Muscle fibre typology affects whole-body metabolic rate during isolated muscle contractions and human locomotion.

The wide variation in muscle fibre type distribution across individuals, along with the very different energy consumption rates in slow versus fast muscle fibres, suggests that muscle fibre typology contributes to inter-individual differences in metabolic rate during exercise. However, this has been hard to demonstrate due to the gap between a single muscle fibre and full-body exercises. We investigated the isolated effect of triceps surae muscle contraction velocity on whole-body metabolic rate during cyclic contractions in individuals a priori selected for their predominantly slow (n = 11) or fast (n = 10) muscle fibre typology by means of proton magnetic resonance spectroscopy (1H-MRS). Subsequently, we examined their whole-body metabolic rate during walking and running at 2 m/s, exercises with comparable metabolic rates but distinct triceps surae muscle force and velocity demands (walking: low force, high velocity; running: high force, low velocity). Increasing triceps surae contraction velocity during cyclic contractions elevated net whole-body metabolic rate for both typology groups. However, the slow group consumed substantially less net metabolic energy at the slowest contraction velocity, but the metabolic difference between groups diminished at faster velocities. Consistent with the more economic force production during slow contractions, the slow group exhibited lower metabolic rates than the fast group while running, whereas metabolic rates were similar during walking. These findings provide important insights into the influence of muscle fibre typology on whole-body metabolic rate and emphasize the importance of considering muscle mechanical demands to understand muscle fibre typology related differences in whole-body metabolic rates. KEY POINTS: Muscle fibre typology is often suggested to affect whole-body metabolic rate, yet convincing in vivo evidence is lacking. Using isolated plantar flexor muscle contractions in individuals a priori selected for their predominantly slow or fast muscle fibre typology, we demonstrated that having predominantly slow muscle fibres provides a metabolic advantage during slow muscle contractions, but this benefit disappeared at faster contractions. We extended these results to full-body exercises, where we demonstrated that higher proportions of slow fibres associated with better economy during running but not when walking. These findings provide important insights into the influence of muscle fibre typology on whole-body metabolic rate and emphasize the importance of considering muscle mechanical demands to understand muscle fibre typology related differences in whole-body metabolic rate.

Faster triceps surae muscle cyclic contractions alter muscle activity and whole body metabolic rate.

Hundred years ago, Fenn demonstrated that when a muscle shortens faster, its energy liberation increases. Fenn's results were the first of many that led to the general understanding that isometric muscle contractions are energetically cheaper than concentric contractions. However, this evidence is still primarily based on single fiber or isolated (ex vivo) muscle studies and it remains unknown whether this translates to whole body metabolic rate. In this study, we specifically changed the contraction velocity of the ankle plantar flexors and quantified the effects on triceps surae muscle activity and whole body metabolic rate during cyclic plantar flexion (PF) contractions. Fifteen participants performed submaximal ankle plantar flexions (∼1/3 s activation and ∼2/3 s relaxation) on a dynamometer at three different ankle angular velocities: isometric (10° PF), isokinetic at 30°/s (5-15° PF), and isokinetic at 60°/s (0-20° PF) while target torque (25% MVC) and cycle frequency were kept constant. In addition, to directly determine the effect of ankle angular velocity on muscle kinematics we collected gastrocnemius medialis muscle fascicle ultrasound data. As expected, increasing ankle angular velocity increased gastrocnemius medialis muscle fascicle contraction velocity and positive mechanical work (P < 0.01), increased mean and peak triceps surae muscle activity (P < 0.01), and considerably increased net whole body metabolic rate (P < 0.01). Interestingly, the increase in triceps surae muscle activity with fast ankle angular velocities was most pronounced in the gastrocnemius lateralis (P < 0.05). Overall, our results support the original findings from Fenn in 1923 and we demonstrated that greater triceps surae muscle contraction velocities translate to increased whole body metabolic rate.NEW & NOTEWORTHY Single muscle fiber studies or research on isolated (ex vivo) muscles demonstrated that faster concentric muscle contractions yield increased energy consumption. Here we translated this knowledge to muscle activation and whole body metabolic rate. Increasing ankle angular velocity increased triceps surae contraction velocity and mechanical work, increasing triceps surae muscle activity and substantially elevating whole body metabolic rate. Additionally, we demonstrated that triceps surae muscle activation strategy depends on the mechanical demands of the task.

