Human Ethology

Human Ethology

IRENAUS EIBL-EIBESFELDT

Routledge Taylor & Francis Group

LONDON AND NEW YORK First published 1989 by Transaction Publishers

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Copyright © 1989 by Irenaus Eibl-Eibesfeldt.

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Library of Congress Catalog Number: 2006048043

Library of Congress Cataloging-in-Publication Data

Eibl-Eibesfeldt, Irenaus. [Biologie des menschlichen Verhaltens. English] Human ethology / Irenaus Eible-Ebesfeldt. p. cm. Originally published: New York: Aldine De Gruyter, c1989. (Foundations of human behavior) Includes bibliographical references (p. ) and indexes. ISBN 978-0-202-30970-5 (alk. paper) 1. Genetic psychology. 2. Human behavior. 3. Psychobiology. 4. Sociobiology. I. Title.

BF70 l.E4313 2007 304.5—dc22 006048043 2006048043

ISBN 13: 978-0-202-30970-5 (pbk) Contents

Preface ix Acknowledgments xiii Mirror-Image Illustrations xv

1 OBJECTIVES AND THEORETICAL BASES OF HUMAN ETHOLOGY 2

1.1. Inquiry Formulation and Definition 3 Summary 1.1 6 1.2. Phylogenetic and Cultural Adaptation 7 Summary 1.2 16

BASIC CONCEPTS OF ETHOLOGY 18

2.1. The Concept of the "Innate" 19 Summary 2.1 23 2.2. Phylogenetic Adaptations in Behavior 23 Summary 2.2 84 2.3. Decoupling of Actions from Drives and Conscious Self-Control: The Neuroethology of Human Freedom 86 Summary 2.3 90 2.4. The Units of Selection—A Critical Evaluation of Sociobiology 90 Summary 2.4 102

METHODOLOGY 104

3.1. Gestalt Perception and Cognition 105 Summary 3.1 107 3.2. Methods of Data Gathering, Observation, and Description 107 Summary 3.2 112 3.3. Film and Sound Documentation 112 Summary 3.3 132 3.4. The Comparative Approach 132 Summary 3.4 147 3.5. Quantifying Ethology 148 Summary 3.5 161

3.6. Models 161 Summary 3.6 165

4 SOCIAL BEHAVIOR 166

4.1. Origins of Sociability 167 Summary 4.1 169 4.2. The Ambivalence of Approach and Withdrawal in Human Interactions 170 Summary 4.2 184 4.3. The Human Family as the Nucleus of Society 184 Summary 4.3 234 4.4. Family and Marriage 235 Summary 4.4 238 4.5. Pair Formation, Courtship, Sexual Love 238 Summary 4.5 260 4.6. Incest Taboos and Incest Avoidance 261 Summary 4.6 264 4.7. Sex Roles and Their Differentiation 265 Summary 4.7 288 4.8. The Individualized Group: Family, Kin, and Alliances 289 Summary 4.8 296 4.9. Rank Order, Dominance 297 Summary 4.9 314 4.10. Group Identity Maintenance 314 Summary 4.10 320 4.11. Territoriality 321 Summary 4.11 339 4.12. Origin and Social Function of Possession 339 Summary 4.12 358

5 INTRASPECIFIC AGGRESSION: CONFLICT AND WAR 360

5.1. Concepts 361 Summary 5.1 364 5.2. Aggression Theories 365 vi Summary 5.2 386 5.3. Functional Aspects of Aggressive Behavior 391 Summary 5.3 396 5.4. Socialization of Aggressive Behavior 396 Summary 5.4 399 5.5. Duels 399 Summary 5.5 402 5.6. Intergroup Aggression—War 402 Summary 5.6 421

6 COMMUNICATION 424

6.1. Olfactory Communication 426 Summary 6.1 429 6.2. Tactile Communication 430

6.3. Visual Communication 437 Summary 6.3 492 6.4. Interaction Strategies—The Universal Grammar of Human Social Behavior 493 Summary 6.4 522 6.5. On the Ethology of Verbal Communication 523 Summary 6.5 546

7 BEHAVIOR DEVELOPMENT (ONTOGENY) 548

7.1. Developmental Theories 549 Summary 7.1 580 7.2. Curiosity and Play 580 Summary 7.2 593 7.3. The Development of Interpersonal Relationships 594

7.4. Child Groups—Child Culture 600 Summary 7.4 605

8 MAN AND HIS HABITAT: ECOLOGICAL CONSIDERATIONS 606

8.1. Ecotype Homo sapiens: Hominization and Behavior 607 Summary 8.1 616 8.2. From the Individualized Society to Industrial Society 616 Summary 8.2 628 8.3. On the Ethology of Settlement and Residence 629 Summary 8.3 648 vii 8.4. Social Order and Human Behavior 649 Summary 8.4 662 8.5. Concluding Thoughts 663

9 THE BEAUTIFUL AND TRUE: THE ETHOLOGICAL CONTRIBUTION TO AESTHETICS 664

9.1. Aesthetics and Fine Arts 665 Summary 9.1 673 9.2. Species-Specific Biases of Perception of Aesthetic Relevance 673 Summary 9.2 677 9.3. Art as Communication 677 Summary 9.3 685 9.4. Cultural Manifestations: An Observation on Style and Stylization 685 Summary 9.4 690 9.5. On the Ethology of Music, Dance, and Poetry 690 Summary 9.5 702

