The Study of Animal Behaviour

1 Introduction.- 1.1 The ethological approach to the study of behaviour.- 1.2 A brief outline of classical ethological theory.- 1.3 The modern study of animal behaviour.- 1.4 An outline of the book.- 2 The description and measurement of behaviour.- 2.1 Describing behaviour by its function.- 2.2 Describing behaviour by its form.- 2.2.1 Behaviour described as a continuous process.- 2.2.2 Behaviour described as a series of discrete events.- 2.3 Describing and measuring the relationship between an animal and its environment.- 2.3.1 Describing the way a single animal uses space.- 2.3.2 Describing the relationship between an animal and neighbouring conspecifics.- 2.4 What is the point of all this sophisticated analysis?.- 3 The study of the causes of behavioural change.- 3.1 What constitutes a causal explanation of behaviour?.- 3.2 The different kinds of causal explanation.- 3.3 Motivational models.- 3.3.1 Descriptive and explanatory models.- 3.3.2 General, theory-based and specific, data-based models.- 3.3.3 Deterministic and probabilistic models.- 3.3.4 Assessing models.- 3.4 Studying external influences on behaviour.- 3.4.1 Prey catching in toads; behavioural studies.- 3.4.2 Attack in Haplochromis burtoni.- 3.4.3 Incubation in herring gulls.- 3.4.4 Prey catching in toads; physiological studies.- 3.4.5 The role of external stimuli in the control of behaviour.- 3.5 Studying internal influences on behaviour.- 3.5.1 Identifying groups of causally related acts.- 3.5.2 The role of internal factors in the control of behaviour.- 3.6 What is the nature of the mechanisms which cause behavioural change?.- 3.6.1 A probabilistic model.- 3.6.2 A deterministic model.- 3.6.3 Modelling motivation as a hierarchy of decisions.- 3.6.4 Control theory models of motivation.- 3.6.5 Space-state models.- 3.7 Studying the physiological bases of behavioural change.- 3.7.1 Physiological explanations of complex behaviour.- 3.7.2 Alteration in behavioural theory as a result of physiological research.- 3.7.3 Identifying fruitful topics for physiological study by behavioural analysis.- 3.7.4 Ethological concepts and techniques can improve the quality of physiological research.- 3.7.5 Ethological ideas can help in the interpretation of physiological results.- 4 The development of behaviour.- 4.1 Problems with the instinct-learning dichotomy.- 4.1.1 Instincts are defined by exclusion.- 4.1.2 Interpreting deprivation experiments is not simple.- 4.1.3 Learning is not the only way the environment influences development of behaviour.- 4.1.4 The instinct-learning dichotomy confounds three distinct consequences of behavioural development.- 4.2 Why is the term innate still used?.- 4.3 Describing the ontogeny of behaviour.- 4.3.1 The emergence of recognizable behaviour patterns in young animals.- 4.3.2 The time of appearance of behaviour patterns during development.- 4.3.3 The development of motivational systems.- 4.3.4 Physiological changes accompanying behavioural development.- 4.4 Characterizing the factors which influence the development of behaviour.- 4.4.1 Deductions from descriptive studies.- 4.4.2 Experimental studies of behavioural development.- 4.4.3 Experimental studies of the development of song in passerines.- 4.4.4 A quantitative model of the development of social preferences in birds.- 4.5 Classifying the factors which influence the development of behaviour.- 4.6 Some general features of behavioural development.- 4.6.1 Sensitive periods.- 4.6.2 Constraints on learning.- 4.6.3 Functional aspects of behavioural development.- 5 The adaptive significance of behaviour.- 5.1 Sources of evidence about the adaptive significance of behaviour.- 5.1.1 Deduction from the theory of natural selection.- 5.1.2 The context in which behaviour is shown.- 5.1.3 The short-term consequences of behaviour.- 5.1.4 Comparisons between species and populations.- 5.1.5 Measuring the fitness of individuals which differ in their behaviour.- 5.1.6 Experimental manipulation of the consequences of behaviour.- 5.1.7 Deductions from the genetic architecture of behaviour.- 5.1.8 Models of the adaptive significance of behaviour.- 5.2 Difficulties in studying the adaptive significance of behaviour.- 5.2.1 Multiple benefits.- 5.2.2 Costs, benefits and optimality models.- 5.2.3 Frequency dependent costs and benefits; games theory.- 5.3 The state of the art.- 5.4 The adaptive significance of the way animals pattern their behavior in time.- 5.4.1 Rhythms of behaviour.- 5.4.2 The adaptive significance of behavioural rhythmicity.- 5.5 The adaptive significance of the way animals use space.- 5.5.1 Dispersal patterns.- 5.5.2 Deciding where to live.- 5.5.3 What to do where; optimal foraging theory.- 5.5.4 Problems with optimal foraging theory.- 5.5.5 Constraints on use of space resulting from the presence of conspecifics.- 5.5.6 Territorial behaviour.- 5.6 The adaptive significance of an animal’s aggressive responses.- 5.6.1 Games theory and the hawk-dove model.- 5.6.2 Weaknesses of the hawk-dove model.- 5.6.3 Games theory models in sexually reproducing species.- 5.6.4 Conflicts between doves; the War of Attrition model.