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Knapen, T.

Publications and source records attributed to Knapen, T..

At least 19 recordsLinked to original sources

Attraction of population receptive fields towards the attended locus is invariant to contrast

Attention enhances perception and spatial resolution. One mechanism by which attention achieves this is by attracting receptive fields towards the attended locus. The literature remains unclear on whether stimulus-driven parameters like contrast should alter this attraction. On one hand, Bayesian interpretation predicts changes in attraction due to contrast, whereas attention field models do not. Here we investigate whether stimulus contrast alters attention-driven attraction towards the attended locus. We used a demanding attentional task at fixation (0.1{degrees}) while mapping population receptive fields (pRFs) using either a low-contrast (5% Michelson contrast) or a high-contrast (80% Michelson contrast) checkerboard bar. Behavioral performance across conditions was matched. We show large and consistent differences in the amplitude of responses, but surprisingly, no difference in the amount of attraction towards the attended locus. The variance explained in this experiment was comparable to that of similar studies, which did observe pRF property changes, suggesting sufficient sensitivity to detect attraction towards the attended locus had it occurred at a similar magnitude. Our results cannot be explained by the Bayesian interpretation that predicts attention-based attraction effects varying with contrast. Instead, attention field models better account for our observations.

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The precision of attention controls attraction of population receptive fields

We alter our sampling of visual space not only by where we direct our gaze but also by where and how we direct our attention. Attention attracts receptive fields toward the attended position, but our understanding of this process is limited. Here we show that the degree of this attraction towards the attended locus is dictated not just by the attended position, but also by the precision of attention. We manipulated attentional precision while using 7T fMRI to measure population receptive field (pRF) properties. Participants performed the same color-proportion detection task either focused at fixation (0.1{degrees} radius) or distributed across the entire display (more than 5{degrees} radius). We observed BOLD response amplitude increases as a function of the task, with selective increases in foveal pRFs for the focused attention task and vice versa for the distributed attention task. Furthermore, cortical spatial tuning changed as a function of attentional precision. Specifically, focused attention more strongly attracted pRFs towards the attended locus compared to distributed attention. This attraction also depended on the degree of overlap between a pRF and the attention field. A Gaussian attention field model with an offset on the attention field explained our results. Together, our observations indicate the spatial distribution of attention dictates the degree of its resampling of visual space.

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Non-invasive mapping of the temporal processing hierarchy in the human visual cortex

Vision is not instantaneous but evolves over time. However, simultaneously capturing the fine spatial details and the rapid temporal dynamics of visual processing remains a major challenge, resulting in a gap in our understanding of spatiotemporal dynamics. Here, we introduce a forward modeling technique that bridges high-spatial resolution fMRI with high-temporal resolution MEG, enabling us to non-invasively measure different levels of the visual hierarchy in humans and their involvement in visual processing with millisecond precision. Using fMRI, levels of the visual hierarchy were identified by measuring individuals population receptive fields and determining visual field maps. We predicted how much the activity patterns in each visual field map would contribute to brain responses measured with MEG. By comparing these predicted responses with the measured MEG responses, we assessed how much a given visual field map contributed to the measured MEG response, and, most importantly, when. We combined information from all MEG sensors and revealed a cortical processing hierarchy across visual field maps. We validated the method using cross-validations and demonstrated that the model generalized across MEG sensor types, stimulus shapes, and was robust to the number of visual field maps included in the model. We found that the primary visual cortex captured most of the variance in the MEG sensors and did so earlier in time than extrastriate regions. We also report a processing hierarchy across extra-striate visual field maps and clusters. We effectively combined the advantages of two very different neuroimaging techniques, opening avenues for answering research questions that require recordings with high spatiotemporal detail. By bridging traditionally separate areas of research, our approach helps close longstanding gaps in our understanding of brain function. Author SummaryVision doesnt happen instantaneously, but unfolds over time. While we understand a lot about how the brain processes visual space, understanding how the brain processes information over both space and time is much more challenging. Vision happens incredibly fast, space and time are tightly linked, but current technology is limited in capturing these spatiotemporal dynamics. Here, we developed a method that combines two non-invasive human brain imaging techniques using computational models: fMRI, which provides high spatial detail, and MEG, which captures millisecond-level timing. We use fMRI to model the detailed layout of the visual system, and predict how this spatial layout responds to specific visual stimuli and compare these predictions with MEG recordings. This allows us to pinpoint not just where visual processing happens, but also when. Our results reveal a clear processing hierarchy: primary visual cortex responds first and explains most of the MEG signal, while higher-level visual areas activate later. We show that our method works across different stimuli, MEG sensor types, and brain areas, proving its robustness. This approach offers a new way to track how information flows through the living human brain, opening up new possibilities to study vision in both health and disease.

