Attentional shifts induced by uninformative number symbols modulate neural activity in human occipital cortex

Number processing interacts with space encoding in a wide variety of experimental paradigms. Most intriguingly, the passive viewing of uninformative number symbols can shift visuo-spatial attention to different target locations according to the number magnitude, i.e., small/large numbers facilitate processing of left/right targets, respectively. The brain architecture dedicated to these attention shifts associated with numbers remains unknown. Evoked potential recordings indicate that both early and late stages are involved in this spatio-numerical interaction, but the neuro-functional anatomy needs to be specified. Here we use, for the first time, functional magnetic resonance imaging (fMRI) to investigate attentional orienting following uninformative Arabic digits. We show that BOLD response in occipital visual regions is modulated by the congruency between digit magnitude (small/large) and target side (left/right). Additionally, we report higher BOLD responses following large (8, 9) compared to small (1, 2) digits in two bilateral parietal regions, yielding a significant effect of digit magnitude. We propose and discuss the view that encoding of semantic representations related to number symbols in parietal cortex leads to shifts in visuo-spatial attention and enhances visual processing in the occipital cortex according to number-space congruency rules.     

Attentional load attenuates synaesthetic priming effects in grapheme-colour synaesthesia

One of the hallmarks of grapheme-colour synaesthesia is that colours induced by letters, digits and words tend to interfere with the identification of coloured targets when the two colours are different, i.e., when they are incongruent. In a previous investigation (Mattingley et al., 2001) we found that this synaesthetic congruency effect occurs when an achromatic-letter prime precedes a coloured target, but that the effect disappears when the letter is pattern masked to prevent conscious recognition of its identity. Here we investigated whether selective attention modulates the synaesthetic congruency effect in a letter-priming task. Fourteen grapheme-colour synaesthetes and 14 matched, non-synaesthetic controls participated. The amount of selective attention available to process the letter-prime was limited by having participants perform a secondary visual task that involved discriminating pairs of gaps in adjacent limbs of a diamond surrounding the prime. In separate blocks of trials the attentional load of the secondary task was systematically varied to yield 'low load' and 'high load' conditions. We found a significant congruency effect for synaesthetes, but not for controls, when they performed a secondary attention-demanding task during presentation of the letter prime. Crucially, however, the magnitude of this priming was significantly reduced under conditions of high-load relative to low-load, indicating that attention plays an important role in modulating synaesthesia. Our findings help to explain the observation that synaesthetic colour experiences are often weak or absent during attention-demanding tasks.     

The syndrome of visual-verbal disjunction following transection of the splenium of the corpus callosum. Difficulty in verbalizing visual information in the non-dominant hemisphere

According to the methods proposed by Gazzaniga et al. (1965, 1967), various visual-language associative functions were examined in 3 patients who had undergone the transection of the splenium of the corpus callosum for a pineal operation.Case 1 presented partial disturbance in reading aloud and also in colour naming in the left visual field with intact object naming capacity. Case 2 presented disturbances in reading aloud and in colour naming as well as partial object naming difficulty in the left visual field. Similar tests for Case 3 revealed disturbances in reading aloud and mild colour naming difficulty in the left visual field. His colour naming disturbance could be observed only by bilateral colour presentation tests, and the results of the unilateral colour presentation tests were normal.Visual matching tests revealed that the perception of, or the recognition of visual information in the left visual field were well preserved in spite of the inability to describe the information verbally.Present investigations did not support the view of Gazzaniga et al. (1967, 1971) that the right hemisphere can understand the meaning of written words.In the colour naming tests of bilateral simultaneous presentation, visual extinction-like phenomena were observed in the left visual field. Visual matching tests by bilateral simultaneous colour presentation revealed, however, that colour perception or recognition is perfectly preserved in both visual fields in spite of the apparent colour neglect in the left visual field.     

Attention and sensory gain control: a peripheral visual process?

