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The Induction of Parental Behavior in the Blue Gourami, Trichogaster Trichopterus (Pisces, Belontiidae)

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The Induction of Parental Behavior in the Blue Gourami, Trichogaster Trichopterus (Pisces, Belontiidae) THE INDUCTION OF PARENTAL BEHAVIOR IN THE BLUE GOURAMI, TRICHOGASTER TRICHOPTERUS (PISCES, BELONTIIDAE) by DONALD LAWRENCE KRAMER B.Sc, Boston College, 1966 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in the Department of ZOOLOGY We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA July, 1971 In presenting this thesis in partial fulfilment of the require• ments for an advanced degree at the University of British Col• umbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the Head of my Department or by his representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permis• sion. Department of The University of British Columbia Vancouver 8, British Columbia ii ABSTRACT The blue gourami, Trichogaster trichopterus, is a small, freshwater tropical fish, native to Southeast Asia. Parental be• havior is normally shown only by males after spawning. Females and non-parental males eat small numbers of test eggs which they are given, while parental males retrieve them to their nests. The goal of this thesis was to investigate the factors responsible for the sudden change in behavior from egg-eating to parental care which takes place at the time of spawning. In the initial section, the patterns of parental behavior are described, and quantitative data on the development and main• tenance of parental behavior in male fish spawning for the first time are presented. The experimental studies reveal that stimuli from the eggs are an important factor in the induction of parental behavior. Many naive males developed parental behavior within 0.5-2.5 min when given eggs spawned by other fish. This response was depen• dent upon the number of eggs presented: a large proportion of fish became parental when given 2000-3000 eggs, a smaller propor• tion did so when given 500-1000 eggs, and none did so when given 100 eggs. However, even 100 eggs evidently affected parental responsiveness because some fish eventually developed complete parental behavior when repeatedly given 100 eggs. These results indicate that eggs can induce parental behav• ior, not that they do so in the natural spawning situation. However, tests of males spawning with females whose oviducts were plugged indicated that exposure to eggs was an important compo- iii nent in the natural development of parental behavior. Spawning by itself did not induce fish to become parental, but it did re• duce the number of eggs necessary for parental behavior to de• velop. Besides spawning and stimuli from the eggs, androgens also play a role in the development of parental behavior. It was shown that, after spawning, males performed more parental beTiav- ior than females, although detailed observations of females re• vealed that they had some capacity to perform parental behavior. The masculinization of females by means of methyl testosterone implants allowed them to develop parental behavior in response to large numbers of eggs. Whether castration of males eliminated their capacity to perform parental behavior could not be deter• mined with certainty. The presence of young facilitates the maintenance of paren• tal responsiveness to eggs when males with developing broods are compared with males whose broods have been removed. Apparently, physical contact v/ith the young is necessary because males lose their parental responsiveness when their broods are placed in, baskets which allow only visual and chemical contact. An important result of this study is the demonstration of the significance of stimulus strength in the induction of paren• tal responsiveness. This factor has not often been taken into consideration in other studies of parental behavior. By varying stimulus strength, repetition of stimuli, and sexual experience, patterns of the induction of parental behavior were found in gouramis which were similar to those observed in other studies on a variety of mammals, birds, and fish. That is, parental iv behavior developed either almost immediately or gradually with the presentation of young to non-parental fish, or it was shown only if the young were presented during the "sensitive period" after spawning. The finding of such different patterns in a single species as a result of varying the strength of stimuli from the eggs sug• gested that the motivating effects of eggs and young on parental responsiveness may represent a common pattern underlying the con• trol of parental behavior in a variety of vertebrate species. This possibility and the relationship between hormones and stim• uli from the young in the control of parental behavior are dis• cussed in the final chapter. V TABLE OF CONTENTS PAGE I. INTRODUCTION 1 Chapter 1. Introduction to the Problem and Methods. 