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.     

Influence of dominance on the development of play fighting in pairs of male Syrian golden hamsters (Mesocricetus auratus)

In the highly social rat, male juvenile and adult subordinates initiate more playful contacts with dominant pairmates than vice versa. This study examined the effect of dominance on playful contacts in the relatively asocial golden hamster. Pairs of male hamsters were reared together from weaning, and their play was filmed in the juvenile (28-36 days) and the young adult (60-70 days) stages of development. By the adult stage, it became clear that one pairmate was dominant over the other. The dominant pairmate launched all aggressive attacks (i. e., bites to the lower flanks and rump), and the subordinate pairmate performed all the submissive gesturing (e. g., tail up submissive posture). Playful contact, which in this species involves gentle nibbling of the posterior cheeks, was more frequently launched by the dominant than by the subordinate. This was not only true at the adult stage, but also at the juvenile stage, before dominance-subordination relationships were sharply polarized. Therefore, it would appear that in the relatively asocial hamster, the subordinates tend to avoid playful contact with dominants. This is markedly different to rats, where the subordinates actively seek out and engage dominants in play. This contrast further supports our hypothesis that subordinate male rats use play as a means of maintaining familiarity with dominants. © 1993 Wiley-Liss, Inc.     

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.     

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.     

Male-female differences and the influence of neonatal and adult testosterone on intraspecies aggression in rats.

Male and female albino rats were tested for intraspecies aggression without the use of shock. In the first experiment, male pairs showed more biting attacks, offensive sideways movements, and self-grooming than did female pairs; male pairs also showed more stereotyped defensive/submissive behaviors and were wounded more frequently. The second experiment examined the effects of neonatal castration and testosterone propionate (TP) administration on fighting. Males castrated at birth attacked other males less frequently than did controls when tested with TP treatment as adults. The TP given at birth to neonatally castrated males restored attacks to control levels. Females given TP as neonates did not differ from either male or female controls. Other aggressive/defensive behaviors, however, did not show this pattern. The results suggest that while the presence of testosterone during a brief postnatal period and during adulthood is necessary for attack behavior to occur, other related behaviors may not be affected in a similar manner.     

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.     

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.     

Steroidal modulation of scent investigation and marking behaviors in male and female ferrets (Mustela putorius furo)

Gonadectomized male and female ferrets (Mustela putorius furo) given either testosterone propionate (TP) or oil vehicle preferred to investigate the side of a test cage previously soiled by a breeding male or female as opposed to a clean side. Male and female ferrets receiving TP showed more urogenital wiping than oil-treated animals in either side of the test cage. In a 2nd experiment, ferrets treated sequentially with TP, oil, and estradiol benzoate (EB) were given simultaneous access to sides of a test cage previously soiled by either a breeding female or male. Either EB or TP treatment of females and TP treatment of males facilitated the investigation of odors of opposite-sex ferrets. Females given TP and males given either TP or EB showed increased urogenital wiping in both sides of the test cage. Sex steroids modulate scent investigation and marking in adult ferrets in a sexually differentiated fashion.     

Role of estrogens in androgen-induced spontaneous activity in male rats.

Three experiments tested the hypothesis that testosterone may be aromatized to an estrogen to stimulate running-wheel activity in rats. Aromatizable (testosterone propionate: TP) and nonaromatizable (dihydrotestosterone propionate; DHTP) androgens were compared with estradiol benzoate (EB) for the ability to induce running in castrated male rats. The DHTP had no effect on running. The TP increased running, but EB was more than 100 times as effective. A relatively small dose of a specific estrogen antagonist, MER-25, was shown to attenuate the effects of both EB and TP on male running. The MER-25 did not affect the running of castrated, oil-treated male rats and did not inhibit the running induced by food deprivation.     

Targets of attack and defense in play-fighting of the Djungarian hamster Phodopus campbelli: Links to fighting and sex

Analysis of the body targets attacked and defended during play-fighting by juvenile Djungarian hamsters Phodopus campbelli revealed that about 70% of all attacks were directed at the mouth. If successfully contacted, the mouth was briefly licked and nuzzled. The remaining playful attacks were gentle bites directed at the rump, and to a lesser extent, the top of the head. During serious fighting the top of the head and the rump are targets of attack, whereas the mouth is not. Licking and nuzzling the mouth was found to be a behavior performed by adult males at the beginning of sexual encounters. Therefore, play-fighting in juvenile hamsters cannot be thought of merely as a form of “mock fighting” since the principal target is seemingly sexual, not agonistic. The data also show that of the sexual body targets contacted, adult females are more likely to defend the mouth. In this way it is suggested that targets attacked and defended during juvenile play-fighting are derived from adult contexts in which such targets are defended. This hypothesis accounts for the prevalence of agonistic targets in the play-fighting of many species, and may provide a rationale for classifying those amicable targets that are competed for during play-fighting.     

