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Khalil, A. M., Murakami, N., & Kaseda, Y. (1998). Relationship between plasma testosterone concentrations and age, breeding season and harem size in Misaki feral horses. J Vet Med Sci, 60(5), 643–645.
Abstract: Jugular vein blood samples were collected from 23 young and sexual mature feral stallions to examine the relationship between plasma testosterone concentration and age, breeding season or harem size. Testosterone concentration increased with the age of the stallions until they formed their own harems, at about 4 to 6 years old. Seasonal variations in testosterone concentrations were observed, and found to be significantly higher (P<0.001) throughout the breeding season than non-breeding season, from 3 years of age. Testosterone levels were correlated with harem size for individual stallions. It can be inferred from these results that there is a relationship between plasma testosterone concentration and age, breeding season and harem size.
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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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Kirkpatrick, J. F., Wiesner, L., Kenney, R. M., Ganjam, V. K., & Turner, J. W. (1977). Seasonal variation in plasma androgens and testosterone in the North American wild horse. J Endocrinol, 72(2), 237–238.
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Kirkpatrick, J. F., Vail, R., Devous, S., Schwend, S., Baker, C. B., & Wiesner, L. (1976). Diurnal variation of plasma testosterone in wild stallions. Biol Reprod, 15(1), 98–101.
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Dzieweczynski, T. L., Eklund, A. C., & Rowland, W. J. (2006). Male 11-ketotestosterone levels change as a result of being watched in Siamese fighting fish, Betta splendens. Gen Comp Endocrinol, 147(2), 184–189.
Abstract: This study investigated the effects of nesting status and the presence of an audience on 11-ketotestosterone (11KT) levels in male Siamese fighting fish, Betta splendens. Prior studies have demonstrated that both nesting status, an indicator of territory-holding power and reproductive state, and the sex of a conspecific audience lead to differences in male behavior during aggressive encounters. Since behavioral changes have already been demonstrated, we chose to investigate whether 11KT levels were also influenced by nesting status and audience presence as 11KT both stimulates, and is stimulated by, reproductive and aggressive behaviors in male teleosts. Male 11KT levels were measured from water samples taken from containers holding fish both before and after interaction. Males interacted under three treatment conditions: no audience, female audience, and male audience. Within these treatments were two nest paradigms: both males had nests or neither male had a nest. 11KT levels varied depending on nesting status and audience type. In general, 11KT levels were lower in interacting males when a female audience was present or when males had nests. Overall, 11KT showed increases or decreases as aggression increased or decreased, as shown by already established behavioral findings [see Dzieweczynski T.L., Green T.M., Earley R.L., Rowland W.J., 2005. Audience effect is context dependent in Siamese fighting fish, Betta splendens. Behav. Ecol. 16, 1025-1030; Doutrelant, C., McGregor, P.K., Oliveira, R.F., 2001. Effect of an audience on intrasexual communication in male Siamese fighting fish (Betta splendens). Behav. Ecol. 12, 283-286.]. Our results suggest that 11KT levels are influenced by reproductive status, as indicated by nest ownership, and audience presence and are most likely modulated by territorial behavior and social environment.
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Chaudhuri, M., & Ginsberg, J. R. (1990). Urinary androgen concentrations and social status in two species of free ranging zebra (Equus burchelli and E. grevyi). J Reprod Fert, 88, 127–133.
Abstract: In both species of zebra, breeding males had higher urinary androgen concentrations (ng androgens/mg Cr) than did non-breeding bachelor males (30.0 +/- 5.0 (N = 9) versus 11.4 +/- 2.8, (N = 7) in the plains zebra; 19.0 +/- 2.2 (N = 17) versus 10.7 +/- 1.2 (N = 14) in the Grevy's zebra). In the more stable family structure of the plains zebra (single male non-territorial groups) variations in androgen concentrations could not be ascribed to any measured variable. In the Grevy's zebra, androgen values were significantly lower in samples taken from territorial (breeding) males which had temporarily abandoned their territories (N = 4) and the urinary androgen concentration for a male on his territory was negatively correlated with the time since females last visited the territory.
Keywords: zebra; testosterone; androgens; territoriality; social behaviour; dominance
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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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Ganswindt, A., Palme, R., Heistermann, M., Borragan, S., & Hodges, J. K. (2003). Non-invasive assessment of adrenocortical function in the male African elephant (Loxodonta africana) and its relation to musth. Gen Comp Endocrinol, 134(2), 156–166.
