Microsaccade Rate Varies with Subjective Visibility during Motion-Induced Blindness

Motion-induced blindness (MIB) occurs when a dot embedded in a motion field subjectively vanishes. Here we report the first psychophysical data concerning effects of microsaccade/eyeblink rate upon perceptual switches during MIB. We find that the rate of microsaccades/eyeblink rises before and after perceptual transitions from not seeing to seeing the dot, and decreases before perceptual transitions from seeing it to not seeing it. In addition, event-related fMRI data reveal that, when a dot subjectively reappears during MIB, the blood oxygen-level dependent (BOLD) signal increases in V1v and V2v and decreases in contralateral hMT+. These BOLD signal changes observed upon perceptual state changes in MIB could be driven by the change of perceptual states and/or a confounding factor, such as the microsaccade/eyeblink rate

Seuratun kappaleen poikkeuttaminen silmänräpäysten aikana: käyttäytymis- ja neuromagneettisia havaintoja

The visual world is perceived as continuous despite frequent interruptions of sensory data due to eyeblinks and rapid eye movements. To create the perception of constancy, the brain makes use of fill-in mechanisms. This study presents an experiment in which the location of an object during smooth pursuit tracking is altered during eyeblinks. The experiment investigates the effects of blink suppression and fill-in mechanisms to cloud the discrimination of these changes. We employed a motion-tracking task, which promotes the accurate evaluation of the object’s trajectory and thus can counteract the fill-in mechanisms. Six subjects took part in the experiment, during which they were asked to report any perceived anomalies in the trajectory. Eye movements were monitored with a video-based tracking and brain responses with simultaneous MEG recordings. Discrimination success was found to depend on the direction of the displacement, and was significantly modulated by prior knowledge of the triggered effect. Eye-movement data were congruent with previous findings and revealed a smooth transition from blink recovery to object locating. MEG recordings were analysed for condition-dependent evoked and induced responses; however, intersubject variability was too large for drawing clear conclusions regarding the brain basis of the fill-in mechanisms.Visuaalinen maailma koetaan jatkuvana, vaikka silmänräpäykset ja nopeat silmänliikkeet aiheuttavat keskeytyksiä sensoriseen tiedonkeruuseen. Luodakseen käsityksen pysyvyydestä, aivot käyttävät täyttömekanismeja. Tämä tutkimus esittelee kokeen, jossa kappaleen seurantaa hitailla seurantaliikkeillä häiritään muuttamalla sen sijaintia silmänräpäysten aikana. Tämä koe tutkii, kuinka silmänräpäysten aiheuttama suppressio ja täyttömekanismit sumentavat kykyä erotella näitä muutoksia. Käytimme liikeseurantatehtävää, joka vastaavasti edistää kappaleen liikeradan tarkkaa arviointia. Kuusi koehenkilöä osallistui kokeeseen, jonka aikana heitä pyydettiin ilmoittamaan kaikki havaitut poikkeamat kappaleen liikeradassa. Silmänliikkeitä tallennettiin videopohjaisella seurannalla, ja aivovasteita yhtäaikaisella MEG:llä. Erottelykyvyn todettiin riippuvan poikkeutuksen suunnasta, sekä merkittävästi a priori tiedosta poikkeutusten esiintymistavasta. Silmänliikedata oli yhtenevää aiempien tutkimusten kanssa, ja paljasti sujuvan siirtymisen silmänräpäyksistä palautumisesta kappaleen paikallistamiseen. MEG-tallenteet analysoitiin ehdollisten heräte- ja indusoitujen vasteiden löytämiseksi, mutta yksilölliset vaste-erot koehenkilöiden välillä olivat liian suuria selkeiden johtopäätösten tekemiseksi täyttömekanismien aivoperustasta

Activity in the human superior colliculus relating to endogenous saccade preparation and execution

