Introduction to Ethology

241 Pages·1980·6.794 MB·English

by  Klaus Immelmann (auth.)

Introduction to
Ethology
Introduction to
Ethology
Klaus Immelmann
University of Bielefeld
Bielefeld, West Germany
Translated from German by
Erich Klinghammer
Purdue University
Lafayette, Indiana
PLENUM PRESS . NEW YORK AND LONDON
Library of Congress Cataloging in Publication Data
Immelmann, Klaus.
Introduction to ethology.
Translation of Einfiihrung in die Verhaltensforschung.
Includes bibliographical references and indexes.
I. Animals, Habits and behavior of. I. Title.
QL751.15213 591.51 80-15721
ISBN-I3: 978-1-4684-1056-3 e-ISBN-I3: 978-1-4684-1054-9
DOl: 10.1007/978-1-4684-1054-9
First Printing-November 1980
Second Printing-September 1983
This volume is a translation of the first edition of Einfiihrung in die Verhaltensforschung,
published by Paul Parey Verlag, Berlin and Hamburg, in 1976. It contains additional ma
terial that was included in the second German edition of the work, which was published in
1979 concurrently with the preparation of this translation.
© 1980 Plenum Press, New York
Softcover reprint of the hardcover 1st edition 1980
A Division of Plenum Publishing Corporation
227 West 17th Street, New York, N.Y. 10011
All rights reserved
No part of the English translation may be reproduced, stored in a retrieval system,
or transmitted, in any form or by any means, electronic, mechanical, photocopying,
microfilming, recording, or otherwise, without written permission from the Publisher
Preface
Ethology, the study of the biology of behavior, has grown tremendously
during the last few decades. The large number of accumulated facts is difficult
to survey, understanding and an appreciation of the ethological approach to
the study of behavior have grown, and the number of attempts at holistic
explanations for certain behavioral phenomena has increased. Because of this
development it has become more difficult to gain an overview of the field, to
keep up with new developments, and to update the subject matter by the
inclusion of new facts in the proper place. The nonspecialist is unable to
evaluate the more general statements in the popular literature, especially
when such works are aimed at a broader audience.
Hence, this book has a dual purpose: (1) to lend some order to the
dizzying array of information and thus simplify inquiry into ethology; and (2)
to present relevant facts and knowledge that will help the reader confronted
with numerous studies and articles in the ethological literature.
It is always difficult to select material for such a survey and to arrange it
in some logical manner. Hence, this volume offers but one of many possible
ways to present this material. This book is based on a course of lectures of the
same title that I have offered during the past 12 years at the universities of
Braunschweig, Berlin, and Bielefeld in West Germany. These lectures were
continually modified in response to questions that indicated what knowledge
in which particular places would enhance the understanding of certain
relationships. Thus, this book can be regarded in its organization as a sort of
stepping-stone on the path to a solution. This, I hope, will be the clearest and
simplest way to present the relationships among facts, hypotheses, and
theories.
For reasons of space, limitations existed on the selection of material. This
is why the physiological basis of behavior, which is generally offered in texts
and surveys in physiology, is treated less comprehensively than the area of
"classical" and comparative ethology. However, even here focal points had to
be chosen; thus, the treatment of topics in the currently popular style of
teaching by illustration and example is somewhat uneven. As a result, some
areas, such as aggression, where general regularities of behavior and methods
of study can be clearly demonstrated, have been presented in more detail than
have some others. The extensive area of learning has been presented in
comparatively condensed form, since numerous texts on the subject are readily
available.
v
vi PREFACE
Since this is an introduction that is primarily designed to provide the
means for a more intensive involvement with the field, the author has not
participated in the sometimes controversial discussions about basic concepts
in the various "schools" of ethology. These are pointed out in those instances
where certain kinds of explanations and theories are still being debated.
With respect to humans it must certainly be accepted that some charac
teristics of human behavior developed as the result of the evolutionary process,
and are hence comparable to corresponding aspects of the behavior of animals.
Hence, ethology is in a position to point out a certain regularity that in some
way is also applicable to humans. Such applications must, however, be made
with great care since insufficiently supported comparisons could, as has been
shown repeatedly in the past, only harm ethology in its attempts to make
more general statements about behavior, and thus to aid in a better understand
ing of human behavior as well.
