本實驗以Wistar品系大白鼠為材料，動物經麻醉後，進行氣管與股動靜脈插管，以gallamine triethiodide麻痺，接人工呼吸器。分離膈神經，舌下神經、喉返神經以及喉返神經內收支與外展支。將人工呼吸器的呼氣端，藉由PE管安置於水面下3公分，使呼氣末壓力（positive end-expired pressure; PEEP）為 3 cmH2O作為對照，實驗時，以所測氣管壓力代表氣道壓力變化，將氣道壓力從對照降低到PEEP= 0 cmH2O，再進一步降低到連續氣道負壓（continuous negative airway pressure; CNAP）或呼氣末負壓（negative end-expired pressure; NEEP）為 -1與 -3 cmH2O，然後，升高氣道壓力為PEEP= 6 cmH2O，研究這些不同氣道壓力（6、0、-1、-3 cmH2O）對喉返神經與舌下神經活動的影響。結果顯示，氣道負壓會引起血壓升高，呼吸頻率加快，膈神經立即反應沒有明顯變化，喉返神經內收支活動增強、活動時間延長，因而轉變為連續性活動，相對地，喉返神經外展支活動高度沒有明顯變化，但是活動開始時間延後，這種反應雖然與興奮C纖維所引起的反應相似但卻無關，舌下神經活動高度降低，且活動開始時間延後。PEEP等於 6 cmH2O時，血壓降低，呼吸頻率降低，膈神經立即反應也是沒有明顯變化，喉返神經內收支與外展支活動降低，舌下神經活動高度無明顯改變，僅活動開始時間提前。喉返神經內收支與舌下神經活動的反應，可能是增強上呼吸道的阻力，以留住肺裡的氣體，顯示可能與防禦機制以維持肺體積的大小有關。

To study the effect of changes in airway pressure on upper airway motor nerve activities, adult rats of Wistar were used. The rat was anesthetized with urethane, paralyzed, and artificially ventilated. Catheterization was performed in the trachea, femoral artery and vein. The phrenic, recurrent laryngeal (RLN), and hypoglossal nerves, as well as the abducent (Abd) and adducent (Add) branches of the RLN were separated and their activities were recorded under normocapnia in hyperoxia. The outlet of the ventilator was placed under water surface with a deep of 3 cm such that a 3-cmH2O positive end-expired pressure (PEEP) was obtained to be used as the control. The airway pressure measured by the tracheal pressure (TP) was decreased to a level of continuous negative airway pressure (CNAP) and of negative end-expired pressure (NEEP) at -1 and -3 cmH2O with a water trap system, and then increased to PEEP at 6 cmH2O. The observed results showed that blood pressure and respiratory frequency were increased, and phrenic burst was not changed in immediate response to CNAP and PEEP, and that activity of the entire RLN was significantly increased during inspiratory and expiratory period. Data from the recording of the intralaryngeal branches showed that activity of the Add RLN was increased and also extended to the inspiratory duration such that it transformed into a tonic discharge pattern with CNAP and NEEP. Activity of the hypoglossal nerve was significantly and immediately decreased with CNAP and NEEP. Increase in PEEP at 6 cmH2O produced a decrease of blood pressure and respiratory frequency without conspicuous change in phrenic burst. Activity of the Add RLN and Abd RLN was decreased. There was no change in hypoglossal activity but displayed an advancement in onset. All these reflexive responses were totally abolished after bilateral vagotomy. These results showed that airway resistance may be increased due to the increase of activity of the Add RLN and the decrease of hypoglossal discharge and that this increase in airway resistance may benefit for the maintenance of lung volume.

Aronson RM, Onal EN, Carley DW, Lopata M. Upper airway and respiratory muscle responses to continuous negative airway pressure. J Appl Physiol 66: 1373-1382, 1989.
Ashton JH, Cassidy SS. Reflex depression of cardiovascular function during lung inflation. J Appl Physiol 58: 137-145, 1985.
