is .

fou nd to pass to the skin of the c ran ial side of the p in n a and in the ca t also sen ds
bran ches alon g the side of the fa ce in c omp an y wi th t he bran ches of the fa cial .

The tymp ani c b u lla m ay n o w be exp osed E m ergi n g fro m the b u lla will be
.

found a slen der nerve whi ch very soon join s the li n gu al bra n ch of the man
d ib u l ar This is the chor da tympan i ( so c alled be c au se it ru n s in the tymp an i c
.

m embrane) a bran ch of the fa cial n erve

,
I ts b ers p ass o u t wi th the li n gu al
.

n erv e and su pp ly t he tas t e b u ds on the an t erior p ar t of the to n gu e ; it also in n er

va tes the su blin gu al an d su b m ax illary salivary glan ds .

34 6 LABORATOR Y MANUAL FOR VE RTF B RATE ANA OM

T Y

in cision thr o u gh the skin above the eye and dee c t the skin downward toward the
eye on whi ch you are worki n g s tre tchin g the S kin away from the head On
, .

the S kin of the in n er s u rfa c e of the u p p er ey e lid n o t e a thin she e t of m u s cle bers ,

p ro c eedi n g in a so m e w ha t c ir cu lar dir ec tion This is

. or b icu l a r is ocu l i a p ar t of ,

the p la tys m a an d has the f u n c tion of c losin g the eyelids

, .

R abb it : S tre tc h t he u pp er eyelid away fro m the head an d c lean away the
c onne c tive tissu e be tween it and the eyeball A thin she e t of m u s cl e will be
.

fou n d ex ten din g from b en ea th the s u p raorbi tal ar ch to the u pp er eyelid Thi s .

is the l evator pal peb r ae s u per ior is whi ch raises the eyelid R ep ea t the fore goin g
, .

dire c tion s on the lower eyelid s tre tchi n g the S kin away from the eyeball 0 n
, .

the in ner s u rfa c e of the lower eyelid no t e the res t of the orbi cu laris o cu li The .

depress or pal peb r ae infer ior is m ay be no t ed ex tendin g fro m the z y go m a ti c ar ch

to the lower eyelid ; it lowers the eyelid R em ove the s u rro u ndin g S kin and
.

eyelids cu ttin g the m away from the eyeball Wi th the bone clipp ers cu t away
,
.

the s u p raorbi tal ar ch and c lean away tiss u e be tw een the dorsal s u rfa ce of the
eyeball and the orbi t A S len der b u t s trong mu s cle will now be seen ex ten din g
.

from abou t the middle of the wall of the orbi t to the dorsal su rfa ce of the eye
ball ; this is the su per ior obliqu e mu s cle I t sep ara tes the thin S hee t of the
.

leva tor p alp ebrae su perioris in to two p ar ts whi ch p ass on ei ther S ide of it Tra ce .

the s u p erior obliqu e to the wall of the orbi t Here there will be fo u n d a ten di
.

n ou s c ord the tr och

, l ea over whi ch the mu s cle p asses
, Nex t rem ove the leva tor
.
,

p pal ebrae s u p erioris an d nd u ndernea t h its p os t erior p or t io n th e t hin at

s u per ior rectu s mu s cle The in ser tion of the su p erior obliqu e on the eyeball is
.

c o n c ealed u nder the m ar g in of the s u p erior re c tu s .

R em ove the half of the ma n dible an d the z y gom a ti c ar ch on the side o n

whi ch you are workin g This fu lly exp oses the ven tral side of the eyeball
. .

Along the ven tral s u rfa ce of the o u ter p ar t of the eyeball ex te n ds the yellowis h
infr aor bital salivary gland M edial to this ex ten din g ben ea th the eyeball is the
.

larger Ha rder ian gland whi ch po u rs its se cre tion o n to the n ic ti ta ting m em bran e
, .

R em ove these glands ; no te the whi te p ar t of the Harderian gland ex ten di n g far
m edially . The inferior obl ique mu s cle is now seen ex ten din g to the eyeball
fro m the an teroven tral region of the orbi t P os terior to it is the inferior rectus
.

mu s cle ori gina tin g fro m the p os teroven tral region of the orbi t No te the bran ch
.

O f the ocu l omotor nerve ru nnin g alon g the an t erior border of the i n ferior re c tu s

and su pp lyin g bo th mu s cles The nerve whi ch ru ns alo n g the p os terior border
.

of the in ferior re c tu s i n n erva t ing the lower eyelid is the zygoma tic bra n ch of
, ,

the m axillary nerve . Im m edia tely behind the in ferior re c tu s is the exter n al or
l a ter a l r ectu s . The nerve p assi n g alo n g the p os terior m argin of the ex t ern al
rec tu s is the l a cr imal bran ch of the m ax illary I t p asses to the l acr imal gla n d
.

and to the skin be tween the eye and base of the p inn a The la crimal gland is .

a small reddish bo dy which will be fou nd by p ress ing the eyeba l l forward an d
searc hin g agai n s t the p os terodorsal wall of the orbi t Two nerves pass the po in t.
 CO M P ARATI VE ANATOM Y OF T HE NE R VOUS S YSTE M 3 47
o
of origin of the ex ternal re c tu s from the orbi t The l ar ger is the ocul omotor the .
,

s maller the a bdu cen s Cu t throu gh the inser tions of the inferior obliqu e and
.

inferior and ex ternal re c ti at the eyeball and dee c t them ven trally Above the .

in ferior re c tu s the in tern al OI medial rectus will be seen inser ted on the eyeball .

Lo ok on the i n n er s u rfa c e of the ex ternal re c tu s and n d the abdu cen s nerve ,

cu rvi n g aro u n d the p os t erior border of the ori gin of this m u s c le and assi
p n g o n

to its s u rfa ce R e tu rn to the dorsal s u rfa c e of the eyeball cu t thro u gh the in ser
.
,

tion of the s u p erior obliqu e a t the eyeball and p ress the eyeball ven trally ,
Two .

n erves will be seen on the m edial wall of the orbi t The lower one is the trochl ear
.

n erve . Tra ce it to the m edial su rfa c e of the sup erior obliqu e The upper nerve .

is the fron tal n erve one of the m ain bran ches of the ophtha l mic branch of the
,

tri g eminu s . I t p asses to the dorsal p ar t of the orbi t and ex i ts thro u gh the
an terior su p raorbi tal foramen to be dis trib u ted to the u pp er eyel id and skin in
fron t of the orbi t I t m ay have been cu t in rem ovin g the su p raorbi tal ar ch
. .

The whi te p ar t of the Harderian gland will be no ted in the an terior p ar t of the
orbi t Cu t thro u gh all of the i n ser tions of the eye mu s cles a t the eyebal l and
.

thro u gh the op ti c nerve re m oving the eyeball , The optic n erve is the s to u t
.

whi te tru nk near the su p erior re c tu s The mu s cles arou nd the Op ti c n erve .
,

ex clu sive of those already iden tied belong to the retr actor b u l b i Find the, .

m ain t ru nk of the o cu lom o tor nerve and t ra c e its bra n c hes to the re tra c tor
b u lbi and s up erior and in ternal re cti The m ain nerve cu rves below the op ti c
.

nerve The n as ocil iary branch of the Op h thalmi c n erve m ay be no t ed p assi n g

.

be tween the su p erior obliqu e and the re tra c tor b u lbi I ts m ain p or tion the .
,

ethmoidal n erve leaves the orbi t by a small foram en in fron t of the sup erior
,

obliqu e mu s cle On tra cing this nerve p os teriorly ne bran ches to the orbi t
.

m ay be see n .

Tra ce the n erves of the orbi t to their ex i ts from the Sku ll The third fou r th .
, ,

and S ix th n erves and the Op h thalm i c and maxillary bran ches of the tri geminu s
p ass t hro u g h th e orbi t al ss u re The m andib. u lar bra n c h of th e tri g e m in u s
p asses t hro u g h th e fora m e n la c er u m .

Cat : R e m ove the eyelids an d the s u rro u ndi n g skin cu t t in g the m away fro m ,

the eyeball . R em ove the half of the m andible an d the z yg o m a ti c ar c h fro m

the side o n whi ch you are workin g P ress the eyeball ven trally away fro m the
.

sup raorbi tal ar ch In the an terodorsal an gle of the orbi t a s trong brou s con
.

n ec tio n will be fo u nd be tween the wall of the orbi t and the eyeb all On inves ti .

g a ti n g thi s it is fo u nd t o c onsis t of t wo bro u s bands whi c h form a p u l ley ; this

is known as the tr ochl ea The ten don of the su per ior obl ique mu s cle p asses over
.

the tro chlea an d is i n ser ted on the eyeball I ts inser tio n is m u ch exp anded a n d
.

ex te n ds cau dad from the tro chlea P os terior to the inser tion of the su p erior
.

obliqu e is a thin at mu s cle the l evator pal peb rae su per ioris or eleva tor of the
, ,

u pp er eyelid . This p asses to the dorsop os terior su rfa ce of the eyeball Cu t .

thro u gh this a t its inser tion P os terior to this m u s c le in the dorso p os terior
.
 3 48 LAB ORATOR Y MANUAL FO R VE R
T T
E B RA E ANATO MY

an gle of the orbi t is the a tten e d l acr imal glan d Cu t o u t the n i c tita ti n g .

m em brane and ex ami n e its in tern al su rfa ce I t is fo u n d to be ro u ghen ed o win g

.
,

to the p res en c e of the Harder ia n g land in its wall .

Tu rn to the ven tral su rfa ce of the eyeball exp osed by the rem oval of the
m an dible the z ygom a tic ar ch and p ar t of the hard p ala t e
, , I den tify a gain the .

sm all reddish infr a or b ital salivary glan d si tu a ted ba ck of the las t too th On
, .

c learin g away c o n n e c tive tiss u e and fa t the infer ior ob l iqu e eye m u s c le will be
,

seen ex ten din g fro m the an terior p ar t of the orbi t to the ve n tral su rfa ce of the
eyeball Ven tral an d a t ri gh t a n gles to the in ferior obliqu e is the inferior r ectu s
. .

The bran ch of the o cu lom o tor whi ch in n erva tes the in ferior obliqu e ru ns alo n g
the p os terior border of the in ferior re c tu s The zygoma tic bran ch of the m axil
.

lary nerve p asses along the p os terior border of the i n ferior re c tu s to the lower
eyelid b u t m ay have bee n des troyed P os t erior to the i n ferior re c tu s is the
, .

exter n al r ectu s ; be tween a n d in ter n al to the m ap p ears o n e of the fo u r p ar ts of the

retr actor b u l b i m u s c le . Alo n g the p os terior border of the e x tern al re c tu s ru n s

the l acr ima l bran ch of the m ax illary n erve s u pp lyi n g the la c ri m al g land and
,

adja c en t ski n On d e ta ching the eyeball from the p os terior wall of the orbi t
.

a n o ther p ar t of the r etractor b u l b i will be seen nex t p os terior to the ex ternal

re ctu s ; dorsal to this is the su perior r ectu s Cu t throu gh bo th obliqu es at their
.

inser tion s an d p ress the eyeball p os teriorly No te the in ter n al rectu s on the
.

an terior su rfa ce of the eye and above it the remain der of the re tra ctor b u lbi .

Cu t thro u gh all of the eye mu s cles and the o p ti c n erve a t their inser tion on
the eyeball an d rem ove the la t ter No te the fo u r p ar ts of the re tra c tor b u lbi
.

arou n d the op ti c n erve D eec t the ex t ern al re c tu s ven trally an d no te the

.

a b ducen s n erve as c e n di n g on its inner s u rfa c e Ru nni n g alo n g the ve n tral s u r

.

fa c e o f the op ti c nerve n o te a slen der n erve the l on g ciliary bran ch of the oph
,

thal mic bra n ch of the t ri geminu s ; it a cco mp anies the o p ti c n erve in to the
eyeball L oo k on the inner su rfa ce of the i nferior rec tu s for the bran ch of the
.

ocu l omotor to t his mu s c le .No te the ciliary ga n gl ion of the sym p a the ti c n ear
thi s bran ch an d obs erve bran ches be t wee n thi s g an glio n a n d the o cu lo m o t or
and lon g ciliary n erves a n d the s hor t ciliary n erves p assi n g from the gan glio n
alon g the op ti c n erve F ind the m ain t ru n k Of the o cu lo mo tor ven tral to the
.

op ti c n erve a t the p la ce of p assage Of bo th throu gh the wall of the o rbi t an d ,

n o te bra n c hes of the o cu lo m o tor to the re tra c t or b u lbi a n d s u p eri o r re c tu s .

B end all eye mu s c les ex c ep t the s u p erior obliqu e ven t rally leav in g the s u p erior ,

ob li qu e a gains t the m edial wall of the orbi t Crossin g the inn er s u rfa c e of the
.

s u p erior obliqu e obliqu ely fo rward are two nerves They are p ar ts of the oph .

tha lmi c bran ch of the t ri gem in u s The lower one is the ethmoidal nerve ; it
.

p asses t hro u g h a fora m en in t o t he nasal c avi t y T h e u pp er one is.th e i nfr a

tr ochl ear n e r ve I t g o es to the an t e rior p art of t he u pp er eye lid

. P os t e rior .

t o and p arall el to the p os t e rior m argin of the s u p erior obliqu e is the fr on ta l

bran ch of the op h thalmi c I t innervat es the upp er eyelid an d in tegum en t
.
 350 LABORATOR Y MANUAL FOR VE R EE B RAT E ANATO M
'
Y

the saw only for the bony p art s Af t er havin g sawed t hrou gh the roof of the
.

sku ll cu t down throu gh the brain wi t h a S in gle slidin g s t roke of a blu nt knife
,

like a table kn ife The brain and S ku ll shou ld t hu s be cu t in tw o one p ar t

.
,

bein g sli gh tly larger than the o ther Wash the cu t su rfa c es gentl y u nder the tap
.
,

and s tu dy the nasal c avi t ies .

The nasal cavi ties are very long in the rabbi t S hor ter in the ca t They are , .

divided in to right and l eft cavi ties or fossae by a p erp endi cu lar p la te the septu m ,

of the nose which is p resen t on the larger se ction of the head The sep tu m
, .

c o n sis ts of c ar tila ge a n teriorly and of t hi n bone p os t eriorly the la t ter bein

g the
,

p per en di c u l a r pl a te of t h e eth m o id bone O n th e. s m aller se c tio n th e la teral and

p os t erior walls of th e nasal fossa are seen to be o c cu p ied by deli c a t e s c rolled a n d

fold ed bones the tu r bin ated b on es or con chae I n the rabb it these are readily
, .

divis ab le in to an an terior c o n cha the infer ior con cha or ma xill otu r bin al m u ch
, ,

folded and lo ca ted on a sep ara te s m all bone of the sku ll ; a middl e con cha or
n a sotu r bin al a lon g sin gle fold dorsal to the p re cedin g and dep e n de n t fro m the
,

n asal bone ; and the s u per ior con cha or ethmotu r bin al p ar t of the e th m oid bo n e , .

In the cat the tu rbinals are c losely c rowded to ge ther b u t by p rying them ap ar t ,

g en tl y there ca n be dis t in g u ished a s m all an t erior m ax il l otu r bi n al o n th e m a x illa ;

abo ve this a single fold the n asotu r bin al dep enden t from the nasal bone ; and a
, ,

g rea t m ass of folds , th e eth m o tu r bi n al llin g th

,
e g rea t er p ar t of th e nasal fossa .

The e thmo tu rbinals are also called the ethmoid l abyrin ths and the sp a ces in closed ,

by the bony folds are called the ethmoid cel ls Den i te p assa ges kn own as the .

mea tu s es of the nose run be tween the co n chae an d c ond uc t the air to the naso
p har y nx . They c o n n e c t wi th th e naso p hary n x below th e e t h m o tu rbi n als .

The pos terior dorsal p ar t of the nasal fossa is closed by the crib riform p la te
of the e thmoid whi ch u ni tes wi th the p erp endi cu lar p la te of the e thmoid bon e
m edially and wi th the p ar ts of the e th moid which bear the l ayr in ths la terally .

The an terior end of the brain ( olfa c tory b u lbs) is readily seen to ab u t again s t
the cribriform p la t e and thro u gh this p la te the bers of the olfa c tory n erve p ass
,

from the olfa c tory m embrane c overing the e thmoid lab yri n ths to the olfa c tory
b u lbs .

c) The s tr uctu r e of the ear : Caref u lly re m ove the brain fro m the two halves

of the sku ll p reservi n g the la tter I n doing this the roof of the sku ll may be
, .

cu t away L oosen the brai n on all sides by p assi n g a blu n t ins tru men t be tween
.

the brain an d the S k u l l The to u gh membra n e the du r a ma ter whi ch c overs

.
, ,

the brain sho u ld b e re tai n ed wi th the brain Carefu lly cu t the c ran ial nerves
.

where they p ass thro u gh the fora mi n a of the Sku ll leavin g their roo ts a tta ched ,

to the brain No te the s mall rou nd reddish body the pitu itary body a tta ched to
.
, ,

the ven tral s u rfa ce of the brain set in to a dep ression in the oor of the sku ll ;
,

keep this body a tta ched to the brain if p ossible P reserve the two halves of the .

brain in a vessel of wa ter or if they are to be kep t for some tim e in weak for m al
, ,

dehyde .
 COMP ARATI VE ANATOM Y OF T HE NE R VO U S S YSTE M 351

Af ter rem oval of the brain ex amin e the c avi ties of the S ku ll on the larger
p ie c e . A n t eriorly behi n d t h e c ribrifor m p la t e is th e s m all an t er ior or o lf a ctory

fossa in whi ch the olfac tory b u lbs are si tu a ted P os terior to this is the large .

middl e or cereb ra l foss a lod gi n g the c erebru m B ehind this is the s maller pos terior
.

or cer eb ell ar foss a for the cerebellu m The c erebral an d cerebellar fossae are .

p ar tl y se p ara t ed by a bony led g e t he ten to r iu,

m whi c h is c on t
, in u ed in life by
the d u ra m a ter I n the oor of the c erebral fossa in the basisp he n oid bo n e is
.

the s ell a tu r cica lod gi n g the p i tu i tary body No te also the optic for amen in fron t
.

of this and behin d this near the ven tral en d of the ten tori u m the foramina for
, ,

the p assa ge of the third to six th cranial nerves In the wall of the c erebellar
.

fossa observe an area of hard whi te bone ; this is the petr ou s p or tion of the
,

te mp oral . I n the cen ter of this is a foram en for the p assa ge of the au di tory
n erve in t o the in ternal ear Above this in the rabbi t is a dep res sion the fl occu l a r
.
,

f ss a whi ch lod ges a p ar t of the c erebellu m called the occu l u s I n removing

o , .

the rabbi t brain the o ccu l u s is lef t behind in the fossa I n fro n t of the ven tral
.

p ar t of th e p e t ro u s bo n e j u s t
,
beh i n d th e t en t ori u m is th e in t ernal ,
o p eni n g of
the facia l can al for the p assa ge of the fa cial nerve B ehind the middle of the
.

p e t ro u s bone is th e j g
u u l a r f or a m e n for t h e p assa g e of th e ni n t h t en th a n d , ,

eleven th nerves B ehind this the twelf th nerve p asses throu gh one or more
.

foramina .

The ear of m amm als consis ts of three p ar ts the extern al the middl e and the , , ,

in tern al ear . The ex ternal ear in clu des the pin n a or au ricl e and the extern al a udi
tory mea tu s lea din g in to the in terior of the b u lla ; t hese have alr eady been no ted .

The middle ear is S i tu a ted in the t ymp ani c b u lla the in ternal ear in the p e tro u s ,

p or tion of t h e t e m p oral bone B o t h are

. c onseq u en t ly in t h e wall of th e c ere
b ellar fossa . Wi th the bone c lipp ers remove this wall in one p ie ce and dis card
the re main der of the S ku ll Clean away the mu s c les fro m its ex tern al s u rfa ce
.
,

exp osi n g the tym p a n i c b u lla .

R abb it : Wi th the bo n e c lipp ers cu t away the ven tral wall of the tymp ani c
b u lla . A lar ge c avi ty the cavity of the middl e ear or tympan ic cavity is revealed .
, ,

In the la teral wall of thi s cavi ty is a rin glike eleva tion of bo n e a cross whi ch is
s tre tched a deli ca te membran e the tympa n ic memb r an e or eardr u m B y p robin g
,
.

in to the ex ternal au di tory m ea tu s de term i n e tha t the m ea tu s term ina tes a t the

eardru m whi ch c loses its in tern al op ening The tymp ani c membrane has a
, .

nearly ver ti c al p osi tion E x tending toward the ty mp an i c membra n e fro m the
.

m edial wall is a shor t cal c areo u s p ro cess whi ch supp or ts the chorda tympan i
bran ch of the fa cial nerve as it crosses fro m the facial to the tympanic m embrane .

