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PubMed1995

Afferent connections of the medial frontal cortex of the rat. II. Cortical and subcortical afferents.

In order to compare the frontal cortex of rat and macaque monkey, cortical and subcortical afferents to subdivisions of the medial frontal cortex (MFC) in the rat were analyzed with fluorescent retrograde tracers. In addition to afferent inputs common to the whole MFC, each subdivision of the MFC has a specific pattern of afferent connections. The dorsally situated precentral medial area (PrCm) was the only area to receive inputs from the somatosensory cortex. The specific pattern of afferents common to the ventrally situated prelimbic (PL) and infralimbic (IL) areas included projections from the agranular insular cortex, the entorhinal and piriform cortices, the CA1-CA2 fields of the hippocampus, the subiculum, the endopiriform nucleus, the amygdalopiriform transition, the amygdalohippocampal area, the lateral tegmentum, and the parabrachial nucleus. In all these structures, the number of retrogradely labeled cells was larger when the injection site was located in area IL. The dorsal part of the anterior cingulate area (ACd) seemed to be connectionally intermediate between the adjacent areas PrCm and PL; it receives neither the somatosensory inputs characteristic of area PrCm nor the afferents characteristic of areas PL and IL, with the exception of the afferents from the caudal part of the retrosplenial cortex. A comparison of the pattern of afferent and efferent connections of the rat MFC with the pattern of macaque prefrontal cortex suggests that PrCm and ACd areas share some properties with the macaque premotor cortex, whereas PL and IL areas may have characteristics in common with the cingulate or with medial areas 24, 25, and 32 and with orbital areas 12, 13, and 14 of macaques.

Afferent Pathways↗

PubMed2005

Early social environment interferes with the development of NADPH-diaphorase-reactive neurons in the rodent orbital prefrontal cortex.

The influence of early parental deprivation on the development of NADPH-diaphorase-(NO-synthase) reactive neuron numbers in subregions of the orbital prefrontal cortex (ventrolateral orbital, lateral orbital, and agranular insular cortex) was quantitatively investigated in the precocious lagomorph Octodon degus. Forty-five-day-old degus from three groups were compared: (1) repeated parental separation: degus that were repeatedly separated from their parents during the first three postnatal weeks and thereafter raised in undisturbed social conditions; (2) chronic isolation: degus that were raised under undisturbed social conditions until postnatal day 21, and then were reared in chronic social isolation; and (3) control: degus that were reared undisturbed in their families. Compared to the control animals the ventrolateral orbital prefrontal cortex and agranular insular cortex of the two deprived groups displayed significantly decreased density of NADPH-diaphorase-reactive neurons (down to 62% in the ventrolateral orbital prefrontal cortex of males, 70% in the agranular insular cortex, and in the lateral orbital prefrontal cortex 80% in both genders). These results confirm that early changes of social environment interferes with the development of limbic circuits, which might determine normal or pathological behaviors in later life.

Animals↗

PubMed1978

Projection from area 3a to the motor cortex by neurons activated from group I muscle afferents.

Two receiving areas in the pericruciate cortex are known for inputs from group I muscle afferents of forelimb nerves. One focus is near the postcruciate dimple of area 3a, and the other in the lateral sigmoid gyrus of the motor cortex (area 4gamma). The cortico-cortical projection of area 3a to 4gamma, and the relay by this projection of group I muscle afferent input to the motor cortex were investigated in cats. The following results were obtained. 1. Seventy-four neurons within area 3a were antidromically activated by intracortical microstimulation of the motor cortex. 2. Although excitation evoked by stimulation of group I muscle afferents could be demonstrated for only a few (8 of 48) cortico-cortical neurons in extracellular recordings, due to the methodological limitations discussed, this input evoked EPSPs in 8 of 9 cortico-cortical neurons recorded intracellularly. Therefore, it is likely that the majority of neurons projecting from area 3a to the motor cortex have an excitatory synaptic input from group I afferents. 3. Neurons projecting from area 3a to the motor cortex were most commonly found in cortical layer III, although some were found in layer V. 4. Five of nine pyramidal tract neurons of area 3a had a strong excitatory synaptic input from group I muscle afferents. 5. A new type of pyramidal tract neuron was found which has cortico-cortical axon collaterals connecting the two cytoarchitectonic regions. These various neurons may be part of a feedback system from muscle afferents to the motor cortex.

