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1254:, and it becomes more difficult for it to activate in future. This mechanism prevents inappropriate responses to self, as self-peptides will not usually be presented with suitable co-stimulation. Once a T cell has been appropriately activated (i.e. has received signal one and signal two) it alters its cell surface expression of a variety of proteins. Markers of T cell activation include CD69, CD71 and CD25 (also a marker for Treg cells), and HLA-DR (a marker of human T cell activation). CTLA-4 expression is also up-regulated on activated T cells, which in turn outcompetes CD28 for binding to the B7 proteins. This is a checkpoint mechanism to prevent over activation of the T cell. Activated T cells also change their cell surface glycosylation profile. 502:β-selection is the first checkpoint, where thymocytes that are able to form a functional pre-TCR (with an invariant alpha chain and a functional beta chain) are allowed to continue development in the thymus. Next, positive selection checks that thymocytes have successfully rearranged their TCRα locus and are capable of recognizing MHC molecules with appropriate affinity. Negative selection in the medulla then eliminates thymocytes that bind too strongly to self-antigens expressed on MHC molecules. These selection processes allow for tolerance of self by the immune system. Typical naive T cells that leave the thymus (via the corticomedullary junction) are self-restricted, self-tolerant, and single positive. 29: 757: 1431: 1372:(Caspase activation and recruitment domains) domains; that then binds TRAF6, which is ubiquitinated at K63. This form of ubiquitination does not lead to degradation of target proteins. Rather, it serves to recruit NEMO, IKKα and -β, and TAB1-2/ TAK1. TAK 1 phosphorylates IKK-β, which then phosphorylates IκB allowing for K48 ubiquitination: leads to proteasomal degradation. Rel A and p50 can then enter the nucleus and bind the NF-κB response element. This coupled with NFAT signaling allows for complete activation of the IL-2 gene. 540: 890:) differ from the other memory subsets in that they do not originate following a strong clonal expansion event. Thus, although this population as a whole is abundant within the peripheral circulation, individual virtual memory T cell clones reside at relatively low frequencies. One theory is that homeostatic proliferation gives rise to this T cell population. Although CD8 virtual memory T cells were the first to be described, it is now known that CD4 virtual memory cells also exist. 1708:
studies have suggested that it is possible to predict relapse of leukemia based on expression of inhibitory receptors PD-1 and TIM-3 by T cells. Many experiments and clinical trials have focused on immune checkpoint blockers in cancer therapy, with some of these approved as valid therapies that are now in clinical use. Inhibitory receptors targeted by those medical procedures are vital in T cell exhaustion and blocking them can reverse these changes.
1155: 392:α-chain, signals are produced which cease rearrangement of the β-chain (and silence the alternate allele). Although these signals require the pre-TCR at the cell surface, they are independent of ligand binding to the pre-TCR. If the chains successfully pair a pre-TCR forms, and the cell downregulates CD25 and is termed a DN4 cell (CD25CD44). These cells then undergo a round of proliferation, and begin to re-arrange the TCRα locus during the 41: 1583: 988:
recognition of peptide antigens in the context of the MHC molecule. Overall, there are three large populations of unconventional T cells: NKT cells, MAIT cells, and gammadelta T cells. Now, their functional roles are already being well established in the context of infections and cancer. Furthermore, these T cell subsets are being translated into many therapies against malignancies such as leukemia, for example.
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must be tested to make sure they work at all. First, the thymocytes attempt to create a functional beta chain, testing it against a 'mock' alpha chain. Then they attempt to create a functional alpha chain. Once a working TCR has been produced, the cells then must test if their TCR will identify threats correctly, and to do this it is required to recognize the body’s
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thought to belong to either the effector or central memory subtypes, each with their own distinguishing set of cell surface markers (see below). Subsequently, numerous new populations of memory T cells were discovered including tissue-resident memory T (Trm) cells, stem memory TSCM cells, and virtual memory T cells. The single unifying theme for all
1222:, like dendritic cells, B cells, and macrophages, to name a few. The peptides presented to CD8 T cells by MHC class I molecules are 8–13 amino acids in length; the peptides presented to CD4 cells by MHC class II molecules are longer, usually 12–25 amino acids in length, as the ends of the binding cleft of the MHC class II molecule are open. 1108:. In rabbits, sheep, and chickens, the number of γδ T cells can be as high as 60% of total T cells. The antigenic molecules that activate γδ T cells are still mostly unknown. However, γδ T cells are not MHC-restricted and seem to be able to recognize whole proteins rather than requiring peptides to be presented by MHC molecules on 1679:
encounter anti-inflammatory cytokines and pro-apoptotic proteins take over to protect the body from damage. Sepsis also carries high antigen load and inflammation. In this stage of sepsis T cell exhaustion increases. Currently there are studies aiming to utilize inhibitory receptor blockades in treatment of sepsis.