An terior to the t ymp a n i c m embrane is a dep ression in which are lodged the three
li ttle ear bon es These bones are so small and so deep ly lodged in the dep ression
.

tha t they canno t be dis tin c tly seen b u t on p i ckin g in the dep ression wi th a
,

for cep s it is u su ally p ossible to ex tra ct one or more of them Co mp are them wi th .

K p age 20 2 Fi gu re 2 1 0
, , .
 3 52 LABORATOR Y MANUAL FOR VE R TEB RATE ANATO M Y

Cat : R em ove the eshy p ar t of the ex ternal au di t ory m ea tu s down to the

t ymp ani c b u lla The m ea tu s will be fou nd to term ina te a t an oval op enin g wi th
.

a sli gh tly eleva ted r im Ac ross the rim is s tre tched the deli c a te tympan ic mem
.

b r an e or ear dr u m The han dl e of the mall eu s or hammer is visible throu gh

.

the eardrum a t t a c hed to its i n tern al s u rfac e Ne x t re m ove wi th the .

,

bone clipp ers the ven tral wall of the b u lla The in terior is the tympan ic .

cavity of the middl e ea r No te tha t it is divided by a bony p la te in to a

.

larger p os terove n t ral chamber and a s maller an terodorsal cham ber The la t ter .

is the on e covered above by the tymp ani c m embrane B reak op en the p la te of .

bon e ex p osin g this cavi ty whi ch is called the tymp anu m p rop er and whi ch con
, ,

tains the ear bo n es No te the m embra n e whi ch l i n es it and the eardru m form ing
.

its an terodorsal wall F ro m the p os terodorsal region of the c avi ty a c al careo u s

.

p ro c ess p j
r o e c t s t o w a r d the eardr u m and c arries th e cho r da t m
y p a n i n er ve a ,

bran ch of the fa cial to the eardru m F rom the in ternal s u rfa ce of the eardru m
, .

the three li t tle ear bo n es are p lai n ly seen ex tendi n g in to a dep ression in the in ter
n a l wall of the ty mp a n u m These may be ex tra c ted and ex amined Comp are
. .

wi th K p age 2 0 2 F igu re 2 1 0
, , .

There now remain s be tween the b u lla and the c erebellar fossa the hard whi te
mass of the p e tro u s bo n e already no ted This c o n tains the in ternal ear Owin g . .

to the co mp lex i ty of the in tern al ear an d its s mall S iz e a disse c tio n of it is imp r ac ,

ti cal b u t its m ain p ar ts can be seen by breakin g away the p e tro u s bone in s m all
,

fragmen ts The tiny spiral ly c oiled cha mber in the bone is the cochl ea; it
.

c o n tains a sp iral tu be the coch l ear duct in which the organ o f so u nd p er c ep tion
, ,

or
( g an of C or t i ) is lo c a t ed I n the t hi c ke r harder p ar t of
. th e p e t ro u s bone are
semi cir cu lar channels the semicircu l ar can al s in closing the s emicircu l ar ducts
, , .

T he in tern al ear is thus han n el s in the p etr ou s bon e co sis tin g

s een to b e in cl o s ed in c ,
n

o f t he co chl ea t he s emicir c u l ar c an al s an d the ves tib u l e o r co n n ec t in g chamb er ; to g ethe

, ,
r,

these co n s tit u te t he b o y l b y i th T he in tern al ear p ro p er o r membran ous l byr in th is

n a r n .
,
a ,

co n t ain ed in t he b o y l abyr in th n I t s p ar ts ar e : t he s a ccul us an d u tr icu l us in cl os d in the

. e

v es tibul e ; t he s emici cu l a d cts a is in g f r o m the u t r ic ul us an d s itu a t ed in s ide o f t he s emi

r r u r

cir c ul ar c an al s a n d t he cochl ear du ct a r is i g f o m the s ac c u l u s an d in c l o s ed in the co c hl ea

,
n r .

T he co chl ear d u ct is a n ew s tru c t u r e chara c ter is tic o f mamm al s a l thou gh it b egin s to a pp ear ,

in bir ds I t is an o u t gr o wth o f t he s cc u l us an d is t he r eal o rg an o f he r in g the s em icir cu l a

. a a ,
r

d Uc ts s ervin g e q u il ib ra t o ry fu c tio n s on ly n .

5 The . r
s t u ctu r e of the brain .

)
a The memb ran es
m en in ge s of th e b r a in : Wi th th
or e t w o halves of the brain
p revio u sly re m oved before y ou s tu dy t h e m e m bra n es of th e brai n The brain is,
.

c overed by a to u gh m e mbrane the du r a ma ter This co n sis ts of the d u ra m a t er

, .

of lower form s fu sed to the in ternal li nin g (p erios teu m) of the sku ll A c onsider .

able sp a ce the s u bdu ral sp a ce is p resen t be tween the du ra m a ter and the o ther
, ,

m em bra n es of the brain The d u ra ma ter dip s down be tween the larger division s
.

of the brai n The su rfa ce of the brai n is c overed by the deli ca te pia mater in
.
,
3 S4 LAB O RA OR T Y MANUAL FOR VE R TJ IBRA TE ANA O M T Y
U

p os t e rior p oi n te d e n d of t he fo u r th ven t ri cle is a t ra c t t e r mi n a t i n g in a c l u b sha p ed -

area the cl ava L a teral to each clava is an o ther elon ga ted area the tu b er cu l u m
,
.
,

cu n ea tu m . These two belon g to the s omatic s en s ory col u mn An terior to th em .

a whi te b un dle p asses toward the c erebell u m disapp eari n g ven tral to an eleva tio n
,

whi c h lies j u s t ve n tral to the hem isp here of the cerebellum The b u n dle is the .

r es tifor m b ody or pos ter ior pedu n cl e of the c ereb ell u m whi c h c o n veys i m u lses
p
fro m the m ed u lla an d sp in al cord to the cerebellu m The eleva tio n over the .

res tiform body is the a rea acus tica or pr ima ry a uditory cen ter The gen eral .

r esem blan ce of these s tru ctu res to those fou n d in the do gS h S hou ld b e eviden t .

Draw the dorsal view of the brain .

c) The ven tral a s pect of the b r a in : No t e the b as il a r ar tery ( c o n ti n u a tio n of

t he two ver tebral ar te ri e s ) r u n n i n g in the m idve n tral li n e an d for m i n g a cir c l e

arou n d so m e s tru c tu res in the ce n ter of the ven tral s u rfa c e This cir cle the .
,

cir cl e of W il lis is joi n e d o n ea c h side by the in tern a l carotid ar t ery

, No te the .

ar teri e s arisin g from the basilar an d cir cle of Willis an d dis trib u ted over the brai n ,

c o u rsi n g in the p ia m a ter The ar teri es sho u ld be re m oved

. .

At the a n terior en d of the ven tral s u rfa ce are the two olfa c tory b u lbs F ro m .

ea ch o n e a denite whi te tra c t the olfactory tr act ex ten ds obliqu ely cau dad an d
, ,

term i n a tes po s t eriorly in a lobe the pyrifor m l ob e whi ch form s the p os te rove n tral
, ,

p ar t of th e c erebral he m is p heres The ss

. u re or s u l cu s w h i c h se p ara t es th e p yri
form lobe fro m the res t of the c erebral he misp here is c alled the r hin al ssu re .

In c losed be tween the two p yriform lobes is the ven tral sid e of the dien cepha l on ,

or thal amen cephal on At the an terior en d of thi s is the opti c chias ma from whi ch
.

the optic n erves p roje c t The region be tween the op ti c c hiasm a an d the olfa c tory
.

tra c ts is c alled the an ter ior perfor a ted s u bs tan ce B ehin d the o p ti c chias m a is a
.

sli gh t rou n ded eleva tio n the tu b er cin ereu m fro m whi ch the pitu ita ry b ody or
, ,

hypophys is d ep en ds by a s talk I n c ase the p i tu i tary body was torn o in

.

rem ovin g the brain a slitlike ap er tu re wi l l be n o ti c ed in the cen ter of the tu ber
,

c in ere u m m arkin g the p la c e of a t ta chm e n t of the p i tu i tary body Imm edia tely .

p os t erior to the a t ta c hm e n t of the p i tu i tary body is an area the ma mmill a r y ,

b ody n o t dis ti n c tly m arked o fro m the tu ber ci n ere um

,
P os terior to thi s is a
.

dep ressed area the pos ter ior perfor a ted s u bs ta nce from whi ch arise the two third
, , ,

or ocu l omotor n erves F rom ben ea th ( dorsal to) the pyriform lobes a thi ck whi te
,
.

b u n dle will be seen p assin g obliqu ely ba ckward on ea ch side of the p os terior
p erfora t ed s u bs t a n ce These
. b u ndles are th e cer eb r a l pedu n cl es belo n g in g t o ,

the midbrain . The fou r th or tr ochl ear nerves arise on the S ide of the brain
be tween the cereb ellu m an d the in ferior colli cu li an d p ass ven trally over the ,

o u ter su rfa ce of the pedu n cles .

The rem ainder of the ven tral su rfa c e of the brai n belon gs to the hin dbrain ,

an d c o n sis ts of the pon s an d the m e d u lla oblo n g a ta The p on s is the heavy ban d
.

O f bers whi c h c rosses the ven tral s u rfa ce of the hi n dbra in i mm edia tely b ehi n d

the p os terior p e rfora ted su bs ta n ce B y followi n g it aro u nd to the S ide s of the

.
 COMP ARA I VE ANA OMT T Y O F T HE NE R VO US S YSTE M 3 55

brain it will be seen to narrow to a whi te cord t he b r achiu m pon tis or middl e ,

p e d u n c l e of th e c erebell u m w,
hi c h p asses i n t o th e s u bs t a n c e of th e c erebell u m .

The p o n s is in fa c t a bridge be tween the two hemisp heres of the cerebellu m

, , .

I t is the ven tral p ar t of the m e te n cep halon of whi ch the c erebell u m is the dorsal
p or tio n . I mm edia t ely p os t erior to the bra c hi u m p on t is a n d p ar t ly c on c eali n g
it is the thi ck roo t of the tr igemin u s nerve On close ex a mina tion this will b e
.

seen to con sis t of a large dorsal por tio n the s en sory roo t (por tio major) whi ch
,

c o n sis t s of the so m a ti c se n sory bers of the t ri gemin u s and a very s m all ve n t ral ,

p or t io n t
,
h e m otor roo t ( por tio min or ) whi c h c on t ains the vis c eral m o tor bers

for the mu s cles of m as ti ca tio n (m asse ter temp oral di gas tri c , ,
P os terior ,

to the p o n s an d of abo u t half its wid th is ano ther b u n dle of tra n sverse bers
the tr apezoid body Close i n s p e ctio n will S how tha t the t rap ez oid body ori gi n a tes
.

from the area a cu s ti ca or au di tory c enter ; it p asses toward the m edia n line b u t
before rea chin g it tu rn s forward and disapp ears d orsal to the p on s The trap e
, .

z oid body is the m ain t ra c t whi c h c arries the a u di tory i mp u lses to the m ore

an terior p or tio n s of the brain Atta ched to the S ide of the area a cu s ti c a is the
.

roo t of the eigh th or au ditory nerve Ju s t ven tral to this and behi n d the roo t
.

of the trigeminu s is the roo t of the facial nerve em ergin g thro u gh the trap ez oid
body I n the m edian ven tral line of the m edu lla is a groove the median ven tr al
.
,

s su r e Alon g ea ch side of this ru ns a n arrow b u n dle of bers ; ea ch emerges

.

dorsal to the p os terior m argin of the p ons and p ro ceeds s trai gh t p os teriorly .

These two tra c ts are the pyr amid s or s oma tic motor tracts ; they c on vey imp u lses
from the c erebral hemisp heres to the volu n tary mu s cles At the p la ce where .

the p yra m ids e m erge fro m above the p on S are the roo ts of the S i x th or a bdu cen s
n erves . The s m all roo t of the nin th or gl os s opharyn geal n erve will be fou n d a t
the p os t erior bo u n dary of the acu s tic area a t the p oin t where the res tiform body
,

p asses dorsal t o i t a n d abo u t on a line wi t h th e roo t of t h e ei g h t h nerve The .

equ ally small roo t of the ten th or vagu s nerve lies im media tely p os terior to a n d
o n a lin e wi th the roo t of the ni n t h n erve P os terior to the vagu s are the n u m er
.

ou s r o o ts o f the eleven th or s pin al a ccess ory n erve arisin g in a line The m ain

.
,

roo t of the a cc essory as cends from the sp in al c ord b u t is p robably m issin g in ,

the sp e ci m en The roo ts of the twelf th or hypogl oss al n erve em erge alon g the
.

la teral border of the p yramid p os terior to the p re cedin g roo ts

,
.

D raw the ven tral view of the brain in clu di n g the roo ts o i the cranial nerves

as far as you have seen them .

d) The median s agittal s ection : Now cu t the larger half of the brain alo n g
the lo n gi tu di n al cerebral ss u re so as to ob tain an ex a c t m edia n sa gi ttal se c tio n .

In m aki n g s u ch a cu t u se a d u ll k n ife an d p ass it thro u gh the brain wi th o n e

slidin g s troke E xamin e the cu t su rfa ce The cerebral hemisp here for m s a
. .

thi ck roof whi c h ar c hes p os teriorly above the dien c ep halo n an d midbrain In .

the c erebral he misp h ere id en tify the se c tion of the cor pu s ca l l os u m This is an ,

obliqu ely p la c ed lon gi tu din al band O fwhi te m a terial B o th an terior a n d p os terior .

356 LAB ORA OR T Y MANUAL FO R VE R E BR A T TE ANATO M Y

en ds are en larged the form er bein g named the gen u the la tter the splen iu m
, , .

F ro m abou t the m iddle of the c orp u s c allosu m a band of bers the forn ix cu rv es , ,

ven trally B e tween the forni x and the a n terior half of the c orpu s callosum
.

s tre tches a thin m em bran e the s eptu m pel l u cidu m co n sis ti n g of two leaves If
, , .

the brai n is se c tio n ed e x a c tly in the m edia n sa gi t tal p lane the se c tion will p ass ,

be tween the two leaves of the sep tu m p ellu cidu m ; b u t of ten the whole sep tu m is
lef t on on e half ; in this case a sli tlike op enin g in to a cavi ty the l ater al ven tricl e

, ,

will app ear on the o ther half be tween the forn ix and the corp u s callosu m The .

fornix p asses downward and soon tu rn s ( as the col u mn of the forn ix ) in to

the i n terior of the brai n where it is los t to view I mm e di a t ely in fron t of the p o in t
.

where it disapp ears is the se ction of a sm all rou n d b u n dle the an ter ior commis su r e , .

F ro m the an t erior c om miss u re a deli ca te m em bra n e the l amin a termin alis , ,

ex ten ds ve n trally to the op ti c chiasm a The forn ix the an terior c ommissu re.
, ,

and the lamin a termi nalis form the a n terior bou n dary O f a deep b u t narrow
c ham ber the thir d ven tr icl e whi c h lies in the middle of the dien c ep halon
, , The .

c avi ty of t he t hird ven tri c le ex tends ven trally i n t o the tu ber c i n ere u m and the

p i tu i t ary body .

The dien cep halon is the m assive region ex ten ding b etw een the for n ix an d
la mi n a termin alis and midbrain I t consis ts of three p ar ts : a dorsal region the
.
,

epitha l a mu s ; a c e n tral and la teral re gion the thal a mu s ; and a ve n tral re gion
, ,

the hypothal amu s The hyp o thalamu s i n clu des the op ti c chiasma the tu ber
.
,

ci n ereu m the m ammillary body and the hyp o p hysis or p i tu i tary body all of
, , ,

whi c h shou ld be ide n tied in the se c tion The epi thalamu s in clu des the s tru c.

tu res in the roof of the dien c ep halon These are : the chorioid pl exu s a thin
.
,

folded vas cu lar m e mbran e be tween the c erebral hemisp here and the die n cep halon ;
the pin eal b ody a s talked body lying in the c horioid p lexu s ; the ha b en u l a a
, ,

s m all m ass j u s t in fron t of the a tta ch m en t of the p in eal body to the dien c ep halon ;
an d the pos ter ior commiss u re a s m all cir cu lar area j u s t p os terior to the habe n u la
,
.

The thal amus co n s ti tu tes the grea ter p ar t of the dien c ep halo n On the cu t .

su rfa ce it p resen ts a lar ge ro u nd m ass the in ter mediate mass or middl e commis
, ,

su r e; this is n o t really a c o mmiss u re b u t m erely the cu t m edian m ass of the

thala m u s . The gr ea ter p ar t of the thalamu s is c o n c ealed by the overha n gin g
c erebral hem isp here On the s m aller pie c e of the brain re m ove the cerebral
.

hemisp here an d th en ex amin e the dorsal an d la teral regions of the thalamu s .

Three low eleva tion s are p resen t The m os t dorsal an d medial one is the pu lvin ar
. .

L a teral to thi s an d whi ter in c olor is the l a ter al gen icu l a te b ody A whi te ban d .
,

the optic tr a ct is p lai nl y see n as c e n di n g from the o p ti c c hias m a a n d termi n a ti n g

,

on the la t eral geni cu la te body P os terior an d ven tral to the la teral gen i cu la te
.

body is a sm all er swellin g the medial gen icu l a te b ody B ehin d the gen i cu la te
,
.

bodies will be re cogn iz ed the c orp ora qu adru gem in a as two low hi llo cks Ven tral .

to them ru n s the s to u t c erebral p ed u n c le the an t erior p ar t of whi ch is c rossed

,

ex ter n ally by the o p ti c tra c t .

358 ,
LAB ORA OR T Y MANUAL FOR VE R E BRA E ANA OM T T T Y
Q

more eviden t in the rabbi t than in the ca t S in c e in , the ca t the bers tu rn dorsall y .

This radia tin g m ass is c alled the cor on a radiata .

f) F u n ction s o f the pa r ts o f the b r a in : As a n aid in t he r

u n de s t an din g o f the an ato my o f
the b ai a few s tatemen ts may b e made con ce n i g the f u c tions of the p ar ts iden tied in
r n, r n n

the p r eced in g sec tions T he o l f ac to ry b ul bs t he o l f a cto ry t a c ts the pyrifo rm l ob e the tu b er

.
,
r , ,

c in er eu m the fo rn i , the hab e ul ae the mammill ary bo dy a n d the hipp oc a mp u s b el o n g to

x, n , ,

the o l f ac to ry a pp a r a tu s o f t he mammal T he o l f actory imp u l s es C o me al o n g t he o l f a c to ry

.

n erves in to the o l f a c to ry b u lb s which a e t he p r ima ry o l f c to ry c en ter s ; they ar e then rel ayed

,
r a

a l o n g the o l f a c t o ry tra c t s to t he py rifo rm l ob e which is t he s ec o n dary o l f a c to ry cen ter ; ,

fro mthe pyrifo rm l obe they p as s to the hipp ocamp u s the tertia ry ol f actory cen ter or con , ,

s c io us c en ter o f s mell T he hipp ocamp u s has e ten s ive co n ec tio ns with o ther p ar ts o f the
. x n

b ain fo r r e e p ur p os es
r x Thes e con n ections o ccur by way o f the mb ria a mass of n erve
.
,

bers The mb ria conn ec ts with the for i an d this in turn with the mammill ary body
. n x .

The hab en ul a is al so co n n ec ted with the hipp o camp u s I n the do g s h p ra c tic a l ly the whol e .

o f the t el en ceph l on is co n c ern ed with s mell whil e her e the o l f a c to ry f u n c tio s o cc u p y b u t

a , n

a p a t o f the t el en c epha l o n the r em ain d er havin g d ev el o p ed

r ,
ew conn ec tion s an d f un c tio n s n .

