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Are visual representations in the human early visual cortex necessary for visual working memory (VWM)? Previous studies suggest that VWM is underpinned by distributed representations across several brain regions, including the early visual cortex. Notably, in these studies, participants had to memorize images under consistent visual conditions. However, in our daily lives, we must retain the essential visual properties of objects despite changes in illumination or viewpoint. The role of brain regions—particularly the early visual cortices—in these situations remains unclear. The present study investigated whether the early visual cortex was essential for achieving stable VWM. Focusing on VWM for object surface properties, we conducted fMRI experiments while male and female participants performed a delayed roughness discrimination task in which sample and probe spheres were presented under varying illumination. By applying multi-voxel pattern analysis to brain activity in regions of interest, we found that the ventral visual cortex and intraparietal sulcus were involved in roughness VWM under changing illumination conditions. In contrast, VWM was not supported as robustly by the early visual cortex. These findings show that visual representations in the early visual cortex alone are insufficient for the robust roughness VWM representation required during changes in illumination.

Previous neuropsychological studies of ideomotor apraxia (IMA) indicated impairments in pantomime actions for tool use for both right and left hands following lesions of parieto-premotor cortices in the left hemisphere. Using functional magnetic resonance imaging (fMRI) with multi-voxel pattern analysis (MVPA), we tested the hypothesis that the left parieto-premotor cortices are involved in the storage or retrieval of hand-independent representation of tool-use actions. In the fMRI scanner, one of three kinds of tools was displayed in pictures or letters, and the participants made pantomimes of the use of these tools using the right hand for the picture stimuli or with the left hand for the letters. We then used MVPA to classify which kind of tool the subjects were pantomiming. Whole-brain searchlight analysis revealed successful decoding using the activities largely in the contralateral primary sensorimotor region, ipsilateral cerebellum, and bilateral early visual area, which may reflect differences in low-level sensorimotor components for three types of pantomimes. Furthermore, a successful cross-classification between the right and left hands was possible using the activities of the left inferior parietal lobule (IPL) near the junction of the anterior intraparietal sulcus. Our finding indicates that the left anterior intraparietal cortex plays an important role in the production of tool-use pantomimes in a hand-independent manner, and independent of stimuli modality.

The cortical motor system follows a modular organization in which different features of executed movements are supported by distinct streams. Accordingly, different levels of action recognition, such as movement characteristics or action semantics may be processed within distinct networks. The present study aimed to differentiate areas related to the analysis of action features involving semantic knowledge from regions concerned with the evaluation of movement characteristics determined by structural object properties. To this end, the assessment of (i) tool-associated actions in relation to semantically, but not functionally inappropriate recipients (factor "Semantics"), and the evaluation of (ii) tool-associated movements performed with awkward versus correct hand postures (factor "Hand") were experimentally manipulated in an fMRI study with an event-related 2 × 2 factorial design. The videos used as stimuli displayed actions performed with the right hand in third-person perspective. Conjunction analysis of all four experimental conditions showed that observing videos depicting tool-related actions compared to rest was associated with widespread bilateral activity within the frontal lobes, inferior and superior parietal lobules, parts of the temporal lobes, as well as the occipital lobes. Viewing actions executed with incorrect compared to correct hand postures (factor "Hand") elicited significantly more activity within right primary sensory cortex (Brodmann area 2) and superior parietal lobule. Conversely, tool-associated actions displayed after semantically incorrect compared to correct recipients elicited higher activation within a left-lateralized network comprising the ventro-lateral prefrontal cortex (VLPFC), parts of the intraparietal sulcus and the angular gyrus (AG), as well as the supplementary motor area (SMA) and pre-SMA. Probabilistic diffusion tensor imaging-based tractography revealed two distinct fiber connections between AG and the frontal activation: A dorsal pathway via the superior longitudinal fascicle to the caudal part of VLPFC and a ventral pathway reaching the more rostral parts of VLPFC via the extreme capsule. The task-dependent relative modulation of activity within these brain networks composed of activated cortical areas connected by specific white matter tracts may indicate that the assessment of semantic action features relies on both dorso-ventral and ventral processing streams, whereas the analysis of hand postures with respect to objects depends on areas within the dorso-dorsal stream.

