David Megirian

Respiratory roles of genioglossus, sternothyroid, and sternohyoid muscles during sleep.

We examined the respiratory activity of the genioglossus, sternothyroid, and sternohyoid muscles of the rat during nonrapid eye movement (non-REM) and REM sleep. Each animal carried implanted electrodes for recording the integrated EMG activity of respiratory muscles, the postural tone (EMG), and electrocortical activity (polygraphic identification of sleep-waking states). The three upper airway muscles exhibited inspiratory activity during non-REM sleep while rats breathed ambient air. Curled up postures promoted inspiratory activity of genioglossus and sternothyroid muscles, an effect enhanced by CO2 breathing but reduced by hypoxic breathing. During REM sleep, genioglossus and sternothyroid muscles lost their activity but the sternohyoid muscles retained their inspiratory activity. We conclude that the genioglossus and sternothyroid muscles contribute to upper airway patency during non-REM sleep, an effect CO2 augments but hypoxia reduces. The sternohyoid muscles have at least two functions during both sleep states: they contribute to maintenance of upper airway patency and to rib cage fixation, thereby optimizing the ventilatory action of the diaphragm.

Spontaneous and reflexly evoked laryngeal abductor and adductor muscle activity of cat

Abstract A study of spontaneous and reflexly evoked activity of laryngeal abductor (posterior cricoarytenoid) and adductor (thyroarytenoid and lateral cricoarytenoid) muscles was carried out in cats anesthetized with chloraloseurethane or made decerebrate and supplemented with ketamine HCl. The posterior cricoarytenoid muscle was active largely during inspiration but showed tonic activity throughout the respiratory cycle; the thyroarytenoid and lateral cricoarytenoid muscles were rhythmically active during expiration. Anesthetic amounts of pentobarbital abolished adductor rhythmicity and enhanced cyclic inspiratory activity of the abductor muscle. Hyperventilation increased the tonic adductor muscle activity while diminishing abductor muscle activity prior to resolution of apnea. Glossopharyngeal (epipharyngeal branch) and superior laryngeal nerve stimulation evoked chiefly excitatory effects on adductors and largely and attenuating effect on the abductor during inspiration. Stimulation of caudal intercostal nerves caused similar effects but to a lesser degree. Peripheral phrenic nerve stimulation during inspiration facilitated reflex abductor muscle activity whereas such stimulation during expiration facilitated reflex adductor muscle activity. The collective evidence further supports the conclusion that the larynx has a dual function: that of a respiratory organ (widening of the glottis during inspiration and its narrowing during expiration) and of a guardian of the lower respiratory tract from invading foreign matter (reflex sphincter action of contracting adductor muscles with relaxation of the abductor).

An electrophysiological analysis of sleep and respiration of rats breathing different gas mixtures: Diaphragmatic muscle function

The effects of breathing 21% O2, 21% O2 + 5% CO2, 10% O2 + 4% CO2 and 10% O2 on the sleep-waking rhythm, respiratory rate, diaphragmatic EMG, inspiratory (Ti) and expiratory (Te) times were studied in rats. They carried chronically implanted electrodes to permit polygraphic recordings of the ECoG, EOG and dorsal neck and integrated diaphragmatic EMG activity. Average respiratory rates, independent of state of consciousness varied depending on the gas mixture breathed. Sleep-waking times, expressed as percentages, were determined as a function of the gas mixture breathed. Oxygen deficiency caused PS deprivation which was partially alleviated by the addition of 4% CO2. Diaphragmatic EMG activity decreased during PS when rats breathed gas mixtures rich in CO2 but increased when they breathed 10% O2. In general, at a given frequency of breathing, Ti was shorter during PS than during SWS except when rats breathed 10% O2. It is concluded that: (1) regardless of the state of consciousness hypoxia is a more potent stimulus of respiratory rate than hypercapnia, (2) diaphragmatic effort is reduced when rats breathe CO2 enriched gas mixtures but is increased by hypoxia due to changes in upper airway resistance, and (3) low O2 content of an inspired gas disrupts the inspiratory and expiratory off-switch mechanisms, this disruption being prevented by the addition of CO2.

