Morphometry of Macaca mulatta forelimb. II. Fiber-type composition in shoulder and elbow muscles

The present study examined the fiber-type proportions of 22 muscles spanning the shoulder and/or elbow joints of three Macaca mulatta. Fibers were classified as one of three types: fast-glycolytic (FG), fast-oxidative-glycolytic (FOG), or slow-oxidative (SO). In most muscles, the FG fibers predominated, but proportions ranged from 25-67% in different muscles. SO fibers were less abundant except in a few deep, small muscles where they comprised as much as 56% of the fibers. Cross-sectional area (CSA) of the three fiber types was measured in six different muscles. FG fibers tended to be the largest, whereas SO fibers were the smallest. While fiber-type size was not always consistent between muscles, the relative size of FG fibers was generally larger than FOG and SO fibers within the same muscle. When fiber CSA was taken into consideration, FG fibers were found to comprise over 50% of the muscle's CSA in almost all muscles.     

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.     

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.     

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.     

Morphology, histochemistry, and function of epaxial cervical musculature in the horse (Equus caballus).

The semispinalis capitis and splenius muscles of the horse were analyzed for gross morphology, microarchitecture, fiber length, and fiber type. Although these two muscles are similar in size and anatomical position, they are very different from one another in structural design and histochemistry, implying diverse functional roles in the animal's behavior. The histochemical staining profile was limited to two fiber types: slow oxidative and fast glycolytic. The splenius muscle has simple architecture, long fibers, and a 60/40 ratio of SO to FG cross-sectional area. The semispinalis capitis has complex architecture with short-fibered, concentric compartments dorsal to its central tendon and longer-fibered compartments ventrally. The entire dorsal region has an increasing gradient of slow oxidative fiber percentage from caudal to cranial (58-71% SO). In contrast, the ventral region has a decreasing gradient of slow oxidative fibers from caudal to cranial (48-67% FG). These patterns can be interpreted within the context of the cervical musculature during locomotion and posture to indicate the functional advantages of this organization.     

Metabolic investigations of fibroblasts from horses, Equus caballus, with hereditary severe combined immunodeficiency

In an attempt to determine the metabolic defect causing severe combined immunodeficiency (SCID) in horses in which altered purine metabolism has been observed, various parameters of purine and pyrimidine metabolism were evaluated. The activities of nine purine enzymes (adenosine kinase, purine nucleoside phosphorylase, deoxyadenosine kinase, deoxycytidine kinase, 5'-nucleotidase, AMP deaminase, hypoxanthine-guanine phosphoribosyl transferase, and adenine phosphoribosyl transferase were measured in fibroblasts. All activities determined for SCID horses were normal. Uptake of 10 microM adenosine or 2'-deoxyadenosine (a growth inhibitory concentration for SCID fibroblasts) by SCID fibroblasts was identical to that found for normal fibroblasts in the presence of both 1 and 50 microM phosphate. The Km determined for the transport of both adenosine and 2'-deoxyadenosine was 35 microM. In the presence of p-nitrobenzylthioguanosine (a nucleoside transport inhibitor), 2'-deoxyadenosine uptake was inhibited to the same extent in all fibroblast lines tested. To determine if the last step in pyrimidine biosynthesis might be altered in SCID fibroblasts, UMP synthase activities were evaluated but found to be normal (0.5 nmol UMP formed/min/mg protein).     

Morphometric and anatomic study of the forelimb of the dog

The object of this study was to obtain the anatomic and morphometric data required for biomechanical analyses of the forelimb in dogs. Following the euthanasia of four healthy, adult, crossbred dogs, 44 muscles of the right forelimb were identified and meticulously removed. Morphometric data for all muscles were collected and physiologic cross-sectional areas (PCSA) and architectural indices (AI) were calculated. The coordinates of the origin and insertion of each muscle were determined using orthogonal, right-handed coordinate systems embedded in the scapula, humerus, and radius-ulna. The PCSA and AI were calculated for all the muscles and coordinates for the origins and insertions of these muscles were determined. Results provide the morphometric and anatomic data necessary for three-dimensional biomechanical studies of the forelimb in dogs.     

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.     

Serum and liver lipid composition and lecithin: cholesterol acyltransferase in horses, Equus caballus.

1. The lipid composition of serum and liver and some properties of serum lecithin: cholesterol acyltransferase of the horse were investigated. 2. Phospholipids and cholesterol were the major components of serum lipids and the concentration of triglyceride was considerably low. The concentration of liver lipids was comparable with that of other mammals. 3. Fatty acid composition of serum cholesterol ester resembled that of the 2-position of lecithin, except palmitic acid. 4. The activity of serum cholesterol esterifying enzyme was found to be 0.03-0.09 mumol/hr per ml. There was an equimolar decrease in free cholesterol and lecithin during incubation, and changes in unsaturated fatty acids in these two components were in good agreement. 5. Cholesterol esterification was reversibly inhibited by 5,5'-dithiobis-(2-nitrobenzoic acid). The acyl-transferase had a specificity for linoleic acid.     

