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This study combines in vivo ultrasound measurements of the Vastus Lateralis (VL) and Gastrocnemius Medialis (GM) muscles with electromyographic, kinematic, and kinetic measurements during treadmill running at different speeds (10, 13, and 16 km/h) to better understand the role of muscle and tendon behavior in two functionally different muscle-tendon units. In addition, the force-length and force-velocity relationships of VL and GM were experimentally assessed by combining dynamometry and EMG data with ultrasound measurements. With increasing running speed, the operating length of the fascicles in the stance phase shifted toward smaller lengths in the GM (P < .05; moving down the ascending limb of the F-L relationship) and longer lengths in the VL (P < .05; moving down the descending limb) at all speeds; however, both muscles contracted close to their optimal length L 0 , where isometric force is maximal. Whereas the length of VL SEE did not change as a function of speed, GM SEE lengthened and shortened more at higher speeds. With increasing running speed, the contribution of elastic strain energy to the positive power generated by the MTU increased more for GM (from 0.75 to 1.56 W/kg) than for VL (from 0.62 to 1.02 W/kg). Notwithstanding these differences, these results indicate that, at increasing running speeds, both the VL and GM muscles produce high forces at low contraction velocities, and that the primary function of both muscle-tendon units is to enhance the storage and recovery of elastic strain energy., (© 2020 John Wiley & Sons A/S. Published by John Wiley & Sons Ltd.)

New Findings: What is the central question of this study? Which structures of the medial gastrocnemius muscle-tendon unit contribute to its lengthening during joint rotation and thus receive the stretching stimulus? What is the main finding and its importance? We show, for the first time, that muscle and tendon lengthen in a different manner in children with cerebral palsy compared with typically developing children during a similar amount of muscle-tendon unit lengthening or joint rotation. This indicates possible differences in mechanical muscle and tendon properties attributable to cerebral palsy, which are not evident by assessment of muscle function at the level of a joint., Abstract: Children with cerebral palsy (CP) commonly present with reduced ankle range of motion (ROM) attributable, in part, to changes in mechanical properties of the muscle-tendon unit (MTU). Detailed information about how muscle and tendon interact to contribute to joint rotation is currently lacking but might provide essential information to explain the limited effectiveness of stretching interventions in children with CP. The purpose of this study was to quantify which structures contribute to MTU lengthening and thus receive the stretch during passive ankle joint rotation. Fifteen children with CP (age, in mean ± SD, 11.4 ± 3 years) and 16 typically developing (TD) children (age, in mean ± SD, 10.2 ± 3 years) participated. Ultrasound was combined with motion tracking, joint torque and EMG to record fascicle, muscle and tendon lengthening of the medial gastrocnemius during passive ankle joint rotations over the full ROM and a common ROM. In children with CP, relative to MTU lengthening, muscle and fascicles lengthened less (CP, 50.4% of MTU lengthening; TD, 63% of MTU lengthening; P < 0.04) and tendon lengthened more (CP, 49.6% of MTU lengthening; TD, 37% of MTU lengthening; P < 0.01) regardless of the ROM studied. Differences between groups in the amount of lengthening of the underlying structures during a similar amount of joint rotation and MTU displacement indicate possible differences in tissue mechanical properties attributable to CP, which are not evident by assessment at the level of a joint. These factors should be considered when assessing and treating muscle function in children with CP, for example during stretching exercises, because the muscle might not receive much of the applied lengthening stimulus., (© 2018 The Authors. Experimental Physiology © 2018 The Physiological Society.)

Background: The ability to rapidly generate and transfer muscle force is essential for effective corrective movements in order to prevent a fall. The aim of this study was to establish the muscle and tendon contributions to differences in rate of torque development (RTD) between younger (YM) and older males (OM)., Method: Twenty-eight young males (23.9 years ± 1.1) and 22 old males (68.5 years ± 0.5) were recruited for assessment of Quadriceps Anatomical CSA (ACSA), maximal voluntary contraction (MVC), rate of torque development (RTD), and tendon biomechanical properties. Activation capacity (AC), maximal muscle twitch df/dt) and time to peak EMG amplitude (TTPE) were also assessed., Results: Absolute RTD (aRTD) was lower in OM (577.5 ± 34.6 Nm/s vs 881.7 ± 45.6 Nm/s, p < .0001). RTD remained lower in OM following normalization (nRTD) for muscle ACSA (9.93 ± 0.7 Nm/s/cm2 vs 11.9 ± 0.6 Nm/s/cm2, p < .05). Maximal muscle twitch df/dt (1,086 Nm∙s-1 vs 2,209 Nm∙s-1, p < .0001), TTPE (109.2 ± 8.6ms vs 154.6 ± 16.6 ms, p < .05), and AC (75.8 ± 1.5% vs 80.1 ± 0.9%, p < .01) were all affected in OM. Tendon stiffness was found to be lower in OM (1,222 ± 78.4 N/mm vs 1,771 ± 154.1 N/mm, p < .004). nRTD was significantly correlated with tendon stiffness (R2 = .15)., Conclusion: These observations provide evidence that in absolute terms, a lower RTD in the elderly adults is caused by slower muscle contraction speeds, slower TTPE, reduced ACSA, reduced MVC, and a decrease in tendon stiffness. Once the RTD is normalized to quadriceps ACSA, only MVC and tendon stiffness remain influential. This strongly reinforces the importance of both muscle and tendon characteristics when considering RTD.

New Findings: What is the central question of this study? Can the increased range of motion seen acutely after stretching in children with cerebral palsy be explained by changes in the stiffness of the medial gastrocnemius fascicles? What is the main finding and its importance? We show, for the first time, that passive muscle and tendon properties are not changed acutely after a single bout of stretching in children with cerebral palsy and, therefore, do not contribute to the increase in range of motion. This contradicts common belief and what happens in healthy adults., Abstract: Stretching is often used to increase or maintain the joint range of motion (ROM) in children with cerebral palsy (CP), but the effectiveness of these interventions is limited. Therefore, our aim was to determine the acute changes in muscle-tendon lengthening properties that contribute to increased ROM after a bout of stretching in children with CP. Eleven children with spastic CP [age 12.1 (3 SD) years, 5/6 hemiplegia/diplegia, 7/4 gross motor function classification system level I/II] participated. Each child received three sets of five × 20 s passive, manual static dorsiflexion stretches separated by 30 s rest, with 60 s rest between sets. Before and immediately after stretching, ultrasound was used to measure medial gastrocnemius fascicle lengthening continuously over the full ROM and an individual common ROM pre- to post-stretching. Simultaneously, three-dimensional motion of two marker clusters on the shank and the foot was captured to calculate ankle angle, and ankle joint torque was calculated from manually applied torques and forces on a six degrees-of-freedom load cell. After stretching, the ROM was increased [by 9.9 (12.0) deg, P = 0.005]. Over a ROM common to both pre- and post-measurements, there were no changes in fascicle lengthening or torque. The maximal ankle joint torque tolerated by the participants increased [by 2.9 (2.4) N m, P = 0.003], and at this highest passive torque the maximal fascicle length was 2.8 (2.4) mm greater (P = 0.009) when compared with before stretching. These results indicate that the stiffness of the muscle fascicles in children with CP remains unaltered by an acute bout of stretching., (© 2017 The Authors. Experimental Physiology © 2017 The Physiological Society.)

