Maternal care and foal social relationships in a herd of Sorraia horses: Influence of maternal rank and experience

The influence of maternal rank and experience on patterns of maternal care and social relationships of foals were investigated in a managed herd of Sorraia horses, Equus caballus. Social interactions and spatial relationships of 13 foals (seven females and six males) born to seven mares were examined from birth to 10 months of life, within the three major periods of foal development. Conflict over suckling between dam and foal was not generally affected by rank and experience, but higher-ranking mothers allowed more suckling during late lactation than lower-ranking mothers. Foals of higher-ranking mares spent more time in proximity to the mother during socialization. Maternal rank and experience did not significantly affect maternal protectiveness, foal independence from the mother or the development of affiliative relationships between foals and group members. Foals of higher-ranking mares received lower frequencies of aggression from other horses only in the first month of life. Dominance relationships among foals depended mainly on aggressiveness and were not associated with maternal rank. The large variability in maternal behaviour, the absence of a significant association between maternal rank and body condition at parturition and the stable social environment within this herd may partly account for the reported results.     

Male infanticide and paternity analyses in a socially natural herd of Przewalski's horses: Sexual selection?

The sexual selection hypothesis explains infanticide by males in many mammals. In our 11-year study, we investigated this hypothesis in a herd of Przewalski's horses where we had witnessed infanticidal attacks. Infanticide was highly conditional and not simply linked to takeovers. Attacks occurred in only five of 39 cases following a takeover, and DNA paternity revealed that, although infanticidal stallions were not the genetic fathers in four cases out of five, stallions present at birth did not significantly attempt to kill unrelated foals. Infanticide did not reduce birth intervals; only in one case out of five was the infanticidal stallion, the father of the next foal; mothers whose foals were attacked subsequently avoided associating with infanticidal stallions. Therefore, evidence for the sexual selection hypothesis was weak. The “human disturbance” hypothesis received some support, as only zoo bred stallions which grew up in unnatural social groups attacked foals of mares which were pregnant during takeovers.     

Age-specific fertility rates of the Jeju pony (<Emphasis Type="Italic"> Equus caballus</Emphasis>)

Age-specific fertility is an essential parameter of life history. Here we report age-specific fertility rates, measured as the number of foals per mare per year, for Jeju ponies aged 2–28 years. The total sample consisted of 545 foals produced by 178 mares from 1988 to 2002. The mean fertility rate across all ages of the mares was 0.65 foals/mare per year (±0.24 SD). The fertility rates were above average for the 7- to 8- and 19- to 20-year-old mares, whereas they were relatively low for mares under 4 years old. The fertility rates tended to increase with the age of the stallion, but the relationship was not significant (n =15, P =0.09). The incidence of inter-birth interval was not associated with the age that the mare first reproduced (n =64 mares, P =0.99). However, mares that reproduced later in life tended to have a reduced fertility rate due to an increase in the duration of inter-birth intervals relative to mares that reproduced earlier (n =4 years of first reproduction, P =0.068). The fertility rates of Jeju ponies were lower than for other horses, perhaps because only one stallion was introduced to a relatively large herd every year. We suggest that the introduction of more stallions to the herd each year would increase fertility rates.     

On lactation and associated behaviour in a natural herd of horses

Developmental changes in time spent suckling and related mother-foal behaviour are described in an unmanaged herd of Camargue horses. Male foals spent about 40% more time suckling than females during the first 8 weeks. Body weight did not differ between the sexes but time-budgets did: males grazed less and were more active. If pregnant, the typical multiparous mare nursed her foals for 35–40 weeks, males and females alike, and weaned them 15 weeks before the next foaling. Primiparae lactated longer and weaned closer to the next foaling by 5 weeks. The mares played an active role in regulating the time spent suckling in early, and particularly in late lactation.     

