Behavioral
Ecology
,H0: Overview of the course
Learning outcomes – 6 credits
• Students acquire a thorough knowledge of topic research in behavioral ecology and ethology
thereby building on related disciplines such as ecology genetics and physiology.
• The student is able to understand recent research papers in the field of behavioral ecology
and can develop a research project around a given hypothesis based on this.
• The student has the expertise to plan, execute, analyze and critically interpret a behavioral
experiment in an international research team, thereby takes a responsible role and is able to
solve problems associated with the research project.
Course content
Professor Dr. Robby Stoks
1. An evolutionary approach to animal behavior
2. Understanding the proximate and ultimate causes of bird song
3. The development of behavior
4. The control of behavior: neural mechanisms
5. The organization of behavior: neurons and hormones
6. Behavioral adaptations for survival
7. Evolution of feeding behavior
Professor Dr. Tom Wenseleers
8. The evolution of communication
9. The evolution of reproductive behavior
10. The evolution of mating systems
11. The evolution of parental care
12. The evolution of social behavior
13. The evolution of human behavior
The exam
The exam will be a written exam:
• Written exam: 16/20 (8/20 per professor)
• Practicals: 4/20
,H1: AN INTRODUCTION TO ANIMAL BEHAVIOR ...............................................................................................................1
1.1 AN EVOLUTIONARY APPROACH TO ANIMAL BEHAVIOR ................................................................................................................ 1
1.1.1 Natural selection and the evolution of behavior .................................................................................................. 1
1.1.2 The cost-benefit approach to behavioral biology ................................................................................................. 2
1.2 BEHAVIORAL ECOLOGY ........................................................................................................................................................ 3
1.2.1 Defining behavior ................................................................................................................................................. 3
1.2.2 Approaches to studying behavior ......................................................................................................................... 5
1.2.3 Group selection and Darwinian puzzles ................................................................................................................ 5
1.2.4 The science of animal behavior ............................................................................................................................ 6
H2: THE DEVELOPMENT OF BEHAVIOR .............................................................................................................................7
2.1 BEHAVIOR REQUIRES GENES AND THE ENVIRONMENT ................................................................................................................ 7
2.1.1 Example: development of bee behavior ............................................................................................................... 7
2.1.2 Nature vs nurture controversy .............................................................................................................................. 8
2.2 ENVIRONMENTAL DIFFERENCES CAN CAUSE BEHAVIORAL DIFFERENCES .......................................................................................... 8
2.3 GENETIC DIFFERENCES CAN CAUSE BEHAVIORAL DIFFERENCES ..................................................................................................... 9
2.3.1 Common garden experiments............................................................................................................................... 9
2.3.2 Gene knockout experiments ............................................................................................................................... 10
2.3.3 Cross-breeding experiments ............................................................................................................................... 10
2.3.4 DNA damage experiments .................................................................................................................................. 11
2.3.5 Reciprocal transplant experiments ..................................................................................................................... 11
2.3.6 Artificial selection experiments ........................................................................................................................... 12
2.3.7 Inbred lines ......................................................................................................................................................... 12
2.3.8 Quantitative genetic experiments....................................................................................................................... 12
2.4 LEARNING ...................................................................................................................................................................... 14
2.4.1 Different types of learning mechanisms ............................................................................................................. 14
2.4.2 The adaptive value of learning ........................................................................................................................... 18
2.4.3 Costs and benefits of learning ............................................................................................................................ 18
H3: THE NEURAL BASIS OF BEHAVIOR ............................................................................................................................ 19
3.1 RESPONDING TO STIMULI .................................................................................................................................................. 19
3.1.1 Complex responses to simple stimuli: FAPs ......................................................................................................... 19
3.1.2 Neural control of escape behavior: the bat-moth arms race .............................................................................. 20
3.2 NEURAL COMMAND AND CONTROL ..................................................................................................................................... 23
3.2.1 Stimulus overload and behavioral prioritization ................................................................................................. 23
3.2.2 Central pattern generators (CPGs) to orchestrate complex behaviors ............................................................... 23
3.2.3 The proximate basis of stimulus filtering ............................................................................................................ 24
