Your body and mine: a neuropsychological perspective.

The concept of "shared representations" suggests the existence of a common representation network between self and other. Strong support for this theory is derived from the discovery in the monkey of a population of neurons, which encode object-directed actions performed by oneself and by others. In the human, the issue of "shared representations" has been raised in brain-damaged studies, in particular in the setting of "pointing to body parts" disorders. According to the dominant view, body part designation engages a shared representation system, which encodes the visuospatial characteristics of both one's own body and the bodies of other individuals. However, the recent observation of two patients, JR and AP, with dissociated performance in pointing to body parts leads to question this model. JR presented a deficit in pointing to his own body parts, while his capacity to point to the body parts of other persons was not altered. AP exhibited the reverse pattern of impairment. Lesion study indicated a putative area of dysfunction setting in the left superior parietal lobule (SPL) in JR, and in the left inferior parietal lobule (IPL) in AP. This double dissociation, along with two subsequent neuroimaging studies, suggests that the left SPL and IPL participate in the building of differential representations between oneself and other individuals.

Relations between the directions of vibration-induced kinesthetic illusions and the pattern of activation of antagonist muscles.

In humans, tendon vibration evokes illusory sensations of movement that are usually associated with an excitatory tonic response in muscles antagonistic to those vibrated (antagonist vibratory response, AVR), i.e., in the muscle groups normally contracted if the illusory movement had been performed. The aim of the present study was to investigate the relation between the parameters of the illusory sensation of movement and those of the AVR and to determine whether vectorial models could account for the integration of proprioceptive inputs from several muscles, as well as for the organization of the elementary motor commands leading to one unified motor response. For that purpose, we analyzed the relations between the anatomical site of the tendon vibration, the direction of the illusory movement, the muscles in which the AVR develops, and the characteristics of the AVR (surface EMG, motor unit types, firing rates, and activation latencies). This study confirmed the close relationship between the parameters of an AVR and those of the kinesthetic illusion. It showed that, during illusions of movements in different directions, motor units are activated according to a specific pattern correlated with their type, with the direction of the illusory movement and with the biomechanical properties of their bearing muscles. Finally, kinesthetic illusions and AVRs can be effectively represented using similar vectorial computations. These strong relations between the perceptual and motor effects of tendon vibration once again suggest that the AVR may result from a perceptual-to-motor transformation of proprioceptive information, rather than from spinal reflex mechanisms.

Antagonist motor responses correlate with kinesthetic illusions induced by tendon vibration.

In humans, vibration applied to muscle tendons evokes illusory sensations of movement that are usually associated with an excitatory tonic response in muscles antagonistic to those vibrated (antagonist vibratory response or AVR). The aim of the present study was to investigate the neurophysiological mechanisms underlying such a motor response. For that purpose, we analyzed the relationships between the parameters of the tendon vibration (anatomical site and frequency) and those of the illusory movement perceived (direction and velocity), as well as the temporal, spatial, and quantitative characteristics of the corresponding AVRs (i.e., surface EMG, motor unit firing rates and activation latencies). Analogies were supposed between the characteristics of AVRs and voluntary contractions. The parameters of the AVR were thus compared with those of a voluntary contraction with similar temporal and mechanical characteristics, involving the same muscle groups as those activated by vibration. Wrist flexor muscles were vibrated either separately or simultaneously with wrist extensor muscles at frequencies between 30 and 80 Hz. The illusory movement sensations were quantified through contralateral hand-tracking movements. Electromyographic activity from the extensor carpi radialis muscles was recorded with surface and intramuscular microelectrodes. The results showed that vibration of the wrist flexor muscle group induced both a kinesthetic illusion of wrist extension and a motor response in the extensor carpi radialis muscles. Combined vibration of the two antagonistic muscle groups at the same frequency evoked neither kinesthetic illusion nor motor activity. In addition, vibrating the same two antagonistic muscle groups at different frequencies induced both a kinesthetic illusion and a motor response in the muscle vibrated at the lowest frequency. The surface EMG amplitude of the extensor carpi radialis as well as the motor unit activation latency and discharge frequency were clearly correlated to the parameters of the illusory movement evoked by the vibration. Indeed, the faster the illusory sensation of movement, the greater the surface EMG in these muscles during the AVRs and the sooner and the more intense the activation of the motor units of the wrist extensor muscles. Moreover, comparison of the AVR with voluntary contraction showed that all parameters were highly similar. Mainly slow motor units were recruited during the AVR and during its voluntary reproduction. That the AVR is observed only when a kinesthetic illusion is evoked, together with the similarities between voluntary contractions and AVRs, suggests that this vibration-induced motor response may result from a perceptual-to-motor transformation of proprioceptive information, rather than from spinal reflex mechanisms.

