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PubMed2000

Characteristics of GABAergic neurons and their synaptic relationships with intrinsic axons in the cat motor cortex.

The intrinsic circuitry of the motor cortex comprises a complex network of connections whose synaptic relationships are poorly understood. This study was designed to determine the characteristics of subsets of GABAergic neurons containing the calcium-binding proteins parvalbumin (PV) and calbindin (CB), and their relationships with intrinsic axons in motor cortex. Immunohistochemically identified PV-containing neuronal profiles were more evenly distributed across cortical laminae (38% in II-III, 32% inV, 30% in VI) and more numerous (2.1/1) than CB-containing neuronal profiles (71% in II-III, 17% in V, 12% in VI). Relationships between neurons and axons intrinsic to motor cortex were visualized with fluorescent markers using the laser scanning confocal microscope. Similar percentages of PV (43%) and CB-immunoreactive (IR) (40%) neurons formed sparsely distributed appositions (1-5/neuron) with anterogradely labeled axons. The mean distances of such appositions from the somata were significantly different for the two groups (PV, mean = 22 microm, range = 1.6-93 microm; CB, mean = 32 microm, range = 6.2-132 microm). PV-IR neurons had a lower ratio of axosomatic/axodendritic appositions (1/99) compared with CB-IR neurons (14/86). Ultrastructural studies confirmed these findings. Fifty-seven percent of CB-IR neurons and 38% of PV-IR neurons formed synapses with intrinsic axons. Both populations received sparse input (1-6 synapses/neuron). Nearly all appositions between labeled terminals and postsynaptic profiles formed one synapse. Postsynaptic dendrites of PV-IR neurons (mean = 1.4 microm diameter) were larger than those of CB-IR neurons (mean = 1.1 microm), indicating more proximal synapses. Distinct input patterns of intrinsic axons to the two populations of neurons suggest unique roles in cortical processing.

Animals↗

PubMed1994

Corticocortical and thalamocortical responses of neurons in the monkey primary motor cortex and their relation to a trained motor task.

1. We studied the responsiveness of neurons in the primary motor cortex (MI) of monkeys (Macacafuscata) to electrical stimulation of the supplementary motor area (SMA), primary sensory cortex (SI), and the ventral subnucleus of the thalamus (VPLo) with chronically implanted electrodes. 2. All neurons examined in this study were characterized by their relation to a motor task performed by the animals. They responded to stimulation of the cortical or thalamic area with excitation from one area alone (n = 128) or from multiple areas (n = 84) of all combinations. In a majority of neurons, response latencies to both cortical and thalamic stimulation were within 5 ms. 3. A vast majority of neurons (80%) that were active during a preparatory period for forthcoming reaching movements were activated by SMA stimulation. They were activated only infrequently by SI or thalamic stimulation. 4. Movement-related neurons (active immediately before and during reaching movements) were activated by thalamic, SI, or SMA stimulation or by any combination of those stimuli. More than half of the movement-related neurons activated exclusively by either thalamic or SMA stimulation exhibited activity onset times earlier than those observed in the earliest muscles. By contrast, most movement-related neurons that responded only to SI stimulation were late in their activity onset. 5. These findings suggest that the SMA input to MI is important in developing a preparatory type of activity in MI, whereas the thalamus (VPLo) provides substantial inputs in movement execution. The roles played by inputs from SI and SMA in relation to motor execution are debatable and are discussed here with reference to previous reports.

Animals↗

PubMed1987

Peripheral nerve injury in developing rats reorganizes representation pattern in motor cortex.

We investigated the effect of neonatal nerve lesions on cerebral motor cortex organization by comparing the cortical motor representation of normal adult rats with adult rats that had one forelimb removed on the day of birth. Mapping of cerebral neocortex with electrical stimulation revealed an altered relationship between the motor cortex and the remaining muscles. Whereas distal forelimb movements are normally elicited at the lowest threshold in the motor cortex forelimb area, the same stimuli activated shoulder and trunk muscles in experimental animals. In addition, an expanded cortical representation of intact body parts was present and there was an absence of a distinct portion of motor cortex. These data demonstrate that representation patterns in motor cortex can be altered by peripheral nerve injury during development.

