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McDonnell, S. M., & Henry, M. B., F. (1991). Spontaneous erection and masturbation in equids Proc 35th. J. Reprod. Fert. Suppl, 44, 664–665.
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Villani, M., Cairoli, F., Kindahl, H., Galeati, G., Faustini, M., Carluccio, A., et al. (2006). Effects of mating on plasma concentrations of testosterone, cortisol, oestrone sulphate and 15-ketodihydro-PGF2alpha in stallions. Reprod Domest Anim, 41(6), 544–548.
Abstract: Very little information is available regarding the physiological mechanisms involved in the normal sexual activity in the stallion and, in particular, the endocrine control of reproduction is still not clearly understood. This experiment was designed to determine the short-term effect of sexual stimulation on plasma concentrations of testosterone, cortisol, oestrone sulphate and 15-ketodihydro-PGF(2alpha) in stallions. Semen samples were collected from 10 lighthorse stallions of proven fertility using a Missouri model artificial vagina. At the same time, blood samples were collected from the jugular vein with heparinized tubes, 20 and 10 min before oestrous mare exposure, at exposure and 10, 20, 30 min after dismounting. Testosterone concentrations showed a sharp rise 10 min after mating (p < 0.001), reached a plateau, and then showed a further increase 30 min after mating (p < 0.001). Cortisol concentrations increased 10 min after mating (p < 0.001) and remained at high levels in the subsequent samples taken. A peak of oestrone sulphate was observed 10 min after mating (p < 0.001). 15-Ketodihydro-PGF(2alpha) concentrations decreased rapidly at the moment of the exposure of the stallions to an oestrous mare (p < 0.05), returned to pre-mating concentrations and then decreased again 30 min after mating (p < 0.05).
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Feist, J. D., & McCullough, D. R. (1976). Behavior patterns and communication in feral horses. Z. Tierpsychol., 41(4), 337–371.
Abstract: The social behavior of feral horses was studied in the western United States. Stable harem groups with a dominant stallion and bachelor hermaphrodite hermaphrodite groups occupied overlapping home ranges. Groups spacing, but not territoriality, was expressed. Harem group, stability resulted from strong dominance by dominant stallions, and fidelity of group members. Eliminations of group members were usually marked by urine of the dominant stallion. Hermaphrodite-hermaphrodite aggression involved spacing between harems and dominance in bachelor groups. Marking with feces was important in hermaphrodite-hermaphrodite interactions. Foaling occurred in May and early June, following the post-partum estrous. All breeding was done by harem stallions. Young were commonly nursed through yearling age. These horses showed social organizations similar to other feral horses and plains zebras.
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Pickerel, T. M., Crowell-Davis, S. L., Caudle, A. B., & Estep, D. Q. (1993). Sexual preference of mares (Equus caballus) for individual stallions. Appl. Anim. Behav. Sci., 38(1), 1–13.
Abstract: Eight mares were tested to determine if they remained near one of two stallions longer than would be expected if association was random. Six stallions were paired in 30 combinations and each mare was tested 30 times. The mares (Equus caballus) demonstrated a definite preference for individual stallions throughout the breeding season. This preference was influenced by the estrous state of the mare. During estrus, mares' preferences for stallions were positively correlated with the rate at which a given stallion vocalized. During diestrus, mares spent significantly less time in the proximity of stallions and did not exhibit any preference for individual stallions.
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Gil, M., Bhatt, R., Picotte, K. B., & Hull, E. M. (2013). Sexual experience increases oxytocin receptor gene expression and protein in the medial preoptic area of the male rat. In Psychoneuroendocrinology (Vol. 38, pp. 1688–1697). Pergamon Press.
Abstract: Oxytocin (OT) promotes social and reproductive behaviors in mammals, and OT deficits may be linked to disordered social behaviors like autism and severe anxiety. Male rat sexual behavior is an excellent model for OT regulation of behavior, as its pattern and neural substrates are well characterized. We previously reported that OT microinjected into the medial preoptic area (MPOA), a major integrative site for male sexual behavior, facilitates copulation in sexually experienced male rats, whereas intra-MPOA injection of an OT antagonist (OTA) inhibits copulation. In the present studies, copulation on the day of sacrifice stimulated OTR mRNA expression in the MPOA, irrespective of previous sexual experience, with the highest levels observed in first-time copulators. In addition, sexually experienced males had higher levels of OTR protein in the MPOA than sexually naïve males and first-time copulators. Finally, intra-MPOA injection of OT facilitated mating in sexually naive males. Others have reported a positive correlation between OT mRNA levels and male sexual behavior. Our studies show that OT in the MPOA facilitates mating in both sexually naive and experienced males, some of the behavioral effects of OT are mediated by the OTR, and sexual experience is associated with increased OTR expression in the MPOA. Taken together, these data suggest a reciprocal interaction between central OT and behavior, in which OT facilitates copulation and copulation stimulates the OT/OTR system in the brain.
