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Chapter 6 : Lymphoid Organs

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Abstract:

The lymphoid cells responsible for specific immune responses are distributed in blood, lymphatics, and a number of tissues known as the lymphoid system. The morphologic characteristics and functional properties of the lymphoid organs that make up the lymphoid system are different. The bone marrow serves as the major source for lymphoid stem cells. T cells develop from stem cells that migrate to the thymus, learn to recognize self and nonself, and subsequently recirculate to home in thymus-dependent areas of other lymphoid organs-spleen, lymph node, and gastrointestinal tract. B cells mature in the bone marrow, liver, or gastrointestinal lymphoid tissue and migrate to B-cell areas (follicles) of the other lymphoid organs. Different immune responses take place in different lymphoid tissue microenvironments. Specific T-dependent B-cell proliferation occurs in germinal centers, T-cell proliferation in the paracortex or periarteriolar sheath, T-independent B-cell proliferation in marginal zones, and antibody secretion in medullary cords. Memory B cells may differentiate in the mantle of germinal centers and be stored in primary follicles. Induction of antibody formation is associated with a hyperplasia of follicles, plasma cell production, and synthesis and secretion of immunoglobulin antibodies into the circulation by plasma cells that remain in the lymph nodes, whereas cellular sensitivity is associated with hyperplasia of thymus-dependent areas and release of sensitized T lymphocytes into the circulation.

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
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Image of Figure 6.1
Figure 6.1

Diagram of the human lymphoid system. The system consists of circulating lymphocytes and the lymphoid organs and includes the network of lymphatic vessels and the lymph nodes stationed along the vessels, the bone marrow (in the long bones, only one of which is illustrated), the thymus, the spleen, the adenoids, the tonsils, the Peyer's patches of the small intestine (GALT), the appendix, the lung (BALT), the skin (SALT), and the mammary gland (DALT). Afferent lymphatic vessels collect the fluid and cells that escape from the blood capillaries and return them via the lymph nodes to the bloodstream at the subclavian veins. In addition, efferent lymphatics collect lymphocytes and antibody molecules from the lymph nodes and deliver them to the blood. The thoracic duct is the largest lymph vessel in the body and joins the left subclavian vein. The right lymphatic duct joins the right subclavian vein. Seventy-five percent of body tissues are drained by the thoracic duct, and 25% are drained by the right lymphatic duct. There is no pump for the lymphatic circulation, such as the heart for the systemic circulation. Lymphatic fluid is propelled by contraction of skeletal muscles or in larger vessels by smooth muscle cells that force the fluid from one level to another past valves that permit passage of fluid and cells in only one direction.

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
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Image of Figure 6.2
Figure 6.2

Lymph node lymphatics. Lymphatic collecting vessels are similar to veins but, except for the larger vessels, do not have muscular walls. Afferent lymphatics deliver to the lymph node lymphatic fluid-containing blood cells that have escaped through capillaries, as well as foreign material that has entered the interstitial spaces of the body. The lymph nodes act as a filter for the lymphatic fluid, which delivers antigens to the node. In response to the antigens, specific antibodies or specifically sensitized lymphocytes are produced in the lymph nodes and delivered to the systemic circulation by efferent lymphatic vessels.

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
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Image of Figure 6.3
Figure 6.3

Thymus. The thymus contains an outside layer of packed lymphoid cells (cortex), an inside layer of less densely packed cells (medulla), a fibrous capsule, prominent trabeculae that divide the organ into lobules, and a hilum with entering arteries and draining veins and lymphatics. The cortex contains thymic nurse cells, which are epithelium-derived cells in the outer cortex. Each thymic nurse cell contains 5 to 20 immature thymocytes within cytoplasmic spaces, as well as thymocytes that are partially enclosed by plasma membranes on the surface of the cell. Thymic nurse cells play a critical role in the early differentiation of thymocytes.

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
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Image of Figure 6.4
Figure 6.4

T-cell differentiation and acquisition of CD markers in the human thymus. Prothymocytes, bearing CD7, CD34, and CD45 markers, arise in the bone marrow and migrate to the thymus, where they mature into T cells. Thymocytes are cells in the thymus. T cells are thymus-derived cells that have matured in the thymus and have migrated to peripheral lymphoid markers. The first identifiable markers in the thymic cortex are CD2 and CD5; CD1, CD4, and CD5 are acquired as the cortical thymocytes mature. In the thymic medulla, CD1 is lost, and CD3, part of the T-cell receptor, is acquired. As a final differentiation step, CD4+ cells and CD8+ cells segregate into two separate populations. CD4 designates the Thelper population; CD8 designates the T-cytotoxic population. After leaving the thymus, CD4+ T cells locate preferentially in the cortex of lymph nodes; CD8+ T cells localize in the medulla, although there is considerable overlap.