Bartlett D. Jr. Respiratory functions of the larynx. Physiol Rev 69: 33-57, 1989.
Badier M, Hammes Y, Ormero-Coloer P, Lemerre C. Tonic activity in inspiratory muscles and phrenic motoneurons by stimulation of vagal afferents. J Appl Physiol 66: 1613-619, 1989.
Bergren DR, Peterson DF. Identification of vagal sensory receptors in the rat lung: are there subtypes of slowly adapting receptors? J Physiol 464: 6814-698, 1993.
Belvisi MG. Sensory nerves and airway inflammation: role of A delta and C-fibres. Pulm Pharmacol Ther 16: 1-7, 2003.
Carley DW, Pavlovic S, Malis M, Knezevic N, Saponjic J, Li C. Radulovacki M. C-fiber activation exacerbates sleep-disordered breathing in rats. Sleep Breath 8: 147-154, 2004.
Chen HH, Lee BP, Kou YR. Mechanisms underlying stimulation of rapidly adapting receptors during pulmonary air embolism in dogs. Repir Physiol 109: 1-3, 1997.
Coleridge HM, Coleridge JC. Reflexes evoked from tracheobronchial tree and lungs. In: Cherniak NS and Widdicombe JG, Handbook of Physiology: The Respiratory System. Control of Breathing, part 1, vol. II. Bethesda, MD: Am. Physiol. Soc., sect. 3, vol. II, pt. 1, chapt. 12, p. 395–429, 1986.
Davies A, Roumy M. The effect of transient stimulation of lung irritant receptors on the pattern of breathing in rabbits. J Physiol 324: 389-401, 1982.
Ezure K, Tanaka I. Lung inflation inhibits rapidly adapting receptor relay neurons in the rat. NeuroReport 11: 1709-1712, 2000.
Ezure K, Tanaka I. GABA, in some cases together with glycine, is used as the inhibitory transmitter by pump cells in the Hering-Breuer reflex pathway of the rat. Neuroscience 127: 409-417, 2004.
Fukuda Y, Honda Y. Differences in respiratory neural activities between vagal (superior laryngeal), hypoglossal, and phrenic nerves in the anesthetized rat. Jpn J Physiol 32: 387-398, 1982.
Glick G, Wechsler AS, Epstein SE. Reflex cardiovascular depression produced by stimulation of pulmonary stretch receptors in the dog. J Clin Invest 48: 467-473, 1969.
Green JF, Kaufman MP. Pulmonary afferent control of breathing as end-expiratory lung volume decreases. J Appl Physiol 68: 2186-2194, 1990.
Gunawardena S, Ravi K, Longhurst JC, Kaufman MP, Ma A, Bravo M, Kappagoda CT. Responses of C fiber afferents of the rabbit airways and lungs to changes in extra-vascular fluid volume. Respir Physiol & Neurobiol 132: 239-251, 2002.
Hargreaves M, Ravi KC, Kappagoda T. Effect of bradykinin on respiratory rate in anaesthetized rabbits; role of rapidly receptors. J Physiol 468: 501-513, 1993.
Higenbottam T. Narrowing of glottis opening in humans associated with experimentally induced bronchoconstriction. J Appl Physiol 49: 403-407, 1980.
Ho CY, Gu Q, Lin YS, Lee LY. Sensitivity of vagal afferent endings to chemical irritants in the rat lung. Respir Physiol 127(2-3), 113-124, 2001.
Hwang JC, St. John WM, Bartlett DJr. Respiratory-related hypoglossal nerve activity: influence of anesthetics. J Appl Physiol 55: 785-792, 1983.
Hwang JC, St. John WM. Alterations of hypoglossal motoneuronal activities during pulmonary inflations. Exp Neurol 97: 615-625, 1987.
Kaczynska K, Szereda-Przestaszewska M. Superior laryngeal nerve section abolishes capsaicin evoked chemoreflex in anaesthetized rats. Acta Neurobiol Exp (Warsz) 62: 19–24, 2002.