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Turner, J. W. J., & Kirkpatrick, J. F. (1982). Androgens, behaviour and fertility control in feral stallions. J Reprod Fertil Suppl, 32, 79–87.
Abstract: This field study of feral stallions in Montana and Idaho examines and correlates the seasonal pattern of plasma androgens and specific sociosexual behaviour and reports the effect of a long-acting androgenic steroid on this behaviour and on fertility. Plasma testosterone was measured by competitive protein binding assay in samples obtained by jugular venepuncture from captured animals. In samples taken from 34 sexually mature stallions in 6 different months during the year, a definite seasonal pattern in testosterone was present, with a peak in May (3.04 +/- 0.63 ng/ml) and a nadir in December (1.55 +/- 0.34 ng/ml). Values were less than 2.0 ng/ml in non-breeding months and greater than 2.4 ng/ml in breeding months. Behavioural endpoints measured were (1) stallion scent marking in response to elimination by mares (elimination marking), (2) mounting and (3) copulation. The frequencies of each of these endpoints followed closely the seasonal pattern seen for plasma androgens. In the fertility study microcapsulated testosterone propionate (microTP) was administered i.m. to 10 harem stud stallions 3 months before the 1980 breeding season. In these stallions and in 10 control harem studs, the above behavioural endpoints were examined in the 1980 and 1981 breeding seasons, and foal counts were made in 1981. There were no direct inhibitory or stimulatory effects of microTP treatment on any of the behavioural endpoints in either year. In 1981 foals were produced in 87.5% of the control bands and 28.4% of the microTP-treated bands. These results indicate that microencapsulated testosterone propionate can provide effective fertility control in feral horses without causing significant alterations in sociosexual behaviour.
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Klingel, H. (1982). Social organization of feral horses. J Reprod Fertil Suppl, 32, 89–95.
Abstract: The basic social unit in feral horses is the family group consisting of one stallion, one to a few unrelated mares and their foals. Surplus stallions associate in bachelor groups. Stallions are instrumental in bringing mares together in a unit which then persists even without a stallion. The similarity of social organization in populations living in a variety of different habitats indicates that feral horses have reverted to the habits of their wild ancestors, and that domestication has had no influence on this basic behavioural feature.
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Kiley, M. (1972). The vocalizations of ungulates, their causation and function. Z. Tierpsychol., 31(2), 171–222.
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Kaseda, Y., Ogawa, H., & Khalil, A. M. (1997). Causes of natal dispersal and emigration and their effects on harem formation in Misaki feral horses. Equine Vet J, 29(4), 262–266.
Abstract: Misaki feral horses were separated into 2 herds and the difference between dispersal from natal group (natal dispersal) and dispersal from natal area (natal emigration) was studied. The causes of dispersal and emigration and their effects on harem formation were studied 1979-1994. The number of horses ranged from 73 (mature males: 8, mature females: 26, young males: 8, young females: 3, colt foals: 6, filly foals: 10 and geldings: 12) in 1979 and 86 (mature males: 14, mature females: 37, young males: 12, young females: 7, colt foals: 5, filly foals: 7 and geldings: 4) in 1994 when the present study ended. All 29 males which survived to age 4 years and 58 females which survived to age 3 years left their natal or mother groups at age one to 3. Seventeen of 22 dispersing males and 29 of 39 dispersing females left their natal groups around the birth of their siblings and significant correlations were found between natal dispersal and birth of a sibling. The number of emigrating young males correlated negatively and significantly with the total number of young males in another herd and the number of emigrating young females correlated positively and significantly with the total number of young females in the natal herd. All 13 emigrating stallions which survived to age 5 years formed stable harem groups and a significant correlation was found between natal emigration and harem formation. Twenty-three of 35 resident mares formed stable consort relations with harem stallions and a significant correlation was found between residence and formation of stable consort relations.
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Hrdy, S. B. (1974). Male-male competition and infanticide among the langurs (Presbytis entellus) of Abu, Rajasthan. Folia Primatol (Basel), 22(1), 19–58.
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