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Decoding peripheral saccade targets from foveal retinotopic cortex

Human vision is characterized by frequent eye movements and constant shifts in visual input, yet our perception of the world remains remarkably stable. Here, we directly demonstrate image-specific foveal feedback to primary visual cortex in the context of saccadic eye movements. To this end, we used a gaze-contingent fMRI paradigm, in which peripheral saccade targets disappeared before they could be fixated. Despite no direct foveal stimulation, we were able to decode peripheral saccade targets from foveal retinotopic areas, demonstrating that image-specific feedback during saccade preparation may underlie this effect. Decoding was sensitive to shape but not semantic category of natural images, indicating feedback of only low-to-mid-level information. Cross-decoding to a control condition with foveal stimulus presentation indicates a shared representational format between foveal feedback and direct stimulation. Moreover, eccentricity-dependent analyses showed a u-shaped decoding curve, confirming that these results are not explained by spillover of peripheral activity or large receptive fields. Finally, fluctuations in foveal decodability covaried with activity in the intraparietal sulcus, thus providing a candidate region for driving foveal feedback. These findings suggest that foveal cortex predicts the features of incoming stimuli through feedback from higher cortical areas, which offers a candidate mechanism underlying stable perception.

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Psilocybin alters visual-contextual computations

Psilocybin alters perception and brain dynamics. Contextual computations are ubiquitous in the brain. Here, we investigate the effects of psilocybin using psychophysics, ultra-high field functional MRI, and computational modeling. We find that 1) psilocybin alters contextual perception in the Ebbinghaus illusion, 2) psilocybin alters contextual modulation in cortical responses to visual stimuli, and 3) we propose a computational model capable of capturing and linking these changes. Leveraging vision as a beachhead, our findings highlight the alteration of contextual computations as a potential general mechanism underlying psychedelic action. TeaserPsilocybin alters visual-contextual computations, a potential general computational mechanism for psychedelic effects in the human brain.

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Vicarious Somatotopic Maps Tile Visual Cortex

Our sensory systems work together to generate a cohesive experience of the world around us. Watching others being touched activates brain areas representing our own sense of touch: the visual system recruits touch-related computations to simulate bodily consequences of visual inputs. One long-standing question is how the brain implements this interface between visual and somatosensory representations. To address this question, we developed a method to simultaneously map somatosensory body part tuning and visual field tuning throughout the brain. Applying this method on ongoing co-activations during rest resulted in detailed maps of the body-part tuning in the brains endogenous somatotopic network. During movie watching, somatotopic tuning explains responses throughout the entire dorsolateral visual system, revealing an array of somatotopic body maps that tile the cortical surface. The tuning of these maps aligned with those of visual maps, and predicted both preferences for visual field locations and the visual-category preferences for body parts. These results reveal a mode of brain organization in which aligned visual-somatosensory topographic maps connect visual and bodily reference frames. This cross-modal interface is ideally situated to translate raw sensory impressions into more abstract formats useful for action, social cognition, and semantic processing.

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Hierarchical cortical plasticity in congenital sight impairment

A robust learning system balances adaptability to new experiences with stability of its foundational architecture. To investigate how the human brain implements this we used a new approach to study plasticity and stability across hierarchical processing stages in visual cortex. We compare the rod system of individuals born with rod-only photoreceptor inputs (achromatopsia) to the typically developed rod system, allowing us to dissociate impacts of life-long versus transient responses to altered input. Cortical input stages (V1) exhibited high stability, with input-deprived cortex showing no retinotopic remapping and exhibiting structural hallmarks of deprivation. However, plasticity manifested as reorganised read-out of these inputs by higher-order cortex, in a pattern that could compensate for the lower resolution of a rod-only system and its lack of high-density foveal input. We propose that these hierarchical dynamics robustly optimize processing of available input and could reflect a broader principle of brain organisation with important implications for emerging sight-rescue therapies.