T he d ien cephal on is the g r ea t c en ter of co rr el a tion s in the mammal ian b r ain I t s r el a tio n .

to the o l f ac to ry s en s e has al r ea dy been n o ted T he l a teral g en icul ate bo dy is t he p rimary .

o p tic c en ter in which as we sa w the O p tic tra c ts t erm in ate in p ar t

, , , F ro m the l a t er al .

gen icul a te b o dy the o p tic imp ul ses p ass to the c ereb r al hem is phe e by way o f t he co ro n a r

radi ta a The p ul vin ar an d the s u p er ior co l l ic u l s o f the midb r a i ar e a ls o co n cer n ed in

. u n

o p tic imp u l s es t he l a t ter b ein g a r e ex c en ter fo r thes e imp ul ses

,
T he prim ary au dito ry .

c en ter is l o c ted in the ar ea a cu s tica ; from here the au dito ry imp u l s es a e ca rrie d in p ar t by
a r ,

the t rap ezo id b o dy to the in ferio r colli cu l u s an d t he medial g en icul a te b o dy which co n s ti

, ,

t u te s eco n dary an d ter tia ry au dito ry cen ters Fro m the medial g en ic ul ate bo dy the a d ito ry . u

imp u l ses ar e c arried in the coro n a radia ta to t he c ereb ral cor tex I n a S imil r w y o ther . a a

s en s a tio n s s u ch as p ain to u ch tem p era tur e p es s u r e cons cio u s n e ss o f mu sc l e a d j o in t

, , ,
r ,
n

movemen ts an d p os ition o f the bo dy in s p a ce ar e car ied by d en ite p a ths ( which are for
,
r

the m o s t p a t in vis ib l e ext ern all y on the brain ) to the thal amu s f ro m which th y ar e t e
r e

l ayed to the c er ebral cor tex ( T he cl ava an d tu ber cu l um cun ea tu m a e co n ce ed with j o in t

. r rn

a n d mu scl e s en se an d s tea d in ess of bo dy moveme t a n d p o s ition ) I t will thu s b e s een that n .

prac tic all y al l s e s atio ns mak e a r el ay in the dien cephal on from which they as cen d by a n ew
n

p ath the coron a radiata to the c er eb al co te T he co on a radiata is thus the gr ea t

, ,
r r x. r

p athway between the d ien ceph l on a d the cer eb al cor tex by mean s of which the se sa a n r n

t ion s ar e p r o j ec ted as it were u p on the co r tex

,
I t is f ur ther well k n o wn o n which ar ea o f
,
.

the cer eb ral co tex each sen s a tion is p ro j ec t ed

r Af ter all o f the s en s a tio n s have thu s b een .

l o c ali ed u p o t he co r tex there s till r emai s a co n s id ab l e p o tio n o f t he c o r tex to which n o

z n ,
n er r

den ite t r ac ts f o m b el o w c a n b e tra ced

r I t is p es u med tha t these ea a e co n cer ed with
. r ar s r n

co - o r d in a t io n , r ea s o n , e mo tio n , e tc .

T he im p u l s es to war d vo l un ta r y
p ar t o f the co r tex
mo vemen ts o rigin a te in a d en ite ,

p as s down war d in the co ro n a radi t in to the c erebr al p edu cl es n d app ea o n the ven tr al
a a n ,
a r

s u rf ace o f t he m e d u ll a as t he p yr am id l t r a c ts o r pyramid s which d es cen d t he who l e l e g th

a ,
n

o f the s p i a l co d a n d m ak e c o
n ec t io s with the m o t or c ell s o f the ven tr al co l u m s o f t he
r nn n n

co r d .T he ce eb al p edun cl es b es id es car ryin g the pyr mid al tra cts al so c a ry l ar g e trac ts

r r ,
a ,
r

f o m the co r te t o the p o ns wher e they p ass in to the cereb ell um

r x ,
.

T he c er eb ell u m is the g rea t c en ter fo r e q uilibra tio n an d mo to r co o r din a tion I ts -

.

fu c tion s ar e in vo l un tary a d un p e ceived by the conscio us min d I t is con n ected with

n n r .

the es t o f the b ra in b y m ean s o f its p ed un cl es : t he p o s terio r p edun cl es o r r es t if o rm bod ies

r ,
 CO MP ARA I VE ANATOM T Y OF T HE NE R VOUS S YSTE M 3 59
O

h
w ic h co nn ec t it wit h the med u ll a an d sp in al co rd ; the br achiu m p on tis o r middl p edun cl e e ,

which j o in s the two sides o f the c ereb ell u m an d al so c o veys t a c ts b etween t he c e eb el l u m n r r

a n d cereb al co r tex ; a d t he an ter io r p edu n c l es ( which were n o t s een in t he dis s ec t io n )

r n

which e ten d be twee the c er eb ell u m a d midb ain a d thal am u s

x n T he ce eb ell m has n r n . r u

ex ten s ive co nn e c tio s with the ar ea acu s tica s in ce the imp u l ses f om the amp u ll ae o f th
n ,
r e

semic ir cu l ar d u c t s which ar e c o n c ern ed with eq u ilib r a tio n

, te min a te in t he a r ea ac s tica ,
r u .

G . S UMMAR Y OF THE NE R VOUS S Y S TE M AND THE S E NS E OR ANS G

I . T he n er vo u s s ys te m an d the n erv ou s p arts o f the sen s e o rgan s are derived from the
ec t o d er m .

2. T he b divided in to c en tral p e ipheral a d symp a thetic n ervou s

n er vou s sys t e m is su ,
r ,
n

sys tems T he r s t in cl des the brain an d s p in al cor d the s eco n d the c an ial an d s p i l
. u ,
r na

n erves an d t he thi d t he symp a the tic c ords g an g l ia an d n erves

,
r , , .

3 T he s p i al cor d c o n s is ts o f a c en tra l g ray r egion a n d a p er iphera l white r eg ion

. n

s u b divid ed in to tr ac ts T he s p i al n erves a ise in p a i s f o m the co d a t s eg men tal in te va l s

. n r r r r r .

4 E a ch s p in al n erve a ris es f r om the co r d by two r oo ts a d o r s al n d a ven tr al

. T he ,
a .

dorsal ro o t is s en so ry a n d b ear s a g an glion co mp os d o f s en s ory c ell s ; the ven tr al roo t is e

mo tor .T he t wo r oo ts u n ite to for m a s p in al n erv e .

5 The s p in al n erve very so on divides in to a do r s al ramus which p as ses to the ep ial

. ax

mus cl es an d adj acen t skin a ven tral r amu s which p as s es to the hyp a ial mu scl es an d
,
x

a dj ac en t s k in an d a co mmu n ica tin g ramu s whic h c o n n ec ts with the symp a the tic s ys tem
,
.
.

6 The ven t al rami of the s p in al n erves are in t ric a tely u n ited by cro s s con n ec tion s i
. r -
n

the region o f the a pp en d g es to fo rm p l exi fr o m which the mo to r n erves to the muscl es of the
a

a pp en dages a rise T he chief p l exi are the brachial p l exus to the a terior app e dages an d
. n n

the l u mb osa cr al p l e us to the p o s terio r app en d ages x

7 T he s ym p a thetic sys tem co n s is ts chiey of a p aired g a n g lio n a ted co r d i the mid

. n

dor sal reg ion of the bo dy c avity These a e con n ec ted with the s p in al an d cran ial n erves
. r

by commun ic atin g bran ches an d with the vis ce a gl an ds bl oo d vessel s etc by m ea s of r , , ,

.
,
n

bran ches n etwo r k s or p lexi an d g a gl ia

, ,
n .

8 T he ve tebra te bra in con s is ts a t rs t o f three v es icl es the fo rebrain o r p r o sen cephal o n

. r , ,

the midbra i o r mesen cephal o n an d the hin db r ain or r ho mben cephal on

n The r s t an d thi d
,
. r

s oo n divide i to t wo ves icl es mak in g ve in all

n ,
.

9 T he ve ves icl es o f the a du l t ver teb rate b ra in ar e ar r an ged in a l on g itu din al s e ies
. r

an d ar e n amed b egin nin g an terio rl y : the tel en c ephal on t he dien cephal on t he mesen cep ha
, , ,

l on the meten c ephal on an d the myel en ceph l o n

, ,
a .

10 The tel en cephal on d iffer en tia tes in to the Ol f ac to ry b u lb s the o l f a c to ry tr ac t

.
,
s,

o l f c to ry l ob es an d cer ebr al hem is pheres

a , T he hemis pher es are l ater al e p an s ion s o f the . x

t el en cephal o T he o l f a c to ry p a t of the tel en cephal on is of p aramou n t imp o tan ce i the

n. r r n

lo wer ver teb ra tes b u t l ter b eco mes s u b o r din ated to the c er ebral hemis phe es
,
a The e r . s

l a t ter in c eas e in s i e in the v er teb r a te s c l e un til in mamm al s they cover mo s t o f t he rem in

r z a a

in g p ar ts o f the b r ain I n m ammal s the t wo hemis pher es ar e conn ec ted by the co r p s

. u

c a l l os u m .

II . p
hyp o thal amus the thal amu s an d the
T he dien ce hal on d ieren tiates in t o the , ,

ep ithal amus T he . ~, hyp o th

l amu s in cl u des the o p tic chias m a o p tic n erves an d n er vous
a , ,

p ar ts of the eye ( these ar e o f ten rega ded however as bel on gi g to the tel en cephalon ) the r , ,
n ,

in f un dibul um an d the m mmil l a ry b o dy

, T he in fun dibu l u m is an evagin a tio n f o m the
a . r

oo r of the dien cephal on ; its mos t ven tr al p ortion u n ites with an ou tgrowt h from the roof
of the mo u th t he two tog ether formin g the p itu itary body
,
T he thal amu in cl u des the . s
3 6 0 LAB ORA OR T Y MANUAL F OR VE R TE BRATE ANATOM Y

g r ea ter cen t al r m ass of the d ien ce phalon an d in mammal s is subdivided in to s ever al p ar ts .

T he p ithal a mu s co n s is ts of the
e p in eal bod y an d o ther evagin ations from the roo f o f th e
i
d ie cephal on an d adj ace t p a ts
n ,
n r .

1 2 T he mes en cephal on or midbr in is co mp os ed do r s all y o f the o p tic l ob es of whic h

. a ,

ther e ar e t wo in mo s t ver teb ra tes four in m amm al s n amed the co r p or a q u ad ru gemin , , a.

Ven t ally the m idb r in co n s is ts o f the teg men tu m an d the cer eb al p ed un cl es

r a r .

I3 T he met e cephal on in cl u des the c er eb el l u m d o rs all y an d the p on s ven tr all y

. n T he .

p on s is d istin ctly devel o p ed o n ly in mammal s .

1 4 T he myel en c ephal on beco mes the medu ll a obl on g a t a

. .

5 T he b ain is ho ll ow its c a vities b ein g k n o wn as v e t ricl es

I . r , T he rs t two ven tr icl es n .

are in the c er eb ral hemis p her es t he thir d ven t icl e in t he dien cephal on , the fo u r th in the r ,

med u ll a I n l o wer v er teb r a tes ven t ricl es ar e al s o p r esen t in the c er eb ell um an d midbr ain
. .

T he ven tr icles are c o ec ted with ea ch o ther nn .

6 T he b rain o f shes an d A mphib ia is p r ovided with ten c an ial

1 . erves that o f r n ,

a mnio tes with twel ve T he l as t two in mn io tes ar e p rob ab l y n o t n ew f or ma tion s

. a .

1 7 T he rs t o r o l f a c to ry
. erve e te d s f o m t he o l f a c to ry ep ithel iu m in t he n os e to
n x n r

the o l f a c to ry bul bs .

18 T he s eco d o r p tic e ve e ten ds fr o m the re tin a to the dien cephal on

. n o n r x I t is n o t .

a tru e n erve b u t a tra c t o f t he b a in r .

9 1 T h e .th ir d o r o c u l o m o to r n erve is a m o t o r n erve t o t he in f e rio r o b l iq u e p rior ,

su e ,

i ferior an d i tern al e c ti an d s o me a c ces s o ry mu s cl es o f the eyeb all

n ,
n r I t o r ig i a tes i t he . n n

midbrain .

2 T he four th o r tro chl ear n erve is a mo tor n erve to t he su p erior o bliqu e mu scl e o f
0 .

the eyeb all I t ar is es f o m the midb ain

. r r .

2r T he fth o r t igemin u s n erve is the chief s o ma tic s e s o ry n erve o f the hea d a n d t he

. r n

n erve o f t he rs t v is cer al or man dib u l ar ar ch I t has in al l ver teb ra t es thr ee b ra ches : . n

the o phthal mic ( d eep o phthal mic o f shes ) b a n ch to the o rb it a n d n as al r egion ; the max il l ary r

to the u pp er j aws a d roo f o f the mou th an d pharyn geal c avities ; the man dib u l ar to the l o we
n r

jaw an d oor of these cavities T he r st two ar e p u r e so ma tic s en s ory n erves ; the ma

. n

dib ul ar al so con t ai s Visceral mo to r b ers to the vis ceral mu s cl es b el o gi g to the l o wer ja w

n n n .

I s hes ther e is an a dditio al br an ch o f the tr ig emin us the su p er cial o phth al mic which
n n , ,

d is pp ear s l a ter
a T he tr ig emi n u s aris es fro m the an terio r e n d of the medu ll a
. .

2 2 T he S i th o ab d cen s n erve is t he mo to r n erve o f the extern al r e ctu s mu s cl e of the

. x r u

e y eb a l l I t t ak es o ig in i t he oor O f t he medu ll a
. r n .

23 T he s ev e th o fa cial n erve is t he n erve o f the s eco n d o hyo id Vis cer al ar ch

. n r I t is r .

l ar g e in s hes b ecau s e it in cl u des bran ches for the l a teral l i e sys tem Thes e dis app ear in n .

l a d v er teb tes l eavin g o ly tha t p a t of the f acial which is

n ra ,
n a med t he hyo m a d ib u l ar
r n n

n erv e in shes This n erve is a s e s ory n erve ( tas te) to the p ha ry geal cavity a d a vis cer al
. n n n

mo to r n erve to the mu s cl es o f t he hyo id ar ch T he f acial s p i gs fr o m t he med ul l a . r n .

4 T he eigh th o r au d ito ry n er ve is a s en sory n erve e te d in g fr o m t he in tern al ea r to

2 . x n

t he ac u s t ic a r e a o f t he medu ll a .

2 5. T he n in t h or g l o sso ph y
ar n g eal n e e is the n e rv rve of the thi d viscer al ar ch In r .

ver tebr a tes

t he l an d it is r edu ced b u t c o n tin u es to s u l pp y sen so y b ers to the oor of the
r

pharyn geal cavity an d vis ceral mo tor b er s to pharyn geal mu s c l es . I t is a tta ched to the
m edu ll a .

26 . T he t en t h or v agu s n e rve is
rve of the rem in in g Vis ceral r ches In shes the ne a a .

it al s o i cl u d es a l ar g e b ran ch for t he l a ter al l in e

n With the l oss Of the l a ter l l in e sys tem n d . a a

the b ra chial mo de o f b r ea thin g the vag u s is mu c h redu c ed b u t c o n ti

n es t o s pp ly t he nu u

c o r es p o d in g reg ion o f the pharynx b ein g a s e so ry

r n e rve t o t he li in g o f t his region a d a
,
n n n n
 AP P E ND I XA
P R ONUN CIATI ON AND DE RI VA TI ON OF TE CHNI CAL W ORD S

S o me co m pou n d wo r d s an d n am es co m bin ed of two or mor e wor d s ar e given un der their

com po n en ts . S om e c o mm on p r x s ar e al o given (L L a tin ; G G
e e s .
, ,
r eek ; F , F r en ch ) .

A ( G pr ex withou t) An s a an n
sah (L a han dl e)

, ,

Ab (L pr ex a way fro m) (G tail )

An u r n ew rah witho u t ;
y
a a on , au r a ,

, ,

Ab d o m e n , d ou gh men , dom i

a b d o min al

ab ab An u s a n us (L r in g )
ti y
,

n al (L , o f un certain o rigin )
(G to li t) f

Ao rta , tab tik a or te,
h d
aor c a or -

i p
, ,

Ab d u c en s a ue

s enz (L , ab , from ; du co, l ead ) Ap o n eu r o s oh n ew ro w

Sis (G po, fr o m ;
k
s a ,
a

(G thorn ) ten don )

Ac an thia s a an the as aca n tha, n eu r on ,

,

(G ( G an ou tgrowth)

wi thou t ; ken tr on ,

Ac en tr ou s a s en trou s a, Ap o h
p y i s s a po y S is

e
, ,

cen ter ) Aqu e d ah

we du ct (L aqua water ; duco
f
c
, , ,

Ac e tab ul u m ass i tab yu l um (L , me o f a kin d

na l ead
of cu p ) Aqu e o u s kwee us ( L a qu a water)

ay

, ,

(L ) a rac k moid ( G ar a chne sp ider)

Ac ip en s er ass i pen set a s tu rg eo n Ar a chn oid

, , ,

krow me (G mos , (G f

Acr o m ion -
a on ,
c hr os , top ; a Ar che n ter on ar ken

ter on ,
archos , c hie ,

s ho u l d er ) rs t ; en teron in tes tin e)

Ac tin o p ter ygiiak

ti n op
'
ter yg

ee eye (G ak
,

b o w)

,
Ar cu alia -
kiu al e cc ah (L

ar ar cu s ,
pter ygion , win g , n )
,
tin os , ray ;
(L a r ter ia , ar t ery )

Ar te rio s u s ar teer cc oh s u s
ko u s (G

a kous tikos , r el ated
,
Acu s tic o -
a ti koh
(G pitch
,
a rytain a ,

Ar yt en o id a r ee tee n oid er ,
t o hea rin g) ,

fu n n el )
p p
Ad (L
Ad d u c tor rex ,

a
to war d ,
d u ck to re (L ,

u on

ad ,
)
to ; duco, l ea d)
As tr a g al u s
Atl t l ss ( G tl to b e r)
as a

a
ass t ra g a l u S

a o,
(G
a
, an a n kl e b o n e)
Ad r en ak -
add r ee

n al (L ad , u p on ; r enes , kid
,

y tr ee m ( L a c o t)

,
Atri m u a u ur
n eys )
,

A dit y
u w d i to e ry ( L
or d it
a

au or iu s , p e rt a in ing
(L ad , fer o b ear )

a err en t
-
A e r en t to ; ,

to hearin g )
, ,

Ali al

cc (L p r e x , al a, win g )
i
l ar ( L ,

Aur icl e , au ricu l a r aw kal , a w rik
-
r ee yu
(G

Auan to l an toe iss all a s , eidos ,
s a , sausa g e ;
a ur icul a , a l ittl e ear )
form) y
( L b irds)

Al ve ol ar , al v e ol u s , al veoli
a1 vee Oh lar ,

- l us , -lie
Ave s a veez ,

(L

Axill a , axill ar y axe il l a h, a xe i l ay r ee a xil l a ,
(L , a li ttl e ca vity ) a l ittl e axis )
,

Amia m (G kin d of sh) a

ai c c ah
-

(G
a

i t m
,
Azyg o s z

ee gos , a, witho u t ; zygon , yo ke)
Am n io n , a mn io ta , a mn o n ee o n , a m

e a n ee
t (G
o h tah a m n ee oa m mb ran e
b

a e of the
mb ryo) ( G b al anos
,

e B al an o g l o s s u s a hl a n o h gl o s s us , ,

Amp l ib ia a m b

cc ah (G a mphi, dou b l e ; bios , a co rn ; gl os s a , t o n gu e)
fea B a s aliab ah s a il
,

(L b as is , the b a se)

li ee a h

G
,

p
)
(L the b as e of

Am p hico el ou s a m fee s ee l o us (

,
a mphi, do u b l e ; B a i b ase
s cc ,
re x, a t

koil os , holl o w) B a s il ar b ass

cc l ar ( L , b as is , the b a se)
Amp hioxu s m a

fee ox

us (G ,
a mphi, b o th; oxys , Bic e p s- b uy

se p s (L bi two ; pu t, hea d)
ca

l b la s
, ,

S ha r p) B l a s to c o e

toe s ea l (G bl a s tos , germ ; hail
Am phip l atyan a m
,

(G am phi,

fee p l a

tee an l os , ho ll o w)

,

b o th; pla tys , a t) m toe d erm (G bl a s tos , germ ;

Bl t d bl s
m p
as o er a ,

Am pu ll a a m p u ll a e a ul l

ah, am p u ll

ee (L d ma s kin )
er

, , ,

as k) Bl t l b las t i
as u a littl e germ)

u l ah (L , a

An am n iaan am

n ee ah (G ,
on , withou t ; a mn ion ,

B r ac hial , b r ac hialis b r achi u m b r ay kee a l , -kee ,

mb ryon ic memb ran e)

e ay l is , -kee u m ( L , b rachiu m, a rm)

An as to mo sis a n ass toh mo w s is (

G an O p en in g ) B r achioc ep halic b r ay kee Oh s e hal ik ( L , br a
-
p
an
,

An con eu s

ko n ee

us (L , a n con , the b en d of chiu m, a r m ; ceph al icu s , er tain in g to the hea d ) p



B r an chia b r an chi ae , b
,
ran c hi al b ra n kee a h,
-kee

An k yl o sis an

kee l o

sis (G ,
a n kyl os , b e n t) cc , -kee al ( ,
b r a nchia , gill s) G
36 2
 AP PE NDIX A 6
3 3

b r an

kee o s s t e g a l

( G br an chia , Cho rio n lio h

( G memb r an e)
ill
Br an c hio s t e g a l , r ee o n ,

gill s ; s tego , t o co v er ) cil iu m, eyel id

(L

Cil iar y s ee a ry )
b r o n c hi al b r o n m i
,
'
B r o n chu s , b r o n c hi, ku s , b ro n
G m (L

