A direct neural measure of variable precision representations in visual working memory.

Visual working memory (VWM) is an active representation enabling the manipulation of item information even in the absence of visual input. A common way to investigate VWM is to analyze the performance at later recall. This approach, however, leaves uncertainties about whether the variation of recall performance is attributable to item encoding and maintenance or to the testing of memorized information. Here, we record the contralateral delay activity (CDA), an established electrophysiological measure of item storage and maintenance, in human subjects performing a delayed orientation precision estimation task. This allows us to link the fluctuation of recall precision directly to the process of item encoding and maintenance. We show that for two sequentially encoded orientation items, the CDA amplitude reflects the precision of orientation recall of both items, with higher precision being associated with a larger amplitude. Furthermore, we show that the CDA amplitudes for the items vary independently from each other, suggesting that the precision of memory representations fluctuates independently.NEW & NOTEWORTHY The present work demonstrates for the first time that the contralateral delay activity (CDA), an online electrophysiological measure of the number of representations maintained in memory, is also a reliable measure of the precision of memory representations. Furthermore, we show that the CDA fluctuates independently for individual items held in memory, thereby providing unambiguous direct neurophysiological support for independently fluctuating memory representations.

Unseen food images capture the attention of hungry viewers: Evidence from event-related potentials.

Motivationally relevant visual targets appear to capture visuospatial attention. This capture is evident behaviorally as faster and more accurate responses, and neurally as an enhanced-amplitude of the N2pc - an index of spatial attention allocation, which is observed even when observers are unaware of the target. In the case of reinforcers such as food or substances of dependence, it is likely that the motivational state of craving accompanying deprivation potentiates this capture. The automaticity of such attentional capture by reward-associated stimuli, as well as its possible interaction with craving, is as yet not completely understood, though it is likely a major explanatory factor in motivated behaviors. For the present experiment, participants completed two EEG recording sessions: one just after eating lunch (sated/non-craving), and the other following a minimum 12-h period of fasting (hungry/craving). For both sessions, participants identified food- and clothing-related targets embedded in an object-substitution masking paradigm, which yielded trials of full target visibility, as well as trials for which targets were present but undetected. Although masking equally disrupted visual awareness of both classes of targets as measured behaviorally, a three-way hunger by visibility by target interaction was observed in the neural data, with unseen food targets eliciting an enhanced N2pc. Interestingly, this subliminal attentional capture by food-related items was observed only during the "hungry" session. No such capture was evident under conditions of full visibility. These findings indicate that attentional capture by food-related images, and reflected in enhancements of the N2pc, is spurred by hunger, and that this effect can be viewed as automatic, or independent of explicit awareness of food-relevant target content.

Neural correlates of multiple object tracking strategies.

Amazingly, human observers can track four independently moving targets. The present study investigated the neural correlates of multiple-object tracking (MOT). Based on previous work we used a modified MOT-task to which subjects exhibited different behaviors. One half of the subjects showed slower RTs and higher error rates with increasing correspondence between tracked items and a probe consisting of 4 highlighted items presented after the tracking. The other half of the subjects had better performance when the probe fully matched the tracked items. Here we sought to investigate the neural representation of the two divergent behavior types. Using multivariate pattern analysis we observed two partly overlapping functional networks associated with the different behaviors. Subjects that responded fast and accurate to full-congruity trials predominantly showed a functional pattern for the full-congruity condition that was very different from patterns associated with any of the partly congruent conditions. This "deviant" pattern was observed in frontal, parietal and extrastriate visual brain areas. In the group of subjects with decreasing performance for increasing target-probe congruity these same regions exhibited a very different functional relationship, in which increasing congruities were associated with linearly changing neural activity patterns. Early low-tier visual areas exclusively exhibited the linear classification pattern while area LO and the primary motor cortex exclusively showed the deviant pattern across all subjects. The coexistence of both networks in groups with different behaviors provides the neural basis for a flexible behavior that can be flexibly adjusted as a function of the strategy employed in the task.

Distinct neural correlates of attending speed vs. coherence of motion.

