Modulations of sensory processing in early visual areas are thought to play an important role in conscious perception. To date, most empirical studies focused on effects occurring before or during visual presentation. By contrast, several emerging theories postulate that sensory processing and conscious visual perception may also crucially depend on late top–down influences, potentially arising after a visual display. To provide a direct test of this, we performed an fMRI study using a postcued report procedure. The ability to report a target at a specific spatial location in a visual display can be enhanced behaviorally by symbolic auditory postcues presented shortly after that display. Here we showed that such auditory postcues can enhance target-specific signals in early human visual cortex (V1 and V2). For postcues presented 200 msec after stimulus termination, this target-specific enhancement in visual cortex was specifically associated with correct conscious report. The strength of this modulation predicted individual levels of performance in behavior. By contrast, although later postcues presented 1000 msec after stimulus termination had some impact on activity in early visual cortex, this modulation no longer related to conscious report. These results demonstrate that within a critical time window of a few hundred milliseconds after a visual stimulus has disappeared, successful conscious report of that stimulus still relates to the strength of top–down modulation in early visual cortex. We suggest that, within this critical time window, sensory representation of a visual stimulus is still under construction and so can still be flexibly influenced by top–down modulatory processes.

Regions of the occipital and temporal lobes, including a region in the fusiform gyrus (FG), have been proposed to constitute a “core” visual representation system for faces, in part because they show face selectivity and face repetition suppression. But recent fMRI studies of developmental prosopagnosics (DPs) raise questions about whether these measures relate to face processing skills. Although DPs manifest deficient face processing, most studies to date have not shown unequivocal reductions of functional responses in the proposed core regions. We scanned 15 DPs and 15 non-DP control participants with fMRI while employing factor analysis to derive behavioral components related to face identification or other processes. Repetition suppression specific to facial identities in FG or to expression in FG and STS did not show compelling relationships with face identification ability. However, we identified robust relationships between face selectivity and face identification ability in FG across our sample for several convergent measures, including voxel-wise statistical parametric mapping, peak face selectivity in individually defined “fusiform face areas” (FFAs), and anatomical extents (cluster sizes) of those FFAs. None of these measures showed associations with behavioral expression or object recognition ability. As a group, DPs had reduced face-selective responses in bilateral FFA when compared with non-DPs. Individual DPs were also more likely than non-DPs to lack expected face-selective activity in core regions. These findings associate individual differences in face processing ability with selectivity in core face processing regions. This confirms that face selectivity can provide a valid marker for neural mechanisms that contribute to face identification ability.

“Regret aversion” is proposed to explain a tendency to avoid future choices that have induced past regret. However, regret might also motivate us to repeat previous regret-related choices to make up for their previous selection, a behavior resembling “chasing” in the context of gambling. In the current experiment, we acquired fMRI brain data while participants placed monetary bets on repeated gambles. Behaviorally, participants showed a tendency to repeat previously regret-related choices (operationalized as those leading to an outcome worse than what might have been), an effect restricted to early runs of the task. At gamble outcome, we show a reduction in ventral striatal activity for regret-related relative to relief-related outcomes. Critically, this modulation was only seen when subjects were responsible for the bet choice. Activity in dorsal striatum was associated with an influence of previous regret on participants' subsequent choices, which is evident in increased activity when regret-related choices were repeated, relative to avoided, on the next trial. Our findings indicate that regret can lead to choice repetition as if seeking to make up for our mistakes and in so doing may lead to subsequent chasing behavior.

We used concurrent TMS–fMRI to test directly for hemispheric differences in causal influences of the right or left fronto-parietal cortex on activity (BOLD signal) in the human occipital cortex. Clinical data and some behavioral TMS studies have been taken to suggest right-hemisphere specialization for top–down modulation of vision in humans, based on deficits such as spatial neglect or extinction in lesioned patients, or findings that TMS to right (vs. left) fronto-parietal structures can elicit stronger effects on visual performance. But prior to the recent advent of concurrent TMS and neuroimaging, it was not possible to directly examine the causal impact of one (stimulated) brain region upon others in humans. Here we stimulated the frontal or intraparietal cortex in the left or right hemisphere with TMS, inside an MR scanner, while measuring with fMRI any resulting BOLD signal changes in visual areas V1–V4 and V5/MT+. For both frontal and parietal stimulation, we found clear differences between effects of right- versus left-hemisphere TMS on activity in the visual cortex, with all differences significant in direct statistical comparisons. Frontal TMS over either hemisphere elicited similar BOLD decreases for central visual field representations in V1–V4, but only right frontal TMS led to BOLD increases for peripheral field representations in these regions. Hemispheric differences for effects of parietal TMS were even more marked: Right parietal TMS led to strong BOLD changes in V1–V4 and V5/MT+, but left parietal TMS did not. These data directly confirm that the human frontal and parietal cortex show right-hemisphere specialization for causal influences on the visual cortex.

