A re-investigation of the histology of certain organs of the viscera of the California anchovy, <italic>Engraulis mordax mordax</italic>, Girard

PDF

Download

Open document

Flip pages

Download a page range

Download transcript

Copy asset link

Request this asset

Request accessible transcript

Transcript (if available)

Content A RE-INVESTIGATION
OP
THE HISTOLOGY 0P CERTAIN ORGANS OF THE VISCERA
OF THE CALIFORNIA ANCHOVY
ENGRAULIS MORDAX MOKDAX, GIRARD
A THESIS
PRESENTED TO
THE FACULTY OF THE DEPARTMENT OF ZOOLOGY
UNIVERSITY OF SOUTHERN CALIFORNIA
In P a r t i a l F u l f i l l m e n t
o f th e r e q u ir e m e n ts f o r th e d e g re e
M a ste r o f A r ts
by
John T. MacDonald
June* 1946
UMI Number: EP67158
All rights reserved
INFORMATION TO ALL USERS
The quality of this reproduction is dependent upon the quality of the copy submitted.
In the unlikely event that the author did not send a complete manuscript
and there are missing pages, these will be noted. Also, if material had to be removed,
a note will indicate the deletion.
UMI
Oissertatk)n Publishing
UMI EP67158
Published by ProQuest LLC (2014). Copyright in the Dissertation held by the Author.
Microform Edition © ProQuest LLC.
All rights reserved. This work is protected against
unauthorized copying under Title 17, United States Code
west
ProQuest LLC.
789 East Eisenhower Parkway
P.O. Box 1346
Ann Arbor. Ml 48106-1346
u
Thh thesis, written by
under the guidance of A,8 Faculty Committee,
and approved by all its members, has been
presented to and accepted by the Council on
Graduate Study and Research in partial fulfill
ment of the requirements for the degree of
y
9m
^>L
MASTER OF ARTS
Secretary
Date.
June 1946
Faculty Committee
ChairnMTi
CHAPTER HEADIHGS
CHAPTER PAGE
I . INTRODUCTION.......................................................................................................1
I I . REVIEW OF THE LITERATURE..................... 3
I I I . MATERIALS 4BD METHODS.............................................................................. 12
IV. HISTOLOGY............................................................................................................16
V. DISCUSSION........................................................................................................ 45
V I. SUMMARY.................................................................................................................47
V I I . BIBLIOGRAPHY.................................................................................................... 49
TABLE OP COÎTTENTS
CHAPTER PAGE
I . INTRODUCTION........................... 1
I I . REVIEW OP THE LITERATURE.............................................................................3
I I I . MATERIALS AND METHODS..................................................................................12
IV. HISTOLOGY.................................. . . . 1 6
I . U r o - g e n i t a l o r g a n s ..............................................................................16
1 . The g o n a d s . . ................................................................................. .16
Shape and a p p e a r a n c e ..........................................................16
S iz e and p o s i t i o n .............................................. 16
The d i f f e r e n c e betw een o v ary and
t e s t i s in v i v o .........................................................................16
2. The t e s t i s . . . . . . .........................................................................17
C a p s u l e ................................... 17
L o b u l e s . ......................................................................................17
S e n a t a ............................................................................................. 18
S p a c e - c h a n n e l s .........................................................................18
Median c o r e .................................................................................18
Germ c e l l s ................................. 18
N e s tin g te n d e n c y ....................................................................18
P o l l i c l e s ......................................................................................19
G e n i ta l d u c t ..............................................................................19
3 . c The 0 v a r v ........................................................................................ 25
C a p su lew .................... 25
L o b u le s ...........................................................................................25
S e p t a ................................................................................................26
S p a c e - c h a n n e l s .........................................................................26
Median c o r e ............................ 26
Germ C e l l s ...................................................................................26
N e s tin g t e n d e n c y ....................................................................26
P o l l i c l e s ......................................................................................27
G e n i t a l d u c t ..............................................................................27
4. Body k i d n e y s ...................................................................................27
S iz e and s h a p e ............................ 27
P o s i t i o n and a p p e a r a n c e ..................................................27
T u b t i l e s . .............................. 27
C e l l s .......................................................................... 27
G l o m e r u l i ......................................................................................27
E x c r e to r y d u c t .........................................................................28
2
TABLE OF CONTENTS
CHAPTER PAGE
IV. HISTOLOGY c o n t.
5 . R enal ly m p h a tic o r g a n ..............................................28
2 . Organs o f th e v i s c e r a .....................................................................29
1 . The l i v e r ......................................................................................... 29
S i z e , s h a p e ,a n d a p p e a r a n c e ........................................29
L o b u le s o r u n i t s .................................................................29
C o n n e c tiv e t i s s u e ...............................................................29
C e l l s ............................................................................................. 29
C o r d s ............................................................................................. 30
C a n a l i c u l i ............................ 30
L y m ph ocytes..............................................................................30
C e n t r a l v e i n ........................................................................... 30
2 . G a ll b l a d d e r ................................................................................. 31
3. S p l e e n .................................................................................................31
S i z e , s h a p e ,a n d p o s i t i o n .............................................31
P u lp p a t t e r n s .........................................................................32
N o d u le s ........................................................................................ 32
W hite p u l p ............................................................. 32
Red p u l p ......................................................................................32
L o b u l e s ........................................................................................ 32
3. D i g e s t i v e c a n a l .................................................................................. .33
1. E so p h a g u s .........................................................................................33
MuCosa............................................................. 33
Subm ucosa. ..............................................................................33
Muscula r i s ........................................................ 36
S e r o s a ...........................................................................................33
2 . C a rd ia c arm o f th e sto m a c h ............................................. 36
M ucosa........................................... 36
Subm ucosa................................................................................... 37
M u s c u l a r i s .................................................................................37
S e r o s a ...........................................................................................36
3. Caecum o f th e sto m ac h ..........................................................40
4. P y l o r i c arm o f th e sto m a c h ............................................. 40
M ucosa .................................................................................41
Subm ucosa................................... 41
3
TABLE OP CONTENTS
CHAPTER PAGE
IV. HISTOLOGY c o n t.
M u s c u l a r i s . . ............................................................ 41
S e r o s a ................................................... 41
5 . P y l o r i c c a e c a . ........................................... 42
6 . P y l o r i c V a lv e ................................... 43
7 . Sm all i n t e s t i n e ....................................................................... .43
M ucosa......................................................................................43
Subm ucosa.............................................................................. 44
M u s c u l a r i s ............................................................................44
S e r o s a ......................................................................... • • • • • 4 4
V. DISCUSSION..........................................................................................................45
V I. SUMMARY................... . . . . 4 7
V I I . BIBLIOGRAPHY.................................................................................................... 49
I . INTRODUCTION
THE PROBLEM. In r e v ie w in g th e a v a i l a b l e l i t e r a t u r e on th e
th e anchovy, a c lu p e o id ty p e o f t e l e o s t f i s h , no s i n g l e r e
f e r e n c e where a l l th e p u b l is h e d r e p o r t s had been c o l l e c t e d
i n t o a s i n g l e , m e a n in g fu l o u t l i n e was d is c o v e r e d .
( l ) An a tte m p t h a s been made by th e p r e s e n t w r i t e r to
summarize in a s i n g l e o u t l i n e th e a v a i l a b l e , known, h i s t o l
o g i c a l f a c t s r e g a r d i n g c e r t a i n o rg a n s o f th e v i s c e r a o f t h i s
f i s h . ( 2 ) An a tte m p t h a s been made to r e - i n v e s t i g a t e c e r t a i n
s t r u c t u r e s o f t h e s e o rg a n s a b o u t w hich th e l i t e r a t u r e a v a i l
a b le to t h i s w r i t e r h a s been uncom m unicative o r f r u g a l in
s u p p ly in g i n f o r m a t i o n , and to p r e s e n t th e f i n d i n g s more c l e a r l y
th a n th e y have b e en h e r e t o f o r e .
THE ECGNOMIC IMPORTANCE OP THE ANCHOVY. In a p e r s o n a l
i n t e r v i e w w ith D r. P r a n c i s C la rk o f th e S o u th e rn C a l i f o r n i a
P is h and Game Commission i t was l e a r n e d t h a t th e anchovy i s
c o n s id e r e d o f econom ic im p o rta n c e , b o th c o m m e rc ia lly and
b i o l o g i c a l l y . Pour m i l l i o n pounds o f E n g r a u l i s t e l o s a a re
u se d y e a r l y a s b a i t f o r fis h e rm e n in I n d ia n w a te r s a l o n e ,
( N a ir 1940) . T h is s p e c i e s i s a so u rc e o f p a s t e f o r c e r t a i n
t a b l e s a u c e s , and o c c a s i o n a l l y f i s h e r i e s can a l i m i t e d amount
o f th e f i s h w ith s a r d i n e s (Cunningham 1889, H ild e b ra n d and
C able 1930, K untz 1933, and B o lin 1 9 3 6 ). The b i o l o g i c a l
2
im p o rta n c e o f th e anchovy a s food f o r l a r g e r f i s h i s c o n s i d e r
a b le .
I t was l e a r n e d a t th e S o u th e rn C a l i f o r n i a F is h and Game
C om m ission’ s l i b r a r y a t T e rm in a l I s l a n d , where D r. C la rk i s
in c h a r g e , t h a t t h e C a l i f o r n i a anchovy i s one o f th e g r e a t e s t
