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We investigated the three-dimensional spatial characteristics of caloric nystagmus during excitation and inhibition of the lateral semicircular canal in five normal human subjects. Each subject was repositioned in 45 degrees steps at 1-min intervals such that the right lateral semicircular canal plane was reoriented in pitch, from 135 degrees backwards from the upright position to 135 degrees forwards, while the right ear was continuously stimulated with air at 44 degrees C. In orientations in which caloric stimulus resulted in excitation of the right lateral semicircular canal, the eye velocity axis was orthogonal to the average orientation of the right lateral semicircular canal plane. However, in orientations in which caloric stimulus resulted in inhibition of the right lateral semicircular canal, the eye velocity axis was orthogonal to the average orientation of the left and not the right lateral semicircular canal plane. These findings suggest that velocity and direction of caloric nystagmus depend not only on the absolute magnitude of vestibular activity on the stimulated side but also on the differences in activity between the left and right vestibular nuclei, most probably mediated centrally via brainstem commissural pathways.

In cerebellar ataxia, kinematic aberrations of multijoint movements are thought to originate from deficiencies in generating muscular torques that are adequate to control the mechanical consequences of dynamic interaction forces. At this point the exact mechanisms that lead to an abnormal control of interaction torques are not known. In principle, the generation of inadequate muscular torques may result from an impairment in generating sufficient levels of torques or from an inaccurate assessment and prediction of the mechanical consequences of movements of one limb segment on adjacent joints. We sought to differentiate the relative contribution of these two mechanisms and, therefore, analyzed intersegmental dynamics of multijoint pointing movements in healthy subjects and in patients with cerebellar degeneration. Unrestrained vertical arm movements were performed at three different target movement velocities and recorded using an optoelectronic tracking system. An inverse dynamics approach was employed to compute net joint torques, muscular torques, dynamic interaction torques and gravitational torques acting at the elbow and shoulder joint. In both groups, peak dynamic interaction forces and peak muscular forces were largest during fast movements. In contrast to normal subjects, patients produced hypermetric movements when executing fast movements. Hypermetric movements were associated with smaller peak muscular torques and smaller rates of torque change at elbow and shoulder joints. The patients' deficit in generating appropriate levels of muscular force were prominent during two different phases of the pointing movement. Peak muscular forces at the elbow were reduced during the initial phase of the movement when simultaneous shoulder joint flexion generated an extensor influence upon the elbow joint. When attempting to terminate the movement, gravitational and dynamic interaction forces caused overshooting extension at the elbow joint. In normal subjects, muscular torque patterns at shoulder and elbow joint were synchronized in that peak flexor and extensor muscular torques occurred simultaneously at both joints. This temporal pattern of muscular torque generation at shoulder and elbow joint was preserved in patients. Our data suggest that an impairment in generating sufficient levels of phasic muscular torques significantly contributes to the patients' difficulties in controlling the mechanical consequences of dynamic interaction forces during multijoint movements.

Kinematic abnormalities of fast multijoint movements in cerebellar ataxia include abnormally increased curvature of hand trajectories and an increased hand path and are thought to originate from an impairment in generating appropriate levels of muscle torques to support normal coordination between shoulder and elbow joints. Such a mechanism predicts that kinematic abnormalities are pronounced when fast movements are performed and large muscular torques are required. Experimental evidence that systematically explores the effects of increasing movement velocities on movement kinematics in cerebellar multijoint movements is limited and to some extent contradictory. We, therefore, investigated angular and hand kinematics of natural multijoint pointing movements in patients with cerebellar degenerative disorders and healthy controls. Subjects performed self-paced vertical pointing movements with their right arms at three different target velocities. Limb movements were recorded in three-dimensional space using a two-camera infrared tracking system. Differences between patients and healthy subjects were most prominent when the subjects performed fast movements. Peak hand acceleration and deceleration were similar to normals during slow and moderate velocity movements but were smaller for fast movements. While altering movement velocities had little or no effect on the length of the hand path and angular motion of elbow and shoulder joints in normal subjects, the patients exhibited overshooting motions (hypermetria) of the hand and at both joints as movement velocity increased. Hypermetria at one joint always accompanied hypermetria at the neighboring joint. Peak elbow angular deceleration was markedly delayed in patients compared with normals. Other temporal movement variables such as the relative timing of shoulder and elbow joint motion onsets were normal in patients. Kinematic abnormalities of multijoint arm movements in cerebellar ataxia include hypermetria at both the elbow and the shoulder joint and, as a consequence, irregular and enlarged paths of the hand, and they are marked with fast but not with slow movements. Our findings suggest that kinematic movement abnormalities that characterize cerebellar limb ataxia are related to an impairment in scaling movement variables such as joint acceleration and deceleration normally with movement speed. Most likely, increased hand paths and decomposition of movement during slow movements, as described earlier, result from compensatory mechanisms the patients may employ if maximum movement accuracy is required.

