Inputs from motor and premotor cortex to the superior colliculus of the macaque monkey

In retrograde studies of corticotectal projections in the monkey using horseradish peroxidase (HRP), projections of the frontal lobes were found to originate not only from the frontal eye fields and prefrontal association cortex but also from both motor and premotor cortex. Even small HRP injections into the superficial layers of the superior colliculus yielded labelled cells in the agranular cortex (area 6) of the anterior bank of the arcuate sulcus. After large collicular injections affecting all layers, labelled cells were found in both motor and premotor cortex. This projection appeared to be topographically organized. Injections into the anterolateral parts of the superior colliculus labelled cells that were distributed within the presumed finger-hand--arm-shoulder representation, whereas after more caudal injections labelled cells occurred more in the presumed arm-trunk representation. The supplementary motor cortex was not found to contain labelled cells. The corticotectal cells in the motor cortex differed from those in the premotor cortex in their size distribution; the former being small, the latter both small and large. The functional significance of the motor and premotor input into the superior colliculus for sensory, and particularly visual, guidance of movements is discussed in view of a collicular role in the extrapersonal space representation and of its possible participation in steering arm and hand movements.     

Motor aspects of cue-related neuronal activity in premotor cortex of the rhesus monkey

Neurons in the premotor cortex of rhesus monkeys were studied under two conditions: (1) visuospatial cues were given to guide the amplitude, direction, and onset time of forearm movements or (2) physically identical visual cues were given when reward was contingent on withholding movement. Neurons with sustained activity following the cues were preferentially active when the cues triggered a movement. Thus, activity of certain neurons in this cortical field is linked to motor set, i.e. intention to make a movement in response to the cue, rather than the visual cue per se.     

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.     

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. © 1995 Wiley-Liss, Inc.     

A visuo‐somatomotor pathway through superior parietal cortex in the macaque monkey: cortical connections of areas V6 and V6A

This report addresses the connectivity of the cortex occupying middle to dorsal levels of the anterior bank of the parieto-occipital sulcus in the macaque monkey. We have previously referred to this territory, whose perimeter is roughly circumscribed by the distribution of interhemispheric callosal fibres, as area V6, or the ‘V6 complex’. Following injections of wheatgerm agglutinin conjugated to horseradish peroxidase (WGA-HRP) into this region, we examined the laminar organization of labelled cells and axonal terminals to attain indications of relative hierarchical status among the network of connected areas. A notable transition in the laminar patterns of the local, intrinsic connections prompted a sub-designation of the V6 complex itself into two separate areas, V6 and V6A, with area V6A lying dorsal, or dorsomedial to V6 proper. V6 receives ascending input from V2 and V3, ranks equal to V3A and V5, and provides an ascending input to V6A at the level above. V6A is not connected to area V2 and in general is less heavily linked to the earliest visual areas; in other respects, the two parts of the V6 complex share similar spheres of connectivity. These include regions of peripheral representation in prestriate areas V3, V3A and V5, parietal visual areas V5A/MST and 7a, other regions of visuo-somatosensory association cortex within the intraparietal sulcus and on the medial surface of the hemisphere, and the premotor cortex. Subcortical connections include the medial and lateral pulvinar, caudate nucleus, claustrum, middle and deep layers of the superior colliculus and pontine nuclei. From this pattern of connections, it is clear that the V6 complex is heavily engaged in sensory-motor integration. The specific somatotopic locations within sensorimotor cortex that receive this input suggest a role in controlling the trunk and limbs, and outward reaching arm movements. There is a secondary contribution to the brain's complex oculomotor circuitry. That the medial region of the cortex is devoted to tightly interconnected representations of the sensory periphery, both visual and somatotopic—which are routinely stimulated in concert—would appear to be an aspect of the global organization of the cortex which must facilitate multimodal integration.     

Sensory input to the motor fields of the agranular frontal cortex: a comparison of the precentral, supplementary motor and premotor cortex

Arm displacements applied to the passive, but awake monkey are powerful stimuli for activating neurones in somatotopically appropriate areas of the precentral cortex. We have found that neurones in medial area 6 (SMA) and in lateral area 6 (PMC) may likewise be activated by such kinesthetic stimuli, at latencies which are only slightly longer than in area 4. Confirming previous findings, PMC neurones were sometimes also responsive to visual stimuli. The 'somatosensory' cells in the SMA were found in the microexcitable zone of the more posterior part of the SMA from where motor effects were elicited in arm and trunk muscles. These sensory neurones tended to be clustered together and they were only exceptionally excited antidromically by peduncular stimulation. Thus, somatosensory signals have access to both the PMC and SMA suggesting that both areas may be implicated in sensory-guided or sensory-triggered movements.     

