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he development of the mesonephros of the pink salmon, Oncorhynchus Gorbuscha (Walbum) Newstead, James Duncan 1956

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THE DEVELOPMENT OP THE MESONEPHROS OP THE PINK SALMON, ONCORHYNCHUS GORBUSCHA (WALBAUM). by James D. Newstead  A thesis submitted i n p a r t i a l fulfilment of the requirements f o r the degree of MASTER OP ARTS i n the Department of Zoology  We accept this thesis as conforming to the standard required from candidates f o r the degree of MASTER OP ARTS.  Members of the Department of Zoology  THE UNIVERSITY OP BRITISH COLUMBIA SEPTEMBER, 1956.  ABSTRACT  The development o f the k i d n e y o f the p i n k salmon was t r a c e d from the e a r l i e s t stages t o the y e a r - o l d f i n g e r ling.  The development o f the pronephros has been t r e a t e d  separately. The mesonephros develops from the i n t e r m e d i a t e c e l l mass o f the t w e l f t h t o t h e t h i r t y - n i n t h segments. D i f f e r e n t i a t i o n o f the unsegmented i n t e r m e d i a t e c e l l mass c l o s e l y f o l l o w s segmentation  o f the somites.  The v a s c u l a r  s t r a n d i s e s t a b l i s h e d beneath the hypochord by m i g r a t i o n o f c e l l s from the i n t e r m e d i a t e mesoderm o f e i t h e r s i d e .  A solid  c y l i n d r i c a l duct rudiment forms as a f o l d o f t h e d o r s o l a t e r a l aspect o f the mass on e i t h e r s i d e .  Mesonephrogenic m a t e r i a l  separates a s a narrow b r i d g e o f c e l l s p a s s i n g from one duct to  the o t h e r between the c a r d i n a l v e i n and the a o r t a .  mesonephrogenic rudiments appear as condensations c e l l s c l o s e s t t o the d u c t s .  Primary  o f the b r i d g e  Tubule rudiments a r e formed by  e l o n g a t i o n o f the condensations.  The f r e e end o f each t u b u l e  rudiment d i l a t e s t o form a t h i n w a l l e d Bowman c a p s u l e whose lumen l a t e r extends i n t o the tubule rudiment and f i n a l l y opens to  the duct.  A glomerulus  develops i n the Bowman capsule as a  s o l i d i n v a g i n a t i o n o f c e l l s from the v a s c u l a r s t r a n d . mass o f c e l l s subsequently  becomes v a s c u l a r i s e d .  This  Differentia-  t i o n o f the t u b u l e i n t o r e g i o n s f o l l o w s the appearance o f t h e glomerulus.  Secondary and subsequent t u b u l e g e n e r a t i o n s a r e  aa?e s i m i l a r l y formed but open i n t o p r e v i o u s l y formed primary tubules.  The i n t e r t u b u l a r spaces become f i l l e d w i t h myeloid  t i s s u e r i c h l y s u p p l i e d w i t h blood s i n u s e s . i s d e r i v e d from the v a s c u l a r s t r a n d .  The myeloid t i s s u e  Three p a i r s o f  corpuscles  o f S t a n n i u s appear e a r l y i n development as outgrowths o f segmental  duct e p i t h e l i u m .  Smaller unpaired corpuscles  the appear  l a t e r to make the t o t a l number twelve. The f u l l y d i f f e r e n t i a t e d nephron has f i v e  regions:-  1. a Bowman capsule and glomerulus 2. a short neck segment which opens d i r e c t l y from the Bowman capsule 3. a segment c h a r a c t e r i s e d by i t s  low columnar e p i t h e l i u m  and h i g h brush border 4. a segment with t a l l columnar c e l l s and a low brush border 5. a segment o f low simple columnar  cells.  S e v e r a l nephrons enter a c o l l e c t i n g t u b u l e . l e a s t three g e n e r a t i o n s  At  o f t u b u l e s open to the segmental  duct  by way o f a common c o l l e c t i n g duct formed by m o d i f i c a t i o n o f the primary c o l l e c t i n g  tubule.  I n v o l u t i o n o f the pronephros commences at about time the f r y n o r m a l l y enter the water a t the r i v e r mouth. t e n t i o n i n f r e s h water does not appear to a f f e c t onset o f i n v o l u t i o n .  Re-  the time o f  The p r o n e p h r i c r e g i o n becomes v e r y r i c h l y  permeated w i t h b l o o d s i n u s e s and may serve a s a blood organ i n o l d e r f i s h .  the  storage  I n the o l d e s t specimens s t u d i e d a r t e r i a l blood i s s u p p l i e d by p a i r e d a r t e r i o l e s i n the i n t e r s e g m e n t a l  septa.  Venous b l o o d from the t a i l passes d i r e c t l y i n t o a l a r g e median v e i n which passes a n t e r i a d i n the k i d n e y t i s s u e .  This v e i n  o r i g i n a t e s as the r i g h t p o s t c a r d i n a l v e i n i n the young embryo. The  l e f t p o s t c a r d i n a l does not develop p o s t e r i o r to the  nephros.  P o r t a l blood from the d o r s a l and  reaches the k i d n e y by way p o s t e r i o r r e g i o n two  pro-  v e n t r a l musculature  o f venules i n the myosepta.  In  the  l a r g e median a r t e r i e s ( a r t e r i a p r i m i t i v a  mesenterica) pass through the k i d n e y to the h i n d gut.  In  pronephric  artery  r e g i o n the l a r g e median c o e l i a c o - m e s e n t e r i c  passes through the kidney to supply  the  the v i s c e r a .  Glomerular counts made on samples taken from f r e s h and  s a l t water over a p e r i o d o f two  i n c r e a s e d osmotic c o n c e n t r a t i o n  months suggest t h a t  o f the marine environment  the e f f e c t o f r e t a r d i n g the r a t e o f development o f new i n a l o g a r i t h m i c r e l a t i o n to  the  length.  has  glomeruli  - i -  TABLE OF CONTENTS PAGE I. II. III. IV.  INTRODUCTION  1  ACKNOWLEDGEMENTS  2  LIFE HISTORY OP THE PINK SALMON  3  HISTORICAL REVIEW  3  A. The Study o f the Kidney i n P i s h  ....  4  B. Osmoregulation i n T e l e o s t s  6  1. Renal P h y s i o l o g y  7  2. C h l o r i d e Balance  9  3. E v o l u t i o n o f the t e l e o s t k i d n e y C. S t r u c t u r e o f the T e l e o s t Kidney  ....  10 12  1. Anatomy  12  2. H i s t o l o g y  14  D. The Development o f the Kidney 1. Nomenclature  18 18  2. The Development o f the Segmental Duct  20  3. Development o f the mesonephros i n Teleosts  22  4. The Development o f the C o r p u s c l e of S t a n n i u s V.  MATERIALS AND METHODS  25 27  A. M a t e r i a l s  27  B. Methods  27  1. F i x a t i o n (a) General h i s t o l o g i c a l d e t a i l .  27 28  - ii  -  (b) Mitochondria and brush borders 2. Embedding and Sectioning  28  3. Staining  29  (a) Bulk Staining  29  (b) General H i s t o l o g i c a l D e t a i l  29  (c) Mitochondria  29  4. Glomerular counts VI.  28  30  OBSERVATIONS  31  A. The Structure of the F u l l y Differentiated Kidney  31  1. Anatomy  31  2. Histology  32  B. The Chloride C e l l s of the G i l l s  ....  C. The Development of the Mesonephros  34 34  D. Comparison of Marine and Fresh Water Series  41  VII.  DISCUSSION  42  VIII.  CONCLUSIONS  44  LITERATURE CITED  45  IX.  - i i i -  L I S T OP  ILLUSTRATIONS PAGE  ANATOMY PLATE I 1-3  Figures  Venous d r a i n a g e mesonephros  of the t e l e o s t 31  PLATE I I <Figure  4  Figure 5  V e n t r a l view o f e n t i r e kidney p i n k salmon Transverse o f 65  o f the 32  s e c t i o n o f mesonephros  mms.  52  fingerling  PHYSIOLOGY TABLE Figure  6  Tabular representation of glomerular c o u n t s i n f r e s h and s a l t w a t e r s e r i e s  53  Graphical representation of r e l a t i o n s h i p o f number o f g l o m e r u l i t o l e n g t h i n f r e s h and s a l t w a t e r s e r i e s (Regression l i n e s )  54  Glomerulus and c a p s u l e o f f u l l y d i f f e r e n t i a t e d nephron  55  PLATE I I I Figure 7  HISTOLOGY PLATE I V Figure 8  Figure 9  Transverse  section of f i r s t  major 55  segment PLATE V Figure  10  Figure  11  Transverse segment Transverse segment  section stained section stained  o f f i r s t major f o r mitochondria o f second major f o r mitochondria  56 56  PLATE V I Figure  12  Transverse s e c t i o n o f second major segment  57  -  F i g u r e 13  i v-  Transverse segment  section of third  major  F i g u r e 14  Transverse duct  section of collecting  F i g u r e 15  Transverse  s e c t i o n o f segmental  57  PIATB V I I  58 duct  58  T r a n s v e r s e s e c t i o n o f 3-45 mm embryo t h r o u g h s i x t e e n t h segment t o show t r i a n g u l a r i n t e r m e d i a t e c e l l mass  59  T r a n s v e r s e s e c t i o n o f 4i4 mm embryo t h r o u g h t w e n t i e t h s o m i t e t o show s e p a r a t i o n o f duct rudiment and vascular strand  59  T r a n s v e r s e s e c t i o n o f 5.24 mm. embryo to show l o c a l i s a t i o n o f s t r u c t u r e s . d e r i v e d from the i n t e r m e d i a t e mesoderm  60  Transverse s e c t i o n o f l e f t lobe o f k i d n e y o f 16.66 mm embryo t o show mespnephric condensation  60  T r a n s v e r s e s e c t i o n o f k i d n e y o f 19.05 mm a l e v i n t o show e a r l y s t a g e i n d e v e l o p m e n t o f t h e Bowman c a p s u l e  61  EMBRYOLOGY PLATE V I I I Figure  Figure  16  17  PLATE IX Figure  Figure  18  19  PLATE X Figure  Figure  20  21  Transverse section o f kidney o f mm a l e v i n t o show i n c i p i e n t glomerular i n v a g i n a t i o n  19.8  61  PLATE X I Figure  Figure  22  23  S e c t i o n o f Bowman c a p s u l e a n d a v a s cular glomerular i n v a g i n a t i o n o f 19.8 mm a l e v i n  62  T r a n s v e r s e s e c t i o n o f k i d n e y o f 26.9 mm f r y t o show b e g i n n i n g v a s c u l a r i t y o f glomerulus  62  -  V  -  PLATE XII Figure 24  Figure 25  Transverse section of kidney of 28.58 mm f r y to show well-formed glomeruli  63  Transverse section of kidney of 29.78 mm f r y to show secondary Bowman capsule  63  Typical transverse section of kidney of 64 mm finger l i n g  64  Longitudinal section of 20.9 mm a l e v i n to show bladder l i k e distention of ducts  64  PLATE XIII Figure 26 Figure 27  CORPUSCLE OP STANNIUS PLATE XIV Figure 28  Figure 29  Longitudinal section of segmental duct i n 7.4 mm embryo to show rudiment of corpuscle of Stannius  65  Transverse section of right lobe of kidney i n 9.54 mm embryo to show separation of corpuscle from duct  65  Frontal section of corpuscle of Stannius i n 59 mm f i n g e r l i n g  66  PLATE XV Figure 30  INTERRENAL TISSUE PLATE XVI Figure 31  Transverse section of part of pronephros i n 100 mm f i n g e r l i n g to show arrangement of i n t e r r e n a l tissue  67  - 1 -  I . INTRODUCTION  I t has been suggested t h a t the k i d n e y i s p r i m a r i l y an osmoregulatory  organ and that i t s e x c r e t o r y r o l e i n h i g h e r  v e r t e b r a t e s i s s e c o n d a r i l y a c q u i r e d ( M a r s h a l l and Smith, 1930; Smith, 1951).  The secondary n a t u r e o f r e n a l e x c r e t i o n i s e s -  p e c i a l l y apparent i n the f i s h s i n c e the m a j o r i t y o f m e t a b o l i c wastes a r e l o s t e x t r a r e n a l l y through the g i l l membranes, (Smith, 1928). E v o l u t i o n a r y m o d i f i c a t i o n o f the t e l e o s t k i d n e y has i n v o l v e d p a r t i c u l a r l y the degree o f development o f t h e glome r u l a r apparatus.  I t was thought t h a t a study o f the k i d n e y o f  the anadromous P a c i f i c Salmon would prove o f i n t e r e s t i n t h i s respect.  Changes i n the kidney s t r u c t u r e d u r i n g development  were a n t i c i p a t e d such as would f a c i l i t a t e a d a p t a t i o n t o the d u a l osmotic environment  o f a migratory f i s h .  T h i s study  includes:1. A d e s c r i p t i v e account o f the development o f the mesonephros. 2. A study o f the s t r u c t u r e o f the f u l l y  differen-  t i a t e d mesonephric nephron. 3* A c o n s i d e r a t i o n o f some o f the p h y s i o l o g i c a l a s p e c t s o f osmoregulation viewed i n the l i g h t o f h i s t o l o g i c a l observations.  -  2  -  I I . ACKNOWLEDGEMENTS  I t i s a p l e a s u r e to acknowledge the a s s i s t a n c e o f Dr. P e t e r Pord o f the Zoology Department who  suggested  and  d i r e c t e d the p r e p a r a t i o n o f t h i s t h e s i s and whose a s s i s t a n c e w i t h t e c h n i c a l and photographic m a t e r i a l has proved i n v a l u a b l e . I am i n d e b t e d to Dr. I . McT. interest  i n t h i s work; Dr. W.  