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.
Jansson A, Lindberg JE.Most athletic horses are fed a high-starch diet despite the risk of health problems. Replacing starch concentrate with high-energy forage would alleviate these health problems, but could result in a shift in major substrates for muscle energy supply from glucose to short-chain fatty acids (SCFA) due to more hindgut fermentation of fibre. Dietary fat inclusion has previously been shown to promote aerobic energy supply during exercise, but the contribution of SCFA to exercise metabolism has received little attention. This study compared metabolic response with exercise and lactate threshold (VLa...
Nostell KE, Essén-Gustavsson B, Bröjer JT.The branched chain amino acid leucine is a potent stimulator of insulin secretion. Used in combination with glucose it can increase the insulin response and the post exercise re-synthesis of glycogen in man. Decreased plasma amino acid concentrations have been reported after intravenous or per oral administration of leucine in man as well as after a single per oral dose in horses. In man, a negative correlation between the insulin response and the concentrations of isoleucine, valine and methionine have been shown but results from horses are lacking. This study aims to determine the effect of ...
Suagee JK, Corl BA, Swyers KL, Smith TL, Flinn CD, Geor RJ.High glycaemic feeds are associated with the development of insulin resistance in horses. However, studies that evaluated the effect of high glycaemic feeds used horses that either ranged in body condition from lean to obese or were fed to increase body condition over a period of months; thus, the ability of high glycaemic feeds to induce insulin resistance in lean horses has not been determined. This study evaluated the insulin sensitivity of 18 lean horses fed a 10% (LO; n = 6), 20% (MED; n = 6) or 60% (HI; n = 6) non-structural carbohydrate complementary feed for 90 days. Alth...
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...
Essén-Gustavsson B, Connysson M, Jansson A.There is little information about the influence of crude protein (CP) intake on glycogen and free pool amino acid concentrations in the muscle of horses in training. High energy forage-only diets may be an alternative to concentrate rich diets and may provide high levels of CP. Objective: To study the effect of feeding 2 forage-only diets, containing either high or moderate CP concentrations on glycogen and free pool amino acid concentrations in the muscle. Methods: Two high energy forage-only diets based on high-energy grass forage were fed for 23 days in a crossover design to 6 Standardbred ...
Stewart-Hunt L, Pratt-Phillips S, McCutcheon LJ, Geor RJ.Starch rich (S) feeds reduce insulin sensitivity in untrained horses when compared to high fat (F) feeds, but insulin sensitivity is not affected when S or F are fed during exercise training. The effects of S vs. F on training-associated alterations in skeletal muscle glucose metabolism are unknown. Objective: To determine the effects of dietary energy source on training-associated changes in insulin sensitivity, skeletal muscle GLUT4 protein and hexokinase (HK) and glycogen synthase (GS) activities in horses. Methods: After a baseline period on an all forage diet (Phase 1), horses were adapte...
Borgia LA, Valberg SJ, Essen-Gustavsson B.Flexor tendon injury may be due to flexor muscle fatigue, contributing to fetlock joint hyperextension and tendon damage. A water treadmill provides resistance training on flexor tendon muscles, which might reduce the risk of tendon injury. Objective: To determine the effect of water treadmill training on the properties of the gluteal and superficial digital flexor (SDF) muscles and on cardiocirculatory response to a standardised exercise test. Methods: Five healthy unfit horses were trained on a water treadmill for 5 days/week for 4 weeks, starting with 5 min/day increasing to 20 min/day. Bef...
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...
Urschel KL, Geor RJ, Waterfall HL, Shoveller AK, McCutcheon LJ.Providing protein or amino acid mixtures in combination with glucose to post exercise in man has resulted in increases in the post feeding insulin response and in muscle glycogen and protein synthesis rates. However, whether protein and/or amino acids can modify the post exercise insulin responses in horses remains to be fully elucidated. Objective: To determine whether whey protein or leucine addition to a glucose solution affects the post gavage plasma insulin, glucose and amino acid responses in horses and whether these responses are different following a period of exercise vs. rest. Method...
