Biology subjects

Chota, S.

Publications and source records attributed to Chota, S..

15 recordsLinked to original sources

bioRxiv2026

Covert attentional tracking with invisible phase tagging

Attention forms a fundamental component of visual processing. The locus and spread of covert attention reveals how specific locations or features in the environment are preferentially processed, however, tracking this process in a time-resolved fashion is not easy. Rapid Invisible Frequency Tagging (RIFT) is a recent advance in localizing covert attention. This technique uses stimuli that flicker at unique frequencies to invisibly "tag" areas of the visual field. The frequency-specific response elicited offers a measure of visual processing (and covert attention) over time. However, the narrow frequency band compatible with RIFT restricts the number of distinct frequency tags that can be simultaneously used. This limitation can be addressed by using oscillatory phase as an additional tagging feature. Here, we conducted a RIFT-EEG experiment (n=24) with 8 unique tags based on both frequency and phase modulation. We show that modulations in visual processing can be measured at several tagged locations with identical frequency but different phases. This also allowed for the use of combined phase- and frequency- modulated RIFT to quantify the distribution of attention across the visual field. By integrating phase-modulation into invisible frequency tagging studies of attention, we bring the field a step closer towards fine-resolution attentional tracking continuously over time and space, allowing novel insights across various domains of visual cognition.

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bioRxiv2026

Visual Attention doesnt Travel, it Teleports

Attention forms a key, yet elusive component of visual processing. We shift attention constantly across the visual field to enhance processing of relevant locations or stimuli in service of goal-directed behavior. Here, we investigate a fundamental property of attentional shifts: when covertly shifting from one location to another, does visual attention "travel" (enhance processing at intermediate locations) or "teleport" (not interact with intermediate locations)? While most shifts of information or movement "travel" (e.g., eye and body movements, neuronal signalling), for attentional shifts such intermediate processing enhancement might not be necessary, nor functional. To answer this question while tackling the difficulty of tracking covert attentional dynamics, we conducted an EEG-eyetracking experiment (n=24) paired with Rapid Invisible Frequency Tagging (RIFT). This let us track attentional enhancement during covert attentional shifts with high temporal and spatial precision. We successfully registered covert shifts, but found no evidence for any attentional modulation in-between the start and end-point of an attentional shift. Additional insilico modelling of attentional shifts confirmed that our method was sensitive enough to pick up on these modulations had they been present. Our results support a "teleporting" model of attention, suggesting that attention is implemented in a fundamentally different manner compared to overt visual behaviours such as eye movements.

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bioRxiv2025

Independent Markers of Attentional Shifts

Visuospatial attention has been extensively studied using a wide variety of markers in the brain and in behaviour. We can broadly classify these markers into two distinct groups: reactive measures, where the presence and strength of attentional biases is inferred from the reaction to a probe stimulus, and proactive measures, which track attentional biases without (or before) the presentation of a probe stimulus. A difference in either type of measure (between two experimental conditions) is typically interpreted to reflect a difference in attentional resources. However, it is not a given that these various reactive and proactive measures of attention all read out the same construct; they may very well tap into independent processes. Here, we measure some of these proactive and reactive signatures of attending a location (changes in cortical excitability as measured by RIFT, alpha lateralization, gaze bias, probe-evoked ERPs, and behavioral reports) in an EEG-eyetracking task (n=23). Though all these metrics indeed correlate with our attentional manipulation, we show that they are not direct correlates of each other, and that of all the proactive measures only the gaze bias predicts subsequent behavioral responses. This demonstrates that existing markers of attention cannot be used interchangeably, and likely capture independent subprocesses of attention.

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bioRxiv2025

Tracking attention using RIFT with a consumer-monitor setup

Rapid Invisible Frequency Tagging (RIFT) is a recent technique that extends the traditional frequency tagging approach by modulating the luminance of distinct locations on a screen at frequencies beyond the threshold of perception ([≥]60Hz). Retrieving the corresponding frequencies from M/EEG recordings provides a time-resolved and location-specific measure of early visual processing, without the confounding effect of introducing visible stimuli. This method is most frequently harnessed as a tracker of covert attention in experimental paradigms across various disciplines in cognitive neuroscience. However, almost all existing RIFT work so far has made use of expensive display hardware limited in its accessibility. Recent work has shown that a RIFT response elicited by a 480Hz refresh-rate consumer monitor can be retrieved from the EEG, but it is not yet clear whether such a setup can be utilized to track the locus of attention. Here, using a spatial cueing paradigm (n=24) while simultaneously tagging two locations (60.0Hz and 65.5Hz), we show that covert attention can be reliably measured with RIFT on a 360Hz refresh rate consumer monitor. We hope that this study will facilitate the widespread application of using consumer-grade high refresh-rate gaming monitors with RIFT for future scientific and applied research.

