Voluntary orienting is dissociated from target detection in human posterior parietal cortex

Human ability to attend to visual stimuli based on their spatial locations requires the parietal cortex. One hypothesis maintains that parietal cortex controls the voluntary orienting of attention toward a location of interest. Another hypothesis emphasizes its role in reorienting attention toward visual targets appearing at unattended locations. Here, using event-related functional magnetic resonance (ER-fMRI), we show that distinct parietal regions mediated these different attentional processes. Cortical activation occurred primarily in the intraparietal sulcus when a location was attended before visual-target presentation, but in the right temporoparietal junction when the target was detected, particularly at an unattended location.     

Lateralized irrelevant speech alters visuospatial selective attention mechanisms

Recent studies indicate that the coordination of spatial attention across modalities may in part be mediated by a supramodal attentional system. We try to extend the concept of a supramodal system and hypothesized that involuntary modulations of auditory attentional processes by irrelevant speech signals influence visuospatial attention, suggesting crossmodal links between vision and speech. In order to test this we recorded event-related brain potentials (ERPs) of 12 healthy subjects in a visuospatial selective attention task. The task to identify target stimuli appearing at lateral visual field locations caused the expected enhancements of the early P1 and N1 ERP components to attended visual stimuli. Understandable and ununderstandable task irrelevant speech was presented either at the visually attended position or in the opposite visual field location. Speech contralateral to unattended visual stimuli led to a decreased N1 amplitude. This effect was stronger for understandable speech. Thus, speech influences the allocation of visual spatial attention if it is presented in the unattended location. The results suggest crossmodal links of speech and visuospatial attention mechanisms at a very early stage of human perception.     

Push-pull mechanism of selective attention in human extrastriate cortex

Selective attention operates in visual cortex by facilitating processing of selected stimuli and by filtering out unwanted information from nearby distracters over circumscribed regions of visual space. The neural representation of unattended stimuli outside this focus of attention is less well understood. We studied the neural fate of unattended stimuli using functional magnetic resonance imaging by dissociating the activity evoked by attended (target) stimuli presented to the periphery of a visual hemifield and unattended (distracter) stimuli presented simultaneously to a corresponding location of the contralateral hemifield. Subjects covertly directed attention to a series of target stimuli and performed either a low or a high attentional-load search task on a stream of otherwise identical stimuli. With this task, target-search-related activity increased with increasing attentional load, whereas distracter-related activity decreased with increasing load in areas V4 and TEO but not in early areas V1 and V2. This finding presents evidence for a load-dependent push-pull mechanism of selective attention that operates over large portions of the visual field at intermediate processing stages. This mechanism appeared to be controlled by a distributed frontoparietal network of brain areas that reflected processes related to target selection during spatially directed attention.     

Mechanisms of feature- and space-based attention: response modulation and baseline increases

Selective attention modulates neural activity in the visual system both in the presence and in the absence of visual stimuli. When subjects direct attention to a particular location in a visual scene in anticipation of the stimulus onset, there is an increase in baseline activity. How do such baseline increases relate to the attentional modulation of stimulus-driven activity? Using functional magnetic resonance imaging, we demonstrate that baseline increases related to the expectation of motion or color stimuli at a peripheral target location do not predict the modulation of neural responses evoked by these stimuli when attended. In areas such as MT and TEO that were more effectively activated by one stimulus type than the other, attentional modulation of visually evoked activity depended on the stimulus preference of a visual area and was stronger for the effective than for the noneffective stimulus. In contrast, baseline increases did not reflect the stimulus preference of a visual area. Rather, these signals were shown to be spatially specific and appeared to be dominated by the location information and not by the feature information of the cue with the experimental paradigms under study. These findings provide evidence that baseline increases in visual cortex during cue periods do not reflect the activation of a memory template that includes particular stimulus properties of the expected target, but rather carry information about the location of an expected target stimulus. In addition, when the stimulus contained both color and motion, an object-based attention effect was observed, with significant attentional modulation in the area that responded preferentially to the unattended feature.     