Purpose: The biarticular rectus femoris (RF), operating on the ascending limb of the force-length curve, produces more force at longer lengths. However, experimental studies consistently report higher knee extension torque when supine (longer RF length) compared to seated (shorter RF length). Incomplete activation in the supine position has been proposed as the reason for this discrepancy, but differences in antagonistic co-activation could also be responsible due to altered hamstrings length. We examined the role of agonist and antagonist muscles in explaining the isometric knee extension torque variation with changes in hip joint angle., Method: Maximum voluntary isometric knee extension torque (joint MVC) was recorded in seated and supine positions from nine healthy males (30.2 ± 7.7 years). Antagonistic torque was estimated using EMG and added to the respective joint MVC (corrected MVC). Submaximal tetanic stimulation quadriceps torque was also recorded., Result: Joint MVC was not different between supine (245 ± 71.8 Nm) and seated (241 ± 69.8 Nm) positions and neither was corrected MVC (257 ± 77.7 and 267 ± 87.0 Nm, respectively). Antagonistic torque was higher when seated (26 ± 20.4 Nm) than when supine (12 ± 7.4 Nm). Tetanic torque was higher when supine (111 ± 31.9 Nm) than when seated (99 ± 27.5 Nm)., Conclusion: Antagonistic co-activation differences between hip positions do not account for the reduced MVC in the supine position. Rather, reduced voluntary knee extensor muscle activation in that position is the major reason for the lower MVC torque when RF is lengthened (hip extended). These findings can assist standardising muscle function assessment and improving musculoskeletal modelling applications.

Aim: To examine the effects of a 16-week resistance exercise training intervention on the speed of ankle and knee strength generation during stair ascent and descent, in people with neuropathy., Methods: A total of 43 people: nine with diabetic peripheral neuropathy, 13 with diabetes but no neuropathy and 21 healthy control subjects ascended and descended a custom-built staircase. The speed at which ankle and knee strength were generated, and muscle activation patterns of the ankle and knee extensor muscles were analysed before and after a 16-week intervention period., Results: Ankle and knee strength generation during both stair ascent and descent were significantly higher after the intervention than before the intervention in the people with diabetes who undertook the resistance exercise intervention (P < 0.05). Although muscle activations were altered by the intervention, there were no observable patterns that underpinned the observed changes., Conclusions: The increased speed of ankle and knee strength generation observed after the intervention would be expected to improve stability during the crucial weight acceptance phase of stair ascent and descent, and ultimately contribute towards reducing the risk of falling. Improvements in muscle strength as a result of the resistance exercise training intervention are likely to be the most influential factor for increasing the speed of strength generation. It is recommended that these exercises could be incorporated into a multi-faceted exercise programme to improve safety in people with diabetes and neuropathy., (© 2015 The Authors. Diabetic Medicine © 2015 Diabetes UK.)

When descending stairs bodyweight becomes supported on a single limb while the forwards-reaching contralateral limb is lowered in order to make contact with the step below. This is associated with lowering of the centre of mass (CoM), which in order to occur in a controlled manner, requires increased ankle and knee joint torque production relative to that in overground walking. We have previously shown that when descending steps or stairs older people operate at a higher proportion of their maximum eccentric capacity and at, or in excess of the maximum passive reference joint range of motion. This suggests they have reduced and/or altered control over their CoM and we hypothesised that this would be associated with alterations in muscle activity patterns and in the CoM vertical acceleration and velocity profiles during both the lowering and landing phases of stair descent. 15 older (mean age 75 years) and 17 young (mean age 25 years) healthy adults descended a 4-step staircase, leading with the right limb on each stair, during which CoM dynamics and electromyographic activity patterns for key lower-limb muscles were assessed. Maximum voluntary eccentric torque generation ability at the knee and ankle was also assessed. Older participants compared to young participants increased muscle co-contraction relative duration at the knee and ankle of the trailing limb so that the limb was stiffened for longer during descent. As a result older participants contacted the step below with a reduced downwards CoM velocity when compared to young participants. Peak downwards and peak upwards CoM acceleration during the descent and landing phases respectively, were also reduced in older adults compared to those in young participants. In contrast, young participants descended quickly onto the step below but arrested their downward CoM velocity sooner following landing; a strategy that was associated with longer relative duration lead-limb plantar flexor activity, increased peak upwards CoM acceleration, and a reduced landing duration. These results suggest that a reduced ability to generate high eccentric torque at the ankle in the forward reaching limb is a major factor for older participants adopting a cautious movement control strategy when descending stairs. The implications of this CoM control strategy on the incidences of falling on stairs are discussed., (Copyright © 2012 Elsevier Inc. All rights reserved.)

The purpose of this study was to determine the effect of dynamometer and joint axis misalignment on measured isometric knee-extension moments using inverse dynamics based on the actual joint kinematic information derived from the real-time X-ray video and to compare the errors when the moments were calculated using measurements from external anatomical surface markers or obtained from the isokinetic dynamometer. Six healthy males participated in this study. They performed isometric contractions at 90° and 20° of knee flexion, gradually increasing to maximum effort. For the calculation of the actual knee-joint moment and the joint moment relative to the knee-joint center, determined using the external marker, two free body diagrams were used of the Cybex arm and the lower leg segment system. In the first free body diagram, the mean center of the circular profiles of the femoral epicondyles was used as the knee-joint center, whereas in the second diagram, the joint center was assumed to coincide with the external marker. Then, the calculated knee-joint moments were compared with those measured by the dynamometer. The results indicate that 1) the actual knee-joint moment was different from the dynamometer recorded moment (difference ranged between 1.9% and 4.3%) and the moment calculated using the skin marker (difference ranged between 2.5% and 3%), and 2) during isometric knee extension, the internal knee angle changed significantly from rest to the maximum contraction state by about 19°. Therefore, these differences cannot be neglected if the moment-knee-joint angle relationship or the muscle mechanical properties, such as length-tension relationship, need to be determined.

Background: Testosterone increases lean mass and may help to counter the changes in muscle architecture associated with sarcopenia. This study was designed to investigate the effects of testosterone replacement therapy on skeletal muscle architecture in intermediate-frail and frail elderly men., Methods: A subgroup of 30 intermediate-frail and frail elderly men (65-89 years) with low to borderline-low testosterone levels were enrolled from a single-center randomized, double-blind placebo-controlled trial. Participants received either a transdermal testosterone (50 mg) or placebo gel daily for 6 months. Architecture (muscle thickness, fascicle length, and pennation angle) of the gastrocnemius medialis muscle was assessed by ultrasound imaging at baseline and after 6 months of treatment., Results: Serum testosterone increased from 11.6 ± 3.5 to 18.0 ± 8.1 nmol/L by 10 days after randomization in the active group (but not the placebo group) and was maintained throughout the treatment period. Testosterone treatment resulted in a preservation of muscle thickness at 6 months while it decreased in the placebo group (effect size 1.4 [95% confidence interval = 0.3-2.5; p = .015]). There was no significant effect of treatment on fascicle length (effect size 1.9 mm [95% confidence interval = -1.2 to 5.0 mm; p = .22]) or pennation angle (effect size 1.2° [95% confidence interval = -1.3 to 3.7°; p = .32])., Conclusions: Testosterone replacement in intermediate-frail and frail elderly men is associated with preservation of muscle thickness. The results suggest that testosterone mitigates sarcopenia by improving muscle tissue to maintain a state of normality in aging men.