Cowan f o r h i s c o n t i n u e d  S. Hoar f o r c o l l e c t i o n  i a l s , many u s e f u l suggestions and a s s i s t a n c e w i t h the c a l a n a l y s i s ; Dr. J . Nash f o r a s s i s t a n c e w i t h the a n a l y s i s ; the d i r e c t o r and s t a f f of the P a c i f i c  of.materstatisti-  statistical  Biological  S t a t i o n f o r c o l l e c t i o n o f specimens; Miss Martha Nagai, Bernard Cox,  Mr.  Miss Janet Goodman and Miss Sachiko Tabata, a l l  o f whom a s s i s t e d i n the p r e p a r a t i o n of the l a r g e number o f s l i d e s r e q u i r e d f o r t h i s p r o j e c t ; my f e l l o w s t u d e n t s , i n p a r t i c u l a r Mrs. Mary H o l l a n d s , Mrs. K a t h e r i n e Newman and A r t h u r Houston, who  Mr.  have c o n t r i b u t e d s e v e r a l i d e a s d u r i n g the  course o f d i s c u s s i o n s .  I am p a r t i c u l a r l y i n d e b t e d t o my  wife  f o r her constant encouragement, a s s i s t a n c e w i t h the p r e p a r a t i o n o f photographs,  and f o r the f i v e l i n e drawings reproduced  here-  in. The work was  a s s i s t e d by two  $1600 g r a n t s g i v e n t o  Dr. P. Pord and by a b u r s a r y o f $800 awarded t o the author the N a t i o n a l Research C o u n c i l o f Canada.  by  - 3 -  III.  L I F E HISTORY OP THE PINK SALMON  The Onchorhyncids a r e anadromous f i s h e s which spend the  e a r l i e r p a r t o f t h e i r l i f e i n f r e s h water.  i n f r e s h water i n the e a r l y f a l l .  P i n k s spawn  The a l e v i n s h a t c h a f t e r  about s i x t y days i n c u b a t i o n i n g r a v e l beds a t stream head waters.  E a r l y the f o l l o w i n g s p r i n g the f r y m i g r a t e seaward.  M a t u r i t y i s a t t a i n e d a t the end o f two years when t h e a d u l t s r e t u r n t o the streams from which they emerged, spawn and d i e . P i n k s a r e the s m a l l e s t o f the P a c i f i c Salmon.  Their  young are r e a d i l y i d e n t i f i e d by the absence o f p a r r marks. A d u l t s a r e i d e n t i f i e d by:1. The number o f r a k e r s on the f i r s t  gill  arch  (26-34) 2. The number o f r a y s i n the a n a l f i n (13-17) 3* The number o f s c a l e s on t h e l a t e r a l  line  (170-229). Metamorphosed  males a r e f u r t h e r i d e n t i f i e d by t h e i r  v e r y l a r g e d o r s a l hump (Clemens and Wilbee, 1946). P i n k s were chosen f o r t h i s study because t h e i r r a p i d development makes them p a r t i c u l a r l y s u i t a b l e f o r developmental study*  IV. HISTORICAL REVIEW  A. The Study o f the.Kidney i n F i s h I n the l a t e n i n e t e e n t h and e a r l y t w e n t i e t h c e n t u r i e s emphasis was p l a c e d on d e s c r i p t i v e s t u d i e s o f r e n a l in fish.  The account by P r i c e (1897, 1904)  o f the  development development  o f the k i d n e y o f Bdellostoma s t o u t i i s one o f the c l a s s i c s t r a t i o n s o f the p r i m i t i v e mode o f r e n a l development. o f t e l e o s t kidney development  illu-  Accounts  were p u b l i s h e d by Rosenberg  (1867),  Emery (1882), Brook (1887), Mcintosh and P r i c e (1887), Swaen and Brachet (1899, 1901) and F e l i x (1906).  More r e c e n t s t u d i e s by  S t r o e r (1933), Maschkowzeff (1934), K i n d a h l (1938) and Moghe (1945) a r e a v a i l a b l e .  A b r i e f review was  compiled by Moghe  (1947). E m b r y o l o g i c a l s t u d i e s encouraged  s p e c u l a t i o n over the  e v o l u t i o n a r y o r i g i n o f the t h r e e v e r t e b r a t e k i d n e y r e g i o n s . Two major t h e o r i e s were proposed:1. that the pronephros, mesonephros and metanephros as d e s c r i b e d by Lankester (1877) were t h r e e d i s t i n c t nonhomologous organs developed i n c h r o n o l o g i c a l s u c c e s s i o n a l o n g the trunk.  T h i s t h e o r y r e c e i v e d s t r o n g support from German  authors o f the time, and p a r t i c u l a r l y from F e l i x  (1906).  2. t h a t the t h r e e r e g i o n s were s e r i a l l y homologous areas o f a s i n g l e p r i m i t i v e l y metameric a r c h i n e p h r o s which tended the e n t i r e l e n g t h o f the trunk.  Audige  ex-  (1910) r e f e r r e d  to the e n t i r e v e r t e b r a t e k i d n e y as a mononephros w h i l e P r i c e  - 5 -  (1904) a p p l i e d the term holonephros to the s e r i a l l y homologous p r o - and mesonephroi o f  Bdellostoma.  D e s c r i p t i v e work on lower c r a n i a t e s (Brauer, Hypogeophis) and l a t e r experimental Gruenwald, 1942-  s t u d i e s (Boyden, 1926  c h i c k ; Cambar, 1948-  Anura) confirmed  homologous nature o f the three kidney r e g i o n s .  F r a s e r (1950) reviewed  q u e s t i o n o f the homology o f the three v e r t e b r a t e kidney gions.  She  emphasised the a r t i f i c i a l i t y  and  the  Goodrich  (1930) d i s c u s s e d the e v o l u t i o n o f the h i g h e r v e r t e b r a t e from a p r i m i t i v e a r c h i n e p h r o s .  1902-  kidney the re-  o f the terms pro-,  meso- and metanephros which imply some d i s t i n c t i o n between the three r e g i o n s .  She  recommended t h a t the e n t i r e v e r t e b r a t e  ney be regarded  as a holonephros and  kid-  that the terms p r o - , meso-  and metanephros be r e t a i n e d f o r convenience of d e s c r i p t i o n o n l y . Gerard in  (1954) d i s t i n g u i s h e d between p r o - , meso- and  t e l e o s t s on the b a s i s o f t h e i r anatomical  structure.  and  metanephroi  histological  Audige (1910) had a l s o d e s c r i b e d the separate  kidney of some t e l e o s t s as metanephroi.  caudal  F r a s e r , however, p r e -  f e r r e d to r e s t r i c t the term to the a d u l t k i d n e y o f Amniotes. She p o i n t e d out t h a t p r o - and mesonephroi are d i s t i n g u i s h a b l e o n l y i n the young o f animals t h a t have a prolonged and  t h a t the d i f f e r e n c e was  a matter o f s i z e and  larval  life  complexity  only. More r e c e n t work on t e l e o s t kidneys  has been concen-  t r a t e d on the h i s t o l o g y and p h y s i o l o g y o f the nephrons. (1897, 1902  i n M a r s h a l l , 1929)  aglomerular  kidneys  Huot  f i r s t r e p o r t e d the e x i s t e n c e o f  i n c e r t a i n s p e c i e s o f lophobranchs.  Guitel  - 6 -  (1906) e s t a b l i s h e d the aglomerular c o n d i t i o n o f some Gobiescoides.  Audige  (1910) reviewed the then a v a i l a b l e m a t e r i a l  on the h i s t o l o g y o f the t e l e o s t k i d n e y and r e p o r t e d an a g l o merular kidney i n another lophobranch, Lophius p i s c a t o r i u s * F u r t h e r r e p o r t s o f aglomerular k i d n e y s have s i n c e been publ i s h e d by Verne (1922), Edwards (1929), Nash (1931), (1937, a ) .  Grafflin  .  The d i s c o v e r y o f the common occurrence o f the a g l o merular tubule i n t e l e o s t s s t i m u l a t e d i n t e r e s t i n the comparat i v e p h y s i o l o g y o f aglomerular and glomerular t u b u l e s .  The  demonstration ( G r a f f l i n , 1933) that the k i d n e y o f the daddy s c u l p i n (Myxocephalus  s c o r p i u s ) , i n which the t u b u l e s behaved  p h y s i o l o g i c a l l y as i f aglomerular, was i n f a c t g l o m e r u l a r was of p a r t i c u l a r i n t e r e s t .  S t u d i e s o f v a r i o u s s p e c i e s by Edwards  (1928), G r a f f l i n and E i s e n b e r g (1934) and Smith were r e p o r t e d . Attempts  have been made to c o r r e l a t e the degree o f  glomerular development with the osmotic c o n c e n t r a t i o n o f the environment 1933;  ( M a r s h a l l and Smith, 1930; Nash, 1931; Edwards,  G r a f f l i n , 1937, a ) . More r e c e n t l y a t t e n t i o n has turned  to the e v o l u t i o n a r y s i g n i f i c a n c e o f the v e r t e b r a t e k i d n e y and e s p e c i a l l y o f the g l o m e r u l a r apparatus (Smith,  1953).  B. Osmoregulation i n T e l e o s t s I t i s claimed t h a t , owing to i t s importance i n osmor e g u l a t i o n , e v o l u t i o n a r y m o d i f i c a t i o n o f the kidney had to p r e cede a l l o t h e r m o d i f i c a t i o n s o f the v e r t e b r a t e s (Smith, M a r s h a l l and Smith  (1930) and G r a f f l i n (1937, b) have  1953).  attempted  - 7 -  to  c o r r e l a t e the degree o f development o f v e r t e b r a t e nephrons,  and p a r t i c u l a r l y o f the g l o m e r u l i , t o the osmotic i n which the animals l i v e .  environment  Nash (1931) c a l c u l a t e d the a r e a o f  glomerular f i l t r a t i o n s u r f a c e p e r u n i t a r e a o f body s u r f a c e o f d i f f e r e n t f r e s h water and marine t e l e o s t s .  These authors a l l  concluded t h a t , i n g e n e r a l , f i s h from f r e s h water  environments  possessed many w e l l v a s c u l a r i s e d g l o m e r u l i w h i l e those from marine environments  tended t o have reduced areas o f g l o m e r u l a r  f i l t r a t i o n o r , i n extreme cases, t o be a g l o m e r u l a r .  I n the  e n t i r e l y aglomerular tubule a s s o c i a t e d s i m p l i f i c a t i o n o f t h e r e n a l t u b u l e s was i n v a r i a b l y found Smith  (Edwards, 1929; 1935).  (1953) c o n s i d e r e d the aglomerular c o n d i t i o n t o be t h e  n a t u r a l r e s u l t o f prolonged a d a p t a t i o n o f a s p e c i e s t o a marine environment. 1. Renal P h y s i o l o g y . Nussbaum (1887, i n Gerard, 1936) o b t a i n e d almost  total  c e s s a t i o n o f u r i n e flow when the glomerular neck o f the Amphib i a n nephron was l i g a t u r e d .  Bensley and Steens  (1928), however,  found continued e x c r e t i o n o f d i l u t e u r i n e a f t e r a s i m i l a r tion.  Gray (1930), as a r e s u l t o f Bensley and Steens'  opera-  experi-  ments and h i s own o b s e r v a t i o n t h a t the mesonephrie tubules i n T r i t o n v u l g a r i s f r e q u e n t l y were n o t open to the Bowman c a p s u l e , denied any e x c r e t o r y o r osmoregulatory  r o l e o f the glomerulus.  He p o s t u l a t e d a r o l e i n the r e s o r b t i o n o f d i s s o l v e d  substances  from the coelomic f l u i d by way o f the open p e r i t o n n e a l f u n n e l s . wearn and R i c h a r d s (1924), by chemical a n a l y s i s o f m i c r o p i p e t t e samples from the Bowman c a p s u l e , demonstrated  that  - 8 -  the glomerular f l u i d was o f approximately the same c o m p o s i t i o n as d e p r o t e i n a t e d b l o o d serum.  Their results strongly  Nussbaum's theory o f glomerular f i l t r a t i o n .  supported  Wearn and R i c h a r d s  (1924) and Gerard and C o r d i e r (1934) demonstrated  by the same  technique the occurrence o f t u b u l a r r e s o r b t i o n o f g l u c o s e and sodium c h l o r i d e i n the p r o x i m a l s e c t i o n o f the u r i n i f e r o u s tubule and o f w a t e r . i n the d i s t a l s e c t i o n o f the Amniote t u b u l e . Tubular r e s o r b t i o n o f water has not been demonstrated i n t e l e o s t s (Gerard, 1 9 3 6 ) . Gerard and C o r d i e r ( 1 9 3 4 ) , (1933) were a b l e to demonstrate  Gerard (1936) and S i n g e r  tubular excretion of v i t a l  i n tubules whose glomerular necks were l i g a t e d .  Their results  were supported by s t u d i e s o f G r a f f l i n and E i s e n b e r g (1934) fluorescent materials.  dyes  with  I n the aglomerular t u b u l e a l l e x c r e t i o n ,  i n c l u d i n g that o f water, must occur by a c t i v e processes i n t h e tubule w a l l s .  M a r s h a l l (1930)  showed t h a t , w i t h the e x c e p t i o n  of g l u c o s e which i s e x c r e t e d o n l y by the glomerulus, the a g l o merular t u b u l e o f marine t e l e o s t s e x c r e t e d a u r i n e o f a p p r o x i mately the same composition as that e x c r e t e d by g l o m e r u l a r tubules o f marine t e l e o s t s .  G r a f f l i n ( 1 9 3 7 , a) p o i n t e d out  that a l l g r a d a t i o n s from the glomerular to the c o m p l e t e l y aglomerular kidney were found i n animals from both marine and f r e s h water environments.  