The 3D in vivo Achilles' tendon moment arm, quantified during active muscle control and compared across sexes

The Achilles' tendon moment arm (ATma) is a critical quantity in that it defines the triceps surae's ability to generate a moment on the calcaneus, which is then transferred to the foot. This measure has been primarily acquired two-dimensionally in small male populations. Thus, the primary purpose of this study was to establish the first in vivo three-dimensional measures of the ATma, measured non-invasively during dynamic activity in a large normative population, inclusive of both males and female subjects (n=20). Subjects were each placed supine in a 1.5 T MRI and asked to repeat a simulated toe-raise while a full sagittal-cine-phase contrast (dynamic) MRI dataset was acquired. From these data, the 3D and 2D ATma was calculated. The ATma was scaled by the distal tibial width, based on a correlation analysis. The 2D ATma overestimated its 3D counterpart by 3.1 mm, on average. The scaled ATma was no different between the male and female cohorts, but the scaled Achilles' tendon area was smaller in the male cohort. The magnitudes of the ATma were most similar to previously reported values when variations in ankle angle were taken into account. The results of this study have important implications for the applicability of ATma data to both clinical questions and modeling. Any future studies should adapt the ATma based on subject size and/or sex, ensure compatibility between the manner in which the ankle angle is defined and the data being used, and account for the influence that muscle force has on the 3D ATma.     

Moment arm of the patellar tendon in the human knee

The moment arm of the knee-extensor mechanism is described by the moment arm of the patellar tendon calculated with respect to the screw axis of the tibia relative to the femur. The moment arm may be found once the line of action of the patellar tendon and the position and orientation of the screw axis are known. In this study, the orientation of the patellar tendon and the position and orientation of the finite screw axis of the tibia relative to the femur were calculated from measurements of the three-dimensional positions of the bones obtained from fresh cadaver specimens. Peak values of the patellar tendon moment arm ranged from 4-6 cm for the six knees tested; the moment arm was maximum near 45 degrees of knee flexion. The moment arm of the patellar tendon was nearly equal to the shortest (perpendicular) distance between the line of action of the patellar tendon and the axis of rotation of the knee at all flexion angles, except near full extension. Near full extension, the angle between the patellar tendon and the screw axis was significantly less than 90 degrees, and the magnitude of the moment arm was then less than the perpendicular distance between these two lines. The patellar tendon moment arm remained roughly constant across individuals when normalized by femoral condyle width, suggesting that anatomical differences play a large role in determining the moment arm of the extensor mechanism.     

Dynamic in vivo 3-dimensional moment arms of the individual quadriceps components

The purpose of this study was to provide the first in vivo 3-dimensional (3D) measures of knee extensor moment arms, measured during dynamic volitional activity. The hypothesis was that the vastus lateralis (VL) and vastus medialis (VM) have significant off-axis moment arms compared to the central quadriceps components. After obtaining informed consent, three 3D dynamic cine phase contrast (PC) MRI sets (x,y,z velocity and anatomic images) were acquired from 22 subjects during active knee flexion and extension. Using a sagittal-oblique and two coronal-oblique imaging planes, the origins and insertions of each quadriceps muscle were identified and tracked through each time frame by integrating the cine-PC velocity data. The moment arm (MA) and relative moment (RM, defined as the cross product of the tendon line-of-action and a line connecting the line-of-action with the patellar center of mass) were calculated for each quadriceps component. The tendencies of the VM and VL to produce patellar tilt were evenly balanced. Interestingly, the magnitude of RM-P_Spin for the VM and VL is approximately four times greater than the magnitude of RM-P_Tilt for the same muscles suggesting that patellar spin may play a more important role in patellofemoral kinematics than previously thought. Thus, a force imbalance that leads to excessive lateral tilt, such as VM weakness in patellofemoral pain syndrome, would produce excessive negative spin (positive spin: superior patellar pole rotates laterally) and to a much greater degree. This would explain the increased negative spin found in recent studies of patellar maltracking. Assessing the contribution of each quadriceps component in three dimensions provides a more complete understanding of muscle functionality.     

Can the patellar tendon moment arm be predicted from anthropometric measurements?

