Linearity,symmetry and additivity of the human eye-movement response to maintained unilateral and bilateral surface galvanic (DC) vestibular stimulation

Recent studies have shown that, although responses to long-duration, constant-current surface galvanic vestibular stimulation (GVS) show substantial interindividual variability, individual subjects show a reliable, repeatable, idiosyncratic oculomotor response pattern to GVS. It follows that GVS may be a more reliable stimulus than may have been anticipated from the literature. The aim of the present study was to examine the metrics of 3D eye-movement responses to maintained (120 s), unilateral and bilateral surface GVS. Eye movements were measured using computerised video-oculography. Two experiments were conducted: Experiment 1 examined whether the normal response is linear over increasing levels of current; and Experiment 2 examined (1) whether the normal response to surface GVS is symmetrical when comparing stimulated sides, (2) whether the normal response to surface GVS is symmetrical when the polarity of the stimulating current was reversed, and (3) whether there is additivity in the normal response to combinations of unilateral/bilateral surface GVS. Five subjects participated in Experiment 1 and eight subjects participated in Experiment 2. In both experiments, the onset of stimulation produced characteristic eye-movement responses: changes in torsional position with the upper pole of both eyes rolling towards the anode and away from the cathode; together with horizontal and torsional nystagmus with slow phases towards the anode and away from the cathode; and negligible vertical nystagmus. These responses reversed direction at stimulus offset. In the fixation condition of Experiment 1, the magnitude of ocular torsional position (OTP) and torsional nystagmus responses showed a linear relationship over conditions of increasing current strength, as did OTP, torsional and horizontal nystagmus responses in darkness. The results of Experiment 2 showed that responses to unilateral stimulation are symmetrical between stimulated sides, symmetrical between stimulating polarities, and additive (with respect to responses to bilateral stimulation). The principles derived from these findings, as well as those of recent studies, provide a foundation for future work investigating eye-movement responses to surface GVS in patients with known types of vestibular dysfunction. Electronic Publication     

Effects of pre-motion electromyographic silent period on dynamic force exertion during a rapid ballistic movement in man

Summary The effects of pre-motion silent period (PSP) on dynamic force exertion were studied in ten healthy subjects performing ballistic elbow extensions. The experiments were designed to evaluate the significance of mean differences between the averaged dynamic force curves of two groups: PSP-presence groups and PSP-absence groups. The presence of PSP was judged quantitatively and automatically by means of a newly developed method using statistical analysis. The results indicated that there were two effects of PSP on dynamic force exertion: one was a reducing effect, observed prior to the movement; the other was a reinforcing effect, observed in the first part of the ballistic movement. The duration of the reinforcement was significantly correlated with the duration of the reducing effect of PSP. The findings suggested that the reinforcement of dynamic force may be produced by the pre-stretch of agonistic muscles caused by prior force reduction due to PSP occurrence. The fact that PSP plays an important role in dynamic force exertion suggests that PSP may be incorporated in the central motor control system designed to interrupt the background activity, to stretch the agonist and to reinforce the dynamic force.     

Age-related changes in smooth pursuit initiation

Quantitative analysis of eye movements is a useful tool for examining the behavioural effects of ageing. Although the effect of ageing on saccadic eye movement has been examined in some detail, the effect of ageing on a second class of eye movement, smooth pursuit (SP), has received less attention. We examined the initiation of SP in a group of fifteen healthy older people (mean age 72 years) and compared their performance with that of ten young controls (mean age 21 years). Although their performance was qualitatively similar, pursuit latency was increased in the older group. Investigation of the gap effect on pursuit revealed that, while the gap effect was present in the older group, it seemed to be directionally asymmetrical. When the longer absolute latencies were taken into account, although the gap effect in the two groups was identical for leftward tasks, for rightward tasks it was reduced in the older group, although this did not reach statistical significance. The difference between the old and young groups was driven by some of the older subjects. At the longest gap duration employed (400 ms), while there was a clear gap effect for leftward tasks in these subjects, there was no reduction in latency, or increases in latency, for rightward tasks. This asymmetry was not related to chronological age within the older group. These results suggest an age-related alteration in SP initiation that is more complex than general slowing of information processing in ageing. They may be indicative of additional ageing effects specific to the oculomotor or closely related systems.     

