On the origin of allostasis and stress-induced pathology in farm animals: Celebrating Darwin’s legacy

Abstract

Darwin's largest contribution to science is without doubt the mechanism of natural selection, an evolutionary game with players, strategies, and pay-offs. Game theory, which attempts to mathematically capture behaviour in situations where an organism's success in making choices depends on the choices of others, is not only important for economists, but also for biologists, veterinarians and other scientists, as it increases understanding of why individual differences exist. John Maynard Smith showed that the success of an individual's behaviour often depends on others and his Hawk-Dove model is one of the best known examples of game theory: the 'hawk' initiates aggressive behaviour (not stopping until injured or until the opponent backs down); the 'dove' retreats immediately if the opponent initiates aggressive behaviour and will not fight under any circumstances. Simultaneous hawkish behaviour has the worst pay-off for both players, whereas hawkish behaviour with a dove opponent has the best pay-off. Maynard Smith showed that natural selection will work towards an evolutionarily stable strategy that, when used by an entire population, is resistant to invasion by new mutant strategies. Thus, natural selection actually favours a particular ratio of aggressive hawkish and non-aggressive dovish behaviours in order to maintain a balance of different characteristics in the population. Natural selection has sculpted physiology and behaviour differently in hawks and doves, each in their own way so as to maintain stability of the internal environment through change--a process which is defined as allostasis. In the short term, allostasis has benefits, but in the long run it produces costs. Farm animals have been genetically selected by man for increased product quantity and quality, such as increased muscle volume, lean meat and egg shell quality, accompanied by altered steroid balance (such as more testosterone and less corticosteroids) and lower brain monoamine concentrations (serotonin and dopamine). It is hypothesised that such genetic selection results in the production of farm animals that prefer the hawk behavioural strategy. There is a growing body of evidence that hawk-like animals (such as laying hens and pigs) are more vulnerable to the development of increased impulsivity and compulsivity (stereotypies) as well as violent behaviour.