The epidemiology and clinical manifestations of hamstring muscle and plantar foot flexor shortening

A population of 920 healthy children was studied with the aim of assessing the incidence of hamstring muscle and plantar foot flexor tightness, and to correlate such symptoms with gait, posture, and low back discomfort or pain. Special attention was paid to the popliteal angle and dorsal foot flexion. The borderline values for the popliteal angle in the following age groups were, boys: 3 to 5 years, 40^o; 6 to 15 years, 50^o; and 16 to 19 years, 40^o; girls: 3 to 5 years, 30^o; 6 to 14 years, 45^o; 15 to 19 years, 30^oo. The borderline values for dorsal foot flexion in the following age groups were 3 to 4 years, 7^o; 5 to 13 years, 10^o; and 14 to 19 years, 5^o. The results obtained indicate a natural increase in hamstring tightness, particularly shortly before the pubertal growth spurt. This seems to be linked with the natural evolution of lumbar lordosis and pelvic tilt. When hamstring tightness surpassed borderline values, dorsiflexion and lumbar lordosis decreased leading to postural deformities, bending-forward deficit, discomfort when sitting, and a shambling gait.     

Continuum of reflex excitability in hemiplegia: influence of muscle length and muscular transformation after heel-cord lengthening and immobilization on the pathophysiology of spasticity and clonus

The electromyographic (EMG) responses to tendon taps at the ankle and ensuing muscular twitch forces and temporal parameters were studied at varying angles across the joint range in 18 children, aged 3 to 14.9 years, with congenital hemiparetic cerebral palsy and 22 healthy (control) children, aged 3 to 13.6 years. Those subjects with hemiparesis were community ambulators without assistance. In all subjects, passive muscle stretch caused a waxing of the reflex EMG and twitch force near neutral (with the sole of the foot at right angles to the tibia) and a diminution of these with further dorsiflexion. Twitch times increased with each dorsiflexing increment, being slowest at maximum dorsiflexion and fastest at the resting plantarflexion angle. Heterogeneity of the hemiparetic-limb data is evident when compared with data of non-paretic and unaffected limbs, with clear differences in the clonic (fast twitch) as opposed to non-clonic (slow twitch) muscles. In four cases with clinical clonus, clonus frequency was reduced by passive dorsiflexion. Plaster immobilization for 1 month produced clonus which was previously absent in one subject, and caused a fast-twitch phenotype to emerge in two subjects. Follow-up after heel-cord lengthening in one subject showed that clonus frequency diminished from 9 to 3 Hz with slowing and strengthening of muscle-twitch phenotype. Short- and long-term peripheral manipulations appear to regulate neuromuscular excitability according to whether muscles are loaded or unloaded. Although damage to the nervous system provides the setting for reflex excitability, the data suggest that the muscle length (which specifies the joint angle) and the muscle-twitch phenotype of any given limb for any given case appear to dictate the actual speed and strength of reflex muscle-twitch and clonus profiles. This study illustrates how peripheral manipulations of muscles and tendons may alter the expression of what have hitherto been considered as exclusively central phenomena.     

Inhibition from the plantar nerve to soleus muscle during the stance phase of walking

The purpose of this study is to investigate the effect and the circuit from the branch of tibial (plantar) nerve to soleus muscle and its modulation during walking in humans. Stimulation of the plantar nerve produced short latency inhibition of soleus EMG activity and the H-reflex in humans. The threshold of afferent fibers was lower than that of motor fibers. This inhibition did not converge to disynaptic reciprocal Ia inhibition nor did inhibition from the cutaneous nerve of the big toe, but to Ib inhibition from the medial gastrocnemius nerve. The inhibitory pathway from the plantar nerve therefore is considered to include Ib inhibitory interneurones. Modulation of the inhibition was investigated during walking. Less EMG depression after plantar nerve stimulation occurred in the stance phase of walking than for tonic or dynamic plantar flexion at similar background EMG activity level. The inhibition of the soleus H-reflex after plantar nerve stimulation was also decreased during the stance phase. For investigating the influence of load on the inhibition from the plantar nerve, more EMG depression occurred in the stance phase with body unloading. Similar findings were observed in Ib inhibition from the medial gastrocnemius nerve, but not in disynaptic reciprocal Ia inhibition to soleus muscle. It is concluded that transmission of inhibition from the plantar nerve to soleus muscle is modulated during walking. It would minimize this inhibition during the stance phase of walking and might enhance soleus muscle activity via this reflex pathway for the support of weight.     