1 A. The Problem 1 B. The Animal 2 C. The Methods 3 1. Culture 3 2. Maintenance 3 3. Introduction of Spawning 6 LL. Collection of Eggs 9 5. Surgical Procedures . 10 6. Behavioral Observations 12 D. Summary 12 Chapter 2. Reproductive Behavior of the Blue Gourami . 13 A. Qualitative Description 13 1. Nest Building 13 2. Spawning 14 3. Parental Behavior 15 LL. Descriptions and Definitions of Behavior Used in Recording , 16 5. Development of the Young. ..... 20 B. A Quantitative Study of the Natural Induction of Parental Behavior 21 1. Introduction 21 2. Methods 21 3. Results and Discussion 23 vi PAGE C. A Quantitative Study of the Maintenance of Parental and Nest Building Behavior 27 1. Introduction 27 2. Methods 27 3. Results and Discussion 29 D. Summary ...... 34 II. FACTORS INVOLVED IN THE INDUCTION OF PARENTAL BEHAVIOR. ........ 36 Chapter 3. The Role of Eggs 36 A. Induction of Parental Behavior v/ith Clutches Spawned by Other Fish 36 1. Introduction 36 2. Methods 37 3. Results and Discussion. ..... 38 B. The Effect of the Number of Stimulus Eggs on the Induction of Parental Behavior. ....... 40 1. Introduction 40 2. Methods 41 3. Results 43; 4. Discussion 51 C. Induction of Parental Behavior by Repeated Exposure to Small Numbers of Eggs 53 1. Introduction. 53 2. Methods 54 3. Results and Discussion 56 D. Summary 60 vii PAGE Chapter L\. The Stimulus for the Induction of Parental Behavior . 6l 1. Introduction 61 2. Results and Discussion 62 Chapter 5« The Role of Performance of the Spawning Act 65 A. The Role of Spawning 65 1. Introduction 65 2. Methods 66 3. Results ..... 68 LL. Discussion 70 B. The Effect of Spawning on the Speed of Develop• ment of Parental Behavior . 72 1. Introduction 72 2. Methods 72 3. Results and Discussion 73 C. Summary 76 Chapter 6. The Role of Androgens 77 A. Sexual Dimorphism in Parental Behavior 77 1. Introduction 77 2. Methods . 78 3. Results and Discussion. 80 B. Effect of Methyl Testosterone on the Parental and Nest Building Behavior and the Secondary Sex Characters of Female Blue Gouramis 85 1. Introduction . 85 viii PAGE 2. Methods 86 3. Results and Discussion 89 C. The Effects of Castration on the Parental and Nest Building Behavior of Male Gouramis 97 1. Introduction. 97 2. Methods 98 3. Results 101 A. Discussion 103 D. Hormones and the Parental and Nest Building Behavior of Fish 106 E. Summary . 109 III. MAINTENANCE OF PARENTAL BEHAVIOR Ill Chapter 7. The Role of Young in the Maintenance of Parental Behavior Ill 1. Introduction HI 2. Methods . HI 3. Results 113 if. Discussion 114 5.- Summary ........ .118 IV. GENER& DISCUSSION 119 Chapter 8. General Conclusions About the Control of Parental Behavior in the Gourami 119 Chapter 9. Effects of Eggs and Young on Parental Responsiveness in Vertebrates 124 A. Introduction. 124 ix PAGE B. Evidence for Motivational Effects of Eggs and Young 125 1. Animals Initially Unresponsive: Parental Behavior After Extended Expo• sure to Young • . 126 2. Animals Initially Responsive to Young: Changes in Responsiveness to Weak Stimuli After Exposure to Strong Stimuli. 130 3. Animals Responsive to Young Only at Particular Times: Sensitive Periods for Effects of Eggs and Young on Responsiveness . 131 if. Changes in the Intensity of Parental Behavior 132 5. Role of Eggs and Young in the Maintenance of Parental Responsiveness 133 C. The Nature of the Motivating Stimuli From the Young 136 D. The Mechanism by Which Young Affect Paren• tal Responsiveness 138 E. The Role of Motivational Effects of the Young in Relation to the Role of Hormones in the Induction of Parental Behavior . 140 F. Suggestion of a Common Pattern in the Motiva• tional Effects of Young 143 BIBLIOGRAPHY. ..... 147 X LIST OF TABLES TABLE PAGE I Number of fish building nests of different types before spawning and from 0 to 13 days after spawning .........32 II The number of eggs surviving out of 100 pre• sented in the overnight test 39 III Number of fish developing parental behavior after presentation of different numbers of stimulus eggs k7 IV The effect of the number of stimulus eggs on measures of latency and strength of parental behavior in male gouramis 48 V The effect of 5 modifications of the stimulus for the induction of parental behavior on egg- eating inhibition in male gouramis 63 VI The effect of exposure to eggs and performance of the spawning act on the induction of egg- eating inhibition in male gouramis without previous reproductive experience. .... 69 VTI Effects of spawning experience on parental behavior in male gouramis presented with 1.5 ml eggs 75 VIII Sexual dimorphism in the parental behavior of gouramis 81 IX Mean and range of weights in mg of steroid implanted into female gouramis . 87 xi table page X Parental behavior of female gouramis implanted vrith methyl testosterone or cholesterol and presented with the stimulus of 1.5 ml recently spawned eggs 93 XI Statistical comparisons of the effect of the presence of young on the maintenance of parental behavior .
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