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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 Michael Jampolsky A Thesis Submitted i n P a r t i a l Fulfilment of the Requirements f o r the Degree of MASTER OF ARTS In the Department of ZOOLOGY The University of B r i t i s h Columbia A p r i l 1951 ABSTRACT In adult g o l d f i s h (Qarassius auratus), embryos of the herring (Clupea p a l l a s i i ) , and alevins of the sockeye salmon (Oncorhynchus nerka), the heart i s inhibited by acetyl -choline but shows no response to adrenalin. The findings are in agreement with previous investigations on the adult teleost but contrary to the condition described f o r the embryo {Fundulus,. I t i s concluded that the t e l e o s t heart lacks a sympathetic innervation, and that the myocardial c e l l s are unresponsive to adrenalin at a l l ages. ACKNOWLEDGEMENTS I would l i k e to take t h i s opportunity to express my appreciation to Dr. W. A. Clemens, Head of the Department of Zoology, f o r h i s kindness during my studies at the University of B r i t i s h Columbia. I wish to thank Dr. W. S. Hoar f o r help i n suggestion of the topic, for assistance and encouragement and for the valuable advice he has offered throughout. I am indebted to Donald Outram and Robert McMynn f o r advice on technique, of f e r t i l i z i n g herring eggs. THE AUTONOMIC CONTROL of the TELEOST HEART INTRODUCTION Botazzi (1901), Armstrong (1951), and Brinley (1932), showed that the heart of the teleost f i s h can be arrested by either d i r e c t or r e f l e x vagal stimulation. Thus there i s l i t t l e doubt as to vagal supply and i n h i b i t i o n by a c e t y l -choline. On the other hand, a sympathetic innervation i s not generally recognized. Young (1931) points out that Stannius i n 1849 and Chevrel i n 1887 doing h i s t o l o g i c a l studies 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 for sympathetic f i b r e s to the heart but did f i n d the cardiac branch of the vagus. He traced the l a t t e r from i t s o r i g i n at several points along the v i s c e r a l branch of the vagus, running ventral to the gut i n the loose tissue just anterior to the ductus Cu v i e r i , pene-tr a t i n g the ductus and f i n a l l y reaching the sinus where i t s p l i t s up into several smaller branches, most of which con-verge on the sino auricular opening, around which there i s a complicated plexus containing many nerve c e l l s . Young, i n stating that the cardiac nerves consist of about 20 medium-sized parasympathetic medullated f i b r e s , suggests that per-haps there may be some sympathetic f i b r e s running along with these parasympathetic f i b r e s . These he says may have joined 2. the vagus v i a the ramus communicans. He then throws doubt on th i s theory when he says that the ramus communicans to the vagus consists only of non-medullated f i b r e s , and since the cardiac nerve contains only 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 also mentions that t h i s cannot be absolute proof since i t i s w e l l known that post-ganglionic f i b r e s may acquire a sheath a f t e r run-ning f o r some distance. Thus afte r t h i s most c a r e f u l study there i s no clear anatomic demonstration of a sympathetic nerve supply to the heart of the t e l e o s t . Brinley (1935) working with Fundulus he t e r o c l i t u s attacked the problem pharmacologically. He found that i n j e c -tions of adrenalin, 1:1000 to 1:50,000 into embryos from 4 - 7 days old produced no change i n the rate of rhythm of the heart; however, with older embryos ( 8 - 1 3 days Inclusive) he found that immediately after i n j e c t i o n the heart stopped for several seconds. Following t h i s arrest, he found that the heart rate quickly increased 10-20% above the i n i t i a l rate. He concluded from t h i s that the sympathetic must become functional on the 8th day. He also describes another method whereby the embryos are immersed i n a solution of the drug. The results obtained from both methods were the same. Brinley goes on to describe a method of trans-planting the embryonic heart into a host embryo. The nerves of the heart were severed i n making the transplant. In about 90% of the cases the drug showed no e f f e c t on the 5 . transplanted heart but i n the remaining 10% the heart rate was greatly accelerated duplicating the results obtained with the innervated heart. Brinley's conclusions obtained from a pharmacological study are contradictory 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 studies. With t h i s i n mind a complete pharmacological study was undertaken i n an attempt to elucidate the problem. 