Afferent Pathways↗

PubMed1996

Supraspinal factors in human muscle fatigue: evidence for suboptimal output from the motor cortex.

1. Voluntary activation of elbow flexor muscles can be optimal during brief maximal voluntary contractions (MVCs), although central fatigue, a progressive decline in the ability to drive the muscle maximally, develops during sustained or repeated efforts. We stimulated the motor cortex and motor point in human subjects to investigate motor output during fatigue. 2. The increment in force (relative to the voluntary force) produced by stimulation of the motor point of biceps brachii increased during sustained isometric MVCs of the elbow flexors. Motoneuronal output became suboptimal during the contraction, i.e. central fatigue developed and accounted for a small but significant loss of maximal voluntary force. During 3 min MVCs, voluntary activation of biceps fell to an average of 90.7% from an average of > 99%. 3. The increment in force (relative to the voluntary force) produced by magnetic cortical stimulation was initially small (1.0%) but also increased during sustained MVCs to 9.8% (with a 2 min MVC). Thus, cortical output was not optimal at the time of stimulation nor were sites distal to the motor cortex already acting maximally. 4. A sphygmomanometer cuff around the upper arm blocked blood supply to brachioradialis near the end of a sustained MVC and throughout subsequent brief MVCs. Neither maximal voluntary force nor voluntary activation recovered during ischaemia after the sustained MVC. However, fatigue-induced changes in EMG responses to magnetic cortical stimulation recovered rapidly despite maintained ischaemia. 5. In conclusion, during sustained MVCs, voluntary activation becomes less than optimal so that force can be increased by stimulation of the motor cortex or the motor nerve. Complex changes in excitability of the motor cortex also occur with fatigue, but can be dissociated from the impairment of voluntary activation. We argue that inadequate neural drive effectively 'upstream' of the motor cortex must be one site involved in the genesis of central fatigue.

Adult↗

PubMed1988

Types and distribution of glutamic acid decarboxylase (GAD)-immunoreactive neurons in mouse motor cortex.

Neuronal structures in mouse motor cortex that contain gamma-aminobutyric acid (GABA), were identified by an immunocytochemical method, using an antiserum to glutamic acid decarboxylase (GAD). GAD-positive cell bodies occurred in all layers of the motor cortex, but were more concentrated in layers III and VI. GAD-positive puncta, presumably axon terminals, were also distributed throughout the cortical layers; a high density of puncta occurred in layer III, whereas a somewhat lower density characterized layer VI. Based on the shapes of their somata and dendritic trees we concluded that all GAD-positive cells were of the non-pyramidal type.

Animals↗

PubMed1997

Force and the motor cortex.

The relation between the activity of cells in the motor cortex and static force has been studied extensively. Most studies have concentrated on the relation to the magnitude of force; this relation is more or less monotonic. The slope of the relation, however, shows considerable variation among different studies and seems to be inversely associated with the range of forces over which the cell activity has been studied. Cells in the motor cortex also show variation in activity with the direction of static force. When both the direction and the magnitude of static force are allowed to vary, a majority of cells show significant changes in activity with direction of force alone, an intermediate number relate to both direction and magnitude, while a small number relate purely to the magnitude. This suggests that the direction of static force can be controlled independently of its magnitude and that this directional signal is especially prominent in the motor cortex. In general, it has been more difficult to study the relations to dynamic force. There is a correlation between motor cortex cell activity and the rate of change of force. The direction of dynamic force is also an important determinant of cell activity. When both static and dynamic force output are required (for example, with arm movement in the presence of gravity) it is the dynamic signal that is most clearly reflected in motor cortex activity. The relations between motor cortex activity and static or dynamic force are not invariant, but may be modified by the behavioral context of the motor output.

Animals↗

PubMed2004

Improvement of motor performance and modulation of cortical excitability by repetitive transcranial magnetic stimulation of the motor cortex in Parkinson's disease.

