PHYSIOLOGY & ELSEVIER

Physiology & Behavior 73 (2001) 435-442

BEHAVIOR

Social

defeat
Kaj

as a stressor
Bjorkqvist* PO. Box 311, FIN-65101

in humans

Abo Akademi Received

University,

Vaasa. Finland 2001

2 November

2000; accepted 22 February

Abstract

Studies on social defeat in humans, and their similarities with studies on social defeat in animals are reviewed. Studies on social defeat in humans typically are conducted as a branch of social psychology, most often focusing on bullying in schools and in workplaces. Victims of bullying are known to suffer from depression, anxiety, sociophobia, loss of self-esteem, psychosomatic diseases, and other behavioral symptoms. On the other hand, animal studies on social defeat, usually based on the rodent resident-intruder paradigm, present fmdings related to physiological rather than to behavioral consequencesof defeat. The two branches use different terminology, e.g., "dominant" and "subordinate" (animal studies) and "bully" and "victim" (human studies). It is suggested that the two fields could benefit from a mutual exchange in theory and methodology. ~ 2001 Elsevier Science Inc. All rights reserved.
Keywords: Social stress; Social defeat; Aggression; Bullying; Victimization; Dominant; Subordinate

I. Introduction:

lessons

from

animal

models

The aim of the present article is to review studies on the effect of social defeat in humans and to compare results from these studies to findings obtained within research on social stress in subhuman species. Research on social stress in animals, particularly in rodents, has developed a tradition with specific models, the most popular being the resident-intruder paradigm (see, e.g., Refs. [46,48]). In this model, a male rodent (the intruder) is put into the home cage of another male (the resident). Alternatively, a cage with two compartments with a removable wall between them is used. As the wall is removed, the animals fight for a predetermined period of time, almost inevitably leading to one of them taking a, usually severe, beating. The winner is often referred to as the dominant male and the defeated animal as the subordinate male. The defeated animal is considered to experience social stress. If they are allowed to fight on a single occasion only, it is regarded as a model of acute stress; if they are allowed to fight at several consecutive occasions stretching over days and perhaps weeks, it is regarded as a model of chronic

stress. The subordinate may be exposed to chronic stress also in the form of threat by, e.g., having to stay in the compartment beside the dominant, exposed to its visual and odor cues, after having been defeated or between fights. The model works well as far as male rodents are concerned, but a problem with the model is that only male rodents may be used, since female rats and mice do not fight each other in a resident- intruder confrontation. Animal studies suggest that social defeat may result in a large variety of severe symptoms. They also suggest that these symptoms differ from those caused by other stressors. In fact, already a single experience of defeat induces remarkable signs of stress in male rats: findings by Koolhaas et al. [45] indicate long-term (in this case, extending over several weeks) effects, reminding of those of depression in humans. Likewise, Blanchard et al. [20] found that subordinate male rats living in colonies together with aggressive males showed a number of symptoms typical of depression. Indications of depression would hardly be seen in connection with plainly physiological stress, neither in animals nor in humans. . Corticosterone is known to playa role in mitigating the adverse effects of stress. Social defeat in male rodents by Albeck et al. [3] was shown to cause an impaired corticosterone response, and they also found that the subordinates expressed a significantly lower number of corticotrophinreleasing factor mRNA grains per cell in the paraventricular

* Tel.: +358-6-3247-469; +358-6-3274-491. fax: E-mail address:kaj.bjorkqvist@abo.fi Bjorkqvist). (K.

0031-9384/01/$ -see front matter (\:)2001 E1sevier Science Inc. All rights reserved. PII: SO03 1-9384(01 )00490-5