p e s t s w ith w hich th e f i s h i n g i n d u s t r y o f th e P a c i f i c c o a s t h a s
to c o n te n d . O ften w i t h i n a few m in u te s a f t e r th e n e t s a re
c a s t a n c h o v ie s c lo g th e m esh e s. The n e t s must th e n be f r e e d
by b e in g ru n th r o u g h l i v e steam a t c o n s i d e r a b le c o s t to th e
f i s h e r i e s . A lth o u g h th e anchovy i s an e x c e l l e n t pan f i s h , i t
i s so f r a g i l e t h a t i t must be p r o c e s s e d im m e d ia te ly a f t e r b e in g
c a u g h t t o p r e v e n t i t from s p o i l i n g , and f o r t h i s re a s o n f i s h
e r i e s do no t t r y to c o m m e rc ia liz e i t on a l a r g e r s c a l e th a n
th e y hav e done he re t o fo r e (Ni shekawa and N a ir 1 9 0 1 -4 0 ).
Cunningham (1889) m e n tio n s th e f a c t t h a t a t t h a t tim e E n g l i s h
f i s h e r i e s were t r y i n g to d e v is e an in e x p e n s iv e means o f
s u p p l a n ti n g s a r d i n e s and h e r r i n g w ith th e anch ovy .
A c co rd in g to H ild e b r a n d and C able (1 9 3 0 ), and B o lin (1 9 3 6 ),
th e anchovy l i v e s on m i c r o s c o p i c a l m arin e a n im a ls such a s cope-
p o d s . I t i s t h e r e f o r e n o t a menace to s m a ll e r f i s h o r to th e
food su p p ly o f l a r g e r o n e s .
In r e g a r d to t h i s r e - i n v e s t i g a t i o n , th e p r e s e n t w r i t e r
w is h e s to e x p r e s s h i s in d e b t e d n e s s to D r. B. M. H a r r i s o n o f
th e zo olo g y d e p a rtm e n t o f th e U n i v e r s i t y o f S o u th e rn C a l i f o r n i a ,
and t o D r. Howard H i l l , m arin e z o o l o g i s t o f th e Los A ng eles
C ounty Museum, f o r t h e i r a s s i s t a n c e .
I I . REVIEW OP THE LITERATURE
THE OLDEST REPORTS ON ANCHOVIES. Cunningham’ s (1869)
h i s t o r y o f t h e anchovy r e v e a l s t h a t t h i s f i s h was known to
th e a n c î é n t t G r e e k s and Romans by th e name cf e n c r a s i c h o l u s ,
th e name b e in g Greek in o r i g i n ; and t h a t M edieval n a t u r a
l i s t s c o n tin u e d to c a l l i t by t h a t name.
A c co rd in g to Cunningham (1889) L in n a e u s c l a s s i f i e d th e
anchovy a s C lu p e a , and gave i t i t s a n c i e n t name, e n c r a s i c h o
l u s . L a t e r , sa y s Cunningham, C u r v i e r removed i t from th e
genus C lu p ea , and p la c e d i t in one o f i t s own, th e genus
E n g r a u l i s , t h e E n g l i s h o r E uropean s p e c i e s b e in g c a l l e d
E n g r a u l i s e n c r a s i c h o l u s .
SPECIES OP NORTH AMERICAN ANCHOVIES. The two s p e c i e s o f
a n c h o v ie s on th e A t l a n t i c c o a s t o f N o rth Am erica a r e M i t c h i l l i ,
I n v e s t i g a t e d by K untz (1 9 3 3 ), a,nd e p s e t u s , i n v e s t i g a t e d by
H ild e b r a n d and C able (1 9 3 0 ).
There a r e f o u r s p e c i e s o f P a c i f i c c o a s t anchovy of N o rth
A m erica, th e l i s t o f them b e in g a s f o l lo w s : ( l ) The o f f i c i a l
C a l i f o r n i a anchovy i s mordax mordax, G i r a r d , o f t e n c a l l e d th e
b lu e anchovy b e c a u se o f i t s c o l o r (Hubbs 1 9 2 5 ). The o t h e r
a n c h o v ie s a re g re e n f i s h e s , th e t y p e s b e in g : (2) d é l i c a t is s i m a ,
G i r a r d ; (3 ) s c i t u l a , P o w ler; and (4) co m pressa, G ir a r d
(B a rn h a r t 1 9 3 0 ). The ty p e g iv e n s p e c i a l s tu d y in t h i s
i n v e s t i g a t i o n i s m .m ., G ir a r d
4
THE SIMILARITY OF ALL ANCHOVIES. A f t e r c o n s i d e r a b l e r e
s e a r c h on th e Am erican A t l a n t i c c o a s t anchovy, e p s e t u s ,
H ild e b r a n d and C able (1930) co n clu d ed t h a t t h i s f i s h i s e s s e n
t i a l l y s i m i l a r in m orphology, h i s t o l o g y , and b e h a v io r h a b i t s to
th e E uropean ty p e , e n c r a s i c h o l u s . These a u t h o r s f u r t h e r con
c lu d e d t h a t th e o n to g en y and em bryology o f th e Am erican A t l a n t i c
a n c h o v ie s , M i t c h i l l i and e p s e t u s , a re th e same, w ith s l i g h t
v a r i a t i o n s ; and t h a t a s f a r a s i s known th e t r u is m s o f e p s e t u s ,
w hich in m orphology i s a lm o st i d e n t i c a l to th e C a l i f o r n i a sp e c
i e s , mordax m ordax, G i r a r d , h o ld f o r any o t h e r t y p e . These
a u t h o r s r e p o r t e d t h a t many o f th e d e t a i l s o f th e h i s t o l o g y ,
spaw ning, and l i f e h i s t o r y o f a n c h o v ie s a re o b s c u r e .
IDENTIFYING THE CALIFORNIA ANCHOVY. The anchovy h a s a p e c u
l i a r sn o u t t h a t d i s t i n g u i s h e s i t from a l l o t h e r t e l e o s t f i s h .
T h is ’’a n ch o v y ” sn o u t i s s h a r k - l i k e , on th e o r d e r o f a cone,
and th e mouth i s v e n t r a l l y p la c e d (Cunningham and C alderw ood
1 8 6 9 -9 1 , H ild e b ra n d and Cable 1930, K untz 1 9 3 3 ).
B a r n h a r t (1930) g i v e s th e f o l lo w in g c h a r a c t e r i s t i c s s p e c i f i c
t o mordax m ordax, G i r a r d , th e o f f i c i a l C a l i f o r n i a anchovy:
(1) The m id d le o f t h e h e a d i s some d i s t a n c e b e h in d th e e y e .
( 2 ) The h e ad i s l o n g e r th a n th e d e p th o f th e bo dy.
( 3 ) The b a se o f th e a n a l f i n i s s h o r t e r th a n th e h e a d .
( 4 ) T here may be two ro d s a t th e c e n t e r o f th e t a i l .
( 5 ) There may be a f a i n t l a t e r a l c o l o r - l i n e . Jo rd a n and
5
Evermann (1858-1900) and B a r n h a rt (1930) s t a t e t h a t th e
C a l i f o r n i a anchovy does n o t have a l a t e r a l l i n e , h u t t h a t
i n s t e a d may have a l a t e r a l c o l o r - l i n e .
(6) The f i s h grows to seven i n c h e s .
( 7 ) The h a c k i s h l u e - g r a y . The s i d e s and th e b e l l y a re
w h i t e .
OTHER PACIFIC COAST ANCHOVIES. A cco rd in g to B a r n h a r t (1930)
th e o f f i c i a l C a l i f o r n i a anchovy (m ordax m ordax, G i r a r d , des
c r i b e d a b o v e , may be d i s t i n g u i s h e d from a l l o t h e r C a l i f o r n i a o r
P a c i f i c c o a s t t y p e s by i t s b lu e c o l o r . The o t h e r s p e c i e s o f
C a l i f o r n i a anchovy a re g r e e n i s h .
The g ree n a n c h o v ie s have a d i s t i n c t w h ite s i d e - b a n d . S c i
t u l a , F o w ler, h a s a w h ite s t r i p e down th e c e n t e r o f i t s b a c k .
A c co rd in g to B a r n h a r t ’ s (1930) m ea su re m e n ts, th e g ree n a n ch o
v i e s do n o t exceed f i v e in c h e s in l e n g t h .
SPERMARIA AND 0VARIA. Cunningham and Calderw ood (1889-91)
r e p o r t e d t h a t th e e m b ry o lo g ic a l developm ent o f th e u r o - g e n i t a l
s t r u c t u r e s o f a l l c lu p e o id f i s h , f o r example a n c h o v ie s , h e r r
in g , salm on, and t r o u t a r e i d e n t i c a l , th o s e s t r u c t u r e s in th e
male and fem ale b e in g h o m olo gues, w ith s l i g h t v a r i a t i o n s , o f
each o t h e r . These a u th o r s found t h a t th e s p e rm a ria and o v a r i a
o f a n c h o v ie s , t h a t i s , th e developm ent o f th e germ c e l l s w i t h i n
th e g o n a d s, a r e , w ith s l i g h t v a r i a t i o n s , d u p l i c a t e s . Cunningham
6
and Calderw ood (18 89 -9 1 ) p o i n te d out t h a t th e d e t a i l s o f germ
c e l l s i z e - t y p e developm ent in th e anchovy gonad was n o t w e ll
u n d e rs to o d due to th e f a c t t h a t th e n u c le u s o f th e germ c e l l in
t h i s p a r t i c u l a r f i s h i s d i f f u s e d in th e cy to p o lasm and v i s i b l e
o n ly a s " n u c l e a r d u s t o r a s a n u c l e a r c lo u d " . H ild e b r a n d and
C able (1930) and Kuntz (1933) c o n c u r.
GERM CELL SIZE-TYPES IN TELEOST FISH. The t h r e e s i z e - t y p e s
o f germ c e l l s , w hich r e p r e s e n t grow th s t a g e s , u s u a l l y found in
th e r i p e o r spaw ning gonads o f t e l e o s t f i s h , w hich i n c l u d e s th e
anchovy, a re a s f o l lo w s : ( Cunningham and Calderw ood 1889-91,
H ild e b r a n d and C able 1930, and K untz 1 9 3 3 ).
(1) The rainüte o r s m a l l - s i z e germ c e l l s .
( 2 ) The m id d le s i z e germ c e l l s .
( 3 ) The l a r g e s i z e germ c e l l s .
These s i z e - t y p e s r e p r e s e n t imm ature germ c e l l s as d i s t i n g
u i s h e d from th e m atu re germ c e l l s o r sp e rm ataz o a and eggs, th e
l a t t e r b e in g to o l a r g e to be c o n fu se d w ith th e fo rm e r.
The anchovy egg h a s a c h e c k e re d a p p e a ra n c e due to th e
se g m e n ta tio n o f th e y o lk g l o b u l e s . So f a r a s i s known, bona
f i d e anchovy m i l t o r sp e rm a ta z o a h a s n e v e r been i d e n t i f i e d
( Cunningham and Calderwood 1 8 8 9 -9 1 , H ild e b r a n d and C able 1930,
K untz 1 9 3 3 ) .
These a u t h o r s r e p o r t t h a t so f a r a s i s known, a l l male germ
c è l l s lo o k l i k e imm ature "fem a le o v a ".
7
HISTOLOGY OF THE ANCHOVY GONADS. From Cunningham and
Calderw ood (18 89-91 ) th e f o llo w in g d a ta h a s been g a th e r e d in
r e g a r d to t h e h i s t o l o g y o f th e anchovy g o n a d s, and r e c a p i t u
l a t e d e i t h e r in te rm s o f g e n e r a l i t i e s o r w ith d e f i n i t e c o r r o b
o r a t i v e d i s c u s s i o n s by Hubbs (1 9 2 5 ), H ild e b r a n d and C able (1 9 3 0 ),
K untz (1 9 3 3 ), and B o lin (1 9 3 6 ).
(1) The anchovy gonad i s c o v ered by a c o n n e c tiv e t i s s u e
c a p s u l e . A p p a re n tly t h e r e i s no s e r o s a .
(2) The anchovy gonad h a s no t r u e s e m e n ife ro u s t u b u l e s .
T u rn e r (1919) o b s e r v e s t h a t t h i s i s th e c o n d i t i o n in r e g a r d to
f r e s h - w a t e r p e r c h .
( 3 ) The anchovy o v a ry i s a t u b u l a r o v a ry , t h a t i s , i t i s
a s a c , and t h i s t r u i s m h o l d s f o r th e t e s t i s ( P a r k e r and H as-
w e ll 1 9 4 0 ).
( 4 ) The anchovy tu b u la r - g o n a d c o n ta in s l o b u l e s pro d u ced by
th e c o n n e c tiv e t i s s u e c a p s u l e , and s e p a r a t e d from each o t h e r
by c o n n e c tiv e t i s s u e s e p t a ( P o t t e r 1938, P a r k e r and H asw ell
1 9 4 0 ). The gonad l o b u l e s a re p yram idal in sh a p e , th e b ro a d
b a s e s b e in g fu s e d w ith th e c o n n e c tiv e t i s s u e o f th e p e r i p h e r y
o f th e o rg a n , th e p o i n te d a p i c e s b e in g c l u s t e r e d ab ou t th e
c o n n e c tiv e t i s s u e core from w hich th e y r a d i a t e .
( 5 ) The g e n i t a l d uct o f th e gonad i s a c o n t i n u a t i o n o f th e
gonad c a p s u l e , and opens to th e e x t e r i o r by way o f th e u r o
g e n i t a l s i n u s , t h a t o r i f i c e b e in g a c lo a c a i n t o w hich opens
th e sm a ll i n t e s t i n e . Each eonad h a s a g e n i t a l d u c t t h a t
8
f u s e s w ith i t s f e l l o w on th e o p p o s i t e s id e e i t h e r c lo s e to
th e c a u d a l t i p of th e gonad o r a t th e u r o - g e n i t a l s i n u s .
P a r k e r and H asw ell (1940) o b se rv e t h a t i t i s i n c o r r e c t to c a l l
th e g e n i t a l d u c ts o f f i s h w ith t r u e o v i d u c t s , such a s th e
anchovy, v a s a d e f e r e n t i a , th e p r e f e r a b l e term b e in g g e n i t a l
d u c t o r o v i d u c t .
(6) T e l e o s t f i s h w ith t r u e o v i d u c t s , a s in th e anchovy, do
n o t have v a s a e f è r e n t i a . The i n t e r t r a b e c u l a r sp a c e s s e rv e a s
th e c h a n n e ls by way o f w hich th e germ c e l l s may e sc a p e i n to
th e g e n i t a l d u c t s . P a r k e r and H asw ell (1940) l i s t L e p i d o s t e u s ,
o r th e g a r - p i k e , a s h a v in g a gonad t h a t i s a h o llo w sac b u t
t h a t , in h a v in g v a s a e f f e r e n t i a , i s th e e x c e p tio n to th e r u l e
t h a t t e l e o s t s w ith t u b u l a r gonads do not have t h e s e t u b u l e s .
T h is was m en tio n ed b e f o r e , b u t i s r e p e a t e d f o r e m p h a sis.
(7) The gonad o f th e a d u l t anchovy does not have a m esorchium ,
t h a t i s , a m e s e n te ry s u p p o r t i n g th e t e s t i s , o r a m esovariura,
t h a t i s , a m e s e n te ry s u p p o r tin g th e o v a ry .
ASYmETRY OP ANCHOVY GONADS. K endal (1921) o b s e rv e d t h a t in
