Cell culture studies have shown that amino acids activate mTORC1 signaling by increasing the interaction between mTOR and its essential activator Rheb. However, the existence of this mechanism in human skeletal muscle remains to be determined.

PURPOSE: To determine if increased mTORC1 signaling in response to amino acids in human skeletal muscle is due to an increased interaction between mTOR and Rheb.

METHODS: Eight well trained men performed resistance exercise on two separate occasions. In connection with the exercise, subjects were supplemented with flavored water (Pla) and essential amino acids (EAA) in a double-blind, randomized cross-over design. Muscle biopsies were taken in the vastus lateralis muscle before, immediately after and 90 and 180 min post exercise. Activity of the mTORC1 pathway was assessed by a radiolabeled in-vitro kinase assay for its immediate downstream target S6K1. Protein-protein interactions were determined by western blot following co-immunoprecipitation of mTOR with Rheb. Co-immunoprecipitation was performed on pooled muscle samples from three of the eight subjects.

RESULTS: Activity of S6K1 remained unchanged immediately after exercise in both trials. However, at 90 min post exercise, S6K1 activity increased by approximately 2- and 8-fold (p<0.05) from baseline the Pla and EAA trials, respectively. At the 180 min time point, S6K1 activity remained elevated in both trials being approx. 3-fold higher in the Pla trial and 5-fold higher (p<0.05) in the EAA trial. The fold-change in mTOR and Rheb interaction largely resembled the activity pattern of S6K1 in both trials; in the Pla trial the fold-change was 0.9, 1.3 and 1.4 while in the EAA trial the fold-change was 1.6, 2.9 and 1.9 immediately after, 90 min after and 180 min after exercise, respectively.

CONCLUSIONS: The large increase in S6K1 activity following EAA intake appears to be mediated by an increased interaction between mTOR and its proximal activator Rheb. This is the first time this mechanism has been demonstrated in human skeletal muscle.

Biomechanically Influenced Differences in O-2 Extraction in Diagonal Skiing: Arm versus Leg. Med. Sci. Sports Exerc., Vol. 42, No. 10, pp. 1899-1908, 2010. Purpose: This study aimed to determine whether the differences in oxygen extraction and lactate concentration in arms and legs during cross-country skiing are related to muscle activation or force production and how these differences are influenced by a reduction in exercise intensity. Methods: Nine well-trained male cross-country skiers (age = 22 +/- 3 yr, (V) over dotO(2max) = 5.3 +/- 0.3 L.min(-1) and 69 +/- 3 mL.kg(-1).min(-1)) performed diagonal skiing on a treadmill for 3 min at 90% followed by 6 min at 70% of (V) over dotO(2max). During the final minute of each workload, arterial, femoral, and subclavian venous blood was collected for determination of blood gases, pH, and lactate. EMG was recorded from six upper-and lower-body muscles, and leg and pole forces were measured. Cardiorespiratory variables were monitored continuously. Results: Oxygen extraction in the legs was higher than that in the arms at both 90% and 70% of (V) over dotO(2max) (92% +/- 3% vs 85% +/- 6%, P < 0.05 and 90% +/- 3% vs 78% +/- 8%, P < 0.001). This reduction with decreased workload was more pronounced in the arms (-9.8% +/- 7.7% vs -3.2% +/- 3.2%, P < 0.01). EMGRMS for the arms was higher, and pole ground contact time was greater than the corresponding values for the legs (both P < 0.01). At both intensities, the blood lactate concentration was higher in the subclavian than that in the femoral vein but was lowered more in the subclavian vein when intensity was reduced (all P < 0.001). Conclusions: The higher muscle activation (percentage of maximal voluntary isometric contraction) in the arms and the longer ground contact time of the poles than the legs contribute to the lower oxygen extraction and elevated blood lactate concentration in the arms in diagonal skiing. The better lactate recovery in the arms than that in the legs is aided by greater reductions in muscle activation and pole force when exercise intensity is reduced.