3.2.4 Sensory tuning .................................................................................................................................................... 24
H4: THE PHYSIOLOGICAL BASIS OF BEHAVIOR ................................................................................................................ 25
4.1 NEURAL COMMAND CENTERS: THE BASIS OF BEHAVIORAL CONTROL ........................................................................................... 25
4.1.1 Case study: decision making in the praying mantis (Roeder’s experiment) ........................................................ 25
4.2 ENDOGENOUS CLOCKS AND BEHAVIORAL RHYTHMS................................................................................................................. 26
4.2.1 Hypotheses explaining daily behavioral changes ............................................................................................... 26
4.2.2 The neurobiology of circadian timing ................................................................................................................. 26
4.2.3 The genetics of circadian timing ......................................................................................................................... 27
4.2.4 The physiology of circadian timing ..................................................................................................................... 27
4.2.5 Seasonal and annual rhythms ............................................................................................................................ 28
4.2.6 Cues that entrain cycles of behavior ................................................................................................................... 28
4.3 HORMONAL MECHANISMS UNDERLYING BEHAVIORAL CHANGE .................................................................................................. 29
4.3.1 Hormones and reproduction ............................................................................................................................... 30
4.3.2 The cost of hormonal regulation ........................................................................................................................ 32
4.3.3 The challenge hypothesis (Wingfield et al. 1990) ............................................................................................... 33
, H5: BEHAVIORAL ADAPTATIONS FOR SURVIVAL ............................................................................................................. 35
5.1 ANTI-PREDATOR BEHAVIORS AND PREDATOR AVOIDANCE.......................................................................................................... 35
5.1.1 Strategy 1: decreasing detection by predators ................................................................................................... 35
5.1.2 Strategy 2: preventing attack or capture ............................................................................................................ 38
5.1.3 Strategy 3: behavior once captured .................................................................................................................... 44
5.2 VARIABILITY IN ANTIPREDATOR BEHAVIOR ............................................................................................................................. 44
5.2.1 Threat-sensitivity hypothesis .............................................................................................................................. 44
5.2.2 Covariation with morphological traits ................................................................................................................ 45
5.2.3 Predation risk allocation hypothesis ................................................................................................................... 45
5.3 ABSENCE OF EFFECTIVE ANTIPREDATOR BEHAVIORS ................................................................................................................. 46
H6: EVOLUTION OF FEEDING BEHAVIOR ........................................................................................................................ 47
6.1 DIET SELECTION MODELS ................................................................................................................................................... 47
6.1.1 Case study: Northwestern crows dropping whelks ............................................................................................. 47
6.1.2 Case study: Eurasian oystercatcher and mussel choice ...................................................................................... 48
6.1.3 Diet choice: Charnov’s optimal diet model ......................................................................................................... 49
6.2 PATCH MODELS ............................................................................................................................................................... 52
6.2.1 Charnov’s marginal value theorem..................................................................................................................... 53
6.2.2 Empirical test of the MVT ................................................................................................................................... 55
6.2.3 Game theory and feeding behavior .................................................................................................................... 55
6.3 DARWINIAN PUZZLES IN FEEDING BEHAVIOR .......................................................................................................................... 56
H7: BEHAVIORAL & PACE-OF-LIFE SYNDROME ............................................................................................................... 58
7.1 BEHAVIORAL SYNDROMES & ANIMAL PERSONALITY................................................................................................................. 58
7.1.1 Personality traits ................................................................................................................................................. 59
7.1.2 Four levels of explanation (Tinbergen’s questions) ............................................................................................. 61
7.1.3 Strategies to study behavioral syndromes .......................................................................................................... 62
7.2 THE PACE-OF-LIFE SYNDROME (POLS)................................................................................................................................ 63
7.2.1 POLS at species level ........................................................................................................................................... 63
7.2.2 POLS at the population level ............................................................................................................................... 64
