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The autonomic control of the teleost heart Jampolsky, Michael 1951

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L c 5 #7  (Gap - /  THE AUTONOMIC CONTROL of the TELEOST HEART by M i c h a e l Jampolsky  A T h e s i s Submitted i n P a r t i a l F u l f i l m e n t o f the Requirements f o r the Degree o f MASTER OF ARTS In the Department of ZOOLOGY  The U n i v e r s i t y o f B r i t i s h Columbia A p r i l 1951  ABSTRACT  In a d u l t g o l d f i s h (Qarassius the h e r r i n g  (Clupea p a l l a s i i ) ,  salmon (Oncorhynchus n e r k a ) ,  and  auratus),  a l e v i n s of the  the h e a r t  The  findings  are  i n v e s t i g a t i o n s on the a d u l t t e l e o s t  but c o n t r a r y to the c o n d i t i o n d e s c r i b e d {Fundulus,.  sockeye  i s i n h i b i t e d by acetyl -  c h o l i n e but shows no response to a d r e n a l i n . i n agreement w i t h p r e v i o u s  embryos of  f o r the  embryo  I t i s concluded t h a t the t e l e o s t heart l a c k s a  sympathetic i n n e r v a t i o n , and  that the m y o c a r d i a l c e l l s  unresponsive to a d r e n a l i n a t a l l ages.  are  ACKNOWLEDGEMENTS  I would l i k e t o take t h i s o p p o r t u n i t y t o express  my  a p p r e c i a t i o n t o Dr. W. A. Clemens, Head o f the Department of Zoology, f o r h i s kindness of  d u r i n g my s t u d i e s a t the U n i v e r s i t y  B r i t i s h Columbia. I wish t o thank Dr. W. S. Hoar f o r h e l p i n s u g g e s t i o n  of  the t o p i c , f o r a s s i s t a n c e and encouragement and f o r the  v a l u a b l e advice he has o f f e r e d throughout. I am indebted t o Donald Outram and Robert McMynn f o r advice on technique, o f f e r t i l i z i n g h e r r i n g eggs.  THE AUTONOMIC  CONTROL  of the TELEOST HEART INTRODUCTION B o t a z z i (1901), Armstrong  (1951), and B r i n l e y  (1932),  showed t h a t the h e a r t o f the t e l e o s t f i s h can be a r r e s t e d by either d i r e c t or r e f l e x vagal stimulation. little  Thus there i s  doubt as t o v a g a l supply and i n h i b i t i o n by a c e t y l -  choline.  On the o t h e r hand, a sympathetic i n n e r v a t i o n i s n o t  generally recognized.  Young (1931) p o i n t s out that S t a n n i u s  i n 1849 and C h e v r e l i n 1887 doing h i s t o l o g i c a l s t u d i e s were unable to f i n d connections between the sympathetic and the heart.  Young (1931) working on Aranoscopus  scaber was unable  to f i n d any h i s t o l o g i c a l evidence f o r sympathetic f i b r e s t o the h e a r t but d i d f i n d the c a r d i a c branch o f the vagus.  He  t r a c e d the l a t t e r from i t s o r i g i n a t s e v e r a l p o i n t s along the v i s c e r a l branch o f the vagus, running v e n t r a l to the gut i n the l o o s e t i s s u e j u s t a n t e r i o r to the ductus C u v i e r i , penet r a t i n g t h e ductus and f i n a l l y r e a c h i n g the s i n u s where i t s p l i t s up i n t o s e v e r a l s m a l l e r branches, most o f which  con-  verge on the s i n o a u r i c u l a r opening, around which t h e r e i s a complicated plexus c o n t a i n i n g many nerve c e l l s .  Young, i n  s t a t i n g t h a t the c a r d i a c nerves c o n s i s t of about 20 mediums i z e d parasympathetic medullated f i b r e s , suggests t h a t p e r haps t h e r e may be some sympathetic f i b r e s running a l o n g w i t h these parasympathetic f i b r e s .  These he says may have j o i n e d  2. the vagus v i a the ramus communicans.  He then throws doubt  on  t h i s t h e o r y when he says t h a t the ramus communicans to the vagus c o n s i s t s o n l y of non-medullated  f i b r e s , and s i n c e the  c a r d i a c nerve c o n t a i n s o n l y medullated f i b r e s , i t i s u n l i k e l y that any of them are from the sympathetic.  He a l s o mentions  that t h i s cannot be a b s o l u t e proof s i n c e i t i s w e l l known that p o s t - g a n g l i o n i c f i b r e s may n i n g f o r some d i s t a n c e .  a c q u i r e a sheath a f t e r r u n -  Thus a f t e r t h i s most c a r e f u l study  there i s no c l e a r anatomic  demonstration of a sympathetic  nerve s u p p l y to the h e a r t of the  teleost.  B r i n l e y (1935) working w i t h Fundulus  heteroclitus  a t t a c k e d the problem p h a r m a c o l o g i c a l l y . He found t h a t  injec-  t i o n s of a d r e n a l i n , 1:1000 to 1:50,000 i n t o embryos from 4-7  days o l d produced no change i n the r a t e of rhythm of  the h e a r t ; however, w i t h o l d e r embryos ( 8 - 1 3  days  Inclusive)  he found t h a t immediately a f t e r i n j e c t i o n the h e a r t stopped f o r s e v e r a l seconds.  F o l l o w i n g t h i s a r r e s t , he found  the h e a r t r a t e q u i c k l y i n c r e a s e d 10-20% above the rate.  that  initial  He concluded from t h i s t h a t the sympathetic must  become f u n c t i o n a l on the 8th day.  He a l s o d e s c r i b e s another  method whereby the embryos are immersed i n a s o l u t i o n of the drug.  The r e s u l t s obtained from both methods were the same. B r i n l e y goes on to d e s c r i b e a method of t r a n s -  p l a n t i n g the embryonic  h e a r t i n t o a host embryo.  The  of the h e a r t were severed i n making the t r a n s p l a n t .  nerves In  about 90% of the cases the drug showed no e f f e c t on the  5.  t r a n s p l a n t e d h e a r t but i n the remaining 10%  the h e a r t r a t e  was  g r e a t l y a c c e l e r a t e d d u p l i c a t i n g the r e s u l t s o b t a i n e d w i t h the innervated heart. B r i n l e y ' s c o n c l u s i o n s obtained from a p h a r m a c o l o g i c a l study are c o n t r a d i c t o r y to those obtained by e a r l i e r workers doing h i s t o l o g i c a l s t u d i e s . With t h i s i n mind a complete p h a r m a c o l o g i c a l was  undertaken  i n an attempt  t o e l u c i d a t e the problem.  