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.     

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.     

Processing of Coherent Visual Motion in Topographically Organized Visual Areas in Human Cerebral Cortex

Recent imaging studies in human subjects have demonstrated representations of global visual motion in medial parieto-occipital cortex (area V6) and posterior parietal cortex, the latter containing at least seven topographically organized areas along the intraparietal sulcus (IPS0–IPS5, SPL1). In this fMRI study we used topographic mapping procedures to delineate a total of 18 visual areas in human cerebral cortex and tested their responsiveness to coherent visual motion under conditions of controlled attention and fixation. Preferences for coherent visual motion as compared to motion noise as well as hemispheric asymmetries were assessed for contralateral, ipsilateral, and bilateral visual motion presentations. Except for areas V1–V4 and IPS3-5, all other areas showed stronger responses to coherent motion with the most significant activations found in V6, followed by MT/MST, V3A, IPS0-2 and SPL1. Hemispheric differences were negligible altogether suggesting that asymmetries in parietal cortex observed in cognitive tasks do not reflect differences in basic visual response properties. Interestingly, areas V6, MST, V3A, and areas along the intraparietal sulcus showed specific representations of coherent visual motion not only when presented in the hemifield primarily covered by the given visual representation but also when presented in the ipsilateral visual field. This finding suggests that coherent motion induces a switch in spatial representation in specialized motion areas from contralateral to full-field coding.     

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.     

Topographic maps in human frontal cortex revealed in memory-guided saccade and spatial working-memory tasks

We used fMRI at 3 Tesla and improved spatial resolution (2 x 2 x 2 mm(3)) to investigate topographic organization in human frontal cortex using memory-guided response tasks performed at 8 or 12 peripheral locations arranged clockwise around a central fixation point. The tasks required the location of a peripheral target to be remembered for several seconds after which the subjects either made a saccade to the remembered location (memory-guided saccade task) or judged whether a test stimulus appeared in the same or a slightly different location by button press (spatial working-memory task). With these tasks, we found two topographic maps in each hemisphere, one in the superior branch of precentral cortex and caudalmost part of the superior frontal sulcus, in the region of the human frontal eye field, and a second in the inferior branch of precentral cortex and caudalmost part of the inferior frontal sulcus, both of which greatly overlapped with activations evoked by visually guided saccades. In each map, activated voxels coded for saccade directions and memorized locations predominantly in the contralateral hemifield with neighboring saccade directions and memorized locations represented in adjacent locations of the map. Particular saccade directions or memorized locations were often represented in multiple locations of the map. The topographic activation patterns showed individual variability from subject to subject but were reproducible within subjects. Notably, only saccade-related activation, but no topographic organization, was found in the region of the human supplementary eye field in dorsomedial prefrontal cortex. Together these results show that topographic organization can be revealed outside sensory cortical areas using more complex behavioral tasks.     

Neural correlates of sustained spatial attention in human early visual cortex

Attention is thought to enhance perceptual performance at attended locations through top-down attention signals that modulate activity in visual cortex. Here, we show that activity in early visual cortex is sustained during maintenance of attention in the absence of visual stimulation. We used functional magnetic resonance imaging (fMRI) to measure activity in visual cortex while human subjects performed a visual detection task in which a variable-duration delay period preceded target presentation. Portions of cortical areas V1, V2, and V3 representing the attended part of the visual field exhibited sustained increases in activity throughout the delay period. Portions of these cortical areas representing peripheral, unattended parts of the visual field displayed sustained decreases in activity. The data were well fit by a model that assumed the sustained neural activity was constant in amplitude over a time period equal to that of the actual delay period for each trial. These results demonstrate that sustained attention responses are present in early visual cortex (including primary visual cortex), in the absence of a visual stimulus, and that these responses correlate with the allocation of visuospatial attention in both the spatial and temporal domains.     

Neural correlates of disparity-defined shape discrimination in the human brain

Binocular disparity, the slight differences between the images registered by our two eyes, provides an important cue when estimating the three-dimensional (3D) structure of the complex environment we inhabit. Sensitivity to binocular disparity is evident at multiple levels of the visual hierarchy in the primate brain, from early visual cortex to parietal and temporal areas. However, the relationship between activity in these areas and key perceptual functions that exploit disparity information for 3D shape perception remains an important open question. Here we investigate the link between human cortical activity and the perception of disparity-defined shape, measuring fMRI responses concurrently with psychophysical shape judgments. We parametrically degraded the coherence of shapes by shuffling the spatial position of dots whose disparity defined the 3D structure and investigated the effect of this stimulus manipulation on both cortical activity and shape discrimination. We report significant relationships between shape coherence and fMRI response in both dorsal (V3, hMT+/V5) and ventral (LOC) visual areas that correspond to the observers' discrimination performance. In contrast to previous suggestions of a dichotomy of disparity-related processes in the ventral and dorsal streams, these findings are consistent with proposed interactions between these pathways that may mediate a continuum of processes important in perceiving 3D shape from coarse contour segmentation to fine curvature estimation.     

