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Oscillatory cortical activity during a motor task in a deafferented patient.

Little is known about the influence of the afferent peripheral feedback on the sensorimotor cortex activation. To answer this open question we investigated the alpha and beta band task-related spectral power decreases (TRPow) in the deafferented patient G.L. and compared the results to those of six healthy subjects. The patient has been deafferented up to the nose for 24 years but her motor fibers are unaffected and she can perform complex motor tasks under visual control. We recorded EEG (58 scalp positions) as well as the exerted force during a visuomotor task. The subjects had to maintain in precision grip an isometric force at 15% of the maximal voluntary contraction. In the patient we found a significantly higher alpha band spectral power during rest and larger alpha TRPow decreases during the motor task when compared to the healthy subjects' data. In contrast, we did not observe any significant differences between patient and controls for the beta band TRPow. The results indicate an altered functional alpha band network state in the patient, probably due to the chronic deafferentation leading to a deep 'idling' state of the contralateral sensorimotor area.

Adult↗

Glutamate receptor expression in rat striatum: effect of deafferentation.

The cerebral cortex is the primary source of glutamatergic afferents to the neostriatum. We used in situ hybridization to examine the effect of removal of the glutamatergic input to the striatum by unilateral frontal cortical ablation on the expression of genes encoding subunits from three families of glutamate receptors: N-methyl-D-aspartate receptors (NMDAR1, NMDAR2A, and NMDAR2B); alpha-amino-3-hydroxy-5-methyl-4-isoxazolepropionate (AMPA) receptors (GluR1-4, flip and flop splice variants); and metabotropic receptors (mGluR1-5). Significant changes were restricted to the dorsolateral quadrant of the ipsilateral striatum, the main projection area of the sensorimotor cortex. The expression of those messages which are normally abundant, NMDAR1, NMDAR2A, GluR1-4 flop and mGluR1, 3 and 5, was decreased in the deafferented dorsolateral striatum by 10-39% at 3 days after cortical ablation and subsequently increased to 120-165% of control at 15 and 60 days. mRNAs encoding the flip isoforms of GluR1-4, mGluR2 and 4, and an alternatively spliced region of NMDAR1 (Insertion I) which are undetectable or present at low levels in the striatum were not induced by cortical ablation. In contrast, both glial fibrillary acid protein and beta-actin mRNA expression were markedly enhanced at 3 and 15 days, returning to near normal at 60 days. Striatal NMDA, AMPA and metabotropic type 1 ligand binding sites were increased as early as 3 days after cortical ablation, reached a peak at 15 days and remained increased for up to 60 days, while metabotropic type 2 binding was slightly but significantly reduced at 3 and 15 days and [3H]kainate binding did not change significantly. These results demonstrate that cortical ablation, and subsequent loss of glutamatergic afferents to the striatum, results in alterations in the expression of genes encoding glutamate receptor subunits in striatal neurons. The regulation of these genes appears to be coordinate, so that the relative abundance of the different messages is preserved.

Animals↗

Hemispheric asymmetry in supplementary motor area connectivity during unilateral finger movements.

Studies of unilateral finger movement in right-handed subjects have shown asymmetrical patterns of activation in primary motor cortex. Some studies have measured a similar asymmetry in the supplementary motor area (SMA), but others have not. To shed more light on the symmetry of function in the SMA, we used path analysis of functional MRI data to investigate effective connectivity during a unilateral finger movement task. We observed a slight asymmetry in task activation: left SMA was equally active during movement of either hand, while right SMA was more active for left-hand movement, suggesting a dominant role of left SMA. In addition, we tested for a corresponding asymmetry in the influence of SMA on sensorimotor cortex (SMC) using a path model based on the well-established principle that SMA is involved in motor control and SMC in execution. We observed that the influence of left SMA on left SMC increased during right-hand movement, and the influence of left SMA on right SMC increased during left-hand movement. However, there was no significant hand-dependent change in the influences of the right SMA. This asymmetry in connectivity implies that left SMA does play a dominant role in unilateral movements of either hand in right handers. The experiment also provides a basis for further studies of motor system connectivity in healthy or patient populations.

Adult↗

Cortical activation during human volitional swallowing: an event-related fMRI study.

