dominance, submission, depression, anxiety

Lounge
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in case it’s not clear, pheromone usage is very closely tied to issues of social rank and dominance. so understanding this dynamic is important.

Delwart

TERRITORY, RANK AND MENTAL HEALTH
Appeasement displays at three levels of the triune brain: the history of an idea
Presented at a seminar of the Jean-Marie Delwart Foundation in Genval, Belgium in October, 2006, on:
“Can Ethology meet Psychiatric expectations?”.
About thirty years ago four psychiatrists, independently, started to think about mood and anxiety disorders from an ethological, functional, adaptational and evolutionary perspective.
Albert Demaret, who is an ornithologist as well as a psychiatrist, observed that his patients with elevated mood behaved rather like birds strutting and swaggering about on their own territories, whereas his depressed patients behaved like birds on another bird’s territory, hopping furtively about and not inclined to break into song (Demaret, 1971, 1979). He thought that if evolution had designed the contrasting social roles of territory owner and non-territory owner, then it should also have designed behaviours to fit those roles, and what could be better fitted to perform that function than elevated and depressed mood? Unfortunately Demaret did not publish in English, and so his ideas have not received the attention they deserve in the UK and US.
Russell Gardner, Jr., who was Professor and Chair of Psychiatry in North Dakota (and later became Harry K. Davis Professor of Psychiatry in Galveston, Texas), was impressed with the similarity between patients with elevated mood and normal leaders, and suggested that mania represented leadership behaviour occurring out of context (Gardner, 1982, 1988). His patients were communicating high rank signals in the absence of high rank; and likewise his depressed patients were communicating inappropriate low rank. He felt that these rank signals were of ancient evolutionary origin.
John Price had come under the influence of Niko Tinbergen while reading Psychology at Oxford, and later, while working in the Emergency Clinic at the Maudsley Hospital in London, he was impressed by the similarity between his depressed patients and the low ranking long-tailed macaque monkeys seen in a film shown by ethopharmacologist Michael Chance (Price, 1972, 1998). Looking back at those hen-pecked animals, he thought that if they could talk and were allowed access, they would be queuing up at the Emergency Clinic to complain of anxiety, depression and widespread aches and pains. It seemed obvious that if you had to have social hierarchies, you had to have behaviours appropriate to the different positions in the hierarchy that individuals came to adopt, and especially you had to have behaviours to mediate going up and down the hierarchy – and it seemed that elevated and depressed mood were “tailor-made” to contribute to the performance of that task. Literature describing hierarchies in non-human primates was scarce at this time, the most informative were those by Zuckerman (1932) and DeVore (1965).
Leon Sloman (1976, 1979, 2000) pointed out that neurotic depression and anxiety impair function, so that, to the extent that depression is the result of failure to achieve, the depressed person will achieve even less than he did before, resulting in a vicious circle or positive-feedback situation between depression and impairment of function, a state of maladaptation which would impair breeding success, compounded also by assortative mating for maladaptation. Moreover, the original causes of failure to achieve, which are likely to be related to small genetic differences in skill and intelligence, will be magnified by this positive-feedback process, thus accelerating the evolution of abilities including intelligence.
These ideas converged on the conclusion that the phenomena we were observing in the clinic were part of an evolved mechanism for creating and managing social asymmetry – the asymmetry between the owners of territories and non-owners, and the asymmetry between high-ranking individuals and low-ranking individuals. States of high and low mood appeared adapted not only to create asymmetry, but also to maintain asymmetry once established, and to reverse asymmetry in the cases of loss of rank or territory. It seemed likely that more intense mood states were required for the creation and reversal of asymmetry than for the maintenance of asymmetry.
After formulating these ideas, it came as a pleasant surprise to learn that they had already been anticipated by a Norwegian zoologist who was the first to describe social hierarchy (or peck order) in animals. Thorlief Schjelderup-Ebbe (1935) described a low-grade depression in his low-ranking hens and a severe, life-threatening depression in alpha birds who were deposed from their high status. It seemed unlikely that “neurotic” and “endogenous” depression had a separate phylogeny stretching back to the common ancestor of domestic hens and human beings, but rather that the very nature of hierarchies was likely to give rise to different mood states to subserve change in rank on the one hand and the maintenance of low rank on the other (Price, 1969, Price & Sloman, 1987).