Afferent Pathways↗

PubMed1994

Are extent and force independent movement parameters? Preparation- and movement-related neuronal activity in the monkey cortex.

Movement extent and movement force can be independently controlled in motor performance. Therefore, independent representations of extent and force should exist in the central nervous system (CNS). To test this hypothesis, microelectrode recordings were made in sensorimotor cortex of monkeys trained to perform visually cued wrist flexion movements of two extents, against two levels of frictional resistance. An initial preparatory signal (PS) provided complete, partial or no information about extent and/or force of the movement, which had to be performed in response to a second, response signal (RS). The activity of 511 neurons of the primary motor cortex (MI), the premotor cortex (PM), the postcentral cortex (PC), and the posterior parietal cortex (PA) was recorded in two monkeys. Both reaction time (RT) and neuronal data suggest that there exist independent neuronal mechanisms responsible for the programming of either parameter. On the one hand, partial information about either movement parameter shortened RT when compared with the condition of no prior information. On the other hand, there were, among others, two discrete populations of neurons, one related only to extent, the other only to force. Preparatory changes in activity related to either movement parameter were mainly located in the frontal cortex, especially in the PM. After occurrence of the RS, the percentage of selective changes in activity increased and tended to extend to the parietal cortex. In particular during the movement, force-related changes in activity have been encountered in PA. Furthermore, we conducted trial-by-trial correlation analyses between RT and preparatory neuronal activity for all conditions of prior information. The mean correlation coefficient was significantly higher in the condition of information about movement extent than of information about movement force and it was significantly higher in MI/PM than in PC/PA.

Animals↗

PubMed2003

Lewy body densities in the entorhinal and anterior cingulate cortex predict cognitive deficits in Parkinson's disease.

Previous studies reported an association between cortical Lewy body (LB) formation and dementia in Parkinson's disease (PD). However, it is unclear whether cognitive decline in this disorder is related to specific patterns of LB distribution within the cerebral cortex. Moreover, the prediction of cognitive status based on concomitant assessment of LB and Alzheimer's disease lesions has led to conflicting results. We performed a clinicopathological study in 22 elderly PD patients in whom parkinsonism preceded cognitive decline by at least 3 years. Cognitive status was assessed prospectively using the clinical dementia rating scale (CDR); quantitative assessment of LB, neurofibrillary tangles (NFT), and senile plaques (SP) was performed in Brodmann areas 9, 21, 24, 40 and the entorhinal cortex. Statistical analysis was performed using both correlation coefficients and logistic regression models. There was a highly significant correlation between CDR scores and regional LB scores in the entorhinal cortex and area 24. LB and SP densities in the entorhinal cortex accounted for 36.2% and 19.3% of the variability in CDR scores. LB densities in area 24 could explain 25.2% of this variability. NFT densities did not predict cognitive status. In multivariate models only LB densities in the entorhinal cortex and anterior cingulate cortex were significantly associated with CDR scores. These results imply that an assessment of LB pathology limited to the entorhinal cortex and area 24 may be sufficient to predict cognition in PD. They also suggest that LB formation in limbic areas may be crucial for the development of PD dementia.

Age Distribution↗

PubMed1991

Differential alterations of second messenger systems and cerebral glucose use following excitotoxic lesion of rat cerebral cortex.

