Glycogen is a polysaccharide that serves as a primary form of energy storage in horses, predominantly found in the liver and muscle tissues. It acts as a readily available energy source that can be mobilized during periods of increased metabolic demand, such as exercise or stress. The synthesis and breakdown of glycogen are regulated by hormonal and enzymatic pathways, which are influenced by factors like diet, exercise, and overall health status. Understanding glycogen metabolism is important for optimizing performance, managing dietary needs, and preventing metabolic disorders in equine populations. This page compiles peer-reviewed research studies and scholarly articles that explore the role, regulation, and physiological implications of glycogen in equine health and performance.
Essén-Gustavsson B, Jensen-Waern M.Glycogenolysis and lipolysis are essential for energy production in muscle during prolonged exercise but less is known about the role of amino acid metabolism. The aim was to study the effect of an endurance race on pro- and macroglycogen, triglycerides and amino acid concentrations in muscle and on selected blood parameters, especially the amino acid response in the blood during the recovery phase. Seven endurance-trained horses completed a race covering 100-120 km. Blood samples were collected one day before the race, after the finish and 1 and 8 days after the race. Needle biopsy specimens ...
Rivero JL, Sporleder HP, Quiroz-Rothe E, Vervuert I, Coenen M, Harmeyer J.The purpose of this study was to determine whether oral L-carnitine supplementation enhances the responses of skeletal muscle to training in seven 2-year-old Standardbreds. Four horses were supplemented with 10 g/day L-carnitine for 10 weeks and 3 horses served as controls. All horses were exercised regularly every second day on a treadmill for 5 weeks (training period) and housed in individual boxes for 5 additional weeks (detraining period). The training period consisted of 8 high- and 8 low-speed exercises carried out in alternating sequence. Gluteus medius muscle biopsies were taken at Wee...
Palmgren Karlsson C, Jansson A, Essén-Gustavsson B, Lindberg JE.The aim of this study was to investigate the effect of partly replacing oats with molassed sugar beet pulp in a traditional hay/oat diet on nutrient utilisation and metabolic parameters in exercising horses. In a change-over experiment, 4 Standardbred geldings were fed a hay and oat-based diet (Oat diet) and a hay and oat-based diet where oats was partially replaced with molassed sugar beet pulp (MSBP diet). Each experimental period was 21 days during which total collection of faeces and urine was made and an exercise test (ET) performed. The crude fat digestibility was lower on the MSBP diet ...
Schuback K, Essén-Gustavsson B, Persson SG.Administration of bicarbonate has been shown to cause metabolic alkalosis both in man and in horses and is, therefore, thought to increase the buffering capacity of the body and thereby delay the onset of fatigue. However, results regarding the influence of sodium bicarbonate loading on performance both in human athletes and in horses are conflicting. The aim of this study was, therefore, to investigate the metabolic response to a standardised treadmill exercise test to fatigue, in horses given bicarbonate (0.6 g/kg bwt), in comparison to horses given placebo (water). Five Standardbred trotter...
Bröjer JT, Stämpfli HR, Graham TE.The objective of this study was to determine whether the concentrations of proglycogen (PG) and macroglycogen (MG) in biopsy samples of horse muscle are influenced by extraction time or perchloric acid (PCA) concentration. In study 1, individual muscle-biopsy samples from 10 horses were divided into 4 parts each and then randomly subjected to 4 periods of extraction (10, 20, 60, or 120 min) with 1.5 M PCA. In study 2, individual muscle-biopsy samples from 6 horses were divided into 24 pieces each and then randomly subjected to 12 combinations of extraction time (10, 20, 30, or 40 min) and PCA ...
Bröjer JT, Stämpfli HR, Graham TE.To determine proglycogen (PG) and macroglycogen (MG) content in equine skeletal muscle and to compare 2 analytical methods (acid hydrolysis [AC] and PG plus MG determination) for measurement of total muscle glycogen content (Gly(tot)) in biopsy specimens. Methods: Muscle biopsy specimens obtained from 41 clinically normal horses. Methods: Forty-five muscle biopsy specimens obtained from the middle gluteal (n = 31) or triceps (14) muscle were analyzed, using AC and MG plus PG determination for Gly(tot). Variability within muscle biopsy specimens for each method was calculated from duplicate ana...
