Cross-modal individual recognition in domestic horses (Equus caballus) extends to familiar humans

It has recently been shown that some non-human animals can cross-modally recognize members of their own taxon. What is unclear is just how plastic this recognition system can be. In this study, we investigate whether an animal, the domestic horse, is capable of spontaneous cross-modal recognition of individuals from a morphologically very different species. We also provide the first insights into how cross-modal identity information is processed by examining whether there are hemispheric biases in this important social skill. In our preferential looking paradigm, subjects were presented with two people and playbacks of their voices to determine whether they were able to match the voice with the person. When presented with familiar handlers subjects could match the specific familiar person with the correct familiar voice. Horses were significantly better at performing the matching task when the congruent person was standing on their right, indicating marked hemispheric specialization (left hemisphere bias) in this ability. These results are the first to demonstrate that cross-modal recognition in animals can extend to individuals from phylogenetically very distant species. They also indicate that processes governed by the left hemisphere are central to the cross-modal matching of visual and auditory information from familiar individuals in a naturalistic setting.     

Social feeding decisions in horses (Equus caballus)

Like many other herbivores, in a natural environment equids feed on rather evenly distributed resources. However, the vegetation in their vast habitats constantly changes. If food is plentiful only little competition occurs over food, and in non-competitive situations domestic horses tend to return to the same feeding site until it is overgrazed. In contrast, they compete over limited food for which the social status of the individuals appears to be important. Especially in ruminants several studies have proved an influence of social organisations, rank, sex and the depletion of feeding sites on the feeding behaviour of individuals. However, it is not yet understood whether and how social aspects affect horses' feeding decisions. Curiosity about the influence of social rank on the horses' feeding decisions between two, equally with high-quality surplus food-filled buckets placed in different social feeding conditions, led us to create the test below. The observer horses were alternately tested with a dominant and a subordinate demonstrator placed in one of the three different positions. We conclude that domestic horses use social cognition and strategic decision making in order to decide where to feed in a social feeding situation. When possible they tend to return to the same, continuously supplied feeding site and switch to an "avoidance tendency" in the presence of dominant horses or when another horse is already feeding there. Thus, the social rank and the position of conspecifics affect the feeding strategy of horses.     

An investigation of colour discrimination with horses (Equus caballus)

The ability of four horses (Equus caballus) to discriminate coloured (three shades of blue, green, red, and yellow) from grey (neutral density) stimuli, produced by back projected lighting filters, was investigated in a two response forced-choice procedure. Pushes of the lever in front of a coloured screen were occasionally reinforced, pushes of the lever in front of a grey screen were never reinforced. Each colour shade was randomly paired with a grey that was brighter, one that was dimmer, and one that approximately matched the colour in terms of brightness. Each horse experienced the colours in a different order, a new colour was started after 85% correct responses over five consecutive sessions or if accuracy showed no trend over sessions. All horses reached the 85% correct with blue versus grey, three horses did so with both yellow and green versus grey. All were above chance with red versus grey but none reached criterion. Further analysis showed the wavelengths of the green stimuli used overlapped with the yellow. The results are consistent with histological and behavioural studies that suggest that horses are dichromatic. They differ from some earlier data in that they indicate horses can discriminate yellow and blue, but that they may have deficiencies in discriminating red and green.     

Elastic extension of leg tendons in the locomotion of horses (Equus caballus)

Several of the distal leg muscles of horses have such extremely short muscle fibres that their changes of length in locomotion must be due almost entirely to elastic extension of their tendons. Films of a horse have been analysed to determine these extensions, using data obtained by experiments on dissected legs. The tendons investigated experience peak strains of 3–6% in walking, 3–7% in trotting and 4–9% in galloping. These strains occur while the foot is on the ground.     

