Some subordinates are more equal than others: Play fihgting amongst adult subordinate male rats

Adult male rats living together form dominance relationships, with one dominant and the remainder adopting subordinate roles. In previous studies, it was shown that in adult male pairs, the subordinate rat initiates more playful contacts and retains a more juvenile response to the playful contacts by the dominant. In this experiment, triads were used to examine the play between subordinate males. The subordinates directed fewer playful contacts to each other than to the dominant rat, and there was a symmetrical play relationship between the subordinates. After the dominant was removed from the colony, one subordinate became the dominant. Playful interactions amongst these pairs increased, with the subordiante intiating more playful contacts than the dominant. Furthermore, from a similarly low frequency of juvenile-type response to playful contact to each other when in triads, the subordinate in the dyads increased its frequency of juvenile responses to the dominant partner. This supports the hypothesis that the playful behavior of subordinate male rats towards the dominant is an adaptive response, playful behavior of subordinate male rats towards the dominant is an adaptive response, serving a “friendship maintenance” function. Finally, when in triads, one subordinate was more playful with the dominant than the other subordinate. It was the least playful subordinate that was the most likely to become the dominant. This sugests that within a colony, not all subordinates are the same. © 1993 Wiley-Liss, Inc.     

Juvenilized play fighting in subordinate male rats

As pairs of male juvenile sibling rats that are housed together become sexually mature, they develop a dominance-subordinance relationship. These dominance relationships appear to be reflected in the play fighting of the pairmates both as juveniles and as young adults, in that the seemingly subordinate partner initiates more playful attacks at both ages. However, as adults, even though it is the subordinate that initiates more playful attacks, it is the subordinate that is pinned on his back by the partner most often. Dominant pairmates were found to switch to defensive patterns typically found in adult males. In contrast, the subordinates, when contacted on the nape, were more likely to retain the juvenile pattern of turning over to supine. Therefore, the subordinate pairmate of an adult pair of male siblings both initiates more playful attacks and defends itself in a more juvenile manner than its dominant partner, and this leads to it being pinned more frequently. This pattern of behavior by subordinate rats is suggested to function as a friendship maintenance mechanism permitting co-existence in multimale colonies.     

Playful defensive responses in adult male rats depend on the status of the unfamiliar opponent

Adult male rats reared as pairmates from weaning were tested in a neutral arena with both members of another pair (one at a time). The unfamiliar pairs were found to engage in play fighting, although they were more likely to escalate the encounter into serious fighting than were pairs of familiar rats. Based on their within‐home pair behavior, each pairmate was designated as a dominant or a subordinate. When the test encounters between unfamiliar males were analyzed with regard to whether the pairings consisted of two dominants, two subordinates, or a mixed pair, the pattern of play fighting was found to be attenuated. Both dominants and subordinates were more likely to initiate playful encounters, to respond defensively during these encounters, and to do so using adult‐typical tactics of defense when paired with an unfamiliar rat that was dominant in its home cage. The mechanisms by which the home status of unfamiliar male rats can be identified by another male are discussed, particularly with regard to the role that play fighting may serve for this function. It is concluded that the data support the hypothesis that play fighting can be used by adult rats for social testing, which in this case seems to involve ascertaining the opponent's fighting capability. Aggr. Behav. 25:141–152, 1999. © 1999 Wiley‐Liss, Inc.     

Neonatal testosterone augmentation increases juvenile play fighting but does not influence the adult dominance relationships of male rats

Neonatal male rats were either injected subcutaneously with testosterone propionate (TP) or oil vehicle. When weaned, each treated pup was paired with an untreated male sibling. The play fighting of TP-and oil-treated rats were compared at the juvenile phase (30–36 days), and in adulthood (84–90 days). In the juvenile phase, the rate of initiating playful attacks was significantly greater for TP-treated rats. Playful defense in response to such attacks did not differ between TP- and oil-treated rats. At the completion of the study, cortical thickness was measured for all the groups of rats. Oil treatment decreased overall cortical thickness relative to untreated pairmates, whereas TP treatment did not. Both oil and TP treatment abolished the asymmetry in hemispheric thickness, which was present in the untreated pairmates. The reversal of at least one of these injectioninduced changes in the cortex by TP provided independent evidence for the effectiveness of the TP treatment. As adults, neither the TP treatment nor the oil treatment influenced which pairmate became dominant. Dominance was judged by which pairmate initiated less playful attacks. Therefore, it is concluded that the early neonatal testosterone surge is not likely to be a factor in influencing the behaviors that lead to adult dominance. In contrast, play fighting is influenced by hormonal events in this early neonatal phase. It thus appears that play fighting and the aggressive systems subserving dominance relationships are differentially controlled. © 1992 Wiley-Liss, Inc.     

