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Lung Cell Biology

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 Basic Plan of the Cell
1.1 Cell Nucleus
1.2 Cytoplasmic Membrane Systems and Granules
1.3 Mitochondria
1.4 Ground Substance and Cytoskeleton
1.5 Plasma Membrane
2 Organization of Lung Cell Population
2.1 Histogenetic Origin of Lung Cells and Tissues
2.2 Differentiation of Functional Zones
2.3 Morphometry of Lung Cell Population
3 Epithelium
3.1 Epithelium of Conducting Airways
3.2 Alveolar Epithelium
4 Vascular Endothelium
4.1 Structure of Capillary Endothelium
4.2 Structure of Arterial and Venous Endothelium
4.3 Metabolic Functions of Endothelial Cells
5 Interstitial Cells
5.1 Cells Related to Connective Tissue Fibers
5.2 Interstitial Cells With Contractile Properties
5.3 Lymphatics and Free Cells
5.4 Nerves
6 Cells of Pulmonary Defense System
6.1 Alveolar and Interstitial Macrophages
6.2 Cells of Immune Defense System
6.3 Granulocytes
7 Mesothelial Cells of Pleura
Figure 1. Figure 1.

Organization of half a type II epithelial cell from human lung (cf. Fig. B). Nucleus is enwrapped by a perinuclear cisterna (PC) made of 2 membranes, which is traversed by nuclear pores (NP), shown (inset, right) at greater magnification; nuclear content is divided into dense heterochromatin (HC) areas and karyoplasm, which contains euchromatin. Cell surface is made of the plasma membrane (PM) with a subjacent ectoplasmic layer of filaments (f) and a fuzzy coat or glycocalyx (arrow) on the outside (inset, left). Cells in epithelia are joined by junctional complexes (J) with terminal bars at the boundary between apical and lateral face; basal face is attached to the basement membrane (BM). Cytoplasm houses a variety of organelles, cf, Collagen fibrils; ER, endoplasmic reticulum; G, Golgi complex; L, lysosomes; LB, lamellar body; M, mitochondrion; mt, microtubule; MV, microvilli. Bars: 1 μm, 0.1 μm (insets).

Figure 2. Figure 2.

Mitochondria (M) and rough endoplasmic reticulum with cisternal space (asterisks) bounded by fine membrane to which ribosomes (arrows) are attached. Some free ribosomes are not attached to membranes. Double membrane of mitochondria with formation of cristae. Plasma cell. Bar, 0.2 μm.

Figure 3. Figure 3.

Golgi‐associated endoplasmic reticulum and lysosomes (GERL) from alveolar type II cell consists of Golgi complex (G) with a stack of cisternae surrounded by vesicles, rough endoplasmic reticulum (ER), and primary lysosomes (L). M, mitochondrion; MV, multivesicular bodies. Bar, 0.2 μm.

Figure 4. Figure 4.

Cytoskeletal 10‐nm filaments (arrows) extend between 2 plasma membranes (PM) in alveolar type I cell. Bar, 0.2 μm.

Figure 5. Figure 5.

A: thin section of cell membrane of erythrocyte with 2 dark staining leaflets (circle). B: freeze‐fracture replica of cleaved membranes of 2 erythrocytes showing that intramembranous particles on inner protoplasmic leaflet (PF) facing the cytoplasm are denser than those on external leaflet (EF). Bars, 0.1 μm.

Figure 6. Figure 6.

Electron micrograph of human ciliated cells. Terminal bar or tight junction (J) appears as lacework of ∼10 rows of densely aligned particles on fracture face of protoplasmic leaflet (PF) and as complementary grooves on external leaflet (EF) of adjacent cell. Tuft of microvilli (V) and base of cilium (C) with ciliary necklace of intramembrane particles (Ne). Bar, 0.2 μm.

Courtesy of E. E. Schneeberger
Figure 7. Figure 7.