Proprioceptive, chemoreceptive and sleep state modulation of expiratory muscle activity in the rat

The purpose of this study was to assess the respiratory and tonic activity of the abdominal muscles and the postinspiratory activity of the diaphragm (stage 1 expiration) in rats during sleep while they breathed air, hypercapnic, and hypoxic gas mixtures. ECoG and neck EMG recordings enabled the differentiation to be made between nonrapid eye movement sleep (nREMS) and rapid eye movement sleep (REMS). EMGs of the rectus abdominis, internal and external oblique, and diaphragm muscles were displayed on a CRT and polygraph. During nREMS the rectus abdominis showed no respiratory activity, whereas the oblique muscles showed activity confined to stage 2 expiration. This activity was modulated by proprioceptive (sleep postures) and chemoreceptive activation (5% CO2 in air and 10-12% O2 in nitrogen): tonic activity was not consistently affected by such inputs. During REMS tonic activity disappeared, whereas phasic activity either remained unchanged or was abolished. If phasic activity ceased it could reappear periodically during the same REMS epoch. While breathing air, rats in nREMS showed postinspiratory diaphragmatic activity which was sustained or slightly increased while breathing a hypoxic gas mixture but was virtually abolished during hypercapnia. In REMS postinspiratory discharges almost disappeared. The data support the conclusion that the diaphragm provides expiratory braking and that the external and internal oblique muscles contribute to active exhalation during nREMS as well as priming the diaphragm for the next inspiration by improving its length-tension relationship. A three-phase neural respiratory pattern generator operates in nREMS: it changes temporarily to a two-phase system while breathing CO2 and during REMS due to the inhibition of the postinspiratory phase.

Analysis of the respiratory role of pharyngeal constrictor motoneurons of cat

Abstract The electromyogram of the middle pharyngeal constrictor muscle and the electroneurogram of the phrenic nerve was recorded in parallel with measurements of tidal inspiration and tracheal airflow in chloralose-urethane anesthetized cats. In eupnea, pharyngeal constrictor exhibited tonic and expiratory phasic activity. Lung inflation inhibited phasic pharyngeal constrictor activity to reveal its tonic component; lung deflation eliminated both phasic and tonic components. Vagotomy eliminated these static lung volume effects and increased spontaneous cyclic activity. Hypocapnia abolished phasic activity and unmasked its tonic component in vagotomized cats. Hypercapnia increased cyclic pharyngeal constrictor activity (vagi intact or cut). Weak superior laryngeal or glossopharyngeal nerve stimulation had little or no effect during inspiration; during expiration, superior laryngeal nerve stimulation evoked a short lasting attenuating effect, and glossopharyngeal nerve stimulation exerted a long lasting blocking effect. (vagi intact or cut). Pharyngeal constrictor, like laryngeal adductors, is classified as an expiratory resisting type muscle. Pharyngeal constrictor reduces dead space during hypercapnia thereby promoting the exodus of CO 2 . Comparison of vagal and other respitatory motoneurons reveals some special features of neural control of respiration heretofore overlooked.

Respiratory functions of the inferior pharyngeal constrictor and sternohyoid muscles during sleep

We studied the respiratory activity of the inferior pharyngeal constrictor and sternohyoid muscles of the rat during non-rapid eye movement (non-REM) and REM sleep. Each animal carried chronically implanted electrodes for recording the integrated EMG activity of respiratory muscles as well as the electrocorticogram (ECoG) and postural tone (dorsal neck EMG). The latter permitted polygraphic identification of sleep states. Curled up postures enhanced inspiratory activity of both upper airway muscles during non-REM sleep, an effect which CO2 breathing failed to augment except in the well curled up position. Hypoxia reduced their activity. During REM sleep, the inferior pharyngeal constrictor and sternohyoid muscles retained their inspiratory activity. No tonic activity could be detected in either muscle. We conclude that the inferior pharyngeal constrictor and sternohyoid muscles safeguard upper airway patency in the two main sleep states.

Analysis of the respiratory role of intrinsic laryngeal motoneurons of cat

Abstract The electromyographic activity of the posterior cricoarytenoid, thyroarytenoid, lateral cricoarytenoid muscles, and the phrenic electroneuronogram were recorded along with measurements of tidal inspiration and trachael airflow in chloralose-urethane anesthetized cats. In eupnea, the posterior cricoarytenoid exhibited tonic and inspiratory phasic activity; the other two muscles showed only expiratory bursts of activity. Lung inflation inhibited phasic posterior cricoarytenoid activity to reveal a tonic component; activity of the other two was inhibited. Lung deflation increased activity in all three. Vagotomy abolished static volume effects and increased spontaneous cyclic activity of laryngeal muscles. Hypercapnia increased posterior cricoarytenoid activity, but decreased thyroarytenoid activity; these effects were enhanced after vagotomy. Hering-Breuer reflexes curb intrinsic laryngeal motoneuronal excitability. The hypercapnic effect on laryngeal motoneurons promotes the egress of CO2 by widening the glottis due to both facilitation of abduction and attenuation of adduction.

The labile respiratory activity of ribcage muscles of the rat during sleep.

