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The central nervous system and fish behaviour
A&m. Behav., 1969, 17, 593-596
BOOK REVIEWS the problem of restoration of function. Their data lend support to a remarkable degree of point-to-point specific regeneration. Aronson and Kaplan survey, in - some detail, the behavioural studies of teleost forebrain function. Their data demonstrate that forebrain lesions can depress or change certain behaviour patterns. The organization of behaviour patterns is conceived as taking place in the lower centrcs, while the forebrain facilitates or inhibits the behaviour in a non-specific manner. The theory is presented that the teleost forebrain is an arousal centre, capable of adjusting to changing conditions. Savage also finds that learning and memory can take place in a forebrain-less fish, but with a deficit in nerformance. In different words. he suonorts the notion of the forebrain as a facilitating ‘centre.-With a feeling of dL;ici vu (harking back to Iashley), one reads: ‘We should avoid the simplistic notion that “memory” is the property of any single structure . . .” In the last paper of this section. Reaestein describes avoidance conditioninn deficits’resufing from hypothalamic lesions. He points out that these deficits are qualitatively different from those produced by forebrain lesions. His experiment and his conclusions are coloured by the implicit acceptance of the ‘fear’ hypothesis of avoidance conditioning, and he ascribes the performance deficit to hyperemotionality, i.e. the fish is ‘scared’. Neither ‘fear’ nor ‘emotionality’ are defined for the fish. Indeed it is difficult to define these essentially anthropomorphic terms in any animal without giving due consideration to the concept of levels of organization. Part III comprises five somewhat unrelated chapters. With exceptional clarity. Bennett summarizes about 14 years of &search on the physiology and behaviour of electric fishes. The problem of synchrony of discharge by electric organs is given considerable attention. The nature of the ‘command’ and ‘pacemaker’ centres is discussed, and qualitative differences in organization of these centres are shown to exist among the different electric species, many of which represent extremely diverse sections of the phylogenetic tree of fishes. The fact that some species can bc conditioned to alter their electric discharge rates is of considerable interest, and this provides a new and promising tool for the study of a variety of bioelectrical phenomena. In the first of two papers on the effects of low temperature on the brain and behaviour of fish,. Prosser and Nagai present data on the complex of relations between nerve conduction, classical and instrumental conditioning as blocked by cold. Elcctrophysiological studies confirm that deeper (pyriform layer) tectal responses and conditioned responses are more sensitive to cold. Ectothermic animals offer some interesting opportunities for the study of temperature effects on physiology and behaviour, and Rozin presents a strong case for this type of research. (He uses the term ‘poikilothermy’ which now seems to be going out of fashion.) Experiments are reviewed in which fish can be instrumentally conditioned to bring their environmental temperature down to a preferred level. Short-duration temporal rhythms could be conditioned, and these appear to be temperature independent. Agranoff and Davis review some of their previous studies of puromycin effects on retention of an avoidance resoonse in goldfish. The results are inter-meted as inhibition of proiein synthesis at the critical- time of memory consolidation. A somewhat low criterion for
The Central Nervous System and Fish Behaviour. Edited by DAVID INGLE. Chicago: University of Chicago Press (1968). Price: $15. This volume, dedicated to the foremost of American neurologists C. Judson Herrick, resulted from a symposium held at the University of Chicago in April 1967. Major topics include visual systems, brain function and behaviour of fishes. The contributtons range widely in topic, coverage and quality: from neurophysiology and biochemistry of brain function to conditioned behaviour: from topical reviews and reports of new data to repetitions of prior publications. The first section of six chapters on vision in teleosts is the most coherent. The introductory chapter by Schwassmann and Kruger is a capsule survey of neurology of the visual system. Electrophysiological mapping of the optic tectum is shown to represent a fairly close point-to-point correspondence with the retina. In a short but meaty chapter by Jacobson, the neurophysiological approach to fish vision is exemplii%d. Colour-coding can be detected in the retina at various levels and in the optic tectum. The exact function of t&al units, however, is-left as a major question. Yager presents a review of his own studies on the behavioural approach to colour