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hypothalamic nucleus as compared to dominants and cagehoused controls. Sgoifo et al. [83] found that social stress, in contrast to other stress contexts, produced a shift of autonomic balance toward sympathetic dominance, associated with the occurrence of cardiac tachyarrythmias, in male rats, especially in those of a wild strain. Social defeat results in clear changes in hormonal and neurotransmitter responses in rats: for instance, Stefanski [85] investigated the specific hormonal response patterns of winners and losers after a 7-day period of chronic confrontation. At Day 7, winner males showed stable concentrations of corticosteroid-binding globulin (CBG) and reduced titers of total corticosterone, while losers had a marked decrease in CBG and unaffected total corticosterone. Increased norepinephrine and epinephrine titers were also evident in losers. Furthermore, reduced testosterone levels were observed in the defeated males. Similar effects have been found in pigs. Otten et al. [69] found socially defeated pigs (in contrast to the rodent studies, the sample consisted in this case of females and castrated males) to show an increase in plasma catecholamines, ACTH, and heart rate. Immediately after the introduction of an intruder, loser pigs responded with a significantly higher increase in plasma epinephrine and norepinephrine concentrations than did winner pigs. Losers also showed reduced locomotor activity and exploring behavior, i.e., they showed not only physiological but also behavioral symptoms, suggesting more emotional distress and fear as compared to winning pigs. Levels of nerve growth factor (NGF), a polypeptide growth factor exerting trophic and differentiative effects on both peripheral and CNS neurons, have been shown to increase subsequentto aggressive encounters in mice [4-6]. They suggested that when released into the bloodstream after psychosocial stress, NGF affects peripheral nervous structures (chromaffm cells and ganglia) and peritoneal mast cells [4]. Others [1,87] have made similar fmdings. There are also clear indications of impaired immunological function as a result of social defeat. For instance, Stefanski and Engler [86] studied the effects on winning and losing among male Long- Evans rats on Days 2 and 7 in a chronic social stress design. On both Days 2 and 7, losers showed reductions in the number of blood CD4 and CD8T cells, and they also showed a suppression of in vitro lymphocyte proliferation. Winners, on the other hand, showed similar immunological changes only on Day 2 of confrontation. Cohen et al. [25] found impaired immunological function and increased susceptibility to respiratory infection in socially defeated male macaques. Weiss and Sundar [91] and Biondi et al. [II], in their reviews of the effects of stress on cellular immune response in animals, report similar findings in other species. A single experience of social defeat has been shown to increase slow-wave sleep in rats as measured by EEG [59]. Since slow-wave activity is thought to be a sign of sleep

intensity, the authors suggestedthat the acute stress of defeat results in a greater need of rest for the animal. Kaplan and Manuck [43] studied the effect of social defeat in male macaques and in socially stressed, socially housed female ovariectomized macaques [41]. In both cases, they found an enhanced disposition for coronary atherosclerosis in subordinated animals, males and females alike -although fertile nonovariectomized female macaques, like women, are resistant to atherosclerosis. They concluded that circulating estrogen is likely to prevent atherosclerosis in nonovariectomized females. Stresscausedby social defeat may also be related to drug abuse. Tidey and Mizcek [89] found that when given the opportunity to administer cocaine to themselves, defeated male rats acquired self-administration habits in half the time ofnondefeated males. They also examined, by use ofin vivo microdialysis, dopamine concentrations in nucleus accumbens. Extracellular dopamine levels in nucleus accumbens increased to 135% of baseline in defeated rats as compared to 125% of baseline in nondefeated rats. Furthermore, when exposed to social threat by the resident, the levels changed to 145% in defeated rats vs. 120% in nondefeated rats. The authors suggested that exposure to stress may induce changes in central dopaminergic activity, rendering the individual more vulnerable to self-stimulation due to a cross-sensitization to the rewarding effects of cocaine. In their review of studies based on mice, Laviola et al. [51] suggest that adolescence, in both animals and humans, may be associatedwith an increased risk of drug abuse. This may be due to the fact that novelty-seeking behavior is typical of this period, but there may also be an underlying, strictly biological, explanation: in animal models of preadolescence, the search for novel stimuli and sensations shares a common neurological substrate with psychostimulants, the reward-related mesolimbic pathways. As previously pointed out, animal models of social defeat, and in particular, those based on rodents, almost exclusively use males as subjects. This weakness has troubled researchers, and Railer et al. [37] underlined that reliable social stress models for females are virtually lacking. They suggestedthat social instability might be a useful alternative as a social stressindicator in female rodents: in order to create a socially unstable environment for female rats, they alternated phases of isolation with mixed-sex crowding, including rotating membership during the crowding phases.The effect of social stress, induced in this manner in females, was compared to the effect of the social defeat paradigm in males. Defeat stress reduced weight gain and increased both adrenals and plasma corticosterone in males, while social stress had an effect only on adrenal weight in females. The question remains, however, to what extent the social stressorsin these two cases really are comparable. Conclusively, the animal social defeat model has revealed a number of quite severe consequences of social defeat in animals of various species. The examples provided above are by no means intended as a comprehensive over-