10 BIOLOGY'S CONTRIBUTION TO ETHICS 704

Summary 10 717 Conclusion 718

Bibliography 720 Film Publications 789 Author Index 805 Subject Index 824

viii Preface

With the discovery of conditioned reflexes by I. P. Pavlov, the possibilities for experimenting, following the example set by the classical, exact sciences, were made available to the behavioral sciences. Many psychologists hoped that the component parts of behavior had also been found from which the entire, multi- faceted cosmos of behavior could then be constructed. An experimentally oriented psychology subsequently developed including the influential school of behaviorism. Motivated by pedagogical idealism, psychologists indulged in the hope that ed­ ucation could achieve every desired result. Among behaviorists, the Lockean conviction that humans come into the world with "clean slates" crystallized into a doctrine. It seems certain that this conceptual approach restricted research in­ terest. Although behaviorists carefully investigated the various aspects of learning behavior, and extracted some fundamental laws of learning, their theoretical fix­ ation prevented their perceiving other possible approaches. The possibility that some aspects of behavior might, through phylogenetic adaptation, be genetically programmed was not even considered. Anyone referring to a possible antecedent innate human nature was, until recently, dismissed as a biological determinist or reductionist. As late as the 1970s, the majority of behaviorists and sociologists viewed humans as primarily passive beings subject to the formative influences of their surroundings. It was believed that through rewards and punishments almost all human behavior could be "conditioned" (B. F. Skinner, 1938, 1971). Such an extreme environmentalist approach would consign all behavior to learning, relegate all power to the educators, and ultimately, legitimize them to establish behavior norms as well. Parallel to the development of behaviorism in psychology there developed in anthropology a pronounced cultural relativism (discussed critically by W. Rudolph, 1968) according to which culture is a construct on which the laws of biology have no influence. This school of thought is traceable to Franz Boas (1911, 1928, 1938) and its tenets must have reflected the spirit of the times since leading anthropologists acknowledged them (A. L. Kroeber, 1915; M. J. Her- skovits, 1950, among others). Margaret Mead can be regarded as one of the victims of this doctrinal teaching; she was so convinced of the relativism of cultures that she saw and recorded only that which fit those theories. Her reports on Samoa, so often cited, could not stand the test of critical review (D. Freeman, 1983). The decisive impetus for revising this extreme environmental approach came from behavioral biology (ethology). The work of Konrad Lorenz (1935, 1937, 1961) and Nikolaas Tinbergen (1948, 1951) clarified the notion of instinct, a concept which, until this time, was a philosophical speculation of the vitalists. There were, of course, exceptions, such as W. James (1890) who defined "instinct" in fairly modern terms, as capacities of organisms based on an innate neuronal organization. Lorenz and Tinbergen demonstrated that phylogenetic adaptations determine be­ havior in well-defined ways, and also voiced the opinion that comparable phy­ logenetic adaptations influence human behavior. By introducing the dimension of phylogeny into the science of behavior their work incorporated Charles Darwin's findings; behaviorists had apparently forgotten to consider the evolutionary aspects of behavior in their theories. About the same time that Lorenz formulated his theories, Erich von Hoist (1935, 1939) refuted the classical reflex theory, according to which all behavior is a re­ sponse to external stimuli. Ethologists can look back over the last decade with a certain sense of satis­ faction, for within a short span of 10 years, six biological researchers have received the Nobel Prize: Konrad Lorenz, Niko Tinbergen, and Karl von Frisch in 1973 (as founders of ethology) and Roger W. Sperry, David Hubel, and Torsten Wiesel in 1981 (as pioneers of neuroethology).1 The discovery that even the behavior of higher vertebrates is preprogrammed in well-definable ways through phylogenetic adaptation has initiated a process of reconsideration in the human sciences, to date so heavily influenced by behav­ iorism. In 1971, R. Sperry wrote that the new discoveries on the self-organization of the nervous system up to the point of functional maturity has hardly been noticed in disciplines beyond the fields of biology and ethology. Currently, however, the biological approach is well acknowledged by most fields of science dealing with humans. Konrad Lorenz indicated in his earlier works that the findings of animal behavior research could contribute to the understanding of human behavior. He devoted a number of important chapters to human behavior in Die angeborenen Formen moglicher Erfahrung (1943) and in 1950 he designated as one of the most important objectives of ethology the testing of the hypotheses derived from the study of animal behavior through the study of human behavior. Clearly, this objective consists of more than simply applying to humans the findings and models of animal behavior research. Ethological research on man was needed, and it truly began in the mid-1960s. I have been able to contribute to the development of this field, bringing to it a considerable background of prac­ tical field and laboratory experience in animal behavior on a broad comparative basis. As a student, beginning in 1946, I had the opportunity to study the behavior of amphibians and mammals at the Wilhelminenberg Biological Station (Vienna). A badger that I hand-raised helped me learn a great deal about animal play behavior. It occurred to me that during the course of its play behavior the badger freely combined behavior patterns from a number of different functional systems. Indeed, in its play behavior we can observe the first manifestations of that freedom to perform derived from voluntary motor control and the ability to detach behavior patterns from emotions which also lies at the root of man's behavioral freedom (I. Eibl-Eibesfeldt, 1950). In 1949-1950, I joined Lorenz at the newly established Research Center for Behavioral Physiology of the Max Planck Society. In addition to my interest in behavioral development, questions regarding communication caught my attention. How do signals guide the course of social interactions? How do they originate, and how does the understanding of signals develop? In this area I was particularly concerned with the ritualization of aggressive behavior (I. Eibl-Eibesfeldt, 1955a, 1959).

x 'Erich von Hoist is only absent from this list since his death took him from us too soon. On a 10-month expedition led by Hans Hass to the Caribbean Sea and the Galapagos Islands, I had the opportunity to study the cleaning symbiosis of fishes and other interspecific relationships of coral fishes (I. Eibl-Eibesfeldt, 1955b). I also observed the ritualized fights of marine iguanas, and became acquainted with adaptive radiation from first-hand observation of Darwin's finches. On another diving expedition (1957-1958), again with Hans Hass, I spend 1 year in the Maldive and Nicobar Islands. My main interest was the behavior and ecology of coral fishes. I assembled a large collection of these fishes and learned much about sys- tematics and functional morphology. On these two research voyages21 sharpened my appreciation for interrelationships, especially those concerning ecological fac­ tors. After returning from this expedition I became increasingly involved in the nature- nurture debate. I experimentally refuted a number of assertions about the ontogeny of certain behavior patterns and showed how innate and acquired elements in mammalian behavior intercalate (I. Eibl-Eibesfeldt, 1963). In the early 1960s I began to include human behavior in my research. Through investigations with children deaf and blind from birth, I was able to demonstrate phylogenetic adaptations in human behavior (I. Eibl-Eibesfeldt, 1973). Between 1963 and 1965 I began with the documentation of unstaged ("natural") human social interactions. The impulse came from my friend Hans Hass, who provided the technical requisites through the development of a specialized reflex lens (p. 112). We tested this technique in 1964 in Africa and in 1965 on a world-wide expedition (I. Eibl-Eibesfeldt and H. Hass, 1966, 1967). From this experience, we developed a program of cross-cultural documentation of human social inter­ actions and rituals on film and tape. For the longitudinal studies, I selected a number of cultures with different sub­ sistence strategies and which, within certain limitations, could also serve as models for different stages of cultural evolution-from Paleolithic hunters and gatherers to farming societies. In 1970, I was entrusted with an Institute for Human Ethology within the organization of the Max Planck Society. Cross-cultural documentation formed a principal part of our efforts.3 We in­ vestigated phylogenetic and cultural adaptations, the development of rituals and their function, and, in general, the behavior patterns of communication. In studying strategies of social interactions, I found that certain verbal and nonverbal inter­ actions are structured according to the same universal rules. The exploration of this universal system of rules that structure social behavior is at present one key aspect of our research efforts. To achieve this goal, we are working in conjunction with linguists. In 1967, I concluded my animal ethological studies with the pub­ lication of Ethology: The Biology of Behavior.4

2 A summary is found in the more recently published books Galapagos (7th ed., Piper, 1984) and Die Maldiven (Piper, 1982). 3I spent over 4 years in the field gathering comparative cultural documentation: 1568 days at the target sites (excluding travel time). During this time I produced approximately 200 km of 16 mm film. Many of my findings are based on this large body of material. My zoological field experience comprises 1488 days—in total over 8 years of expedition ex­ perience. 4Grundrip der vergleichenden Verhaltensforschung, Piper, Munich. The 6th edition of this work appeared in 1980. xi As a supplement to my textbook on animal behavior, this new text on human ethology is based on the latter half of my life's work.5 The findings on animal ethology are only occasionally cited in this work and then only to the extent that they contribute to the understanding of ethological concepts. I hope that this first text on human ethology presents itself as a unified work, even though not every area could be treated with equal depth. For example, a branch of ethology has developed in the past decades which places particular emphasis on ecology and population genetics. This field, known as sociobiology, has enriched discussion beyond the boundaries of behavioral biology through its stimulating, and often provocative, theses. I will discuss some of its contributions when appropriate in this volume, but, of course, cannot cover this new field in depth. After vigorous debates between behaviorists, anthropologists, and sociologists, we have entered a period of exchange of thoughts and a mutual approach, which in many instances has led to cooperative projects of researchers from different disciplines. This work offers a biological point of view for discussion and includes data from my own cross-cultural work and research from the staff of our institute. It confirms, above all else, the astonishing unity of mankind and paints a basically positive picture of how we are moved by the same passions, jealousies, friendliness, and active curiosity. It is hoped that it encourages further discussion and thought across traditional academic boundaries. Before turning to the text, I would like to add that the need to understand our­ selves has never been so great as it is today. An ideologically torn humanity strug­ gles for its survival. Our species, who is in a position to send satellites to Mars and Venus and to broadcast photographs of Saturn and Jupiter from space, is helpless when confronted with its social problems. It does not know how it should compensate its workers, and it experiments with various economic systems, con­ stitutions, and forms of government. It struggles for freedom and stumbles into newer conflicts. Population growth is apparently completely out of hand, and at the same time many resources are being depleted. With the continuation of the present exponential growth of the world population looms the catastrophe of a global breakdown of populations. In his book Mutations of Mankind, Pierre Ber- taux (1963) writes: "Assessing contemporary human existence as a biological phenomenon is not only permissible but mandatory. Anyone rejecting this approach fails to recognize the significance of our existence and thus the extent of human responsibility. ..." Our hope is to develop an ethic for human survival through gaining insight into the biological processes governing our lives. We must consider our existence rationally in order to understand it, but certainly not with cold, calculating reason but with the warm feeling of a heart concerned for the welfare of later generations. May this book contribute to such an outlook.

