Hormonal Aspects of Aggression and Violence

Paul Fredric Brain

Relationships between aggressive behavior and the endocrine system have been studied intensively in recent years. This interest has occurred presumably because hormones are naturally occurring secretions of the bodies' endocrine or ductless glands, and are perceived as providing possibly reversible (certainly when compared to psychosurgery) therapies for some clinical conditions that include hyperaggressiveness as a symptom. Hormones are transported throughout the body by the blood stream and represent the slow and chronic component of the neuroendocrine coordinating system that regulates physiological and behavioral activities.

AGGRESSION AND VIOLENCE

TERMINOLOGIES OF AGGRESSION

To provide a critical evaluation of developments within this area, bringing together material from animal and human studies, it is essential initially to clarify some terminologies. Brain (1990b), as well as others (Buss, 1971; Kutash et al., 1978; Goldstein,

Paul Fredric Brain is at the School of Biomedical Sciences, University College of Swansea, United Kingdom.



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Understanding and Preventing Violence: Volume 2, Biobehavioral Influences Hormonal Aspects of Aggression and Violence Paul Fredric Brain Relationships between aggressive behavior and the endocrine system have been studied intensively in recent years. This interest has occurred presumably because hormones are naturally occurring secretions of the bodies' endocrine or ductless glands, and are perceived as providing possibly reversible (certainly when compared to psychosurgery) therapies for some clinical conditions that include hyperaggressiveness as a symptom. Hormones are transported throughout the body by the blood stream and represent the slow and chronic component of the neuroendocrine coordinating system that regulates physiological and behavioral activities. AGGRESSION AND VIOLENCE TERMINOLOGIES OF AGGRESSION To provide a critical evaluation of developments within this area, bringing together material from animal and human studies, it is essential initially to clarify some terminologies. Brain (1990b), as well as others (Buss, 1971; Kutash et al., 1978; Goldstein, Paul Fredric Brain is at the School of Biomedical Sciences, University College of Swansea, United Kingdom.

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Understanding and Preventing Violence: Volume 2, Biobehavioral Influences 1986; Huntingford and Turner, 1987; Archer, 1988; Klama, 1988; Browne and Archer, 1989), have recently reexamined the nature of aggression for a general audience. In both animal and human sciences, terms such as "aggression" and "violence" are used with enormous flexibility, making it difficult to tie down firm associations with biologic factors (such as hormones). Potential for Harm or Damage The one attribute of aggression about which everyone agrees is that the action must, at least, have the potential for harm or damage. Yet what do we mean by harm? Does harm include only physical harm, or can it include emotional damage or reduced breeding potential? There are behavioral responses that clearly involve harm or potential harm and receive labels other than aggression. For example, harm is definitely involved in predation—an activity that is generally distinguished from aggression by ethologists (students who emphasize behavior's role within the organism's natural environment). Predation is often, but not exclusively, an activity involving members of different species and generally does not involved marked arousal (see below). Harm is also a potential consequence of defensive responses by animals. Consequently, the potential for harm is insufficient cause for an action to be labeled aggression. Having said this, one could make a convincing case for examining the possibility that behaviors more associated with predation deserve consideration in accounts of human violence. Humans are clearly designed to be omnivorous and do (in most cultures) obtain at least part of their food by predatory behaviors. Indeed, a surprisingly large number of infrahuman primate species are also not averse to taking the occasional prey item. Although accounts of human cannibalism are often rendered rather lurid in popular writing and the activity clearly has a semireligious component in many of the cultures that practice (or used to practice) it, this kind of "predation" has been described in a range of cultures. One can also add that the behaviors of certain psychopaths (efficient killing without many visible signs of emotional arousal) seem to fit the ethological description of predation rather than that of social aggression. Perhaps the detailed studies of the biologic factors involved in activities such as mouse killing by rats (e.g., Karli, 1981) are of relevance to some types of human violence even if they are not aggression.

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Understanding and Preventing Violence: Volume 2, Biobehavioral Influences Intentionality Intentionality is another feature necessary, according to some authorities, to identify aggression, but it is generally difficult to establish whether responses are deliberate or not. Some authorities maintain that the motives of the "aggressor" are actually unimportant—what matters is whether the "victim" regards the action as intentional or not. Others go one stage further and maintain that a "dispassionate observer'' (i.e., an individual outside the encounter) is a better judge of aggression. Although the best way of distinguishing between intentional and accidental acts is to consider the probability of the particular event, one must note that different individuals vary in their willingness to see particular responses as intentional. Arousal Biologists generally maintain that aggression has to involve arousal. Arousal is a psychological term applied to evidence of internal changes including alterations in heart rate, respiration, and the distribution of blood in the tissues. Charles Darwin, as early as 1872, advocated that one could deduce something about the arousal state of animals by looking for evidence in postures; the position of hairs, feathers, or combs; and the production of sounds (e.g., spitting, snarling, and crowing). Indeed, one can see animals (e.g. "cornered," subordinate dogs) that are simultaneously fearful and likely to attack. Some authorities (e.g., Scott, 1981; Huntingford and Turner, 1987) have expressed a preference for the term agonistic behavior to cover the range of activities (from overt attack to fleeing, submission, and defense) evident in social conflict. There certainly is some merit in this position since organisms in social conflict encounters generally fluctuate between a range of activities. One should not rely exclusively on this "body language" because one may misinterpret postures and "facial" expressions in animals. In studies of humans, individuals may simulate emotional expressions. Aversiveness A final proviso needed, before some authorities will accept that an act is aggressive, is that the "victim" must regard the action as something to be avoided. This requirement is intended to get around the difficulties of sadomasochism in humans and