Attention-related sensory gain control in human extrastriate cortex is believed to improve the acuity of visual perception. Yet given wide variance in the spatial resolution of vision across the retina, it remains unclear whether sensory gain operates homogenously between foveal and nonfoveal retinotopic locations. To address this issue, we used event-related potentials (ERPs) in a variant of the canonical spatial attention task. Participants were cued to expect targets at either fixation (foveal targets) or at a location several degrees above fixation (parafoveal targets). At both target locations, manual reaction times were shorter for cued relative to uncued targets, indicating that attention was consistently oriented to the cued location. Nevertheless, attention-related increases in sensory-evoked cortical activity were only observed at the parafoveal target location, as measured by the amplitude of the lateral occipital P1 ERP component. A second experiment replicated this data pattern using targets with lower stimulus contrast, indicating that the absence of a P1 effect for foveal targets could not be attributed to a saturated P1 response under higher-contrast stimulus conditions. When considered in light of retinogeniculate projections to cortex showing systematic changes in their physiological organization beginning within a degree of visual angle of the fovea, our findings support the proposal that the strategic functions of visual attention may vary with the retinotopic location involved.     

Shifting visual attention in space: an electrophysiological analysis using high spatial resolution mapping.

OBJECTIVES: Evidence from cortical electrophysiology and functional imaging converges on the view that visual spatial selective attention results in a facilitation of early sensory processing in visual cortical structures. Little is known, however, about the neural control processes that lead to this facilitation. The present study was aimed at further investigating these control processes and their neural correlates by analyzing high spatial resolution maps of brain activity that were evoked by attention-directing cues, but occurred prior to presentation of the target stimulus. METHODS: Subjects (n=14) were presented with central arrow cues that instructed them to attend covertly to either a left or right field location in order to compare two subsequent target stimuli simultaneously presented to the location. On half of the trials, targets were presented to the cued location, while in the other half, targets were presented to the opposite visual field location. Subjects had to respond via button press on 16% of the trials when target stimuli were identical. Event-related potentials (ERPs) were recorded from 92 scalp electrodes which allowed a sufficiently finegrained analysis of the regional specificity of the ERP components. RESULTS: In response to the cues, an initial component over occipital-parietal electrode sites was consistent with an early involvement of the posterior-parietal cortex, perhaps in the initial step of attentional orienting. A second component over the lateral-prefrontal cortex is consistent with the voluntary control and maintenance of attention, a function known to be subserved by frontal cortical structures. A late component narrowly focussed over occipital-temporal electrode sites is most plausibly related to activation of parts of the ventral extrastriate cortex. CONCLUSIONS: The data support the current view that voluntarily orienting visual attention in space leads to top-down modulations in cortical excitability of ventral extrastriate regions initiated by posterior-parietal and mediated by lateral-prefrontal cortical structures.     

The "zoom lens" of focal attention in visual search: changes in aging and Alzheimer's disease

Visual search for a target in an array of distractors relies upon flexible shifts between global and local modes of attentional processing. Visual search is slowed in patients with Alzheimer's disease (AD), in part due to an increase in the number and duration of eye fixations made en route to a target (R?sler et al., 2000). This phenomenon may represent a compensatory adaptation to a narrowing of the zone of focal attention, necessitating more shifts of gaze in order to attend to the global workspace. Eye fixations were analyzed in two regions of interest (ROIs, central fixation and peripheral target locations) in 9 patients with mild AD, 9 cognitively intact age-matched control subjects, and 9 young controls, while they searched for a target object in a radial array that contained from 1 to 6 stimuli. Contrasted with young subjects, the search strategy of older controls and, to a greater extent, AD patients showed an increase in the average number and duration of peripheral fixations. Reduced efficiency of visual search in AD may be contributed to by reduced ability to dynamically adjust the attentional zoom, coupled with the inability to disengage attention from peripheral targets.     

Processing of multisensory spatial congruency can be dissociated from working memory and visuo-spatial attention

Tactile stimuli at the same location as a visual target can increase activity in the contralateral occipital cortex compared with spatially incongruent bimodal stimulation. Does this cross‐modal congruency effect in the visual cortex depend on available cognitive resources? Visual attention and working memory can modulate responses to visual stimuli in the occipital cortex, with attenuation in conditions of high‐attentional or low‐memory load. Here, we asked if these two cognitive factors affect multisensory processing in the visual cortex. Using functional magnetic resonance imaging in humans, we manipulated the load of working memory or visuo‐spatial attention. Concurrently with these primary tasks we presented same‐side (congruent) or opposite‐side (incongruent) visuo‐tactile stimuli. Both experiments revealed enhancement of brain activity for congruent vs incongruent bimodal stimulation in the contralateral occipital cortex. Critically, these cross‐modal effects occurred irrespective of the level of load. We conclude that processing of visuo‐tactile spatial congruency in the visual cortex does not depend on available visuo‐spatial and memory resources.     