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.     

Modifications induced by neonatal steroids in reproductive organs and behavior of male rats.

Male rats injected on Day 3 neonatally with .01, .1, 1, 10, 100, or 1,000 micrograms of estradiol benzoate (EB), 10,000 microgram of testosterone propionate (TP), or sesame oil were subsequently examined for testicular, penile, and accessory organ development. Sexual behavior was evaluated during therapy with fluoxymesterone (FM) and then with TP. Estradiol benzoate in dosages greater than 1.0 micrograms delayed testicular descent, reduced the size and hormone responsiveness of reproductive organs, and decreased sexual behavior in a dose-dependent manner. The 10,000-microgram dosage of neonatal TP delayed testicular descent and reduced sexual behavior to levels near those of the 10--100 micrograms EB groups, but it produced no significant penile or accessory organ changes. Neither reduced peripheral organ development nor inhibited neonatal testicular secretions fully explain reductions in male behavior following large dosages of neonatal TP. Neonatal androgen may reduce the responsiveness of central nervous system neurons governing male sexual behavior after being converted to estrogen or by directly altering steroid receptor systems.     

Organisational effects of neonatal and pubertal testosterone on sexually differentiated behaviours in the open-field and head-dip apparatus

On days 1 and 4 after birth rats were injected with 100 μg of testosterone propionate (TP) or vehicle, and at 35 days of age they were injected intramuscularly with 400 μg of testosterone oenanthate (TO), a long acting androgen, or the vehicle. There were four groups (oil-oil, TP-oil, oil-TO, TP-TO), each group subdivided by sex. Females treated with testosterone neonatally or at puberty were masculinised or defeminised on adult open-field behaviours, being less active and rearing less than oil-oil females; the oil-TO group also defaecated significantly more than controls. The TP-TO female group was indistinguishable from the oil-TO group. In a second experiment, sex differences were found in head-dipping behaviour as well as in activity and rearing. Females treated with TP or TO reared less and defaecated more than controls, and TP also decreased activity, but neither hormone treatment affected head-dipping behaviour. There is thus a peripubertal as well as a neonatal period when testosterone can act organisationally to masculinise or defeminise female rats. Potentiation between effects of neonatal and pubertal androgens was found on female body weights. TO alone had no effect, but TP-TO females were significantly heavier than controls at 90 days of age and by 130 days of age the TP-oil group was also heavier than controls.     

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.     

Gonadal hormones and intrasexual aggressive behavior in female bank voles (Clethrionomys glareolus)

The effects of gonadal hormones on aggressive behavior in the female bank vole was investigated in 10 min home cage tests. Ovariectomized (ovx) or intact females injected with oil, with progesterone (P), with a mixture of progesterone and estrogen (P+E), or with testosterone (T) alone were confronted in a resident-intruder test with unfamiliar, nonoperated females as intruders. Intact females showed aggressive behavior more frequently than ovx females. Ovx females injected with P, with P+E, or with T made significantly more attacks, and these attacks lasted longer than those observed for oil-treated voles. The results indicate that P, the typical female hormone, is responsible for aggressive behavior in female bank voles; however, only T increased the duration of interfemale aggression. Aggr. Behav. 24:63–70, 1998. © 1998 Wiley-Liss, Inc.     

The use of the bared-teeth display during play fighting in Tonkean macaques (Macaca tonkeana): sometimes it is all about oneself

Play signals are viewed as important means by which animals inform each other that bites, strikes, and throws that occur during play fighting are indeed playful rather than serious. One such signal is the open mouth play face that is common in primates and many other mammals. Unfortunately, as most play fighting involves biting, it can be ambiguous as to whether any instance of opening the mouth is performed to communicate playful intent or is simply a preparation for biting. In this study, open mouths co-occurring with the bared-teeth display (teeth-baring) in Tonkean macaques were used to assess the context in which facial gestures only relevant for signaling (i.e., teeth-baring is not necessary for biting) are used during play. Two predictions arising from the hypothesis that play signals are used to facilitate playful contact were tested: that the open mouth with teeth-baring should (1) be most frequent preceding contact, and (2) that it should be performed most often when bites are directed at orientations that is visible to the recipient. The data only partially support these predictions. The open mouth with teeth-baring is also frequently used when a monkey withdraws from playful contact. Moreover, it is associated with bites to body targets, such as the rump, that offer little prospect for detection by the recipient; this supports the possibility that play signals may sometimes be emitted not to communicate with the partner but with the performer itself. Thus, play signals serve multiple functions during play fighting.     

