Creatine Supplementation: A Comparison of Loading and Maintenance Protocols on Creatine Uptake by Human Skeletal Muscle

2003 ◽  
Vol 13 (1) ◽  
pp. 97-111 ◽  
Author(s):  
David Preen ◽  
Brian Dawson ◽  
Carmel Goodman ◽  
John Beilby ◽  
Simon Ching
Keyword(s):  
Skeletal Muscle ◽  
Human Skeletal Muscle ◽  
Vastus Lateralis ◽  
Effective Means ◽  
Creatine Supplementation ◽  
Whole Body ◽  
Physically Active ◽  
Treatment Conditions ◽  
Creatine Uptake ◽  
The Impact

The purposes of this investigation were first to determine the impact of 3 different creatine (Cr) loading procedures on skeletal muscle total Cr (TCr) accumulation and, second, to evaluate the effectiveness of 2 maintenance regimes on retaining intramuscular TCr stores, in the 6 weeks following a 5-day Cr loading program (20 g · day−1). Eighteen physically active male subjects were divided into 3 equal groups and administered either: (a) Cr (4 X 5 g · day−1 X 5 days), (b) Glucose+Cr (1 g · kg−1 of body mass twice per day), or (c) Cr in conjunction with 60 min of daily muscular (repeated-sprint) exercise. Following the 5-day loading period, subjects were reassigned to 3 maintenance groups and ingested either 0 g · day−1, 2 g · day−1 or 5 g · day−1 of Cr for a period of 6 weeks. Muscle biopsy samples (vastus lateralis) were taken pre- and post-loading as well as post-maintenance and analyzed for skeletal muscle ATP, phosphocreatine (PCr), Cr, and TCr concentrations. Twenty-four hour urine samples were collected for each of the loading days and last 2 maintenance days, and used to determine whole body Cr retention. Post-loading TCr stores were significantly (p < .05) increased in all treatment conditions. The Glucose+Cr condition produced a greater elevation (p < .05) in TCr concentrations (25%) than the Cr Only (16%) or Exercise+Cr (18%) groups. Following the maintenance period, muscle TCr stores were still similar to post-loading values for both the 2 g · day−1 and 5 g · day−1 conditions. Intramuscular TCr values for the 0 g · day−1 condition were significantly lower than the other conditions after the 6-week period. Although not significantly different from pre-loading concentrations, muscle TCr for the 0 g · day−1 group had not fully returned to baseline levels at 6 weeks post-loading. The data suggests that Glucose+Cr (but with a much smaller glucose intake than currently accepted) is potentially the most effective means of elevating TCr accumulation in human skeletal muscle. Furthermore, after 5 days of Cr loading, elevated muscle TCr concentrations can be maintained by the ingestion of small daily Cr doses (2-5 g) for a period of 6 weeks and that TCr concentrations may take longer than currently accepted to return to baseline values after such a Cr loading regime.

Effect of α-Lipoic Acid Combined with Creatine Monohydrate on Human Skeletal Muscle Creatine and Phosphagen Concentration

2003 ◽  
Vol 13 (3) ◽  
pp. 294-302 ◽  
Author(s):  
Darren G. Burke ◽  
Philip D. Chilibeck ◽  
Gianni Parise ◽  
Mark A. Tarnopolsky ◽  
Darren G. Candow
Keyword(s):  
Skeletal Muscle ◽  
Lipoic Acid ◽  
Human Skeletal Muscle ◽  
Vastus Lateralis ◽  
Creatine Supplementation ◽  
Creatine Monohydrate ◽  
Glucose Disposal ◽  
Creatine Uptake ◽  
Total Creatine ◽  
Muscle Creatine

