Social isolation increases male aggression toward females in the field cricket Gryllus bimaculatus

Social isolation has often been reported to facilitate male aggressiveness in various animal species. If social isolation also escalates male aggressive behavior towards females, the mating success of the aggressive males will be low. This study evaluated the effect of social isolation on mating behavior in the field cricket Gryllus bimaculatus, which has traditionally been considered to be an asocial species. The results showed that social isolation from same-sex individuals enhanced male aggressiveness to females, and the mating success of aggressive males was reduced under the experimental conditions. More aggressive males exhibited a longer latency to court than less aggressive males. These results suggest that because male aggressiveness causes a delay in courtship, aggressive males may have reduced mating success. This demonstrated that social relationships are a critical factor affecting male mating success, even if the species is normally considered solitary.     

Crustacean social behavioral changes in response to isolation

Periods of isolation during which animals have no social contact are common in the design of behavioral experiments. They are used, for example, to test memory and recognition responses, or to ensure a baseline condition before experimental manipulations commence. We investigated the effect of isolation periods on the aggressive behavior of matched pairs of the crayfish Cherax destructor in two contexts. The first experiment tested the effects of a period of isolation between two encounters. The second experiment tested the effects of isolation before an encounter by pairing one crayfish from a communal living environment with another crayfish from an isolated one. Fight outcome and aggression levels were analyzed, resulting in three conclusions about the social biology of C. destructor. First, encounters between familiar opponents are influenced by the outcome of the familiarization fight for about 2 weeks. Second, the level of aggression and the outcome of an encounter are affected over different time frames. Third, individuals that are isolated before an encounter can be disadvantaged. These data suggest that isolation, or events that occur during periods of isolation, affect multiple elements of social behavior in C. destructor. This suggestion has implications for the interpretation of previous results and future studies in crustaceans and other taxa.     

Untersuchungen zum Aggressionsverhalten von Poecilia sphenops (Pisces,Poeciliidae) vor der Geschlechtsreife3

After 14 days of social isolation, half-grown Poecilia sphenops still showed no sexual activity but an increase in biting and tail beating. In relation to this result data on diurnal rhythms of the observed activities, the biological significance of the aggressivity in immature fish, and the ontogeny of the aggressive and sexual behaviour are presented. In contrast to the aggressive behaviour no sexual activity could be observed before maturity.     

Ontogenetic and gender-related variation in the isotopic niche within and between three species of fur seals (genus <Emphasis Type="Italic">Arctocephalus</Emphasis>)

Differences in aggressiveness when competing for environmental resources are the main factor leading to social hierarchy in group living fish. Social status acquired is related to changes in physiological parameters, as metabolic rate. Habitat variation can interfere with aggressive behaviour and promote changes in physiological parameters associated with social status. The primary goal of our study was to investigate how differences in habitat complexity affect the relationship between resting metabolic rate (RMR) and social status in the Amazonian dwarf cichlid Apistogramma agassizii. We compared agonistic interactions between pairs of males in aquaria with different habitat enrichment levels, manipulated by adding shelters. RMR was measured before and after hierarchy establishment. Habitat enrichment promotes changes in aggressive behaviour and influences differences in metabolic rate between dominant and subordinate fish. We observed an increase in biting by dominant fish at high enrichment habitat, which could be related to the increase in territory value. We observed an increase in metabolic rate in dominant fish after hierarchy establishment. However, it occurs only in enriched habitats. We concluded that habitat structure interfere with behavioural characteristics in social hierarchies, as aggressiveness, and changes in aggressive interactions affect metabolic rate in different social ranks in the dwarf cichlid Apistogramma agassizii.     

Effects of social isolation on aggressiveness in fish with special reference to the swordtail (Xiphophorus helleri)

