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van de Ven, V.

Publications and source records attributed to van de Ven, V..

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Theta-Phase Targeted Encoding Gates Memory Sensitivity

Theta phase has been proposed to play an important role in memory. It may gate encoding and retrieval, such that certain phases are more favorable for memory processing than others, or become part of the memory representation, such that reinstating the encoding phase benefits retrieval. To test and distinguish these competing accounts, we used a closed-loop EEG system, allowing us to control the phase at which items were presented at encoding and retrieval. In an associative memory task, visual targets were presented at predefined phases of the ongoing mid-frontal theta rhythm, thereby pairing individual memory items with specific encoding phases. During retrieval, targets appeared either at the same phase as during encoding or at an alternative phase. We show that theta phase during encoding predicted subsequent memory sensitivity, while finding no evidence that retrieval phase nor the match between encoding and retrieval phases influenced performance. Therefore, our findings are consistent with a gating account, in which the rhythmic state at the moment of encoding influences the efficacy of memory formation.

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Stimulus dependent modulation of perceptual filling-in is predicted by the properties of early visual cortex

Filling-in occurs during the perceptual disappearance of a blank figure presented on a textured background. Current models of perceptual filling-in are based on a two-stage model where the figure boundary weakens after a period of adaptation, followed by the spreading of the background representation into the region representing the figure. This suggests a competition between figure boundary and background representations whereby filling-in is facilitated by a weaker boundary representation and a stronger background representation. Here, we test this interpretation, by using the oblique effect and surround-modulation suppression, which are functional properties of early visual cortex that modulate the expected strengths of the responses to the background texture and to the figure boundary. In a sample of N=58 participants, we found more filling-in with background textures of cardinal compared to oblique orientations (earlier onset time, with more and longer episodes of filling-in per trial), in line with a known, stronger neuronal response for cardinal than for oblique orientation in early visual cortex. We found more filling-in when the main axis of the rectangular figure was iso-oriented rather than cross-oriented with the background texture (more and longer episodes of filling-in per trial, but no change in onset time), in line with a lower response to oriented stimuli when surrounded by iso-oriented flankers compared to cross-oriented flankers. Overall, our results support the two-stage model and suggest the involvement of early visual cortical areas characterized by the oblique effect and orientation-tuned surround-suppression.

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Feedback to deep layers in human V1 during perceptual filling-in

Visual surface perception is a fundamental aspect of vision, yet its neural implementation remains poorly understood. Troxlers perceptual filling-in paradigm provides a tractable illusion for studying surface perception, in which a peripheral figure becomes perceptually assimilated into the surrounding background after a period of sustained fixation. Although neural correlates of this phenomenon have been reported in early visual cortex, the underlying mechanisms, particularly the contribution of feedback signaling, remain unresolved. Here we use ultra-high-field (7T) layer-fMRI to investigate perceptual filling-in in the human visual cortex. While experimentally controlling perceptual filling-in, we measured GE-BOLD responses in ten participants. Analyses across cortical depth in the independently localized figure representation in primary visual cortex (V1) revealed neural correlates of filling-in in deep cortical layers, which are associated with feedback input. These findings provide evidence that perceptual filling-in and visual surface perception in general are supported by feedback signals to early visual cortex.

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Neural correlates of implicit memory for reoccurring sound patterns

The human brain exhibits remarkable sensitivity to auditory patterns. Previous research has shown that beyond the detection of such patterns, humans form memories for them that are implicit, robust to interference, and persistent over time. Here, we examine the mechanisms underlying implicit memory formation for reoccurring regular tone patterns using UHF fMRI. Prior to scanning, implicit memory was induced through a standard behavioral task where participants detected regular patterns embedded in random tone sequences. Unbeknownst to participants, five regular patterns reoccurred sporadically (every 1.5 min) across trials. Consistent with previous reports, behavioral performance indicated a reaction time advantage to reoccurring patterns, confirming the formation of implicit memory traces. During passive listening in the scanner, regularity engaged the superior temporal gyrus, inferior frontal gyrus, insula, putamen, and hippocampus, a result we interpret as reflecting the higher perceptual precision of regular patterns. Reoccurrence interacted with regularity in the superior temporal gyrus and a frontal cluster, showing higher activity for reoccurring compared to novel regular patterns. The hippocampus showed sensitivity to reoccurrence in both regular and random patterns. Together, these results indicate that the neocortex may selectively amplify reoccurring inputs in line with their enhanced (memory driven) perceptual precision.

