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Key points Sudden unloading of preloaded wrist muscles elicits motion to a new wrist position. Such motion is prevented if subjects unload muscles using the contralateral arm (self-unloading)., Corticospinal influences originated from the primary motor cortex maintain tonic influences on motoneurons of wrist muscles before sudden unloading but modify these influences prior to the onset and until the end of self-unloading., Results are interpreted based on the previous finding that intentional actions are caused by central, particularly corticospinal, shifts in the spatial thresholds at which wrist motoneurons are activated, thus predetermining the attractor point at which the neuromuscular periphery achieves mechanical balance with environment forces., By maintaining or shifting the thresholds, descending systems let body segments go to the equilibrium position in the respective unloading tasks without the pre-programming of kinematics or muscle activation patterns., The study advances the understanding of how motor actions in general, and anticipation in particular, are controlled., Abstract The role of corticospinal (CS) pathways in anticipatory motor actions was evaluated using transcranial magnetic stimulation (TMS) of the primary motor cortex projecting to motoneurons (MNs) of wrist muscles. Preloaded wrist flexors were suddenly unloaded by the experimenter or by the subject using the other hand (self-unloading). After sudden unloading, the wrist joint involuntarily flexed to a new position. In contrast, during self-unloading the wrist remained almost motionless, implying that an anticipatory postural adjustment occurred. In the self-unloading task, anticipation was manifested by a decrease in descending facilitation of pre-activated flexor MNs starting ∼72 ms before changes in the background EMG activity. Descending facilitation of extensor MNs began to increase ∼61 ms later. Conversely, these influences remained unchanged before sudden unloading, implying the absence of anticipation. We also tested TMS responses during EMG silent periods produced by brief muscle shortening, transiently resulting in similar EMG levels before the onset and after the end of self-unloading. We found reduced descending facilitation of flexor MNs after self-unloading. To explain why the wrist excursion was minimized in self-unloading due to these changes in descending influences, we relied on previous demonstrations that descending systems pre-set the threshold positions of body segments at which muscles begin to be activated, thus predetermining the equilibrium point to which the system is attracted. Based on this notion, a more consistent explanation of the kinematic, EMG and descending patterns in the two types of unloading is proposed compared to the alternative notion of direct pre-programming of kinematic and/or EMG patterns. [ABSTRACT FROM AUTHOR]

Key Points: The fluctuations in force during a steady isometric contraction (force steadiness) are associated with oscillations in common synaptic input to the involved motor neurons. Decreases in force steadiness are associated with increases in pegboard times in older adults, although a mechanism for this link has not been established. We used a state-space model to estimate the variability in common synaptic input to motor neurons during steady, isometric contractions. The estimate of common synaptic input was derived from the discharge times of motor units as recorded with high-density surface electrodes. We found that the variability in common synaptic input to motor neurons modulates force steadiness for young and older adults, as well as pegboard time for older adults., Abstract: We investigated the associations between grooved pegboard times, force steadiness (coefficient of variation for force) and variability in an estimate of the common synaptic input to motor neurons innervating the wrist extensor muscles during steady contractions performed by young and older adults. The discharge times of motor units were derived from recordings obtained with high-density surface electrodes when participants performed steady isometric contractions at 10% and 20% of maximal voluntary contraction force. The steady contractions were performed with a pinch grip and wrist extension, both independently (single action) and concurrently (double action). The variance in common synaptic input to motor neurons was estimated with a state-space model of the latent common input dynamics. There was a statistically significant association between the coefficient of variation for force during the steady contractions and the estimated variance in common synaptic input in young (r 2  = 0.31) and older (r 2  = 0.39) adults, although not between either the mean or the coefficient of variation for interspike interval of single motor units with the coefficient of variation for force. Moreover, the estimated variance in common synaptic input during the double-action task with the wrist extensors at the 20% target was significantly associated with grooved pegboard time (r 2  = 0.47) for older adults but not young adults. These findings indicate that longer pegboard times of older adults were associated with worse force steadiness and greater fluctuations in the estimated common synaptic input to motor neurons during steady contractions., (© 2018 The Authors. The Journal of Physiology © 2018 The Physiological Society.)

Key Points: Activity in the primary motor cortices of both hemispheres increases during unilateral movement preparation, but the functional role of ipsilateral motor cortex activity is unknown. Ipsilateral motor cortical activity could represent subliminal 'motor planning' for the passive limb. Alternatively, it could represent the state of the active limb, to support coordination between the limbs should a bimanual movement be required. Here we assessed how preparation of forces toward different directions, with the left wrist, alters evoked responses to transcranial magnetic stimulation of left motor cortex. Preparation of a unilateral movement caused excitability increases in ipsilateral motor cortex that reflected forces produced with the active limb in an intrinsic (body-centred), rather than an extrinsic (world-centred), coordinate system. These results suggest that ipsilateral motor cortical activity prior to unilateral action reflects the state of the active limb, rather than subliminal motor planning for the passive limb., Abstract: Corticospinal excitability is modulated for muscles on both sides of the body during unilateral movement preparation. For the effector, there is a progressive increase in excitability, and a shift in direction of muscle twitches evoked by transcranial magnetic stimulation (TMS) toward the impending movement. By contrast, the directional characteristics of excitability changes in the opposite (passive) limb have not been fully characterized. Here we assessed how preparation of voluntary forces towards four spatially distinct visual targets with the left wrist alters muscle twitches and motor-evoked potentials (MEPs) elicited by TMS of left motor cortex. MEPs were facilitated significantly more in muscles homologous to agonist rather than antagonist muscles in the active limb, from 120 ms prior to voluntary EMG onset. Thus, unilateral motor preparation has a directionally specific influence on pathways projecting to the opposite limb that corresponds to the active muscles rather than the direction of movement in space. The directions of TMS-evoked twitches also deviated toward the impending force direction of the active limb, according to muscle-based coordinates, following the onset of voluntary EMG. The data indicate that preparation of a unilateral movement increases task-dependent excitability in ipsilateral motor cortex, or its downstream projections, that reflects the forces applied by the active limb in an intrinsic (body-centred), rather than an extrinsic (world-centred), coordinate system. The results suggest that ipsilateral motor cortical activity prior to unilateral action reflects the state of the active limb, rather than subliminal motor planning for the passive limb., (© 2018 The Authors. The Journal of Physiology © 2018 The Physiological Society.)

