The effect of the type of opponent in tests of murine aggression

The interactions between isolated resident mice and anosmic, juvenile, submissive and dominant intruders were studied; the responses of intruder mice to being attacked and the distribution of bites on these subjects were investigated. Anosmic, juvenile and (trained) submissive intruders did not attack residents; they did, however, show differences both in thir responses to being attacked and their ability to avoid bites to particular regions of their body surface. These differences are likely to make it difficult to extrapolate between tests that employ different types of intruder. Trained dominant intruders showed substantial attack on resident mice and a very different pattern of attack was generated by this type of encounter. The study confirms that, while “standard opponents” are useful means of assessing aggressive motivation, the pattern of fighting they produce differs depending on their background.     

Aggressive behaviour of female prairie deer mice in laboratory populations

Prairie deer mice (Peromyscus maniculatus bairdii), living in asymptotic laboratory populations established two years earlier, were observed for agonistic responses to conspecific intruders. In the first experiment, intruders of six age-sex classes were placed into 10 of the populations for 10 min. The sex of the intruder did not influence the behaviour of the residents, but juveniles elicited more aggression than did adults. A second experiment revealed that female residents were responsible for almost all of the attacks upon juveniles. Experiment 3, in which the responses of pairs of deer mice to juvenile intruders were recorded, demonstrated that the aggressiveness of a female was enhanced by the presence of a male. In the final experiment, females were observed to be highly aggressive during the first few days after giving birth. The aggressive behaviour of the female deer mouse may have greater significance for population dynamics than that of the male.     

Aggression persists after ovariectomy in female rats

Aggression occurs not only in males but also in females, however, under different sex-specific stimulus and endocrine conditions. After being housed with males, female rats exhibit frequent and intense aggressive behavior toward unfamiliar rats. However, the female residents primarily attack female intruder rats, while the male residents attack males and not females. Altering the hormonal condition of the intruders can modify the behavior that they provoke from the residents. Castration of the male intruders reduces aggression from male residents, but ovariectomy of the female intruders does not alter the behavior of the female residents. Treatment of the gonadectomized intruders with gonadal steroids significantly alters the response of the male residents. Resident-intruder aggressive behavior depends on the presence of the testes in the male residents but not on the ovaries or on lactation in the female residents. Even 7 weeks after ovariectomy the female residents continue to show aggressive behavior toward female intruders. In the same time period the castrated male residents show a marked decrease in aggressive and sexual behavior.     

Age and brood defence in male CRL:NMRI BR laboratory mice, Mus musculus domesticus

According to Lack's principle, older mice should invest more in their offspring than younger ones. I tested this principle with pairs of CRL:NMRI BR mice protecting their young against an intruder, an unfamiliar same-stock male or female. The intruder was presented within a small wire-mesh cage in an extension, a cage the same size as the pair's home cage. The dam, her mate and the young in the nest were present in all tests. Although both parents had full access to the intruder, only the male interacted with it. Young breeding pairs and young intruders were 6 months old, the old pairs and old intruders 12 months. Each breeding pair was tested with both young and old male intruders. Older males defended their pups more vigorously than younger ones. The old males showed a higher preference to stay near (to interact with) the opponent; they also showed more tail rattling and biting attempts than the young ones. This indicates that older males take greater risks in defending their offspring than younger males. Older intruders also elicited a stronger brood defence in older males and a weaker response in younger ones.     

Effects of familiarity on the behaviour towards intruders in captive common marmosets (Callithrix jacchus)

A study on the behaviour of a group of common marmosets (Callithrix jacchus) towards unrelated intruding conspecifics (juvenile male twins) was performed. The group members' proximity to the intruders during two consecutive study periods was analyzed. During the first period in which the experimental conditions as well as the intruders were unfamiliar, all but one group member were equally likely to stay near the intruders and usually more than three individuals were present simultaneously. During the second period the group members showed different scores of proximity towards the intruders. One individual, an adult male, was in close proximity nearly the whole time. Additionally, the number of individuals simultaneously staying near the intruders was clearly reduced. It is argued that (1) the change in behaviour of the group members was affected by the increasing familiarity and (2) intruder experiments with unfamiliar conspecifics are only of partial value for clarification of the process of immigration.     

