Reproductive behavior of the stallion

The reproductive behavior of semi-wild and feral horses is described briefly in order to provide a background on the reproductive patterns and the specific behavioral elements comprising courtship and mating in the horse. The reproductive behavior of the domestic stallion is discussed at length as are various breeding conditions and normal breeding performance. Finally, sexual behavior problems are considered.     

Sexual behavior of mares

Behavior during the estrous phase of the ovulatory cycle of the mare is analogous in most ways to that of estrous females of other species. Proceptive behaviors bring the mare into the proximity of the male and attract his attention. Positioning facilitates mounting, intromission, and ejaculation. Estrous signs appear to be more intense in the few days prior to ovulation than during the transition periods that separate the recurring estrous and diestrous phases. Sexual behavior is absent during diestrus. Detection of estrus in mares is problematic in that it requires the presence (or at least facsimile acoustic or tactile stimuli) or a stallion. Unexplained conditions such as silent or subestrus make reliable determinations even more difficult. The choice of methods for distinguishing estrous from nonestrous mares depends, among other factors, upon characteristics of individual mares, as well as the number of mares to be evaluated. Detection of true estrus--that is, sexual behavior that is associated with follicular growth and ovulation--is confounded by the occurrence of estrus-like behavior independent of gonadal stimulation. Seasonally anovulatory and ovariectomized mares may regularly solicit and accept copulation despite low or even no circulating levels of ovarian steroids. Instead, endocrine support of sexual behavior in these mares appears to be provided by adrenal cortical androgens and/or estrogens. A smaller percentage of pregnant mares may also show some signs of estrus but are not likely to be receptive to mating. The ability of the mare to dissociate sexual behavior from ovulation is unique among infraprimate species studied. The display of estrous behavior and willingness to copulate, even at times when conception is impossible, may have functional significance in promoting band cohesiveness in wild and feral populations. Even without copulation, continued motivation to remain in proximity to the band stallion may contribute to the stability of the social unit outside the ovulatory season.     

Comparative reproductive biology of North American feral horses

Recent studies have suggested that various aspects of reproductive biology are strikingly different among the many herds of feral horses and ponies (E. caballus) in North America. The greatest differences include (1) sharply seasonal versus year-round mating and foaling patterns, (2) mare behavior at the time of parturition, (3) forced copulation and incest, (4) exclusive breeding by a single harem stallion vs breeding by two or more stallions, and (5) fecundity. The causes for these differences are discussed in terms of genetic origins of the various herds, the length of time each herd has been in a free-roaming state and subject to the forces of natural selection, the ecology of the ranges inhabited by these horses, population density, and sex ratios.     

Injuries and blemishes in a semi-feral herd of ponies

With the goal of characterizing the type and degree of injuries and blemishes incurred by horses living under natural social conditions, all members of a semi-feral herd of ponies were inspected on 4 occasions over a period of 28 months. Two occasions were during breeding/foaling season and 2 during non–breeding/foaling season months. On each occasion, each animal was examined (N = 47-65) and all injuries or blemish marks were recorded. Based on the types and number of injuries or blemishes, an injury/blemish grade was assigned for each inspection, ranging from 1 for no blemishes to 6 for more than 1 open wound. Almost all injuries and blemishes recorded were extremely mild. Only 12 of the 213 inspections and 14 of the 858 total injuries and blemishes involved wounds affecting tissues other than hair and skin.Males had significantly greater mean injury/blemish grade than females (P < .0001). For all social categories and ages, the mean injury/blemish grade was significantly greater in the breeding/foaling season months than during the non–breeding/foaling season (P < .0001).All foals were blemish-free at the non–breeding/foaling season inspections (n = 24) and relatively blemish-free (9 of 14) at the breeding/foaling season inspections. The rump and the barrel areas were the most common site of injuries or blemishes, both as a percentage of the total injury count and as a percentage of inspections involving those areas. These findings are consistent with the seasonal and gender patterns of aggressive behavior seen in this herd. Compared with reports of truly feral horse populations, it appears that injuries are fewer and less severe in this semi-feral herd. In addition, there appear to be fewer leg injuries in this semi-feral herd than has been described for truly feral horse populations.     