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view, but they show evidence of at least the following effects of social defeat in animals: impaired corticosterone response in male rats [3,85], decreasein testosterone levels in male rats (e.g., Ref. [85]), increase in plasma epinephrine and norepinephrine in male rats [85] and in female and sterilized male pigs [69], increase in plasma catecholamines, ACTH, and heart rate in female and castrated male pigs [69], a general shift of autonomic balance toward sympathetic dominance, associated with the occurrence of cardiac tachyarrythmias in male rats [83], enhanced disposition for atherosclerosis in both male and ovariectomized female macaques [43,44], increased NGF expression in male mice [4-6], impaired immunological function in male rats [85,86], male macaques [25], and in a number of other species as well [8,91], increase in slow-wave sleep in male rats [59], increased self-administration of cocaine and changes in dopaminergic activity, which may cross-sensitize to cocaine self-administration, in male rats [89], reduced weight gain in both male and female rats [37], and decreased locomotion and exploring behavior in female and castrated male pigs [69]. Now turning our focus toward research on social defeat in humans, the first observation made is the relative absence of laboratory experiments. Experiments in which humans experience severe social defeat will hardly be approved by the ethics board of any present-day university. Accordingly, the main part of these studies is correlational, and accordingly, they usually lack the baseline measurements possible in laboratory designs. However, a few experiments on social defeat in humans have been conducted.

Jeffcoat et al. [42] investigated whether endocrine changes could be induced in five men confined on a boat for 14 days. The participants were secretly ranked for dominant/aggressive behavior toward the other men. A significant correlation was found between day-to-day changes in self-assessedanxiety and stress hormones, cortisol, and prolactin. A significant correlation was found also between plasma prolactin, testosterone, and rank position for dominance/aggression. The authors concluded that under at least some circumstances, social interaction might modify endocrine status in human males. One of the earliest studies on the relationship between testosterone, aggression, and dominance among human males was conducted by Ehrenkranz et al. [31 ], with male prisoners as participants. They found a significant relationship between testosterone and both aggression and dominance. Dabbs et al. [29] found higher levels of testosterone among prisoners convicted for violent crimes than among prisoners convicted for nonviolent crimes. Kreuz and Rose [47] made a similar study but failed to find any such relationship. However, they found higher levels of testosterone than among others among a small subgroup of 10 inmates who had committed violent crime during their adolescence. A link between high testosterone and violence was found by Rada et al. [74] among a small group (n = 5) of sexual offenders (rapists and molesters) judged by police records to be more violent than other offenders. Another study of offenders by Olweus et al. [67] provided mixed results. Bain et al. [9] did not fmd any difference in testosterone level between three groups of offenders convicted either for property offence, assault, or repeated assault. Accordingly, there are studies suggesting a link between testosterone and aggression in human males convicted for sexual offense or violent crime, but the findings are equivocal. However, since these studies concern samples of prison inmates, generalizations to the population-at-large must be made with caution. Dabbs [26] conducted a similar study with female prison inmates as participants. Testosterone was measured from radioimmunoassay of saliva samples. He concluded that, like in male prisoners, testosterone indeed was also linked to aggressive dominance in female inmates. Cashdan [24] investigated hormonal correlates of dominance and status in coresidential college women. She found that estradiol, total testosterone, and free testosterone all were related to high self-regard, number of sexual partners, and infrequent smiling, a behavior usually associated with dominance. Dabbs [28], for instance, found that low-t~stosterone men were smiling more frequently and with broader smiles than did high-testosterone men. Although there may be a link between social dominance and testosterone in small-group interaction among both males and females, there is no evidence of any correlation between testosterone and social/occupational status. The opposite may in fact be the case: Dabbs [27], who inves-