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 ...
Yamano S, Kawai M, Minami Y, Hiraga A, Miyata H.We evaluated differences in muscle fiber recruitment patterns between continuous and interval training to develop an optimal training program for Thoroughbred horses. Five well trained female thoroughbred horses (3-4 years old) were used. The horses performed two different exercises on a 10% inclined treadmill: 90%VO2 max for 4 min (continuous) and 90% VO2 max for 2 min × 2 times with 10-min interval (interval). Muscle samples were obtained from the middle gluteal muscle before and immediately after the exercises. Four muscle fiber types (type I, IIA, IIA/X, and IIX) were immunohistochemicall...
Borgia L, Valberg S, McCue M, Watts K, Pagan J.The aim of this study was to determine whether the glycaemic/insulinaemic responses to hay with non-structural carbohydrate (NSC, soluble carbohydrate) of 17% (HC), 10% (MC) or 4% (LC) differs in control horses and whether these responses differ between control and horses with polysaccharide storage myopathy (PSSM). Five clinically normal control horses and seven PSSM horses, all unfit and of Quarter Horse breeding (age 9.4 ± 3.4 years, body condition score range: 4.5-6). A crossover design compared the HC and LC hay, with horses randomly assigned to hay type for 5 days, and all horses fed th...
Isgren CM, Upjohn MM, Fernandez-Fuente M, Massey C, Pollott G, Verheyen KL, Piercy RJ.Exertional rhabdomyolysis syndrome is recognised in many athletic horse breeds and in recent years specific forms of the syndrome have been identified. However, although Standardbred horses are used worldwide for racing, there is a paucity of information about the epidemiological and performance-related aspects of the syndrome in this breed. The objectives of this study therefore were to determine the incidence, risk factors and performance effects of exertional rhabdomyolysis syndrome in Standardbred trotters and to compare the epidemiology and genetics of the syndrome with that in other bree...
Waller AP, Lindinger MI.Carbohydrate (CHO) stored in the form of skeletal muscle glycogen is the main energy source for glycolytic and oxidative ATP production during vigorous exercise in mammals. In man, horse and dog both short-term high intensity and prolonged submaximal exercise deplete muscle glycogen. In horses, however, muscle glycogen synthesis is 2-3-fold slower than in man and rat, even when a diet high in soluble CHO is fed. There appear to be significant differences in CHO and glycogen metabolism between horses and other mammals, and it is becoming increasingly clear that many conclusions drawn from human...
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...
Minami Y, Yamano S, Kawai M, Hiraga A, Miyata H.To find a new parameter indicating muscle fitness in Thoroughbred horses, we examined time-dependent recovery of glycogen content and sarcoplasmic reticulum (SR) Ca(2+)-ATPase activity of skeletal muscle after intensive treadmill running. Two repeated 50-sec running sessions (13 m/sec) were performed on a flat treadmill (approximately 90%VO2max). Muscle samples of the middle gluteal muscle were taken before exercise (pre) and 1 min, 20 min, 60 min, and 24 hr after exercise. Muscle fiber type composition was determined in the pre muscle samples by immunohistochemical staining with monoclonal an...
Barrey E, Mucher E, Jeansoule N, Larcher T, Guigand L, Herszberg B, Chaffaux S, Guérin G, Mata X, Benech P, Canale M, Alibert O, Maltere P, Gidrol X.Several cases of myopathies have been observed in the horse Norman Cob breed. Muscle histology examinations revealed that some families suffer from a polysaccharide storage myopathy (PSSM). It is assumed that a gene expression signature related to PSSM should be observed at the transcriptional level because the glycogen storage disease could also be linked to other dysfunctions in gene regulation. Thus, the functional genomic approach could be conducted in order to provide new knowledge about the metabolic disorders related to PSSM. We propose exploring the PSSM muscle fiber metabolic disorder...