Muscle performance is closely related to the architecture and dimensions of the muscle-tendon unit and the effect of maturation on these architectural characteristics in humans is currently unknown. This study determined whether there are differences in musculo-tendinous architecture between adults and children of both sexes. Fascicle length and pennation angle were measured from ultrasound images at three sites along the length of the vastus intermedius, vastus lateralis, vastis medialis and rectus femoris muscles. Muscle volume and muscle-tendon length were measured from magnetic resonance images. Muscle physiological cross-sectional area (PCSA) was calculated as the ratio of muscle volume to optimum fascicle length. Fascicle length was greater in the adult groups than in children (P < 0.05) but pennation angle did not differ between groups (P > 0.05). The ratios between fascicle and muscle length and between fascicle and tendon length were not different (P > 0.05) between adults and children for any quadriceps muscle. Quadriceps volume and PCSA of each muscle were greater in adults than children (P < 0.01) but the relative proportion of each head to the total quadriceps volume was similar in all groups. However, the difference in PCSA between adults and children (men approximately 104% greater than boys, women approximately 57% greater than girls) was greater (P < 0.05) than the difference in fascicle length (men approximately 37% greater than boys, women approximately 10% greater than girls). It is concluded that the fascicle, muscle and tendon lengthen proportionally during maturation, thus the muscle-tendon stiffness and excursion range are likely to be similar in children and adults but the relatively greater increase in PCSA than fascicle length indicates that adult muscles are better designed for force production than children's muscles.

To better understand the effects of pubertal maturation on the contractile properties of skeletal muscle in vivo, the present study investigated whether there are any differences in the specific tension of the quadriceps muscle in 20 adults and 20 prepubertal children of both sexes. Specific tension was calculated as the ratio between the quadriceps tendon force and the sum of the physiological cross-sectional area (PCSA) multiplied by the cosine of the angle of pennation of each head within the quadriceps muscle. The maximal quadriceps tendon force was calculated from the knee extension maximal voluntary contraction (MVC) by accounting for EMG-based estimates of antagonist co-activation, incomplete quadriceps activation using the interpolation twitch technique and magnetic resonance imaging (MRI)-based measurements of the patellar tendon moment arm. The PCSA was calculated as the muscle volume, measured from MRI scans, divided by optimal fascicle length, measured from ultrasound images during MVC at the estimated angle of peak quadriceps muscle force. It was found that the quadriceps tendon force and PCSA of men (11.4 kN, 214 cm(2)) were significantly greater than those of the women (8.7 kN, 152 cm(2); P < 0.01). Both adult groups had greater values than the children (P < 0.01) but there were no differences between boys (5.2 kN, 99 cm(2)) and girls (6.1 kN, 102 cm(2)). Agonist activation was greater in men and women than in girls (P < 0.05), and antagonist activation was greater in men than in boys (P < 0.05). Moment arm length was greater in men than in boys or girls and greater in women than in boys (P < 0.05). The angle of pennation did not differ between the groups in any of the quadriceps heads. The specific tension was similar (P > 0.05) between groups: men, 55 +/- 11 N cm(-2); women, 57.3 +/- 13 N cm(-2); boys, 54 +/- 14 N cm(-2); and girls, 59.8 +/- 15 N cm(-2). These findings indicate that the increased muscle strength with maturation is not due to an increase in the specific tension of muscle; instead, it can be attributed to increases in muscle size, moment arm length and voluntary activation level.

It is not known to what extent the inter-individual variation in human muscle strength is explicable by differences in specific tension. To investigate this, a comprehensive approach was used to determine in vivo specific tension of the quadriceps femoris (QF) muscle (Method 1). Since this is a protracted technique, a simpler procedure was also developed to accurately estimate QF specific tension (Method 2). Method 1 comprised calculating patellar tendon force (F (t)) in 27 young, untrained males, by correcting maximum voluntary contraction (MVC) for antagonist co-activation, voluntary activation and moment arm length. For each component muscle, the physiological cross-sectional area (PCSA) was calculated as volume divided by fascicle length during MVC. Dividing F (t) by the sum of the four PCSAs (each multiplied by the cosine of its pennation angle during MVC) provided QF specific tension. Method 2 was a simplification of Method 1, where QF specific tension was estimated from a single anatomical CSA and vastus lateralis muscle geometry. Using Method 1, the variability in MVC (18%) and specific tension (16%) was similar. Specific tension from Method 1 (30 +/- 5 N cm(-2)) was similar to and correlated with that of Method 2 (29 +/- 5 N cm(-2); R (2) = 0.67; P < 0.05). In conclusion, most of the inter-individual variability in MVC torque remains largely unexplained. Furthermore, a simple method of estimating QF specific tension provided similar values to the comprehensive approach, thereby enabling accurate estimations of QF specific tension where time and resources are limited.

The present study examined the effect of agonist activation and antagonist co-activation on the shape of the knee extension moment-angle relationship in adults and children. Isometric knee extension maximum voluntary contractions (MVCs) were performed at every 5 degrees of knee flexion between 55 degrees and 90 degrees (full extension = 0 degrees) by ten men, ten women, ten boys and ten girls. For each trial, the knee extensors' voluntary activation level was quantified using magnetic stimulation and the level of antagonist co-activation was quantified from their electromyographical activity. Peak MVC moment was greater for men (264 +/- 63 N m) than women (177 +/- 60 N m), and greater for adults than children (boys 78 +/- 17 N m, girls 91 +/- 28 N m) (p < 0.01). The agonistic activation level was greater for adults (approximately 85%) than children (approximately 70%). Similarly, antagonist co-activation was greater for adults than children, but relative to the agonist moment there were no differences between groups (all groups 7-8%). Correcting the peak moment for agonist and antagonist activation levels resulted in moments produced by fully activated agonist muscles of 334 +/- 83, 229 +/- 70, 114.2 +/- 32 and 147 +/- 46 N m, for men, women, boys and girls, respectively. Although correcting for shifts in joint angle during contraction altered the angle of peak moment by approximately 10 degrees (p < 0.01), the peak moment occurred at approximately 60 degrees for all groups. Changes in tendon stiffness, muscle size and architecture, and the pattern of the moment arm-angle relationship may in combination occur so that as children develop and mature into adults the shape of the moment-angle relationship is not altered.

Humans can stand using sensory information solely from the ankle muscles. Muscle length and tension in the calf muscles (gastrocnemius and soleus) are unlikely to signal postural sways on account of balance-related modulation in agonist activity. These facts pose two questions: (1) Which ankle muscles provide the proprioceptive information? (2) Which peripheral mechanism could modulate agonist activity? To address these issues, subjects were asked to stand normally on two force plates. Ultrasound and surface EMG were recorded from the calf and tibialis anterior (TA) muscles. For all nine subjects, changes in muscle length of TA were mainly (84 +/- 9% whole trial duration) orthodoxly correlated with bodily sway (centre of gravity, CoG), i.e. in accordance with passive ankle rotation. When orthodox, TA had the highest correlation with CoG (-0.66 +/- 0.07, deep compartment, P < 0.001). For five subjects, the superficial TA compartment showed counter-intuitive changes in muscle length with CoG, probably due to the flattening of the foot and proximal attachment geometry. Gastrocnemius and soleus were usually (duration 71 +/- 23 and 81 +/- 16%, respectively) active agonists (paradoxically correlated with CoG) but, for short periods of time, they could be orthodox and then presented a moderate correlation (0.38 +/- 0.16 and 0.28 +/- 0.09, respectively) with CoG. Considering the duration and extent to which muscle length is orthodox and correlated with CoG, TA may be a better source of proprioceptive information than the active agonists (soleus and gastrocnemius). Therefore, if a peripheral feedback mechanism modulates agonist activity then reciprocal inhibition acted by TA on the calf muscles is more likely to be effective than the autogenic pathway.