c lu p e o id f i s h one gonad i s u s u a l l y s h o r t e r th a n th e o t h e r in
th e same i n d i v i d u a l . He found t h a t th e r i g h t gonad o f salmon
i s c o n s i s t e n t l y s h o r t e r th a n i t s f e l l o w on th e l e f t .
The p r e s e n t w r i t e r d id no t f i n d s t a t i s t i c s in th e a v a i l a b l e
l i t e r a t u r e r e g a r d i n g th e l e n g t h o f each o f a p a i r of anchovy
gonads to th e o t h e r . S am p ling s o f th e f i s h showed t h a t th e
r i g h t gonad i s u s u a l l y s h o r t e r th a n th e l e f t .
9
THE ANCHOVY KIDNEY. The l i t e r a t u r e on th e anchovy k id n e y s
was s c a r c e . Cunningham and C alderw ood (1 8 8 9 -9 1 ), P o t t e r (1 9 3 8 ),
P a r k e r and H a sw ell ( 1 9 4 0 ), and H e g n e r(l9 4 2 ) s t a t e t h a t th e body
k id n e y o r m eson ep hros i s a m a ssiv e p a i r e d organ ru n n in g a lo n g
th e backbone b e n e a th th e p e rito n e u m from th e extrem e c au d a l
end o f th e body c a v i t y to a p o s i t i o n o p p o s ite th e o p e r c l e .
The p r o n e p h ro s o r h e ad k id n e y o f t e l e o s t s d e g e n e r a t e s in th e
a d u l t f i s h i n t o lym phoid t i s s u e t h a t c o n ti n u e s i n to th e head
c a v i t y to a p o s i t i o n above th e h e a r t .
P a r k e r and H asw ell (1940) s t a t e t h a t in th e a d u l t t e l e o s t th e
m eso n ep h ric d u c t i s th e e x c r e t o r y d u c t b u t t h a t in some s p e c i e s
th e p r o n e p h r ic d u ct may p e r s i s t . A cco rd in g t o t h e s e a u th o r s
th e s e c r e t o r y d u c ts o r u r i n i f e r o u s t u b u l e s have b l i n d ends t h a t
u s u a l l y a re d i l a t e d i n to a Bowman’ s c a p s u le o r p r i m i t i v e glom e
r u l u s . These t u b u l e s a r e h e l d t o g e t h e r by c o n n e c tiv e t i s s u e .
Z o eth o u t end T u t t l e (1943) s t a t e t h a t some f i s h , th e s p e c i e s
i s n o t named, do n o t have a Bowman’ s c a p s u l e .
THE TELEOST LIVER. P o t t e r (1 9 3 8 ), P a r k e r and H asw ell (1 9 4 0 ),
and H egner (1942) d e s c r i b e th e t e l e o s t l i v e r a s a d i g e s t i v e
g la n d t h a t s e c r e t e s b i l e , and th e h i s t o l o g y o f which i s t y p i c a l
o f t h a t org an in o t h e r a n im a l s . The l i v e r t i s s u e o f f i s h i s
composed o f h e p a t i c c e l l s , th e b i l e c a n a l i c u l i a r e form ed in
th e u s u a l way, and p r i m i t i v e l o b u l e s a re i n d i c a t e d by th e
p re s e n c e o f c e n t r a l v e i n s .
10
THE GALL BLADDER IN FISH. The l i t e r a t u r e on th e g a l l b la d d e r
in t e l e o s t f i s h i s n o t c o p io u s , and th e p r e s e n t w r i t e r c o u ld
n o t f i n d t h a t org an l i s t e d a s b e in g p r e s e n t in th e anchovy.
P a r k e r and H asw ell (1940) and H egner (1942) d e s c r i b e th e g a l l
b l a d d e r in t e l e o s t f i s h a s a b l i n d o u tp u s h in g o f th e g u t , th e
f u n c t i o n o f w hich i s to s t o r e b i l e from th e l i v e r . These
a u t h o r s l i s t th e p r e s e n c e o f a g a l l b l a d d e r in Salmo f a r i o o r
b ro o k t r o u t , a t e l e o s t t h a t i s a r e l a t i v e o f th e anchovy.
THE TELEOST SPLEEN. P o t t e r (1 9 3 8 ), P a r k e r and H asw ell (1 9 4 0 ),
and H egner (1942) s t a t e t h a t th e s p le e n i n t e l e o s t f i s h i s
more on th e o r d e r o f a haemolymph node r a t h e r th a n in th e form
o f mammalian s p l e e n i c t i s s u e t h a t shows d i s t i n c t n o d u le s o r
m a lp ig h ia n b o d i e s . These a u t h o r s s t a t e t h a t in some t e l e o s t
f i s h th e s p l e e n , a d u c t l e s s g l a n d , i s a t t a c h e d by a t h r e a d o f
m e s e n te ry to th e d o r s a l w a ll o f th e d i g e s t i v e c a n a l n e a r th e
a n u s , b u t t h a t in o t h e r form s i t i s a w edge-shaped body imbedded
i n th e l i v e r t i s s u e , and a t t a c h e d by a t h r e a d o f m e s e n te ry to
th e duodenum. So f a r a s i s known to th e p r e s e n t w r i t e r th e
p o s i t i o n o f th e s p le e n in th e anchovy i s not l i s t e d in th e
l i t e r a t u r e .
THE PYLORIC CAECA. R ogick and B lake (19 30 -3 1 ) l i s t th e s e
t u b e s as b l i n d o u tp u s h in g s o f th e p y l o r i c arm o f th e stom ach
th e f u n c t i o n o f w hich i s to i n c r e a s e th e s e c r e t i n g s u r f a c e o f
t h e i n t e s t i n e .
11
HISTOLOGY OF THE DIGESTIVE CANAL. So f a r a s i s known to
th e p r e s e n t w r i t e r , th e a v a i l a b l e l i t e r a t u r e does n o t o f f e r
an o u t l i n e o r a d e t a i l e d d e s c r i p t i o n o f th e t i s s u e l a y e r s o f
th e d i g e s t i v e c a n a l o f th e anchovy.
B lake (l9 30) s u b m itte d a com plete o u t l i n e o f th e h i s t o l o g y
o f th e w a l l s o f th e d i g e s t i v e c a n a l o f s e a - b a s s , th e ma.in
f e a t u r e s b e in g a s f o l l o w s :
(1) The t u n i c a s o r t i s s u e - l a y e r s o f th e d i g e s t i v e c a n a l o f
t h e s e a - b a s s c o n s i s t o f : ( l ) The m ucosa, (2) th e submucosa
w hich i s a c o n t i n u a t i o n o f th e la m in a p r o p r i a o f th e m ucosa,
( 3 ) th e t w o - l a y e r e d m u s c u l a r i s , and (4) th e s e r o s a o f a s i n g l e
l a y e r o f e p i t h e l i a l c e l l s .
( 2 ) The e p i t h e l i a l c e l l s a re colum nar and g o b l e t t y p e s .
( 3 ) The g a s t r i c g la n d s a re s t r a i g h t t u b u l e s .
( 4 ) The t w o - l a y e r e d m u s c u la r is c o n s i s t s o f a wide i n t e r n a l
m uscle l a y e r o f c i r c u l a r , s t r i a t e d , smooth m uscle f i b e r s ; and
a n a rro w e x t e r n a l m uscle l a y e r o f l o n g i t u d i n a l , u s u a l l y non-
s t r i a t e d , smooth m uscle f i b e r s .
B o lto n ( 1 9 3 3 ) found t h a t in k in g salmon and s p e c k le d t r o u t ,
form s t h a t a re r e l a t i v e s o f th e anchovy, th e e n t i r e l e n g t h o f
th e d i g e s t i v e c a n a l was m ark e d ly g r a n u l a r in a p p e a ra n c e due to
th e p r e s e n c e o f an enormous q u a n t i t y o f ly m p h o cy tes w hich he
c a l l e d b a s o p h i l e s o r m ast c e l l s .
12
I I I . MATERIALS AND METHODS
SECURING AND FIXING MATERIAL. The m a t e r i a l f o r t h i s i n
v e s t i g a t i o n was s e c u re d from th e s e a - w a t e r t a n k s o f th e S a n ta
Monica B a it and T as^e Company t h a t , in t u r n , s e c u r e d th e l i v e
a n c h o v ie s from th e C oast Guard b o a t s a t S a n ta M onica, C a l i f o r n i a .
The l i v e f i s h would be l i f t e d from th e ta n k s o f s e a - w a t e r ,
c a re b e in g ta k e n to s e l e c t o n ly i n d i v i d u a l s t h a t a p p e a re d h e a l
t h y , and l e f t in a i r u n t i l th e y c e a se d to r e a c t to s t i m u l i .
The abd om inal c a v i t y o f each i n d i v i d u a l specim en o f A nchovia
m ordax m ordax, G i r a r d , was th e n opened, a f t e r w hich th e f i s h
was p la c e d f o r f i x i n g in B o u in ’ s s o l u t i o n . Four o r f i v e dozen
a n c h o v ie s would be s e c u re d in a s i n g l e sa m p lin g a s t h i s q u a n t i t y
c o u ld be h e l d c o n v e n i e n t l y , w ith o u t c a u s in g a d i s t o r t i o n o f th e
b o d i e s o f th e f i s h , in a g a l l o n - s i z e , g l a s s , s a l a d j a r .
The g r o s s s t r u c t u r e s o f th e specim ens were c a r e f u l l y d i s s e c
t e d a t th e l a b o r a t o r y o f th e zoology d e p a rtm e n t of th e U n iv e r
s i t y o f S o u th e rn C a l i f o r n i a , th e v a r i o u s o rg a n s o f th e v i s c e r a
b e in g e x t i r p a t e d and p l a c e d in r e s p e c t i v e v i a l s c o n t a i n i n g
e i g h t y p e r c e n t a l c o h o l .
PREPARATION QE THE TISSUE. When th e v a r i o u s t i s s u e s were
re a d y to be imbedded in wax, a m ix tu re o f one p a r t b a y b e r ry
wax to n in e p a r t s p a r a f f i n , th e y were f i r s t d e h y d r a te d . E th y l
a lc o h o l was th e d e h y d r a t in g a g e n t u s e d , th e t i s s u e # b e in g ta k e n
13
th ro u g h th e f o llo w in g p e r c e n t a g e s in th e o r d e r i n d i c a t e d in
th e o u t l i n e h e lo w .
1 . 8 0 ^ a lc o h o l 2 h r s .
2 . 90 ^ a l c o h o l 2 h r s .
3. 95^ a l c o h o l 2 h r s .
4 . A b s o lu te a lc o h o l 2 h r s .
I t was found t h a t th e b e s t r e s u l t s were o b ta in e d when th e
m a t e r i a l was l e f t in a b s o l u t e a lc o h o l o v er n i g h t . To c l e a r
them o f th e d e h y d r a tin g a g e n t , th e t i s s u e s were ta k e n z y len e
a s f o l l o w s .
1. Z ylene 2 h r s .
2. Z ylene 2 h r s .
3. The m a t e r i a l was th e n p la c e d in a b e a k e r c o n ta in i n g a
one to n in e b a y b e r r y - p a r a f f i n m ix tu re f o r i n f i l t r a t i o n and
l e f t in th e oven two d a y s, th e te m p e r a tu r e b e in g k e p t a t 70®C.
IMBEDDING IN W AX MOLDS. A f t e r i n f i l t r a t i o n w ith th e one to
n in e b a y b e r r y - p a r a f f i n m ix tu re a s d e s c r i b e d ab o v e, th e t i s s u e s
were removed from th e oven and p la c e d in sm a ll p a p e r b o a t s t h a t
were f i l l e d w ith th e s to c k b a y b e r r y - p a r a f f i n m ix tu r e t h a t had
been u se d f o r i n f i l t r a t i o n . The one to n in e b a y b e r r y - p a r a f f i n
m ix tu re in t h e b o a ts c o n t a i n i n g th e t i s s u e s was a llo w e d to
h a rd e n i n t o " b lo c k s " each o f w hich was l a b e l e d and num bered.
C e r t a i n o rg a n s were cu t i n t o segm ents f o r co n v en ien c e in h a n d l
in g b e f o r e b e in g p la c e d in th e m old.
14
MOUNTING THE SECTIONS. The p ro c e e d u re f o r m ounting th e
s e c t i o n s a s th e y came from th e m icrotom e in th e f car m o f a "wax
rib b o n " was a s f o l l o w s :
A g l a s s s l i d e , a f t e r b e in g f i r s t c le a n e d w ith a one to one
m ix tu re o f a lc o h o l and z y le n e , was c o v e re d w ith a f i l m o f
M ayer’ s Albumin F i x a t i v e , and th e n f lo o d e d w ith c o o l, d i s t i l l e d
w a te r a f t e r w hich th e s e c t i o n s were p la c e d upon i t in such a
m anner t h a t th e y re a d from l e f t to r i g h t . The p r e p a r e d s e c t i o n s
were th e n p la c e d upon g l a s s p l a t e s t h a t in t u r n c o v ered a pan
o f warm w a te r t h a t was k e p t a t 48®C. As soon a s th e s e m ounted
s e c t i o n s were d ry th e y were p la c e d in an a e r a t e d d raw er to
" c u re " f o r s e v e r a l days b e f o r e b e in g s t a i n e d .
STAINING. Most o f th e s e c t i o n s were s t a i n e d w ith D e l a f i e l d ’ s
h e m a to x y l i n - e o s in in th e u s u a l way. I t was d is c o v e r e d t h a t
anchovy t i s s u e s were too d e l i c a t e to be run th ro u g h th e r e s
p e c t i v e s o l u t i o n s in th e s t a i n i n g t r a y w ith o u t th e u s e o f c e l -
l o i d i n to h o l d them to th e s l i d e s so t h a t th e y would no t wash
o f f .
CONTROLS. The m ic r o te c h n iq u e c l a s s o f 1946 s u p p lie d th e
p r e s e n t w r i t e r w ith s t a i n e d s e c t i o n s o f th e o rg a n s o f th e r a t
and c a t a s mammalian s t a n d a r d s o f h i s t o l o g i c a l s t r u c t u r e w ith
w hich to compare s e c t i o n s o f anchovy t i s s u e .
15
G ross d i s s e c t i o n r e v e a l e d t h a t i t was im p o s s ib le to i s o l a t e
such o rg a n s in s i t u a s th e l i v e r , s p le e n , and d i g e s t i v e c a n a l
from each o t h e r by means o f a p r o b e , n e e d l e , o r s c a p e l w ith o u t
m u t i l a t i n g them . The l i v e r c o v ered and e n fo ld e d th e two arms
o f th e stom ach, th e s p le e n was imbedded in th e l i v e r t i s s u e ,
and th e p y l o r i c c ae ca were h e l d i n e x t r i c a b l y in th e t i s s u e -
m asses o f th e l i v e r and s p l e e n .
As a c o n t r o l , th e d i g e s t i v e c a n a l was c a r e f u l l y l i f t e d from
i t s r e l a t e d o rg a n s , t h e l i v e r , s p l e e n ,a n d p y l o r i c c a e c a ; and
s e r i a l s e c t i o n s were made o f th e e so p h a g u s, th e c a r d i a c and py