McCue ME, Valberg SJ, Pagan JD, Essén-Gustavsson B, Roe CR.To compare effects of corn oil or a 7-carbon fat (triheptanoin) on acylcarnitine, lipid, and carbohydrate metabolism in plasma or muscle of exercising horses. Methods: 8 Thoroughbred geldings. Methods: Horses received isocaloric diets containing 650 mL of oil (triheptanoin or corn oil)/d for 18 or 25 days in a crossover design with a 26-day washout period. On day 17 or 24 of each feeding period, the respective oil (217 mL) was nasogastrically administered; 120 minutes later, horses performed a 90-minute submaximal exercise test (SET). Blood and muscle samples were obtained before oil administr...
McCue ME, Armién AG, Lucio M, Mickelson JR, Valberg SJ.Polysaccharide storage myopathy (PSSM) has been found in more than 35 different horse breeds through identification of abnormal storage of polysaccharide in muscle biopsies. A dominant mutation in the glycogen synthase 1 gene (GYS1) accounts for a substantial proportion of PSSM cases in at least 17 breeds, including Quarter Horses, but some horses diagnosed with PSSM by muscle histopathologic analysis are negative for the mutation. We hypothesized that a second distinct form of glycogen storage disease exists in GYS1-negative horses with PSSM. The objectives of this study were to compare the h...
Waller AP, Geor RJ, Spriet LL, Heigenhauser GJ, Lindinger MI.Oral acetate supplementation enhances glycogen synthesis in some mammals. However, while acetate is a significant energy source for skeletal muscle at rest in horses, its effects on glycogen resynthesis are unknown. We hypothesized that administration of an oral sodium acetate-acetic acid solution with a typical grain and hay meal after glycogen-depleting exercise would result in a rapid appearance of acetate in blood with rapid uptake by skeletal muscle. It was further hypothesized that acetate taken up by muscle would be converted to acetyl CoA (and acetylcarnitine), which would be metaboliz...
McCue ME, Valberg SJ, Jackson M, Borgia L, Lucio M, Mickelson JR.In this study we examined a family of Quarter Horses with Polysaccharide Storage Myopathy (PSSM) with a dominant mutation in the skeletal muscle glycogen synthase (GYS1) gene. A subset of horses within this family had a more severe and occasionally fatal PSSM phenotype. The purpose of this study was to identify a modifying gene(s) for the severe clinical phenotype. A genetic association analysis was used to identify RYR1 as a candidate modifying gene. A rare, known equine RYR1 mutation, associated with malignant hyperthermia (MH), was found to segregate in this GYS1 PSSM family. Retrospective ...
Waller AP, Heigenhauser GJ, Geor RJ, Spriet LL, Lindinger MI.We hypothesized that postexercise rehydration using a hypotonic electrolyte solution will increase the rate of recovery of whole body hydration, and that this is associated with increased muscle glycogen and electrolyte recovery in horses. Gluteus medius biopsies and jugular venous blood were sampled from six exercise-conditioned Standardbreds on two separate occasions, at rest and for 24 h following a competitive exercise test (CET) designed to simulate the speed and endurance test of a 3-day event. After the CETs, horses were given water ad libitum, and either a hypotonic commercial electrol...
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...
Hoffmann C, Bazer FW, Klug J, Aupperle H, Ellenberger C, Schoon HA.Although alterations in patterns of protein secretion revealed in uterine flushings from mares suffering from endometrosis have been described, little is known about alterations at the cellular level. Hence, the aim of this study was to characterize deviations in patterns of uterine gland secretion patterns using endometrial biopsies, histochemical and newly established immunohistochemical methods. Forty-eight endometrial biopsies were obtained from mares suffering from various types of endometrosis (active and inactive, destructive and non-destructive) and degree (mild to severe) were analyze...
Firshman AM, Valberg SJ, Baird JD, Hunt L, DiMauro S.To determine insulin sensitivity, proportions of muscle fiber types, and activities of glycogenolytic and glycolytic enzymes in Belgians with and without polysaccharide storage myopathy (PSSM). Methods: 10 Quarter Horses (QHs) and 103 Belgians in which PSSM status had been determined. Methods: To determine insulin sensitivity, a hyperinsulinemic euglycemic clamp (HEC) technique was used in 5 Belgians with PSSM and 5 Belgians without PSSM. Insulin was infused i.v. at 3 mU/min/kg for 3 hours, and concentrations of blood glucose and plasma insulin were determined throughout. An i.v. infusion of g...
