Abstract: Many mammalian species are highly social, creating intra-group competition for such things as food and mates. Recent research with nonhuman primates indicates that in competitive situations individuals know what other individuals can and cannot see, and they use this knowledge to their advantage in various ways. In the current study, we extended these findings to a non-primate species, the domestic goat, using the conspecific competition paradigm developed by Hare et al. (2000). Like chimpanzees and some other nonhuman primates, goats live in fission-fusion societies, form coalitions and alliances, and are known to reconcile after fights. In the current study, a dominant and a subordinate individual competed for food, but in some cases the subordinate could see things that the dominant could not. In the condition where dominants could only see one piece of food but subordinates could see both, subordinates' preferences depended on whether they received aggression from the dominant animal during the experiment. Subjects who received aggression preferred the hidden over the visible piece of food, whereas subjects who never received aggression significantly preferred the visible piece. By using this strategy, goats who had not received aggression got significantly more food than the other goats. Such complex social interactions may be supported by cognitive mechanisms similar to those of chimpanzees. We discuss these results in the context of current issues in mammalian cognition and socio-ecology.

Abstract: Mammalian social systems rely on signals passed between individuals conveying information including sex, reproductive status, individual identity, ownership, competitive ability and health status. Many of these signals take the form of complex mixtures of molecules sensed by chemosensory systems and have important influences on a variety of behaviours that are vital for reproductive success, such as parent-offspring attachment, mate choice and territorial marking. This article aims to review the nature of these chemosensory cues and the neural pathways mediating their physiological and behavioural effects. Despite the complexities of mammalian societies, there are instances where single molecules can act as classical pheromones attracting interest and approach behaviour. Chemosignals with relatively high volatility can be used to signal at a distance and are sensed by the main olfactory system. Most mammals also possess a vomeronasal system, which is specialized to detect relatively non-volatile chemosensory cues following direct contact. Single attractant molecules are sensed by highly specific receptors using a labelled line pathway. These act alongside more complex mixtures of signals that are required to signal individual identity. There are multiple sources of such individuality chemosignals, based on the highly polymorphic genes of the major histocompatibility complex (MHC) or lipocalins such as the mouse major urinary proteins. The individual profile of volatile components that make up an individual odour signature can be sensed by the main olfactory system, as the pattern of activity across an array of broadly tuned receptor types. In addition, the vomeronasal system can respond highly selectively to non-volatile peptide ligands associated with the MHC, acting at the V2r class of vomeronasal receptor.The ability to recognize individuals or their genetic relatedness plays an important role in mammalian social behaviour. Thus robust systems for olfactory learning and recognition of chemosensory individuality have evolved, often associated with major life events, such as mating, parturition or neonatal development. These forms of learning share common features, such as increased noradrenaline evoked by somatosensory stimulation, which results in neural changes at the level of the olfactory bulb. In the main olfactory bulb, these changes are likely to refine the pattern of activity in response to the learned odour, enhancing its discrimination from those of similar odours. In the accessory olfactory bulb, memory formation is hypothesized to involve a selective inhibition, which disrupts the transmission of the learned chemosignal from the mating male. Information from the main olfactory and vomeronasal systems is integrated at the level of the corticomedial amygdala, which forms the most important pathway by which social odours mediate their behavioural and physiological effects. Recent evidence suggests that this region may also play an important role in the learning and recognition of social chemosignals.

Abstract: Parent-offspring communication has evolved under strong selection to guarantee that the valuable resource of parental care is expended efficiently on raising offspring. To ensure allocation of parental care to their own offspring, individual recognition becomes established in higher vertebrates when the young become mobile at a time when a nest site can no longer provide a safe cue to recognition. Such recognition needs to be established by rapid, sometimes imprinting-like, processes in animals producing precocial offspring. In parents, offering strategies that stimulate feeding and entice offspring to approach the right site have evolved. Such parental signals can be olfactory, acoustic, or visual. In offspring, begging strategies involve shuffling for the best place to obtain food – be this the most productive teat or the best position in the nest. This involves signals that make the offspring particularly obvious to the parent. Parents often feed young according to their signaling intensity but may also show favoritism for weaker offspring. Offspring signals also serve to communicate the continuing presence of the young and may thereby maintain brood-care behavior in parents. Internal processes in parents may end parental care irrespective of further signaling by offspring, thus ensuring that offspring cannot manipulate parents into providing substantially more care than is optimal for their own fitness.

Abstract: Social relationships are essential for maintaining human mental health, yet little is known about the brain mechanisms involved in the development and maintenance of social bonds. Animal models are powerful tools for investigating the neurobiological mechanisms regulating the cognitive processes leading to the development of social relationships and for potentially extending our understanding of the human condition. In this review, we discuss the roles of the neuropeptides oxytocin and vasopressin in the regulation of social bonding as well as related social behaviors which culminate in the formation of social relationships in animal models. The formation of social bonds is a hierarchical process involving social motivation and approach, the processing of social stimuli and formation of social memories, and the social attachment itself. Oxytocin and vasopressin have been implicated in each of these processes. Specifically, these peptides facilitate social affiliation and parental nurturing behavior, are essential for social recognition in rodents, and are involved in the formation of selective mother-infant bonds in sheep and pair bonds in monogamous voles. The convergence of evidence from these animal studies makes oxytocin and vasopressin attractive candidates for the neural modulation of human social relationships as well as potential therapeutic targets for the treatment of psychiatric disorders associated with disruptions in social behavior, including autism.

Abstract: Transitive inference (TI) involves using known relationships to deduce unknown ones (for example, using A > B and B > C to infer A > C), and is thus essential to logical reasoning. First described as a developmental milestone in children, TI has since been reported in nonhuman primates, rats and birds. Still, how animals acquire and represent transitive relationships and why such abilities might have evolved remain open problems. Here we show that male fish (Astatotilapia burtoni) can successfully make inferences on a hierarchy implied by pairwise fights between rival males. These fish learned the implied hierarchy vicariously (as 'bystanders'), by watching fights between rivals arranged around them in separate tank units. Our findings show that fish use TI when trained on socially relevant stimuli, and that they can make such inferences by using indirect information alone. Further, these bystanders seem to have both spatial and featural representations related to rival abilities, which they can use to make correct inferences depending on what kind of information is available to them. Beyond extending TI to fish and experimentally demonstrating indirect TI learning in animals, these results indicate that a universal mechanism underlying TI is unlikely. Rather, animals probably use multiple domain-specific representations adapted to different social and ecological pressures that they encounter during the course of their natural lives.