In recent years a small number of studies have applied functional imaging techniques to investigate visual responses in the human superior colliculus (SC), but few have investigated its oculomotor functions. Here, in two experiments, we examined activity associated with endogenous saccade preparation. We used 3-T fMRI to record the hemodynamic activity in the SC while participants were either preparing or executing saccadic eye movements. Our results showed that not only executing a saccade (as previously shown) but also preparing a saccade produced an increase in the SC hemodynamic activity. The saccade-related activity was observed in the contralateral and to a lesser extent the ipsilateral SC. A second experiment further examined the contralateral mapping of saccade-related activity with a larger range of saccade amplitudes. Increased activity was again observed in both the contralateral and ipsilateral SC that was evident for large as well as small saccades. This suggests that the ipsilateral component of the increase in BOLD is not due simply to small-amplitude saccades producing bilateral activity in the foveal fixation zone. These studies provide the first evidence of presaccadic preparatory activity in the human SC and reveal that fMRI can detect activity consistent with that of buildup neurons found in the deeper layers of the SC in studies of nonhuman primates

The interaction between human vision and eye movements in health and disease

Human motor behaviour depends on the successful integration of vision and eye movements. Many studies have investigated neural correlates of visual processing in humans, but typically with the eyes stationary and fixated centrally. Similarly, many studies have sought to characterise which brain areas are responsible for oculomotor control, but generally in the absence of visual stimulation. The few studies to explicitly study the interaction between visual perception and eye movements suggest strong influences of both static and dynamic eye position on visual processing and modulation of oculomotor structures by properties of visual stimuli. However, the neural mechanisms underlying these interactions are poorly understood. This thesis uses a range of fMRI methodologies such as retinotopic mapping, multivariate analsyis techniques, dynamic causal modelling and ultra high resolution imaging to examine the interactions between the oculomotor and visual systems in the normal human brain. The results of the experiments presented in this thesis demonstrate that oculomotor behaviour has complex effects on activity in visual areas, while spatial properites of visual stimuli modify activity in oculomotor areas. Specifically, responses in the lateral geniculate nucleus and early cortical visual areas are modulated by saccadic eye movements (a process potentially mediated by the frontal eye fields) and by changes in static eye position. Additionally, responses in oculomotor structures such as the superior colliculus are biased for visual stimuli presented in the temporal rather than nasal hemifield. These findings reveal that although the visual and oculomotor systems are spatially segregated in the brain, they show a high degree of integration at the neural level. This is consistent with our everyday experience of the visual world where frequent eye movements do not lead to disruption of visual continuity and visual information is seamlessly transformed into motor behaviour