Professor Dr. Irendus Eibl-Eibesfeldt

5My prior books on this subject include: Love and Hate. The Natural History of Ele­ mentary Behavioral Patterns (10th ed., 1982); The !Ko Bushman Society. Group Formation and Aggression Control in a Hunting and Gathering People (1973); Pre-programmed Man xii (4th ed., 1982); New Pathways of Human Research (2nd ed., 1984). Acknowledgments

In the past 20 years, I have spent nearly one-fourth of my time in societies whose members still follow traditional living patterns. Few of them will read the lines in this book, but perhaps some of their children will, which would make me happy indeed. It would be one of the ways in which my gratitude to them could be expressed—for their hospitality and for what they taught me. I am especially indebted to my wife Lorle and my children Bernolf and Roswitha for their courage and good spirits during the extended absences necessitated by my expeditions and field studies. Konrad Lorenz, whom I have regarded as my fatherly friend for the past 40 years, promoted my work in many ways. We engaged in endless conversations and he was open to my ideas. His openness always remained a model for me. I also express my special thanks to another close friend: Hans Hass, who, 35 years ago, led me through the reefs of the Caribbean Sea, and who developed in the early 1960s the reflex lens, the key to our cross-cultural documentary work. The unforgettable diving expeditions on the "Xarifa" were followed by no less fas­ cinating expeditions to human communities where we tested new methods of doc­ umentation. Some of the thoughts in this volume were developed on these shared journeys. We have also worked together as editors of the human ethology film archives of the Max Planck Society. I thank my expedition companions of many years. Quite aside from their as­ sistance, they have enhanced, with their human presence, the pleasure of con­ ducting field work. Thus, I thank from my heart: Kuno Budack, Volker Heeschen, Hans Joachim Heinz, Harald Herzog, Dieter Heunemann, Wulf Schiefenhovel, and Heide Sbreszny-Klein. I thank my co-workers at the Institute and my many friends in Europe and overseas, especially my colleagues Jiirgen Aschoff, Ingrid Bell-Krannhals, Norbert Bischof, William Charlesworth, Mario von Cranach, De­ rek Freeman, Elke and Karl-Friedrich Fuhrmeister, Inga Goetz, Karl Grammer, Anna Guggenberger, Bernhard Hassenstein, Klaus Helfrich, Eckhardt Hess, Les Hiatt, Barbara Hold, Franz Huber, Hermann Kacher, Erich Klinghammer, Gerd Koch, Otto Koenig, Renate Krell, Cornelia von Laue-Canady, Ernie Reese, Mar­ garet and Wolfgang Schleidt, Reinhard Schropp, Gunter Senft, Christa Siitterlin, Pauline Wiessner-Larsen and Wolfgang Wickler. Special thanks are due to the governments of Australia, Botswana, Indonesia, Namibia, Papua New Guinea, the Philippines, South Africa, and Venezuela, who fostered our work through granting our work permits and who also supported us to the best of their ability. In the field we had continuous help from the local missions. Individual examples include the help of the Salesian Mission in Vene­ zuela; in the early 1970s, the New Tribes Mission in Serra Parima was also a friendly host. Missions in Irian Jaya and Papua New Guinea also helped with their hospitality, by providing transportation assistance and a well-developed infra­ structure. I would like to mention especially the assistance of the German and Austrian embassies, without whom many plans would not have been realized. The scientific organizations of the Federal Republic of Germany have consis­ tently fostered our work most generously. The Max Planck Society established an independent research center for human ethology and also made possible the creation of a film documentation program. For this sustained support, I thank the Max Planck Society and its President. Without the financial assistance of the German Research Society (Deutsche Forschungsgemeinschaft) a number of important interdisciplinary projects would not have been carried out. Individual projects were also supported by the Fritz von Thyssen Fund, the Werner von Reimers Fund, and the Maison de l'Homme. The A. von Gwinner Fund provided me with the decisive initial assistance during the early years of my human ethology field research. I sincerely appreciate the support of Inter Nationes for subsidizing this trans­ lation. It is, furthermore, my great pleasure to thank the Institute for Scientific Film in Gottingen and its director, Hans Karl Galle, for many years of fruitful coop­ eration. My heartfelt thanks to Pauline Wiessner-Larsen and Anette Heunemann for their faithful translation of this book. In addition, I wish to express my deepest appreciation to William Charlesworth, Tom Pitcairn, Anne Rasa, and Wulf Schie- fenhovel, for consulting and assisting with specific sections of the text. Last, but not least, I want to thank Melvin Konner, and the Editors of the Foundation of Human Behavior series for their interest in my work. I also wish to thank the publisher, and, in particular, Ms. Sheila Johnston, who invested so much work in the final editing of the book.

xiv Mirror-1 mage Illustrations

The illustrations listed below appear as mirror images when printed since the film sequences from which they were copied were taken with the mirror lens.

2.10, 2.11, 2.15, 2.17, 3.18, 3.19, 3.20c, 3.21A, 3.21B, 3.21C, 3.22, 3.23, 4.1, 4.2, 4.3, 4.5, 4.24, 4.25, 4.26, 4.27, 4.28, 4.30, 4.31, 4.32, 4.33, 4.34, 4.36b,d,e, 4.37, 4.38, 4.40A, 4.40B, 4.41A, 4.41B, 4.46, 4.48, 4.49, 4.50B, 4.67b, 4.68, 4.69, 4.70, 4.77, 4.78, 4.80, 4.81, 4.82, 4.83, 5.1, 5.3, 5.4, 5.5, 5.6, 5.7, 5.8, 5.9, 5.12, 5.13, 5.15, 5.16, 5.18, 5.23, 5.24, 5.26, 5.29, 6.5, 6.13, 6.19, 6.22, 6.23, 6.24A, 6.33, 6.34, 6.35, 6.36, 6.37, 6.38, 6.39a-c, 6.40, 6.45A, 6.45B, 6.47, 6.48, 6.49, 6.54, 6.55b, 6.56, 6.62, 6.64, 6.65, 6.70, 6.71, 6.72, 6.73, 6.74, 6.76, 7.2, 7.3, 7.4, 7.5, 7.6, 7.8, 7.9, 7.10, 7.11, 7.12, 7.13, 7.14A, 7.14B, 7.15, 7.17b, 7.20, 7.21, 7.22, 7.24B, 7.25, 7.29, 7.30, 7.31, 7.32, 7.33, 7.34. FOUNDATIONS OF HUMAN BEHAVIOR An Aldine de Gruyter Series of Texts and Monographs

Series Editors Sarah Blaffer Hrdy, University of California, Davis Richard W. Wrangham, University of Michigan

Human Ethology. 1989 Irendus Eibl-Eibesfeldt Evolutionary History of the "Robust" Australopithecines. 1988 Frederick E. Grine, Editor Early Hominid Activities at Olduvai. 1988 Richard Potts Homicide. 1988 Martin Daly and Margo Wilson The Biology of Moral Systems. 1987 Richard D. Alexander Child Abuse and Neglect: Biosocial Dimensions. 1987 Richard J. Gelles and Jane B. Lancaster, Editors Parenting Across the Life Span: Biosocial Dimensions. 1987 Jane B. Lancaster, Jeanne Altmann, Alice S. Rossi, and Lonnie R. Sherrod, Eds. Primate Evolution and Human Origins. 1987 Russell L. Ciochon and John G. Fleagle, Editors Human Birth: An Evolutionary Perspective. 1987 Wenda R. Trevathan School-Age Pregnancy and Parenthood: Biosocial Dimensions. 1986 Jane B. Lancaster and Beatrix A. Hamburg, Editors Despotism and Differential Reproduction: A Darwinian View of History. 1986 Laura L. Betzig Sex and Friendship in Baboons. 1985 Barbara Boardman Smuts Infanticide: Comparative and Evolutionary Perspectives. 1984 Glenn Hausfater and Sarah Blaffer Hrdy, Editors Navajo Infancy: An Ethological Study of Child Development. 1983 James S. C his holm