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A retinotopic reference frame for space throughout human visual cortex

We perceive the world as stable despite our rapid eye movements. To explain our sense of visual stability, it has been suggested that the brain encodes the location of attended visual stimuli in an external, or spatiotopic, reference frame. However, such spatiotopy is seemingly at odds with the fundamental retinotopic organization of visual inputs. Here, we probe the spatial reference frame of vision using ultra-high-field (7T) fMRI and voxel-level receptive field modeling, while manipulating both gaze direction and spatial attention. To manipulate spatial attention, participants performed an equally demanding visual task on either a bar stimulus that traversed the visual field, or a small stimulus at fixation. To dissociate retinal stimulus position from its real-world position the entire stimulus array was placed at one of three distinct horizontal screen positions in each run. We found that population receptive fields in all cortical visual field maps are pinioned to the retina, irrespective of how spatial attention is deployed. This pattern of results is strong evidence for a fully retinotopic reference frame for visual-spatial processing. Reasoning that a spatiotopic reference frame could independently be computed at the population level of entire visual areas rather than in individual voxels, we additionally used Bayesian decoding of stimulus location from the BOLD response patterns in visual areas. We found that decoded stimulus locations also adhere to the retinotopic frame of reference. Again, this result holds for all visual areas and irrespective of the deployment of spatial attention. Our findings reorient the search for visual stability mechanisms toward transient sensorimotor interactions rather than static spatiotopic maps.

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Hierarchical cortical and subcortical mechanisms underlying binocular rivalry

Conscious perception alternates between the two eyes images during binocular rivalry. How hierarchical processes in our brain interact to resolve visual competition to generate conscious perception remains unclear. Here we investigated the mesoscale neural circuitry for binocular rivalry in human cortical and subcortical areas using high-resolution functional MRI at 7 Tesla. Eye-specific response modulation in binocular rivalry was strongest in the superficial layers of V1 ocular dominance columns (ODCs), and more synchronized in the superficial and deep layers. The intraparietal sulcus (IPS) generated stronger eye-specific response modulation and increased effective connectivity to the early visual cortex during binocular rivalry compared to monocular "replay" simulations. Although there was no evidence of eye-specific rivalry modulation in the lateral geniculate nucleus (LGN) of the thalamus, strong perceptual rivalry modulation can be found in its parvocellular (P) subdivision. Finally, IPS and ventral pulvinar showed robust perceptual rivalry modulation and increased connectivity to the early visual cortex. These findings demonstrate that local interocular competition arises from lateral mutual inhibition between V1 ODCs, and feedback signals from IPS to visual cortex and visual thalamus further synchronize and resolve visual competition to generate conscious perception. Graphical abstract O_FIG O_LINKSMALLFIG WIDTH=200 HEIGHT=166 SRC="FIGDIR/small/528110v2_ufig1.gif" ALT="Figure 1"> View larger version (37K): org.highwire.dtl.DTLVardef@af6feaorg.highwire.dtl.DTLVardef@1dc8cd3org.highwire.dtl.DTLVardef@a1905forg.highwire.dtl.DTLVardef@10d7b75_HPS_FORMAT_FIGEXP M_FIG C_FIG HighlightsO_LIEye-specific rivalry modulation is strongest in the superficial layers of V1 ODCs and more synchronized in superficial and deep layers C_LIO_LIIPS generates stronger eye-specific response modulation and increases connectivity to V1 during rivalry compared to replay C_LIO_LILGN activity shows no evidence of eye-specific rivalry modulation but strong perceptual rivalry modulation in its P subdivision C_LIO_LIIPS and ventral pulvinar show robust perceptual rivalry modulation and increased connectivity to the early visual cortex C_LI

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Naturalistic Audiovisual Stimulation Reveals the Topographic Organization of Human Auditory Cortex

Despite the importance of audition in spatial, semantic, and social function, there is no consensus regarding the detailed organisation of human auditory cortex. Using a novel application of a population receptive field model to a high-powered naturalistic audiovisual movie-watching dataset, we simultaneously estimate the basic spectral tuning properties and category selectivity of human auditory cortex. This revealed unprecedentedly clear tonotopic maps which showcase the modes of organization and computational motifs of the auditory cortex. Specifically, we find that regions more remote from the auditory core exhibit more compressive, non-linear response properties with finely-tuned, speech selectivity in low frequency portions of their tonotopic maps. These patterns of organisation mirror aspects of the visual cortical hierarchy, wherein tuning properties progress from a stimulus category-agnostic front end towards more advanced regions increasingly optimised for behaviorally relevant stimulus categories.

