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The effect of perturbation acceleration and advance warning on the neck postural responses of seated subjects.

The muscle and kinematic responses of subjects exposed to postural perturbations have been shown to vary with platform acceleration when this acceleration was covaried with platform velocity or displacement. The purpose of the current study was to isolate platform acceleration and examine its effect on the neck muscle response and head kinematics of seated subjects exposed to anterior perturbations. Thirty-six subjects (20 females, 16 males) underwent two blocks of 36 perturbations. Three different perturbations with peak accelerations of 7.7, 14.7, and 21.7 m/s(2) up to a common velocity of 0.5 m/s were used. In one block, subjects received an audible warning corresponding to the platform acceleration magnitude, and in the other block, no advance warning was given. Onset and amplitude of the sternocleidomastoid and cervical paraspinal muscle responses were measured using surface electromyography. Kinematic measures included linear and angular accelerations and displacements of the head. The results showed no differences in either the preperturbation posture or the muscle or kinematic responses between the warned and unwarned trials. Significant differences were observed in the onset and amplitude of the muscle and kinematic variables with perturbation acceleration, although these response differences were not linearly graded with perturbation acceleration. Gradation of muscle activation times has not been previously observed in postural perturbation studies, and their gradation with platform acceleration in the current study suggested that platform acceleration was a strong regulator of the reflex muscle response in postural perturbations.

Acceleration↗

Experimental muscle pain changes feedforward postural responses of the trunk muscles.

Many studies have identified changes in trunk muscle recruitment in clinical low back pain (LBP). However, due to the heterogeneity of the LBP population these changes have been variable and it has been impossible to identify a cause-effect relationship. Several studies have identified a consistent change in the feedforward postural response of transversus abdominis (TrA), the deepest abdominal muscle, in association with arm movements in chronic LBP. This study aimed to determine whether the feedforward recruitment of the trunk muscles in a postural task could be altered by acute experimentally induced LBP. Electromyographic (EMG) recordings of the abdominal and paraspinal muscles were made during arm movements in a control trial, following the injection of isotonic (non-painful) and hypertonic (painful) saline into the longissimus muscle at L4, and during a 1-h follow-up. Movements included rapid arm flexion in response to a light and repetitive arm flexion-extension. Temporal and spatial EMG parameters were measured. The onset and amplitude of EMG of most muscles was changed in a variable manner during the period of experimentally induced pain. However, across movement trials and subjects the activation of TrA was consistently reduced in amplitude or delayed. Analyses in the time and frequency domain were used to confirm these findings. The results suggest that acute experimentally induced pain may affect feedforward postural activity of the trunk muscles. Although the response was variable, pain produced differential changes in the motor control of the trunk muscles, with consistent impairment of TrA activity.

Adult↗

Postural adjustments due to external perturbations during sitting in 1-month-old infants: evidence for the innate origin of direction specificity.

The aim of the study was to examine whether infants, at an age when they have no or little experience in sitting, can produce direction specific postural adjustments, i.e. synergies of muscle activity on the ventral side of the body during backward sway and on the dorsal side during forward sway. In addition, we addressed the question whether postural adjustments at this young age are restricted to single muscle responses or consist of a variable repertoire of muscle activation patterns including one during which all direction specific muscles participate ('complete' pattern). Postural adjustments due to external perturbations in a sitting position were studied in eight healthy infants aged 1 month. Multiple surface EMGs of neck, trunk and leg muscles and kinematics were recorded while the infants were exposed to horizontal forward (Fw) and backward (Bw) displacements of the surface of support. Direction specific postural adjustments, defined as adjustments during which agonist activation or antagonist inhibition preceded antagonist activation, were present in 85% of Bw and 72% of Fw translations. The direction specific adjustments showed a large variability with the repertoire of adjustments including the activation of one, two or all of the recorded direction specific muscles. The finding of direction specific adjustments at 1 month of age support the opinion that the basic level of organisation of postural adjustments has an innate origin. The finding of a variable repertoire of muscle response patterns, including the 'complete' pattern, refutes the idea that the development of postural adjustments results from gradual addition of appropriate muscles to the synergies.

Biomechanical Phenomena↗

Muscle modes during shifts of the center of pressure by standing persons: effect of instability and additional support.