OBJECTIVE: To assess the effects of focal motor cortex stimulation on motor performance and cortical excitability in patients with Parkinson's disease (PD). METHODS: Repetitive transcranial magnetic stimulation (rTMS) was performed on the left motor cortical area corresponding to the right hand in 12 'off-drug' patients with PD. The effects of subthreshold rTMS applied at 0.5 Hz (600 pulses) or at 10 Hz (2000 pulses) using a 'real' or a 'sham' coil were compared to those obtained by a single dose of l-dopa. The assessment included a clinical evaluation by the Unified Parkinson's Disease Rating Scale and timed motor tasks, and a neurophysiological evaluation of cortical excitability by single- and paired-pulse TMS techniques. RESULTS: 'Real' rTMS at 10 or 0.5 Hz, but not 'sham' stimulation, improved motor performance. High-frequency rTMS decreased rigidity and bradykinesia in the upper limb contralateral to the stimulation, while low-frequency rTMS reduced upper limb rigidity bilaterally and improved walking. Concomitantly, 10 Hz rTMS increased intracortical facilitation, while 0.5 Hz rTMS restored intracortical inhibition. CONCLUSIONS: Low- and high-frequency rTMS of the primary motor cortex lead to significant but differential changes in patients with PD both on clinical and electrophysiological grounds. The effects on cortical excitability were opposite to previous observations made in healthy subjects, suggesting a reversed balance of cortical excitability in patients with PD compared to normals. However, the underlying mechanisms of these changes remain to determine, as well as the relationship with clinical presentation and response to l-dopa therapy. SIGNIFICANCE: The present study gives some clues to appraise the role of the primary motor cortex in PD. Clinical improvement induced by rTMS was too short-lasting to consider therapeutic application, but these results support the perspective of the primary motor cortex as a possible target for neuromodulation in PD.

Aged↗

PubMed2002

Increase in tibialis anterior motor cortex excitability following repetitive electrical stimulation of the common peroneal nerve.

The purpose of this study was to investigate whether repetitive electrical stimulation of the common peroneal nerve (CPN) is associated with changes in the motor response of the tibialis anterior (TA) muscle elicited by focal magnetic stimulation of the motor cortex. Motor evoked potentials (MEP) with a stimulation intensity of 125% of the threshold of the relaxed right TA were obtained before, during, and after repetitive electrical stimulation of the CPN (trains of five pulses of 1 ms, at a frequency of 200 Hz, repeated every second with a 30-min duration). The MEP of the TA muscle elicited after repetitive electrical stimulation were increased by 104% (range: 18-263%), and the increase was maintained for up to 110 min (range: 15-110 min) after the end of nerve stimulation. This increase in the MEP of the TA muscle was associated with a decrease in the threshold from the stimulation-response curve. Furthermore, during that period the early component of the TA stretch reflex as well as the latency of the MEP did not significantly change. To further test the origin of the increased MEP, complementary experiments showed that MEP elicited by transcranial electrical stimulation (TES) were also increased, but to a lesser degree (approximately 50%) than MEP elicited by TMS. It can be concluded that short-term nerve repetitive electrical stimulation of the lower extremities in healthy human participants can lead to a long-term increase in the contralateral MEP. As TES is believed to mainly activate the axon and not the soma of the cortical cells, the increased MEP cannot be explained exclusively by changes in the motor cortex cell excitability, but also by changes in subcortical neural structures involved in the excitation of spinal motoneurons. The results of this study allow the speculation that it would be possible to use repetitive electrical stimulation in the rehabilitation of patients with lower limb muscle weakness and spasticity.

Adult↗

PubMed2003

Level of action of cathodal DC polarisation induced inhibition of the human motor cortex.