α-lipoic acid has been found to enhance glucose uptake into skeletal muscle in animal models. Studies have also found that the co-ingestion of carbohydrate along with creatine increases muscle creatine uptake by a process related to insulin-stimulated glucose disposal. The purpose of this study was to determine the effect of α-lipoic acid on human skeletal muscle creatine uptake by directly measuring intramuscular concentrations of creatine, phosphocreatine, and ad-enosine triphosphate when creatine monohydrate was co-ingested with α-lipoic acid. Muscle biopsies were acquired from the vastus lateralis m. of 16 male subjects (18–32 y) before and after the experimental intervention. After the initial biopsy, subjects ingested 20 g · d−1 of creatine monohydrate, 20 g · d−1 of creatine monohydrate + 100 g · d−1 of sucrose, or 20 g · d−1 of creatine monohydrate + 100 g · d−1 of sucrose + 1000 mg · d−1 of α-lipoic acid for 5 days. Subjects refrained from exercise and consumed the same balanced diet for 7 days. Body weight increased by 2.1% following the nutritional intervention, with no differences between the groups. There was a significant increase in total creatine concentration following creatine supplementation, with the group ingesting α-lipoic acid showing a significantly greater increase (p < .05) in phosphocreatine (87.6 → 106.2 mmol · kg−1 dry mass [dm]) and total creatine (137.8 → 156.8 mmol · kg−1 dm). These findings indicate that co-ingestion of α-lipoic acid with creatine and a small amount of sucrose can enhance muscle total creatine content as compared to the ingestion of creatine and sucrose or creatine alone.

Parallel Increases in Phosphocreatine and Total Creatine in Human Vastus Lateralis Muscle during Creatine Supplementation

2007 ◽  
Vol 17 (6) ◽  
pp. 624-634 ◽  
Author(s):  
Jeffrey J. Brault ◽  
Theodore F. Towse ◽  
Jill M. Slade ◽  
Ronald A. Meyer
Keyword(s):  
Skeletal Muscle ◽  
Free Energy ◽  
Gibbs Free Energy ◽  
Human Skeletal Muscle ◽  
Atp Hydrolysis ◽  
Vastus Lateralis ◽  
Creatine Supplementation ◽  
Vastus Lateralis Muscle ◽  
Lateralis Muscle ◽  
Total Creatine

Short-term creatine supplementation is reported to result in a decreased ratio of phosphocreatine (PCr) to total creatine (TCr) in human skeletal muscle at rest. Assuming equilibrium of the creatine kinase reaction, this decrease in PCr:TCr implies increased cytoplasmic ADP and decreased Gibbs free energy of ATP hydrolysis in muscle, which seems contrary to the reported ergogenic benefits of creatine supplementation. This study measured changes in PCr and TCr in vastus lateralis muscle of adult men (N = 6, 21–35 y old) during and 1 day after 5 d of creatine monohydrate supplementation (0.43 g·kg body weight−1·d−1) using noninvasive 31P and 1H magnetic-resonance spectroscopy (MRS). Plasma and red-blood-cell creatine increased by 10-fold and 2-fold, respectively, by the third day of supplementation. MRS-measured skeletal muscle PCr and TCr increased linearly and in parallel throughout the 5 d, and there was no significant difference in the percentage increase in muscle PCr (11.7% ± 2.3% after 5 d) vs. TCr (14.9% ± 4.1%) at any time point. The results indicate that creatine supplementation does not alter the PCr:TCr ratio, and hence the cytoplasmic Gibbs free energy of ATP hydrolysis, in human skeletal muscle at rest.

Evidence against a role for insulin-signaling proteins PI 3-kinase and Akt in insulin resistance in human skeletal muscle induced by short-term GH infusion

2005 ◽  
Vol 288 (1) ◽  
pp. E194-E199 ◽  
Author(s):  
Niels Jessen ◽  
Christian B. Djurhuus ◽  
Jens O. L. Jørgensen ◽  
Lasse S. Jensen ◽  
Niels Møller ◽  
...  
Keyword(s):  
Insulin Resistance ◽  
Skeletal Muscle ◽  
Insulin Signaling ◽  
Infusion Rate ◽  
Human Skeletal Muscle ◽  
Vastus Lateralis ◽  
Fold Increase ◽  
Saline Infusion ◽  
Short Term ◽  
The Impact

