Parveen Bawa

Bilateral responses of upper limb muscles to transcranial magnetic stimulation in human subjects

Abstract.Anatomical and behavioural work on primates has shown bilateral innervation of axial and proximal limb muscles, and contralateral control of distal limb muscles. The following study examined if a clear boundary exists between the distal and proximal upper limb muscles that are controlled contralaterally or bilaterally. The right motor cortical area representing the upper limb was stimulated, while surface EMG was recorded bilaterally from various upper limb muscles during rest and phasic voluntary contractions. Peak-to-peak amplitude of motor evoked potential (MEP) was measured for each muscle on both sides. The ratio R = (ipsilateral MEP: contralateral MEP) was calculated for seven pairs of muscles. For each of the seven pairs, R was less than 1.0, implying that for each muscle and subject, the contralateral control is stronger. The boundary where R changed from almost zero to a clearly measurable magnitude depended on the subject. Ipsilateral MEPs from trapezius and pectoralis could be recorded with a small background contraction from almost all subjects; on the other hand, in deltoid and biceps brachii, ipsilateral MEPs were observed only with bimanual phasic contractions. The forearm and hand muscles, in general, did not show any ipsilateral MEPs. Major differences between subjects lay in the presence or the absence of ipsilateral MEPs in biceps brachii and deltoid, without defining a sharp boundary between proximal and distal muscles.

Motor unit recruitment during lengthening contractions of human wrist flexors

The purpose of this study was to revisit the question of recruitment of motor units during lengthening contractions because of conflicting views in the literature on this subject. Motor unit activity was recorded from the flexor carpi radialis muscle of four human subjects to compare the patterns of recruitment during lengthening and isometric contractions. Lengthening contractions were produced either when the subject voluntarily stopped opposing a background load or when an additional load was imposed on the already contracting muscle. In both cases, lengthening of the active muscle was produced at a variety of speeds, from quite slow to “as fast as possible.” No differences in recruitment order were observed between isometric and lengthening contractions at any speed of lengthening contraction. It is concluded that all contractions in normal humans recruit motor units in an orderly fashion from small to large, according to the size principle of motor unit recruitment.

Comparison of the depression of H-reflexes following previous activation in upper and lower limb muscles in human subjects

Abstract When conditioning-testing (C-T) stimuli are applied to Ia afferents to elicit H-reflexes, the test reflex is abolished immediately following the conditioning reflex. As the C-T interval is increased, the test response slowly begins to recover, taking several hundred milliseconds to attain control values. The time course of this recovery is known as the H-reflex recovery curve. H- reflex recovery curves were compared using surface EMG and single motor unit activities in lower limb soleus and upper limb flexor carpi radialis (FCR) muscles in seven healthy human subjects. Under rest conditions, the recovery of H-reflexes and single motor unit activity was slow for soleus; the recovery was not complete even in 1 s. In comparison, the recovery was very fast for FCR motor units, occurring in 200–300 ms. The effects of rate of stimulation (0.1–10.0 imp/s) were also examined on the magnitude of H-reflex responses. The reflex response declined with increasing rate of stimulation, the decline being slightly greater in soleus than in FCR. When these phenomena were examined with voluntary facilitation of the spinal cord, the time of recovery shortened and the effect of stimulus rate also diminished. Changes with background facilitation were greater in FCR than in soleus. The differences between the two muscles are attributed mainly to differences in presynaptic inhibition in the two spinal segments, and/or to the differences in dynamics of the transmitter release in terminals of Ia afferents synapsing with slow soleus motoneurons and those synapsing with the fast FCR motoneurons.

Electromyographic activity, H-reflex modulation and corticospinal input to forearm motoneurones during active and passive rhythmic movements

Four subjects produced coordinated movements, consisting of flexion and extension of the wrist in ipsilateral (right wrist only), contralateral (left wrist only), inphase (both wrists in flexion or both in extension) and antiphase (one wrist in flexion, the other in extension) conditions. Electromyographic (EMG) activity was recorded from right wrist flexor and extensor muscles. In one session, transcranial magnetic stimuli (TMS) of the left motor cortex, around threshold intensity, evoked short-latency responses in the right wrist extensors and flexors. In another session, the median nerve at the cubital fossa was stimulated to elicit an H-reflex in the right flexor carpi radialis (rFCR). A movement cycle was divided into 8 segments. In total, 10 identical stimuli were delivered during each segment in each condition, at two movement frequencies. The magnitude of the EMG reponses to TMS was modulated markedly during movements made in the ipsilateral condition, and in both bimanual conditions. EMG activity was greater, and motor-evoked potentials (MEPs) were larger in the antiphase condition than in the inphase condition. When the amplitudes of the MEPs were normalised with respect to background EMG, no significant differences between the bimanual conditions were obtained. For H-reflexes, significant differences between the two bimanual conditions were observed, suggesting differences in levels of excitability of the Ia afferent pathway. These differences were attributed to segmental input associated with changes in muscle length arising from limb movement, and upon descending input to the spinal cord, possibly mediated by Renshaw cell inhibition. During rhythmic passive movement of the right limb, H-reflexes were inhibited and MEPs potentiated in a cyclic fashion. Passive movement of the contralateral left limb resulted in inhibition of both responses

Control of the wrist joint in humans.

