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Boyd, L. (1986). Behavior problems of equids in zoos. Vet Clin North Am Equine Pract, 2(3), 653–664.
Abstract: Behavior problems in zoo equids commonly result from a failure to provide for needs basic to equine nature. Equids are gregarious, and failure to provide companions may result in pacing. Wild equids spend 60 to 70 per cent of their time grazing, and failure to provide ad libitum roughage contributes to the problems of pacing, cribbing, wood chewing, and coprophagia. Mimicking the normal processes of juvenile dispersal, bachelor-herd formation, and mate acquisition reduces the likelihood of agonistic and reproductive behavior problems. Infanticide can be avoided by introducing new stallions to herds containing only nonpregnant mares and older foals.
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Kirkpatrick, J. F., & Turner, A. (2002). Reversibility of action and safety during pregnancy of immunization against porcine zona pellucida in wild mares (Equus caballus). Reprod Suppl, 60, 197–202.
Abstract: Contraceptive management of publicly valued wildlife species requires safeguards to ensure that these populations are preserved in a healthy state. In addition, reversibility of contraceptive effects and safety in pregnant animals are major concerns. A population of wild horses has been immunized against porcine zona pellucida (PZP) over a 12 year period on Assateague Island National Seashore, MD (ASIS). Mares initially received one or two 65 microg inoculations and once a year 65 microg booster inoculations, all delivered by dart. All young mares aged > 2 years were treated with PZP for 3 consecutive years regardless of whether they have bred successfully and they were then removed from treatment until they had foaled. All mares vaccinated for 1 or 2 consecutive years became fertile again and 69% of mares treated for 3 consecutive years returned to fertility. All five mares treated for 4 or 5 consecutive years have also returned to fertility, but over longer periods of time. Mares treated for 7 consecutive years have not returned to fertility, but several, while still infertile, have started ovulating again. There was no difference in survival rates between foals born to treated and untreated mares, and PZP treatment of pregnant mares did not affect subsequent fertility of their female offspring.
Keywords: Animals; *Animals, Wild; Antigens/administration & dosage; Contraception, Immunologic/methods/*veterinary; Egg Proteins/administration & dosage; Female; Fertility; *Horses; Immunization, Secondary/veterinary; Membrane Glycoproteins/administration & dosage; Population Control; Pregnancy; *Receptors, Cell Surface; Safety; Swine; Time Factors; Vaccines, Contraceptive/*administration & dosage
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Kirkpatrick, J. F., & Turner, A. (2003). Absence of effects from immunocontraception on seasonal birth patterns and foal survival among barrier island wild horses. J Appl Anim Welf Sci, 6(4), 301–308.
Abstract: Despite a large body of safety data, concern exists that porcine zonae pellucidae (PZP) immunocontraception--used to manage wild horse populations--may cause out-of-season births with resulting foal mortality. Our study at Assateague, Maryland indicated the effects of immunocontraception on season of birth and foal survival between 1990 and 2002 on wild horses from Assateague Island. Among 91 mares never treated, 69 (75.8%) of foals were born in April, May, and June (in season). Among 77 treated mares, 50 (64.9%) were born in season. Of 29 mares foaling within 1 year after treatment (contraceptive failures), 20 (68.9%) were born in season. Of 48 mares treated for greater than 2 years then withdrawn from treatment, 30 (62.5%) of 48 foals were born in season. There were no significant differences (p <.05) between either treatment group or untreated mares. Survival did not differ significantly among foals born in or out of season or among foals born to treated or untreated mares. Data indicate a lack of effect of PZP contraception on season of birth or foal survival on barrier island habitats.