vision, and reports his data on relative photopic spectral sensitivity. The relation of these and other data to pigment absorption curves shows some significant departures, but the basic trichromatic system is supported. The next three papers, also on vision, are more concerned with higher level integrative functions of the visual svstem. In what is nrimarilv a review of his own extensive work, Sutherland ~ummhzes his studies on shape discrimination in gold&h, as compared to octopus and rats. Aside from the well-known fact that fish do not generalize as easily as mammals, it is quite apparent that fish use different kinds of cues in learning and discriminating shapes. Considering the vast phylogenetic gulf between fish and mammal, and fish and octopus, such major qualitative differences in perception are not surprising. Unfortunately, the evolutionary and ecological aspects of these differences are not treated. Some concern with ecology is shown by Ingle in his chapter on visuomotor nroblems in fish. He also points out major differences between species and shows that orientation and identification of obiects is coded differently in fish and in birds. With so much weight of evidence on response specificity and coding in the sense organs, it is surprising that Ingle falls back on the well-worn lock-and-key analogy of stimulus and response in the central nervous system. The multitude of experiential factors that channel the development of the response system are not even mentioned. The major chapter (33 pages) by Trevarthen includes several very elegant diagrams of the visual field of fish and its relation to locomotion. This is a most interesting and challenging review of the optomotor response, its relation to locomotion and to ecological characteristics. A hypothesis of the evolution of vision in fish is presented, along with many stimulating and novel ideas on visuomotor integration. This chapter is clearly the highlight of the section. Part II is on the teleost forebrain, and is introduced bv a short statement bv Schnitzlein in which the problems of nomenclature and homology of the telencephalon are brought into focus. Buamgarten and Miessner summarize their data on olfactory tract regeneration and pose 593
594
ANIMAL
BEHAVIOUR,
acquisition is considered acceptable in these experiments. Shashoua presents a most unusual task for a goldfishto learn to swim upright with a buoyant float attached under its chin. Tritiated erotic acid is administered, and trained fish show a significant increase in labelled RNA in the brain. Puromycin reduces this increase, with a correlated decrement in the performance of the task. This is indeed a drastic treatment to the fish, and certainly involves adjustments in several sensory and motor systems. ‘Memory’ in this instance undoubtedly involves many peripheral as well as central factors. The conclusions of both these papers tend to be somewhat oversirnolified and reductionistic. ‘Memorv’ is confounded with a phenomenon that is far more complex than the word. It is evident that protein synthesis is involved in retention of performance, but it is hard to find any cytoplasmic function in which protein synthesis is not involved. We still seem to be searchine for the enzram. Part IV consists of four chapters on molar aspects of fish behaviour. In a somewhat rambling, instinctivist approach Barlow presents arguments toward a search for behavioural units-fixed action patterns. What makes certain actions unitary and ‘natural’ is not clear. His conclusions show an attempt to reconcile the more rigid aspects of ethology with the discovery that quantitative studies of behaviour reveal extensive variability. Unfortunately, there seems to be no appreciation of the role of development of behaviour, and the discredited dichotomy of nature v. nurture is still central. Some experiments on conditioning of sticklebacks are reported by Sevenster. Essentially, he demonstrates that fish can be conditioned to perform a variety of tasks, some of which seem complex but are actually no more so than many other operants, such as paddle-pushing. In a concise chapter, Hasler summarizes much of the work on homing and sun orientation in fishes. The tinal paper is a summary by Bitterman of the studies coming from his laboratory on reversal and probability conditioning in fishes. Although the qualitative differences in learning between fish and mammals are evident, the evolutionary and developmental factors that underlie these differences need to be explored further. In summary, as is the case with many symposium volumes, there is unevenness in quality and coverage. Much of the material has been published elsewhere. In spite of this, the volume represents a unique compilation of recent advances in the field, especially in the section on vision. It will have a long life as a valuable reference work and source book for any serious student of animal behaviour. WILLIAM
N.
