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Effects of anodal transcranial direct current stimulation over the right primary motor cortex on a sequential motor finger tapping task in developmental stuttering.

INTRODUCTION: This study investigates the impact of anodal transcranial direct current stimulation (tDCS) on non-speech sequential motor practice in adults who stutter (AWS), compared to non-stuttering controls (ANS). Recent research has explored the effects of tDCS on speech fluency in stuttering. However, its effect on non-speech motor tasks has not yet been studied. METHODS: 20 AWS and 30 ANS right-handed participants were randomly assigned to anodal or sham tDCS conditions, performing a sequential finger tapping task. We targeted over the right primary motor cortex, stimulating at 2 mA for 20 min. Sequence duration and reaction time were analyzed. RESULTS: AWS analysis revealed that the anodal condition had significantly slower reaction times in the second half of the task compared to sham. For sequence durations, AWS in the anodal condition had slower overall sequence durations than the sham condition. However, there were no block-by-block differences in sequence duration. When comparing AWS and ANS, no significant differences were observed for sequence duration. However, there were significant differences in reaction time between AWS and ANS, specifically in earlier blocks. Additionally, there was no significant Group × Condition interaction. DISCUSSION: The findings suggest that anodal stimulation impeded finger sequencing in AWS, showing overall slower sequence durations and a diminishing effect on reaction times in the second half of the experiment, suggesting anodal tDCS may interact uniquely with the neural mechanisms in stuttering. Future studies should explore the effects of anodal tDCS on non-speech motor tasks to gain a broader understanding of its impact on motor control and motor learning.

Humans

High-frequency contralesional dorsal premotor cortex and low-frequency contralesional primary motor cortex rTMS in subacute stroke with severe upper limb impairment: comparable motor outcomes and differential regional degree centrality changes.

BACKGROUND: The contralesional dorsal premotor cortex has been proposed as a potential neuromodulatory target for patients with severe upper limb impairment due to subacute ischemic stroke. This proof-of-concept study aimed to compare behavioral outcomes and resting-state neuroimaging findings between high-frequency repetitive transcranial magnetic stimulation (rTMS) over the contralesional dorsal premotor cortex and guideline-supported low-frequency stimulation over the contralesional primary motor cortex. METHODS: In this randomized trial, 46 patients with severe upper limb impairment in the subacute stage after ischemic stroke were randomly assigned to receive either high-frequency rTMS over the contralesional dorsal premotor cortex or low-frequency rTMS over the contralesional primary motor cortex. Low-frequency stimulation over the contralesional primary motor cortex served as an evidence-supported active comparator for poststroke upper limb motor recovery. Stimulation was administered five times per week for two weeks using magnetic resonance imaging-guided neuronavigation. All participants received concurrent standard rehabilitation therapy. The primary outcome was the Fugl-Meyer Assessment for Upper Extremity. Secondary outcomes included the Arm Subscore of the Motricity Index, the Hong Kong version of the Functional Test for the Hemiplegic Upper Extremity, the Modified Barthel Index, and resting-state functional magnetic resonance imaging-derived degree centrality. RESULTS: Both groups showed significant improvements in the primary and secondary behavioral measures (p&#x202f;<&#x202f;0.01), with no significant between-group differences in the magnitude of change (p&#x202f;>&#x202f;0.05). In neuroimaging analyses, patients receiving high-frequency rTMS over the contralesional dorsal premotor cortex showed significantly greater degree centrality changes in the ipsilesional middle occipital gyrus, contralesional medial superior frontal gyrus, and contralesional middle frontal gyrus than those receiving low-frequency rTMS over the contralesional primary motor cortex (p&#x202f;<&#x202f;0.05). Within the high-frequency stimulation group, degree centrality changes in the ipsilesional middle occipital gyrus were positively correlated with improvements in the Fugl-Meyer Assessment for Upper Extremity (r&#x202f;=&#x202f;0.619, false discovery rate-corrected p&#x202f;=&#x202f;0.018). CONCLUSIONS: High-frequency rTMS over the contralesional dorsal premotor cortex produced behavioral improvements comparable to guideline-supported low-frequency rTMS over the contralesional primary motor cortex, without establishing superiority or formal non-inferiority. Exploratory neuroimaging analyses showed greater degree centrality changes in the ipsilesional middle occipital gyrus after high-frequency premotor stimulation, and these changes correlated with upper-limb motor improvement. These findings support further investigation of contralesional dorsal premotor cortex-targeted high-frequency rTMS for severe subacute post-stroke upper limb impairment. REGISTRATION: URL: http://www.chictr.org.cn; Unique identifier: ChiCTR2000038049.