It may seem inconceivable now, but at that time (in the 1970s) it was politically incorrect to write about social hierarchy. The buzz-word was “equality” and those who wrote about inequality were assumed to approve of it and were labelled as fascist. It therefore seemed more tactful to frame our ideas in terms of the behaviour which leads to the formation of territories and hierarchies; namely, ritual agonistic behaviour. Depression, and neuroticism, could be thought to have evolved as part of the yielding component of ritual agonistic behaviour, leading to low rank and loss of territory. There is a ritual quality to depression, which gives the impression to sufferers and observers that it is not “real” in some way, and that all the sufferer needs to do is to “snap out of it” or “pull himself together” and the whole thing would vanish like smoke. In this way we could see depression as being the result of ritual combat, so that the psychological incapacity of depression represents a ritual equivalent of the physical incapacity which may follow unritualised combat, or we could even say that it was a ritual equivalent of death. When this idea was put to some of our depressed patients, they replied that they did indeed feel “dead”.
These ideas depended on the plentiful observations of ethologists on ritual agonistic behaviour in a wide variety of species, and soon they were supported by the mathematical calculations of behavioural ecologists (Krebs & Davies, 1981, 1997; Maynard Smith, 1982), who showed that in the analysis of pairwise contests it was necessary to have both “hawk” genotypes who always escalated the contest and “dove” genotypes who always capitulated (or you could have mixed genotypes who played hawk on some occasions and dove on others). The idea of mutually incompatible alternative behavioural strategies was becoming popular in ecology, and it seemed appropriate to describe elevation and depression of mood as related, respectively, to escalating and de-escalating strategies, which had evolved in relation to ritual agonistic behaviour, but had become relevant to other and more diffuse types of competition, such as competition for prestige.
The match of escalation/de-escalation with elevation/depression of mood
Many of the features of mood disorders could be accounted for by changes in the three variables which are needed to predict winning and losing in pairwise contests: Resource Holding Potential (RHP), Resource Value (RV), and ownership. RHP is a self-concept representing fighting capacity, and in some complex way animals are able to compare their estimate of their own RHP with that of their rival, so avoiding unevenly matched contests. It seems likely that RHP is one of the forerunners of human self-esteem, and we know that self-esteem is markedly influenced by mood, so that our manic patients have an elevated sense of their own power and importance, while our depressed patients tend to feel inferior and insignificant. The second variable predicting the outcome of contests is Resource Value, or the value which the contestant attaches to whatever is being fought about; and we know that to manic patients things tend to seem significant and important, while to depressed patients the various prizes of life seem so much dust and ashes – to them, nothing seems worth doing, let alone fighting for. The only other relevant variable is ownership, in that, in most species, the owner of a territory tends to win a fight, while the intruder loses – and here there is a vast difference between manic patients, who tend to think they own the whole world, and depressed patients who may feel they own nothing at all, and do not deserve even the room they take up in space. The only other important difference between manic and depressed patients is energy, and we think it is reasonable to assume that when the “can” of RHP is present together with the “will” of Resource Value, nature adds the energy to carry out the task.
Thus, when a person adopts an escalating strategy, he or she undergoes an increase in RHP, Resource Value and sense of entitlement (Ownership), experienced as a surge of elevated mood, and in such a state is likely to win whatever conflict he or she is engaged on. Conversely, an individual adopting a de-escalating strategy undergoes a decrease in self-esteem, Resource Value and Ownership, and these changes are experienced as depressed mood, which makes that person more likely to lose a conflict, and to accept that he or she did not deserve to win and so accept reduced social rank and reduced access to resources.
Contributions from younger men
At this stage we were joined in our efforts by three younger men, who all made important contributions (we say men deliberately, for it is a sad fact that no member of the female sex has shown any interest in our ideas). Dan Wilson, then a psychiatrist at MacLean Hospital in Boston, took an epidemiological approach and showed that the genetic tendency to affective disorders was more prevalent in the population than would be expected by random mutation, and thus would be likely to confer some fitness advantage (Wilson, 1989, 1993, 1994 a & b). Paul Gilbert, a clinical psychologist in Derby, UK, carried out surveys on depressed patients, showing that they do indeed show characteristics associated with low hierarchical rank (Gilbert, 1992, 1993, Gilbert & Allan, 1994, Gilbert, Price & Allen, 1995); he also clarified the evolutionary switch in social competition from agonistic to prestige competition, an important theoretical development to which we will return. Peter Rohde, a psychiatrist working for the NHS in London and also in Harley Street, reported that our hypothesis was useful in his clinical work, and he added the observation that the agonistic encounter important to the patient can be with an element of his or her own mind rather than with an external person; he also pointed to the lack of attention to hierarchy in social science literature (Rohde, 2001).
Catalytic influences during these years were the Birmingham Group which revolved around the late ethopharmacologist Michael Chance, and the ASCAP Society (Across Species Comparisons and Psychopathology). In particular, the ASCAP Newsletter (followed by the ASCAP Bulletin) edited by Russell Gardner, Jr. provided an informal and hedonic forum for the exchange of ideas.
In spite of these contributions, our theory did not gain wide acceptance, and the time came for a critical examination.
Problems with the hypothesis