Everesting: cycling the elevation of the tallest mountain on Earth.

PURPOSE: With few cycling races on the calendar in 2020 due to COVID-19, Everesting became a popular challenge: you select one hill and cycle up and down it until you reach the accumulated elevation of Mt. Everest (8,848 m or 29,029ft). With an almost infinite number of different hills across the world, the question arises what the optimal hill for Everesting would be. Here, we address the biomechanics and energetics of up- and downhill cycling to determine the characteristics of this optimal hill. METHODS: During uphill cycling, the mechanical power output equals the power necessary to overcome air resistance, rolling resistance, and work against gravity, and for a fast Everesting time, one should maximize this latter term. To determine the optimal section length (i.e., number of repetitions), we applied the critical power concept and assumed that the U-turn associated with an additional repetition comes with a 6 s time penalty. RESULTS: To use most mechanical power to overcoming gravity, slopes of at least 12% are most suitable, especially since gross efficiency seems only minimally diminished on steeper slopes. Next, we found 24 repetitions to be optimal, yet this number slightly depends on the assumptions made. Finally, we discuss other factors (fueling, altitude, fatigue) not incorporated in the model but also affecting Everesting performances. CONCLUSION: For a fast Everesting time, our model suggests to select a hill climb which preferably starts at (or close to) sea level, with a slope of 12-20% and length of 2-3 km.

Triceps surae muscle force potential and force demand shift with altering stride frequency in running.

While it is well recognized that the preferred stride frequency (PSF) in running closely corresponds to the metabolically optimal frequency, the underlying mechanisms are still unclear. Changes in joint kinematics when altering stride frequency will affect the muscle-tendon unit lengths and potentially the efficiency of muscles crossing these joints. Here, we investigated how fascicle kinematics and forces of the triceps surae muscle, a highly energy consuming muscle, are affected when running at different stride frequencies. Twelve runners ran on a force measuring treadmill, adopting five different frequencies (PSF; PSF ± 8%; PSF ± 15%), while we measured joint kinematics, whole-body energy expenditure, triceps surae muscle activity, and soleus (SOL; N = 10) and gastrocnemius medialis (GM; N = 12) fascicle kinematics. In addition, we used dynamic optimization to estimate SOL and GM muscle forces. We found that SOL and GM mean muscle fascicle length during stance followed an inverted U-relationship with the longest fascicle lengths occurring at PSF. Fascicle lengths were shortest at frequencies lower than PSF. In addition, average SOL force was greater at PSF-15% compared with PSF. Overall, our results suggest that reduced SOL and GM muscle fascicle lengths, associated with reduced muscle force potential, together with greater SOL force demand, contribute to the increased whole-body energy expenditure when running at lower than PSF. At higher stride frequencies, triceps surae muscle kinematics and force production were less affected suggesting that increased energy expenditure is rather related to higher cost of leg swing and greater cost of force production.

Changing Stride Frequency Alters Average Joint Power and Power Distributions during Ground Contact and Leg Swing in Running.

PURPOSE: Runners naturally adopt a stride frequency closely corresponding with the stride frequency that minimizes energy consumption. Although the concept of self-optimization is well recognized, we lack mechanistic insight into the association between stride frequency and energy consumption. Altering stride frequency affects lower extremity joint power; however, these alterations are different between joints, possibly with counteracting effects on the energy consumption during ground contact and swing. Here, we investigated the effects of changing stride frequency from a joint-level perspective. METHODS: Seventeen experienced runners performed six running trials at five different stride frequencies (preferred stride frequency (PSF) twice, PSF ± 8%, PSF ± 15%) at 12 km·h-1. During each trial, we measured metabolic energy consumption and muscle activation, and collected kinematic and kinetic data, which allowed us to calculate average positive joint power using inverse dynamics. RESULTS: With decreasing stride frequency, average positive ankle and knee power during ground contact increased (P < 0.01), whereas average positive hip power during leg swing decreased (P < 0.01). Average soleus muscle activation during ground contact also decreased with increasing stride frequency (P < 0.01). In addition, the relative contribution of positive ankle power to the total positive joint power during ground contact decreased (P = 0.01) with decreasing stride frequency, whereas the relative contribution of the hip during the full stride increased (P < 0.01) with increasing stride frequency. CONCLUSIONS: Our results provide evidence for the hypothesis that the optimal stride frequency represents a trade-off between minimizing the energy consumption during ground contact, associated with higher stride frequencies, without excessively increasing the cost of leg swing or reducing the time available to produce the necessary forces.

