Reflections on mare behavior: Social and sexual perspectives

Among horse owners “mare-ishness” is easily recognized, if poorly defined, as generally awkward or recalcitrant behavior. In common with other mammals, however, horses do show a range of female-specific behavior patterns concerned particularly with sexual and maternal functions but also encompassing aspects of social peer interaction. Many of these behavior patterns are observed more easily in free living feral populations and are repressed at least partially under standard domestic management regimes. Feral horses form small stable breeding groups within which mares create linear dominance hierarchies based on the outcomes of aggressive encounters and positive social bonding. Rank within the social hierarchy may also influence reproductive success, with dominant mares having shorter foaling intervals and earlier foaling than lower ranking subordinate mares. Advantageous early foaling may reflect preferential mating by the group stallion based either simply on enhanced physical condition leading to earlier reproductive cycling in dominant mares or on an increased attractiveness to dominant mares by the stallion contingent on some aspect of dominance characteristics. The endocrine axis responsible for regulation of sexual behavior in the mare shows a degree of commonality with systems underlying social dominance. Both these behavior patterns are driven by a complex interaction of endocrine and neuro-endocrine factors sensitive to physical and social environmental cues and in feral free-living horses these are interlinked to an extent that may be under appreciated in domestic breeding populations.     

京海黄鸡公鸡性行为的观察研究

采用全天观察法,对京海黄鸡公鸡的性行为进行记录,获得京海黄鸡公鸡性行为的过程,总结京海黄鸡公鸡性行为规律,分析京海黄鸡公鸡个体间性行为的差异,并通过检测精液品质,研究京海黄鸡公鸡性行为与精液质量的关系;通过观察晴天与阴天时的性行为,分析不同的天气对京海黄鸡公鸡性行为的影响,结果显示：京海黄鸡性行为个体间差异不大,受天气的影响较为明显。本文通过对京海黄鸡公鸡性行为的观察研究,为进一步提高京海黄鸡公鸡自然交配的授精率和繁殖性能提供一定的科学依据。     

Male pseudohermaphroditism in a Labrador Retriever,and a review of mammalian sexual differentiation

CASE HISTORY: An 8-month-old Labrador Retriever was referred with a history of ambiguous external genitalia.

CLINICAL FINDINGS AND TREATMENT: Clitoromegaly within apparent vulval folds, and an adjacent subcutaneous mass were noticed on external examination. An intra-abdominal testicle, with epididymis and suspected vas deferens ducts, was found during exploratory celiotomy. Incision over the subcutaneous mass revealed the accompanying testicle. Clitoridectomy was performed and an os clitoris removed. Normal juvenile testes were diagnosed on histology of the gonads. Chromosomal studies revealed a normal 78, XY male chromosomal constitution. Due to the combination of a male karyotype (78, XY), the presence of testicular tissue in the gonads, and the appearance of the external genitalia, a diagnosis of male pseudohermaphroditism (MPH) was made.

CLINICAL RELEVANCE: This case presents the first report of MPH in a Labrador Retriever, and highlights the diagnostic steps recommended when confronted with a dog with ambiguous external genitalia.     

Male pseudohermaphroditism in a Labrador Retriever, and a review of mammalian sexual differentiation

CASE HISTORY: An 8-month-old Labrador Retriever was referred with a history of ambiguous external genitalia. CLINICAL FINDINGS AND TREATMENT: Clitoromegaly within apparent vulval folds, and an adjacent subcutaneous mass were noticed on external examination. An intra-abdominal testicle, with epididymis and suspected vas deferens ducts, was found during exploratory celiotomy. Incision over the subcutaneous mass revealed the accompanying testicle. Clitoridectomy was performed and an os clitoris removed. Normal juvenile testes were diagnosed on histology of the gonads. Chromosomal studies revealed a normal 78, XY male chromosomal constitution. Due to the combination of a male karyotype (78, XY), the presence of testicular tissue in the gonads, and the appearance of the external genitalia, a diagnosis of male pseudohermaphroditism (MPH) was made. CLINICAL RELEVANCE: This case presents the first report of MPH in a Labrador Retriever, and highlights the diagnostic steps recommended when confronted with a dog with ambiguous external genitalia.     

