LYMPHOID TISSUES

Lymphocytes are located most obviously at strategic sites which are liable to infection. The main areas of
lymphocyte concentration are classified as primary or secondary lymphoid organs, according to whether
they are involved in de novo lymphocyte generation (e.g. bone marrow, thymus) or the site of mature
lymphocyte activation (e.g. lymph nodes, spleen).
All lymphocytes arise from pluripotent haemopoietic stem cells in the bone marrow. The B lymphocyte
lineage develops through differentiation stages within the bone marrow. The newly formed B cells then
migrate to peripheral sites through the circulation. In contrast, bone marrow-derived stem cells of T
lymphocyte must therefore migrate via the blood circula- tion to the thymus. After have survived thymic
selection processes (1–3%), they are transported to peripheral secondary lymphoid organs via blood and
lymphatic circulation systems.
The secondary or peripheral lymphoid organs are the specialized sites where B and T lymphocytes and
antigen presenting cells come together to initiate immune responses to foreign antigens. These include
lymph nodes, spleen, and mucosa-associated lymphoid tissue (MALT), e.g. the palatine and
nasopharyngeal tonsils, Peyer’s patches in the small intestine, lymphoid nodules in the respiratory and
urogenital systems, the skin and conjunctiva of the eye.
Lymphocytes enter secondary lymphoid tissues from the blood. In the spleen, lymphocyte entry and exit
is via the marginal zone and venous drainage respectively. Antigen presenting cells (dendritic cells) enter
via the lymphatics, bringing with them antigen from peripheral infected sites. After activation, functionally
competent lymphocytes migrate to other sites in the body, where they combat the original infection.

LYMPH NODES
Lymph nodes are encapsulated centres of antigen presen- tation and lymphocyte activation,
differentiation and proliferation. They generate mature, antigen-primed, B and T cells, and filter particles,
including microbes, from the lymph by the action of numerous phagocytic macrophages. A normal young
adult body contains up to 450 lymph nodes, of which 60–70 are found in the head and neck, 100 in the
thorax and as many as 250 in the abdomen and pelvis. Lymph nodes are particularly numerous in the
neck, mediastinum, posterior abdominal wall, abdominal mesenteries, pelvis and proximal regions of the
limbs (axillary and inguinal lymph nodes).

Microstructure

Lymph nodes are small, oval bodies, 0.1– 2.5 cm long, lying along the course of the lymphatic vessels.
Each usually has a slight indentation, the hilum, through which blood vessels enter and leave and the
efferent lymphatic vessel leaves. Several afferent lymphatic vessels enter the capsule around the
periphery. Lymph nodes have a highly cellular cortex and a medulla which contains a network of
lymphatic channels through which lymph from the afferent lymphatics is filtered, to be collected by the
efferent lymphatic. At the hilum, the medulla reaches the surface.
The capsule is composed mainly of collagen fibres, elastin fibres and fibroblasts. From the capsule,
trabeculae of dense connective tissue extend radially into the interior of the node.
The fine reticulin bundles branch and interconnect to form a very dense network in the cortex. They
provide attachment for various cells, mostly dendritic cells, macrophages and lymphocytes.

Lymphatic and vascular supply
Lymph nodes are permeated by channels through which lymph percolates after its entry from the afferent