Effect of book size and age of mare and stallion on foaling rates in thoroughbred horses

The entire population data for Thoroughbred horses in the United States, Canada and Puerto Rico registered with the Jockey Club of America were utilized to study the effect of stallion book size, age of mare and age of stallion on live foal percentage for the 1987 and 1988 breeding seasons. There were 17,260 stallions bred to 179,009 mares, with a resulting overall live foaling rate of 58.1%. There was an increase in live foal percentage as book size increased (P<0.001) and a decrease in live foal percentage as mare age increased (P<0.0001). Stallion age had no effect on live foal percentage.     

Reproductive phenology of Creole horses in Ecuador in the absence of photoperiod variation: The effects of forage availability and flooding affecting body condition of mares

Horse reproduction tends to be seasonal. The main adjusting factor in their original temperate ranges is photoperiod variation, although it is absent in equatorial areas where horses were introduced by European colonizers. Hence, dates of reproduction in these areas may be influenced by factors affecting mares’ conditions and the success of foaling. Here we study reproductive timing in Creole horses in Ecuador reared in an extensive production system. We found that foaling peaked in August. Mares’ conditions showed one peak in June‐July, before the start of the breeding season, and another in December, and it was highly variable along the year. Mares’ conditions increased after a period of vegetation growth and thus appeared negatively associated with the increment of grass greenness (normalized difference vegetation index data). Seasonal flooding of some pasturelands during March and April appeared to seriously impair mares’ conditions and probably influenced the timing of foaling toward the dry season. Our results evidenced that horse breeding in these equatorial areas tended to be seasonal and point to some key factors that influence phenology by affecting body condition of mares, which may have implications for horse biology and management.     

Fertility control as a management tool for feral horse populations

This report addresses the basic reproductive biology of the North American feralhorse. With a basis of population records to establish a framework, experimental studies on specific and seasonal aspects of sexual behavior and social behavior are discussed with special attention to elimination marking (scent marking) behavior in stallions and estrus behavior in mares. In addition, the physiological basis for the hormonal control of reproduction in male and female feralhorses is presented, with discussion of specific experiments. Definite and correlated seasonal patterns of plasma testosterone and elimination marking behavior in stallions are examined, and hormonal patterns and events of the estrus cycle of the mare are presented. These aspects of feral horsereproduction are then interpreted with attention to integration of processes, and comparisons between the feral and domestic horse are made.     

Some Factors Associated With Fertility of Thoroughbred Stallions

The results of 3 years (2005–2007) of observations and mating (5,646 estrous cycles of 3,788 mares bred to 1 of 15 stallions) at one Thoroughbred breeding farm in central Kentucky were analyzed by a multiple logistic regression model using Bayesian statistics to evaluate the relationship between data entries (factors) and pregnancy outcomes. Factors found to be significantly (P < .05) associated with pregnancy outcome included stallion (one stallion had lower OR for pregnancy higher odds ratio [OR] for pregnancy, and one had, than other stallions), date of mating (OR for pregnancy declined slightly in May – July), mare age (OR for pregnancy were higher for mares <13 years old, and lower for mares >18 years old), mare beginning status (foaling mares had a higher OR for pregnancy), mating on foal heat (lowered OR for pregnancy), mating of the day for the stallion (OR for pregnancy was 4.16 times lower for fifth compared with first mating of day), reinforcement breeding (increased OR for pregnancy), dismount semen neutrophil score (lowered OR for pregnancy when neutrophils were present in dismount semen samples), and tranquilization before breeding (lowered OR for pregnancy in foaling and barren mares). The influence of dismount sample sperm motility scores on OR for pregnancy was weak, so motility scores were not included in the final logistic regression model. The majority of variation in pregnancy outcome was because of mare factors, with only approximately one-third of the variation in fertility explained by stallion.     

