Quiet breathing chiefly uses active inspiration and passive expiration; deeper or more forceful breathing recruits additional muscles and changes their coordination. The vertical, anteroposterior and transverse dimensions of the thorax can all change, and different patterns need not represent different mechanisms of gas exchange.
The respiratory diaphragm is the principal inspiratory muscle. Its contraction lowers and flattens its domes, increases the thoracic vertical dimension and displaces abdominal contents so the relaxed abdominal wall moves outwards. Its lower-rib attachments also contribute to expansion of the costal margin. The phrenic nerves, chiefly C3–C5, provide its motor supply. The external intercostal muscles elevate and stabilize the ribs, while the interchondral or parasternal portions of the internal intercostal muscles also assist inspiration. It would be wrong to give every internal intercostal fibre an exclusively expiratory function.
Elevation of upper ribs and the sternum increases the anteroposterior dimension, the pump-handle component. Outward movement of the lateral portions of lower ribs increases transverse diameter, the bucket-handle component. These are components of oblique rib rotation rather than two mechanically isolated systems. Abdominal or diaphragmatic breathing makes descent of the respiratory diaphragm particularly conspicuous; costal breathing makes chest-wall expansion more conspicuous. Both contribute in ordinary respiration, with their proportions changing with posture, mechanical loading and individual anatomy.
As the thorax expands, pleural pressure becomes more negative and transpulmonary pressure increases. The lungs expand and alveolar pressure briefly falls below atmospheric pressure, admitting air. Relaxation of inspiratory muscles lets elastic recoil reverse the flow in quiet expiration. The abdominal wall and respiratory diaphragm return towards their resting arrangement without requiring active abdominal compression for each quiet breath.[ref]
22.3 The Process of Breathing. Mechanisms of Breathing; Pulmonary Ventilation: pressure changes and inspiratory versus expiratory activity
With deeper inspiration, the scalene muscles help stabilize or elevate the upper ribs, and the sternocleidomastoid muscle can elevate the sternum when the head is fixed. With the upper limbs or shoulder girdle fixed, muscles such as pectoralis minor and serratus anterior can assist expansion in greater effort. Recruitment is graded; an anatomical list of possible accessory muscles does not mean all are active in every ordinary breath.
Active expiration uses the abdominal muscles, including rectus, oblique and transversus components, to compress the abdominal contents and drive the respiratory diaphragm upwards. The interosseous portions of the internal intercostal muscles help depress the ribs. This is important in vigorous exercise and expulsive actions. At rest, elasticity supplies much of the expiratory work; during high demand, active muscle contraction adds to it.
The observable pattern also changes with the physiological task. Exercise increases breath depth and usually frequency. A sigh is an enlarged inspiration followed by expiration; yawning includes a prolonged deep inspiratory movement. Coughing coordinates an inspiration, closure of the glottis, pressure-generating expiratory effort and sudden glottic opening. Speech imposes a controlled prolonged expiration with laryngeal regulation. Swallowing briefly interrupts ordinary airflow. Thus respiratory observation should include phase, depth, frequency and coordination, not merely count which ribs move. Fixed claims that men must breathe abdominally and women must breathe costally overstate a variable tendency. Pregnancy or a change of posture can alter mechanical contributions without replacing the basic pressure mechanism.
The two pneumogastric nerves are the right and left vagus nerves, the tenth cranial nerves: mixed sensory, branchial motor and parasympathetic pathways from the medulla oblongata to the neck, thorax and abdomen. A useful description separates their apparent origin at the surface of the brain from the origins and terminations of their component fibres.
Rootlets emerge in the groove behind the inferior olive, between the glossopharyngeal nerve above and the accessory rootlets below. Branchial motor fibres arise chiefly in the nucleus ambiguus; preganglionic parasympathetic fibres arise in the dorsal motor nucleus of the vagus and, particularly for cardiac control, the nucleus ambiguus. Visceral sensory fibres have their cell bodies chiefly in the inferior vagal ganglion and terminate centrally in the nucleus of the solitary tract; somatic sensory fibres associated with the superior vagal ganglion project to the spinal trigeminal nucleus. Consequently a sensory fibre is not accurately described as originating in a motor nucleus simply because its peripheral nerve emerges nearby.[ref]
Neuroanatomy, Nucleus Ambiguus (StatPearls). Introduction and Structure and Function: branchial motor output and cardiac vagal control
Each vagus nerve leaves the skull through the jugular foramen, with the glossopharyngeal nerve and accessory nerve. It bears the small superior vagal ganglion at the foramen and the larger inferior vagal ganglion below it. Connections include communications with adjacent cranial nerves and the cervical sympathetic nervous system. Traditional descriptions assign fibres joining the vagus nerve from the cranial part of the accessory nerve to the latter peripheral nerve; their modern classification is more nuanced and does not change the peripheral branches being described. In the neck the vagus nerve descends in the carotid sheath, behind and between the internal jugular vein and the internal, then common, carotid artery.
