Attentional modulation of oscillatory activity in human visual cortex

The effects of attentional modulation on activity within the human visual cortex were investigated using magnetoencephalography. Chromatic sinusoidal stimuli were used to evoke activity from the occipital cortex, with attention directed either toward or away from the stimulus using a bar-orientation judgment task. For five observers, global magnetic field power was plotted as a function of time from stimulus onset. The major peak of each function occurred at about 120 ms latency and was well modeled by a current dipole near the calcarine sulcus. Independent component analysis (ICA) on the non-averaged data for each observer also revealed one component of calcarine origin, the location of which matched that of the dipolar source determined from the averaged data. For two observers, ICA revealed a second component near the parieto-occipital sulcus. Although no effects of attention were evident using standard averaging procedures, time-varying spectral analyses of single trials revealed that the main effect of attention was to alter the level of oscillatory activity. Most notably, a sustained increase in alpha-band (7-12 Hz) activity of both calcarine and parieto-occipital origin was evident. In addition, calcarine activity in the range of 13-21 Hz was enhanced, while calcarine activity in the range of 5-6 Hz was reduced. Our results are consistent with the hypothesis that attentional modulation affects neural processing within the calcarine and parieto-occipital cortex by altering the amplitude of alpha-band activity and other natural brain rhythms. © 2003 Elsevier Inc. All rights reserved.

Colour and luminance interactions in the visual perception of motion

We sought to determine the extent to which red–green, colour–opponent mechanisms in the human visual system play a role in the perception of drifting luminance–modulated targets. Contrast sensitivity for the directional discrimination of drifting luminance–modulated (yellow–black) test sinusoids was measured following adaptation to isoluminant red–green sinusoids drifting in either the same or opposite direction. When the test and adapt stimuli drifted in the same direction, large sensitivity losses were evident at all test temporal frequencies employed (1–16 Hz). The magnitude of the loss was independent of temporal frequency. When adapt and test stimuli drifted in opposing directions, large sensitivity losses were evident at lower temporal frequencies (1–4 Hz) and declined with increasing temporal frequency. Control studies showed that this temporal–frequency–dependent effect could not reflect the activity of achromatic units. Our results provide evidence that chromatic mechanisms contribute to the perception of luminance–modulated motion targets drifting at speeds of up to at least 32°s-1. We argue that such mechanisms most probably lie within a parvocellular–dominated cortical visual pathway, sensitive to both chromatic and luminance modulation, but only weakly selective for the direction of stimulus motion.

Distinct contrast response functions in striate and extra-striate regions of visual cortex revealed with magnetoencephalography (MEG)

Objective: To spatially and temporally characterise the cortical contrast response function to pattern onset stimuli in humans. Methods: Magnetoencephalography (MEG) was used to investigate the human cortical contrast response function to pattern onset stimuli with high temporal and spatial resolution. A beamformer source reconstruction approach was used to spatially localise and identify the time courses of activity at various visual cortical loci. Results: Consistent with the findings of previous studies, MEG beamformer analysis revealed two simultaneous generators of the pattern onset evoked response. These generators arose from anatomically discrete locations in striate and extra-striate visual cortex. Furthermore, these loci demonstrated notably distinct contrast response functions, with striate cortex increasing approximately linearly with contrast, whilst extra-striate visual cortex followed a saturating function. Conclusions: The generators that underlie the pattern onset visual evoked response arise from two distinct regions in striate and extra-striate visual cortex. Significance: The spatially, temporally and functionally distinct mechanisms of contrast processing within the visual cortex may account for the disparate results observed across earlier studies and assist in elucidating causal mechanisms of aberrant contrast processing in neurological disorders. © 2005 International Federation of Clinical Neurophysiology. Published by Elsevier Ireland Ltd. All rights reserved.

Are poor mathematics skills associated with visual deficits in temporal processing?

