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The lymphatic system, or lymphoid system, is an organ system in vertebrates that is part of the
immune system The immune system is a network of biological processes that protects an organism from diseases. It detects and responds to a wide variety of pathogens, from viruses to parasitic worms, as well as Tumor immunology, cancer cells and objects such ...
, and complementary to the
circulatory system The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, tha ...
. It consists of a large network of lymphatic vessels, lymph nodes, lymphatic or lymphoid organs, and lymphoid tissues. The vessels carry a clear fluid called lymph (the Latin word ''lympha'' refers to the deity of fresh water, " Lympha") back towards the
heart The heart is a muscular organ found in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as carbon diox ...
, for re-circulation. Unlike the circulatory system that is a closed system, the lymphatic system is open. The human circulatory system processes an average of 20 litres of
blood Blood is a body fluid in the circulatory system of humans and other vertebrates that delivers necessary substances such as nutrients and oxygen to the cells, and transports metabolic waste products away from those same cells. Blood in th ...
per day through capillary filtration, which removes plasma from the
blood Blood is a body fluid in the circulatory system of humans and other vertebrates that delivers necessary substances such as nutrients and oxygen to the cells, and transports metabolic waste products away from those same cells. Blood in th ...
. Roughly 17 litres of the filtered blood is reabsorbed directly into the
blood vessel Blood vessels are the structures of the circulatory system that transport blood throughout the human body. These vessels transport blood cells, nutrients, and oxygen to the tissues of the body. They also take waste and carbon dioxide away from ...
s, while the remaining three litres are left in the
interstitial fluid In cell biology, extracellular fluid (ECF) denotes all body fluid outside the cells of any multicellular organism. Total body water in healthy adults is about 60% (range 45 to 75%) of total body weight; women and the obese typically have a l ...
. One of the main functions of the lymphatic system is to provide an accessory return route to the blood for the surplus three litres. 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 Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...
and
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respon ...
s. 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 (where the highest lymphocyte concentration is found), the spleen, the thymus, and the tonsils. Lymphocytes are initially generated in the
bone marrow Bone marrow is a semi-solid tissue found within the spongy (also known as cancellous) portions of bones. In birds and mammals, bone marrow is the primary site of new blood cell production (or haematopoiesis). It is composed of hematopoieti ...
. 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 (MALT). Fluid from circulating blood leaks into the tissues of the body by capillary action, carrying nutrients to the cells. The fluid bathes the tissues as interstitial fluid, collecting waste products, bacteria, and damaged cells, and then drains as lymph into the lymphatic capillaries and lymphatic vessels. 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 (or central) lymphoid organs generate lymphocytes from immature progenitor cells. The thymus and the
bone marrow Bone marrow is a semi-solid tissue found within the spongy (also known as cancellous) portions of bones. In birds and mammals, bone marrow is the primary site of new blood cell production (or haematopoiesis). It is composed of hematopoieti ...
constitute the primary lymphoid organs involved in the production and early clonal selection of lymphocyte tissues.


Bone marrow

Bone marrow is responsible for both the creation of
T cell A T cell is a type of lymphocyte. T cells are one of the important white blood cells of the immune system and play a central role in the adaptive immune response. T cells can be distinguished from other lymphocytes by the presence of a T-cell ...
precursors and the production and maturation of B cells, which are important cell types of the immune system. From the 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 develop further and mature. Mature T cells then join B cells in search of pathogens. The other 95% of T cells begin a process of apoptosis, a form of
programmed cell death Programmed cell death (PCD; sometimes referred to as cellular suicide) is the death of a cell (biology), cell as a result of events inside of a cell, such as apoptosis or autophagy. PCD is carried out in a biological process, which usually confers ...
.


