Attenuation of the effect of remote muscle contraction on the soleus H-reflex during plantar flexion.

OBJECTIVE: We investigated to what extent the facilitation of the soleus (Sol) Hoffmann (H-) reflex during a phasic voluntary wrist flexion (Jendrássik maneuver, JM) can be modulated by graded plantar flexion force and conditioning wrist flexion force. METHODS: The subjects were asked to perform phasic wrist flexion under a reaction time condition. Sol H-reflex was evoked by stimulating the right tibial nerve at various time intervals (50-400ms) after the 'Go' signal for initiating JM while the ankle was at rest and while plantarflexing. The level of tonic plantar flexion force (isometric contraction of 10, 20 and 30% of maximal EMG) and conditioning wrist flexion (isometric contraction of 30, 50 and 80% of maximum voluntary contraction) during JM was graded systematically. RESULTS: Although JM facilitation could be seen 80-120ms after the flexor carpi radialis (FCR) EMG onset even while plantarflexing, the magnitude of JM facilitation under plantar flexion was significantly decreased compared to that at rest. The degree of decrease in JM facilitation did not depend on the level of plantar flexion force. In contrast, the degree of JM facilitation was proportional to the level of wrist flexion force while the ankle was at rest and while plantarflexing, though the amount of JM facilitation significantly decreased while plantarflexing. CONCLUSIONS: JM facilitation of Sol H-reflex is decreased while performing tonic voluntary contraction of the homonymous muscle. The degree of decrease in JM facilitation is independent of the level of homonymous muscle contraction, but depends on the level of remote FCR contraction. In clinical application, when we intend to elicit a maximum stretch reflex by JM, full relaxation of homonymous muscle should be carefully confirmed. SIGNIFICANCE: Our results provide evidence for better understanding of the features of JM and insight into its clinical application.

Patterns of muscle activation in human hopping.

In the present study, we examined the electromyogram (EMG) patterns of the soleus and medial gastrocnemius (MG) muscles during rhythmical, two-legged hopping to investigate the contributions of the monosynaptic short- and long-latency stretch reflexes during such a natural movement in human. During rhythmical hopping, soleus muscle is activated reflexly at near-monosynaptic latency by stretch resulting from passive ankle flexion upon landing. Soleus muscle also contracts voluntarily in order to launch the body into the next hop. This is part of the rhythmical bursts of activity producing the hops. Depending on the hopping interval, this phase of activation can follow the short-latency phase or precede landing at very short hopping intervals. In MG, there is an initial phase of activity that stiffens the muscle in preparation for landing, and continues through the contact phase. The monosynaptic reflex response to landing is usually superimposed on this activity. Depending on the hopping interval, both of these responses may be overlaid with activity that is time-locked to the take-off into the next hop, and serves to launch the body into the next hop. However, no evidence for a long-latency stretch reflex was found. In addition, the preferred hopping frequency for all subjects was about 2 Hz. This frequency is associated with a pattern of EMG activity the timing of which indicates that it balances the requirement for a comfortable landing from a hop with the optimal muscle activation required for launching the following hop.

Selective modification of somatosensory evoked potential during voluntary finger movement in humans.

We examined changes in somatosensory evoked potentials (SEPs) during voluntary movement of fingers innervated by the stimulated nerve and those not innervated by the stimulated nerve and the relationship to the kind of movement modality. Analysis showed that the amplitude of most components at F3, C3', and P3, except for P45 at C3, N35 and P45 at P3, decreased during voluntary finger movement tasks. Further, we found that the components of P40 at F3, P45 at C3', and N35 at P3 were increased during the voluntary pulling movement of the second and the third digits compared to those during the voluntary pushing movement of the fourth and the fifth digits, whereas all other components were decreased at F3, C3', and P3. We also found that not all components of SEPs were decreased while some SEPs in middle latency were increased. In conclusion, we confirmed the selectivity in attenuation of the SEPs. Moreover, we noted an interesting finding that the selectivity of attenuation of the SEPs was most frequently observed in the N20, P30 (P25 at F3), N35 (N30 at F3), and P45 (P40 at F3) components at F3, C3', and P3.