Functional magnetic resonance imaging (fMRI) provides a safe, noninvasive method for studying task-related cortical neuronal activity. Because the cerebral cortex is strongly implicated in the control of human swallowing, we sought to identify its functional neuroanatomy using fMRI. In 10 healthy volunteers, a swallow event-related paradigm was performed by injecting 5 ml water bolus into the oral cavity every 30 s. Whole brain functional magnetic susceptibility -weighted spiral imaging data were simultaneously acquired over 600 s on a 1.5-T magnetic resonance scanner, utilizing the blood oxygenation level-dependent technique, and correlation maps were generated using both >99% percentile rank and spatial extent thresholding. We observed areas of increased signal change consistently in caudal sensorimotor cortex, anterior insula, premotor cortex, frontal operculum, anterior cingulate and prefrontal cortex, anterolateral and posterior parietal cortex, and precuneus and superiomedial temporal cortex. Less consistent activations were also seen in posterior cingulate cortex and putamen and caudate nuclei. Activations were bilateral, but almost every region, particularly the premotor, insular, and frontal opercular cortices, displayed lateralization to one or the other hemisphere. Swallow-related cortical activity is multidimensional, recruiting brain areas implicated in processing motor, sensory, and attention/affective aspects of the task.

Adult↗

High-resolution CMR(O2) mapping in rat cortex: a multiparametric approach to calibration of BOLD image contrast at 7 Tesla.

The blood oxygenation level-dependent (BOLD) functional magnetic resonance imaging (fMRI) method, which is sensitive to vascular paramagnetic deoxyhemoglobin, is dependent on regional values of cerebral metabolic rate of oxygen utilization (CMR(O2)), blood flow (CBF), and volume (CBV). Induced changes in deoxyhemoglobin function as an endogenous contrast agent, which in turn affects the transverse relaxation rates of tissue water that can be measured by gradient-echo and spin-echo sequences in BOLD fMRI. The purpose here was to define the quantitative relation between BOLD signal change and underlying physiologic parameters. To this end, magnetic resonance imaging and spectroscopy methods were used to measure CBF, CMR(O2), CBV, and relaxation rates (with gradient-echo and spin-echo sequences) at 7 Tesla in rat sensorimotor cortex, where cerebral activity was altered pharmacologically within the autoregulatory range. The changes in tissue transverse relaxation rates were negatively and linearly correlated with changes in CBF, CMR(O2), and CBV. The multiparametric measurements revealed that CBF and CMR(O2) are the dominant physiologic parameters that modulate the BOLD fMRI signal, where the ratios of (deltaCMR(O2)/CMR(O2)/(deltaCBF/ CBF) and (deltaCBV/CBV)/(deltaCBF/CBF) were 0.86 +/- 0.02 and 0.03 +/- 0.02, respectively. The calibrated BOLD signals (spatial resolution of 48 microL) from gradient-echo and spin-echo sequences were used to predict changes in CMR(O2) using measured changes in CBF, CBV, and transverse relaxation rates. The excellent agreement between measured and predicted values for changes in CMR(O2) provides experimental support of the current theory of the BOLD phenomenon. In gradient-echo sequences, BOLD contrast is affected by reversible processes such as static inhomogeneities and slow diffusion, whereas in spin-echo sequences these effects are refocused and are mainly altered by extravascular spin diffusion. This study provides steps by which multiparametric MRI measurements can be used to obtain high-spatial resolution CMR(O2) maps.

Animals↗

Visual cortex activation in kinesthetic guidance of reaching.

The purpose of this research was to determine the cortical circuit involved in encoding and controlling kinesthetically guided reaching movements. We used (15)O-butanol positron emission tomography in ten blindfolded able-bodied volunteers in a factorial experiment in which arm (left/right) used to encode target location and to reach back to the remembered location and hemispace of target location (left/right side of midsagittal plane) varied systematically. During encoding of a target the experimenter guided the hand to touch the index fingertip to an external target and then returned the hand to the start location. After a short delay the subject voluntarily moved the same hand back to the remembered target location. SPM99 analysis of the PET data contrasting left versus right hand reaching showed increased (P < 0.05, corrected) neural activity in the sensorimotor cortex, premotor cortex and posterior parietal lobule (PPL) contralateral to the moving hand. Additional neural activation was observed in prefrontal cortex and visual association areas of occipital and parietal lobes contralateral and ipsilateral to the reaching hand. There was no statistically significant effect of target location in left versus right hemispace nor was there an interaction of hand and hemispace effects. Structural equation modeling showed that parietal lobe visual association areas contributed to kinesthetic processing by both hands but occipital lobe visual areas contributed only during dominant hand kinesthetic processing. This visual processing may also involve visualization of kinesthetically guided target location and use of the same network employed to guide reaches to visual targets when reaching to kinesthetic targets. The present work clearly demonstrates a network for kinesthetic processing that includes higher visual processing areas in the PPL for both upper limbs and processing in occipital lobe visual areas for the dominant limb.