Valberg SJ, Ward TL, Rush B, Kinde H, Hiraragi H, Nahey D, Fyfe J, Mickelson JR.Seven related Quarter Horse foals that died by 7 weeks of age were examined for glycogen branching enzyme (GBE) deficiency. Clinical signs varied from stillbirth, transient flexural limb deformities, seizures, and respiratory or cardiac failure to persistent recumbency. Leukopenia (5 of 5 foals) as well as high serum creatine kinase (CK; 5 of 5), aspartate transaminase (AST; 4 of 4), and gamma glutamyl transferase (GGT; 5 of 5) activities were present in most foals, and intermittent hypoglycemia was present in 2 foals. Gross postmortem lesions were minor, except for pulmonary edema in 2 foals....
Hinchcliff KW, Lauderdale MA, Dutson J, Geor RJ, Lacombe VA, Taylor LE.High intensity exercise is associated with production of energy by both aerobic and anaerobic metabolism. Conditioning by repeated exercise increases the maximal rate of aerobic metabolism, aerobic capacity, of horses, but whether the maximal amount of energy provided by anaerobic metabolism, anaerobic capacity, can be increased by conditioning of horses is unknown. We, therefore, examined the effects of 10 weeks of regular (4-5 days/week) high intensity (92+/-3 % VO2max) exercise on accumulated oxygen deficit of 8 Standardbred horses that had been confined to box stalls for 12 weeks. Exercise...
Jose-Cunilleras E, Hinchcliff KW, Sams RA, Devor ST, Linderman JK.In a randomized, balanced, crossover study each of six fit, adult horses ran on a treadmill at 50% of maximal rate of oxygen consumption for 60 min after being denied access to food for 18 h and then 1) fed corn (51.4 kJ/kg digestible energy), or 2) fed an isocaloric amount of alfalfa 2-3 h before exercise, or 3) not fed before exercise. Feeding corn, compared with fasting, resulted in higher plasma glucose and serum insulin and lower serum nonesterified fatty acid concentrations before exercise (P < 0.05) and in lower plasma glucose, serum glycerol, and serum nonesterified fatty acid conce...
Quiroz-Rothe E, Rivero JL.Combined methodologies of immunohistochemistry, histochemistry and photometric image analysis were applied: (1) to characterise control equine skeletal muscle fibres according to their myosin heavy chain (MyHC) composition and (2) to determine on a fibre-to-fibre basis the correlation between contractile [i.e. MyHC(s), myofibrillar ATPase (mATPase) and sarco(endo)plasmic reticulum Ca(2+)-ATPase (SERCA) isoforms], metabolic [i.e. succinate dehydrogenase (SDH) and alpha-glycerophosphate dehydrogenase (GPD) activities, glycogen and phospholamban (PLB) contents], and morphological [i.e. cross-sect...
Lacombe VA, Hinchcliff KW, Geor RJ, Baskin CR.The purpose of this study was to determine the effect of muscle glycogen depletion and subsequent replenishment on anaerobic capacity of horses. In a blinded crossover study, seven fit horses performed glycogen-depleting exercise on two occasions. Horses were infused after glycogen-depleting exercise with either 6 g/kg body wt of glucose as a 13.5% solution in 0.9% NaCl (Glu) or with 0.9% NaCl (Sal) of equivalent volume. Subsequently, horses performed a high-speed exercise test (120% of maximal rate of oxygen consumption) to estimate maximum accumulated oxygen deficit. Replenishment of muscle ...
Hyyppä S.To test the effect of nandrolone on their recovery, six adult half-bred riding horses performed a competition exercise test (CET) and a standardized exercise test (SET) on consecutive days before and after a 2-week treatment with the anabolic steroid nandrolone laurate. Blood samples were collected during and between these tests for the determination of red cell volume and concentrations of blood lactate, plasma glucose, non-esterified fatty acids, glycerol, triglycrides, erythropoietin, cortisol, insulin, and glucagon. Muscle biopsy specimens were taken immediately after the CET and before th...
Klomkleaw W, Kasashima Y, Kobayashi A, Fuller GA, Morimoto M, Nakade T, Muto M, Oba T, Hamlin RL, Yamaguchi M.Tubular aggregates (TAs) originate from the sarcoplasmic reticulum (SR) and form polymorphic double (or single) -walled structures in cross section. TAs are involved in various human skeletal muscle disorders including periodic paralysis, congenital myasthenic syndromes, inflammatory myopathies, and malignant hyperthermias. Horse lumbrical muscle (LM) is a slender fusiform muscle that shows varying degrees of regression due to its limited activity in the limb. Double-walled TAs were found in degenerating spindle fibers and with a range of 80-116 nm (average 92 nm, n=135) for outer layer and 50...