Increased H‐reflex excitability is not accompanied by changes in neural drive following 24 days of unilateral lower limb suspension

The purpose of this study was to determine whether the gain in soleus H‐reflex excitability induced by unilateral lower limb suspension (ULLS) is associated with changes in neural drive to the plantar flexor muscles. Six male subjects (23 ± 2 years, 187 ± 7 cm, 79 ± 9 kg) underwent 24 days of ULLS of the dominant limb. Plantar flexor maximal voluntary contraction (MVC) torque, activation capacity (twitch interpolation), soleus maximal electromyographic (EMG) activity, Hoffman (H)‐reflex, and the first volitional (V) wave normalized to the compound muscle action potential (M‐wave) were quantified before and after ULLS. Following ULLS, MVC torque decreased by 15% (P < 0.05). However, neither activation capacity nor EMG activity was significantly altered after the suspension. The V‐wave remained unchanged consistently after ULLS, whereas the H‐reflex increased significantly (+20%). Furthermore, there was no significant relationship between changes in H‐reflex and V‐wave over the ULLS period. These findings indicate that 24 days of ULLS can result in a substantial reduction of muscle strength without any apparent change in voluntary activation capacity. H‐reflex and V‐wave findings suggest that the spinal adaptations that underlie the unloading‐induced increase in resting soleus H‐reflex excitability did not significantly affect the efferent motor output to the plantar flexor muscles. Muscle Nerve, 2009     

Sex differences in spinal excitability during observation of bipedal locomotion

This study investigated whether there are sex differences in the spinal excitability of the human mirror-neuron system. We measured the modulation of spinal excitability, elicited by Hoffmann reflex in the left plantar flexor muscle (soleus), when women and men participants observed videos of bipedal heel-stepping (plantar dorsiflexion), standing still, and bipedal toe-stepping (plantar flexion). Men and women were similarly divided in their sex judgments of the observed legs. Our results indicate that whereas both men and women mimicked spinal excitability to the observed bipedal step, only women participants produced stronger modulation of spinal excitability. These findings demonstrate that the human mirror-neuron system exhibits sex differences in spinal excitability.     

Modulation of spinal excitability during observation of bipedal locomotion

This study investigated whether a mirror mechanism exists for bipedal locomotion. We employed the soleus (plantar flexor) Hoffman reflex to investigate corticospinal excitability at the spinal level during observation of bipedal locomotion. The differential amplitude modulation of the left soleus Hoffman reflex during observation of bipedal heel-stepping (plantar dorsiflexion) (324+/-53 microV), standing still (383+/-60 microV), and bipedal toe-stepping (plantar flexion) (419+/-53 microV) reached significance (P < 0.05). The observation of bipedal toe-stepping produced a greater increase in spinal excitability than the observation of bipedal heel-stepping. These findings support the suggestion that there is a mirror mechanism for bipedal locomotion and they demonstrate that spinal excitability for observation of bipedal locomotion mirrors that for execution of bipedal locomotion.     

Attenuation of the effect of remote muscle contraction on the soleus H-reflex during plantar flexion.