Prolonged growth hormone (GH) excess is known to be associated with insulin resistance, but the underlying mechanisms remain unknown. The aim of this study was to assess the impact of GH on insulin-stimulated glucose metabolism and insulin signaling in human skeletal muscle. In a cross-over design, eight healthy male subjects (age 26.0 ± 0.8 yr and body mass index 24.1 ± 0.5 kg/m2) were infused for 360 min with either GH (Norditropin, 45 ng·kg−1·min−1) or saline. During the final 180 min of the infusion, a hyperinsulinemic euglycemic clamp was performed (insulin infusion rate: 1.2 mU·kg−1·min−1). Muscle biopsies from vastus lateralis were taken before GH/saline administration and after 60 min of hyperinsulinemia. GLUT4 content and insulin signaling, as assessed by insulin receptor substrate (IRS)-1-associated phosphatidylinositol 3-kinase and Akt activity were determined. GH levels increased to a mean (±SE) level of 20.0 ± 2.3 vs. 0.5 ± 0.2 μg/l after saline infusion ( P < 0.01). During GH infusion, the glucose infusion rate during hyperinsulinemia was reduced by 38% ( P < 0.01). In both conditions, free fatty acids were markedly suppressed during hyperinsulinemia. Despite skeletal muscle insulin resistance, insulin still induced a similar ∼3-fold rise in IRS-1-associated PI 3-kinase activity (269 ± 105 and 311 ± 71% compared with baseline, GH vs. saline). GH infusion did not change Akt protein expression, and insulin caused an ∼13-fold increase in Akt activity (1,309 ± 327 and 1,287 ± 173%) after both GH and saline infusion. No difference in total GLUT4 content was noted (114.7 ± 7.4 and 107.6 ± 16.7 arbitrary units, GH vs. saline, compared with baseline). In conclusion, insulin resistance in skeletal muscle induced by short-term GH administration is not associated with detectable changes in the upstream insulin-signaling cascade or reduction in total GLUT4. Yet unknown mechanisms in insulin signaling downstream of Akt may be responsible.

Effect of exercise and training on phospholemman phosphorylation in human skeletal muscle

2011 ◽  
Vol 301 (3) ◽  
pp. E456-E466 ◽  
Author(s):  
Boubacar Benziane ◽  
Ulrika Widegren ◽  
Sergej Pirkmajer ◽  
Jan Henriksson ◽  
Nigel K. Stepto ◽  
...  
Keyword(s):  
Skeletal Muscle ◽  
Human Skeletal Muscle ◽  
Acute Exercise ◽  
Vastus Lateralis ◽  
Exercise Bout ◽  
Short Term ◽  
Atpase Subunit ◽  
Acute Bout ◽  
Subunit Expression ◽  
The Impact

Phospholemman (PLM, FXYD1) is a partner protein and regulator of the Na+-K+-ATPase (Na+-K+ pump). We explored the impact of acute and short-term training exercise on PLM physiology in human skeletal muscle. A group of moderately trained males ( n = 8) performed a 1-h acute bout of exercise by utilizing a one-legged cycling protocol. Muscle biopsies were taken from vastus lateralis at 0 and 63 min (non-exercised leg) and 30 and 60 min (exercised leg). In a group of sedentary males ( n = 9), we determined the effect of a 10-day intense aerobic cycle training on Na+-K+-ATPase subunit expression, PLM phosphorylation, and total PLM expression as well as PLM phosphorylation in response to acute exercise (1 h at ∼72% V̇o2peak). Biopsies were taken at rest, immediately following, and 3 h after an acute exercise bout before and at the conclusion of the 10-day training study. PLM phosphorylation was increased both at Ser63 and Ser68 immediately after acute exercise (75%, P < 0.05, and 30%, P < 0.05, respectively). Short-term training had no adaptive effect on PLM phosphorylation at Ser63 and Ser68, nor was the total amount of PLM altered posttraining. The protein expressions of α1-, α2-,and β1-subunits of Na+-K+-ATPase were increased after training (113%, P < 0.05, 49%, P < 0.05, and 27%, P < 0.05, respectively). Whereas an acute bout of exercise increased the phosphorylation of PKCα/βII on Thr638/641 pre- and posttraining, phosphorylation of PKCζ/λ on Thr403/410 was increased in response to acute exercise only after the 10-day training. In conclusion, we show that only acute exercise, and not short-term training, increases phosphorylation of PLM on Ser63 and Ser68, and data from one-legged cycling indicate that this effect of exercise on PLM phosphorylation is not due to systemic factors. Our results provide evidence that phosphorylation of PLM may play a role in the acute regulation of the Na+-K+-ATPase response to exercise.