The neural mechanisms of top-down attentional control

Selective visual attention involves dynamic interplay between attentional control systems and sensory brain structures. We used event-related functional magnetic resonance imaging (fMRI) during a cued spatial-attention task to dissociate brain activity related to attentional control from that related to selective processing of target stimuli. Distinct networks were engaged by attention-directing cues versus subsequent targets. Superior frontal, inferior parietal and superior temporal cortex were selectively activated by cues, indicating that these structures are part of a network for voluntary attentional control. This control biased activity in multiple visual cortical areas, resulting in selective sensory processing of relevant visual targets.     

Involvement of striate and extrastriate visual cortical areas in spatial attention

We investigated the cortical mechanisms of visual-spatial attention while subjects discriminated patterned targets within distractor arrays. Functional magnetic resonance imaging (fMRI) was used to map the boundaries of retinotopic visual areas and to localize attention-related changes in neural activity within several of those areas, including primary visual (striate) cortex. Event-related potentials (ERPs) and modeling of their neural sources, however, indicated that the initial sensory input to striate cortex at 50-55 milliseconds after the stimulus was not modulated by attention. The earliest facilitation of attended signals was observed in extrastriate visual areas, at 70-75 milliseconds. We hypothesize that the striate cortex modulation found with fMRI may represent a delayed, re-entrant feedback from higher visual areas or a sustained biasing of striate cortical neurons during attention. ERP recordings provide critical temporal information for analyzing the functional neuroanatomy of visual attention.     

Transient neural activity in human parietal cortex during spatial attention shifts

Observers viewing a complex visual scene selectively attend to relevant locations or objects and ignore irrelevant ones. Selective attention to an object enhances its neural representation in extrastriate cortex, compared with those of unattended objects, via top-down attentional control signals. The posterior parietal cortex is centrally involved in this control of spatial attention. We examined brain activity during attention shifts using rapid, event-related fMRI of human observers as they covertly shifted attention between two peripheral spatial locations. Activation in extrastriate cortex increased after a shift of attention to the contralateral visual field and remained high during sustained contralateral attention. The time course of activity was substantially different in posterior parietal cortex, where transient increases in activation accompanied shifts of attention in either direction. This result suggests that activation of the parietal cortex is associated with a discrete signal to shift spatial attention, and is not the source of a signal to continuously maintain the current attentive state.     

Frontoparietal activation with preparation for antisaccades

Several current models hold that frontoparietal areas exert cognitive control by biasing task-relevant processing in other brain areas. Previous event-related functional magnetic resonance imaging (fMRI) studies have compared prosaccades and antisaccades, which require subjects to look toward or away from a flashed peripheral stimulus, respectively. These studies found greater activation for antisaccades in frontal and parietal regions at the ends of long (>or=6 s) preparatory periods preceding peripheral stimulus presentation. Event-related fMRI studies using short preparatory periods (相似文献   

Comparison of memory- and visually guided saccades using event-related fMRI

Previous functional imaging studies have shown an increased hemodynamic signal in several cortical areas when subjects perform memory-guided saccades than that when they perform visually guided saccades using blocked trial designs. It is unknown, however, whether this difference results from sensory processes associated with stimulus presentation, from processes occurring during the delay period before saccade generation, or from an increased motor signal for memory-guided saccades. We conducted fMRI using an event-related paradigm that separated stimulus-related, delay-related, and saccade-related activity. Subjects initially fixated a central cross, whose color indicated whether the trial was a memory- or a visually guided trial. A peripheral stimulus was then flashed at one of 4 possible locations. On memory-guided trials, subjects had to remember this location for the subsequent saccade, whereas the stimulus was a distractor on visually guided trials. Fixation cross disappearance after a delay period was the signal either to generate a memory-guided saccade or to look at a visual stimulus that was flashed on visually guided trials. We found slightly greater stimulus-related activation for visually guided trials in 3 right prefrontal regions and right rostral intraparietal sulcus (IPS). Memory-guided trials evoked greater delay-related activity in right posterior inferior frontal gyrus, right medial frontal eye field, bilateral supplementary eye field, right rostral IPS, and right ventral IPS but not in middle frontal gyrus. Right precentral gyrus and right rostral IPS exhibited greater saccade-related activation on memory-guided trials. We conclude that activation differences revealed by previous blocked experiments have different sources in different areas and that cortical saccade regions exhibit delay-related activation differences.     