He observed.that the u n i f o r m l y brush-  bordered aglomerular t u b u l e was a b l e to perform most o f t h e f u n c t i o n s o f glomerular t u b u l e s . t h a t the a b i l i t y to adapt  G r a f f l i n ( 1 9 3 7 , b) suggested  to a wide range o f osmotic c o n d i t i o n s  was a f u n c t i o n o f e x t r a r e n a l f a c t o r s a s s o c i a t e d w i t h  osmoregula-  -  tion.  He a p p a r e n t l y  9  considered  -  c h l o r i d e r e g u l a t i o n by means  o f the c h l o r i d e c e l l s o f the g i l l s  to he o f more importance  i n osmoregulation than water r e g u l a t i o n by means o f the glomerulus.  Smith (1953) s t r o n g l y emphasised  the r o l e o f the  glomerulus i n osmoregulation. 2. C h l o r i d e Balance. The r o l e o f the g i l l s i n c h l o r i d e uptake i n f r e s h water t e l e o s t s has been demonstrated (Keys, 1931). (1932) c l a i m e d that c h l o r i d e uptake occurred gill  epithelium.  described  Bevelander  through the  general  Burns and Copeland (1950) and Copeland (1948)  s p e c i a l i s e d c e l l s i n the g i l l  epithelium  and w e l l  v a s c u l a r i s e d areas o f the o r a l e p i t h e l i u m which showed a h i g h degree o f e o s i n o p h i l i a and s t a i n e d s t r o n g l y w i t h a s i l v e r nique.  The authors concluded that these were c h l o r i d e  tech-  secreting  c e l l s s i m i l a r to the g a s t r i c p a r i e t a l c e l l s and that they were responsible  f o r the c h l o r i d e uptake observed by Keys.  Krogh  (1939) was unable to d e t e c t measurable c h l o r i d e uptake i n the f r e s h water adapted e e l but observed c h l o r i d e a b s o r b t i o n other t e l e o s t s i n c l u d i n g Pundulus.  Wikgren  in  (1953) found i n  the lamprey a dynamic e q u i l i b r i u m between a c t i v e uptake and d i f f u s i v e l o s s o f s a l t s from the g i l l s i n f r e s h water  acclima-  tisation. Marine t e l e o s t s c o n s t a n t l y water l o s t by exosmosis.  d r i n k s e a water to r e p l a c e  The contained  c h l o r i d e , which i s ab-  sorbed from the i n t e s t i n e w i t h the water, i s removed from the blood by a c t i v e e x c r e t i o n i n the g i l l r e g i o n  (Black,  1951).  Copeland (1950) was a b l e to demonstrate r e v e r s a l o f f u n c t i o n i n  - 10 -  the c h l o r i d e c e l l s o f ffundulus when t r a n s f e r r e d from f r e s h to s a l t water and back. sal i n Anguilla. in  Getman  (1950)  described  Both authors d e s c r i b e d  a similar  rever-  c y t o l o g i c a l changes  the c h l o r i d e c e l l s which accompanied the r e v e r s a l o f f u n c -  tion.  Keys and Wilmer  (1932)  and B l a c k  (1951)  emphasised  the  importance o f r e v e r s i b i l i t y o f c h l o r i d e c e l l f u n c t i o n to t h e osmoregulation o f e u r y h a l i n e  teleosts.  B l a c k suggested a  q u a n t i t a t i v e r e l a t i o n s h i p between degree o f e u r y h a l i n i t y and rate o f reversal of chloride c e l l function. and Keys species  (1933)  Getman  (1950)  observed that c h l o r i d e s e c r e t i o n began i n  the  s t u d i e d o n l y a f t e r the body f l u i d s had become r a t h e r  concentrated.  These authors c o n s i d e r e d that a d a p t a b i l i t y o f  the c h l o r i d e c e l l s was p r i m a r i l y r e s p o n s i b l e  f o r the mainten-  ance o f homeostasis i n t e l e o s t s . 3. E v o l u t i o n o f the t e l e o s t k i d n e y . I t i s g e n e r a l l y accepted (Smith, M o r g u l i s , 1952)  1953r  that f i s h e s evolved from the salt  Florkin  and  protovertebrate  ft****  stock o n l y a f t e r m i g r a t i o n from figesh to oalrfc water had been initiated.  I n f r e s h water the p r o t o v e r t e b r a t e s  to osmotic f l o o d i n g owing t o t h e i r r e l a t i v e l y body f l u i d s .  were  subject  concentrated  To m a i n t a i n homeostasis i t was n e c e s s a r y :  a) t o exclude from the body as much water a s p o s s i ble.  T h i s was achieved w i t h the e v o l u t i o n o f the  armour o f the Ostracoderms ( P l o r k i n and M o r g u l i s , b) to e l i m i n a t e  impervious  1952).  osmotic water which p e n e t r a t e d t h e  permeable g i l l and o r a l membranes.  T h i s was made p o s s i b l e by  - l i -  the i n t r o d u c t i o n o f a g l o m e r u l a r u l t r a f i l t e r  i n t o the aglomer-  u l a r prot©vertebrate nephridium (Smith, 1953). f u n c t i o n o f the n e p h r i d i a l e p i t h e l i u m , for  The a b s o r b t i v e  which was r e s p o n s i b l e  r e s o r b t i o n o f u s e f u l s o l u t e s from the coelomic f l u i d , was  r e t a i n e d i n the glomerular t u b u l e o f f i s h e s f o r the r e s o r b t i o n o f m a t e r i a l s from the g l o m e r u l a r f i l t r a t e .  A high rate of  glomerular f i l t r a t i o n together w i t h a c t i v e t u b u l a r enabled the e a r l y f i s h to excrete  resorbtion  osmotic water w h i l e r e t a i n i n g  essential chloride. c) to o b t a i n c h l o r i d e from the environment an adverse d i f f u s i o n g r a d i e n t . chloride absorbtion  T h i s was achieved by a c t i v e  i n the g i l l and o r a l r e g i o n s  pense o f r e s p i r a t o r y energy.  against  a t t h e ex-  C o n s i d e r a b l e amounts o f c h l o r i d e  were a l s o obtained from t h e f o o d . T h e i r l a t e r r e t u r n to s e a water exposed t h e ancest r a l f i s h t o osmotic d e h y d r a t i o n . t r a t i o n became a l i a b i l i t y .  Excessive  glomerular  fil-  The e v o l u t i o n o f the u r e a r e t e n -  t i o n h a b i t u s made unnecessary any m o d i f i c a t i o n o f t h e elasmobranch nephron (Smith, 1936). l o n g term m o d i f i c a t i o n s an organ f o r m e r l y  T e l e o s t s , however, underwent  which made u s e f u l i n water  s p e c i a l i s e d f o r water e l i m i n a t i o n .  t i o n i n v o l v e d p r i m a r i l y r e d u c t i o n o f the glomerular rate.  conservation Modificafiltration  With the reduced u r i n e f l o w a c t i v e e x c r e t i o n by the  tubular epithelium  became o f more s i g n i f i c a n c e .  Reversibility  o f the c h l o r i d e c e l l f u n c t i o n presumably made p o s s i b l e the i n i t i a l c o l o n i s a t i o n o f t h e new environment.  - 12  -  C. S t r u c t u r e o f the T e l e o s t Kidney 1. Anatomy. The d e f i n i t i v e kidney o f most t e l e o s t s i s the mesonephros.  Audige  (1910) showed the presence of a h i n d  k i d n e y w i t h a d i s t i n c t duct system i n s e v e r a l s p e c i e s .  Some  s p e c i e s possessed f u n c t i o n a l t u b u l e s i n both mid and h i n d k i d neys w h i l e some possessed t u b u l e s o n l y i n the h i n d k i d n e y . the a d u l t t e l e o s t the f o r e kidney was ate.  In  u s u a l l y e n t i r e l y degener-  Tubules and glomerulus were r e p l a c e d by myeloid  tissue.  T e l e o s t s are more v a r i e d i n t h e i r r e n a l anatomy and h i s t o l o g y than any o t h e r taxonomic group. c r i b e d two  Audige  (1910) des-  d i s t i n c t forms o f a d u l t k i d n e y : 1) mononephric kidneys such as were found i n the  genus Cyclothone  (Owen, 1938).  A s i n g l e v e r y l a r g e median  glomerulus p r o j e c t e d on e i t h e r s i d e i n t o a n e p h r i c chamber from which a s i n g l e u r i n i f e r o u s t u b u l e l e d i n t o Gerard  the duct o f t h a t  (1954) c o n s i d e r e d the mononephric nephron to be a  ous c o n d i t i o n i n Cyclothone.  side.  neoten-  A s i m i l a r mononephric f o r e k i d n e y  was found i n a l l s p e c i e s o f a d u l t Lepadogaster. c o n j u n c t i o n w i t h a f u n c t i o n a l p o l y n e p h r i c mid  sometimes i n  kidney.  2) p o l y n e p h r i c k i d n e y s , l o c a t e d i n the mid o r h i n d kidney r e g i o n .  They c o n s i s t e d of many nephrons e n t e r i n g a p a i r  o f ducts which emptied i n t o the c l o a c a . Gerard (1953) found f o u r f a i r l y r e p r e s e n t a t i v e arrangements o f the kidney i n t e l e o s t s : -  - 13 -  1) fore and mid kidneys i n the adult. kidney might (a) be mononephric (e.g.  The fore  Lepadogaster),  no nephrons, but consist of a mass of myeloid tissue  (b) have (e.g.  Barbus) (c) possess both a t y p i c a l mononephric unit and a group of glomerular nephrons (e.g. Fundulus) 2) fore kidney degenerate; mid and hind (post  cloaca!)  kidneys both with functional tubules (e.g. Auguilla) 3) fore kidney a mass of myeloid t i s s u e .  Mid and  hind kidneys with functional tubules (e.g. Apola) 4) mid kidney only i n the adult (e.g. Lophius, Opsanus). Gerard found the hind kidney to be t y p i c a l l y a compact mass which was located i n a post cloacal diverticulum of the coelom.  A r t e r i a l c i r c u l a t i o n was supplied by one or two  medial*.arteries.  In those forms where the hind kidney was pre-  sent but aglomerular no a r t e r i a l c i r c u l a t i o n was found (Hippocampus - Edwards, 1929).  A p a i r of ducts which developed as  buds of the Wolffian ducts drained the hind kidney.  The mid  kidney was often an elongate organ, tapering posteriad, which extended most of the length of the trunk. no outward sign of metamerism.  There was generally  The separation of r i g h t and  l e f t kidneys was also obscured but Edwards (1928) showed by i n j e c t i o n of the Wolffian ducts that there was no connection between the nephrons of the two sides i n the species studied by him.  The kidney was well supplied with wide venous sinuses  around which myeloid tissue was packed.  - 14 -  Audige (1910) d e s c r i b e d three p r i n c i p a l arrangements of  the venous c i r c u l a t i o n i n the mesonephros:1) blood from the caudal v e i n reached the kidney  way  o f a t r u e p o r t a l system.  developed  by  The r i g h t post c a r d i n a l was b e t t e r  than the l e f t but both p e r s i s t e d and extended f a r  p o s t e r i a d i n the k i d n e y to r e c e i v e b l o o d from the r e n a l p o r t a l veins (figure 1). 2) a p o r t a l v e i n e x i s t e d o n l y on the l e f t w h i l e r i g h t post c a r d i n a l was  the  continuous w i t h the caudal v e i n ( f i g u r e  2). 3) the r i g h t post c a r d i n a l alone p e r s i s t e d t o the adult stage i n the trunk r e g i o n .  I t was  continuous w i t h the  caudal v e i n and passed a n t e r i a d through the mesonephros. the p r o n e p h r i c r e g i o n the l e f t branched  c a r d i n a l e x i s t e d as a s h o r t  v e s s e l r e c e i v i n g blood from the s i n u s o i d s ( f i g u r e 3 ) .  A type o f p o r t a l system was  formed by the i n t e r s e g m e n t a l v e i n s  from the d o r s a l and v e n t r a l body w a l l which d r a i n e d i n t o kidney s i n u s o i d s . way  In  the  A r t e r i a l blood reached the mesonephros by  o f a r t e r i o l e s i n the i n t e r s e g m e n t a l c o n n e c t i v e t i s s u e .  2. H i s t o l o g y . S t r u c t u r e o f the nephron. L i Ko^e Viltner  Tchang (1923), M a r s h a l l and Smith (1930) and  (1935) r e p o r t e d the presence o f an a v a s c u l a r core o f  dense s y n c i t i a l t i s s u e i n the a v i a n glomerulus.  Nash (1931)  r e p o r t e d m o d i f i c a t i o n s i n the g l o m e r u l i o f marine t e l e o s t s  - 15 -  r e l a t i v e to those of fresh water species.  He observed i n par-  t i c u l a r reduction of v a s c u l a r i t y and the presence of  quantities  of fibrous material scattered through the glomerular t i s s u e . He interpreted the presence of quantities of fibrous matter as an i n d i c a t i o n of degeneracy.  G r a f f l i n (1929) observed hyaline  degeneration i n the centre of the pseudo-glomerulus of Lophius piscatorius.  The core lacked c a p i l l a r i e s , c e l l s ,  or f i b r e s ,  and i n cases of extreme degeneration was f i l l e d with granules. A r t e r i a l vascularisation was l i m i t e d to the peripheral areas of the pseudo-glomeruli.  