Purpose This study aims to investigate fluctuations in total mechanical energy of the body (E-body) in relation to external ergometer work (W-erg) during the poling and recovery phases of simulated double-poling cross-country skiing. Methods Nine male cross-country skiers (mean SD age, 24 5 yr; mean +/- SD body mass, 81.7 +/- 6.5 kg) performed 4-min submaximal tests at low-intensity, moderate-intensity, and high-intensity levels and a 3-min all-out test on a ski ergometer. Motion capture analysis and load cell recordings were used to measure body kinematics and dynamics. From these, W-erg, E-body (sum of the translational, rotational, and gravitational potential energies of all segments), and their time differentials (power P) were calculated. P(tot)the rate of energy absorption or generation by muscles-tendonswas defined as the sum of P-body and P-erg. ResultsE(body) showed large fluctuations over the movement cycle, decreasing during poling and increasing during the recovery phase. The fluctuation in P-body was almost perfectly out of phase with P-erg. Some muscle-tendon energy absorption was observed at the onset of poling. For the rest of poling and throughout the recovery phase, muscles-tendons generated energy to do W-erg and to increase E-body. Approximately 50% of cycle P-tot occurred during recovery for all intensity levels. Conclusions In double poling, the extensive contribution of the lower extremities and trunk to whole-body muscle-tendon work during recovery facilitates a direct transfer of E-body to W-erg during the poling phase. This observation reveals that double poling involves a unique movement pattern different from most other forms of legged terrestrial locomotion, which are characterized primarily by inverted pendulum or spring-mass types of movement.

Several investigators have demonstrated that chronic eccentric leg cycling is an effective method for improving lower body neuromuscular function (e.g., quadriceps muscle size, strength, and mobility) in a variety of patient and athletic populations. To date, there are no reports of using eccentricarm cycling (ECarm) as an exercise modality, probably in large part because of the lack of commercially available ECarm ergometers. Purpose: Our purposes for conducting this study were to 1) describe the design and construction of an ECarm ergometer and 2) compare ECarm to traditional concentric arm cycling (CCarm). Methods: All of the parts of a Monark 891E cycle ergometer (Monark Exercise AB, Vansbro, Sweden) were removed, leaving the frame and flywheel. An electric motor (2.2 kW) was connected to the flywheel via a pulley and a belt. Motor speed and pedaling rate were controlled by a variable frequency drive. A power meter quantified power and pedaling rate, and provided feedback to the individual. Eight individuals performed 3-min ECarm and CCarm trials at 40, 80, and 120 W (60 rpm) while V̇O₂ was measured. Results: The ECarm ergometer was simple to use, was adjustable, provided feedback on power output to the user, and allowed for a range of eccentric powers. V̇O₂ during ECarm was substantially lower compared with CCarm (P < 0.001). At similar V̇O₂ (0.97 ± 0.18 vs 0.91 ± 0.09 L•min, for ECarm and CCarm, respectively, P = 0.26), power absorbed during ECarm was approximately threefold greater than that produced during CCarm (118 ± 1 vs 40 ± 1 W, P < 0.001). Conclusion: This novel ECarm ergometer can be used to perform repetitive, high-force, multijoint, eccentric actions with the upper body at a low level of metabolic demand and may allow researchers and clinicians to use ECarm as a training and rehabilitation modality. © 2012 by the American College of Sports Medicine.

Purpose: Repeated-sprint training in hypoxia (RSH) was recently shown to improve repeated-sprint ability (RSA) in cycling. This phenomenon is likely to reflect fiber type-dependent, compensatory vasodilation, and therefore, our hypothesis was that RSH is even more beneficial for activities involving upper body muscles, such as double poling during cross-country skiing. Methods: In a double-blinded fashion, 17 competitive cross-country skiers performed six sessions of repeated sprints (each consisting of four sets of five 10-s sprints, with 20-s intervals of recovery) either in normoxia (RSN, 300 m; FiO2, 20.9%; n = 8) or normobaric hypoxia (RSH, 3000 m; FiO2, 13.8 %; n = 9). Before (pre) and after (post) training, performance was evaluated with an RSA test (10-s all-out sprints-20-s recovery, until peak power output declined by 30%) and a simulated team sprint (team sprint, 3×3-min all-out with 3-min rest) on a double-poling ergometer. Triceps brachii oxygenation was measured by near-infrared spectroscopy. Results: From pretraining to posttraining, peak power output in the RSA was increased (P < 0.01) to the same extent (29% ± 13% vs 26% ± 18%, nonsignificant) in RSH and in RSN whereas the number of sprints performed was enhanced in RSH (10.9 ± 5.2 vs 17.1 ± 6.8, P < 0.01) but not in RSN (11.6 T 5.3 vs 11.7 ± 4.3, nonsignificant). In addition, the amplitude in total hemoglobin variations during sprints throughout RSA rose more in RSH (P < 0.01). Similarly, the average power output during all team sprints improved by 11% T 9% in RSH and 15% T 7% in RSN. Conclusions: Our findings reveal greater improvement in the performance of repeated double-poling sprints, together with larger variations in the perfusion of upper body muscles in RSH compared with those in RSN. © 2014 by the American College of Sports Medicine.

