{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,1]],"date-time":"2026-04-01T22:31:45Z","timestamp":1775082705890,"version":"3.50.1"},"reference-count":58,"publisher":"MDPI AG","issue":"1","license":[{"start":{"date-parts":[[2021,12,23]],"date-time":"2021-12-23T00:00:00Z","timestamp":1640217600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Cells"],"abstract":"<jats:p>The purpose of our study was to identify the features of metabolic regulation in highly trained cross-country skiers of different qualifications at different stages of the maximum load test. We examined 124 highly trained cross-country skiers (male, ages 17\u201324). The group consisted of two subgroups based on their competition performance: 61 nonelite athletes (Group I) and 63 elite athletes (group II), who were current members of the national team of the Komi Republic and Russia. The bicycle ergometer test was performed by using the OxyconPro system (Erich Jaeger, Hoechberg, Germany). All the examined athletes performed the exercise test on a cycle ergometer \u201cuntil exhaustion\u201d. The results of our research indicate that the studied groups of athletes with high, but different levels of sports qualifications are a convenient model for studying the molecular mechanisms of adaptation to physical loads of maximum intensity. Athletes of higher qualifications reveal additional adaptive mechanisms of metabolic regulation, which is manifested in the independence of serum lactate indicators under conditions of submaximal and maximum power from maximal oxygen uptake, and they have an NO-dependent mechanism for regulating lactate levels during aerobic exercise, including work at the anaerobic threshold.<\/jats:p>","DOI":"10.3390\/cells11010039","type":"journal-article","created":{"date-parts":[[2021,12,23]],"date-time":"2021-12-23T21:40:21Z","timestamp":1640295621000},"page":"39","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":6,"title":["Features of Metabolic Support of Physical Performance in Highly Trained Cross-Country Skiers of Different Qualifications during Physical Activity at Maximum Load"],"prefix":"10.3390","volume":"11","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-1862-6936","authenticated-orcid":false,"given":"Olga I.","family":"Parshukova","sequence":"first","affiliation":[{"name":"Institute of Physiology of Komi Science Centre of the Ural Branch of the Russian Academy of Sciences, FRC Komi SC UB RAS, 50 Pervomayskaya Str., 167982 Syktyvkar, Russia"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1444-4684","authenticated-orcid":false,"given":"Nina G.","family":"Varlamova","sequence":"additional","affiliation":[{"name":"Institute of Physiology of Komi Science Centre of the Ural Branch of the Russian Academy of Sciences, FRC Komi SC UB RAS, 50 Pervomayskaya Str., 167982 Syktyvkar, Russia"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4804-6908","authenticated-orcid":false,"given":"Natalya N.","family":"Potolitsyna","sequence":"additional","affiliation":[{"name":"Institute of Physiology of Komi Science Centre of the Ural Branch of the Russian Academy of Sciences, FRC Komi SC UB RAS, 50 Pervomayskaya Str., 167982 Syktyvkar, Russia"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4849-4735","authenticated-orcid":false,"given":"Aleksandra Y.","family":"Lyudinina","sequence":"additional","affiliation":[{"name":"Institute of Physiology of Komi Science Centre of the Ural Branch of the Russian Academy of Sciences, FRC Komi SC UB RAS, 50 Pervomayskaya Str., 167982 Syktyvkar, Russia"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8027-898X","authenticated-orcid":false,"given":"Evgeny R.","family":"Bojko","sequence":"additional","affiliation":[{"name":"Institute of Physiology of Komi Science Centre of the Ural Branch of the Russian Academy of Sciences, FRC Komi SC UB RAS, 50 Pervomayskaya Str., 167982 Syktyvkar, Russia"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2021,12,23]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"148","DOI":"10.3389\/fcvm.2019.00148","article-title":"Influence of vigorous physical activity on structure and function of the cardiovascular system in young athletes\u2014The MuCAYA-Study","volume":"6","author":"Baumgartner","year":"2019","journal-title":"Front. Cardiovasc. Med."},{"key":"ref_2","first-page":"31","article-title":"Possible mechanisms of sudden cardiac death in top athletes: A basic cardiac electrophysiological point of view","volume":"460","author":"Varro","year":"2010","journal-title":"Cardiovasc. Physiol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"14","DOI":"10.14529\/hsm190402","article-title":"Anxiety level and parameters of the cardiovascular system in athletes of various qualifications","volume":"19","author":"Turbasova","year":"2019","journal-title":"Hum. Sport Med."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1152\/jappl.1967.23.3.353","article-title":"Maximal oxygen uptake in athletes","volume":"23","author":"Saltin","year":"1967","journal-title":"J. Appl. Physiol."},{"key":"ref_5","first-page":"1","article-title":"Nitric oxide production in professional skiers during physical activity at maximum load","volume":"7","author":"Parshukova","year":"2020","journal-title":"Front. Cardiovasc. Med. Hypertens."