Muscle synergies in postural tasks have recently been studied using the framework of the uncontrolled manifold (UCM) hypothesis. A set of three hypothetical control variables, named M-modes, derived from the activity of 11 postural muscles, were identified. It was shown that postural synergies composed of these three M-modes preserve a certain shift of the center of pressure (COP) when subjects perform postural tasks while standing on a stable surface. In the present study we investigated the effects of support surface instability and availability of a light touch or grasp of a stable external support on the M-modes and their co-variation. The study was performed in two sessions. In the first session subjects released a load behind the body under four conditions: standing on a stable surface with no support (ST), standing on an unstable surface with no support (UN), standing on an unstable surface with a light touch (UN,T) and standing on an unstable surface with grasp of a stable object (UN,G). In the second session subjects performed two tasks: an arm movement backward and voluntary sway forward (towards the toes) under three conditions--ST, UN and UN,T. Principal component analysis was used to identify M-modes from data in the first session, and a UCM analysis was performed to study M-mode synergies in postural stabilization from data in the second session. A 'menu' of five M-modes was found, which were named either reciprocal M-modes or co-contraction M-modes based on the agonist-antagonist relationship of muscles comprising each mode. For a given task, subjects chose any three of these five M-modes in a subject- and task-specific manner. The reciprocal and co-contraction M-modes occurred equally frequently whether subjects stood on a stable or unstable support surface or whether a light touch was available or not. However, the co-contraction M-modes predominated when grasp of an object was available. In this condition, when the arm could be used for stabilization, there were M-modes uniting hip and shoulder muscles. However, the identified M-mode synergies were not found to lead to a consistent shift in the COP in any of the stability conditions. Possible reasons for this finding are discussed.

Adult↗

Feedforward activity of the cervical flexor muscles during voluntary arm movements is delayed in chronic neck pain.

The objective of this study was to compare onset of deep and superficial cervical flexor muscle activity during rapid, unilateral arm movements between ten patients with chronic neck pain and 12 control subjects. Deep cervical flexor (DCF) electromyographic activity (EMG) was recorded with custom electrodes inserted via the nose and fixed by suction to the posterior mucosa of the oropharynx. Surface electrodes were placed over the sternocleidomastoid (SCM) and anterior scalene (AS) muscles. While standing, subjects flexed and extended the right arm in response to a visual stimulus. For the control group, activation of DCF, SCM and AS muscles occurred less than 50 ms after the onset of deltoid activity, which is consistent with feedforward control of the neck during arm flexion and extension. When subjects with a history of neck pain flexed the arm, the onsets of DCF and contralateral SCM and AS muscles were significantly delayed ( p<0.05). It is concluded that the delay in neck muscle activity associated with movement of the arm in patients with neck pain indicates a significant deficit in the automatic feedforward control of the cervical spine. As the deep cervical muscles are fundamentally important for support of the cervical lordosis and the cervical joints, change in the feedforward response may leave the cervical spine vulnerable to reactive forces from arm movement.

Adult↗

Modulatory effect of repetitive peripheral magnetic stimulation on skeletal muscle tone in healthy subjects: stabilization of the elbow joint.

To investigate the role of repetitive peripheral magnetic stimulation (RPMS) on the postural component of motor performances, the long-lasting modulatory effect of RPMS on the stabilization of the elbow joint was examined in 13 healthy subjects. The resistance against very slow passive movements in the relaxed state was recorded simultaneously with the electromyogram (EMG) of the forearm extensor and flexor muscles. The experiments show that RPMS performed on the forearm flexor muscles increased the degree of stabilization of the elbow joint, whereas RPMS on the forearm extensor muscles caused a decrease in stabilization. This leads to the assumption that the postural component of motor tasks depends on the motor task itself: motor tasks like manipulation, pointing or grasping which are fine skilled movements require an increase in stabilization while goal-directed movements require a decrease in stabilization. Therefore RPMS is involved in sensorimotor integration and may modulate the motor program at the cortical level.

Adult↗

Can prepared responses be stored subcortically?

