sleep/wake cycle, as sleep stages often alternate with a On the other hand, however, many problems concerning shorter time constant, with DS, in particular, normally not cerebral blood flow and metabolism remain to be solved lasting that long. A common strategy in most sleep studies (e.g. the regulation of blood flow, the nature of the has been, therefore, to correlate glucose utilization during flow-metabolism coupling, the existence of capillary measurement with the time spent in different sleep/wake recruitment, and the extent of non-oxidative metabolism).states (weighted for variations in glucose plasma concentraSleep is a natural, stereotyped, repetitive phenomenon, tion (in animals) and positron emission tomography (in humans) model with which to address these specific problems, while are hereafter considered separately, the same common logic findings about brain circulation and metabolism during sleep applies and their results are, in principle, comparable. will foster our understanding of brain circulation and metabolism in general.(a) Autoradiography studies Rat: Ramm and Frost (1983) found no correlation between this reason experimental data in this field have been tightly graphy), and (ii)
C E R E B R A L E N E R G Y METABOLISM
Direct measurements2.1.1 Local cerebral glucose utilization. When Greenberg wrote his review in 1980, very few attempts had been made to measure directly cerebral metabolism during sleep (Mangold et al. 1955;Meyer and Toyoda 1971) and, due to limitations in methodology, none could address the issue of regional differences in metabolic activity. The rather long list of papers considered in this section bears witness to the mean cerebral glucose utilization (reflected by mean optical density in the autoradiographs) and the different sleep/wake states. They also considered the "relative metabolic activity", which was calculated as the regional optic density given as a proportion of the mean brain optic density; this expresses the level of metabolic activity in a region relative to the mean level in the brain. During SS, relative metabolic activity decreases in the cerebellum, in most thalamic nuclei and in cortical layer IV, and increases in some hippocampal and hypothalamic structures. DS is associated with a relative decrease in metabolic activity in the cerebellum, and a relative increase in some brainstem regions (e.g. pontine Pavlides et al. (1987) reported a decreased cerebral glucose Correspondence: Institute Fisiologia Umana, Porta S. Donato 2, raphe, gigantocellular tegmental field, substantia nigra).