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 adult life, 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 Salmon () is the common name In biology, a common name of a taxon or organism (also known as a vernacular name, English name, colloquial name, country name, popular name, or farmer's name) is a name that is based on the normal language of ...
and showed that there are not many T cells in non-lymphoid tissues. The thymus provides an inductive environment for the development of 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 (or peripheral) lymphoid organs (SLO), 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
antigen In immunology, an antigen (Ag) is a molecule or molecular structure or any foreign particulate matter or a pollen grain that can bind to a specific antibody or T-cell receptor. The presence of antigens in the body may trigger an immune response. ...
s. Activation leads to clonal expansion 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
antigen In immunology, an antigen (Ag) is a molecule or molecular structure or any foreign particulate matter or a pollen grain that can bind to a specific antibody or T-cell receptor. The presence of antigens in the body may trigger an immune response. ...
s # to remove
particulate matter Particulates – also known as atmospheric aerosol particles, atmospheric particulate matter, particulate matter (PM) or suspended particulate matter (SPM) – are microscopic particles of solid or liquid matter suspended in the air. The t ...
and aged blood cells, mainly
red blood cell Red blood cells (RBCs), also referred to as red cells, red blood corpuscles (in humans or other animals not having nucleus in red blood cells), haematids, erythroid cells or erythrocytes (from Greek ''erythros'' for "red" and ''kytos'' for "holl ...
s # to produce blood cells during fetal life. The spleen synthesizes antibodies in its white pulp and removes antibody-coated bacteria and antibody-coated blood cells by way of blood and lymph node circulation. The white pulp of the spleen provides immune fucntion due to the lymphocytes that are housed there. The spleen also consists of red pulp which is responsible for getting rid of aged red blood cells, as well as pathogens. This is carried out by macrophages present in the red pulp. A study published in 2009 using mice found that the spleen contains, in its reserve, half of the body's
monocyte Monocytes are a type of leukocyte or white blood cell. They are the largest type of leukocyte in blood and can differentiate into macrophages and conventional dendritic cells. As a part of the vertebrate innate immune system monocytes also i ...
s within the red pulp. These monocytes, upon moving to injured tissue (such as the heart), turn into
dendritic cell Dendritic cells (DCs) are antigen-presenting cells (also known as ''accessory cells'') of the mammalian immune system. Their main function is to process antigen material and present it on the cell surface to the T cells of the immune system. ...
s and macrophages while promoting tissue healing. The spleen is a center of activity of the mononuclear phagocyte system and can be considered analogous to a large lymph node, as its absence causes a predisposition to certain
infection An infection is the invasion of tissues by pathogens, their multiplication, and the reaction of host tissues to the infectious agent and the toxins they produce. An infectious disease, also known as a transmissible disease or communicable d ...
s. Notably, the spleen is important for a multitude of functions. The spleen removes pathogens and old erythrocytes from the blood (red pulp) and produces lymphocytes for immune response (white pulp). The spleen also is responsible for recycling some erythrocytes components and discarding others. For example, hemoglobin is broken down into amino acids that are reused. Research on bony fish has shown that a high concentration of T cells are found in the white pulp of the spleen.ref name="Koppang"/> Like the thymus, the spleen has only
efferent lymphatic vessels The lymphatic vessels (or lymph vessels or lymphatics) are thin-walled vessels (tubes), structured like blood vessels, that carry lymph. As part of the lymphatic system, lymph vessels are complementary to the cardiovascular system. Lymph vessel ...
. Both the short gastric arteries and the splenic artery supply it with blood. The germinal centers are supplied by arterioles called ''penicilliary radicles''. In the human until the fifth month of
prenatal development Prenatal development () includes the development of the embryo and of the fetus during a viviparous animal's gestation. Prenatal development starts with fertilization, in the germinal stage of embryonic development, and continues in fetal devel ...
, the spleen creates red blood cells; after birth, the
bone marrow Bone marrow is a semi-solid tissue found within the spongy (also known as cancellous) portions of bones. In birds and mammals, bone marrow is the primary site of new blood cell production (or haematopoiesis). It is composed of hematopoieti ...
is solely responsible for
hematopoiesis Haematopoiesis (, from Greek , 'blood' and 'to make'; also hematopoiesis in American English; sometimes also h(a)emopoiesis) is the formation of blood cellular components. All cellular blood components are derived from haematopoietic stem cells ...
. 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. 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 (groin, armpits) and in the neck, where lymph is collected from regions of the body likely to sustain pathogen contamination from injuries. Lymph nodes are particularly numerous in the mediastinum in the chest, neck, pelvis,
axilla The axilla (also, armpit, underarm or oxter) is the area on the human body directly under the shoulder joint. It includes the axillary space, an anatomical space within the shoulder girdle between the arm and the thoracic cage, bounded supe ...
, inguinal region, 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 mostly immature T cells, or thymocytes, 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, or ''B lymphocytes'', occurs in the germinal centre of the lymph nodes. Secondary lymphoid tissue provides the environment for the foreign or altered native molecules (antigens) to interact with the lymphocytes. It is exemplified by the lymph nodes, and the lymphoid follicles in tonsils, Peyer's patches, spleen, adenoids,
skin Skin is the layer of usually soft, flexible outer tissue covering the body of a vertebrate animal, with three main functions: protection, regulation, and sensation. Other cuticle, animal coverings, such as the arthropod exoskeleton, have diffe ...
, etc. that are associated with the mucosa-associated lymphoid tissue (MALT). In the
gastrointestinal wall The gastrointestinal wall of the gastrointestinal tract is made up of four layers of specialised tissue. From the inner cavity of the gut (the lumen) outwards, these are: # Mucosa # Submucosa # Muscular layer # Serosa or adventitia The mucosa is ...
, the
appendix Appendix, or its plural form appendices, may refer to: __NOTOC__ In documents * Addendum, an addition made to a document by its author after its initial printing or publication * Bibliography, a systematic list of books and other works * Index (pub ...
has mucosa resembling that of the colon, but here it is heavily infiltrated with lymphocytes.


