Postexercise depression of motor evoked potentials: a measure of central nervous system fatigue

Fatigue of voluntary muscular effort is a complex and multifaceted phenomenon. Fatigue of peripheral nervous system components, including the contractile apparatus and the neuromuscular junction, has been well studied. Central nervous system components also fatigue, but studies have lagged for want of objective methods. Transcranial magnetic stimulation is a relatively new technique that can be used to assess central nervous system excitability from the motor cortex to the alpha-motoneuron. In six normal volunteers, including four of the investigators, the amplitudes of motor evoked potentials elicited by transcranial magnetic stimulation were transiently decreased after exercise, indicating fatigue of motor pathways in the central nervous system. The decrease in amplitude was associated with a feeling of fatigue. The mechanism of this phenomenon is apparently decreased efficiency in the generation of the motor command in the motor cortex.     

Automatic postural responses in the cat: responses of distal hindlimb muscles to paired vertical perturbations of stance

Summary The active components of the quadrupedal diagonal stance response to rapid removal of the support from beneath a single limb were studied in cats to further define the mechanisms that trigger and generate the response. We recorded EMG activity from lateral gastrocnemius and tibialis anterior muscles in awake, behaving cats while they stood on an hydraulic posture platform. By dropping the support from beneath a single limb, we evoked the diagonal stance response, with its characteristic changes in vertical force and EMG patterns. As the animal responded to this drop, a second perturbation of posture was then presented at intervals of 10 to 100 ms following the first. This second perturbation, which consisted of dropping the support from beneath the two limbs that were loaded as a result of the initial limb drop, made the first response biomechanically inappropriate. The EMG responses observed in both muscles during paired perturbations were triggered by the somatosensory events related to the perturbations. Muscle responses that were appropriate for the first perturbation always occurred with amplitudes and latencies similar to control trials. This was true even when the second perturbation occurred 10–20 ms after the first, that is, when this perturbation either preceded or was coincident with the response to the initial limb drop. The EMG responses that were normally associated with the second perturbation were delayed and/or reduced in amplitude when the time interval between perturbations was short. As the inter-perturbation interval was lengthened beyond 60–100 ms, however, EMG responses to the second perturbation were unaffected by the occurrence of the first perturbation. When the hindlimb containing the recording electrodes was dropped as part of the second perturbation, a myotatic latency response was observed in tibialis anterior. The amplitude of this response to the second perturbation was greater than controls when this displacement was presented during the period between initiation of the first perturbation and execution of the response to it. When the second displacement was presented after execution of the first response began, the amplitude of the myotatic response was reduced below control levels. While the results do not preclude the possibility that these automatic postural responses are segmental or suprasegmental reflexes, they support the hypothesis that the active component of the response to drop of the support beneath a single limb is centrally programmed and that the appropriate response can be riggered very rapidly by the somatosensory information signalling the perturbation.Supported by NIH grants NS19484 and RR05593 as well as the Medical Research Foundation of Oregon, and the Neurological Sciences Center of Good Samaritan Hospital and Medical Center     

Development of postural control in children: short-, medium-, and long latency EMG responses of leg muscles after perturbation of stance

Summary 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.     

Visual evoked responses to the upper and lower half-field stimulation in a dark-adapted man

The averaged visual evoked responses (VERs) to flashed blank and checkerboard-patterned (32 checkers) stimulation of the upper and lower half-field of 6° angular subtense were recorded referentially (O_z–A₁₊₂) in seven dark adapted subjects using a luminance range of 3.69 log units (maximum luminance=2220 cd/m²).With the logarithmical rise of stimulation luminance the peak latencies of the maximum positive-negative deflection of the VERs to the upper and lower half-field blank and to the upper half-field patterned stimulation display a monotonick shortening. These three response types do not differ in waveform, polarity and amplitude. Their amplitudes show no significant luminance-dependent changes.The peak latencies of the VERs to the lower half-field patterned stimulation exhibit an U shaped course with the increase of the luminance. At lower luminances they are by about 30 ms shorter in comparison with the VERs to the other three stimulation types, at higher luminances a gradual lengthening is observable. Consequently, at low luminances these VERs have a reversed polarity, at higher luminances the same polarity as the VERs to the other three stimulation types. The amplitude values of the lower half-field patterned responses are the highest and show a triphasic luminance-dependent course.From these and further differences between the VERs to the upper and lower half-field patterned stimulation in connection with the reaction to defocusing and to the subtraction of the luminance-related part of the response it is concluded that the VERs to the lower half-field patterned stimulation only contain a pattern-related component.     

