Comparative ultrastructural morphology of human basophils stimulated to release histamine by anti-IgE, recombinant IgE-dependent histamine-releasing factor,

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Comparative ultrastructural morphology of human basophils stimulated to release histamine by anti-IgE, recombinant IgE-dependent histamine-releasing factor, or monocyte chemotactic protein-1  Ann M. Dvorak, MDa, John T. Schroeder, PhDb, Donald W. MacGlashan, MD, PhDb, Karen P. Bryan, BSa, Ellen S. Morgan, BAa, Lawrence M. Lichtenstein, MD, PhDb, Susan M. MacDonald, MDb  Journal of Allergy and Clinical Immunology  Volume 98, Issue 2, Pages 355-370 (August 1996) DOI: 10.1016/S0091-6749(96)70160-4 Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 1 Control human basophil shows a polylobed nucleus (N) and numerous large secretory granules filled with electron-dense particles. Another granule type is small, rests near the nucleus, and contains homogeneous, poorly electron-dense particles (arrow). Focal electron-dense particles and aggregates of glycogen are present in the cytoplasm. (Original magnification ×26,500.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 2 Portions of basophils (2 minutes after stimulation with anti-IgE) show granule-vesicle attachments (A and B), cytoplasmic perigranular vesicles (C), and exocytosis of a single granule (D). In A and B, vesicles attached to granules contain altered granule contents, and granules contain particles. Granule-attached vesicles are surrounded by large, electron-dense aggregates of glycogen and are oriented toward the overlying plasma membrane. Similar glycogen clusters, which are free in the cytoplasm (arrows), enclose electron-lucent vesicles. In C, one granule has completely released its dense particle contents and contains larger, electron-dense glycogen particles. Small, smooth membrane–bound, electron-lucent vesicles surround this granule pole, just beneath the overlying plasma membrane. Two granule types of human basophils are shown in D. One particle-containing granule, devoid of its membrane, has been extruded to the cell surface (arrow); a small, homogeneous granule (arrowhead) rests beside the nucleus (N). (Original magnifications: A, ×62,500; B, ×76,000; C, ×58,000; D, ×50,000.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 2 Portions of basophils (2 minutes after stimulation with anti-IgE) show granule-vesicle attachments (A and B), cytoplasmic perigranular vesicles (C), and exocytosis of a single granule (D). In A and B, vesicles attached to granules contain altered granule contents, and granules contain particles. Granule-attached vesicles are surrounded by large, electron-dense aggregates of glycogen and are oriented toward the overlying plasma membrane. Similar glycogen clusters, which are free in the cytoplasm (arrows), enclose electron-lucent vesicles. In C, one granule has completely released its dense particle contents and contains larger, electron-dense glycogen particles. Small, smooth membrane–bound, electron-lucent vesicles surround this granule pole, just beneath the overlying plasma membrane. Two granule types of human basophils are shown in D. One particle-containing granule, devoid of its membrane, has been extruded to the cell surface (arrow); a small, homogeneous granule (arrowhead) rests beside the nucleus (N). (Original magnifications: A, ×62,500; B, ×76,000; C, ×58,000; D, ×50,000.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 2 Portions of basophils (2 minutes after stimulation with anti-IgE) show granule-vesicle attachments (A and B), cytoplasmic perigranular vesicles (C), and exocytosis of a single granule (D). In A and B, vesicles attached to granules contain altered granule contents, and granules contain particles. Granule-attached vesicles are surrounded by large, electron-dense aggregates of glycogen and are oriented toward the overlying plasma membrane. Similar glycogen clusters, which are free in the cytoplasm (arrows), enclose electron-lucent vesicles. In C, one granule has completely released its dense particle contents and contains larger, electron-dense glycogen particles. Small, smooth membrane–bound, electron-lucent vesicles surround this granule pole, just beneath the overlying plasma membrane. Two granule types of human basophils are shown in D. One particle-containing granule, devoid of its membrane, has been extruded to the cell surface (arrow); a small, homogeneous granule (arrowhead) rests beside the nucleus (N). (Original magnifications: A, ×62,500; B, ×76,000; C, ×58,000; D, ×50,000.