Pregrouping aggression and defense scores influences alcohol consumption for dominant and subordinate rats in visible burrow systems

Five male/two female rat colonies were established in visible burrow systems, with males selected for pregrouping attack scores and also evaluated in open field and cat odor tests. Dominant-subordinate pregrouping attack differences suggested that the males becoming dominant are those showing more persistent and higher level attack. For six colonies showing dominant-subordinate behavioral differences, pregrouping defense tests failed to predict subordinate status. However, pregrouping defense scores were reliably correlated with subordinate pre-postgrouping change scores for voluntary ethanol consumption. Subordinates showed higher ranked ethanol consumption than dominants, but these groups were not different on pregrouping ethanol consumption. Subordinate postgrouping ethanol consumption was positively correlated with pregrouping attack toward an adult intruder, consonant with previous findings that highly aggressive subordinates are the targets of more intense attack by dominants. These results provide further support for a view that subordination stress increases voluntary ethanol consumption in male rats and suggest some additional individual differences factors that may be involved in increased ethanol consumption for male subordinates. © 1992 Wiley-Liss, Inc.     

Effects of fluoxetine on play dominance in juvenile rats

In a series of three studies, we investigated the influence of a selective serotonin re-uptake inhibitor (fluoxetine) on the rough-and-tumble play of juvenile rats. In Experiment 1, both members of eight pairs of solitary-housed juvenile rats received either vehicle, 2.5, 5, or 10 mg/kg fluoxetine in a counterbalanced within-subject design 20 min before being allowed to play for 5 min periods on four successive test days. The 5 and 10 mg/kg pretreatments significantly reduced incidence of pins during play without affecting dorsal contacts. In Experiment 2, one member of each of 19 established play pairs received 5 mg/kg fluoxetine 20 min before play, while the other member received vehicle. Dominant rats showed no reduction in pins as a result of fluoxetine treatment, but subordinate rats who received fluoxetine exhibited significant reductions in pins. Subsequent dyadic analyses indicated that in pairs where the subordinate animal received fluoxetine, dominant animals maintained their pinning advantage over the 10 days of testing, but in pairs where the dominant animals received fluoxetine, this pinning asymmetry diminished. In Experiment 3, we replicated the above procedure with inexperienced play pairs, to control for the effects of prior social learning. Fluoxetine treatment (5 mg/kg) significantly reduced both pins and dorsal contacts in all treated rats. The results indicate that fluoxetine can reduce the playful pins of juvenile rats, but that prior social learning mediates the strength of these effects. © 1996 Wiley-Liss, Inc.     

Aversive potency of urine from dominant and subordinate male laboratory mice (Mus musculus): Resolution of a conflict

The present study assessed the aversive potency of urine collected from male albino mice that had been clearly identified as dominants/winners or subordinates/losers of paired aggression tests and then housed either individually or in a quasi-paired situation in which only a wire-mesh divider separated the two mice. This divider permitted constant visual, olfactory, auditory, and some tactile contact. The responses of individually tested, group-housed males were recorded when half of the substrate in a test box was treated with either water or one of the four urine types; the other half remained untreated. Significant preferences for the untreated half were found when the urine of winners or losers housed in individual metabolism cages or that of pair-housed dominants was used as the test stimulus. On the other hand, neither water nor the urine of cohabiting subordinate males was avoided. The present findings confirmed our earlier reports that the urine of dominant male mice was aversive, whereas that of their cohabiting subordinate partners was not. They also identified Sawyer's [1978] procedure of housing winners and losers in individual cages, with the consequent interruption of social contact as the likely reason for his failure to replicate our reports that subordinate male urine lacked aversive properties. The territorial implications of the aversive factor and other urinary signals are discussed.     