Schematic representation of airway wall structure from bronchus to alveolus. EP, epithelial cells; BM, basement membrane; SM, smooth muscle cells; FC, fibrocartilaginous coat.

From Weibel
Figure 8. Figure 8.

Mucous membrane of small human bronchus. A: thin section showing ciliated cell (CC) with cilia (C) and microvilli (arrows), goblet cell (GC) with apical mucous plug (MU), and basal cell (BC). Fibers and fibroblasts (FB) in connective tissue. MP, macrophage. B: scanning electron micrograph of epithelial surface showing ciliary tufts (C) and mucous plug (MU) of goblet cell in process of extrusion. Bars, 5 μm.

Figure 9. Figure 9.

Scanning electron micrograph (A) and longitudinal (B) and transverse (C) thin sections of cilia of human bronchial epithelium. Microvilli (arrows) between cilia and axonemal complex (A) with its different structure in the shaft, neck (Ne), and basal body or kinetosome (K). mt, Microtubule; S, satellite of ciliary kinetosome. D: greater magnification of cross‐sectioned shaft of a normal cilium with dynein arms (arrowhead). E: dynein arms are lacking in a cilium from a patient with immotile cilia syndrome. Bars: 0.5 μm (A‐C); 0.1 μm (D, E).

E, courtesy of J. M. Sturgess
Figure 10. Figure 10.

Scanning electron micrograph of brush cell from small bronchiole of rat lung. Compare brush with cilia (C) and ordinary microvilli (arrow). Bar, 1 μm.

Figure 11. Figure 11.

Basal part of neuroendocrine cell of human bronchiolar epithelium showing part of nucleus (N), mitochondria (M), and secretory vesicles (arrows) with dark core, which are seen at greater magnification in inset. Bars, 0.2 μm.

Figure 12. Figure 12.

Scanning electron micrograph of surface of alveolar epithelium of human lung with two type II epithelial cells (EP2). Arrows, boundary (terminal bar) of type I cell. Asterisk, nucleus of type I cell. Bar, 5 μm.

From Weibel
Figure 13. Figure 13.

A: squamous type I epithelial cell (EP1). B: capillary endothelial cell (EN) from human lung. Nuclei (N) are surrounded by little cytoplasm, which continues into squamous extensions. Circles, fused basement membranes (BM) in barrier parts. Capillary contains neutrophil granulocyte (GR) with 2 lobes of nucleus sectioned (N′). F, fiber; J, intercellular junction. Bars, 1 μm.

Figure 14. Figure 14.

Squamous extensions of endothelial cell (EN) and type I pneumocyte (EP1). Cytoplasm, bounded by plasma membrane similar to that of erythrocyte (EC), contains microfilaments (f), microtubules (mt), and pinocytotic vesicles (arrows). BM, basement membrane. Bar, 0.1 μm.

Figure 15. Figure 15.

Scanning electron micrograph (A) and thin section (B) of type II pneumocyte with microvilli (V) on peripheral zone of apical cell surface. BM, basement membrane; J, intercellular junction; LB, lamellar body; N, nucleus. Detail of this cell is shown in Fig. . Bars, 1 μm.

Figure 16. Figure 16.

Characteristic organelles of type II cell related to surfactant production and storage. Cone‐shaped lamellar bodies from rat lung (A, B) and sequence of multivesicular bodies (C‐E) with stacks of lamellae (arrows) and membrane envelope (arrowheads). A, C, D: thin sections. B, E: freeze‐fracture preparation. Bars, 0.2 μm.

Figure 17. Figure 17.

Release of lamellar body (LB) from type II epithelial cell into surface lining layer, which contains tubular myelin (TM) continuous with the phospholipid surface film (arrows). Bar, 0.2 μn.

Figure 18. Figure 18.