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 2 Portions of basophils (2 minutes after stimulation with anti-IgE) show granule-vesicle attachments (A and B), cytoplasmic perigranular vesicles (C), and exocytosis of a single granule (D). In A and B, vesicles attached to granules contain altered granule contents, and granules contain particles. Granule-attached vesicles are surrounded by large, electron-dense aggregates of glycogen and are oriented toward the overlying plasma membrane. Similar glycogen clusters, which are free in the cytoplasm (arrows), enclose electron-lucent vesicles. In C, one granule has completely released its dense particle contents and contains larger, electron-dense glycogen particles. Small, smooth membrane–bound, electron-lucent vesicles surround this granule pole, just beneath the overlying plasma membrane. Two granule types of human basophils are shown in D. One particle-containing granule, devoid of its membrane, has been extruded to the cell surface (arrow); a small, homogeneous granule (arrowhead) rests beside the nucleus (N). (Original magnifications: A, ×62,500; B, ×76,000; C, ×58,000; D, ×50,000.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 3 Human basophils, stimulated for 7 minutes with anti-IgE, show AND. At low magnification (A), a completely degranulated basophil displays a polylobed nucleus (N), marked shape change with numerous elongated cell processes, focal dense aggregates of cytoplasmic glycogen, and extruded granules (arrowhead) and dense concentric membranes (arrows) adjacent to the cell surface. At higher magnifications (B and C), exocytosis of membrane-free granules (arrows), dense concentric membranes (open arrowheads), and a CLC (closed arrowhead) are evident. Note that several unaltered, large granules remain in the cytoplasm of these degranulating basophils. (Original magnifications: A, ×18,000; B, ×30,000; C, ×33,000.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 3 Human basophils, stimulated for 7 minutes with anti-IgE, show AND. At low magnification (A), a completely degranulated basophil displays a polylobed nucleus (N), marked shape change with numerous elongated cell processes, focal dense aggregates of cytoplasmic glycogen, and extruded granules (arrowhead) and dense concentric membranes (arrows) adjacent to the cell surface. At higher magnifications (B and C), exocytosis of membrane-free granules (arrows), dense concentric membranes (open arrowheads), and a CLC (closed arrowhead) are evident. Note that several unaltered, large granules remain in the cytoplasm of these degranulating basophils. (Original magnifications: A, ×18,000; B, ×30,000; C, ×33,000.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 3 Human basophils, stimulated for 7 minutes with anti-IgE, show AND. At low magnification (A), a completely degranulated basophil displays a polylobed nucleus (N), marked shape change with numerous elongated cell processes, focal dense aggregates of cytoplasmic glycogen, and extruded granules (arrowhead) and dense concentric membranes (arrows) adjacent to the cell surface. At higher magnifications (B and C), exocytosis of membrane-free granules (arrows), dense concentric membranes (open arrowheads), and a CLC (closed arrowhead) are evident. Note that several unaltered, large granules remain in the cytoplasm of these degranulating basophils. (Original magnifications: A, ×18,000; B, ×30,000; C, ×33,000.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 4 This basophil (anti-IgE, 7-minute stimulation) shows PMD characterized by secretion of granule contents with retention and enlargement of granule containers. Some residual granule particles remain in one enlarged granule (closed arrowhead); another granule shows only focal piecemeal loss (arrow) of its granule particles. One empty granule contains a CLC (open arrowhead). Note that this cell is round and lacks the extensive shape change and elongation of surface processes that characterize cells undergoing AND (compare with Fig. 3, A). N, Nucleus. (Original magnification ×25,500.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 5 This basophil (rHRF, 7 minutes) shows a motile configuration without morphologic evidence of PMD or AND. A full complement of unaltered, particle-filled granules is present. The nucleus (N) has several lobes. An elongated uropod (arrow) is evident. (Original magnification ×13,500.