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Effect of thumb anaesthesia on weight perception, muscle activity and the stretch reflex in man.

1. We have confirmed the results of Gandevia & McCloskey (1977) on the effect of thumb anaesthesia on perception of weights lifted by the thumb. Weights lifted by flexion feel heavier and weights lifted by extension feel lighter. 2. The change in size of the long-latency stretch reflex in flexor pollicis longus or extensor pollicis longus after thumb anaesthesia cannot explain the effect on weight perception by removal or augmentation of the background servo assistance to muscular contraction. 3. During smooth thumb flexion, thumb anaesthesia increases e.m.g. activity in flexor pollicis longus and extensor pollicis longus for any given opposing torque. 4. During smooth thumb extension the opposite occurs: e.m.g. activity in both extensor and flexor pollicis longus decreases. 5. Clamping the thumb at the proximal phalanx to limit movement solely to the interphalangeal joint reduces or abolishes the effect of anaesthesia on both weight perception and e.m.g. activity during both flexion or extension tasks. 6. Gandevia & McCloskey's findings on the distorting effects of thumb anaesthesia on weight perception cannot be used to support the hypothesis of an efferent monitoring system of the sense of effort. Our results emphasize the close functional relationship between cutaneous and joint afferent information and motor control.

Anesthesia, Local↗

Stretch reflexes of the proximal arm in a patient with mirror movements: absence of bilateral long-latency components.

The stretch reflex responses evoked by unilateral limb displacement in distal (first dorsal interosseus (FDI)) and in proximal (biceps brachii (Bb)) arm muscles were studied during matched bilateral contractions in a patient with congenital mirror movements. In this patient unilateral transcortical magnetic stimulation (TMS) elicited not only the normal contralateral EMG response but also a clear ipsilateral component in the EMG of both proximal and distal arm muscles. As expected from previous studies, the ipsilateral FDI muscle responded to stretch of the index finger with short- (M1) and long-latency (M2) reflex components. In addition, the FDI contralateral to displacement exhibited an abnormal mirrored response corresponding to the M2 interval. In contrast, whereas the ipsilateral Bb responded to imposed elbow extension with a marked M1/M2 reflex response, no mirroring of either reflex component was apparent in the contralateral Bb EMG. If the mirroring of the M2 in the FDI is accepted as evidence for the transcortical nature of the M2 reflex response, then it follows that the absence of such mirroring in the Bb indicates that a transcortical mechanism cannot play a major role in the generation of long-latency stretch reflex responses in proximal arm muscles.

Adult↗

Tonic stretch reflexes in older able-bodied people.

It is a general assumption that, in able-bodied persons, tonic stretch reflex (TSR) activity is not elicited during stretching of relaxed muscles and that the presence of TSR activity following brain damage is, therefore, indicative of spasticity. However, a variety of studies have reported age-related changes in reflex activity, raising the question of whether this assumption is justified in older subjects. The aim of this study was to determine if TSRs were activated in the relaxed elbow flexors of able-bodied people in an age-group at risk of stroke. Electromyographic (EMG) activity was recorded in 30 able-bodied subjects aged 46 to 78 years when their relaxed elbow flexors were subjected to ramp and sinusoidal stretches of different amplitudes and velocities. It was found that these subjects did not exhibit TSR activity under these conditions. Therefore, the practice of measuring TSR activity as a means of quantifying spasticity in stroke patients appears justified.

Aged↗

Behavior of the stretch reflex in a multi-jointed limb.

When an external perturbation acts on a segment of a multi-jointed limb, angular motion results in all limb segments which are inertially linked to the one which is perturbed. In such a situation, different hypotheses on the functional role of the stretch reflex lead to different predictions on reflex responses by the stretched muscles. These predictions were examined experimentally and it was found that the EMG responses of elbow muscles evoked by forces applied to either the upper arm or the forearm were not uniquely related to changes in elbow angular position. They were instead correlated with the direction of changes in the net elbow torque resulting from the perturbation.

Arm↗

Influence of posture and muscle length on stretch reflex activity in poststroke patients with spasticity.

