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of animal welfare is considered again in Chapter 10.
SUMMARY AND CONCLUSIONS
Motivation refers to the immediate causes of behavior, that is, to Tinbergen’s causal question. All behavior is caused by the action of a combination of internal and external causal factors, some of which have very specific effects on behavior and others more general effects. Stimuli can release, direct, inhibit, and prime behavior. These effects all depend on the internal state of the animal, which is controlled by hormones and other substances and by the intrinsic activity of the nervous system. The “psychohydraulic” model of behavior proposed by Lorenz provides a useful analogy for understanding how all these factors interact with each other. In general, causal factors for more than one behavior system are present at the same time. Sometimes the system with the strongest causal factors inhibits all the other systems, but most of the time animals engage in some type of ambivalent, redirected, or displacement behavior.
Emotions are the subjective aspect of strongly activated behavior systems. They can be studied in animals using Tinbergen’s motivation analysis (p. 62). Operant training techniques can be used to assess animal welfare.
FURTHER READING
Amplification of many of the ideas in this chapter can be found in The Study of Behavior (Hogan 2017). An Introduction to Behavioral Endocrinology (Nelson 2016) provides an excellent source for details on the physiological control of most of the behavior systems mentioned in this chapter. Toates’ (1986) book Motivational Systems presents a review of behavior systems with an emphasis on control theory variables, and Enquist and Ghirlanda (2005) discuss behavior systems from a neural network point of view. Lorenz’ (1966) controversial, but entertaining, book On Aggression gives his views on how ethological concepts can be applied to human behavior. James’ (1890) chapter on emotion should be read by anyone interested in the subject.
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