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bioRxiv2025

Dynamic competition between bottom-up saliency and top-down goals in early visual cortex

Task-irrelevant yet salient stimuli can elicit automatic, bottom-up attentional capture and compete with top-down, goal-directed processes for neural representation. However, the temporal dynamics underlying this competition, and how they influence early visual processing, remain poorly understood. Here, we combined electroencephalography (EEG) with Rapid Invisible Frequency Tagging (RIFT) to non-invasively and simultaneously track early visual cortex responses to target and distractor. Both target and distractor evoked stronger initial RIFT responses than nontargets, reflecting top-down and bottom-up attentional effects on early visual processing. Importantly, the presence of a distractor attenuated the initial RIFT response to the target, reflecting competition during the initial stages of visual processing and predicting subsequent behavioral performance. RIFT responses to the distractor eventually even decreased below responses to the target and nontarget, representing active suppression of task-irrelevant but salient stimuli. We show that the dynamic interplay between top-down control and bottom-up saliency directly impacts early visual responses, thereby illuminating a complete timeline of attentional competition in visual cortex.

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bioRxiv2025

Anticipated Relevance Prepares Visual Processing for Efficient Memory-Guided Selection

Finding an object typically involves using working memory to select relevant visual information at the right time. Using EEG, we investigated how predictable changes in stimulus relevance influence top-down visual selection. Participants memorized an oriented grating, followed by a cue indicating which of two sequentially presented probes was relevant for a memory-match/-mismatch judgment. Consistent with earlier work, relevant probes evoked stronger univariate responses than irrelevant probes. Furthermore, responses to (memory-matching and memory-mismatching) probes were more distinct when they were relevant compared to irrelevant. Crucially, using rapid invisible frequency tagging (RIFT), we found that visual responses to the (empty) probe location were enhanced even before the presentation of relevant (compared to irrelevant) probes. Predictable changes in stimulus relevance, therefore, induce both pro-active and re-active changes in visual processing. We conclude that anticipating the (ir)relevance of upcoming visual events enables the visual system to prepare for efficient memory-guided visual selection.

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bioRxiv2024

Gaze biases can reflect task-specific spatial memorization strategies

Previous work has suggested that small directional eye movements not only reveal the focus of external spatial attention towards visible stimuli, but also accompany shifts of internal attention to stimuli in visual working memory (VWM)(van Ede et al., 2019). When the orientations of two bars are memorized and a subsequent retro-cue indicates which orientation needs to be reported, participants gaze is systematically biased towards the former location of the cued item (Figure 1AB). This finding was interpreted as evidence that the oculomotor system indexes internal attention; that is, attention directed at the location of stimuli that are no longer presented but are maintained in VWM. Importantly, as the location of the bars is presumably not relevant to the memory report, the authors concluded that orientation features in VWM are automatically associated with locations, suggesting that VWM is inherently spatially organized. This conclusion depends on the key assumption that participants indeed memorize and subsequently attend orientation features. Here we re-analyse Experiment 1 by van Ede et al. (2019) and demonstrate that this assumption does not hold. Instead of memorizing orientation features, participants deployed an alternative spatial strategy by memorizing bar endpoints. Although we do not call into question the conclusion that internal attention is inherently spatially organized, our results do imply that directional gaze biases might also reflect attention directed at task-relevant stimulus endpoints, rather than internal attention directed at memorized orientations. O_FIG O_LINKSMALLFIG WIDTH=200 HEIGHT=161 SRC="FIGDIR/small/610231v2_fig1.gif" ALT="Figure 1"> View larger version (43K): org.highwire.dtl.DTLVardef@940e51org.highwire.dtl.DTLVardef@37ec3dorg.highwire.dtl.DTLVardef@176b186org.highwire.dtl.DTLVardef@180e8a7_HPS_FORMAT_FIGEXP M_FIG O_FLOATNOFigure 1.C_FLOATNO Gaze density maps from Experiment 1 by van Ede et al. (2019) (N = 23, trials included = 20.864, 400 to 1000 ms). AB. Original reported effect of cued item location on gaze bias. Calculated by subtracting cued-item-left and cued-item-right gaze density maps. Rectangles indicate used stimulus positions and orientation ranges (min: 20{degrees}, mean: 45{degrees}, max: 70{degrees}; min: 110{degrees}, mean: 135{degrees}, max: 160{degrees}) of bar stimuli. C. Normalized Gaze bias vectors per condition (red dotted lines), horizontal vectors (dotted black lines) and average vectors pointing towards most foveal bar endpoints (solid black lines). Gaze bias vector endpoints were calculated from the centre of mass of each condition, ignoring negative values. Circular t-tests revealed that individual gaze bias vector angles (red dotted lines) were significantly different from horizontal vectors (dotted black lines) but not significantly different from endpoint vectors (solid black lines). FI. Vertical gaze bias revealed by separating trials based on bar orientations. Red dotted lines depict group average gaze bias vectors. F. Both bar endpoints "upwards" (left: 20{degrees} to 70{degrees} right: 110{degrees} to 160{degrees}) minus both bars endpoints "downwards" (left: 110{degrees} to 160{degrees}, right: 20{degrees} to 70{degrees}). I. Both "downwards" minus both "upwards". DEGH. Individual gaze density maps for each attention (left versus right) and bar endpoint direction (upwards versus downwards) separately. Solid black Lines show average vector pointing towards closest 45{degrees}/135{degrees} bar endpoint (i.e., average optimal gaze location for solving the memory task through memory maintenance of a spatial location). Red dotted lines depict group average gaze bias vectors (calculated from the centre of mass of each condition, ignoring negative values). C_FIG