  1. A lot of high ranking people are depressed.
  2. A lot of low ranking people are happy.
  3. Ritual agonistic behaviour has been called “inter male aggression” because in most species it is the prerogative of the male, but depression is commoner in human females in a ratio between 2:1 and 3:2.
  4. Depression tends to follow what social scientists call “exit events” such as death and desertion by loved ones, rather than by the “entry” of a rival or competitor.
  5. Depressed patients can be quite powerful and stubborn. Aaron Beck advised, “Do not lock horns with a depressed patient or you will be pushed right out of the office (consulting room).”
  6. Depressed patients do not express submission to individual people, and their skills at appeasement displays such as flattery are impaired.
  7. In a different logical category, is it a good thing to classify depressed patients as “losers”, and so possibly stigmatise them?
    There are a number of reasons why there is an imperfect correlation between mood and social rank. One is ambition or “up-hierarchy motivation” – if one has no ambition, then low rank may not be depressing. Another is the existence of multiple hierarchies – for instance, a man who is high-ranking at work may be depressed because is dissatisfied with his low rank at home, and vice-versa.
    The human female may be not much less aggressive than the male, but the aggression is expressed in different ways – for instance by social exclusion rather than by fighting, and the weapons used are verbal rather than physical. Also, the female is exposed to situations which the male does not encounter, such as domination by a mother-in-law. Moreover, because of the lesser variation in fertility among females, a de-escalating strategy may be less of a disadvantage.
    There is no doubt that depression follows exit events, such as bereavement, separation, rejection and betrayal. This may be because loss of an ally is a good predictor of loss of rank. Even among Old World monkeys, rank is determined by patronage, kinship and alliances as much, or even more, that by individual qualities. It is not surprising therefore that human should be exquisitely sensitive to the social effects of bereavement and other exit events (Price & Gardner, 1995b).
    We dealt at some length with the problem of the power apparently wielded by depressed patients and the fact that they often appear aggressive (Price & Gardner, 1995a). The important thing is the object of the aggression. If we divide the social environment into loving supporters on the one hand and competitors on the other, then there are two categories of people who are the recipients of the aggression of depressed people. One such group is the loving supporters, and the other is lower-ranking competitors. That this simple distinction is not generally understood is revealed by the surprising fact that in all the extensive work that his been carried out on the aggressiveness of depressed patients, the relative rank of the recipient has never been recorded, nor whether that person was a supporter or competitor. For one thing, a contestant in a fight or other conflict is often acting as a representative of a group of allies, who are supporting the contestant and making, as it were, aggressive noises in the background. So that when the protagonist de-escalates he or she must force the supporters to de-escalate too, and this may require some forcefulness – the message is, “Do not force me into the arena to fight on your behalf, and stop making those aggressive noises, otherwise my submission will not be accepted, and moreover, you might like to treat me as sick and succour me.” And thus we are in the common situation of having a patient apparently suffering form a physical disease, and who forcefully insists on a physical diagnosis which will support the de-escalating strategy of being “off sick” or “out of action” (Price, Gardner & Erickson, 2004).
    Another reason for the aggressiveness of depressed patients is that the de-escalating strategy applies to those who are higher-ranking and/or have just defeated them, and not to those lower in the hierarchy. If alpha falls to beta, then gamma may hope to get in on the action and rise to beta, so the new beta may be aggressive to gamma in order to prevent a further fall. Thus lower-ranking competitors may experience an increase in aggression from depressed patients.
