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 |
Dynamically similar locomotion in horses. It is possible for animals of very different sizes to use the same patterns of locomotion, i.e. to move in a ;dynamically similar fashion'. This will only occur, however, if relevant biomechanical parameters scale with size in such a way that they compensate for the effects of size differences. Here we apply this principle to understanding the effects of size on locomotion within a species: the domestic horse. We predict that, without any factor to compensate for size differences, detectable deviations from dynamically similar locomotion would occur over the size range present in adult horses....
A method for deriving displacement data during cyclical movement using an inertial sensor. Biomechanical studies often employ optical motion capture systems for the determination of the position of an object in a room-based coordinate system. This is not ideal for many types of study in locomotion since only a few strides may be collected per ;trial', and outdoor experiments are difficult with some systems. Here, we report and evaluate a novel approach that enables the user to determine linear displacements of a proprietary orientation sensor during cyclical movement. This makes experiments outside the constraints of the laboratory possible, for example to measure mechanical energy ...
In vivo muscle function vs speed. II. Muscle function trotting up an incline. Different locomotor tasks, such as moving up or down grades or changing speed, require that muscles adjust the amount of work they perform to raise or lower, accelerate or decelerate the animal's center of mass. During level trotting in the horse, the triceps had shortening strains of around 10.6% while the vastus shortened 8.1% during the stance phase. Because of the 250% increase in metabolic rate in horses trotting up a 10% incline which is, presumably, a result of the increased requirement for mechanical work, we hypothesized that muscle strain during trotting would be increased in both th...
In vivo muscle function vs speed. I. Muscle strain in relation to length change of the muscle-tendon unit. The activity of muscles can be concentric (shortening), eccentric (lengthening) or isometric (constant length). When studying muscle function it is important to know what the muscle fascicles are actually doing because the performance of muscle is strongly influenced by the type of activity: force decreases as a function of shortening velocity during concentric contractions; force produced during eccentric contractions can be stronger than maximum isometric force, and force production is enhanced if a concentric contraction follows an eccentric phase. It is well known that length changes of mu...
Contribution of the forelimbs and hindlimbs of the horse to mechanical energy changes in jumping. The purpose of the present study was to gain more insight into the contribution of the forelimbs and hindlimbs of the horse to energy changes during the push-off for a jump. For this purpose, we collected kinematic data at 240 Hz from 23 5-year-old Warmbloods (average mass: 595 kg) performing free jumps over a 1.15 m high fence. From these data, we calculated the changes in mechanical energy and the changes in limb length and joint angles. The force carried by the forelimbs and the amount of energy stored was estimated from the distance between elbow and hoof, assuming that this part of the le...
Biomechanical and energetic determinants of the walk-trot transition in horses. We studied nine adult horses spanning an eightfold range in body mass (M(b)) (90-720 kg) and a twofold range in leg length (L) (0.7-1.4 m). We measured the horses' walk-trot transition speeds using step-wise speed increments as they locomoted on a motorized treadmill. We then measured their rates of oxygen consumption over a wide range of walking and trotting speeds. We interpreted the transition speed results using a simple inverted-pendulum model of walking in which gravity provides the centripetal force necessary to keep the leg in contact with the ground. By studying a large size range of ...
Determination of peak vertical ground reaction force from duty factor in the horse (Equus caballus). Measurement of peak vertical ground reaction force (GRFz) from multiple limbs simultaneously during high-speed, over-ground locomotion would enhance our understanding of the locomotor mechanics of cursorial animals. Here, we evaluate the accuracy of predicting peak GRFz from duty factor (the proportion of the stride for which the limb is in contact with the ground). Foot-mounted uniaxial accelerometers, combined with UHF FM telemetry, are shown to be practical and accurate for the field measurement of stride timing variables, including duty factor. Direct comparison with the force plate produc...
Ground reaction forces in horses trotting up an incline and on the level over a range of speeds. Although the forces required to support the body mass are not elevated when moving up an incline, kinematic studies, in vivo tendon and bone studies and kinetic studies suggest there is a shift in forces from the fore- to the hindlimbs in quadrupeds. However, there are no whole-animal kinetic measurements of incline locomotion. Based on previous related research, we hypothesized that there would be a shift in forces to the hindlimb. The present study measured the force produced by the fore- and hindlimbs of horses while trotting over a range of speeds (2.5 to 5 m s(-1)) on both level and up an...
