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In this paper, we analyze the relationship between head and chest movements and gaze direction in both walking and non-walking conditions. In a different approach from existing studies, we aim to analyze behavior when humans intentionally gaze at a certain target from two perspectives: (1) the relationship between gaze and body movements and (2) the effects of walking on body motion. We performed three experiments: fixed target scenes (Experiment 1), moving target scenes (Experiment 2) and more realistic gazing scenes (Experiment 3). The experimental results showed a linear relationship between the head and chest directions and gaze directions regardless of walking, non-walking situations, or target movements, and stronger gaze-head correlations than gaze-chest correlations. Further, we found effects of walking that constrained rotational body movements, and that body parts with larger moments were easily affected by walking. These results suggest that the findings of existing studies in non-walking situations may be applicable to walking situations directly or with simple modifications.

Shifting the direction of the line of sight in everyday life often involves rotations not only of the eyes and head but also of the trunk. Here, we investigated covariation patterns of eye-in-orbit, head-on-trunk and trunk-in-space angular horizontal displacements during whole-body rotations to targets of up to 180 degrees eccentricity performed by standing healthy human subjects. The spatial covariation was quantified statistically across various behavioral task conditions (unpredictable, memory driven predictable, visual feedback) and constraints (accuracy) by principal components (PC) analysis. Overall, the combined movement was stereotyped such that the first two PCs accounted for essentially the whole data variance of combined gaze transfers up to about 400 ms, suggesting that the three mechanical degrees of freedom under consideration are reduced to two kinematic degrees of freedom. Moreover, quantification of segment velocity variability across repetitions showed that velocities of eye-in-space and head-in-space (i.e. 'end-point' velocity) were less variable than those of the elemental variables composing them. In contrast, three statistically significant PCs accounted for the covariation of the three segments during presumably vestibularly mediated nystagmic transfers, suggesting control by a separate driving circuit. We conclude that progression of the line of sight is initially stereotypic and fulfills criteria defining a motor synergy.

Precision of limb control is associated with increased joint stiffness caused by antagonistic co-activation. The aim of this study was to examine whether this strategy also applies to precision of trunk postural control. To this end, thirteen subjects performed static postural tasks, aiming at a target object with a cursor that responded to 2D trunk angles. By manipulating target dimensions, different levels of precision were imposed in the frontal and sagittal planes. Trunk angle and electromyography (EMG) of abdominal and back muscles were recorded. Repeated measures ANOVAs revealed significant effects of target dimensions on kinematic variability in both movement planes. Specifically, standard deviation (SD) of trunk angle decreased significantly when target size in the same direction decreased, regardless of the precision demands in the other direction. Thus, precision control of trunk posture was directionally specific. However, no consistent effect of precision demands was found on trunk muscle activity, when averaged over time series. Therefore, it was concluded that stiffness regulation by antagonistic co-activation was not used to meet increased precision demands in trunk postural control. Instead, results from additional analyses suggest that precision of trunk angle was controlled in a feedback mode.

The described experiment investigated whether adaptation to successive Achilles tendon vibration depends on the postural challenge. In phase 1, thirty-six participants were divided into three groups, each with a different postural challenge. After being blindfolded, participants received 15 trials of Achilles tendon vibration (10 s-90 Hz) while standing upright in either a free-standing condition (FS group), a light finger touch condition (LFT group), or a restrained-standing condition (RS group) in which a dorsal stand prevented body displacement. In phase 2, all participants were freely standing and received 15 additional trials of vibration. Antero-posterior trunk tilt was measured at the level of the trunk (C7). Although there were limited postural readjustments in LFT, adaptation was observed in phase 1 in both FS and LFT groups. In phase 2, the evoked postural displacement was greater in the RS group than in the other two groups. We concluded that although the absence of any postural challenge concomitant to the sensory stimulation prevented adaptation to occur, a minimal challenge was sufficient.

Stabilising shifts of the centre of mass (COM) are observed during balance recovery when subjects simultaneously execute voluntary unilateral knee flexion or unilateral arm raising. Here, we examined whether voluntary lateral trunk bending provided more beneficial stabilising effects, and how motor programs of balance corrections are combined with those of the focal voluntary action. The upright balance of 24 healthy young subjects (19-33 years of age) was perturbed using multi-directional rotations of the support-surface. The perturbations consisted of combined pitch and roll rotations (7.5 degrees and 60 degrees/s) presented randomly in six different directions. Three conditions were tested: perturbation of stance only (PO); combined balance perturbation and cued uphill bending of the trunk (CONT); and combined perturbation and cued downhill bending of the trunk (IPS). For comparison, subjects were required to perform trunk bending alone (TO). Outcome measures were biomechanical responses and surface EMG activity of several muscles. Calculated predicted outcomes (PO + TO) were compared with combined measures (CONT or IPS). CONT trunk bending uphill showed two phases of benefit in balance recovery for laterally but, in contrast to voluntary knee bending, not for posterior directed components of the perturbations. IPS trunk bending had negative effects on balance. Early balance correcting muscle responses were marginally greater than PO responses. Prominent secondary balance correcting responses, having a similar timing as voluntary responses observed under TO conditions, were seen under CONT only in trunk muscles. These, and later stabilising, responses had amplitudes as expected from PO + TO conditions being significantly greater than PO responses. The ability with which different muscle synergies for balance corrections and voluntary trunk bending were integrated into one indicates a flexible adjustment of the CNS programs to the demands of both tasks.

The aim of this study was to determine whether the timing of the muscular synergies was influenced by the reduction of the base of support when we initiate a whole body reaching movement. To answer this question, we performed a principal component analysis on electromyographic activities of 24 muscles recorded on the leg, the trunk, and the arm. Our results demonstrated that during the initiation of the whole body pointing movement, only three principal components accounted for at least 95% of the variance for the overall muscular data, both when the equilibrium constraints were normal and when the base of support was reduced. These principal components were strongly correlated despite the fact that the center of mass forward displacement and the center of pressure backward displacements significantly decreased when the base of support was reduced. It suggests that the central nervous system did not change the overall timing of the muscular synergies when new equilibrium constraints were introduced in the task but was rather able to tune their amplitude as evidenced by the modification of the center of mass and center of pressure displacements.

