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Net joint moments and powers in the equine forelimb during the stance phase of the trot.

The objective of this study was to provide normative data describing the net joint moments and joint powers for the stance phase of the forelimb in trotting horses. Kinematic and force plate data, synchronised in time and space, were collected for the right forelimb of 6 Warmblood horses moving at a trot. The 3-D kinematic data were collapsed onto a sagittal plane, and were combined with the vertical and longitudinal ground reaction forces and with segment morphometric data to calculate net joint moments in the sagittal plane across the distal interphalangeal (coffin), metacarpophalangeal (fetlock), carpal, elbow and shoulder joints. The joint mechanical power was calculated as the product of the joint moment and the joint's angular velocity. Major peaks on the moment and power curves were identified. Each joint showed consistent and repeatable patterns in the net joint moments and joint powers. During most of stance the net joint moment was on the caudal/palmar side of all joints except the shoulder. At the coffin joint the power profile indicated an energy absorbing function that peaked at 74% stance, which coincided with the maximal longitudinal propulsive force. The fetlock joint behaved as an elastic spring; energy was absorbed in the first half of stance as the flexor tendons and SL stored elastic energy, which was released in the second half of stance as a result of elastic recoil. The carpus did not appear to play an important role in energy absorption or propulsion. Both the elbow and shoulder joints showed what appeared to be phases of elastic energy storage and release in the middle part of the stance phase, followed by a propulsive function at the shoulder in the later part of stance. The fetlock, carpus and elbow showed virtually no net generation or absorption of energy. The net energy generation at the shoulder joint was approximately equal to the energy absorption at the coffin joint. In human subjects specific gait pathologies produce characteristic alterations in the shape of the power profile as well as changes in the amount of energy absorbed and generated at the joints. In horses evaluation of net joint moments and joint powers will further our understanding of the mechanics and energetics of lameness, and may prove to be a useful diagnostic tool. An understanding of the function and dysfunction of different anatomical structures will facilitate the interpretation of clinical findings in terms of mechanical deficits.

Animals↗

Supporting forelimb lameness: clinical judgement vs. computerised symmetry measurement.

The aim of this study was to compare supporting forelimb lameness determined by a motion analysis system with the subjective grading of a trained equine orthopaedic surgeon. Trotting on a treadmill, 29 individuals with a supporting forelimb lameness were measured with the SELSPOT II system and judged by the clinician. The vertical motion of the head was measured, analysed using Fourier transform, and the percentages of symmetry determined. The veterinarian evaluated the lameness and graded it according to a clinical routine. Veterinarian and system for motion analysis assigned the lameness to the same leg in all cases, but the grading of the lameness differed in 6 out of 29 cases. The results of this study indicate that motion analysis can be used as an informative tool supporting the subjective veterinary judgement.

Animals↗

Forelimb kinematics and net joint moments during the swing phase of the trot.

The purpose of this study was to calculate net moments of force at the joints of the forelimb during the swing phase of the stride. An optoelectronic system was used to measure segmental kinematics for 3 strides in 5 sound, Warmblood horses trotting at a mean velocity +/- s.d. of 3.03 +/- 0.16 m/s. A link segment model was used to determine the net moments of force about the joints of the left forelimb. The model combined kinematic data with morphometric data describing the inertial parameters of the limb segments of warmblood horses, and incorporated correction factors for skin displacement. At each joint the net moment of force was on the cranial/dorsal side during the early swing phase and on the caudal/palmar side during the later swing phase. The transition (time of zero moment) occurred between 35-52% of the swing phase. The peak magnitude of the net joint moments decreased progressively in a proximal to distal direction. Published electromyographic (EMG) data correlated well with the timing of muscular activity required to generate the calculated net joint moments. The moments in the proximal limb are indicative of muscular activity accelerating the limb forward during the first 30-40% of the swing phase, then decelerating the forward swing of the upper limb segments. The net joint moments at all of the joints except the elbow work to slow the motion of the joints. The net joint moment about the elbow actively flex and then extend the joint. The low net joint moments at the distal joints during the first half of swing are consistent with their motion being primarily a result of inertial forces. Flexor muscle activity during the last half of swing indicate active control in preparation for ground contact.

Animals↗

In vitro transmission and attenuation of impact vibrations in the distal forelimb.

