Inter-male aggression with regard to polygynous mating system in Pampean grassland mouse, Akodon azarae (Cricetidae: Sigmodontinae)

Based on the hypothesis that, in Akodon azarae, polygyny operates through female defence, we studied inter-male aggression in order to test the following predictions: during the breeding period (1) resident males are more aggressive than intruder males in the presence of females (FP), and (2) aggressive behaviour is independent of male condition (resident or intruder) in the absence of females (FA). To test our predictions, we used the resident male behavioural response towards an intruder male in relation to FP or FA. We conducted 30 encounters in FP and 27 in FA in 0.79-m² round enclosures placed in the Espinal Reservation. Our results support the prediction that, in FP, the intensity of aggressive behaviour exhibited by males varied in relation to resident or intruder condition. Resident males showed high levels of aggression towards intruders, and intruders exhibited the greatest values of submissive behaviours with residents. In FA, the intensity of aggressive behaviour did not vary in relation to resident or intruder condition. Both resident and intruder males exhibited low aggressive behaviour and inter-male encounters resulted mainly in non interactive behaviours. Our results support the hypothesis that, in A. azarae, the polygynous mating system operates through female defence.     

Territorial aggression can be sensitive to the status of heterospecific intruders

Territorial animals are known to be able to differentiate between intruding individuals posing a low or high threat and adjust their aggressive response accordingly. However, plastic territorial aggression based on recognising individuals with different attributes is typically assumed to be relevant only in the context of conspecific interactions. In this study, we investigated territorial aggression of neotropical cichlid fish in their natural habitat to assess whether responses to different types of individuals of another species can also be plastic. We show that arrow cichlids (Amphilophus zaliosus) adjusted their territorial aggression regarding the status of heterospecific intruders: breeding individuals of Amphilophus astorquii received a lower level of aggression than non-breeders. The same pattern was also found for the two different types of A. astorquii individuals intruding into conspecific territories. These results suggest that heterospecific individuals should not be ignored when considering selection pressures shaping plasticity of aggressive behaviour in territorial animals.     

Aggression patterns in adaptation and speciation of subterranean mole rats

The patterns and correlates of aggressionwithin andbetween actively speciating subterranean mole rats of theSpalax ehrenbergi superspecies in Israel were tested in an attempt to approach an evolutionary theory of aggression related to both adaptation and speciation. Laboratory experiments were conducted on 314 adult mole rats (188 males and 126 females) caught in nature, representing 12 populations and comprising four chromosomal species (2n = 52, 54, 58, 60). The present analysis concentrates on “total aggression” comprising four aggressive variables: attacks, bitings, head-ons and sniffing with open mouth, out of 23 recorded variables. The results indicate that (a) aggression is distributed as a multipeak, discontinuous phenotypic parameter, displaying polymorphism within all 12 populations and 4 species tested; (b) the level of aggression was higher in males than in females, in the breeding than in the nonbreeding season, and displayed clinal geographic variation in both sexes across the superspecies range, where “militancy” increased northwards; (c) aggression was significantly correlated with ecological, physiological, genetical and ethological factors. These results support an evolutionary theory of aggression related to the intertwined processes of adaptation and speciation.     

Aggressive behaviour and nest-site defence during the breeding season in an island kestrel population

We examined the changes in the intensity of intra-specific aggression (ISA) in a kestrel (Falco tinnunculus canariensis) population throughout the breeding season, in order to evaluate the main hypotheses regarding ISA (defence of parentage, food, nest-site and offspring). Each pair was presented with a live caged adult male or female as a conspecific intruder. The intensity of ISA did not vary significantly during the breeding season, and nor did it correlate with offspring value, or food availability. Our island-dwelling kestrel population showed low fluctuation in ISA during the breeding season, and similar levels of aggression in both members of the pair towards male or female intruders. These findings support the nest-site defence hypothesis. Pairs with more exposed nest-sites showed increased ISA, unexplained by offspring value. We suggest that this behaviour is related with nest-site conspicuousness and vulnerability. The pairs displaying greater ISA also showed greater aggression towards humans, the main local predator. This may indicate that defensive behaviour evolved in the past when it was advantageous against predators, and became a trait. Compared with the non-island reference population, we only found differences in ISA during the pre-laying phase: lower intensity of aggressive behaviour between males, and higher between females.     