Aggression and social spacing in light horse (Equus caballus) mares and foals

Aggression and social spacing were studied in 14 light horse mares and their foals living at pasture. Focal samples were collected on each mare-foal dyad for 6 to 10.5 h from 2 months of foal age until weaning at approximately 4 months of age. Observations on foals continued until approximately 6 months of age for 7.5 to 10.5 h per foal. Every 2 min the identities of all individuals within 5 m were recorded. All occurrences of agonistic behavior, and the participants, were recorded during the focal samples. In addition, during feeding of supplemental grain, all occurrences of agonistic behavior by all subjects were recorded. Significant correlations were found between mare rank and the rank of foals both prior to and after weaning. Before weaning, the rank of the foal was significantly correlated with birth order. No significant correlation between birth order and foal rank was found for the post-weaning hierarchy. An animal's gender had no significant effect on foal rank or the choice of preferred associate. Both prior to and after weaning, foals associated preferentially with the foal of their dam's most preferred associate. In addition, significant positive correlations were found between rank of mares and foals and the rate at which they directed aggression to other herd members.     

Karyotypes of cattle (Bos taurus) and horses (Equus caballus) on the basis of synaptonemal complexes

The cytogenetic study performed has shown that karyotyping of meiotic cells can be based on the synaptonemal complexes (SC) of spreading pachytene spermatocytes of bull and of horse. The horse SC karyotype has not been previously described. A comparison of the relative length of SC with metaphase chromosomes of bull and horse somatic cells has revealed the correspondence of the chromosome length in pachytene of meiosis and metaphase, which is in agreement with the data on house mouse and Chinese hamster. The method of spreading pachytene cells may be of great practical importance in studies of the fertility disturbances in farm animals.     

Meal size and feeding management strategies influence the daily rhythm of total locomotor activity in horses (Equus caballus)

Keeping and management of horses can induce changes to instinctive and innate behavioural patterns. We investigated the effect of five different management conditions in five groups of horses. All groups were housed in individual boxes under natural environmental and lighting conditions. They were fed three times a day (07:00, 12:30 and 20:00) and had free access to water. Group A was fed with 8 kg/capo/die of hay divided in the three meals. Group B was fed with 8 kg/capo/die of an unifeed divided in the three meals. Group C was fed with unifeed at 07:00 and 12:30 and with hay at 20:00. They were kept in wood-bedded boxes. Groups D and E were fed with unifeed at 07:00 and 12:30, respectively, and in the other, two meals received hay. They were kept in straw-bedded boxes. Our results showed a daily rhythm of total locomotor activity in all groups, influenced by management conditions. Group A engaged in meal patterns similar to those seen in grazing animals. Groups B and C showed the highest MESOR values due to a high searching behaviour. Group C showed a nocturnal acrophase contrary to the other groups. Groups D and E showed a total locomotor activity pattern similar to that observed in Group A probably due to an increase in straw-bedding consuming. The reduction of fibre in diet has an impact on physiology and behaviour of horses. The valuation of diet and in bedding provided to horses kept in box is useful to guarantee the maintenance of the physiological daily rhythm of total locomotor activity.     

Architectural properties of distal forelimb muscles in horses,Equus caballus

Articular injuries in athletic horses are associated with large forces from ground impact and from muscular contraction. To accurately and noninvasively predict muscle and joint contact forces, a detailed model of musculoskeletal geometry and muscle architecture is required. Moreover, muscle architectural data can increase our understanding of the relationship between muscle structure and function in the equine distal forelimb. Muscle architectural data were collected from seven limbs obtained from five thoroughbred and thoroughbred-cross horses. Muscle belly rest length, tendon rest length, muscle volume, muscle fiber length, and pennation angle were measured for nine distal forelimb muscles. Physiological cross-sectional area (PCSA) was determined from muscle volume and muscle fiber length. The superficial and deep digital flexor muscles displayed markedly different muscle volumes (227 and 656 cm3, respectively), but their PCSAs were very similar due to a significant difference in muscle fiber length (i.e., the superficial digital flexor muscle had very short fibers, while those of the deep digital flexor muscle were relatively long). The ulnaris lateralis and flexor carpi ulnaris muscles had short fibers (17.4 and 18.3 mm, respectively). These actuators were strong (peak isometric force, Fmax=5,814 and 4,017 N, respectively) and stiff (tendon rest length to muscle fiber length, LT:LMF=5.3 and 2.1, respectively), and are probably well adapted to stabilizing the carpus during the stance phase of gait. In contrast, the flexor carpi radialis muscle displayed long fibers (89.7 mm), low peak isometric force (Fmax=555 N), and high stiffness (LT:LMF=1.6). Due to its long fibers and low Fmax, flexor carpi radialis appears to be better adapted to flexion and extension of the limb during the swing phase of gait than to stabilization of the carpus during stance. Including muscle architectural parameters in a musculoskeletal model of the equine distal forelimb may lead to more realistic estimates not only of the magnitudes of muscle forces, but also of the distribution of forces among the muscles crossing any given joint.     

