Muscle architecture of the upper limb in the orangutan

We dissected the left upper limb of a female orangutan and systematically recorded muscle mass, fascicle length, and physiological cross-sectional area (PCSA), in order to quantitatively clarify the unique muscle architecture of the upper limb of the orangutan. Comparisons of the musculature of the dissected orangutan with corresponding published chimpanzee data demonstrated that in the orangutan, the elbow flexors, notably M. brachioradialis, tend to exhibit greater PCSAs. Moreover, the digital II-V flexors in the forearm, such as M. flexor digitorum superficialis and M. flexor digitorum profundus, tend to have smaller PCSA as a result of their relatively longer fascicles. Thus, in the orangutan, the elbow flexors demonstrate a higher potential for force production, whereas the forearm muscles allow a greater range of wrist joint mobility. The differences in the force-generating capacity in the upper limb muscles of the two species might reflect functional specialization of muscle architecture in the upper limb of the orangutan for living in arboreal environments.     

Effect of resistance training on skeletal muscle-specific force in elderly humans.

This study assessed muscle-specific force in vivo following strength training in old age. Subjects were assigned to training (n = 9, age 74.3 +/- 3.5 yr; mean +/- SD) and control (n = 9, age 67.1 +/- 2 yr) groups. Leg-extension and leg-press exercises (2 sets of 10 repetitions at 80% of the 5 repetition maximum) were performed three times/wk for 14 wk. Vastus lateralis (VL) muscle fascicle force was calculated from maximal isometric voluntary knee extensor torque with superimposed stimuli, accounting for the patella tendon moment arm length, ultrasound-based measurements of muscle architecture, and antagonist cocontraction estimated from electromyographic activity. Physiological cross-sectional area (PCSA) was calculated from the ratio of muscle volume to fascicle length. Specific force was calculated by dividing fascicle force by PCSA. Fascicle force increased by 11%, from 847.9 +/- 365.3 N before to 939.3 +/- 347.8 N after training (P < 0.05). Due to a relatively greater increase in fascicle length (11%) than muscle volume (6%), PCSA remained unchanged (pretraining: 30.4 +/- 8.9 cm(2); posttraining: 29.1 +/- 8.4 cm(2); P > 0.05). Activation capacity and VL muscle root mean square electromyographic activity increased by 5 and 40%, respectively, after training (P < 0.05), indicating increased agonist neural drive, whereas antagonist cocontraction remained unchanged (P > 0.05). The VL muscle-specific force increased by 19%, from 27 +/- 6.3 N/cm(2) before to 32.1 +/- 7.4 N/cm(2) after training (P < 0.01), highlighting the effectiveness of strength training for increasing the intrinsic force-producing capacity of skeletal muscle in old age.     

Scaling of peak moment arms of elbow muscles with upper extremity bone dimensions.

It is often assumed that moment arms scale with size and can be normalized by body segment lengths or limb circumferences. However, quantitative scaling relationships between moment arms and anthropometric dimensions are generally not available. We hypothesized that peak moment arms of the elbow flexor and extensor muscles scale with the shorter distance (D(s)) between the elbow flexion axis and a muscle's origin and insertion. To test this hypothesis, we estimated moment arms of six muscles that cross the elbow, digitized muscle attachment sites and bone surface geometry, and estimated the location of the elbow flexion axis in 10 upper extremity cadaveric specimens which ranged in size from a 5'0" female to a 6'4" male. D(s) accurately reflected the differences in peak moment arms across different muscles, explaining 93-99% of the variation in peaks between muscles in the same specimen. D(s) also explained between 55% and 88% of the interspecimen variation in peak moment arms for brachioradialis, biceps, and ECRL. Triceps peak moment arm was significantly correlated to the anterior-posterior dimension of the ulna measured at the olecranon (r(2)=0.61, p=0.008). Radius length provides a good measure of the interspecimen variation in peaks for brachioradialis, biceps, and ECRL. However, bone lengths were not significantly correlated to triceps moment arm or anterior-posterior bone dimensions. This work advances our understanding of the variability and scaling dimensions for elbow muscle moment arms across subjects of different sizes.     

Gastrocnemius muscle specific force in boys and men.

