Effects of muscle glycogen depletion on some metabolic and physiological responses to submaximal treadmill exercise.

The aim of this study was to investigate the effects of reduced muscle glycogen concentration on some physiological and metabolic responses during moderate intensity treadmill exercise in horses. Six Thoroughbred geldings were randomly allocated to 2 treatments (protocols A and B) or control in a 3 x 3 replicated Latin square design. In protocol A, horses performed low intensity exercise while horses in protocol B performed short bursts of high intensity exercise. Protocol A was designed to induce glycogen depletion mainly of slow twitch muscle fibers while protocol B aimed to deplete mainly fast twitch muscle fibers. Horses in the control group did not undergo exercise prior to the exercise test. Five hours after glycogen depletion, horses performed treadmill exercise at 60% VO2max at a treadmill slope of 10% until fatigue (20-30 min). The induced glycogen depletion prior to exercise had no significant effect on plasma glucose, insulin, or lactate concentrations during the exercise test, and there was no effect on glycogen utilization rate, although respiratory exchange ratios were lower in the glycogen-depleted groups. The VO2, heart rate and central blood temperature did not vary significantly between the protocols A and B and control throughout the exercise test. It was concluded that 20-30% depletion of glycogen concentration in the middle gluteal muscle resulted in a shift towards fat metabolism, but does not significantly affect heart rate, oxygen uptake, or concentrations of plasma glucose and lactate during moderate intensity exercise.     

Effects of cool and hot humid environmental conditions on neuroendocrine responses of horses to treadmill exercise

To determine the effects of exercise, high heat and humidity and acclimation on plasma adrenaline, noradrenaline, beta-endorphin and cortisol concentrations, five horses performed a competition exercise test (CET; designed to simulate the speed and endurance test of a three-day event) in cool dry (CD) (20 degrees C/40% RH) and hot humid (30 degrees C/80% RH) conditions before (pre-acclimation) and after (post-acclimation) a 15 day period of humid heat acclimation. Plasma adrenaline and noradrenaline concentrations pre-acclimation were significantly increased compared with exercise in the CD trial at the end of Phases C (P<0.05) and D (P<0.05 and P<0.01, respectively) and at 2 min recovery (P<0.01), with adrenaline concentrations still elevated after 5 min of recovery (P<0.001). Plasma beta-endorphin concentrations were increased at the end of Phases C (P<0.05) and X (P<0.01) and at 5 and 30 min recovery (P<0.05) in the pre-acclimation session. Plasma cortisol concentrations were elevated after the initial warm up period pre-acclimation (P<0.01) and at the end of Phase C (P<0.05), compared with the CD trial. A 15 day period of acclimation significantly increased plasma adrenaline concentrations at 2 min recovery (P<0.001) and plasma cortisol concentration at the end of Phase B (P<0.01) compared with pre-acclimation. Acclimation did not significantly influence noradrenaline or beta-endorphin responses to exercise, although there was a trend for plasma beta-endorphin to be lower at the end of Phases C and X and after 30 min recovery compared with pre-acclimation. Plasma adrenaline, noradrenaline, beta-endorphin and cortisol concentrations were increased by exercise in cool dry conditions and were further increased by the same exercise in hot humid conditions. Exercise responses post-acclimation suggest that adrenaline and noradrenaline may play a role in the adaptation of horses to thermal stress and that changes in plasma beta-endorphin concentrations could be used as a sensitive indicator of thermal tolerance before and after acclimation. The use of plasma cortisol as a specific indicator of heat stress and thermal tolerance before or after acclimation in exercising horses appears limited.     

Physiological responses to treadmill exercise and ambient temperature in normal and malignant hyperthermia susceptible pigs.

Ten susceptible and ten resistant pigs to malignant hyperthermia were used to observe the effects of exercise and ambient temperature on selected physiological parameters. Pigs were submitted to a ten minute exercise on a treadmill operating at a speed of 1.8 km/h and inclined to 11 degrees. Exercise in the first group was at an ambient temperature of 14 degrees C, and in the second at 29 degrees C. The right carotid artery was previously cannulated for blood pressure measurements and for repeated blood sampling during exercise. Arterial pressure, heart rate, rectal and cutaneous temperatures were recorded. Levels of cortisol, creatine kinase and its isoenzymes were measured. At 14 degrees C, exercise caused some physiological adjustments in susceptible animals; heart rate, skin temperature and cortisol levels increased (P less than 0.05). In resistant pigs, only the heart rate was elevated significantly following exertional stress at 14 degrees C. Exercise at 29 degrees C produced severe stress and marked physiological changes: heart rate, rectal and skin temperatures and cortisol levels increased significantly in both susceptible and resistant swine. At 29 degrees C, susceptible pigs also had higher levels of serum cortisol, total creatine kinase and MM isoenzyme (P less than 0.05) compared to resistant pigs. The results indicate that, following exertional or thermal stress, susceptible pigs undergo more extensive physiological changes than do resistant pigs. Similar levels of stress prior to slaughter may trigger physiological changes which in the susceptible pigs would likely result in pale, soft exudative myopathy.     