The purpose of this study was to examine the relations between patellar tendon moment arm length and several relevant anthropometric characteristics of 22 healthy men. The patellar tendon moment arm length was measured using magnetic resonance imaging with two different methods: (1) measurement of patellar tendon moment arm length (d(PT)) with respect to the tibiofemoral contact point (d(PTCP)) and (2) measurement of d(PT) with respect to the intersection point of the anterior and posterior cruciate ligament (d(PTIP)). Pearson correlation coefficients and a stepwise linear regression analysis were used to examine the relationships between the d(PT) and anthropometric measurements taken. Furthermore, a Student's t-test was used to determine differences between the d(PTCP) and d(PTIP) values. Only knee circumference was a significant d(PTCP) predictor (P < 0.05) but with a very low R2 (0.139). None of the anthropometric parameters examined was found to be a significant d(PTIP) predictor. The correlation coefficients ranged from -0.04 to 0.42. The d(PTIP) values were significantly higher (by 0.84-1.89 cm) than the d(PTCP) values (P < 0.05). These results are in disagreement with previous in vitro findings that d(PT) variance may be explained by knee joint size differences. Hence, existing imaging-based methodologies remain necessary for accurate quantification of the patellar tendon moment arm.     

Influence of patellar position on the knee extensor mechanism in normal and crouched walking

Patella alta is common in cerebral palsy, especially in patients with crouch gait. Correction of patella alta has been advocated in the treatment of crouch, however the appropriate degree of correction and the implications for knee extensor function remain unclear. Therefore, the goal of this study was to assess the impact of patellar position on quadriceps and patellar tendon forces during normal and crouch gait. To this end, a lower extremity musculoskeletal model with a novel 12 degree of freedom knee joint was used to simulate normal gait in a healthy child, as well as mild (23 deg min knee flexion in stance), moderate (41 deg), and severe (67 deg) crouch gait in three children with cerebral palsy. The simulations revealed that quadriceps and patellar tendon forces increase dramatically with crouch, and are modulated by patellar position. For example with a normal patellar tendon position, peak patellar tendon forces were 0.7 times body weight in normal walking, but reached 2.2, 3.2 and 5.4 times body weight in mild, moderate and severe crouch. Moderate patella alta acted to reduce quadriceps and patellar tendon loads in crouch gait, due to an enhancement of the patellar tendon moment arms with alta in a flexed knee. In contrast, patella baja reduced the patellar tendon moment arm in a flexed knee and thus induced an increase in the patellar tendon loads needed to walk in crouch. Functionally, these results suggest that patella baja could also compromise knee extensor function for other flexed knee activities such as chair rise and stair climbing. The findings are important to consider when using surgical approaches for correcting patella alta in children who exhibit crouch gait patterns.     

Knee muscle moment arms from MRI and from tendon travel.

We tested magnetic resonance imaging (MRI) as a means to collect geometric data for moment arm estimation. A knee specimen in five successive flexion postures was scanned by MRI, while simultaneously tendon positions of loaded muscles were measured (long head of biceps femoris, lateral and medial gastrocnemius, gracilis, rectus femoris, sartorius, semimembranosus, semitendinosus, and tensor fasciae latae). Discrete rotation centres were derived from MRI pictures. Moment arms were estimated as the distances from these centres to the tendons. The ratio of tendon travel over the increment of joint angulation was the alternative, more reliable estimate of the moment arm. An important principal shortcoming of MRI is the impossibility of accounting for force distribution in taut tissue. As a consequence, for some muscles, considerable inaccuracies in moment arm estimation are found in a relatively small range of joint angulation (up to about 30% for the rectus femoris and semimembranosus). For the tensor fasciae latae, the moment arm cannot be estimated by MRI, while the estimate by tendon travel is unreliable owing to the deformability and attachments of the fascia lata.     

The human patellar tendon moment arm assessed in vivo using dual-energy X-ray absorptiometry

Accurate assessment of muscle–tendon forces in vivo requires knowledge of the muscle–tendon moment arm. Dual-energy X-ray absorptiometry (DXA) can produce 2D images suitable for visualising both tendon and bone, thereby potentially allowing the moment arm to be measured but there is currently no validated DXA method for this purpose. The aims of this study were (i) to compare in vivo measurements of the patellar tendon moment arm (d_PT) assessed from 2D DXA and magnetic resonance (MR) images and (ii) to compare the reliability of the two methods. Twelve healthy adults (mean±SD: 31.4±9.5 yr; 174.0±9.5 cm; 76.2±16.6 kg) underwent two DXA and two MR scans of the fully extended knee at rest. The tibiofemoral contact point (TFCP) was used as the centre of joint rotation in both techniques, and the d_PT was defined as the perpendicular distance from the patellar tendon axis to the TFCP. The d_PT was consistently longer when assessed via DXA compared to MRI (+3.79±1.25 mm or +9.78±3.31%; P<0.001). The test–retest reliability of the DXA [CV=2.13%; ICC=0.94; ratio limits of agreement (RLA)=1.01 (?/÷1.07)] and MR [(CV=2.27%; ICC=0.96; RLA=1.00 (?/÷1.07)] methods was very high and comparable between techniques. Moreover, the RLA between the mean DXA and MRI d_PT values [1.097 (?/÷1.061)] demonstrated very strong agreement between the two methods. In conclusion, highly reproducible d_PT measurements can be determined from DXA imaging with the knee fully extended at rest. This has implications for the calculation of patellar tendon forces in vivo where MR equipment is not available.     