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 6 Higher magnification views of human basophils (2 minutes [A and B] or 7 minutes [C to F] after stimulation with rHRF) are shown. Glycogen-rich GVAs are illustrated (A to D). Attached vesicles show a variable degree of altered contents similar to that underlying them in the particulate granule material, and vesicles are encased with dense glycogen aggregates. Some of these attached vesicles are oriented toward the cell surface (A,B). Note the separate vesicle adjacent to one of the attached vesicles, which spans the cytoplasm to abut on the plasma membrane (arrow, B). The entire content of one granule is altered (E), as in PMD, and an extensive glycogen aggregate surrounds this granule. In F, granule exocytosis, as in AND, is illustrated (arrow). (Original magnifications: A, ×53,500; B, ×56,000; C, ×52,000; D, ×42,000; E, ×35,500; F, ×51,500.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 6 Higher magnification views of human basophils (2 minutes [A and B] or 7 minutes [C to F] after stimulation with rHRF) are shown. Glycogen-rich GVAs are illustrated (A to D). Attached vesicles show a variable degree of altered contents similar to that underlying them in the particulate granule material, and vesicles are encased with dense glycogen aggregates. Some of these attached vesicles are oriented toward the cell surface (A,B). Note the separate vesicle adjacent to one of the attached vesicles, which spans the cytoplasm to abut on the plasma membrane (arrow, B). The entire content of one granule is altered (E), as in PMD, and an extensive glycogen aggregate surrounds this granule. In F, granule exocytosis, as in AND, is illustrated (arrow). (Original magnifications: A, ×53,500; B, ×56,000; C, ×52,000; D, ×42,000; E, ×35,500; F, ×51,500.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 6 Higher magnification views of human basophils (2 minutes [A and B] or 7 minutes [C to F] after stimulation with rHRF) are shown. Glycogen-rich GVAs are illustrated (A to D). Attached vesicles show a variable degree of altered contents similar to that underlying them in the particulate granule material, and vesicles are encased with dense glycogen aggregates. Some of these attached vesicles are oriented toward the cell surface (A,B). Note the separate vesicle adjacent to one of the attached vesicles, which spans the cytoplasm to abut on the plasma membrane (arrow, B). The entire content of one granule is altered (E), as in PMD, and an extensive glycogen aggregate surrounds this granule. In F, granule exocytosis, as in AND, is illustrated (arrow). (Original magnifications: A, ×53,500; B, ×56,000; C, ×52,000; D, ×42,000; E, ×35,500; F, ×51,500.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 6 Higher magnification views of human basophils (2 minutes [A and B] or 7 minutes [C to F] after stimulation with rHRF) are shown. Glycogen-rich GVAs are illustrated (A to D). Attached vesicles show a variable degree of altered contents similar to that underlying them in the particulate granule material, and vesicles are encased with dense glycogen aggregates. Some of these attached vesicles are oriented toward the cell surface (A,B). Note the separate vesicle adjacent to one of the attached vesicles, which spans the cytoplasm to abut on the plasma membrane (arrow, B). The entire content of one granule is altered (E), as in PMD, and an extensive glycogen aggregate surrounds this granule. In F, granule exocytosis, as in AND, is illustrated (arrow). (Original magnifications: A, ×53,500; B, ×56,000; C, ×52,000; D, ×42,000; E, ×35,500; F, ×51,500.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 6 Higher magnification views of human basophils (2 minutes [A and B] or 7 minutes [C to F] after stimulation with rHRF) are shown. Glycogen-rich GVAs are illustrated (A to D). Attached vesicles show a variable degree of altered contents similar to that underlying them in the particulate granule material, and vesicles are encased with dense glycogen aggregates. Some of these attached vesicles are oriented toward the cell surface (A,B). Note the separate vesicle adjacent to one of the attached vesicles, which spans the cytoplasm to abut on the plasma membrane (arrow, B). The entire content of one granule is altered (E), as in PMD, and an extensive glycogen aggregate surrounds this granule. In F, granule exocytosis, as in AND, is illustrated (arrow). (Original magnifications: A, ×53,500; B, ×56,000; C, ×52,000; D, ×42,000; E, ×35,500; F, ×51,500.