Attack and defense in conspecific fighting in tree shrews (Tupaia belangeri)

Principles of conspecific defense have been analyzed for rodents, in which specific target sites for biting by attackers on defenders serve as an important determinant of the actions involved in both attacker and defender behavior. In an effort to determine the generality of these principles, attack and defensive behaviors and target sites for biting attack were evaluated in a nonrodent species, the tree shrew (Tupaia belangeri). Brief daily and repeated conspecific dyadic encounters between adult, socially experienced males (dominants, attackers), and adult, socially naive males (subordinates, defenders) that had been transferred into the territory of the dominants, produced a polarization of attack and defense. The dominant males showed chase, chase attack, jump attack, and biting behaviors, while the subordinates displayed flight and freezing. The vast majority of bites, as well as wounds and bruises, were on the subordinates’ backs. These patterns are very similar to those previously found in rats and mice and suggest that the organization of fighting, with targets of biting (or other painful) attack serving as an important determinant of both attacker (dominant) and defender (subordinate) behavior, may show considerable generality across nonrodent as well as rodent species. Although relatively few wounds were found after 28 days of repeated and daily encounters, the subordinate tree shrews show a variety of behavioral, neuroendocrine, and central nervous changes, indicating that they are stressed by these encounters per se. Aggr. Behav. 27:139–148, 2001. © 2001 Wiley‐Liss, Inc.     

Social dominance and individual aggressiveness

Male rats exhibiting high, moderate, or low levels of offensive aggressive behavior in interactions with intruders in their home cage were grouped in mixed-sex colonies with 1 male of each aggression-level group in each colony. Agonistic interactions measured 1, 2, 3, 7, 14, 21, and 22 days after colony formation indicated that highly aggressive males on pretests continued to be more aggressive, becoming the dominant colony male in five of seven colonies and attacking intruders more often than less aggressive males. In the two remaining colonies the moderately aggressive male became dominant. This relationship, which was consistent over a number of indices, including offensive and defensive behaviors, and wound counts and wound sites, was seen even when a substantial weight differential favored the less aggressive animal. Dominance relationships were rapidly established and within-group fighting declined significantly over the 21-day test period. Pretest offensive levels also influenced the behavior of subordinates, with high or moderately aggressive subordinates showing more defense in interactions with dominants and receiving more wounds than did low-aggression subordinates. Dominant males also showed more defense in interacting with those subordinates which had been more aggressive during pretests. This pattern of results suggests that aggression level of the subordinate as well as the dominant may be an important factor determining the intensity of agonistic interactions in male rats.     

Dominance and age-related changes in the play fighting of intact and post-weaning castrated male rats (Rattus norvegicus)

The play fighting behaviour of male rats (Rattus norvegicus) castrated at weaning was compared to that of intact controls during the juvenile and post-pubertal phases of development. Following puberty, both the castrated and intact animals exhibited an age-related change in their play fighting; the frequency of initiating play fighting decreased and juvenile patterns of playful defense were replaced by more adult-like patterns. As these changes occurred even in the absence of the pubertal surge of gonadal hormones, they were more likely to result from the organizational effects of gonadal hormones in the perinatal period than the activational effects of these hormones at puberty. Although the castrated animals exhibited the age-related changes in behaviour, they did not exhibit the asymmetries in play associated with dominance relationships. As demonstrated in previous studies, in pairs of intact rats, the animal that attacks the most and uses more juvenile defenses during play fighting and weighs the least is typically the subordinate. In the castrates, asymmetries in weight and playful defense are not related to play frequency, indicating the absence of a dominance relationship. Although the characteristic changes in male play fighting at puberty are independent of the activational effects of gonadal hormones, dominance relationships and their associated changes in play fighting are dependent on these hormones. Therefore, in the perinatal period gonadal hormones most likely organize the age-related changes in play behaviour, whereas post-pubertally gonadal hormones activate dominance relationships and thus, indirectly modify play fighting by affecting dominance-associated assymetries in behaviour. © 1996 Wiley-Liss, Inc.     

Role reversal changes during the ontogeny of play fighting in male rats: Attack vs. defense

From weaning until sexual maturity, the rates at which young male rats hold each other supine during play fighting appear to become progressively asymmetrical. These changes have been previously thought to reflect an initial lack of dominance and a later development of dominance-subordinance relationships. In this paper it is shown that pairs of male rats exhibit asymmetries in playful attack and playful defense throughout development. The changes, resulting in greater asymmetry of pinning rates, are shown to result from age-dependent changes in defensive tactics; the relationship, therefore, remains constant while the form of the behavior changes. Furthermore, it is not the animals showing the highest rates of playful attack who become dominant in older ages.     