The aim of this study was to assess whether the in vivo specific force and architectural characteristics of the lateral gastrocnemius (GL) muscle of early pubescent boys (n = 11, age = 10.9 +/- 0.3 yr, Tanner stage 2) differed from those of adult men (n = 12, age = 25.3 +/- 4.4 yr). Plantarflexor torque was 55% lower in the boys (77.4 +/- 21.4 N x m) compared with the adults (175.6 +/- 31.7 N x m, P < 0.01). Physiological cross-sectional area (PCSA), determined in vivo using ultrasonography and MRI, was 52% smaller in the boys (P < 0.01). No difference was found in pennation angle, or in the ratio of fascicle length (L(f)) to muscle length between the boys and men. Moment arm length was 25% smaller in the boys (P < 0.01). Antagonist coactivation, assessed using surface EMG on the dorsiflexors, was not different between the boys and men (11.8 +/- 6.7% and 13.5 +/- 5.8%, respectively). Surprisingly, GL force normalized to PCSA (specific force) was significantly higher (21%) in the boys than in the men (13.1 +/- 2.0 vs. 15.9 +/- 2.7 N/cm(2), P < 0.05). This finding could not be explained by differences in moment arm length, muscle activation, or architecture, and other factors, such as tendinous characteristics and/or changes in moment arm length with contraction, may be held responsible. These observations warrant further investigation.     

Influence of different shortening velocities preceding stretch on human triceps surae moment generation in vivo

The purpose of this study was to examine the influence of different shortening velocities preceding the stretch on moment generation of the triceps surae muscles and architecture of the m. gastrocnemius medialis after shortening-stretch cycles of equal magnitude in vivo. Eleven male subjects (31.6+/-5.8 years, 178.4+/-7.3cm, 80.6+/-9.6kg) performed a series of electro-stimulated (85Hz) shortening-stretch plantar flexion contractions. The shortening-stretch cycles were performed at three constant angular velocities (25, 50, 100 degrees /s) in the plantar flexion direction (shortening) and at 50 degrees /s in the dorsiflexion direction (stretching). The resultant ankle joint moments were calculated through inverse dynamics. Pennation angle and fascicle length of the m. gastrocnemius medialis at rest and during contractions were measured using ultrasonography. The corresponding ankle moments, kinematics and changes in muscle architecture were analysed at seven time intervals. An analysis of variance for repeated measurements and post hoc test with Bonferroni correction was used to check the velocity-related effects on moment enhancement (alpha=0.05). The results show an increase in pennation angles and a decrease in fascicle lengths after the shortening-stretch cycle. The ankle joint moment ratio (post to pre) was higher (p<0.01) than 1.0 indicating a moment enhancement after the shortening-stretch cycle. The found ankle joint moment enhancement was 2-5% after the shortening-stretch cycle and was independed of the shortening velocity. Furthermore, the decrease in fascicle length after the shortening-stretch cycle indicates that the moment enhancement found in the present study is underestimated at least by 1-3%. Considering that the experiments have been done at the ascending limb of the force-length curve and that force enhancement is higher at the descending and the plateau region of the force-length curve, we conclude that the moment enhancement after shortening-stretch cycle can have important physiological affects while locomotion.     

Morphometry of Macaca mulatta forelimb. I. Shoulder and elbow muscles and segment inertial parameters

The present study examined the morphometric properties of the forelimb, including the inertial properties of the body segments and the morphometric parameters of 21 muscles spanning the shoulder and/or elbow joints of six Macaca mulatta and three M. fascicularis. Five muscle parameters are presented: optimal fascicle length (L(0)(M)), tendon slack length (L(S)(T)), physiological cross-sectional area (PCSA), pennation angle (alpha(0)), and muscle mass (m). Linear regressions indicate that muscle mass, and to a lesser extent PCSA, correlated with total body weight. Segment mass, center-of-mass, and the moment of inertia of the upper arm, forearm, and hand are also presented. Our data indicate that for some segments, radius of gyration (rho) predicts segment moment of inertia better than linear regressions based on total body weight. Key differences between the monkey and human forelimb are highlighted.     

EMG and MMG of agonist and antagonist muscles as a function of age and joint angle.

