Thalamic projections to areas 3a, 3b,and 4 in the sensorimotor cortex of the mature and infant macaque monkey

Area 3a in the macaque monkey, located in the fundus of the central sulcus, separates motor and somatosensory cortical areas 4 and 3b. The known connections of areas 4 and 3b differ substantially, as does the information which they receive, process, and transfer to other parts of the central nervous system. In this analysis the thalamic projections to each of these three cortical fields were examined and compared by using retrogradely transported fluorescent dyes (Fast Blue, Diamidino Yellow, Rhodamine and Green latex microspheres) as neuron labels. Coincident labeling of projections to 2–3 cortical sites in each monkey allowed the direct comparison of the soma distributions within the thalamic space of the different neuron populations projecting to areas 3a, 3b, and 4, as well as to boundary zones between these cortical fields. The soma distribution ofthalamic neurons projecting to a small circumscribed zone (diameter = 0.5–1.0 mm) strictly within cortical area 3a (in region of hand representation) filled out a “territory” traversing the dorsal half of the cytoarchitectonically defined thalamic nucleus, VPLc (abbreviations as in Olszewski [1952] The Thalamus of the Macaca mulatta. Basel: Karger). This elongate, rather cylindrical, territory extended caudally into the anterior pulvinar nucleus, but not forward into VPLo. The rostrocaudal extent of the thalamic territory defining the soma distribution of neurons projecting to small zones of cortical area 3b was similar, but typically extended into the ventral part of VPLc, filling out a medially concavo-convex laminar space. Two such territories projecting to adjacent zones of areas 3a and 3b, respectively, overlapped and shared thalamic space, but not thalamic neurons. Contrasting with the 3a and 3b thalamic territories, the soma distribution of thalamic neurons projecting to a circumscribed zone in the nearby motor cortex (area 4) did not penetrate into VPLc, but instead filled out a mediolaterally flattened territory extending from rostral VLo, VLm, VPLo to caudal and dorsal VLc, LP, and Pulo. These territories skirted around VPLc. All three cortical areas (4, 3a, and 3b) also received input from distinctive clusters of cells in the intralaminar Cn.Md. It is inferred that, in combination, the thalamic territories in areas 3a, 3b, and 4 (and also area 1 and 2), which would be coactive during the execution of a manual task, constituted a lamellar space extending from VLo rostrally to Pul.o caudally. How Pul.o neuron populations relate to the more rostral populations within the same thalamic territory projecting to a localized cortical zone remains uncertain. Within the medially located territories the distribution of the neuron population in Pul.o was spatially continuous with the more rostral thalamic cells projecting to the same localized cortex, but in lateral thalamic territories these 2 populations were usually spatially discontinous. In the newborn macaque an orderly change in the territorial projections to localized zones in area 4, 3a, and 3b was also demonstable. However, thalamic nuclear projections were more expansive than in the mature animal. As well as the VPLc input, a third of the thalamic input to area 3a was now from VLo, VPLo, and VLm. Area 4 also had a significant input from VPLc, an input not observed in the mature macaque. © 1993 Wiley-Liss, Inc.     