Alveolar brush cells from rat lung. A: scanning electron micrograph showing brush (B) on narrow apical surface bent to the side, whereas cell body is covered by smooth type I cell extension (asterisks). B: thin section shows brush beneath surface film (arrows) and covered by type I cell (asterisks). C: squat brush microvilli contain bundles of filaments (f) that extend deep into cell. Bars: 1 μm (A, B); 0.5 μm (C).

B from Weibel
Figure 19. Figure 19.

Kinetics of transformation of type II to type I cells. Rats exposed to NO2 to induce lung damage were injected with [3H]thymidine at time 0 and killed from 1 h to 14 days later.

From Evans et al.
Figure 20. Figure 20.

Endothelium (EN) and single layer of smooth muscle (SM) from small pulmonary artery of human lung. Thick endothelial cytoplasm and wealth of organelles (inset) comprising mitochondria (M), endoplasmic reticulum (ER), lipid droplet (Li), specific granules (asterisks), microtubules (mt), and many vesicles (arrows). Cross‐sectioned smooth muscle cells show central nucleus, mitochondria, sarcoplasmic reticulum (SR), membrane‐bounded caveolae (arrows), filamentous matter with dense bodies (db), and cell‐to‐cell contacts (circle), cf, Collagen fibrils; el, elastic fibers. Bars, 0.5 μm.

Figure 21. Figure 21.

Fibroblast engaged in fibrogenesis contains much endoplasmic reticulum (ER) with ribosomes and large Golgi complex (G), as well as mitochondria (M). cf, Collagen fibrils. Bar, 0.2 μm.

Figure 22. Figure 22.

Septal fibroblast (FB) ramifies with slim extensions (arrows) into interstitial spaces between capillary endothelium (EN) and alveolar epithelium (EP), following the fiber strands (F). Circles, cytoplasmic areas with condensed filaments spanning the interstitial space crosswise. Bar, 2 μm.

Figure 23. Figure 23.

A: smooth muscle cells from small pulmonary artery of human lung cut longitudinally, db, Dense bodies of filamentous matter; db′, dense bands on plasma membrane; M, mitochondria; N, central elongated nucleus; SR, sarcoplasmic reticulum. Greater magnification shows filaments on longitudinal (B) and transverse (C) sections together with caveolae (arrows) of plasma membrane, which is coated by basement membrane. Bars: 1 μm (A); 0.2 μm (B, C).

Figure 24. Figure 24.

Myofibroblasts from human alveolar septa with bundles of filaments (f) extending obliquely or crosswise between dense insertion spots on plasma membrane, which appears affixed to basement membrane (arrows). Bars, 0.2 μm.

Figure 25. Figure 25.

Capillary pericyte (PC) with extension (arrows) that lies between the endothelial cell (EN) and its basement membrane in its peripheral part. Other processes of same cell make contact with epithelial (EP) basement membrane (arrowheads). Inset, pericyte filaments (f). Bars: 1 μm; 0.1 μm (inset).

Figure 26. Figure 26.

Mast cell from human alveolar septum contains granules (arrows) with scroll‐like substructure. Inset, scroll‐like substructure of mast cell at higher magnification, cf, Collagen fibrils. Bars, 1 μm; 0.1 μm (inset).

Figure 27. Figure 27.

Scanning electron micrograph of human alveolar macrophage on alveolar epithelial surface. Thin advancing cytoplasmic lamella appears partly attached to epithelium. Bar, 5 μm.

From Weibel
Figure 28. Figure 28.

Alveolar macrophage of human lung apposed to epithelium in several places (arrows). Advancing lamella (AL) is free of organelles that are rich in perinuclear cytoplasm. Bar, 2 μm.

Figure 29. Figure 29.

Alveolar macrophage lies beneath thin film of surface lining layer (arrows). Exclusion of organelles from short peripheral cytoplasmic flaps (asterisks). Bar, 1 μm.

Figure 30. Figure 30.

Interstitial macrophage (MP) from human lung contains heterogeneous dark material in phagolysosomes or residual bodies (arrows). Macrophage is in group of free cells with lymphocytes (LC) and plasma cell (PC). Bar, 2 μm.