Fibre operating lengths of human lower limb muscles during walking

Muscles actuate movement by generating forces. The forces generated by muscles are highly dependent on their fibre lengths, yet it is difficult to measure the lengths over which muscle fibres operate during movement. We combined experimental measurements of joint angles and muscle activation patterns during walking with a musculoskeletal model that captures the relationships between muscle fibre lengths, joint angles and muscle activations for muscles of the lower limb. We used this musculoskeletal model to produce a simulation of muscle-tendon dynamics during walking and calculated fibre operating lengths (i.e. the length of muscle fibres relative to their optimal fibre length) for 17 lower limb muscles. Our results indicate that when musculotendon compliance is low, the muscle fibre operating length is determined predominantly by the joint angles and muscle moment arms. If musculotendon compliance is high, muscle fibre operating length is more dependent on activation level and force-length-velocity effects. We found that muscles operate on multiple limbs of the force-length curve (i.e. ascending, plateau and descending limbs) during the gait cycle, but are active within a smaller portion of their total operating range.     

Four forearm flexor muscles of the horse, Equus caballus: anatomy and histochemistry.

Two of the forearm flexors of the horse, the superficial and deep digital flexor muscles, are critical to support the digital and fetlock joints, exhibit differing insertions, and are passively supported by the proximal and distal check ligaments, respectively. These two muscles differ in histochemical composition and architecture. The differences are correlated with the different stress levels transmitted through their tendons, and the different frequencies of clinical breakdown that have been reported. Both muscles contain type I and type IIa fibers. A few type IIb fibers occurred in the deep digital flexor. The superficial digital flexor contained approximately 56% type I fibers, extremely short muscle fibers, and extensive connective tissue investment. In contrast, the deep digital flexor had three muscle heads: ulnar, radial, and "long" and "short" regions of the humeral head. The "long" and "short" regions of the humeral head contained 33% and 44% type I fibers, respectively, fiber lengths three to four times as long as those in the superficial digital flexor, and relatively less connective tissue investment. Flexor carpi radialis and flexor carpi ulnaris compared most closely with the humeral head of the deep digital flexor. These data suggest a correlation of the unique architecture of superficial digital flexor with its proposed elastic storage properties during locomotion in horses, and an explanation for the frequent breakdown of the superficial digital flexor in athletic horses.     

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.     

Effects of enrichment items on activity and social interactions in domestic horses (Equus caballus)

The aim of this study was to investigate the use of items intended to provide enrichment during turnout, both for individual and group kept horses in an attempt to reduce the amount of passive behaviours. The study was divided into two parts, where study 1 involved eight horses rotated through eight individual paddocks, each containing one of seven enrichment items and one paddock being kept without item, functioning as a control. The horses’ item-directed behaviours; passive behaviours or other non-item related activities were scored using instantaneous sampling, every minute for 1 h at the beginning and the end of the turnout period. Study 2 involved six horse groups (3–6 horses) and the same scoring methods and ethogram as in study 1. The four items that the horses interacted the most with during study 1 (straw STRA, ball filled with concentrates CBALL, branches BRAN and scratching pole POLE) are investigated in study 2. In addition, the amount of social interactions was recorded.Both horses kept individually (P < 0.05) and in groups (P < 0.0001) performed significantly more item-directed behaviours towards edible items like STRA and CBALL than other objects. There was, however, no overall relation between the numbers of item-directed behaviours and the number of passive behaviours observed, indicating that the enrichment items did not alone reduce the amount of passive behaviours during turnout periods. Such a reduction was, however, only apparent when horses spent more time eating green leaves growing on the paddock surface (R = ?0.97 study 1, R = ?0.67 study 2, P < 0.0001). Access to STRA in group kept horses also seemed to reduce the amount of agonistic behaviours (P < 0.0001). In conclusion, if grass is not available in paddocks, the provision of roughage reduces the amount of passive behaviours in singly kept horses and it also reduces the risk of agonistic interactions between horses kept in group.     

Contractile behavior of the forelimb digital flexors during steady-state locomotion in horses (Equus caballus): an initial test of muscle architectural hypotheses about in vivo function

The forelimb digital flexors of the horse display remarkable diversity in muscle architecture despite each muscle-tendon unit having a similar mechanical advantage across the fetlock joint. We focus on two distinct muscles of the digital flexor system: short compartment deep digital flexor (DDF(sc)) and the superficial digital flexor (SDF). The objectives were to investigate force-length behavior and work performance of these two muscles in vivo during locomotion, and to determine how muscle architecture contributes to in vivo function in this system. We directly recorded muscle force (via tendon strain gauges) and muscle fascicle length (via sonomicrometry crystals) as horses walked (1.7 m s(-1)), trotted (4.1 m s(-1)) and cantered (7.0 m s(-1)) on a motorized treadmill. Over the range of gaits and speeds, DDF(sc) fascicles shortened while producing relatively low force, generating modest positive net work. In contrast, SDF fascicles initially shortened, then lengthened while producing high force, resulting in substantial negative net work. These findings suggest the long fibered, unipennate DDF(sc) supplements mechanical work during running, whereas the short fibered, multipennate SDF is specialized for economical high force and enhanced elastic energy storage. Apparent in vivo functions match well with the distinct architectural features of each muscle.     

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.     

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.     

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.