Optokinetic nystagmus deficits following parieto-occipital cortex lesions in monkeys

Summary Optokinetic nystagmus (OKN) was induced in six monkeys by rotation of a full field drum. After unilateral lesions of the inferior parietal lobule and prestriate cortex (IPL-PS lesion), three monkeys had diminished velocity of slow components when the drum rotated toward the side of the lesion. OKN slow components appeared normal when the drum rotated in the opposite direction, while fast components appeared normal in both directions. The severity of the slow component deficit was greater at higher rates of stimulus rotation than at lower ones. Recovery occurred within 7–10 days for two monkeys; no recovery was evident after 2 weeks for the third. Subsequent bilateral IPL-PS lesions in two of these monkeys reduced OKN slow phase velocity in both directions. Two monkeys with unilateral lesions limited to the inferior parietal lobule (IPL lesions) had only very mild and transient deficits lasting 1–3 days. Bilateral IPL lesions also produced only slight OKN deficits. One monkey had a lesion which destroyed most of the lateral striate cortex of one hemisphere and had no discernable OKN deficit. These results demonstrate that, in contrast to earlier reports, lesions of parieto-occipital cortex in monkeys produce deficits which are qualitatively similar to, although of shorter duration than, those OKN deficits which commonly are associated with posterior parietal damage in humans.Supported by NIH grants EY-2640, EY-5318, 5-S01-RR-5530-14, and a grant from the Mayo Foundation     

Enhanced visual spatial attention ipsilateral to rTMS-induced 'virtual lesions' of human parietal cortex

The breakdown of attentional mechanisms after brain damage can have drastic behavioral consequences, as in patients suffering from spatial neglect. While much research has concentrated on impaired attention to targets contralateral to sites of brain damage, here we report the ipsilateral enhancement of visual attention after repetitive transcranial magnetic stimulation (rTMS) of parietal cortex at parameters known to reduce cortical excitability. Normal healthy subjects received rTMS (1 Hz, 10 mins) over right or left parietal cortex. Subsequently, detection of visual stimuli contralateral to the stimulated hemisphere was consistently impaired when stimuli were also present in the opposite hemifield, mirroring the extinction phenomenon commonly observed in neglect patients. Additionally, subjects' attention to ipsilateral targets improved significantly over normal levels. These results underline the potential of focal brain dysfunction to produce behavioral improvement and give experimental support to models of interhemispheric competition in the distributed brain network for spatial attention.     

Restoration of visual orienting into a cortically blind hemifield by reversible deactivation of posterior parietal cortex or the superior colliculus

A contralateral hemineglect of the visual field can be induced by unilateral cooling deactivation of posterior middle suprasylvian (pMS) sulcal cortex of the posterior parietal region, and this neglect can be reversed by additional cooling deactivation of pMS cortex in the opposite hemisphere. The purpose of the present study was to test whether an enduring hemianopia induced by removal of all contiguous visual cortical areas of one hemisphere could be reversed by local cooling of pMS cortex in the opposite hemisphere. Two cats sustained large unilateral ablations of the contiguous visual areas, and cooling loops were placed in the pMS sulcus, and in contact with adjacent area 7 or posterior ectosylvian (PE) cortex of the opposite hemisphere. In both instances cooling of pMS cortex, but neither area 7 nor PE, restored a virtually normal level of orienting performance to stimuli presented anywhere in the previously hemianopic field. The reversal was highly sensitive to the extent of cooling deactivation. In a third cat, cooling deactivation of the superficial layers of the contralateral superior colliculus also restored orienting performance to a cortical ablation-induced hemianopia. This reversal was graded from center-to-periphery in a temperature-dependent manner. Neither the cortical ablation nor any of the cooling deactivations had any impact on an auditory detection and orienting task. The deactivations were localized and confirmed by reduced uptake of radiolabeled 2-deoxyglucose to be limited to the immediate vicinity of each cooling loop. The results are discussed in terms of excitation and disinhibition of visual circuits.     

