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Transient visually evoked potentials to the pattern reversal and onset of sinusoidal gratings

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      They are stimuli that at high contrasts interfere with the visibility of small low contrast target stimuli, masking them effectively (Chronicle and Wilkins, 1996). The stimuli evoke high-amplitude electrical signals from the scalp (Plant et al., 1983) and large blood oxygenation dependent signals during functional magnetic resonance imaging (Huang et al., 2003). They induce perceptual distortions (Wilkins et al., 1984) and, in those who are susceptible, migraine (Huang et al., 2003) and seizures (Wilkins et al., 1980).

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      Clear evidence has been provided that different spatial frequencies result in different VEPs. The most salient difference occurs between 70 and 120 ms after stimulus onset over occipital–temporal areas, consisting of an increased negative (or decreased positive) amplitude to high as compared to low spatial frequencies (e.g., Bodis-Wollner et al., 1992; Harter and White, 1970; Kenemans et al., 1993; Plant et al., 1983; Proverbio et al., 1996, 1993; Reed et al., 1884; Zani and Proverbio, 1995). Several investigators have suggested that these differences in amplitude are related to different brain areas (e.g., Aine et al., 1990; Drasdo, 1980; Hudnell et al., 1990).

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      Specifically, around 80 ms high spatial frequencies evoke more negativity over occipital areas than do low spatial frequencies. This early sensory difference between high and low spatial frequency processing expressed in VEP amplitude is robust and has been found repeatedly in adults and children, using both gratings and checkerboards, with both sinusoidal and square-wave stimuli, and irrespective of attention manipulation (Rietveld et al., 1967; Harter and White, 1970; Drasdo, 1980; Plant et al., 1983; Skrandies, 1984; Reed et al., 1984; Ossenblok and Spekreijse, 1991; Ossenblok, 1992; Kenemans et al., 1993; Ossenblok et al., 1994; Kenemans and Lorist, 1995; Zani and Proverbio, 1995; Proverbio et al., 1996; Kenemans et al., 2000; Martinez et al., 2001; Boeschoten et al., 2005b). Further, dipole-source analyses in typical subjects have indicated that the larger negativity during high spatial frequency processing around 80 ms originates from relatively (peri-) striate areas, while the larger positivity seen during low spatial frequency processing around this latency originates from relatively extra-striate areas (Ossenblok and Spekreijse, 1991; Ossenblok, 1992; Ossenblok et al., 1994; Kenemans et al., 2000; Martinez et al., 2001; Baas et al., 2002; Boeschoten et al., 2005b).

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      Part of the conflict in the literature is a consequence of attempts to relate components in the waveform across studies from different laboratories that have used very different stimuli. Stimulus types (checkerboard vs gratings), spatial frequencies (low vs high), contrast (low vs high), the location of the stimulus field (upper vs lower or right vs left), eccentricities (central vs peripheral) are important factors that have an impact on whether one sees CI and CII as predominantly a striate or extrastriate source (Aine et al., 1995; Jeffreys, 1977; Kenemans et al., 2000; Plant et al., 1983). The multiplicity of the extrastriate cortices in humans is slowly being unravelled and will require additional studies.

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