l o r i c arm s o f th e stom ach, t h e caecum o f th e stom ach, and th e
sm a ll i n t e s t i n e . P y l o r i c c a e c a would c l i n g to th e p y l o r i c
arm o f th e stom ach w henever t h a t r e g io n was e x t i r p a t e d f o r th e
p u rp o se o f s e r v i n g a s a c o n t r o l ; and h en ce t h e i r e x a c t r e l a
t i o n s h i p to t h e i r p a r e n t t u b e , th e p y l o r i c arm, c o u ld be d e t e r
m ined b e f o r e s e r i a l s e c t i o n s were made o f th e u n d i s t u r b e d mass
o f th e stom ach and i t s r e l a t e d o rg a n s .
In making s e r i a l s e c t i o n s o f th e whole m ass o f th e stom ach
and i t s r e l a t e d o rg a n s such a s th e l i v e r , s p l e e n , and p y l o r i c
c a e c a , (and th e p r e s e n t w r i t e r hoped to l o c a t e th e g a l l b l a d d e r
i f such a s t r u c t u r e were p r e s e n t ) i t was found t h a t th e t i s s u e s
were so d e l i c a t e t h a t when m ounted th e y were d e ta c h e d from
t h e i r p r o p e r p l a c e - l o c a t io n s and o c c u p ie d abnorm al p o s i t i o n s .
W itho ut h a v in g f i r s t e s t a b l i s h e d a c o n t r o l a s d e s c r i b e d a b o v e ,
d e ta c h e d o rg a n s in a s e c t i o n c o u ld n o t have been i d e n t i f i e d .
16
IV. HISTOLOGY
THE GONADS. T here i s a p a i r o f gonads in each f i s h , th e
male and fem ale t y p e s b e in g s i m i l a r in a p p e a r a n c e . The gonads
a r e w h ite o r c re a m -c o lo r e d in v i v o , f l a t t e n e d c y l i n d e r s in sh a p e ,
and a re f r e q u e n t l y a s lo n g a s th e body c a v i t y . The m ale gonads
in th e a d u l t a r e f r e q u e n t l y l o n g e r th a n th o s e in a fem ale f i s h
o f th e same s k e l e t a l l e n g t h . The gonads a re p la c e d on edge in
t h e abdom inal c a v i t y , each o rg an b e in g convex on th e l a t e r a l o r
b o d v -w a ll s id e and concave on th e m e d ia l s i d e , th e d o r s a l edge
ru n n in g p a r a l l e l t o th e backbone and th e v e n t r a l edge ru n n in g
p a r a l l e l to th e m i d - v e n t r a l l i n e o f th e b e l l y . A lth o u g h f i s h
gonads a re s a i d to c o l l a p s e a f t e r spaw ning, th e p r e s e n t w r i t e r
found t h a t due t o th e f a c t t h a t th e c o n n e c tiv e t i s s u e l o b u l e s
o f th e t e s t i s do n o t a p p r e c i a b l y d e g e n e ra te a t t h a t t im e , and
due to th e t h i c k n e s s o f th e f i b r o u s , c o n n e c tiv e t i s s u e c a p s u le
o f th e o v a ry , t e n d i n g to m a in ta in i t s shape a s th e o v a ria n
l o b u l e s d e g e n e r a t e , anchovy g onads do n o t s h r i n k a p p r e c i a b l y
a t spaw ning tim e .
The m ale and fem ale gonads may be d i s t i n g u i s h e d from each
o t h e r in th e f o l lo w in g m anner: I n g e n e r a l , when th e f i s h t e s t i s
i s b ro k e n open w ith th e f i n g e r s i t i s more e l l i p t i c a l in c r o s s -
s e c t i o n th a n i s th e o v a ry w hich i s round when s i m i l a r l y exam ined,
a r u l e g iv e n to t h i s w r i t e r by th e f i e l d - w o r k e r s , and i t a p p l i e s
to th e anchov y. In vivo th e anchovy t e s t i s shows l e s s
17
b u l g in g o f th e l o b u l e s th a n i s th e case in th e o v a ry where
t h e r e i s u s u a l l y much l o b u l a t i o n on th e m e d ia l s id e and where
t h e l o b u l e s b u lg e t o such a d e g re e t h a t th e e l e v a t e d m edian
c o re o f th e organ i s o b sc u re d from im m ediate v iew .
The t e s t i s i s s o l i d t o t h e to u c h , and i s e a s i l y e x t i r p a t e d
and h a n d le d w ith th e f o r c e p s , w hereas t h e o v a ry i s spongy,
" r o t t e n " in a p p e a ra n c e , and so d e l i c a t e t h a t i t may be e x t i r
p a te d and h a n d le d o n ly w ith g r e a t d i f f i c u l t y .
THE TESTIS. The f o l lo w in g i s a summary o f th e s t r u c t u r e s
t h a t were seen i n a c r o s s - s e c t i o n o f anchovy t e s t i s .
(1) T here was a c o n n e c tiv e t i s s u e c a p s u l e . The spawning
t e s t i s o f t e n a p p e a re d to have no c a p s u l e , b u t t h i s a p p e a ra n c e
was c r e a t e d by th e f a c t t h a t t h a t s t r u c t u r e was f r e q u e n t l y v e ry
n a rro w , i n f i l t r a t e d w i t h m i n u t e - s i z e germ c e l l s , and th e t r i a n g
u l a r l o b u l e s , o f t e n crowded c l o s e l y t o g e t h e r , were c o n t i n u i t i e s
o f c a p s u l a r t i s s u e . T here was a p p a r e n t l y no c a p s u l a r s e r o s a ,
and to t h i s w r i t e r ’ s k n o w le d g e , none h a s been l i s t e d by th e
l i t e r a t u r e .
( 2 ) The t e s t i c u l a r l o b u l e s c o n s i s t e d o f f i b r o u s , c o n n e c tiv e
t i s s u e m a s s e s , and were p y ra m id a l in sh a p e, t h e i r b a s e s a t th e
p e r i p h e r y o f th e org an and t h e i r a p i c e s r a d i a l l y d is p o s e d a b o u t
t h e m edian c o r e . The l o b u l e s were m ark e d ly wavy in o u t l i n e .
18
(3) The s e p t a betw een th e t e s t i c u l a r l o b u l e s were n o t
a lw ay s p r e s e n t a s i n d i v i d u a l s t r u c t u r e s . More f r e q u e n t l y th e y
c lu n g t i g h t l y to th e m a rg in s o f th e l o b u l e s , th e o n ly b o un dary
betw een th e l o b u l e s b e in g an i n t e r t r a b e c u l a r s p a c e - c h a n n e l .
( 4 ) As r e p o r t e d in th e l i t e r a t u r e a s a t r u i s m o f a lu p e o id
f i s h , th e anchovy t e s t i s d id n o t have v a s a e f f e r e n t i a . I n s t e a d
t h e r e were s p a c e - c h a n n e ls betw een th e l o b u l e s fo rm in g t o r t u o u s
p a ssa g e -w a y s to th e m edian c o r e .
( 5 ) The m edian core o f th e t e s t i s was a t h i c k e n i n g o f th e
f i b r o u s , c o n n e c tiv e t i s s u e c a p s u le a lo n g th e m id d le l i n e o f
th e m edian s u r f a c e . A c r o s s - s e c t i o n o f t e s t i s showed no
c o n tin u o u s lumen w i t h in th e m edian c o r e . I t was f i l l e d w ith
i r r e g u l a r - s h a p e d s m a ll, i n t e r t r a b e c u l a r s p a c e s t h a t c o a le s c e d
and o f f e r e d a t o r t u o u s c h an n e l th r o u g h o u t th e e n t i r e l e n g t h
o f th e c o r e . The m edian c o re s e rv e d a s a p r i m i t i v e r e c e i v i n g
tu b e t h a t c o n d u c te d th e germ c e l l s to th e g e n i t a l d u c t .
(6) T hree s i z e - t y p e s o f germ c e l l s were found in th e r i p e
o r spaw ning t e s t i s , nam ely, th e s m a ll, th e m id d le , and th e
l a r g e . The m ale germ c e l l s were round and s i m i l a r in a p p e a r
ance t o th e ova in th e o v a r y . The p r e s e n t w r i t e r was n o t a b le
to d i s c o v e r a male germ c e l l t h a t c o u ld be i d e n t i f i e d a s a
m atu re sp e rm a ta z o a o r m i l t . Male germ c e l l s lo o k e d l i k e o va.
( 7 ) Germ c e l l s i z e ty p e s te n d e d to c l u s t e r t o g e t h e r in
r e s p e c t i v e p l a c e - l o c a t i o n s , b u t a s id e from a te n d e n c y to w ard i t .
19
t h e r e was a p p a r e n t l y no n e s t i n g b e h a v i o r .
1. Germ c e l l s i z e - t y p e s te n d e d to l a y in s t r a t a ,
a p a r t i c u l a r s i z e - t y p e r e p r e s e n t i n g a p a r t i c u l a r s t r a tu m .
The s t r a t a te n d e d to run c r o s s - w i s e th e t e s t i c u l a r l o h u l e , t h a t
i s , from th e l a t e r a l s id e to w ard th e m e d ia l s i d e , en d in g a t
th e m e d ia l c o r e . T h is gave th e c r o s s - s e c t i o n o f t e s t i s a handed
a p p e a ra n c e , th e sequence b e in g an i n n e r re g io n w i t h in a l o b u le
o f s m a l l - s i z e germ c e l l s bounded by a m arg in of m i d d l e - s i z e
germ c e l l s . A p p a r e n tly an e n t i r e l o b u le c o u ld c o n s i s t o f l a r g e -
s i z e male germ c e l l s . Immature male germ c e l l s lo o k ed l i k e ova.
2. I t was o b se rv e d t h a t th e s t r a t i f i e d p a t t e r n o f
germ c e l l s i z e - t y p e s in a r e s p e c t i v e s tr a tu m d e s c r i b e d above
o f t e n showed a v a r i a t i o n , b e in g so a r r a n g e d t h a t th e c r o s s -
s e c t i o n o f t e s t i s had a c i r c l e - w i t h i n - c i r c l e a p p e a ra n c e , th e
c e n t r a l r e g io n c o n t a i n i n g s m a l l - s i z e germ c e l l s .
An a rra n g e m e n t o f r e g i o n s in th e o r d e r l y sequence o f s m a ll,
m id d le , and l a r g e s i z e - t y p e s was som etim es s e e n , b u t a s many
tim e s t h e r e was a p p a r e n t l y a random d i s t r i b u t i o n in w hich l a r g e
germ c e l l s were b e s i d e sm a ll o n e s.
(8) The spaw ning t e s t i s showed th e p r e s e n c e o f male f o l l i c l e s .
The empty t e s t i c u l a r f o l l i c l e was s i m i l a r in a p p e a ra n c e to t h a t
o f th e o v a ry , l o o k in g much l i k e p a tc h e s o f c r o c h e t work in th e
l o b u l e s .
(9) The m ale g e n i t a l d u c t was a c o n t i n u a t i o n o f th e f i b r o u s .
20
FIGURE I
B .........................................................Basement membrane
C.........................................................C u t ic u l a
C E .....................................................Columnar e p i t h e l i a l c e l l s
GT......................................................C o n n e c tiv e t i s s u e w a ll o f th e d u ct
L. . ........................................... . .Lamina p r o p r i a
LD......................................................Lumen o f th e d u c t
L S ......................................................Lymph-space c h an n e l
N o te : I t i s th e m orphology o f th e t i s s u e - l a y e r s and th e
shape o f th e la m in a p r o p r i a in th e form o f f o l d s t h a t i s
im p o r ta n t in i d e n t i f y i n g d u c ts in t h i s f i s h .
21
F i e . 1
C r o s s - s e c t i o n o f a p o r t i o n o f th e w a ll o f th e l e f t male
g e n i t a l d u c t showing t h r e e f o l d s o f th e la m in a p r o p r i a . On
th e l e f t o f th e f i g u r e i s th e c o n n e c tiv e t i s s u e o u t e r - l a y e r
of th e d u c t; on th e r i g h t i s th e lum en. The m a g n i f i c a t i o n i s
10X44X 1 .8 8 .
22
c o n n e c tiv e t i s s u e c a p s u l e . The m edian c o re , a t h i c k e n i n g of
th e c a p s u le i n to a r i d g e t h a t ra n a lo n g th e m id d le l i n e of
th e m e d ia l s u r f a c e o f th e t e s t i s , and f i l l e d w ith s p a c e s , le a d
d i r e c t l y i n t o th e g e n i t a l d u c t -
A com plete c r o s s - s e c t i o n o f th e m ale g e n i t a l d u c t, t h a t i s ,
th e com plete c i r c l e o f th e w a l l s , was n o t s e c u r e d . C r o s s - s e c t
io n s o f one w a ll were o b t a i n e d , how ever, and a summary o f th e
h i s t o l o g y i t r e p r e s e n t e d , a s a f ra g m e n t, i s a s f o l lo w s :
1. There was a f i b r o u s , c o n n e c tiv e t i s s u e c a p s u l e .
2 . T here was a la m in a p r o p r i a o f c o n n e c tiv e t i s
sue t h a t was throw n up i n t o th e lumen in w id e, low f o l d s ,
th e f r e e ends o f w hich w ere s l i g h t l y l o b u l a t e d .
3. The tu b e was l i n e d w ith low , co lum n ar, e p i
t h e l i a l c e l l s t h a t were c o v e re d w ith a d e l i c a t e c u t i c u l a and
t h a t r e s t e d upon a basem ent membrane.
S e r i a l s e c t i o n s were not s e c u re d o f th e g e n i t a l duct owing
to th e f r a g i l i t y o f t h a t o rg a n . The p r e s e n t w r i t e r su b m its
th e above d a ta on t h i s o rg an w ith th e wide r e s e r v a t i o n t h a t
i t have s t a t i s t i c a l b u t no a c t u a l s i g n i f i c a n c e , and t h a t , in
th e a b sen ce o f s e r i a l s e c t i o n s f o r v e r i f i c a t i o n , no c o n c lu s io n
can be re a c h e d a s to th e r e a l s t r u c t u r e o f th e male g e n i t a l
d u c t .
23
FIGURE 2
G.............................................C a p su le o r o u t s i d e l i m i t o f ovary
D.................................. . . . .D e b ris
L .................................................L obule in a s t a t e o f d e g e n e ra tio n showing