Comparison of tracheal aspirates before and after high-speed treadmill exercise in racehorses

OBJECTIVE: To determine whether percentages of neutrophils in tracheal aspirate (TA) samples collected from racehorses are increased after exercise and whether interpretation of results from TA samples taken before and after exercise agree. DESIGN: Case series of 40 young Thoroughbred and Standardbred racehorses in race training presented for evaluation of poor performance. PROCEDURE: TA samples were collected endoscopically from racehorses presented for poor performance 24 h before and 1 to 2 h after high speed treadmill exercise testing. Aliquots of the retrieved fluid were cytocentrifuged and smears were stained with Diff-Quik. Mean neutrophil counts were expressed as percentages of the total number of inflammatory cells counted and subsequently were categorised as either above or below an accepted cut-off of 20%. Comparisons between percentages of neutrophils before and after exercise were made. RESULTS: Percentage of neutrophils from TA samples obtained from racehorses after exercise was significantly higher than neutrophil percentages from TA samples collected from the same horse before exercise. In horses with TA specimens that were categorised as having < or = 20% neutrophils before treadmill exercise, the percentage of neutrophils in their TA specimens after exercise was, on average, significantly higher and was greater than the cut-off value of 20%. CONCLUSION: Recent strenuous exercise may change the proportion of neutrophils in lower airways of racehorses and practitioners should be aware of this when collecting and interpreting the results from TA samples. The most practical time for collection of a TA sample to obtain the most diagnostically useful information might be after a suitable washout period of at least 1 to 2 h post-exercise.     

The effects of short intensive exercise on plasma free amino acids in standardbred trotters

The aim of this study was to investigate the effect of short intense exercise on plasma amino acid concentrations in trotters and to test the repeatability of plasma amino acids concentration in samples obtained on two independent days under field conditions. Plasma amino acid concentrations were analysed in blood samples of 36 standardbred trotters before and after intense exercise over a distance of 2000 m. Sampling was repeated in 20 horses after 35 days. Exercise intensity was estimated from post‐exercise lactate levels. Horses were divided in two groups according to a cut‐off lactate concentration at 15 mmol/l. The plasma concentrations of alanine, aspartate, glutamate, isoleucine, leucine, lysine and taurine increased and arginine, asparagine, citrulline, glutamine, glycine, histidine, methionine, serine, tryptophan and 3‐methylhistidine decreased after exercise. Ornithine, threonine, tyrosine, phenylalanine and valine concentrations remained constant. Higher intensity of exercise significantly decreased tryptophan and increased taurine concentrations. Sampling day had a significant effect on the absolute pre‐ and post‐exercise amino acid concentrations. Exercise had a significant influence on the concentrations of most plasma amino acids in trotters. These changes could reflect shifts between the free amino acid compartments, but there were also some indications for muscle catabolism. The amino acid supply of sporting horses could be of specific significance for maintaining muscle integrity and for the improvement of post‐exercise recovery of competition horses.     

Effects of two field continuous incremental tests on cardiorespiratory responses in Standardbred trotters

In humans, cardiorespiratory responses are widely evaluated from field incremental exercise tests. On the contrary, equine exercise physiology faces a huge lack of oxygen consumption measurements (VO₂) in field conditions due to technical concerns. The aim of this study was to test the effects of two incremental continuous field tests on cardiorespiratory responses in Standardbred trotters. The two protocols were realized at trot and ended when horses galloped. The tests started at 4.2 m/s (T1) and 6.4 m/s (T2), with speed increments of 1.4 m/s every 3 min for T1 and 0.8 m/s every 2 min for T2. Velocity (v), heart rate (HR) and gas exchanges were recorded continuously, and blood lactate concentration [La^?] was measured before and after tests. Values recorded at the end of the tests were considered as peak values. The v_peak values were 10.6 ± 0.3 and 10.7 ± 0.7 m/s for T1 and T2 respectively. Horses reached higher VO_2peak (T1: 116.6 ± 11.5 ml/min/kg; T2: 88.9 ± 10.2 ml/min/kg; p < 0.05) and HR_peak (T1: 217 ± 5 bpm; T2: 209 ± 3 bpm; p < 0.05) during T1 compared with T2. T1 was significantly longer than T2 (17.5 ± 1.9 vs. 12.9 ± 1.6 min respectively, p < 0.01), and the number of steps entirely ran tended to be different (T1: 5.6 ± 0.6; T2: 6.2 ± 0.8, p = 0.07). Compared to T2, the design of T1 appeared easier to implement and allowed higher cardiorespiratory responses. The relationship between HR‐VO₂ obtained through T1 gave a better correlation between the two variables than T2. These findings suggest that T1 might be better than T2 for evaluating cardiorespiratory adaptations to exercise and for estimating aerobic energy expenditure in exercising trotters.     