Developmental learning disabilities such as dyslexia and dyscalculia have a high rate of co-occurrence in pediatric populations, suggesting that they share underlying cognitive and neurophysiological mechanisms. Dyslexia and other developmental disorders with a strong heritable component have been associated with reduced sensitivity to coherent motion stimuli, an index of visual temporal processing on a millisecond time-scale. Here we examined whether deficits in sensitivity to visual motion are evident in children who have poor mathematics skills relative to other children of the same age. We obtained psychophysical thresholds for visual coherent motion and a control task from two groups of children who differed in their performance on a test of mathematics achievement. Children with math skills in the lowest 10% in their cohort were less sensitive than age-matched controls to coherent motion, but they had statistically equivalent thresholds to controls on a coherent form control measure. Children with mathematics difficulties therefore tend to present a similar pattern of visual processing deficit to those that have been reported previously in other developmental disorders. We speculate that reduced sensitivity to temporally defined stimuli such as coherent motion represents a common processing deficit apparent across a range of commonly co-occurring developmental disorders.

Topographic mapping and source localization of the pattern onset visual evoked magnetic response

The topography of the visual evoked magnetic response (VEMR) to a pattern onset stimulus was investigated using 4 check sizes and 3 contrast levels. The pattern onset response consists of three early components within the first 200ms, CIm, CIIm and CIIIm. The CIIm is usually of high amplitude and is very consistent in latency within a subject. Half field (HF) stimuli produce their strongest response over the contralateral hemisphere; the RHF stimulus exhibiting a lower positivity (outgoing field) and an upper negativity (ingoing field), rotated towards the midline. LHF stimulation produced the opposite response, a lower negative and an upper positive. Larger check sizes produce a single area of ingoing and outgoing field while smaller checks produce on area of ingoing and outgoing field over each hemisphere. Latency did not appear to vary with change in contrast but amplitudes increased with increasing contrast. A more detailed topographic study incorporating source localisation procedures suggested a source for CIIm - 4cm below the scalp, close to the midline with current flowing towards the lateral surface. Similar depth and position estimates but with opposite polarity were obtained for the pattern shift P100m previously. Hence, the P100m and the CIIm may originate in similar areas of visual cortex but reveal different aspects of visual processing. © 1992 Human Sciences Press, Inc.

The influence of age on the pattern and flash visual evoked magnetic response (VEMR)

The visual evoked magnetic response (VEMR) was measured over the occipital cortex to pattern and flash stimuli in 86 normal subjects aged 15-86 years. The latency of the major positive component (outgoing magnetic field) to the pattern reversal stimulus (P100M) increased with age, particularly after 55 years, while the amplitude of the P100M decreased more gradually over the lifespan. By contrast, the latency of the major positive component to the flash stimulus (P2M) increased more slowly with age after about 50 years, while its amplitude may have decreased in only a proportion of the elderly subjects. The changes in the P100M with age may reflect senile changes in the eye and optic nerve, e.g. senile miosis, degenerative changes in the retina or geniculostriate deficits. The P2M may be more susceptible to senile changes in the visual cortex. The data suggest that the contrast channels of visual information processing deteriorate more rapidly with age than the luminance channels.

Normative MEG data to flash and pattern reversal stimuli

We have attempted to establish normative values of components of the magnetic evoked field to flash and pattern reversal stimuli prior to clinical use of the MEG. Full visual field, binocular evoked magnetic fields were recorded from 100 subjects 16 to 86 years of age with a single channel dc Squid (BTI) second-order gradiometer at a point 5-6cm above the inion. The majority of subjects showed a large positive component (out going magnetic field) of mean latency 115 ms (SD range 2.5 -11.8 in different decades of life) to the pattern reversal stimulus. In many subjects, this P100M was preceeded and succeeded by negative deflections (in going field). About 6% of subjects showed an inverted response i.e. a PNP wave. Waveforms to flash were more variable in shape with several positive components; the most consistent having a mean latency of 110ms (SD range 6.4-23.2). Responses to both stimuli were consistent when measured on the same subject on six different occasions (SD range 4.8 to 7.3). The data suggest that norms can be established for evoked magnetic field components, in particular for the pattern reversal P100M, which could be used in the diagnosis of neuro-opthalmological disease.