7.2.3 POLS at the individual level................................................................................................................................. 67
7.2.4 Evolutionary and ecological implications of fast POLS ....................................................................................... 68
H8: PRINCIPLES OF COMMUNICATION ........................................................................................................................... 69
8.1 COMMUNICATION AND ANIMAL SIGNALS .............................................................................................................................. 69
8.2 THE EVOLUTIONARY ORIGIN OF SIGNALS ............................................................................................................................... 69
8.2.1 Co-option and pre-adaptation ............................................................................................................................ 70
8.2.2 Sensory exploitation and preexisting biases ....................................................................................................... 70
8.2.3 Ambiguity: trait or bias first? .............................................................................................................................. 71
8.3 MAINTAINING SIGNAL HONESTY .......................................................................................................................................... 71
8.3.1 Honest communication due to common interest ............................................................................................... 72
8.3.2 Honest communication due to costs ................................................................................................................... 75
8.3.3 Honest communication due to uncheatable indices of quality ........................................................................... 77
8.4 DECEITFUL SIGNALING ...................................................................................................................................................... 78
8.5 SIGNAL EXPLOITATION AND EAVESDROPPING.......................................................................................................................... 80
H9: EVOLUTION OF REPRODUCTIVE BEHAVIOR .............................................................................................................. 81
9.1 EVOLUTIONARY ORIGIN OF TWO SEXES: SMALL AND LARGE GAMETES.......................................................................................... 81
9.2 THE EVOLUTION OF TYPICAL SEX ROLES ................................................................................................................................. 82
9.2.1 From gametes to gendered behavior.................................................................................................................. 82
9.2.2 Typical sex roles in nature ................................................................................................................................... 82
9.2.3 Sexual conflict and arms race ............................................................................................................................. 83
9.2.4 When sex roles reverse ....................................................................................................................................... 83
9.2.5 Conflicts over sex roles in hermaphrodites ......................................................................................................... 84
Ecology
,H0: Overview of the course
Learning outcomes – 6 credits
• Students acquire a thorough knowledge of topic research in behavioral ecology and ethology
thereby building on related disciplines such as ecology genetics and physiology.
• The student is able to understand recent research papers in the field of behavioral ecology
and can develop a research project around a given hypothesis based on this.
• The student has the expertise to plan, execute, analyze and critically interpret a behavioral
experiment in an international research team, thereby takes a responsible role and is able to
solve problems associated with the research project.
Course content
Professor Dr. Robby Stoks
1. An evolutionary approach to animal behavior
2. Understanding the proximate and ultimate causes of bird song
3. The development of behavior
4. The control of behavior: neural mechanisms
5. The organization of behavior: neurons and hormones
6. Behavioral adaptations for survival
7. Evolution of feeding behavior
Professor Dr. Tom Wenseleers
8. The evolution of communication
9. The evolution of reproductive behavior
10. The evolution of mating systems
11. The evolution of parental care
12. The evolution of social behavior
13. The evolution of human behavior
The exam
The exam will be a written exam:
• Written exam: 16/20 (8/20 per professor)
• Practicals: 4/20
,H1: AN INTRODUCTION TO ANIMAL BEHAVIOR ...............................................................................................................1
1.1 AN EVOLUTIONARY APPROACH TO ANIMAL BEHAVIOR ................................................................................................................ 1
1.1.1 Natural selection and the evolution of behavior .................................................................................................. 1
1.1.2 The cost-benefit approach to behavioral biology ................................................................................................. 2
1.2 BEHAVIORAL ECOLOGY ........................................................................................................................................................ 3
1.2.1 Defining behavior ................................................................................................................................................. 3
1.2.2 Approaches to studying behavior ......................................................................................................................... 5
1.2.3 Group selection and Darwinian puzzles ................................................................................................................ 5
1.2.4 The science of animal behavior ............................................................................................................................ 6
H2: THE DEVELOPMENT OF BEHAVIOR .............................................................................................................................7
2.1 BEHAVIOR REQUIRES GENES AND THE ENVIRONMENT ................................................................................................................ 7
2.1.1 Example: development of bee behavior ............................................................................................................... 7
2.1.2 Nature vs nurture controversy .............................................................................................................................. 8