Previous studies have shown that preplanned movements can be rapidly released when a startling acoustic stimulus (SAS) is presented immediately prior to, or coincident with, the imperative signal to initiate movement. Based on the short latency of the onset of muscle activity (typically in less than 90 ms) and the frequent co-expression of startle responses in the neck and eye muscles, it has been proposed that the release of planned movements by a SAS is mediated by subcortical, possibly brainstem, pathways. However, a role for cortical structures in mediating these responses cannot be ruled out based on timing arguments alone. We examined the role of the cortex in the mediation of these responses by testing if a suprathreshold transcranial magnetic stimulation applied over the primary motor cortex, which suppresses voluntary drive and is known to delay movement initiation, could delay the release of movement by a SAS. Eight subjects performed an instructed-delay task requiring them to make a ballistic wrist movement to a target in response to an acoustic tone (control task condition). In a subset of trials subjects received one of the following: (1) suprathreshold TMS over the contralateral primary motor cortex 70 ms prior to their mean response time on control trials (TMS(CT)), (2) SAS 200 ms prior to the go cue (SAS), (3) suprathreshold TMS 70 ms prior to the mean SAS-evoked response time (TMS(SAS)), or (4) TMS(SAS) and SAS presented concurrently (TMS+SAS). Movement kinematics and EMG from the wrist extensors and flexors and sternocleidomastoid muscles were recorded. The application of TMS(CT) prior to control voluntary movements produced a significant delay in movement onset times (P < 0.001) (average delay = 37.7 ± 12.8 ms). The presentation of a SAS alone at -200 ms resulted in the release of the planned movement an average of 71.7 ± 2.7 ms after the startling stimulus. The early release of movement by a SAS was significantly delayed (P < 0.001, average delay = 35.0 ± 12.9 ms) when TMS(SAS) and SAS were presented concurrently. This delay could not be explained by a prolonged suppression of motor unit activity at the spinal level. These findings provide evidence that the release of targeted ballistic wrist movements by SAS is mediated, in part, by a fast conducting transcortical pathway via the primary motor cortex.

Motor skill acquisition in the lower limb may induce modifications of spinal network excitability. We hypothesized that short-term motor adaptation in precision grip tasks would also induce modifications of cervical spinal network excitability. In a first series of experiments, we studied the impact of two different precision grip force control tasks (a visuomotor force-tracking task and a control force task without visual feedback) on cervical spinal network excitability in healthy subjects. We separately tested the efficacy of two key components of the spinal circuitry: (i) presynaptic inhibition on flexor carpi radialis (FCR) Ia terminals, and (ii) disynaptic inhibition directed from extensor carpi radialis (ECR) to FCR. We found that disynaptic inhibition decreased temporarily after both force control tasks, independently of the presence of visual feedback. In contrast, the amount of presynaptic inhibition on FCR Ia terminals decreased only after the visuomotor force tracking task. This temporary decrease was correlated with improved tracking accuracy during the task (i.e. short-term motor adaptation). A second series of experiments confirmed these results and showed that the visuomotor force-tracking task resulted also in an increase of the Hmax/Mmax ratio and the slope of the ascending part of the H-reflex recruitment curve. In order to address the role of presynaptic inhibition in the motor adaptation process, we conducted a third series of experiments during which presynaptic inhibition was recorded before and after two consecutive sessions of visuomotor force tracking. The results showed that (i) improved tracking accuracy occurred during both sessions, and (ii) presynaptic inhibition decreased only after the first session of visuomotor force tracking. Taken together, these results suggest thus that the nature of the motor task performed has a specific impact on the excitability of these cervical spinal circuits. These findings also suggest that early motor adaptation is associated with a modulation of presynaptic Ia inhibition in the upper limb.

It has previously been established that muscles become active in response to deviations from a threshold (referent) position of the body or its segments, and that intentional motor actions result from central shifts in the referent position. We tested the hypothesis that corticospinal pathways are involved in threshold position control during intentional changes in the wrist position in humans. Subjects moved the wrist from an initial extended to a final flexed position (and vice versa). Passive wrist muscle forces were compensated with a torque motor such that wrist muscle activity was equalized at the two positions. It appeared that motoneuronal excitability tested by brief muscle stretches was also similar at these positions. Responses to mechanical perturbations before and after movement showed that the wrist threshold position was reset when voluntary changes in the joint angle were made. Although the excitability of motoneurons was similar at the two positions, the same transcranial magnetic stimulus (TMS) elicited a wrist extensor jerk in the extension position and a flexor jerk in the flexion position. Extensor motor-evoked potentials (MEPs) elicited by TMS at the wrist extension position were substantially bigger compared to those at the flexion position and vice versa for flexor MEPs. MEPs were substantially reduced when subjects fully relaxed wrist muscles and the wrist was held passively in each position. Results suggest that the corticospinal pathway, possibly with other descending pathways, participates in threshold position control, a process that pre-determines the spatial frame of reference in which the neuromuscular periphery is constrained to work. This control strategy would underlie not only intentional changes in the joint position, but also muscle relaxation. The notion that the motor cortex may control motor actions by shifting spatial frames of reference opens a new avenue in the analysis and understanding of brain function.