Does the central nervous system (CNS) independently control roll and pitch movements of the human body during balance corrections? To help provide an answer to this question, we perturbed the balance of 16 young healthy subjects using multi-directional rotations of the support surface. All rotations had pitch and roll components, for which either the roll (DR) or the pitch (DP) component were delayed by 150 ms or not at all (ND). The outcome measures were the biomechanical responses of the body and surface EMG activity of several muscles. Across all perturbation directions, DR caused equally delayed shifts (150 ms) in peak lateral centre of mass (COM) velocity. Across directions, DP did not cause equally delayed shifts in anterior-posterior COM velocity. After 300 ms however, the vector direction of COM velocity was similar to the ND directions. Trunk, arm and knee joint rotations followed this roll compared to pitch pattern, but were different from ND rotation synergies after 300 ms, suggesting an intersegmental compensation for the delay effects. Balance correcting responses of muscles demonstrated both roll and pitch directed components regardless of axial alignment. We categorised muscles into three groups: pitch oriented, roll oriented and mixed based on their responses to DR and DP. Lower leg muscles were pitch oriented, trunk muscles were roll oriented, and knee and arm muscles were mixed. The results of this study suggest that roll, but not pitch components, of balance correcting movement strategies and muscle synergies are separately programmed by the CNS. Reliance on differentially activated arm and knee muscles to correct roll perturbations reveals a dependence of the pitch response on that of roll, possibly due to biomechanical constraints, and accounts for the failure of DP to be transmitted equally in time across all limbs segments. Thus it appears the CNS preferentially programs the roll response of the body and then adjusts the pitch response accordingly.

As individuals stand or walk in an environment their gaze may be reoriented from one location to another in response to auditory or visual stimuli. In order to reorient gaze, the eyes and/or the head and trunk must rotate. However, what determines the exact degree of rotation of each segment while standing or walking is not fully understood. In the current study we show that when participants were asked to reorient their gaze towards light cues positioned at eccentric locations of up to 90 degrees while standing or walking on a treadmill their eyes and head mainly facilitated the action. Rotations of the head-in-space were similar for both tasks, but the rotation of the shoulders- and hips-in-space were lower for the treadmill walking condition. It is argued that this difference in the level of head-on-trunk rotation during the two tasks is controlled by the vestibular feedback loop. The regulation of this feedback loop is performed by the cerebellum in response to the level of threat to postural stability.

Dynamic stability can be threatened by various travel surface changes that humans encounter on a daily basis. The central nervous system (CNS) must acquire appropriate information about upcoming surface changes and provide necessary proactive and reactive changes to maintain stability. The purpose of this study was to examine stability control by characterizing adaptations in step patterns, center of mass (COM) trajectory, and lower limb muscle activity when stepping onto and walking on a compliant surface. Eight young adults walked under two conditions: baseline ground walking and while walking on a large foam mat (compliant surface). Optotrak system was used to collect 3D-full body kinematics and electromyography was collected for the rectus femoris, biceps femoris, tibialis anterior, medial gastrocnemius, and soleus bilaterally. Vertical COM decreased on the compliant surface while medio-lateral COM was not affected. This lowering of the vertical COM peak would provide a more stable posture when walking on the surface. Toe trajectory during the swing phase was elevated to avoid tripping on the deformable compliant surface. Step width and length increased on the compliant surface which would increase base of support and provide better control of COM. Increases in gastrocnemius and soleus activity during push-off accounted for increases in step length seen on the compliant surface. Dynamic stability margin in the anterior-posterior direction demonstrated a constant overcompensation and subsequent correction in COM control. These proactive and reactive changes in motor patterns show how the CNS actively coordinates all body segments while traveling on a compliant surface in order to maximize stability.

The present study focused on the influence of neck vibration on the perception of the head and trunk midline position (orientation and localization). The orientation of the head and trunk was investigated by the rolling adjustment of a rod on their midline while their localization was investigated by the adjustment of the position of a visual dot as being straight-ahead the eyes or the sternum. The first experiment investigated whether a head-trunk dissociation was induced by the unilateral vibration of neck muscles in upright and restrained subjects. Results showed that the subjective orientation and localization of whole-body midline were shifted toward the vibrated side. The second experiment determined the effect of the neck muscles vibration when the subjects were lying on their side. The effect of vibration disappeared when the side of vibration was opposed to the side of postural inclination and it was stronger than in the upright position when the side of vibration and the side of postural inclination were congruent. Whereas, results suggested that the input from neck muscle proprioceptors participates directly to the elaboration of the egocentric space, the question may be raised as to how the sensory cues interacted in their contribution to the neural generation of an egocentric, body centred coordinate system.

The properties of adaptation within the locomotor and balance control systems directed towards improving one's recovery strategy for fall prevention are not well understood. The purpose of this study was to examine adaptive control of gait stability to repeated slip exposure leading to a reduction in backward loss of balance (and hence in protective stepping). Fourteen young subjects experienced a block of slips during walking. Pre- and post-slip onset stability for all slip trials was obtained as the shortest distance at touchdown (slipping limb) and lift-off (contralateral limb), respectively, between the measured center of mass (COM) state, that is, position and velocity relative to base of support (BOS) and the mathematically predicted threshold for backward loss of balance. An improvement in pre- and post-slip onset stability correlated with a decrease in the incidence of balance loss from 100% (first slip) to 0% (fifth slip). While improvements in pre-slip stability were affected by a proactive anterior shift in COM position, the significantly greater post-slip onset improvements resulted from reductions in BOS perturbation intensity. Such reactive changes in BOS perturbation intensity resulted from a reduction in the demand on post-slip onset braking impulse, which was nonetheless influenced by the proactive adjustments in posture and gait pattern (e.g., the COM position, step length, flat foot landing and increased knee flexion) prior to slip onset. These findings were indicative of the maturing process of the adaptive control. This was characterized by a shift from a reliance on feedback control for postural correction to being influenced by feedforward control, which improved pre-slip stability and altered perturbation intensity, leading to skateover or walkover (>0.05 m or <0.05 m displacement, respectively) adaptive strategies. Finally, the stability at contralateral limb lift-off was highly predictive of balance loss occurrence and its subsequent rapid reduction, supporting the notion of the internal representations of stability limits that could be modified and updated, as a key component in the adaptive control.

