Individual differences in visual field shape modulate the effects of attention on the lower visual field advantage in crowding.

It has previously been reported that visual crowding of a target by flankers is stronger in the upper visual field than in the lower, and this finding has been attributed to greater attentional resolution in the lower hemifield (He, Cavanagh, & Intriligator, 1996). Here we show that the upper/lower asymmetry in visual crowding can be explained by natural variations in the borders of each individual's visual field. Specifically, asymmetry in crowding along the vertical meridian can be almost entirely accounted for by replacing the conventional definition of visual field location, in units of degrees of visual angle, with a definition based on the ratio of the extents of an individual's upper and lower visual field. We also show that the upper/lower crowding asymmetry is eliminated when stimulus eccentricity is expressed in units of percentage of visual field extent but is present when the conventional measure of visual angle is used. We further demonstrate that the relationship between visual field extent and perceptual asymmetry is most evident when participants are able to focus their attention on the target location. These results reveal important influences of visual field boundaries on visual perception, even for visual field locations far from those boundaries.

White matter microstructure throughout the brain correlates with visual imagery in grapheme-color synesthesia.

In this study we show, for the first time, a correlation between the neuroanatomy of the synesthetic brain and a metric that measures behavior not exclusive to the synesthetic experience. Grapheme-color synesthetes (n=20), who experience colors triggered by viewing or thinking of specific letters or numbers, showed altered white matter microstructure, as measured using diffusion tensor imaging, compared with carefully matched non-synesthetic controls. Synesthetes had lower fractional anisotropy and higher perpendicular diffusivity when compared to non-synesthetic controls. An analysis of the mode of anisotropy suggested that these differences were likely due to the presence of more crossing pathways in the brains of synesthetes. Additionally, these differences in white matter microstructure correlated negatively, and only for synesthetes, with a measure of the vividness of their visual imagery. Synesthetes who reported the most vivid visual imagery had the lowest fractional anisotropy and highest perpendicular diffusivity. We conclude that synesthetes as a population vary along a continuum while showing categorical differences in neuroanatomy and behavior compared to non-synesthetes.

Exploring the edges of visual space: the influence of visual boundaries on peripheral localization.

Previous studies of localization of stationary targets in the peripheral visual field have found either underestimations (foveal biases) or overestimations (peripheral biases) of target eccentricity. In the present study, we help resolve this inconsistency by demonstrating the influence of visual boundaries on the type of localization bias. Using a Goldmann perimeter (an illuminated half-dome), we presented targets at different eccentricities across the visual field and asked participants to judge the target locations. In Experiments 1 and 2, participants reported target locations relative to their perceived visual field extent using either a manual or verbal response, with both response types producing a peripheral bias. This peripheral localization bias was a non-linear scaling of perceived location when the visual field was not bounded by external borders induced by facial features (i.e., the nose and brow), but location scaling was linear when visual boundaries were present. Experiment 3 added an external border (an aperture edge placed in the Goldmann perimeter) that resulted in a foveal bias and linear scaling. Our results show that boundaries that define a spatial region within the visual field determine both the direction of bias in localization errors for stationary objects and the scaling function of perceived location across visual space.

Attentional selection of relative SF mediates global versus local processing: evidence from EEG.

Previous research on functional hemispheric differences in visual processing has associated global perception with low spatial frequency (LSF) processing biases of the right hemisphere (RH) and local perception with high spatial frequency (HSF) processing biases of the left hemisphere (LH). The Double Filtering by Frequency (DFF) theory expanded this hypothesis by proposing that visual attention selects and is directed to relatively LSFs by the RH and relatively HSFs by the LH, suggesting a direct causal relationship between SF selection and global versus local perception. We tested this idea in the current experiment by comparing activity in the EEG recorded at posterior right and posterior left hemisphere sites while participants' attention was directed to global or local levels of processing after selection of relatively LSFs versus HSFs in a previous stimulus. Hemispheric asymmetry in the alpha band (8-12 Hz) during preparation for global versus local processing was modulated by the selected SF. In contrast, preparatory activity associated with selection of SF was not modulated by the previously attended level (global/local). These results support the DFF theory that top-down attentional selection of SF mediates global and local processing.

