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Perception and Motor Control in Birds: An Ecological Approach PDF

374 Pages·1994·8.065 MB·English
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M.N.O. Davies P.R. Green (Eds.) Perception and Motor Control in Birds An Ecological Approach With 129 Figures and 3 Tables Springer-Verlag Berlin Heidelberg New York London Paris Tokyo Hong Kong Barcelona Budapest Dr. MARK N.O. DAVIES University College London Department of Psychology Gower St London, WClE 6BT UK Dr. PATRICK R. GREEN University of Nottingham Department of Psychology Nottingham NG72RD UK ISBN-13:978-3-642-75871-3 e-ISBN-13:978-3-642-75869-0 DOl: 10.1007/978-3-642-75869-0 Library of Congress Cataloging-in-Publication Data. Davies, Mark N.O., 1960- Perception and motor control in birds: an ecological approach / Mark N.O. Davies, Patrick R. Green. p. em. Includes bibliographical references and index. ISBN-13:978-3-642-75871-3 1. Birds - Sense organs. 2. Birds - Behavior. 3. Perceptual-motor processes. I. Green, Patrick R. II. Title. QL698.D22 1994 598.2'18 - dc20 93-33022 This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilm or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9,1965, in its current version, and permission for use must always be obtained from Springer-Verlag. Violations are liable for prosecution under the German Copyright Law. © Springer-Verlag Berlin Heidelberg 1994 Softcover reprint of the hardcover 1st edition 1994 The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. Typesetting: Macmillan India Ltd., Bangalore -25 31/3145/SPS - 5 4 3 2 1 0 - Printed on acid-free paper Preface The scope of the chapters in this book will seem at once broad and narrow. They encompass topics in both perception and motor organization, but are restricted to just one group of animals. Our reason for bringing the areas of perception and motor control together is to make the case that each should be considered in the context of the other, rather than separately. Psychologists will recognize the provenance of this view in Gibsonian or "ecological" theory, and our subtitle reflects this influence on our selection of research topics. Many features of "ecological psychology" are familiar to biologists as principles which should underlie any good biology - the analysis of structure and function in the context of the whole animal and its ecological niche - and these principles guide all the research described in this book. By restricting the scope of the book to research on birds, we hope to provide enough points of contact between different areas of research to convince readers that fruitful interactions between them are a practical possibility. Such topics as the control of flight manoeuvres or pecking provide concrete examples of how problems in perception and motor organization can be treated together. The chapters fall into three sections, and each begins with a short introduction drawing out some of the links between the chapters it contains. Section I contains reviews of recent research on a variety of sensory and perceptual processes in birds, which all involve subtle analyses of the relationships between species' perceptual mechanisms and their ecology and behaviour. Chapters in Section II describe research using a variety of methods - behavioural, neurophysio logical, anatomical and comparative - but all dealing with the common problem of understanding how the activities of large numbers of muscles are co-ordinated to generate adaptive behavi our. In Section III, chapters are concerned with a range of ap proaches to analyzing the links between perceptual and motor processes, through cybernetic modelling, neurophysiological analysis, or behavioural methods. We are grateful to Dr Dieter Czeschlik and his colleagues at Springer-Verlag for their support and advice throughout the VI Preface planning and preparation of this book. We would also like to thank Carlos Martinoya, who was prevented by illness from contributing a chapter, for his advice on the selection of topics. The compilation and editing of the chapters was made far easier by using manuscripts on disc, and we thank Karen Sherlock and Howard Martin for their help in handling different disc and document formats. We are also grateful to Charlotte Dewey for handling all our editorial corres pondence so efficiently. Nottingham, UK MARK N.O. DAVIES January 1994 PATRICK R. GREEN Contents Introduction to Section I . . . . . . . . . . . . . . . . . . . . . . 1 1 Form and Function in the Optical Structure of Bird Eyes G.R. Martin (With 9 Figures) . . . . . . . . . . . . . . 