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Animal Behaviour
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e Animal Behaviour
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Mechanism, Development, Function and Evolution hn
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CHRIS BARNARD
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m Mechanism, Development, Function and Evolution
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‘Animal Behaviour is an important and popular subject, but all too often core texts focus on easy-to-digest
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aspects of the field. Barnard's book is different. He covers all the current, as well as historically important, e
aspects of animal behaviour with a clarity that encourages curiosity. This book takes a much needed, deep lo
and serious look at the most popular subject in organismal biology.’ pl
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Mike Siva-Jothy, University of Sheffield
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‘This excellent textbook deals with the complexities of animal behaviour in a readable and informative style. t
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Its balanced treatment of mechanisms, development, functions and evolution provides an overview which is
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lacking in most of the other textbooks in this area. It could be the main text for any animal behaviour course u
in biological sciences or psychology, but would also greatly benefit students taking courses in behavioural n
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ecology, evolution, experimental psychology and others.’ t
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Ian R Hartley, Biological Sciences, Lancaster University o
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Animal behaviour has been one of the fastest-growing scientific disciplines of recent years. Its impact nv
on the way we think about biology has spawned lucid ‘best-sellers’ like The Selfish Geneand d
widespread scientific and public debate about our view of the natural world and our place in it. This Ei
book provides a comprehensive introduction to the study of behaviour, from its basis in the animal’s v
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anatomy and physiology to its adaptive value in the environment. It is aimed at undergraduate
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students in the biological sciences and psychology and is designed to serve as both a detailed
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introduction and an extensive, up-to-date source of reference enabling students to pursue topics in io
the primary literature. n
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Features include:
• Comprehensive coverage of the four major levels of enquiry – mechanism,
development, function and evolution
• Integration of ideas and approaches from different disciplines of behaviour
study
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• An up-to-date focus on current debates
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• Extensive boxed features explaining underlying theory
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• Wide-ranging and extensively illustrated examples from international
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research literature
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• Clear introductory synopses and summaries for each chapter
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• Suggestions for further reading and numerous supporting references
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Chris Barnardis Professor of Animal Behaviour at the University of Nottingham and a recent
Executive Editor of the journal Animal Behaviour.
Cover image: Spoonbills. We have been unable to locate the
copyright holder of this image but if notified we should be CHRIS BARNARD
pleased to amend the acknowledgement in any future edition.
www.pearson-books.com
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Animal Behaviour
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We work with leading authors to develop the
strongest educational materials in animal behaviour
bringing cutting-edge thinking and best
learning practice to a global market.
Under a range of well-known imprints, including
Prentice Hall, we craft high quality print and
electronic publications which help readers to understand
and apply their content, whether studying or at work.
To find out more about the complete range of our
publishing, please visit us on the World Wide Web at:
www.pearsoned.co.uk
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Animal Behaviour
Mechanism, Development, Function and Evolution
Chris Barnard
Professor of Animal Behaviour, University of Nottingham, UK
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For my family
Pearson Education Limited
Edinburgh Gate
Harlow
Essex CM20 2JE
England
and Associated Companies throughout the world
Visit us on the World Wide Web at:
www.pearsoned.co.uk
First published 2004
© Pearson Education Limited 2004
The right of Chris Barnard to be identified as author of this work has been asserted
by him in accordance with the Copyright, Designs and Patents Act 1988.
All rights reserved. No part of this publication may be reproduced, stored in a retrieval
system, or transmitted in any form or by any means, electronic, mechanical,
photocopying, recording or otherwise, without either the prior written permission of the
publisher or a licence permitting restricted copying in the united Kingdom issued by the
Copyright Licensing Agency Ltd, 90 Tottenham C ourt Road, London W1T 4LP.
ISBN 0130899364
British Library Cataloguing-in-Publication Data
A catalogue record for this book is available from the British Library
Library of Congress Cataloging-in-Publication Data
Barnard, C. J. (Christopher J.)
Animal behavior : mechanism, development, function, and evolution / Chris Barnard.
p. cm.
Includes bibliographical references and index.
ISBN 0-13-089936-4
1. Animal behavior. I. Title.
QL751.B18563 2003
591.5–dc22 2003060928
10 9 8 7 6 5 4 3 2 1
09 08 07 06 05 04
Typeset in 9.5/12pt Sabon by 35
Printed and bound by Ashford Colour Press, Gosport, Hants
The publisher’s policy is to use paper manufactured from sustainable forests.
