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Development and maturation of Philonema species (Nematoda: Philometridae) in salomid hosts with different… Bashirullah, Abul Kashem Mohammed 1966

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THE  DEVELOPMENT AND MATURATION OF PHILONEMA  SPECIES  (NEMATODA: PHILOMETRIDAE) IN SALMONID HOSTS WITH DIFFERENT L I F E HISTORIES  by ABUL KASHEM B.  MOHAMMED BASHIRULLAH  Sc. Dacca U n i v e r s i t y ,  1958  M. S c . D a c c a U n i v e r s i t y ,  1960  A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS OF THE DEGREE OF Doctor in  of Philosophy  t h e Department of Zoology  We a c c e p t t h i s t h e s i s required standard  The  University  as c o n f o r m i n g t o the  of B r i t i s h  A u g u s t , 1966  Columbia  i n presenting  t h i s thesis i n p a r t i a l f u l f i l m e n t of the requirements  for an advanced degree at the U n i v e r s i t y of B r i t i s h Columbia, I agree that the L i b r a r y s h a l l make i t f r e e l y available f o r reference and study,,  L further agree that permission f o r extensive copying of t h i s  thesis f o r s c h o l a r l y purposes may be granted by the Head of my Department or by h i s representatives.  I t i s understood that copying  or p u b l i c a t i o n of t h i s thesis f o r f i n a n c i a l gain s h a l l not be allowed without my written permission.  Department of  ZOOLOGY  The U n i v e r s i t y of B r i t i s h Columbia Vancouver 8 , Canada Date  August  §1,1966  The University of B r i t i s h Columbia FACULTY OF GRADUATE STUDIES PROGRAMME OF THE FINAL ORAL EXAMINATION FOR THE DEGREE OF DOCTOR OF PHILOSOPHY  of  ABUL KASHEM MOHAMMED BASHIRULLAH B.Sc., University of Dacca, 1958 M.Sc., University of Dacca, 1960 MONDAY, AUGUST 29, 1966 AT 3:30 P.M. IN ROOM 3332, BIOLOGY SCIENCES BUILDING COMMITTEE IN CHARGE Chairman:  W. Gage  J. R. Adams P. A. Dehnel L. Margolis External Examiner:  D. J . Randall W. B. Schofield J. D. McPhail Z. Kabata  Aberdeen Marine Laboratory Aberdeen, Scotland. Research Supervisor:  J . R. Adams  "The Development and Maturation of Philonema Species (Nematoda: Philometridae) i n Salmonid Hosts ' with Different L i f e Histories^." ABSTRACT This study was undertaken to determine the i d e n t i t y of Philonema oncorhynchi from anadromous sockeye and Philonema agubernaculum from non-anadromous trout, and also to test the hypothesis that the worm i n salmon i s dependent on hormonal stimulus from the host for synchronization of reproduction. On the basis of differences found i n l i f e cycles, cross i n f e c t i o n and starch gel electrophoresis, P. oncorhynchi and P. agubernaculum are considered to be d i f f e r e n t species. The hypothesis was tested experimentally, using salmon p i t u i t a r y extracts and synthetic s t i l b e s t r o l . P i t u i t a r y extracts accelerated the production of larvae i n the uterus, of the worm and s t i l b e s t r o l inhibited the gonadal development of the f i s h , but had no apparent effect on the worms. The hypothesis was further supported by the r e s u l t s of transplantation of adult but non-larvigerous worms from maturing sockeye into immature trout. Larval development f a i l e d to take place i n r e c i p i e n t hosts. As -well, larvigerous worms were c o l l e c t e d only from sexually mature f i s h , whether 3, 4 or 5 years old. Rapid development of the worm paralleled.the rapid gonadal.development of.-the f i s h during the last 6 months of the f i s h ' s life. Sexually immature sockeye had only immature worms.  GRADUATE STUDIES F i e l d of Study: Introductory  Zoology Parasitology  J. R. Adams  Advance Parasitology  J. R. Adams  Experimental Zoology  Wo S. Hoar  Quantitative Methods of Zoology Ecology Marine F i e l d Course  J. T. McFadden W. Murdoch P. A. Dehnel  Chairman:  P r o f e s s o r James R. Adams ABSTRACT  T h i s study was undertaken t o determine the i d e n t i t y o f Philonema oncorhynchi from anadromous sockeye and Philonema agubernaculum from non-anadromous t r o u t , and a l s o t o t e s t t h e h y p o t h e s i s t h a t the worm i n salmon i s dependent on hormonal stimulus  from the h o s t  On cross  f o r synchronization  o f reproduction.  the b a s i s o f d i f f e r e n c e s found i n t h e l i f e  cycles,  i n f e c t i o n and s t a r c h g e l e l e c t r o p h o r e s i s , Philonema  o n c o r h y n c h i and Philonema agubernaculum a r e c o n s i d e r e d different  species.  The h y p o t h e s i s was t e s t e d e x p e r i m e n t a l l y , salmon p i t u i t a r y e x t r a c t and s y n t h e t i c P i t u i t a r y extract accelerated the u t e r u s  t o be  stilbestrol.  the p r o d u c t i o n  o f the worm and s t i l b e s t r o l  using  of larvae i n  i n h i b i t e d the gonadal  development o f f i s h b u t had no apparent e f f e c t on t h e worms. The h y p o t h e s i s was f u r t h e r supported by t h e r e s u l t s of t r a n s p l a n t a t i o n of adult b u t non-larvigerous maturing sockeye i n t o immature t r o u t . failed  t o take p l a c e  worms from  L a r v a l development  i n the r e c i p i e n t host.  As w e l l , the  l a r v i g e r o u s worms were c o l l e c t e d o n l y from s e x u a l l y mature f i s h whether 3, 4 o r 5 years o l d .  Rapid development o f t h e  worm p a r a l l e l e d the r a p i d gonadal development o f t h e f i s h during  the l a s t 6 months o f the f i s h ' s l i f e .  immature sockeye had o n l y immature worms.  Sexually  TABLE O F CONTENTS PAGE GENERAL INTRODUCTION OUTLINE OF  THE  .  1  PROBLEM  2  SECTION I A c o m p a r i s o n o f P h i l o n e m a o n c o r h y n c h i i n n a t u r a l l y i n f e c t e d sockeye of C o i t u s Lake and P h i l o n e m a a g u b e r n a c u l u m i n e x p e r i m e n t a l l y infected trout 7 INTRODUCTION MATERIALS AND  . . METHODS  7  1.  Source of e x p e r i m e n t a l  fish  2.  Source of e x p e r i m e n t a l  copepods  3.  Source of L a r v a e of Philonema  4.  Infection  and m a i n t e n a n c e  5.  Infection  of t r o u t  6.  Fixation, o f worms  Staining  . . . .  3.  and  o f Copepods  . 10  Measurements 10  D e s c r i p t i o n of t h i r d stage larvae  and  11  fourth . . . . .  11  Developmental stages of Philonema from sockeye  13  Development in trout  16  o f P^  agubernaculum  . . . . . . . 1.  9 9  .  2.  DISCUSSION  7  10  RESULTS 1.  7  . . . . . . . .  I n f e c t i o n of j u v e n i l e sockeye under n a t u r a l c o n d i t i o n  2.  Route  3.  Growth  4.  Escapement  5.  V i s c e r a l adhesions  salmon 18  of i n f e c t i o n of the swimbladder and d e v e l o p m e n t  18  of Philonema  o f worm f r o m t h e h o s t  . 21  „ . . 23  . . . .  27 30  TABLE OF CONTENTS SUMMARY  (continued)  ,  SECTION I I  Experimental and salmon  . . . cross infection  between  strains  INTRODUCTION  trout  . . . . . . .  . . . . . . . . .  32 .  MATERIALS AND METHODS . . . . . . . . . . . . . . . . . 1. 2.  S o u r c e and h u s b a n d r y o f f i s h I n f e c t i o n and m a i n t e n a n c e o f experimental f i s h  32 32  . . . . . .  32 33  RESULTS  34 1. 2.  DISCUSSION SECTION  31  Infection f i s h with  o f d i f f e r e n t s a l m o n i d and g o l d Philonema from sockeye . . . .  I n f e c t i o n of sockeye, kokanee and g o l d f i s h  pink, s p r i n g , with t r o u t s t r a i n s  .  47  INTRODUCTION  47  MATERIALS AND METHODS  . . . . .  . . . . . . .  P r e p a r a t i o n o f salmon  47  pituitary  extracts 2.  36 33  I I I Hormonal t r e a t m e n t s  1.  34  43  Injection  o f salmon  a) Shuswap L a k e b) G r e a t  pituitary  fish  C e n t r a l Lake  3.  Injection  4.  Implantation  5.  Feeding  48  fish  49  of 1 7 - b - e s t r a d i o l of s t i l b e s t r o l  Shuswap Lake  2.  Great  fish  50 pills  . . .  50  powder  . . . . . 50  ,  51  . . . 1.  48  . . .  of s t i l b e s t r o l  RESULTS  extracts .  . . .  C e n t r a l Lake f i s h  a) E f f e c t  o f salmon  pituitary  b) E f f e c t  of s t i l b e s t r o l  51 . „  51  extracts .  51  .  53  iv TABLE OF CONTENTS  (Continued)  DISCUSSION  55  SECTION IV  Transplantation  o f worms  65  INTRODUCTION  65  MATERIALS AND METHODS  65  1.  Worms u s e d  for transplantation  .......  2.  Recipient  3.  Transplantation  4.  T r a n s p l a n t a t i o n o f l a r v a l worm i n t o f i s h  5.  T r a n s p l a n t a t i o n of a d u l t worms i n t o f r o g s . ,  hosts  66 of a d u l t  worms i n t o f i s h .  66 67  and l a r v a l 67  RESULTS  68 1.  T r a n s p l a n t a t i o n o f a d u l t P. o n c o r h y n c h i f r o m m a t u r e d s o c k e y e i n t o immature rainbow t r o u t  68  2 ( a ) . T r a n s p l a n t a t i o n of t h i r d stage l a r v a e i n t o the stomachs o f s t e e l h e a d , rainbow t r o u t and s o c k e y e s a l m o n  69  2 ( b ) . T r a n s p l a n t a t i o n of t h i r d i n t o body c a v i t y o f t r o u t  71  3.  DISCUSSION  65  stage  larvae  T r a n s p l a n t a t i o n of f o u r t h stage larvae i n t o t h e s t o m a c h and body c a v i t y o f trout  72  4.  Transfer  72  5.  T r a n s p l a n t a t i o n of a d u l t l a r v a e o f P^_ o n c o r h y n c h i  6.  T r a n s p l a n t a t i o n of f o u r t h stage larvae i n t o the abdominal c a v i t y of g o l d f i s h  of i n f e c t i o n  by p r e d a t i o n and f o u r t h into frogs  . . .  stage . . 73 .  73 74  V  TABLE OF CONTENTS  (Continued)  SECTION V D e v e l o p m e n t o f P h i l onema o n c o r h y n c h i i n naturally  infected  sockeye  o i Rivers  Inlet  .  79  INTRODUCTION  79  MATERIALS AND METHODS  80  1.  Collection Philonema  and measurement oncorhynchi  of adult  . . . .  81  RESULTS  82  DISCUSSION  85  GENERAL  91  SUMMARY AND CONCLUSIONS  LITERATURE APPENDIX  CITED  93 100  ERRATA p. 4  last line  p. 7  Para 5 l i n e 1  p. 13 p. 15  Para 3 l i n e 10 Para 2 l i n e 9  p. 20  Para i  p. 23 p. 27 p. 31  Para 3 l i n e 3 line 4 Para 2 l i n e 6  p. 34 p. 34  line 3 Para 2 l i n e 3  p. 49 p. 51 p. 53  line 8 line 1 Para 3 l i n e 1  p. 57  last line  p. 58 p. 75  Para 2 l i n e 4 l i n e 15  p. 77  line 1  p- 78 p. 79  l a s t para l i n e 1 and 3I Para 3 l i n e 8  p. 84  Para 2 l i n e 6  p. 88  Para 2 l i n e 2  p. 95  Ishida, T and e t a l , 1958  l i n e 10  variation in for variation from' insert "infection of trout with, e x p e r i m e n t a l l y " a f t e r experimental v'rs'ceral f o r v i s e r a l i n s e r t " i n diameter" a f t e r 5.5 cm' i n s e r t "the l i f e c y c l e " i n place of "that" i n s e r t " e i t h e r " a f t e r "due" read P h i l o m e t r a f o r Philonema read "mature o n l y " f o r 'is matured' read "uneaten" f o r 'unfed' i n s e r t "amount" a f t e r minimal' i n s e r t "were" a f t e r 'fish' i n s e r t "were" a f t e r 'fish' read "between 1 and 23 days" for'between 173 days' i n s e r t "According t o Gorshkov" b e f o r e 'ontogenesis' i n s e r t "occurs" a f t e r 'this' i n s e r t "Transplanted" b e f o r e 'fourth stage' and d e l e t e " t r a n s p l a n t e d " from l i n e 16 substitute "tissue" f o r 'epithelial' d e l e t e "had" s u b s t i t u t e " i s " f o r may be (merely)' i n s e r t "were c a l c u l a t e d a l s o by r e g r e s s i o n a g a i n s t s i z e of the worms" a f t e r 'worms' i n s e r t "The r e p r o d u c t i v e c y c l e " b e f o r e 'of oncorhynchi' i n s e r t "waters"  a f t e r 'offshore''  LIST OF FIGURES PAGE  FIGURE 1.  Development of Philonema oncorhynchi i n sockeye salmon of v a r i o u s ages from C u l t u s Lake  2.  Development of Philonema agubernaculum i n experiment a l l y infected trout 19  3.  Diagrammatic comparison of the spawning areas and the m i g r a t i o n s of t r o u t and salmon with those of P. agubernaculum and P. oncorhynchi i n t h e i r t r o u t and salmon h o s t s r e s p e c t i v e l y ~. . 26  4.  Development of P. oncorhynchi i n e x p e r i m e n t a l l y i n f e c t e d t r o u t , I T two years o l d  35  5.  Development of P. oncorhynchi from sockeye i n experimentally" i n f e c t e d t r o u t f r y .  37  6. 7.  8.  9.  17  salmon  Development of P. oncorhynchi from t r o u t i n e x p e r i m e n t a l l y i n f e c t e d f r y of sockeye salmon  . .  H y p o t h e t i c a l r e p r o d u c t i v e i s o l a t i o n of P. oncorhynchi and P_^ agubernaculum i n salmonids (based on 4 year r e p r o d u c t i v e c y c l e of sockeye salmon)  39  41  A comparison of s t a r c h g e l electropherograms of P. agubernaculum from t r o u t and I\_ oncorhynchi From sockeye salmon  46  E f f e c t of crude salmon p i t u i t a r y e x t r a c t s on the p r o d u c t i o n of l a r v a e i n P. oncorhynchi  52  10. E f f e c t of s y n t h e t i c s t i l b e s t r o l on the p r o d u c t i o n of l a r v a e i n P_^ oncorhynchi  54  11. E f f e c t of implanted s y n t h e t i c s t i l b e s t r o l on the p r o g r e s s i v e growth of t e s t e s of sockeye salmon from the Great C e n t r a l Lake  56  12. Development of P. oncorhynchi i n d i f f e r e n t p l a n t e d salmonids  70  trans-  vi FIGURE 13.  14.  15.  R e l a t i o n s h i p between the d e g r e e o f m a t u r a t i o n o f s o c k e y e salmon o f R i v e r s I n l e t and t i m e o f c a t c h  PAGE  . 83  R e l a t i o n s h i p between t h e o v a r y w e i g h t and t h e p r o d u c t i o n o f l a r v a e i n P. o n c o r h y n c h i f r o m s o c k e y e salmon o f R i v e r s I n l e t '. . '. 7  86  R e l a t i o n s h i p between the t e s t i s w e i g h t of f i s h and t h e p r o d u c t i o n o f l a r v a e i n P. o n c o r h y n c h i f r o m s o c k e y e salmon o f R i v e r s I n l e t  87  L I S T OF TABLES TABLE 1.  2.  3.  4.  PAGE Stages of Philonema Lake s o c k e y e s a l m o n  oncorhynchi found o f v a r i o u s ages  i n Cultus 100  Measurements o f P h i l o n e m a o n c o r h y n c h i o f s o c k e y e salmon o f d i f f e r e n t ages f r o m C u l t u s L a k e . . .  101  Development of Philonema mentally infected trout  103  agubernaculum  i n experi-  E f f e c t o f 1 7 - b - e s t r a d i o l , salmon p i t u i t a r y e x t r a c t s and s t i l b e s t r o l on P h i l o n e m a s p e c i e s i n immature 2 y e a r o l d s o c k e y e f r o m Shuswap Lake .  104  E f f e c t on p i t u i t a r y e x t r a c t s on f e m a l e P h i l o n e m a r e c o v e r e d from the t r e a t e d f i s h . . . ~ ". '. T  105  6.  -Effect  106  7.  E f f e c t o f s t i l b e s t r o l on f e m a l e ed from the i m p l a n t e d f i s h  5.  8. 9.  o f salmon  pituitary  on s o c k e y e  salmon  Philonema  . .  recover107  E f f e c t o f i m p l a n t e d s t i l b e s t r o l on t h e t e s t e s o f " p r e s p a w n i n g s o c k e y e f r o m G r e a t C e n t r a l Lake . .  108  R e c o v e r y o f P h i l o n e m a o n c o r h y n c h i f r o m t h e body c a v i t y of t r a n s p l a n t e d t r o u t .  109  10.  Recovery  111  11.  R e c o v e r y o f t r a n s p l a n t e d a d u l t and f o u r t h l a r v a e f r o m t h e body c a v i t y o f f r o g s  12.  of t r a n s p l a n t e d  worms f r o m t h e s a l m o i d s stage  Summary o f m e r i s t i c measurements o f f i s h o f d i f f e r e n t age c o m p o s i t i o n o f R i v e r s I n l e t , . .  112 113  13.  Summary o f m e r i s t i c measurements o f P i " b n c d r h y n c h i o f " d i f f e r e n t age c o m p o s i t i o n o f R i v e r s I n l e t . . 114  14.  Measurements o f Philonema R i v e r s I n l e t Sockeye '. '.  15.  R e s u l t s of r e g r e s s i o n time caught  oncorhynchi  from 115  o f worms on age o f f i s h and 116  ACKNOWLEDGMENT The for  author i s gr> a t e f u l t o P r o f e s s o r James R,. Adams  h i s s u p e r v i s i o n , f o r h i s unbiased c r i t i c i s m , , and f o r h i s  h e l p f u l s u g g e s t i o n s i n the p r e p a r a t i o n of t h i s manuscript* S i n c e r e g r a t i t u d e i s e s p e c i a l l y extended of  t o Dr. Leo M a r g o l i s  the F i s h e r i e s Research Board of Canada, Nanaimo, from whom  the author always r e c e i v e d s t i m u l a t i n g encouragement, who allowed f r e e access t o unpublished data,, and who helped i n many more ways than the author can put i n t o words.  Many  t e c h n i c a l d i f f i c u l t i e s d u r i n g the e a r l y phase of t h i s work were overcome and new methods developed through d i s c u s s i o n with Dr. G. G. Gibson. his  critical  The author wishes t o express  a p p r e c i a t i o n t o Drs. C. C. L i n d s a y , P. A. Dehnel, I . A.  E f f o r d , and W. B. S c h o f i e l d f o r t h e i r c o n s t r u c t i v e suggestions i n the p r e p a r a t i o n of t h i s The  thesis.  author wishes t o thank P r o f e s s o r W. S. Hoar f o r  providing synthetic s t i l b e s t r o l ,  Dr. M. Smith f o r s u p p l y i n g  crude salmon p i t u i t a r y e x t r a c t s , and Dr. H. T s y u k i and Mrs. E. Roberts f o r a s s i s t a n c e with the e l e c t r o p h o r e s i s .  Dr. D. J .  R a n d a l l who was always prepared t o supply experimental and a d v i c e , i s acknowledged with much a p p r e c i a t i o n .  fish  The  author wishes t o thank Mr. Melvin Weisbert f o r s u p p l y i n g h a t c h e r y - r e a r e d salmonids. i n the s t a t i s t i c a l extended of  The a s s i s t a n c e of Dr. J . T. McFadden  a n a l y s i s i s acknowledged.  t o Dr. M. Shepgard  Thanks are  of the F i s h e r i e s Research  Canada, Nanaimo, f o r h i s i n t e r e s t i n Philonema  s u g g e s t i n g the problem  i n sockeye  Board  and f o r  salmon of R i v e r s I n l e t .  Mr. T. Bq:lton, Mr. N. Boyce, personnel of the F i s h C u l t u r e Room, and many o t h e r s of the B i o l o g i c a l S t a t i o n p r o v i d e d a s s i s t a n c e d u r i n g the l a t e r phase of t h i s study at Nanaimo. The  author acknowledges the c o n t r i b u t i o n s of the f o l l o w i n g  o r g a n i z a t i o n s who made d i f f e r e n t k i n d s of f i s h a v a i l a b l e f o r t h i s study:  The B r i t i s h Columbia  F i s h and Game Branch,  I n t e r n a t i o n a l North P a c i f i c Salmon Commission, Department of F i s h e r i e s , B i o l o g i c a l S t a t i o n of the F i s h e r i e s Research  Board  df Canada at Nanaimo, and the B r i t i s h Columbia  Council.  Research  The author extends h i s s i n c e r e a p p r e c i a t i o n t o a l l those who helped him i n the f i e l d ,  and e s p e c i a l l y t o Mr. James  Adams, Mr. Gordon Halsey, and Mr. B i l l  Woodall.  Mrs. Z. J u r i c i c  took care of experimental f i s h , and her a s s i s t a n c e i n the l a b o r a t o r y i s acknowledged. Acknowledged with thanks i s the f i n a n c i a l  assistance  p r o v i d e d by the Canadian Commonwealth S c h o l a r s h i p and Fellowss h i p Committee. Thanks are due t o a l l those who have given the author h e l p and advice d u r i n g the course of t h i s study. The author wishes t o thank Mrs. R. C o l l i s f o r her e x c e l l e n t t y p i n g of t h i s  manuscript.  F i n a l l y , the author i s indebted t o P r o f e s s o r H. K. Y o s u f z a i f o r h i s encouragement.  GENERAL INTRODUCTION  The  literature  demonstrations  of P a r a s i t o l o g y c o n t a i n s s e v e r a l  or suggestions  that the c o r r e l a t i o n  observed  between t h e r e p r o d u c t i o n o f a p a r a s i t e and t h e s e a s o n a l reproduction to  of i t s host  t h e h o s t s ' hormones  Miretski,  the coelomic  parasite life,  (Smyth & M o f t y ,  of the p a r a s i t e  1964; B a r r o w , 1962;  1951; R o t h s c h i l d , 1 9 6 4 a ) . A parasitic  in  i s due t o a r e s p o n s e  cavity  infects  very slowly.  the maturation  salmon c o n t a i n r i p e  char,  during their Final  early  host.  to release their  larvae.  and w h i t e f i s h a r e a l s o  in  salmon.  of  Philonema with  t h e worm i n salmon host  was u n d e r t a k e n  particular  fresh-water  Spawning  a P h i l o n e m a w h i c h may o r may n o t be c o n s p e c i f i c  study  parasitized with  that  t o e l u c i d a t e the l i f e  reference t o the hypothesis  i s dependent  The  o f t h e worm  by  This  i s found  salmon.  maturation  of the f i s h  worms r e a d y  Non-anadromous t r o u t ,  oncorhynchi  o f t h e anadromous s o c k e y e  t h e young f i s h  but develops  coincides with  nematode, P h i l o n e m a  cycle that  on h o r m o n a l s t i m u l u s f r o m t h e  f o r s y n c h r o n i z a t i o n of reproduction.  Outline  of the Problem Kuitunen-Ekbaum  from the abdominal c a v i t y E n g l i s h Bay, B r i t i s h same s p e c i e s  (1933) d e s c r i b e d P h i l o n e m a o n c o r h y n c h i of adult  Columbia.  Smedley  o f worm f r o m t h e body  from C u l t u s Lake, B r i t i s h  another s p e c i e s ,  fontinalis  ( J o r d a n ) w h i c h were c o l l e c t e d this  cavity  of sockeye  Simon and Simon  and m u s c l e s o f t h e a b d o m i n a l w a l l  erected  (1933) d e s c r i b e d t h e  Philonema agubernaculum  Salvelinus  (1936)  of a n t e r i o r  Platzer  salmon  of Prosopium  (Mitchell)  cavity  williamsoni  and Salmo  i n Wyoming N a t i o n a l  to posterior  described  f r o m t h e body  new s p e c i e s m a i n l y on t h e b a s i s  and t h e r a t i o  nerka from  Columbia.  Three years l a t e r ,  (Girard),  Oncorhynchus  shasta  Forest.  of smaller  They  size  oesophagus.  and Adams ( i n p r e s s ) worked  out the l i f e  cycle  of P h i l o n e m a o n c o r h y n c h i and f o u n d C y c l o p s b i c u s p j d a t u s was intermediate larvae occur  host.  are the i n f e c t i v e  the  fresh  which  migration. water.  cycle  Life  Adult  O n l y t h o s e worms  spawns c a n p e r p e t u a t e t h e s p e c i e s  on t h e o t h e r hand h a s been  i n which f i s h cycle  only  t h e worms must a d j u s t t o  of the f i s h . i  a r e g r a v i d when t h e f i s h  localities  molts  o f P. o n c o r h y n c h i o c c u r s  water, i t i s obvious that  P. a g u b e r n a c u l u m  Two f u r t h e r  T h i r d stage  i  the transmission  long reproductive  i n the copepod.  stage f o r f i s h .  i n the salmon. Since  in  Two m o l t s o c c u r  an  reported  from  h o s t s a r e u n a b l e t o make a s e a w a r d  must be c o m p l e t e d e n t i r e l y  P^ a g u b e r n a c u l u m  have  than a year o l d ( T a y l o r , u n p u b l i s h e d  been  i n fresh  found i n t r o u t  data).  