Although it is known that sounds can affect visual perception, the neural correlates for crossmodal interactions are still disputed. Previous tracer studies in non-human primates revealed direct anatomical connections between auditory and visual brain areas. We examined the structural connectivity of the auditory cortex in normal humans by diffusion-weighted tensor magnetic resonance imaging and probabilistic tractography. Tracts were seeded in Heschl's region or the planum temporale. Fibres crossed hemispheres at the posterior corpus callosum. Ipsilateral fibres seeded in Heschl's region projected to the superior temporal sulcus, the supramarginal gyrus and intraparietal sulcus and the occipital cortex including the calcarine sulcus. Fibres seeded in the planum temporale terminated primarily in the superior temporal sulcus, the supramarginal gyrus, the central sulcus and adjacent regions. Our findings suggest the existence of direct white matter connections between auditory and visual cortex--in addition to subcortical, temporal and parietal connections.

How do we achieve unitary perception of an object when it touches two parts of the sensory epithelium that are not contiguous? We investigated this problem with a simple psychophysical task, which we then used in an fMRI experiment. Two wooden rods were moved over two digits positioned to be spatially adjacent. The digits were either from one foot (or hand) or one digit was from either foot (or hand). When the rods were moving in phase, one object was reliably perceived. By contrast, when the rods were moving out of phase, two objects were reliably perceived. fMRI revealed four cortical areas where activity was higher when the moving rods were perceived as one object relative to when they were perceived as two separate objects. Areas in the right inferior parietal lobule, the left inferior temporal sulcus and the left middle frontal gyrus were activated for this contrast regardless of the anatomical configuration of the stimulated sensory epithelia. By contrast, the left intraparietal sulcus was activated specifically when integration across the midline was required, irrespective of whether the stimulation was applied to the hands or feet. These results reveal a network of brain areas involved in generating a unified percept of the presence of an object that comes into contact with different parts of the body surface.

Processing of one visual target (T1) makes it difficult to become aware of a second target (T2), when two targets, embedded in a stream of distractor stimuli, occur within about 500 ms. This phenomenon is known as attentional blink (AB) and reflects the temporal limitation in allocating visual attention. Although several studies suggest that parietal regions are concerned with the AB phenomenon, their functional relevance remains unclear. We investigated whether left and/or right intraparietal sulcus (IPS) contributed to the AB bottleneck using transcranial magnetic stimulation (TMS). The course of recovery from the AB deficit was facilitated when single pulse TMS induced a transient interruption of left or right IPS activity at a T1-TMS stimulus onset asynchrony of 350 ms, while there was no effect of TMS or sham stimulation delivered over Cz with the same timing. These results provide direct evidence that activation of left as well as right IPS is involved in the genesis of AB. This finding supports the idea that the IPS plays a critical role in the cortical network controlling the temporal dynamics of visual awareness.

Blindness is known to alter the responsiveness of visual cortex. Recently, reversible visual deprivation by blindfolding has been shown to affect non-visual abilities as well as visual cortical function. Here we investigated the effect of 2 h of blindfolding on cerebral cortical activation patterns during tactile form perception, using functional magnetic resonance imaging. Two form tasks were used, one requiring discrimination of global stimulus form and the other, detection of a gap in a bar. Blindfolded subjects showed significant deactivation during these tasks in regions that are intermediate in the hierarchy of visual shape processing: probable V3A and ventral intraparietal sulcus (vIPS). These regions lacked signal changes in controls. There were also task-specific increases in activation in blindfolded relative to control subjects, favoring the form over the gap task, along the IPS and in regions of frontal and temporal cortex. We also found alterations of functional connectivity that corresponded to the activity differences, with the emergence of correlated activity between the vIPS and V3A in blindfolded subjects. We conclude that blindfolding sighted individuals for a 2-h period induces significant changes in the neural processing of tactile form, probably reflecting short-term neural plasticity.

In primates, the frontal eye field (FEF) contains separate representations of saccadic and smooth-pursuit eye movements. The smooth-pursuit region (FEFsem) in macaque monkeys lies principally in the fundus and deep posterior wall of the arcuate sulcus, between the FEF saccade region (FEFsac) in the anterior wall and somatomotor areas on the posterior wall and convexity. In this study, cortical afferents to FEFsem were mapped by injecting retrograde tracers (WGA-HRP and fast blue) into electrophysiologically identified FEFsem sites in two monkeys. In the frontal lobe, labeled neurons were found mostly on the ipsilateral side in the (1) supplementary eye field region and lateral area F7; (2) area F2 along the superior limb of the arcuate sulcus; and (3) in the buried cortex of the arcuate sulcus extending along the superior and inferior limbs and including FEFsac and adjacent areas 8, 45, and PMv. Labeled cells were also found in the caudal periprincipal cortex (area 46) in one monkey. Labeled cells were found bilaterally in the frontal lobe in the deep posterior walls of the arcuate sulcus and postarcuate spurs and in cingulate motor areas 24 and 24c. In postcentral cortical areas all labeling was ipsilateral and there were two major foci of labeled cells: (1) the depths of the intraparietal sulcus including areas VIP, LIP, and PEa, and (2) the anterior wall and fundus of the superior temporal sulcus including areas PP and MST. Smaller numbers of labeled cells were found in superior temporal sulcal areas FST, MT, and STP, posterior cingulate area 23b, area 3a within the central sulcus, areas SII, RI, Tpt in the lateral sulcus, and parietal areas 7a, 7b, PEc, MIP, DP, and V3A. Many of these posterior afferent cortical areas code visual-motion (MT, MST, and FST) or visual-motion and vestibular (PP, VIP) signals, consistent with the responses of neurons in FEFsem and with the overall physiology and anatomy of the smooth-pursuit eye movement system.