Abstract As one considers changes in motor activity from lower mammals to higher primates, one of the major changes one observes lies in the cortical control of forelimb muscles. There has been a shift from disynaptic control of spinal motoneurons in, for example, the cat, to a greater and greater percentage of monosynaptic control of hand and forelimb motoneurons in the primate. In spite of the species and evolutionary changes in the synaptic connections of the corticospinal tract, it appears that the interneurons identified in the cat are retained in the monkey and human. These interneurons, under the influence of descending pathways, modulate the output of motoneuron pools. Perhaps the control of these interneurons has also changed towards finer control of movement, as has been suggested by recent studies in the monkey. Whether in cat or human, the recruitment pattern for motor units is the same; the change from disynaptic to monosynaptic connections has not changed the recruitment pattern of muscles. Differences in the recruitment patterns of muscles may lie in the finer control of inputs to motoneurons in the primate. This review seeks to integrate the current knowledge of the mechanisms involved in the motor control of the wrist joint and especially in the recruitment patterns of the muscles. These motor control mechanisms include the biomechanics of the wrist joint, recruitment patterns of wrist muscles, interneurons and spinal cord circuits in the cervical regions mediating the output of spinal motoneurons, and the supraspinal control of these muscles.

Neural development in children: A neurophysiological study☆

In adult human subjects torque motor imposed angular displacements of the upper limb joints result in two major reflex EMG components as identified by their latency. The shorter latency (SL) component is probably the spinal stretch reflex and the longer latency (LL) reflex component may involve supraspinal structures. Tendon jerk, arising via the spinal reflex pathway, is present in children at birth. In this study the presence and the development of the LL component in wrist flexors was investigated in children between the ages of 2 and 13 years. Kinaesthetic reaction times were tested simultaneously. In young children, the duration of the LL component was much longer than that in adults. The duration decreased slowly from 2 to 6 years, following which a relatively abrupt decrease between the ages of 6 and 8 years took place. After 8 years of age responses looked more adult-like. Kinaesthetic reaction times attained adult-like values after 10 years of age. Various possibilities underlying these observations are discussed.

Chapter 19 Do Lengthening Contractions Represent a Case of Reversal in Recruitment Order

Publisher Summary The central nervous system (CNS) recruits spinal motoneurons in an orderly fashion under most of the tested experimental paradigms. Experimental observations suggesting selective excitation/recruitment of larger motor units during the electrical stimulation of cutaneous nerves, electrical stimulation of the rubrospinal tract, and lengthening contractions have been reported. The results utilizing synchronous electrical stimulation of cutaneous nerves, or the red nucleus, have to be interpreted as nonphysiological. Instead, under physiological activation of these pathways, the preferential inhibition of small motoneurons by these pathways would change the slope of the recruitment curve instead of causing selective recruitment of large motoneurons. The implications and mechanisms for the selective recruitment reported during lengthening contractions have to be reconsidered. This chapter presents the data obtained with two different experimental paradigms, both of which result in lengthening of active muscles. These observations do not support selective recruitment of fast twitch motor units during lengthening contractions. Two main papers where single motor unit data have been produced and interpreted to suggest selective recruitment of large motor units are central to the discussion in the chapter.

Motor Unit Rotation in a Variety of Human Muscles

The phenomena of substitution and rotation among motor units of a muscle were examined in seven different muscles. Intramuscular motor unit activity and surface electromyographic (EMG) activity were recorded from one of the following muscles: abductor digiti minimi, first dorsal interosseous, extensor digitorum communis, flexor and extensor carpi radialis, tibialis anterior, and soleus. The subject was asked to discharge a discernible unit at a comfortable constant or rhythmically (pseudosinusoidally) modulated rate with audio and visual feedback. Results are reported from a total of 42 sets of motor units from all seven muscles. We observed that when a subject fired a motor unit for a long period, an additional motor unit frequently started to discharge after a few minutes. When the subject was asked to keep activity down to one unit, very often it was Unit 1 that dropped and Unit 2 continued to fire. Whereas Unit 2 had fired for a few minutes, Unit 1 resumed firing without any conscious effort by the subject. If the subject was then asked to retain just one unit, it was Unit 2 that dropped. Rhythmic modulation of firing rate of a tonically firing unit showed that whereas the threshold of this unit increased, the threshold of a phasically discharging unit decreased substantially. The increase in threshold of a tonically discharging unit is suggested to arise from inactivation of Na(+) and Ca(2+) channels and the decrease in threshold of higher-threshold units is suggested to arise from an increase in persistent inward currents that may occur during prolonged contractions. Whether a unit stops or starts to fire is suggested to depend on a balance between the strength of the central motor command, persistent inward currents, and inactivation of voltage-gated channels. Such rotations among low-threshold motoneurons would ensure low-level sustained contractions to be viable not only in small hand muscles but also in larger limb muscles.

Contribution of joint and cutaneous afferents to longer-latency reflexes in man

Electromyographic records from wrist extensors and flexors show a short latency reflex response, M1 and a longer latency response comprising of M2 and M3 peaks. M1 corresponds to the spinal stretch reflex and hence mediated by spindle afferents. In order to determine the contribution of various afferent types to M2-M3 components and simple reaction times, reflexes were elicited before and after anaesthetic blocks of palm cutaneous and wrist joint afferents in human subjects. The results show that joint and cutaneous afferents have no significant contribution to the longer latency reflexes or simple reaction times.

Recruitment order of motor units during the stretch reflex in man

The recruitment pattern of single motor units contributing to the short- and long-latency periods of the stretch reflex was investigated in human adults. When non-tonic, a given unit responded to stretch during the long-latency reflex period. When made tonic, the same unit showed a reflex response during the short-latency period, while a higher threshold, non-tonic unit would respond during the long-latency period. The data show an orderly recruitment of motor units during both phases of the stretch reflex in man.