Older people may operate much closer to their maximum capabilities than young adults when ascending stairs due to their lower maximum musculoskeletal capabilities. The purpose of this study was to establish the joint moment and range of motion demands of stair ascent relative to maximum capabilities in elderly and young adults. Fifteen elderly (mean age 75 years) and 17 young adult (mean age 25 years) participants ascended a purpose-built 4-step staircase with force platforms embedded into the steps and kinematic data was acquired using motion capture. Maximum musculoskeletal capabilities were assessed using a dynamometer. This study showed for the first time that stair ascent approaches the joint moment limits at the ankle in both young and older participants ( approximately 90%). One of the most important and novel findings of this study was that elderly people were only capable of meeting the high demands by adopting a number of alternative strategies not observed in young adults: (i) applying the joint moments differently than young adults across the knee and ankle, (ii) translocating energy from the knee to the ankle, thereby enhancing the ankle joint moment upon maximum demand and (iii) by enabling the plantarflexors to act over a more favourable portion of the moment-angle relation upon maximum ankle joint moment demand. The elderly displayed a more cautious strategy to optimize positional stability during stair ascent, by maintaining a smaller separation between the centre of mass and centre of pressure in the frontal plane. It seems that elderly people may meet the demands of unaided stair ascent by adopting a number of alternative strategies to compensate for their reduced musculoskeletal capabilities.

The gastrocnemius medialis (GM) muscle plays an important role in stair negotiation. The aim of the study was to investigate the influence of cadence on GM muscle fascicle behaviour during stair ascent and descent. Ten male subjects (young adults) walked up and down a four-step staircase (with forceplates embedded in the steps) at three velocities (63, 88 and 116 steps/min). GM muscle fascicle length was measured using ultrasonography. In addition, kinematic and kinetic data of the lower legs, and GM electromyography (EMG) were measured. For both ascent and descent, the amount of fascicular shortening, shortening velocity, knee moment, ground reaction force and EMG activity increased monotonically with gait velocity. The ankle moment increased up to 88 steps/min where it reached a plateau. The lack of increase in ankle moment coinciding with further shortening of the fascicles can be explained by an increased shortening of the GM musculotendon complex (MTC), as calculated from the knee and ankle angle changes, between 88 and 116 steps/min only. For descent, the relative instant of maximum shortening, which occurred during touch down, was delayed at higher gait velocities, even to the extent that this event shifted from the double support to the single support phase.

Undisturbed human standing is primarily characterized by sway of the whole body about the ankle joints and is regulated primarily by the calf muscles. Traditionally, in accord with normal ideas of postural control, ankle stiffness, enhanced by spindle mediated muscle stretch reflexes, has been considered to be important for maintaining the upright human stance. This idea predicts that during forward sway, the calf muscles are stretched and the mechanoreflex response enhances muscle activity to maintain posture and balance. Muscle contractile displacement is expected to be positively correlated with bodily sway. However, recent experiments have revealed problems with these ideas. Using a new ultrasound technique for viewing and measuring the dynamic contractile displacements of the calf muscles, it has been shown that calf muscle movement is usually poorly or negatively correlated with bodily sway. The shortening of the contractile tissue during forward sway and vice versa is described as paradoxical muscle movements. This paradoxical muscle movement can be explained by the fact that the Achilles tendon, which transmits the calf muscle force, is compliant in relation the bodily load. There are two main consequences of the compliant Achilles tendon. First, the body is unstable: it cannot be stabilized by intrinsic ankle stiffness alone and thus requires modulation of muscle activity to maintain balance. Second, contractile displacement is mechanically decoupled from bodily sway, which implies that stretch-reflex mechanisms mediated by the calf muscle spindles are unable to successfully modulate muscle activity to maintain balance. This leaves uncertain the postural role of the numerous calf muscle spindles: it is predicted that they signal the effective motor output rather than bodily sway.

Aim: Loss of muscle strength and balance are main characteristics of physical frailty in old age. Postural sway is associated with muscle contractile capacity and to the ability of rapidly correcting ankle joint changes. Thus, resistance training would be expected to improve not only strength but also postural balance., Methods: In this study, age-matched older individuals (69.9+/-1.3 years) were randomly assigned to flywheel (n=12), or weight-lifting (n=12) groups, training the knee extensors thrice weekly for 12 weeks. The hypotheses were that owing to a larger eccentric loading of the knee extensors, flywheel training would result in (a) greater gains in quadriceps strength; (b) greater improvements in balance performance compared with weight-lifting training. Isokinetic dynamometry, B-mode ultrasonography, electromyography, percutaneous muscle stimulation and magnetic resonance imaging were employed to acquire the parameters of interest., Results: Following training, knee extensors peak isokinetic power increased by 28% (P<0.01) in the flywheel group with no change in the weight-lifting group. Adaptations of the gastrocnemius muscle also occurred in both groups. The gastrocnemius characteristic with the highest response to training was tendon stiffness, with increases of 54% and 136% in the weight-lifting and flywheel groups, respectively (P<0.01). The larger increase in tendon stiffness in the flywheel group was associated with an improvement in postural balance (P<0.01)., Conclusion: Quadriceps flywheel loading not only produces a greater increase in power than weight training but its physiological benefits also transfer/overspill to the plantarflexor muscle-tendon unit resulting in a significantly improved balance. These findings support our initial hypotheses.

Aim: To investigate the influence of tendinous and synaptic changes induced by unilateral lower limb suspension (ULLS) on the tendon tap reflex., Methods: Eight young men underwent a 23-day period of ULLS. Muscle cross-sectional area (CSA), torque and electromyographic (EMG) activity of the plantar flexor muscles (normalized to the M wave), Achilles tendon-aponeurosis mechanical properties, soleus (SOL) H and T reflexes and associated peak twitch torques were measured at baseline, after 14 and 23 days of ULLS, and 1 week after resuming ambulatory activity., Results: Significant decreases in muscle CSA (-9%), in maximal voluntary torque (-10%) and in the associated SOL EMG activity (-16%) were found after ULLS (P < 0.05). In addition to a 36% (P < 0.01) decrease in tendon-aponeurosis stiffness, normalized H reflex increased by 35% (P < 0.05). An increase in the slope (28%, P < 0.05) and intercept (85%, P < 0.05) of the T reflex recruitment curve pointed to an increase in the gain and to a decrease in the sensitivity of this reflex, possibly resulting from the decrease in the tendon-aponeurosis stiffness at low forces. Following ULLS, changes in tendinous stiffness correlated with changes in neuromuscular efficiency (peak twitch torque to reflex ratio) at higher tendon tap forces., Conclusion: These findings point out the dual and antagonistic influences of spinal and tendinous adaptations upon the tendon tap reflex in humans under conditions of chronic unloading. These observations have potential implications for the sensitivity of the short-latency Ia stretch response involved in rapid compensatory contractions to unexpected postural perturbations.

Aim: The present study investigated the influence of muscle architectural changes on muscle torque during 3-week unilateral lower limb suspension (ULLS)., Methods: Plantarflexion maximal voluntary contraction (MVC), soleus (SOL), gastrocnemius medialis (GM) and lateralis (GL) muscle volume (VOL), GL fascicle length (L(f)) and pennation angle (theta), physiological cross-sectional area (PCSA), and electromyographic (EMG) activity were assessed in eight healthy men (aged 19 +/- 0 years) after days 14 and 23 of ULLS., Results: After 14 day of ULLS, MVC and SOL EMG decreased (P < 0.05) by 10% and 29%, respectively, but did not further decline between days 14 and 23. SOL, GM and GL muscle VOL decreased by 5%, 6% and 5%, respectively (P < 0.05), on day 14, and by 7% (SOL), 10% (GM) and 6% (GL) on day 23. In GL, theta and L(f) were reduced by 3% (P < 0.05) and 2% (NS), respectively, on day 14, and by 5% (P < 0.05) and 4% (P < 0.05), respectively, on day 23. Consequently, GL PCSA declined by 3% (P < 0.05) on day 14, but did not further decrease on day 23. Similarly, the 7% (P < 0.05) loss in GL force/PCSA observed on day 14 persisted until the end of the unloading period., Conclusion: These findings suggest that rapid muscle architecture remodelling occurs with lower limb unloading in humans, with changes occurring within 14 days of weight bearing removal. These adaptations, mitigating the decline in muscle PCSA, might protect from a larger loss of muscle force.