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Understanding and Preventing Violence: Volume 2, Biobehavioral Influences the use of "love darts" by snails, which cause slight tissue damage but appear to facilitate courtship in these hermaphrodite animals. AGGRESSION AS A CONCEPT A basic problem with the everyday use of the term aggression is that people generally think they are discussing an entity ("thing") rather than using a concept. We humans essentially have to deal with a complex world in which a vast array of so-called independent variables (potential causes) may be related to an equally large collection of dependent variables (potential consequences). Humans are not computers, and they attempt to make sense of the world by creating intervening variables that link together groups of independent and dependent variables. The concept of aggression is one of these intervening constructs. The trouble with concepts is that they are theoretically definable in many ways—one does not assess a concept by its accuracy but by its usefulness (as an explanatory device). Aggression and Communication Aggression in animals involves communication with any or all of the sensory modalities (as we shall see later, hormones can influence the cues used in such communication and the sensitivities of the sensory systems that respond to them). It has also become apparent that most species have a range of threatening and attack-related activities that can be used in different contexts or for different purposes. Utilities of Aggression It is recognized that animals fight and threaten for a wide range of reasons, such as selection of mates, obtaining exclusive access to an area (territory) that is a prerequisite for breeding, gaining status within a social hierarchy, or defending themselves from conspecifics and predators. Status determines the animal's ease of access to a mate, food, water, or nest sites. One misconception is the view that, because particular animals may employ aggression to obtain a mate, territory, or elevated social status, behaviors receiving the same label in humans necessarily serve one or more of these functions. There is little evidence that humans are intrinsically territorial, always obtain their mates by

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Understanding and Preventing Violence: Volume 2, Biobehavioral Influences crude physical competition, or attain high social status by attacking other individuals. The serious dangers of simplistic extrapolations from animals to humans have been well explored. Different Tests for Animal Aggression Another complicating feature of dealing with animal aggression is the striking diversity of tests said to measure this attribute in particular species (see Brain, 1981, 1989b). In, for example, the laboratory mouse, aggression is said to be generated by pairing preisolated males (intermale aggression), by exposure of paired males or females to unavoidable foot or tail shock (shock-elicited aggression), by arranging for an unfamiliar intruder to enter the nest area of a lactating female with her offspring (maternal aggression), by placing a lactating female (or an animal marked with her urine) into an established group of females or castrated males, by giving the subject the opportunity to kill a locust or a cricket (predatory aggression), and by confining subjects in a narrow tube where they may bite a target suspended in front of them thus activating a telegraph key (instrumental aggression). Thus even in the "simple" mouse, the tests used to generate aggression are so varied (and the responses generated so qualitatively different), it is highly improbable that all measure the same motivation. Certainly, housing conditions (Brain and Benton, 1983), genes (Jones and Brain, 1987), hormones (Brain et al., 1983), and drugs do not have consistent influences across these different tests. It has been argued (Brain, 1984a) to be highly probable that these diverse harm-directed activities variously tap offensive, defensive, or even predatory motivations. In some cases, mixtures of motivations appear to be involved. Support for this view is provided by the use of video analysis, which reveals that, in some "ritualized" responses, vulnerable areas (i.e., the head and ventral surface) of the opponent's body are rarely bitten (in so-called offensive intermale aggression); in other tests, vulnerable areas are frequently bitten (e.g., "defensive" maternal attack on a potentially cannibalistic male intruder), and a third category involves directed killing strategies (e.g., predatory aggression). Perhaps one should limit the term aggression to offensive displays, and thus clearly separate these utilities of attack and threat from defensive and predatory functions? Having said this, one can make a strong case for the detailed investigation of offense, defense and predation in animals being of great relevance to understanding the possible roles

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Understanding and Preventing Violence: Volume 2, Biobehavioral Influences of biological factors in human behavior. One should note that the terms "offensive" and "defensive" are essentially based on functional explanations of particular events; it is not easy to operationalize them. For example, the action of biting can be used in rodents to carry offspring, to eat, to kill prey, to defend a nest site, or to attack a conspecific. Brain (1984a) has suggested that one can define a number of categories of behavior that all employ fighting and/or threat. These include the following: Social conflict: generally intraspecific phenomena involving competition for a substrate (e.g., a mate, territory, social status, or food), the possession of which increases the organism's relative fitness. These are generally ritualized responses in which the potential for serious damage is limited. Parental defense: behaviors in both inter- and intraspecific contexts that serve to protect the attacker's young or nest sites from potentially destructive intruders. Self-defense: behaviors in both inter- and intraspecific contexts that normally serve to protect the organism per se from potential predators or attacking conspecifics. Such behaviors are generally limited to situations where flight is difficult or precluded, and do not involve ritualization. Infanticide: an intraspecific phenomenon involving the killing of young. In males, this may be a method of increasing the individual's reproductive fitness, whereas in females, it is commonly a response to stress or disturbance (recycling of resources?). Predation: an inter- or intraspecific response that involves efficient killing and is often followed by feeding activity. One should note that it is extremely rare (in animal studies) to find purely offensively or purely defensively motivated behavior. The "ethoexperimental" approach to the analysis of animal behavior (Blanchard et al., 1989) seems to offer advantages in studying animal conflict. It basically attempts to fuse the positive features of ethology and experimental psychology by creating laboratory environments that reflect the natural requirements of the feral ancestors of laboratory animals. For example, when dealing with a socially living primate species from an arboreal habitat, it seems appropriate to study mixed-sex groups in complex environments, offering a range of tactile experience. When dealing with animals such as the laboratory rat, it seems appropriate to offer the species the opportunity to construct burrow and nest systems (or to provide an equivalent) and to investigate