The attack readiness of socially isolated male swordtails ($\text{Xiphophorus helleri}$) was measured in two different aggression tests: (a) In the standard–opponent test the aggressive responses to a small, rather passive opponent swimming in a small adjacent compartment of the test aquarium were counted. (b) In the mirror test the subjects were confronted with their own mirror image. Social isolation for 4 weeks resulted in a decrease of attack readiness in both test situations. It is concluded that -- similar to cichlids -- external stimuli from conspecifics are essential for maintaining the level of aggression readiness characteristic for socially living males. There are good reasons to propose that the effect of external stimuli is mediated by the pituitary-gonadal axis. On the other hand, it is well known from previous studies that, in escalated encounters between freely swimming males, isolated males fight longer and at higher intensities for rank-order position than non-isolated males. The hypothesis is proposed that the high fighting intensity of previously isolated males could be due to an experience levelling effect of social isolation, i.e. the influence of different agonistic experience is reduced by isolation.A critical review of the literature on isolation-induced decremental and incremental effects on aggression in teleosts is presented. Often it is extremely difficult to compare the results of different authors because the methods of behavioural measurement differ to a large extent. However, it is a striking fact that social isolation of immature fish in all known cases has resulted in an increase of aggressiveness. Probably in mature males, depending on species and circumstances, either an incremental or a decremental effect of social isolation may be adaptive. On the one hand, a spontaneous increase of attack readiness may be advantageous for defence of a territory or a position in the social hierarchy and improves the ability of a single individual to join a group of strange conspecifics. On the other hand, it may be economic to lower the level of androgenic hormones to some extent in the absence of conspecifics, and this could lead to a reduction of the aggression level.     

Nonaggressive interventions by third parties in conflicts among captive Bornean orangutans (<Emphasis Type="Italic">Pongo pygmaeus</Emphasis>)

Whereas orangutans are regarded as semisolitary animals in the wild, several studies have reported frequent social interactions, including aggression, among orangutans in captivity. As yet, there is a lack of knowledge about how they cope with aggression. In this report, we provide a number of new observations of interventions by third parties in aggressive interactions within a captive group of Bornean orangutans (Pongo pygmaeus) in the Tama Zoological Park, Japan. We observed that an adult female and a juvenile male orangutan intervened in aggressive interactions. The victim was a newly introduced juvenile female who was unrelated to anyone in the zoo. The ways in which the orangutans intervened were not aggressive, as the interveners simply aimed to separate the opponents, and these interventions did not lead to further aggression in almost every case. Our observations suggest that third parties can play an important role in managing aggressive conflicts among captive orangutans and, under conditions in which orangutans share limited space, nonaggressive interventions by third parties for settling conflicts appear. It is possible that orangutans may actively promote the peaceful coexistence of other individuals.     

Ontogenie und Funktion der Rangordnung innerhalb einer Geschwisterschar von Graugänsen (Anser anser L.)

Between the siblings of a family of greylag geese (Anser anser L.) a rank order is established by fighting during the first days after hatching, and also by other, more complex interactions. The position of each goose in the rank order can be defined by three groups of behaviour patterns (aggressive behaviour, “greetings”, escape responses). A clear-cut ranking relation between two individuals is a prerequisite for their close cohesion. The survival function of rank order lies in diminishing the intensity and frequency of agonistic interactions, in diminishing social stress situations and in guaranteeing the close family cohesion.     

Laboratory studies of the effects of short‐term isolation on aggressive behaviour in fish

A review of the literature on the effects of short‐term isolation on aggressive behaviour in a variety of fishes reveals different approaches and often conflicting results. This diversity is attributed to a number of factors, including different experimental conditions and specific differences in habitat, behavioural repertoire and social system. Some common trends are identified and the main hypotheses proposed to account for the different results are discussed. Some methodological issues as well as proximate and ultimate mechanisms are examined, and the importance of a standard experimental paradigm in comparative research emphasised.     

Social interactions associated with reproduction in the bushveld gerbil <Emphasis Type="Italic">Gerbilliscus leucogaster</Emphasis>

The study of social behaviour can give us important insights into the social and mating system of a species or population. We investigated male–female interactions in captive bushveld gerbils, Gerbilliscus leucogaster, to gain insight into the social behaviour and mating structure. We conducted two experiments. In experiment 1, we studied the interactions between two strangers for a week in a three-tank setup, which tested behavioural variation spatially and temporally. Although tolerance between strange males and females increased with time, it remained generally low throughout the experiment. Females appeared to be cautious of males initially, but they later became aggressive towards them. Males showed an increase in submissive behaviour over time. The ‘home’ cage did not appear to be defended by either sex. In experiment 2, we investigated the social interactions of male–female pairs during pregnancy and lactation. Aggression persisted throughout the study and amicable behaviour was low; females were much more aggressive than males. We did not detect changes in social behaviour with the progression of pregnancy and lactation. We suggest that pair bonding is unlikely and that promiscuity is the most probable mating system. Female aggression may be related to mate choice prior to mating and mate exclusion thereafter, while it may be a response to infanticide risk during pregnancy and lactation.     

