Adaptive immune system - Knowledge (XXG)

1518:, which induce the production of interferons and other cytokines increasing resistance of cells such as monocytes to infections. Cytokines produced during innate immune responses are among the activators of adaptive immune responses. Antibodies exert additive or synergistic effects with mechanisms of innate immunity. Unstable HbS clusters Band-3, a major integral red cell protein; antibodies recognize these clusters and accelerate their removal by phagocytic cells. Clustered Band 3 proteins with attached antibodies activate complement, and complement C3 fragments are opsonins recognized by the CR1 complement receptor on phagocytic cells. 186: 1210:. Throughout the lifetime of an animal these memory cells form a database of effective B and T lymphocytes. Upon interaction with a previously encountered antigen, the appropriate memory cells are selected and activated. In this manner, the second and subsequent exposures to an antigen produce a stronger and faster immune response. This is "adaptive" in the sense that the body's immune system prepares itself for future challenges, but is "maladaptive" of course if the receptors are autoimmune. Immunological memory can be in the form of either 1388:, in which one gene segment recombines with other gene segments to form a single unique gene. This assembly process generates the enormous diversity of receptors and antibodies, before the body ever encounters antigens, and enables the immune system to respond to an almost unlimited diversity of antigens. Throughout an animal's lifetime, lymphocytes that can react against the antigens an animal actually encounters are selected for action—directed against anything that expresses that antigen. 1133: 377: 368:(NK) cells with so-far unexplained specificity for antigens, expansion of NK cells expressing germ-line encoded receptors, and activation of other innate immune cells to an activated state that confers a short-term "immune memory". In this sense, "adaptive immunity" more closely resembles the concept of "activated state" or "heterostasis", thus returning in sense to the physiological sense of "adaptation" to environmental changes. 560: 900: 724: 1346: 43: 1404:(which is considered "non-self") from attack by the immune system, are particularly interesting. Although no comprehensive explanation has emerged to explain this mysterious, and often repeated, lack of rejection, two classical reasons may explain how the fetus is tolerated. The first is that the fetus occupies a portion of the body protected by a non-immunological barrier, the 352:, a previously marginal organism for the study of immunology. The term "adaptive" as used in immunology is problematic as acquired immune responses can be both adaptive and maladaptive in the physiological sense. Indeed, both acquired and innate immune responses can be both adaptive and maladaptive in the evolutionary sense. Most textbooks today, following the early use by 140: 1101:. Those domains can be alternatively spliced reaching high numbers of variations. It was shown that after exposure to different pathogens there are different splice forms of dscam produced. After the animals with different splice forms are exposed to the same pathogen only the individuals with the splice form specific for that pathogen survive. 1365:, or antigenic determinants. Most antigens contain a variety of epitopes and can stimulate the production of antibodies, specific T cell responses, or both. A very small proportion (less than 0.01%) of the total lymphocytes are able to bind to a particular antigen, which suggests that only a few cells respond to each antigen. 1369:

different pathogen that might ever be encountered. Even in the absence of antigen stimulation, a human can produce more than 1 trillion different antibody molecules. Millions of genes would be required to store the genetic information that produces these receptors, but, the entire human genome contains fewer than 25,000

1408:, which the immune system does not routinely patrol. The second is that the fetus itself may promote local immunosuppression in the mother, perhaps by a process of active nutrient depletion. A more modern explanation for this induction of tolerance is that specific glycoproteins expressed in the uterus during 1391:

The innate and acquired portions of the immune system work together, not in spite of each other. The acquired arm, B, and T cells could not function without the innate system input. T cells are useless without antigen-presenting cells to activate them, and B cells are disabled without T cell help. On

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Like the T cell, B cells express a unique B cell receptor (BCR), in this case, a membrane-bound antibody molecule. All the BCR of any one clone of B cells recognizes and binds to only one particular antigen. A critical difference between B cells and T cells is how each cell "sees" an antigen. T cells

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discovery, "antigen-specific immunity mediated by somatic gene rearrangements that create clone-defining antigen receptors". In the last decade, the term "adaptive" has been increasingly applied to another class of immune response not so-far associated with somatic gene rearrangements. These include

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Immunity can be acquired either actively or passively. Immunity is acquired actively when a person is exposed to foreign substances and the immune system responds. Passive immunity is when antibodies are transferred from one host to another. Both actively acquired and passively acquired immunity can

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For the acquired response to "remember" and eliminate a large number of pathogens the immune system must be able to distinguish between many different antigens, and the receptors that recognize antigens must be produced in a huge variety of configurations, in essence one receptor (at least) for each

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CD4+ lymphocytes, also called "helper" T cells, are immune response mediators, and play an important role in establishing and maximizing the capabilities of the acquired immune response. These cells have no cytotoxic or phagocytic activity; and cannot kill infected cells or clear pathogens, but, in

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Several T cells subgroups can be activated by professional APCs, and each type of T cell is specially equipped to deal with each unique toxin or microbial pathogen. The type of T cell activated, and the type of response generated, depends, in part, on the context in which the APC first encountered

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in reference to antibody responses in frogs as a synonym for "acquired immune response" in 1964. Good acknowledged he used the terms as synonyms but explained only that he preferred to use the term "adaptive". He might have been thinking of the then not implausible theory of antibody formation in

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cannot do sufficiently, as certain blood cells specialize to insert themselves between adjacent epithelial cells). The immunodepressive action was the initial normal behavior of the virus, similar to HIV. The fusion proteins were a way to spread the infection to other cells by simply merging them

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Classically, two types of effector CD4 T helper cell responses can be induced by a professional APC, designated Th1 and Th2, each designed to eliminate different types of pathogens. The factors that dictate whether an infection triggers a Th1 or Th2 type response are not fully understood, but the

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in bacteria, that is, enzymes whose expression could be induced by their substrates. The phrase was used almost exclusively by Good and his students and a few other immunologists working with marginal organisms until the 1990s when it became widely used in tandem with the term "innate immunity"

1380:. According to the clonal selection theory, at birth, an animal randomly generates a vast diversity of lymphocytes (each bearing a unique antigen receptor) from information encoded in a small family of genes. To generate each unique antigen receptor, these genes have undergone a process called 735:

Helper T cells express T cell receptors (TCR) that recognize antigen bound to Class II MHC molecules. The activation of a naive helper T-cell causes it to release cytokines, which influences the activity of many cell types, including the APC (Antigen-Presenting Cell) that activated it. Helper

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Naive cytotoxic T cells are activated when their T-cell receptor (TCR) strongly interacts with a peptide-bound MHC class I molecule. This affinity depends on the type and orientation of the antigen/MHC complex, and is what keeps the CTL and infected cell bound together. Once activated, the CTL

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are a critical part of the adaptive immune system. Adaptive immunity can provide long-lasting protection, sometimes for the person's entire lifetime. For example, someone who recovers from measles is now protected against measles for their lifetime; in other cases it does not provide lifetime

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receptors, Ig and TCR, are found in all jawed vertebrates. The most ancient Ig class, IgM, is membrane-bound and then secreted upon stimulation of cartilaginous fish B cells. Another isotype, shark IgW, is related to mammalian IgD. TCRs, both α/β and γ/δ, are found in all animals from

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and discards the rest, which produces a highly unique combination of antigen-receptor gene segments in each lymphocyte. This mechanism allows a small number of genetic segments to generate a vast number of different antigen receptors, which are then uniquely expressed on each individual

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antigens are produced by intracellular bacteria and viruses replicating within a host cell. The host cell uses enzymes to digest virally associated proteins and displays these pieces on its surface to T-cells by coupling them to MHC. Endogenous antigens are typically displayed on

528:. These antigens are different from those on the surface of bacteria or on the surface of virus-infected host cells ("non-self" or "foreign" antigens). The acquired immune response is triggered by recognizing foreign antigen in the cellular context of an activated dendritic cell. 1288:) is the deliberate induction of an immune response, and represents the single most effective manipulation of the immune system that scientists have developed. Immunizations are successful because they utilize the immune system's natural specificity as well as its inducibility. 1071:

they are able to develop a memory of that infection that allows them to withstand otherwise lethal dose of the same bacteria they were exposed to before. Unlike in vertebrates, insects do not possess cells specific for adaptive immunity. Instead those mechanisms are mediated by

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A population study has shown that the protective effect of the sickle-cell trait against falciparum malaria involves the augmentation of acquired as well as innate immune responses to the malaria parasite, illustrating the expected transition from innate to acquired immunity.

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infection. HIV is able to subvert the immune system by specifically attacking the CD4 T cells, precisely the cells that could drive the clearance of the virus, but also the cells that drive immunity against all other pathogens encountered during an organism's lifetime.