Geelen SN, Blázquez C, Geelen MJ, Sloet van Oldruitenborgh-Oosterbaan MM, Beynen AC.The metabolic effects of feeding soyabean oil instead of an isoenergetic amount of maize starch plus glucose were studied in ponies. Twelve adult Shetland ponies were given a control diet (15 g fat/kg DM) or a high-fat diet (118 g fat/kg DM) according to a parallel design. The diets were fed for 45 d. Plasma triacylglycerol (TAG) concentrations decreased by 55 % following fat supplementation. Fat feeding also reduced glycogen concentrations significantly by up to 65 % in masseter, gluteus and semitendinosus muscles (P < 0.05 and P < 0.01 and P < 0.01 respectively). The high-fat diet significan...
Geelen SN, Sloet van Oldruitenborgh-Oosterbaan MM, Beynen AC.Feeding fat-supplemented diets to horses has drawn considerable interest. One of the advantages of such diets is that the energy density is increased, so that less feed is needed to meet energy requirements. In addition, adding fat to the diet enhances the contribution of fat oxidation to energy production, thus sparing muscle glycogen. The 'spared' glycogen is available for energy metabolism when the acutely exercising horse reaches a point of oxygen deficit and must rely on anaerobic metabolism. This appears to be beneficial for both aerobic and anaerobic performance. Fats are readily digest...
Valentine BA, Habecker PL, Patterson JS, Njaa BL, Shapiro J, Holshuh HJ, Bildfell RJ, Bird KE.Skeletal muscle samples from 38 draft horse-related animals 1-23 years of age were evaluated for evidence of aggregates of glycogen and complex polysaccharide characteristic of equine polysaccharide storage myopathy (EPSSM). Cardiac muscle from 12 of these horses was also examined. Antemortem serum levels of creatine kinase (CK) and aspartate aminotransferase (AST) from 9 horses with EPSSM and 5 horses without EPSSM were compared. Skeletal muscle from 17 horses contained inclusions of periodic acid-Schiff (PAS)-positive, amylase-resistant complex polysaccharide. Similar inclusions were also pr...
Serrano AL, Quiroz-Rothe E, Rivero JL.Twenty-four 4-year-old Andalusian (Spanish breed) stallions were used to examine the plasticity of myosin heavy chain (MHC) phenotype and the metabolic profile in horse skeletal muscle with long-term endurance-exercise training and detraining. Sixteen horses underwent a training programme based on aerobic exercises for 8 months. Afterwards, they were kept in paddocks for 3 months. The remaining eight horses were used as controls. Three gluteus medius muscle biopsy samples were removed at depths of 20, 40 and 60 mm from each horse before (month 0), during (month 3) and after (month 8) training,...
Schuback K, Essén-Gustavsson B, Persson SG.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 horse...
Geor RJ, Hinchcliff KW, Sams RA.To examine the role of beta-adrenergic mechanisms in the regulation of endogenous glucose (Glu) production [rate of appearance (R(a))] and utilization [rate of disappearance (R(d))] and carbohydrate (CHO) metabolism, six horses completed consecutive 30-min bouts of exercise at approximately 30% (Lo) and approximately 60% (Hi) of estimated maximum O(2) uptake with (P) and without (C) prior administration of the beta-blocker propranolol (0.22 mg/kg iv). All horses completed exercise in C; exercise duration in P was 49.9 +/- 1.2 (SE) min. Plasma Glu was unchanged in C during Lo but increased prog...
Geor RJ, Hinchcliff KW, McCutcheon LJ, Sams RA.This study examined the effects of preexercise glucose administration, with and without epinephrine infusion, on carbohydrate metabolism in horses during exercise. Six horses completed 60 min of treadmill exercise at 55 +/- 1% maximum O(2) uptake 1) 1 h after oral administration of glucose (2 g/kg; G trial); 2) 1 h after oral glucose and with an intravenous infusion of epinephrine (0.2 micromol. kg(-1). min(-1); GE trial) during exercise, and 3) 1 h after water only (F trial). Glucose administration (G and GE) caused hyperinsulinemia and hyperglycemia ( approximately 8 mM). In GE, plasma epine...
Valentine BA, McDonough SP, Chang YF, Vonderchek AJ.Polysaccharide storage myopathy is an equine neuromuscular disorder characterized by accumulation of glycogen-related polysaccharide inclusions within skeletal muscle fibers. The pathologic criteria for diagnosis of this disorder are somewhat controversial; however, periodic acid-Schiff-positive, amylase-resistant inclusions are considered pathognomonic. Although these inclusions are most often found in affected horses related to the Quarter Horse, draft horse, and Warmblood breeds, this report describes these characteristic inclusions in muscle of five horses from nonrelated breeds (two Morga...
