Abstract: Whereas mammalian mothers and young may retain long-term social affiliations in nature, the management of animals in captivity typically dictates that offspring are abruptly and permanently separated from their mothers at a relatively early age, often prior to the time of natural weaning. For animal breeders, this strategy can enhance the yield of offspring from a breeding population. Morbidity and mortality can also precipitate severance of mother-young bonds. Although it is recognized that early weaning provides nutritional challenges for the young, relatively little attention has been paid to the psychological consequences and long-term impacts of breaking the mother-young bond in non-human mammals. Furthermore, whereas great strides are being made in our understanding of the neurobiological and genetic underpinnings of social bonding, the mechanisms underlying the process of detachment following establishment of a mother-young bond remain relatively unexplored, although parallels can be drawn with processes involved in withdrawal from addictive substances. In this review, we outline mechanisms involved in social bonding. We consider the diversity in extent and duration of mother-young attachment across mammalian lineages and implications for predicting the outcome of severing ties between mothers and young at different times post-partum. We identify characteristics signalling emotional distress resulting from separation of mothers and young and discuss strategies for mitigating separation-induced distress. These include postponement of separation, ensuring high-quality maternal care of young prior to separation, providing bonded individuals with opportunities to separate voluntarily for brief periods prior to permanent separation, use of anti-suck devices prior to separation, allowing a period of partial (fence line) contact prior to full separation, providing substitutes for stimuli previously exchanged between mother and young, providing social buffers, gradual introduction to new housing arrangements, and pharmacological intervention. Areas for future research are proposed, including the use of functional neuroimaging technologies and functional genomics approaches, in combination with behavioural assessments of reinstatement motivation, individual recognition memory and long-term consequences of early separation, to shed further light on the nature of mother-young bonding and detachment in animals.

Abstract: Four methods of ranking horses on reactivity were evaluated and compared: isolation from conspecifics, presentation of a static novel stimulus, traversing a novel stimulus in a runway (isolation, novel stimulus and runway tests, respectively) and assigning subjective emotionality scores. In all tests, horses' heart rates were recorded and behaviour was videotaped. To be considered a valid test of reactivity, at least one heart rate and one behavioural measurement in the test had to change significantly between treatments (tranquilizer administation versus sham tranquilizer administration), and behavioural measures had to be displayed in at least 75% of the trials. Forty horses performed each of the three tests daily on three different days in a switchback design. Horses were assigned randomly to a daily test sequence, which was maintained throughout the study. In the runway test, no significant difference in heart rate values in tranquilized and non-tranquilized horses was found, and no behavioural attribute was displayed in more than 52% of the trials; therefore it was rejected as a valid test of reactivity. Both isolation and novel stimulus tests produced valid measurements. Mean heart rate was the most precise physiological measure for these tests, and walking and defecation frequency were the most precise behavioural measures for novel stimulus and isolation tests, respectively. Mean heart rates on the novel stimulus and isolation tests were correlated (rs = 0.79, P < 0.01) indicating that these tests produced similar rankings based on physiological responses. However, behavioural measures ranked horses differently (rs = 0.27, P < 0.10) on the tests. Rank correlations between mean heart rates and behavioural measures were higher in the novel stimulus (rs = 0.66, P < 0.01) than the isolation test (rs = 0.55, P < 0.01), indicating that the novel stimulus test ranked horses based on either physiological or behavioural responses more similarly than did the isolation test. Therefore, the novel stimulus test was considered the more accurate evaluation of reactivity. Subjective emotionality scores were correlated moderately with mean heart rates (rs > 0.33, P < 0.01) from the novel stimulus and isolation tests and with walking scores (rs = 0.47, P < 0.01) from the novel stimulus test. Assignment of subjective emotionality scores was not as accurate as the novel stimulus or isolation tests in ranking horses for reactivity. Using physiological data alone, combining physiological and behavioural measurements or using more than one behavioural measurement in reactivity tests may reflect the reactivity of the horse better than a single behavioural measurement.

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: The horse's temperament, including its manageability and reactivity and/or fearfulness, is of importance as it can result in problems and can render horses unsuitable for inexperienced riders. Early experience, including handling during infancy, may influence the horse's adult behaviour and reduce its fear of humans and other potentially frigthening situations. In the various species studied, handling has generally been undertaken during the neonatal period. The aim of the present study was to test the effects of handling young horses around the time of weaning, a period which has been demonstrated to be effective in increasing ease of handling in cattle and goats. Sixteen Anglo-Arab foals were handled for 12 days either immediately following weaning (early handled: EH) or 21 days later (late handled: LH); eight additional non-handled foals served as controls (C). Handling consisted of haltering, gently petting all parts of the body, picking up feet and leading the foal over 120 m. During handling sessions, EH were easier to handle than LH: time taken to fit them with a halter, to pick up feet, and “walk-ratio” (time walking under constraint/total time walking) were significantly lower for EH. During subsequent tests conducted over 2 days, 4, and 7 months, as well as 10 months and to some extent 18 months after the end of handling period, EH and LH were easier to handle and less reactive than controls, although differences diminished with time. The period following weaning can therefore be qualified as an “optimal period” for handling. Some of the effects persist for at least 18 months.

Abstract: Handling is a crucial component of the human-horse relationship. Here, we report data from an experiment conducted to assess and compare the effect of two training methods. Two groups of six Welsh mares were trained during four sessions of 50 min, one handled with traditional exercises (halter leading, grooming/brushing, lifting feet, lunging and pseudo-saddling (using only girth and saddle pad) and the second group with natural horsemanship exercises (desensitization, yielding to body pressure, lunging and free-lunging). Emotional reactivity (ER) and the human-horse relationship (HHR) were assessed both prior to and following handling. A social isolation test, a neophobia test and a bridge test were used to assess ER. HHR was assessed through test of spontaneous approach to, and forced approach by, an unknown human. Horses' ER decreased after both types of handling as indicated by decreases in the occurrence of whinnying during stressful situations. Head movement (jerk/shake) was the most sensitive variable to handling type. In the spontaneous approach tests, horses in the traditional handling group showed higher latencies to approach a motionless person after handling than did the natural horsemanship group. Our study suggests that natural horsemanship exercises could be more efficient than traditional exercises for improving horses' HHR.