Perspectives on Animal Behavior

Chapter 1 Introduction 1 Four Questions About Animal Behavior 2 Animal Behavior as an Interdisciplinary Study 2 The Interplay of Questions: A Case Study 3 Hypothesis Testing 5 Part 1 Approaches to the Study of Animal Behavior Chapter 2 History of the Study of Animal Behavior 11 The Beginnings 11 Intellectual Continuity in the Animal World 11 Darwin's Evolutionary Framework 12 Classical Ethology 13 The Approach: Evolutionary, Comparative, Descriptive, Field-Oriented 13 Classical Ethological Concepts 15 Comparative Psychology 17 The Approach: Physiological, Developmental, Quantitative, Laboratory-Oriented 17 Early Concepts of Comparative Psychology 17 The Roots of Physiological Psychology 20 Sociobiology and Behavioral Ecology 20 More Recent Trends 22 Field Studies 22 Cellular and Molecular Bases of Behavior 22 Behavioral Biology 23 Applied Animal Behavior 23 Chapter 3 Genetic Analysis of Behavior 27 Basics of Gene Action 28 Goals of Behavioral Genetics 31 Methods of Behavioral Genetics 31 Inbreeding 31 Artificial Selection 33 Inducing Mutations and Screening for Change in Behavior 35 Finding Natural Variants and Looking for Genetic Differences 36 Hybridization 37 The Foraging Gene as an Example of Behavioral Genetics in Action 38 Candidate Genes 38 Linking a Protein to a Trait 40 Locating all the Genes Associated with a Trait 41 Microarray Analysis 41 Important Principles of Behavioral Genetics 42 One Gene Usually Affects Several Traits 42 Genes Work in Interacting Networks 44 Behavioral Variation and Genes 45 Environmental Regulation of Gene Expression 47 Dominance Relationships in Cichlid Fish 47 Song Learning in Male Songbirds 48 The Importance of Genetic Background to Behavioral Genetics 50 Networks of Genes are Responsive to the Environment 51 Epigenetics and Behavioral Genetics 52 Complex Relationships Among Genes 54 A Broader Perspective 54 Chapter 4 Natural Selection and Behavior 57 Natural Selection 58 Common Misunderstandings About Natural Selection 60 Genetic Variation 61 Variation is Common 61 The Raw Material of Genetic Variation 61 Variation and the Response to Natural Selection 62 The Maintenance of Variation 63 Gene Flow and Genetic Drift 63 Correlated Traits 65 Changing Environmental Conditions 65 Frequency-Dependent Selection 67 Negative-Assortative Mating 68 Evolutionarily Stable Strategies: Fitness and the Behavior of Others 68 Testing Hypotheses About Natural Selection and Adaptation 70 The Experimental Approach 72 The Comparative Approach 73 Monitoring Selection in the Field 75 Modeling the Costs and Benefits of Traits 75 Chapter 5 Learning and Cognition 77 Definition of Learning 78 Types of Learning 78 Habituation 78 Classical Conditioning 80 Operant Conditioning 82 Latent Learning 84 Social Learning 85 Species Differences in Learning: Comparative Studies 87 The Ability to Learn as a Heritable Trait 87 Evolution and the Variation in Learning Across Species 88 Other Evidence of Cognitive Abilities in Animals 91 Tool Use 92 Detours 94 Understanding Numbers and Other Abstract Concepts 95 Self-Recognition and Perspective Taking 97 Chapter 6 Physiological Analysis—Nerve Cells and Behavior 99 Concepts from Cellular Neurobiology 100 Types of Neurons and Their Jobs 100 The Message of a Neuron 101 Ions, Membrane Permeability, and Behavior 104 Behavioral Change and Synaptic Transmission 104 The Structure of the Synapse 104 Integration 106 Specializations for Perception of Biologically Relevant Stimuli—Sensory Processing 107 Processing of Sensory Information for Sound Localization 108 Predators and Prey: The Neuroethology of Life-and- Death Struggles 109 Processing in the Central Nervous System 113 Brain Changes Underlying Behavioral Change 113 Social Behavior Network 120 Responding—Motor Systems 122 Neural Control in Motor Systems 122 Locust Flight 122 Chapter 7 Physiological Analysis of Behavior—The