Figure 31. Figure 31.

A: phagocytosis of erythrocyte (EC) by alveolar macrophage. Ectoplasmic “lips” (asterisks) are pinching off fragment. B: phagolysosome may contain tubular myelin (TM) and lamellar (L) osmiophilic material, possibly the result of enzymatic disassembly of surface lining material. Bars: 1 μm (A); 0.2 μm (B).

Figure 32. Figure 32.

Plasma cell from small rat lymph node contains massive rough endoplasmic reticulum (ER). Golgi complexes (G) are in adjacent plasma cell. Bar, 2 μm.



Figure 1.

Organization of half a type II epithelial cell from human lung (cf. Fig. B). Nucleus is enwrapped by a perinuclear cisterna (PC) made of 2 membranes, which is traversed by nuclear pores (NP), shown (inset, right) at greater magnification; nuclear content is divided into dense heterochromatin (HC) areas and karyoplasm, which contains euchromatin. Cell surface is made of the plasma membrane (PM) with a subjacent ectoplasmic layer of filaments (f) and a fuzzy coat or glycocalyx (arrow) on the outside (inset, left). Cells in epithelia are joined by junctional complexes (J) with terminal bars at the boundary between apical and lateral face; basal face is attached to the basement membrane (BM). Cytoplasm houses a variety of organelles, cf, Collagen fibrils; ER, endoplasmic reticulum; G, Golgi complex; L, lysosomes; LB, lamellar body; M, mitochondrion; mt, microtubule; MV, microvilli. Bars: 1 μm, 0.1 μm (insets).



Figure 2.

Mitochondria (M) and rough endoplasmic reticulum with cisternal space (asterisks) bounded by fine membrane to which ribosomes (arrows) are attached. Some free ribosomes are not attached to membranes. Double membrane of mitochondria with formation of cristae. Plasma cell. Bar, 0.2 μm.



Figure 3.

Golgi‐associated endoplasmic reticulum and lysosomes (GERL) from alveolar type II cell consists of Golgi complex (G) with a stack of cisternae surrounded by vesicles, rough endoplasmic reticulum (ER), and primary lysosomes (L). M, mitochondrion; MV, multivesicular bodies. Bar, 0.2 μm.



Figure 4.

Cytoskeletal 10‐nm filaments (arrows) extend between 2 plasma membranes (PM) in alveolar type I cell. Bar, 0.2 μm.



Figure 5.

A: thin section of cell membrane of erythrocyte with 2 dark staining leaflets (circle). B: freeze‐fracture replica of cleaved membranes of 2 erythrocytes showing that intramembranous particles on inner protoplasmic leaflet (PF) facing the cytoplasm are denser than those on external leaflet (EF). Bars, 0.1 μm.



Figure 6.

Electron micrograph of human ciliated cells. Terminal bar or tight junction (J) appears as lacework of ∼10 rows of densely aligned particles on fracture face of protoplasmic leaflet (PF) and as complementary grooves on external leaflet (EF) of adjacent cell. Tuft of microvilli (V) and base of cilium (C) with ciliary necklace of intramembrane particles (Ne). Bar, 0.2 μm.

Courtesy of E. E. Schneeberger


Figure 7.

Schematic representation of airway wall structure from bronchus to alveolus. EP, epithelial cells; BM, basement membrane; SM, smooth muscle cells; FC, fibrocartilaginous coat.

From Weibel


Figure 8.

Mucous membrane of small human bronchus. A: thin section showing ciliated cell (CC) with cilia (C) and microvilli (arrows), goblet cell (GC) with apical mucous plug (MU), and basal cell (BC). Fibers and fibroblasts (FB) in connective tissue. MP, macrophage. B: scanning electron micrograph of epithelial surface showing ciliary tufts (C) and mucous plug (MU) of goblet cell in process of extrusion. Bars, 5 μm.