empty f o l l i c l e s
MO........................................... Mass o f ova.
L arge s i z e ova
M iddle s i z e ova
S . ........................................... Space
T ..............................................T ra b e c u la e o r s e p ta in a, s t a t e of d e g e n e r
a t i o n
24
%
g
X
m
Fig. 2
C r o s s - s e c t i o n o f th e m i d - s a g i t a l r e g io n o f th e l e f t o v a ry .
On th e ex trem e l e f t i s shown th e l a t e r a l s i d e . The m a g n i f i
c a t i o n i s 5X10X . 188.
25
THE OVARY The f o llo w in g i s a summary o f th e s t r u c t u r e s
t h a t were seen in a c r o s s - s e c t i o n o f spaw ning o v a ry :
(1) The f i b r o u s , c o n n e c tiv e t i s s u e c a p s u le o f th e o v ary
was w id e r, and more e a s i l y seen th a n t e s t i c u l a r c a p s u l e .
C l u s t e r s o f sm all s i z e ova were f r e q u e n t l y seen in th e o v a ria n
c a p s u l e . The o v a - c l u s t e r te n d e d to be more in th e form o f a
n e s t th a n was th e c a se in r e g a r d to male germ c e l l s in t e s t i
c u l a r c a p s u l e .
( 2 ) The s t r u c t u r e o f o v a r ia n l o b u l e s was e s s e n t i a l l y th e
same a s t h a t o f t e s t i c u l a r l o b u l e s . The o v a r ia n l o b u le was
m a rk e d ly s t r a i g h t in o u t l i n e , and narrow in c o n t r a s t to th e
w ide, wavy t e s t i c u l a r l o b u l e . The o v a r ia n l o b u le d id not have
th e banded o r s t r a t i f i e d a p p e a ra n c e o f t e s t i c u l a r l o b u l e , b u t
i n s t e a d c o n s i s t e d o f a mass o f ova showing g ro w th -d e g re e s i z e s
w ith much s i z e - t y p e o v e r l a p p i n g .
D i s t i n c t i v e o f th e spaw ning ov ary was th e marked d e g e n e r a tio n
o f th e l o b u l e s , o n ly fra g m e n ts b e in g p r e s e n t ; th e h u g e , o v a ria n
sp a c e s l e f t by t i s s u e d e g e n e r a t i o n ; and th e d e b r i s f i l l i n g
th e s p a c e s .
D i s t i n c t i v e o f th e o v a ry were th e g l o b u l a r and p o l y h e d r a l
shaped c l u s t e r s o f m id d le and l a r g e s i z e - t y p e ova in th e
i n t e r t r a b e c u l a r s p a c e s a s w e ll a s in th e fra g m e n ts o f l o b u l e s .
In th e t e s t i s t h e r e were no g l o b u l a r o r p o ly h e d r a l- s h a p e d
m asses o f male germ c e l l s c l u s t e r e d t o g e t h e r .
26
(3) O v a ria n s e p t a , a s l i n e s t h a t were n o t i n t e g r a l m a rg in s
o f l o b u l e s , were n o t s e e n . The o v a r ia n s p a c e s were f i l l e d w ith
f ra g m e n ts o f s e p t a , b u t th e i n d i c a t i o n s were t h a t t h e s e s e p ta
were m e re ly th e rem n an ts o f th e m a rg in s of l o b u l e s . I t was
p r e v i o u s l y m en tio n ed t h a t i n t h e c r o s s - s e c t i o n o f t e s t i s ,
s e p t a t h a t were n o t i n t e g r a l p a r t s o f th e l o b u l a r m a rg in s were
f r e q u e n t l y s e e n . The o v a r ia n s e p t a were m ark ed ly s t r a i g h t in
c o n t r a s t to th e wavy c o n to u r s o f th e t e s t i c u l a r s e p t a .
( 4 ) The o v a r ia n i n t e r t r a b e c u l a r s p a c e s were on th e o r d e r o f
th o s e in th e t e s t i s , d e s c r i b e d u n d e r t h a t h e a d in g .
( 5 ) The o v a r i a n m edian c o re was s i m i l a r in s t r u c t u r e to
t h a t r i d g e in th e t e s t i c u l a r c a p s u l e , th e d e t a i l s b e in g g iv e n
u n d e r t h a t h e a d in g . The o v a r ia n i n t e r t r a b e c u l a r sp ace w i t h i n
th e m edian c o r e , how ever, was m ark e d ly o v a l in s h a p e , and te n d e d
to show th e c o n t i n u i t y o f a t u b u l a r lum en. There o f t e n were
in th e o v a ry , as was r e g u l a r l y th e case in th e t e s t i c u l a r
m edian c o r e , many s m a ll, i n t e r t r a b e c u l a r s p a c e s .
(6) Ova showed th e t h r e e germ c e l l si 7. ^ - t y pas a s was shown
by m ale germ c e l l s .
( 7 ) Ova o f a s i m i l a r s i z e - t y p e te n d e d to c l u s t e r t o g e t h e r
i n r e s p e c t i v e p l a c e - l o c a t i o n s , and showed more o f a te n d e n c y
to w a rd what m ight be term ed n e s t i n g b e h a v io r th a n was th e c ase
i n r e g a r d to male germ c e l l s . L arge s i z e - t y p e ova were h a b i t
u a l l y found in g l o b u l a r m asses in th e o v a r ia n s p a c e s , o f t e n
s u rro u n d e d by s m a ll e r s i z e - t y p e s showing grow th s t a g e s
27
(8) O varian empty f o l l i c l e s were s i m i l a r in a p p e a ra n c e to
t e s t i c u l a r , d e s c r i b e d u n d e r t h a t h e a d i n g .
( 9 ) C r o s s - s e c t i o n s o f th e fem ale g e n i t a l d u ct were n o t s e
c u re d .
THE BODY KIDNEYS. The body k id n e y s were two p a i r e d o rg a n s
t h a t in v iv o were in th e shape o f f l a t , n arrow s t r i p s o f t i s s u e
e x te n d in g from th e extrem e end o f th e body c a v i t y , and ru n n in g
a n t e r i o r l y a lo n g th e backbone and u n d e rn e a th th e m e s e n te r y ,
to a p o s i t i o n o p p o s i t e th e o p e r c l e s . The w id th and l e n g t h o f
th e s e o rg a n s were v a r i a b l e . In v iv o th e body k id n e y s were
b lu e o r a b l u e - g r e e n in c o l o r , and u s u a l l y c o u ld n ot be d i s
t i n g u i s h e d from th e b l ti e - b l a c k a p p e a r in g r e n a l ly m p h a tic o rg a n s
r u n n in g a lo n g th e back b o n e .
In c r o s s - s e c t i o n th e k id n e y s were seen to c o n s i s t o f r e n a l
t u b u l e s th e w a l l s o f which were composed of a s i n g l e l a y e r o f
c u b o id a l e p i t h e l i u m . The e p i t h e l i a l c e l l s r e s t e d on a b a s e
ment membrane and were c o v e re d by a t h i n c u t i c u l a . The c e l l s
w ere g r a n u l a r , and th e n u c l e i n e a r th e basem ent membrane.
A p p a r e n tly each t u b u le was an e x te n s i o n o f a b l i n d end. No
c i l i a o r g lo m e r u li were fo u n d . The t u b u l e s were h e l d t o g e t h e r
by a mass o f a r e o l a r c o n n e c tiv e t i s s u e . A r e n a l c a p s u l e , o t h e r
th a n a m a rg in a l t h i c k e n i n g o f th e a r e o l a r c o n n e c tiv e t i s s u e
r e t i c u l u m was not a p p a r e n t .
28
Each k id n e y o f th e p a i r was ro u g h ly t r i a n g u l a r - s h a p e d , th e
b ro a d b a se r e s t i n g a g a i n s t t h e r e n a l ly m p h a tic t i s s u e , t h e ap ex
p o i n t i n g to w a rd th e body w a l l .
There was, a c o n tin u o u s m eso n ep h ric d u c t r u n n in g th r o u g h th e
d i s t a l end o f th e c r o s s - s e c t i o n . A d i s c o n ti n u o u s p r o - n e p h r i c
d u c t was f r e q u e n t l y found in th e p ro x im a l end o f th e c r o s s -
s e c t i o n . The m e so n e p h ric d u ct was l i n e d w ith c u b o id a l e p i t h e -
lia m , b u t th e p r o n e p h r ic d u c t , when p r e s e n t i n th e a r e a , was
u n l i n e d .
THE RENAL LYMPHATIC ORGANS. The a p p e a ra n c e o f th e r e n a l
ly m p h a tic o rg a n s in v iv o was d e s c r i b e d a b o v e. In c r o s s - s e c
t i o n t h e s e were seen to be two p a i r e d o rg a n s composed of
ly m p h a tic t i s s u e l y i n g betw een th e two body k i d n e y s . Each
r e n a l ly m p h a tic o rg an was ro u g h ly w edge-shaped or b e a n -s h a p e d ,
and d id not show t r a c i n g s o f n o d u le s o r M a lp ig h ia n b o d i e s .
The spongy, re d p u lp a p p a r e n t l y c o n s i s t e d of e n d o t h e l i a l c e l l s
h e l d t o g e t h e r in a f i n e r e t i c u l u m .
The ly m p h a tic o rg a n s e x te n d e d from th e extrem e c a u d a l end
o f th e body c a v i t y to a p o s i t i o n w e ll w i t h i n th e head c a v i t y .
S e c t i o n s of th e c o n t i n u a t i o n o f th e r e n a l ly m p h a tic o rg an s
i n t o th e h e ad c a v i t y where th e y a re o f t e n c a l l e d th e d e g e n e ra
t e d h ead k id n e y s were not s e c u re d owing to t h e i r f r a g i l i t y .
29
THE LIVER» The l i v e r was so f r a g i l e , and in most c a s e s so
d e g e n e r a te d and r e p l a c e d in so l a r g e a d e g re e w ith lym phoid
t i s s u e , t h a t com plete c r o s s - s e c t i o n s c o u ld n o t he s e c u r e d .
The l i v e r in many f i s h , e s p e c i a l l y c lu p e o id c and e e l s , hccomco
so " r o t t e n " , and so i n f i l t r a t e d w ith ly m p h a tic t i s s u e , e s p e c
i a l l y d u rin g spawning tim e t h a t i t canno t he s u c c e s s f u l l y
s e c t i o n e d . A summary o f th e f i n d i n g s i s as f o llo w s :
(1) O c c a s i o n a l l y th e anchovy l i v e r was d i v id e d i n to two
l o b e s , b u t u s u a l l y t h e r e was no marked l o b u l a t i o n , th e organ
c o n s i s t i n g o f a s o l i d ,s p o n g y mass of l i v e r c e l l s e n c l o s i n g th e
arms of th e stom ach and r e l a t e d s t r u c t u r e s . In some i n d i v i d
u a l s th e l i v e r was a p a le o l i v e - c o l o r in v iv o , b u t u s u a l l y i t
was b l u e - b l a c k . One specim en o f l i v e r showed an a n t e r i o r lo b e
t h a t was e m e rald g re e n in c o l o r .
( 2 ) A c r o s s - s e c t i o n of l i v e r shov;ed t h a t t h e r e were no u n i t s
o r l o b u l e s o f l i v e r marked o f f in any v/ay such a s by a co n n ec
t i v e t i s s u e b o u n d a ry . A p p a r e n tly t h e r e were no i s l a n d s , a s in
human l i v e r , or lo n g t r a b e c u l a e , as in c a t l i v e r , o f f i b r o u s
c o n n e c tiv e t i s s u e . The o n ly c o n n e c tiv e t i s s u e d i s c e r n a b l e was
t h a t o f th e f i n e r e t i c u l u m .
( 3 ) A c r o s s - s e c t i o n o f anchovy l i v e r showed t h a t t i s s u e to
be in th e form o f w ide s h e e t s o f c e l l s . These h e p a t i c o r l i v e r
c e l l s were l a r g e and o v al o r i r r e g u l a r in s h a p e . They were
g r a n u l a r and c o n ta in e d d i s t i n c t l y s e e n , i f n o t l a r g e , n u c l e i .
30
B a i l e y (1 9 1 3 -4 3 ) s t a t e s t h a t th e d e g re e o f g r a n u l a r i t y of
l i v e r c e l l s depends upon th e f u n c t i o n a l c o n d i t i o n o f t h a t o rg an
when th e an im al d i e d .
( 4 ) The l i v e r c e l l s were l o o s e l y h e l d t o g e t h e r , and were
a r r a n g e d in lo n g c o r d s , th e h e p a t i c c o r d s , h u t so l o o s e l y were
th e c e l l s h e l d t o g e t h e r t h a t th e c o rd d e s ig n was more o r l e s s
l o s t in th e l a r g e r p a t t e r n o f a wide s h e e t o f c e l l s , m en tio n ed
a b o v e .
( 5 ) B i le c a n a l i c u l i were form ed in th e u s u a l way by th e
c e l l s o f th e c o rd s b e in g p la c e d back to back to c r e a t e an u n
l i n e d t u b u l e .
(6) No K u'pffer c e l l s a s d e s c r i b e d by Maximow and Bloom
( 1 9 4 4 ) were s e e n .
( 7 ) Lym phocytes in s i n g l e f i l e a p p e a re d to f o llo w th e m ar
g i n s o f th e h e p a t i c c o r d s , and in t h i s way t r a c e d c o r d - p a t t e r n s .
O c c a s io n a lly ly m p h o c y tes a p p e a re d to c l u s t e r t o g e t h e r in sm a ll
t r i a n g u l a r o r d i f f u s e - s h a p e d i s l a n d s t h a t were s c a t t e r e d a t
random th ro u g h o u t th e l i v e r t i s s u e .
(e ) C e n tr a l v e in s were ab u n d an t in some l o c a t i o n s and s c a r c e
in o t h e r s The c e n t r a l v e in s were ro u n d , f r e q u e n t l y p o ly h e
d r a l in sh a p e , th e lumen a, c o n t a i n i n g a sm a ll g l o b u l a r mass o f
lym phoid t i s s u e . These sp a c e s c o u ld be c o n s tr u e d a s p r i m i t i v e
c e n t r a l v e in s in a s much a s th e h e p a t i c c o rd s were o f t e n seen
to be r a d i a l l y d i s p o s e d , such lumena w hich, in t u r n , were l i n e d
w ith a d e l i c a t e c u t i c u l a .
31
(9) A c r o s s - s e c t i o n of l i v e r showed, n o th in g more s u b s t a n
t i a l a s a c o v e rin g a n sw e rin g t h e d e s c r i p t i o n o f G l i s s o n 's
c a p s u le th a n a t h i c k e n i n g o f th e r e t i c u l u m h o l d in g t o g e t h e r th e