Male infanticide in captive plains zebra, Equus burchelli

On the assumption that infanticide exists in plains zebra, as reported for horses, Equus caballus, we tested the following hypothesis. Introducing a new zebra male into a herd of breeding females should increase foal mortality in comparison with herds in which the sire of the foals is still present. The younger the foal, the more likely infanticide should be. We collected data from five herds in two zoological gardens in the Czech Republic. We found nine records of infanticide in plains zebra and three cases of abortions that were probably induced by forced copulation. We analysed additional indirect data to investigate the possibility of introduced males causing other abortions. Abortions were three times more likely in herds with introduced males than with only fathers present. Postnatal mortality of the foals was four times greater with introduced males than with fathers. No indication of a sex preference was observed for infanticide by a new male for either abortions or postpartum deaths. When we combined all records involving introduced males, the probability of foal death was greatest when the new male joined the herd just after conception and decreased with increasing time between conception and date of the new male introduction (the chance of a foal surviving was less than 5% just after conception and more than 50% at the time of delivery). Mortality of foals did not depend on whether the new male was introduced before or after the foal was born. Survival increased to more than 60% after the foal reached 1 month of age. Our results suggest that captive plains zebra show the highest occurrence of infanticide reported among ungulates. Copyright 2000 The Association for the Study of Animal Behaviour.     

Alliances and reproductive success in Camargue stallions

A study of a herd of Camargue horses Equus caballus, showed that while the majority of high-ranking stallions held single-male harems, some sons of low-ranking mares, being low ranking themselves, formed alliances that could last a lifetime. The two stallions were each other's closest associate and preferential grooming partner. Alliances were based on coalitions in which either both partners confronted an intruder synchronously or the dominant of the pair tended the female(s) while the subordinate simultaneously displayed towards the rival. Alliance partners were of similar age but were not more closely related to each other than to other stallions in the herd. Long-term paternity data revealed that subordinates sired close to a quarter of the foals born into the alliance group, and significantly more foals than low-ranking stallions in the herd adopting a 'sneak'-mating strategy. The dominant appeared to benefit from the presence of his subordinate partner. Fights occurred all year round, and the subordinate stallion of each alliance pair fought outside competitors more than twice as often as the dominant. Forming short-term alliances before defending mares on their own may enhance long-term reproductive success for both partners. Other benefits to both partners include higher survivorship of their foals and increased access to proven reproductive mares. These results suggest that the relationship between alliance partners is based on mutualism, but several conditions for reciprocity seem to be fulfilled: the benefit to the dominant (assistance in fights), and the benefit to the subordinate (access to reproduction), are both costly to the other partner and delayed in time. Copyright 1999 The Association for the Study of Animal Behaviour.     

Common white facial markings in bay and chestnut Arabian horses and their hybrids

Common white facial and leg markings have a multifactorial mode of inheritance in Equus caballus. Evidence for the complexity of the genetic component is the observation that chestnut (e/e) horses have more extensive white markings than do bay (E/-) horses. Computerized records obtained from the Arabian Horse Registry of America, Inc., were used to determine if heterozygous (E/e) bay horses have more extensive white facial markings than do homozygous (E/E) bay horses. Thirty-five sire families were analyzed. Each sire family consists of a sire, his foals, and the dams of those foals. The facial region was divided into five areas, and each horse was given a score from 0 to 5 according to the number of areas with whiteness. Since dams and foals with E/E genotypes cannot be identified in these sire families, mean facial scores were compared in dams and foals that were E/e and E/-. It was assumed that if a difference exists between E/e and E/E horses, the presence of E/E horses in the E/- group would reduce the mean of the E/- group. The results show that Arabian horses with the genotype E/e have more white markings than do horses with the genotype E/-, leading to the conclusion that horses with the genotypes e/e, E/e, and E/E vary as to the quantitative expression of white facial markings, with heterozygotes having an intermediate expression.     

T lymphocyte development and maturation in horses

Monoclonal antibodies specific for equine T lymphocyte subpopulations were produced and procedures for the continuous culture of equine lymphocytes were developed. These reagents and procedures were used to analyse the appearance, maturation and functions of T lymphocytes in normal horses and in T lymphocyte deficient horses with severe combined immunodeficiency (SCID). T lymphocytes appeared as early as the 75th day of fetal development and were normally distributed prior to birth of normal foals. Analysis of thymic T lymphocyte differentiation in SCID foals revealed the presence of both prothymocytes and mature thymocytes, but a virtual absence of cortical thymocytes. The data obtained support the hypothesis that two distinct pathways of T lymphocyte differentiation exist within the thymus. Although the gene defect in foals with SCID blocks the production of mature B and T lymphocytes, such foals do possess large granular lymphocytes which are cytotoxic following induction with interleukin 2. This suggests that lymphoid cells with natural killer cell activity are spared by the gene defect resulting in SCID in horses.     

