The Muscle Fiber Profiles, Mitochondrial Content, and Enzyme Activities of the Exceptionally Well-Trained Arm and Leg Muscles of Elite Cross-Country Skiers
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As one of the most physically demanding sports in the Olympic Games, cross-country skiing poses considerable challenges with respect to both force generation and endurance during the combined upper- and lower-body effort of varying intensity and duration. The isoforms of myosin in skeletal muscle have long been considered not only to define the contractile properties, but also to determine metabolic capacities. The current investigation was designed to explore the relationship between these isoforms and metabolic profiles in the arms (triceps brachii) and legs (vastus lateralis) as well as the range of training responses in the muscle fibers of elite cross-country skiers with equally and exceptionally well-trained upper and lower bodies. The proportion of myosin heavy chain (MHC)-1 was higher in the leg (58±2% [34-69%]) than arm (40±3% [24-57%], although the mitochondrial volume-percentages (8.6±1.6 (leg) and 9.0±2.0 (arm)), and average number of capillaries per fiber (5.8±0.8 (leg) and 6.3±0.3 (arm)) were the same. In these comparable highly-trained leg and arm muscles, the maximal citrate synthase activity was the same. Still, 3-hydroxy-acyl-CoA-dehydrogenase (HAD) capacity was 52% higher (P<0.05) in the leg compared to arm muscles, suggesting a relatively higher capacity for lipid oxidation in leg muscle, which cannot be explained by the different fiber type distributions. For both limbs combined, HAD activity was correlated with the content of MHC-1 (r2= 0.32, P = 0.011), whereas citrate synthase activity was not. Thus, in these highly trained cross-country skiers capillarization of and mitochondrial volume in type 2 fiber can be at least as high as in type 1 fibers, indicating a divergence between fiber type pattern and aerobic metabolic capacity. The considerable variability in oxidative metabolism with similar MHC profiles provides a new perspective on exercise training. Furthermore, the clear differences between equally well-trained arm and leg muscles regarding HAD activity cannot be explained by training status or MHC distribution, thereby indicating an intrinsic metabolic difference between the upper and lower body. Moreover, trained type 1 and type 2A muscle fibers exhibited similar aerobic capacity regardless of whether they were located in an arm or leg muscle.
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Elite cross-country skiers, characterized by equally trained upper- and lower-body muscles, exhibit similar mitochondrial volume fractions, capillarization, and citrate synthase activity, despite a higher proportion of MHCII fibers in the arms.
“Elite cross‐country skiers, who are characterized by equally trained upper‐ and lower‐body, arm and leg muscles, have been shown to exhibit similar mitochondrial volume fractions, capillarization, and citrate synthase activity, despite a higher proportion of MHCII fibers in the arms [30].”
From Carbohydrate Storage and Supplementation Strategies for Peak Performance in Cross-Country Skiing (2026) · cites this paper as [30]
The greater proportion of type II fibers in upper body musculature might account for the improved bench press and inconsistent effects on squat performance after NO3− supplementation.
“Given that there is evidence to suggest that NO3− supplementation may be more effective at enhancing physiological responses in type II muscle fibers [124] and since the proportion of type II muscle fibers may be greater in the upper body musculature, i.e., [125], this might account for the improved bench press and the inconsistent effects observed on squat performance after NO3− supplementation.”
From Effects of Dietary Nitrate Supplementation on Weightlifting Exercise Performance in Healthy Adults: A Systematic Review (2020) · cites this paper as [125]
The proportion of type II muscle fibers may be greater in the upper body musculature.
“Given that there is evidence to suggest that NO3− supplementation may be more effective at enhancing physiological responses in type II muscle fibers [124] and since the proportion of type II muscle fibers may be greater in the upper body musculature, i.e., [125], this might account for the improved bench press and the inconsistent effects observed on squat performance after NO3− supplementation.”
From Effects of Dietary Nitrate Supplementation on Weightlifting Exercise Performance in Healthy Adults: A Systematic Review (2020) · cites this paper as [125]
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- Authors
- Ørtenblad, N., Nielsen, J., Boushel, R., Söderlund, K., Saltin, B., & Holmberg, H.-C.
- Affiliations
- University of British Columbia (CA)
- University of Southern Denmark (DK)
- Swedish School of Sport and Health Sciences (SE)
- University of Copenhagen (DK)
- Mid Sweden University (SE)
- Corresponding author
- Ørtenblad, Niels
- Venue
- Frontiers in Physiology
- Publisher
- Frontiers Media SA
- Published
- 2018-08-02
- Volume
- 9
- Pages
- 1031-1031
- ISSN
- 1664-042X (ISSN-L)
- PMC
- PMC6084043
- OpenAlex
- W2883867909
- Cited by
- 102
- References
- 38
- Citations by year
- 2026: 7 · 2025: 9 · 2024: 11 · 2023: 15 · 2022: 21 · 2021: 19 · 2020: 8 · 2019: 12
- FWCI
- 13.38
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Locations
- Frontiers in Physiology (publishedVersion, cc-by)
- Europe PMC (PubMed Central) (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- University of Southern Denmark Research Portal (University of Southern Denmark) (acceptedVersion, cc-by)
- Diva portal (Dalarna University Library) (submittedVersion, other-oa)
- KTH Publication Database DiVA (KTH Royal Institute of Technology) (submittedVersion, other-oa)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
- NORA - Norwegian Open Research Archives (submittedVersion, cc-by-nc-nd)
- Funders
- Kulturministeriet
- Topics
- Health Sciences › Medicine › Complementary and alternative medicine › Cardiovascular and exercise physiology
- Health Sciences › Medicine › Orthopedics and Sports Medicine › Sports Performance and Training
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Cell Biology › Muscle metabolism and nutrition
- Keywords
- cross-country skiing, myosin heavy chain isoforms, muscle fiber type distribution, mitochondrial content, capillarization, citrate synthase, 3-hydroxyacyl-CoA dehydrogenase, lipid oxidation, cardiorespiratory fitness, triceps brachii, vastus lateralis, elite athletes, exercise training response, fast-twitch muscle
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