less  3 Platzer  (1964) made a m o r p h o m e t r i c  a n a l y s i s of  P h i l o n e m a t a k e n f r o m s a l m o n o i d s i n two g e o g r a p h i c a l l y a r e a s , C u l t u s and K o o t e n a y L a k e s , B r i t i s h Lake  contained  species.  Columbia.  anadromous f i s h ; K o o t e n a y L a k e  Adult  only  separated Cultus  landlocked  worms t a k e n f r o m w h i t e f i s h , P r o s o p i u m  w i l l i a m s o n i , D o l l y V a r d e n , S a l v e l i n u s malma and kamloops Salmo g a i r d n e r i , salmon. ring,  were g e n e r a l l y  Comparisons  of t o t a l  m u s c u l a r and g l a n d u l a r  length  revealed  aspects  taxonomically  body  length,  distance  tail  Platzer  morphological  morphometric  d i d n o t f i n d any differences  specimens o f Philonema o n c o r h y n c h i Kuitunen-Ekbaum Lake  and t h e t y p e s p e c i m e n  and Simon. separated  on m o r p h o l o g i c a l Seven  other  to  d a t e on t h e b a s i s  to  glandular  1937;  Fujita,  of t o t a l  p a r e d them w i t h  body  length  Simon and Simon,  been  and r a t i o  Simon  that  Philonema e l o n g a t a  (Richardson  Philonema Fujita  B a u e r were synonyms o f P h i l o n e m a  1965).  Oncorhynchus  l e u c o m a e n i s and com-  t h a t were d e s c r i b e d  He s t a t e d  of muscular  1960 and Rumyantsev,  and S a l v e l i n u s  the s p e c i e s  described  and f e m a l e worms  (1955) s t u d i e d P h i l o n e m a f r o m  and Simon and Simon.  Cultus  c o u l d n o t be  o f P h i l o n e m a have  i n b o t h male  keta,  from  agubernaculum  t h e two s p e c i e s  1940; B a u e r , 1946; Meyer,  nerka , Oncorhynchus  between  grounds,  species  oesophagus  Akhmerov  sibirica  of Philonema  He c o n c l u d e d t h a t  t o nerve  and s p i c u l e  v a r i a b i l i t y of those  Philonema.  significant  than those from sockeye  oesophagus,  a tremendous  i n t h e genus  smaller  trout,  by F u j i t a ,  Bauer  agubernaculum  and Coregonema oncorhynchi Kuitunen  4 Ekbaum. P.  He  d i d not  tenuicaUda,  since  the  P.  kondai,  difference noted  Akhmerov  their  considered  and  available  oncorhynchi  P.  to  (1955) d i d n o t  h o s t s and  therefore that  other four s p e c i e s ,  salvelini,  between P h i l o n e m a  that  proposed  P.  d e s c r i p t i o n s were n o t Although  he  synoriomize F u j i t a ' s  ochotense  him.  find  and  any  morphological  Coregonema  sibirica,  t h e i r e c o l o g y were d i f f e r e n t .  t h e nematode Coregonema s i b i r i c a  as a s u b s p e c i e s of P.  and  be  named P h i l o n e m a  oncorhynchi  sibirica.  and  (1960) a c c e p t e d Akhmerov's synonyms b u t  a contrary  formed, by Rumyantsev  the  Shulman  opinion  was  Spasskii  oncorhynchi  He  e t a l (1958, 1959)  (1965) who  reduced  Coregonema B a u e r , 19^46  i n t o a synonym o f t h e  K u i t u n e n - E k b a u m , 1933,  and  established  (synonym Coregonema s i b i r i c a ) stated  that Philonema  a g u b e r n a c u l u m , and t o be  and  i n North be  of the  differences America  biologically  However, two  independent  that  on  i n the b i o l o g y of h o s t s .  s p e c i e s can  from  to  He P.  t h e y may  i n d i c a t e s both  genus P h i l o n e m a m a i n l y  distinct  species.  prove  1965).  of the l i t e r a t u r e  the Philonema  sibirica  shows some s i m i l a r i t y  t h e same s p e c i e s (Rumyantsev,  splitting  size  sibirica  genus  genus P h i l o n e m a  Philonema  considered i t possible  This review and  as an  and S t r e l k o v  the Both  lumping  grounds of in Asia  anadromous s a l m o n a p p e a r  and to  f r o m thostf.< of non-anadromous h o s t s . n o t be  s e p a r a t e d by  morphological  criteria. Reported cycle  could result  differences  in size  from v a r i a t i o n  from  and  p a t t e r n of  the  response  life  of a  single  5  s p e c i e s of Philonema  t o the environments  p h y s i o l o g i c a l l y different hosts.  p r o v i d e d by the  The d i f f e r e n c e s might  equally  be due t o the e x i s t e n c e of d i f f e r e n t b i o l o g i c a l s p e c i e s which are g e n e t i c a l l y  distinct.  The presence of g r a v i d worms o n l y i n s e x u a l l y maturing sockeye salmon ( P l a t z e r , 1964), but i n t r o u t as young as one year ( T a y l o r , unpublished d a t a ) , suggest the following hypotheses: i ) That there are two p h y s i o l o g i c a l l y s t r a i n s of Philonema biological  different  which may deserve the s t a t u s of d i f f e r e n t  species, i i ) that the form i n t r o u t  ( P ^ agubernaculum)has  a  developmental c y c l e of a year or l e s s i n l e n g t h and i s i n d i f f e r ent t o the hormonal changes i n i t s h o s t s , iii)  that the form i n salmon (P. oncorhynchi) has a  l o n g developmental p e r i o d dependent  on hormonal changes i n i t s  h o s t s f o r s y n c h r o n i z a t i o n of i t s r e p r o d u c t i o n process t o t h a t of the salmon. In support of these hypotheses f i v e l i n e s of i n v e s t i g a t i o n s w i l l be presented. i ) A comparison  of the developmental r a t e and maturation  of the two s t r a i n s i n t h e i r normal h o s t s , i i ) the r e s u l t s of i n f e c t i n g salmon with the t r o u t strain  (Pj_ agubernaculum) and i n t r o u t with the salmon s t r a i n  (P. o n c o r h y n c h i ) , iii)  the e f f e c t on the worm i n salmon of treatments of  the hosts with p i t u i t a r y e x t r a c t s and gonadal hormones,  6 iv) normal a d u l t v) oh  Philonema  the e f f e c t  on  t h e worm o f t r a n s p l a n t a t i o n  hosts to other salmonids a study  and  abnormal h o s t s ,  of t h e p a t t e r n o f d e v e l o p m e n t  in a natural  w h i c h mature v a r i o u s l y Each of these  from  and  maturation  p o p u l a t i o n o f salmon c o m p r i s i n g  in their  3rd, 4th  i s presented  or 5th  inaseparate  years. section.  fish  7 SECTION I .  A comparison of the development oncorhynchi  in naturally  and o f P h i l o n e m a  infected  agubernaculum  of Philonema  sockeye of C u l t u s  i n experimentally  Lake,  infected  trout.  INTRODUCTION Platzer development  and Adams  ( i n press) described  o f P^ o n c o r h y n c h i i n e x p e r i m e n t a l l y  sockeye  salmon.  They showed t h a t  inhabit  the t i s s u e  of the swimbladder.  were f o u n d i n t h e body c a v i t y after  23 months.  because  development  greater  precision  sockeye  salmon m a i n t a i n e d i n s a l t  of  stage  larvae  infected  rate  larvae fish  of development  of the i n f r e q u e n c y  i n the f i s h  infected  Fourth stage  of n a t u r a l l y  However, t h e p r e c i s e  c o u l d n o t be s t a t e d The  the t h i r d  the general  of sampling.  i s reported  b a s e d on s a m p l i n g o f n a t u r a l l y  here  with  infected  w a t e r , some f o r a p e r i o d  over 2 y e a r s . The  n e v e r been  reported.  successfully agubernaculum young  fish  development  infected  Meyer  i n t r o u t has  (1958, 1960) and V i k (1964)  Cyclops species with  b u t were u n a b l e t o t r a n s m i t  larvae  o f P.  the p a r a s i t e t o  host. This section  Cyclops  o f P^ a g u b e r n a c u l u m  and d e s c r i b e s  also reports  the experimental  the development  infected  o f I\_ a g u b e r n a c u l u m  8 over  the p e r i o d The  hosts  o f 182 d a y s w h i c h t h e f i s h  development  survived.  o f t h e two f o r m s i n t h e i r  normal  i s compared.  MATERIALS AND METHODS Source  of Experimental F i s h a)  Downstream m i g r a n t s o c k e y e s a l m o n were  from C u l t u s Lake ined  at this  i n May, 1965. A l l one y e a r o l d f i s h ,  t i m e , were i n f e c t e d  Philonema o n c o r h y n c h i . water  and e x a m i n e d  the development b) C u l t u s Lake  obtained  with  third  exam-  stage larvae of  The s m o l t s were m a i n t a i n e d i n s e a  infrequently until  June  1966, t o d e t e r m i n e  of Philonema.  Three year o l d n a t u r a l l y - i n f e c t e d maintained i n s a l t  water s i n c e  sockeye  capture  from  a s down-  s t r e a m m i g r a n t s i n 1963. c)  Wild  adult  p r e c o c i o u s m a l e s , and p r e - s p a w n i n g  f o u r y e a r o l d male and female- s o c k e y e s a l m o n were from Sweltzer Creek a t C u l t u s Lake, B r i t i s h provided data  on t h e f i n a l  captured  Columbia.  s t a g e s o f development  They  o f P.  oncorhynchi. d)  Y e a r l i n g h a t c h e r y - r e a r e d rainbow t r o u t  obtained  f r o m t h e Summerland H a t c h e r y o f B. C. F i s h  Branch.  T h e y were u s e d f o r e x p e r i m e n t a l  agubernaculum.  infection  were and Game  w i t h P.  9 Source  o f E x p e r i m e n t a l Copepods Plankton  was  Cyclops bicuspidatus those  of P l a t z e r  stant t  temperature  Source a)  Creek  sockeye fish  were i s o l a t e d by methods s i m i l a r t o  o o f 10 C as s t o c k c u l t u r e of  for infection.  Philonema  gravid salmon  trap  f e m a l e s were o b t a i n e d g e n e r a l l y (Oncorhynchus  nerka)  at C u l t u s Lake, B r i t i s h  Only g r a v i d  Columbia.  females c o n t a i n i n g  moving f i r s t  i n t h e u t e r u s were s e l e c t e d f o r t h e c o l l e c t i o n  larvae  to infect  burst  at b)  a minute  larvae  t h r o u g h #12 the  Philonema  easily.  The  a gill  net.  Columbia  Philonema  water. The  f e m a l e s were  stage larvae  larvae  of  f e m a l e s were  lake  Worms  water  con-  filtered  passed  were k e p t  through  in a refrigerator  before i n f e c t i o n .  rainbow  t r o u t , Salmo g a i r d n e r i o f t h i r t e e n  c o l l e c t e d from T r o u t Lake, B r i t i s h The  fish  was  brought  i n an i c e p a c k e d  same p r o c e d u r e was for  First  stage  agubernaculum: i  A female  British  gravid  the l a r v a e .  of g r a v i d  bolting silk.  a p p r o x i m a t e l y 8°C  using  to five  to release  and d e b r i s  bolting silk  pounds was  One  i n t o a f i n g e r bowl c o n t a i n i n g  within  taining  copepods.  from  from the S w e l t z e r  larvae  transferred  con-  oncorhynchi:  Fully female  at C u l t u s Lake.  (1964) and m a i n t a i n e d i n a room w i t h a  of L a r v a e  Philonema  c o l l e c t e d i n S m i t h Bay  used  to collect  oncorhynchi.  cooler  Columbia  by  t o the U n i v e r s i t y of on May  the f i r s t  27,  1965.  stage larvae  The as  10 Infection  and M a i n t e n a n c e Approximately  out of  of a stock four  1600 for  inch  E a c h d i s h was i n o c u l a t e d  larvae  and p l a c e d  or f o r t y - e i g h t hours.  dish  and c u l t u r e d  at 10°C.  capsule.  transferred  t o complete within a  with  t o an 8 i n c h i n copepods  t h i s operation  into a  small  t o a #5 g e l a t i n  The c a p s u l e was p u s h e d t h r o u g h  a glass  fish.  as the c a p s u l e  tube  inserted  I t t o o k two p e o p l e becomes s o f t and  minute. Fish  the f i s h  Fixation,  interval  i n 37 d a y s .  i n t o t h e stomach o f an a n e s t h e t i z e d  were t a g g e d  and t r a n s f e r r e d  were f e d s t a n d a r d wet f o o d  Staining, Live  formalin-acetic ethanol  Larvae  c o p e p o d s were c o n c e n t r a t e d  volume o f w a t e r and q u i c k l y  All  The l a t t e r  of Trout Fifty  wet  i n the r e f r i g e r a t o r  c o p e p o d s were t r a n s f e r r e d  i n f e c t i v e 3rd stage  Infection  with  per copepod.  Infected  reached  filtered  t o p r o d u c e 80 t o 1 0 0 % i n f e c t i o n o f c o p e p o d s  2 larvae  stacking  h u n d r e d c o p e p o d s were  a #12 n e t i n t o e a c h o f a s e r i e s  f i n g e r bowls.  twenty-four  about  four  j a r through  t o 2000 P h i l o n e m a  was f o u n d  o f Copepods  tanks.  e v e r y a l t e r n a t e day.  and Measurements o f Worms,  larvae  and a d u l t  P h i l o n e m a were f i x e d i n  a c i d and p r e s e r v e d  w h e r e a s worms f r o m f r o z e n  were t r a n s f e r r e d  into holding  directly  i n e i t h e r FAA o r 7 0 % and f o r m a l i n  t o 80% e t h a n o l .  preserved  fish  11 For  t h e s t a i n i n g and measurement  i n g m o d i f i c a t i o n s o f Goodey's Platzer's  (1964) t e c h n i q u e s 1)  2)  larvae,  ( F r a n k l i n & Goodey, 1949) and were made:  A d r o p o r two o f 0.0005% c o t t o n  p h e n o l was p l a c e d  analytical  of l a r v a e ^ f o l l o w -  on a c l e a n  Glass  filter  L a r v a e were  wool, g l a s s r o d or a s m a l l  added.  piece of  i n the drop as support  of the  for a coverslip.  The c o v e r s l i p was a p p l i e d and s e a l e d w i t h  mastix p a r a f f i n  (50 p . c . p a r a f f i n  plus  50 p . c . gum  L a r v a e were measured u n d e r a compound using  i n lacto-  p a p e r , d e p e n d i n g on t h e t h i c k n e s s  were p l a c e d 3)  slide.  blue  an o c u l a r m i c r o m e t e r ;  measured by u s i n g c v s c a l e  gum  mastix).  microscope  somewhat l a r g e r male worms were  projector; very  were m e a s u r e d u n d e r a b i n o c u l a r  l a r g e female  worms  microscope.  RESULTS Description Third  o f T h i r d and F o u r t h  oncorhynchi  (1966) d e s c r i b e d  from f i s h  third  stage  w h i c h were e x p e r i m e n t a l l y  These l a r v a e  are morphologically  experimentally  from those The  intestinal  of Philonema  i n copepods.  t o those  i n f e c t e d f o r 55 d a y s .  taken  Third  sockeye d i f f e r  stage only  i n copepods.  body i s f i l i f o r m  mean body l e n g t h  larvae  reared  identical  t a k e n f r o m downstream m i g r a t i n g  slightly  The  Larvae  Stage: Ko  larvae  Stage  i n shape, t a p e r i n g  i s 1.16 mm,;  j u n c t i o n i s 16.66 / u .  body w i d t h The c u t i c l e  posteriorly.  a t the oesophageal i s d i l a t e d at  12 the  buccal  region  has  rounded s h o u l d e r s  is  104.3  /u  is  248.9 /u  developed The  from the long,  i n t o the  lip-like  intestine.  a fine  structures.  i t s tapered  a n t e r i o r end. and  the  tip.  The  The  glandular  and  The  intestine.  and  tube c o n n e c t i n g  of two  n a r r o w lumen and  t o the  exterior.  well  posteriorly. pro-  genital  anterior portion  r e c t u m i s made up  ring  oesophagus  The  tail  oesophagus  i s well developed,  m o s t l y a t the  thick w a l l s  The  nerve  muscular  l o n g ) , g r a d u a l l y expanding  lumen o f t h e  i s oval  region with  of  regions:  anterior  posterior region,  Rectal  glands  are  developed. The  the  near  slender  (480.6 yu  primordium  well  two  o e s o p h a g e a l — i n t e s t i n a l valve  truding  the  forming  third  stage 1)  f r o m the slender  third  i n the  stage  prior  well  to molting  differs  from  f o l l o w i n g ways:  Mean body l e n g t h ,  a n t e r i o r end; and  773.6 /u  late  muscular  separated  3.32  mm.;  nerve r i n g ,  o e s o p h a g u s , 312.7  from the  grandular  /u  143.3 long,  oesophagus,  long. 2)  Cuticle dilated  rounded head without 3)  at b u c c a l  region  to  produce  lips.  Oesophageal--intestinal  valve  becomes more  prominent. Fourth  Stage: The  of the  f o u r t h stage 1)  4.1  mm.  f o l l o w i n g a d d i t i o n i s made t o t h e d e s c r i p t i o n  The  l a r v a e by  f o u r t h stage  i n l e n g t h , the  P l a t z e r and i n the  nerve r i n g  149  Adams ( i n p r e s s ) :  swimbladder  averaged  /u  anterior  from the  13 end,  muscular  881.6  /u  2)  A n t e r i o r end  3)  Tail  4)  Sex  size  (5.22  and  stage  - 9.5  Stages  larvae undifferentiated  mm.)  1965,  larvae  stage  show r u d i m e n t a r y  sampling  of Philonema  larvae  from  oncorhynchi fourth  stage l a r v a  One  female  s u b - a d u l t and  the p e r i t o n e a l sampling  was  were t w e n t y  cavity  had  third, and  late  months o l d .  larvae  a fourth  s t a g e s o f P.  third,  fourth  of  stage  and larvae  Philonema of s m o l t s  i n t o t h e body  stage  larva  January,  Some male and  1966, female  i n t h e body c a v i t y  i n t h e body c a v i t y  mm. and  By  and  cavity.  were f o u n d i n  a month l a t e r .  grown t o a l e n g t h o f 89.9  The  However,  (Table I . ) .  migrated  impossible u n t i l  worms were f o u n d appeared.  a few  of a f i s h  o n c o r h y n c h i were f o u n d  worm had  s m o l t s showed o n l y  showed g r o w t h i n t h e s w i m b l a d d e r  one  sexual characters.  i n the swimbladder.  s u b - a d u l t s i n t h e body c a v i t y  of  Sockeye  of y e a r l i n g  i n the swimbladder  1965,  while i n  Those i n t h e body c a v i t y  of Philonema  By A u g u s t ,  P.  rounded.  y e a r o l d downstream m i g r a n t s had  fourth  g l a n d u l a r oesophagus,  changed t o p o i n t e d t i p . of the  I n May,  two  bluntly  of the swimbladder.  Developmental  third  and  long.  the t i s s u e larger  o e s o p h a g u s , 329.8 /u  Further  when t h e  fish  sub-adult and  one  April,  viseral  female  1966,  more  adhesions  m a j o r i t y o f t h e worms p r e s e n t were male..  o n c o r h y n c h i w i t h the e x c e p t i o n of a d u l t  All  females  14 were f o u n d visceral sex.  in fish  adhesion  when e x a m i n e d i n J u n e , was o b s e r v e d  F r e s h l y encased  1966.  i n these f i s h  More  r e g a r d l e s s of  male worms were f o u n d when t h e a d h e s i o n  was t e a s e d a p a r t . D u r i n g t h e same p e r i o d , tanks  i n the l a b o r a t o r y -  fish  grew f r o m  maintained  an a v e r a g e  8.2 cm. t o 20.8 cm. and i n c r e a s e d i n w e i g h t  i n seawater  f o r k l e n g t h of  from  an  average  8.1 gm. t o 88.8 gm. Three as s m o l t s  immature  sockeye  i n 1963 and m a i n t a i n e d  laboratory until  o n c o r h y n c h i were f o u n d stage  larvae  were f o u n d  adhesions  encased  worms were f o u n d  adhered  mass.  adult  and a d u l t  cavity. larvae  stage  i n the  i n loose fibrous  larvae  tanks  i n the  date.  Philonema  of these  fish.  swimbladders. and dead and tissue  were e x a m i n e d  of the  i n October,  o c c u r r e d i n the swimbladder;  number o f 335 f o u r t h  i n the swimbladder  stage  o f one o f t h e s e  A month l a t e r  (November, 1965) most o f t h e l a r v a e h a d  migrated  the swimbladder  from  sub-  o h c o r h y n c h i were p r e s e n t i n t h e body  An e s t i m a t e d maximum were f o u n d  males of  were p r e s e n t i n t h e f i s h  P r e c o c i o u s male s o c k e y e fourth  i n seawater  i n t h e body c a v i t y  Viseral  1965,  from C u l t u s Lake  May, 1965 were e x a m i n e d on t h a t  F o u r t h s t a g e , s u b - a d u l t and a d u l t  No t h i r d  collected  fish  when c o l l e c t e d  than  female  from  i n t o t h e body c a v i t y  the spawning ground.  worms o c c u r r e d i n t h e body c a v i t y .  fish.  of the  More male During the  15 study three gravid  female  worms were f o u n d  A few s u b - a d u l t male worms were c o l l e c t e d Visceral c o c i o u s males. absent The  The i n t e n s i t y  t o heavy i n October  adhesions  caeca.  viseral  from  the  cavity.  swimbladder.  were p r e s e n t i n 8 o f 10 p r e o f a d h e s i o n was s c o r e d a s f r o m  and a b s e n t  occurred primarily  T h i n s t r a n d s of t i s s u e  abdominal  to light  i n November.  i n the r e g i o n of the p y l o r i c  were f o u n d  t o bridge the  c a v i t y when t h e two body w a l l s o f f i s h c o n t a i n i n g  a d h e s i o n were t e a s e d a p a r t . In  encysted taken  adhesions  i n t h e body  the f i s h  female  e x a m i n e d i n November, 1965, f r e s h l y  worms were f o u n d .  out of the incomplete  with whitish  fluid.  Live  cysts.  Thin fibrous  females  c o u l d be  F r e s h c y s t s were layers  of t i s s u e  filled  were  found  i  from and  t h e e n d o f t h e p e r i c a r d i u m t o t h e e n d o f t h e stomach this  layer  of t i s s u e  c o u l d be t a k e n  out s e p a r a t e l y .  c y s t s w i t h u n r e c o g n i z a b l e worms were f o u n d liver,  t o t h e body w a l l  or free  Many  attached t o the  i n t h e body c a v i t y .  Cysts  measured f r o m 0.5 t o 5.5 cm. Larvae (adult in  spawning f i s h )  the swimbladder Gravid  in 10%  were n o t g e n e r a l l y  female  than  b u t one f o u r t h  female  Philonema  larvae  swimbladder, was f o u n d  .  o n c o r h y n c h i were more common  In female  worms h a r b o u r e d  i n the  stage  o f a 4g male s o c k e y e .  i n male f i s h .  of the adult  found  fish,  i n September,  were g r a v i d whereas i n  November o f t h e same y e a r , 9 0 % o f t h e a d u l t s h a r b o u r e d gravid.  were  16 No v i s c e r a l adhesions were observed  i n spawning  female f i s h b u t v e r y l i g h t adhesions were observed i n some spawning male f i s h .  Encysted worms o c c u r r e d i n both_male and  female f i s h , b u t were more common i n the males. Measurements o f worms were taken and t h e i r growth i s p r e s e n t e d i n Table I I and F i g u r e I . Development o f Philonema  agubernaculum i n t r o u t  T h i r t y - t w o slow maturing, h a t c h e r y - r e a r e d rainbow t r o u t , Salmo g a i r d n e r i , o n e year o l d , were dosed w i t h 50 i n f e c t e d copepods i n a g e l a t i n c a p s u l e .  Three o f t h e f i s h  d i e d on the day o f i n f e c t i o n , u n d i g e s t e d copepods and l a r v a e of  Philonema were found i n t h e stomach o f these f i s h b u t no  l a r v a e were found i n t h e t i s s u e o f the swimbladder.  Eight  o t h e r f i s h were found t o be u n i n f e c t e d when examined l a t e r i n the experiment. No growth o f l a r v a e o c c u r r e d i n the f i r s t 30 days of  infection.  The f o u r t h stage l a r v a e were recovered from  the body c a v i t y o f one f i s h on the 58th day a f t e r  infection.  A f t e r t h i s time, t h i r d stage l a r v a e were found i n the swimbladder o f o n l y f i v e f i s h which a l s o c o n t a i n e d s u b - a d u l t worms i n t h e body c a v i t y .  Sub-adult worms were found i n t h e  body c a v i t y from the 118th day onward. female Philonema  Mature male and  agubernaculum were found i n tie coelom  from  the 151st day b u t no g r a v i d worms were recovered from t h e f i s h which s u r v i v e d up t o 182 days (Table I I I ) . Loss o f f i s h due t o a contaminated water supply, terminated the experiment b e f o r e any worm became g r a v i d . the t r o u t  strain  A l l stages o f  17  Fig.  