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