A primary follicle is uniformly populated by small. In the ‘dark zone’ the B cells (centroblasts) undergo rapid proliferation which is associated with hypermutation of their antibody molecules. Capillaries are especially profuse around the follicles. . some of which become memory B cells and join the recirculating pool. These dendritic cells include Langerhans cells from the skin and other squamous epithelia which have migrated into the draining lymph nodes. T cells are also present. where they interact with the FDCs which carry intact unprocessed antigen on their surface: the centrocytes compete for binding to the antigen and only the most suitable B cells can survive. while others leave to mature as antibody. They are exposed to antigens present in the lumen because the epithelium samples and transfers these antigens to antigen-presenting cells in the underlying tissues.and B-cell immunity. Anatomically. The latter become confluent at the hilum with the efferent vessel which drains the node. Appropriate clones of T and B cells in local lymphoid tissues are then activated and amplified prior to their exit via the lymphatics. which drain interstitial fluid as lymph. The density of the capillary beds increases greatly when lymphocytes multiply in response to antigenic stimulation. notably the peripharyngeal lymphoid (Waldeyer’s) ring of tonsillar tissue (palatine. Follicle-associated epithelium (?) The epithelium covering MALT is unusual in possessing cells which are involved in sampling antigens and passing them to the underlying tissues. Throughout the body. They then move into the light zone (as centrocytes). which are populated mainly by B cells and specialized follicular dendritic cells (FDC). tubal and lingual). which are not organized into follicles and interdigitating dendritic cells. Cells and cellular zones of lymph nodes In the cortex. where they coalesce as larger medullary sinuses. After antigen activation. The function of cells in these zones is similar to that found in lymph nodes. whereas a secondary follicle has a germinal centre which is composed mainly of antigen-stimulated B cells. eventually returning to venules and veins. Their role is to present processed antigen to T cells. branching medullary cords between which the reticulin network is seen. FDCs and macrophages. MUCOSA-ASSOCIATED LYMPHOID TISSUE (MALT) Large amounts of unencapsulated lymphoid tissue exist in the walls of the alimentary. and is populated mainly by T cells. quiescent lymphocytes. Arteries and veins serving lymph nodes pass through the hilum and traverse the medulla. After numerous mitotic divisions the selected B cells give rise to small lymphocytes. In the small and large intestine these specialized epithelial cells have characteristic short microvilli on their luminal surface and are known as microfold (M) cells. forming irregular. helping the survival of the B cells and inducing class switching. The deep cortex or paracortex lies between the cortical follicles and the medulla. cells are densely packed and in the outer cortical area they form lymphoid follicles or nodules. MALT includes an extremely large population of lymphocytes. In the palatine tonsils they include modified stratified squamous reticulated epithelial cells. lymphocytes are much less densely packed. Numerous radial cortical sinuses lead to the medulla. LYMPHOID TISSUES AND HAEMOPOIESIS The role of the germinal centre is to provide a microenvironment which allows the affinity maturation of the B cell response. Postcapillary HEV are abundant in the paracortical zones. The mantle zone surrounding the rapidly growing germinal centre is populated by mainly quiescent B cells. They form an important site of blood-borne lymphocyte extravasation into lymphoid tissue. The close proximity of lymphocytes within the MALT to an epithelial surface facilitates their access to pathogens. a few helper T cells. Macrophages in the germinal centre phagocytose apoptotic lymphocytes. In the cortex. In the medulla. respiratory. the main subclasses are gut-associated lymphoid tissue (GALT) and bronchusassociated lymphoid tissue (BALT). Lymphocyte populations are supported by a network of fine type III collagen fibres. MALT lacks afferent lymphatic vessels. The main function of B lymphocytes in MALT is to produce IgA for secretion into the lumen of the tracts which they line. BLOOD. MALT includes macroscopically visible lymphoid masses. Afferent lymphatic vessels enter at the periphery and then open into the subcapsular sinus. lymphocytes travel via the regional lymph nodes to disperse widely along mucosal surfaces to provide protective T. and in the skin: they are collectively termed mucosa-associated lymphoid tissue (MALT).vessels. and the Peyer’s patches of the small intestine. arteries form dense arcades of numerous anastomosing loops. Lymphocytes migrate into MALT through its HEV and leave mainly via its efferent lymphatics. The lymphoid cells are located in the lamina propria and in the submucosa as discrete follicles or nodules and in the lamina propria and in the base of the epithelium as scattered cells. Macrophages and B and T lymphocytes contact the lymph when flowing in the endothelium of channels or along their reticular fibers. nasopharyngeal.secreting plasma cells. reproductive and urinary tracts.

Thus. are also scattered throughout the entire mucosal lamina propria. Similar cells. many of the lymphocytes migrating between cells in the basal regions of epithelia are effector cytotoxic and helper T cells that have already been selected in lymphoid nodules and are engaged in immune responses. . and activated IgA-producing B cells and plasma cells.