study  4. MATERIAL AND METHODS G o l d f i s h ( C a r a s s i u s auratus) were o b t a i n e d from the G o l d f i s h Supply Company, S t o u f f v i l l e , v a r i e d from 5 t o 10 cms. the  Ontario.  Their lengths  They were kept i n a g l a s s  temperature of the water being t h e r m o s t a t i c a l l y  between E0°C. and  aquarium; controlled  E1°C.  A d u l t h e r r i n g (Clupea p a l l a s i i ) were procured from the  Pacific  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  of Canada and t r a n s p o r t e d to the u n i v e r s i t y i n 100 g a l l o n cans of sea water.  Eggs were s t r i p p e d from the females and spread  in layers ( 1 - 4  eggs t h i c k ) on the bottom o f Syracuse watch  g l a s s e s ; approximately 15 eggs were placed i n each.of watch g l a s s e s .  The watch g l a s s e s were placed i n a l a r g e tub  and covered w i t h 2 inches of sea water. cut tub.  100  The male f i s h were  open, the t e s t e s removed and the sperm squeezed i n t o the A f t e r 7 minutes the watch g l a s s e s were removed to a  b a t t e r y j a r c o n t a i n i n g f r e s h c o n t i n u o u s l y a e r a t e d sea water. "Tae b a t t e r y j a r was  kept i n a water bath at temperature  v a r y i n g between 8 and 8.5°C. and 80% s u c c e s s f u l . v a r i e d between  F e r t i l i z a t i o n was  between 70  The diameter of the f e r t i l i z e d eggs  and 1 mm.  Eggs were used f o r v a r y i n g p e r i o d s  up to 20 days f o l l o w i n g f e r t i l i z a t i o n . The salmon used were f r y of the sockey (Oncorhynchus  nerka).  salmon  These were o b t a i n e d from a stock main-  t a i n e d f o r experimental purposes by Grant Robertson. ranged from 1.5  to 2.5  and 8 weeks i n age.  cms.  Larvae  i n l e n g t h and were between 2 weeks  5. Two drugs were used: A c e t y l c h o l i n e (B.D.H. 0.1 gm. per ml.)  and a d r e n a l i n (Parke D a v i s 1:1000 s o l u t i o n ) .  Experiments on G o l d f i s h The r a t e of h e a r t b e a t of the g o l d f i s h was determined before and a f t e r a d m i n i s t e r i n g the drug.  F i s h were p i t h e d  w i t h a sharp probe i n s e r t e d i n t o the b r a i n a t a p o i n t  immedia-  t e l y p o s t e r i o r t o the cranium, and p l a c e d on a f r o g beard w i t h the  v e n t r a l s u r f a c e upwards.  An i n c i s i o n was made i n the  v e n t r a l w a l l extending about one cm. back from the c o r a c o i d r e g i o n o f the p e c t o r a l g i r d l e , the body w a l l was spread apart and secured t o the board by p a s s i n g two probes through i t . The exposed h e a r t c o u l d be seen b e a t i n g .  A stream of water  con-  t r o l l e d a t 21°C. (equal t o the a c c l i m a t i o n temperature o f tie g o l d f i s h ) was passed over the g i l l s and through the mouth o f the  fish.  The drug i n d i f f e r e n t c o n c e n t r a t i o n s was dropped on  the  exposed h e a r t from a hypodermic n e e d l e .  The number o f  beats were counted a t r e g u l a r i n t e r v a l s w i t h a stop watch. Four groups o f experiments comprised the f i r s t The f i r s t was a c o n t r o l group was  series.  (5 f i s h ) i n which the h e a r t r a t e  counted over p e r i o d s of between 50 and 95 minutes w i t h o u t  a d m i n i s t e r i n g any drugs. I n the second (4 f i s h ) , f o u r t h (3 f i s h ) ,  t h i r d (2 f i s h ) ,  c o n c e n t r a t i o n s of a d r e n a l i n 1:2,000, 1:24,000,  and 1:42,000 were t e s t e d . In the  a second s e r i e s of experiments the h e a r t beat o f  g o l d f i s h was recorded on a kymograph... L a r g e r , g o l d f i s h  about 10 cms. l o n g were chosen f o r kymograph r e c o r d s .  The  6. f i s h were secured fish firmly.  i n a v i c e which could be adjusted to h o l d  A stream of water t h e r m o s t a t i c a l l y c o n t r o l l e d  the a c c l i m a t i o n temperature was fish.  The h e a r t was  was  recorded  exposed and a f i n e p i n bent at one The  other end  end  was  of the  a t t a c h e d by a f i n e thread to a h e a r t l e v e r which the h e a r t c o n t r a c t i o n s on the smoked paper of a slow  speed kymograph drum. as b e f o r e .  Drugs were dropped on the exposed h e a r t  With the use of a stop watch, h e a r t r a t e  counted before and was  at  passed over the g i l l s of the  i n s e r t e d i n t o the t i p of the v e n t r i c l e . pin  the  administered  a f t e r a d m i n i s t e r i n g the drug.  was  Adrenalin  i n c o n c e n t r a t i o n s of 1:1000, 1:5,000, 1:7,500,  1:20,000, 1:50,000, 1:100,000, 1:250,000, 1:500,000 and c h o l i n e i n c o n c e n t r a t i o n s of 1:10,  1:50,  and  1:150.  In a t h i r d s e r i e s of experiments the h e a r t was away from i t s surrounding The h e a r t continued  acetyl-  cut  t i s s u e and removed to a g l a s s s l i d e .  b e a t i n g r h y t h m i c a l l y a f t e r i t s removal and  the e f f e c t of a c e t y l c h o l i n e 1:10 Experiments on H e r r i n g Larvae  was  determined as b e f o r e .  -  J u s t p r i o r to the experiment a Syracuse d i s h , cont a i n i n g f e r t i l i z e d h e r r i n g eggs i n which l a r v a e were d e v e l o p i n g was  removed from the b a t t e r y j a r , and p l a c e d i n a c l e a r  plastic  box  14 cms.  ends  by 18 cms.  with no  top and openings at both  a l l o w i n g f o r the f r e e passage of water through the box. maintained  This  the temperature of the eggs i n the Syracuse d i s h  between 8 and 8.5°C. Syracuse d i s h .  