Motor cortex involvement during choice reaction time: a transcranial magnetic stimulation study in man.

It has been shown that transcranial magnetic stimulation can delay simple reaction time; this happens when the stimulation is delivered during the reaction time and over the cortical area which commands the prime mover of the required response. Although it is agreed that magnetic stimulation could be a useful tool for studying information processing in man, we argue that, because of the use of simple reaction time, the results reported so far are difficult to interpret within this theoretical framework. In the present paper, three experiments are reported. Six subjects participated in experiment 1 in which magnetic stimulation was delivered, at different times, during choice reaction time. The effects of the magnetic stimulation of the cortical area involved in the response (induced current passing forward over the motor representation of the responding hand), were compared to the effects of an electrical stimulation of the median nerve (H-reflex). In a first control experiment (experiment 2a; 5 subjects), the coil was placed over the ipsilateral motor cortex (induced current passing backward over the motor representation of the non-responding hand) thus minimizing the probability to excite the same neural nets as in the first experiment. In a second control experiment (experiment 2b; 4 subjects), the coil was placed a few centimeters away from the subject's scalp thus ensuring no stimulation of any neural nets. The results show that: (1) the noise and the smarting of the skin associated with the coil discharge produce an intersensory facilitation thereby shortening reaction time (experiment 2a), (2) actually, the noise produced by the stimulation is sufficient to produce such a facilitatory effect (experiment 2b), (3) a stimulation over the area at the origin of the motor command causes a reaction time delay which counteracts this intersensory facilitation effect (experiment 1), (4) in this latter case, the closer the stimulation to the actual overt response, the longer the delay and (5) there is no trace of correlation between the amplitude of the motor evoked potential and the reaction time change.

Suppression of voluntary motor activity revealed using transcranial magnetic stimulation of the motor cortex in man.

1. Suppression of voluntary muscle activity of hand and arm muscles in response to transcranial magnetic stimulation (TMS) of the motor cortex has been investigated in man. 2. Suppression could be elicited by low levels of TMS without any prior excitatory response. The latency of the suppression was 3-8 ms longer than the excitation observed at a higher stimulus intensity. The duration of the suppression ranged from 8 to 26 ms. 3. A circular stimulating coil was used to determine threshold intensity for excitation and suppression of contraction of thenar muscles in response to TMS at different locations over the motor cortex. The locations for lowest threshold excitation coincided with those for lowest threshold suppression. Suppression was elicited at a lower threshold than excitation at all locations. 4. A figure-of-eight stimulating coil was positioned over the left motor cortex at the lowest threshold point for excitation of the right thenar muscles. The orientation for the lowest threshold excitatory and inhibitory responses was the same for all subjects. That orientation induced a stimulating current travelling in an antero-medial direction. Suppression was invariably elicited at lower thresholds than excitation. 5. When antagonistic muscles (second and third dorsal interosseus) were co-contracted, TMS evoked coincident suppression of voluntary EMG in the two muscles without prior excitation of either muscle. This suggests that the suppression is not mediated via corticospinal activation of spinal interneurones. 6. Test responses to electrical stimulation of the cervical spinal cord were evoked in both relaxed and activated thenar muscles. In the relaxed muscle, prior TMS at an intensity that would suppress voluntary activity failed to influence the test responses, suggesting absence of inhibition at a spinal level. However, in the activated muscle, prior TMS could reduce the test response. This may be explained by disfacilitation of motoneurones due to inhibition of corticospinal output. 7. We propose that suppression of voluntary muscle activity by TMS is due in large part to activation of a mechanism within the motor cortex that reduces the corticospinal output to the muscle. It is concluded that TMS evokes excitation and inhibition via neuronal structures lying close to one another and having similar orientations.