Females in charge: Effects of sex of subordinate and romantic attachment status upon self-ratings of dominance

Female, male, and mixed-sex dyads in which one member was assigned the leader role interacted and rated their own dominance throughout the interaction. The effects of gender and romantic attachment status (whether one has an exclusive dating partner or is “unattached” and free to go out with someone new) upon these self-ratings of dominance within the interaction were examined. The results showed that both leaders and subordinates perceived female leaders to be less dominant than male leaders. In addition, members of mixed-sex pairs rated themselves as less dominant than did those in same-sex pairs. Female leaders paired with males rated themselves least dominant and unattached female leaders interacting with males rated themselves least dominant of all. Female subordinates rated themselves as less dominant when with male leaders than when with female leaders, while the effect of the gender of the leader was insignificant for male subordinates. The results are discussed as evidence of a role conflict created by the contradictory roles of “dominant” leader and “subordinate” female, roles described by the sex role stereotypes prevalent in our culture.     

Social structure features in three inbred strains of mice, C57Bl/6J, Balb/cj, and NIH: a comparative study

Some features of social living were investigated in three inbred strains of mice: C57Bl/6j, Balb/cj, and NIH. Five social behaviors--Aggression, Submission, Allogrooming, Social Rest and Rest Alone, as well as Wheel Running Activity--were recorded 2 h daily for a week, in 26 groups of seven male mice living in an enriched environment. Dominants and subordinates among strains shared some behavioral characteristics, but strain contributions upon social structure were also detected. Similarities among strains included the appearance of a single dominant mouse per group rated as the most aggressive, besides from being the animal presented with most Submissions. Allogrooming was more frequent among subordinates than among dominants and subordinates. Differences among strains were that agonistic behavior among subordinates was higher in the Balb/cj and NIH groups than in the C57Bl/6j mice. NIH mice were recorded in Rest Alone more often than mice from the other strains. C57Bl/6j dominants showed higher rates of Wheel Running Activity than subordinates.     

Play fighting and the development of agonistic behavior in male and female rats

Two longitudinal studies were conducted to quantify the social behaviors exhibited by both male and female Long-Evans rats from the immediate postweaning period until young adulthood. In Experiment 1, male sibling pairs engaged in a high level of play fighting during the early juvenile period but such activity declined to a level significantly lower than that of female and mixed-sex pairs after 54 days of age. In Experiment 2, social exchanges during maturation were examined during the presence and absence of the piloerection response in an effort to distinguish play fighting from agonistic interactions. In male pairs, piloerection was rarely seen before 55–75 days of age but thereafter occurred with increasing frequency especially among dominant males. Furthermore, subordinate males retreated from their dominant partners and remained in an escape chamber for a significant amount of time only during encounters involving the exhibition of piloerection. This finding suggests that piloerection can be useful in identifying play and aggressive interactions. In female and heterosexual pairs, piloerection was observed infrequently during social encounters occurring throughout maturation. In addition, when given the opportunity to escape, females were less likely to retreat from play activity if their partner was another female than a male.     

Aggressiveness and memory: subordinate crabs present higher memory ability than dominants after an agonistic experience

A relationship between aggressiveness and memory has been proposed in several studies with different animal species. Here, we study this possibility in the crab Chasmagnathus granulatus, using the context-signal memory model (CSM) that involves an association between the learning context and a visual danger stimulus. Each experiment consisted of an agonistic phase and a memory one. During the former, matched pairs of male crabs were staged in two 10-min encounters and the dominant or subordinate condition of each member of the dyad was determined. During the memory phase, crabs were trained to acquire CSM and tested 24 h later. Results showed that the agonistic encounter, staged 48 h before the acquisition of CSM, can modulate memory according to the dominance condition of the fighter; in such a way that memory retention of subordinates results higher than that of dominants. By contrast, when the memory phase preceded the agonist one, forthcoming dominants and subordinates did not differ in their memory ability. The memory modulation would not be linked to a dominance status but to a persistent dominance relationship fully reconstructed in each encounter between the same opponents. Therefore, the crab's CSM would not depend directly on predetermined intrinsic properties, but on the outcome of the fight, which would be determined in turn by the relative aggressiveness of the fighters. The finding that the agonistic episode modulates memory opens the possibility of using this episodic interference to probe the function of diverse phases of CSM.     