The influence of age on the pattern and flash visual evoked field

We have investigated the effect of ageing on the visual system using the relatively new technique of magentoencephalography (MEG). This technique measures the magnetic signals produced by the visual system using a SQUID magnetometer. The magnetic visual evoked field (VEF) was measured over the occipital cortex to pattern and flash stimuli in 86 normal subjects aged 15 - 86 years. Factors that influenced subject defocussing or defixating the stimulus or selective attention were controlled as far as possible. The latency of the major positive component to the pattern reversal stimulus (P100M) increased with age particularly after the age of 55 years while the amplitude of the P100M decreased over the life span. The latency of the major flash component (P2M) increased much more slowly with age, while its amplitude decreased in only a proportion of elderly subjects. Changes in the P100M with age may reflect senile changes in the eye and optic nerve, e.g. senile miosis or degenerative changes in the retina. The P2M may be more susceptible to senile changes in the retina. The data suggest that the spatial frequency channels deteriorate more rapidly with age than the luminance channels and that MEG may be an effective method of studying ageing in the visual system.

Topography of the visual evoked magnetic response and hemispherical specialization

The visual evoked magnetic response CIIm component to a pattern onset stimulus presented half field produced a consistent scalp topography in 15 normal subjects. The major response was seen over the contralateral hemisphere, suggesting a dipole with current flowing away from the medial surface of the brain. Full field responses were more unpredictable. The reponses of five subjects were studied to the onset of a full, left half and right half checkerboard stimuli of 38 x 27 min arc checks appearing for 200 ms. In two subjects the full field CIIm topography was consistent with that of the mathematical summation of their relevant half field distribution. The remaining subjects had unpredictable full field topographies, showing little or no relationship to their half or summated half fields. In each of these subjects, a distribution matching that of the summated half field CIIm distribution appears at an earlier latency than that of the predominant full field waveform peak. By examining the topography of the full and half field responses at 5 ms intervals along the waveform for one such subject, the CIIm topography of the right hemisphere develops 10 ms before that of the left hemisphere, and is replaced by the following CIIIm component 20 ms earlier. Hence, the large peak seen in full field results from a combination of the CIIm component of the left hemisphere plus that of the CIIIm from the right. The earlier peak results from the CIIm generated in both hemispheres, at a latency where both show similar amplitudes. As the relative amplitudes of these two peaks alter with check and field size, topographic studies would be required for accurate CIIm identification. In addition. the CIIm-CIIIm complex lasts for 80 ms in the right hemisphere and 135 ms in the left, suggesting hemispherical apecialization in the visual processing of the pattern onset response.

Magnetoencephalography:technical improvements in image co-registration and studies of visual cortical oscillations

The work presented in this thesis is divided into two distinct sections. In the first, the functional neuroimaging technique of Magnetoencephalography (MEG) is described and a new technique is introduced for accurate combination of MEG and MRI co-ordinate systems. In the second part of this thesis, MEG and the analysis technique of SAM are used to investigate responses of the visual system in the context of functional specialisation within the visual cortex. In chapter one, the sources of MEG signals are described, followed by a brief description of the necessary instrumentation for accurate MEG recordings. This chapter is concluded by introducing the forward and inverse problems of MEG, techniques to solve the inverse problem, and a comparison of MEG with other neuroimaging techniques. Chapter two provides an important contribution to the field of research with MEG. Firstly, it is described how MEG and MRI co-ordinate systems are combined for localisation and visualisation of activated brain regions. A previously used co-registration methods is then described, and a new technique is introduced. In a series of experiments, it is demonstrated that using fixed fiducial points provides a considerable improvement in the accuracy and reliability of co-registration. Chapter three introduces the visual system starting from the retina and ending with the higher visual rates. The functions of the magnocellular and the parvocellular pathways are described and it is shown how the parallel visual pathways remain segregated throughout the visual system. The structural and functional organisation of the visual cortex is then described. Chapter four presents strong evidence in favour of the link between conscious experience and synchronised brain activity. The spatiotemporal responses of the visual cortex are measured in response to specific gratings. It is shown that stimuli that induce visual discomfort and visual illusions share their physical properties with those that induce highly synchronised gamma frequency oscillations in the primary visual cortex. Finally chapter five is concerned with localization of colour in the visual cortex. In this first ever use of Synthetic Aperture Magnetometry to investigate colour processing in the visual cortex, it is shown that in response to isoluminant chromatic gratings, the highest magnitude of cortical activity arise from area V2.

