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.     

Play-fighting differs from serious fighting in both target of attack and tactics of fighting in the laboratory rat Rattus norvegicus

Play-fighting is often difficult to differentiate from inhibited or immature serious fighting because both may utilize many of the same behavior patterns. In the rat the two behaviors involve different targets of attack. During play-fighting, snout or oral contact is directed at the opponent's nape of the neck, whereas during serious fighting, male residents mostly direct their bites at the intruder's rump. Although similar to those used in serious fighting, the behavior patterns used during play-fighting are modified to achieve the different targets of attack. Even though the tactics of attack and defense appear more adult-like with increasing age, the playful targets persist well into adulthood.     

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.     

The organization of play fighting in the grasshopper mouse (Onychomys leucogaster): Mixing predatory and sociosexual targets and tactics

The body targets contacted, the type of contact made, and the patterns of defense and counterattack elicited by those attacks are examined in the play fighting of captive male and female pairs of grasshopper mice. The nape was the most frequently contacted body target, irrespective of the type of contact made, be it nosing, allogrooming, biting, or striking with a forepaw. The types of defense varied with both body area contacted and type of attack performed. Based on the topography and pattern of contact, it was concluded that grasshopper mice, as is the case for many other muroid rodents, primarily attack and defend targets otherwise contacted during precopulatory encounters. However, grasshopper mice, which are obligate carnivores, also attack and defend predatory targets, although less frequently than sociosexual targets. Surprisingly, predatory attacks were more likely to be counterattacked with predatory attacks, whereas sociosexual attacks were more likely to be counterattacked with sociosexual attacks. Conspecific aggression involves bites directed at the face, lower flanks, and dorsum. Neither the biting of these areas nor the tactics of attack and defense usually associated with such bites were observed during the juvenile interactions. There were no sex differences in either frequency or patterns of attack and defense in play fighting. The data presented for grasshopper mice shed light on the issue of mixing behavior patterns from multiple functional systems during play. Aggr. Behav. 26:319–334, 2000. © 2000 Wiley‐Liss, Inc.     

Targets and tactics: The analysis of moment-to-moment decision making in animal combat

Even though injury and death are more common consequences of fighting among animals than once believed, they are still relatively infrequent. Modern evolutionary models of animal combat have emphasized that given the threat of retaliation, animals only escalate to more injurious fighting if the benefits outweigh the costs, and then only if threat and bluff fail to achieve the goal. Such models stress the role of communication as to whether animals decide to escalate or not. An alternative view is that failure to produce injury or death arises from the neutralization of one animal's attack by another's defense. That is, attack and defense end in a stalemate that may be misinterpreted by outside observers as an absence of injury producing behavior. As attack typically involves the biting or striking of specific body targets, movements and postures occurring during combat need to be analyzed with respect to their role in gaining or averting such contact. For example, in the combat of muroid rodents the attacker targets the lower dorsum and flanks (low threshold) or face (high threshold), whereas a defender may defensively launch counterstrikes against the attacker's face. Two combat tactics (supine defense and lateral attack) typically present in the fighting of muroid rodents are analyzed in detail to illustrate how targets constrain the movements of combatants. Such a functional analysis of combat assumes that the movements and postures performed are related to their role in the attack and defense of targets. Deviations from such a strict functional interpretation reveal some of the other factors that may constrain the combatants' behavior. For example, body morphology and the aggressiveness of the opponent are shown to be important in deciding the type of combat tactic to use and how it is performed. Finally, movements and postures that are neutral or even counterproductive for attack and defense may be revealed as communicatory. This approach provides a means of analyzing behavior during the "heat of combat" that is typically not dealt with in traditional evolutionary models. Aggr. Behav. 23:107–129, 1997.© 1997 Wiley-Liss, Inc.     