We recorded reaching movements from nine infants longitudinally from the onset of reaching (5th postnatal month) up to the age of 3 years. Here we analyze hand and proximal joint trajectories and examine the emerging temporal coordination between arm segments. The present investigation seeks (a) to determine when infants acquire consistent, adult-like patterns of multijoint coordination within that 3-year period, and (b) to relate their hand trajectory formation to underlying patterns of proximal joint motion (shoulder, elbow). Our results show: First, most kinematic parameters do not assume adult-like levels before the age of 2 years. At this time, 75% of the trials reveal a single peaked velocity profile of the hand. Between the 2nd and 3rd year of life, "improvements" of hand- or joint-related movement units are only marginal. Second, infant motor systems strive to obtain velocity patterns with as few force reversals as possible (uni- or bimodal) at all three limb segments. Third, the formation of a consistent interjoint synergy between shoulder and elbow motion is not achieved within the 1st year of life. Stable patterns of temporal coordination across arm segments begin to emerge at 12-15 months of age and continue to develop up to the 3rd year. In summary, the development toward adult forms of multijoint coordination in goal-directed reaching requires more time than previously assumed. Although infants reliably grasp for objects within their workspace 3-4 months after the onset of reaching, stereotypic kinematic motor patterns are not expressed before the 2nd year of life.

Nine young infants were followed longitudinally from 4 to 15 months of age. They performed multi-joint reaching movements to a stationary target presented at shoulder height. Time-position data of the hand, shoulder, and elbow were collected using an optoelectronic measurement system. In addition, we recorded electromyographic activity (EMG) from arm extensors and flexors. This paper documents how control problems of proximal torque generation may account for the segmented hand paths seen during early reaching. Our analysis revealed the following results: first, muscular impulse (integral of torque) increased significantly between the ages of 20 (reaching onset) and 64 weeks. That is, as infants got older they produced higher levels of mean muscular flexor torque during reaching. Data were normalized by body weight and movement time, so differences are not explained by anthropometric changes or systematic variations in movement time. Second, while adults produced solely flexor muscle torque to accomplish the task, infants generated flexor and extensor muscle torque at shoulder and elbow throughout a reach. At reaching onset more than half of the trials revealed this latter kinetic profile. Its frequency declined systematically as infants got older. Third, we examined the pattern of muscle coordination in those trials that exhibited elbow extensor muscle torque. We found that during elbow extension coactivation of flexor and extensor muscles was the predominant pattern in 67% of the trials. This pattern was notably absent in comparable adult reaching movements. Fourth, fluctuations in force generation, as measured by the rate of change of total torque (NET) and muscular torque (MUS), were more frequent in early reaching (20-28 weeks) than in the older cohort (52-64 weeks), indicating that muscular torque production became increasingly smoother and task-efficient. Our data demonstrate that young infants have problems in generating smooth profiles of proximal joint torques. One possible reason for this imprecision in infant force control is their inexperience in predicting the magnitude and direction of external forces. That infants learned to consider external forces is documented by their increasing reliance on these forces when performing voluntary elbow extensions. The patterns of muscle coordination underlying active elbow extensions were basically the same as during the prereaching phase, indicating that the formation of functional synergies is based on a basal repertoire of innervation patterns already observable in very early, spontaneous movements.