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 6 Higher magnification views of human basophils (2 minutes [A and B] or 7 minutes [C to F] after stimulation with rHRF) are shown. Glycogen-rich GVAs are illustrated (A to D). Attached vesicles show a variable degree of altered contents similar to that underlying them in the particulate granule material, and vesicles are encased with dense glycogen aggregates. Some of these attached vesicles are oriented toward the cell surface (A,B). Note the separate vesicle adjacent to one of the attached vesicles, which spans the cytoplasm to abut on the plasma membrane (arrow, B). The entire content of one granule is altered (E), as in PMD, and an extensive glycogen aggregate surrounds this granule. In F, granule exocytosis, as in AND, is illustrated (arrow). (Original magnifications: A, ×53,500; B, ×56,000; C, ×52,000; D, ×42,000; E, ×35,500; F, ×51,500.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 7 MCP-1–stimulated basophils are shown. In A (5 seconds after stimulation), a basophil with a polylobed nucleus (N) displays PMD with granule particle losses from approximately 50% of visible granules. Partially and completely empty granules are moderately increased in size and have not fused with adjacent granules or plasma membrane. In B (30 seconds after stimulation), another cell with a polylobed nucleus (N) shows extensive PMD involving nearly all secretory granules. Empty granule chambers are swollen and remain in the cytoplasm. Amplification of surface processes and shape changes are absent from these cells, which are actively undergoing PMD. (Original magnifications: A, ×14,500; B, ×15,000.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 7 MCP-1–stimulated basophils are shown. In A (5 seconds after stimulation), a basophil with a polylobed nucleus (N) displays PMD with granule particle losses from approximately 50% of visible granules. Partially and completely empty granules are moderately increased in size and have not fused with adjacent granules or plasma membrane. In B (30 seconds after stimulation), another cell with a polylobed nucleus (N) shows extensive PMD involving nearly all secretory granules. Empty granule chambers are swollen and remain in the cytoplasm. Amplification of surface processes and shape changes are absent from these cells, which are actively undergoing PMD. (Original magnifications: A, ×14,500; B, ×15,000.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 8 Basophils (30 seconds, MCP-1 stimulation) display AND. In A several membrane-free granules are being extruded through a single wide opening in the plasma membrane (arrow). In B extensive shape change and surface process amplification are evident. Electron-dense cytoplasmic aggregates of glycogen are present. Exocytosis of granules and concentric dense membranes has occurred at several openings in the cell surface (arrows). A single unaltered granule, containing dense particles and a CLC, remains in the cytoplasm (arrowhead). (Original magnifications: A, ×37,500; B, ×16,500.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 8 Basophils (30 seconds, MCP-1 stimulation) display AND. In A several membrane-free granules are being extruded through a single wide opening in the plasma membrane (arrow). In B extensive shape change and surface process amplification are evident. Electron-dense cytoplasmic aggregates of glycogen are present. Exocytosis of granules and concentric dense membranes has occurred at several openings in the cell surface (arrows). A single unaltered granule, containing dense particles and a CLC, remains in the cytoplasm (arrowhead). (Original magnifications: A, ×37,500; B, ×16,500.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 9 A higher magnification montage of basophils, stimulated with MCP-1 for 5 seconds (A to G) or 30 seconds (H and I), shows glycogen-rich, granule-vesicle attachments that accompany PMD. Some of the attached, elongated vesicles contain electron-dense granule particles (arrows in A to C); some attached vesicles contain less dense (or electron-lucent) altered materials (arrows in D to I). Larger, electron-dense glycogen particles are interposed at the necks of tubular-vesicular granule extensions (A, B, and E) or encase the outer surface of attached vesicles (D, F to I). Also noted are single (A to C, E, F, and I), double (H), and triple (G) glycogen-rich vesicles attached to granules. Some granules, with vesicles attached, have a full complement of dense particles (A to D), focal, underlying piecemeal losses of granule particles (E, G to I), or diminished particles throughout (F). One empty granule (D) has individual glycogen particles within it. A, ×50,000; B, ×57,500; C and D, ×49,000; E, ×70,000; F and G ×63,000; H, ×51,000; I, ×54,500.