Covariation of distributions of callosal cell bodies and callosal axon terminals in layer III of cat primary auditory cortex

Primary auditory cortex in the cat is both the source and target of callosal fibers. Injection of horseradish peroxidase (HRP) in the high frequency representation of AI in one hemisphere retrogradely labels callosal cell bodies and anterogradely labels callosal axon terminals in AI of the opposite hemisphere. In tissue sections cut through layer III parallel to the cortical surface, elongated patches composed of dense aggregates of callosal cell bodies and callosal axon terminals alternate with regions containing lower concentrations of these elements. Labeling in AI is most dense in regions corresponding to the frequency representation of the injected site. In layer III of the densely labeled region, patches of high concentrations of labeled callosal axon terminals correspond with high concentrations of labeled callosal cell bodies. On the other hand, little correspondence is apparent between the distributions of the two elements in layer III in the sorrounding area of lighter labeling. Layers V and VI contain relatively few labeled callosal axon terminals and cell bodies, and our data do not suggest whether the two distributions covary in these layers.     

Effects of premotor cortex cooling upon visually initiated hand movements in the monkey

The premotor cortex was temporarily impaired with a local cooling method and effects of the cooling upon visually initiated hand movement and upon cortical field potentials associated with the movement were examined in the monkey. Bilateral cooling of the premotor cortex (the dorsolateral part of presumed area 6) disorganized the well-trained reaction-time movement, in which a lever was lifted within duration of the light stimulus (about 0.5 s) delivered at random time intervals. Accordingly the monkey showed nearly self-paced, random movements with little regard to the light stimulus during the cooling. To obtain a certain number of appropriate reaction-time movements, about twice as many as trials in normal conditions were required during the premotor cooling. Unilateral (contralateral to the moving hand) cooling of the premotor cortex produced similar but weak effects. No appreciable paresis was observed by cooling. Such effects of cooling the premotor cortex faded in successive experimental days of several weeks or even in successive cooling sessions on the same day. This was in contrast with effects of motor cortex cooling which were reported to be almost unfaded on repetition. It is suggested that compensatory actions are quickly elicited in some other structure even on temporal impairment of the premotor cortex and that the actions are gradually accumulated on repetitive cooling with days and weeks.     

胼胝体梗死导致的异手综合征(3例报告和文献复习)

目的:分析异手综合征(AHS)的发病特点,探讨胼胝体梗死与AHS的关系。方法:对胼胝体梗死患者进行前瞻性的研究。结果:我院2004至2006年收治的1860例住院脑梗死患者中有25例胼胝体梗死,约占同期脑梗死患者的1.34%(25/1860)。有3例患者出现AHS,占0.16%(3/1860),占胼胝体梗死患者的12%(3/25)。属于前型AHS2例,后型AHS1例;3例患者均有双手矛盾动作,1例有左侧失用和拟人格化;1例有未受累侧肢体的模仿动作。结论:高血压、严重的动脉粥样硬化是胼胝体梗死并发AHS主要原因,双手的矛盾动作是AHS常见表现。     

Relationship between the corticostriatal terminals from areas 9 and 46, and those from area 8A, dorsal and rostral premotor cortex and area 24c: an anatomical substrate for cognition to action

Our previous data indicate that there are specific features of the corticostriatal pathways from the prefrontal cortex. First, corticostriatal pathways are composed of focal, circumscribed projections and of diffuse, widespread projections. Second, there is some convergence between terminal fields from different functional regions of the prefrontal cortex. Third, anterior cingulate projections from area 24b occupy a large region of the rostral striatum. The goal of this study was to determine whether these features are also common to the corticostriatal projections from area 8A (including the frontal eye field; FEF), the supplementary eye field (SEF), dorsal and rostral premotor cortex (PMdr) and area 24c. Using a new approach of three-dimensional reconstruction of the corticostriatal pathways, along with dual cortical tracer injections, we mapped the corticostriatal terminal fields from areas 9 and 46, 8A-FEF, SEF, PMdr and 24b and c. In addition, we placed injections of retrogradely transported tracers into key striatal regions. The results demonstrated that: (i) a diffuse projection system is a common feature of the corticostriatal projections from different frontal regions; (ii) key striatal regions receive convergent projections from areas 9 and 46 and from areas 8A-FEF, SEF, PMdr and 24c, suggesting a potential pivotal role of these striatal regions in integrating cortical information; (iii) projections from area 24c, like those from area 24b, terminate widely throughout the striatum, interfacing with terminals from several frontal areas. These features of the corticostriatal frontal pathways suggest a potential integrative striatal network for learning.     

