Comprehensive Physiology Wiley Online Library

Clearance Mechanisms in the Respiratory Tract

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 Conducting Airways
1.1 Ciliated Cells
1.2 Goblet Cells and Mucus
1.3 Serous Cells and Water
1.4 Transport and Clearance Dynamics
2 Fluid Absorption and Mucociliary Clearance
2.1 null
2.2 Other Physical Mechanisms
2.3 Leukocyte Recruitment and Endocytosis in Conducting Airways
3 Alveolar Ducts and Alveoli
3.1 Physical Removal
3.2 Solubility and Permeability
3.3 Digestion
3.4 Retention
4 Implications for Environmental Diseases
4.1 Pulmonary
4.2 Systemic Disorders
Figure 1. Figure 1.

The two distal zones of the lung appear to have different modes of clearance of particles. The alveolar zone depends upon digestion by macrophages and clearance via lymph and blood. The terminal bronchioles have ciliary clearance of particles attached to mucus and recruitment of polymorphonuclear leukocytes to their lumens.

From Mangos and Talamo (editors) with permission of Academic Press
Figure 2. Figure 2.

Several tall columnar ciliated cells are attached to the basal lamina and extend to the bronchial lumen. Interspersed are secretory cells, some of which contain mucous vacuoles, Golgi apparatus, and endoplasmic reticulum. A few flattened basal cells with scanty cytoplasm form an incomplete layer beneath the columnar cells. The lamina propria beneath the basal lamina has collagen and elastin fibrils and fibroblasts. Not shown are the capillaries and smooth muscle of the deeper lamina propria. Hamster bronchus fixed with OsO4, stained with uranyl acetate and lead citrate. x 4,400.

Figure 3. Figure 3.

Pseudostratified ciliated columnar epithelium of the human trachea. In addition to basal cells there are intermediate cells which differentiate into either ciliated or goblet cells. They form an incomplete layer of less differentiated cells.

From Rhodin
Figure 4. Figure 4.

The ciliated cell as it appears by light microscopy, upper left, contrasts with a drawing based on electron microscopy. The proximity of mitochondria to rootlets and basal bodies of cilia is important for ciliary energetics. The filaments of the ciliary stalk have been shown to be microtubules. RNP, ribosomes.

From Satir by permission of Scientific American.
Figure 5. Figure 5.

Resolution of this cross‐section of flagellar axoneme of a blowfly sperm has been enhanced by rotational and translational image reinforcement of positive and negative images which show important details. The nine pairs of doublet microtubules (mt A and B) with their side arms appear to be linked to the central sheath. It has a wide y portion opposed by a narrow x portion. The x portion is linked to doublet tubule 1. Details of transitional links are still only tentative as are details of the coarse outer fibers.

Adapted from Warner
Figure 6. Figure 6.

Drawing of a goblet cell based on electron photomicrographs and biochemical studies. It shows the synthesis pathway of mucin from precursors entering the cell from the blood (1), proteins assembled from amino acids on ribosomes (2) move up rough endoplasmic reticulum (3) to enter Golgi saccules (4). Meanwhile simple sugars enter the saccules to combine with proteins by glycosylation and sulfate is added (5). The saccules of glycoprotein are transformed and hydrated into globules of mucus (6). Mucin lobules move toward the cell apex (7) for subsequent release from the cell into the lumen (8).

From Neutra and Leblond by permission of Scientific American.
Figure 7. Figure 7.

Stages in formation and rupture of a meniscus of mucus in a small airway reflect the viscosity of the mucus. Retraction and thinning occur in the third dimension as well.

From Besarab & Litt
Figure 8. Figure 8.

This section from a small bronchiole shows cuboidal ciliated cells and secretory cells with apical knobs packed with small Golgi vesicles and occasional lysosomes. Such cells resemble the serous or albuminous cells of bronchial glands and salivary glands and presumably secrete watery serous fluids. Osmium tetroxide‐fixed hamster lung, stained with uranyl acetate and lead citrate. x 5,250.

Figure 9. Figure 9.

Polymorphonuclear leukocyte recruitment from capillaries into the lamina propria (1) and across the epithelium (2) into airway lumens (3) appears to be an important early response of the lung and upper airways to particle deposition. The leukocyte in the center is crossing the basal lamina. The one on the left is in the intercellular space. Vacuoles often appear in the cytoplasm when the leukocytes reach the lumenal surface. Osmium tetroxide fixation, uranyl acetate, and lead citrate. x 8,500.