Humans

Anatomical analysis of ventrolateral thalamic input to primate motor cortex.

1. The origin and topographical organization of input to the arm area of the primate motor cortex from the ventrolateral thalamus were examined using the method of retrograde transport of horseradish peroxidase (HRP). 2. A thin, continuous slab of labeled neurons was found in the ventrolateral thalamus followingmultiple injections of HRP into the arm area of the motor cortex. The slab of labeled neurons was flanked, medially and laterally, by groups of unlabeled neurons. 3. The origin of ventrolateral thalamic input was more extensive than previously thought. Labeled neurons were found from A10.0 to A6.0 and occurred in three ventolateral thalamic subdivisions: ventralis lateralis pars oralis (VLo), ventralis lateralis pars caudalis (VLc), and ventralis posterior lateralis pars oralis (VPLo). For simplicity this region containing labeled neurons has been termed the ventrolateral thalamic (VL) arm area. 4. Injections of HRP into the somatic sensory cortex indicated that the thalamic regions which project to the somatic sensory cortex are separate from the VL arm area. 5. The distribution of labeled neurons following single injections of HRP into different regions of the motor cortex arm area indicated that the VL arm area is topographically organized, particularly its caudal part. Ventral regions of the VL arm area were labeled following HRP injections into motor cortex regions adjacent to the central sulcus where the representation of largely distal musculature is localized. Dorsal regions of the VL arm area were labeled following HRP injections into motor cortex regions more rostral to the central sulcus where the representation of more proximal musculature is localized. 6. A larger region of the VL arm area was labeled following HRP injections adjacent to the central sulcus than following the more rostral motor cortex injections. This suggests that, like the arm area of the motor cortex, more of the VL arm area is allotted to the representation of distal than proximal musculature. 7. Following very small cortical HRP injections, isolated labeled thalamic neurons were diffusely scattered throughout a 3-mm rostrocaudal extent of the VL arm area. In addition, a small focal cluster of labeled thalamic neurons was also seen. The labeled cluster was limited to 0.5 mm rostrocaudally and 300 mum in width. The focal distribution of labeled thalamic neurons suggests that aspects of a point to point organization may exist in the connection between VL and the motor cortex arm area.

Afferent Pathways

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

Responses of neurones in motor cortex and in area 3A to controlled stretches of forelimb muscles in cebus monkeys.

1. The experiments were designed to investigate the effects of longitudinal muscle displacements on neurones of the motor cortex of anaesthetized Cebus monkeys and thus test the hypothesis that signals from muscle spindles may modify motor cortical output. The effects of sinusoidal stretching of the extensor digitorum communis (EDC) at frequencies varying from 6 to 300 Hz and of step and rhomboidal stretches were studied in neurones of the motor cortex. For comparison, neurones of the primary receiving area for low-threshold muscle afferents, cortical area 3a, were also included in this study. Neurones of the motor cortex were subdivided into corticospinal (PT) neurones and non-corticospinal (non-PT) neurones. 2. Threshold stretch amplitudes were clearly higher for neurones of area 4 (PT and non-PT) than for 3a neurones. However, a conspicuous fall in threshold stretch amplitude was observed for all three neurone populations when the frequency of sinusoidal stretching was increased (highest frequency: 300 Hz). A small number of non-PT and PT neurones responded to vibration amplitudes of less than 100 mum and some of these low-threshold cells of area 4 also responded to rhomboidal stretches of 8 mm/sec ramp velocity and 80 mum plateau amplitude. Increasing the stretch amplitude to twice threshold nearly doubled the output magnitude in all three cell types. Neurones of area 3a and non-PT neurones of area 4 had similar latencies, and these were significantly shorter than the latencies of PT neurones tested with trains of high frequency vibration. Dynamic response patterns were observed in all three cell types, but most frequently in 3a neurones. 3. It is concluded that, in Cebus monkeys, signals from both primary and secondary muscle spindle endings from forelimb muscles reach the motor cortex. Under the present experimental conditions, the input from the primaries to the motor cortex was effective only if these spindle receptors were driven maximally by vibratory stimuli. The particularly low probability of stretch-evoked discharges of cortico-spinal neurones in the anaesthetized preparation may be explained by a low gain in transmission from input to output cells of the motor cortex.