Moments and power generated by the horse (Equus caballus) hind limb during jumping. The ability to jump over an obstacle depends upon the generation of work across the joints of the propelling limb(s). The total work generated by one hind limb of a horse and the contribution to the total work by four joints of the hind limb were determined for a jump. It was hypothesized that the hip and ankle joints would have extensor moments performing positive work, while the knee would have a flexor moment and perform negative work during the jump. Ground reaction forces and sagittal plane kinematics were simultaneously recorded during each jumping trial. Joint moment, power and work wer...
Are uniform regional safety factors an objective of adaptive modeling/remodeling in cortical bone? It has been hypothesized that a major objective of morphological adaptation in limb-bone diaphyses is the achievement of uniform regional safety factors between discrete cortical locations (e.g. between cranial and caudal cortices at mid-diaphysis). This hypothesis has been tested, and appears to be supported in the diaphyses of ovine and equine radii. The present study more rigorously examined this question using the equine third metacarpal (MC3), which has had functionally generated intracortical strains estimated by a sophisticated finite element model. Mechanical properties of multiple mid...
The energetics of the trot-gallop transition. Two studies have focused on potential triggers for the trot-gallop transition in the horse. One study concluded that the transition was triggered by metabolic economy. The second study found that it was not metabolic factors but, rather, peak musculoskeletal forces that determine gait transition speeds. In theory, peak musculoskeletal forces should be the same when trotting up an incline as when trotting at the same speed on the level. Assuming this is the case, we hypothesized that if peak forces determine gait transition speeds then horses should switch from a trot to a gallop at the same sp...
The effect of gait and digital flexor muscle activation on limb compliance in the forelimb of the horse Equus caballus. A horse's legs are compressed during the stance phase, storing and then returning elastic strain energy in spring-like muscle-tendon units. The arrangement of the muscle-tendon units around the lever-like joints means that as the leg shortens the muscle-tendon units are stretched. The forelimb anatomy means that the leg can be conceptually divided into two springs: the proximal spring, from the scapula to the elbow, and the distal spring, from the elbow to the foot. In this paper we report the results of a series of experiments testing the hypothesis that there is minimal scope for muscle cont...
Torque patterns of the limbs of small therian mammals during locomotion on flat ground. In three species of small therian mammals (Scandentia: Tupaia glis, Rodentia: Galea musteloides and Lagomorpha: Ochotona rufescens) the net joint forces and torques acting during stance phase in the four kinematically relevant joints of the forelimbs (scapular pivot, shoulder joint, elbow joint, wrist joint) and the hindlimbs (hip joint, knee joint, ankle joint, intratarsal joint) were determined by inverse dynamic analysis. Kinematics were measured by cineradiography (150 frames s(-1)). Synchronously ground reaction forces were acquired by forceplates. Morphometry of the extremities was perfo...
The anisotropic Young’s modulus of equine secondary osteones and interstitial bone determined by nanoindentation. The equine radius is a useful subject for examining the adaptation of bone histology to loading because in life the anterior cortex is loaded almost entirely in tension, the posterior cortex in compression. The histology of the two cortices is correspondingly different, the osteones and the interstitial lamellae in the posterior cortex having a more transversely oriented fibre arrangement than those in the anterior cortex. Presumably as a result of this histological difference, the posterior cortex is stronger in compression than the anterior cortex; the anterior cortex is stronger in tension ...
Preferred speed and cost of transport: the effect of incline. Preferred speed is the behavioral tendency of animals to utilize a relatively narrow set of speeds near the middle of a much broader range that they are capable of using within a particular gait. Possible explanations for this behavior include minimizing musculoskeletal stresses and maximizing energetic economy. If preferred speed is determined by energetic economy (cost of transport, C(T)), then shifts in preferred speed should produce shifts in C(T). To test this hypothesis, preferred speeds were measured in trotting horses on the level and on an incline. The preferred trotting speed decreas...
Oxygen-sensitive membrane transporters in vertebrate red cells. Oxygen is essential for all higher forms of animal life. It is required for oxidative phosphorylation, which forms the bulk of the energy supply of most animals. In many vertebrates, transport of O(2) from respiratory to other tissues, and of CO(2) in the opposite direction, involves red cells. These are highly specialised, adapted for their respiratory function. Intracellular haemoglobin, carbonic anhydrase and the membrane anion exchanger (AE1) increase the effective O(2)- and CO(2)-carrying capacity of red cells by approximately 100-fold. O(2) also has a pathological role. It is a very reac...