Its meningeal and auricular branches supply small sensory territories of the cranial covering and external ear. Pharyngeal branches contribute to the pharyngeal plexus, supplying much of the soft palate and pharynx; the stylopharyngeus muscle belongs to the glossopharyngeal nerve and the tensor veli palatini muscle to the mandibular division of the trigeminal nerve. The internal branch of the superior laryngeal nerve supplies sensation above the vocal folds; its external branch supplies the cricothyroid muscle. The recurrent laryngeal nerve supplies the other intrinsic laryngeal muscles and sensation below the vocal folds. The right recurrent laryngeal nerve loops beneath the right subclavian artery; the left loops under the aortic arch beside the ligamentum arteriosum. Both return towards the larynx near the groove between trachea and oesophagus.[ref]
Neuroanatomy, Cranial Nerve 10 (Vagus Nerve) (StatPearls). Structure and Function; Vagus Nerve Branches; Muscles
In the thorax, both vagus nerves pass behind the roots of the lungs, contrasting with the anterior course of the phrenic nerves. Cardiac branches join the cardiac plexus; pulmonary branches join the pulmonary plexus. The trunks break up into the oesophageal plexus, then collect principally as an anterior trunk derived from the left vagus nerve and a posterior trunk derived from the right. Through the oesophageal hiatus they reach the stomach; hepatic and coeliac branches distribute to abdominal viscera. Their parasympathetic territory includes foregut and midgut derivatives, extending into the proximal colon, rather than every abdominal and pelvic organ.[ref]
Neuroanatomy, Cranial Nerve 10 (Vagus Nerve) (StatPearls). Structure and Function: thoracic course and oesophageal plexus
This extensive distribution explains why the level of experimental section is indispensable information.The main consequences of section can be organized by the interrupted functions:
- Heart: cutting both cervical vagus nerves removes tonic inhibition and usually increases heart rate. The heart continues beating because its rhythm is intrinsic. Electrical stimulation of the peripheral cut end can slow the heart, whereas stimulation of the central end tests reflex effects through the brainstem. Cutting a pathway and stimulating its intact output are different experiments.
- Breathing: bilateral cervical section commonly produces slower, deeper breaths in experimental mammals. Sensory information from the lungs has been interrupted; the respiratory diaphragm is not directly paralysed, because its motor peripheral nerves are the phrenic nerves. Persistence of breathing establishes that the vagus nerves modify the respiratory rhythm rather than being its sole generator. The subsequently named Hering-Breuer inflation reflex provides a later explanation of one component of this feedback; its formal description belongs to 1868, after this examination.[ref]Reflexes from the lungs and airways: historical perspective. Discussion of the 1868 lung inflation and deflation experiments
- Larynx and lungs: section proximal to the recurrent laryngeal nerves can impair opening and closure of the vocal folds, causing altered voice, obstruction and impaired airway protection. Classical bilateral cervical experiments sometimes ended in death with pulmonary congestion, oedema or aspiration changes. Such outcomes must be distinguished from an immediate necessary arrest of breathing: species, age, lesion level and airway management affect them.
- Stomach: bilateral section can reduce vagally driven gastric acid secretion and disturb gastric motor coordination and emptying. Nevertheless digestion and nutrient absorption are not abolished as chemical possibilities: enteric nervous system activity, local stimuli and hormones remain. Likewise, interruption of gastric sensory fibres does not establish that the entire experience of hunger originates in them.
A discriminating historical experiment would record respiratory movements and heart rate before and after section, distinguish unilateral from bilateral section, and compare an intact airway with an airway bypass. A tracheal cannula can separate obstruction at the glottis from changes in the respiratory rhythm, while the central-versus-peripheral stimulation comparison separates incoming reflex traffic from outgoing motor effects. Later separate-lung recordings confirmed that a change following unilateral section could affect both lungs, supporting a central reflex interpretation rather than simple local paralysis.[ref]
A Study of the Hering-Breuer Reflex. Summary and Conclusions, points 1–4; separate recordings of the two lungs, 1927
The gastric distinction is also borne out by measurements of persistent acidity after surgical vagotomy, despite major reductions in stimulated gastric acid secretion.[ref]Twenty-four hour gastric acidity after vagotomy. Abstract: hourly intragastric sampling, seven patients, four years after operation