    Regarding the lack of focussed submissive behaviour in depressed patients, we consulted with Tyge Schelde in Denmark, who has spent many years carrying out ethological studies of depressed patients. He told us that when he divides appeasement into active and passive, he finds that the depressed patients are low on active submission but high on passive submission: they showed a lot of involuntary submissive behaviours but not a single voluntary submissive behaviour. We developed the idea that the submission of depressed patients is a passive, involuntary process, which we variously called the “yielding subroutine” or “the Involuntary Subordinate Strategy (ISS)”. Then we read about Paul MacLean’s triune brain theory. It was like shining the white light of escalation/de-escalation theory through the prism of triune brain theory, revealing a double triptych in which the escalatory and de-escalatory behaviours at the three brain levels were revealed in their primary colours (Table 1).
    A triune mind in a triune brain
    The idea that the mind consists of two or more relatively independent entities has been around at least since the time of Plato. It has been most pithily expressed by Pascal in his aphorism, “The heart has its reasons which the Reason knows nothing of.” Ancient Eastern philosophers, whose ideas were largely promulgated in the West by Gurdjieff, used the metaphor of the cart, horse and driver. The driver represented Reason, or the rational mind, but he had only limited control over the horse, who represented the emotional mind (located in the heart), who in turn had limited control of the cart, representing the instinctive mind, located by some in the gut. Plato likened the three minds to different organs of state.
    (Table 1 about here)
    The work of the evolutionary neuroanatomist Paul MacLean has given support to the idea of the triune mind by his demonstration of a triune brain (MacLean 1985, 1990). Prior to MacLean, it was thought that over the course of evolution the brain had gradually grown in size, with the later additions on the whole controlling the earlier parts, largely by inhibition. MacLean pointed out that the forebrain had grown in three distinct stages, leaving three “central processing assemblies” which relatively independently respond to changes in the environment. Firstly, the reptilian forebrain evolved from the fish and amphibian brains and concerned itself, as far as social relations went, with the courtship of the opposite sex, and competition with the same sex by means of agonistic behaviour. This brain is present in all reptiles, birds and mammals, and in humans it occupies the basal ganglia or corpus striatum. Then, instead of a homogeneous accretion of additional brain volume, there developed a “paleomammalian brain” which dealt with mammalian social life, the family, the parent/offspring bond, play, and such social matters as were not part of reptilian social life. This brain is situated in the limbic system. And not only did it deal with mammalian matters, but it also dealt with those problems which had been faced by reptiles and were also faced by mammals, such as the courtship of the opposite sex and competition with the same sex. In higher mammals there developed the neomammalian brain which subserves what we recognise as rational thought and decision-making, and it brings these capacities to bear not only on modern problems such as technology and litigation, but also on the older problems which are addressed by the reptilian and paleomammalian brains such as courtship and competition. This neomammalian brain is situated in the neocortex.
    Thus we have three brains dealing with the same problems, and to some extent they co-operate, but also to some extent they act independently. They have different sources of information, they make different executive decisions, and they have different representations in awareness. This is quite a surprising situation, one that would not have been predicted, say, by an engineer accustomed to designing robots. The most surprising thing is that the rational brain, which appears to be the most sophisticated thinking machine ever to have evolved, has so little control over the two lower brains. The driver is not in control of the horse or the cart. It would have been easy for such control to have evolved, so the fact that it has not evolved suggests that there is some advantage in having one or more relatively independent lower “central processing assemblies”. In competitive relations with conspecifics, a decision frequently has to be made between escalation (fighting harder) and de-escalation (fleeing or submitting) and this decision appears to be made, relatively independently, by each of the three brains, sometimes sequentially, sometimes simultaneously.
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part 2

