Maintenance Of Social Behavior Research Articles

Oxytocin receptors influence the development and maintenance of social behavior in zebrafish (Danio rerio)

Zebrafish are highly social teleost fish and an excellent model to study social behavior. The neuropeptide Oxytocin is associated different social behaviors as well as disorders resulting in social impairment like autism spectrum disorder. However, how Oxytocin receptor signaling affects the development and expression kinetics of social behavior is not known. In this study we investigated the role of the two oxytocin receptors, Oxtr and Oxtrl, in the development and maintenance of social preference and shoaling behavior in 2- to 8-week-old zebrafish. Using CRISPR/Cas9 mediated oxtr and oxtrl knock-out fish, we found that the development of social preference is accelerated if one of the Oxytocin receptors is knocked-out and that the knock-out fish reach significantly higher levels of social preference. Moreover, oxtr−/− fish showed impairments in the maintenance of social preference. Social isolation prior to testing led to impaired maintenance of social preference in both wild-type and oxtr and oxtrl knock-out fish. Knocking-out either of the Oxytocin receptors also led to increased group spacing and reduced polarization in a 20-fish shoal at 8 weeks post fertilization, but not at 4. These results show that the development and maintenance of social behavior is influenced by the Oxytocin receptors and that the effects are not just pro- or antisocial, but dependent on both the age and social context of the fish.

Venom and Social Behavior: The Potential of Using Spiders to Evaluate the Evolution of Sociality under High Risk.

Risks of sociality, including competition and conspecific aggression, are particularly pronounced in venomous invertebrates such as arachnids. Spiders show a wide range of sociality, with differing levels of cannibalism and other types of social aggression. To have the greatest chance of surviving interactions with conspecifics, spiders must learn to assess and respond to risk. One of the major ways risk assessment is studied in spiders is via venom metering, in which spiders choose how much venom to use based on prey and predator characteristics. While venom metering in response to prey acquisition and predator defense is well-studied, less is known about its use in conspecific interactions. Here we argue that due to the wide range of both sociality and venom found in spiders, they are poised to be an excellent system for testing questions regarding whether and how venom use relates to the evolution of social behavior and, in return, whether social behavior influences venom use and evolution. We focus primarily on the widow spiders, Latrodectus, as a strong model for testing these hypotheses. Given that successful responses to risk are vital for maintaining sociality, comparative analysis of spider taxa in which venom metering and sociality vary can provide valuable insights into the evolution and maintenance of social behavior under risk.

Variable ecological conditions promote male helping by changing banded mongoose group composition

Ecological conditions are expected to have an important influence on individuals' investment in cooperative care. However, the nature of their effects is unclear: both favorable and unfavorable conditions have been found to promote helping behavior. Recent studies provide a possible explanation for these conflicting results by suggesting that increased ecological variability, rather than changes in mean conditions, promote cooperative care. However, no study has tested whether increased ecological variability promotes individual-level helping behavior or the mechanisms involved. We test this hypothesis in a long-term study population of the cooperatively breeding banded mongoose, Mungos mungo, using 14 years of behavioral and meteorological data to explore how the mean and variability of ecological conditions influence individual behavior, body condition, and survival. Female body condition was more sensitive to changes in rainfall leading to poorer female survival and pronounced male-biased group compositions after periods of high rainfall variability. After such periods, older males invested more in helping behavior, potentially because they had fewer mating opportunities. These results provide the first empirical evidence for increased individual helping effort in more variable ecological conditions and suggest this arises because of individual differences in the effect of ecological conditions on body condition and survival, and the knock-on effect on social group composition. Individual differences in sensitivity to environmental variability, and the impacts this has on the internal structure and composition of animal groups, can exert a strong influence on the evolution and maintenance of social behaviors, such as cooperative care.