Purpose The present study investigated the effects of periodic carbohydrate (CHO) restriction on endurance performance and metabolic markers in elite endurance athletes. Methods Twenty-six male elite endurance athletes (maximal oxygen consumption (VO2max), 65.0 mL O(2)kg(-1)min(-1)) completed 4 wk of regular endurance training while being matched and randomized into two groups training with (low) or without (high) CHO manipulation 3 dwk(-1). The CHO manipulation days consisted of a 1-h high-intensity bike session in the morning, recovery for 7 h while consuming isocaloric diets containing either high CHO (414 2.4 g) or low CHO (79.5 1.0 g), and a 2-h moderate bike session in the afternoon with or without CHO. VO2max, maximal fat oxidation, and power output during a 30-min time trial (TT) were determined before and after the training period. The TT was undertaken after 90 min of intermittent exercise with CHO provision before the training period and both CHO and placebo after the training period. Muscle biopsies were analyzed for glycogen, citrate synthase (CS) and -hydroxyacyl-coenzyme A dehydrogenase (HAD) activity, carnitine palmitoyltransferase (CPT1b), and phosphorylated acetyl-CoA carboxylase (pACC). Results The training effects were similar in both groups for all parameters. On average, VO2max and power output during the 30-min TT increased by 5% +/- 1% (P < 0.05) and TT performance was similar after CHO and placebo during the preload phase. Training promoted overall increases in glycogen content (18% +/- 5%), CS activity (11% +/- 5%), and pACC (38% +/- 19%; P < 0.05) with no differences between groups. HAD activity and CPT1b protein content remained unchanged. Conclusions Superimposing periodic CHO restriction to 4 wk of regular endurance training had no superior effects on performance and muscle adaptations in elite endurance athletes.

PURPOSE: In the classical style of cross-country skiing, the double-poling (DP) technique, which is regarded as an upper-body exercise, is used on the flatter parts of a course. Limited biomechanical and physiological data are available about DP compared with other cross-country skiing techniques. The purpose of the present study was to evaluate the possible role of the lower body during DP. METHODS: Eleven elite cross-country skiers performed two incremental tests using DP roller skiing at 1 degree inclination on a treadmill with or without locking the knee and ankle joints (DPLOCKED and DPFREE). Maximal and peak oxygen uptake (VO2max and VO2peak) during classic diagonal skiing and DP, respectively, were measured. In addition, heart rate, blood lactate concentration, and maximal DP velocity (Vmax) were determined. Pole-ground reaction forces and joint angles (elbow, hip, knee, and ankle) were analyzed. RESULTS: The skiers obtained 7.7% higher VO2peak, 9.4% higher Vmax, and 11.7% longer time to exhaustion during DPFREE compared with DPLOCKED (all P < 0.05). There was a higher heart rate and blood lactate concentration in DPLOCKED at submaximal stages (all P < 0.05), with no difference in oxygen consumption. At 85% Vmax, corresponding to approximately 81% VO2peak FREE, the differences in physiological variables were accompanied by a 13.6% higher poling frequency, a 4.9% shorter poling phase, 13.3% shorter recovery phase, and 10.9% lower relative pole force in DPLOCKED (all P < 0.05). CONCLUSIONS: Movements of the knee and ankle joints are an integrative part in the skillful use of the DP technique, and restriction of the motion in these joints markedly affects both biomechanical and physiological variables, impairing DP performance.