Selective visual neglect in right brain damaged patients with splenial interhemispheric disconnection

Left unilateral neglect is frequently reported after right hemispheric lesions of the middle cerebral artery (MCA) damaging the parietal–frontal cortical–subcortical network subserving space representation and awareness. However, accumulating evidence shows that neglect can also follow lesions of the posterior cerebral artery (PCA) that do not directly affect this parietal–frontal network. Surgical studies in the monkeys have demonstrated that complete callosal resection combined with lesion of the right optic tract entirely deprives the right hemisphere of visual inputs from the left hemispace provoking severe left unilateral neglect. Here, through the detailed study of two patients we show, for the first time, that PCA lesions selectively affecting the splenium of the corpus callosum and the adjacent right primary visual cortex provoke severe neglect selectively restricted to the visual domain. No trace of personal, motor or representational-imagery neglect was found. Also at variance with previous case studies in which neglect followed lesion of the trunk or the genu of the corpus callosum, no restriction of neglect to tasks performed with the right hand, no left hemispatial limb akinesia, no tactile extinction for the left hand and no tactile anomia for stimuli explored with the left hand were observed. These findings demonstrate that brain lesions depriving intact parietal and frontal attentional areas from specific sensory inputs can yield spatial neglect limited to specific sensory modalities or sectors of space.     

Hemiextinction induced by transcranial magnetic stimulation over the right temporo-parietal junction

Whereas it is widely accepted that the parietal cortex is crucial for visual attention, the role of the temporal cortex and the temporo-parietal junction (TPJ) is less clear. There are clinical reports of patients with lesions in different posterior temporal areas which exhibit contralateral visual neglect but this syndrome seems to be less frequent than in patients with parietal lesions. In a previous study, we could show that single-pulse transcranial magnetic stimulation (TMS) over the right inferior parietal cortex is capable to induce both neglect-like and extinction-like impairments of performance in normal subjects. In the present study, we used this method to examine the functional role of the superior temporal gyrus (STG) and the TPJ of the right hemisphere for visuo-spatial attention. Healthy volunteers were asked to detect small dots appearing for 40 ms unilaterally on right or left side or bilaterally on a computer screen. TMS was applied over the TPJ or STG. TMS over the TPJ induced an extinction-like behavioral pattern to the contralateral hemifield. TMS over the STG had no effect. The results demonstrate a functional involvement of the TPJ in visuo-attentional processing of competing stimuli in both hemifields. This region is part of the cortical network mediating stimulus-driven attention which is relevant for processing of competing stimuli.     

Unilateral neglect: a theory of proprioceptive space of a stimulus as determined by the cerebellar component of motor efference copy (and is autism a special case of neglect)