Quantitative autoradiography of [3H]forskolin and [3H]phorbol 12,13 dibutyrate (PDBu) binding was examined 21 days after unilateral lesioning of the rat visual cortex using ibotenic acid. In the same animals, the functional deficit was assessed using quantitative [14C]-2-deoxyglucose autoradiography. [3H]Forskolin binding was significantly reduced in each layer of the lesioned visual cortex by at least 40% compared to the contralateral hemisphere. Significant reductions in [3H]forskolin binding were observed in the superior colliculus (-15%) and dorsal lateral geniculate body (-12%) ipsilateral to the lesioned cortex. [3H]PDBu binding was significantly reduced in the lesioned visual cortex (layers V-VI) by 34%, compared with the control hemisphere. There were no significant alterations in [3H]DPBu binding in any other brain regions. Following ibotenate-induced lesioning of the visual cortex, glucose use was significantly reduced throughout the lesioned cortex by at least 25% with minor alterations in glucose use in the ipsilateral dorsal lateral geniculate body and superior colliculus. The present study highlights the relative robustness of [3H]PDBu binding compared to [3H]forskolin binding after excitotoxic damage to the cerebral cortex and suggests that [3H]forskolin binding sites are present on cortical efferent fibres.

Animals↗

PubMed1991

Time course of interictal EEG patterns induced by a penicillin injection into the olfactory cortex.

In awake rats, the time courses of behavioural and cortical interictal hyperactivity patterns were investigated following an injection of the GABA antagonist Na-penicillin (PCN) (125 IU/0.5 microliters) into the olfactory or motor cortex. The cortical EEG was recorded by means of 6 AgAgCl-electrodes, behaviour was videotaped simultaneously. Behavioural hyperactivity developed immediately after PCN injection. It lasted longer than 2.5 to 4 h in the olfactory but less than 30 min in the motor cortex group of rats. The interictal EEG pattern of the olfactory group was characterized by a slow establishing of three centers of high cortical activity. They were located in the ipsilateral central and the contralateral frontal and occipital cortex. In the motor cortex group, however, a cortical center of high activity developed immediately after the PCN injection near the injection site only, or additionally, over the homotopic area of the contralateral hemisphere. The results indicate different susceptibility properties in the underlying neuronal networks. Ongoing epileptiform activity obviously modifies this susceptibility in a site-specific manner. Moreover, the time-correlated occurrence of high activity in the frontal motor and occipital cortex evoked by a PCN injection into the olfactory cortex suggests a close coupling of these three areas. A coupling between the frontal motor, occipital and focal area could not be shown, if PCN was injected into the motor cortex.

Animals↗

PubMed2004

Increased facilitation of the primary motor cortex following 1 Hz repetitive transcranial magnetic stimulation of the contralateral cerebellum in normal humans.

Connections between the cerebellum and the contralateral motor cortex are dense and important, but their physiological significance is difficult to measure in humans. We have studied a group of 10 healthy subjects to test whether a modulation of the excitability of the left cerebellum can affect the excitability of the contralateral motor cortex. We used repetitive transcranial magnetic stimulation (rTMS) at 1 Hz frequency to transiently depress the excitability of the left cerebellar cortex and paired-pulse TMS testing of intracortical inhibition (ICI) and intracortical facilitation (ICF) to probe the excitability of cortico-cortical connections in the right motor cortex. The cortical silent period was also measured before and after cerebellar rTMS. Motor evoked potentials (MEPs) were significantly larger after than before conditioning rTMS trains (p < 0.01). Moreover, left cerebellar rTMS increased the ICF of the right motor cortex as measured with paired-pulses separated by an interstimulus interval (ISI) of 15 ms. The effect lasted for up to 30 min afterward and was specific for the contralateral (right) motor cortex. The cortical silent period was unaffected by cerebellar rTMS. The implication is that rTMS of the cerebellar cortex can shape the flowing of inhibition from Purkinje cells toward deep nuclei, thereby increasing the excitability of interconnected brain areas.