Intensity to force translation: a new effect of stimulus-response compatibility revealed by analysis of response time and electromyographic activity of a prime mover.

In reaction time studies of stimulus-response compatibility, emphasis has been placed on the influence of spatial stimulus-response relationships, but what seems to be essential for the emergence of an effect of stimulus-response compatibility is the existence of a conceptual match between stimulus and response variables. This notion was at the origin of the present study to assess the compatibility relationship between the intensity of a visual stimulus and the force of a voluntary muscle contraction. A stimulus-response compatibility effect was demonstrated. This effect was entirely due to premotoric processes.

Effects of tonic vibration reflex on motor unit recruitment in human wrist extensor muscles.

Tonic vibration reflex was used to investigate the effects of muscle spindle Ia afferent activation on motor unit (MU) recruitment in human wrist extensor muscles. The MU force recruitment threshold recorded in the extensor carpi radialis muscles were quantitatively compared under two experimental situations: (1) during tonic isometric reflex contractions induced by mechanical tendon vibration and during voluntary contractions performed at the same velocity; (2) during two voluntary imposed ramp contractions (0.25 N.s-1) performed the one immediately before, and the other immediately after a tonic vibration reflex. In the first situation, it was observed that the Ia afferents activated by tendon vibration exerted a strong homonymous facilitatory action on their bearing muscles (extensor carpi radialis longus and brevis), while their heteronymous action on the synergistic muscle (extensor carpi ulnaris) was very weak. The MU recruitment thresholds in the extensor carpi radialis muscles were therefore significantly lower during the tonic reflex contraction than during the voluntary contraction. In the second situation, the tonic vibration reflex induced a facilitatory after-effect which decreased the MU recruitment thresholds during the subsequent voluntary imposed ramp contraction. It is suggested that this post-vibratory effect may have been due either to a postsynaptic potentiation of the motoneurones or to a reflex sensitization of the muscle spindles increasing their response to voluntary isometric contraction and consequently, increasing their facilitatory reflex action on the motoneurone pool.

Comparison of fluctuations of motor unit recruitment and de-recruitment thresholds in man.

Recruitment and de-recruitment thresholds of motor units in the wrist extensor muscles can undergo important random fluctuations, even when they are measured during stereotyped contractions and relaxations. These fluctuations were statistically quantified and compared. The statistical analysis indicated that recruitment and de-recruitment thresholds display the same kind of fluctuations, and that the successive measurements are randomly distributed following a quasi-normal law. We suggest that the notion of force threshold for motor unit recruitment and de-recruitment might be oversimplified and that a motor unit seems to have a range of force in which it can be recruited or de-recruited. Comparison of the mean values of recruitment and de-recruitment thresholds of the motor units in the extensor carpi radialis muscles showed that de-recruitment thresholds were significantly lower than recruitment thresholds. This difference in the thresholds, together with the difference in the motor unit discharge frequency during a contraction and a relaxation, suggests a differential control of the motoneurone activity during contractions and relaxations.

Differential activation of motor units in the wrist extensor muscles during the tonic vibration reflex in man.