OBJECTIVE: To investigate the influence of different positions on stretch reflex activity of knee flexors and extensors measured by electromyography in poststroke patients with spasticity and its expression in the Ashworth Scale. DESIGN: Crossover trial with randomized order of positioning. SETTING: Outpatient rehabilitation center in the Netherlands. PARTICIPANTS: Poststroke patients (N = 19) with lower-limb spasticity. INTERVENTION: Changing position: sitting versus supine. MAIN OUTCOME MEASURES: Root mean square (RMS) values of muscle activity and goniometric parameters, obtained during the pendulum test and passive knee flexion and extension, and Ashworth scores. RESULTS: RMS values of bursts of rectus femoris activity were significantly higher in the supine compared with the sitting position (P = .006). The first burst of vastus lateralis activity during the pendulum test (P = .049) and semitendinous activity during passive stretch (P = .017) were both significantly higher in the supine versus the sitting position. For both the pendulum test and passive movement test, the duration and amplitude of the cyclic movement of the lower leg changed significantly as well. In the supine position, we found significantly higher Ashworth scores for the extensors (P = .001) and lower scores for the flexors (P = .002). CONCLUSIONS: The outcome of clinical and neurophysiologic assessment of spasticity is influenced considerably by subject positioning.

Cross-Over Studies↗

Nonlinear properties of stretch reflex studied in the decerebrate cat.

1. Pairs of brief stretches or a series of stretches at random intervals (Poisson process) were applied to a slow (soleus) and a fast (plantaris) muscle in decerebrate cats to analyze the nonlinear effects of one stretch on the reflex responses to subsequent stretches. Neural activity, electromyogram (EMG), and force were recorded. The reflex responses due to stretch were compared with reflexes as a result of electrical stimulation of nerves. Nonlinearities of muscle were also examined in the absence of reflexes. Short-latency neural activity produced by the stimuli at all intervals studied was quite constant, so changes in sensory activity cannot account for the nonlinearities. Three phases of nonlinear interactions were observed, and mechanisms for these nonlinearities are suggested. 2. For short intervals (less than 100 ms) following a stretch the force and EMG produced by a second stretch is depressed. This early depression could be due to the after hyperpolarization of the motoneuron cel body or to synaptic mechanisms, since the depression of EMG is seen with electrical stimulation of Ia sensory, but not alpha-motor axons. In addition, a second stretch can disrupt the reflex contraction produced by the first stretch if it occurs at a time when new actomyosin bonds are not readily formed. Because of this force suppression, the total reflex force produced in response to two stretches may be less than the response to a single stretch. 3. For intervals between 100 and 300 ms the force and EMG produced by a second stretch is enhanced. This potentiation is also seen with electrical stimulation of large sensory but not motor axons and could result from a synchronization of motoneuronal excitability cycles. It is more prominent in the homogeneous (soleus) muscle than the mixed (plantaris) muscle, probably because the motoneuron cell bodies will reach a period of high excitability at more nearly the same time in the homogeneous muscle. 4. For longer intervals the force produced by a second stretch is reduced even when the EMG is close to control values. This late depression is also observed with electrical stimulation of cut motor axons and therefore arises from the contractile properties of muscles. 5. With a random series of stretches, the same time course of nonlinear interactions is observed. However, as the mean rate of the random stretches is increased, the average response of the reflex decreases. Thus, the stretch reflex will be most effective in correcting for occasional perturbations to a movement, rather than for continuously varying disturbances.

Animals↗

Relationship between the degree of inhibited stretch reflex activities of the wrist flexor and reaction time during quick extension movements.