5 1.1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . 5 1.2 The Bases of Diversity in Avian Eye Structure . . . . 6 1.3 Quantitative Descriptions of Eye Structures and Their Properties . . . . . . . . . . . . . . . . . . . . . . 7 1.4 Interpretations of Diversity. . . . . . . . . . . . . . . . 8 1.4.1 Shape and Size of Eyes. . . . . . . . . . . . . . . 9 1.4.2 The Optical Design of Eyes . . . . . . . . . . . . 12 1.5 The Role of the Iris . . . . . . . . . . . . . . . . . . .. 16 1.5.1 Pupil Size and Image Brightness. . . . . . . . . 16 1.5.2 Pupil Size and Image Quality. . . . . . . . . . . 18 1.5.3 Pupil Size and Depth of Field. . . . . . . . . .. 19 1.6 Visual Fields . . . . . . . . . . . . . . . . . . . . . . .. 23 1.6.1 Monocular Fields. . . . . . . . . . . . . . . . .. 23 1.6.2 Binocular and Panoramic Fields . . . . . . . .. 24 1.6.3 Visual Fields and Amphibious Habits . . . . .. 28 1.7 Concluding Remarks . . . . . . . . . . . . . . . . . .. 30 References. . . . . . . . . . . . . . . . . . . . . . . . .. 30 2 Functional Accommodation in Birds F. Schaeffel (With 10 Figures) . . . . 35 2.1 The Power and Precision of Accommodation as a Distance Cue . . . . . . . . . . . . . . . . . . . . . . .. 35 2.2 A Technique to Measure Accommodation in Unrestrained, Alert Birds ................ . 37 2.3 Mechanisms of Accommodation in Terrestrial Birds. 39 2.3.1 Speed of Accommodation ............ . 41 2.3.2 Coupled and Uncoupled Accommodation and the Convergence of Information. . . . . . . . . . . . . 43 2.4 Visual Guidance of Pecking Behaviour ........ . 43 VIII Contents 2.5 Lower Field Myopia: an Adaptation That "Keeps the Ground in Focus"? ............ . 45 2.6 The Role of Accommodation in Judging Distances. 48 References. . . . . . . . . . . . . . . . . . . . . . . . . . 51 3 Binocular Depth Perception S.A. McFadden (With 5 Figures) 54 3.1 Introduction . . . . . . . . . . . . 54 3.2 What Exactly is Stereopsis? . . . 56 3.2.1 Retinal Disparity and Stereopsis .. 56 3.2.2 Types of Stereopsis . . . . . . . . . . 57 3.3 Stereopsis in Birds. . . . . . . . . . . . .. 58 3.3.1 Neural Mechanisms for Local Stereopsis in Birds ................... . 59 33.2 Behavioural Tests of Stereopsis in Birds. 59 3.4 Binocular Vision and the Oculomotor System in Birds .................... . 62 3.4.1 The Position of the Binocular Field 62 3.4.2 The Visual Trident in Birds . . . 63 3.4.3 Binocular Fixation and Fusion. . . 64 3.4.4 Vergence Eye Movements ..... . 65 3.4.5 Stereoscopic Limits Imposed Through the Oculomotor System. . . . . . . . . . . . 66 3.5 Role of Binocular Vision in the Guidance of Avian Behaviour. . . . . . . . . . . . . . . . . 67 3.5.1 Guidance of the Peck Movement. . . . 68 3.5.2 Dependence of Behaviour on the Frame of Reference . 69 3.6 Conclusions. 70 References. . . . 70 4 Sound Cues to Distance: The Perception of Range P.K. McGregor (With 5 Figures) 74 4.1 Introduction . 74 4.2 Why Range? ........... . 74 4.3 Ranging Cues. . . . . . . . . . . . 75 4.4 The Experimental Evidence for Ranging Ability 79 4.5 Mechanisms of Degradation Perception . 84 4.6 Ranging and Honesty. . . . . . . . . . . . . . .. 87 4.7 Some Developments of Ranging Studies . . . . . 89 4.7.1 Ranging as a Component of Other Signalling Behaviour ........ . 89 4.7.2 Resolution of Ranging ............. . 90 Contents IX 4.8 Conclusions. 91 References. . 91 5 Av ian Orientation: Multiple Sensory Cues and the Advantage of Redundancy R. Wiltschko and W. Wiltschko (With 13 Figures) . 95 5.1 Theoretical Considerations. . . . . . . . . . . . 95 5.2 Compass Mechanisms and Their Interrelation . . .. 97 5.2.1 The Magnetic Compass of Birds . . . . . . . .. 97 5.2.2 The Interrelation Between Magnetic Compass and Sun Compass. . . . . . . . . . . 99 5.2.3 Directional Orientation at Night. . . . . . . 102 5.2.4 Integrating Directional Orientation . . . . . 104 5.3 Mechanism for Determining the Home Direction. 104 5.3.1 Navigation by Route-Specific Information . 105 5.3.2 Site-Specific Information ~ the Navigational "Map". . . . . . . . . . . . . . . . . . . . . . . 107 5.3.3 Different Strategies Supplement Each Other . 110 5.4 Determining the Migratory Direction. . . . . . . . . 111 5.4.1 Reference Systems for the Migratory Direction. 112 5.4.2 The Interrelation Between Celestial Rotation and the Magnetic Field During Ontogeny. 113 5.5 Conclusion 115 References. . . . . . . . . . . . . . . . . . . . . 116 Introduction to Section II. . . . . . . . . . . . . . . . . . 121 6 Neuroembryology of Motor Behaviour in Birds A. Bekoff .............. . 