Tertiary lymphoid organs

Tertiary lymphoid organs (TLOs) are abnormal lymph node-like structures that form in peripheral tissues at sites of chronic inflammation, such as chronic infection,
transplanted organs Organ transplantation is a medical procedure in which an organ is removed from one body and placed in the body of a recipient, to replace a damaged or missing organ. The donor and recipient may be at the same location, or organs may be transpor ...
undergoing graft rejection, some
cancer Cancer is a group of diseases involving abnormal cell growth with the potential to invade or spread to other parts of the body. These contrast with benign tumors, which do not spread. Possible signs and symptoms include a lump, abnormal bl ...
s, and
autoimmune In immunology, autoimmunity is the system of immune responses of an organism against its own healthy cells, tissues and other normal body constituents. Any disease resulting from this type of immune response is termed an " autoimmune disease" ...
and autoimmune-related diseases. TLOs are regulated differently from the normal process whereby lymphoid tissues are formed during
ontogeny Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the egg to adult. The term can also be used to refer to the stu ...
, being dependent on cytokines and
hematopoietic Haematopoiesis (, from Greek , 'blood' and 'to make'; also hematopoiesis in American English; sometimes also h(a)emopoiesis) is the formation of blood cellular components. All cellular blood components are derived from haematopoietic stem cells ...
cells, but still drain
interstitial fluid In cell biology, extracellular fluid (ECF) denotes all body fluid outside the cells of any multicellular organism. Total body water in healthy adults is about 60% (range 45 to 75%) of total body weight; women and the obese typically have a l ...
and transport lymphocytes in response to the same chemical messengers and gradients. TLOs typically contain far fewer lymphocytes, and assume an immune role only when challenged with
antigen In immunology, an antigen (Ag) is a molecule or molecular structure or any foreign particulate matter or a pollen grain that can bind to a specific antibody or T-cell receptor. The presence of antigens in the body may trigger an immune response. ...
s that result in
inflammation Inflammation (from la, wikt:en:inflammatio#Latin, inflammatio) is part of the complex biological response of body tissues to harmful stimuli, such as pathogens, damaged cells, or Irritation, irritants, and is a protective response involving im ...
. They achieve this by importing the lymphocytes from blood and lymph. TLOs often have an active germinal center, surrounded by a network of follicular dendritic cells (FDCs). TLOs are thought to play an important role in the immune response to cancer and to have possible implications in immunotherapy. They have been observed in a number of cancer types such as melanoma, non-small cell lung cancer and colorectal cancer (reviewed in ) as well as glioma. Patients with TLOs in the vicinity of their tumors tend to have a better prognosis, although the opposite is true for certain cancers. TLOs that contain an active germinal center tend to have a better prognosis than those with TLOs without a germinal center. The reason that these patients tend to live longer is thought to be the immune response against the tumor, which is mediated by the TLOs. TLOs may also promote an anti-tumor response when patients are treated with immunotherapy. TLOs have been referred to in many different ways, including as tertiary lymphoid structures (TLS) and ectopic lymphoid structures (ELS). When associated with colorectal cancer, they are often referred to as a Crohn's-like lymphoid reaction.


Other lymphoid tissue

Lymphoid tissue associated with the lymphatic system is concerned with immune functions in defending the body against
infection An infection is the invasion of tissues by pathogens, their multiplication, and the reaction of host tissues to the infectious agent and the toxins they produce. An infectious disease, also known as a transmissible disease or communicable d ...
s and the spread of
tumour A neoplasm () is a type of abnormal and excessive growth of tissue. The process that occurs to form or produce a neoplasm is called neoplasia. The growth of a neoplasm is uncoordinated with that of the normal surrounding tissue, and persists ...
s. It consists of
connective tissue Connective tissue is one of the four primary types of animal tissue, along with epithelial tissue, muscle tissue, and nervous tissue. It develops from the mesenchyme derived from the mesoderm the middle embryonic germ layer. Connective tissue ...
formed of reticular fibers, with various types of
leukocyte White blood cells, also called leukocytes or leucocytes, are the cells of the immune system that are involved in protecting the body against both infectious disease and foreign invaders. All white blood cells are produced and derived from mul ...
s (white blood cells), mostly lymphocytes enmeshed in it, through which the lymph passes. Regions of the lymphoid tissue that are densely packed with lymphocytes are known as ''lymphoid follicles''. Lymphoid tissue can either be structurally well organized as lymph nodes or may consist of loosely organized lymphoid follicles known as the mucosa-associated lymphoid tissue (MALT). The
central nervous system The central nervous system (CNS) is the part of the nervous system consisting primarily of the brain and spinal cord. The CNS is so named because the brain integrates the received information and coordinates and influences the activity of all p ...
also has lymphatic vessels. The search for T cell gateways into and out of the meninges uncovered functional
meningeal lymphatic vessels The meningeal lymphatic vessels (or meningeal lymphatics) are a network of conventional lymphatic vessels located parallel to the dural venous sinuses and middle meningeal arteries of the mammalian central nervous system (CNS). As a part of the lym ...
lining the dural sinuses, anatomically integrated into the membrane surrounding the brain. *