Understanding the physiological and psychological factors that contribute to healthy and pathological balance control in man has been made difficult by the confounding effects of the perturbations used to test balance reactions. The present study examined how postural responses were influenced by the acceleration-deceleration interval of an unexpected horizontal translation. Twelve adult males maintained balance during unexpected forward and backward surface translations with two different acceleration-deceleration intervals and presentation orders (serial or random). "SHORT" perturbations consisted of an initial acceleration (peak acceleration 1.3 m s(-2); duration 300 ms) followed 100 ms later by a deceleration. "LONG" perturbations had the same acceleration as SHORT perturbations, followed by a 2-s interval of constant velocity before deceleration. Surface and intra-muscular electromyography (EMG) from the leg, trunk, and shoulder muscles were recorded along with motion and force plate data. LONG perturbations induced larger trunk displacements compared to SHORT perturbations when presented randomly and larger EMG responses in proximal and distal muscles during later (500-800 ms) response intervals. During SHORT perturbations, activity in some antagonist muscles was found to be associated with deceleration and not the initial acceleration of the support surface. When predictable, SHORT perturbations facilitated the use of anticipatory mechanisms to attenuate early (100-400 ms) EMG response amplitudes, ankle torque change and trunk displacement. In contrast, LONG perturbations, without an early deceleration effect, did not facilitate anticipatory changes when presented in a predictable order. Therefore, perturbations with a short acceleration-deceleration interval can influence triggered postural responses through reactive effects and, when predictable with repeated exposure, through anticipatory mechanisms.

We have used the phenomenon that speed increases with movement amplitude as a rehabilitation strategy. We tested the hypothesis that the generalized training of amplitude in the limb motor system may reduce bradykinesia and hypokinesia in the upper and lower limbs in subjects with Parkinson's disease (PD) across disease severity (Stage I, n=6; Stage II, n=7; Stage III, n=5). While studies have separately examined the relationship of amplitude to speed in reaching and gait, the same study has not reported the relationship for both limb systems. Moreover, the rehabilitation intervention, Training BIG, is unique in that it applies well-established treatment concepts from a proven treatment for the speech motor system in PD [Lee Silverman Voice Treatment (LSVT)] to the limb motor system. Subjects (n=18) participated in intense practice (1-h sessions/4x week/4 weeks) of large amplitude movements involving the whole body (i.e., head, arm, trunk, and leg) while focusing on the sensory awareness of "movement bigness." Testing procedures were designed to demonstrate the transfer of generalized amplitude practice to speed improvements during functional "untrained" tasks in "uncued" conditions with blinded testers. After therapy, the subjects significantly increased their speed of reaching and gait for the preferred speed condition. This effect was greater when the severity of the disease was less. The results support further application and efficacy studies of Training BIG. Amplitude-based behavioral intervention in people with PD appears to be a simple target that may be applied in different contexts for multiple tasks and results in improved speed-amplitude scaling relations across the upper and lower limbs.

Tripping during gait occurs frequently. A successful balance recovery implies that the forward body rotation is sufficiently reduced. In view of this, adequate control of the trunk momentum is important, as the trunk has a high inertia. The aim of this study was to establish out-of-plane trunk movements after a trip and to determine trunk muscle responses. Ten male volunteers repeatedly walked over a platform in which 21 obstacles were hidden. Each subject was tripped over one of these obstacles at mid-swing of the left foot in at least five trials. Kinematics, dynamics, and muscle activity of the main trunk muscles were measured. After a trip, an increase in trunk flexion was observed (peak flexion 37 degrees). In addition, considerable movements outside the sagittal plane (up to 20 degrees) occurred. Already before landing of the blocked foot, the trunk forward bending movement was reduced, while trunk torsion and lateral rotation were still increasing. Fast responses were seen in both abdominal and back muscles, indicating stiffening of the trunk. These muscle responses preceded the mechanical trunk disturbances, which implies that these responses were triggered by other mechanisms (such as afferent signals from the extremities) rather than a simple stretch reflex. A second burst of predominantly trunk muscle extensor activity was seen at landing, suggesting specific anticipation of the trunk muscles to minimize trunk movements due to landing. In conclusion, despite large movements outside the sagittal plane, it appears that trunk muscle responses to trips are aspecific and especially aimed at minimizing trunk forward bending.

In real life situations large gaze saccades may involve rotations of the trunk, as well as the eyes and head. When this happens the rotation of the head-in-space is similar whether or not the trunk is also rotating. However, the rotation of the head on the trunk (i.e. the neck movement) is very different in the two circumstances. For similar head-in-space rotations to occur, the neck and trunk movements cannot simply add independently: they must be coordinated. It is argued that this is achieved via a feedback loop in which the semi-circular canals monitor the rotation of the head-in-space, and the neck is driven by an error signal representing the difference between the intended head-in-space trajectory and the actual trajectory. This mechanism, which is essentially the same as the vestibulo-collic reflex, nulls out disturbances to the head-in-space trajectory, whether these are caused by active or passive trunk rotation.

