Location- and Object-Based Representational Mechanisms Account for Bilateral Field Advantage in Multiple-Object Tracking.

Keeping track of multiple visually identical and independently moving objects is a remarkable feature of the human visual system. Theoretical accounts for this ability focus on resource-based models that describe parametric decreases of performance with increasing demands during the task (i.e., more relevant items, closer distances, higher speed). Additionally, the presence of two central tracking resources, one within each hemisphere, has been proposed, allowing for an independent maintenance of moving targets within each visual hemifield. Behavioral evidence in favor of such a model shows that human subjects are able to track almost twice as many targets across both hemifields compared with within one hemifield. A number of recent publications argue for two separate and parallel tracking mechanisms during standard object tracking tasks that allow for the maintenance of the relevant information in a location-based and object-based manner. Unique electrophysiological correlates for each of those processes have been identified. The current study shows that these electrophysiological components are differentially present during tracking within either the left or right hemifield. The present results suggest that targets are mostly maintained as an object-based representation during left hemifield tracking, while location-based resources are preferentially engaged during right hemifield tracking. Interestingly, the manner of representation does not seem to have an impact on behavioral performance within the subjects, while the electrophysiological component indicating object-based tracking does correlate with performance between subjects. We propose that hemifield independence during multiple-object tracking may be an indication of the underlying hemispheric bias for parallel location-based and object-based tracking mechanisms.

Serial attentional resource allocation during parallel feature value tracking.

The visual system has evolved the ability to track features like color and orientation in parallel. This property aligns with the specialization of processing these feature dimensions in the visual cortex. But what if we ask to track changing feature-values within the same feature dimension? Parallel tracking would then have to share the same cortical representation, which would set strong limitations on tracking performance. We address this question by measuring the precision of color representations when human observers track the color of two superimposed dot clouds that simultaneously change color along independent trajectories in color-space. We find that tracking precision is highly imbalanced between streams and that tracking precision changes over time by alternating between streams at a rate of ~1 Hz. These observations suggest that, while parallel color tracking is possible, it is highly limited, essentially allowing for only one color-stream to be tracked with precision at a given time.

A cortical zoom-in operation underlies covert shifts of visual spatial attention.

Shifting the focus of attention without moving the eyes poses challenges for signal coding in visual cortex in terms of spatial resolution, signal routing, and cross-talk. Little is known how these problems are solved during focus shifts. Here, we analyze the spatiotemporal dynamic of neuromagnetic activity in human visual cortex as a function of the size and number of focus shifts in visual search. We find that large shifts elicit activity modulations progressing from highest (IT) through mid-level (V4) to lowest hierarchical levels (V1). Smaller shifts cause those modulations to start at lower levels in the hierarchy. Successive shifts involve repeated backward progressions through the hierarchy. We conclude that covert focus shifts arise from a cortical coarse-to-fine process progressing from retinotopic areas with larger toward areas with smaller receptive fields. This process localizes the target and increases the spatial resolution of selection, which resolves the above issues of cortical coding.

Electrophysiological hallmarks of location-based and object-based visual multiple objects tracking.

Converging evidence shows that our visual system can track multiple visual, independently moving items over time. This is accomplished location-based by maintaining the individual spatial information of each target item or object-based by constructing an abstract object-based representation out of the tracked items. Previous work showed specific behavioural, electrophysiological and haemodynamic markers for location-based or object-based representations of the relevant targets by probing the encoded information subsequently after tracking. However, domain-specific differences of representational correlates during visual tracking itself have not been reported yet. The current study aims to identify spectral properties of the electrophysiological signal during tracking that might indicate location-based versus object-based maintenance of visual information. Subjects had to covertly track four out of eight visually identical items for several seconds while electrophysiological signals were recorded. Subsequently, a probe consisting of four items appeared and the subjects had to indicate with a button press whether the probe matched all targets or not. Subjects employing an object-based strategy showed an enhanced gamma response during the presentation of the target items at the beginning of the trial. On the other hand, subjects using a location-based strategy showed enhanced gamma synchronization throughout the tracking itself. Both the object- and location-based gamma responses yielded identical spatial topographical field distributions. These results indicate that object-based tracking is supported by enhanced encoding during the initial presentation of the targets to be tracked. Location-based tracking is characterized by the sustained maintenance of the individual targets during the entire tracking period in that same processing network.

Attention expedites target selection by prioritizing the neural processing of distractor features.