Fourteen patients with a chronic, unilateral lesion restricted to the frontal lobe (twelve involving the frontal eye field (FEF)), nine patients with a chronic, unilateral lesion restricted to posterior association cortex (eight involving the intraparietal sulcus (IPS)), and twelve neurologically normal control subjects were studied in an anti-saccade task. A combination of manipulating cuing and fixation offset enabled us to examine the effects of chronic oculomotor lesions on both saccade preparation and voluntary control over ocular fixation. Patients with lesions of the FEF made more errors (reflexive glances) toward contralesional targets, whereas patients with IPS lesions made fewer errors toward contralesional targets. Patients with IPS lesions had increased latencies to initiate saccades away from contralesional targets. For FEF patients, the presence of a fixation point inhibited the initiation of contralesionally directed saccades less than those directed ipsilesionally. Saccade preparation in response to a cue did not reduce the inhibitory effect of a fixation point on initiating anti-saccades directed either ipsilesionally or contralesionally for either patient group. We conclude that chronic IPS lesions result in a reduced contralesional visual grasp reflex (VGR) and delayed utilization of visual signals in the contralesional field for planning voluntary eye movements. In contrast, patients with chronic FEF lesions are impaired in inhibiting the VGR toward contralesional signals, and manifest an asymmetry in the balance between fixation and saccade activity. Moreover, voluntary control of fixation is compromised after chronic damage to either frontal or parietal oculomotor cortex.

We used event-related functional magnetic resonance imaging to study the neural correlates of endogenous spatial attention for vision and touch. We examined activity associated with attention-directing cues (central auditory pure tones), symbolically instructing subjects to attend to one hemifield or the other prior to upcoming stimuli, for a visual or tactile task. In different sessions, subjects discriminated either visual or tactile stimuli at the covertly attended side, during bilateral visuotactile stimulation. To distinguish cue-related preparatory activity from any modulation of stimulus processing, unpredictably on some trials only the auditory cue was presented. The use of attend-vision and attend-touch blocks revealed whether preparatory attentional effects were modality-specific or multimodal. Unimodal effects of spatial attention were found in somatosensory cortex for attention to touch, and in occipital areas for attention to vision, both contralateral to the attended side. Multimodal spatial effects (i.e. effects of attended side irrespective of task-relevant modality) were detected in contralateral intraparietal sulcus, traditionally considered a multimodal brain region; and also in the middle occipital gyrus, an area traditionally considered purely visual. Critically, all these activations were observed even on cue-only trials, when no visual or tactile stimuli were subsequently presented. Endogenous shifts of spatial attention result in changes of brain activity prior to the presentation of target stimulation (baseline shifts). Here, we show for the first time the separable multimodal and unimodal components of such preparatory activations. Additionally, irrespective of the attended side and modality, attention-directing auditory cues activated a network of superior frontal and parietal association areas that may play a role in voluntary control of spatial attention for both vision and touch.

The functional properties of motion selective areas in human visual cortex, including V3A, MT+, and intraparietal sulcus (IPS) are not fully understood. To examine the functional specialization of these areas for global and local motion processing, we used off-line, neuronavigated, continuous theta burst (cTBS) transcranial magnetic stimulation to temporarily alter neural activity within unilateral V3A, MT+, and IPS. A within-subjects design was employed and stimulation sessions were separated by at least 24 h. In each session, subjects were asked to discriminate the global motion directions of successively presented random dot kinematograms (RDKs) before and after cTBS. RDKs were presented at either 100 or 40 % coherence in either the left or right visual field. We found that V3A stimulation selectively impaired discrimination of 100 % coherent motion, while MT+ stimulation selectively impaired discrimination of 40 % coherent motion. IPS stimulation impaired discrimination of both motion stimuli. All cTBS effects were specific to stimuli presented contralaterally to the stimulation site and vertex stimulation had no effect. The double dissociation between the cTBS effects on MT+ and V3A indicates distinct roles for these two regions in motion processing. Judging the direction of 100 % coherent motion can rely on local motion processing because every dot moves in the same direction. However, judging the global direction of 40 % coherent motion requires global processing. Thus, our results suggest separate, parallel processing of local and global motion in V3A and MT+, respectively, with the outputs of these two areas being combined within the IPS.