Changes in central motor-sensory system following voluntary movement of stimulated finger.

OBJECTIVE: We examined which components of somatosensory evoked potentials (SEP) were modulated during the voluntary movement of stimulated fingers and discuss our findings with respect to the decrease in SEPs in terms of both centrifugal and centripetal mechanisms. METHODS: The study was performed in the Laboratory of Physiology, Institute of Health and Sport Sciences, University of Tsukuba. Experiments were performed on 10 healthy young male subjects, aged 19-21 years. All subjects were right-handed and were free of neurological disease. RESULTS: We found that N20-P25 (P22 at frontal area) -N35-(N30 at frontal) P45 (P40 at frontal area) components at the F3, the C3' and the P3 were consistently evoked, but N35 was increased at P3 and N30 attenuation at F3 during a sustained voluntary movement. Furthermore, the present experiments showed that frontal N30 was markedly decreased during a sustained voluntary movement. We also found that all SEPs components, except for parictal N35, were significantly decreased during sustained voluntary movement tasks. CONCLUSIONS: We concluded that the suppression of SEPs appeared to be due to the interaction of both centrifugal and centripatal mechanisms. Namely, in a particular type of movement task, more prenounced afferent mechanisms may contribute to the decrease in SEPs, while in the other types of tasks, more pronounced afferent activities may contribute to the decrease in SEPs.

Inhibition of the soleus H-reflex during dorsiflexion is dependent on individual differences in maximal soleus H-reflex as a test reflex.

The quantitative differences among individuals in the natural reciprocal inhibition of the soleus H-reflex during dorsiflexion were examined, in conjunction with the maximal H-reflex as the test reflex size in each individual. Maximal H-reflex was expressed relative to the maximal M-response (H(max)) when compared among individuals. Analysis showed that with increases in H(max) at rest in each individual, the inhibitory effect was first enhanced, then reached a peak, and was finally alleviated. This pattern was similar to the intraindividual pattern of the inhibitory effect induced by specific conditioning stimulus as a function of the test reflex size.

Evident difference in the excitability of the motoneuron pool between normal subjects and patients with spasticity assessed by a new method using H-reflex and M-response.

The excitability of the motoneuron (MN) pool in the resting state was compared between normal control subjects and patients with spasticity resulting from HTLV-I-associated myelopathy, using a new parameter, Hslp/Mslp, and the conventional parameters Hmax/Mmax and Hth/Mth. Differences in the excitability of the MN pool between these two groups reached a high degree of statistical significance only when assessed with the new parameter. This suggests the methodological advantage of the Hslp/Mslp over both Hmax/Mmax and Hth/Mth for evaluation of the excitability of the MN pool in the resting state.

Somatosensory evoked potentials following voluntary movement during upper arm compression.

We examined the modulation of somatosensory evoked potentials (SEPs) during upper arm compression and following voluntary movement during upper arm compression. Most SEPs were significantly decreased, although some SEPs showed a slight, non-significant diminution. SEPs are mediated not only by myelinated fibers but also by mixed nerves and afferents from cutaneous, joints, and deep tissues, these being dependent upon the dorsal column-medical lemniscal system. Therefore, most of the diminution in SEPs found here may have been due to afferent occlusion from muscle, cutaneous, joints, and deep tissues, since normal SEPs are selectively modulated, corresponding to motor or mental tasks, regardless of whether gating is centrifugal or centripetal. In addition, the present experiments showed that all the SEPs at FZ, C3' and CZ were significantly decreased following voluntary movement at a pressure 25%-30% higher than the subject's systolic blood pressure. Comparing SEPs during upper arm compression and those following voluntary movement during upper arm compression at these pressures, we found that the SEPs at C3' were significantly diminished following voluntary movement during upper arm compression, with other SEPs showing slight, non-significant attenuation. In conclusion, it is possible that the diminution in SEPs following voluntary movement could be responsible for sensory inputs, however, when sensory inputs are present, centrifugal modulation would also be responsible for this diminution.