Influence of interaction force levels on degree of motor adaptation in a stable dynamic force field

Studies have shown that the point-to-point reaching movements of subjects seated in a dark, rotating room demonstrate errors in movement trajectories and endpoints, consistent with the direction of the Coriolis force perturbations created by room rotation. Adaptation of successive reaches and the presence of postrotation aftereffects have indicated that subjects form internal models of the Coriolis field dynamics in order to make appropriate movement corrections. It has been argued that these findings are inconsistent with predictions of peripheral stabilization assumed in equilibrium-point models of motor control. A possibility that has been raised, however, is that the Coriolis field findings may in fact stem from changes in control commands elicited due to the magnitude and destabilizing nature of the Coriolis perturbations. That is, it has been suggested that a perturbation threshold exists, below which central reactions are not necessary in order to maintain movement stability. We tested the existence of a perturbation threshold in normal-speed reaching movements. Twelve normal human subjects performed non-visually guided reaching movements while grasping a robotic manipulandum. The endpoints and trajectory deviations of their movements were measured before, during, and after a position-dependent force field (similar to a Coriolis field in terms of the time history of applied forces) was applied to their movements. We examined the responses to a range of perturbation field strengths from small to considerable. Our experimental results demonstrated a substantial adaptation response over the entire range of perturbation field magnitudes examined. Neither the amount of adaptation after 5 trials nor after 25 trials was found to change as disturbance magnitudes decreased. These findings indicate that there is an adaptive response even for small perturbations; i.e., threshold behavior was not found. This result contradicts the assertion that peripheral stabilization mechanisms enable the central controller to ignore small details of peripheral or environmental dynamics. Our findings instead point to a central dynamic modeler that is both highly sensitive and continually active. The results of our study also showed that subjects were able to maintain baseline pointing accuracies despite exposure to perturbation forces of sizeable magnitude (more than 7 N).     

Parabrachial neuron discharge in the cat is altered during the carbachol-induced REM sleep-like state (DCarb)

Pontine parabrachial neurons have been suggested to play a regulatory role in both respiratory and sleep cycle control. Encouraged by the finding that microinjections of the cholinergic agonist carbachol into the medial pontine reticular formation (mPRF) of the cat produced respiratory changes paralleling those observed during rapid eye movement (REM) sleep (Neurosci. Lett., 102 (1989) 211–216), this study tested the hypothesis that cholinergic mechanisms in the mPRF can also cause state-dependent changes in the discharge of parabrachial neurons. This paper describes extracellular recordings of parabrachial neurons during REM sleep and during the carbachol-induced REM sleep-like state (DCarb). Cells which were activated (REM-on) or inactivated (REM-off) during REM maintained these same state-dependent firing patterns during the DCarb state. These results support the hypothesis that cholinergic mechanisms in the mPRF can cause state-dependent changes in the discharge of parabrachial neurons.     

Coordination of multiple muscles in two degree of freedom elbow movements

The present study quantifies electromyographic (EMG) magnitude, timing, and duration in one and two degree of freedom elbow movements involving combinations of flexion-extension and pronation-supination. The aim is to understand the organization of commands subserving motion in individual and multiple degrees of freedom. The muscles tested in this study fell into two categories with respect to agonist burst magnitude: those whose burst magnitude varied with motion in a second degree of freedom at the elbow, and those whose burst magnitude depended on motion in one degree of freedom only. In multiarticular muscles contributing to motion in two degrees of freedom at the elbow, we found that the magnitude of the agonist burst was greatest for movements in which a muscle acted as agonist in both degrees of freedom. The burst magnitudes for one degree of freedom movements were, in turn, greater than for movements in which the muscle was agonist in one degree of freedom and antagonist in the other. It was also found that, for movements in which a muscle acted as agonist in two degrees of freedom, the burst magnitude was, in the majority of cases, not different from the sum of the burst magnitudes in the component movements. When differences occurred, the burst magnitude for the combined movement was greater than the sum of the components. Other measures of EMG activity such as burst onset time and duration were not found to vary in a systematic manner with motion in these two degrees of freedom. It was also seen that several muscles which produced motion in one degree of freedom at the elbow, including triceps brachii (long head), triceps brachii (lateral head), and pronator quadratus displayed first agonist bursts whose magnitude did not vary with motion in a second degree of freedom. However, for the monoarticular elbow flexors brachialis and brachioradialis, agonist burst magnitude was affected by pronation or supination. Lastly, it was observed that during elbow movements in which muscles acted as agonist in one degree of freedom and antagonist in the other, the muscle activity often displayed both agonist and antagonist components in the same movement. It was found that, for pronator teres and biceps brachii, the timing of the bursts was such that there was activity in these muscles concurrent with activity in both pure agonists and pure antagonists. The empirical summation of EMG burst magnitudes and the presence in a single muscle of both agonist and antagonist bursts within a movement suggest that central commands associated with motion in individual degrees of freedom at the elbow may be superimposed to produce elbow movements in two degrees of freedom.     