Antecubital Vein Cannula Position Impacts Assessment of Forearm Glucose Uptake During an Oral Glucose Challenge in Healthy Volunteers

2020 ◽  
Author(s):  
Cécile Bétry ◽  
Aline V. Nixon ◽  
Paul L. Greenhaff ◽  
Elizabeth J. Simpson
Keyword(s):  
Skeletal Muscle ◽  
Blood Glucose ◽  
Glucose Uptake ◽  
Venous Blood ◽  
Whole Body ◽  
Oral Glucose ◽  
Antecubital Vein ◽  
Glucose Challenge ◽  
The Impact

Abstract Introduction Skeletal muscle is a major site for whole-body glucose disposal, and determination of skeletal muscle glucose uptake is an important metabolic measurement, particularly in research focussed on interventions that impact muscle insulin sensitivity. Calculating arterial-venous difference in blood glucose can be used as an indirect measure for assessing glucose uptake. However, the possibility of multiple tissues contributing to the composition of venous blood, and the differential in glucose uptake kinetics between tissue types, suggests that sampling from different vein sites could influence the estimation of glucose uptake. This study aimed to determine the impact of venous cannula position on calculated forearm glucose uptake following an oral glucose challenge in resting and post-exercise states. Materials and Methods In 9 young, lean, males, the impact of sampling blood from two antecubital vein positions; the perforating vein (‘perforating’ visit) and, at the bifurcation of superficial and perforating veins (‘bifurcation’ visit), was assessed. Brachial artery blood flow and arterialised-venous and venous blood glucose concentrations were measured in 3 physiological states; resting-fasted, resting-fed, and fed following intermittent forearm muscle contraction (fed-exercise). Results Following glucose ingestion, forearm glucose uptake area under the curve was greater for the ‘perforating’ than for the ‘bifurcation’ visit in the resting-fed (5.92±1.56 vs. 3.69±1.35 mmol/60 min, P<0.01) and fed-exercise (17.38±7.73 vs. 11.40±7.31 mmol/75 min, P<0.05) states. Discussion Antecubital vein cannula position impacts calculated postprandial forearm glucose uptake. These findings have implications for longitudinal intervention studies where serial determination of forearm glucose uptake is required.

RPS6 phosphorylation occurs to a greater extent in the periphery of human skeletal muscle fibers, near focal adhesions, after anabolic stimuli

2021 ◽  
Author(s):  
Nathan Hodson ◽  
Michael Mazzulla ◽  
Maksym N. H. Holowaty ◽  
Dinesh Kumbhare ◽  
Daniel R. Moore
Keyword(s):  
Skeletal Muscle ◽  
Focal Adhesion ◽  
Human Skeletal Muscle ◽  
Muscle Fibers ◽  
Focal Adhesions ◽  
Immunofluorescent Staining ◽  
Whole Body ◽  
Vastus Lateralis Muscle ◽  
Cell Periphery ◽  
Rps6 Phosphorylation

Following anabolic stimuli (mechanical loading and/or amino acid provision) the mechanistic target of rapamycin complex 1 (mTORC1), a master regulator of protein synthesis, translocates toward the cell periphery. However, it is unknown if mTORC1-mediated phosphorylation events occur in these peripheral regions or prior to translocation (i.e. in central regions). We therefore aimed to determine the cellular location of a mTORC1-mediated phosphorylation event, RPS6Ser240/244, in human skeletal muscle following anabolic stimuli. Fourteen young, healthy males either ingested a protein-carbohydrate beverage (0.25g/kg protein, 0.75g/kg carbohydrate) alone (n=7;23±5yrs;76.8±3.6kg;13.6±3.8%BF, FED) or following a whole-body resistance exercise bout (n=7;22±2yrs;78.1±3.6kg;12.2±4.9%BF, EXFED). Vastus lateralis muscle biopsies were obtained at rest (PRE) and 120 and 300min following anabolic stimuli. RPS6Ser240/244 phosphorylation measured by immunofluorescent staining or immunoblot was positively correlated (r=0.76, p<0.001). Peripheral staining intensity of p-RPS6Ser240/244 increased above PRE in both FED and EXFED at 120min (~54% and ~138% respectively, p<0.05) but was greater in EXFED at both post-stimuli time points (p<0.05). The peripheral-central ratio of p-RPS6240/244 staining displayed a similar pattern, even when corrected for total RPS6 distribution, suggesting RPS6 phosphorylation occurs to a greater extent in the periphery of fibers. Moreover, p-RPS6Ser240/244 intensity within paxillin-positive regions, a marker of focal adhesion complexes, was elevated at 120min irrespective of stimulus (p=0.006) before returning to PRE at 300min. These data confirm that RPS6Ser240/244 phosphorylation occurs in the region of human muscle fibers to which mTOR translocates following anabolic stimuli and identifies focal adhesion complexes as a potential site of mTORC1 regulation in vivo.