Hemispace asymmetries and laterality effects in arm positioning.

Hemispace asymmetries and laterality effects were examined on an arm positioning reproduction task. Sixteen male subjects were asked to reproduce both abductive and adductive positioning movements with the left or right arm within either the left or the right hemispace. Hemispace was manipulated using a 90 degrees head-rotation paradigm. A left hemispace advantage in positioning accuracy was predicted for both left and right arm movements on the grounds that the perceptual-motor control of positioning movements made in left hemispace is primarily mediated by the right hemisphere which is known to be advantageous for tasks which are spatial in nature (Heilman, Bowers, & Watson, 1984). No arm laterality effects were predicted to occur because the proximal musculature involved in the control of arm movements is innervated from both contralateral and ipsilateral cerebral hemispheres (Brinkman & Kuypers, 1973). Results showed that the predicted left hemispace advantage was evident for the right arm on the positioning variability measure alone, whereas it was absent for all other possible conditions on all error measures. Laterality (arm) effects were absent as predicted. The experiment also demonstrated a greater degradation of reproduction performance under the "crossed" arm-hemispace conditions than under the "uncrossed" conditions. A plausible explanation for the uncrossed advantage for the task is that under normal conditions, a single hemisphere is primarily responsible for both controlling the contralateral arm and directing attention to the contralateral hemispace, and consequently potential interhemispheric interference is minimized. A clear response bias effect in movement reproduction was also evident as a function of the direction of concurrent arm movement and head rotation. Arm movements made in the same direction as head rotation were systematically undershot in reproduction to a much greater degree than arm movements made in the opposite direction to head rotation.

Frontal N30 potential following a sustained voluntary movement.

With 6 male subjects we investigated whether the decreased amplitude in the frontal N30 potential of Somatosensory Evoked Potentials is related to a sustained voluntary movement or not. We concluded that the diminution of frontal N30 was also closely related to the sustained voluntary movement.

Excitability of the soleus motoneuron pool revealed by the developmental slope of the H-reflex as reflex gain.

The excitability of a motoneuron (MN) pool was evaluated by the developmental slope of H-reflexes (Hslp) evoked at a range of a stimulus intensity less than the threshold of an M-response. The Hslp has been regarded as the "reflex gain", which is the changing rate in MN excitability as a function of the increase of Ia input to an MN pool. In a comparison of two parameters used in the H-reflex technique, such as the ratio of the maximal H-reflex to the maximal M-response and the ratio of the threshold of an H-reflex to that of an M-response, the Hslp was predicted to be a reasonable parameter to evaluate motoneuronal excitability, because the Hslp is free from the effect of any collision between the H-reflex discharge and the antidromic volley drived from the occurrence of an M-response within the alpha-efferents, and the Hslp can estimate the recruitment properties of a whole MN pool. The Hslp was alleviated during dorsi-flexion and steepened during plantar-flexion, according to the inhibitory or facilitatory synaptic modifications onto soleus MNs. The developmental slope of an M-response (Mslp), which shows the recruitment property of axons of soleus MNs, was alleviated especially in plantar-flexion. In order to exclude the peripheral neuromuscular factors in evaluating substantial MN excitability, the Hslp/Mslp is proposed as a more effective parameter than the Hslp.

Auditory middle latency responses under different task conditions.

We examined the relationship between the Na and Pa components of human MLRs and the performance of different tasks. We also investigated whether MLRs are reliable indices of activity in the central motor-sensory system. The click stimuli we used consistently evoked the Na and Pa components. At CZ, the Na and Pa components significantly decreased for all tasks other than pegging with right hand while at FZ, these components were significantly decreased for all tasks. The Na and Pa latencies were slightly increased during task performances. These results indicate that the Na and Pa components of human MLRs decreased when various tasks were performed, while subjects were concentrating. A general principle of evoked potentials is that latencies decrease as amplitudes increase in excitation due to neural activation. Thus, it would appear that, under the conditions of this study, the pathways from the reticular formation and the thalamus to the primary auditory cortex were inhibited. Since the thalamus is considered to be the relay region for poly-sensory inputs, it is thought that the attenuation of the MLRs and SEPs occurs at the level of cerebral cortex, including the reticular formation, the thalamus, and the primary auditory cortex. Accordingly, since it is inferred that central factors are responsible for the attenuation of the MLRs, Na and Pa components observed during the performance of tasks carried out in the present experiment, it may be concluded that MLRs are reliable indices of activity in the central-motor system.