2. Dominance, defeat, and testosterone A popular research topic in the area has been the investigation of whether defeat is related to reduced testosterone levels based on the rationale that testosterone, dominance, and aggression ~ay be related not only in subhuman species but also in humans. Rejeski et al. [77] examined cardiovascular and testosterone responses in dominant and submissive male undergraduates, who engaged in a debate with a trained female. While dominant males reported a subjective greater loss of dominant status as a result of the debate, the submissiye males manifested greater decrease in testosterone levels. Heart rate data revealed that, as the debate continued, the dominant males habituated to the situation, while the submissives experienced increased threat. In another study, Rejeski et al. [78] studied cardiovascular and testosterone responses in dominant and submissive female undergraduates in a similar design. Dominant females reacted more to the social stress with cardiovascular parameters than submissive ones. However, serum testosterone did not vary as a function of the experimental manipulation.

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tigated 4462 foffiler US military servicemen, found that participants with higher levels of serum testosterone had lower-status occupations. As an explanation, Dabbs suggested that since serum testosterone is thought to be heritable and available at an early developmental stage, high levels presumably evolved in association with dominance in individual and small-group settings. Thus, testosterone, which perhaps evolved in order to support a primitive kind of status, in fact, in today's society, is associated with low socioeconomic status. Schaal et al. [82] and Tremblay et al. [90] conducted a longitudinal study of boys from 6 to 13 years of age, following their development with regards to testosterone, dominance, and aggression. At age 13, social dominance and testosterone levels were assessedduring a I-day visit to the laboratory, together with four unfamiliar peers. Boys perceived by the peers as dominant had higher levels of testosterone than others. However, a positive correlation between testosterone and aggressivenesswas not found: in fact, boys who had a history of high physical aggression had lower levels of testosterone than boys with no such history. Bjorkqvist et al. [16] administered testosterone, placebo, or no drug to adult human males. They did not observe any increasein aggression in the testosteronegroup; however, the placebo group (!) showed and increase in aggression, both in comparison with the controls and the testosteronegroup. Mazur and Lamb [58] found increased levels oftestosterone among winners in a tennis competition but not among winners in a lottery. Also, new recipients of an MD degreehad increased testosterone. They concluded that when a rise in status is achieved, testosterone levels increase. Conclusively, there appears to be support for the notion that testosterone is linked to dominance in humans, although mainly in small-group settings; the findings are so far inconclusive with regards to social/occupational status. The link to aggressiveness is less clear, demonstrated in some studies (e.g., Refs. [31,67)), but not in others (e.g., Refs. [16,82,90)). In his review, Archer [8] suggests that there is reasonable evidel;lce for a link between aggressiveness and testosterone among adult human males but not among adolescents.As Benton [10] points out, the claim of a relationship between testosterone and aggression is unambiguously demonstrated only in animal data. Human aggression is often instrumental and calculated rather than emotogenic, i.e., related to anger. As far as human aggression is concerned, social and cognitive factors playa much greater role than physiological factors. The higher up on the phylogenetic scale, the more uncertain the relationship between aggression and testosterone appears to be. Testosterone-relateddominance in small groups may be observed in nonverbal and verbal communication, such as less smiling and more frequent interruption of others [24,26,28]. With regards to small-group interaction, there appears to exist a testosterone-dominance link similar to that found in subhuman species. However, with respect to social/occupational status, there is no evidence at all of

testosterone being linked to high status. Instead, the opposite appears to be the case [27]. Summing up, when groups of males (and surprisingly, also females) spend much time together and interact intensively with each other, like in prison, on a boat, or in a coresidential college -perhaps also in the army, in school, or at the workplace -social defeat appears to reduce testosterone from baseline levels. On the other hand, individuals low in testosterone may also appear less threatening to their peers, and, accordingly, they may become socially defeated more easily. Thus, the testosterone-dominance relation brings to mind the old question of which is hen and which is egg: clearly, testosteroneand dominance seemto interact with each other, and a circular effect may also occur.