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Separable pupillary signatures of perception and action during perceptual multistability

The pupil provides a rich, non-invasive measure of the neural bases of perception and cognition, and has been of particular value in uncovering the role of arousal-linked neuromodulation, which alters cortical processing as well as pupil size. But pupil size is subject to a multitude of influences, which complicates unique interpretation. We measured pupils of observers experiencing perceptual multistability -- an ever-changing subjective percept in the face of unchanging but inconclusive sensory input. In separate conditions the endogenously generated perceptual changes were either task-relevant or not, allowing a separation between perception-related and task-related pupil signals. Perceptual changes were marked by a complex pupil response that could be decomposed into two components: a dilation tied to task execution and plausibly indicative of an arousal-linked noradrenaline surge, and an overlapping constriction tied to the perceptual transient and plausibly a marker of altered visual cortical representation. Constriction, but not dilation, amplitude systematically depended on the time interval between perceptual changes, possibly providing an overt index of neural adaptation. These results show that the pupil provides a simultaneous reading on interacting but dissociable neural processes during perceptual multistability, and suggest that arousal-linked neuromodulation shapes action but not perception in these circumstances.

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Topographic connectivity reveals task-dependent retinotopic processing throughout the human brain.

The human visual system is organized as a hierarchy of maps that share the topography of the retina. These retinotopic maps have been identified throughout the brain, but how much of the brain is visually organized remains unknown. Here we demonstrate widespread stable visual organization beyond the traditional visual system. We analyzed detailed topographic connectivity with primary visual cortex during moviewatching, rest, and retinotopic mapping experiments to reveal that visual-spatial representations are warped by experimental condition and cognitive state. Specifically, traditionally visual regions alternate with default mode network and hippocampus in preferentially representing the center of the visual field. This visual role of hippocampus would allow it to implement sensory predictions by interfacing between abstract memories and concrete perceptions. These results indicate that pervasive sensory coding facilitates the communication between far-flung brain regions.

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Ultra-high resolution fMRI reveals origins of feedforward and feedback activity within laminae of human ocular dominance columns

Ultra-high field MRI can functionally image the cerebral cortex of human subjects at the submillimeter scale of cortical columns and laminae. Here, we investigate both in concert, by, for the first time, imaging ocular dominance columns (ODCs) in primary visual cortex (V1) across different cortical depths. We ensured that putative ODC patterns in V1 (a) are stable across runs, sessions, and scanners located in different continents (b) have a width ([~]1.3 mm) expected from post-mortem and animal work and (c) are absent at the retinotopic location of the blind spot. We then dissociated the effects of bottom-up thalamo-cortical input and attentional feedback processes on activity in V1 across cortical depth. Importantly, the separation of bottom-up information flows into ODCs allowed us to validly compare attentional conditions while keeping the stimulus identical throughout the experiment. We find that, when correcting for draining vein effects and using both model-based and model-free approaches, the effect of monocular stimulation is largest at deep and middle cortical depths. Conversely, spatial attention influences BOLD activity exclusively near the pial surface. Our findings show that simultaneous interrogation of columnar and laminar dimensions of the cortical fold can dissociate thalamocortical inputs from top-down processing, and allow the investigation of their interactions without any stimulus manipulation. Significance StatementThe advent of ultra-high field fMRI allows for the study of the human brain non-invasively at submillimeter resolution, bringing the scale of cortical columns and laminae into focus. De Hollander et al imaged the ocular dominance columns and laminae of V1 in concert, while manipulating top-down attention. This allowed them to separate feedforward from feedback processes in the brain itself, without resorting to the manipulation of incoming information. Their results show how feedforward and feedback processes interact in the primary visual cortex, highlighting the different computational roles separate laminae play.

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Learning in visual regions as support for the bias in future value-driven choice

Reinforcement learning can bias decision-making towards the option with the highest expected outcome. Cognitive learning theories associate this bias with the constant tracking of stimulus values and the evaluation of choice outcomes in the striatum and prefrontal cortex. Decisions however first require processing of sensory input, and to-date, we know far less about the interplay between learning and perception. This fMRI study (N=43), relates visual BOLD responses to value-beliefs during choice, and, signed prediction errors after outcomes. To understand these relationships, which co-occurred in the striatum, we sought relevance by evaluating the prediction of future value-based decisions in a separate transfer phase where learning was already established. We decoded choice outcomes with a 70% accuracy with a supervised machine learning algorithm that was given trial-by-trial BOLD from visual regions alongside more traditional motor, prefrontal, and striatal regions. Importantly, this decoding of future value-driven choice outcomes again highligted an important role for visual activity. These results raise the intriguing possibility that the tracking of value in visual cortex is supportive for the striatal bias towards the more valued option in future choice.