Adult↗

PubMed2006

Oxygenation in the motor cortex during exhaustive pinching exercise.

The purpose of this study was to examine the effect of fatigue resulting from exhaustive pinching exercise on frontal and motor cortex activity. Eight healthy subjects (four male and four female) participated in the present study. All subjects performed at 70% of maximal voluntary contraction (MVC) pinching exercise until reaching a state of volitional fatigue. Frontal cortex and motor cortex oxygenation was measured by near-infrared spectroscopy throughout the exhaustive exercise. Frontal cortex and motor cortex oxygenation increased significantly at the 90 and 120 s after the start of exercise compared with the pre-exercise values and these decreased with the time passage. Frontal cortex oxygenation at exhaustion was significantly lower than the 90 and 120 s after the start of exercise, while motor oxygenation at exhaustion was the same value with the pre-exercise value. These findings suggest that the exhaustive exercise induces the decrease of cerebral function and that the fatigue resulting from dynamic exercise decreases the motor cortex activity.

Adult↗

PubMed1986

The organization of the rat motor cortex: a microstimulation mapping study.

In conclusion, the rat primary motor cortex appears to be organized into irregularly shaped patches of cortex devoted to particular movements. The location of major subdivisions such as the forelimb or hindlimb areas is somatotopic and is consistent from animal to animal, but the internal organization of the pattern of movements represented within major subdivisions varies significantly between animals. The motor cortex includes both agranular primary motor cortex (AgL) and, in addition, a significant amount of the bordering granular somatic sensory cortex (Gr(SI)), as well as the rostral portion of the taste sensory insular or claustrocortex (Cl). The rat frontal cortex also contains a second, rostral motor representation of the forelimb, trunk and hindlimb, which is somatotopically organized and may be the rat's supplementary motor area. Both of these motor representations give rise to direct corticospinal projections, some of which may make monosynaptic connections with cervical enlargement motoneurons. Medial to the primary motor cortex, in cytoarchitectonic field AgM, is what appears to be part of the rat's frontal eye fields, a region which also includes the vibrissae motor representation. The somatic motor cortical output organization pattern in the rat is remarkably similar to that seen in the primate, whose primary, supplementary and frontal eye field cortical motor regions have been extensively studied.

Animals↗

PubMed1986

Adult rat motor cortex connections to thalamus following neonatal and juvenile frontal cortical lesions: WGA-HRP and amino acid studies.

Frontal cortex was removed in 1- and 30-day-old rats. When both groups reached 90 days of age, the forelimb motor/sensory cortex in the unlesioned hemisphere was injected with wheat germ agglutinin-horseradish peroxidase (WGA-HRP) or tritiated leucine. Thalamic neurons were retrogradely labeled only ipsilateral to the WGA-HRP injection site in both neonatally and juvenile-lesioned subjects. Ventrolateral (VL), ventromedial (VM), centromedial (CM), centrolateral (CL), parafascicular (PF), posteromedial (POm), and posterior (PO) thalamic nuclei were labeled. This and the demonstration of only ipsilateral thalamocortical connections at birth helped explain the marked thalamic atrophy which developed ipsilateral to neonatal frontal cortex lesions. Death of thalamic neurons after neonatal removal of their normal cortical target could be due to their failure to sprout into the opposite cortex because that cortex was already innervated by the opposite thalamus at birth. Leucine motor/sensory cortex injections in both neonatally and juvenile-lesioned subjects labeled the ipsilateral VL, VM, CM, CL, PF, POm, and PO thalamic nuclei; contralateral CM, CL, and PF thalamic nuclei; ipsilateral medial, ventral, and lateral pontine nuclei; and parts of the contralateral pontine nuclei. The ipsilateral connections were always more robust than the contralateral connections. The contralateral corticothalamic and corticopontine projections, however, were much more numerous and widespread in neonatally compared to juvenile-lesioned subjects. The greater sparing of some motor functions said to occur in neonatal compared to adult motor cortex-lesioned subjects could be due to the plasticity of corticothalamic, corticopontine, and other corticofugal pathways, but not to the plasticity of thalamocortical pathways.

