Gender differences in active musculoskeletal stiffness. Part II. Quantification of leg stiffness during functional hopping tasks.

Leg stiffness was compared between age-matched males and females during hopping at preferred and controlled frequencies. Stiffness was defined as the linear regression slope between the vertical center of mass (COM) displacement and ground-reaction forces recorded from a force plate during the stance phase of the hopping task. Results demonstrate that subjects modulated the vertical displacement of the COM during ground contact in relation to the square of hopping frequency. This supports the accuracy of the spring-mass oscillator as a representative model of hopping. It also maintained peak vertical ground-reaction load at approximately three times body weight. Leg stiffness values in males (33.9+/-8.7 kN/m) were significantly (p<0.01) greater than in females (26.3+/-6.5 kN/m) at each of three hopping frequencies, 3.0, 2.5 Hz, and a preferred hopping rate. In the spring-mass oscillator model leg stiffness and body mass are related to the frequency of motion. Thus male subjects necessarily recruited greater leg stiffness to drive their heavier body mass at the same frequency as the lighter female subjects during the controlled frequency trials. However, in the preferred hopping condition the stiffness was not constrained by the task because frequency was self-selected. Nonetheless, both male and female subjects hopped at statistically similar preferred frequencies (2.34+/-0.22 Hz), therefore, the females continued to demonstrate less leg stiffness. Recognizing the active muscle stiffness contributes to biomechanical stability as well as leg stiffness, these results may provide insight into the gender bias in risk of musculoskeletal knee injury.     

Neural control of leg stiffness during hopping in boys and men

The purpose of the study was to investigate whether boys and men utilise different control strategies whilst hopping. Eleven boys (11–12 yr old) and ten men completed hopping at 1.5 Hz, 3.0 Hz and at their preferred frequency. A footswitch measured contact and flight times, from which leg stiffness was calculated. Simultaneously, surface electromyograms (EMGs) of selected lower limb muscles were recorded and quantified for each 30 ms period during the first 120 ms post-ground contact. At 1.5 Hz there were no differences between the groups in relative stiffness or muscle activity. At 3.0 Hz men had significantly shorter contact times (P = 0.013), longer flight times (P = 0.002), greater relative stiffness (P = 0.01) and significantly greater soleus (P = 0.012) and vastus lateralis (P < 0.001) activity during the initial 30 ms post-ground contact. At the preferred frequency men hopped significantly faster than the boys (P = 0.007), with greater leg stiffness (P < 0.01) and with more extensor activity in most time periods. Boys and men demonstrated similar control strategies when hopping at a slow frequency, but when hopping frequency increased men were able to better increase feedforward and reflex muscle activity to hop with greater relative stiffness.     

Leg stiffness adjustment for a range of hopping frequencies in humans

The purpose of the present study was to determine how humans adjust leg stiffness over a range of hopping frequencies. Ten male subjects performed in place hopping on two legs, at three frequencies (1.5, 2.2, and 3.0 Hz). Leg stiffness, joint stiffness and touchdown joint angles were calculated from kinetic and/or kinematics data. Electromyographic activity (EMG) was recorded from six leg muscles. Leg stiffness increased with an increase in hopping frequency. Hip and knee stiffnesses were significantly greater at 3.0 Hz than at 1.5 Hz. There was no significant difference in ankle stiffness among the three hopping frequencies. Although there were significant differences in EMG activity among the three hopping frequencies, the largest was the 1.5 Hz, followed by the 2.2 Hz and then 3.0 Hz. The subjects landed with a straighter leg (both hip and knee were extended more) with increased hopping frequency. These results suggest that over the range of hopping frequencies we evaluated, humans adjust leg stiffness by altering hip and knee stiffness. This is accomplished by extending the touchdown joint angles rather than by altering neural activity.     

