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5.2 HUMAN RESPIRATORY TRACT PHYSIOLOGY                                    203
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                   and hydrostatic pressure.  Estimates of daily mucus production range
                   from 7-12 mL/day in healthy individuals to > 100 mL/day in cystic fibro-
                   sis patients.
                      Airway Epithelial Cell Types
                      Conducting airway passages are generally composed of ciliated
                   pseudostratified cuboidal columnar epithelial cells interspersed with
                   basal, brush, and secretory cells (goblet, serous, Clara). Cilia found on
                   ciliated epithelial cell apical surfaces (along the lumen) provide motive
                   force for propelling mucus gel along the airway. Extrathoracic airway
                   surfaces are lined with ciliated epithelium, except for squamous epithe-
                   lium covering the nasal vestibule, nasopharynx, oral cavity, orophar-
                   ynx, and portions of the larynx. Squamous epithelium protects airway
                   surfaces against mechanical impact or shear in areas where relatively
                   large inspired particles usually impact. Also, nasal olfactory surfaces
                   are not lined with ciliated epithelium but covered instead by special
                   sensory cells. These specialized olfactory receptor cells react with in-
                   haled odorant molecules, generating neural signals sent to the olfactory
                   bulbs of the brain that produce a sense of smell. All tracheobronchial
                   surfaces are lined with ciliated epithelium down to the pulmonary air
                   ways. Proximal airway epithelium is thickest and progressively flattens
                   and thins toward the lung parenchyma, gradually transitioning into al-
                   veolar endothelium.
                      Secreting cells found along the conducting airways include nonciliated
                  goblet and serous cells. Goblet cells produce glycoproteins that form droplets
                   or sheets of mucus gel floating on periciliary fluid. Serous cell exudates are
                   believed to include periciliary fluid, various proteins and peptides (including
                   lysozyme and lactoferrin), and protease inhibitors. Periciliary fluid also de-
                   rives from interstitial fluid transudate. Glycosaminoglycans, Jipids, serum
                   proteins, and ions found in ASL appear to originate from all surface epithelial
                  cells and submucosal glands (serous and mucous). The quantity of submit-
                   cosal glands decreases in more distal airways and are absent from pulmonary
                   airways.
                      Microvilli, approximately 2 jjim long, give a "brush-like" appearance as
                   they project from the apical surface of brush cells. These cells contribute to
                   fluid regulation along the luminal surface by absorbing excess periciliary fluid
                  either secreted by neighboring serous cells or transported from distal airways
                   by the mucociliary elevator. Basal cells are progenitors of the other epithelial
                  cells and are the most actively mitotic epithelial cells. Lymphocytes also ap-
                  pear in ASL as either migratory or basal cells.
                      Pulmonary airways are lined with specialized cells generally not found
                  in the conducting airways. Alveolar epithelium, composed of thin sheet-like
                  cells separated from pulmonary capillaries by only a basement membrane,
                   permits easy exchange of gases between alveolar sacs and blood (Fig. 5.17).
                  Secretory Clara and Type II pneumonocyte cells produce surfactant, lipids,
                  and protease inhibitors within the pulmonary airways. Macrophages are
                  scavenger cells that remove microorganisms and particulates depositing
                  along alveolar surfaces.
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