The aim of this study was to determine the effect of elbow joint position on electromyographic (EMG) and mechanomyographic (MMG) activities of agonist and antagonist muscles in young and old women. Surface EMG and MMG were recorded from the triceps and biceps brachii, and brachioradialis muscles during isometric elbow extensions in young and old women. The measurements were carried out at an optimal joint angle (A(o)), as well as at smaller (A(s) = A(o) - 30 degrees ) and larger (A(l) = A(o) + 30 degrees ) angles. The normalized to force EMG amplitude (RMS-EMG/F) was smaller in old women compared to young in all muscles. The RMS-EMG/F of the triceps brachii muscle was not affected by muscle length while that of the biceps brachii and brachioradialis muscles increased at shortest muscle length in both groups. The normalized to force MMG amplitude (RMS-MMG/F) was smaller in old than in young in the triceps brachii muscle only. There was an increase in RMS-MMG/F with triceps brachii and biceps brachii muscle shortening in both groups, and in the brachioradialis muscle -- in young only. Compared to young, older women exhibited a bigger force fluctuation during maximum voluntary contraction, but these did not contribute significantly to the RMS-MMG. Skinfold thickness accounted for the RMS-EMG/F and RMS-MMG/F differences seen between old and young women in the biceps brachii muscle only. It is concluded that, the EMG and MMG response to muscles length change in agonist and antagonist muscles is generally similar in old and young women but the optimal angle shifts toward a bigger value in older women.     

In vivo measurements of moment arm lengths of three elbow flexors at rest and during isometric contractions

The purpose of this study was to determine in vivo moment arm lengths (MAs) of three elbow flexors at rest and during low- and relatively high-intensity contractions, and to examine the contraction intensity dependence of MAs at different joint positions. At 50°, 80° and 110° of elbow flexion, MAs of the biceps brachii, brachialis and brachioradialis were measured in 10 young men using sagittal images of the right arm obtained by magnetic resonance imaging, at rest and during 20% and 60% of isometric maximal voluntary elbow flexion. In most conditions, MAs increased with isometric contractions, which is presumably due to the contraction-induced thickening of the muscles. This phenomenon was especially evident in the flexed elbow positions. The influence of the contraction intensities on the increases in MAs varied across the muscles. These results suggest that in vivo measurements of each elbow flexor MA during contractions are essential to properly examine the effects on the interrelationships between elbow flexion torque and individual muscle forces.     

Architectural Properties of Sloth Forelimb Muscles (Pilosa: Bradypodidae)

Tree sloths have reduced skeletal muscle mass, and yet they are able to perform suspensory behaviors that require both strength and fatigue resistance to suspend their body mass for extended periods of time. The muscle architecture of sloths is hypothesized to be modified in ways that will enhance force production to compensate for this reduction in limb muscle mass. Our objective is to test this hypothesis by quantifying architecture properties in the forelimb musculature of the brown-throated three-toed sloth (Bradypus variegatus: N = 4). We evaluated architecture from 52 forelimb muscles by measuring muscle moment arm (r _m), muscle mass (MM), belly length (ML), fascicle length (L_F), pennation angle (θ), and physiological cross-sectional area (PCSA), and these metrics were used to estimate isometric force, joint torque, and power. Overall, the musculature becomes progressively more pennate from the extrinsic to intrinsic regions of the forelimb, and the flexors are more well developed than the extensors as predicted. However, most muscles are indicative of a mechanical design for fast joint rotational velocity instead of large joint torque (i.e., strength), although certain large, parallel-fibered shoulder (e.g., m. latissimus dorsi) and elbow (e.g., m. brachioradialis) flexors are capable of producing appreciable torques by having elongated moment arms. This type of functional tradeoff between joint rotational velocity and mechanical advantage is further exemplified by muscle gearing in Bradypus that pairs synergistic muscles with opposing L_F/r _m ratios in each functional group. These properties are suggested to facilitate the slow, controlled movements in sloths. In addition, the carpal/digital flexors have variable architectural properties, but their collective PCSA and joint torque indicates the capability for maintaining grip force and carpal stability while distributing load from the manus to the shoulder. The observed specializations provide a basis for understanding sustained suspension in sloths.     

Individual muscle force parameters and fiber operating ranges for elbow flexion-extension and forearm pronation-supination

We have quantified individual muscle force and moment contributions to net joint moments and estimated the operating ranges of the individual muscle fibers over the full range of motion for elbow flexion/extension and forearm pronation/supination. A three dimensional computer graphics model was developed in order to estimate individual muscle contributions in each degree of freedom over the full range of motion generated by 17 muscles crossing the elbow and forearm. Optimal fiber length, tendon slack length, and muscle specific tension values were adjusted within the literature range from cadaver studies such that the net isometric joint moments of the model approximated experimental joint moments within one standard deviation. Analysis of the model revealed that the muscles operate on varying portions of the ascending limb, plateau region, and descending limb of the force-length curve. This model can be used to further understand isometric force and moment contributions of individual muscles to net joint moments of the arm and forearm and can serve as a comprehensive reference for the forces and moments generated by 17 major muscles crossing the elbow and wrist.     

