There is good evidence that early visual processing involves the coding of different features in independent brain regions. A major question, then, is how we see the world in an integrated manner, in which the different features are “bound” together. A standard account of this has been that feature binding depends on attention to the stimulus, which enables only the relevant features to be linked together [Treisman, A., & Gelade, G. A feature-integration theory of attention. Cognitive Psychology, 12, 97–136, 1980]. Here we test this influential idea by examining whether, in patients showing visual extinction, the processing of otherwise unconscious (extinguished) stimuli is modulated by presenting objects in their correct (familiar) color. Correctly colored objects showed reduced extinction when they had a learned color, and this color matched across the ipsi- and contralesional items (red strawberry + red tomato). In contrast, there was no reduction in extinction under the same conditions when the stimuli were colored incorrectly (blue strawberry + blue tomato; Experiment 1). The result was not due to the speeded identification of a correctly colored ipsilesional item, as there was no benefit from having correctly colored objects in different colors (red strawberry + yellow lemon; Experiment 2). There was also no benefit to extinction from presenting the correct colors in the background of each item (Experiment 3). The data suggest that learned color–form binding can reduce extinction even when color is irrelevant for the task. The result is consistent with preattentive binding of color and shape for familiar stimuli.

The ability to search efficiently for visual targets among distractors can break down after a variety of brain lesions, but the specific processes affected by the lesions are unclear. We examined search over space (conjunction search) and over time plus space (preview search) in a consecutive series of patients with acquired brain lesions. We also assessed performance on standard neuropsychological measures of visuospatial short-term memory (Corsi Block), sustained attention and memory updating (the contrast between forward and backward digit span), and visual neglect. Voxel-based morphometry analyses revealed regions in the occipital (middle occipital gyrus), posterior parietal (angular gyrus), and temporal cortices (superior and middle temporal gyri extending to the insula), along with underlying white matter pathways, associated with poor search. Going beyond standard voxel-based morphometry analyses, we then report correlation measures of structural damage in these regions and the independent neuropsychological measures of other cognitive functions. We find distinct patterns of correlation in areas linked to poor search, suggesting that the areas play functionally different roles in search. We conclude that neuropsychological disorders of search can be linked to necessary and distinct cognitive functions, according to the site of lesion.

Deficits in the ability to draw objects, despite apparently intact perception and motor abilities, are defined as constructional apraxia. Constructional deficits, often diagnosed based on performance on copying complex figures, have been reported in a range of pathologies, perhaps reflecting the contribution of several underlying factors to poor figure drawing. The current study provides a comprehensive analysis of brain–behavior relationships in drawing disorders based on data from a large cohort of subacute stroke patients ( n = 358) using whole-brain voxel-wise statistical analyses linked to behavioral measures from a complex figure copy task. We found that (i) overall poor performance on figure copying was associated with subcortical lesions (BG and thalamus), (ii) lateralized deficits with respect to the midline of the viewer were associated with lesions within the posterior parietal lobule, and (iii) spatial positioning errors across the entire figure were associated with lesions within visual processing areas (lingual gyrus and calcarine) and the insula. Furthermore, deficits in reproducing global aspects of form were associated with damage to the right middle temporal gyrus, whereas deficits in representing local features were linked to the left hemisphere lesions within calcarine cortex (extending into the cuneus and precuneus), the insula, and the TPJ. The current study provides strong evidence that impairments in separate cognitive mechanisms (e.g., spatial coding, attention, motor execution, and planning) linked to different brain lesions contribute to poor performance on complex figure copying tasks. The data support the argument that drawing depends on several cognitive processes operating via discrete neuronal networks and that constructional problems as well as hierarchical and spatial representation deficits contribute to poor figure copying.

Humans typically rely upon vision to identify object shape, but we can also recognize shape via touch (haptics). Our haptic shape recognition ability raises an intriguing question: To what extent do visual cortical shape recognition mechanisms support haptic object recognition? We addressed this question using a haptic fMRI repetition design, which allowed us to identify neuronal populations sensitive to the shape of objects that were touched but not seen. In addition to the expected shape-selective fMRI responses in dorsal frontoparietal areas, we observed widespread shape-selective responses in the ventral visual cortical pathway, including primary visual cortex. Our results indicate that shape processing via touch engages many of the same neural mechanisms as visual object recognition. The shape-specific repetition effects we observed in primary visual cortex show that visual sensory areas are engaged during the haptic exploration of object shape, even in the absence of concurrent shape-related visual input. Our results complement related findings in visually deprived individuals and highlight the fundamental role of the visual system in the processing of object shape.