4. MATERIAL AND METHODS Goldfish (Carassius auratus) were obtained from the Goldfish Supply Company, S t o u f f v i l l e , Ontario. Their lengths varied from 5 to 10 cms. They were kept in a glass aquarium; the temperature of the water being thermostatically controlled between E0°C. and E1°C. Adult herring (Clupea p a l l a s i i ) were procured from the P a c i f i c B i o l o g i c a l Station of the Fisheries Research Board of Canada and transported to the unive r s i t y i n 100 gallon cans of sea water. Eggs were stripped from the females and spread in layers ( 1 - 4 eggs thick) on the bottom of Syracuse watch glasses; approximately 15 eggs were placed i n each.of 100 watch glasses. The watch glasses were placed i n a large tub and covered with 2 inches of sea water. The male f i s h were cut open, the testes removed and the sperm squeezed into the tub. A f t e r 7 minutes the watch glasses were removed to a battery jar containing fresh continuously aerated sea water. "Tae battery jar was kept in a water bath at temperature varying between 8 and 8.5°C. F e r t i l i z a t i o n was between 70 and 80% successful. The diameter of the f e r t i l i z e d eggs varied between and 1 mm. Eggs were used f o r varying periods up to 20 days following f e r t i l i z a t i o n . The salmon used were f r y of the sockey salmon (Oncorhynchus nerka). These were obtained from a stock main-tained f o r experimental purposes by Grant Robertson. Larvae ranged from 1.5 to 2.5 cms. i n length and were between 2 weeks and 8 weeks i n age. 5. Two drugs were used: Acetylcholine (B.D.H. 0.1 gm. per ml.) and adrenalin (Parke Davis 1:1000 s o l u t i o n ) . Experiments on Goldfish -The rate of heartbeat of the go l d f i s h was determined before and a f t e r administering the drug. F i s h were pithed with a sharp probe inserted into the brain at a point immedia-t e l y posterior to the cranium, and placed on a frog beard with the ventral surface upwards. An i n c i s i o n was made i n the ventral wall extending about one cm. back from the coracoid region of the pectoral g i r d l e , the body wall was spread apart and secured to the board by passing two probes through i t . The exposed heart could be seen beating. A stream of water con-t r o l l e d at 21°C. (equal to the acclimation temperature of tie goldfish) was passed over the g i l l s and through the mouth of the f i s h . The drug i n d i f f e r e n t concentrations was dropped on the exposed heart from a hypodermic needle. The number of beats were counted at regular intervals with a stop watch. Four groups of experiments comprised the f i r s t s e r i e s . The f i r s t was a control group (5 fish) i n which the heart rate was counted over periods of between 50 and 95 minutes without administering any drugs. In the second (4 f i s h ) , t h i r d (2 fish), fourth (3 f i s h ) , concentrations of adrenalin 1:2,000, 1:24,000, and 1:42,000 were tested. In a second series of experiments the heart beat of the g o l d f i s h was recorded on a kymograph... Larger,goldfish about 10 cms. long were chosen f o r kymograph records. The 6. f i s h were secured i n a vice which could be adjusted to hold the f i s h f i r m l y . A stream of water thermostatically controlled at the acclimation temperature was passed over the g i l l s of the f i s h . The heart was exposed and a fine pin bent at one end was inserted into the t i p of the v e n t r i c l e . The other end of the pin was attached by a fine thread to a heart lever which recorded the heart contractions on the smoked paper of a slow speed kymograph drum. Drugs were dropped on the exposed heart as before. With the use of a stop watch, heart rate was counted before and after administering the drug. Adrenalin was administered i n concentrations 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 a c e t y l -choline in concentrations of 1:10, 1:50, and 1:150. In a t h i r d series of experiments the heart was cut away from i t s surrounding tissue and removed to a glass s l i d e . The heart continued beating rhythmically a f t e r i t s removal and the e f f e c t of acetylcholine 1:10 was determined as before. Experiments on Herring Larvae -Just p r i o r to the experiment a Syracuse dish, con-taining f e r t i l i z e d herring eggs i n which larvae were developing was removed from the battery j a r , and placed i n a clear p l a s t i c box 14 cms. by 18 cms. with no top and openings at both ends allowing f o r the free passage of water through the box. This maintained the temperature of the eggs i n the Syracuse dish between 8 and 8.5°C. Exactly 20 cc. of water was placed i n the Syracuse dish. No mixing of the water i n the Syracuse dish 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 placed on the platform of a binocular microscope. One egg was selected for study. The beat was timed f o r a period and then the drug was added to the 20 cc. of water making the solu-t i o n up to a d e f i n i t e concentration. Again the number of beats was recorded and the eff e c t was noted. The f i n a l concentra-tions of drugs used i n the watch glass were adrenalin 1:1,000, 1:5,000, and 1:25,000; and