Ipsilateral cortical projections to areas 3a, 3b,and 4 in the macaque monkey

In the macaque monkey area 3a of the cerebral cortex separates area 4, a primary motor cortical field, from somatosensory area 3b, which has a subcortical input mainly from cutaneous mechanoreceptive neurons. That each of these cortical areas has a unique thalamic input was illustrated in the preceding paper. In the present experiments the cortical afferent projections to these 3 areas of the sensorimotor cortex monkey were visualized and compared, using 4 differentiable fluorescent dyes as axonal retrogradely transported labels. The cortical projection patterns to areas 3a, 3b, and 4 were similar in that they each consisted of (a) a “halo” of input from the immediately surrounding cortex, and (b) discrete projections from one or more remote cortical areas. However, the pattern of remote inputs from precentral, mesial, and posterior parietal cortex was different for each of the 3 cortical target areas. The cortical input configuration was least complex for area 3b, its remote input projecting mainly from insular cortex. The pattern of discrete cortical inputs to the motor area 4, however, was more complex, with projections from the cingulate motor area (24c/d), the supplementary motor area, postarcuate cortex, insular cortex, and postcentral areas 2/5. Area 3a, in addition to the proximal projections from the immediately surrounding cortex, also received input from the supplementary motor area, cingulate motor cortex, insular cortex, and areas 2/5. Thus, this pattern of cortical input to area 3a resembled more closely that of the adjacent motor rather than that of the somatosensory area 3b. Contrasting with this, however, the thalamic input to area 3a was largely from somatosensory VPLc (abbreviations from Olszewski [1952] The Thalamus of the Macaca mulatta. Basel: Karger) and not from VPLo (with input from cerebellum, and projecting to precentral motor areas). © 1993 Wiley-Liss, Inc.     

Thalamic projections to sensorimotor cortex in the newborn macaque.

In the present experiments thalamocortical projections to different functional areas of the newborn (or prematurely delivered) macaque's sensorimotor cortex were labeled using retrogradely transported fluorescent dyes. Several dyes were used in each animal to (1) enable the direct comparison of the soma distributions of different thalamocortical projections within thalamic space, and (2) identify by double labeling neurons shared between these distributions. The projection patterns in the newborn macaque were compared with those of the mature animal reported by Darian-Smith et al. (J. Comp. Neurol. 1990;298:000-000). The main observations were (1) all thalamocortical projections to the sensorimotor cortex of the mature macaque are well established by embryonic days 146-150, as was shown by labeling these pathways in infants delivered by cesarean section, (2) a significant number of thalamocortical neurons in the newborn were double-labeled following dye injections into different pre- or postcentral areas, and where the margins of the dye uptake zones were separated by 3-8 mm, and (3) extensive projections from the anterior pulvinar nucleus to the motor and premotor cortex, and to the supplementary motor cortex were labeled in the newborn macaque. Both the exuberant terminal arborizations, and the precentral pulvinar projections were diminished by the 6th postnatal month, and absent in the mature macaque. The role of epigenetic determinants of these postnatal events is briefly considered.     

Neural circuits and functional organization of the striatum

The basal ganglia and motor thalamic nuclei are functionally and anatomically divided into the sensorimotor, supplementary motor, premotor, associative and limbic territories. There exist both primary segregated basal ganglia-thalamocortical loops and convergence of functionally related information from different cortical areas onto these cortical basal gaglia-thalamocortical loops. The basal ganglia-thalamocortical loop arising from the sensorimotor area, supplementary motor area (SMA), premotor area and cingulate motor area provides distinct segregated subloops through the functionally distict stritial, pallidal and thalamic regions with partial overlap. The subthalamic nucleus (STN) is also topographically organized. The ventrolateral part of the caudal 2/3 levels of the medial pallidal segment (GPi) projects to the primary motor area via the oral part of the ventral lateral thalamic nucleus (VLo) (Voa, Vop by Hassler's nomenclature). The thalamic relay nuclei of the GPi projection to SMA are identified in the transitional zoe of the VApc (parvicellular part of the anterior ventral nucleus)-VLo and in the rostromedical part of the VLo. The thalamic nuclei relaying the cingulate subloop are not yet clearly defined. The supplementary motor subloop appears to be divided into the pre-SMA and SMA proper subloops. The premotor area is also divided into the dorsal premotor area subloop and the ventral premotor area subloop. It is suggested that the limbic loop consists of a number of subloops in the monkey as indicated by Haber et al. [67] and in rats [64]. We review here the microcircuitry of the striatum, as well as the convergence and integration between the functionally segregated loops. Finally, we discuss the functional implications of stritial connections.

     

How do the basal ganglia and cerebellum gain access to the cortical motor areas?

