Lymphatic system
The lymphatic system, or lymphoid system, is an organ system in vertebrates that is part of the immune system and complementary to the circulatory system. It consists of a large network of lymphatic vessels, lymph nodes, lymphoid organs, lymphatic tissue and lymph. The Latin word for lymph, lympha, refers to the deity of fresh water, "Lympha".
Unlike the circulatory system, which is a closed system, the lymphatic system is open. Lymph originates in the interstitial fluid that leaks from blood in the circulatory system into the tissues of the body. This fluid carries nutrients to the cells and collects waste products, bacteria, and damaged cells, before draining into the lymphatic vessels as lymph. The circulatory system processes an average of of blood per day through capillary filtration, which removes plasma from the blood. Roughly of the filtered blood is reabsorbed directly into the blood vessels, while the remaining are left in the interstitial fluid. The lymphatic system provides an accessory return route to the blood for this remainder.
The other main function is that of immune defense. Lymph is very similar to blood plasma, in that it contains waste products and cellular debris, together with bacteria and proteins. The cells of the lymph are mostly lymphocytes. Associated lymphoid organs are composed of lymphoid tissue, and are the sites either of lymphocyte production or of lymphocyte activation. These include the lymph nodes, the spleen, the thymus, and the tonsils. Lymphocytes are initially generated in the bone marrow. The lymphoid organs also contain other types of cells such as stromal cells for support. Lymphoid tissue is also associated with mucosas such as mucosa-associated lymphoid tissue.
These vessels carry the lymph throughout the body, passing through numerous lymph nodes which filter out unwanted materials such as bacteria and damaged cells. Lymph then passes into much larger lymph vessels known as lymph ducts. The right lymphatic duct drains the right side of the region and the much larger left lymphatic duct, known as the thoracic duct, drains the left side of the body. The ducts empty into the subclavian veins to return to the blood circulation. Lymph is moved through the system by muscle contractions. In some vertebrates, a lymph heart is present that pumps the lymph to the veins.
The lymphatic system was first described in the 17th century independently by Olaus Rudbeck and Thomas Bartholin.
Structure
The lymphatic system consists of a conducting network of lymphatic vessels, lymphoid organs, lymphoid tissues, and the circulating lymph.Primary lymphoid organs
The primary lymphoid organs, including the thymus, bone marrow, fetal liver and yolk sac, are responsible for generating lymphocytes from immature progenitor cells in the absence of antigens. The thymus and the bone marrow constitute the primary lymphoid organs involved in the production and early clonal selection of lymphocyte tissues.Bird species' primary lymphoid organs include the bone marrow, thymus, bursa of Fabricius, and yolk sac.
Red bone marrow
Bone marrow- specifically red bone marrow- is responsible for both the creation of T cell precursors and the production and maturation of B cells, which are known as lymphocytes, important cell types of the immune system. From the red bone marrow, B cells immediately join the circulatory system and travel to secondary lymphoid organs in search of pathogens. T cells, on the other hand, travel from the bone marrow to the thymus, where they undergo a multistage process to develop immunocompetency. Within the thymic cortex, developing T cells rearrange their T-cell receptor genes and are subjected to positive selection, which ensures that only T cells capable of recognizing self-major histocompatibility complex molecules survive. Those that bind too weakly die off. Surviving T cells then move to the thymic medulla, where negative selection removes T cells that bind self-antigens with high affinity, preventing autoimmune disorders. Less than 5% of T cells pass both selection tests and reach maturity; these mature T cells then join B cells in search of pathogens.Thymus
The thymus increases in size from birth in response to postnatal antigen stimulation. It is most active during the neonatal and pre-adolescent periods. The thymus is located between the inferior neck and the superior thorax. At puberty, by the early teens, the thymus begins to atrophy and regress, with adipose tissue mostly replacing the thymic stroma. However, residual T cell lymphopoiesis continues throughout adulthood, providing some immune response. The thymus is where the T lymphocytes mature and become immunocompetent. The loss or lack of the thymus results in severe immunodeficiency and subsequent high susceptibility to infection. In most species, the thymus consists of lobules divided by septa, which are made up of epithelium, which is often considered an epithelial organ. T cells mature from thymocytes, proliferate, and undergo a selection process in the thymic cortex before entering the medulla to interact with epithelial cells.Research on bony fish showed a buildup of T cells in the thymus and spleen of lymphoid tissues in salmon and showed that there are not many T cells in non-lymphoid tissues.
