Functional muscle synergies constrain force production during postural tasks

We recently demonstrated that a set of five functional muscle synergies were sufficient to characterize both hindlimb muscle activity and active forces during automatic postural responses in cats standing at multiple postural configurations. This characterization depended critically upon the assumption that the endpoint force vector (synergy force vector) produced by the activation of each muscle synergy rotated with the limb axis as the hindlimb posture varied in the sagittal plane. Here, we used a detailed, 3D static model of the hindlimb to confirm that this assumption is biomechanically plausible: as we varied the model posture, simulated synergy force vectors rotated monotonically with the limb axis in the parasagittal plane (r2=0.94+/-0.08). We then tested whether a neural strategy of using these five functional muscle synergies provides the same force-generating capability as controlling each of the 31 muscles individually. We compared feasible force sets (FFSs) from the model with and without a muscle synergy organization. FFS volumes were significantly reduced with the muscle synergy organization (F=1556.01, p<0.01), and as posture varied, the synergy-limited FFSs changed in shape, consistent with changes in experimentally measured active forces. In contrast, nominal FFS shapes were invariant with posture, reinforcing prior findings that postural forces cannot be predicted by hindlimb biomechanics alone. We propose that an internal model for postural force generation may coordinate functional muscle synergies that are invariant in intrinsic limb coordinates, and this reduced-dimension control scheme reduces the set of forces available for postural control.     

Inter-joint coupling effects on muscle contributions to endpoint force and acceleration in a musculoskeletal model of the cat hindlimb

The biomechanical principles underlying the organization of muscle activation patterns during standing balance are poorly understood. The goal of this study was to understand the influence of biomechanical inter-joint coupling on endpoint forces and accelerations induced by the activation of individual muscles during postural tasks. We calculated induced endpoint forces and accelerations of 31 muscles in a 7 degree-of-freedom, three-dimensional model of the cat hindlimb. To test the effects of inter-joint coupling, we systematically immobilized the joints (excluded kinematic degrees of freedom) and evaluated how the endpoint force and acceleration directions changed for each muscle in 7 different conditions. We hypothesized that altered inter-joint coupling due to joint immobilization of remote joints would substantially change the induced directions of endpoint force and acceleration of individual muscles. Our results show that for most muscles crossing the knee or the hip, joint immobilization altered the endpoint force or acceleration direction by more than 90° in the dorsal and sagittal planes. Induced endpoint forces were typically consistent with behaviorally observed forces only when the ankle was immobilized. We then activated a proximal muscle simultaneous with an ankle torque of varying magnitude, which demonstrated that the resulting endpoint force or acceleration direction is modulated by the magnitude of the ankle torque. We argue that this simple manipulation can lend insight into the functional effects of co-activating muscles. We conclude that inter-joint coupling may be an essential biomechanical principle underlying the coordination of proximal and distal muscles to produce functional endpoint actions during motor tasks.     

In vivo 3-dimensional measurement of the force exerted on a tooth during clenching

We have developed a three-dimensional (3D) force-measuring device for teeth and used it to measure functional forces in vivo. It comprises an inner part forming a metal core (abutment), a 3D piezoelectric force transducer, and an outer part forming a metal crown, all joined together with a steel screw. The force transducer can measure +/- 500 N along the z-axis and +/- 150 N along the x- and y-axes. We evaluated the relationship between output and load and the effects of hysteresis and temperature on the output. The transducer had high linearity (r>0.9999), low hysteresis (1.7% at maximum), and high thermal stability (0.05% per degree) along each axis. The measuring device was mounted on the maxillary left second molar of a healthy male subject; the tooth had been endodontically treated (neurovascular bundle removed) and prepared for metal abutment and a crown. The 3D load calculated from the outputs of the transducer was expressed as a vector of the coordinates based on the Frankfort horizontal (x-y) and sagittal (y-z) planes. The force measured during maximum voluntary clenching was about 170 N; the force vector was directed from the crown to the root medially at an angle of about 10 degrees from the y-z plane and posteriorly at an angle of about 3 degrees from the x-z plane. This transducer will enable measurement of forces applied to different types of prosthetic appliances and has the potential to provide important basic in vivo data for analysis using computer simulation.     

Total shoulder and relative muscle strength in the scapular plane.