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"165913","DOI":"10.1016\/j.bbadis.2020.165913","article-title":"The effects of regular aerobic exercise training on blood nitric oxide levels and oxidized LDL and the role of eNOS intron 4a\/b polymorphism","volume":"1866","author":"Yol","year":"2020","journal-title":"BBA Molec. Bas. Dis."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"1783","DOI":"10.1007\/s00421-021-04601-y","article-title":"Bicycle spiroergometry: Comparison of standardized examination protocols for adolescents: Is it necessary to define own standard values for each protocol?","volume":"121","author":"Windhaber","year":"2021","journal-title":"Eur. J. Appl. Physiol."},{"key":"ref_8","first-page":"38","article-title":"A comparison of methodologies in detection of the anaerobic threshold","volume":"81","author":"Dickstein","year":"1990","journal-title":"Circulation"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"787","DOI":"10.1378\/chest.111.3.787","article-title":"Dangerous curves. A perspective on exercise, lactate, and the anaerobic threshold","volume":"111","author":"Myers","year":"1997","journal-title":"Chest"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"142","DOI":"10.1016\/S0076-6879(96)68016-1","article-title":"Measurement of nitrate and nitrite in biological samples using nitrate reductase and Griess reaction","volume":"268","author":"Granger","year":"1996","journal-title":"Methods Enzymol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"542","DOI":"10.1177\/1941738113479763","article-title":"What performance characteristics determine elite versus nonelite athletes in the same sport?","volume":"5","author":"Lorenz","year":"2013","journal-title":"Sports Health"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"635651","DOI":"10.3389\/fpsyg.2021.635651","article-title":"Differences in the psychological profiles of elite and non-elite athletes","volume":"12","author":"Mitic","year":"2021","journal-title":"Front. Psychol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1113\/jphysiol.2003.059287","article-title":"Maximal muscular vascular conductances during whole body upright exercise in humans","volume":"558","author":"Calbet","year":"2004","journal-title":"J. Physiology"},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Martin, S.A., and Hadma\u0219, R.M. (2019). Individual adaptation in cross-country skiing based on tracking during training conditions. Sports, 7.","DOI":"10.3390\/sports7090211"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1003","DOI":"10.1123\/ijspp.2016-0749","article-title":"Physiological capacity and training routines of elite cross-country skiers: Approaching the per limits of human endurance","volume":"1","author":"Sandbakk","year":"2017","journal-title":"Int. J. Space Physiol. Perform."},{"key":"ref_16","first-page":"616","article-title":"Acute hemodynamic and cardiovascular responses following resistance exercise to voluntary exhaustion. Effects of different loadings and exercise durations","volume":"56","author":"Gjovaag","year":"2016","journal-title":"J. Sports Med. Phys. Fit."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"56","DOI":"10.17116\/terarkh2017891256-63","article-title":"Annual dynamics of blood pressure and me-Teosensitivity in women","volume":"12","author":"Varlamova","year":"2017","journal-title":"Ther. Arch."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1016\/S0049-3848(99)00028-6","article-title":"The effect of short-term cold exposure on risk factors for cardiovascular diseases from China","volume":"95","author":"Mercer","year":"1999","journal-title":"Thromb. Res."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"3197","DOI":"10.3390\/ijerph9093197","article-title":"Artificial cold air increases the cardiovascular risks in spontaneously hypertensive rats","volume":"9","author":"Luo","year":"2012","journal-title":"Int. J. Environ. Res. Public Health"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"42","DOI":"10.12965\/jer.150258","article-title":"The effects of strenuous exercises on resting heart rate, blood pressure, and maximal oxygen uptake","volume":"12","author":"Oh","year":"2016","journal-title":"J. Exerc. Rehabil."},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"de Sire, A., Marotta, N., Marinaro, C., Curci, C., Invernizzi, M., and Ammendolia, A. (2021). Role of physical exercise and nutraceuticals in modulating molecular pathways of osteoarthritis. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22115722"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1435","DOI":"10.1007\/s001250051551","article-title":"Tetrahydrobiopterin improves endothelium-dependent vasodilation by increasing nitric oxide activity in patients with type II Diabetes mellitus","volume":"43","author":"Heitzer","year":"2000","journal-title":"Diabetologia"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1515\/jomb-2015-0008","article-title":"NO-synthase activity in patients with coronary heart disease associated with hypertension of different age Groups","volume":"35","author":"Besedina","year":"2016","journal-title":"J. Med. Biochem."