Gehlen H, Oey L, Rohn K, Bilzer T, Stadler P.Chronic pulmonary diseases (recurrent airway obstruction [RAO]) have been reported to alter skeletal muscle cells in humans. The purpose of this study was to evaluate a potential relationship between pulmonary and muscle variables in horses with a clinical diagnosis of RAO. Muscle biopsies from healthy horses and from horses with RAO were investigated and the relationship between the severity of lung disease and the degree of muscular changes was determined. Objective: We hypothesized that chronic pulmonary disease can lead to changes of the skeletal muscle in horses. Methods: Fifteen healthy ...
Aleman M.Muscle disorders are a common cause of disability in horses. For many years, clinical manifestations such as muscle pain, exercise intolerance, weakness, and stiffness were believed to be caused by a single syndrome. However, in the past years a broad spectrum of muscle disorders have been recognized including glycogen and polysaccharide storage myopathies, malignant hyperthermia, mitochondrial myopathy, hyperkalemic periodic paralysis and others. For some, a specific mutation has been identified. Recognition of the myopathic clinical phenotype and thorough clinical, electrodiagnostic, and his...
McCue ME, Valberg SJ, Miller MB, Wade C, DiMauro S, Akman HO, Mickelson JR.Polysaccharide storage myopathy (PSSM) is a novel glycogenosis in horses characterized by abnormal glycogen accumulation in skeletal muscle and muscle damage with exertion. It is unlike glycogen storage diseases resulting from known defects in glycogenolysis, glycolysis, and glycogen synthesis that have been described in humans and domestic animals. A genome-wide association identified GYS1, encoding skeletal muscle glycogen synthase (GS), as a candidate gene for PSSM. DNA sequence analysis revealed a mutation resulting in an arginine-to-histidine substitution in a highly conserved region of G...
Tiley HA, Geor RJ, McCutcheon LJ.To determine the effects of dexamethasone treatment on selected components of insulin signaling and glucose metabolism in skeletal muscle obtained from horses before and after administration of a euglycemic-hyperinsulinemic clamp (EHC). Methods: 6 adult Standardbreds. Methods: In a balanced crossover study, horses received either dexamethasone (0.08 mg/kg, IV, q 48 h) or an equivalent volume of saline (0.9% NaCl) solution, IV, for 21 days. A 2-hour EHC was administered for measurement of insulin sensitivity 1 day after treatment. Muscle biopsy specimens obtained before and after the EHC were a...
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...
Valberg SJ, Macleay JM, Billstrom JA, Hower-Moritz MA, Mickelson JR.Polysaccharide storage myopathy (PSSM) is a distinct cause of exertional rhabdomyolysis in Quarter Horses that results in glycogen and abnormal polysaccharide accumulation. The purpose of this study was to determine if excessive glycogen storage in PSSM is due to a glycolytic defect that impairs utilisation of this substrate during exercise. Muscle biopsies, blood lactates and serum CK were obtained 1) at rest from 5 PSSM Quarter Horses, 4 normal Quarter Horses (QH controls) and 6 Thoroughbreds with recurrent exertional rhabdomyolysis (TB RER) and 2) after a maximal treadmill exercise test in ...
Borgia LA, Valberg SJ, Essen-Gustavsson B.Flexor tendon injury may be due to flexor muscle fatigue, contributing to fetlock joint hyperextension and tendon damage. A water treadmill provides resistance training on flexor tendon muscles, which might reduce the risk of tendon injury. Objective: To determine the effect of water treadmill training on the properties of the gluteal and superficial digital flexor (SDF) muscles and on cardiocirculatory response to a standardised exercise test. Methods: Five healthy unfit horses were trained on a water treadmill for 5 days/week for 4 weeks, starting with 5 min/day increasing to 20 min/day. Bef...