Adult↗

Varieties of paw and digit movement during spontaneous food handling in rats: postures, bimanual coordination, preferences, and the effect of forelimb cortex lesions.

This study describes how rats use their paws and digits when handling a wide range of foodstuffs, including food pellets, grapes, sunflower seeds, shelled and unshelled peanuts, and different sized pastas, etc. Analysis of videorecordings show that the rats display digit postures that include variations in the spacing of the digits, differences in the relative use of different digits, and interlimb differences in paw and digit posture. The rats also display limb preferences in that one paw is used in a supporting function while the other rotates, flips, or pushes the food as is required by the shape of the item. There is a significant correlation between the paw used for manipulation and food items of similar shape but no correlation between the limb used for manipulation and that used for skilled reaching. Small unilateral lesions to the forepaw area of somatic sensorimotor cortex produced impairments in use of the paw contralateral to the lesions. These results: (1) reveal a surprising complexity in the way in which rats use their paws and digits in manipulating food; (2) show that rats have limb preferences in spontaneous food handling; and (3) show that manipulatory dexterity is dependent upon the integrity of the forelimb area of motor cortex. The results are discussed in relation to the evolution of motor skill, the use of rats for investigating questions of motor system organization, neural plasticity, and recovery of function after brain damage.

Animals↗

An aberrant crossed visual corticotectal pathway in albino rats.

This study investigates the dynamic nature of the developing corticotectal pathway arising in the visual cortex. Special attention is given to the interaction occurring between the corticotectal pathways of each side of the brain and between corticotectal and retinotectal terminations. Normally the visual cortex of rats projects only to the ipsilateral superior colliculus. If one visual cortex is removed at birth, the remaining visual cortex subsequently shows a bilateral projection to the superior colliculus. The aberrant corticotectal pathway is heavier if the cortical ablation is accompanied by eye removal at birth but eye enucleation alone is not a sufficient stimulus for production of a crossed corticotectal projection. The aberrant crossed pathway shows a topographic order which appears to correspond to that of the normal ipsilateral corticotectal pathway. The pathway differs from the aberrant projections from the retina in that lesions done as late as 20 days postnatal still result in an aberrant crossed corticotectal pathway. This is similar to the aberrant crossed cortical projections from sensorimotor cortex. The pathway would appear to arise as a result of lack of competition from corticotectal axons normally present contralaterally or from attraction of denervated corticotectal sites. While denervated retinotectal sites stimulate sprouting of the corticotectal axons once in the deafferented colliculus, they do not stimulate crossing of the corticotectal projection.

Age Factors↗

Spinal cord plasticity in response to unilateral inhibition of the rat motor cortex during development: changes to gene expression, muscle afferents and the ipsilateral corticospinal projection.

In developing Wistar albino rats, ventral horn muscle afferent boutons are lost following corticospinal innervation. Motor cortex lesions rescue a proportion of these boutons and perturb activity dependent expression of cJun and parvalbumin (PV) in the spinal cord. Therefore, we tested whether activity-dependent competition between corticospinal and proprioreceptive afferents determines the balance of these inputs to motor output pathways by delivering the inhibitory GABA agonist muscimol unilaterally to the forelimb motor cortex using slow release polymer implants from postnatal day 7 (P7) coincident with corticospinal synaptogenesis. Controls received saline. Inhibition of immature cortical neurons by muscimol was confirmed with separate in vitro electrophysiological recordings. After P28, spinal cord sections were immunostained for PV, cJun and muscle afferents transganglionically labelled with cholera toxin-B (CTB). Unilateral inhibition reduced contralaterally the number of PV positive spinal cord neurons and muscle afferent boutons in the dorsolateral ventral horn, compared to controls, and significantly altered the distribution of motoneuronal cJun expression. Separately, descending tracts were retrogradely traced with CTB from the cervical hemicord contralateral to implants. Forelimb sensorimotor cortex sections were immunostained for either CTB or PV. In muscimol treated animals, significantly fewer neurons expressed PV in the inhibited hemicortex, but as many CTB labelled corticospinal neurons were present as in controls, along with an equally large corticospinal projection from contralateral to the implant, significantly greater than in controls. Unexpectedly, unilateral inhibition of the motor cortical input did not lead to an expanded muscle afferent input. Instead, this was reduced coincident with development of a bilateral corticospinal innervation.