Respiratory responses to exercise in the horse

Horses are elite athletes when compared with other mammalian species. In the latter, performance is limited by cardiovascular or musculoskeletal performance whereas in athletic horses it is the respiratory system that appears to be rate limiting and virtually all horses exercising at high intensities become hypoxaemic and hypercapnoeic. This is due to both diffusion limitation and a level of ventilation inadequate for the metabolic level that enables horses to exercise at these intensities. In conjunction with these blood gas changes, total pulmonary resistance increases and the work of breathing rises exponentially and airflow eventually plateaus despite increases in inspiratory and expiratory intrapleural pressures. Horses breathe at comparatively high frequencies when galloping due to the tight 1:1 coupling of strides to breathing. Whether this effects gas exchange and, if so, to what extent, has not been fully elucidated.     

The effect of interval versus continuous exercise on plasma leptin and ghrelin concentration in young trotters

The effect of interval vs. continuous exercise on plasma leptin and ghrelin concentration in young Standardbred horses was studied. The experiment was conducted on 27 trotters, in the age between 2 and 3 years. They were divided into two groups according to the type of exercise. Blood samples were collected through jugular venipuncture in the following experimental conditions: at rest, immediately after exercise and 30 minutes after the end of the effort. Plasma leptin and ghrelin concentrations were determined using RIA tests. The continuous exercise induced an increase in plasma leptin concentration whereas the interval type of exercise did not influence the level of this hormone (3.47 +/- 0.78 vs. 4.07 +/- 0.94 and 2.31 +/- 0.15 vs. 2.36 +/- 0.21 ng/mL, respectively). The plasma ghrelin concentration measured after the continuous exercise, significantly increased (720 +/- 27.4 vs. 814 +/- 13.8; p < or = 0.05) whereas concentration of this hormone assessed after the interval exercise, significantly dropped (982 +/- 56.5 vs. 842 +/- 35.6 pg/mL; p < or = 0.05). The changes in plasma ghrelin concentration measured after the end of the effort correlated inversely with blood lactic acid concentration. In conclusion, the obtained results showed that medium-intensive type of exercise, such as trot, interval or continuous, slightly affected plasma leptin level but significantly affected plasma ghrelin concentration in young Standardbred trotters.     

Effects of phlebotomy on haemodynamic characteristics during exercise in standardbred trotters with red cell hypervolaemia

Five Standardbred trotters with red cell hypervolaemia (RCHV) were compared before and after removal of approximately 22% (36 ml/kg bwt) of the total blood volume in order to evaluate the haemodynamic responses, haemorheological alterations and oxygen transport during exercise to fatigue. Data were recorded during submaximal exercise at 4 different speeds on a treadmill and then during continued running at the highest speed step until fatigue. Oxygen uptake (VO2), pulmonary artery pressure (PAP), systemic artery pressure (SAP), heart rate (HR), haematocrit and haemoglobin concentrations (Hb) were measured. Arteriovenous O2 content difference (C(a-v)O2), pulmonary vascular resistance (PVR) and total systemic resistance (TSR) were calculated. Whole blood and plasma viscosity and erythrocyte aggregation tendency were determined with a rotational viscometer. Endoscopy was performed after exercise. ANOVA was used for statistical analysis. Phlebotomy resulted in a decrease in haematocrit and Hb during the course of exercise. Blood and plasma viscosity were lower and erythrocyte aggregation tendency was higher after phlebotomy. Throughout exercise, including submaximal work and continued running to fatigue, PAP, SAP, PVR, TSR and C(a-v)O2 were lower after phlebotomy. HR was higher after phlebotomy during submaximal exercise. Oxygen delivery and VO2 were lower after phlebotomy in the period from submaximal exercise to fatigue. Run time to fatigue was shorter after phlebotomy. Four horses showed exercise-induced pulmonary haemorrhage (EIPH) before phlebotomy and the degree of bleeding was diminished but not abolished after phlebotomy. The reductions in PVR, TSR, PAP and SAP after phlebotomy were probably a result of reduced blood viscosity. In conclusion, although a 22% reduction in blood volume improved the haemodynamic and haemorheological parameters and the degree of EIPH, it was found that RCHV trotters have to rely on high oxygen delivery to the working muscles for maintenance of maximal performance.     