Valberg SJ, McKenzie EC, Eyrich LV, Shivers J, Barnes NE, Finno CJ.Although exertional rhabdomyolysis (ER) is common in Arabian horses, there are no dedicated studies describing histopathological characteristics of muscle from Arabian horses with ER. Objective: To prospectively identify distinctive histopathological features of muscle from Arabian endurance horses with a history of ER (pro-ER) and to retrospectively determine their prevalence in archived samples from Arabian horses with exertional myopathies (retro-ER). Methods: Prospective and retrospective histopathological description. Methods: Middle gluteal muscle biopsies obtained from Arabian controls ...
Ringmark S, Roepstorff L, Essén-Gustavsson B, Revold T, Lindholm A, Hedenström U, Rundgren M, Ogren G, Jansson A.The aim of this study was to, from a holistic perspective, describe the effects of a forage-only feeding system and a conventional training program on young Standardbred horses and compare data with similar observations from the literature. Sixteen Standardbred colts fed a forage-only diet for 4 months from breaking (August to December) and with the goal to vigorously trot 5 to 7 km at a speed of 5.6 m/s (3 min/km) were studied. The horses were fed grass haylage (56 to 61% dry matter (DM), 2.80 to 3.02 Mcal DE/kg DM and 130 to 152 g CP/kg DM) ad libitum, 1 kg of a lucerne product and minerals....
Chanda M, Srikuea R, Cherdchutam W, Chairoungdua A, Piyachaturawat P.The match play patterns in equestrian polo are unique and require specific training programs to ensure sport performance. The effect of commonly used exercise training regimens on the adaptation of skeletal muscle is unclear. The present study investigated the modulating effects of the classic training regimen, comprised of aerobic exercise training with increasing exercise intensities and varying duration combined with match play, on the properties of muscle in polo ponies. Nine healthy adult female polo ponies were subjected to four consecutive subsets of 1 year classic training regimen incl...
Annandale EJ, Valberg SJ, Mickelson JR, Seaquist ER.Equine polysaccharide storage myopathy (PSSM) is an inherited disorder characterized by the accumulation of glycogen and abnormal polysaccharide in muscle with normal glyco(geno)lytic enzyme activities. The purpose of this study was to evaluate in vivo insulin sensitivity and glucose excursion in PSSM using a euglycemic hyperinsulinemic clamp. In addition, the content of muscle glucose transporters (GLUT1 and GLUT4) and the insulin receptor was determined in muscle biopsies using Western blot analysis. The glycogen content was 1.8-fold higher, and isolated polysaccharide analyzed by iodine abs...
McCue ME, Ribeiro WP, Valberg SJ.Controversy exists as to the prevalence of polysaccharide storage myopathy (PSSM) in breeds of horses and its impact on performance. Objective: To determine 1) the prevalence of PSSM in horses that presented with a neuromuscular disorder, as well as breed, sex and age distributions and clinical signs 2) effect of diagnostic criteria on prevalence, breed distribution and age of horses diagnosed with PSSM. Methods: Fresh frozen biopsies (n = 1426) submitted to the Neuromuscular Diagnostic Laboratory at the University of Minnesota were searched to identify horses diagnosed with PSSM. Horses with ...
Gottlieb M, Essén-Gustavsson B, Lindholm A, Persson SG.Five Standardbred trotters were trained on a treadmill 3 times/wk for 12 wk by intervals of draft-loaded exercise. The draft load was 34 kp and the velocity approximately 7 m/s. Muscle biopsies were taken from the gluteus medius and longissimus muscles before training and after 2, 4, 8, and 12 wk of training and from the brachiocephalicus muscle before and after training. Both the percentage and the area of type IIa fibers increased and the percentage of type IIb fibers decreased in the gluteus medius muscle during the first 2 wk of training, and then no further significant difference was note...