h e p a t i c c e l l s and c o r d s ; b u t no d e f i n i t e c o n c lu s io n c o u ld be
made a s com plete c r o s s - s e c t i o n s were n ot s e c u r e d .
THE GALL BLADDER. S e r i a l s e c t i o n s f a i l e d to shew th e p r e
sence o f a g a l l b l a d d e r .
THE SPLEEN. The f i n d i n g s in r e g a r d to th e s p le e n may be o u t
l i n e d a s f o llo w s :
( l ) The l i v e r - s p l e e n was imbedded in l i v e r t i s s u e in th e
v e n t r a l - c a u d a l r e g io n o f t h a t o r g a n , r e s t i n g upon and s u r r o u n
d in g th e p y l o r i c arm o f th e stom ach- I t was o n ly on r a r e o c c
a s i o n s , h ow ever, t h a t th e s p le e n c o u ld be seen in v iv o . Us
u a l l y th e s p le e n was b l u e , th e c o l o r of th e d e g e n e r a te d l i v e r ,
and i n d i s t i n g u i s h a b l e from th e t i s s u e o f t h a t o rg a n .
O c c a s i o n a l l y th e l i v e r w ould be a p a le o liv e -b ro w n in c o l o r ,
th e s p le e n b e in g a p a le p in k in v iv o , and u n d e r such c o n d i
t i o n s i t s s i z e , p o s i t i o n to th e stom ach and l i v e r , and shape
were as d e s c r i b e d a b o v e. Ullien th e s p le e n co u ld be r e c o g n iz e d
a s such in v iv o , i t c o n s i s t e d of l o b e s in th e form o f a f o u r -
l e a f c l u s t e r o f iv y l e a v e s s t a n d in g on end. I t was not uncommon
to f i n d a b l u e - b l a c k l i v e r and a j e t b l a c k , h a r d - t o - t h e - t o u c h
32
s p le e n c o n s i s t i n g o f s h a r p l y - p o i n t e d l o b e s t u r n e d downward
in th e d i r e c t i o n o f th e b e l l y .
( 2 ) In c r o s s - s e c t i o n th e s p le e n was r e a d i l y r e c o g n iz e d a s
such even though th e s p l e e n i c - t i s s u e was o f a p r i m i t i v e ty p e .
( 3 ) When th e s e c t i o n was s t a i n e d w ith h e m a to x y l i n - e o s in ,
a n etw ork o f round p a t t e r n s t r a c e d in b lu e was seen m arking
o f f th e to p o g ra p h y o f th e r e d p u lp w ith d e s ig n s s i m i l a r in
a p p e a ra n c e to th o s e made by th e n o d u le s o r M a lp ig h ia n b o d ie s
seen in mammalian s p l e e n . These b lu e c i r c u l a r and s e m i - c i r
c u l a r p a t t e r n s were o n ly shadowy o u t l i n e s and la c k e d th e su b
s t a n t i a l i n n e r m a t e r i a l o f i n f i l t r a t e d ly m p h o c y tes c h a r a c t e r i s
t i c o f th e t y p i c a l s p l e e n i c n o d u l e . The shadowy, p r i m i t i v e
s p l e e n i c n o d u le s o f th e anchovy d id not have more th a n a t r a c e ,
and f r e q u e n t l y l e s s th a n t h a t , o f w h ite pu lp c e n t e r s , and t h e r e
were no s h e a th e d a r t e r i e s .
( 4 ) The spongy, p i n k - c o l o r e d re d p u lp c o n s i s t e d o f e n d o th e
l i a l c e l l s h e l d t o g e t h e r in a f i n e r e t i c u l u m , th e c e l l s and th e
f i b e r s o f th e r e t i c u l u m ru n n in g in a l l d i r e c t i o n s . Maximow’ s
and Bloom’ s (1944) d e s c r i p t i o n o f th e c o rd s o f B i l l r o t h .
( 5 ) The s p l e e n i c o rg an had an abundance o f l o b u l e s ; and in
r e g a r d to th e p r o p r i e t y o r i m p r o p r i e t y o f c a l l i n g them l o b u l e s
o f M a ll, th e m a t t e r a p p e a re d to be one in w hich e i t h e r d e c i s i o n
would be J u s t i f i a b l e .
33
THE ESOPHAGUS> The t i s s u e l a y e r s o r t u n i c a s o f t h e esop h ag u s
c o n s i s t e d o f ; ( l ) th e m ucosa, (2) th e suhm ucosa, and th e t u n i c a
m u s c u l a r i s . (4) A s e r o s a was not fo u n d .
The h i s t o l o g y o f th e t u n i c a s i s a s f o llo w s ;
(1) The mucosa c o n s i s t e d o f e p i t h e l i u m u n d e r l i n e d w ith a
la m in a p r o p r i a o f c o n n e c tiv e t i s s u e .
The e p i t h e l i a l c e l l s l i n i n g th e lumen o f th e tu b e were s i m i
l a r in s t r u c t u r e and a p p e a ra n c e to th o s e in th e c a r d i a c arm o f
th e stom ach, to be d e s c r i b e d in d e t a i l p r e s e n t l y .
The la m in a p r o p r i a o f th e mucosa was throw n up i n t o t a l l , w ide,
p y ra m id a l f o l d s th e b a s e s o f w hich r e s t e d upon th e subm ucosa,
th e a p i c e s o r f r e e ends b e in g t u r n e d tow ard th e lum en. In
c r o s s - s e c t i o n th e s i d e s of t h e f o l d s n e a r th e b a se f r e q u e n t l y
o v e rla p p e d due to s l i g h t c o n s t r i c t i o n s in th e tu b e and to th e
b u l g in g o f th e b a s e s . B lake (1930) m en tio n ed th e same p a t t e r n
o c c u r r i n g in s e a - b a s s .
S t r a i g h t t u b u l a r g la n d s p e n e t r a t e d th e la m in a p r o p r i a a t
v a r y i n g d e p t h s , so t h a t in c r o s s - s e c t i o n th e c u t ends o f t u
b u l e s were f r e q u e n t l y se e n . The g la n d s were e s s e n t i a l l y l i k e
th o s e in th e c a r d i a c arm, th e d e t a i l s o f w hich a re p r e s e n t e d
u n d e r t h a t h e a d in g .
( 2 ) The submucosa was a c o n t i n u i t y of th e c o n n e c tiv e t i s s u e
o f th e lam ina o r o p r i a , b u t m o d ifie d i n t o a r e o l a r c o n n e c tiv e
t i s s u e . I t was e v id e n t a s R ogick and B lake (193 0-31) and
B o lto n ( 1 9 3 3 ) p o i n te d out t h a t t h e s e t i s s u e l a y e r s , a lth o u g h
34
FIGURE 3
B ............................................ Basement membrane
G..............................................C u t i c u l a
CE........................................... Columnar e p i t h e l i a l c e l l s
E P . . ................E p i t h e l i a l pads
L ..............................................Lamina p r o p r i a
LC........................................... Lumen o f th e canari
M..............................................M u s c u la ri s
S ..............................................Submu c osa
35
F i g . 3
C r o s s - s e c t i o n o f a p o r t io n of one w a ll o f th e c a r d i a c arm o f
th e stom ach. On th e l e f t o f th e f i g u r e i s th e m u s c u l a r i s , on
th e r i g h t , th e lum en. The m a g n i f i c a t i o n i s lOXlOX . 188,
36
c o n t i n u i t i e s , se rv e d i f f e r e n t p u rp o se s f u n c t i o n a l l y a s w e ll
a s make d i f f e r e n t t o p o g r a p h i c a l p a t t e r n s in th e w a l ls o f th e
d i g e s t i v e c a n a l .
( 3 ) The re was a w ide, s i n g l e - 1 a y e r t u n i c a m u s c u la r is t h a t
c o n s i s t e d o f c i r c u l a r r u n n in g , sm ooth, n o n - s t r i a t e d f i b e r s .
A narrow e x t e r n a l m uscle l a y e r o f sm ooth, s t r i a b e d , l o n g i t u
d i n a l l y - r u n n i n g f i b e r s c o u ld o c c u r in l i m i t e d a r e a s .
B lak e (1930) n o te d t h a t in s e a - b a s s th e w id e , c i r c u l a r ,
i n t e r n a l m uscle l a y e r c o n s i s t e d o f sm ooth, s t r i a t e d f i b e r s .
THE CARDIAC ARM OF THE STOMACH. The t u n i c a s o f th e c a r d i a c
arm c o n s i s t e d o f t h e f o llo w in g s t r u c t u r e s : (1) th e m ucosa,
( 2 ) th e subm ucosa, ( 3 ) th e m u s c u l a r i s . ( 4 ) A s e r o s a was not
i d e n t i f i e d .
The h i s t o l o g y o f th e t u n i c a s i s a s f o llo w s :
( 1 ) The mucosa c o n s i s t e d o f t a l l , colum nar e p i t h e l i a l c e l l s
c o v e re d by a t h i n t o p - p l a t e o r c u t i c u l a and r e s t i n g on a b a s e
ment membrane. G o b le t c e l l s f r e q u e n t l y o c c u r r e d . The C u t i
c u la was n o t s t r i a t e d . B lake (1930) r e p o r t e d t h a t in s e a b a s s
th e c u t i c u l a c o n ta in e d s t r i a t i o n s t h a t were p e r p e n d i c u l a r to
th e lumen o f th e tu b e . The c e l l s o f th e e p i t h e l i u m were g r a n u
l a r , and c o n ta in e d l a r g e , round o r i r r e g u l a r - s h a p e d n u c le i
n e a r th e basem ent m em brane.
The g a s t r i c g la n d s were s t r a i g h t t u b u l e s p e n e t r a t i n g th e
la m in a p r o p r i a o f th e mucosa t h a t c o n s i s t e d o f dense c o n n e c tiv e
37
t i s s u e , and was throw n up i n t o t a l l , oblo n g f o l d s . The c r y p t s
o r t u b u l e s te n d e d to narrow i n t o a neck a s th e y e n te r e d th e
la m in a p r o p r i a . The e p i t h e l i a l c e l l s above th e neck were
m ark e d ly t a l l . Those below t h e c o n s t r i c t i o n g r a d u a l l y assumed
th e shape o f low colum nar o r c u b o id a l c e l l s o f a p y ram id al
s h a p e , th e b ro a d b a s e s r e s t i n g upon th e basem ent membrane, th e
a p i c e s t u r n e d to w a rd th e lumen o f th e c r y p t . The e p i t h e l i a l
c e l l s below th e n e c k s o f t h e t u b u l e s were m ark ed ly more g r a n u
l a r th a n th o s e a b o v e.
The la m in a n r o p r i a o f th e c a r d i a c arm was d i s t i n c t i v e f o r th e
t a l l , w id e, o b lo n g f o l d s i t p r o j e c t e d i n t o t h e lumen o f th e
d i g e s t i v e c a n a l . The b l u n t , f r e e edg es o f th e f o l d s were i n
d e n te d to form w id e , roun d o r p y ra m id a l l o b e s . D i s t i n c t i v e o f
t h e s e f o l d s were th e e p i t h e l i a l p a d s , c o n s i s t i n g o f s t r a t i f i e d ,
squamous e p i t h e l i u m .
( 2 ) The submucosa c o n s i s t e d o f a w e ll-m a rk e d l a y e r o f
a r e o l a r c o n n e c tiv e t i s s u e , b u t t h e r e were a r e a s where t h i s t u n i
ca n arro w ed to o n ly a t h r e a d - l i k e c o n t i n u i t y . Lymph space
b o u n d a r i e s b etw een th e la m in a p r o p r i a and th e submucosa were
s e e n .
( 3 ) The t u n i c a M u s c u la r is c o n s i s t e d o f two m uscle l a y e r s ,
an i n n e r and an o u t e r . The i n n e r m uscle l a y e r was w id e , and
c o n s i s t e d o f c i r c u l a r r u n n in g , s t r i a t e d , smooth f i b e r s . The
o u t e r m uscle l a y e r was n a rro w , and c o n s i s t e d o f l o n g i t u d i n a l ,
sm ooth, n o n - s t r i a t e d f i b e r s .
38
F IG U R E 4
B............................................................Basem ent m em b ran e
C .............................................................Cuticula
C E ..................... ....................................C olum nar epithelial cells
1............................................................Indentation or constriction in the wall
of the m u s c u l a r i s
L............................................................L am ina propria
L C .........................................................L u m en of the canal
M ............................................. .Muscularis
S............................................ Subm ucosa
39
m
m
F i g . 4
C r o s s - s e c t i o n o f a p o r t i o n o f th e w a ll o f th e p y l o r i c arm
o f th e stom ach. At th e l e f t o f th e f i g u r e i s th e m u s c u l a r i s ;
on th e r i g h t i s th e lum en. The m a g n i f i c a t i o n i s lOxlOx . 168.
40
The two extrem e ends o f t h e c a r d i a c arm , th e a n t e r i o r and th e
c a u d a l e n d s , were s i m i l a r in t h e i r c r o s s - s e c t i o n a p p e a ra n c e to
t h a t o f t h e e so p h a g u s .
The m id dle r e g i o n , d e s c r i b e d a b o v e, d i f f e r e d s u f f i c i e n t l y ,
p a r t i c u l a r l y in r e g a r d to th e shape o f th e oblong f o l d s and
t h e p r e s e n c e o f e p i t h e l i a l p a d s , to be d i s t i n g u i s h e d from any
o t h e r r e g i o n o f th e d i g e s t i v e c a n a l .
THE CAECUM OF THE STOMACH. The caecum o f th e stom ach i s a
s h o r t , b l i n d pouch p r o j e c t i n g c a u d a l l y from th e elbow curve
made by th e j u n c t u r e o f th e c a r d i a c and p y l o r i c arm s of th e
sto m ach . I t s p r e s e n c e i s a c h a r a c t e r i s t i c o f t e l e o s t f i s h .
A c r o s s - s e c t i o n o f t h i s r e g i o n i s a lm o st a d u p l i c a t e o f th e
e so p h ag u s in a p p e a ra n c e . The t u b u l e s p e n e t r a t i n g th e la m in a
p r o p r i a a p p e a re d to be more s l e n d e r th a n th o s e in th e e so p h a
g u s .
T here was no subm ucosa.
The t u n i c a m u s c u l a r i s was in d e n te d e x t e r n a l l y to shov; deep
c o n s t r i c t i o n s . There were two m uscle l a y e r s , a wide i n n e r l a y e r