Differentiation of sexual behavior in cattle, sheep and swine

Gonadal steroid hormones influence an animal's sexual behavior through two primary means. During development they affect differentiation of the brain (primarily in males), and after puberty, circulating concentrations of steroids influence expression of sexual behaviors. In mammals, sexual behaviors of females are regarded as inherent (independent of steroids secreted by the developing ovary). Males, on the other hand, must undergo active differentiation that is brought about by actions of testicular steroids on the brain during discrete sensitive periods of early development. Sexual differentiation in mammals is referred to as processes of defeminization and masculinization. Defeminization is loss of behavioral traits inherent to females and occurs prenatally in sheep but postnatally in swine. Comparative data are lacking for cattle, but preliminary evidence indicates prenatal defeminization of sexual behaviors. Masculinization is acquisition of behavioral traits characteristic of males. Adult females of all three species show mounting behavior after prolonged treatment with testosterone, and in cattle and swine, there is, as yet, no strong evidence that males undergo masculinization of sexual behaviors. In sheep, limited observations are suggestive of two sensitive periods for masculinization; one prenatally and one postnatally. The earlier concept that differentiation of sexual behaviors in mammals occurs prenatally in species with a long gestation and postnatally in species with a short gestation must now be modified in view of recent findings in swine, in which defeminization occurs postnatally. Further studies will likely identify additional species differences with respect to differentiation of sexual behaviors.     

Rhythmic occurrence of sexual behavior and egg laying activity of Muscovy ducks

1. The existence of nychthemeral rhythms in the sexual behaviour of Muscovy ducks, kept under intensive husbandry conditions, was investigated. Studies were also carried out of the relationships between artificial insemination or natural mating times, oviposition time and female fertility. 2. Sexual activity (as measured by the number of complete matings observed) followed a trimodal pattern with peaks of activity in the second to third, 5th to 6th and 10th to 15th hours of the photoperiod. The last peak in sexual activity was the most important in both length and intensity. 3. The rhythm in sexual activity was entrained by the light-dark transition. 4. Female fertility was lowest during the period from 4 h before to 1 h after oviposition and greatest in the periods 8 to 4 h before and 1 to 7 h after oviposition. 5. The periods of maximum female fertility did not coincide with the periods of greatest sexual activity. More than 60% of all matings observed occurred during periods of intermediate female fertility. It would therefore appear that natural mating is not consistent with optimal female reproductive performance under artificial husbandry conditions. 6. The results are discussed in relation to a possible reduction in the number of males necessary in breeder populations in Muscovy ducks with a view to reducing the cost of producing ducklings.     

Effect of cortisol on luteinizing hormone release during sexual behavior in the boar.

The effect of inhibiting the rise in cortisol concentrations that occurs at copulation upon luteinizing hormone release was studied in seven adult boars. Plasma samples were collected for assay of luteinizing hormone, testosterone and cortisol on a control day and before, during and after exposure to an estrous sow. The area under the curve was used to evaluate hormone production and treatment effects were evaluated by a paired Student's t-test. The 11 beta-hydroxylase inhibitor, metyrapone, was used to suppress glucocorticoid hormone production. Cortisol concentrations increased significantly (p less than 0.05) after breeding compared to values on the control day while treatment with metyrapone prior to breeding prevented the cortisol increase (p greater than 0.05). Although luteinizing hormone production increased significantly after copulation in both breeding experiments, metyrapone pretreatment resulted in a reduction of luteinizing hormone secretion (p less than 0.05). Testosterone production was also reduced in boars pretreated with metyrapone. The results suggest that the increased levels of cortisol occurring at copulation may enhance luteinizing hormone release in boars.     

Effect of prepubertal versus postpubertal castration on sexual and aggressive behavior in male horses

Males horses castrated prepubertally occasionally display stallion-like sexual and aggressive behavior as adults. A retrospective survey of 140 horse owners was undertaken to compare the effectiveness of prepubertal versus postpubertal castration on objectionable sexual and aggressive behavior. Of 94 geldings that had been castrated prepubertally (before 2 years of age), 20% to 30% displayed stallion-like sexual behavior and aggression toward horses, and 5% were aggressive toward people in a stallion-like manner. This occurrence of problem behavior was not significantly different from the occurrence of these behavioral patterns in 46 geldings that had been castrated as stallions (over 3 years of age). When castration was examined as a therapeutic measure to eliminate sexual and aggressive behavior in stallions in which the behavior had become objectionable, it was found that the operation was effective on sexual behavior and aggression toward people in 60% to 70% of horses and on aggression toward horses in 40% of horses.     

Effects of various steroid milieus or physiological states on sexual behavior of Holstein cows.