Immunocontraception in wild horses (Equus caballus) extends reproductive cycling beyond the normal breeding season

Background

Although the physiological effects of immunocontraceptive treatment with porcine zona pellucida (PZP) have been well studied, little is known about PZP''s effects on the scheduling of reproductive cycling. Recent behavioral research has suggested that recipients of PZP extend the receptive breeding period into what is normally the non-breeding season.

Methodology/Principal Findings

To determine if this is the case, we compiled foaling data from wild horses (Equus caballus) living on Shackleford Banks, North Carolina for 4 years pre- and 8 years post-contraception management with PZP (pre-contraception, n = 65 births from 45 mares; post-contraception, n = 97 births from 46 mares). Gestation lasts approximately 11–12 months in wild horses, placing conception at approximately 11.5 months prior to birth. Since the contraception program began in January 2000, foaling has occurred over a significantly broader range than it had before the contraception program. Foaling in PZP recipients (n = 45 births from 27 mares) has consistently occurred over a broader range than has foaling in non-recipients (n = 52 births from 19 mares). In addition, current recipients of PZP foaled later in the year than did prior recipient and non-recipient mares. Females receiving more consecutive PZP applications gave birth later in the season than did females receiving fewer applications. Finally, the efficacy of PZP declined with increasing consecutive applications before reaching 100% after five consecutive applications.

Conclusions/Significance

For a gregarious species such as the horse, the extension of reproductive cycling into the fall months has important social consequences, including decreased group stability and the extension of male reproductive behavior. In addition, reproductive cycling into the fall months could have long-term effects on foal survivorship. Managers should consider these factors before enacting immunocontraceptive programs in new populations. We suggest minor alterations to management strategies to help alleviate such unintended effects in new populations.     

Dominance relationships and patterns of aggression in a bachelor group of Sorraia horses (Equus caballus)

The influence of individual factors on dominance rank and the relationship between rank distance and patterns of aggression predicted by models of evolutionarily stable strategies (ESS) of animal conflict were investigated in a managed bachelor group of Sorraia horses, Equus caballus. The group was composed of four to six stallions 3- to 12-years-old during the study period. The dominance hierarchy was significantly linear and rank was not related to age, weight, height or aggressiveness. Frequency and intensity of agonistic interactions were low, but higher-ranking stallions did not receive lower aggressiveness than lower-ranking stallions. There was some evidence that dominance relationships were more contested among close-ranking stallions, as predicted. Agonistic-related interactions among close-ranking stallions served similar functions to those among distant-ranking stallions, but the latter interacted more frequently than expected for access to resting sites and/or resting partners. Therefore, we found some evidence that agonistic-related interactions among distant-ranking stallions play a larger role in providing access to valuable and defendable resources than those among close-ranking stallions. Nevertheless, the fact that space to escape from aggression was limited and breeding access was independent from dominance rank may have reduced the benefits relative to costs of aggression and therefore limited the occurrence of contests over dominance and resources.     

Successful transfer of the embryos of Przewalski's horses (Equus przewalskii) and Grant's zebra (E. burchelli) to domestic mares (E. caballus)

Blastocysts were collected non-surgically from 2 Przewalski's horse and 2 Grant's zebra mares and transferred extra-specifically to domestic horse and donkey recipients. Nine Przewalski's horse embryos were transferred surgically, and 2 non-surgically, to domestic Welsh-type pony mares. After surgical transfer, 7 (77.8%) pregnancies were established and 4 foals were born. Twelve Grant's zebra embryos were transferred surgically to 5 pony and 7 domestic donkey recipients respectively and 1 non-surgically to a donkey; 3 (60%) zebra-in-horse pregnancies were established and 2 went to term. Only 2 (28.6%) zebra-in-donkey pregnancies were established but neither went to term, although one zebra foal was aborted alive at Day 292 but failed to survive. No pregnancies resulted from the non-surgical transfers. Measurement of chorionic gonadotrophin concentrations and parental-specific lymphocytotoxic antibodies in the serum of the recipient animals indicated a pronounced maternal immunological response to the extra-specific embryo, but this could not be correlated with success or failure of pregnancy. The results indicate that extra-specific embryo transfer may be a useful aid to breeding exotic equids in captivity.     