De La Corte FD, Valberg SJ, Mickelson JR, Hower-Moritz M.Polysaccharide storage myopathy (PSSM) in Quarter Horses (QH) and QH crosses is a glycogen storage disorder in which blood glucose clearance and insulin sensitivity, following an i.v. or oral glucose challenge, are enhanced. Exercise is known also to enhance glucose uptake into skeletal muscle in many animal species. Therefore, the purpose of this study was to compare the effect of exercise on glucose clearance in PSSM and control horses when an oral carbohydrate meal (8 Mcal sweet feed) was fed following either 12 h fasting alone (NEX protocol) or following fasting and a standard exercise pro...
Essén-Gustavsson B, Gottlieb-Vedi M, Lindholm A.The aim was to investigate metabolic response in muscle during submaximal treadmill exercise to fatigue, with a special emphasis on adenine nucleotide degradation products such as inosine monophosphate (IMP) in muscle and hypoxanthine, xanthine and uric acid in plasma. Five Standardbred trotters performed treadmill exercise on 2 occasions, once at 7 m/s and once at 10 m/s. Venous blood samples were taken at rest, during exercise and at the end of exercise. Muscle biopsies were taken before and after exercise and muscle temperature was measured before and after exercise. Running time differed a...
Gansen S, Lindner A, Marx S, Mosen H, Sallmann HP.The effects of 3 different conditioning programmes on muscle glycogen concentration in horses were examined. Speed of exercise was selected according to the blood lactate values for each horse derived from a standardised exercise test before beginning a conditioning programme. Six 2-year-old Haflinger stallions were assigned randomly to one of 3 conditioning programmes according to a 6 x 3 latin square design: 45 min at their individual v1.5 or v2.5 and 25 min at v4. Each conditioning programme lasted 6 weeks (21 exercise sessions), followed by 5 weeks without conditioning (resting period). Al...
Hyyppä S, Saastamoinen M, Reeta Pösö A.Low muscle glycogen at the beginning of exercise may adversely affect performance, increase protein degradation and contribute to the onset of fatigue. As horses are sometimes required to compete on consecutive days both in racing and endurance types of competition, optimal muscle glycogen repletion may improve performance on the day following a race day. The purpose of this experiment was to study the effects of fat supplementation on repletion of muscle glycogen. Twelve Finnhorses performed an exercise test on a treadmill, and 2 and 4 h later they received hay and concentrate (Trial A). Two ...
Valberg SJ, Mickelson JR, Gallant EM, MacLeay JM, Lentz L, de la Corte F.The purpose of this study was to determine if chronic exertional rhabdomyolysis (ER) in Quarter Horses and Thoroughbreds represents one or several distinct myopathies. Eighteen Quarter Horses and 18 Thoroughbreds with ER were selected from cases presented to the Veterinary Hospital on the basis of a history of ER, assessment of muscle histopathology, and serum CK activity before and 4 h post exercise. In addition, 2 of 3 of the following parameters were evaluated: muscle glycogen concentrations, thyroid hormones (T3, T4), fractional excretion (FE) of sodium, potassium and chloride. The CK resp...
Schuback K, Essén-Gustavsson B, Persson SG.The aim was to study metabolic response and locomotion pattern in Standardbred trotters during incremental treadmill exercise performed by increasing speed by 1 m/s in 1 min steps (start 7 m/s) until the onset of fatigue. The test protocol included determination of oxygen uptake, heart rate (HR), stride length (SL) and stride frequency (SF). Venous blood samples were collected at rest, at the end of each exercise step and after 30 min of recovery. Muscle biopsies were taken at rest and post exercise and muscle temperature was measured after exercise. As horses fatigued at different speed steps...
Lacombe V, Hinchcliff KW, Geor RJ, Lauderdale MA.The purpose of this study was to develop a model of muscle glycogen depletion and to study the effect of this model on aerobic and anaerobic capacity of horses. The maximal rate of oxygen consumption (VO2max), maximal accumulated oxygen deficit (MAOD), muscle glycogen concentration and blood lactate concentration of 6 fit Standardbred horses were measured on 3 occasions 7 days apart (Trials 1, 2 and 3). Between Trials 2 and 3, strenuous exercise intended to deplete muscle glycogen was performed by exercising horses on the treadmill on 3 consecutive days. Strenuous exercise resulted in reductio...
Jensen-Waern M, Lindberg A, Johannisson A, Gröndahl G, Lindgren JA, Essén-Gustavsson B.The effects of an endurance ride on neutrophil functions in endurance-trained horses were evaluated and related to metabolic changes and changes in cortisol concentrations. Blood samples were taken from 7 horses (aged 9-15 years) one day before, and then 30-60 min, 1 day and 8 days after the ride. The race resulted in elevated serum cortisol levels (< 465 nmol/l) and an increased neutrophil:lymphocyte ratio. Immediately post race, the neutrophil ability to engulf yeast was increased. One day after the race, a decrease in leukotriene B4 production (approximately 40%) and in the respiratory b...