We recently argued that the specific compensation of spatial neglect by manipulating neck proprioceptive and vestibular input is due to a central "correction" of the disturbed neural transformation process converting the afferent input coordinates from the peripheral sensory organs into a central representation of egocentric space. Both types of stimulation were proposed to induce a reorientation of the deviated or distorted egocentric spatial reference frame. The aim of the present study was to observe this process of reorientation under a condition in which no visual stimulus can attract the subject's attention and thus influence exploration behaviour from outside. We recorded eye movements of normal subjects and of three patients with spatial neglect after right parietal lesions while searching for a non-existent target in complete darkness. It was assumed that the area of the outer space that subjects spontaneously explore under this condition is a direct function of the subject's representation of egocentric space. Ocular space exploration was biased and confined almost entirely to the right side of the midsagittal plane in patients with neglect. This spatial distribution of exploratory eye movements changed remarkably with left-sided neck muscle vibration as well as with left-sided vestibular stimulation using ice water calorics. The spatial area of exploration was significantly enlarged to the contralesional side and the exploration maximum shifted in the same direction. Whereas with both types of stimulation space exploration of patients with neglect was similar to that of normal subjects when not being stimulated, neck proprioceptive and vestibular stimulation in normal subjects induced a quasi neglect-like exploration pattern, i.e. a bias to one side of the objective midsagittal plane. If ocular space exploration was, however, related to the subjectively perceived position of the midsagittal plane in space, eye movements were symmetrically distributed and carried out to both sides of subjective "straight ahead" in all experimental conditions, in normal subjects as well as in patients with neglect. The present results support the above hypothesis and indicate that neck proprioceptive as well as vestibular input directly contribute to the computation of the subject's central representation of egocentric space used for localizing body orientation and for guiding motor behaviour in space.

The otolith-semicircular canal interaction during postrotatory nystagmus was studied in ten normal human subjects by applying fast, short-lasting, passive head and body tilts (15, 30, 45, or 90 degrees in the roll or pitch plane) 2 s after sudden stop from a constant-velocity rotation (100 degrees/s) about the earth-vertical axis in yaw. Eye movements were measured with three-dimensional magnetic search coils. Following the head tilt, activity in the semicircular canal primary afferents continues to reflect the postrotatory angular velocity vector in head-centered coordinates, whereas otolith primary afferents signal a different orientation of the head relative to gravity. Despite the change in head orientation relative to gravity, postrotatory eye velocity decayed closely along the axis of semicircular canal stimulation (horizontal in head coordinates) for large head tilts (90 degrees) and also for small head tilts (15-45 degrees) for reorientations in the pitch plane. Only for small head tilts (15-45 degrees) in the roll plane was there a reorientation of the eye rotation axis toward the gravitational vector. This reorientation was approximately compensatory for 15 degrees head tilts. For 30 degrees and 45 degrees head tilts the eye rotation axis tilted toward the gravitational vector by about the same amount as for 15 degrees head tilts. These results suggest that, with the exception of small head tilts in the roll plane, there was no compelling data showing a relationship between the eye rotation axis and head tilt and that postrotatory nystagmus is largely organized in head-centered rather than gravity-centered coordinates in humans. This indicates a rudimentary, nonlinear, and direction-specific interaction of semicircular canal and otolith signals in the central vestibular system in humans.

Nine young infants were followed longitudinally from 4 to 15 months of age. We recorded early spontaneous movements and reaching movements to a stationary target. Time-position data of the hand (endpoint), shoulder, and elbow were collected using an optoelectronic measurement system (ELITE). We analyzed the endpoint kinematics and the intersegmental dynamics of the shoulder and elbow joint to investigate how changes in proximal torque control determined the development of hand trajectory formation. Two developmental phases of hand trajectory formation were identified: a first phase of rapid improvements between 16 and 24 weeks of age, the time of reaching onset for all infants. During that time period the number of movement units per reach and movement time decreased dramatically. In a second phase (28-64 weeks), a period of "fine-tuning" of the sensorimotor system, we saw slower, more gradual changes in the endpoint kinematics. The analysis of the underlying intersegmental joint torques revealed the following results: first, the range of muscular and motion-dependent torques (relative to body weight) did not change significantly with age. That is, early reaching was not confined by limitations in producing task-adequate levels of muscular torque. Second, improvements in the endpoint kinematics were not accomplished by minimizing amplitude of muscle and reactive torques. Third, the relative timing of muscular and motion-dependent torque peaks showed a systematic development toward an adult timing profile with increasing age. In conclusion, the development toward invariant characteristics of the hand trajectory is mirrored by concurrent changes in the control of joint forces. The acquisition of stable patterns of intersegmental coordination is not achieved by simply regulating force amplitude, but more so by modulating the correct timing of joint force production and by the system's use of reactive forces. Our findings support the view that development of reaching is a process of unsupervised learning with no external or innate teacher prescribing the desired kinematics or kinetics of the movement.