Time of contact and step length: the effect of limb length, running speed, load carrying and incline. Using published values for twelve species of birds and mammals, we investigated the effects of limb length and running speed on time of contact and step length. In addition, we measured the time of contact in horses trotting up a 10 % incline and when carrying a load averaging 19 % of their body mass. From these values, we calculated stride period and step length. Our analysis of the interspecific data yielded the following relationship between time of contact (t(c) in s) and leg length (L in m) and running speed (v in m s(-)(1)): t(c)=0.80L(0.84)/v(0.87) (r(2)=0.97). Both exponents in this re...
The relationship between mechanical work and energy expenditure of locomotion in horses. Three-dimensional motion capture and metabolic assessment were performed on four standardbred horses while walking, trotting and galloping on a motorized treadmill at different speeds. The mechanical work was partitioned into the internal work (W(INT)), due to the speed changes of body segments with respect to the body centre of mass, and the external work (W(EXT)), due to the position and speed changes of the body centre of mass with respect to the environment. The estimated total mechanical work (W(TOT)=W(INT)+W(EXT)) increased with speed, while metabolic work (C) remained rather constant. A...
The development of microcracking and failure in bone depends on the loading mode to which it is adapted. During locomotion, the anterior cortex of the equine radius is loaded predominantly in tension, the posterior predominantly in compression. The anterior cortex is relatively strong in tension, the posterior in compression. We investigated the pattern of failure of specimens from the two cortices using laser scanning confocal microscopy. All specimens were loaded in four-point bending to increasingly higher loads. We quantified the amount of diffuse microcracking on the tensile side of these specimens by observing the amount of light emitted under laser illumination. The amount of light emitted...
Micromechanics of the equine hoof wall: optimizing crack control and material stiffness through modulation of the properties of keratin. Small-scale components of the equine hoof wall were tested to determine their mechanical roles in the morphological hierarchy. Macroscale tensile tests conducted on samples of the inner wall tubules and intertubular material showed a sixfold difference in mean initial stiffnesses (0.47 and 0.08 GPa, respectively), indicating that the inner wall tubules stiffen the wall along its longitudinal axis. The similarity in material properties of tubule and intertubular samples from the mid-wall suggests that tubules in this region offer only minor reinforcement along the longitudinal axis. Microscale ...
Stress/strain behaviour of the equine laminar junction. The equine laminar junction plays a vital role in transferring the forces of weight-bearing between the epidermal hoof wall and the bone of the third phalanx, but the way in which it performs this function is poorly understood. Using samples from sites varying proximodistally and circumferentially around the hoof, the stress/strain behaviour of this tissue was characterised in three directions: radial tension and proximodistal and mediolateral shear. The influences of toe angle and length were also examined. For all three test directions, the modulus of elasticity increased with increasing str...
Persistent desensitisation of the beta 2 adrenoceptors expressed by cultured equine sweat gland epithelial cells. Adrenaline, forskolin and ATP all evoked accumulation of cyclic AMP in equine sweat gland epithelial cells, although the response to adrenaline was more transient than that to forskolin and ATP. Cells preincubated in adrenaline (10 micromol l-1, 32 min) showed essentially complete, homologous desensitisation, and this phenomenon reversed slowly (half-time 6.3+/-0.9 h). After 10 min of recovery from preincubation in adrenaline, isobutylmethylxanthine (IBMX, 5 mmol l-1) had no effect upon the desensitisation and the cells showed no loss of sensitivity to ATP and forskolin. After 10 h, however, t...
The effect of exercise-induced localised hyperthermia on tendon cell survival. Tendons that store energy during locomotion, such as the equine superficial digital flexor tendon (SDFT) and human Achilles tendon, suffer a high incidence of central core degeneration which is thought to precede tendon rupture. Although energy storage contributes to the efficiency of locomotion, tendons are not perfectly elastic and some energy is lost in the form of heat. Recent studies have shown that the central core of equine SDFT reaches temperatures as high as 45 degrees C during high-speed locomotion. In this study, we test the hypothesis that hyperthermia causes tendon cell death and ...