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The principle behind immunization is to introduce an antigen, derived from a disease-causing organism, that stimulates the immune system to develop protective immunity against that organism, but that does not itself cause the pathogenic effects of that organism. An

1529:. By school age most children have developed efficacious adaptive immunity against malaria. These observations raise questions about mechanisms that favor the survival of most children in Africa while allowing some to develop potentially lethal infections. 963:

recognize their cognate antigen in a processed form – as a peptide in the context of an MHC molecule, whereas B cells recognize antigens in their native form. Once a B cell encounters its cognate (or specific) antigen (and receives additional signals from a

1627:. Though the molecules of the AIS are well-conserved, they are also rapidly evolving. Yet, a comparative approach finds that many features are quite uniform across taxa. All the major features of the AIS arose early and quickly. Jawless fishes have a 575:

signals, to the T cell-enriched lymph nodes. During migration, dendritic cells undergo a process of maturation in which they lose most of their ability to engulf other pathogens, and develop an ability to communicate with T-cells. The dendritic cell uses

701:(cell death). To limit extensive tissue damage during an infection, CTL activation is tightly controlled and in general requires a very strong MHC/antigen activation signal, or additional activation signals provided by "helper" T-cells (see below). 708:

clear them away—but a few of these cells remain as memory cells. On a later encounter with the same antigen, these memory cells quickly differentiate into effector cells, dramatically shortening the time required to mount an effective response.

1431:(ERVs) are activated and produced in high quantities during the implantation of the embryo. They are currently known to possess immunosuppressive properties, suggesting a role in protecting the embryo from its mother's immune system. Also, 653:

Cytotoxic T cells (also known as TC, killer T cell, or cytotoxic T-lymphocyte (CTL)) are a sub-group of T cells that induce the death of cells that are infected with viruses (and other pathogens), or are otherwise damaged or dysfunctional.

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The acquired immune system, which has been best-studied in mammals, originated in jawed fish approximately 500 million years ago. Most of the molecules, cells, tissues, and associated mechanisms of this system of defense are found in

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Rogozin IB, Iyer LM, Liang L, Glazko GV, Liston VG, Pavlov YI, Aravind L, Pancer Z (June 2007). "Evolution and diversification of lamprey antigen receptors: evidence for involvement of an AID-APOBEC family cytosine deaminase".

1076:. Hemocytes function similarly to phagocytes and after priming they are able to more effectively recognize and engulf the pathogen. It was also shown that it is possible to transfer the memory into offspring. For example, in 2970:

Mi S, Lee X, Li X, Veldman GM, Finnerty H, Racie L, LaVallie E, Tang XY, Edouard P, Howes S, Keith JC, McCoy JM (February 2000). "Syncytin is a captive retroviral envelope protein involved in human placental morphogenesis".

939:, differing in biological properties; each has evolved to handle different kinds of antigens. Upon activation, B cells produce antibodies, each of which recognize a unique antigen, and neutralizing specific pathogens. 1665:– involves a natural transfer of antibodies from a mother to her infant. The antibodies cross the woman's placenta to the fetus. Antibodies can also be transferred through breast milk with the secretions of colostrum. 865:, γδ T cells exhibit characteristics that place them at the border between innate and acquired immunity. On one hand, γδ T cells may be considered a component of adaptive immunity in that they rearrange TCR genes via 1444:

with the infected one (HIV does this too). It is believed that the ancestors of modern viviparous mammals evolved after an infection by this virus, enabling the fetus to survive the immune system of the mother.

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One of the most interesting developments in biomedical science during the past few decades has been elucidation of mechanisms mediating innate immunity. One set of innate immune mechanisms is humoral, such as

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because the fetus does not actually make any memory cells or antibodies: It only borrows them. Short-term passive immunity can also be transferred artificially from one individual to another via antibody-rich

857:(TCR) as opposed to CD4+ and CD8+ αβ T cells and share characteristics of helper T cells, cytotoxic T cells and natural killer cells. Like other 'unconventional' T cell subsets bearing invariant TCRs, such as 1532:

In malaria, as in other infections, innate immune responses lead into, and stimulate, adaptive immune responses. The genetic control of innate and acquired immunity is now a large and flourishing discipline.

662:, in which it gains functions and divides rapidly to produce an army of "armed" effector cells. Activated CTL then travels throughout the body searching for cells that bear that unique MHC Class I + peptide. 251:, which are proteins also known as immunoglobulins. Antibodies travel through the bloodstream and bind to the foreign antigen causing it to inactivate, which does not allow the antigen to bind to the host. 535:), all cells are capable of presenting antigen through the function of major histocompatibility complex (MHC) molecules. Some cells are specially equipped to present antigen, and to prime naive T cells. 384:

Acquired immunity is triggered in vertebrates when a pathogen evades the innate immune system and (1) generates a threshold level of antigen and (2) generates "stranger" or "danger" signals activating

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differs in cartilaginous fishes, which have a cluster form as compared to the translocon form in bony fish to mammals. Like TCR and Ig, the MHC is found only in jawed vertebrates. Genes involved in

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Hibino T, Loza-Coll M, Messier C, Majeske AJ, Cohen AH, Terwilliger DP, Buckley KM, Brockton V, Nair SV, Berney K, Fugmann SD, Anderson MK, Pancer Z, Cameron RA, Smith LC, Rast JP (December 2006).

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Rubio M, Maestro JL, Piulachs MD, Belles X (June 2018). "Conserved association of Argonaute 1 and 2 proteins with miRNA and siRNA pathways throughout insect evolution, from cockroaches to flies".

873:, and develop a memory phenotype. On the other hand, however, the various subsets may also be considered part of the innate immune system where a restricted TCR or NK receptors may be used as a 919:. Antibodies (also known as immunoglobulin, Ig), are large Y-shaped proteins used by the immune system to identify and neutralize foreign objects. In mammals, there are five types of antibody: 727:

The T lymphocyte activation pathway. T cells contribute to immune defenses in two major ways: some direct and regulate immune responses; others directly attack infected or cancerous cells.

985:. Already primed to produce specific antibodies, these cells can be called upon to respond quickly if the same pathogen re-infects the host, while the host experiences few, if any, symptoms. 2028:. National Institute of Allergy and Infectious Diseases National Cancer Institute, U.S. Department of Health and Human Services National Institutes of Health. September 2003. Archived from 1268:

In general, active immunity is long-term and can be acquired by infection followed by B cell and T cell activation, or artificially acquired by vaccines, in a process called immunization.

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Humoral and cell-mediated immune responses limit malaria parasite multiplication, and many cytokines contribute to the pathogenesis of malaria as well as to the resolution of infections.

459:. In humans, approximately 1–2% of the lymphocyte pool recirculates each hour to increase the opportunity for the cells to encounter the specific pathogen and antigen that they react to. 1481:

of the variable portion of an antibody) and 'anti-idiotypes' (antigen receptors that react with the idiotype as if it were a foreign antigen). This theory, which builds on the existing

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are any substances that elicit the adaptive immune response. Sometimes the adaptive system is unable to distinguish harmful from harmless foreign molecules; the effects of this may be

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Historically, infectious disease has been the leading cause of death in the human population. Over the last century, two important factors have been developed to combat their spread:

1108:(RNAi). RNAi is a form of antiviral immunity with high specificity. It has several different pathways that all end with the virus being unable to replicate. One of the pathways is 247:, two different types of lymphocytes, carry out the main activities: antibody responses, and cell-mediated immune response. In antibody responses, B cells are activated to secrete 1016:, have an adaptive immune system that shows 3 different cell lineages, each sharing a common origin with B cells, αβ T cells, and innate-like γΔ T cells. Instead of the classical 439:

are the cells of the adaptive immune system. The human body has about 2 trillion lymphocytes, which are 20–40% of white blood cells; their total mass is about the same as the

212:, which is pre-programmed to react to common broad categories of pathogen, the adaptive immune system is highly specific to each particular pathogen the body has encountered. 3570: 1248:

contains antibodies (mainly IgA) that are transferred to the gut of the infant, protecting against bacterial infections, until the newborn can synthesize its own antibodies.

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The B lymphocyte activation pathway. B cells function to protect the host by producing antibodies that identify and neutralize foreign objects like bacteria and viruses.

4314: 1677:– This involves the introduction of antibodies rather than antigens to the human body. These antibodies are from an animal or person who is already immune to the disease. 1112:

in which long double stranded RNA is cut into pieces that serve as templates for protein complex Ago2-RISC that finds and degrades complementary RNA of the virus.

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if the queen is infected with bacteria then the newly born workers have enhanced abilities in fighting with the same bacteria. Other experimental model based on

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Gesner E.M; Schellenberg M.J; Garside E.L; George M.M. & Macmillan A.M. 2011. Recognition and maturation of effector RNAs in a CRISPR interference pathway.

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CRISPR has a lot of short repeated sequences. These sequences are part of an adaptive immune system for prokaryotes. It allows them to remember and counter the

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are another distinct population of effector CD4 T cells that develop from naive T cells post-antigen activation. Tfh cells are specialized in helping B cell

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Acquired immunity relies on the capacity of immune cells to distinguish between the body's own cells and unwanted invaders. The host's cells express "self"

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components. Many antigens derived from acellular vaccines do not strongly induce an adaptive response, and most bacterial vaccines require the addition of

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T-cells require a much milder activation stimulus than cytotoxic T cells. Helper T cells can provide extra signals that "help" activate cytotoxic cells.

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which antibodies were plastic and could adapt themselves to the molecular shape of antigens, and/or to the concept of "adaptive enzymes" as described by

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are short-lived cells (2–3 days) that secrete antibodies. These antibodies bind to antigens, making them easier targets for phagocytes, and trigger the

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and other mobile elements. Despite the research the exact mechanisms responsible for immune priming and specificity in insects are not well described.

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the immune system to control aberrant immune responses to self-antigens; an important mechanism in controlling the development of autoimmune diseases.

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have had no prior exposure to microbes and are particularly vulnerable to infection. Several layers of passive protection are provided by the mother.