Effects of treadmill exercise on sweating in three breeds of goats

OBJECTIVE: To compare the sweating responses of three breeds of goats to exercise at 30 degrees C. DESIGN: Factorial experiment with two goats of each of three breeds exercised for 60 min at 3 km/h and 30 degrees C on 6 days. PROCEDURE: Two mature females of the Anglo-Nubian, Saanen and Toggenburg breeds were used. Rectal temperature, respiration rate and sweating rate at three sites were recorded every 20 min during six replicates of exercise at 48 h intervals on a treadmill. RESULTS: Respiration rate varied with time and breed (P < 0.01) and increased from 20 +/- 2 breaths/min to 135, 195 and 260/min after 60 min exercise in Anglo-Nubian, Saanen and Tottenburg goats, respectively. Bread differences in rectal temperature were small but significant (P < 0.001), and mean values increased from 38.9 degrees C before exercise to 39.7 degrees C after 1 h exercise. The breed x sites interaction for sweating was significant (P < 0.01). On the rump breed differences in sweating rate were not significant. On the loin, Toggenburg goats started to sweat most rapidly and reached maximal values of 80.2 +/- 10.6 g/m2/h after 20 min and then decreased to 70.0 +/- 4.3 g/m2/h at 60 min. Sweating rate on the ear was highest in Toggenburg goats, followed by those of Saanen and Anglo-Nubian goats (P < 0.05), but the differences were small (7 g/m2/h); peak values of 67.3 to 76.1 g/m2/h were recorded after 20 min. CONCLUSION: Respiration and sweating rates increased significantly during exercise in all three breeds of goats, but breed differences were marked only for respiration rate. The goats sweated more on the rump than on the loin on ear, with peak values after 40 min of exercise.     

The opioid haemorphin-7 in horses during low-speed and high-speed treadmill exercise to fatigue

The opioid neuropeptide haemorphin-7 was measured, by immunoreactivity, in Standardbred horses during low-speed (7 m/s) and high-speed (10 m/s) endurance exercises, lasting 49-58 and 12-16 min respectively. In parallel, heart rate, muscle temperature and plasma lactate concentrations were measured. The profile of the low-speed exercise showed significantly increased heart rate after 10 min [154 beats per minute (bpm)]. After the exercise, muscle temperature (42.1 degrees C) and plasma lactate (4.8 mmol/l) were significantly increased. The profile of the high-speed exercise was comparatively characterized by a higher increase of heart rate after 5 min (194 bpm) and higher increases of muscle temperature (43.2 degrees C) and lactate levels (15.8 mmol/l) after the exercise. The horses were probably exhausted by glycogen depletion in the low-speed exercise and by muscle pH decrease in the high-speed exercise. Haemorphin-7 increased significantly during the high-speed exercise (274.8 fmol/ml) but not during low speed (108.3 fmol/ml), coincident with the results of lactate. These results suggest that plasma haemorphin-7 is measurable in the horse by immunoreactivity, and that intense exercise stimulates release of this opioid. Such endogenous opioids are most likely involved in regulatory functions associated with pain, physical effort, inflammation, and blood pressure variation in horses, as have been established in other species.     

Effects of treadmill exercise on cortical bone in the third metacarpus of young horses.

The effects of exercise and relative inactivity on cortical bone were compared in young horses. Two groups were used; one was given a 14-week programme of exercise (n = 6) and the other kept as unexercised controls (n = 6). The first nine weeks of exercise involved trotting and cantering (2 to 4 km d-1 at speeds up to 12 m s-1) on a treadmill set at an incline of 3 degrees. Over the next five weeks the horses were trained at near maximal speeds (that is, up to 14.5 m s-1) with no incline of the treadmill. At the end of the programme marked differences in cortical porosity and distribution of subperiosteal osteogenesis at the mid-shaft of the third metacarpal bone were found between the groups. Histomorphometrical examination of the dorsal cortex showed minimal bone remodelling in the exercised horses, but extensive modelling as evidenced by the large amount of subperiosteal bone formation. In contrast, the unexercised horses had significantly more bone remodelling and less formation of subperiosteal bone. The histomorphometric and microradiographic findings provided an explanation for changes in the non-invasive bone measurements that occurred during training. Bone mineral content of the mid-metacarpus was found to increase more in the exercised than the unexercised horses despite a lower overall growth in bodyweight. In those horses that completed the full training programme, ultrasound speed increased significantly by the end of the training programme. It remained unchanged in the horse that did not complete the full exercise programme and decreased slightly in the unexercised horses. The difference in ultrasound speed between the groups was considered to reflect differences in intracortical bone porosity, endosteal bone formation and alterations in skin thickness. The stiffness of cortical bone increased significantly in the exercised horses but remained unaltered in the unexercised horses.     