The function of threat display in wintering great tits

It is generally accepted that threat displays have signal function and serve to repel opponents, but why they should have such an effect is a matter of debate. Using videotaped interactions from 2 years and nine flocks each year, we analysed the pattern of occurrence of agonistic displays and attacks in captive flocks of wintering great tits, Parus major. One currently influential perspective is to view threat interactions as sequences of distinct moves and countermoves, in which stronger threats can reliably indicate aggressive motivation because they carry a cost of eliciting counterattacks from certain opponents. We found little evidence for this kind of mechanism operating in great tits. Another possibility is that social dominance is important in ensuring reliable communication, for instance because an individual needs both to display and to attack to maintain its rank. It would be hard to test the importance of this kind of mechanism directly, but our observations were consistent with such an idea. Dominants won the majority of bouts of interaction and had higher rates of display and attack than subordinates, but a subordinate could temporarily overcome a dominant through intense aggression. We also found that the level of displaying in one bout provided information about the rate of aggression in following bouts. As a general interpretation of great tit aggressive behaviour, we suggest that particular displays form part of a graded signal of motivational state, so that a greater rate of displaying or a greater proportion of intense displays, together with a greater rate of attacking, correspond to a greater aggressive motivation. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Intraspecific competition and aggression for shells in the hermit crab Pagurus samuel1s

Many studies have investigated shell‐related behaviour in hermit crabs. Few studies, however, have focused specifically on the intraspecies aggression associated with shell competition. We examined intraspecies aggression in hermit crab (Pagurus samuelis) pairs as it relates to competition for a limiting resource, gastropod shells. Pairs of hermit crabs were observed in the laboratory in four different treatments that varied the presence or absence of shells for one or both of the crabs. Measurements of the latency to respond, the number of bouts, and the fight durations were recorded. There was a significant difference among treatments for all three measurements, and naked hermit crabs were much more aggressive than housed hermit crabs. There was no significant difference in aggression between males and females in any of the three treatments. The heightened aggression observed in naked P. samuelis is likely in service of acquiring a protective shell.     

Contexts and predictability of aggression in chimpanzees

Reasons for aggression may be deduced from the situations preceding aggressive behaviour. This we may call the retrospective approach. In addition to results from this conventional procedure the present paper investigates the predictability of aggressive behaviour. In this so-called anticipatory approach, particular non-agonistic events were used as predictors of aggression. These events had in common that they were intuitively judged to be unpleasant for one of the chimpanzees. This individual was observed during the next 10 min in order to establish if its inclination to exhibit aggression had increased shortly after that seemingly frustrating event. The studies reported here were carried out on a colony of 20 semi-free-living chimpanzees Pan troglodytes in Arnhem Zoo. The data show that: (1) half of all aggressive actions arose in agonistic context: thus aggression was contagious; (2) adult males performed apparent spontaneous aggression remarkably often: this result is suggested to reflect dominance-rivalry among males; (3) aggressive behaviour was predictable on the basis of intuition, but attempts to account for the observer's intuitive knowledge were not entirely successful.     

Intra- and interspecific agonistic behaviour in sympatric harriers during the breeding season

We analysed the temporal and sexual patterns of intra- and interspecific aggression in sympatric harriers during the breeding season, to determine the main resource defended (food, nest sites, mates) and how factors such as body size or breeding system (territorial versus colonial) influence aggressive behaviour. We predicted that if aggression is (at least partly) related to competition for food, the hen harrier, Circus cyaneus, because of its large size and territorial system, should be more aggressive (both intra- and interspecifically) than the smaller, colonial Montagu's harrier, Circus pygargus. The intraspecific aggression rate of both harriers peaked early in the season, was mainly intrasexual and increased with the number of neighbours. These patterns support the mate competition hypothesis to explain intraspecific aggression in both species. Montagu's harriers were more aggressive towards conspecifics than hen harriers. Their aggression rates towards heterospecifics were high at the start of the season then decreased, supporting the hypothesis that interspecific aggression serves primarily for nest site defence. Hen harriers showed lower interspecific aggression rates in the prelaying period and a gradual increase throughout the breeding cycle, particularly by females, who hunt around the nest for food for the nestlings. These patterns correspond to food competition, as food resources around the nest are probably more important and interspecific intrusion more costly for territorial hen harriers than for colonial Montagu's harriers. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