Cin er eu s ) n ee r ee u ashy
ka i, b ro n kee

al ( ,
b ron chos , wi n d i e) pp
Cio n a s igh oh n a ( G hion p il l ar)

,

Bu c cal b u ck
al (L b u cca , c hee k or mou th)
, ,

Cir c u l ato ry s ir kiu l a to w r ee ( L cir cul o to fo r m

B u ccin ator b ck u

s i n ay

to r (L , p er ta in in g to a cir cl e)
, ,

b l o win g a t ru m p et )
Cirru s c irr i s ir ru s r ee (L cirr u s tuft l ock o f

b u ll
-
, , , , ,
B ul l a

ah (L a r o u n d s eal o r l oc ket) hair)
b Cl avacl ay va (L b ran ch cl u b )
,

B ur s a u rr

s ah (L a p u rs e )

, , ,

Cl avicl e kl a v i kel ( L cl a r icu l a l it tl e key)
Ca e c u m, c a e ca
s ee

ku m , see
I
ka h ( L , caecu s ,
Cl avo c l ay voh ( L c omb in in g fo r m cl a vis key

, ,

, , ,
b l in d) refer r in g to the cl avicl e)

Cal can eu s , ka l ka y

n ee u s (L , cal x , heel ) i
Cl e d o kl y do u gh ( G c omb in in g fo rm kl eis key

Can in e k y a n in e

(L ca n is , dog) ref errin g to the c l avicl e)
, , ,

pil l u s , hair) Cl eithr u m kl yth ru m ( G kl cithr on b a r ga te)

(L

Ca p il l ar r -
kap ee l ay ry ca

mka
, , , ,

kl y to riss ( G kl eio to Cl os e)

- u l um

Ca p itu l ar , c a p itu l u p it yu l a r y (L , Clitoris

, , , ,

a s mall head) Cl oa ca kl o ay kah ( L s ewer)

Ca r a p ace k a ir p

a a ce ( F, p r ob a b l y fr o m L ,
ca pa, Co cc yx coccyg eal kock s ix S Ij c c a1 ( G wo rd)
,

-

, ,
ho od )
k
Cochl ea ko ck l ee ah ( L sn ail )
'

( G kardia
,
Car dia , car d ia c d ee a h, - d ee ah
Co eliac see l ee ak ( G koil ia s to m ach)
ar , ,

hear t)

, ,

Car in a k a rye

n ah (L , keel ) Co el o m s ee l oam ( G koil os holl ow)

, ,

Car n ivor a , c a r n iv or e
k a r n iv

O h ra h, ka r

n i vo r e
Col o n -
koh l on ( G kol on memb er)

Col u m el l akol yu mell ah (L littl e col u mn )

, ,

(L c ar o , e s h; vor o, d evou r)

G Con chakon ka (L shell )

, ,

Car o tidka p
( )

ro t id ka r o s , s tu or
ka G
, ,

Co n d yl e kon dill (L condyl u s kn u c kl e)

(
'
Car p al e s , c ar p u s r p ay l es , kar p us har , ,

p os , wris t) Con ju n ctiva kon junk tie va (L conju nctu s

d alk
, ,

Cau aw

d al ( L , ca ud a , tail ) join together)
Cava, cava l kave
ah, -
al (L cavu s , holl o w) Co p u l a kop yu l ah ( L cu m together ; apo b in d )
-

k
, , ,
,

( G hor akoeides like a

Cor ac oid kor a koid
(L

Cav ern o s a , cav er n o u s av er n o s ah, -n
ou s ,
, ,

c r o w s b eak)

hol l o w)

cavern a ,

Cen tr al e , c e n tr alia
t ray l ee, -tr ay

l ee ah (L , Cor i u m koh ree u m (L l ea ther)

s en ,

Co r n ea cor n eu m kor n ee ah u m ( L cor n eu s

cen tr al is , c en t ral ) ~

m
, , , ,

t ru m
horn y)
(L )

C en tru sen c en ter

G
,
Corn u corn u a kor n iu kor n iu ah ( L ho r n )
S e ph l ik ( kephal e, head)
k h
'
Ce p h laic a
, , ,

,
Cor o n a , ro w n ah, kor o h n a y ry
(G
c or o n ar y o
a1

Ce p h lai tiza ph
on se i za tio n kephal o,
wrea th)
hea d)
,
(L , cr o wn o r

r d a tas ep h
(G

(G
Co r on o id -
ko r oh n oid kor on e, cr o w )
k

Ce p ha l o c ho a l o w chor day tah ,
,

kephal o, hea d ; chor de, s tr in g ) Cor p or a , c or pu s or p o ra h, ko r p u s ( L, b ody)

(G (L )

Cer a to s er

a to e x her a s , horn ) Co s ta l ko s tal cos ta , rib
p re ,

s
, ,

Ce r e ea r (L , cer a , wax ) Cr an ial , c r an ia t e , c an iu m

r k r ay
n ee a l, -n ee a te ,

-
n ee u G k
m ( r on i on , s ku ll )
mk ib
m um

C er e b e l l u s er e b el l (L a littl e Dr ain ) ,

form (L
,
cr ib ru m, sieve

C er e b r al , c er e b ru m

b ral ,

b ru m (L
Cr ib r ifor r ree , )
b rain )
ser e s er e ,
Cr ico id kry

co id (G kr iks os , rin g )
kris tah (L cre
,

C ervic al s ex vi kal ( L cer vix n eck)

Cr is ta

st )

,
, ,

Chel o n ia kee l o w n ee ah ( G chel on c

tur tl e) Cr o c o dili krok ohdil l
a

ee ah (L cr oco d il e )
G
,

, ,

( G kr os

Chia s m a kai as ma ( c r o ss ma r k) Cr o s s o pter ii ter yg

- yg cross s op ee eye ,

koh frin ge ;
,
s oi , pter on , win g)
(G fun n el )

Choan a , c ho a n ae a n ah, -
a n ee

mko
Cho n d r o cr an iu n

dro cray

n ee
,

u m ( G Cr ur a , c ru r al , c ru s kru

r ah, kru

ral , krus s (L ,

,
cr u s , l eg )
idten
chondr os , ca r til ag e ; kr on ion , k u ll )
(G
s
G mb )

Cten o o id ktein , co

,
Cho n d r o s te i ko n dr o ss tee eye ( chondros ,
m ki i fo r m ( L ,

,
Cun eifo r u n ee cu neu s , wedg e)
ca r til ag e ; os teon ,
ko
b on e)
ki

Cho r d ata , c hor d at e r

da y tah, kor d a te ( L , Cu tan eo u s n tay

n ee u s (L , cu tis , s kin )
cho rda , c o r d o r s trin g ) Cu vie r kin vyay na me o fFren ch an ato mis t)
Cho r d o c e n tr o u s kor
d ou g h s en

trous ( L , chor da , Cycl o
id s igh

kl oid ( G lay/
al as , c ircl e )
mes igh
,

cord ; cen tr u m, cen ter ) Cycl o s to m ata , kl o s to w ma tah,

cycl o s to

Chorio id ( or h
c o r o id
) koh

r ee o l d , ko h

r oid (G ,
-s to me

G ( ,
lay/
al as , c ir c l e ; s toma , m o u t h)

chor ion , mem b ran e) Cys tic s is

G
tik ( , kys tis , b la dd er )
 6 L AB ORA OR T Y MANUAL FOR VE R E T RA E ANA OM T T Y
3 4 B
De fer en s
Q
d e

er enz (L defor o, ca r ry a way ) F ac e t f (L facies face)
a s et

G
, , ,

De l toid d ell

del ta , ou r th l etter
toid ( f of the F acial fay S hal ( L facies fa c e)

G k p a l hab et, t r ian g u l a r in o rm)

,

f
F al cifo r m fal s ee fo r m ( L fa l x s ickl e)
'
, ,

r ee

, ,

(L den s too th)

D en tar y d en ta r ee F al l o p ian fa l oh pee an ( af ter Fal l o p io

d
, ,
, an

en tin ( o r t een ) ( L den s t oo th) a n a t o mis t)

D e n tin e
F as ciaf as h ee ah ( L a b an d o r b an d ag e)
, ,

Der m al ,
d e r mi s d er r mal derr miss ( G der ma ,
'

, , ,

s kin )

F a s cic u l u s , fa s c icu li ia s ik yu l us , -yu l ye ( L ,
d
a
D er m a t o m e ma tome (G der ma , kin ; lit tl e b u n d l e)

err s
f
,

tomos , cu ttin g )
F au ce s ( L thr o a t)

a sees

(G
,

a fr amm dia , b et ween ;

Dia p hr a g m dye F e m o r al , fe m or fm ( L femur .

, i
s e o h r al , o
o h r is s
phr a gn ymi, to in cl ose)
,

thig h)

Dia s te ma dye

a s tee

mah ( G in ter val ) Femu r fee
mu r ( L , thigh)
dy
,

k (G
l l y) F e n e s tr afee n es

Dig as tr ic e g as di t wo ; gas tetr i
r b e
tr ah ( L , win do w)

, , ,

Digitig r a d e dij i ti gr ade ( L , digitu s , n ger , toe ;

F ib l a b u l ar e b
u

yu l ah, yu l ay
-

r ee ( L b u l a
gr ad u s , s te p wal ) k
,

c l a s p b uc kl e)
, ,

di ll
, ,

(G diphyes , twofol d ; ( o r l e) ohpl u me ( L l u m threa d

Dip hycer c al c c s ir kel , Fil o p l u m e

, ,
her kos , tail )
d l
pl u ma fea ther)

,

Dipl o s p on y y l ow S on dy l ee
di p
(G ,
dipl oos , Fimb r ia m b ree a h ( L b o rd er)

F l oc cu l u so c k yu l us ( L a lit tl e piec e o f woo l )

,
d oub l e ; s pond y os , ver teb ra)

diu
F oll icl efoll i kel ( L foll icu l u s a l ittl e b ag )
,
Du o d en u m oh dee n u m (L du oden i, t wel ve)

diu
,

, ,

(L hard)

Du r a rah
F on tan e ll e fo n t a n el l ( F a l ittl e fou n tain )
, '

t kt

,

Gp F or a m en for a m in a f0 r ay men -ra m ee n ah ( L

toh ( o u ts ide )

E c t, ec o e eh re x , the ou ter , , , ,

a n open i n g )
c c
,

den tay tah (L ,

E d en tata e, withou t ; den s ,
F or n ix for n iks (L a n a r ch o r vau l t)

too th)

,

te e en t (L eero F o s s a fos s ah ( L a p it o r c a vity)

to b ea r away fro m)
E er en r

,

ti e er e
, ,

F r e n u l u m fr en yu l u m ( L a littl e b r idl e o r b it)

s hi a ( L
'
E er e n to b ear a way

a n ,
e ero, ,

from) F r o n tal fru n tal (L fr ons b ro w)

, ,

mow F u n d u s fun d us ( L the b o t to m)

E l a s mo b r an ch, m o b r an c hii

e la s -
cc las

,

k
b ran , mow b ran kee (G el a s mos , pl

a te ; F u n g ifo r m fu n ji f o r m ( L , fu n gu s a mu s hr oo m)

-
eye ,

,
br anchia , gil ls )
F u n ic u l u s u n ik yu l u s ( L a s ma l l r ope or co r d)

E n c ephal on en sef a

l on (G en ke phal os , b rain )
,

F u r cu l a fu r kiu l ah ( L a littl e fo rk)

Gp
,

r ex, within , in
,
d ou gh (

en d , en
-
En d, en do

S ide )
e (G
G an gl i
g a gl
on o n ( G a t u mo ) n

ee r
endo,
G

E n d o s tyl e n d o ugh s tyl e within ; ,

id g n o l d ( G g
,
s b ig ht)

s tyl os , c ol u mn ) an o a an o r

G i g a s ee r i an ( f o m a phys ic ian G a
, ,

m (G
'

toe d er m en d o, within ; der ma , as s e r a n r sser )
E n to d er en ,

G as r t ic g tr og as t rik g as troh ( L g
as

as ter ,

, , ,

s to ma ch
E p pi epp PP 6 6 ( G p u po ) )

.e

x a
.e

E p a i lepp a e i al ( G epi po ; L x

,
re x ,

u n
n

a xis ) G as tr o c e n tr ou s g as

troh s en

tr ou s (L , gas ter ,
E pid mi e p ee d err miss ( G
, , ,
s to ma c h; cen tr u m, c en ter )

G g

er s pi,
e u po n ;
k (G
,
a s tr o cn e m iu s

me

der m kin a, S
as tr o n ee us , gas ter ,
cc d id y mis s ( G p s to m a ch; kneme, s han k)
E pid id ymi
G

epp s e pi, u on ;
(G
,
s to mac h;

didymos , tes tis ) a s tr o c o el g as t r oh s eal , gas ter ,
pp G po meros , par t)
koil os , holl o w)
E p im er e mere (
G

e cc
e p i
,
e pi, u n;
as tr u l a
gas
trew l ah ( L , a lit tl e s to mach)
G
"
E p ip hys is , e p ip hys e al cc S is , epp S e al
G
cc

( an o u tg ro wth ) e n icu l a teji n ik yu l a te (L gen u

-
kn ee)
G ji n ye ohhigh o id ( G
, ,

,

e p ip l o icu m kum e n io hyo id gen ei on , c hin ;

E p ip l oic, p l o ik, pl o
epp cc - cc ,

(G pipl oon , o men tu m ) u p s il on , a Y-s hap ed l etter of the G reek alpha b et )

G l
e
epp
,

E p is tr o p he u s

fee (G pi, po e n ita j en

i tal ( L , gign o, to r e p odr u ce )
s trepho, tu rn )
e s tro us ,
e u n;
G en u
jee
n ew ( L , kn ee)

E s o pha g u s , e s o p ha g e al
-
cc so h

a g us , cc

so faj

G er m in a tivu mj er t

mi n ay tiv u m ( L , ger m m o ,
c c al (G ois opha gos , gu ll et to sp t) r ou

G gl
,

Et h m o id eth m o id ( G

eth moe, s ieve) l an s anz (L an a corn )
E u s ta c hia n yo u s tay G
, ,

kee an ( a f ter E u s ta chius l en o id g l ee

n o id (G l
g en e, a s oc ket)
G gl
, ,

a n an a to mis t) l ome rul u s ow mer yu l u s ( L , gl omus ,

b al l
( L p ex
a
Ex r o u t , o u ts id e ) o f yar n )

,

E x cr e to r k k

toh ry (L h
G o o p ar y g l g los s

o h fa r in

jee ai ( o r fa I

(G
y e s r ee , ex , o u t ; cer no , l s s n e a re

sepa r a te ) in jee

, gl os s a ,
t o n g u e ; pha r yn x , ph arynx )
3 6 6 LABORAT OR Y MANUAL FOR VE RT W RATE ANA OM T Y
o

L ym p ha ti r -
lim m fa t ik (L , l ympha ,

cl ear wa ter ) M et
m e ta me t me t

ah (L Gp b e tween
limph
, , or re x , .

Lym p ho c yte s Oh s ites ( L , l ymph

c l ea r a f ter rev ersel y )
G Gm
a, ,

wa ter ; kytos , holl o w l ac e)

, p M e ta c ar p a l m et

a ka r p al (

, eta , a fter ; har
pos w r is t )
G
,

M ajor m
a y jo r (L the g r ea ter )
' M e ta cr o m io n met (

a k ro w

mee on ,

a f te r ; a kr omion p o in t O f the s ho u l d er )
,

M al ar ma y l ar ( L , mal a cheek)
- ,

M e ta m e r e m e ta m er is m met a mere met a me:

,

M al l e ol u s m all e oli m a l ee oh l u s
litt l e ha mmer )
,

,
-
oh l ye (L ,
iz m ( G ,

meta af ter ; mer os pa r t )

,

M e ta pl e u r a lmet a p l u ra l ( G meta af ter

, , ,

M all e u s mahl ee u s ( L ha mm er )
, ,

pleu r a s ide)
,

M al p ig hia n m al p ig hiimahl p ig ee
,

M e tap te ry g iu m met a p ter yg cc u m ( G meta

an , -cc eye
,

( af ter M al pig hi an I talian b iol og is t)

, ,

af ter ; pter on win g )

,

,
M a m il l ar y ( or m a m mil l a r y) ma m ill lay r ee ( L

,
M e tatar s al met a ta r sa l ( G meta af ter ; tar

m a mma b r eas t )
, ,

s os a a t s u r fa c e )
M a mm alma m mal ( L mo mma b r eas t)
. ,

Min imu s min i mu s ( L the l eas t)

, ,

M a m ma l ia ma m a y l ee a h ( L mamma b rea s t)
,

M a m m ar yma m a r ee ( L mo mma b r ea s t)
, ,
M in or my n o r ( L the l es s er )

' ,

M itr al my tral ( F mitr e a p eaked ca p )

M a n d ib l e m a n dib u l ar man d i b l man d ib yu

, ,

, ,

, ,
M o l ar mo w l a r ( L mol a mil l s to n e)

l a r ( L ma ndib ul a ja w) , ,

M on otr e m ata m o n o tr e m e mo n oh trem m a

M a n u b riu mma n iu b ree u m (L ha n dl e)
, ,

G
-
,

tab -t r eem ( mon os o n e ; trema hol e o r

,
, ,
M a n u s ma y n u ss ( L ha n d)
, ,

open in g )

,

,

M ar s u p ial m a r s u p ial ia marr s iu pee ai ma r s iu
,

M iil l e r ia n me l er r i an ( a f ter M ull er a G er man

p ee a y l ee a h ( L mars u piu m po u c h)
,

physio logis t)

, ,

M a s s e t er ma s ee ter ( G ma s eter a che w) M u l ti d u smu l l ti i d u s ( L ma n y c l eft)

-
"

, ,
,

M a s toid mass toid ( G mos tos b r eas t)

M yel o n my e lo n ( G marro w)

G
, ,

M yl ohyo idmy lo w hig h o id (

,

M a te r ma y ter ( L mo ther )
myle mill

,
, ,

M ax ill a m ax il l ar a s ill a h, ma ks m k u ps il on the l e t ter y)

, i l ay y r ee ,

( L maxil l a j a wb o n e)
M yo co mm a m yo co m m ata my o ko mm ah

G
, ,
, ,

M ax imu s maks i mus (L , the l arg es t)

-ko mm
a ta b ( , mys , mu s c l e ; komma , tha t
M -
m ee a t e u s ( L p ass a ge) whic h is cu t o )
my
e a tu s ,

M yo to m e t o me ( G mys mu s cl e ; tome,

M e dia s tin al m e dias tin u m mee d ee a ss tie nal
-
oh , ,
, ,
-
n u m ( L medias tin u s b ein g in the middl e) c u t tin g )

M e diu s
, ,

mee d i u s ( L middl e)

Na r e s n a y
,

(L

Na r is

r is s , n a y r ees n o s t r il )
M e d ul l a m e d u l l ar y m e d u ll l ah medd u l ay ree

, ,

(G
, ,

( L med u l l a ma r r o w pit h) Ne ctu r u s s wim min g ;

n ek to o rus , neletos ,

M e n in x m e n in g e smee n in ks mee n in j ees ( G

, , ,
ou ra , ta il )
G

,

Nep hr o r n e r o s s ( , kidn ey)

, ,
-
memb r an e)

M e n ta l men t al ( L men tu m chin )
Ne p hr o s t o m e n e r o w s to me ( , n ephr os kid
"
G
G
,

M er o b l a s ti cmerr Oh b l as tik (
, ,
n ey ; s toma , mo u th)
mer os pa r t ;

bl a s tos g er m)
, ,
Ne p hr o to m e n e ro w t o me (
-
n ephr os kid n ey ; G , ,

,
tome, c u t tin g )
M e s m e s o mes s m ess o h ( G p r ex the middl e)
,

, ,

Ne u r al n iu r al ( , n eu r on , n er ve)

G
M e s e n c hym e m ess en kime ( G mes os middl e ;
G

en chyma in a uid )
, ,
Ne u r o c o e l i n u

r o w s ea l ( , n eu r on , n er v e ; hoi

,
l os , hol l o w)
M e s e n te r ic m e s e n t er y mes s en ta r e ih -y ( G

k)
, , ,
Nictita tin

mes os mid d l e ; en ter on g u t) g ik n ti tay tin g ( L , n icto , win
,

,
k

M e s o d e r m mess o h d er m ( G mes os middl e ; No d o s al n o d o se )

al (L n odos u s , n o t ty

,
, ,

d er ma S kin ) No to c e n tr ou s (G k

no toe sen t ro us n otos , b ac ;
G
, ,

M e s o g a s te r mes s Oh g as t er ( , me s os middl e ;
hen tron , cen ter )
ga s ter , s t o ma ch)
mess
,