acetylcholine 1:10 and 1:30. EXPERIMENTS on Salmon Fry -A microscope was not required since the heart was c l e a r l y v i s i b l e macroscopically. As in the previous experi-ments the salmon f r y was placed i n a Syracuse dish containing 20 cc. of water. The same transparent glass box was used as a constant temperature water bath and the temperature of the f r y kept between 8.0 and 8.5°C. The heart beat of each f r y was timed for a period before adding the drug to the 20 cc. of water and making the solution up to the required concentration. Afte r addition of the drug the number of beats was counted ai d the e f f e c t of the drug noted. The f i n a l concentrations of drugs used were adrenalin 1:1,000, 1:5,000, 1:7,500, 1:20,000, 1:50,000, 1:100,000 and acetylcholine 1:1,000, 1:500, 1:330, 1:200, and 1:75. Control tests were run when i t was noted that the pH of the adrenalin was 2.4 and that of the acetylcholine was 3.3 Two solutions were made up, one equivalent to a pH of 2.4, and the other 3.3, and diluted to the same concentrations as use! fo r the drugs, namely 1:1,000, 1:500, 1:330, 1:200, 1:75 f o r the acetylcholine and 1:7,500, 1:20,000, 1:50,000, 1:33,000 and 1:100,000 f o r the adrenalin. RESULTS 9. Goldfish Heart Rate -The results are summarized i n figure $ and tabulated in d e t a i l i n the Appendices Tables 1,2,3 and 4. The control group showed a pattern of gradual i n h i b i t i o n i n heart rate. The experimental groups when exposed to concentrations of adrenalin 1:1,000, 1:24,000 and 1:42,000 showed the same pat-tern of gradual i n h i b i t i o n as was shown by the control group. The adrenalin had no v i s i b l e e f f e c t on heart rate. 90 Figure $ - Goldfish heart rate i n beats per minute. a. control group c. adrenalin 1:42,000 b. adrenalin 1:2,000 &* adrenalin 1:24,000 The arrows mark the points where the adrenalin was adminis-tered. Each l i n e represents one f i s h . Goldfish Heart Kymograph Recordings - 10. The results are summarized below and tabulated i n d e t a i l i n the Appendices Figures 1,2,3,4,5 and 6. Concentration Adrenalin 1:1,000 11 1:5,000 " 1:7,500 " 1:20,000 " 1:50,000 " 1:100,000 " 1:250,000 " 1:500,000 Acetylcholine 1:10 " 1:50 " 1:150 Ef f e c t s No change i n heart rate or amplitude of contractions. No change i n heart rate or s l i g h t i n h i -b i t i o n . No change i n amplitude. One f i s h showed an increase i n heart rate, the second one showed a decrease i n heart rate. No change i n amplitude. I n h i b i t i o n of heart rate. No change i n amplitude. No change i n heart rate or amplitude. No change i n heart rate or amplitude. No change i n heart rate or amplitude. No change i n heart rate or amplitude. Gradual decrease i n rate. Gradual decrease i n amplitude. Decrease i n rate. No change i n amplitude. Decrease i n rate. No change i n amplitude. Rate of Beat of the Isolated Goldfish Heart -The e f f e c t of concentrated acetylcholine was tested on the is o l a t e d heart of two g o l d f i s h . In the f i r s t , the average rate f o r ten determinations was 40 beats per minute, the range 35 - 45. The addition of concentrated acetylcholine caused immediate cessation of beat for about t h i r t y seconds; following t h i s there was a gradual recovery to the normal beat. In the second, the average rate f o r ten determina-11. tions was 50 beats per minute, the range 40 - 60. The addition of concentrated acetylcholine caused an immediate i n h i b i t i o n to 20 beats per minute. No recovery was noted for a period of 10 minutes. 12. Herring Heart Rate -2r 3 The r e s u l t s are summarized in Figures & and •£ and tabulated in d e t a i l i n the Appendices Figures 5,6,7,8,9,10 and 11. I /a 3*. Figure 2 - Heart rate of herring eggs i n beats per minute. a. adrenalin 1:1,000 b. adrenalin 1:5,000 c. adrenalin 1:25,000 The arrow marks the point where the adrenalin was administered. Each l i n e represents one embryo. 13. i i i i » ' ' ' ' 1 JZO SO +O 3To 40 *0 8o So s+o sso 7~/A7£ //V A7//MTES Figure 3 - Heart rate of herring eggs i n beats per minute„ a„ acetylcholine 1:10 b 0 acetylcholine 1:100 c. acetylcholine 1:300 The arrow marks the point where the acetylcholine was adminis-tered o Each l i n e represents one embryo0 14. F e r t i l i z e d herring eggs when exposed to concentrations of adrenalin 1:1,000, 1:5,000 and 1:25,000 showed no change of heart rate. When, however, the eggs were exposed to concentrations of acetylcholine 1:10, 1:100 and 1:300 an i n h i b i t i o n i n heart rate was noted. The concentrated acetylcholine 1:10 produced a stoppage i n heart rate. 1:100 acetylcholine showed an imme-diate i n h i b i t i o n , followed by a gradual return to normal rate and sometimes acceleration. 