Lymphatic vessels

The lymphatic vessels, also called lymph vessels, are thin-walled vessels that conduct lymph between different parts of the body. They include the tubular vessels of the lymph capillaries, and the larger collecting vessels–the right lymphatic duct and the thoracic duct (the left lymphatic duct). The lymph capillaries are mainly responsible for the absorption of interstitial fluid from the tissues, while lymph vessels propel the absorbed fluid forward into the larger collecting ducts, where it ultimately returns to the bloodstream via one of the subclavian veins. The tissues of the lymphatic system are responsible for maintaining the balance of the body fluids. Its network of capillaries and collecting lymphatic vessels work to efficiently drain and transport extravasated fluid, along with proteins and antigens, back to the circulatory system. Numerous intraluminal valves in the vessels ensure a unidirectional flow of lymph without reflux. Two valve systems, a primary and a secondary valve system, are used to achieve this unidirectional flow. The capillaries are blind-ended, and the valves at the ends of capillaries use specialised junctions together with anchoring filaments to allow a unidirectional flow to the primary vessels. The collecting lymphatics, however, act to propel the lymph by the combined actions of the intraluminal valves and lymphatic muscle cells.


Development

Lymphatic tissues begin to develop by the end of the fifth week of embryonic development. Lymphatic vessels develop from
lymph sacs Lymph sacs are a part of the development of the lymphatic system, known as lymphangiogenesis. The lymph sacs are precursors of the lymph vessels. These sacs develop through the processes of vasculogenesis and angiogenesis. However, there is ...
that arise from developing veins, which are derived from
mesoderm The mesoderm is the middle layer of the three germ layers that develops during gastrulation in the very early development of the embryo of most animals. The outer layer is the ectoderm, and the inner layer is the endoderm.Langman's Medical Emb ...
. The first lymph sacs to appear are the paired jugular lymph sacs at the junction of the internal jugular and subclavian veins. From the jugular lymph sacs, lymphatic capillary plexuses spread to the thorax, upper limbs, neck, and head. Some of the plexuses enlarge and form lymphatic vessels in their respective regions. Each jugular lymph sac retains at least one connection with its jugular vein, the left one developing into the superior portion of the thoracic duct. The spleen develops from mesenchymal cells between layers of the dorsal mesentery of the stomach. The thymus arises as an outgrowth of the third pharyngeal pouch.


Function

The lymphatic system has multiple interrelated functions: * It is responsible for the removal of
interstitial fluid In cell biology, extracellular fluid (ECF) denotes all body fluid outside the cells of any multicellular organism. Total body water in healthy adults is about 60% (range 45 to 75%) of total body weight; women and the obese typically have a l ...
from tissues * It absorbs and transports
fatty acid In chemistry, particularly in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated or unsaturated. Most naturally occurring fatty acids have an unbranched chain of an even number of carbon atoms, ...
s and fats as chyle from the digestive system * It transports white blood cells to and from the lymph nodes into the bones * The lymph transports antigen-presenting cells, such as
dendritic cells Dendritic cells (DCs) are antigen-presenting cells (also known as ''accessory cells'') of the mammalian immune system. Their main function is to process antigen material and present it on the cell surface to the T cells of the immune system. ...
, to the lymph nodes where an immune response is stimulated.