This study examined whether the head of elderly subjects was less stable in space when the trunk was free to move than when the trunk was fixed to a linearly moving platform. Fourteen healthy elderly subjects were seated on a linear sled with their trunk either fixed to the seat or free to move. Subjects received 10 cm, 445 cm/s2 anterior-posterior ramps and 0.35-4.05 Hz sum-of-sines translations while performing a mental distraction task in the dark. Kinematics of the head and trunk were derived from an Optotrak motion analysis system and a linear accelerometer placed on the head. Electromyographic (EMG) signals were collected for neck and paraspinal muscles. Data were tested for significance with paired t-tests corrected for multiple testing and compared (Mann-Whitney U-test) with previously published data from 12 healthy young adults (Keshner 2003). Linear acceleration trajectories of the head corresponded to the direction of sled linear acceleration when the trunk was fixed and countered the direction of the sled when the trunk was free. Angular head accelerations countered the sled or the trunk when the trunk was fixed or free, respectively. Peak amplitudes of head angular acceleration in space were greater with a fixed trunk. With the trunk free, amplitudes of head linear peak acceleration, angular accelerations, and response gains exceeded those of the young adults. Muscle EMG response latencies did not vary with the timing of head acceleration onset but the neck muscles were activated more frequently in a direction consistent with a vestibulocollic or cervicocollic reflex. Differences in angular motion of the head could be explained by the biomechanical constraints of the two tasks having one freely moving mass (head) with the trunk fixed and two freely moving masses (head and trunk) with the trunk free. These data suggest that elderly subjects rely upon active trunk mechanics in order to coordinate their head and trunk motion; however, a contribution by vestibular and ascending segmental inputs cannot be completely ruled out. A less flexible trunk in the elderly could explain why they were not as successful as young adults in stabilizing their heads in space when the trunk was free to move.

Pain changes postural activation of the trunk muscles. The cause of these changes is not known but one possibility relates to the information processing requirements and the stressful nature of pain. This study investigated this possibility by evaluating electromyographic activity (EMG) of the deep and superficial trunk muscles associated with voluntary rapid arm movement. Data were collected from control trials, trials during low back pain (LBP) elicited by injection of hypertonic saline into the back muscles, trials during a non-painful attention-demanding task, and during the same task that was also stressful. Pain did not change the reaction time (RT) of the movement, had variable effects on RT of the superficial trunk muscles, but consistently increased RT of the deepest abdominal muscle. The effect of the attention-demanding task was opposite: increased RT of the movement and the superficial trunk muscles but no effect on RT of the deep trunk muscles. Thus, activation of the deep trunk muscles occurred earlier relative to the movement. When the attention-demanding task was made stressful, the RT of the movement and superficial trunk muscles was unchanged but the RT of the deep trunk muscles was increased. Thus, the temporal relationship between deep trunk muscle activation and arm movement was restored. This means that although postural activation of the deep trunk muscles is not affected when central nervous system resources are limited, it is delayed when the individual is also under stress. However, a non-painful attention-demanding task does not replicate the effect of pain on postural control of the trunk muscles even when the task is stressful.

In the present study we investigated the extent to which individuals suffering from spastic tetraparesis as a consequence of cerebral palsy tune their trunk involvement to accuracy demands in a spoon-handling task. Twenty-two participants (ten adolescents with spastic tetraparesis and 12 control participants) had to transport a spoon filled with water or sugar to a small or a large bowl that was placed within reach. Even though trunk displacement was larger in the tetraparetic participants than it was in the control participants, the effects of the imposed accuracy constraints were remarkably similar. Participants in both groups increased trunk displacement with increasing precision requirements. Furthermore, in both groups the largest trunk involvement was found in the initial and final part of the substance-transporting phase, when wrist velocity was lowest. We propose several explanations for these findings and conclude that the large trunk involvement in individuals with tetraparetic cerebral palsy should, in any case, not be regarded as a primary symptom of the disorder, but rather as an adaptive reaction to increased task demands.

This study evaluated the degree to which the disturbance to posture from respiration is compensated for in healthy normals and whether this is different in people with recurrent low back pain (LBP), and to compare the changes when respiratory demand is increased. Angular displacement of the lumbar spine and hips, and motion of the centre of pressure (COP), were recorded with high resolution and respiratory phase was recorded from ribcage motion. With subjects standing in a relaxed posture, recordings were made during quiet breathing, while breathing with increased dead-space to induce hypercapnoea, and while subjects voluntarily increased their respiration to match ribcage expansion that was induced in the hypercapnoea condition. The relationship between respiration and the movement parameters was measured from the coherence between breathing and COP and angular motion at the frequency of respiration, and from averages triggered from the respiratory data. Small angular changes in the lumbopelvic and hip angles were evident at the frequency of respiration in both groups. However, in quiet standing, the LBP subjects had a greater displacement of their COP that was associated with respiration than the control subjects. The LBP group had a trend for less hip motion. There were no changes in the movement parameters when respiratory demand increased involuntarily via hypercapnoea, but when respiration increased voluntarily, the amplitude of motion and the displacement of the COP increased in both groups. The present data suggest that the postural compensation to respiration counteracts at least part of the disturbance to posture caused by respiration and that this compensation may be less effective in people with LBP.

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.

This study evaluated the extent to which movement of the lower limbs and pelvis may compensate for the disturbance to posture that results from respiratory movement of the thorax and abdomen. Motion of the neck, pelvis, leg and centre of pressure (COP) were recorded with high resolution in conjunction with electromyographic activity (EMG) of flexor and extensor muscles of the trunk and hip. Respiration was measured from ribcage motion. Subjects breathed quietly, and with increased volume due to hypercapnoea (as a result of breathing with increased dead-space) and a voluntary increase in respiration. Additional recordings were made during apnoea. The relationship between respiration and other parameters was measured from the correlation between data in the frequency domain (i.e. coherence) and from time-locked averages triggered from respiration. In quiet standing, small angular displacements ( approximately 0.5 degrees ) of the trunk and leg were identified in raw data. Correspondingly, there were peaks in the power spectra of the angular movements and EMG. While body movement and EMG were coherent with respiration (>0.5), the coherence between respiration and COP displacement was low (<0.2). The amplitude of movement and coherence was increased when respiration was increased. The present data suggest that the postural disturbance that results from respiratory movement is matched, at least partly, and counteracted by small angular displacements of the lower trunk and lower limbs. Thus, stability in quiet stance is dependent on movement of multiple body segments and control of equilibrium cannot be reduced to control of a single joint.