Investigating equine ingestive, maternal, and sexual behavior in the field and in the laboratory.

Some of the techniques that may be used to study social, reproductive, and ingestive behavior in horses are described in this paper. One of the aspects of equine social behavior is the dominance hierarchy or patterns of agonistic behavior. Paired or group feeding from a single food source may be used to determine dominance hierarchies quickly. Focal animal studies of undisturbed groups of horses may also be used; this method takes longer, but may reveal affiliative as well as agonistic relationships among the horses. Reproductive behavior includes flehmen, the functional significance of which can be determined using combinations of field observations of harem groups and laboratory studies of stallions exposed to female urine or feces in the absence of the donor mare. Ingestive behavior may include food, salt, or water intake. Direct and indirect measurements of intake can be made and used to answer questions regarding the ability of horses to control their energy intake when the diet is diluted, the effect of feral equids on the ecology of an area, and the abilities of horses to compensate for dehydration and hypovolemia.     

Reproductive performance and factors that decrease pregnancy rate in heavy draft horses bred at the foal heat

Reproductive performance and factors that decrease the pregnancy rate in heavy draft horses bred at the foal heat were investigated. The study was based on a total of 422 mares varying from 4 to 20 years in age, which had foaled during the period 1991 to 1994. Over the four years, the percentage of the mares bred at the foal heat was 62.6% and the pregnancy rate was 60.2%. The pregnancy rate following breeding at the foal heat in 1991 (67.7%) was significantly higher than that in 1992 (44.1%). Significant correlation was observed between the pregnancy rate and the mean placental retention time and also between the pregnancy rate and the mean temperature during the winter season. When compared on a monthly basis, the pregnancy rate following breeding at the foal heat in February was 53.3% and in May, 48.2%. The mean number of covers per mare at the foal heat was 2.3; this figure decreased to 1.8 for those bred in May. The mean number of days from parturition to the first breeding date was 11.3 days and the number of days from parturition to the last breeding day at the foal heat was 14.0 days. The number of days from parturition to the last breeding day at the foal heat decreased to 12.5 days in May. The latter may have been influenced by factors involving the stallion. When compared on a monthly basis, the mean placental retention time was longest in January (219 min.) following which it decreased gradually up to mid-summer. The mean placental retention time of mares which were not bred at the foal heat was significantly longer (197 min.) than in mares bred at the foaling heat (143 min.) and also in mares that conceived following breeding at the foal heat (135 min.). The pregnancy rate was less in mares bred shortly after parturition. The findings of this study would suggest that certain factors decrease the pregnancy rate in mares bred at the foal heat. The stress of low temperature in winter, prolonged placental retention time, breeding too early after parturition and factors pertaining to individual stallions may be involved. Expulsion of the placenta as soon as possible after foaling and appropriate breeding timing are significant factors that can influence the pregnancy rate following breeding at the foal heat.     

Reproductive performance of Thoroughbred mares in the Waikato region of New Zealand: 2. Multivariable analyses and sources of variation at the mare,stallion and stud farm level

Abstract

AIM: The objective of this study was to utilise multivariable statistical methods appropriate for clustered data to identify mare-related explanatory variables that significantly affected the reproductive performance of Thoroughbred mares in the Waikato region of New Zealand. In addition, we aimed to determine the relative contribution of the mare, stallion and stud farm to reproductive performance.

METHODS: A prospective cohort study was performed involving five stud farms in the Waikato region of New Zealand during three consecutive breeding seasons (2006–2008). A total of 1,482 individual mares contributed 2007 mare years and 3,402 oestrous cycles over the three breeding seasons. Reproductive performance was measured using three parameters; (a) first-cycle pregnancy rate (FCPR), (b) end-of-season pregnancy rate (SPR), and (c) the start-of-mating to conception interval.

