Stride parameters and hindlimb length in horses fatigued on a treadmill and at an endurance ride.

REASON FOR PERFORMING STUDY: The relationship between fatigue and stride and/or muscle stiffness requires further study. OBJECTIVE: To measure stride parameters in horses undergoing fatigue associated with running at submaximal speeds both on a treadmill and in an endurance ride. HYPOTHESIS: Stride frequencies and estimates of hindlimb stiffness would be decreased in fatigued horses. METHODS: Horses were fatigued using 2 paradigms: run to exhaustion at a treadmill (4.5 m/sec, 6% incline) and finishing an 80 km endurance ride. Videos were digitised before and after fatigue and analysed for stride parameters: hind limb length, stride frequency, time of contact, step length, duty factor and stride length. RESULTS: In fatigued horses, stride durations were 5% longer (P = 0.007) resulting in lower stride frequencies (P = 0.016) and longer stride lengths (P = 0.006). The time of contacts (tc) for stance phase were not different (P = 0.108) nor was duty factor (tc/stride period, P = 0.457). Step length (speed x tc) and hindlimb lengths were also not different (P = 0.104, P = 0.8). For endurance horses, stride data for nonfatigued horses were consistent with data extrapolated to 4.5 m/sec from nonfatigued horses on the treadmill. Endurance horses slowed (P = 0.002) during the race from 4.55 to 4.03 m/sec and stride lengths were shorter. Despite a slower speed, other stride parameters were unchanged. Hindlimb length was shorter in fatigued horses. CONCLUSION: Horses fatigued on a treadmill and during the natural course of an endurance ride responded differently, biomechanically. On the treadmill, where speed is constrained, stride frequencies decreased and stride lengths increased. During one endurance ride, stride frequencies were the same, although speeds were substantially reduced. Limb length was shorter in fatigued endurance horses. It remains to be determined if these changes in mechanics are advantageous or disadvantageous in terms of energetics or injury. Further examination of endurance rides is also warranted.

Oxygen consumption (VO2) during trotting on a 10% decline.

REASONS FOR PERFORMING STUDY: Although there have been reports of oxygen consumption measurements of horses running on the level and incline, there are no measurements during decline locomotion. This may be due, in part, to the potential for muscle damage produced by eccentric contractions. In man, running on a 10% decline, VO2 decreased by 35% and stride frequency (SF) decreased by 3% when compared to level locomotion. HYPOTHESIS: The rate of O2 consumption and SF would be decreased in horses on a 10% decline when compared to the level. METHODS: Six horses (average 467 +/- 68 kg) were acclimated to trotting on the level and decline prior to data collection. VO2 under moderate conditions was measured (using open flow respirometry) during trotting between 2.25 and 4.0 m/sec (at 0.25 m/sec increments) on a treadmill on the level and declined 10%. Stride frequencies were counted manually. RESULTS: VO2 decreased (P<0.009) on the decline by an average of 45% (range 42-47%), and SF was 2.7% slower. The speed at which the minimum Cost of Transport occurs on the decline was faster than on the level. SF was reduced on the decline. No evidence of muscle soreness was noted in response to the downhill running. CONCLUSIONS AND POTENTIAL RELEVANCE: Downhill trotting, eccentric exercise, can be done safely in the horse and requires almost half the energetic costs as trotting on the level. It is not known whether this is the optimum downhill gradient or if the horse adjusts its preferred speed to accommodate downhill trotting.

Energetic and kinematic consequences of weighting the distal limb.

REASON FOR PERFORMING STUDY: It is well known that adding a load to a horse's back increases its energetic costs of locomotion, but the magnitude of increase obtained by loading the most distal portion of limb has not been measured. OBJECTIVES: To measure oxygen consumption in horses with mass added to the back and hooves. Because such mass distribution alters inertial parameters of the limbs, kinematic measurements were made to quantify the magnitude of change in limb movement. METHODS: Steady-state oxygen consumption was measured in 6 horses with a load of 2.4 kg. The load was either carried on the back or distributed equally between the 4 limbs. Modified bell boots kept the mass at the level of P3. Horses trotted on a treadmill at speeds ranging from 2 to 5 m/sec (in 0.5 m/sec increments). High-speed (250 Hz) digital images were recorded in a sagittal plane and the positions of retroreflective markers located on standard positions on the limbs were digitised for kinematic analysis. RESULTS: Loading of the distal limbs produced a 6.7% increase in metabolic rate, an order of magnitude higher than when the mass was added over the back. Although the stride period was 2% longer in horses with loads on the distal limbs, time of contact and duty factor were not different. Distal limb loading increased the range of motion in hind- but not forelimbs. CONCLUSIONS: The costs of swinging the limbs in the horse are considerable and the addition of weights to the distal limb can have a profound effect on not only the energetics of locomotion but also the kinematics, at least in the hindlimb. POTENTIAL RELEVANCE: The use of weighted shoes, intended to increase animation of the gait, increases the metabolic effort of performance horses a disproportionate amount. The additional mass also increases the joint range of motion and, potentially, the likelihood of injury.