The postural response to translation of the support surface may be influenced by the performance of an ongoing voluntary task. This study was designed to test this proposal by applying lateral perturbations while subjects handled a load in the frontal plane. Measurements were made of medio-lateral displacement of the centre of pressure, angular displacement of the trunk and thigh in the frontal plane and intra-abdominal pressure. Subjects were translated randomly to the left and right in a variety of conditions that involved standing either quietly or with a 5 kg load in their left hand, which they were required either to hold statically or to lift or lower. The results indicate that when the perturbation occurred towards the loaded left side the subjects were able to return their centre of pressure, trunk and thigh rapidly and accurately to the initial position. However, when the perturbation occurred towards the right (away from the load) this correction was delayed and associated with multiple changes in direction of movement, suggesting decreased efficiency of the postural response. This reduced efficiency can be explained by a conflict between the motor commands for the ongoing voluntary task and the postural response, and/or by the mechanical effect of the asymmetrical addition of load to the trunk.

In this study, we examined flexibility in postural coordination by inducing transitions between postural patterns. Previous work demonstrated that the postural control system produces two task-specific postural patterns as a function of the frequency of support surface translation. For slow translation frequencies (<0.5 Hz), subjects ride on the platform reminiscent of upright stance (ride pattern), and for fast frequencies (> or =0.75 Hz) subjects actively fixed the head and trunk in space (head fixed pattern) during anterior-posterior platform motion. To study the adaptation of the postural control system, we had subjects stand on a support surface undergoing increases (from 0.2 to 1.0 Hz in 0.1-Hz steps) and decreases (from 1.0 to 0.2 Hz in 0.1-Hz steps) in translation frequency with the eyes open and closed. Kinematic measures of sagittal plane body motion revealed a gradual transition between these two postural patterns as a function of frequency scaling. In both the increasing and decreasing frequency conditions with visual input, center of mass displacements gradually decreased and increased, respectively, whereas upper-trunk (and head) displacement decreased gradually within the ride pattern until a head fixed pattern was observed without any significant changes in displacement for translation frequencies at and above 0.6 Hz. Without visual input, the scaling of the ride pattern was similar except the transition to the head fixed pattern never emerged with increasing frequency; instead, a less stable pattern exhibiting slow drift in head-trunk anterior-posterior motion (drift pattern) was observed at and above 0.5 Hz oscillations. The stability of the head fixed pattern at fast frequencies was clearly dependent on visual input suggesting that vision was more critical for trunk and head control in space at high than low translation frequencies. Head velocity was kept constant, and lower with vision, as translation frequency (and velocity) changed suggesting a head velocity threshold constraint across postural patterns. The gradual transition from the ride to the head fixed pattern was made possible by the recruitment of available degrees of freedom in the form of ankle, then knee, and then hip joint motion. In turn, the transition from the head fixed or drift pattern was made possible by the gradual suppression of available degrees of freedom in the form of reducing hip, then knee, and then ankle motion. The gradual change in postural kinematics without instabilities and hysteresis suggests that the ability to recruit and suppress biomechanical degrees of freedom allows the postural control system to gradually change postural strategies without suffering a loss of stability. The results are discussed in light of possible self-organizing mechanisms in the multisensory control of posture.

Movements of different body segments may be combined in different ways to achieve the same motor goal. How this is accomplished by the nervous system was investigated by having subjects make fast pointing movements with the arm in combination with a forward bending of the trunk that was unexpectedly blocked in some trials. Subjects moved their hand above the surface of a table without vision from an initial position near the midline of the chest to remembered targets placed within the reach of the arm in either the ipsi- or contralateral workspace. In experiment 1, subjects were instructed to make fast arm movements to the target without corrections whether or not the trunk was arrested. Only minor changes were found in the hand trajectory and velocity profile in response to the trunk arrest, and these changes were seen only late in the movement. In contrast, the patterns of the interjoint coordination substantially changed in response to the trunk arrest, suggesting the presence of compensatory arm-trunk coordination minimizing the deflections from the hand trajectory regardless of whether the trunk is recruited or mechanically blocked. Changes in the arm interjoint coordination in response to the trunk arrest could be detected kinematically at a minimal latency of 50 ms. This finding suggests a rapid reflex compensatory mechanism driven by vestibular and/or proprioceptive afferent signals. In experiment 2, subjects were required, as soon as they perceived the trunk arrest, to change the hand motion to the same direction as that of the trunk. Under this instruction, subjects were able to initiate corrections only after the hand approached or reached the final position. Thus, centrally mediated compensatory corrections triggered in response to the trunk arrest were likely to occur too late to maintain the observed invariant hand trajectory in experiment 1. In experiment 3, subjects produced similar pointing movements, but to a target that moved together with the trunk. In these body-oriented pointing movements, the hand trajectories from trials in which the trunk was moving or arrested were substantially different. The same trajectories represented in a relative frame of reference moving with the trunk were virtually identical. We conclude that hand trajectory invariance can be produced in an external spatial (experiment 1) or an internal trunk-centered (experiment 3) frame of reference. The invariance in the external frame of reference is accomplished by active compensatory changes in the arm joint angles nullifying the influence of the trunk motion on the hand trajectory. We suggest that to make a transition to the internal frame of reference, control systems suppress this compensation. One of the hypotheses opened to further experimental testing is that the integration of additional (trunk) degrees of freedom into movement is based on afferent (proprioceptive, vestibular) signals stemming from the trunk motion and transmitted to the arm muscles.