Age Factors↗

Early exclusive use of the affected forelimb after moderate transient focal ischemia in rats : functional and anatomic outcome.

BACKGROUND AND PURPOSE: Previous work by researchers in our laboratory has shown that in the rat, the exclusive use of the affected forelimb during an early critical period exaggerates lesion volume and retards functional recovery after electrolytic lesions of the forelimb sensorimotor cortex. In the present study, we examined the effects of exclusive use of the affected forelimb after middle cerebral artery occlusion (MCAO). METHODS: Ischemia of moderate severity was produced in male Long-Evans rats through 45 minutes of occlusion of the left middle cerebral and both common carotid arteries. Exclusive use of either the affected or unaffected forelimb was forced through immobilization of either the ipsilateral (MCAO+ipsi) or contralateral (MCAO+contra) forelimb, respectively, for 10 days in a plaster cast, or the animal was left uncasted (MCAO+nocast). Sham surgeries were performed, and animals were also casted for 10 days or left uncasted. Sensorimotor testing was performed during days 17 to 38. At the end of sensorimotor testing, cognitive performance was tested with use of the Morris water maze. In a separate experiment, temperatures and corticosterone levels were measured during the 10-day period after 45-minute ischemia and casting. RESULTS: The MCAO+ipsi group performed worse on sensorimotor tasks than the MCAO+contra, MCAO+nocast, and sham groups. Infarct volume was significantly larger in the MCAO+ipsi group than in the sham and MCAO+contra groups but not in the MCAO+nocast group. No group differences were found with the Morris water maze, and no group differences were found in either temperature or plasma corticosterone level. CONCLUSIONS: The exclusive use of the affected forelimb immediately after focal ischemia has detrimental effects on sensorimotor function that cannot be attributed to hyperthermia or stress.

Animals↗

Cortical reorganization of hand motor function to primary sensory cortex in hemiparetic patients with a primary motor cortex infarct.

OBJECTIVE: To show cortical reorganization in hemiparetic patients with a primary motor cortex (M1) infarct including the precentral knob by using functional magnetic resonance imaging (fMRI). DESIGN: Case-control. SETTING: Outpatient clinics in the rehabilitation department of a university hospital. PARTICIPANTS: Two stroke patients and 20 control subjects. INTERVENTIONS: By using fMRI, we evaluated the hand motor function of 2 hemiparetic stroke patients, who had made some recovery from complete paralysis of the affected hand, and 20 control subjects. MAIN OUTCOME MEASURES: fMRI was performed by using the blood oxygen level-dependent technique at 1.5 T with a standard head coil. The motor task paradigm consisted of hand grasp-release movements. RESULTS: The contralateral primary sensorimotor cortex was activated by the hand movements of the control subjects and of the unaffected side of the 2 patients. Only the contralateral (infarct side) primary sensory cortex (S1) was activated by the movements of the affected hand of the 2 patients, a result that was not observed in the control subjects or with the unaffected hand in the stroke patients. CONCLUSIONS: The hand motor function associated with the infarcted M1 in our patients was reorganized into the S1. These results suggest cortical reorganization in patients with an M1 infarct.

Adult↗

Mu and beta rhythm topographies during motor imagery and actual movements.

