Sleep stage N2 is not discriminated from SWS in rodents. Almost 50% of sleep in adult humans is marked by a lighter form of non-REM sleep (stage “N2”) that is characterized by the occurrence of distinct (waxing and waning) sleep spindles ( FIGURE 1 B) and K-complexes in the EEG, but minor SWA. In addition, REM sleep is characterized by phasic REMs and by muscle atonia.
SWS is hallmarked by slow high-amplitude EEG oscillations (slow wave activity, SWA), whereas REM sleep (also termed paradoxical sleep) is characterized by wakelike fast and low-amplitude oscillatory brain activity. In human nocturnal sleep, SWS is predominant during the early part and decreases in intensity and duration across the sleep period, whereas REM sleep becomes more intense and extensive towards the end of the sleep period. Sleep in mammals consists of two core sleep stages: slow-wave sleep (SWS) and rapid-eye-movement (REM) sleep, which alternate in a cyclic manner ( FIGURE 1 A). Sleep probably occurs in all vertebrates, including birds, fishes, and reptiles, and sleeplike states are similarly observed in invertebrates like flies, bees, and cockroaches ( 209). Sleep deprivation and sleep disruptions cause severe cognitive and emotional problems ( 142, 634, 1243), and animals deprived of sleep for several weeks show temperature and weight dysregulation and ultimately die of infections and tissue lesions ( 973). Sleep occurs in regular intervals and is homeostatically regulated, i.e., a loss or delay of sleep results in subsequently prolonged sleep ( 113). Sleep is defined as a natural and reversible state of reduced responsiveness to external stimuli and relative inactivity, accompanied by a loss of consciousness.
While elaborated with respect to hippocampus-dependent memories, the concept of an active redistribution of memory representations from networks serving as temporary store into long-term stores might hold also for non-hippocampus-dependent memory, and even for nonneuronal, i.e., immunological memories, giving rise to the idea that the offline consolidation of memory during sleep represents a principle of long-term memory formation established in quite different physiological systems. Ensuing REM sleep may stabilize transformed memories. Consolidation originates from reactivation of recently encoded neuronal memory representations, which occur during SWS and transform respective representations for integration into long-term memory. Specifically, newer findings characterize sleep as a brain state optimizing memory consolidation, in opposition to the waking brain being optimized for encoding of memories. Whereas older research concentrated on the role of rapid-eye-movement (REM) sleep, recent work has revealed the importance of slow-wave sleep (SWS) for memory consolidation and also enlightened some of the underlying electrophysiological, neurochemical, and genetic mechanisms, as well as developmental aspects in these processes. Whereas initial theories posed a passive role for sleep enhancing memories by protecting them from interfering stimuli, current theories highlight an active role for sleep in which memories undergo a process of system consolidation during sleep. In this review we aim to comprehensively cover the field of “sleep and memory” research by providing a historical perspective on concepts and a discussion of more recent key findings. Over more than a century of research has established the fact that sleep benefits the retention of memory.