We investigated whether a tap with a reflex hammer to the forehead can elicit responses in the leg muscles and whether vestibular stimulation is the crucial prerequisite for eliciting these responses. We also measured the postural changes caused by the tap and by the compensatory, presumably reflex-like reactions of the subject. Tap-evoked activity of leg muscles was easily elicited during upright stance in normal subjects and was also seen in two subjects without vestibular function. The pattern of muscle activation clearly showed a counteraction to the tap-evoked perturbation of stance. Taps applied to the chest elicited similar reflexes. Since these two conditions imply a different activation of the vestibular apparatus, the vestibular input alone cannot account for the observed leg muscle reflexes. We suggest that multisensory reflex pathways that integrate vestibular and proprioceptive inputs account for these reflexes.

The purpose of this study is to analyze the coordination patterns of the elevation angles of lower limb segments following the onset of unsupported walking in children and to look for the existence of a planar covariation rule as previously described in adult human locomotion. The kinematic patterns of locomotion were recorded in 21 children (11-144 months of age) and 19 adults. In 4 children we monitored the very first unsupported steps. The extent to which the covariation of thigh, shank, and foot angles was constrained on a plane in 3D space was assessed by means of orthogonal regression and statistically quantified by means of principal component analysis. The orientation of the covariation plane of the children was compared with the mean value of the adults' plane. Trunk stability with respect to the vertical was assessed in both the frontal (roll) and sagittal (pitch) planes. The evolution with walking experience of the plane orientation and trunk oscillations demonstrated biexponential profiles with a relatively fast time constant (< 6 months after the onset of unsupported locomotion) followed by a much slower progression toward adult values. The initial fast changes of these walking parameters did not parallel the slow, monotonic maturation of anthropometric parameters. The early emergence of the covariation plane orientation and its correlation with trunk vertical stability reflect the dynamic integration of postural equilibrium and forward propulsion in a gravity-centered frame. The results support the view that the planar covariation reflects a coordinated, centrally controlled behavior, in addition to biomechanical constraints. The refinement of the planar covariation while morphological variables drastically change as the child grows implies a continuous update of the neural command.

Trunk and head movements were characterized over a wide range of walking speeds to determine the relationship between stride length, stepping frequency, vertical head translation, pitch rotation of the head, and pitch trunk rotation as a function of gait velocity. Subjects (26-44 years old) walked on a linear treadmill at velocities of 0.6-2.2 m/s. The head and trunk were modeled as rigid bodies, and rotation and translation were determined using a video-based motion analysis system. At walking speeds up to 1.2 m/s there was little head pitch movement in space, and the head pitch relative to the trunk was compensatory for trunk pitch. As walking velocity increased, trunk pitch remained approximately invariant, but a significant head translation developed. This head translation induced compensatory head pitch in space, which tended to point the head at a fixed point in front of the subject that remained approximately invariant with regard to walking speed. The predominant frequency of head translation and rotation was restricted to a narrow range from 1.4 Hz at 0.6 m/s to 2.5 Hz at 2.2 m/s. Within the range of 0.8-1.8 m/s, subjects tended to increase their stride length rather than step frequency to walk faster, maintaining the predominant frequency of head movement at close to 2.0 Hz. At walking speeds above 1.2 m/s, head pitch in space was highly coherent with, and compensatory for, vertical head translation. In the range 1.2-1.8 m/s, the power spectrum of vertical head translation was the most highly tuned, and the relationship between walking speed and head and trunk movements was the most linear. We define this as an optimal range of walking velocity with regard to head-trunk coordination. The coordination of head and trunk movement was less coherent at walking velocities below 1.2 m/s and above 1.8 m/s. These results suggest that two mechanisms are utilized to maintain a stable head fixation distance over the optimal range of walking velocities. The relative contribution of each mechanism to head orientation depends on the frequency of head movement and consequently on walking velocity. From consideration of the frequency characteristics of the compensatory head pitch, we infer that compensatory head pitch movements may be produced predominantly by the angular vestibulocollic reflex (aVCR) at low walking speeds and by the linear vestibulocollic reflex (1VCR) at the higher speeds.

The major purpose of this research was to determine whether kinesthetic/proprioceptive perceptions of the earth-fixed vertical axis are more accurate than perceptions of intrinsic axes. In one experiment, accuracy of alignment of the forearm to earth-fixed vertical and head- and trunk-longitudinal axes by seven blindfolded subjects was compared in four tasks: (1) Earth-Arm--arm (humerus) orientation was manipulated by the experimenter; subjects aligned the forearm parallel to the vertical axis, which was also aligned with the head and trunk longitudinal axis; (2) Head--head, trunk, and upper-limb orientations were manipulated by the experimenter, subjects aligned the forearm parallel to the longitudinal axis of the head using only elbow flexion/extension and shoulder internal/external rotation; (3) Trunk--same as (2), except that subjects aligned the forearm parallel to the trunk-longitudinal axis; (4) Earth--same as (2), except that subjects aligned the forearm parallel to the earth-fixed vertical. Head, trunk, and gravitational axes were never parallel in tasks 2, 3, and 4 so that subjects could not simultaneously match their forearm to all three axes. The results showed that the errors for alignment of the forearm with the earth-fixed vertical were lower than for the trunk- and head-longitudinal axes. Furthermore, errors in the Earth condition were less dependent on alterations of the head and trunk orientation than in the Head and Trunk conditions. These data strongly suggest that the earth-fixed vertical is used as one axis for the kinesthetic sensory coordinate system that specifies upper-limb orientation at the perceptual level. We also examined the effects of varying gravitational torques at the elbow and shoulder on the accuracy of forearm alignment to earth-fixed axes. Adding a 450 g load to the forearm to increase gravitational torques when the forearm is not vertical did not improve the accuracy of forearm alignment with the vertical. Furthermore, adding small, variably sized loads (between which the subjects could not distinguish at the perceptual level) to the forearm just proximal to the wrist produced similar errors in aligning the forearm with the vertical and horizontal. Forearm-positioning errors were not correlated with the size of the load, as would be expected if gravitational torques affected forearm-position sense. We conclude that gravitational torques exerted about the shoulder and elbow do not make significant contributions to sensing forearm-orientation relative to earth-fixed axes when the upper-limb segments are not constrained by external supports.