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bioRxiv2024

Dissociating External and Internal Attentional Selection

Visual Working Memory (VWM) stores visual information for upcoming actions. Just as attention can shift externally towards relevant objects in the visual environment, attention can shift internally towards (i.e., prioritize) VWM content that is relevant for upcoming tasks. Internal and external attentional selection share a number of key neural and functional characteristics, which include their spatial organization: recent work has shown that spatial attention is directed towards the previous location of a prioritized memory item, similar to how a perceived stimulus is prioritized. Attending stimuli that are physically present is useful, as it enhances processing of the relevant visual input. When prioritizing items in memory, however, attending the prior stimulus location cannot serve this purpose, as there is no visual input to enhance. Here, we address this apparent contradiction which highlights the gaps in our understanding of the mechanisms underlying external and internal visual attention. In two EEG experiments, we compare location-specific sensory enhancement during the attentional selection of external (perceived) as compared to internal (memorized) stimuli. During both internal and external selection we observed a lateralization of alpha oscillations and gaze position bias toward the previous locations of prioritized items, confirming earlier findings that suggested an inherent spatial organization within VWM. Critically, using Rapid Invisible Frequency Tagging (RIFT), we show that sensory enhancement at the attended location is only observed during external attentional selection of (perceived) stimuli. No such location-specific sensory enhancement was observed during attentional selection of items in VWM. Furthermore, we found no clear relationship across trials between alpha lateralization and sensory enhancement (measured through RIFT) during external attention, suggesting that these two metrics indeed reflect distinct cognitive mechanisms. In sum, using a novel combination of EEG and RIFT, we demonstrate a fundamental distinction between the neural mechanisms underlying the selection of perceived and memorized objects. Both types of selection operate within a spatial reference frame, but only external selection modulates early sensory processing. Our findings suggest that the visual system is not vestigially recruiting existing mechanisms of external attention for prioritization in VWM, but is instead using space as an organizational principle to store and select items in VWM.

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bioRxiv2024

Rapid Invisible Frequency Tagging (RIFT) in a novel setup with EEG

Steady-State Visual Evoked Potentials (SSVEPs) provide a report-free and continuous measure of neural processing. Recent progress in display technology has allowed for the tagging of multiple stimuli simultaneously at >60Hz frequencies - high enough to evade perceptibility, while still evoking an oscillatory neural response. Known as Rapid Invisible Frequency Tagging (RIFT), this technique has currently only been used in combination with Magnetoencephalography (MEG), which is less accessible compared to Electroencephalography (EEG). Although responses to LEDs flickering at similar frequencies have been shown in EEG, it is currently unclear whether RIFT, using a more conventional stimulus display, can sufficiently evoke a response in EEG, and therefore whether it is worth adding the RIFT-EEG pairing to the cognitive neuroscientists toolkit. Here, we successfully implement the first RIFT-EEG setup. We show that the oscillatory input is measurable in the EEG trace, what its topographical spread is, a rough range of applicable frequencies, and that this response is comparable to that evoked in MEG.