Energetics and Biomechanics of Running Footwear with Increased Longitudinal Bending Stiffness: A Narrative Review.

In the wake of the quest to break the 2-h marathon barrier, carbon-fiber plates have become commonplace in marathon racing shoes. Despite the controversy surrounding this shoe technology, studies on the effects of increased longitudinal bending stiffness on running economy report mixed results. Here, we provide a comprehensive review of the current literature on midsole bending stiffness and carbon-fiber plates in distance running shoes, focusing on how longitudinal bending stiffness affects running energetics and lower limb mechanics. The current literature reports changes in running economy with increased longitudinal bending stiffness ranging from ~ 3% deterioration to ~ 3% improvement. In some studies, larger improvements have been observed, but often those shoes varied in many aspects, not just longitudinal bending stiffness. Biomechanically, increased longitudinal bending stiffness has the largest impact on metatarsal-phalangeal (MTP) and ankle joint mechanics. Plate location [top loaded (an insole), embedded (in between midsole foam), and bottom loaded (along the bottom of the shoe)] and geometry (flat/curved) affect joint moments and angular velocities at the MTP and ankle joint differently, which partly explains the mixed running economy results. Further research investigating how carbon-fiber plates interact with other footwear features (such as foam and midsole geometry), scaling of those with shoe size, body mass, and strike pattern, and comparing various plate placements is needed to better understand how longitudinal bending stiffness affects running economy.

Habitual foot strike pattern does not affect simulated triceps surae muscle metabolic energy consumption during running.

Foot strike pattern affects ankle joint work and triceps surae muscle-tendon dynamics during running. Whether these changes in muscle-tendon dynamics also affect triceps surae muscle energy consumption is still unknown. In addition, as the triceps surae muscle accounts for a substantial amount of the whole-body metabolic energy consumption, changes in triceps surae energy consumption may affect whole-body metabolic energy consumption. However, direct measurements of muscle metabolic energy consumption during dynamic movements is difficult. Model-based approaches can be used to estimate individual muscle and whole-body metabolic energy consumption based on Hill type muscle models. In this study, we use an integrated experimental and dynamic optimization approach to compute muscle states (muscle forces, lengths, velocities, excitations and activations) of 10 habitual midfoot/forefoot striking and nine habitual rearfoot striking runners while running at 10 and 14 km h-1 The Achilles tendon stiffness of the musculoskeletal model was adapted to fit experimental ultrasound data of the gastrocnemius medialis muscle during ground contact. Next, we calculated triceps surae muscle and whole-body metabolic energy consumption using four different metabolic energy models provided in the literature. Neither triceps surae metabolic energy consumption (P>0.35) nor whole-body metabolic energy consumption (P>0.14) was different between foot strike patterns, regardless of the energy model used or running speed tested. Our results provide new evidence that midfoot/forefoot and rearfoot strike patterns are metabolically equivalent.

Effect of habitual foot-strike pattern on the gastrocnemius medialis muscle-tendon interaction and muscle force production during running.