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

At whole muscle level, the reduction in intrinsic force observed with ageing is probably the result of the combined effect of changes in: (i) muscle architecture, (ii) tendon mechanical properties, (iii) neural drive (reduced agonist and increased antagonist muscles' activity), and (iv) single fibre specific tension. Only recently have alterations in muscle architecture and in tendon mechanical properties been shown to contribute to the reduction in intrinsic muscle force, and tendon stiffness changes play an important role. Of note is the fact that most of these changes may be reversed by 14 weeks of resistive training, for both fibre fascicle length and tendon stiffness were found to be increased by 10% and 64%, respectively. Surprisingly, however, training had no effect on the estimated relative length-tension properties of the muscle, indicating that the effects of increased tendon stiffness and increased fascicle length cancelled out each other. It seems that natural strategies may be in place to ensure that the relative operating range of muscle remains unaltered by changes in physical activity, and perhaps age.

The effect of spastic cerebral palsy on in vivo gastrocnemius muscle fascicle length is not clear. Similarity of fascicle lengths in children with diplegia and typically developing children, but shortening of fascicle lengths in the paretic legs of children with hemiplegia compared with the non-paretic legs, are both reported. In the former case, comparisons were made between fascicle lengths normalized to leg length, whereas in the latter case, absolute fascicle lengths were compared. The inherent assumptions when normalizing fascicle length (measured via ultrasonography) were not validated, raising the possibility that inappropriate normalization contributed to the controversy. We used statistical methods to control the potential confounding effect of leg length on fascicle length, and tested the feasibility of the normalization method for a group of 18 children with diplegia (nine males, nine females; mean age 8y 7mo [SD 3y 11mo], range 2-15y; Gross Motor Function Classification System levels II and III) and 50 typically developing children (20 males, 30 females; mean age 9y 1mo [SD 2y 4mo], range 4-14y). Children with diplegia, as a group, had shorter absolute and normalized fascicle lengths (p<0.05) but we could not refute the appropriateness of the normalization method. Other methodological issues (such as sample characteristics) might have contributed to the apparent controversy between the studies.

To better understand the role of the ankle plantar flexor muscles in stair negotiation, we examined the effects of manipulation of kinematic and kinetic constraints on the behavior of the gastrocnemius medialis (GM) muscle during stair ascent. Ten subjects ascended a four-step staircase at four different step-heights (changing the kinematic constraints): standard (17 cm), 50% decreased, 50% increased and 75% increased. At the standard height, subjects also ascended the stairs wearing a weighted jacket, adding 20% of their body mass (changing the kinetic constraints). During stair ascent, kinematics and kinetics of the lower legs were determined using motion capture and ground reaction force measurements. The GM muscle fascicle length was measured during the task with ultrasonography. The amount of GM muscle fascicle shortening increased with step-height, coinciding with an increase in ankle joint moment. The increase in body mass resulted in an increased ankle joint moment, but the amount of GM muscle fascicle shortening during the lift-off phase did not increase, instead, the fascicles were shorter over the whole stride cycle. Increasing demands of stair ascent, by increasing step-height or body mass, requires higher joint moments. The increased ankle joint moment with increasing demands is, at least in part, produced by the increase in GM muscle fascicle shortening.

Passive stretching is commonly used to increase limb range of movement prior to athletic performance but it is unclear which component of the muscle-tendon unit (MTU) is affected by this procedure. Movement of the myotendinous junction (MTJ) of the gastrocnemius medialis muscle was measured by ultrasonography in eight male participants (20.5 +/- 0.9 years) during a standard stretch in which the ankle was passively dorsiflexed at 1 deg s(-1) from 0 deg (the foot at right angles to the tibia) to the participants' volitional end range of motion (ROM). Passive torque, muscle fascicle length and pennation angle were also measured. Standard stretch measurements were made before (pre-) and after (post-) five passive conditioning stretches. During each conditioning stretch the MTU was taken to the end ROM and held for 1 min. Pre-conditioning the extension of the MTU during stretch was taken up almost equally by muscle and tendon. Following conditioning, ROM increased by 4.6 +/- 1.5 deg (17%) and the passive stiffness of the MTU was reduced (between 20 and 25 deg) by 47% from 16.0 +/- 3.6 to 10.2 +/- 2.0 Nm deg(-1). Distal MTJ displacement (between 0 and 25 deg) increased from 0.92 +/- 0.06 to 1.16 +/- 0.05 cm, accounting for all the additional MTU elongation and indicating that there was no change in tendon properties. Muscle extension pre-conditioning was explicable by change in length and pennation angle of the fascicles but post-conditioning this was not the case suggesting that at least part of the change in muscle with conditioning stretches was due to altered properties of connective tissue.

During human standing, tonic ankle extensor torque is required to support the centre of mass (CoM) forward of the ankles, and dynamic torque modulation is required to maintain unstable balance. Passive mechanisms contribute to both but the extent is controversial. Some groups have revealed a substantial intrinsic stiffness (65-90%) normalized to load stiffness, 'mgh'. Others regard their methodology as unsuitable for the low-frequency conditions of quiet standing and believe the passive contribution to be small (10-15%). Here we applied low-frequency ankle rotations to upright subjects who were supported at the waist allowing the leg muscles to be passive and we report normalized stiffness. The passive calf muscles provided: (i) an extensor torque capable of sustaining unstable balance without tonic activity at a mean CoM-ankle angle of 1.6 deg, (ii) a long range stiffness of 13 +/- 2% and (iii) a short range (< 0.2 deg) stiffness of 67 +/- 8%. Chordal ankle stiffness, derived from the torque versus angle relationship for 7 deg rotations, shows a non-linear decrease (stiffness alpha rotation(-0.33+/-0.04)) from 101 +/- 9% to 19 +/- 5% for rotations of 0.03-7 deg, respectively. Thus, passive stiffness is well adapted for the continuum of postural and movement activity and has a substantial postural role eliminating the need for continuous muscle activity and increasing the unstable time constant of the human inverted pendulum. Ignoring the non-linear dependence of passive stiffness on sway size could lead to serious misinterpretation of experiments using perturbations and sensory manipulations such as eye closure, sway referencing and altered support surfaces.

Using short duration perturbations, previous attempts to measure the intrinsic ankle stiffness during human standing have revealed a substantial stabilizing contribution (65-90% normalized to load stiffness 'mgh'). Others regard this method as unsuitable for the low-frequency conditions of quiet standing and believe the passive contribution to be small (10-15%). This latter view, consistent with a linear Hill-type model, argues that during standing, the contractile portion of the muscle is much less stiff than the tendon. Here, for upright subjects, we settle this issue by measuring the stiffness of the contractile portion of the passive calf muscles using low-frequency ankle rotations. Using ultrasound we tracked the changes in muscle contractile length and partitioned the ankle rotation into contractile and extra-contractile (series elastic) portions. Small ankle rotations of 0.15 and 0.4 deg show a contractile to series elastic stiffness ratio (K(ce)/K(se)) of 12 +/- 9 and 6.3 +/- 10, respectively, with both elements displaying predominantly elastic behaviour. Larger, 7 deg rotations reveal the range of this ratio. It declines in a non-linear way from a high value (K(ce)/K(se) = 18 +/- 11) to a low value (K(ce)/K(se) = 1 +/- 0.4) as rotation increases from 0.1 to 7 deg. There is a marked transition at around 0.5 deg. The series elastic stiffness (K(se)/mgh) remains largely constant (77 +/- 13%) demonstrating the contractile component origin of passive, short range stiffness. The linear Hill-type model does not describe the range-related stiffness relevant to the progression from quiet standing to perturbed balance and movement and can lead to inaccurate predictions regarding human balance.