Valberg SJ, Townsend D, Mickelson JR.To determine whether polysaccharide storage myopathy (PSSM) in Quarter Horses is attributable to a defect in glycolysis or in the allosteric regulation of phosphofructokinase (PFK) enzyme. Methods: Muscle biopsy specimens were obtained from 6 Quarter Horses with PSSM and 8 Quarter Horse or Thoroughbred control horses. Methods: Maximal activity of glycogenolytic and glycolytic enzymes was determined spectrophotometrically. Maximal activity of PFK was determined for each horse at pH 8.0, and at pH 7.0 when variable concentrations of the activators, fructose 6 phosphate, fructose 2,6 bisphosphate...
Snow DH, Mackenzie G.The effects of intermittent maximal exercise (galloping) before and after a 10 week training programme were studied in 6 horses. Determinations were carried out on venous blood for packed cell volume, total plasma protein, glucose, glycerol, free fatty acids, lactate, 11-hydroxycorticosteroids, blood gases and pH. There were marked changes associated with galloping and some of these could be modified with training. The major findings included (i) an elevated blood glucose, (ii) a large increase in glycerol, which was greatest at 30 min post-exercise and was higher following training, (iii) sma...
McCue ME, Valberg SJ.To estimate the prevalence of polysaccharide storage myopathy (PSSM) among Quarter Horses in the United States and evaluate possible relationships between muscle glycogen concentration, turnout time, and exercise level. Methods: Cross-sectional study. Methods: 164 overtly healthy Quarter Horses > 2 years old from 5 states. Methods: Horses with a history of exertional rhabdomyolysis or any other muscular disease were excluded. Muscle biopsy specimens were examined histologically for evidence of PSSM and were submitted for determination of muscle glycogen concentration. A diagnosis of PSSM wa...
Geor RJ, Hinchcliff KW, Sams RA.To examine the role of beta-adrenergic mechanisms in the regulation of endogenous glucose (Glu) production [rate of appearance (R(a))] and utilization [rate of disappearance (R(d))] and carbohydrate (CHO) metabolism, six horses completed consecutive 30-min bouts of exercise at approximately 30% (Lo) and approximately 60% (Hi) of estimated maximum O(2) uptake with (P) and without (C) prior administration of the beta-blocker propranolol (0.22 mg/kg iv). All horses completed exercise in C; exercise duration in P was 49.9 +/- 1.2 (SE) min. Plasma Glu was unchanged in C during Lo but increased prog...
Geor RJ, Hinchcliff KW, McCutcheon LJ, Sams RA.This study examined the effects of preexercise glucose administration, with and without epinephrine infusion, on carbohydrate metabolism in horses during exercise. Six horses completed 60 min of treadmill exercise at 55 +/- 1% maximum O(2) uptake 1) 1 h after oral administration of glucose (2 g/kg; G trial); 2) 1 h after oral glucose and with an intravenous infusion of epinephrine (0.2 micromol. kg(-1). min(-1); GE trial) during exercise, and 3) 1 h after water only (F trial). Glucose administration (G and GE) caused hyperinsulinemia and hyperglycemia ( approximately 8 mM). In GE, plasma epine...
Schwarz B, Ertl R, Zimmer S, Netzmann Y, Klein D, Schwendenwein I, Hoven RV.The aim of this study was to determine the occurrence and frequency of a mutation in the gene coding for skeletal muscle glycogen synthase type 1 (GYS-1), which is the cause of equine polysaccharide storage myopathy (PSSM) type 1 in a population of 50 Haflingers. GYS-1 genotyping of 50 Haflingers was performed with a validated restriction fragment length polymorphism (RFLP) assay. The second aim was to compare resting and post-exercise muscle enzyme activities as well as parameters of glucose metabolism in blood between horses with and without the mutation. Nine of the 50 Haflingers were ident...
De La Corte FD, Valberg SJ, MacLeay JM, Williamson SE, Mickelson JR.To determine whether excessive glycogen accumulation in skeletal muscle of Quarter Horses with polysaccharide storage myopathy (PSSM) is a result of enhanced cellular uptake of glucose. Methods: 6 horses with PSSM and 10 healthy (control) horses. Methods: Intravenous glucose tolerance tests (IVGTT), oral glucose tolerance tests (OGTT), and modified insulin tolerance tests (MITT) were performed. Plasma glucose and insulin concentrations were measured in blood samples collected before and for up to 8 hours after glucose or insulin administration. Results: Peak glucose concentrations during IVGTT...