- 5.6.5 Games with progressive escalation.- 5.6.6 Games with asymmetries between opponents.- 5.6.7 Games in which information is acquired during a fight.- 5.6.8 Games with non-random encounters.- 5.6.9 The contribution of games theory to our understanding of the adaptive significance of the way animals fight.- 5.7 The adaptive significance of an animal’s breeding habits; mating systems.- 5.7.1 Life history parameters.- 5.7.2 Questions about sex.- 5.7.3 Mating systems.- 5.8 Adaptive significance of behaviour accompanying mating.- 5.8.1 Identifying the correct species.- 5.8.2 Preventing mating with conspecifics of the wrong sex.- 5.8.3 Synchronizing the behaviour of potential mates.- 5.8.4 The impact ofintra-sexual competition for mates.- 5.8.5 The impact of selectiveness in potential mates.- 5.8.6 Alternative mating strategies.- 5.8.7 Darwin, natural selection and sexual selection.- 5.8.8 Conflicts of interest between the sexes.- 5.9 The adaptive significance of parental care.- 5.10 The adaptive significance of living in groups.- 5.10.1 Studies of the adaptive significance of group life.- 5.10.2 The benefits resulting from group life.- 5.10.3 The complex costs and benefits of group life; Caraco’s model (1979).- 5.11 Behaviour which cannot be explained by classic natural selection theory.- 5.11.1 An outline of the problem.- 5.11.2 Animals may ultimately benefit from their ‘altruistic’ behaviour.- 5.11.3 Group selection.- 5.11.4 Kin selection.- 5.11.5 Parental manipulation.- 5.11.6 Has altruism been explained?.- 5.12 Overview; sociobiology and behavioural ecology.- 5.12.1 Weaknesses in the sociobiological approach to behaviour.- 5.12.2 Reasons why behavioural differences may not be adaptive.- 6 The phylogeny of behaviour.- 6.1 Sources of evidence about the phylogeny of behaviour.- 6.1.1 The fossil record.- 6.1.2 Ontogeny.- 6.1.3 The behaviour of interspecific hybrids.- 6.1.4 Comparative studies.- 6.2 Some representative behavioural phylogenies.- 6.2.1 Tongue movements in reptiles.- 6.2.2 Predatory behaviour in gastropods.- 6.2.3 Aggressive interactions in fish.- 6.2.4 Hymenopteran social systems.- 6.3 Deriving general principles of behavioural evolution.- 6.3.1 Sound production in crickets.- 6.3.2 Courtship in peacocks.- 7 The role of behaviour in the evolutionary process.- 7.1 The behaviour of other animals as a major selective force.- 7.1.1 The behaviour of conspecifics.- 7.1.2 The behaviour of heterospecifics.- 7.2 Behaviour dictates the selection pressures to which an animal is exposed.- 7.3 The impact of behaviour on population structure.- 7.3.1 Behavioural influences on gene flow between populations.- 7.3.2 Behavioural influences on population size.- 7.3.3 Behavioural influences on panmixis.- 8 Behavioural genetics.- 8.1 The objectives of research into the inheritance of behaviour.- 8.2 Potential contributions of genetics to the study of animal behaviour.- 8.2.1 As an additional research tool in the analysis of behaviour.- 8.2.2 As a source of essential information about behaviour.- 8.3 Quantitative genetics.- 8.4 Techniques for identifying the genetic origin of a known behavioural difference.- 8.4.1 Crosses between behavioural variants.- 8.4.2 Selective breeding from extremes of a behavioural continuum.- 8.5 Screening known genetic variants for behavioural differences.- 8.5.1 Known single gene mutations which influence behaviour.- 8.5.2 Visible differences in chromosome structure with behavioural effects.- 8.5.3 Comparing naturally occurring, isolated populations.- 8.5.4 Correlating degree of relatedness and behavioural similarity.- 8.5.5 Inbred strains.- 8.5.6 Recombinant strains.- 8.6 Characterizing the precise behavioural effects of genetic differences.- 8.7 The mechanisms whereby genes influence behaviour.- 8.7.1 Effects on the structures necessary for performing the behaviour.- 8.7.2 Effects on muscles or neuromuscular junctions.- 8.7.3 Effects on the sense organs.- 8.7.4 Effects on the CNS.- 8.7.5 Effects on the endocrine system.- 8.7.6 Effects on some totally unknown mechanism.- 8.8 Genetic mosaics.- 8.9 Animal behaviour and behavioural genetics.- 9 Applied ethology.- 9.1 Clarification of terms; what is applied ethology?.- 9.2 Ways in which etiological research can be applied to practical problems.- 9.2.1 Direct application of the results of etiological research.- 9.2.2 Using the ideas and concepts of ethology.- 9.2.3 Using etiological techniques.- 9.3 Pest control.- 9.4 Increasing the productivity of commercially important species.- 9.4.1 Identifying suitable species for exploitation.- 9.4.2 Selection for desired behavioural attributes.- 9.4.3 Controlling dispersal and movement.- 9.4.4 Promoting survival.- 9.4.5 Promoting growth.- 9.4.6 Promoting successful reproduction.- 9.5 Animal welfare.- 9.5.1 Identifying suffering in animals.- 9.5.2 Alleviating animal suffering.- 9.6 Conservation.- 9.7 Human behaviour.- 9.7.1 Applying ethological techniques to human behaviour.- 9.7.2 Ethological concepts as a source of hypotheses about human behaviour.- 9.7.3 Making direct use of the facts collected by ethologists.- 9.7.4 Sociobiology and human behaviour.- References.- Author index.- Species index.