Lindholm A, Saltin B.LINDHOLM, ARNE and BENGT SALTIN: The physiological and biochemical response of standardbred horses to exercise of varying speed and duration. Acta vet. scand. 1974, 15, 310–324. — Welltrained standardbred horses were studied to examine the metabolic response to excercise of various speeds and duration. Comparisons between interval (400, 700, 1,000 and 2,000 m) and continuous trotting (1 hr., 2 hrs.) and racing were made. Muscle and rectal temperatures were recorded before and immediately after each work bout. Heart rate was linearly related to trotting speed, and maximal heart rate (240 be...
Valentine BA, Cooper BJ.Muscle samples were obtained at necropsy from 225 horses and ponies 1 year of age or older. Samples were processed in routine manner and were stained with hematoxylin and eosin and with periodic acid-Schiff for glycogen. Sections were examined for abnormal glycogen content and amylase-resistant complex polysaccharide and for chronic myopathic change (excessive fiber size variation, increase in number of internal nuclei). A total of 101 horses and ponies with lesions of polysaccharide storage myopathy were identified. Age of affected horses ranged from one to 30 years, with a mean of 14.7 years...
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 ...
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...
Pinzon-Arteaga C, Snyder MD, Lazzarotto CR, Moreno NF, Juras R, Raudsepp T, Golding MC, Varner DD, Long CR.Phenotypic selection during animal domestication has resulted in unwanted incorporation of deleterious mutations. In horses, the autosomal recessive condition known as Glycogen Branching Enzyme Deficiency (GBED) is the result of one of these deleterious mutations (102C > A), in the first exon of the GBE1 gene (GBE1). With recent advances in genome editing, this type of genetic mutation can be precisely repaired. In this study, we used the RNA-guided nuclease CRISPR-Cas9 (clustered regularly-interspaced short palindromic repeats/CRISPR-associated protein 9) to correct the GBE1 mutation in a...
Ward TL, Valberg SJ, Lear TL, Guérin G, Milenkovic D, Swinburne JE, Binns MM, Raudsepp T, Skow L, Chowdhary BP, Mickelson JR.Comparative biochemical and histopathological data suggest that a deficiency in the glycogen branching enzyme (GBE) is responsible for a fatal neonatal disease in Quarter Horse foals that closely resembles human glycogen storage disease type IV (GSD IV). Identification of DNA markers closely linked to the equine GBE1 gene would assist us in determining whether a mutation in this gene leads to the GSD IV-like condition. FISH using BAC clones as probes assigned the equine GBE1 gene to a marker deficient region of ECA26q12-->q13. Four other genes, ROBO2, ROBO1, POU1F1, and HTR1F, that flank GB...
Stewart-Hunt L, Pratt-Phillips S, McCutcheon LJ, Geor RJ.Starch rich (S) feeds reduce insulin sensitivity in untrained horses when compared to high fat (F) feeds, but insulin sensitivity is not affected when S or F are fed during exercise training. The effects of S vs. F on training-associated alterations in skeletal muscle glucose metabolism are unknown. Objective: To determine the effects of dietary energy source on training-associated changes in insulin sensitivity, skeletal muscle GLUT4 protein and hexokinase (HK) and glycogen synthase (GS) activities in horses. Methods: After a baseline period on an all forage diet (Phase 1), horses were adapte...
Yamano S, Kawai M, Minami Y, Hiraga A, Miyata H.We evaluated differences in muscle fiber recruitment patterns between continuous and interval training to develop an optimal training program for Thoroughbred horses. Five well trained female thoroughbred horses (3-4 years old) were used. The horses performed two different exercises on a 10% inclined treadmill: 90%VO2 max for 4 min (continuous) and 90% VO2 max for 2 min × 2 times with 10-min interval (interval). Muscle samples were obtained from the middle gluteal muscle before and immediately after the exercises. Four muscle fiber types (type I, IIA, IIA/X, and IIX) were immunohistochemicall...
Rose RJ, Sampson D.Total and individual non-esterified fatty acids (NEFA), pyruvate, lactate, alpha ketoglutarate, acetoacetate, beta hydroxybutyrate, glucose and insulin were measured in horses during an 80 km endurance ride and during four days of food deprivation. In the latter group venous blood-gas and acid-base parameters, and plasma cortisol concentrations were also measured. During exercise and food deprivation the NEFA became considerably elevated, the predominant of which were oleic (35 per cent), palmitic (24 per cent), linoleic (19 per cent) and linolenic (10 per cent). By one hour after feeding tota...
