Millivolt-Scale DC Shifts in the Human Scalp EEG: Evidence for a Nonneuronal Generator

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Abstract
Slow shifts in the human scalp-recorded EEG, including those related to changes in brain CO2 levels, have been generally assumed to result from changes in the level of tonic excitation of apical dendrites of cortical pyramidal neurons. We readdressed this issue using DC-EEG shifts elicited in healthy adult subjects by hypo- or hypercapnia. A 3-min period of hyperventilation resulted in a prompt negative shift with a rate of up to 10 μV/s at the vertex (Cz) and an extremely steep dependence (up to 100 μV/mmHg) on the end-tidal Pco2. This shift had a maximum of up to −2 mV at Cz versus the temporal derivations (T3/T4). Hyperventilation-like breathing of 5% CO2-95% O2, which does not lead to a significant hypocapnia, resulted in a near-complete block of the negative DC shift at Cz. Hypoventilation, or breathing 5% CO2 in air at normal respiratory rate, induced a positive shift. The high amplitude of the voltage gradients on the scalp induced by hyperventilation is not consistent with a neuronal origin. Instead, the present data suggest that they are generated by extracortical volume currents driven by a Pco2-dependent potential difference across epithelia separating the cerebrospinal fluid and blood. Since changes in respiratory patterns and, hence, in the level of brain Pco2, are likely to occur under a number of experimental conditions in which slow EEG responses have been reported (e.g., attention shifts, preparatory states, epileptic seizures, and hypoxic episodes), the present results call for a thorough reexamination of the mechanisms underlying scalp-recorded DC-EEG responses.