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Understanding and Preventing Violence: Volume 2, Biobehavioral Influences the organism under seminocturnal (its major activity occurs at night) conditions. carrying out detailed, inclusive analyses of behavior that simultaneously record other categories of activity in addition to threat and attack (these can be very revealing in determining how a biological manipulation changes aggressiveness). HETEROGENEOUS NATURE OF HUMAN AGGRESSION AND VIOLENCE Aggressive behavior is certainly no less heterogeneous in our own species. The classification of Buss (1971), based on three dichotomies, provides a clear indication of the diversity of human aggression as viewed through the eyes of a social psychologist. Aggression, according to Buss, may be physical or verbal, active or passive, and direct or indirect. Although it is easy to think of animal analogies for punching, stabbing, or shooting (physical/active/direct aggression), it is much harder to think of animal analogies for "failing to carry out a necessary task" (physical/passive/indirect aggression) or "refusing consent" (verbal/passive/indirect aggression). Obviously, the social psychologist includes a much wider range of activities under the heading aggression than does the biologist. This diversity of human aggression, has led to an enormous range of methods for assessing the attribute in our species. These broadly fall into two categories. In the first, behavior is assessed in situ by seeking the opinions of peers or by questionnaires. These include examining interactions in preschool play groups, determining the reactions (verbally or physiologically) to films or written material, creating experimental conflict situations (such as use of the "hostility" machine), studying individuals in natural high-stress situations, looking at participation in group activities in which hostile outcomes are probable (certain sports and committees), responding as observers in sporting situations (e.g. "football hooliganism"), and even investigating participation in riots. The second category involves relating physiological events to behavioral characteristics largely determined on the basis of past events. These include studies of convicted criminals (here the material is often divided into largely sexual and nonsexual, and a distinction is made between "impulsive" and "premeditated" crime). One should note here that studying the "same'' crime does not always mean that one is dealing with the same phenomenon (e.g., rape is said by several authorities such as Groth, 1979, to have several etiologies and is generated by a plethora of influences).

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Understanding and Preventing Violence: Volume 2, Biobehavioral Influences Similar critiques could be advanced for homicide, assault, etc., before adding the complications introduced by procedures such as plea bargaining. Other studies involve investigations of persons resisting attack on themselves or on property, analysis of individuals with a variety of psychiatric and clinical disorders, and comparisons of male and female individuals (e.g., variations in "rough-and-tumble play" and correlating premenstrual tension with crime). The focus of this review is human violence. Violence is used almost interchangeably with aggression in most reviews. One might add that violence is a term often applied to aggressive actions that attract greater than normal social disapproval. In this respect, the term clearly fulfills Felson's role of being used to label behavior that transgresses normative values. Obviously, since judgments of sections of society may be involved in determining which behaviors receive the labels aggression and violence, relating hormones to such human activities is not easy. It is doubtful whether one can always differentiate adaptive forms of aggression from maladaptive violent and aggressive acts, because this implies a very accurate knowledge of the motivations of all participants at all times. The definition of the Panel on the Understanding and Control of Violent Behavior of violent human behavior as "threatened, attempted, or completed intentional infliction of physical harm by persons against persons" is eminently reasonable and is broadly the type of behavior referred to throughout this account. One must note, however, that even within such a framework, "appropriate" vigor grades into violence. Obviously, areas of contention include physical punishment of children and activities in a range of contact sports. HORMONES AND AGGRESSION It is necessary to note initially that what we call aggression is (like any other behavioral concept) influenced by diverse factors that are difficult (impossible?) to disentangle. These include biological factors (i.e., genes, neural systems, neurotransmitters, and hormones); situational determinants (i.e., the environmental or social context); and the accumulated experiences of individuals. Figure 1 is a schema of the relationship between biology and behavior. If one looks at interindividual forms of aggression, one is