People can learn to control the 8-12 Hz mu rhythm and/or the 18-25 Hz beta rhythm in the EEG recorded over sensorimotor cortex and use it to control a cursor on a video screen. Subjects often report using motor imagery to control cursor movement, particularly early in training. We compared in untrained subjects the EEG topographies associated with actual hand movement to those associated with imagined hand movement. Sixty-four EEG channels were recorded while each of 33 adults moved left- or right-hand or imagined doing so. Frequency-specific differences between movement or imagery and rest, and between right- and left-hand movement or imagery, were evaluated by scalp topographies of voltage and r spectra, and principal component analysis. Both movement and imagery were associated with mu and beta rhythm desynchronization. The mu topographies showed bilateral foci of desynchronization over sensorimotor cortices, while the beta topographies showed peak desynchronization over the vertex. Both mu and beta rhythm left/right differences showed bilateral central foci that were stronger on the right side. The independence of mu and beta rhythms was demonstrated by differences for movement and imagery for the subjects as a group and by principal components analysis. The results indicated that the effects of imagery were not simply an attenuated version of the effects of movement. They supply evidence that motor imagery could play an important role in EEG-based communication, and suggest that mu and beta rhythms might provide independent control signals.

Adolescent↗

Early appearance of functional deficits after neonatal excitotoxic and hypoxic-ischemic injury: fragile recovery after development and role of the NMDA receptor.

We sought to determine whether neonatal rats that sustain unilateral cerebral hypoxic-ischemic or excitotoxic insults (1) manifest contralateral sensorimotor deficits during development or in adulthood and (2) recover from those deficits. Seven-day-old (P7) rats received a right intrastriatal injection of the glutamate analog N-methyl-D-aspartate (NMDA). Unilateral hypoxia-ischemia (HI) was induced by right carotid ligation followed by 1.5 h in 8% O2. Both procedures produce neuronal loss in the striatum and sensorimotor cortex. Nonlesioned controls were included. We scored percent forepaw placement on the edge of a horizontal surface, with lateral vibrissa stimulation, from P9 to P19, and at P33 and P50. Then, on P50, rats were treated with the NMDA antagonist MK-801 to determine whether deficits could be reinstated. NMDA- and HI-lesioned rats exhibited a deficit in contralateral vibrissa-stimulated forepaw placing that emerged during the second week of life. Yet, by P33 and P50, the lesioned groups and controls were indistinguishable. MK-801 injection on P50 resulted in transient reinstatement of the placing deficit. After unilateral neonatal excitotoxic or HI brain injury, contralateral sensorimotor deficits are detected, but in many animals, these deficits have resolved by adulthood. Thus, timing of sensorimotor tests may influence their sensitivity for detection of focal neuropathology originating in the neonatal period.

Animals↗

[Regional cerebral blood flow in limbic seizures induced by microinjection of kainic acid into amygdala in cats].

The relationship between neuronal epileptic activity and regional cerebral blood flow was studied by means of hydrogen clearance method (rCBF). Measurements of rCBF in the limbic structure and cerebral cortex were performed during limbic seizures induced by a microinjection of kainic acid to the left amygdala under concurrent monitoring of polygraph. Pentobarbital (35 mg/kg, i.p.) anesthesia was induced and cats were artificially ventilated. Physiological parameters such as blood pressure, body temperature, PaO2, PaCO2 and HCO3 were kept stable. After fixation of the cat's head in a stereotaxic device, stainless screw electrodes were placed over bilateral anterior sigmoid gyrus so as to touch the dura mater for cortical EEG monitoring. Bipolar needle electrodes were stereotaxically placed to both amygdala and the left dorsal hippocampus. Platinum electrodes were stereotaxically placed to the left amygdala, dorsal hippocampus and sensorimotor cortex. After measurements of control rCBF, a microsyringe was stereotaxically inserted into the left amygdala and kainic acid solution (2 micrograms) was injected. Polygraphic monitoring followed. Five to 30 minuted after kainic acid injection, continuous multiple spikes appeared in the left amygdala. Forty to 120 minutes later, continuous multiple spikes transmitted to the left hippocampus. Two to five hours later, limbic seizure occurred. The limbic seizure lasted five to eight hours and interictal discharges appeared. Repeated measurements of rCBF were done. In the left amygdala, rCBF increased about two-fold of the control immediately after development of continuous multiple spikes and remained increased as long as the seizure persisted.(ABSTRACT TRUNCATED AT 250 WORDS)

Amygdala↗

Representation of nociceptive stimuli in primary sensory cortex.

Using fMRI, we observed cortical activity associated with nociceptive hot and cold sensations applied to hand and foot that are not spatially restricted to the corresponding regions of the primary somatosensory cortex (SI). Hot (55-57 degrees C) and cold (0-2 degrees C) tactile stimuli were applied separately to the right hand and foot of eight right-handed subjects. Although somatotopic mapping of hand and foot was observed as expected based on the Penfield homunculus, activations associated with hot during both hand and foot stimulation and subsequently, cold, activated regions unique to each thermal modality irrespective of the body part. This distributed system for thermal information is present at both nociceptive and more neutral thermal intensities (i.e. warm and cool sensations) indicating the presence of distributed sensory processing associated with thermal-related sensations in human primary sensorimotor cortex.