In our movements and posture, we always act against a physical load. A key property of any load is its elastic stiffness (K), which describes how the force required to hold it must change with position. Here we examine how load stiffness affects the ability to maintain a stable posture at the wrist. Loads having positive (like a spring) and negative stiffness (like an inverted pendulum) were created by varying the position of weights on multiarm rigid pendulum. Subjects (n = 9) held 15 loads (K = +/- 0.04, +/- 0.01 and 0 N m deg(-1) at mean torques of 0.2, 0.4 and 0.6 N m) still for 60 s. Residual wrist movement (sway) increased with mean torque and increased as stiffness became more negative. Large effects of load stiffness were seen at low frequencies (< 1.5 Hz) but not at higher frequencies that reflect load resonance and reflex activity. Subjects accurately perceived their postural sway while holding the loads but measured psychophysical thresholds showed that load stiffness was not perceived. We conclude that load stiffness, independent of force levels, affects the ability to control a load and that the postural control process relies on perception and volitional tracking rather than more automatic reflex pathways. Despite an awareness of their postural errors, we see no evidence for adaptation of postural control processes to compensate for changes in load properties. This is unlike the adaptation of feedforward control processes that produce targeted volitional movements when load properties are altered. We propose that postural control and movement control are fundamentally different neural processes.

The equilibrium point hypothesis (EPH) was conceived as a means whereby the central nervous system could control limb movements by a relatively simple shift in equilibrium position without the need to explicitly compensate for task dynamics. Many recent studies have questioned this view with results that suggest the formation of an internal dynamics model of the specific task. However, supporters of the EPH have argued that these results are not incompatible with the EPH and that there is no reason to abandon it. In this study, we have tested one of the fundamental predictions of the EPH, namely, equifinality. Subjects learned to perform goal-directed wrist flexion movements while a motor provided assistance in proportion to the instantaneous velocity. It was found that the subjects stopped short of the target on the trials where the magnitude of the assistance was randomly decreased, compared to the preceding control trials (P = 0.003), i.e. equifinality was not achieved. This is contrary to the EPH, which predicts that final position should not be affected by external loads that depend purely on velocity. However, such effects are entirely consistent with predictions based on the formation of an internal dynamics model.

In order to document the effects of recurrent inhibition on the firing times of human alpha-motoneurones during natural motor behaviour, a case study was performed on a deafferented patient. The fact that this subject had completely lost the large-diameter sensory afferents provided us with a unique opportunity of selectively stimulating the motor axons in the nerves. The tonic activity of single motor units (n = 21) was recorded in the extensor carpi radialis muscles while applying randomly timed antidromic electrical stimuli to the radial nerve. The peristimulus time histogram analysis showed the presence of biphasic inhibitory effects, including an early, short-lasting component followed by a longer-lasting component occurring 20-40 ms later. The interspike interval (ISI) during which the stimulation occurred was generally lengthened as compared to the previous ISIs. The stimulation was most effective when delivered early (20-30 ms) after a spike. It was also effective, although less so, when delivered at the end of the ISI (70-100 ms after a spike). The lengthening effect sometimes extended over one or two of the subsequent ISIs. The lengthening effect of the motor axon stimulation was followed by an excitatory-like effect, which took the form of a shortening that affected up to five ISIs after the stimulation. The biphasic inhibitory effects and the subsequent facilitatory effects are discussed in terms of the dual nature of the synaptic processes involved in the recurrent inhibitory network, the postactivation facilitation/depression processes and the mutual inhibition occurring between Renshaw cells.

Voluntary contractions induce thixotropic changes in intrafusal muscle fibres and hence, by induction or removal of "slack", the background discharge and sensitivity of spindle endings to stretch is altered. This study assessed whether such changes also altered the "excitability" of the motor cortex. Eleven subjects performed a series of voluntary conditioning contractions of the wrist flexors designed to remove slack in the intrafusal fibres (contract and test at intermediate length, termed "contract-test") or to introduce slack (contract at long length and test at intermediate length, termed "contract-long"). Surface electromyographic recordings were made from one wrist flexor, flexor carpi radialis. Subjects relaxed after each contraction, and 10 s later a test stimulus was applied to elicit a tendon tap response, H-reflex, or motor-evoked potential (MEP) to transcranial magnetic stimulation in the flexor carpi radialis. Each of the three test stimuli was applied during 15 consecutive pairs of contractions ("contract-long" and "contract-test"). Three subjects repeated the protocol using transmastoid electrical stimulation as the test stimulus to evoke a cervicomedullary motor-evoked potential (CMEP). For the group of subjects, after conditioning contractions designed to induce slack there was a significant reduction in the amplitude of the tendon reflex, no significant change in the H-reflex, and a small but significant reduction in the amplitude of the MEP. In one subject the CMEP was significantly reduced, while it was unchanged in two others. In the absence of corresponding changes in the H-reflex (or CMEP), changes in the size of the response to motor cortical stimulation suggest that the level of motor cortical "excitability" changes according to naturally induced variations in the discharge of muscle spindle afferents.