Identification of the possible origin of the body target that differentiates play fighting from serious fighting in syrian golden hamsters (Mesocricetus auratus)

Play fighting in the Syrian Golden hamster Mesocricetus auratus can be distinguished from serious fighting by the targets attacked in each case. In play fighting, the animals attack and defend the cheeks and cheek pouches, whereas in serious fighting they attack and defend the rump and lower flanks. Since play typically involves the use of behaviors borrowed from other functional contexts, this paper investigates the origin of the cheek target during play fighting. Comparison of resident-intruder serious fighting with awake and anesthetized intruders does not reveal the cheek to be an inhibited target for serious attack. Similarly, analysis of social investigation and allog-rooming, while revealing the ears to be important targets, do not show the cheeks to be targets in these behaviors. Sniffing, licking, and nibbling of the cheek area appear to occur mainly during sexual encounters by males. This area, seemingly a sexual target, may be the one utilized during play fighting.     

Analysis of the targets and tactics of conspecific attack and predatory attack in northern grasshopper mice onychomys leucogaster

Comparisons of tactics of fighting between species are often difficult to make since the body targets attacked may differ. Thus it becomes difficult to assess whether differences in fighting tactics are due to species-specific differences in the tactics themselves or due to the different targets attacked. A solution to this problem is to analyse the tactics of a species that attacks different targets under different circumstances. In this way, differences in tactics can be more readily attributed to differences in targets. In this study, resident male northern grasshopper mice (Onychomys leucogaster) were tested against intruding male conspecifics and against laboratory mice (Mus musculus domesticus). Conspecifics were mainly bitten on the lower dorsum, whereas prey were bitten and killed by bites to the nape of the neck. Therefore, it was possible to analyze the tactics of attack by grasshopper mice when attacking different body targets. For example, in order to defend the lower dorsum and the nape, both intruding conspecifics and prey adopted an upright defensive posture. Resident grasshopper mice used the lateral attack tactic to gain access to the lower flanks but not the nape. This illustrates that the lateral attack tactic is not merely a tactic suitable for overcoming the upright defense tactic, but is used in this context only when the target attacked is on the opponent's posterior dorsum. Such withinpecies comparison enables the identification of the contextual rules which govern the use of fighting tactics. © 1992 Wiley-Liss, Inc.     

Different levels of complexity in the play-fighting by muroid rodents appear to result from different levels of intensity of attack and defense

Play-fighting in deer mice, Peromyscus maniculatus bairdii, prairie voles, Microtus ochrogaster, and montane voles, M. Montanus, was compared to that of laboratory rats, Rattus norvegicus. Play in rats appears more complex for two reasons: 1) more of the playful contacts elicit defensive behaviors, and 2) more of these defenses lead to counterattacks, and hence, role reversals between attackers and defenders. Neither high levels of defense, as shown by montane voles, nor high levels of counterattack, as shown by prairie voles, produce rat-like play-fighting. This only occurs when high rates of defense involving turning to face the attacker and counterattack are combined, as in rats. These two components are rarely combined together by deer mice, and so this species rarely exhibits rat-like play-fighting. Furthermore, playful counterattack appears to arise from playful attack, and not from an escalation of defense. These data suggest that the more complex forms of social play, such as play-fighting, have evolved, in part, via the escalation of defense in response to playful attack.     

Motivational systems of agonistic behavior in muroid rodents: A comparative review and neural model

The data on agonistic behavior of muroid rodents that have been obtained from field observations and laboratory experiments are reviewed and compared in terms of a hypothetical model of the neural organization of these behaviors. The neural model has been presented elsewhere and is used here only as a way to organize the data. The data are organized in terms of four hypothetical motivational systems: Offense, defense, submission, and patrol/marking. The various behaviors are considered as motor patterns and are compared and analyzed in terms of the proposed motivating, releasing, and directing stimuli of the motivational systems. Interactions and overlaps between the motivational systems are also considered. It is concluded that the organization of agonistic behavior may be similar across all species of muroid rodents. Generalizations are complicated by the profound effects of ontogenetic factors. Four categories of behaviors differ from species to species: Scent-marking, submissive behaviors, threat behaviors, and alarm signals. The possible phylogenetic and ontogenetic factors in these differences are considered.     

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.     