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role in tolerance of a graft mainly by depletion of alloreactive CD8 T cells. Several studies showed positive effect of chronic infection on graft acceptance and its long-term survival mediated partly by T cell exhaustion. It was also shown that recipient T cell exhaustion provides sufficient conditions for
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While during infection T cell exhaustion can develop following persistent antigen exposure after graft transplant similar situation arises with alloantigen presence. It was shown that T cell response diminishes over time after kidney transplant. These data suggest T cell exhaustion plays an important
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About 98% of thymocytes die during the development processes in the thymus by failing either positive selection or negative selection, whereas the other 2% survive and leave the thymus to become mature immunocompetent T cells. The thymus contributes fewer cells as a person ages. As the thymus shrinks
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cells can be a source of IL-10 and TGF-β and therefore they can play a role in T cell exhaustion. Furthermore, T cell exhaustion is reverted after depletion of Treg cells and blockade of PD1. T cell exhaustion can also occur during sepsis as a result of cytokine storm. Later after the initial septic
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T cell exhaustion can be triggered by several factors like persistent antigen exposure and lack of CD4 T cell help. Antigen exposure also has effect on the course of exhaustion because longer exposure time and higher viral load increases the severity of T cell exhaustion. At least 2–4 weeks exposure
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into the cytosol. Low calcium in the endoplasmic reticulum causes STIM1 clustering on the ER membrane and leads to activation of cell membrane CRAC channels that allows additional calcium to flow into the cytosol from the extracellular space. This aggregated cytosolic calcium binds calmodulin, which
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During cancer T cell exhaustion plays a role in tumor protection. According to research some cancer-associated cells as well as tumor cells themselves can actively induce T cell exhaustion at the site of tumor. T cell exhaustion can also play a role in cancer relapses as was shown on leukemia. Some
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after they encounter their cognate antigen within the context of an MHC molecule on the surface of a professional antigen presenting cell (e.g. a dendritic cell). Appropriate co-stimulation must be present at the time of antigen encounter for this process to occur. Historically, memory T cells were
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T cells are grouped into a series of subsets based on their function. CD4 and CD8 T cells are selected in the thymus, but undergo further differentiation in the periphery to specialized cells which have different functions. T cell subsets were initially defined by function, but also have associated
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on their cell surface and although the T cell antigen receptor can interact with at least a subset of these self pMHC, the T cell generally ignores these healthy cells. However, when these very same cells contain even minute quantities of pathogen derived pMHC, T cells are able to become activated
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Negative selection removes thymocytes that are capable of strongly binding with "self" MHC molecules. Thymocytes that survive positive selection migrate towards the boundary of the cortex and medulla in the thymus. While in the medulla, they are again presented with a self-antigen presented on the
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and constant region genes in an attempt to create a functional TCRβ chain. As the developing thymocyte progresses through to the DN3 stage (CD44CD25), the thymocyte expresses an invariant α-chain called pre-Tα alongside the TCRβ gene. If the rearranged β-chain successfully pairs with the invariant
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A thymocyte can only become an active T cell when it survives the process of developing a functional TCR. The TCR consists of two major components, the alpha and beta chains. These both contain random elements designed to produce a wide variety of different TCRs, but due to this huge variety they
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production. Optimal CD8 T cell response relies on CD4 signalling. CD4 cells are useful in the initial antigenic activation of naive CD8 T cells, and sustaining memory CD8 T cells in the aftermath of an acute infection. Therefore, activation of CD4 T cells can be beneficial to the action of CD8 T
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subtypes is that they are long-lived and can quickly expand to large numbers of effector T cells upon re-exposure to their cognate antigen. By this mechanism they provide the immune system with "memory" against previously encountered pathogens. Memory T cells may be either CD4 or CD8 and usually
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T cell exhaustion is a poorly defined or ambiguous term. There are three approaches to its definition. "The first approach primarily defines as exhausted the cells that present the same cellular dysfunction (typically, the absence of an expected effector response). The second approach primarily
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on MHC molecules, which reside on the surface of cortical epithelial cells. Only thymocytes that interact well with MHC-I or MHC-II will receive a vital "survival signal", while those that cannot interact strongly enough will receive no signal and die from neglect. This process ensures that the
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Regulatory T cells can develop either during normal development in the thymus, and are then known as thymic Treg cells, or can be induced peripherally and are called peripherally derived Treg cells. These two subsets were previously called "naturally occurring" and "adaptive" (or "induced"),
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represent some subsets of T cells that behave differently in immunity. They trigger rapid immune responses, regardless of the major histocompatibility complex (MHC) expression, unlike their conventional counterparts (CD4 T helper cells and CD8 cytotoxic T cells), which are dependent on the
192:. After migration to the thymus, the precursor cells mature into several distinct types of T cells. T cell differentiation also continues after they have left the thymus. Groups of specific, differentiated T cell subtypes have a variety of important functions in controlling and shaping the 1617:
defines as exhausted the cells that are produced by a given cause (typically, but not necessarily, chronic exposure to an antigen). Finally, the third approach primarily defines as exhausted the cells that present the same molecular markers (typically, programmed cell death protein 1 )."