To understand the internal representations used by the nervous system to coordinate multijoint movements, we examined the coordination among the body segments during reach-to-grasp movements which involve grasping by the hand and reaching by the arm and trunk. Subjects were asked to reach and grasp an object using the arm only, the trunk only, and some combinations of both arm and trunk. Results showed that kinematic parameters related to the transport component of the arm and the trunk, such as peak velocity and time to peak velocity, varied across conditions and that the coordination pattern between the arm and trunk was different across conditions. However, parameters related to the grasp component, such as peak aperture, time to peak aperture, and closing distance, were invariant, regardless of whether the hand was delivered to the target by the arm only, the trunk only, or both. We hypothesize that a hierarchy of motor control processes exists, in which the reach and grasp components are governed by independent neuromotor synergies, which in turn are coordinated temporally and spatially by a higher-level synergy.

The contribution of cervical and vestibular cues in signaling the changes in target-trunk relative positions during self-motion was investigated. Normal subjects (Ss) were shown a LED flashed in the peripheral visual field in a dark room. Ss were then passively rotated about the vertical axis in one of three different conditions: (1) head chair-fixed (vestibular condition); (2) head earth-fixed (relaxed neck condition); and (3) head earth-fixed, but with the Ss actively attempting to turn it (activated neck condition). The Ss were then required to indicate, with their unseen index finger, the position of the previously flashed target. It was found that pointing at the memorized target was similarly accurate in the relaxed neck condition and in the activated neck condition. In the vestibular condition, pointing accuracy dropped significantly. These results suggest that neck proprioceptive signals are more effective than vestibular ones in signaling relative changes in the position of stationary objects with respect to the body during head-trunk motion. The finding that cervically mediated estimates were unchanged during active contraction of the neck muscles may suggests that efference copy signals may help interpret the change in the afferent signals caused by voluntary neck muscle activation.

Upper trunk bending movements were accompanied by opposite movements of the lower body segments. These axial kinematic synergies maintained equilibrium during the movement performance by stabilizing the center of gravity (CG), which shifted on average across all the subjects by 1 +/- 4 cm in the anteroposterior direction and thus always remained within the support area. The aim of the present investigation was to provide an insight into the central control responsible for the performance of these synergies. The kinematic analysis was performed by the method of principal components (PC) analysis applied to the covariation between ankle, knee and hip joint angles and compared with CG shifts during upper trunk bending. Subjects were asked to perform backward or forward upper trunk bending in response to a tone. They were instructed to move as fast as possible or slowly (2 s), with high or low movement amplitudes. PC analysis showed a strong correlation between hip, knee and ankle joint changes. The first principal component (PC1) representing a multijoint movement with fixed ratios between joint angular changes, accounted, on average, for 99.7% +/- 0.2% of the total angular variance in the forward trunk movements and for 98.4% +/- 1.4% in the backward movements. The instructed voluntary regulation of the amplitude and velocity of the movement was achieved by adapting the bell-shaped profile of the velocity time course without changes in interjoint angular relations. Fixed ratios between changes in joint angles, represented by PC1, ensured localization of the CG within the support area during trunk bending. The ratios given by PC1 showed highly significant dependence on subjects, suggesting the adaptability of the central control to each subject's biomechanical peculiarities. Subject's intertrial variability of PC1 ratios was small, suggesting a stereotyped automatic interjoint coordination. When changing velocity and amplitude of the movement, the ratios remained the same in about half the subjects while in others slight variations were observed. A weak second principal component (PC2) was shown only for fast movements. In forward movements PC2 reflected the early knee flexion that seems related to the disturbances caused by the passive interaction between body segments, rather than to the effect of a central command. In fast backward movements, PC2 reflected the delay in hip extension relative to the movement onset in the ankle and knee that mirrors intersubject differences in the initiation process of the axial synergy. The results suggest that PC1 reflects the centrally controlled multijoint movement, defining the time course and amplitude of the movement and fixing the ratios between changes in joint angles. They support the hypothesis that the axial kinematic synergies result from a central automatic control that stabilizes the CG shift in the anteroposterior direction while performing the upper trunk bending.

Neuronal activity was recorded from the superior colliculus (SC) and the underlying reticular formation in two monkeys during an arm reaching task. Of 744 neurons recorded, 389 (52%) clearly modulated their activity with arm movements. The temporal activity patterns of arm-movement-related neurons often had a time course similar to rectified electromyograms (EMGs) of particular muscles recorded from the shoulder, arm or trunk. These reach cells, as well as the muscles investigated, commonly exhibited mono- or biphasic (less frequently tri- or polyphasic) excitatory bursts of activity, which were related to the (pre-)movement period, the contact phase and/or the return movement. The vast majority of reach cells exhibited a consistent activity pattern from trial to trial as did most of the muscles of the shoulder, arm and trunk. Similarities between the activity patterns of the neurons and the muscles were sometimes very strong and were especially notable with the muscles of the shoulder girdle (e.g. trapezius descendens, supraspinatus, infraspinatus or the anterior and medial deltoids). This high degree of co-activation suggests a functional linkage, though not direct, between the collicular reach cells and these muscles. Neuronal activity onset was compared with that of 25 muscles of the arms, shoulders and trunk. The majority of cells (78.5%) started before movement onset with a mean lead time of 149+/-90 ms, and 36.5% were active even before the earliest EMG onset. The neurons exhibited the same high degree of correlation (r=0.97, Spearman rank) between activity onset and the beginning of the arm movement as did the muscles (r=0.98) involved in the task. The mean neuronal reach activity (background subtracted) ranged between 7 and 193 impulses/s (mean 40.5+/-24.2). The mean modulation index calculated [(reach activity background activity)/reach activity+background activity)] was 0.75+/-0.23 for neurons (n=358) and 0.87+/-0.14 for muscles (n=25). As the monkeys fixated the reach target constantly during an arm movement, neuronal activity which was modulated in this period was not related to eye movements. The three neck muscles investigated in the reach task exhibited no reach-related activity modulation comparable to that of either the reach cells or the muscles of the shoulder, arm and trunk. However, tonic neck muscle EMG was monotonically related to horizontal eye position. The clear skeletomotor discharge characteristics of arm-movement-related SC neurons revealed in this study agree with those already known from other sensorimotor regions (for example the primary motor, the premotor and parietal cortex, the basal ganglia or the cerebellum) and are consistent with the possible role of this population of reach cells in the control of arm movements.

