Naokazu Goda

Hierarchical Bayesian estimation for MEG inverse problem.

Source current estimation from MEG measurement is an ill-posed problem that requires prior assumptions about brain activity and an efficient estimation algorithm. In this article, we propose a new hierarchical Bayesian method introducing a hierarchical prior that can effectively incorporate both structural and functional MRI data. In our method, the variance of the source current at each source location is considered an unknown parameter and estimated from the observed MEG data and prior information by using the Variational Bayesian method. The fMRI information can be imposed as prior information on the variance distribution rather than the variance itself so that it gives a soft constraint on the variance. A spatial smoothness constraint, that the neural activity within a few millimeter radius tends to be similar due to the neural connections, can also be implemented as a hierarchical prior. The proposed method provides a unified theory to deal with the following three situations: (1) MEG with no other data, (2) MEG with structural MRI data on cortical surfaces, and (3) MEG with both structural MRI and fMRI data. We investigated the performance of our method and conventional linear inverse methods under these three conditions. Simulation results indicate that our method has better accuracy and spatial resolution than the conventional linear inverse methods under all three conditions. It is also shown that accuracy of our method improves as MRI and fMRI information becomes available. Simulation results demonstrate that our method appropriately resolves the inverse problem even if fMRI data convey inaccurate information, while the Wiener filter method is seriously deteriorated by inaccurate fMRI information.

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.

Transformation from image-based to perceptual representation of materials along the human ventral visual pathway

Every object in the world has its own surface quality that is a reflection of the material from which the object is made. We can easily identify and categorize materials (wood, metal, fabric etc.) at a glance, and this ability enables us to decide how to interact appropriately with these objects. Little is known, however, about how materials are represented in the brain, or how that representation is related to material perception or the physical properties of material surface. By combining multivoxel pattern analysis of functional magnetic resonance imaging data with perceptual and image-based physical measures of material properties, we found that the way visual information about materials is coded gradually changes from an image-based representation in early visual areas to a perceptual representation in the ventral higher-order visual areas. We suggest that meaningful information about multimodal aspects of real-world materials reside in the ventral cortex around the fusiform gyrus, where it can be utilized for categorization of materials.

Estimation of the Timing of Human Visual Perception from Magnetoencephalography

To explore the timing and the underlying neural dynamics of visual perception, we analyzed the relationship between the manual reaction time (RT) to the onset of a visual stimulus and the time course of the evoked neural response simultaneously measured by magnetoencephalography (MEG). The visual stimuli were a transition from incoherent to coherent motion of random dots and an onset of a chromatic grating from a uniform field, which evoke neural responses in different cortical sites. For both stimuli, changes in median RT with changing stimulus strength (motion coherence or chromatic contrast) were accurately predicted, with a stimulus-independent postdetection delay, from the time that the temporally integrated MEG response crossed a threshold (integrator model). In comparison, the prediction of RT was less accurate from the peak MEG latency, or from the time that the nonintegrated MEG response crossed a threshold (level detector model). The integrator model could also account for, at least partially, intertrial changes in RT or in perception (hit/miss) to identical stimuli. Although we examined MEG–RT relationships mainly for data averaged over trials, the integrator model could show some correlations even for single-trial data. The model predictions deteriorated when only early visual responses presumably originating from the striate cortex were used as the input to the integrator model. Our results suggest that the perceptions for visual stimulus appearances are established in extrastriate areas [around MT (middle temporal visual area) for motion and around V4 (fourth visual area) for color] ∼150–200 ms before subjects manually react to the stimulus.

Neural Selectivity and Representation of Gloss in the Monkey Inferior Temporal Cortex

When we view an object, its appearance depends in large part on specific surface reflectance properties; among these is surface gloss, which provides important information about the material composition of the object and the fine structure of its surface. To study how gloss is represented in the visual cortical areas related to object recognition, we examined the responses of neurons in the inferior temporal (IT) cortex of the macaque monkey to a set of object images exhibiting various combinations of specular reflection, diffuse reflection, and roughness, which are important physical parameters of surface gloss. We found that there are neurons in the lower bank of the superior temporal sulcus that selectively respond to specific gloss. This neuronal selectivity was largely maintained when the shape or illumination of the object was modified and perceived glossiness was unchanged. By contrast, neural responses were significantly altered when the pixels of the images were randomly rearranged, and perceived glossiness was dramatically changed. The stimulus preference of these neurons differed from cell to cell, and, as a population, they systematically represented a variety of surface glosses. We conclude that, within the visual cortex, there are mechanisms operating to integrate local image features and extract information about surface gloss and that this information is systematically represented in the IT cortex, an area playing an important role in object recognition.

Distribution of colour-selective activity in the monkey inferior temporal cortex revealed by functional magnetic resonance imaging

Previous electrophysiological, neuroimaging and lesion studies have suggested that the anterior part of the monkey inferior temporal (IT) cortex, or area TE, plays an important role in colour processing. However, little is known about how colour information is distributed in these cortical regions. Here, we explored the distribution of colour‐selective activity in alert macaque monkeys using functional magnetic resonance imaging (fMRI) with two types of stimuli: a multicoloured (‘Mondrian’) pattern and an isoluminant colour grating. These two types of stimuli are both commonly used in human fMRI studies, but Mondrian stimuli, which contain a richer variety of hues and hence might be more suitable for activating higher‐order areas than grating stimuli, have not been used to examine colour‐selectivity in higher‐order areas in earlier monkey studies. With the Mondrian stimuli, we observed that areas along the ventral pathway, V1, V2/V3, V4 and the IT cortex, responded more strongly to colour stimuli than to luminance stimuli. In the IT cortex, we found that colour‐selective activities are not distributed uniformly, but are localized in discrete regions, each extending several millimetres in the anterior or posterior part of the IT cortex. The colour‐selective activation in the anterior IT was observed only with the Mondrian stimuli, whereas the colour‐selective activation in the posterior IT was observed with both the Mondrian and grating stimuli, with little overlap. These findings suggest that there are multiple subregions with differing stimulus selectivities distributed in the IT cortex, and that colour information is processed in these discrete subregions.