1100:(γδ T cells) represent a small subset of T cells which possess a γδ TCR rather than the αβ TCR on the cell surface. The majority of T cells express αβ TCR chains. This group of T cells is much less common in humans and mice (about 2% of total T cells) and are found mostly in the gut 5634:
Liu L, Chang YJ, Xu LP, Zhang XH, Wang Y, Liu KY, Huang XJ (May 2018). "T cell exhaustion characterized by compromised MHC class I and II restricted cytotoxic activity associates with acute B lymphoblastic leukemia relapse after allogeneic hematopoietic stem cell transplantation".
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While in most cases activation is dependent on TCR recognition of antigen, alternative pathways for activation have been described. For example, cytotoxic T cells have been shown to become activated when targeted by other CD8 T cells leading to tolerization of the latter.
1116:γδ T cells recognize MHC class IB molecules. Human γδ T cells that use the Vγ9 and Vδ2 gene fragments constitute the major γδ T cell population in peripheral blood. These cells are unique in that they specifically and rapidly respond to a set of nonpeptidic phosphorylated 1025:. Once activated, these cells can perform functions ascribed to both helper and cytotoxic T cells: cytokine production and release of cytolytic/cell killing molecules. They are also able to recognize and eliminate some tumor cells and cells infected with herpes viruses. 417:
surviving thymocytes will have an 'MHC affinity' that means they can serve useful functions in the body, responding to MHC molecules to assist immune responses. The vast majority of developing thymocytes will not pass positive selection, and die during this process.
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Pekalski ML, García AR, Ferreira RC, Rainbow DB, Smyth DJ, Mashar M, Brady J, Savinykh N, Dopico XC, Mahmood S, Duley S, Stevens HE, Walker NM, Cutler AJ, Waldron-Lynch F, Dunger DB, Shannon-Lowe C, Coles AJ, Jones JL, Wallace C, Todd JA, Wicker LS (August 2017).
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and initiate immune responses. The ability of T cells to ignore healthy cells but respond when these same cells contain pathogen (or cancer) derived pMHC is known as antigen discrimination. The molecular mechanisms that underlie this process are controversial.
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molecules mature into CD8 "killer" cells. A thymocyte becomes a CD4 cell by down-regulating expression of its CD8 cell surface receptors. If the cell does not lose its signal, it will continue downregulating CD8 and become a CD4, both CD8 and CD4 cells are now
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Kjer-Nielsen L, Patel O, Corbett AJ, Le Nours J, Meehan B, Liu L, Bhati M, Chen Z, Kostenko L, Reantragoon R, Williamson NA, Purcell AW, Dudek NL, McConville MJ, O'Hair RA, Khairallah GN, Godfrey DI, Fairlie DP, Rossjohn J, McCluskey J (November 2012).
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molecules (positively selected CD4 cells must interact with these MHC class II molecules, thus APCs, which possess MHC class II, must be present for CD4 T-cell negative selection). Thymocytes that interact too strongly with the self-antigen receive an
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Woo SR, Turnis ME, Goldberg MV, Bankoti J, Selby M, Nirschl CJ, Bettini ML, Gravano DM, Vogel P, Liu CL, Tangsombatvisit S, Grosso JF, Netto G, Smeltzer MP, Chaux A, Utz PJ, Workman CJ, Pardoll DM, Korman AJ, Drake CG, Vignali DA (February 2012).
253:" response. For this reason, these regulatory T cells have also been called "suppressor" T cells. These same regulatory T cells can also be co-opted by cancer cells to prevent the recognition of, and an immune response against, tumor cells. 1692:
transfer. While there are data showing that induction of T cell exhaustion can be beneficial for transplantation it also carries disadvantages among which can be counted increased number of infections and the risk of tumor development.
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A critical step in T cell maturation is making a functional T cell receptor (TCR). Each mature T cell will ultimately contain a unique TCR that reacts to a random pattern, allowing the immune system to recognize many different types of
1328:); PI3K also acts on PIP2, phosphorylating it to produce phosphatidlyinositol-3,4,5-trisphosphate (PIP3). DAG binds and activates some PKCs. Most important in T cells is PKC-θ, critical for activating the transcription factors 787:
molecules, present on the surface of all nucleated cells. Cytotoxic T cells also produce the key cytokines IL-2 and IFNγ. These cytokines influence the effector functions of other cells, in particular macrophages and NK cells.