OBJECTIVE: To induce prolonged motor cortical excitability reductions by transcranial direct current stimulation in the human. METHODS: Cathodal direct current stimulation was applied transcranially to the hand area of the human primary motor cortex from 5 to 9 min in separate sessions in twelve healthy subjects. Cortico-spinal excitability was tested by single pulse transcranial magnetic stimulation. Transcranial electrical stimulation and H-reflexes were used to learn about the origin of the excitability changes. Neurone specific enolase was measured before and after the stimulation to prove the safety of the stimulation protocol. RESULTS: Five and 7 min direct current stimulation resulted in motor cortical excitability reductions, which lasted for minutes after the end of stimulation, 9 min stimulation induced after-effects for up to an hour after the end of stimulation, as revealed by transcranial magnetic stimulation. Muscle evoked potentials elicited by transcranial electric stimulation and H-reflexes did not change. Neurone specific enolase concentrations remained stable throughout the experiments. CONCLUSIONS: Cathodal transcranial direct current stimulation is capable of inducing prolonged excitability reductions in the human motor cortex non-invasively. These changes are most probably localised intracortically.

Adult↗

PubMed2003

Subthreshold high-frequency TMS of human primary motor cortex modulates interconnected frontal motor areas as detected by interleaved fMRI-TMS.

To elucidate changes in human brain activity evoked by repetitive transcranial magnetic stimulation (rTMS), sub- and suprathreshold rTMS (4 Hz, 10 s) over the left primary sensorimotor cortex (M1/S1) was interleaved with blood-oxygenation-level-dependent (BOLD) echo-planar imaging of primary and secondary motor areas. Suprathreshold rTMS over left M1/S1 caused marked increases in BOLD signal in the stimulated area and SMA-proper in seven of eight subjects. By contrast, we found no change in BOLD signal in the stimulated M1/S1, when rTMS was given at intensities that were subthreshold for inducing motor responses in the contralateral hand. However, five of eight subjects showed consistent increases in BOLD MRI signal in the SMA-proper and, to a lesser extent, in bilateral lateral premotor cortex (LPMC) during subthreshold rTMS. A decrease in BOLD MRI signal was found in contralateral (right) M1/S1 in 6/8 subjects across all conditions. No significant changes were observed in the pre-SMA. The results support the notion that BOLD MRI responses to suprathreshold rTMS over M1/S1 are dominated by neuronal activity related to reafferent processing of TMS-induced hand movements. At subthreshold intensity, a short train of high-frequency rTMS seems to predominantly modulate activity of corticocortical connections which link the stimulated area with remote frontal premotor areas.

Adult↗

PubMed2005

Skilled reaching impairments from the lateral frontal cortex component of middle cerebral artery stroke: a qualitative and quantitative comparison to focal motor cortex lesions in rats.

The classical approach to investigating brain contributions to behavior has been to localize function to a region. In clinical investigations, however, injury is frequently multifocal, raising the question of how individual brain regions contribute to a resulting behavioral syndrome. For example, middle cerebral artery (MCA) ischemia in humans can concurrently damage a number of cortical and subcortical areas and the same areas are damaged in rat models of MCA stroke. In the rat, MCA occlusion produces severe motor deficits, but the cortical area of damage is the lateral neocortex, sparing motor cortex. This anatomical finding raises the question of whether the rat lateral neocortex contributes to MCA-related motor impairments, a question that was investigated in the present study. Rats received unilateral neocortical lesions via electrocoagulation of the MCA and were compared to rats with standard motor cortex lesions produced by devascaulrization of the overlaying blood vessels. The MCA group was as impaired as the motor cortex group in skilled reaching movements as assessed by quantitative measures of the contralateral-to-lesion forelimb in a single pellet task and in a tray-reaching task. Although there was improvement in success scores over a 2-week period in both groups, the groups were characterized by distinctive and enduring qualitative impairments. The motor cortex deficit was exemplified by use of trunk musculature and head movements to assist the reaching limb while the MCA impairment included sensory abnormalities. The results are discussed in relation to the contribution of lateral frontal cortex injury to MCA stroke sensorimotor syndromes.

Animals↗

PubMed2006

Evidence against a single coordinate system representation in the motor cortex.

