Development and application of magnetoencephalographic methods in the investigation of the human visual system

This thesis is an exploration of the organisation and functioning of the human visual system using the non-invasive functional imaging modality magnetoencephalography (MEG). Chapters one and two provide an introduction to the ‘human visual system and magnetoencephalographic methodologies. These chapters subsequently describe the methods by which MEG can be used to measure neuronal activity from the visual cortex. Chapter three describes the development and implementation of novel analytical tools; including beamforming based analyses, spectrographic movies and an optimisation of group imaging methods. Chapter four focuses on the use of established and contemporary analytical tools in the investigation of visual function. This is initiated with an investigation of visually evoked and induced responses; covering visual evoked potentials (VEPs) and event related synchronisation/desynchronisation (ERS/ERD). Chapter five describes the employment of novel methods in the investigation of cortical contrast response and demonstrates distinct contrast response functions in striate and extra-striate regions of visual cortex. Chapter six use synthetic aperture magnetometry (SAM) to investigate the phenomena of visual cortical gamma oscillations in response to various visual stimuli; concluding that pattern is central to its generation and that it increases in amplitude linearly as a function of stimulus contrast, consistent with results from invasive electrode studies in the macaque monkey. Chapter seven describes the use of driven visual stimuli and tuned SAM methods in a pilot study of retinotopic mapping using MEG; finding that activity in the primary visual cortex can be distinguished in four quadrants and two eccentricities of the visual field. Chapter eight is a novel implementation of the SAM beamforming method in the investigation of a subject with migraine visual aura; the method reveals desynchronisation of the alpha and gamma frequency bands in occipital and temporal regions contralateral to observed visual abnormalities. The final chapter is a summary of main conclusions and suggested further work.

Neuro-electromagnetic imaging of the human visual cortex

Methods of solving the neuro-electromagnetic inverse problem are examined and developed, with specific reference to the human visual cortex. The anatomy, physiology and function of the human visual system are first reviewed. Mechanisms by which the visual cortex gives rise to external electric and magnetic fields are then discussed, and the forward problem is described mathematically for the case of an isotropic, piecewise homogeneous volume conductor, and then for an anisotropic, concentric, spherical volume conductor. Methods of solving the inverse problem are reviewed, before a new technique is presented. This technique combines prior anatomical information gained from stereotaxic studies, with a probabilistic distributed-source algorithm to yield accurate, realistic inverse solutions. The solution accuracy is enhanced by using both visual evoked electric and magnetic responses simultaneously. The numerical algorithm is then modified to perform equivalent current dipole fitting and minimum norm estimation, and these three techniques are implemented on a transputer array for fast computation. Due to the linear nature of the techniques, they can be executed on up to 22 transputers with close to linear speedup. The latter part of the thesis describes the application of the inverse methods to the analysis of visual evoked electric and magnetic responses. The CIIm peak of the pattern onset evoked magnetic response is deduced to be a product of current flowing away from the surface areas 17, 18 and 19, while the pattern reversal P100m response originates in the same areas, but from oppositely directed current. Cortical retinotopy is examined using sectorial stimuli, the CI and CIm ;peaks of the pattern onset electric and magnetic responses are found to originate from areas V1 and V2 simultaneously, and they therefore do not conform to a simple cruciform model of primary visual cortex.

Neuromagnetic investigations of functional organisation within human visual cortex

This thesis describes a series of experimental investigations into the functional organisation of human visual cortex using neuromagnetometry.This technique combines good spatial and temporal resolution enabling identification of the location and temporal response characteristics of cortical neurones within alert humans. To activate different neuronal populations and cortical areas a range of stimuli were used, the parameters of which were selected to match the known physiological properties of primate cortical neurones. In one series of experiments the evoked magnetic response was recorded to isoluminant red/green gratings. Co-registration of signal and magnetic resonance image data indicated a contribution to the response from visual areas V1, V2 and V4. To investigate the spatio-temporal characteristics of neurones within area V1 the evoked response was recorded for a range of stimulus spatial and temporal frequencies. The response to isoluminant red/green gratings was dominated by a major component which was found to have bandpass spatial frequency tuning with a peak at 1-2 cycles/degree, falling to the level of the noise at 6-8 cycles/degree. The temporal frequency tuning characteristics of the response showed bimodal sensitivity with peaks at 0-1Hz and 4Hz. In a further series of experiments the luminance evoked response was recorded to red/black, yellow/black and achromatic gratings and in all cases was found to be more complex than the isoluminant chromatic response, comprising up to three distinct components. The major response peak showed bandpass spatial frequency tuning characteristics, peaking at 6-8 cycles/degree, falling to the level of the noise at 12-16 cycles/degree. The results provide evidence to suggest that within area V1 the same neuronal population encodes both chromatic and luminance information and has spatial frequency tuning properties consistent with single-opponent cells. Furthermore, the results indicate that cells within area V1 encode chromatic motion information over a wide range of temporal frequencies with temporal response characteristics suggestive of the existence of a sub-population of cells sensitive to high temporal frequencies.