Selective Attention in Auditory Processing as Reflected by Event-Related Brain Potentials

Measures of event-related brain potentials (ERPs) have revealed two kinds of selective-attention mechanisms that operate on attended and unattended auditory stimuli. The processing negativity of the ERP reveals a mechanism of intramodal selective attention in the auditory cortex controlled by the frontal cortex. This mechanism selects attended auditory stimuli for further processing when they differ from unattended stimuli in location or tonal frequency. Studies of intermodal selective attention have compared auditory ERPs during auditory and visual attention. At least in part different brain mechanisms may be involved in the selection of auditory stimuli among other auditory stimuli (intramodal selective attention) and in the selection of auditory stimuli among visual stimuli (intermodal selective attention). This is suggested by the results showing that the earlier component of the processing negativity, which is generated in the auditory cortex during intramodal selective attention, differs in scalp distribution from the early attention-related negativity elicited during intermodal selective attention. With respect to the unattended auditory stimuli, ERP studies of selective attention suggest that physical features of these stimuli are extensively processed. This is shown by the mismatch negativity component of the ERP, which is usually elicited by infrequent physical deviations in an auditory stimulus sequence both when this sequence is attended and when it is ignored. This would be impossible if the physical stimulus features were not extensively processed, even in the absence of attention.     

The time course of cortical facilitation during cued shifts of spatial attention

Adaptive behavior requires the rapid switching of attention among potentially relevant stimuli that appear in the environment. The present study used an electrophysiological approach to continuously measure the time course of visual pathway facilitation in human subjects as attention was shifted from one location to another. Steady-state visual evoked potentials (SSVEPs) were recorded to rapidly flickering lights at attended and unattended locations, and variations in SSVEP amplitude over time were calculated after a cue to shift attention. The build-up of cortical facilitation reflected in SSVEP amplitude was found to bear a close temporal relationship with the emergence of accurate target discriminations at the newly attended location.     

Effects of task difficulty and target likelihood in area V4 of macaque monkeys

Spatial attention improves performance at attended locations and correspondingly modulates firing rates of cortical neurons. The size of these behavioral and neuronal effects depends on the difficulty of the task performed at the attended location. Psychological theorists have attributed this to a tighter focus of a fixed amount of processing resource at the attended location, but the effects of task difficulty on the distribution of neuronal effects of attention across the visual field have not been fully explored. We trained rhesus monkeys to do a detection task in which difficulty and spatial attention were manipulated independently. Probe stimuli were used to measure behavioral performance in different conditions of attention and difficulty. Animals performed better at attended locations and this advantage increased with difficulty, consistent with data from human psychophysics. Neuronal modulation by spatial attention was larger with greater difficulty. In two animals, increasing difficulty caused a modest increase in neuronal responses to visual stimuli regardless of the locus of spatial attention. In a third animal, which was previously trained to ignore multiple distracting stimuli, increasing task difficulty increased responses at the focus of attention and suppressed responses away from the focus of attention. The results show that difficulty can modulate effects of spatial attention in V4; it can alter the distribution of sensory responses across the visual scene in ways that may depend on the subject's behavioral strategy.     

Dynamic shifts of visual receptive fields in cortical area MT by spatial attention

Voluntary attention is the top-down selection process that focuses cortical processing resources on the most relevant sensory information. Spatial attention--that is, selection based on stimulus position--alters neuronal responsiveness throughout primate visual cortex. It has been hypothesized that it also changes receptive field profiles by shifting their centers toward attended locations and by shrinking them around attended stimuli. Here we examined, at high resolution, receptive fields in cortical area MT of rhesus macaque monkeys when their attention was directed to different locations within and outside these receptive fields. We found a shift of receptive fields, even far from the current location of attention, accompanied by a small amount of shrinkage. Thus, already in early extrastriate cortex, receptive fields are not static entities but are highly modifiable, enabling the dynamic allocation of processing resources to attended locations and supporting enhanced perception within the focus of attention by effectively increasing the local cortical magnification.     