I  Development o f Philonema o n c o r h y n c h i i n sockeye o f v a r i o u s ages f r o m C u l t u s L a k e .  salmon  i  LOG 1 — i — r  O I II I  LENGTH O  IN M M .  oo  O)  o  TT  o o  JO  m E TO  > m  o po CO  2  O  z —i i  to  IV) 01  IV) CD  CO  o IV)  at  .ADDER  J COELO  _  LAPVAE IN SWIh  o  a-  C-  cr  > Pi  oo  01  18 developing of  i n trout  sockeye s t r a i n  were m o r p h o l o g i c a l l y  developing  identical  i n sockeye.  The  two  t o those forms  differ,  however i n t h e d e v e l o p m e n t a l r a t e . No mentally  visceral  infected  f o u n d dead  One  adult  growth  t h e 182  15.4  mm.  f e m a l e worm  of a female f i s h  of Philonema agubernaculum  b o t h male and f e m a l e f i s h .  over  non-gravid  experiwas  on t h e  infection.  The  to  fish.  i n the abdominal c a v i t y  1 6 0 t h day o f  in  a d h e s i o n s were o b s e r v e d i n any  day d u r a t i o n f o r males  The  size  t h e same  a c h i e v e d by t h e worm  o f t h e e x p e r i m e n t was  and f r o m 26.mm. t o 70.9  Philonema agubernaculum  was  f r o m 11.8  mm.  mm.  f o r female  (Figure I I ) .  DISCUSSION Infection  of j u v e n i l e Platzer  the  life  cycle  bicuspidatus initial  Beach  of Philonema  oncorhynchi using  as t h e i n t e r m e d i a t e  host  i n the connective  t o s i x metres  November t o mid December  (Foerster,  tissue  1925).  of sockeye  lake  (Foerster,  They  spawn a t L i n d e l  remain i n the  eleven  and o n e - h a l f months and o c c a s i o n a l l y  The  latter  constitutes  (Foerster  1929).  The  early  swimming i n t h e  for  o n l y 26.7%  salmon.  The e g g s h a t c h e d  and t h e f r y a r e f o u n d f r e e 1938).  Cyclops  i n depth from  d u r i n g the w i n t e r May  completed  and f o l l o w i n g i t s  C u l t u s L a k e , the sockeye salmon  i n w a t e r o f one  in early  conditions  and Adams ( i n p r e s s ) s u c c e s s f u l l y  development In  sockeye salmon under n a t u r a l  two  lake  years.  of the t o t a l m i g r a n t s  seaward m i g r a t i o n  commences on A p r i l  12  19  Fig.  II  Development o f Philonema infected trout.  agubernaculum  in experimentally  20 and  continues throughout  the f i r s t in  and May r e a c h i n g a peak d u r i n g  (Foerster,  1925).,  t h e s e a f o r two y e a r s r e t u r n i n g  the f a l l time, to  week o f May  April  of t h e i r  Philonema  the s e a .  rapidly life  during the l a s t  oncorhynchi  t o t h e l a k e t o spawn i n  (Foerster, i s carried  1938).  A t t h e same  by downstream m i g r a n t s  T h e y grow v e r y s l o w l y i n t h e h o s t and mature  physiological  t o that  The December  f o u r t o s i x months o f t h e h o s t ' s  i n f r e s h w a t e r when t h e h o s t s  i s a complete  7.7°C.  year  oncorhynchi  and r e p r o d u c e  There  and  third  They f e e d and grow  spawn.  a d a p t a t i o n of Philonema  o f i t s anadromous h o s t s  mean t e m p e r a t u r e (average  o f C u l t u s L a k e d u r i n g November  of a l l depths  ( R i c k e r , 1937a).  Philonema  from  0 t o 40 m e t r e s ) i s  oncorhynchi experimentally o  reared third the  i n C y c l o p s b i c u s p i d a t u s a t 10 C r e a c h e d stage  culture  tion  by 37 d a y s and s u r v i v e d f o r 85 d a y s , a f t e r w h i c h o f c o p e p o d s was u s e d  of f i s h .  Ko  eight  that  live  infected  up f o r e x p e r i m e n t a l  c o p e p o d s may  8 months and d u r i n g t h i s of Philonema  time  months a t 4 ° C .  fry  This  infected suggests  i n c o l d e r l a k e s f o r p e r i o d s of  are capable  of t r a n s f e r r i n g  t o the f i s h .  S o c k e y e s a l m o n spawn i n t h e o u t l e t River  infec-  ( p e r s o n a l communication) maintained  copepods f o r a p p r o x i m a t e l y  infection  the i n f e c t i v e  d u r i n g August t o October  each  year  of the Babine  and e m e r g i n g  swimming  e n t e r t h e l a k e i n t h e f o l l o w i n g May and J u n e , where  t a k e up a p e l a g i c , Groot,  1963).  zooplankton-eating habit  Sockeye f r y o f Babine  (Johnson  Lake were  found  they  and infected  21 with  third  stage  larvae  when c a p t u r e d  in July.  Infected  copepods  and c l a d o c e r a n s were f o u n d i n t h e s t o m a c h  of  fry.  these  infected ward  T h i s means t h a t  copepods  fingerlings  Survival  f r y i n B a b i n e Lake survive  t h e n e x t age  free-swimming larger  of e x p e r i m e n t a l l y  their  sized  native  in July  sea-  that  of j u v e n i l e  i n May.  T h i s would  have  of I n f e c t i o n The  abundant occur  third  o f t h e two been  tissue  which  gases.  The  that  stage  accountkon the  i s probably sub-mucosa  infected some  size  classes  occasional  classes  and B a b i n e L a k e .  An  o f down alter-  of Philonema i s  by b o t h P^ o n c o r h y c h i  and  rates.  larvae layer  o f P^ o n c o r h y n c h i  a r e most  of swimbladder but  p e r i t o n e u m and t u n i c a  tunica  infect-  Swimbladder  i n the submucosal  (1966) s t a t e s  of  to infect  and two y e a r  infected  of the  i n the p a r i e t a l  copepods  s o c k e y e s a l m o n w h i c h become  a g u b e r n a c u l u m w h i c h grow a t d i f f e r e n t  Route  a few  from January t o August  worms f o u n d i n one  the f i s h  of sockeye  infected  suggest that  class  explanation  infection  months, and t h e o c c u r r e n c e  stream m i g r a n t s i n C u l t u s Lake  P.  before  d u r i n g t h e month o f J a n u a r y i n  approximately eight  c o p e p o d s may of  the winter  (1964) s u g g e s t e d t h a t  takes place  C u l t u s Lake.  ed  again during  w o u l d be e x p o s e d t o  migration. Platzer  for  the f i s h  contents  externa a barrier  consists against  externa. o f dense diffusion  i s a loose j e l l y - l i k e  w h i c h n e r v e s and v e s s e l s p a s s t o t h e mucosa.  tissue,  also Fange connective of through  T h i s might  be  22 the reason for abundance of l a r v a e i n sub-mucosal l a y e r . T h i r d stage l a r v a e migrate t o the sub-mucosal l a y e r of the swimbladder from the stomach w i t h i n seventeen hours of being eaten by the f i s h i n the i n f e c t e d copepods or i n t a c t i n f e c t e d f i s h .  Ricker  (1937b) s t a t e d t h a t f i n g e r -  l i n g sockeye salmon take f o u r t o eight- hours t o d i g e s t a small p l a n k t o n i c meal.  T h i s shows t h a t l a r v a e may not r e q u i r e t o o  l o n g a time t o move t o the swimbladder. avoid complicated  routes of m i g r a t i o n  shown e x p e r i m e n t a l l y .  The l a r v a e  probably  but i t has not been  Kuitunen-Ekbaum (1937) s t a t e d t h a t  l a r v a e made t h e i r way through the i n t e s t i n a l w a l l i n t o the abdominal c a v i t y of the f i s h .  P l a t z e r (1964) suggested four  routes of m i g r a t i o n : 1)  The l a r v a e may enter the deep v i s c e r a l  lymph-  a t i c s and proceed t o the p o s t e r i o r c a r d i n a l v e i n s v i a the coeliaco-mesenteric 2)  and s u b v e r t e b r a l  lymph  trunks.  The l a r v a e may enter the h e p a t i c p o r t a l system  andb»transported t o the swimbladder by the c o e l i a c o - m e s e n t e r i c a r t e r y where they c o u l d penetrate  the v e s s e l w a l l s and enter  the t u n i c a a d v e n t i t i a . 3)  Active migration  through the i n t e s t i n a l w a l l .  4)  A p e n e t r a t i o n through the w a l l of the i n t e s t i n e  i n t o the coelom and then p e n e t r a t i o n  i n t o the p a r i e t a l  peritoneum or t u n i c a a d v e n t i t i a . A f u r t h e r p o s s i b i l i t y i s t h a t the l a r v a e from the stomach migrate i n t o the swimbladder through the pneumatic duct  of the f i s h .  23 Larvae mucosal canal  are found f r e e  and s u b - m u c o s a l  i n t o the swimbladder  17 and 23 h o u r s a f t e r found  contents.  around the o u t l e t  of the  of the fish,, that  layer  live  or v i s c e r a  Udvardy the Salmonidae  No l a r v a e a r e time.  Dead  that  in fecal  materials  they could  pene-  wall.  (unpublished) stated  that  t h e stomach o f  i s very strong walled. P e n e t r a t i o n through  strong walled  migration.  between  ones were f o u n d i n t h e i n t e s t i n a l  and i t i s u n l i k e l y  the i n t e s t i n a l  died  at that  They were a l w a y s f o u n d t r a p p e d  of the i n t e s t i n e trate  canal,  attempted i n f e c t i o n .  i n the p e r i t o n e a l  l a r v a e , but r a r e l y  this  tissue  i n the pneumatic  stomach  would  n o t be an e a s y r o u t e f o r  I n an e x p e r i m e n t , l a r v a e  were f o u n d t o m i g r a t e  i n t o the swimbladder  when t h e p o s t e r i o r e n d o f t h e stomach  was t i e d  the passage o f l a r v a e  to restrict  Physocl- i s t i c when l a r v a e copepods logical duct.  fish  were f e d t o t h e f i s h . incompatability  The open  route  o r when  infected  orfbthe c l o s e d  pneumatic  o f s a l m o n i d s a p p e a r s t o be  of m i g r a t i o n . of Philonema  In t h e p r e s e n t  study, larvae  i n the swimbladder  began  were 15 months o l d and s u b - a d u l t in  infected  T h i s may be due t o t h e p h y s i o -  of the host  pneumatic duct  and D e v e l o p m e n t  oncorhynchi  c o u l d n o t be  were s q u i r t e d i n t o t h e stomach  t h e most l i k e l y Growth  gold  i n t o the i n t e s t i n e .  of Philonema  t o grow when t h e f i s h  worms a p p e a r e d i n abundance  t h e a b d o m i n a l c a v i t y by t h e t i m e f i s h  were 20 months o l d .  24 Adult  male P h i l o n e m a  when f i s h  oncorhynchi  appeared  i n t h e body  were 25 months o l d . A few t w o - y e a r o l d downstream m i g r a n t s  were f o u n d  t o harbour  the swimbladder adult  large  been e x p o s e d  than  t o t h e progeny  spawning salmon. percentage  of infection  i n the sockeye  year  so t h a t  stage  s t a g e and s u b -  (1937) o b s e r v e d  that  two i n f e c t e d  fish  of e i g h t  larvae  one-year o l d  o u t o f 100 o n e two-year o l d s  i n t h e w a l l s of the swimbladder t o the macroscopic  i n keeping with the f i n d i n g s stage  a greater  water as i n the a d u l t  s t a g e o r s u b - a d u l t worms i n t h e body c a v i t y .  No t h i r d  from  K u i t u n e n - E k b a u m was unaware o f t h e  a l l of her records r e f e r  therefore  They have  p a r a s i t e s were n o t a s numerous  occuring i n fresh  i n C u l t u s Lake. third  larvae i n  o f two g e n e r a t i o n s o f P h i l o n e m a  o l d s m o l t s e x a m i n e d , and s e v e n  microscopic  stage  i n two-year o l d smolts than  I n 1937 she f o u n d  examined  i n C u l t u s Lake  one y e a r o l d m i g r a n t s .  Kuitunen-Ekbaum  s m o l t s and she a l s o s t a t e d  sockeye.  numbers o f t h i r d  and g r e a t e r numbers o f f o u r t h  worms i n t h e c o e l o m  per host  cavity  larvae  of t h i s  are found  fourth  Her r e c o r d s are study.  i n swimbladder o f  p r e c o c i o u s m a l e s ( 2 9 - 3 0 months) and immature t h r e e - y e a r o l d sockeye  (35 months) f r o m C u l t u s Lake, b u t by t h i s  grown t o t h e f o u r t h appeared are  found  stage.  i n t h e body c a v i t y i n t h e coelom  immature f i s h  Fourth stage of both  of f i s h  i s that  t h e y have  and s u b - a d u l t worms 1  Adult gravid  o f p r e c o c i o u s male s o c k e y e  o f t h e same a g e .  t h e s e two g r o u p s  fish.  time  The o n l y d i f f e r e n c e  females  but not i n between  p r e c o c i o u s males a r e young  25 by  age  of  C u l t u s L a k e w h i c h a r e 40-43 months o l d , g e n e r a l l y h a r b o u r  adult  but s e x u a l l y matured.  the  the  3-5  year  life  cycle  30  days of i n f e c t i o n .  swimbladder  of i n f e c t i o n  as t h i r d  i s not  due  oncorhynchi. 182  shown by  P.  and  the presence  System.  T h e r e a f t e r , from  The  dracunculoids  life  t h e worms a r e g e n e r a l l y p r e s e n t  as  oncorhynchi  longer  developmental  occur  This differential  to difference  i n sex  infected  become g r a v i d  of  adult  than  differs  from  period  and  of l e s s the r e s t  than  In t h e  of and  Similar  host of  P.  between  a year  as  infected  R i v e r of the  one-  Kootenay that  of other  1 year f o r completion,  i n the requirement  in i t s physiological  of a  adaptation  I  Male worms o f most d r a c u n c u l o i d s a r e r a r e are r i p e .  rate  stage  agubernaculum resembles  the h o s t ' s m a t u r i t y .  females  i n the  of the host  adult  in less  from the Lardeau  i n i t s requirement  b u t P^  synchron-  to  occasionally  the  o f g r a v i d worms i n n a t u r a l l y  cycle  On  58th  the n a t u r a l l y  may  o l d immature t r o u t  host.  (Figure I I I ) .  a g u b e r n a c u l u m grows t o t h e  days,  i s adapted  t o some o t h e r unknown f a c t o r ( s ) .  o b s e r v a t i o n s a r e made on  for  only  o f P^ a g u b e r n a c u l u m o c c u r s d u r i n g  stage l a r v a e .  therefore attributed  and  host  However, t h e worms may  growth of l a r v a e  year  sockeye  o f P^_ a g u b e r n a c u l u m i s n o t  growth o f l a r v a e  sub-adults.  oncorhynchi  o f i t s anadromous s o c k e y e  No  t h e 182nd day  151  Philonema  w i t h i t s non-anadromous t r o u t  the f i r s t  is  that  cycle  c o n t r a r y , the l i f e  ized  female  worms. Thus i t i s s e e n  to  Spawning male and  case  o f P.  when t h e  oncorhynchi males are  found  26  Fig.  I l l D i a g r a m m a t i c c o m p a r i s o n o f t h e s p a w n i n g a r e a s and t h e m i g r a t i o n s o f t r o u t and s a l m o n w i t h t h o s e - o f P h i l o n e m a a g u b e r n a c u l u m and P h i l o n e m a o n c o r h y n c h i i n t h e i r t r o u t and s a l m o n h o s t s r e s p e c t i v e l y . £  spawning  area  juvenile  migration i n freshwater  \  _ —  D  adult  ^  free  L""  _ —  _ —  spawning swimming  t o sea migration larvae  infected  copepods  infected  juvenile  transfer  of i n f e c t i o n  fish  i n lake  SALMO G AIRDNERI -SPAWN MORE.THAN ONCE LIFE  LIFE  HISTORY OF TROUT  TRIBUTARY  A  LAKE  • •  HISTORY  ONCORHYNC HUS NERKA - SPAWN ONCELIFE  OF P.AGUBERNACULUM  A  m  f  <  \  < <  .  —  —  LIFE HISTORY OF P.ONCORHYNCHI  r  •  ffl  t  m •  HISTORY OF SOCKEYE  ®  ^  ffl ' J RIVER  ESTUARY  OCEAN  •  \) m  j  27 commonly w i t h r i p e  females  i n t h e body c a v i t y  of adult  t h e y a r e n o t a s numerous a s i n immature f i s h . a b l y mature much more e a r l i e r Wierzbicki  which  and  a r e r e - a b s o r b e d by t h e o r g a n  pretation  and t h e n d i e o f f .  the males of Philonema  sanguinea later  die after  Male worms p r o b -  t h a n the, f e m a l e  (1960) h a s shown t h a t  f i s h but  the f e r t i l i z a t i o n ,  become e n c y s t e d  of the host.  c o u l d p r o b a b l y be a l s o a p p l i e d  This  inter-  t o male worms o f P.  oncorhynchi. Escapement  o f Worm f r o m t h e H o s t Few  escapement  s u g g e s t i o n s have been made s o f a r r e g a r d i n g t h e  of gravid dracunculoids. Nybelin  the  abdominal  ation  c a v i t y never  way  within  out.  that  female  (1871) s u g g e s t s t h a t the f i s h  Linstaw  the dracunculoids i n h a b i t i n g  leave the l a t t e r ,  i s g i v e n how t h e r i p e  Willemoes-Sum burst  (1931) s t a t e s  the r i p e  the i n t e s t i n a l  reaching  with the faeces.  upon t h e f a c t had,  that  with i t s e x t e r i o r  fish.  The p a r a s i t e  suggested  that  liberate  larvae  female  the r i p e  wall  female  of the f i s h ,  He b a s e d  was f o u n d dead Philonema  when o b s e r v e d . might  pass  that  i n t h e body c a v i t y w h i c h  parasite;; thus  which  wall  of the  Meyer  (1960)  out with the roe  the gravid  escapes  a s he h a s f o u n d l a r v a e  their  h i s statement  the i n t e s t i n a l  V i k (1964) s u g g e s t e d  r o e s o f salmon,  parasite  d r a c u n c u , l o i d was f o u n d  end, p i e r c e d  the gravid  d u r i n g spawning.  and  a ripe  female  young a c t i v e l y make  that  makes i t s way t h r o u g h the e x t e r i o r  no e x p l a n -  p a r a s i t e s reach the e x t e r i o r .  and t h e l i b e r a t e d  (1874) s u g g e s t e d  though  worms  through  i n the m i l t  the m i l t  and r o e s  28 of  stripped  fish.  of  maturation  oncorhynchi the  Kuitunen-Ekbaum  of the host  dorsal of  fish  (1921) o b s e r v e d  a r e suspended  peritoneum.  an immature  from  that  ovary.  in size.  dorsally. trout  anical  Philonema  of r i p e  i n ripe  female  o v a r i e s and a l s o  sockeye  salmon.  the author  t o pass  trout  and s o c k e y e .  structure  roe duringfcspawning a c t .  the g r a v i d  Gravid  i n t h e mesovarium of are very  o r any o t h e r mech-  o r body w a l l and  s u p p o r t s the assumption  (1960) t h a t  out with roe  G r a v i d Philonema  means t o p e n e t r a t e t h e i n t e s t i n a l  (1937) and Meyer  i n salmonids  o b s e r v a t i o n s i n matured  are observed  and p o s s e s s no d e n t i c u l a r  therefore  as the o v a r y  salmon.  o v a r i e s of r i p e  fragile  thus  o v a r i e s a r e c n o t c o v e r e d m e s i o - v e n t r a l l y and  during the s t r i p p i n g  the  t h e mem-  and f u l l y m a t u r e d o v a r i e s a r e n o t c o v e r e d  and s o c k e y e  worms a r e f o u n d  forward  (unpublished) observed  The a u t h o r h a s made s i m i l a r  Gravid  surface  however, t h e  The s u r f a c e o f t h e o v a r y ,  increases  Udvardy  i n the  a narrow area of the ovary  g r a d u a l l y w i d e n s and e x t e n d s  dorso-laterally  the e n t i r e  of the o u t e r s u r f a c e p a r t s from  membranous c o v e r i n g .  the maturing  the o v a r i e s of the  As d e v e l o p m e n t s p r o c e e d ,  uncovered  that  during  a membrane o r i g i n a t i n g  of the inner surface l e a v i n g  without  time  of the Philonema  reach the e x t e r i o r  T h i s membrance e n v e l o p e s  edge o f t h e membrane brane  might  that  activities.  Kendall salmonid  coincides with that  and t h e p a r a s i t e  spawning  (1937) s u g g e s t e d  of Kuitunen-Ekbaum  female  pass  out with  30 On The  ripe  t h e o t h e r hand t h e r e r e m a i n s  individuals  abdominal  cavity  of the f i s h ,  g r a d u a l b r e a k up free  of Philonema  Ekbaum  host*.  (1937).  explain  The  trout  or  time  dead  above a s s u m p t i o n  the is  stated  that  not s u c c e s s f u l i s a "dead  Visceral  to  the l i f e  i n t h e body  i n immature  cycle  end"  A t t h e same cavity.  sockeye  salmon  I t i s assumed  of Philonema  from  agubernaculum  t h e r b y the r e p r o d u c t i v e  i n immature hosts..  adhesions  maturing  the  found  i n immature h o s t s and  Visceral  adhesions are observed  anadromous s o c k e y e  C u l t u s Lake.  ponding  dead  Kuitunen-  n o t be u s e d  i n s i x months.  worms a r e a l s o o b s e r v e d  facts  of  worms i n s e x u a l l y  Shuswap L a k e maybe P^ a g u b e r n a c u l u m .  cycle  in  could  become  in experimentally infected  t o mature  Dead and e n c y s t e d P h i l o n e m a from  b u r s t s i n coelom  of r i p e  agubernaculum  are found  adult  the l a r v a e  and  salmon.  Philonema immature t r o u t  and  a l s o been s u g g e s t e d by  the means o f l i b e r a t i o n  immature  i n the  await the decomposition  when t h e f e m a l e T h i s has  possibility.  oncorhynchi l y i n g  o f t h e body o f t h e f i s h  swimming i n l a k e  disintegrated  may  another  These  salmon  from  and  oncorhynchi corres-  the swimbladder  into  cavity. Severe  in January i n August  i n f e c t e d w i t h P^  o c c u r i n 23 month o l d f i s h ,  t o the m i g r a t i o n of l a r v a e  body  i n immature  visceral  and A p r i l , from C o a s t a l  1965  adhesions are observed from  fishing  in fish  captured  t h e G u l f o f A l a s k a b u t much and  none i n f i s h  caught  from  less the  31 spawning ground. male s o c k e y e  Occasionally,  are found  t o posses  above o b s e r v a t i o n s u g g e s t adhesions ing  are t r a n s i t o r y .  t o the  gonadal  i n matured sockeye  fish  are  and  male and  very l i g h t  though  North  precocious  adhesions.  They b e g i n t o d i s a p p e a r  noted  adhesions and  Pacific  The  t h e wide s p r e a d cf. v i s c e r a l  (1965) , who  immature Oncorhyrius n e r k a Sea  that  adult  development of the h o s t s .  agreement w i t h F r e n c h  Bering  the  correspond-  This i s in  the. a b s e n c e  of  full  adhesions  a r e more p r e v e l a n t i n  Oncorhyrius k e t a c a u g h t Ocean and  i n the  the m a j o r i t y of the  adhered  males.  Summary Philonema  oncorhynchi  requires  32 t o 35 months t o  mature i n i t s anadromous h o s t , whereas P h i l o n e m a  agubernaculum  r e q u i r e s 6 t o 12 months i n i t s non-anadromous t r o u t developmental The  form  salmon  in trout  i s matured The  result in  stages through  from  different  distinct  c a n mature i n maturing  variation  which they pass i n immature  cycles  physiological^different  species.  fish,  The  identical.  but  the form  in  fish.  in life  h o s t s or from  are  host.  of the  responses  two  forms  of a s i n g l e  t h e e x i s t e n c e o f two  may species  biologically  32  SECTION I I Experimental  cross  infection  between t r o u t  and salmon  strains.  INTRODUCTION The oncorhynchi inherent  longer developmental  pattern o f P h i l o n e m a  i n i t s anadromous h o s t may be e i t h e r  because o f  c h a r a c t e r i s t i c s o r due t o a p h y s i o l o g i c a l  Hot' t h e l o n g l i f e  history  This series Philonema  from t r o u t  information Philonema  of experiments and salmon  regarding  from  and anadromous h a b i t s of cross  were c a r r i e d  the d i f f e r e n c e s  adjustment  of salmon. infection  between  out t o obtain  i n the l i f e  c y c l e s of  anadromous and non-anadromous hosts.  