The structure and usage of female and male mouse ultrasonic vocalizations reveal only minor differences

Ultrasonic vocalizations (USV) of mice are increasingly recognized as informative dependent variables in studies using mouse models of human diseases. While pup vocalizations primarily serve to re-establish contact with the mother, adult male "songs" were considered to be courtship signals. Alternatively, mouse USVs may generally function as territorial signals. To distinguish between these two hypotheses, we compared the structure and usage of adult male and female USVs in staged resident-intruder encounters. If calls function primarily as courtship signals, males should respond stronger than females, specifically when presented with a female intruder. Refuting this hypothesis, we found that in response to female intruders, females called more than males (228±32 calls/min vs. 71±15 calls/min), and males called more to female than to male intruders (14±7.5 calls/min). There were no significant differences in the acoustic characteristics of the calls given by females and males. To control for the influence of the intruder's behavior on calling, we repeated the experiments using anaesthetized intruders. Again, females produced more calls to female than male intruders (173±17 calls/min vs. 71±15 calls/min), while males called more in response to female than male intruders (39±17 calls/min), and there were no acoustic differences in female and male calls. The vocal activity did not differ significantly with regard to intruder state (awake or anaesthetized), while the acoustic structure exhibited significant differences. Taken together, our findings support the view that calls do not mainly function as courtship signals, although they might serve both a territorial (sex-independent) and a courtship function. The comparison of responses to awake vs. anaesthetized intruders suggests that the latter are sufficient to elicit vocal activity. The subtle acoustic differences, however, indicate that the subject differentiates between intruder states.     

Sexual dominance in hermit crab shell fights: asymmetries in owner–intruder status,crab size,and resource value between sexes

We studied sexual dominance and seasonal differences in aggressiveness of individuals in intraspecific competition for shells of the hermit crab Pagurus filholi in terms of size of contestants and duration of the attempt to deprive other crabs of their shell. Experiments were conducted using paired intrasexual and intersexual contests in the pre-breeding and post-breeding seasons. Size ratios between contestants were systematically varied to assess the sexual difference in size and owner advantages. In both intrasexual and intersexual contests intruder crabs tended to win the contests more often as their size increased, that is, size advantage overcame owner advantage. Although we did not recognize a sexual difference in size and owner advantages in contest outcomes, male intruder crabs took a shorter time to deprive female owners of a shell than to deprive male owners. Furthermore, male individuals in the pre-breeding season had significantly longer fight durations. Fighting is costly. Thus males can afford to expend more energy and time fighting, indicating that males are dominant over females in shell fights as both intruders and owners. Electronic Publication     

Sexual Differences in Offspring Defense in a Monogamous Cichlid Fish

Solitary male and female Texas cichlids, Cichlasoma cyanoguttatum employ different tactics when defending their offspring. Males emphasize the lateral display against an intruder and make infrequent return visits to the young while the intruder is present. The female is more likely to attack, attempting to bite the intruder and also making frequent return visits to the offspring. These differences disappear with increasing age of the offspring as the male becomes more like the female with more violent attacks. Both defense systems appear effective, in the short term, against intruders.     

Postpartum aggression in mice: Experiential and environmental factors

Six experiments were conducted to assess the influence of duration of lactation, the presence of young, and the stimulus characteristics of intruder animals upon postpartum aggression of mice. The first experiment showed that postpartum aggression toward conspecifics was highest between Day 3 and Day 8, declined between Day 9 and Day 14, and was present toward males but absent toward females between Day 15 and Day 21 of the lactation period. Experiment 2 showed that lactating mice rarely attacked conspecifics to which they had been previously exposed but would readily attack strangers. Experiment 3 and 4 demonstrated that lactating animals never attacked intruders when tested 5 hr after pup removal. However, placement of young behind a wire partition in the home-cage for 5 hr or replacement of the offspring for as little as 5 min following 5 hr of separation restored postpartum aggression. The fifth experiment showed that 1- and 10-day old intruders were seldom attacked while intense aggression was directed against 14- and 20-day old intruders. Finally, Experiment 6 demonstrated that 14-day old intruders whose hair was removed were rarely attacked.     

Imperfect assessment and limited information preclude optimal strategies in male-male fights in the orb-weaving spider Metellina mengei

Agonistic behaviour between male orb-web spiders Metellina mengei competing for access to female webs was examined in field experiments to test the major predictions of game theory. Winners of fights were significantly larger than losers, particularly with respect to the length of the first pair of legs, which are sexually dimorphic in this species and used extensively in agonistic encounters. The size of the winning male had no influence on contest intensity or duration, and neither did relative size. However, fight intensity and duration were both positively correlated with the size of the losing male. Resident males won significantly more contests than intruders. Winning intruders were significantly larger than winning residents and it was these winning intruders that tended to produce the longer fights. Female weight and hence reproductive value had a marked influence on fight intensity and duration of fights won by the intruder but not those won by the resident. This indicates that only the resident obtains information about the female. These data are discussed with reference to the discrepancy with theory and a failure of some contestants to obtain information on resource value and relative contestant size necessary to optimize fight strategy.     