Figure 10. Figure 10.

This latticework pattern represents a phospholipid array on the alveolar surface of an epithelial cell fixed by osmium tetroxide. This is an artifact due to surface forces during fixation and dehydration.



Figure 1.

The two distal zones of the lung appear to have different modes of clearance of particles. The alveolar zone depends upon digestion by macrophages and clearance via lymph and blood. The terminal bronchioles have ciliary clearance of particles attached to mucus and recruitment of polymorphonuclear leukocytes to their lumens.

From Mangos and Talamo (editors) with permission of Academic Press


Figure 2.

Several tall columnar ciliated cells are attached to the basal lamina and extend to the bronchial lumen. Interspersed are secretory cells, some of which contain mucous vacuoles, Golgi apparatus, and endoplasmic reticulum. A few flattened basal cells with scanty cytoplasm form an incomplete layer beneath the columnar cells. The lamina propria beneath the basal lamina has collagen and elastin fibrils and fibroblasts. Not shown are the capillaries and smooth muscle of the deeper lamina propria. Hamster bronchus fixed with OsO4, stained with uranyl acetate and lead citrate. x 4,400.



Figure 3.

Pseudostratified ciliated columnar epithelium of the human trachea. In addition to basal cells there are intermediate cells which differentiate into either ciliated or goblet cells. They form an incomplete layer of less differentiated cells.

From Rhodin


Figure 4.

The ciliated cell as it appears by light microscopy, upper left, contrasts with a drawing based on electron microscopy. The proximity of mitochondria to rootlets and basal bodies of cilia is important for ciliary energetics. The filaments of the ciliary stalk have been shown to be microtubules. RNP, ribosomes.

From Satir by permission of Scientific American.


Figure 5.

Resolution of this cross‐section of flagellar axoneme of a blowfly sperm has been enhanced by rotational and translational image reinforcement of positive and negative images which show important details. The nine pairs of doublet microtubules (mt A and B) with their side arms appear to be linked to the central sheath. It has a wide y portion opposed by a narrow x portion. The x portion is linked to doublet tubule 1. Details of transitional links are still only tentative as are details of the coarse outer fibers.

Adapted from Warner


Figure 6.

Drawing of a goblet cell based on electron photomicrographs and biochemical studies. It shows the synthesis pathway of mucin from precursors entering the cell from the blood (1), proteins assembled from amino acids on ribosomes (2) move up rough endoplasmic reticulum (3) to enter Golgi saccules (4). Meanwhile simple sugars enter the saccules to combine with proteins by glycosylation and sulfate is added (5). The saccules of glycoprotein are transformed and hydrated into globules of mucus (6). Mucin lobules move toward the cell apex (7) for subsequent release from the cell into the lumen (8).

From Neutra and Leblond by permission of Scientific American.


Figure 7.

Stages in formation and rupture of a meniscus of mucus in a small airway reflect the viscosity of the mucus. Retraction and thinning occur in the third dimension as well.

From Besarab & Litt


Figure 8.

This section from a small bronchiole shows cuboidal ciliated cells and secretory cells with apical knobs packed with small Golgi vesicles and occasional lysosomes. Such cells resemble the serous or albuminous cells of bronchial glands and salivary glands and presumably secrete watery serous fluids. Osmium tetroxide‐fixed hamster lung, stained with uranyl acetate and lead citrate. x 5,250.



Figure 9.

Polymorphonuclear leukocyte recruitment from capillaries into the lamina propria (1) and across the epithelium (2) into airway lumens (3) appears to be an important early response of the lung and upper airways to particle deposition. The leukocyte in the center is crossing the basal lamina. The one on the left is in the intercellular space. Vacuoles often appear in the cytoplasm when the leukocytes reach the lumenal surface. Osmium tetroxide fixation, uranyl acetate, and lead citrate. x 8,500.



Figure 10.

This latticework pattern represents a phospholipid array on the alveolar surface of an epithelial cell fixed by osmium tetroxide. This is an artifact due to surface forces during fixation and dehydration.

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Kaye H. Kilburn. Clearance Mechanisms in the Respiratory Tract. Compr Physiol 2011, Supplement 26: Handbook of Physiology, Reactions to Environmental Agents: 243-262. First published in print 1977. doi: 10.1002/cphy.cp090116