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: This article posits that stereotypical behavior patterns and the overall psychological well being of today's performance horse could be substantially enhanced with care that acknowledges the relationship between domesticated horses and their forerunners. Feral horses typically roam in stable, social groups over large grazing territories, spending 16-20 hr per day foraging on mid- to poor-quality roughage. In contrast, today's elite show horses live in relatively small stalls, eat a limited-but rich-diet at specific feedings, and typically live in social isolation. Although the horse has been domesticated for more than 6000 years, there has been no selection for an equid who no longer requires an outlet for these natural behaviors. Using equine stereotypies as a welfare indicator, this researcher proposes that the psychological well being of today's performance horse is compromised. Furthermore, the article illustrates how minimal management changes can enhance horses' well being while still remaining compatible with the requirements of the sport-horse industry. The article discusses conclusions in terms of Fraser, Weary, Pajor, and Milligan's “integrative welfare model” (1997).

Abstract: From observations of intra-specific social grooming in cattle and studies on human stroking in other species, we hypothesised that cows' reactions to human stroking differ depending on the body regions being stroked. Moreover, we tested, whether cows `reactions to stroking change with the animals' experience of stroking. Sixty dairy cows were stroked in three different body regions, i.e. the withers, W, neck ventral, NV (both licked often in social grooming) and the lateral chest, LC (licked rarely), in a balanced order during 10-min sessions. Behavioural reactions and heart rate during stroking as well as reactions to the human just after stroking were recorded. Two test sessions were carried out with 3 weeks of treatment in-between. During this period, the cows were randomly allocated to four treatment groups: three groups received 5 min of daily stroking in either W, NV or LC and the last one (control group) was exposed to simple human presence. During stroking W and NV, cows showed longer neck stretching and ear hanging than during stroking LC (P < 0.001). Moreover, ear hanging was shown longer when W was stroked as compared to NV (P < 0.001), but neck stretching was observed longer during stroking NV as compared to W only after the treatment period (P < 0.01). In the first test session, more animals showed head shaking and head throwing during stroking W and NV than LC (P < 0.01), whereas in the second test session these behaviours were observed only in few animals. In the first test session heart rate was higher during stroking W than the other body regions (P <= 0.01). In the second test session, the lowest heart rate was found during stroking NV (P < 0.05). Finally, contact with the experimenter following stroking differed for the three body regions stroked in both test sessions (first: P = 0.06, second: P < 0.01); contact was shortest for LC. When comparing both test sessions, stretching the neck and ear hanging increased (P < 0.001), while head shaking and head throwing decreased (P < 0.05); there was no difference for heart rate and contact with the experimenter. The four treatments had no influence on reactions to stroking. To conclude, stroking of body regions often licked during social grooming led to more responses than stroking the one licked rarely. Some reactions, such as stretching the neck, are also observed during social licking. This suggests that cows may in part perceive human stroking of body regions often licked similarly to social licking. This knowledge could be of interest for an improvement in quality of human-cattle interactions.

Abstract: The human-animal relationship is an important factor when considering animal welfare at herd level. In the present study, two behavioural tests for the on-farm assessment of the human-animal relationship at herd level of dairy cows housed in loose housing cubicle systems were evaluated with respect to inter-observer reliability, test-retest reliability, effect of familiarity of test person as well as inter-correlation of the two tests. In a voluntary animal approach (VAA) test, the number of cows and the latencies to approach and touch a stationary test person was measured. In an avoidance (AV) test, the cows' avoidance reactions to an approaching test person were categorised. A first study was carried out in 12 commercial Danish and Austrian dairy farms. On each farm, both behaviour tests were carried out on the same day and repeated within 4-5 days. For each test, cows were tested by both an unfamiliar and a familiar test person (the stock-person) and two observers simultaneously registered the animals' test responses. The inter-observer reliability of both behavioural tests was found to be high (VAA: 2.5-m approach r=0.98 (P<0.001) and touch r=0.97 (P<0.001); AV: Kappa coefficientweighted=0.886 (prevalence index for flight distance>=2 m is 0.636)). The cows at herd level showed shortest latency for touching an unfamiliar test person on the first test day (P=0.006). Further, the AV test had a high test-retest reliability (Kappa coefficientweighted=0.503 (prevalence index for flight distance>=2 m is -0.660)) and results indicated no effect of familiarity of test person (Kappa coefficientweighted=0.463 (prevalence index for flight distance>=2 m is -0.677)). In a second study, the correlation between the two behavioural tests (similar measures) was evaluated. On each of 10 commercial Danish dairy farms with loose housing cubicle systems at four repeated sessions, both behaviour tests were carried out on the same day. For each test cows were tested by the stock-person. The VAA and AV tests at herd level were highly correlated (rs=-0.84; P=0.002). The results suggest that the AV test is valid and applicable for on-farm assessment of the human-animal relationship at herd level. This accounts only partly for the VAA test, which seem to be more unclear regarding motivation for the animals' approach behaviour.

Abstract: Learning to associate sensory cues with threats is critical for minimizing aversive experience. The ecological benefit of associative learning relies on accurate perception of predictive cues, but how aversive learning enhances perceptual acuity of sensory signals, particularly in humans, is unclear. We combined multivariate functional magnetic resonance imaging with olfactory psychophysics to show that initially indistinguishable odor enantiomers (mirror-image molecules) become discriminable after aversive conditioning, paralleling the spatial divergence of ensemble activity patterns in primary olfactory (piriform) cortex. Our findings indicate that aversive learning induces piriform plasticity with corresponding gains in odor enantiomer discrimination, underscoring the capacity of fear conditioning to update perceptual representation of predictive cues, over and above its well-recognized role in the acquisition of conditioned responses. That completely indiscriminable sensations can be transformed into discriminable percepts further accentuates the potency of associative learning to enhance sensory cue perception and support adaptive behavior.