2.2 ENVIRONMENTAL DIFFERENCES CAN CAUSE BEHAVIORAL DIFFERENCES .......................................................................................... 8
2.3 GENETIC DIFFERENCES CAN CAUSE BEHAVIORAL DIFFERENCES ..................................................................................................... 9
2.3.1 Common garden experiments............................................................................................................................... 9
2.3.2 Gene knockout experiments ............................................................................................................................... 10
2.3.3 Cross-breeding experiments ............................................................................................................................... 10
2.3.4 DNA damage experiments .................................................................................................................................. 11
2.3.5 Reciprocal transplant experiments ..................................................................................................................... 11
2.3.6 Artificial selection experiments ........................................................................................................................... 12
2.3.7 Inbred lines ......................................................................................................................................................... 12
2.3.8 Quantitative genetic experiments....................................................................................................................... 12
2.4 LEARNING ...................................................................................................................................................................... 14
2.4.1 Different types of learning mechanisms ............................................................................................................. 14
2.4.2 The adaptive value of learning ........................................................................................................................... 18
2.4.3 Costs and benefits of learning ............................................................................................................................ 18
H3: THE NEURAL BASIS OF BEHAVIOR ............................................................................................................................ 19
3.1 RESPONDING TO STIMULI .................................................................................................................................................. 19
3.1.1 Complex responses to simple stimuli: FAPs ......................................................................................................... 19
3.1.2 Neural control of escape behavior: the bat-moth arms race .............................................................................. 20
3.2 NEURAL COMMAND AND CONTROL ..................................................................................................................................... 23
3.2.1 Stimulus overload and behavioral prioritization ................................................................................................. 23
3.2.2 Central pattern generators (CPGs) to orchestrate complex behaviors ............................................................... 23
3.2.3 The proximate basis of stimulus filtering ............................................................................................................ 24
3.2.4 Sensory tuning .................................................................................................................................................... 24
H4: THE PHYSIOLOGICAL BASIS OF BEHAVIOR ................................................................................................................ 25
4.1 NEURAL COMMAND CENTERS: THE BASIS OF BEHAVIORAL CONTROL ........................................................................................... 25
4.1.1 Case study: decision making in the praying mantis (Roeder’s experiment) ........................................................ 25
4.2 ENDOGENOUS CLOCKS AND BEHAVIORAL RHYTHMS................................................................................................................. 26
4.2.1 Hypotheses explaining daily behavioral changes ............................................................................................... 26
4.2.2 The neurobiology of circadian timing ................................................................................................................. 26
4.2.3 The genetics of circadian timing ......................................................................................................................... 27
4.2.4 The physiology of circadian timing ..................................................................................................................... 27
4.2.5 Seasonal and annual rhythms ............................................................................................................................ 28
4.2.6 Cues that entrain cycles of behavior ................................................................................................................... 28
4.3 HORMONAL MECHANISMS UNDERLYING BEHAVIORAL CHANGE .................................................................................................. 29
4.3.1 Hormones and reproduction ............................................................................................................................... 30
4.3.2 The cost of hormonal regulation ........................................................................................................................ 32
4.3.3 The challenge hypothesis (Wingfield et al. 1990) ............................................................................................... 33
, H5: BEHAVIORAL ADAPTATIONS FOR SURVIVAL ............................................................................................................. 35
5.1 ANTI-PREDATOR BEHAVIORS AND PREDATOR AVOIDANCE.......................................................................................................... 35
5.1.1 Strategy 1: decreasing detection by predators ................................................................................................... 35
5.1.2 Strategy 2: preventing attack or capture ............................................................................................................ 38
5.1.3 Strategy 3: behavior once captured .................................................................................................................... 44
5.2 VARIABILITY IN ANTIPREDATOR BEHAVIOR ............................................................................................................................. 44
5.2.1 Threat-sensitivity hypothesis .............................................................................................................................. 44