The role of the frontal cortex of the left and right hemispheres in regulating the intraspecific interaction of rats with differing zoosocial experience

By means of the analysis of probabilistic ethological structures of the intraspecies rats behaviour in the test "stranger-resident" and by the method of isolated destruction of the frontal cortex or the left and right hemispheres,--it is established that the frontal cortex of the left hemisphere regulates the intraspecies sociability and keeps it on a certain level, because the change (increase) of the intraspecies sociability is observed only at the unilateral destruction of the left frontal cortex. Isolation from the relatives in early ontogenesis leads to the loss of regulating influence of the left frontal cortex on the intraspecies sociability. In isolated animals the frontal cortex of the left hemisphere regulates the level and probability of the appearance of aggression. The right hemisphere ensures the adequate inclusion of the aggression in the probabilistic ethological structure of the total intraspecies behaviour; at its damage the forms appear of an adequate pathological aggression, and connection is lost between the elements of aggressive behaviour and the acts of intraspecies sociability.     

Play-like behaviour: An essay in speculative ethology

It is claimed that certain processes of individual behaviour and of interaction between individuals run parallel. Such parallels are seen along three axes: antagonism-coordination, constriction-expansion and neutral-play-like. Characteristics of ritualized behaviour and play are analysed and the two categories of behaviour are compared in detail. They are shown to differ largely in degree of expansion. They also differ along the antagonism-coordination axis. Both are play-like. A theory comprising both ritualized behaviour and play is attempted. The concept of field interaction is introduced. The physical theory of elect ro-magnetism is used as a conceptual model. Positive and problematic aspects of the theory are discussed: Is the behavioural field polarised or not? The relationship between aggression and play-like behaviour is considered, both as a motivational problem and in phylogenetic perspective. Some limits to the theory are indicated. Relationships of the theory to some other theories are briefly discussed.     

Trade-off between growth rate and aggression in juvenile coho salmon, Oncorhynchus kisutch

In juvenile salmon and trout, there seems to be a positive phenotypic correlation between individual aggression level and growth rate. Aggressive fish are dominant, and they obtain and defend territories, giving them access to good feeding sites. Being aggressive may increase predation risk, and may also carry costs such as increased metabolic demand, with effects on growth. To test the hypothesis that there is a trade-off between individual growth rate and aggression, we mated 12 female coho salmon with two unique males each, creating 24 full-sibling families. Growth of individually marked fish from each family was estimated in a situation where food could not be monopolized. Thereafter, individual fish were tested for mirror-elicited agonistic behaviour. We found significant variation between families in early growth rate, with a high heritability (1.04). There was also significant between-family variation in agonistic behaviour, but activity was generally low and heritability was low (0.25) and not significant. Growth rate and agonistic behaviour were negatively correlated. These results imply that aggressive behaviour has an energetic cost. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

Manipulation of colony environment modulates honey bee aggression and brain gene expression

The social environment plays an essential role in shaping behavior for most animals. Social effects on behavior are often linked to changes in brain gene expression. In the honey bee (Apis mellifera L.), social modulation of individual aggression allows colonies to adjust the intensity with which they defend their hive in response to predation threat. Previous research has showed social effects on both aggression and aggression‐related brain gene expression in honey bees, caused by alarm pheromone and unknown factors related to colony genotype. For example, some bees from less aggressive genetic stock reared in colonies with genetic predispositions toward increased aggression show both increased aggression and more aggressive‐like brain gene expression profiles. We tested the hypothesis that exposure to a colony environment influenced by high levels of predation threat results in increased aggression and aggressive‐like gene expression patterns in individual bees. We assessed gene expression using four marker genes. Experimentally induced predation threats modified behavior, but the effect was opposite of our predictions: disturbed colonies showed decreased aggression. Disturbed colonies also decreased foraging activity, suggesting that they did not habituate to threats; other explanations for this finding are discussed. Bees in disturbed colonies also showed changes in brain gene expression, some of which paralleled behavioral findings. These results show that bee aggression and associated molecular processes are subject to complex social influences .     

