Abstract: Animals that live in stable social groups need to gather information on their own relative position in the group`s social hierarchy, either by directly threatening or challenging others, or in a less costly manner, by observing interactions among others. Such indirect inference of dominance relationships has previously been reported from primates, rats, and birds and fish. Here, we show that domestic horses, Equus caballus, are similarly capable of social cognition. Taking advantage of a specific “following behavior” that horses show towards humans in a riding arena, we investigated whether bystander horses adjust their response to an experimenter according to the observed interaction and their own dominance relationship with the horse whose reaction to the experimenter they had observed before. Horses copied the following behavior towards an experimenter after watching a dominant horse following, but did not follow after observing a subordinate horse or a horse from another social group doing so. The following behavior which horses show towards an experimenter therefore appears to be affected by the demonstrator`s behavior and social status relative to the observer.

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: PENN, D. AND W. K. POTTS. Untrained mice distinguish MHC-determined odors. PHYSIOL BEHAV 64(3) 235-243, 1998.--Immune recognition occurs when foreign antigens are presented to T-lymphocytes by molecules encoded by the highly polymorphic genes of the major histocompatibility complex (MHC). House mice (Mus musculus) prefer to mate with individuals that have dissimilar MHC genes. Numerous studies indicate that mice recognize MHC identity through chemosensory cues; however, it is unclear whether odor is determined by classical, antigen-presenting MHC loci or closely linked genes. Previous studies have relied on training laboratory mice and rats to distinguish MHC-associated odors, but there are several reasons why training experiments may be inappropriate assays for testing if MHC genes affect odor. The aim of this study was to determine whether classical MHC genes affect individual odors and whether wild-derived mice can detect MHC-associated odors without training. In the first experiment, we found that wild-derived mice can be trained in a Y-maze to detect the odors of mice that differ genetically only in the MHC region. In the second and third experiments, we used a naturalistic habituation assay and found that wild-derived mice can, without training, distinguish the odors of mice that differ genetically only at one classical MHC locus (dm2 mutants).

Abstract: Olfaction is a major sensory element in intraspecies recognition and communication in mice. The present study investigated scent marking behaviors of males of the highly inbred C57BL/6J (C57) strain in order to evaluate the ability of these behaviors to provide clear and consistent measures of social familiarity and response to social signals. C57 males engage in scent marking when placed in a chamber with a wire mesh partition separating them from a conspecific. Male mice (C57 or outbred CD-1 mice) showed rapid habituation of scent marking (decreased marking over trials) with repeated exposure at 24-h intervals, to a stimulus animal of the C57 or CD-1 strains, or to an empty chamber. Subsequent exposure to a genetically different novel mouse (CD-1 after CD-1 exposure, or CD-1 after C57 exposure) or to a novel context (different shaped chamber) produced recovery of marking, while responses to a novel but genetically identical mouse (C57 after C57 exposure) or to the empty chamber did not. This finding demonstrated that male mice differentiate familiar and novel conspecifics as expressed by habituation and recovery of scent marking, but neither C57 or CD-1 mice can differentiate new vs. familiar C57 males; likely due to similarities in their odor patterns. The data also indicate that scent marking can differentiate novel from familiar contexts.

Abstract: Newborn rabbits, Oryctolagus cuniculus, discriminate between different categories of adult conspecifics on the basis of their abdominal odour cues. Whether these cues can support the development of filial preferences has not been adequately tested. Using a two-choice paradigm, we assessed the ability of 3-8-day-old pups to orient selectively to the mother versus an unfamiliar female, either spontaneously or after odour conditioning. In experiment 1, nonconditioned pups roamed indifferently over the mother and an unfamiliar female. In experiment 2, pups conditioned to a neutral odorant while nursing or with the mammary pheromone became attracted by the odorant. In experiment 3, pups that had learned the odorant while nursing oriented for longer to any female carrying it, but the unscented mother and a scented unfamiliar female were equally attractive. Finally, in experiment 4, pups that had learned the odorant paired with the mammary pheromone showed a preference for their scented mother, but not systematically for a scented unfamiliar female; furthermore, they were equally attracted by the unscented mother and a scented unfamiliar female. In sum, pups did not spontaneously evince an olfactory preference for the mother when opposed to an unfamiliar female, although they seemed able to detect individual maternal odours. In fact, they appeared to react to both species-specific cues and individual cues that they had learned, and their responses depended on their degree of familiarity with the cues and on the context. The mammary pheromone by itself might act as both a releasing and a reinforcing signal in these early socially oriented behaviours.