Fat absorption

Lymph vessels called lacteals are at the beginning of the
gastrointestinal tract The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans a ...
, predominantly in the small intestine. While most other nutrients absorbed by the
small intestine The small intestine or small bowel is an organ in the gastrointestinal tract where most of the absorption of nutrients from food takes place. It lies between the stomach and large intestine, and receives bile and pancreatic juice through t ...
are passed on to the portal venous system to drain via the portal vein into the
liver The liver is a major organ only found in vertebrates which performs many essential biological functions such as detoxification of the organism, and the synthesis of proteins and biochemicals necessary for digestion and growth. In humans, it ...
for processing, fats ( lipids) are passed on to the lymphatic system to be transported to the blood circulation via the thoracic duct. (There are exceptions, for example medium-chain triglycerides are fatty acid esters of glycerol that passively diffuse from the GI tract to the portal system.) The enriched lymph originating in the lymphatics of the
small intestine The small intestine or small bowel is an organ in the gastrointestinal tract where most of the absorption of nutrients from food takes place. It lies between the stomach and large intestine, and receives bile and pancreatic juice through t ...
is called chyle. The nutrients that are released into the circulatory system are processed by the
liver The liver is a major organ only found in vertebrates which performs many essential biological functions such as detoxification of the organism, and the synthesis of proteins and biochemicals necessary for digestion and growth. In humans, it ...
, having passed through the systemic circulation.


Immune function

The lymphatic system plays a major role in the body's immune system, as the primary site for cells relating to
adaptive immune system The adaptive immune system, also known as the acquired immune system, is a subsystem of the immune system that is composed of specialized, systemic cells and processes that eliminate pathogens or prevent their growth. The acquired immune system ...
including
T-cell A T cell is a type of lymphocyte. T cells are one of the important white blood cells of the immune system and play a central role in the adaptive immune response. T cells can be distinguished from other lymphocytes by the presence of a T-cell ...
s and B-cells. Cells in the lymphatic system react to antigens presented or found by the cells directly or by other
dendritic cells Dendritic cells (DCs) are antigen-presenting cells (also known as ''accessory cells'') of the mammalian immune system. Their main function is to process antigen material and present it on the cell surface to the T cells of the immune system. ...
. When an antigen is recognized, an immunological cascade begins involving the activation and recruitment of more and more cells, the production of antibodies and cytokines and the recruitment of other immunological cells such as
macrophages Macrophages (abbreviated as M φ, MΦ or MP) ( el, large eaters, from Greek ''μακρός'' (') = large, ''φαγεῖν'' (') = to eat) are a type of white blood cell of the immune system that engulfs and digests pathogens, such as cancer ce ...
.


Clinical significance

The study of lymphatic drainage of various organs is important in the diagnosis, prognosis, and treatment of cancer. The lymphatic system, because of its closeness to many tissues of the body, is responsible for carrying cancerous cells between the various parts of the body in a process called
metastasis Metastasis is a pathogenic agent's spread from an initial or primary site to a different or secondary site within the host's body; the term is typically used when referring to metastasis by a cancerous tumor. The newly pathological sites, then, ...
. The intervening lymph nodes can trap the cancer cells. If they are not successful in destroying the cancer cells the nodes may become sites of secondary tumours.


Enlarged lymph nodes

Lymphadenopathy refers to one or more enlarged lymph nodes. Small groups or individually enlarged lymph nodes are generally ''reactive'' in response to
infection An infection is the invasion of tissues by pathogens, their multiplication, and the reaction of host tissues to the infectious agent and the toxins they produce. An infectious disease, also known as a transmissible disease or communicable d ...
or
inflammation Inflammation (from la, wikt:en:inflammatio#Latin, inflammatio) is part of the complex biological response of body tissues to harmful stimuli, such as pathogens, damaged cells, or Irritation, irritants, and is a protective response involving im ...
. This is called ''local'' lymphadenopathy. When many lymph nodes in different areas of the body are involved, this is called ''generalised'' lymphadenopathy. Generalised lymphadenopathy may be caused by
infection An infection is the invasion of tissues by pathogens, their multiplication, and the reaction of host tissues to the infectious agent and the toxins they produce. An infectious disease, also known as a transmissible disease or communicable d ...
s such as infectious mononucleosis,
tuberculosis Tuberculosis (TB) is an infectious disease usually caused by ''Mycobacterium tuberculosis'' (MTB) bacteria. Tuberculosis generally affects the lungs, but it can also affect other parts of the body. Most infections show no symptoms, in w ...
and
HIV The human immunodeficiency viruses (HIV) are two species of '' Lentivirus'' (a subgroup of retrovirus) that infect humans. Over time, they cause acquired immunodeficiency syndrome (AIDS), a condition in which progressive failure of the immu ...
, connective tissue diseases such as SLE and
rheumatoid arthritis Rheumatoid arthritis (RA) is a long-term autoimmune disorder that primarily affects synovial joint, joints. It typically results in warm, swollen, and painful joints. Pain and stiffness often worsen following rest. Most commonly, the wrist and ...
, and
cancer Cancer is a group of diseases involving abnormal cell growth with the potential to invade or spread to other parts of the body. These contrast with benign tumors, which do not spread. Possible signs and symptoms include a lump, abnormal bl ...
s, including both cancers of tissue within lymph nodes, discussed below, and
metastasis Metastasis is a pathogenic agent's spread from an initial or primary site to a different or secondary site within the host's body; the term is typically used when referring to metastasis by a cancerous tumor. The newly pathological sites, then, ...
of cancerous cells from other parts of the body, that have arrived via the lymphatic system.