Adult↗

Effects of NGF, b-FGF, and cerebrolysin on water maze performance and on motor activity of rats: short- and long-term study.

The effects of 14-day treatments with nerve growth factor (NGF), basic fibroblast growth factor (b-FGF), or the peptidergic drug Cerebrolysin on postlesion acquisition of a water maze task and on motor activity were evaluated. Rats were tested in the Morris water maze 14 days (early test) and 7 to 8 months (delayed test) after a bilateral lesion of the frontoparietal (sensorimotor) cortex. Only the rats treated with Cerebrolysin performed the water maze task at the level of the nonlesioned controls in the early test. No short-term effect of NGF (6.5 ng/14 days; 38 ng/ml) or b-FGF (17 ng/14 days; 100 ng/ml) treatment was found. The delayed test revealed that water maze performance was restored in rats treated with b-FGF in comparison with intact controls. The data showed that b-FGF can support or initiate processes in the CNS that lead to a delayed functional amelioration and/or compensation for a water maze performance deficit. NGF did not influence the acquisition impairment caused by a sensorimotor cortical lesion. Two-week administration of Cerebrolysin had a time-dependent influence: it attenuated the acquisition deficit and increased the motor activity of rats, both effects declined to the level of lesioned controls within 8 months.

Aging↗

Frontal cortex and the programming of repetitive tapping movements in man: lesion effects and functional neuroimaging.

Two studies examined the contribution of human frontal cortical areas to the programming of trains of repetitive movements. The first study compared the performance of patients with unilateral frontal excisions, unilateral temporal excisions and controls on the speed of initiation of discrete vs. sequential tapping movements to visual stimuli. The frontal group showed normal initiation times in single taps and a normal execution (pace and accuracy) in sequential taps but they were slower than the other groups at initiating sequential taps indicating a sequence programming problem for repetitions of a single response. A second study examined the functional anatomy of single and sequential taps in eight control subjects using fMRI. Subjects performed flexion/extension movements of the right thumb at either 1 movement/s or as trains of four closely spaced movements at a rate of 1 train/4 s. Statistical analyses revealed that primary sensorimotor cortex and a dorsolateral premotor cortex region were activated in both conditions. Medial frontal activation was not significant in discrete movements but was clearly present in sequential movements and involved SMA and cingulate regions bilaterally. In addition, two other dorsolateral premotor foci of activation were observed in the sequential taps condition. Results from these two experiments converge toward establishing a significant role of dorsolateral and medial premotor regions in the programming of trains of repetitive responses.

Adult↗

Inhibitory actions of motor cortex following unilateral brain lesions as studied by magnetic brain stimulation.

Noninvasive transcranial magnetic stimulation (TMS) of the motor cortex not only induces short-latency, motor-evoked potentials (MEP) in contralateral muscles, but also inhibitory phenomena. One type of inhibitory action appears directly after the MEP in contralateral muscles and can be visualized by blockade of tonic, voluntary electromyographic (EMG) activity (postexcitatory inhibition, PI). Evidence for a cortical origin of PI, especially in its later part, was derived from double cortical stimulation in previous studies and is further supported by examination of PI in patients with focal hemispheric unilateral brain lesions in the present study. Thirty patients with different sites of vascular or tumour lesions were studied by TMS. In 6 patients with circumscribed lesions of the primary sensorimotor cortex a significant shortening of PI to contralateral muscles was observed. In 7 patients with focal lesions of the thalamus or internal capsule, in 6 patients with lesions of the premotor cortex and in 5 patients with lesions restricted to the parietal or temporal lobe, a significant prolongation of PI to the contralateral muscles was detected. Six patients with transient ischemic attacks showed either prolongation or shortening of PI. We conclude that PI is predominantly generated in the primary motor cortex, correspondingly its damage causes shortening of PI. In contrast, damage to brain areas that project to the primary motor cortex is followed by prolongation of PI. This remote effect on the primary motor cortex may result from disinhibition of cortical interneurones.

Adult↗