Active leg stiffness and energy stored in the muscles during maximal counter movement jump in the aged

The purpose of this study was to examine the effects of age on active leg stiffness adjustment, electromyogram (EMG) activities and energy stored during eccentric and concentric phases in performing a maximal functional task involving stretch-shorten cycle. Ten young (24.3 ± 2 years) and 10 old (68.6 ± 5 years) healthy male subjects were filmed during maximal performance of counter movement jump (CMJ) and squat jump (SJ) on force plate. Integrated EMG (IEMG), ground reaction force (GRF), active leg stiffness, energy stored/returned and active work done by the muscles were compared between two groups on eccentric (ECC) and concentric (CON) phases of CMJ. The GRF, leg stiffness and energy stored in ECC and GRF, IEMG, energy returned and active work in CON were less in the elderly (p < 0.05). These results demonstrate that the neuromuscular function of adjusting active stiffness, storing elastic energy and optimizing the performance may decrease with age during CMJ.     

Leg stiffness primarily depends on ankle stiffness during human hopping

When humans hop in place or run forward, they adjust leg stiffness to accommodate changes in stride frequency or surface stiffness. The goal of the present study was to determine the mechanisms by which humans adjust leg stiffness during hopping in place. Five subjects hopped in place at 2.2 Hz while we collected force platform and kinematic data. Each subject completed trials in which they hopped to whatever height they chose ("preferred height hopping") and trials in which they hopped as high as possible ("maximum height hopping"). Leg stiffness was approximately twice as great for maximum height hopping as for preferred height hopping. Ankle torsional stiffness was 1.9-times greater while knee torsional stiffness was 1.7-times greater in maximum height hopping than in preferred height hopping. We used a computer simulation to examine the sensitivity of leg stiffness to the observed changes in ankle and knee stiffness. Our model consisted of four segments (foot, shank, thigh, head-arms-trunk) interconnected by three torsional springs (ankle, knee, hip). In the model, increasing ankle stiffness by 1.9-fold, as observed in the subjects, caused leg stiffness to increase by 2.0-fold. Increasing knee stiffness by 1.7-fold had virtually no effect on leg stiffness. Thus, we conclude that the primary mechanism for leg stiffness adjustment is the adjustment of ankle stiffness.     

Influence of muscle activity on musculotendinous stiffness quantification in stunted,prepubertal children

The quick-release technique to estimate musculotendinous (MT) stiffness has been extensively used over the last years, in both animals and humans, to gain insights in the adaptive process of the series elastic component (SEC). Recently, MT stiffness quantification, i.e., SEC behavior, has been revisited for subjects not able to fully activate their muscles (effects of long-term spaceflight or non-mature muscles). Such a phenomenon can also be encountered in stunted children. So, the aim of the present study was to analyze the effect of stunting on MT stiffness taking into account possible defect in muscle activation. For this study, 20 eutrophic children (EU) with an average age of 9 years ± 4 months were compared to 11age matched stunted children (S) evaluated by the height-to-age index. The MT stiffness index was obtained with regard to stiffness–torque and stiffness–soleus EMG relationships. The children of the S group presented a significantly lower Maximal Voluntary Contraction (MVC) in plantar flexion in comparison with children of the EU group (?37.8%). The significantly lower MT stiffness index for S children (?42.6%) was evidenced only when quantified with regard to the stiffness–soleus EMG relationship (66.5 ± 42.8 vs. 38.2 ± 19.9 Nm rad^?1%^?1). Possible delay in fiber type differentiation or tendinous structure maturation can account for the lower MT stiffness index in S children. In conclusion, stunting during early childhood delays the differentiation and maturation processes of musculotendinous structures as shown by the lower MT stiffness quantified with regards to muscle activity, also altered for stunted prepubertal children.     

Knee stiffness is a major determinant of leg stiffness during maximal hopping

Understanding stiffness of the lower extremities during human movement may provide important information for developing more effective training methods during sports activities. It has been reported that leg stiffness during submaximal hopping depends primarily on ankle stiffness, but the way stiffness is regulated in maximal hopping is unknown. The goal of this study was to examine the hypothesis that knee stiffness is a major determinant of leg stiffness during the maximal hopping. Ten well-trained male athletes performed two-legged hopping in place with a maximal effort. We determined leg and joint stiffness of the hip, knee, and ankle from kinetic and kinematic data. Knee stiffness was significantly higher than ankle and hip stiffness. Further, the regression model revealed that only knee stiffness was significantly correlated with leg stiffness. The results of the present study suggest that the knee stiffness, rather than those of the ankle or hip, is the major determinant of leg stiffness during maximal hopping.     