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Grissa I; Vergnaud G. & Pourcel C. 2007. The CRISPRdb database and tools to display CRISPRs and to generate dictionaries of spacers and repeats.

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as an abnormal expansion of Tfh cell numbers can lead to unrestricted autoreactive antibody production causing severe systemic autoimmune disorders.

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Increasingly, there is strong evidence from mouse and human-based scientific studies of a broader diversity in CD4 effector T helper cell subsets.

3563: 278:). This acquired response is called "adaptive" because it prepares the body's immune system for future challenges (though it can actually also be 60: 1616:

The evolution of the AIS, based on Ig, TCR, and MHC molecules, is thought to have arisen from two major evolutionary events: the transfer of the

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An antibody is made up of two heavy chains and two light chains. The unique variable region allows an antibody to recognize its matching antigen.

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Antibody-dependent cell-mediated cytotoxicity: Antibodies attached to target cell cause destruction by macrophages, eosinophils, and NK cells

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that relies on gene rearrangement to generate diverse immune receptors with a functional dichotomy that parallels Ig and TCR molecules. The

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the other hand, the innate system would likely be overrun with pathogens without the specialized action of the adaptive immune response.

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in secondary lymphoid organs and provide them positive paracrine signals to enable the generation and recall production of high-quality

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The cornerstone of the immune system is the recognition of "self" versus "non-self". Therefore, the mechanisms that protect the human

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Immune memory in insects was discovered through the phenomenon of priming. When insects are exposed to non-lethal dose or heat killed

592:(HLA)). This MHC-antigen complex is recognized by T-cells passing through the lymph node. Exogenous antigens are usually displayed on 274:

are "acquired" during the lifetime of the organism (whereas in innate immunity pathogen-specific receptors are already encoded in the

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Mojica F.J.M. et al 2000. Biological significance of regularly spaced repeats in the genomes of Archaea, Bacteria and mitochondria.

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of "acquired immunity " vs. "adaptive immunity". The peak for "adaptive" in the 1960s reflects its introduction to immunology by

126: 31: 584:. In the lymph node, the dendritic cell displays these non-self antigens on its surface by coupling them to a receptor called the 489:

In an adult animal, the peripheral lymphoid organs contain a mixture of B and T cells in at least three stages of differentiation:

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are equipped with special "co-stimulatory" ligands recognized by co-stimulatory receptors on T cells, and are termed professional

3902: 1826: 797:. Like cytotoxic T cells, most of the CD4 helper cells die on resolution of infection, with a few remaining as CD4 memory cells. 585: 482:, B cells and T cells are produced by stem cells in the bone marrow. T cell progenitors then migrate from the bone marrow to the 1904: 1525:

Repeated malaria infections strengthen acquired immunity and broaden its effects against parasites expressing different surface

1244:, so that, at birth, human babies have high levels of antibodies, with the same range of antigen specificities as their mother. 4132: 2403:"Insect immunity: oral exposure to a bacterial pathogen elicits free radical response and protects from a recurring infection" 1949: 1361:, can serve as antigens. The parts of an antigen that interact with an antibody molecule or a lymphocyte receptor, are called 1056:. However, in recent years some of the basic hallmarks of adaptive immunity have been discovered in insects. Those traits are 1028:(VLRs for short) that, like the antigen receptors of jawed vertebrates, are produced from only a small number (one or two) of 500:, which have left the bone marrow or thymus and entered the lymphatic system, but have yet to encounter their matching antigen 86: 4267: 3897: 786: 219:

after an initial response to a specific pathogen, and leads to an enhanced response to future encounters with that pathogen.

64: 3049: 4004: 3840: 1617: 1610: 471: 4335: 4197: 1586: 1025: 874: 447:. The peripheral bloodstream contains only 2% of all circulating lymphocytes; the other 98% move within tissues and the 144: 93: 196:

and use by colleagues; the explosive increase in the 1990s was correlated with the use of the phrase "innate immunity".

2714:"RNA interference-based antiviral immune response against the salivary gland hypertrophy virus in Glossina pallidipes" 1494: 809: 185: 5036: 3726: 1116:

pathway in cytoplasm binds to Ago1-RISC complex and functions as a template for viral RNA degradation. Last one is

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Williams TN, Mwangi TW, Roberts DJ, Alexander ND, Weatherall DJ, Wambua S, Kortok M, Snow RW, Marsh K (May 2005).

2663:"AgDscam, a hypervariable immunoglobulin domain-containing receptor of the Anopheles gambiae innate immune system" 53: 4996: 4078: 1304:

erator), is defined as any substance that binds to a specific antibody and elicits an adaptive immune response.

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Generation of responses that are tailored to maximally eliminate specific pathogens or pathogen-infected cells.

3163:"Band 3/complement-mediated recognition and removal of normally senescent and pathological human erythrocytes" 2823: 881:

produced by microbes, and highly restricted intraepithelial Vδ1 T cells respond to stressed epithelial cells.

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Wiedenheft B; Sternberg S.H; Doudna J.A. 2012. RNA-guided genetic silencing systems in bacteria and archaea.

2097:"Single-cell transcriptomics of human T cells reveals tissue and activation signatures in health and disease" 4064: 4046: 4029: 3906: 3664: 1439:

to limit exchange of migratory cells between the developing embryo and the body of the mother (something an

1428: 589: 467: 3399: 5127: 4930: 4835: 4687: 3928: 3646: 2855: 1758: 1478: 762: 356:, use "adaptive" almost exclusively and noting in glossaries that the term is synonymous with "acquired". 205: 4737: 3875: 3731: 3709: 2712:

Meki IK, Kariithi HM, Parker AG, Vreysen MJ, Ros VI, Vlak JM, van Oers MM, Abd-Alla AM (November 2018).

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Antigen presentation stimulates T cells to become either "cytotoxic" CD8+ cells or "helper" CD4+ cells.

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that have been activated by their matching antigen, and are actively involved in eliminating a pathogen

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in the antibody-coding genes, which allows antibodies with novel specificity to be created. Second,

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also showed pathogen specific primed memory transfer into offspring from both mothers and fathers.

866: 821: 365: 302: 3024:"The Viruses That Make Us: A Role For Endogenous Retrovirus In The Evolution Of Placental Species" 4907: 4778: 4540: 4531: 4517: 3772: 3535: 3492: 3448: 3284: 3004: 2798: 2491: 1602: 1563: 1515: 1312: 1197: 978: 850: 845: 817: 801: 171: 1132: 322:(offspring) of that cell inherit genes that encode the same receptor specificity, including the 1060:

and specificity. Although the hallmarks are present the mechanisms are different from those in

804:, have been identified as important negative regulators of adaptive immunity as they limit and 5106: 5096: 4808: 4786: 4627: 4424: 4182: 4115: 3820: 3586: 3527: 3484: 3440: 3380: 3336: 3276: 3235: 3184: 3143: 3099: 2996: 2952: 2790: 2745: 2694: 2640: 2611:"Paternally derived immune priming for offspring in the red flour beetle, Tribolium castaneum" 2591: 2542: 2483: 2434: 2383: 2334: 2326: 2285: 2236: 2187: 2136: 2077: 1993: 1910: 1832: 1753: 1733: 1511: 256: 1469:

A theoretical framework explaining the workings of the acquired immune system is provided by

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Passive memory is usually short-term, lasting between a few days and several months. Newborn

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In humans, it takes 4–7 days for the adaptive immune system to mount a significant response.

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Recognition of specific "non-self" antigens in the presence of "self", during the process of

5073: 4580: 4563: 4444: 4394: 4389: 4272: 4215: 4192: 4152: 4021: 3940: 3880: 3852: 3847: 3815: 3802: 3792: 3519: 3476: 3430: 3422: 3370: 3326: 3318: 3307:"Origin and evolution of the adaptive immune system: genetic events and selective pressures" 3266: 3225: 3215: 3174: 3133: 3089: 2988: 2942: 2780: 2772: 2735: 2725: 2684: 2674: 2630: 2581: 2573: 2532: 2522: 2473: 2424: 2414: 2373: 2316: 2275: 2267: 2226: 2218: 2177: 2167: 2126: 2116: 2067: 2059: 1748: 1482: 1377: 1105: 1081: 932: 916: 894: 825: 813: 805: 749: 659: 648: 448: 201: 1097:

gene also known as Down syndrome cell adhesive molecule is a gene that contains 3 variable

376: 5081: 5029: 4981: 4912: 4902: 4499: 4327: 4277: 4237: 4177: 4172: 4137: 3835: 3119:"Excess heme in sickle erythrocyte inside-out membranes: possible role in thiol oxidation" 2891: 2859: 1743: 1601:

use variable lymphocyte receptors (VLRs) for antigen binding. Diversity is generated by a

1021: 854: 694: 678: 536: 385: 360: 353: 271: 1635:, which has an important role in AIS activation, is the most important defense system of 3418: 2984: 2938: 2626: 2469: 2369: 2154:

Magen A, Nie J, Ciucci T, Tamoutounour S, Zhao Y, Mehta M, et al. (December 2019).

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response generated does play an important role in the clearance of different pathogens.

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Szabo PA, Levitin HM, Miron M, Snyder ME, Senda T, Yuan J, et al. (October 2019).

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essence "manage" the immune response, by directing other cells to perform these tasks.