Strength profiles of the shoulder joint are measured experimentally for two arm positions in "the scapular plane" in order to present quantitative data on the shoulder strength. Apart from yielding the actual force a subject can exert in various directions, these measurements also exhibit e.g. the strongest and weakest directions, in fact the relative strength in all directions. The inter-individual variation of the direction of maximal force was at most 14 degrees (sd). The experimental profiles are compared with the corresponding theoretical profiles, obtained by using a shoulder model. The calculations were made both with default muscle parameters and individually adapted parameters. The results show that the employed shoulder model, which is based on data from an elderly population, may be adapted to other populations and that the necessary changes in relative muscle strength are those expected on biomechanical grounds. Without model changes the difference between measured (in the mean) and predicted maximal force directions was at most 50 degrees. Muscle parameter adjustment reduced this difference to 23 degrees. The strength profiles clearly indicate in what direction a person can produce larger forces and which muscles that contribute.     

Internal forces and moments in the femur of the rat during gait

The rat is of increasing importance for experimental studies on fracture healing. The healing outcome of long bone fractures is strongly influenced by mechanical factors, such as the interfragmentary movement. This movement depends on the stability of the fracture fixation and the musculoskeletal loads. However, little is known about these loads in rats.The musculoskeletal loads during gait were estimated using an inverse-dynamic musculoskeletal model of the right hindlimb of the rat. This model was based on a micro-CT scan of the lower extremities and an anatomical study using 15 rat cadavers. Kinematics were reconstructed from X-ray movies, taken simultaneously from two perpendicular directions during a gait cycle. The ground reaction forces were taken from the literature. The muscle forces were calculated using an optimization procedure.The internal forces and moments varied over the gait cycle and along the femoral axis. The greatest internal force (up to 7 times bodyweight) acted in the longitudinal direction. The greatest internal moment (up to 13.8 bodyweight times millimeter) acted in the sagittal plane of the femur. The validity of the model was corroborated by comparing the estimated strains caused by the calculated loads on the surface of the femoral mid-shaft with those from the literature.Knowledge of the internal loads in the femur of the rat allows adjustment of the biomechanical properties of fixation devices in fracture healing studies to the desired interfragmentary movement.     

Biomechanical study of the programming of anticipatory postural adjustments associated with voluntary movement

The present research concerns anticipatory postural adjustments (APA), with the purpose of determining whether they are preprogrammed and of specifying their biomechanical finality. The experimental situation allowed us to distinguish between the voluntary movement itself (an upper limb elevation) and the postural adjustments associated with it. To this aim, the upper limb kinematics, evaluated from an accelerometer fixed at wrist level, were compared to the whole body dynamics, recorded by means of a force platform. Movements, executed in series of five, were studied according to three conditions: bilateral flexions (BF) and unilateral flexions (UF), with (IUF) and without (OUF) an additional inertia, of the stretched upper limb(s). Six right handed adults were tested twice. Results showed that the ground reaction resultant forces as well as the ground reaction resultant moment about the vertical axis presented reproducible variations before and after the onset of upper limb acceleration. The biomechanical organization of APA corresponded, for the three experimental conditions, to an upward and forward acceleration of the body center of gravity, and also, for UF, to a resultant moment directed towards the contralateral side. The duration of APA varied with the characteristics of the forthcoming voluntary movement, increasing significantly from BF to OUF and from OUF to IUF. It is concluded that APA correspond to dynamic phenomena which are centrally preprogrammed. The inertia forces associated with APA may, when the time comes, balance the inertia forces due to the movement of the mobile limb therefore counteracting the disturbance to postural equilibrium.(ABSTRACT TRUNCATED AT 250 WORDS)     

Sensitivity of the knee joint kinematics calculation to selection of flexion axes

Various flexion axes have been used in the literature to describe knee joint kinematics. This study measured the passive knee kinematics of six cadaveric human knee specimens using two widely accepted flexion axes; transepicondylar axis and the geometric center axis. These two axes were found to form an angle of 4.0 degrees +/- 0.8 degrees. The tibial rotation calculated using the transepicondylar axis was significantly different than the rotation obtained using the geometric center axis for the same knee motion. At 90 degrees of flexion, the tibial rotation obtained using the transepicondylar axis was 4.8 degrees +/- 9.4 degrees whereas the rotation recorded using the geometric center axis at the same flexion angle was 13.8 degrees +/- 10.2 degrees. At 150 degrees of knee flexion, the rotations obtained from the transepicondylar and the geometric center axes were 7.2 degrees +/- 5.7 degrees and 19.9 degrees +/- 6.9 degrees, respectively. The data suggest that a clear definition of the flexion axis is necessary when reporting knee joint kinematics.     