Selective responses to specular surfaces in the macaque visual cortex revealed by fMRI

The surface properties of objects, such as gloss, transparency and texture, provide important information about the material characteristics of objects in our visual environment. However, because there have been few reports on the neuronal responses to surface properties in primates, we still lack information about where and how surface properties are processed in the primate visual cortex. In this study, we used functional magnetic resonance imaging (fMRI) to examine the cortical responses to specular surfaces in the macaque visual cortex. Using computer graphics, we generated images of specular and matte objects and prepared scrambled images by locally randomizing the luminance phases of the images with specular and matte objects. In experiment 1, we contrasted the responses to specular images with those to matte and scrambled images. Activation was observed along the ventral visual pathway, including V1, V2, V3, V4 and the posterior inferior temporal (IT) cortex. In experiment 2, we manipulated the contrasts of images and found that the activation observed in these regions could not be explained solely by the global or local contrasts. These results suggest that image features related to specular surface are processed along the ventral visual pathway from V1 to specific regions in the IT cortex. This is consistent with previous human fMRI experiments that showed surface properties are processed in the ventral visual pathway.

Anterior and superior lateral occipito-temporal cortex responsible for target motion prediction during overt and covert visual pursuit

In smooth-pursuit eye movements (SPEM) with gain close to one, SPEM should be controlled mainly by prediction of target motion because retinal slip is nearly zero. We investigated the neural mechanisms of visual-target prediction by the three fMRI experiments. (1) Overt pursuit task: subjects pursued a sinusoidally moving target which blinked (blink condition) or did not blink (continuous condition). (2) Covert pursuit task: subjects covertly pursued the same target with eyes gazed at fixation point. (3) Attend-to-stationary target task: subjects brought attention on a stationary target with eyes gazed at fixation point. In the overt pursuit task, the SPEM gain and the delay in the blink condition were not very different from the continuous condition, indicating good prediction of the blinking target motion. Activities in the dorsolateral prefrontal, precentral, medial superior frontal, intraparietal, and lateral occipito-temporal cortexes increased in the blink-continuous subtraction. The V1 activity decreased for this contrast. In the covert pursuit task, only the anterior/superior LOTC activity remained in the blink-continuous subtraction. In the attend-to-stationary target task, the blink-continuous subtraction elicited no activation. Consequently, the a/sLOTC activity is responsible for target prediction rather than motor commands for eye movements or just target blinking such as visual saliency.

Wiener filter-magnetoencephalography of visual cortical activity.

This paper reports a revised Wiener filter to resolve the inverse problem for magnetoencephalograms (MEGs) according to the structural and functional constraints based on magnetic resonance imaging (MRI) and functional magnetic resonance imaging (fMRI). Wiener filter-MEG imaging for half field stimulation with the chromatic stimulus resolved fast, slow and late responses in V1, V4 and the inferotemporal cortex, respectively. The time courses of these responses were roughly comparable with those reported by unit recording studies of the corresponding monkey visual cortical areas. Wiener filter-MEG imaging had comparable spatial resolution and better signal to noise ratio than fMRI. The background noise was robust in fMRI responses, but became virtually eliminated in Wiener filter responses. Wiener filter-MEG imaging with upper and lower quadrant field stimulation demonstrated V1 responses differentially distributed respectively in the lower and upper banks of the calcarine sulcus. These results demonstrate that responses in two cortical areas facing close to each other can be resolved by Wiener filter-MEG. The present method provides a way to image brain activities with millisecond- and millimeter-order spatiotemporal resolution.

Mechanisms underlying the representation of angles embedded within contour stimuli in area V2 of macaque monkeys.

We previously found that surprisingly many V2 neurons showed selective responses to particular angles embedded within continuous contours [ M. Ito & H. Komatsu (2004)Journal of Neuroscience, 24, 3313–3324]. Here, we addressed whether the selectivity is dependent on the presence of individual constituent components or on the unique combination of these components. To reveal roles of constituent half‐lines in response to whole angles, we conducted a quantitative model study after the framework of cascade models. Our linear–non‐linear summation model implemented a few subunits selective to particular half‐lines and was fitted to neuronal responses for each neuron. The study indicates that the best‐fitting models well replicate the selectivity in the majority of V2 neurons and that the angle selectivity is dependent on a linear combination of responses to individual half‐line components of the angles. The implication is that optimal angles are given by a combination of two preferred half‐line components and the selectivity is sharpened by introducing suppression to non‐preferred half‐line components, rather than a specific facilitatory interaction between two preferred half‐line components. The study indicates the participation of the gain control of responsiveness according to the number of half‐line components. We also showed that the selectivity to acute angles depends on a combination of responses to one preferred component and weak responses to another component. Therefore, we concluded that the angle selectivity is dependent on selective responses to individual half‐line components of the angles rather than a unique combination between them, whereas neurons could be selective to various angle widths at area V2.