3. Bullying in schools There exists a large body of research on social defeat in humans, which has been totally neglected by social stress researchers. This is the research on bullying in schools and work places. Bullying was defined by Sullivan [88] as a conscious and willful act of aggression and/or manipulation by one or more people against another person or people. Bullying can last for a short period or go on for years and is an abuse of power by those who carry it out. Others (e.g., Refs. [13,15,63-65]) explicitly stress that it is an unequal-power relationship between aggressor (the bully) and victim, and the victim is, for some reasonor another, not able to defend him/herself. It is usually an ongoing process, with the bullying increasing in severity. However, also, single acts may be regarded as bullying when characterized by abuse of power. Often, bullying does not appear to have any other meaningful purpose than to humiliate the victim, thereby increasing the bully's self-esteem and scaring others ("look what may happen to you if you oppose me"). In a school class with bullying problems, there typically is one main bully with a few armor-bearers or supporters and one or two victims, and roughly 60- 70% of the pupils are silent onlookers or bystanders [14,63,65,81]. However, as the bullying increases in severity, more and more pupils typically take part, turning it into a process of group aggression [49]. Typically, besides being beaten up, the victim often is pressured to bring money to the bully or to do other services. He is also threatened with revenge in case he attempts to get help from teachers or other adults. Suicides caused by school bullying have been reported in several countries [60,66]. At fIrst, the research focus was on physical bullying [63,65]; however, soon it was discovered that much of the bullying occurring in schools was in fact of psychological nature, conducted as indirect aggression, manipulating the social fabric of the class in order to exclude the victim from the social groups of the class, and having him/her rejected

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by peers. This type of bullying was found to be particularly prominent among girls [10,19,50,68,92]. Research on bullying in schools commenced in the 1970s [56,63,65]. Most of the early studies were conducted in Scandinavia [12,17,49,63-66,80,81]; later on, research was taken up in the UK, Australia, and New Zealand [22,70,79,84,88,92]. The phenomenon is well known also in Japaneseschools, where it is called ijime [60]. Research on bullying in the USA has been scarce,but some studies do exist; (for a review, see Harachi et al. [39]). Slightly more research in the area has been conducted in Canada [38,73]. However, there is a related body of research in US focusing on peer rejection [21,71,72]. Partly, it appears to be a question of terminology. At present date, research on school bullying is going on in a large variety of countries on all continents, indicating that the phenomenon is not limited to any specific culture, but it appears to be universally human [62,84]. Bullying has interesting parallels with animal models of social defeat, the bully being equivalent to the dominant animal and the victim to the subordinate one. A single incident of bullying may be compared to the acute stress model and ongoing bullying to the chronic stress model. There can be no doubt about the fact that victims of school bullying experience severe stressdue to social defeat. What is known about the consequences of bullying? Bjorkqvist et al. [14] found victims to be depressed, have poor self-esteem, feel themselves as failures, and feel themselves (wrongly) to be unintelligent, poor academic achievers, and ugly. Lagerspetz et al. [49] found victims likewise to have low self-esteem, having a subjective feeling of maladjustment, and showing grave dissatisfaction with school and peer relations. Olweus [63,65] found high levels of anxiety in victims. Hawker and Boulton [40], in their review of others' studies, conclude that victims typically show the following characteristics: loneliness, depression, anxiety, low self-esteem, submissiveness,social withdrawal, and unpopularity among peers. These symptoms occur among male and femal~ victims alike. Accordingly, chronic stress in the form of social defeat -victimization to bullying -has severe behavioral consequences.A problem, of course, is that since the research usually (ifnot always) is post hoc, no baseline (prebullying) measures of symptoms exist. Furthermore, so far, there are few, if any, studies using physiological measures, like in research based on animal models.