We report an event-related functional magnetic-resonance-imaging (fMRI) experiment that investigates the relationship of transient visual object memory, visuospatial orienting, and object recognition. Delayed object matching and visuospatial orienting involved a highly overlapping network of brain areas. Common areas were the frontal eye fields (FEF), the pre-supplementary motor area (pre-SMA)/SMA complex, the precentral gyri, and the horizontal and descending branches of the intraparietal sulcus (IPS). Selective delay activation was observed anterior to the FEF and in the ascending part of the IPS. Right dorsolateral prefrontal cortex was involved in goal-directed visual search, but showed no delay activity.

Two functionally different cortical areas are located in the rostral part of the intraparietal sulcus (IP): the ventral intraparietal area (VIP), along the fundus of the sulcus, and the anterior intraparietal area (AIP), rostral in the lateral bank. VIP and AIP have functional properties comparable to those of the ventral premotor areas, F4 and F5, respectively. The aim of this study was to establish whether these intraparietal and premotor areas have direct and specific anatomical connections. Neural tracers were injected in F4, F5, and AIP in three macaque monkeys. The results showed that F4 and F5 are targets of strong projections from VIP and AIP, respectively, and that the linkage between F5 and AIP is highly selective. These data support the notion that parietofrontal connections selectively link areas displaying similar functional properties and form largely segregated anatomical circuits. Each of these circuits is possibly dedicated to specific aspects of sensorimotor transformations. In particular, the AIP-F5 circuit should play a crucial role in visuomotor transformation for grasping, the VIP-F4 circuit is possibly involved in peripersonal space coding for movement.

Functional magnetic resonance imaging (fMRI) was used to assess cerebral activation during manipulation of various complex meaningless objects as compared to manipulation of a single simple object (a sphere). Significant activation was found bilaterally in the ventral premotor cortex (Brodmann's area 44), in the cortex lining the anterior part of the intraparietal sulcus (most probably corresponding to monkey anterior intraparietal area, AIP), in the superior parietal lobule and in the opercular parietal cortex including the secondary somatosensory area (SII). We suggest that the cortex lining the anterior part of the intraparietal sulcus and area 44 are functionally connected and mediate object manipulation in humans.

A scheme for the cortical control of saccadic eye movements is proposed based partly on defects revealed by specific test paradigms in humans with discrete lesions. Three different cortical areas are capable of triggering saccades. The frontal eye field disengages fixation, and triggers intentional saccades to visible targets, to remembered target locations, or to the location where it is predicted that the target will reappear (i.e., saccades concerned with intentional exploration of the visual environment). The parietal eye field triggers saccades made reflexively on the sudden appearance of visual targets (i.e., saccades concerned with reflexive exploration of the visual environment). The supplementary eye field is important for triggering sequences of saccades and in controlling saccades made during head or body movement (i.e., saccades concerned with complex motor programming). Three other areas contribute to the preparation of certain types of saccades. The prefrontal cortex (area 46 of Brodmann) plays a crucial role for planning saccades to remembered target locations. The inferior parietal lobule is involved in the visuospatial integration used for calculating saccade amplitude. The hippocampus appears to control the temporal working memory required for memorization of the chronological order of sequences of saccades.

Fifty to 85 % of patients with sensorimotor hemiparesis following stroke encounter impaired tactile processing and proprioception. Sensory feedback is, however, paramount for motor recovery. Sensory feedback through passively guided somatosensory discrimination exercises has been used in therapy, but so far, no studies have investigated which brain areas are involved in this process. Therefore, we performed a study with functional magnetic resonance imaging (fMRI) to examine brain areas related to discriminating passively guided shape and length discrimination in stroke patients and evaluate whether they differed from healthy age-matched controls. Eight subcortical stroke patients discriminated different shapes or length based on passive finger movements provided by an fMRI compatible robot. The data were contrasted to a control condition whereby patients discriminated music fragments. Passively guided somatosensory discrimination versus music discrimination elicited activation in similar frontoparietal areas in stroke patients compared to the healthy control group. Still, patients had increased activation in the right angular gyrus, left superior lingual gyrus, and right cerebellar lobule VI compared to healthy volunteers. Conversely, healthy volunteers activated the right precentral gyrus to a greater extent than patients. In both groups, shape discrimination resulted in anterior intraparietal sulcus and premotor activation, while length discrimination elicited a more medially located parietal activation with mainly right-sided premotor activity. The current study is a first step in clarifying brain activations during passively guided shape and length discrimination in subcortical stroke patients. Research into the effects of the use of sensory discrimination exercises on brain reorganization and brain plasticity is encouraged.