Unilateral neglect is a devastating condition, which manifests as a loss of a person's spatial awareness opposite the damaged side of the brain. It challenges our conception of the seat of the soul and its explanation is at the heart of the mind-body problem. A heuristic definition of the dorsal stream of a modality is here based on the categorization of parietal networks by the cerebellar component of motor efference copy. Taking this premise, the proprioceptive space of a stimulus is established as a concept in this paper. It is proposed that unilateral neglect is typically a dysfunction of proprioceptive space of a stimulus associated with lesions of the dorsal stream. Furthermore, most experimental findings of unilateral visual neglect (and by extrapolation, other sensory modalities), can be explained by two developmental mechanisms by which the proprioceptive space of a stimulus is encoded in the parietal cortex. Its right and left hemisphere representation can be dissociated from the hemifield of presentation of perceptual information, such that the left hemifield can have a left hemisphere representation through callosal connections and likewise, the right hemifield can have a right hemifield representation. The processing of a sensory stimulus in either parietal hemisphere is dynamically determined as shown by experimental modulation of performance. A theory of historical precedence will provide a developmental background to the organization of proprioceptive space and will invoke separate models according to specified terms of engagement. A model based on the expansion and contraction of the proprioceptive space of a stimulus as a gradient across both hemispheres and modulated by concurrent proprioceptive state will be differentiated from a model that is non-graduating but competitive and lacks such modulation. In other words, the dorsal representation of a sensory stimulus in the former case is shared to a varying degree by the two parietal hemispheres, whereas in the latter case the representation of left and right aspects of the same object stimulus is strictly divided between the two hemispheres. A further hypothesis of a dorsoventral gradient of the peripheral and foveal components of proprioceptive space characterizing dorsal stream networks will predict the double dissociation revealed by experimental paradigms. It will explain why some patients show neglect only in foveal while others only in peripheral vision. The paper proposes to unify neglect, extinction and optic ataxia on the one hand, and spatial and object-based neglect on the other hand, under a singularly proficient paradigmatic structure. A binding model as described is a component theory that acknowledges how the involved pathways or transitional zones in a pathway may contribute to a differential clinical picture as one progresses from posterior to anterior parietal cortex. Finally, a brief discussion is given on how autistic subjects neglect spatial cues and the inability of a spatial cognitive transformation underlies the impairments postulated for 'a theory of mind'.     

Bilateral parietal and contralateral responses during maintenance of unilaterally encoded objects in visual short-term memory: Evidence from magnetoencephalography

A component of the event-related magnetic field (ERMF) response was observed in magnetoencephalographic signals recorded during the maintenance of information in visual short-term memory (VSTM). This sustained posterior contralateral magnetic (SPCM) field is likely the magnetic equivalent of the sustained posterior contralateral negativity (SPCN) found in electrophysiology. Magnetoencephalography data showed, at the sensor level, a bilateral activation over the parietal cortex that increased in amplitude for higher memory load. Others sensors, also over the parietal cortex, showed an activation pattern similar to the SPCN with higher activation for the hemisphere contralateral to the visual field from which visual information was encoded. These two activation patterns suggest that the SPCN and SPCM are generated by a network of cortical sources that includes bilateral parietal loci, likely intra-parietal/intra-occipital cortex, and contralateral parietal sources.     

Remapping the remembered target location for anti-saccades in human posterior parietal cortex

We used functional magnetic resonance imaging (fMRI) to investigate the role of the human posterior parietal cortex (PPC) in anti-saccades. To do so, we exploited the laterality of a subregion of the PPC for remembered target location. Using an event-related design, we tracked fMRI signal changes in this region while subjects remembered the location of a flashed target, then were instructed to plan either an anti- or pro-saccade to that location, and finally were instructed to execute the movement. At first, the region responded preferentially to the memory of a target location presented in the contralateral visual field. However, when an anti-cue specified a saccadic response into the opposite visual field, we observed a dynamic shift in cortical activity from one hemisphere to the other. This shows that this region within the human posterior parietal cortex codes the target location for an upcoming saccade, rather than the location of the remembered visual stimulus in an anti-saccade task.     

Central core control of developmental plasticity in the kitten visual cortex: I. Diencephalic lesions

Summary In five, dark-reared, 4-week-old kittens the posterior two thirds of the corpus callosum were split, and a lesion comprising the intralaminar nuclei was made of the left medial thalamic complex. In addition, the right eye was closed by suture. Postoperatively, the kittens showed abnormal orienting responses, neglecting visual stimuli presented in the hemifield contralateral to the side of the lesion. Sudden changes in light, sound, or somatosensory stimulation elicited orienting responses that all tended toward the side of the lesion. These massive symptoms faded within a few weeks but the kittens continued to neglect visual stimuli in the hemifield contralateral to the lesion when a second stimulus was presented simultaneously in the other hemifield. Electrophysiologic analysis of the visual cortex, performed after the end of the critical period, revealed marked interhemispheric differences. In the visual cortex of the normal hemisphere most neurons were monocular and responded exclusively to stimulation of the open eye, but otherwise had normal receptive field properties. In the visual cortex of the hemisphere containing the thalamic lesion, the majority of the neurons remained binocular. In addition, the selectivity for stimulus orientation and the vigor of responses to optimally aligned stimuli were subnormal on this side. Thus, the same retinal signals, which in the control hemisphere suppressed the pathways from the deprived eye and supported the development of normal receptive fields, failed to do either in the hemisphere containing the thalamic lesion. Apparently, experience-dependent changes in the visual cortex require both retinal stimulation and the functioning of diencephalic structures which modulate cortical excitability and control selective attention.     