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Understanding and Preventing Violence: Volume 2, Biobehavioral Influences FIGURE 1 Schema showing the relationship(s) between biology and aggression: factors to be considered in rating "simple" interactions between individuals. SOURCE: Brain (1989a). really dealing with some quite complex interactions between biology and experience. Some of these effects are mediated by changes in aggressive motivation, some by influencing other behaviors that compete for expression with the aggression, others by changing the social signals that organisms direct toward each other, and yet others by the way in which individuals perceive those social signals. There are also changes over time and the impact of the particular environment to consider. One has to add to this complex mix that whether one chooses to call a behavior aggression or not, is based the observer's value judgment. It is consequently highly improbable that one will find simple relationships between any one biological factor and expressed behavior. NEUROENDOCRINOLOGY Although the neural and endocrine systems have tended to be regarded separately, it is currently thought by a variety of authorities that they are best considered components with differing characteristics (in terms of the speed, duration, and diffuseness of their actions) of a single neuroendocrine coordinating system. Certainly there is an intimate relationship between neural and endocrine factors. Indeed, hormones, neurotransmitters, and neuromodulators can all be defined as information-transferring molecules. Cues received and integrated by the central nervous system (CNS) can

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Understanding and Preventing Violence: Volume 2, Biobehavioral Influences be passed by the neural elements to specialized neurosecretory cells that essentially convert nerve impulses into hormonal output. These cells (in evolutionary terms, the oldest glands) are modified neurons (with many of the elements of such cells) that secrete protein-derived (peptidergic) or amino-derived (aminergic) material. This material may be transported along the modified axonal elements that end in close association with blood vessels (some of these are called neurohemal organs: e.g., the posterior lobe of the pituitary). It is now apparent that many endocrine glands are innervated by conventional neurons, which suggests that direct neural input can modify their secretory activity. Some neurosecretory cells may have direct effects on muscles and other effector organs, such as exocrine ("ducted") glands. Although some endocrine glands are primarily controlled by direct neural input (the adrenal medulla), others are controlled by tropic hormones from the pituitary (the adrenal cortex, the gonads, and the thyroid), and a third category largely responds to blood-borne metabolites (the pancreas and the parathyroids). Many endocrine systems maintain homeostasis (a balance vis-à-vis the internal environment) by employing negative feedback mechanisms (see Figure 2). In some cases, positive feedback mechanisms may also operate (e.g., involvement of luteinizing hormone (LH) in ovulation in mammals). DEFINING "HORMONE" As mentioned earlier, hormones are secretions of endocrine glands that are passed into the bloodstream and are accumulated by target tissues (including the CNS), where they induce particular physiological or behavioral responses. The use of the term hormone originally implied (1) a natural chemical structure that had been extracted from a recognized endocrine gland and (2) the use of the blood system as the transport mechanism employed to reach the target tissue. The term hormone has, however, recently become less precise. Synthetic hormones, fragments of peptide factors, analogues of hormones, and parahormones (e.g., prostaglandins and opioids) have been included within this heading. Is the neurally located material that is immunoreactive to an anti-body to adrenocorticotropic hormone (ACTH) truly a hormone, even if the substance is chemically identical to ACTH? This peptide may never get near the bloodstream. Are the bodies' own pain killers, enkephalins and endorphins, hormones? They may be derived from the peptide hormone ß-lipotropin, but are they

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Understanding and Preventing Violence: Volume 2, Biobehavioral Influences FIGURE 2 Negative feedback arrangement evident between an endocrine gland and a tropic gland (e.g., the adrenal cortex and the anterior pituitary). SOURCE: Brain (1989c). Reprinted by permission of Kluwer Academic Publishers. best regarded as degradation products? A certain degree of flexibility seems consequently appropriate in this area at the present time. The main systems discussed in this review are the HPG (hypothalamus-pituitary-gonadal axis, which includes hypothalamic luteinizing hormone (LH) releasing factor (LHRF), gonadotropins from the pituitary, and sex steroids from the gonads) and the HPA

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Understanding and Preventing Violence: Volume 2, Biobehavioral Influences Edwards, D.A. 1968 Mice: Fighting by neonatally androgenized females. Science 161:1027–1028. 1969 Early androgen stimulation and aggressive behavior in male and female mice. Physiology and Behavior 4:333–338. 1971 Neonatal administration of androstenedione, testosterone or testosterone propionate: Effects on ovulation, sexual receptivity, and aggressive behavior in female mice. Physiology and Behavior 6:223–228. Ehrhardt, A.A. 1969 Zur wirkung fötaler Hormone auf Intelligenz und geschlechts-spezifisches Verhalten. Doctoral thesis, Universitat Dusseldorf. Ehrhardt, A.A., and S.W. Baker 1974 Fetal androgen, human CNS differentiation and behavior sex differences. Pp. 53–76 in R.C. Friedman, R.M. Richart, R.L. Van de Wiele, eds., Sex Differences in Behavior. New York: John Wiley and Sons. Ehrhardt, A.A., and J. Money 1967 Progestin-induced hermaphroditism: IQ and psychosexual identity in a study of ten girls. Journal of Sex Research 3:83–100. Elias, M. 1981 Serum cortisol, testosterone, and testosterone-binding globulin responses to competitive fighting in human males. Aggressive Behavior 7:215–224. Epple, G. 1978 Lack of effects of castration on scent marking displays and aggression in a South American primate Saguinus fuscicollis. Hormones and Behavior 11:139–150. Farrington, D.P. 1989 Early predictors of adolescent aggression and adult violence. Violence and Victims 4:79–100. Field, L.H., and M. Williams 1970 The hormonal treatment of sexual offenders. Medicine, Science, and the Law 10:27–34. Floody, O.R., and D.W. Pfaff 1977 The hormonal basis for fluctuations in female aggressiveness correlated with estrous state. Journal of Comparative and Physiological Psychology 91:443–464. Frodi, A. 1977 Sexual arousal, situational restrictiveness, and aggressive behavior. Journal of Research in Personality 11:48–58. Gadelman, R. 1981 Androgens and fighting behavior. Pp. 215–230 in P.F. Brain and D. Benton, eds., The Biology of Aggression. Alphen aan den Rijn, Holland: Sijthoff and Noordhoof b.v.