In Preparation Inujjuamiut Foraging Strategies Eric Alden Smith Human Reproductive Ecology James W. Wood Ecology, Evolution, and Human Behavior Eric A. Smith and Bruce Winterhalder, Editors To the Memory of

My Beloved Friend and Teacher

KONRAD LORENZ (1903 - 1989) 1

Objectives and Theoretical Bases of Human Ethology The animal is taught by its organs; man teaches his and controls them. Johann Wolfgang von Goethe

Whether mankind considers himself the son of God or a successful ape will make a clear difference in his behavior towards actual facts; in both cases he will also hear very different commands from within himself. Arnold Gehlen (1970, p. 1)

1.1. Inquiry Formulation and Definition

There has never been a shortage of attempts to define man and, by interpretation, attribute meaning to his life. For millennia, priests, artists, and philosophers have considered these questions. Religious revelation has often been contrasted with the efforts to gain insight into human nature through observation and introspection, that is, through experience and with the aid of reason. To this empirical-rational approach, biology gave new impetus with the theory of evolution, a theory that shook our anthropocentric perception of the world. Through evolutionary theory, man became aware not only of his animal heritage, but also of the reality of his incompleteness—man could no longer consider himself as the final "crowning achievement" of creation, but rather, at best, as an intermediate link on the path toward a higher level of humanity. Of course, this new insight led to a new perspective of man, one of a progres­ sively developing creature, setting ever higher goals and striving toward these objectives. But, at the same time, man became painfully aware that he travels on a narrow ridge, risks toppling off, and is burdened with responsibility, in as much as he accepts his further development as his mission. If, for some, man was simply one animal species among others, a "naked ape" as Desmond Morris provocatively expressed it, others, in contrast, felt that man had progressed so far above animals in his evolution that he no longer had anything in common with them; that he had gone beyond the biological process of evolution by virtue of his culture. Man, therefore, was considered free to arrange his life in accordance with reason, without any limitations on constraints. These alternate viewpoints became unnecessarily polarized, for both approaches have validity. Biological inheritance determines human behavior, as we will show, in precisely definable parameters. But it is equally true that only man possesses a language, with which he can creatively formulate new statements and thus pass on his cultural heritage by tradition. Only Man can be defined as a cultural being even though some primates show some modest cultural beginnings. Art, reason, and responsible morality, as well as open-mindedness and adaptability, are further distinctive hu­ man traits, and no reasonable biologist doubts their special role.1 It is important, however, that we are also aware of the more primitive action and reaction patterns that determine our behavior, and to not pretend as if they did not exist. It is especially in the area of social behavior that we are less free to act than we generally assume. This is poignantly demonstrated by the astonishing

'Konrad Lorenz wrote (1971, p. 509): "Far from underestimating the differences between the previously described behavioral patterns of higher animals and human achievements, molded by reason and morality, I maintain: no one is able to appreciate the uniqueness of these specific human attributes as clearly as those who can perceive them against the back­ ground of those much more primitive actions and reactions, which we also share with the higher animals." 3 discrepancy between our ability to control the external environment and our in­ ability to shape our social life satisfactorily. At the same time that we enthusi­ astically view the space photographs of Jupiter from the Voyager space probe, we read in our newspapers about executions in Iran, terrorist acts in Ireland, and mass murder in Kampuchea. Of course we should be hopeful for our future based on our potential for responsible morality, but this is possible only when we also recognize the inherited basis of our actions and take them into consideration. In this sense biology can contribute to our enlightenment and emancipation. Human ethology can be defined as the biology of human behavior. Like its parent discipline ethology, it too is divided into specialties, since the question as to why humans behave one way rather than another can be defined and answered in different ways. If one's interest lies in the function of the underlying physiological mechanisms, then human ethology becomes allied with traditional behavioral physiology. Here we are dealing with an explanation of the proximate cause of a behavior pattern. Inquiry is directed, for example, toward the stimuli that trigger specific responses, how the coordination for muscle action is achieved, what mo­ tivates and terminates behavior, as well as other questions. One can also ask how and why specific behavioral patterns evolved. In order to answer these questions, which aim at the ultimate causes, one must first understand in which way the behavior in question contributes to fitness, as measured by the number of surviving offspring. In short, one must establish its function, or in this sense its task. Ex­ perimental and ecological designs, including the study of the ontogeny, are used to answer this question. The observation of behavior in the natural context is an important starting point for such an investigation. Whenever we encounter a structure or a behavioral pattern on a regular basis it is common sense to ask what that pattern's function is, in other words to begin with the presumption that the behavior fulfills some task. It can, of course, be the case that the particular behavior under observation from a selectionistic point of view is either neutral or even a disadvantageous heritage, dragged along as a burden of history, or as a by-product of other ad­ aptations. This is, however, only rarely the case. The various cost-benefit cal­ culations used by sociobiologists constitute a promising new way to study adap- tiveness; we shall explore this topic in a special section later in this volume. Some functions are immediately obvious. If a paleontologist finds a petrified wing impression he needs no experimental procedure to state that this is a flight organ. Should one find the fossil imprint of a structure looking like a camera eye, i.e., with lens, glass body, accommodation structures, and a projection surface like the retina, the investigator can state that he found an eye, an organ used for visual perception. This statement can be made even if the organ was found in a meteorite and it was thus impossible to make any experimental verification of the structure's function. Questions about function and development can be asked for cultural behavior patterns just as they can for phylogenetically evolved characteristics. We em­ phasize this because one occasionally comes across the point of view that human ethologists deal only with the basic "animal" heritage in human behavior. This is false. We also investigate man's cultural behavior. Thus questions as to specific adaptiveness can be posed both for phylogenetically and culturally evolved patterns and achievements from the biological point of view mentioned above. O. Koenig (1970) even coined the term "cultural ethology." Excellent examples of the eth- 4 ological approach to the study of cultural behavior include Koenig's investigations on the biology of uniforms and the ethological analysis of soccer by D. Morris (1981). We must also correct the frequent equating of phylogenetic adaptation with animal heritage. This is unwarranted because there are many phylogenetic adaptations that are specific to Homo sapiens alone. Consider, for example, our innate propensity for speech or the expression of crying. Human ethologists investigate complex behavioral sequences of individuals and of interactions among people and groups of people. They thus work on higher integration levels than physiologists, who are concerned with the elemental life processes such as stimulus perception, muscle contraction, and the conduction of nerve impulses. Although these processes are an important prerequisite to the understanding of behavioral events, one cannot deduce all the laws underlying any given social interaction from these elemental processes. Each higher level of integration has its own laws that cannot be derived from those of the levels below. Special emphasis must therefore be placed on the necessity of formulating questions appropriate to each level of behavioral organization. Human ethology makes use of experimental and analytical methods of behav­ ioral research developed in related disciplines, including techniques of data sam­ pling developed in anthropology and psychology, such as, the interviewing of informants. From animal ethology we take the methods of nonparticipant obser­ vation, techniques of recording and documenting, and the comparative approach. Human ethology also places value in studying behavioral patterns in their natural context (Section 3.3) and thereafter proceeding to experimental analysis. The theoretical basis of human ethology is critical realism (K. R. Popper, 1973; K. Lorenz, 1973, 1983). Hence, the basic assumption is that every "adaptation" reflects features of a reality outside the subject (p. 8). In order to make objective statements about the real world, our perception must be capable of reconstructing this world from the sensory data perceived. In order to reconstruct the real world we must be able to recognize constancy, even under changing conditions. Special mechanisms of constancy perception as well as our ability to recognize configurations (Gestalt perception, Section 2.2.2) enable us to do this. Since these are phylogenetic adaptations, we can say that the con­ stancy hypothesis is innate. The relationships between reality, perception, and knowledge can be demon­ strated by using a graphic projection model (G. Vollmer, 1983). If a cube is optically projected onto a screen, we can reconstruct it if we know its structure, the type of the projection, and the characteristics of the screen. In this manner we can accurately reproduce three-dimensional objects from two-dimensional projections on our retina. Even when we look at pictures, we interpret the objects portrayed on them in three dimensions. In the reconstruction of three-dimensional objects, the information lost in the projection process must be recovered. This takes place relatively reliably during the course of the perception process, with the exception of special cases, such as optical illusions. Cognition can be seen as an effort to reconstruct the real, "true" structures of reality outside the subject from the sensory impressions perceived as projections of these structures. Thus we interpret the cosmic signals perceived by our sensory organs as projections of astronomical objects. That the interpretation of these projections fits the outer world is proved by the success of interplanetary space travel. The ability to reconstruct a real world from sensory data presupposes a knowl­ edge about this world. This knowledge is based in part on individual experience 5 and, in part, on the achievements of data processing mechanisms, which we in­ herited as phylogenetic adaptations. Knowledge about the world in the latter in­ stance was acquired during the course of evolution. It is, so to speak, a priori knowledge—prior to all individual experience—but certainly not prior to any ex­ perience (Section 2.1).