Attention to specific features of moving visual stimuli modulates the activity in human cortical motion sensitive areas. In this study we employed combined event-related electrophysiological, magnetencephalographic (EEG, MEG) and hemodynamic functional magnetic resonance imaging (fMRI) measures of brain activity to investigate the precise time course and the neural correlates of feature-based attention to speed and coherence. Subjects were presented with an aperture of dots randomly moving either slow or fast, at the same time displaying a high or low level of coherence. The task was to attend either the speed or the coherence and press a button upon the high speed or high coherence stimulus respectively. When attention was directed to the speed of motion enhanced neural activity was found in the dorsal visual area V3a and in the IPL, areas previously shown to be specialized for motion processing. In contrast, when attention was directed to the coherence of motion significant hemodynamic activity was observed in the parietal areas fIPS and SPL that are specialized for the processing of complex motion patterns. Concurrent recordings of the event-related electro- and magnetencephalographic responses revealed that the speed-related attentional modulations of activity occurred at an earlier time range (around 240-290 ms), while the coherence-related ones occurred later (around 320-370 ms) post-stimulus. The current results suggest that the attentional selection of motion features modulates neural processing in the lowest-tier regions required to perform the task-critical discrimination.

Pinning down response inhibition in the brain--conjunction analyses of the Stop-signal task.

Successful behavior requires a finely-tuned interplay of initiating and inhibiting motor programs to react effectively to constantly changing environmental demands. One particularly useful paradigm for investigating inhibitory motor control is the Stop-signal task, where already-initiated responses to Go-stimuli are to be inhibited upon the rapid subsequent presentation of a Stop-stimulus (yielding successful and unsuccessful Stop-trials). Despite the extensive use of this paradigm in functional neuroimaging, there is no consensus on which functional comparison to use to characterize response-inhibition-related brain activity. Here, we utilize conjunction analyses of successful and unsuccessful Stop-trials that are each contrasted against a reference condition. This conjunction approach identifies processes common to both Stop-trial types while excluding processes specific to either, thereby capitalizing on the presence of some response-inhibition-related activity in both conditions. Using this approach on fMRI data from human subjects, we identify a network of brain structures that was linked to both types of Stop-trials, including lateral-inferior frontal and medial frontal cortical areas and the caudate nucleus. In addition, comparisons with a reference condition matched for visual stimulation identified additional activity in the right inferior parietal cortex that may play a role in enhancing the processing of the Stop-stimuli. Finally, differences in stopping efficacy across subjects were associated with variations in activity in the left anterior insula. However, this region was also associated with general task accuracy (which furthermore correlated directly with stopping efficacy), suggesting that it might actually reflect a more general mechanism of performance control that supports response inhibition in a relatively nonspecific way.

Sensory MEG responses predict successful and failed inhibition in a stop-signal task.

In the present study magnetoencephalographic recordings were performed to investigate the neural mechanisms underlying the stopping of manual responses. Subjects performed in a Stop-signal task in which Go-stimuli (S1), requiring a rapid motor response, were sometimes rapidly followed by a Stop-stimulus (S2) indicating to withhold the already initiated response to S1. Success of stopping strongly depended on the early perceptual processing of S1 and S2 reflected by the magnetic N1 component. Enhanced processing of S1 facilitated the execution of the movement, whereas enhanced processing of S2 favored its inhibition. This suggests that the processing resources for the subsequent stimuli are limited and need to be shared. This sharing of resources appeared to arise from adjustments made on a trial-by-trial basis, in that systematic reaction time prolongations on Go-trials following Stop-trials versus following Go-trials were accompanied by attenuated sensory processing to the Go-stimulus similar to that seen in successful versus unsuccessful stopping in Stop-trials.

The center-surround profile of the focus of attention arises from recurrent processing in visual cortex.

We recently demonstrated with magnetoencephalographic recordings in human observers that the focus of attention in visual search has a spatial profile consisting of a center enhancement surrounded by a narrow zone of sensory attenuation. Here, we report new data from 2 experiments providing insights into the cortical processes that cause the surround attenuation. We show that surround suppression appears in search tasks that require spatial scrutiny, that is the precise binding of search-relevant features at the target's location but not in tasks that permit target discrimination without precise localization. Furthermore, we demonstrate that surround attenuation is linked with a stronger recurrent activity modulation in early visual cortex. Finally, we show that surround suppression appears with a delay (more than 175 ms) that is beyond the time course of the initial feedforward sweep of processing in the visual system. These observations together indicate that the suppressive surround is associated with recurrent processing and binding in the visual cortex.

Rapid recurrent processing gates awareness in primary visual cortex.