Animals

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

Correlation of neural discharge with pattern and force of muscular activity, joint position, and direction of intended next movement in motor cortex and cerebellum.

1. Monkeys were trained to grasp a rod movable in a horizontal arc (Fig. 1), and to hold the rod by angulation of the wrist in each of three positions (A,B, C). A maintained load was placed on the rod alternately to oppose flexion and extension. At a light signal, the monkey had to move to the next position in a prescribed sequence (ABCBABCBA, ETC.). The task was designed to dissociate, while holding in position, the following variables: 1) pattern of muscular activity in the forearm required to hold the wrist in position, determined by the direction of the load (flexor or extensor muscles); 2) position of the rod, and thus angulation of the wrist joint (A, B, and C); and 3) set for the direction of the intended next movement (flexor or extensor). These variables are subsequently referred to as MPAT, JPOS, and DSET, respectively. 2. After training, recordings were made of the EMG activity of muscles used in the task and of the discharge of single neurons in the motor cortex of the cerebrum and the interposed and dentate nuclei of the cerebellum. 3. While holding the wrist in position, EMG and interpositus behaved uniformly, with higher discharge frequency under load in one direction and lower discharge frequency under load in the opposite direction. This relation was relatively independent of the position held and of the direction of the intended next movement. Thus, interpositus and EMG both seemed best related to the MPAT variable, as opposed to JPOS and DSET variables. By contrast, neurons in motor cortex and in dentate fell into three categories: one category discharged in relation to the pattern of muscular activity (MPAT), a second to the position of the wrist (JPOS), and a third to the direction of the intended next movement (DSET). While MPAT neurons formed a distinct dissociated group, neurons that were best related to JPOS were often related to DSET, and vice versa. 4. A few of the MPAT neurons in interpositus and motor cortex were further studied by varying the magnitude (as well as the direction) of the loads. Both interpositus and motor cortex MPAT neurons changed firing frequency in relation to the magnitude of load, and though few neurons were thus studied, the relation seemed clearer for interpositus than for motor cortex. 5. Anatomically, the three types of neurons thus classified by firing pattern during the hold periods were intermixed in the arm area of motor cortex. In dentate and interpositus, those neurons thus related to the performance were localized to a narrow strip across the posterior part of both nuclei. Neurons apparently related to eye and drinking movements were located more posteriorly still, suggesting somatotopic representation.

Animals

Pyramidal cell abnormalities in the motor cortex of a child with Down's syndrome. A Golgi study.