o f sm ooth, c i r c u l a r - r u n n i n g , n o n - s t r i a t e d f i b e r s ; and a n arro w
o u t e r l a y e r o f l o n g i t u d i n a l , sm ooth, n o n - s t r i a t e d f i b e r s .
THE PYLORIC ARM OF THE STOMACH. The two ends o f th e p y l o r i c
arm were s i m i l a r in a p p e a ra n c e to c r o s s - s e c t i o n s o f th e e so p h a
gus and caecum. The m id d le s e c t i o n showed a h i s t o l o g y s p e c i f i c
41
to i t s e l f , and t h i s i s th e r e g i o n r e f e r r e d in t h e f o llo w in g
o u t l i n e . The t u n i c a s o f th e p y l o r i c arm c o n s i s t e d o f : ( l ) th e
m ucosa, (2) th e suhm ucosa, and (3) th e m u s c u l a r i s . (4) A s e r o s a
was n o t i d e n t i f i e d a s su c h .
The h i s t o l o g y o f th e p y l o r i c arm i s as f o l lo w s :
(1) The e p i t h e l i a l c e l l s o f th e mucosa were l i k e th o s e d e s
c r i b e d in t h e c a r d i a c arm .
The g a s t r x a g la n d s w ere s t r a i g h t t u b u l e s , b u t were crowded
so c l o s e l y t o g e t h e r t h a t t h e co n seq u e n t f o l d s o f th e la m in a
p r o p r i a d i v i d i n g them h a d th e c r o s s - s e c t i o n a p p e a ra n c e o f t a l l
v i l l i . E p i t h e l i a l .pads were n o t found in t h i s r e g i o n . B i f u r
c a t i o n o f a f o l d , h u t n o t o f a t u b u l e , o c c a s i o n a l l y o c c u r r e d .
The la m in a p r o p r i a . The la m in a p r o p r i a c o n s i s t e d o f compact
c o n n e c tiv e t i s s u e o f t e n s e p a r a t e d from th e basem ent membrane o f
th e e p i t h e l i u m o f th e mucosa by a lym p h -sp ace b o u n d a ry . The
la m in a p r o p r i a , a s was m e n tio n e d a b o v e, was throw n up i n t o t a l l ,
n a rro w f o l d s t h a t in c r o s s - s e c t i o n h ad th e a p p e a ra n c e o f v i l l i .
( 2 ) The submucosa c o n s i s t e d o f a r e o l a r c o n n e c tiv e t i s s u e ,
o c c a s i o n a l a r e a s showing t h i s t i s s u e - l a y e r n a rro w in g to a t h r e a d
l i k e t h i c k n e s s .
( 3 ) The t u n i c a m u sc u la rim was in d e n te d e x t e r n a l l y by c o n s t
r i c t i o n s . The wide i n t e r n a l m uscle l a y e r c o n s i s t e d o f c i r c u
l a r r u n n in g , sm ooth, s t r i a t e d f i b e r s . The narrow e x t e r n a l
m uscle l a y e r c o n s i s t e d o f l o n g i t u d i n a l , sm ooth, n o n - s t r i a t e d
f i b e r s •
42
THE PYLORIC CAECA. The p y l o r i c c a e c a were b l i n d t u b e s t h a t
were o u tp o u c h in g s o f th e p y l o r i c arm o f th e sto m ach . They a r e
c a l l e d by many a u t h o r s th e h e p a t o - p a n c r e a s .
H egner (1 9 4 2 ) , B o lto n (1 9 3 3 ), and R ogick and B lake (1 9 3 0 -3 1 )
i n t e r p r e t e d t h e s e b l i n d s a c s a s m echanism s f o r i n c r e a s i n g th e
s e c r e t i n g s u r f a c e o f th e p y l o r i c arm o f th e stom ach, r e p o r t i n g
t h a t th e y s e c r e t e d i g e s t i v e enzym es, and t h a t t h e r e can be from
two to two h u n d re d in num ber, d e p en d in g upon th e s p e c i e s of
f i s h . The p r e s e n t w r i t e r c o u n te d n o t l e s s th a n s i x in th e anchovy,
and t h e r e were o f t e n t w e n ty - f o u r l a r g e o n e s . T here were o f t e n
c l u s t e r s o f c a e c a to o sm all to be c o u n te d w ith th e naked ey e.
A c r o s s - s e c t i o n o f th e p y l o r i c caecum a p p e a re d to be th e dup
l i c a t e o f t h a t o f th e p y l o r i c arm o f th e sto m a c h . The p y l o r i c
c a e c a d i f f e r e d from t h e i r p a r e n t tu b e in t h a t th e lumena of th e
s t r a i g h t t u b u l e s v;ere n a rro w e r p e r m i t t i n g a g r e a t e r number o f
t u b u l e s p e r u n i t of l e n g t h o f th e lum en. T h is r e s u l t e d in th e
la m in a p r o p r i a b e in g throw n up i n t o n a rro w e r f o l d s th a n were
found in th e p y l o r i c arm.
S e c t i o n s of t h i s l o c a l i t y o f th e d i g e s t i v e c a n a l u s u a l l y
showed t h e p r e s e n c e o f c r o s s - s e c t i o n s o f th e sm a ll i n t e s t i n e .
The d i f f e r e n c e betw een a c r o s s - s e c t i o n o f th e p y l o r i c c a e ca and
th e sm a ll i n t e s t i n e was t h a t in th e c a e c a th e f o l d s were t a l l
and n a rro w , and th e m uscle l a y e r was wide and c o n s i s t e d o f two
l a y e r s w h e rea s in th e i n t e s t i n e th e f o l d s were s h o r t and w ide,
and th e s i n g l e - l a y e r m u s c u l a r i s was n a rro w .
43
THE PYLORIC VALVE. T h is r e g io n o f th e p y l o r i c arm o f th e
stom ach s e r v e s th e same p u rp o se in f i s h t h a t th e p y l o r i c s p h i n
c t e r does in mammals. I t i s th e extrem e a n t e r i o r end o f th e
p y l o r i c arm a t th e j u n c t u r e o f t h a t organ w ith th e duodenum.
A c r o s s s e c t i o n o f th e p y l o r i c v a lv e was s i m i l a r to t h a t of
th e eso p h ag u s and th e caecum o f th e stom ach e x c e p t t h a t t h e r e
was a m u s c u la r is c o n s i s t i n g o f a s i n g l e l a y e r o f c i r c u l a r l y -
r u n n in g , sm ooth, n o n - s t r i a t e d f i b e r s , th e t u n i c a b e in g to o wide
and t h e lumen to o c o n s t r i c t e d to p e rm it t h i s r e g i o n b e in g co n
fu s e d w ith t h a t o f any o t h e r s e c t i o n o f th e d i g e s t i v e c a n a l .
T here was no subm ucosa.
THE SMALL INTESTINE. Sample c r o s s - s e c t i o n s o f sm a ll i n t e s t i n e
w ere ta k e n from d i f f e r e n t r e g i o n s in t h a t o rg a n . The h i s t o l o g y
was e s s e n t i a l l y th e same in a l l th e s a m p lin g s . An o u t l i n e o f
th e t u n i c a s o f th e sm a ll i n t e s t i n g i s a s f o l lo w s :
( l ) The mucosa c o n s i s t e d o f e p i t h e l i u m , e s s e n t i a l l y l i k e
t h a t o f t h e c a r d i a c arm o f th e stom ach, r e s t i n g upon a la m in a
p r o p r i a .
The la m in a p r o p r i a was throw n up i n t o s h o r t , wide f o l d s t h a t
f r e q u e n t l y showed l o b u l a t i o n . F o ld s in th e a n t e r i o r r e g io n o f
th e sm a ll i n t e s t i n e were o b lo n g in sh a p e , th e f r e e ends b e in g
b l u n t o r ro u n d e d . In th e p o s t e r i o r r e g io n and th e a n u s , th e
f o l d s were o f t e n o a k - l e a f in sh a p e, th e b r o a d , lo b e d b a s e s
a t t a c h e d to th e lam in a p r o p r i a o f th e t u b e .
44
The la m in a p r o p r i a th ro u g h o u t th e sm a ll i n t e s t i n e o f th e a n
chovy showed a h i s t o l o g y s i m i l a r , in r e g a r d to th e p re s e n c e o f
m ast c e l l s , in a p p e a ra n c e to t h a t o f th e salmon and t r o u t a s
r e p o r t e d by B o lto n ( 1 9 3 3 ).
In th e anchovy sm a ll i n t e s t i n e , th e la m in a p r o p r i a o f a f o l d
c o n s i s t e d o f dense c o n n e c tiv e t i s s u e , and was d i v id e d by a
t h r e a d o f d e n s e r c o n n e c tiv e t i s s u e i n t o two r e g i o n s , an o u t e r
and an i n n e r . The c e n t r a l p a r t of t h e i n n e r r e g io n had a den se
c o n n e c tiv e t i s s u e d o r e . U s u à lly th e e n t i r e f o l d was a g r a n u l a r
mass a r e s u l t o f th e p r e s e n c e o f m ast c e l l s . F r e q u e n t l y th e
m ast c e l l s would be m assed a lo n g th e t h r e a d - l i k e b o u n d ary l i n e
betw een th e o u t e r and i n n e r zone c r e a t i n g a s t r a tu m g ra n u le su m .
The la m in a p r o p r i a was bounded on i t s d i s t a l s id e by th e
b asem ent membrane o f th e e p i t h e l i u m , and on i t s p ro x im a l s id e
by a ly m p h -sp ac e b o u n d a ry , o f t e n f i l l e d w ith d i s c o n ti n u o u s
s h e e t s o f a r e o l a r c o n n e c tiv e t i s s u e .
( 2 ) T here was no submucosa o t h e r th a n th e d i s c o n t i n u o u s ,
f r u g a l s u p p ly o f a r e o l a r c o n n e c tiv e t i s s u e in th e ly m p h -sp ace
b o u n d a ry betw een th e la m in a p r o p r i a and th e t u n i c a m u s c u l a r i s .
( 3 ) The t u n i c a m u s c u l a r i s was t h e n a r r o w e s t m u s c u la r is found
in th e e n t i r e d i g e s t i v e c a n a l . I t c o n s i s t e d f o r th e most p a r t
o f a wide i n t e r n u s o f c i r c u l a r l y r u n n in g , sm ooth, s t r i a t e d f i
b e r s . A d i s c o n t i n u o u s n a rro w e x t e r n u s co u ld o c c u r .
( 4 ) No s e r o s a was fo u n d .
45
V . DISCUSSION
1 . So f a r a s i s known to th e p r e s e n t w r i t e r , Cunningham’ s
and C ald erw o o d ’ s (1 8 8 9 -9 1 ) r e p o r t s a r e th e o l d e s t r e f e r e n c e s
d e s c r i b i n g th e h i s t o l o g y o f t h e o v a ry o f th e anchovy, and i n d i
c a t i n g t h a t th e h i s t o l o g y o f th e t e s t i s i s t h e same a s th e o v a ry
w ith s l i g h t v a r i a t i o n s . S u b seq u e n t i n v e s t i g a t o r s have c o n c u rr e d ,
and t h i s p r e s e n t i n v e s t i g a t i o n i s c o r r o b o r a t o r y .
2 . So f a r a s c o u ld be a s c e r t a i n e d , no m en tio n i s made in th e
l i t e r a t u r e o f any one h a v in g i d e n t i f i e d bona f i d e anchovy s p e r -
m ataz o a o r m i l t . The p r e s e n t w r i t e r f a i l e d to do s o .
3 . P r e v io u s r e p o r t s s t a t e d t h a t s a t i s f a c t o r y c r o s s - s e c t i o n s
o f th e v i s c e r a o f th e anchovy were d i f f i c u l t to o b t a i n owing to
th e i n h e r e n t f r a g i l i t y o f th e t i s s u e . From th e e a r l i e s t r e p o r t s
down to th e p r e s e n t ones i t was rem arked t h a t s a t i s f a c t o r y
c r o s s - s e c t i o n s o f th e l i v e r , k id n e y s , and s p le e n were a lm o st
im p o s s ib le to s e c u re due to t h e f a c t t h a t th ro u g h o u t th e y e a r
t h e s e o rg a n s a r e i n t e r m i t t e n t l y i n f i l t r a t e d w i t h ly m p h a tic t i s s u e ,
and t h i s was th e f i n d i n g o f th e p r e s e n t w r i t e r .
4 . The p r e s e n t i n v e s t i g a t i o n o f t h e l i v e r , r e n a l ly m p h a tic
o rg a n , s p l e e n , and d i g e s t i v e c a n a l o f th e anchovy showed t h a t
t h e s e o rg a n s a r e m e re ly v a r i a t i o n s o f th e form s o f th o s e o rg a n s
in o t h e r w ell-know n c lu p e o id f i s h .
5 . So f a r a s c o u ld be l e a r n e d , t h e a v a i l a b l e l i t e r a t u r e d oes
n o t m en tio n any one h a v in g v e r i f i e d th e p re s e n c e o f o r ab sen c e
46
o f th e g a l l b l a d d e r i n a n c h o v i e s . The p r e s e n t i n v e s t i g a t i o n
f a i l e d to f i n d such an org an p r e s e n t .
7 . No s e r o s a a s an o u t s i d e c o v e r in g o f th e d i g e s t i v e c a n a l
■was fo u n d . T here was no p o s i t i v e e v id e n c e to show w h e th e r i t
was d esq uam ated by th e b la d e o f th e m icrotom e o r w h e th e r i t was
a b s e n t in v i v o . A s e r o s a i s r e p o r t e d p r e s e n t i n o t h e r fo rm s o f
c lu p e o id f i s h . So f a r a s i s known t o t h i s w r i t e r , no p r e v i o u s
r e p o r t h a s been made in r e g a r d to t h e s e r o s a o f th e g a s t r i c
c a n a l o f a n c h o v i e s .
47
V I . SUM M ARY
1 . The u r o - g e n i t a l sy ste m , and o t h e r o rg a n s o f th e v i s c e r a
such a s th e l i v e r , g a l l b l a d d e r , s p le e n , and the d i g e s t i v e c a n a l
o f E n g r a u l i s m ordax m ordax, G i r a r d , have been exam ined and t h e i r
h i s t o l o g i c a l s t r u c t u r e d e s c r i b e d .
2 . The h i s t o l o g y o f th e a b o v e -m e n tio n e d sy ste m s i s found to
be o n ly a s l i g h t v a r i a t i o n o f t h a t i n o t h e r c l o s e l y - r e l a t e d
s p e c i e s o f t e l e o s t s .
3 . The d i g e s t i v e c a n a l from th e e so p h ag u s to th e duodenum
in E n g r a u l i s m ordax m ordax i s found to be l i k e th e e n t i r e d i
g e s t i v e c a n a l o f s e a b a s s i n t h a t t h e r e i s a s c a r c i t y o f m ast
c e l l s in th e t i s s u e l a y e r s . From th e duodenum to th e an us in
th e d i g e s t i v e c a n a l o f E n g r a u l i s m ordax m ordax t h e r e i s an a b