It has been reported that stretch reflex responses, including the long latency component, are modulated by motor preparation for the direction and type of movement. In the present study, human subjects were required to make a reaction movement in the direction of the wrist extension following a muscle stretch to the wrist flexor, and we investigated the relationship between the modulation of reflex activities of the wrist flexor and the length of reaction time (premotor time) of the wrist extensor. Twenty-five healthy males, ranging in age from 20 to 28, participated in the experiments. A DC torque motor was used to evoke the reflex EMG responses on the flexor. Averaging the rectified EMG, recorded with the surface electrodes over the flexor, showed short and long latency reflexes (M1 and M2 components) in response to the muscle stretch. For all subjects, the amplitudes of the reflex components during the extension reaction movement decreased, compared to those amplitudes in the non-reaction tasks. The decrease in the M2 component, which is considered a transcortical reflex, was significantly larger than the decrease in the M1 component, which is a spinal reflex. Moreover, there were correlations between reaction time to muscle stretch and the degree of decrease in reflex activities with the extension reaction (r = 0.652 for M1, r = 0.813 for M2, P < 0.01). It became clear that the subjects with shorter reaction times inhibited their reflex activities of the flexor, particularly the M2 component which prevents the extension movement, to a greater degree than the subjects with longer reaction times. Therefore, our results suggest that the degree of M2 modulation directly reflects the individual motor control required to perform quick movements.

Adult↗

Observations on stretch reflexes in lumbar back muscles of the cat.

Reflex responses to brief muscle stretch and to electrical stimulation of dorsal roots were studied in cat longissimus and iliocostalis muscles by myographic and electromyographic recordings in spinal and anesthetized preparations. Brief stretch applied simultaneously to both muscles, by pulling at an isolated segment of the iliac bone, elicited contractions only in the central region of longissimus, composed of slowly contracting fibers, the time to peak tension varying between 70 and 100 ms. No reflex responses were observed in the faster contracting parts of this muscle or in the iliocostalis under the stimulation conditions used. The total reflex time usually varied from 4.5 to 7 ms which is shown to correspond to intra-spinal conduction times, approximately between 2 and 5 ms. Only exceptionally was a central reflex delay corresponding to a monosynaptic transmission observed. Reflexes evoked by dorsal root simulation (L4, L5) have a central conduction time similar to those elicited by adequate stimulation. The possibility that the central pathway for stretch reflexes in longissimus may involve more than two neurons is considered. In decerebrate preparations the central region of longissimus displays a prominent tonic stretch reflex. With the longissimus in situ the magnitude of the reflex is highly dependent on the position of the lumbar spine relative to the pelvic girdle.

Anesthesia↗

Interaction between voluntary contraction and tonic stretch reflex transmission in normal and spastic patients.

Transmission characteristics of tonic stretch reflex (TSR) pathways in both normal and spastic patients have been measured during different levels of sustained voluntary contraction of the biceps brachii muscle. A cross-correlation technique of analysis was used to separate reflex responses from the total electromyographic activity. TSR transmission in normal subjects was observed to change with the level of voluntary contraction; sensitivity to stretch increased approximately three-fold as the subject stiffened the arm by simultaneously contracting flexor and extensor muscles. In contrast, TSR transmission did not alter during voluntary contraction in spastic patients. It is proposed that in spastic patients supraspinal modulation of reflex transmission is impaired because hypersensitive spinal reflexes short-circuit long loop pathways.

Cerebral Palsy↗

Stretch reflex of quadriceps femoris in normal man: methodological considerations and normative data.

Short- and long-latency responses to stretching have been evoked in various muscles in the healthy subject. The quadriceps femoris, however, has never been investigated in this connection, despite its functional importance. We studied its stretch reflex in 17 healthy subjects with a torque motor to determine its morphological features in relation to the level of background EMG activity, the stretching velocity and duration and the instructions given to the subject. Two bursts of EMG activity were constantly evoked when the muscle was voluntarily activated. Response magnitude was proportional to the background activity and displacement velocity. Short stretching evoked only one EMG burst. These findings are similar to those from other muscles. Intra-individual variability was evaluated repeating the experiment in the same session without changing position of the electrodes, and then after two months. The long-latency response showed a good intra-individual reproducibility.

Adult↗

Modulation of the mandibular stretch reflex sensitivity during various phases of rhythmic open-close movements in humans.