In vivo physiological cross-sectional area and specific force are reduced in the gastrocnemius of elderly men.

Sarcopenia and muscle weakness are well-known consequences of aging. The aim of the present study was to ascertain whether a decrease in fascicle force (Ff) could be accounted for entirely by muscle atrophy. In vivo physiological cross-sectional area (PCSA) and specific force (Ff/PCSA) of the lateral head of the gastrocnemius (GL) muscle were assessed in a group of elderly men [EM, aged 73.8 yr (SD 3.5), height 173.4 cm (SD 4.4), weight 78.4 kg (SD 8.3); means (SD)] and for comparison in a group of young men [YM, aged 25.3 yr (SD 4.4), height 176.4 cm (SD 7.7), weight 79.1 kg (SD 11.9)]. GL muscle volume (Vol) and Achilles tendon moment arm length were evaluated using magnetic resonance imaging. Pennation angle and fiber fascicle length (Lf) were measured using B-mode ultrasonography during isometric maximum voluntary contraction of the plantar flexors. PCSA was estimated as Vol/Lf. GL Ff was calculated by dividing Achilles tendon force by the cosine of theta, during the interpolation of a supramaximal doublet, and accounting for antagonist activation level (assessed using EMG), Achilles tendon moment arm length, and the relative PCSA of the GL within the plantar flexor group. Voluntary activation of the plantar flexors was lower in the EM than in the YM (86 vs. 98%, respectively, P < 0.05). Compared with the YM, plantar flexor maximal voluntary contraction torque and Ff of the EM were lower by 47 and 40%, respectively (P < 0.01). Both Vol and PCSA were smaller in the EM by 28% (P < 0.01) and 16% (P < 0.05), respectively. Also, pennation angle was 12% smaller in the EM, whereas there was no significant difference in Lf between the YM and EM. After accounting for differences in agonists and antagonists activation, the Ff/PCSA of the EM was 30% lower than that of the YM (P < 0.01). These findings demonstrate that the loss of muscle strength with aging may be explained not only by a reduction in voluntary drive to the muscle, but mostly by a decrease in intrinsic muscle force. This phenomenon may possibly be due to a reduction in single-fiber specific tension.     

Nature of the coupling between neural drive and force-generating capacity in the human quadriceps muscle

The force produced by a muscle depends on both the neural drive it receives and several biomechanical factors. When multiple muscles act on a single joint, the nature of the relationship between the neural drive and force-generating capacity of the synergistic muscles is largely unknown. This study aimed to determine the relationship between the ratio of neural drive and the ratio of muscle force-generating capacity between two synergist muscles (vastus lateralis (VL) and vastus medialis (VM)) in humans. Twenty-one participants performed isometric knee extensions at 20 and 50% of maximal voluntary contractions (MVC). Myoelectric activity (surface electromyography (EMG)) provided an index of neural drive. Physiological cross-sectional area (PCSA) was estimated from measurements of muscle volume (magnetic resonance imaging) and muscle fascicle length (three-dimensional ultrasound imaging) to represent the muscles'' force-generating capacities. Neither PCSA nor neural drive was balanced between VL and VM. There was a large (r = 0.68) and moderate (r = 0.43) correlation between the ratio of VL/VM EMG amplitude and the ratio of VL/VM PCSA at 20 and 50% of MVC, respectively. This study provides evidence that neural drive is biased by muscle force-generating capacity, the greater the force-generating capacity of VL compared with VM, the stronger bias of drive to the VL.     

Muscle synergy in isometric flexion of the elbow]

A technique was developed for calculating the torque generated by two individual muscles (biceps brachii and brachioradialis) that contribute to the isometric flexion of the elbow. The external torque is the sum of individual torques which are unknown. Each individual torque (CB or CBR) can be related to the corresponding integrated surface EMG (QB or QBR) by means of coefficients (pB or PBR). A block of several equations C = pB QB + pBR QBR is obtained by exploring several experimental conditions. In these conditions, isometric flexion efforts of the elbow were associated to isometric efforts of supination or pronation so as to vary integrated EMG by reciprocal inhibition. By means of a least square method it was possible to know the coefficients PB and PBR. With these coefficients, it was possible to calculate the individual torques generated by the biceps brachii and brachioradialis muscles in each experimental condition.     