Quick voluntary responses to environmental stimuli are required of people on a daily basis. These movements have long been thought to be controlled via cortical loops involving processing of the stimulus and generation of a suitable response. Recent experiments have shown that in simple reaction time (RT) tasks, the appropriate response can be elicited much earlier (facilitated) when the "go" signal is replaced by a startling (124 dB) auditory stimulus. In the present experiment we combined a startling acoustic stimulus with an established RT paradigm that involved simple and choice RT. In a simple RT condition the prepared voluntary response was elicited at very short latencies following the startle. However, when cortical processing was required prior to responding (choice RT task), the startle did not facilitate the voluntary response, and gave rise to more movement production errors. Since movements requiring ongoing cortical processing following the stimulus are not facilitated by startle, it is unlikely that the startle facilitation is due to increased neural activation. In contrast, it appears more likely that the startle acts as an early trigger for subcortically stored prepared movements since movements that are prepared in advance can be initiated at such short latencies (<60 ms).

Acoustic Stimulation↗

Muscle synergies involved in shifting the center of pressure while making a first step.

We used the framework of the uncontrolled manifold (UCM) hypothesis to analyze multi-muscle synergies involved in making a step by a standing person. We hypothesized that leg and trunk muscles are organized into stable groups (muscle modes, M-modes) related to shifts of the center of pressure (COP) in the anterior-posterior and medio-lateral directions. Another hypothesis was that the magnitudes of the modes co-vary across repetitive trials to stabilize a certain magnitude of the COP shift in both directions. M-modes were defined using principal component analysis applied to indices of changes in the electromyographic (EMG) activity prior to releasing variable loads that were held by the subject using a pulley system. For the task of releasing the load behind the body three M-modes associated with a backward COP shift were defined. Four M-modes were defined for the task of releasing the load at the body side associated with a lateral COP shift. Multiple regression analysis was used to relate changes in the M-mode magnitudes to COP shifts. EMG changes prior to making a step were quantified over five 100 ms time windows before the lift-off of the stepping leg. Two components of the variance in the M-mode space computed across repetitions of a stepping task were quantified-a component that did not affect the average COP shift in a particular direction (variance within the UCM, V (UCM)), and a component that affected the COP shift (variance orthogonal to the UCM, V (ORT)). V (UCM) was significantly higher than V (ORT) for both directions of the COP shifts. This relation was observed for the M-modes in the stepping leg as well as in the support leg. The stepping leg showed a different time evolution of the ratio V (UCM)/V (ORT) such that the difference between the two variance components disappeared closer to the time of the lift-off. The findings corroborate both main hypotheses. The study supports a view that control of whole-body actions involves grouping the muscles, using fewer elemental variables to scale the muscle activity, and forming synergies in the space of the elemental variables that stabilize time profiles of important performance variables.

Adult↗

Impedance is modulated to meet accuracy demands during goal-directed arm movements.

The neuromuscular system is inherently noisy and joint impedance may serve to filter this noise. In the present experiment, we investigated whether individuals modulate joint impedance to meet spatial accuracy demands. Twelve subjects were instructed to make rapid, time constrained, elbow extensions to three differently sized targets. Some trials (20 out of 140 for each target, randomly assigned) were perturbed mechanically at 75% of movement amplitude. Inertia, damping and stiffness were estimated from the torque and angle deviation signal using a forward simulation and optimization routine. Increases in endpoint accuracy were not always reflected in a decrease in trajectory variability. Only in the final quarter of the trajectory the variability decreased as target width decreased. Stiffness estimates increased significantly with accuracy constraints. Damping estimates only increased for perturbations that were initially directed against the movement direction. We concluded that joint impedance modulation is one of the strategies used by the neuromuscular system to generate accurate movements, at least during the final part of the movement.

Adult↗

Startle produces early response latencies that are distinct from stimulus intensity effects.

Recent experiments pairing a startling stimulus with a simple reaction time (RT) task have shown that when participants are startled, a prepared movement was initiated earlier in comparison to voluntary initiation. It has been argued that the startle acts to trigger the response involuntarily. However, an alternative explanation is that the decrease in RT may be due to stimulus intensity effects, not involuntary triggering. Thus the aim of the current investigation was to determine if RT simply declined in a linear fashion with increasing stimulus intensity, or if there was a point at which RT dramatically decreased. In the present experiment participants completed 50 active wrist extension trials to a target in response to an auditory stimulus of varying stimulus intensity (83-123 dB). The presented data show that RTs associated with a startle response are separate from stimulus intensity facilitated responses. Furthermore, this startle facilitation is more highly associated with sternocleidomastoid electromyographic (EMG) activity, rather than the EMG from the widely used startle response indicator muscle orbicularis oculi.