A system for automatic recording of social behavior in a free-living wild house mouse population

Our research focuses on mechanisms that promote and stabilize social behavior, fitness consequences of cooperation, and how interactions with conspecifics structure groups and populations. To this end, we studied wild house mice (Mus musculus domesticus) in the laboratory, in semi-natural enclosures and in the field. In 2002, we initiated a project on a free-living population of house mice in a barn near Zurich, Switzerland, where mice were equipped with RFID transponders and were provided with 40 nest boxes for resting and breeding. The population typically comprised between 250 and 400 mice. To analyze social group membership, social interactions and social preferences of the mice in our study population over their lifespan, we installed a continuous transponder reading system (AniLoc, FBI Science GmbH, Germany). Mice accessed nest boxes through tunnels equipped with two antennas each. When a mouse implanted with an RFID transponder passed the electromagnetic field of an antenna, its identity was transmitted and registered in real time with AniLoc. Additionally, body weights of mice were automatically registered at eight drinking facilities (Intelliscale, FBI Science GmbH, Germany). Here, a mouse sits on a freely movable platform that connects to a scale registering body weight when drinking, and an antenna around the head of the water bottle registers the drinking individual’s RFID transponder. The system enabled continuous remote monitoring of the behavior of a free-living, open population of house mice, when using nest boxes and when drinking. Since such safe places are an important resource for survival and reproduction, time of day, duration and frequency of meetings with conspecifics reveal information about the function of their interactions. Trigger efficiency of antennas was 98.2 %. Mice entered and left the nest boxes with an average speed of 0.03 m/s, which is within the antennas’ detection capacity (detection speed of 1 m/s or 3.6 km/h). The antenna devices documented not only social structuring of our study population but also spatial genetic structuring. The observation that mice lived in rather closed social groups and tended to share nest boxes with relatives highlights the importance of kin selection for the evolution and maintenance of social behavior. We suggest that such automatic recording of activity, spatial distribution and social interactions is helpful not only in field studies, for a variety of species, but also in captivity or laboratory studies, to answer basic questions in behavioral ecology, population ecology, population genetics, conservation biology, disease ecology, or animal welfare.

Social evolution and inclusive fitness theory: an introduction

Social behavior has long puzzled evolutionary biologists, since the classical theory of natural selection maintains that individuals should not sacrifice their own fitness to affect that of others. Social Evolution and Inclusive Fitness Theory argues that a theory first presented in 1963 by William D. Hamilton—inclusive fitness theory—provides the most fundamental and general explanation for the evolution and maintenance of social behavior in the natural world. James Marshall guides readers through the vast and confusing literature on the evolution of social behavior, introducing and explaining the competing theories that claim to provide answers to questions such as why animals evolve to behave altruistically. Using simple statistical language and techniques that practicing biologists will be familiar with, he provides a comprehensive yet easily understandable treatment of key concepts and their repeated misinterpretations. Particular attention is paid to how more realistic features of behavior, such as nonadditivity and conditionality, can complicate analysis. Marshall highlights the general problem of identifying the underlying causes of evolutionary change, and proposes fruitful approaches to doing so in the study of social evolution. Social Evolution and Inclusive Fitness Theory describes how inclusive fitness theory addresses both simple and complex social scenarios, the controversies surrounding the theory, and how experimental work supports the theory as the most powerful explanation for social behavior and its evolution.

Levels of social behaviors and genetic structure in a population of round-tailed ground squirrels (Xerospermophilus tereticaudus)

Ground-dwelling sciurids exhibit a continuum of sociality and several models predict levels of sociality within this taxon. Models of ground squirrel sociality predict round-tailed ground squirrels (Xerospermophilus tereticaudus) to be solitary; however, previous behavioral studies suggest round-tailed ground squirrels have a matrilineal social structure. To resolve this discrepancy, we combined behavioral observations with genetic analyses of population structure. We assessed levels of agonistic and amicable behaviors combined with fine-scale population genetic structure of round-tailed ground squirrels in a multi-year study in AZ. Only 45 agonistic and 40 amicable interactions were observed between adults in over 137 h of observations. Overall rates of agonistic or amicable interactions between adults were low (≤0.69/h), with no relationship between relatedness of individuals and rates of either amicable or agonistic interactions. Interactions between juvenile littermates were predominantly amicable. Population substructure was not evident with Bayesian analyses, global or pairwise F ST values; average relatedness among females was not different from males. However, in 2006, the year after a population reduction through targeted animal elimination, a population bottleneck was detected within at least five of seven loci. Contrary to previous behavioral studies, this population of round-tailed ground squirrels, although aggregated spatially, did not exhibit high levels of social behavior nor subpopulation genetic structure. Analyses of the genetic relationships and sociality along a continuum, particularly within aggregates of individuals, may lead to insights into the origin and maintenance of social behaviors by elucidating the mechanisms by which aggregates with intermediate social levels are formed and maintained.

