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planation may be that a unilateral lesion interrupts the connections between each of the paired groups of nuclei, which normally synchronize the two sides in the generation of rhythmic bursts of excitatory impulses to spinal motor neurons It is of interest that in a case of a very delimited metastasis to the NTS there was no apparent impact on the breathing pattern until a terminal respiratory arrest (Rhodes and Wightman) Voluntary Control of Breathing During speech, swallowing, breath-holding, or voluntary hyperventilation, the automaticity of the brainstem mechanisms of respiration is arrested in favor of re exive or of conscious control of diaphragmatic contraction The observations of Colebatch and coworkers, utilizing PET scanning, indicate that voluntary control of breathing is associated with activity in the motor and premotor cortex The experiments of Maskill and associates demonstrated that magnetic cortical stimulation of a region near the cranial vertex activates the diaphragm Although automatic and voluntary breathing utilize the same pools of cervical motor neurons that give rise to the phrenic nerves, the descending cortical pathways for voluntary breathing are distinct from those utilized by automatic brainstem mechanisms It is not known whether the voluntary signal bypasses the brainstem mechanisms or is possibly integrated there When both dorsal descending tracts subserving voluntary control are interrupted, as in the locked-in syndrome, the independent, automatic respiratory system in the medulla is capable of maintaining an almost perfectly regular breathing rate of 16 per minute with uniform tidal volumes These essential facts do not fully depict the rich interactions between the neuronal groups governing respiration and between the neurons for laryngeal and glottic activity that come into play during such coordinated acts as swallowing, sneezing, and coughing, and speaking The brainstem regions involved in holding breathing in abeyance while swallowing occurs are pertinent to aspiration, a common feature of many neurologic diseases, as discussed further on The drive applied to these systems is damped in processes such as Parkinson disease and may contribute to the problem of aspiration, as also discussed further on Afferent Respiratory In uences A number of signals that modulate respiratory drive originate in chemoreceptors located in the carotid artery These receptors are in uenced both by changes in pH and by hypoxia Chemoreceptor afferents pass along the carotid sinus nerves, which join the glossopharyngeal nerves and terminate in the NTS Aortic body receptors, which are less important as detectors of hyopxia, send afferent volleys to the medulla through the aortic nerves, which join the vagus nerves There are also chemoreceptors in the brainstem, but their precise location is uncertain Their main locus is thought to be in the ventral medulla, but other areas that are responsive to changes in pH have been demonstrated in animals What is clear is that these regions are sensitive not to the pH of CSF, as had been thought, but to the hydrogen content of the extracellular uid of the medulla Numerous stretch receptors within smooth muscle cells of the airways also project via the vagus nerves to the NTS and in uence the depth and duration of breathing Afferent signals from these specialized nerve endings mediate the Hering-Breuer re ex, described in 1868 a shortened inspiration and decreased tidal volume triggered by excessive lung expansion The Hering-Breuer mechanism seems not to be important at rest, since bilateral vagal section has no effect on the rate or depth of respiration These aspects of afferent pulmonary modulation of breathing have been.

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reviewed by Berger and colleagues It is interesting, however, that patients with high spinal transections and inability to breathe can still sense changes in lung volume, attesting to a nonspinal afferent route to the brainstem from lung receptors, probably through the vagus nerves In addition, there are receptors located between pulmonary epithelial cells that respond to irritants such as histamine and smoke They have been implicated in the genesis of asthma Also there are J-type receptors that are activated by substances in the interstitial uid of the lungs These are capable of inducing hyperpnea and probably play a role in driving ventilation under conditions of pulmonary edema Both the diaphragm and the accessory muscles of respiration contain conventional spindle receptors, but their role is not clear; all that can be said is that the diaphragm has a paucity of these receptors compared with other skeletal muscles (a property shared with extraocular muscles) and is therefore not subject to spasticity with corticospinal lesions or to the loss of tone in states such as REM sleep, in which gamma motor neuron activity is greatly diminished Dyspnea The common respiratory sensations of breathlessness, air hunger, chest tightness, or shortness of breath, all subsumed under the term dyspnea, have de ed neurophysiologic interpretation In animals, Chen and colleagues from Eldridge s laboratory have demonstrated that neurons in the thalamus and central midbrain tegmentum re in a graduated manner as respiratory drive is increased These neurons are in uenced greatly by afferent information from the chest wall, lung, and chemoreceptors and are postulated to be the thalamic representation of sensation from the thorax that is perceived at a cortical level as dyspnea However, functional imaging studies indicate that various areas of the cerebrum are activated by dyspnea, mainly the insula and limbic regions Aberrant Respiratory Patterns Many of the most interesting respiratory patterns observed in neurologic disease are found in comatose patients, and several of these patterns have been assigned localizing value, some of uncertain validity: central neurogenic hyperventilation, apneusis, and ataxic breathing These are discussed in relation to the clinical signs of coma (Chap 17) and sleep apnea (Chap 19) Some of the most bizarre cadences of breathing those in which unwanted breaths intrude on speech or those characterized by incoordination of laryngeal closure, diaphragmatic movement, or swallowing or by respiratory tics have occurred in paraneoplastic brainstem encephalitis Similar incoordinated patterns occur in certain extrapyramidal diseases Patterns such as episodic tachypnea up to 100 breaths per minute and loss of voluntary control of breathing were, in the past, noteworthy features of postencephalitic parkinsonism In Leeuwenhoek disease, named for the discoverer of the microscope who described and suffered from the disease, there is a continuous epigastric pulsation and dyspnea in association with rhythmic bursts of activity in the inspiratory muscles a respiratory myoclonus akin to palatal myoclonus (Phillips and Eldridge) Two such cases in our clinical material followed in uenza-like illnesses and resolved slowly over months Cheyne-Stokes breathing, the common and well-known waxing and waning type of cyclic ventilation reported by Cheyne in 1818 and later elaborated by Stokes, has for decades been ascribed to a prolongation of circulation time, as in congestive heart failure; but there are data that support a primary neural origin of the dis-.

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