Maternal aggression toward infanticidal males of different social status in wild house mice (Mus musculus domesticus)

Maternal aggression was examined in wild female mice (Mus musculus domesticus) derived from animals trapped in Alberta, Canada. Lactating females were tested for their behavior toward intruder males during the time of postpartum estrus while housed in a two-cage apparatus containing a defensible nest area. Prior to being used as intruders, sexually naive males were screened for their behavior toward a newborn pup (83% exhibited infanticide). Only infanticidal males were then housed in pairs and allowed to establish a dominance hierarchy. Dominance status was further verified by a urine marking test. The dominant and subordinate infanticidal males were then placed into a lactating female's cage and observed for 1 hr. The test was terminated immediately when a male began to attack the pups. Lactating females attacked the males in both groups, but subordinate males received more intense attacks than dominant males. Dominant males elicited significantly more fear/defense behavior than subordinate intruders. All of the dominant males and only one submissive male attacked the pups. Females were thus successful in blocking infanticide only by infanticidal subordinate males. Since females do not persist in attacking males with high fighting ability, one function of maternal aggression could be to assess the fighting, and resource holding, potential of a future mate. © 1994 Wiley-Liss, Inc.     

Ultrasounds produced by rats accompany decreases in intraspecific fighting

Male intruder rats were placed individually into the cage of an established resident on 2 occasions separated by a 7–8 day interval. Residents readily attacked intruders and both animals lost weight during the first encounter. In contrast, no serious fighting occurred on the second encounter, and both intruders and residents maintained their body weight during the 24-hr test. Observation of the intruder's behavior during the first 30 min of each encounter indicated that defensive-submissive postures represent a response to an attack that only temporarily inhibits aggression whereas the emission of 22 kHz calls by the intruder is associated with a relatively permanent decrease in the resident animal's aggressive response.     

Response of colony mice to intruders with different fighting experience

Six male mice placed in a large, moderately complex enclosure formed a stable dominance hierarchy in which two mice defended adjacent floor areas and the remaining four mice were subordinate and did not form territories. Intruder mice with winning or losing experience in prior paired encounters, or those with no fighting experience, were introduced individually into the colony for 30 minute periods. These intruders were attacked by the dominant members of the colony, and the fighting outcomes were strongly dependent upon the fighting experience of the intruder. Intruders with losing or no fighting experience engaged in little mutual fighting with residents, were easily defeated, and terminated attacks by engaging in subordinate behaviors. Intruders with winning experience fought vigorously with residents, attacked and, in many cases, defeated residents. These results suggest that relatively little winning experience gained in earlier paired encounters may be sufficient to overcome the various fighting advantages enjoyed by a dominant territorial holding member of a colony.     

The meaning of observed violence: Real vs. fictional violence and consequent effects on aggression and emotional arousal

Sixty male subjects were either attacked or treated neutrally by a confederate, after which each saw a videotape of two men fighting. Subjects were informed that fight was either real or fictitious or were given no explanation of it. Subjects who had previously been attacked and had observed the fight under a set to perceive it as real were subsequently more punitive in their treatment of the confederate than subjects in all other conditions. The combination of prior attack and observation of real violence also sustained blood pressure (BP) at near the level produced by the attack, whereas BP of attacked subjects in the other conditions declined during the time the fight was observed. Palmar sweat measures revealed that observation of real violence was more arousing than observation of fictitious fighting. The results are discussed in terms of the effects that the reality of observed violence has on emotional arousal.     