Figure 9.

Scanning electron micrograph (A) and longitudinal (B) and transverse (C) thin sections of cilia of human bronchial epithelium. Microvilli (arrows) between cilia and axonemal complex (A) with its different structure in the shaft, neck (Ne), and basal body or kinetosome (K). mt, Microtubule; S, satellite of ciliary kinetosome. D: greater magnification of cross‐sectioned shaft of a normal cilium with dynein arms (arrowhead). E: dynein arms are lacking in a cilium from a patient with immotile cilia syndrome. Bars: 0.5 μm (A‐C); 0.1 μm (D, E).

E, courtesy of J. M. Sturgess


Figure 10.

Scanning electron micrograph of brush cell from small bronchiole of rat lung. Compare brush with cilia (C) and ordinary microvilli (arrow). Bar, 1 μm.



Figure 11.

Basal part of neuroendocrine cell of human bronchiolar epithelium showing part of nucleus (N), mitochondria (M), and secretory vesicles (arrows) with dark core, which are seen at greater magnification in inset. Bars, 0.2 μm.



Figure 12.

Scanning electron micrograph of surface of alveolar epithelium of human lung with two type II epithelial cells (EP2). Arrows, boundary (terminal bar) of type I cell. Asterisk, nucleus of type I cell. Bar, 5 μm.

From Weibel


Figure 13.

A: squamous type I epithelial cell (EP1). B: capillary endothelial cell (EN) from human lung. Nuclei (N) are surrounded by little cytoplasm, which continues into squamous extensions. Circles, fused basement membranes (BM) in barrier parts. Capillary contains neutrophil granulocyte (GR) with 2 lobes of nucleus sectioned (N′). F, fiber; J, intercellular junction. Bars, 1 μm.



Figure 14.

Squamous extensions of endothelial cell (EN) and type I pneumocyte (EP1). Cytoplasm, bounded by plasma membrane similar to that of erythrocyte (EC), contains microfilaments (f), microtubules (mt), and pinocytotic vesicles (arrows). BM, basement membrane. Bar, 0.1 μm.



Figure 15.

Scanning electron micrograph (A) and thin section (B) of type II pneumocyte with microvilli (V) on peripheral zone of apical cell surface. BM, basement membrane; J, intercellular junction; LB, lamellar body; N, nucleus. Detail of this cell is shown in Fig. . Bars, 1 μm.



Figure 16.

Characteristic organelles of type II cell related to surfactant production and storage. Cone‐shaped lamellar bodies from rat lung (A, B) and sequence of multivesicular bodies (C‐E) with stacks of lamellae (arrows) and membrane envelope (arrowheads). A, C, D: thin sections. B, E: freeze‐fracture preparation. Bars, 0.2 μm.



Figure 17.

Release of lamellar body (LB) from type II epithelial cell into surface lining layer, which contains tubular myelin (TM) continuous with the phospholipid surface film (arrows). Bar, 0.2 μn.



Figure 18.

Alveolar brush cells from rat lung. A: scanning electron micrograph showing brush (B) on narrow apical surface bent to the side, whereas cell body is covered by smooth type I cell extension (asterisks). B: thin section shows brush beneath surface film (arrows) and covered by type I cell (asterisks). C: squat brush microvilli contain bundles of filaments (f) that extend deep into cell. Bars: 1 μm (A, B); 0.5 μm (C).

B from Weibel


Figure 19.

Kinetics of transformation of type II to type I cells. Rats exposed to NO2 to induce lung damage were injected with [3H]thymidine at time 0 and killed from 1 h to 14 days later.

From Evans et al.


Figure 20.