In the common s c u l p i n (Myxocephalus  octodecemspinosus) G r a f f l i n (1937, c) found considerable v a r i a b i l i t y i n s i z e , degree of v a s c u l a r i t y , degree of l o b u l a t i o n and c e l l u l a r i t y of the glomerulus.  Scattered groups of  connective  tissue c e l l s were common, but no fibrous core was reported.  In  a few glomeruli the centre was occupied by a large f l u i d - f i l l e d cyst.  The glomeruli of the daddy sculpin (Myxocephalus scorpius)  (Grafflin,  1933)  showed more advanced degeneration.  of the Bowman capsules was common.  Four stages of  Distortion degeneration  were found:1) glomeruli consisted of a few n u c l e i i n a large mass of s y n c i t i a l tissue 2) glomeruli were without nuclei or f i b r e s and cons i s t e d of a mass of green caseous matter 3) glomeruli consisted of a mass of  structureless  membranes 4) the peripheral region of the glomerulus was  - 16 -  v a s c u l a r but the c e n t r e was f i l l e d by a l a r g e c y s t . In  a l l cases the g l o m e r u l i were connected  to t u b u l e s  whose necks were c l o s e d . G r a f f l i n (1937, d) observed g r e a t v a r i a b i l i t y i n t h e s i z e and degree o f l o b u l a t i o n o f the g l o m e r u l i o f the c a t a d r o mous common e e l ( A n g u i l l a r o s t r a t a ) .  In larger glomeruli  a v a s c u l a r c o n n e c t i v e t i s s u e masses o c c u p i e d some o f the l o b u l e s but no c e n t r a l cores were found.  Some g l o m e r u l i were found to  possess two d i s t i n c t v a s c u l a r t u f t s , each w i t h an a f f e r e n t and an e f f e r e n t a r t e r i o l e . n e p h r i c glomerulus  My o b s e r v a t i o n s o f the i n v o l u t i n g p r o -  o f 0 . gorbuscha  i n d i c a t e d that g l o m e r u l a r  degeneration was accompanied by i n v a s i o n o f the glomerulus by f i b r o u s connective  tissue.  Edwards (1937) found no j u x t a g l o m e r u l a r s t r u c t u r e s i n association with teleost glomeruli.  I n other v e r t e b r a t e s he  found e i t h e r a plaque, a d i s c , o r both. Edwards and S c h n i t t e r (1933) remarked that a t u b u l e segment c y t o l o g i c a l l y e q u i v a l e n t to the brush bordered  proximal  c o n v o l u t i o n o f the mammalian nephron was common t o t h e nephrons of  a l l vertebrates.  The aglomerular t u b u l e s o f some marine  t e l e o s t s were u n i f o r m l y brush bordered over t h e i r whole l e n g t h . One  o r more such t u b u l e s entered a common c o l l e c t i n g d u c t u l e .  Glomerular  t u b u l e s were o f v a r i a b l e s t r u c t u r e but were always  more complex than were aglomerular t u b u l e s . 1935)  Edwards (1933,  found up to f i v e segments i n the g l o m e r u l a r t e l e o s t t u b u l e : 1) the glomerular neck ( f i r s t minor segment).  The  squamous e p i t h e l i a l c e l l s o f the Bowman capsule changed g r a d -  - 17 -  u a l l y to the columnar l a t e r a l l y compressed c e l l s of the neck segment.  I f the tubules of a given species possessed an i n t e r -  mediate segment, the glomerular neck was i n v a r i a b l y c i l i a t e d ; i f not, i t was never c i l i a t e d . 2) the f i r s t major segment.  According to Edwards,  the c e l l s of t h i s segment were low columnar with a high w e l l defined brush border at least one t h i r d the height of the Their large e l l i p s o i d a l n u c l e i were basally located. plasm was f a i n t l y b a s o p h i l i c .  cells.  The cyto-  Numerous short rod l i k e mito-  chondria were concentrated toward the base of the c e l l s (Cowdry and C o v e l l , 1928). 3) the second major segment, which usually preceded the second minor segment.  T a l l columnar c e l l s with a low brush  border characterised t h i s region of the tubule. nuclei were basally located.  The round  Thread-like mitochondria were  distributed throughout the eosinophilic cytoplasm.  The mito-  chondria - cytoplasmic r a t i o was considerably lower i n t h i s segment (Cowdry and C o v e l l , 1928). 4) the second minor segment, or intermediate segment, was absent i n many species.  I f present, i t was i n v a r i a b l y of  c i l i a t e d t a l l columnar l a t e r a l l y compressed  cells.  5) the t h i r d major segment resembled the d i s t a l convolution of the mammalian tubule.  Its c e l l s were low columnar  with central nuclei and basophilic cytoplasm. region of the tubule was large.  The lumen of t h i s  - 18 -  Several tubules drained into a c o l l e c t i n g tfoufabule characterised by i t s columnar epithelium and very narrow lumen which increased i n diameter further along i t s  length.  Kempton (194-3) reported extreme elongation of the neck segment of the elasmobranch nephron. boidal with depressed basal n u c l e i . heavily c i l i a t e d .  Its c e l l s were cu-  The entire length was  It was believed that this s p e c i a l segment  was related to the resorbtion of urea from the urine. G-rafflin (1937, c) observed numerous c i l i a t e d c e l l s i n the second major segment of the nephron of the common s c u l p i n . Edwards (1929) found the entire length of the f i r s t and second major segments i n Muraena was uniformly c i l i a t e d .  He noted that  only the f i r s t major segment was common to a l l vertebrate nephrons and that t h i s was the only segment occurring i n aglomerular tubules.  He concluded that the glomerulus acted as the  stimulus for c y t o l o g i c a l d i f f e r e n t i a t i o n of the tubule but that functional differences were not attributable to the presence of the glomerulus.  D. The Development of the Kidney 1. Nomenclature. The entire kidney develops from the mesodermal materi a l located between the somite and the l a t e r a l p l a t e .  This  material was c a l l e d by Balfour (1877) the intermediate c e l l mass. P r i m i t i v e l y , the mass becomes segmented and arranged into somatic and splanchnic layers.  The narrow cavity between the layers  is  continuous v e n t r a l l y with the splanchnocoele and d o r s a l l y with  - 19 -  the myocoele.  The a r e a n e a r e s t the somite d i l a t e s t o form a  conspicuous eoelomic chamber o r nephrotome which e n c l o s e s a nephrocoele (Ruckert, 1888).  The c o n n e c t i o n w i t h the myocoele  i s l o s t v e r y e a r l y i n development.  The c o n n e c t i o n w i t h the  splanchnocoele i s reduced to a narrow p e r i t o n e a l f u n n e l and p e r i t o n e a l c a n a l by ingrowth o f c e l l s from e i t h e r end o f the nephrotome.  D i l a t i o n o f the nephrotome and f l a t t e n i n g o f i t s  e p i t h e l i u m forms a Bowman c a p s u l e . E a r l y i n the development o f the k i d n e y amoeboid  cells  migrate from the medial s u r f a c e o f the i n t e r m e d i a t e c e l l mass, and l a t e r o f the nephrotomes, a l o n g the e n t i r e trunk. c e l l s form beneath the hypochord  These  a d a r k l y s t a i n i n g mass which  Maschkowzeff (1929) c a l l e d the venous s t r a n d .  The s t r a n d g i v e s  r i s e not o n l y to the c a r d i n a l v e i n s , as was supposed  by Masch-  kowzeff, but a l s o to the a o r t a , g l o m e r u l i , myeloid t i s s u e and c e r t a i n s k e l e t a l elements  ( F r a s e r , 1950).  A c c o r d i n g l y the s t r a n d  i s r e f e r r e d to as the v a s c u l a r s t r a n d ("masse v a s c u l a i r e " ) as suggested by Swaen and Brachet  (1902).  The medial w a l l o f the Bowman capsule i s invaded by a v a s c u l a r t u f t , t h e glomerulus, which i s s u p p l i e d w i t h a r t e r i a l blood from the a o r t a . A s o l i d outgrowth  (rudiment o f the u r i n i f e r o u s t u b u l e )  o f the d o r s a l w a l l o f the e a r l y nephrotome becomes e l o n g a t e d , c a v i t a t e d and opens from t h e nephrocoele by a nephrostome. Goodrich (1930) p r e f e r s the term nephrocoelostome  to d i f f e r e n t -  i a t e i t from t h e nephrostome o f Amphibian p r o n e p h r o i but F r a s e r (1950) has p o i n t e d out the homology o f the s t r u c t u r e i n Amphibia  - 20 -  and other vertebrates.  Fraser describes two types of glomerulus:  1) Internal glomerulus, entirely enclosed in a Bowman capsule. 2) External glomerulus which projects into a diverticulum of the general coelom. glomus to this structure.  McEwen (1949) applies the term  Fraser's nomenclature is preferred  since i t emphasises the homology of the two types of glomerulus. Emery (1882) described the appearance of mesonephrogenic condensations in a narrow band of cells passing from one segmental duct to the other between the aorta and the cardinal. Hall (1904) found a similar condition in Rana.  Hall called the  bridge mesonephric blastema and the condensations blastulae. Moghe (1945) referred to a similar bridge of cells in Thynnichthys as bridge cells and to the mesonephric rudiments simply as condensations. 2. The development of the segmental duct. In the primitive animals Bdellostoma (Price, 1899, 1904) and Hypogeophis (Brauer, 1897) each of the segmentally placed uriniferous tubules grows posteriorly to unite with the next tubule.  A duet is formed along the length of the trunk by  the union of the tubules.  The free end of the most posterior  tubule grows actively ventrad to enter the cloaca.  It is be-  lieved (Goodrich, 1930) that the segmental ducts of the early vertebrates were similarly formed by union of segmental contributions. The segmental duct is formed in Elasmobranchs by  - 21 -  active growth from the anterior region.  A small knob of tissue  develops from the intermediate mesoderm just posterior to the heart.  From the knob a s o l i d rod of c e l l s grows posteriad to  the cloaca.  The rod l a t e r becomes tubulated and opens at the  anterior end to the coelom.  Mesonephric rudiments form and  l a t e r open to the duct (Bates, 1914). Maschkowzeff (1926) found the segmental duct of Acipenser (Chondrostei) to be formed as a thickening of the dorsal aspect of the intermediate c e l l mass i n each segment. The thickening l a t e r becomes folded off as a s o l i d rod which i n turn acquires a lumen.  Union with the cloaca i s  accomplished  by active growth. In Polypterus (Kerr, 1907 i n Moghe, 1947)  the  segmental  n  duct i s formed posterior to the pronephros by bodily conversion and fusion of segmental  nephrotomes".  Mcintosh and Price (1887) and Brook (1887) claimed that the segmental duct of teleosts i s formed as a thickening of the ectoderm from which the duct l a t e r becomes separated. Swaen and Brachet (1899, 1901) described a d o r s o - l a t e r a l cleavage of the intermediate c e l l mass which sets aside a s o l i d mass of t i s s u e .  Later, the mass of tissue acquires a lumen  which the authors claimed i s l i n e d p a r t l y with somatic and p a r t l y with splanchnic mesoderm and therefore i s the morphological equivalent of the coelom.  Swaen and Brachet suggested that  tubule and pronephric chamber differentiate end of the duct.  the  from the anterior  Stroer (1930) found that the duct i n Perca  forms as a f o l d of the dorsolateral aspect of the intermediate  - 22 -  mesoderm i n each segment a f t e r f o r m a t i o n o f the p r o n e p h r i c chamber. Recent experimental p h i b i a ; Cambar, 1948,  s t u d i e s (O'Connor, 1935  Gruenwald, 1942,  on  Am-  on b i r d s ) show that the  duct i s formed i n l e s s p r i m i t i v e animals by union o f the f r e e ends o f the f o r e kidney rudiments and a c t i v e growth p o s t e r i a d of  the l a s t rudiment to the c l o a c a .  There are no  segmental c o n t r i b u t i o n s o f m a t e r i a l .  The  significant  supposed segmental  o r i g i n of the t e l e o s t segmental duct i s y e t to be t e s t e d e x p e r i mentally. 3. Development o f the mesonephros i n t e l e o s t s . The i n t e r m e d i a t e c e l l mass appears e a r l y i n development as a l o o s e t r i a n g u l a r mass o f t i s s u e l a t e r a l to the and medial  to the l a t e r a l p l a t e ( B r a c h e t , 1935  - Salmo t r u t t a ) .  