No to cho r d n o

toe ko rd ( G , notos , b ac ; k chor de,

M e s o m er e oh mer e ( G mes os , middl e ; s trin g )

mer os , par t)
,

Nu c h l a n ew

ka l ( L , n ucha , na p e o fthe n ec k)

M es opt er yg iu m mes s op terr yg cc u m ( G

,

mes os , mid dl e ; ptefon , win g)
)

O b l on g a ta oh lo n g ay t a h ( L , obl on gu s , o b l o n g

M e s or chi u m mes s or kee u m ( mes os middl e ;

G , , O b tu r a to r ob tiu r ay

t er (L , ob tu r o, to c l o se ,
or chis , t es t is ) s hu t )
M e s otu b aria mmess
o h tiu b ay

ri u m( G , mes os , O cc
ipital o h s ip

i ta l ( L , occi u t, p b a ck of the
middl e ; L , tu b u s , tu b e) hea d)
M es ovariu mm ess oh vay

ree um ( G mes os , , O cul o m o tor ok
yu loh mo w to r ( L ,

oculu s , eye
mi ddl e ; L , ovu m , egg ) motor mo ver) ,
 APPE NDIX A

O d o n toido d o n

to l d (G odon s , too th) P e tr o s al , p e tr o u s pe tro w

sal , pee

t ro u s (L
Oh l
,

(G petr os u s , ro cky)
k
f y l

ol ene, u ln a ;

O l e c r an o n
G
ee r ay n on ,

k ll) ks
1f k

kran ion , s u P ha l a n g e s , p hal a n x a an gees , fay l an ( ,

ph l b t tl -l in
e)
to h r ee ( L , s mell )
a
O l fa ctor y f
ol acer e, a n x a e
i
0 a to ,

t mo h m ( o r far in jee

a rin jee

O me n u t m ( L f t kin ) en

u a s
P ha r yn g e al , p har yn x al

mo h mo w o hm ( G p ( G pharyn x throa t)
,

) fa r

a ll yn ks
mo s ,
P hr e n icfren ik ( G phr en d ia p hr ag m)
O m o,

o , re x, o s ho u l , ,

der )
G
, ,

O p er cul ar , O p e r cu l u m
o p er

kiu lar , -l u m ( L , P hyl u m fye l u m (

phyl on ra ce tr ib e) ,

, ,

per cu l u m lid ) P ia p ie a h ( L piou s k in d )

o
o d G
, , ,

O phidia phis , s erp en t)

ee a h ( o P in e al p in

cc a l ( L p in ea p in e con e)

P in n a
, , ,
f thal mik ( , ophthal mos
O p hthal m ic Of
'
G ) f
er)
eye
p in a h ( L ea t h
to P is c e s
, ,

(G p piss ees (L s h)

O p is to , p is tho , p is t rex , o pis

O p is 0

then , b ehin d)
P itu itar y p i tiu i t ay r ee ( L pitu ita mu cus )
Ora l oh

r al (L , os , mou th) P l a c e ta p l a c en tal iap l a s en ta h pl as s
,

,

tay

i o
n , , en

(L l ee ah ( L pl acen ta c ake)

O r b icu l ar s r b ik yu l a y r iss or b icu l u s , a

P l a coidpl ak O id ( G pl ax pl a te)
, , ,

lit tl e c ir cl e)

O r b ital , o
b i ta l ,

b i toe (L, or bita i pl
, ,

or b ito
( L pl an ta so l e of the foo t)

r or P l a n tar s an tay r iss
pl
, , ,
o rb it )
t i g rad e ( L

P l a n tig r a d e an pl anta so l e ; , ,
O s tiu m o s s tee u m ( L , mo u tho r en tran ce)

gr ad u s , wa l k)
G p pl
O ti c o h

tik ( otikos , er ta in in g to the ear ) P l a s tr o n

tro n (F b r eas t l ate) p
l itho h to e lith ( G
, as s , a
l ithos , s to n e) P l a ty s m ap l a t iz ma h ( G a t p l a t e)

O to


ou s , ea r ;

G
,

P l e u r a pl eu r alpl ew r ah ra l (
,

m,

O var ia n , o var y o va r ee a n , o h va r ee (L , ovu
, pl eu r a

-
r ib s id e )

, , , ,

)
h
eg g
P l ex u s pl eks u s ( L in ter wea vin g )

pl
,

(L m ductu s , d u c t )

O vid u ct o vi d u c t ovu egg ;
miu l ee ( L , lit tl e fea ther )
m
P l u mu l ae

ew
, ,

P n eu m atic mat (G pn eu ma , b rea th

ih
P al a tal p al a te , p a l a tin e
pal

ta l , p a l

a t e, pal

u , ,

, a a s p ir it )
tiyn ( o ( L pal a tu m
r -
tin ) p a te)
al
P ol yo d o n po l ee o h d o n

( G polys ma n y ; odon s ,

, ,

P all iu m pa hl c c u m ( L c l o a k )
, ,
'

t ee th)
Gp
,

P al p e b r a pa hl p e b r ah ( L eyelid )

P ol yp t er u s
p o l ip

te r ( ol ys , ma n y ; pteron
ti p
, us , ,

P an c r e a s , p a n c r e a c an
'
kree as s , p an

k r ee a t

win g )
G
ik ( , pa s , a l l ; hr ea s , e sh ) P on s po n z ( L b r idg e)
p p l po u
,
.

( L , n i p l e)

P a pill a , p a p il l a e p ill a h, -ee
a P o r ta ta l ( L , porta , g a te)
p a
r

( G prex b eside n ea r) ( L prex b ehin d af ter)

P ar a r ah P os t -

P ar ap hys isp a r a c c s is ( G
, ,
,

P r e p ree ( L prex b efo r e)

para b esid e ; phya , , ,

produce)
P r e p u c e pree pyu se ( F p ro b ab l y

,

fr o m L , pr e,
P ar ie tal pa r ye cc tal ( L paries wall )
,

b efo r e ; an d
pos thion , p en is ) G
P ar o tidpa r o t id ( G para b es ide ; ou s ea r)
, ,

P r im ate p ry ma te ( L , pr imu s , rs t)

, , ,

P a tel l apa tell a h ( L a s mal l pan o r d ish) p

,
P ro ( L or G pr ex b efore)
r oh

P r o c o e l o u s
,

p ek ten ( L c o mb )

P e ct en

,
proh s ee l ou s ( G pr o b efo re ; koil as

pek ti n ee al (L pecten co mb )
, , ,

P ec tin e al , , holl o w)
P r o cto d a e u mpr o k toe dee u m ( G pr ohtos

P e c tor al p e c tor al is- p ek toe ral -r ay l iss ( L pec

, , , , ,

tor alis r efer r in g to t he ches t) a n us ; daio d ivid e)

P r o s ta t epr o s s tay te ( G pr os tates in the fr o n t
, ,

P ell u c id u m p e l iu s ee d u m ( L tran s l u c en t)

.
, ,

P el vic p el vis pell vik -viss ( L pel vis b asin the

r a n k)

, , , , ,

pel vis ) P r o ve n tr icu l u s p r o h ven t r ick yu l us ( L pr o

P e n ispee n is ( L ma l e ext ern al s ex o rg an )

, ,

b efo re ; ven ter b elly)

,
,

P er i- per r cc ( L o r G pr ex a r o u n d ) P s e u d o s u e d o ( G pr ex ps eu dos fa l s e)


, ,

te ryg ee u m ( G pter ygion a lit tl e

P e ric ar d ial , r i ar d i m

kar d ee a1, -u m
P te r yg iu m
p e c u p er ee , ,

( G per i a ro u n d ; hardia , hear t) win g o r n o r p ro j ec t io n fr o m pter on win g ) , ,

, ,

P e rin e u m p er r

i n ee

um ( L per ineon ,
, or ig in P te yg o p t yg o
r er , id t er r i g o g o id ( G pter on

,
-
, ,

u n c er ta in ) W in g )

P e r ip he r al pe r i
er al (G , p er i, ar o u n d ; pher o, P te r yl a eterr y l ee ( pter on fea ther win g ;

G , , ,

b ea r ) hyl e wo od )
P u b ic p u b isp iu b ik b is ( L pu b es hair b y
,

P e rito n e al , p en t o n e u m er r i to e n ee
p
al , -
u m( G ,
,

,
- .
, ,

in f eren c e, ma tu rity)
per i, a ro u n d ; l ein o, s tretch)
p p ll
(G P ul m on ar y
m o w n ay ( L pu l mon l un g)

p m

P e r o n e al , p e r o n e u s ert oh n ee al , -U S , er u ree , ,
,

pin ofa b rooc h, r e er rin gf to the b ul a ) Pul vin ar p u l vin e

ar ( L pul vinu s cu shion )
, ,
3 6 8 LAB ORA OR T Y MANUAL F OR VE RTE B M T E ANA OM T Y
mm i li
O

P ys al Pyc g al ( G.t ree. ru m p)

S e mil u n ar

mi, half; l u na ,

so n ar (L
py
u , se

( G pyg e, ru m p; mo o n )

P yg o s tyl e e g o s tyl e s tyl os ,
S e min al s emm
,

co lu mn )
i n al ( L , s emen , s eed)
P yl o r u s py e l oh ru s

( G pyl or os
, , g a te keeper ) S e min ifer o u s mm se

i n i

er ou s (L , s emen ,
s eed ; fer o to b ear )

,

kwad r a te ( L qu ad ratu s s q uar e) s epp tu m ( L fe ce w ll )

Qu a d r a t e Se p tu m
n a
ee row ah (L s m se
, , , ,

kwad r ee s eps ( L qu attu or fo ur ;

d m)

Qu a r i e
c p s , , S er o s a s s , er u ,
ru
ca p u t, head )
S err atu s se rate u s (L s er r a , s a w )
G
,

Rac hi s ray

kis ( G s pi ) ne
S e s am oid s ess
-
a moid (
p
, s es a mon , a pl an t,

y f
re er r in g to the s ha )
,
e o fthe s eed s
igh

R a dial , r a d ial e , r a d iu s ra d ee a l , ray d ee ay
(L )

S in u s S n us s in u s , c u r ve
d ee (L , r adiu s , ray ) k
- ,
ee , us

R a mu s ,
mu s , ray mee ( L , a b r an ch)
ra mi ray
S kel e to g e n o u s s ell

e to j
'
cc n ou s (G , s kel eton ,

R a p lie
ray fee ( , sea m) G fro m s k ll e o, make d ry)
l ar ( L , s ol , t he su n )
-
S ol ar s oh
(L )

rek tr y sees rectu s , s t r a ig ht
-
R e c tric e s
kt foo t)
,
l ee u s ( L , s ol ea , so l e o fthe
'
S ol e u s S oh
u m, - t u s
R e c tu m , (L )

r e ct us r ectu s , s t r a ig ht
m
re
o b m a t ik ( G s m
,
S oma b ody)

ti c s o a,
i j eez ( L , remu s , ) soh ma toe pl G

R e m ig e s re o ar
,

S o ma

( ma , b od y
R e n al ree
t pl
o eu r e ure so

n al (L r en es , kidn eys ) ,

ep til l pl eur a , s id e)
,

R e p til ia , r e p til e r ee ah, r ep till (L , rep S o mite S oh might (

G s oma , b ody)
til u s , p til f om r p p)
m ati cs per r mat
,

pi
re e, r e o, c ree

R e s p ir a tor y

toh ( or

rah
S p er

ik ( L , s per ma , s p er m )
re s re a ree r es p ee
S p he n o d on s fee

n oh do n (G phen , wedg e ;
) (L b r ea the b a ck) s

to h r ee res pir o, ,

to o th)
,

ree tee (L n et) odon ,

R e te
( G s phen wedge)
,
s fee n o id
S p he n oid
)
,
R e tin a re t i n ah ( L r ete
,
n et
S p ir a cl e spear a h kel ( or s pir e ah kel ) ( L s pi
et
, ,

( L prex b ac k b a ckwa rd)

,
R e tr o r ro h
r a cu l u m air hol e)
, ,

rye n al ( G r hi s n os e)
,
Rhi n a l
S pl an chn i c S pl an k n ik ( G s pl a nchn on o n e of

Rho m b r ho mb oro mb ro mm b o h ( L o r G p r ex
, ,
, ,

, , , the vis c er a )
f
r e err in g to a g eo m et r ic gu re a kin d o fparallel
I

S pl an chn o pl eu r e s pl an k n o h p l u r e ( G s pla nch

,

o g ra m )
,

ro m b oy d ee us (L G n on a vis c u s ; pl eu r a s id e)
R ho mb mb u s , o r
, ,
o id e us rho
S pl e n ial s pl e n ic s p l ee n ee a l s pl en n ih ( G
, ,

r homb os , r ho mb o id in fo r m) , , ,

s pl en s pl een )

R hyn c o c ep hal ia r in ko h see fa y l ee a h ( , r hyn

G ,

S p l e n iu s - s pl ee n ee u s ( G s pl en ion b an dage)

chos , s n o u t ; kepha l e, hea d ) , ,

S qu a m ata S kwa m ay t ah ( L s qu ama s c a l e)

r o w d en t , r o w d en s he a h ( L

R o d e n t , r o d e n tia

, ,
,
S qu a m o s S kwa mo h s a l ( L s qu a ma s c al e)

r od en s , g n a win g ) a l , ,

R o s tr a l , r o s tr u m

tra l , tru m (L r os tr u m, S ta p e s s tay p eez ( L s tirr u p )
-

G
r o ss -

, ,

b eak) S te g o c e p hal a s teg o hs e ahl ah (

s tego co ver ;

, ,

kephal e hea d)
S ac c u l u s s ak

yu l u s ( L , littl e s a c )
,

S ter n e b r a s ter n e b r a e s tir n e b r ah b r ee (

- G
S a c r al , s a cr u m
s ay
c r al , -
kru m (L ,
s ac er , s ac red ) ,
,

s ter n on b r ea s tb o n e ; L ver teb ra j oin t) , ,

, ,

S a g itta l s aj

i tal ( L , s agitta , ar r o w ) St -
s t ir n o h ( G co mb in in g

wo r d s ter n on
ll
e r n o , ,

( L s al iva Spit) b r eas tb o n e)

S al ivar y sa i vay r ee

S a r to r iu ss a r to e r ec u s ( L s ar tor t a il o r)

, ,

s tir mu mm ( G s ter n on b r ea s t b o n e)


S t er n u m
G
, ,
, ,

S au r o p s id a s ah r o p s i d ah ( G s au r os m mo h dee u m ( s toma

-
liza r d ; S to m o d ae u s to w

)
, ,
, ,

ps is , a pp ea r a n ce mo u th; daio d ivide)

o

,

s tray tu m ( L a s pr ead o r co ver )

(G

S c al e n e , s c al e n e s k ay
l ean , kay
l ean s S tr atu m

s s ,
,

) S tr y ay tu m ( L s tr ia furro w)

s hal en os , u n even S tr iatu m
k p
, ,

yu l ah ( L , s ho u l der b l ad e) S tyl o- s ty l o h ( G co mb inin g wo r d s tyl os col u mn )

S cap u l a s a

i ti ig h t
, ,

Sc a c s a

ik ( L , s cia ticu s , o r igin a ll y is chia S ub s u b b ( L p r ex u n d er )

,

dic f om G
us, r , is chion , hip ) Sub l i s u b b cl ay vee a n ( L s u b un der ;

kl y
c a v a n , ,

( G s kl er os , hard) cl a vis key refer rin g to the c l a vic l e)

S cl er a S a r ah

ti kl G
, , ,

i s ull ku furro w o

s s , s ul l s igh (L ,

s kl eros har d )

S cl er o c s e ro t ih ( S ul cu s , s ul c a r

, ,

S cl er o to m e s kl e

r o h t o me ( G s kl er os har d ; g roo ve)
to me c u t tin g )
, ,

S u p in e s u e

pi ne (L su pino, to pu t on the

, ,

S c r o tal s cr o tu m s kr oh tal -tu m ( L , u n cer ta in

b ac k)
o r ig in )
, ,

S u p r a S iu p
( L pr ex ab o ve)

r ah

S u tu r e
,

S c u t e s kiu t ( L s cu tu m s hiel d) s iu c hu r ( L s uo t o se w)

G
, , , ,

e

S l l a sell a h ( L a s ea t o r s a ddl e)

,
S ymphys is simm fee sis ( , union )
-
 App en dix B

P RE P ARA T I ON OF MA TE RIA L S

1 . Kill in g the S p e cim e n s Nectu r us is

best k ill ed b y pl in g in ho t wa ter ; t tl es b y ac ur

in j c tio o f ther o r hl o f m in to the cl o aca or better the tr ach a ; bi d a d m mmal s by

e n e c r o or e r s n a

i cl sin g them in a tightly cl o sed ves sel with a w d o f co tto n s oak ed in e th o chl o o f m
n o I a er r r or . n

han dl i g tu r tl es pu l l the head forwa d by in ser tin g a s tou t hoo k b ehin d the jaw an d p ry o pen
n r
,

the mo u th .

P p 2. ti f k l t s
re
ar a S k el to s ar e bes t p p ar ed fr o m f e h ma t i l o in the
on o s e e on e n re r s er a s r,

ca es o f m a i e f o rms thos e tha t hav e been p r s erved in b ri e

r n , cul t to p p a e s k l
I t is d if e n . re r e e

t n s fro m s p cimen s tha t h ve been p es er ved in fo malin

o e To p ep r a s k el et on emo v th
a r r . r a e r e e

S kin a ll o f the vi c e an d as m u ch o f the mu s cl s as p oss ibl e an d s o k the s e cim

s ra , in wa ter e a
, p en .

The r em ain in g esh will decay an d may b e r emov d with a s tif f br us h o r fo cep s This proce s e r . s

o f mace a t io n i c o l d wa ter ta k es s ome tim e

r n The p o c s s may b e mu ch sho t d n d to u gh . r e r en e ,
a

s p ecim s a m en s il y p ep ared by immers i g the s pe cimen f r a few h u s i

re o r e ea r h t o b o ili g n o o r n o r n

wa ter to whi ch g l d d u t or the f ll o wi g s o ap s ol tio n ha s b e n dded

o s K i g l ey gives the
o n u e a . n s

foll o win g formul a fo th s oap s ol u tion : r e

7 5 gm s . of hard s oap
12 gm s . o f p o tass iu m n i tra te ( sal tp e ter)

I 50 c c . . o f s tr o n g ammon ia

c c . . o f dis till ed o r s o f t wa ter .

M ix t horoughly . In us in g , ta k e on e p ar t o f the so ap s ol u tion to thr ee o r fou r p ar t s o f wa ter .

T he l en g th of time re quir ed before the esh wil l separa te eas ily fro m the b o n es var ies with
d iffer en t animal s yo un g sp ecimens than fo l a ge a d ol d o n es Da vi
an d is sho r ter for s mal l or r r n .

son gives the tim e for the ca t as two to fou r ho u s in b o il in g w ter or be t te th ee to s i h rs in r a r r x ou

wa ter kep t a t 75 to 9 0 C He al so s ta tes tha t if the bo n s ar hea t d fo r n ly on e to two ho urs

. e e e o

a t a tem p er a tur e n o t abo ve 8 5 C the l ig ame ts will b e p s e ved a d wh n the S k el eton is

.
, n re r ,
n e

d ri d the ligam en ts wil l har den an d hol d the b o es to g ether

e S k el ton s con tain i g a con s id r n . e n e

abl e a m oun t o f ca til age S hou l d n o t o f cou rse b e bo il d b u t t ea ted o nl y with m d ra tely ho t
r , ,
e r o e

For shes a f w min u tes tr ea tm n t with ho t w ter is g en e all y s u i ci n t Car til ag i

wa t r e . e e a r e .

n o us S k el ton s sh u ld b e pres erved in weak formalin or in 70 p er cen t l cohol

o e a .