Acoustic Stimulation↗

Time to reconfigure balancing behaviour in man: changing visual condition while riding a continuously moving platform.

While balancing on a continuously antero-posterior (A-P) translating platform (10 cm, 0.5 Hz), the head normally oscillates with the platform without vision but is stabilized in space with vision. We estimated the time to shift from one to the other balancing behaviour when visual condition changed at some stage during the balancing trials. Ten subjects performed randomly 50 balancing trials (each lasting 18 s): 10 trials with eyes open (EO), 10 with eyes closed (EC), 15 in which participants started with EO and closed their eyes (condition EO-->EC) in response to an acoustic signal delivered during the trial, and 15 starting with EC and closing their eyes (EC-->EO) in response to the same signal. No other specific instruction was given. Displacements of malleolus, hip and head, and EMG from leg and axial muscles were recorded. Indexes of amplitude of A-P head and hip oscillation and of amplitude of EMG activity were computed. All variables were larger with EC than EO. On changing visual condition during the trial, the pattern of head and hip movement and of muscle activity turned into that appropriate for the new visual condition in a time-interval ranging from about 1 to 2.5 s. For each subject, the mean latency of the change in the balancing behaviour was assessed by statistical methods. On average, the latencies of kinematics and EMG changes proved to be longer for the EO-->EC condition than vice versa. Further, the latencies of the changes were also measured across all EO-->EC and EC-->EO individual trials. These values were clustered around particular epochs of the first few oscillation cycles following the shift in visual condition. The results show that subjects can rapidly adapt their balancing behaviour to the new visual condition. However, they appear to refrain from releasing the new behaviour were this unfit, and unfastened it at appropriate time in the next platform translation cycle. These findings reveal the temporal and spatial features of the automatic release of the new balancing strategy in response to a shift in the ongoing sensory set, and emphasize the swiftness in the change in balancing behaviour when subjects pass from a non-visual to a visual reference frame.

Adult↗

The effect of voluntary arm abduction on balance recovery following multidirectional stance perturbations.

The goal of this study was to investigate how voluntarily abducting one arm, 90 degrees at onset of a rotational perturbation of the support surface, influences the recovery of upright stance. Young adults were tested under four stance conditions: abducting one arm to the horizontal only (AO); perturbation of stance using a support surface rotation only (PO); combined support surface rotation and abduction of the downhill arm, ipsilateral to tilt (IPS); and fourth abduction of the uphill, contralateral arm (CON). Simultaneous auditory and visual trigger cues were used for arm raising. Perturbations consisted of six directions of combined support surface roll and pitch rotation (7.5 degrees and 60 degrees/s). Outcome measures were whole body centre of mass (COM) movements and body segment angular displacements recorded with a motion analysis system, as well as leg, trunk, and arm EMG responses. Arm raises contralateral and ipsilateral to the direction of support surface roll were more rapid than in the AO condition and significantly reduced or increased, respectively, COM lateral displacements relative to the PO condition. The changes in COM displacements and velocities during combined CON arm raise and perturbation were greater than expected from the sum of displacements for AO and PO conditions alone, but less for the IPS condition. Arm raising increased trunk roll in a direction opposite arm raising was more than for the AO and PO conditions. Robust effects were also observed for hip abduction but not for leg flexion. Early balance correcting activity was enhanced on the side opposite arm raising and later stabilising activity reduced bilaterally in lower trunk muscles compared to summed activity for the AO and PO conditions. Similar effects were observed in gluteus medius muscles but effects were weak in ankle muscles. EMG onsets in muscles of the raised arm were earlier than in the AO conditions. We conclude that triggered arm abduction, contralateral to the direction of support surface rotation, had significant stabilization benefits for young adults and ipsilateral arm movements had destabilizing effects. The arm raises could be simultaneously executed with balance corrections. These results provide insights into the integration of balance corrections and voluntary commands into one automatic reaction that may be useful in training fall avoidance.

Adaptation, Physiological↗

Adaptation of postural orientation to changes in surface inclination.