339: 327: 193: 3400:"Somatic diversification of variable lymphocyte receptors in the agnathan sea lamprey" 2609:

Roth O, Joop G, Eggert H, Hilbert J, Daniel J, Schmid-Hempel P, Kurtz J (March 2010).

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pathogens, such as bacteria, parasites or toxins in the tissues and then migrate, via

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randomly selects one variable (V), one diversity (D), and one joining (J) region for

279: 235:

The cells that carry out the adaptive immune response are white blood cells known as

163: 3539: 3288: 2495: 1709:– Antibodies pass from mother to fetus via placenta or infant via the mother's milk. 559: 5086: 4409: 4357: 4187: 3797: 3782: 3777: 3714: 3612: 3496: 3008: 2802: 1953: 1671:– is done by vaccination (introducing dead or weakened antigen to the host's cell). 1636: 1575: 1555: 1281: 1257: 1162: 1049: 974: 912: 753: 666: 593: 497: 493: 283: 189: 3452: 3398:

Pancer Z, Amemiya CT, Ehrhardt GR, Ceitlin J, Gartland GL, Cooper MD (July 2004).

2454:"Insect immunity shows specificity in protection upon secondary pathogen exposure" 348:

which became a popular subject after the discovery of the Toll receptor system in

3989: 3220: 2776: 2679: 2527: 2172: 17: 5014: 4863: 4843: 4658: 4653: 4593: 4242: 4038: 3375: 3138: 3057: 1930:

Charles A Janeway, Jr; Travers, Paul; Walport, Mark; Shlomchik, Mark J. (2001).

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which prey on them. They work as a kind of acquired immune system for bacteria.

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Alder MN, Rogozin IB, Iyer LM, Glazko GV, Cooper MD, Pancer Z (December 2005).

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Antigen and antibody binding would cause five different protective mechanisms:

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Hernández López J, Schuehly W, Crailsheim K, Riessberger-Gallé U (June 2014).

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Bacteria use CRISPR as part of their adaptive immune system to defend against

1061: 908: 674: 612: 475: 430: 311: 290: 248: 236: 225: 175: 148: 2354:"Diversity and function of adaptive immune receptors in a jawless vertebrate" 2330: 1803:. Institute for Quality and Efficiency in Health Care (IQWiG). 4 August 2016. 1659:– when a person is naturally exposed to antigens, becomes ill, then recovers. 166:

that is composed of specialized, systemic cells and processes that eliminate

4885: 4680: 4461: 3978: 3683: 2880: 2511:"A specific primed immune response in Drosophila is dependent on phagocytes" 2419: 2378: 2353: 1474: 1436: 1424: 1409: 1316: 1077: 705: 698: 568: 319: 139: 4292: 3531: 3488: 3444: 3384: 3340: 3280: 3239: 3188: 3103: 3094: 3077: 3000: 2956: 2794: 2749: 2698: 2644: 2595: 2577: 2546: 2487: 2438: 2387: 2338: 2289: 2240: 2191: 2140: 2081: 170:

or prevent their growth. The acquired immune system is one of the two main

3147: 981:. About 10% of plasma cells survive to become long-lived antigen-specific 760:(marking for phagocytosis) and complement-fixing antibodies, and leads to 624:+ cytotoxic T-cells. With the exception of non-nucleated cells (including 30:

This article is about human immunity. For the same concept in botany, see

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in a similar way to antibodies, and with the same degree of specificity.

1017: 690: 670: 525: 298: 252: 220: 167: 3426: 2947: 2922: 2785: 2305:"Evolution of innate and adaptive immune systems in jawless vertebrates" 1578:

genes, are closely linked within the MHC of almost all studied species.

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and water to flow into the infected cell, and causing it to burst or

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Boehm T (May 2011). "Design principles of adaptive immune systems".

3356:"The immune gene repertoire encoded in the purple sea urchin genome" 3322: 3271: 3254: 1825:

Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walters P (2002).

1715:– Preformed antibodies in immune serum are introduced by injection. 1477:(unique molecular features of one clonotype, i.e. the unique set of 3480: 1936:

Immunobiology: The Immune System in Health and Disease. 5th Edition

5056: 5051: 4936: 4663: 4570: 4471: 4429: 4419: 4368: 3026:. University of California, Irvine (lecture notes). Archived from 1640: 1401: 1344: 1320: 1131: 1104:

Other mechanisms supporting the specificity of insect immunity is

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Agglutination: Reduces number of infectious units to be dealt with

898: 794: 722: 686: 558: 444: 440: 375: 184: 138: 3204:"An immune basis for malaria protection by the sickle cell trait" 2921:

International Human Genome Sequencing Consortium (October 2004).

2207:"T cells that promote B-Cell maturation in systemic autoimmunity" 1952:. University of South Carolina School of Medicine. Archived from 1181:. Its structure and function was discovered in the 21st century. 958:

Neutralization: Blocks adhesion of bacteria and viruses to mucosa

952:

Opsonization: Coating antigen with antibody enhances phagocytosis

380:

Overview of the processes involved in the primary immune response

200:

Like the innate system, the adaptive immune system includes both

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Biochimica et Biophysica Acta (BBA) - Gene Regulatory Mechanisms

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encapsulated in a granule that enters cells via pores to induce

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Pham LN, Dionne MS, Shirasu-Hiza M, Schneider DS (March 2007).

2401:

Mikonranta L, Mappes J, Kaukoniitty M, Freitak D (March 2014).

1514:. Another set comprises pattern recognition receptors such as 4858: 4675: 4439: 3050:"Persisting Viruses Could Play Role in Driving Host Evolution" 1154: 1120:

where small RNA binds to the Piwi protein family and controls

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Most commonly accepted theory of the specificity is based on

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The relevance of CD4 T helper cells is highlighted during an

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pathogens (viruses and bacteria that are inside host cells).

359:

The classic sense of "acquired immunity" came to mean, since

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to mammals. The organization of gene segments that undergo

824:

antibodies. Similar to Tregs, Tfh cells also play a role in

704:

On resolution of the infection, most effector cells die and

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Activation of complement: Cause inflammation and cell lysis

391:

The major functions of the acquired immune system include:

1024:, these animals possess a large array of molecules called 1376:

Myriad receptors are produced through a process known as

1159:

clustered regularly-interspaced short palindromic repeats

2923:"Finishing the euchromatic sequence of the human genome" 1581:

Lymphoid cells can be identified in some pre-vertebrate

1493:, is seen as being relevant to the understanding of the 907:

B Cells are the major cells involved in the creation of

1988:

Janeway CA, Travers P, Walport M, Shlomchik MJ (2005).

1903:

Janeway CA, Travers P, Walport M, Shlomchik MJ (2001).

1485:

hypothesis and since 1974 has been developed mainly by

789:. In general, Th2 responses are more effective against 766:. In general, Th1 responses are more effective against 756:

activities of macrophages, and induces B cells to make

748:

The Th1 response is characterized by the production of

208:

components and destroys invading pathogens. Unlike the

1909:(5th ed.). New York and London: Garland Science. 1831:(4th ed.). New York and London: Garland Science. 1315:

viruses, whereas many bacterial vaccines are based on

2048:"Eosinophils and Th2 immunity: contemporary insights" 531:

With the exception of non-nucleated cells (including

773:

The Th2 response is characterized by the release of

228:. This process of adaptive immunity is the basis of 5072: 4995: 4957: 4834: 4777: 4736: 4701: 4642: 4579: 4539: 4530: 4498: 4470: 4343: 4334: 4258: 4206: 4163: 4086: 4077: 4055: 4037: 4028: 3961: 3919: 3861: 3762: 3692: 3600: 3593: 1202:When B cells and T cells are activated some become 330:that are the keys to long-lived specific immunity. 67:. Unsourced material may be challenged and removed. 1609:-based DNA segments. There is no evidence for the 1329:that activate the antigen-presenting cells of the 1032:. These molecules are believed to bind pathogenic 677:: cytotoxins that form pores in the target cell's 162:, or specific immune system is a subsystem of the 3255:"Immunological processes in malaria pathogenesis" 1319:components of microorganisms, including harmless 580:to chop the pathogen into smaller pieces, called 2562:"Trans-generational immune priming in honeybees" 2205:Weinstein JS, Hernandez SG, Craft J (May 2012). 781:in the clearance of parasites. Th2 also produce 665:When exposed to these infected or dysfunctional 3300: 3298: 1177:have it. It is their defence against attack by 971:into an effector cell, known as a plasma cell. 628:), MHC class I is expressed by all host cells. 2019:"Understanding the Immune System How It Works" 1950:"Microbiology and Immunology On-Line Textbook" 1353:Most large molecules, including virtually all 509:Memory cells, the survivors of past infections 462:B cells and T cells are derived from the same 409:, in which pathogens are "remembered" through 4308: 4005: 3564: 2661:Dong Y, Taylor HE, Dimopoulos G (July 2006). 1585:(i.e., sea urchins). These bind antigen with 816:as they are uniquely capable of migrating to 470:, whereas T cells are intimately involved in 8: 3117:Kuross SA, Rank BH, Hebbel RP (April 1988). 3078:"Toll-like receptors and Type I interferons" 1932:"Principles of innate and adaptive immunity" 1652:be obtained by natural or artificial means. 793:bacteria, parasites including helminths and 1983: 297:is a process of accelerated random genetic 4536: 4340: 4315: 4301: 4293: 4083: 4034: 4012: 3998: 3990: 3597: 3571: 3557: 3549: 3071: 3069: 3067: 2256:"A cold-blooded view of adaptive immunity" 1981: 1979: 1977: 1975: 1973: 1971: 1969: 1967: 1965: 1963: 1898: 1896: 1894: 1892: 1890: 1888: 1886: 1884: 1882: 1880: 1878: 1876: 1874: 1872: 1870: 1868: 1412:suppress the uterine immune response (see 3434: 3374: 3330: 3270: 3229: 3219: 3178: 3137: 3093: 2946: 2835:Nature Structural & Molecular Biology 2784: 2739: 2729: 2688: 2678: 2634: 2585: 2536: 2526: 2477: 2428: 2418: 2377: 2320: 2279: 2230: 2181: 2171: 2130: 2120: 2071: 1866: 1864: 1862: 1860: 1858: 1856: 1854: 1852: 1850: 1848: 1820: 1818: 1816: 1814: 1812: 1810: 406: 127:Learn how and when to remove this message 3161:Arese P, Turrini F, Schwarzer E (2005). 1680: 1284:. Immunization (commonly referred to as 1000:Adaptive immunity in jawless vertebrates 270:In adaptive immunity, pathogen-specific 3305:Flajnik MF, Kasahara M (January 2010). 3253:Schofield L, Grau GE (September 2005). 2013: 2011: 2009: 1790: 1701:– Antigens are introduced in vaccines. 1240:IgG is transported directly across the 2452:Sadd BM, Schmid-Hempel P (June 2006). 1800:The innate and adaptive immune systems 1675:Artificially Acquired Passive Immunity 967:(predominately Th2 type)), it further 338:The term "adaptive" was first used by 2656: 2654: 1669:Artificially Acquired Active Immunity 1435:cause the formation of the placental 885:B lymphocytes and antibody production 7: 3167:Cellular Physiology and Biochemistry 2254:Flajnik, Martin F. (19 March 2018). 1044:For a long time it was thought that 853:(γδ T cells) possess an alternative 740:Th1 and Th2: helper T cell responses 464:multipotent hematopoietic stem cells 65:adding citations to reliable sources 3082:The Journal of Biological Chemistry 1695:– Antigen enters the body naturally 1663:Naturally Acquired Passive Immunity 1623:(possibly of viral origin) and two 643:CD8+ T lymphocytes and cytotoxicity 1657:Naturally Acquired Active Immunity 1396:Acquired immunity during pregnancy 588:, or MHC (also known in humans as 25: 1992:(6th ed.). Garland Science. 32:Plant-induced systemic resistance 2636:10.1111/j.1365-2656.2009.01617.x 2566:Proceedings. Biological Sciences 2322:10.1111/j.1348-0421.2012.00500.x 2223:10.1111/j.1600-065x.2012.01122.x 1505:Stimulation of adaptive immunity 1473:, based on interactions between 586:major histocompatibility complex 282:when it results in allergies or 41: 4133:Periarteriolar lymphoid sheaths 3076:Uematsu S, Akira S (May 2007). 2303:Kasamatsu, Jun (January 2013). 810:Follicular helper T (Tfh) cells 293:because of two factors. First, 52:needs additional citations for 3898:Immunoglobulin class switching 2046:Spencer LA, Weller PF (2010). 1611:recombination-activating genes 486:, where they develop further. 472:cell-mediated immune responses 27:Subsystem of the immune system 1: 2615:The Journal of Animal Ecology 1828:Molecular Biology of the Cell 1587:pattern recognition receptors 1455:classified into 24 families. 1386:combinatorial diversification 1026:variable lymphocyte receptors 3221:10.1371/journal.pmed.0020128 2777:10.1016/j.bbagrm.2018.04.001 2680:10.1371/journal.pbio.0040229 2528:10.1371/journal.ppat.0030026 2173:10.1016/j.celrep.2019.10.131 1423:mammals (all mammals except 875:pattern recognition receptor 145:scanning electron microscope 3376:10.1016/j.ydbio.2006.08.065 3139:10.1182/blood.V71.4.876.876 2309:Microbiology and Immunology 2052:Immunology and Cell Biology 2026:NIH Publication No. 03-5423 1605:-mediated rearrangement of 785:, which facilitates B cell 658:undergoes a process called 5144: 3727:Polyclonal B cell response 3512:Nature Reviews. Immunology 3259:Nature Reviews. Immunology 3048:Villarreal LP (Oct 2001). 2122:10.1038/s41467-019-12464-3 1647:Types of acquired immunity 1462: 1195: 1143: 997: 888: 843: 716: 646: 635: 596:molecules, which activate 517: 428: 215:Adaptive immunity creates 29: 4198:Waldeyer's tonsillar ring 2731:10.1186/s12866-018-1298-1 2479:10.1016/j.cub.2006.04.047 2272:10.1038/s41577-018-0003-9 2260:Nature Reviews Immunology 1625:whole genome duplications 802:Regulatory T (Treg) cells 4226:High endothelial venules 3311:Nature Reviews. Genetics 2852:Nobel Prize in Chemistry 1161:. These are part of the 1157:research. It stands for 620:molecules, and activate 545:antigen-presenting cells 147:image of a single human 76:"Adaptive immune system" 4047:Hematopoietic stem cell 2420:10.1186/1742-9994-11-23 2379:10.1126/science.1119420 1451:found several thousand 1429:endogenous retroviruses 1341:Immunological diversity 826:immunological tolerance 669:, effector CTL release 590:human leukocyte antigen 567:Dendritic cells engulf 468:humoral immune response 4767:Lymphoid immune system 4336:Musculoskeletal system 3841:Tolerance in pregnancy 3583:adaptive immune system 3095:10.1074/jbc.R700009200 2906:Molecular Microbiology 2856:Emmanuelle Charpentier 2578:10.1098/rspb.2014.0454 1759:Original antigenic sin 1687:Artificially