Whether doing the shopping, or driving the car - to navigate daily life, our brain has to rapidly identify relevant color signals among distracting ones. Despite a wealth of research, how color attention is dynamically adjusted is little understood. Previous studies suggest that the speed of feature attention depends on the time it takes to enhance the neural gain of cortical units tuned to the attended feature. To test this idea, we had human participants switch their attention on the fly between unpredicted target color alternatives, while recording the electromagnetic brain response to probes matching the target, a non-target, or a distracting alternative target color. Paradoxically, we observed a temporally prioritized processing of distractor colors. A larger neural modulation for the distractor followed by its stronger attenuation expedited target identification. Our results suggest that dynamic adjustments of feature attention involve the temporally prioritized processing and elimination of distracting feature representations.

Parallel fast and slow recurrent cortical processing mediates target and distractor selection in visual search.

Visual search has been commonly used to study the neural correlates of attentional allocation in space. Recent electrophysiological research has disentangled distractor processing from target processing, showing that these mechanisms appear to operate in parallel and show electric fields of opposite polarity. Nevertheless, the localization and exact nature of this activity is unknown. Here, using MEG in humans, we provide a spatiotemporal characterization of target and distractor processing in visual cortex. We demonstrate that source activity underlying target- and distractor-processing propagates in parallel as fast and slow sweep from higher to lower hierarchical levels in visual cortex. Importantly, the fast propagating target-related source activity bypasses intermediate levels to go directly to V1, and this V1 activity correlates with behavioral performance. These findings suggest that reentrant processing is important for both selection and attenuation of stimuli, and such processing operates in parallel feedback loops.

Decoding the covert shift of spatial attention from electroencephalographic signals permits reliable control of a brain-computer interface.

OBJECTIVE: One of the main goals of brain-computer interfaces (BCI) is to restore communication abilities in patients. BCIs often use event-related potentials (ERPs) like the P300 which signals the presence of a target in a stream of stimuli. The P300 and related approaches, however, are inherently limited, as they require many stimulus presentations to obtain a usable control signal. Many approaches depend on gaze direction to focus the target, which is also not a viable approach in many cases, because eye movements might be impaired in potential users. Here we report on a BCI that avoids both shortcomings by decoding spatial target information, independent of gaze shifts. APPROACH: We present a new method to decode from the electroencephalogram (EEG) covert shifts of attention to one out of four targets simultaneously presented in the left and right visual field. The task is designed to evoke the N2pc component-a hemisphere lateralized response, elicited over the occipital scalp contralateral to the attended target. The decoding approach involves decoding of the N2pc based on data-driven estimation of spatial filters and a correlation measure. MAIN RESULTS: Despite variability of decoding performance across subjects, 22 out of 24 subjects performed well above chance level. Six subjects even exceeded 80% (cross-validated: 89%) correct predictions in a four-class discrimination task. Hence, the single-trial N2pc proves to be a component that allows for reliable BCI control. An offline analysis of the EEG data with respect to their dependence on stimulation time and number of classes demonstrates that the present method is also a workable approach for two-class tasks. SIGNIFICANCE: Our method extends the range of strategies for gaze-independent BCI control. The proposed decoding approach has the potential to be efficient in similar applications intended to decode ERPs.

Modulating the global orientation bias of the visual system changes population receptive field elongations.

The topographical structure of the visual system in individual subjects can be visualized using fMRI. Recently, a radial bias for the long axis of population receptive fields (pRF) has been shown using fMRI. It has been theorized that the elongation of receptive fields pointing toward the fovea results from horizontal local connections bundling orientation selective units mostly parallel to their polar position within the visual field. In order to investigate whether there is a causal relationship between orientation selectivity and pRF elongation the current study employed a global orientation adapter to modulate the orientation bias for the visual system while measuring spatial pRF characteristics. The hypothesis was that the orientation tuning change of neural populations would alter pRF elongations toward the fovea particularly at axial positions parallel and orthogonal to the affected orientation. The results indeed show a different amount of elongation of pRF units and their orientation at parallel and orthogonal axial positions relative to the adapter orientation. Within the lower left hemifield, pRF radial bias and elongation showed an increase during adaptation to a 135° grating while both parameters decreased during the presentation of a 45° adapter stimulus. The lower right visual field showed the reverse pattern. No modulation of the pRF topographies were observed in the upper visual field probably due to a vertical visual field asymmetry of sensitivity toward the low contrast spatial frequency pattern of the adapter stimulus. These data suggest a direct relationship between orientation selectivity and elongation of population units within the visual cortex.

Electroencephalography reveals a selective disruption of cognitive control processes in craving cigarette smokers.