MATERIALS AND METHODS Source  and H u s b a n d r y o f F i s h The  a)  following  T r o u t , Salmo i)  different  species  of f i s h  Columbia,  Hatchery reared  rainbow  Trout Hatchery of the F i s h Dept. ii)  obtained b)  trout  f r y which  i) collected  were 4  o b t a i n e d from  and Game B r a n c h  of B r i t i s h  o f R e c r e a t i o n and C o n s e r v a t i o n . Year  o l d slow maturing hatchery reared  f r o m t h e Summerland H a t c h e r y o f t h e above  Salmon,  used.  gairdneri  months o l d a t t h e b e g i n n i n g o f t h e e x p e r i m e n t , Abbotsford  were  trout,  organization.  Oncorhynchus Sockeye  (Oncorhynchus  nerka)  from the w i n t e r r u n sockeye  reared  from  eggs  f r o m C u l t u s Lake and  33 hatched fish  i n the U n i v e r s i t y  of B r i t i s h  December  Spring  (Oncorhynchus  iii) Department  Pink  State,  of Zoology, U n i v e r s i t y  Campbell  Chum  v)  Kokanee  from Oregon F i s h British  c)  Gold F i s h ,  Columbia  Gold Infection  hatched Columbia thirty  i n the on J a n u a r y  miles north  Research C o u n c i l .  The  k i s u t c h ) s i x months o f age when  Squamish, B r i t i s h  Columbia i n  i n J a n u a r y , 1965.  (Oncorhynchus  nerka k e n n e r l y i ) ,  a year  and Game H a t c h e r y , were o b t a i n e d t h r o u g h Research  Council.  Carassius carassius f i s h were o b t a i n e d l o c a l l y  and M a i n t e n a n c e Fish  Hatchery,  k e t a ) h a t c h e d i n O c t o b e r , 1964  Columbia  (Oncorhynchus  f r o m Cheakamus R i v e r ,  the  of B r i t i s h  (Oncorhynchus  September, 1964 and u t i l i z e d  old,  State  on J a n u a r y 17, 1965. Coho  vi)  h a t c h e d on  I s l a n d , B.C.  were o b t a i n e d f r o m t h e B r i t i s h were u s e d  gorbuscha)  from Baer R i v e r ,  R i v e r , Vancouver  iv)  caught  tshawytscha)  U.S.A.  (Oncorhynchus  1, 1965 f r o m e g g s c o l l e c t e d  The  infected.  18, 1964 were o b t a i n e d f r o m t h e Samish  B u r l i n g t o n , Washington  fish  Hatchery.  were s i x and o n e - h a l f months o f age when ii)  of  Columbia  of  were i n f e c t e d  from a p e t shop.  Experimental Fish iy, t h e f o l l o w i n g  different  methods: a)  F r y and f i n g e r l i n g  transferring  f i s h were s t a r v e d  f o r two d a y s b e f o r e  them i n t o 8" g l a s s s t o c k i n g d i s h  i n three l i t r e s  34 of  dechlorinated'  were p o u r e d The  water  ferring  into  was  the  fish  clipping.  b)  Fingerling of  the  fish  stomach  anaethetized operation.  tank.  c)  trout  in  Section  infected  allowing  recover  fed  thirty  222  (1  were  were  to  tagged  infected  fifty  with  by  on  copepods.  after  trans-  were marked  help  The  of  fish  by  on  a  the  poly-  were  carrying  transferred  gelatin  fish  (depending  the  before  and  feed  fish  water.  g./litre)  to  copepods  The  fish  i n a minimal  copepods per  fish  unfed  tanks.  copepods per  fish  recirculating Yearling  were  tube  i n MS The  to  for  into holding  infection)  ethylene  Thirty  dish  filtered  fin  rate  water.  out  into  capsule  this  a  as  holding  described  I.  RESULTS Infection from  of  different  a)  served  as  Infection  the of  Fish per  and  Gold Fish  with  Philonema  sockeye Development  Lake  Salmonid  fish  in  the  were f o u n d  larvae  were  i n the  IV)  which  copepods.  no  Salmo  infected  swim b l a d d e r .  which  infected  salmon from  Cultus  control.  Rainbow T r o u t ,  days d u r i n g  (Figure  in naturally  The  development  third WCLCU  stage  with  an  fish  survived  of  with  comparable  gairdneri  to  larvae  average  for  9.1  stage  of  larvae  forty-six  wase f o u n d .  a mean l e n g t h third  of  A l l  0.98  larvae  mm. in  35  Fig.  IV  Development of Philonema o n c o r h y n c h i i n f e c t e d t r o u t o f two y e a r s o l d .  in experimentally  o  LENGTH  IN MM  CP  T  o  en  oo ao 0|  o  o3»<eoa ooo oo #o o  oh  5a OIL  o  36 b)  Infection  o f Coho, Oncornyncfaus  Sixteen exposed of  for infection.  infection  average larvae c)  o f 3.6  larvae  per f i s h  were i n t h e t h i r d  Infection  the f i s h  o f Chum fish  (Oncorhynchus k i s u t c h ) w e r e  A l l the f i s h  and 8 1 % o f t h e f i s h  The of  f i n g e r l i n g Coho  kisutch  died  after  fifty-six  were f o u n d i n f e c t e d  i n the swimbladder.  days  w i t h an A l l the  stage.  (Oncorhynchus  keta)  died eighteen hours a f t e r  were i n f e c t e d  infection  w i t h an a v e r a g e o f 3.1  and 7 5 %  larvae  i n the  swimbladder. d)  Infection  of t r o u t f r y  Eleven larvae  per  o f t w e l v e f r y were f o u n d i n f e c t e d  and 84 h o u r s a f t e r copepods.  recovered found  fish  placing Two,  the f i s h  from the swimbladder.  i n 149 d a y s .  larva  Infection Trout  was  a)  B o t h dead  time. fish  fish.  with  respectively  and l i v e  were  larvae  were  Heavier i n f e c t i o n  thus  No growth  survived  of water  of l a r v a e  was  346 d a y s and one  from i t s swimbladder  fourth  (Figure V ) .  o f S o c k e y e , P i n k , S p r i n g , Kokanee and G o l d F i s h  with  Strain Trout  I  One  recovered  i n the d i s h  2, 6, and 11 l a r v a e  occurs with longer exposure  stage  were a u t o p s i e d - 17, 32, 62,  i n t h e stomach o f t h e f i r s t  observed  2.8  fish.  A group of f o u r  infected  with  (page  infected  16) s e r v e d  Infection  with Philonema  as c o n t r o l  of Sockeye  in this  (Oncorhynchus  agubernaculum  experiment, nerka) f r y  i n Section  37  Fig.  V  Development of Philonema o n c o r h y n c h i from sockeye in experimentally infected trout f r y .  salmon  o  "WW Nl H19N31  38 Five (13.5%) up  of t h i r t y - s e v e n  with Philonema  t o the 157th  agubernaculum.  f o u n d i n t h e body c a v i t y  VI)  after  b)  Infection  days  of  larvae  of a sockeye  (Oncorhynchus  was  observed  mm.  f r y o f 8.5  long cm.  gorbuscha), Spring  (0. nerka k e n n e r l y i )  of three  p i n k salmon  recovered after  ii) larvae  growth  (Figure  and G o l d  (0.  Fish  carassius) i ) Two  stage  No  infected  infection.  of P i n k  t s h a w y t s c h a ) Kokanee (Carassius  f r y were  day b u t a s u b - a d u l t f e m a l e o f 32.1  was  161  sockeye  One  of three  twelve  and  third  days.  s p r i n g s were i n f e c t e d  were f o u n d i n t h e s w i m b l a d d e r iii)  were i n f e c t e d  on t h e 5 5 t h  T h r e e kokanee and g o l d  fish  and  third  stage  day.  were f o u n d t o be  uninfected. DISCUSSION Though a c o m p l e t e made due is  to unavailability  sufficient  and s a l m o n result  s e t of r e c i p r o c a l  of m a t e r i a l s at s p e c i f i c  d a t a t o show t h a t  ^strains"  the d i f f e r e n c e  are inherent  of v a r i a t i o n s  due  i s specific  and n o t  to i t s typical  host.  o r P^ The  salmonids.  Neither  strain  was  able  i n the experiment.  agubernaculum  the  from t r o u t  to infect  agubernaculum  experiments  t h a t each form i s capable of i n f e c t i n g  t h a t P.  trout  to host.  demonstrate  salmonid used  not  times, t h e r e  between t h e  characteristics  N e i t h e r P^ o n c o r h y n c h i f r o m salmon from t r o u t  i n f e c t i o n s was  several  the s i n g l e  non-  However, t h e d a t a s u g g e s t s does not i n f e c t  sockeye  salmon  39  Fig.  VI  Development o f Philonema agubernaculum from t r o u t e x p e r i m e n t a l l y i n f e c t e d f r y of sockeye salmon.  in  LENGTH cn  > -<  CO  81  T  CJl  IN MM. 6 T"  OJ  o  <J1  40 as r e a d i l y infected that  as o t h e r h o s t s .  as compared  died within  figure  O n l y 13.5%  t o 65.6%  page 1 6 ) . fry.  The  required larvae The is  and  360  contrasted fourth  t o 450  o f t h e two  oncorhynchi  species  i n Figure VII.  The  I  days i n sockeye stage  or sockeye  infections  during  oncorhynchi  in trout  and s t a g e u n t i l  times required  grew much f a s t e r  mm.  t o molt  parasitized.  P_._ a g u b e r n a c u l u m  P^ a g u b e r n a c u l u m  t o 15.4  mm.  in trout  (4.1  P. mm.)  produced  f o r m a l e s and 26.8  f o r f e m a l e s i n 182 d a y s ; i n s a l m o n  to  distinct  ( f o u r t h s t a g e ) by 58 d a y s .  the 346th day.  salmon  t h a n P^ o n c o r h y n c h i  d i d n o t r e a c h t h e same s i z e  s u b - a d u l t s m e a s u r i n g 11.8  and  stage are  and a r e i n d e p e n d a n t o f t h e h o s t  t o 4.1  (Section I ) .  o f worms i n t r o u t  t o the sub-adult  and s o c k e y e s a l m o n .  grown t o 3.3  mm.  trout  days t o produce s u b - a d u l t s  agubernaculum  both t r o u t  70.9  (Section  observed i n the t h i r d  In n a t u r a l  s t a g e and a g a i n  P.  had  r e a c h e d i n 161  sub-adult  a p p r o x i m a t e l y 240-360 d a y s t o p r o d u c e f o u r t h s t a g e  f o r each s p e c i e s  in  s t a g e was  of time.  development  the  s t a g e by 58 d a y s and  of Philonema oncorhynchi i n e i t h e r  t h e same p e r i o d  coho  Philonema  and 182 d a y s i n immature t r o u t  C o n v e r s e l y , no growth was  larvae  produced the  chum s a l m o n 75%,  shown e x p e r i m e n t a l l y t h a t  sub-adult  latter  r a i n b o w f r y 88%.  r e a c h e s the f o u r t h  s t a g e s 118  fish  are e x c l u d e d the  i n a l l hosts,  81%, r a i n b o w y e a r l i n g s 82%, I t h a s been  ( I f three  o n c o r h y n c h i from salmon  same p e r c e n t a g e i n f e c t i o n  and a d u l t  o f 32 t r o u t  24 h o u r s o f i n f e c t i o n  becomes 7 4 . 8 % ) .  agubernaculum  o f 37 s o c k e y e were  to  (O. n e r k a ) a s u b  41  Fig.  VII  H y p o t h e t i c a l r e p r o d u c t i v e i s o l a t i o n of Philonema o n c o r h y n c h i and P h i l o n e m a a g u b e r n a c u l u m i n s a l m o n i d ( b a s e d on f o u r y e a r r e p r o d u c t i v e c y c l e o f s o c k e y e salmon).  Trout  Sockeye Philonema oncorhynchi  time i n days  Philonema agubernaculum  Philonema oncorhynchi  Philonema agubernaculum 3rd  stage  4th  stage  30 50  100  3rd  stage  sub  150  adult adult  200 250  3rd  stage 300  10% 4 t h stage  gravid  ?  4th  stage gravid (at l a t e s t )  350 400 450  10% sub adult  500 550 600 dead and encysted  gravid  650  1100  42 adult  female  measuring  The ing f i e l d  observations.  which  landlocked  do n o t have any l i n k  Only  with sea.  explained  i f a l l Philonema  are a l s o  Lake  b u t no  was f o u n d  Lake.  i n Kootenay  British  There  i n Kootenay  i n Kootenay  puzzl-  found i n  Lakes,  one a d u l t P h i l o n e m a  kokanee e x a m i n e d  several  are g e n e r a l l y  sockeye, 0^ nerka k e n n e r l y i  spawning  easily  Philonema  f r o m T r o u t and K o o t e n a y  anadromous s o c k e y e . 200  was p r e s e n t i n 161 d a y s .  above e x p e r i m e n t a l d a t a e x p l a i n ;  non-andromous t r o u t Columbia  32.1 mm.  These  Lake  i n one o f f a c t s are  a r e P.  agubernaculum. C u l t u s Lake Philonema Platzer  may be c o n s i d e r e d a t y p e l o c a l i t y f o r  oncorhynchi,.  Kuitunen-Ekbaum  and Adams ( i n p r e s s ) and t h i s  P. o n c o r h y n c h i i s a s p e c i e s w e l l Though t r a n s m i t t e d slowly and  return  recorded cavity  t o spawn.  hosts are s e x u a l l y  fish  a p p r o x i m a t e l y one y e a r o l d .  from C u l t u s Lake. in this  study.  Similar  two p e r i o d s o f i n f e c t i o n  May t o J u l y  from  spawning. that  infected  copepods  An a l t e r n a t i v e  C u l t u s Lake  o f 17 month  p r e c o c i o u s Philonema  T h e i r p r e s e n c e may be e x p l a i n e d by  postulating  fact  mature  However, K u i t u n e n - E k b a u m  n o t e d s u b - a d u l t male worms i n t h e c o e l o m  were n o t e d  the  salmon.  t h e p r e s e n c e o f a few m a c r o s c o p i c worms i n t h e body  sockeye  salmon  t o the sockeye  t h e worms grow and mature  o n l y when t h e i r  o f downstream m i g r a n t  Platzer old  t o the lake  (1964) ,  s t u d y have shown t h a t t h e  adapted  i n f r e s h water,  and a r e g r a v i d  (1937) , P l a t z e r  (Section  surviving  explanation  contains small  I ) , the f i r s t i n since  the previous  i s s u g g e s t e d by  p o p u l a t i o n s of rainbow  43  trout, and by  Salmo g a i r d n e r i ; D o l l y V a r d e n ;  white  fish,  Prosopium w i l l i a m s o n i •  of m i g r a n t  Philonema found  sockeye  agubernaculum.  Data  reaching  well  maturity.  oncorhynchi  possible  during  salmon  One o r more o f t h e s e the l i f e  cycle  and e v e n p r o b a b l e  probably  by h o s t  species  salmonids,  could  occurrence  be  of both  of B r i t i s h from  of lake  P.  Columbia anadro-  residence.  b o t h t h e j u v e n i l e t r o u t and s o c k e y e  on p l a n k t o n i c  they  o f t h e worm.  and P. a g u b e r n a c u l u m f r o m t r o u t  foraging  P.  reaction  several  o f P^ oncorhynchi  i n f e c t i o n s o f P. o n c o r h y n c h i their  bladders  p a p e r show t h a t  but are destroyed  of i n t e r b r e e d i n g  i s presumed t h a t  acquire  I I I of t h i s  and P^_ a g u b e r n a c u l u m i n some l a k e s  a question  mous s o c k e y e  i n t h e swim  Shuswap L a k e c o n t a i n s  f o r maintaining  The  as l a r v a e  i n Section  as white f i s h .  responsible  rises  i n sockeye  l e a v i n g Shuswap L a k e a r e v e r y  develop r a p i d l y i n the f i s h before  are p a r a s i t i z e d  species.  The  It  I f these  P^ a g u b e r n a c u l u m , t h e r a p i d l y g r o w i n g worms  may be t h a t  as  c h a r , S a l v e l i n u s malma ;  may  and P ^ a g u b e r n a c u l u m food  i n the lake.  Inter-  b r e e d i n g may o c c u r w h i c h would i n v a l i d a t e t h e r e c o g n i t i o n o f genetically of  distinct  species.  available information  trout  that  I t i s a r g u e d , h e r e , on t h e b a s i s interbreeding  may n e v e r  or s o c k e y e , even though they c a r r y both s p e c i e s  same t i m e .  Figure  VII presents  P. a g u b e r n a c u l u m oncorhynchi  i s very  slow.  culum r e a c h e s a r e p r o d u c i n g  t h e argument  i s fast  at the  graphically.  m a t u r i n g whereas P.  In s o c k e y e stage,  occur i n  salmon when Pj_ a g u b e r n a -  P. o n c o r h y n c h i  continue t o  44 live P.  i n the  swimbladder  agubernaculum w i l l  oncorhynci confirmed Lake  reaches by  the  theoretical mentally  become e n c y s t e d  by  larvae. the  All  time  This i s in  a v a i l a b l e from sockeye  12  f a t e of the  of r e a c h i n g  between P^  reached  of  P. fact  Shuswap  3 year  d i d not  worms i s unknown.  o n l y the  fourth larval  a reproducing  stage  (page  o r no  and  probability  species.  i n t e r b r e e d i n g between t h e  due  If there  two  c y c l e i n the  the  cross possi-  of i n t e r b r e e d i n g  i s any  occurrence  o f an  salmon  isolating  forms i n t r o u t , t h e r e  non-spawning  stage.  30).  to temporal  t o a common gene p o o l , b e c a u s e  May.ir  o r no  P_^ a g u b e r n a c u l u m i n s o c k e y e  isolation  346  P.  o f a d u l t s and  is little  of r e p r o d u c t i v e  reproductive  beyond At  developmental p e r i o d of  a coincidence  on  trout experi-  survive  there  oncorhynchi  contribution  because the  s p e c i e s , but  is little  mechanism o f t h e s e  been p r e s e n t e d  agubernaculum superimposed d u r i n g  might produce two  but  Vj_ o n c o r h y n c h i  had  w i t h P^  There  because  with  ultimate  of the  argument has  (Figure VII)  months o f t h e  breeding bility  trout, a similar  oncorhynchi  oncorhynchi,  ful  or f o u r t h s t a g e  i n matured h o s t s .  information  infected  infection  last  and  maturity  grounds  days the  d a y s P^ An  die  third  (Section I I I ) . For  346  as  is  of no  unsuccess-  host.  (1940) d e f i n e d s p e c i e s as " g r o u p s t f a c t u a l l y  or p o t e n t i a l l y  interbreeding natural populations  reproductively  isolated  (1950) d e f i n e d t h e  from other  same as  "The  which  s u c h g r o u p s . " and  largest  and  most  are  Dobzhansky  inclusive...  45 reproductive  community o f s e x u a l  u a l s which share author confirms Simon and Simon,  and c r o s s  i n a common gene p o o l . " the v a l i d i t y 1936  from Philonema oncorhynchi,  there  characteristics  by w h i c h t h e two  Electrophoretic  a n a l y s i s o f t h e two s p e c i e s  difference  (Figure  VIII).  species  the  agubernaculum,  of developmental  although  individ-  In c o n c l u s i o n ,  of Philonema  on t h e b a s i s  fertilizing  a r e no  c a n be  differences  morphometric separated.  confirms  their  46  Fig.  VIII  A comparison of s t a r c h g e l e l e c t r o p h e r o g r a m s of P h i l o n e m a a g u b e r n a c u l u m f r o m t r o u t and P h i l o n e m a o n c o r h y n c h i from sockeye salmon. T = Philonema  agubernaculum  S = Philonema oncorhynchi  47  SECTION I I I Hormonal  Treatments  INTRODUCTION It ion  was d e m o n s t r a t e d  of Philonema  i n Section  I that  the r e p r o d u c t -  oncorhynchi coincides with that  of i t s  anadromous h o s t . The  n e x t two s e c t i o n s  were u n d e r t a k e n is  dependent  to test  describe experiments  the h y p o t h e s i s t h a t P^ o n c o r h y n c h i  on h o r m o n a l  stimulation  synchronization  of r e p r o d u c t i o n .  salmon  with adult  infected  treated with several eration  MATERIALS AND The salmon a)  c o u l d be  following  were u s e d  strains  b)  fish  sockeye  hormones t o s e e i f any  accel-  produced.  of n a t u r a l l y  year o l d sockeye Columbia  infected  salmon  i n May,  were  sockeye  1964.  collected At the  c a p t u r e t h e y were m i g r a t i n g downstream.  s t o c k s were m a i n t a i n e d i n s e a w a t e r Board  green  as e x p e r i m e n t a l a n i m a l s :  f r o m Shuswap L a k e , B r i t i s h of t h e i r  Pre-spawning  METHODS  A p p r o x i m a t e l y l\  time  from the h o s t f o r  b u t n o n - l a r v i g e r o u s worms were  different  of development  which  a t the F i s h e r i e s  The Research  hatchery.  Pre-spawning  sockeye  C e n t r a l Lake, Vancouver  salmon Island,  were c o l l e c t e d  from Great  and m a i n t a i n e d i n a  large  48 rectangular the  tank i n f r e s h w a t e r at the B i o l o g i c a l  F i s h e r i e s R e s e a r c h B o a r d , Nanaimo.  a saltwater  bath three  S t a t i o n of  The f i s h  t i m e s a week t o p r e v e n t  were  given  fungal  infection. Preparation  o f Salmon P i t u i t a r y  Extracts  E x t r a c t s from the p i t u i t a r y  of  Oncorhynchus  t s h a w y t s c h a were p r e p a r e d by Mr. P. S c h m i d t of  and D r . M.  the F i s h e r i e s R e s e a r c h Board of Canada, Vancouver,  Columbia,  u s i n g p r e v i o u s l y p u b l i s h e d methods  Smith British  (Schmidt, e t a l  1965) . Injection a)  of P i t u i t a r y  Shuswap L a k e  Fish  Twenty f i s h in  Extract  the experiment.  o f a p p r o x i m a t e l y e q u a l l e n g t h were  Thirty untreated  fish  were r e s e r v e d  used as a  control. Each This per  amount fish  period  fish  was  given  i n eight equal i n j e c t i o n s  to injection  o f 0.1  gm.  benzoate  per l i t r e  was made i n t o t h e d o r s a l 0.25 m l . s y r i n g e inserted  the f i s h  o f an aqueous s o l u t i o n  o f m-amino e t h y l  tration  ml. of p i t u i t a r y  extract.  a t 0.05 m l .  day and s p r e a d o v e r a  weeks.  Prior  salt  0.4  a d m i n i s t e r e d on e v e r y t h i r d  of f o u r  2 litres  received  diagonally  o f methane  of seawater.  Each  (Von T h e r i n g ,  gauge n e e d l e .  on e i t h e r  sulphonic  acid  (MS 222, S a n d o z ) a t a c o n c e n -  muscle  and a #30  were a n a e s t h e t i z e d i n  side  injection  1937) w i t h a  The n e e d l e  of the d o r s a l  fin.  was Some  49 t i m e was body. lost  allowed  This  before  precaution  from the f i s h .  of t h e f i s h diffusion  of the f l u i d .  gm.  a total  two e q u a l  doses  i n s e r t e d about  before  bacterial  was  taken  tank along  administered  \ inch  injection  with  inches As  the hypodermic  tags  the c o n t r o l  \\  was made.  each  as i n (a) t o a v o i d spear  on  of i n s e r t i o n of the  infection,  F i s h were t a g g e d w i t h  extract  and a 10 m l . SESI  i n a u r e o m y c i n powder b e f o r e  same p r e c a u t i o n  to a holding  o f t h e hormone.  a n t e r i o r l y f o r the f u l l  of the needle  n e e d l e was d i p p e d  submerged i n t h e  o f 4.0 m l . o f p i t u i t a r y  the p o s t e r i o r p o i n t  against  o f aqueous  I n j e c t i o n s were  administration  The n e e d l e was  f i n and were d r i v e n  of the l e n g t h  i n 10 l i t r e s  #22 h y p o d e r m i c n e e d l e  opposite  i n j e c t i o n s . Fourteen  per l i t r e ) .  o f two weeks u s i n g  syringe.  a precaution  the muscle  as a c o n t r o l .  facilitated  received  a l | inch  hormone.  hormone  and t h e h e a d was k e p t  This  with  The  received  were a n a e s t h e t i z e d  a period  dorsal  fish  Fish  over  side  was  Fish  used  anaesthetic.  the  and a f t e r an i n j e c t i o n ,  fish  made a s b e f o r e ,  plastic  o r no hormone  (Burrows e t a l , 1952).  o f (MS 222, 0.1  Each f i s h  little  was massaged i n an a n t e r i o r d i r e c t i o n t o a s s i s t  Fifteen  solution  was w i t h d r a w n f r o m t h e  ensured that  During  b) G r e a t C e n t r a l L a k e  untreated  the needle  injection.  losing  and t r a n s f e r r e d  fish.  50 Injection  of  17-b-Estradiol  Twenty f i s h injections  f r o m Shuswap L a k e r e c e i v e d  and 20 u n t r e a t e d A suspension  0.05 m l . o f c o r n using fish  o f 0.05 mg.  received  a total  o f 0.3 mg.  i n s i x equal  hormone. salt  Each  over  a 3  i n j e c t i o n s were a d m i n i s t r a t i o n of and m a i n t a i n e d  Pills  4 t o 5 months p r i o r  s a l m o n were o b t a i n e d fish  used  25 mg. e a c h were p l a c e d  was t h e n s p r e a d infection.  of approximately one-half  The f i s h  aqueous s o l u t i o n o f MS  from the i n c i s i o n  from Great C e n t r a l Lake.  on t h e m i d l i n e  t o the p e c t o r a l f i n s .  with  t o spawning, Seven-  as c o n t r o l .  incision  was made v e r t i c a l l y  222.  a b o u t two i n c h e s  were h e l d Two  an i n c h posterior  v e n t r a l s i d e up i n  Stilbestrol  pills  i n t h e body c a v i t y and p u s h e d the help  of a g l a s s r o d .  