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: To investigate the functions of social licking in cattle, four calves (one heifer and one steer in each of two herds), known to exhibit frequent social licking were observed continuously for 2 h before sunset for 13 days, using the focal animal sampling method. Calves were observed under various environmental conditions. Social licking significantly decreased on rainy days and tended to increase in a dirty barn and when food was restricted. Solicitation for social licking occurred not only from dominant animals of pairs but also from subordinates. Of the licking interactions, 31% occurred following solicitation, and these accounted for 39% of the total time spent licking. Following solicitation, 78% of social licking was oriented to the head and the neck regions that were inaccessible to self-licking animals. Unsolicited licking, however, was oriented not only to the head and the neck but also to the back and the rump regions, and these two latter regions were the major ones to receive licking. The effect of social relationships on social licking was investigated using least-squares analysis of variance. Social factors investigated were the difference of dominance values, the dominance-subordinance relationship, and kinship and familiarity; the sex of calves involved was also considered. Only familiarity had a significant effect on licking; exchanges of social licking increased with length of cohabitation. We suggest that social licking may have a cleaning effect, a tension-reducing effect and a bonding effect.

Abstract: Herding behavior in ungulates is executed mainly by males. There are several forms of herding: guarding a single estrous female; rounding up a bunch of females during the rutting season; territorial herding by which a male keeps females inside his territory; herding of a moving, permanent, harem group; social herding in which group members of both sexes are herded by one dominant male. When put into this sequence, a phenotypical trend is illustrated, leading from an intimate connection of herding with mating behavior toward an increasing independence from sexual behavior and culminating in a complete socialization of herding. Aspects and problems of herding behavior are the recognition of, and the animal's respect for, partners in gregarious species; the animal “taking offense” at activities of others which deviate from its own activity; the active coordination and synchronization of group activities; the use of, and the effects of, threat, dominance and courtship displays in herding; the possibility of substitution among expressive displays in relation to partners of different sex; the communicative function of the animal's orientation relative to the partner; social hierarchy and leadership in a group; the possibility of interspecific herding, particularly in man-animal relationships, which is closely linked to the process of domestication.

Abstract: The aim of this study was to investigate social behaviour in differently reared stallions in their respective environments; one group of stallions was reared under typical domestic conditions whereas the other group was reared and lives under natural conditions. The domestic group consisted of 19, 2-year-old stallions (Equus caballus), which were all weaned at 4 months of age and experienced either individual or group housing facilities before being pastured with the other similarly aged stallions. The natural living and mixed age group of Przewalski stallions (E. przewalskii) consisted of 13 stallions, most of which were juveniles (n=11, <=4 years; n=2, >9 years). The domestic group was studied in a 4-ha enclosure at the Danish Institute of Agricultural Sciences and the Przewalski group under free-ranging conditions in a 75-ha enclosure in the Askania Nova Biosphere Reserve, Ukraine. Behavioural data was collected during 168 h of direct observation. The occurrence of 14 types of social interactions was recorded and group spacing behaviour was studied using nearest neighbour recordings. In spite of very different environments, reflecting domestic and natural rearing conditions, many similarities in behaviour was found. Play and play fight behaviour was very similar in the two stallion groups. Quantitative differences were found in social grooming since Przewalski stallions groomed more frequently (P=0.004), and in investigative behaviours, since domestic stallions showed more nasal (P=0.005) and body sniffing (P<0.001), whereas Przewalski stallions directed more sniffing towards the genital region (P<0.001). These differences may, however, be attributed to environmental factors and in the period of time the stallions were together prior to the study period. Quantitative differences appeared in some agonistic behaviours (kick threat, P<0.001; and kick, P<0.001), but data do not support earlier findings of Przewalski horses being significantly more aggressive than domestic horses. In general, Przewalski stallions engaged in more social interactions, and they showed less group spacing, i.e. maintained a significantly shorter distance between neighbours (P<0.001). The study indicates that also domestic horses, which have been reared under typical domestic conditions and allowed a period on pasture, show social behaviour, which is very similar to that shown by their non-domestic relatives.

Abstract: Nursing behaviour and related aggression of mare-foal pairs was studied from birth (n = 21) to 24 weeks of age (n = 15) of the foal. Foals exhibited a decreasing length and frequency of nursing as they grew older. Mares rarely aggressed against their foals during nursing in the foal's first 4 weeks of life, but did so increasingly through Weeks 13-16, after which the rate of aggression during nursing decreased. Mares terminated nursing primarily by moving away, and were most likely to do so during the foal's first 4 weeks of life. They became gradually less likely to do so as the foal grew older. It was concluded that mares sometimes flex their hind limb on the side opposite the foal during nursing in order to conserve energy in a situation in which they would be remaining still anyway. There was no difference between colts and fillies in the frequency or duration of nursing or in the frequency with which their mothers aggressed against them or terminated nursing.