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Understanding and Preventing Violence: Volume 2, Biobehavioral Influences Gladue, B.A., M. Boechler, and K.D. McCaul 1989 Hormonal response to competition in human males. Aggressive Behavior 15:409–422. Goldman, L., and H.H. Swanson 1975 Developmental changes in pre-adult behavior in confined colonies of Golden hamsters. Developmental Psychology 8:137–150. Goldstein, J.H. 1986 Aggression and Crimes of Violence. New York: Oxford University Press . Gotz, F., W. Rohde, and C. Dorner 1990 Neuroendocrine differentiation of sex-specific gonadotrophin secretion, sexual orientation, and gender role behavior. Pp. 167–194 in M. Haug, P.F. Brain, and C. Aron, eds., Heterotypical Behavior in Man and Animals. London: Chapman and Hall. Goy, R.W. 1968 Organizing effects of androgen on the behavior of rhesus monkeys. Pp. 12–31 in R.P. Michael, ed., Endocrinology and Human Behavior. London: Oxford University Press. Goy, R.W., and M. Roy 1990 Heterotypic sexual behavior in female mammals. Pp. 71–97 in M. Haug, P.F. Brain, and C. Aron, eds., Heterotypical Behaviour in Man and Animals. London: Chapman and Hall. Green, R., R.E. Whalen, B. Butley, and C. Battie 1972 Dominance hierarchy in squirrel monkeys: Role of gonads and androgen on genital display and feeding order. Folia Primatologia 18:185–195. Groth, A.N. 1979 Men Who Rape: The Psychology of the Offender. New York: Plenum Press. Hamburg, D.A. 1971 Recent research on hormonal factors relevant to human aggressiveness. International Social Science Journal 23:36–47. Hammer, R.P., K.M. Hori, P. Cholvanich, D.C. Blanchard, and R.J. Blanchard 1990 Opiate, serotonin, and benzodiazepine receptor systems in rat brain defense circuits. Pp. 201–217 in P.F. Brain, S. Parmigiani, R. Blanchard, and D. Mainardi, eds., Fear and Defence. Chur, Switzerland: Harwood Academic Press, gmbh. Harding, C.F. 1981 Social modulation of circulating hormone levels in the male. American Zoologist 21:223–231. 1989 Interactions of androgens and estrogens in the modulation of social behavior in male songbirds. Pp. 558–579 in R.J. Blanchard, P.F. Brain, D.C. Blanchard, and S. Parmigiani, eds., Ethoexperimental Approaches to the Study of Behavior. Dordrecht, Holland: Kluwer Academic Press. Hasan, S.A., P.F. Brain, and D. Castano 1988 Studies of the effects of Tamoxifen ICI 46474 on agonistic encounters

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Understanding and Preventing Violence: Volume 2, Biobehavioral Influences between pairs of intact mice. Hormones and Behavior 22:178–185. Haug, M., and P.F. Brain 1989 Psychobiological influences of attack on lactating females: A varient on "typical" house mouse aggression. Pp. 205–222 in P.F. Brain, D. Mainardi, and S. Parmigiani, eds., House Mouse Aggression . Chur, Switzerland: Harwood Academic Publishers gmbh. Hays, S.E. 1978 Strategies for psychoendocrine studies of puberty. Psychoeuro-endocrinology 3:1–15. Hellhammer, D.H., W. Hubert, and T. Schurmeyer 1985 Changes in saliva testosterone after psychological stimulation in men. Psychoeuroendocrinology 10:77–81. Hucklebridge, F.H., L. Gamal el Din, and P.F. Brain 1981 Social status and the adrenal medulla in the house mouse (Mus musculus L.). Behavioral and Neural Biology 33:345–363. Huntingford, F., and A. Turner 1987 Animal Conflict. London: Chapman and Hall. Jeffcoate, W.J., N.B. Lincoln, C. Selby, and M. Herbert 1986 Correlation between anxiety and serum prolactin in humans. Journal of Psychosomatic Research 30:217–222. Jones, S.E., and P.F. Brain 1987 Performances of inbred and outbred laboratory mice in putative tests on aggression. Behavior Genetics 17:87–96. Joslyn, W.D. 1973 Androgen-induced social dominance in infant female rhesus monkeys. Journal of Child Psychology and Psychiatry 14:137–145. Julian, T., and P.C. McKenry 1989 Relationship of testosterone to men's family functioning at midlife: A research note. Aggressive Behavior 15:281–289. Kaplan, H., and S.O. Hyland 1972 Behavioral development in the Mongolian gerbil Meriones unguiculatus. Animal Behaviour 20:147–154. Karli, P. 1981 Conceptual and methological problems associated with the study of brain mechanisms underlying aggressive behavior. Pp. 323–362 in P.F. Brain and D. Benton, eds., The Biology of Aggression. Alphen aan den Rijn, Holland: Sijthoff and Noordhoof b.v. Kelley, D.B. 1981 Social signals—an overview. American Zoologist 21:111–116. Kesler, P., R. Green, S.J. Finch, and K. Williams 1980 Prenatal "female hormone" administration and psychosexual development in human males . Psychoeuroendocrinology 5:269–285.