We tested the hypothesis that the importance of calf muscle-tendon properties for maintaining balance during single-leg stance increases in the absence of visual feedback. Trial duration, centre of pressure displacement normalized for trial duration (nD), electromyographic (EMG) activity of the main ankle plantarflexors and dorsiflexors, and ground reaction forces (F(P)), were measured in 20 younger (aged 18+/-1 years; mean+/-S.E.M.) and 28 older (aged 68+/-1 years) healthy participants during single-leg stance in eyes-open (EO) and eyes-closed (EC) conditions. Plantarflexor muscle strength, activation capacity and tendon stiffness were assessed by dynamometry, electrical stimulation and ultrasonography, respectively. Muscle-tendon characteristics in the older participants were up to 55% (P<0.0001) lower compared with their younger counterparts. Trial duration, F(P), nD and EMG changed in EC compared with EO by 21% and up to approximately 4.6 times (P<0.01) in the two population groups. Multiple linear regression with age and the three muscle-tendon properties showed a substantial increment in EC compared to EO for trial duration (R(2)=0.86 versus R(2)=0.72), but a similarity for nD (R(2)=0.36 versus R(2)=0.33). These results suggest that factors other than the ones that we examined become important when steadiness rather than stance duration is the object of single-leg stance in the absence of vision.

Muscles and tendons are highly adaptive tissues in response to chronic changes in loading and to aging. A remarkable reorganization in muscle architecture occurs in both conditions together with significant alterations in tendon mechanical properties. This review discusses the possible mechanisms underlying these myotendinous changes and the influence thereof on the behavior of the muscle-tendon complex as a whole.

Background: The aim of this study was to investigate the architectural alterations of skeletal muscle following hemiplegic cerebral palsy. If associated with functional and clinical measures of disability, information on muscle architecture could then be used as an objective tool in the assessment of motor disability in these patients., Methods: Ultrasonography was used to assess in vivo the gastrocnemius muscle architecture in the paretic and non-paretic legs of eight children with cerebral palsy., Findings: Fascicle length and muscle thickness at the resting ankle position were reduced in the paretic compared to the non-paretic legs by up to 18% and 20%, respectively (P<0.05), indicating a loss of both in-series and in-parallel sarcomeres in the affected muscles. However, pennation angle was similar (P>0.05) in the two legs., Interpretation: The present results indicate that paresis in hemiplegic cerebral palsy may affect the geometry of skeletal muscle. Further studies are required to examine the relation between muscle architecture, severity of motor disability, and treatment.

The aim of the present study was to establish the behavior of human medial gastrocnemius (GM) muscle fascicles during stair negotiation. Ten healthy male subjects performed normal stair ascent and descent at their own comfortable speed on a standard-dimension four-step staircase with embedded force platforms in each step. Kinematic, kinetic, and electromyographic data of the lower limbs were collected. Real-time ultrasound scanning was used to determine GM muscle fascicle length changes. Musculotendon complex (MTC) length changes were estimated from ankle and knee joint kinematics. The GM muscle was mainly active during the push-off phase in stair ascent, and the muscle fascicles contracted nearly isometrically. The GM muscle was mainly active during the touch-down phase of stair descent where the MTC was lengthened; however, the GM muscle fascicles shortened by approximately 7 mm. These findings show that the behavior and function of GM muscle fascicles in stair negotiation is different from that expected on the basis of length changes of the MTC as derived from joint kinematics.

Activation capacity has traditionally been assessed using the interpolated twitch technique (ITT) and central activation ratio (CAR). However, the quantitative agreement of the two methods and the physiological mechanisms underpinning any possible differences have not been fully elucidated. The aim of this study was to compare and assess the sensitivity of the ITT and CAR to potential errors introduced by (1) evoking inadequate force, by manipulating the number of stimuli, and (2) neglecting differences in series elasticity between conditions, by manipulating joint angle. Ten subjects performed knee extension contractions at 30 degrees and 90 degrees knee-joint angles during which the ITT and CAR methods were applied using 1, 2, 4, and 8 electrical stimuli. Joint angle influenced the ITT outcome with higher values taken at 90 degrees (P < 0.05), while the number of stimuli influenced the CAR outcome with a higher number of stimuli yielding lower values (P < 0.05). For any given joint angle and stimulus number, the CAR method produced higher activation values than the ITT method by 8%-16%. Therefore, in the quantification of voluntary drive with the ITT and CAR methods consideration should be given not only to the number of stimuli applied but also to the effect of series elasticity due to joint-angle differences, since these factors may differently affect the outcome of the calculation, depending on the approach followed.

Bone loss from the paralysed limbs after spinal cord injury (SCI) is well documented. Under physiological conditions, bones are adapted to forces which mainly emerge from muscle pull. After spinal cord injury (SCI), muscles can no longer contract voluntarily and are merely activated during spasms. Based on the Ashworth scale, previous research has suggested that these spasms may mitigate bone losses. We therefore wished to assess muscle forces after SCI with a more direct measure and compare it to measures of bone strength. We hypothesized that the bones in SCI patients would be in relation to the loss of muscle forces. Six male patients with SCI 6.4 (SD 4.3) years earlier and 6 age-matched, able-bodied control subjects were investigated. Bone scans from the right knee were obtained by pQCT. The knee extensor muscles were electrically stimulated via the femoral nerve, isometric knee extension torque was measured and patellar tendon force was estimated. Tendon force upon electrical stimulation in the SCI group was 75% lower than in the control subjects (p<0.01). Volumetric bone mineral density of the patella and of the proximal tibia epiphysis were 50% lower in the SCI group than in the control subjects (p<0.01). Cortical area was lower by 43% in the SCI patients at the proximal tibia metaphysis, and by 33% at the distal femur metaphysis. No group differences were found in volumetric cortical density. Close curvilinear relationships were found between stress and volumetric density for the tibia epiphysis (r(2)=0.90) and for the patella (r(2)=0.91). A weaker correlation with the tendon force was found for the cortical area of the proximal tibia metaphysis (r(2)=0.63), and none for the distal femur metaphysis. These data suggest that, under steady state conditions after SCI, epiphyseal bones are well adapted to the muscular forces. For the metaphysis of the long bones, such an adaptation appears to be less evident. The reason for this remains unclear.

We tested the hypothesis that compromised postural balance in older subjects is associated with changes in calf muscle-tendon physiological and mechanical properties. Trial duration and center of pressure (COP) displacements were measured in 24 younger (aged 24+/-1 yr), 10 middle-aged (aged 46+/-1 yr), and 36 older (aged 68+/-1 yr) healthy subjects under varying levels of postural difficulty. Muscle-tendon characteristics were assessed by dynamometry, twitch superimposition, and ultrasonography. In tandem and single-leg stances, trial duration decreased (

The age-related loss of muscle mass known as senile sarcopenia is one of the main determinants of frailty in old age. Molecular, cellular, nutritional and hormonal mechanisms are at the basis of sarcopenia and are responsible for a progressive deterioration in skeletal muscle size and function. Both at single-fibre and at whole-muscle level, the loss of force exceeds that predicted by the decrease in muscle size. For single fibres, the loss of intrinsic force is mostly due to a loss in myofibrillar protein content. For whole muscle, in addition to changes in neural drive, alterations in muscle architecture and in tendon mechanical properties, exemplified by a reduction in tendon stiffness, have recently been shown to contribute to this phenomenon. Resistance training can, however, cause substantial gains in muscle mass and strength and provides a protective effect against several of the cellular and molecular changes associated with muscle wasting and weakness. In old age, not only muscles but also tendons are highly responsive to training, since an increase in tendon stiffness has been observed after a period of increased loading. Many of the myotendinous factors characterizing ageing can be at least partly reversed by resistance training.