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Shi XL, de Mare-Bredemeijer EL, Tapirdamaz Ö, Hansen BE, van Gent R, van Campenhout MJ, Mancham S, Litjens NH, Betjes MG, van der Eijk AA, Xia Q, van der Laan LJ, de Jonge J, Metselaar HJ, Kwekkeboom J (September 2015).
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The second signal comes from co-stimulation, in which surface receptors on the APC are induced by a relatively small number of stimuli, usually products of pathogens, but sometimes breakdown products of cells, such as
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by about 3% a year throughout middle age, a corresponding fall in the thymic production of naive T cells occurs, leaving peripheral T cell expansion and regeneration to play a greater role in protecting older people.
363:. This process is essential in developing immunity to threats that the immune system has not encountered before, since due to random variation there will always be at least one TCR to match any new pathogen. 286:. The process of differentiation then proceeds to a common lymphoid progenitor (CLP), which can only differentiate into T, B or NK cells. These CLP cells then migrate via the blood to the thymus, where they 1136:) and corresponding mononucleotide conjugates, in addition to IPP and DMAPP. Plant cells produce both types of phosphoantigens. Drugs activating human Vγ9/Vδ2 T cells comprise synthetic phosphoantigens and 1620:
Dysfunctional T cells are characterized by progressive loss of function, changes in transcriptional profiles and sustained expression of inhibitory receptors. At first, cells lose their ability to produce
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Marusina AI, Ono Y, Merleev AA, Shimoda M, Ogawa H, Wang EA, Kondo K, Olney L, Luxardi G, Miyamura Y, Yilma TD, Villalobos IB, Bergstrom JW, Kronenberg DG, Soulika AM, Adamopoulos IE, Maverakis E (2017).
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A unique feature of T cells is their ability to discriminate between healthy and abnormal (e.g. infected or cancerous) cells in the body. Healthy cells typically express a large number of self derived
223:, to recruit other types of cells when mounting an immune response. A different population of T cells, the CD4+ T cells, function as "helper cells". Unlike CD8+ killer T cells, the CD4+ helper T (T 5069:
Wei Z, Li P, Yao Y, Deng H, Yi S, Zhang C, Wu H, Xie X, Xia M, He R, Yang XP, Tang ZH (July 2018). "Alpha-lactose reverses liver injury via blockade of Tim-3-mediated CD8 apoptosis in sepsis".
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Bianchini E, De Biasi S, Simone AM, Ferraro D, Sola P, Cossarizza A, Pinti M (March 2017). "Invariant natural killer T cells and mucosal-associated invariant T cells in multiple sclerosis".
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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.
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exists as a complex of several proteins. The actual T cell receptor is composed of two separate peptide chains, which are produced from the independent T cell receptor alpha and beta (
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that regulate or assist the immune response. These cells can differentiate into one of several subtypes, which have different roles. Cytokines direct T cells into particular subtypes.
1629:, which is followed by the loss of high proliferative capacity and cytotoxic potential, and eventually leads to their deletion. Exhausted T cells typically indicate higher levels of 3695:
Janssen EM, Lemmens EE, Wolfe T, Christen U, von Herrath MG, Schoenberger SP (February 2003). "CD4+ T cells are required for secondary expansion and memory in CD8+ T lymphocytes".
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cells, which are thought to originate during an immune response and act by producing suppressive molecules. Tr1 cells are associated with IL-10, and Th3 cells are associated with
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Abbas AK, Benoist C, Bluestone JA, Campbell DJ, Ghosh S, Hori S, Jiang S, Kuchroo VK, Mathis D, Roncarolo MG, Rudensky A, Sakaguchi S, Shevach EM, Vignali DA, Ziegler SF (2013).
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bacterially-infected cells. MAIT cells can also be activated through MR1-independent signaling. In addition to possessing innate-like functions, this T cell subset supports the
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co-receptor. The newly arrived CLP cells are CD4CD8CD44CD25ckit cells, and are termed early thymic progenitor (ETP) cells. These cells will then undergo a round of division and
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de Mare-Bredemeijer EL, Shi XL, Mancham S, van Gent R, van der Heide-Mulder M, de Boer R, Heemskerk MH, de Jonge J, van der Laan LJ, Metselaar HJ, Kwekkeboom J (August 2015).