지각판단 중 인간 1차시각피질의 역할 규명을 위한 뇌영상연구

학위논문 (박사)-- 서울대학교 대학원 : 뇌인지과학과, 2015. 8. 이상훈.Primary visual cortex (V1) forms the initial cortical representation of objects and events in our visual environment, and it distributes information about that representation to higher cortical areas within the visual hierarchy. Decades of work have established tight linkages between neural activity occurring in V1 and features comprising the retinal image, but it remains debatable how that activity relates to perceptual decisions. An actively debated question is the extent to which V1 responses determine, on a trial-by-trial basis, perceptual choices made by observers. By inspecting the population activity of V1 from human observers engaged in a difficult visual discrimination task, we tested one essential prediction of the deterministic view: choice-related activity, if it exists in V1, and stimulus-related activity should occur in the same neural ensemble of neurons at the same time. Our findings do not support this prediction: while cortical activity signifying the variability in choice behavior was indeed found in V1, that activity was dissociated from activity representing stimulus differences relevant to the task, being advanced in time and carried by a different neural ensemble. Moreover, realizing that small deviations in fixational eye movements could affect our fMRI measurements, we tested and confirmed that this pattern of results cannot be attributed to fixational eye movements. The spatiotemporal dynamics of population responses suggest that short-term priors, perhaps formed in higher cortical areas involved in perceptual inference, act to modulate V1 activity prior to stimulus onset without modifying subsequent activity that actually represents stimulus features within V1.1. INTRODUCTION 1 2. EXPERIMENT 1: NEURAL SIGNATURES OF STIMULUS AND CHOICE ARE DISSOCIATED IN POPULATION ACTIVITY OF HUMAN V1 DURING PERCEPTUAL DECISION-MAKING 4 2.1. FINE RING-SIZE CLASSIFICATION TASK 4 2.2. DEFINITION OF ECCENTRICITY-TUNING FOR INDIVIDUAL VOXELS 8 2.3. DEFINITION OF TRIAL-RELATED MATRICES OF POPULATION RESPONSES 11 2.4. NEITHER STIMULI NOR CHOICES SIGNIFICANTLY CORRELATED WITH RAW RESPONSES 13 2.5. DISSOCIATED SIGNATURES OF STIMULUS AND CHOICE IN TUNED RESPONSES 18 2.6. DECODING STIMULUS AND CHOICE INFORMATION FROM RAW RESPONSES WITH POPULATION READ-OUT WEIGHTS 23 3. EXPERIMENT 2: PUPIL SIZE DYNAMICS DURING FIXATION IMPACT THE ACCURACY AND PRECISION OF VIDEO-BASED GAZE ESTIMATION 29 3.1. SPURIOUS EYE MOVEMENT SIGNALS IN VIDEO-BASED GAZE ESTIMATION 30 3.2. PUPIL SIZE VARIATION AND GAZE POSITION ESTIMATES DURING FIXATION 32 3.3. CORRECTION OF GAZE POSITION MEASUREMENTS FOR PUPIL SIZE ARTIFACT 37 3.4. COMPARISON BETWEEN PUPIL CENTER ESTIMATION METHODS 44 4. EXPERIMENT 3: CHOICE SIGNATURES IN V1 CANNOT BE ATTRIBUTED TO FIXATIONAL EYE MOVEMENTS 47 4.1. CORRECTION OF GAZE POSITION MEASUREMENTS DURING THE FINE RING-SIZE CLASSIFICATION TASK FOR PUPIL SIZE ARTIFACT 49 4.2. LACK OF CHOICE SIGNATURE IN FIXATIONAL EYE MOVEMENTS 54 5. DISCUSSIONS 59 5.1. THE ROLE OF V1 IN PERCEPTUAL DECISION-MAKING 59 5.2. POTENTIAL ORIGIN OF THE CHOICE SIGNATURE IN V1 64 5.3. PUPIL SIZE DYNAMICS DURING FIXATION 67 5.4. PUPIL SIZE ARTIFACT IN VIDEO-BASED GAZE ESTIMATION 70 6. EXPERIMENTAL PROCEDURES 74 6.1. OBSERVERS 74 6.2. FMRI EXPERIMENTAL SETUP 74 6.3. FMRI BEHAVIORAL PROTOCOL (EXPERIMENT 1) 75 6.4. ACQUISITION AND PREPROCESSING OF MRI DATA 77 6.5. ECCENTRICITY-TUNING MAPPING IN V1 VOXELS 79 6.6. DEFINITION OF ECCENTRICITY BINS 83 6.7. COMPUTATION OF STIMULUS AND CHOICE PROBABILITIES 84 6.8. PREDICTION OF V1 POPULATION RESPONSES TO RING STIMULI 87 6.9. DECOMPOSITION OF FMRI POPULATION RESPONSES INTO TUNED AND UNTUNED COMPONENTS 88 6.10. POPULATION DECODING OF STIMULUS AND CHOICE INFORMATION 89 6.11. EYE-TRACKING EXPERIMENTAL SETUP 90 6.12. EYE-TRACKING BEHAVIORAL PROTOCOL (EXPERIMENT 2 AND 3) 93 6.13. EYE-TRACKING DATA PREPROCESSING 94 6.14. CORRECTION OF GAZE POSITION MEASUREMENTS FOR PUPIL SIZE ARTIFACT 97 6.15. ACCURACY AND PRECISION ANALYSIS OF GAZE POSITION SIGNAL 98 6.16. STATISTICAL POWER ANALYSIS OF GAZE POSITION SIGNAL 100 6.17. MICROSACCADES ANALYSIS 101 6.18. PUPIL SIZE ANALYSIS 102 6.19. ANALYSIS OF GAZE POSITION AND VERGENCE ANGLE 103 BIBLIOGRAPHY 104 국문초록 116Docto

