Voluntary orienting is dissociated from target detection in human posterior parietal cortex

Human ability to attend to visual stimuli based on their spatial locations requires the parietal cortex. One hypothesis maintains that parietal cortex controls the voluntary orienting of attention toward a location of interest. Another hypothesis emphasizes its role in reorienting attention toward visual targets appearing at unattended locations. Here, using event-related functional magnetic resonance (ER-fMRI), we show that distinct parietal regions mediated these different attentional processes. Cortical activation occurred primarily in the intraparietal sulcus when a location was attended before visual-target presentation, but in the right temporoparietal junction when the target was detected, particularly at an unattended location.     

Integration of target and effector information in human posterior parietal cortex for the planning of action

Recently, using event-related functional MRI (fMRI), we located a bilateral region in the human posterior parietal cortex (retIPS) that topographically represents and updates targets for saccades and pointing movements in eye-centered coordinates. To generate movements, this spatial information must be integrated with the selected effector. We now tested whether the activation in retIPS is dependent on the hand selected. Using 4-T fMRI, we compared the activation produced by movements, using either eyes or the left or right hand, to targets presented either leftward or rightward of central fixation. The majority of the regions activated during saccades were also activated during pointing movements, including occipital, posterior parietal, and premotor cortex. The topographic retIPS region was activated more strongly for saccades than for pointing. The activation associated with pointing was significantly greater when pointing with the unseen hand to targets ipsilateral to the hand. For example, although there was activation in the left retIPS when pointing to targets on the right with the left hand, the activation was significantly greater when using the right hand. The mirror symmetric effect was observed in the right retIPS. Similar hand preferences were observed in a nearby anterior occipital region. This effector specificity is consistent with previous clinical and behavioral studies showing that each hand is more effective in directing movements to targets in ipsilateral visual space. We conclude that not only do these regions code target location, but they also appear to integrate target selection with effector selection.     

Topographic organization for delayed saccades in human posterior parietal cortex

Posterior parietal cortex (PPC) is thought to play a critical role in decision making, sensory attention, motor intention, and/or working memory. Research on the PPC in non-human primates has focused on the lateral intraparietal area (LIP) in the intraparietal sulcus (IPS). Neurons in LIP respond after the onset of visual targets, just before saccades to those targets, and during the delay period in between. To study the function of posterior parietal cortex in humans, it will be crucial to have a routine and reliable method for localizing specific parietal areas in individual subjects. Here, we show that human PPC contains at least two topographically organized regions, which are candidates for the human homologue of LIP. We mapped the topographic organization of human PPC for delayed (memory guided) saccades using fMRI. Subjects were instructed to fixate centrally while a peripheral target was briefly presented. After a further 3-s delay, subjects made a saccade to the remembered target location followed by a saccade back to fixation and a 1-s inter-trial interval. Targets appeared at successive locations "around the clock" (same eccentricity, approximately 30 degrees angular steps), to produce a traveling wave of activity in areas that are topographically organized. PPC exhibited topographic organization for delayed saccades. We defined two areas in each hemisphere that contained topographic maps of the contra-lateral visual field. These two areas were immediately rostral to V7 as defined by standard retinotopic mapping. The two areas were separated from each other and from V7 by reversals in visual field orientation. However, we leave open the possibility that these two areas will be further subdivided in future studies. Our results demonstrate that topographic maps tile the cortex continuously from V1 well into PPC.     

Specialization of reach function in human posterior parietal cortex

Posterior parietal cortex (PPC) plays an important role in the planning and control of goal-directed action. Single-unit studies in monkeys have identified reach-specific areas in the PPC, but the degree of effector and computational specificity for reach in the corresponding human regions is still under debate. Here, we review converging evidence spanning functional neuroimaging, parietal patient and transcranial magnetic stimulation studies in humans that suggests a functional topography for reach within human PPC. We contrast reach to saccade and grasp regions to distinguish functional specificity and also to understand how these different goal-directed actions might be coordinated at the cortical level. First, we present the current evidence for reach specificity in distinct modules in PPC, namely superior parietal occipital cortex, midposterior intraparietal cortex and angular gyrus, compared to saccade and grasp. Second, we review the evidence for hemispheric lateralization (both for hand and visual hemifield) in these reach representations. Third, we review evidence for computational reach specificity in these regions and finally propose a functional framework for these human PPC reach modules that includes (1) a distinction between the encoding of reach goals in posterior–medial PPC as opposed to reach movement vectors in more anterior–lateral PPC regions, and (2) their integration within a broader cortical framework for reach, grasp and eye–hand coordination. These findings represent both a confirmation and extension of findings that were previously reported for the monkey.     