Keywords: Animals; Animals, Newborn; Animals, Wild/*physiology; Birth Rate; Case-Control Studies; Contraception, Immunologic/methods/*veterinary; Egg Proteins/administration & dosage; Female; Horses/*physiology; Maryland/epidemiology; Membrane Glycoproteins/administration & dosage; Population Control; Pregnancy; *Receptors, Cell Surface; *Reproduction; Seasons; Vaccines, Contraceptive/administration & dosage
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Schwarzenberger, F., Mostl, E., Bamberg, E., Pammer, J., & Schmehlik, O. (1991). Concentrations of progestagens and oestrogens in the faeces of pregnant Lipizzan, trotter and thoroughbred mares. J Reprod Fertil Suppl, 44, 489–499.
Abstract: Faecal samples were collected at weekly intervals from pregnant Lipizzan mares during Weeks 7-16 following mating and from Lipizzan, Trotter and Thoroughbred mares during the last 3 months of gestation. After parturition, samples were taken daily from the Thoroughbred mares for another 6 days. Non-pregnant mares served as controls. The concentrations of unconjugated oestrogens (Eg), 20 alpha-OH-progestagens (20 alpha-G) and 20 beta-OH-progestagens (20 beta-G) were measured by enzyme immunoassay. In the faeces of Lipizzan mares, immunoreactive progestagens were significantly (P less than 0.01) elevated above the levels in non-pregnant mares by Week 11, and Eg by Week 13 of pregnancy onwards. During the last 3 months of gestation, concentrations of Eg were significantly higher in Trotter mares than in Lipizzan and Thoroughbred mares. Concentrations of 20 alpha-G and 20 beta-G increased to maximal values in the last month of gestation. There was no significant difference among the 3 breeds with respect to 20 alpha-G but, during the 10 weeks before parturition, concentrations of 20 beta-G in the Lipizzan mares were significantly lower (P less than 0.05) than those in the Thoroughbred mares. They were also significantly lower than those of the Trotter mares during the last 4 weeks of gestation. After parturition, the concentrations of Eg and progestagens had declined to baseline values by Days 3 and 4 respectively. From these results we conclude that high concentrations of progestagens with 20 alpha- and 20 beta-hydroxyl groups are present in the faeces of pregnant mares, especially during the last month of gestation.
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Lucas, Z., Raeside, J. I., & Betteridge, K. J. (1991). Non-invasive assessment of the incidences of pregnancy and pregnancy loss in the feral horses of Sable Island. J Reprod Fertil Suppl, 44, 479–488.
Abstract: Field observations of 400 totally unmanaged feral horses on Sable Island, Nova Scotia, were complemented by oestrogen determinations in faecal samples from 154 identified females over a 4-year period (454 mare-years). Of mares that were sampled throughout the year and subsequently produced foals, 92.1% exhibited elevated faecal oestrogens between 15 October and 30 March. The results confirm that faecal oestrogens are a useful indicator of pregnancy after approximately 120 days gestation. Distribution of foaling resembled that seen in other feral populations, with 95% of births occurring from April through July. The foaling rate for mares aged 3 years or older was 62.0%, with 50.7% of mares foaling in 3 or 4 years. Foaling rates were low (4.1%) in mares bred as yearlings and rose with age to 70.8% in those bred as 4-year-olds. Fetal loss after Day 120 was deduced from faecal oestrogens to be 26.0% overall, with marked variation from year to year (9.6-37.3%) and with age (70.0% in those bred as yearlings, decreasing to 5.6% in those bred as 4-year-olds). Of 58 mares aged 2 years or older that were sampled every year, about half (49.6%) the barren years were attributable to fetal loss after 120 days gestation. All mares conceived in at least 2 of the 4 years, suggesting that pregnancy loss, even after Day 120, is as important as failure to conceive in causing barren years.
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Franceschini, C., Siutz, C., Palme, R., & Millesi, E. (2007). Seasonal changes in cortisol and progesterone secretion in Common hamsters. Gen Comp Endocrinol, 152(1), 14–21.
