The Journal of experimental biology.
Publisher:
Cambridge Univ. Press.. London : Company Of Biologists Limited
Frequency: Twenty four no. a year
Country: England
Language: English
Author(s):
Company of Biologists.
Start Year:1930 -
ISSN:
0022-0949 (Print)
1477-9145 (Electronic)
0022-0949 (Linking)
1477-9145 (Electronic)
0022-0949 (Linking)
Impact Factor
2.8
2022
NLM ID: | 0243705 |
(DNLM): | J20480000(s) |
(OCoLC): | 01754580 |
Coden: | JEBIAM |
Classification: | W1 JO644H |
Roles of collagen cross-links and osteon collagen/lamellar morphotypes in equine third metacarpals in tension and compression tests. Many bones experience bending, placing one side in net compression and the other in net tension. Because bone mechanical properties are relatively reduced in tension compared with compression, adaptations are needed to reduce fracture risk. Several toughening mechanisms exist in bone, yet little is known of the influences of secondary osteon collagen/lamellar 'morphotypes' and potential interplay with intermolecular collagen cross-links (CCLs) in prevalent/predominant tension- and compression-loaded regions. Paired third metacarpals (MC3s) from 10 adult horses were prepared for mechanical test...
Metabolomic responses to high-intensity interval exercise in equine skeletal muscle: effects of rest interval duration. High-intensity interval training has attracted considerable attention as a time-efficient strategy for inducing physiological adaptations, but the underlying mechanisms have yet to be elucidated. By using metabolomics techniques, we investigated changes in the metabolic network responses in Thoroughbred horses to high-intensity interval exercise performed with two distinct (15 min or 2 min) rest intervals. The peak plasma lactate level was higher during high-intensity exercise with a 2 min rest duration than that with a 15 min rest duration (24.5±6.8 mmol/L vs. 13.3±2.7 mmol/L). The arterial...
Why don’t horseflies land on zebras? Stripes deter horseflies (tabanids) from landing on zebras and, while several mechanisms have been proposed, these hypotheses have yet to be tested satisfactorily. Here, we investigated three possible visual mechanisms that could impede successful tabanid landings (aliasing, contrast and polarization) but additionally explored pattern element size employing video footage of horseflies around differently patterned coats placed on domestic horses. We found that horseflies are averse to landing on highly but not on lightly contrasting stripes printed on horse coats. We could find no evidence for ...
The effect of diet change and insulin dysregulation on the faecal microbiome of ponies. The equine microbiome can change in response to dietary alteration and may play a role in insulin dysregulation. The aim of this study was to determine the effect of adding pasture to a hay diet on the faecal bacterial microbiome of both healthy and insulin-dysregulated ponies. Faecal samples were collected from 16 ponies before and after dietary change to enable bacterial 16S rRNA sequencing of the V3-V4 region. The dominant phyla in all samples were the Firmicutes and Bacteroidetes. The evenness of the bacterial populations decreased after grazing pasture, and when a pony was moderately insu...
Ground reaction forces of overground galloping in ridden Thoroughbred racehorses. The horse has evolved to gallop economically at high speed. Limb force increases with speed but direct measures of limb ground reaction forces (GRFs) at gallop are sparse. This study reports GRFs for multiple limbs, using force plates, across seven Thoroughbred racehorses during ridden galloping. The results show peak vertical GRF values of 13.6 N kg (non-lead hindlimb), 12.3 N kg (lead hindlimb), 14.0 N kg (non-lead forelimb) and 13.6 N kg (lead forelimb) at 11.4 m s and recorded values are consistent with those predicted from duty factor. The distribution of body weight b...
Mammals repel mosquitoes with their tails. The swinging of a mammal's tail has long been thought to deter biting insects, which, in cows, can drain up to 0.3 liters of blood per day. How effective is a mammal's tail at repelling insects? In this combined experimental and theoretical study, we filmed horses, zebras, elephants, giraffes and dogs swinging their tails. The tail swings at triple the frequency of a gravity-driven pendulum, and requires 27 times more power input. Tails can also be used like a whip to directly strike at insects. This whip-like effect requires substantial torques from the base of the tail on the order of 10-1...