Sources of attention-sensitive visual event-related potentials

Summary In a study of the neural processes that mediate visual attention in humans, 32-channel recordings of event-related potentials were obtained from 14 normal subjects while they performed a spatial attention task. The generator locations of the early C1, P1, and Nl components of the visual evoked response were estimated by means of topographic maps of voltage and current source density in conjunction with dipole modelling. The topography of the C1 component (ca. 85 ms post-stimulus) was consistent with a generator in striate cortex, and this component was unaffected by attention. In contrast, the P1 and Nl components (ca. 95 and 170ms) exhibited current density foci at scalp sites overlying lateral extrastriate cortex and were larger for attended stimuli than for unattended stimuli. The voltage topographies in the 75–175 ms latency range were modeled with a 5-dipole configuration consisting of a single striate dipole and left-right pairs of dipoles located in lateral extrastriate and inferior occipito-temporal areas. This model was found to account for the voltage topographies produced by both attended and unattended stimuli with low residual variance. These results support the proposal that visual-spatial attention modulates neural activity in extrastriate visual cortex but does not affect the initial evoked response in striate cortex.This study was supported by ONR Contract N00014-89-J-1806, by grants from NIMH (MH-25594), NINCDS (NS 17778), the Human Frontier Science Program, DGICYT (PM92-0128), and by a Fulbright scholarship to the first author.     

Does focused endogenous attention prevent attentional capture in pop‐out visual search?

To investigate whether salient visual singletons capture attention when they appear outside the current endogenous attentional focus, we measured the N2pc component as a marker of attentional capture in a visual search task where target or nontarget singletons were presented at locations previously cued as task-relevant, or in the uncued irrelevant hemifield. In two experiments, targets were either defined by color or by a combination of color and shape. The N2pc was elicited both for attended singletons and for singletons on the uncued side, demonstrating that focused endogenous attention cannot prevent attentional capture by salient unattended visual events. However, N2pc amplitudes were larger for attended and unattended singletons that shared features with the current target, suggesting that top-down task sets modulate the capacity of visual singletons to capture attention both within and outside the current attentional focus.     

Neural correlates of sustained spatial attention in human early visual cortex

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

Predictive coding for motion stimuli in human early visual cortex

The current study investigates if early visual cortical areas, V1, V2 and V3, use predictive coding to process motion information. Previous studies have reported biased visual motion responses at locations where novel visual information was presented (i.e., the motion trailing edge), which is plausibly linked to the predictability of visual input. Using high-field functional magnetic resonance imaging (fMRI), we measured brain activation during predictable versus unpreceded motion-induced contrast changes during several motion stimuli. We found that unpreceded moving dots appearing at the trailing edge gave rise to enhanced BOLD responses, whereas predictable moving dots at the leading edge resulted in suppressed BOLD responses. Furthermore, we excluded biases in directional sensitivity, shifts in cortical stimulus representation, visuo-spatial attention and classical receptive field effects as viable alternative explanations. The results clearly indicate the presence of predictive coding mechanisms in early visual cortex for visual motion processing, underlying the construction of stable percepts out of highly dynamic visual input.     

ERP correlates of anticipatory attention: spatial and non-spatial specificity and relation to subsequent selective attention

Brain-based models of visual attention hypothesize that attention-related benefits afforded to imperative stimuli occur via enhancement of neural activity associated with relevant spatial and non-spatial features. When relevant information is available in advance of a stimulus, anticipatory deployment processes are likely to facilitate allocation of attention to stimulus properties prior to its arrival. The current study recorded EEG from humans during a centrally-cued covert attention task. Cues indicated relevance of left or right visual field locations for an upcoming motion or orientation discrimination. During a 1 s delay between cue and S2, multiple attention-related events occurred at frontal, parietal and occipital electrode sites. Differences in anticipatory activity associated with the non-spatial task properties were found late in the delay, while spatially-specific modulation of activity occurred during both early and late periods and continued during S2 processing. The magnitude of anticipatory activity preceding the S2 at frontal scalp sites (and not occipital) was predictive of the magnitude of subsequent selective attention effects on the S2 event-related potentials observed at occipital electrodes. Results support the existence of multiple anticipatory attention-related processes, some with differing specificity for spatial and non-spatial task properties, and the hypothesis that levels of activity in anterior areas are important for effective control of subsequent S2 selective attention.     