1:300 acetylcholine showed a lesser i n h i b i t i o n followed by a s i m i l a r return to normal rate and sometimes acceleration. E f f e c t s of Adrenalin and pH on Salmon Fry Oncorhynchus nerka -The re s u l t s are summarized below and tabulated i n d e t a i l i n the Appendices Tables 12,13,14,15 and 16. Concentration Adrenalin 1:7,500 " 1:20,000 E f f e c t I n h i b i t i o n No e f f e c t I n h i b i t i o n " 1:33,000 " 1:50,000 " 1:100,000 pH 2.4 1:7,500 " 1:20,000 " 1:33,000 " 1:50,000 " 1:100,000 " Adrenalin i n a range of concentration between 1:7,500 and 1:100,000 when administered to salmon f r y (Oncorhynchus Acceleration I n h i b i t i o n nerka)produced an i n h i b i t i o n of heart rate. When, however., a solution of the same pH as adrenalin (pH 2.4) was administered, the same i n h i b i t i o n of heart rate was noted. I t appears then that the i n h i b i t i o n was due to the pH and not due to the adrenalin. E f f e c t s of Acetylcholine and pH on Salmon Fry Oncorhynchus nerka The results are summarized below and tabulated i n d e t a i l i n the Appendices Tables 17 and 18. Concentration Acetylcholine 1:75 " 1:20 " 1:33 " 1:50 " 1:100 pH 3.3 1:75 1:20 1:33 1:50 1:100 E f f e c t Marked i n h i b i t i o n Gradual i n h i b i t i o n S l i g h t acceleration followed by gradual i n h i b i t i o n . Fluctuations, no pattern, no ef f e c t tt tt tt it it Acceleration S l i g h t i n h i b i t i o n Fluctuations, no pattern, no e f f e c t Acetylcholine when administered i n concentrations between 1:7.5 and 1:100 showed i n h i b i t i o n . A solution of pH 3.3 which i s the same pH as the acetylcholine caused f l u c -tuations without any d e f i n i t e pattern. I t i s concluded then that the acetylcholine causes i n h i b i t i o n of heart rate of the salmon f r y . 16 Figure ^ - Kymograph recording of the cardiac cycle of the Goldfish Heart. K B Figure £ - One cardiac cycle of the Goldfish Heart enlarged from Figure *f. AB - auricular systole BC - f i r s t part of auricular diastole which i s completed at F CDE - ve n t r i c u l a r systole EF - f i r s t part of v e n t r i c u l a r diastole which i s com-pleted at G During systole the heart i s contracting and i s repas -sented on the kymograph by a l i n e going up (AB and CD) a u r i -cular and ven t r i c u l a r systole respectively. During diastole the heart i s relaxing and i s repre-sented on the kymograph by a l i n e going down (BC and EF) auri -cular and ventricular diastole respectively. 1 7 . D i s c u s s i o n -T h e h e a r t o f t h e t e l e o s t i s c o m p o s e d o f a s i n u s v e n o s u s , a s i n g l e a t r i u m a n d a s i n g l e v e n t r i c l e . T h e r e i s n o c o n u s b u t t h e p r o x i m a l e n d o f t h e v e n t r a l a o r t a i s d i l a t e d t o f o r m a b u l b u s . H i s ( 1 8 9 3 ) , a f t e r s t u d i e s o n t h e t e l e o s t T r u t o c o n -c l u d e d t h a t t h e e m b r y o n i c h e a r t i n v a r i a b l y f u n c t i o n e d b e f o r e t h e . n e r v e s r e a c h e d i t . T h e a u t h o r f o u n d t h a t a f t e r t h e h e a r t w a s c o m p l e t e l y s e p a r a t e d f r o m t h e c e n t r a l n e r v o u s s y s t e m a n d r e m o v e d f r o m t h e b o d y i t c o n t i n u e d t o b e a t r h y t h m i c a l l y . T h i s i n h e r e n t r h y t h m m a y b e m o d i f i e d b y n e r v e s . T h e m e t h o d i n v o l v e d i n t h i s s t u d y d e p e n d s u p o n a d r e n a l i n h a v i n g t h e s a m e a c t i o n o n t h e h e a r t a s t h e s t i m u l a -t i o n o f s y m p a t h e t i c n e r v e s , a n d a c e t y l c h o l i n e h a v i n g t h e s a m e a c t i o n o n t h e h e a r t a s t h e s t i m u l a t i o n o f p a r a s y m p a t h e t i c n e r v e s . T h i s t h e o r y i s n o w g e n e r a l l y a c c e p t e d ( W i n t o n a n d B a y l i s s , 1 9 4 8 ) . T o d a y , t h e b e l i e f i s t h a t t h e s u b s t a n c e r e -l e a s e d f r o m t h e p a r a s y m p a t h e t i c v a g u s e n d i n g s i s a c e t y l c h o l i n e ; t h e s u b s t a n c e r e l e a s e d f r o m t h e s y m p a t h e t i c f i b r e s w h i c h C a n n o n c a l l e d s y m p a t h i n i s c l o s e l y r e l a t e d c h e m i c a l l y t o a d r e n a l i n a n d m a y b e i d e n t i c a l w i t h i t . S h o u l d t h e r e t h e n b e a s y m p a t h e t i c a c c e l e r a t o r s y s t e m t o t h e h e a r t , a d r e n a l i n w i l l c a u s e a n i n c r e a s e d r a t e o f b e a t o f t h e h e a r t a n d i f t h e r e i s a p a r a s y m p a t h e t i c i n h i b i t o r s y s t e m a c e t y l c h o l i n e w i l l c a u s e a d e c r e a s e d r a t e o f b e a t . T h e r e i s t h e f u r t h e r p o s s i b i l i t y t h a t t h e m y o c a r d i a l c e l l s m a y r e s p o n d t o t h e s e d r u g s i n t h e a b s e n c e o f n e r v e s . 