An in vitro model was developed and validated in vivo to quantify the attenuation of impact vibrations, transmitted through the lower equine forelimb and to assess the effects of horseshoeing on this attenuation. The transsected forelimbs of 13 horses were equipped with custom-made hollow bone screws in the 4 distal bones, on each of which a tri-axial accelerometer could be mounted. The limbs were then preloaded while the impact was simulated by dropping a weight on the steel plate on which the hoof was resting. At the hoof wall, the distal, middle and proximal phalanx and at the metacarpal bone, the shock waves resulting from this impact were quantified. To assess the damping effects of shoeing, measurements were performed with unshod hooves, hooves shod with a normal flat shoe and hooves shod with an equisoft pad and a silicone packing between hoof and pad. The in vitro model was validated by performing in vivo measurements using one horse, and subjecting the limb of this horse to the same in vitro measurements after death. Approximately 67% of the damping of impact vibrations took place at the interface between the hoof wall and the distal phalanx. The attenuation of impact vibrations at the distal and proximal interphalangeal joints was considerably less (both 6%), while at the metacarpophalangeal joint 9% of the amplitude of that at the hoof wall was absorbed, leaving approximately 13% of the initial amplitude at the hoof wall detectable at the metacarpus. Compared to unshod hooves the amplitude at the hoof wall is 15% higher in shod hooves. No differences could be observed between shoe types. At the level of the first phalanx and metacarpus the difference between shod and unshod vanished; it was therefore concluded that, although shoeing might influence the amplitude of impact vibrations at the hoof wall, the effect of shoeing on the amplitude at the level of the metacarpophalangeal joint is minimal.

Animals↗

In vivo transmission of impact shock waves in the distal forelimb of the horse.

There is a high prevalence of lameness among Standardbred trotters, most commonly caused by noninfectious joint diseases, mainly related to training and competition. In this context, impact-related shock waves transmitted through the skeleton and joints have been proposed to be one important factor in the development of osteoarthritis. The aim of the present study was to investigate the characteristic pattern of the events immediately following first contact, with a focus on the in vivo transmission of impact shock waves in the distal forelimb. Two horses were trotted by hand over a force plate. Recordings of 3-D kinematics of the distal forelimb were carried out by use of a 240 Hz video system. Tri-axial accelerometer data were collected from a bone-mounted accelerometer on the midlateral side of the third metacarpal bone (McIII) and from another accelerometer attached to the lateral side of the hoof. Force plate and accelerometer data were sampled at 4.8 kHz using a 16-bit A/D-converter, synchronised with the kinematic data. The results indicate that the time lapse of the horizontal retardation of the hoof is an important factor in the attenuation of the impact. A shorter period of hoof braking showed higher amplitudes in the longitudinal retardation of McIII and a more rapid oscillation. This makes all parameters that affect the horizontal hoof braking potentially important to the orthopaedic health of the horse.

Animals↗

Sensitivity of forelimb swing phase inverse dynamics to inertial parameter errors.

Estimations of segmental inertial parameters are required for true inverse dynamics calculations during the swing phase of locomotion. This study attempts to quantify the effect of inertial parameter errors on inverse dynamic solutions. Swing phase forelimb net joint moments and powers at the trot (mean +/- s.d 3.03 +/- 0.16 m/s) were calculated with sagittal plane kinematic data from 5 Dutch Warmbloods using inertial parameters based on published regression equations for the breed. Significant peaks in the net moment and power curves for each forelimb joint were identified and measured. Net joint moments and powers were then recalculated after varying the segment mass, location of the segment centre of mass and the mass moment of inertia separately for each of the limb segments. Peak values for the net joint moments and joint powers were determined after each variation, and the percent change in peak value per percent change in inertial parameter was calculated. Segment mass was the most influential parameter, followed by location of the centre of mass. Changes in the mass moment of inertia showed little effect on peak values. The most influential single inertial parameter was the mass of the hoof segment with a mean +/- s.d effect 0.74 +/- 0.22 and 0.69 +/- 0.18 percent peak change per percent parameter change on net joint moments and powers, respectively, across all joints. The results demonstrate the need for an accurate approximation of segment masses during the swing phase, especially the hoof, and the need to account for any additional masses in the model, such as shoes.