Behavioural and physiological consequences of home advantage resource holding in male guinea pigs

This study tested the physiological and behavioural effects of a resource asymmetry between guinea pig males. Asymmetry was defined via cohabitation with a female in a home area during a 5-day period. An experimental setup was designed in which the same male alternated between being in this home area (and monopolizing a female therein) and playing the role of an intruder. The confrontation phase between the resource holder and the intruder lasted 5 days. Behavioural data were collected for 0.5 h at the start of the confrontation and again for 0.5 h at 3, 24, 72, and 120 h after the onset of confrontation. Blood samples were taken before the test started and after 3, 72, and 120 h. Body mass was registered before and after the confrontation phase. The results indicate that resource holders were more highly motivated than intruders in displaying aggressiveness. They exhibited more chasing behaviour and won more confrontations. Both male groups performed almost identical frequencies of threat displays and courtship behaviour. Physiologically, the resource holders lost less body mass and showed increased plasma testosterone concentrations after 120 h. Resource holders also exhibited steady plasma corticosteroid concentrations, whereas the intruders showed elevated corticosteroid levels at 72 and 120 h. The results show that when acting as a resource holder, males react according to the resident-always-wins hypothesis, and when acting as an intruder, lose most confrontations. We conclude that the behavioural and physiological reaction of males during confrontations is determined to a greater extent by environmental asymmetries than by experiences or physical conditions before the conflict. Communicated by R.F. Oliveira     

Effects of resource distribution on activity and territory defence in house mice, Mus domesticus

How much an animal invests in defending a territory depends, in part, on the quality, quantity and distribution of resources, but do animals target their investment in defence within the territory according to the location of different resources? We examined whether wild-caught male and female house mice increased aggression towards intruders of the same or opposite sex when in areas containing their food/water or nest site. Pairs of resident mice were established in enclosures consisting of two areas. In split enclosures, food and water were in one area and a nest site in the other, while in clumped enclosures all resources were in one area with no resources in the other. Residents of both sexes were quicker to attack intruders, and were more aggressive, in areas containing resources than in an area containing no resources. There was no difference in aggression between areas containing food/water, a nest site or both. The effect of resources on territorial defence was due largely to changes in aggression against intruders of the opposite sex. Residents, especially males, attacked intruders of their own sex more than those of the opposite sex. This bias was much stronger when no resources were present, owing to low aggression against opposite-sex intruders in the absence of resources. There were no differences in intruder aggression, and the relative size of the intruder (heavier or lighter than the resident) had no effect on resident aggression. Residents also spent less time out of their nest in clumped enclosures owing to low activity in the nonresource area.     

Crowding pregnant mice affects attack and threat behavior of male offspring

Attack and threat behavior of adult male offspring of female mice crowded during the final third of pregnancy was investigated. In 5-min test pairings with an anosmic "standard opponent" which had 50 microliter of male mouse urine applied to its fur, the prenatally stressed group of males showed significantly less attack behavior; attack latency was longer and number of attacks, bites, amount of time spent attacking, and composite aggression scores were all lower, compared with the control group. Similarly, less threat behavior was observed in offspring from crowded dams; there were lower frequencies of tail rattles, rough grooms, and upright threats. Additionally, proportionally fewer males in the prenatally stressed group attacked or displayed threats. A second experiment was designed to investigate the effects of exogenous androgen on the aggressiveness of males from crowded mice: testosterone propionate administration (500 micrograms/animal/day, for 5 days prior to testing) abolished differences both in the proportion of males from crowded mice that fought and also apparently abolished differences in intensities of attack and threat behavior between groups. However, trends toward reduced aggression in prenatally crowded males remained. More detailed analysis of these responses, based only on animals that displayed aggression, revealed significantly reduced intensity of aggression in offspring from females crowded during pregnancy, indicating that testosterone propionate therapy did not completely restore this behavior. In order to reduce postnatal effects due to possible differences in mothering, all offspring were fostered to untreated mothers at birth. The results are discussed in terms of in utero exposure of male fetuses of crowded dams to stress-liberated adrenal steroids of maternal origin, and the possible consequences for the endocrine integrity of these offspring.     