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 9 A higher magnification montage of basophils, stimulated with MCP-1 for 5 seconds (A to G) or 30 seconds (H and I), shows glycogen-rich, granule-vesicle attachments that accompany PMD. Some of the attached, elongated vesicles contain electron-dense granule particles (arrows in A to C); some attached vesicles contain less dense (or electron-lucent) altered materials (arrows in D to I). Larger, electron-dense glycogen particles are interposed at the necks of tubular-vesicular granule extensions (A, B, and E) or encase the outer surface of attached vesicles (D, F to I). Also noted are single (A to C, E, F, and I), double (H), and triple (G) glycogen-rich vesicles attached to granules. Some granules, with vesicles attached, have a full complement of dense particles (A to D), focal, underlying piecemeal losses of granule particles (E, G to I), or diminished particles throughout (F). One empty granule (D) has individual glycogen particles within it. A, ×50,000; B, ×57,500; C and D, ×49,000; E, ×70,000; F and G ×63,000; H, ×51,000; I, ×54,500.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 9 A higher magnification montage of basophils, stimulated with MCP-1 for 5 seconds (A to G) or 30 seconds (H and I), shows glycogen-rich, granule-vesicle attachments that accompany PMD. Some of the attached, elongated vesicles contain electron-dense granule particles (arrows in A to C); some attached vesicles contain less dense (or electron-lucent) altered materials (arrows in D to I). Larger, electron-dense glycogen particles are interposed at the necks of tubular-vesicular granule extensions (A, B, and E) or encase the outer surface of attached vesicles (D, F to I). Also noted are single (A to C, E, F, and I), double (H), and triple (G) glycogen-rich vesicles attached to granules. Some granules, with vesicles attached, have a full complement of dense particles (A to D), focal, underlying piecemeal losses of granule particles (E, G to I), or diminished particles throughout (F). One empty granule (D) has individual glycogen particles within it. A, ×50,000; B, ×57,500; C and D, ×49,000; E, ×70,000; F and G ×63,000; H, ×51,000; I, ×54,500.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 9 A higher magnification montage of basophils, stimulated with MCP-1 for 5 seconds (A to G) or 30 seconds (H and I), shows glycogen-rich, granule-vesicle attachments that accompany PMD. Some of the attached, elongated vesicles contain electron-dense granule particles (arrows in A to C); some attached vesicles contain less dense (or electron-lucent) altered materials (arrows in D to I). Larger, electron-dense glycogen particles are interposed at the necks of tubular-vesicular granule extensions (A, B, and E) or encase the outer surface of attached vesicles (D, F to I). Also noted are single (A to C, E, F, and I), double (H), and triple (G) glycogen-rich vesicles attached to granules. Some granules, with vesicles attached, have a full complement of dense particles (A to D), focal, underlying piecemeal losses of granule particles (E, G to I), or diminished particles throughout (F). One empty granule (D) has individual glycogen particles within it. A, ×50,000; B, ×57,500; C and D, ×49,000; E, ×70,000; F and G ×63,000; H, ×51,000; I, ×54,500.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 9 A higher magnification montage of basophils, stimulated with MCP-1 for 5 seconds (A to G) or 30 seconds (H and I), shows glycogen-rich, granule-vesicle attachments that accompany PMD. Some of the attached, elongated vesicles contain electron-dense granule particles (arrows in A to C); some attached vesicles contain less dense (or electron-lucent) altered materials (arrows in D to I). Larger, electron-dense glycogen particles are interposed at the necks of tubular-vesicular granule extensions (A, B, and E) or encase the outer surface of attached vesicles (D, F to I). Also noted are single (A to C, E, F, and I), double (H), and triple (G) glycogen-rich vesicles attached to granules. Some granules, with vesicles attached, have a full complement of dense particles (A to D), focal, underlying piecemeal losses of granule particles (E, G to I), or diminished particles throughout (F). One empty granule (D) has individual glycogen particles within it. A, ×50,000; B, ×57,500; C and D, ×49,000; E, ×70,000; F and G ×63,000; H, ×51,000; I, ×54,500.