5022:"Frontline Science: Defects in immune function in patients with sepsis are associated with PD-1 or PD-L1 expression and can be restored by antibodies targeting PD-1 or PD-L1" 4922:
Boomer JS, To K, Chang KC, Takasu O, Osborne DF, Walton AH, Bricker TL, Jarman SD, Kreisel D, Krupnick AS, Srivastava A, Swanson PE, Green JM, Hotchkiss RS (December 2011).
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Haynes BF, Markert ML, Sempowski GD, Patel DD, Hale LP (2000). "The role of the thymus in immune reconstitution in aging, bone marrow transplantation, and HIV-1 infection".
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Halloran PF, Chang J, Famulski K, Hidalgo LG, Salazar ID, Merino Lopez M, Matas A, Picton M, de Freitas D, Bromberg J, Serón D, Sellarés J, Einecke G, Reeve J (July 2015).
2798:"CD4+ virtual memory: Antigen-inexperienced T cells reside in the naïve, regulatory, and memory T cell compartments at similar frequencies, implications for autoimmunity" 219:– this means that they are able to directly kill virus-infected cells, as well as cancer cells. CD8+ T cells are also able to use small signalling proteins, known as 5959: 4819:
Penaloza-MacMaster P, Kamphorst AO, Wieland A, Araki K, Iyer SS, West EE, O'Mara L, Yang S, Konieczny BT, Sharpe AH, Freeman GJ, Rudensky AY, Ahmed R (August 2014).
1645:. Exhaustion can develop during chronic infections, sepsis and cancer. Exhausted T cells preserve their functional exhaustion even after repeated antigen exposure. 1250:
and ICOS, but these largely depend upon CD28 for their expression. The second signal licenses the T cell to respond to an antigen. Without it, the T cell becomes
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The first signal is provided by binding of the T cell receptor to its cognate peptide presented on MHCII on an APC. MHCII is restricted to so-called professional
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Sallusto F, Lenig D, Förster R, Lipp M, Lanzavecchia A (1999). "Two subsets of memory T lymphocytes with distinct homing potentials and effector functions".
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that activates the transcription of a pleiotropic set of genes, most notable, IL-2, a cytokine that promotes long-term proliferation of activated T cells.
375:, called negative selection. If both positive and negative selection are successful, the TCR becomes fully operational and the thymocyte becomes a T cell. 3845:
Jennifer Rolland and Robyn O'Hehir, "Turning off the T cells: Peptides for treatment of allergic Diseases," Today's life science publishing, 1999, Page 32
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cell depends on its subtype (such as T-helper1, T-helper2, T-helper17, regulatory T-cell), which is distinguished by the types of cytokines they secrete.
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Gassa A, Jian F, Kalkavan H, Duhan V, Honke N, Shaabani N, Friedrich SK, Dolff S, Wahlers T, Kribben A, Hardt C, Lang PA, Witzke O, Lang KS (2016).
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are able to distinguish invading cells from "self". This prevents immune cells from inappropriately reacting against one's own cells, known as an "
5829: 1364:. DAG activates PKC-θ, which then phosphorylates CARMA1, causing it to unfold and function as a scaffold. The cytosolic domains bind an adapter 404:
The process of positive selection takes 3 to 4 days and occurs in the thymic cortex. Double-positive thymocytes (CD4/CD8) migrate deep into the
5345:"CMV Primary Infection Is Associated With Donor-Specific T Cell Hyporesponsiveness and Fewer Late Acute Rejections After Liver Transplantation" 1274: 1124:, which are produced by virtually all living cells. The most common phosphoantigens from animal and human cells (including cancer cells) are 3427:
Serriari NE, Eoche M, Lamotte L, Lion J, Fumery M, Marcelo P, Chatelain D, Barre A, Nguyen-Khac E, Lantz O, Dupas JL, Treiner E (May 2014).
3031:"Invariant and noninvariant natural killer T cells exert opposite regulatory functions on the immune response during murine schistosomiasis" 4431:"Hepatitis B Virus-Specific CD8+ T Cells Maintain Functional Exhaustion after Antigen Reexposure in an Acute Activation Immune Environment" 3029:
Mallevaey T, Fontaine J, Breuilh L, Paget C, Castro-Keller A, Vendeville C, Capron M, Leite-de-Moraes M, Trottein F, Faveeuw C (May 2007).
5262:"Cytomegalovirus-Induced Expression of CD244 after Liver Transplantation Is Associated with CD8+ T Cell Hyporesponsiveness to Alloantigen" 2000:"Distinct phases in the positive selection of CD8+ T cells distinguished by intrathymic migration and T-cell receptor signaling patterns" 4621:
Okagawa T, Konnai S, Nishimori A, Maekawa N, Goto S, Ikebuchi R, Kohara J, Suzuki Y, Yamada S, Kato Y, Murata S, Ohashi K (June 2018).