Plasticity of human Achilles tendon mechanical and morphological properties in response to cyclic strain

The purpose of the current study in combination with our previous published data (Arampatzis et al., 2007) was to examine the effects of a controlled modulation of strain magnitude and strain frequency applied to the Achilles tendon on the plasticity of tendon mechanical and morphological properties. Eleven male adults (23.9±2.2 yr) participated in the study. The participants exercised one leg at low magnitude tendon strain (2.97±0.47%), and the other leg at high tendon strain magnitude (4.72±1.08%) of similar frequency (0.5 Hz, 1 s loading, 1 s relaxation) and exercise volume (integral of the plantar flexion moment over time) for 14 weeks, 4 days per week, 5 sets per session. The exercise volume was similar to the intervention of our earlier study (0.17 Hz frequency; 3 s loading, 3 s relaxation) allowing a direct comparison of the results. Before and after the intervention ankle joint moment has been measured by a dynamometer, tendon–aponeurosis elongation by ultrasound and cross-sectional area of the Achilles tendon by magnet resonance images (MRI). We found a decrease in strain at a given tendon force, an increase in tendon–aponeurosis stiffness and tendon elastic modulus of the Achilles tendon only in the leg exercised at high strain magnitude. The cross-sectional area (CSA) of the Achilles tendon did not show any statistically significant (P>0.05) differences to the pre-exercise values in both legs. The results indicate a superior improvement in tendon properties (stiffness, elastic modulus and CSA) at the low frequency (0.17 Hz) compared to the high strain frequency (0.5 Hz) protocol. These findings provide evidence that the strain magnitude applied to the Achilles tendon should exceed the value, which occurs during habitual activities to trigger adaptational effects and that higher tendon strain duration per contraction leads to superior tendon adaptational responses.     

Lower limb joint stiffness and muscle co-contraction adaptations to instability footwear during locomotion

Unstable shoes (US) continually perturb gait which can train the lower limb musculature, but muscle co-contraction and potential joint stiffness strategies are not well understood. A shoe with a randomly perturbing midsole (IM) may enhance these adaptations. This study compares ankle and knee joint stiffness, and ankle muscle co-contraction during walking and running in US, IM and a control shoe in 18 healthy females. Ground reaction forces, three-dimensional kinematics and electromyography of the gastrocnemius medialis and tibialis anterior were recorded. Stiffness was calculated during loading and propulsion, derived from the sagittal joint angle-moment curves. Ankle co-contraction was analysed during pre-activation and stiffness phases. Ankle stiffness reduced and knee stiffness increased during loading in IM and US whilst walking (ankle, knee: p = 0.008, 0.005) and running (p < 0.001; p = 0.002). During propulsion, the opposite joint stiffness re-organisation was found in IM whilst walking (both joints p < 0.001). Ankle co-contraction increased in IM during pre-activation (walking: p = 0.001; running: p < 0.001), and loading whilst walking (p = 0.003), not relating to ankle stiffness. Results identified relative levels of joint stiffness change in unstable shoes, providing new evidence of how stability is maintained at the joint level.     

Vitamin D sufficiency associates with an increase in anti-inflammatory cytokines after intense exercise in humans

The purpose of this study was to identify the influence of vitamin D status (insufficient vs. sufficient) on circulating cytokines and skeletal muscle strength after muscular injury. To induce muscular injury, one randomly selected leg (SSC) performed exercise consisting of repetitive eccentric–concentric contractions. The other leg served as the control. An averaged serum 25(OH)D concentration from two blood samples collected before exercise and on separate occasions was used to establish vitamin D insufficiency (<30 ng/mL, n = 6) and sufficiency (>30 ng/mL, n = 7) in young, adult males. Serum cytokine concentrations, single-leg peak isometric force, and single-leg peak power output were measured before and during the days following the exercise protocol. The serum IL-10 and IL-13 responses to muscular injury were significantly (both p < 0.05) increased in the vitamin D sufficient group. The immediate and persistent (days) peak isometric force (p < 0.05) and peak power output (p < 0.05) deficits in the SSC leg after the exercise protocol were not ameliorated with vitamin D sufficiency. We conclude that vitamin D sufficiency increases the anti-inflammatory cytokine response to muscular injury.     