The possible impact upon human visual cortex from saccades to remembered target locations was investigated using functional magnetic resonance imaging (fMRI). A specific location in the upper-right or upper-left visual quadrant served as the saccadic target. After a delay of 2400 msec, an auditory signal indicated whether to execute a saccade to that location (go trial) or to cancel the saccade and remain centrally fixated (no-go). Group fMRI analysis revealed activation specific to the remembered target location for executed saccades, in the contralateral lingual gyrus. No-go trials produced similar, albeit significantly reduced, effects. Individual retinotopic mapping confirmed that on go trials, quadrant-specific activations arose in those parts of ventral V1, V2, and V3 that coded the target location for the saccade, whereas on no-go trials, only the corresponding parts of V2 and V3 were significantly activated. These results indicate that a spatial–motor saccadic task (i.e., making an eye movement to a remembered location) is sufficient to activate retinotopic visual cortex spatially corresponding to the target location, and that this activation is also present (though reduced) when no saccade is executed. We discuss the implications of finding that saccades to remembered locations can affect early visual cortex, not just those structures conventionally associated with eye movements, in relation to recent ideas about attention, spatial working memory, and the notion that recently activated representations can be “refreshed” when needed.

We compared the contribution of featural information and second-order spatial relations (spacing between features) in face processing. A fully factorial design has the same or different “features” (eyes, mouth, and nose) across two successive displays, whereas, orthogonally, the second-order spatial relations between those features were the same or different. The range of such changes matched the possibilities within the population of natural face images. Behaviorally, we found that judging whether two successive faces depicted the same person was dominated by features, although second-order spatial relations also contributed. This influence of spatial relations correlated, for individual subjects, with their skill at recognition of faces (as famous, or as previously exposed) in separate behavioral tests. Using the same repetition design in functional magnetic resonance imaging, we found feature-dependent effects in the lateral occipital and right fusiform regions. In addition, there were spatial relation effects in the bilateral inferior occipital gyrus and right fusiform that correlated with individual differences in (separately measured) behavioral sensitivity to those changes. The results suggest that featural and second-order spatial relation aspects of faces make distinct contributions to behavioral discrimination and recognition, with features contributing most to face discrimination and second-order spatial relational aspects correlating best with recognition skills. Distinct neural responses to these aspects were found with functional magnetic resonance imaging, particularly when individual skills were taken into account for the impact of second-order spatial relations.

Right hemisphere lesions often lead to severe disorders in spatial awareness and behavior, such as left hemispatial neglect. Neglect involves not only pathological biases in attention and exploration but also deficits in internal representations of space and spatial working memory. Here we designed a new paradigm to test whether one potential component may involve a failure to maintain an updated representation of visual locations across delays when a gaze-shift intervenes. Right hemisphere patients with varying severity of left spatial neglect had to encode a single target location and retain it across an interval of 2 or 3 sec, during which the target was transiently removed, before a subsequent probe appeared for a same/different location judgment. During the delay, gaze could have to shift to either side of the remembered location, or no gaze-shift was required. Patients showed a dramatic loss of memory for target location after shifting gaze to its right (toward their “intact” ipsilesional side), but not after leftward gaze-shifts. Such impairment arose even when the target initially appeared in the right visual field, before being updated leftward due to right gaze, and even when gaze returned to the screen center before the memory probe was presented. These findings indicate that location information may be permanently degraded when the target has to be remapped leftward in gaze-centric representations. Across patients, the location-memory deficit induced by rightward gaze-shifts correlated with left neglect severity on several clinical tests. This paradoxical memory deficit, with worse performance following gaze-shifts to the “intact” side of space, may reflect losses in gaze-centric representations of space that normally remap a remembered location dynamically relative to current gaze. Right gaze-shifts may remap remembered locations leftward, into damaged representations, whereas left gaze-shifts will require remapping rightward, into intact representations. Our findings accord with physiological data on normal remapping mechanisms in the primate brain but demonstrate for the first time their impact on perceptual spatial memory when damaged, while providing new insights into possible components that may contribute to the neglect syndrome.