On the synchronization of breathing with running in wallabies (Macropus spp.) and horses (Equus caballus)

Previous investigations have shown that, when they run, wallabies and various quadrupedal mammals take exactly one breath per stride. It has been suggested that their breathing movements may be driven by the movements of running, and three mechanisms have been proposed: the lungs may be compressed and expanded by vibrations of a 'visceral piston', by bending and extension of the back or by the forces imposed on the rib cage by the forelegs.
The three mechanisms are tested against published observations on wallabies and horses. Simple mathematical models are used to predict the phase relationship between running and breathing for each mechanism, and the amplitudes of breathing that they could cause. The observed breathing movements of hopping wallabies are consistent with a visceral piston mechanism tuned to the stride frequency, but cannot be explained by either of the other mechanisms. Those of cantering and galloping horses are not consistent with a tuned visceral piston mechanism, but could be driven by the bending and extension of the back.     

Genetic diversity of Cheju horses (Equus caballus) determined by using mitochondrial DNA D-loop polymorphism

We used sequence polymorphism of the mitochondrial DNA D-loop (968 bp excluding the tandem repeat region) to determine genetic diversity of horses inhabiting Cheju (a southern island of Korea). Seventeen haplotypes with frequencies from 1.5 to 21.5% were found among 65 Cheju horse samples. Genetic diversity (h) of the 17 haplotypes was calculated to be 0.91, indicating that the extant Cheju horse population consists of diverse genetic groups in their maternal lineage. Phylogenetic analysis showed that 17 types of Cheju (D-loop sequences determined), 5 Mongolian, 6 Arabian, 3 Belgian, 2 Tsushima, 2 Yunnan, 1 Przewalskii, and 3 Thoroughbred horses (published sequences for the latter seven breeds) showed that Cheju horses were distributed into many different clusters in the tree. Four Mongolian horses clustered with separate Cheju horse groups, showing that some Cheju horses are clearly of Mongolian origin. The analysis of partial sequences (284 bp) of the D-loop of 109 horses showed that Thoroughbred, Mongolian, Lipizzan, and Arabian breeds are as diverse as Cheju horses. Our data together with others' suggest that most horse breeds tested with reasonably sufficient numbers of samples are diverse in their maternal lineages and also are not uniquely different from each other.     

Preliminary evaluation of porcine zona pellucida (PZP) immunocontraception for behavioral effects in feral horses (Equus caballus)

Successful management of captive populations of wild animals requires effective control of reproduction. Contraception is one tool for controlling reproduction of animals in zoos; however, the options available to the animal manager are limited. Contraceptives vary in efficacy, reversibility, and side effects, and thus may not be suitable for widespread use. One consideration when selecting a contraceptive is its potential for side effects on behavior, especially given the fact that reproduction plays such a prominent role in the biology of any species. To date, there have been few evaluations of contraceptives for behavioral effects, and those that have been conducted have focused on hormone-based contraceptives. This study sought to evaluate a novel method of population control, immunocontraception, for behavioral effects in a population of feral horses. Porcine zona pellucida (PZP) immunocontraception prevents fertilization of ova and does not alter normal hormone secretion patterns. It therefore should leave the animal behaviorally intact in terms of reproductive behavior. The study examined the behavior of 43 sexually mature mares on Assateague Island during the 1997 breeding season and, with help from Earthwatch volunteers, collected observations over a 3-month period. The study found no significant differences between treated and untreated mares in general activity budget, aggression given or received, and spatial relationships relative to the stallion. These preliminary findings indicate that PZP contraception seems to have no acute behavioral effects on the behavior of individuals. The study findings also suggest that PZP could be a desirable and effective management tool for captive species in which social behavior plays an integral role in group dynamics. Analyses of group level effects and population level effects are continuing.     