Local or global? Attentional selection of spatial frequencies binds shapes to hierarchical levels.

Contrary to the traditional view that shapes and their hierarchical level (local or global) are a priori integrated in perception, recent evidence suggests that the identity of a shape and its level are encoded independently, implying the need for shape-level binding to account for normal perception. What is the binding mechanism in this case? Using hierarchically arranged letter shapes, we obtained evidence that the left hemisphere has a preference for binding shapes to the local level, whereas the right hemisphere has a preference for binding shapes to the global level. More important, binding is modulated by attentional selection of higher or lower spatial frequencies. Attention to higher spatial frequencies facilitated subsequent binding by the left hemisphere of elements to the local level, whereas attention to lower spatial frequencies facilitated subsequent binding by the right hemisphere of elements to the global level.

Implicit representation and explicit detection of features in patients with hemispatial neglect.

Despite profound inattention to the side of space opposite a brain lesion in patients with unilateral neglect, priming studies demonstrate that undetected stimuli are capable of influencing subsequent behaviour. However, the nature of implicit processing of neglected stimuli is poorly understood. In the current study, we examined implicit processing in five patients with neglect using both visual search and priming methods. A psychophysical staircase method varying time of presentation was first used to establish a high (75%) and low (25%) detection probability for targets in both a feature and a conjunction search array. The arrays were then used in a priming task to examine how a difference in the level of overt detection of a feature or a conjunction presented in neglected space influenced subsequent discrimination speed to a single probe presented at fixation. The results showed that priming effects with feature primes were independent of their explicit detection rates (high versus low), but priming effects with conjunction primes reflected the pattern of explicit detection. These findings are discussed as they relate to availability versus accessibility of neglected stimuli.

Where does attention go when it moves? Spatial properties and locus of the attentional repulsion effect.

Reliable effects of spatial attention on perceptual measures have been well documented, yet little is known about how attention affects perception of space per se. The present study examined the effects of involuntary shifts of spatial attention on perceived location using a paradigm developed by S. Suzuki and P. Cavanagh (1997) that produces an attentional repulsion effect (ARE). The ARE refers to the illusory displacement of two vernier lines away from briefly presented cues. In Experiment 1, we show that the magnitude of the ARE depends on cue-target distance, indicating that the effects of attention on perceived location are not uniform across the visual field. Experiments 2 and 3 tested whether repulsion occurs away from cue center of mass or from cue contour. Perceived repulsion always occurred away from the cues' center of mass, regardless of the arrangement of the cue contours relative to the vernier lines. Moreover, the magnitude of the ARE varied with shifts in the position of the cues' center of mass. These experiments suggest that the onset of the cue produces a shift of attention to its center of mass rather than to the salient luminance contours that define it, and that this mechanism underlies the ARE.

The attentional repulsion effect and relative size judgments.

Rapid shifts of involuntary attention have been shown to induce mislocalizations of nearby objects. One pattern of mislocalization, termed the Attentional Repulsion Effect (ARE), occurs when the onset of peripheral pre-cues lead to perceived shifts of subsequently presented stimuli away from the cued location. While the standard ARE configuration utilizes vernier lines, to date, all previous ARE studies have only assessed distortions along one direction and tested one spatial dimension (i.e., position or shape). The present study assessed the magnitude of the ARE using a novel stimulus configuration. Across three experiments participants judged which of two rectangles on the left or right side of the display appeared wider or taller. Pre-cues were used in Experiments 1 and 2. Results show equivalent perceived expansions in the width and height of the pre-cued rectangle in addition to baseline asymmetries in left/right relative size under no-cue conditions. Altering cue locations led to shifts in the perceived location of the same rectangles, demonstrating distortions in perceived shape and location using the same stimuli and cues. Experiment 3 demonstrates that rectangles are perceived as larger in the periphery compared to fixation, suggesting that eye movements cannot account for results from Experiments 1 and 2. The results support the hypothesis that the ARE reflects a localized, symmetrical warping of visual space that impacts multiple aspects of spatial and object perception.

Different effects of voluntary and involuntary attention on EEG activity in the gamma band.