Evidence of respiratory tract infection induced by equine herpesvirus, type 2, in the horse.

Five horses were experimentally exposed to equine herpesvirus 2 strain LK. Two young foals developed chronic pharyngitis (98 and 232 days, respectively). Growth characteristics, cytopathic effects (CPE), inclusion body formation, ether sensitivity, and immunofluorescent analysis indicated that the virus recovered from infected animals was a herpesvirus serologically identical with, or at least antigenically related to EHV-2 strain LK. No significant complement-fixing (CF) or virus-neutralizing (VN) antibody responses were observed in adult horses while both foals demonstrated a rise in CF antibody titer. One of the two foals demonstrated a rise in VN antibody only. The results suggest that EHV-2 virus induced chronic pharyngitis, primarily the result of lymphoid proliferation, with no overt clinical signs.     

Linkage of tobiano coat spotting and albumin markers in a pony family

Genetic segregation patterns among blood type markers and various phenotypically observed traits were studied in a small herd of ponies. The herd consisted of 10 mares without white spotting and a single stallion with the dominant pattern of tobiano spotting. Comparison of segregation patterns at loci for which the stallion was heterozygous showed tight linkage for the Alb-B and tobiano markers. In 17 cases in which the Alb contribution of the sire could be determined, all 10 foals that inherited AlbB from him were tobiano spotted, and all 7 non-spotted foals inherited his AlbA. The use of the symbol To is proposed for dominantly inherited tobiano spotting linked to the albumin.     

Inheritance of guttural pouch tympany in the arabian horse

The objective of the present study was to analyze the mode of inheritance of guttural pouch tympany (GPT) using pedigrees of Arabian horses. Complex segregation analyses were employed to test for the significance of nongenetic transmission and for monogenic, polygenic, and mixed monogenic-polygenic modes of inheritance. Horses affected by GPT comprised 27 Arabian purebred foals. Of these 27 animals, 22 were patients at the Clinic for Horses, School of Veterinary Medicine Hannover, Hannover, Germany, between 1994 and 2001 and 5 Arabian foals were from stud farms. Information on the pedigrees of these patients allowed us to classify the affected foals into four families with a total of 276 animals. The regressive logistic model analysis took into account the nonrandomness of the pedigrees through multiple single ascertainment correction. The complex segregation analysis showed that, among all other models employed, a polygenic and a mixed monogenic-polygenic model best explained the segregation of Arabian foals with GPT. Models including only nongenetic distributions and monogenic inheritance could be significantly rejected. This is the first report in which a genetic component could be shown to be responsible for GPT in horses.     

A missense mutation in the endothelin-B receptor gene is associated with Lethal White Foal Syndrome: an equine version of Hirschsprung Disease

Lethal White Foal Syndrome is a disease associated with horse breeds that register white coat spotting patterns. Breedings between particular spotted horses, generally described as frame overo, produce some foals that, in contrast to their parents, are all white or nearly all white and die shortly after birth of severe intestinal blockage. These foals have aganglionosis characterized by a lack of submucosal and myenteric ganglia from the distal small intestine to the large intestine, similar to human Hirschsprung Disease. Some sporadic and familial cases of Hirschsprung Disease are due to mutations in the endothelin B receptor gene (EDNRB). In this study, we investigate the role of EDNRB in Lethal White Foal Syndrome. A cDNA for the wild-type horse endothelin-B receptor gene was cloned and sequenced. In three unrelated lethal white foals, the EDNRB gene contained a 2-bp nucleotide change leading to a missense mutation (I118K) in the first transmembrane domain of the receptor, a highly conserved region of this protein among different species. Seven additional unrelated lethal white foal samples were found to be homozygous for this mutation. No other homozygotes were identified in 138 samples analyzed, suggesting that homozygosity was restricted to lethal white foals. All (40/40) horses with the frame overo pattern (a distinct coat color pattern that is a subset of overo horses) that were tested were heterozygous for this allele, defining a heterozygous coat color phenotype for this mutation. Horses with tobiano markings included some carriers, indicating that tobiano is epistatic to frame overo. In addition, horses were identified that were carriers but had no recognized overo coat pattern phenotype, demonstrating the variable penetrance of the mutation. The test for this mutant allele can be utilized in all breeds where heterozygous animals may be unknowingly bred to each other including the Paint Horse, Pinto horse, Quarter Horse, Miniature Horse, and Thoroughbred. Received: 25 November 1997 / Accepted: 3 February 1998     