Cuticular hydrocarbons as potential kin recognition cues in a subsocial spider

In animal societies, recognition of group members and relatives is an important trait for the evolution and maintenance of social behavior. In eusocial insects, nest mate recognition is based on cuticular hydrocarbons and allows colony members to reject competitors and parasites. The study of recognition cues in subsocial species can provide insights into evolutionary pathways leading to permanent sociality and kin-selected benefits of cooperation. In subsocial spiders, empirical evidence suggests the existence of both kin recognition and benefits of cooperating with kin, whereas the cues for kin recognition have yet to be identified. However, cuticular hydrocarbons have been proposed to be involved in regulation of tolerance and interattraction in spider sociality. Here, we show that subsocial Stegodyphus lineatus spiderlings have cuticular hydrocarbon profiles that are sibling-group specific, making cuticular hydrocarbons candidates for kin recognition cues. Our behavioral assays indicate that spiderlings can discriminate between cuticular cues from kin and nonkin: In a choice set-up, spiderlings more often chose to reside near cuticular chemical extracts of siblings compared with nonsiblings. Furthermore, we show that cuticular chemical composition changes during development, especially around the stage of dispersal, supporting the hypothesis that cuticular cues are involved in regulating conspecific tolerance levels. Lastly, our results indicate that the potential kin recognition cues might be branched alkanes that are influenced very little by rearing conditions and may be genetically determined. This indicates that a specific group of cuticular chemicals, namely branched alkanes, could have evolved to act as social recognition cues in both insects and spiders

Examining potential benefits of group living in a sawfly larva, Perga affinis

Although defense may serve as a primary selective force behind the evolution and maintenance of social behavior, numerous other benefits play a role as well. Larvae of the Australian sawfly, Perga affinis, live in colonies and retain their gregarious lifestyle through pupation. To evaluate selective pressures maintaining social behavior in P. affinis, I investigated potential benefits of group living. These included predation protection, immune function, thermoregulation, foraging facilitation, and pupation success where I compared treatments between individuals versus groups and between groups of different sizes. I found no evidence of predation on larvae; however, the mortality risk was significantly higher for single versus grouped larvae, suggesting that other grouping benefits are important. Immune function was not a benefit as hemolymph from single versus grouped larvae did not differ in its ability to limit bacterial growth. Investigating thermoregulation revealed that grouped larvae attained significantly higher temperatures than single individuals and that large groups reached higher temperatures than small groups. Larvae grew significantly faster at higher temperatures, indicating that the increased heat absorption and retention capacity of large groups may speed up development. Group size did not affect foraging facilitation (measured via weight gain) except during the last time period, where individuals in large groups gained significantly more weight. Weight gain in the last instar may be critical for pupation as high larval weight significantly increased the odds of pupation success. Thus, the group size effect on weight gain during the last instar provides an adaptive explanation for the observed ontogenetic increase in colony size and coalescence. Key words: group living, growth rate, group size, larvae, ontogenic shift, thermoregulation. [Behav Ecol]