1. The primary aim of the present study was to explore whether in healthy subjects the muscle contractions required for unrestrained voluntary wrist dorsiflexions are adjusted in strength to thixotropy-dependent variations in the short-range stiffness encountered in measurements of passive torque resistance to imposed wrist dorsiflexions. 2. After a period of rest, only the first movement in a series of passive wrist dorsiflexions of moderate amplitude exhibited clear signs of short-range stiffness in the torque response. During analogous types of voluntary movements, the extensor EMG during the first movement after rest showed a steep initial rise of activity, which apparently served to compensate for the short-range stiffness. 3. The passive torque resistance to minute repetitive wrist dorsiflexions (within the range of short-range stiffness) was markedly reduced after various types of mechanical agitation. During analogous low-amplitude voluntary wrist dorsiflexions the extensor EMG signals were weaker after than before agitation. 4. Mechanical agitation also led to enhancement of passive dorsiflexion movements induced by weak constant torque pulses. In an analogous way, the movement-generating capacity of weak voluntary extensor activations (as determined by EMG recordings) was greatly enhanced by mechanical agitation. 5. The signals from a force transducer probe pressed against the wrist flexor tendons--during passive wrist dorsiflexions--revealed short-range stiffness responses which highly resembled those observed in the torque measurements, suggesting that the latter to a large extent emanated from the stretched, relaxed flexor muscles. During repetitive stereotyped voluntary wrist dorsiflexions, a close correspondence was observed between the degree of short-range stiffness as sensed by the wrist flexor tension transducer and the strength of the initial extensor activation required for movement generation. 6. The results provide evidence that the central nervous system in its control of voluntary movements takes account of and compensates for the history-dependent degree of inherent short-range stiffness of the muscles antagonistic to the prime movers.

Impulses of human single muscle spindle afferents were recorded from the m. extensor carpi radialis, while 1 Hz sinusoidal movements for a wide range of amplitudes (0.05-10 deg, half of the peak-to-peak amplitude) were imposed at the wrist joint. The response was considered as linear when the discharge was approximately sinusoidally modulated. The linearity was further checked by a linear increase in the response with the amplitude and a constancy of the phase and mean level. Fifteen of 25 primary afferents were active at rest with a mean rate of 10.6 impulses s-1 (median). The linear response to sinusoidal stretching was limited to amplitudes lower than about 1.0 deg. The sensitivity was 5.6 impulses s-1 deg-1 (median) in the linear range and decreased at larger amplitudes. The other 10 primary afferents were silent at rest and lacked a linear response at low amplitudes. Nine secondary afferents were active at rest with a mean rate of 9.5 impulses s-1. The linear range extended up to about 4.0 deg with a sensitivity of 1.4 impulses s-1 deg-1. In the linear range, the phase advance of the response to sinusoidal stretching was about 50 deg and was similar between the two types of spindle afferents. In primary afferents, the phase advance increased to nearly 90 deg outside the linear range. The findings suggest that high sensitivity to small stretches is important in determining primary afferent firing during natural movements in intact humans.

1. A postural hand tremor of enhanced size was induced in eleven subjects. There was rhythmic activity in the posturally active extensor muscles synchronised to the tremor oscillation, which implied an oscillatory modulation of motor neurones. 2. The subjects executed single rapid wrist flexion movements in response to a flash of light. The light flash was presented at an instant when the wrist was spontaneously moving in the flexion direction, extension direction or at random. The time taken to generate a movement was not significantly different or more consistent in any of the conditions. 3. Inspection of individual and averaged acceleration and EMG records strongly suggests that the movement is made at a time that is independent of the tremor cycle at the time of stimulus presentation or at the moment of movement initiation. 4. These observations suggest that the central or spinal mechanism generating tremor does not gate the central mechanism that produces voluntary movement.

1. The activity of 36 pairs of single motor units were recorded with intramuscular wire electrodes from m. extensor carpi radialis while subjects performed slow wrist extension and flexion movements. Periods of steady position holding were interposed between movements. 2. The discharge trains from pairs of motor units were analysed statistically in the time and frequency domains. During extension movements, when the muscle recorded from was the agonist, coherence between motor units was significant below 12 Hz, with a peak at 6-12 Hz in 30 of 36 pairs (83 %). The magnitude of coherence decreased during position holding compared to movements in 26 pairs, while the difference in average firing rate was small. 3. During movements, but not during position holding, coherence estimates between single motor units and acceleration showed a significant peak at 6-12 Hz in 56 out of 62 motor units, suggesting that a modulation of motor unit discharge contributed to angular acceleration at these frequencies. Common motor unit modulation was present at 3 Hz as well, although the coupling between motor unit activity was weaker than at 6-12 Hz. 4. It is concluded that a 6-12 Hz common modulation of agonist motor units is a distinguishing feature of slow voluntary wrist movements, extending the previously established notion of an 8-10 Hz rhythmic organization of slow finger movements to more proximal limb segments. It is suggested that the 6-12 Hz input is specific for movements and is normally absent or much weaker during steady maintenance of position or force.

