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Effect of Speed Endurance Training and Reduced Training Volume On View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Victoria University Eprints Repository Physiological Reports ISSN 2051-817X ORIGINAL RESEARCH Effect of speed endurance training and reduced training volume on running economy and single muscle fiber adaptations in trained runners Casper Skovgaard1,2, Danny Christiansen3, Peter M. Christensen1,2, Nicki W. Almquist1, Martin Thomassen1 & Jens Bangsbo1 1 Department of Nutrition, Exercise and Sports, Section of Integrative Physiology, University of Copenhagen, Copenhagen, Denmark 2 Team Danmark (Danish Elite Sports Organization), Copenhagen, Denmark 3 Institute of Sport, Exercise and Active Living (ISEAL), Victoria University, Melbourne, Australia Keywords Abstract Intense training, muscle fiber type-specific adaptations, muscular adaptations, sprint The aim of the present study was to examine whether improved running interval training. economy with a period of speed endurance training and reduced training vol- ume could be related to adaptations in specific muscle fibers. Twenty trained Correspondence male (n = 14) and female (n = 6) runners (maximum oxygen consumption Jens Bangsbo, University of Copenhagen, (VO2-max): 56.4 Æ 4.6 mL/min/kg) completed a 40-day intervention with 10 Department of Nutrition, Exercise and Sports, sessions of speed endurance training (5–10 9 30-sec maximal running) and a Section of Integrative Physiology, August reduced (36%) volume of training. Before and after the intervention, a muscle Krogh Building, Universitetsparken 13, 2100 Copenhagen O, Denmark. biopsy was obtained at rest, and an incremental running test to exhaustion Tel: +45 35 32 16 23 was performed. In addition, running at 60% vVO2-max, and a 10-km run was Fax: +45 35 32 16 00 performed in a normal and a muscle slow twitch (ST) glycogen-depleted con- E-mail: [email protected] dition. After compared to before the intervention, expression of mitochondrial uncoupling protein 3 (UCP3) was lower (P < 0.05) and dystrophin was higher Funding Information (P < 0.05) in ST muscle fibers, and sarcoplasmic reticulum calcium ATPase 1 The study was supported by a grant from (SERCA1) was lower (P < 0.05) in fast twitch muscle fibers. Running econ- Team Danmark (Danish elite sports Æ Æ organization), Copenhagen, Denmark. omy at 60% vVO2-max (11.6 0.2 km/h) and at v10-km (13.7 0.3 km/h) was ~2% better (P < 0.05) after the intervention in the normal condition, but Received: 26 October 2017; Revised: 7 unchanged in the ST glycogen-depleted condition. Ten kilometer performance January 2018; Accepted: 9 January 2018 was improved (P < 0.01) by 3.2% (43.7 Æ 1.0 vs. 45.2 Æ 1.2 min) and 3.9% (45.8 Æ 1.2 vs. 47.7 Æ 1.3 min) in the normal and the ST glycogen-depleted doi: 10.14814/phy2.13601 condition, respectively. VO2-max was the same, but vVO2-max was 2.0% higher (P < 0.05; 19.3 Æ 0.3 vs. 18.9 Æ 0.3 km/h) after than before the inter- Physiol Rep, 6 (3), 2018, e13601, https://doi.org/10.14814/phy2.13601 vention. Thus, improved running economy with intense training may be related to changes in expression of proteins linked to energy consuming pro- cesses in primarily ST muscle fibers. Introduction However, the mechanisms causing the improved running economy are not clearly identified, but may be related to Speed endurance training (SET; 10–40 sec repeated “all- metabolic changes in the trained muscles (Saunders et al. out” efforts with rest periods lasting >5 times the exercise 2004). bouts) with a concomitant reduced training volume has Training-induced improvement in running economy been found to improve endurance performance in associ- may be due to higher mitochondrial efficiency, that is, ation with better running economy at submaximal speeds higher ATP/O2, which could be due to reduced uncou- in trained runners (Bangsbo et al. 2009; Bangsbo 2015). pled respiration. The mitochondrial uncoupling protein 3 ª 2018 The Authors. Physiological Reports published by Wiley Periodicals, Inc. on behalf of 2018 | Vol. 6 | Iss. 3 | e13601 The Physiological Society and the American Physiological Society Page 1 This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. Muscle Fiber Type Adaptations to Speed Endurance Training in Runners C. Skovgaard et al. (UCP3) is suggested to be involved in thermogenesis by activity between ST and FT fibers in gastrocnemius mus- dispersing energy as heat instead of converting it to ATP cle of elite orienteers, suggesting that FT fibers have the (Gong et al. 1997; Boss et al. 2000) and improved run- ability to metabolically adapt to high oxidative demands ning economy may therefore be related to reduced levels (Jansson and Kaijser 1977). Metabolic adaptations in FT of muscle UCP3. In agreement, cross-sectional studies fibers may therefore contribute to improving running have shown that endurance-trained subjects have lower economy after intense