TAVOLGA
Ecological Adaptations to Breeding in Birds. By DAVID LACK. London: Methuen (1969). Price 84s. This is not so much a book as an extended scientific paper. It is also a further round in a polemical duel, mainly between Dr Lack and his supporters and Professor Wynne-Edwards and his supporters, that many people will now feel has been carried about as far as is usefully possible using the present weapons. The weapons may briefly be described as bombardment by enormous mounds of facts. The dispute is about whether, as WynneEdwards claims, animal populations are controlled by self-balancing behavioural devices, or, as Lack reioins. purely by adaptations basically operated by food supply: Wvnne-Edwards would agree that food is the basic controlling factor,. but safs that in many animals the self-balancing devices come into play to forestall the
17,
3
ultimate arbitrament of hunger. Lack, on the other hand, says that this involves group selection, which is impossible, because we know that inheritance is in fact by individuals. However, one cannot help feeling that this is one of the arguments where both sides are right in some degree and wrong in some degree. The solution will probably be found in the direction of further work on the adaptive significance of social and solitary habits; will somebody please look more closely at the relationship between Wynne-Edwards’s supposed group selection (the term is his opponents’ rather than his own) and the entirely respectable (from the point of view of orthodox Darwinians) adaptive value of animals living socially? This extended introduction is necessary to explain why David Lack has found it necessary to read through the enormous amount of literature and review, most valuably, the breeding adaptations of the class Aves as a whole. Though one may feel it is flogging a dead horse to go to all this trouble to establish the adaptive value of all all these aspects of a bird’s breeding cycle, nesting dispersion, the pair bond, clutch size and so forth, the value of the work when done is undoubted. It shows that the amateurs who over the past hundred years have been steadily amassing facts about birds as and when they came across them have not by any means been wasting their time. Indeed, they have not done enough of it, for there are many birds and even whole families for which our information is sketchy in the extreme. David Lack has nroduced an invaluable source-book. in the course of providing himself with a large pile of &nmunition wherewith to bombard the Wynne-Edwards School. R. S. R. FIITER Behavioral
Pharmacology. By T. THOMPSON & C. R. Englewood Cliffs: Prentice Hall Inc., New Jersey. (1968). Price: $10.50. This is an excellent, pragmatic text which brings to the student and research worker a clear statement of the potential and the aims of this relatively new, but very active field of behavioural pharmacology. Such a text was surely needed by teachers and students both in psychology and in pharmacology. The authors are realistic in recognizing that ‘it would be misleading to suggest that a &fled-discipline concerned with the behavioral actions of drugs has already evolved’. They have instead tried to present the important variables inherent in any study of the effects of drugs on animal behaviour. In recognizing the potential complexity of these relationships, one may hope that experimenters will strive to uncover the lawfulness of such relationships and will not be satisfied merely to observe interesting drug effects. The text consists of 297 pages divided into five sections. with a glossary of technic& terms, bibliography and index. Section I consists of a short introduction justifying the existence of behavioural pharmacology as a basic science. Section II presents very briefly, in chapter 2, the pharmacological principles of drug action. Chapter 3 is a short introduction to the classification of the major groups of behaviourally active drugs. The student should be recommended to pharmacological texts for more detailed information. Section III includes chapters on the principles of experimental analysis of behaviour and the classification of behaviour. These chapters reflect the influence of Skinner’s descriptive behaviourism on the authors and serve SCHUSTER.
BOOK REVIEWS the problem of restoration of function. Their data lend support to a remarkable degree of point-to-point specific regeneration. Aronson and Kaplan survey, in - some detail, the behavioural studies of teleost forebrain function. Their data demonstrate that forebrain lesions can depress or change certain behaviour patterns. The organization of behaviour patterns is conceived as taking place in the lower centrcs, while the forebrain facilitates or inhibits the behaviour in a non-specific manner. The theory is presented that the teleost forebrain is an arousal centre, capable of adjusting to changing conditions. Savage also finds that learning and memory can take place in a forebrain-less fish, but with a deficit in nerformance. In different words. he suonorts the notion of the forebrain as a facilitating ‘centre.-With a feeling of dL;ici vu (harking back to Iashley), one reads: ‘We should avoid the simplistic notion that “memory” is the property of any single structure . . .” In the last paper of this section. Reaestein describes avoidance conditioninn deficits’resufing from hypothalamic lesions. He points out that these deficits are qualitatively different from those produced by forebrain lesions. His experiment and his conclusions are coloured by the implicit acceptance of the ‘fear’ hypothesis of avoidance conditioning, and he ascribes the performance deficit to hyperemotionality, i.e. the fish is ‘scared’. Neither ‘fear’ nor ‘emotionality’ are defined for the fish. Indeed it is difficult to define these essentially anthropomorphic terms in any animal without giving due consideration to the concept of levels of organization. Part III comprises five somewhat unrelated chapters. With exceptional clarity. Bennett summarizes about 14 years of &search on the physiology and behaviour of electric fishes. The problem of synchrony of discharge by electric organs is given considerable attention. The nature of the ‘command’ and ‘pacemaker’ centres is discussed, and qualitative differences in organization of these centres are shown to exist among the different electric species, many of which represent extremely diverse sections of the phylogenetic tree of fishes. The fact that some species can bc conditioned to alter their electric discharge rates is of considerable interest, and this provides a new and promising tool for the study of a variety of bioelectrical phenomena. In the first of two papers on the effects of low temperature on the brain and behaviour of fish,. Prosser and Nagai present data on the complex of relations between nerve conduction, classical and instrumental conditioning as blocked by cold. Elcctrophysiological studies confirm that deeper (pyriform layer) tectal responses and conditioned responses are more sensitive to cold. Ectothermic animals offer some interesting opportunities for the study of temperature effects on physiology and behaviour, and Rozin presents a strong case for this type of research. (He uses the term ‘poikilothermy’ which now seems to be going out of fashion.) Experiments are reviewed in which fish can be instrumentally conditioned to bring their environmental temperature down to a preferred level. Short-duration temporal rhythms could be conditioned, and these appear to be temperature independent. Agranoff and Davis review some of their previous studies of puromycin effects on retention of an avoidance resoonse in goldfish. The results are inter-meted as inhibition of proiein synthesis at the critical- time of memory consolidation. A somewhat low criterion for