Ropka-Molik K, Stefaniuk-Szmukier M, Musiał AD, Piórkowska K, Szmatoła T.The ACTN3 gene codes for α-actinin-3, a protein localized in the Z-line in the skeletal muscle. Actinin-3 is critical in anchoring the myofibrillar actin filaments and plays a key role in muscle contraction. ACTN3 (α-actinin-3) cross-links glycogen phosphorylase (GP), which is the key enzyme catalysing glycogen metabolism. The aim of present study was to establish the expression level of the ACTN3 gene (for both isoforms separately and together in the gene expression analysis) in the gluteus medius muscle in order to verify if the α-actinin-3 gene can be related to training intensity in Ara...
Snow DH, Baxter P, Rose RJ.An investigation into fibre composition and glycogen depletion pattern within the middle gluteal of 16 horses participating in an 80 km endurance ride was carried out. Although the proportion of slow twitch high oxidative (ST) fibres in the horses varied between 7 and 38 per cent, it was found that the horses with the highest proportion of these fibres usually had the best performance records. The cross-sectional area of the fast twitch low oxidative (FT) fibres was greatest, with the ST and fast twitch high oxidative (FTH) being similar in size. Most marked histological evidence of glycogen d...
Urschel KL, Geor RJ, Waterfall HL, Shoveller AK, McCutcheon LJ.Providing protein or amino acid mixtures in combination with glucose to post exercise in man has resulted in increases in the post feeding insulin response and in muscle glycogen and protein synthesis rates. However, whether protein and/or amino acids can modify the post exercise insulin responses in horses remains to be fully elucidated. Objective: To determine whether whey protein or leucine addition to a glucose solution affects the post gavage plasma insulin, glucose and amino acid responses in horses and whether these responses are different following a period of exercise vs. rest. Method...
McCue ME, Valberg SJ, Jackson M, Borgia L, Lucio M, Mickelson JR.In this study we examined a family of Quarter Horses with Polysaccharide Storage Myopathy (PSSM) with a dominant mutation in the skeletal muscle glycogen synthase (GYS1) gene. A subset of horses within this family had a more severe and occasionally fatal PSSM phenotype. The purpose of this study was to identify a modifying gene(s) for the severe clinical phenotype. A genetic association analysis was used to identify RYR1 as a candidate modifying gene. A rare, known equine RYR1 mutation, associated with malignant hyperthermia (MH), was found to segregate in this GYS1 PSSM family. Retrospective ...
Lewis SS, Nicholson AM, Williams ZJ, Valberg SJ.OBJECTIVE To characterize clinical findings for polysaccharide storage myopathy (PSSM) in warmblood horses with type 1 PSSM (PSSM1; caused by mutation of the glycogen synthase 1 gene) and type 2 PSSM (PSSM2; unknown etiology). SAMPLE Database with 3,615 clinical muscle biopsy submissions. PROCEDURES Reported clinical signs and serum creatine kinase (CK) and aspartate aminotransferase (AST) activities were retrospectively analyzed for horses with PSSM1 (16 warmblood and 430 nonwarmblood), horses with PSSM2 (188 warmblood and 646 nonwarmblood), and warmblood horses without PSSM (278). Lameness e...
The Journal of heredityNovember 8, 2013
Volume 105, Issue 2 163-172 doi: 10.1093/jhered/est075
McCoy AM, Schaefer R, Petersen JL, Morrell PL, Slamka MA, Mickelson JR, Valberg SJ, McCue ME.A dominantly inherited gain-of-function mutation in the glycogen synthase (GYS1) gene, resulting in excess skeletal muscle glycogen, has been identified in more than 30 horse breeds. This mutation is associated with the disease Equine Polysaccharide Storage Myopathy Type 1, yet persists at high frequency in some breeds. Under historical conditions of daily work and limited feed, excess muscle glycogen may have been advantageous, driving the increase in frequency of this allele. Fine-scale DNA sequencing in 80 horses and genotype assays in 279 horses revealed a paucity of haplotypes carrying th...