Behavioural and neuroimaging investigations of the relationship between visual attention, affordance and action

In this thesis the relationship between visual attention, affordance and action was investigated using a combination of neuroimaging and behavioural studies. Neuronal activity and movement construction were assessed when individuals passively viewed or produced action towards stimuli varying in their affordance and/or attentional attributes. The main findings were: (i) the passive perception of both object and abstract visual patterns was associated with decreased alpha and/or beta activity in sensori-motor cortex, occipito-temporal cortex and cerebellum. These are brain regions associated with the planning and production of visually guided action; (ii) for object patterns, decreased alpha and beta activity was also observed in regions of superior parietal and premotor cortex. These regions contain neurons argued to be essential for matching hand kinematics with manipulate objects; and (iii) in both control participants and a deafferented individual, studies of planned and unplanned pointing manoeuvres revealed that the attentional bias of a stimulus was critical for fast, efficient action production whereas the affordance bias was critical in determining end-point accuracy. Taken together, these findings demonstrate that affordance is not a necessary prerequisite for the potential of motor codes. Rather, affordance enables the construction of motor responses that reflect object functionality and/or manipulability. They further demonstrate that visual attention is associated with the potentiation of motor codes. Indeed, directed visual attention would appear critical for speeded responses. These findings provide new insights into the roles of directed visual attention and affordance upon action.

A comparison between static and kinetic visual attention as a means of detecting age-related deterioration of the visual system and driving performance

A critical review of previous research revealed that visual attention tests, such as the Useful Field of View (UFOV) test, provided the best means of detecting age-related changes to the visual system that could potentially increase crash risk. However, the question was raised as to whether the UFOV, which was regarded as a static visual attention test, could be improved by inclusion of kinetic targets that more closely represent the driving task. A computer program was written to provide more information about the derivation of UFOV test scores. Although this investigation succeeded in providing new information, some of the commercially protected UFOV test procedures still remain unknown. Two kinetic visual attention tests (DRTS1 and 2), developed at Aston University to investigate inclusion of kinetic targets in visual attention tests, were introduced. The UFOV was found to be more repeatable than either of the kinetic visual attention tests and learning effects or age did not influence these findings. Determinants of static and kinetic visual attention were explored. Increasing target eccentricity led to reduced performance on the UFOV and DRTS1 tests. The DRTS2 was not affected by eccentricity but this may have been due to the style of presentation of its targets. This might also have explained why only the DRTS2 showed laterality effects (i.e. better performance to targets presented on the left hand side of the road). Radial location, explored using the UFOV test, showed that subjects responded best to targets positioned to the horizontal meridian. Distraction had opposite effects on static and kinetic visual attention. While UFOV test performance declined with distraction, DRTS1 performance increased. Previous research had shown that this striking difference was to be expected. Whereas the detection of static targets is attenuated in the presence of distracting stimuli, distracting stimuli that move in a structured flow field enhances the detection of moving targets. Subjects reacted more slowly to kinetic compared to static targets, longitudinal motion compared to angular motion and to increased self-motion. However, the effects of longitudinal motion, angular motion, self-motion and even target eccentricity were caused by target edge speed variations arising because of optic flow field effects. The UFOV test was more able to detect age-related changes to the visual system than were either of the kinetic visual attention tests. The driving samples investigated were too limited to draw firm conclusions. Nevertheless, the results presented showed that neither the DRTS2 nor the UFOV tests were powerful tools for the identification of drivers prone to crashes or poor driving performance.