Comparison between kinematic and kinetic methods for computing the vertical displacement of the center of mass during human hopping at different frequencies

Kinematic and kinetic methods (sacral marker, reconstructed pelvis, segmental analysis, and force platform methods) have been used to calculate the vertical excursion of the center of mass (COM) during movement. In this study we compared the measurement of vertical COM displacement yielded by different methods during able-bodied subjects' hopping at different frequencies (varying between 1.2 and 3.2 Hz). ANOVA revealed a significant interaction between hopping frequency and method (p < 0.001), showing that increasing hopping frequency reduced the differences between methods. A post hoc analysis revealed a significant difference between all methods at the lowest hopping frequency and between the force platform and both the sacral marker and reconstructed pelvis methods at the intermediate hopping frequencies, with differences ranging from 16 to 67 millimeters (all p < 0.05). Results are discussed in view of each methods' limits. We conclude that the segmental analysis and force platform methods can be considered to provide the most accurate results for COM vertical excursion during human hopping in a large range of hopping frequency.     

Vertical stiffness during one-legged hopping with and without using a running-specific prosthesis

Although athletes with unilateral below-the-knee amputations (BKAs) generally use their affected leg, including their prosthesis, as their take-off leg for the long jump, little is known about the spring-like leg behavior and stiffness regulation of the affected leg. The purpose of this study was to investigate vertical stiffness during one-legged hopping in an elite-level long jump athlete with a unilateral BKA. We used the spring-mass model to calculate vertical stiffness, which equals the ratio of maximum vertical ground reaction force to maximum center of mass displacement, while the athlete with a BKA hopped on one leg at a range of frequencies. Then, we compared the vertical stiffness of this athlete to seven non-amputee elite-level long-jumpers. We found that from 1.8 to 3.4 Hz, the vertical stiffness of the unaffected leg for an athlete with a BKA increases with faster hopping frequencies, but the vertical stiffness of the affected leg remains nearly constant across frequencies. The athlete with a BKA attained the desired hopping frequencies at 2.2 and 2.6 Hz, but was unable to match the lowest (1.8 Hz) and two highest frequencies (3.0 and 3.4 Hz) using his affected leg. We also found that at 2.5 Hz, unaffected leg vertical stiffness was 15% greater than affected leg vertical stiffness, and the vertical stiffness of non-amputee long-jumpers was 32% greater than the affected leg vertical stiffness of an athlete with a BKA. The results of the present study suggest that the vertical stiffness regulation strategy of an athlete with a unilateral BKA is not the same in the unaffected versus affected legs, and compared to non-amputees.     

An effective balancing response to lateral perturbations at pelvis level during slow walking requires control in all three planes of motion

In this study we investigated balancing responses to lateral perturbations during slow walking (0.85 m/s). A group of seven healthy individuals walked on an instrumented treadmill while being perturbed at the level of waist at left heel strike in outward and inward lateral directions. Centre of mass (COM) and centre of pressure (COP), rotation of pelvis around vertical axis, step lengths, step widths and step times were assessed. The results have shown that beside control of COP in lateral direction, facilitated by adequate step widths, control of COP in sagittal direction, slowing down movement of COM was present after commencement of lateral perturbations. Sagittal component of COM was significantly retarded as compared to unperturbed walking for both inward (4.32 ± 1.29 cm) and outward (9.75 ± 2.17 cm) perturbations. This was necessary since after an inward perturbation first step length (0.29 ± 0.04 m compared to 0.52 ± 0.02 m in unperturbed walking) and step time (0.45 ± 0.05 s compared to 0.61 ± 0.04 s in unperturbed walking) were shortened while after an outward perturbation first two step lengths (0.36 ± 0.05 m and 0.32 ± 0.11 m compared to 0.52 ± 0.03 m in unperturbed walking) were shortened that needed to be accommodated by the described modulation of COP in sagittal plane. In addition pronounced pelvis rotation assisted in bringing swing leg to new location. The results of this study show that counteracting lateral perturbations at slow walking requires adequate response in all three planes of motion.     