RESULTS: When controlled for the effects of serving stallion, stud farm and year of study the only significant mare-related variables included in the final models of FCPR, SPR and conception interval were the age of the mare and her reproductive status (classified as dry or foaling). Advancing mare age significantly reduced reproductive performance regardless of reproductive status and foaling mares had significantly poorer reproductive outcomes compared with dry mares when controlled for age. For each additional increase in year of age, the FCPR was reduced by a factor of 0.94 (95% CI=0.92–0.96) and the SPR was reduced by a factor of 0.91 (95% CI=0.88–0.93). Mares older than 14 years of age took longer to conceive after the start-of-mating compared with younger mares. The daily hazard of conception for mares 14 years and older was 0.64 (95% CI=0.47–0.83) times less than mares younger than 9 years of age. Determining the relative contribution of the mare, stallion and stud farm to the FCPR indicated that 95.9% of the variation was at the mare level, 4.1% was at the stallion level and 0% was at the stud farm level. For the SPR the variance components indicated that 92.5% of the variation was at the mare level, 6.7% was at the stallion level and 0.8% was at the stud farm level.

CONCLUSIONS: The reproductive performance of Thoroughbred mares in the Waikato region of New Zealand is influenced by two main mare-related factors; the age of the mare and her reproductive status (dry or foaling). The majority of variation in reproductive performance was associated with mare-level factors and the contribution of the stallion and stud farm was relatively minor.     

Method of predicting stallion to mare ratio for natural and artificial insemination programs

A method of predicting a stallion to mare ratio for horses entering either natural service or artificial insemination programs was discussed. Limiting factors for stallions in hand-breeding situations were age, libido and physical ability to cover mares presented for breeding. Other limiting factors for stallions involved in artificial breeding programs were scrotal width, daily sperm output, and the number of live spermatozoa inseminated per mare. Considerations were given for the seasonal effect upon pregnancy rate of the mare, but no considerations were made for seasonal effect upon the stallion. This method must only be applied to stallions that have undergone a thorough breeding soundness examination. It must be emphasized that it served only as a guide; the calculated values were merely useful estimates rather than firm numbers. The multitude of uncontrollable factors involved in breeding farm management, especially with regard to the mares that are presented for breeding were influences that could not be directly considered by this method.     

A cross-sectional survey of Thoroughbred stud farm management in the North Island of New Zealand

AIM: To obtain initial baseline data on the management of Thoroughbred stud farms in the North Island of New Zealand.

METHODS: Data on the management of Thoroughbred stud farms were collected from a sample of 22 stud farms located in the south Auckland/Waikato region (n=15) and lower North Island (n=7) of New Zealand, using a face-to-face survey. The studmaster provided information on the size, scope and management of the farms during the 2004/2005 breeding season. Analysis was based on the location of the farm and size of the breeding operation (number of resident mares).

RESULTS: Effective farm size ranged from 20 to 526 ha and averaged 167 (standard error (SE) 36) and 88 (SE 49) ha in the south Auckland/Waikato and lower North Island areas, respectively. Some farms in the Auckland/Waikato region stood shuttle stallions. The median number of stallions per farm was three (range 0.9), and the median mare-to-stallion ratio was 43 (range 10.250). The farms had a mean of 50 (range 7.180) wet mares and 21 (range 0.100) dry mares. The number of mares per breeding stallion increased with increasing size of breeding operation (p=0.04), being 28 (95% confidence interval (CI) = 10.56) vs 40 (95% CI=16.74) vs 74 (95% CI=44.113) for moderate (≤70 mares), medium (90–199 mares) and large (≥200 mares in total) operations, respectively. Seventy-one percent of farms aimed to breed dry mares early in the breeding season, and used a combination of lights, hormone therapy, and rising plane of nutrition to achieve this.

