Although low-ranking animals are commonly stressed, they are not necessarily depressed. Moreover, socially anxious people and low self-esteem people can feel inferior and behave submissively but may not be depressed (Gilbert, 2001b). Indeed, Brown and Harris (1978) suggest that low self-esteem is a vulnerability factor that can fuel depression in the face of provoking agents. There are, in fact, two other ingredients that may need to be present for depression: defeat and entrapment. Price (1972, Price & Sloman, 1987) first argued that depression might be an evolved response to social defeats where either there is a loss in social standing (rank) and/or an animal is being harassed from above and cannot escape to somewhere else or try something else. He argued that under such conditions the best response may be a form of demobilisation which stops the animal from engaging in behaviour that elicits further attacks, and signals to potential attackers that 'one is defeated and out of action'. Price originally called this kind of demobilisation a yielding subroutine of a subordinate strategy set, where the downregulation of PA forces the animal to yield and stay out of action for a while (Price et al., 1994). However, we have subsequently called it an involuntary defeat strategy (see Sloman & Gilbert, 2000). One reason for this is that socially anxious and low self-esteem people may use some subordinate strategies, and they may yield (be non-assertive) in many contexts, but it is the involuntary defeat strategy that may underpin some depressions.

Social defeats in animals have major physiological effects and can produce quite rapid and profound effects; that is, they set the system in a new state. Levitan et al. (2000) have reviewed data on the biology of submissive and defeat states, finding that defeated and/or harassed subordinates are chronically stressed, with increased HPA activation (Ray & Sapolsky, 1992; Sapolsky, 1989). Laboratory studies on rodents show that defeat experiences reliably result in physiological and behavioural consequences, including a decrease in offensive aggression, an increase in defensive responses, decreases in subsequent exploratory behaviour, increases in freezing, weight loss, reduction of appetitive behaviours, and disruption of escape learning (Gilbert, 2000a). Social defeats are also associated with retardation (Sloman et al., 2003). Importantly, however, within populations, there are individual differences in strategies (perhaps gene or early history related) for coping with down-rank agonism, rejection, and social defeats. For example, Von Holst (1986) studied tree shrews and found that defeated losers adopted one of two different strategies. Some continued with activities but in a timid and cautious way. They showed an elevated stress response and elevated tyrosine hydroxylase activity. However, other defeated animals became seriously demobilised, with greatly elevated cortisol responses and reductions in tyrosine hydroxylase. They died within 14 days of the confrontation. Even separating victor and loser by a wire mesh did not save these animals.

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