Two experiments were conducted to determine how steroid milieus and pregnancy affect sexual behavior. Experiment 1 was arranged as a Latin square with five ovariectomized cows and five steroid milieus: no steroid (N); progesterone (P4); estradiol benzoate (EB); P4 + EB; and P4 followed by EB (P4:EB). Progesterone was administered via pessary (2 g of P4) for 5 d and EB was injected (1 mg i.m.) on the day before a test day. On a test day, cows were exposed for four 30-min periods, twice each with a tied or a loose estrual test cow (prepared using P4:EB). Sexual behaviors recorded were attempted mounts, successful mounts, front mounts, stands, head butts, chin rests, and vulvar sniffs. Cows exhibited more (P less than .05) sexual behavior during periods with the loose estrual test cow than with the tied estrual test cow. Cows receiving P4 alone ranked lowest among treatments for each behavior, whereas cows receiving EB or P4:EB ranked highest or second-highest. Progesterone prevented stands in cows given P4 + EB, but these cows displayed mounting behaviors similar to those of cows given EB and P4:EB. Cows given P4 + EB were similar to those given N for most behaviors. In Exp. 2, 118 intact, lactating cows were observed in groups of four or five for mounting of estrual test cows during 24, 30-min observation periods on 8 d over 2 yr. The design was an incomplete block with physiological state, parity, estradiol, progesterone, and a calculated estrogen:progesterone ratio included in the model. Each block included one or two cows at 23 +/- .8 d after insemination, divided retrospectively into one pregnant and two non-pregnant groups (low [less than 1 ng/mL] vs high progesterone), and other cows at 89 +/- 1.0, 152 +/- 1.2, and 234 +/- 1.7 d of gestation (six physiological groups). Most cows were observed once, but 27 cows were included twice during 2 yr. Only 60% of the 118 cows made attempted or successful mounts even though estrual test cows were always receptive. Physiological state was not associated with amount of mounting because very active (greater than or equal to five attempts) and inactive cows were represented in all physiological groups. The estrogen:progesterone ratio on test day accounted for small, but significant, variation in mounting behavior. For cows observed on two different days, correlations between successive observations were .46 for attempted mounts, .78 for successful mounts, and .71 for total mounts.(ABSTRACT TRUNCATED AT 400 WORDS)     

Moxidectin interference on sexual behavior, penile erection and hypothalamic GABA levels of male rats

The moxidectin (MXD) is an antiparasitic drug used in domestic animals. The mechanism of action, in mammals, involves GABA, a neurotransmitter with an important role in the sexual behavior control. Presently, the effects of 0.2 mg/kg therapeutic dose were studied on sexual behavior, sexual motivation, penile erection and central GABA levels. Sexual behavior results showed increased latencies to the first mount and intromission as well as in inter-intromission interval; a reduction in total mounts was detected on the drug post-treatment. No difference was observed between sexual motivation of control and experimental animals. MXD treatment reduced penile erection and hypothalamic GABA levels. The results suggest that MXD reduced sexual behavior and penile erection by an action on the hypothalamic GABA system. Probably, the lack of effects in the motivational test and the increased mount and intromission latencies as well as decreased total mounts could be explained as a consequence of reduced male rat erection process.     

Relationship between sexual behavior classifications of rams and lambs sired in a competitive breeding environment

The objectives for this study were to 1) determine the relationship between sexual performance class and lambs sired in a competitive mating environment, and 2) determine whether the male-oriented ram test is valid. Fifteen 2- to 3-yr-old whitefaced rams classified as female-oriented, with high or low sexual performance, or classified as male-oriented were used in a multiple-sire breeding arrangement. Five groups of approximately 200 ewes each were exposed for 21 d to 3 rams per group consisting of 1 ram from each class. Rams were blocked for sexual class, and those with close genetic relationship were assigned to different pens. Genomic DNA was prepared from blood collected from 15 rams, 934 ewes, and 1,757 lambs. Up to 4 microsatellite markers were used to determine a lamb's sire. Of 884 ewes with identifiable lambs (known sires), 178 ewes had single lambs, 408 had multiples sired by 1 ram, and 298 had multiples sired by more than 1 ram. The sexual partner preference test used to identify male-oriented rams did not absolutely reflect their sexual performance during competitive breeding. In contrast to only mounting and servicing males in preference tests before breeding, male-oriented rams sired 480 lambs from 330 ewes. Serving capacity tests predicted sexual performance of high and low sexual performance rams. High performance rams impregnated more ewes (499 vs. 258; P < 0.05) and sired more lambs (756 vs. 357; P < 0.05) than did low performance rams, respectively. Low performance and male-oriented rams did not differ for ewes impregnated or lambs sired. We conclude that 1) sexual partner preference tests used to classify male-oriented rams were not absolute in reflecting their breeding performance in a competitive breeding environment; 2) serving capacity tests predicted that high performance rams would breed more ewes than low performance rams and sire more lambs than either low performance or male-oriented rams; and 3) under the conditions of this study, low performance and male-oriented rams did not have an adverse impact on the overall breeding outcome. Combined, low performance and male-oriented rams sired 81 more lambs than did high performance rams, but this required twice as many rams to obtain approximately equal breeding results. Therefore, we suggest that serving capacity tests should be used to select high performance rams, reduce number of rams with marginal sexual performance, and make decisions on ram numbers needed.     

