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de Waal, F. B. (1996). Macaque social culture: development and perpetuation of affiliative networks. J Comp Psychol, 110(2), 147–154.
Abstract: Maternal affiliative relations may be transmitted to offspring, similar to the way in which maternal rank determines offspring rank. The development of 23 captive female rhesus monkeys (Macaca mulatta) was followed from the day of birth until adulthood. A multivariate analysis compared relations among age peers with affiliative relations, kinship, and rank distance among mothers. Maternal relations were an excellent predictor of affiliative relations among daughters, explaining up to 64% of the variance. Much of this predictability was due to the effect of kinship. However, after this variable had been controlled, significant predictability persisted. For relations of female subjects with male peers, on the other hand, maternal relations had no significant predictive value beyond the effect of kinship. One possible explanation of these results is that young rhesus females copy maternal social preferences through a process of cultural learning.
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de Waal, F. B., Uno, H., Luttrell, L. M., Meisner, L. F., & Jeannotte, L. A. (1996). Behavioral retardation in a macaque with autosomal trisomy and aging mother. Am J Ment Retard, 100(4), 378–390.
Abstract: The social development of a female rhesus monkey (Macaca mulatta) was followed from the day of birth until her death, at age 32 months. The subject, born to an older mother, had an extra autosome (karyotype: 43, XX, +18), an affliction that came about spontaneously. MRI scans revealed that she was also hydrocephalic. Compared to 23 female monkeys growing up under identical conditions, the subject showed serious motor deficiencies, a dramatic delay in the development of social behavior, poorly established dominance relationships, and greater than usual dependency on mother and kin. The subject was well-integrated into the social group, however.
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Henry, S., Hemery, D., Richard, M. - A., & Hausberger, M. (2005). Human-mare relationships and behaviour of foals toward humans. Appl. Anim. Behav. Sci., 93(3-4), 341–362.
Abstract: We studied experimentally whether horse dams influenced foals' relationships with humans. We investigated the influence of the establishment of positive human-mare relationships on foals' behaviour toward humans. Forty-one foals and their dams were involved in this experiment. Half of the mares were softly brushed and fed by hand during a short period (total of 1.25 h) during the first 5 days of their foals' lives (experimental group, n = 21). The other mares were not handled experimentally and their foals received no contact with the experimenter (control group, n = 20). The reactions of both experimental and control foals were recorded under various conditions, first, for 5 min in the presence of a motionless experimenter, when foals were 15 and 30-35 days old, then in an approach test when they were 15 days old and in a saddle-pad tolerance test when they were 30-35 days old. Finally, approach-stroking tests were performed successively by the familiar experimenter when foals were 11-13 months old and by an unfamiliar person when they were 13-15 months old. Several observations strongly suggest that mares can influence their foals' behaviour toward humans: (1) during the handling procedure, experimental foals of protective mares were further from the handler than foals of calm mares (p < 0.001); (2) experimental foals remained, at all ages, closer to the experimenter (p < 0.05) and initiated more physical contacts (sniffing, licking, etc.) with the experimenter (p < 0.05) than control foals; (3) avoidance and flight responses of experimental foals were considerably reduced during approaches by the experimenter (p < 0.01) and they accepted saddle-pads on their backs more easily (p < 0.01) and more quickly (p < 0.01) than control foals. Lastly, the consequences of handling mares had effects that lasted at least until foals were one year old (p < 0.05) and became generalized from experimenter to unfamiliar humans, who could approach and stroke experimental foals rapidly during a test (p < 0.05). This is the first report of an attempt to use observation of mother by foals to facilitate human-foal relationships. The procedure is simple, takes little time and can easily be applied to any dam-foal pair, as it is not intrusive and presents no risks of disrupting mare-foal bonds.
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McGee, S., & Smith, H. V. (2004). Accompanying pre-weaned Thoroughbred (Equus caballus) foals while separated from the mare during covering reduces behavioural signs of distress exhibited. Appl. Anim. Behav. Sci., 88(1-2), 137–147.