PURPOSE: To further the understanding of double poling (DP) through biomechanical analysis of upper and lower body movements during DP in cross-country (XC) skiing at racing speed. METHODS: Eleven elite XC skiers performed DP at 85% of their maximal DP velocity (V85%) during roller skiing at 1 degrees inclination on a treadmill. Pole and plantar ground reaction forces, joint angles (elbow, hip, knee, and ankle), cycle characteristics, and electromyography (EMG) of upper and lower body muscles were analyzed. RESULTS: 1) Pole force pattern with initial impact force peak and the following active force peak (PPF) correlated to V85%, (r = 0.66, P < 0.05); 2) active flexion-extension pattern in elbow, hip, knee, and ankle joints with angle minima occurring around PPF, correlated to hip angle at pole plant (r = -0.89, P < 0.01), minimum elbow angle (r = -0.71), and relative poling time (r = -0.72, P < 0.05); 3) two different DP strategies (A and B), where strategy A (best skiers) was characterized by higher angular elbow- and hip-flexion velocities, smaller minimum elbow (P < 0.01) and hip angles (P < 0.05), and higher PPF (P < 0.05); 4) EMG activity in trunk and hip flexors, shoulder, and elbow extensors, and several lower body muscles followed a specific sequential pattern with changing activation levels; and 5) EMG activity in lower body muscles showed DP requires more than upper body work. CONCLUSIONS: DP was found to be a complex movement involving both the upper and lower body showing different strategies concerning several biomechanical aspects. Future research should further investigate the relationship between biomechanical and physiological variables and elaborate training models to improve DP performance.

PURPOSE: Orienteering athletes must adapt to running on various surfaces, with biomechanics likely contributing to performance. Here, our aims were to identify the effect of athletic status and of surface on the running biomechanics of orienteers. METHODS: Seven elite and seven amateur male orienteers ran 20 m on road, path, and forest surfaces at maximal, 3.8 m·s, and 85% of maximal speeds. A three-dimensional motion capturing system monitored temporal gait and lower extremity kinematic parameters. Data were analyzed using mixed effects models that considered surface (road-path-forest), group (elite-amateur), and surface-group interaction effects. RESULTS: Forest running at maximal speed was slower and involved longer step and cycle times, greater knee extension at foot strike, smaller peak hip flexion and dorsiflexion during stance, and increased ranges of vertical pelvis motion compared with those observed on the road. Elites specifically exhibited greater hip extension at foot strike, larger dorsiflexion at toe-off, and lower pelvis at foot strike and toe-off, whereas amateurs displayed longer stance, greater plantarflexion at foot strike, and greater knee with lesser ankle motion. At the slowest speed, subjects exhibited greater knee flexion at foot strike, greater dorsiflexion at toe-off, shorter strides, smaller peak dorsiflexion during stance, and greater hip, knee, and vertical pelvis motions on forest than on road surfaces. Elites specifically demonstrated shorter stance, step, and cycle times whereas amateurs did not. CONCLUSIONS: Orienteering athletes adjusted their running biomechanics when off-road, with distinct adaptations observed in elite versus amateur competitors. The vertical pelvis motion was consistently greater when running off-road, coherent with reported increases in energy expenditure. However, our athletes did not exhibit more crouched lower limb postures when sprinting in the forest, indicating alternative responses to off-road running to that previously proposed by "Groucho" running.

PURPOSE: Athletes who use anabolic steroids get larger and stronger muscles. How this is reflected at the level of the muscle fibers has not yet been established and was the topic of this investigation. METHODS: Muscle biopsies were obtained from the trapezius muscles of high-level power lifters who have reported the use of anabolic steroids in high doses for several years and from high-level power lifters who have never used these drugs. Enzyme-immunohistochemical investigation was performed to assess muscle fiber types, fiber area, myonuclear number, frequency of satellite cells, and fibers expressing developmental protein isoforms. RESULTS: The overall muscle fiber composition was the same in both groups. The mean area for each fiber type in the reported steroid users was larger than that in the nonsteroid users (P < 0.05). The number of myonuclei and the proportion of central nuclei were also significantly higher in the reported steroid users (P < 0.05). Likewise, the frequency of fibers expressing developmental protein isoforms was significantly higher in the reported steroid users group (P < 0.05). CONCLUSION: Intake of anabolic steroids and strength-training induce an increase in muscle size by both hypertrophy and the formation of new muscle fibers. We propose that activation of satellite cells is a key process and is enhanced by the steroid use. The incorporation of the satellite cells into preexisting fibers to maintain a constant nuclear to cytoplasmic ratio seems to be a fundamental mechanism for muscle fiber growth. Although all the subjects in this study have the same level of performance, the possibility of genetic differences between the two groups cannot be completely excluded.