over the i n c i s i o n  F i s h were t a g g e d  as a p r e c a u t i o n  (B.D.H.) away  Aureomycin against  and t r a n s f e r r e d t o h o l d i n g  tank.  Stilbestrol,Powder Thirty  mixed  Further  mortality during  of S t i l b e s t r o l  unteated  Feeding  a 0.25 m l . s y r i n g e .  water.  A small  an  muscularly,  of 1 7 - b - e s t r a d i o l  doses.  Seventeen f i s h ,  teen  injected intra  F i s h were t a g g e d by f i n c l i p p i n g  Implantation  sockeye  as c o n t r o l .  of 1 7 - b - e s t r a d i o l per  o i l s u s p e n s i o n was  abandoned b e c a u s e o f h i g h  in  served  a #30 h y p o d e r m i c n e e d l e w i t h  week p e r i o d  the  fish  hormone  t a b l e t s of S t i l b e s t r q l  i n 1 3/4 pounds o f wet f o o d  i n g machine.  F o r t y grams of t h i s  (25 mg.  by an e l e c t r i c food  e a c h ) were food  was f e d t w i c e  grinddaily  51 t o twenty tagged fish  sockeye  salmon  over a p e r i o d  and m a i n t a i n e d i n s a l t  of the experiment  water  o f 14 d a y s .  f o r 8 weeks.  (3) were u s e d  The f i s h Untreated  as a c o n t r o l .  RESULTS Shuswap L a k e  Fish  Three  groups  of s e x u a l l y  e d s e p a r a t e l y w i t h salmon synthetic  Stilbestrol No a d u l t  tal  and c o n t r o l  were f o u n d in  worms were f o u n d  control  Effect  o f Crude All  days  fish  Fish  adult  fish.  whereas none were  found  ( T a b l e V. & F i g u r e IX)  Salmon P i t u i t a r y fish  treatment.  female  Philonema  (Table I V . ) .  died  Fifty  o n c o r h y n c h i were f o u n d i n f i f t e e n Sixty  1 7 - b - e s t r a d i o l and  O n l y two s u b - a d u l t f e m a l e  the i n j e c t e d  after  treat-  i n any o f t h e e x p e r i m e n -  i n the experimental f i s h ,  the t h i r t y  seven  pituitary,  were  between F e b r u a r y 23 and March 19, 1965.  groups.  G r e a t C e n t r a l Lake a)  immature s o c k e y e  Extract between s e v e n  adult  female  Pituitary  and f o r t y -  Philonema  injected  fish.  worms were o b t a i n e d f r o m t h e c o n t r o l  One g r a v i d worm was f o u n d  i n the control  fish  during  the t e n u r e o f t h e e x p e r i m e n t . Five  developmental  stages could  t h e u t e r u s o f t h e f e m a l e worms: larvae,  and t a i l e d  larvae  (free  be r e c o g n i z e d i n  e g g , m o r u l a , embryo, swimming).  No t a i l e d  tailless larvae  were f o u n d i n t h e f e m a l e worms between 7 and 12 d a y s o f t r e a t m e n t . Of t h o s e f i s h  that  died  between 19 and 47 d a y s  52  F i g . IX  E f f e c t of crude salmon p i t u i t a r y e x t r a c t s on the u c t i o n of l a r v a i n Philonema oncorhynchi. 0 r e p r e s e n t s the number of worms.  prod-  50|-  45f-  40 CD  35  30  0 CCD  25  o  o  to >-  §  a m mama  20  LU  ceo cam  cccco t IN  10 •ceo  j  EGG  cr»  i  MOR  0  CONTROL  •  PITUITARY  i  1 NO  1  EMB  TAIL  TAIL  DEVELOPMENTAL  STAGES  1  53 after of  t r e a t m e n t , 33 f e m a l e  t h e s e worms h a d t a i l e d  worms were c o l l e c t e d . larvae  f e m a l e worms h a d e g g s i n t h e i r ment  ( T a b l e V. and F i g u r e  larvae  uteri  IX ) .  after  male f i s h  sexes  changes  fish.  i n body w e i g h t  and u n t r e a t e d f i s h  None o f t h e  12 d a y s  of t r e a t -  a t P<^.05 l e v e l i n  The n o n s i g n i f i c a n t  i s p r o b a b l y due t o v e r y l o w sample  The treated  and f e m a l e  uteri.  Jhe, abundance o f t a i l e d  i n t h e f e m a l e worms i s s i g n i f i c a n t  b o t h combined in  in their  Eighteen  number.  and gonad w e i g h t  were n o t s i g n i f i c a n t .  Nuptial  was o b s e r v e d  after  t r e a t m e n t b u t no s u c h s e c o n d a r y s e x u a l c h a r a c t e r s  nuptial +++  i n the untreated f i s h coloration  (Pickford,  b) E f f e c t  Development of  died  between 173 d a y s .  i n 17 o f t h e S t i l b e s t r o l  basis  from + t o  Seventy implanted  sockeye.  No s t a g e s beyond o f t h e 70 f e m a l e  Philonema  implanted f i s h .  o f 60 a d u l t  t h e embryo were f o u n d  i n the u t e r i  o n c o r h y n c h i taken from t h e S t i l b e s -  A l s o , no d i f f e r e n c e  worms f r o m male and f e m a l e  hosts  was f o u n d  i n the  (Table VII.and Figure X ) .  worms r e c o v e r e d f r o m t h e c o n t r o l  f i s h had  was no d i f f e r e n c e  o r gonad  larvae. There  weight  were  of S t i l b e s t r o l  pre-spawning  tailed  (Table V I . ) .  two weeks  1953a).  f e m a l e worms were f o u n d  One  within  was r e c o r d e d on an a r b i t r a r y  A l l treated f i s h  trol  fish  of the  color  observed  i n male and f e m a l e  result  between t h e t r e a t e d  i n body w e i g h t  and u n t r e a t e d f e m a l e  fish.  Body  54  Fig.  X  Effect larvae  of synthetic i n Philonema  0 represents  s t i l b e s t r o l on t h e p r o d u c t i o n oncorhynchi.  t h e number o f worms.  of  50r  45  40 OCD  35  30  o OCD  aoocniD  COD  oocco  a) 25 <  UJ  20  1  ODD  I5h  0 03)  i  CD  0 CONTROL  "  • STILBESTROL  i  i  1  1  NO  EGG  MOR  EMB  TAIL  DEVELOPMENTAL  TAIL STAGES  »  55 weight  of  implanted  regression  male f i s h  of t e s t i s  ( T a b l e V I I I . and  was  Figure  d i d not  found XI).  i n the  100/body w e i g h t ) of  salmon d i f f e r e d  significantly  male  implanted  T e s t i s weight  (gonad w e i g h t X  treated  change b u t  the  a  notable  male  and  maturity  Stilbestrol  a t P<^.05 l e v e l  fish index  treated  from the  un-  fish.  DISCUSSION "In f o r m the the  the  first  majority  link  of a n i m a l s ,  between t h e  neurosecretary  environmental  p h y s i o l o g i c a l machinery concerned with  (Scharer  and  reproduction  Scharer,  1963).  The  the  (from  t o Arthropod) are  Protozoan  effects  endocrine  of host  ence the  number o f p a r a s i t e s p e r  directly  or  indirectly  the  r e g u l a t i o n of So  far, i t  hormones on  o f two host  g r o w t h and  has  parasites  kinds.  They  they  control  and  and  reproduction"  i n p a r a s i t e s i s p o o r l y known.  been f o u n d t h a t  trigger  cells  reproduction  of  influ-  the  parasites. Several cycle an  of the  and  of s y n c h r o n i z a t i o n  the  p a r a s i t e are  i n t e r e s t i n g phenomanon.  sporidian  species  i t s definite  (Smyth, 1 9 6 2 ) .  between t h e the  with  Cleveland  reproductive  moulting  life  cycle that  termite  hosts.  c y c l e of O p a l i n a ,  a haemois  host,  (1957) showed a parasitic  life  represent  simondi,  intermediate  c y c l e of the  c y c l e s of t h e i r  (1961) r e s t u d i e d t h e  a life  and  between t h e  known and  Leucocytozoon  p a r a s i t e o f d u c k s , has  synchronized  and  host  cases  Simulian  relation  flagellates Wessenberg  which l i v e s  in  56  Fig.  XI  E f f e c t o f i m p l a n t e d s y n t h e t i c s t i l b e s t r o l on the p r o g r e s s i v e growth o f t e s t e s o f s o c k e y e salmon f r o m the G r e a t C e n t r a l Lake.  I  if) |V)  I  O  I  1  1  1  <P  O  tp  O  OJ  CVI  —  —  X3QNI  Aiiamvw  L.  uO O  57 the rectum reproduce winter  o f f r o g s and t o a d s . a s e x u a l l y by f i s s i o n  and s e x u a l l y  water t o b r e e d .  He f o u n d  d u r i n g t h e summer, autumn and  Bieniarz  (1950) h a d s u g g e s t e d  and  parasite  activity, Mofty  (1956) s t a t e d life  (1964) showed  hormones,  integerrimum,  develop  Stunkard  i s related,  Gallien  of c y s t  not t o the l e v e l of of gonadal  hormones  of  Polystoma  o f an E u r o p e a n host  frog,  and p a r a s i t e this  the maturation  of  obserPolystoma  d e p e n d e n t on t h e host^s e n d o c r i n e t o demonstrate  o f Polystomum  of s c a r c i t y  cycle  (1935) c o n f i r m e d  (1951) shows t h a t  (1959) t r i e d  i n nature  stellai  of m a t e r i a l s .  the f i r s t  the r o l e  cycle.  of p i t u i t a r y i n  but d i d not suceed  G o r s h k o v (1964) r e p o r t e d  r e p r o d u c t i o n o f Polystomum  takes  n o t a t t h e e n d o f 3 r d , b u t a t t h e e n d o f t h e 2nd y e a r o f  life,  frog.  host  pituitary  the i n d u c t i o n  a n d he a l s o r e p o r t e d t h a t  i s almost  the m a t u r a t i o n  its  that  (1876) d e s c r i b e d t h e l i f e  Miretski  integerrimum  place  between  t o seasonal  but t o the l e v e l s  simultaneously.  vation.  that  the c o r r e l a t i o n  i n the u r i n a r y bladder  temporaria,  because  during copulat-  i n i t s host. Zeller  Rana  that  pituitary  c y c l e s was r e l a t e d  i n the o p a l i n i d s  gonadotropic present  this  and p r o b a b l y due t o t h e p r o d u c t i o n o f g o n a d o t r o p i n .  and Smyth  formation  that  i n d u c e d by t h e g o n a d o t r o p i c hormones o f  the h o s t r e l e a s e d by t h e a n t e r i o r Cehovic  the o p a l i n i d s  d u r i n g t h e s p r i n g when t h e h o s t s move t o  change was p r o b a b l y  ion.  that  i . e . one y e a r  Ontogenesis  prior  t o the f i r s t  spawning of the  and a n n u a l  sexual cycle  o f Polystomum  do  58 not  depend upon g o n a d o t r o p i n s Raikova  relationship  between t h e e g g  hydriforme  Polypodium  stolons  becomes r i p e .  fifth  and  The  sturgeon eggs.  of  (1956, 1958)  cyprinids  that migrate  because and  trematode  e v e r s i o n of  of the s t o l o n s  the  female  c o u l d be  the eggs t o  d e s c r i b e d the b i o l o g y of  G e n a r c h e l l a which l i v e s  the  The  pass  maturation.  trematode  time  close  b e f o r e s p a w n i n g when t h e  inversion  frog.  coelenterate parasite  i n j e c t i o n s which caused  stage  Szidat  of the  occurs j u s t  i n d u c e d by h y p o p h y s i s into their  s e x u a l hormones o f t h e  (1959) e x p e r i m e n t a l l y c o n f i r m e d a  Polypodium  fish  and  from  develops  i n the  intestines  sea t o f r e s h water. rapidly.  of i n c r e a s e d a c t i v i t y  and  He  of  During  concludes  development  the  that  this  of the  thyroid  hypophysis. R o t h s c h i l d ' s work  rabbits  showed t h a t  an e c t o p a r a s i t e  hormones f o r m a t u r a t i o n . most e f f e c t i v e rabbit  fleas  also rely  (1965b) h a s control  are c o r t i c o s t e r o i d s  and  on  shown t h a t  maturation  host the  i n the  estradiol.  of salmon p i t u i t a r i e s  sexual characters.  t e n d a y s and  separated color.  may  of  into  prespawning  from G r e a t C e n t r a l Lake s t i m u l a t e d development  secondary after  She  hormones t h a t  Injection sockeye  (1964b, 1965a) on f l e a s  from  pituitary  the r e s t  after  Nuptial injected 21  injected  adult  blueback  fish  d a y s on  B u r r o w s e t a l (1952) o b s e r v e d  pituitary  coloration  was  apparent  could readily  the b a s i s o f  spawning c o l o r  sockeye,  of  be  nuptial in  Oncorhynchus  nerka  59 a f t e r , 12 d a y s o f i n j e c t i o n . injection hastened  o f salmon p i t u i t a r y the development  development was j u d g e d 'tailed'  on t h e b a s i s  larvae  of Philonema  o f absence  second-  as 3 days  in pituitary  has s t i m u l a t e d the oncorhynchi.  or presence  injected  treated  of v i a b l e  worms. fish  Development  Development  i s very  signifi-  development  h a s a marked e f f e c t  significant erence  changes  was f o u n d  o f P ^ o n c o r h y n c h i may be s u p p r e s s e d in fish  i n the o v a r i e s of implanted  The r e s u l t s  extract  oncorhynchi  differs  or i n d i r e c t l y ,  Stil-  b u t no  fish.  No  diff-  significantly  d i s c u s s e d above s u g g e s t  s t i m u l a t e s t h e development  directly  sexes.  o f i m p l a n t e d and c o n t r o l  and m a t u r i t y i n d e x o f t h e f o r m e r  pituitary  o f both  on t h e growth o f t e s t e s ,  between body w e i g h t  the l a t t e r .  and f r o m  fish.  the i m p l a n t a t i o n of S t i l b e s t r o l  bestrol  by  and o t h e r  and r i p e n e d m a l e s a s e a r l y  i n the u t e r u s of female  Larval  from  markedly  when compared w i t h worms f r o m u n t r e a t e d c o n t r o l s  Stilbestrol  fish,  salmon  coloration  o f salmon p i t u i t a r y  of the larvae  o f P. o n c o r h y n c h i  by  adult  that  injection. Injection  cant  into  of n u p t i a l  ary sexual c h a r a c t e r i s t i c s after  P a l m e r e t a l (1954) s t a t e d  that the  of Philonema  whereas i t i s s u p p r e s s e d  Stilbestrol implantation. Experiments  Shuswap L a k e f a i l e d lack  of l i v e  Sub-adult imately  adult  female  fifty  using sexually  (Table IV). female  immature  sockeye  from  T h i s may have r e s u l t e d  worms i n t r e a t e d  worms were f o u n d  percent of the f i s h ,  and u n t r e a t e d  i n t h e body c a v i t y upon e x a m i n a t i o n  from fish.  of approxbefore  60 experiments  began.  But  a l l t h e worms were f o u n d e n c y s t e d i n  both e x p e r i m e n t a l  and  the  similar  situation  was  found  sockeye  i n Shuswap L a k e .  ing  fish  out  The  above f a c t s  Philonema of  the  control  oncorhynchi  i n J u n e , 1965.  A  i n p r e c o c i o u s male a d u l t  Only  o f 25 e x a m i n e d suggest  fish  one  cyst  was  found  (Margolis, personal  that  Philonema  spawning  i n a spawncommunication).  i n Shuswap L a k e a r e  which i s s y n c h r o n i z e d w i t h  the l i f e  o f P.  oncorhynchi  i s probably  synchronized  behavior  sockeye.  with  s e a s o n a l c h a n g e s i n p h y s i o l o g y and  Hoar  (1965a) d i s c u s s e d t h e s e a s o n a l c h a n g e s i n d e t a i l s .  spawn once d u r i n g t h e i r fourth  year  and  oncorhynchi  Philonema  but  bladder t i s s u e molt  d u r i n g the  sea  the  after  c o u l d not  be  spring  swimming.  this  fifth  i n the sea  There  in fresh  molts  o c c u r s d u r i n g the  first  The  o r soon a f t e r  w h i c h P^_ o n c o r h y n c h i oncorhynchi  first  stage  of the  occurs  the  first  gonadal  The growth  are e m i t t e d along with l a r v a e then  become  by C y c l o p s b i c u s p i d a t u s w h i c h  i n abundance d u r i n g t h i s p e r i o d . i s s e a s o n a l and  the  grows r a p i d l y .  parallels  i s correlated  Reproduction  with the  in  swim-  t i m i n g of  i t probably  the  water,  a r e two  but  G r a v i d P^_ o n c o r h y n c h i  Sockeye  i s g e n e r a l l y on year  cavity.  determined  They are e a t e n  oncorhynchi  of  w h i c h t h e l a r v a e move o u t  e g g s i n t o t h e w a t e r , and  available  the f i r s t  o f l a r v a e i n P^  fish.  and  follows i t closely.  i n t o the p e r i t o n e a l  first  production  cycle  i n some s t o c k i n t h e  molting period,after  in  life  larvae in f i s h :  autumn i n t h e  second  cycle  host. Development  P.  not  cycle  fish  freeare of of  P.  61 sockeye  salmon.  made by Hoar  I t i s an i n s t a n c e o f t h e g e n e r a l  (1966),  " I f r e p r o d u c t i o n i s s e a s o n a l or occurs  once  i n the animals  that  young a p p e a r when f o o d i s a b u n d a n t  optimal  the  cycle,  Seasonal  i t must be p r e c i s e l y  cycle  f o o d , and c h e m i c a l s may s e r v e  photo-  as t r i g g e r s f o r  changes."  i n t e r - r e l a t i o n s h i p between hormones o f t h e f i s h  work h a s been done on t h e p i t u i t a r y  of f i s h  timed s o  changes i n t e r m p e r a t u r e ,  g r o w t h and r e p r o d u c t i o n o f P. o n c o r h y n c h i  extensive  only  and o t h e r c o n d i t i o n s a r e  timing of the associated p h y s i o l o g i c a l The  and  life  for survival.  period, moisture,  statement  (Pickford  reviewed  the endocrine  He n o t e s  that  and A t z , 1 9 5 7 ) . regulation  the p i t u i t a r y  i s not c l e a r , but and r e p r o d u c t i v e  R e c e n t l y Hoar  (1965b)  of r e p r o d u c t i o n i n the f i s h .  i s necessary  f o r maturation of  gonads i n a l l v e r t e b r a t e s above t h e C y c l o s t o m e s .  Therefore, i t  may be s a i d  i s directly  or  that  indirectly  the maturation  dependent  on t h e p i t u i t a r y  w i t h some k i n d s o f f e e d b a c k tropic  hormones.  effect  effect  I t h a s been d e m o n s t r a t e d  of p i t u i t a r y  on t h e h o s t The  the  effect  work.  to s i x fold  that  by t h e i r  on t h e worms.  o f 1 7 - b - e s t r a d i o l on P_^ o n c o r h y n c h i  needs  1 7 - b - e s t r a d i o l h a s been r e p o r t e d i n  o f b o t h male and f e m a l e  show a f i v e  study  o f P^ o n c o r h y n c h i .  on p a r a s i t e s may be e i t h e r  o v a r i e s of the Oncorhynchus nerka  estrogens  and gonado-  in this  the maturation  o r by d i r e c t  effect  more e x p e r i m e n t a l  hormones o f f i s h o r  mechanism o f g o n a d a l  t h e p i t u i t a r y hormone e f f e c t s The  o f P^ o n c o r h y n c h i  (Hisaw, 1 9 6 3 ) .  Atlantic  increase a t t h e t i m e  The b l o o d  Salmon, Salmo  salar  o f s p a w n i n g and  62 reach f i v e The  t o s e v e n mg.  concentration  /100  of 1 7 - b - e s t r a d i o l  known, t h o u g h i t i s b e l i e v e d spawning P a c i f i c if  maturation  ml. b l o o d  Salmon  o f P.  that  (Cedar e t a l , 1961).  in Pacific  there  as i n A t l a n t i c  oncorhynchi  Salmon  i s not  i s an i n c r e a s e i n Salmon.  i n adult  I t i s unknown  salmon i s e f f e c t e d  by  17-b-estradiol. Implanted maturity  S t i l b e s t r o l had no a p p a r e n t e f f e c t  o f P_^ o n c o r h y n c h i  salmon, but had s u p p r e s s e d testes.  But o v a r i e s  Stilbestrol the  given  The  have  function of the p i t u i t a r y  the  b o t h t h e gonads  of f i s h  of  o f P^_ o n c o r h y n c h i  literature  histological and  hyperplasia  s e x u a l l y mature increased  salmon?  of the a d r e n a l (Robertson  (Hane  produce  A review of hormone  changes d u r i n g t h e i r N  concentration  that  levels  question.  ( R o b e r t s o n and<£al 1961  fish  s a l m o n ' s bfood  this  have  o f worms.  (Genus O n c o r h y n c h u s ) e x h i b i t e d  and p h y s i o l o g i c a l  spawning p e r i o d  marked  ing  i n adult  s p a w n i n g s a l m o n m i g h t answer salmon  and g o n a d a l  and t h e m a t u r i t y  c o n c e r n i n g changes i n d i f f e r e n t  Pacific  suppressed  o f g o n o d o t r o p i c hormones) w h i c h  What o t h e r hormones c o u l d be i n v o l v e d maturation  growth of  l a r g e doses of  t o r e p r o d u c i n g s o c k e y e may  (or suppression  regressed  and f e m a l e p r e s p a w n i n g  the normal p r o g r e s s i v e  are u n a f f e c t e d .  normal c o - o r d i n a t i n g  hormones  i n b o t h male  on t h e  a, b) .  cortical tissue  maturation T h e r e was  present  and W e x l e r 1 9 5 9 ) , and a  of 1 7 - h y d r o x y c o r t i c o s t e r o i d s and R o b e r t s o n , 1 9 5 9 ) .  The  both  a  i n the  greatly  i n spawn-  changes i n  63 concentration represented in  o f 17-OHCS f r o m t h e s e a t o t h e s p a w n i n g  an i n c r e a s e  o f a s much a s s e v e n f o l d .  s p a w n i n g f e m a l e s was 79 ^ug p e r c e n t ,  Robertson  and W e x l e r  i n the c o r t i s a l  during  period  completion the 41  i n m a l e s 31 /ug p e r c e n t ) .  concentration  immediately  of the spawning a c t .  concentration  (The a v e r a g e  (1960) and I d l e r e t a l (1959) a l s o f o u n d a  sharp increase a brief  grounds  of c o r t i s o n e  i n sockeye  preceeding  plasma  s p a w n i n g and d u r i n g  Female p l a s m a c o n t a i n e d and C o r t i s o l  i n male  plasma,  ^ u g and 26 ^ug/lOO m l . r e s p e c t i v e l y a s compared w i t h  and  11 /ug/100 m l .  extremely high metabolism  Idler  levels  e t a l (1959) s u g g e s t e d  o f c o r t i s o n e and C o r t i s o l  i n salmon, a t l e a s t  during  the r i v e r  T h e r e a r e o s m o t i c and m e t a b o l i c they and  enter  an e s t u a r y  fish?  a similar  Is there  of Philonema during that  this  osmotic  change.  of f i s h  oncorhynchi  the migration  that the regulate  mineral  migration.  changes i n f i s h  change  when  between s h a r p  oncorhynchi  along  increase  and t h e r a p i d d e v e l o p m e n t  with the o f 17-  of larvae  i n the p e r i t o n e a l c a v i t y of the host  of salmon?  t h e most e f f e c t i v e  Does P h i l o n e m a  and m e t a b o l i c  any c o r r e l a t i o n  OHCS i n t h e b l o o d  22 /ug  o r f r e s h w a t e r f r o m t h e s e a , and C o r t i s o l  cortisone regulate  also face  double  Rothschild  hormones c o n t r o l l i n g  (1965b) showed maturation  i n the  rabbit  flea  glands  u n d e r t h e c o n t r o l o f t h e a d r e n o c o r t i c o t r o p i c hormones  which ary.  a r e t h e c o r t i c o s t e r o i d s , p r o d u c e d by t h e a d r e n a l  are secreted  i n t u r n by t h e a n t e r i o r l o b e  of the p i t u i t -  64 There whether gonadal a role of  as y e t , t o  hormone o r hormones f r o m  a d r e n a l organ  i n the m a t u r i t y of Philonema  these  ism.  i s no e x p e r i m e n t a l e v i d e n c e ,  e x e r t an e f f e c t  So f a r , i t i s known t h a t  stimulating effect more t h a n mental  this  on  work i s t o be  oncorhynchi, through  the h o s t ' s  salmon p i t u i t a r y  I t i s obvious  that  done b e f o r e s p e c i f i c  oncorhynchi  play  or whether  extract  much more  any  metabolhas  t h e m a t u r i t y of P^_ o n c o r h y n c h i , b u t  i s known.  t h e m a t u r i t y o f P^ known.  indirectly  the  indicate  a  no  experi-  hormones r e q u i r e d f o r  i n anadromous s o c k e y e  salmon  are  65 SECTION IV Transplantation  o f Worm.  INTRODUCTION A second  series  o f e x p e r i m e n t s watte d e s i g n e d t o t e s t  the h y p o t h e s i s t h a t Philonema hormones o f s o c k e y e different and  salmon  o n c o r h y n c h i i s dependent  on  f o rp r o d u c t i o n of l a r v a e .  ages were t r a n s p l a n t e d  Worms o f  t o o t h e r h o s t s o f v a r y i n g age  species. The  information  purpose  o f t h e e x p e r i m e n t s was t o g a i n  on t h e d e p e n d e n c e  hormones f o r t h e d e v e l o p m e n t whether a s i n g l e The  o f P_._ o n c o r h y n c h i on h o s t of i t s larvae  stimulus or sustained  experiments had a d d i t i o n a l  tion  as t o the s u r v i v a l  different  further  stimulation  advantages  and g r o w t h  and t o d e t e r m i n e i s required.  