Neuromechanical adaptation induced by jumping on an elastic surface

Jumping on an elastic surface produces a number of sensory and motor adjustments. This effect caused by jumping on the trampoline has been called “trampoline aftereffect”. The objective of the present study was to investigate the neuromuscular response related with this effect. A group of 15 subjects took part in an experimental session, where simultaneous biomechanical and electromyographic (EMG) recordings were performed during the execution of maximal countermovement jumps (CMJs) before and after jumping on an elastic surface. We assessed motor performance (leg stiffness, jump height, peak force, vertical motion of center of mass and stored and returned energy) and EMG activation patterns of the leg muscles. The results showed a significant increase (p ? 0.05) of the RMS EMG of knee extensors during the eccentric phase of the jump performed immediately after the exposure phase to the elastic surface (CMJ₁), and a significant increase (p ? 0.05) in the levels of co-activation of the muscles crossing the ankle joint during the concentric phase of the same jump. Results related with motor performance of CMJ₁ showed a significant increase in the leg stiffness (p ? 0.01) due to a lower vertical motion of center of mass (CoM) (p ? 0.005), a significant decrease in jump height (p ? 0.01), and a significantly smaller stored and returned energy (p ? 0.01). The changes found during the execution of CMJ₁ may result from a mismatch between sensory feedback and the efferent copy.     

Spontaneous whole body rotations and classical dance expertise: How shoulder–hip coordination influences supporting leg displacements

The link between supporting leg stability and individual trunk strategies used during spontaneous whole-body rotations was studied according to visual and kinesthetic imagery styles for classical dancers and untrained female participants. Shoulders–hip angles in the horizontal plane and supporting leg (SL) displacements were analyzed with three-dimensional kinematic at the beginning and end of the four turns, identified according to their SL (left vs. right) and turn direction (clockwise, CW vs. counterclockwise, CCW). To begin a turn in CCW on left SL, all the participants turned shoulders before hips (?25° angle), p < 0.01. Untrained participants yielded the reverse (+30° angle) in CW – their non-preferred turn - whereas dancers maintained their trunk en bloc. In the turn slow down, in their preferred direction all the participants adopted en bloc behavior to avoid imbalance. Dancers kept this pattern in CCW but untrained participants separated shoulders and hips, p < 0.01; on left SL (+20° angle) hips finished before shoulders and on right SL (?25° angle), shoulders finished before hips. Both mental imagery styles and spatial context link reduction of shoulder–hip angle and stability of SL. Daily expertise, not only dance training, facilitates the en bloc shoulder–hip coordination to maintain equilibrium.     

Differences in scapular kinematics and scapulohumeral rhythm during elevation and lowering of the arm between typical children and healthy adults

Scapular kinematics in healthy adults is well described in the literature but little is known on typical children. This study aimed to compare the three-dimensional (3-D) scapular kinematics and scapulohumeral rhythm during the elevation and lowering of the arm in the scapular plane in typical children and healthy adults. Twenty-six healthy adults (35.34 ± 11.65 years, 1.70 ± 0.10 m, 70.00 ± 12.30 kg) and 33 typical children (9.12 ± 1.51 years, 1.40 ± 0.10 m, 35.40 ± 10.45 kg) participated in this study. 3-D scapular kinematics were obtained using an electromagnetic tracking device. The subjects were asked to elevate and lower their arm in the scapular plane. Children showed less scapular protraction compared to adults at 120° during arm elevation, more anterior tilt than adults in the elevation and also at 60°, 90° and 120° during lowering of the arm. Children also showed higher scapulohumeral rhythm during lowering of the arm compared to adults from 90° to 60°. It was also found a low to little correlation between scapular position and age. The study showed small but significant differences in scapular kinematics and scapulohumeral rhythm between children and adults. These results can help clinicians to improve diagnosis and treatment protocols directed to children with dysfunction, as reference values on scapular kinematics in healthy children are also provided in this study.     