The mechanism of generation and transmission of forces in leg extension.

This paper deals with the mechanical and electromyographic evaluation of the mechanism generating and transmitting the resultant leg extension force by maximal isometric contraction in two directions, the knee and hip joint being kept at 90 degrees. The two directions were a) from the center of gravity of the body to the ankle joint and b) from a point near the knee to the ankle. Six male subjects in a supine position exerted a maximal leg extension force of 47-112 kg for a) and 51-73 kg for b). These values were close to the smaller values of two forces estimated at the knee and at the hip from maximal isometric forces at the corresponding joint of the same joint angle. It was thus suggested that the joint limiting the resultant leg extension force was the knee for a) and the hip for b). The single joint muscles exhibited almost maximal activities when they concerned the joint which limited the resultant leg extension force. The double joint muscles were often contracted only moderately during the maximal isometric leg extension, indicating a different role of double joint muscles even at the maximal force production at a particular joint.     

Analysis of forces of ACL reconstructions at the tunnel entrance: is tunnel enlargement a biomechanical problem?

Bone tunnel enlargement is a common phenomenon following reconstruction of the anterior cruciate ligament (ACL). Biomechanical and biological factors have been reported as potential causes of this problem. However, there is no analysis of forces between the graft and bone, as the graft changes direction at the bone tunnel entrance. The purpose of this study was to study these 'redirecting forces'. Magnetic resonance images of 10 patients with an ACL reconstruction (age: 26+/-6.8 years) were used to determine the angle between graft and drill holes. Vector analysis was used to calculate the direction and magnitude of the perpendicular component of the force between the bone tunnel and the graft at the entrance of the bone tunnel. Force components were projected into the radiographically important sagittal and coronal planes. Tension of ACL reconstructions was recorded during passive knee motion in 10 cadaveric knee experiments (age: 28.9+/-10.6 years) and the tension multiplied with the force component for each plane. Results are reported for the coronal and sagittal planes, respectively: For -10 degrees of extension, the percentages of graft tension were determined to be 17+/-7 (max: 26; min: 7%) and 26+/-9 (max: 39; min: 16%) for the tibia. They were 59+/-6 (max: 66; min: 48%) and 99+/-1 (max: 1.00; min: 99%) for the femur. Force components were 14.68+/-6.54 and 25.73+/-12.96 N for the tibial tunnel. For the femoral tunnel, they were 52.48+/-19.03 and 90.77+/-32.06 N. Percentages of graft tension and force components were significantly higher for the femoral tunnel compared with the tibial tunnel. Moreover, in the sagittal direction, force components for the femoral tunnel were significantly higher compared with the coronal plane (Wilcoxon test, p < 0.01). The differences in force components calculated in this study corresponds with the amount of tunnel enlargement in the radiographic planes in the literature providing evidence that biomechanical forces play a key role in postoperative tunnel expansion.     

Effect of eccentric exercise on position sense at the human forearm in different postures.

This is a study of the ability of blindfolded human subjects to match the position of their forearms before and after eccentric exercise. The hypothesis tested was that the sense of effort contributed to forearm position sense. The fall in force after the exercise was predicted to alter the relationship between effort and force and thereby induce position errors. In the arms-in-front posture, subjects had their unsupported reference arm set to one of two angles from the horizontal, 30 or 60 degrees , and they matched its position by voluntary placement of their other arm. Matching errors were compared with a task where the arms were counterweighted, so could be moved in the vertical plane with minimal effort, and where the arms were moved in the horizontal plane. In these latter two tasks, the intention was to test whether removal of an effort sensation from holding the arm against gravity influenced matching performance. It was found that, although absolute errors for counterweighted and horizontal matching were no larger than for unsupported matching, their standard deviations, 6.1 and 6.8 degrees , respectively, were significantly greater than for unsupported matching (4.6 degrees ), indicating more erratic matching. The eccentric exercise led, the next day, to a fall in maximum voluntary muscle torque of >or=15%. This was accompanied by a significant increase in matching errors for the unsupported matching task from 2.7 +/- 0.5 to 0.8 +/- 0.7 degrees but not for counterweighted (1.4 +/- 0.2 to -0.2 degrees +/- 1.1 degrees ) or horizontal matching (-1.3 +/- 0.7 degrees to -1.8 +/- 0.7 degrees ). This, it is postulated, is because the reduced voluntary torque after exercise was accompanied by a greater effort required to support the arms, leading to larger matching errors. However, effort is only able to provide positional information for unsupported matching where gravity plays a role. In gravity-neutral tasks like counterweighted or horizontal matching, a change in the effort-force relationship after exercise leaves matching accuracy unaffected.     