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Milstein O, Hagin D, Lask A, Reich-Zeliger S, Shezen E, Ophir E, Eidelstein Y, Afik R, Antebi YE, Dustin ML, Reisner Y (January 2011).
1234:. The only co-stimulatory receptor expressed constitutively by naive T cells is CD28, so co-stimulation for these cells comes from the 4723:"Cell-intrinsic transforming growth factor-beta signaling mediates virus-specific CD8+ T cell deletion and viral persistence in vivo" 317:(DN1) cells. To become T cells, the thymocytes must undergo multiple DN stages as well as positive selection and negative selection. 1758: 440:. The potentially autoimmune cells are removed by the following process of negative selection, which occurs in the thymic medulla. 866:
stands for terminally differentiated effector memory cells re-expressing CD45RA, which is a marker usually found on naive T cells.
6236: 5719: 5672:"PD-1(hi)TIM-3(+) T cells associate with and predict leukemia relapse in AML patients post allogeneic stem cell transplantation" 5163:"Disappearance of T Cell-Mediated Rejection Despite Continued Antibody-Mediated Rejection in Late Kidney Transplant Recipients" 4821:"Interplay between regulatory T cells and PD-1 in modulating T cell exhaustion and viral control during chronic LCMV infection" 543:
Depiction of the various key subsets of CD4-positive T cells with corresponding associated cytokines and transcription factors.
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Godfrey DI, Uldrich AP, McCluskey J, Rossjohn J, Moody DB (November 2015). "The burgeoning family of unconventional T cells".
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Wu H, Arron JR (November 2003). "TRAF6, a molecular bridge spanning adaptive immunity, innate immunity and osteoimmunology".
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and in the peripheral circulation. (Note- CD44 expression is usually used to distinguish murine naive from memory T cells).
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and serves to prevent the formation of self-reactive T cells that are capable of inducing autoimmune diseases in the host.
371:(MHC) in a process known as positive selection. The thymocyte must also ensure that it does not react adversely to "self" 6092: 3185:
Eckle SB, Corbett AJ, Keller AN, Chen Z, Godfrey DI, Liu L, Mak JY, Fairlie DP, Rossjohn J, McCluskey J (December 2015).
2142: 1765:. Bethesda, MD: National Institutes of Health, U.S. Department of Health and Human Services. 17 June 2001. Archived from 1466: 1203: 1034: 1674:
are also able to trigger exhaustion. Last known factors that can play a role in T cell exhaustion are regulatory cells.
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protein, (B7.1 and B7.2, respectively) on the APC. Other receptors are expressed upon activation of the T cell, such as
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A thymocyte's fate is determined during positive selection. Double-positive cells (CD4/CD8) that interact well with MHC
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and cytotoxic T cells, which leads to a larger immune response. The specific adaptive immune response regulated by the T
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of T cells and/or defects on function of individual T cells. Complete insufficiency of T cell function can result from
862:. These memory T cells lack lymph node-homing receptors and are thus found in the peripheral circulation and tissues. T 5444:"Immune inhibitory molecules LAG-3 and PD-1 synergistically regulate T-cell function to promote tumoral immune escape" 5212:"Exhaustive differentiation of alloreactive CD8+ T cells: critical for determination of graft acceptance or rejection" 771:
cells, CTLs, T-killer cells, killer T cells) destroy virus-infected cells and tumor cells, and are also implicated in
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Zelle-Rieser C, Thangavadivel S, Biedermann R, Brunner A, Stoitzner P, Willenbacher E, et al. (November 2016).
6575: 6128: 1717: 1454: 1320:. PLC-γ cleaves PI(4,5)P2 on the inner leaflet of the membrane to create the active intermediaries diacylglycerol ( 1305: 1018: 413: 368: 129: 876:) occupy tissues (skin, lung, etc.) without recirculating. One cell surface marker that has been associated with T 1671: 1059:
metabolites to MAIT cells. After the presentation of foreign antigen by MR1, MAIT cells secrete pro-inflammatory
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Several other types of T cells have suppressive activity, but do not express FOXP3 constitutively. These include
912:. Their major role is to shut down T cell–mediated immunity toward the end of an immune reaction and to suppress 3746:
Shedlock DJ, Shen H (April 2003). "Requirement for CD4 T cell help in generating functional CD8 T cell memory".
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is needed to establish exhaustion. Another factor able to induce exhaustion are inhibitory receptors including
1561: 1294: 869: 3799:"CD4+ T cells are required for the maintenance, not programming, of memory CD8+ T cells after acute infection" 6513: 6051: 4025: 3855:
Maverakis E, Kim K, Shimoda M, Gershwin M, Patel F, Wilken R, Raychaudhuri S, Ruhaak LR, Lebrilla CB (2015).