Plantarflexor moment arms estimated from tendon excursion in vivo are not strongly correlated with geometric measurements

Geometric and tendon excursion methods have both been used extensively for estimating plantarflexor muscle moment arm in vivo. Geometric measures often utilize magnetic resonance imaging, which can be costly and impractical for many investigations. Estimating moment arm from tendon excursion measured with ultrasonography may provide a cost-effective alternative to geometric measures of moment arm, but how well such measures represent geometry-based moment arms remains in question. The purpose of this study was to determine whether moment arms from tendon excursion can serve as a surrogate for moment arms measured geometrically. Magnetic resonance and ultrasound imaging were performed on 19 young male subjects to quantify plantarflexor moment arm based on geometric and tendon excursion paradigms, respectively. These measurements were weakly correlated that approached statistical significance (R² = 0.21, p = 0.052), and moment arm from tendon excursion under-approximated geometric moment arm by nearly 40% (p < 0.001). This weak correlation between methods is at odds with a prior report (N = 9) of a strong correlation (R² = 0.94) in a similar study. Therefore, we performed 92,378 regression analyses (19 choose 9) to determine if such a strong correlation existed in our study population. We found that certain sub-populations of the current study generated similarly strong coefficients of determination (R² = 0.92), but 84% of all analyses revealed no correlation (p > 0.05). Our results suggest that the moment arms from musculoskeletal geometry cannot be otherwise obtained by simply scaling moment arms estimated from tendon excursion.     

Influence of patella alta on knee extensor mechanics

The purpose of this study was to compare the knee extensor mechanics in persons with and without patella alta. Thirteen subjects with patella alta and 14 subjects with normal patellar position participated in the study. Sagittal and axial MR images of the knee were acquired at 0°, 20°, 40°, and 60° of knee flexion. Measurements of actual moment arm, patellar ligament/quadriceps tendon force ratio, quadriceps effective moment arm, and joint reaction force/quadriceps force ratio were obtained. There were no differences between groups in terms of actual moment arm. However, subjects with patella alta had significantly larger patellar ligament/quadriceps tendon force ratios (1.04±0.02 vs. 0.92±0.02) and quadriceps effective moment arms (4.40±0.09 vs. 4.00±0.09 cm) when averaged across the range of knee flexion angles tested. There was no difference in the joint reaction force/quadriceps force ratio between groups. The observed differences in knee extensor mechanics suggest that individuals with patella alta have a more efficient knee extensor mechanism and would be expected to generate similar joint reaction forces per unit quadriceps force compared to subjects with normal patellar position. Therefore, persons with patella alta may experience less patellofemoral joint reaction force to overcome the same knee flexion moment in the range of 0°–60° of knee flexion.     

In vivo moment arm determination using B-mode ultrasonography

The tendon excursion of the tibialis anterior (TA) muscle was measured in vivo using B-mode ultrasonography in seven subjects under three force levels (0, 30 and 60% maximal voluntary contraction, MVC). For each force level, the TA moment arm (m) was determined by calculating the derivative of the tendon excursion relative to the ankle angle (a). A dynamometer controlled the ankle angle while force levels were monitored. The parametric model proposed by Miller and Dennis (1996), m = R sin(a + delta), where R is the largest moment arm and delta represents the offset angle of R from 90 degrees, was used in a least-squares fit of the relationship between moment arm and ankle angle. The R values at 0% MVC were significantly smaller than those at 30 and 60% MVC. The values of calculated moment arm at 0% MVC were not considered adequate estimates of the TA moment arm because of the possible confounding effect of the slackness of the relaxed muscle-tendon unit in more dorsiflexed positions. The moment arm values at 30 and 60% MVC were believed to provide reliable estimates of those of TA since the application of tension probably reduced the effects of the slackness of the muscle-tendon unit and tendon elongation on tendon excursion measurement at these force levels. Since the ultrasonographic technique is an in vivo application of the tendon excursion technique and therefore takes the functional meaning into consideration, it can yield more significant moment arms than other in vivo or cadaver techniques.     