Non-pulsatile tinnitus is considered to be an auditory phantom percept. The extremely emotional context of disabling tinnitus often leads to a higher level of selective attention directed toward the tinnitus. As such, tinnitus is a continuously distracting auditory event. Auditory attention is associated with the activation of the intraparietal sulcus (IPS), and modulating the IPS with 10 Hz transcranial magnetic stimulation (TMS) creates the ability to ignore salient distractors. Thus, it can be expected that modulating the parietal area might interfere with the perception of tinnitus. The effect of TMS on tinnitus is evaluated using a double-cone coil tilted to the left parietal area in 24 individuals (study 1) and in 40 individuals with the double-cone coil symmetrically overlying both parietal areas (study 2). When transient tinnitus suppression is noted, the patient is asked to estimate the decrease in tinnitus in percentage using the numeric rating scale. The procedure is repeated with stimulations at sham, 1 and 10 Hz, each stimulation session consisting of 200 pulses for study 1 and for study 2 stimulations at sham, 1, 5, and 10 Hz, each stimulation session consisting of 200 pulses. For both studies, the order of the different stimulation frequencies was randomized over the participants. For study 1, patients report a significant transient reduction of the tinnitus percept for 10 Hz stimulation in comparison with, respectively, pre-treatment, sham, and 1 Hz stimulation, with a suppression effect of 11.36 %. No significant effect was obtained for 1 Hz stimulation with the coil tilted toward the left parietal area. For study, 2 patients revealed a significant suppression effect on 1, 5, and 10 Hz in comparison with pre-treatment. However, only stimulation at 5 and 10 Hz had a significant difference in comparison with sham with a suppression effect of, respectively, 8.78 and 9.50 %. Our data suggest that the parietal area is involved in tinnitus perception and that 10 Hz TMS using the double-cone coil overlying the parietal area can modulate tinnitus.

We investigated associations between olfactory function and gray matter thickness in 46 healthy young subjects by means of an automated technique for measuring cortical thickness. We used an extended version of the Sniffin’ Sticks test to assess olfactory function, including odor threshold, concentration discrimination, quality discrimination, and odor identification. We observed a correlation between olfactory performance and cortical thickness of structures involved in earlier and later stages of chemosensory processing such as right medial orbitofrontal cortex, right insula, and adjacent cortex. Furthermore, we found significant bilateral correlations of olfactory performance with cortical thickness of areas around the central sulcus bilaterally, structures responsible for voluntary respiration and sniffing. In addition to expected general sex effects on cortical thickness, we observed areas, such as the entorhinal cortex, occipital cortex, intraparietal sulcus and insula (all in the right hemisphere), where the correlation between higher order olfactory functions and cortical thickness differed between women and men. These data demonstrate, for some neuroanatomical structures, a link between cortical thickness and olfactory function, in that thicker cortex is usually associated with better performance, but not always. This association between anatomy and olfactory performance suggests a possible biological explanation for the high degree of individual differences and sex effects observed in higher order olfactory tasks.

The simulation concept suggested by Jeannerod (Neuroimage 14:S103-S109, 2001) defines the S-states of action observation and mental simulation of action as action-related mental states lacking overt execution. Within this framework, similarities and neural overlap between S-states and overt execution are interpreted as providing the common basis for the motor representations implemented within the motor system. The present brain imaging study compared activation overlap and differential activation during mental simulation (motor imagery) with that while observing gymnastic movements. The fMRI conjunction analysis revealed overlapping activation for both S-states in primary motor cortex, premotor cortex, and the supplementary motor area as well as in the intraparietal sulcus, cerebellar hemispheres, and parts of the basal ganglia. A direct contrast between the motor imagery and observation conditions revealed stronger activation for imagery in the posterior insula and the anterior cingulate gyrus. The hippocampus, the superior parietal lobe, and the cerebellar areas were differentially activated in the observation condition. In general, these data corroborate the concept of action-related S-states because of the high overlap in core motor as well as in motor-related areas. We argue that differential activity between S-states relates to task-specific and modal information processing.