The process of cognition consists of a step-by-step reconstruction of an hypothetically postulated reality, a stepwise liberation from the limitations of our sensory organs (the "screen" . . . ). This reconstruction process works counter to the chain of projections. While each projection effectively reduces information, in the process of cognition we attempt a reconstruction. Naturally, such a reconstruction must remain partial and hypothetical" (G. Vollmer, 1983, p. 64, translation of the German original).

In recent years, promising relationships have developed with other disciplines of human behavior and human culture. In this book we will discuss the work of many researchers who would not regard themselves as ethologists, but whose findings are ethologically relevant. This holds true for psychologists as well as ethnologists, sociologists, political and legal scientists, and art historians. A particularly close relationship exists with ethnologists and social anthropol­ ogists (E. Goffman, 1963, 1967; K. Jettmar, 1973; M. Godelier, 1978; H. Schindler, 1980). Common interests arise from comparative cultural work and from a mutual interest in the general, universal laws of human behavior, as demonstrated, for example, by the structuralism of C. Levi-Strauss (p. 517). These relationships are so varied that they cannot all be cited here. Our common interests touch on ques­ tions of early childhood development and socialization, societal structure, hier­ archical organization, aggression, ethical norms, and many more. Ties also exist with psychology and sociology. We will cite specific examples of this in the dis­ cussion of these themes. There are further common interests with linguistics, both at the level of concept formation and that of speech itself (p. 523). There have long been mutual relationships with medicine, especially psychiatry and psycho­ analysis (D. Ploog, 1964, 1966, 1969; J. Bowlby, 1969, 1973). The growing interest in ethological findings by political scientists is noteworthy. In America, biopolitics has developed as a new discipline (R. D. Alexander, 1979; C. Barner-Barry, 1983; P. A. Corning, 1981, 1983b; H. Flohr and W. Tdnnesmann, 1983; R. D. Masters, 1976, 1981b, c; G. Schubert, 1973, 1975, 1983b; A. Somit, 1976; A. Somit and R. Slagter, 1983). Art history and archeology have also been inspired by findings from ethology (D. Fehling, 1974; G. C. Rump, 1978, 1980; M. Schuster and H. Beisl, 1978).

Summary 1.1

Human ethology can be defined as the biology of human behavior. The research objectives are the elucidation of the physiological mechanisms underlying behavior, the discovery of the functions fulfilled by behavioral patterns and thus the unveiling of the selective pressures to which the behavior in question owes its existence, and, finally, the investigation of behavioral development in ontogeny, phylogeny, and cultural history, with emphasis on the question as to how and to what extent Man became programmed to act through phylogenetically acquired adaptations or by learning during individual ontogeny. Human ethology makes use of the con­ cepts and methods developed in animal ethology, but adapts them to the special 6 requirements associated with the unique position of man within the Animal King- dom. Methodological techniques are also utilized from the related disciplines of psychology, anthropology, and sociology. Through points of mutual interest human ethology thus endeavors to bridge the gap between the different human sciences. Human ethologists study not only the phylogenetically evolved behavior of man but also its individual and cultural modifiability. Critical realism is the episte- mological basis of human ethology.

Oh, great star! What would you be, had you not those on which to shine? Friedrich Nietzsche (from Zarathustra's prologue)