Herszberg B, McCue ME, Larcher T, Mata X, Vaiman A, Chaffaux S, Chérel Y, Valberg SJ, Mickelson JR, Guérin G.Glycogen storage diseases or glycogenoses are inherited diseases caused by abnormalities of enzymes that regulate the synthesis or degradation of glycogen. Deleterious mutations in many genes of the glyco(geno)lytic or the glycogenesis pathways can potentially cause a glycogenosis, and currently mutations in fourteen different genes are known to cause animal or human glycogenoses, resulting in myopathies and/or hepatic disorders. The genetic bases of two forms of glycogenosis are currently known in horses. A fatal neonatal polysystemic type IV glycogenosis, inherited recessively in affected Qu...
Lawrence LM.Substrate depletion and end product accumulation are two important factors in exercise fatigue. Fatigue during long-term exercise results from a depletion of muscle and liver glycogen and coincides with an inability to maintain blood glucose levels. During high intensity exercise, the rapid catabolism of carbohydrate and the resultant production of lactate and hydrogen ions cause a reduction in muscle pH that inhibits maximum force generation. Dietary manipulations that can influence carbohydrate status or lactate accumulation may be beneficial to performance. In human athletes, carbohydrate l...
Jose-Cunilleras E, Hinchcliff KW, Sams RA, Devor ST, Linderman JK.In a randomized, balanced, crossover study each of six fit, adult horses ran on a treadmill at 50% of maximal rate of oxygen consumption for 60 min after being denied access to food for 18 h and then 1) fed corn (51.4 kJ/kg digestible energy), or 2) fed an isocaloric amount of alfalfa 2-3 h before exercise, or 3) not fed before exercise. Feeding corn, compared with fasting, resulted in higher plasma glucose and serum insulin and lower serum nonesterified fatty acid concentrations before exercise (P < 0.05) and in lower plasma glucose, serum glycerol, and serum nonesterified fatty acid conce...
Essén-Gustavsson B, Jensen-Waern M.Glycogenolysis and lipolysis are essential for energy production in muscle during prolonged exercise but less is known about the role of amino acid metabolism. The aim was to study the effect of an endurance race on pro- and macroglycogen, triglycerides and amino acid concentrations in muscle and on selected blood parameters, especially the amino acid response in the blood during the recovery phase. Seven endurance-trained horses completed a race covering 100-120 km. Blood samples were collected one day before the race, after the finish and 1 and 8 days after the race. Needle biopsy specimens ...
Bröjer JT, Stämpfli HR, Graham TE.To determine proglycogen (PG) and macroglycogen (MG) content in equine skeletal muscle and to compare 2 analytical methods (acid hydrolysis [AC] and PG plus MG determination) for measurement of total muscle glycogen content (Gly(tot)) in biopsy specimens. Methods: Muscle biopsy specimens obtained from 41 clinically normal horses. Methods: Forty-five muscle biopsy specimens obtained from the middle gluteal (n = 31) or triceps (14) muscle were analyzed, using AC and MG plus PG determination for Gly(tot). Variability within muscle biopsy specimens for each method was calculated from duplicate ana...
Valberg S, Essén-Gustavsson B, Lindholm A, Persson S.The skeletal muscle metabolism of horses with a wide range of cardiocirculatory capacities was studied during a standardised near-maximal exercise test in relation to muscle fibre composition. Although the same amount of work was performed by all horses the amount of triglycerides and glycogen utilised and lactate accumulated differed widely. Both blood and muscle lactate accumulation were positively related to the amount of glycogen utilised. These factors were in turn positively associated with the percentage of Type IIB fibres and consequently negatively associated with the percentage of Ty...
Williams ZJ, Bertels M, Valberg SJ.Type 1 polysaccharide storage myopathy (PSSM1) is a glycogen storage disorder of known cause whereas the basis for type 2 PSSM (PSSM2) is unknown. The same diet and exercise regime prescribed for PSSM1 is recommended for PSSM2; however, the benefit of these recommendations for PSSM2 is undocumented. The objectives of this study were to determine traits of PSSM2 Warmblood horses (WB), determine the changes in exercise responses that occur with a recommended low-starch/fat-supplemented diet and exercise regime, and determine if glycogen concentrations correspond to the severity of signs. Owners ...