Ethology is a biological science. Each behavioral characteristic of an
animal can be understood and interpreted properly only when its biological
significance, i.e., what its importance is for the particular species in its natural
environment, is known. This has not always been the case in the history of
the study of animal behavior to the extent that is necessary. The continued
reference to such ecological relationships, and to adaptations that have
developed during the course of evolution, is an additional aim of this book.
Since an introduction of this kind is also directed to readers interested in
ethology who may not have any special biological background, it was neces
sary to explain certain basic concepts that are important for an understanding
of the biological significance of a particular behavior. These are found in the
text and in footnotes. This also applies to special terms that are not properly
a part of ethology.
The scientific names of the animal species cited are given only when a
proper common name is not available.
I am indebted to the illustrator, Mr. Klaus Weigel, of the Faculty of
Biology of the University of Bielefeld, for the preparation of the drawings,
which he carried out with great care and understanding. Thanks are also due
my colleagues and assistants in Bielefeld for their stimulating ideas and
discussions during the writing of this book.
Klaus Immelmann
Bielefeld, March 1976
This English edition contains additional material that was included in
the second German edition, which appeared while the English translation was
in progress. Two new chapters, one on domestication and one dealing with
ethology and human behavior, were added. These chapters deal with research
PREFACE vii
that may lead to a better understanding of human behavior as well. However,
the facts that have been collected so far permit only limited conclusions to
date. For this reason, these chapters are of a more speculative nature than the
previous chapters. Indeed, the last chapter was added only in response to
requests for its inclusion, although there has been no change in the views
expressed on this subject in the introduction to the first edition.
Klaus Immelmann
Bielefeld, July 1980
Contents
1 Aims, Methods, and Areas of Ethology ................. 1
1.1. Definitions of Concepts .................................... 1
1.2. Descriptive Ethology ...................................... 2
1.3. Experimental Ethology ..................................... 4
1.4. Areas of Ethological Research .............................. 7
2 Basic Ethological Concepts ........................... 11
2.1. Reflexes ................................................. 11
2.2. Threshold Changes ....................................... 13
2.3. Specific Fatigue .......................................... 15
2.3.1. Action-Specific Fatigue ............................. 15
2.3.2. Stimulus-Specific Fatigue ........................... 15
2.4. Appetitive Behavior and Consummatory Acts ............... 17
2.5. Spontaneity of Behavior .................................. 19
2.6. Motivation ............................................... 22
2.7. Motivational Analysis .................................... 24
2.8. Motivational Systems; "Drives" ............................ 24
3 External Stimuli ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
3.1. Stimulus Filtering ........................................ 27
3.1.1. Peripheral Filtering ................................. 27
3.1.2. Central Filtering ................................... 29
3.2. Releasing Mechanisms .................................... 33
3.3. Key Stimuli .............................................. 34
3.4. Releasers ................................................ 34
3.5. Interspecific Releasers .................................... 37
3.6. Stimulus Summation ..................................... 38
3.7. Supernormal Releasers .................................... 39
3.8. Differential Effects of External Stimuli ..................... 41
3.9. Dependence upon External Stimuli ........................ 42
3.9.1. Inherited Movement Coordinations (Fixed Action
Patterns) .......................................... 42
3.9.2. Taxis Components .................................. 43
ix
x CONTENTS
4 Temporal and Hierarchical Organization of Behavior . . 45
4.1. Categorization of Behavior Patterns ........................ 45
4.2. Conflict Behavior ......................................... 47
4.2.1. Ambivalent Behavior ............................... 47
4.2.2. Redirected Behavior ................................ 47
4.2.3. Displacement Behavior ............................. 48
4.3. Models of Instinctive Behavioral Organization .............. 50
5 Behavioral Physiology ............................... 55
5.1. The Nervous System and Behavior ......................... 55
5.1.1. Methods ........................................... 56
5.1.2. Results ............................................ 59
5.2. Hormones and Behavior .................................. 68
5.2.1. Methods ........................................... 68
5.2.2. Influences of Hormones ............................. 70
5.2.3. Effects of Hormones ................................ 75
5.2.4. Hormone Release Controlled by Behavior ............ 79
6 Ontogeny of Behavior ................................ 81
6.1. Inborn or Acquired? ...................................... 81
6.1.1. Sources of Information .............................. 81
