Competing visual flicker reveals attention-like rivalry in the fly brain

After 40 years of investigation, steady-state visually evoked potentials (SSVEPs) have been shown to be useful for many paradigms in cognitive (visual attention, binocular rivalry, working memory, and brain rhythms) and clinical neuroscience (aging, neurodegenerative disorders, schizophrenia, ophthalmic pathologies, migraine, autism, depression, anxiety, stress, and epilepsy). Recently, in engineering, SSVEPs found a novel application for SSVEP-driven brain-computer interface (BCI) systems. Although some SSVEP properties are well documented, many questions are still hotly debated. We provide an overview of recent SSVEP studies in neuroscience (using implanted and scalp EEG, fMRI, or PET), with the perspective of modern theories about the visual pathway. We investigate the steady-state evoked activity, its properties, and the mechanisms behind SSVEP generation. Next, we describe the SSVEP-BCI paradigm and review recently developed SSVEP-based BCI systems. Lastly, we outline future research directions related to basic and applied aspects of SSVEPs. Copyright 2009 Elsevier Ltd. All rights reserved.

Event-related brain potentials (ERPs) provide high-resolution measures of the time course of neuronal activity patterns associated with perceptual and cognitive processes. New techniques for ERP source analysis and comparisons with data from blood-flow neuroimaging studies enable improved localization of cortical activity during visual selective attention. ERP modulations during spatial attention point toward a mechanism of gain control over information flow in extrastriate visual cortical pathways, starting about 80 ms after stimulus onset. Paying attention to nonspatial features such as color, motion, or shape is manifested by qualitatively different ERP patterns in multiple cortical areas that begin with latencies of 100-150 ms. The processing of nonspatial features seems to be contingent upon the prior selection of location, consistent with early selection theories of attention and with the hypothesis that spatial attention is "special."

Detection of a visual signal requires information to reach a system capable of eliciting arbitrary responses required by the experimenter. Detection latencies are reduced when subjects receive a cue that indicates where in the visual field the signal will occur. This shift in efficiency appears to be due to an alignment (orienting) of the central attentional system with the pathways to be activated by the visual input. It would also be possible to describe these results as being due to a reduced criterion at the expected target position. However, this description ignores important constraints about the way in which expectancy improves performance. First, when subjects are cued on each trial, they show stronger expectancy effects than when a probable position is held constant for a block, indicating the active nature of the expectancy. Second, while information on spatial position improves performance, information on the form of the stimulus does not. Third, expectancy may lead to improvements in latency without a reduction in accuracy. Fourth, there appears to be little ability to lower the criterion at two positions that are not spatially contiguous. A framework involving the employment of a limited-capacity attentional mechanism seems to capture these constraints better than the more general language of criterion setting. Using this framework, we find that attention shifts are not closely related to the saccadic eye movement system. For luminance detection the retina appears to be equipotential with respect to attention shifts, since costs to unexpected stimuli are similar whether foveal or peripheral. These results appear to provide an important model system for the study of the relationship between attention and the structure of the visual system.