Understanding the coordinate systems in which the motor cortical cells encode movement parameters such as direction is a fundamental yet unresolved issue. Although many studies have assumed that motor cortex encodes direction in an extrinsic, Cartesian (CA) coordinate system, other studies have provided evidence for encoding in intermediate coordinate systems such as a shoulder-centered (SC) or in a purely intrinsic, joint-angle-based (JA) coordinate frame. By simultaneously recording from multiple single units in primary motor cortex, we examined movement direction encoding under each of these three coordinate systems. We directly compared the degree of directional tuning invariance over multiple sub-regions in the workspace. We also compared the mutual information between neuronal firing rate and movement direction in the three systems. We observed a broad range of directional invariance in all three coordinate systems with no strong dominance of any single coordinate system. The mutual information analyses corroborated this observation. However, we found a small but significant bias toward the SC coordinate frame, which was also supported by population vector decoding. Similar results were found when we compared hand/torque force direction encoding in all three coordinate systems. These results suggest that the motor cortex employs a coordinate system that is yet to be discovered or perhaps that the motor cortex should not be viewed as a substrate for any coordinate system representation.

Action Potentials↗

PubMed2004

Improvements in the signal-to-noise ratio of motor cortex cells distinguish early versus late phases of motor skill learning.

There are numerous experience-driven changes in cortical circuitry that correlate with improved performance. Improved motor performance on a reach-to-grasp task in rodents is associated with changes in long-term potentiation (LTP), synaptogenesis, and movement representations in primary motor cortex (M1) by training days 3, 7, and 10, respectively. We recorded single-cell activity patterns in M1 during reach-to-grasp training to test how neural-spiking properties change with respect to LTP, synaptogenesis, and motor map changes. We also tested how neural-spiking changes relate directly to improved performance by monitoring muscle activity patterns. We found that signal-to-noise ratios (SNRs) of M1 spiking were significantly improved with practice but only after 7-12 d. Three sources of noise were assessed: signal-dependent noise exemplified by the slope of the relationship between mean spike count and count variance per burst, signal-independent noise exemplified by the offset of this relationship, and background firing rates before and after bursts. Signal-independent noise and pre-burst firing rates were reduced with practice. Early performance gains (days 1-6) were dissociated from SNR improvements, whereas later performance gains (day 7-12) were related directly to the magnitude of improvement in both muscle recruitment reliability and success rates. With training, an increased number of cells exhibited firing rates that were correlated with muscle recruitment patterns, with lags suggesting a primary direction of influence from M1 to muscles. These results suggest a functional linkage from local synaptogenesis in M1 to improved spiking reliability of M1 cells to more reliable recruitment of muscles and finally to improved behavioral performance.

Action Potentials↗

PubMed1997

Force and the motor cortex.

Animals↗

PubMed1999

Automatic activation in the human primary motor cortex synchronized with movement preparation.

The human primary motor cortex during a unilateral finger reactive movement to visual stimuli was examined by magnetoencephalography (MEG) measurement. The brain activity related to movement execution (the motor activity contralateral to the movement side) was estimated based on movement onset conditions and reaction times. The movement onset conditions were: (1) a simple reaction time task with a visual stimulus, (2) a Go/NoGo task with different colored stimuli and (3) a Go/NoGo task with different position stimuli. Dipole source estimation was done, and the time course of the motor activity was calculated. The results showed that not only the visual response but also the contralateral motor activity was evoked by the stimulus in all cases, and even when the NoGo stimulus was given. The motor activity in the primary motor cortex was conjectured to consist of two dominant components: the first component for the movement preparation and the second component for the movement execution. Because the first component happened with a constant delay time from the stimulus even in the NoGo case, the first component, coming through a fast pathway for signals from visual stimulus processing to the motor cortex without any intervening cognitive processing, was conjectured to make the motor cortex prepare for the forthcoming movement onset automatically regardless of the stimulus instruction.

Adult↗

PubMed1991

Evidence for a contribution of the motor cortex to the long-latency stretch reflex of the human thumb.