Preparatory activity in visual cortex indexes distractor suppression during covert spatial orienting

The deployment of spatial attention induces retinotopically specific increases in neural activity that occur even before a target stimulus is presented. Although this preparatory activity is thought to prime the attended regions, thereby improving perception and recognition, it is not yet clear whether this activity is a manifestation of signal enhancement at the attended locations or suppression of interference from distracting stimuli (or both). We investigated the functional role of these preparatory shifts by isolating a distractor suppression component of selection. Behavioral data have shown that manipulating the probability that visual distractors will appear modulates distractor suppression without concurrent changes in signal enhancement. In 2 experiments, functional magnetic resonance imaging revealed increased cue-evoked activity in retinotopically specific regions of visual cortex when increased distractor suppression was elicited by a high probability of distractors. This finding directly links cue-evoked preparatory activity in visual cortex with a distractor suppression component of visual selective attention.     

Inhibition of return in microsaccades

Inhibition of return (IOR) is the term used to describe the phenomenon whereby stimuli appearing at recently attended locations are reacted to less efficiently than stimuli appearing at locations that have not yet been attended. In the present study, we employed a typical IOR paradigm with peripheral uninformative cues while participants maintained their eyes at fixation. Eye position was monitored at a high sampling rate (500 Hz) in order to detect miniature eye movements called microsaccades, which have been shown to be crucial for avoiding disappearance of visual image. However, recent studies have demonstrated a close relationship between covert endogenous attentional shifts and the direction of microsaccades. Here, we demonstrate that the direction of microsaccades can be biased away from the peripheral location occupied by a salient, although task-irrelevant, visual signal. Because microsaccades are known not to be under conscious control, our results suggest strong links between IOR and unconscious oculomotor programming.     

The nature of selective attention effects on auditory event-related potentials

The purpose of the research reported here was to examine a number of issues relating to the nature of selective attention effects on auditory event-related potentials (ERPs), namely, to determine the relative contribution of N1 and slow wave (SW) to the early and late components of Nd respectively, where Nd is defined as the negative shift of attended ERPs relative to unattended ERPs; to examine whether individual differences in Nd morphology are related to performance and the strategies that subjects use; and to determine the contribution of changes in the attended and unattended ERPs to Nd. Auditory ERPs were recorded from subjects as they carried out an auditory selective attention task and a visual target detection task. The auditory selective attention task was a multidimensional task in which stimuli varied on location, pitch and duration and in which the subject's task was to pay attention to a particular location/pitch combination and respond whenever they detected a long-duration target tone. In the visual target detection task, subjects were required to respond whenever they detected a colour change in a light-emitting diode which also acted as a fixation point. Auditory ERPs recorded during the visual task were used to provide a measure of exogenous components uncontaminated by differential effects of selective processing of auditory stimuli. The results suggested that early Nd and N1 are independently generated as Nd did not exhibit the contralateral scalp focus typical of N1, and that late Nd is independent of SW. While substantial differences in Nd morphology were observed over subjects, these differences showed no consistent relationships to performance or to task strategies. Comparison of auditory ERPs during active auditory attention with auditory ERPs recorded during the visual control task indicated that there was an early negative shift of the attended ERP, a later negative shift of the attended ERP which had a frontal focus and a later positive shift of the unattended ERP. These results suggest that there are active processes involved in the processing of stimuli from both the attended and unattended source.     