Abstract: Some Thoroughbred stallion farms separate the mare from the foal during breeding, leaving the foal alone in its stall. Such separation often leads to distress in the foal, as evidenced by behavioural indicators such as vocalisations, pacing or moving about, pawing and striking the body against solid objects. This study examined whether the foal's distress is reduced if a handler accompanies it during the period of separation. A total of 57 foals on an Irish stud farm were observed: 27 foals were accompanied during separation and 30 were left in isolation but discretely observed from outside the stall. On average, unaccompanied foals vocalised at significantly (P<0.001) shorter intervals (14.8 s versus 26.8 s), and spent significantly (P<0.05) more time pawing (29.6 s versus 6.8 s) than accompanied ones. In addition, 17 of the unaccompanied foals hurled themselves against a solid object on at least one occasion, while not one of the accompanied foals did this (P<0.001). Age, gender and number of times the foal had been previously separated from the mare while she was being teased or covered were not significantly related to any of the behavioural indicators of distress. Overall, the data provide clear evidence that accompanying a foal reduces the distress that it exhibits when separated from the mare. This procedure has obvious welfare benefits in the short term, in that the foal both experiences less distress and has a reduced likelihood of injuring itself, and it may have longer-term benefits by reducing the chance of the animal later developing a locomotory stereotypical behaviour.
Keywords: Distress; Foals; Maternal separation; Stereotypies; Welfare
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Vieuille, C., Berger, F., Le Pape, G., & Bellanger, D. (2003). Sow behaviour involved in the crushing of piglets in outdoor farrowing huts--a brief report. Appl. Anim. Behav. Sci., 80(2), 109–115.
Abstract: This study focuses on maternal activities involved in the trapping of piglets by the sow's body in outdoor farrowing systems and examine the mother-piglet context leading either to the death of piglets or to their survival. The behaviour of six Large-WhitexLandrace sows and their litters was continuously video recorded at their first and second parity, during the 40 h following parturition. Crushing mainly occurred at evening and night, during the first 12 h of farrowing and involved changes between lying, sitting and standing positions, as well as between udder and side lying. No piglet died from savaging. Nevertheless, aggressive behaviours of sows were observed, particularly in their first maternal experience. The immediate context of trapping was related to the labour of the sow and to the feeding and resting of piglets. The immediate crushing context was related to active avoidance of restless piglets while lying down, as well as sitting and standing behaviours. These results are discussed in terms of differential reactions of the sow to suckling attempts of piglets.
Keywords: Pig-maternal behaviour; Crushing; Free-ranging; Welfare
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Houpt, K. A. (2002). Formation and dissolution of the mare-foal bond. Appl. Anim. Behav. Sci., 78(2-4), 319–328.
Abstract: The behavior of mares at foaling and their behavior during subsequent weekly brief separations from their foals was quantified. Thirteen multiparous pony mares were observed for the first 30 min after foaling. Activities directed toward the foal and toward the fetal membranes were recorded. There were 16+/-8 contacts with the fetal membranes, most in the first 10 min post-partum. There were 81+/-12 activities directed toward the foal. There was a downward trend in foal contacts over the first 30 min. Eight of the 13 mares were observed for the 30-60 min post-partum during which licking and touching the foal continued. Only one of the foals suckled in the first 30 min, but half had suckled by 60 min. Ten mares and foals were separated for 5 min each week for 9 weeks. Although mare responses (neighs and steps) decreased with age of the foal, the foals' responses increased from weeks 1 to 2 and then decreased with foal age. This indicates that foal attachment to the mare is not complete until it is 2 weeks old.
Keywords: Maternal; Horse; Foal; Separation; Ontogeny
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Cameron, E. Z., Linklater, W. L., Stafford, K. J., & Minot, E. O. (2008). Maternal investment results in better foal condition through increased play behaviour in horses. Anim. Behav., 76(5), 1511–1518.
Abstract: Play behaviour is widespread in mammals, but benefits to play have been difficult to demonstrate. Physical training is one of the many proposed hypotheses, suggesting that males and females should play differently, that increased maternal investment should lead to increases in play, and that increases in play should result in physical advantages. In a population of feral horses, Equus caballus, males and females did not differ in their play behaviour except that males initiated more of their play bouts than females. Maternal condition influenced play behaviour only in males, with sons of mothers in good condition playing more. However, when we controlled for maternal effects by comparing a son and a daughter of the same mother, daughters played more when their mother was in poor condition and sons played more when their mother was in good condition. Mothers of foals that played more lost more condition. Therefore, the difference in play behaviour could not be explained by offspring sex or maternal condition alone, but play behaviour mirrored variation in maternal investment. In addition, those individuals that played more survived better and had better body condition as yearlings despite weaning earlier. Since increased activity has been linked to enhanced musculoskeletal development in domestic horses, we suggest that play provides a link between increased maternal investment, increased body condition and future reproductive success in feral horses, and probably in other species.