Callosal connections of the posterior neocortex in normal-eyed, congenitally anophthalmic, and neonatally enucleated mice

Following multiple injections of horseradish peroxidase into the posterior neocortex of one hemisphere, we examined the distribution of retrogradely labeled cells and anterogradely labeled terminations in tangential and coronal sections through contralateral areas 17 and 18 in three groups of adult mice: normal-eyed (ZRDCT-n and C57Bl/6J strains), congenitally anophthalmic (ZRDCT-an strain), neonatally enucleated (ZRDCT-n strain). In agreement with previous studies, we observed that the pattern of callosal connections in areas 17 and 18 of normal-eyed mice contains the following features: (1) a dense band of callosal cells and terminations separating the interiors of areas 17 and 18, which have relatively few callosal connections, (2) a ring-like configuration anterolateral to area 17, (3) a region of dense labeling lateral to area 18, (4) a narrow band of labeling bridging the posterior portion of area 18, and (5) a region of labeling anteromedial to area 17. We find that all these features of the normal callosal pattern are recognizable in congenitally anophthalmic mice. Their presence in mice that never had eyes supports the hypothesis that central visual pathways can develop many aspects of their connectivity in the absence of input from the periphery. However, we also find that the details of certain features of the callosal pattern in congenitally eyeless mice often differ from those of the same features in normal-eyed mice, and that the between-animal variability in the appearance of these features is higher in eyeless mice. These latter findings indicate that the eyes are needed during normal development to fine-tune the pattern of callosal connections. Our results also reveal that the callosal pattern in neonatally enucleated mice does not differ significantly from that in congenitally anophthalmic mice, indicating that the period in which the eyes guide callosal development extends into postnatal life. While the present data do not delineate the time course of this period, the finding of similarly abnormal callosal patterns in congenitally anophthalmic and neonatally enucleated mice suggests that the eyes exert little if any influence prenatally. Finally, examination of coronal sections indicates that the laminar distribution of callosal connections develops normally in both groups of eyeless mice.     

Ventral premotor cortex neuronal activity matches perceptual decisions

The relationship between neuronal activity and psychophysical judgments is central to understanding the brain mechanisms responsible for perceptual decisions. The ventral premotor cortex is known to be involved in representing different components of the decision-making process. In this cortical area, however, neither the neuronal ability to discriminate nor the trial-to-trial relationship between neuronal activity and behavior have been studied during visual decision-making. We recorded from single neurons while monkeys reported a decision based on the comparison of the orientation of two lines shown sequentially and separated by a delay. Analyses based on signal detection theory provided both the behavioral and neuronal sensitivities (d') and the coherence between behavioral and neuronal choices. To determine the temporal evolution of neuronal sensitivity and of coherence, the optimal size and position of the encoding windows were assessed. For a subset of neurons from the premotor ventral cortex, neuronal sensitivity was close to behavioral sensitivity and the trial-to-trial coherence between the neuronal and behavioral choices was close to 100%. By comparing these results with those obtained in a motor control task we ruled out the possibility of this activity being explained by the motor component of the task. These results suggest that activity in the ventral premotor cortex explains behavioral performance and predicts trial-to-trial subject choices.     