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Unveiling early visual cortical mechanisms in perceptual filling-in: a parametric study of eccentricity and movement dependence

Surface perception relies on interactions between boundary encoding and surface filling-in. In a perceptual filling-in paradigm, a blank figure becomes perceptually replaced by a textured background after prolonged fixation of a point away from the figure. Filling-in begins when neurons representing the figures boundary adapt, allowing background-related activity to spread into the figures retinotopic representation. Adaptation proceeds faster the better the boundary is stabilized in a neurons receptive field (RF). We hypothesized that moving the figure boundary beyond a neurons RF would reduce adaptation and hinder filling-in, with larger movements permitting progressively less filling-in. As RF size increases with eccentricity, we further hypothesized that greater eccentricities would require larger movements to interfere with filling-in. Our results confirmed both predictions. The reduction in filling-in duration with increasing motion range permitted estimating RF size at each eccentricity. The slope of a linear function relating RF size to eccentricity matched values reported in human fMRI studies of V1/V2, suggesting that boundary adaptation involves early visual areas. We also explored whether microsaccade amplitude affects filling-in, but found no supporting evidence. Thus, external figure motion and microsaccades may disrupt adaptation through different mechanisms. These findings provide new insights into neural adaptation processes preceding perceptual filling-in.

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Transcranial alternating current stimulation at theta frequency to the parietal cortex impairs associative, but not perceptual, memory encoding

Neural oscillations in the theta range (4-6 Hz) are thought to underlie associative memory function in the hippocampal-cortical network. While there is ample evidence supporting a role of theta oscillations in animal and human memory, most evidence is correlational. Non-invasive brain stimulation (NIBS) can be employed to modulate cortical oscillatory activity to influence brain activity, and possibly modulate deeper brain regions, such as hippocampus, through strong and reliable cortico-hippocampal functional connections. We applied high-definition transcranial alternating current stimulation (HD-tACS) at 6 Hz over left parietal cortex to modulate brain activity in the putative cortico-hippocampal network to influence associative memory encoding. After encoding and brain stimulation, participants completed an associative memory and a perceptual recognition task. Results showed that theta tACS significantly decreased associative memory performance but did not affect perceptual memory performance. These results show that parietal theta tACS modulates associative processing separately from perceptual processing, and further substantiate the hypothesis that theta oscillations are implicated in the cortico-hippocampal network and associative encoding.

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Hippocampus plays a role in speech feedback processing

There is increasing evidence that the hippocampus is involved in language production and verbal communication, although little is known about its possible role. According to one view, hippocampus contributes semantic memory to spoken language. Alternatively, hippocampus is involved in the processing the (mis)match between expected sensory consequences of speaking and the perceived speech feedback. In the current study, we re-analysed functional magnetic resonance (fMRI) data of two overt picture-naming studies to test whether hippocampus is involved in speech production and, if so, whether the results can distinguish between a "pure memory" versus an "expectation" account of hippocampal involvement. In both studies, participants overtly named pictures during scanning while hearing their own speech feedback unimpededly or impaired by a superimposed noise mask. Results showed decreased hippocampal activity when speech feedback was impaired, compared to when feedback was unimpeded. Further, we found increased functional coupling between auditory cortex and hippocampus during unimpeded speech feedback, compared to impaired feedback. Finally, we found significant functional coupling between a hippocampal/supplementary motor area (SMA) interaction term and auditory cortex, anterior cingulate cortex and cerebellum during overt picture naming, but not during listening to ones own pre-recorded voice. These findings indicate that hippocampus plays a role in speech production that is in accordance with an "expectation" view of hippocampal functioning.

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Hippocampal-striatal functional connectivity supports processing of temporal expectations from associative memory

The hippocampus and dorsal striatum are both associated with temporal processing, but they are thought to play distinct roles. The hippocampus has been reported to contribute to storing temporal structure of events in memory, whereas the striatum contributes to temporal motor preparation and reward anticipation. Here, we asked whether the striatum cooperates with the hippocampus in processing the temporal context of memorized visual associations. In our task, participants were trained to implicitly form temporal expectations for one of two possible time intervals associated to specific cue-target associations, and subsequently were scanned using 7T functional magnetic resonance imaging. During scanning, learned temporal expectations could be violated when the pairs were presented at either the learned or not-learned time intervals. When temporal expectations were not met during testing trials, activity in hippocampal subfields CA3/CA2 and CA1 decreased while right putamen activity increased, compared to when temporal expectations were met. Further, psycho-physiological interactions showed that functional connectivity between left CA1 and caudate, as well as between putamen and caudate, decreased when temporal expectations were not met. Our results indicate that the hippocampus and striatum cooperate to process implicit temporal expectation from mnemonic associations, with different but complementary contributions from caudate and putamen. Our findings provide further support for a hippocampal-striatal network in temporal associative processing.

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