Gustavsson [55] as the pioneers. Again, research in the area began in Scandinavia [7,13,18,19,32,33,52-55], from where it spread to the Gennan-speaking areas of Europe [34,61] and to the UK [2,76]. Research on workplace bullying took school bullying as its starting point and source of inspiration, defining bullying in the same way. Bjorkqvist [13] described three distinct stagesof workplace bullying: (1) Stage 1 is characterized by indirect aggression, such as gossiping and backbiting; (2) Stage 2 is characterized by open verbal confrontations, followed by social rejection of the victim, who typically is not spoken to or looked at; (3) during Stage 3, the bully tries, and often succeeds, in forcing the victim out of the workplace, losing his/her employment. Although workplace bullying reminds of school bullying, there are also important differences. At workplaces, the competition about jobs is an important intervening factor, and bullying often occurs between employees of similar or close occupational status, competing for the same position. Both women and men are victimized by workplace bullying, and unlike school bullying, which mostly takes place between members of the same sex, workplace bullying occurs almost as frequently between members of different sexesas between employees of the same sex. The competition about position is a factor more important than gender [13,18]. The social group formed by colleagues at work is, besides the family, probably the most important reference group for the adult individual. Each person's self-image is dependent on how (s)he is treated by colleagues. Furthermore, losing one's job may imply losing much more economy, career, and future. No wonder that exposure to workplace bullying has been shown to have quite severe consequences, and suicides are not uncommon [52,53]. Leymann [52] provides evidence that victims of workplace bullying frequently show symptoms typical of posttraumatic stress disorder. If an employee is exposed to bullying by colleagues, especially by the boss, not being able to defend him/herself, the mean length of time for serious symptoms to appear is as short as 15 months [52]. Other typical consequences reported are depression, sociophobia, anxiety, loss of self-esteem, psychosomatic disorders, and sleep disorders. All these symptoms have been demonstrated among males as well as females [7,13,18,32,33,52,53].

5. Social defeat in females 4. Bullying in work places The f1fStstudy of workplace bullying -sometimes also called work harassment (not to be confused with sexual harassmentat workplaces, a sexist variety) -was conducted by Brodsky [23] in 1976. His study was ahead of time, and researchon workplace bullying did not gain momentum until the late 1980s, with Leymann [52-54] and Leymann and As indicated above, human females are not unaggr~ssive. During the last decades, female aggression has been studied extensively, especially the female propensity for indirect aggression [12,15,19,50,68]. In her review, Hyde [41] concluded that when aggressive, females try to cause psychological rather than physical harm to their enemies, and Eagly and Steffen [30] and Frodi et al. [35], in their reviews, came to similar conclusions. Although less physically aggressive

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than males, the human female has found other ways to aggressagainst her enemies. In fact, laboratory studies show that when not in danger of being recognized, hence of being retaliated against, females behave equally aggressively as males [36,57,75]. Human females, accordingly, are victimized to aggression by other females. They are also victimized to aggression by males. As research on bullying in schools and workplaces has shown, human females frequently experience severe stress in the form of social defeat, with similar symptoms as seen in males. However, we also do know that females in some respects react differently to social stress than males: for instance, human males may develop coronary atherosclerosis, a disease that nonovariectomized females before the menopause are protected against [43,44]. Hence, there is a need for female social stress models in animals. As mentioned in the introduction, the rodent residentintruder paradigm is not possible to use with female rats and mice as subjects, since they do not aggressagainst each other in that particular situation. Accordingly, alternative animal models of female social stressneed to be discovered. Hailer et al. [37] suggested that an unstable social situation might be one possibility. However, this source of stress appears less dramatic than for instance bullying among humans, and certainly, it does not represent social defeat. Perhaps mice and rats are not very well suited for the purpose, and other animals, in which females are known to show visible aggression- such as the golden hamster, the pig, and the hyena would be better suited for the purpose.

describe "bullies" and "victims." With the wrong key words, a search in psychological abstracts may be futile. Animal and human studies on social defeat should be able to benefit from each other by a mutual understanding of and communication about both theory and methodology. As an example, a worthwhile pursuit would be to investigate whether findings related to social defeat in animals may be replicated with humans, in research on defeat in sports, and in studies on bullying and victimization in schools and workplaces.