Addiction to nicotine is extremely challenging to overcome, and the intense craving for the next cigarette often leads to relapse in smokers who wish to quit. To dampen the urges of craving and inhibit unwanted behaviour, smokers must harness cognitive control, which is itself impaired in addiction. It is likely that craving may interact with cognitive control, and the present study sought to test the specificity of such interactions. To this end, data from 24 smokers were gathered using EEG and behavioural measures in a craving session (following a three-hour nicotine abstention period) and a non-craving session (having just smoked). In both sessions, participants performed a task probing various facets of cognitive control (response inhibition, task switching and conflict processing). Results showed that craving smokers were less flexible with the implementation of cognitive control, with demands of task switching and incongruency yielding greater deficits under conditions of craving. Importantly, inhibitory control was not affected by craving, suggesting that the interactions of craving and cognitive control are selective. Together, these results provide evidence that smokers already exhibit specific control-related deficits after brief nicotine deprivation. This disruption of cognitive control while craving may help to explain why abstinence is so difficult to maintain.

High Working Memory Capacity at the Cost of Precision?

Working memory capacity (WMC) varies tremendously among individuals. Here, we investigate the possibility that subjects with high WMC use this limited resource more efficiently by reducing the precision with which they store information in demanding tasks. Task difficulty was increased by (a) presenting more objects to be memorized, (b) informing subjects only after the encoding phase about the relevant objects, and (c) delivering distracting features at retrieval. Precision was assessed by means of a continuous delayed-estimation task, in which object features had to be estimated from memory. High WMC subjects did not show a stronger drop in precision in difficult tasks. Instead, a positive correlation between precision and general WMC emerged. These findings suggest that high WMC subjects do not necessarily trade in quantity for quality when forming working memory (WM) representations under increasing demand. Instead, they seem to be able to devote more cognitive resources to support WM storage.

A neural hallmark of auditory implicit learning is altered in older adults.

Temporal regularities in the environment are often learned implicitly. In an auditory target-detection paradigm using EEG, Jongsma and colleagues (2006) showed that the neural response to these implicit regularities results in a reduction of the P3-N2 complex. Here, we utilized the same paradigm, this time in both young and old participants, to determine if this EEG signature of implicit learning was altered with age. Behaviorally, both groups of participants showed similar benefits for the presence of temporal regularity, with faster and more accurate responses given when the auditory targets were presented in a temporally regular vs. random pattern. In the brain, the younger adults showed the expected decrease in amplitude of this complex for regular compared to irregular trials. Older adults, in contrast, showed no difference in the amplitude of the P3-N2 complex between the irregular and regular condition. These data suggest that, although auditory implicit learning may be behaviorally spared in aging, older adults are not using the same neural substrates as younger adults to achieve this.

Dissociating Reward- and Attention-driven Biasing of Global Feature-based Selection in Human Visual Cortex.

Objects that promise rewards are prioritized for visual selection. The way this prioritization shapes sensory processing in visual cortex, however, is debated. It has been suggested that rewards motivate stronger attentional focusing, resulting in a modulation of sensory selection in early visual cortex. An open question is whether those reward-driven modulations would be independent of similar modulations indexing the selection of attended features that are not associated with reward. Here, we use magnetoencephalography in human observers to investigate whether the modulations indexing global color-based selection in visual cortex are separable for target- and (monetary) reward-defining colors. To assess the underlying global color-based activity modulation, we compare the event-related magnetic field response elicited by a color probe in the unattended hemifield drawn either in the target color, the reward color, both colors, or a neutral task-irrelevant color. To test whether target and reward relevance trigger separable modulations, we manipulate attention demands on target selection while keeping reward-defining experimental parameters constant. Replicating previous observations, we find that reward and target relevance produce almost indistinguishable gain modulations in ventral extratriate cortex contralateral to the unattended color probe. Importantly, increasing attention demands on target discrimination increases the response to the target-defining color, whereas the response to the rewarded color remains largely unchanged. These observations indicate that, although task relevance and reward influence the very same feature-selective area in extrastriate visual cortex, the associated modulations are largely independent.

Enhanced spatial focusing increases feature-based selection in unattended locations.

Attention is a multifaceted phenomenon, which operates on features (e.g., colour or motion) and over space. A fundamental question is whether the attentional selection of features is confined to the spatially-attended location or operates independently across the entire visual field (global feature-based attention, GFBA). Studies providing evidence for GFBA often employ feature probes presented at spatially unattended locations, which elicit enhanced brain responses when they match a currently-attended target feature. However, the validity of this interpretation relies on consistent spatial focusing onto the target. If the probe were to temporarily attract spatial attention, the reported effects could reflect transient spatial selection processes, rather than GFBA. Here, using magnetoencephalographic recordings (MEG) in humans, we manipulate the strength and consistency of spatial focusing to the target by increasing the target discrimination difficulty (Experiment 1), and by demarcating the upcoming target's location with a placeholder (Experiment 2), to see if GFBA effects are preserved. We observe that motivating stronger spatial focusing to the target did not diminish the effects of GFBA. Instead, aiding spatial pre-focusing with a placeholder enhanced the feature response at unattended locations. Our findings confirm that feature selection effects measured with spatially-unattended probes reflect a true location-independent neural bias.

