Cortical development, electroencephalogram rhythms, and the sleep/wake cycle

Abstract

During adulthood, electroencephalogram (EEG) recordings are used to distinguish wake, non-rapid eye movement sleep, and rapid eye movement sleep states. The close association between behavioral states and EEG rhythms is reached late during development, after birth in humans and by the end of the second postnatal week in rats and mice. This critical time is also when cortical activity switches from a discontinuous to a continuous pattern. We review the major cellular and network changes that can account for this transition. After this close link is established, new evidence suggests that the slow waves of non-rapid eye movement sleep may function as markers to track cortical development. However, before the EEG can be used to identify behavioral states, two distinct sleep phases--quiet sleep and active sleep--are identified based on behavioral criteria and muscle activity. During this early phase of development, cortical activity is far from being disorganized, despite the presence of long periods of neuronal silence and the poor modulation by behavioral states. Specific EEG patterns, such as spindle bursts and gamma oscillations, have been identified very early on and are believed to play a significant role in the refinement of brain circuits. Because most early EEG patterns do not map to a specific behavioral state, their contribution to the presumptive role of sleep in brain maturation remains to be established and should be a major focus for future research.

Keywords: Active sleep; Gamma activity; NREM sleep; Quiet sleep; REM sleep; Spindle bursts; Theta activity.

Figure 1

Schematic representations of cortical EEG recordings in a rat pup. During the first postnatal week transient events such as spindle bursts and evoked gamma oscillations can be observed, but EEG activity is mainly independent of behavioral states (see text for details). Traces are redrawn from (46) (spindle bursts), (64) (evoked gamma), and (13) (sleep/wake traces). Behavioral states during the first postnatal week were defined based on behavioral observation (respiration pattern, body, vibrissae, and tail movements, eyes open/closed).