Essén-Gustavsson B, Gottlieb-Vedi M, Lindholm A.The aim was to investigate metabolic response in muscle during submaximal treadmill exercise to fatigue, with a special emphasis on adenine nucleotide degradation products such as inosine monophosphate (IMP) in muscle and hypoxanthine, xanthine and uric acid in plasma. Five Standardbred trotters performed treadmill exercise on 2 occasions, once at 7 m/s and once at 10 m/s. Venous blood samples were taken at rest, during exercise and at the end of exercise. Muscle biopsies were taken before and after exercise and muscle temperature was measured before and after exercise. Running time differed a...
Valentine BA, Habecker PL, Patterson JS, Njaa BL, Shapiro J, Holshuh HJ, Bildfell RJ, Bird KE.Skeletal muscle samples from 38 draft horse-related animals 1-23 years of age were evaluated for evidence of aggregates of glycogen and complex polysaccharide characteristic of equine polysaccharide storage myopathy (EPSSM). Cardiac muscle from 12 of these horses was also examined. Antemortem serum levels of creatine kinase (CK) and aspartate aminotransferase (AST) from 9 horses with EPSSM and 5 horses without EPSSM were compared. Skeletal muscle from 17 horses contained inclusions of periodic acid-Schiff (PAS)-positive, amylase-resistant complex polysaccharide. Similar inclusions were also pr...
Chen J, Gollnick PD.Horses were subjected to treadmill running at 65% (submaximal) or 100% (maximal) VO2,max to examine the effects of exercise on subcellular distribution of hexokinase (HK) and on mitochondrial respiration. It is hypothesized that the fraction of HK bound to mitochondria will be reduced due to an elevation of glucose-6-phosphate (G-6-P) concentration in the exercising muscle and that such release of HK from mitochondria will depress oxidative phosphorylation. Changes in muscle G-6-P concentration, pH, subcellular HK distribution, mitochondrial respiration and other metabolites were determined in...
Gansen S, Lindner A, Marx S, Mosen H, Sallmann HP.The effects of 3 different conditioning programmes on muscle glycogen concentration in horses were examined. Speed of exercise was selected according to the blood lactate values for each horse derived from a standardised exercise test before beginning a conditioning programme. Six 2-year-old Haflinger stallions were assigned randomly to one of 3 conditioning programmes according to a 6 x 3 latin square design: 45 min at their individual v1.5 or v2.5 and 25 min at v4. Each conditioning programme lasted 6 weeks (21 exercise sessions), followed by 5 weeks without conditioning (resting period). Al...
Freeman KP, Roszel JF, Slusher SH, Castro M.Histochemical stains were applied to six equine uterine biopsies representative of the physiologic breeding season, Spring and Fall transition, and Winter anestrus periods. These were compared with uterine biopsies from six mares with intrauterine urine pooling, eight mares used to study the uterine response to indwelling catheterization, and necropsy specimens from four pregnant mares at approximately 60 or 100 d of gestation. Alcian blue staining at pH 2.5 or 1.0 was used to identify the presence of carboxylated and sulfated acid mucins or only suflated acid mucins, respectively. Periodic ac...
Bröjer JT, Nostell KE, Essén-Gustavsson B, Hedenström UO.To determine whether repeated oral administration of glucose and leucine during the period immediately after intense exercise would increase the release of insulin and thereby enhance glycogen synthesis in horses. Methods: 12 Standardbred horses. Methods: In a crossover study design, after glycogen-depleting exercise, horses received oral boluses of glucose (1 g/kg at 0, 2, and 4 hours) and leucine (0.1 g/kg at 0 and 4 hours) or boluses of water (10 mL/kg at 0, 2, and 4 hours; control treatment). Blood samples for determination of glucose, insulin, and leucine concentrations were collected pri...
Bergh A, Nordlöf H, Essén-Gustavsson B.Neuromuscular electrical stimulation (NMES) is used to increase or maintain muscle strength during rehabilitation. Human studies investigating different protocols show that some treatments induce changes in muscle characteristics. Despite the frequent use of NMES in horses, no studies have been published describing its efficacy. Objective: To investigate the effects of a NMES protocol on equine fibre types and areas, glycogen concentrations and enzyme activities. Methods: NMES was administrated to m. gluteus medius and m. longissimus dorsi, on one side of 6 healthy Standardbred horses. The con...
Nam GH, Ahn K, Bae JH, Han K, Lee CE, Park KD, Lee SH, Cho BW, Kim HS.Muscle glycogen Phosphorylase (PYGM) has been shown to catalyze the degradation of glycogen to glucose-1-phosphate. The PYGM gene can contribute to providing energy to the body by disassembling the glycogen in muscle. Here, we analyzed the genomic structure and expression of the PYGM gene in the thoroughbred horse. The PYGM gene, containing several transposable elements (MIRs, LINEs, and MERs), was highly conserved in mammalian genomes. In order to understand the expression of the horse PYGM gene, we performed quantitative RT-PCR using 11 thoroughbred horse tissue samples. The horse PYGM gene ...