Comparative ergoespirometric adaptations to a treadmill exercise test in untrained show Andalusian and Arabian horses

Significant differences exist in the respiratory adaptation to exercise in different equine breeds. This research describes the ergoespirometric response to exercise of Andalusian (AN) and Arabian (A) horses, both selected according to morphological criteria. Thirteen untrained male horses (6 AN and 7 A) performed a treadmill exercise test (TET) with a slope of 6%, with workloads starting from 5 m/s and increasing 1 m/s every 3 min until the horses were not able to keep the required velocity. Tidal volume (TV), respiratory rate, minute ventilation (VE), oxygen uptake (VO2), carbon dioxide production, peak oxygen uptake (VO2peak), respiratory exchange ratio (RER), exercise time to fatigue (ETF) and respiratory aerobic threshold (RAT) were determined. AN horses presented higher TV and VE, whereas respiratory rate, VO2 and VCO2 were lower at the same velocities. RER was similar between breeds. ETF was longer in A horses (556.7 ± 66.5 in AN vs. 607.1 ± 71.1 s in A) and no significant differences were found in RAT (5.50 ± 0.50 in AN vs. 5.86 ± 1.07 m/s in A). In summary, despite the more intense ventilatory response to exercise at the same velocity, AN horses had lower VO2. The AN horse develops a more intense ventilatory response to fixed velocities than the A horse and it could be interesting to clarify the role of the locomotion characteristics in this response.     

Effect of creatine supplementation on muscle metabolic response to a maximal treadmill exercise test in Standardbred horses

The aim of the present study was to investigate the effect of creatine (Cr) supplementation on muscle metabolic response in connection with a maximal treadmill exercise test, known to cause a marked anaerobic metabolic response and adenine nucleotide degradation. First, 6 Standardbred trotters performed a standardised maximal exercise test until fatigue (baseline test). The test used was an inclined incremental treadmill test in which the speed was increased by 1 m/s, starting at 7 m/s, every 60 s until the horse could no longer keep pace with the treadmill. After this baseline test, the horses were separated into 2 equal groups. One half received a dose of 25 g creatine monohydrate twice daily, and the other group were given the same dose of lactose (placebo). The supplementation period was 6.5 days, after which the maximal treadmill exercise test was performed again. A washout period of 14 days was allowed before treatments were switched between groups and a new supplementation period started. After this second supplementation period a new maximal exercise test was performed. After supplementation with creatine or placebo, horses were stopped after performing the same number of speed steps and duration of exercise as they had in the baseline test. Blood samples for analysis of plasma lactate, creatine (Cr), creatinine, hypoxanthine, xanthine and uric acid concentrations were collected at rest, during each speed step and during recovery. The total blood volume (TBV) was also determined. Muscle biopsies for analysis of muscle metabolites (adenosine triphosphate [ATP], adenosine diphosphate [ADP], adenosine monophosphate [AMP], inosine monophosphate [IMP], creatine phosphate [CP], lactate [La] and glycogen) were taken at rest, immediately post exercise and after 15 min recovery. The results showed no significant increase in plasma Cr or muscle total creatine concentration (TCr) after supplementation with Cr. At the end of exercise ATP and CP concentrations had decreased and IMP and lactate concentrations increased in muscle in all groups. Plasma lactate concentration increased during exercise and recovery and plasma uric acid concentration increased during recovery in all groups. No influence could be found in TBV after supplementation with creatine. These results show that creatine supplementation in the dosage used in this study had no influence on muscle metabolic response or TBV.     

Stress resistant and stress susceptible landrace pigs: comparison of blood variables after exposure to halothane or exercise on a treadmill

Stress susceptible pigs develop pale, soft, exudative pork as a consequence of exposure to natural stress or exposure to halothane. Hyperstimulation of glycolysis is a major component of the stress reaction. Whether natural and artificial stress have other similarities was investigated by exposing pigs to halothane or to physical exercise, and measuring their blood composition. There were many similarities between the two groups but the significant differences suggest that although both halothane and exercise induce glycolysis, the mechanisms involved are different.