3 In j ti
. n f th ir
ec l at ry y t m
o o T O en de c the b l d ves s l s co n sp icu o us a d m
cu o s s e . r er oo e n o re

eas il y f oll o wed it is ad visabl e a n d in fac t p r ac tic l ly eces sary tha t the te ies a t l ea s t b a n ar r e

i j cted with a col or ed s ol u ti n

n e I j ec ti n yr in g s for thi p u rp o s m y b e O b tain d f o m
o . n o s e s e a e r

deal ers in l ab o a to ry s u pp lies or an o di ary ru bber a to mi er b u lb m ay b e u s d

r ,
A g l ass r n z e .

c nn ul a i i ser ted in the v es s l t

a s n b e i j ec ted A can n ul a is S imp l y a p i ce o f gl a
e o tu b i g n . e ss n

drawn u t in the ame a t o e e d to a i suit b l e fo r the bl oo d v e

o n l in to which it is to b e
n s ze a s se

i er t d
ns I t is al s o d s i ab l tha t the e d t b e in s e ted in the ves s el b e s l ightly en la rged as it
e . e r e n o r

wil l the h l d m o e s cu ely in the b l ood ves s el

n o r e L oosen the ves s l to b e in j ec ted fro m the s ur
r . e

rou n di g ti s u s p as s a co r d un d e it an d ti the co d in a l oo s e s i gl
n s e ,
kn o t b o ve the ves sel
r , e r n e a .

With a n e s cissors make a V sh p ed c u t i to the bl ood ves sel h vi g th cu t e ten d n o t mo e

- a n ,
a n e x r

than hal fw y thr u gh the v s s l a d im media tel y i er t t he can ul a i t the v s sel

a o e e ,
n Tig ht n ns n n o e . e

the k n o t o u d the ca n u l a an d bl o od vess el b y p u llin g o n the two e ds o f th co r d b u t m ak e

ar n n n e ,

o nl y a s in g l e k n o t T he ca n n ul a is con n e ted w ith th i j e c ti

. v i g or ru b ber bu lb b y a c e n on s r n e

$
70
 APPE NDIX B 37 1

piece of rubber tubin g The whol e sys tem shou ld b e ll ed with the in j ec tion uid in adva e
. nc ,

in o d r to av id for cing air in to the bl d ves sel s

r e o Af ter every thin g is eady in j ec t the s l u ti n
oo . r o o

in to the b l o d ves sel by a s teady b u t n o t too fo ceful p es sur e on the syr i g

o bul b The su c r r n e or .

cess o f the in j ec tio n shou l d b e d e ter mi ed by exa min in g the s mall ves sel s o f th s k in or in then e

in tes ti al wall s
n When these are deeply col ored the in j ection is compl ete an d the c n n ul a is
. a

th n withd awn the cor d bein g immediately tigh ten ed aroun d the bl ood ves s el an d tied with a
e r ,

d ou bl e k n o t to p even t l eak ag e I n case o n ly the ar terial sys tem is in j ec t d s is u su ll y the

r . e ,
a a

c s e the bl oo d shou l d n o t b e l et ou t o f the an imal ; the in j ection wil l f o c e the bl o o d

a , r u d r a o n

in to the vein s an d dis ten d them If the sys temic vein s are to b e in j ec ted the bl oo d sho ul d b e
.
,

d ain ed from the animal by op en in g the vein a t the p l ace wher e it is to b e in j ec ted
r ,
.

T he foll wi g fo m ul a for the in j ec tion uid is given by Davis n :

o n r o

1 00 c c . . o f wa ter

20 c c . . o f gl ycerin
20 c c . . o f co n cen t r a ted formal in
8 5 gms . o f co rn s tarc h
s u i cien t col o r in g ma t ter to g ive a p c ol or
d ee .

S tir thor ou ghl y an d s tr ain throu g h n e heesecl th

c o . Al l l u mp s mu s t b e r em o ved . As
the s ta r ch s ettl es to the bo ttom o n s tan din g ,
the mix ture mu s t b e s tirr ed bf e o r e u s in g Co l o r
in g ma tter u s ed in the in j ec tion mas s is : for r ed co l or , ver m il io n ( mer cu ric s ul phide) ; fo r
yell ow r chrome yell ow (l ead chr omate) ; for bl ue col or P ru s s ian blue The i j ecti
co l o , ,
. n on

uid will k eep i de itely n n .

In j ec tion is u su al ly made in to the foll o win g vess el s : in el asmobr an chs the ar teries are ,

in j ec ted by way o f the ca u dal ar tery exp os ed by cu ttin g acro ss the tail ; the hep a tic po r tal ,

sys tem is in j ec ted by way o f the hep a tic p or tal vein n ear the l iver b ack war d or in bo th d irec

tion s or in to the pos terior mes en ter ic vein f orward I n j ec tion o f the sys temic v ein s is mo r e .

diicu l t an d is u su ally n o t d on e ; a metho d is giv en by Ran d in the Amer ica n Na t al is t Vol ur ,

.

XXXIX I n Necturus the ar teries ar e mos t eas ily in j ec ted b y way o f the bulbu s
a terios u s
r T he hep a tic por tal sys tem shou l d b e in j ec ted in bo th dir ec tion s in to the hep a tic
.

p or tal vein j us t before it en ters the l iver T he p o s tcaval can b e in j ec ted if des ired thr ou gh .
, ,

the l ar g e hep a tic vein s een o n the v en tr al s u f ace o f the l iver j u s t b ehin d the tr an sv e s e s ep tu m r r .

The ar terial sys tem o f the tur tl e is in j ec ted either back wa d by way o f on e o f the caro tids or r

in to the l e f t aor ta away fro m the hear t I t is prac ticall y imp os s ibl e to wor k ou t the r en l . a

p r tal an d hep a tic por tal sys tems on pres erved turtl es un l es s they ar e in j ec ted In j ection is
o .

very eas il y mad e o f bo th o f thes sys tems by way of the abdo min al vein e T he p l as tron is o f .
,

c ou r s e to b e removed by s awi g acr os s the b ridg es

, I n s e t the cann ul a in to o n e o f the a b domi
n . r

n al vein s abou t hal fway b etween the hear t an d p el vis an d i j ec t fo r war d I n j ec tion o f the n .

ar terial sys tem of the p ig eon c an b e d on e by way o f the p ec to ral ar tery to war d the hea t r .

A cording to P ar k er the sys temic vein s an d the hep a tic p ortal sys tem may b e in j ec ted by way o f
c

the co ccyg eo mes en teric vein in bo th direc tions

o
The arterial sys tem o f mammal s is i j ec ted
,
. n

either b y way o f the c ar o tid o r the fem oral ar tery the can n ul a bein g di ec ted t war d the hea t ,
r o r .

T he vei s a e u su al l y n o t i j ec ted b u t it is s ta ted tha t they m ay b e in j ec ted by way of the

n r n ,

e tern al j u gul ar to war d the hear t

x The he p a tic p or tal sys tem may b e in j ec ted by way o f the
.

h p a tic p o r tal vein n ear its en tran c e in to the liver

e I n in j ec ting ar teries an d vein s in the s ame .

s p ecimen diffe en tl y c l o ed in j ec tio n uid shou ld o f co ur se b e us ed

,
r o r s , ,
.

4 Pr . v tio o f s p e im
e s er s
a S p ecime s a e usu all y p es er ved in 5 p er cen t fo r mali
n c en . n r r n

( 5 c c o f co mmercial formali pl u s 9 5 c c o f wa te ) A s lit should b e made in the b ody cavity

. . n . . r .

o r the fo rmal in sho uld b e in j ec ted in to the bo dy ca vity th o u gh a s m all o p enin g to ins ur e p r es r

erv a tion o f the vis c era A p r ti n o f the r oo f o f the s k ull s hou l d b e remo ved in the s m al l r
. o o e
37 2 LAB ORAT OR Y MANUAL FOR W RT E
W TE ANA OM T Y
specimen s in o der tha t th b ain may harden In the case of mamm l s addition al measures
r e r . a

a e n ces s ary in o r der to in s u r e p res rv a tion

r e Mammal s Shoul d b e emb al med This is don e . . e

by in j ctin g a embal mi g uid i to the blood vessel s b efore the in j ec tion u id is sen t in The
e n n n .

em b ahni g uid m ay c o s is t o f 5 p r c en t for malin o r b etter 5 p er cen t fo r m al in p l u s on e s i th

n n e -
x

its vol u m e o fglycerin A s til l b et ter b u t more expen s ve emb al mi g uid is o ne u s ed fo r hu man
. i n

bodies with the foll owi g fo mul a :

,
n r

P ar ts b y Vo l u me
Fo r malin
Car bol ic acid ( mel ted cr ys tal s)
G lyce in r

Wa ter

T he em bal mi g u id i in j ec ted pr fe ably in to the femoral ar tery thr ough a can n ul a in the sam
n s e r e

way as al ea dy d s c ibed fo r the in j ec tion u id

r e r B en s l ey r ecommen ds tha t the in j ec tion o f .

th emb al min g u id shou l d b e d on e n o t with a syr in g e b u t by a tta chin g the can n u l a to a

e , ,

recep tacl e con tainin g the emb al min g uid el ev a ted ab ou t thr ee f eet an d a ll o win g the uid to

r u n i to the v s s el u n der this p res sur e for abo u t two ho u rs

n e The a n imal Shou l d b e arr an g d . e

in a p osition s itabl e for dis s ec tio n with the l imbs s p r ead well ap ar t an d the head til ted back
u ,

war d .Af ter the embal mi g the in j ection u id is ru n in to the s ame cann ul a by a t tachi g the
n n

syrin g to it ; B en s l ey rec o mm en d s tha t twen ty fo ur hou rs el ap s e b e tween the e mb al min g an d

e -

the in j ec tio n with the c ol o r ed u id .

I f the a imal has been thor ou ghly embal m d it wil l k eep withou t bein g immer s ed i a
n e , n

pres ervin g uid I t Shoul d b e preven ted from becomin g dry by bein g p l aced in air tight
.
-

recep ta cl s o r wrapp ed in cl o ths s at ur a ted with the f or malin g lyc er in s ol u tion

e I t is bes t to -
.

s p o n ge the hair wi th a mi tu r e o f al cohol an d wa ter c on tain in g x p er cen t f or malin The 2 .

animal s m ay ho wever if p r eferr ed b e i mmersed in I p er c en t fo r malin s ol u tio n

, , ,
D u i g the . r n

dissec tion the an imal s shoul d b e k p t w apped in cl o ths mois ten ed with formal in glycerin e r .

Fur ther detail s o n thes e ma tter s will b e fo u d in : B en sl ey P actical An atomy f the Ra b bit n ,
r o

( Un ivers ity of To on to Pr s s) ; Reighar d a d J e i g s Ana tomy of the Ca t ( Hen ry Hol t an d

r e n nn n ,

Comp an y) ; an d Davison Mammal ia n A na t my ( P B l kis to n s S o n an d Co mp a y)

,
o . a n .

5 Deal e s
.
P eserved an d i j ec ted ma ter ial n c s s ary in the c ou rs e can b e o b tain ed f o m
r r n e e r :

The Gen e al B iol ogical Su pp ly Ho u se 1 1 7 7 E as t 55th S tr eet Chica g o

r , ,
.

S u pp ly Dep ar tmen t Marin e B iol ogical L abo a tory Woods Hol e Mas sachus etts
,
r , , .

The An gl er s Co mp an y 1 53 4 W es t L ak e S tr et Chicago

, e , .

L ive Nectu r us t ur tl s p igeo s e tc a e al s o ob tain abl e fr om the rs t an d thir d deal e s

, e ,
n ,
.
,
r r

n amed ab o ve The G en ral B i l o gical S u pp ly Hou se c n al o fu r nish p p a ed S k l eto n s o f

. e o a s re r e

a n y f o ms F o r the ames o f o ther deal er s con sul t S cien ce Tran sacti n s of the A m ic
r . n ,
o er an

Micr oscopical S ociety an d simil ar j ourn al s , .

3 74 LABORATORY MANUAL F OR VERTE BRATE ANATOMY

9 3 9 4 ; hea rt 88 , 2 48 , 2 6 5- 6 6 hi d C hl 3 Cr os s o pter ygii: l i ti f 6;
C hl d t 3 5
1 ; n oc ea 2 c as s ca on o
li m b 8 48 5; o hy id a p p ara t u s 1 26 oc ear uc 2 n s 79 8 9 r ib 6 7 ; k l l s s u 102
1 8 6 ; ja w 1 1 8- 1 9 , 1 24 ; m l u sc es 136
,

C li t y t 6 ; d g h 9
oe ac a r e r : ca 2 2 o s 21 C ru ra f d i p h gm 3 3 8

o
,

f p i
,

a ra ; o en s
yt m 3 3 3 59 ; p ; pig bbi t 6 6 ;
,

57 : n e r v ou s s s e -
ec 2 20 4 5; eo n 2 ra 2 0- 1 9 2 2
l gi d l 9 9 3 ; p l i g i d l 8 k t tl l i ti
t o ra
84 ;
7 5;
r

p i t y y t m 8 6 89 ; i
res
g 3 4 5 5; k ll 6
sen s e o r
e
ra o r
2

an s
s s e
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s
-
r

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t 3
s

C li p l
oe
; t a e 2 20
C li g g li 3 3 8
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36

m mm l 3 3 8 ; p ig
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exu s :
on
ur e 2

a a eo n
C y pt b
r
f limb
o re
r
h
9 ;
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e
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s
93 ;
33 7 39 ; t t
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,

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d i i f 33 4
ee

e asl m b h 3 7 9 ; f t i l
o ran c s 0 -
unc on a C l d i ti f 6 ; m mm l
o on: e n on o 1 1 a a
e
D ml b
res s o r
M mb
e n t on o 1 , 1 1

p t ar s f 97 98;
o m mm l 3 4
2 a a 1 9 3 9 4; t
1 l 74 u rt e 1
b
er a 58 S o ne . ee e ra n e

3 5 59 ; N

2 -
3 9 ; p ig
ectu r u s 1 21 eo n
,

C l m l l p ig 3 3 ; t tl 3 4
o u e a

eo n 0 ur e 2
D ml
on e
k l t d i ti f 4 7
3 3 03 33 133 ; t t l 2 3 4
-
5 3 7 ur e 2 2 2 C l m
o u 47
n a e ca r n a c 2
er a
D m l p p il l f f t h 5 53
exo s e e on , e n on o

C t l 79 8 8 S d L i m b
en ra es
.

ee u n er
.

s C mm
o d i l i d l pm t
o n ca r na ve n : ev e o en
er a
h i 55; f p l i d l 4 7
a a: or ea er 2-

C t m f m t i f6 ; typ f76 9 ; m mm l
f
.
,
or a r or aco s ca e
en ru
C p h li t i 3
: or a on o 1 es o f
o
69 ; N t
3 ; d g h
20 7 o

5; p ig
s 20 , 20 a a
D m t m 43 4 5
er a o e
D m i 4 5 46 i f m t i
,
e a za on 2 ec u r u s 222, 22 eo n
f
C 5 k t tl
er s : n or a on o
t
f t h 5 54 : i f m t i f h
,
e re 2 s a e 210 : 4 ur e 2 0

C b ll f C mm ti d t y m mm l ea e rs
l 4 7 49 5 ; i f m ti f
1- n or a on o s

h
ere e ar 3 o ss a 1 2 1 1 o on ca ro ar er : a a
C b ll m h 3 8; 59 6 ; p i g
,
s ca e s 0 n or a on o
l m
i 55; i f m ti p t il
- - , ,
2
57
b bi t 6 3
er e e u : e as o ra n c s 0 2 40 eo n 2 4 4
m mm l 3 53 3 57 ; N t 3 ;
a a ec u r u s 20 C mm i l
o i
.

on t y ac a r er : ra 2
pl t
a r f n or a on o re e

p ig 333 ; t tl 3 4 tl
,
a es5 0

C b l
eo n
3 1 , ur e 2 t
C mm i l i
ur 37 38 e 2
i D es c e n di g n S P
v e n a c av a l ec r eca v a
t 65
D
.

er e ra 35
ossa 1 2 1 1 o on a c ve n : c a 2
C b l h mi ph
er e ra e l mb h s
,

e re
:
e as o ra n c s C mp i
o i
a r so n s , l t y y t m c rc u a o r s s e
es c e n t ft t
D l p m t f h d t 3 44
o 9 9 es es , 2 0 , 2 1

3 7 3 8 ; m mm l 3 53
0

1 3 55 56 a a -
m mm l 66 7 N t
a a 2 9 3 ; o ; ec u r u s 2 2 -
o
ev e o
Di p h g m 59 8 8
en o c or a es 1
-

3 57 58 ; N t ; p ig p ig tl
. . .
a ra 1 1
47 4 8 ; t Di t m 5
- , ,
3 ec u r u s
33 20 eo n 1 eo n 2 4 4 ur e 2 0 1
tl 3 4 3 7 C h 5 3 ; m mm l
h
.
as e a 1 2

Di p h l 9 7 3 596 ; l m
h
-
33 3 33 ; t
2. ur e 2 2 on c a 10 00 a a 1 21 2 2,
C b lp d l l m b 3 5 ; p ig tl 3 3
. ,
en c e a on 2 0 e as o

3 7 8 3 7 3 8 ; m mm l
3 8 b
,
ere ra e un c e e as o ra n c 1 ; 0 3 9; t eo n 2 ur e 2

m mm l 3 54 ; N t 3 C dy l f l i m b 8 3 8 4 ; f k ll
.

ra n c 0 1 1 a a

i
, ,
a 21 on e: o s o s u
3 53 3 54 3 59 3 57 3 58 ; N t
a ec u ru s

C b p i l i d 3 4 3 53
,
ec u r u s ,

; p g 3 3 3 ; t tl
. , . .
ere ro s na u 1 99
C i l pl m mm l 3 3 4 C dyl i d p
h
,
3 203 21 33 eo n 1 ur e
j w 4
i i
, ,
erv ca exu s a a on o ro cess , a 1 2
C i b i l pl
erv c o ra c d g h
a exu s : o s C j ti
on unc l mb h 3 4
va e as o ra n c 0
3 4 3 7
Dg t
2
g
2
y t m 59 9 9 l m
m mm l 3 4 5 3 49 ; p -
.