Responses of Snow Voles, Chionomys nivalis, Towards Conspecific Cues Reflect Social Organization during Overwintering Periods

Among microtine rodents, reaction to chemical cues from conspecifics is assumed to reflect social and spatial relationships. Generally, strong attraction of particular odours correlates with non‐aggressive behaviour and high spatial tolerance towards odour donors, whereas weak attraction correlates with greater levels of aggression and spatial segregation. In the present study, we examined whether winter odour preferences of the snow vole Chionomys nivalis, a rock‐dwelling microtine principally found at high‐mountainous regions, differ from that of other vole species, owing to their different social organization during overwintering periods. The social structure of C. nivalis over the winter period is relatively unusual among vole species in that they become nomadic and solitary. In odour choice trials under laboratory conditions, we found that both males and females avoided zones with conspecific odours of both sexes in comparison with unscented control zones or own odours. These results are consistent with the elevated levels of intraspecific aggression and spatial isolation of C. nivalis during overwintering periods. Furthermore, scent‐elicited self‐grooming increased when their own odour was offered against conspecific cues. This, in combination with an active avoidance of conspecific odours, might functionally contribute to minimize direct confrontations between solitary individuals, thereby reducing the risks of aggressive encounters during overwintering periods.     

Absence of a prezygotic behavioural barrier to gene flow between the two sympatric morphs of the squat lobster <Emphasis Type="Italic">Munida gregaria</Emphasis> (Fabricius, 1793) (Decapoda: Anomura: Galatheidae)

Munida gregaria and M. subrugosa have been considered two different species for more than a century; however, after a recent molecular phylogenetic study, they are considered a single polymorphic species. Yet, the use of markers to diagnose species may be misleading when divergence between species is recent, since a speciation event may be obscured by the retention and stochastic sorting of ancestral polymorphisms. The morphs gregaria and subrugosa of Munida gregaria constitute an interesting case for the study of behavioural isolation since they are sympatric, breed at the same time of the year, and might have experienced a recent speciation. Mating behaviour observations and mate choice mating trials were conducted in order to investigate the potential existence of a behavioural prezygotic barrier to gene flow between these two morphs. Since factors involved in mate choice in galatheids are unknown, the four possible combinations of the two different morphs in trios were used to test for the existence of mate choice. Video recordings of all the possible trio combinations revealed that there was cross-attraction between males and females of different morphs. Females bearing partial broods participated in encounters as well as non-ovigerous females. The frequency and duration of homo- and heterotypic encounters were registered, and a reproductive isolation index was calculated for each variable for each trio. The isolation indexes calculated were not different from zero indicating random mating, and were not affected by the composition of the trio or the partial ovigerous condition of females. These results provided evidence of the absence of behavioural prezygotic barriers to gene flow between the morphs gregaria and subrugosa of M. gregaria.     

Intrinsic survival advantage of social insect queens depends on reproductive activation

The central trade‐off between reproduction and longevity dominates most species' life history. However, no mortality cost of reproduction is apparent in eusocial species, particularly social insects in the order Hymenoptera: one or a few individuals (typically referred to as queens) in a group specialize on reproduction and are generally longer lived than all other group members (typically referred to as workers), despite having the same genome. However, it is unclear whether this survival advantage is due to social facilitation by the group or an intrinsic, individual property. Furthermore, it is unknown whether the correlation between reproduction and longevity is due to a direct mechanistic link or an indirect consequence of the social role of the reproductives. To begin addressing these questions, we performed a comparison of queen and worker longevity in the ant Cardiocondyla obscurior under social isolation conditions. Survival of single queens and workers was compared under laboratory conditions, monitoring and controlling for brood production. Our results indicate that there is no intrinsic survival advantage of queens relative to workers unless individuals are becoming reproductively active. This interactive effect of caste and reproduction on life expectancy outside of the normal social context suggests that the positive correlation between reproduction and longevity in social insect queens is due to a direct link that can activate intrinsic survival mechanisms to ensure queen longevity.     

Bemerkungen zur Evolution von., Sprachen bei Säugetieren1 Zur Variabilität innerartlicher Kommunikation bei Caniden