Remapping in human visual cortex

With each eye movement, stationary objects in the world change position on the retina, yet we perceive the world as stable. Spatial updating, or remapping, is one neural mechanism by which the brain compensates for shifts in the retinal image caused by voluntary eye movements. Remapping of a visual representation is believed to arise from a widespread neural circuit including parietal and frontal cortex. The current experiment tests the hypothesis that extrastriate visual areas in human cortex have access to remapped spatial information. We tested this hypothesis using functional magnetic resonance imaging (fMRI). We first identified the borders of several occipital lobe visual areas using standard retinotopic techniques. We then tested subjects while they performed a single-step saccade task analogous to the task used in neurophysiological studies in monkeys, and two conditions that control for visual and motor effects. We analyzed the fMRI time series data with a nonlinear, fully Bayesian hierarchical statistical model. We identified remapping as activity in the single-step task that could not be attributed to purely visual or oculomotor effects. The strength of remapping was roughly monotonic with position in the visual hierarchy: remapped responses were largest in areas V3A and hV4 and smallest in V1 and V2. These results demonstrate that updated visual representations are present in cortical areas that are directly linked to visual perception.     

Activation of the human posterior parietal cortex by median nerve stimulation

We recorded somatosensory evoked magnetic fields from ten healthy, right-handed subjects with a 122-channel whole-scalp SQUID magnetometer. The stimuli, exceeding the motor threshold, were delivered alternately to the left and right median nerves at the wrists, with interstimulus intervals of 1, 3, and 5 s. The first responses, peaking around 20 and 35 ms, were explained by activation of the contralateral primary somatosensory cortex (SI) hand area. All subjects showed additional deflections which peaked after 85 ms; the source locations agreed with the sites of the secondary somatosensory cortices (SII) in both hemispheres. The SII responses were typically stronger in the left than the right hemisphere. All subjects had an additional source, not previously reported in human evoked response data, in the contralateral parietal cortex. This source was posterior and medial to the SI hand area, and evidently in the wall of the postcentral sulcus. It was most active at 70–110 ms.     

Role of the posterior parietal cortex in updating reaching movements to a visual target.

The exact role of posterior parietal cortex (PPC) in visually directed reaching is unknown. We propose that, by building an internal representation of instantaneous hand location, PPC computes a dynamic motor error used by motor centers to correct the ongoing trajectory. With unseen right hands, five subjects pointed to visual targets that either remained stationary or moved during saccadic eye movements. Transcranial magnetic stimulation (TMS) was applied over the left PPC during target presentation. Stimulation disrupted path corrections that normally occur in response to target jumps, but had no effect on those directed at stationary targets. Furthermore, left-hand movement corrections were not blocked, ruling out visual or oculomotor effects of stimulation.     

Directional selectivity of BOLD activity in human posterior parietal cortex for memory-guided double-step saccades

We used functional magnetic resonance imaging (fMRI) to investigate the role of the human posterior parietal cortex (PPC) in storing target locations for delayed double-step saccades. To do so, we exploited the laterality of a subregion of PPC that preferentially responds to the memory of a target location presented in the contralateral visual field. Using an event-related design, we tracked fMRI signal changes in this region while subjects remembered the locations of two sequentially flashed targets, presented in either the same or different visual hemifields, and then saccaded to them in sequence. After presentation of the first target, the fMRI signal was always related to the side of the visual field in which it had been presented. When the second target was added, the cortical activity depended on the respective locations of both targets but was still significantly selective for the target of the first saccade. We conclude that this region within the human posterior parietal cortex not only acts as spatial storage center by retaining target locations for subsequent saccades but is also involved in selecting the target for the first intended saccade.     