In this study we attempted to differentiate distinct components of the saccade network, namely cortical ocular motor centers and parieto-occipital brain regions, by means of a “minimal design” approach. Using a blocked design fMRI paradigm we evaluated the BOLD changes in a 2 × 2 factorial design experiment which was performed in complete darkness: while looking straight ahead with eyes open (OPEN) or closed (CLOSED) as well as during the execution of self-initiated horizontal to-and-fro saccades with the eyes open (SACCopen) or closed (SACCclosed). Eye movements were monitored outside the scanner via electro-oculography and during scanning using video-oculography. Unintentional eye-drifts did not differ during OPEN and CLOSED and saccade frequencies, and amplitudes did not vary significantly between the two saccade conditions. The main findings of the functional imaging study were as follows: (1) Saccades with eyes open or closed in complete darkness lead to distinct differences in brain activation patterns. (2) A parieto-occipital brain region including the precuneus, superior parietal lobule, posterior part of the intraparietal sulcus (IPS), and cuneus was relatively deactivated during saccades performed with eyes closed but not during saccades with eyes open or when looking straight ahead. This could indicate a preparatory state for updating spatial information, which is active during saccades with eyes open even without actual visual input. The preparatory state is suppressed when the eyes are closed during the saccades. (3) Selected ocular motor areas, not including the parietal eye field (PEF), show a stronger activation during SACCclosed than during SACCopen. The increased effort involved in performing saccades with eyes closed, perhaps due to the unusualness of the task, may be the cause of this increased activation.

It has been widely argued that the intraparietal sulcus (IPS) is involved in tasks that evoke representations of numerical magnitude, among other cognitive functions. However, the causal role of this parietal region in processing symbolic and non-symbolic numerosity has not been established. The current study used repetitive Transcranial Magnetic Stimulation (rTMS) to the left and right IPS to investigate the effects of temporary deactivations of these regions on the capacity to represent symbolic (Arabic numbers) and non-symbolic (arrays of dots) numerosities. We found that comparisons of both symbolic and non-symbolic numerosities were impaired after rTMS to the left IPS but enhanced by rTMS to the right IPS. A signature effect of numerical distance was also found: greater impairment (or lesser facilitation) when comparing numerosities of similar magnitude. The reverse pattern of impairment and enhancement was found in a control task that required judging an analogue stimulus property (ellipse orientation) but no numerosity judgements. No rTMS effects for the numerosity tasks were found when stimulating an area adjacent but distinct from the IPS, the left and right angular gyrus. These data suggest that left IPS is critical for processing symbolic and non-symbolic numerosity; this processing may thus depend on common neural mechanisms, which are distinct from mechanisms supporting the processing of analogue stimulus properties.

The spatial location of activation for writing individual letters and for writing simple dots was studied using event-related functional MRI. Ten healthy right-handed subjects were scanned while performing two different protocols with self-paced repetitive movement. In the first protocol with self-paced dot writing, we observed significant activation in regions known to participate in motor control: contralateral to the movement in the primary sensorimotor and supramarginal cortices, the supplementary motor area (SMA) with the underlying cingulate, in the thalamus and, to a lesser extent, in the ipsilateral inferior parietal and occipital cortices. In the second protocol, we investigated an elemental writing feature—writing single letters. We observed statistically significant changes in the premotor, sensorimotor and supramarginal cortices, the SMA and the thalamus with left predominance, and in the bilateral premotor and inferior/superior parietal cortices. The parietal region that was active during the writing of single letters spanned the border between the parietal superior and inferior lobuli Brodmann area (BA 2, 40), deep in the intraparietal sulcus, with a surprising right-sided dominance. The direct comparison of the results of the two protocols was not significant with a confidence level of P

Mirror apraxia is a condition in which patients with lesions of the posterior parietal cortex have deficits in reaching to objects presented through a mirror. The aim of the present study was to investigate possible mechanisms underlying this disorder. First, we addressed the question of whether mirror apraxia is exhibited to the same extent in peripersonal and in body space. Four patients with lesions of the posterior parietal lobe on either side and with marked mirror apraxia were required to reach for objects that were presented to them through a mirror and located either in body space (i.e. on the body surface) or in peripersonal space (i.e. in the reaching distance). Whereas reaching for objects located in body space was flawless in all patients, the performance deteriorated when the same objects were transferred to the peripersonal space. Although the objects were located only a few centimetres above the body surface, the patients reached towards the virtual object in the mirror. Based on these results we suggest that mirror apraxia may originate from a dissociation between the representations of body schema and peripersonal space and that objects located on the body surface become integrated into the body schema. In the second part of the study, using positron emission tomography study (PET), we studied the cerebral activation pattern during reaching to objects presented through a mirror in the peripersonal space in healthy subjects. The results show that increased neural activity in the anterior part of the intraparietal sulcus and in the dorsal premotor cortex was bound to the transformation of the target position from the mirror space to the real space. In contrast, the activity related to object localization in the mirror occurred at the parieto-occipital junction. Both mirror and arm transformation involved the medial posterior part of the superior parietal lobule, putatively area V6a. The results demonstrate that acting through a mirror is processed in a number of cortical areas of the dorsal stream.