Alterations in Skeletal Muscle Repair in Young Adults with Type 1 Diabetes Mellitus

2021 ◽  
Author(s):  
Athan G. Dial ◽  
Grace K. Grafham ◽  
Cynthia MF. Monaco ◽  
Jennifer Voth ◽  
Linda Brandt ◽  
...  
Keyword(s):  
Electron Microscopy ◽  
Skeletal Muscle ◽  
Type 1 Diabetes ◽  
Young Adults ◽  
Gene Networks ◽  
Vastus Lateralis ◽  
Post Exercise ◽  
Ultrastructural Damage ◽  
The Impact

Though preclinical models of type 1 diabetes (T1D) exhibit impaired muscle regeneration, this has yet to be investigated in humans with T1D. Here we investigated the impact of damaging exercise (eccentric quadriceps contractions) in eighteen physically-active young adults with and without T1D. Pre- and post-exercise (48h and 96h), participants provided blood samples, vastus lateralis biopsies and performed maximal voluntary quadriceps contractions (MVC). Skeletal muscle sarcolemmal integrity, extracellular matrix content (ECM), and satellite cell (SC) content/proliferation were assessed by immunofluorescence. Transmission electron microscopy was used to quantify ultrastructural damage. MVC was comparable between T1D and controls before exercise. Post-exercise, MVC was decreased in both groups, but T1D subjects exhibited moderately lower strength recovery at both 48h and 96h. Serum creatine kinase, an indicator of muscle damage, was moderately higher in T1D participants at rest, and exhibited a small elevation 96h post-exercise. T1D participants showed lower SC content at all timepoints and demonstrated a moderate delay in SC proliferation after exercise. A greater number of myofibers exhibited sarcolemmal damage (disrupted dystrophin) and increased ECM (laminin) content in participants with T1D despite no differences between groups in ultrastructural damage as assessed by electron microscopy. Finally, transcriptomic analyses revealed dysregulated gene networks involving RNA translation and mitochondrial respiration, providing potential explanations for previous observations of mitochondrial dysfunction in similar T1D cohorts. Our findings indicate that skeletal muscle in young adults with moderately-controlled T1D is altered after damaging exercise; suggesting that longer recovery times following intense exercise may be necessary.

Phosphorylase a in human skeletal muscle during exercise and electrical stimulation

1978 ◽  
Vol 45 (6) ◽  
pp. 852-857 ◽  
Author(s):  
P. D. Gollnick ◽  
J. Karlsson ◽  
K. Piehl ◽  
B. Saltin
Keyword(s):  
Skeletal Muscle ◽  
Electrical Stimulation ◽  
Human Skeletal Muscle ◽  
Vastus Lateralis ◽  
Muscular Activity ◽  
Voluntary Exercise ◽  
Vastus Lateralis Muscle ◽  
Minor Importance ◽  
Muscle Lactate ◽  
Needle Technique

Experiments were conducted to examine the conversions of phosphorylase b to phosphorylase a in human skeletal muscle during bicycle exercise or isometric contractions. Muscle biopsies were obtained from the vastus lateralis with the needle technique at rest and either during or immediately after activity and frozen in liquid nitrogen within 2--4 s. Total phosphorylase and phosphorylase a activities were differentiated by measurement in the presence and absence of AMP, respectively. At rest 8.5% of the total phosphorylase activity existed in the a form. Little or no change in the percent of phosphorylase in the a form occurred during voluntary dynamic or static muscular activity that produced muscle lactate concentrations in excess of 18 mmol.kg-1 wet muscle. Electrical stimulation of the vastus lateralis muscle also failed to produce an increase in the percentage of phosphorylase a. These data suggest that during exercise the conversion of phosphorylase to the a form is of minor importance. An increased activity of phosphorylase b due to changes in muscle concentrations of ATP, AMP, and inorganic phosphate may regulate glycogenolysis during voluntary exercise in man.

scholarly journals Contractile activity-specific transcriptome response to acute endurance exercise and training in human skeletal muscle