Lymphedema

Lymphedema is the swelling caused by the accumulation of lymph, which may occur if the lymphatic system is damaged or has malformations. It usually affects limbs, though the face, neck and abdomen may also be affected. In an extreme state, called elephantiasis, the edema progresses to the extent that the skin becomes thick with an appearance similar to the skin on
elephant Elephants are the largest existing land animals. Three living species are currently recognised: the African bush elephant, the African forest elephant, and the Asian elephant. They are the only surviving members of the family Elephantidae ...
limbs. Causes are unknown in most cases, but sometimes there is a previous history of severe infection, usually caused by a parasitic disease, such as
lymphatic filariasis Lymphatic filariasis is a human disease caused by parasitic worms known as filarial worms. Usually acquired in childhood, it is a leading cause of permanent disability worldwide. While most cases have no symptoms, some people develop a syndrome ...
. Lymphangiomatosis is a disease involving multiple cysts or lesions formed from lymphatic vessels. Lymphedema can also occur after surgical removal of lymph nodes in the armpit (causing the arm to swell due to poor lymphatic drainage) or groin (causing swelling of the leg). Conventional treatment is by manual lymphatic drainage and compression garments. Two drugs for the treatment of lymphedema are in clinical trials: Lymfactin and Ubenimex/ Bestatin. There is no evidence to suggest that the effects of manual lymphatic drainage are permanent.


Cancer

Cancer Cancer is a group of diseases involving abnormal cell growth with the potential to invade or spread to other parts of the body. These contrast with benign tumors, which do not spread. Possible signs and symptoms include a lump, abnormal bl ...
of the lymphatic system can be primary or secondary.
Lymphoma Lymphoma is a group of blood and lymph tumors that develop from lymphocytes (a type of white blood cell). In current usage the name usually refers to just the cancerous versions rather than all such tumours. Signs and symptoms may include enl ...
refers to cancer that arises from lymphatic tissue. Lymphoid leukaemias and lymphomas are now considered to be tumours of the same type of cell lineage. They are called "leukaemia" when in the blood or marrow and "lymphoma" when in lymphatic tissue. They are grouped together under the name "lymphoid malignancy". Lymphoma is generally considered as either
Hodgkin lymphoma Hodgkin lymphoma (HL) is a type of lymphoma, in which cancer originates from a specific type of white blood cell called lymphocytes, where multinucleated Reed–Sternberg cells (RS cells) are present in the patient's lymph nodes. The condition ...
or non-Hodgkin lymphoma. Hodgkin lymphoma is characterised by a particular type of cell, called a Reed–Sternberg cell, visible under microscope. It is associated with past infection with the
Epstein–Barr virus The Epstein–Barr virus (EBV), formally called ''Human gammaherpesvirus 4'', is one of the nine known human herpesvirus types in the herpes family, and is one of the most common viruses in humans. EBV is a double-stranded DNA virus. It is ...
, and generally causes a painless "rubbery" lymphadenopathy. It is staged, using Ann Arbor staging.
Chemotherapy Chemotherapy (often abbreviated to chemo and sometimes CTX or CTx) is a type of cancer treatment that uses one or more anti-cancer drugs (chemotherapeutic agents or alkylating agents) as part of a standardized chemotherapy regimen. Chemothe ...
generally involves the ABVD and may also involve
radiotherapy Radiation therapy or radiotherapy, often abbreviated RT, RTx, or XRT, is a therapy using ionizing radiation, generally provided as part of cancer treatment to control or kill malignant cells and normally delivered by a linear accelerator. Ra ...
. Non-Hodgkin lymphoma is a cancer characterised by increased proliferation of B-cells or T-cells, generally occurs in an older age group than Hodgkin lymphoma. It is treated according to whether it is '' high-grade'' or ''low-grade'', and carries a poorer prognosis than Hodgkin lymphoma. Lymphangiosarcoma is a malignant soft tissue tumour, whereas lymphangioma is a benign tumour occurring frequently in association with
Turner syndrome Turner syndrome (TS), also known as 45,X, or 45,X0, is a genetic condition in which a female is partially or completely missing an X chromosome. Signs and symptoms vary among those affected. Often, a short and webbed neck, low-set ears, low h ...
. Lymphangioleiomyomatosis is a benign tumour of the smooth muscles of the lymphatics that occurs in the lungs. Lymphoid leukaemia is another form of cancer where the host is devoid of different lymphatic cells.