18. Brinley (1935), i n his work on the teleost embryo, showed adrenalin to have an excitatory action. He therefore postulated the presence of a sympathetic system. Similar work was carried out by the author on sockeye salmon f r y and herring embryos but no increase in rate was obtained when adrenalin was administered. Concentrations of adrenalin 1:1,000, 1:5,000 and 1:25,000 when administered to the herring eggs caused no noticeable change i n heart rate. In the salmon experiments a wider range of concentrations were used, 1:7,500, 1:20,000, 1:33,000, 1:50,000 and 1:100,000. The t y p i c a l effects here were i n h i b i t i o n . When, however, a solution of the same pH as the adrenalin (pH 2.4) was administered the same i n h i b i t i o n of heart rate was noted. I t appears then, that the i n h i b i -t i o n was due to the pH and not due to the adrenalin. This i s i n l i n e with the h i s t o l o g i c a l evidence (Young, 1931) but con-tra r y to the r e s u l t s of Brinley (1935). Acetylcholine was tested on the herring embryos and the salmon f r y . In both cases t y p i c a l vagal effects were attained. In a l l three concentrations (1:10, 1:100 and 1:30D) the rate was depressed, the greatest slowing was obtained with the highest concentration. For the salmon experiments a wider range of concentrations was used namely, 1:75, 1:200, 1:330, 1:500 and 1:1,000. In a l l cases the beat was depressed. A s o l u t i o n of pH 3.3 which i s the same pH as the acetylcholih e caused fluctuations without any d e f i n i t e pattern. The con-clusion i s then that the acetylcholine causes i n h i b i t i o n of heart rate of the salmon f r y . This i s i n l i n e with the 19. findings of workers already referred to i n the introduction. Similar conclusions were drawn from experiments on the g o l d f i s h . Adrenalin i n concentrations of 1:1,000, 1:24,000 and 1:42,000 showed no excitatory e f f e c t on the heart rate of the g o l d f i s h . Kymograph records show that adrenalin had no e f f e c t either on the rate or amplitude of contractions (see Kymograph recordings Appendices Figures 1,2,3 and 4). When acetylcholine was administered a decrease in rate was noted. A decrease i n both rate and amplitude of g o l d f i s h heart was noted with the strongest concentration (see Kymograph record-ings Appendices Figures 5 and 6). Hogben and Hobson (1924), working with certain invertebrates which lack a sympathetic innervation to the heart found that adrenalin caused an acceleration i n beat. H i a t t and G-arrey (1942) showed that adrenalin has a s l i g h t augmentor action on the contractions 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 believe to be without autonomic innervation. The s i t e of action of adrenalin must be either on the vagus endings or d i r e c t l y on the myocardial c e l l s . The f a c t that adrenalin e f f e c t s , both augmentor and i n h i b i t o r y , are unaffected by atropine would appear to i n d i -cate that the action i s not on vagus endings, therefore s i t e of action of adrenalin must be on the myocardial c e l l . The e f f e c t of adrenalin on the myocardium of the elasmobranch a u r i c l e (Hiatt 1942), which i s supposed to lack a sympathetic innervation, i s greater than the e f f e c t on the t u r t l e v e n t r i -cle which also lacks a sympathetic innervation; therefore, i t 2 0 . appears that the myocardial cells of the elasmobranch auricle have a special sensitivity to adrenalin. Thus i t would seem in some invertebrates and in elasmobranch and some reptiles, the contractile elements are responsive to adrenalin even though there are no nerves. This does not seem to be true for the teleost myocardial c e l l s . A review of the literature on the nervous control of the heart in various phyla of animals does not show any phylogenetic pattern. In many of the crustaceans (Conant and Clark, 1896) the heart is controlled by both accelerator and inhibitor 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 inhibitor, and the mollusc heart controlled by only inhibitcr fibres. The annelids (Rogers, 1938) have both accelerator and inhibitor nerves. On the basis of work done by Botazzi (1902), and Lutz (1930) on the elasmobranch, i t is concluded that there is no sympathetic inns rvation to the heart although there is a parasympathetic inhibitor system. Among the amphi-bians and reptiles (Rogers, 1938) inhibitory and accelerator fibres pass to the heart. In the mammals (Winton and Bayliss, 1948) there is both sympathetic and parasympathetic control of the heart. It would seem that both vertebrate and inverte-brate groups have members which possess both excitatory and inhibitory nerves as well as those in which one or the other is lacking. Among vertebrates the teleosts are unique in that they lack both excitatory fibres and a sensitivity to adrenalin. CONCLUSIONS The hearts of Embryo herring ( 3 - 1 4 days sifter f e r t i l i z a t i o n ) , sockeye alevins ( 2 - 8 weeks after f e r t i l i z a tion) and adult g o l d f i s h are in h i b i t e d by acetylcholine but unaffected by