2019 ◽  
Vol 316 (4) ◽  
pp. E605-E614 ◽  
Author(s):  
Daniil V. Popov ◽  
Pavel A. Makhnovskii ◽  
Elena I. Shagimardanova ◽  
Guzel R. Gazizova ◽  
Evgeny A. Lysenko ◽  
...  
Keyword(s):  
Skeletal Muscle ◽  
Transcription Factors ◽  
Human Skeletal Muscle ◽  
Acute Exercise ◽  
Contractile Activity ◽  
Vastus Lateralis ◽  
Aerobic Exercise Training ◽  
Vastus Lateralis Muscle ◽  
Transcriptome Response ◽  
The One

Reduction in daily activity leads to dramatic metabolic disorders, while regular aerobic exercise training is effective for preventing this problem. The purpose of this study was to identify genes that are directly related to contractile activity in human skeletal muscle, regardless of the level of fitness. Transcriptome changes after the one-legged knee extension exercise in exercised and contralateral nonexercised vastus lateralis muscle of seven men were evaluated by RNA-seq. Transcriptome change at baseline after 2 mo of aerobic training (5/wk, 1 h/day) was evaluated as well. Postexercise changes in the transcriptome of exercised muscle were associated with different factors, including circadian oscillations. To reveal transcriptome response specific for endurance-like contractile activity, differentially expressed genes between exercised and nonexercised muscle were evaluated at 1 and 4 h after the one-legged exercise. The contractile activity-specific transcriptome responses were associated only with an increase in gene expression and were regulated mainly by CREB/ATF/AP1-, MYC/MAX-, and E2F-related transcription factors. Endurance training-induced changes (an increase or decrease) in the transcriptome at baseline were more pronounced than transcriptome responses specific for acute contractile activity. Changes after training were associated with widely different biological processes than those after acute exercise and were regulated by different transcription factors (IRF- and STAT-related factors). In conclusion, adaptation to regular exercise is associated not only with a transient (over several hours) increase in expression of many contractile activity-specific genes, but also with a pronounced change (an increase or decrease) in expression of a large number of genes under baseline conditions.

Eccentric exercise markedly increases c-Jun NH2-terminal kinase activity in human skeletal muscle

1999 ◽  
Vol 87 (5) ◽  
pp. 1668-1673 ◽  
Author(s):  
Marni D. Boppart ◽  
Doron Aronson ◽  
Lindsay Gibson ◽  
Ronenn Roubenoff ◽  
Leslie W. Abad ◽  
...  
Keyword(s):  
Skeletal Muscle ◽  
Protein Kinase ◽  
Eccentric Exercise ◽  
Intracellular Signaling ◽  
Human Skeletal Muscle ◽  
Vastus Lateralis ◽  
Fold Increase ◽  
Mitogen Activated Protein Kinase ◽  
Eccentric Contractions ◽  
Vastus Lateralis Muscle

Eccentric contractions require the lengthening of skeletal muscle during force production and result in acute and prolonged muscle injury. Because a variety of stressors, including physical exercise and injury, can result in the activation of the c-Jun NH2-terminal kinase (JNK) intracellular signaling cascade in skeletal muscle, we investigated the effects of eccentric exercise on the activation of this stress-activated protein kinase in human skeletal muscle. Twelve healthy subjects (7 men, 5 women) completed maximal concentric or eccentric knee extensions on a KinCom isokinetic dynamometer (10 sets, 10 repetitions). Percutaneous needle biopsies were obtained from the vastus lateralis muscle 24 h before exercise (basal), immediately postexercise, and 6 h postexercise. Whereas both forms of exercise increased JNK activity immediately postexercise, eccentric contractions resulted in a much higher activation (15.4 ± 4.5 vs. 3.5 ± 1.4-fold increase above basal, eccentric vs. concentric). By 6 h after exercise, JNK activity decreased back to baseline values. In contrast to the greater activation of JNK with eccentric exercise, the mitogen-activated protein kinase kinase 4, the immediate upstream regulator of JNK, was similarly activated by concentric and eccentric exercise. Because the activation of JNK promotes the phosphorylation of a variety of transcription factors, including c-Jun, the results from this study suggest that JNK may be involved in the molecular and cellular adaptations that occur in response to injury-producing exercise in human skeletal muscle.

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