Animals↗

Effect of walking velocity on forelimb kinematics and kinetics.

A database of biomechanical variables obtained from normal horses walking at a range of velocities is needed for comparison with the variables obtained from lame horses in which velocity cannot be predetermined. The objective was to investigate velocity-dependent changes in selected kinematic variables, ground reaction forces (GRF) and net joint energies in the forelimb and to develop statistical equations to calculate expected values of these variables for horses walking at different velocities. Five sound horses walked at a range of velocities (0.82 to 1.91 m/s) over a force plate. Kinematic data were recorded simultaneously for 51 trials. Kinematic, GRF and energetic variables were determined using standard methods. Correlation and simple regression analyses between velocity and measured variables were performed. An increase in walking velocity was correlated with an increase in stride length and decreases in stride and stance duration. Vertical, braking and propulsive impulses decreased as a consequence of the large reduction in stance duration, even though peak vertical, braking and propulsive GRFs increased. There was no significant increase in energy generation at any of the forelimb joints, indicating that muscle activity was not the source of the increase in GRFs. Changes in the longitudinal GRFs appeared to be influenced by velocity-dependent increases in head and neck oscillations. The equations obtained in this study can be used to calculate the expected normal variables from a range of walking velocities and to detect deviations from normal values in lame horses.

Animals↗

Electromyographic responses evoked in muscles of the forelimb by intracortical stimulation in the cat.

Chronically implanted microwires were used to deliver brief trains of electrical stimuli (11 cathodal pulses at 330 Hz and intensity 5-35 microA) to sixty-two locations in the grey matter of the pericruciate cortex in cats. Electromyographic (e.m.g.) responses in the contralateral forelimb were recorded from a total of ten muscles (four to eight in each animal) acting about the shoulder, elbow and wrist and on the digits. The animals were relaxed with little background e.m.g. in the muscles and as a result only excitatory effects could be described. Five muscles which are flexors in the locomotor context were excited from more electrodes, distributed more widely across the motor cortex, than another five muscles which are extensors during locomotion; this difference in 'accessibility' was present both at 35 microA stimulus intensity and at 15 microA. At a stimulus intensity of 15 microA, effective cortical electrodes tended to cluster either in the most lateral part of the anterior sigmoid gyrus (rostromedial focus) or in the coronal gyrus just caudal to a line prolonged beyond the lateral end of the cruciate sulcus (caudolateral focus). This is consistent with the existence of a double motor representation within the forelimb motor cortex (Pappas & Strick, 1981). The two foci were similar in that both gave rise to more flexor than extensor responses and to fewer responses in digit or wrist muscles than in muscles acting about more proximal joints (elbow and shoulder). At stimulus intensity 35 microA the latency of the earliest e.m.g. responses ranged from 11 to 14 ms in different muscles. For some muscles and electrodes the amplitude of the e.m.g. responses was substantially altered by a quite small postural change. After pyramidectomy the cortical thresholds and the e.m.g. latencies were both greatly increased.

Action Potentials↗

Effect of norepinephrine on lymph flow and edema formation in the canine forelimb.

In the present study we sought evidence for the hypothesis that norepinephrine (NE) can cause constriction of lymph vessels in the canine forelimb perfused at constant flow. Mechanical venous compression (small-vein pressure approximately equal to 45 mmHg), intra-arterial infusion of histamine (16 micrograms base/min), and intra-arterial infusion of NE (16 micrograms base/min) caused the limbs to gain weight at similar rates due to edema formation; the first two maneuvers caused a sustained increase in skin lymph flow, but the increase in lymph flow with NE was only transient. Similar changes were seen during infusion of NE at 1 and 2 micrograms/min, and studies with radioactive microspheres indicated that NE increased capillary blood flow. When venous pressure was elevated and held constant at 45 mmHg. NE (4 micrograms base/min ia) antagonized and phentolamine (400 micrograms/min ia) potentiated the increase in skin lymph flow. Finally, NE (16 micrograms base/min ia) caused a reduction in histamine (16 micrograms base/min)-elevated skin lymph flow, despite the fact that capillary pressure was greatly increased due to active venous constriction. We conclude that intra-arterial infusion of norepinephrine in the dog forelimb perfused at constant flow causes constriction of lymphatic trunk vessels in skin, resulting in increased resistance to the flow of lymph.