We have used retrograde transport of wheat germ agglutinin conjugated to horseradish peroxidase to examine the origin of the thalamic input to the two premotor areas with the densest projections to the motor cortex. These are: arcuate premotor area (APA) and the supplementary motor area (SMA). Retrograde transport demonstrated that the two premotor areas and the motor cortex each receive thalamic input from separate, cytoarchitectonically well-defined subdivisions of the ventrolateral thalamus. According to the nomenclature of Olszewski (1952), input to the APA originates largely from area X; input to the SMA originates largely from the pars oralis subdivision of the nucleus ventralis lateralis (VLo); and that to the motor cortex is largely from the pars oralis subdivision of the nucleus ventralis posterior lateralis (VPLo). These observations, when combined with prior studies on the termination of various subcortical efferents in the thalamus, lead to the following scheme of projections: rostral portions of the deep cerebellar nuclei project to motor cortex via VPLo, caudal portions of the deep cerebellar nuclei project to the APA via area X; and the globus pallidus projects to the SMA via VLo. Thus each thalamocortical pathway is associated with a distinct subcortical input.     

The origin of thalamic inputs to the arcuate premotor and supplementary motor areas

We have used retrograde transport of wheat germ agglutinin conjugated to horseradish peroxidase to examine the origin of thalamic input to the two premotor areas with the densest projections to the motor cortex: the arcuate premotor area (APA) and the supplementary motor area (SMA). Retrograde transport demonstrated that the two premotor areas and the motor cortex each receive thalamic input from separate, cytoarchitectonically well defined subdivisions of the ventrolateral thalamus. According to the nomenclature of Olszewski (Olszewski, J. (1952) The Thalamus of the Macaca mulatta. An Atlas for Use with the Stereotaxic Instrument, S. Karger, AG, Basel), input to the APA originates largely from area X, input to the SMA originates largely from the pars oralis subdivision of nucleus ventralis lateralis (VLo), and that to the motor cortex originates largely from the pars oralis subdivision of nucleus ventralis posterior lateralis (VPLo). These observations, when combined with prior studies on the termination of various subcortical efferents in the thalamus, lead to the following scheme of connections: (1) rostral portions of the deep cerebellar nuclei project to the motor cortex via VPLo; (2) caudal portions of the deep cerebellar nuclei project to the APA via area X; and (3) the globus pallidus projects to the SMA via VLo. Thus, each thalamocortical pathway is associated with a distinct subcortical input.     

Comparison of cerebral activation between motor execution and motor imagery of self-feeding activity

Motor imagery is defined as an act wherein an individual contemplates a mental action of motor execution without apparent action. Mental practice executed by repetitive motor imagery can improve motor performance without simultaneous sensory input or overt output. We aimed to investigate cerebral hemodynamics during motor imagery and motor execution of a self-feeding activity using chopsticks. This study included 21 healthy right-handed volunteers. The self-feeding activity task comprised either motor imagery or motor execution of eating sliced cucumber pickles with chopsticks to examine eight regions of interest: pre-supplementary motor area, supplementary motor area, bilateral prefrontal cortex, premotor area, and sensorimotor cortex. The mean oxyhemoglobin levels were detected using near-infrared spectroscopy to reflect cerebral activation. The mean oxyhemoglobin levels during motor execution were significantly higher in the left sensorimotor cortex than in the supplementary motor area and the left premotor area. Moreover, significantly higher oxyhemoglobin levels were detected in the supplementary motor area and the left premotor area during motor imagery, compared to motor execution. Supplementary motor area and premotor area had important roles in the motor imagery of self-feeding activity. Moreover, the activation levels of the supplementary motor area and the premotor area during motor execution and motor imagery are likely affected by intentional cognitive processes. Levels of cerebral activation differed in some areas during motor execution and motor imagery of a self-feeding activity. This study was approved by the Ethical Review Committee of Nagasaki University(approval No. 18110801) on December 10, 2018.     