Intravaginal impedance and sexual behavior of ovariectomized goats given estrogen alone or in combination with progesterone

Intravaginal impedance (IVI) fluctuates during the goat estrous cycle. To understand which ovarian steroids are responsible for IVI changes and whether IVI variations are associated with precopulatory and copulatory behaviors, 8 ovariectomized females were assigned to 4 treatments in a 4 x 4 Latin square replicated over four 8-d periods. The treatments were as follows: progesterone plus estradiol-17beta (P4 + E2), oil plus estradiol-17beta (E2), progesterone plus oil (P4), or oil (OIL). Daily IVI measurements at the vagino-cervical junction were taken at 1 and 70 KHz. Progesterone was given on d 2 and 3. Estradiol was given in the evening of d 5. On d 1 to 8, goats were group-exposed to a sexually experienced male and observed for the expression of sexual behaviors. On d 6 and 7, IVI was less when goats received P4 + E2 or E2 compared with goats given P4 or OIL (P < 0.05). Impedance measured at 1 kHz tended to remain lower on d 8 in P4 + E2-treated females compared with those given P4 or OIL (P < 0.055). Like previous results, P4 + E2 or E2 treatment induced behavioral estrus; 5 of 8 P4 + E2-treated and 5 of 8 E2-treated females were sexually receptive on d 6. On d 7, although IVI remained low and 2 of 8 P4 + E2-treated goats and 4 of 8 E2-treated goats remained sexually receptive, no additional females were in estrus. No IVI decreases and no estrous behavior were observed in goats given P4 or OIL. This experiment demonstrated that E2 initiates the periestrous drop in IVI, and P4 may delay baseline return.     

Early embryonic administration of xenoestrogens alters vasotocin system and male sexual behavior of the Japanese quail

The copulatory behavior and the parvocellular vasotocin (VT) system of the nucleus of the stria terminalis (BST) are sexually dimorphic in the Japanese quail. Embryonic administration of estradiol benzoate (EB) induces an organizational effect determining the disappearance of such a dimorphism (male shows behavior and cerebral phenotype of the female). The VT parvocellular system can therefore be considered an accurate marker of the sexual differentiation of brain circuits and a very sensitive indicator of the activity of estrogen-like substances on neural circuits. To test this hypothesis we administered diethylstilbestrol (DES), a powerful synthetic xenoestrogen, genistein (GEN), a phytoestrogen produced by soy, and bisphenol A (BPA). After 3 days of incubation, quail eggs were injected with vehicle, EB, DES, GEN or BPA. Administration of BPA caused an early blockage of development and no further analyses were done on the BPA groups. At puberty, the copulatory behavior of EB- or DES-treated male quail was totally abolished, whereas only the highest doses of GEN determined a significant decrease of the behavior. After the tests, the animals were sacrificed and perfused. The fractional area (FA) covered by VT immunoreactivity was analyzed in BST, medial preoptic nucleus, and lateral septum by computerized image analysis. The FA was significantly reduced after treatment with EB, DES and GEN at high doses. These results confirm that the sexually dimorphic VT system of the Japanese quail is a sensible indicator of the effects of xenoestrogens at the level of the central nervous system.     

Ivermectin impairs sexual behavior in sexually naïve, but not sexually experienced male rats

Ivermectin (IVM) is an antiparasitic drug, widely used in domestic animals. In mammals, IVM act as a GABA agonist. This neurotransmitter has an important role in the regulation of sexual behavior. Thus, this study sought to investigate the effects of various medically relevant doses IVM on the sexual behavior of male rats. In particular, we also wished to examine if previous sexual experience modulated responses to IVM. In the first experiment, the sexual behavior of inexperienced male rats was analyzed after they received 0.2, 0.6, 1.0 or 2.0 mg/kg IVM, 15 min prior to behavioral testing. In the second experiment, the effects of four previous sexual experiences on IVM treated rats (1.0 or 2.0 mg/kg, 15 min prior to the 5th session) were assessed. The standard therapeutic dose (0.2 mg/kg) did not impair the sexual behavior of inexperienced male rats. At a more concentrated dose (0.6 mg/kg), which is still within the therapeutic range, the appetitive phase of sexual behavior of inexperienced male rats was impaired. Likewise, 1.0 mg/kg impaired the appetitive phase. Previous sexual experience blocked almost entirely this sexual impairment, suggesting that previous sexual experience exerts a positive effect in attenuating the sexual impairment produced by IVM treatment. Therefore, the standard therapeutic dose of IVM can be used without producing side effects on sexual behavior. Use of more concentrated therapeutic doses is not recommended during reproductive periods, unless the animals have had previous sexual experience.