Hierarchical models of visual processing assume that global pattern recognition is contingent on the progressive integration of local elements across larger spatial regions, operating from early through intermediate to higher-level cortical regions. Here, we present results from neuropsychological fMRI that refute such models. We report two patients, one with lesions to intermediate ventral regions and the other with damage around the intraparietal sulcus (IPS). The patient with ventral damage showed normal behavioral and BOLD responses to global Glass patterns. The patient with IPS damage was impaired in discriminating global patterns and showed a lack of significant responses to these patterns in intermediate visual regions spared by the lesion. However, this patient did show BOLD activity to translational patterns, where local element relations are important. These results suggest that activation of intermediate ventral regions is not necessary to code global patterns; instead global patterns are coded in a heterarchical fashion. High-level regions of dorsal cortex are necessary to generate global pattern coding in intermediate ventral regions; in contrast, local integration processes are not sufficient.

There is considerable evidence that there are anatomically and functionally distinct pathways for action and object recognition. However, little is known about how information about action and objects is integrated. This study provides fMRI evidence for task-based selection of brain regions associated with action and object processing, and on how the congruency between the action and the object modulates neural response. Participants viewed videos of objects used in congruent or incongruent actions and attended either to the action or the object in a one-back procedure. Attending to the action led to increased responses in a fronto-parietal action-associated network. Attending to the object activated regions within a fronto-inferior temporal network. Stronger responses for congruent action–object clips occurred in bilateral parietal, inferior temporal, and putamen. Distinct cortical and thalamic regions were modulated by congruency in the different tasks. The results suggest that (i) selective attention to action and object information is mediated through separate networks, (ii) object–action congruency evokes responses in action planning regions, and (iii) the selective activation of nuclei within the thalamus provides a mechanism to integrate task goals in relation to the congruency of the perceptual information presented to the observer.

This study is the first to assess lesion–symptom relations for subitizing and counting impairments in a large sample of neuropsychological patients (41 patients) using an observer-independent voxel-based approach. We tested for differential effects of enumerating small versus large numbers of items while controlling for hemianopia and visual attention deficits. Overall impairments in the enumeration of any numbers (small or large) were associated with an extended network, including bilateral occipital and fronto-parietal regions. Within this network, severe impairments in accuracy when enumerating small sets of items (in the subitizing range) were associated with damage to the left posterior occipital cortex, bilateral lateral occipital and right superior frontal cortices. Lesions to the right calcarine extending to the precuneus led to patients serially counting even small numbers of items (indicated by a steep response slope), again demonstrating an impaired subitizing ability. In contrast, impairments in counting large numerosities were associated with damage to the left intraparietal sulcus. The data support the argument for some distinctive processes and neural areas necessary to support subitization and counting with subitizing relying on processes of posterior occipital cortex and with counting associated with processing in the parietal cortex.

Because of our limited processing capacity, different elements of the visual scene compete for the allocation of processing resources. One of the most striking deficits in visual selection is simultanagnosia, a rare neuropsychological condition characterized by impaired spatial awareness of more than one object at time. To decompose the neuroanatomical substrates of the syndrome and to gain insights into the structural and functional organization of visuospatial attention, we performed a systematic evaluation of lesion patterns in a group of simultanagnosic patients compared with patients with either (i) unilateral visuospatial deficits (neglect and/or extinction) or (ii) bilateral posterior lesions without visuospatial deficits, using overlap/subtraction analyses, estimation of lesion volume, and a lesion laterality index. We next used voxel-based morphometry to assess the link between different visuospatial deficits and gray matter and white matter (WM) damage. Lesion overlap/subtraction analyses, lesion laterality index, and voxel-based morphometry measures converged to indicate that bilateral parieto-occipital WM disconnections are both distinctive and necessary to create symptoms associated with simultanagnosia. We also found that bilateral gray matter damage within the middle frontal area (BA 46), cuneus, calacarine, and parieto-occipital fissure as well as right hemisphere parietal lesions within intraparietal and postcentral gyri were associated with simultanagnosia. Further analysis of the WM based on tractography revealed associations with bilateral damage to major pathways within the visuospatial attention network, including the superior longitudinal fasciculus, the inferior fronto-occipital fasciculus, and the inferior longitudinal fasciculus. We conclude that damage to the parieto-occipital regions and the intraparietal sulcus, together, with bilateral WM disconnections within the visuosptial attention network, contribute to poor visual processing of multiple objects and the loss of processing speed characteristic of simultanagnosia.