Continuous measurement of contractile length has been traditionally achieved using animal preparations in which the muscle and tendon are exposed. More modern methods, e.g., sonomicroscopy, are still invasive. There is a widely perceived need for a noninvasive, in vivo method of measuring continuous changes of human muscle contractile length. Ultrasonography has been used for several years to measure relatively static, discrete changes in tendon, aponeurosis, and muscle fascicle length. We have recently developed this technique to continuously track changes in muscle contractile length during quiet standing. Here, we present the tracking algorithm and use externally applied perturbations to establish the spatial and temporal resolution of the technique. Subjects maintained a low level of ankle torque while a pneumatic actuator applied rapid, square-pulse ankle rotations of defined magnitude and 0.2-s duration. Tracked changes in gastrocnemius and soleus contractile length follow the temporal profile of the perturbations and scale progressively (5-400 microm) with the size of the ankle rotation (0.03-0.7 degrees ). In a second experiment, we tracked a wire oscillating in water with known peak to peak amplitudes of 1.5 microm to 8 mm. The ultrasound tracking procedure had near 100% accuracy at all amplitudes for frequencies up to 3 Hz and showed attenuation at higher frequencies consistent with an effective sampling frequency of 12 Hz and sampling time of 80 ms. This noninvasive technique is sensitive, without averaging, to changes as small as 1 microm and is suitable for observing neuromotor activity in posture and locomotion.

Senile sarcopenia, the loss of muscle mass associated with aging, is one of the main causes of muscle weakness and reduced locomotor ability in old age. Although this condition is mainly driven by neuropathic processes, nutritional, hormonal and immunological factors, as well as a reduction in physical activity, contribute to this phenomenon. Sarcopenia alone, however, does not fully account for the observed muscle weakness, as the loss of force is greater than that accounted for by the decrease in muscle size. As a consequence, a reduction in the force per unit area, both at single fibre and at whole muscle level, is observed. We recently suggested that at whole muscle level, this reduction in intrinsic force is the result of the combined effect of changes in (1) muscle architecture, (2) tendon mechanical properties, (3) neural drive (reduced agonist and increased antagonist muscle activity) and (4) single fibre-specific tension. Whereas several studies support the role of the last two factors in the loss of intrinsic muscle force with aging, alterations in muscle architecture and in tendon mechanical properties have also been shown to contribute to the above phenomenon. Indeed, sarcopenia of the human plantarflexors, represented by a 25% reduction in muscle volume, was found to be associated with a 10% reduction in fibre fascicle length and 13% reduction in pennation angle. These architectural alterations were accompanied by a 10% decrease in tendon stiffness, attributable to alterations in tendon material properties, as suggested by a 14% decrease in Young's modulus. Most of these changes may be reversed by 14 weeks of resistive training; both fibre fascicle length and tendon stiffness were found to be increased by 10 and 64%, respectively. Surprisingly, however, training had no effect on the estimated relative length-tension properties of the muscle, indicating that the effects of greater tendon stiffness and increased fascicle length cancelled out each other. It seems that natural strategies may be in place to ensure that the relative operating range of muscle remains unaltered by changes in physical activity, in old age.

The purpose of the present study was to investigate the effects of a soccer training session on the balance ability of the players and assess whether the effectiveness of a balance program is affected by its performance before or after the regular soccer training. Thirty-nine soccer players were randomly divided into three subject groups (n=13 each), one control group (C group), one training group that followed a balance program (12 weeks, 3 times per week, 20 min per session) before the regular soccer training (TxB group), and one training group that performed the same balance program after the soccer training (TxA group). Standard testing balance boards and the Biodex Stability System were used to assess balance ability in the C, TxB, and TxA groups at baseline (T0) and after completing the balance program (T12). The same tests and additional isokinetic knee joint moment measurements were carried out in the TxB and TxA groups pre- and post-soccer training. Two main results were obtained: (1) No differences (p>0.05) were found in balance ability and knee joint moment production between pre- and post-soccer training. (2) The balance program increased (p<0.01) the balance ability in the TxB and TxA groups, and the improvement in the TxA group was greater (p<0.05) than that in the TxB group post-soccer training. Result (1) is in contrast to the notion of a link between fatigue induced by a soccer training session or game and injury caused by impaired balance, and result (2) has implications for athletic training and rehabilitation.

In the present study, we measured the contraction-induced shortening (dL) of individual synergistic human muscles in a repeated motor task to assess their contractile behaviour. Ultrasonography was used to obtain dL measurements in the gastrocnemius (GS) and soleus (SOL) muscles of six men performing 11 consecutive isometric plantarflexions. Contractions 1 and 11 were performed with maximal effort, and contractions 2-4, 5-7 and 8-10 were performed with efforts generating 50, 70 and 90%, respectively, of the plantarflexion moment produced in contraction 1. In contractions 5-10, the SOL muscle dL was similar (p > 0.05) to that produced in contraction 1 (approximately 6 mm), indicating that the SOL muscle became fully activated at 70% of the maximum plantarflexion moment. The GS muscle dL in contractions 10 and 11 exceeded by approximately 0.5 mm (p < 0.05) and 1.3 mm (p < 0.01), respectively, that generated in contraction 1 (approximately 10 mm), despite evidence obtained by superimposed stimulation that contraction 1 was produced with full motor unit activation. The consequent paradox that the GS muscle would produce in contractions 10 and 11 a greater activation and therefore more force than its actual potential is resolved when considering the interaction between the time-dependent tensile response of tendon and the performance of muscle as dictated by the sliding filament mechanism of contraction.

Muscles of individuals with a spinal cord injury (SCI) exhibit an unexpected leftward shift in the force (torque)-frequency relationship. We investigated whether differences in torque-angle relationships between SCI and able-bodied control muscles could explain this shift. Electrically stimulated knee-extensor contractions were obtained at knee flexion angles of between 30 degrees and 90 degrees. Torque-frequency relationships were obtained at 30 degrees, 90 degrees, and optimum angle. Optimum angle was not different between groups but SCI-normalized torques were lower at the extreme angles. At all angles, SCI muscles produced higher relative torques at low stimulation frequencies. Thus, there was no evidence of a consistent change in the length of paralyzed SCI muscles, and the anomalous leftward shift in the torque-frequency relationship was not the result of testing the muscle at a relatively long length. The results provide valuable information about muscle changes occurring in various neurological disorders.