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bioRxiv2023

SSVEPs reveal dynamic (re-)allocation of spatial attention during maintenance and utilization of visual working memory

Visual Working Memory (VWM) allows us to temporarily store goal-relevant information to guide future behavior. Prior work has established that VWM and spatial attention are intrinsically connected - even if location is irrelevant for responses: behavioral and neural correlates of attention show spatial biases, specific to the location at which memory items were encoded. This suggests that VWM is spatially organized and that maintaining information might rely on the allocation of spatial attention towards the location of memory items. Importantly, attention often needs to be dynamically redistributed between several locations, e.g. in preparation for an upcoming probe. Very little is known about how attentional resources are distributed between multiple locations during a VWM task and even less about the dynamic changes governing such attentional shifts over time. This is largely due to the inability to use behavioral outcomes to reveal fast dynamic changes within trials. We here demonstrate EEG Steady-State Visual Evoked Potentials (SSVEPs) to successfully track the dynamic allocation of spatial attention during a VWM task. Participants were presented with to-be-memorized gratings and distractors at two distinct locations. During maintenance and retrieval, each location was tagged with a disk flickering at either 10 or 13 Hz. This allowed us to dynamically track attention allocated to memory and distractor items via their coupling with space by quantifying the amplitude and coherence of SSVEP responses in the EEG signal to flickering stimuli at the former memory and distractor locations. SSVEP responses at memory locations did not differ from distractor locations during early parts of the maintenance window. However, shortly before probe comparison, we observed a decrease in SSVEP coherence over distractor locations indicative of a reallocation of spatial attentional resources. Reaction times were shorter when preceded by stronger decreases in SSVEP coherence at distractor locations, reflecting the reallocation of attention from the distractor location to the memory location or towards the distinct upcoming probe location. Broader Significance We demonstrate that SSVEPs can inform about dynamic processes in VWM, even if location does not have to be reported by participants. This finding not only supports the notion of a spatially organized VWM, but also reveals that SSVEPs betray a dynamic prioritization process of working memory items and locations over time that is directly predictive of memory performance.

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bioRxiv2022

A matter of availability: Neural representations of task-relevant stimulus features are sharper when stimuli are memorized rather than perceived.

Our visual environment is relatively stable over time and an optimized visual system ought to capitalize on this by not devoting any representational resources to objects that are still present. Subjective experience, however, suggests that externally available (i.e., perceived) information is more strongly represented in neural signals than memorized information. To distinguish between these possibilities, we use EEG multivariate pattern analysis to quantify the strength of representation of task-relevant features (color or spatial frequency) in anticipation of a change-detection task. Perceptual availability was manipulated between experimental blocks by either keeping the stimulus on the screen during a two second delay period (perception) or removing it shortly after its initial presentation for the same time period (memory). We find that task-relevant (i.e., attended) memorized features are more strongly represented than irrelevant features. More importantly, we find significantly weaker representations for available (perceived and attended) features than for unavailable (memorized and attended) features. Contrary to what subjective experience suggests, our findings demonstrate that vividly perceived and attended stimuli elicit weaker neural representations (in terms of detectable multivariate information) than stimuli maintained in visual working memory. We hypothesize that an efficient visual system spends little of its limited resources on the internal representation of information that is externally available anyway.

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bioRxiv2022

Random Tactile Noise Stimulation Reveals Beta-Rhythmic Impulse Response Function of the Somatosensory System

Both passive tactile stimulation and motor actions result in dynamic changes in beta-band (15-30 Hz Hz) oscillations over somatosensory cortex. Similar to alpha-band (8-12 Hz) power decrease in the visual system, beta-band power also decreases following stimulation of the somatosensory system. This relative suppression of alpha and beta oscillations is generally interpreted as an increase in cortical excitability. Here, next to traditional single-pulse stimuli, we employed a random intensity continuous right index finger tactile stimulation (white noise), which enabled us to uncover an impulse response function (IRF) of the somatosensory system. Contrary to previous findings, we demonstrate a burst-like initial increase rather than decrease of beta activity following white noise stimulation (human participants, N = 18, 8 female). These beta bursts, on average, lasted for 3 cycles and their frequency was correlated with resonant frequency of somatosensory cortex, as measured by a multi-frequency steady-state somatosensory evoked potential (SSSEP) paradigm. Furthermore, beta-band bursts shared spectro-temporal characteristics with evoked and resting-state beta oscillations. Taken together, our findings not only reveal a novel oscillatory signature of somatosensory processing that mimics the previously reported visual IRFs, but also point to a common oscillatory generator underlying spontaneous beta bursts in the absence of tactile stimulation and phase-locked beta bursts following stimulation, the frequency of which is determined by the resonance properties of the somatosensory system. Significance StatementThe investigation of the transient nature of oscillations has gained great popularity in recent years. The findings of bursting activity rather than sustained oscillations in the beta-band has provided important insights into its role in movement planning, working memory, inhibition and reactivation of neural ensembles. In this study, we show that also in response to tactile stimulation the somatosensory system responds with [~]3 cycle oscillatory beta-band bursts, whose spectro-temporal characteristics are shared with evoked and resting-state beta-band oscillatory signatures of the somatosensory system. As similar bursts have been observed in the visual domain, these oscillatory signatures might reflect an important supramodal mechanism in sensory processing.