Abstract: In this study, we investigated endocrine factors and behaviour in free-living Common hamsters (Cricetus cricetus) during reproductive and non-reproductive periods of the annual cycle. We applied a non-invasive method to gain information on seasonal changes in adrenocortical activity in male and female hamsters by analysing faecal glucocorticoid metabolite concentrations (FCM). In addition, plasma progesterone concentrations were monitored in females throughout the non-hibernation season. The animals were live-trapped from spring emergence until the onset of hibernation in autumn. Reproductive status was determined at capture and blood and faecal samples were collected. During behavioural observations, agonistic and sexual interactions were recorded. FCM concentrations were significantly higher in males than in females during the reproductive period. In males, a pronounced increase in FCM during the reproductive period coincided with high frequencies of intrasexual aggression. In females, FCM levels remained relatively constant. Aggressive behaviour in females increased during the reproductive period, but was much less frequent than in males. Females, which successfully raised a second litter after a postpartum oestrus and concurrent lactation and gestation had lower FCM levels than individuals, which lost their second litter after parturition. As expected, plasma progesterone concentrations were low before and after the reproductive period. During gestation, levels peaked and remained elevated during lactation. The results of this field study provide insight in critical periods associated with reproduction in male and female Common hamsters.
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Brennan, P. A. (2004). The nose knows who's who: chemosensory individuality and mate recognition in mice. Horm Behav, 46(3), 231–240.
Abstract: Individual recognition is an important component of behaviors, such as mate choice and maternal bonding that are vital for reproductive success. This article highlights recent developments in our understanding of the chemosensory cues and the neural pathways involved in individuality discrimination in rodents. There appear to be several types of chemosensory signal of individuality that are influenced by the highly polymorphic families of major histocompatibility complex (MHC) proteins or major urinary proteins (MUPs). Both have the capability of binding small molecules and may influence the individual profile of these chemosignals in biological fluids such as urine, skin secretions, or saliva. Moreover, these proteins, or peptides associated with them, can be taken up into the vomeronasal organ (VNO) where they can potentially interact directly with the vomeronasal receptors. This is particularly interesting given the expression of major histocompatibility complex Ib proteins by the V2R class of vomeronasal receptor and the highly selective responses of accessory olfactory bulb (AOB) mitral cells to strain identity. These findings are consistent with the role of the vomeronasal system in mediating individual discrimination that allows mate recognition in the context of the pregnancy block effect. This is hypothesized to involve a selective increase in the inhibitory control of mitral cells in the accessory olfactory bulb at the first level of processing of the vomeronasal stimulus.
Keywords: Animals; Chemoreceptors/physiology; Discrimination Learning/*physiology; Embryo Implantation/physiology; Female; Individuality; Major Histocompatibility Complex/physiology; Male; Mice; Neurons, Afferent/physiology; Nose/cytology/physiology; Perception/physiology; Pregnancy; Pregnancy Maintenance/physiology; Pregnancy, Animal/*physiology; Receptors, Odorant/*physiology; Recognition (Psychology)/*physiology; Sexual Behavior, Animal/*physiology; Smell/*physiology; Urine/physiology; Vomeronasal Organ/cytology/physiology
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De Boyer Des Roches, A., Richard-Yris, M. - A., Henry, S., Ezzaouia, M., & Hausberger, M. (2008). Laterality and emotions: visual laterality in the domestic horse (Equus caballus) differs with objects' emotional value. Physiol. Behav., 94(3), 487–490.
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.
Keywords: Animals; Animals, Newborn; Behavior, Animal/physiology; Dominance, Ocular/*physiology; *Emotions; Exploratory Behavior/physiology; Female; Horses/*physiology; Olfactory Pathways/physiology; Pattern Recognition, Visual/*physiology; Photic Stimulation; Pregnancy; Statistics, Nonparametric; Visual Fields/physiology
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Brennan, P. A., & Kendrick, K. M. (2006). Mammalian social odours: attraction and individual recognition. Phil. Trans. Biol. Sci., 361(1476), 2061–2078.