Frequency-Following and Connectivity of Different Visual Areas in Response to Contrast-Reversal Stimulation

The sensitivity of visual areas to different temporal frequencies, as well as the functional connections between these areas, was examined using magnetoencephalography (MEG). Alternating circular sinusoids (0, 3.1, 8.7 and 14 Hz) were presented to foveal and peripheral locations in the visual field to target ventral and dorsal stream structures, respectively. It was hypothesized that higher temporal frequencies would preferentially activate dorsal stream structures. To determine the effect of frequency on the cortical response we analyzed the late time interval (220–770 ms) using a multi-dipole spatio-temporal analysis approach to provide source locations and timecourses for each condition. As an exploratory aspect, we performed cross-correlation analysis on the source timecourses to determine which sources responded similarly within conditions. Contrary to predictions, dorsal stream areas were not activated more frequently during high temporal frequency stimulation. However, across cortical sources the frequency-following response showed a difference, with significantly higher power at the second harmonic for the 3.1 and 8.7 Hz stimulation and at the first and second harmonics for the 14 Hz stimulation with this pattern seen robustly in area V1. Cross-correlations of the source timecourses showed that both low- and high-order visual areas, including dorsal and ventral stream areas, were significantly correlated in the late time interval. The results imply that frequency information is transferred to higher-order visual areas without translation. Despite the less complex waveforms seen in the late interval of time, the cross-correlation results show that visual, temporal and parietal cortical areas are intricately involved in late-interval visual processing.     

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.     

Acetylcholinesterase histochemistry in the macaque thalamus reveals territories selectively connected to frontal, parietal and temporal association cortices

The patterns of histochemical staining for acetylcholinesterase (AChE) activity in the macaque thalamus were analyzed and compared with the distribution of cells and terminals labeled from injections of axonal tracers in the dorsolateral and orbital prefrontal cortex, in area 7a of the posterior parietal cortex and in the polysensory cortex of the superior temporal sulcus. AChE histochemistry is very useful in delineating the thalamic nuclei connected with the association cortex and in uncovering thalamic subdivisions that are barely evident on cytoarchitectonic grounds. Moreover, AChE activity reveals previously unrecognized heterogeneities within several thalamic nuclei, like the ventral anterior (VA), where a new ventromedial subdivision (VAvm) is described, the medial pulvinar (PuIM) or the mediodorsal nucleus (MD). In this nucleus three distinct chemical domains are present: the medial, ventral and lateral sectors characterized by low, moderate and high AChE activities, respectively. The staining pattern of the lateral sector is markedly heterogeneous with patches of intense AChE activity surrounded by a moderately stained matrix. The MD medial sector is connected with the orbitofrontal cortex, whereas the AChE-rich patches in the lateral sector are selectively connected with the dorsolateral prefrontal, parietal and temporal association cortices. In the PulM, a dorsomedial AChE-rich patch is selectively connected with the orbitofrontal cortex, whereas the surrounding territory, which shows moderate AChE activity, is preferentially connected with the parietal and temporal cortices. Chemically specific domains in the anterior, ventral anterior, midline, and intralaminar thalamic nuclei are also connected with the examined association cortices. These findings indicate that the topographic patterns of the thalamo-cortical connections of primate association areas conform to the chemical architecture of the thalamus. This implies that because each cortical area is connected to a particular set of thalamic regions, the influence of the thalamus on cortical function is exclusive for each area, highly diverse among the various association areas, and subject to a wide range of modulation at the thalamic level.     

fMRI signal increases and decreases in cortical areas during small-field optokinetic stimulation and central fixation

Small-field optokinetic nystagmus (OKN) was performed in seven healthy volunteers in order to analyze the activation and deactivation patterns of visual motion, ocular motor, and multisensory vestibular cortex areas by means of fMRI during coherent visual motion stimulation. BOLD signal decreases (deactivations) were found in the first and second long insular gyri and retroinsular areas (the human homologue of the parietoinsular vestibular cortex and the visual posterior sylvian area in the monkey) of both hemispheres, extending into the transverse temporal gyrus and inferior-anterior parts of the superior temporal gyrus (BA 22), and the precentral gyri at two separate sites (BA 4 and 6). Further deactivations were found in cranioposterior parts of the superior temporal gyrus (BA 22) and the adjacent inferior parietal lobule (BA 40), anterior cingulate gyrus, hippocampus, and corpus callosum. Most of these BOLD signal decreases involved parts of the "multisensory vestibular cortical circuit". These findings support the concept of a reciprocally inhibitory visual–vestibular interaction that has now been demonstrated not only for large-field visual motion stimulation that induces vection (without eye movements) but also for optokinetically induced eye movements (without vection). The functional significance of this concept may be related to the perception of self-motion, since both large-field visual motion stimulation and optokinetic nystagmus are linked to the visual control of self-motion. With respect to activation of the cortical ocular motor system two separate and distinct areas of activations were delineated in the precentral sulcus of both hemispheres, one ventrolaterally (in BA 9) and the other dorsomedially at the junction of the superior frontal sulcus with the precentral sulcus (in BA 6). Both probably correspond to different subregions of the frontal eye field and the premotor cortex for the ocular motor performance of OKN. Electronic Publication     