Previous studies have shown that EEG activity in the gamma range can be modulated by attention. Here, we compared this activity for voluntary and involuntary spatial attention in a spatial-cueing paradigm with faces as targets. The stimuli and trial timing were kept constant across attention conditions with only the predictive value of the cue changing. Gamma-band response was linked to voluntary shifts of attention, but not to the involuntary capture of attention. The presence of increased gamma responses for the voluntary allocation of attention, and its absence in cases of involuntary capture suggests that the neural mechanisms governing these two types of attention are different. Moreover, these data allow a description of the temporal dynamics contributing to the dissociation between voluntary and involuntary attention. The distribution of this correlate of voluntary attention is consistent with a top-down process involving contralateral anterior and posterior regions.

Using spatial frequency scales for processing face features and face configuration: an ERP analysis.

In the present study we examined the influence of spatial filtering on the N170-effect, a relatively early face-selective ERP difference associated with face detection. We compared modulation of the N170-effect using spatially filtered stimuli that either facilitated feature analysis or impeded configural analysis. The salience of inner face components was enhanced by presenting them in isolation. Configural processing was manipulated by face inversion. The N170-effects elicited by upright faces and isolated inner components were similar across low- and high-spatial frequency scales. In contrast, the inversion effect (enhanced N170 amplitude for inverted compared with upright faces) was only observed with broadband and low-spatial frequency stimuli. These findings demonstrate that the N170-effect can be influenced by both low- and high-spatial frequency channels. Moreover, they indicate that different configural manipulations (isolated features vs. face inversion) affect face detection in distinct ways, consistent with separate processing mechanisms for different types of configural encoding.

Visual hemispatial neglect, re-assessed.

Increased computer use in clinical settings offers an opportunity to develop new neuropsychological tests that exploit the control computers have over stimulus dimensions and timing. However, before adopting new tools, empirical validation is necessary. In the current study, our aims were twofold: to describe a computerized adaptive procedure with broad potential for neuropsychological investigations, and to demonstrate its implementation in testing for visual hemispatial neglect. Visual search results from adaptive psychophysical procedures are reported from 12 healthy individuals and 23 individuals with unilateral brain injury. Healthy individuals reveal spatially symmetric performance on adaptive search measures. In patients, psychophysical outcomes (as well as those from standard paper-and-pencil search tasks) reveal visual hemispatial neglect. Consistent with previous empirical studies of hemispatial neglect, lateralized impairments in adaptive conjunction search are greater than in adaptive feature search tasks. Furthermore, those with right hemisphere damage show greater lateralized deficits in conjunction search than do those with left hemisphere damage. We argue that adaptive tests, which automatically adjust to each individual's performance level, are efficient methods for both clinical evaluations and neuropsychological investigations and have the potential to detect subtle deficits even in chronic stages, when flagrant clinical signs have frequently resolved.

Attentional selection and the representation of holes and objects.

We examine whether holes (two separate cutout rectangles in a surface) appearing as if on a homogeneous background produce object-based effects similar to those observed when the same regions appear as separate items in front of that surface (commonly called objects). We used a version of the two-rectangle design described by R. Egly, J. Driver, and R. D. Rafal (1994). Viewing modified patterns through stereoscopic goggles created the perception of the rectangles as either part of the background or as foreground objects. In Experiment 1, we replicated Egly et al. when the regions were perceived as objects but not when they were perceived as holes. In Experiment 2, we included a condition where the background was split: The rectangles in the holes condition were perceived as part of two separate background regions. In this case, the object-based effects were the same as when the rectangles were foreground objects. The findings of Experiment 2 demonstrate that those of Experiment 1 were not due to depth per se, but rather to the background being treated as a single region. More importantly, these results demonstrate that identically shaped regions in the stimulus engage object-based attention differently, depending on how the regions are perceptually organized.

Implicit integration in a case of integrative visual agnosia.