OBJECTIVE: We investigated to what extent the facilitation of the soleus (Sol) Hoffmann (H-) reflex during a phasic voluntary wrist flexion (Jendrássik maneuver, JM) can be modulated by graded plantar flexion force and conditioning wrist flexion force. METHODS: The subjects were asked to perform phasic wrist flexion under a reaction time condition. Sol H-reflex was evoked by stimulating the right tibial nerve at various time intervals (50-400ms) after the 'Go' signal for initiating JM while the ankle was at rest and while plantarflexing. The level of tonic plantar flexion force (isometric contraction of 10, 20 and 30% of maximal EMG) and conditioning wrist flexion (isometric contraction of 30, 50 and 80% of maximum voluntary contraction) during JM was graded systematically. RESULTS: Although JM facilitation could be seen 80-120ms after the flexor carpi radialis (FCR) EMG onset even while plantarflexing, the magnitude of JM facilitation under plantar flexion was significantly decreased compared to that at rest. The degree of decrease in JM facilitation did not depend on the level of plantar flexion force. In contrast, the degree of JM facilitation was proportional to the level of wrist flexion force while the ankle was at rest and while plantarflexing, though the amount of JM facilitation significantly decreased while plantarflexing. CONCLUSIONS: JM facilitation of Sol H-reflex is decreased while performing tonic voluntary contraction of the homonymous muscle. The degree of decrease in JM facilitation is independent of the level of homonymous muscle contraction, but depends on the level of remote FCR contraction. In clinical application, when we intend to elicit a maximum stretch reflex by JM, full relaxation of homonymous muscle should be carefully confirmed. SIGNIFICANCE: Our results provide evidence for better understanding of the features of JM and insight into its clinical application.     

Control of soleus motoneuron excitability during muscle stretch in man.

The relative contributions of intramuscular and extramuscular receptors to changes in the reflex excitability of soleus motoneurons, following muscle stretch, have been studied in man. It was found that reflex excitability was decreased by muscle stretch. The extent of the decrease was related to the amount of stretch, irrespective of whether the latter was produced by dorsiflexion of the ankle or by depression of the Achilles tendon with the ankle joint fixed. The results were unaffected by anaesthesia of the skin. It would appear that neither joint receptors nor cutaneous mechanoreceptors contribute significantly to the decrease in reflex excitability during ankle dorsiflexion and that the intramuscular receptors are mainly responsible for the effects observed.     

The action of plantar pressure on flexion reflex pathways in the isolated human spinal cord.

OBJECTIVE: To investigate the conditioning effects of plantar pressure on flexion reflex excitability in patients with motor complete spinal cord injury (SCI). METHODS: In five motor complete SCI subjects, the non-nociceptive flexion reflex was evoked via electrical stimulation of the right sural nerve and was recorded from the ipsilateral tibialis anterior muscle. Pressure ranging from 25 to 80kPa was applied to the metatarsal heads through an adjustable platform incorporated into a foot rest and a comparison of the reflex size made between control conditions and during pressure application. RESULTS: In all subjects, a significant depression of the long latency flexion reflex was observed when pressure was applied to the foot sole. The short latency flexion reflex appearing at latencies less than 100ms was absent in all patients. CONCLUSIONS: The results demonstrate that flexion reflex excitability in the isolated human spinal cord can be modulated by adequate activation of plantar mechanoreceptors. SIGNIFICANCE: Activation of plantar mechanoreceptors is a feature of normal standing and walking. Rehabilitation for standing and walking in SCI commonly uses body weight support based protocols. The strong inhibitory actions of plantar pressure on reflex pathways in the isolated human spinal cord suggest that sensory feedback from the foot sole may be an important factor in successful rehabilitation of standing and stepping in SCI patients.     