Brain activity was recorded while participants engaged in a difficult visual search task for a target defined by the spatial configuration of its component elements. The search displays were segmented by time (a preview then a search display), by motion, or were unsegmented. A preparatory network showed activity to the preview display, in the time but not in the motion segmentation condition. A region of the precuneus showed (i) higher activation when displays were segmented by time or by motion, and (ii) correlated activity with larger segmentation benefits behaviorally, regardless of the cue. Additionally, the results revealed that success in temporal segmentation was correlated with reduced activation in early visual areas, including V1. The results depict partially overlapping brain networks for segmentation in search by time and motion, with both cue-independent and cue-specific mechanisms.

Visual evoked responses were monitored while participants searched for a target (e.g., bird ) in a four-object display that could include a semantically related distractor (e.g., fish ). The occurrence of both the target and the semantically related distractor modulated the N2pc response to the search display: The N2pc amplitude was more pronounced when the target and the distractor appeared in the same visual field, and it was less pronounced when the target and the distractor were in opposite fields, relative to when the distractor was absent. Earlier components (P1, N1) did not show any differences in activity across the different distractor conditions. The data suggest that semantic distractors influence early stages of selecting stimuli in multielement displays.

Working memory (WM) and visual selection processes interact in a reciprocal fashion based on overlapping representations abstracted from the physical characteristics of stimuli. Here, we assessed the neural basis of this interaction using facial expressions that conveyed emotion information. Participants memorized an emotional word for a later recognition test and then searched for a face of a particular gender presented in a display with two faces that differed in gender and expression. The relation between the emotional word and the expressions of the target and distractor faces was varied. RTs for the memory test were faster when the target face matched the emotional word held in WM (on valid trials) relative to when the emotional word matched the expression of the distractor (on invalid trials). There was also enhanced activation on valid compared with invalid trials in the lateral orbital gyrus, superior frontal polar (BA 10), lateral occipital sulcus, and pulvinar. Re-presentation of the WM stimulus in the search display led to an earlier onset of activity in the superior and inferior frontal gyri and the anterior hippocampus irrespective of the search validity of the re-presented stimulus. The data indicate that the middle temporal and prefrontal cortices are sensitive to the reappearance of stimuli that are held in WM, whereas a fronto-thalamic occipital network is sensitive to the behavioral significance of the match between WM and targets for selection. We conclude that these networks are modulated by high-level matches between the contents of WM, behavioral goals, and current sensory input.

Attentional cues can trigger activity in the parietal cortex in anticipation of visual displays, and this activity may, in turn, induce changes in other areas of the visual cortex, hence, implementing attentional selection. In a recent TMS study [Mevorach, C., Humphreys, G. W., & Shalev, L. Opposite biases in salience-based selection for the left and right posterior parietal cortex. Nature Neuroscience, 9, 740–742, 2006b], it was shown that the posterior parietal cortex (PPC) can utilize the relative saliency (a nonspatial property) of a target and a distractor to bias visual selection. Furthermore, selection was lateralized so that the right PPC is engaged when salient information must be selected and the left PPC when the salient information must be ignored. However, it is not clear how the PPC implements these complementary forms of selection. Here we used on-line triple-pulse TMS over the right or left PPC prior to or after the onset of global/local displays. When delivered after the onset of the display, TMS to the right PPC disrupted the selection of the more salient aspect of the hierarchical letter. In contrast, left PPC TMS delivered prior to the onset of the stimulus disrupted responses to the lower saliency stimulus. These findings suggest that selection and suppression of saliency, rather than being “two sides of the same coin,” are fundamentally different processes. Selection of saliency seems to operate reflexively, whereas suppression of saliency relies on a preparatory phase that “sets up” the system in order to effectively ignore saliency.