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Retinotopic maps of visual space in the human cerebellum

While the cerebellum is instrumental for motor control, it is not traditionally implicated in vision. Here, we report the existence of 5 ipsilateral visual field maps in the human cerebellum. These maps are located within the oculomotor vermis and cerebellar nodes of the dorsal attention and visual networks. These findings imply that the cerebellum is closely involved in visuospatial cognition, and that its contributions are anchored in sensory coordinates.

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Dopaminergic medication reduces striatal sensitivity to negative outcomes in Parkinson’s disease

Reduced levels of dopamine in Parkinsons disease (PD) contribute to changes in learning, resulting from the loss of midbrain dopamine neurons that transmit a teaching signal to the striatum. Dopamine medication used by PD patients has previously been linked to either behavioral changes during learning itself or adjustments in approach and avoidance behavior after learning. To date, however, very little is known about the specific relationship between dopaminergic medication-driven differences during learning and subsequent changes in approach/avoidance tendencies in individual patients. We assessed 24 PD patients on and off dopaminergic medication and 24 healthy controls (HC) performing a probabilistic reinforcement learning task, while undergoing functional magnetic resonance imaging. During learning, medication in PD reduced an overemphasis on negative outcomes. When patients were on medication, learning rates were lower for negative (but not positive) outcomes and concurrent striatal BOLD responses showed reduced prediction error sensitivity. Medication-induced shifts in negative learning rates were predictive of changes in approach/avoidance choice patterns after learning, and these changes were accompanied by striatal BOLD response alterations. These findings highlight dopamine-driven learning differences in PD and provide new insight into how changes in learning impact the transfer of learned value to approach/avoidance responses in novel contexts.

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Implicit and explicit learning in reactive and voluntary saccade adaptation

Saccades can either be elicited automatically by salient peripheral stimuli or can additionally depend on explicit cognitive goals. Similarly, it is thought that motor adaptation is driven by the combination of a more automatic, implicit process and a more explicit, cognitive process. However, the degree to which such implicit and explicit learning contribute to the adaptation of more reactive and voluntary saccades remains elusive. To study this question, we employed a global saccadic adaptation paradigm with both increasing and decreasing saccade amplitudes. We assessed the resulting adaptation using a dual state model of motor adaptation. This model decomposes learning into a fast and slow process, which are thought to constitute the explicit and implicit learning, respectively. Our results show that adaptation of reactive saccades is equally driven by fast and slow learning, while fast learning is nearly absent when adapting voluntary (i.e. scanning) saccades. This pattern of results was present both when saccade gain was increased or decreased. Our results suggest that the increased cognitive demands associated with voluntary compared to reactive saccade planning interfere specifically with explicit learning.

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No evidence that frontal eye field tDCS affects latency or accuracy of prosaccades

Transcranial direct current stimulation (tDCS) may be used to directly affect neural activity from outside of the skull. However, its exact physiological mechanisms remain elusive, particularly when applied to new brain areas. The frontal eye field (FEF) has rarely been targeted with tDCS, even though it plays a crucial role in control of overt and covert spatial attention. Here we investigate whether tDCS over the FEF can affect the latency and accuracy of saccadic eye movements. 26 participants performed a prosaccade task in which they made eye movements to a sudden-onset eccentric visual target (lateral saccades). After each lateral saccade, they made an eye movement back to the center (center saccades). The task was administered before, during and after anodal or cathodal tDCS over the FEF, in a randomized, double-blind, within-subject design. One previous study (Kanai et al., 2012) found that anodal tDCS over the FEF decreased the latency of saccades contralateral to the stimulated hemisphere. We did not find the same effect: neither anodal nor cathodal tDCS influenced the latency of lateral saccades. tDCS also did not affect accuracy of lateral saccades (saccade endpoint deviation and saccade endpoint variability). For center saccades, we found some differences between the anodal and cathodal sessions, but these were not consistent across analyses (latency, endpoint variability), or were already present before tDCS onset (endpoint deviation). We tried to improve on the design of Kanai et al. (2012) in several ways, including the tDCS duration and electrode montage, which could explain the discrepant results. Our findings add to a growing number of null results, which have sparked concerns that tDCS outcomes are highly variable. Future studies should aim to establish the boundary conditions for frontal eye field tDCS to be effective, in addition to increasing sample size and adding additional controls such as a sham condition. At present, we conclude that it is unclear whether eye movements or other aspects of spatial attention can be affected through tDCS of the frontal eye fields.

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