The Central Nervous System and Fish Behaviour. Edited by DAVID INGLE. Chicago: University of Chicago Press (1968). Price: $15. This volume, dedicated to the foremost of American neurologists C. Judson Herrick, resulted from a symposium held at the University of Chicago in April 1967. Major topics include visual systems, brain function and behaviour of fishes. The contributtons range widely in topic, coverage and quality: from neurophysiology and biochemistry of brain function to conditioned behaviour: from topical reviews and reports of new data to repetitions of prior publications. The first section of six chapters on vision in teleosts is the most coherent. The introductory chapter by Schwassmann and Kruger is a capsule survey of neurology of the visual system. Electrophysiological mapping of the optic tectum is shown to represent a fairly close point-to-point correspondence with the retina. In a short but meaty chapter by Jacobson, the neurophysiological approach to fish vision is exemplii%d. Colour-coding can be detected in the retina at various levels and in the optic tectum. The exact function of t&al units, however, is-left as a major question. Yager presents a review of his own studies on the behavioural approach to colour vision, and reports his data on relative photopic spectral sensitivity. The relation of these and other data to pigment absorption curves shows some significant departures, but the basic trichromatic system is supported. The next three papers, also on vision, are more concerned with higher level integrative functions of the visual svstem. In what is nrimarilv a review of his own extensive work, Sutherland ~ummhzes his studies on shape discrimination in gold&h, as compared to octopus and rats. Aside from the well-known fact that fish do not generalize as easily as mammals, it is quite apparent that fish use different kinds of cues in learning and discriminating shapes. Considering the vast phylogenetic gulf between fish and mammal, and fish and octopus, such major qualitative differences in perception are not surprising. Unfortunately, the evolutionary and ecological aspects of these differences are not treated. Some concern with ecology is shown by Ingle in his chapter on visuomotor nroblems in fish. He also points out major differences between species and shows that orientation and identification of obiects is coded differently in fish and in birds. With so much weight of evidence on response specificity and coding in the sense organs, it is surprising that Ingle falls back on the well-worn lock-and-key analogy of stimulus and response in the central nervous system. The multitude of experiential factors that channel the development of the response system are not even mentioned. The major chapter (33 pages) by Trevarthen includes several very elegant diagrams of the visual field of fish and its relation to locomotion. This is a most interesting and challenging review of the optomotor response, its relation to locomotion and to ecological characteristics. A hypothesis of the evolution of vision in fish is presented, along with many stimulating and novel ideas on visuomotor integration. This chapter is clearly the highlight of the section. Part II is on the teleost forebrain, and is introduced bv a short statement bv Schnitzlein in which the problems of nomenclature and homology of the telencephalon are brought into focus. Buamgarten and Miessner summarize their data on olfactory tract regeneration and pose 593
594
ANIMAL
BEHAVIOUR,
acquisition is considered acceptable in these experiments. Shashoua presents a most unusual task for a goldfishto learn to swim upright with a buoyant float attached under its chin. Tritiated erotic acid is administered, and trained fish show a significant increase in labelled RNA in the brain. Puromycin reduces this increase, with a correlated decrement in the performance of the task. This is indeed a drastic treatment to the fish, and certainly involves adjustments in several sensory and motor systems. ‘Memory’ in this instance undoubtedly involves many peripheral as well as central factors. The conclusions of both these papers tend to be somewhat oversirnolified and reductionistic. ‘Memorv’ is confounded with a phenomenon that is far more complex than the word. It is evident that protein synthesis is involved in retention of performance, but it is hard to find any cytoplasmic function in which protein synthesis is not involved. We still seem to be searchine for the enzram. Part IV consists of four chapters on molar aspects of fish behaviour. In a somewhat rambling, instinctivist approach Barlow presents arguments toward a search for behavioural units-fixed action patterns. What makes certain actions unitary and ‘natural’ is not clear. His conclusions show an attempt to reconcile the more rigid aspects of ethology with the discovery that quantitative studies of behaviour reveal extensive variability. Unfortunately, there seems to be no appreciation of the role of development of behaviour, and the discredited dichotomy of nature v. nurture is still central. Some experiments on conditioning of sticklebacks are reported by Sevenster. Essentially, he demonstrates that fish can be conditioned to perform a variety of tasks, some of which seem complex but are actually no more so than many other operants, such as paddle-pushing. In a concise chapter, Hasler summarizes much of the work on homing and sun orientation in fishes. The tinal paper is a summary by Bitterman of the studies coming from his laboratory on reversal and probability conditioning in fishes. Although the qualitative differences in learning between fish and mammals are evident, the evolutionary and developmental factors that underlie these differences need to be explored further. In summary, as is the case with many symposium volumes, there is unevenness in quality and coverage. Much of the material has been published elsewhere. In spite of this, the volume represents a unique compilation of recent advances in the field, especially in the section on vision. It will have a long life as a valuable reference work and source book for any serious student of animal behaviour. WILLIAM
N.
