02 November 2009

Notes on electrical rhythms in the brain

I'm reading George Buszáki's Rhythms of the Brain at the moment. Here are some notes on what I've read so far. Please correct me if I've got something wrong.

Electrical fluctuations in the cortex are organized into rhytmic oscillations at different spatial and temporal scales. The resting cortex is characterized by oscillations primarily in the alpha band (8-12 Hz, the brain's 'default network'). The active (i.e. behaving, percieving) cortex is characterized by oscillations primarily in the gamma band (25-100 Hz). Buszáki and others argue that cortical neurons that synchronize their membrane oscillations in the gamma band 'bind' their respective functions (e.g. visual feature detection) together into cognitive processes (e.g. object perception). Such formations of neurons are called neuronal groups or assemblies. Particularly striking are neuronal groups in the gamma range emerging in the prefrontal cortex for the duration of time that human patients are asked to hold items in working memory.

(alpha)

(gamma)

Cortical neurons sponaneously synchronize their membrane oscillations in the gamma range and form transient neuronal groups even in the absence of stimuli. This is, at least in part, due to the time constants of GABA curents, synaptic delays and synaptic potentiation. Buszáki writes:
"If neurons are already engaged in internal synchronization, the external stimulus will compete with the central oscillator, and the coutcome depends on the relative timing and strenght of the external input and the propensity of the internal oscillator. The stimulus may be ignored, or it may enhance or quench the internal oscillation." p.255
In other words, the effect of a stimulus on cortical activity depends strongly on the prior state of the brain. This explains the significant variability in brain activity (e.g. on EEG/MEG/fMRI) seen within and between subjects in response to invariant stimuli. Buszáki laments the fact that this variability is usually averaged out and treated as 'noise'. Björn Brembs often makes a similar argument.

Stimuli interact with ongoing cortical activity in various ways. Whereas a weak stimulus may reset the phase of ongoing oscillations, a strong or salient stimulus may completely change the type and distribution of oscillations in the cortex. Several studies have found that strong ongoing oscillations in the gamma, theta or alpha ranges prior to stimulus presentation promote efficient memory encoding. A stimulus that resets a strong rhythm presumably has a larger impact on brain activity than one that resets a weak rhythm. The presence or absence of strong rhythms in the brain is directly related to states of attention and catecholamine concentrations.
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