Attending to the location of an expected visual target can lead to anticipatory activations in spatiotopic occipital cortex, emerging before target onset. But less is known about how the brain may prepare for a distractor at a known location remote from the target. In a psychophysical experiment, we found that trial-to-trial advance knowledge about the presence of a distractor in the target-opposite hemifield significantly reduced its behavioral cost. In a subsequent functional magnetic resonance imaging experiment with similar task and stimuli, we found anticipatory activations in the occipital cortex contralateral to the expected distractor, but no additional target modulation, when participants were given advance information about a distractor's subsequent presence and location. Several attention-related control structures (frontal eye fields and superior parietal cortex) were active during attentional preparation for all trials, whereas the left superior prefrontal and right angular gyri were additionally activated when a distractor was anticipated. The right temporoparietal junction showed stronger functional coupling with occipital regions during preparation for trials with an isolated target than for trials with a distractor expected. These results show that anticipation of a visual distractor at a known location, remote from the target, can lead to (1) a reduction in the behavioral cost of that distractor, (2) preparatory modulation of the occipital cortex contralateral to the location of the expected distractor, and (3) anticipatory activation of distinct parietal and frontal brain structures. These findings indicate that specific components of preparatory visual attention may be devoted to minimizing the impact of distractors, not just to enhancements of target processing.

Attention can enhance processing for relevant information and suppress this for ignored stimuli. However, some residual processing may still arise without attention. Here we presented overlapping outline objects at study, with subjects attending to those in one color but not the other. Attended objects were subsequently recognized on a surprise memory test, whereas there was complete amnesia for ignored items on such direct explicit testing; yet reliable behavioral priming effects were found on indirect testing. Event-related fMRI examined neural responses to previously attended or ignored objects, now shown alone in the same or mirror-reversed orientation as before, intermixed with new items. Repetition-related decreases in fMRI responses for objects previously attended and repeated in the same orientation were found in the right posterior fusiform, lateral occipital, and left inferior frontal cortex. More anterior fusiform regions also showed some repetition decreases for ignored objects, irrespective of orientation. View-specific repetition decreases were found in the striate cortex, particularly for previously attended items. In addition, previously ignored objects produced some fMRI response increases in the bilateral lingual gyri, relative to new objects. Selective attention at exposure can thus produce several distinct long-term effects on processing of stimuli repeated later, with neural response suppression stronger for previously attended objects, and some response enhancement for previously ignored objects, with these effects arising in different brain areas. Although repetition decreases may relate to positive priming phenomena, the repetition increases for ignored objects shown here for the first time might relate to processes that can produce “negative priming” in some behavioral studies. These results reveal quantitative and qualitative differences between neural substrates of long-term repetition effects for attended versus unattended objects.

We examined priming of visual search by repeated target location or color in two patients with left visual neglect and extinction, following strokes centered on the right inferior parietal lobe. Both patients, like the healthy controls we tested, showed intact priming, with performance speeded when either the location or color of a singleton target was repeated over successive trials in a standard search condition (Experiment 1). This was observed both from and to targets on the contralesional (left) side. Moreover, priming of search was still observed even when a return of fixation back to display-center was required between successive trials (Experiment 2). When briefer displays were used (Experiment 3), the patients often failed to detect left targets. This situation revealed an important dissociation: Whereas location priming only arose from preceding left targets that had been consciously detected, color priming (possibly arising within the intact ventral stream) did not depend on awareness of the preceding target. There was considerable color priming from missed targets. These findings demonstrate relatively intact priming of visual search by color and location in patients with right parietal damage, and also reveal that location priming may differ from color priming in requiring awareness.