Influences of lesions of parietal cortex on visual spatial attention in humans

Summary Several brain areas have been identified with attention, because damage to these regions leads to neglect and extinction. We have tested elements of visual attentional processing in patients with parietal, frontal, or temporal lesions and compared their responses to control subjects. Normal humans respond faster in a reaction time task when the spatial location of a target is correctly predicted by an antecedent stimulus (valid cue) than when the location is incorrectly predicted (invalid cue). The cue is hypothesized to shift attention towards its location and thereby facilitate or impede response latencies. The reaction times of individuals with damage to the parietal lobe are somewhat slowed for targets ipsilateral or contralateral to the side of the lesion if the targets are preceded by valid cues. These same patients are extremely slow in responding to targets in the visual field contralateral to the lesion when the cue has just appeared in the unaffected (ipsilateral) visual field. In addition, these individuals are especially slow in responding to targets in either visual field when the lights are preceded by weak, diffuse illumination of the entire visual field. Patients with lesions of the frontal lobe have very slow reaction times in general and, as is the case for patients with lesions of the temporal lobe, are slow in all conditions for targets in the field contralateral to the lesion. These patterns are probably not associated with attentional defects. For patients with parietal lesions, these studies demonstrate a further deficit in a cued reaction-time task suggesting abnormal visual attention. Since different sites of brain damage yield different patterns of responses, tests such as these could be of analytic and diagnostic value.     

Control of fixation and saccades in humans with chronic lesions of oculomotor cortex

To elucidate the dynamic interactions of cortical and subcortical oculomotor systems, the authors investigated reflexive and strategic control over fixation and saccades in patients with chronic unilateral lesions that involved either frontal or parietal cortex. They measured the effects of indicating the location of the forthcoming target and removing the fixation stimulus on the latencies of eye movements toward a peripheral visual target in 12 patients with frontal eye field (FEF) lesions, 9 patients with lesions restricted to parietal cortex, and 12 neurologically healthy controls. They found that chronic damage to FEF cortex disrupts cortico-collicular interactions, resulting in hypoactivity in the contralesional superior colliculus and a loss of strategic control over the intrinsic collicular circuits that regulate fixation.     

Organization of corticopontocerebellar connections to the paramedian lobule in the cat

Summary To reveal the organization and relative magnitude of connections from various parts of the cerebral cortex to the cerebellar paramedian lobule via the pontine nuclei, horseradish peroxidase conjugated to wheat germ agglutinin was injected in the paramedian lobule in conjunction with injection of the same tracer in various parts of the cerebral cortex in 14 cats. Termination areas of cortical fibres (anterogradely labelled) and pontine neurons projecting to the paramedian lobule (retrogradely labelled) were carefully plotted in serial sections through the pons. On the average 89% of all labelled cells were found in the pontine nuclei contralateral to the cerebellar injection, 11% in the ipsilateral pontine nuclei. The highest degree of overlap between anterograde and retrograde labelling was found after injections in the posterior sigmoid gyrus (SmI), while less overlap was found after injections of the anterior sigmoid gyrus (MsI). Injections of the second somatosensory area (SmII) and the parietal association cortex (areas 5 and 7) gave moderate degrees of overlap. Very little or no overlap was found after injections of the premotor cortex (area 6), the visual areas 17, 18 and 19 and the auditory cortex (AI and AII).It is concluded that a major cortical input to the paramedian lobule arises in the posterior sigmoid gyrus (SmI), but that additional contributions arise in the anterior sigmoid gyrus (MsI), the parietal areas 5 and 7 and the second somatosensory cortex (SmII). Among the latter regions probably the parietal areas contribute most. Overlap between terminal regions of cortical fibres and cells projecting to the paramedian lobule takes place at numerous discrete sites at virtually all rostrocaudal levels of the pons. Cerebrocortical afferents via the pontine nuclei to the intermediate zone of the posterior lobe are organized according to the same principles as described previously for cortical afferents to the hemispheral parts of the posterior lobe (crus I and II).     