undance o f m ast c e l l s in th e la m in a p r o p r i a , and in t h i s r e
s p e c t th e c a n a l i s s i m i l a r to th e e n t i r e d i g e s t i v e c a n a l o f
salm on and t r o u t a s d e s c r i b e d by B o lto n (1 9 3 3 ).
4 . The o u t s i d e T is s u e l a y e r o f th e g e n i t a l d u c t o f th e m ale
was found to be c o n tin u o u s w ith th e c a p s u le o f th e t e s t i s , and
i s l i n e d w ith co lum nar e p i t h e l i a l c e l l s .
5 . The w a l l s o f th e c a r d i a c arm o f th e stom ach were found
to c o n ta in oblo n g f o l d s w hich a r e a s s o c i a t e d w ith e p i t h e l i a l
p a d s , and which a re a d i s t i n g u i s h i n g c h a r a c t e r i s t i c o f t h i s
r e g i o n .
6 . Long, narro w f o l d s o f th e la m in a p r o p r i a a r e d i s t i n c t i v e
f o r th e p y l o r i c arm o f t h e stom ach, and th e y e x te n d i n t o th e
48
p y l o r i c c a e c a of w hich t h e r e a re from s i x to t h i r t y - s i x in
num ber.
7 . The m orphology and h i s t o l o g i c a l s t r u c t u r e s o f E n g r a u l i s
m ordax mordax were found to show wide v a r i a t i o n , a te n d e n c y
t h a t i s a c h a r a c t e r i s t i c o f c lu p e o id f i s h .
49
V I I . BIBLIOGRAPHY
AUTHOR
Adams, L. A.
B a i l e y , P. R.
B a r n h a r t , P e rc y S p e n c e r
B la k e , I r v i n g H.
PUBLICATION
B o l i n , R o lf L.
B o lto n , L loyd L .
Bowie, D. J .
B r e d e r ,
R .P .
C• M• J r .,
B r id g e , Thomas Wm.
1933 N e c tu r u s , A L ab . S tu d y . The
M acM illan Co. N.Y. 1933.
1943 T extbook of H i s t o l o g y . The
M acM illan Co. 1 9 1 3 -4 3 .
1930 M arine P i s h e s o f S o u th e rn
C a l i f o r n i a , U n i v . o f C a l i f .
P r e s s . 1930
1930 S t u d i e s on t h e C om parative
H i s t o l o g y o f th e D i g e s t iv e
Tube of C e r t a i n T e l e o s t
P i s h : S e a - b a s s , C e n t r o p r i s -
t i s S t r i a t u s Jo u rn .M o rp h .
1930
1936 Em bryonic and E a r l y L a r v a l
S ta g e s o f th e C a l i f o r n i a
Anchovy, E n g r a u l i s Mordax
M o r d a x , G i r a r d , C a l i f . P i s h and
Game Pub. V .2 , No.4 ,1 9 3 6
1933 B a s o p h i l i c C e l l s i n th e A l i
m e n ta ry C anal o f Salm onoid
P is h e s .J o u r .M o r p h .V .6 4 ,N o .
3 , June 1933.
1924 C y t o l o g i c a l S t u d i e s in th e
I s l e t s o f L a n g e rh a n s in a
T e l e o s t , Neomaens G r i s e u s .
A n a t . R e c .V .2 9 ,No.1 ,1 9 2 4 .
and N i g r e l l i ,
1935 The I n f l u e n c e o f T em p erature
and O th e r F a c t o r s on th e
W in te r A g g r e g a tio n o f Sun-
f i s h , L e p o n is A u r e t i s . E c o
lo g y , V. 16 .pp . 33-47 ,1935 .
1890 S t r u c t u r e and F u n c tio n o f
The A ir b l a d d e r in P i s h . P r o c .
P h il.S o c .B irm in g h a m , 1890-
9 7 . p p . 1 4 4 - 8 7 .1 8 9 0 .2 .
50
V II. BIBLIOGRAPHY
AUTHOR
Buchbaum, R alph
C alderw ood, W.L.
PUBLICATION
Chapman, W il b e r t
Cunningham, Jo sep h
1938 A nim als W ith o u t B a c k b o n e s .U n i.
C h i . P r e s s .1 9 3 8 .
1891 On t h e Ovary an d I n t r a - O v a -
r i a n Egg in T e l e o s t e a n s . J o u r n .
M arine B i o l . A s s o c . 2 , p p . 298-
3 1 3 .1 8 9 1 .2 .N .S .
1895 O v a rie s o f F i s h e s . J o u r n . M a r .
B i o l . A s s o c .1 8 9 3 - 9 5 .N .S . 3 , pp.
1 5 4 - 1 6 5 .P l a t e s .
1891 The Head K idney o f T e l e o s t e a n s
Jo u rn .M a rin e B i o l . A s s o c .1 6 9 1 -
2 , p p . 43.
1944 The C om p arativ e O s te o lo g y o f
H e r r i n g - l i k e F i s h e s , C lu p e i-
d a e . C a l i f . F i s h and Game Pub.
V .3 0 .N o .1 .1 9 4 4 .
1884 C r i t i c a l Note on th e L a t e s t
T heory o f V e r t e b r a t e M orpholo
gy .Pro c .R o y .S o c .E d b .1 8 8 4 -1 2 ,
p p .7 5 6 - 6 5 .1 8 8 4 .1 .
1893 The Im m ig sa tio n o f A ncho vies in
th e E n g l i s h C h a n n e l.J o u r n .M a r .
B i o l . A s s o c .1 8 9 3 - 9 5 .N .S . 3 .p p .
3 0 0 - 3 0 3 .1 8 9 3 .7 .
1889 A n chov ies in th e E n g l i s h Chan
n e l .J o u r n . M a r .B io l.A s s o c .
1 8 8 9 -9 0 . N . S . I . p p . 3 28-33 9,
1 8 8 9 .1 .
1897 On th e H i s t o l o g y o f th e Ovary
and O v a ria n Ovum i n T e l e o s t e a n s
Jo u rn .R o y .M i c r.S o c .L o n d o n .
1 8 9 7 .p p . 353-354
1887 On th e Developm ent o f th e O vi
d u c t in T e l e o s t a n s . P r o c . R o y .
P h y s .S o c .E d b .U 8 8 7 .p p .116 ;
3 4 2 - 4 5 .1 8 8 7 .3 .
51
V II. BIBLIOGRAPHY
AUTHOR
Dean, B a s h fo rd
G ray, Henry
G reen , C.W.
PUBLICATION
G re g o ry , Wm.K.
H a r g i t t , G e o .T .
H eg ner, R o b e rt W.
1895 P i s h e s , L iv in g and F o s s i l .T h e
M acM illan C o.N .Y .1895.
1944 Human Anatom y.Lea and P e b ig e r
C o .1944.
1912 The Anatomy and H i s t o l o g y o f
The A lim e n ta ry T r a c t o f King
S a l m o n . B u l l . U . S . F i s h e r i e s . V .
3 2 ,p p . 7 3 -1 0 0 .1 9 1 2 .
1931 F i s h e s , C a ta lo g u e s , and C o l l
e c t i o n s . Amer .Mu s . Nat . H i s t .
N .Y .1931.
1920 Germ C e l l s o f C o e l e n t e r a t e s .
J o u r .M orph .V .3 3 .1 9 1 9 - 2 0 . p p . l .
1942 C o lle g e Z oology.T he M acM illan
C o .1942.
H ild e b r a n d , Samuel P . , and C a b le , L o u e l la E .
1930 A t l a n t i c C oast A nchov ies .Re
p o r t in B u r .F is h e r ie s .D o c u m e n t
No.1 0 9 3 .V.XLVI.1930
Holm es, Samuel J .
Hubbs, C a r l . L.
H u x ley , Thomas
1935 B io lo g y o f th e F ro g .T h e Mac
M illa n Co. N.Y. 1935.
1925 M arine F i s h e s of C a l i f o r n i a .
Pub. i n Z o o l.V .1 6 .N o .1 3 .1 9 2 5 .
1925 P u b .i n Z o o l.V .1 6 . p p .1 5 3 -1 3 8 .
1925.
1883 C o n t r i b u t i o n s to M orphology;
O v id u c ts of s m e l t . P r o c . Z o o l .
Soc.L ond on, p p . 1 3 2 -1 3 9 .1 8 8 3 .
J o r d a n , D avid S. and Everm ann, B. W.
1858 E n g r u a l i s C om pressa, G i r a r d ,
An A n c h o v y .P a c .R .R .S u r v .1 0.
p p . 3 3 6 -4 4 7 .San D i e g o .1858.
52
V II. BIBLIOGRAPHY
AUTHOR
K e llo g g , Vernon L.
K e n d a l, W.C.
K u n tz , A l b e r t
PUBLICATION
L o c k in g to n to n ,W .H .
M acC ulloch, John
M anning, F .B .
Maximow, A .A .,a n d Bloom,Wm.
M o r r is , C h a r le s
N a i r , K. K ris h n a n
N a k a h a ra , Waro
N ishekaw a, T.
Norman, J . R.
P a i n t e r , T .S .
1906 E le m e n ta ry Z oolo gy.H enry H a l l
and C o .N .Y .1906.
1921 O v id u c ts o f T e l e o s t e i . B u i . U . S .
B u r . F i s h e r i e s .1 9 2 1 .V .3 7 .p p .
189.
1933 The Em bryology and L a r v a l De
velopm ent o f B a i r d i e l l a Chry-
s u r a , and A nchovia M i t c h i l l i .
B u l l . U . S . B u r . F i s h e r i e s .1 9 3 3 .
p p .1 - 1 9 .
1880 No T i t l e ; On F i s h . P r o c . N a t .Mus.
P a m p h le t .p p .6 3 -6 8 .1 8 8 0 .
1893 On H e r r i n g . Q u a r t . J o u r . S c i . A r t s ,
L o n d o n .1 8 9 3 .V .1 6 .p p .2 1 0 -2 8 .
1924 H e a rin g in G o l d f i s h . J o u r . E x p .
Z o o l.V .4 1 . p p . 5 .1 9 2 4 - 5 .
1944 T extbook o f H is to lo g y .W .B .
S a u n d e rs C o .1944.
1895 SwimBladder of F i s h e s . P r o c .
P h i l a d . 1 8 8 5 -9 2 .V .6 . p p . 1 2 4 -3 5 .
1940 R e co rd s o f th e I n d i a n Museum.
V .X L I I .P a r t I I . p p . 2 7 7 - 8 8 .On
Anchovy E n g r a u l i s T e lo s a ;
C a l c u t t a , Ju n e , 1940.
1917 On th e P h y s io lo g y o f th e
N u c l e o l i in S i l k G lan d s o f
C e r t a i n I n s e c t s .J o u r .Morpti.
p p . 5 5 , J u n e , 1917.
1901 On th e D evelopm ent o f th e
Anchovy, E n g r a u l i s J a p o n i c u s .
J o u r .F i s h . B u r .T o k y o . 1 0 . N r .190 1.
1931 A H i s t o r y o f F i s h e s . The F r e
d e r i c k S to k e s C o .N .Y .1931.
1921 R e p t i l l i a n S p e r m a to g e n e s is .
J o u r . E x p . Z o o l . V . 3 4 . p p . 281 -1 9 2 1 .
53
V II. BIBLIOGRAPHY
AUTHOR PUBLICATION
P a r k e r , T. J . and H a s w e ll, W.A.
1940 T extbook o f Zoology.T he Mac
M illa n C o .N .Y .1940.
P a r k e r , Wm. N.
P a r r , A. E .
Payne, PernanduH
1882 K idney s o f T e l e o s t e i . R p t . B r i t
A s s o c .A d v .S c i .p p .5 7 7 .1 8 8 2 .1 .
1927 Sex Dim orphism an d S c h o o lin g
B e h a v io r of P i s h e s . O c c a s i o n a l
P a p e r o f th e Bingham Oceano
g r a p h i c C o l l . V . l . p p . 3 6 .1 9 2 7 .
1916 Germ C e l l s o f C r y l l o t a l p a and
B o r e a l i s .J o u r . M o r p h .V .2 8 .1 9 1 6 •
p p . 287.
P i e r s o l (No i n i t i a l s l i s t e d ) 1921 Normal H i s t o l o g y . J . B . L i p p i n c o t t
C .1 9 2 1 .
P o t t e r , G eo.E.
R e g n a r t ,H .C .
R ic h , W i l l i s , H.
Ridewood, W.G.
R o g ick , M.D.
S p o o n e r, G.
S t a r k s , E. C.
1938 T extbook o f Z oology.T he G.V.
Mosby C o . S t . L o u i s .1 9 3 8 .
1931 The Lower L im its o f P e r c e p t io n
o f E l e c t r i c a l C u r r e n t s by P i s h .
J o u r .M a r .B io l.A p S o c .o f th e
U n ite d K ingdom .V .1 7 . 2 . 1 9 3 1 .p p .
4 1 5 -2 0 .
1920 E a r l y H i s t o r y of Seaw ard Mi
g r a t i o n o f C hinook Salm on.
B u l l . B u r . F i s h e r i e s . V . 3 7 .19 19-20
1904 On th e C r a n ia l O s te o lo g y o f th e
C lu p e o id F i s h e s . P r o c . R o y . Z o o l .
S o c .L o n d o n .V .2 .1 9 0 4 .
1931 S t u d i e s on th e C om parative
H i s t o l o g y of th e D i g e s t i v e Tube
o f C e r t a i n T e l e o s t F i s h . A
Minnow. J o u r .M o r p h .1931.
1931 Some O b s e r v a tio n s on S c h o o lin g
in F i s h . Mar. B i o l .A s s o c .U n i t e d
K ingdom .V .1 7 . p p . 4 2 1 -8 1 .1 9 3 1 .
1918 The C a l i f o r n i a A n c h o v y .C a l i f .
F i s h and Game M ag.V .4 .1 9 1 8 .
54
V II. BIBLIOGRAPHY
AUTHOR PUBLICATION
S t a r k s , E d .C . and Mann, Wm.M.1930 New and R are F i s h o f S o u th e rn
C a l i f cr n i a . U n i. o f C a l i f .
P u h l. in Z o o l.V .8 .1 9 3 0 .
S t o r r e r , T rac y I .
S u r r a r e r , T.
T ak u , Komai
Townsend, H enry C.
T ra c y , H enry
T u r n e r , C la re n c e L.
U lr e y , A l b e r t B.
W illia m s o n , Henry C.
1943 G e n e ra l Z oology.T he McGraw-
H i l l Book Co. N.Y. 1943.
1931 I n t e s t i n a l T r a c t o f Two Min
nows. Ohio J o u r , o f S c i e n c e .
1931.
1920 S p e rm a to g e n e s is o f S q u i l l a
O r o t o r i a . J o u r .M o r p h .V .3 4 .
p p . 3 0 7 .1 9 2 0 .
1919 H i b e r n a t i n g F i s h e s . B u l l . U . S .
Z o o l.S o c .V .1 9 . p p . 3 4 5 .1 9 1 9 .
1920 The C lu p e o id C ranium and Swim
B la d d e r . J o u r .Morph. V. 33. pp.
4 3 9 .1 9 1 9 -2 0 .
1919 The S e a s o n a l C ycle in th e
Sperm ary o f P e r c h . Jo u r.M o rp h .
V .2 2 . p p . 6 8 1 .1 9 1 9 .
1928 F i s h e s o f S o u t h .C a l i f . T h e
Calif.Acad.Sci.Bull.B.2 8 .1 8 2 8 .
1896 Stom . and R e p ro d u c tiv e O rgans
o f t h e S e a - t r o u t and H e r r i n g s .
14 A n n .R p t.F is h e r y B o a rd ,
S c o t l a n d . 1 8 9 6 .1 .
Z o e th o u t, Wm.D.and T u t t l e , W.W.
1943 T extbook o f P h y s io lo g y .T h e G.
V.Mosby C o . S t . L o u i s .1 9 4 3 .

Asset Metadata

Creator MacDonald, J. T. (author)

Core Title A re-investigation of the histology of certain organs of the viscera of the California anchovy, Engraulis mordax mordax, Girard

Contributor Digitized by ProQuest (provenance)

Degree Master of Arts

Publisher University of Southern California (original), University of Southern California. Libraries (digital)

Tag Biological Sciences,OAI-PMH Harvest

Format application/pdf (imt)

Language English

Permanent Link (DOI) https://doi.org/10.25549/usctheses-c39-274079

Unique identifier UC11315775

Identifier EP67158.pdf (filename),usctheses-c39-274079 (legacy record id)

Legacy Identifier EP67158.pdf

Dmrecord 274079

Document Type Thesis

Format application/pdf (imt)

Rights MacDonald, J. T.

Type texts

Source University of Southern California (contributing entity), University of Southern California Dissertations and Theses (collection)

Access Conditions The author retains rights to his/her dissertation, thesis or other graduate work according to U.S. copyright law. Electronic access is being provided by the USC Libraries in agreement with the au...

Repository Name University of Southern California Digital Library

Repository Location USC Digital Library, University of Southern California, University Park Campus, Los Angeles, California 90089, USA

Linked assets

University of Southern California Dissertations and Theses