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.     

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.     

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.     

Sexual and aggressive play fighting of sibling Richardson’s ground squirrels

Play fighting in many species of squirrels can involve sexual play and aggressive play, both of which can lead to wrestling which appears superficially similar. Such convergence can make scoring of the relative frequencies of these two types of play difficult and can lead to the mistaken conclusion that they grade into one another. In this study, both staged laboratory encounters between sibling pairs and spontaneous encounters between siblings in free‐living litters of Richardson’s ground squirrels (Spermophilus richardsonii) were videotaped. Frame‐by‐frame analyses using the Eshkol‐Wachman Movement Notation were employed to record the correlated movements of attack and defense by the partners and to reveal the body areas targeted during each play bout. Whereas sexual play was organized around access to the rump, aggressive play was organized around the shoulders. Although in most cases the defender’s tactics blocked access to the respective target, when contact did occur, it involved mounting in sexual play and nosing or biting in aggressive play. Eighty‐six percent of play fights could be unambiguously categorized as either sexual or aggressive play. Of these, the majority (?80%) involved sexual play. The sex of the participants did not affect the frequency of aggressive play, but in sexual play, males initiated more attacks than females. Once initiated, each form of play fighting remained distinct—if a bout began as sexual play, it would end as sexual play. Furthermore, a counterattack following sexual play was significantly more likely to be sexual than aggressive, and vice versa for counterattacks following aggressive play. Therefore, all the evidence suggested that the two forms of play fighting were not intermixed in Richardson’s ground squirrels. Aggr. Behav. 27:323–337, 2001. © 2001 Wiley‐Liss, Inc.     

Reflexive fighting in response to aversive stimulation

Reflexive fighting was elicited between paired rats as a reflex reaction to electric shock prior to any specific conditioning. Such fighting was fairly stereotyped and easily differentiated from the rats' usual behavior. The strength of this reflex was not attributable to any apparent operant reinforcement. Elicitation of fighting was a direct function of the enclosed floor area and a nonmonotonic function of the shock intensity.

Failure to scramble the polarity of the electrified grid produced inconsistent fighting. Under optimal conditions fighting was consistently elicited by shock regardless of the rat's sex, strain, previous familiarity with each other, or the number present during shock. Repeated shock presentations did not produce an appreciable decrease in fighting until signs of physical debility appeared. Although shock did not cause a rat to attack inanimate objects, it did produce attack movements toward other small animals. Failure of guinea pigs to defend themselves revealed that the elicitation of fighting from the rat does not require reciprocal attack. Paired hamsters showed fighting reactions similar to those of the rats, whereas guinea pigs failed to fight. Electrode shock and a heated floor elicited fighting between the rats, but intense noise and a cooled floor did not.

     

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.     

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.     

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.     

Agonistic versus amicable targets of attack and defense: Consequences for the origin,function, and descriptive classification of play-fighting

Play-fighting appears to involve the behavior patterns of attack and defense otherwise seen in serious fighting. The degree of similarity, however, depends on the body targets attacked and defended during these forms of fighting. For many taxa, including diverse mammalian families and some birds, the same targets are attacked and defended during both play-fighting and serious fighting. However, for several species of muroid rodents, the targets of play-fighting are not the same as those of serious fighting. In these cases, the tactics of attack and defense are also different. It is argued that for these muroid species the playful targets have arisen from amicable behavior (e.g., social investigation, greeting, allogrooming) rather than, as appears to be the case in so many other taxa, from agonistic behavior. These data strongly suggest that “play-fighting” has evolved from different precursors in different taxa and thus has multiple origins. Furthermore, these data have an important bearing on the universal applicability of many of the suggested functions of play-fighting and also on how such behavior is to be described and classified.     

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.     

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.     

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.     

Play fighting and real fighting: Using video playback methodology with young children

This study develops a video playback methodology: children aged five to eight years viewed taped play fighting and real fighting bouts in which they were both participants and nonparticipants. Views of participants were also compared for immediate and delayed viewing. The methodology examines the criteria used to distinguish play fighting and real fighting; views concerning the characteristics of such episodes; and the motivations involved. The methodology was found to be feasible at this age; and useful in terms of differing views of participants and nonparticipants. There was evidence for increased insight from participants, who used more criteria to make their judgments, and more informative criteria such as knowledge of the rules of a game being played. Participant knowledge was most evident at the immediate viewings, but was partially retained one week later. Participants (more than nonparticipants) mostly described play fighting as friendly, and not involving hurt or showing off. The methodology could usefully be applied to examine further developmental changes in older children. Aggr. Behav. 30:164–173, 2004. © 2004 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.     

Influence of timing of post-weaning isolation on play fighting and serious aggression in the male golden hamster (Mesocricetus auratus)

Effects of timing of social isolation on play fighting and serious fighting were studied at different ages in male golden hamsters. Litters were isolated at 21, 35, and 65 days of age, and tested in a resident-intruder paradigm. Behaviors were compared within grous and with a fourth group of socially reared conspecifics. The earlier the pups were isolated, the more they engaged in play activities. Later, in adulthood, the aggression level of the same animals was retested using the same paradigm. The three isolated groups showed a high level of aggression, with significant differences among them. When compared with socially reared subjects, a reliable difference in the level of aggression was also found. These results support the view that early social experience is important, suggesting that isolation during early critical periods of socialization has a significant impact on play fighting, whereas short periods of isolation may be enough to trigger adult agonistic behavior. © 1994 Wiley-Liss, Inc.     

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.     

Competitive fighting in the rat.

Competitive fighting was obtained in pairs of like-sexed laboratory rats by placing a single piece of food into the food hopper following 48 hr. of food deprivation. The fighting was characterized by offensive sideways posture, full aggressive posture, and bite and kick attack. Tests were conducted at 110-120 days of age on pairs of animals that had been housed together since weaning. Fighting was more frequent in pairs consisting of nonlittermates than in pairs of littermates, and it was equally frequent in male and female pairings. Probability of fighting was enhanced by prior experience with food deprivation, and attack was most often initiated by the heavier animal of the pair.