Can Achilles tendon moment arm be predicted from anthropometric measures in pre-pubescent children?

Muscle-tendon moment arm magnitudes are essential variables for accurately calculating muscle forces from joint moments. Their measurement requires specialist knowledge and expensive resources. Research has shown that the patellar tendon moment arm length is related to leg anthropometry in children. Here, we asked whether the Achilles tendon moment arm (MA(AT)) can be accurately predicted in pre-pubescent children from surface anthropometry. Age, standing height, mass, foot length, inter-malleolar ankle width, antero-posterior ankle depth, tibial length, lower leg circumference, and distances from the calcaneus to the distal head of the 1st metatarsal and medial malleolus were determined in 49 pre-pubescent children. MA(AT) was calculated at three different ankle positions (neutral, 10° plantarflexion, and 10° dorsiflexion) by differentiating tendon excursion, measured via ultrasonography, with respect to ankle angle change using seven different differentiation techniques. Backwards stepwise regression analyses were performed to identify predictors of MA(AT.) When all variables were included, the regression analysis accounted for a maximum of 49% of MA(AT) variance at the neutral ankle angle when a third-order polynomial was used to differentiate tendon excursion with respect to ankle angle. For this condition, foot length and the distance between calcaneus and 1st metatarsal were the only significant predictors, accounting for 47% of the variance (p<0.05). The absolute error associated with this regression model was 3.8±4.4 mm, which would result in significant error (mean=14.5%) when estimating muscle forces from joint moments. We conclude that MA(AT) cannot be accurately predicted from anthropometric measures in children.     

A comparison of the triceps surae and residual muscle moments at the ankle during cycling

The rigid linked system model and principles of inverse dynamics have been widely used to calculate residual muscle moments during various activities. EMG driven models and optimization algorithms have also been presented in the literature in efforts to estimate skeletal muscle forces and evaluate their possible contribution to the residual muscle moment. Additionally, skeletal muscle-tendon forces have been measured, directly, in both animals and humans. The purpose of this investigation was to calculate the moment produced by the triceps surae muscles and compare it to the residual muscle moment at the ankle during cycling at three power outputs (90, 180 and 270 W). Inferences were made regarding the potential contribution made by each triceps surae component to the tendon force using EMG and muscle-tendon length changes. A buckle-type transducer was surgically implanted on the right Achilles tendon of one male subject. Achilles tendon forces measured in vivo were multiplied by their corresponding moment arms to yield the triceps surae moment during the three working conditions. Moment arm lengths were obtained in a separate experiment using magnetic resonance imaging (MRI). Pedal reaction forces, body segment accelerations (determined from high speed film), and appropriate mass parameters served as input to the inverse solution. The triceps surae moment was temporally in phase with and consistently represented approximately 65% of the residual muscle moment at the ankle. These data demonstrate the feasibility of using implanted transducers in human subjects and provide a greater understanding of musculoskeletal mechanics during normal human movements.     

Antibody neutralization of TGF-beta enhances the deterioration of collagen fascicles in a tissue-cultured tendon matrix with ex vivo fibroblast infiltration

A tissue-cultured tendon matrix infiltrated with cultured fibroblasts can be regarded as an ideal tissue-engineered tendon model. To clarify the role of TGF-beta in a tissue-cultured tendon matrix during ex vivo cellular infiltration, the present ex vivo study was conducted to test the following hypothesis that antibody neutralization of TGF-beta enhances weakening of the collagen fascicles of the patellar tendon matrix in response to ex vivo fibroblast infiltration. In skeletally mature female rabbits, fibroblasts were isolated from the right patellar tendons using an explant culture technique, and the left patellar tendons underwent multiple freeze/thaw treatment with liquid nitrogen to obtain an acellular tendon matrix. Each acellular tendon was placed in a collagen gel containing cultured fibroblasts and then incubated with or without anti-TGF-beta1 antibody for 6 weeks. We found that antibody neutralization of TGF-beta enhanced the decrease in the tensile strength and tensile modulus of the collagen fascicles of the patellar tendon matrix in response to ex vivo fibroblast infiltration. The present study indicates a possibility that TGF-beta may have a role in suppressing the material deterioration of the fascicles in the tendon during ex vivo cellular infiltration.     