Interlimb symmetry of dynamic knee joint stiffness and co-contraction is maintained in early stage knee osteoarthritis

Individuals with knee OA often exhibit greater co-contraction of antagonistic muscle groups surrounding the affected joint which may lead to increases in dynamic joint stiffness. These detrimental changes in the symptomatic limb may also exist in the contralateral limb, thus contributing to its risk of developing knee osteoarthritis. The purpose of this study is to investigate the interlimb symmetry of dynamic knee joint stiffness and muscular co-contraction in knee osteoarthritis.Muscular co-contraction and dynamic knee joint stiffness were assessed in 17 subjects with mild to moderate unilateral medial compartment knee osteoarthritis and 17 healthy control subjects while walking at a controlled speed (1.0 m/s). Paired and independent t-tests determined whether significant differences exist between groups (p < 0.05).There were no significant differences in dynamic joint stiffness or co-contraction between the OA symptomatic and OA contralateral group (p = 0.247, p = 0.874, respectively) or between the OA contralateral and healthy group (p = 0.635, p = 0.078, respectively). There was no significant difference in stiffness between the OA symptomatic and healthy group (p = 0.600); however, there was a slight trend toward enhanced co-contraction in the symptomatic knees compared to the healthy group (p = 0.051).Subjects with mild to moderate knee osteoarthritis maintain symmetric control strategies during gait.     

Endothelial actin and cell stiffness is modulated by substrate stiffness in 2D and 3D

There is a growing appreciation of the profound effects that passive mechanical properties, especially the stiffness of the local environment, can have on cellular functions. Many experiments are conducted in a 2D geometry (i.e., cells grown on top of substrates of varying stiffness), which is a simplification of the 3D environment often experienced by cells in vivo. To determine how matrix dimensionality might modulate the effect of matrix stiffness on actin and cell stiffness, endothelial cells were cultured on top of and within substrates of various stiffnesses. Endothelial cells were cultured within compliant (1.0–1.5 mg/ml, 124±8 to 202±27 Pa) and stiff (3.0 mg/ml, 502±48 Pa) type-I collagen gels. Cells elongated and formed microvascular-like networks in both sets of gels as seen in previous studies. Cells in stiffer gels exhibited more pronounced stress fibers and ～1.5-fold greater staining for actin. As actin is a major determinant of a cell's mechanical properties, we hypothesized that cells in stiff gels will themselves be stiffer. To test this hypothesis, cells were isolated from the gels and their stiffness was assessed using micropipette aspiration. Cells isolated from relatively compliant gels were 1.9-fold more compliant than cells isolated from relatively stiff gels (p<0.05). Similarly, cells cultured on top of 1700 Pa polyacrylamide gels were 2.0-fold more compliant that those cultured on 9000 Pa (p<0.05). These data demonstrate that extracellular substrate stiffness regulates endothelial stiffness in both three- and two-dimensional environments, though the range of stiffnesses that cells respond to vary significantly in different environments.     

Braking characteristics during cutting and pivoting in female soccer players

Most biomechanical studies into changing direction focus on final contact (FC), whilst limited research has examined penultimate contact (PEN). The aim of this study was to explore the kinematic and kinetic differences between PEN and FC of cutting and pivoting in 22 female soccer players (mean ± SD; age: 21 ± 3.1 years, height: 1.68 ± 0.07 m, mass: 58.9 ± 7.3 kg). Furthermore, the study investigated whether horizontal force–time characteristics during PEN were related to peak knee abduction moments during FC. Three dimensional motion analyses of cutting and pivoting on the right leg were performed using Qualysis ‘Proreflex’ infrared cameras (240 Hz). Ground reaction forces (GRF) were collected from two AMTI force platforms (1200 Hz) to examine PEN and FC. Both manoeuvres involved significantly (P < 0.05) greater knee joint flexion angles, peak horizontal GRF, but lower average horizontal GRF during PEN compared to FC. Average horizontal GRF during PEN (R = −0.569, R² = 32%, P = 0.006) and average horizontal GRF ratio (R = 0.466, R² = 22%, P = 0.029) were significantly related to peak knee abduction moments during the FC of cutting and pivoting, respectively. The results indicate PEN during pre-planned changing direction helps reduce loading on the turning leg where there is greater risk of injuries to knee ligaments.