The involvement of monkey premotor cortex neurones in preparation of visually cued arm movements

Some neurones in macaque postarcuate premotor area modulate their firing frequency in relation to motor tasks which require visual information. We previously reported that a large proportion of these neurones modulate during execution of a detour reaching task in which the movement phase was separated in time from the phase in which the monkey received a visual cue for the movement required to retrieve a food reward. A large proportion of task-related neurones (75%) modulated during this 'visual' phase, in which no task-related movements were made. This modulation was related to the position of the food reward, which served as the visual cue. Most of these neurones were located in cortical area 6, close to the arcuate curvature and its spur, but also more caudally in area 4 and rostrally in area 8. In the present chronic recording experiments in monkeys, several variations of the original task were used in order to test whether the 'visual'-related neuronal modulation could be involved in preparation of the upcoming movement. This modulation is unlikely to be related to any eye or arm movements occurring during the visual phase or to changes in environmental illumination. Neither can it be related to the presence of the visual cue in a particular part of the visual field, since the pattern of neuronal modulation was similar when a cue with a fixed position was used. This modulation was, however, contingent upon the occurrence of food retrieval during the subsequent 'movement phase', since it was abolished or diminished during presentation of a 'food-reward' which the monkey did not retrieve. For several neurones, modulation pattern during the visual phase depended on whether the food reward was to be retrieved with a gross hand movement or with relatively independent finger movements. It is likely, therefore, that neurones in the postarcuate premotor cortex are involved in preparation for arm movements with the help of visual cues. The results are discussed in view of corticocortical pathways which might be involved in transmission of visual information from visual areas through parietal association areas and premotor cortex to the primary motor cortex.     

Interhemispheric connections in the visual cortex of the squirrel monkey (Saimiri sciureus)

The callosal connections within the posterior parietal and occipital cortices were studied in the squirrel monkey with horseradish peroxidase tracing techniques. The data were evaluated with particular emphasis on the relationship of major callosal connections along the 17-18 border. The overall pattern of callosal connections in the squirrel monkey also was compared with callosal patterns in other New World simians. Our results show that the dense band of callosal connections along the 17-18 border in the squirrel monkey differs from the connections observed in other New World monkeys in that it is virtually confined to area 18 and avoids area 17. In addition to a continuous band of callosal connections in area 18 that parallels the 17-18 border, rostral extensions of the band are oriented perpendicular to the 17-18 border and present an obvious periodicity. The remaining parieto-occipital cortex contains a complex pattern of callosal connections that is strikingly similar to patterns reported for other New World monkeys. Thus, it is likely that the dorsolateral extrastriate visual cortex in the squirrel monkey is organized in a manner similar to that found within other New World monkeys.     

The relationship of corpus callosum connections to electrical stimulation maps of motor, supplementary motor, and the frontal eye fields in owl monkeys

Microstimulation and anatomical techniques were combined to reveal the organization and interhemispheric connections of motor cortex in owl monkeys. Movements of body parts were elicited with low levels of electrical stimulation delivered with microelectrodes over a large region of precentral cortex. Movements were produced from three physiologically defined cortical regions. The largest region, the primary motor field, M-I, occupied a 4-6-mm strip of cortex immediately rostral to area 3a. M-I represented body movements from tail to mouth in a grossly somatotopic mediolateral cortical sequence. Specific movements were usually represented at more than one location, and often at as many as six or seven separate locations within M-I. Although movements related to adjoining joints typically were elicited from adjacent cortical sites, movements of nonadjacent joints also were produced by stimulation of adjacent sites. Thus, both sites producing wrist movements and sites producing shoulder movements were found next to sites producing digit movements. Movements of digits of the forepaw were evoked at several locations including a location rostral to or within cortex representing the face. Overall, the somatotopic organization did not completely correspond to previous concepts of M-I in that it was neither a single topographic representation, nor two serial or mirror symmetric representations, nor a "nesting about joints" representation. Instead, M-I is more adequately described as a mosaic of regions, each representing movements of a restricted part of the body, with multiple representations of movements that tend to be somatotopically related. A second pattern of representation of body movements, the supplementary motor area (SMA), adjoined the rostromedial border of M-I. SMA represented the body from tail to face in a caudorostral cortical sequence, with the most rostral portion related to eye movements. Movements elicited by near-threshold levels of current were often restricted to a single muscle or joint, as in M-I, and the same movement was sometimes multiply represented. Typically, more intense stimulating currents were required for evoking movements in SMA than in M-I. A third motor region, the frontal eye field (FEF), bordered the representation of eyelids and face in M-I. Eye movements elicited from this cortex consisted of rapid horizontal and downward deviation of gaze into the contralateral visual hemifield.     