adrenalin. The teleost-heart apparently lacks sympathetic innervation and the myocardial c e l l s are unres-ponsive to adrenalin. •?M. &3 bac^Ts: * j . /OOP /ID. /At-Appending© Figure 1. Kymograph Recordings of the goldfish heart Carasgius Jforatus before and after administering adrenalin i n concentration of: a. 1:1000 b. 1:^000 c. l:fj0U0 d. 1:5000 5s ^ • v Appendictee F igu re 2 . Kymograph r eco rd ings o f the g o l d f i s h hea r t C a r a s s i u s S t r a t u s before 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 concen t r a t i ono f 1:7500 1 Is 20,000 55 beats per minute 1$ beats per minute \f\j\f % - £ j„, .... AAAAAAAAAAAAAA <^ / £> (S. &** 73 '/• c . i Appendijfee F igu re 3. Kymograph r eco rd ings of the g o l d f i s h hear t Garass ius a a r a t u s before 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 : a . 1:20,000 b . 1:20,000 c . 1:50,000 * d. L:500,000 55 beats per minute 55 bea ts per minute Apoendiate* F i g u r e U . Kymograph r eco rd ings o f the g o l d f i s h hea r t Garass ius ^ u r a t u s before 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 : 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 eco rd ings o f the g o l d f i s h hea r t Garas s ius a a r a t u g before 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 b e a t s p e r m i n u t e 32 b e a t s p e r m i n u t e 1:50 b e a t s p e r m i n u t e 32.beats p e r m i n u t e A p p e n d ! * * F i g u r e 6 . K y m o g r a p h 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 a n d 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 i n c o n c e n t r a t i o n o f : a . 1:50 b. 1:50 c . 1:150 28. A p p e n d i f f c * T a b l e 1 - C o n t r o l s - h e a r t r a t e o f t h e g o l d f i s h C a r a s s i u s a u r a t u s i n b e a t s p e r m i n u t e c o u n t e d o v e r a p e r i o d b e t w e e n 50 a n d 95 m i n u t e s . N o . o f b e a t s p e r m i n u t e T i m e i n N o . o f b e a t s N o . o f b e a t s N o . o f b e a t s N o , m i n u t e s p e r m i n u t e . p e r m i n u t e p e r m i n u t e p e r m i n u t e : 0 53 31 36 7U 50 5 66 63 ks 56 82 10 • 6k 62 55 5U 7U 15 6h 62 62 50 69 20 62 63 59 h9 65 25 61 62 57 hi 62 30 60 60 56 h6 62 35 58 58 5U U5 63 UO 53 56 5U U3 63 U5 51 5U 5U U2 59 50 U9 5U 51 U0 5U 55 hi 52 50 50 60 hi 50 U8 U9 65 UU 50 hi U8 70 k2 k9 hi U9 75 U2 hi 80 U6 85 l i l * 90 U3 95 1*1 AppendiJ |p» Table 2 - Hear t r a t e of the g o l d f i s h i n beats pe r minute counted before 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 1:2,000 Time i n minutes No .o f beats pe r minute 0 5 10 15 20 25 30 35 1*0 1*5 50 55 60 65 70 75 80 85 90 95 29 U5 51* 50 A d r e n a l i n 1*3 1*3 1*0 37 36 No.o f bea ts pe r minute 52 58 57 56 A d r e n a l i n 50 51 1*9 1*5 1*1-i*o 39 hp 39 38 1*0 1*2 1*2 1*2 1*0 No.of bea ts pe r minute 32 1*2 1*6 1*0 31* A d r e n a l i n 32 11 No.o f bea ts p e r minute 75 71* 66 72 69 63 A d r e n a l i n 62 60 58 53 53 51 50 1*6 1*3 1*1 1*0 1*0 &8 Appendixes Table 3 - Hear t r a t e of the C a r a s s i u s auratus i n beats pe minute counted before 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 of 1:2U,000 Time i n No .o f bea ts No .o f bea ts minutes pe r minute p r minute 0 71 83 5 83 68 A d r e n a l i n 10 77 6U A d r e n a l i n 15 76 60 20 76 60 25 73 55 30 72 55 35 70 56 bP 68 5U U5 67 5L 63 U7 55 U2 60 35 65 31 70 3U 75 32 35 39 Appendiflfe-Table k - Hear t r a t e o f the g o l d f i s h i n beats pe r minute counted before 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 of 1:U2,000 Time i n minutes N o . o f beats No .o f beats No .o f beats pe r minute per minute pe r minute 0 105 5 101 10 95 15 90 20 80 25 75 A d r e n a l i n 30 67 35 60 U0 56 1*5 1*8 50 I46 55 1*3 60 U3 65 U5 1*1* 1*2 ho 33 32 32 35 73 A d r e n a l i n ho 56 Adrei 38 56 37 50 1*8 51 50 51 5U 1*9 56 1*7 65 1*1* 68 1*1 69 38 68 35 61* 3k 63 33 61* 32 6k 32 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 0 2 u 6 8 10 12 1U 16 18 20 22 2h 26 28 No.of beats No.of beats No.of beats per minute pr minute per minute 30 lil 32 31 111 33 32 Ul 33 32 uu 3U 33 uu 35 33 uu UO 3U ui UD 35 38 UO 35 UO 39 3U 38 38 3U 37 3U 35 35 35 36 J 3*. Appendi&ss Table 6 - Hear t r a t e o f h e r r i n g eggs (Clupea p a l l a s i i ) before 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 Time 0 2 h 6 8 10 12 11* 16 18 20 22 2k 26 28 30 32 3k 36 38 UO 1*2 hh hS 1*8 SO 52 5U 56 58 of 1:1,000 Beats 2k 2k 2k A d r e n a l i n 25 25 26 26 25 22 22 Beats Beats Beats Beats Beats 3k 33 30 32 2k 3k 3k 30 32 2k 3k 35 30 32 2k A d r e n a l i n 35 36 30 32 29 35 37 30 33 28 37 38 30 3k 33 39 kO 30 32 31 A d r e n a l i n 39 ko 29 3k 31 A d r e n a l i n 37 35 29 3U 27 38 38 29 3k 30 A d r e n a l i n 38 38 29 32 30 38 1*0 28 30 30 39 UO 28 29 30 39 1*1 28 29 29 35 1*1 27 31* 29 kk 31* 30 kk 36 31 1*6 38 32 Uo UO 33 1*1 3k 31* la 3k 31* 1*2 29 3k A d r e n a l i n 1*3 35 1*3 35 U3 36 1*3 1*3 U6 k6 ia