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Gayet, S.

Publications and source records attributed to Gayet, S..

9 recordsLinked to original sources

Decoding auditory working memory load from EEG alpha oscillations

Working memory (WM) enables temporary retention of task-relevant information for imminent use. Increases in visual WM load are accompanied by elevated contralateral delay activity (CDA), and EEG alpha-band power. While most WM research focuses on the visual domain, it remains unknown whether similar EEG responses also reflect WM load in the auditory domain. Using EEG, we set out to establish such neural markers of auditory WM load. Participants memorized the pitches of 1 to 4 pure tones presented to one ear, with 1 to 4 identical distractor tones presented to the other ear. Behaviorally, auditory WM capacity plateaued between set-sizes two and three. Unlike for visual WM, auditory WM load was not reflected in lateralized EEG responses. This shows that the CDA is a vision-specific rather than domain-general neural marker of WM load. Applying multivariate pattern analyses on the delay activity revealed that auditory WM load is reflected in patterns of alpha-band oscillations. Surprisingly, a temporal generalization analysis revealed that the alpha patterns reflecting specific load conditions changed throughout the maintenance period (despite load being inherently constant), revealing dynamic coding of auditory WM load.

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Dynamic context-based updating of object representations in visual cortex

Objects in real-world scenes are often poorly or partially visible, for example because they are occluded or appear in the periphery. An additional challenge of real-world vision is that it is dynamic, causing the appearance of objects (e.g., their size and orientation) to change as we move. Importantly, however, these changes are predictable from the 3D structure of the surrounding scene. In two fMRI studies, we find that visual cortex dynamically updates object representations using this predictive contextual information. Firstly, visual cortical representations of objects were enhanced when they rotated congruently (versus incongruently) with the surrounding scene. Secondly, the inferred orientation of the object could be decoded from visual cortex activity even when the object was fully occluded. These findings indicate that predictive processes in visual cortex follow the geometric structure of the environment, providing a mechanism to support object perception in dynamic natural vision. TeaserObjects are mentally rotated together with the changing viewpoint on a scene, affecting their representation in visual cortex.

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The neural time course of size constancy in natural scenes

Accurate real-world size perception relies on size constancy, a mechanism that integrates an objects retinal size with distance information. The neural time course of extracting pictorial distance cues from scenes and integrating them with retinal size information - a process referred to as scene-based size constancy - remains unknown. In two experiments, participants viewed objects with either large or small retinal sizes, presented at near or far distances in outdoor scene photographs, while performing an unrelated one-back task. We applied multivariate pattern analysis (MVPA) to time-resolved EEG data to decode the retinal size of large versus small objects, depending on their distance (near versus far) in the scenes. The objects were either perceptually similar in size (large-near versus small-far) or perceptually dissimilar in size (large-far versus small-near), reflecting size constancy. We found that the retinal size of objects could be decoded from 80 ms after scene onset onwards. Distance information modulated size decoding at least 120 ms later: from 200 ms after scene onset when objects were fixated, and from 280 ms when objects were viewed in the periphery. These findings reveal the neural time course of size constancy based on pictorial distance cues in natural scenes.

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Numerosity adaptation suppresses early visual responses

Humans and many animals rapidly and accurately perceive numerosity, the number of objects, in a visual image. The numerosity of recently viewed images influences our perception of the current images numerosity: numerosity adaptation. How does numerosity adaptation affect responses to numerosity in the brain? Recent studies show both early visual responses that monotonically increase with numerosity, and later numerosity-tuned responses that peak at different (preferred) numerosities in different neural populations. We have recently shown that numerosity adaptation affects the preferred numerosity of numerosity-tuned neural populations. We have also shown that early visual monotonic responses reflect image contrast, which follows numerosity closely. Here we ask how monotonic responses in the early visual cortex are affected by adaptation to different numerosities, using ultra-high field (7T) fMRI and neural model-based analyses. FMRI response amplitudes increased monotonically with numerosity throughout the early visual field maps (V1-V3, hV4, LO1-LO2 & V3A/B). This increase in response amplitudes becomes less steep after adaptation to higher numerosities, with this effect becoming stronger through the early visual hierarchy. This suppression of responses to numerosity is consistent with perceptual effects where adaptation to high numerosities reduces the perceived numerosity. These results imply that numerosity adaptation effects in later numerosity-tuned neural populations may originate in early visual areas that respond to image contrast in the adapting image.

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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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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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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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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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Preparatory attention incorporates contextual expectations

Humans are remarkably proficient at finding objects within a complex visual world. Current theories of attentional selection propose that this ability is mediated by target-specific preparatory activity in visual cortex, biasing visual processing in favor of the target object. In real-world situations, however, the retinal image that any object will produce is unknown in advance; its size, for instance, varies dramatically with the objects distance from the observer. Using fMRI, we show that preparatory activity is systematically modulated by expectations derived from scene context. Human participants searched for objects at different distances in scenes. Activity patterns in object-selective cortex during search preparation (while no objects were presented), resembled activity patterns evoked by viewing targets object in isolation. Crucially, this preparatory activity was modulated by distance, reflecting the predicted retinal image of the object at each distance. These findings reconcile current theories of attentional selection with the challenges of real-world vision. HighlightsO_LIVisual cortex contains object-specific representations during search preparation. C_LIO_LIWe demonstrate this for the first time during concurrent visual scene processing. C_LIO_LIPreparatory object representations are scaled to account for viewing distance. C_LIO_LIPreparatory biases reflect the predicted retinal image inferred from scene context. C_LI eTOC blurbAttentional selection is thought to be mediated by target-specific preparatory activity in visual cortex. Gayet and Peelen provide evidence that such preparatory biases incorporate contextual expectations about object appearance, reconciling attention theories with the challenges of naturalistic vision.

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