acquired 1497:and the search for an 1479:antigenic determinants 1350: 1141: 994:In jawless vertebrates 904: 869:, which also produces 863:natural killer T cells 763:cell-mediated immunity 752:, which activates the 728: 564: 381: 206:cell-mediated immunity 197: 160:acquired immune system 156:adaptive immune system 151: 4755:Myeloid immune system 4541:Cardiovascular system 3876:Somatic hypermutation 3710:Polyclonal antibodies 3705:Monoclonal antibodies 3363:Developmental Biology 2211:Immunological Reviews 2101:Nature Communications 1764:Somatic hypermutation 1512:complement activation 1471:immune network theory 1465:Immune network theory 1459:Immune network theory 1433:viral fusion proteins 1348: 1214:short-term memory or 1196:Further information: 1135: 902: 726: 562: 451:, which includes the 379: 307:genetic recombination 295:somatic hypermutation 289:The system is highly 188: 178:(the other being the 142: 5107:Islets of Langerhans 4967:Genitourinary system 4703:Integumentary system 4065:Hassall's corpuscles 3893:Junctional diversity 3661:Antigen presentation 2407:Frontiers in Zoology 2166:(10): 3019–3032.e6. 2064:10.1038/icb.2009.115 1956:on 2 September 2008. 1776:Notes and references 1633:innate immune system 1591:innate immune system 1547:cartilaginous fishes 1491:Geoffrey W. Hoffmann 1449:human genome project 1419:During pregnancy in 1331:innate immune system 1192:Immunological memory 1054:innate immune system 915:and lymph, known as 871:junctional diversity 520:Antigen presentation 514:Antigen presentation 407:immunological memory 397:antigen presentation 224:protection, as with 217:immunological memory 210:innate immune system 180:innate immune system 174:strategies found in 158:, also known as the 61:improve this article 4997:Reproductive system 4716:Subcutaneous tissue 4485:Cartilaginous joint 4148:Trabecular arteries 3888:V(D)J recombination 3871:Affinity maturation 3623:Antigenic variation 3427:10.1038/nature02740 3419:2004Natur.430..174P 2985:2000Natur.403..785M 2948:10.1038/nature03001 2939:2004Natur.431..931H 2627:2010JAnEc..79..403R 2470:2006CBio...16.1206S 2370:2005Sci...310.1970A 2113:2019NatCo..10.4706S 1769:Polyclonal response 1729:Affinity maturation 1516:toll-like receptors 1382:V(D)J recombination 1006:jawless vertebrates 989:Alternative systems 867:V(D)J recombination 851:Gamma delta T cells 840:Gamma delta T cells 608:Endogenous antigens 303:V(D)J recombination 4779:Respiratory system 4532:Circulatory system 2817:BMC Bioinformatics 2572:(1785): 20140454. 1684:Naturally acquired 1603:cytosine deaminase 1564:antigen processing 1560:gene rearrangement 1351: 1311:are based on live 1218:long-term memory. 1198:Immunity (medical) 1142: 979:complement cascade 911:that circulate in 905: 846:Gamma delta T cell 818:follicular B cells 729: 565: 555:Exogenous antigens 382: 318:of each cell, all 198: 152: 5115: 5114: 4638: 4637: 4628:Glymphatic system 4526: 4525: 4290: 4289: 4286: 4285: 4116:Cords of Billroth 4073: 4072: 3987: 3986: 3915: 3914: 3665:professional APCs 3469:Nature Immunology 3180:10.1159/000089839 2364:(5756): 1970–73. 1754:Immunosuppression 1734:Allelic exclusion 1719: 1718: 1597:, two subsets of 1570:, as well as the 787:isotype switching 137: 136: 129: 111: 18:Adaptive immunity 16:(Redirected from 5135: 5074:Endocrine system 4836:Digestive system 4581:Lymphatic system 4564:Lymphatic vessel 4537: 4341: 4317: 4310: 4303: 4294: 4216:Lymphatic vessel 4193:Tonsillar crypts 4153:Trabecular veins 4084: 4079:Secondary organs 4035: 4022:lymphatic system 4014: 4007: 4000: 3991: 3881:Clonal selection 3853:Immune privilege 3848:Immunodeficiency 3803:Cross-reactivity 3793:Hypersensitivity 3598: 3573: 3566: 3559: 3550: 3544: 3543: 3507: 3501: 3500: 3463: 3457: 3456: 3438: 3413:(6996): 174–80. 3404: 3395: 3389: 3388: 3378: 3360: 3351: 3345: 3344: 3334: 3302: 3293: 3292: 3274: 3250: 3244: 3243: 3233: 3223: 3199: 3193: 3192: 3182: 3158: 3152: 3151: 3141: 3123: 3114: 3108: 3107: 3097: 3088:(21): 15319–23. 3073: 3062: 3061: 3056:. Archived from 3045: 3039: 3038: 3036: 3035: 3019: 3013: 3012: 2993:10.1038/35001608 2979:(6771): 785–89. 2967: 2961: 2960: 2950: 2933:(7011): 931–45. 2918: 2912: 2902: 2896: 2888: 2882: 2879:(7385): 331–338. 2869: 2863: 2848: 2842: 2831: 2825: 2813: 2807: 2806: 2788: 2760: 2754: 2753: 2743: 2733: 2724:(Suppl 1): 170. 2718:BMC Microbiology 2709: 2703: 2702: 2692: 2682: 2658: 2649: 2648: 2638: 2606: 2600: 2599: 2589: 2557: 2551: 2550: 2540: 2530: 2506: 2500: 2499: 2481: 2449: 2443: 2442: 2432: 2422: 2398: 2392: 2391: 2381: 2349: 2343: 2342: 2324: 2300: 2294: 2293: 2283: 2251: 2245: 2244: 2234: 2202: 2196: 2195: 2185: 2175: 2151: 2145: 2144: 2134: 2124: 2092: 2086: 2085: 2075: 2043: 2037: 2036: 2034: 2023: 2015: 2004: 2003: 1985: 1958: 1957: 1946: 1940: 1939: 1927: 1921: 1920: 1900: 1843: 1842: 1822: 1805: 1804: 1795: 1749:Immune tolerance 1681: 1495:HIV pathogenesis 1483:clonal selection 1378:clonal selection 1253:passive immunity 1106:RNA interference 1082:red flour beetle 1022:T cell receptors 917:humoral immunity 895:Humoral immunity 879:common molecules 822:affinity-matured 814:humoral immunity 777:, which induces 750:Interferon-gamma 660:clonal selection 649:Cytotoxic T cell 449:lymphatic system 202:humoral immunity 132: 125: 121: 118: 112: 110: 69: 45: 37: 21: 5143: 5142: 5138: 5137: 5136: 5134: 5133: 5132: 5118: 5117: 5116: 5111: 5068: 5030:Seminal vesicle 4991: 4953: 4903:Small intestine 4830: 4773: 4732: 4697: 4634: 4575: 4522: 4500:Muscular system 4494: 4466: 4345:Skeletal system 4330: 4321: 4291: 4282: 4278:Germinal center 4254: 4238:Germinal center 4202: 4159: 4138:Germinal center 4069: 4051: 4024: 4018: 3988: 3983: 3957: 3911: 3857: 3836:Clonal deletion 3764: 3758: 3688: 3589: 3577: 3547: 3524:10.1038/nri2944 3509: 3508: 3504: 3465: 3464: 3460: 3402: 3397: 3396: 3392: 3358: 3353: 3352: 3348: 3323:10.1038/nrg2703 3304: 3303: 3296: 3272:10.1038/nri1686 3252: 3251: 3247: 3201: 3200: 3196: 3173:(4–6): 133–46. 3160: 3159: 3155: 3121: 3116: 3115: 3111: 3075: 3074: 3065: 3047: 3046: 3042: 3033: 3031: 3022:Villarreal LP. 3021: 3020: 3016: 2969: 2968: 2964: 2920: 2919: 2915: 2903: 2899: 2892:Jennifer Doudna 2889: 2885: 2870: 2866: 2860:Jennifer Doudna 2849: 2845: 2832: 2828: 2814: 2810: 2762: 2761: 2757: 2711: 2710: 2706: 2660: 2659: 2652: 2608: 2607: 2603: 2559: 2558: 2554: 2508: 2507: 2503: 2464:(12): 1206–10. 