Abstract: Mammalian social systems rely on signals passed between individuals conveying information including sex, reproductive status, individual identity, ownership, competitive ability and health status. Many of these signals take the form of complex mixtures of molecules sensed by chemosensory systems and have important influences on a variety of behaviours that are vital for reproductive success, such as parent-offspring attachment, mate choice and territorial marking. This article aims to review the nature of these chemosensory cues and the neural pathways mediating their physiological and behavioural effects. Despite the complexities of mammalian societies, there are instances where single molecules can act as classical pheromones attracting interest and approach behaviour. Chemosignals with relatively high volatility can be used to signal at a distance and are sensed by the main olfactory system. Most mammals also possess a vomeronasal system, which is specialized to detect relatively non-volatile chemosensory cues following direct contact. Single attractant molecules are sensed by highly specific receptors using a labelled line pathway. These act alongside more complex mixtures of signals that are required to signal individual identity. There are multiple sources of such individuality chemosignals, based on the highly polymorphic genes of the major histocompatibility complex (MHC) or lipocalins such as the mouse major urinary proteins. The individual profile of volatile components that make up an individual odour signature can be sensed by the main olfactory system, as the pattern of activity across an array of broadly tuned receptor types. In addition, the vomeronasal system can respond highly selectively to non-volatile peptide ligands associated with the MHC, acting at the V2r class of vomeronasal receptor.The ability to recognize individuals or their genetic relatedness plays an important role in mammalian social behaviour. Thus robust systems for olfactory learning and recognition of chemosensory individuality have evolved, often associated with major life events, such as mating, parturition or neonatal development. These forms of learning share common features, such as increased noradrenaline evoked by somatosensory stimulation, which results in neural changes at the level of the olfactory bulb. In the main olfactory bulb, these changes are likely to refine the pattern of activity in response to the learned odour, enhancing its discrimination from those of similar odours. In the accessory olfactory bulb, memory formation is hypothesized to involve a selective inhibition, which disrupts the transmission of the learned chemosignal from the mating male. Information from the main olfactory and vomeronasal systems is integrated at the level of the corticomedial amygdala, which forms the most important pathway by which social odours mediate their behavioural and physiological effects. Recent evidence suggests that this region may also play an important role in the learning and recognition of social chemosignals.
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Arnold, W., Ruf, T., & Kuntz, R. (2006). Seasonal adjustment of energy budget in a large wild mammal, the Przewalski horse (Equus ferus przewalskii) II. Energy expenditure. J Exp Biol, 209(Pt 22), 4566–4573.
Abstract: Many large mammals show pronounced seasonal fluctuations of metabolic rate (MR). It has been argued, based on studies in ruminants, that this variation merely results from different levels of locomotor activity (LA), and heat increment of feeding (HI). However, a recent study in red deer (Cervus elaphus) identified a previously unknown mechanism in ungulates--nocturnal hypometabolism--that contributed significantly to reduced energy expenditure, mainly during late winter. The relative contribution of these different mechanisms to seasonal adjustments of MR is still unknown, however. Therefore, in the study presented here we quantified for the first time the independent contribution of thermoregulation, LA and HI to heart rate (f(H)) as a measure of MR in a free-roaming large ungulate, the Przewalski horse or Takhi (Equus ferus przewalskii Poljakow). f(H) varied periodically throughout the year with a twofold increase from a mean of 44 beats min(-1) during December and January to a spring peak of 89 beats min(-1) at the beginning of May. LA increased from 23% per day during December and January to a mean level of 53% per day during May, and declined again thereafter. Daily mean subcutaneous body temperature (T(s)) declined continuously during winter and reached a nadir at the beginning of April (annual range was 5.8 degrees C), well after the annual low of air temperature and LA. Lower T(s) during winter contributed considerably to the reduction in f(H). In addition to thermoregulation, f(H) was affected by reproduction, LA, HI and unexplained seasonal variation, presumably reflecting to some degree changes in organ mass. The observed phase relations of seasonal changes indicate that energy expenditure was not a consequence of energy uptake but is under endogenous control, preparing the organism well in advance of seasonal energetic demands.
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