The long head of the triceps brachii as a free functioning muscle transfer

This anatomic study investigates the possibility of using the long head of the triceps brachii muscle as a free functioning muscle transfer for the upper limb. It has been reported that the long head is not difficult to harvest and that its loss does not create significant donor-site morbidity. The muscle was studied in 23 fresh frozen upper limbs. The long head in all 23 specimens had a constant and proximal vascular pedicle from the profunda brachii artery and vein. The mean pedicle was long (4 cm) and had large-caliber vessels (diameter, 3-mm artery and 4-mm vein). Angiograms were carried out in five specimens and dye perfusion studies in six specimens. A single branch from the radial nerve of at least 7 cm in length innervated the muscle. Muscle architecture was studied in 12 specimens and revealed that the long head of the triceps is better suited for forearm reconstruction than either the gracilis or the latissimus dorsi muscles. The mean physiologic cross-sectional area (8.36 cm(2)) and fiber length (10.8 cm on the superficial surface and 8.2 cm on the deep surface) of the long head match more closely those of the flexor digitorum profundus and the extensor digitorum communis, the muscles most commonly replaced.     

Muscle architecture and out‐force potential of the thoracic limb in the eastern mole (Scalopus aquaticus)

Moles have modified thoracic limbs with hypertrophied pectoral girdle muscles that allow them to apply remarkably high lateral out‐forces during the power stroke when burrowing. To further understand the high force capabilities of mole forelimbs, architectural properties of the thoracic limb muscles were quantified in the Eastern mole (Scalopus aquaticus). Architectural properties measured included muscle mass, moment arm, belly length, fascicle length, and pennation angle, and these were used to provide estimates of maximum isometric force, joint torque, and power. Measurements of muscle moment arms and limb lever lengths were additionally used to analyze the out‐force contributions of the major pectoral girdle muscles. Most muscles have relatively long fascicles and little‐to‐no pennation. The humeral abductor/rotators as a functional group are massive and are capable of relatively high force, power, and joint torque. Of this group, the bipennate m. teres major is the most massive and has the capacity to produce the highest force and joint torque to abduct and axially rotate the humerus. In general, the distal limb muscles are relatively small, but have the capacity for high force and mechanical work by fascicle shortening. The muscle architectural properties of the elbow extensors (e.g., m. triceps brachii) and carpal flexors (e.g., m. palmaris longus) are consistent with the function of these muscles to augment lateral out‐force application. The humeral abductor/rotators m. latissimus dorsi, m. teres major, m. pectoralis, and m. subscapularis are calculated to contribute 13.9 N to out‐force during the power stroke, and this force is applied in a ‘frontal’ plane causing abduction of the humerus about the sternoclavicular joint. Moles have several specializations of their digging apparatus that greatly enhance the application of out‐force, and these morphological features suggest convergence on limb form and burrowing function between New and Old World moles. J. Morphol. 274:1277–1287, 2013. © 2013 Wiley Periodicals, Inc.     

Electromyographic measures of muscle activation and changes in muscle architecture of human elbow flexors during fatiguing contractions.

The study compared changes in intramuscular and surface recordings of EMG amplitude with ultrasound measures of muscle architecture of the elbow flexors during a submaximal isometric contraction. Ten subjects performed a fatiguing contraction to task failure at 20% of maximal voluntary contraction force. EMG activity was recorded in biceps brachii, brachialis, and brachioradialis muscles using intramuscular and surface electrodes. The rates of increase in the amplitude of the surface EMG for the long and short heads of biceps brachii and brachioradialis were greater than those for the intramuscular recordings measured at different depths. The amplitude of the intramuscular recordings from three muscles increased at a similar rate (P = 0.13), as did the amplitude of the three surface recordings from two muscles (P = 0.83). The increases in brachialis thickness (27.7 +/- 5.7 to 30.9 +/- 3.5 mm; P < 0.05) and pennation angle (10.9 +/- 3.5 to 16.5 +/- 4.8 degrees ; P = 0.003) were not associated with the increase in intramuscular EMG amplitude (P > 0.58). The increase in brachioradialis thickness (22.8 +/- 4.8 to 25.5 +/- 3.4 mm; P = 0.0075) was associated with the increase in the amplitude for one of two intramuscular EMG signals (P = 0.007, r = 0.79). The time to failure was more strongly associated with the rate of increase in the amplitude of the surface EMG than that for the intramuscular EMG, which suggests that the surface measurement provides a more appropriate measure of the change in muscle activation during a fatiguing contraction.     