Paw-shake responses with joint immobilization: EMG changes with atypical feedback

Summary To determine the effects of atypical motion-related feedback on motor patterns of the paw shake, EMG patterns of selected flexor and extensor muscles were recorded under four conditions of joint immobilization (hip and ankle alone, hip-knee, hip-knee-ankle) and compared to responses evoked in the freely-moving hindlimb of the chronic-spinal cat. With only the ankle joint casted, paw shaking was easily evoked by applying tape to the paw, and cyclic characteristics were not altered. However, under the three conditions with hip-joint immobilization (hip alone, hip-knee, hip-knee-ankle), responses were difficult to obtain, and if elicited, the number of cycles within a response decreased and cycle periods were prolonged. The temporal organization of consecutive cycles, however, was not altered by immobilization of any joint(s). Ankle (LG) and hip (GM) extensor activity was relatively unaffected by conditions of joint immobilization. In contrast, hip flexor (IP) and knee extensor (VL) bursts were often absent under all three conditions of hip-joint immoblization, and if present, VL burst durations decreased under the casted hip-knee-ankle condition, while the onset of IP activity occurred early in the cycle with prolonged bursts under casted ankle and casted hip-knee-ankle conditions. The coactivity of the knee extensor (VL) and ankle flexor (TA) was disrupted by conditions of hip-joint immobilization: VL onset was dissociated from TA onset and coincident with LG onset. These results suggest that motion-related feedback from the hip joint is particularly important in the initiation, cycle frequency, and the number of cycles of paw-shake responses. The presence of atypical motion-dependent feedback from the hip joint altered activity of knee and ankle anterior muscles, while motion-dependent feedback from the ankle joint changed activity of the anterior hip muscle. Moreover, the results suggest a differential control of posterior and anterior muscles of the hindlimb, consistent with paw-shake limb dynamics.     

A simple method for computer quantification of stage REM eye movement potentials

We describe a simple method for computer quantification of eye movement (EM) potentials during REM sleep. This method can be applied by investigators using either period-amplitude (PA) or Fast Fourier Transform (FFT) spectral EEG analysis without special hardware or computer programming. It provides good correlations with visual ratings of EM in baseline sleep and after administration of GABAergic hypnotics. We present baseline data for both PA and FFT measures for 16 normal subjects, studied for 5 consecutive nights. Both visually rated and computer-measured EM density (EMD) showed high night-to-night correlations across baseline and drug nights and the computer measures detected the EMD suppression that is produced by GABAergic drugs. Measurement of EM in addition to stage REM provides biologically significant information and application of this simple computer method, which does not require pattern recognition algorithms or special hardware, could provide reliable data that can be compared across laboratories.     

Second-order vestibular neuron morphology of the extra-MLF anterior canal pathway in the cat

Second-order vestibular neurons form the central links of the vestibulo-oculomotor three-neuron arcs that mediate compensatory eye movements. Most of the axons that provide for vertical vestibulo-ocular reflexes ascend in the medial longitudinal fasciculus (MLF) toward target neurons in the oculomotor and trochlear nuclei. We have now determined the morphology of individual excitatory second-order neurons of the anterior semicircular canal system that course outside the MLF to the oculomotor nucleus. The data were obtained by the intracellular horseradish peroxidase method. Cell somata of the extra-MLF anterior canal neurons were located in the superior vestibular nucleus. The main axon ascended through the deep reticular formation beneath the brachium conjunctivum to the rostral extent of the nucleus reticularis tegmenti pontis, where it crossed the midline. The main axon continued its trajectory to the caudal edge of the red nucleus from where it coursed back toward the oculomotor nucleus. Within the oculomotor nucleus, collaterals reached superior rectus and inferior oblique motoneurons. Some axon branches recrossed the midline within the oculomotor nucleus and reached the superior rectus motoneuron subdivision on that side. Since these neurons did not give off a collateral toward the spinal cord, they were classified as being of the vestibulo-oculomotor type and are thought to be involved exclusively in eye movement control. The signal content and spatial tuning characteristics of this anterior canal vestibulo-oculomotor neuron class remain to be determined.     

Evidence for high-velocity smooth pursuit in the trained cat

It is generally accepted that in cats smooth pursuit velocity of the eye never exceeds a few degrees per second. This is in contrast with observations in primates, where smooth pursuit velocity can reach values as high as 100°/s. Cats were trained to fixate and pursue spots of light appearing on a translucent screen. Spots were moved in the horizontal and vertical planes at different constant velocities up to 80°/s. Eye position was recorded with the sclerai search coil technique. Naive cats did not pursue moving targets with high efficiency. Smooth eye movement velocity saturated at 5°/s. After a few days of training, smooth-pursuit eye velocity increased with target velocity and saturated at 25°/s on average. However, velocities twice as high have been observed frequently. When the target was unexpectedly extinguished, smooth eye movement velocity dropped to values close to 0°/s in approximately 350 ms. After a short training period (usually 5 times the same target presentation), the eye continued to move smoothly until the target reappeared. These data suggest that smooth pursuit eye movements of the cat are qualitatively similar to those of primates, but reach lower velocities and are more variable in their characteristics.