1. Growing evidence points toward involvement of the human motor cortex in the control of the ipsilateral hand. We used focal transcranial magnetic stimulation (TMS) to examine the pathways of these ipsilateral motor effects. 2. Ipsilateral motor-evoked potentials (MEPs) were obtained in hand and arm muscles of all 10 healthy adult subjects tested. They occurred in the finger and wrist extensors and the biceps, but no response or inhibitory responses were observed in the opponens pollicis, finger and wrist flexors and the triceps. 3. The production of ipsilateral MEPs required contraction of the target muscle. The threshold TMS intensity for ipsilateral MEPs was on average 1.8 times higher, and the onset was 5.7 ms later (in the wrist extensor muscles) compared with size-matched contralateral MEPs. 4. The corticofugal pathways of ipsilateral and contralateral MEPs could be dissociated through differences in cortical map location and preferred stimulating current direction. 5. Both ipsi- and contralateral MEPs in the wrist extensors increased with lateral head rotation toward, and decreased with head rotation away from, the side of the TMS, suggesting a privileged input of the asymmetrical tonic neck reflex to the pathway of the ipsilateral MEP. 6. Large ipsilateral MEPs were obtained in a patient with complete agenesis of the corpus callosum. 7. The dissociation of the pathways for ipsilateral and contralateral MEPs indicates that corticofugal motor fibres other than the fast-conducting crossed corticomotoneuronal system can be activated by TMS. Our data suggest an ipsilateral oligosynaptic pathway, such as a corticoreticulospinal or a corticopropriospinal projection as the route for the ipsilateral MEP. Other pathways, such as branching of corticomotoneuronal axons, a transcallosal projection or a slow-conducting monosynaptic ipsilateral pathway are very unlikely or can be excluded.

1. The reaction time to a visual stimulus shortens significantly when an unexpected acoustic startle is delivered together with the 'go' signal in healthy human subjects. In this paper we have investigated the physiological mechanisms underlying this effect. If the commands for the startle and the voluntary reaction were superimposed at some level in the CNS, then we would expect to see alterations in the configuration of the voluntary response. Conversely, if the circuit activated by the startling stimulus is somehow involved in the execution of voluntary movements, then reaction time would be sped up but the configuration of the motor programme would be preserved. 2. Fourteen healthy male and female volunteers were instructed to react as fast as possible to a visual 'go' signal by flexing or extending their wrist, or rising onto tiptoe from a standing position. These movements generated consistent and characteristic patterns of EMG activation. In random trials, the 'go' signal was accompanied by a very loud acoustic stimulus. This stimulus was sufficient to produce a startle reflex when given unexpectedly on its own. 3. The startling stimulus almost halved the latency of the voluntary response but did not change the configuration of the EMG pattern in either the arm or the leg. In some subjects the reaction times were shorter than the calculated minimum time for processing of sensory information at the cerebral cortex. Most subjects reported that the very rapid responses were produced by something other than their own will. 4. We conclude that the very short reaction times were not produced by an early startle reflex adding on to a later voluntary response. This would have changed the form of the EMG pattern associated with the voluntary response. Instead, we suggest that such rapid reactions were triggered entirely by activity at subcortical levels, probably involving the startle circuit. 5. The implication is that instructions for voluntary movement can in some circumstances be stored and released from subcortical structures.

1. Muscle receptors play an important role in our conscious perception of movement, but there are no published accounts of our ability to detect their signals during different motor tasks. The present experiments introduce a method to test muscular sense when humans move. 2. Muscle receptors were excited by an electrically induced twitch of the right extensor carpi ulnaris muscle. The muscle was stimulated via percutaneously inserted intramuscular electrodes or using surface stimulation through anaesthetized skin. Muscular sense was represented by the ability to detect the twitch and was compared between various tasks and stationary control trials. 3. Three hertz voluntary wrist movements significantly attenuated muscular sense to 37 % of control. This velocity-dependent attenuation was present over a range of twitch amplitudes suggesting it does not simply reflect a masking of low intensity stimuli. Perceptual ratings of twitch amplitude during fast imposed passive movements were reduced by 40 %, though this did not quite reach statistical significance. However, perceptual ratings of twitches evoked up to 2 s after the termination of the passive movements were significantly different from control. 4. Reaching with the stimulated, but not the contralateral, arm also significantly reduced muscular sense (to 40 %). 5. Attenuation to 58 % of control during cyclic stretching of the skin on the dorsum of the hand showed that signals from peripheral receptors may play a role. Attenuation prior to a single wrist flexion movement indicated that central sources can also contribute. 6. The results are consistent with current findings of a general attenuation of sensory feedback during movement and raise questions regarding the role of muscular sense in movement control.

1. Single motor unit activity was investigated in the extensor carpi radialis muscles during voluntary isometric contraction involving either the coactivation of the wrist agonist extensor muscles (wrist extension) or the coactivation of the wrist and finger antagonist extensor and flexor muscles (hand clenching). 2. The motor units were found to be activated at a similar level of motoneurone pool drive during both wrist extension and hand clenching, as indicated by the fact that the EMG activity at which they were recruited was practically the same in both cases (mean +/- S.D.: 20 +/- 26 and 21 +/- 25 mV, respectively). In addition, the net excitatory drive exerted on the motoneurones, as assessed from the mean interspike intervals, did not differ significantly between the two tasks (mean +/- S.D.: 104.57 +/- 17.24 and 103.01 +/- 16.26 ms, for wrist extension and hand clenching, respectively). 3. However, the discharge variability, in terms of the coefficient of variation of the interspike intervals, was slightly but significantly greater during hand clenching than during wrist extension (0.213 +/- 0.049 and 0.198 +/- 0.045, respectively). This increase involved all types of motor units, regardless of their contractile force. 4. We suggest that the greater motoneurone discharge variability observed during hand clenching may be attributable to an increase in the synaptic noise. This increase might be due to the activation of numerous afferent pathways mediating reciprocal interactions between antagonist motoneurone pools, as well as to the activation of hand cutaneous receptors that play a major role in the regulation of handling and gripping motor activities.