The representation of the visual field in the primate lateral intraparietal area (LIP) was examined, using a rapid, computer-driven receptive field (RF) mapping procedure. RF characteristics of single LIP neurons could thus be measured repeatedly under different behavioral conditions. Here we report data obtained using a standard ocular fixation task during which the animals were required to monitor small changes in color of the fixated target. In a first step, statistical analyses were conducted in order to establish the experimental limits of the mapping procedure on 171 LIP neurons recorded from three hemispheres of two macaque monkeys. The characteristics of the receptive fields of LIP neurons were analyzed at the single cell and at the population level. Although for many neurons the assumption of a simple two-dimensional gaussian profile with a central area of maximal excitability at the center and progressively decreasing response strength at the periphery can represent relatively accurately the spatial structure of the RF, about 19% of the cells had a markedly asymmetrical shape. At the population level, we observed, in agreement with prior studies, a systematic relation between RF size and eccentricity. However, we also found a more accentuated overrepresentation of the central visual field than had been previously reported and no marked differences between the upper and lower visual representation of space. This observation correlates with an extension of the definition of LIP from the posterior third of the lateral intraparietal sulcus to most of the middle and posterior thirds. Detailed histological analyses of the recorded hemispheres suggest that there exists, in this newly defined unitary functional cortical area, a coarse but systematic topographical organization in area LIP that supports the distinction between its dorsal and ventral regions, LIPd and LIPv, respectively. Paralleling the physiological data, the central visual field is mostly represented in the middle dorsal region and the visual periphery more ventral and posterior. An anteroposterior gradient from the lower to the upper visual field representations can also be identified. In conclusion, this study provides the basis for a reliable mapping method in awake monkeys and a reference for the organization of the properties of the visual space representation in an area LIP extended with respect to the previously described LIP and showing a relative emphasis of central visual field.

In the first part of this article, we review our neurophysiological studies of the hand-manipulation-related neurons in the anterior part of the lateral bank of the intraparietal sulcus (area AIP). We describe the properties of visually responsive neurons in area AIP. Object-type visual-dominant neurons responded to the sight of objects and showed selectivity not only for simple geometrical shapes, but also for complex objects such as a knob-in-groove and a plate-in-groove. Some of the object-type visual-dominant neurons showed selectivity for the orientation of the longitudinal axis or the plane (surface) of a plate or a ring. In the second part of this article, we review our study of binocular visual neurons in the caudal part of the lateral bank of the intraparietal sulcus (c-IPS area), in particular, of axis-orientation-selective (AOS) neurons and surface-orientation-selective (SOS) neurons. AOS neurons preferred long and thin stimuli, were sensitive to binocular disparity, and tuned to the axis orientation in three-dimensional (3D) space. SOS neurons preferred broad and flat stimuli and were tuned to the surface orientation in depth. Some SOS neurons responded to a square in a random dot stereogram (RDS) with orientation tuning, suggesting that they encode surface orientation from a disparity gradient. Others responded to solid figure stereograms with orientation disparity and/or width disparity. It was concluded that the c-IPS area is a higher center for stereopsis, which integrates various binocular disparity signals received from the V3 complex and other prestriate areas to represent the neural code for 3D features. It may send projections to the AIP area and contribute to visual adjustment of the shape of the handgrip and/or hand orientation for manipulation and grasping. Neurons of the AIP area may also receive monocular cues of depth from the ventral visual pathway to discriminate the 3D shape of the object of manipulation.

The activity of each of 99 intraparietal neurons was studied in three awake-behaving rhesus monkeys (Macaca mulatta) while subjects performed 100-900 delayed saccade trials. On each trial, a saccadic target was presented at one location selected randomly from a grid of 441 locations spanning 40 degrees of horizontal and vertical visual space. Individual neurons in our population were sensitive to both the direction and amplitude of saccades. Response fields, which plotted firing rate as a function of the horizontal and vertical amplitude of movements for each neuron, were characterized by a Cartesian two-dimensional gaussian model. The goodness-of-fit of these gaussian models was tested by: (1) comparing observed responses with predicted responses for each movement; and (2) by computing the percentage of variance explained by each model. Cartesian Gaussian models provided a good fit to the response fields of most neurons. Across our population, the Gaussian fit to the response field of each neuron accounted for more of the variance in neuronal activity when the data were plotted with regard to the horizontal and vertical amplitude of the saccade than when the same data were plotted with regard to the position of the saccadic target. The Gaussian functions were used to estimate the eccentricity and spatial tuning breadth of each neuronal response field. Modal response field radius was less than 5 degrees, whereas mean response field radius was about 10 degrees. Linear regression analysis demonstrated that response field eccentricity accounted for less than 30% of the variance in response field radius. Analysis of the horizontal distribution of response field centers showed an approximately normal distribution around central fixation. Most histologically recovered neurons were located on the lateral bank of the intraparietal sulcus, although a small number of saccade-related neurons were recorded from Brodmann's area 5 on the medial bank of the intraparietal sulcus.