The ipsilateral cortico-cortical connexions between the cytoarchitectonic subdivisions of the primary somatic sensory cortex in the monkey

The callosal connexions of the primary somatic sensory cortex, SI, of the monkey have been studied with axonal degeneration methods after the placement of lesions of varying size in the cortex of one hemisphere and after section of the corpus callosum. For the correlation of the distribution of the degeneration with the cytoarchitectonic subdivisions of SI and with their boundaries, planar reconstructions of the extents of the subdivisions and of area 5 were made. The extent of area 5 is surprisingly large, being about the same as SI, and area 3a can be recognized as a distinct subdivision along the entire mediolateral extent of SI. The callosal fibres end in narrow, irregular bands aligned in the medio-lateral dimension and there are accentuations at the boundaries of the cytoarchitectural subdivisions. In the representations of the trunk and face, the bands of degeneration are present across the entire antero-posterior extent of SI and with increases at the boundaries, while in the limb regions the degeneration becomes restricted to the boundaries. It is suggested that the callosal connexions of the somatic sensory cortex, like those in the visual and auditory areas, are connecting those parts of the cortex in the two hemispheres that are concurrently activated by a peripheral stimulus. The parts of SI that are devoid of callosal connexions are related to the distal limbs. The callosal connexions are homo- and heterotopical; an architectonic subdivision within the callosally connected regions projects to the same and other architectonic subdivisions at the same medio-lateral level in the opposite hemisphere; the cortex containing the representation of the caudal trunk near the post-central dimple is connected with the same region in the other hemisphere and with that of the separate representation of the caudal trunk in the posterior part of the cingulate sulcus, while the representation of the occipital region at the post-central dimple is connected both with the homotopical site in the other hemisphere and with the other representation of this part of the periphery at the level of the lower end of the intraparietal sulcus.     

The primate premotor cortex fifty years after Fulton

Fifty years ago, F.M.R. Walshe launched his famous diatribe against J.F. Fulton and his thesis that a premotor area with specialized functions exists within the agranular frontal cortex of primates. Now there is general agreement that the agranular frontal cortex contains at least 3 cortical fields: the primary motor cortex, the supplementary motor cortex, and the premotor cortex. This paper summarizes some of the evidence that the premotor cortex plays a role in the preparation for limb movements and presents speculation concerning the aspects of motor preparation in which the premotor cortex might be involved.     

Callosal connections of the cortical taste area in rats

The granular and dysgranular insular subregions of the cortical taste area in rats are shown to connect anatomically with the homotopical regions in the opposite hemisphere through the corpus callosum. Cells of callosal efferents and terminals of callosal fibers were found in almost all cortical layers. The findings clarify the current understanding of the morphological substrate of callosal interactions in the gustatory system.     

Interhemispheric connections of the visual cortex in the grey squirrel (Sciurus carolinensis)

The total pattern of visual callosal connections was studied in the grey squirrel by using the Fink-Heimer technique for axonal and terminal degeneration and the autoradiographic and horseradish peroxidase techniques for axonal transport. The pattern of terminations was correlated with architectonic landmarks. The results show that callosal terminations are distributed in a complex fashion within the visual cortical areas. The major terminations form a band in area 17 along its border with area 18. This band is contiguous rostrally with the callosal terminations in area L that extend caudomedially onto the medial wall of the hemisphere. Caudally the band in area 17 wraps around the ventral aspect of the occipital pole and ends medially at the level of the hippocampus. This band exhibits a distinct periodicity in the density of terminations. The callosal terminations in area 18 are usually found along the lateral and medial borders and are concentrated in discrete patches. The pattern in area 19 exhibits two or three primary patches and only loosely corresponds to the borders of the area. Few callosal terminations are found in area 19p and the posterior temporal area, Tp, while the intermediate temporal area, Ti, receives an extensive input. The laminar distribution of callosal terminations is different in each area studied. Characteristically, area 18 has dense terminations in layers III, II, and the inner one-half of layer I, with less dense terminations in layers V and VI, and sparse terminations in layer IV. Area 17 has a similar pattern in the supragranular and infragranular layers but also has dense terminations in layer IV. The patterns in area 19 are intermediate between these extremes but are more similar to those in area 17. The cells that give rise to the callosal projections were found primarily in layers III and V and occasionally in layers II, IV, and VI. The distribution of the callosal efferent neurons is more extensive than the areas of terminations. The distribution of callosal terminations suggests that the organization of visual cortical areas in the grey squirrel is more complex than had been previously recognized. This finding is discussed with reference to the general organization of the mammalian visual cortical areas, and a need for more extensive analyses of visual cortical areas in the grey squirrel, particularly with respect to extrastriate visual areas, is indicated.