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Brain–Computer Interfaces - Index of

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52 G. Pfurtscheller and C. Neuper<br />

regions. Once the movement sequence has been learned and the movement is performed<br />

more “automatically,” the ERD is reduced. These ERD findings strongly<br />

suggest that activity in primary sensorimotor areas increases in association with<br />

learning a new motor task and decreases after the task has been learned [41].<br />

The involvement <strong>of</strong> the primary motor area in learning motor sequences was also<br />

suggested by Pascual-Leone [42], who studied motor output maps using TMS.<br />

The opposite <strong>of</strong> desynchronization is synchronization, when the amplitude<br />

enhancement is based on the cooperative or synchronized behavior <strong>of</strong> a large number<br />

<strong>of</strong> neurons. When the summed synaptic events become sufficiently large, the<br />

field potentials can be recorded not only with macro electrodes within the cortex,<br />

but also over the scalp. Large mu waves on the scalp need coherent activity<br />

<strong>of</strong> cell assemblies over at least several square centimeters [43, 44]. When patches <strong>of</strong><br />

neurons display coherent activity in the alpha frequency band, active processing <strong>of</strong><br />

information is very unlikely and probably reflects an inhibitory effect.<br />

However, inhibition in neural networks is very important, not only to optimize<br />

energy demands but also to limit and control excitatory processes. Klimesch [45]<br />

suggested that synchronized alpha band rhythms during mental inactivity (idling)<br />

are important to introduce powerful inhibitory effects, which could block a memory<br />

search from entering irrelevant parts <strong>of</strong> neural networks. Combined EEG and TMS<br />

studies demonstrated that the common pathway from motor cortex to the target hand<br />

muscle was significantly inhibited during the movement inhibition conditions, and<br />

the upper mu components simultaneously displayed synchronization in the hand<br />

area. In contrast, the TMS response was increased during execution <strong>of</strong> the movement<br />

sequence, and a mu ERD was observed in the hand representation area [46].<br />

Adrian and Matthews [47] described a system that is neither receiving nor processing<br />

sensory information as an idling system. Localized synchronization <strong>of</strong><br />

12–14 Hz components in awake cats was interpreted by Case and Harper [48] as<br />

a result <strong>of</strong> idling cortical areas. Cortical idling can thus denote a cortical area <strong>of</strong> at<br />

least some cm 2 that is not involved in processing sensory input or preparing motor<br />

output. In this sense, occipital alpha rhythms can be considered idling rhythms <strong>of</strong><br />

the visual areas, and mu rhythms as idling rhythms <strong>of</strong> sensorimotor areas [49]. Also,<br />

sleep spindles during early sleep stages can occur when signals through the thalamus<br />

are blocked [40].<br />

5 “Focal ERD/Surround ERS”<br />

Localized desynchronization <strong>of</strong> the mu rhythms related to a specific sensorimotor<br />

event does not occur in isolation, but can be accompanied by increased synchronization<br />

in neighboring cortical areas that correspond to the same or to another<br />

information processing modality. Lopes da Silva [38] introduced the term “focal<br />

ERD/surround ERS” to describe this phenomenon. Gerl<strong>of</strong>f et al. [50] reported an<br />

antagonistic behavior with desynchronization <strong>of</strong> central mu rhythm and synchronization<br />

<strong>of</strong> parieto-occipital alpha rhythms during repetitive brief finger movement.

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