2458:Current Biology 2451: 2450: 2446: 2400: 2399: 2395: 2351: 2350: 2346: 2302: 2301: 2297: 2253: 2252: 2248: 2204: 2203: 2199: 2153: 2152: 2148: 2094: 2093: 2089: 2045: 2044: 2040: 2032: 2021: 2017: 2016: 2007: 2000: 1987: 1986: 1961: 1948: 1947: 1943: 1929: 1928: 1924: 1917: 1902: 1901: 1846: 1839: 1824: 1823: 1808: 1797: 1796: 1792: 1778: 1773: 1744:Immune response 1724: 1649: 1542: 1507: 1467: 1461: 1398: 1359:polysaccharides 1343: 1274: 1266: 1224: 1200: 1194: 1148: 1130: 1042: 1002: 996: 991: 897: 889:Main articles: 887: 855:T cell receptor 848: 842: 742: 721: 715: 695:serine protease 679:plasma membrane 651: 645: 640: 634: 610: 557: 539:, B-cells, and 537:Dendritic cells 522: 516: 433: 427: 405:Development of 386:dendritic cells 374: 336: 263:, or any other 204:components and 133: 122: 116: 113: 70: 68: 58: 46: 35: 28: 23: 22: 15: 12: 11: 5: 5141: 5139: 5131: 5130: 5120: 5119: 5113: 5112: 5110: 5109: 5104: 5099: 5094: 5089: 5084: 5078: 5076: 5070: 5069: 5067: 5066: 5065: 5064: 5059: 5054: 5049: 5044: 5034: 5033: 5032: 5027: 5022: 5017: 5012: 5001: 4999: 4993: 4992: 4990: 4989: 4984: 4979: 4974: 4969: 4963: 4961: 4959:Urinary system 4955: 4954: 4952: 4951: 4950: 4949: 4944: 4939: 4927: 4926: 4925: 4920: 4915: 4910: 4905: 4895: 4894: 4893: 4888: 4883: 4881:Laryngopharynx 4878: 4868: 4867: 4866: 4861: 4856: 4851: 4849:Salivary gland 4840: 4838: 4832: 4831: 4829: 4828: 4827: 4826: 4821: 4816: 4806: 4805: 4804: 4799: 4794: 4783: 4781: 4775: 4774: 4772: 4771: 4770: 4769: 4759: 4758: 4757: 4746: 4744: 4742:immune systems 4738:Haematopoietic 4734: 4733: 4731: 4730: 4729: 4728: 4718: 4713: 4707: 4705: 4699: 4698: 4696: 4695: 4690: 4688:Somatic system 4685: 4684: 4683: 4678: 4671:Sensory system 4668: 4667: 4666: 4661: 4656: 4648: 4646: 4644:Nervous system 4640: 4639: 4636: 4635: 4633: 4632: 4631: 4630: 4623:CNS equivalent 4620: 4619: 4618: 4613: 4603: 4602: 4601: 4596: 4585: 4583: 4577: 4576: 4574: 4573: 4568: 4567: 4566: 4561: 4556: 4545: 4543: 4534: 4528: 4527: 4524: 4523: 4521: 4520: 4515: 4510: 4504: 4502: 4496: 4495: 4493: 4492: 4490:Synovial joint 4487: 4482: 4476: 4474: 4468: 4467: 4465: 4464: 4459: 4458: 4457: 4452: 4447: 4442: 4437: 4432: 4427: 4422: 4417: 4412: 4407: 4402: 4397: 4392: 4387: 4382: 4377: 4372: 4366: 4360: 4349: 4347: 4338: 4332: 4331: 4322: 4320: 4319: 4312: 4305: 4297: 4288: 4287: 4284: 4283: 4281: 4280: 4275: 4270: 4264: 4262: 4256: 4255: 4253: 4252: 4251: 4250: 4245: 4240: 4230: 4229: 4228: 4218: 4212: 4210: 4204: 4203: 4201: 4200: 4195: 4190: 4185: 4180: 4175: 4169: 4167: 4161: 4160: 4158: 4157: 4156: 4155: 4150: 4144:Blood vessels 4142: 4141: 4140: 4135: 4125: 4124: 4123: 4118: 4108: 4107: 4106: 4101: 4092: 4090: 4081: 4075: 4074: 4071: 4070: 4068: 4067: 4061: 4059: 4053: 4052: 4050: 4049: 4043: 4041: 4032: 4030:Primary organs 4026: 4025: 4020:Organs of the 4019: 4017: 4016: 4009: 4002: 3994: 3985: 3984: 3982: 3981: 3976: 3971: 3965: 3963: 3959: 3958: 3956: 3955: 3950: 3949: 3948: 3938: 3937: 3936: 3925: 3923: 3917: 3916: 3913: 3912: 3910: 3909: 3900: 3895: 3890: 3885: 3884: 3883: 3878: 3867: 3865: 3863:Immunogenetics 3859: 3858: 3856: 3855: 3850: 3845: 3844: 3843: 3838: 3833: 3828: 3823: 3811: 3810: 3808:Co-stimulation 3805: 3800: 3795: 3790: 3785: 3780: 3775: 3768: 3766: 3760: 3759: 3757: 3756: 3751: 3749:Immune complex 3745: 3744: 3739: 3734: 3729: 3724: 3723: 3722: 3717: 3712: 3707: 3696: 3694: 3690: 3689: 3687: 3686: 3681: 3676: 3671: 3669:Dendritic cell 3657: 3656: 3651: 3650: 3649: 3647:Conformational 3644: 3633: 3632: 3627: 3626: 3625: 3620: 3615: 3604: 3602: 3595: 3591: 3590: 3578: 3576: 3575: 3568: 3561: 3553: 3546: 3545: 3502: 3481:10.1038/ni1463 3458: 3390: 3346: 3294: 3245: 3194: 3153: 3109: 3063: 3060:on 2009-05-08. 3040: 3014: 2962: 2913: 2897: 2883: 2864: 2843: 2826: 2808: 2755: 2704: 2650: 2601: 2552: 2515:PLOS Pathogens 2501: 2444: 2393: 2344: 2295: 2246: 2197: 2146: 2087: 2038: 2035:on 2007-01-03. 2005: 1998: 1959: 1941: 1922: 1915: 1844: 1837: 1806: 1789: 1788: 1787: 1783: 1782: 1777: 1774: 1772: 1771: 1766: 1761: 1756: 1751: 1746: 1741: 1736: 1731: 1725: 1723: 1720: 1717: 1716: 1710: 1703: 1702: 1696: 1689: 1688: 1685: 1679: 1678: 1672: 1666: 1660: 1648: 1645: 1595:jawless fishes 1589:(PRRs) of the 1541: 1538: 1506: 1503: 1463:Main article: 1460: 1457: 1397: 1394: 1342: 1339: 1335:immunogenicity 1273: 1270: 1265: 1262: 1223: 1222:Passive memory 1220: 1208:memory T cells 1204:memory B cells 1193: 1190: 1186:bacteriophages 1144:Main article: 1138:bacteriophages 1129: 1126: 1041: 1038: 1008:, such as the 998:Main article: 995: 992: 990: 987: 983:memory B cells 969:differentiates 960: 959: 956: 953: 950: 947: 886: 883: 844:Main article: 841: 838: 741: 738: 717:Main article: 714: 713:Helper T-cells 711: 689:. CTL release 647:Main article: 644: 641: 636:Main article: 633: 630: 609: 606: 602:T helper cells 556: 553: 518:Main article: 515: 512: 511: 510: 507: 504:Effector cells 501: 429:Main article: 426: 423: 419: 418: 415:memory T cells 411:memory B cells 403: 400: 373: 370: 366:natural killer 335: 332: 328:memory T cells 324:memory B cells 194:Robert A. Good 135: 134: 49: 47: 40: 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 5140: 5129: 5128:Immune system 5126: 5125: 5123: 5108: 5105: 5103: 5100: 5098: 5095: 5093: 5090: 5088: 5085: 5083: 5080: 5079: 5077: 5075: 5071: 5063: 5060: 5058: 5055: 5053: 5050: 5048: 5045: 5043: 5040: 5039: 5038: 5035: 5031: 5028: 5026: 5023: 5021: 5018: 5016: 5013: 5011: 5008: 5007: 5006: 5003: 5002: 5000: 4998: 4994: 4988: 4985: 4983: 4980: 4978: 4975: 4973: 4970: 4968: 4965: 4964: 4962: 4960: 4956: 4948: 4945: 4943: 4942:Biliary tract 4940: 4938: 4935: 4934: 4933: 4932: 4928: 4924: 4921: 4919: 4916: 4914: 4911: 4909: 4906: 4904: 4901: 4900: 4899: 4896: 4892: 4889: 4887: 4884: 4882: 4879: 4877: 4874: 4873: 4872: 4869: 4865: 4862: 4860: 4857: 4855: 4852: 4850: 4847: 4846: 4845: 4842: 4841: 4839: 4837: 4833: 4825: 4822: 4820: 4817: 4815: 4812: 4811: 4810: 4807: 4803: 4800: 4798: 4795: 4793: 4790: 4789: 4788: 4785: 4784: 4782: 4780: 4776: 4768: 4765: 4764: 4763: 4760: 4756: 4753: 4752: 4751: 4748: 4747: 4745: 4743: 4739: 4735: 4727: 4726:Mammary gland 4724: 4723: 4722: 4719: 4717: 4714: 4712: 4709: 4708: 4706: 4704: 4700: 4694: 4691: 4689: 4686: 4682: 4679: 4677: 4674: 4673: 4672: 4669: 4665: 4662: 4660: 4657: 4655: 4652: 4651: 4650: 4649: 4647: 4645: 4641: 4629: 4626: 4625: 4624: 4621: 4617: 4614: 4612: 4609: 4608: 4607: 4604: 4600: 4597: 4595: 4592: 4591: 4590: 4587: 4586: 4584: 4582: 4578: 4572: 4569: 4565: 4562: 4560: 4557: 4555: 4552: 4551: 4550: 4547: 4546: 4544: 4542: 4538: 4535: 4533: 4529: 4519: 4516: 4514: 4511: 4509: 4506: 4505: 4503: 4501: 4497: 4491: 4488: 4486: 4483: 4481: 4480:Fibrous joint 4478: 4477: 4475: 4473: 4469: 4463: 4460: 4456: 4453: 4451: 4448: 4446: 4443: 4441: 4438: 4436: 4433: 4431: 4428: 4426: 4423: 4421: 4418: 4416: 4413: 4411: 4408: 4406: 4403: 4401: 4398: 4396: 4393: 4391: 4388: 4386: 4383: 4381: 4378: 4376: 4373: 4370: 4367: 4364: 4361: 4359: 4356: 4355: 4354: 4351: 4350: 4348: 4346: 4342: 4339: 4337: 4333: 4329: 4325: 4324:Human systems 4318: 4313: 4311: 4306: 4304: 4299: 4298: 4295: 4279: 4276: 4274: 4273:Peyer's patch 4271: 4269: 4266: 4265: 4263: 4261: 4257: 4249: 4248:Marginal zone 4246: 4244: 4241: 4239: 4236: 4235: 4234: 4231: 4227: 4224: 4223: 4222: 4219: 4217: 4214: 4213: 4211: 4209: 4205: 4199: 4196: 4194: 4191: 4189: 4186: 4184: 4181: 4179: 4176: 4174: 4171: 4170: 4168: 4166: 4162: 4154: 4151: 4149: 4146: 4145: 4143: 4139: 4136: 4134: 4131: 4130: 4129: 4126: 4122: 4121:Marginal zone 4119: 4117: 4114: 4113: 4112: 4109: 4105: 4102: 4100: 4097: 4096: 4094: 4093: 4091: 4089: 4085: 4082: 4080: 4076: 4066: 4063: 4062: 4060: 4058: 4054: 4048: 4045: 4044: 4042: 4040: 4036: 4033: 4031: 4027: 4023: 4015: 4010: 4008: 4003: 4001: 3996: 3995: 3992: 3980: 3977: 3975: 3972: 3970: 3967: 3966: 3964: 3960: 3954: 3951: 3947: 3944: 3943: 3942: 3939: 3935: 3932: 3931: 3930: 3927: 3926: 3924: 3922: 3918: 3908: 3904: 3901: 3899: 3896: 3894: 3891: 3889: 3886: 3882: 3879: 3877: 3874: 3873: 3872: 3869: 3868: 3866: 3864: 3860: 3854: 3851: 3849: 3846: 3842: 3839: 3837: 3834: 3832: 3831:Clonal anergy 3829: 3827: 3824: 3822: 3819: 3818: 3817: 3813: 3812: 3809: 3806: 3804: 3801: 3799: 3796: 3794: 3791: 3789: 3786: 3784: 3781: 3779: 3776: 3774: 3770: 3769: 3767: 3761: 3755: 3752: 3750: 3747: 3746: 3743: 3740: 3738: 3735: 3733: 3730: 3728: 3725: 3721: 3720:Microantibody 3718: 3716: 3713: 3711: 3708: 3706: 3703: 3702: 3701: 3698: 3697: 3695: 3691: 3685: 3682: 3680: 3677: 3675: 3672: 3670: 3666: 3662: 3659: 3658: 3655: 3652: 3648: 3645: 3643: 3640: 3639: 3638: 3635: 3634: 3631: 3628: 3624: 3621: 3619: 3616: 3614: 3611: 3610: 3609: 3606: 3605: 3603: 3599: 3596: 3592: 3588: 3584: 3581: 3574: 3569: 3567: 3562: 3560: 3555: 3554: 3551: 3541: 3537: 3533: 3529: 3525: 3521: 3518:(5): 307–17. 