1. In normal subjects, transcranial magnetic stimulation of the hand region of the motor cortex evokes motor responses only in contralateral hand muscles at a latency of about 19-24 ms. In contrast, stimulation of the motor cortex of three mirror movement subjects evoked, nearly simultaneously, motor responses in hand muscles on both sides of the body at latencies similar to those of normal subjects. In these subjects no other neuroanatomical pathways appear to be abnormally directed across the mid-line. Thus, their mirror movements are probably due to a projection of the corticospinal tract to homologous motoneurone pools on each side of the body. 2. We reasoned that if the motor cortex contributes to the generation of long-latency stretch reflex responses then in these mirror movement subjects stretching a muscle on one side of the body should produce long-latency reflex responses in the ipsilateral and the homologous contralateral muscle. 3. To test this idea experiments were done on normal human subjects and on the subjects with mirror movements. The electromyographic (EMG) activity of the flexor pollicis longus muscle (FPL) on each side of the body was recorded. Stretch of the distal phalanx of the thumb of one hand produced a series of distinct reflex EMG responses in the ipsilateral FPL. The earliest response, when present, began at 25 ms (S.D. = 3.5 ms) and was followed by responses at 40 (S.D. = 3.9 ms) and 56 ms (S.D. = 4.3 ms). There was no difference, either in timing or intensity, between the ipsilateral FPL EMG responses of normal subjects and those of the mirror movement subjects. 4. No response of any kind was observed in the contralateral (unstretched) FPL of normal subjects. In contrast, we observed in all three mirror movement subjects EMG responses in the contralateral (unstretched) FPL beginning at 45-50 ms. The latency of this response is considerably shorter than the fastest voluntary kinaesthetic reaction time, which was on average 130 ms (S.D. = 11 ms). The contralateral long-latency EMG response observed in the mirror movement subjects was on average 30% (range 5-60%) of that on the ipsilateral side. No short-latency response (25 ms) was ever observed in the contralateral FPL of these subjects. 5. These observations are quite consistent with the idea that the long-latency stretch reflex responses of hand and finger muscles are produced, at least in part, by the motor cortex.

Electromyography↗

PubMedDate not supplied

The effects of high-frequency microstimulation of the cortex on interhemisphere synchronization in the rat motor cortex.

Studies were carried out on long-term changes in the synchronization of neuronal activity in networks including callosal cells of the opposite hemispheres evoked by high-frequency microstimulation in the motor cortex of anesthetized rats. The level of synchronization was assessed in terms of the amplitude and width of peaks located symmetrically on cross-correlograms relative to the coordinate origin. Tetanization predominantly decreased synchronization in a group of initially background-active neurons, while there was a significant number of synchronously firing neurons in a group of cells which became activated. "Super-narrow" peaks appeared in interhemisphere interactions. There was a correlation between the type of modification of "narrow" (<20 msec) and "intermediate" (30-80 msec) peaks and changes in the efficiencies of mono- and polysynaptic connections.

Animals↗

PubMed2006

Reliability of motor cortex transcranial magnetic stimulation in four muscle representations.

OBJECTIVE: Motor cortex plasticity may underlie motor recovery after stroke. Numerous studies have used transcranial magnetic stimulation (TMS) to investigate motor system plasticity. However, research on the reliability of TMS measures of motor cortex organization and excitability is limited. We sought to test the reliability of these TMS measurements. METHODS: Twenty healthy volunteers were tested twice over a two-week period using TMS to determine motor threshold, map topography, and stimulus-response curves for first dorsal interosseous (FDI), abductor pollicis brevis (APB), extensor digitorum communis (EDC), and flexor carpi radialis (FCR) muscles. RESULTS: We found moderate to good test-retest reliability TMS measurements of motor threshold (ICC=0.90-0.97), map area (ICC=0.63-0.86) and location (ICC=0.69-0.86), and stimulus-response curves (ICC=0.60-0.83). CONCLUSIONS: TMS assessments of motor representation size, location, and excitability are generally reliable measures, although their reliability may vary according to the muscle under investigation. SIGNIFICANCE: These results suggest that TMS measurements of motor cortex function are reliable enough to be potentially useful in investigation of motor system plasticity.

Adult↗