Agonistic competition at three brain levels
Decisions to escalate or de-escalate take place either simultaneously or consecutively at all three levels of the triune brain (Table 1).
At the rational, or neomammalian level, the decision is made consciously and voluntarily either to fight harder or to back off; when backing off, the appeasement display may take the form of a graciously worded apology, or a flowery speech of submission. At the emotional or limbic level, escalation takes the form of anger, indignation and the exhilaration of combat, with its associated bodily changes; de-escalation at this level may recruit the dysphoric emotions of anxiety and the sense of being chastened. At the instinctive level, we hypothesize that escalation in the reptilian brain takes the form of elevated mood, giving the individual a prolonged increase in energy, optimism, self-confidence and heightened sociability all of which function to recruit allies. Conversely, de-escalation at the instinctive level takes the form of depressed mood and may include unfocused anxiety, fatigue and a sense of physical disability. The appeasement display at this level communicates this impairment and disability to any rival or to society as a whole. Parenthetically, when directed at friends and allies, the appeasement display takes the form of a distress signal, sending the message, “I am sick, care for me, and do not send me into the arena to fight on your behalf”.
Prestige competition overtakes agonistic competition
Methods of competition have become more complex over the course of evolution. Group living lengthened the duration of contests, so that even in apes a struggle for dominance may take several months to be resolved. And, instead of fleeing, as happens in territorial species, the loser could remain in the group with the winner of the contest, and this gave rise to appeasement or submissive behaviour, which reflects the capacity to live in a subordinate social role. Anxiety and fear of the dominant individual, together with relatively low self-esteem and lowered mood, enabled the social hierarchy to maintain stability, and prevent rebellion. At some stage in evolution, this stabilising anxiety gave rise to a new way of relating to a higher-ranking individual: respect. The leaders of the group made themselves attractive to the group members instead of (or in addition to) intimidating them. Social rank was then determined by the choice of the group rather than by agonistic dyadic encounters. The new self-concept of Social Attention Holding Power (suggested by Paul Gilbert) began to replace RHP, as group members evaluated themselves according to their power to attract interest and investment (such as votes or other forms of political support). Related to SAHP is the concept of prestige, which is the extent to which the group is prepared to invest in the individual. Prestige competition was added to, but did not entirely replace, agonistic competition.
The capacity for escalation and de-escalation appears to have survived the switch to prestige competition, but takes different forms, at least at the upper two forebrain levels. At the highest level, pursuit of goals replaces the decision to attack, so that escalation consists in the adoption of new goals, and de-escalation consist of giving up goals. The goals are usually ones that lead to prestige, if achieved. Also, on social occasions, escalation takes the form of self-assertion, such as standing up to speak, and promoting one’s own goals, whereas de-escalation takes the form of self-effacement, and allowing other people’s goals to take precedence in the group.
At the emotional level, escalation is less dramatic than the anger of agonistic competition; it takes the form of exhilaration, enthusiasm and self-confidence. De-escalation reflects the fact that punishment comes from the group rather than from a dominant individual, so there is social anxiety, guilt and shame. This is an appeasement display to the group, expressing contrition for breaking group rules, or for failing to come up to group standards.
At the instinctive, reptilian level of the forebrain, little seems to have changed: elevation of mood represents escalation, and depression of mood de-escalation. However, the information which leads to the activation of the strategy set is clearly different. Instead of measuring punishment received from the rival, the reptilian brain in some way monitors social standing in the group, and is sensitive to group approbation and disapprobation, to comparison of self with other group members, and with one’s own aspirations, and to the knowledge of having failed the group in some way and the likelihood of being found out. Note that depressed and elevated mood are “all or none” things; whereas at the higher levels it is possible to escalate in some areas of life and de-escalate in others, in the reptilian brain the mood change is pervasive and affects all aspects of life – it is not situation dependent. This may reflect the pervasive change in the defeated reptile, who often loses his gaudy adult colouring and reverts to the dull brown or green of the adolescent colouration.
The manifestation of escalation and de-escalation at the three brain levels are shown for agonistic competition in Table 1 and for prestige competition in Table 2.
(Table 2 about here)
Different decisions at the three levels
Normally a “resource challenge” will activate only one or two of the three levels, and then, if anger accompanies rational escalation, the individual is likely to win the conflict and the resource challenge is dealt with. Or, if chastened mood accompanies rational submission, the individual loses the conflict and becomes reconciled to the loss of whatever was at stake.