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protein on their cell surface. Cytotoxic T cells recognize their targets by binding to short peptides (8-11
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mission to study how "deficiencies in the human immune system are affected by a microgravity environment".
4251: 2352:"T Helper (Th) Cell Profiles in Pregnancy and Recurrent Pregnancy Losses: Th1/Th2/Th9/Th17/Th22/Tfh Cells" 278:. The HSC then differentiate into multipotent progenitors (MPP) which retain the potential to become both 6161: 6156: 5996: 4531:"Progressive loss of memory T cell potential and commitment to exhaustion during chronic viral infection" 1626: 1511: 1384: 1336:
is released from the membrane by PLC-γ and diffuses rapidly to activate calcium channel receptors on the
1301: 1290: 1129: 708: 5544:"Cancer-associated fibroblasts induce antigen-specific deletion of CD8 T Cells to protect tumour cells" 1642: 1407: 6399: 1543: 1380: 1273:
proteins: CD3εγ and CD3εδ heterodimers and, most important, a CD3ζ homodimer, which has a total of six
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immune response and has a memory-like phenotype. Furthermore, MAIT cells are thought to play a role in
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At the DN2 stage (CD44CD25), cells upregulate the recombination genes RAG1 and RAG2 and re-arrange the
5840: 3647: 2194:"Neonatal and adult recent thymic emigrants produce IL-8 and express complement receptors CR1 and CR2" 110: 5866: 5394:"Recipient T Cell Exhaustion and Successful Adoptive Transfer of Haploidentical Natural Killer Cells" 4482:"CD4+ T cells are required to sustain CD8+ cytotoxic T-cell responses during chronic viral infection" 3294:"MR1-Restricted Mucosal-Associated Invariant T Cells and Their Activation during Infectious Diseases" 2896:"Modulation of autoimmune diseases by interleukin (IL)-17 producing regulatory T helper (Th17) cells" 1548: 1317: 1191: 1125: 643:
Produce an inflammatory response, key for defense against intracellular bacteria, viruses and cancer.
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Conforti L (February 2012). "The ion channel network in T lymphocytes, a target for immunotherapy".
3478:"MR1 antigen presentation to mucosal-associated invariant T cells was highly conserved in evolution" 3345:"Mucosal-associated invariant T cells in autoimmunity, immune-mediated diseases and airways disease" 424:
molecules will eventually become CD4 "helper" cells, whereas thymocytes that interact well with MHC
3977:"CTLs respond with activation and granule secretion when serving as targets for T cell recognition" 1149: 1109: 957: 267: 199:
One of these functions is immune-mediated cell death, and it is carried out by two major subtypes:
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Williams RL, Cooley S, Bachanova V, Blazar BR, Weisdorf DJ, Miller JS, Verneris MR (March 2018).
4870:"The late phase of sepsis is characterized by an increased microbiological burden and death rate" 4429:
Wang Q, Pan W, Liu Y, Luo J, Zhu D, Lu Y, Feng X, Yang X, Dittmer U, Lu M, Yang D, Liu J (2018).
3429:"Innate mucosal-associated invariant T (MAIT) cells are activated in inflammatory bowel diseases" 2978:
de Araújo ND, Gama FM, de Souza Barros M, Ribeiro TL, Alves FS, Xabregas LA, et al. (2021).
2141:
Hinterberger M, Aichinger M, Prazeres da Costa O, Voehringer D, Hoffmann R, Klein L (June 2010).
1655: 1395: 1219: 1179: 909: 594: 470: 4868:
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6682: 6601: 6319: 6255: 6043: 2648:"Multiparameter flow cytometric analysis of CD4 and CD8 T cell subsets in young and old people" 2596:"Molecular signatures distinguish human central memory from effector memory CD8 T cell subsets" 1820:
T cells ... derive their from the organs in which they develop. T cells develop in the thymus
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Starr TK, Jameson SC, Hogquist KA (2003-01-01). "Positive and negative selection of T cells".
6548: 6404: 6382: 6082: 6059: 6035: 5493:"T cells in multiple myeloma display features of exhaustion and senescence at the tumor site" 4971:
Shindo Y, McDonough JS, Chang KC, Ramachandra M, Sasikumar PG, Hotchkiss RS (February 2017).
3673: 2070: 1478: 1337: 1333: 1002: 997: 582:
glycoprotein on their surfaces. Helper T cells become activated when they are presented with
275: 170: 5211: 3187:"Recognition of Vitamin B Precursors and Byproducts by Mucosal Associated Invariant T Cells" 457:
positive (AIRE) to properly express self-antigens from all tissues of the body on their MHC
6677: 6565: 6498: 6409: 6333: 6106: 5923: 5859: 5757: 5555: 3755: 3704: 3601: 3489: 2503: 2011: 1536: 1494: 1450: 1354: 1043:, effector-like qualities. In humans, MAIT cells are found in the blood, liver, lungs, and 1040: 1014: 936: 454: 8: 6687: 6625: 6560: 6543: 6377: 6295: 6115: 5954: 4331:
Blank CU, Haining WN, Held W, Hogan PG, Kallies A, Lugli E, et al. (November 2019).