PURPOSE: Double poling (DP) as a main technique in cross-country skiing has developed substantially over the last 15 yr. The purpose of the present study was to analyze the question, "How do modern elite skiers control DP speed?" METHODS: Twelve male elite cross-country skiers roller skied using DP at 9, 15, 21, and 27 km.h(-1) and maximum velocity (V(max)). Cycle characteristics, pole and plantar forces, and elbow, hip, and knee joint angles were analyzed. RESULT: Both poling frequency and cycle length increased up to 27 km.h (-1)(P < 0.05), with a further increase in poling frequency at V(max) (P < 0.05). Peak pole force, rate of force development, and rearfoot plantar force increased with submaximal velocities (V(sm)), whereas poling time and time-to-peak pole force gradually shortened (P < 0.05). Changes in elbow joint kinematics during the poling phase were characterized by a decreased angle minimum and an increased flexion and extension ranges of motion as well as angular velocities across V(sm) (P < 0.05), with no further changes at V(max). Hip and knee joint kinematics adapted across V(sm) by 1) decreasing angles at pole plant and angle minima during the poling phase, 2) increasing the ranges of motion and angular velocities during the flexion phases occurring around pole plant, and 3) increasing extension ranges of motion and angular velocities during the recovery phase (all P values <0.05), with no further changes at V(max). CONCLUSIONS: Elite skiers control DP speed by increasing both poling frequency and cycle length; the latter is achieved by increased pole force despite reduced poling time. Adaptation to higher speeds was assisted by an increased range of motion, smaller angle minima, and higher angular velocities in the elbow, the hip, and the knee joints.

Introduction: In light of the recent revolutionary change in the use of the doublepoling (DP) technique in cross-country skiing, our purpose was to compare the associated kinetics and kinematics on flat (DPflat) and uphill terrain (DPup), as well as to identify factors that determine performance. Methods: Thirteen elite male cross-country skiers completed two incremental speed tests (V-peak) involving roller skiing with the DP technique at moderate (13 and 24 km.h(-1)) and high speed (15 and 28.5 km.h(-1)) on a treadmill that was flat (1 degrees) or tilted uphill (7 degrees). Pole forces and three-dimensional whole-body kinematics were monitored simultaneously. Results: In comparison to DPflat, during DPup, swing times were much shorter (-48%) and peak pole forces greater (+13%) and generated later during the poling phase (+68%), with higher impulses for all force components (+87%-123%). Furthermore, pole forces were 18% more effectively oriented for propulsion. During DPup, the skiers demonstrated more flexed elbows, as well as shoulder angles that were less flexed in the forward direction and less abducted throughout the poling phase, together with more highly flexed knee and ankle joints, a more upright thorax, less flexed hips, and a shortened backward swing after pole off. With DPup, the skiers raised their center of mass 25% more, attaining maximal heel raise and maximal vertical position at a timepoint closer to pole plant compared with flat. On the uphill incline, the magnitude of V-peak was positively related to body mass, relative pole length (% body height), and magnitude of heel raise. Conclusions: The present findings provide novel insights into the coordination, kinetics and kinematics of elite skiers while DP on flat and uphill terrain.