Visual awareness has been proposed to depend on recurrent processing in early visual cortex areas including the primary visual cortex (V1). Here, we address this hypothesis with high spatiotemporal resolution magnetoencephalographic recordings in subjects performing a substitution masking paradigm. Neural activity reflecting awareness is assessed by directly comparing the neuromagnetic response elicited by effectively and ineffectively masked targets after the proportion of trials leading to masking was individually adjusted to match the proportion of trials without masking. This revealed a modulation of recurrent activity in the primary visual cortex rapidly after the onset of the feedforward sweep of processing in striate and extrastriate areas but significantly before the onset of attention-dependent recurrent modulations in V1. Our data provide direct support for the notion that (i) recurrent processing in V1 correlates with visual awareness and (ii) that attention and awareness involve distinct recurrent processing operations.

Spatio-temporal analysis of feature-based attention.

The cortical mechanisms of feature-selective attention to color and motion cues were studied in humans using combined electrophysiological, magnetoencephalographic, and hemodynamic (functional magnetic resonance imaging) measures of brain activity. Subjects viewed a display of random dots that periodically either changed color or moved coherently. When attention was directed to the color change it elicited enhanced neural activity in visual area V4v, previously shown to be specialized for processing color information. In contrast, when dot movement was attended it produced enhanced activity in the motion-specialized area human MT. Parallel recordings of event-related electrophysiological and magnetoencephalographic responses indicated that the attention-related facilitation of neural activity in these specialized cortical areas occurred rapidly, beginning as early as 90-120 ms after stimulus onset. We conclude that selection of an entire feature dimension (motion or color) boosts neural activity in its specialized cortical module much more rapidly than does selection of one feature value from another (e.g., one color from another), as reported in previous electrophysiological studies. By combining methods with high spatial and temporal resolution it is possible to analyze the precise time course of feature-selective processing in specialized cortical areas.

Direct neurophysiological evidence for spatial suppression surrounding the focus of attention in vision.

The spatial focus of attention has traditionally been envisioned as a simple spatial gradient of enhanced activity that falls off monotonically with increasing distance. Here, we show with high-density magnetoencephalographic recordings in human observers that the focus of attention is not a simple monotonic gradient but instead contains an excitatory peak surrounded by a narrow inhibitory region. To demonstrate this center-surround profile, we asked subjects to focus attention onto a color pop-out target and then presented probe stimuli at various distances from the target. We observed that the electromagnetic response to the probe was enhanced when the probe was presented at the location of the target, but the probe response was suppressed in a narrow zone surrounding the target and then recovered at more distant locations. Withdrawing attention from the pop-out target by engaging observers in a demanding foveal task eliminated this pattern, confirming a truly attention-driven effect. These results indicate that neural enhancement and suppression coexist in a spatially structured manner that is optimal to attenuate the most deleterious noise during visual object identification.

Functional motor compensation in amyotrophic lateral sclerosis.

The present study investigated the fMRI correlates of functional compensation/neural reorganization of the motor system in patients with amyotrophic lateral sclerosis (ALS). The hypothesis was that ALS patients would recruit additional brain regions compared with controls in a motor task and that activity in these regions would vary as a function of task difficulty. Patients and controls executed a motor task with two sequences (a simple and a more difficult one) of consecutive button presses. Patients and controls both activated brain regions known to be involved in motor execution and control. Activity in ipsilateral motor areas as well as difficulty-related activity in the left cerebellum could only be observed in patients. The behavioral data indicated that the motor task was much more difficult for patients than for controls. At nearly equal difficulty the observed patterns of hemodynamic activity in controls were very similar to those observed in ALS. The findings suggest that functional compensation in ALS relies on existing resources and mechanisms that are not primarily developed as a consequence of the lesion.

Functional magnetic resonance tomography correlates of taste perception in the human primary taste cortex.

The present study investigated the functional magnetic resonance tomography correlates of taste perception in the human primary taste cortex. There is conflicting evidence in the literature about chemotopical organization in this brain region. The topography of hemodynamic activity elicited by five taste stimuli (sweet, sour, salty, bitter and umami) was analyzed on the flattened cortical surfaces of six single subjects. A high inter-individual topographical variability had to be noted. The results showed different patterns of hemodynamic activity for the investigated tastes with some considerable overlap. However, the taste specific patterns were stable over time in each subject. Such an individual taste specific pattern was also found for the umami taste within the primary taste cortex of each subject. These results suggest that input from glutamate receptors on the tongue might be processed in an exclusive way in the primary taste cortex rather than as a combination of inputs from the classical taste receptors.

Dynamics of feature binding during object-selective attention.