TAVOLGA
Ecological Adaptations to Breeding in Birds. By DAVID LACK. London: Methuen (1969). Price 84s. This is not so much a book as an extended scientific paper. It is also a further round in a polemical duel, mainly between Dr Lack and his supporters and Professor Wynne-Edwards and his supporters, that many people will now feel has been carried about as far as is usefully possible using the present weapons. The weapons may briefly be described as bombardment by enormous mounds of facts. The dispute is about whether, as WynneEdwards claims, animal populations are controlled by self-balancing behavioural devices, or, as Lack reioins. purely by adaptations basically operated by food supply: Wvnne-Edwards would agree that food is the basic controlling factor,. but safs that in many animals the self-balancing devices come into play to forestall the
17,
3
ultimate arbitrament of hunger. Lack, on the other hand, says that this involves group selection, which is impossible, because we know that inheritance is in fact by individuals. However, one cannot help feeling that this is one of the arguments where both sides are right in some degree and wrong in some degree. The solution will probably be found in the direction of further work on the adaptive significance of social and solitary habits; will somebody please look more closely at the relationship between Wynne-Edwards’s supposed group selection (the term is his opponents’ rather than his own) and the entirely respectable (from the point of view of orthodox Darwinians) adaptive value of animals living socially? This extended introduction is necessary to explain why David Lack has found it necessary to read through the enormous amount of literature and review, most valuably, the breeding adaptations of the class Aves as a whole. Though one may feel it is flogging a dead horse to go to all this trouble to establish the adaptive value of all all these aspects of a bird’s breeding cycle, nesting dispersion, the pair bond, clutch size and so forth, the value of the work when done is undoubted. It shows that the amateurs who over the past hundred years have been steadily amassing facts about birds as and when they came across them have not by any means been wasting their time. Indeed, they have not done enough of it, for there are many birds and even whole families for which our information is sketchy in the extreme. David Lack has nroduced an invaluable source-book. in the course of providing himself with a large pile of &nmunition wherewith to bombard the Wynne-Edwards School. R. S. R. FIITER Behavioral
Pharmacology. By T. THOMPSON & C. R. Englewood Cliffs: Prentice Hall Inc., New Jersey. (1968). Price: $10.50. This is an excellent, pragmatic text which brings to the student and research worker a clear statement of the potential and the aims of this relatively new, but very active field of behavioural pharmacology. Such a text was surely needed by teachers and students both in psychology and in pharmacology. The authors are realistic in recognizing that ‘it would be misleading to suggest that a &fled-discipline concerned with the behavioral actions of drugs has already evolved’. They have instead tried to present the important variables inherent in any study of the effects of drugs on animal behaviour. In recognizing the potential complexity of these relationships, one may hope that experimenters will strive to uncover the lawfulness of such relationships and will not be satisfied merely to observe interesting drug effects. The text consists of 297 pages divided into five sections. with a glossary of technic& terms, bibliography and index. Section I consists of a short introduction justifying the existence of behavioural pharmacology as a basic science. Section II presents very briefly, in chapter 2, the pharmacological principles of drug action. Chapter 3 is a short introduction to the classification of the major groups of behaviourally active drugs. The student should be recommended to pharmacological texts for more detailed information. Section III includes chapters on the principles of experimental analysis of behaviour and the classification of behaviour. These chapters reflect the influence of Skinner’s descriptive behaviourism on the authors and serve SCHUSTER.