Endocrine System 127 The Endocrine System 129 Endocrine Glands and Hormones 129 Hormonal Versus Neural Communication 129 Types of Hormones and Their Modes of Action 130 How Hormones Influence Behavior 132 Effects on Sensation and Perception 132 Effects on Development and Activity of the Central Nervous System 133 Effects on Muscles 133 Methods of Studying Hormone–Behavior Relationships 135 Interventional Studies 135 Correlational Studies 137 Organizational and Activational Effects of Hormones 139 Defining the Dichotomy 139 Sex Differences in the Behavior of Norway Rats 139 Individual Differences in the Behavior of Male Tree Lizards 141 Questioning the Dichotomy 141 The Dynamic Relationship Between Hormones and Behavior 142 A Reciprocal Relationship 142 Hormonal Suppression of Behavior 142 Interactions Between Hormones, Behavior, and the Environment 142 Adjusting to the Harshness and Predictability of the Physical Environment 143 Adjusting to Onlookers in the Social Environment 145 A Detailed Look at the Hormonal Basis of Selected Behaviors 146 Helping at the Nest 146 Scent-Marking 147 Migrating 148 Chapter 8 The Development of Behavior 151 Influences on Behavioral Development 152 Development of the Nervous System 152 Development of Nonneural Structures 155 Hormonal Milieu 155 Physical Characteristics of the Environment 157 Experience Through Play 158 The Concept of Sensitive Periods 159 Changing Terminology—From Critical Periods to Sensitive Periods 159 Timing of Sensitive Periods 160 Multiple Sensitive Periods 161 Some Examples of Sensitive Periods in Behavioral Development 161 Pulling It All Together—The Development of Bird Song 171 Genetic, Hormonal, and Neural Control of Song 172 Role of Learning in Song Development 174 Sensitive Periods in Song Learning 176 Own-Species Bias in Song Learning 178 Social Factors and Song Development 178 A Diversity of Song Learning Strategies 179 Developmental Homeostasis 180 Rehabilitation of Chimpanzees After Long-Term Isolation 180 Part 2 Survival Chapter 9 Biological Clocks 185 Defining Properties of Clock-Controlled Rhythms 186 Persistence in Constant Conditions 186 Entrainment by Environmental Cycles 187 Temperature Compensation 188 Rhythmic Behavior 189 Daily Rhythms 189 Lunar Day Rhythms 189 Semilunar Rhythms 189 Monthly Rhythms 190 Annual Rhythms 191 The Clock Versus the Hands of the Clock 192 Advantages of Clock-Controlled Behavior 192 Anticipation of Environmental Change 192 Synchronization of a Behavior with an Event That Cannot Be Sensed Directly 193 Continuous Measurement of Time 194 Adaptiveness of Biological Clocks 194 Organization of Circadian Systems 194 Multiple Clocks 195 Coordination of Circadian Timing 195 Human Implications of Circadian Rhythms 200 Jet Lag 200 Human Health 200 Chapter 10 Mechanisms of Orientation and Navigation 203 Levels of Navigational Ability 204 Piloting 204 Compass Orientation 204 True Navigation 206 Multiplicity of Orientation Cues 206 Visual Cues 206 Landmarks 206 Sun Compass 207 Star Compass 211 Polarized Light and Orientation 212 Magnetic Cues 214 Cues from the Earth’s Magnetic Field 214 Directional Information from the Earth’s Magnetic Field: A Magnetic Compass 217 Positional Information from the Earth’s Magnetic Field: A Magnetic Map? 218 Magnetoreception 222 Chemical Cues 224 Olfaction and Salmon Homing 224 Olfaction and Pigeon Homing 226 Electrical Cues and Electrolocation 229 Chapter 11 The Ecology and Evolution of Spatial Distribution 233 Remaining at Home Versus Leaving 234 Costs and Benefits of Natal Philopatry 234 Costs and Benefits of Natal Dispersal 235 Sex Biases in Natal Dispersal 238 Natal Dispersal and Conservation Biology 240 Habitat Selection 241 Indicators of Habitat Quality 241 Search Tactics 243 Effects of Natal Experience 243 Habitat Selection and Conservation Biology 244 Migration 245 Costs of Migration 246 Benefits of Migration 248 Migration