Valentine BA, Credille KM, Lavoie JP, Fatone S, Guard C, Cummings JF, Cooper BJ.A severe myopathy leading to death or euthanasia was identified in 4 Belgian and 4 Percheron draught horses age 2-21 years. Clinical signs ranged from overt weakness and muscle atrophy in 2 horses age 2 and 3 years, to recumbency with inability to rise in 6 horses age 4-21 years. In 5 horses there was mild to severe increases in muscle enzyme levels. Clinical diagnoses included equine motor neuron disease (2 horses), post anaesthetic myopathy (2 horses), exertional myopathy (2 horses), myopathy due to unknown (one horse), and equine protozoal myelitis (one horse). Characteristic histopathology...
Nimmo MA, Snow DH, Munro CD.The effect of 11 weekly injections of nandrolone phenylpropionate (400 mg) on some skeletal muscle parameters was investigated in 6 Thoroughbred geldings undergoing training. Three muscles were sampled, the middle gluteal, the biceps femoris and the semitendinosus. Training alone produced increases in the percentage of fast twitch high oxidative fibres (FTH), glycogen content and the activities of citrate synthase, 3-hydroxyacl CoA dehydrogenase and cytochrome oxidase. In contrast the training programme did not alter water content, total protein content, the activities of lactate dehydrogenase...
Naylor RJ, Livesey L, Schumacher J, Henke N, Massey C, Brock KV, Fernandez-Fuente M, Piercy RJ.Equine type 1 polysaccharide storage myopathy (PSSM1), a common glycogenosis associated with an R309H founder mutation in the glycogen synthase 1 gene (GYS1), shares pathological features with several human myopathies. In common with related human disorders, the pathogenesis remains unclear in particular, the marked phenotypic variability between affected animals. Given that affected animals accumulate glycogen and alpha-crystalline polysaccharide within their muscles, it is possible that physical disruption associated with the presence of this material could exacerbate the phenotype. The aim ...
Valberg S.The glycogen depletion patterns in the gluteus muscle of Standardbred horses were studied under different trotting intensities. After racing significant glycogen depletion was found in all Type I and IIA fibres and in a varying percentage of Type IIB fibres, depending on the individual horse. When horses performed exercise over a short distance (5 to 8 km) at three different speeds, glycogen depletion was difficult to detect for the faster speeds (10 and 8 m/sec) but notably involved a major proportion of Type I fibres at the slowest speed (6 m/sec). When exercise was prolonged over a greater ...
Echigoya Y, Okabe H, Itou T, Endo H, Sakai T.Muscle glycogen synthase (GYS1) is the rate-limiting enzyme in glycogen synthesis, and its activity is regulated by the phosphorylation states of certain amino acid residues encoded by the GYS1 gene. In the present study, the authors molecularly characterized the full-length equine GYS1 (eGYS1) cDNA and found that it contains a less common polyadenylation signal (AATACA). An amino acid alignment with other mammalian GYS1 showed that the phosphorylation sites in eGYS1 are completely conserved. Genomic DNA analysis revealed that the equine-specific substitutions (Glu 16 Asp and Ala 252 Thr) were...
Lacombe V, Hinchcliff KW, Geor RJ, Lauderdale MA.The purpose of this study was to develop a model of muscle glycogen depletion and to study the effect of this model on aerobic and anaerobic capacity of horses. The maximal rate of oxygen consumption (VO2max), maximal accumulated oxygen deficit (MAOD), muscle glycogen concentration and blood lactate concentration of 6 fit Standardbred horses were measured on 3 occasions 7 days apart (Trials 1, 2 and 3). Between Trials 2 and 3, strenuous exercise intended to deplete muscle glycogen was performed by exercising horses on the treadmill on 3 consecutive days. Strenuous exercise resulted in reductio...
Jansson A, Nyman S, Lindholm A, Lindberg JE.Studies of the effect of varying dietary carbohydrate compositions on exercise metabolism of horses are scarce. In the present study, the starch and sugar proportions were altered in the diet to 4 Standardbred horses. In a crossover experiment, the horses were offered a hay and oat diet, where the oats were substituted for barley syrup (BS) at 4 levels (BS 0, 0.5, 1.0 and 1.5 kg). Each diet was consumed for 21 days and both an incremental treadmill exercise test (IE) and a 40 min submaximal exercise test (SE) were performed. During the IE there was a significant (P<0.05) increase in mean VO...