1. Effects of conditioning motor volleys of increasing amplitude on antagonistic motor nuclei were tested at the wrist and elbow level in man. 2. The modifications of biceps, triceps, flexor carpi radialis (FCR) and extensor carpi radialis (ECR) motoneurone excitability were tested both in post-stimulus time histogram (PSTH) and rectified averaged EMG experiments. 3. Conditioning motor volleys were evoked in biceps, triceps, FCR and ECR muscles by electrical stimuli applied to the corresponding nerves. The intensity of the conditioning stimuli was systematically varied from an intensity just below motor threshold to one giving rise to a motor response equal to half of the maximal direct motor response (Mmax). 4. The effect of the injection of a cholinergist agonist, L-acetylcarnitine (L-Ac), on antagonistic conditioning motor volleys was tested for each motor nucleus. 5. Results obtained at the wrist and elbow level were strikingly different. Antagonistic motor volleys resulted in an early and long-lasting inhibition enhanced by L-Ac injection in wrist motor nuclei, while no modification was observed in elbow motor nuclei. 6. The characteristics of the early and long-lasting inhibition evoked in wrist motor nuclei by antagonistic motor volleys suggest that they are due to the activation of Renshaw cells by the conditioning motor volleys. 7. It is therefore concluded that these experiments provide further evidence that the interconnections between motoneurones, interneurones mediating reciprocal inhibition and Renshaw cells at the wrist level are different from those described elsewhere. Indeed, it has already been shown that the interneurones mediating reciprocal inhibition between FCR and ECR are not inhibited by Renshaw cells, and the results presented here suggest that FCR-coupled Renshaw cells inhibit ECR motoneurones and vice versa.

1. The possibility was investigated that, in man, some of the descending command for tonic voluntary wrist extension is transmitted to extensor motoneurones over a non-monosynaptic pathway. 2. Stimulation of the cutaneous superficial radial nerve at 3 times perceptual threshold depressed the electromyogram (EMG) of extensor carpi radialis (ECR) and the discharge of single ECR motor units, both with a mean central delay of 4.2 ms. Such stimuli depressed the response to transcranial magnetic stimulation of the motor cortex, but had little effect on the H reflex. 3. The possibility that the relative sparing of the H reflex was due to an alteration in transmission of the afferent volley for the H reflex was excluded. 4. The central latency of the cutaneous-induced depression of the discharge of single motor units in biceps brachii (C5-C6) was shorter by about 1 ms than that of the more caudal wrist and finger extensor motor units. This suggests that the locus for the cutaneous-induced effects was spinal but above the cervical enlargement. 5. The pattern of EMG depression (evoked by superficial radial but not palmar stimuli, in wrist extensors but not wrist flexors) is that previously described for the presumed propriospinal system of human subjects. 6. It is concluded that a significant component of the voluntary command for tonic wrist extension reaches the relevant motoneurone pool via a non-monosynaptic pathway. It is suggested that the interposed neurones could be C3-C4 propriospinal neurones.

1. The threshold for obtaining EMG responses after transcranial magnetic stimulation of the brain is reduced by voluntary contraction of the target muscle. The present experiments tested whether some of this effect is due to increased cortical, as opposed to spinal, excitability during the contraction. 2. Magnetic stimulation was delivered with a figure-of-eight coil oriented with the junction region along the interaural line and also (in 4 of 7 subjects) with a circular coil centred at the vertex. The intensity of the conditioning stimulus was subthreshold for evoking a motor response in the relaxed wrist flexor muscles of the forearm. The presence of a small descending corticospinal volley in both the relaxed and active conditions was detected by measuring the facilitation of test H reflexes elicited in the flexor muscles of the forearm. 3. In all subjects, magnetic stimulation with either coil facilitated the H reflex at conditioning-test intervals of -1 to -3 ms (median nerve stimulus before magnetic). This was followed by a long-lasting facilitation. In three of the seven subjects stimulation with the figure-of-eight coil elicited an additional, earlier peak of facilitation at a conditioning-test interval of -3 to -5 ms. 4. In all subjects, the threshold for obtaining facilitation of the H reflex using a conditioning-test interval of -1 to -3 ms was reduced, and the amount of facilitation was larger, if subjects performed a weak tonic voluntary contraction. In contrast, with a conditioning-test interval of -3 to -5 ms voluntary contraction had no effect on the threshold. 5. It is suggested that H reflex facilitation at the conditioning-test interval of -1 to -3 ms was produced by indirect activation of corticospinal neurones by the magnetic stimulus, whereas at -3 to -5 ms, the facilitation was produced by direct activation of corticospinal axons. It is concluded that tonic voluntary contraction of a target muscle decreases the threshold for indirect activation of corticospinal neurones but not for direct stimulation of their axons.