Animals↗

Effects of ANP on venous pressures and microvascular protein permeability in dog forelimb.

Intravenous administration of alpha-atrial natriuretic peptide (ANP) produces a decrease in arterial blood pressure due to a decrease in cardiac output. The mechanism of the decrease in cardiac output is unknown but has been suggested to result from transcapillary fluid efflux caused by venoconstriction and/or increased permeability of the microvascular membrane to plasma proteins. We investigated these possibilities in the dog forelimb perfused at constant flow and prepared to measure 1) large and small vessel pressures in the two parallel skin and skeletal muscle vascular beds, and limb weight; or 2) skin lymph flow and skin lymph protein concentration. In both preparations, human ANP was injected and infused into the brachial artery. Bolus injections of ANP (0.1-0.4 micrograms) depressed perfusion pressure much less than acetylcholine or an isotonic solution of potassium chloride and did not raise small vein pressures. ANP infused at nine different rates ranging from 0.1 ng to 38.8 micrograms/min failed to influence skin and muscle small and large vein pressures, limb weight, skin lymph flow, and skin lymph protein concentration. It also failed to affect perfusion pressure and skin and muscle small artery pressures until the infusion was shut off, at which time they increased above control levels. These studies suggest that human alpha-ANP is neither a direct venous constrictor nor a potent arteriolar dilator and does not directly influence the permeability of the microvascular membrane to plasma proteins in the skin and skeletal muscle of the dog forelimb.

Animals↗

Comparison of the exercise pressor reflex between forelimb and hindlimb muscles in cats.

In thirteen cats anesthetized with alpha-chloralose, we compared the cardiovascular and ventilatory responses to both static contraction and tendon stretch of a hindlimb muscle group, the triceps surae, with those to contraction and stretch of a forelimb muscle group, the triceps brachii. Static contraction and stretch of both muscle groups increased mean arterial pressure and heart rate, and the responses were directly proportional to the developed tension. The cardiovascular increases, however, were significantly greater (P < 0.05) when the triceps brachii muscles were contracted or stretched than when the triceps surae muscles were contracted or stretched, even when the tension developed by either maneuver was corrected for muscle weight. Likewise, the ventilatory increases were greater when the triceps brachii muscles were stretched than when the triceps surae muscles were stretched. Contraction of either muscle group did not increase ventilation. Our results suggest that in the anesthetized cat the cardiovascular responses to both static contraction and tendon stretch are greater when arising from forelimb muscles than from hindlimb muscles.

Animals↗

In vivo recordings of bulbospinal excitation in adult mouse forelimb motoneurons.

Here we report on pyramidal and reticulospinal excitation in forelimb motoneurons in the adult mouse using intracellular recordings in vivo. The results have been obtained in BALB/C mice, which were anesthetized with midazolam fentanyl/fluanison. In contrast to the rat, only weak and infrequent pyramidal excitation could be evoked with a minimal trisynaptic linkage. Disynaptic reticulospinal excitation could always be evoked, as well as monosynaptic excitation from the medial longitudinal fasciculus. The results suggest that the reticulospinal pathway in the mouse is important in voluntary motor control of the forelimbs and that the role of the corticospinal tract might be different in mouse compared with rat. Our study provides an opening for studying the effect of genetic manipulation on specified descending systems in the mouse in vivo.

Animals↗

Forelimb movements and muscle responses evoked by microstimulation of cervical spinal cord in sedated monkeys.