Topographic maps of visual spatial attention in human parietal cortex

Functional magnetic resonance imaging (fMRI) was used to measure activity in human parietal cortex during performance of a visual detection task in which the focus of attention systematically traversed the visual field. Critically, the stimuli were identical on all trials (except for slight contrast changes in a fully randomized selection of the target locations) whereas only the cued location varied. Traveling waves of activity were observed in posterior parietal cortex consistent with shifts in covert attention in the absence of eye movements. The temporal phase of the fMRI signal in each voxel indicated the corresponding visual field location. Visualization of the distribution of temporal phases on a flattened representation of parietal cortex revealed at least two distinct topographically organized cortical areas within the intraparietal sulcus (IPS), each representing the contralateral visual field. Two cortical areas were proposed based on this topographic organization, which we refer to as IPS1 and IPS2 to indicate their locations within the IPS. This nomenclature is neutral with respect to possible homologies with well-established cortical areas in the monkey brain. The two proposed cortical areas exhibited relatively little response to passive visual stimulation in comparison with early visual areas. These results provide evidence for multiple topographic maps in human parietal cortex.     

BOLD repetition decreases in object-responsive ventral visual areas depend on spatial attention

Functional imaging studies of priming-related repetition phenomena have become widely used to study neural object representation. Although blood oxygenation level-dependent (BOLD) repetition decreases can sometimes be observed without awareness of repetition, any role for spatial attention in BOLD repetition effects remains largely unknown. We used fMRI in 13 healthy subjects to test whether BOLD repetition decreases for repeated objects in ventral visual cortices depend on allocation of spatial attention to the prime. Subjects performed a size-judgment task on a probe object that had been attended or ignored in a preceding prime display of 2 lateralized objects. Reaction times showed faster responses when the probe was the same object as the attended prime, independent of the view tested (identical vs. mirror image). No behavioral effect was evident from unattended primes. BOLD repetition decreases for attended primes were found in lateral occipital and fusiform regions bilaterally, which generalized across identical and mirror-image repeats. No repetition decreases were observed for ignored primes. Our results suggest a critical role for attention in achieving visual representations of objects that lead to both BOLD signal decreases and behavioral priming on repeated presentation.     

Target processing is facilitated by motivationally relevant cues

The effect that motivationally relevant stimuli have on processes of attentional engagement and disengagement was investigated during two modified peripheral cueing paradigms. Sexual, mutilation, threatening, and neutral stimuli served as peripheral cues in both experiments. Responses were made to target location in Experiment 1 (N=19 female) and target identity in Experiment 2 (N=18 female). As indexed by enhanced target-evoked P1 and P3b component amplitudes, target processing was facilitated by the presentation of sexual and mutilation stimuli in both experiments. This facilitation in response to targets cued by sexual and mutilation stimuli occurred regardless of whether cueing was valid or invalid as demonstrated by the non-significant cue validity x picture-type interaction. As such, the processes of attentional engagement (as inferred by responses to validly cued targets) and attentional disengagement (as inferred by responses to invalidly cued targets) were not differentially affected by the motivational relevance of the preceding cue. These results indicate that in a non-clinical sample, participants can shift attention rapidly to process information following the onset of motivationally relevant stimuli at attended (valid) and unattended (invalid) locations and that target processing is facilitated by the presence of appetitive and aversive cues.     

Correspondence of Visual Evoked Potentials with FMRI Signals in Human Visual Cortex

Functional magnetic resonance imaging (FMRI) and event related potentials (ERPs) are tools that can be used to image brain activity with relatively good spatial and temporal resolution, respectively. Utilizing both of these methods is therefore desirable in neuroimaging studies to explore the spatio-temporal characteristics of brain function. While several studies have investigated the relationship between EEG and positive (+) BOLD (activation), little is known about the relationship between EEG signals and negative (−) BOLD (deactivation) responses. In this study, we used a visual stimuli designed to shift cortical activity from anterior to posterior regions of the visual cortex. Using EEG and FMRI, we investigated how shifts in +BOLD and −BOLD location were correlated to shifts in the N75 and P100 visual evoked potential (VEP) dipolar sources. The results show that the N75 dipole along with +BOLD, were indeed shifted from posterior to anterior regions of the visual cortex. The P100 VEP component, along with the −BOLD were not shifted to the same extent, indicating that N75 is better correlated to +BOLD than to −BOLD. These findings indicate how different components of the EEG signal are related to the positive and negative BOLD responses, which may aid in interpreting the relationship between visually evoked EEG and FMRI signals. An erratum to this article can be found at