The muscle spindles of the jaw elevator muscles provide positive feedback to the alpha motoneurons. It is generally assumed that the feedback is modulated during chewing so that counterproductive forces of the jaw elevator muscles can be avoided during jaw opening. Our aim was to investigate the modulation of the muscle spindle input to the alpha motoneurons during various phases of open-close movements in man. To that end, subjects made rhythmic open-close movements at their natural chewing frequency. A force impulse (5 N, 10 ms), eliciting a jaw-jerk reflex, was unexpectedly applied. The impulse was applied to the mandible at 8 different phases during an open-close cycle, but only 1 impulse per cycle. Jaw movement and surface EMG of the masseter and temporal muscles on both sides were recorded during 3 cycles without an impulse and 3 succeeding cycles with an impulse. To examine whether the modulation of the mandibular stretch reflex sensitivity depends on the food resistance, we applied an additional external force on the mandible, counteracting closing of the jaw each cycle. Two experimental sessions were performed in random order, i.e., without force and with an additional force of 20 N. We observed pronounced reflexes at the onset of jaw closing, during the closing phase, and at occlusion. No or only weak jaw-jerk reflexes were present during jaw opening. The reflex amplitudes at occlusion were larger when an external force was present. This increase in reflex amplitude may be the result of an adjusted gamma motoneuron activity, from pre-motor inhibition, or from both. The reflex amplitudes elicited during jaw closing were not correlated with the phase of the movement.

Adult↗

Reducing human biceps brachii spinal stretch reflex magnitude.

1. Nine subjects received 6 baseline and 24 training sessions, each consisting of 250 elbow joint stretch perturbations into extension. The training sessions were designed to reduce the subjects' biceps brachii spinal stretch reflexes (SSR). Changes in longer-latency responses and short-latency brachioradialis responses were also monitored. Background electromyogram activity was recorded from the lateral head of triceps brachii during the biceps sampling intervals. These data were compared with those form 12 control subjects who received equal numbers of stretches over an extended baseline (i.e., without application of a training paradigm). 2. Training subjects reduced their mean biceps SSR responses by 24%, whereas control subjects increased their responses by 12% When changes in activity were grouped by sets of six consecutive sessions following the baseline interval, group-by-time interaction was observed. Training subjects showed significant reductions from baseline after the first raining set. The increased magnitude of biceps SSR for control subjects was significantly larger than baseline in sets 2-5. After the first training interval, all subsequent differences between groups were statistically significant. 3. The brachioradialis showed greater response variability, but these responses paralleled those seen in biceps brachii. The training group reduced their brachioradialis responses by 18%, whereas the control group increased their responses by 12%. Background activity recorded over the lateral head of triceps brachii during the biceps brachii SSR window became smaller in both groups. 4. Comparison of data between control subjects and subjects who had undergone biceps SSR up-training in previous studies suggests that the small increase seen among the present control subjects was probably not the result of a direct training effect. 5. Baseline differences in longer-latency response onset time were seen between groups. Over extended baseline sessions, longer-latency responses showed only a 1% onset, representing a change of 0.3 ms, in the control group. A 1.3% later onset, equaling a change of 1.1 ms, was observed over conditioning sessions in the training group. When analyzed by sets following baseline, neither group showed significant within- or between-group changes over time. 6. The magnitude of the longer-latency biceps brachii response showed 7% and 37% reductions for the control and training groups, respectively. No difference in set averages was seen within groups, but a significant difference was measured over sets 2-4 between groups. Reductions in the magnitude of longer-latency responses in training subjects were more obvious in later training sessions. 7. These data suggest that 1) the biceps brachii SSR can be reduced in human subjects; 2) concurrent changes are often observed in the synergist brachioradialis; and 3) the reduced biceps SSR magnitude is not linked to a compensatory increase in antagonist muscle activity. Reductions in the magnitude of longer-latency biceps brachii activity seen in parallel with the decreased SSR may imply that some degree of supraspinal processing is required to achieve this task.

Adolescent↗

Modulation of the stretch reflex of jaw-closing muscles in different modes and phases of respiration.