Tonotopic organization in human auditory cortex revealed by progressions of frequency sensitivity

Functional neuroimaging experiments have revealed an organization of frequency-dependent responses in human auditory cortex suggestive of multiple tonotopically organized areas. Numerous studies have sampled cortical responses to isolated narrow-band stimuli, revealing multiple locations in auditory cortex at which the position of response varies systematically with frequency content. Because appropriate anatomical or functional grouping of these distinct frequency-dependent responses is uncertain, the number and location of tonotopic mappings within human auditory cortex remains unclear. Further, sampling does not address whether the observed mappings exhibit continuity as a function of position. This functional magnetic resonance imaging study used frequency-swept stimuli to identify progressions of frequency sensitivity across the cortical surface. The center-frequency of narrow-band, amplitude-modulated noise was slowly swept between 125 and 8,000 Hz. The latency of response relative to sweep onset was determined for each cortical surface location. Because frequency varied systematically with time, response latency indicated the frequency to which a location was maximally sensitive. Areas of cortex exhibiting a progressive change in response latency with position were considered tonotopically organized. There exist two main findings. First, six progressions of frequency sensitivity (i.e., tonotopic mappings) were repeatably observed in the superior temporal plane. Second, the locations of the higher- and lower-frequency endpoints of these progressions were approximately congruent with regions reported to be most responsive to discrete higher- and lower-frequency stimuli. Based on these findings and previous anatomical work, we propose a correspondence between these progressions and anatomically defined cortical areas, suggesting that five areas in human auditory cortex exhibit at least six tonotopic organizations.     

Retinotopic maps and foveal suppression in the visual cortex of amblyopic adults

Amblyopia is a developmental visual disorder associated with loss of monocular acuity and sensitivity as well as profound alterations in binocular integration. Abnormal connections in visual cortex are known to underlie this loss, but the extent to which these abnormalities are regionally or retinotopically specific has not been fully determined. This functional magnetic resonance imaging (fMRI) study compared the retinotopic maps in visual cortex produced by each individual eye in 19 adults (7 esotropic strabismics, 6 anisometropes and 6 controls). In our standard viewing condition, the non-tested eye viewed a dichoptic homogeneous mid-level grey stimulus, thereby permitting some degree of binocular interaction. Regions-of-interest analysis was performed for extrafoveal V1, extrafoveal V2 and the foveal representation at the occipital pole. In general, the blood oxygenation level-dependent (BOLD) signal was reduced for the amblyopic eye. At the occipital pole, population receptive fields were shifted to represent more parafoveal locations for the amblyopic eye, compared with the fellow eye, in some subjects. Interestingly, occluding the fellow eye caused an expanded foveal representation for the amblyopic eye in one early–onset strabismic subject with binocular suppression, indicating real-time cortical remapping. In addition, a few subjects actually showed increased activity in parietal and temporal cortex when viewing with the amblyopic eye. We conclude that, even in a heterogeneous population, abnormal early visual experience commonly leads to regionally specific cortical adaptations.     

Differentiated parietal connectivity of frontal regions for “what” and “where” memory

In a previous meta-analysis across almost 200 neuroimaging experiments, working memory for object location showed significantly stronger convergence on the posterior superior frontal gyrus, whereas working memory for identity showed stronger convergence on the posterior inferior frontal gyrus (dorsal to, but overlapping with Brodmann’s area BA 44). As similar locations have been discussed as part of a dorsal frontal—superior parietal reach system and an inferior frontal grasp system, the aim of the present study was to test whether the regions of working-memory related “what” and “where” processing show a similar distinction in parietal connectivity. The regions that were found in the previous meta-analysis were used as seeds for functional connectivity analyses using task-based meta-analytic connectivity modelling and task-independent resting state correlations. While the ventral seed showed significantly stronger connectivity with the bilateral intraparietal sulcus (IPS), the dorsal seed showed stronger connectivity with the bilateral posterior inferior parietal and the medial superior parietal lobule. The observed connections of regions involved in memory for object location and identity thus clearly demonstrate a distinction into separate pathways that resemble the parietal connectivity patterns of the dorsal and ventral premotor cortex in non-human primates and humans. It may hence be speculated that memory for a particular location and reaching towards it as well as object memory and finger positioning for manipulation may rely on shared neural systems. Moreover, the ensuing regions, in turn, featured differential connectivity with the bilateral ventral and dorsal extrastriate cortex, suggesting largely segregated bilateral connectivity pathways from the dorsal visual cortex via the superior and inferior parietal lobules to the dorsal posterior frontal cortex and from the ventral visual cortex via the IPS to the ventral posterior frontal cortex that may underlie action and cognition.