The ability to inhibit saccades toward suddenly appearing peripheral stimuli (prosaccades) and direct them to contralateral locations instead (antisaccades) is a crucial marker of eye movement control. Typically developing infants as young as 4-month-olds can learn to inhibit reflexive saccades to peripheral stimuli, but they do not produce antisaccades, whose development later in infancy and its underlying neural computations remain unexplored. Here we tested oculomotor control in typically developing toddlers and toddlers with fragile X syndrome (FXS), a disorder of known genetic origin that allows the investigation of the neuro-computational properties contributing to the development of saccadic control. Typically developing toddlers decreased looking toward peripheral cues that predicted contralateral rewards, whose appearance they anticipated. Furthermore, this correlated with age, indicating a gradual development of saccadic control. In contrast with the typical case, toddlers with FXS did not decrease their looks to peripheral onsets that predicted contralateral events. Importantly, the atypical pattern of performance was also evident in the elimination of the correlation with mental or chronological age found in controls. Taken together, the findings suggest that control of saccades and its developmental trajectory is atypical in toddlers with FXS, consistent with inhibitory deficits previously shown at later ages in this condition. Potential implications for the neural mechanisms underlying the typical and atypical development of oculomotor control are discussed.

Right-hemisphere patients with left neglect often demonstrate abnormal visual search, re-examining stimuli to the right while ignoring those to the left. But re-fixations alone do not reveal if patients misjudge whether they have searched a location before. Here, we not only tracked the eye movements of 16 neglect patients during search, but also asked them to click a response button only when they judged they were fixating a target for the very first time. “Re-clicking” on previously found targets would indicate that patients erroneously respond to these as new discoveries. Lesions were mapped with high-resolution MRI. Neglect patients with damage involving the right intraparietal sulcus or right inferior frontal lobe “re-clicked” on previously found targets on the right at a pathological rate, whereas those with medial occipito-temporal lesions did not. For the intraparietal sulcus patients, the probability of erroneous re-clicks on an old target increased with time since first discovering it; whereas for frontal patients it was independent of search time, suggesting different underlying mechanisms in these two types of patient. Re-click deficits correlated with degree of leftward neglect, mainly due to both being severe in intraparietal cases. These results demonstrate that misjudging previously searched locations for new ones can contribute to pathological search in neglect, with potentially different mechanisms being involved in intraparietal versus inferior frontal patients. When combined with a spatial bias to the right, such deficits might explain why many neglect patients often re-examine rightward locations, at the expense of items to their left.

Previous ERP studies have uncovered cross-modal interactions in endogenous spatial attention. Directing attention to one side to judge stimuli from one particular modality can modulate early modality-specific ERP components not only for that modality, but also for other currently irrelevant modalities. However, past studies could not determine whether the spatial focus of attention in the task-irrelevant secondary modality was similar to the primary modality, or was instead diffuse across one hemifield. Here, auditory or visual stimuli could appear at any one of four locations (two on each side). In different blocks, subjects judged stimuli at only one of these four locations, for an auditory (Experiment 1) or visual (Experiment 2) task. Early attentional modulations of visual and auditory ERPs were found for stimuli at the currently relevant location, compared with those at the irrelevant location within the same hemifield, thus demonstrating within-hemifield tuning of spatial attention. Crucially, this was found not only for the currently relevant modality, but also for the currently irrelevant modality. Moreover, these within-hemifield attention effects were statistically equivalent regardless of the task relevance of the modality, for both the auditory and visual ERP data. These results demonstrate that within-hemifield spatial attention for one task-relevant modality can transfer cross-modally to a task-irrelevant modality, consistent with spatial selection at a multimodal level of representation.

Visual extinction after right parietal damage involves a loss of awareness for stimuli in the contralesional field when presented concurrently with ipsilesional stimuli, although contralesional stimuli are still perceived if presented alone. However, extinguished stimuli can still receive some residual on-line processing, without awareness. Here we examined whether such residual processing of extinguished stimuli can produce implicit and/or explicit memory traces lasting many minutes. We tested four patients with right parietal damage and left extinction on two sessions, each including distinct study and subsequent test phases. At study, pictures of objects were shown briefly in the right, left, or both fields. Patients were asked to name them without memory instructions (Session 1) or to make an indoor/outdoor categorization and memorize them (Session 2). They extinguished most left stimuli on bilateral presentation. During the test (up to 48 min later), fragmented pictures of the previously exposed objects (or novel objects) were presented alone in either field. Patients had to identify each object and then judge whether it had previously been exposed. Identification of fragmented pictures was better for previously exposed objects that had been consciously seen and critically also for objects that had been extinguished (as compared with novel objects), with no influence of the depth of processing during study. By contrast, explicit recollection occurred only for stimuli that were consciously seen at study and increased with depth of processing. These results suggest implicit but not explicit memory for extinguished visual stimuli in parietal patients.