1.2. Phylogenetic and Cultural Adaptation

Life is defined today as an energetic process during which organisms, bearers of this process, extract more potential energy from their environment than they must expend in the acquisition of this energy. In other words, organisms are energy- acquiring systems with a positive energy balance. H. Hass (1970), the first to clearly formulate this concept to my knowledge, originated the term "Energon" for such energy-acquiring systems. The life processes are maintained by the mul­ tiplicity of organisms and by those energy-acquiring systems developed by them. Although we can trace the development of the process throughout evolution, we have yet to understand its ultimate causes. The energetic process presupposes structures at its disposal that are "adapted" to the appropriate energy transformations. Each organism must possess structures with which it can extract energy from the environment. These structures are adapted to the appropriate energy sources, that is, they are so constructed that they can tap the specific energy sources and thus aid in the maintenance of the energy-acquiring system. The environment is not only the source of energy but also the source of a multitude of interfering and even harmful influences against which the organism must protect itself. Organisms must also repair damage and be able to transform their positive energy balance into the procreation of their species. In short, there are a multitude of adaptations that require energy investment, such as reproduction, growth, protection, and similar tasks, which enter the balance as costs (H. Hass, 1970). Since organisms live in an ever-changing environment, they must be able to modify their adaptations in response to these changes. This may even require changes in their basic construction. Furthermore, there are a variety of changes to which an organism must continually adapt during its lifetime. Thus it must be able to institute short-term, reversible changes in response to temporary envi­ ronmental changes. The musculature and circulatory systems must be able to adapt to new stresses, and calluses must be able to form on the skin in response to pressure. Finally, an animal must be able to learn from experience in such a way that it modifies its behavior adaptively. Adaptive here refers to fitness as measured by reproductive success or, ultimately, the propagation of the genes of the indi­ vidual in question (see inclusive fitness, p. 92). Adaptation is thus a central problem for all organisms. Since the entire nature- nurture controversy centers around the question of the source of adaptation, we will add a few basic remarks about this subject here. Adaptations depict features of the environment relevant to survival. They reflect facets of the external reality, such as, characteristics of the energy source they 7 tap or the environment in which they move. Thus, certain characteristics of water are reflected in the anatomy of fishes and dolphins, by adaptations that are func­ tionally related to the movement of these organisms in this medium. This is what is meant when we say that an adaptation reflects features of the environment. The degree of detail and exactness in which an adaptation depicts the outer world varies from case to case since it is determined by function. The organism only reflects characteristics of the environment relative to fitness and not every detail of the environment. However, the match can be amazingly complete, as for ex­ ample in the case of mimicry, in which a mimic imitates its model even down to the smallest details. The leaf insects, for example, mimic the leaves of the shrubs on which they live. Interesting examples of mimicry are reviewed in W. Wickler (1968). K. Lorenz (1941, 1973) has made a significant contribution to the question of what constitutes reality in his biological epistemology. In our adaptations we do not depict the external world—pictorially (iconically), but our thought processes and our perceptual capabilities fit as adaptations in the same way as do our physical characteristics. "Our cognitive and perceptual categories, given to us prior to individual experience, are adapted to the environment for the same reasons that the horse's hoof is suited for the plains before the horse is born, and the fin of a fish is adapted for water before the fish hatches from its egg" (K. Lorenz, 1941, p. 99, translation of the German original). The adaptations of an organisms, whether these are bodily structures or ad­ aptations in behavior, represent assumptions (hypothesis) about the world in which it is going to live—assumptions which have been tested by selection. Our perceptual organizations come into disarray when we move into other en­ vironments to which we are not phylogenetically adapted, as for example in the atomic or astronomical environments. Our causal thought processes are proof of this. We associate events by means of rigidly determined programs and extrapolate from the coincidence of events: When-then (coincidence) to causes (because— then). We can only think in causal sequences, and this becomes a problem when we use these intellectual tools to inquire into the origin of the universe. David Hume states that causality may not actually exist in nature, but that it is just a "necessity of the soul." The remarkable essay by R. Riedl (1981) contains a chapter on the consequences of causal thought: "The Superstition of Causes." This chapter title is misleading and more appropriately should have been "The Established Faith in Causes," since for the middle level—the "mesocosmos"—causal thinking is an appropriate adaptation. Accurate knowledge based on an iconic depiction of reality cannot be accomplished and is not really necessary for research. We perceive the world with preconceived perceptual and thought processes that strongly influence the postulation of our hypotheses (p. 105). Our perceptive and cognitive mechanisms, nonetheless, fit well enough to an outer world to enable us to send satellites into outer space, which will eventually arrive at their goal even after years of travel. That our cognition is capable of this is a matter of interest and it is more reasonable to assume that our cognition corresponds to external reality than to take the standpoint which disavows the existence of a world independent of a perceiving subject. In P. Watzlawick's (1981) Die erfundene Wirklichkeit ("Invented Reality"), these problems are precisely and intelligibly discussed by the various authors. Other notable contributions have been made by R. Riedl (1979), G. Vollmer (1975, 1983), D. Campbell (1974), and G. Radnitzky 8 and W. W. Bartley (1987). In their organization, organisms mirror facets of the extrasubjective environ­ ment. Even though the postulates of critical realism are neither demonstrable nor refutable, as Karl R. Popper states, we can "argue in their favor, and the arguments are convincing" (K. R. Popper, 1973, p. 50, translated from the original German text). This is the position any sensible scientist must take. Whether this is "critical realism" or "hypothetical realism" is, from my point of view, of little significance. G. Vollmer (1983) is of the opinion that one must differentiate between these, since critical realism at least considers the existence of the world as evident, ret­ rospectively unquestioned and intuitively guaranteed, while hypothetical realism differentiates between psychological certainty (hypothesis about the world) and epistemological uncertainty. But critical realists like Popper have a clear under­ standing of the epistemological uncertainty, and the hypothetical realists proceed from the postulation of an extrasubjective reality, which we reflect incompletely but nevertheless fittingly in our adaptations. This is not surprising, for, using an illustration from Ernst Mayr: the monkey without a realistic perception of the branch on which he wants to jump would soon be a dead monkey—and thus would not be one of our ancestors. From the above it is evident that every adaptive system must have acquired "information" from its environment at some time relevant to the organism's ad­ aptations, which reflect these environmental features. For this to occur, the or­ ganism has had to have interacted with the environment (K. Lorenz, 1961). Such an interaction can occur during phylogeny through the mechanisms of mutation, recombination, and selection. Information relative to survival has thus been col­ lected each generation and became codified genetically, providing the develop­ mental instructions that guide the process of self-differentiation by which the in­ dividual organisms or phenotypes develop. This developmental scheme is so strongly controlled by the genes that each individual develops the various species- specific structures during ontogeny. Special feedback mechanisms prevent ex­ cessive deviations from the blueprint. This is why sparrows hatch from sparrow eggs and chaffinches from chaffinch eggs and not, accidentally, a duck. Each characteristic develops within a specified range of variation. This range— the reaction norm—is genetically determined and thus the result of phylogenetical adaptation. The modifications of plants in different habitats illustrate this point. The mountain forms of many plants often have feltlike hairs and are generally shorter-stemmed and more compact than conspecifics growing in valleys. Such variability is clearly adaptive in most cases. However, it can also be an epiphe- nomenon. There are plants that develop red flowers in acid soil and blue in alkaline soil. The anthocyanins (flower pigments) turn red in acidic cell fluids and blue in weakly alkaline ones. We have not been able to deduce the selective advantage for certain plants to possess both color morphs depending on habitat. Perhaps this characteristic of the range of variation is neutral in terms of selection and may even be an epiphenomenon. The range of variation during development is limited and the environmental pressures cannot force modifications to occur in any arbitrary direction. However, the genetically determined prospective potential is often greater than what is normally observed. Gall wasps possess chemical means of stimulating plant tissue to grow galls, which they would not normally do. Generally the range of modifiability of genetically determined processes of differentiation is rather limited, and the prospective potential decreases with each further step in differentiation. Phylogenetic development is not directed toward specific goals. The direction 9 taken is dictated more by selection contingencies. In a process analogous to trial- and-error learning, an exploration of possibilities takes place at the level of the species. Many writers have been disturbed by the decisive role of "chance" in the development of the amazing diversity of organisms. Once, however, it is made clear that basically it is a question of species maintaining themselves in an en­ vironment of unforeseen changes, then it is understandable that such adaptation only works when a continual testing of all possibilities of constructive change takes place. This is the only way that "promising monsters" (R. B. Goldschmidt, 1940) can be created and open new pathways in evolution.2 These can be ethological (p. 14) or morphological monsters. Thus in every generation of flies there are wing­ less mutants that are weeded out from the population as being malformed. How­ ever, that even such monsters of evolution have a chance can be seen in the subarctic Kerguelen Islands, where only wingless insects live (beetles, butterflies, and flies). The winged forms are continually blown off the island by storms. From generation to generation, new genetic variants are thus created and tested by selection. They are, in a way, set into the world to test for new changes that might have come about. Extreme specialization limits the potential for further evolution. Whereas the pentadactylid foreleg of the vertebrate had the potential to evolve into the fin of the whale, the wing of a bat or the hooves of a horse, the potential of these latter specialized structures is greatly limited. A fin cannot become reconstructed into an extremity with five fingers or a wing. Specialists, therefore, run a greater risk of extinction when their environment changes drast­ ically. The future lies with the generalists. During the daily life of organisms there are numerous situations in which a rapid adaptability is advantageous. An insect seeking flowers would benefit from remembering where a plant had just blossomed. It is also advantageous for animals to remember where predators are or which pathway is a safe escape route. It is useful when an animal can learn from individual experience, i.e., is able to retain information about specific data or experiences, to store this information, and to adapt its behavior accordingly. During the course of phylogeny, structures of the central nervous system evolved that permitted individual experiences to be stored in recallable engrams so that future behavior could be adjusted accordingly on the basis of these experiences. Here, as well, phylogenetic adaptations determine whether what is learned is obligatory or optional. Animals learn what contributes to their fitness and, since this varies from species to species, innate learning dis­ positions vary accordingly (p. 74). Even though learning is based on phylogenetic adaptations and channeled by