i*i . Appendix©* Table 7 - Heart rate of herring eggs (Clupea pallasii) in beats per minute counted before and after administering adrenalin in concentration of 1:5,000 Time in minutes No.of beats per minute No.of beats per minute 0 23 29 2 2k 29 k 26 30 6 28 31 8 29 31 10 29 31 12 28 32 Adrenalin lk 28 32 16 22 32 18 20 32 20 21 33 22 23 33 2k 2k 33 26 2k 33 28 2k 33 30 2k 33 Adrenalin 32 2k 3k 3k 2k 3k 36 2k 38 2k Appendidte*. Table 8 - Hear t r a t e of h e r r i n g eggs (Clupea p a l l a s i i ) i n beats pe r minute before 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 of 1:25,000 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 n N o . o f beats No .o f beats N o . o f beats N o l o f beats s pe r minute pe r minute pe r minute pe r minute 3U U8 U2 5U 35 U8 U2 5U 3U U8 U2 5U 3U U9- U2 52 35 50 U3 52 A d r e n a l i n 35 52 UU 50 35 52 U7 . 50 35 52 U7 50 36 5U. U5 U8 35 5U UU U8 A d r e n a l i n 3U 56 UU 50 3U 56 U6 50 A d r e n a l i n 38 56 U6 50 38 56 U6 5o 39 U6 50 37 U5 50 37 U5 51 A d r e n a l i n 38 U8 52 39 U8 50 39 U8 50 UO U8 50 U9 50 U8 U7 U6 U5 U5 U5 Appendix©*- Table 9 - Hear t r a t e of h e r r i n g eggs (Clupea p a l l a s i i ) beats pe r minute counted before 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 1:10 Time i n minutes No. o f bea ts pe r minute 0 78 A c e t y l c h o l i n e 3 Hear t stopped comple t e ly AppendiKoe Table 10 - Hear t r a t e o f h e r r i n g eggs (Clupea p a l l a s i i ) beats pe r minute counted before 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 i n c o n c e n t r a t i o n o f 1:100 Time i n N o . o f bea ts i n N o . o f bea ts Time i n N o . o f beats minutes pe r minute minutes p e r minute minutes pe r minute 0 U3 0 37 0 • 3U 5 10 3 37 3 3k 10 k3 6 37 8 3k 15 I4.3 10 37 A c e t y l c h o l i n e 20 k3 13 37 9 36 A c e t y l c h o l i n e l U U2 10 35 21 k3 19 kk 12 3k 23 U2 A c e t y l c h o l i n e 13 33 2k k0 29 30 16 30 29 39 30 30 31 37 35 k3 3k 31 U0 kk 38 25 U5 27 53 36 50 36 59 39 55 30 6k k3 60 28 69 k$ 65 25 Ik U5 79 k9 8k k9 89 50 9k 50 10U kl 109 k6 119 U6 Appendiafc* Table 11 - Hear t r a t e of h e r r i n g eggs (Clupea p a l l a s i i ) i n beats pe r minute counted before 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 i n c o n c e n t r a t i o n of 1:300 Time i n No. o f beats Time i n N o . o f beats minutes pe r minute minutes pe r minute 0 U2 0 U8 2 U2 2 U8 3 U2 U U9 8 U2 6 U9 A c e t y l c h o l i n e 9 U8 13 33 11 k8 18 27 A c e t y l c h o l i n e 23 26 13 hh 28 30 Iii U3 15 U3 17 UO 18 39 19 38 22 39 23 UO 27 UI 28 UI 29 U3 32 U6 35 U8 37 53 39 52 U2 53 UU 55 U7 55 57 55 37. Appendices Table 12 a . Hear t r a t e o f salmon f r y (Oncorhynchus nerka) i n beats pe r minute counted before 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 1:100,000 b . Hear t r a t e o f salmon f r y (Oncorhynchus nerka) i n beats per minute counted before 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 c o n c e n t r a t i o n o f 1:100,000 a . a . a . a . a . a . a . a . a . a . b . J D C C t l / D . D B c l 0 5? 55 59 B l — W 1 65 68 69 78 65 2 6h 68 70 78 6k 3 65 69 71 79 6U . ... A d r e n a l i n k 68 69 71 77 55 68 6 1   •     69 72 69 73 1 70 7k I 70 76 1 A d r e n a l i n 70 83 ! Beats Beats Beats Beats Beats Beats 55 68 60 71 71 6U 60 65 59 6% 77 6k 6k 62 60 68 79 67 A d r e n a l i n 60 65 62 73 77 67 i 63 68 60 71 77 67 A d r e n a l i n 58 67 62 73 7U 68 59 67 6U 71 77 68 A d r e n a l i n 60 67 58 72 80 70 65 67 67 68 73 - 6U A d r e n a l i n 65 60 70 70 82 5 67 75 67 6 65 69 83 65 7 67  81 63 8 64 9 62 A d r e n a l i n ' A d r e n a l i n 1:100,000 10 60 70 82 68 6U 65 79 59 A d r e n a l i n 11 55 73 79 77 67 65 77 55 12 50 73 86 75 6k 6k 79 50 A d r e n a l i n 13 U8 73 83 75 65 65 77 61 1U 18 71 79 75 6U 65 79 59 71 82 62 6k Si !6 83 61 6k 17 58 68 18 58 70 ^ 58 20 l9 % <*> 22 60 23 60 $0. Appendixes Table 13 a . Hear t r a t e o f salmon f r y (Ohcorhynchus nerka) i n beats pe r minute counted before 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 of 1:50,000 b . Hear t r a t e of salmon f r y (Oncorhynchus nerka) i n beats pe r minute counted before 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 c o n c e n t r a t i o n of 1:50,000 Time i n minutes 0 1 2 3 u 5 6 7 8 9 10 11 12 13 I k 15 16 a. a. b . b . N o . o f beats N o . o f beats pe r minute pe r minute N o . o f beats No.of beats pe r minute p e r minute 56 57 58 60 59 59 67 6k 62 65 65 A d r e n a l i n 65 68 68 67 67 68 63 6k 65 62 60 67 6k 67 70 67 A d r e n a l i n 67 65 67 68 68 68 68 69 A c i d PH 2.U 1:50,000 67 71 60 60 6k 65 66 68 67 65 57 58 56 A c i d PH 2.k 1:50,000 65 65 65 6k 60 60 59 56 55 o Appendices- Table l k . a . Hear t r a t e of salmon f r y (Oncorhynchus nerka) i n beats p e r minutecounted before 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 of 1:33,000 b .Hea r t r a t e o f salmon f r y (Oncorhynchus nerka) i n beats pe r