Age Factors↗

PubMed1999

Motor cortex modulation of exteroceptive information at bulbar and thalamic lemniscal relays in the cat.

We have studied the control of the primary motor cortex on the activity of lemniscal neurons in semi-chronic implanted cats. In each experiment, two to three foci in the primary motor cortex were identified by intracortical microstimulation at low threshold (up to 20 microA) for their capacity to evoke movements of contralateral single limb joints. Neurons belonging to the dorsal column nuclei (main cuneate nucleus and gracile nucleus), or to the ventral posterolateral nucleus, were sampled for their response to stimulation of the peripheral cutaneous fields, as well as the antidromic response to stimulation of the contralateral medial lemniscus and ipsilateral somatosensory cortex, respectively. These neurons were then tested for stimulation of the cortical foci using a current intensity equal to the threshold needed to evoke motor effects, although we reduced the duration of the stimulating trains; thus, we avoided evoking movements which could elicit afferent volleys along the somatosensory paths. It was found that the primary motor cortex was able to modulate the transmission of exteroceptive signals at the level of both dorsal column nuclei and ventral posterolateral nucleus with analogous modalities. In particular: (i) a high percentage of responses, with a prevalence of excitatory effects, was observed when the receptive field of the neurons topographically corresponded to, or was very close to, the joint controlled by a given cortical focus; (ii) in these cases, higher percentages of excitations were observed in tests which concerned the distal segments of limbs than the proximal segments; (iii) the percentage of responses became lower as the neuronal receptive field was located further from the cortical motor target, the pattern being more frequently inhibitory in nature. From a functional point of view, the motor cortex control appears to be organized in a very precise manner. Its excitatory nature might subserve integrative mechanisms by which exteroceptive information arising in a given limb segment would be enhanced by a motor command inducing movements of the same body part. Moreover, a better definition of the afferent input could be obtained by a simultaneous depression of neurons, which send towards the cortex signals from adjacent or more distant cutaneous regions. It can be hypothesized that such an organization of the cortical control could improve the discriminative somatosensory aspects during the execution of explorative movements, besides supplying a sharper cutaneous feedback to the motor cortex.

Animals↗

PubMed1999

Spontaneous activity of the perirhinal cortex in behaving cats.

The perirhinal cortex lies at the interface between the neocortex and allocortex. Whether the perirhinal cortex expresses spontaneous electroencephalographic rhythms that are characteristic of the allocortex and/or of the neocortex is unknown. Thus, the present investigation was undertaken to characterize the activity of the perirhinal cortex with respect to various electroencephalographic rhythms that are displayed by neocortical areas or the entorhino-hippocampal system during different behavioral states of vigilance in chronically-implanted cats. Although perirhinal and neocortical electroencephalograms underwent similar state-dependent changes in amplitude, the ubiquitous neocortical sleep spindles were absent from the perirhinal cortex. In addition, while the slow sleep oscillation (0.5-1 Hz), which is pervasive in the neocortex, was present in the perirhinal cortex, its temporal relation to the neocortical oscillation was highly variable. In contrast, a high degree of correlation was found between perirhinal and entorhinal electroencephalographic activities in all behavioral states. In particular, during waking and paradoxical sleep, multiple simultaneously recorded entorhinal and perirhinal sites displayed an oscillation in the theta range which was highly correlated. To rule out the possibility that the perirhinal theta oscillation reflected volume conduction from neighboring structures, single-unit recordings were performed. Spike-triggered averages and peri-event histograms revealed that perirhinal cells displayed a statistically significant theta-related modulation of their spontaneous activity, albeit weaker than that observed in the entorhinal cortex. Thus, from the standpoint of spontaneous electroencephalographic rhythms, the perirhinal cortex is more closely related to the entorhino-hippocampal system than to the neocortex.

