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: Weaving is a common locomotor stereotypy in horses, which has been shown to be reduced by the presence of a mirror in the stable; however, it remains unclear whether this effect is due to the reflected image of a horse or some other property associated with the introduction of a mirror. To investigate this further, the response of six known weaving horses to each of three different types of poster was recorded. The poster images consisted of a life-size image of a horse's face (true image), a version of the same image cut into 54 squares, which were then randomly rearranged (pixilated image) and a blank (white image) display of the same size as the other posters. Four observation periods were included in each of the two days of observation per treatment condition: 08:00-08:30 h, 10:00-10:30 h, 12:00-12:30 h and 16:00-17:00 h. The first observation period was followed by the provision of concentrate feed and forage, the second observation period by exercise, the third by further forage and the fourth by concentrate feed and forage. The horses' activities and positions in the box were scanned at 1 min intervals and the number of observations registering each activity and position was calculated as a percentage of the total number of observations for each observation period. Weaving was significantly less when the horses were provided with the image of a horse's face (mean percentage of observations +/- S.E.M.; 5.56 +/- 1.57), compared to both the pixilated (14.85 +/- 3.06) and white (20.52 +/- 4.12) images. Nodding was significantly less when provided with the true image of a horse (0.45 +/- 0.15) compared to the pixilated image (2.15 +/- 0.67). The pattern of weaving throughout the day differed from that of nodding, and together these results suggest that the two behaviours are not analogous. The presence of the image of the horse's face was associated with a significant increase in alertness (F112,2 = 11.31, p < 0.001) and recorded time spent looking at the poster(F112,2 = 3.46, p < 0.05). The timing of stereotypic head movements in the horse in this study suggests that they are not associated with boredom, or lack of stimulation but rather acute frustration.

Abstract: Weaning foals marks a stressful event in horses' lives. Limited research exists regarding different housing methods post-weaning and the long-term implications on horse behavior and welfare. The purpose of this study was to monitor behavior and physiological stress markers in horses weaned individually in solid partition box stalls versus horses weaned in small groups and housed in paddocks. Both treatment groups underwent maternal deprivation stress, but the stalled weanlings had the additive effects of social isolation which prevented them from performing social behaviors. Quarter Horse weanlings from the Michigan State University, Merillat Equine Center, average age 4.5 months, were weaned in 13.4 m2 box stalls (n=6) or in groups of three in a 992 m2 paddock, which had very limited grazing forage and an open shelter available (n=6). Subjects were fed concentrate and hay to National Research Council recommendations. A time budget for 31 observed behaviors was developed. Behavioral observations were made 2 days per week, approximately 6 h per day, for the duration of the 56 days study. Instantaneous samples were recorded every 5 min on each observation day, with equal division between the two treatment groups (n=35 scans per horse per observation day). Focal data were recorded continuously between scans to provide a more detailed ethogram. On each observation day, fecal samples were collected to measure 11,17-dioxoandrostanes, an indicator of glucocorticoid metabolite concentration. Regarding the fecal 11,17-dioxoandrostanes, there was no discernible treatment difference either immediately post-weaning or at the conclusion of the 56 days study. Interestingly, all 12 weanlings showed a 4 week post-weaning increase in 11,17-dioxoandrostanes. The reason for this peak was unclear. Behavioral observations demonstrated a significantly different time budget in paddock-housed weanlings than in stall-housed weanlings (P<0.0001). Paddock-housed weanlings displayed a time budget more like a feral horse time budget, showing more time spent moving and less time spent lying. Paddock-housed weanlings, who had the option of selectively engaging in a broader range of behaviors, showed strong motivation to graze and be near conspecifics. Stalled weanlings spent significantly more time engaged in aberrant behaviors: licking or chewing the stall/shed wall, kicking at the stall/shed wall, pawing, and bucking/rearing bouts (P<0.03). Based on the variety of behaviors shown, the ability to engage in strongly preferred behaviors, and freedom from aberrant behavior, we conclude that the paddock-reared, group-housed weanlings had better welfare. However, there was insufficient evidence to conclude that the stalled weanlings had poor welfare.

Abstract: We observed the behaviour of six stabled horses (stallions n = 3; geldings n = 3) in an attempt to identify behavioural measures of eating satisfaction. The horses were required to perform an operant response (pressing a button with the muzzle) in order to access a food reward in an experimental box stall. After each horse had successfully learned the experimental situation, it participated in the experimental protocol on 4 days. Horses were brought to the experimental box stall for the operant response sessions (1 h duration per session), and upon completion, they were returned to their own (home) box stalls. The number of presses for the reward was a Fixed Ratio schedule of either 3 or 12 muzzle presses (FR3, FR12) and the FR procedure for each horse was as follows: FR3 FR12 FR12 FR3 or FR12 FR3 FR3 FR12. Number of rewards obtained during each session, and behaviour and heart rate after each session were recorded for each horse. A repeated measures ANOVA showed that the number of rewards obtained in FR3 was higher than in FR12 (P < 0.05). The horses spent more time in standing-rest, (with ears rotating laterally and exhibiting a low neck position) indicating sleep, in the home box stall, after FR3 compared to FR12 treatments (P < 0.05). Mean heart rate after standing-sleep was significantly lower than mean heart rate in the home box stall (P < 0.01). These results suggest that eating satisfaction induces sleep in stabled horses, and that episodes of standing-sleep behaviour may be a useful indicator of appropriate or enhanced welfare in the horse.

Abstract: We investigated lateralisation in horses because it is likely to be important in training and athletic performance. Thoroughbred horses (n = 106) were observed every 60 s for 2 h, when they were at pasture, and the position of the forelimbs in relation to one another was recorded. There was a population bias skewed to standing with the left forelimb advanced over the right (i.e. directional lateralisation). Using the first 50 observations, the distribution of preferences was 43 significantly left, 10 significantly right with 53 being non-significant (i.e. ambidextextrous). The strength of motor bias increased with age, suggesting maturation or an influence of training. The horses were also presented with an olfactory stimulus (stallion faeces) to score the tendency to use one nostril rather than the other. A significant preference to use the right nostril first was shown in horses under 4 years of age (n = 61) but not in older horses. Of the 157 horses tested for nostril bias, 76 had been assessed for motor bias and so were used for analysis of the relationship between laterality in the two modalities. There was no significant relationship between direction of foreleg motor bias and first nostril used, total number of inhalations or laterality index of nostril use. The absence of a correlation between laterality of nostril use and motor bias indicates that lateralisation of the equine brain occurs on at least two levels of neural organisation--sensory and motor--a finding that is consistent with other examples of lateralisation in species that have been examined in more detail.