3517: 3513: 3506: 3503: 3498: 3494: 3490: 3486: 3482: 3478: 3475:(6): 647–56. 3474: 3470: 3462: 3459: 3454: 3450: 3446: 3442: 3437: 3436:2027.42/62870 3432: 3428: 3424: 3420: 3416: 3412: 3408: 3401: 3394: 3391: 3386: 3382: 3377: 3372: 3369:(1): 349–65. 3368: 3364: 3357: 3350: 3347: 3342: 3338: 3333: 3328: 3324: 3320: 3316: 3312: 3308: 3301: 3299: 3295: 3290: 3286: 3282: 3278: 3273: 3268: 3265:(9): 722–35. 3264: 3260: 3256: 3249: 3246: 3241: 3237: 3232: 3227: 3222: 3217: 3213: 3209: 3208:PLOS Medicine 3205: 3198: 3195: 3190: 3186: 3181: 3176: 3172: 3168: 3164: 3157: 3154: 3149: 3145: 3140: 3135: 3132:(4): 876–82. 3131: 3127: 3120: 3113: 3110: 3105: 3101: 3096: 3091: 3087: 3083: 3079: 3072: 3070: 3068: 3064: 3059: 3055: 3051: 3044: 3041: 3030:on 2007-07-15 3029: 3025: 3018: 3015: 3010: 3006: 3002: 2998: 2994: 2990: 2986: 2982: 2978: 2974: 2966: 2963: 2958: 2954: 2949: 2944: 2940: 2936: 2932: 2928: 2924: 2917: 2914: 2910: 2907: 2901: 2898: 2895: 2893: 2887: 2884: 2881: 2878: 2874: 2868: 2865: 2861: 2857: 2853: 2847: 2844: 2841:(6): 688–692. 2840: 2836: 2830: 2827: 2824: 2821: 2818: 2812: 2809: 2804: 2800: 2796: 2792: 2787: 2782: 2778: 2774: 2771:(6): 554–60. 2770: 2766: 2759: 2756: 2751: 2747: 2742: 2737: 2732: 2727: 2723: 2719: 2715: 2708: 2705: 2700: 2696: 2691: 2686: 2681: 2676: 2672: 2668: 2664: 2657: 2655: 2651: 2646: 2642: 2637: 2632: 2628: 2624: 2621:(2): 403–13. 2620: 2616: 2612: 2605: 2602: 2597: 2593: 2588: 2583: 2579: 2575: 2571: 2567: 2563: 2556: 2553: 2548: 2544: 2539: 2534: 2529: 2524: 2520: 2516: 2512: 2505: 2502: 2497: 2493: 2489: 2485: 2480: 2475: 2471: 2467: 2463: 2459: 2455: 2448: 2445: 2440: 2436: 2431: 2426: 2421: 2416: 2412: 2408: 2404: 2397: 2394: 2389: 2385: 2380: 2375: 2371: 2367: 2363: 2359: 2355: 2348: 2345: 2340: 2336: 2332: 2328: 2323: 2318: 2314: 2310: 2306: 2299: 2296: 2291: 2287: 2282: 2277: 2273: 2269: 2266:(7): 438–53. 2265: 2261: 2257: 2250: 2247: 2242: 2238: 2233: 2228: 2224: 2220: 2217:(1): 160–71. 2216: 2212: 2208: 2201: 2198: 2193: 2189: 2184: 2179: 2174: 2169: 2165: 2161: 2157: 2150: 2147: 2142: 2138: 2133: 2128: 2123: 2118: 2114: 2110: 2106: 2102: 2098: 2091: 2088: 2083: 2079: 2074: 2069: 2065: 2061: 2058:(3): 250–56. 2057: 2053: 2049: 2042: 2039: 2031: 2027: 2020: 2014: 2012: 2010: 2006: 2001: 1999:0-443-07310-4 1995: 1991: 1990:Immunobiology 1984: 1982: 1980: 1978: 1976: 1974: 1972: 1970: 1968: 1966: 1964: 1960: 1955: 1951: 1945: 1942: 1937: 1933: 1926: 1923: 1918: 1916:0-8153-4101-6 1912: 1908: 1907: 1906:Immunobiology 1899: 1897: 1895: 1893: 1891: 1889: 1887: 1885: 1883: 1881: 1879: 1877: 1875: 1873: 1871: 1869: 1867: 1865: 1863: 1861: 1859: 1857: 1855: 1853: 1851: 1849: 1845: 1840: 1838:0-8153-3218-1 1834: 1830: 1829: 1821: 1819: 1817: 1815: 1813: 1811: 1807: 1802: 1801: 1794: 1791: 1785: 1784: 1780: 1779: 1775: 1770: 1767: 1765: 1762: 1760: 1757: 1755: 1752: 1750: 1747: 1745: 1742: 1740: 1737: 1735: 1732: 1730: 1727: 1726: 1721: 1714: 1711: 1708: 1705: 1704: 1700: 1697: 1694: 1691: 1690: 1686: 1683: 1682: 1676: 1673: 1670: 1667: 1664: 1661: 1658: 1655: 1654: 1653: 1646: 1644: 1642: 1638: 1637:invertebrates 1634: 1630: 1629:different AIS 1626: 1622: 1619: 1614: 1612: 1608: 1604: 1600: 1596: 1592: 1588: 1584: 1583:deuterostomes 1579: 1577: 1573: 1569: 1565: 1561: 1557: 1552: 1548: 1539: 1537: 1534: 1530: 1528: 1523: 1519: 1517: 1513: 1504: 1502: 1500: 1496: 1492: 1488: 1484: 1480: 1476: 1472: 1466: 1458: 1456: 1454: 1450: 1445: 1442: 1438: 1434: 1430: 1426: 1422: 1417: 1415: 1411: 1407: 1403: 1395: 1393: 1389: 1387: 1383: 1379: 1374: 1372: 1366: 1364: 1360: 1356: 1347: 1340: 1338: 1336: 1332: 1328: 1327: 1322: 1318: 1314: 1310: 1305: 1303: 1299: 1295: 1289: 1287: 1283: 1279: 1271: 1269: 1264:Active memory 1263: 1261: 1259: 1254: 1249: 1247: 1243: 1239: 1235: 1234: 1229: 1221: 1219: 1217: 1213: 1209: 1205: 1199: 1191: 1189: 1187: 1182: 1180: 1176: 1172: 1168: 1164: 1160: 1156: 1153:is a term in 1152: 1147: 1139: 1134: 1127: 1125: 1123: 1119: 1115: 1111: 1107: 1102: 1100: 1096: 1092: 1091: 1085: 1083: 1079: 1075: 1070: 1065: 1063: 1059: 1058:immune memory 1055: 1052:possess only 1051: 1050:invertebrates 1047: 1039: 1037: 1035: 1031: 1027: 1023: 1019: 1015: 1011: 1007: 1001: 993: 988: 986: 984: 980: 976: 972: 970: 966: 965:helper T cell 957: 954: 951: 948: 945: 944: 943: 940: 938: 934: 930: 926: 922: 918: 914: 910: 901: 896: 892: 884: 882: 880: 876: 872: 868: 864: 860: 856: 852: 847: 839: 837: 834: 829: 827: 823: 819: 815: 811: 807: 803: 798: 796: 792: 791:extracellular 788: 784: 783:Interleukin 4 780: 776: 775:Interleukin 5 771: 769: 768:intracellular 765: 764: 759: 755: 751: 746: 739: 737: 733: 725: 720: 719:T helper cell 712: 710: 707: 702: 700: 696: 692: 688: 684: 680: 676: 672: 668: 667:somatic cells 663: 661: 655: 650: 642: 639: 632:T lymphocytes 631: 629: 627: 623: 619: 614: 607: 605: 603: 599: 595: 591: 587: 583: 579: 574: 570: 561: 554: 552: 551:the antigen. 548: 546: 542: 538: 534: 529: 527: 521: 513: 508: 505: 502: 499: 498:naive T cells 495: 492: 491: 490: 487: 485: 481: 477: 473: 469: 465: 460: 458: 454: 450: 446: 442: 438: 432: 424: 422: 416: 412: 408: 404: 401: 398: 394: 393: 392: 389: 387: 378: 371: 369: 367: 364:expansion of 362: 357: 355: 351: 346: 341: 333: 331: 329: 325: 321: 317: 313: 308: 304: 300: 296: 292: 287: 285: 281: 277: 273: 268: 266: 262: 258: 254: 250: 246: 242: 238: 233: 231: 227: 222: 218: 213: 211: 207: 203: 195: 191: 187: 183: 181: 177: 173: 169: 165: 164:immune system 161: 157: 150: 146: 141: 131: 128: 120: 109: 106: 102: 99: 95: 92: 88: 85: 81: 78: – 77: 73: 72:Find sources: 66: 62: 56: 55: 50:This article 48: 44: 39: 38: 33: 19: 4929: 4897: 4870: 4622: 4605: 4588: 4548: 3798:Inflammation 3783:Alloimmunity 3778:Autoimmunity 3763:Immunity vs. 3715:Autoantibody 3613:Superantigen 3582: 3515: 3511: 3505: 3472: 3468: 3461: 3410: 3406: 3393: 3366: 3362: 3349: 3317:(1): 47–59. 3314: 3310: 3262: 3258: 3248: 3211: 3207: 3197: 3170: 3166: 3156: 3129: 3125: 3112: 3085: 3081: 3058:the original 3053: 3043: 3032:. 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