S e p a r a t i o n from the somite occurs v e r y e a r l y but the w i t h the l a t e r a l p l a t e i s r e t a i n e d l o n g e r .  Formation  connection of the  v a s c u l a r s t r a n d commences soon a f t e r s e p a r a t i o n from the Swaen and Brachet  cell  Masehkowzeff (1929) showed t h e i r c o n c l u s i o n  to be based on erroneous  i n t e r p r e t a t i o n of the prolonged a t t a c h -  ment o f i n t e r m e d i a t e c e l l mass and l a t e r a l p l a t e .  Nephrotomes  are s a i d not to appear d u r i n g the development o f the kidney  somites.  (1899) d e s c r i b e d the appearance o f the v a s -  c u l a r s t r a n d p r i o r to d i f f e r e n t i a t i o n o f the i n t e r m e d i a t e mass i n S. t r u t t a .  somites  ( F r a s e r , 1950).  teleost  F e l i x (1897), however, d e s c r i b e d the  appearance o f segmental s t r u c t u r e s i n the p r o n e p h r i c r e g i o n o f S.  t r u t t a and S t r o e r (1932) r e f e r r e d to a "nephrotome" i n P e r c a . Nussbaum ( i n Moghe, 1940)  d e s c r i b e d the o r i g i n o f  - 23 -  mesonephric Felix  rudiments as outgrowths  o f the duct e p i t h e l i u m .  (1906) d e s c r i b e d a s i m i l a r o r i g i n o f primary aglomerular  mesonephric  t u b u l e s i n S. t r u t t a .  He claimed that the primary  tubules are l a t e r r e p l a c e d by permanent t u b u l e s of unknown origin.  Maschkowzeff (1934) t r a c e d the o r i g i n of the permanent  mesonephric  t u b u l e s to condensations of mesonephric  developed from the i n t e r m e d i a t e c e l l mass i n  mesenchyme  S. t r u t t a .  He d i d  not comment on F e l i x ' s d e s c r i p t i o n o f the o r i g i n o f primary tubules. cal  Brachet (1935) observed the appearance  of hemispheri-  e v a g i n a t i o n s o f the duct e p i t h e l i u m s i m i l a r to those des-  c r i b e d by F e l i x .  He t r a c e d t h e i r f u r t h e r development and found  that they s e p a r a t e from the duct, round up and migrate out i n t o the kidney t i s s u e .  In l a t e r stages they become l o b u l a t e d and  well vascularised.  He  c o n s i d e r e d them to be the rudimentary  corpuscles of Stannius. Rosenberg (1867) thought that the mesonephric  rudi-  ments are a g g r e g a t i o n s o f c e l l s e i t h e r from the v e n t r a l w a l l of  the a o r t a or from " p a r i e t a l " c e l l s l o c a t e d i n the w a l l s o f  the c a r d i n a l v e i n s . mesonephric  Emery (1882) d e s c r i b e d the appearance  condensations i n a b r i d g e o f c e l l s which passes  of be-  tween the a o r t a and c a r d i n a l v e i n from one duct to the o t h e r . He thought that the b r i d g e o r i g i n a t e d from the p e r i t o n e a l epithelium. H a l l (1904) found i n Rana a s i m i l a r b r i d g e o f mesonephrogenic c e l l s i n which mesonephric densation. to  rudiments formed by  con-  He t r a c e d the o r i g i n o f the mesonephrogenic b r i d g e  the i n t e r m e d i a t e c e l l mass.  Gray (1930) v e r i f i e d h i s con-  - 24 -  elusion i n a different species of Rana. Moghe described the appearance as condensations of the mesonephrogenic bridge c e l l s of hemispherical caps of closely applied to the ducts.  cells  He traced the o r i g i n of the bridge  c e l l s to the region of the intermediate c e l l mass closest to the somite.  This i s the region from which nephrotomes develop i n  those species where mesonephric nephrotomes form (Fraser, 1950). Brachet (1935) and F e l i x (1906) described the elaboration of the mesonephrogenic condensations i n Salmo t r u t t a .  The hemispherical  condensations become closely applied to the ducts. d i v i s i o n r e s u l t s i n elongation of the rudiments.  Rapid c e l l The end of the  newly formed cylinder of c e l l s penetrates the wall of the segmental duct and, after tubulation, becomes open to the duct. The free end of the tubule d i l a t e s to form a Bowman capsule which i s l a t e r invaginated by a glomerulus.  Moghe described a  s i m i l a r s i t u a t i o n i n Thynniehthys, but reported that the glomerulus and capsule are formed independently of the tubules as condensations of the c e l l s of the mesonephrogenic bridge.  The  condensations grow to large s o l i d spherical masses which become attached to the tubule and subsequently become hollow.  The  glomerulus appears as a s o l i d mass of basophilic c e l l s which i n vaginateSthe wall of the Bowman capsule.  Gray (1930) found  that the tubules i n Rana form by elongation and tubulation of the mesonephrogenic condensations.  As a r e s u l t of more rapid  d i v i s i o n of the l a t e r a l l y located c e l l s , the developing tubules c u r l ventro-medially around the duct.  At the free end, d i v i s i o n  occurs exclusively i n a tangential plane and r e s u l t s i n the  - 25 -  formation of a large t h i n walled capsule.  The area of the wall  opposite the tubule mouth i s invaginated by a s o l i d mass of c e l l s derived from the mesonephric bridge.  Vascularisation of  the mass by an a r t e r i a l shoot completes the development of the glomerulus.  Andige (1910) remarked that a l l mesonephric tubules  are i n i t i a l l y aglomerular.  F e l i x (1906) and Moghe (1945) ob-  served the formation of secondary and t e r t i a r y mesonephric tubules which open into the connecting segment of previously formed tubules.  Gray (1953) pointed out that no secondary  tubules can form i n Triton since the entire mesonephrogenic mass differentiates  into primary v e s i c l e s .  In t h i s species, rudiments  are formed by fragmentation of the mass rather than by condensation of c e l l s as i n Rana.  Secondary generations are formed i n  Rana. Peritoneal funnels, which Gray reported i n the mesonephros of Rana and T r i t o n , were not observed by F e l i x , Brachet, Moghe or Maschkowzeff i n the mesonephroi of t e l e o s t s . 4. The development of the Corpuscle of Stannius. Garrett (1942) reviewed the available l i t e r a t u r e on the development and structure of the corpuscle of Stannius. When o r i g i n a l l y reported by Stannius the corpuscle was thought to be the homologue of the mammalian adrenal cortex  (interrenal).  More recent research located both suprarenal and i n t e r r e n a l bodies i n the anterior region of the teleost kidney.  It i s now  believed that the corpuscle of Stannius i s unrelated to the adrenal system.  I t s function i s unknown but apparently endocrine.  The corpuscles have been reported only i n teleosts and holostean  - 26 -  ganoids.  Up to f i f t y pairs are reported i n Holostei; one  i n most t e l e o s t s .  pair  In the c a t f i s h , a single corpuscle i s formed  by fusion of the two members of a p a i r .  Salmonids commonly  possess three or four p a i r s . Garrett  (1942) discussed the o r i g i n of the corpuscles  of Stannius i n Amia calva as paired outgrowths of the duct epithelium  p r i o r to the appearance of mesonephric condensations.  Their c e l l s are early d i f f e r e n t i a t e d from those of the duct by their very palely stained cytoplasm.  The outgrowths appear i n  regions where the cardinal sinusoids are c l o s e l y applied to  the  ducts. The rudiments increase greatly i n size and acquire a s l i t l i k e lumen.  At this stage they have the appearance of early  mesonephric rudiments ( c f . F e l i x , 1906). separation  At about the time of  from the ducts fibrous trabeculae carrying a r t e r i a l  c a p i l l a r i e s invade the tissue of the corpuscles and them to lobules.  subdivide  F i n a l l y the developing corpuscles are forced  away from the ducts into the kidney tissue where they increase considerably  i n s i z e , v a s c u l a r i t y and degree of lobulation.  - 27 -  V. MATERIALS AND METHODS  A. Materials Eggs of the pink salmon (Onchorhynous gorbuscha) were collected at Cultus Lake i n November, 1953, and a r t i f i c i a l l y f e r t i l i s e d before transportation to the University hatchery. They were maintained i n fresh water tanks at temperatures which ranged from 1 2 . 5 ° C i n November through 5°C i n January to 15°C i n August. tion. Pablum.  Hatching began on the f i f t i e t h day after f e r t i l i z a -  The alevins were fed on a mixture of canned salmon and Samples were taken every hour i n the early stages.  Thereafter, the i n t e r v a l s were increased to two weeks f o r the l a t e s t stages of the series sampled. was continued u n t i l August, 1954.  This fresh water s e r i e s  Migrating f r y were trapped  i n A p r i l , 1955 as they entered the sea at Port John, B . C . , and were held i n s a l t water pens i n the bay.  These specimens were  sampled weekly at f i r s t , but l a t e r bimonthly u n t i l November 1955.  Further sea water specimens were obtained during t h e i r  migration into the P a c i f i c i n J u l y and August of 1955.  A  second series was maintained at the University hatchery during the f a l l and winter of 1955.  In the l a t e spring of 1956 they  were transferred to the Vancouver Public Aquarium and divided into s a l t and fresh water populations which were sampled weekly.  B. Methods 1. F i x a t i o n .  - 28 -  (a) General h i s t o l o g i c a l  detail.  Eggs and f r y were fixed whole i n Bouin's f l u i d or i n Smith's formal bichromate.  These f i x a t i v e s gave rather poor  preservation of c y t o l o g i c a l d e t a i l .  They proved adequate for  demonstration of general h i s t o l o g i c a l structure.  Smith-fixed  material hardened less during subsequent treatment and was found more satisfactory for yolky material.  Pry and fingerlings  collected i n the f i e l d were fixed i n Bouin's. (b) Mitochondria and brush borders. Regaud's f i x a t i v e  (Krajian, 1940) was found to give  very satisfactory preservation of mitochondria and brush borders. The material was f i x e d four days i n Regaud's followed by postchroming f o r 10 days i n 3 percent potassium bichromate.  The  kidney was p a r t i a l l y dissected out of the f r e s h l y k i l l e d animal to aid penetration of the  fixative.  2. Imbedding and sectioning. Material was embedded i n a 50:50 mixture of 5 0 - 5 3 ° C m.p.  and 5 4 - 5 6 ° C m.p.' tissuemat.  The mixture was found to be  most satisfactory f o r the summer conditions under which much of the work was performed. not containing yolk. off  Routine methods were followed f o r tissues  Whenever possible,  the yolk p r i o r to embedding.  embryos were dissected  Where t h i s was not  possible  either the P e t e r f i double embedding technique (Pantin, 1948)  or  the method of c l e a r i n g with terpineol from 90% alcohol was employed.  The kidney was dissected out of older embryos before  embedding.  The need for d e c a l c i f i c a t i o n was thereby avoided.  - 29 -  Yolk-containing blocks were trimmed down to the yolk on one side and soaked 24 hours i n softening agent p r i o r to sectioning.  Material for demonstration of general structure  was cut at 10 u , that for demonstration of mitochondria at 5 u . Sections were adhered to s l i d e s with Mayer's albumen.  In the  case of yolk-containing sections i t was necessary to coat  the  adhered sections with a 0.5% solution of c e l l o i d i n i n 50% etheralcohol immediately after dewaxing to prevent t h e i r becoming detached.  The c e l l o i d i n was then removed with acetone p r i o r to  clearing the stained sections.  3. Staining. (a) Bulk s t a i n i n g . Bulk staining with Grenadier's borax carmine was t r i e d to simplify glomerular counting.  The attempt was unsuccessful  owing to heavy uptake of s t a i n by the kidney tubules and myeloid tissue.  Staining of thick sections (40 microns) was also un-  successful. (b) General h i s t o l o g i c a l  detail.  Heidenhain's i r o n haematoxylin was used f o r demonstration of general d e t a i l owing to i t s good photographic properties. For general purposes no counterstain was used.  