1. Sleep‐waking states of chronically implanted rats were identified polygraphically while recording the integrated electromyogram (e.m.g.) of extrinsic (scalenus medius and levator costae) and intrinsic (external and internal interosseous intercostal and parasternal) muscles of the thoracic cage. Rats breathed air, air enriched in CO2 (5%) or air deficient in O2 (10% O2 in N2) and were free to adopt any desired posture. 2. In non‐rapid eye movement (non‐r.e.m.) sleep, the scalenus medius and intercostal muscles of the cephalic spaces were always inspiratory; intercostal muscles of the mid‐thoracic spaces were commonly expiratory while the more caudal ones were only occasionally expiratory. Expiratory activity, when present in quiet wakefulness, extended for a variable period of time into non‐r.e.m. sleep and always disappeared in r.e.m. sleep regardless of the ribcage muscle under study. 3. Inspiratory activity, when present in non‐r.e.m. sleep, was unaffected, partially attenuated or abolished at entry into r.e.m. sleep. The peak integrated e.m.g. activity of ribcage muscles was measured as a function of posture, gas mixture breathed and ribcage site: (a) the greater the degree of curled‐up posture, the greater the respiratory activity of scalenus medius, an effect augmented by CO2 but depressed by hypoxia, and (b) the more caudally placed ribcage muscles exhibited respiratory activity which was essentially unaffected by posture and gas mixture inspired. 4. The presence or absence of tonic activity in ribcage respiratory muscles during non‐r.e.m. sleep was unrelated to posture. When tonic activity was present, it always disappeared in r.e.m. sleep. When expiratory activity was present in non‐r.e.m. sleep, it too always disappeared in r.e.m. sleep. Inspiratory activity present in non‐r.e.m. sleep was variably affected at entry into r.e.m. sleep; it was unchanged, partially attenuated or abolished. 5. It is concluded that thoracic cage muscles exhibit marked variability in their respiratory activity depending on posture, sleep‐waking states and gas mixture breathed. It is postulated that the presence of tonic and/or expiratory activity in ribcage muscles during non‐r.e.m. sleep reflects an increase in functional residual capacity (F.R.C.).

Spontaneous and reflexly evoked activity in pharyngeal, laryngeal, and phrenic motoneurons of cat

Abstract Experiments were performed on spontaneously breathing cats that were anesthetized with chloralose-urethane in order to study changing excitability in pharyngeal, laryngeal, and phrenic motoneurons. Pharyngeal constrictor motoneurons chiefly were spontaneously active during expiration; the stylopharyngeus and certain motoneurons serving the superior pharyngeal constrictors were inactive during either phase of tidal respiration. While confirming work of previous investigators that feeble glossopharyngeal (epipharyngeal branch) and superior laryngeal nerve stimulation transiently attenuated on-going phrenic nerve activity, such evoked afferent volleys excited laryngeal motoneurons independent of phases of respiration. On-going pharyngeal constrictor nerve activity (expiration) was attenuated for 20–50 msec by superior laryngeal nerve stimulation and for 100–300 msec by glosso-pharyngeal nerve stimulation. Intercostal nerve stimulation evoked occasional pharyngeal constrictor nerve discharges but more commonly attenuated its spontaneous nerve activity. Stylopharyngeus and an unknown portion of the superior pharyngeal constrictor motoneurons were excited by afferent volleys evoked in glossopharyngeal and superior laryngeal nerves. These aforementioned changes in excitability were unaffected by total neuromuscular paralysis while maintaining artificial respiration. It is concluded that certain pharyngeal constrictor motoneurons of the cat are spontaneously active during tidal expiration, are under central respiratory control, and that their intermittence along with those serving the diaphragm and larynx by modest afferent volleys in the ninth and tenth cranial nerves stage the act of swallow or expulsive reflexes, or both.

An analysis of reflex changes in excitability of phrenic, laryngeal, and intercostal motoneurons

Abstract An investigation was carried out in anesthetized cats to ascertain whether self-excitation of phrenic motoneurons is a specific or generalized reflex mechanism for motoneurons allied to respiration. Whereas stimulation of only caudal intercostal nerves evoked discharge of phrenic motoneurons (intercostal-to-phrenic reflex), stimulation of all intercostal nerves elicited discharges in the recurrent laryngeal nerve (intercostal-to-recurrent laryngeal reflex). Weak superior laryngeal nerve stimulation provoked short-latency discharges in the recurrent laryngeal nerve but inhibited on-going inspiratory activity in phrenic and external intercostal motoneurons. In the presence of self-excitation of phrenic motoneurons (phrenophrenic system), there was concomitant excitation of laryngeal motoneurons. In contrast, when self-excitation of laryngeal motoneurons occurred (laryngolaryngeal system) there was concomitant inhibition of inspiratory activity (phrenic and external intercostal motoneurons). Paired shocks delivered to superior laryngeal and intercostal nerves while recording from phrenic, recurrent laryngeal, and intercostal nerves failed to reveal convergent interaction. It is concluded that self-excitation is a generalized reflex mechanism for certain motoneurons allied to respiration.