Investigating tiredness and ad complexity effects on visual processing of advertisements:an eye-tracking study

Although advertising is pervasive in our daily, it proves to be not necessarily efficient all the times due to bad conditions or bad contexts of reception. Indeed, the communication process might be jeopardized at its very last stage because of advertising exposure quality. However critical it may be, ad exposure quality is not very much examined by researchers or practitioners. In this paper, we investigate how tiredness combined with ad complexity might influence the way consumers extract and process ad elements. Investigating tiredness is useful because it is a common daily state experienced by everyone at various moments of the day. And although it might drastically alter ad reception, it has not been studied in advertising for the moment. In this regards, we observe eye movement patterns of consumers viewing simple or complex advertisements being tired or not. We surprisingly find that tired subjects viewing complex ads don’t adopt a lessening effort visual strategy. They rather use a resource demanding one. We assume that the Sustained Attention strategy occurring is a kind of adaptive strategy allowing to deal with an anticipated lack of resource.

Enhanced access to early visual processing of perceptual simultaneity in autism spectrum disorders

We compared judgements of the simultaneity or asynchrony of visual stimuli in individuals with autism spectrum disorders (ASD) and typically-developing controls using Magnetoencephalography (MEG). Two vertical bars were presented simultaneously or non-simultaneously with two different stimulus onset delays. Participants with ASD distinguished significantly better between real simultaneity (0 ms delay between two stimuli) and apparent simultaneity (17 ms delay between two stimuli) than controls. In line with the increased sensitivity, event-related MEG activity showed increased differential responses for simultaneity versus apparent simultaneity. The strongest evoked potentials, observed over occipital cortices at about 130 ms, were correlated with performance differences in the ASD group only. Superior access to early visual brain processes in ASD might underlie increased resolution of visual events in perception. © 2012 Springer Science+Business Media New York.

Detection of Glass pattern configurations by the primary visual cortex

The neurons in the primary visual cortex that respond to the orientation of visual stimuli were discovered in the late 1950s (Hubel, D.H. & Wiesel, T.N. 1959. J. Physiol. 148:574-591) but how they achieve this response is poorly understood. Recently, experiments have demonstrated that the visual cortex may use the image processing techniques of cross or auto-correlation to detect the streaks in random dot patterns (Barlow, H. & Berry, D.L. 2010. Proc. R. Soc. B. 278: 2069-2075). These experiments made use of sinusoidally modulated random dot patterns and of the so-called Glass patterns - where randomly positioned dot pairs are oriented in a parallel configuration (Glass, L. 1969. Nature. 223: 578-580). The image processing used by the visual cortex could be inferred from how the threshold of detection of these patterns in the presence of random noise varied as a function of the dot density in the patterns. In the present study, the detection thresholds have been measured for other types of patterns including circular, hyperbolic, spiral and radial Glass patterns and an indication of the type of image processing (cross or auto-correlation) by the visual cortex is presented. As a result, it is hoped that this study will contribute to an understanding of what David Marr called the ‘computational goal’ of the primary visual cortex (Marr, D. 1982. Vision: A Computational Investigation into the Human Representation and Processing of Visual Information. New York: Freeman.)

Perceptually-based Comparison of Image Similarity Metrics

The image comparison operation ??sessing how well one image matches another ??rms a critical component of many image analysis systems and models of human visual processing. Two norms used commonly for this purpose are L1 and L2, which are specific instances of the Minkowski metric. However, there is often not a principled reason for selecting one norm over the other. One way to address this problem is by examining whether one metric better captures the perceptual notion of image similarity than the other. With this goal, we examined perceptual preferences for images retrieved on the basis of the L1 versus the L2 norm. These images were either small fragments without recognizable content, or larger patterns with recognizable content created via vector quantization. In both conditions the subjects showed a consistent preference for images matched using the L1 metric. These results suggest that, in the domain of natural images of the kind we have used, the L1 metric may better capture human notions of image similarity.