Counterstaining  with eosin was used for demonstration of acidophila i n the c e l l s of the g i l l and renal epithelium.  Heidenhain's azan was used  f o r demonstration of the f i n e structure of the glomeruli. (c) Mitochondria.  - 30 -  Altmann's acid fuchsin a n i l i n e (Krajian, counterstained with methyl green gave excellent  1940)  results.  4. Glomerular counts. It was found that, with the exercise of a reasonable amount of caution, accurate counts (4%) could be obtained by counting glomeruli i n every f i f t h section i n the larger f i s h . In the smaller f i s h , each section was studied.  -  31  -  V I . OBSERVATIONS  A. The structure of the f u l l y differentiated kidney. 1. Anatomy. The d e f i n i t i v e kidney i s a mesonephros.  I t i s an  elongate organ extending from the twelfth to the f o r t i e t h segment.  In i t s early development i t i s d i s t i n c t l y  bilobate  but by the time the f i s h i s a year o l d the lobes are obscured along most of the length of the kidney.  The fore kidney region  (fourth to twelfth segments) i s flattened and tends to be b i lobate.  The post cardinal veins pass out of the kidney i n the  f i f t h segment at which point the kidney tissue grows ventrol a t e r a l ^ along the veins as they pass around the anterior end of the l i v e r .  The right cardinal v e i n i s continuous with the  caudal vein i n the f o r t i e t h segment.  It enters the dorsal as-  pect of the kidney i n the t h i r t y - e i g h t h segment and passes anteriad i n the centre of the kidney t i s s u e .  Opposite the  sixteenth somite i t curves to the r i g h t before entering the region of the fore kidney (figure 4 ) .  The entire fore kidney  i s permeated by a network of venous sinusoids.  Blood from the  r i g h t postcardinal passes direct to the r i g h t common c a r d i n a l . Blood drains from the sinusoids into the r i g h t and l e f t postcardinal veins.  Patches of well-vascularised i n t e r r e n a l tissue (h$VT€.  are distributed throughout the pronephric region^  31)'  Arterial  blood reaches the i n t e r r e n a l tissue from a r t e r i o l e s which develop only after occlusion of the main pronephric a r t e r i o l e .  - 32 -  Paired intersegmental renal veins drain blood from the kidney tissue into the cardinal v e i n (figure 5).  Portal  blood from the dorsal and ventral body walls reaches the kidney sinusoids by way of intersegmental venules which enter the lower angles of the kidney.  A r t e r i a l blood reaches the kidney from  paired a r t e r i o l e s which enter at the dorsal angles opposite alternate segments.  Two median a r t e r i a p r i m i t i v a mesenterica  pass through the kidney tissue to supply the swim bladder and hind gut.  The median coeliaco-mesenteric artery passes through  the fore kidney to supply the anterior v i s c e r a .  The lower  angles of the kidney are usually indented corresponding with the myosepta. The Wolffian ducts pass posteriad along the lower surface of the kidney to the t h i r t y - n i n t h segment where they unite p r i o r to passing ventrad to the cloaca.  The common duct  enters a bladder-like distention (figure 25) but no sphincter i s present. 2. Histology. The excretory r o l e of the kidney i s performed by a number of non-metameric renal units made up of several glomerul a r tubules and a c o l l e c t i n g duct.  The c o l l e c t i n g ducts d i s -  charge into the segmental ducts on either side.  Bach unit  comprises several nephrons w i t h : a) glomerulus and Bowman capsule. wide d i l a t i o n of the free end of the tubule.  The capsule i s a It i s l i n e d with  simple squamous epithelium which undergoes a gradual t r a n s i t i o n  - 33 -  to the l a t e r a l l y compressed columnar epithelium of the neck segment.  The glomerular pole i s usually diametrically opposite  the tubular pole.  The well developed glomerulus i s an e l l i p s o i d -  a l mass, about 70 microns X 50 microns, composed of a r t e r i a l c a p i l l a r i e s among which are d i s t r i b u t e d connective tissue c e l l s and f i b r e s .  The whole structure i s enclosed by the epithelium  of the Bowman capsule which i s reflected into the glomerular tissue around the c a p i l l a r i e s (figure 8). are usually almost e n t i r e l y avascular.  The smaller glomeruli  The centre of many larger  glomeruli i s taken up by a v e s i c l e . b) a short neck segment of low simple columnar l a t e r a l l y compressed c e l l s with e l l i p t i c a l nuclei basally located i n the c e l l s (figure  8).  c) a f i r s t major segment of low columnar weakly basop h i l i c c e l l s with a high brush border at least one quarter the height of the c e l l s . (figure 9).  The spherical nuclei are basally located  Granular mitochondria are concentrated i n the basal  region of the c e l l (figure 10).  The lumen of the tubule i s  rather wide throughout the length of t h i s segment. d) a second major segment of t a l l e r columnar c e l l s with a low brush border (figure 12).  Rodlike mitochondria are  distributed throughout the c e l l (figure 11).  The tubule lumen  i s narrow i n t h i s segment. e) a t h i r d major segment of low columnar unbordered © e l l s with spherical basally placed n u c l e i .  The tubule lumen  -34-  i s very wide (figure  13).  No intermediate segment i s found i n 0. gorbuscha. A c o l l e c t i n g duct of t a l l columnar epithelium 14) drains into the segmental duct (figure 15),  (figure  also of t a l l  columnar epithelium. The intertubular spaces are packed with myeloid tissue permeated by a r i c h net of venous sinusoids.  Post glomerular  blood apparently enters the sinusoids since no c a p i l l a r y nets are to be found associated with the  tubules.  In the twenty-seventh to the twenty-ninth segments large e l l i p s o i d a l highly lobulated corpuscles of Stannius appear. The corpuscles are r i c h l y vascularised.  Between s i x and twelve  corpuscles are found.  B. The Chloride C e l l s of the G i l l s . In the eight month old sea water acclimatised  specimen  the "chloride" c e l l s appear as weakly a c i d o p h i l i c c e l l s located i n the g i l l epithelium and i n well vascularised areas of the mouth.  However, an eight month o l d embryo of the fresh water  series (67 mms) also showed weak a c i d o p h i l i a of c e r t a i n c e l l s i n the g i l l and o r a l region.  C. The Development of the Mesonephros. 3*33 mms.  16 somites The pronephros of t h i s embryo has commenced to d i f f e r -  entiate.  The nephrotomes of the s i x t h and seventh segments,  - 35 -  from which the d e f i n i t i v e pronephric chamber develops,  are  fused together and open over their whole length to the coelom. The tubule rudiment^or pronephric fold,appears as a thickening of the dorso l a t e r a l surface of the fused nephrotomes.  This  i s continuous i n the eighth and ninth segments with a s o l i d uniform rod of c e l l s (duct rudiment) which formed as a f o l d of the intermediate c e l l mass i n t h i s region.  The vascular strand  appears as a darkly staining mass of c e l l s beneath the hypochord i n the fourth to the seventh segments.  Posterior to the  ninth segment the intermediate c e l l mass i s an undifferentiated unsegmented triangular mass of c e l l s between the somite and the hypomere (figure 16).  D i f f e r e n t i a t i o n of mesoderm has not pro-  ceeded beyond the sixteenth segment. 4.4 mms.  23 somites The duct rudiment i s set aside i n the anterior meso-  nephric segments as a s o l i d cylinder which forms as a f o l d of the dorsolateral surface of the intermediate c e l l mass (figure 17).  In the pronephric region the duct i s becoming hollow.  Migration of c e l l s from the medial surface of the intermediate c e l l mass has established the vascular strand i n the more ant e r i o r mesonephric segments. 7.4 mms. The pronephros i s well formed with large pronephric chambers and single median glomerulus. nephric tubule has commenced.  Convolution of the pro-  The segmental ducts are complete  to the l e v e l of the cloaca where they unite and pass v e n t r a l l y  - 36 -  to enter the cloaca.  A narrow hand of p a l e l y stained meso-  nephric bridge c e l l s derived from the intermediate c e l l mass stretches uninterrupted the length of the mid kidney from one duct to the other between the aorta and the post cardinal vein (figure 18 - 5.24 mms.).  The aorta has differentiated from the  intermediate c e l l mass along the length of the trunk.  Ventro-  l a t e r a l to i t on the r i g h t i s located the r i g h t post cardinal vein which i s continuous caudally with the caudal vein and a n t e r i o r l y with the pronephric p o r t a l system.  P r o l i f e r a t i o n of  myeloid tissue has commenced from the vascular strand tissue just posterior to the pronephric chambers. Prom the dorsal surface of each duct i n the f i f t h to the twenty-seventh  twenty-  segments rudimentary corpuscles of  Stannius appear as palely-stained hemispherical outgrowths of the duct epithelium (figure  27).  16.66 mms. Condensations of c e l l s of the mesonephrogenic bridge •nearest the ducts form hemispherical caps of basophilic c e l l s which become closely applied to the ducts (figure 19).  In t h i s  embryo three such condensations are apparent on the l e f t and two on the r i g h t . l o c a t e d i n the posterior region of the midkidney.  The corpuscles of Stannius are separated from the ducts  by this time as large e l l i p s o i d a l bodies (figure  28).  19.05 mms. The pronephros of this embryo has attained i t s peak development.  The pronephric chamber and glomerulus have attained  - 37 -  t h e i r maximum length of 400,11.  The uriniferous tubule i s high-  l y convoluted and c y t o l o g i c a l d i f f e r e n t i a t i o n i n t o two regions has occurred.  Fifteen mesonephric blastulae are apparent, nine  on the right and s i x on the l e f t .  Most of the new blastulae  are added anterior to those present i n the 10.95 mm. embryo. Elongation of the more posterior rudiments has commenced.  More  rapid c e l l d i v i s i o n i n the l a t e r a l aspects causes the forming tubules to curve around the ducts i n a medio-ventral d i r e c t i o n . The proximal ends of the s o l i d tubule rudiments are fused to the ducts.  The mesonephric bridge i s obscured by myeloid t i s s u e ,  p r o l i f e r a t e d from the vascular strand. to f i l l  Myeloid c e l l s have begun  the spaces between the post cardinal v e i n , ducts and  tubules.  The corpuscles of Stannius are becoming penetrated by  a r t e r i o l e s direct from the aorta which follow connective trabeculae into the corpuscular stroma.  tissue  No other a r t e r i a l blood  supply to the mesonephros has developed. 19.80 mms. Forty^six mesonephric rudiments appear, the new ones being added anterior to those observed at 19.05 mms. f o r t y - s i x only the most anterior are unelaborated. der show varying degrees of tubular development.  Of the The remain-  In the  first  formed rudiments the free end i s d i l a t e d into a wide Bowman capsule (figure 20).  The lumen of the capsule extends i n t o  the tubule i n some cases and some tubules are open to the ducts. Ten such rudimentary capsules are apparent. 20.41 mms. The antitubular aspect of some of the newly formed  - 38 -  Bowman capsules i s invaginated by a s o l i d mass of l i g h t l y staining c e l l s , (figure 21).  the rudiments of the mesonephric glomeruli  There i s as yet no sign of vascularisation of  the g l o m e r u l i nor of a r t e r i a l blood supply to the mesonephros. There are seventeen rudimentary glomeruli i n t h i s specimen. 20.9 mms. Thirty-one i n c i p i e n t glomeruli are apparent.  The  epithelium of the expanded Bowman capsules i s squamous changing gradually to t a l l columnar at the point from which the tubule leaves.  The c e l l s of the neck segment are very basophilic and  l a t e r a l l y compressed.  The remainder of the tubule i s unborder-  ed t a l l columnar weakly basophilic epithelium. low columnar epithelium.  