We previously showed that standing on an inclined surface resulted in an after-effect of leaning in many healthy, blindfolded subjects when they returned to standing on a horizontal surface (Kluzik et al. in Exp Brain Res 162:474-489, 2005). The direction of leaning depended on the direction of prior surface inclination, always in a direction that preserved the relative alignment between the body and the support surface. For example, subjects leaned forward after they stood on a toes-up-inclined surface. In the present study, we investigated how the amplitude of surface inclination affected postural muscle activity, joint position, body segment orientation, and body center of mass (CoM) and foot center of pressure (CoP) locations before, during, and after subjects stood on an inclined surface. We asked whether the mechanism that underlies the lean after-effect involves regulation of local postural variables, such as the position of the ankle joint or the level of muscle activity, or whether instead, the mechanism involves regulation of global, whole-body postural variables that can only be determined by multisensory processing, such as orientation of the trunk or the body's CoM. In one experiment, we found that varying the amplitude of a toes-up surface inclination between 2.5 degrees and 10 degrees had a systematic, linear, effect on the post-incline orientation of the trunk and head, but did not systematically affect the post-incline orientation of the legs, position of the ankle joint, the level of EMG activity, or the location of the CoP. In a second experiment, we found that preventing the legs from leaning in the post-incline period did not abolish leaning of the upper body. These findings suggest that (1) the body-to-support-surface relationship is an important reference for the CNS internal representation of postural orientation which is subject to adaptive modification and (2) the adaptive mechanism underlying the post-incline after-effect of leaning acts at the level of global, whole-body postural variables.

Adaptation, Physiological↗

Experimental muscle pain results in reorganization of coordination among trapezius muscle subdivisions during repetitive shoulder flexion.

The aim of the study was to examine the effect of experimental unilateral upper trapezius muscle pain on the relative activation of trapezius muscle subdivisions bilaterally during repetitive movement of the upper limb. Surface EMG signals were detected from nine healthy subjects from the upper, middle and lower divisions of trapezius during a repetitive bilateral shoulder flexion task. Measurements were performed before and after injection of 0.5 ml hypertonic (pain condition) and isotonic (control) saline into the upper division of the right trapezius muscle in two experimental sessions. On the painful side, upper trapezius showed decreased EMG amplitude (average rectified value, ARV) and lower trapezius increased ARV throughout the entire task following the injection of hypertonic saline (40.0 +/- 22.2 vs. 26.0 +/- 17.4 microV, and 12.5 +/- 7.6 vs. 25.6 +/- 14.8 microV, respectively, at the beginning of the contraction). On the side contralateral to pain, greater estimates of ARV were identified for the upper division of trapezius as the task progressed (37.4 +/- 20.2 vs. 52.7 +/- 28.4 microV, at the end of the contraction). Muscle fiber conduction velocity did not change with pain in all three divisions of the right trapezius muscle. The results suggest that local elicitation of nociceptive afferents in the upper division of the trapezius induces reorganization in the coordinated activity of the three subdivisions of the trapezius in repetitive dynamic tasks.

Adaptation, Physiological↗

Is the human masticatory system devoid of recurrent inhibition?

The aim of the present study was to investigate the existence or otherwise of a functional recurrent inhibitory system (Renshaw cell system) in the motoneurons that innervate human masticatory muscles. In a previous study, L: -acetylcarnitine (L: -Ac), a substance known to potentiate recurrent inhibition in humans was found to alter, in a specific way, the discharge variability, and the synchronous activity of motor units depending on the presence or absence of recurrent inhibition in the corresponding motoneuron pool. Using a similar paradigm, we have recorded the tonic discharge activity of motor unit pairs from the masseter muscle during voluntary isometric contraction while subjects were undergoing continuous intravenous saline (SAL, NaCl 0.9%) perfusion. Following a brief baseline-recording period, the subjects were given a test injection of either L: -Ac or isotonic saline (SAL) in a double blind manner. The variability, synchronization, and coherence between the motor unit discharges were analysed during three successive periods: pre-injection, during injection, and post-injection, each lasting 2-3 min. Neither L: -Ac nor SAL injection induced a significant change in the inter-spike interval (ISI) or the coefficient of variation of the ISIs in the motor units tested. There were also no significant changes in the pattern of synchronous activity or in the coherence, which reflects the common frequency content of the unit discharges. Reminiscent of what had been observed previously with motoneurons without recurrent inhibition in the Abductor Digitorum Minimi muscle, the lack of effects of L: -Ac injection on the firing behaviour of masseter motoneurons may suggest that classical Renshaw cell inhibition is lacking in this motoneuron pool.