Keywords: Equus caballus; feral horse; maternal investment; play
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Houpt, K. A. (1976). Animal behavior as a subject for veterinary students. Cornell Vet, 66(1), 73–81.
Abstract: Knowledge of animal behavior is an important asset for the veterinarian; therefore a course in veterinary animal behavior is offered at the New York State College of Veterinary Medicine as an elective. The course emphasizes the behavior of those species of most interest to the practicing veterinarian: cats, dogs, horses, cows, pigs and sheep. Dominance heirarchies, animal communication, aggressive behavior, sexual behavior and maternal behavior are discussed. Play, learning, diurnal cycles of activity and sleep, and controls of ingestive behavior are also considered. Exotic and zoo animal behaviors are also presented by experts in these fields. The critical periods of canine development are related to the optimum management of puppies. The behavior of feral dogs and horses is described. The role of the veterinarian in preventing cruelty to animals and recognition of pain in animals is emphasized. Whenever possible behavior is observed in the laboratory or on film.
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Houpt, K. A., Parsons, M. S., & Hintz, H. F. (1982). Learning ability of orphan foals, of normal foals and of their mothers. J. Anim Sci., 55(5), 1027–1032.
Abstract: The maze learning ability of six pony foals that had been weaned at birth was compared to that of six foals reared normally. The foals' learning ability was also compared to their mothers' learning ability at the same task; the correct turn in a single choice point maze. The maze learning test was conducted when the foals were 6 to 8 mo old and after the mothered foals had been weaned. There was no significant difference between the ability of orphaned (weaned at birth) and mothered foals in their ability to learn to turn left (6 +/- .7 and 5.1 +/- .1 trials, respectively) or to learn the reversal, to turn right (6.7 +/- .6 and 6.2 +/- .6 trials, respectively). The orphan foals spent significantly more time in the maze in their first exposure to it than the mothered foals (184 +/- 42 vs 55 +/- 15 s. Mann Whitney U = 7, P less than .05). The mothers of the foals (n = 11) learned to turn left as rapidly as the foals (5.9 +/- .7 trials), but they were slower to learn to turn right (9.8 +/- 1.4 vs 6.4 +/- .4 trials, Mann Whitney U = 33, P less than .05), indicating that the younger horses learned more rapidly. There was no correlation between the trials to criteria of the mare and those of her foal, but there was a significant negative correlation between rank in trials to criteria and age (r = -65, P less than .05) when data from the mare and foal trials were combined. The dominance hierarchy of the mares was determined using a paired feeding test in which two horses competed for one bucket of feed. Although there was no correlation between rank in the hierarchy and maze learning ability, there was a correlation between body weight and rank in the hierarchy (r = .7, P less than .05). This may indicate either that heavier horses are likely to be dominant or that horses high in dominance gain more weight. Maternal deprivation did not appear to seriously retard learning of a simple maze by foals, although the orphans moved more slowly initially. The lack of maternal influence on learning is also reflected in the lack of correlation between the mare's learning ability and that of her foal. Young horses appear to learn more rapidly than older horses.
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Khalil, A. M., & Kaseda, Y. (1997). Behavioral patterns and proximate reason of young male separation in Misaki feral horses. Appl. Anim. Behav. Sci., 54(4), 281–289.
Abstract: The present investigation was undertaken to study the proximate reasons why and the behavioral patterns of young male Misaki feral horses when they left their natal band or mothers. We observed a total of ten young males twice a month from January 1988 to December 1995. Almost all young males left their natal band or mothers at between 1 and 4 years of age. We found that, during the separation process, all the young males from first parity dams returned several times after the initial separation, indicating a strong attachment between primiparous mares and their male offspring. The other five separated only once without rejoining. Our observations showed five variable behavior patterns of young males at separation time, depending on the consort relation between their mothers and harem stallion and the reason for separation at that time. Eight young males separated in the non-breeding season at average 2.1 years and the other two separated in the breeding season at average 3 years and the average difference was not significant. These results revealed that 80% of the young males separated voluntarily when the natural resources become poor whereas 20% separated when their siblings were born.
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