Quantitative evaluation of electromyogram activity in rat extensor and flexor muscles immobilized at different lengths

Because immobilization of muscles in the "long" position mitigates the effects of inactivity and rapid wasting occurs when muscles are immobilized in the "short" position, a study was made of the EMG activity in the soleus (SOL)--an extensor muscle--and the tibialis anterior (TA)--a flexor muscle--in order to clarify the possible role of muscle function in modifying the course of disuse atrophy. EMG activity was recorded in the SOL and TA muscles in adult rats in which the ankle had been immobilized in a plaster cast either in plantar flexion or dorsiflexion. The number of action potentials per minute in samples of the EMG activity from control and immobilized muscles was assessed before, for 10 days during immobilization, and up to 9 days after removal of the cast. Immobilization in the short position (plantar flexion) led to a dramatic reduction in the EMG activity of the SOL (to 10% of the control). On the other hand, fixation of the SOL in the long position was without effect upon resting EMG activity. In the TA, EMG activity was exclusively phasic in character and corresponded to about 3% of that of the SOL. Neither the fixation of the ankle in plantar flexion nor dorsiflexion had any appreciable effect upon EMG activity in the TA. We conclude that, because immobilization in the lengthened position does not increase EMG activity in either extensor or flexor muscles, passive stretch appears to be the factor mainly responsible for mitigating the effects of disuse in this situation. On the other hand, when a typical extensor muscle (SOL) is immobilized in the shortened position and undergoes rapid wasting, an accessory role of decreased activity cannot be excluded.     

The influence of experimental muscle pain on the human soleus stretch reflex during sitting and walking.

OBJECTIVES: The stretch reflex is functionally important during human locomotion. Muscle pain has been found to increase the stretch reflex amplitude during sitting, possibly due to an altered fusimotor drive. To further study the importance of altered fusimotor activity due to muscle pain we investigated the combined effect of muscle pain and motor task on the soleus stretch reflex. METHODS: Stretch reflexes were elicited before, during and after experimentally induced muscle pain in soleus (i.m. infusion of 6% saline) in 3 experiments: (1) in the relaxed soleus muscle and before, during and after an isometric ramp contraction (500 ms, 0-10 Nm), (2) at 3 different time periods during walking, and (3) at matched pain intensity and soleus activity during sitting and walking. RESULTS: Infusion of hypertonic saline into the soleus muscle caused a significant facilitated stretch reflex in the relaxed muscle (P<0.01), but not during walking or during sitting and walking at matched soleus EMG and matched pain levels. The infusion of isotonic saline (non-painful) did not cause any changes (P = 0.75). CONCLUSIONS: The main findings of the present study were that experimental muscle pain facilitated the stretch reflex during pain in the relaxed muscle, but caused no changes in stretch reflex amplitude during sitting and walking at higher "functional" background EMG levels.     

Hyperactive stretch reflexes, co-contraction, and muscle weakness in children with cerebral palsy

The aim of this study was to examine the repeatability of and relationships among spasticity, co-contraction of agonist–antagonist, and muscle strength in children with cerebral palsy (CP). Eight children with spastic diplegic CP (five males, three females; Gross Motor Function Classification System [GMFCS] Levels I–III; mean age 10y 2mo [SD 2y 9mo], range 6–13y) and nine children in a comparison group (six males, three females; mean age 8y 10mo [SD 2y 4mo], range 6y to 12y 6mo) were assessed twice to examine repeatability of Composite Spasticity Scale, soleus stretch reflexes, electromyography (EMG) co-contraction ratio, and torque recorded during maximal isometric voluntary contraction of ankle dorsiflexors and plantarflexors. Sixty-one children with spastic CP, (54 diplegic, seven hemiplegic; 32 males, 29 females; GMFCS levels I–III; mean age 10y 8mo [SD 2y 9mo], range 6–15y) were then assessed to delineate possible correlations among these measures. Intraclass correlation coefficients (0.78–0.97) showed high data repeatability in both groups. Children with spastic CP demonstrated significantly larger soleus stretch reflex/M-response areas smaller torques, but larger EMG co-contraction ratios during both voluntary dorsiflexion and plantarflexion (all p <0.05). Children with spastic CP who had larger soleus stretch reflex/M-response areas demonstrated larger plantarflexion co-contraction ratio ( r = 0.28), and produced smaller plantarflexion and dorsiflexion torques ( r = –0.48 and –0.27 respectively). However, no correlation was noted between soleus stretch reflex and clinical spasticity. Our findings demonstrated that hyperactive soleus stretch reflex affected torque production of ankle muscles. Moreover, the severity of spasticity may not be fully described by either stretch reflex or tone measure alone.     