In vivo fascicle behavior of synergistic muscles in concentric and eccentric plantar flexions in humans.

Ultrasonography was used to directly measure in vivo fascicle behavior of the medial gastrocnemius (MG) and soleus (SOL) muscles while the subjects (n=6 men) performed maximal voluntary concentric and eccentric plantar flexions at 60, 120, 180 and 240 deg/s. Fascicle shortening and lengthening velocities of MG, obtained from fascicle length changes over time, were significantly higher than those of SOL at +/-120, +/-180 and +240 deg/s, possibly reflecting physiological and mechanical differences between these muscles. On the other hand, the effective fascicle shortening and lengthening velocities, defined as the velocities in the longitudinal direction of muscle belly, were not significantly different between MG and SOL. This could be due to difference in fascicle architecture and/or the existence of mechanical linkages between these muscles. Moreover, when the contribution of tendinous tissues to muscle-tendon complex length change was determined from fascicle length, pennation angle, moment arm and joint angle, it accounted for approximately 50% in both concentric and eccentric trials, but showed considerable intra-subject variations. This result quantifiably demonstrates the importance of tendinous tissues in isokinetically controlled joint movements.     

The effect of handgrip position on upper extremity neuromuscular responses to arm cranking exercise.

The purpose of this study was to determine if handgrip position during arm cranking exercise influences the neuromuscular activity of muscles biceps brachii (BB), lateral head of triceps brachii (TB), middle deltoid (DT), infraspinatus (IS) and brachioradialis (BR). Fifteen participants cranked an arm ergometer using three different handgrip positions (supinated, pronated, and neutral). Electromyographic (EMG) data were recorded from the aforementioned muscles, and relative duration of EMG activation and amplitude were quantified for the first and second 180 degrees of crank angle. EMG measures were analyzed with MANOVA and follow-up univariate procedures; alpha was set at 0.01. The relative durations of EMG activation did not differ between handgrip positions. Muscle IS exhibited 36% less amplitude in the supinated versus neutral handgrip position (second half-cycle), and muscle BR displayed 63% greater amplitude across cycles in the neutral versus supinated and pronated handgrip positions. The greater BR activity displayed in the neutral handgrip position may reflect its anatomical advantage as an elbow flexor when the forearm is in neutral position. Muscle IS exhibited less activity in the supinated position and may be clinically relevant if it allows arm cranking to occur without subsequent shoulder pain, which is often the aim of shoulder rehabilitation.     

Moment distribution among human elbow extensor muscles during isometric and submaximal extension

To understand better how the central nervous system (CNS) distributes a joint moment among muscles, moment distribution among the three heads of the triceps and the anconeus muscles during isometric elbow extension was quantified in vivo and noninvasively. Electrical stimulation was used to activate an individual muscle selectively at various contraction levels, and the relationship between the peak M-wave amplitude and peak elbow extension moment was established across various contraction levels for each muscle. The relationship was then used to calibrate the corresponding EMG signal and determine moment distribution among the muscles during voluntary isometric elbow extension. Results showed that moment distribution among muscles was not proportional to the muscles' physiological cross-sectional areas (PCSA) and the CNS favored uniarticular muscles for the isometric task performed: the uniarticular lateral and medial heads of the triceps were dominant (contributing approximately 70-90% of the total elbow extension moment) and the anconeus contributed significantly, especially at the lower levels of elbow extension moment (up to approximately 15% of the extension moment). In contrast, the two-joint long head of the triceps contributed significantly less than the uniarticular heads of the triceps. While the absolute contributions of all the muscles increased with the total elbow extension moment, the relative contributions of the muscles may increase or decrease with the elbow extension moment. Cross-validation using fresh data (not used in determining the moment distribution) showed close match between the measured and predicted elbow extension moment except for trials in which fatigue became significant.