Reconsidering the 90 degrees head-rotation paradigm used in neuropsychological research: are there reflexive rather than hemispatial effects?

The 90 degrees head-rotation paradigm has often been used in neuropsychological studies to manipulate external hemispace (circumcorporeal space) relative to the head. Under the 90 degrees head-rotation paradigm, the performance of limb and hand movements carried out within the left or right hemispace as defined by head positions relative to the body is likely to be affected by the reflexive effect due to the neck and vestibular afferent inputs elicited by the head rotations, as well as by the hemispatial effect. Using the H-reflex technique, the present study examined whether the reflexive effect on the spinal motoneuron excitability occurred with head rotations under the 90 degrees head-rotation paradigm. The results showed that the amplitudes of H-reflexes evoked on both the thumb flexor and soleus muscles were not affected by head rotations, indicating no reflexive change in the spinal motoneuron excitability for both the thumb and soleus muscles. This finding suggests that the reflexive effect due to neck and vestibular afferent inputs can be ruled out from possible causal factors influencing the motor performance of limb and hand movements performed within the left or right hemispace as manipulated by the 90 degrees head-rotation paradigm.

Threshold of the soleus muscle H-reflex is less sensitive to the change in excitability of the motoneuron pool during plantarflexion or dorsiflexion in humans.

In the present study, we investigated whether weak (10% of maximal voluntary contraction) tonic dorsiflexion (DF) and plantarflexion (PF) affects the two conventional parameters used for evaluating the excitability of the soleus motoneuron (MN) pool, i.e. the ratio of the threshold of H-reflex to that of M-response (Hth:Mth) and the ratio of the maximal amplitude of H-reflex to that of M-response (Hmax:Mmax) in human subjects. The results showed that the Hmax:Mmax decreased during DF and increased during PF compared with that during rest, whereas no clear alteration was observed in Hth:Mth. These results are consistent with the scheme proposed by earlier workers, who have argued that neither inhibitory nor facilitatory effects of the conditioning stimulus apply to specific spinal reflex circuits occurring around the threshold of the test H-reflex. It is suggested, therefore, that the conventional use of the Hth:Mth ratio as a parameter reflecting the excitability of the MN pool should be reconsidered.

Separation of human tear proteins with ceramic hydroxyapatite high-performance liquid chromatography.

Human tear protein, which consists mainly of albumin, lysozyme, and lactoferrin, was assayed with high-performance liquid chromatography using a new ceramic hydroxyapatite column. Proteins were eluted at room temperature using a 20-min linear gradient from 95:5% A/B buffer to 0:100% A/B buffer (buffer A, distilled water; buffer B, 400 mM KH2PO4 containing 240 mM NaOH). The proteins eluted at 1.2 min for albumin, 8.5 min for lysozyme, and 20.4 and 21.7 min for lactoferrin, respectively. The assays may be performed in 30 min.

Information-processing mediating the location-distance interference in motor short-term memory.

We reviewed the literature on basic psychological correlates of the well-known phenomenon of the location-distance interference in motor short-term memory (Kerr, 1978; Walsh, Russell, Imanaka, & James, 1979). The location-distance interference in motor short-term memory has frequently been demonstrated as an unavoidable interference phenomenon observed in the reproduction of movement location and distance in arm positioning. The most important aspect of this phenomenon is that even when a subject concentrates on a specific cue (i.e., either end-location or distance) the other cue is also coded unintentionally and, as a result, the reproduction movement guided on the basis of the specific cue is unavoidably influenced by the other nonspecific cue. In this review article, we first reviewed the literature on the basic theories and nature of short-term memory, particularly on the limited processing capacity. We then referred to the unlimited, automatic processing in visual-verbal domains, referring to the Stroop phenomenon. Finally, in conjunction with the notion of automatic processing, we examined the possible aspects of information processing which may be responsible for mediating the location-distance interference in motor short-term memory.