E x a c t l y 20 c c . of water was  No mixing  p l a c e d i n the  of the water i n the Syracuse d i s h  7. with  the water i n the p l a s t i c  box was allowed.  The p l a s t i c box  was p l a c e d on the p l a t f o r m of a b i n o c u l a r microscope. was s e l e c t e d f o r study.  One egg  The beat was timed f o r a p e r i o d and  then the drug was added to the 20 c c . of water making the s o l u t i o n up t o a d e f i n i t e c o n c e n t r a t i o n . was recorded  and the e f f e c t was noted.  Again the number o f beats The f i n a l  concentra-  t i o n s o f drugs used i n the watch g l a s s were a d r e n a l i n 1:1,000, 1:5,000, and 1:25,000; and a c e t y l c h o l i n e 1:10 and 1:30. EXPERIMENTS  on Salmon F r y -  A microscope was not r e q u i r e d s i n c e the heart was c l e a r l y v i s i b l e macroscopically. ments the salmon f r y was p l a c e d 20 c c . o f water. constant  As i n the p r e v i o u s  experi-  i n a Syracuse d i s h c o n t a i n i n g  The same t r a n s p a r e n t  g l a s s box was used as a  temperature water bath and the temperature o f the f r y  kept between 8.0 and 8.5°C.  The heart beat o f each f r y was  timed f o r a p e r i o d before adding the drug t o the 20 c c . of water and making the s o l u t i o n up t o the r e q u i r e d  concentration.  A f t e r a d d i t i o n o f the drug the number of beats was counted ai d the e f f e c t o f the drug noted.  The f i n a l c o n c e n t r a t i o n s  of  drugs used were a d r e n a l i n 1:1,000, 1:5,000, 1:7,500, 1:20,000, 1:50,000, 1:100,000 and a c e t y l c h o l i n e 1:1,000, 1:500, 1:330, 1:200, and 1:75. C o n t r o l t e s t s were r u n when i t was noted t h a t the pH of the a d r e n a l i n was 2.4 and t h a t of the a c e t y l c h o l i n e was 3.3 Two s o l u t i o n s were made up, one e q u i v a l e n t to a pH of 2.4, and the other 3.3, and d i l u t e d t o the same c o n c e n t r a t i o n s for  as use!  the drugs, namely 1:1,000, 1:500, 1:330, 1:200, 1:75 f o r  the a c e t y l c h o l i n e and  and 1:7,500, 1:20,000, 1:50,000, 1:33,000  1:100,000 f o r the a d r e n a l i n .  RESULTS  9.  G o l d f i s h H e a r t Rate The r e s u l t s are summarized i n f i g u r e $ i n d e t a i l i n the Appendices Tables 1,2,3 and 4. group showed a p a t t e r n  and  tabulated  The c o n t r o l  of gradual i n h i b i t i o n i n h e a r t r a t e .  The experimental groups when exposed t o c o n c e n t r a t i o n s of adrenalin  1:1,000, 1:24,000 and 1:42,000 showed the same p a t -  t e r n of g r a d u a l i n h i b i t i o n as was shown by the c o n t r o l group. The a d r e n a l i n  had no v i s i b l e e f f e c t on h e a r t  rate.  90  Figure  $ - G o l d f i s h h e a r t r a t e i n beats per minute. a. c o n t r o l group c. a d r e n a l i n 1:42,000 b. a d r e n a l i n 1:2,000 &* a d r e n a l i n 1:24,000 The arrows mark the p o i n t s where the a d r e n a l i n was administered. Each l i n e r e p r e s e n t s one f i s h .  G o l d f i s h Heart Kymograph Recordings -  10.  The r e s u l t s are summarized below and t a b u l a t e d i n d e t a i l i n the Appendices F i g u r e s 1,2,3,4,5 and 6. Concentration  Effects No change i n h e a r t r a t e or amplitude of contractions.  A d r e n a l i n 1:1,000 1:5,000  No change i n h e a r t r a t e or s l i g h t b i t i o n . No change i n amplitude.  "  1:7,500  One f i s h showed an i n c r e a s e i n h e a r t r a t e , the second one showed a decrease i n h e a r t r a t e . No change i n amplitude.  "  1:20,000  I n h i b i t i o n of h e a r t r a t e . amplitude.  "  1:50,000  No change i n h e a r t r a t e or amplitude.  "  1:100,000  No change i n h e a r t r a t e or amplitude.  "  1:250,000  No change i n h e a r t r a t e or amplitude.  "  1:500,000  No change i n h e a r t r a t e or amplitude.  11  A c e t y l c h o l i n e 1:10  Gradual decrease i n r a t e . decrease i n amplitude.  inhi-  No change i n  Gradual  "  1:50  Decrease  i n r a t e . No change i n amplitude.  "  1:150  Decrease  i n r a t e . No change i n amplitude.  Rate o f Beat o f the I s o l a t e d G o l d f i s h Heart The e f f e c t o f concentrated a c e t y l c h o l i n e was t e s t e d on the i s o l a t e d h e a r t o f two g o l d f i s h . In the f i r s t ,  the average r a t e f o r t e n d e t e r m i n a t i o n s  was 40 beats per minute,  the range 35 - 45.  The a d d i t i o n of  concentrated a c e t y l c h o l i n e caused immediate c e s s a t i o n of beat f o r about  t h i r t y seconds; f o l l o w i n g t h i s there was a g r a d u a l  recovery t o the normal  beat.  In the second, the average r a t e f o r t e n determina-  11. t i o n s was  50 beats per minute,  of c o n c e n t r a t e d a c e t y l c h o l i n e 20 beats per minute. minutes.  the range 40 - 60.  The  addition  caused an immediate i n h i b i t i o n t o  No recovery was  noted f o r a p e r i o d  of 10  12.  H e r r i n g Heart Rate 2r  The tabulated  3  r e s u l t s are summarized i n F i g u r e s & and •£ and  in detail  i n the Appendices F i g u r e s  5,6,7,8,9,10  and  11.  I /a  3*.  F i g u r e 2 - Heart r a t e of h e r r i n g eggs i n beats per minute. a. a d r e n a l i n 1:1,000 b. a d r e n a l i n 1:5,000 c. a d r e n a l i n 1:25,000 The arrow marks the p o i n t where the a d r e n a l i n was administered. Each l i n e r e p r e s e n t s one embryo.  13.  