Targets,tactics, and the open mouth face during play fighting in three species of primates

The play fighting of many mammals involves the nonserious use of behavior patterns derived from serious fighting. A major question of theoretical importance has been that of how, given this overlap in patterns of behavior, the animals can distinguish between playful and nonplayful intent. One proposed solution is that animals use play signals to inform each other about the playful intent of their actions. The most widely reported play signal amongst primates is the open mouth play face. The manner in which this so-called signal functions is based on correlational evidence, with most reports simply noting its presence or absence in a given species. This study involved a detailed video-based analysis of the occurrence of open mouths during the play fighting of three species of primates. One captive troop each of ring-tailed lemurs, black-handed spider monkeys, and patas monkeys was used. By examining all open mouths in the context of the species-typical style of play fighting, several conclusions were empirically verified. 1) Most open mouths occur as a functionally necessary precursor for biting. 2) Some open mouths occur as a defensive threat which deters further contact. 3) The residual open mouths which may function as contact promoting play signals, constituted about 20–25% of all open mouths by the lemurs and patas monkeys, but less than 5% for spider monkeys. These species differences appeared to arise from two causes. Firstly, the spider monkeys used another signal, the head shake, in situations where lemurs and patas monkeys used open mouths. Secondly, the style of play fighting greatly influenced the frequency and duration of open mouths. This was most marked in the face-to-face combat style of patas monkeys. These findings show that comparative studies of the occurrence and function of play signals need to take into account species-typical styles of playful combat. Aggr. Behav. 23:41–57, 1997. © 1997 Wiley-Liss, Inc.     

Threats and promises: Reputational consequences and transfer of credibility

This study was designed to investigate the reputational consequences of making threats and promises and how they affect subsequent credibility. Eighty-eight high school girls were given predetermined information about the past behavior of their pairmates prior to receiving influence attempts from them. Each first learned that her pairmate had (1) threatened or promised in the past and then learned whether she had (2) fulfilled or failed to fulfill her threat or promise. The subject herself then received a threat or promise from her pairmate. As anticipated, threateners were viewed less favorably than promisers. These impressions were maintained when it was learned that commitments had been fulfilled, but altered decidedly when it was learned that they had not. Credibility or lack of it was found to generalize from threats to promises and from promises to threats only under some conditions. It was concluded that the fit between existing impressions and proclaimed future activity, not a person's record for past reliability, is often the critical determinant of whether or not she is judged to be credible.     

Children's hostile attribution bias is reduced after watching realistic playful fighting, and the effect is mediated by prosocial thoughts

Hostile attribution bias (HAB) has been found to characterize aggressive children. Watching prosocial media has been shown to have positive effects on children, and the general learning model has been used to account for these observations. This study tested the hypotheses derived from this theory that exposure to playful fighting would lead to a reduction in HAB, both immediately and after a 1-day delay, and that this effect would be mediated by positive thoughts. Four studies exposed child participants (N=242) to playful fighting versus neutral behavior primes and then tested their HAB. In two studies, thoughts about playful fighting and about children were assessed and tested as mediators. The main hypotheses were supported. The positive effect of watching playful fighting on HAB was evident immediately but not after 1 day. This effect was mediated by positive thoughts. In line with the general learning model, watching playful fighting reduced HAB in children, and positive thoughts contribute to this effect. This extends the realm of the general learning model and suggests interventions to help children avoid aggression.     

Effects of neonatal handling on play behavior and fear towards a predator odor in juvenile rats (Rattus norvegicus)

The effects of brief daily separation, also known as "handling," during the first 2 weeks of life on play behavior and fearfulness toward a predatory odor were assessed in juvenile rats. Handled rats were more playful than nonhandled control rats, and while handling had no effect on the direct response of these rats toward a predatory odor, handled rats did not exhibit a conditioned suppression of play when tested later in the same context where they had been exposed to the predatory odor. Handled rats were still wary of the environment in that they continued to show a heightened level of risk assessment behavior. These data suggest that early postnatal experiences may play a significant role in determining how an animal deals with predatory threats later in life.