There are several controversies concerning the organization and induction of postural adjustments in standing humans. Some investigators suggest the responses are triggered by somatosensory inputs (especially from the ankle in standing subjects), while others emphasize the vestibular input induced by head acceleration. We examined postural responses in sitting subjects in order to describe the muscle activation pattern during various perturbations and to test whether somatosensory or vestibular stimulation elicited the responses. The kinematics and EMG patterns in response to perturbations caused by movements of the support surface were studied in adults. The postural muscle activation following a backward sway was mainly the same, whether it was elicited by a forward translation or a legs-up rotation. This is remarkable, since, except for pelvis rotation, the movements of all body segments including the head differed in the two conditions. Furthermore, a second experiment showed that the direction of the initial head movement could be reversed with retainment of the same postural muscle activation pattern. The results suggest that somatosensory signals derived from the backward rotation of the pelvis, and not vestibular information from the head, trigger postural responses during sitting. There was a slight but consistent difference in the muscle activation pattern, whether the backward sway was elicited by a forward translation or legs-up rotation. The difference seemed to reflect the sensory information from head and other body parts (except the pelvis). This finding allowed us to speculate in a central pattern generator for postural adjustments containing two levels. At the first level, a simple format of the muscle activation would be generated; at the second level, the centrally generated pattern could be shaped and timed by interaction from the entire somatosensory, vestibular, and visual input.

This study examined whether postural responses emerge in children in a predetermined way before independent sitting is achieved, and in what respect postural responses in infants differ from those in adults. Children just able to sit independently and children not yet able to sit were exposed to surface perturbations (translation and rotation) while body movement and electromyographic (EMG) responses were recorded. Perturbations causing a backward sway of the body (i.e., forward translation and legs-up rotation), elicited consistent patterns of muscle activity in ventral hip, trunk, and neck muscles in the independently sitting children. A high tonic EMG background activity in trunk and neck extensor muscles was inhibited at the onset of the ventral muscle activity. Kinematic analysis revealed that backward rotation of the pelvis was the first detectable body movement, while head movements (linear and angular displacement) were irregular and occurred later than the pelvis movement. Perturbations in the opposite direction, causing a forward sway, evoked variable responses in dorsal trunk and neck muscles, suggesting that the excitability level for postural responses was set according to the stability limits of the body. Children not yet able to sit without support were tested when the support around the waist, given by the experimenter's hands, was released prior to the onset of the platform perturbation. Postural responses were elicited in ventral muscles following a backward sway in all children and in about 60% of all trials. Often, only some of the ventral muscles were activated. No distinct responses were evoked during perturbations imposing a forward sway. These results suggest that (1) backward rotation of the pelvis triggers the postural adjustments in the independently sitting children; (2) a basic form of the postural adjustment develops in a predetermined manner before children practice independent sitting; and (3) the basic structure of ventral muscle activation pattern resembles that of adults, while the activation of the dorsal muscles (inhibition) differs in several aspects. These findings are in agreement with a recent model of central pattern generators for postural responses consisting of two operative levels. At the first level, which is triggered by backward rotation of the pelvis, the basic activation pattern is generated. At the second level, the pattern is shaped and fine-tuned by multisensory interactions from all activated sensory systems.(ABSTRACT TRUNCATED AT 400 WORDS)

Chronic loss of vestibular function modifies the role of neck afferents in human perception of self-motion. We characterized this change by comparing the self-motion perception of patients with chronic vestibular loss (Ps) to that of normal subjects (Ns). Stimuli consisted of sinusoidal horizontal rotations (0.025-0.4 Hz) of the trunk relative to the head (neck stimulation) and/or of the head in space (vestibular stimulation). Perception of head rotation relative to the trunk, of trunk rotation in space, or of head rotation in space was assessed in terms of gain and phase (veridical perception, G = 1 and phi = 0 degree) as well as detection threshold using a pointing procedure. (1) Perception of head rotation relative to the trunk (neck proprioception). Ps' detection threshold of head-to-trunk rotation was normal (i.e. similar to that of Ns) across all frequencies tested. Also, with peak angular velocities above 5 degrees/s, the gain of their perception was approximately normal. When peak velocity was decreased below this value, however, either by lowering stimulus frequency with peak displacement kept constant (+/- 8 degrees) or by decreasing peak displacement at constant frequency (0.05 Hz), the gain increased above unity, unlike in Ns. In contrast, the phase remained normal (approximately 0 degree). (2) Perception of trunk rotation in space. Ps perceived their trunks as stationary during neck stimulation and all vestibular-neck combinations at medium to low frequencies. At 0.4 Hz, however, Ps consistently perceived the trunk rotation, conceivably due to somatosensory self-motion cues arising from high body acceleration. In contrast, Ns perceive a trunk-in-space rotation with the neck stimulation and most of the stimulus combinations across the whole frequency range tested. Ns perceived their trunks as stationary only during head rotation on the stationary trunk (presumed to reflect a mutual cancellation of neck and vestibular signals). (3) Perception of head rotation in space. In Ps, unlike Ns, this perception always resembled that of head rotation relative to the trunk. (4) When Ps were presented with a visual or somatosensory space reference (not motion cues), their perception of trunk and head rotation in space became approximately normal. (5) We suggest that there are basically two changes in the neck-induced self-motion perception associated with chronic vestibular loss. First, neck proprioception shows a non-linear gain that overemphasizes low stimulus velocities, for unknown reasons.(ABSTRACT TRUNCATED AT 400 WORDS)