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 9 A higher magnification montage of basophils, stimulated with MCP-1 for 5 seconds (A to G) or 30 seconds (H and I), shows glycogen-rich, granule-vesicle attachments that accompany PMD. Some of the attached, elongated vesicles contain electron-dense granule particles (arrows in A to C); some attached vesicles contain less dense (or electron-lucent) altered materials (arrows in D to I). Larger, electron-dense glycogen particles are interposed at the necks of tubular-vesicular granule extensions (A, B, and E) or encase the outer surface of attached vesicles (D, F to I). Also noted are single (A to C, E, F, and I), double (H), and triple (G) glycogen-rich vesicles attached to granules. Some granules, with vesicles attached, have a full complement of dense particles (A to D), focal, underlying piecemeal losses of granule particles (E, G to I), or diminished particles throughout (F). One empty granule (D) has individual glycogen particles within it. A, ×50,000; B, ×57,500; C and D, ×49,000; E, ×70,000; F and G ×63,000; H, ×51,000; I, ×54,500.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 9 A higher magnification montage of basophils, stimulated with MCP-1 for 5 seconds (A to G) or 30 seconds (H and I), shows glycogen-rich, granule-vesicle attachments that accompany PMD. Some of the attached, elongated vesicles contain electron-dense granule particles (arrows in A to C); some attached vesicles contain less dense (or electron-lucent) altered materials (arrows in D to I). Larger, electron-dense glycogen particles are interposed at the necks of tubular-vesicular granule extensions (A, B, and E) or encase the outer surface of attached vesicles (D, F to I). Also noted are single (A to C, E, F, and I), double (H), and triple (G) glycogen-rich vesicles attached to granules. Some granules, with vesicles attached, have a full complement of dense particles (A to D), focal, underlying piecemeal losses of granule particles (E, G to I), or diminished particles throughout (F). One empty granule (D) has individual glycogen particles within it. A, ×50,000; B, ×57,500; C and D, ×49,000; E, ×70,000; F and G ×63,000; H, ×51,000; I, ×54,500.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 9 A higher magnification montage of basophils, stimulated with MCP-1 for 5 seconds (A to G) or 30 seconds (H and I), shows glycogen-rich, granule-vesicle attachments that accompany PMD. Some of the attached, elongated vesicles contain electron-dense granule particles (arrows in A to C); some attached vesicles contain less dense (or electron-lucent) altered materials (arrows in D to I). Larger, electron-dense glycogen particles are interposed at the necks of tubular-vesicular granule extensions (A, B, and E) or encase the outer surface of attached vesicles (D, F to I). Also noted are single (A to C, E, F, and I), double (H), and triple (G) glycogen-rich vesicles attached to granules. Some granules, with vesicles attached, have a full complement of dense particles (A to D), focal, underlying piecemeal losses of granule particles (E, G to I), or diminished particles throughout (F). One empty granule (D) has individual glycogen particles within it. A, ×50,000; B, ×57,500; C and D, ×49,000; E, ×70,000; F and G ×63,000; H, ×51,000; I, ×54,500.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 9 A higher magnification montage of basophils, stimulated with MCP-1 for 5 seconds (A to G) or 30 seconds (H and I), shows glycogen-rich, granule-vesicle attachments that accompany PMD. Some of the attached, elongated vesicles contain electron-dense granule particles (arrows in A to C); some attached vesicles contain less dense (or electron-lucent) altered materials (arrows in D to I). Larger, electron-dense glycogen particles are interposed at the necks of tubular-vesicular granule extensions (A, B, and E) or encase the outer surface of attached vesicles (D, F to I). Also noted are single (A to C, E, F, and I), double (H), and triple (G) glycogen-rich vesicles attached to granules. Some granules, with vesicles attached, have a full complement of dense particles (A to D), focal, underlying piecemeal losses of granule particles (E, G to I), or diminished particles throughout (F). One empty granule (D) has individual glycogen particles within it. A, ×50,000; B, ×57,500; C and D, ×49,000; E, ×70,000; F and G ×63,000; H, ×51,000; I, ×54,500.) Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions

FIG. 10 Kinetic and quantitative relationships of four activated morphologic features induced in human basophils by anti-IgE, rHRF, or MCP-1 compared with unstimulated human basophils. Journal of Allergy and Clinical Immunology 1996 98, 355-370DOI: (10.1016/S0091-6749(96)70160-4) Copyright © 1996 Mosby, Inc. Terms and Conditions