Helicobacter equorum: prevalence and significance for horses and humans

Helicobacter equorum colonizes the caecum, colon and rectum of horses. The agent is highly prevalent in <6-month-old foals. In adult horses, the prevalence of H. equorum seems to be rather low, but these animals may harbour low, subdetectable numbers of this microorganism in their intestines. So far, no association could be made between the presence of H. equorum and clinical disease or intestinal lesions in adult horses. Further research is necessary to elucidate the pathogenic potential of this bacterial species towards young foals. Helicobacter equorum DNA was not detected in human faeces, indicating that this microorganism does not commonly spread from horses towards humans.     

Regulatory T Cells in Early Life: Comparative Study of CD4+CD25high T Cells from Foals and Adult Horses

The immune system of mammals is subject to continuous development during the postnatal phase of life. Studies following the longitudinal development of the immune system in healthy children are limited both by ethical considerations and sample volumes. Horses represent a particular valuable large animal model for T regulatory (Treg) cells and allergy research. We have recently characterised Treg cells from horses, demonstrated their regulatory capability and showed both their expansion and induction in vitro. Insect bite hypersensitivity (IBH) is a common allergy in horses resembling atopic dermatitis and studies have shown that first exposure to allergens in adult life results in an increased incidence of IBH. The aim of the present study was to characterize circulating CD4⁺CD25^highFoxP3⁺cells in foals, evaluate their suppressive capability and their in vitro induction compared to adult horses. 19 foals (age range, 1–5 months), their adult mothers and six one-year-old horses (yearlings) were included in the study. The proportion of FoxP3⁺ cells within the circulating CD4⁺CD25^high population was significantly higher in foals (47%) compared to their mothers (18%) and to yearlings (26%). Treg cells from foals also displayed a higher suppressive capability. Furthermore, CD4⁺CD25^high cells in foals could be induced in vitro from CD4⁺CD25⁻ cells in a significantly higher proportion compared to mares. These cells also displayed a significantly enhanced suppressive capability. In summary these findings support the notion that exposure of horses to allergens during maturation of the immune system assists the establishment of induced (i)Treg driven tolerance.     

Risk Factors for High Endoparasitic Burden and the Efficiency of a Single Anthelmintic Treatment of Danish Horses

A questionnaire survey regarding endoparasite control practices in Danish horse herds was carried out in 1995. The participating veterinarians and herd owners were sampled using convenience and purposive sampling. In the analysis of risk factors for development of a high endoparasitic burden (>200 eggs per gram faeces) 903 horses were sampled and the analysis of the efficiency of a single anthelmintic treatment was based on 605 horses. The following factors had a significant effect on the endoparasitic burden: herd type, age of the horses, use of pasture rotation, anthelmintic treatment of horses visiting the herd, use of an adviser in the planning of endoparasite control and advice regarding pasture rotation. An interaction between pasture rotation and advice regarding pasture rotation was found, but due to high collinearity this was not reported. The factors influencing significantly on the reduction of the faecal egg count after a single anthelmintic treatment were the type of herd, the age of the horse, the drug used, and the anthelmintic-resistance-status of the herd. A negative effect of permanent pastures was observed. If pasture hygiene was performed on the advice of the veterinarian, the effect of a single anthelmintic treatment was less compared to a single anthelmintic treatment without any advice. An interaction between the treatment group and the resistance-status of the herd was found. Additional factors, normally accounted for, when endoparasites and anthelmintic resistance is discussed, were investigated, but not found significant in this study.     