Evidence for interhemispheric processing of inputs from the hands in human S2 and PV

In the present investigation, we identified cortical areas involved in the integration of bimanual inputs in human somatosensory cortex. Using functional magnetic resonance imaging (fMRI) and magnetoencephalography (MEG), we compared the responses to unilateral versus bilateral stimulation in anterior parietal cortex and areas in the Sylvian fissure of the contralateral hemisphere. The extent of fMRI activation on the upper bank of the Sylvian fissure, in the second somatosensory (S2) and the parietal ventral (PV) areas, was significantly larger for bilateral stimulation than for unilateral stimulation. Using MEG, we were able to describe the latency of response in S1 and S2/PV to unilateral and bilateral stimulation. The MEG response had three components under both stimulus conditions. An early peak in S1 at 40 ms, a middle peak in S2/PV at 80-160 ms, and three late peaks in S2/PV at 250-420 ms. There was an increase in magnetic field strength in S2/PV to bilateral stimulation at 300-400 ms post stimulus. The fMRI results indicate that, as in monkeys, S2/PV receives inputs from both the contralateral and ipsilateral hand. The MEG data suggest that information is processed serially from S1 to S2. The very late response in S2/PV indicates that extensive intrahemispheric processing occurs before information is transferred to the opposite hemisphere. The neural substrate for the increased activation and field strength at long latencies during bilateral stimulation can be accounted for in three ways. Under bilateral stimulus conditions, more neurons may be active, neuronal firing rate may increase, and/or neural activity may be more synchronous.     

Effects of posterior parietal lesions on visually guided movements in monkeys

Summary Four monkeys were trained to position, with either hand, a vertical rod in front of one of 5 target lights spaced 20° apart on a semicircular screen. After the monkeys had reached the preoperative criterion (80% trials correct per session) they received a 1- or 2-stage bilateral lesion of posterior parietal cortex restricted to area 7. The lesion produced in all the monkeys considerable but temporary changes in movement latency, accuracy, velocity and duration. Latency increase appeared to be independent of changes in the other parameters. After the first lesion, movement latency increased for the contralateral arm in both left and right working spaces, from 100 ms up to 400 ms depending on the animal. A second lesion symmetrical to the first one increased movement latency of the arm contralateral or ipsilateral to the last lesion, depending on the time interval between the two lesions. In addition, unilateral lesions of area 7 induced a gross inaccuracy in movements of the arm contralateral to the lesion, more marked in the contralateral working space. These lesions also increased movement peak velocity and simultaneously decreased movement duration for the arm contralateral to the lesion. The increase in velocity appeared to be related to the decrease in duration. A second lesion of area 7 in the opposite hemisphere similarly affected accuracy, velocity and duration but for the arm contralateral to the second lesion.     