Quantifying deficits in the 3D force capabilities of a digit caused by selective paralysis: application to the thumb with simulated low ulnar nerve palsy

We present the development of a vision-feedback method to characterize how selective paralysis distorts the three-dimensional (3D) volume representing digit-tip force production capability and its application to healthy individuals producing thumb-tip force with and without simulated low ulnar nerve palsy (LUNP). Subjects produced maximal static voluntary force spanning the transverse, sagittal and frontal planes of the thumb (16, 15 and 10 subjects, respectively). Subjects produced thumb-tip force tasks in guided and self-selected directions. The envelope (convex hull) of extreme forces in each plane approximated that cross-section of the 3D volume of force capability. Some subjects repeated the tasks with a temporary ulnar nerve block applied at the wrist to simulate complete acute LUNP. Three geometric properties of the force convex hull characterized each cross-section's shape: the ratios of its principal moments of inertia (RPMIs), the orientation of its principal axis (OPA), and its centroid location. Our results show that force production in the thumb's sagittal plane may be a reproducible and objective test to grade motor impairment in LUNP: paired t-tests of the larger RPMI in this plane best distinguished the nerve-blocked cases from the control cases in the guided task (p = 0.012), and Discriminant Analysis of the centroid location for the self-selected task in this plane correctly classified 94.7% of subjects into the control and ulnar nerve-blocked groups. We show that our method measures and detects changes in a digit's force production capabilities. Towards a clinical test of motor impairment in LUNP, this biomechanical study dictates which 3D thumb-tip forces to measure (those in the sagittal plane) and how to measure them (using the self-selected task).     

Evolutionary implications of the unusual walking mechanics of the common marmoset (C. jacchus)

Several features that appear to differentiate the walking gaits of most primates from those of most other mammals (the prevalence of diagonal-sequence footfalls, high degrees of humeral protraction, and low forelimb vs. hindlimb peak vertical forces) are believed to have evolved in response to requirements of locomotion on thin arboreal supports by early primates that had developed clawless grasping hands and feet. This putative relationship between anatomy, behavior, and ecology is tested here by examining gait mechanics in the common marmoset (Callithrix jacchus), a primate that has sharp claws and reduced pedal grasping, and that spends much of its time clinging on large trunks. Kinematic and kinetic data were collected on three male Callithrix jacchus as they walked across a force platform attached to the ground or to raised horizontal poles. The vast majority of all walking gaits were lateral-sequence. For all steps, the humerus was retracted (<90 degrees relative to a horizontal axis) or held in a neutral (90 degrees ) position at forelimb touchdown. Peak vertical forces on the forelimb were always higher than those on the hindlimb. These three features of the walking gaits of C. jacchus separate it from any other primate studied (including other callitrichids). The walking gaits of C. jacchus are mechanically more similar to those of small, nonprimate mammals. The results of this study support previous models that suggest that the unusual suite of features that typify the walking gaits of most primates are adaptations to the requirements of locomotion on thin arboreal supports. These data, along with data from other primates and marsupials, suggest that primate postcranial and locomotor characteristics are part of a basal adaptation for walking on thin branches.     

Orientation of the tyrosyl D, pheophytin anion, and semiquinone Q(A)(*)(-) radicals in photosystem II determined by high-field electron paramagnetic resonance

The radical forms of two cofactors and an amino acid in the photosystem II (PS II) reaction center were studied by using high-field EPR both in frozen solution and in oriented multilayers. Their orientation with respect to the membrane was determined by using one-dimensionally oriented samples. The ring plane of the stable tyrosyl radical, Y(D)(*), makes an angle of 64 degrees +/- 5 degrees with the membrane plane, and the C-O direction is tilted by 72 degrees +/- 5 degrees in the plane of the radical compared to the membrane plane. The semiquinone, Q(A)(*)(-), generated by chemical reduction in samples lacking the non-heme iron, has its ring plane at an angle of 72 degrees +/- 5 degrees to the membrane plane, and the O-O axis is tilted by 21 degrees +/- 5 degrees in the plane of the quinone compared to the membrane plane. This orientation is similar to that of Q(A)(*)(-) in purple bacteria reaction centers except for the tilt angle which is slightly bigger. The pheophytin anion was generated by photoaccumulation under reducing conditions. Its ring plane is almost perpendicular to the membrane with an angle of 70 degrees +/- 5 degrees with respect to the membrane plane. This is very similar to the orientation of the pheophytin in purple bacteria reaction centers. The position of the g tensor with respect to the molecule is tentatively assigned for the anion radical on the basis of this comparison. In this work, the treatment of orientation data from EPR spectroscopy applied to one-dimensionally oriented multilayers is examined in detail, and improvements over previous approaches are given.     