Visual performance of the human eye with soft hydrophilic contact lenses

The recording of visual acuity using the Snellen letter chart is only a limited measure of the visual performance of an eye wearing a refractive aid. Qualitative in addition to quantitative information is required to establish such a parameter: spatial, temporal and photometric aspects must all be incorporated into the test procedure. The literature relating to the correction of ametropia by refractive aids was reviewed. Selected aspects of a comparison between the correction provided by spectacles and contact lenses were considered. Special attention was directed to soft hydrophilic contact lenses. Despite technological advances which have produced physiologically acceptable soft lenses, there still remain associated with this recent form of refractive aid unpredictable visual factors. Several techniques for vision assessment were described, and previous studies of visual performance were discussed. To facilitate the investigation of visual performance in a clinical environment, a new semi-automated system was described: this utilized the presentation of broken ring test stimuli on a television screen. The research project comprised two stages. Initial work was concerned with the validation of the television system, including the optimization of its several operational variables. The second phase involved the utilization of the system in an investigation of visual performance aspects of the first month of regular daily soft contact lens wear by experimentally-naive subjects. On the basis of the results of this work an ‘homoeostatic’ model has been proposed to represent the strategy which an observer adopts in order to optimize his visual performance with soft contact lenses.

Automated assessment of visual fields and their inter-relation to evoked potentials in visual disorders

The Octopus Automated Perimeter was validated in a comparative study and found to offer many advantages in the assessment of the visual field. The visual evoked potential was investigated in an extensive study using a variety of stimulus parameters to simulate hemianopia and central visual field defects. The scalp topography was recorded topographically and a technique to compute the source derivation of the scalp potential was developed. This enabled clarification of the expected scalp distribution to half field stimulation using different electrode montages. The visual evoked potential following full field stimulation was found to be asymmetrical around the midline with a bias over the left occiput particularly when the foveal polar projections of the occipital cortex were preferentially stimulated. The half field response reflected the distribution asymmetry. Masking of the central 3° resulted in a response which was approximately symmetrical around the midline but there was no evidence of the PNP-complex. A method for visual field quantification was developed based on the neural representation of visual space (Drasdo and Peaston 1982) in an attempt to relate visual field depravation with the resultant visual evoked potentials. There was no form of simple, diffuse summation between the scalp potential and the cortical generators. It was, however, possible to quantify the degree of scalp potential attenuation for M-scaled full field stimuli. The results obtained from patients exhibiting pre-chiasmal lesions suggested that the PNP-complex is not scotomatous in nature but confirmed that it is most likely to be related to specific diseases (Harding and Crews 1982). There was a strong correlation between the percentage information loss of the visual field and the diagnostic value of the visual evoked potential in patients exhibiting chiasmal lesions.

The development of calibration and adaptive filtering procedures for neuromagnetic studies of human visual function

This thesis first considers the calibration and signal processing requirements of a neuromagnetometer for the measurement of human visual function. Gradiometer calibration using straight wire grids is examined and optimal grid configurations determined, given realistic constructional tolerances. Simulations show that for gradiometer balance of 1:104 and wire spacing error of 0.25mm the achievable calibration accuracy of gain is 0.3%, of position is 0.3mm and of orientation is 0.6°. Practical results with a 19-channel 2nd-order gradiometer based system exceed this performance. The real-time application of adaptive reference noise cancellation filtering to running-average evoked response data is examined. In the steady state, the filter can be assumed to be driven by a non-stationary step input arising at epoch boundaries. Based on empirical measures of this driving step an optimal progression for the filter time constant is proposed which improves upon fixed time constant filter performance. The incorporation of the time-derivatives of the reference channels was found to improve the performance of the adaptive filtering algorithm by 15-20% for unaveraged data, falling to 5% with averaging. The thesis concludes with a neuromagnetic investigation of evoked cortical responses to chromatic and luminance grating stimuli. The global magnetic field power of evoked responses to the onset of sinusoidal gratings was shown to have distinct chromatic and luminance sensitive components. Analysis of the results, using a single equivalent current dipole model, shows that these components arise from activity within two distinct cortical locations. Co-registration of the resulting current source localisations with MRI shows a chromatically responsive area lying along the midline within the calcarine fissure, possibly extending onto the lingual and cuneal gyri. It is postulated that this area is the human homologue of the primate cortical area V4.