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: Lateralization of emotions has received great attention in the last decades, both in humans and animals, but little interest has been given to side bias in perceptual processing. Here, we investigated the influence of the emotional valence of stimuli on visual and olfactory explorations by horses, a large mammalian species with two large monocular visual fields and almost complete decussation of optic fibres. We confronted 38 Arab mares to three objects with either a positive, negative or neutral emotional valence (novel object). The results revealed a gradient of exploration of the 3 objects according to their emotional value and a clear asymmetry in visual exploration. When exploring the novel object, mares used preferentially their right eyes, while they showed a slight tendency to use their left eyes for the negative object. No asymmetry was evidenced for the object with the positive valence. A trend for an asymmetry in olfactory investigation was also observed. Our data confirm the role of the left hemisphere in assessing novelty in horses like in many vertebrate species and the possible role of the right hemisphere in processing negative emotional responses. Our findings also suggest the importance of both hemispheres in the processing positive emotions. This study is, to our knowledge, the first to demonstrate clearly that the emotional valence of a stimulus induces a specific visual lateralization pattern.

Abstract: Among mammals, lactation is the most energy demanding part of parental care and so parentoffspring conflict should arise over milk provided by the mother. Mother and offspring should disagree over the length and amount of the milk provision. We focused on effect of pregnancy on suckling behaviour variables as indicators of mother-offspring conflict in domestic horses. We presumed shorter suckling bouts and higher rates of rejected and/or terminated suckling in pregnant mares compared to non-pregnant ones. Increasing conflict over amount of maternal investment between mother and her young are to be expected because of her parallel investment into a nursed foal and a foetus. Eight groups of loose housed lactating mares with foals of Kladruby horse were studied at the National Stud Kladruby nad Labem (Czech Republic) from deliveries to abrupt weaning (at the age of 127 to 210 days). We recorded 10 848 suckling solicitations of 79 mare-foal pairs, from which 10 607 resulted in a suckling bout. In 41 cases a nursing mare became pregnant during lactation. We found no significant effect of pregnancy either on probability of the mother rejecting suckling solicitation of her foal or probability that she terminated a suckling bout. However the overall effect of mother’s pregnancy on suckling bout duration was not significant, there were considerable differences in pregnant and non-pregnant mares according to who terminated a suckling bout, whether the mother or the foal (F(1, 9776) = 12.1, P < 0.001). In case it was the mother then the suckling bout was longer if she was pregnant (65.36 ± 1.25 s) than barren (60.55 ± 1.36 s). We found no impact of pregnancy on duration of suckling bouts terminated by the foal. Further, nursing a foal during the first two trimesters of pregnancy had no negative impact on birth weight of the foetus. In conclusion, we found not higher, but a lower mother-offspring conflict in pregnant than in non-pregnant lactating mares while expecting just the opposite. We suggest that pregnant mares compensate their nursed foals during intensive stages of lactation through a relaxed mother-offspring conflict for later decrease in investment due to increasing demands of the foetus and/or for the shorter period of milk supply. Our results (partly published in Bartosova et al. 2011, PLoS ONE 6(8): e22068) are of high importance in horse breeding. One of the main arguments for early weaning of the foals is regeneration of their pregnant mothers before upcoming delivery. Here we present evidence that a pregnant mare “counts” with her dual maternal investment and “employs” evolutionary mechanisms enabling her to rear a vital foetus. From this point of view there is no objective reason for stressful weaning of her nursed offspring practised in conventional breeding. Supported by AWIN, EU FP7 project No. 266213.

Abstract: Six pregnant mares were used to determine what level of water restriction causes physiological and/or behavioural changes indicative of stress. Nonlegume hay was fed ad libitum. During the first week of restriction, 5 l water/100 kg bwt was available, during the second week 4 l/100 kg bwt and, during the third week, 3 l/100 kg bwt. Ad libitum water intake was 6.9 l/100 kg bwt; at 3 l/100 kg bwt water intake was 42% of this. Daily hay intake fell significantly with increasing water restriction from 12.9 +/- 0.75 kg to 8.3 +/- 0.54 kg; bodyweight fell significantly for a total loss of 48.5 +/- 8.3 kg in 3 weeks. Daily blood samples were analysed; osmolality rose significantly with increasing water restriction from 282 +/- 0.7 mosmols/kg to 293.3 +/- 0.8 mosmols/kg bwt, but plasma protein and PCV did not change significantly. Cortisol concentrations fell from 8.1 ng/ml to 6.4 ng/ml over the 3 week period. Aldosterone fell from 211.3 +/- 74.2 pg/ml to 92.5 +/- 27.5 pg/ml at the end of the first week. The behaviour of 4 of the 6 mares was recorded 24 h/day for the duration of the study. The only significant difference was in time spent eating, which decreased with increasing water restriction from 46 +/- 3% to 30 +/- 3%. It is concluded that water restriction to 4 l/100 kg bwt dehydrates pregnant mares and may diminish their welfare, but is not life- or pregnancy-threatening.