Estimating maximum and psychophysically acceptable hand forces using a biomechanical weakest link approach

Accurate estimation of occupational performance capability facilitates better job (re-) design by informing workplace parties about the potential mismatches between job demands and the capability of their labour force. However, estimating occupational performance requires consideration of multiple factors that may govern capacity. In this paper, a novel model is described that uses a stochastic algorithm to estimate how variability in underlying biomechanical constraints affects hand force capability. In addition, the model estimates psychophysically acceptable hand force capacity thresholds by applying a biomechanical weakest link approach. Model estimates were tested against experimentally determined maximal and psychophysically determined hand forces in two exertion directions in constrained postures. The model underestimated maximum hand force capacity relative to measured maximum hand force by 30% and 35% during downward pressing and horizontal pulling, respectively. These values are consistent with those observed using previous two-dimensional models. Psychophysically acceptable hand forces were also underestimated by 29% during both pressing and pulling. Since the psychophysical estimates were scaled as a percentage of the estimated maximum capacity, this suggests that the underestimation in both predictions may be corrected by improving estimates of maximum hand force. Psychophysically acceptable forces were observed to be partially governed by demands at the biomechanical weakest link.     

External forces on the limbs of jumping lemurs at takeoff and landing

Ground reaction forces were recorded for jumps of three individuals each of Lemur catta and Eulemur fulvus. Animals jumped back and forth between a ground-mounted force plate and a 0.5-m elevated platform, covering horizontal distances of 0.5-2 m. In total, 190 takeoffs and 263 landings were collected. Animals typically jumped from a run up and into a run out, during which they gained or into which they carried horizontal impulse. Correspondingly, vertical impulses dominated takeoffs and landings. Peak forces were moderate in magnitude and not much higher than forces reported for quadrupedal gaits. This is in contrast to the forces for standing jumps of specialized leapers that considerably exceed forces associated with quadrupedal gaits. Force magnitudes for the lemur jumps are more comparable to peak forces reported for other quadrupeds performing running jumps. Takeoffs are characterized by higher hindlimb than forelimb peak forces and impulses. L. catta typically landed with the hindlimbs making first contact, and the hindlimb forces and impulses were higher than the forelimb forces and impulses at landing. E. fulvus typically landed with the forelimbs striking first and also bearing the higher forces. This pattern does not fully conform to the paradigm of primate limb force distribution, with higher hindlimb than forelimb forces. However, the absolute highest forces in E. fulvus also occur at the hindlimbs, during acceleration for takeoff.     

Nasal vestibule wall elasticity: interactions with a nasal dilator strip.

We studied the effect of an adhesive external nasal dilator strip (ENDS) on external nasal geometry in 20 healthy Caucasian adults (10 men, 10 women; age 21-45 yr). The recoil force exerted by ENDS was estimated by bending the device (n = 10) with known weights. In the horizontal direction, a small/medium-sized ENDS in situ exerted a unilateral recoil force of 21.4-22.6 g. Application of ENDS resulted in a displacement of the lateral nasal vestibule walls that had both anterosuperior and horizontal components and that was maintained over an 8-h period. The resultant unilateral nasal vestibule wall displacement at the tip of the device was at 47.6 +/- 2.0 degrees to the horizontal (as related to the plane of the device when in situ) and had a magnitude of 3.5 +/- 0.1 mm. ENDS increased external nasal cross-sectional area by 23.0-65.3 mm2. Nasal vestibule wall compliance was estimated at 0.05-0.16 mm/g. Thus ENDS applies a relatively constant abducting force irrespective of nasal width. Variable responsiveness to ENDS may be related to differences in elastic properties of the nasal vestibule wall.     