1. The alterations in voluntary wrist extension and flexion movement trajectories induced by application of vibration to the tendon of flexor carpi radialis throughout the course of the movement, together with the associated EMG patterns, have been studied in normal human subjects. Both extension and flexion movements were routinely of a target amplitude of 30 deg and made against a torque load of 0.32 N m. Flexor tendon vibration consistently produced undershooting of voluntary extension movements. In contrast, voluntary flexion movements were relatively unaffected. 2. The degree of vibration-induced undershooting of 1 s voluntary extension movements was graded according to the amplitude (0.75, 1.0 and 1.5 mm) of flexor tendon vibration. 3. As flexor vibration was initiated progressively later (at greater angular thresholds) during the course of 1 s voluntary extension movements, and the period of vibration was proportionately reduced, so the degree of vibration-induced undershooting showed a corresponding decline. 4. Varying the torque loads (0.32, 0.65 and 0.97 N m) against which 1 s extension movements were made, and thereby the strength of voluntary extensor contraction, produced no systematic changes in the degree of flexor vibration-induced undershooting. 5. Analysis of EMG patterns recorded from wrist flexor and extensor muscles indicated that vibration-induced undershooting of extension movements resulted largely from a reduction in activity in the prime-mover rather than increased antagonist activity. The earliest reductions in extensor EMG commenced some 40 ms after the onset of vibration, i.e. well before voluntary reaction time; these initial responses were considered to be 'automatic' in nature. 6. These results support the view that the central nervous system utilizes proprioceptive information in the continuous regulation of moderately slow voluntary wrist movements. Proprioceptive sensory input from the passively lengthening antagonist muscle, presumably arising mainly from muscle spindle I a afferents, appears to be particularly important and to act mainly in the reciprocal control of the prime-mover.

1. Experiments were performed to check the validity of previous claims concerning the ;long-loop' aetiology of ;late' reflex electromyogram (e.m.g.) responses to muscle stretch in man. The primary aim was to investigate whether observations previously presented in favour of the ;long-loop hypothesis' are explicable also in terms of the ;resonance hypothesis', according to which the ;late' reflex components represent spinal, short-latency responses to intramuscular oscillations initiated by the impact.2. The contracting wrist flexors of healthy subjects were exposed to trains of recurrent 25-50 Hz stretch stimuli (wrist torque pulses). Each of the initial two or three pulses in the train was followed by e.m.g. peaks with a latency of 20-25 msec. The e.m.g. peaks driven in this way had the following characteristics in common with the successive two or three e.m.g. peaks which were induced by single ramp stretches or tendon taps. (a) Changes in stimulus parameters which altered the strength of the initial e.m.g. peak often had an opposite effect on the strength of the succeeding peak(s). Muscle vibration which attenuated the initial peak often enchanced the succeeding one(s). (b) The initial e.m.g. peak was less affected than the succeeding peak(s) by the subjects' attempts to respond with rapid ;resist' or ;let go' reactions.3. Intramuscular oscillations (monitored by a needle accelerometer) and e.m.g. responses evoked by single ramp stretches and/or tendon taps were also studied in the long thumb flexor, the calf muscles and the masseter muscle. In the thumb flexor, the initial accelerometer deflexion was only rarely succeeded by a short latency e.m.g. peak, but the succeeding wave in the needle accelerogram was followed by such a peak, appearing about 40 msec after stimulus application. By contrast, the calf muscles and the jaw elevators exhibited a high amplitude, short-latency e.m.g. response to the first but only rarely to the second intramuscular oscillation wave.4. The interval between initial and second e.m.g. peaks following tendon taps was longer for calf muscles than for wrist flexors and longer for wrist flexors than for jaw elevators. Similar differences were observed with respect to the intervals between the damped intramuscular oscillations initiated by the impacts.5. Without denying the existence of ;long-loop reflexes' it is concluded that the characteristics of the ;late' reflex responses to muscle stretch in man are explicable also in terms of the ;resonance hypothesis'.

The resonance of the relaxed wrist for flexion-extension movements in the horizontal plane has been investigated by using rhythmic torques generated by a printed motor. In the normal subject the resonant frequency of the wrist is ca. 2 Hz unless the torque is reduced below a certain critical value when the system is no longer linear and the resonant frequency rises. This critical torque level, and the damping are both less in women than men. The resonant frequency is uninfluenced by surgical anaesthesia. With added bias the increase of resonant frequency at low torques still occurs although the hand is now oscillating about a displaced mean position. It follows that the stiffening implied by this elevation of resonant frequency for small movements is neither the result of pre-stressing of the muscles nor of reflex activity. With velocity feed-back of appropriate polarity the system will oscillate spontaneously at its resonant frequency. If the peak driving torque is progressively reduced the resonant frequency increases abruptly, indicating that the system has stiffened. Perturbations delivered to the wrist may reduce its stiffness. The postural system is thixotropic with a 'memory time' of 1-2 s. The resonant frequency is elevated in voluntary stiffening.

1. Brisk joint displacements and tendon taps were applied to voluntarily contracting wrist flexor muscles in subjects who did not attempt to react to them. Different types of mechanotransducers, some of them attached to an intramuscular needle, were used to detect mechanical oscillations in the wrist flexors, resulting from the imposed impacts. The transducer responses to the perturbations were compared with simultaneously recorded reflex electromyogram (e.m.g.) responses. Experiments were also carried out on a rubber band model, exposed to similar mechanical stimuli.2. During brisk ramp wrist extensions the transducers signalled damped muscular oscillations at 30-50 Hz. The oscillations grew in amplitude with increasing speed of onset of the stretch movement and at angular accelerations exceeding about 2 x 10(4) deg/sec(2) the e.m.g. response changed from a non-segmented to a progressively more pronounced segmented pattern. Peak intervals in the segmented reflex e.m.g. responses were similar to those of the mechanical oscillations and did not change significantly with small or moderate variations in background contraction force. Latencies from successive deflexions in the accelerometer records to corresponding deflexions in the e.m.g. were 20-25 msec.3. Damped muscular oscillations in the 30-50 Hz range were also initiated by sudden halts of voluntary wrist movements, by electrically induced twitches, and by voluntary brisk contractions. In these instances too, the mechanical oscillations were reflected in the shape of the succeeding e.m.g. response.4. The interval between the two initial peaks in the accelerometer records was always shorter with tendon taps than in ramp stretch experiments. A corresponding difference was noted in the intervals between the following two peaks in the reflex e.m.g. response.5. The initial peak in the accelerometer records could be ascribed to a wave propagated at about 40 m/sec in the wrist flexor muscles. Inconclusive results were obtained in attempts to determine whether the subsequent oscillations represented reflexions of the propagated wave at the ends of the muscle.6. The muscles were also exposed to recurrent stretch stimuli (torque pulses) with a repetition rate varying between 15 and 100 Hz. At 30-50 Hz the intramuscular oscillations reached their maximal amplitude, and such repetition rates were also most efficient in producing synchronized e.m.g. bursts, time-locked to the oscillations. The mechanical responses of the wrist flexors to single or recurrent perturbations were to a large extent mimicked by a rubber band model with a longitudinal resonance frequency of about 40 Hz.7. It is concluded that segmentation of reflex e.m.g. responses to sudden joint displacements and other types of brisk muscle perturbations to a large extent depends on the inherent resonance characteristics of musculo-tendinous structures. Primary spindle endings with their high vibration sensitivity and their segmental projections to alpha-motoneurones are believed to be the receptors primarily responsible for reflex entrainment of the motor impulses.