Split of spatial attention as predicted by a systems-level model of visual attention

Can we attend to multiple distinct spatial locations at the same time? According to a recent psychophysical study [J. Dubois et?al. (2009)Journal of Vision, 9, 3.1-11] such a split of spatial attention might be limited to short periods of time. Following N. P. Bichot et?al. [(1999)Perception & Psychophysics, 61, 403-423] subjects had to report the identity of multiple letters that were briefly presented at different locations, while two of these locations (targets) were relevant for a concurrent shape comparison task. In addition to the design used by Bichot et?al. stimulus onset asynchrony between shape onset and letters was systematically varied. In general, the performance of subjects was superior at target locations. Furthermore, for short stimulus onset asynchronies, performance was simultaneously increasing at both target locations. For longer stimulus onset asynchronies, however, performance deteriorated at one of the target locations while increasing at the other target location. It was hypothesized that this dynamic deployment of attention might be caused by competitive processes in saccade-related structures such as the frontal eye field. Here we simulated the task of Dubois et?al. using a systems-level model of attention. Our results are consistent with recent findings in the frontal eye field obtained during covert visual search, and they support the view of a transient deployment of spatial attention to multiple stimuli in the early epoch of target selection.     

Objections against the view of visual extinction as an attentional disengagement deficit: the interaction between spatial position and temporal modulation

Visual extinction is a common consequence of brain injury where individuals fail to detect a contralesional target when it is presented with a competing ipsilesional target. This disorder is often seen as either a consequence of biased competitive interactions or as a consequence of an attentional disengagement deficit. A study of neurological patients by Rorden et al. (2008) found that extinction is maximal at stimulus simultaneity when the target stimuli straddle the central gaze fixation location, while it is maximal when the ipsilesional stimulus has a temporal lead on the contralesional stimulus when both target stimuli are presented in the ipsilesional visual field. The authors argued that these results are most parsimoniously explained as a consequence of low-level biased competitive interactions due to cortical magnification, but acknowledged that an attentional disengagement deficit could also accommodate the results. The present study set out to adjudicate between these models by examining the performance of neurologically healthy subjects who exhibit normal cortical magnification but do not have pathological attentional biases. We presented (a)synchronous double stimulation trials where both targets to be identified could straddle the central gaze fixation location or be presented in either the left or the right visual field. We found that in both the left and the right visual field target performance accuracy was poorest when the more peripheral stimulus led. This suggests that the findings from Rorden et al. can indeed be explained by a low-level physiological bias due to cortical magnification and argues against the notion that neurological extinction represents an attentional disengagement deficit.     

Fearful,surprised, happy,and angry facial expressions modulate gaze-oriented attention: Behavioral and ERP evidence

The impact of emotions on gaze-oriented attention was investigated in non-anxious participants. A neutral face cue with straight gaze was presented, which then averted its gaze to the side while remaining neutral or expressing an emotion (fear/surprise in Exp.1 and anger/happiness in Exp.2). Localization of a subsequent target was faster at the gazed-at location (congruent condition) than at the non-gazed-at location (incongruent condition). This Gaze-Orienting Effect (GOE) was enhanced for fear, surprise, and anger, compared to neutral expressions which did not differ from happy expressions. In addition, Event Related Potentials (ERPs) to the target showed a congruency effect on P1 for fear and surprise and a left lateralized congruency effect on P1 for happy faces, suggesting that target visual processing was also influenced by attention to gaze and emotions. Finally, at cue presentation, early postero-lateral (Early Directing Attention Negativity (EDAN)) and later antero-lateral (Anterior Directing Attention Negativity (ADAN)) attention-related ERP components were observed, reflecting, respectively, the shift of attention and its holding at gazed-at locations. These two components were not modulated by emotions. Together, these findings show that the processing of social signals such as gaze and facial expression interact rather late and in a complex manner to modulate spatial attention.