Male Defense of the Breeding Cavity and Factors Affecting the Persistence of Breeding Pairs in the Stomatopod,Gonodactylus bredini (Manning) (Crustacea: Hoplocarida)

In Caribbean Panama, nonreproductive male and female stomatopods are solitary and defend their own coral-rubble cavities. When breeding pairs form, however, males assume all responsibility for cavity defense. To compare success in cavity defense and defensive tactics among paired and unpaired males, and to examine the tendency for paired stomatopods to exchange their present mates for larger (higher quality) individuals, we introduced same-sized and 15% larger male, and same-sized and 15% larger reproductive female intruders to paired and unpaired male residents in a balanced design. Paired males were more successful at cavity defense than unpaired males, evidently because paired males strike intruders more than unpaired males, and because intruders fight less intensely against paired males than against unpaired males. Paired males occasionally attempted extrapair copulations, but showed little tendency to abandon their mates in favor of larger females. Paired females, however, mated readily with intruder males that evicted resident males. Populationwide female breeding synchrony and prolonged female receptivity before oviposition reduce variance in male mating success and may force males to guard the breeding cavity to assure their paternity. Uncertainty about the reproductive condition of intruder females may prevent males from exchanging mates.     

Asymmetrical Resource Ownership Increases Owners’ and Reduces Non‐Owners’ Motivation to Fight in the Mangrove Rivulus,Kryptolebias marmoratus

Resource ownership often increases an individual's aggressiveness and its probability of defeating a competitor. Individuals contesting resource owners could therefore incur higher costs, making individuals reluctant to compete with owners. We tested the hypothesis that animals use asymmetry in resource ownership as a cue for contest costs and adjust contest decisions accordingly. Using a mangrove rivulus fish (Kryptolebias marmoratus), we staged (1) contests with a randomly assigned asymmetry in resource ownership (one fish was provided with a shelter) and (2) contests in which neither fish had a shelter. Owners that were in their shelters at the contest start showed a greater tendency to fight and won more contests than their intruder opponents; those outside the shelter at the start did not. Compared with fish in contests with no shelters at stake, shelter owners had a higher tendency to fight whether or not they were in their shelters at the start; intruders, however, had a lower tendency to fight only against owners that were inside the shelter at the start. These results demonstrate (1) that ownership status influences both owners’ and intruders’ contest decisions (and in opposite directions), producing a detectable ownership advantage and (2) that intruders required confirmation of their opponents’ ownership status before retreating without challenging them. Ownership status per se is therefore important to the fish's contest decisions.     

Aggressive behaviour of male mice (Mus musculus) towards familiar and unfamiliar opponents.

Two types of colonies of laboratory mice were employed; hierarchically organized ones formed by placing five unfamiliar 8-week-old mice in a cage together and amicably organized colonies in which four male litter mates were kept together throughout the whole period of the experiment. During a 21-day pre-experimental period intra-colony aggressive behaviour was recorded. A dominant animal and ranked subordinates occurred in every hierarchical colony, whilst no aggression was recorded in any of the amicable colonies. During a 25-day period unfamiliar adult male or female mice were introduced daily into the amicable or hierarchical colonies for 10 min. In a third experiment juvenile mice 17, 24, 31 or 38 days old were introduced into hierarchically organized colonies during a 20-day period. In all hierarchical colonies the stranger was attacked irrespective of sex and age; the majority of attacks were carried out by the dominant mouse. Aggression by the dominant declined exponentially throughout the experimental period and regression analyses compared the different data. Unfamiliar adult females were the recipients of fewer attacks than unfamiliar adult males and the age of juvenile strangers was found to be an important factor. Amicably organized mice initially did not attack strangers, but over a period of 25 days the number of attacks on unfamiliar males gradually increased.     

Winning fights elevates testosterone levels in California mice and enhances future ability to win fights

The 'winner effect' has been studied in a variety of species, but only rarely in mammals. We compared effects of winning three, two, one, or zero resident-intruder encounters on the likelihood of winning a subsequent aggressive encounter in the California mouse (Peromyscus californicus). During the training phase, we ensured that resident males won all encounters by staging contests with mildly sedated, smaller intruders. During the test phase, the resident male encountered an unfamiliar, more evenly matched intruder that had experience winning an encounter and was larger than the resident. Testosterone (T) plasma levels significantly increased after the final test when they had experienced two prior winning encounters, and the probability of winning a future encounter increased significantly after three prior wins independent of intrinsic fighting ability. We hypothesize a 'winner-challenge' effect in which increased T levels serve to reinforce the winner effect in male California mice.     