and Conservation Biology 251 Chapter 12 Foraging Behavior 253 Obtaining Food 253 Suspension Feeding 254 Omnivory 254 Herbivory 254 Carnivory 256 Adaptations for Detecting Prey 259 Optimal Foraging 264 Diet Selection: A Simple Model 265 Deciding When to Leave a Patch: The Marginal Value Theorem 267 Adding Complexity and Realism 268 The Utility of Models 274 Chapter 13 Antipredator Behavior 275 Camouflage 277 Coloration Matching the Visual Background 277 Disruptive Coloration 280 Countershading 282 Transparency 283 Masquerade 284 Other Functions of Color 284 Polymorphism 285 Warning Coloration 287 Batesian Mimicry 289 Diverting Coloration, Structures, and Behavior 290 False Heads 290 Autotomy 291 Feigning Injury or Death 292 Intimidation and Fighting Back 294 Enhancement of Body Size and Display of Weaponry 294 Eyespots 294 Chemical Repellents 296 Pronouncement of Vigilance 297 Group Defense 298 Alarm Signals 298 Improved Detection 299 Dilution Effect 299 Selfish Herd 300 Confusion Effect 301 Mobbing 301 Maintenance of Antipredator Behavior 302 Part 3 Interactions Between Individuals Chapter 14 Reproductive Behavior 305 Sexual Selection: Historical and Theoretical Background 306 Explanations for Sex Differences in Reproductive Behavior 306 Revisiting the Ideas of Bateman 308 Intrasexual Selection—Competition for Mates 309 Adaptations That Help a Male Secure Copulations 309 Adaptations That Favor the Use of a Male’s Sperm 313 Sexual Interference: Decreasing the Reproductive Success of Rival Males 318 Intersexual Selection—Mate Choice 320 Criteria by Which Females Choose Mates 320 Origin and Maintenance of Mate-Choice Preferences 327 Cryptic Female Choice 329 Sexual Conflict 330 Chapter 15 Parental Care and Mating Systems 333 Parental Care 334 Conflicts Among Family Members Over Parental Investment 334 Some Factors That Influence the Allocation of Parental Resources 336 Overall Patterns of Parental Care 339 Dispensing with Parental Care—Brood Parasitism 343 Mating Systems 345 Classifying Mating Systems 345 Monogamy 346 Polygyny 349 Polyandry 353 Chapter 16 Communication: Channels and Functions 355 The Definition of Communication 356 Channels for Communication 356 Vision 356 Audition 358 Substrate Vibrations 361 Chemical Senses 361 Touch 364 Electrical Fields 365 Multimodal Communication 366 Functions of Communication 367 Species Recognition 368 Mate Attraction 369 Courtship and Mating 369 Maintaining Social Bonds 372 Alarm 372 Aggregation 373 Agonistic Encounters 374 Communication about Resources: A Case Study 374 Chapter 17 The Evolution of Communication 381 The Changing Views of Communication 381 Sharing Information 381 Manipulating Others 382 Signals and Honesty 382 When are Honest Signals Likely? 382 When are Dishonest Signals Likely? 386 Can Honest and Dishonest Signals Coexist? 386 The Evolutionary Origins of Signals 387 Ritualization 387 Receiver-Bias Mechanisms 390 Selective Forces That Shape Signals 391 Characteristics of the Sender 391 Characteristics of the Environment 392 Characteristics of the Receiver 395 Language and Apes 396 What is Language? 396 Ape Language Studies 397 Communication and Animal Cognition 401 Chapter 18 Conflict 405 Aggression and Conflict 406 Why Do Animals Fight? 406 An Evolutionary View of Conflict 406 The Evolution of Fighting Behavior 406 Using Game Theory to Understand the Evolution of Conflict 406 Asymmetries in Contests 410 Conflict Among Group Members 413 How Dominance is Determined 413 The Benefits of Being Dominant 413 The Benefits of Being Subordinate 414 Conflict Over Space 415 Home Ranges, Core Areas, and Territories 415 The Ideal Free Distribution and Space Use 415 The Economics of Holding a Territory 416 The Economics of Territory Size 416 Strategies for Reducing the Cost of Territorial Defense 417 A Proximate View of Conflict 419 Aggression and Testosterone 419 Stress, Aggression, and Dominance 420