Other

* Castleman's disease * Chylothorax *
Kawasaki disease Kawasaki disease is a syndrome of unknown cause that results in a fever and mainly affects children under 5 years of age. It is a form of vasculitis, where blood vessels become inflamed throughout the body. The fever typically lasts for more t ...
* Kikuchi disease * Lipedema * Lymphangitis *
Lymphatic filariasis Lymphatic filariasis is a human disease caused by parasitic worms known as filarial worms. Usually acquired in childhood, it is a leading cause of permanent disability worldwide. While most cases have no symptoms, some people develop a syndrome ...
* Lymphocytic choriomeningitis *
Solitary lymphatic nodule The Solitary lymphatic nodules (or solitary follicles) are structures found in the small intestine and large intestine. Small intestine The solitary lymphatic nodules are found scattered throughout the mucous membrane of the small intestine, but ...


History

Hippocrates Hippocrates of Kos (; grc-gre, Ἱπποκράτης ὁ Κῷος, Hippokrátēs ho Kôios; ), also known as Hippocrates II, was a Greek physician of the classical period who is considered one of the most outstanding figures in the history o ...
, in the 5th century BC, was one of the first people to mention the lymphatic system. In his work ''On Joints'', he briefly mentioned the lymph nodes in one sentence. Rufus of
Ephesus Ephesus (; grc-gre, Ἔφεσος, Éphesos; tr, Efes; may ultimately derive from hit, 𒀀𒉺𒊭, Apaša) was a city in ancient Greece on the coast of Ionia, southwest of present-day Selçuk in İzmir Province, Turkey. It was built i ...
, a Roman physician, identified the axillary, inguinal and mesenteric lymph nodes as well as the thymus during the 1st to 2nd century AD. The first mention of lymphatic vessels was in the 3rd century BC by Herophilos, a Greek anatomist living in
Alexandria Alexandria ( or ; ar, ٱلْإِسْكَنْدَرِيَّةُ ; grc-gre, Αλεξάνδρεια, Alexándria) is the second largest city in Egypt, and the largest city on the Mediterranean coast. Founded in by Alexander the Great, Alexandr ...
, who incorrectly concluded that the "absorptive veins of the lymphatics," by which he meant the lacteals (lymph vessels of the intestines), drained into the hepatic portal veins, and thus into the liver. The findings of Ruphus and Herophilos were further propagated by the Greek physician
Galen Aelius Galenus or Claudius Galenus ( el, Κλαύδιος Γαληνός; September 129 – c. AD 216), often Anglicized as Galen () or Galen of Pergamon, was a Greek physician, surgeon and philosopher in the Roman Empire. Considered to be on ...
, who described the lacteals and mesenteric lymph nodes which he observed in his dissection of apes and pigs in the 2nd century AD. In the mid 16th century, Gabriele Falloppio (discoverer of the
fallopian tube The fallopian tubes, also known as uterine tubes, oviducts or salpinges (singular salpinx), are paired tubes in the human female that stretch from the uterus to the ovaries. The fallopian tubes are part of the female reproductive system. In ...
s), described what is now known as the lacteals as "coursing over the intestines full of yellow matter." In about 1563 Bartolomeo Eustachi, a professor of anatomy, described the thoracic duct in horses as ''vena alba thoracis.'' The next breakthrough came when in 1622 a physician,
Gaspare Aselli Gaspare Aselli (or Asellio) ( – 9 September 1625). was an Italian physician noted for the discovery of the lacteal vessels of the lymphatic system. Aselli discovered (or rediscovered) the chylous vessels, and studied systematically the signifi ...
, identified lymphatic vessels of the intestines in dogs and termed them ''venae albae et lacteae,'' which are now known as simply the lacteals. The lacteals were termed the fourth kind of vessels (the other three being the artery, vein and nerve, which was then believed to be a type of vessel), and disproved Galen's assertion that chyle was carried by the veins. But, he still believed that the lacteals carried the chyle to the liver (as taught by Galen). He also identified the thoracic duct but failed to notice its connection with the lacteals. This connection was established by Jean Pecquet in 1651, who found a white fluid mixing with blood in a dog's heart. He suspected that fluid to be chyle as its flow increased when abdominal pressure was applied. He traced this fluid to the thoracic duct, which he then followed to a chyle-filled sac he called the ''chyli receptaculum,'' which is now known as the cisternae chyli; further investigations led him to find that lacteals' contents enter the venous system via the thoracic duct. Thus, it was proven convincingly that the lacteals did not terminate in the
liver The liver is a major organ only found in vertebrates which performs many essential biological functions such as detoxification of the organism, and the synthesis of proteins and biochemicals necessary for digestion and growth. In humans, it ...
, thus disproving Galen's second idea: that the chyle flowed to the liver. Johann Veslingius drew the earliest sketches of the lacteals in humans in 1641. The idea that blood recirculates through the body rather than being produced anew by the liver and the heart was first accepted as a result of works of
William Harvey William Harvey (1 April 1578 – 3 June 1657) was an English physician who made influential contributions in anatomy and physiology. He was the first known physician to describe completely, and in detail, the systemic circulation and prope ...
—a work he published in 1628. In 1652, Olaus Rudbeck (1630–1702), a Swede, discovered certain transparent vessels in the liver that contained clear fluid (and not white), and thus named them ''hepatico-aqueous vessels''. He also learned that they emptied into the thoracic duct and that they had valves. He announced his findings in the court of Queen Christina of Sweden, but did not publish his findings for a year, and in the interim similar findings were published by Thomas Bartholin, who additionally published that such vessels are present everywhere in the body, not just in the liver. He is also the one to have named them "lymphatic vessels." This had resulted in a bitter dispute between one of Bartholin's pupils, Martin Bogdan, and Rudbeck, whom he accused of
plagiarism Plagiarism is the fraudulent representation of another person's language, thoughts, ideas, or expressions as one's own original work.From the 1995 '' Random House Compact Unabridged Dictionary'': use or close imitation of the language and though ...
. Galen's ideas prevailed in medicine until the 17th century. It was thought that blood was produced by the liver from chyle contaminated with ailments by the intestine and stomach, to which various spirits were added by other organs, and that this blood was consumed by all the organs of the body. This theory required that the blood be consumed and produced many times over. Even in the 17th century, his ideas were defended by some physicians. Alexander Monro, of the University of Edinburgh Medical School, was the first to describe the function of the lymphatic system in detail. Image:Galen detail.jpg, "Claude Galien". Lithograph by Pierre Roche Vigneron. (Paris: Lith de Gregoire et Deneux, ca. 1865) Gabriele_Falloppio.jpg, Gabriele Falloppio Image:Bartolomeus Eustachius.jpg, Portrait of Eustachius File:Olaus Rudbeck Sr (portrait by Martin Mijtens Sr, 1696).jpg, Olaus Rudbeck in 1696. File:Thomas bartholin.jpg, Thomas Bartholin