The duct i s of  The corpuscles of Stannius appear as  large e l l i p s o i d a l masses of vascularised tissue located dorsol a t e r a l ^ to the ducts. 26.9 mms. A r t e r i a l blood i s supplied to the mesonephros by paired mesonephric a r t e r i o l e s which enter through the i n t e r segmental connective tissue of each segment.  Venous blood from  the dorsal and ventral myosepta enters the four corners of the trapezoid mesonephros.  The larger glomeruli i n the posterior  regions are becoming vascularised (figure 22).  Mesonephric  rudiments extend a n t e r i o r l y to the twelfth segment on the l e f t though the more anterior rudiments are not elaborated.  The area  between the tenth (last pronephric) and twelfth segments never shows any development of kidney structures.  Glomerular tubules  - 39 -  extend only as f a r as the twentieth segment at t h i s  time.  Development of rudiments i n the right lobe extends only to the sixteenth segment since anterior to t h i s the lobe i s f i l l e d by the very large r i g h t post cardinal v e i n .  F i f t y - f i v e meso-  nephric glomeruli are found i n t h i s specimen. 28.58 mms. Secondary mesonephric condensations i d e n t i f i e d by the deep basophilia of their c e l l s are s t a r t i n g to appear c l o s e l y applied to the primary tubules.  The more posterior primary  glomeruli are large and quite vascular (figure 23)• tubules are differentiated into (1) neck segment; (2)  Their first  major segment of low columnar c e l l s with a low brush border and 1  large e l l i p s o i d a l basally located n u c l e i ; (3) second major segment of t a l l columnar c e l l s with a low brush border and smaller spherical muclei; (4) t h i r d major segment of low columnar unbordered c e l l s and a connecting segment of t a l l columnar c e l l s . The duct epithelium i s now t a l l columnar. es seventy-six glomeruli.  This specimen possess-  The more posterior glomeruli are high-  l y vascular and apparently f u n c t i o n a l . 29.78 mms. Incipient secondary Bowman capsules are commencing to appear by this time though d i f f e r e n t i a t i o n of the secondary tubules has not yet occurred (figure 24).  One hundred and twelve  primary and nine secondary glomeruli appear. pronephric a r t e r i o l e has commenced. 35.O mms  Occlusion of the  - 40 -  Mesonephric glomeruli extend as far as the sixteenth segment on the l e f t .  Secondary condensations are as f a r a n t e r i -  ad as the twenty-sixth segment.  P o s t e r i o r l y , no tubules appear  behind the t h i r t y - e i g h t h segment at which point the ducts leave the kidney.  Myeloid tissue extends to the l e v e l of the cloaca  (segment f o r t y ) .  Further new primary oondonoationo and secondary  condensations appear i n the posterior regions of the mid kidney. The pronephric a r t e r i o l e i s p a r t l y occluded and the glomerulus shows reduction of v a s c u l a r i t y .  In a second speci-  men the artery i s e n t i r e l y occluded and the glomerulus reduced i n length to 210 microns by invasion of i t s anterior and posterior ends with fibrous connective tissue.  The nephrostomes are  Heir  p a r t l y closed by ingrowth of c e l l s from i-t* epithelium. 49.6 mms. The pronephric artery i s completely occluded.  The  glomerulus i s much reduced i n size and i t s v a s c u l a r i t y reduced by a core of fibrous connective t i s s u e . tubular borders has commenced.  Degeneration of the  There has been l i t t l e  change i n  the mesonephros beyond an increase i n the size and v a s c u l a r i t y of the already formed glomeruli, addition of new primary and secondary glomeruli and the appearance of new condensations. The t o t a l number of glomeruli i s now three hundred and s i x t y nine.  The corpuscles of Stannius are large and highly lobulated  with a very r i c h a r t e r i a l supply (figure 67 mms.  29).  The mesonephros i s becoming t y p i c a l l y trapezoid i n  -  41 -  cross section (figure 26).  One thousand mesonephric glomeruli  were counted i n one specimen, twelve hundred and ten i n another. Masses of very vascular palely stained i n t e r r e n a l tissue appear scattered throughout the pronephric region (figure 30).  The  pronephric chambers are open, but the glomerulus i s very v e s i cular and degenerate i n appearance.  The entire kidney i s r i c h l y  permeated with blood sinuses.  D. Comparison of Marine and Fresh Water Series. There i s no s i g n i f i c a n t difference i n the histology of glomeruli or tubules i n specimens taken from fresh or s a l t water.  No aglomerular tubules are apparent even i n the most  anterior mesonephric region.  Occlusion of the pronephric  a r t e r i o l e commences at the same stage of development i n both series.  Glomerular counts indicate a rather larger number of  glomeruli i n fresh water f i s h than i n marine f i s h of a given size.  - 42 -  VII.  DISCUSSION  The prolonged attachment of the teleost intermediate c e l l mass to the l a t e r a l plate has caused considerable confusion i n the i n t e r p r e t a t i o n of the early phases of teleost kidney development.  The o r i g i n of the mesonephric blastema from the  intermediate c e l l mass i n teleosts was established only i n 1934 (Maschkowzeff, 1934).  During the course of the present  investi-  gation i t was found that the mesonephric blastema i n the pink salmon was derived from that portion of the intermediate c e l l mass nearest to the somite. The development of the Bowman capsule i s  somewhat  different than that of any species hitherto reported.  In general  the Bowman capsule i s reported as a r i s i n g as a distention of the already hollow tubule.  Moghe (1940) reported the o r i g i n of  Bowman capsules i n Thynnichthys as separate condensations of the mesonephrogenic bridge c e l l s .  These condensations  subsequently  became attached to the already formed tubule and opene* to i t . In the pink salmon the Bowman capsule develops as a distention of the tubule rudiment p r i o r to i t s  tubulation.  The cavity of  the Bowman capsule subsequently extends into the tubule rudiment and f i n a l l y breaks through into the lumen of the segmental duct. The glomerulus forms as a separate condensation apparently of vascular strand c e l l s .  Vascularisation of the glomeruli p r i o r  to their connection with the aorta by c a p i l l a r i e s developed i n s i t u has been reported i n birds (Davies i n Praser, 1950) but this i s apparently the f i r s t time that i t has been observed i n  - 43 -  teleosts. Reference to the table (figure 6) indicates a rather higher number of glomeruli i n f r e s h water than i n marine s p e c i mens of comparable length. (figure 7) and efficients  11  t  H  The logarithmic regression curves  values calculated f o r the regression co-  indicate that there i s s i g n i f i c a n t difference i n the  rate of development of glomeruli i n marine and fresh water f i s h . The lower figures f o r the marine series may be, i n p a r t , a r e s u l t of a temporary retardation of the rate of development of glomerul i r e l a t i v e to length, perhaps associated with the abrupt transf e r from fresh to s a l t water.  The experiment as performed here  i s not e n t i r e l y satisfactory since the f i s h ceased to thrive when maintained i n fresh water past the time of their normal migration.  It i s suspected that a species of the P a c i f i c salmon  which remains t r u l y euryhaline for a prolonged period might prove more satisfactory for an experiment of t h i s type.  The present  results should be viewed with caution since no attempt was made to assess the effect of the different temperatures of the fresh water and marine series on the rate of appearance of glomeruli.  - 44 -  VIII.  CONCLUSIONS  1. The development of the mesonephros i n the pink salmon closely p a r a l l e l s that of teleosts described by other authors. 2. The mesonephros i s glomerular during i t s ment.  develop-  Since glomeruli continue to develop following the trans-  f e r from fresh to s a l t water i t seems l i k e l y that the mesonephros of the adult w i l l be glomerular.  No aglomerular tubules  were observed even i n the oldest specimens studied.  No s i g -  n i f i c a n t difference i n the s i z e of glomeruli developed i n s a l t or fresh water was observed. 3. It appears that the increased osmotic concentration of the marine environment has the effect of retarding rate of development of glomeruli. 4. 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The vascular pole of the glomerulus i n the kidney of vertebrates. Anat. R e c , 76:381-389.  21.  and L . C o n d o r e l l i . 1928. Studies on aglomerul a r and glomerular kidneys. II. Physiological. Am. J . P h y s i o l . , 86:383-398.  22.  and C. Schnitter. 1933. The renal unit i n the vertebrate kidney. Am. J . Anat., 53:55*  23. Emery, C. 1882. Etudes sur le developpement et l a morphologie du r e i n des poissons osseux. Arch. i t a l . biologique. i i . i n Moghe, 1947. 24. P e l i x , W. 1906. Die Entwicklung der Harnapparatus. Handbk. v e r g l . exp. Bntwicklungslehre der W i r b e l t i e r e , ^(l):81-422. 25. P l o r k i n , M. and S. Morgulis. 1949. Biochemical Evolution. V. Biochemical Adaptations. Academic Press, New York, N.Y. 26. F r a s e r , E . A . 1950. Development of the vertebrate system. B i o l . Rev., 25:159-187.  excretory  27. Garrett, F . D . 1942. The development and phylogeny of the Corpuscle of Stannius i n Ganoid and Teleostean fishes. J . Morph., 70:41-67. 28. Gerard, P. 1936. Comparative histophysiology of the vertebrate nephron. J . Anat., 70:354-379.  -  29.  47 -  G e r a r d , P. 1954. Organes U r o g e n i t a u x . I . Organes Excreteurs. I n T r a i t e de Z o o l o g i e , T. X I I . Vertebres: G e n e r a l i t e s , Embryologie Topographique, A n a t o m i e Comparee. Masson e t C i e , P a r i s , e d . P-P. Grasse.  30.  a n d R. C o r d i e r . 1934. E s q u i s s e d'une h i s t o p h y s i o l o g i e compare'e d u r e i n d e s vertebrates.  B i o l . Rev., 9:110-131. 31.  Getman, H.C. 1950. A d a p t i v e changes i n t h e c h l o r i d e c e l l s of A n g u i l l a r o s t r a t a . B i o l . B u l l . , 99:439-445.  32.  G o o d r i c h , E . S . 1930. S t u d i e s oJi-the s t r u c t u r e a n d d e v e l o p ment o f v e r t e b r a t e s . X I I I . E x c r e t o r y o r g a n s a n d g e n i t a l ducts. M a c m i l l a n Co., L o n d o n .  33.  Grafflin,  34.  A . L . 1929. The p s e u d o g l o m e r u l i o f t h e k i d n e y o f Lophius p i s c a t o r i u s . Am. J . A n a t . , 44:441. 1931. The s t r u c t u r e o f t h e r e n a l t u b u l e o f the t o a d f i s h . B u l l . J o h n s H o p k i n s H o s p . , 48:  269-271. 35. of  1933. Glomerular degeneration i n the kidney t h e daddy s c u l p i n ( M y x o c e p h a l u s scorpius).  A n a t . Rec. ,57:59-80.  ,g  1937ft». O b s e r v a t i o n s u p o n t h e a g l o m e r u l a r nature o f certain  61:165-173.  37.  teleostean kidneys.  J . Morph.  1937 Jb The p r o b l e m o f a d a p t a t i o n t o f r e s h and s e a water i n t h e t e l e o s t s , viewed from t h e standpoint o f thestructure o f the r e n a l tubules. J . C e l l . Comp. P h y s i o l . , 9:469-476.  38.  1937e. T h e s t r u c t u r e o f t h e n e p h r o n i n t h e s c u l p i n , Myxocephalus octodecemspinosus.  Rec.  68:145-163.  Anat.  39.  1937«t. O b s e r v a t i o n s u p o n t h e s t r u c t u r e o f t h e n e p h r o n i n t h e common e e l . Am. J . A n a t . , 61:21-62,  40.  a n d T . E . Moses. 1934. A microfluoroscopic study o f the t e l e o s t e a n kidney. A n a t . R e c , 59:  449.  41.  G r a y , P. 1930. The development o f t h e amphibian k i d n e y . I . The d e v e l o p m e n t o f t h e mesonephros o f Rana temporaria. Q u a r t . J . M i c r . S c . , 73:508-.  - 48 -  42. Gray, P. 1932. The development of the Amphibian kidney. I I . The development of the kidney of Triton vulgaris and a comparison of this form with Rana temporaria. Quart. J . Micr. S c . , 75: 423^446. 43. Gruenwald, P. 1942. Common t r a i t s i n development and structure of the organs o r i g i n a t i n g from the coelomic w a l l . J . Morph., 70:353-587. 