3 0 1- 2 ; k t 3 s a e 02 a a ig 3 9 eo n 2
es
b h 64 66 6
ve s
68; g
s e
l 1 : e as o

Ch b ll i g t 7 m m C t f th 5 5 5 - -
.
1 en era

ra n c 1

9 6 3 ; m mm l
,
ev r o n o n es : a a or o; a on o u r ea er 2 2 4
C t i l mb h 7 86 8

h
, - 1
3 a9 94 a 1
f 6
1
m l 76 a o n u s a r er o s u s : e a s o ra n c 20
,

d iti f 8 ; m mm l 63
69 7 ; p t -
C ; N t tl
,
t ar s o 1 1

cc u ru : 1 0
i g
,
221 9; t
ec u r u s 2 2 2 , 2 2 3 ur e 2 0
"6
an a e n on o 1 1 a a

6 8 8 8 ; m 1-

h
,
7 77 1 0 1 2 su
m y 89 9 t tl 73 75
p eo n 1

w
, ,
1 3
39
C d i m b d i d fm
-
on ro c ra n u : o n es er ve ro C o n ver g 4 en ce
ar 1
lk 8
: ur e 1

10 3 5; d l pm t f 9 6 9 7 ; ev e o en o C p d m 87
o ro a eu 2
Dig itig
Di p h y
ra
l t il 3
e a 2

d g h 9 7
o s ; N t
-
91 00 ec u r u s 1 0 1 1 1 C o ra co i d 8 6 S P t l gi d l ee ec o ra r e
c erca a 2

C h d t di m mm l 6 6 ; C id p Dip l p dyly 6 6 4
.
,
os on 1
86 9
Di g 4
,
or ae en n ae : a a 2 o r ac o ro ces s 2

p ig C i m 4 46 ,
ver en c e
k ll 6
g l p m t f3 ; l m
or u
Ch d t h t f 5; l i
eo n 2
4 D g
C
i
,
o : s u 10
D g h t i l y t m
or a a : c a ra c er s o c a ss ca o r n ea : ve o en o 00 e as o
ti on f 6 7; o mb y i d l p
-
e r on c eve o b h 3 5; m mm l 3 49 ; p g
ran c 0 a a eo n
b i 3 7
o s : a r er a
h d
4 ; s s e 21 21

m t f3 4 4
en o 1- 3 9 2
ra n 9 3 7 8;
i m 9 7 9 9 ; l i ti f 6
0 , 1 -
1 c on ro

t b C y t y fh t l m -

h
c ra n u c as s ca on o
Ch d or t
oc en r o u s ver e
C i id t y d l pm t 3 ;
65 ra o ro n a r
b h 5 6 7 ; m mm l 57 ;
a r er , o e ar : e as o
l m 6 3 6 4 6 6 6 7;
c oe o 1 i l-
1
-
c ran a
or o
l m b
c oa
h 3 5; m mm l 3 49 :
, e e: e ve o en 00
t tl
ra n c 21 , 21 -1 a a 2
n erv es 3 9 5; dig t i 0 ytm -
1
,

es ve s s e
e as

p ig
o ra n c 0 a a
C
ur 35 e 2
y l ig m t 6 6 6 9 59 6 ; 64 6 6 6 7 6 8; k l t 48 ;

h
-

l t m y 6 7 ; 7
-

h
1 1 exo s e e on

h
- ,
eo n
3 9 3 2 0 o ro n a r a en 1 0 1 1

C h i id p l b i l mb l mb 6 6 ; m mm l
t
, ,
ex e r n a an a o 1 1 ns 1
gill 6 8 ; t 7
,
or o ex u s , ra n : e as o ra n c s e as o ra n c 9 ; 1 a a 1 2

3 8 ; m mm l 3 53 3 56 ; N t N t 7 ; pig 8 ; t tl
s 1 ; ea r 20 221- 22
0
tl 3 4
a a , ec u r us ec u r u s 1 0 eo n 1 1 ur e
m t i 65 6 6 ; m l 8
es e n er es 1 usc es
,

1 2

3 ; my t m
; t
Ch i 9
3 20 ur e 2 73
I
64
or on 2 94 2 C o ro n a r y i 53 65
s nus 2 2
0
y t m 3 8 ; p t l 8 9 ;
9;
o o es ,
1 2 n er vo u s

C i l i y b dy m mm l 3 49 ; p i g C y i bb i t 5 ec o r a n
,
s s e 0 1 1
ar o : a a eo n
C
o ro n a r
id p
ve n : c a t 2
53 ; ra 2 1
p t l gi dl 8 8 8 9 ; p l i
ec o ra r e - e v c n

pi t y
f j w f
8 ; p l i g i d l 79 8 ;
3 o o ro n o ro c ess :4; o a 1 2 o -
C i y g g li
l ar an t 3 48 ; l m on : ca e as o l 93
u na
0 e v c
y t m 6 7 6 8 ; ib 6 6 ; p i l
r e 0 res ra o r

b h3 C p bi g m i 3 8 S Op ti -

h
s s e 1 r s s na
ra n c 1 1
t 3 48 ; l m b
or
lb
o ra e na 0 . ee c
n er ves ; g it l y t m
0 1- 2 u ro en a s s e

8
3
Cih y ar n er ves : ca e as o ra n c
C p
o es
t tl
2 83 ;
0 y t m 7 v en o u s s s e 20
-
10

Ci 3 1 1 o r o ra
m mm l 9 9
ca v er n o sa
05 8 5; 2 ; ur e 2
2 1 2- 14 t b :
l l m 64 6 6
v er e ra co u n
on a 1 2 - 1 3
C i l ti t h g h h t l C p l t 89
a a 2 1 2
i k l t 99 -

h
v sc era s e e on 1 00
rcu a
b
on

; m mm l
m
ro u ea r : e as o o r o ra
C p q d g mi 3 53 3 6
u ea 2
D o rs al t ao r a l mb h 20 2 : e as o ra n c s

; m mm l 6 6 4; N t
ra n c 2 21- 2 2 66 6 a a 2 or o ra ua ru e na 0

N t 9 ; p ig C p ll m 3 53 3 55 56 8 0 - ec u r u s
,
21 21 a a 2
ec u r u s 22 47 ; t t eo n 2 ur e or us ca osu
pig
t tl
t i t m
8 46 ;
C p f ti ur e
-
,
7
22 ; 2 4 eo n 2 4

ii
-
or s r a u 33 : unc on ,

C i l t y y t m 7 d l p
2 39 4 0 us
rcu a o r s s e 20 0 2 2: e ve o m mm l 3 58 ; p ig 3 3 3 ; t t
a a eo n ur e
3 7 38
2
D l l m p l d 97
-

m t l mb h 7 o rsa
D l m p l
co u n, s na co r 2
7;
w
en 20 1 e as o ra n c s 20 3 7 2
22 ; m mm l 48 7 ; N t
a a 2 -
2 er u r u s C p l
or u s c es 2 0 0
o rsa
D f t h 5 53
ra us , s 99 na n erve 2

2 2 2-
3 ; p t f ar s ; p ig 20 0 - 20 1 C t b i 3 7 o
D t fC i
n ea er
C mm
2-

l i ti f 7 ; j w
0 o eo n o r ex , ra n
S
2 4 48 ;
1 mm y 7 7 ; t t l su ar 2 0- 2 ur e C tyl
o w
os a u r z c a ss ca on o a
uc
di l i
o 3 u v er 20 ee o on

23 0 1 108 p t l gi d l 8 7 8 8 ; k ll
ec o ra r e s u
ca r
D d m l m b h 64; m m
na ve n

w
, ,
uo en u : e as o ra n c 1 a
Clados e che 79 m l 9 9 3: N t 6 9 ; p ig
Cl i h d t 6 C p gl d 9 1 eo n
-
a 1 2 ec u r u s

8 8 ; t tl

a ss i ca t o n o f c 8 o r a es o er s an 2 1
C l 3 54 3 58 C il 1 1
-
2 74 ur e 1
a va
C l i l 8 7 8 8 bi d 9 ; f g
,
ra n a
l m b h 3 9 5; m mm l 3 3 4
9 6
n e rv e s 99 3 6 6 2 , 2 , 0 1
D u ra m m mm l 3 4 3 5
a t er a a 1 2,

p ig tl 3 4
,

$
av c e r ra n c -1 a a
; 1 ro 90 ; e as o 0
m mm l 9
a a
,

2 37 338 34 45 3 4 6 47

3 4 8 2- -
eo n
33 t 0 , ur e 2

Cl g 3 3 4 g
.
. . .

t 20 - 2 1
d l pm t f
ea va e 2 ce u ra s
3 ; p 33 3 ; i eo n 1 2
C l it h m 8 7 8 8
e ru t l 3 5 7
ur t e 2 -2 E ar 3 36
-
0 1 1 : e ve o en o

Cht i m mm l 8 9 9 ; t t l C i m S Sk ll
i
,

or s : a ur e 28 5 ra n u ee u 3
00 ;
-
0 1 m b h 3 5 6; e as o ra n c 0 -

a 2 2 0
Cl C b if m pl t ti f 3 6 ; li d 3; m m
.
,

o a ca 16 79 8 d g h 6 5
1 2 0 o s 1 28 1 ; r r or 35 a e 1 21 0 un c o ns o 0 zar 2 a .

m mm l 79 8 N t C i id t il g m m mal 1 8 7 ,

i
.
,

a a s 2 -
0 ec u r u s 16 9 r co ca r a e a m l 7 35 5 ; N t
a 2 0- 2 ec u r u s

g l 36 ; p g
,

8 4 ; p ig k t i
i
, ,

8 8 65 7 8 ;
eon 1t t 76 ur e 1 p t ar s of3 3 3 0 00 - 01 1 eo n
2 eo n 1 2, 2 s a e 1
p
5 6 33 ; t l tl
, ,

t tl C t 3 6
,

28 2; 75 8 5 6 ur e 1 2 r s a 0 2 -2 , t ; t
0 4 e eos 20 ur e 2 ,

C y f g 6 9 m 76 C p 6 78
,

oc c x : ro ; an ro 1 1. 1 3 23
 INDEX 3 75

Ea r
3 51
b o n es

3 52

1 27
N
30 1 :
ectu r u s
m mm l
a

1 10 ;
a

p ig
1 20 ,
eo n
E xt ern a
b ra n c
l
h 7; 21
ca ro t imd mm l y a
art e r
a 2 59 - 6 0
el a s

;
0
mo
Nec
G g li d i ti f 9 6
an
G id
ano
on ,
l i ti s h e
f 6
es :
n on o
c a ss
2
ca on o
ig klt l t my
.

3 3 o ; t u rt e 3 2 4 l tu r u s 2 27; p
45 eo n 2 exos e e ont e x ern a
d m 6 S ymp i liz d
a na o
E ar ru 1 2 ee T an c mem E l
x t er n a 3 3 36 ea r 0 01 1: ar ; l im b
2 1- 22 m 79 ; p t l s ro ns
b m mm l p ig k ll
ec o ra
.
, ,

ra n e 3;2 7 35 ; a a 2 1 eo n 89 ;
n 3; t b ls u 1 0 1- ver e ra
E t d m d i ti
c o er : e n on of 34; d i er va
5 6 33
2 2 0
,

co ul m 6 3 64 n -

i E t l g it l i 9 8 9 9
E
b
t y es o f 4 1
ere n t
h 6 8; N t
ra n c
b i
s 21
l m
ra n c

7
hi l 1
a a r te r es :
ec u r u s
e as
22
o .
x er n a
E t l li
x er n a
6 4 ; t tl 38
t y
ur
i
m mm l 6 3
en
a c a r er
e 2
a a 2 ,

:
2
a
, 2 0
a 2
G id l
G i
G
an o
an o n 2 1
ar pik
9
49 sca es 2 1

L pid t
e
,

cc
,

e os eus
Eg f3 ; k i d E t l i li i m mm l 6 4 6 5 G i g g li S S m i l
.

l: c eav a e o f3 2 n s o 1 x ern a ac ve n : a a 2 -
as s er a n an on. ee
i m 97
e un ar
E m b ii
o ra n ch d : c on r oc ra n u Et l j g l
x ern a i t 54;
u u ar ve n : ca 2 g g
an on
1 00 ; i lt y ytm 7 ;
c rcu a o r s s e 20 - 22
N t 5;
ec u rus bbit 5 ; t tl 22 ra 2 2 ur e G t i pl
as r c 338 ex u s
l i ti f 6 ; l m 6 3 6 4
c ass ca on o c oe o 1 -
342 G t t
as roc en rou s ver e t b 7 ra
y t m 64 66
1
66 6 7 ; di g t i d l pm t f 3 l 3 4 m i t 3 5; Am
h
,

1 -
es ve s s e 1
-
Ey e: eve o 36 en o 00 1
'

G t as ro co e : a n o es
l mb m mm
i
,

k l t 4 7 48 ; t
,

6 68; - -
ex er -
phib i A mphi
7 t my 6 9 ; m t
1 exos e e on e as o
3 4 5; ran c 0 a a a 34 oxu s
G t l 3 4 m i t 3 6 ; Amphibi
na
1 65 66 ;
p tec o ral
-

ana o

8
y t m 3 8;
p t l g i d
n ervou s
l n
1 1
s s e
ec o r a
esen er es
0 1- 1
r e
Ey
3 5 49 : p i g
ll
e l mb
a
m l 3 49 ; p i g
a
-
s c a ; t tl 3 3
3 4 5; m m
: e as

3 9 3 ; t tl
eo n
o
eon
ran c
a - o
h 2
0
-
-

0
ur e
a
ur
2

e G
a s ru a

i
3 4; A mphi
en cu a e
34
l t b d i 3 56 3 58
: a

o
oxu s
es
n o es a

88 89 ; p l i 8 ; p l i gi dl G i l t g g li 3 3
,

e v c n 0 e v c r e 3 3 2 en c u a e an on 1
pi t y y t m E y l i d d g h 7 ; l i z d 3 ; m m

79 8 ; 6 7 6 8; ra o r
-
G mi l d i k 3
er
g i t l y t m 8 8 3 ;
0 r es s s e 1 e s: o s 1 ar 2 a na s 1
u ro en a t s s e 2 0 ver e m l 7 ; m l f i m mm l
a s 2 u s c es o n a a s G m l y 3 4 3 6 ; d l pm t f
er a ers eve o en o
b l l m 64 6 6 ; i l k l 3 46 3 4 7 ; p i g s; t tl
i h
, ,

ra co u n -
v s c era s e e 4 eon a ur e 2 6 44 -

Ey m l 99 3 G l 6 l mb
w h
.

t on9 0
-
1 00 e u s c es t 3 4 7 48 ; 2 11 : ca - 1 1 68; N
e as o ran c 1 ec
E l b bi d 9 ; m mm l 9 3
o r 2 a a s l m b 3 4 5; p i g 3 9 ;
e as o ran c
,
-
eo n 2 t u rus 2 2; t l t
G ill h 9 9
s
:
0 e eo s 20
El t d i ti f 3 4
eva o r , e n on o 1 1 0 bbi t 3 46 4 7 ; t tl 3 3
ra ur e 2 a rc 5 7 6 7 1 00 10 10 1 2 - 2
Em b y l l ig t ti l g
FF
, , ,

t 94; d g 5 6;
Em b y l g y 3 -
-
r o : ca 94 2 o s 2 a a or
3 1 1 1 1 1 ca r a e
m i t 34 36 b f m 5; d g h 9 9 ;
h
,

r o o43 1 : a n o es acet7 x. 7 4 . 7 s o n es ro 10 o s 1 00
il l mb m mm l m mb b
, ,

37 39 ; A
-
mph t b t a 2 -
3 . 3 4 . 3 5. 3 7 3 0 : ac a n erve 3 1 3 , 36 0 : e a s o ra n c a a 1 26 ; e ra n e o n es to
Amphioxu s 3 2 , 3 4 3 6 3 3:
1 a a -
m mm l
3 43 44 . 3 55; Nec 1 0 7; N ectu r us 1 1 1 - 1 2 ;
3 1 5: n erves
Emb y i m mb ig l l
,

r on c 9 94 e ra n es 2 2- tu r u s 3 2 1 ; p eo n 3 3 2 ; t u r t e 3 2 6 , ig eo n 1 7 7- 78 ; t u r t e 1 1 5- 1 6 , 1 75
E ml l 4 t th 4 9 G l li s ts 1 6 1 : A mphzox us 1 0 - 1 1 ;
5
na e : s ca e ee 37
E d l ym p h ti l ig m
h
,

n o t 7 98 3 6
a c uc s 1 0 F a cifo rm a en t 1 96 : el a s mo Bala n og 0 53 243 1 d o g s h 1 7 , 1 6 8 ;
E d l y m p h ti f 9 8
n o a c o ss a
, ,

bra n c 1 66 ; a a 192; Nec m mm l m y

l a p re 1 5; cetu r a s 2 2, 1 71 ;
l ig k l
E d k l t 57 7 h d
n
u
os e e on
m 9 6 9 9 ; d i ti
gi ll h 9 9 ; l i m b 78 9 4
f 4 5 57;
i -

e
1 2
n
:
on
c on
o
roc ra n
FFtu ru s 1 70 ; p eo n 1 8 0 ; t u rt e 1 73
ll i
a op a n tu e
i 3 34 36
S ee t er n e t u e b b U i s a te 1 9 , 1 6 8 ; te eo s t 2 0
G i zz d 8 ar 1 0 1 81 1 82
m mm l
FF
.
, , ,

a rc es -
1 00 s -
a sc a 1 0 1 1 C l an s , of penis : a a 29 1 ;
p t ar s o f 57 ; p t l g i d l 8 59 4 ; ec o ra r e i l 35 36
as c c u u s 1
,
-
,

l 86
tu r t e 2

p l i g i d l 78 8 5; ib 57 7 7 th Gl id f 9 S P l gi dl
s
e v c
k ll 6 7; t
u 0
-

l m 57 6 6 ; i l
r
m 85 9 4;
1 2
e
s er n u
-
r
-
s
ver F ea
e o ra
5 6 5 54
ers 2
m l t y m mm l 6 4 ; N
8 ; p ig
a r er
-
2 .

:
2-
a a 2 ec
en o
G l m l 73 74 75
o
Gl p h y g l
eru u s 2
to ssa 0 .

2
ec
2
ec o ra r e

F
, ,

t b e ra l co u n v s cera t u ru s 2 2 46 eo n 2 o sso 36 arl m n ea n erve o : e as o

s k l t 99
e e on - 1 00
m l i m mm l 6 5; p ig
e o ra ve n : a a 2 eo n b h 3 4 ; m mm l 3 43 3 5
ra n c 1 a a
E d ty l
n os 3 6
e 1 1 1 1 1 43 ; t
2 tl 3 ur e 2 1 N t 3 ; p ig
ec u r u s 33 ; t t 21 eo n 2
,

ur e
E t d m d i ti f 3 4 : d i m 79 S Hi d l im b
F
,

n o er : e n on o er va e ur ec n
G l tti m mm l 8 6 ; N t
ti
E t gl
n o
ves o f

Ep i dyl 8 4 9 3
co n
4
o ssatil g
es
77
1-
ca r
2
a e 1
t
e n es ra , o
u a
F ib l 79 S Hi d lim b
u a re
f k ll 9 7
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 376 LAB ORATORY MANUAL FOR VERT E BRATE ANATOMY
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I t d l gi dl 88 ; l 3 5 5 8 3 ; m mm l
, , ,

n ern a 3 ea r poo eve o r e sca es 2 0- 1 d g h 8

o s 2 1 29 a a 2 2

m t f 3 3 ; l m b h L zi i m p ll 3 ; l 3 N t 8 4 ; p ig k t
, , ,

en o 00 01 e as o ra n c o ren n : a u ae 0 2 ca n a s 0 2 ec u r u s 2 8 7; eo n 2 s a e

3 5 6;
0 f t i 3 6 ; m mm l
-
un c on 0 a a L mbu l p l d g h 3 ;
o sac r a exus : o s 0 1- 2 8
2 8 3 ; t tl
2, 2 86 ur e 2

35 ; N t 2 3 9 ; p ig
ec u r u s 33 ; 1 eo n 0 migmm l 83 3 9k 4t ; N tt tl 3 9
a a 0 ec u r u s 1 M es o n e ph
8 ; m mm 7 9 ; N t
ros 4 75 95 2d g h -
2 : o s

tl 3 4 3 6
,

t 8 ;
L g 6 6 3 9 6
ur e 2 2 p 3 eo n; 3 ; 2 3 s a e 0 2 ur e 22 2 o a a 2 2 ec u r u s 2 4
I t l i l i t y m mm l 6 3 6 4 m mm l k t 8 ; t tl 86
i
,

n er n a ac a r er : a a 2 -
un 1 2-
97 99 1 1 : a a s a e 2 2 ur e 2

p ig 46 ; t
eo n 2 tl 38 ur 88 89 ; N t 69 7
,

7 ; ec u r u s
,

1 0 1 1 M p t yg m 8 9
eso er u

i m mm l 6 46 5
e 2 1 1 1

p ig M t p l 8 8 S F li m b
.

I t l ili
, ,

n ern a a c ve n : a a 2 8 3 ; t tl
eo n 1 76 ur e 1 e a ca r a s ee o re

p ig 43 ; t t l Lym p h M t mi p
L ym p h ti y t m
eo n 2 3 ur e 2 2 20 1 e a c ro on 93 r ocess
I t lj g l
n er n a i 7 ; t 55
u u ar ve n 2 1 ca 2 a c s s e 20 0 20 1 Mtm e a ere 2
Ly m p h g l d M t m im
N t
t
n ero r
tl
ur 34
I t bi t l p t m p ig
5
ec u ru s 2 2
e 2
7;

a
bbi t 5 53
se
3 9
, 22

u :
ra

eo n
2 2, 2

2
L ymp h h t
L ym p h d l 9 4
;

no
fian ti
ea r s 20 0
94

u es 1
s 20 0 o n t es ne 1 e a
M t ph i d t S
e an e
M t ph
e an e
er s

76
r c
r os 2
94
2
t uc
m mm l 8 8
.

2
ee

:
U re er
a a 2
tl 3 3 Ly m p h y t p ig 8 7 ; t tl
, ,

t ur e 2 oc es 20 0 eo n 2 85 86 ur e 2 . 2
378 LAB ORATORY MANUAL FOR VE R E BRA E ANA OM T T T Y
8 3 - 8 4 ; o ri n o f 7 9 ; p a rt s o f 7 9 ;
tu r t e 8 1 ; u ro e es 8 0
l
gi
dl
Nectu rus 2 2 5- 2 6 ; p ig eo n 2 4 2 , 2 43 R ena l
po r ta ve n 2 0 6 : e as o ra n c l i l m b h
P enis 2 9 5:
l
t u r t e 2 8 5, 2 8 6
a a
81
29 m mm l
29 1 - 9 2 ; ,
P 44 .
os te r i t u rt e
3 5. 2 3 7 . 2 4 x
o r ca r i n a v e n 2 0 3 . 2 70 , 2 7 1 :
d o g s b 2 0 8 ; m a 2 6 9 ; Nec
d l
l
i
mm
-

l
21 4; N
ectu r u s 2 2 3 - 2 4
, 2 29 ; pi eo n
2 4 3 44 , 2 4 7 ; t u r t e 2 3 1 - 3 2 , 2 40 - 4 1
7
Repl il za : c l a ssr ca tio n o f 7 ; exo
. l
g

a
P i i l iy ig k kl l im b
er ca r d
1 9 9 : e as
a ca

ma l 1 8 7 , 1 8 8 ;
v t
o ra n c
1 59 ,
l mb
1 6

ectu ru s
0 , 1 9 5 9 7,
1 6 6 - 6 7 ; ma m
1 70 7 1 ;
hN P i

tu ru s 2 2 6 , 2 2 9 ; p eon 2 4 7 ; s a te
l
2 1 0 1 1 ; t u r t e 2 4o
o s te r o r c a mb h
e r o f ey e 3 0 5, 3 4 9
h
s e e to n
n

9
d mb
i
li
0
9 ;
50 52 ;

p l i
81-82;
gi p
1 1
o re
gi l
ec to ra
8 ;l k ll
91 ;
rd e
e v c r(
f
ig
p eo n 1 8 0 ; tu r t e 1 7 2 l P io s ter o r ven a ca va P
S ee o s tca va l 6;
1 1 2-1 t m 9 ; t b l s er n u
e .
1
ver e ra
s u

P i il mm l P i l b d f l im b 3 l m 7 7 S f th d
1
.

er ca r d a sa c 1 6 0 , 1 9 6 9 7 ; a ma rea x a or er o 2 co u n 0- 2 ee er
P l i 7 ; t 53 55; p i g ur un er
ig l Al lig t L i z d d T tl
R pi t y y t m 6
.