We present a case (SE) with integrative visual agnosia following ischemic stroke affecting the right dorsal and the left ventral pathways of the visual system. Despite his inability to identify global hierarchical letters [Navon, D. (1977). Forest before trees: The precedence of global features in visual perception. Cognitive Psychology, 9, 353-383], and his dense object agnosia, SE showed normal global-to-local interference when responding to local letters in Navon hierarchical stimuli and significant picture-word identity priming in a semantic decision task for words. Since priming was absent if these features were scrambled, it stands to reason that these effects were not due to priming by distinctive features. The contrast between priming effects induced by coherent and scrambled stimuli is consistent with implicit but not explicit integration of features into a unified whole. We went on to show that possible/impossible object decisions were facilitated by words in a word-picture priming task, suggesting that prompts could activate perceptually integrated images in a backward fashion. We conclude that the absence of SE's ability to identify visual objects except through tedious serial construction reflects a deficit in accessing an integrated visual representation through bottom-up visual processing alone. However, top-down generated images can help activate these visual representations through semantic links.

A mismatch negativity study of local-global auditory processing.

We used mismatch negativity (MMN) to examine structural encoding of local and global auditory patterns in perceptual memory. Unlike previous MMN studies of local-global auditory perceptual organization that used interval-contour stimuli, here we presented hierarchical stimuli in which local pattern organization formed global patterns. Importantly, our stimuli allowed independent manipulation of the two structural levels. In separate blocks, participants were exposed to frequent local standard patterns and rare local deviant patterns, or to frequent global standard patterns and rare global deviant patterns. Within each deviant pattern, the variation from the standard pattern could occur at onset (early), towards the end of the pattern (late) or over both time windows (both). To isolate pattern indexing at one level, the other level continuously changed (e.g., in a global standard block, local elements varied trial-by-trial). MMN was found only for global deviant patterns, and only when deviation occurred late in the pattern. In a separate behavioral experiment, global deviants were detected more often than local ones, although initial similarity followed by a late deviation from the standard pattern was not required for explicit deviant detection (as with the MMN). This report demonstrates neural structural encoding for global information, when independently manipulated from local information. Furthermore, it extends previous MMN findings that have revealed indexing of complex abstract auditory information to the realm of hierarchical perceptual organization.

Inhibition of return and object-based attentional selection.

Visual attention research has revealed that attentional allocation can occur in space- and/or object-based coordinates. Using the direct and elegant design of R. Egly, J. Driver, and R. Rafal (1994), the present experiments tested whether space- and object-based inhibition of return (IOR) emerge under similar time courses. The experiments were capable of isolating both space- and object-based effects induced by peripheral and back-to-center cues. The results generally support the contention that spatially nonpredictive cues are effective in producing space-based IOR at a variety of stimulus onset asynchronies (SOAs) and under a variety of stimulus conditions. Whether facilitatory or inhibitory in direction, the object-based effects occurred over a very different time course than did the space-based effects. Reliable object-based IOR was only found under limited conditions and was tied to the time since the most recent cue (peripheral or central). The finding that object-based effects are generally determined by SOA from the most recent cue may help to resolve discrepancies in the IOR literature. These findings also have implications for the search facilitator role that IOR is purported to play in the guidance of visual attention.

Is filtering difficulty the basis of attentional deficits in schizophrenia?

The distractibility that schizophrenia patients display may be the result of a deficiency in filtering out irrelevant information. The aim of the current study was to assess whether patients with schizophrenia exhibit greater difficulty when task-irrelevant features change compared to healthy participants. Thirteen medicated outpatients with a diagnosis of schizophrenia and thirteen age- and parental education-matched controls performed a target selection task in which the task-relevant letter or the task-irrelevant features of color, and/or location repeated or switched. Participants were required to respond by pressing the appropriate key associated with the target letter. These patients with schizophrenia were slower when the task-relevant target letter switched than when it repeated. In contrast, schizophrenia patients performed similarly to controls when task-irrelevant information changed. Thus, we found no evidence that patients with schizophrenia were impaired in inhibiting irrelevant perceptual features. In contrast, changes in task-relevant features were problematic for patients relative to control participants. These results suggest that medicated outpatients who are mild to moderately symptomatic do not exhibit global impairments of feature processing. Instead, impairments are restricted to situations when task-relevant features vary. The current findings also suggest that when a course of action is not implied by an irrelevant feature, outpatients' behavior is not modulated by extraneous visual information any more than in healthy controls.

Changes in the distribution of sustained attention alter the perceived structure of visual space.