Orientation of tendons in vivo with active and passive knee muscles

Tendon orientations in knee models are often taken from cadaver studies. The aim of this study was to investigate the effect of muscle activation on tendon orientation in vivo. Magnetic resonance imaging (MRI) images of the knee were made during relaxation and isometric knee extensions and flexions with 0 degrees , 15 degrees and 30 degrees of knee joint flexion. For six tendons, the orientation angles in sagittal and frontal plane were calculated. In the sagittal plane, muscle activation pulled the patellar tendon to a more vertical orientation and the semitendinosus and sartorius tendons to a more posterior orientation. In the frontal plane, the semitendinosus had a less lateral orientation, the biceps femoris a more medial orientation and the patellar tendon less medial orientation in loaded compared to unloaded conditions. The knee joint angle also influenced the tendon orientations. In the sagittal plane, the patellar tendon had a more anterior orientation near full extension and the biceps femoris had an anterior orientation with 0 degrees and 15 degrees flexions and neutral with 30 degrees flexions. Within 0 degrees to 30 degrees of flexion, the biceps femoris cannot produce a posterior shear force and the anterior angle of the patellar tendon is always larger than the hamstring tendons. Therefore, co-contraction of the hamstring and quadriceps is unlikely to reduce anterior shear forces in knee angles up to 30 degrees . Finally, inter-individual variation in tendon angles was large. This suggests that the amount of shear force produced and the potential to counteract shear forces by co-contraction is subject-specific.     

A musculoskeletal model of the human lower extremity: the effect of muscle, tendon, and moment arm on the moment-angle relationship of musculotendon actuators at the hip, knee, and ankle

We have developed a musculoskeletal model of the human lower extremity for computer simulation studies of musculotendon function and muscle coordination during movement. This model incorporates the salient features of muscle and tendon, specifies the musculoskeletal geometry and musculotendon parameters of 18 musculotendon actuators, and defines the active isometric moment of these actuators about the hip, knee, and ankle joints in the sagittal plane. We found that tendon slack length, optimal muscle-fiber length, and moment arm are different for each actuator, thus each actuator develops peak isometric moment at a different joint angle. The joint angle where an actuator produces peak moment does not necessarily coincide with the joint angle where: (1) muscle force peaks, (2) moment arm peaks, or (3) the in vivo moment developed by maximum voluntary contractions peaks. We conclude that when tendon is neglected in analyses of musculotendon force or moment about joints, erroneous predictions of human musculotendon function may be stated, not only in static situations as studied here, but during movement as well.     

Tensile properties of the in vivo human gastrocnemius tendon

In the present experiment we obtained the tensile properties of the human gastrocnemius tendon, a high-stressed tendon suitable for spring-like action during locomotion. Measurements were taken in vivo in six men. The gastrocnemius tendon elongation during tendon loading−unloading induced by muscle contraction−relaxation was measured using real-time ultrasonography. Tendon forces were calculated from the moment generated during isometric plantarflexion contraction, using tendon moment arm length data obtained in vivo with the tendon travel method. Tendon stiffness data were calculated from the slope of the tendon force−elongation curve, and were then normalized to the tendon's original dimensions, obtained from morphometric analysis of sonographs, to estimate the tendon Young's modulus. Mechanical hysteresis values were obtained from area calculations by numerical integration. The elongation of the tendon increased curvilinearly with the force acting upon it, from 1.7±1 mm (0.8±0.3% strain) at 87.5±8.5 N to 11.1±3.1 mm (4.9±1% strain) at 875±85 N. The tendon Young's modulus and mechanical hysteresis were 1.16±0.15 GPa and 18±3%, respectively. These values fall within the range of values obtained from in vitro experiments and are very similar to the respective values recently obtained from in vivo measurements in the less highly stressed human tibialis anterior tendon (1.2 GPa and 19%), thus indicating that the material properties of tendon are independent of physiological loading and function. Combining the present tendon force−elongation data with previously reported Achilles tendon force data recorded during walking indicates that the gastrocnemius tendon would provide 6% of the total external work produced by the locomotor system. This estimate illustrates the contribution of passive elastic mechanisms on the economy and efficiency of walking. The contributions would be greater in more active exercise such as running.     