The effect of digital nerve stimulation on recruitment order of motor units in the first deep lumbrical muscle of the cat

The normal recruitment order of EMG spikes of the first deep lumbrical muscle of the cat's hindpaw, usually seen during cortical stimulation, pad pinch and weak plantar nerve stimulation, was temporarily reversed after stimulation of the medial digital nerve of the foot at 50 Hz for 2 min, and normal order was recovered in 2 to 10 min. The longer the period of stimulation of the medial digital nerve was, the longer the time for recovery. min. After prolonged medial digital nerve stimulation, EMG responses to pad pinch and plantar nerve stimulation were facilitated. Combination of some stimuli (e.g. cortical stimulation and plantar nerve stimulation, pad pinch and plantar nerve stimulation, plantar nerve and medial digital nerve stimulation) also produced reversal of recruitment order during the period of stimulation.A functionally single motor nerve fiber to the first deep lumbrical muscle was isolated from the motor nerve axons in L7 and S1 of the spinal cord of cats, and physiological properties of pairs of motor units whose recruitment order was temporarily altered by plantar nerve stimulation after prolonged stimulation of the medial digital nerve of a foot were examined. Motor units with large action potentials were more facilitated than motor units with small action potentials after prolonged stimulation of the medial digital nerve. The former motor units showed fast contraction time, large twitch tension, low resistance to fatigue and presence of sag-behavior. The latter motor units showed slow contraction time, small twitch tension and high resistance to fatigue.High threshold motor units with middle-sized surface EMG records were recruited after motor units with large potentials had been recruited; motor units with middle-size action potentials were beta motor axons branching to both intrafusal and extrafusal muscle fibers. The beta motor axons also showed fast conduction velocity and the presence of sag-behavior.     

Antagonist inhibition during rest and precontraction.

The excitability of antagonist soleus motoneurons was tested during fast voluntary contractions of tibialis anterior (TA). Contractions of TA started either from zero or higher levels of tonic contraction of ankle extensors. Results showed that H reflex depression under 3 different conditions: 25% (R25) or 50% (R50) of the maximal isometric dorsiflexion from the resting state and the sequential isometric response (SW) appeared about 20-40 msec prior to the EMG onset of TA. In particular, H reflex depression was clear in the SW response, and the stronger the prior contraction of extensors the greater the H reflex depression. There was a significant difference in the integrated EMG between R25 and SW but none between R50 and SW. The different amounts of inhibition found for dorsiflexion from the resting state and sequential movement is most probably explained by presynaptic inhibition. If presynaptic inhibition were increased throughout the sequential movement, in comparison with the resting state, change in the H reflex gain would occur.     

Effects of initial conditions on the Hoffman reflex

The Hoffmann reflex is a monosynaptic reflex elicited by electrical stimulation of afferent nerve fibres. The amplitude of the reflex response may be measured both by EMG recording from the muscle and by sensing the muscle twitch. These two effects are dependent, not only on the amplitude of the stimulus, but on the state of excitability of the afferent-efferent synaptic pools and on the mechanical state of the muscle. Voluntary control of the stimulated muscle influences these conditions but the effects on the EMG are quite different from those on the twitch. This paper discusses these effects under isometric conditions.     