Kappagoda CT, Skepper JN, McNaughton L, Siew EE, Navaratnam V. Morphology of presumptive rapidly adapting receptors in the rat bronchus. J Anat 168: 265-276, 1990.
Kaufman MP, Coleridge HM, Coleridge JC, Baker DG. Bradykinin stimulates afferent vagal C-fibers in intrapulmonary airways of dogs. J Appl Physiol 48: 511-517, 1980.
Knowlton GC, Larrabee MG. A unitary analysis of pulmonary volume receptors. Am J Physiol 147: 100–114, 1946.
Kubin, L., Alheid, G.F., Zuperku, E.J., & McCrimmon, D.R. Central pathways of pulmonary and lower airway vagal afferents. J Appl Physiol 101: 618-627, 2006.
Lai CJ, Kou YR. Stimulation of pulmonary rapidly adapting receptors by inhaled wood smoke in rats. J Physiol 508: 597-607, 1998.
Lara JP, Dawid-Milner MS, Lopez MV, Montes C, Spyer KM, Gonzalez-Baron S. Laryngeal effects of stimulation of rostral and ventral pons in the anaesthetized rat. Brain Res 934: 97–106, 2002.
Lee KZ, Fuller DD, Lu IJ, Lin JT, Hwang JC. Neural drive to tongue protrudor and retractor muscles following pulmonary C-fiber activation. J Appl Physiol 102: 434–444, 2007a.
Lee KZ, Fuller DD, Tung LC, Lu IJ, Ku LC, Hwang JC. Uncoupling of upper airway motor activity from phrenic bursting by positive end-expired pressure in the rat. J Appl Physiol 102: 878-889, 2007b.
Lee KZ, Fuller DD, Lu IJ, Ku LC, Hwang JC. Pulmonary C-fiber receptor activation abolishes uncoupled facial nerve activity from phrenic bursting during positive end-expired pressure in the rat. J Appl Physiol. 104: 119-29, 2008.
Lin YS, Kou YR. Reflex apneic response evoked by laryngeal exposure to wood smoke in rats. J Appl Physiol 83: 723–730, 1997.
Lopes J, Muller NL, Bryan MH, Bryan AC. Importance of inspiratory muscle tone in maintenance of FRC in the newborn. J Appl Physiol Physiol 51: 830-834, 1981.
Lorino AM, Hamoudi K, Lofaso F, Dahan E, Mariette C, Harf A, Lorino H. Effects of continuous negative airway pressure on lung volume and respiratory resistance. J Appl Physiol 87: 605–610, 1999.
Lu IJ, Ku LC, Lin JT, Lee KZ, Hwang JC. Pulmonary C-fiber activation enhances respiratory-related activities of the recurrent laryngeal nerve in rats. Chin J Physiol 45: 143–154, 2002.
Lu IJ, Lee KZ, Lin JT, Hwang JC. Capsaicin administration inhibits the abducent branch but excites the thyroarytenoid branch of the recurrent laryngeal nerves in the rat. J Appl Physiol 98: 1646-1652, 2005.
Lu IJ, Lee KZ, Hwang JC. Capsaicin-induced activation of pulmonary vagal C-fibers produces reflex laryngeal closure in the rat. J Appl Physiol 101: 1104-1112, 2006.
Meessen NEL, Grinten CPM, Luijendijk SCM, Folgering HTM. Continuous negative airway pressure increases tonic activity in diaphragm and intercostals muscles in humans. J Appl Physiol 77: 1256-1262, 1994.
Muller N, Bryan AC, Zamel N. Tonic activity as a cause of hyperinflation in histamine-induced asthma. J Appl Physiol 49: 869-874, 1980.
Matshmoto S, Ikeda M, Nishikawa T, Yoshida S, Tanimoto T. Excitatory Mechanism of Deflationary Slowly Adapting Pulmonary Stretch Receptors in the Rat Lung. J Pharmacol Expt Ther 300: 597-604, 2002.