Effects of cerebellar lesions on the production of isometric pinch force and the coordination of pinch and lift force were examined. Twenty-one patients, mostly with degenerative cerebellar disorders, and ten healthy controls lifted an instrumented test object using the precision grip of thumb and index finger. The load of the object could be varied to study the adaptation of pinch force generation. The results were: (1) Cerebellar patients were able to adapt their pinch force levels to the different object loads. (2) Patients showed a longer latency between the onset of pinch force and onset of life force than controls. The level of pinch force at the start of lift force was elevated. (3) Patients were able to use sensorimotor memory about object load to adapt force output based on previous experience through repetitive testing, but they were significantly less efficient than healthy controls. (4) The temporal profile of pinch force rate of change featured an irregular pattern characteristic for a lack of sufficient anticipatory parameterization.

Automatic postural responses of leg muscles to the sudden displacement of standing support were investigated under four different conditions of information given to subjects in advance. Results from three groups of subjects were compared: 6 normal subjects, 10 patients with cerebellar disease, and 9 patients with Parkinson's disease. Specifically, each subject was provided with visual information about the direction and/or the amplitude of an upcoming platform tilt. For the control situation no advance information on the characteristics of platform tilt was provided. Neither the latencies nor the integrals of postural EMG-responses showed alterations with advance information. In contrast, in a control experiment in which 3 normal subjects had to perform large or small forward or backward voluntary movements of the body around the ankle joint, shorter onset-latencies of leg muscle EMG responses were observed with increasing complexity of the advance information. These results suggest that, unlike voluntary movements, postural responses to rapid surface tilts do not benefit from advance visual information on direction or amplitude of a postural disturbance.

Apparent velocities of moving visual stimuli are known to be different depending on whether the subject pursues the stimulus (efferently controlled motion perception) or whether the eye is stationary and the image moves across the retina (afferent motion perception). Afferent motion perception of a periodic pattern or a moving single object causes overestimation of velocity (magnitude estimations) as compared to smooth pursuit. This socalled Aubert-Fleischl phenomenon is shown to depend on local temporal frequency stimulation on the retina caused by the repetitive passage of contrast borders of the moving periodic pattern. This is evidenced by the fact that for a given stimulus speed the amount of overestimation is a function of the spatial frequency of the pattern (or the angular subtend of a single moving object) and that the Aubert-Fleischl phenomenon is not observed if a single edge moves. Background characteristics seem not to influence the apparent velocity during smooth pursuit.

Displacements of the center of foot pressure, the hip and the head were recorded in six subjects standing on a platform, sinusoidally tilting in pitch (anterior-posterior). Stimulus frequencies ranged between 0.01 and 1 Hz. Stimulus amplitudes were 2, 4 and 6 degrees. With eyes open the displacements were minimal at 0.3 Hz. With eyes closed, however, induced sway was maximal at this frequency. The apparent lack of visual stabilization at the lowest frequency (0.01 Hz) might be attributed to a subthreshold velocity of the retinal image motion induced by the swaying body. A similar absence of visual stabilization of 1 Hz is assumed to indicate the limit of the working range of visual stabilization of posture. Independent of stimulus amplitude a phase lead of about 90 degrees was found at 0.01 Hz. This decreased with increasing frequency up to a phase lag of 100 degrees at the highest frequency (1 Hz). Head stabilization was generally more effective than hip stabilization. EMG recordings from the leg muscles suggest that with eyes closed the center of force is mainly stabilized by leg muscle activity, while with eyes open this stabilization is best, when vision allows for stabilization of body posture by intersegmental movements between head, trunk and legs.

Short (SL), medium (ML), and long (LL) latency EMG responses of leg muscles were recorded after perturbation of stance by means of a sudden toe-up tilt of a movable platform. 56 healthy children varying in age between 14 months and 15 years were investigated. All three responses were present when children were able to stand on the recording platform. The SL-response in the triceps surae muscle, which corresponds to the mono- and oligo-synaptic spinal stretch reflex, showed a decreasing latency up to the age of 5 years. This reflects the increasing peripheral nerve conduction velocity. The ML-response in the triceps surae muscle, which as the SL-response has no stabilizing effect in this experiment, showed somewhat delayed maturational changes. The LL-response in the relaxed anterior tibial muscle helps to restore upright posture even in the youngest children. Its maturational changes in terms of latency by far exceed the range that can be explained by the increase of peripheral and spinal conduction velocities. Its mechanisms of maturation, besides the biophysical optimalization of a polysynaptic network, may include learning in terms of selecting the shortest pathways by way of synaptic potentiation within structures involved in the supposedly transcortical pathway of the LL-response. Qualitative observations made during the trials showed that the pattern of postural adaptation changed with age, suggesting the development of additional intersegmental mechanisms.