The cost of transport in an extended trot.

We hypothesised that trotters during an extended trot have lower energetic costs of locomotion (CT) than horses not bred for this behaviour. VO2 was measured as a function of speed in 7 Arabian horses (3 trained to extend their trotting speeds) and in 2 horses, of similar mass, bred to trot (Hackney). Both oxygen consumption and CT increased with speed and there was, contrary to our hypothesis, no difference between breeds. In Arabians at 6.5 m/s, CT had increased 25% above the CT at 5.0 m/s (normal transition speed). For Hackneys at 6.8 m/s, the CT was almost 35% higher. Stride frequencies increased linearly in all horses up to 5.0 m/s. At the canter at 5.0 m/s, the frequency increased 9% to 111 strides/min, but then increased minimally with speed. In the Hackneys and the Arabians that extended the trot, stride frequencies were approximately 102 and did not increase with speed. Stride length (SL) increased linearly with speed in both trotting and cantering horses, and cantering SL were lower than trotting (at 5.0 m/s, SL for trotting = 3.04 m and for cantering SL = 2.68 m). There were no differences between breeds in stride frequency or stride length. Extending the trot can have profound energetic requirements that could limit athletic performance and may lead to increased concussive impact on the limbs.

Effect of trotting speed, load and incline on hindlimb stance-phase kinematics.

The objective was to understand how the stance-phase kinematics of the hindlimb of trotting horses change with speed under 3 conditions (level, loaded and incline), to compare our results with the predictions of the spring-mass model and to help focus our future studies of muscle function. Video recordings were made of 5 Arabian horses trotting on a treadmill. Five consecutive strides were digitised and averaged for each trial. The angle-time diagrams were qualitatively similar to those reported previously. As speed increases, the range of motion of the hindlimb increases, as predicted by the spring-mass model. This is the result of increased range of motion in the coxofemoral and tarsal joints. The hindlimb does not 'land short-take off long'. When trotting up an incline, the hindlimb undergoes a greater range of motion because of increased retraction resulting from increased extension of the coxofemoral joint. At hoof contact on an incline, the 3 proximal joints are more flexed than on the level. Carrying a load had no effect on kinematics. These results suggest that there may be larger changes in strain with speed in muscles acting at the coxofemoral and tarsal joints than at the femorotibial joint, and that locomotion up an incline will change muscle strain more than carrying a load.

Hindlimb net joint energies during swing phase as a function of trotting velocity.

Net joint powers and energies have been described in walking horses during the swing phase of the stride in the fore- and hindlimb (Clayton et al. 2001). During trotting, swing phase net joint powers have been described in the forelimb but not in the hindlimb. The effects of velocity on power profiles and energy patterns are important in relation to locomotor energetics. The objective of this study was to evaluate velocity-dependent changes in hindlimb net energy profiles of the swing phase during trotting. Inverse dynamic analysis was used to calculate net joint energies at the hindlimb joints of 6 horses trotting overground at velocities ranging from 2.27-5.17 m/s. At all velocities, there was net energy generation at the hip and tarsus and net energy absorption at the stifle, fetlock and coffin joints. Velocity-dependent bursts of energy generation at the hip actively protracted the limb in early swing and initiated retraction in late swing. Synchronous with the bursts of energy generation at the hip were velocity-dependent bursts of energy absorption across the stifle that acted to control flexion in early swing and extension in late swing. The distal limb was raised and lowered by velocity-dependent bursts of energy generation that flexed the tarsus in early swing and extended it in late swing. The energy bursts in early swing increased linearly with velocity, whereas the energy bursts in late swing increased as a function of the square or cube of velocity. The results contribute to understanding the mechanisms used to accelerate and decelerate the limbs more rapidly as velocity increases, which is an important consideration in racing and sporting performance.

Effect of load on preferred speed and cost of transport.

