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on-off switches in the transition between inspiration and expiration Inspiratory neurons are concentrated in the dorsal respiratory group and in the rostral portions of the ventral group, some of which have monosynaptic connections to the motor neurons of the phrenic nerves and the nerves to the intercostal muscles Normal breathing is actively inspiratory and only passively expiratory; however, under some circumstances of increased respiratory drive, the internal intercostal muscles and abdominal muscles actively expel air The expiratory neurons that mediate this activity are concentrated in the caudal portions of the ventral respiratory group and in the most rostral parts of the dorsal group On the basis of both neuroanatomic tracer and physiologic studies, it has been determined that these expiratory neurons project to spinal motor neurons and have an inhibitory in uence on inspiratory neurons The pathway of descending bers that arises in the inspiratory neurons and terminates on phrenic nerve motor neurons lies just lateral to the anterior horns of the upper three cervical cord segments When these tracts are damaged, automatic but not voluntary diaphragmatic movement on that side is lost As noted below, the bers carrying voluntary motor impulses to the diaphragm course more dorsally in the cord The phrenic motor neurons form a thin column in the medial parts of the ventral horns, extending from the third through fth cervical cord segments Damage to these neurons, of course, precludes both voluntary and automatic breathing The exact locus from which the breathing rhythm is generated, if there is such a site, is not known The conventional teaching has been that the dorsal respiratory group (DRG) was the dominant generator of the respiratory rhythm, but the situation is certainly more complex Animal experiments have focused attention instead on the rostral ventrolateral medulla (VRG) This region contains a group of neurons in the vicinity of the Botzinger complex (which itself contains neurons that re mainly during expiration) Cooling of this area or injection with neurotoxins causes the respiratory rhythm to cease (see the review by Duf n et al) It has been shown that the paired respiratory nuclei in the pons that are thought to act as switches between inspiration and expiration also possess a degree of autonomous rhythmicity, but their role in engendering cyclic breathing has not been clari ed Some workers are of the opinion that two or more sets of neurons in the VRG create a rhythm by their reciprocal activity or that oscillations arise within even larger networks (see Blessing for details) There are also centers in the pons that do not generate respiratory rhythms but may, under extreme circumstances, greatly in uence them One pontine group, the pneumotaxic center, modulates the response to hypoxia, hyopcapnia, and lung in ation In general, expiratory neurons are located laterally and inspiratory neurons medially in this center, but there is an additional group that lies between them and remains active during the transition between respiratory phases Also found in the lower pons is a group of neurons that prevent unrestrained activity of the medullary inspiratory neurons ( apneustic center ) In addition to these ambiguities regarding a center for the generation of respiratory rhythm, there is the dif culty that the nuclei described above are not well de ned in humans As to the effects of a unilateral brainstem lesion on ventilation, numerous cases of hypoventilation or total loss of automatic ventilation ( Ondine s curse see further on) have been recorded (Bogousslavsky et al) We have observed several such remarkable cases as well, due in most instances to a large lateral medullary infarction If the neural oscillators on each side were totally independent, such a syndrome should not be possible The likely ex-.

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