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Brain Signals for Brain–Computer <strong>Interfaces</strong> 33<br />

cortex (area V1). It is followed by a positive component at ~100 ms (P1 or P100)<br />

and a negative complex at ~145 msec (N1 or N145). While the neural generators for<br />

P1 and N1 are still debated, they are probably striate and extrastriate visual areas.<br />

The steady-state visual evoked potential (SSVEP) is elicited by repetitive patternreversal<br />

stimulation [12]. It is thought to arise from the same areas that produce the<br />

VEP, plus the motion sensitive area MT/V5.<br />

The VEP and SSVEP depend mainly on the properties <strong>of</strong> the visual stimulus.<br />

They have not been shown to vary with intent on a trial-by-trial basis. Nevertheless,<br />

the first BCI systems [13–15] and some modern systems [16, 17] are VEP- or<br />

SSVEP-based. For example, the BCI user can modulate the SSVEP by looking at<br />

one <strong>of</strong> several visual stimuli, each with different stimulus properties (i.e., different<br />

flash rates). SSVEP features vary according to the stimulus properties and are<br />

measured to determine which target is being looked at. The computer then executes<br />

the command associated with the target that the user is looking at. This communication<br />

system is equivalent to systems that determine gaze direction from the eyes<br />

themselves. Since it depends on the user’s ability to control gaze direction, it is<br />

categorized as a dependent BCI system, that is, it requires muscle control. Recent<br />

evidence suggests that it is possible to modulate SSVEP features by shifting attention<br />

(as opposed to gaze direction), and thus that the SSVEP might support operation<br />

<strong>of</strong> an independent BCI (i.e., a BCI that does not depend on neuromuscular function)<br />

[16, 18, 19].<br />

P300<br />

When an auditory, visual, or somatosensory (touch) stimulus that is infrequent,<br />

desirable, or in other ways significant is interspersed with frequent or routine stimuli,<br />

it typically evokes a positive peak at about 300 ms after stimulus onset (i.e., a<br />

P300 ERP) in the EEG over centroparietal cortex [20–22]. A stimulation protocol <strong>of</strong><br />

this kind is known as an ‘oddball’ paradigm [23, 24]. Evocation <strong>of</strong> the P300 in the<br />

oddball paradigm requires that the subject attend to the target stimulus. While the<br />

underlying neural generators <strong>of</strong> the P300 are debated, it is thought that the signal<br />

reflects rapid neural inhibition <strong>of</strong> ongoing activity and that this inhibition facilitates<br />

transmission <strong>of</strong> stimulus/task information from frontal to temporal-parietal cortical<br />

areas [25, 26].<br />

In a P300-based BCI system, the user is presented with an array <strong>of</strong> auditory,<br />

visual, or somatosensory stimuli, each <strong>of</strong> which represents a particular output (e.g.,<br />

spelling a particular letter), and pays attention to the stimulus that represents the<br />

action s/he desires. That attended stimulus elicits a P300 and the other stimuli do<br />

not (Fig. 2a). The BCI recognizes the P300 and then executes the output specified<br />

by the eliciting stimulus. Since it requires only that a user modulate attention, rather<br />

than any muscular output, a P300-based BCI is an independent BCI.<br />

The first P300-based BCI system was developed by Donchin and his colleagues<br />

[27, 28]. It presents the user with a 6 × 6 matrix <strong>of</strong> letters, numbers, and/or<br />

other symbols. The individual rows and columns <strong>of</strong> the array flash in succession<br />

as the user attends to the desired item and counts how many times it flashes. The

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