Documenting the forelimb responses evoked by stimulating sites in primate cervical spinal cord is significant for understanding spinal circuitry and for potential neuroprosthetic applications involving hand and arm. We examined the forelimb movements and electromyographic (EMG) muscle responses evoked by intraspinal microstimulation in three M. nemestrina monkeys sedated with ketamine. Trains of three stimulus pulses (10-80 muA) at 300 Hz were delivered at sites in regularly spaced tracks from C6 to T1. Hand and/or arm movements were evoked at 76% of the 745 sites stimulated. Specifically, movements were evoked in digits (76% of effective sites), wrist (15% of sites), elbow (26%), and shoulder (17%). To document the muscle activity evoked by a stimulus current just capable of eliciting consistent joint rotation, stimulus-triggered averages of rectified EMG were calculated at each site where a movement was observed. Typically, many muscles were coactivated at threshold currents needed to evoke movements. Out of the 13-15 muscles recorded per animal, only one muscle was active at 14% of the effective sites and two to six muscles were coactivated at 47% of sites. Thus intraspinal stimulation at threshold currents adequate for evoking movement typically coactivated multiple muscles, including antagonists. Histologic reconstruction of stimulation sites indicated that responses were elicited from the dorsal and ventral horn and from fiber tracts in the white matter, with little somatotopic organization for movement or muscle activation. The absence of a clear somatotopic map of output sites is probably a result of the stimulation of complex mixtures of fibers and cells.

Action Potentials↗

Unilateral forelimb amputation affects protein synthesis in ipsilateral and contralateral spinal cord and spinal ganglia neurons of the newt in vivo.

The aim of this investigation was to analyze the effect of urodele limb amputation and blastema growth on the protein synthesis in spinal cord and spinal ganglia neurons corresponding to the 3rd and 4th spinal segment of young, postmetamorphic Triturus vulgaris, in vivo. Protein synthesis was studied on both trunk sides as a function of right forelimb amputation at the distal stylopodium, and blastema development, during a critical period of 22 days post amputation (dpa). Protein synthesis was assessed by counting silver grains on autoradiographs obtained after pulse labeling with tritiated phenylalanine. An electronic image analyzer was used to evaluate the results. During the period of observation, several statistically significant, yet moderate changes were observed in the protein synthesis of motor neurons. In contrast, the studied metabolic parameter exhibited considerable augmentation in sensory neurons and spinal nerve. The main results may be summarized as follows: (a) Limb amputation triggers protein synthesis in both ipsilateral and contralateral neural elements; (b) protein synthesis is higher on the ipsilateral than on the contralateral side; (c) as the aftermath of forelimb amputation, the 4th ipsilateral ganglion responds with an approximately 20% higher protein synthesis in comparison to the 3rd one; (d) peaks in protein synthesis are encountered mainly at two time points: 3 hpa and 14 dpa. Protein synthesis returns to control levels after 22 days of limb regeneration.

Amputation, Surgical↗

Functional implications of felid forelimb anatomy.

The elbow and wrist anatomy of 17 felid species were studied and compared with that of other representative mammaliam carnivores. Based on the shape and position of the olecranon fossa, it was determined that for felids the forelimb cannot travel in a "pendulum-like" motion during locomotion, but must travel through an arch away from the parasagittal plane of the body. For the anterior limb, the degree of deviation from the parasagittal plane was correlated with habitat preference. In this regard, those felids that are exclusive forest dwellers (found exclusively in high, densely structured habitats) had the greatest angle of inclination of the olecranon fossa. In addition, these, species had a large lateral olecranon tuberosity for the attachment of the lateral head of the triceps muscle. For those felids that inhabit more open terrain (low-structured habitat), the olecranon fossa was less inclined, the medical olecranon tuberosity relatively large, and the medial head of the triceps was significantly heavier than those of the forest felids. Both the wrist and elbow joints exhibited a large degree of mobility which was reflective of the claw-equipped forelimb being used as a hunting weapon.

Anatomy, Comparative↗

Evidence that serotonin receptors mediate the cutaneous vasoconstriction produced by 5-hydroxytryptamine in canine forelimbs.