Seasonal changes in the white blood cell system, lyzozyme activity and cortisol level in Arabian brood mares and their foals

In 34 pure-breed Arabian horses divided into four groups (Gr. I, ten pregnant mares; Gr. II, seven barren mares; Gr. III, ten foals born in 1981; Gr. IV, seven foals born in 1982) seasonal changes in the white blood cell system, cortisol level and lyzozyme activity were studied. Seasonal periodicity was found in all groups for the number of lymphocytes, segmented neutrophils and eosinophils and cortisol level. Leukocyte periodicity was found in three groups, but not in the barren mares. In lyzozyme activity there was periodicity in three groups but not in the youngest foals. In the stab neutrophils, basophils and monocytes no cycle was observed. The behaviour of the indices studied showed the influence of age of the horses (mature vs young) and the physiological state of the mares (pregnancy or barrenness).     

Specific Immune Response of Mares and their Newborn Foals to Actinobacillus spp. Present in the Oral Cavity

Oral swab samples, serum and colostrum was taken from 15 mares and 14 of their foals, within 24 h of birth. The presence of antibody against Actinobacillus spp. isolated from the oral cavity was investigated using agar gel immunodiffusion. Antibodies against 48 out of the 77 Actinobacillus isolates from all horses in the study were present in the respective sera of 13 mares and 9 foals. In 11 mother-foal pairs, the antibody content of the foal serum was similar to that of the mare, and in 9 cases this was reflected in the antibody content of colostrum from the mare. The results indicate that an immune response to Actinobacillus spp. colonising the oral cavity is present in many adult horses and that this immune response can be transferred from mother to foal via colostrum.     

Continuous cultivation of equine lymphocytes: evidence for occasional T cell-like maturation events in horses with hereditary severe combined immunodeficiency

Peripheral blood mononuclear cells (PBMC) from 14 foals with hereditary severe combined immunodeficiency (SCID) were studied to determine the extent of lymphocyte differentiation that occurs in this disorder. PBMC from all 14 horses had the morphologic characteristics of large granular lymphocytes (LGL). Cells from only one of 14 horses were responsive to phytolectin stimulation in a standard blastogenesis assay; however, PBMC from all 14 horses proliferated in continuous culture in the presence of partially purified interleukin 2. Furthermore, there were differences in the growth patterns of these cultured cells that correlated with their ability to respond to phytolectin stimulation. PBMC obtained from the 13 phytolectin-unresponsive foals survived in culture for only 4 to 5 wk, divided very slowly, developed large granules composed primarily of calcium phosphate, and accumulated high concentrations of histamine. In contrast, PBMC from the phytolectin-responsive SCID foal proliferated in continuous culture for over 100 days, divided as rapidly as normal equine PBMC under identical culture conditions, and did not accumulate granules or histamine. These observations indicate that lymphoid cell differentiation occurs in some horses with SCID even though the identity of the LGL is unresolved. Two possibilities are that LGL are products of a pathway separate from that of lymphocytes or that LGL are precursors of mature lymphocytes.     

Lethal white foals in matings of overo spotted horses

Overo is a variable pattern of white coat color spotting which occurs in several breeds of horses in the United States. Occasionally, when overos are crossed inter se white foals are born which die soon after birth. Both intestinal tract abnormalities and isoerythrolysis have been reported in these foals. This report presents data which show that neonatal isoerythrolysis (NI) is not involved in the death of the white foals. Further research is needed to define the nature of the lethal anomaly, as well as the genetics, of overo and lethal white foals.     

Spatial relations between mares and foals of the Welsh pony (Equus caballus)

Welsh pony mares and foals (Equus caballus) were usually found to be within 1 or 5 m of each other during the first week of the foal's life and gradually spent more time at greater distances as the foals became older. There was an overall levelling of the trend during the 9th–15th weeks of life of the foal, followed by a second period of change during weeks 16–24. Through weeks 21–24, mares and foals spent at least half of their time within 5 m of each other. Proximity was primarily due to foal activity except during foal recumbency. During the first 8 weeks of the foal's life, a mare remained close by when it was recumbent, either by grazing in a circle around it or by standing upright beside it. Mares and foals were most likely to be close together when they were resting upright with the other ponies in the herd and most likely to be far apart when the foal was playing. Similarities in patterns of spatial relationship between the foals of a given mare were demonstrated. There was no difference between colts and filies in the development of independence.