Engineers and technicians : the effect of cognitive style and gender on visual information processing

The purpose of this project was to identify in a subject group of engineers and technicians (N = 62) a preferred mode of representation for facilitating correct recall of information from complex graphics. The modes of representation were black and white (b&w) block, b&w icon, color block, and color icon. The researcher's test instrument included twelve complex graphics (six b&w and six color - three per mode). Each graphics presentation was followed by two multiple-choice questions. Recall performance was better using b&w block mode graphics and color icon mode graphics. A standardized test, the Group Embedded Figures Test (GEFT) was used to identify a cognitive style preference (field dependence). Although engineers and technicians in the sample were strongly field-independent, they were not significantly more field-independent than the normative group in the Witkin, Oltman, Raskin, and Karp study (1971). Tests were also employed to look for any significant difference in cognitive style preference due to gender. None was found. Implications from the project results for the design of visuals and their use in technical training are discussed.

The quiet eye without a target: The primacy of visual information processing

Motor-performance-enhancing effects of long final fixations before movement initiation – a phenomenon called Quiet Eye (QE) – have repeatedly been demonstrated. Drawing on the information-processing framework, it is assumed that the QE supports information processing revealed by the close link between QE duration and task demands concerning, in particular, response selection and movement parameterisation. However, the question remains whether the suggested mechanism also holds for processes referring to stimulus identification. Thus, in a series of two experiments, performance in a targeting task was tested as a function of experimentally manipulated visual processing demands as well as experimentally manipulated QE durations. The results support the suggested link because a performance-enhancing QE effect was found under increased visual processing demands only: Whereas QE duration did not affect performance as long as positional information was preserved (Experiment 1), in the full vs. no target visibility comparison, QE efficiency turned out to depend on information processing time as soon as the interval falls below a certain threshold (Experiment 2). Thus, the results rather contradict alternative, e.g., posture-based explanations of QE effects and support the assumption that the crucial mechanism behind the QE phenomenon is rooted in the cognitive domain.

The representation of the ipsilateral visual field in human cerebral cortex

Previous studies of cortical retinotopy focused on influences from the contralateral visual field, because ascending inputs to cortex are known to be crossed. Here, functional magnetic resonance imaging was used to demonstrate and analyze an ipsilateral representation in human visual cortex. Moving stimuli, in a range of ipsilateral visual field locations, revealed activity: (i) along the vertical meridian in retinotopic (presumably lower-tier) areas; and (ii) in two large branches anterior to that, in presumptive higher-tier areas. One branch shares the anterior vertical meridian representation in human V3A, extending superiorly toward parietal cortex. The second branch runs antero-posteriorly along lateral visual cortex, overlying motion-selective area MT. Ipsilateral stimuli sparing the region around the vertical meridian representation also produced signal reductions (perhaps reflecting neural inhibition) in areas showing contralaterally driven retinotopy. Systematic sampling across a range of ipsilateral visual field extents revealed significant increases in ipsilateral activation in V3A and V4v, compared with immediately posterior areas V3 and VP. Finally, comparisons between ipsilateral stimuli of different types but equal retinotopic extent showed clear stimulus specificity, consistent with earlier suggestions of a functional segregation of motion vs. form processing in parietal vs. temporal cortex, respectively.

The effects of alcohol on the driver's visual information processing. Final report.

National Highway Traffic Safety Administration, Washington, D.C.

Investigations of attentional processing in parietal and occipital human cortical regions with magnetoencephalography

Attention defines our mental ability to select and respond to stimuli, internal or external, on the basis of behavioural goals in the presence of competing, behaviourally irrelevant, stimuli. The frontal and parietal cortices are generally agreed to be involved with attentional processing, in what is termed the 'fronto-parietal' network. The left parietal cortex has been seen as the site for temporal attentional processing, whereas the right parietal cortex has been seen as the site for spatial attentional processing. There is much debate about when the modulation of the primary visual cortex occurs, whether it is modulated in the feedforward sweep of processing or modulated by feedback projections from extrastriate and higher cortical areas. MEG and psychophysical measurements were used to look at spatially selective covert attention. Dual-task and cue-based paradigms were used. It was found that the posterior parietal cortex (PPC), in particular the SPL and IPL, was the main site of activation during these experiments, and that the left parietal lobe was activated more strongly than the right parietal lobe throughout. The levels of activation in both parietal and occipital areas were modulated in accordance with attentional demands. It is likely that spatially selective covert attention is dominated by the left parietal lobe, and that this takes the form of the proposed sensory-perceptual lateralization within the parietal lobes. Another form of lateralization is proposed, termed the motor-processing lateralization, the side of dominance being determined by handedness, being reversed in left- relative to right-handers. In terms of the modulation of the primary visual cortex, it was found that it is unlikely that V1 is modulated initially; rather the modulation takes the form of feedback from higher extrastriate and parietal areas. This fits with the idea of preattentive visual processing, a commonly accepted idea which, in itself, prevents the concept of initial modulation of V1.