Learned kin recognition cues in a social bird

In many cooperatively breeding birds, kin selection has an important role in the evolution and maintenance of social behaviour, and 'helpers' can maximize indirect fitness gains by preferentially allocating care to close relatives. Although there is evidence for kin-biased helping behaviour in several species, the mechanism of kin recognition underlying this behaviour is poorly understood. Vocalizations are the most commonly used cues in avian recognition systems, but the effectiveness of vocal signals as reliable recognition cues must depend on how they are acquired. However, there have been no experimental studies of the development of vocal recognition cues in cooperative birds; indeed, the ontogeny of all bird vocalizations other than song is poorly known in any species. Here, we show that cooperatively breeding long-tailed tits (Aegithalos caudatus) can discriminate between kin and non-kin according to the individual-specific characteristics of contact calls, and show experimentally that individuals learn these calls from provisioning adults during the nestling period. Finally, we show that the pattern of cooperative behaviour in this species is consistent with the use of recognition cues learned through association.

Reduced worker relatedness does not affect cooperation in honey bee colonies

Although kin selection is commonly regarded as a major force in the evolution and maintenance of social behaviour in eusocial insects, recent controversy regarding whether honey bee (Apis mellifera L., 1758) workers can discriminate close kin from more distant relatives casts doubt on the extent to which cooperation among individuals within honey bee colonies is controlled by genetic relatedness. We contrasted brood and honey production in colonies where we diluted worker relatedness with those elements of productivity in colonies where relatedness was unmanipulated. Relatedness manipulation did not affect overall brood production, worker or drone brood production, the worker to drone brood sex ratio, or the volume of honey produced. Thus, there is no evidence that honey bees discriminate close from distant relatives or, more importantly, that dilution of the coefficient of relatedness within a colony has any impact on the efficiency of that colony.

On the relative importance of haplodiploidy, assortative mating and social synergy on the evolutionary emergence of social behavior.

Advances in multiagent simulation techniques make it possible to study more realistic dynamics of complex systems and allow evolutionary theories to be tested. Here I use simulations to assess the relative importance of reproductive systems (haplodiploidy vs. diploidy), mate selection (assortative mating vs. random mating) and social economics (pay-off matrices of evolutionary games) in the evolutionary dynamics leading to the emergence of social cooperation in the provision of parental care. The simulations confirm that haplo-diploid organisms and organisms mating assortatively have a higher probability for fixing alleles and require less favorable conditions for their fixation, than diploids or organisms mating randomly. The simulations showed that social behavior was most likely to emerge a) when the cost for parental investment was much lower than the benefits to the offspring, b) when cooperation improved synergistically the fitness of offspring compared to the corresponding egoistic behavior and c) when alleles coding for altruistic or social behavior could be rapidly fixed in the population, thanks to mechanisms such as haplo-diploidy and/or assortative mating. Cooperative social behavior always appeared if sociality conferred much higher fitness gains compared to non cooperative alternatives suggesting that the most important factors for the emergence and maintenance of social behavior are those based on energetic or efficiency considerations. The simulations, in congruence with the scant experimental evidence available, suggest that economic considerations rather than genetic ones are critical in explaining the emergence and maintenance of sociality.

Reproductive caste determination in eusocial wasps (Hymenoptera: Vespidae).

Wasps (Vespidae) exhibit a range of social complexity, from solitary living to eusocial colonies, and thus are exemplary for studies of the evolutionary origin and maintenance of social behavior in animals. Integral to the definition of eusociality is the presence of reproductive castes, group members that differ qualitatively in their ability to reproduce in a social setting. Behavioral and morphological evidence suggests that caste determination, the developmental process by which differences in fecundity are established, occurs to a large extent before adult emergence (pre-imaginally) in many species of Vespidae, in both basal and advanced taxa within the clade (Vespinae+Polistinae), which includes most eusocial species. Pre-imaginal determination has been documented in many taxa (e.g. independent-founding Polistinae) where it was not thought to occur. Correlative and experimental studies indicate that differences in nutrition during larval development are often the basis of pre-imaginal caste determination. Pre-imaginal caste determination has important implications for the roles of subfertility and manipulation by nest mates in the evolution of eusocial behavior.