The neuronal organization of the motor cortex of a 19-month old child with Down's syndrome (mongolism) has been studied with the rapid Golgi method. This congenital syndrome, also known as 21 Trisomy is caused by a chromosomal abnormality consisting of the presence of an extra chromosome in the group 21. Various structural abnormalities have been found in the dendritic spines (postsynaptic structures) of the pyramidal neurons of the motor cortex of this child. The axo-spinous synapses of these neurons are considered to be altered by these spine abnormalities. In addition, a peculiar form of intrinsic vacuolar change affecting the dendrites and scattered neuronal fragmentation and necrosis have also been found. At least three different types of abnormality involving the spines--(the unusually long spine, the very short spine and a reduction in the number of spines)--are recognized among the pyramidal cells of the motor cortex. It is postulated herein: that a basic anomaly, possibly related to the genetic disorder affects primarily some cortical neurons which undergo progressive degenerative changes terminating in cell fragmentation and death. The different spine abnormalities are considered to represent various developmental stages of the common genetic anomaly. These changes might be structural correlates of the motor incoordination and mental retardation which are characteristic of this genetic disorder, but, final conclusions should await the investigation of other cases with this or similar methods capable of demonstrating the normal as well as the abnormal structural organization of the human cerebral cortex.

Dendrites

Does the supplementary motor area play a part in modifying motor cortex reflexes?

Neuronal activity in the supplementary motor area was recorded from a monkey performing a trained motor task that required readiness for proper usage of sensory inputs. Thirty-two neurons exhibited activity changes, which supports the hypothesis that the SMA is part of the system involved in modulating responsiveness of the motor cortex to sensory inputs in association with learned movements.

Animals

Comparative cellular analysis of motor cortex in human, marmoset and mouse.

The primary motor cortex (M1) is essential for voluntary fine-motor control and is functionally conserved across mammals1. Here, using high-throughput transcriptomic and epigenomic profiling of more than 450,000 single nuclei in humans, marmoset monkeys and mice, we demonstrate a broadly conserved cellular makeup of this region, with similarities that mirror evolutionary distance and are consistent between the transcriptome and epigenome. The core conserved molecular identities of neuronal and non-neuronal cell types allow us to generate a cross-species consensus classification of cell types, and to infer conserved properties of cell types across species. Despite the overall conservation, however, many species-dependent specializations are apparent, including differences in cell-type proportions, gene expression, DNA methylation and chromatin state. Few cell-type marker genes are conserved across species, revealing a short list of candidate genes and regulatory mechanisms that are responsible for conserved features of homologous cell types, such as the GABAergic chandelier cells. This consensus transcriptomic classification allows us to use patch-seq (a combination of whole-cell patch-clamp recordings, RNA sequencing and morphological characterization) to identify corticospinal Betz cells from layer 5 in non-human primates and humans, and to characterize their highly specialized physiology and anatomy. These findings highlight the robust molecular underpinnings of cell-type diversity in M1 across mammals, and point to the genes and regulatory pathways responsible for the functional identity of cell types and their species-specific adaptations.

Animals

Influences of cerebellar hemispherectomy on slow potentials in the motor cortex preceding self-paced hand movements in the monkey.

With chronically implanted electrodes, surface negative and deep positive, slowly increasing potentials were recorded in the forelimb area of the motor cortex prior to self-paced movements of the contralateral hand in monkeys. The slow premovement potentials were markedly reduced in size after ablation of the cerebellar hemisphere on the contralateral side to the motor cortex under recording. It was suggested that the cerebellar hemisphere (neocerebellum) participates in preparing the activity of the motor cortex prior to voluntary movements.

Animals

Reversibility of lipofuscin accumulation caused by protein malnutrition in the motor cortex of squirrel monkeys, Saimiri scireus.

Lipofuscin pigment has been demonstrated histochemically in the motor cortex by the use of several histochemical and cytochemical methods in the healthy (maintained on a diet with 25% protein content) and protein malnourished (maintained on a diet with 2% protein content) adult animals. 4 animals in the latter category were rehabilitated over a period of 11 months and various histochemical techniques were repeated on the motor cortex of these animals. The healthy animals showed a somewhat uniform distribution of lipofuscin pigment in the neuronal perikarya with the perineuronal glia showing only slight occurrence of pigment bodies. The malnourished animals exhibited a significantly larger number of lipofuscin bodies in the neuronal as well as glial perikarya. The neurons, especially, showed aggregations of lipofuscin bodies characterized by a large increase in the activity of acid phosphatase and simple esterases. The rehabilitated animals, however, showed a decrease of lipofuscin pigment in the neuronal perikarya with a concomitant loss of lysosomal enzymes, while a significant increase of these bodies was observed in the perineuronal glial cells. It is evident that the formation of lipofuscin pigment gets accelerated under the extrinsic influence of dietary protein deprivation in the adult animals, but the process is reversed at least to some extent by halting the dietary deficiency or its correction by rehabilitation. The perineuronal glial cells appear to play a significant role in the removal of lipofuscin bodies from the neuronal perikarya. The significance of these observations has been discussed.