minute counted before 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 of 2.k i n c o n c e n t r a t i o n of 1:33,000 a . b . b . Time i n No .o f beats N o . o f beats No .o f beats minutes pe r minute pe r minute pe r minute 0 67 70 55 1 77 75 55 \ 2 77 75 55 3 77 71 57 A d r e n a l i n k 71 73 51 A d r e n a l i n A c i d PH 2.k 1:33,000 5 75 67 56 6 19 65 60 7 78 71 58 8 75 65 57 9 77 71 59 10 19 70 58 11 11 70 56 N 12 67 52 13 67 k6 l k 65 15 60 Appendices Table 15 a . Hear t r a t e o f salmon f r y (Oncorhynchus nerka) i n bea ts p e r minute counted before 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 of 1:20,000 b .Hea r t r a t e o f salmon f r y (Oncorhynchus nerka) i n bea ts pe r minute counted be fo re 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 of 2.U i n c o n c e n t r a t i o n o f 1:20,000 Time i n minutes 0 1 2 3 k 5 6 7 8 9 10 11 12 13 Hi a . N o . o f beats pe r minute 73 71 71 71 A d r e n a l i n 67 65 65 6U 65 6k 65 6U 6h 60 58 b . N o . o f beats pe r minute 6U 65 A c i d PH 2.U 1:20,000 59 61 65 67 56 58 Appendices- Table 16 a . Hear t r a t e of salmon f r y (Oncorhynchus nerka) i n beats pe r minute counted before 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 1:7,500 b . Hear t r a t e o f salmon f r y (Oncorhynchus nerka) i n beats per minute counted before 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 c o n c e n t r a t i o n o f 1:7,500 a . b . Time i n N o . o f bea ts No .o f bea ts minutes pe r minute pe r minute 0 71 68 1 73 67 A c i d PH 2.k 1:7,500 2 75 65 A d r e n a l i n 3 71 67 k 71 60 5 70 58 6 65 55 7 60 53 8 57 9 56 10 52 AppendiJSe* Table 17 Hear t r a t e o f salmon f r y (Oncorhynchus nerka) before 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 i n concen t r a t i on of a . 1:100 b . 1:500 c . 1:330 a . d . 1:200 e. 1:75 b . c . d . e. Time i n No .o f beats No .o f hea ts No .o f beats No.of beats No.of beats minute per minute per minute pe r minute pe r minute minutes per : 0 59 1 59 2 59 3 61 h 58 5 57 6 59 7 58 Acetylch< 1:1000 8 55 9 6h 10 68 11 62 12 71 13 6h i n 67 15 60 16 5U 17 57 18 68 65 59 58 55 58 59 62 6U 60 6U 70 60 61 60 67 60 75 61 75 A c e t y l c h o l i n e A c e t y l c h o l i n e 1:200 1:75 67 109 A c e t y l c h o l i n e A c e t y l c h o l i n e 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 fry (Oncorhynchus nerka) i n beats per minute counted before and after administering acid 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 0 1 2 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 62 67 60 3 75 U < 75 5 58 Acid PH 3-3 1:1000 6 75 7 ,75 8 75 9 75 10 75 11 75 12 75 13 75 Iii 75 15 75 16 61 17 68 18 62 19 70 20 75 21 75 76 72 75 68 77 81 79 Acid PH 1:330 81 3.3 72 72 68 72 Acid PH 3.3 1:200 72 86 68 Acid PH 3.3 1:500 72 81 68 72 86 75 72 86 70 72 86 70 68 86 65 61 86 70 67 86 63 68 86 70 61 67 63 60 65 70 68 67 75 Acid PH 3.3 1:75 65 73 65 56 60 35 35 Armst rong , P . B . 1931 B o t a z z i , F . 1901 B o t a z z i , F . 1902 B r i n l e y , F . J . 1932 B r i n l e y , F . J . 1935-Conant , F . S . and C l a r k , H . L ' . 1896 H i a t t , E . P . and G a r r y , ¥ . E . 1942 H i a t t , E . P . 1942 H i s , W.J. 1893 BIBLIOGRAPHY F u n c t i o n a l r e a c t i o n s i n the embryonic h e a r t , accompanying the ingrowth 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. C e n t l . F . P h y s i o l . ,14 (Quoted from B r i n l e y 1935). Z e i t s c h r . F . B i o l . ,43 (Quoted from L u t z 1930). A p h y s i o l o g i c a l s tudy o f the i n n e r v a t i o n o f the 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. Evidence f o r a sympathet ic 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 note 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. On the a c c e l e r a t o r and i n h i b i t o r nerves o f the c r a b ' s h e a r t . J o u r . E x p . M e d . , 1 t 341-347. Drug a c t i o n s on the spontaneously bea t ing t u r t l e v e n t r i c l e i n d i c a t i n g l a c k o f i n n e r v a t i o n . Amer. J o u r . P h y s i o l . , 138 : 758-762. 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 potass ium 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 d o g f i s h (Squalus  a c a n t h i a s ) . Amer. J o u r . P h y s i o l . , 139 t 45-48. Math . - P h y s . C l a s s e , B d . 18 S . 1-64 (Quoted from Armstrong J o u r . E x p . Z o o l . 58 t 43-61). Hogben, L.T. and Hobson, A.D. 1924 Lutz, B.R. 1930 Rogers, C.G. 1938 Winton, F.R. and Bayliss, L.E. 194-8 Young, J.Z. 1931" 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. The effect of adrenalin on the auricle of elasmobranch fishes. Amer. Jour. Physiol., 94 : 135-139. Textbook of Comparative Physiology, McGraw H i l l Book Co. Inc. Human Physiology, Blakiston. On the autonomic system of the teleost fish Aranoscopus scaber. Quart. Jour. Micro. Sci., 74 : 492-535. 

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