Title:	Electrophysiological Signatures of Active Vision
Authors:	Carl, Christine
Thesis advisor:	Prof. Dr. Peter König
Thesis referee:	Prof. Dr. Andreas K. Engel
Abstract:	Active movements are a key feature of human behavior. Even when we do not move our limbs we almost never stop guiding our eyes. As a minimal but omnipresent form of behavior, fast eye movements, called saccades, sample the visual world and determine to a large extent what we perceive. Despite being an integral part of visual perception, prevalent research practice treats the human subject as a passive observer who fixates a spot on the screen and is not allowed to move. Yet, learning sensorimotor interactions by active exploration in order to predict future changes and guide actions seems to be a fundamental principle of neural organization. This results in neural patterns of active behavior that can be fundamentally different from the neural processes revealed in movement-restricted laboratory settings questioning the transferability of results from experimental paradigms demanding fixation to real-world free viewing behavior. In this thesis, we aim at studying the neural mechanisms underlying free viewing behavior. In order to assess the fast, flexible and possibly distributed neural dynamics of active vision, we established a procedure for studying eye movements in magnetoencephalography (MEG) and investigated oscillatory signatures associated with sensorimotor processes of eye movements and saccade target selection, two fundamental processes of active vision. Electroencephalography (EEG) and MEG can non-invasively measure fast neural dynamics and hence seem ideally suited for studying active vision in humans. However, artifacts related to eye movements confound both EEG and MEG signals, and a thorough handling of these artifacts is crucial for investigating neural activities during active movements. Mostly, cleaning of ocular artifacts has been performed for occasional eye movements and blinks in fixation paradigms in EEG. Less is known about the impact of ocular artifacts and especially the saccadic spike on MEG. As a first step to enable active vision studies in MEG, we investigated ocular artifacts and possible ways of their separation from neural signals in MEG. We show that the saccadic spike seriously distorts the spatial and spectral characteristics of the MEG signal (Study 2). We further tested if electrooculogram (EOG) based regression is feasible for corneo-retinal artifact removal (Study 1). Due to an often-raised concern, we addressed if EOG regression eliminates neural activity when applied for MEG. Our results do not indicate such susceptibility and we conclude that EOG regression for removing the corneo-retinal artifact in MEG is suitable. Based on insights from both studies, we established an artifact handling procedure including EOG regression and independent component analysis (ICA) to assess the neural dynamics of active vision. In Study 3, we investigated spectral signatures of neuronal activity across cortex underlying saccade preparation, execution and re-fixation in a delayed saccade task. During preparation and execution, we found a dichotomic signature of gamma power increases and beta power decreases in widespread cortical areas related to saccadic control, including fronto-parietal structures. Saccade direction specific signatures resided in hemisphere lateralized changes in low gamma and alpha power in posterior parietal cortex during preparation extending to extrastriate areas during re-fixation. Real-world behavior implies the constant need to flexibly select actions between competing behavioral alternatives depending on both sensory input and internal states. In order to assess internally motivated viewing behavior, we compared neuronal activity of externally cued saccades with saccades to freely chosen, equally valuable targets. We found gamma band specific power increases in fronto-parietal areas that are likely to reflect a fast transient process of action guidance for sensory-guided saccades and a sustained process for internally selecting between competing behavioral alternatives. The sustained signature of internal action selection suggests that a decision between spatially oriented movements is mediated within sensorimotor structures by neural competition between assemblies encoding parallel evolving movement plans. Since our observations support the assumption that a decision emerges through the distributed consensus of neural activities within effector specific areas rather than in a distinct decision module, they argue for the importance of studying mental processes within their ecologically valid and active context. This thesis shows the feasibility of studying neural mechanisms of active vision in MEG and provides important steps for studying neurophysiological correlates of free viewing in the future. The observed spectrally specific, distributed signatures highlight the importance of assessing fast oscillatory dynamics across the cortex for understanding neural mechanisms mediating real-world active behavior.
URL:	https://osnadocs.ub.uni-osnabrueck.de/handle/urn:nbn:de:gbv:700-2014042912431
Subject Keywords:	cortical saccade generation; action selection; saccadic spike potential; MEG; gamma band; EEG
Issue Date:	29-Apr-2014
Type of publication:	Dissertation oder Habilitation [doctoralThesis]
Appears in Collections:	FB08 - E-Dissertationen

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