Automatic postural responses in the cat: responses of hindlimb muscles to horizontal perturbations of stance in multiple directions

Summary The effect of the direction of unexpected horizontal perturbations of stance on the organization of automatic postural responses was studied in cats. We recorded EMG activity in eight proximal and distal muscles of the hindlimb along with vertical forces imposed by the limbs in awake behaving cats while they stood on an hydraulic platform. Postural responses to motion of the platform in 16 different horizontal directions were recorded. Vertical force changes were consistent with passive shifts of the center of mass and active correction of stance by the animals. When the perturbation was in the sagittal plane, vertical force changes began about 65 ms following initial platform movement. When the perturbation contained a component in the lateral direction, latency for vertical force changes was about 25 ms and an inflection in the vertical force trace was observed at 65 ms. No EMG responses were observed with latencies that were short enough to account for the early force component and it was concluded that this force change was due to passive shifts of the center of mass. The amplitude of the EMG responses of each muscle recorded varied systematically as perturbation direction changed. The directions for which an individual muscle showed measurable EMG activity were termed the muscle's angular range of activation. No angular range of activation was oriented strictly in the A-P or lateral directions. Most muscles displayed angular ranges of activation that encompassed a range of less than 180°. Onset latencies of EMG responses also varied systematically with perturbation direction. The amplitude and latency relationships between muscles, which made up the organization of postural responses, also varied systematically as perturbation direction was changed. This result suggests that direction of perturbation determines organizational makeup of postural responses, and for displacements in the horizontal plane, is considered a continuous variable by the nervous system.     

Automatic postural responses in the cat: responses of proximal and distal hindlimb muscles to drop of support from a single hind- or forelimb

Summary Cats respond to drop of the support from beneath a single limb with the diagonal stance response (Coulmance et al. 1979). They load the limbs on the diagonal opposite to the one containing the dropped limb and unload the third supporting limb in the diagonal containing the dropped limb. Characteristic biomechanical delays in limb motion and in vertical force changes imposed upon the limbs are observed. These delays range from 30 to 45 ms, depending upon the location of the dropped limb. This study describes the kinematics of the diagonal stance response and the activation of selected agonist-antagonist muscle pairs acting on the joints of the hindlimb during the response. Proximal and distal hindlimb muscles respond to perturbations in groups that are appropriate to the vertical forces imposed upon the limb. When the hindlimb containing the recording electrodes is loaded by drop of the contralateral hindlimb or the ipsilateral forelimb medium latency (25–45 ms) EMG responses occur in the extensors. This response serves to stiffen the limb against the increased vertical force of loading. A similar response is observed when the hindlimb is reloaded after being dropped. In this case, however, short latency responses precede the medium latency responses in muscles that are passively stretched by the limb drop. When drop of the diagonal forelimb unloads the hindlimb containing the electrodes, medium latency responses are observed in the distal hindlimb flexors, which indicates that the unloading is evoked in part by active lifting of the limb. In most cases, the medium latency responses precede or are coincident with the changes in force imposed on the limb, suggesting that the observed responses are centrally programmed.     

Human postural responses to motion of real and virtual visual environments under different support base conditions