Animals↗

PubMed2005

The organization and connections of anterior and posterior parietal cortex in titi monkeys: do New World monkeys have an area 2?

We used multiunit electrophysiological recording techniques to examine the topographic organization of somatosensory area 3b and cortex posterior to area 3b, including area 1 and the presumptive area 5, in the New World titi monkey, Callicebus moloch. We also examined the ipsilateral and contralateral connections of these fields, as well as those in a region of cortex that appeared to be similar to both area 7b and the anterior intraparietal area (7b/AIP) described in macaque monkeys. All data were combined with architectonic analysis to generate comprehensive reconstructions. These studies led to several observations. First, area 1 in titi monkeys is not as precisely organized in terms of topographic order and receptive field size as is area 1 in macaque monkeys and a few New World monkeys. Second, cortex caudal to area 1 in titi monkeys is dominated by the representation of the hand and forelimb, and contains neurons that are often responsive to visual stimulation as well as somatic stimulation. This organization is more like area 5 described in macaque monkeys than like area 2. Third, ipsilateral and contralateral cortical connections become more broadly distributed away from area 3b towards the posterior parietal cortex. Specifically, area 3b has a relatively restricted pattern of connectivity with adjacent somatosensory fields 3a, 1, S2 and PV; area 1 has more broadly distributed connections than area 3b; and the presumptive areas 5 and 7b/AIP have highly diverse connections, including connections with motor and premotor cortex, extrastriate visual areas, auditory areas and somatosensory areas of the lateral sulcus. Fourth, the hand representation of the presumptive area 5 has dense callosal connections. Our results, together with previous studies in other primates, suggest that anterior parietal cortex has expanded in some primate lineages, perhaps in relation to manual abilities, and that the region of cortex we term area 5 is involved in integrating somatic inputs with the motor system and across hemispheres. Such connections could form the substrate for intentional reaching, grasping and intermanual transfer of information necessary for bilateral coordination of the hands.

Animals↗

PubMed1969

The characteristics of acetylcholine release mechanisms in the auditory cortex.

1. The characteristics of the acetylcholine (ACh) release mechanism have been studied in the auditory cortex of rabbits on stimulation of the ipsilateral medial geniculate nucleus.2. On stimulation of the medial geniculate nucleus the mean release of ACh from the auditory receiving cortex was 6.1 times the spontaneous release; the mean release from other parts of the cortex was 2.2 times the spontaneous release.3. The frequency of stimulation most effective in evoking ACh release was found to be 10/sec.4. Both the spontaneous and evoked release of ACh were reduced by 40-65% in the absence of calcium from the solution bathing the auditory cortex, and increased by 15-25% when the calcium concentration in the bathing solution was doubled.5. The presence of low concentrations of magnesium in the fluid bathing the cortex was essential for the optimal release of ACh, but high magnesium concentrations lowered this release.6. The presence of triethylcholine (TEC) in the fluid bathing the auditory cortex reduced both the spontaneous and evoked release of ACh. This reduction was reversed in the presence of choline.7. The effects of calcium, magnesium and TEC on the ACh release mechanism in the cerebral cortex and at the neuromuscular junction are compared.

Acetylcholine↗

PubMed1987

Comparison of penicillin epileptogenesis in rat somatosensory and motor cortex.

The relative sensitivities of somatosensory and motor areas of the cerebral cortex in penicillin epileptogenesis were compared in urethane-anaesthetized rats. Penicillin was applied electrophoretically from a fluid-filled micro-electrode. Spontaneous focal interictal epileptiform discharges were detected by a nearby recording electrode. In motor cortex, every cortical layer was less sensitive in penicillin epileptogenesis than the corresponding layer in somatosensory cortex; epileptic spikes occurred later, were of lower amplitude and were less frequent. In motor cortex, the sensitive depth extended from the deep part of layer III to the upper part of layer V. It seemed possible that penicillin applied to motor cortex might be producing its effects by diffusing back to the sensitive somatosensory area. This was excluded by applying penicillin to motor cortex whilst recording from both somatosensory and motor areas and demonstrating that the spikes were found in motor but not in somatosensory cortex.