1 88 ; p eo n 1 8 0 ; t u r t e 1 7 2 73
- -
reca va ve n 2 1 ca 2 eo n a or, ar an ur e
P i d i l
er ca r i o -p e r to n ea l ca n a 1 6 7 : d o g 2
4 43 2 47; bbi t 5 53 ; t tl
2 ra 2 1 ur e es ra o r 63 9 8 99 s s e
,

1 1 1 -
:
m b h 6 7 6 8 ; m mm l
k
h i i l
, ,

s h 2 2 2 ; s a te 2 1 0 , 2 2 2 33 34
2 -
4 2 e as o ra n c -

16
s 1
P i d m l mb
e r ca r i u : e as o ran c
6 6 7; P h d l ti l g 9 7 ; b d d
r ec or
7

a ca r a e o n es er ve 8 6 88 ; N t
1 7 ; p ig 77 ec u r u s 1 eo ns 1
a a

f m 4
1
m mm l g mm 9 89 9 ; t tl 75 7 6
R i f m b ?y l m b h 3 8 ;
a a 1 8 8 ; Ncc tu r us 1 70 ; p eo n ro 10 su a 1 -

l
1 8 0 ; tu r t e 1 7 2 P m l t t h 5
re o ar ee 1 2 es t or o : e as
ur e1

P id m
er er 52 . 53 . 54 P p 9 9 9
re uce 2 2 1 2 mt mm l 3 54 3 58
a a
o ra n c 0

P i my i m P l t b R t ti 76
R ti 3 3 6 l m b h 3 5
, ,

er s u 136 ro c oe o u s ver e 69 7 76 ra 0 e e es s 2
i m
Pe r n eu 29 P roc o ra c o id 8 6 S d P t l ee
,

un
,

er ec o ra e na 01 1 : e as o ra n c 0
P ih l y m gi d l mt mm l 3 49 ; p ig 3 3
h
.
,

er p era n ervo us s s te 200 , 29 6 , r e a a eo n 0

2 9 8- 99 , 3 59 , 3 6 0 l mb ti l g R p i t l d i ti f 7 6
Rh ph l
'
6 1 : e as o ra n c s P i id
roc r co 78 ca r a e 1 e ro er o n ea e n on o 1
s o r 2 , 3 0 9 1 6 ; rn
- -
a n n n a l -
3 3 4 4 5. P t d m 6 ; p ig 8 7
roc o aeu 1 o eo n 2 i n en ce l m b h3 7 a o n : e as
,

o ran c
P d i ti f 8 R h m b ph l 9 7
s 0
34
p
6 -

ig
eo
49 ;
n 3 2 8
N e ctu

, 33 1
r
-
us
2
3 ;
3 1 8
t u r
1 9, 3
tl e 3
2
2
0
1
-

-
21 ;
23 .
ro n e ,
P p i d t 73 7 5 9 4 9 5
ro n e
P p h 73 74 75 9 4
h e
r c
n
uc
on o
2
2
2 2 2 R i b 6 7 7 llig t 7 7 ; Am
o
s
e n ce
2, :
a on 2
a a or 1- 2

i
,

bi d 73 ; d g h 6 6 ;
, ,

h bi
'

ron e -
6
P t l i ty 59 6 9 69 7
ro s 2 2 2 z a r s o s
er o n ea ca v 1 0 1 P p t ygi m 8 8 9
ro er u o,
, ,

6
n 77;
s m mm l 75;
o 2, a a s
m mm l 8 9 9 ; p ig 8 , ,

P ph l ig i f6 ; t l t 6 7 6 8 ; t t l 7

-
99 z a a 1 0 eo n 1 0 ro sen ce 97 a on 2 or n o 2 e eo s -
ur e 2
P t t gl d 9
p g
2
P t l g gl i l m b
e r o sa
l eo n
33
an
2
3 4; o n : e as o r an c h 1 P
ro s a e

P d t
seu
t i l 8 8
ro ven r c u u s 1
t b
oc en r o us ver e
an

69
2 1
1 , 1 2
r ae
R t f pi l
oo s o
m b
R t m 7 97
o s ru
o
9 8 3 59
ra n c
s
h 3 5 3 6 ; m mm l 3 4
1
l na
s 1
n er ves 2

, 1 a
,

a
: e as
1

P y p t h 94 P t yg i d f
i l m b h 3 6;
,

8

e er s a c es 1 er o 3 o ss a 1 1 1 1
P h l g 79 8 8 S d Li m b
a an es ee u n er P t yg q d t
er o ua t l g 99 ra e 5 ca r
,

a e 10 S l accu us , of ea r : e as o ra n c 0
Ph y 6 6 98 9 9 l m m mm l 3 5 ; N t m 3 9 ; p ig
h
, . , ,
-
ar nx 1 1 2, 1 : e as o a a 2 ec u 5 1 eo n
b ra n c 6 8 ; m mm l 8 5 8 6 ; N
1 a a 1
-
ec P bi 79 S P l i g i dl
u s ee e v c r e 330
7 ; p ig tl P d d l pl pig S l d 93
.

t u r us 1 177 ; t 75 eo n 1 ur e 1 u en a 3 8 exu s , eo n 2 t
a cc u u s ro u n us 1
Ph i r en c n erv e 33 35 4
-
P lm y t y 6 7 7 m mm l
u on a r ar er 2 2 2: a a S l pl
ac ra pig 3 8 exus , eo n 2
Pi a m t m mm l 3 4 3 5 53
a er : a a 1 2- 2 48 57 ; N l 2 7; p ig ec u ru s
,

22 eo n S m 69 S
a c ru d V tb l ee u n er er e ra
N t p ig l tl l m
, ,

ec u r us 3 33 ; 20 eo n 0 t urt e 2 44 4 5; t 2 35 36 ur e 2 2 co u n
P lm y pl 3 3 7 S li y g l d 6 m mm l 8 3 84
, , ,

-
52 4 u on a r exu s a va r an s 1 1 : a a 1
Pi g eo n i a r78 8 ; i l
sa c s 1
-
1 a r t er a P lm y i
u on a r 7 m mm l
ve n s 2 0 0 2 2: a a S p id d i ti f 8
auro s a, e n on o
y t m 46 ; i l y y m 56 57 ; N t 6 ; pig
,

s s e 2 44 c r c u a to r s s te 2 -
ec u r u s 2 2
4 eo n 2 2, S l t
ca es : i d 49 5 ; y l i d 4 9 ;
c eno -
0 c c o

48 l i i f 7 44 ; t t l h 47 5 ; g i d 49 ;
-
ca t o n ur -
2 4 1 c a ss 5; o 2 2 35 e 2 s es 2 1 2 2, 0 a no 21
c oe o l m 8 8 8 8 3 ; d i g ti
,

1 0 1 1 2 -
,

es ve P l p f th 5 ; l 47
u
:
ea er 2 sca e m mm l a a s
,

55; rep t il es 23 2 4,
,

2 5,
y t m 76 7 7 8 8 8 P l i 3 56 3 58
h
, ,
-
u v n ar
-
s s e 0 2 e xo
l
1 1 1 1 50 2
k l t 5 54 ; t l t my P pi l l m b S P l
, , , ,

s e e on 2- ex ern a an a o u : e as 3 5; o ra n c 0 ca p u a 8 6 S ee ec to ra
5 6 ; f th 5 5 5 ; f S i i m mm l
.

c a t c n erve : 3 4 0 ; N6 6
- -
2 2 ea er s 2 2 4 o re 3 49 a a
lim b 9 ; h t 4 4 PPyygl ty l {3 m ig l
h
,

2 6 47 ; ear 2 1
-
2, 24 - os e tu r u s 3 1 9 ; p eo n 3 2 8 ; tu r t e 3 2 2
i d limb 83 ; m
n l 79 usc es 1 o r u s : e as b h 6 4 ; m mm l o ran c 1 a a S i
cia t c ve n : p i ig
eo n 2 4 3 ; t u r t e 2 3 1 l
n er v o u s y t m 3 8 3 3 ; p t l
s s e 2 -
ec o ra 1 9 94; N t
0 1 69 ec u r us 1 S l
c era 3 0 0 : e a s o ra n c l mb
3 0 5; mam h
g i d l 9 p l i g i dl 8 8 3 ;
r e 1 e v c r e 2 P y m i d f b i 3 55 3 58
ra
,

o ra n ma 1 3 4o; p eo n 3 2 9 ig
pi t y y t m 7 7 8 ; ib 73 ; P y if m l b 3 54 3 58 S i Sl
, ,

cl ero t c coa t S oc
-
r es ra o r s s e 1 1 r s r or o e c era
m 9 ; y i 83 ; g i t l S l m d l pm
.
,

t
s e rn u 1 s r nx 1 u ro en a c ero to e 43 ; in eve o en t of
s y tm 86 8 7 ;
s e 2 yt m
-
v en o u s s s e R bbi t t i l y t m 57 58 6
a : a r er a s s e 2 -
2 0 vert e b ra e 5 6
8 -
1
t b l l m 7 73 64; b i 35 5 ; i l t y y S m
,

ve r e 2- c rc u a o r s cro t u

2 2 44 ; co u ra n s
9 ti
4 ra n 2 29 29 0
P i l b d) S E p i p h y i
n ea o ee s s t m e
4 9 6 5; 2 f 7 c ca on o
'
S c u te : a r 55; m di ll
il
,

a 5 5 o rep t es 0 2
Pi 7 3 3 3 5 3 6 l m 8 78 9 9 6 9 8 ; S gm
.

c ra t

nna 2 0 0 1 1 1 co e o 1 1 e en 2
Pi 6 S Fi h S g m t ti f h d
i i
, , , , ,

s ces ee s es n erv es 3 3 4 3 37 3 3 8 3 4 45 3 46 2- e en a on99 3 2 o ea 2 00

P i t i t y b dy S Hyp p hy i 4 7 3 4 55; d g t y t m f i lt y y m 4 6
.
. . . . ,

83 s te

-
u ar o ee o s s es ve s s e 1 3 5; 1 o c r cu a o r s 20 20
Pl l t my 7 S gm t ti i ty 3
. ,
,

a cen t a 2 94 86 9 9 4; 1 t ex er n a an a o 2 e en a o n ca v 2
P l dt l m mm l 7 9; f lim b 9 3 4 ; h t 88 S ll t i
,

ac e n a a a 2 o re ea r 1 e a 35
u rc ca 1 2 1 1
P l i l 4 7 48
aco s ca e 2 48 6 5 6 6 ; hi 2 li m b 8 4 8 5;
-
n -
,

S mi i l
e c r cu ar l m mm l 3 5 ca n a s :
,

a a 2

Pl f b dy p ig
a n es o
P l ti g d l k 8
an
Pl m
ra e
o
w a
1
2
hi d
1 1 8 9

pp
-1
4;
t
m
a 6
l
86 ; j w
y
1 2
t m 3 3 3 59 ;
a ra u s
t m
1 2
usc u ar
, 1
s s e
a
S mi i
e
3
c rc
eo n
d t
3 6 ; m mm l 3 5 ; N t
0
l m b h
r uc s e as o ra n c

y
i
,

3 4 56 ;
-
as a 20 0 1 n ervou s s s e 0 a 3 a 2 ec u r us 1 9;
Pl t 5 5
as ro n 2 2 p t l gi d l 9 9 3 ; p l
ec o ra r e 2 e v c p ig 33 ; teo n tl 3 6 o ur e 2
8 9 p ig gi d l 8 3 8 4 ; p i t y y t m S mi l g g li N t
,

Pl ei r a 1 8 8 ; m mm l 88 -
ra o r s e un ar an
3 on : ec u r u s 2o ;
3
a a r 1 eo n r e r es s e
tl 3 5 3 6
ib 75; g
h
. ,

86 8 9 ; p ig 33 ; t
-
I 1 r s sen se o r an s eo n 2 ur e 2 2
P l l i ty 59 9 69 7 99 k ll l S ml l l m b
,

6 6; pi
-
eu r a ca v 1 1 1 : 3 45 5 ; 2 s u 1 1 2 s na e i u n a r va ves : e a s ; o ra n c 221
m mm l 8 7 89 ; p ig 8 8 ,

m 9 3 3 ym m mm l 6 6 ; N t 9 ; p ig
hi
,

t
3 3 7-
- ec u r u s 2 2 eo n
a a 1 eo n 1 1 1 2 n er v es 334 4 ; 2 s ern u s a a 2
t m 334 35
y
h i h
,

tl
.

83 p t a t c s 39 s e
-
46 2 47; t 2 39 ur e 2

e
P l p i t l i t y 59 9 5 9 9 g t l S mi l i l l mb
. .

h
,

eu r o er o n ea ca v 1 1 -
: 3 43 44 ; t t 5; ee 1 2 u ro en a e na ves c e : 83 e as o ra n c 2
e asl mb o 63 6 4;
ra n c N t 1
,

ec u r u s s y t m 8 89 ;
s e 2 ytm 2 v en o u s s s e bb i t 9
ra 2 1
1 6 8 6 9 ; t tl
-
7 73 ur e 1 2- 2 49 s
-
5 53 o, 55 57 2 6 4 6 5;
0- 2 -
2 -
S mi if
e n t b l 76
er ou s u u es 2
Pl f pi l l l m 73 76 S g d l pm t f 3
h
.

F
.

ex u s o s 99 3 59 na n e r ves 2 : t b
ve r e ra co u n en s e o r a ns : ev e o en o 00
d g h 3
o s ; m mm 0 1- 2
l 33 5 3 7 a a
,

R di l 8 8 S
a a e lim b ec o re 3 36 ;
01 l mb 1 3 6 e as o ra n c s 0 2-

N t 3 9 ; pi g 3 8 ; l 89 ; p l i m mm l 345 5 t ; tlN t
.
, ,

3 39 4
-
0 ; ec u r u s 1 eo n 2 R di h
a a p t a: ec o ra n e v c n a a 3 9;
-
2 ec u rus 1
s k t 3 ; t tl 3
a e 02 ur e 22 8 o p ig 9 3 ;
eo n 3 3 2 4 0 ur e 2 -
2
P m t i f m 9 79
neu a c o ra en 2, 1 R d i l S R di l i
a a s ee a a a S y m ti f 9 7 9 8
en so r e on o 2
-

l i i f6 ; t l R di 8 8 S F l m b S p t m f b i m mm l 3 56
.
,

P ly d
o o orr c as s ca t o n o ex e rn a a us ee o re i e u o ra n : a a
t my R m mm i 99 m mm l 3 3 7 pig 3 3 ; t tl 3 7
.
,

an a o 21 a u s co u n ca n s 2 a a eo n ur e 2
S pt m m mm l 7
:
P l ypt
o l i ti f 6 ; p
eru s
:
c a ss ca on o ec t tl 3 3
ur e 2 e u O n o se : a a 1 1 1 21

l d gi d l 89 ; i b 6 7 ; R ph 3 3 36 tl 3 3
, , ,

t o ra n an r e r s a e 1 1 35 ; t 0 ur e 2
s k ll u 101 R t l gl d 6 5
ec a an
,

1 S p t m l l id m 3 56
e u e uc u

PP t 54y 55m
o ns
-
R ti 6
ec r c es 2 S er o sa ,
S m id b 83
iti f 58 e n on o 1
or a s s te 4 4 6 S 200 20 20 -
ec R m ig 6
e es 2 esa o one
t l y t m R l p l 75 l gi d l 88 ; k ll
.
, ,

f th
ur d H
er u n ti p er e c or a s s e en a co r us c e 2 S yme i p t
ou r a : e c o ra r e s u
d R lp t y t m R lp tl ytm 46 4 4
an
P i l b d f lim b 4
o s ta x a
P t l
os ca v a
en a

d l pm
or
ve n :
or a

i
er o
f
s s e

eve o
2
en t
en a
229
d g h
4 , 4
or a

47
2
m mm
69 7
0-
l 6 7
s

9 7 ;
1
s e

. 2
20

. 2
-
,
-
21
0,
, 22
2 1
Si
h 610
S l l gl d 8
e
n us
an
l m b
v en o s u s
2
7
1
e as o ra n c h 20

; m mm l 4 8 ; N t
o o 4; s 21 a a 2 0
68 7 N t 9 ; p ig ec u ru s 222
-
22 9 3 4 0 47 2 7 ;
r. 2 2 -
0 2 1 ec u r u s 3 22 47 ;
-
2 22 eo n 2 221 a a 2
m mm l 5556 6 4 6 5 6 8 7 ;
a a
,

2
.

2 -
, 2
.
-
0 sk t 4; t t 3 3 4 4
a e 21 ur e 2 0- 2, 2 0 - 1 822 9 t -tl 2 3 .38 ur e 2 0 , 2
 INDEX 3 79

Sk a te :

3 7 9 0
ar t er a

3 1 6
il y t m
8; i l 1
s s e
c rc u a o r
21 5 9; b i
-

t y y t m
1
s s e
ra n S b di l
u ca r
l m b h 4 ; i f m ti
e as
i
o
na
ra n c
4 v e n 2 0 4- 6
7 7
s 21
, 22
n
,

or
2 0-
a on
1 T
T
hh eco
o rac c
d
i d
ont 1 1
uc t 2 6 0
6, 1 24

2 1 0 -1 1
,

21 21 4, 21 5
1 9; Cl a sS l f p t
o l i
o s ca v a 9 4 68 ve n 22 2 1 2 T h o ra x 2 7, 1 8 7- 8 9 : m l u s c es of 1
l m , 39,

hym
, , ,

c a tio n 6 3 6 7; 6; -
7 7 0- r 4 4-

o c oe o 1 2 1 so
il
c ra n a 3 9
6;
n e rv es d i g ti 0 1 es ve S bl i
u c av a n t y t 8
5 59 ;
a r er ca 2 T us 1 62 m mm l 8 a a tu rt e l
l m b h 8; N t
1 9;
y t m 6 46 6 ; kl t 4 7 ;
s
t
s e
ex er n a
68; h
1
l
t

t my
7 ea r
1
8
;
an a o
9; g il l
20
e xo s

221- 22
1
e e on
1
n e rvo u s
s pig
e as

2
3
eo n
44 ;
o ra n c
bbi t 57 58 ; t tl
2
7;
ra
21
2
ec u r u s
-
ur
22
e T
T
hhyy iid
23 5
ro
ro d
ca r til g
gl d an
a e 1 42 , 1
1 61,
87
162 el a s m o
y t m 3 8 ; pi t y y t m i i b h 5; m mm l 8 7 ; t tl
,

S b t 54 ; d g h
g 3 -
s s e 0 1- 1 res ra o r s s e u c av a n ve n ca 2 o s ra n c 21 a a 1 ur e
l 47 ; 5; p i g
68; 6;
i 20 N t
8bit 5 ; t tl 3 4
ec u r u s 2 2 43 ; 2- 5
'

1 s en s e o r a n s 02 eon 2 2
T i bi 79 8
s c a es 4
pi l na n e rv es
; g t l -
2 u ro en a ra 2 1 ur e 2 a S Hi d l im b
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Un ive rs ity of Cal ifo rn ia Lib ra ry
o r to the

O O
N RTHERN RE G I NAL LIB RARY FAC ILITY
Bld g 4 00 Ric hmo nd Field S tatio n
.
,

Un ive rs ity o f Cal ifo rnia

/
l x
Ric hmo nd CA 9 48 04 46 9 8
,
-

ALL B OO KS MAY BE RECALLE D AFTE R 7 DAY S

2 -
mo nth loa ns may b e re n e we d b y call ing
( 51 0) 6 4 2 6-
7 53
0
1 yea r loa ns may b e
-
rec ha rge d b y b ring ing
b oo ks to NRLF
Re ne wals and rec ha rges may b e mad e
4 d ays prio r to d ue d ate
l
l
DUE AS S TAMP E D BELOW

DDZO 1 5M 4 02 -