i JZO  i  SO  i  +O  i  3To  7~/A7£ //V  »  '  *0 A7//MTES 40  '  '  '  8o  So  s+o  1  sso  F i g u r e 3 - Heart r a t e of h e r r i n g eggs i n beats per minute„ a„ a c e t y l c h o l i n e 1:10 b a c e t y l c h o l i n e 1:100 c. a c e t y l c h o l i n e 1:300 The arrow marks the p o i n t where the a c e t y l c h o l i n e was administ e r e d o Each l i n e r e p r e s e n t s one embryo 0  0  14. F e r t i l i z e d h e r r i n g eggs when exposed t o c o n c e n t r a t i o n s of a d r e n a l i n heart  1:1,000, 1:5,000 and 1:25,000 showed no change o f  rate. When, however, the eggs were exposed t o c o n c e n t r a t i o n s  of a c e t y l c h o l i n e 1:10, 1:100 and 1:300 an i n h i b i t i o n i n h e a r t r a t e was noted.  The concentrated a c e t y l c h o l i n e  1:10 produced  a stoppage i n h e a r t r a t e .  1:100 a c e t y l c h o l i n e  showed an imme-  diate  by a g r a d u a l r e t u r n  t o normal r a t e  and  i n h i b i t i o n , followed  sometimes a c c e l e r a t i o n .  l e s s e r i n h i b i t i o n followed and  sometimes  1:300 a c e t y l c h o l i n e by a s i m i l a r r e t u r n  showed a  to normal r a t e  acceleration.  E f f e c t s of Adrenalin  and pH on Salmon F r y Oncorhynchus nerka -  The r e s u l t s are summarized below and t a b u l a t e d i n d e t a i l i n the Appendices Tables 12,13,14,15 and 16. Concentration Adrenalin  1:7,500  Inhibition  "  1:20,000  "  1:33,000  "  1:50,000  No e f f e c t  "  1:100,000  Inhibition  pH 2.4  1:7,500  "  1:20,000  "  1:33,000  Acceleration  "  1:50,000  Inhibition  "  1:100,000 Adrenalin  and  Effect  "  i n a range of c o n c e n t r a t i o n  between 1:7,500  1:100,000 when administered t o salmon f r y (Oncorhynchus  nerka)produced an i n h i b i t i o n of h e a r t s o l u t i o n o f the same pH as a d r e n a l i n the  same i n h i b i t i o n of heart  rate.  When, however., a  (pH 2.4) was a d m i n i s t e r e d ,  r a t e was n o t e d .  I t appears then  t h a t the i n h i b i t i o n was due t o the pH and n o t due t o the adrenalin. E f f e c t s of Acetylcholine The  results  and pH on Salmon F r y Oncorhynchus nerka  are summarized below and t a b u l a t e d i n  d e t a i l i n the Appendices Tables 17 and 18. Concentration  Effect  Acetylcholine  1:75  "  1:20  "  1:33  "  1:50  "  1:100  S l i g h t a c c e l e r a t i o n followed gradual i n h i b i t i o n .  1:75  F l u c t u a t i o n s , no p a t t e r n , no e f f e c t  pH 3.3  Marked  inhibition  Gradual  inhibition  tt  1:20  tt  tt  by  it  1:33  Acceleration  1:50  Slight  1:100  F l u c t u a t i o n s , no p a t t e r n ,  it  inhibition no e f f e c t  A c e t y l c h o l i n e when administered i n c o n c e n t r a t i o n s between 1:7.5 and 1:100 showed i n h i b i t i o n .  A s o l u t i o n of  pH 3.3 which i s the same pH as the a c e t y l c h o l i n e caused t u a t i o n s w i t h o u t any d e f i n i t e  pattern.  I t i s concluded then  t h a t the a c e t y l c h o l i n e causes i n h i b i t i o n of heart salmon f r y .  fluc-  r a t e o f the  16  F i g u r e ^ - Kymograph r e c o r d i n g G o l d f i s h Heart.  of the c a r d i a c  c y c l e of the  K B  F i g u r e £ - One c a r d i a c c y c l e o f the G o l d f i s h Heart enlarged from F i g u r e *f. AB - a u r i c u l a r s y s t o l e BC - f i r s t p a r t o f a u r i c u l a r d i a s t o l e which i s completed at F CDE - v e n t r i c u l a r s y s t o l e EF - f i r s t p a r t of v e n t r i c u l a r d i a s t o l e which i s completed at G During s y s t o l e the h e a r t i s c o n t r a c t i n g  and i s repas -  sented on the kymograph by a l i n e going up (AB and CD) a u r i c u l a r and v e n t r i c u l a r s y s t o l e  respectively.  During d i a s t o l e the h e a r t i s r e l a x i n g and i s r e p r e sented on the kymograph by a l i n e going down (BC and EF) auri c u l a r and v e n t r i c u l a r d i a s t o l e  respectively.  1 7 . D i s c u s s i o n  -  The v e n o s u s , c o n u s f o r m  a  b u t a  h e a r t  s i n g l e t h e  c l u d e d  t h a t  ( 1 8 9 3 ) , t h e  n e r v e s  w a s  c o m p l e t e l y  r e m o v e d  r h y t h m  a d r e n a l i n o f  a c t i o n  a  o f  t h e  i s  c o m p o s e d  s i n g l e t h e  o f  a  v e n t r i c l e .  v e n t r a l  1 9 4 8 ) .  f r o m  s u b s t a n c e  C a n n o n  c a l l e d  a d r e n a l i n  a n d  s y m p a t h e t i c a n  t h e  s i n u s T h e r e  a o r t a  i s  i s  n o  d i l a t e d  t o  t h e  a b s e n c e  t h e  f o u n d  m o d i f i e d  b y  i n  a n d  a f t e r  n e r v o u s  b e a t  c o n b e f o r e  t h e  h e a r t  s y s t e m  r h y t h m i c a l l y .  o n  s t u d y t h e  d e p e n d s  h e a r t  a s  a c e t y l c h o l i n e  s t i m u l a t i o n  o f  t h e  s t i m u l a -  h a v i n g  t h e  ( W i n t o n  T o d a y ,  t h e  b e l i e f  t h a t  s u b s t a n c e  s y m p a t h i n m a y  be  f r o m i s  o f  n e r v e s .  v a g u s  t h e  r a t e  b e a t  i n h i b i t o r b e a t . c e l l s  t o o f  s y s t e m  T h e r e m a y  e n d i n g s  r e l a t e d  w i t h  s y s t e m  o f  t h e  s y m p a t h e t i c  c l o s e l y  i d e n t i c a l  a c c e l e r a t o r  i s  i s  i t .  t h e  i s  h e a r t , h e a r t  t h e r e  r e s p o n d  t o  t h e s e  t o t h e n  a d r e n a l i n  a n d  f u r t h e r  r e -  w h i c h  c h e m i c a l l y  a c e t y l c h o l i n e t h e  a n d  a c e t y l c h o l i n e ;  f i b r e s  S h o u l d  t h e  s a m e  p a r a s y m p a t h e t i c  a c c e p t e d  r e l e a s e d  T h i s  u p o n  g e n e r a l l y  p a r a s y m p a t h e t i c  a n d  n e r v e s .  t h i s  a c t i o n  t h e  t o  f u n c t i o n e d  t h a t  c e n t r a l  T r u t o  n o w  m y o c a r d i a l o f  a s  t e l e o s t  i n v a r i a b l y  c o n t i n u e d  same  t h e  i s  p a r a s y m p a t h e t i c r a t e  i t  o n  a u t h o r  f r o m  n e r v e s ,  i n c r e a s e d  d e c r e a s e d  h e a r t  i n v o l v e d  t h e o r y  t h e  s t u d i e s  The  be  h e a r t  B a y l i s s , l e a s e d  m a y  s y m p a t h e t i c o n  a f t e r  b o d y  h a v i n g  T h i s  t h a t  e n d  i t .  