44. G u i t e l , P. 1906. Recherches sur l'anatomie des reins de quelques Gobiescoides. Arch. Zool. Exp. et Gen. £:505-698. 45. H a l l , R.W. 1904. Development of the mesonephros and Mullerian ducts i n Amphibia. B u l l . Mus. Comp. Zool. Harvard. 45:321. 46. Kempton, R . T . 1943. Studies on the elasmobranch kidney. I . The structure of the renal tubule of the spring dogfish (S. acanthias). J . Morph., 73: 247-263. ~ 47. Keys, A . B . 1931. Chloride and water secretion and absorbt i o n by the g i l l s of the e e l . Zeitschr. v e r g l . P h y s i o l . , 15:364-388. 48.  1933. The mechanism of adaptation to varying s a l i n i t y i n the common eel and the general problem of osmotic regulation i n f i s h e s . Proc. Roy. Soc. london, Ser. B, 112:184-199.  49.  and E . N . Willmer. 1932. Chloride secreting c e l l s i n the g i l l s of fishes with s p e c i a l reference to the common e e l . J . P h y s i o l . , 76:368-578.  50. Kindahl, M. 1938. Zur Entwicklung der Exkretionsorgane von Dipndern und Amphibien. Acta Zool. Stockholm, 19:1-190. 51. K r a j i a n , A . A . 1940. H i s t o l o g i c a l technique. S t . Louis.  C . V . Mosby,  52. Krogh, A. 1937. Osmotic regulation i n fresh water fishes by active absorbtion of chloride ions. Zeitschr. v e r g l . P h y s i o l . , 24:656-666. 53. Lankester, E . R . 1877. Notes on the embryology and c l a s s i f i cation of the animal kingdom. In Praser, 1950. 54. L i Kowe Tchang. In Marshall, 1934. Recherches histologiques sur l a structure des reins des oiseaux. These medicale. Lyon, 1923.  -  49  -  55. Marshall, E . K . 1929. The aglomerular kidney of the toadf i s h (Opsanus tau). B u l l . Johns Hopkins Hosp., 45:95-100. 56.  1930. A Comparison of the glomerular and aglomerular kidney. Am. J . P h y s i o l . , 94:1-10.  57.  and H. W. Smith. 1930. The glomerular development of the vertebrate kidney i n r e l a t i o n to habitat. B i o l . B u l l . , 59:135-153.  58. Maschkowzeff, A. 1926. Zur Phylogenie des Urogenitalsysterns der Wirbeltiere auf Grund der Entwicklung des Mesoderms, der Pronephros, der Analfiffnung und der Abdominalporen bei Acipenser s t e l l a t u s . Zool. J a h r b . , Abt. Anat. u Ontog., 51:9. 59.  1934. Zur Phylogenie der Geschlechtsdrttsen und der Geschlechts aus fuhr gange bei den Vertebrata auf Grund von Porschungen betreffeud die Entwicklung des Mesonephros und der Geschleehtsorgane bei den Acipenseridae, Salmoniden und Amphibien. I . Die Entwicklung des Mesonephros und der Genitaldrttse bei den Acipenseridae und Salmonidae. Zool. Jahrb. Abt. Anat. u. Ontog.,  59(i):1-68.  60. McEwen, R.S. 1949.  Vertebrate Embryology.  H o l t , New York.  61. Mcintosh, W., and E . Prince. 1887. On the development and l i f e history of the teleostean food and other fishes. Trans. Roy. Soc. E d i n . , 35:. 62. Moghe, M.A. 1945. Development of the mesonephros i n a telestean, Thynnichthys sandkhol. Quart. J . Micr. S c . , 85:129-151. 63.  1947. The development of the kidney i n f i s h e s . Proc. Indian S c i . Congress, 113-122.  64. Nash, J . 1931. The number and size of glomeruli i n the kidneys of f i s h e s , with observations on the morphology of the r e n a l tubules of f i s h e s . Am. J . A n a t . , 47:425-445. 65. Nussbaum, M. 1878. Uber der Niere der Wirbeltiere. Moghe, 1945.  In  66. O'Connor, R . J . 1939. Experiments on the development of the pronephric duct. J . A n a t . , London, 73: 145-154. ~~ 67. Owen, B.B. 1938. A simple teleost kidney i n the genus Cyclothone. B i o l . B u l l . Woods Hole, 74:349-565.  - 50 -  68.  P a n t i n , C P . A . 1948. Notes o n m i c r o s c o p i c a l technique f o r zoologists. Cambridge U n i v e r s i t y P r e s s .  69. P r i c e ,  G.C. 1904. A f u r t h e r study o f t h e development o f the e x c r e t o r y organs i n B d e l l o s t o m a s t o u t i .  Am. J . A n a t . , 4:117-138.  70.  R o s e n b e r g , A . 1867. 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The development o f t h e pronephros i n t h e common p e r c h ( P e r c a f l u v i a t i l i s ) . Quart. J .  M i c r . S c . , 75:557.  78.  Swaen, A. a n d A . B r a c h e t . 1899, 1901. Etudes sur l e s premieres phases du developpement des organes d e r i v e s d u mesoblaste chez l e s p o i s s o n s T e l e ostiens. A r c h . B i o l . P a r i s , 16:173-311; 18:  73-190.  79. V e r n e ,  80. V i t t e r ,  —  J . 1922. C o n t r i b u t i o n s a 1'etude des r e i n s a g l o merulaires. L ' a p p a r e i l des poissons Lophobranches. A r c h , de A n a t . M i c r o s c o p . , T. 18. p p . 357-407. R.W. 1935. The m o r p h o l o g y a n d d e v e l o p m e n t o f t h e metanephric glomerulus o f the pigeon. Anat. R e c ,  61:12.  81.  Wearn, J . T . , a n d A.N. R i c h a r d s . 1924. O b s e r v a t i o n s on t h e composition o f glomerular urine with p a r t i c u l a r r e f e r e n c e t o the problem o f r e a b s o r b t i o n i n t h e r e n a l t u b u l e s . Am. J . P h y s i o l . , 71:209-227.  82.  W i k g r e n , B o - J u n g a r . 1953. O s m o t i c r e g u l a t i o n i n some a q u a t i c animals with s p e c i a l r e f e r e n c e to the i n f l u e n c e o f temperature. A c t a Z o o l . P e n n i c a . 71:1.  -  PLATE I  51a -  ANATOMY  Venous drainage  of the teleost  mesonephros.  Figure 1  True p o r t a l system i n a d u l t .  Figure  2  Left p o r t a l vein only i n adult.  Pigure  3  Right post c a r d i n a l only i n adult no t r u e p o r t a l s y s t e m .  *cv  caudal  pcv^  left  pcv  right  *pv c  r  vein  post  cardinal vein  post  cardinal vein  portal vein common c a r d i n a l  •adapted from Audige,  (1910).  -  PLATE I I  Pigure  Pigure  52a  -  ANATOMY  4  5  V e n t r a l view o f e n t i r e kidney 65 mms. f i n g e r l i n g .  b  bladder  pcv^  left  pcv  r i g h t post  r  post  sd  segmental  ugp  urogenital  cardinal cardinal  of  vein vein  duct papilla  T r a n s v e r s e s e c t i o n o f mesonephros 65 mms. f i n g e r l i n g . ( i d e a l i s e d ) .  a  aorta  pcv  post  cardinal  ra  renal arteriole  rv  renal  v  portal  vein veim  vein  ANT  cigp  Figure-  4  Fi g u  P  e  5  - 53a  TABLE  -  - PHYSIOLOGY  Figure  6  Tabular representation of glomerular c o u n t s i n f r e s h and s a l t w a t e r s e r i e s .  Fresh  D at e 4/6/56  18/6/56  Ma r i n e __S e r i e s  Water . S e r i e s  Length  No. Glomeruli  32.0 36.2 33.4 33.4 2 7.7 27 .8 30.9 32.1  2 91 1 94 2 63 135 249 2 21 2 17  37.5 33 .5 3 2.8  32 0 268 2 1 9  i*  •  #  Length  No. Glomeruli  3 7. 6 36.1 35. 2 3 1.0 36.0 29.6 30;2  2 13 194 18 1 14 6 17 6 129 15 3  43.7 3 9 .8 40 .2 41 .2.  3 2 2 2  1 0 79 8 30 6  2 5/6/56  38.6 37.4  325 310  39.8 34.4 3 4.0 3 3-4  2 51 208 2 48 2 |9  2/7/56  303 3 8 .7 331 3 3.9  303 34 2 23 7 2 86  4 5.8 4 1 .7 44. 6 32.3  307 2 88 293 2 04 °  9/7/5 6  4 0.8 36 7  33 2 2 95  5 5 3 3  576 3 76 2 29 2 77  4.9 1.7 6.0 66  -  PLATE I I I  Pigure 7  54a -  PHYSIOLOGY  Graphical representation of relationship o f number o f g l o m e r u l i t o l e n g t h i n f r e s h and s a l t w a t e r s e r i e s ( R e g r e s s i o n lines).  Regression  Equation?  I.F.W. Y -  2.36X-  1.12 t  2.S.W. Y »  2.44X-  2.43  d 8 f f  - 0.0 4 8  .  - 55a  P L A T E IV  i I  -  HISTOLOGY  Pigure 8  Glomerulus and c a p s u l e o f d i f f e r e n t i a t e d nephron. Heidenhaihn haematoxylin. x  800  a  Pigure 9  fully-  arteriole  be  Bowman c a p s u l e  e  epithelial cell  ns  neck  nucleus  segment  Transverse section of f i r s t m a j o r segment. Heidenhaihn haematoxylin. x  b  800  brush  border  - 56a  PLATE V  HISTOLOGY  P i g u r e 10  Transverse s e c t i o n o f f i r s t major segment s t a i n e d f o r m i t o c h o n d r i a . Altmann a c i d f u c h s i n . x  Pigure  -  11  800  T r a n s v e r s e s e c t i o n o f second major segment s t a i n e d f o r m i t o c h o n d r i a . Altmann a c i d f u c h s i n . x  800  - 57a  PLATE V I  -  HISTOLOGY  P i g u r e 12  T r a n s v e r s e s e c t i o n o f second major segment. H e i d e n h a i h n H a e m a t o x y l i n . x  P i g u r e 13  800  T r a n s v e r s e s e c t i o n o f t h i r d major segment. H e i d e n h a i h n H a e m a t o x y l i n . x  s  800  secondary condensation  - 58a -  PLATE VII  HISTOLOGY  Figure 14  Transverse section of c o l l e c t i n g Heidenhaihn Haematoxylin.  duct.  x 800  Figure 15  Transverse section of segmental duct. Heidenhaihn Haematoxylin. x 600  - 59a -  PLATE VIII  EMBRYOLOGY  Figure 16  Transverse section of 3.45 nun. embryo through sixteenth segment to show triangular intermediate c e l l mass. x 350  Figure 17  ch  notochord  ent  entoderm  icm  intermediate  IP  lateral,plate  n  neural keel  s  somite  c e l l mass  Transverse section of 4*4 mm embryo through twentieth somite to show separation of duct rudiment and vascular strand. x 350  d  duct rudiment  m  mesonephric bridge rudiment  vs  vascular strand  ic  m  I P  ent  m  - 60a -  PLATE IX  EMBRYOLOGY  Pigure 18  Transverse section of 5.24 mm embryo to show l o c a l i s a t i o n of structures derived from the intermediate mesoderm. x 125  Pigure 19  a  aorta  ch  notochord  g  gut  mb  mesonephrogenic bridge  pcv  post cardinal vein  sd  segmental duct  Transverse section of l e f t lobe of kidney of 16.66 mm embryo to show mesonephric condensation. x 700  a  aorta  mb  mesonephric bridge  mc  mesonephric condensation  pcv  post cardinal vein  sd  segmental duct  - 61a -  PLATE X  EMBRYOLOGY  Pigure 20  Transverse section of kidney of 19.05 mm a l e v i n to show early stage i n development of the Bowman capsule. x 350  Pigure 21  be  Bowman capsule  go  gonad  mt  myeloid tissue  sb  swim bladder  sd  segmental duct  tr  tubule rudiment  Transverse section of kidney of 19.8 mm alevin to show i n c i p i e n t glomerular invagination. x 700  be  Bowman capsule glomerular rudiment  tr  tubule rudiment  -  PLATE X I  Figure  62a -  EMBRYOLOGY  22  S e c t i o n o f Bowman c a p s u l e a n d avascular glomerular invagination, o f 19.8 mm a l e v i n . x 800  Figure  25  be  Bowman  capsule  gr  avascular  mt  myeloid  ns  n e c k segment o f t u b u l e  glomerulus  tissue  Transverse s e c t i o n o f kidney o f 26.9 mm f r y t o show b e g i n n i n g v a s c u l a r i t y o f glomerulus. x 700  c  capillary  gl  glomerulus  sd  segmental  t  tubule  containing  duct  erythrocyte  - 63a -  PLATE XII  EMBRYOLOGY  Figure 24  Transverse section of kidney of 28.58 mm f r y to show well-formed glomeruli. x 400  Figure 25  be  Bowman capsule  gl  glomerulus  meg  secondary mesonephric condensation  sd  segmental duct  t  tubule  Transverse section of kidney of 29.78 mm f r y to show secondary Bowman capsule. x 400  beg  secondary capsule  sd  segmental duct  t  primary tubule  t  9  secondary tubule  - 64a -  PIATE XIII  EMBRYOLOGY  Figure 26  Typical transverse section of kidney of 64 mm f i n g e r l i n g . x 200  Figure 27  a  arteriole  gl  glomerulus  pcv  post cardinal vein  sd  segmental duct  t  tubule  Longitudinal section of 20.9 mm a l e v i n to show bladder l i k e d i s tention of ducts. x 100  a  anus  b  bladder  g  gut  m  mesonephros  sd  segmental duct  sdc  common urinary duct  ugp  urogenital p a p i l l a  - 65a -  PLATE XIV  CORPUSCLE OP STANNIUS  Pigure 28  Longitudinal section of segmental duct i n 7.4 mm embryo to show rudiment of corpuscle of Stannius. x 800  Pigure 29  c  corpuscular rudiment  ep  epithelium of duct  1  lumen of duct  Transverse section of right lobe of kidney i n 9.54 mm embryo to show separation of corpuscle from duct. x 400  c  corpuscle  pcv  post cardinal vein  sd  segmental duct  - 66a -  PLATE XV  CORPUSCLE OP STANNIUS  Pigure 30  F r o n t a l section of corpuscle of Stannius i n 59 mm f i n g e r l i n g . x 100  c  capillary  cap  capsule  t  fibrous trabeculum  - 67a -  PLATE XVI  Figure  INTERRENAL  51  TISSUE  Transverse section of part o f p r o n e p h r o s i n 100 mm- f i n g e r l i n g t o show a r r a n g e m e n t o f i n t e r renal tissue. x 50  c  c a p i l l a r y net  it  interrenal  mt  myeloid  s  blood  tissue  tissue  sinusoid  

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