不同地位克氏原螯虾对社会环境改变的行为反应

为探究不同社会地位的亚成体克氏原螯虾(Procambarus clarkii)对利益的分配,以及对社会环境改变(体型较大的入侵同类)各自做出的行为反应,本文统计已确立等级序列的成对同性别螯虾对隐蔽所的占有情况,并视频拍摄同性别螯虾入侵时,入侵者及不同地位的原居者的格斗行为,分析格斗次数、格斗时间、首次攻击及被首次攻击对象多个参数。在隐蔽所占有实验中,71%为优势者占有隐蔽所,7%为从属者占有,共同占有或没有占有的各11%。在入侵-反入侵实验中,雄性入侵者对2只原居虾之间的攻击选择无显著差异,雌性入侵者首次攻击显著选择与原优势者格斗;雄性原居者中优势者对入侵者和原从属者的攻击无显著差异,雌性优势者则显著选择与入侵者格斗;雄性原居者中从属者显著选择与入侵者格斗,雌性从属者对入侵者和原优势者的攻击无显著差异。上述结果表明,优势者优先占有资源;雌雄螯虾在入侵和面临入侵时有各自不同的策略;不同地位的螯虾在反入侵时行为反应也各有不同。     

The Costs of Confronting Infanticidal Intruders in a Burying Beetle

Infanticide by unrelated adults is a complex behavior in burying beetles (Nicrophorus spp.) serving multiple functions (resource competition, access to mates, cannibalism). The costs of confronting an infanticidal intruder are likely to vary with context. To assess these costs for a single female parent (Nicrophorus pustulatus), we systematically manipulated the sex of a conspecific intruder and the timing of the intrusion. Male intruders were a greater takeover threat than female intruders, and infanticidal takeovers were more common earlier in the breeding cycle. Even though a male intruder posed a greater threat to the brood, a female intruder was a greater threat to the reproductive success of the resident female. Female intruders that took over a carcass excluded the resident female from the resource. When a male intruder took over a carcass, the resident female was able to recover much of her loss by producing a replacement brood. Even when females successfully defended their broods, they suffered decreased reproductive output relative to control females that never had to confront an intruder (expt 1), but the mechanisms underlying this cost were unclear. To test the hypothesis that defeated female intruders attempt to parasitize late‐stage broods, female intruders whose eggs could be identified by a fat‐soluble dye were introduced to resident females caring for larvae (expt 2). Fifteen of 20 intruders oviposited eggs and the number of eggs were related to intrusion pressure. Because resident females rarely produced eggs while caring for larvae, it is uncertain whether the behavior of the defeated female should be characterized as brood parasitism, a failed takeover attempt, or an attempt to use the remains of the depleted resource. This study provides the most complete picture of the changing costs of confronting an infanticidal threat throughout the vulnerable period of offspring development.     

Male <Emphasis Type="Italic">Betta splendends</Emphasis> are equally aggressive toward neighbors and strangers

Male Siamese fighting fish, Betta splendens, are known for their ferociousness when defending their territories against male or female conspecific intruders. The aim of this study was to investigate whether this species exhibits the dear enemy phenomenon, where territorial males would be less aggressive toward neighbors with already established territories than toward complete stranger males. In experiment 1, a male Betta was placed in an aquarium. A second male was placed in a glass jar that was fitted in one corner of the aquarium. After 24 h, a cover was placed around the jar, and then removed after 1 h. Alternatively, the jar male was replaced with a different male of a different color while the jar was covered. The frequency and duration of opercular expansion by the aquarium male were recorded. In experiment 2, two male Bettas were placed, each in a half of a large aquarium that was equally divided by a clear, perforated Plexiglas divider. After 24 h, one male (intruder) was transferred into the other male’s (resident) half. In another variation of the experiment, the intruder was a stranger male Betta that the resident male had never encountered before. The opercular expansion and duration were scored for the resident male. The results indicated that male Siamese fish reacted similarly to familiar and stranger males. This lack of dear enemy effect in Siamese fish could be due to an inability to discriminate between neighboring males and non-neighboring males. Alternatively, territorial males could be equally aggressive to all intruders because all intruders represent equal danger.