2Chance is creative in this way. H. Markl (1980) has provided a pertinent definition: "I have dealt with these problems so exhaustively because chance stands in such poor stead with learned people: chance, it is said, is never creative. It is merely known as the un­ imaginative shepherd pushing his herd of events toward the gate of average expectation, lacking any originality. Rarely has something been so badly misjudged. For it is only through chance that something new can arise, for novel value only exists in terms of that which does not follow the undeviating path of the standard laws. If we termed it 'spontaneity,1 we would recognize the unpredictable character that only freedom of behavior can produce. However, it is important from the molecular level to that of behavior: chance, left to itself, can create nothing lasting, for it destroys as it builds. A mechanism that evaluates chance, retaining the useful and rejecting the unusable, is required to put to use the innovative potential of chance. Chance produces nothing of value without a selection process occurring, 10 but selection has nothing to select without chance" (translated from the German original). them, new perspectives of evolution were opened with learning, which, among others, is one of the prerequisites for cultural evolution. In general, each animal learns from experience for itself only. However, some species can also learn by imitation. In a free-living group of Japanese macaques on the Japanese island of Koshima, a female discovered the washing of sweet potatoes before eating them. The others saw this and imitated her. Today this behavior has become established as a group-specific characteristic and is a tradition passed on from one generation to the next. Such behavior is sometimes described as "protocultural," but it was only with the evolution of speech and the subsequent development of writing and other information storage systems that true culture could arise. In the protocultural stage, one individual has to show another the behavior to be learned. With the development of speech it became possible to carry on a tradition independently of an object by verbal instruction alone. One Japanese macaque must always see another washing potatoes, but one person can instruct another by saying, "We wash potatoes before we eat them." He could also per­ manently record this information in writing so that it would be available for cen­ turies to literate individuals. Thus, with the introduction of writing, a new, more powerful data bank becomes available to assist the memory enabling knowledge to be passed on without the presence of another society member. Thus accu­ mulative culture develops. With cultural evolution a new level of existence is achieved, since culturally acquired knowledge can maintain existence, to a large extent, independently of its inventors. If the inventor of the radio or another technical innovation died without heirs, the invention could still be perpetuated. Theoretically it could even be utilized by genetically unrelated inhabitants of another planet. Ideologies can survive in the same way and thus have an existence of their own. K. R. Popper (1973) speaks of a world of objective thought content, which exists even though it is a product of our mental activity. The products of this world (theories, books, and others) can be studied without knowledge of the ways in which they originated, in the same way as we can investigate spider webs and bird nests without having seen the animals that made them. Books can outlive us and be studied by other intelligent beings. Popper categorizes this world of objective ideas as World 3, with the world of states of consciousness and behavioral dispositions (World 2), and the physical world (World l).3 All our "artificial organs" (H. Hass, 1970), from tools to books and from fac­ tories to expressways, belong to Popper's World 3. They demonstrate effectively how much and in which ways our accumulating culture dominates human evolution. New ideas, theories, and inventions act like mutations in the biological realm, and they have to stand the test of selection. The problem to be solved is the survival in offspring and this is achieved by the elimination of mistakes. Popper mentions that anatomical and behavioral adaptations of plants and animals are biological analogs of theories: "Theories correspond (like many external objects such as honeycombs . . . ) to bodily organs and their functions. Theories and organs alike are experimental adaptations to the world in which we live" (K. Popper, 1973, p. 125).

3One can also describe World 2 in broader terms as the world of the organic, which differs from World 1 (physical) since its organization is derived from selective pressure (N. Bischof, 1981). 11 Progress in knowledge is the result of a process quite similar to natural selection but consists of the selection of competing hypotheses. "This interpretation applies to what animals know, prescientific knowledge and also scientific cognition. While the animal and prescientific knowledge grow mainly because those with unsuitable hypotheses are themselves selected out, scientific critics often let our theories die instead of us; they select against our wrong ideas before we ourselves are eliminated because of them (Popper, 1973, p. 289). Cultural evolution is by no means always a product of rational planning. F. A. von Hayek (1979) pointed out clearly that mankind, with the ability to imitate and to carry on traditions, develops traditions of learned rules of behavior, whose purposes are, in most instances, not understood. Von Hayek continues that in the past, Man has often learned appropriate behavior without understanding why it is correct, and even today many habits usually serve him better than does rational understanding. The system of traditional rules was tested and developed in a process of screen­ ing, which is analogous to selection on at the biological level. Cultural evolution, therefore, is mainly a result of the forces of selection and is, to a lesser extent, the result of conscious rational planning. In a competitive process, the more suc­ cessful characteristics prevail. Von Hayek is undoubtedly correct when he states that cultural evolution could proceed without any kind of insight and also when he expresses the opinion it has also done so for the majority of mankind's existence. If we consider culture to be the totality of traditional adaptations, then his statement that we were not even capable of designing culture is also true. Even today we can only understand partial aspects of our own culture.4 Nonetheless we must come to terms mentally with the problems of our existence, and through reason come up with solutions. Man has had great success with rational planning in many realms of science, economics, and technology. Furthermore, we can set goals for ourselves and thus significantly influence our further cultural develop­ ment. The developments initiated by founders of states and ideologists testify to such developments. Ideologies can reverse developmental trends completely by setting new goals and thus by altering contingencies of selection. Whether this is of advantage or not from the point of view of the fitness of the society would not be determined until later. For those Russian sects that forbade their members to engage in sexual intercourse—and who disappeared soon afterward because of so few new recruits— ideology was shown to be deleterious. The experiments of modern mass societies, whose ideologies stress the equality of all of its members, are fascinating. With few exceptions, in the majority of cultures, successful men, those controlling the natural resources, also produced more descendants. They could afford to have more women, and the women of their subordinates also stood at their disposal. Successful Yanomami chieftains have more wives and children than others in the tribe (N. A. Chagnon, 1979). In

4Cause and effect are so intertwined here that we cannot usually use our linear causal reasoning to understand all the relationships operating here. D. Dorner and R. Reither (1978) contributed to this concept in their experiment giving twelve intelligent students the task of improving the living conditions on the fictitious developing country "Tana." The conditions in the country were known and could be varied. In almost all cases, the student destroyed the originally stable structure and often produced catastrophic consequences for 12 the country. China the wealthy not only had supplemental wives and concubines but also access to the wives of their subordinates. The reigning ideology of the Chou Dynasty (1100-222 B.C.) stated that this was their subjects' obligation and fostered among subordinates the wish to comply since they believed that only the ruling class possessed certain magical powers, which they transmitted for the common good through reproduction. In the above instances the ideology supported those controlling the society's resources and led to growth in the population of the ruling class at the expense of the lower classes (further examples in K. MacDonald, 1983). With Christianity, which interestingly enough spread first among the slaves and the poor of Rome, the ideology of equality—of "leveling" as Nietzsche expressed it—came into ex­ istence. It was directed against those controlling the resources and attempted to construct an ethic of sharing and equalization as well as of enforced monogamy. A papal ban against princes was one means of stopping polygamy in the ruling European classes. The ideology of monogamy with its prohibition against birth control was the most decisive step since it removed the privilege of reproduction from the upper classes. Anyone who was capable of providing nourishment for a family could sire offspring. The division of property was less important, and such differences were even­ tually tolerated by Christianity. Such an ideology leads to the situation in which there is not a limited number of privileged families reproducing at a higher rate relative to other classes, but one in which all levels of society reproduce. It remains to be established whether the single wife in a monogamous society without birth control has more children than those in polygamous societies. I believe it is possible but cannot cite any figures for this. Perhaps this was a factor responsible for the great success of Europeans throughout the world. The Hutterites of Canada can act as a model as to how a group with a strict monogamy and no birth control achieves a higher birth rate than other groups in the Canadian population. The question here is what sort of effect this pattern of behavior will have in the long term. Populations who obey an ideology of reproductive egalitarity may experience a rapid increase in number over other groups following other prescriptions. At the same time, however, the further evolution of certain traits may be slowed down.5 Man is certainly the first being who consciously set goals. Some went as far as to argue that Man, therefore, overpowered nature—that he emancipated himself from the contraints of biology. But whatever man does will be measured by the yardstick of selection. By striving toward a particular goal set by an ideology man directs his future evolution and secures its course within certain limits against the unpredictabilities of chance. However, it will always be selection that will decide whether his choice was correct. Any kind of concept, even the ideas of an insane person, can influence the fate of a people, for better or worse, just as mutations can in the realm of biology. Today we are experiencing how selection is sifting out the newly established