5.2.2 Covariation with morphological traits ................................................................................................................ 45
5.2.3 Predation risk allocation hypothesis ................................................................................................................... 45
5.3 ABSENCE OF EFFECTIVE ANTIPREDATOR BEHAVIORS ................................................................................................................. 46
H6: EVOLUTION OF FEEDING BEHAVIOR ........................................................................................................................ 47
6.1 DIET SELECTION MODELS ................................................................................................................................................... 47
6.1.1 Case study: Northwestern crows dropping whelks ............................................................................................. 47
6.1.2 Case study: Eurasian oystercatcher and mussel choice ...................................................................................... 48
6.1.3 Diet choice: Charnov’s optimal diet model ......................................................................................................... 49
6.2 PATCH MODELS ............................................................................................................................................................... 52
6.2.1 Charnov’s marginal value theorem..................................................................................................................... 53
6.2.2 Empirical test of the MVT ................................................................................................................................... 55
6.2.3 Game theory and feeding behavior .................................................................................................................... 55
6.3 DARWINIAN PUZZLES IN FEEDING BEHAVIOR .......................................................................................................................... 56
H7: BEHAVIORAL & PACE-OF-LIFE SYNDROME ............................................................................................................... 58
7.1 BEHAVIORAL SYNDROMES & ANIMAL PERSONALITY................................................................................................................. 58
7.1.1 Personality traits ................................................................................................................................................. 59
7.1.2 Four levels of explanation (Tinbergen’s questions) ............................................................................................. 61
7.1.3 Strategies to study behavioral syndromes .......................................................................................................... 62
7.2 THE PACE-OF-LIFE SYNDROME (POLS)................................................................................................................................ 63
7.2.1 POLS at species level ........................................................................................................................................... 63
7.2.2 POLS at the population level ............................................................................................................................... 64
7.2.3 POLS at the individual level................................................................................................................................. 67
7.2.4 Evolutionary and ecological implications of fast POLS ....................................................................................... 68
H8: PRINCIPLES OF COMMUNICATION ........................................................................................................................... 69
8.1 COMMUNICATION AND ANIMAL SIGNALS .............................................................................................................................. 69
8.2 THE EVOLUTIONARY ORIGIN OF SIGNALS ............................................................................................................................... 69
8.2.1 Co-option and pre-adaptation ............................................................................................................................ 70
8.2.2 Sensory exploitation and preexisting biases ....................................................................................................... 70
8.2.3 Ambiguity: trait or bias first? .............................................................................................................................. 71
8.3 MAINTAINING SIGNAL HONESTY .......................................................................................................................................... 71
8.3.1 Honest communication due to common interest ............................................................................................... 72
8.3.2 Honest communication due to costs ................................................................................................................... 75
8.3.3 Honest communication due to uncheatable indices of quality ........................................................................... 77
8.4 DECEITFUL SIGNALING ...................................................................................................................................................... 78
8.5 SIGNAL EXPLOITATION AND EAVESDROPPING.......................................................................................................................... 80
H9: EVOLUTION OF REPRODUCTIVE BEHAVIOR .............................................................................................................. 81
9.1 EVOLUTIONARY ORIGIN OF TWO SEXES: SMALL AND LARGE GAMETES.......................................................................................... 81
9.2 THE EVOLUTION OF TYPICAL SEX ROLES ................................................................................................................................. 82
9.2.1 From gametes to gendered behavior.................................................................................................................. 82
9.2.2 Typical sex roles in nature ................................................................................................................................... 82
9.2.3 Sexual conflict and arms race ............................................................................................................................. 83
9.2.4 When sex roles reverse ....................................................................................................................................... 83
9.2.5 Conflicts over sex roles in hermaphrodites ......................................................................................................... 84