Retrograde neuronal labeling of motoneurons in the rat by fluorescent tracers, and quantitative analysis of oxidative enzyme activity in labeled neurons.

Extensor digitorum longus motoneurons in the rat spinal cord were identified by retrograde labeling with two fluorescent tracers, Fast blue (FB) and Nuclear yellow (NY). Labeled motoneurons had a blue fluorescent cytoplasm at 360 nm excitation wavelength with FB, and a golden-yellow fluorescent nucleus with NY on the cryostat section. Labeled motoneurons were further examined for succinate dehydrogenase activity on the same section used for identification of the motoneurons. This study demonstrates that fluorescent dyes are useful for neuroanatomical studies by the retrograde axonal transport method, and that quantitative analysis of metabolic activity in labeled motoneurons is also possible.

Attenuation of somatosensory evoked potentials immediately following rapid reaction movement.

We investigated the changes in the somatosensory evoked potentials (SEPs) and particularly which components of the SEPs altered immediately after rapid reaction movements. N20, P23, N35, and P45 components measured over the scalp were all attenuated except for the P45 component measured at CZ. Particularly, the N20 and P23 components were markedly attenuated at both C3' and CZ immediately following rapid reaction movements. N18 and N20 components are thought to represent thalamocortical and primary sensory cortex activities. Therefore, the present results suggest that following rapid reaction movements, the thalamocortical and primary sensory activities are significantly attenuated. Moreover, following rapid reaction movements, the onset of EMG activity preceded the afferent discharge of the muscle spindle. Therefore, it may be concluded that central factors were principally responsible for the attenuation of the SEPs immediately following rapid reaction movements.

Suppression of cerebral evoked potentials preceding rapid reaction movement.

In this study we examined the gating of the cerebral evoked potentials during warning-imperative stimulus intervals and preceding the rapid movement. We also examined the possibility of the gating phenomenon occurring in the precentral cortex. The experiment showed that the amplitudes of cerebral evoked potentials increase at all C3 and CZ during the warning-imperative stimulus intervals. Therefore, it seems that gating does not occur within the central nervous system during warning-imperative stimulus intervals. Furthermore, we observed that preceding rapid movement, the amplitudes of the cerebral evoked potentials decreased at all C3 and CZ. In the present experiment, it does not seem that a peripheral mechanism plays a role in the rapid movement. Therefore, it can be concluded that the decrease in amplitude of the cerebral evoked potentials preceding rapid movement may be due to a central mechanism irrespective of afferent information. In addition, we confirmed that the amplitude of cerebral evoked potentials decreases considerably at the precentral cortex and that the attenuation in amplitude of cerebral evoked potentials is more pronounced over the CZ than over the C3.

Cerebral motor potential preceding grip strength movement.

The relationship between the force of the movement and the motor potential (MP), and that between the degree of the muscular discharge and the MP were examined by the method of bipolar and monopolar derivation. The MP amplitudes were significantly higher at C3' and Cz than at other lead placements for right grip and higher at C4' and Cz for left grip. A significant difference was observed among all lead placements except at Cz-C4'. The MP amplitudes were significantly higher for a strong grip task than for a weak grip tasks. Moreover, the MP amplitudes were higher at higher degrees of muscular discharge. The MP usually occurred about 100 msec prior to the EMG discharge, as also confirmed by Deecke et al. Therefore, from the present results and other studies, we postulated that the MP may reflect the activity of the neurons in the motor and premotor cortices, which might contribute to movement initiation. In addition, the MP amplitudes measured using bipolar derivation were more explicit than those measured using monopolar derivation.