Neurophysiology and neuroimaging research implicates distinct regions of posterior parietal cortex for reaching versus grasping and for completing these movements in central versus peripheral space. Typically, visuomotor tasks only examine movements made in the frontoparallel plane. We examined a patient with a right superior parietal lesion encompassing the parietal-occipital junction, the intraparietal sulcus and the putative human homologue of V6A on pointing tasks in the sagittal or frontoparallel planes. The patient did not demonstrate a speed-accuracy trade-off, but did show larger times post-peak velocity for all movement directions. Her movements in the sagittal axis were more disordered than movements in the frontoparallel plane. These data indicate a role for superior parietal cortex in fine tuning of visually guided movements and more particularly for movements made back towards the body.

A cyto- and myeloarchitectonic parcellation of the intraparietal sulcus in the rhesus monkey was correlated with the pattern of afferent connections from the parietal lobe as determined by autoradiographic techniques. Area PEa in the upper bank receives topographically-organized input from the ventral and caudal superior parietal lobule and the medial surface of the parietal lobe. Area POa in the lower bank is the recipient of a projection from the rostral inferior parietal lobule. Area IPd, situated in the depth of the intraparietal sulcus, receives converging input from the caudal superior parietal lobule, medial surface of the parietal lobe, and mid- and caudal inferior parietal lobule. Finally, intrinsic sequences of connections, directed from rostral to caudal and caudal to rostral, exist within both areas PEa and POa, each having a distinctive laminar pattern of origins and terminations.

The origin, course, and termination of parieto-temporal connections in the rhesus monkey were studied by autoradiographic techniques. The caudal third of the inferior parietal lobule (including the adjacent lower bank of the intraparietal sulcus) is the chief source of these projections. It projects to three separate architectonic areas in the superior temporal sulcus and to three different areas on the ventral surface of the temporal lobe: the parahippocampal gyrus, presubiculum, and perirhinal cortex. The mid-inferior parietal lobule and medial surface of the parietal lobe, by contrast, project only to the caudal upper bank of the superior temporal sulcus. The rostral inferior parietal lobule and the superior parietal lobule, as well as the postcentral gyrus and rostral parietal operculum, do not project to the temporal lobe. Fibers travel from the posterior parietal region to temporal cortex by way of several different routes. One fiber bundle courses in the superior temporal gyrus and terminates in the superior temporal sulcus. Another proceeds ventrally, between the depth of the superior temporal sulcus and the geniculocalcarine tract, to the parahippocampal area. A separate bundle, coursing part of the way in the company of the cingulum bundle, conveys posterior parietal fibers to the presubiculum.

Corticopontine projection patterns were studied after injections of an 3H-leucine and 3H-proline mixture into each of four distinct cortical fields within the inferior parietal lobule and dorsal prelunate gyrus. Different preferential patterns of pontine labeling were observed for each of the four cortical areas studied. Multiple injections across the dorsal aspect of the prelunate gyrus (area DP) yielded scattered patches of label limited to the dorsolateral pontine nuclear region. A single injection within the lateral intraparietal area (area LIP), located in the caudal portion of the lateral bank of the intraparietal sulcus resulted in a series of labeled patches across the dorsal tier of cells stretching across the dorsal portions of the dorsolateral, peduncular and dorsal pontine nuclei. Injection of the cortex on the caudal aspect of the inferior parietal convexity (area 7a) produced multiple patches of label along the lateral margin of the ventral, lateral, and dorsolateral nuclei. Injection of area 7b resulted in label along the lateral aspects of the ventral, lateral and dorsolateral nuclei, as seen with area 7a injections, as well as additional label in the ventromedial portions of the ventral, peduncular and paramedian pontine nuclei. These results provide supporting anatomic evidence for the functional subdivision of the inferior parietal lobule and dorsal aspect of the prelunate gyrus and provide new information about the organization of cortical projections to the primate pontine nuclei.