6.1.2. Recognition of Inborn Behavior Patterns .............. 83
6.1.3. What Is Inborn? .................................... 85
6.2. Maturation of Behavior Patterns ........................... 88
7 Learning............................................ 91
7.1. The Biological Significance of Individually Acquired
Information .............................................. 91
7.2. Sensitive Phases in Learning .............................. 92
7.3. Learning Predispositions .................................. 94
7.4. Learning Processes ....................................... 97
7.4.1. Habituation ........................................ 97
7.4.2. Classical Conditioning .............................. 98
7.4.3. Instrumental Conditioning .......................... 98
7.4.4. Play Behavior ...................................... 99
7.4.5. Imitation, Social Facilitation ....................... 102
7.4.6. Insight Learning ................................... 102
7.4.7. Imprinting ........................................ 104
7.5. Behavioral Traditions .................................... 110
7.6. Tool Using .............................................. 113
CONTENTS xi
8 Social Behavior ..................................... 115
8.1. Fighting Behavior ....................................... 115
8.1.1. Definition ........................................ 115
8.1.2. Biological Significance ............................ 116
8.1.3. Threat and Intimidation Displays .............. , .... 117
8.1.4. Submission and Appeasement Gestures ............. 119
8.1.5. Territoriality ...................................... 119
8.1.6. Individual Distance ................................ 129
8.1.7. Ritualized Fights .................................. 130
8.1.8. Damaging Fights and Killing of Conspecifics ........ 131
8.1.9. Motivational Aspects ............................... 133
8.2. Sexual Behavior ......................................... 138
8.2.1. Biological Significance ............................ 138
8.2.2. Function of Sexual Behavior ....................... 140
8.2.3. Pair-Bonding Behavior ............................. 152
8.2.4. Forms of Heterosexual Bonds ....................... 153
8.2.5. Mechanisms of Maintaining Bonds ................. 154
8.2.6. Motivational Aspects .............................. 156
8.3. Behavior in Care of Young ............................... 157
8.3.1. Functions in Care of Young ........................ 157
8.3.2. Formation of Families ............................. 159
8.3.3. Behavioral Adaptations with Respect to Care
of Young ......................................... 160
8.4. Behavior of Groups ...................................... 162
8.4.1. Biological Function of Social Groups ............... 162
8.4.2. Social Structure ................................... 166
8.4.3. Altruism .......................................... 167
8.4.4. Types of Groups ................................... 170
8.4.5. Origin of Groups .................................. 172
8.4.6. Mechanisms of Group Formation ................... 173
8.4.7. Motivational Aspects .............................. 176
9 Behavior Genetics .................................. 179
9.1. Methods ................................................ 179
9.2. Behavior of Hybrids ..................................... 180
9.3. Crossbreeding Experiments .......................... " ... 182
9.4. Conclusions ............................................. 184
10 Phylogenetic Development of Behavior ............... 185
10.1. Methods ................................................ 185
10.2. Problems of Homology ................................... 186
10.3. Species Comparisons .................................... 187
10.4. Studies in the Ontogeny of Behavior ...................... 188
xii CONTENTS
10.5. Behavioral Rudiments ................................... 189
10.6. Ritualization ............................................ 190
10.7. Convergence of Behavior Patterns ......................... 192
10.8. Conclusions ............................................. 194
11 Influence of Domestication on Behavior .............. 197
11.1. Problems of Definition ................................... 197
11.2. Characteristics of Domestication .......................... 198
11.2.1. Changes in the Readiness to Act ................... 198
11.2.2. Changes in Innate Releasing Mechanisms .......... 199
11.3. Causes of Changes during Domestication .................. 200
11.4. Conclusions ............................................. 202
12 Ethology and Psychology ............................ 203
12.1. Introduction ............................................ 203
12.2. Level of Comparison ..................................... 203
12.3. Application ............................................. 204
12.3.1. Animal Experiments as a Model ................... 204
12.3.2. Research in Human Ethology ...................... 206
12.4. Phylogenetic Preprogramming of Human Behavior ......... 207
12.4.1. Examples ........................................ 207
12.4.2. Research in Behavior Genetics in Humans .......... 208
12.5. Prospects for the Future .................................. 210
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
Figure Credits .......................................... 217
Author Index ............................. , ............. 219
Animal Index .......................................... 221
Subject Index ....................., . . . . . . . . . . . . . . . . . . . . . . . .. 227