Visual spatial attention is a critical process that allows for the selection and enhanced processing of relevant objects and locations. While studies have shown attentional modulations of perceived location and the representation of distance information across multiple objects, there remains disagreement regarding what influence spatial attention has on the underlying structure of visual space. The present study utilized a method of magnitude estimation in which participants must judge the location of briefly presented targets within the boundaries of their individual visual fields in the absence of any other objects or boundaries. Spatial uncertainty of target locations was used to assess perceived locations across distributed and focused attention conditions without the use of external stimuli, such as visual cues. Across two experiments we tested locations along the cardinal and 45° oblique axes. We demonstrate that focusing attention within a region of space can expand the perceived size of visual space; even in cases where doing so makes performance less accurate. Moreover, the results of the present studies show that when fixation is actively maintained, focusing attention along a visual axis leads to an asymmetrical stretching of visual space that is predominantly focused across the central half of the visual field, consistent with an expansive gradient along the focus of voluntary attention. These results demonstrate that focusing sustained attention peripherally during active fixation leads to an asymmetrical expansion of visual space within the central visual field.

Spatial frequency selection and integration of global and local information in visual processing: A selective review and tribute to Shlomo Bentin.

Previous research has suggested a relationship between processing lower versus higher spatial frequencies (SFs) and global/local perception, respectively. Here we honor Shlomo Bentin by reviewing the work we conducted with him regarding this issue. This work was aimed at investigating the mechanisms by which selective attention to spatial frequency (SF) mediates global and local perception in general and how these perceptual levels are integrated with the shapes that define them. The experiments demonstrate that attention to global and local aspects of a hierarchical display biases the flexible selection of relatively lower and relatively higher SFs during image processing. Additionally, attentional selection of SF allows for the shapes in a hierarchical display to be integrated with the level (global/local) at which they occur. The studies reviewed here provide strong evidence that the flexible, top-down selection of low-level SF channels mediates the perception of global and local elements of visual displays. The studies also support a hemisphere asymmetry in this process, with right hemisphere functions biased toward global perception and left hemisphere functions biased toward local.

Crossing the midline: reducing attentional deficits via interhemispheric interactions.

Patients with unilateral neglect and extinction show a profound lack of awareness of stimuli presented contralateral to their lesion. However, many processes of perception are intact and contralesional stimuli seem to reach a high level of representation, perceptual and semantic. Some of these processes can work to decrease the magnitude of the attentional deficit. Here, we examine two of these intact processes, feature detection and perceptual grouping. First, we demonstrate that feature detection occurs in parallel in the contralesional visual fields of neglect and extinction patients. Second, we attempt to dissociate the influence of perceptual contours across the vertical meridian from the presence of an object or higher-level perceptual unit (or group) that may be created by these contours. We find that connections across the midline affect attentional deficits independently of the objects they may create. This suggests that several effects of grouping on neglect and extinction may be mediated by long-range cortical interactions that arise from connections across the vertical meridian.

Spatial distortions in localization and midline estimation in hemianopia and normal vision.

Studies have shown that individuals with hemianopia tend to bisect a line toward their blind, contralesional visual field, termed the hemianopic line bisection error (HLBE). One theory proposes that the HLBE is a perceptual distortion resulting from expansion of the central region of visual space. If true, perceptual expansions of the central regions in the intact hemifield should also be present and observable across different tasks. We tested this hypothesis using a peripheral localization task to assess localization and midpoint estimation along the horizontal axis of the visual field. In this task, participants judged the location of a target dot presented inside a Goldmann perimeter relative to their perceived visual field boundary. In Experiment 1, we tested neurologically healthy participants on the peripheral localization task as well as a novel midpoint assessment task in which participants reported their perceived midpoint along the horizontal axis of their left and right visual fields. The results revealed consistency in individual biases across the two tasks. We then used the peripheral localization task to test whether two patients with hemianopia showed a selective expansion of central visual space. For these patients, three axes were tested: the spared temporal horizontal axis and the upper and lower vertical axes. The results support the notion that the HLBE is due to expansion of perceived space along the spared temporal axis. Together, the results of both experiments validate the use of these novel paradigms for exploring perceptual asymmetries in both healthy individuals and patients with visual field loss.