training, such as SET, targeting muscle UCP3 expression and better running economy both ST and FT fibers (Egan and Zierath 2013). In sup- than untrained subjects (Russell et al. 2003a,b; Mogensen port, augmented mRNA response related to mitochon- et al. 2006). However, Iaia et al. (2009) found no change drial biogenesis (peroxisome proliferator-activated in whole muscle UCP3 level although running economy receptor-c coactivator-1, PGC-1a) and metabolism (hex- improved after 4 weeks of SET and a 65% reduced train- okinase II and pyruvate dehydrogenase kinase-4, PDK4) ing volume. Thus, studies should investigate whether in trained subjects was observed following a SET session changes in the single muscle fiber expression of UCP3 (Skovgaard et al. 2016). Furthermore, PGC-1a mRNA has could be related to changes in running economy. been shown to increase in an exercise intensity-dependent The transfer of muscle force produced by the acto- manner (Egan et al. 2010; Nordsborg et al. 2010). Regular myosins involves a secondary matrix of proteins that trans- intense training may therefore lead to higher oxidative mit the muscle force along and between muscle fibers and capacity, possibly due to oxidative adaptations in FT out to the extracellular matrix. Cytoskeleton proteins, such fibers, which in turn could contribute to the improved as dystrophin, have been identified as playing a role in this running economy as a result of the intense training (Iaia muscle force transmission (Rybakova et al. 2000; Prins et al. 2008; Bangsbo et al. 2009; Iaia and Bangsbo 2010; et al. 2009) and changes in the expression of these proteins Skovgaard et al. 2014). could influence the integrity and the strength of the muscle In vitro studies have shown that the energy cost of con- (Hughes et al. 2015). Hence, increased expression of mus- traction is higher in FT than ST fibers (Crow and Kushmer- cle dystrophin may result in increased rate of force devel- ick 1982; Barclay et al. 1993; He et al. 2000). This was opment, increased muscular power output and greater confirmed in vivo by Krustrup et al. (2008) who observed storage and return of elastic energy thereby lowering the that the oxygen uptake for at given exercise intensity was cost of running (i.e., improve running economy). higher when ST fibers were blocked by a neuromuscular Another potential cause of training-induced improve- blocking agent. And reports by Krustrup et al. (2004), who ments in running economy is lowered muscle expression depleted the ST fibers the day before submaximal exercise, of the sarcoplasmic reticulum (SR) Ca2+-ATPase (SERCA) that the glycogen depletion of ST fibers enhanced the pumps, as they are suggested to be responsible for up to recruitment of FT fibers and elevated the energy require- 50% of the ATP used during muscle activity (Clausen ment by 7% (Krustrup et al. 2004). By using the approach, et al. 1991; Walsh et al. 2006; Smith et al. 2013). Studies of depleting ST fibers the day before exercise (Krustrup have shown that speed endurance training modulates et al. 2004), before and after a SET period, it may be possi- skeletal muscle fiber type distribution in soccer players ble to study whether a change in running economy is (Gunnarsson et al. 2012) and runners (Skovgaard et al. caused by specific adaptations in FT fibers. 2014), which has been found together with lowered Thus, the aims of the present study were in trained run- SERCA1 expression (Skovgaard et al. 2014) and improved ners to investigate the effect of intensified training, in the running economy. Muscle fibers with high SERCA1 form of speed endurance training and a reduced volume of expression have a faster release and uptake of Ca2+ (Del- aerobic training, on running economy and adaptation of bono and Meissner 1996; Froemming et al. 2000) and single muscle fibers. We hypothesized that FT muscle fibers lowered expression of SERCA1 may therefore reduce the would adapt to the training by lowered expression of energy turnover during exercise. UCP3 and SERCA1, and increased expression of dys- An increase in the respiratory capacity of skeletal mus- trophin and CS, which would be associated with improved cle permits the use of less oxygen per mitochondrial res- running economy and 10-km running performance. piratory chain for a given submaximal running speed (Saunders et al. 2004). Slow twitch (ST) muscle fibers Methods have higher mitochondrial content and are more depen- dent on oxidative metabolism than fast twitch (FT) mus- Subjects cle fibers (Berchtold et al. 2000; Schiaffino and Reggiani 2011). However, Jansson and Kaijser (1977) reported that, Twenty-six trained runners commenced the study. Six sub- unlike a control group of varying physical fitness, there jects did not complete the intervention period due to per- was no difference in succinate dehydrogenase muscle sonal circumstances (n = 4) or low adherence to the 2018 | Vol.
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