How parasites can promote the expression of social behaviour in their hosts

Recent theory on the role of parasites in the evolution of social behaviour has emphasized the costs of social behaviour to hosts. However, parasites whose primary effect on host fitness is to reduce fecundity can favour the evolutionary origin or maintenance of social behaviour, including eusociality, under certain conditions. If the parasites are not readily transmitted among members of social groups, then other group members will not be selected to reject infected individuals as social partners, nor will adaptive suicide or avoidance of grouping be selectively favoured for infected individuals. Rather, total or partial parasitic castration may promote the expression of helping behaviour by infected individuals. Some parasites may therefore act to increase variance in direct reproductive value within populations or societies, and to promote reproductive division of labour. The necessary conditions of reduced host fecundity and low within-group transmission are met in some insect-parasite systems, and could occur in other host-parasite systems as well.

Generalization and maintenance of preschool children's social skills: a critical review and analysis.

This paper summarizes the results of a retrospective review of generalization in the context of social skills research with preschool children. A review of studies from 22 journals (1976 to 1990) that assessed generalization as part of social interaction research provided information concerning the prevalence of studies that have assessed generalization, common practices concerning the production and assessment of generalization, and the overall success of obtaining generalization and maintenance of social behaviors. A comparison of the most and least successful studies, with respect to generalization, revealed some differences concerning the practices employed by studies within each group. Differences differentially related to the production of generalization are discussed and recommendations are provided to guide and support future research efforts.

Philopatry and nest founding in a primitively social bee, Halictus rubicundus

Observations of inter- and intraseasonal dispersal patterns in the primitively social sweat bee Halictus rubicundus in New York reveal considerable philopatry in both sexes. Females overwinter away from the nest aggregation, in diapause for 8–11 months, and return to dig new nests close to the site of their natal nests (typically within 50 cm). Nests are all founded by siggle females (haplometrosis). Clumping of nests may reflect patterns of soil vegetation. Workers rarely enter or take up residence in non-natal nests. Males commonly return to both natal and non-natal nests, and actively patrol vegetation near the natal aggregation for females. These philopatric tendencies, in combination with the haplometrotic mode of nest founding, should have important effects on population structure, particularly in terms of population subdivision, mate choice, and intracolony interactions. The resulting population structure may be conducive to the evolution and maintenance of social behavior.

Measuring genetic relatedness in natural populations: Methodology

The estimation of relatedness within social groups, such as the colonies of a population of social insects, is an important field for evaluating hypotheses concerning the evolution and maintenance of social behaviour. The methodology of this estimation from genetic data in the absence of pedigree information has been poorly understood; we develop this methodology for b, the regression coefficient of relatedness, and discuss its applications. Both b and G (the pedigree coefficient of relatedness) are potentially asymmetric coefficients, whereas φ, r, and F ST are necessarily symmetric. We develop an estimator for b suitable for small samples, and also one for standard deviation, and examine the properties of both using sampling simulations. The b estimator returns values slightly below E( b), and the standard deviation estimator yields conservative confidence intervals. A comparative study of b and F ST shows that, given the same set of data, b is estimated with greater reliability than is F ST . As is the case for F ST , b can be used to examine population structure at various levels, and b possesses the advantage of an estimator for its standard error, which can also be used to test for heterogeneity among the loci surveyed. The actual numbers of identical genes held in common by interacting individuals, and not simply their proportions, need to be considered in using coefficients of relatedness in inclusive fitness calculations. This necessity is handled by the weighted coefficients of relatedness, G′ and b′, which have been referred to in the literature as r (as have most relatedness measures).

Effects of hippocampal lesion on cardiovascular, adrenocortical and behavioral response patterns in mice

It was found that animals with a hippocampal lesion developed high blood pressure, low heart rates, and high plasma corticosterone during social interaction in a territorial situation. However, hippocampal animals kept in a nonterritorial situation with minimal social interaction did not show significant cardiovascular or plasma corticosterone changes as compared to unoperated controls or cortically lesioned controls. The socially interacting hippocampal animals failed to develop a social hierarchy and did not respond aggressively to an intruder placed in the colony. The data suggests that the hippocampus is involved in the maintenance of social behavior which in turn may modulate autonomic nervous system activity.