Microsaccades are very small, involuntary flicks in eye position that occur on average once or twice per second during attempted visual fixation. Microsaccades give rise to EMG eye muscle spikes that can distort the spectrum of the scalp EEG and mimic increases in gamma band power. Here we demonstrate that microsaccades are also accompanied by genuine and sizeable cortical activity, manifested in the EEG. In three experiments, high-resolution eye movements were corecorded with the EEG: during sustained fixation of checkerboard and face stimuli and in a standard visual oddball task that required the counting of target stimuli. Results show that microsaccades as small as 0.15°generate a field potential over occipital cortex and midcentral scalp sites 100 -140 ms after movement onset, which resembles the visual lambda response evoked by larger voluntary saccades. This challenges the standard assumption of human brain imaging studies that saccade-related brain activity is precluded by fixation, even when fully complied with. Instead, additional cortical potentials from microsaccades were present in 86% of the oddball task trials and of similar amplitude as the visual response to stimulus onset. Furthermore, microsaccade probability varied systematically according to the proportion of target stimuli in the oddball task, causing modulations of late stimulus-locked event-related potential (ERP) components. Microsaccades present an unrecognized source of visual brain signal that is of interest for vision research and may have influenced the data of many ERP and neuroimaging studies.
Although it is well established that attention affects visual performance in many ways, by using a novel paradigm [Carrasco, M., Ling, S., & Read. S. (2004). Attention alters appearance. Nature Neuroscience, 7, 308-313.] it has recently been shown that attention can alter the perception of different properties of stationary stimuli (e.g., contrast, spatial frequency, gap size). However, it is not clear whether attention can also change the phenomenological appearance of moving stimuli, as to date psychophysical and neuro-imaging studies have specifically shown that attention affects the adaptability of the visual motion system. Here, in five experiments we demonstrated that attention effectively alters the perceived speed of moving stimuli, so that attended stimuli were judged as moving faster than less attended stimuli. However, our results suggest that this change in visual performance was not accompanied by a corresponding change in the phenomenological appearance of the speed of the moving stimulus.
The same object produces quite distinct images in the cortical representation, depending on whether it is looked at foveally or with the periphery, yet some form of size constancy prevents us from experiencing objects inflating or deflating as we move our eyes. According to the prominent sensorimotor account of vision by O'Regan and Noë [1], we constantly learn to discount the predictable sensory effects of motor actions, such as the projection of a stimulus on a larger cortical area as it gets foveated. Although previous studies have shown that foveal and parafoveal inputs can be associated in visual memory [2, 3], trans-saccadic prediction error could in principle re-calibrate even the appearance of peripheral and foveal stimuli. Here we introduce a new paradigm that induces such changes in the relative appearance of peripheral and foveal stimuli when directly compared. Repeated exposure to a trans-saccadic change in size, though unnoticed by most observers, induced a substantial modification of perceived size that lasted at least 1 day. Prediction is not limited to the motor system but can also occur for the sensory effects of external events, such as stimulus motion. We show that perceptual re-calibration can occur in the absence of eye movements if the change in size occurs predictably while objects move across the visual field. Perceptual uniformity emerges due to the continuously updated prediction of foveal appearance based on peripheral appearance.
Eyes never stop moving. Even when asked to maintain the eyes at fixation, the oculomotor system produces small and rapid eye movements called microsaccades, at a frequency of about 1.5-2 s(-1). The frequency of microsaccades changes when a stimulus is presented in the visual field, showing a stereotyped response pattern consisting of an early inhibition of microsaccades followed by a rebound, before the baseline is reached again. Although this pattern of response has generally been considered as a sort of oculomotor reflex, directional biases in microsaccades have been recently linked to the orienting of spatial attention. In the present study, we show for the first time that regardless of any spatial bias, the pattern of absolute microsaccadic frequency is different for oddball stimuli compared to that elicited by standard stimuli. In a visual-oddball task, the oddball stimuli caused an initial prolonged inhibition of microsaccades, particularly when oddballs had to be explicitly recognized and remembered. The present findings suggest that high-order cognitive processes, other than spatial attention, can influence the frequency of microsaccades. Finally, we also introduce a new method for exploring the visual system response to oddball stimuli.
The variable resolution and limited processing capacity of the human visual system requires us to sample the world with eye movements and attentive processes. Here we show that where observers look can strongly modulate their reports of simple surface attributes, such as lightness. When observers matched the color of natural objects they based their judgments on the brightest parts of the objects; at the same time, they tended to fixate points with above-average luminance. When we forced participants to fixate a specific point on the object using a gaze-contingent display setup, the matched lightness was higher when observers fixated bright regions. This finding indicates a causal link between the luminance of the fixated region and the lightness match for the whole object. Simulations with rendered physical lighting show that higher values in an object's luminance distribution are particularly informative about reflectance. This sampling strategy is an efficient and simple heuristic for the visual system to achieve accurate and invariant judgments of lightness.lightness constancy | lightness perception | visual perception | attention J udging the lightness of visual stimuli has been studied for centuries, since the original investigations by Weber (1) and Fechner (2). The light reaching the eye is the product of the illumination and the reflectance of the object, and also depends on the scene geometry (3). However, only the proportion of reflected light is an invariant property of the object and thus of great importance for vision. There are several well-established factors that support lightness constancy in the face of these challenges. On the one hand, lateral inhibition between retinal neurons filters out shallow intensity gradients, which are mostly caused by illumination effects (4, 5). On the other hand, more complex factors also have an effect on lightness perception, such as object shape (6-9) or the interpretation of transparent surfaces (10, 11). However, eye movements have been almost completely neglected so far, even though a general influence of viewing behavior has been shown for some color constancy tasks (12)(13)(14)(15). This finding is surprising because the visual system needs to sample the local properties of objects and this is accomplished by moving the eyes and the focus of spatial attention around. Because visual acuity, luminance sensitivity, contrast sensitivity, and color sensitivity change with retinal eccentricity (16-18), our visual system has to stitch together its representation of the world from many small samples to analyze the visual scene in detail. Peripheral vision is not only characterized by poor resolution, but also the appearance of basic visual features-like spatial frequency, luminance, or chromatic saturation-is distorted in the periphery of the visual field (19-22). Eye movements may then be used to select relevant information, even for stimuli that are above threshold in peripheral vision. We investigated whether the distribution of fixations on an object has an e...
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