Thyroid hormones correlate with field metabolic rate in ponies, Equus ferus caballus. During winter, free-living herbivores are often exposed to reduced energy supply at the same time that energy needs for thermoregulation increase. Several wild herbivores as well as robust horse breeds reduce their metabolism during times of low ambient temperature and food shortage. Thyroid hormones (THs) affect metabolic intensity and a positive effect of THs on basal metabolic rate (BMR) has been demonstrated in mammals and birds. As BMR and field metabolic rate (FMR) are often assumed to be intrinsically linked, THs may represent a reliable indicator for FMR. To test this hypothesis, 10 Sh...
Effects of acceleration on gait measures in three horse gaits. Animals switch gaits according to locomotor speed. In terrestrial locomotion, gaits have been defined according to footfall patterns or differences in center of mass (COM) motion, which characterizes mechanisms that are more general and more predictive than footfall patterns. This has generated different variables designed primarily to evaluate steady-speed locomotion, which is easier to standardize in laboratory conditions. However, in the ecology of an animal, steady-state conditions are rare and the ability to accelerate, decelerate and turn is essential. Currently, there are no data availa...
Saving energy during hard times: energetic adaptations of Shetland pony mares. Recent results suggest that wild Northern herbivores reduce their metabolism during times of low ambient temperature and food shortage in order to reduce their energetic needs. It is, however, not known whether domesticated animals are also able to reduce their energy expenditure. We exposed 10 Shetland pony mares to different environmental conditions (summer and winter) and to two food quantities (60% and 100% of maintenance energy requirement) during low winter temperatures to examine energetic and behavioural responses. In summer, ponies showed a considerably higher field metabolic rate (FM...
Modelling biomechanical requirements of a rider for different horse-riding techniques at trot. The simplest model possible for bouncing systems consists of a point mass bouncing passively on a mass-less spring without viscous losses. This type of spring-mass model has been used to describe the stance period of symmetric running gaits. In this study, we investigated the interaction between horse and rider at trot using three models of force-driven spring (-damper)-mass systems. The first system consisted of a spring and a mass representing the horse that interact with another spring and mass representing the rider. In the second spring-damper-mass model, dampers, a free-fall and a forcin...
Forelimb muscle activity during equine locomotion. Few quantitative data exist to describe the activity of the distal muscles of the equine forelimb during locomotion, and there is an incomplete understanding of the functional roles of the majority of the forelimb muscles. Based on morphology alone it would appear that the larger proximal muscles perform the majority of work in the forelimb, whereas the smaller distal muscles fulfil supplementary roles such as stabilizing the joints and positioning the limb for impact with the ground. We measured the timing and amplitude of the electromyographic activity of the intrinsic muscles of the forelim...
Adaptation strategies to seasonal changes in environmental conditions of a domesticated horse breed, the Shetland pony (Equus ferus caballus). Recent results suggest that the wild ancestor of the horse, the Przewalski horse, exhibits signs of a hypometabolism. However, there are speculations that domestic animals lost the ability to reduce energy expenditure during food shortage and adverse environmental conditions. Therefore, we investigated physiological and behavioural strategies employed by a robust domesticated horse breed, the Shetland pony, over the course of a year under temperate conditions by measuring ambient temperature (T(a)), subcutaneous temperature (T(s)), locomotor activity (LA), lying time, resting heart rate, body ...
Intraspecific scaling of chewing cycle duration in three species of domestic ungulates. In mammals, chewing cycle duration (CCD) increases with various measures of size, scaling with body mass(0.13-0.28) and jaw length(0.55). Proposed explanations for these scaling relationships include the allometry of body size, basal metabolic rate and tooth size, on the one hand, and pendular mechanics treating the jaw as a gravity-driven pendulum, on the other. Little is known, however, about the relationship between CCD and size within species. Recent research in dogs demonstrates altogether different scaling exponents and weaker correlations. This research suggests that breed-specific grow...
Relationship between muscle forces, joint loading and utilization of elastic strain energy in equine locomotion. Storage and utilization of strain energy in the elastic tissues of the distal forelimb of the horse is thought to contribute to the excellent locomotory efficiency of the animal. However, the structures that facilitate elastic energy storage may also be exposed to dangerously high forces, especially at the fastest galloping speeds. In the present study, experimental gait data were combined with a musculoskeletal model of the distal forelimb of the horse to determine muscle and joint contact loading and muscle-tendon work during the stance phase of walking, trotting and galloping. The flexor te...