The interaction between gastrocnemius medialis (GM) muscle and Achilles tendon, i.e., muscle-tendon unit (MTU) interaction, plays an important role in minimizing the metabolic cost of running. Foot-strike pattern (FSP) has been suggested to alter MTU interaction and subsequently the metabolic cost of running. However, metabolic data from experimental studies on FSP are inconsistent, and a comparison of MTU interaction between FSP is still lacking. We, therefore, investigated the effect of habitual rearfoot and mid-/forefoot striking on MTU interaction, ankle joint work, and plantar flexor muscle force production while running at 10 and 14 km/h. GM muscle fascicles of 9 rearfoot and 10 mid-/forefoot strikers were tracked using dynamic ultrasonography during treadmill running. We collected kinetic and kinematic data and used musculoskeletal models to determine joint angles and calculate MTU lengths. In addition, we used dynamic optimization to assess plantar flexor muscle forces. During ground contact, GM fascicle shortening ( P = 0.02) and average contraction velocity ( P = 0.01) were 40-45% greater in rearfoot strikers than mid-/forefoot strikers. Differences in contraction velocity were especially prominent during early ground contact. Moreover, GM ( P = 0.02) muscle force was greater during early ground contact in mid-/forefoot strikers than rearfoot strikers. Interestingly, we did not find differences in stretch or recoil of the series elastic element between FSP. Our results suggest that, for the GM, the reduced muscle energy cost associated with lower fascicle contraction velocity in mid-/forefoot strikers may be counteracted by greater muscle forces during early ground contact. NEW & NOTEWORTHY Kinetic and kinematic differences between foot-strike patterns during running imply (not previously reported) altered muscle-tendon interaction. Here, we studied muscle-tendon interaction using ultrasonography. We found greater fascicle contraction velocities and lower muscle forces in rearfoot compared with mid-/forefoot strikers. Our results suggest that the higher metabolic energy demand due to greater fascicle contraction velocities might offset the lower metabolic energy demand due to lower muscle forces in rearfoot compared with mid-/forefoot strikers.

Comparison of running and cycling economy in runners, cyclists, and triathletes.

PURPOSE: Exercise economy is one of the main physiological factors determining performance in endurance sports. Running economy (RE) can be improved with running-specific training, while the improvement of cycling economy (CE) with cycling-specific training is controversial. We investigated whether exercise economy reflects sport-specific skills/adaptations or is determined by overall physiological factors. METHODS: We compared RE and CE in 10 runners, 9 cyclists and 9 triathletes for running at 12 km/h and cycling at 200 W. Gross rates of oxygen consumption and carbon dioxide production were collected and used to calculate gross metabolic rate in watts for both running and cycling. RESULTS: Runners had better RE than cyclists (917 ± 107 W vs. 1111 ± 159 W) (p < 0.01). Triathletes had intermediate RE values (1004 ± 98 W) not different from runners or cyclists. CE was not different (p = 0.20) between the three groups (runners: 945 ± 60 W; cyclists: 982 ± 44 W; triathletes: 979 ± 54 W). CONCLUSION: RE can be enhanced with running-specific training, but CE is independent of cycling-specific training.

Endotoxemia shifts neutrophils with TIMP-free gelatinase B/MMP-9 from bone marrow to the periphery and induces systematic upregulation of TIMP-1.

Lipopolysaccharides or endotoxins elicit an excessive host inflammatory response and lead to life-threatening conditions such as endotoxemia and septic shock. Lipopolysaccharides trigger mobilization and stimulation of leukocytes and exaggerated production of pro-inflammatory molecules including cytokines and proteolytic enzymes. Matrix metalloproteinase-9 (MMP-9) or gelatinase B, a protease stored in the tertiary granules of polymorphonuclear leukocytes, has been implicated in such inflammatory reactions. Moreover, several studies even pinpointed MMP-9 as a potential target molecule to counter excessive inflammation in endotoxemia. Whereas the early effect of lipopolysaccharide-induced inflammation in vivo on the expression of MMP-9 in various peripheral organs has been described, the effects on the bone marrow and during late stage endotoxemia remain elusive. We demonstrate that TIMP-free MMP-9 is a major factor in bone marrow physiology and pathology. By using a mouse model for late-stage endotoxemia, we show that lipopolysaccharides elicited a depletion of neutrophil MMP-9 in the bone marrow and a shift of MMP-9 and MMP-9-containing cells towards peripheral organs, a pattern which was primarily associated with a relocation of CD11bhighGr-1high cells. In contrast, analysis of the tissue inhibitors of metalloproteinases was in line with a natural, systematic upregulation of TIMP-1, the main tissue inhibitor of TIMP-free MMP-9, and a general shift toward control of matrix metalloproteinase activity by tissue inhibitors of metalloproteinases.