Fowden AL, Mundy L, Ousey JC, McGladdery A, Silver M.Glycogen and glucose 6-phosphatase levels were measured in a variety of tissues from fetal, newborn and adult pregnant mares. Hepatic and renal glucose 6-phosphatase activities increased with increasing gestational age and rose further in the 24 h after birth to values similar to those observed in maternal tissues. The glycogen content of fetal liver and skeletal muscle also increased in late gestation, whereas that of fetal lung and cardiac muscle decreased toward term. No significant change in renal glycogen content with fetal age was observed. The values of hepatic glycogen and glucose 6-ph...
Lindholm A, Johansson HE, Kjaersgaard P.LINDHOLM, A., H.-E. JOHANSSON & P. KJÆRSGAARD: Acute rhabdomyolysis (“tying-up”) in standardbred horses. A morphological and biochemical study. Acta vet. scand. 1974, 15, 325–339. — Morphological, biochemical and histochemical changes were studied in muscle needle biopsy specimens (gluteus medius) from 59 standardbred trotters with acute clinical symptoms of the “tying-up” disease. All horses had increased levels of serum enzymes SGOT and SCPK. The biopsy specimens were taken at various intervals after onset of clinical symptoms (1–4 hrs., 18–24 hrs. and 2–20 days). Ry light...
Valberg SJ, Velez-Irizarry D, Williams ZJ, Pagan JD, Mesquita V, Waldridge B, Maresca-Fichter H.Horses have a slow rate of muscle glycogen repletion relative to other species for unknown reasons. Our aim was to determine the expression of glucose transporters () and genes impacting GLUT4 expression and translocation in the gluteal muscle. Five fit Thoroughbred horses performed glycogen-depleting exercises on high-starch (HS, 2869 g starch/day) and low-starch, high-fat diets (LS-HF, 358 g starch/d) with gluteal muscle biopsies obtained before and after depletion and during repletion. Muscle glycogen declined by ≈30% on both diets with little increase during repletion on LS-HF. Transcrip...
Essén-Gustavsson B, Connysson M, Jansson A.There is little information about the influence of crude protein (CP) intake on glycogen and free pool amino acid concentrations in the muscle of horses in training. High energy forage-only diets may be an alternative to concentrate rich diets and may provide high levels of CP. Objective: To study the effect of feeding 2 forage-only diets, containing either high or moderate CP concentrations on glycogen and free pool amino acid concentrations in the muscle. Methods: Two high energy forage-only diets based on high-energy grass forage were fed for 23 days in a crossover design to 6 Standardbred ...
Firshman AM, Valberg SJ, Baird JD, Hunt L, DiMauro S.To determine insulin sensitivity, proportions of muscle fiber types, and activities of glycogenolytic and glycolytic enzymes in Belgians with and without polysaccharide storage myopathy (PSSM). Methods: 10 Quarter Horses (QHs) and 103 Belgians in which PSSM status had been determined. Methods: To determine insulin sensitivity, a hyperinsulinemic euglycemic clamp (HEC) technique was used in 5 Belgians with PSSM and 5 Belgians without PSSM. Insulin was infused i.v. at 3 mU/min/kg for 3 hours, and concentrations of blood glucose and plasma insulin were determined throughout. An i.v. infusion of g...
Schuback K, Essén-Gustavsson B.The purpose of this study was to develop a standardised maximal treadmill exercise test performed until fatigue in order to find reproducible markers for anaerobic metabolism, specifically adenine nucleotide degradation. Six Standardbred trotters performed an incremental maximal treadmill exercise test in 1 min steps (starting with 7 m/s) until they could no longer keep pace with the treadmill. The test was performed twice with at least one week between the tests. Heart rate was recorded and venous blood samples were obtained during the test and in the recovery period for determination of plas...