Intensity of activation and timing of deactivation modulate elastic energy storage and release in a pennate muscle and account for gait-specific initiation of limb protraction in the horse. The equine biceps brachii (biceps) initiates rapid limb protraction through a catapult mechanism. Elastic strain energy is slowly stored in an internal tendon and is then rapidly released to protract the forelimb. The muscle fibres are short, have little scope for length change and can therefore only shorten slowly compared with the speed at which the whole muscle must shorten, which makes them poor candidates for driving rapid limb protraction. We suggest that the muscle fibres in the biceps act to modulate the elastic energy output of the muscle-tendon unit (MTU) to meet the demands of locom...
Limits to running speed in dogs, horses and humans. Are there absolute limits to the speed at which animals can run? If so, how close are present-day individuals to these limits? I approach these questions by using three statistical models and data from competitive races to estimate maximum running speeds for greyhounds, thoroughbred horses and elite human athletes. In each case, an absolute speed limit is definable, and the current record approaches that predicted maximum. While all such extrapolations must be used cautiously, these data suggest that there are limits to the ability of either natural or artificial selection to produce ever fast...
Stabilization and mobility of the head, neck and trunk in horses during overground locomotion: comparisons with humans and other primates. Segmental kinematics were investigated in horses during overground locomotion and compared with published reports on humans and other primates to determine the impact of a large neck on rotational mobility (> 20 deg.) and stability (< or = 20 deg.) of the head and trunk. Three adult horses (Equus caballus) performing walks, trots and canters were videotaped in lateral view. Data analysis included locomotor velocity, segmental positions, pitch and linear displacements and velocities, and head displacement frequencies. Equine, human and monkey skulls and cervical spines were measured to es...
High-speed gallop locomotion in the Thoroughbred racehorse. II. The effect of incline on centre of mass movement and mechanical energy fluctuation. During locomotion on an incline, mechanical work is performed to move an animal up the slope and increase the potential energy (PE) of the trunk and hence the centre of mass (CoM). Thus, at a given speed the total net mechanical work increases with the PE of the animal. In this study we investigate the mechanical energy (ME) fluctuations and the mechanical cost of transport (MCT) in six horses galloping up a range of gradients. We captured trunk movement with a six degrees-of-freedom inertial sensor mounted over the dorsal spinous process of the fourth to sixth thoracic vertebrae of the horse....
High-speed gallop locomotion in the thoroughbred racehorse. I. The effect of incline on stride parameters. During locomotion up an incline, power is required to elevate the centre of mass. This is provided when the animal's limbs are in contact with the ground. Measurements of stride timing variables from multiple limbs during high speed, over-ground locomotion would enhance our understanding of locomotor powering during changes in terrain. This study measured foot-on and foot-off times from galloping horses using a previously validated system of limb-mounted accelerometers and a global positioning system data logger. A detailed track survey provided incline information from all areas of the track....
Colour perception in a dichromat. Most mammals have dichromatic colour vision based on two different types of cones: a short-wavelength-sensitive cone and a long-wavelength-sensitive cone. Comparing the signal from two cone types gives rise to a one-dimensional chromatic space when brightness is excluded. The so-called ;neutral point' refers to the wavelength that the animal cannot distinguish from achromatic light such as white or grey because it stimulates both cone types equally. The question is: how do dichromats perceive their chromatic space? Do they experience a continuous scale of colours or does the neutral point divi...
Validation of vertical ground reaction forces on individual limbs calculated from kinematics of horse locomotion. The purpose of this study was to determine whether individual limb forces could be calculated accurately from kinematics of trotting and walking horses. We collected kinematic data and measured vertical ground reaction forces on the individual limbs of seven Warmblood dressage horses, trotting at 3.4 m s(-1) and walking at 1.6 m s(-1) on a treadmill. First, using a segmental model, we calculated from kinematics the total ground reaction force vector and its moment arm relative to each of the hoofs. Second, for phases in which the body was supported by only two limbs, we calculated the individu...
Modulation of mandibular loading and bite force in mammals during mastication. Modulation of force during mammalian mastication provides insight into force modulation in rhythmic, cyclic behaviors. This study uses in vivo bone strain data from the mandibular corpus to test two hypotheses regarding bite force modulation during rhythmic mastication in mammals: (1) that bite force is modulated by varying the duration of force production, or (2) that bite force is modulated by varying the rate at which force is produced. The data sample consists of rosette strain data from 40 experiments on 11 species of mammals, including six primate genera and four nonprimate species: goat...