References
[1] Abramchik GV, Yennakova SS, Kaliunov VN, Tanina RM, Tumilovich MK. The immunomodulatory effect of nerve growth factor. J Neurosci Res 1988;19:349-56. [2] Adams A. Bullying at work: how to confront and overcome it. London: Virago, 1992. [3] Albeck DS, McKittric CR, Blanchard CD, Blanchard RJ, Nikulina J, McEwen BS, Sakai RR. Chronic social stress alters levels of corticotrophin-releasing factor and arginine vasopressin rnRNA in rat brain. J Neurosci 1997;17:4895-903. [4] Alleva E, Aloe L. Physiological roles of nerve growth factor in adult rodents: a biobehavioral perspective. Int J Comp Psychol 1989;2:213-30. [5] Alleva E, Petruzzi S, Cirulli F, Aloe E. NGF regulatory role in stress and coping of rodents and humans. Phannacol, Biochem Behav 1996;54:65-72. [6] Aloe E, Levi-Montalcini R. Mast cell increase in tissues of neonatal rats injected with the nerve growth factor. Brain Res 1977;133:358-66. [7] Appelberg K, Romanov K, Honkasalo M, Koskenvuo M. Interpersonal conflicts at work and psychosocial characteristics of employees. Soc Sci Med 1991;32:1051-6. [8] Archer J. The influence of testosterone on human aggression. Br J PsychoI1991;82:1-28. [9] Bain J, Langevin R, Dickey R, Ben-Aron M. Sex honnones in murderers and assaulters. Behav Sci Law 1987;5:95-101. [10] Benton D. .Honnones and human aggression. In: Bjorkqvist K, Niemelii P, editors. Of mice and women: aspects of female aggression. San Diego, CA: Academic Press, 1992. pp. 37-48. [II] Biondi M, Massimo Z, Luca G. Psychological stress, neuroimmunomodulation, and susceptibility to infectious diseases in animals and man: a review. Psychother Psycllosom 1997;66:3-26. [12] Bjorkqvist K. Sex differences in physical, verbal, and indirect aggression: a review of recent research. Sex Roles 1994;30:177-88. [13] Bjorkqvist K. Trakassering forekommer bland anstiillda vid AA. Med AA 1992;9:14-7. [14] Bjorkqvist K, Ekrnan K, Lagerspetz KMJ. Bullies and victims: their ego picture, ideal ego picture and nonnative ego picture. Scand J PsychoI1982;23:307-13. [15] Bjorkqvist K, Lagerspetz KMJ, Kaukiainen A. Do girls manipulate and boys fight? Developmental trends in regard to direct and indirect aggression. Aggressive Behav 1992;18:117-27. [16] Bjorkqvist K, Nygren T, Bjorklund A-C, Bjorkqvist S-E. Testosterone intake and aggression: real affect or anticipation? Aggressive Behav 1994;20:17-26. [17] Bjorkqvist K, Ostennan K. Finland. In: Smith PK, Morita Y, Junger-Tas D, Olweus D, Catalano R, Slee P, editors. The nature of school bullying: a crQss-national perspective. London: Routledge, 1999. pp. 56-67. [18] Bjorkqvist K, Ostennan K, Hjelt-Biick M. Aggression among university employees. Aggressive Behav 1994;20:173-84. [19] Bjorkqvist K, Ostennan K, Lagerspetz KMJ. Sex differences in covert aggression among adults. Aggressive Behav 1994;20:27-33.

6. Final

comments

The present article was written with the specific purpose of attempting to find common ground between studies on social defeat in humans and in animals. Since I am conducting research with human subjects, my comments and reflections about subhuman species run the obvious risk of appearing naive and simplistic. However, this is a risk that has to be taken in order to enhance communication between the two fields. Researchon social defeat in animals is usually conducted by biologists or physiological psychologists, within the broader framework of social stress studies, often with the purpose of using the model for the testing ofpsychopharmacological agents against anxiety and depression. Researchon social defeat in humans, on the other hand, has developed as a branch of social psychology, with little awareness,if any at all, of the studies made with subhuman species as subjects, despite the fact that these should be of great relevance for the understanding of the consequencesof victimization. Accordingly, one obvious problem in the communication between human and animal studies on social defeat is that of terminology. For example, the animal literature uses terms like "dominant" and "subordinate," while human studies

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