The role of visual orientation cues for human control of upright stance is still not well understood. We, therefore, investigated stance control during motion of a visual scene as stimulus, varying the stimulus parameters and the contribution from other senses (vestibular and leg proprioceptive cues present or absent). Eight normal subjects and three patients with chronic bilateral loss of vestibular function participated. They stood on a motion platform inside a cabin with an optokinetic pattern on its interior walls. The cabin was sinusoidally rotated in anterior-posterior (a-p) direction with the horizontal rotation axis through the ankle joints (f=0.05-0.4 Hz; A (max)=0.25 degrees -4 degrees ; v (max)=0.08-10 degrees /s). The subjects' centre of mass (COM) angular position was calculated from opto-electronically measured body sway parameters. The platform was either kept stationary or moved by coupling its position 1:1 to a-p hip position ('body sway referenced', BSR, platform condition), by which proprioceptive feedback of ankle joint angle became inactivated. The visual stimulus evoked in-phase COM excursions (visual responses) in all subjects. (1) In normal subjects on a stationary platform, the visual responses showed saturation with both increasing velocity and displacement of the visual stimulus. The saturation showed up abruptly when visually evoked COM velocity and displacement reached approximately 0.1 degrees /s and 0.1 degrees , respectively. (2) In normal subjects on a BSR platform (proprioceptive feedback disabled), the visual responses showed similar saturation characteristics, but at clearly higher COM velocity and displacement values ( approximately 1 degrees /s and 1 degrees , respectively). (3) In patients on a stationary platform (no vestibular cues), the visual responses were basically similar to those of the normal subjects, apart from somewhat higher gain values and less-pronounced saturation effects. (4) In patients on a BSR platform (no vestibular and proprioceptive cues, presumably only somatosensory graviceptive and visual cues), the visual responses showed an abnormal increase in gain with increasing stimulus frequency in addition to a displacement saturation. On the normal subjects we performed additional experiments in which we varied the gain of the visual response by using a 'virtual reality' visual stimulus or by applying small lateral platform tilts. This did not affect the saturation characteristics of the visual response to a considerable degree. We compared the present results to previous psychophysical findings on motion perception, noting similarities of the saturation characteristics in (1) with leg proprioceptive detection thresholds of approximately 0.1 degrees /s and 0.1 degrees and those in (2) with vestibular detection thresholds of 1 degrees /s and 1 degrees , respectively. From the psychophysical data one might hypothesise that a proprioceptive postural mechanism limits the visually evoked body excursions if these excursions exceed 0.1 degrees /s and 0.1 degrees in condition (1) and that a vestibular mechanism is doing so at 1 degrees /s and 1 degrees in (2). To better understand this, we performed computer simulations using a posture control model with multiple sensory feedbacks. We had recently designed the model to describe postural responses to body pull and platform tilt stimuli. Here, we added a visual input and adjusted its gain to fit the simulated data to the experimental data. The saturation characteristics of the visual responses of the normals were well mimicked by the simulations. They were caused by central thresholds of proprioceptive, vestibular and somatosensory signals in the model, which, however, differed from the psychophysical thresholds. Yet, we demonstrate in a theoretical approach that for condition (1) the model can be made monomodal proprioceptive with the psychophysical 0.1 degrees /s and 0.1 degrees thresholds, and for (2) monomodal vestibular with the psychophysical 1 degrees /s and 1 degrees thresholds, and still shows the corresponding saturation characteristics (whereas our original model covers both conditions without adjustments). The model simulations also predicted the almost normal visual responses of patients on a stationary platform and their clearly abnormal responses on a BSR platform.     

Effect of contraction strength on responses in biceps brachii and adductor pollicis to transcranial magnetic stimulation

The sizes of the motor-evoked potentials (MEPs) and the durations of the silent periods after transcranial magnetic stimulation were examined in biceps brachii, brachioradialis and adductor pollicis in human subjects. Stimuli of a wide range of intensities were given during voluntary contractions producing 0–75% of maximal force (maximal voluntary contraction, MVC). In adductor pollicis, MEPs increased in size with stimulus intensity and with weak voluntary contractions (5% MVC), but did not grow larger with stronger contractions. In the elbow flexors, MEPs grew little with stimulus intensity, but increased in size with contractions of up to 50% of maximal. In contrast, the duration of the silent period showed similar changes in the three muscles. In each muscle it increased with stimulus intensity but was unaffected by changes in contraction strength. Comparison of the responses evoked in biceps brachii by focal stimulation over the contralateral motor cortex with those evoked by stimulation with a round magnetic coil over the vertex suggests an excitatory contribution from the ipsilateral cortex during strong voluntary contractions. Received: 12 August 1996 / Accepted: 14 May 1997     

Non-Painful and Painful Surface and Intramuscular Electrical Stimulation at the Thenar and Hypothenar Sites: Differential Cerebral Dynamics of Early to Late Latency SEPs

Little is known about somatosensory evoked potentials (SEPs) from muscle stimulation compared to that from skin stimulation. The current study examined this issue in the full SEP spectrum (0 - 440 ms). The aims of the study were to (1) establish the dynamics of early to late latency SEPs from intramuscular stimulation in contrast to surface stimulation, (2) compare the effect of non-painful and painful stimuli on SEP latencies and amplitudes of the two methods, and (3) investigate to which extent these results can be shared between the median nerve innervated thenar site and ulnar nerve innervated hypothenar site. Stimuli were delivered (2 Hz) at a non-painful and a painful intensity above or within the thenar and hypothenar muscles of the hand. Maximas of the SEPs were extracted by a combination of global field power and visual inspection of the topographies. Amplitudes and latencies of the maximas were analysed by a two-way ANOVA with repeated measures. In the early phase (0 - 50 ms) the topographic patterns showed different dynamics between surface and intramuscular stimulation and in the late phase (100- 440 ms) prolonged latencies were found for intramuscular stimulation. Apart from this, similar topographic patterns and time sequences were obtained. Significant higher SEP amplitudes for most of the isolated components (C4'/P25, Fz/N35, C4'/P45, Fc2/N65, P4/P90, T4/N137, F3/P150, Cz/P240-P270) were found with surface stimulation compared to intramuscular stimulation. In contrast to surface stimulation, intramuscular stimulation at a stimulation frequency of 2 Hz did not result in a differentiation in amplitude for any of the isolated components. These results indicate differences in the early and late processing of sensory input from skin and muscle.