Pharmacoethological analysis of the action of beta-carbolines

The ethological spectrum of four beta-carbolines was studied for intraspecific aggression, sociability integration and individual behavior. It was established that beta-carboline-3-carboxyethyl ether given at subconvulsant doses increases aggression, lowers sociability as a diazepam antagonist. Diazepam reduces aggression and increases sociability. It is suggested that benzodiazepine receptors are involved in the integration of interspecific sociability and aggressive behavior of the "attack" type. Aggression of the defence type has no inhibitory beta-carboline control (1-methyl-6-methoxy-dihydro-beta-carboline). The differences in the ethological spectra of beta-carbolines have been demonstrated.     

Differential gene expression in brain tissues of aggressive and non-aggressive dogs

Background  

Canine behavioural problems, in particular aggression, are important reasons for euthanasia of otherwise healthy dogs. Aggressive behaviour in dogs also represents an animal welfare problem and a public threat. Elucidating the genetic background of adverse behaviour can provide valuable information to breeding programs and aid the development of drugs aimed at treating undesirable behaviour. With the intentions of identifying gene-specific expression in particular brain parts and comparing brains of aggressive and non-aggressive dogs, we studied amygdala, frontal cortex, hypothalamus and parietal cortex, as these tissues are reported to be involved in emotional reactions, including aggression. Based on quantitative real-time PCR (qRT-PCR) in 20 brains, obtained from 11 dogs euthanised because of aggressive behaviour and nine non-aggressive dogs, we studied expression of nine genes identified in an initial screening by subtraction hybridisation.     

Coping with social tension: Sex differences in the effect of food provision to small rhesus monkey groups

In previous research, male chimpanzees (Pan troglodytes) were found to be more ‘conciliatory’ than females, in that after aggressive incidents males more often engaged in socially positive contact with former opponents. The first part of the present paper presents similar results for a large breeding troop of rhesus monkeys (Macaca mulatta). To investigate further the possible sex difference in postconflict behaviour, a series of experiments was carried out on small isosexual groups of young rhesus monkeys, three male and three female groups. Food competition was created by providing either a single apple piece (Monopoly test) or a handful of small pieces (Equality test). Observations lasted for 30 min, and behaviour in tests without food provisioning was also recorded. The predicted response to food was an increase in aggression followed by restorative behaviour, such as grooming. In Equality tests both sexes showed similar responses, i.e. increased aggression and decreased grooming and cohesiveness. In Monopoly tests, however, their responses differed. After the aggression increase, which occurred in all groups, males showed a significant increase in grooming and cohesiveness, whereas females showed the (non-significant) opposite trend. Another sex difference was that changes in grooming and aggression frequencies were related to the rank order in female groups, but not in male groups. Since there was no evidence for a direct causal connection between grooming and aggression, a new model is introduced which links grooming behaviour to the type of food provision rather than to the social disturbance by aggression. According to this model inequality in food distribution causes social tensions. Males actively try to reduce these tensions, whereas females do not. Some alternative explanations are also discussed.     

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.     

Sex-Specific Differences in Agonistic Behaviour,Sound Production and Auditory Sensitivity in the Callichthyid Armoured Catfish Megalechis thoracata

Background

Data on sex-specific differences in sound production, acoustic behaviour and hearing abilities in fishes are rare. Representatives of numerous catfish families are known to produce sounds in agonistic contexts (intraspecific aggression and interspecific disturbance situations) using their pectoral fins. The present study investigates differences in agonistic behaviour, sound production and hearing abilities in males and females of a callichthyid catfish.