m e t h o d  n e r v e s .  a  a n d  s e p a r a t e d  t h e  T h e  c a u s e  t e l e o s t  e m b r y o n i c  r e a c h e d  f r o m  i n h e r e n t  a  a t r i u m  p r o x i m a l  t h e .  t h e  t h e  b u l b u s . H i s  t i o n  o f  i f  b e w i l l  t h e r e  w i l l  i s  c a u s e  p o s s i b i l i t y d r u g s  i n  t h e  a  18. B r i n l e y (1935), i n h i s work on the t e l e o s t embryo, showed a d r e n a l i n t o have an e x c i t a t o r y a c t i o n .  He  p o s t u l a t e d the presence of a sympathetic system. was  S i m i l a r work  c a r r i e d out by the author on sockeye salmon f r y and  embryos but no i n c r e a s e i n r a t e was was  therefore  administered.  herring  obtained when a d r e n a l i n  C o n c e n t r a t i o n s o f a d r e n a l i n 1:1,000, 1:5,000  and 1:25,000 when a d m i n i s t e r e d t o the h e r r i n g eggs caused no n o t i c e a b l e change i n heart r a t e .  In the salmon  experiments  a wider range of c o n c e n t r a t i o n s were used, 1:7,500, 1:20,000, 1:33,000, 1:50,000 and 1:100,000. were i n h i b i t i o n .  When, however, a s o l u t i o n of the same pH as  the a d r e n a l i n (pH 2.4) was of h e a r t r a t e was noted. t i o n was  The t y p i c a l e f f e c t s here  a d m i n i s t e r e d the same i n h i b i t i o n I t appears then, t h a t the i n h i b i -  due to the pH and not due t o the a d r e n a l i n .  i n l i n e w i t h the h i s t o l o g i c a l evidence (Young, t r a r y t o the r e s u l t s of B r i n l e y A c e t y l c h o l i n e was the salmon f r y . attained.  1931)  t e s t e d on the h e r r i n g embryos and  In both cases t y p i c a l v a g a l e f f e c t s were and 1:30D)  depressed, the g r e a t e s t slowing was obtained  w i t h the h i g h e s t c o n c e n t r a t i o n .  F o r the salmon  a wider range of c o n c e n t r a t i o n s was 1:330, 1:500  but con-  (1935).  In a l l t h r e e c o n c e n t r a t i o n s (1:10, 1:100  the r a t e was  This i s  and 1:1,000.  experiments  used namely, 1:75,  In a l l cases the beat was  1:200,  depressed.  A s o l u t i o n of pH 3.3 which i s the same pH as the a c e t y l c h o l i h e caused f l u c t u a t i o n s without any d e f i n i t e p a t t e r n .  The con-  c l u s i o n i s then t h a t the a c e t y l c h o l i n e causes i n h i b i t i o n of h e a r t r a t e of the salmon f r y .  T h i s i s i n l i n e w i t h the  19. f i n d i n g s of workers already  r e f e r r e d to i n the  introduction.  S i m i l a r c o n c l u s i o n s were drawn from experiments the g o l d f i s h . and  Adrenalin  i n concentrations  on  of 1:1,000, 1:24,000  1:42,000 showed no e x c i t a t o r y e f f e c t on the h e a r t r a t e  the g o l d f i s h .  Kymograph r e c o r d s  show that a d r e n a l i n had  e f f e c t e i t h e r on the r a t e or amplitude of c o n t r a c t i o n s Kymograph r e c o r d i n g s a c e t y l c h o l i n e was  Appendices F i g u r e s 1,2,3  administered  A decrease i n both r a t e and noted w i t h the s t r o n g e s t ings Appendices F i g u r e s  and  When noted.  amplitude of g o l d f i s h heart  concentration 5 and  no  (see  4).  a decrease i n r a t e was  of  was  (see Kymograph r e c o r d -  6).  Hogben and Hobson (1924), working w i t h c e r t a i n i n v e r t e b r a t e s which l a c k a sympathetic i n n e r v a t i o n to the heart  found t h a t a d r e n a l i n caused an a c c e l e r a t i o n i n beat.  H i a t t and G-arrey (1942) showed t h a t a d r e n a l i n has augmentor a c t i o n on the c o n t r a c t i o n s  a slight  of spontaneously  beating  s t r i p s of t u r t l e v e n t r i c l e which they b e l i e v e to be without autonomic i n n e r v a t i o n .  The  s i t e of a c t i o n of a d r e n a l i n must  be e i t h e r on the vagus endings or d i r e c t l y on the m y o c a r d i a l cells.  The  f a c t t h a t a d r e n a l i n e f f e c t s , both augmentor  i n h i b i t o r y , are u n a f f e c t e d  by a t r o p i n e would appear to  and indi-  cate t h a t the a c t i o n i s not on vagus endings, t h e r e f o r e  site  of a c t i o n of a d r e n a l i n must be  The  on the m y o c a r d i a l c e l l .  e f f e c t of a d r e n a l i n on the myocardium of the  elasmobranch  a u r i c l e ( H i a t t 1942), which i s supposed to l a c k a sympathetic innervation, i s greater  than the  e f f e c t on the  turtle  ventri-  c l e which a l s o l a c k s a sympathetic i n n e r v a t i o n ; t h e r e f o r e , i t  20.  appears that the myocardial c e l l s of the elasmobranch  auricle  have a special s e n s i t i v i t y to adrenalin.  Thus i t would seem  in some invertebrates and i n elasmobranch  and some r e p t i l e s ,  the c o n t r a c t i l e elements are responsive to adrenalin even though there are no nerves.  This does not seem to be true f o r  the teleost myocardial c e l l s . A review of the l i t e r a t u r e on the nervous control of the heart i n various phyla of animals does not show any phylogenetic pattern. Clark, 1896)  In many of the crustaceans (Conant and  the heart i s controlled by both accelerator and  i n h i b i t o r nerves.  Brinley (1935) points out that Carlson in  1905 and 1906, working on arthropods and molluscs, found the heart of arthropods to be controlled by both accelerator and i n h i b i t o r , and the mollusc heart controlled by only inhibitcr fibres.  The annelids (Rogers, 1938) have both accelerator  and i n h i b i t o r nerves.  