The responses of vestibular nuclei neurons of relaxed unanaesthetized goldfish have been examined with trapezoid velocity stimuli under three conditions. Responses to horizontal body rotation in the dark (pure vestibular stimulation) resemble those observed in vestibular nerve afferents. Optokinetic responses to exclusive visual surround-motion are also direction-specific and, in contrast to vestibular responses, exhibit a tonic response to constant velocity. They show three different response profiles, classified A,B or C, based on the neuron's discharge rate: either increasing, decreasing or remaining constant once surround motion is maintained at constant velocity. Following these dynamic effects, optokinetic responses have a maintained modulation of resting discharge until deceleration commences. The time constants associated with the dynamic effects vary between 1 and 11 seconds. Steady-state modulation of optokinetic responses shows a weak relation to stimulus velocities exceeding 10 deg/sec. Responses to body rotation in the light were found to linearly combine the weighted vestibular and optokinetic responses so that accurate velocity information is available for sensory and motor functions independent of the neuron's vestibular (I,II) or optokinetic (A,B,C) response type. The principle of this visual-vestibular interaction is discussed with respect to multisensory processing within the vestibular nuclei.

The gain of the vestibulo-ocular reflex in the sagittal plane may be due to a cooperation between otoliths and the vertical semi-circular canals. The present space experiment was aimed at studying the influence of the absence of gravity stimulation on the otoliths, by comparing VOR gain and phase in space and on ground. Measurements were taken the 5th and the 7th day of flight, the subject being asked to perform, eyes closed, active head oscillations in pitch while fixating an imaginary target in front of him. No significant decrease of the VOR gain was found in space, but a change in phase was noted. A significant increase of the VOR gain was found 14 h after landing. Control experiments have been done on ground on several subjects. They indicate that pitch VOR gain during active head movements is about one, with eyes open in darkness at 1 Hz.

The excitability of hindlimb extensor and flexor motoneurons is tonically modulated by animal tilt and a large visual stimulus rotating about the animal's line of sight. This direction-specific modulation is opposite for extensor and flexor motoneurons and opposite for optokinetic and vestibular stimuli, thus combining to a functionally significant pattern of postural reflexes.

The functional role of spinal and supraspinal EMG-responses for the maintenance of upright human posture was investigated in ten healthy subjects standing on a force measuring platform, which could be rotated in pitch around an axis aligned with the subject's ankle joint. Voluntary changes of body posture prior to the platform movement by leaning forward or backward led to a change in the amplitude and temporal organization of EMG-responses as compared to platform movements starting from a neutral position. Tilting the platform toe-up while leaning backward led to an increase of the latency of the short- and medium-latency responses in the triceps surae muscle and to a decrease of the latency of the stabilizing response in the anterior tibial muscle. Functionally, a cocontraction of both antagonistic muscles could be observed which partly compensated for the destabilizing action of the "reflex" response in the stretched triceps surae muscle. In analogy, leaning forward and tilting the platform toe-down led to a cocontraction of the two antagonistic muscles. The observed changes of latencies of short-, medium-, and long-latency response show the functional variability of segmental and suprasegmental "reflex" mechanisms. EMG-activities, which are functionally destabilizing posture, can be suppressed or compensated by reflexive cocontractions of antagonists.

The functional role of short-, medium- and long-latency responses for the maintenance of upright posture was investigated in twenty healthy subjects standing on a platform which could be rotated in pitch around the subject's ankle joints. Tilting the platform toe-up evokes a stretch reflex in the triceps surae muscle (TS, latency 55-65 ms) and at higher speeds and amplitudes of platform displacement a medium-latency response (latency 108-123 ms). Both responses functionally destabilize posture, since they enforce the induced backward displacement of the body. Compensation of body displacement in this situation is achieved by a long-latency EMG response in the anterior tibial muscle (TA 130-145 ms). Platform movement toe-down elicits a rather small medium-latency response in TA (103-118 ms), but no short-latency response. A late compensatory response occurs in the triceps surae muscle (latency 139-170 ms). The mean latency of the late antagonistic EMG response was significantly shorter than that of a voluntary movement triggered by a somatosensory stimulus. Integrals of rectified EMG responses from the two muscles were linearly related to the amplitude and to a smaller degree to the velocity of platform displacement. The slope of this function (gain) varied depending on the direction of ankle displacement and the functional importance of the subsequent EMG responses. Destabilizing short- and medium-latency responses of the stretched muscle had a lower gain relative to amplitude than the late stabilizing response of the antagonist. This functionally adaptive modulation of gain was not seen in relation to the rate of platform displacement.