Role for human posterior parietal cortex in visual processing of aversive objects in peripersonal space

The posterior parietal cortex of both human and non-human primates is known to play a crucial role in the early integration of visual information with somatosensory, proprioceptive and vestibular signals. However, it is not known whether in humans this region is further capable of discriminating if a stimulus poses a threat to the body. In this functional magnetic resonance imaging (fMRI) study, we tested the hypothesis that the posterior parietal cortex of humans is capable of modulating its response to the visual processing of noxious threat representation in the absence of tactile input. During fMRI, participants watched while we "stimulated" a visible rubber hand, placed over their real hand with either a sharp (painful) or a blunt (nonpainful) probe. We found that superior and inferior parietal regions (BA5/7 and BA40) increased their activity in response to observing a painful versus nonpainful stimulus. However, this effect was only evident when the rubber hand was in a spatially congruent (vs. incongruent) position with respect to the participants' own hand. In addition, areas involved in motivational-affective coding such as mid-cingulate (BA24) and anterior insula also showed such relevance-dependent modulation, whereas premotor areas known to receive multisensory information about limb position did not. We suggest these results reveal a human anatomical-functional homologue to monkey inferior parietal areas that respond to aversive stimuli by producing nocifensive muscle and limb movements.     

An 'automatic pilot' for the hand in human posterior parietal cortex: toward reinterpreting optic ataxia

We designed a protocol distinguishing between automatic and intentional motor reactions to changes in target location triggered at movement onset. In response to target jumps, but not to a similar change cued by a color switch, normal subjects often could not avoid automatically correcting fast aiming movements. This suggests that an 'automatic pilot' relying on spatial vision drives fast corrective arm movements that can escape intentional control. In a patient with a bilateral posterior parietal cortex (PPC) lesion, motor corrections could only be slow and deliberate. We propose that 'on-line' control is the most specific function of the PPC and that optic ataxia could result from a disruption of automatic hand guidance.     

Non-spatial,motor-specific activation in posterior parietal cortex

A localized cluster of neurons in macaque posterior parietal cortex, termed the parietal reach region (PRR), is activated when a reach is planned to a visible or remembered target. To explore the role of PRR in sensorimotor transformations, we tested whether cells would be activated when a reach is planned to an as-yet unspecified goal. Over one-third of PRR cells increased their firing after an instruction to prepare a reach, but not after an instruction to prepare a saccade, when the target of the movement remained unknown. A partially overlapping population (two-thirds of cells) was activated when the monkey was informed of the target location but not the type of movement to be made. Thus a subset of PRR neurons separately code spatial and effector-specific information, consistent with a role in specifying potential motor responses to particular targets.     

Thalamo-cortical projections to the posterior parietal cortex in the monkey

Thalamo-cortical projections to the posterior parietal cortex (PPC) were investigated electrophysiologically in the monkey. Cortical field potentials evoked by the thalamic stimulation were recorded with electrodes chronically implanted on the cortical surface and at a 2.0-3.0 mm cortical depth in the PPC. The stimulation of the nucleus lateralis posterior (LP), nucleus ventralis posterior lateralis pars caudalis (VPLc), and nucleus pulvinaris lateralis (Pul.l) and medialis (Pul.m) induced surface-negative, depth-positive potentials in the PPC. The LP and VPLc projected mainly to the superior parietal lobule (SPL) and the anterior bank of the intraparietal sulcus (IPS), and the Pul.m mainly to the inferior parietal lobule (IPL) and the posterior bank of the IPS. The Pul.l had projections to all of the SPL, the IPL and both the banks. The significance of the projections is discussed in connection with motor functions.     

Preparative activities in posterior parietal cortex for self-paced movement in monkeys

Cortical field potentials were recorded by electrodes implanted chronically on the surface and at a 2.0-3.0 mm depth in various cortices in monkeys performing self-paced finger, toe, mouth, hand or trunk movements. Surface-negative, depth-positive potentials (readiness potential) appeared in the posterior parietal cortex about 1.0 s before onset of every self-paced movement, as well as in the premotor, motor and somatosensory cortices. Somatotopical distribution was seen in the readiness potential in the posterior parietal cortex, although it was not so distinct as that in the motor or somatosensory cortex. This suggests that the posterior parietal cortex is involved in preparation for self-paced movement of any body part. This study contributes to the investigation of central nervous mechanisms of voluntary movements initiated by internal stimulus.     

