Doris Zumpe and Richard P. Michael Emory University School of Medicine Atlanta, Georgia Kluwer Academic/Plenum Publishers Copyright 2001 In fond memory of my parents and of my first great teacher, Konrad Lorenz. D.Z. PREFACE These notes are intended to help undergraduates who need to understand something of behavior both for its intrinsic interest and for their future careers in medicine, biology, psychology, anthropology, veterinary medicine, and nursing. In Emory University's Biology Department, a single-semester course called Evolutionary Perspectives on Behavior is given to undergraduates. It amounts to four, not eight months of study, so a great deal of compression is essential. There are several excellent textbooks available that deal with behavioral science from different perspectives, but we have found them too compendious for use in a short course when students are so heavily burdened; it is unsatisfactory to direct them to a chapter here and there in several different books or to this or that review article and original paper. In this volume, we have tried effectively and inexpensively to put in one place what we know is needed. The topics we have selected deal with their subjects in a simple, straightforward way without being too superficial. We could not cover everything and the gaps are not entirely idiosyncratic but reflect what students are given very well in other courses. Thus, there is no mention of the physiology of the axon and synapse; learning, memory, cognition, and basic genetics are hardly touched upon because students know about these matters from elsewhere. This volume particularly emphasizes certain physiological mechanisms when they are known, and it also draws attention to the application of what is known about animal behavior to the human, sometimes even to the clinical situation. The illustrative examples are taken from both the classical behavioral literature and newer work. It has given us considerable pleasure to write this volume and we trust it will find a useful niche in the education of our students. CONTENTS 1. The Study of Behavior: History 1 A Brief History 1 Classical Ethology 1 Comparative Psychology 4 Emergence of Modern Behavioral Research 5 Some Theory and Terminology 6 Fitness and Inclusive Fitness 6 Benefit-Cost Ratios 7 Optimality Theory 7 Game Theory 8 Evolutionarily Stable Strategies 9 General Life Strategies: K and r Selection 10 Warnings, Fallacies, and Pitfalls 11 Correlational Studies 11 Genes: What They Do and Do Not Do 12 Understanding Optimality Theory 12 2. Some Ethological Concepts 15 Evolutionary Basis of Behavior 17 Ethology's Objectives 17 General Methods 17 Fixed Action Patterns (FAPs) 18 Unvarying Form 18 Coordination of Several Muscle Groups 19 Environmental Influences 19 Genetic Factors 22 Brain Stimulation 22 Conflict Behaviors 22 Redirection 24 Displacement 25 Intention Movement 27 Alternation 27 Ambivalence 27 Compromise 28 Conflict in Psychiatry 28 Vacuum Activity 30 Ritualization and Displays 30 Stereotypy 31 Typical Intensity 31 Association with Conspicuous Morphological Features 31 Threshold Changes 32 Motivational Changes 32 3. Some More Ethological Concepts 37 Sign Stimuli, Releasers, and Innate Releasing Mechanisms 37 Perception versus Attention 37 Definitions 38 Properties of Sign Stimuli and Releasers 38 Modifying Influences 41 Supranormal Stimuli 43 Advantages and Disadvantages of Innate Releasing Mechanisms 43 Programmed Learning and Imprinting 44 Programmed Learning 44 Imprinting 45 Drive or Motivation 48 Illustrative Model for Thirst 49 Drive Models 51 4. Assessment of Hereditary Influences 55 Indirect Methods 57 Presence in Geographically Isolated Populations 57 Presence in Individuals Raised in Isolation from Conspecifics 58 Phylogenetic Comparisons: Presence in Closely Related Species 58 Association with Specialized Morphological or Physiological Traits 58 Studies on Human Twins 59 Direct Methods 59 Artificial Selection 60 Inbreeding 60 Hybridization 64 Molecular Changes 64 5. Behavioral Endocrinology: Gonadal Hormones 67 Synthesis and Major Sites of Production 69 Estrogens and Progestins 69 Androgens 70 Transport 71 Mechanisms of Action 71 Hormone Receptors 71 Effects on Tissues 71 Organizational Effects During Development 74 Sexual Differentiation: Somatic Modifications 74 Sexual Differentiation: Behavioral Modifications 75 Sexual Dimorphism in Brain Structures 77 Activational Effects in Adults 77 Breeding Seasonality 78 Activational Effects in the Female 78 Activational Effects in the Male 83 6. Behavioral Endocrinology: Stress and Adrenal Hormones 89 Definition 89 Types of Stress 89 The Stress Responses of the Body 90 The Adrenal Medulla and Sympathetic Arousal 91 The Hypothalamus and the Adrenal Cortex 91 Corticosteroid Production 92 Corticosteroid Metabolism 93 Habituation to Stress 95 Functions of Stress 96 Psychosomatic Medicine 96 Two Psychiatric Syndromes 97 7. Biological Rhythms 99 Functions of Biological Rhythms 99 Circadian Rhythms (23-26 Hours) 101 Studying Circadian