Female behavior in populations of mice in the presence and absence of male hierarchy

Female aggression may be the regulator of population size in small mammals. Freely growing populations of house mice showed several differences in aggressive female behavior in the presence and the absence of a male hierarchy. Territoriality in females and not in males appeared to maintain social order and regulate population density. Certain females were seen patrolling and guarding the territory and chasing and fighting with both male and female intruders. These females did not fight amongst themselves, suggesting that they were not fighting for rank (as do the males) but for territory. Although these aggressive females produced young, the pups were neglected, and few were weaned. The non-aggressive females were the successful breeders. Aggression by the females only occurred when there was reproduction and increased densities. Assembled females with no males present never show this aggression. The occurrence of “male-type” behavior became most apparent when the males were removed at peak population densities. The removed males were then castrated and injected with testosterone cyprionate. Doses were increased by population cage, and therefore all males returned to each freely growing population were given the same dose. The males given oil placebo injections showed no return of a male hierarchy and the females showed high levels of aggression toward them. Males injected with testosterone cyprionate showed return of male aggression and fighting and mounting of females. But the new “dominant” females continued their patrols and chased males away from their territories and did not permit these males to mount. Male-male fighting consisted primarily of frontal attacks to the face and roll and tumble fights. Female-male aggression consisted primarily of attacks to the posterior region targeted at the base of the tail and the genitals of the male. The males were rarely seen attacking females and then only during mating. Females only attacked each other in defense of their territories.     

Estimation of the probability of fighting in fallow deer (Dama dama) during the rut

Fighting between males is a frequent component of the rutting behavior of Cervidae. Frequent conflicts are exhausting; fighting may be risky and can lead to serious injuries or even death. We focused on the process of assessment of the opponent's fighting ability and escalation of the combat, estimating the probability of fighting based on the encounter components such as groaning and parallel walk. In this study, we observed the agonistic behavior of fallow deer bucks (Dama dama) during the rut over four seasons. During this time, we recorded 205 encounters between bucks. Non-contact display, which allows contestants to assess their opponents fighting ability, occurred in 83% of the encounters. The highest predicted probability of a fight was found when both of the males vocalized and turned into the parallel walk. The chance of a clear outcome decreased when the males were fighting in comparison to when they did not fight. The initiator of the competitive encounter won 41% of the cases, while the attacked buck won 23% of the encounters. If the contestants avoided fighting, however, the initiator won 78% of encounters. Therefore, the initiator was more successful when no fight occurred compared to when the encounters escalated into fighting. In most cases where ritualized behavior occurred, one of the opponents left after vocalization or parallel walk occurred. Thus, vocalization and parallel walk increased the probability for a clear outcome. The probability of a fight was lowest in situations where the males displayed asymmetric behavior. Increased symmetry of the contestants' behavior was strongly correlated with a higher probability of a fight. Thus, these results indicate that fallow deer bucks use efficient tactic during the rut, which, in turn, minimizes the chance of injury while fighting during the breeding season.     

MISUNDERSTOOD IN THE DIASPORA: THE EXPERIENCE OF ORTHODOX SIKHS IN VANCOUVER

Heroism and martyrdom are central values in contemporary Sikh historiography and primary symbols in their collective memory. Both are replete with the stories of numerous heroes and martyrs who faced death, unflinchingly fighting in defense of the Sikh faith and its believers (constituting the Panth or community) against overwhelming odds. Conversely, it is considered shameful for Sikhs to become mere victims, succumbing to attacks against faith and community without fighting to the death. It is also insisted that true Sikhs do not themselves attack those who are weak or harmless; on the contrary true Sikhs come to the defense of such persons even if they are not of the same faith. However, during the last half century, Sikhs have many times become victims, as well as perpetrators of violence, being unable to act in a way consistent with their central values. The consequence has been an inability to integrate their modern history successfully into a narrative consistent with those values and to find appropriate ways of memorializing what has been done to many Sikhs and what many Sikhs have done to others.     

Attack and defensive behaviors in the albino mouse

Attack by dominant male colony mice on intruders included chasing and lateral attack behaviors, while the corresponding intruder behaviors were flight, boxing, and checking. Both of these are similar to the attack and defensive behaviors of colony rats and intruders. However, mice did not show a significant constraint on bites to ventral areas, and the rat defensive behavior of lying on the back, which is effective because of this constraint, was rare; the corresponding “on-top” behavior of attackers was almost absent in mice. These findings strongly support the view that intraspecific attack and defensive behaviors, and target sites for bites, are interrelated factors facilitating effective but nonlethal agonistic interactions in muroid rodents.     

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.