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Understanding and Preventing Violence: Volume 2, Biobehavioral Influences Klama, J. 1988 Aggression: Conflict in Animals and Humans Reconsidered. Burnt Mill, Harlow, England: Longman Group. Kutash, I.L., S.B. Kutash, L.B. Schlesinger and Associates, eds. 1978 Violence: Perspectives on Murder and Aggression. San Francisco: Jossey-Bass Publishers. Langevin, R., J. Bain, M. Ben-Aron, R. Coulthard, D. Day, L. Handy, G. Heasman, S. Hucker, J. Purins, V. Roper, A. Russan, C. Webster, and G. Wortzman 1985 Sexual aggression: Constructing a predictive equation. Pp. 50–93 in R. Langevin, ed., Erotic Preference Gender Identity and Aggression in Men. Hillsdale, N.J.: Lawrence Erlbaum Associates. Larsson, K., P. Sodersten, and C. Beyer 1973 Induction of male sexual behavior by oestradiol benzoate in conjunction with dihydrotestosterone. Journal of Endocrinology 57:563–564. Laskowski, W. 1954 Einige verhaltensstudien an Platypoecilus variatus. Biologisches Zentralblatt 73:429–438. Lehrman, D.S. 1965 Interaction between internal and external environments in the regulation of the reproductive cycle of the ring dove. Pp. 355–380 in F.A. Beach, ed., Sex and Behavior. New York: John Wiley & Sons. Leshner, A.I. 1980 Interaction of experience and neuroendocrine factors in determining behavioral adaptations to aggression. Pp. 427–438 in P.S. McConnell et al., eds., Progress in Brain Research, Vol. 53. Amsterdam, Holland: Elsevier/North-Holland. 1981 The role of hormones in the control of submissiveness. Pp. 309–322 in P.F. Brain and D. Benton, eds., Multidisciplinary Approaches to Aggression Research. Amsterdam, Holland: Elsevier/N North-Holland. Leshner, A.I., and K.E. Roche 1977 Comparison of the effects of ACTH and lysine vasopressin on avoidance-of-attack in mice. Physiology and Behavior 18:879–833. Levander, S., A. Mattsson, D. Schalling, and A. Dalteg 1987 Psychoendocrine patterns within a group of male juvenile delinquents as related to early psychosocial stress, diagnostic classification, and follow-up data. Pp. 235–252 in D. Magnusson and A. Ohman, eds., Psychopathology: An International Perspective. Orlando, Florida: Academic Press. Lincoln, G.A. 1987 Long-term stimulatory effects of a continuous infusion of LHRH

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Understanding and Preventing Violence: Volume 2, Biobehavioral Influences Meyer-Bahlburg, H.F.L., and A.A. Ehrhardt 1982 Prenatal sex hormone and human aggression: A review and new data on progestogen effects. Aggressive Behavior 8:39–62. Meyerson, B.J. 1967 Relationship between the anesthetic and gestagenic action and oestrous behavior-inducing activity of different progestins. Endocrinology 81:369–374. 1983 Endorphin-monoamine interaction and steroid-dependent behavior. Pp. 111–117 in J. Balthazart, E. Prove, and R. Gilles, eds., Hormones and Behavior in Higher Vertebrates. Berlin: Springer-Verlag. Miczek, K.A., and M. Krsiak 1981 Pharmacological analysis of attack and flight. Pp. 341–354 in P.F. Brain and D. Benton, eds., Multidisciplinary Approaches to Aggression Research. Amsterdam, Holland: Elsevier/North Holland. Mirsky, A.F. 1955 The influence of sex hormones on social behavior in monkeys. The Journal of Comparative and Physiological Psychology 48:327–335. Money, J. 1990 The development of sexuality and eroticism in humankind. Pp. 127–166 in M. Haug, P.F. Brain, and C. Aron, eds., Heterotypical Behavior in Man and Animals. London: Chapman and Hall. Money, J., and A.A. Ehrhardt 1972 Man and Woman, Boy and Girl. Baltimore, Md.: Johns Hopkins University Press. Money, J., and C. Ogunro 1974 Behavioral sexology: Ten cases of genetic male intersexuality with impaired prenatal and pubertal androgenization. Archives of Sexual Behavior 3:181–205. Money, J., and M. Schwartz 1976 Fetal androgens in the early treated adrenogenital syndrome of 46 hermaphroditism: Influence on assertive and aggressive types of behavior. Aggressive Behavior: 2:19–30. Monti, P.M., W.A. Brown, and D.P. Corriveau 1977 Testosterone and components of aggressive and sexual behavior in man. American Journal of Psychiatry 134:692–694. Mugford, R.A. 1974 Androgenic stimulation of aggression eliciting cues in adult opponent mice castrated at birth, weaning or maturity. Hormones and Behavior 5:93–102. Naess, O., and A. Attramadal 1974 Uptake and binding of androgens in the anterior pituitary gland, hypothalmus, preoptic and brain cortex of rats. Acta endocrinologica Kbn. 76:417–430.