Rhythms 102 Location of the Internal Clock 105 Circadian Rhythms in Humans 108 Circatidal Rhythms (12.4 Hours) 111 Circalunar Rhythms (14.8 Days) 111 Monthly Rhythms (29.5 Days) 112 Circannual Rhythms (365 Days) 113 Rhythms in Human Disease 113 8. Orientation and Navigation 117 Orienting Responses 117 Kinesis 118 Taxis 118 Navigation 121 Navigational Mechanisms 121 The Compass Mechanism 124 Navigational Cues 125 Migration 128 9. Feeding, Foraging, and Predation 131 Feeding Behavior 132 Social Learning and Facilitation of Feeding 132 Foraging 134 Optimal Foraging 135 Constraints on Optimal Foraging 138 Coping with Changes in Food Supply 139 Feeding in Humans 144 Some Physiological Aspects of Feeding 145 Predatory Techniques and Antipredator defense 146 Somatic Adaptations 146 Predatory Techniques 148 Antipredator Defense 149 10. Social Behavior 151 Social Systems 151 Coelenterate Colonies 152 Eusociality in Insects 152 Vertebrates 154 Benefits of Sociality 155 Reduction in Predator Pressure 155 Improved Foraging and Hunting Efficiency 156 Improved Defense of Limited Resources 156 Improved Care of Offspring 157 Costs of Sociality 157 Increased Competition between Conspecifics 157 Increased Risk of Infection 157 Increased Risk of Mating Interference and Parental Exploitation by Conspecifics 159 Increased Risk That Offspring Are Killed by Conspecifics 159 Philopatry and Dispersal 159 Dispersal Hypotheses 160 Evolution of Cooperative Behavior 162 Cooperation (Mutualism) 163 Reciprocity (Reciprocal Altruism) 163 Altruism (Kin Selection) 164 Mechanisms of Kin Recognition 165 Location 165 Familiarity 165 Phenotype Matching 166 Allele Recognition 167 Environmental and Cultural Influences in Primates 167 Nonhuman Primates 167 Humans 168 11. Communication 171 Definition 171 Functions of Communication 171 "Honesty" and "Deception" in Communication 172 Communicatory Signals 174 Sensory Channels of Communication 176 Visual Communication by Reflected Light 177 Auditory Communication 180 Chemical Communication (Olfaction) 188 Tactile Communication 195 Electrical Communication 197 12. Agonistic Behavior 199 Interspecific Agonism 200 Predatory Aggression 202 Antipredatory Agonism 203 Intraspecific Agonism 203 Individual Distance 203 Intraspecific Aggression 204 Categories of Intraspecific Aggression 206 Intraspecific Submission and Flight 215 Comparisons between Interspecific and Intraspecific Agonism 216 Human Aggression 216 13. Sexual Selection 221 Asexual and Sexual Reproduction 221 Sex Determination 222 Sex Ratio (SR) 223 Theoretical Considerations 224 Bateman's Principle 224 Trivers's Theory of Parental Investment 224 Trivers-Willard Hypothesis 226 Intrasexual Selection 227 Competition among Males 227 Competition among Females 223 Intersexual (Epigamic) Selection 233 Desirable Male Qualities 233 Markers of Male Qualities 233 Evaluation of Male Qualities 234 Evolution of Male Traits and Female Preferences for Them 234 Mate Choice by Males 236 14. Courtship and Mating 237 Factors Important for the Onset of Courtship and Mating 238 Seasonal Factors 238 Hormonal Stimulation 238 Social Stimulation 239 Functions of Courtship 239 Species (and Strain) Identification 239 Gender Identification 240 Aggression Reduction between the Male and Female 240 Individual Recognition 241 Behavioral and Physiological Synchronization between the Male and Female 241 Signaling Competitive and Parental Abilities 242 Mating Categories 243 External and Internal Fertilization 243 Copulatory Patterns in Mammals 243 Bisexual Behavior 244 Nonhuman Animals 244 Humans 245 15. Parental Behavior and Mating Systems 249 Models of the Parent-Offspring Relationship 250 The Parental Provision Model 250 The Mutual Benefit (Symbiosis) Model 250 The Conflict Model 250 Evolution of Parental Care 252 Sex Differences in Parental Care 252 Selection Pressures for Parental Care 255 Mating Systems 258 Definition of Mating Systems 258 Classification of Mating Systems 259 16. Nonhuman Primates 265 Ethology 265 Fixed Action Patterns (FAPs) 267 Conflict Behaviors 268 Ritualization 268 Releasers 269 Sensitive Periods, Imprinting 272 Sociobiology 273 Social Systems 273 Mating Systems 277 Mate Competition and Mate Choice 277 Parental Investment 278 Dispersal and Inbreeding Avoidance 279 Hormonal and Seasonal Influences 282 Language in Apes 286 17. Humans 289 Human Ethology 289 Fixed Action Patterns (FAPs) 290 Conflict Behaviors 291 Ritualization 292 Releasers 294 Ethology in Clinical Settings 298 Sensitive Periods, Imprinting 298 Human Sociobiology 299 Mating Systems 299 Mate Competition and Mate Choice 299 Parental Investment 303 Incest Avoidance 305 Hormonal and Seasonal Influences 308 References 313 Author Index 323 Subject Index 327 WHERE TO ORDER: Kluwer Academic Publishers Order Department P.O. Box 358, Accord Station Hingham, MA 02018-0358, USA Tel: (781) 871-6600 Fax: (781) 871-6528 E-mail: kluwer@wkap.com http://www.wkap.com Price: $85.00 ISBN: 0-306-46577-9 (hardbound)
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