Absorption and fluorescence spectra and molecular organization of bacteriochlorophyll in reaction centers of Rhodopseudomonas spheroides

Second derivative spectroscopy, computer curve analysis and Stepanov's equation show that the absorbance and fluorescence spectra of primary electron donor in reaction center of Rhodopseudomonas sphaeroides are splitting each into two asymmetric Gaussian components. Their absorption maxima at -196 degrees are 880 and 896 nm and emission maxima-906 and 923 nm, respectively. The absorption spectrum of Bchl-800 splits in the near infrared region into two bands with maxima at 790 and 803 nm. These components are ascribed to an exciton coupling in the two dimers of bacteriochlorophyll in the reaction center. The Qy transition moments of the two bacteriochlorophyll molecules of primary electron donor make an angle of 110 degrees and the angle between two Qy transitions of the pigment in Bchl-800 dimer is 150 degrees. The distance between the centers of chromophores in the dimers is estimated to be 8-11 A.     

Effect of body tilt on calf muscle performance and blood flow in humans.

To explore the effect of posture on muscle performance, we tested the effects of body tilt angle on the strength, endurance, and fatigue of, and blood flow into, the plantar flexors. Human subjects were fixed to a tilt table that could tilt them from the horizontal (0 degrees ) to upright (90 degrees ) position and enabled force to be applied to a footplate through isometric action of the right calf muscle. In experiment 1, six subjects performed a strength test and graded test (intermittent contractions) to the point of failure at three tilt angles (0, 47, and 90 degrees ). In Experiment 2, seven subjects performed a strength test and constant-force test [70% maximum force (F(max)); intermittent contractions] to the point of failure in the horizontal and three inclined positions (32, 47, and 67 degrees ). In experiment 3, leg blood flow was assessed during constant-force exercise at two intensities (30 and 70% F(max)) and two tilt angles (0 and 67 degrees ) in six subjects. Strength was not affected (P > 0.05) by tilt angle. Time to failure during the graded test was significantly higher at 47 degrees (25.9 +/- 2.0 min) and 90 degrees (25.1 +/- 3.0 min) than 0 degrees (22.2 +/- 2.6 min). Time to failure during the constant-force test was also significantly higher at 32 degrees (7.1 +/- 3.6 min), 47 degrees (8.0 +/- 5.2 min), and 67 degrees (8.6 +/- 5.6 min) compared with 0 degrees (4.0 +/- 2.6 min). When graded or constant-force exercise was performed with arterial flow to the leg eliminated, there were no differences in exercise time between the horizontal and an inclined position. During nonischemic exercise, leg blood flow was significantly higher during exercise in the inclined position. These results demonstrate that head-up tilt improves endurance of the plantar flexors, that this effect occurs in the absence of an effect on strength, and that it depends on an intact peripheral circulation. Moreover, the postural effect on muscle endurance appears to be due to a greater blood flow into the leg, an effect that is established during the initial contractions.     

Spatial, temporal and muscle action patterns of Tai Chi gait.

This study was to quantitatively characterize the spatial, temporal, and neuromuscular activation patterns of Tai Chi gait (TCG). Ten healthy young subjects were tested. The kinematics of TCG and normal gait (NG) were measured using a marker-based motion analysis system and two biomechanical force plates. Surface electromyography (EMG) was recorded from six left-side muscles: tibialis anterior, soleus, peronaeus longus, rectus femoris, semitendinosus, and tensor fasciae latae. The results showed that TCG had (1) a longer cycle duration (11.9+/-2.4 vs. 1.3+/-0.2 s) and a longer duration of single-leg stance time (1.8+/-0.6 vs. 0.4+/-0.05 s); (2) a larger joint motion in ankle dorsi/plantar flexion (40+/-9 degrees vs. 20+/-8 degrees), knee flexion (82+/-8 degrees vs. 53+/-10 degrees), hip flexion (81+/-7 degrees vs. 24+/-4 degrees), and hip abduction (20+/-8 degrees vs. 0+/-3 degrees); (3) a larger lateral body shift (>25% vs. 5% body height); and (4) significant involvement of ankle dorsiflexors, knee extensors/hip flexors and hip abductors, as indicated by significantly higher peak (88+/-14%, 80+/-18% and 83+/-17% vs. 35+/-10%, 14+/-8% and 28+/-19% peak amplitude, respectively) and root-mean-square values of their EMG (37+/-6%, 32+/-7% and 33+/-7% vs. 23+/-7%, 11+/-8% and 22+/-11% peak amplitude, respectively), longer proportions of action (76+/-19%, 68+/-8% and 65+/-19% vs. 59+/-23%, 16+/-23% and 40+/-32% gait cycle duration, respectively), longer proportions of isometric and eccentric actions, and longer proportions of co-activations. These results demonstrate that the biomechanical characteristics of TCG can be quantified. The quantification of TCC movements is important for understanding its effect on balance, flexibility, strength, and health.