The objective of this study was to investigate whether and how changes in the mode of respiration affect the electromyographic activity of human jaw-closing muscles. Fifteen men were examined in this study. A pair of surface electrodes was attached bilaterally to the masseter and anterior and posterior temporalis muscles for electromyographic recording. Respiratory movements of the chest wall and nasal airflow were recorded simultaneously. Recordings were performed with subjects in the sitting position during quiet nasal and oral respiration. The stretch reflex of jaw-closing muscles was elicited by randomly tapping the chin with an impulse hammer. In 11 subjects, we measured nasal resistance with a rhinomanometer. The amplitude of electromyographic activities of the masseter and anterior temporalis muscles during oral respiration was significantly less than that during nasal respiration, whereas that of the posterior temporalis muscle showed no significant difference between the different modes of respiration. Furthermore, the reduction in the amplitude of the electromyographic activity was more evident in the inspiratory phase during oral respiration. There was a significant positive correlation between the ratio of the reflex amplitude during inspiration in the 2 respiratory modes and nasal resistance for the masseter muscle, but not for the anterior temporalis muscle. These results suggest that the reflexive electromyographic activity of some human jaw-closing muscles is modulated during oral respiration.

Adult↗

Long-latency stretch reflexes of two intrinsic muscles of the human hand analysed by cooling the arm.

1. The arm was cooled to examine the effect of slowing nervous conduction on the latency of various reflex responses of the abductor digiti minimi (ADM) and the first dorsal interosseus (FDI) muscles. A reflex dependent upon slow afferents acting with a short central delay should be slowed by more than one of comparable initial latency, but dependent upon fast afferents acting with a long central delay. The F wave seen in the EMG on stimulating the ulnar nerve was used to assess the effect of cooling in slowing fast fibres. 2. Cooling showed that the prominent long-latency excitation evoked on stretching one or other muscle by displacing the appropriate finger is medicated by fast afferent fibres. 3. Both fast cutaneous and group I muscles afferents were found to be capable of contributing to the long-latency responses. The muscle afferent contribution was consistently dominant for slow 'stretches', while a cutaneous afferent contribution was sometimes recognizable for finger taps. 4. The distinction proved possible because the cutaneous afferents from the relevant finger run in the median nerve for FDI and the ulnar nerve for ADM. The cooling produced much greater slowing of conduction for the ulnar nerve. 5. The prior suggestion that spindle group II afferents are responsible for the major part of the long-latency component of the human stretch reflex thus fails to stand; however, some contribution has not been finally excluded.

Cold Temperature↗

Different mechanisms underlie the long-latency stretch reflex response of active human muscle at different joints.

1. Stretch of voluntarily activated human muscle results in a reflex response consisting of short-latency (M1) and delayed long-latency (M2) components. The mechanism of the M2 response remains the subject of controversy. The present study tested the universality of the hypothesis that the M2 response results from the transmission of low-threshold muscle afferent input travelling over a long-loop supraspinal pathway. Muscle reflex responses resulting from imposed stretch were obtained from the first dorsal interosseus (FDI), biceps brachii (BB), triceps brachii (TB) and triceps surae (TS) muscles. 2. Patients suffering from Huntington's disease (HD) show a selective loss of FDI-M2 responses, with sparing of the M1. This has been attributed to disruption of supraspinal pathways as a part of the disease pathology. Accordingly, HD has been used in the present study as a model to test the universality of the long-loop hypothesis: if this is so, then HD patients with an absent FDI M2 should also fail to show an M2 response in other muscles. 3. It is shown that a group of HD patients in whom the FDI-M2 response was absent or residual developed clear M2 responses in the TB, BB and TS muscles following stretch sufficient to invariably evoke this component in normal subjects. 4. It is thus concluded that longer-latency stretch reflex components are not invariably mediated over long-loop supraspinal pathways, but that this mode of control is dominant only in muscles, such as those of the hand, whose function depends largely on direct cortical control.

Adult↗

Gating of the afferent volley of the monosynaptic stretch reflex during movement in man.