Recent behavioral and event-related brain potential (ERP) studies have revealed cross-modal interactions in endogenous spatial attention between vision and audition, plus vision and touch. The present ERP study investigated whether these interactions reflect supramodal attentional control mechanisms, and whether similar cross-modal interactions also exist between audition and touch. Participants directed attention to the side indicated by a cue to detect infrequent auditory or tactile targets at the cued side. The relevant modality (audition or touch) was blocked. Attentional control processes were reflected in systematic ERP modulations elicited during cued shifts of attention. An anterior negativity contralateral to the cued side was followed by a contralateral positivity at posterior sites. These effects were similar whether the cue signaled which side was relevant for audition or for touch. They also resembled previously observed ERP modulations for shifts of visual attention, thus implicating supramodal mechanisms in the control of spatial attention. Following each cue, single auditory, tactile, or visual stimuli were presented at the cued or uncued side. Although stimuli in task-irrelevant modalities could be completely ignored, visual and auditory ERPs were nevertheless affected by spatial attention when touch was relevant, revealing cross-modal interactions. When audition was relevant, visual ERPs, but not tactile ERPs, were affected by spatial attention, indicating that touch can be decoupled from cross-modal attention when task-irrelevant.

We examined the effects of chronic unilateral lesions to either the inferior parietal lobe, or to the dorsolateral prefrontal cortex including the frontal eye fields (FEFs), upon human visual perception and saccades in temporal-order-judgment (TOJ) tasks. Two visual events were presented on each trial, one in each hemifield at various stimulus onset asynchronies (SOAs). In the saccade task, patients moved their eyes to whichever stimulus attracted gaze first. In the perceptual-manual task, they pressed a button to indicate which stimulus was perceived first. Frontal patients showed appropriate TOJs for visual targets in both tasks. Parietal patients showed appropriate TOJs in the perceptual-manual but not the saccade task; their saccades tended to be ipsilesional unless the contralesional target led substantially. This reveals a bias in saccade choice after parietal damage that cannot be attributed to deficient visual perception. These results challenge previous claims that only anterior lesions produce motoric spatial biases in humans. However, they are in accord with recent neurophysiological evidence for parietal involvement in saccade generation, and also with suggestions that visuomotor transformations in the parietal lobe serving direct spatial motor responses can dissociate from conscious perception as indicated by indirect arbitrary responses.

Tactile-visual links in spatial attention were examined by presenting spatially nonpredictive tactile cues to the left or right hand, shortly prior to visual targets in the left or right hemifield. To examine the spatial coordinates of any cross-modal links, different postures were examined. The hands were either uncrossed, or crossed so that the left hand lay in the right visual field and vice versa. Visual judgments were better on the side where the stimulated hand lay, though this effect was somewhat smaller with longer intervals between cue and target, and with crossed hands. Event-related brain potentials (ERPs) showed a similar pattern. Larger amplitude occipital N1 components were obtained for visual events on the same side as the preceding tactile cue, at ipsilateral electrode sites. Negativities in the Nd2 interval at midline and lateral central sites, and in the Nd1 interval at electrode Pz, were also enhanced for the cued side. As in the psychophysical results, ERP cueing effects during the crossed posture were determined by the side of space in which the stimulated hand lay, not by the anatomical side of the initial hemispheric projection for the tactile cue. These results demonstrate that crossmodal links in spatial attention can influence sensory brain responses as early as the N1, and that these links operate in a spatial frame-of-reference that can remap between the modalities across changes in posture.

Five patients with visual extinction following unilateral brain injury were briefly presented with colored letters in either or both visual fields, and required to report and locate the colors or the shapes. On double simultaneous stimulation, they tended to miss the event contralateral to their lesion. This extinction was increased when the two stimuli were the same on the reported dimension, Similarity on the irrelevant dimension had no effect. These data suggest that extinguished colors and shapes may be correctly extracted by the visual system (when task-relaant) even though they are unavailable for verbal report. An analogy is made with the phenomena of “repetition blindness” in normal observers, and it is proposed that extinction may reflect failure in a token-individuation process for correctly extracted visual types.