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
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Image of Figure 6.5
Figure 6.5

Scheme of selection of T-cell repertoire in the thymus. Pre-T cells from the bone marrow enter the thymus and develop a large variety of specificities of Tcell αβ receptor and express both CD4 and CD8. Cells at this stage of differentiation that react with MHC on thymic nurse cells are positively selected, proliferate, and express the CD3 component of the T-cell receptor. Those that do not react are eliminated by apoptosis. After this stage, CD3+, CD4+, and CD8+ cells that react “strongly” with MHC on dendritic cells or macrophages are “negatively selected” and eliminated (clonal deletion). Those that react “weakly” with MHC (in a manner similar to that of MHC modified with foreign peptide) are allowed to survive and differentiate into CD3+ CD4+ or CD3+ CD8+ cells, which now leave the thymus for further development into T cells in the peripheral lymphoid organs.

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
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Image of Figure 6.6
Figure 6.6

Normal lymph node. Nodes are made up of lymphoid cells contained in meshwork of reticular fibers surrounded by connective tissue capsule. Most lymph nodes are bean-shaped, with an indented area known as the hilum. The cortex (outer layer) contains densely packed lymphoid cells and includes germinal centers responsible for production of antibody-synthesizing plasma cells and paracortical areas where lymphocytes are produced. The medulla (central area) consists of sinusoidal channels maintained by reticular cells. Columns of lymphoid cells are found between sinusoids in areas containing reticular macrophages. Afferent lymphatics drain through cortex around germinal centers into medullary sinusoids. Since the medullary sinusoids contain lymphatic fluid and not blood, there are normally very few red blood cells in the medullary sinusoids. Medullary sinusoids drain into efferent lymphatics and are collected by main efferent lymphatic that drains from the hilum. The main artery divides into capillaries supplying the cortex. These capillaries drain into veins that follow the trabeculae and exit at the hilum

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
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Image of Figure 6.7
Figure 6.7

Normal splenic lobule. The spleen is composed of a network of sinusoidal channels filled mainly with red blood cells (red pulp). There are no lymphatic vessels. Blood enters through arteries that may empty directly into splenic sinusoids or into the reticular area between sinusoids. The sinusoids are drained by veins that exit via trabecular veins to a large vein that leaves the spleen at the hilus. A zone of densely packed lymphocytes surrounding the central arteriole contains T cells (thymus-dependent area), whereas B cells are found surrounding the germinal center. The mantle surrounding the germinal center is composed mainly of B cells but also T cells, which are believed to be nonactivated cells pushed aside from the B-cell zone by formation of the germinal center. Overlying the mantle is the marginal zone, containing venous capillaries that permit circulating cells to enter the white pulp. Blood flows from the central arteriole through small follicular arterioles into the marginal sinus, which drains into the red pulp. The central arteriole continues through the white pulp. Upon exiting the white pulp, the central arteriole is divided into many small branches—the penicillus arterioles—which drain into the sinusoids or the medullary cords of Billroth.

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
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Image of Figure 6.8
Figure 6.8

Structure of the Peyer's patch. Peyer's patches are collections of lymphoid tissue in the submucosa of the small intestine containing follicles (germinal centers) and interfollicular lymphoid tissue (thymus-dependent zones). The overlying mucosa is covered by a dome of epithelial cells, some of which (M cells) have the specialized property of transporting antigens from the lumen into the Peyer's patch. Lacteals drain filtrated from the intestinal contents into the lymphatics. In addition, both T and B cells are produced in the Peyer's patch and added to the lymphatic fluid.

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
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Image of Figure 6.9
Figure 6.9

Cellular traffic in the secretory system. T and B cells produced in the GALT are delivered to the systemic circulation through the thoracic duct. These cells in the blood then may localize in other secretory lymphatic tissue, such as mammary glands, or in systemic lymph nodes. For more on the cell recognition systems involved in specific localization of lymphocytes, see the text.