Foaling took place in foaling paddocks monitored using a night foaling attendant (17/22) or with foaling alarms (5/22). At birth, 17/22 studmasters routinely administered antibiotics, 14/22 administered tetanus antitoxin, 9/22 administered an enema to foals, and 2/22 did not routinely administer prophylactic treatments. Weaning occurred at 5 (range 3.7–7) months of age, and foals were confined to a box for 1–2 weeks on 16/22 farms. Weaned foals were drenched with anthelmintics every 7 (range 4–9) weeks, and were fed 2.9 (range 1–6) kg of concentrate feed while at pasture until intensive management associated with preparation of the horses for auction began 13 (range 6–20) weeks before the yearling sales. Eight farms weighed the weanlings, at least monthly, to monitor growth.

CONCLUSIONS AND CLINICAL RELEVANCE: The management of Thoroughbred horses was relatively consistent throughout the regions surveyed. Utilisation of breeding stallions tended to be more efficient on the larger stud farms in the south Auckland/Waikato region. Even though foals are grown at pasture they are often provided with large quantities of concentrate feed.     

A cross-sectional survey of Thoroughbred stud farm management in the North Island of New Zealand

AIM: To obtain initial baseline data on the management of Thoroughbred stud farms in the North Island of New Zealand. METHODS: Data on the management of Thoroughbred stud farms were collected from a sample of 22 stud farms located in the south Auckland/Waikato region (n=15) and lower North Island (n=7) of New Zealand, using a face-to-face survey. The studmaster provided information on the size, scope and management of the farms during the 2004/2005 breeding season. Analysis was based on the location of the farm and size of the breeding operation (number of resident mares). RESULTS: Effective farm size ranged from 20 to 526 ha and averaged 167 (standard error (SE) 36) and 88 (SE 49) ha in the south Auckland/Waikato and lower North Island areas, respectively. Some farms in the Auckland/Waikato region stood shuttle stallions. The median number of stallions per farm was three (range 0-9), and the median mare-to-stallion ratio was 43 (range 10-250). The farms had a mean of 50 (range 7-180) wet mares and 21 (range 0-100) dry mares. The number of mares per breeding stallion increased with increasing size of breeding operation (p=0.04), being 28 (95% confidence interval (CI) = 10-56) vs 40 (95% CI=16-74) vs 74 (95% CI=44-113) for moderate (or=200 mares in total) operations, respectively. Seventy-one percent of farms aimed to breed dry mares early in the breeding season, and used a combination of lights, hormone therapy, and rising plane of nutrition to achieve this. Foaling took place in foaling paddocks monitored using a night foaling attendant (17/22) or with foaling alarms (5/22). At birth, 17/22 studmasters routinely administered antibiotics, 14/22 administered tetanus antitoxin, 9/22 administered an enema to foals, and 2/22 did not routinely administer prophylactic treatments. Weaning occurred at 5 (range 3.7-7) months of age, and foals were confined to a box for 1-2 weeks on 16/22 farms. Weaned foals were drenched with anthelmintics every 7 (range 4-9) weeks, and were fed 2.9 (range 1-6) kg of concentrate feed while at pasture until intensive management associated with preparation of the horses for auction began 13 (range 6-20) weeks before the yearling sales. Eight farms weighed the weanlings, at least monthly, to monitor growth. CONCLUSIONS AND CLINICAL RELEVANCE: The management of Thoroughbred horses was relatively consistent throughout the regions surveyed. Utilisation of breeding stallions tended to be more efficient on the larger stud farms in the south Auckland/Waikato region. Even though foals are grown at pasture they are often provided with large quantities of concentrate feed.     

Management Strategies Aiming to Improve Horse Welfare Reduce Embryonic Death Rates in Mares