125 6.1 Introduction . . . . . . . . . . . . . 125 6.2 The Environment Within the Egg. 126 6.3 Embryonic Motor Behaviours ... 126 6.3.1 Type I Embryonic Motility . 127 6.3.2 Type II and Type III Embryonic Motility 128 6.4 Role of Sensory Information During Ongoing Embryonic Behaviours .............. . 129 6.4.1 What Sensory Information Is Available? . 129 6.4.2 How Is Sensory Information Used? .... 130 6.5 Role of Sensory Input at Transitions in Behaviour. 131 6.6 Role of Prior Sensory Input in Development of Later Behaviours. 132 6.7 Conclusions. 132 References. . . . . 133 x Contents 7 Pre- and Postnatal Development of Wing-Flapping and Flight in Birds: Embryological, Comparative and Evolutionary Perspectives R.R. Provine (With 7 Figures) . . . . . . 135 7.1 Introduction. . . . . . . . . . . . . . . . 135 7.2 Prenatal Development of Spontaneous Wing-Flapping. . . . . . . . . . . . . . . 135 7.3 Neural Basis of Embryonic Behaviour. 137 7.4 Effect of Spontaneous Embryonic Behaviour on Muscle and Joint Development. . . . . . . . . . . . . . . . . 139 7.5 Naturally Occurring Motor Neuron Death . . . . . 139 7.6 Comparative Development of Wing-Flapping and Flight: Effects of Domestication. . . . . . . . . . . . 140 7.7 Experimental Studies of the Postnatal Development of Wing-Flapping and Flight. . . . . . . . . . . . . . .. 144 7.8 Bilateral Wing Coordination: Studies of Induced Bilateral Asymmetry. . . . . . . . . . . . . . . . 147 7.9 Development of Wing-Flapping and Flight in Dystrophic Chickens. . . . . . . . . . . . . . . . 150 7.10 Wing-Flapping in Flightless Birds: Evolutionary Insights . . . . . . . . . . . . . . . . . . . . . . 151 7.11 Centripetal Hypothesis of Neurobehavioural Evolution . 154 References. . . . . . . . . . . . . . . . . . . . . 155 8 Development of Prehensile Feeding in Ring Doves (Streptopelia risoria): Learning Under Organismic and Task Constraints J.D. Deich and P.D. Balsam (With 7 Figures). . 160 8.1 Introduction..................... 160 8.2 Thrusting and Grasping During Feeding in the Adult. . . . . . . . . . . . . . . . . . . . . . 161 8.3 Evidence for Plasticity and Skill in Adult Columbidae. . . . . . . . . . . . . . . . . . 162 8.4 The Transition from Dependent to Independent Feeding in the Ring Dove . . . . . . . . . . . . . . . 164 8.5 Development of Pecking. . . . . . . . . . . . . . . . 165 8.5.1 Behavioural Analysis of the Development of Pecking. . . . . . . . . . . . . . . . . . . . . . . 166 8.6 Behavioural Processes Underlying Development of Prehensile Feeding. . . . . . . . . . . . . . . . 171 8.7 The Viewpoint That Prehensile Feeding Is a Preorganized Response . 173 8.8 Task Analysis. . . . . . . . . . . . . . . . . . . 175 Contents XI 8.9 Summary 178 References. 179 9 Ingestive Behaviour and the Sensorimotor Control of the Jaw H.P. Zeigler, R. Bermejo and R. Bout (With 11 Figures) ............ . 182 9.1 Introduction............... 182 9.2 Ingestive Behaviour: Descriptive Analysis 183 9.3 Functional Considerations. . . . . . . . . 185 9.4 Kinematic Analysis of Ingestive Jaw Movement Patterns. . . . . . . . . . . . . 185 9.4.1 Kinematics of Drinking . . . . . . . . . . 186 9.4.2 Kinematics of Eating. . . . . . . . . . . . 187 9.5 Morphology and Myology of the Pigeon Jaw. 190 9.6 Electromyographic Analysis of Ingestive Jaw Movements. . . . . . . . . . . . . . . . . . . . . 191 9.6.1 Jaw Muscle Activity Patterns During Eating.. 191 9.6.2 Jaw Muscle Activity Patterns During Drinking 195 9.7 Response Topography and the Modulation of Jaw Movement Patterns. 196 9.8 Conclusions. 198 References. . . . . . . 198 10 Motor Organization of the Avian Head-Neck System G. Zweers, R. Bout and 1. Heidweiller (With 10 Figures) 201 10.1 Introduction........................ 201 10.2 Osteo-Muscular Design of the Avian Cervical Column 202 10.2.1 Osteology. 202 10.2.2 Arthrology . . . . . . . . . . . . . . . . . . . .. 202 10.2.3 Myology. . . . . . . . . . . . . . . . . . . . . .. 203 10.3 Design Modifications of the Avian Cervical Column. 205 10.3.1 Ligamentum Elasticum Cervicale . . . . . . .. 205 10.4 Patterning Head-Neck Movement and Motor Action 206 10.4.1 Postures: Minimal Flexion Model. 206 10.4.2 Motion: Least Motion Model . 207 10.4.3 Major Motion Principles. . . 209 10.4.4 Motor Patterns. . . . . . . . . . 212 10.5 Control of Head-Neck Movements. . 213 10.5.1 Comparator Model of Head-Neck Control. 213 10.5.2 Connections in the Central Nervous System. 214 10.5.3 Network Control. 216 10.6 Conclusions. 218 References. . . . . . . . . 219

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