Functionally relevant responses to human facial expressions of emotion in the domestic horse (Equus caballus)

Whether non-human animals can recognize human signals, including emotions, has both scientific and applied importance, and is particularly relevant for domesticated species. This study presents the first evidence of horses'' abilities to spontaneously discriminate between positive (happy) and negative (angry) human facial expressions in photographs. Our results showed that the angry faces induced responses indicative of a functional understanding of the stimuli: horses displayed a left-gaze bias (a lateralization generally associated with stimuli perceived as negative) and a quicker increase in heart rate (HR) towards these photographs. Such lateralized responses towards human emotion have previously only been documented in dogs, and effects of facial expressions on HR have not been shown in any heterospecific studies. Alongside the insights that these findings provide into interspecific communication, they raise interesting questions about the generality and adaptiveness of emotional expression and perception across species.     

Plasma lipid transport in the horse (Equus caballus)

• 1.1. Equine plasma contains lipoproteins corresponding to very low density (VLDL), low density (LDL) and high density lipoproteins (HDL).
• 2.2. HDL accounts for approximately 60% of plasma lipoprotein mass and consists of a single population of particles.
• 3.3. LDL is heterogeneous comprising three discrete subfractions.
• 4.4. Two proteins are found in the region of apolipoprotein (apo) B-100 in VLDL and LDL and a third similar to apo B-48 is in VLDL.
• 5.5. Lecithin:cholesterol acyl transferase is active in plasma and hepatic lipase and lipoprotein lipase are evident in post-heparin plasma.
• 6.6. There is no significant cholesteryl ester transfer protein activity.

     

Suckling behaviour does not measure milk intake in horses, Equus caballus

Studies of parental investment in mammals have frequently used suckling behaviour to estimate energy transfer from mother to offspring, and consequently to measure maternal input. Such studies assume that the more an offspring sucks, the more milk it will receive. This assumption has been questioned, and a review of the literature found little support for it. To test if suckling behaviour provided an accurate index of milk or energy intake we used a radioactive isotope technique to label the milk of thoroughbred mares and to measure milk transfer to foals. We found no significant linear relationship between usual measures of suckling behaviour and milk or energy intake. No behaviours associated with suckling nor with characteristics of mares and foals improved the relationship; only the number of butts associated with each suck episode even approached significance. If we had used suckling behaviour to test theories on differential maternal investment our conclusions would have been in error. For example, female foals tended to suck for longer than males did but there was no difference in the amount of milk transferred. Consequently, we show that measures of suckling behaviour do not adequately predict milk intake in the domestic horse and we suggest that conclusions about differential maternal investment in mammals based on suckling behaviour are likely to be in error. Copyright 1999 The Association for the Study of Animal Behaviour.     

Quantitative characteristics of the differential banding pattern prometaphase chromosomes of the domestic horse (Equus caballus)

A high-resolution cytogenetic map (670 bands per haploid set) of RBA-banded chromosomes has been constructed in the domestic horse Equus caballus. The size and distribution of the replication-based R(G)-bands were analyzed using the computer program VideoTest-Karyo. The obtained data were compared to the results of cytogenetic mapping in other mammalian species and human.     

Long-term effects of porcine zonae pellucidae immunocontraception on ovarian function in feral horses (Equus caballus).

Ten feral mares free-roaming in Maryland, USA, were inoculated with porcine zonae pellucidae (PZP) protein before the breeding season for three consecutive years (1988-90). Ovarian function was monitored for 51 days during the peak of the breeding season after the third annual PZP inoculation, in seven of these mares and in four untreated control mares, by means of urinary oestrone conjugates and nonspecific progesterone metabolites. None of the ten inoculated mares became pregnant in 1990, compared with 55% of 20 control mares, which included two of the four monitored for ovarian function. Three of the untreated mares demonstrated apparent normal ovarian activity, characterized by preovulatory oestrogen peaks, concurrent progesterone nadirs at ovulation, breeding activity, and luteal-phase progesterone increases after ovulation. Two of the seven monitored PZP-treated mares demonstrated ovulatory cycles that did not result in conception. One was pregnant as a result of conception in 1989 and demonstrated a normal, late-gestation, endocrine profile. The remaining four PZP-treated mares revealed no evidence of ovulation, and urinary oestrogen concentrations were significantly depressed. The experiments indicated that (i) a third consecutive annual PZP booster inoculation is greater than 90% effective in preventing pregnancies in mares and (ii) three consecutive years of PZP treatment may interfere with normal ovarian function as shown by markedly depressed oestrogen secretion.