The introduction of a group of alien mice (Mus musculus L.) into hierarchically organized groups of five male mice

Groups of 3 male or female Mus musculus were introduced into hierarchically organized colonies of 5 male mice for a period of 8 days. The colonies were of 2 kinds; high-level aggressive (HLA), which had been set up 3 days prior to the introduction of the aliens, and low-level aggressive (LLA), which had been established for 21 days before strangers were introduced. Both males and anestrous females were attacked, but males were attacked 5 times more frequently than females. In HLA colonies most of the attacks on the aliens were by the dominant; in LLA colonies there was no difference in the behavior of dominant and subordinate mice toward strangers. Both aggression toward aliens and intracolony aggression declined over the 8 day experimental period. In HLA colonies the dominant mouse mounted females 5 times more frequently than did either his subordinates or male mice in the LLA colonies. In HLA colonies aliens huddled with subordinates to form a single colony with 1 dominant and 7 subordinates. In LLA colonies alien males remained as a discrete group spatially separated in the cage. Female aliens were incorporated into the main colony. In all cases dominant male mice made more attacks on aliens as compared with familiar mice.     

Male aggression in the presence of an estrous female.

The relationship between male rat aggressiveness and the presence of a female was studied. Twenty 100-day-old dominant male albino rats were individually exposed to an inaccessible female rat in a specially constructed apparatus. The males were then given free-forced trials to an empty compartment or a compartment housing a submissive male. The results indicate that following exposure to an estrous female, dominant males increased their percentage of choices to the submissive male. Moreover, the incidences of aggressions by the dominant animal toward the submissive animal increased sharply after prior exposure to the estrous female. These data suggest that one antecedent source of aggression among male rodents may be the presence of a sexually receptive female.     

Targets and tactics of agonistic and precopulatory behavior in montane and prairie voles: Their relationship to juvenile play-fighting

Play-fighting by juvenile montane and prairie voles involves attack and defense of the head, neck and shoulders. Since during play animals typically borrow behavior patterns from other functional contexts, two adult behavioral contexts were compared to juvenile play-fighting. These were serious fighting and sexual encounters. During serious fighting in a resident-intruder paradigm, most bites are directed at the rump and lower flanks. During sexual encounters, especially in precopulatory behavior, the head, neck and shoulders are gently contacted. Therefore, play-fighting by juveniles would appear to involve attack and defense of areas of the body contacted in adult precopulatory behavior, not adult fighting. Furthermore, the species-specific differences in juvenile play-fighting were also found to be matched by species-specific differences in precopulatory behavior. In both playful and precopulatory encounters, montane voles contacted the head and used upright defensive behaviors more often than prairie voles. In contrast, prairie voles made mutual contact more often and were more likely to rotate to supine in defense of contact to the nape and head. These findings support our hypothesis that juvenile play-fighting in muroid rodents involves the precocial expression of precopulatory, not agonistic behavior.     

Is Transformational Leadership Always Perceived as Effective? Male Subordinates' Devaluation of Female Transformational Leaders1

This study investigated the impact of the gender composition of the leader–subordinate dyad on the relationship between leaders' transformational leadership behavior and their subordinates' ratings of the leaders' effectiveness. There were 109 dyads of leaders (58 male, 51 female) paired with a subordinate who was either the same or a different gender from themselves. The relationship between a leader's self‐report on transformational leadership and their subordinates' evaluation of their performance was significantly less positive for female leaders with male subordinates than for female leaders with female subordinates. The male and female subordinates of male leaders rated their performance as equally effective, regardless of their levels of transformational leadership.     

Masculinity,status, and subordination: Why working for a gender stereotype violator causes men to lose status

Occupying gender stereotype-incongruent roles can lead individuals to lose status and earn a lower salary. The present research examined whether merely working for a supervisor in a gender-atypical occupational role leads a subordinate to lose status. Two studies found that male subordinates of gender deviants (i.e., a female supervisor in a masculine domain or a male supervisor in a feminine domain) were accorded lower status and were paid less than male subordinates of supervisors in gender-congruent roles (i.e., a female supervisor in a feminine domain or a male supervisor in a masculine domain). However, the status of female subordinates was unaffected by working for a gender atypical supervisor. Moreover, the status loss for male subordinates was mediated by a perceived lack of masculinity. Thus, establishing the male subordinate's masculine credentials eliminated the bias.