However, two very human tendencies may lead to trouble. Our often implacable ambition and stubbornness may lead to prolonged escalation at the rational level in situations in which victory is extremely unlikely, and then the anticipation of losing may activate the reptilian level strategy set and select for de-escalation at that level. The resulting incapacitating depression makes winning even less likely, and a chronic situation results in which there is continued escalation at the rational level and continued de-escalation at the instinctive level. This is a common manifestation of depressed mood as seen in the clinic.
The other human tendency is our desire to see fair play and our intolerance of injustice – this manifests at the emotional, limbic level, which seems finely tuned to evaluate the fairness of events and particularly of other people’s actions. If we feel we have been treated unfairly we feel angry, and if this anger is ineffective in righting the situation, our reptilian strategy set may be activated and we have a mood change. If elevation of mood is selected, we may then have enough energy to right the wrong, but if depression is selected, the depressive incapacity then makes effective action even more impossible. Then, again, we get chronic reptilian de-escalation which presents in the clinic as depressive illness.
Case examples
In order to give an idea of how these things appear in the clinic, we will give some case examples, disguised to maintain confidentiality.
Case A : Rational escalation and instinctive de-escalation. A highly moral lady had a job taking classified ads for a magazine. The magazine started losing money and the manager was desperate to increase the advertisement revenue. He required the patient to take ads from call girls. She refused to do this and received a lot of negative signals from the manager and other staff and eventually went off sick with depression. In the treatment of such a patient, we do not direct attention to the depressive symptoms, but rather ask the patient why they have not dealt with the problem at a higher brain level. We say to them, “If you came to us complaining of shivering, we would not rub ointment on your shivering muscles, but rather we would ask why you had not turned on the central heating.”
This patient had refused to submit to her manager because of moral scruples. We have seen a case of depression in a health service clerk who refused to “fiddle” waiting list data in order to make the waiting time appear to be getting shorter. Sometimes the required submission is not to management but to colleagues. We have seen depression in workers who refused to take part in systematic theft. Stevens & Price (2000) described a hospital porter who became depressed because he refused to take sick leave when he was not sick (and so allow his colleagues to claim overtime pay). These are all cases in which adherence to moral values prevents voluntary submission. Other blocks to voluntary submission are pride, stubbornness, sense of fair play, and in general the sort of mental state which made Job say, “I will never submit” and Garibaldi to say, “Rome or death!”. In these cases, while voluntary submission is blocked, the involuntary submission of depression may take over the direction of behaviour.
(Table 3 about here)
Case B: Emotional escalation and instinctive de-escalation. A couple had a daughter who at the age of nine developed a pituitary tumour. This led to premature puberty and menstrual bleeding. The parents took the girl to hospital, where the father was detained by social services, aided by the police, for twenty four hours, accused of sexually assaulting his daughter, and he was not allowed to return to the family home until the diagnosis of pituitary tumour was made some weeks later. The parents were overwhelmingly angry at the treatment of the father – it was the mother who came to the clinic some years later suffering from depressive illness.
Common causes of emotional escalation (anger) are mistreatment by management or official bodies, or by fate, as when parents suffer the death of a child at the hands of a drunken driver. In Palestine depression may be due to punishment by the occupying power (Costin, 2005). In these cases it is helpful for the patient to join a group of those who are similarly afflicted. Then the anger can become group anger, and the depression may remit as depression is part of ritual agonistic behaviour and not of inter-group conflict. Also the anger may be directed into useful channels, such as lobbying for stricter drink/driving laws. In Palestine, the groups organised by the health service also reduce the chances that victims will join groups for training suicide bombers.
(Table 4 about here)
Case C: Triune de-escalation but submission not accepted. A morbidly jealous man had financial difficulties and so his wife went out to work, and the husband fantasised that she was meeting other men. He would not admit to these feelings, but criticised his wife for being stupid, ugly and slovenly. Although the wife submitted totally, she did not know that the submission he required was for her to give up work. Eventually he got his way when she went off sick with depression.
Submission may also be blocked by a third party. We remember a patient who tried to submit to her husband but was prevented by an even more dominant mother, who insisted that her daughter visit her (and do housework) at a time when the husband wanted her at home (in this case the patient recovered when her mother died).
(Table 5 about here)
Case D: Inappropriate triune de-escalation. A female patient had been told by her mother that she was unwanted, and would have been aborted if the mother could have afforded it. Then she was badly bullied by her peers in adolescence. As a result she had very low self-esteem, and became depressed as a result of trivial setbacks. She required individual psychotherapy to give her a successful “rerun” of the parent child relationship, and group therapy to give her a successful rerun of the peer-group relationship.
In these cases the agonistic strategy set (of escalation and de-escalation) is too easily accessed or triggered, a problem well described by Sloman (1998).