1. The time course of the changes in presynaptic inhibition of Ia fibres to soleus motoneurones has been investigated during a voluntary ramp-and-hold plantar flexion. 2. Monosynaptic Ia facilitation of the soleus H reflex was evoked by stimulation of the inferior soleus (homonymous) and femoral (heteronymous) nerves. Changes in presynaptic inhibition of the Ia fibres mediating the conditioning volleys were inferred from changes in the amount of reflex facilitation evoked by such constant conditioning stimulations. 3. At the beginning of the voluntary contraction, both homonymous and heteronymous Ia facilitations were markedly increased with respect to their rest values. Then, they dropped abruptly in the middle of the ramp, whatever the ramp duration. 4. The stronger the contraction at the end of the ramp, the larger the initial increase in reflex facilitation. 5. These results indicate that presynaptic inhibition of Ia terminals on motoneurones of the contracting muscle is decreased during the first half of the ramp and then returns to its rest level. It is argued that these changes in presynaptic inhibition are centrally programmed, and it is suggested that, under certain conditions, presynaptic inhibition can be modulated according to both the strength and timing of the contraction. 6. The functional role of the resulting changes in the gain of the stretch reflex is discussed.

Adult↗

Firing patterns of human flexor carpi radialis motor units during the stretch reflex.

Single motor unit and gross surface electromyographic responses to torque motor-produced wrist extensions were studied in human flexor carpi radialis muscle. Surface EMG typically showed two "periods" of reflex activity, at a short and long latency following stretch, but both periods occurring before a subject's voluntary reaction to the stretch. The amplitude of EMG activity in both reflex periods increased monotonically with an increase in the torque load. The amplitude of the short-latency reflex response was very dependent on the motoneuron pool excitability, or preload. The amplitude of the long-latency reflex response also varied with the preload, but could, in addition, be modulated by the subject's preparatory set for a voluntary response to the imposed displacement. When a single motor unit that was not tonically active began to fire during the stretch reflex, it did so primarily during the long-latency period. When caused to fire repetitively by voluntary facilitation of the motoneuron pool, that same unit now showed activity during both periods of the stretch reflex. Further increases in either motoneuron pool facilitation or in perturbation strength resulted in a monotonic increase in response probability of a single motor unit during the short-latency period. However, the response probability of a single unit during the long-latency reflex period did not always vary in a monotonic way with increases in either torque load or motoneuron pool facilitation. For an additional series of experiments, the subject was instructed on how to respond voluntarily to the upcoming wrist perturbation. The three instructions to the subject had no effect on the response probability of a single motor unit during either the background or short-latency periods of the stretch reflex. However, prior instruction clearly affected a unit's response probability during the long-latency reflex period. Changes in the firing rate of motor units, and in the recruitment or derecruitment of nontonic units, contributed to this modulation of reflex activity during the long-latency period.

Adult↗

Separable least squares identification of nonlinear Hammerstein models: application to stretch reflex dynamics.

The Hammerstein cascade, consisting of a zero-memory nonlinearity followed by a linear filter, is often used to model nonlinear biological systems. This structure can represent some high-order nonlinear systems accurately with relatively few parameters. However, it is not possible, in general, to estimate the parameters of a Hammerstein cascade in closed form. The most effective method available to date uses an iterative approach, which alternates between estimating the linear element from a crosscorrelation, and then fitting a polynomial to the nonlinearity via linear regression. This paper proposes the use of separable least squares optimization methods to estimate the linear and nonlinear elements simultaneously in a least squares framework. A separable least squares algorithm for the identification of Hammerstein cascades is developed and used to analyze stretch reflex electromyogram data from two experimental subjects. The results show that in each case the proposed algorithm produced a better model, in that it predicted the system's response to novel inputs more accurately, than did models estimated using the traditional iterative algorithm. Monte-Carlo simulations demonstrated that when the input is a non-Gaussian, nonwhite signal, as is often the case experimentally, the traditional iterative identification approach produces biased models, whereas the separable least squares approach proposed in this paper does not.

Algorithms↗