Design complexity and fracture control in the equine hoof wall. Morphological and mechanical studies were conducted on samples of equine hoof wall to help elucidate the relationship between form and function of this complex, hierarchically organized structure. Morphological findings indicated a dependence of tubule size, shape and helical alignment of intermediate filaments (IFs) within the lamellae on the position through the wall thickness. The plane of the intertubular IFs changed from perpendicular to the tubule axis in the inner wall to almost parallel to the tubule axis in the outer wall. Morphological data predicted the existence of three crack dive...
Activation of apical P2U purine receptors permits inhibition of adrenaline-evoked cyclic AMP accumulation in cultured equine sweat gland epithelial cells. Experiments were undertaken using cultured equine sweat gland epithelial cells that express purine receptors belonging to the P2U subclass which allow the selective agonist uridine triphosphate (UTP) to increase the concentration of intracellular free Ca2+ ([Ca2+]i). Experiments using pertussis toxin (Ptx), which inactivates certain guanine-nucleotide-binding proteins (G-proteins), showed that this response consisted of Ptx-sensitive and Ptx-resistant components, and immunochemical analyses of the G-protein alpha subunits present in the cells showed that both Ptx-sensitive (alpha i1-3) and Ptx...
The modulus of elasticity of equine hoof wall: implications for the mechanical function of the hoof. During normal weight-bearing and locomotion, the equine hoof wall deforms in a consistent pattern; the proximal dorsal wall rotates caudo-ventrally about the distal dorsal border and there is latero-medial flaring posteriorly. The aim of this study is to examine whether there are regional differences in the modulus of elasticity of hoof wall material and whether such differences correlate with the pattern of deformation which occurs in vivo. The modulus of elasticity of equine hoof wall was determined in tension and compression for samples from six forefeet. Samples were tested at the mid-poin...
Strain-rate-dependent mechanical properties of the equine hoof wall. The mechanical properties of fully hydrated equine hoof wall were examined at various loading rates in compact tension (CT) fracture, tensile and three-point bending dynamic tests to determine possible effects of hoof wall viscoelasticity on fracture toughness and tensile parameters. Four cross-head rates were used in CT tests: 1.7 x 10(-5), 1.7 x 10 (-3), 1.7 x 10(-2) and 2.5ms-1; four strain rates were used in tensile tests: 1.6 x 10(-3), 3.2 x 10(-2), 0.33 and 70s(-1). Speeds for the highest test rates were achieved using a large, custom-built impact pendulum. Bending test frequencies range...
Capillary blood transit time in muscles in relation to body size and aerobic capacity. The mean minimal transit time for blood in muscle capillaries (tc) was estimated in six species, spanning two orders of magnitude in body mass and aerobic capacity: horse, steer, dog, goat, fox and agouti. Arterial (CaO2) and mixed venous (CvO2) blood O2 concentrations, blood hemoglobin concentrations ([Hb]) and oxygen uptake rates were measured while the animals ran on a treadmill at a speed that elicited the maximal oxygen consumption rate (VO2max) from each animal. Blood flow to the muscles (Qm) was assumed to be 85% of cardiac output, which was calculated using the Fick relationship. Total...
Extracellular ATP can activate autonomic signal transduction pathways in cultured equine sweat gland epithelial cells. Changes in intracellular free calcium concentration ([Ca2+]i) were monitored in a cell line that was derived from the equine sweat gland epithelium. ATP and closely related compounds could increase [Ca2+]i with a rank order of potency of UTP > or = ATP > ADP >> AMP = adenosine = alpha,beta-methylene-ATP. The responses to ATP and to UTP were initiated by the release of calcium from an internal store and subsequently sustained by calcium influx. The rise in [Ca2+]i thus seems to be mediated by P2U receptors that are coupled to phosphoinositidase C. Some desensitisation of this respon...
Investigation of stimulus-secretion coupling in equine sweat gland epithelia using cell culture techniques. When sweat glands isolated from samples of horse skin were explanted and cultured under favourable conditions, they could exhibit cellular outgrowth. This growth could be maintained for 2-4 weeks and these primary cultures were then disaggregated and the resultant cell suspensions used to initiate epithelial cell lines. Secretion from intact equine sweat glands is regulated by beta 2-adrenoceptors and appears to be mediated by cyclic AMP, but there is evidence that calcium may also play a role. Adrenaline could increase the cyclic AMP content of the cultured cells and this response was mediate...