Endothelium (EN) and single layer of smooth muscle (SM) from small pulmonary artery of human lung. Thick endothelial cytoplasm and wealth of organelles (inset) comprising mitochondria (M), endoplasmic reticulum (ER), lipid droplet (Li), specific granules (asterisks), microtubules (mt), and many vesicles (arrows). Cross‐sectioned smooth muscle cells show central nucleus, mitochondria, sarcoplasmic reticulum (SR), membrane‐bounded caveolae (arrows), filamentous matter with dense bodies (db), and cell‐to‐cell contacts (circle), cf, Collagen fibrils; el, elastic fibers. Bars, 0.5 μm.



Figure 21.

Fibroblast engaged in fibrogenesis contains much endoplasmic reticulum (ER) with ribosomes and large Golgi complex (G), as well as mitochondria (M). cf, Collagen fibrils. Bar, 0.2 μm.



Figure 22.

Septal fibroblast (FB) ramifies with slim extensions (arrows) into interstitial spaces between capillary endothelium (EN) and alveolar epithelium (EP), following the fiber strands (F). Circles, cytoplasmic areas with condensed filaments spanning the interstitial space crosswise. Bar, 2 μm.



Figure 23.

A: smooth muscle cells from small pulmonary artery of human lung cut longitudinally, db, Dense bodies of filamentous matter; db′, dense bands on plasma membrane; M, mitochondria; N, central elongated nucleus; SR, sarcoplasmic reticulum. Greater magnification shows filaments on longitudinal (B) and transverse (C) sections together with caveolae (arrows) of plasma membrane, which is coated by basement membrane. Bars: 1 μm (A); 0.2 μm (B, C).



Figure 24.

Myofibroblasts from human alveolar septa with bundles of filaments (f) extending obliquely or crosswise between dense insertion spots on plasma membrane, which appears affixed to basement membrane (arrows). Bars, 0.2 μm.



Figure 25.

Capillary pericyte (PC) with extension (arrows) that lies between the endothelial cell (EN) and its basement membrane in its peripheral part. Other processes of same cell make contact with epithelial (EP) basement membrane (arrowheads). Inset, pericyte filaments (f). Bars: 1 μm; 0.1 μm (inset).



Figure 26.

Mast cell from human alveolar septum contains granules (arrows) with scroll‐like substructure. Inset, scroll‐like substructure of mast cell at higher magnification, cf, Collagen fibrils. Bars, 1 μm; 0.1 μm (inset).



Figure 27.

Scanning electron micrograph of human alveolar macrophage on alveolar epithelial surface. Thin advancing cytoplasmic lamella appears partly attached to epithelium. Bar, 5 μm.

From Weibel


Figure 28.

Alveolar macrophage of human lung apposed to epithelium in several places (arrows). Advancing lamella (AL) is free of organelles that are rich in perinuclear cytoplasm. Bar, 2 μm.



Figure 29.

Alveolar macrophage lies beneath thin film of surface lining layer (arrows). Exclusion of organelles from short peripheral cytoplasmic flaps (asterisks). Bar, 1 μm.



Figure 30.

Interstitial macrophage (MP) from human lung contains heterogeneous dark material in phagolysosomes or residual bodies (arrows). Macrophage is in group of free cells with lymphocytes (LC) and plasma cell (PC). Bar, 2 μm.



Figure 31.

A: phagocytosis of erythrocyte (EC) by alveolar macrophage. Ectoplasmic “lips” (asterisks) are pinching off fragment. B: phagolysosome may contain tubular myelin (TM) and lamellar (L) osmiophilic material, possibly the result of enzymatic disassembly of surface lining material. Bars: 1 μm (A); 0.2 μm (B).



Figure 32.

Plasma cell from small rat lymph node contains massive rough endoplasmic reticulum (ER). Golgi complexes (G) are in adjacent plasma cell. Bar, 2 μm.

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Ewald R. Weibel. Lung Cell Biology. Compr Physiol 2011, Supplement 10: Handbook of Physiology, The Respiratory System, Circulation and Nonrespiratory Functions: 47-91. First published in print 1985. doi: 10.1002/cphy.cp030102