The effects of local intra-arterial infusions of serotonin (5 or 25 micrograms base/min) or norepinephrine (1 or 5 micrograms base/min) on cutaneous (skin) and skeletal muscle vasculatures were investigated in canine forelimbs perfused at constant flow in dogs anesthetized with pentobarbital. Norepinephrine produced dose-related constriction of the skin and skeletal muscle vasculatures. In the cutaneous vascular circuit, norepinephrine produced large artery, small vessel, and large vein constriction. The increase in cutaneous vascular resistance was primarily due to an increase in small vessel resistance. Serotonin did not increase skeletal muscle vascular resistance but produced marked cutaneous vasoconstriction subsequent to large artery and large vein constriction. The small vessels, if anything, tended to dilate. The skin and skeletal muscle vascular responses to serotonin and norepinephrine were similar in innervated and acutely denervated forelimbs. Phentolamine pretreatment completely blocked all vascular actions of norepinephrine, and largely inhibited the cutaneous vasoconstriction produced by the infusion of the low dose of serotonin. However, the cutaneous large artery and large vein constriction produced by the infusion of the high dose of serotonin was not affected by phentolamine pretreatment. Cyproheptadine pretreatment blocked or largely inhibited the cutaneous vasoconstriction produced by serotonin only in doses which also inhibited norepinephrine and vasopressin cutaneous vasoconstriction. Pretreatment with methysergide blocked or largely inhibited the cutaneous large artery and large vein constriction produced by infusions of serotonin. Norepinephrine and vasopressin produced significant vasoconstriction in the presence of methysergide. These data suggest that the cutaneous large artery and large vein constriction produced by serotonin is not due to the activation of postjunctional alpha-adrenergic receptors.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Specific and redundant functions of the paralogous Hoxa-9 and Hoxd-9 genes in forelimb and axial skeleton patterning.

Using gene targeting, we have produced mice with a disruption of Hoxa-9 or Hoxd-9, two paralogous Abdominal B-related genes. During embryogenesis, these genes are expressed in limb buds and along the vertebral axis with anterior expression boundaries at the level of prevertebra #20 for Hoxa-9 and #23 for Hoxd-9. Skeletal analysis revealed homeotic transformations corresponding to anteriorisations of vertebrae #21 to #25 (L1 to L5) in the lumbar region of Hoxa-9-/- mutants; vertebrae #23 to #25 (L3 to L5) in the lumbar region together with vertebrae #28, #30 and #31 (S2, S4 and Ca1) in the sacrum and tail were anteriorized in Hoxd-9-/- mutants. Thus, anteriorisation of vertebrae #23 to #25 were common to both phenotypes. Subtle forelimb (but not hindlimb) defects, corresponding to a reduction of the humerus length and malformation of its deltoid crest, were also observed in Hoxd-9-/-, but not in Hoxa-9-/-, mutant mice. By intercrosses between these two lines of mutant mice, we have produced Hoxa-9/Hoxd-9 double mutants which exhibit synergistic limb and axial malformations consisting of: (i) an increase of penetrance and expressivity of abnormalities present in the single mutants, and (ii) novel limb alterations at the level of the forelimb stylopod and additional axial skeleton transformations. These observations demonstrate that the two paralogous genes Hoxa-9 and Hoxd-9 have both specific and redundant functions in lumbosacral axial skeleton patterning and in limb morphogenesis at the stylopodal level. Taken all together, the present and previously reported results show that disruption of different Hox genes can produce similar vertebral transformations, thus supporting a combinatorial code model for specification of vertebral identity by Hox genes.

Abnormalities, Multiple↗

EphA4/ephrin-A5 interactions in muscle precursor cell migration in the avian forelimb.

Limb muscles derive from muscle precursor cells that lie initially in the lateral portion of the somitic dermomyotome and subsequently migrate to their target limb regions, where muscle-specific gene transcription is initiated. Although several molecules that control the generation and delamination of muscle precursor cells have been identified, little is known about the mechanisms that guide muscle precursor cell migration in the limb. We have examined the distribution of members of the Eph family during muscle precursor cell development. The EphA4 receptor tyrosine kinase and its ligand, ephrin-A5, are expressed by muscle precursor cells and forelimb mesoderm in unique spatiotemporal patterns during the period when muscle precursors delaminate from the dermomyotome and migrate into the limb. To test the function of EphA4/ephrin-A5 interactions in muscle precursor migration, we used targeted in ovo electroporation to express ephrin-A5 ectopically specifically in the presumptive limb mesoderm. In the presence of ectopic ephrin-A5, Pax7-positive muscle precursor cells are significantly reduced in number in the proximal limb, compared with controls, and congregate abnormally near the lateral dermomyotome. In stripe assays, isolated muscle precursor cells avoid substrate-bound ephrin-A5 and this avoidance is abolished by addition of soluble ephrin-A5. These data suggest that ephrin-A5 normally restricts migrating, EphA4-positive muscle precursor cells to their appropriate territories in the forelimb, disallowing entry into abnormal embryonic regions.

Animals↗