Remarks on the evolution of “languages” of mammals. The variability of the intraspecific communication of canids The canid species mainly considered in this study, the golden jackal (Canis aureus), the cojote (Canis latrans), and especially the wolf (Canis lupus) have an extremely variable species-specific intraspecific communication. Communities, but also differences between the species become evident; the wolf with its distinct social behavior has the greatest differentiation. These results make also clear, that forms of communication in mammals develop in evolutionary processes and events. A safe statement on the beginning of the forms of communication in canids cannot be given so far. This is also true for humans. The ideas of Lenneberg (1967) of the parallel and disjunct origin of the forms of communication in mammals has a high probability for higher systematic units, but on lower systematic level a gradual change can be observed. The forms of communication in mammals are highly differentiated structures, where specific pecularities of the brain are of a special importance. The observations on the variability of intraspecific communication of canids, representing nonhuman mammals, suggest to define “language” in a very general form. In this way, similarities, analogies, and differences between the species of mammals, inclusive man, can be seen more clearly. In only this way the material can be gained to answer the question, if the language of man has in principle exceptional position biologically. It is important for the valuation of forms of intraspecific communication, the “languages”, of mammals not to choose human theory of life and human thinking as a basis, but to start with the biological pecularities of the species. Also canids, as higher mammals, have the ability to abstract, a form of thinking. In their special, species-specific way also non-human mammals arrange their area of experience and thinking. They form for themselves a species-specific “theory of live”. This will of course be different from the human one, according to the systematic position and the biological environment of the species. It is generally accepted that man has special abilities of thinking, he possesses an extraordinary developed brain, compared to all other mammals. This must effect his communication, his language, and result in differences to other mammal species. Gipper (1977) pointed out especially these differences, but they should not be interpreted as differences in principle. Canids use different gestures and sounds for communication within the species; dialog relations can be observed. Different connections of gestures and sounds have different contents of information. It is remarkable that canids do not use their total abilities in using the means of communication in the same way. This is very striking in the change from the wolf, as the only ancestor, to the domestic dog. It is evident that the brain has a greater importance in developing intraspecific communication, than do the peculuarities of organs causing the understanding. A dependanc of intraspecific communication from the environment can be observed either in differences of wird species of canids as well as in the domestication. In the domestic dog, but also in other domestic animals, there is an increase in vocalisations. But the content of information of sounds and gestures decreases in domestication, expressions of arousal become more important. A remarkable plasticity of intraspecific communication becomes evident by domestication. The transformations can be evaluated in the sense of Kosswig (1963, 1965) as first stages of a regressive evolution. Experiments of cross-breeding canid species show exemplarily that forms of communication within these mammal species have a highly polygene regulation, and that processes of learning have an influence. Also from this point of view, no basic differences between nonhuman mammals and humans can be assured, especially when taking into account the facts and thoughts described by Lenneberg. The findings in the canids give useful hints for a better understanding of the human language and its relations to the forms of communication in other mammals. Altogether one can conclude that genetic variability, numerous recombinations, the synorganisation of the processes caused in this way, and natural selection plays definite role in the evolution of “languages”.     

Detailed analysis of paternal knockout Grb10 mice suggests effects on stability of social behavior,rather than social dominance

Imprinted genes are highly expressed in monoaminergic regions of the midbrain and their functions in this area are thought to have an impact on mammalian social behaviors. One such imprinted gene is Grb10, of which the paternal allele is generally recognized as mediating social dominance behavior. However, there has been no detailed study of social dominance in Grb10 ^+/p mice. Moreover, the original study examined tube‐test behavior in isolated mice 10 months of age. Isolation testing favors more territorial and aggressive behaviors, and does not address social dominance strategies employed in group housing contexts. Furthermore, isolation stress impacts midbrain function and dominance related behavior, often through alterations in monoaminergic signaling. Thus, we undertook a systematic study of Grb10 ^+/p social rank and dominance behavior within the cage group, using a number of convergent behavioral tests. We examined both male and female mice to account for sex differences and tested cohorts aged 2, 6 and 10 months to examine any developments related to age. We found group‐housed Grb10 ^+/p mice do not show evidence of enhanced social dominance, but cages containing Grb10 ^+/p and wild‐type mice lacked the normal correlation between three different measures of social rank. Moreover, a separate study indicated isolation stress induced inconsistent changes in tube test behavior. Taken together, these data suggest future research on Grb10 ^+/p mice should focus on the stability of social behaviors, rather than dominance per se.     

Zur Funktion von Warnfarben: Die Reaktion junger Stare auf wespenähnlich schwarz-gelbe Attrappen

Social regulation of developmental rate of the African cichlid fish, Haplochromis bur-toni, was studied experimentally. Broods were raised under three different social conditions: total isolation, with no visual, physical or chemical contact with siblings; physical isolation, with no physical or chemical contact with siblings; and in social groups. Development of behavior and color patterns were observed in fish from 2 to 14 weeks of age. Males were sacrificed at 4-week intervals to measure growth and maturation state. Results showed that developmental rate is influenced by the social environment. Males with no territories (raised in groups) grew and matured more slowly than those in all other conditions. These males were not prevented from maturing but both the rate and phenotypic expression of maturation were inhibited.     