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"643","DOI":"10.1161\/01.HYP.31.2.643","article-title":"Alterations of nitric oxide synthase expression with aging and hypertension in rats","volume":"31","author":"Chou","year":"1998","journal-title":"Hypertension"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"287","DOI":"10.15421\/021745","article-title":"Protein kinase C (PKC) involved in enhancement of alpha (1)-adrenoceptor-mediated responses of the main pulmonary artery in rats with diabetes mellitus","volume":"8","author":"Kizub","year":"2017","journal-title":"Regul. Mech. Biosyst."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"356","DOI":"10.1161\/01.CIR.93.2.356","article-title":"Increased release of NO during ischemia reduces myocardial contractility and improves metabolic dysfunction","volume":"93","author":"Node","year":"1996","journal-title":"Circulation"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"1337","DOI":"10.1152\/physrev.2000.80.4.1337","article-title":"Role of nitric oxide in the pathogenesis of chronic pulmonary hypertension","volume":"80","author":"Hampl","year":"2000","journal-title":"Physiol. Rev."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"452","DOI":"10.1016\/j.molmed.2009.08.002","article-title":"Hemoglobin, nitric oxide and molecular mechanisms of hypoxic vasodilation","volume":"15","author":"Allen","year":"2009","journal-title":"Trends Mol. Med."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"105","DOI":"10.1006\/niox.1999.0213","article-title":"NO-dependent mechanisms of adaptation to hypoxia","volume":"3","author":"Malyshev","year":"1999","journal-title":"Nitric Oxide"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1186\/gm117","article-title":"Metabolic adaptation of skeletal muscle to high altitude hypoxia: How new technologies could resolve the controversies","volume":"1","author":"Murray","year":"2009","journal-title":"Genome Med."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1309","DOI":"10.1152\/jappl.1988.64.4.1309","article-title":"Operation Everest II: Oxygen transport during exercise at extreme simulated altitude","volume":"64","author":"Sutton","year":"1988","journal-title":"J. Appl. Physiol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"1551","DOI":"10.1016\/j.freeradbiomed.2009.03.009","article-title":"Enhanced leukocyte HIF-1alpha and HIF-1 DNA binding in humans after rapid ascent to 4300 m","volume":"46","author":"Serkova","year":"2009","journal-title":"Free Radic. Biol. Med."},{"key":"ref_33","first-page":"147","article-title":"Nitric oxide metabolites level in human serum in acute normobaric hypoxia","volume":"98","author":"Boiko","year":"2012","journal-title":"Rossiiskii Fiziologicheskii Zhurnal Imeni IM Sechenova"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"693","DOI":"10.3357\/AMHP.4172.2015","article-title":"A meta-analysis of exhaled nitric oxide in acute normobaric hypoxia","volume":"86","author":"MacInnis","year":"2015","journal-title":"Aerosp. Med. Hum. Perform."},{"key":"ref_35","first-page":"207","article-title":"Nitric oxide in acute brain injury: A pilot study of NO(x) concentrations in human brain microdialysates and their relationship with energy metabolism","volume":"102","author":"Carpenter","year":"2008","journal-title":"Acta Neurochir."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"549","DOI":"10.1016\/S0031-9422(99)00582-8","article-title":"Biosynthesis of brassinosteroids in cultured cells of Catharanthus roseus","volume":"53","author":"Fujioka","year":"2000","journal-title":"Phytochemistry"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"98","DOI":"10.1097\/00024382-199902000-00005","article-title":"Beneficial effects of L-canavanine, a selective inhibitor of inducible nitric oxide synthase, on lactate metabolism and muscle high energy phosphates during endotoxic shock in rats","volume":"11","author":"Levy","year":"1999","journal-title":"Shock"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"172","DOI":"10.1056\/NEJMra044389","article-title":"Tyrosine kinases as targets for cancer therapy","volume":"353","author":"Krause","year":"2005","journal-title":"N. Engl. J. Med."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1042\/bj3150295","article-title":"Rapid reduction of nitric oxide by mitochondria, and reversible inhibition of mitochondrial respiration by nitric oxide","volume":"1","author":"Borutaite","year":"1996","journal-title":"Biochem. J."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"381","DOI":"10.1016\/S0014-5793(98)00053-2","article-title":"Protein oxidation, tyrosine nitration, and inactivation of sarcoplasmic reticulum Ca21-ATPase in low-frequency stimulated rabbit muscle","volume":"422","author":"Klebl","year":"1998","journal-title":"FEBS Lett."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"7631","DOI":"10.1073\/pnas.95.13.7631","article-title":"Persistent inhibition of cell respiration by nitric oxide: Crucial role of Snitrosylation of mitochondrial complex I and protective action of glutathione","volume":"95","author":"Clementi","year":"1998","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"209","DOI":"10.1152\/physrev.2001.81.1.209","article-title":"Physiology of nitric oxide in skeletal muscle","volume":"81","author":"Stamler","year":"2001","journal-title":"Physiol. Rev."