Mutoh T, Bonham AC, Joad JP. Substance P in the nucleus of the solitary tract augments bronchopulmonary C fiber reflex output. Am J Physiol 279: R1215-1223, 2000.
Pack AI, Delaney RG. Response of pulmonary rapidly adapting receptors during lung inflation. J Appl Physiol 55: 955-963, 1983.
Paintal A S. Mechanism of stimulation of type J pulmonary receptors. J Physiol 203: 511-532, 1969.
Poliacek I, Stransky A, Jakus J, Barani H, Tomori Z, Halasova E. Activity of the laryngeal abductor and adductor muscles during cough, expiration and aspiration reflexes in cats. Physiol Res 52: 749-762, 2003.
Proctor DF. The upper airways. II. The larynx and trachea. Am Rev Respir Dis 115: 315-342, 1977.
Ravi K, Singh M, Julka DB. Properties of rapidly adapting receptors of the airways in monkeys (Macaca mulatta). Respir Physiol 99: 51-62, 1995.
Remmers JE, deGroot WJ, Sauerland EK, Anch AM. Pathogenesis of upper airway occlusion during sleep. J Appl Physiol 44: 931-938, 1978.
Richardson CA, Herbert DA, Mitchell RA. Modulation of pulmonary stretch receptors and airway resistance by parasympathetic efferents. J Appl Physiol 57: 1842-1849, 1984.
Sammon M, Romaniuk JR, Bruce EN. Role of deflation- sensitive feedback in control of end-expiratory volume in rats. J Appl Physiol 75: 902-911, 1993.
Sant'Ambrogio G, Widdicombe J. Reflexes from airway rapidly adapting receptors. Respir Physiol 125: 33-45, 2001.
Schelegle ES, Green JF. An overview of the anatomy and physiology of slowly adapting pulmonary stretch receptors. Respir Physiol 125: 17-31, 2001.
Sériés F, Marc I. Influence of lung volume dependence of upper airway resistance during continuous negative airway pressure. J Appl Physiol 77: 840-844, 1994.
Shiba K, Satoh I, Kobayashi N, Hayashi F. Multifunctional laryngeal motoneurons: an intracellular study in the cat. J Neurosci 19: 2717-2727, 1999.
Sellick H, Widdicombe JG. Vagal deflation and inflation reflexes mediated by lung irritant receptors. Q J Exp Physiol 55: 153-163, 1970.
Stransky A, Szereda-Przestaszewska M, Widdicombe JG. The effects of lung reflexes on laryngeal resistance and motoneurone discharge. J Physiol 231: 417-438, 1973.
Sun QJ, Berkowitz RG, Pilowsky PM. GABAA mediated inhibition and post-inspiratory pattern of laryngeal constrictor motoneurons in rat. Respir Physiol Neurobiol 162: 41-47, 2008.
Tsai CY, Lee KZ, Lu IJ, Hwang JC. Neural mechanism of glottal closure evoked by anandamine in the rat (Abstract). FASEB J 21: 1b590, 2007.
Tsubone H. Characteristics of vagal afferent activity in rats: Three types of pulmonary receptors responding to collapse, inflation, and deflation of the lung. Exptl Neurol 92: 541-552, 1986.
Widdicombe J. Receptors in the trachea and bronchi of the cat. J Physiol 123: 71-104, 1954.
Widdicombe J . Airway receptors. Respir Physiol 125: 3-15, 2001.
Widdicombe J. Functional morphology and physiology of pulmonary rapidly adapting receptors (RARs). Anat Rec A Discov Mol Cell Evol Biol 270: 2-10, 2003.
Zhou D, Huang Q, St John WM, Bartlett DJr. Respiratory activities of intralaryngeal branches of the recurrent laryngeal nerve. J Appl Physiol 67: 1171-1178, 1989.