Relations between the basal ganglia and the thalamus of the primate. New morphologic data. New physiopathologic interpretations

Considerable progress has been made over the last few years in our knowledge of the thalamus and basal ganglia and their relationships to the cerebral cortex. More detailed topographic studies in the macaque have demonstrated the separation, in the lateral region of the thalamus, between afferent cerebellar and basal ganglia territories. These territories fail to correlate with the subdivision between ventral and dorsal elements or the limits of a single cytoarchitectonic nucleus. The cerebellar territory corresponds to VIL (or VPLo) which projects towards the primary cortex, and to VIM (or area X) and DI (or VLc) which project towards premotor cortex. The nigral (and tectal) territory corresponds to VOM (or VAmc) and to some parts of the medial nucleus and projects mainly towards the oculomotor area, supplementary motor area and prefrontal cortex. In return, the oculomotor area and substantia nigra project towards the colliculus superior. Several thalamic nuclei constitute the pallidal territory: VOL (or VLo) projects mainly towards supplementary motor area, LPo (or VApc) and Do towards the prefrontal cortex. The median center, which receives afferents from pallidum and motor cortex, projects towards the striatum but also the motor cortex. The parafascicular nucleus projects towards the striatum and premotor cortex. It is still not possible to transpose data acquired in the macaque to man, but functional reinterpretations are possible. A system which involves the median pallidum, VOL and supplementary motor area could control motor initiative and flow of movement. A second system, involving the substantia nigra, colliculus superior, thalamic relay and oculomotor area could control posture. The pallidum and substantia nigra, anterior part of lateral mass, medial nucleus and prefrontal cortex could elaborate motor programmes.     

Spatial distribution of thalamic projections to the supplementary motor area and the primary motor cortex: A retrograde multiple labeling study in the macaque monkey

The exact knowledge on spatial organization of information sources from the thalamus to the supplementary motor area (SMA) and to the primary motor cortex (MI) has not been established. We investigated the distribution of thalamocortical neurons projecting to forelimb representations of the SMA and the MI using a multiple retrograde labeling technique in the monkey. The forelimb area of the SMA, and the distal and proximal forelimb areas of the MI were identified by electrophysiological techniques of intracortical microstimulation and single neuron recording. Injections were made into these three representations with three different dyes in the same animal (horseradish peroxidase conjugated to wheat germ agglutinin, diamidino yellow, and fast blue), and the thalamic neurons were retrogradely labeled. Injections into the SMA densely labeled thalamic neurons in nuclei ventralis lateralis pars oralis (VLo), ventralis lateralis pars medialis (VLm) and ventralis lateralis pars caudalis (VLc), but not in nucleus ventralis posterior lateralis pars oralis (VPLo). Injections into the MI labeled thalamic neurons primarily in VLo, VLc, and VPLo. We found that the distribution of projection neurons to the three areas was largely separate in the thalamus. However, in the middle part of VLo, and in a limited portion of VLc, thalamic neurons projecting to the SMA partially overlapped with those to the distal forelimb area of the MI. They overlapped little with those to the proximal forelimb area of the MI. We noted no overlap between the distributions of thalamic projection neurons to the distal and proximal forelimb areas of the MI. These findings suggest that the SMA and MI receive separate information from the thalamus, while sharing minor sources of common inputs. © 1995 Wiley-Liss, Inc.     

Topography and collateralization of the dopaminergic projections to motor and lateral prefrontal cortex in owl monkeys.