On the basis of work done by Botazzi  (1902), and Lutz (1930) on the elasmobranch,  i t is concluded  that there i s no sympathetic inns rvation to the heart although there i s a parasympathetic i n h i b i t o r system. Among the amphibians and r e p t i l e s (Rogers, 1938) i n h i b i t o r y and accelerator f i b r e s pass to the heart.  In the mammals (Winton and B a y l i s s ,  1948) there i s both sympathetic and parasympathetic control of the heart.  I t would seem that both vertebrate and inverte-  brate groups have members which possess both excitatory and i n h i b i t o r y nerves as well as those in which one or the other i s lacking.  Among vertebrates the teleosts are unique i n that  they lack both excitatory f i b r e s and a s e n s i t i v i t y to adrenalin.  CONCLUSIONS The  h e a r t s of Embryo h e r r i n g ( 3 - 1 4 days sifter  fertilization),  sockeye a l e v i n s ( 2 - 8 weeks a f t e r  fertiliza  t i o n ) and a d u l t g o l d f i s h are i n h i b i t e d by a c e t y l c h o l i n e but u n a f f e c t e d by a d r e n a l i n .  The t e l e o s t - h e a r t apparently  sympathetic i n n e r v a t i o n and the myocardial c e l l s ponsive to a d r e n a l i n .  lacks  are u n r e s -  •?M.  &3 bac^Ts:  *  j . /OOP  /ID.  /At-  Appending© Figure 1. Kymograph Recordings of the goldfish heart Carasgius Jforatus before and a f t e r administering adrenalin i n concentration of: a. 1:1000 b. 1:^000 c. l:fj0U0  d. 1:5000  5  s  ^  •  v Appendictee F i g u r e 2 . Kymograph r e c o r d i n g s o f t h e g o l d f i s h h e a r t C a r a s s i u s S t r a t u s b e f o r e and a f t e r a d m i n i s t e r i n g a d r e n a l i n i n c o n c e n t r a t i o n o f 1:7500  1  Is 20,000 55 b e a t s p e r minute  \f\j\f %  -  £  1$ b e a t s p e r minute  j„, ....  AAAAAAAAAAAAAA  <^ /  £> (S. &** 73  '/•  c.  i  Appendijfee F i g u r e 3. Kymograph r e c o r d i n g s o f the g o l d f i s h h e a r t G a r a s s i u s a a r a t u s b e f o r e and a f t e r a d m i n i s t e r i n g a d r e n a l i n i n concentration of:  a . 1:20,000 b . 1:20,000 c . 1:50,000  L:500,000  *  d.  55 b e a t s p e r minute  55 b e a t s p e r minute  Apoendiate* F i g u r e U . Kymograph r e c o r d i n g s o f the g o l d f i s h h e a r t G a r a s s i u s ^ u r a t u s b e f o r e and a f t e r a d m i n i s t e r i n g a d r e n a l i n inconcentration of:  a . 1:100,000 b . 1:100,000 c . 1:250,000 d. 1:500,000  Apoendiitefl F i g u r e $ . Kymograph r e c o r d i n g s o f the g o l d f i s h h e a r t G a r a s s i u s a a r a t u g b e f o r e and a f t e r a d m i n i s t e r i n g a c e t y l c h o l i n e m c o n c e n t r a t i o n o f 1:10  a  1:^0 hh  beats  per  minute  beats  per  minute  32.beats  per  minute  32  1:50  A p p e n d ! * *  beats  per  Figure  6.  minute  Kymograph  recordings  of  S t r a t u s before and a f t e r i n concentration of: a.  1:50 b. 1:50 c. 1:150  the  g o l d f i s h  heart  administering  Carassius  a c e t y l c h o l i n e  28.  Appendiffc*  Time  i n  Table  No.of  minutes  per  0 5 10 • 15 20 25 30 35  53  UO  U5 50 55 60 65 70 75 80 85 90 95  66  6k 6h  62 61 60 58 53 51 U9 hi hi  UU  k2  U2  1  -  beats  minute.  Controls  -  heart  r a t e  auratus  i n  beats  per  between  50  and  No.of per  31 63 62 62 63 62  60 58 56 5U 5U 52 50 50 k9 hi  U6 lil*  U3 1*1  beats  minute  of  the  minute  95  minutes.  No.of  beats  per  36 ks  55 62 59 57 56 5U 5U 5U 51 50 U8 hi hi  minute  g o l d f i s h counted  No.of p e r  7U 56 5U 50 h9 hi h6  U5 U3 U2 U0  Carassius  over  beats minute  a  p e r i o d  No, p ee :r  50 82 7U 69 65 62 62 63 63 59 5U 50 U9 U8 U9  minute  AppendiJ|p» T a b l e 2 - H e a r t r a t e o f t h e g o l d f i s h i n b e a t s p e r m i n u t e c o u n t e d b e f o r e and a f t e r a d m i n i s t e r i n g a d r e n a l i n i n concentration of  1:2,000  Time i n minutes  No.of beats p e r minute  No.of beats p e r minute  No.of beats pe r m i n u t e  No.of beats p e r minute  0 5 10 15  29 U5 51* 50  52 58 57 56  32 1*2 1*6 1*0  75 71*  20 25 30 35 1*0  1*5  Adrenalin  Adrenalin  1*3  50  1*3  51  1*0 37 36  1*9 1*5 1*1i*o  50 55  39  65 70 75 80 85 90 95  38 1*0 1*2 1*2  60  hp 39  1*2 1*0  31*  Adrenalin 32  11  66 72  69 63 Adrenalin 62  60 58 53 53 51 50  1*6  1*3 1*1 1*0 1*0  &8  Appendixes T a b l e 3 - H e a r t r a t e o f t h e C a r a s s i u s a u r a t u s i n b e a t s pe minute c o u n t e d b e f o r e and a f t e r adrenalin i n concentration of  1:2U,000  Time i n minutes  No.of beats pe r m i n u t e  No.of beats p r minute  0 5  71 83  83 68  10  77  15 20 25 30 35  bP  U5 55 60 65 70 75  Adrenalin  76 76 73 72 70 68 67 63  Adrenalin  6U  60 60 55 55 56 5U 5L  U7 U2 35 31 3U 32 35 39  administering  Appendiflfe-Table k - H e a r t r a t e o f t h e g o l d f i s h i n b e a t s p e r minute counted before and a f t e r i n c o n c e n t r a t i o n of Time i n m i n u t e s  No.of beats p e r minute  0  105  5  101  10 15 20 25  95 90 80 75  30 35 U0  1*5  50 55 60 65  Adrenalin  67 60 56 1*8  I46  1*3  U3 U5  1*1*  1*2  ho  33 32 32  administering adrenalin  1:U2,000  No.of beats per minute  No.of beats pe r m i n u t e  35  73  Adrenalin  ho  56  Adrei  38 37  1*8  50  56 50 51 51  5U 56 65 68 69 68  1*9 1*7  63  33 32 32  1*1*  1*1  38 35 3k  61*  61*  6k 82  87 90 88 85 83 80 71 72 77 80 75 7h  78 80 80 75 68 61*  6k  1  3fe AppendiBfee- Table 5 - Controls - heart rate of herring eggs (CJLupea pallasii) in beats per minute counted over a period between 20 and 28 minutes. Time in minutes  No.of beats per minute  No.of beats pr minute  0 2 u 6 8  30 31 32 32 33  lil  12  3U 35 35 3U 3U 35 35 35 36  10 1U 16  18 20 22 2h  26 28  111  Ul uu uu uu ui  33  38  UO 38 37  J  No.of beats per minute 32 33 33 3U 35  UO UD UO 39 38 3U  3*. Appendi&ss T a b l e 6 - H e a r t r a t e o f h e r r i n g eggs ( C l u p e a p a l l a s i i ) and a f t e r  before  administering adrenalin i n concentration  of 1:1,000 Time  Beats  Beats  Beats  Beats  Beats  Beats  0 2  2k 2k 2k  3k 3k 3k  33 3k 35  30 30 30  32 32 32  2k 2k 2k  25 25 26 26  35 35 37 39  36 37 38 kO  30 30 30 30  32 33 3k 32  31  25  39  ko  3k  31  22 22  37 38  35 38  29  3U 3k  30  38 38  1*0  38  39 35  1*1 1*1  29 28 28  h  6 8  10 12 11*  16 18 20 22  2k  26 28 30 32  Adrenalin  Adrenalin  39  3k  UO  la  1*2 hh hS  1*8 SO  52 5U 56 58  29  29  UO  kk kk 1*6 Uo  36 38  Adrenalin  28  27  32  30  29 29 31* 31*  36  1*1  1*2  Adrenalin  1*3 1*3  38 UO 3k 3k 29  29 28 33  27  30 30 30 29 29  30 31  32 33 31* 31* 3k 35 35 36  U3  1*3  1*3  U6 k6 ia i*i  Adrenalin  Adrenalin  .  