Animals

Commissural columns in the sensory-motor cortex of monkeys.

Callosally projecting cells and the terminal ramifications of their axons were identified in the monkey sensory-motor cortex by retrograde and anterograde labeling techniques, often by double labeling cells and axons in the same animal. Bundles of callosal fibers terminate in small column-like zones 0.5-1 mm wide in the motor cortex (area 4) and in the first (SI) and second (SII) somatic sensory areas. Such columns are aligned in register to form elongated strips extending mediolaterally in the long axes of the pre- and postcentral gyri. Significant portions of area 4, SI and SII, in regions corresponding to the representations of the hand and foot, are not callosally connected. The cells of origin of callosal fibers in SI are largely confined to layer IIIB and form columns and strips corresponding to the above. In connected zones of SI, the callosal connection is reciprocal and precisely point-to point. This and the laminar distribution of the terminal ramifications of callosal fibers (to layers I-IV) suggest that callosal fibers may arise from the terminate upon exactly homotopic, column-like groups of layer IIIB pyramidal cells. Commissurally projecting cells and their terminal ramifications are not limited to particular architectonic fields or particular parts of fields in SI. All architectonic fields of SI project heterotopically to the contralateral SII.

Animals

Projections of the precentral motor cortex and other cortical areas of the frontal lobe to the subthalamic nucleus in the monkey.

Corticofugal projections were examined by means of the autoradiographic tracer method in 21 macaca fascicularis. The labeled material was injected into the main body representation areas of the precentral motor cortex and into various regions of Brodmann's areas 6, 8 and 9 of the frontal lobe. The ipsilateral subthalamic nucleus receives a moderately strong and somato-topically organized projection from Woolsey's precentral motor cortex. This projection is mainly restricted to the lateral moiety. The remaining nucleus is occupied by less intensive projections from premotor and prefrontal areas. It is concluded that the subthalamic nucleus is a convergence site of pallidal and corticomotor and frontal projections. Cortical afferents may exert an influence on the pallido-subthalamic-pallidal inhibitory feedback loop.

Animals

Properties of the pyramidal tract neuron system within the precentral wrist and hand area of primate motor cortex.

1. To obtain basic anatomical data that will be useful in interpreting the results of studies of primate pyramidal tract neurons (PTNs), extracellular, single-unit recording techniques were used to determine a number of the properties of the PTN population within the electrically defined, precentral wrist zone of the monkey's motor cortex. 2. Recordings were obtained from a total of 1,375 antidromically identified PT and corticospinal tract (CST) cells. A mathematical model was then used to correct the statistics of the sample for variations in the probability of unit detection, which arise from variations in neuronal size and extracellular field dimensions. 3. Both the experimentally observed and theoretically corrected results suggest that the PT projection from this cortical zone is derived principally from slowly conducting, and presumably small to medium-sized cells (an estimated 85% of the resident PTN population). 4. Both the fast and slow cell subpopulations were found to be concentrated within cortical layer V, where they tend to congregate in small, mixed clusters of 2 to 5 neurons. Estimates of the total packing density of PTNs within layer V of this cortical zone suggest that they account for only 10-20% of the neurons within this major efferent layer. 5. 70% of the slow and 82% of the fast PT neurons within this cortical area were found to send their axons into the contralateral, lateral corticospinal tract. Thus, in futur functional studies of PTNs in this cortical area, it can be assumed that three of every four neurons will in fact influence segmental cells of one category or another directly. 6. Extensive data are also presented on the incidence of axon collateral branching from PT and CST cells to the red nucleus, the medial medullary reticular formation and the cuneate nucleus. 7. Some general implications of these findings for the design of future functional studies of anatomically identified motor cortex cell systems are then discussed.