Cerebello-thalamo-cortical projections to the posterior parietal cortex in the macaque monkey

This study reinvestigated the functional neuroanatomy of phonological and visual working memory in humans. Articulatory suppression was used to deprive the human subjects of species-specific verbal strategies in order to make the functional magnetic resonance imaging results more comparable to findings in non-human primates. Both phonological and visual working memory processes activated similar prefronto-parietal networks but were found to be differentially distributed along several cortical structures, in particular along the anterior and posterior parts of the intermediate frontal sulcus. These results suggest that a domain-specific topographical organization of neural working memory mechanisms in the primate brain is conserved in evolution. However, the findings also underline the critical dynamic influence that the additional availability of language may have on working memory processes and their functional implementation in the human brain.     

Thalamocortical connections of rat posterior parietal cortex.

The neuronal connections of rat posterior parietal cortex (PPC) have been examined using retrograde fluorescent axonal tracers. We have found that PPC receives thalamic input predominantly from the lateral posterior and lateral dorsal nuclei, and not from the ventrobasal nucleus, which projects to the rostrally adjacent hindlimb cortex, or from the dorsal lateral geniculate nucleus, which projects to the caudally adjacent visual association area. PPC has reciprocal corticocortical connections with medial agranular cortex and orbital cortex; together, these three cortical areas may function as a network for directed attention in rats.     

Opposite biases in salience-based selection for the left and right posterior parietal cortex

Visual selection is determined in part by the saliency of stimuli. We assessed the brain mechanisms determining attentional responses to saliency. Repetitive transcranial magnetic stimulation (rTMS) was applied to the left and right posterior parietal cortices (PPC) immediately before participants were asked to identify a compound letter. rTMS to the right PPC disrupted the guidance of attention toward salient stimuli, whereas rTMS to the left PPC affected the ability to bias selection away from salient stimuli. We conclude that right and left PPC have opposite roles in biasing selection to and from salient stimuli in the environment.     

Transcranial magnetic stimulation disrupts eye-hand interactions in the posterior parietal cortex

Recent neurophysiological studies have started to shed some light on the cortical areas that contribute to eye-hand coordination. In the present study we investigated the role of the posterior parietal cortex (PPC) in this process in normal, healthy subjects. This was accomplished by delivering single pulses of transcranial magnetic stimulation (TMS) over the PPC to transiently disrupt the putative contribution of this area to the processing of information related to eye-hand coordination. Subjects made open-loop pointing movements accompanied by saccades of the same required amplitude or by saccades that were substantially larger. Without TMS the hand movement amplitude was influenced by the amplitude of the corresponding saccade; hand movements accompanied by larger saccades were larger than those accompanied by smaller saccades. When TMS was applied over the left PPC just prior to the onset of the saccade, a marked reduction in the saccadic influence on manual motor output was observed. TMS delivered at earlier or later periods during the response had no effect. Taken together, these data suggest that the PPC integrates signals related to saccade amplitude with limb movement information just prior to the onset of the saccade.     

Integration of target and hand position signals in the posterior parietal cortex: effects of workspace and hand vision

Previous findings suggest the posterior parietal cortex (PPC) contributes to arm movement planning by transforming target and limb position signals into a desired reach vector. However, the neural mechanisms underlying this transformation remain unclear. In the present study we examined the responses of 109 PPC neurons as movements were planned and executed to visual targets presented over a large portion of the reaching workspace. In contrast to previous studies, movements were made without concurrent visual and somatic cues about the starting position of the hand. For comparison, a subset of neurons was also examined with concurrent visual and somatic hand position cues. We found that single cells integrated target and limb position information in a very consistent manner across the reaching workspace. Approximately two-thirds of the neurons with significantly tuned activity (42/61 and 30/46 for left and right workspaces, respectively) coded targets and initial hand positions separably, indicating no hand-centered encoding, whereas the remaining one-third coded targets and hand positions inseparably, in a manner more consistent with the influence of hand-centered coordinates. The responses of both types of neurons were largely invariant with respect to the presence or absence of visual hand position cues, suggesting their corresponding coordinate frames and gain effects were unaffected by cue integration. The results suggest that the PPC uses a consistent scheme for computing reach vectors in different parts of the workspace that is robust to changes in the availability of somatic and visual cues about hand position.