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
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Image of Figure 6.10
Figure 6.10

Germinal center formation. Localization of labeled antigen in lymph node following immunization demonstrates distribution in both medullary and cortical macrophages. Antigen first appears in lining cells of the subcortical sinus. On day 2, labeled macrophages are scattered through the cortex. By day 4, the label appears in cells in developing follicles (FDCs), and blast cells can be identified underlying the antigen-containing cells. Blast cells and their progeny increase in number until a typical germinal center (secondary follicle) is formed. By day 7, plasma cells and memory cells appear deep to the germinal center. Plasma cells then migrate into medullary cords and secrete antibody into medullary sinusoids. Memory B cells move to the marginal zone. During involution of the follicle, there are many tingible body macrophages that phagocytose numerous cells, even dividing cells.

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
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Image of Figure 6.11
Figure 6.11

Germinal center B cells. The B cells seen in a germinal center range in size and shape from small round cells to large irregular “cleaved” cells on the basis of nuclear appearance. Primary follicles are composed of small round lymphocytes, almost all B cells. Germinal centers contain a mixture of B cells: small round, intermediate round, large round, medium cleaved, and large cleaved. Cleaved cells are believed to represent “activated” B cells; large round cells are “blast” cells that divide to produce two daughter B cells that are small and round. These morphologic cell types have been used to classify tumors arising from B cells (B-cell lymphomas). Small round B-cell tumors have a good prognosis; large round or cleaved cells have a poor prognosis; cell types in between have an intermediate prognosis.

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
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Image of Figure 6.12
Figure 6.12

Morphologic response of lymph node to antigen stimulus. Induction of pure cell-mediated immunity leads to proliferation of lymphocytes in the paracortical zone. Induction of pure antibody formation results in germinal center formation and appearance of plasma cells in medullary cords. Immunization with most antigens produces both changes with enlargement of paracortical zones and production of germinal centers.

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
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Image of Figure 6.13
Figure 6.13

Postulated migration of hematopoietic stem cells. Primitive hematopoietic cells may arise within the neural crest and migrate to the ventral mesenteric region or arise in the ventral mesoderm. From the ventral mesoderm, the cells migrate to the fetal yolk sac or to the primitive liver and then to fetal bone marrow and spleen. Prothymocytes arising in the dorsal mesoderm or bone marrow migrate to the thymus (T-cell development); pro-B cells in the bone marrow migrate to the gastrointestinal tract or mature in the liver to functional B cells. Maturing T and B cells migrate to the spleen or lymph nodes, where final differentiation steps occur. BM, bone marrow; GI, gastrointestinal tract; L, liver; N, neural crest; S, spleen; T, thymus; V, ventral mesoderm, YS, yolk sac.

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
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Image of Figure 6.14
Figure 6.14

Scheme of hematopoiesis and differentiation of T and B cells. Precursors of all blood cells, as well as tissue T and B cells, arise from a common stem cell in the bone marrow. The first step in determination is between lymphoid stem cells and myeloid stem cells. Cells determined to become T cells (prothymocytes) migrate to the thymus. The interaction of these cells with thymic epithelium induces maturation of thymocytes to T cells. T cells are thymus-derived cells, which leave the thymus and migrate to thymus-dependent zones of the spleen and lymph nodes, where they undergo further maturation. B-cell maturation occurs in the bone marrow, liver, or gastrointestinal tract (cloacal bursa in fowl) and migrate to the bursa-dependent zones of peripheral organs. Myeloid differentiation occurs in the bone marrow, and normally only mature cells exit from the bone marrow and enter the blood (see chapter 2).

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
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Tables

Generic image for table
Table 6.1

Characterization of the thymic stroma

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
Generic image for table
Table 6.2

Comparison of differentiation stages of skin and thymic medullary epithelium

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
Generic image for table
Table 6.3

CD markers of T-cell differentiation in the thymus

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
Generic image for table
Table 6.4

Comparison of spleen to lymph node: structure and function

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
Generic image for table
Table 6.5

Some characteristics of lymphoid organs a

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
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Table 6.6

Lymphocyte surface molecules reacting with HEVs

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
Generic image for table
Table 6.7

Chemokines and receptors in lymphoid organs

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6
Generic image for table
Table 6.8

Functional lymphoid organ microenvironments

Citation: Sell S. 2001. Lymphoid Organs, p 198-232. In Immunology, Immunopathology, and Immunity, Sixth Edition. ASM Press, Washington, DC. doi: 10.1128/9781555818012.ch6

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