Gait characterisation and classification in horses. Although a large number of foot-fall sequences are possible in quadrupeds, few sequences are routinely used. The aim of this paper is to characterise, by foot-fall pattern, the gaits used by horses and develop a novel technique to classify symmetric and asymmetric gaits using one common criterion. To achieve this speed and relative foot-fall, timings of all four limbs of eight Icelandic horses were measured using accelerometers. Linear discriminant analysis (LDA) was performed to find criteria that are optimal for discriminating between the different gaits. This also allowed us to evaluate whe...
Seasonal adjustment of energy budget in a large wild mammal, the Przewalski horse (Equus ferus przewalskii) II. Energy expenditure. Many large mammals show pronounced seasonal fluctuations of metabolic rate (MR). It has been argued, based on studies in ruminants, that this variation merely results from different levels of locomotor activity (LA), and heat increment of feeding (HI). However, a recent study in red deer (Cervus elaphus) identified a previously unknown mechanism in ungulates--nocturnal hypometabolism--that contributed significantly to reduced energy expenditure, mainly during late winter. The relative contribution of these different mechanisms to seasonal adjustments of MR is still unknown, however. Therefore,...
Seasonal adjustment of energy budget in a large wild mammal, the Przewalski horse (Equus ferus przewalskii) I. Energy intake. Large ruminants respond to changing plant phenology during winter by decreasing voluntary food intake, increasing gut passage time and utilizing body fat reserves. It is uncertain, however, how other large mammals with a non-ruminant digestive physiology cope with winter forage conditions. Therefore, we investigated seasonality of energy intake in a large herbivorous wild mammal, the Przewalski horse (Equus ferus przewalskii). Throughout all seasons we used the n-alkane method to measure daily dry matter intake (DMI), diet composition and digestion, and determined an index of gut passage time ...
Effect of speed on stride parameters in racehorses at gallop in field conditions. Stride duration, stance duration and protraction duration are key variables when describing the gaits of terrestrial animals. Together, they determine the duty factor (the fraction of the stride for which the limb maintains contact with the ground surface), from which the peak vertical force can be estimated. When an animal changes speed, these variables change at different proportions. Limited measurements of these variables and predictions of peak limb force have been undertaken for large mammals performing high-speed over-ground exercise. This study set out to make such measurements, employ...
Joint work and power for both the forelimb and hindlimb during trotting in the horse. The net work of the limbs during constant speed over level ground should be zero. However, the partitioning of negative and positive work between the fore- and hindlimbs of a quadruped is not likely to be equal because the forelimb produces a net braking force while the hindlimb produces a net propulsive force. It was hypothesized that the forelimb would do net negative work while the hindlimb did net positive work during trotting in the horse. Because vertical and horizontal impulses remain unchanged across speeds it was hypothesized that net work of both limbs would be independent of speed. ...
Centre of mass movement and mechanical energy fluctuation during gallop locomotion in the Thoroughbred racehorse. During locomotion cyclical interchange between different forms of mechanical energy enhances economy; however, 100% efficiency cannot be achieved and ultimately some mechanical work must be performed de novo. There is a metabolic cost associated with fluctuations in mechanical energy, even in the most efficient animals. In this study we investigate the exchanges between different forms of mechanical energy involved in high-speed gallop locomotion in Thoroughbred race horses during over-ground locomotion using innovative, mobile data collection techniques. We use hoof-mounted accelerometers to ...
The influence of collagen fiber orientation and other histocompositional characteristics on the mechanical properties of equine cortical bone. This study examined relative influences of predominant collagen fiber orientation (CFO), mineralization (% ash), and other microstructural characteristics on the mechanical properties of equine cortical bone. Using strain-mode-specific (S-M-S) testing (compression testing of bone habitually loaded in compression; tension testing of bone habitually loaded in tension), the relative mechanical importance of CFO and other material characteristics were examined in equine third metacarpals (MC3s). This model was chosen since it had a consistent non-uniform strain distribution estimated by finite ele...
Adaptive value of ambling gaits in primates and other mammals. At speeds between the walk and the gallop, most mammals trot. Primates almost never trot, and it has been claimed that they transition directly from a walk to a gallop without any distinctive mid-speed running gait. If true, this would be another characteristic difference between the locomotion of primates and that of most other quadrupedal mammals. Presently, however, few data exist concerning the actual presence or absence of intermediate-speed gaits (i.e. gaits that are used between a walk and a gallop) in primates. Video records of running in twelve primate species reveal that, unlike most...