Research
A lot of animal research is going on concerning the biology of social rank change (Wilson & Price, in press), using particularly the resident/intruder paradigm in rats. It would be nice to think that our evolutionary speculations had given impetus to this research, but this is not the case. In the 1970’s research was funded by departments of internal medicine, because it was noted that defeated and low-ranking animals developed stomach ulcers and hypertension. More recently the researchers realised that the defeated animals were also depressed, and so the models have been specifically used to study depression in animals. However, no research has been carried out on one of the most promising animal models of depression since the Medical Research Council terminated the funding of one of us (JSP) who was working on the vervet monkey, Cercopithecus aethiops pygerythrus. This animal was reported to become depressed when it fell in rank, while at the same time its bright blue scrotal skin turned white. Blue skin is common in cercopithcine monkeys, a Tyndall blue caused by the selective scattering of short wavelengths by bundles of collagen of a diameter near to the wavelength of blue light, lying over a deep dermal deposit of melanin which absorbs the transmitted red light. This vervet monkey is unique in having a blue scrotal skin, which is a dominance signal, and is attended to closely by other males. When the monkey loses rank, the scrotal skin turns white over the course of about two weeks due to hydration of the dermis with resulting change in the diameter of the collagen bundles. Genital skin hydration is common in cercopithecine females at oestrus, but the vervet females do not show this. It seems possible that a mechanism of genital skin swelling which subserves the sexual changes in many monkey females has been taken over in this particular vervet to subserve dominance changes in the males (Price, 1989). Much excellent work has been done on the vervet monkey by McGuire and his colleagues (McGuire, 1988), but unfortunately the strain they used does not have the blue scrotal skin (which can be readily seen in wild vervets in East Africa).
Our theorising also indicates the possible use of reptiles as animal models of depression. Reptiles get “depressed” (McLean, 1990) and when they are depressed they may change colour; for instance, the lizard Anolis carolinensis loses its bright adult colouring when it loses rank, and reverts to the dull brown adolescent colouring (Greenberg & Crews, 1983). Reptiles are simpler to study than mammals, having mainly visual rather than olfactory signalling, and lacking the two higher levels of the triune brain. And yet PubMed cites no papers reporting research on depression in reptiles, while there are over 600 reports of depression research in rodents.
Conclusion
Perhaps the musings of Demaret, Gardner, Price and Sloman, supplemented by those of Wilson, Gilbert and Rohde, have borne fruit. We should like to think that they have solved two puzzles, which in their solution have cancelled each other out. One puzzle was the biological function of those mood changes from elation to depression and back again which we see exaggerated and out of context in our bipolar patients. The other puzzle was the behaviours which have evolved to deal with the much contrasted roles of high ranking territory-owners and low-ranking non-territory owners, and the transitions between these roles. Proof that the mood changes evolved to subserve the rank changes is difficult to come by, but we hope that our discussion has illuminated the relation between mood and social competition to such an extent that it will help in the treatment of afflicted individuals.
Summary
Fighting to the death, winning at all costs, taking one’s case to the highest court in the land, these are escalating strategies which serve to obtain resources for the individual. But everyone cannot win. Giving in, submission and appeasement are vital social behaviours, without which social life would be impossible. They are manifestations of a de-escalating strategy, in which the chances of winning are much reduced but the costs of losing are also reduced. In the course of its evolution from reptiles through mammals to primates, the human forebrain has developed three “central processing assemblies”, each of which makes a decision between escalation and de-escalation when confronted by challenge or stress. If the higher assembly does not deal with the problem, the lower assemblies are activated and select more primitive forms of escalation and de-escalation, which in the case of the lowest of the three assemblies may well take the form of elevation and depression of mood, respectively. The appeasement display of the reptilian de-escalating strategy is a sophisticated message which says, in effect, “I am too incapacitated to put on an appeasement display.” This is the ultimate in reassurance.
For the therapist, the aim is to guide the patient away from conflict towards negotiation, compromise, reframing and if necessary leaving the arena. The therapist can reinforce the tendency of depression and anxiety to promote reconciliation, depression by reducing expectation, and anxiety by helping the patient to seek comfort and safety from a former antagonist (Price, 2003; Sloman et al., 2006)).
Perhaps it might not be hubristic to envisage a wider application of our ideas. The notion that depression may be a magnifier of small genetic differences is surely relevant to the speed of evolutionary change. And, in addition, the mechanisms here described may have facilitated group living, by promoting the smooth functioning of hierarchies within groups, with depressed mood in particular helping to reconcile low-ranking individuals to forego the rewards of high rank such as food, shelter and mates; and without group living, if our ancestors had been as solitary as the orang-utan, we might not be in possession of the faculties required to write this article.

