The ontogeny of mammalian sleep: a response to Frank and Heller (2003)
In a recent review, Frank and Heller (2003) provided support for their ‘presleep theory’ of sleep development. According to this theory, rapid eye movement (REM) and non-rapid eye movement (Non-REM) sleep in rats emerge from a common ‘dissociated’ state only when the neocortical EEG differentiates at 12 days of age (P12). Among the assumptions and inferences associated with this theory is that sleep before EEG differentiation is only ‘sleep-like’ and can only be characterized using behavioral measures; that the neural mechanisms governing presleep are distinct from those governing REM and Non-REM sleep; and that the presleep theory is the only theory that can account for developmental periods when REM and Non-REM sleep components appear to overlap. Evidence from our laboratory and others, however, refutes or casts doubt on these and other assertions. For example, infant sleep in rats is not ‘sleep-like’ in that it satisfies nearly every criterion used to characterize sleep across species. In addition, beginning as early as P2 in rats, myoclonic twitching occurs only against a background of muscle atonia, indicating that infant sleep is not dissociated and that electrographic measures are available for sleep characterization. Finally, improved techniques are leading to new insights concerning the neural substrates of sleep during early infancy. Thus, while many important developmental questions remain, the presleep theory, at least in its present form, does not accurately reflect the phenomenology of infant sleep.