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Understanding and Preventing Violence: Volume 2, Biobehavioral Influences Naftolin, F., and R.J. Ryan 1975 The metabolism of androgen in central neuroendocrine tissues. Journal of Steroid Biochemistry 6:993–997. O'Connor, M., and H.W.G. Baker 1983 Depo-medroxy progesterone acetate as an adjunctive treatment in three aggressive schizophrenic patients. Acta Psychiatrica Scandinavica 67:399–402. Olweus, D. 1984 Development of stable aggressive reaction patterns in males. Pp. 103–137 in R.J. Blanchard and D.C. Blanchard, eds., Advances in the Study of Aggression, Vol. 1. Orlando, Florida.: Academic Press. 1986 Aggression and hormones: Behavioral relationship with testosterone and adrenaline. Pp. 51–72 in D. Olweus, J. Block, and M. Radke-Yarrow, eds., Development of Antisocial and Prosocial Behavior: Research, Theories, and Issues. New York: Academic Press. Olweus, D., A. Mattsson, D. Schalling, and H. Low 1980 Testosterone, aggression, physical and personality dimensions in normal adolescent males. Psychosomatic Medicine 42:253–269. 1988 Circulating testosterone levels and aggression in adolescent males. Psychosomatic Medicine 50:261–272. Owens, N.W. 1975 A comparison of aggressive play and aggression in free-living baboons Papio anubis. Animal Behaviour 23:757–765. P'an, S.Y., and G.D. Laubach 1964 Steroid central depressants. Pp. 415–475 in R.I. Dorfman, ed., Methods in Hormone Research, Vol. 3. New York: Academic Press. Parrott, R.F. 1975 Aromatizable and 5a-reduced androgens: Differentiation between central and peripheral effects on male rat sexual behavior. Hormones and Behavior 6:99–108. 1976 Homotypical sexual behavior in gonadectomized female and male rats treated with 5a-19-hydroxytestosterone: Comparison with related androgens. Hormones and Behavior 7:207–215. Payne, A.P. 1976 A comparison of the effects of neonatally administered testosterone, testosterone propionate and dihydrotestosterone on aggressive and sexual behavior in female Golden hamster. Journal of Endocrinology 69:23–31. Payne, A.P., and H.H. Swanson 1972 The effect of sex hormones on the agonistic behavior of the male Golden hamster. Physiology and Behavior 8:687–691. Pfaff, D.W. 1971 Steroid sex hormones in the rate brain: Specificity of uptake

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Understanding and Preventing Violence: Volume 2, Biobehavioral Influences and physiological effects. Pp. 103–112 in C.H. Sawyer and R.A. Gorski, eds., Steroid Hormones and Brain Function. Berkeley.: University of California Press. Poole, T.B. 1985 Social Behavior in Mammals. New York: Chapman and Hall. Popova, N.K., N.N. Voitenko, S.I. Pavlova, E.V. Naumenko, and D.K. Belyaev 1980 Genetics and phenogenetics of hormonal characteristics in animals. VII. Relationship between brain serotonin and hypothalamic-pituitary-adrenal axis under emotional stress in domesticated and non-domesticated silver foxes. Genetics 16:1865–1870 (in Russian). Poshivalov, V.P. 1982 Ethological analysis of neuropeptides and psychotropic drugs: Effects on intraspecies aggression and sociability of isolated mice. Aggressive Behavior 8:355–369. Reinisch, J.M. 1981 Prenatal exposure to synthetic progestins increases potential for aggression in humans. Science 211:1171–1173. Resko, J.A. 1974 The relationship between fetal hormones and the differentiation of the central nervous system in primates. Pp. 211–222 in W. Montagna and W.A. Sadler, eds., Reproductive Behavior. New York: Plenum Press. Roberts, S.S. 1990 Murder, mayhem, and other joys of youth. The Journal of NIH Research 2:67–72. Rose, R.M., T.P. Gordon, and I.S. Bernstein 1978 Diurnal variation in plasma testosterone and cortisol in rhesus monkeys living in social groups. Journal of Endocrinology 76:67–74. Salvador, A., V. Simon, F. Suay, and L. Llorens 1987 Testosterone and cortisol responses to competitive fighting in human males: A pilot study. Aggressive Behavior 13:9–13. Schallert, T. 1977 Reactivity to food odours during hypothalamic stimulation in rats not experienced with stimulation-induced eating. Physiology and Behavior 18:1061–1066. Scott, J.P. 1981 The evolution of function in agonistic behavior. Pp. 129–157 in P.F. Brain and D. Benton, eds., Multidisciplinary Approaches to Aggression Research. Amsterdam, Holland: Elsevier/North-Holland Biomedical Press. Schultz, L., and R. Lore 1987 Jolly fat rats? The effects of diet-induced obesity on fighting. Aggressive Behavior 13:359–366.

