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Tsuchiya, N.

Publications and source records attributed to Tsuchiya, N..

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Expectation and attention increase the integration of top-down and bottom-up signals in perception through different pathways

Perception results from the integration of incoming sensory information with pre-existing information available in the brain. In this EEG (electroencephalography) study we utilised the Hierarchical Frequency Tagging method to examine how such integration is modulated by expectation and attention. Using intermodulation (IM) components as a measure of non-linear signal integration, we show in three different experiments that both expectation and attention enhance integration between top-down and bottom-up signals. Based on multispectral phase coherence, we present two direct physiological measures to demonstrate the distinct yet related mechanisms of expectation and attention. Specifically, our results link expectation to the modulation of prediction signals and the integration of top-down and bottom-up information at lower levels of the visual hierarchy. Meanwhile, they link attention to the propagation of ascending signals and the integration of information at higher levels of the visual hierarchy. These results are consistent with the predictive coding account of perception.

neuroscience

Attention periodically samples competing stimuli during binocular rivalry

The attentional sampling hypothesis suggests that attention rhythmically enhances sensory processing when attending to a single (~8 Hz), or multiple (~4 Hz) objects. Here we investigated using binocular rivalry whether attention samples sensory representations that are not part of the conscious percept, during competition for perceptual dominance. When crossmodally cued toward a conscious image, subsequent changes in consciousness occurred at ~8 Hz, consistent with rates of undivided attentional sampling. However, when attention was cued toward the suppressed image, changes in consciousness slowed to ~3.5 Hz, indicating the division of attention away from the conscious visual image. In the electroencephalogram, we found that at 3.5 and 8 Hz, the strength of inter-trial phase coherence over fronto-temporal and parieto-occipital regions correlated with behavioral measures of changes in perception. When cues were not task-relevant, these effects disappeared, confirming that perceptual changes were dependent upon the allocation of attention, and that attention can flexibly sample away from a conscious image in a task-dependent manner.

neuroscience

Deep Brain Stimulation for Parkinson’s disease changes perception in the Rubber Hand Illusion

Parkinsons disease (PD) alters cortico-basal ganglia-thalamic circuitry and susceptibility to an illusion of bodily awareness, the Rubber Hand Illusion (RHI). Bodily awareness is thought to result from multisensory integration in a predominantly cortical network; the role of subcortical connections is unknown. We studied the effect of modulating cortico-subcortical circuitry on multisensory integration for bodily awareness in PD patients treated with subthalamic nucleus (STN) deep brain stimulation (DBS) using the RHI experiment. Typically, synchronous visuo-tactile cues induce a false perception of touch on the rubber hand as if it were the subjects hand, whereas asynchronous visuo-tactile cues do not. However, we found that in the asynchronous condition, patients in the off-stimulation state did not reject the RHI as strongly as healthy controls; switching on STN-DBS partially normalised their responses. Patients in the off-stimulation state also misjudged the position of their hand, indicating it to be closer to the rubber hand than controls. However, STN-DBS did not affect proprioceptive judgements or subsequent arm movements altered by the perceptual effects of the illusion. Our findings support the idea that the STN and subcortical connections have a key role in multisensory integration for bodily awareness. Decisionmaking in multisensory bodily illusions is discussed.

neuroscience

A unified framework for dissecting the effects of common signals on functional and effective connectivity analyses: power, coherence, and Granger causality

When analyzing neural data it is important to consider the limitations of the particular experimental setup. An enduring issue in the context of electrophysiology is the presence of common signals. For example a non-silent reference electrode adds a common signal across all recorded data and this adversely affects functional and effective connectivity analysis. To address the common signals problem, a number of methods have been proposed, but relatively few detailed investigations have been carried out. We address this gap by analyzing local field potentials recorded from the small brains of fruit flies. We conduct our analysis following a solid mathematical framework that allows us to make precise predictions regarding the nature of the common signals. We demonstrate how a framework that jointly analyzes power, coherence and quantities from the Granger causality framework allows us to detect and assess the nature of the common signals. Our analysis revealed substantial common signals in our data, in part due to a non-silent reference electrode. We further show that subtracting spatially adjacent signals (bipolar rereferencing) largely removes the effects of the common signals. However, in some special cases this operation itself introduces a common signal. The mathematical framework and analysis pipeline we present can readily be used by others to detect and assess the nature of the common signals in their data, thereby reducing the chance of misinterpreting the results of functional and effective connectivity analysis.

neuroscience

Isoflurane reduces feedback in the fruit fly brain

Hierarchically organized brains communicate through feedforward and feedback pathways. In mammals, feedforward and feedback are mediated by higher and lower frequencies during wakefulness. Feedback is preferentially impaired by general anesthetics. This suggests feedback serves critical functions in waking brains. The brain of Drosophila melanogaster (fruit fly) is also hierarchically organized, but the presence of feedback in these brains is not established. Here we studied feedback in the fruit fly brain, by simultaneously recording local field potentials (LFPs) from low-order peripheral structures and higher-order central structures. Directed connectivity analysis revealed that low frequencies (0.1-5Hz) mediated feedback from the center to the periphery, while higher frequencies (10-45Hz) mediated feedforward in the opposite direction. Further, isoflurane anesthesia preferentially reduced feedback. Our results imply that similar spectral characteristics of feedforward and feedback may be a signature of hierarchically organized brains and that general anesthetics may induce unresponsiveness by targeting the mechanisms that support feedback.

neuroscience

Hierarchical Frequency Tagging reveals neural markers of predictive coding under varying uncertainty

Understanding the integration of top-down and bottom-up signals is essential for the study of perception. Current accounts of predictive coding describe this in terms of interactions between state units encoding expectations or predictions, and error units encoding prediction error. However, direct neural evidence for such interactions has not been well established. To achieve this, we combined EEG methods that preferentially tag different levels in the visual hierarchy: Steady State Visual Evoked Potential (SSVEP at 10Hz, tracking bottom-up signals) and Semantic Wavelet-Induced Frequency Tagging (SWIFT at 1.3Hz tracking top-down signals). Importantly, we examined intermodulation components (IM, e.g., 11.3Hz) as a measure of integration between these signals. To examine the influence of expectation and predictions on the nature of such integration, we constructed 50-second movie streams and modulated expectation levels for upcoming stimuli by varying the proportion of images presented across trials. We found SWIFT, SSVEP and IM signals to differ in important ways. SSVEP was strongest over occipital electrodes and was not modified by certainty. Conversely, SWIFT signals were evident over temporo- and parieto-occipital areas and decreased as a function of increasing certainty levels. Finally, IMs were evident over occipital electrodes and increased as a function of certainty. These results link SSVEP, SWIFT and IM signals to sensory evidence, predictions, prediction errors and hypothesis-testing - the core elements of predictive coding. These findings provide neural evidence for the integration of top-down and bottom-up information in perception, opening new avenues to studying such interactions in perception while constraining neuronal models of predictive coding.\n\nSIGNIFICANCE STATEMENTThere is a growing understanding that both top-down and bottom-up signals underlie perception. But how do these signals interact? And how does this process depend on the signals probabilistic properties? Predictive coding theories of perception describe this in terms how well top-down predictions fit with bottom-up sensory input. Identifying neural markers for such signal integration is therefore essential for the study of perception and predictive coding theories in particular. The novel Hierarchical Frequency Tagging method simultaneously tags top-down and bottom-up signals in EEG recordings, while obtaining a measure for the level of integration between these signals. Our results suggest that top-down predictions indeed integrate with bottom-up signals in a manner that is modulated by the predictability of the sensory input.

neuroscience