Appendix©* Table 7 - Heart rate of herring eggs (Clupea pallasii) i n beats per minute counted before and after administering adrenalin i n concentration of 1:5,000 Time i n minutes  No.of beats per minute  0 2 k 6 8 10 12  23 2k 26 28 29 29 28  lk 16 18 20 22 2k 26 28 30  28 22 20 21 23 2k 2k 2k 2k Adrenalin 2k 2k 2k 2k  32 3k 36 38  No.of beats per minute 29 29 30 31 31 31 32 Adrenalin 32 32 32 33 33 33 33 33 33 3k 3k  Appendidte*. T a b l e 8 - H e a r t r a t e o f h e r r i n g eggs ( C l u p e a p a l l a s i i ) i n b e a t s p e r minute b e f o r e and a f t e r adrenalin i n concentration of n s  0 2  U 6  8  10  12  11* 16 18  20 22 2k 26 28 30 32 3k  36 38  UO  U2  UU U6 U8 50 52  administering  1:25,000  No.of beats p e r minute  No.of beats per minute  No.of beats p e r minute  Nolof beats per minute  3U 35 3U 3U 35  U8 U8 U8 U950  U2 U2 U2 U2 U3  5U 5U 5U 52 52  52 52 52 5U. 5U  U7 U7 U5  UU  50 . 50 50 U8 U8  3U 3U  56 56  UU U6  50 50  38 38 39 37 37  56 56  U6 U6 U6 U5 U5  50 5o 50 50 51  U8 U8 U8 U8 U9 U8 U7 U6 U5 U5 U5  52 50 50 50 50  35 35 35 36 35 Adrenalin  38 39 39  UO  Adrenalin  UU  Adrenalin  Adrenalin  Appendix©*- T a b l e 9 - H e a r t r a t e o f h e r r i n g eggs ( C l u p e a p a l l a s i i ) b e a t s p e r minute c o u n t e d b e f o r e and a d m i n i s t e r i n g a c e t y l c h o l i n e 1:10  Time i n m i n u t e s  0  No. o f beats p e r minute  78 Acetylcholine  3  Heart stopped completely  after  AppendiKoe T a b l e 10 - H e a r t r a t e o f h e r r i n g eggs ( C l u p e a p a l l a s i i ) b e a t s p e r minute counted b e f o r e and  after  administering acetylcholine i n concentration of  1:100  Time i n N o . o f b e a t s m i n u t e s p e r minute  minutes  No.of beats p e r minute  Time i n N o . o f b e a t s m i n u t e s p e r minute  0 5 10  k3  0 3 6  37 37 37  0 3 8  20  k3  13  37  9  36  12  3k  16  30  15  21  U3 10  I4.3  10  Acetylcholine  lU  k3  23  U2  29 31  39 37 31 25 36 39  2k  3k  38 53 59 6k  69 Ik  79 8k  89  9k  10U  109  119  i  k0  k3 k$  U5 k9 k9  50 50 kl k6  U6  19 29  30 35 U0 U5 50 55 60 65  n  37 U2  10  Acetylcholine  13  kk  30 30 k3 kk  27 36 30 28 25  • 3U 3k 3k  Acetylcholine 35  33  Appendiafc* T a b l e 11 - H e a r t r a t e o f h e r r i n g eggs ( C l u p e a p a l l a s i i ) i n b e a t s p e r minute c o u n t e d b e f o r e and a f t e r administering acetylcholine i n concentration of  1:300  Time i n minutes  No. of beats pe r minute  Time i n minutes  No.of beats p e r minute  0 2 3 8  U2 U2 U2 U2  0 2  U8 U8  33 27 26 30  11  13 18 23 28  Acetylcholine  U 6 9  13  U9 U9  U8 k8  Acetylcholine  Iii  hh U3  18  UO 39  28  UO UI UI  15 17 19 22 23 27 29 32 35 37 39 U2  UU U7 57  U3  38 39 U3 U6 U8 53 52 53 55 55 55  37. Appendices T a b l e 12 a . H e a r t r a t e o f salmon f r y (Oncorhynchus n e r k a ) i n b e a t s p e r m i n u t e c o u n t e d b e f o r e and a f t e r administering adrenalin i n concentration of  1:100,000 b . H e a r t r a t e o f salmon f r y (Oncorhynchus n e r k a ) i n b e a t s p e r m i n u t e c o u n t e d b e f o r e and a f t e r a d m i n i s t e r i n g a c i d a t a PH o f 2.1*. i n  a.  a.  concentration of a. a. a.  a.  JDCCtl/D  .DBcl  1 2  6h  68 55 68 68 68 68  3  65  69 69  71 71  79  k  68  69 69  71 71  77  67 65  69 69 69 69  72 72 73 73  183  75  67 65  67  70 70 70  7k  76  I 81  1  63  62 70 Adrenalin  83  !  0  .  65 ...  5 6 7 8 9  5?  64  10  60  11 12  55 50  13 1U  U8  !6  17 18 ^ 20 %  22 23  18  69 59 69 69 70 70  1B l — W •78 65  78  a.  a.  6k  59 60  6U  60  65  62  55  i  68  60  67  62 6U  6k  55 60  Adrenalin  Adrenalin  a.  a.  Beats Beats Beats Beats Beats  68 65 62  Adrenalin '  1:100,000  63 58 59  67  60 65  67 67  60  60  6%  68  58 67  70  71 77 79  Adrenalin  Beats  6U 6k  67  73  77  67  71  77  67  73 71  7U 77  68 68  72 68  80  73 -  70 6U  82  65 1:100,000 59  Adrenalin  Adrenalin  Adrenalin  70  71  b.  70  82  68  6U  65  79  73 73  79  77 75  67  65 6k  77 79  55 50  73 71  83  75 75  65 65  77 79  61 59  Adrenalin  71  86  79 82 83  6k  Adrenalin  65 6U 62  61  58 58 58 l  9  <*>  60 60  6k 6k  68 70  Si  $0.  A p p e n d i x e s T a b l e 13 a . H e a r t r a t e o f salmon f r y (Ohcorhynchus n e r k a ) i n b e a t s p e r minute c o u n t e d b e f o r e and a f t e r administering adrenalin i n concentration of  1:50,000 b . H e a r t r a t e o f salmon f r y (Oncorhynchus n e r k a ) i n b e a t s p e r minute c o u n t e d b e f o r e and a f t e r a d m i n i s t e r i n g a c i d a t a PH o f 2.k i n concentration of  a.  1:50,000  a.  b.  b.  