Horses have a tendency to utilize a relatively narrow set of speeds near the middle of a much broader range they are capable of using within a particular gait, i.e., a preferred speed. Possible explanations for this behavior include minimizing musculoskeletal stresses and maximizing metabolic economy. If metabolic economy (cost of transport, CT) and preferred speeds are linked, then shifts in CT should produce shifts in preferred speed. To test this hypothesis, preferred speed was measured in trotting horses (n = 7) unloaded on the level and loaded with 19% of their body weight on the level. The preferred speed on the level was 3.33 +/- 0.09 (SE) m/s, and this decreased to 3.13 +/- 0.11 m/s when loaded. In both conditions (no load and load), the rate of O2 consumption (n = 3) was a curvilinear function of speed that produced a minimum CT (i.e., speed at which trotting is most economical). When unloaded, the speed at which CT was minimum was very near the preferred speed. With a load, CT decreased and the minimum was also near the preferred speed of horses while carrying a load.

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 decreased from 3.29+/-0.24 m s(-)(1) on the level to 3.05+/-0.30 m s(-)(1) (means +/- s.d., N=6) on an 11.8 % incline. The rate of oxygen consumption was measured as a function of trotting speed on a treadmill and was a curvilinear function of speed in all horses under both conditions (level and 10 % incline). This curvilinear relationship resulted in a C(T) that was a U-shaped function of speed. The speed at which C(T) was minimal (i.e. at which trotting was most energetically economical) was very near the preferred speed on the level and decreased on the incline, again to a speed near the preferred speed on the incline.

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 relationship are significantly different from 1.0, indicating that step length increases with speed and that small species use a step length that, relative to their leg length, is longer than the relative step length used by larger species. Time of contact increased when a horse carried a load but not when it trotted up an incline.

Disuse atrophy in the hibernating golden-mantled ground squirrel, Spermophilus lateralis.

Disuse (inactivity, bed rest, and spaceflight) may lead to a loss of muscle mass and a decrease in oxidative capacity in skeletal muscle. If such changes were to occur in hibernating animals, both locomotor and thermogenic function would be compromised. Muscle masses and oxidative capacities (as assessed by citrate synthase activity) were measured in the gastrocnemius and semitendinosus muscles, cardiac muscle (ventricle), and brown fat (axillary pad) in a group (n = 7) of prehibernating ground squirrels (Spermophilus lateralis) and after 6 mo of hibernation (n = 8). Hibernation produced significant atrophy in the gastrocnemius (14%) and semitendinosus (42%) muscles. Cardiac tissue increased (21%) in mass, as did brown adipose tissue (150%). That such changes were not due simply to fluid shifts was evidenced by similar protein concentrations between groups. In contrast to many other disuse studies, oxidative capacity was increased significantly in the gastrocnemius (65%) and semitendinosus (37%). Citrate synthase was also higher in cardiac tissue of hibernators (20%) but was not significantly different in brown fat.

A new model of avian embryonic metabolism.

A new model of avian embryonic metabolism is proposed that more accurately describes the available data. The model assumes that embryonic metabolism can be described by the equation VO2 = A X massB + C X growth rate. Empirically determined values of A, B, and C were used to predict the values of these parameters for the species whose preinternal pipping rate of O2 consumption (PIP) and total energy used during incubation (TOT) have been measured. Different patterns of growth were assumed for altricial and precocial species. Predicted values of PIP and TOT are highly correlated with observed values (r2 = 0.966 and 0.979, respectively). The model predicts the pattern of metabolism in precocial species will vary with hatchling mass and incubation period; TOT will be greater in eggs with longer incubation periods and greater in precocial species than in altricial species because 70-80% of TOT is devoted to maintenance; PIP will be relatively insensitive to interspecific differences in incubation period, because the 80% of PIP devoted to maintenance is insensitive to incubation period; the energy cost of biosynthesis will average only one fifth of the energy content of the tissues produced; and the maintenance metabolic rate of an avian embryo is more like that of an adult reptile than that of an adult bird. Shell conductance may not be exactly inversely proportional to incubation period, because it represents a compromise between the requirements for limiting water loss and permitting adequate exchange of respiratory gases. The model overestimates PIP and TOT in species with very long incubation periods; these embryos may have unusually low metabolic intensities.

Respiration of avian embryos - a comparative analysis.

All published values of the metabolic rates of avian embryos are brought together for analysis. A generalized pattern of O2 consumption during development of precocial eggs is determined and used to estimate the total amount of O2 consumed duirng development. The metabolic rates of avian eggs just prior to internal pipping (preIP MO2 is inversely proportional to the length of the incubation period. As in mammals, the mass specific metabolism of avian embryos is approximately equal to that of their parents, even though the parents are at least five times heavier. The total amount of O2 consumed per gram of fresh egg mass during development appears to be independent of egg mass and incubation period and averages 102 cm3 O2 . G-1. Calculated air cell O2 and CO2 tensions average 101 Torr and 40 Torr, respectively. Calculated air cell gas tensions are significantly correlated with egg mass but directly observed values are not, a discrepancy which remains to be resolved.