Abstract: Adult male elephants periodically show the phenomenon of musth, a condition associated with increased aggressiveness, restlessness, significant weight reduction and markedly elevated androgen levels. It has been suggested that musth-related behaviours are costly and that therefore musth may represent a form of physiological stress. In order to provide data on this largely unanswered question, the first aim of this study was to evaluate different assays for non-invasive assessment of adrenocortical function in the male African elephant by (i) characterizing the metabolism and excretion of [3H]cortisol (3H-C) and [14C]testosterone (14C-T) and (ii) using this information to evaluate the specificity of four antibodies for determination of excreted cortisol metabolites, particularly with respect to possible cross-reactions with androgen metabolites, and to assess their biological validity using an ACTH challenge test. Based on the methodology established, the second objective was to provide data on fecal cortisol metabolite concentrations in bulls during the musth and non-musth condition. 3H-C (1 mCi) and 14C-T (100 microCi) were injected simultaneously into a 16 year old male and all urine and feces collected for 30 and 86 h, respectively. The majority (82%) of cortisol metabolites was excreted into the urine, whereas testosterone metabolites were mainly (57%) excreted into the feces. Almost all radioactive metabolites recovered from urine were conjugated (86% 3H-C and 97% 14C-T). In contrast, 86% and >99% of the 3H-C and 14C-T metabolites recovered from feces consisted of unconjugated forms. HPLC separations indicated the presence of various metabolites of cortisol in both urine and feces, with cortisol being abundant in hydrolysed urine, but virtually absent in feces. Although all antibodies measured substantial amounts of immunoreactivity after HPLC separation of peak radioactive samples and detected an increase in glucocorticoid output following the ACTH challenge, only two (in feces against 3alpha,11-oxo-cortisol metabolites, measured by an 11-oxo-etiocholanolone-EIA and in urine against cortisol, measured by a cortisol-EIA) did not show substantial cross-reactivity with excreted 14C-T metabolites and could provide an acceptable degree of specificity for reliable assessment of glucocorticoid output from urine and feces. Based on these findings, concentrations of immunoreactive 3alpha,11-oxo-cortisol metabolites were determined in weekly fecal samples collected from four adult bulls over periods of 11-20 months to examine whether musth is associated with increased adrenal activity. Results showed that in each male levels of these cortisol metabolites were not elevated during periods of musth, suggesting that in the African elephant musth is generally not associated with marked elevations in glucocorticoid output. Given the complex nature of musth and the variety of factors that are likely to influence its manifestation, it is clear, however, that further studies, particularly on free-ranging animals, are needed before a possible relationship between musth and adrenal function can be resolved. This study also clearly illustrates the potential problems associated with cross-reacting metabolites of gonadal steroids in EIAs measuring glucocorticoid metabolites. This has to be taken into account when selecting assays and interpreting results of glucocorticoid metabolite analysis, not only for studies in the elephant but also in other species.
Keywords: Adrenal Cortex/*metabolism/secretion; Adrenal Cortex Function Tests/methods/*veterinary; Adrenocorticotropic Hormone/physiology; Animals; Carbon Isotopes/diagnostic use; Chromatography, High Pressure Liquid/veterinary; Elephants/*metabolism/urine; Feces/*chemistry; Glucocorticoids/analysis/urine; Hydrocortisone/*analysis/diagnostic use/urine; Immunoenzyme Techniques/methods/veterinary; Male; Reproduction/physiology; Sexual Behavior, Animal/physiology; Stress, Psychological/diagnosis/*physiopathology; Testosterone/*analysis/diagnostic use/urine
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Kralj-Fiser, S., Scheiber, I. B. R., Blejec, A., Moestl, E., & Kotrschal, K. (2007). Individualities in a flock of free-roaming greylag geese: behavioral and physiological consistency over time and across situations. Horm Behav, 51(2), 239–248.
Abstract: The concept of personality implies individual differences in behavior and physiology that show some degree of repeatability/consistency over time and across contexts. Most studies of animal personality, particularly studies of individuals' variation in physiological mechanisms, have been conducted on selected individuals in controlled conditions. We attempted to detect consistent behaviors as well as physiological patterns in greylag ganders (Anser anser) from a free-roaming flock living in semi-natural conditions. We tested 10 individuals repeatedly, in a handling trial, resembling tests for characterization of “temperaments” in captive animals. We recorded the behavior of the same 10 individuals during four situations in the socially intact flock: (1) a “low density feeding condition”, (2) a “high density feeding condition”, (3) a “low density post-feeding situation” and (4) while the geese rested. We collected fecal samples for determination of excreted immuno-reactive corticosterone (BM) and testosterone metabolites (TM) after handling trials, as well as the “low density feeding” and the “high density feeding” conditions. BM levels were very highly consistent over the repeats of handling trials, and the “low density feeding condition” and tended to be consistent over the first two repeats of the “high density feeding condition”. Also, BM responses tended to be consistent across contexts. Despite seasonal variation, there tended to be inter-test consistency of TM, which pointed to some individual differences in TM as well. Aggressiveness turned out to be a highly repeatable trait, which was consistent across social situations, and tended to correlate with an individual's resistance during handling trials. Also, “proximity to the female partner” and “sociability” – the average number of neighboring geese in a close distance while resting – were consistent. We conclude that aggressiveness, “affiliative tendencies” and levels of excreted corticosterone and testosterone metabolites may be crucial factors of personality in geese.
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Baum, M. J. (2006). Mammalian animal models of psychosexual differentiation: when is 'translation' to the human situation possible? Horm Behav, 50(4), 579–588.
Abstract: Clinical investigators have been forced primarily to use experiments of nature (e.g., cloacal exstrophy; androgen insensitivity, congenital adrenal hyperplasia) to assess the contribution of fetal sex hormone exposure to the development of male- and female-typical profiles of gender identity and role behavior as well as sexual orientation. In this review, I summarize the results of numerous correlative as well as mechanistic animal experiments that shed significant light on general neuroendocrine mechanisms controlling the differentiation of neural circuits controlling sexual partner preference (sexual orientation) in mammalian species including man. I also argue, however, that results of animal studies can, at best, provide only indirect insights into the neuroendocrine determinants of human gender identity and role behaviors.
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