Perspectives on Animal Behavior

JUDITH GOODENOUGH, from the Department of Biology at the University of Massachusetts, has studied biological rhythms in creatures from planaria to finches to deer mice, and even in the green alga Chlamydomonas. BETTY MCGUIRE is from the Department of Ecology and Evolutionary Biology at Cornell University. Her research focuses on parental behavior, reproduction, and ecology of small mammals, and she occasionally dabbles in work with larger domestic animals such as dogs and horses. ELIZABETH JAKOB, from the Department of Psychology at the University of Massachusetts, studies the behavior of spiders, asking questions about their learning, perception, and interactions with conspecifics and with other species. She has carried out field projects in California, Mexico, Massachusetts, and Maine.

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