of a c q u i r i n g  o f worm i n a l t e r e d  informas i t e s of  recipient hosts.  MATERIALS AND METHODS Worms u s e d a)  Adult  salmon  f o r Transplantation P_. o n c o r h y n c h i n o t y e t g r a v i d ,  a t S w e l t z e r Creek  months p r i o r b)  Third  of sockeye c) of  fish  collected  t r a p , C u l t u s Lake  from  sockeye  one t o two  t o spawning.  stage larvae  o f P. o n c o r h y n c h i f r o m t h e s w i m b l a d d e r  smolts from C u l t u s Lake.  Fourth stage larvae  o f P ^ o n c o r h y n c h i f r o m t h e body  a p r e c o c i o u s male s o c k e y e  salmon  from C u l t u s Lake.  cavity  66 Recipient a)  Hosts  Rainbow t r o u t  Coquitlam,  Rainbow t r o u t ,  c)  Steelhead,  were a t l e a s t  Sun V a l l e y T r o a t Farm,  2| years  of age, from  t h e above  t h e anadromous v a r i e t y two y e a r s  d)  Sockeye salmon  old  sockeye  e)  Gold  f)  of age, from  B.C.  b)  ercial  two y e a r s  o f age when  of  gairdneri,  (Oncorhynchus n e r k a ) .  Approximately  2\  year  i n 1964.  ( C a r a s s i u s c a r a s s i u s ) were o b t a i n e d f r o m  a comm-  pet shop.  Frogs,, (Rana p j p i e n s ) were s u p p l i e d b y V a n c o u v e r  The water).  o f a d u l t Worms i n t o  fish  c l o t h , h o l d e r , w h i c h was During  t h e -mouth w i t h  were p l a c e d on t h e i r  as  backs i n a  were i r r i g a t e d  o f MS  system, p r o v i d i n g approximately  long  (0.1 g m . / l i t .  a d j u s t e d a c c o r d i n g t o the s i z e  a 1:10,000 s o l u t i o n  c o u l d be m a i n t a i n e d  alive  of the through  222 by means o f a pump 10 l i t r e s p e r m i n u t e .  on t h e o p e r a t i n g t r a y  f o r as  necessary. An i n c i s i o n  was u s e d  fish  the o p e r a t i o n , the g i l l s  Fish  ventral  Fish  were a n a e s t h e t i s e d w i t h MS "222  The i m m o b i l i z e d  recycling  Biological  Victoria.  Transplantation  fish.  which  used.  o f Shuswap L a k e were o b t a i n e d a s s m o l t  fish  Suppliers,  line,  of about \  a v o i d i n g the p e l v i c  t o widen the i n c i s i o n . .  - 1 i n c h was made i n t h e m i d bones o f the f i s h . Required  were t r a n s p l a n t e d i n t o t h e body c a v i t y The  source.  worms were p l a c e d i n s i d e ,  either  A  retractor  numbers o f P h i l o n e m a  i n Ringer's  posterior  solution.  or a n t e r i o r t o  67 the  incision  had  t o be t a k e n  very  with  fragile.  treated  t h e h e l p o f a smooth g l a s s r o d .  i n p l a c i n g the female Sutures  surgical  wound w a t e r - t i g h t . switching thetic  from  of f i s h  a minimal  worms i n t o  was b r o u g h t  about  by  Both  anaes-  fish  pillow.  smolts  lining  and t h e h e a d was r a i s e d secure with rubber  fed naturally  infected  deflated  from  C u l t u s Lake through Free  solution  Transplantation  o f A d u l t and L a r v a l  Operations  and an i n c i s i o n the v e n t r a l out with  were s q u i r t e d  a p o l y e t h y l e n e stomach  i n t o the  Frogs  a retractor  tray  t o the o p e r a t i n g  o f 1 i n c h was made on t h e v e n t r a l abdominal v e i n s .  tube  tube.  Worms i n t o  The l i m b s were t i e d  bands.  the swimbladder i n  on f r o g s were made on a d i s s e c t i n g  anaesthesia.  with  swimbladders  ja smooth g l a s s  l a r v a e from  through  abdominal c a v i t y  s i d e up i n t o  was k e p t  amount o f R i n g e r ' s  were p u l l e d  ventral  The f i s h  i n t o the stomach.  under e t h e r  fish.  was s l i p p e d  stomach of f i s h  avoiding  at a  were a n a e s t h e t i z e d i n MS 222 (0.1 g m . / l i t .  a foam r u b b e r  was t h e n  sockeye  inserted  fish  with  a foam r u b b e r fish  of l a r v a l  The i m m o b i l i z e d  the t r o u g h  tray  silicone  temperature.  The water) .  were  o t h e r t o make t h e  a n a e s t h e t i c t o d e c h l o r i n a t e d water.  Transplantation  of  Recovery  as they  "measuroll" black  t o each  care  and d e c h l o r i n a t e d w a t e r was a e r a t e d and m a i n t a i n e d  constant  The  worms i n s i d e  were made w i t h  thread very close  Great  The s k i n  side,  and body w a l l  and worms were p l a c e d i n t h e  of the f r o g with the help of a p a i r  of f o r c e p s .  68 Larvae  were t r a n s p l a n t e d i n t h e a b d o m i n a l  Ringer's solution out  with a medical pipette.  i n t h e same manner a s i n t h e F r o g s were m a i n t a i n e d  spray  of water,  in a controlled  cavity in  S u t u r i n g was  carried  fish. i n a wooden box environment  with a  room a t  light  10°C.  RESULTS Transplantation sockeye  salmon Two  adult  of a d u l t into  immature r a i n b o w  trout.  t o f o u r non-gravid female  of each  o f 37  and  matured  (Table  IX.)  three to  five  female  P^  after  the second  i n which female  oncorhynchi  female uterine female Male F\  died  between 2 and  The fish  two  f o r 114  worms t h a t  female  survived  show any  c o n t e n t s , whereas i n the normal  oncorhynchi  as a c t i v e  as t h o s e  days.  female  days.  had  reached  the  fish  one female  A-lthough  f o r a longer  i n immature  development of  erence  i n the l o n g e v i t y  female  recipient  the normal  final  sockeye  o f P_„ o n c o r h y n c h i  h o s t s was  the  spawning h o s t a l l stage of m a t u r i t y .  r e c o v e r e d from the t r a n s p l a n t e d host from  No  fish  The  f o r n i n e d a y s i n male f i s h .  days d i d not  P^_ o n c o r h y n c h i  the  114  w i t h the e x c e p t i o n of  i n b o t h male and  female  f o r 114  day,  worms s u r v i v e d  survived  M a l e worms s u r v i v e d period.  trout  o n c o r h y n c h i were r e c o v e r e d f r o m  which d i e d  abdominal  fish.  Transplanted  P.  oncorhynchi from  male P_;_ o n c o r h y n c h i were t r a n s p l a n t e d i n t o t h e  cavity  fish  Philonema  insignificant.  host.  The  i n t h e male  were  diffand  69 It ion  that  was  o b s e r v e d d u r i n g the  b o t h male and  toward the  anterior  around the  liver  t o be  more a c t i v e  recipient  the  cavity  days a f t e r  arbitrarily in a)  the  abdominal c a v i t y ,  p y l o r i c caecae.  through  the  Male worms were control  t o come out  abdominal c a v i t y  tissue  Visceral  and  of  a d h e s i o n was  T h r e e k i n d s of  of  found  the  incisions.  transplantation.  solid  entangling  c h a n g e s were s e e n i n r e c i p i e n t  f r o m + t o +++.  C y s t s of  the  Female worms were s e e n  Pathological 77  of  except-  o n c o r h y n c h i moved  t h a n f e m a l e worms i n b o t h t h e  hosts.  peritoneal  female Philonema  portion  and  autopsy without  transplanted  cysts  were  hosts recorded found  trout:  i n which the  worms c o u l d  hardly  be  identified. b) on c)  F e m a l e worms e n c a s e d the  liver  Soft,  or  the  salmon  stomach of  (Figure The  recipient and  lake),  were of  stage any  the  rate  of  control  single  group of  female  Stage Larvae  of  Philoneifla o n c o r h y n c h i  worms.  and  sockeye  X.)  development  of  P_^ o n c o r h y n c h i i n  salmon, rainbow t r o u t ,  sockeye  (naturally  same p a t t e r n  (Figure  fish  variability fish.  either  of  transplanted  great  attached  tissue  Table  sockeye  the  and  s t e e l h e a d , rainbow t r o u t , and  a l l the  larvae,  Third  XII  hosts,  trout,  Although  of  tissue  wall.  partly disintegrated  Transplantation i n t o the  body  in a fibrous  and  were i n f e c t e d  i n g r o w t h was  Third,  fourth,  steelhead  infected  XII  and  the  in  Table with  the X.). third  observed sub-adult  in P.  70  Fig.  XII  Development of Philonema transplanted salmonids.  oncorhynchi  in different  350  250 L A R V . IN S W I M B L A D D E R  D  HI  1  150  1T» S T A G E L A R V . IN COELOM O" WORMS IN C O E L O M  2  6cJ  z  55  J  i  Is p  I i- 50 O LU _1  Z < LU  2  45 40 -  la  35  li Is  |i  to 3 0  Is  i  E 0!  2  Is  Is Is Is I  25  LU  1  20 15  i i  li  10 5 Control-1  Control-2  SOCKEYE  E H STEELHEAD  PAIN BOW  1L  SOCKEYE  WORMS IN C O E L O M  T I M E IN D A Y S  71 o n c o r h y n c h i were f o u n d sizes of  of l a r v a e  fish  in a single  were f o u n d  i n the swimbladders  w i t h the e x c e p t i o n of the  the  infection  was  found  was  55 d a y s o l d .  i n t h e body c a v i t y  days of t r a n s p l a n t a t i o n .  transplanted f i s h .  Two  of a l l groups  Shuswap L a k e s o c k e y e  One  adult  female  o f a male r a i n b o w  In a l l g r o u p s  where  oncorhynchi  trout  of f i s h ,  after  232  male worms  were g e n e r a l l y more p r e v a l e n t i n t h e body c a v i t y  than  female  worms. Third of  rainbow  hours of  trout  after  sockeye  The  stage l a r v a e  were r e c o v e r e d f r o m  the  when t h e y were a u t o p s i e d a t 16,  24,  b e i n g f e d with the n a t u r a l l y  infected  and  live  larvae  stomach of the r e c i p i e n t  host not l a t e r  Transplantation  stage Larvae  of t h i r d  i n t o t h e body c a v i t y  of  Approximately  10  were f o u n d  than  10  of Philonema  35  larvae  trout  hours,  40  bladder  11  (Salmo g a i r d n e r i i ) .  larvae  8 days,  the v i s c e r a l  13  respectively  of the r e c i p i e n t mass o f t h e  i n the  from the swimbladders  five  and  hours.  trout  of  4,  92  oncorhynchi  s m o l t j were t r a n s p l a n t e d i n t o t h e body c a v i t y  hours,  and  hours.  sockeye  7  36  swimbladders  s a l m o n , b u t none were r e c o v e r e d a t 5 and  u n d i g e s t e d swimbladder  swimbladders  days,  and  were e x a m i n e d 14  days,  were r e c o v e r e d f r o m  hosts. fish.  Fish  and the  of  of  each  after 0,  5,  swim-  Dead l a r v a e were f o u n d  in  6,  .72 Transplantation  of f o u r t h stage l a r v a e  i n t o t h e s t o m a c h and  body c a v i t y o f t r o u t a)  Approximately  of each  of the f i v e  intervals; recipient  trout.  no l a r v a e hosts.  those f i s h b)  30 l a r v a e were s q u i r t e d i n t o t h e stomach F i s h were s a c r i f i c e d  were f o u n d  i n the swimbladders  Dead l a r v a e were f o u n d  which  30 f o u r t h s t a g e l a r v a e  i n t o t h e body c a v i t y o f e a c h  o f the f i v e  e x a m i n e d 1, 3, 5, 50,.and 223 d a y s No l a r v a e were f o u n d with  the exception  stage all  l a r v a was  fish.  average  The l a r v a e  length  Transfer  of the f o u r t h f i s h Larvae  o f 11.78  of i n f e c t i o n  lived  i n the l a s t  by  One  n a t u r a l l y i n f e c t e d sockeye  smolt from  autopsied  f o r two d a y s  tank.  trout.  i n which  F i s h were  one l a t e  third  i n t h e body c a v i t y o f  fish  e x a m i n e d had an  smolt  Both  rainbow  t r o u t o f 24.0  i n two s e p a r a t e t a n k s . o f 8.5  cm. l o n g  t r o u t chased  the a n t e r i o r end w i t h i n  a t 19 and 32 h o u r s ;  was  and c a p t u r e d  10 m i n u t e s .  F i s h were  53 and 22 l a r v a e r e s p e c t i v e l y  were r e c o v e r e d f r o m t h e s w i m b l a d d e r s . were a l s o f o u n d  transplanted  predation  l o n g were s t a r v e d  the  were  mm.  cm.  i n t o each  hours.  o f any o f t h e f i s h  Two h a t c h e r y - r e a r e d u n i n f e c t e d  transferred  of the  after transplantation.  i n the swimbladder  found.  hour  i n the stomachs o f  were e x a m i n e d a t 24 and 48  Approximately  a t 24  i n the stomachs of both  Dead and l i v e fish.  larvae  73 Transplantation oncorhynchi a)  of Adult  and F o u r t h  i n t o F r o g s , Rana  A l l transplanted  adult  stage  16, and 18 d a y s .  f e m a l e P^_ o n c o r h y n c h i were  O n l y one a d u l t  frogs.  Cyst  occurred  The  f e m a l e worms were e n c a s e d  and  the cysts  liver.  No c y s t  i n t h i n brownish  was f o u n d  i n f r o g #4, b u t l o o s e  i n t h e body c a v i t y .  b)  larvae  Fourth  planted  stage  Xlb).  No v i s c e r a l a d h e s i o n  Transplantation of Gold Ten  i n t h e r e c i p i e n t abnormal  of Fourth  (  Stage Larvae  i n one l a r g e  decomposed  sacrificed, pathological host.  i n t o the Abdominal  were a u t o p s i e d Five  i n t o each  were f o u n d  of 6  fish.  5, 10, 4 7 , 6 7 , 7 5 , a n d 80 d a y s (2 d e a d ) , 6, and 2 l a r v a e  f r o m t h e body c a v i t y o f t h e f i r s t  No l a r v a e  or t o the  fish  transplantation.  recovered  f r o g was  o r any o t h e r  worms were t r a n s p l a n t e d  Fish  ly.  fibrous tissue  i n t h e body c a v i t y o f t r a n s -  f r o g s f o r 18 d a y s when t h e l a s t  c h a n g e s were o b s e r v e d  after  survived  f e m a l e worms.  t o t h e body w a l l  worms were f o u n d  Cavity  in a l l recipient  Worms i n f r o g #5 were f o u n d e n c a p s u l a t e d  cyst.  (Table  (Table X I a ) .  around the a d u l t  were e i t h e r a t t a c h e d  after  male worm was r e c o v e r e d  a d h e s i o n was a p p a r e n t  formation  found  were a u t o p s i e d  a l i v e . - a f t e r 13 d a y s o f t r a n s p l a n t a t i o n , Visceral  of Philonema  pipiens  t o be dead and e n c y s t e d when t h e f r o g s 13,  larvae  i n any f i s h  3 fish  were  respective-  a f t e r 47 d a y s o f t r a n s -  plantation. No v i s c e r a l adhesions o r any o t h e r c h a n g e s were  observed.  pathological  74 DISCUSSION Philonema gically than  adapted  oncorhynchi  three years.  oncorhynchi  During t h i s  i s correlated  and  to establish  of the r e c i p i e n t  h o s t s and  of a r a b b i t  animals  showed any  and  that  f o r more  of the  i n the  died within  a male and  microfilariae  (  host.  were t r a n s p l a n t e d i n t o  abnormal f r o g h o s t s , the  Setaria  grows  m a t u r a t i o n of  w i t h the m a t u r a t i o n  themselves  (1958) t r a n s p l a n t e d b o v i n e cavity  I t i s found  oncorhynchi  immature t r o u t  worms f a i l e d  physiolo-  p e r i o d P^_ o n c o r h y n c h i  salmon.  When a d u l t P^ sexually  and  f o r r e s i d e n c e i n anadromous s o c k e y e  a l o n g with the sockeye P.  i s physically  female  abdominal  cavity  a short period.  digitata female  i n t o the  goat.  Victor  peritoneal  None o f  i n t h e b l o o d and  the  none o f  the  a d u l t s were r e c o v e r e d . In t h e p r e s e n t s t u d y two survived er due  f o r 114  period.  The  to their  to their  female  large  active  defensive  d a y s and  gically  specific will  oncorhynchi  survived  for a long-  survival  o f male worms may  be  due  the  host. P.  oncorhynchi  survived  i n immature t r o u t .  worms i n s p a w n i n g s o c k e y e that  adult  i n immature  f o r 114  In a  days  similar  w o u l d have become  P_._ o n c o r h y n c h i  t o the donor sockeye  n o t mature  i s probably  w h i c h e n a b l e s them t o e s c a p e  female  This suggests  therefore  The  become g r a v i d  a l l female  gravid.  size.  mechanism o f t h e  they d i d not period  males g e n e r a l l y  female  worms a r e v e r y s l u g g i s h w h i c h  motility  Although  adult  i s physiolo-  host f o r m a t u r i t y  and  recipient  It  trout.  75 a l s o be p o s s i b l e t h a t l a r v a l dependent  on c o n t i n u e d  of the matured h o s t Larka Setaria  Noda  o f P_. o n c o r h y n c h i i s  s t i m u l a t i o n o f some e s s e n t i a l  which are m i s s i n g  hormones  i n t h e immature  host.  e t a l (1963) t r a n s p l a n t e d male and f e m a l e  species into rabbits.  were p r e s e n t and  production  i n the r a b b i t  (1960) f o u n d  Microfilariae  of  cervi  b u t showed no p e r i o d i c i t y .  that Philophthalmus  Alicata  from c h i c k s  reached  maturity  and p r o d u c e d e g g s i n h e t e r o t r a n s p l a n t e d e y e s o f r a t s  at  t h e same t i m e  about  Third the  l a r v a e of  oncorhynchi  s w i m b l a d d e r when t r a n s p l a n t e d i n t o e i t h e r  body c a v i t y The  stage  as i n c h i c k s .  of t r o u t  t o those Fourth  to live  i n donor stage  to infect  the f i s h  fish  were  the stomach.  l a r v a e have l o s t  original  continued  (donor), but they  when i m p l a n t e d , i n t o  words, t h e f o u r t h s t a g e  stage  i s com-  o f t r a n s p l a n t e d t r o u t and  as i n the sockeye h o s t  back t o t h e i r  host  l a r v a e f r o m t h e body c a v i t y  i n the abdominal c a v i t y  It  t h e stomach or  hosts.  developed  migrate  their  capacity to  t o observe  that fourth  site.  i s not very s u r p r i s i n g  f o r 47 d a y s and i n f r o g s f o r 18 d a y s . have l i v e d Generally,  so f a r there  longer  unable  In other  l a r v a e have s u b s i s t e d i n t h e p e r i t o n e a l c a v i t y  probably  into  and s u r v i v e d a s l o n g a s t h e f i s h d i d .  r a t e o f growth o f l a r v a e i n t h e r e c i p i e n t  parable  migrate  Larvae  of gold  would  i n f r o g s i f the h o s t s had s u r v i v e d .  salmonids  serve  as h o s t s  a r e no r e p o r t s o f P h i l o n e m a  f o r Philonema  i n hosts  other  than  76 salmonids reported author but  w i t h t h e e x c e p t i o n o f s t i c k l e b a c k s w h i c h have from  has t r i e d  failed  or because fore  t h e U.S.S.R. by S p a s s k y to infect  t o do s o .  of fourth  cavities  of g o l d f i s h  cavities  of these  observed  visceral  adhesions  failed  of female  encapsulates  (1937),  infected  Encystment of trout  Prespawning  Needham  ( 1 9 6 5 ) , and  p a t h o g e n i c i t y and v i s c e r a l with  adhesions  Philonema  to observe  any p a t h o -  study. P. o n c o r h y n c h i  i n the abdominal  and f r o g s i s p r o b a b l y due t o h o s t  between h o s t resulted  much s h o r t e r p e r i o d t h a n  and  of l a r v a e .  a l s o observed  of t h i s  Philonema,  nutri-  adhesions  i n the trout  t h e body c a v i t y  adult  the p e r i t o n e a l  visceral  s p e c i e s of t r o u t  Incompatability in  that  exhibited  Richardson  agubernaculum but the author  cavities  specificity  i s c l o s e d . There-  i n high sea salmon.  The a u t h o r  (1966) o b s e r v e d  genicity  host  larvae i n the p e r i t o n e a l  and f r o g s s u g g e s t  adhesions  fishes.  different  stage l a r v a e  (1965)c^iMargolis ( p e r s o n a l communication)  1964).  similar  The  a b d n o r m a l h o s t s c a n meet some e s s e n t i a l  visceral  (Platzer,  in  of gold f i s h  stage  male and p r e c o c i o u s male s o c k e y e  Pippy  third  and o t h e r p h y s i c o - c h e m i c a l r e q u i r e m e n t s French  in  with  T h i s may be due t o e i t h e r  o f the pneumatic duct  the s u r v i v a l  tional  gold f i s h  e t a l i n 1961.  been  the host  ( f r o g ) and i m p l a n t e d i n death  i n trout.  nematodes  of the p a r a s i t e s In response  a c t i v e movement.  in a  t o implanted  p r o b a b l y r e a c t s around  i t . Male worms m o s t l y e s c a p e  p r o b a b l y because of t h e i r  reaction.  the p a r a s i t e  encystment  I t i s easier for  77 the host  t o develop e p i t h e l i a l  outgrowths  inactive  parasites  male worms.  differ other  from  the adult  than s i z e  female  than  females  worms i n f r o g s a p p e a r  of phagocytes trout  Fawcett  similar  with the bulk of f o r e i g n  w i t h t h e worms may c o n t r i b u t e  p h a g o c y t o s i s and e a r l y Reichkenbach-Kline does n o t f o l l o w  that  as a r e a c t i o n  by  tissues  to a specific  i n the presence  f e e d i n g the small i n f e c t e d  Salmonidae. in  Third  the swimbladder  naturally smolt  infected  stage  sockeye  of  trout  smolt.  fish  i n the r e c i p argued by  intro-  efficient  On t h e o t h e r hand, formation"  factory.  I t only  chemical stimulus  from  one h o s t  suffered  t o another  to large carnivorous oncorhynchi 19 h o u r s  are found  after  This suggests  a meal o f  that  sockeye  can a c t as p a r a t e n i c h o s t ^ t o  T h i s would p r o b a b l y h e l p t o e x p l a i n  f e e d on p l a n k t o n .  stroma  Bacteria  the i n f e c t i o n s  o f P. a g u b e r n a c u l u m i n l a r g e non-anadromous t r o u t not  Goodchi  of p a r a s i t e s . "  fish  larvae  of rainbow  or other i n f e c t e d  Philonema.  in a  the "cyst  t o mechanical  P h i l o n e i q a c a n be t r a n s f e r r e d by  r e p o r t e d by  t o t h e prompt,  demise o f p a r a s i t e .  as a response  way  the host responses are  implant.  (1955) s u g g e s t s t h a t  appears fish  to that  The f o r m a t i o n o f c y s t s  e t a l (1947) who s u g g e s t e d  may  and d e c o m p o s i t i o n o f  " p a r a s i t e s became immeshed  and f i b r e s " .  or  biological  and f r o g may be due t o t h e same r e a s o n  correlated duced  Adhesions  immobile  Male worms  i n some i m p o r t a n t  and a c t i v i t y .  ( 1 9 5 4 ) , where he f o u n d  ient  active  around  w h i c h do  78 "Host s p e c i f i c i t y but  less  ates  well recognized  in parasitic  organ  of the h o s t /  specificity  larvae either intact  fish  host  the  body c a v i t y ,  bladder.  host  and o r g a n  when t r a n s p l a n t e d . stage  but the f o u r t h stage hosts  I t suggests  P. o n c o r h y n c h i  specificity. Third  stage  or from swimbladder  of  from t h e stomach i n t o t h e swimbladder o f  the t h i r d  of r e c i p i e n t  oper-  in specific  h a s been d i s c u s s e d i n s e c t i o n I I .  migrated  recipient  cavity  s t u d i e s both  which  1 9 5 4 ) . P„ o n c o r h y n c h i h a s  from d e f l a t e d swimbladder  the  swimbladder  t o and m a i n t e n a n c e  (Goodchild,  8  i n Parasitology  i s an o r g a n s p e c i f i c i t y  orientation  shown i n t h e p r e s e n t Host  i s well recognized  without  that t h i r d  When t r a n s p l a n t e d  larvae migrated larvae lived migrating  i n t o the  i n t h e body  i n t o t h e swim-  and f o u r t h s t a g e  a r e v e r y much s p e c i f i c  into  to their  larvae of  site  and h o s t .  Summary Transplanted gravid the  i n immature  experiment.  