Animals↗

PubMed2006

Dopamine has bidirectional effects on synaptic responses to cortical inputs in layer II of the lateral entorhinal cortex.

Dopaminergic modulation of neuronal function has been extensively studied in the prefrontal cortex, but much less is known about its effects on glutamate-mediated synaptic transmission in the entorhinal cortex. The mesocortical dopamine system innervates the superficial layers of the lateral entorhinal cortex and may therefore modulate sensory inputs to this area. In awake rats, systemic administration of the dopamine reuptake inhibitor GBR12909 (10 mg/kg, ip) enhanced extracellular dopamine levels in the entorhinal cortex and significantly facilitated field excitatory postsynaptic potentials (fEPSPs) in layer II evoked by piriform cortex stimulation. An analysis of the receptor subtypes involved in the facilitation of evoked fEPSPs was conducted using horizontal slices of lateral entorhinal cortex in vitro. The effects of 15-min bath application of dopamine on synaptic responses were bidirectional and concentration dependent. Synaptic responses were enhanced by 10 microM dopamine and suppressed by concentrations of 50 and 100 microM. The D(1)-receptor antagonist SCH23390 (50 microM) blocked the significant facilitation of synaptic responses induced by 10 microM dopamine and the D(2)-receptor antagonist sulpiride (50 microM) prevented the suppression of fEPSPs observed with higher concentrations of dopamine. We propose here that dopamine release in the lateral entorhinal cortex, acting through D(1) receptors, can lead to an enhancement of the salience of sensory representations carried to this region from adjacent sensory cortices.

Animals↗

PubMed1975

Afferent-efferent linkages in motor cortex for single forelimb muscles.

1. In locally anesthetized cats, extracellular recordings were made from single neurons in the lateral cruciate gyrus of cerebral cortex. These neurons responded to natural activation of stretch receptors in single, contralateral, forelimb wrist muscles, typically with phasic excitation. Low-velocity stretches, which activate primary endings of muscle spindles, excited one set of neurons at a mean latency of 11 ms; high-velocity stretches, which principally activate Golgi tendon organs and/or secondary spindle endings, excited a second set at 18 ms. The cortical neurons showing threshold responses to low-velocity stretches were found exclusively within restricted columns, 0.5-2.0 mm in diameter, which were spatially separate for each muscle. Neurons exhibiting threshold responses to high-velocity stretches were present in high density within the same columns and were also distributed, although more sparsely, outside the columns. 2. These afferent columns were located in cytoarchitectonic area 4gamma, and were shown by intracortical microstimulation to coincide with the efferent columns for contraction of the same muscle from which in input rose. Discrete afferent columns were also found for single muscles in the peridimple region of sensory cortex (area 3a), spatially separate from the columns in motor cortex. The excitation of the columns in motor cortex by these inputs from muscle was independent of that in sensory cortex. 3. The role of the cerebellum in controlling these feedback systems to motor cortex was investigated by selective cooling of interpositus and dentate nucleus, respectively. Cooling of interpositus markedly reduced transmission in the high-threshold system; cooling of dentate had a similar effect on the low-threshold system. 4. The latency, threshold, and cooling data indicated that the low-threshold system to motor cortex utilizes extracerebellar pathways including medial lemniscus and is facilitated by dentate nucleus. The high-threshold system involves a transcerebellar pathway including interpositus nucleus. Both systems transmit velocity-related information, with each showing different and complementary sensitivity and dynamic range. 5. The results are discussed with reference to the cortical load-compensation mechanism postulated by Phillips (37-38).

Animals↗