Theory of mind (ToM) reasoning may involve a multiplicity of processes, including an initial stage, where cues relevant for social processes are detected and decoded, and a mentalizing stage, where the decoded information is used to reason about mental states. Here we report that the processing of lower-order facial cues relevant to social judgments can be relatively spared in patients with impaired ToM reasoning. We discuss the implications for understanding the mechanisms underlying social judgments in brain-lesioned patients.

Neuropsychological and functional imaging studies have suggested a general right hemisphere advantage for processing global visual information and a left hemisphere advantage for processing local information. In contrast, a recent transcranial magnetic stimulation study [Mevorach, C., Humphreys, G. W., & Shalev, L. Opposite biases in salience-based selection for the left and right posterior parietal cortex. Nature Neuroscience, 9, 740–742, 2006b] demonstrated that functional lateralization of selection in the parietal cortices on the basis of the relative salience of stimuli might provide an alternative explanation for previous results. In the present study, we applied a whole-brain analysis of the functional magnetic resonance signal when participants responded to either the local or the global levels of hierarchical figures. The task (respond to local or global) was crossed with the saliency of the target level (local salient, global salient) to provide, for the first time, a direct contrast between brain activation related to the stimulus level and that related to relative saliency. We found evidence for lateralization of salience-based selection but not for selection based on the level of processing. Activation along the left intraparietal sulcus (IPS) was found when a low saliency stimulus had to be selected irrespective of its level. A control task showed that this was not simply an effect of task difficulty. The data suggest a specific role for regions along the left IPS in salience-based selection, supporting the argument that previous reports of lateralized responses to local and global stimuli were contaminated by effects of saliency.

Visual search is facilitated if half the distractors are presented as a preview prior to the presentation of the target and second set of distractors-the preview benefit [Watson, D. G., & Humphreys, G. W. Visual marking: Prioritizing selection for new objects by top-down attentional inhibition of old objects. Psychological Review, 104, 90-122, 1997]. On one account, the preview advantage is due to automatic capture of attention by the onsets in the second, search display [Donk, M., & Theeuwes, J. Visual marking beside the mark: Prioritizing selection by abrupt onsets. Perception & Psychophysics, 93, 891-900, 2001]. We provide a neuropsychological test of this assertion. We examined onset capture and preview benefits in search in a group of neuropsychological patients with unilateral parietal damage. We demonstrate a normal pattern of performance when patients detected targets defined by onsets relative to those defined by offsets, irrespective of whether the onset target fell contra-or ipsilateral to the lesion. In contrast, there was a normal preview benefit in search only for ipsilesional targets, and preview search was impaired in the contralesional field. The data demonstrate that the preview benefit can dissociate from the onset advantage in search, and that onsets remain strongly weighted for attention even in the contralesional field of patients with parietal lesions.

A model of the functional and anatomical basis of belief reasoning is essential for understanding the relationship between belief reasoning and other cognitive processes in both normal development and pathology. Studies of brain-damaged patients can give valuable insights into the nature of belief processing but pose unique methodological problems. The current study addresses these problems by using a nonlinguistic belief-reasoning task with substantially reduced executive demands. A case series of 12 brain-damaged patients is presented. The belief-reasoning errors of four patients with damage to the prefrontal cortex appeared to arise from these patients' executive function problems. The belief-reasoning errors of three patients with damage to the temporo-parietal junction could not easily be accounted for in this way, raising the possibility that this brain region has a necessary role in representing beliefs, rather than handling the executive demands of belief-reasoning tasks. We discuss the importance of gaining empirical evidence about the scope of “theory of mind” impairments, and the important role for neuropsychological studies in this project.

The early stages of visual word recognition were investigated by scanning participants using PET as they took part in implicit and explicit reading tasks with visually disrupted stimuli. CaSe MiXiNg has been shown in behavioral studies to increase reaction times (RTs) in naming and other word recognition tasks. In this study, we found that during both an implicit (feature detection) task and an explicit word-naming task, mixed-case words compared to same-case words produced increased activation in an area of the right parietal cortex previously associated with visual attention. No effect of case was found in this area for pseudowords or consonant strings. Further, lowering the contrast of the stimuli slowed RTs as much as case mixing, but did not lead to the same increase in right parietal activation. No significant effect of case mixing was observed in left-hemisphere language areas. The results suggest that reading mixed-case words requires increased attentional processing. However, later word recognition processes may be relatively unaffected by the disruption in presentation.