The sources and histochemical characteristics of dopaminergic projections to motor and premotor areas of cortex were investigated in owl monkeys in which information from related studies was used to subdivide cortex into motor fields. Brainstem projections to frontal cortex were identified by injections of different fluorescent dyes in the primary motor cortex (M1) and the supplementary motor area (SMA), first identified by microstimulation. Injections were also placed in dorsal premotor cortex and lateral prefrontal cortex. The distribution of retrogradely labeled neurons was related to the location of tyrosine hydroxylase immunolabeled neurons on the same or alternate brain sections to identify the dopamine (DA) neurons. All DA cortically projecting neurons were located in the A8-A10 complex, largely in its dorsal components, including the parabrachial pigmented n. of the ventral tegmental area (VTA), pars gamma of the substantia nigra compacta, and the dorsal part of the retrorubral area (A8). Fewer cells were in the midline groups of VTA (n. linearis rostralis and caudalis) and in the n. paranigralis. DA neurons projecting to M1, SMA, and prefrontal cortex were largely intermixed, and some of these neurons were double or triple labeled by the fluorescent dyes, indicating collateralization to two or three fields; DA cells projecting to M1 were more numerous than to the other locations. The dorsal components of the A8-A10 complex from which arose the DA mesocortical projection were also characterized by the presence of calbindin-immunoreactive neurons and by a dense neurotensin and noradrenergic terminal innervation. Compared to rodents or felines, the DA neurons projecting to the lateral frontal lobe of primates appear to be shifted dorsally and laterally in the nigral complex. The topographic overlap, partial collateralization, and common histochemical characteristics of the DA mesocortical neurons projecting to different fields of the lateral frontal lobe suggest that some degree of functional unity exists within this projection.     

Cortical activation in patients with functional hemispherectomy

Functional hemispherectomy, a safe and effective therapeutical procedure in medically intractable epilepsy, offers the chance to investigate a strictly unilateral cortical activation in ipsilateral limb movement. We assessed the pattern of cortical activation in a group of patients following functional hemispherectomy. We measured regional cerebral blood flow (rCBF) in 6 patients postoperatively and 6 normal subjects with positron emission tomography using ¹⁵[O]H₂O as a tracer. Brain activation was achieved by passive elbow movements of the affected arm. Analysis of group results and between-group comparisons were performed with statistical parametric mapping, (SPM96). In normal subjects brain activation was found contralaterally in the cranial sensorimotor cortex and the supplementary motor area and ipsilaterally in the inferior parietal cortex. In patients significant rCBF increases were found in the inferior parietal cortex, caudal sensorimotor cortex and the supplementary motor area ipsilaterally. The activation was weaker than in normal subjects. Compared with normal subjects patients showed additional activation in the premotor cortex, caudal sensorimotor cortex and the inferior parietal cortex of the remaining hemisphere. Less activation compared with normal subjects was found in the cranial sensorimotor cortex and the supplementary motor area. A functional network connecting the inferior parietal cortex, premotor cortex and the supplementary motor area as well as the existence of ipsilateral projections originating from these regions may explain why these areas are predominantly involved in reorganization confined to a single hemisphere. Received: 24 November 2000, Received in revised form: 8 March 2001, Accepted: 10 April 2001     

hTe cortical activation pattern during bilateral arm raising movements

Bilateral arm raising movements have been used in brain rehabilitation for a long time. However, no study has been reported on the effect of these movements on the cerebral cortex. In this study, using functional near infrared spectroscopy (fNIRS), we attempted to investigate cortical activation generated during bilat-eral arm raising movements. Ten normal subjects were recruited for this study. fNIRS was performed using an fNIRS system with 49 channels. Bilateral arm raising movements were performed in sitting position at the rate of 0.5 Hz. We measured values of oxyhemoglobin and total hemoglobin in ifve regions of interest:the primary sensorimotor cortex, premotor cortex, supplementary motor area, prefrontal cortex, and pos-terior parietal cortex. During performance of bilateral arm raising movements, oxyhemoglobin and total hemoglobin values in the primary sensorimotor cortex, premotor cortex, supplementary motor area, and prefrontal cortex were similar, but higher in these regions than those in the prefrontal cortex. We observed activation of the arm somatotopic areas of the primary sensorimotor cortex and premotor cortex in both hemispheres during bilateral arm raising movements. According to this result, bilateral arm raising move-ments appeared to induce large-scale neuronal activation and therefore arm raising movements would be good exercise for recovery of brain functions.     