Investigating the relationship between microsaccades and oscillations in the human visual cortex

Neural oscillations play important roles in vision and attention. Most studies of oscillations use visual fixation to control the visual input. Small eye movements, called microsaccades, occur involuntarily ~ 1-2 times per second during fixation and they are also thought to play important roles in vision and attention. The aim of the work described in this thesis was to explore the relationship between microsaccades and oscillations in the human visual cortex. In Chapter 2, I describe how remote video eye tracking can be used to detect and characterize microsaccades during MEG recordings. Tracking based on the pupil position only, without corneal reflection, and with the participant’s head immobilized in the MEG dewar, resulted in high precision gaze tracking and enabled the following investigations. In Chapter 3, I investigated the relationship between induced visual gamma oscillations and microsaccades in a simple visual stimulation paradigm. I did not find evidence for the relationship. This finding supports the view that sustained gamma oscillations reflect local processing in cortical columns. In addition, early transient gamma response had a reduced amplitude on trials with microsaccades, however the exact nature of this effect will have to be determined in future studies. In Chapter 4, I investigated the relationship between alpha oscillations and microsaccades in covert spatial attention. I did not find evidence for a relationship between hemispheric lateralization of the alpha amplitude and the directional bias of microsaccades. I propose that microsaccades and alpha oscillations represent two independent attentional mechanisms - the former related to early attention shifting and the latter to maintaining sustained attention. In Chapter 5, I recorded, for the first time, microsaccade-related spectral responses. Immediately after their onset, microsaccades increased amplitude in theta and beta bands and this effect was modulated by stimulus type. Moreover, microsaccades reduced alpha amplitude ~ 0.3 s after their onset and this effect was independent of stimulus type. These results have important implications for the interpretation of the classical oscillatory effects in the visual cortex as well as for the role of microsaccades in vision and attention

Naturalistic viewing conditions can increase task engagement and aesthetic preference but have only minimal impact on EEG quality

Free gaze and moving images are typically avoided in EEG experiments due to the expected generation of artifacts and noise. Yet for a growing number of research questions, loosening these rigorous restrictions would be beneficial. Among these is research on visual aesthetic experiences, which often involve open-ended exploration of highly variable stimuli. Here we systematically compare the effect of conservative vs. more liberal experimental settings on various measures of behavior, brain activity and physiology in an aesthetic rating task. Our primary aim was to assess EEG signal quality. 43 participants either maintained fixation or were allowed to gaze freely, and viewed either static images or dynamic (video) stimuli consisting of dance performances or nature scenes. A passive auditory background task (auditory steady-state response; ASSR) was added as a proxy measure for overall EEG recording quality. We recorded EEG, ECG and eye tracking data, and participants rated their aesthetic preference and state of boredom on each trial. Whereas both behavioral ratings and gaze behavior were affected by task and stimulus manipulations, EEG SNR was barely affected and generally robust across all conditions, despite only minimal preprocessing and no trial rejection. In particular, we show that using video stimuli does not necessarily result in lower EEG quality and can, on the contrary, significantly reduce eye movements while increasing both the participants’ aesthetic response and general task engagement. We see these as encouraging results indicating that — at least in the lab — more liberal experimental conditions can be adopted without significant loss of signal quality