Estimates of Achilles tendon moment arm differ when axis of ankle rotation is derived from ankle motion

The plantarflexor moment arm of the Achilles tendon determines the mechanical advantage of the triceps surae and also indirectly affects muscle force generation by setting the amount of muscle-tendon shortening per unit of ankle joint rotation. The Achilles tendon moment arm may be determined geometrically from an axis (or center) of joint rotation and the line of action of the tendon force, but such moment arms may be sensitive to the location of the joint axis. Using motion analysis to track an ultrasound probe overlying the Achilles tendon along with markers on the shank and foot, we measured Achilles tendon moment arm during loaded and unloaded dynamic plantarflexion motions in 15 healthy subjects. Three representations of the axis or center of rotation of the ankle were considered: (1) a functional axis, defined by motions of the foot and shank; (2) a transmalleolar axis; and (3) a transmalleolar midpoint. Moment arms about the functional axis were larger than those found using the transmalleolar axis and transmalleolar midpoint (all p < 0.001). Moment arms computed with the functional axis increased with plantarflexion angle (all p < 0.001), and increased with loading in the most plantarflexed position (p < 0.001) but these patterns were not observed when either using a transmalleolar axis or transmalleolar midpoint. Functional axis moment arms were similar to those estimated previously using magnetic resonance imaging, suggesting that using a functional axis for ultrasound-based geometric estimates of Achilles tendon moment arm is an improvement over landmark-based methods.     

Influence of aging on the in vivo properties of human patellar tendon

Tendons are important for optimal muscle force transfer to bone and play a key role in functional ability. Changes in tendon properties with aging could contribute to declines in physical function commonly associated with aging. We investigated the in vivo mechanical properties of the patellar tendon in 37 men and women [11 young (27 +/- 1 yr) and 26 old (65 +/- 1 yr)] using ultrasonography and magnetic resonance imaging (MRI). Patella displacement relative to the tibia was monitored with ultrasonography during ramped isometric contractions of the knee extensors, and MRI was used to determine tendon cross-sectional area (CSA) and signal intensity. At peak force, patellar tendon deformation, stress, and strain were 13 (P = 0.05), 19, and 12% less in old compared with young (P < 0.05). Additionally, deformation, stiffness, stress, CSA, and length were 18, 35, 41, 28, and 11% greater (P < 0.05), respectively, in men compared with women. After normalization of mechanical properties to a common force, no age differences were apparent; however, stress and strain were 26 and 22% higher, respectively, in women compared with men (P < 0.05). CSA and signal intensity decreased 12 and 24%, respectively, with aging (P < 0.05) in the midregion of the tendon. These data suggest that differences in patellar tendon in vivo mechanical properties with aging are more related to force output rather than an age effect. In contrast, the decrease in signal intensity indirectly suggests that the internal milieu of the tendon is altered with aging; however, the physiological and functional consequence of this finding requires further study.     

Neandertal knees: power lifters in the Pleistocene?

It has been proposed (Trinkaus, 1983 a; Miller & Gross, 1998) that the marked thickness of Neandertal patellae and/or the posterior displacement of their tibial condyles increased their relative M. quadriceps femoris moment arms, thereby making their legs powerful in extension. However, it is necessary to compare these reflections of muscle moment arm length to appropriate measures of the body weight moment arm and body mass estimates, both of which are influenced by ecogeographically determined body proportions. Reassessment of tibial condylar displacement and patellar thickness, as well as patellar height, relative to an appropriate measure of the moment arm for the baseline load on the knee (body weight), to that moment arm times estimated body mass, and to that moment arm times a skeletal reflection of body mass (femoral head diameter) rejects the hypothesis that the Neandertals had exceptionally powerful knee extension. Relative tibial condylar displacement remains above that of a modern industrial society sample, but similar to that of the Broken Hill tibia, Late Pleistocene early modern humans and a recent human nonindustrial sample. Relative patellar thickness is similar to that of early modern humans, who have relatively thick patellae compared to the late Holocene human samples. Consequently, once body proportions are taken into account, there is little difference between the Neandertals and other later Pleistocene humans in knee extensor mechanical advantage, and all of these fossil hominids are similar in the more important proximal tibial proportions to those of nonindustrial recent humans.     

Variation in the human Achilles tendon moment arm during walking

The Achilles tendon (AT) moment arm is an important determinant of ankle moment and power generation during locomotion. Load and depth-dependent variations in the AT moment arm are generally not considered, but may be relevant given the complex triceps surae architecture. We coupled motion analysis and ultrasound imaging to characterize AT moment arms during walking in 10 subjects. Muscle loading during push-off amplified the AT moment arm by 10% relative to heel strike. AT moment arms also varied by 14% over the tendon thickness. In walking, AT moment arms are not strictly dependent on kinematics, but exhibit important load and spatial dependencies.