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Understanding and Preventing Violence: Volume 2, Biobehavioral Influences Selmanoff, M.K., J.E. Jumonville, S.C. Maxson, and B.E. Ginsburg 1975 Evidence for a Y-chromosome contribution to an aggressive phenotype in inbred mice. Nature 253-529–530. Sharp, P.J., J. Culbert, and J.W. Wells 1977 Variations in stored and plasma concentrations of androgens and luteinizing hormone during sexual development in the cockerel. Journal of Endocrinology 74:467–476. Sheard, M.H. 1987 Psychopharmacology of aggression in humans. Pp. 257–266 in B. Olivier, J. Mos, and P.F. Brain, eds., Ethopharmacology of Agonistic Behavior in Animals and Humans. Dordecht, Holland: Martinus Nijhoff Publishers. Sherwin, B.B., M.M. Gelfand, and B. Brender 1985 Androgen enhances sexual motivation in females: A prospective crossover study of sex steroid administration in the surgical menopause. Psychosomatic Medicine 47:339–351. Siegel, J.M. 1986 The multidimensional anger inventory. Journal of Personality and Social Psychology 51:191–200. Sigg, E.B., C. Day, and C. Colombo 1966 Endocrine factors in isolation-induced aggressiveness in rodents. Endocrinology 78:679–684. Silver, R. 1983 Biparental care in birds: Mechanisms controlling incubation bout duration. Pp. 451–462 in J. Balthazart, E. Prove, and R. Gilles, eds., Hormones and Behavior in Higher Vertebrates. Berlin: Springer-Verlag. Simon, N.G., and R.E. Whalen 1986 Hormonal regulation of aggression: Evidence for a relationship among genotype, receptor binding and behavior sensitivity to androgen and estrogen. Aggressive Behavior 12:255–266. Simon, V., M. Martinez, D. Castano, P.F. Brain, and S. Hasan 1987 Studies on the effects of the anti-androgen cyproterone acetate on social encounters between pairs of male mice. International Journal of Neuroscience 41:231–240. Steiner, M., and B.J. Carroll 1977 The psychobiology of premenstrual dysphoria: Review of theories and treatments. Psychoneuroendocrinology 2:321–325. Straus, M. 1979 Measuring intrafamily conflict violence: The conflict tactics CT scales. Journal of Marriage and the Family 41:75–88. Suchowsky, G.K., L. Pegrassi, and A. Bonsignori 1969 The effect of steroids on aggressive behavior in isolated male mice. Pp. 161–171 in S. Garattini and E.B. Sigg, eds., Aggressive Behavior. Amsterdam, Holland: Excerpta Medica Foundation. Suffrin, G., and D.S. Coffey 1973 A new model for studying the effect of drugs on prostatic growth

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Understanding and Preventing Violence: Volume 2, Biobehavioral Influences I. Antiandrogens and DNA synthesis. Investigative Urology II:45–54. Susman, E.J., G. Inoff-Germain, E.D. Nottelmann, D.L. Loriaux, G.B. Cutler, and G.P. Chrousos 1987 Hormones, emotional dispositions, and aggressive attributes in young adolescents. Child Development 58:1114–1134. Svare, B. 1977 Maternal aggression in mice: Influence of the young. Biobehavioral Reviews I:151–164. 1989 Recent advances in the study of female aggressive behavior in mice. Pp. 135–159 in P.F. Brain, D. Mainardi, and S. Parmigiani, eds., House Mouse Aggression. Chur, Switzerland: Harwood Academic Publishers gmbh. Svare, B.B., and M.A. Mann 1983 Hormonal influences on maternal aggression. Pp. 91–104 in B.B. Svare, ed., Hormones and Aggressive Behavior. New York: Plenum Press. Symons, J. 1973 Aggressive Play in a Free-Ranging Group of Rhesus Monkeys Macaca mulatta. Ph.D. thesis, University of California, Berkeley. Thomas, D.A., R.J. Talala, and R.J. Barfield 1981 Effect of devocalization of the male on mating behavior in rats. Journal of Comparative and Physiological Psychology 95:630–637. Tiwary, C.M. 1974 Testosterone, LHRH, and behavior. Lancet May 18:993. Tonks, C.M. 1977 Psychiatric aspects of endocrine disorders. Practioner 218:526–531. Valenti, G., and M. Mainardi 1989 Aggressiveness in mice and thyroid hormones. Pp. 293–309 in P.F. Brain, D. Mainardi, and S. Parmigiani, eds., House Mouse Aggression . Chur, Switzerland: Harwood Academic Publishers gmbh. van de Poll, N.E., and H. van Dis 1977 Hormone induced lordosis and its relation to masculine sexual activity in male rats. Hormones and Behavior 8:17–7. van de Poll, N.E., J.P.C. de Bruin, H. van Dis, and H.G. Van Oyen 1978 Gonadal hormones and the differentiation of sexual and aggressive behavior and learning in the rat. Pp. 309–327 in Progress in Brain Research, Vol. 18. Amsterdam: Elsevier/North-Holland. van de Poll, N.E., F. de Jonge, H.G. Van Oyen, J. Van Pelt, and J.P.C. de Bruin 1981 Failure to find sex differences in testosterone-activated aggression in two strains of rats. Hormones and Behavior 15:94–105.

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Understanding and Preventing Violence: Volume 2, Biobehavioral Influences vom Saal, F.S. 1983 Models of early hormonal effects on intersex aggression in mice. Pp. 197–222 in B.B. Svare, ed., Hormones and Aggressive Behavior. New York: Plenum Press.