Time i n minutes  No.of beats p e r minute  No.of beats p e r minute  No.of beats p e r minute  No.of beats p e r minute  0 1 2  56 57 58  63 6k 65  68 68 69  57 58 56  3  u  5 6 7 8 9 10 11 12 13 Ik  15 16  60 59 59 67 6k  62 65 65  Adrenalin  65 68 68 67 67 68  62 60 67 6k  67 70 67  Adrenalin  67  65 67 68 68  A c i d PH 2.U  1:50,000 67 71 60 60 6k 65 66 68  A c i d PH 2.k  1:50,000 65 65 65 6k 60 60 59 56  67 65  55  o  Appendices- T a b l e l k . a . H e a r t r a t e of salmon f r y (Oncorhynchus n e r k a ) i n b e a t s p e r m i n u t e c o u n t e d b e f o r e and a f t e r administering adrenalin i n concentration of  1:33,000 b . H e a r t r a t e o f salmon f r y (Oncorhynchus n e r k a ) i n b e a t s p e r minute counted b e f o r e and a f t e r a d m i n i s t e r i n g a c i d a t a PH o f 2.k i n concentration of  1:33,000  Time i n minutes  a. No.of beats p e r minute  b. No.of beats p e r minute  b. No.of beats p e r minute  0 1 2 3  67 77 77 77  70 75 75 71  55 55 55 57  k  5 6 7 8 9 10 11 12 13 lk  15  Adrenalin  71  75 19  78 75 77 19 11  73  Adrenalin  67 65 71 65 71 70 70 67 67 65 60  \  51  A c i d PH 2.k  1:33,000 56 60 58 57 59 58 56 52 k6  N  A p p e n d i c e s Table 15 a . H e a r t r a t e o f salmon f r y (Oncorhynchus n e r k a ) i n b e a t s p e r minute c o u n t e d b e f o r e a n d  after  administering adrenalin i n concentration  of 1:20,000 b . H e a r t r a t e o f salmon f r y (Oncorhynchus n e r k a ) i n b e a t s p e r m i n u t e counted b e f o r e and a d m i n i s t e r i n g a c i d a t a PH o f 2.U c o n c e n t r a t i o n o f 1:20,000 Time i n m i n u t e s  0 1 2 3 k 5  6 7  8 9  10 11 12 13  Hi  a. No.of beats p e r minute  73 71 71 71  Adrenalin  67 65 65 6U 65 6k 65 6U 6h 60 58  b. No.of beats p e r minute  6U 65  A c i d PH 2.U  1:20,000 59 61 65 67 56 58  in  after  Appendices- Table 16 a .  H e a r t r a t e o f salmon f r y (Oncorhynchus n e r k a ) i n b e a t s p e r minute c o u n t e d b e f o r e and  after  administering adrenalin i n concentration of  1:7,500 b . H e a r t r a t e o f salmon f r y (Oncorhynchus n e r k a ) i n b e a t s p e r minute c o u n t e d b e f o r e and a d m i n i s t e r i n g a c i d a t a PH o f 2.k i n concentration of  1:7,500  Time i n minutes  a. No.of beats per minute  b. No.of beats p e r minute  0 1  71 73  68 67  2  75  3 k  5 6 7 8 9  10  Adrenalin  71 71 70 65 60 57 56 52  A c i d PH 2.k  1:7,500 65 67 60 58 55 53  after  AppendiJSe* Table 17  H e a r t r a t e o f salmon f r y (Oncorhynchus nerka) b e f o r e and a f t e r  administering acetylcholine  i n c o n c e n t r a t i o n of a . 1:100 b . 1:500 d.  c  . 1:330  1:200 e . 1:75  Time i n minutes  a. No.of beats p e r :minute  b. No.of heats p e r minute  c. No.of beats p e r minute  d. No.of beats p e r minute  e. No.of beats p e r minute  0 1 2  59 59 59  62 6U 60  70  60 60 61  67 75 75  3  61  6U  61  1:200 67  1:75 109  h  58 57 59 58  5 6 7 8 9 10 11 12 13 in  15 16  17 18  Acetylch<  1:1000 55 6h  68 62 71 6h  67 60 5U 57 68  65 59 58 55 58 59  60  Acetylcholine Acetylcholine  Acetylcholine Acetylcholine  1:500 75 68 61 58 56 52  :330 65 56 60  58 57 5H 56  5U  he  38  AppendiJ^* Table 18 - Heart rate of salmon f r y (Oncorhynchus nerka) i n beats per minute counted before and after administering a c i d PH 3.3 i n concentration of  a. 1:1000 b. 1:500 c. 1:330 d. 1:200 e. 1:75 Time i n minutes  a. b. c d. e. No.of beats No.of beats No.of beats No.of beats No.of beats per minute per minute per minute per minute per minute  0 1 2  62 67 60  3 U  5 6 7 8 9 10 11 12  13  Iii  15 16  17  18 19 20 21  75 <  75 58  Acid PH 3-3  1:1000 75 ,75 75 75 75 75 75 75 75 75 61 68 62 70  75 75  76 72 75  77 81  68 72 Acid PH 3.3  1:500 72 72  72  72  68 61 67 68 61 63 60  79  Acid PH  1:330 81  86  3.3  72 72 68  65  72  67  1:200 68  75  Acid PH 3.3  70  68  Acid PH 3.3  81  68  86  75 70 70 65 70 63 70 67  86 86  86 86 86 86  1:75 65 73 65 56 60 35 35  BIBLIOGRAPHY A r m s t r o n g , P . B . 1931  F u n c t i o n a l r e a c t i o n s i n t h e embryonic h e a r t , accompanying t h e i n g r o w t h and development o f the vagus i n n e r v a t i o n .  J o u r . E x p . Z o o l . , 58 :  43-61. B o t a z z i , F . 1901  C e n t l . F . P h y s i o l . , 1 4 (Quoted from B r i n l e y 1935).  B o t a z z i , F . 1902  Z e i t s c h r . F . B i o l . , 4 3 (Quoted from L u t z 1930).  B r i n l e y , F . J . 1932  A p h y s i o l o g i c a l study o f t h e i n n e r v a t i o n o f t h e h e a r t o f f i s h embryos.  •  P h y s i o . Z o o l . , 5 : 527-537. B r i n l e y , F . J . 1935-  Evidence f o r a sympathetic i n n e r v a t i o n o f the t e l e o s t hea,rtj w i t h a n o t e on a method o f t r a n s p l a n t i n g the h e a r t o f fundulus  embryos.  P h y s i o l . Z o o l . , 8 t 360-373. C o n a n t , F . S . and C l a r k , H . L ' . 1896  On t h e a c c e l e r a t o r and i n h i b i t o r n e r v e s o f t h e  H i a t t , E . P . and G a r r y , ¥ . E . 1942  Drug a c t i o n s on t h e s p o n t a n e o u s l y  crab's heart.  J o u r . E x p . M e d . , 1 t 341-347. beating  turtle ventricle indicating lack of innervation. Amer. J o u r . P h y s i o l . , 138 : 758-762.  H i a t t , E . P . 1942  The a c t i o n o f a d r e n a l i n a c e t y l c h o l i n e and potassium i n r e l a t i o n t o the i n n e r v a t i o n o f the i s o l a t e d a u r i c l e o f the s p i n y dogfish acanthias).  H i s , W.J. 1893  (Squalus  Amer. J o u r . P h y s i o l . , 139 t 45-48.  M a t h . - P h y s . C l a s s e , B d . 18 S . 1-64 (Quoted from A r m s t r o n g J o u r . E x p . Z o o l . 58 t 43-61).  Hogben, L.T. and Hobson, A.D. 1924  Studies on Internal secretion 111. - The action of pituitary extract and adrenalin on contractile tissues of certain invertebrata.  British Jour.  Exp. Biol., 1 : 4-87-500. Lutz, B.R. 1930  The effect of adrenalin on the auricle of elasmobranch fishes. Amer. Jour. Physiol., 94 : 135-139.  Rogers, C.G. 1938  Textbook of Comparative Physiology, McGraw H i l l Book Co. Inc.  Winton, F.R. and Bayliss, L.E. 194-8  Human Physiology, Blakiston.  Young, J.Z. 1931"  On the autonomic system of the teleost fish Aranoscopus scaber. Sci., 74 : 492-535.  Quart. Jour. Micro.  

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