Parallel facilitatory reflex pathways from the foot and hip to flexors and extensors in the injured human spinal cord

Spinal integration of sensory signals associated with hip position, muscle loading, and cutaneous sensation of the foot contributes to movement regulation. The exact interactive effects of these sensory signals under controlled dynamic conditions are unknown. The purpose of the present study was to establish the effects of combined plantar cutaneous afferent excitation and hip movement on the Hoffmann (H) and flexion reflexes in people with a spinal cord injury (SCI). The flexion and H-reflexes were elicited through stimulation of the right sural (at non-nociceptive levels) and posterior tibial nerves respectively. Reflex responses were recorded from the ipsilateral tibialis anterior (TA) (flexion reflex) and soleus (H-reflex) muscles. The plantar cutaneous afferents were stimulated at three times the perceptual threshold (200 Hz, 24-ms pulse train) at conditioning-test intervals that ranged from 3 to 90 ms. Sinusoidal movements were imposed to the right hip joint at 0.2 Hz with subjects supine. Control and conditioned reflexes were recorded as the hip moved in flexion and extension. Leg muscle activity and sagittal-plane joint torques were recorded. We found that excitation of plantar cutaneous afferents facilitated the soleus H-reflex and the long latency flexion reflex during hip extension. In contrast, the short latency flexion reflex was depressed by plantar cutaneous stimulation during hip flexion. Oscillatory joint forces were present during the transition phase of the hip movement from flexion to extension when stimuli were delivered during hip flexion. Hip-mediated input interacts with feedback from the foot sole to facilitate extensor and flexor reflex activity during the extension phase of movement. The interactive effects of these sensory signals may be a feature of impaired gait, but when they are appropriately excited, they may contribute to locomotion recovery in these patients.     

Effects of muscle relaxant E-0646 on human stretch reflex and responses

The experiment examined the effects of a muscle relaxant E-0646 on short latency stretch reflex (SLR) and long latency stretch responses (LLRs) of lower leg muscles in 5 healthy male subjects, aged 24-41 years. An AC servomotor produced an ankle rotation of comfortably seated subjects in the direction to stretch the triceps surae muscle for the SLR and the tibialis anterior muscle for the LLRs. Following control measurements, after the oral administration of 50, 100, 150, and 300 mg of E-0646, the SLR and LLRs were recorded every 2 hours for 4 or 6 hours with simultaneous blood sampling to measure E-0646 plasma levels. Electromyograms (EMGs) of the soleus and tibialis anterior muscles were recorded with surface electrodes on each muscle belly. A precision potentiometer was used to measure the ankle angle. The soleus SLR, evoked by a torque pulse (7 msec duration) of one of 5 different amplitudes, decreased 2 hours after the 300 mg administration. A SLR decrease was also found in a simple relationship between the integrated SLR EMG and the angular velocity at 20 msec after the torque onset, in which the slope and intercept of the relationship changed. The plasma E-0646 level roughly correlated with the SLR decrease. No significant change of the SLR was observed when dosage was less than 150 mg. The LLRs of the tibialis anterior muscle were tested under the instruction "resist when perturbed" during a weak isometric contraction against a constant plantar flexion torque.(ABSTRACT TRUNCATED AT 250 WORDS)     

Cortical and segmental excitability during fatiguing contractions of the soleus muscle in humans

ObjectiveThe aim of this study was to examine the cortical and segmental excitability changes during fatigue of the soleus muscle.MethodsTen healthy young subjects performed 45 plantar flexion maximal voluntary contractions (MVCs) (7-s on/3-s off) in 9 epochs of five contractions. Motor evoked potentials (MEPs) using transcranial magnetic stimulation and H-reflexes were assessed during the task.ResultsThe torque and the soleus EMG activity both showed the greatest decline during the 1st epoch, followed by a gradual, but significant decrease by the end of the task (～70% pre-fatigue). The H-reflex sampled at rest after each epoch decreased to 66.6 ± 18.3% pre-fatigue after the first epoch, and then showed no further change. The MEP on 10% pre-fatigue MVC after each epoch increased progressively (252.9 ± 124.2% pre-fatigue). There was no change in the MEPs on the 3rd MVC in each epoch. The silent period on the MVC increased (109.0 ± 9.2% pre-fatigue) early with no further changes during the task.ConclusionsThese findings support that the motor cortex increases excitability during fatigue, but with a concomitant inhibition.SignificanceThese findings are in contrast to upper extremity muscles and may reflect a distinct response specific to postural, fatigue-resistant muscle.     