Objects in the environment may be attended selectively and perceived as unified ensembles of their constituent features. To investigate the timing and cortical localization of feature-integration mechanisms in object-based attention, recordings of event-related potentials and magnetic fields were combined with functional MRI while subjects attended to one of two superimposed transparent surfaces formed by arrays of dots moving in opposite directions. A spatiotemporal analysis revealed evidence for a rapid increase in neural activity localized to a color-selective region of the fusiform gyrus when the surface moving in the attended direction displayed an irrelevant color feature. These data provide support for the "integrated-competition" model of object-selective attention and point to a dynamic neural substrate for the rapid binding process that links relevant and irrelevant features to form a unified perceptual object.

How does attention attenuate target-distractor interference in vision?. Evidence from magnetoencephalographic recordings.

This study used magnetoencephalographic and electroencephalographic recordings to investigate the neural mechanisms that underlie the attentional resolution of ambiguous feature coding in visual search. We addressed this issue by comparing neural activity related to target discrimination under conditions of more versus less feature overlap between the target and distractor items. The results show that increasing feature overlap leads to a focal enhancement of neural activity in ventral occipito-temporal areas, consistent with the larger need to attenuate distractor interference. Furthermore, the results suggest that distractor attenuation proceeds as a stepwise operation, with different spatial locations containing interfering features being suppressed successively. These findings support theories of visual search that emphasize location-based attentional selection as a key mechanism in resolving ambiguous feature coding in vision.

Neural sources of focused attention in visual search.

Previous studies of visual search in humans using event-related potentials (ERPs) have revealed an ERP component called 'N2pc' (180-280 ms) that reflects the focusing of attention onto potential target items in the search array. The present study was designed to localize the neuroanatomical sources of this component by means of magnetoencephalographic (MEG) recordings, which provide greater spatial precision than ERP recordings. MEG recordings were obtained with an array of 148 magnetometers from six normal adult subjects, one of whom was tested in multiple sessions so that both single-subject and group analyses could be performed. Source localization procedures revealed that the N2pc is composed of two distinct neural responses, an early parietal source (180-200 ms) and a later occipito-temporal source (220-240 ms). These findings are consistent with the proposal that parietal areas are used to initiate a shift of attention within a visual search array and that the focusing of attention is implemented by extrastriate areas of the occipital and inferior temporal cortex.

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.

fMRI during word processing in dyslexic and normal reading children.

The present study addresses phonological processing in children with developmental dyslexia. Following the hypothesis of a core deficit of assembled phonology in dyslexia a set of hierarchically structured tasks was applied that specifically control for different kinds of phonological coding (assembled versus addressed phonological strategies). Seventeen developmental dyslexics and 17 normal reading children were scanned during four different tasks: (1) passive viewing of letter strings (control condition), (2) passive reading of non-words, (3) passive reading of legal words, and (4) a task requiring phonological transformation. Statistical analysis of the data was performed using statistical parametric mapping (SPM96). Comparison of patterns of activation in dyslexic and normal reading children revealed significant differences in Broca's area and the left inferior temporal region for both, non-word reading and the phonological transformation task. The present data provide new evidence for alteration of the phonological system in dyslexic children, and in particular, the system that mediates assembled phonological coding.

Event-related brain potentials and case information in syntactic ambiguities.

In an ERP study, German sentences were investigated that contain a case-ambiguous NP that may be assigned accusative or dative case. Sentences were disambiguated by the verb in final position of the sentence. As our data show, sentences ending in a verb that assigns dative case to the ambiguous NP elicit a clear garden-path effect. The garden-path effect was indicated by a broad centro-posterior negative shift that occurred between 300 and 900 msec after the dative-assigning verb was presented. No enhanced P600 following the misanalysis was observed. Noun phrases whose case ambiguity was resolved in favor of accusative case and unambiguously dative-marked NPs did not trigger significant ERP differences. We will discuss the implications of our results for parsing and its neuropsychological correlates. The results of this study support a parser design according to which the so-called structural case (nominative or accusative) is assigned without any delay in the absence of morpho-lexical counterevidence. It is argued that the enhancement of a negative ERP component with a "classical" N400 topography reflects the difficulty of reanalysis due to reaccessing morpho-lexical information that lies outside the domain of the parsing module. Consequently, ERP responses to garden-path effects are not confined to a late positivity but vary depending on the level of processing involved in reanalysis. The fact that garden-path effects may also elicit an N400 can be linked to the nonhomogeneous linguistic properties of the constructions from which they arise.