Comparing thalamocortical and corticothalamic microstructure and spatial reciprocity in the macaque ventral posterolateral nucleus (VPLc) and medial pulvinar.

The detailed morphology of thalamocortical (TC) and corticothalamic (CT) pathways connecting the ventral posterolateral nucleus (VPLc) with the primary somatosensory cortex (areas 3b and 1) and the thalamic pulvinar with the posterior parietal cortex (primarily area 7a), was compared. Each pathway processes information relevant to directed reaching tasks, but whereas VPLc receives its major input from the spinal cord and external environment, the primary afferent to the pulvinar is cortical. Using combined tracer and thick fixed slice procedures, the soma/dendritic morphology of TC neuron populations (with known destination) was shown to be quantitatively similar within VPLc and the pulvinar. This implies that differences in information processing in VPLc (a primary relay) and the pulvinar (an integrative thalamic nucleus) are not defined by a distinctive TC morphology, but rather by the connections of these neuron populations. Two morphologically distinct types of CT axon were observed within the medial pulvinar and VPLc. The more common "Type E" were fine, had boutons en passant and diffuse terminal bifurcations ending in masses of tiny boutons. "Type R" axons were thicker, smooth, and terminated in localised clusters of large terminal boutons. Each type had a unique pattern of termination reflecting a distinct action on target neuron populations. The spatial relationship between TC distribution territories and CT terminal fields was examined within the medial pulvinar and VPLc by using anterograde and retrograde tracers injected together within cortical areas 7a, and 3b/1, respectively. Spatial overlap was incomplete within both thalamic nuclei. Our findings show a more complex relationship between TC and CT neuron populations than previously demonstrated.     

Segregated pathways carrying frontally derived top-down signals to visual areas MT and V4 in macaques

The bottom-up processing of visual information is strongly influenced by top-down signals, at least part of which is thought to be conveyed from the frontal cortex through the frontal eye field (FEF) and the lateral intraparietal area (LIP). Here we investigated the architecture of multisynaptic pathways from the frontal cortex to the middle temporal area (MT) of the dorsal visual stream and visual area 4 (V4) of the ventral visual stream in macaques. In the first series of experiments, the retrograde trans-synaptic tracer, rabies virus, was injected into MT or V4. Three days after rabies injections, the second-order (disynaptically connected) neuron labeling appeared in the ventral part of area 46 (area 46v), along with the first-order (monosynaptically connected) neuron labeling in FEF and LIP. In the MT-injection case, second-order neurons were also observed in the supplementary eye field (SEF). In the next series of experiments, double injections of two fluorescent dyes, fast blue and diamidino yellow, were made into MT and V4 to examine whether the frontal inputs are mediated by distinct or common neuronal populations. Virtually no double-labeled neurons were observed in FEF or LIP, indicating that separate neuronal populations mediate the frontal inputs to MT and V4. The present results define that the multisynaptic frontal input to V4 arises primarily from area 46v, whereas the input to MT arises from not only area 46v but also SEF, through distinct FEF and LIP neurons. Segregated pathways from the frontal cortex possibly carry the functionally diverse top-down signals to each visual stream.     