Cortical Mechanisms of Prioritizing Selection for Rejection in Visual Search.

In visual search, the more one knows about a target, the faster one can find it. Surprisingly, target identification is also faster with knowledge about distractor-features. The latter is paradoxical, as it implies that to avoid the selection of an item, the item must somehow be selected to some degree. This conundrum has been termed the "ignoring paradox", and, to date, little is known about how the brain resolves it. Here, in data from four experiments using neuromagnetic brain recordings in male and female humans, we provide evidence that this paradox is resolved by giving distracting information priority in cortical processing. This attentional priority to distractors manifests as an enhanced early neuromagnetic index, which occurs before target-related processing, and regardless of distractor predictability. It is most pronounced on trials for which a response rapidly occurred, and is followed by a suppression of the distracting information. These observations together suggest that in visual search items cannot be ignored without first being selected.SIGNIFICANCE STATEMENT How can we ignore distracting stimuli in our environment? To do this successfully, a logical hypothesis is that as few neural resources as possible should be devoted to distractor processing. Yet, to avoid devoting resources to a distractor, the brain must somehow mark what to avoid; this is a philosophical problem, which has been termed the "ignoring paradox" or "white bear phenomenon". Here, we use MEG recordings to determine how the human brain resolves this paradox. Our data show that distractors are not only processed, they are given temporal priority, with the brain building a robust representation of the to-be-ignored items. Thus, successful suppression of distractors can only be achieved if distractors are first strongly neurally represented.

Spatial elongation of population receptive field profiles revealed by model-free fMRI back-projection.

Estimates of visual field topographies in human visual cortex obtained through fMRI traveling wave techniques usually provide the parameters of population receptive field (pRF) location (polar angle, eccentricity) and receptive field size. These parameters are obtained by fitting the recorded data to a standard model population receptive field. In this work, pRF profiles are measured directly by back-projecting preprocessed fMRI time-series to sweeps of a bar across the visual field in different angles. The current data suggest that the model-free pRF profiles contain information not only about receptive field location and size but also about the pRF shape characteristics. The elongation (ellipticity) of pRFs decreases along the early visual hierarchy to a different degree for the ventral and the dorsal stream. Furthermore, ellipticity changes as a function of eccentricity. pRF orientation shows a high degree of collinearity with its angular position within the visual field. Using model-free pRF measurements, the traveling wave technique provides additional characteristics of pRF topographies that are not restricted to size and provide robust measures within the single subject.

Attention to Color Sharpens Neural Population Tuning via Feedback Processing in the Human Visual Cortex Hierarchy.

Attention can facilitate the selection of elementary object features such as color, orientation, or motion. This is referred to as feature-based attention and it is commonly attributed to a modulation of the gain and tuning of feature-selective units in visual cortex. Although gain mechanisms are well characterized, little is known about the cortical processes underlying the sharpening of feature selectivity. Here, we show with high-resolution magnetoencephalography in human observers (men and women) that sharpened selectivity for a particular color arises from feedback processing in the human visual cortex hierarchy. To assess color selectivity, we analyze the response to a color probe that varies in color distance from an attended color target. We find that attention causes an initial gain enhancement in anterior ventral extrastriate cortex that is coarsely selective for the target color and transitions within ∼100 ms into a sharper tuned profile in more posterior ventral occipital cortex. We conclude that attention sharpens selectivity over time by attenuating the response at lower levels of the cortical hierarchy to color values neighboring the target in color space. These observations support computational models proposing that attention tunes feature selectivity in visual cortex through backward-propagating attenuation of units less tuned to the target.SIGNIFICANCE STATEMENT Whether searching for your car, a particular item of clothing, or just obeying traffic lights, in everyday life, we must select items based on color. But how does attention allow us to select a specific color? Here, we use high spatiotemporal resolution neuromagnetic recordings to examine how color selectivity emerges in the human brain. We find that color selectivity evolves as a coarse to fine process from higher to lower levels within the visual cortex hierarchy. Our observations support computational models proposing that feature selectivity increases over time by attenuating the responses of less-selective cells in lower-level brain areas. These data emphasize that color perception involves multiple areas across a hierarchy of regions, interacting with each other in a complex, recursive manner.