The thymus provides an inductive environment for developing T cells from hematopoietic progenitor cells. In addition, thymic stromal cells allow for the selection of a functional and self-tolerant T cell repertoire. Therefore, one of the most important roles of the thymus is the induction of central tolerance. However, the thymus is not where the infection is fought, as the T cells have yet to become immunocompetent.
Secondary lymphoid organs
The secondary lymphoid organs, which include lymph nodes and the spleen, maintain mature naive lymphocytes and initiate an adaptive immune response. The secondary lymphoid organs are the sites of lymphocyte activation by antigens. Activation leads to clonal selection and affinity maturation. Mature lymphocytes recirculate between the blood and the secondary lymphoid organs until they encounter their specific antigen.Spleen
The main functions of the spleen are:- to produce immune cells to fight antigens,
- to remove particulate matter and aged blood cells, mainly red blood cells, and
- to produce blood cells during fetal life.
Research on bony fish has shown that a high concentration of T cells is found in the spleen's white pulp.
Like the thymus, the spleen has only efferent lymphatic vessels. Both the short gastric arteries and the splenic artery supply it with blood. The germinal centers are supplied by arterioles called penicilliary radicles.
In humans, until the fifth month of prenatal development, the spleen creates red blood cells; after birth, the bone marrow is solely responsible for hematopoiesis. As a major lymphoid organ and a central player in the reticuloendothelial system, the spleen retains the ability to produce lymphocytes. The spleen stores red blood cells and lymphocytes. It can store enough blood cells to help in an emergency. During acute blood loss the spleen contracts to release stored erythrocytes, helping to temporarily maintain blood volume and oxygen delivery. Up to 25% of lymphocytes can be stored at any one time.
Lymph nodes
A lymph node is an organized collection of lymphoid tissue through which the lymph passes on its way back to the blood. Lymph nodes are located at intervals along the lymphatic system. Several afferent lymph vessels bring in lymph, which percolates through the substance of the lymph node and is then drained out by an efferent lymph vessel. Of the nearly 800 lymph nodes in the human body, about 300 are located in the head and neck. Many are grouped in clusters in different regions, as in the underarm and abdominal areas. Lymph node clusters are commonly found at the proximal ends of limbs and in the neck, where lymph is collected from body regions likely to sustain pathogen contamination from injuries. Lymph nodes are particularly numerous in the mediastinum in the chest, neck, pelvis, axilla, groin, and in association with the blood vessels of the intestines.The substance of a lymph node consists of lymphoid follicles in an outer portion called the cortex. The inner portion of the node is called the medulla, which is surrounded by the cortex on all sides except for a portion known as the hilum. The hilum presents as a depression on the surface of the lymph node, causing the otherwise spherical lymph node to be bean-shaped or ovoid. The efferent lymph vessel directly emerges from the lymph node at the hilum. The arteries and veins supplying the lymph node with blood enter and exit through the hilum. The region of the lymph node called the paracortex immediately surrounds the medulla. Unlike the cortex, which has primarily immature T cells, the paracortex has a mixture of immature and mature T cells. Lymphocytes enter the lymph nodes through specialised high endothelial venules found in the paracortex.
A lymph follicle is a dense collection of lymphocytes, the number, size, and configuration of which change in accordance with the functional state of the lymph node. For example, the follicles expand significantly when encountering a foreign antigen. The selection of B cells occurs in the germinal centre of the lymph nodes.
Secondary lymphoid tissue provides the environment for the foreign or altered native molecules to interact with the lymphocytes. It is exemplified by the lymph nodes, and the lymphoid follicles in tonsils, Peyer's patches, spleen, adenoids, skin, etc. that are associated with the mucosa-associated lymphoid tissue.
In the gastrointestinal wall, the appendix has mucosa resembling that of the colon, but it is heavily infiltrated with lymphocytes here.