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bioRxiv2022

Attention Samples Items in Visual Working Memory Rhythmically

Attention allows us to selectively enhance the processing of specific locations or features in our external environment while simultaneously filtering out momentarily irrelevant information. It is currently hypothesized that this is achieved through the boosting of relevant sensory signals which biases the competition between competing neural representations. Recent neurophysiological and behavioral studies have revealed that attention is a fundamentally rhythmic process, tightly linked to neural oscillations in fronto-parietal networks. Instead of continuously highlighting a single object or location, attention rhythmically alternates between multiple relevant representations at a frequency of 3 - 6 Hz. However attention can not only be directed towards the external world but also towards internal visual working memory (VWM) representations, e.g. when selecting one of several search templates to find corresponding objects in the external world. Two recent studies have revealed that objects in VWM are attended in a similarly rhythmic fashion as perceived objects. We add to the current literature by showing that retro-cues towards multi-feature gratings in VWM initiate a similar theta-rhythmic competition, modulating reaction times in an anti-phasic manner. Our findings add to the converging body of evidence that external and internal visual representations are accessed by highly similar, rhythmic attentional mechanisms.

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bioRxiv2020

Occipital Alpha-TMS causally modulates Temporal Order Judgements: Evidence for discrete temporal windows in vision

Recent advances in neuroscience have challenged the view of conscious visual perception as a continuous process. Behavioral performance, reaction times and some visual illusions all undergo periodic fluctuations that can be traced back to oscillatory activity in the brain. These findings have given rise to the idea of a discrete sampling mechanism in the visual system. In this study we seek to investigate the causal relationship between occipital alpha oscillations and Temporal Order Judgements using neural entrainment via rhythmic TMS. We find that certain phases of the entrained oscillation facilitate temporal order perception of two visual stimuli, whereas others hinder it. Our findings support the idea that the visual system periodically compresses information into discrete packages within which temporal order information is lost. Significance StatementCorrelational evidence for a relationship between brain states and perception is abundant. However, in order to truly understand the causal relationship between the two, we need to be able to manipulate one and observe changes in the other. Neural entrainment via TMS serves as a valuable tool to interfere with cortical rhythms and observe changes in perception. Here, using rhythmic TMS-pulses at 10 Hz, we investigate the effect of the phase of entrained oscillations on performance in a temporal order judgement (TOJ) task. We observe that the causally entrained oscillation indeed modulates time perception rhythmically. On top of previous work on discrete perception we were able to 1. causally influence brain rhythms in a far more direct fashion using TMS, and 2. show that previous work on discrete perception cannot simply be explained by rhythmic fluctuations in visibility. In conclusion our findings support the long discussed idea that the temporal organization of visual processing is discrete rather than continuous, and is causally modulated by cortical rhythms. To our knowledge, this is the first study providing causal evidence via TMS for an endogenous periodic modulation of time perception.

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bioRxiv2019

Visual Entrainment at 10 Hz causes periodic modulation of the Flash Lag Illusion

It has long been debated whether visual processing is, at least partially, a discrete process. Although vision appears to be a continuous stream of sensory information, sophisticated experiments reveal periodic modulations of perception and behavior. Previous work has demonstrated that the phase of endogenous neural oscillations in the 10 Hz range predicts the \"lag\" of the flash lag effect, a temporal visual illusion in which a static object is perceived to be lagging in time behind a moving object. Consequently, it has been proposed that the flash lag illusion could be a manifestation of a periodic, discrete sampling mechanism in the visual system. In this experiment we set out to causally test this hypothesis by entraining the visual system to a periodic 10 Hz stimulus and probing the flash lag effect (FLE) at different time points during entrainment. We hypothesized that the perceived FLE would be modulated over time, at the same frequency as the entrainer (10 Hz). A frequency analysis of the average FLE time-course indeed reveals a significant peak at 10 Hz as well as a strong phase consistency between subjects (N=26). Our findings provide evidence for a causal relationship between alpha oscillations and fluctuations in temporal perception.

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