The objective of this retrospective study was to evaluate the effect of management strategies aiming to improve animal well‐being on pregnancy and embryonic death (ED) rates. Breeding records of a cohort of 1206 Thoroughbred mares brought to a stallion station facility, to be bred with the stallions housed there, were evaluated during ten breeding seasons. Mares were blocked according to management strategies in two groups: Stress and Relax. Strategies used to improve animal well‐being (Relax group) were as follows: stopping the teasing routine, reducing or eliminating stall confinement, reducing the number of mares per group and maintaining herd stability during the breeding season. In barren mares, the pregnancy rate was higher in the Relax group (91.8%) when compared to the observed in Stress group (84.7%). However, no difference in pregnancy rates were observed (Stress = 85.2% vs. Relax = 86.2) in foaling mares. ED rate was higher in barren and foaling mares of the Stress group mares (25.5% and 26.8%, respectively) compared with the Relax group (16.1% and 14.7%, respectively). No significant differences were observed on foal heat pregnancy rate between groups; yet, the embryo loss on foal heat was significant reduced in Relax mares (Relax = 8.7% vs Stress = 24.5%). In conclusion, management strategies aimed to reduce social stress can reduce early pregnancy losses and the average cycles per pregnancy, improving reproductive performance in mares.     

Studies on Motility and Fertility of Cooled Stallion Spermatozoa

This study on extended, cooled stallion spermatozoa aimed to compare the ability of three extenders to maintain sperm motility during 24 h of preservation, and to describe pregnancy and foaling rates after artificial insemination (AI) of stallion spermatozoa stored and transported in the extender chosen from the in vitro study. After 6 and 24 h of preservation, motility, both subjective and evaluated by the motility analyzer (total, progressive and rapid), was lower in non-fat, dried skim milk-glucose than in both other extenders: dried skim milk-glucose added to 2% centrifuged egg yolk, and ultra high temperature treated skim milk-sugar-saline solution added to 2% centrifuged egg yolk (INRA82-Y). Rapid spermatozoa and sperm velocity parameters, after 24 h, were significantly higher in INRA82-Y. In the fertility trial, semen collected from three Maremmano stallions, diluted in INRA82-Y, and transported in a refrigerated Styrofoam box, was used to inseminate 56 mares of the same breed. Pregnancy rates after the first cycle and per breeding season were significantly higher for the 31 mares inseminated in three AI centres (54.8 and 80.6%, respectively) than for the 25 mares inseminated at the breeder's facilities (28.0 and 52.0%). Foaling rates were not significantly different between the AI centres mares (54.8%) and the other mares (44.0%). In conclusion, INRA82-Y yielded satisfactory pregnancy and foaling rates, especially when employed in the more controlled situation of an AI centre, and can therefore be included among those available for cooled stallion semen preservation.     

Inheritance of myotonic discharges in American quarter horses and the relationship to hyperkalemic periodic paralysis.

Electromyography (EMG) was used to detect myotonic discharges in Quarter Horse breeding stock and to follow the results of mating horses with hyperkalemic periodic paralysis (HPP). The studies were performed on two brood mare farms. A total of six breeding stock showed myotonic discharges and 15 were nonmyotonic. Myotonic discharges were seen in five of six horses belonging to the blood line previously implicated as being predisposed to HPP. Two of these horses had shown clinical signs of HPP. Only one of 15 breeding horses unrelated to the HPP predisposed blood line showed myotonic discharges. When both parents were non-myotonic on EMG than the F1 generation (n = 6) were also nonmyotonic. When a stallion with HPP and myotonic discharges was mated to eight nonmyotonic mares over a six year period half the animals of the F1 generation (n = 25) showed myotonic discharges. When both parents showed myotonic discharges four F1 offspring were myotonic and two were nonmyotonic on EMG testing. There was no evidence of sex linkage. The results are consistent with an autosomal dominant mode of inheritance. Hyperkalemic periodic paralysis and myotonic discharges on EMG may be different manifestations of the same underlying defect.     

Genital injuries from mating in the mare

Minor accidents during natural mating are common occurrences during the breeding of horses. Mares may suffer from a variety of genital injuries including vulval separations, vaginal lacerations and, less commonly, vaginal rupture. Possible sequelae to, and management of, these injuries are described, with additional mention of false entry by the stallion and persistent hymen in maiden mares. Vaginal rupture during natural mating can lead to death of the mare. The implications for litigation involving veterinary surgeons and horse breeders are discussed.