Methodology/Principal Findings

Eight males and nine females of the armoured catfish Megalechis thoracata were investigated. Agonistic behaviour displayed during male-male and female-female dyadic contests and sounds emitted were recorded, sound characteristics analysed and hearing thresholds measured using the auditory evoked potential (AEP) recording technique. Male pectoral spines were on average 1.7-fold longer than those of same-sized females. Visual and acoustic threat displays differed between sexes. Males produced low-frequency harmonic barks at longer distances and thumps at close distances, whereas females emitted broad-band pulsed crackles when close to each other. Female aggressive sounds were significantly shorter than those of males (167 ms versus 219 to 240 ms) and of higher dominant frequency (562 Hz versus 132 to 403 Hz). Sound duration and sound level were positively correlated with body and pectoral spine length, but dominant frequency was inversely correlated only to spine length. Both sexes showed a similar U-shaped hearing curve with lowest thresholds between 0.2 and 1 kHz and a drop in sensitivity above 1 kHz. The main energies of sounds were located at the most sensitive frequencies.

Conclusions/Significance

Current data demonstrate that both male and female M. thoracata produce aggressive sounds, but the behavioural contexts and sound characteristics differ between sexes. Sexes do not differ in hearing, but it remains to be clarified if this is a general pattern among fish. This is the first study to describe sex-specific differences in agonistic behaviour in fishes.     

Animal signals: Ethological and games-theory approaches are not incompatible

Recent authors have contrasted the ‘traditional ethological approach’ to the study of animal signals with that derived from games theory. It is argued here that the ‘traditonal ethological approach’ they portray is not in keeping with main stream of ethological research on animal signals. In particular, it has not been generally assumed that the evolution of animal signals was based on selection for mutual benefit of actor and reactor, nor that signals carry precise information of what the actor will do next. A synthesis of the ethological and games-theory approaches is possible. It is suggested that many threat displays may signal ‘Will stay, but attack if provoked’ or ‘Will stay, but will flee if provoked’, and that the subsequent behaviour of the displaying bird depends in part on that of the reactor.     

Interspecific aggressive behaviour by long-toed lapwings (Vanellus crassirostris)

In this paper interspecific aggression is related to ecological relationships between bird species. Three kinds of relationships are distinguished according to the presence or absence of aggression and the degree to which resources are exclusively utilized: interspecific territoriality (aggression, exclusive use), partial exclusion (aggression, no exclusive use), and tolerance (no aggression, no exclusive use). Aggression by long-toed lapwings (Vanellus crassirostris) is examined within this framework, using quantitative data gathered through observational sampling. Lapwings were interspecifically territorial with one species, partially excluded five species, and tolerated four species. The results indicate that the function of interspecific aggressive behaviour may be related to competition, predation, or both.     

Indirect genetic effects contribute substantially to heritable variation in aggression-related traits in group-housed mink (Neovison vison)

Background

Since the recommendations on group housing of mink (Neovison vison) were adopted by the Council of Europe in 1999, it has become common in mink production in Europe. Group housing is advantageous from a production perspective, but can lead to aggression between animals and thus raises a welfare issue. Bite marks on the animals are an indicator of this aggressive behaviour and thus selection against frequency of bite marks should reduce aggression and improve animal welfare. Bite marks on one individual reflect the aggression of its group members, which means that the number of bite marks carried by one individual depends on the behaviour of other individuals and that it may have a genetic basis. Thus, for a successful breeding strategy it could be crucial to consider both direct (DGE) and indirect (IGE) genetic effects on this trait. However, to date no study has investigated the genetic basis of bite marks in mink.

Result and discussion

A model that included DGE and IGE fitted the data significantly better than a model with DGE only, and IGE contributed a substantial proportion of the heritable variation available for response to selection. In the model with IGE, the total heritable variation expressed as the proportion of phenotypic variance (T²) was six times greater than classical heritability (h²). For instance, for total bite marks, T² was equal to 0.61, while h² was equal to 0.10. The genetic correlation between direct and indirect effects ranged from 0.55 for neck bite marks to 0.99 for tail bite marks. This positive correlation suggests that mink have a tendency to fight in a reciprocal way (giving and receiving bites) and thus, a genotype that confers a tendency to bite other individuals can also cause its bearer to receive more bites.

Conclusion

Both direct and indirect genetic effects contribute to variation in number of bite marks in group-housed mink. Thus, a genetic selection design that includes both direct genetic and indirect genetic effects could reduce the frequency of bite marks and probably aggression behaviour in group-housed mink.