1. Single motor unit activity was recorded in the extensor carpi radialis longus and extensor carpi radialis brevis muscles of five healthy human subjects, using metal microelectrodes. 2. Motor units were characterized on the basis of their twitch contraction times and their force recruitment thresholds during voluntary imposed-ramp contractions. 3. The discharge patterns of forty-three motor units were studied during tonic vibration reflex elicited by prolonged (150 s) trains of vibration (30 Hz) applied to the distal tendons of the muscles. The temporal relationships between the individual small tendon taps of the vibratory stimulus and the motor unit impulses were analysed on dot raster displays and post-stimulus time histograms. 4. After tendon taps, the impulses of motor units with long twitch contraction times (mean +/- S.D., 47.2 +/- 10.7 ms) and low recruitment thresholds (0.88 +/- 0.6 N) formed a single narrow peak (P1) with a latency (22.7 +/- 1.4 ms) which was comparable to that of the tendon jerk in the extensor carpi radialis muscles. These motor units were named 'P1 units'. On the other hand, the response of motor units with shorter twitch contraction times (31.1 +/- 3.3 ms) and higher recruitment thresholds (3.21 +/- 1.3 N) showed two peaks: a short latency (23.4 +/- 1.3 ms) P1 peak similar to the previous one and a P2 peak occurring 9.4 +/- 1.2 ms later. These motor units were named 'P1-P2 units'. 5. When the reflex contraction increased slowly, the P1 peaks of 'P1-P2 units' were clearly predominant at the beginning of the contraction, during the rising phase of the motor unit discharge frequency, while the P2 peaks became predominant when the units had reached their maximal discharge frequency. 6. Increasing the tendon vibration frequency (35, 55, 75, 95 Hz) did not modify the 'P1 unit' discharge pattern. Due to interference between vibration period and peak latencies, increasing the vibration frequency caused the P1 and P2 peaks of 'P1-P2 units' to overlap. 7. Superficial cutaneous stimulation of the dorsal side of the forearm during tendon vibration noticeably decreased the P1 peaks in both types of motor units. In the P2 peaks it could result in either a decrease or an increase but the average effect was a slight increase. 8. When applied 10 s before tendon vibration, cutaneous stimulation considerably suppressed the tonic vibration reflex.(ABSTRACT TRUNCATED AT 400 WORDS)

Fluctuations in motor unit recruitment threshold during slow isometric contractions of wrist extensor muscles in man.

Motor unit activity was recorded in the two extensor carpi radialis muscles with metal microelectrodes during isometric contractions. When tested at different 'free' increasing contraction velocities the recruitment thresholds (rt) systematically decreased, whereas when tested at different 'imposed' increasing velocities the rt exhibited great variability, with no observable overall tendency. Moreover, when measured at a given test velocity imposed periodically during MU investigation, rt also displayed great variability. These results suggest that the notion of rt tends to be oversimplified. The motoneurone pool excitability is highly dependent on both the experimental situation and the complexity of the motor task.

Physiological properties of the motor units of the wrist extensor muscles in man.

The physiological properties of 355 motor units (MUs) recorded in the extensor carpi radialis muscles were studied in 34 healthy human subjects during isometric contractions. MU selective twitches were educed from the whole muscle force using the spike-triggered averaging method. The twitch contraction times and twitch forces were measured. From these data it was attempted to estimate the distribution of fast and slow MUs in the muscles studied. Mu recruitment thresholds were systematically measured during stereotyped slow ramp contractions (force increase = 0.25 N. s-1). Degrees of correlation between contraction times, twitch forces and recruitment thresholds were pair analysed by computing simple regression curves and correlation coefficients. The degrees of correlation were compared between 245 Mus recorded in 34 subjects and 66 MUs recorded in a single subject. Analysis of the instantaneous discharge frequency of 132 MUs showed the existence of a remarkable degree of correlation (correlation coefficient, r = -0.75) between the "frequency rise times" (discharge onset to maximal frequency) and the MU twitch contraction times; i.e., the "frequency rise times" increase when the twitch contraction time decrease. The possibility that muscle contraction may be differentially modulated on the basis of this discharge property of the M Us is discussed. The results are compared to previous data and the limitations of the spike-triggered averaging method applied to long muscles in man are extensively discussed.