Habitat preference and flowering‐time variation contribute to reproductive isolation between diploid and autotetraploid Anacamptis pyramidalis

Tetraploid lineages are typically reproductively isolated from their diploid ancestors by post‐zygotic isolation via triploid sterility. Nevertheless, polyploids often also exhibit ecological divergence that could contribute to reproductive isolation from diploid ancestors. In this study, we disentangled the contribution of different forms of reproductive isolation between sympatric diploid and autotetraploid individuals of the food‐deceptive orchid Anacamptis pyramidalis by quantifying the strength of seven reproductive barriers: three prepollination, one post‐pollination prezygotic and three post‐zygotic. The overall reproductive isolation between the two cytotypes was found very high, with a preponderant contribution of two prepollination barriers, that is phenological and microhabitat differences. Although the contribution of post‐zygotic isolation (triploid sterility) is confirmed in our study, these results highlight that prepollination isolation, not necessarily involving pollinator preference, can represent a strong component of reproductive isolation between different cytotypes. Thus, in the context of polyploidy as quantum speciation, that generates reproductive isolation via triploid sterility, ecological divergence can strengthen the reproductive isolation between cytotypes, reducing the waste of gametes in low fitness interploidy crosses and thus favouring the initial establishment of the polyploid lineage. Under this light, speciation by polyploidy involves ecological processes and should not be strictly considered as a nonecological form of speciation.     

The origins of postmating reproductive isolation: testing hypotheses in the grasshopper <Emphasis Type="Italic">Chorthippus parallelus</Emphasis>

 Although there are several well-established hypotheses for the origins of postmating isolation during allopatric divergence, there have been very few attempts to determine their relative importance in nature. We have developed an approach based on knowledge of the differing evolutionary histories of populations within species that allows systematic comparison of the predictions of these hypotheses. In previous work, we have applied this methodology to mating signal variation and premating reproductive isolation between populations of the meadow grasshopper Chorthippus parallelus. Here we review the principles behind our approach and report a study measuring postmating isolation in the same set of populations. The populations have known and differing evolutionary histories and relationships resulting from the colonization of northern Europe following the last glaciation. We use a maximum-likelihood analysis to compare the observed pattern of postmating isolation with the predictions of the hypotheses that isolation primarily evolves either as a result of gradual accumulation of mutations in allopatry, or through processes associated with colonization, such as founder events. We also quantify the extent to which degree of postmating isolation can be predicted by genetic distance. Our results suggest that although there is only a weak correlation between genetic distance and postmating isolation, long periods of allopatry do lead to postmating isolation. In contrast to the pattern of premating isolation described in our previous study, colonization does not seem to be associated with increased postmating isolation. Received: January 24, 2002 / Accepted: July 26, 2002 Acknowledgments Numerous people helped with collecting and rearing grasshoppers. We are grateful to NERC for funding. Correspondence to:R.K. Butlin     

Enhancement of social isolation-induced aggressive behavior of young mice by zinc deficiency

Neuropsychological behavior via activation of the hypothalamic-pituitary-adrenal (HPA) axis was analyzed using young mice fed a zinc-deficient diet for 2 weeks. Serum corticosterone concentration was significantly increased after 2-week zinc deprivation, whereas zinc concentration in the brain was not decreased. In the resident-intruder test, the rate of mice that exhibited aggressive behavior to the total mice was significantly higher in isolated zinc-deficient mice than in isolated control mice. The duration of aggressive behavior was more in isolated zinc-deficient mice. These results indicate that aggressive behavior of young mice elicited by social isolation is enhanced by zinc deficiency. On the other hand, social isolation-induced aggressive behavior was enhanced in isolated pair-fed mice with food restriction that can activate the HPA axis. Serum corticosterone concentration was also significantly higher in isolated zinc-deficient mice. To see the effect of the increased serum corticosterone on behavioral abnormality, neurotransmitter concentrations in brain tissue were checked. The concentrations of glutamate and GABA in brain tissue were significantly higher in both grouped and isolated zinc-deficient mice. Furthermore, the concentration of extracellular glutamate in the amygdala before the resident-intruder test was significantly higher in isolated zinc-deficient (aggressive) mice and the higher concentration was maintained during the test. The changes in neurotransmitter homeostasis, probably via the increase in serum corticosterone, seem to be linked to aggressive behavior elicited by social isolation in zinc deficiency.