The cortical activation pattern during bilateral arm raising movements

Bilateral arm raising movements have been used in brain rehabilitation for a long time. However, no study has been reported on the effect of these movements on the cerebral cortex. In this study, using functional near infrared spectroscopy(f NIRS), we attempted to investigate cortical activation generated during bilateral arm raising movements. Ten normal subjects were recruited for this study. f NIRS was performed using an f NIRS system with 49 channels. Bilateral arm raising movements were performed in sitting position at the rate of 0.5 Hz. We measured values of oxyhemoglobin and total hemoglobin in five regions of interest: the primary sensorimotor cortex, premotor cortex, supplementary motor area, prefrontal cortex, and posterior parietal cortex. During performance of bilateral arm raising movements, oxyhemoglobin and total hemoglobin values in the primary sensorimotor cortex, premotor cortex, supplementary motor area, and prefrontal cortex were similar, but higher in these regions than those in the prefrontal cortex. We observed activation of the arm somatotopic areas of the primary sensorimotor cortex and premotor cortex in both hemispheres during bilateral arm raising movements. According to this result, bilateral arm raising movements appeared to induce large-scale neuronal activation and therefore arm raising movements would be good exercise for recovery of brain functions.     

Thalamic projections to S-I in macaque monkey.

The organization of thalamic input to functionally characterized zones in primary somatosensory cerebral cortex (S-I) of macaque monkeys (Macaca mulatta) was investigated using the method of labelling by retrograde transport of horseradish peroxidase (HRP). It was found that the cell columns positioned at the posterior margin of the band of cortex representing a given body region receive thalamic input from a posterior level of the ventroposterior thalamic nucleus (VP), and that cell columns at successively more anterior positions within that band receive input from successively more anterior levels of VP. The extreme posterior and anterior margins of the S-I hand, foot and face areas receive input from neuron populations which are not as widely separated in the anteroposterior dimension of VP as the neurons projecting to the extreme anterior and posterior margins of the proximal limb and trunk representations in S-I. These characteristics of the organization of the projections from VP to S-I are consistent with the view that the body representations in VP and S-I have the same connectivity and differential submodality distribution; and with the idea that thalamocortical conncetions only exist between functionally equivalent neuron populations in VP and S-I.     

Cortical and subcortical distribution of middle and long latency auditory and visual evoked potentials in a cognitive (CNV) paradigm

OBJECTIVE: This study concerned sensory processing (post-stimulus late evoked potential components) in different parts of the human brain as related to a motor task (hand movement) in a cognitive paradigm (Contingent Negative Variation). The focus of the study was on the time and space distribution of middle and late post-stimulus evoked potential (EP) components, and on the processing of sensory information in the subcortical-cortical networks. METHODS: Stereoelectroencephalography (SEEG) recordings of the contingent negative variation (CNV) in an audio-visual paradigm with a motor task were taken from 30 patients (27 patients with drug-resistant epilepsy; 3 patients with chronic thalamic pain). The intracerebral recordings were taken from 337 cortical sites (primary sensorimotor area (SM1); supplementary motor area (SMA); the cingulate gyrus; the orbitofrontal, premotor and dorsolateral prefrontal cortices; the temporal cortex, including the amygdalohippocampal complex; the parietooccipital lobes; and the insula) and from subcortical structures (the basal ganglia and the posterior thalamus). The concurrent scalp recordings were obtained from 3 patients in the thalamic group. In 4 patients in the epilepsy group, scalp recordings were taken separately from the SEEG procedure. The middle and long latency evoked potentials following an auditory warning (S1) and a visual imperative (S2) stimuli were analyzed. The occurrences of EPs were studied in two time windows (200-300 ms; and over 300 ms) following S1 and S2. RESULTS: Following S1, a high frequency of EP with latencies over 200 ms was observed in the primary sensorimotor area, the supplementary motor area, the premotor cortex, the orbitofrontal cortex, the cingulate gyrus, some parts of the temporal lobe, the basal ganglia, the insula, and the posterior thalamus. Following S2, a high frequency of EP in both of the time windows over 200 ms was observed in the SM1, the SMA, the premotor and dorsolateral prefrontal cortex, the orbitofrontal cortex, the cingulate gyrus, the basal ganglia, the posterior thalamus, and in some parts of the temporal cortex. The concurrent scalp recordings in the thalamic group of patients twice revealed potentials peaking approximately at 215 ms following S1. Following S2, EP occurred with latencies of 215 and 310 ms, respectively. Following S1, separate scalp recordings in 4 patients in the epilepsy group displayed EP 3 times in the 'over 300 ms' time window. Following S2, EP were presented once in the '200-300 ms' time window and 3 times in the 'over 300 ms' time window. CONCLUSIONS: The SM1, the SMA, multiple sites of the frontal lobe, some parts of the temporal lobe, the cingulate gyrus, the basal ganglia, the insula, and the posterior thalamus all participate in a cortico-subcortical network that is important for the parallel cognitive processing of sensory information in a movement related task.     