Anti-extinction Following Unilateral Parietal Damage

Following unilateral damage to the parietal cortex, extinction of sensory events on the side of space opposite the lesion frequently occurs: A contralesional event is detected when it occurs alone, but missed when it is accompanied by an ipsilesional event. We describe a patientwith left visuospatial neglect subsequent to a right parietal infarct, who shows the opposite of this typical extinction pattern: On brief visual presentations, he consistently shows better detection and identification of contralesional targets when they appear simultaneously with an ipsilesional target, compared to when the contralesional target appears alone. This benefitfor contralesional targets on simultaneous presentation is found most clearly when the same task is performed with both contra- and ipsilesional items. These findings first provide a very clear demonstration of neglect in the absence of extinction. Second, our results demonstrate that visuospatial neglect can be mitigated by non-perceptual task variables.     

Visually guided locomotion, distractibility, and the missing-stimulus effect in hooded rats with unilateral or bilateral lesions of parietal cortex.

When hooded rats with bilateral lesions of Krieg's area 7 (parietal cortex) were trained to locomote toward visual targets in a runway, they ran less accurately than did controls, although unilaterals ran accurately. When flashing lights were presented unexpectedly during their run, bilateral parietals were less disrupted than were controls, but they failed to show total neglect. Unilateral paritals turned toward distracters on either side but turned preferentially toward distracters contralateral to the intact hemisphere, particularly when distracters occurred bilaterally and simultaneously. Effects due to the omission of expected distracters were similar in parietals and controls. Rat parietal cortex is not essential for the redirection of attention to stimuli notable for their unexpected presence or absence, but parietal cortex may resolve interhemispheric competition. The results suggest a homology between parietal cortex in rat and primate.     

Spatial and temporal attention deficits following brain injury: A neuroanatomical decomposition of the temporal order judgement task

We investigated spatial and temporal deficits following brain injury using the temporal order judgement (TOJ) task. Patients judged the order in which two letters appeared to the left and right of fixation. We measured the extent of any spatial bias and the temporal resolution of the decision. Temporal and spatial deficits on the TOJ task were significantly correlated. The spatial bias on the TOJ task was also correlated with the spatial bias on a neglect task and with unilateral deficits on an extinction task, but not with extinction itself. These spatial deficits were all associated with damage to contralateral temporoparietal cortex. In contrast, the temporal resolution of TOJs was linked specifically to deficits in processing multiple stimuli on the neglect and extinction tasks and to damage to the right parietal lobe and the cerebellum. These data suggest that spatial and temporal deficits on the TOJ task reflect different underlying processes.     

Somatosensory detection in patients with circumscribed lesions of the brain

Summary The outer phalanx of the index finger was stimulated with a 1.0 ms electric pulse in 92 patients. All had a unilateral circumscribed and operatively verified lesion of one cerebral hemisphere. The relation between stimulus strength in mA and probability of stimulus detection, the psychometric curve, was measured and the detection threshold was calculated. In the mid-part (hand area) of the postcentral gyrus lesions of the cortex lining the posterior wall of the central sulcus increased the detection threshold of the contralateral index-finger, and the psychometric curve showed a poor discrimination between signal and noise. The larger the lesion of this part of cortex, the greater the threshold increase. Lesions of other parts of the parietal lobe did not affect somatosensory detection. Larger lesions of the frontal centrum semiovale and lesions that destroyed or undercut the superior and middle frontal gyrus increased the thresholds; right sided lesions gave bilateral defects, left sided lesions gave contralateral defects. Depending on whether low noise alternative routes between the prefrontal cortex and the postcentral gyrus were available or not, the psychometric curves either showed a sharp or a poor discrimination between signal and noise. This indicated that large fiber bundles in the frontal centrum semiovale and large cortical fields in the primary somatosensory cortex, middle and superior frontal gyrus, were involved in the transmission, decoding and detection of a single impulse. Lesions of the left hippocampus, right inferior frontal and orbitofrontal cortex increased the thresholds bilaterally. This was shown to be due to lack of task related attention and unstable selective attention. Lesions of the right hippocampus increased the threshold contralaterally.This work was supported by grants from the Foundation of Experimental Neurological Research, The P. Carl Petersen Foundation, and The Danish Medical Research Council     

14C-deoxyglucose mapping of the monkey brain during reaching to visual targets.