transplanted  a d u l t P_._ o n c o r h y n c h i  recipient  Adult  stage  d u r i n g t h e 114 d a y s o f  worms d i e d i n a s h o r t e r p e r i o d when  i n abnormal host  Third  host  d i d n o t become  (frog).  l a r v a e had migrated  i n t o the t i s s u e of  s w i m b l a d d e r when t r a n s p l a n t e d i n t o t h e s t o m a c h and t h e c o e l o m of  fish.  Fourth  stage  l a r v a e had l i v e d  were f l u s h e d o u t f r o m t h e s t o m a c h .  i n the coelom but  79 SECTION Development infected  sockeye  V.  of P h i l o n e m a  of R i v e r s  oncorhynchi  in naturally  Inlet.  INTRODUCTION The  third  line  of e v i d e n c e  bearing  hormonal r e g u l a t i o n of l a r v a l p r o d u c t i o n is  t o be  found  in certain  British  l e n g t h of t h e i r  commercial  Columbia,  a d u l t sockeye  of ocean r e s i d e n c e .  spawn i n t h e i r 6th  year.  P.  Rivers  the  hypothesis  its  host.  and  as  f i s h r e t u r n t o the  old  in  the  the  few  f i s h can  f i s h population should  occurrence  spawning sockeye  (merely)  a very  result  f i s h and  the  host  population. test on  hormonal  contain maturing the  ( S e c t i o n I ) may  developmental  their  groups.in  On  3  to  used t o  o f l a r v i g e r o u s worms i n t h e  a 3 year  2 or  oncorhynchi  a l l age  a coincidence  to  after  in this  o f P^  on  cycle.  Inlet,  f i s h return in  spawning grounds.  of C u l t u s L a k e of  sockeye  males r e t u r n  t h e r e f o r e be  i t s l a r v a e , then  Inlet  year,  precocious  worm i s d e p e n d e n t  t o produce  hand, i f the  or.5th  of hormonal dependence  I f the  of  at R i v e r s  i s known t o o c c u r  Inlet  Rivers the  4th  of  oncorhynchi  salmon u s u a l l y r e t u r n  A few  oncorhynchi  The  stimulus  3rd year  question  reproductive  salmon f i s h e r y  Owikeno L a k e t o spawn i n t h e i r years  i n I\_  natural populations  salmon which d i f f e r i n the In t h e  on  the  worms other 4  year  be  between a 4 y e a r  cycle  p a t t e r n i n the  worm,  80 only  one  contain  age  class  the g r a v i d  of the R i v e r s  from  longitude  Inlet  fish  salmon  popula-  of development  1965  of  nerka  1965;  56  H  in April, fishing  Government  number 9 r e s p e c t i v e l y . the f i s h  Inlet  origin  a r e a number 9 o f the  i n October,  and  5^ s o c k e y e  and  were aged  and  1965  1965  spawn-  in  area  frozen. as h a v i n g been  scale c h a r a c t e r i s t i c s  Laboratory, B i o l o g i c a l  Research Board  and  0  identified  by d i s t i n g u i s h i n g  Rivers  Department  fishing  were k e p t  were 02"  were c o l l e c t e d  commercial  A l l the f i s h  48'  f r o m t h e mouth o f  of Canada, i n August,  by t h e members o f M e r i s t i c the F i s h e r i e s  Oncorhynchus  39'  f r o m Owikeno Lake  All  salmon,  Ocean between l a t i t u d e  f r o m t h e Owikeno Lake  of R i v e r s  Inlet  the N o r t h P a c i f i c  P r e c o c i o u s sockeye August,  the s t a t e  year o l d sockeye  i n the commercial  of F i s h e r i e s , ing  of the R i v e r s  METHODS  Four captured  should  harboured.  MATERIALS AND  and  classes  were e x a m i n e d t o d e t e r m i n e  worms t h e y  population  Philonema.  V a r i o u s age tion  Inlet  o f C a n a d a , Nanaimo,  Station  of  British  Columbia. In t h e p r e s e n t s t u d y , t h e G i l b e r t age  designation  cates in  total  which  age  i s used,  the l a r g e  when c a u g h t  and  (1913) s y s t e m  type of A r a b i c  the s u b s c r i p t  i t migrated from the f r e s h  water.  figure  the year of  of indilife  81 Collection  and Measurements o f A d u l t P h i l o n e m a Frozen  and  fish  autopsied with  pectoral  fins  were l e f t  a mid-ventral  t o the anus.  bowl o f R i n g e r ' s  solution.  membrane were s e p a r a t e d slide.  The t i s s u e  Another  piece of s l i d e  out  swimbladder  pressed.  n i g h t a t room  incision  Complete  from  viscera  The s w i m b l a d d e r  temperature  t h e base  of the  was p l a c e d i n a and p e r i t o n e a l  and mounted on a 25 x 13 x 3 mm.  was s p r e a d  on t h e p l e x i  o f t h e same s i z e  slide  with  needles.  ends t o keep t h e t i s s u e  was e x a m i n e d f o r l a r v a e u n d e r a b i n o c u l a r  microscope.  The v i s c e r a  was t e a s e d  apart i n  Ringer's  solution  and t h e a d u l t worms were s e p a r a t e d o u t .  Ringer's  solution  with  allowed  to settle.  c o n t e n t s was p o u r e d  When s e t t l e d ,  into  o u t and a l l t h e s e d i m e n t washed t w i c e  colour  f o reasier  examination  All  was  t o remove t h e d a r k  a partitioned  After, plexiglass  A l l t h e worms were  and s t o r e d i n a v i a l .  t h e a d u l t worms were c l e a r e d i n l a c t o p h e n o l f o r  2 t o 5 days beforeimeasurement. and  into  and e x a m i n e d f o r s m a l l P h i l o n e m a . i n 80% ethanol  solution  under t h e microscope.  w a s h i n g , t h e c o n t e n t s were p o u r e d  presetted  a c y l i n d e r and  the supernatent  poured  petridish  plexi  was p l a c e d on t h e t e a s e d  and s c r e w e d down on b o t h  The p l a t e  dissecting  over  oncorhynchi  The t o t a l  l e n g t h o f t h e male  immature P h i l o n e m a were m e a s u r e d by t h e method u s e d f o r  large  l a r v a e as d e s c r i b e d i n S e c t i o n I . Measurements o f t h e t o t a l  were c a r r i e d slide  board  l e n g t h o f t h e a d u l t worms  o u t on t h e m i l l i m e t e r s c a l e under a b i n o c u l a r  microscope.  mounted on a p l e x i  82 RESULTS (Tables XII, XIII, The  and X I V . F i g u r e s X I I I , X I V , XV)  adult  sockeye  i n Owikeno L a k e , B r i t i s h t o e a r l y December  The and m a t u r i t y itude 1.49  Columbia  i n their  t h r e e y e a r s o f ocean  salmon  of R i v e r s during  fourth  Inlet  the p e r i o d ,  or f i f t h  average  fork  length,  index of the f i s h  year, after  body w e i g h t s , gonad  captured i n A p r i l ,  f o r male f i s h  9.45 gnu , and 0.81 r e s p e c t i v e l y  Three males  and one f e m a l e were f o u n d immature  streams  a t t h e time  1965  only three  and gonad  (Table X I I . ) . Owikeno Lake weight of f i s h 73.4 gm.  4  2  prior  and f i v e  of R i v e r s  fish  were  into  y e a r s o f age c a p t u r e d i n A u g u s t , Inlet  were d e f i n i t e l y  mature and  The f o r k  length,  w e i g h t s o f 52 s o c k e y e were h i g h e r t h a n 4g  sockeye o f O c t o b e r , 1965 t o spawning.  but increased increased  These  spawning.  t o f o u r months b e f o r e t h e s p a w n i n g .  body w e i g h t  fish.  of capture a c c o r d i n g t o the m a t u r i t y  and s i x t o n i n e months b e f o r e  from the o u t l e t  gm.,  and t h e r e s t  t o s i x months b e f o r e t h e e n t r a n c e  of four  weights  ,1965 a t l a t -  f o r female  ( G o d f r e y 1961 and I s h i d a e t a l 1 9 5 8 ) .  Fish  two o r  and 48.0 cm.,  1170.0 gm.,  approximately f i v e  September  39' 56" was 44.4 cm., 1002.8  gm. and 0.14 r e s p e c t i v e l y  indie:es  spawns  residence.  48' 02" and l o n g i t u d e  'maturing'  origin  weight  They  of ovary.  were c a p t u r e d f r o m  showed d e c r e a s e d body The mean w e i g h t  f r o m 1.49 gm, when c a u g h t  in April,  of t e s t e s 1965 t o  gm. i n O c t o b e r , 1965, and t h e o v a r y f r o m 9.45 gm. t o 327.3 (Figure  XIII).  83  Fig.  XIII  R e l a t i o n s h i p between t h e d e g r e e o f m a t u r a t i o n o f s o c k e y e salmon o f R i v e r s I n l e t and t h e t i m e o f catch.  20  TIME OF CATCH  84 The 1965  p r e c o c i o u s sockeye  in their  were f o u n d mature  i n August,  t h i r d y e a r , h a v i n g mean t e s t e s w e i g h t  o f 20.6 gm.  and m a t u r i t y i n d e x o f 3.63. An oncorhynchi  e s t i m a t e o f growth from both sexes  by c a l c u l a t i n g the  fish  a series  i n size  of f i s h  from a n t e r i o r  against  t h e age and t i m e  i n distance  of nerve  e n d and t o t a l oesophagus o f d i f f e r e n t  worms o b t a i n e d f r o m  the d i f f e r e n t  t i m e s o f t h e y e a r and d i f f e r e n t  P.  were o b t a i n e d s e p a r a t e l y  of r e g r e s s i o n s  were c a p t u r e d . The g r o w t h  o f male and f e m a l e  sexes  ring  sexes of  of the f i s h  at d i f f e r e n t  age c o m p o s i t i o n o f s o c k e y e  (Table XV). The oncorhynchi the  progressive  growth  are s i g n i f i c a n t  i n the s i z e  of Philonema  i n b o t h male and f e m a l e  f i s h of  same a g e , ( i . e . 4g) when c a p t u r e d i n A p r i l , A u g u s t , and  O c t o b e r , 1965, b u t n o t s i g n i f i c a n t when compared different captured  age g r o u p s  of f i s h  ( i . e . 32's, 2 4  S  '  a  n  against d  5  2  S  ^  a t t h e same t i m e . Growth i n d i s t a n c e  from the a n t e r i o r  of nerve  ring  and t o t a l  e n d o f worms were d e p e n d e n t  oesophagus  on t h e s i z e o f  worms b u t were u n a f f e c t e d b y t h e age o f t h e f i s h or worms. I n other  words, the d i s t a n c e  agus i n c r e a s e An of The in  female growth  t o nerve r i n g  when t h e t o t a l l e n g t h apparent r e l a t i o n s h i p  Philonema  o f worms exists  Philonema  growth  of  oesoph-  increases.  between t h e g r o w t h  o n c o r h y n c h i a n d gonad o f f i s h  of female  the p r o g r e s s i v e  and l e n g t h  rate  o f same a g e .  oncorhynchi i n size  o f t h e gonad o f t h e f i s h .  was  dependent  Gravidity  85 of  the  female  to  the  r i p e n e s s of e g g s o f t h e  ity  Philonema  of female  worms was  Visceral total in  168  fish  was  fish  was  dependent  irrespective  on  but  o f age  examined.  the  proportional  ( F i g u r e X I V ) . Random  i n male f i s h  age  tended  and  directly  were o b s e r v e d  of R i v e r s I n l e t  not  was  fish  observed  adhesions  v e r y much v a r i a b l e  fish  oncorhynchi  and  gravid-  ( F i g u r e XV).  i n o n l y seven The  sex  occurrence  the  of  cysts  of t h e f i s h .  t o show more c y s t s p e r  time  of  of c a p t u r e  of  It male  fish.  DISCUSSION The present and  study  Miyaguchi  fish  caught  sharp  i n c r e a s e i n gonad w e i g h t  i s i n g e n e r a l agreement w i t h t h e (1958) and  in April The  to  f o r male and  During which  gm. this  fish.  on  the  female also  on  on  (1961).  i n c r e a s e d from  9.45  gm.  proportional  maturation  ripening  t o 77.5  t o the  of f e m a l e  that  gonadal  pituitary  r i p e n e s s of t e s t e s  hormones b u t with  gm.  of  of  of P^ also  as t h e  are not  f o r female  fish.  growth  i n c r e a s e of  gonad  dependent  oncorhynchi  gravidity parallel. i s not  of This  on hormone o r hormones o f  of but  P, sugg-  dependent  some o t h e r mechanism o f r e l a t i o n  the  in April  d e p e n d s not o n l y on m a t u r i t y of t e s t e s  the m a t u r a t i o n  and  t o August  showed r a p i d  sharp  of I s h i d a  gonad w e i g h t  1=49  gm.  the  Conversely, maturation  some o t h e r unknown f a c t o r ( s ) and  findings  worms i s d i r e c t l y  of o v a r i e s of f i s h .  P_^ o n c o r h y n c h i  The  i n comparison  s h o r t p e r i o d , P_^ o n c o r h y n c h i  The  oncorhynchi ests  gonad w e i g h t  is directly  of  Godfrey  i s v e r y low  same y e a r . 65.4  of s o c k e y e i n  the  between  fully  86  Fig.  XIV  R e l a t i o n s h i p between t h e o v a r y w e i g h t and t h e p r o d u c t i o n o f l a r v a e i n P h i l o n e m a o n c o r h y n c h i f r o m s o c k e y e salmon of R i v e r s I n l e t .  4501  03X0  O  O CD  OX00  0  coo  OOCD  oa  00  OOCO CCO  000  ticab  oXBb  EGG  JL  JL  MOR  EMB  JL  NO  TAIL  DEVELOPMENTAL  TAIL STAGES  8?  Fig.  XV  R e l a t i o n s h i p between t h e t e s t i s w e i g h t o f f i s h and the p r o d u c t i o n of l a r v a e i n Philonema o n c o r h y n c h i from sockeye salmon. 0  r e p r e s e n t s t h e number o f worms.  TESTIS o  ~l  o  o  1  g  1  9  o  0  o  1  8  §  8  § 8  _  o  I  0  §  WEIGHT IN GRAMS o  I  1  I  o  f  §8§*  8 8 8  0  8° 9  < m I-  o  I ml  o  o  o  5|  gpo  8  88 pituitary tion in  and  gonad o f f i s h .  i n Philonema  oncorhynchi  S e c t i o n I I I of p r e s e n t 42 male and  mostly 1961  immature  and  immature of  the  fish  same y e a r  sockeye  i n the  when f i s h  oncorhynchi  fish  1961). that  body c a v i t y  i n the  would p r o b a b l y  The  comparable  the  Generally,  sockeye  c y c l e , of P_^ o n c o r h y n c h i cycle  Question: cycle  in fish  of  One  Can  grown a d u l t  in August.  These f i s h  than  U s u a l l y female  i n 42  fish  spawn u n t i l No  one  of f i s h  Larvae  fish  in  from  year's Philonema  of the A p r i l  i n October  year  Philonema  Young  from  adult  fall  female  sexual maturity.  form  catch  which would  be  October,  a f o u r t o s i x year r e p r o d u c t i v e shown t o be  adapted  host. delay their  fifth  oncorhynchi  year?  reproductive  Present studies  i n §^ s o c k e y e  w/veft e x a m i n e d  w o u l d have spawned d u r i n g September  e a r l y December o f t h e same y e a r gravid.  two  three f i s h .  P._ o n c o r h y n c h i  P^  though  i n one  w h i c h spawn i n t h e i r  show f u l l y  of A p r i l ,  stage  cavity  has  are  (Godfrey,  male and  o f C u l t u s L a k e was  the  catch  would have spawned i n t h e  of these  adult  t o worms c o l l e c t e d  f o u r year  fish  gonad w e i g h t .  also reach  abdominal  reach  produc-  been d i s c u s s e d  to the m a t u r i t y i n d i c e s  c a t c h p r o b a b l y would not  would  of l a r v a l  of the A p r i l  s w i m b l a d d e r would p r o b a b l y r e a c h  time  to  hormones has  c o n s i d e r i n g t h e v e r y low  were f o u n d the  the  on  t h e y were c a p t u r e d .  of the A p r i l  later,  female  according  m i g h t be  dependence  studies.  I s h i d a e t a l 1958, t  The  and  worms i n 5  when c a p t u r e d  their  worms w o u l d a l s o  salmon are  a t t h e same t i m e .  larger  in  to become  size  Body w e i g h t  and  89 and  gonad w e i g h t  reached  the s i m i l a r  reproductive cycle  o f 5g s o c k e y e sexual  are g r e a t e r than  maturity,,  c y c l e o f P_^ o n c o r h y n c h i  o f i t s host>  P. o n c o r h y n c h i  4g b u t have  This suggests  that the  f o l l o w s the r e p r o d u c t i v e  b u t i s n o t d e p e n d e n t on age o f worm,  are found  i n matured sockeye  ^2  of  a n c i  (Adult ^2  age) . Larvae bladder  o f Owikeno L a k e ,  observation  larvae  in fish  The is  probably  in  the lake. which  variable Lake.  i n t h e swim-  at t h i s  I t i s not p o s s i b l e t o e x p l a i n  stage.  do n o t s u c e e d  low i n f e c t i o n  rate  due t o d i f f e r e n t Ruggles  However, i t i s c l e a r  i n reaching reproducing  p e r h a p s due t o unknown e c o l o g i c a l  ters  are not found  o f s p a w n i n g s o c k e y e f r o m C u l t u s L a k e b u t a r e common i n  spawning f i s h this  o f Po_ o n c o r h y n c h i  or p h y s i o l o g i c a l i n sockeye  important  food  abundant z o o p l a n k t e r s  exception  of northern  vernalis.  These  young s o c k e y e So  abundance o f  catch sockeye  zooplank-  o f young s o c k e y e , i s basins  o f Owikeno  i n the l a k e , with the  e n d , a r e D i a p t o m u s s p e c i e s and C y c l o p s  two t y p e s  of copepods are important  food f o r  i n Owikeno L a k e . f a r i t i s known t h a t C y c l o p s host  of  bicuspidatus i sa  suitable  intermediate  infected  Diaptomus s p a t u l o c r e n a t u s , E u c y c l o p s  lacustris,  factors.  of the A p r i l  i n s e a s o n s and a l s o i n f o u r d i f f e r e n t  The most  stage„  f o r a g i n g zones of j u v e n i l e  (1966) s t a t e d t h a t  a r e t h e most  that a l l  oncorhynchi.  M e s o c y c l o p s e d a x and C y c l o p s  Philonema^ a g u b e r n a c u l u m .  The a u t h o r  Meyer agilis,  scutifer  infected  with  (1960) Epischura larvae of  M e s o c y c l o p s edax  90 with  l a r v a e of Philonema  very  low r a t e o f i n f e c t i o n  Diaptomus  oregonensis  Lake, B r i t i s h This  oncorhynchi  e x p e r i m e n t a l l y but had  i n comparison  of Cyclops  bicuspidatus.  and M e s o c y c l o p s edax a r e common i n B a b i n e  Columbia  and^lalso i n f e c t a b l e  by P^  oncorhynchi.  o b s e r v a t i o n l e a d s t o a c o n c l u s i o n t h a t s e v e r a l s p e c i e s of  c o p e p o d may Therefore vernalis  serve  as an i n t e r m e d i a t e  i t c a n be s u g g e s t e d a c t as i n t e r m e d i a t e  Lake, B r i t i s h  Columbia.  host  of Philonema  species.  t h a t D i a p t o m u s s p e c i e s and hosts f o r P^ o n c o r h y n c h i  Cyclops  i n Owikeno  91 SUMMARY AND CONCLUSIONS  1)  Philonema  oncorhynchi  The  g r o w t h and m a t u r a t i o n  the  last  i n F\_ o n c o r h y c h i  fish.  Larvae  freshwater. oncorhynchi in  i s synchronized  i s different  from  i s completed  of the f i s h with  the onset  are found 3)  reaches  The  i n t o the adhesions  i n t h e f i s h and  towards the e s t u a r i e s .  No  fish.  stages  of t h i s  o f P^ a g u b e r n a c u l u m  o f P^ o n c o r h y n c h i .  the a d u l t stage  rainbow t r o u t  the swimbladder  are the normal h o s t s  A l l developmental  t h e same a s t h o s e  i s correlated  a g u b e r n a c u l u m Simon and Simon, 1936.  Non-anadromous s a l m o n o i d s dracunculoid.  dracunculoids,  adhesions  of m a t u r i t y  i n the spawning  Philonema  spawn i n  p a t t e r n o f P.  a t t h e age o f 23 months.  the m i g r a t i o n of the f i s h  adhesions  the r i p e n i n g of the  i n 12 months.  The a p p e a r a n c e o f v i s c e r a l  seem t o d i s a p p e a r with  with  t h a t of the other  the m i g r a t i o n of the l a r v a e from  body c a v i t y  during  The p r o d u c t i o n o f  The l o n g d u r a t i o n o f d e v e l o p m e n t a l  2)  are  life.  are r a p i d  o f t h e worms a r e r e l e a s e d when t h e f i s h  which development  with  o f P^ o n c o r h y c h i  6 months o f t h e h o s t ' s  larvae  K u i t u n e n - E k b a u m , 1933.  P. a g u b e r n a c u l u m  i n 182 d a y s i n e x p e r i m e n t a l l y  and g r a v i d worms o c c u r  i n immature  infected  yearling  wild  trout. 4) from other  cycle  t h a t o f P^ o n c o r h y n c h i dracunculoids.  dependent of  The l i f e  but i s s i m i l a r  i s different  t o those  The c o n t i n u a t i o n o f t h e l i f e  on t h e d i s c h a r g e  spawning.  o f P^ a g u b e r n a c u l u m  o f t h e worm i n t o  of the cycle i s  a l a k e a t t h e time  92 5) ted  The d i f f e r e n c e s between t h e two f o r m s a r e a t t r i b u -  t o the f o l l o w i n g : a) L e n g t h o f r e p r o d u c t i v e  cycle,  b)  i n the host,  reproductive  isolation  c) e l e c t r o p h o r e t i c p a t t e r n s , a'ad d)  lastly,  6)  different  The h y p o t h e s i s ,  b i o l o g y of hosts. t h a t t h e worm i s d e p e n d e n t on  hormonal s t i m u l u s from the host uction, uteri  was t e s t e d e x p e r i m e n t a l l y .  period, larvae.  fish,  Treatment  gonadal  support  11 t o 47 d a y s a f t e r  of the'green'  3,  treatment.  fish  Two o t h e r  the hypothesis  lines  i n the  f r o m 11 p i t u i t a r y -  with  I n t h e same  of evidence  fish  contained  stilbestrol  d e v e l o p m e n t b u t i t h a d no a p p a r e n t  effect  inhibit-  on t h e worm.  were d i s c u s s e d t o  o f h o r m o n a l dependence  o f P.  oncorhynchi  i t s host. Firstly,  the  were p r e s e n t  o n l y 2 o f 55 f e m a l e worms f r o m u n t r e a t e d  7)  on  Larvae  o f 33 o f 45 f e m a l e worms r e c o v e r e d  injected  ed  f o r s y n c h r o n i z a t i o n of r e p r o d -  ripe  l a r v i g e r o u s female  sockeye salmon p o p u l a t i o n o f R i v e r s I n l e t ,  4 or 5 years  during the l a s t to the r a p i d  old.  fish  immature  gonadal  failed  trout.  Rapid  development  6 months o f t h e f i s h ' s  Secondly, green  worms were f o u n d  whether  o f t h e worm life  only i n  occurred  which i s p a r a l l e l e d  development. non-larvigerous  P. o n c o r h y n c h i  from the  t o become g r a v i d when t r a n s p l a n t e d  into  93 LITERATURE  CITED  Akhmerov, A. Kh. 1955. P a r a s i t e fauna of the f i s h e s of the r i v e r Kamchatka. I z v e s t . T i l c h o o k e a n s k . N. 1. I n s t . Rybn. K h o z . O k e a n o g r . , 43: 99-137. Alicata,  J . E . and K. Noda 1960. 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Changes i n p l a s m a 17h y d r o x y - c o r t i c o s t e r o i d s accompanying s e x u a l m a t u r a t i o n and s p a w n i n g o f t h e P a c i f i c salmon ( O n c o r h y n c h u s  95 t s h a w y t s c h a ) and r a i n b o w t r o u t (Salmo P r o c . N a t l . A c a d . S c i . , 45: 8 8 6 - 8 9 X Hisaw,  gaidneri),  F. L . 1963. P h y s i o l o g y o f r e p r o d u c t i o n . P r o c . 22nd. Ann. B i o l o g y C o l l o q . Oregon S t a t e U n i v e r s i t y P r e s s .  H o a r , W. S. 1965a. The e n d o c r i n e s y s t e m as a c h e m i c a l l i n k between t h e o r g a n i s m and i t s e n v i r o n m e n t . Trans. RoyaL, S o c . Canada. 4 t h s e r i e s v o l . I l l , s e c t . I I I . , 1965b. Comparative P h y s i o l o g y : Hormones and reproduction i n fishes. Ann. Rev. P h y s i o l . , 27: 1966. G e n e r a l and Comp. P h y s i o l o g y . I n c . , New J e r s e y , C h a p t . 23.  Prentice-Hall  I d l e r , D. R., A. P. R o n a l d , and P. S. S c h m i d t , 1959. B i o c h e m i c a l s t u d e s on s o c k e y e salmon d u r i n g s p a w n i n g m i g r a t i o n . V l l - S t e r o i d homones i n p l a s m a . Canada J . B i o c h e m . P h y s i o l . , 37: 1227-1238. I s h i d a , T. and K. M i y a g u c h i , 1 9 5 8 . On t h e m a t u r i t y o f P a c i f i c salmon i n o f f s h o r e , w i t h r e f e r e n c e t o t h e s e a s o n a l v a r i a t i o n o f gonad w e i g h t . B u l l . Hokkaido Regional F i s h . R e s . L a b . , 18: 11-22. I s h i d a , R., K, T a k a g i , and S. A r i t a 1961. C r i t e r i a f o r t h e d i f f e r e n t i a t i o n o f mature and immature f o r m s o f chum and s o c k e y e s a l m o n i n n o r t h e r n s e a s . Internat i o n a l North P a c i f i c F i s h . Comm. B u l l e t i n No. 5, 27-47. J o h n s o n , W. E . and C. G r o o t 1963. O b s e r v a t i o n s o f t h e m i g r a t i o n o f young s o c k e y e salmon t h r o u g h a l a r g e , complex l a k e system. J . F i s h . R e s . B d . C a n a d a , 2 0 ( 4 ) : 919-938. Kendall,  Ko,  W. C. 1921. P e r i t o n e a l membranes, o v a r i e s and o v i d u c t s o f s a l m o n o i d f i s h e s and t h e i r s i g n i f i c a n c e i n f i s h cultura.4 p r a c t i c e s . B u l l . B u r e a u F i s h . , U.S., 37: 183-208.  