The effect of posture on the normal and pathological auditory startle reflex.

The effect of posture on the EMG pattern of the normal auditory startle reflex was investigated. The startle response to an unexpected auditory tone was studied in eleven normal subjects when standing, and in six normal subjects when sitting relaxed or tonically plantar flexing both feet. Reflex EMG activity was recorded in the tibialis anterior and soleus about twice as frequently when standing, than when sitting relaxed. In addition, the median latencies to onset of reflex EMG activity in the tibialis anterior and soleus were about 40 and 60 ms shorter during standing, than when sitting relaxed. No short latency EMG activity was recorded in the calf muscles during tonic plantar flexion of the feet, while sitting. The effect of posture on the EMG pattern of the pathological auditory startle reflex was studied in five patients with hyperekplexia. In three patients the latency to onset of reflex EMG activity in the tibialis anterior was shorter when standing, than when sitting relaxed. The EMG pattern of the reflex response to sound was studied in detail in two of these patients and consisted of up to three successive components. The expression of each EMG component depended on the postural set of the limbs. In particular, a distinct short latency component was found in posturally important muscles following auditory stimulation. This short latency component was not recorded when sitting relaxed. It is concluded that the EMG pattern of the physiological and pathological auditory startle response is not fixed, but may change with the postural stance of the body. This finding supports the theory that the normal startle reflex and the abnormal startle reflex in hyperekplexia have a common brainstem origin.     

Joint dependent passive stiffness in paretic and contralateral limbs of spastic patients with hemiparetic stroke.

Torque-angle relations at the elbow and ankle joints of relaxed normal controls and patients with hemiparetic stroke were compared. Low velocity flexion/hold/extension angular perturbations were applied to the joint under examination. The resulting torque-angle profiles described a hysteresis loop with similar slopes during the extension and flexion stages but separated by a vertical torque offset. Torque-angle responses obtained in the absence of significant muscle activation, as recorded by surface electromyographic activity, were designated as passive. Elbow passive stiffness estimates were calculated from the slope of the torque-angle response during the flexion stage of the perturbation. The elbow torque-angle plots exhibited linear passive stiffness with magnitude significantly lower than the passive stiffness of the ankle in both normal subjects and spastic patients. Changing ramp velocity had no significant effect on the passive torque-angle hysteresis loop at the elbow. A comparison of the torque-angle relations between hemiparetic spastic and normal control arms showed no significant differences in passive stiffness. Furthermore, no significant differences were found between paretic and contralateral upper limbs of a given hemiparetic subject. By contrast, significant differences in the torque-angle hysteresis loop were present between the paretic and contralateral ankles in all hemiparetic patients tested. These differences were more significant during dorsiflexion, and therefore seem to be related to preferential changes in mechanical properties of plantar flexor muscles. It is hypothesised that the differences in the torque-angle hysteresis loop between elbow and angle joints are related primarily to the larger amount of connective tissue in the calf muscles, as well as to a larger total physiological cross sectional area of calf muscles compared with elbow muscles. It is further hypothesized that the preferential increases in passive stiffness at the ankle in spastic legs result from immobilisation induced changes in muscle connective tissue, which are most prominent in muscles with predominantly slow-twitch fibres (such as soleus). Connective tissue surrounding such slow twitch muscle fibres have been shown to be more sensitive to immobilisation than those in fast twitch muscle. The functional, pathophysiological, and clinical implications of our findings are reviewed.