Action Potentials

Conditioned changes of synaptic transmission in the motor cortex of the cat.

Intracellular recordings were made from 117 neurons in the motor cortex of anesthetized cats. The pyramidal tract (PT) and VL nucleus of thalamus were stimulated in order to activate the neurons from two directions. 1. PT cells were conditioned by antidromic trains (10--50 cps for 4--15 s) and by paired PT and VL stimuli with different intervals and sequences. The VL-EPSPs were examined before and after conditioning, to find differences in efficacy in giving rise to spikes. The conditioning procedures resulted in a remarkable facilitation of VL-EPSPs, manifesting itself as a significant rise of efficacy in generating spikes, a shortening of peak latency and in some cases, an enhancement of background firing. 2. In non-PT neurons the same conditioning procedures elicited heterosynaptic facilitation and a rise in firing activity. 3. Intracellularly injected square wave pulses also resulted in facilitation of VL-EPSPs. 4. Pairings of PT and VL stimuli were more effective than trains in evoking conditioned changes. 5. Plastic modifications were observed in the 13.7% of the neurons subjected to conditioning procedures. 6. The authors assume that synchronous activity of the pre- and postsynaptic neurons is a highly important condition for plastic changes in the efficacy of synaptic transmission.

Animals

Intracellular synaptic potentials of primate motor cortex neurons during voluntary movement.

An intracellular recording technique was applied to the precentral motor cortex of the unanesthetized, chronically behaving monkey. Postsynaptic potentials, responsible for an initiation of the voluntary movement, were recorded. In total, 22 pyramidal tract neurons (PTNs) and 40 non-pyramidal tract neurons (non-PTNs) were successfully penetrated in 5 monkeys while the monkey was performing a flexion-extension wrist movement after a visual cue (reaction time, 200--350 msec). The neurons showed a negative membrane potential shift of at least 30 mV for more than 30 sec. A slowly rising PSP appeared 80--180 msec after the visual cue, and was 70--180 msec prior to an onset of the movement. Spike activities were superimposed upon this slow PSP with 20--80 msec rise time and 2--6 mV depolarization (8 PTNs and 6 non-PTNs). Since these depolarizations were variable in magnitude and latency, these were considered to be summated potentials of small EPSPs and hidden IPSPs. Membrane resistance was measured from an IR drop by a hyperpolarizing current (1.2 X 10(-9) A) passed through a recording electrode, and was 3.5 +/- 1.7 Momega in 18 PTNs and 4.5 +/- 2.5 Momega in 28 non-PTNs. There was a linear relationship in PTNs between membrane resistance and antidromic latency from the pontine pyramid. Because of the time course of PSPs, their possible dendritic origins were discussed.

Animals

Dendro-dendritic and reciprocal synapses in the primate motor cortex.

Dendro-dendritic synapses have been observed infrequently in the deep layers of the motor cortex. The presynaptic dendrites are of a varicose type and themselves receive a considerable density of synapses both of the asymmetric and symmetrical type. The ultrastructure of the dendro-dendritic synapse itself shows the typical arrangement of presynaptic and postsynaptic membrane densities, often with presynaptic dense projections, and the membrane specialization is of the symmetrical type. There is the usual cleft containing electron-dense material between the presynaptic and postsynaptic profiles. The synaptic vesicles occur in a small cluster confined to a region close to the presynaptic membrane specialization; some of the vesicles are flattened and were shown by tilt analysis to be of the discoid type. Two examples were found of reciprocal dendro-dendritic synapses, both components being of the symmetrical type. A single axon terminal may make a synapse on to both dendrites involved in a dendro-dendritic synapse.

Animals