Etymology

''Lymph'' originates in the
Classical Latin Classical Latin is the form of Literary Latin recognized as a literary standard by writers of the late Roman Republic and early Roman Empire. It was used from 75 BC to the 3rd century AD, when it developed into Late Latin. In some later pe ...
word ' "water", which is also the source of the English word ''limpid''. The spelling with ''y'' and ''ph'' was influenced by
folk etymology Folk etymology (also known as popular etymology, analogical reformation, reanalysis, morphological reanalysis or etymological reinterpretation) is a change in a word or phrase resulting from the replacement of an unfamiliar form by a more famili ...
with
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
(') "
nymph A nymph ( grc, νύμφη, nýmphē, el, script=Latn, nímfi, label=Modern Greek; , ) in ancient Greek folklore is a minor female nature deity. Different from Greek goddesses, nymphs are generally regarded as personifications of nature, are ty ...
". The adjective used for the lymph-transporting system is ''lymphatic''. The adjective used for the tissues where lymphocytes are formed is ''lymphoid''. Lymphatic comes from the Latin word ', meaning "connected to water."


See also

* List of lymphatic vessels of the human body * American Society of Lymphology * Glymphatic system and
Meningeal lymphatic vessels The meningeal lymphatic vessels (or meningeal lymphatics) are a network of conventional lymphatic vessels located parallel to the dural venous sinuses and middle meningeal arteries of the mammalian central nervous system (CNS). As a part of the lym ...
- equivalent for the
central nervous system The central nervous system (CNS) is the part of the nervous system consisting primarily of the brain and spinal cord. The CNS is so named because the brain integrates the received information and coordinates and influences the activity of all p ...
* Innate lymphoid cells * Lymphangiogenesis *
Lymphangion A lymphangion is the functional unit of a lymph vessel that lies between two semilunar (half moon-shaped) valves. Lymph vessels are channels larger than the lymph capillaries that have thicker walls, valves in their lumen and smooth muscles in ...
* Mononuclear phagocyte system *
Waldemar Olszewski Prof. Waldemar Lech Olszewski (born 3 September 1931 in Piastów, Warsaw, Poland) is a Polish lymphologist. His main area of study is the human lymphatic system. Clinical and research interests comprise vascular surgery, transplantation, physiol ...
– discovered fundamental processes in human tissues connected with function of the lymphatic system * Trogocytosis


References


External links


Lymphatic System


(innerbody.com) {{DEFAULTSORT:Lymphatic System Angiology Animal anatomy