Purpose: This study aimed to assess the effect of carrying a rifle on the physiological and biomechanical responses of well-trained biathletes. Methods: Ten elite biathletes (five men and five women) performed ski skating with (R) or without a rifle (NR) on a treadmill using the V2 (5- incline) and V1 techniques (8-) at 8 and 6 kmIhj1, respectively, as well as at racing intensity (approximately 95% of peak oxygen uptake (VO2peak), 10.7 T 0.8 and 7.7 T 0.9 kmIhj1, respectively). VO2, ventilation (VE), HR, blood lactate concentration (BLa), and cycle characteristics as well as pole and leg kinetics were evaluated during these trials. Results: Metabolic data were all higher for R than for NR, as follows:VO2, +2.5%;VE, +8.1%; RER, +4.2%; all P G 0.001; HR, +1.7%; and BLa, +15.1%; both P G 0.05. Biomechanically, carrying a rifle reduced cycle time and length, poling and arm swing times, and leg ground contact time and increased cycle rate, the peak and impulse of leg force, average cycle force, and impulse of forefoot force (all P G 0.05). With the exception of elevated pole forces when V2 skating at racing velocity, there were no differences between the peak and impulse of pole force. The difference inVE between R and NR was greater for the women than that for men (P G 0.05), and the difference in BLa also tended to be larger for the women (P G 0.1). Conclusions: Carrying a rifle elevated physiological responses, accelerated cycle rate, and involved greater leg work, with no differences between the V1 and V2 techniques.

Purpose: To describe the detailed kinetics and kinematics associated with use of the V1 skating technique at high skiing speeds and to identify factors that predict performance. Methods: Fifteen elite male cross-country skiers performed an incremental roller-skiing speed test (V-peak) on a treadmill using the V1 skating technique. Pole and plantar forces and whole-body kinematics were monitored at four submaximal speeds. Results: The propulsive force of the "strong side'' pole was greater than that of the "weak side'' (P < 0.01), but no difference was observed for the legs. The poles generated approximately 44% of the total propulsion, being more effective than the legs in this respect (similar to 59% vs 11%, P < 0.001). Faster skiers exhibited more well-synchronized poling, exhibited more symmetric edging by and forces from the legs, and were more effective in transformation of resultant forces into propulsion. Cycle length was not correlated with either V-peak or the impulse of total propulsive forces. Conclusions: The present findings provide novel insights into the coordination, kinetics, and kinematics of the arm and leg motion by elite athletes while V1 skating at high speeds. The faster skiers exhibit more symmetric leg motion on the "strong'' and "weak'' sides, as well as more synchronized poling. With respect to methods, the pressure insoles and three-dimensional kinematics in combination with the leg push-off model described here can easily be applied to all skating techniques, aiding in the evaluation of skiing techniques and comparison of effectiveness.

PURPOSE: In cross-country (XC) skiing, the V1 and V2 alternate skate techniques are asymmetric, and skiers can choose either the right or left side for pole support. The overall purpose of this study was to investigate V1 side preference in elite XC skiers, notably by documenting V1 skate side preference, dominant and nondominant V1peak speeds, left- to right-side differences (ΔL-R) in laboratory-based measurements, and relationships between side preference data. METHODS: Sixteen male elite XC skiers completed one incremental speed test using V1 on their dominant side and another incremental speed test using V1 on their nondominant side while roller-skiing on a treadmill. During these tests, V1peak speed, pole forces, and plantar forces were measured. A whole-body dual-energy x-ray absortiometry (DXA) scan measured anthropometric parameters and questionnaires established side preference for V2 alternate, overall laterality in XC skiing, handedness, footedness, and injury prevalence. RESULTS: Left-to-right V1 side preference was equally distributed among skiers. V1peak speed was approximately 4.5% greater on the dominant versus nondominant sides. V1peak Δ L-R were positively related to Δ L-R in V1-dominant peak pole forces only. Questionnaire data indicated that more skiers preferred V2 alternate right, with moderate correlations between preferred V1 and V2 alternate sides. The expression of a dominant side in V1 and V2 alternate increased as skiing speed increased from moderate to 15-km endurance-race to sprint-race speeds. However, no relationships were established between V1 or V2 side preference and handedness, footedness, or number of one-sided injuries. CONCLUSIONS: ΔL-R in measurements provide limited explanations for V1 side preferences in elite XC skiers. In fact, no systematic relations exist between V1 side preferences and anthropometric, biomechanical, or questionnaire data. Copyright © 2013 by the American College of Sports Medicine.