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1194","DOI":"10.1038\/4531194a","article-title":"Drug discovery: A lifeline for suffocating tissues","volume":"453","author":"Mazzone","year":"2008","journal-title":"Nature"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"752","DOI":"10.1016\/j.bbagen.2011.04.002","article-title":"Regulation cardiovascular processes by S-nitrosylation","volume":"1820","author":"Schulman","year":"2012","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"11482","DOI":"10.1073\/pnas.97.21.11482","article-title":"Role of circulating nitrite and S-nitrosohemoglobin in the regulation of the regional blood floow in humans","volume":"97","author":"Gladwin","year":"2000","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"2636","DOI":"10.1182\/blood-2008-01-115261","article-title":"The functional nitrite reductase activity of the heme globins","volume":"112","author":"Gladwin","year":"2008","journal-title":"Blood"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"1322","DOI":"10.1161\/01.RES.0000078171.52542.9E","article-title":"Nitric oxide regulation of myocardial contractility and calcium cycling","volume":"92","author":"Khan","year":"2003","journal-title":"Circ. Res."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"457","DOI":"10.1113\/jphysiol.2001.014126","article-title":"Role cyclic GMP-dependent protein kinase in the contractile response to exogenous nitric oxide in rat cardiac myocytes","volume":"540","author":"Layland","year":"2002","journal-title":"J. Physiol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"7757","DOI":"10.1074\/jbc.275.11.7757","article-title":"Reduction of nitrite to nitric oxide catalyzed by xanthine oxidoreductase","volume":"275","author":"Godber","year":"2000","journal-title":"J. Biol. Chem."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"305","DOI":"10.1080\/17461391.2014.949310","article-title":"Omega-3 fatty acids supplementation improves endothelial function and maximal oxygen uptake in endurance-trained athletes","volume":"15","author":"Zebrowska","year":"2015","journal-title":"Eur. J. Sport Sci."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"2031","DOI":"10.1046\/j.1538-7836.2003.00350.x","article-title":"Strenuous, acute exercise a Vects reciprocal modulation of platelet and polymorphonuclear leukocyte activities under shear Xow in men","volume":"1","author":"Wang","year":"2003","journal-title":"J. Thromb. Haemost."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"1454","DOI":"10.1097\/00005768-199711000-00011","article-title":"Time course of enhanced endothelium mediated dilation in aorta of trained rats","volume":"29","author":"Delp","year":"1997","journal-title":"Med. Sci. Sports Exerc."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"1164","DOI":"10.1016\/j.freeradbiomed.2004.12.018","article-title":"Regional and whole-body markers of nitric oxide production following hyperemic stimuli","volume":"38","author":"Allen","year":"2005","journal-title":"Free Radic. Biol. Med."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"790","DOI":"10.1016\/S0891-5849(03)00406-4","article-title":"Plasma nitrite resects constitutive nitric oxide synthase activity in mammals","volume":"35","author":"Kleinbongard","year":"2003","journal-title":"Free Radic. Biol. Med."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"173","DOI":"10.1177\/174182679400100212","article-title":"Exercise increases systemic nitric oxide production in men","volume":"1","author":"Boger","year":"1994","journal-title":"J. Cardiovasc. Risk"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"698","DOI":"10.1038\/s41467-020-14433-7","article-title":"Nitric oxide orchestrates metabolic rewiring in M1 macrophages by targeting aconitase 2 and pyruvate dehydrogenase","volume":"4","author":"Palmieri","year":"2020","journal-title":"Nat. Commun."},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Ilari, S., Dagostino, C., Malafoglia, V., Lauro, F., Giancotti, L.A., Spila, A., Proietti, S., Ventrice, D., Rizzo, M., and Gliozzi, M. (2020). Protective effect of antioxidants in nitric oxide\/COX-2 interaction during inflammatory pain: The role of nitration. Antioxidants, 9.","DOI":"10.3390\/antiox9121284"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"1317","DOI":"10.1111\/1756-185X.14194","article-title":"A bond between rheumatic diseases and cancer in the elderly: The interleukin-6 pathway","volume":"24","author":"Iorio","year":"2021","journal-title":"Int. J. Rheum. Dis."}],"container-title":["Cells"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2073-4409\/11\/1\/39\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T07:52:26Z","timestamp":1760169146000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2073-4409\/11\/1\/39"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,12,23]]},"references-count":58,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2022,1]]}},"alternative-id":["cells11010039"],"URL":"https:\/\/doi.org\/10.3390\/cells11010039","relation":{},"ISSN":["2073-4409"],"issn-type":[{"value":"2073-4409","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,12,23]]}}}