Acetylcarnitine↗

The role of synchrony and oscillations in the motor output.

There is currently much interest in the synchronisation of neural discharge and the potential role it may play in information coding within the nervous system. We describe some recent results from investigations of synchronisation within the motor system. Local field potentials (LFPs) and identified pyramidal tract neurones (PTNs) were recorded from the primary motor cortex of monkeys trained to perform a precision grip task. The LFPs showed bursts of oscillatory activity at 20-30 Hz, which were coherent with the rectified electromyographs (EMG) of contralateral hand and forearm muscles. This oscillatory synchronisation showed a highly specific task dependence, being present only during the part of the task when the animal maintained a steady grip and not during the movement phases before or after it. PTNs were phase-locked to LFP oscillations, implying that at least part of the coherence between cortical activity and EMG was mediated by corticospinal fibres. The phase locking of the PTNs to LFP oscillations produced task-dependent oscillatory synchronisation between PTN pairs, as assessed by the single-unit cross-correlation histogram. Recordings were also made from normal human subjects performing a precision grip similar to that used in the monkey recordings. Pairs of EMGs recorded from intrinsic hand and forearm muscles showed 20-30 Hz coherence, which modulated during task performance, being present only during periods of steady contraction. We suggest that these changes in EMG-EMG synchronisation reflect changing levels of synchronous drive from the corticospinal system. The generation of oscillations in the cortex is discussed in the light of results from a model of local cortical circuits. Other modelling work has shown that synchrony in the corticospinal inputs could act to recruit motoneurones more efficiently, producing more output force from a muscle than asynchronous inputs firing at the same mean rate. A speculative hypothesis is presented on the role of synchronous oscillations in the motor system, which is consistent with experimental observations to date.

Animals↗

The role of action in postural preparation for loading and unloading in standing subjects.

The main purpose of the present study has been to find an answer to the question: Can the subject generate anticipatory postural adjustments (APAs) when a predictable postural perturbation occurs in the absence of a voluntary action? Answering this question would allow us to distinguish between two competing hypotheses on the relation between APAs and voluntary movements. One hypothesis considers both APAsigma and voluntary "focal" movements different peripheral patterns associated with a single control process, while the alternative hypothesis considers them outcomes of two parallel control processes. Healthy subjects performed series of loading and unloading trials that included: (1) catching a falling load onto another load held in extended hands; (2) catching a falling load onto a tray attached to the trunk; (3) allowing a falling load to hit another load out of the extended hands, causing an unloading; and (4) releasing a load held in extended hands by a voluntary shoulder movement. In series 1, precautions were taken to avoid possible small hand movements prior to the impact of the falling load. Available visual information on the trajectory of the falling load was manipulated. In all conditions, except when the subject's eyes were closed, APAs were seen with patterns that were adequate for counteracting expected perturbations. Quantitative electromyographic indices of APAs depended on the availability of visual information and particular methods of introducing postural perturbations despite the fact that the magnitude of the perturbation was always the same. Our findings support a hypothesis that control processes resulting in APAs can be different from control processes associated with focal voluntary movements.

Adult↗

Proprioceptive regulation of interlimb behavior: interference between passive movement and active coordination dynamics.

The coordination of homolateral effectors (right arm/right leg) according to the in-phase or anti-phase mode was perturbed through passive movement of a third segment (left arm or left leg) imposed by the experimenter. The manipulated parameters of the passive segment were frequency and amplitude along with their degree of scaling. Results showed that passive movement degraded anti-phase patterns more than in-phase patterns. Furthermore, the anti-phase mode deteriorated profoundly during frequency manipulation, but scaling did not induce additional effects, whereas a linear association was observed between anti-phase deterioration and amplitude manipulation. Together, these data indicate that passive movement disturbed the coordination dynamics of an actively performed task. The fact that interference depended on the manipulated parameter suggests a distinction in the degree of intrusiveness of the irrelevant afferent information induced by the passive limb. It is concluded that sensory discrimination between irrelevant and relevant input is critical in performing a coordinated task adequately under perturbed conditions.

Adult↗