Active prosthesis dependent functional cortical reorganization following stroke.

The present study investigated the neural correlates associated with gait improvements triggered by an active prosthesis in patients with drop-foot following stroke during the chronic stage. Eleven patients took part in the study. MEG recordings in conjunction with somatosensory stimulation of the left and right hand as well as gait analyses were performed shortly before or after prosthesis implantation surgery and 3-4 months later. Plastic changes of the sensorimotor cortex of the ipsi- and contralesional hemisphere were revealed. Gait analysis indicated that all patients improved their gait with the active prosthesis. Patients with larger plastic changes within the lesioned hemisphere maintained their improved gait performance even when the prosthesis was turned off. Patients with larger contralesional changes also improved their gait with the active prosthesis. However, their gait measures decreased when the prosthesis was turned off. The current data provide the neural basis of gait improvement triggered by an active prosthesis and has important implications with respect to the choice of the type of active prosthesis (implantable vs removable) and to the selection procedure of the patients (length of testing period).

Somatosensory Misrepresentation Associated with Chronic Pain: Spatiotemporal Correlates of Sensory Perception in a Patient following a Complex Regional Pain Syndrome Spread.

Chronic pain is suggested to be linked to reorganization processes in the sensorimotor cortex. In the current study, the somatosensory representation of the extremities was investigated in a patient with a complex regional pain syndrome (CRPS) that initially occurred in the right hand and arm and spread later into the left hand and right leg. After the spread, magnetoencephalographic recordings in conjunction with somatosensory stimulation revealed that the clinical symptoms were associated with major changes in the primary somatosensory representation. Tactile stimulation of body parts triggering CRPS-related pain elicited activity located in the left primary somatosensory region corresponding to the right hand representation, where the CRPS initially appeared. Solely the unaffected left foot was observed to have a regular S1 representation. The pain distribution pattern was matching the cortical somatosensory misrepresentation suggesting that cortical reorganization processes might contribute and possibly underlie the development and spread of the CRPS.

Spatio-temporal dynamics of attentional selection stages during multiple object tracking.

Subjects can visually track several moving items simultaneously, a fact that is difficult to explain by classical attention models. Previous work revealed that building a global shape based on the spatial position of the tracked items improves performance. Here we investigated the involved neural processes and the role of attention. A task-irrelevant probe stimulus was presented during multiple objects tracking at a fixed spatial location. Depending on the tracked item's trajectories the probe appeared either outside, inside, or on the edge of aforementioned global shape. Event-related potentials to the probe stimulus revealed two subsequent stages of attentional selection during multiple object tracking. After 100ms attention was deployed on the edge/boundary of the figure formed by the tracked items. In the following 80ms, attention spread from the outline to the full figure. These findings clarify the eminent contribution of attentional mechanisms in multiple objects tracking.

Filtering and storage working memory networks in younger and older age.

INTRODUCTION: Working memory (WM) is a multi-component model that among others involves the two processes of filtering and storage. The first reflects the necessity to inhibit irrelevant information from entering memory, whereas the latter refers to the active maintenance of object representations in memory. In this study, we aimed at a) redefining the neuronal networks sustaining filtering and storage within visual working memory by avoiding shortcomings of prior studies, and b) assessing age-related changes in these networks. METHODS: We designed a new paradigm that strictly controlled for perceptual load by presenting the same number of stimuli in each of three conditions. We calculated fMRI contrasts between a baseline condition (low filter and low storage load) and conditions that posed high demands on filtering and storage, respectively, in large samples of younger (n = 40) and elder (n = 38) participants. RESULTS: Our approach of comparing contrasts between groups revealed more extensive filter and storage WM networks than previous studies. In the younger group, filtering involved the bilateral insulae, the right occipital cortex, the right brainstem, and the right cerebellum. In the elder group, filtering was associated with the bilateral insulae, right precuneus, and bilateral ventromedial prefrontal cortex. An extensive neuronal network was also found during storage of information in the bilateral posterior parietal cortex, the left ventromedial prefrontal cortex, and the right precuneus in the younger participants. In addition to these brain regions, elder participants recruited the bilateral ventral prefrontal cortex, the superior, middle and inferior and temporal cortex, the left cingulum and the bilateral parahippocampal cortex. CONCLUSIONS: In general, elder participants recruited more brain regions in comparison to younger participants to reach similar accuracy levels. Furthermore, in elder participants one brain region emerged in both contrasts, namely the left ventromedial prefrontal cortex. Hence, elder participants seem to routinely recruit this brain region in demanding tasks, irrespective of whether filtering or storing is challenged.