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Brief contents
1 Questions about behaviour 1
2 Evolution and behaviour 33
3 Physiological mechanisms and behaviour 94
4 Motivation and cognition 167
5 From genes to behaviour 220
6 Maturation and learning 255
7 Habitat choice and migration 308
8 Predators and prey 357
9 Social behaviour 406
10 Mating and parental care 470
11 Communication 533
12 Human behaviour 588
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Contents
Preface xv
Acknowledgements xviii
Chapter 1 Questions about behaviour 1
1.1 What is behaviour? 2
1.1.1 Repeatable measurement 3
1.1.2 The function of behaviour patterns 3
1.1.2.1 Purpose and goal-directedness: means to ends 4
1.1.2.2 Purpose and mind 7
1.2 Questions about behaviour 9
1.2.1 Tinbergen’s Four Whys 9
1.2.1.1 Socially mediated changes in brain and behaviour 11
1.2.2 Addressing the Four Whys 13
1.2.2.1 Experiments 13
1.2.2.2 Observation 13
1.2.2.3 Comparison 14
1.2.2.4 Theoretical models 14
1.3 Approaches to the study of behaviour 24
1.3.1 Ethology 25
1.3.2 Comparative psychology 26
1.3.3 Behavioural ecology 27
1.3.4 Evolutionary psychology 29
Summary 31
Further reading 32
Chapter 2 Evolution and behaviour 33
2.1 NeoDarwinism and the modern synthesis 34
2.1.1 The neoDarwinian synthesis 34
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viii x Contents
2.2 Phylogeny and behaviour 35
2.2.1 Phylogeny and courtship displays 37
2.2.2 Phylogeny and song development 39
2.3 Variation and heredity 41
2.3.1 Mendelian genetics and behaviour 41
2.3.1.1 Genes and Mendelian inheritance 41
2.3.1.2 Quantitative characters and heritability 44
2.3.2 Evidence that differences in behaviour are due to genes 45
2.3.2.1 Single genes and behaviour 46
2.3.2.2 Multiple genes 50
2.4 Natural selection and ‘selfish genes’ 53
2.4.1 Natural selection and sexual selection 56
2.4.2 Responses to selection 57
2.4.2.1 Responses to artificial selection 57
2.4.2.2 Adaptive differences between species populations 61
2.4.3 Fitness and adaptation 63
2.4.3.1 What is fitness? 63
2.4.3.2 Outlaw genes and extended phenotypes 64
2.4.4 Levels of selection 66
2.4.4.1 Group selection 66
2.4.4.2 Evolutionarily stable strategies 68
2.4.4.3 Optimality theory 77
2.4.4.4 Evolutionary models and the phenotypic gambit 82
2.4.4.5 Inclusive fitness 83
2.4.5 Life history strategies and behaviour 86
2.4.5.1 Life histories and alternative strategies 88
2.5 Cultural evolution 91
Summary 92
Further reading 93
Chapter 3 Physiological mechanisms and behaviour 94
3.1 Nervous systems and behaviour 96
3.1.1 Nerve cells and synapses 96
3.1.1.1 Nerve cell structure 96
3.1.1.2 Nerve cell function 98
3.1.1.3 Communication between neurons 98
3.1.2 Evolutionary trends in nervous systems and behaviour 99
3.1.2.1 Unicellular organisms 99
3.1.2.2 Nervous systems and behaviour in invertebrates 99
3.1.2.3 Nervous systems and behaviour in vertebrates 104
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Contents x ix
3.1.3 Nervous systems and the adaptive organisation of behaviour 109
3.1.3.1 Neuroanatomy and adaptive behaviour 109
3.1.3.2 Is adaptive behaviour ‘hard-wired’? 118
3.1.3.3 Command centres and neural hierarchies 125
3.1.3.4 Deducing mechanism from behaviour 128
3.2 Sensory mechanisms, perception and behaviour 133
3.2.1 Sensory mechanisms, stimulus filtering and perception 133
3.2.1.1 Visual stimulus filtering 133
3.2.1.2 Auditory stimulus filtering 138
3.2.1.3 Other mechanoperception 140
3.2.1.4 Olfactory and gustatory perception 141
3.2.1.5 Electromagnetic perception 142
3.3 Hormones and behaviour 144
3.3.1 How hormones affect behaviour 144
3.3.1.1 Effects on the nervous system 144
3.3.1.2 Effects on sensory perception 146
3.3.1.3 Effects on effector systems 147
3.3.1.4 Effects on development 148
3.3.1.5 Factors influencing relationships between hormones and behaviour 149
3.4 Neural and hormonal mechanisms and the long-term control
of behaviour: biological rhythms 154
3.4.1 Rhythmic behaviour 154
3.4.1.1 Interactions between clocks 157
3.4.1.2 The physiological basis of endogenous clocks 157
3.5 Mechanism and constraint 159
3.5.1 Mechanism as a con straint on function 160
3.5.1.1 Perceptual constraints 160
3.5.1.2 Motor constraints 161
3.5.2 Function as a constraint on mechanism 163
3.5.2.1 Testosterone, immunity and behaviour 163
Summary 165
Further reading 166
Chapter 4 Motivation and cognition 167
4.1 Causation and motivation 167
4.1.1 Models of motivation 168
4.1.1.1 Intervening variables, drive and energy models of motivation 168
4.1.1.2 Homeostatic models of motivation 174
4.1.1.3 The state-space approach 176
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