To test the hypothesis that a transcortical reflex contributes to the stretch-evoked long-latency electromyographic (e.m.g.) response we documented the responses of identified corticomotoneuronal (c.m.) cells and their target muscles to perturbations of active wrist movements. Macaque monkeys performed ramp-and-hold wrist movements against elastic loads, alternating between flexion and extension zones; brief (25 ms) torque pulses were intermittently applied during the hold period. C.m. cells were identified by a clear post-spike facilitation in spike-triggered averages of forelimb muscle e.m.g. activity. Activity of c.m. cells and twelve wrist and digit flexor and extensor muscles was recorded during: (a) active ramp-and-hold wrist movements, (b) passive ramp-and-hold wrist movements, and (c) torque perturbations applied during the hold phase of active flexion and extension which either lengthened or shortened the c.m. cell's target muscles. Muscle-lengthening perturbations evoked a reproducible pattern of average e.m.g. activity in the stretched muscles, consisting of two peaks: the first response (M1) had an onset latency of 11.2 +/- 2.1 ms (mean +/- S.D.), and the second (M2) began at 27.9 +/- 5.1 ms. Torque perturbations which shortened the active muscles also evoked a characteristic e.m.g. response consisting of an initial cessation of activity at 13.5 +/- 3.4 ms followed by a peak beginning at 33.9 +/- 3.0 ms. The responses of twenty-one c.m. cells which facilitated wrist muscles were documented with torque pulse perturbations applied during active muscle contraction. Twenty of twenty-one c.m. cells responded at short latency (23.4 +/- 8.8 ms) to torque perturbations which stretched their target muscles. For each c.m. cell-target muscle pair, transcortical loop time was calculated as the sum of the onset latency of the c.m. cell's response to lengthening perturbations (afferent time) and the onset latency of post-spike facilitation (efferent time). The mean transcortical loop time was 30.4 +/- 10.2 ms, comparable to the mean onset latency of the M2 peak (27.9 +/- 5.1). The duration of a c.m. cell's response to torque perturbations provides a further measure of the extent of its potential contribution to the M2 muscle response. In all cases but two, the c.m. cell response, delayed by the latency of the post-spike facilitation, overlapped the M2 e.m.g. peak.

1. Impulses in single muscle afferents were recorded from the median nerves of waking human subjects with percutaneously inserted tungsten needle electrodes. During isometric voluntary contractions, unitary discharges were analysed from muscle spindle endings in the wrist and finger flexor muscles and the electromyographic activity from these muscles was recorded simultaneously.2. When the subject activated the muscle portion in which a spindle was located, the afferent discharge increased in spite of the mechanical unloading effects of the skeletomotor contraction indicating a concomitant fusimotor activation. This was valid for slowly rising contractions as well as small fast rising twitches.3. The time of onset of spindle acceleration was determined in relation to the time of onset of the electromyographic activity for thirty-one units studied altogether in more than seven hundred contractions. It was found that spindle acceleration regularly occurred after the onset of the electromyographic activity.4. There was a considerable variation from one test to the other, for the individual units, with regard to the exact time of onset of spindle acceleration, although spindle acceleration occurred mostly within 0.5 sec after the onset of the electromyographic activity in sustained contractions and within 0.1 sec in small fast rising twitches. It was not possible to assess to what extent this variation was accounted for by variations in the mechanical unloading effects of the skeletomotor contraction or variations in the timing of the fusimotor outflow.5. For many units, spindle acceleration did not occur until 10-50 msec after the onset of the skeletomotor contraction. This time is of the same order of magnitude as the time difference in latency from the spinal cord to the recording points in the two systems, as estimated from reasonable assumptions.6. It was concluded that the fusimotor system does not participate in the initiation of voluntary contractions in man, but that the skeletomotor activity is initiated by descending impulses from supraspinal structures and their effects on the neuronal organization within the spinal cord.7. The fact that fusimotor activation occurs also in very small and short lasting twitches, when spindle acceleration must have a negligible influence on the skeletomotor outflow, suggests that the fusimotor and the skeletomotor systems are rigidly co-activated in voluntary contractions.8. The finding that spindle acceleration does not occur until 10-50 msec after the onset of the electromyographic activity suggests that there is an approximately simultaneous onset of the fusimotor and the skeletomotor outflows from the spinal cord.