The strategies used by the macaca monkey brain in controlling the performance of a reaching movement to a visual target have been studied by the quantitative autoradiographic 14C-DG method. Experiments on visually intact monkeys reaching to a visual target indicate that V1 and V2 convey visuomotor information to the cortex of the superior temporal and parietoccipital sulci which may encode the position of the moving forelimb, and to the cortex in the ventral part and lateral bank of the intraparietal sulcus which may encode the location of the visual target. The involvement of the medial bank of the intraparietal sulcus in proprioceptive guidance of movement is also suggested on the basis of the parallel metabolic effects estimated in this region and in the forelimb representations of the primary somatosensory and motor cortices. The network including the inferior postarcuate skeletomotor and prearcuate oculomotor cortical fields and the caudal periprincipal area 46 may participate in sensory-to-motor and oculomotor-to-skeletomotor transformations, in parallel with the medial and lateral intraparietal cortices. Experiments on split brain monkeys reaching to visual targets revealed that reaching is always controlled by the hemisphere contralateral to the moving forelimb whether it is visually intact or 'blind'. Two supplementary mechanisms compensate for the 'blindness' of the hemisphere controlling the moving forelimb. First, the information about the location of the target is derived from head and eye movements and is sent to the 'blind' hemisphere via inferior parietal cortical areas, while the information about the forelimb position is derived from proprioceptive mechanisms and is sent via the somatosensory and superior parietal cortices. Second, the cerebellar hemispheric extensions of vermian lobules V, VI and VIII, ipsilateral to the moving forelimb, combine visual and oculomotor information about the target position, relayed by the 'seeing' cerebral hemisphere, with sensorimotor information concerning cortical intended and peripheral actual movements of the forelimb, and then send this integrated information back to the motor cortex of the 'blind' hemisphere, thus enabling it to guide the contralateral forelimb to the target.     

C-Deoxyglucose mapping of the monkey brain during reaching to visual targets

The strategies used by the macaca monkey brain in controlling the performance of a reaching movement to a visual target have been studied by the quantitative autoradiographic ¹⁴C-DG method.Experiments on visually intact monkeys reaching to a visual target indicate that V1 and V2 convey visuomotor information to the cortex of the superior temporal and parietoccipital sulci which may encode the position of the moving forelimb, and to the cortex in the ventral part and lateral bank of the intraparietal sulcus which may encode the location of the visual target. The involvement of the medial bank of the intraparietal sulcus in proprioceptive guidance of movement is also suggested on the basis of the parallel metabolic effects estimated in this region and in the forelimb representations of the primary somatosensory and motor cortices. The network including the inferior postarcuate skeletomotor and prearcuate oculomotor cortical fields and the caudal periprincipal area 46 may participate in sensory-to-motor and oculomotor-to-skeletomotor transformations, in parallel with the medial and lateral intraparietal cortices.Experiments on split brain monkeys reaching to visual targets revealed that reaching is always controlled by the hemisphere contralateral to the moving forelimb whether it is visually intact or ‘blind'. Two supplementary mechanisms compensate for the ‘blindness' of the hemisphere controlling the moving forelimb. First, the information about the location of the target is derived from head and eye movements and is sent to the ‘blind' hemisphere via inferior parietal cortical areas, while the information about the forelimb position is derived from proprioceptive mechanisms and is sent via the somatosensory and superior parietal cortices. Second, the cerebellar hemispheric extensions of vermian lobules V, VI and VIII, ipsilateral to the moving forelimb, combine visual and oculomotor information about the target position, relayed by the ‘seeing' cerebral hemisphere, with sensorimotor information concerning cortical intended and peripheral actual movements of the forelimb, and then send this integrated information back to the motor cortex of the ‘blind' hemisphere, thus enabling it to guide the contralateral forelimb to the target.