5In this context some recent developments are of interest. D. R. Vining (1986) drew attention to the fact that in modern society the less economically successful produce more children than those better endowed. This leads to a decline of the IQ from one generation to the next in modern societies. Presently this does not need to bea^natter of grave concern, since such a trend might be temporary. However, it might be useful to observe further developments. 13 social and economic systems of the world according to their adaptiveness. We cannot know the results of this experiment, but without doubt various goals have selective consequences, and they may ultimately even influence biological evo­ lution. In the animal world, behavioral changes are the pacemakers of phylogenetic development (E. Mayr, 1950, 1970). If an insect larva shifts to a plant species not yet utilized by the species and becomes imprinted on the new host plant so that the imago henceforth selects this host plant, this new preference will certainly result in other adaptations in the insects' life habits and anatomy. In choosing this new host plant, the insect was placed under new selective conditions. For example, if the epidermis of the new host plant is firmer, the insect may develop stronger mandibles for biting. K. R. Popper (1973), in his "speartip theory" of behavioral evolution, inde­ pendently developed the same ideas as Ernst Mayr. He presumes that specific innate tendencies (the avoidance of danger, search for food, etc.) are subject to mutations without organs of the body being simultaneously affected. Any changes affect only the apparatus controlling behavior—in vertebrates it is the central ner­ vous system and not the morphological structures that carry out the actions. These can be independently altered by other mutations. Mutative changes in the mech­ anism's guiding behavior (Popper speaks of a "central tendency structure"6) gen­ erally disturb the functions of an organism less than mutations in structures ex­ ecuting behavior. It is easier to correct maladaptive changes in the central tendency structure with compensating behaviors. New behavioral tendencies can determine further development. Popper cites changes in feeding habits as an example and mentions in this context the classic Lamarckian example: the giraffe. He is of the opinion that changes in the eating habits must have preceded the anatomical neck modifications. A Darwinian explanation can thus be found for the much discussed topic of orthogenesis (goal-directed evolution).

If a new goal, tendency, disposition, capability or behavior pattern develops in the central tendency structure, this would influence the effects of natural selection in such a way that previously maladaptive (but potentially favorable) mutations now become adaptive ones if they support the newly developed tendency. This means, however, that the development of the executing organs will be guided by this new tendency or objective and is thus 'goal-directed' (K. R. Popper, 1973, p. 305).

Popper (1973) speaks of "promising ethological monsters" (see also p. 10), which deviate markedly from their parents in their behavior and thus introduce new developments. Certainly a new development is more likely to be preceded by a new behavior pattern than it is by a change in structure, since a new behavior is less likely to fatally disturb the functioning of an organism as a whole and is also less likely to prove lethal than a monstrous anatomical change. A new behavior pattern that takes advantage of previously existing light-sensitive structures could significantly enhance their selective value. Thus an "interest" in vision could

This "central tendency structure" consists of, on the one hand, built-in instructions, for example, to flee from one situation or to attack in another (Popper speaks of goal structures), and, on the other hand, of situations enabling one to interpret external stimuli (ability structures). These two, however, cannot be distinguished from one another, so 14 Popper introduces the concept of "central tendency structure" to unite both structures. become one of the key factors in the orthogenesis of the eye.7 In this way, according to Popper, changes in the goal structure lead and the development of the anatom­ ical-physiological structures follow. "As the vitalists claim, the direction can ac­ tually be determined by a tendency similar to consciousness—by a goal structure or the capability structure of the organism, which can develop a tendency or a 'wish' to use an eye, and an ability to interpret the perceptions which it perceives from it" (K. R. Popper, 1973, p. 307). I have referred to these concepts in some depth because I feel they are highly significant for Man. What has been stated here does not only apply to genetically controlled behavioral mutations. In the human sphere, new ideas, ideologies, and other mental concepts, as stated earlier, can also become evolutionary pacemakers and determine its future course. Developments like these lead to cultural delin­ eations and thus initiate separate developments, termed aptly "cultural pseudo- speciation" by E. H. Erikson (1966). To what extent innate values channel the course of evolution will be discussed later. There are indications that our concepts of "differentiated" and "higher" levels are universal goal structures and thus determine our behavior (p. 659). Cul­ tural evolution is also strongly influenced by altruism, which does not appear to be exclusively a product of culture. With cultural evolution, mankind developed an adaptive mechanism that in historical times has no doubt exceeded the biological ones in importance. It is probable that cultural changes in life style are followed by subsequent genetic modifications and there are sound indications for this. Lactose tolerance is ob­ served in the majority of adults in peoples living on milk products—a trait lacking in hunter/gatherer societies. Presumably with the discovery of agriculture and the growth of the habit of drinking milk as an adult, a corresponding genetic adaptation was selected for (L. L. Cavalli-Sforza, 1981; further citations therein). In general we have not changed much in our anatomy in the last 20,000 years. And from the way we handle our culturally acquired knowledge we may even assume that our basic behavioral outfit has remained the same. People with the motivations and intellectual capacity of late upper paleolithic man are now flying jet bombers and confronted with the task of surviving in anonymous society. Since phylogenetic and cultural evolution are shaped by similar selection pres­ sures, cultural evolution phenocopies the biological evolution in many ways. Ex­ amples of this are found in the comparison of cultural and phylogenetic ritualization (p. 439). The same functional laws apply to both areas. Thus the rate of change or of mutation should in neither case be excessive, for an excessive mutation rate is dangerous. Progress depends on the balance achieved between the preserving "conservative" forces and those promoting change. We stick to the proven, but experiment with change in small doses. This is certainly "adaptive," because it is improbable that the entire store of cultural tradition should have lost its adaptive value from one generation to the next. Our need for security makes us cling passionately to our "beloved" customs. It is from this secure base that we experiment with new ideas and insights.

7"Thus an interest in seeing could be fixed and may become a significant factor in the orthogenesis of the eye; even the smallest improvements in its anatomy could have a high survival value if they were utilized by the organism's goal and capability structure" (K. R. Popper, 1973, p. 311). 15 Summary 1.2 Living systems owe their organization to processes of adaptation through se­ lection. Adaptations are all structures which contribute to the survival of the in­ dividual in question and ultimately to its survival in offspring or in short to its fitness. In their adaptations, organisms mirror features of their outer subjective world relevant to fitness. They represent sets of hypotheses about this world which were tested by selection. Thus a fin of a fish, as well as the motor program for its use, both develop prior to hatching within the egg, represent assumptions about the environment in which the fish is going to live. As a prerequisite for adaptation to occur, an interaction between the adapted organismic system and its environ­ ment must take place. During phylogeny, variants of the phenotype, brought about by genetic changes, are tested by selection as to their adaptedness and the geno­ types for adaptive traits retained. This way the organisms can be said to inform themselves in a process analogous to a trial and error learning about its environment and that information is coded and stored genetically. Phylogenetic adaptation is supplemented by adaptation via learning and indi­ vidual experience, and, in humans, also via traditions. It is in the latter case that information transfer occurs. Cultural evolution phenocopies phylogenesis in many respects, since in both cases the shape of the resulting pattern is determined by selection. Customs must prove themselves in order to be retained. We may, how­ ever, learn from failures by eliminating customs without sacrificing the life of the individual at the same time. Behavior patterns are often the pacemakers of further development in both cultural and phylogenetic evolution. Cultural evolution has certainly proceeded in an unplanned manner for most of human existence, and we tend to observe certain customs without insight into their rationale since they have proved themselves beneficial. Culture, because of its complexities, is difficult to plan, but by the setting of goals its development can be directed.

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