R. 1966. Some a s p e c t s o f t h e d e v e l o p m e n t of P h i l o n e m a (Nematodes.: Dracunculoidea) larvae i n Cyclops b i c u s p i d a t u s £laus. M.Sc. T h e s i s , D e p t . o f Z o o l o g y , U n i v e r s i t y of B r i t i s h Columbia.  Kuitjonen-Ekbaum, E . 1933. P h i l o n e m a o n c o r h y n c h i g. n o v . e t . nov. C o n t r i b . Canada B i o l . F i s h . , 8 ( 4 ) : 71-75.  sp.  , 1937. N o t e s on t h e b i o l o g y and p a r t i a l l i f e h i s t o r y of t h e d r a c u n c u l i d , P h i l o n e m a o n c o r h y n c h i K u i t u n e n Ekbaum. M a n u s c r i p t R e p o r t s of the B i o l . S t a t i o n No. 304. F . R. B. o f Canada.  96 Larka,  P.  and S. P. S i n g h 1963. Some o b s e r v a t i o n s on t h e i m p l a n t a t i o n o f S e t a r i a c e r v i (Rud. 1819 and S e t a r i a digitata in rabbits. Indian Veterinary Journal, 4 0 ( 6 ) : 333-336.  Linstow,  . 1874.  Cited  from Kuitunen-Ekbaum,  Mayr, E . 1940. Speciation 74: 249-278. Meyer, M.  phenomenon  1937.  in birds.  Am.  Nat.,  C. 1958. S t u d i e s on P h i l o n e m a a g u b e r n a c u l u m , a d r a c u n c u l o i d nematode i n f e c t i n g s a i m o n i d s . J . P a r a s i t o l . 4 4 ( S u p p l . ) , 42.  , 1960. N o t e s on P h i l o n e m a a g u b e r n a c u l u m a n d o t h e r r r e l a t e d d r a c u n c u l o i d i n f e c t i n g saimonids. 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F u r t h e r s t u d i e s on t h e r e a c t i o n s o f a d u l t b l u e b l a c k s a l m o n t o i n j e c t e d salmon and mammalian gonadotropins. P r o g . F i s h . C u l t . ,16,99-107.  Pickford,  G. E . 1953. A s t u d y o f t h e h y p o p h y s e c t o m i z e d male k i l l i f f i s h , Fundulus h e t e r o c l i t u s ( L i n n . ) . Bull. Bingham O c e a n o g r . L a b . 14.(2), 5-41.  , and J . W. A t z 1957. The P h y s i o l o g y o f t h e p i t u i t a r y g l a n d of f i s h e s . N. Y. Z o o l . S o c . N. Y. 613 pp. P l a t z e r , E . G. 1964. The l i f e h i s t o r y o f P h i l o n e m a o n c o r h y n c h i i n s o c k e y e s a l m o n f r o m C u l t u s L a k e and t h e morphomet r i c v a r i a t i o n of a d u l t nematoes. M.Sc. T h e s i s , D e p t . of Z o o l o g y , U n i v e r s i t y of B r i t i s h Columbia.  97 Platzer,  PiPPy>  E . G. a n d J . R. A d a m s ( i n p r e s s ) . 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S o c , 66: 343-356. Ricker,  W.  E. 1937a. P h y s i c a l and chemical c h a r a c t e r i s t i c s o f C u l t u s L a k e , B . C . J . B i o l . B d . C a n a d . , 3: 3 6 3 402.  , 1937b. The f o o d a n d t h e f o o d s u p p l y o f s o c k e y e s a l m o n ( 0 . n e r k a Valbaum) i n C u l t u s L a k e , B r i t i s h C o l u m b i a J . B i o l . B o a r d , C a n a d . , 3: 4 5 0 - 4 6 8 . R e i c h e n b a c h - K l i n e , K. 1 9 5 5 . I n p a r a s i t e s o f f r e s h w a t e r f i s h and b i o l o g i c a l b a s i s f o r t h e i r c o n t r o l , V o l . XLIX: 155 pub. N a t i o n . S c i . Found'n. 1962 W a s h i n g t o n . Robertson,  0. H. 1 9 6 1 a . H y p e r a d r e n o c o r t i c i s m i n s p a w n i n g m i g r a t i n g r a i n b o w t r o u t ( S a l m o g a i r d n e r i ) ; comp a r i s o n w i t h P a c i f i c Salmon ( O n c o r h y n c h u s ) . Gen. Comp. E n d o c r i n . , 1 ( 5 / 6 ) : 473-4~5¥;  Robertson,  0. H. a n d B . C. W e x l e r , 1 9 5 9 . H y p e r p l a s i a o f t h e a d r e n a l c o r t i c a l t i s s u e i n P a c i f i c salmon (genus Oncorhynchus) and rainbow t r o u t (Salmo g a i r d n e r i ) accompanying s e x u a l m a t u r a t i o n and spawning E n d o c r i n o l o g y , 65: 225-238.  , 1960. H i s t o l o g i c a l changes i n t h e organs and t i s s u e of m i g r a t i n g and spawning P a c i f i c salmon (genus O n c o r h y n c h u s ) . E n d o c r i n o l o g y , 66: 222-239. Robertson,  0. H. a n d M. A . K r u p p , C. P. F a v o u r , S. H a n e a n d S. F . T h o m a s 1 9 6 1 b . P h y s i o l o g i c a l c h a n g e s o c c u r i n g i n t h e b l o o d o f t h e P a c i f i c S a l m o n (Q^_ t s h a w y t s c h a ) accompanying s e x u a l m a t u r a t i o n and spawning. E n d o c r i n o l o g y 68: 733-746.  98 R o t h s c h i l d , M. a n d B. F o r d 1964a. B r e e d i n g of r a b b i t (S. c u t i c u l i ) c o n t r o l l e d by t h e r e p r o d u c t i v e of t h e h o s t . N a t u r e , 201: 103-104.  flea hormones  1964b. M a t u r a t i o n and e g g l a y i n g o f t h e r a b b i t f l e a i n d u c e d by t h e e x t e r n a l a p p l i c a t i o n o f h y d r o c o r t i s o n e N a t u r e , 203: 210-211. 1965a. R e p r o d u c t i o n hormones o f t h e h o s t c o n t r o l l i n g the s e x u a l c y c l e of the r a b b i t f l e a (S. c u n i f c u l i ) . P r o c . X I I I n t . C o n g r . E n t . L o n d o n , 1961" ( 1 9 6 5 ) , p. 802. R o t h s c h i l d , M. 1965b. The r a b b i t XXIV ( 9 3 ) : 162-168.  flea  and hormones. E n d e a v o u r ,  R u g g l e s , C. P. 1966. J u v e n i l e s o c k e y e s t u d i e s i n Owikeno L a k e , B r i t i s h C o l u m b i a . C a n a d . F i s h C u l t u r i s t , V o l . 36: 3-21. Rumyantsev, E . A. 1965. The o c c u r r e n c e o f P h i l o n e m a s i b i r i c a ( B a v e r , 1946) (Nematoda, D r a c u n c u l i d a e ) i n C o r e g o n u s albula i n Karelia. Zoologicheskaya Zhurnal, V o l . , 4 4 ( 7 ) : 1082-1083. S c h a r r e r , E . and B. S c h a r r e r 1963. N e u r o e n d o c r i n o l o g y . C o l u m b i a U.P., New Y o r k . , 289 p. S c h m i d t , P. J . , B. S. M i t c h e l l , M. S m i t h and H. T s u y u k i , 1 9 6 5 . P i t u i t a r y hormone*of t h e P a c i f i c s a l m o n . 1. Response o f gonads i n immature t r o u t (Salmo g a i r d n e r i ) t o e x t r a c t s o f p i t u i t a r y g l a n d s f r o m a d u l t s a l m o n . Gen. Comp. E n d o c r i n o l . , 5 ( 2 ) : 1 9 7 - 2 0 6 . Simon, J . R. and F . Simon 1936. P h i l o n e m a a g u b e r n a c u l u m s p e c i e s nov. ( D r a c u n c u l i d a e ) , a nematode f r o m t h e body c a v i t y o f f i s h e s . P a r a s i t o l o g y , 28: 440-442. Smedley, E . M. 1933. Nematode p a r a s i t e s f r o m C a n a d i a n m a r i n e and f r e s h w a t e r f i s h e s . C o n t r i b . C a n a d . B i o l , and F i s h . n . s . V o l . 8 a n d 14 pp. 169-179. Smyth,  J . D. 1962. I n t r o d u c t i o n t o a n i m a l p a r a s i t o l o g y . The E n g l i s h U n i v e r s i t i e s P r e s s L t d . London, E . C . I . C h a p t . 10.  Strelkov,  I . A. and Shulman 1960 . P a r a s i t e i n f e s t a t i o n o f f i s h i n t h e p r o j e c t e d l a k e — c o n s t r u c t i o n a r e a o f t h e Amur R i v e r B a s i n . 1 0 t h C o n f . P a r a s i t o l . P r o l . , USSR, V o l . 2: 418.  99  Stunkard,  H. W. 1959. Induced gametogenesis i n a monogenetic t r e m a t o d e , P o l y s t o m a s t e l k a i V i g u e r a s , 1955. J . P a r a s i t . 45: 389-394.  S p a s s k i i , A. A. and V. M. Rakova 1958. Nematode f a u n a o f P a c i f i c Ocean f i s h e s . Work f r o m t h e h e l m i n t h . L a b . A k a d . Nauk, USSR, 1945-1957. S p a s s k i i , A. A. and V. A. Roitman and V. G. Shageava 1961. H e l m i n t h f a u n a of f i s h o f P l o t n i k h o v a B a s i n of Kamchatka D i s t r i c t . Trudy Gelminth. Laborat., 2: 270-285. S z i d a t , L.  On  1956. U b e r den E n t w i c k l u n g s z y k l u s m i t progenetischen l a r v e n s t a d i e n ( C e r c a r i a e n ) von G e r a r c h e l l a g e n a r c h e l l a T r a v a s s o s 1928 ( T r e m a t o d a , H e m i u r i d a e ) und d i e M o g l i c h k e i t e i n e r hormonalen B e e i n f l u s s u n g der p a r a s i t e n d u r c h i h r e W i r t s t i e r e . Z e i t s c h r . Tropenmed u. P a r a s i t o l o g i e , Bd. 7, N2. 1958. V e r s u c h e i n e r A n a l y s e des p r o b l e m s d e r A n p a s s u n g von M e e r e s t i e r e n an das S u s s w a s s e r . f . H y d r o b i o l o g i e , Bd., 54: 1-2.  Achiv.  Victor,  D.  V i k , R.  1964. N o t e s on the l i f e h i s t o r y of P h i l o n e m a a g u b e r n a c u l u m Simon and Simon, 1936 (Nematoda). Canad. J . Z o o l . , 42: 511-512.  Von  A. 1958. Cerebrospinal nematodiasis. I I . Transp l a n t a t i o n of S e t a r i a d i g i t a t a i n t o e x p e r i m e n t a l h o s t s . I n d i a n V e t e r i n a r y J o u r n a l , 35: 224-226.  I h e r i n g , R. 1937. Prog. F i s h .  A method f o r i n d u c i n g f i s h C u l t . , 34: 15-16.  t o spawn.  W e s s e n b e r g , H. 1961. S t u d i e s on the l i f e c y c l e and morphog e n e s i s of O . Q p a l i n a . U n i v . of C a l i f . P u b l i c a t i o n s i n Z o o l . 61: 315-370. W i e r z b i c k i , K. 1960. P h i l o m e t r o s i s o f c r u c i a n C a r p . P a r a s i t o l o g i c a p o l o n i c a , 8: 181-194. W i l l e m o e s - S u m , 1871.  C i t e d from Kuitunen-Ekbaum,  Acta  1937.  Table  Date Exam'd Smolts, 1965  Smolts, 2 y r . 1965  May  '65  12  4  2200  20  0  0  June  '65  13  5  1850  20  0  0  Aug.  '65  15  7  450  1+  1  5  0  0  S e p t . '65  16  8  335  3+  1  1  5  0  0  Jan.  '66  20  12  1050  20+  5  3  2  10  0  0  Apr.  '66  23  15  1238  20+  24  26  2  10  7  +  June  '66  25  17  64  20+  12  37  5  4  ++  May '65  24  650  3+  1  4  2  7  0  0  3  2  0  Smolts,'63 k e p t i n sea May water i n Nanaimo  4 & 16  '65  35  28  7  2  4  5  1  P r e c o c i o u s O c t . '65 male '65 Nov. '65  29  21  769  45  9  9  30  3  5  4  +++  30  22  225  32  56  11 135  9  5  4  ++  Prespawn-  S e p t . '64  40  32  15  30  59  5  0  0  '64  41  33  5  37  162  10  1  0  '64  42  34  1  53  203  10  0  0  '65  43  35  29  137  10  0  0  ing Oct. Nov. Dec.  *  approximate estimate  1  o o  Table  I I . Measurements o f P h i l o n e m a Sockeye o f d i f f e r e n t ages Mean s i z e s  History and age of f i s h S m o l t , 1965 12 months  S m o l t , 1965 13 months  Date worms collected May 1965  June 1965  S m o l t , 1965 15 months  August 1965  S m o l t , 1964 16 months  Sept. 1964  S m o l t , 1965 20 months  Jan. 1966  Stages i n swimbladder 3rd 4th  of Philonema  4th  oncorhynchi of from C u l t u s Lake.  oncorhynchi  Stages i n body c a v i t y M  10 1.16 104.26 248.93 480.62  Features measured  F 1. 2. 3. 4. 5.  10 1.02 100.4 228.6 468.3 10 1 3.4 1.18 222.0 108.9 244.2 399.6 492.0 971.2 10 1.08 102.53 225.9 481.2  2 6.27 180.3 410.7 929.6  10 10 1.19 4.6 104.22 152.8 236.38 334.4 512.0 871.7  5 3 5.22 14.3 195.9 222.3 445.8 422.4 818.5 1082.2  2 50. 01 259. 3 516. 4 936. 0  No. m e a s u r e d Length (mm.) N e r v e R i n g (/u) Muse. O e s . (/u) Gland.Oes. (/u)  Table  I I . Continued  S m o l t , 1965 23 months  April 1966  20 1.79 124.4 306.9 544.7  S m o l t , 1965 25 months  June 1966  20 3.32 143.3 312.7 773.6  S m o l t , 1963 35 months  April 1965  Precocious male 29-30 months  Oct.Nov. 1965  male a d u l t 4 41-42" months  Oct.Nov. 1965  female a d u l t 4 41-42 months  Oct.Nov. 1965  0  2  5 8.78 213. 7 451. 7 652. 1  20 2 18 .74 17.0 283.0 234 .9 566 .0 521.7 1536 .1 1172.3  10 4 .1 149 .0 329.8 881 .6  10 15 9. 5 21 .7 256 .7 192. 5 422. 2 502 .4 992. 42 1521 .0  7 3 .43 129 .6 298 .1 823 .1  2 4. 4 151. 2 302. 5 823. 4  20 7 .3 179 .2 360 .0 947 .9  5 5 17 .83 26.52 238 .1 252.4 474 .9 531.8 1534 .3 1325.4  10 8 15 6. 49 19 .15 146.82 189. 7 260 .0 345.0 491 .6 669.5 365. 6 1022. 2 1424 .1 1648.3 20 28 .7 312 .1 576 .8 1654 .0  20 119.7 384.4 771.0 1758.4  20 20 30 .95 202.95 364.6 326 .1 617 .8 738.9 1677 .0 1737.9  103 Table  Sex  III. Infection  ofjitrout  with Philonema  D u r a t i o n o f No. Worms infection Recovered  agubernaculm  Philonema 3rd stage  4th stage M F  30  5  5  _  M  152  7  -  -  -  2  2  1  2  M  152  7  -  -  -  3  2  1  1  M  162  6  M  169  1  M  176  4  M  178  2  M  181  2  -  -  -  M  182  3  -  -  -  -  F  3  1  F  58  3  2  2  -  _  F  118  9  -  -  -  F  151  3  -  -  F  151  3  -  -  -  F  160  2  -  -  - _  F  163  1  -  -  -  1  -  -  -  F  166  1  -  -  -  -  -  -  1  F  168  5  1  -  -  2  1  -  -  F  169  1  1  -  -  F  173  7  2  -  -  -  F  178  1  -  -  -  -  Rate  of i n f e c t i o n  1  _  _ -  _  _  adult M F  M  4  _  sub-adult M F _  1  _  -  _  percent  1 _  2  65.6  _  _  1  _  1  3  -  -  -  _ 3 2  -  1  1  2  _ 1  -  -  2  1  -  -  i  _  _  -  w i t h mean l a r v a e  _  5  1  _  _  i  _  _  1  2  -  1  2.3  Table  Treatments  IV. E f f e c t o f 17-b E s t r a d i o l , Salmon P i t u i t a r y e x t r a c t s and S t i l b e s t r o l on P h i l o n e m a s p e c i e s i n immature two y e a r o l d Sockeye f r o m Shuswap L a k e . No.of fish used  No. o f Infected fish  No. o f live worms  No. o f d e a d and e n c y s t e d f e m a l e worms  Duration of experiment  Remarks  17-b Estradiol  20  Salmon Pituitary  20  10  5M  Stilbestrol mixed f o o d  20  12  IF  12  103  No e f f e c t on f e m a l e worms. Infantile gonads a n d no change was o b s e r v e d on f i s h ,  Control  30  8  3F  8  103  No e f f e c t on f e m a l e worms. Infantile gonads a n d no change was o b s e r v e d on f i s h .  I F & 2M  103  94  No e f f e c t on f e m a l e worms. Infantile gonads a n d no change was o b s e r v e d on f i s h , I n f a n t i l e g o n a d s and no change was o b s e r v e d on f i s h .  T a b l e V.  E f f e c t o f salmon p i t u i t a r y e x t r a c t on f e m a l e P h i l o n e m a o n c o r h y n c h i r e c o v e r e d from the treated f i s h . (Time i n d a y s ) Developmental Stages* 7-12 19-22 47 Total  Egg  4  0  0  4  Morula  7  5  0  12  Tailless  6  4  6  16  Tailed  0  10  8  18  17  19  14  50  Total  106 Table  VI(a). Effect  Sex  Duration  F F F F F F F F F F F F M M M  3 7 10 11 11 12 19 20 22 47 47 47 10 22 47  1928.0 1701.0 2495.0 1587.5 2041.5 1587.5 2041.5 2948.5 2041.5 1928.0 1474.5 2041.5 2495.0 2268.0 2154.0  VI(b).  Control  7 7 15 15 16 31 31 32 49 7 15 24 31 49  1928.0 1928.0 2722.0 1814.5 1361.0 1588.0 1996.0 1474.5 1724.0 1474.5 2041.5 1928.0 1905.0 2948.5  Table F F F F F F F F F M M M M M  o f Salmon P i t u i t a r y  Body wt. (gm.)  Gonad wt. (gm.)  extracts  Mat. Index  82.0 62.0 132.5 129.0 115.5 97.0 120.0 339.0 157.0 159.0 106.0 137.0 59.0 10.0 47.0  4.25 3.64 5.3 8.12 5.66 6.1 5.88 11.5 7.7 8.24 7.2 6.7 2.36 0.44 2.18  76.0 83.0 119.0 93.0 60.5 83.0 165.0 94.0 157.5 21.0 31.0 35.0 15.0 37.0  3.94 4.3 4.33 5.12 4.44 5.22 8.26 6.37 9.13 1.4 1.5 1.8 0.79 1.25  on S o c k e y e  Nuptial  Color — —  ,  + +++ ++  ++ ++  ++ +++ +++ +++ + ++ ++  +++  — _ — — — _ —  ± _ _ —  —  T a b l e V I I . E f f e c t o f s t i l b e s t r o l on f e m a l e Philonema o n c o r h y n c h i r e c o v e r e d from the t r e a t e d f i s h . Developmental stages  Egg  (Time i n d a y s ) 1-9 11-18 20-23  Total  Fish  12  2  1  15  M  4  1  0  5  F  10  11  8  29  M  4  7  2  13  F  0  3  1  4  M  0  3  1  4  F  -  -  -  -  M  -  -  F  Morula  Embryo  Tailless  M Tailed -  -  F  22  16  10  48  M  8  11  3  22  F  Total  108 Table  V I I I . E f f e c t of implanted  stilbestrol  on  of prespawning sockeye Days  s  F i s h Weight (gms.)  ad Weight (gms.)  Mat.  Index  T  3  1474  5.5  .37  I  6  2268  9.5  .42  B  9  1906  5.44  .285  E  11  1361  2.8  .208  S  11  1928  6.5  .34  T  17  1814  8.0  .44  R  18  1814  9.2  .507  0  20  1474  2.0  .135  L  23  2268  6.5  .28  C  7  1474  21.0  1.4  0  15  2041  31.0  1.5  N  16  2041  23.5  1.15  T  24  1928  35.0  1.8  R  32  1769  22.5  1.27  0  37  907  20.5  2.27  L  37  1701  26,5  1.56  49  2948  37.0  1.25  49  2041  83.5  4.09  49  1928  50.5  2.62  testes  Table  IX. T r a n s p l a n t a t i o n o f a d u l t male and f e m a l e oncorhynchi  Sex  Length (cm.)  Mat. Index  Days Elapsed  F F M M M F M F F M M M F F M M F F M M M F F F F M  27.0 28.5 27.4 28.0 27.5 29.0 25.2 30.2 26.5 29.0 28.5 29.5 30.6 29.5 27.8 26.5 29.5 30.5 30.5 31.5 28.5 30.0 30.3 30.5 29.4 28.5  .061 .003 .191 .005 .247 .118 .01 .838 .047 .117 .173 .092 1.25 .87 .305 .19 .017 .03 .214 .205 .004 .03 .03 .023 .029 .02  2 3 9 10 12 20 24 32 32 37 48 57 68 70 77 80 80 81 83 85 85 85 85 86 86 86  i n body c a v i t y  Worms Transplanted 5 4 4 4 4 4 4 3 3 3 4 4 4 3 3 3 3 3 2 3 3 4 3 2 2 3  F F F F F F F F F F F F F F F F F F F F F F F F F F  & & & & & & & & & & & & & & & & & & & & & & & & & &  5 3 4 4 4 4 4 4 4 3 4 4 4 4 4 3 3 3 2 3 3 4 3 3 2 3  M M M M M M M M M M M M M M M M M M M M M M M M M M  Philonema  of t r o u t .  Worms Recovered (alive) 5 F & 5 M 1 M 2 F & 2 M —  3 M — —  2 M 1 M 2 M — — — —  I n t e n s i t y of Encysted adhesions worms __ — — •—  — •—  —  __ __  — —  1 F  .—  — — — — —  —  -__ — — —  —  + + +  — — — —  -  1 M — —  + + ++ —  +  — —  —  1 M  +  2 M  +  —  1 F — •—  2 2 2 2 2 1  F F F F F F —  2 F  Table M M M F M F F F F M F  32.0 31.2 31.0 28.0 29.5 29.8 29.5 31.4 33.0 30.0 30.8  IX.  Continued  .01 .007 .002 .052 .235 .057 .045 .024 .023 .228 .226  94 98 99 99 100 101 101 102 106 110 114  2 2 2 2 2 3 3 2 3 3 3  F F F F F F F F F F F  & & & & & & & & & & &  2 2 2 2 2 3 3 2 3 3 3  M M M M M M M M M M M  2 M —  2 M 4 M  —-  _«. —  +  1 M 3 M  + ++  2 M  +  —  2 F &  2 M  1  F  ++  «---==  2 F 1 F 2 F 1 F 1 F 1 F 2 F  —  I-  1  o  T a b l e X. Summary o f r e c o v e r y o f t r a n s p l a n t e d l a r v a e Mean S i z e o f Worms P h i l o n e m a i n "B.C. No. o f Larvae i n f i s h with 4th swimbladder F a d h e s i o n s days 3rd 4th stage M Sockeye, 1.19 4.6 C u l t u s Lake * (10) Control I (10)  1.64 Sockeye C u l t u s Lake Control II (25)  2.43  Steelhead Trout  1.35 <4)  5.42  Rainbow Trout  1.67 (6)  6.81  Sockey Shuswap Lake  1.25 (10) 1.37 (2)  * Number  measured  5.22 (5)  (7) (6)  (10)  4.07 (1)  14.3 (3)  50.01  17.0  (20)  (2)  20.62 (2)  7.72 (12)  15.46 (6)  8.38 (3)  age o f fish  No. o f f i s h used  perce: i n f ec  0  237  20' mo.  10  100  7  311  23  10  100  0  238240  2  2  223252  2  10  100  0  55  2  8  100  207  2  1  100  (2)  18.74  7.0 (1)  from salmonid f i s h .  56.46 (5)  mo.  5  80'  Table XI. T r a n s p l a n t a t i o n  of a d u l t  and f o u r t h s t a g e  larvae  i n t o body c a v i t y o f F r o g s Number of f r o g s Group A  Group B  I  Sex of f r o g s M  Duration (days)  No. o f worms transplanted  13  2 F & 2 M  1 live M 2 F in 1 dead M 1 c y s t  +  Recovery  Cyst  Adhesions  2  M  16  2 F & 2 M  2 live M 1 dead F  1 F  +  3  M  18  2 F & 2 M  2 dead M  2 F  ++  4  F  18  2 F & 2 M  2 dead M 2 dead F  +  5  F  18  2 F & 2 M  2 dead M a l l i n 2 do ad F 1 c y s t  +  6  M  11  7  M  11  8  M  13  9  F  13  10  F  18  0  0  7 dead  0  0  3 live 2 dead  0  0  tf  5 live  0  0  ft  5 live 2 dead  0  0  4 dead 10 f o u r t h stage larvae ft »»  T a b l e X I I . Summary o f m e r i s t i c of Age of  4.  Time fish caught  Sex of fish  August 1965  M  April 1965  M F  August 1965  M F  October 1965  M F  August 1965  M F  different'age  Area of catch  Average length of f i s h (cm.)  measurements o f  c o m p o s i t i o n of R i v e r s Average body weight (gm.)  Average gonad weight (gm.)  fish Inlet.  Mean M.I.  No. exam'd  No. infected  3.63  20  17  .14 .81  7 14  7 13  35.2  555.6  L a t .48'02 Long.39'56  44.4 48.0  1002.8 1170.0  O u t l e t of Rivers In.  52.9 51.2  2113.7 1810.7  65.4 77.5  3.03 4.13  12 15  10 15  50.4 52.4  1502.7 1793.9  73.4 327.3  5.06 18.33  37 38  36 36  61.5 60.3  3203.3 3005.4  91.0 139.3  2.89 4.51  6 19  6 18  Owikeno Lake  Owikeno Lake Outlet of Rivers In.  20.6 1.49 9.45  T a b l e X I I I . Summary o f o c c u r r e n c e in  of d i f f e r e n t  Philonema Av. No. Av. No. of F. o f M.  of R i v e r s  A v . No. o f larvae i n swimbladder  Inlet.  Time fish caught  3  August 1965  M  20  0  1.5  .6  April 1965  M F  7 14  1 1  1.2 .63  1.1 1.08  5.57 5.1  14 49  2.0 3.5  August 1965  M F  12 15  4 0  18.0 17.8  7.0 7.8  .75 , .13  15 4  1.25 .26  October 1965  M F  37 38  0 0  14.6 16.13  10.8 9.89  August 1965  M F  6 19  1 0  9.3 15.4  5.0 6.3  No. examM  No. w i t h visceral adhesions  age c o m p o s i t i o n  oncorhynchi  Age of fish  2  Sex of Fish  fish  of Philonema  12.9  7.0 2.1 0 .3  Cysts Total _% 15  .75  124 73  3.34 1.92  23 45  3.82 2.36  T a b l e XIV. Measurements o f P h i l o n e m a  oncorhynchi from R i v e r s  Measurements o f f e m a l e P h i l o n e m a Sex o f fish  Age of fish  M  M  M  M  Time  Mean length mm.  Mean Nervering /u  Mean t o t a l Oe s. mm.  August 1965  81.8 (3)  314.5 (3)  1.99 (3)  Inlet  Sockeye  M e a s u r e m e n t s o f male Mean length mm. 23.65 (3)  Philonema  Mean N e r v e r i n g Mean t o t ; Oes. mm /u 312.6 (3)  2.12 (3)  April 1965  5.5* (3)  April 1965  40.74 (3)  _  August 1965  76.77 (38)  280.6 (40)  2.08 (39)  19.23 (27)  265.8 (27)  1.96 (27)  August 1965  59.02 (60)  260.9 (68)  1.97 (67)  19.94 (44)  255.0 (44)  2.01 (41)  October 1965  111.09 (112)  326.5 (126)  2.37 (123)  21.85 (117)  289.51 (117)  2.15 (116)  October 1965  140.59 (134)  346.6 (137)  2.39 (136)  22.68 (110)  297.7 (108)  2.17 (108)  August 1965  91.0 (19)  313.5 (24)  2.25 (23)  21.81 (16)  302.4 (16)  2.182 (15)  August 1965  95.64 (78)  292.5 (83)  2.17 (80)  22.46 (45)  306.7 (44)  2.25 (42)  9.5* (3) _  20.65 (2)  * 4th stage en  T a b l e XV.  R e s u l t s o f Regression o f S i z e o f Worm on age o f f i s h and time  Simple r e g r e s s i o n : Standard error b  Standard error Y Male worms Age Time  13.24 12.27  5.23 4.28  24.84 4.86  50.79 34.76  df  F - value calculated tabulated  statistical significance  -.61 3.03  3.63 .65  130  .02 21.18  3.84 3.0  NS **  1.83 1.94  .92 .20  173  4.1 90.5  3.84 3.0  * **  -10.31 25.22  7.34 1.54  233  1.97 267.45  3.84 3.0  NS **  -0.73 17.72  11.76 1.6  142  .003 116.29  3.84 3.0  NS **  fish  13.01 4.16  Female worms female Age Time  Y = A + bx Y = s i z e o f worms  Male f i s h  Male worms female Age Time  caught  fish  144.80 -116.28  Female worms Male f i s h 46.68 94.43 Age -62.77 Time 34.55  NS * **  -  not s i g n i f i c a n t .05 l e v e l .01 l e v e l  

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