It has been widely assumed for nearly a century, that postural muscles operate in a spring-like manner and that muscle length signals joint angle (the mechano-reflex mechanism). Here we employ automated analysis of ultrasound images to resolve calf muscle (soleus and gastrocnemius) length changes as small as 10 mum in standing subjects. Previously, we have used balancing of a real inverted pendulum to make predictions about human standing. Here we test and confirm these predictions on 10 subjects standing quietly. We show that on average the calf muscles are actively adjusted 2.6 times per second and 2.8 times per unidirectional sway of the body centre of mass (CoM). These alternating, small (30-300 microm) movements provide impulsive, ballistic regulation of CoM movement. The timing and pattern of these adjustments are consistent with multisensory integration of all information regarding motion of the CoM, pattern recognition, prediction and planning using internal models and are not consistent with control solely by local reflexes. Because the system is unstable, errors in stabilization provide a perturbation which grows into a sway which has to be reacted to and corrected. Sagittal sway results from this impulsive control of calf muscle activity rather than internal sources (e.g. the heart, breathing). This process is quite unlike the mechano-reflex paradigm. We suggest that standing is a skilled, trial and error activity that improves with experience and is automated (possibly by the cerebellum). These results complement and extend our recent demonstration that paradoxical muscle movements are the norm in human standing.

In humans, during standing the calf muscles soleus and gastrocnemius actively prevent forward toppling about the ankles. It has been generally assumed that these postural muscles behave like springs with dynamic stiffness reflecting their mechanical properties, reflex gain including higher derivatives, and central control. Here, for the first time, we have used an ultrasound scanner and automated image analysis to record the tiny muscular movements occurring in normal standing. This new, non-invasive technique resolves changes in muscle length as small as 10 mum without disturbing the standing process. This technical achievement has allowed us to test the long-established mechano-reflex, muscle spring hypothesis that muscle length changes in a spring-like way during sway of the body. Our results contradict that hypothesis. Muscle length changes in a non-spring-like manner: on average, shortening during forward sway and lengthening during backwards sway (paradoxical movements). This counter-intuitive result is a consequence of the fact that calf muscles generate tension through a series elastic component (SEC, Achilles tendon and foot) which limits maximal ankle stiffness to 92 +/- 20% of that required to balance the body. Paradoxical movements cannot be generated by stretch reflexes with constant intrafusal drive but might be produced by reflex coupling of extrafusal (alpha) and intrafusal (beta, gamma) drive or by positive force feedback. Standing requires the predictive ability to produce the observed muscle movements preceded (110 +/- 50 ms) by corresponding changes in integrated EMG signal. We suggest higher level anticipatory control is more plausible.

Data are scarce relating to the plasticity with strength training of dynamic muscle performance in older humans. Hence, we investigated alterations in the torque-velocity relation with strength training in old age, and their origin. Knee extension and leg-press exercises were performed three times per week for 14 weeks. Maximal isokinetic knee extension torque was assessed during concentric and eccentric muscle actions. Agonist-antagonist muscle activation was assessed using electromyography. Vastus lateralis muscle architecture was examined in vivo using ultrasonography. Training increased concentric torque by 22-37% (P < 0.01), but failed to alter eccentric torque (P > 0.05). Increased agonist muscle activation, increased muscle fascicle lengths, and greater elastic energy recovered from tendinous structures may explain the adaptations during concentric actions, whereas the failure of eccentric torque to increase might be explained by the preservation of eccentric force with aging and an underloading of the eccentric movement phase during training. These findings may have important implications for dynamic muscle performance in old age.

This study investigated changes in elderly muscle joint angle-torque relation induced by resistance training. Older adults were assigned to either training (n = 9, age 74.3 +/- 3.5 years; mean +/-s.d.) or to control groups (n = 9, age 67.1 +/- 2 years). Leg-extension and leg-press exercises were performed three times per week for 14 weeks. Maximal isometric knee extension torque was measured across the knee joint angle range of movement. Vastus lateralis muscle architecture was examined in vivo using ultrasonography. The vastus lateralis muscle fascicle force was estimated from the measured joint torque, enabling construction of the fascicle length-force relation. Electromyographic (EMG) activity was measured from representative agonist and antagonist muscles. Training altered the angle-torque relation: (a) displacing it by 9-31% towards higher torque values (P < 0.05); and (b) shifting the optimal angle from 70 deg (corresponding torque: 121.4 +/- 61 N m) before to 60 deg (134.2 +/- 57.2 N m; P < 0.05) after training. Training also altered the fascicle length-force relation: (a) displacing it by 11-35% towards higher force values; and (b) shifting the optimal fascicle length from 83.7 +/- 8 mm (corresponding force: 847.9 +/- 365.3 N) before to 93.2 +/- 12.5 mm (939.3 +/- 347.8 N; P < 0.01) after training. The upward displacement of the angle-torque relation was mainly due to a training-induced increase in agonist activation, whilst the shift in the optimal angle was associated with changes in muscle-tendon properties.

In the present work, a generic model for the prediction of moment-angle characteristics in individual human skeletal muscles is presented. The model's prediction is based on the equation M = V x Lo(-1)sigma c cos phi x d, where M, V, and Lo are the moment-generating potential of the muscle, the muscle volume and the optimal muscle fibre length, respectively, and sigma, phi and d are the stress-generating potential of the muscle fibres, their pennation angle and the tendon moment arm length, respectively, at any given joint angle. The input parameters V, Lo, sigma, phi and d can be measured or derived mechanistically. This eliminates the common problem of the necessity to estimate one or more of the input parameters in the model by fitting its outcome to experimental results often inappropriate for the function modelled. The model's output was validated by comparisons with the moment-angle characteristics of the gastrocnemius (GS) and tibialis anterior (TA) muscles in six men, determined experimentally using voluntary contractions at several combinations of ankle and knee joint angles for the GS muscle and electrical stimulation for the TA muscle. Although the model predicted realistically the pattern of moment-angle relationship in both muscles, it consistently overestimated the GS muscle M and consistently underestimated the TA muscle M, with the difference gradually increasing from dorsiflexion to plantarflexion in both cases. The average difference between predicted and measured M was 14% for the GS muscle and 10% for the TA muscle. Approximating the muscle fibres as a single sarcomere in both muscles and failing to achieve complete TA muscle activation by electrical stimulation may largely explain the differences between theory and experiment.

Neuropathic, metabolic, hormonal, nutritional and immunological factors contribute to the development of sarcopenia. This loss of muscle mass associated with ageing, is a main cause of muscle weakness, but the loss of muscle strength typically exceeds that of muscle size, with a resulting decrease in force per unit of muscle cross-sectional area. Recent evidence suggests that, in addition to a reduction in neural drive and in fibre specific tension, changes in muscle architecture contribute significantly to the loss of muscle force through alterations in muscle mechanical properties. Older muscle, however, maintains a high degree of plasticity in response to increased loading since considerable hypertrophy and a reversal of the alterations in muscle architecture associated with ageing are observed with resistive training.

In human standing, gravity causes forward toppling about the ankle joint which is prevented by activity in the soleus and gastrocnemius muscles. It has long been assumed that when people sway forwards the calf muscles are stretched and conversely that they shorten with backward sway. Consequently, for many years, two explanations for standing stabilization have flourished. First, tonic muscle activity itself may generate adequate intrinsic ankle stiffness. Second, if intrinsic ankle stiffness is inadequate, the resistance to stretch of the calf muscles may be augmented by stretch reflexes or by central control. These explanations require that the passive tissue (Achilles' tendon, foot) transmitting the calf muscle tension is stiff. However, our recent measurements have indicated that this passive tissue is not stiff during standing. Accordingly, we predicted a counterintuitive mode of control where the muscles and body must, on average, move in opposite directions (paradoxical movements). Here we use dynamic ultrasound imaging in vivo with novel automated tracking of muscle length to test our hypothesis. We show that soleus and gastrocnemius do indeed move paradoxically, shortening when the body sways forward and lengthening when the body returns. This confirms that intrinsic ankle stiffness is too low to stabilize human standing. Moreover, it shows that the increase in active tension is associated with muscle shortening. This pattern cannot be produced by muscle stretch reflexes and can only arise from the anticipatory neural control of muscle length that is necessary for balance.

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