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ruzzi and Magoun that electrical stimulation of the medial midbrain tegmentum and adjacent areas just above this level caused a lightly anesthetized animal to become suddenly alert and its EEG to change correspondingly, ie, to become desynchronized, in a manner identical to normal arousal by sensory stimuli The sites at which stimulation led to arousal consisted of a series of points extending from the nonspeci c medial thalamic nuclei down through the caudal midbrain These points were situated along the loosely organized core of neurons that anatomists had referred to as the reticular system or formation The anatomic studies of the Scheibels have described widespread innervation of the reticular formation by multiple bifurcating and collateral axons of the ascending sensory systems, implying that this area was maintained in a tonically active state by ascending sensory stimulation Because this region, especially the medial thalamus, projects widely to the cerebral hemispheres, there arose the concept of a reticular activating system (RAS) that maintained the alert state and the inactivation of which led to an unarousable state In this way, despite a number of experimental inconsistencies (see Steriade) the paramedian upper brainstem tegmentum and lower diencephalon came to be conceived as the locus of the alerting system of the brain The anatomic boundaries of the upper brainstem reticular activating system are somewhat indistinct This system is interspersed throughout the paramedian regions of the upper (rostral) pontine and midbrain tegmentum; at the thalamic level, it includes the functionally related posterior paramedian, parafascicular, and medial portions of the centromedian and adjacent intralaminar nuclei In the brainstem, nuclei of the reticular formation receive collaterals from the spinothalamic and trigeminal-thalamic pathways and project not just to the sensory cortex of the parietal lobe, as do the thalamic relay nuclei for somatic sensation, but to the whole of the cerebral cortex Thus, it would seem that sensory stimulation has a double effect it conveys information to the brain from somatic structures and the environment and also activates those parts of the nervous system on which the maintenance of consciousness depends The cerebral cortex not only receives impulses from the ascending reticular activating system but also modulates this incoming information via corticofugal projections to the reticular formation Although the physiology of the reticular activating system is far more complicated than this simple formulation would suggest, it nevertheless, as a working idea, retains a great deal of clinical credibility and makes comprehensible some of the neuropathologic observations noted further on, under Pathologic Anatomy of Coma Although for many years it has been taught that arousal causes a desynchronization of brainwave activity (in distinction to the synchronized activity of sleep), it has become apparent that during wakefulness, there is also a widespread low-voltage fast rhythm (a gamma rhythm that has a frequency of 30 to 60 Hz) This activity, coordinated by the thalamus, has been theorized to synchronize widespread cortical activity and to account perhaps for the uni cation of modular aspects of experience (color, shape, motion) that are processed in different cortical regions In this way the rhythm has been theorized to bind various aspects of a sensory experience or a memory This fast and widespread electrographic activity is not easily appreciable with EEG surface recordings but it can be extracted by sophisticated mathematical tools Using such electrophysiologic methods, Meador and colleagues have shown that the rhythm can be detected over the primary somatosensory cortex after an electrical stimulus on the contralateral hand is perceived, but.

not if the patient fails to perceive it The clinical meaning of the rhythm is controversial, but it has elicited great interest because it may give insight into several intriguing questions about the states of consciousness

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In a number of disease processes that disturb consciousness, there is direct interference with the metabolic activities of the nerve cells in the cerebral cortex and the central nuclei of the brain Hypoxia, global ischemia, hypoglycemia, hyper- and hypo-osmolar states, acidosis, alkalosis, hypokalemia, hyperammonemia, hypercalcemia, hypercarbia, drug intoxication, and severe vitamin de ciencies are well-known examples (see Chap 40 and Table 40-1) In general, the loss of consciousness in these conditions parallels the reduction in cerebral metabolism or blood ow For example, in the case of global ischemia, an acute drop in cerebral blood ow (CBF) to 25 mL/min/100 g brain tissue from its normal 55 mL/ min/l00 g causes slowing of the EEG and syncope or impaired consciousness; a drop in CBF below 12 to 15 mL/min/100 g causes electrocerebral silence, coma, and cessation of most neuronal metabolic and synaptic functions Lower levels are tolerated if arrived at more slowly, but neurons cannot survive when ow is reduced below 8 to 10 mL/min/100 g There is a corresponding reduction in the cerebral metabolic rate In other types of metabolic encephalopathy or with widespread anatomic damage to the hemispheres, blood ow stays near normal levels while metabolism is greatly reduced Oxygen consumption of 2 mg/min/100 g (approximately half of normal) is incompatible with an alert state An exception to these statements is the coma that arises from seizures, in which metabolism and blood ow are greatly increased during the seizure Extremes of body temperature (above 41 C or below 30 C) also induce coma through a nonspeci c effect on the metabolic activity of neurons These metabolic changes are probably epiphenomena re ecting, in each particular encephalopathy, a speci c type of dysfunction in neurons and their supporting cells The endogenous metabolic toxin(s) that are responsible for coma cannot always be identi ed In diabetes, acetone bodies (acetoacetic acid, -hydroxybutyric acid, and acetone) are present in high concentration; in uremia, there is probably an accumulation of dialyzable small molecular toxins, notably phenolic derivatives of the aromatic amino acids In hepatic coma, elevation of blood NH3 to ve to six times normal levels corresponds roughly to the level of coma Lactic acidosis may affect the brain by lowering arterial blood pH to less than 70 The impairment of consciousness that accompanies pulmonary insuf ciency is related mainly to hypercapnia (see page 964) In hyponatremia (Na 120 meq/L) of whatever cause, neuronal dysfunction is probably due to the intracellular movement of water, leading to neuronal swelling and loss of potassium chloride from the cells The mode of action of bacterial toxins is not fully understood Drugs such as general anesthetics (see below), alcohol, opiates, barbiturates, phenytoin, antidepressants, and diazepines induce coma by their direct effects on neuronal membranes in the cerebrum and reticular activating system or on neurotransmitters and their receptors Others, such as methyl alcohol and ethylene glycol, act by producing a metabolic acidosis Although the coma of toxic and metabolic diseases usually evolves through stages of drowsiness, confusion, and stupor (and the reverse sequence occurs during emergence from coma), each disease imparts its own char-.

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