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PheroQuirk are you specialized in the area of Psychology? I ask because that will be my major, and I’d like to maybe get a few pointers from you if you are involved in this field of work, or are going to school.

Btw, I’ll read the article when I get some time. Thanks.

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Hi Mindset.

I’m just an amateur in that field. My area of specialisation is finance but I’m interested in psychology and read a lot.

Pheroquirk

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Ah, ok. I was going to major in accounting/finance, but I have always been fascinated with Psychology, and it has always been my desire to become at the very least, a councelor to help others who have been through as much inner-turmoil as I have. I wasn’t sure, but I thought I had heard Jasmin mention your name in regards to Psychology, but I suppose that was just as a general knowledge assessment as you do seem to be well read on the subject. Anyways, I guess I’ll try some Psyche forums. Thanks, Pheroquirk.

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One piece of advice I can give you is to get as much work experience as you can in fields close to those that you are considering. If you can’t get actual experience then try to arrrange some kind of work shadowing.

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Yes, talking to others has definately rubbed that impression onto me. Do you have any recommendations on some good psychology books that you have read?

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For preparing for a degree course or to counsel people - that I can’t help you with. For understanding other people better… start with a textbook on social psychology. ‘Influence’ by Cialdini. A bunch of books on evolutionary psychology (maybe start with the Red Queen although there are lots of others. I just can’t remember everything I have read). Ekman on facial expressions.

Ask tacitus - he has an immense amount of expertise here. Also read some of MHarris old posts (a good idea to do that anyway!)

Look at community forums on nlpweekly.com - some interesting stuff there.

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