Cortico-cortical projections of the motorcortical larynx area in the rhesus monkey

The efferent cortico-cortical projections of the motorcortical larynx area were studied in three rhesus monkeys (Macaca mulatta), using biotin dextranamine as anterograde tracer. Identification of the larynx area was made with the help of electrical brain stimulation and indirect laryngoscopy. Heavy projections were found into the surrounding ventral and dorsal premotor cortex (areas 6V and D), primary motor cortex (area 4), the homolog of Broca's area (mainly area 44), fronto- and parieto-opercular cortex (including secondary somatosensory cortex), agranular, dysgranular and granular insula, rostral-most primary somatosensory cortex (area 3a), supplementary motor area (area 6M), anterior cingulate gyrus (area 24c) and dorsal postarcuate cortex (area 8A). Medium projections could be traced to the ventrolateral prefrontal and lateral orbital cortex (areas 47L and O), the primary somatosensory areas 3b and 2, the agranular and dysgranular insula, and the posteroinferior parietal cortex (area 7; PFG, PG). Minor projections ended in the lateral and dorsolateral prefrontal cortex (areas 46V and 8B), primary somatosensory area 1 and cortex within the intraparietal sulcus (PEa) and posterior sulcus temporalis superior (TPO). Due to its close spatial relationship to the insula on the one hand and the premotor cortex on the other, the larynx area shows projections which, in some respects, are not typical for classical primary motor cortex.     

Interconnections within the postarcuate cortex (area 6) of the macaque monkey

Small amounts of horseradish peroxidase conjugated with wheat germ were injected in restricted parts of the postarcuate premotor area of the macaque monkey. It was found that regions of this area having different somatotopic representations are richly interconnected among them. This pattern of intra-areal connectivity was not observed in the precentral motor area. It appears therefore that the postarcuate area is organized according to anatomical principles which are different from those of the primary motor cortex.     

Impaired mesial frontal and putamen activation in Parkinson's disease: a positron emission tomography study.

Selection of movement in normal subjects has been shown to involve the premotor, supplementary motor, anterior cingulate, posterior parietal, and dorsolateral prefrontal areas. In Parkinson's disease (PD), the primary pathological change is degeneration of the nigrostriatal dopaminergic projections, and this is associated with difficulty in initiating actions. We wished to investigate the effect of the nigral abnormality in PD on cortical activation during movement. Using C15O2 and positron emission tomography (PET), we studied regional cerebral blood flow in 6 patients with PD and 6 control subjects while they performed motor tasks. Subjects were scanned while at rest, while repeatedly moving a joystick forward, and while freely choosing which of four possible directions to move the joystick. Significant increases in regional cerebral blood flow were determined with covariance analysis. In normal subjects, compared to the rest condition, the free-choice task activated the left primary sensorimotor cortex, left premotor cortex, left putamen, right dorsolateral prefrontal cortex and supplementary motor area, anterior cingulate area, and parietal association areas bilaterally. In the patients with PD, for the free-choice task, compared with the rest condition, there was significant activation in the left sensorimotor and premotor cortices but there was impaired activation of the contralateral putamen, the anterior cingulate, supplementary motor area, and dorsolateral prefrontal cortex. Impaired activation of the medial frontal areas may account for the difficulties PD patients have in initiating movements.