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3759: 3708: 3605: 3493: 2507: 2015: 1926:"Impaired CD8 T cell memory and CD4 T cell primary responses in IL-7R alpha mutant mice" 756: 28: 6445: 6087: 6074: 5809: 5780: 5745: 5696: 5671: 5576: 5543: 5519: 5492: 5468: 5443: 5418: 5393: 5374: 5242: 5187: 5162: 5138: 5113: 5094: 5046: 5021: 4997: 4972: 4948: 4923: 4896: 4869: 4845: 4820: 4796: 4771: 4747: 4722: 4698: 4673: 4649: 4622: 4603: 4555: 4530: 4457: 4430: 4406: 4381: 4357: 4332: 4308: 4281: 4178: 4153: 4129: 4104: 4077: 4050: 4001: 3976: 3957: 3881: 3856: 3823: 3798: 3779: 3728: 3625: 3566: 3512: 3477: 3453: 3428: 3369: 3344: 3320: 3293: 3264: 3237: 3213: 3186: 3162: 3137: 3113: 3086: 3055: 3030: 3006: 2979: 2960: 2912: 2895: 2876: 2822: 2797: 2772: 2747: 2723: 2698: 2674: 2647: 2628: 2576: 2527: 2430: 2405: 2378: 2351: 2327: 2302: 2218: 2193: 2173: 2123: 2034: 1999: 1950: 1925: 1924:
Osborne LC, Dhanji S, Snow JW, Priatel JJ, Ma MC, Miners MJ, et al. (March 2007).
1860: 1833: 1341: 1190:. The signalling pathways downstream from co-stimulatory molecules usually engages the 1097: 1092: 1076: 1072: 947: 4506: 4481: 3476:
Huang S, Martin E, Kim S, Yu L, Soudais C, Fremont DH, Lantz O, Hansen TH (May 2009).
2110: 2094:"The MHC reactivity of the T cell repertoire prior to positive and negative selection" 2093: 1552: 1182:. Both are required for production of an effective immune response; in the absence of 241:
are yet another distinct population of T cells that provide the critical mechanism of
6493: 6259: 6138: 6019: 5938: 5918: 5815: 5785: 5701: 5652: 5616: 5581: 5524: 5473: 5423: 5366: 5324: 5283: 5234: 5192: 5143: 5086: 5051: 5002: 4953: 4901: 4850: 4801: 4752: 4703: 4654: 4595: 4560: 4511: 4497: 4462: 4411: 4397: 4362: 4313: 4231: 4183: 4134: 4105:"Quantitative challenges in understanding ligand discrimination by alphabeta T cells" 4082: 4006: 3949: 3912: 3886: 3828: 3771: 3720: 3677: 3617: 3558: 3517: 3458: 3409: 3374: 3325: 3269: 3218: 3167: 3118: 3060: 3011: 2952: 2917: 2868: 2827: 2777: 2728: 2679: 2620: 2568: 2519: 2476: 2466: 2435: 2383: 2332: 2283: 2279: 2246: 2223: 2165: 2115: 2074: 2039: 1978: 1955: 1906: 1865: 1675: 1596: 1556: 1470: 1442: 1425: 905: 900: 567: 491: 482:
signal that leads to cell death. However, some of these cells are selected to become
238: 5246: 5098: 4607: 3961: 3783: 3570: 2964: 2880: 2632: 2580: 2177: 2127: 700:
Defense against helminths (parasitic worms) and cell-dependent allergic inflammation
270:(HSC) which reside in the bone marrow. In some cases, the origin might be the fetal 6672: 6613: 6553: 6525: 6520: 6488: 6475: 6465: 6011: 5987: 5775: 5765: 5691: 5683: 5644: 5608: 5571: 5563: 5514: 5504: 5463: 5455: 5413: 5405: 5378: 5356: 5314: 5273: 5226: 5182: 5174: 5133: 5125: 5078: 5041: 5033: 5020:
Patera AC, Drewry AM, Chang K, Beiter ER, Osborne D, Hotchkiss RS (December 2016).
4992: 4984: 4943: 4935: 4891: 4881: 4840: 4832: 4791: 4783: 4742: 4734: 4693: 4685: 4644: 4634: 4587: 4550: 4542: 4501: 4493: 4452: 4442: 4401: 4393: 4352: 4344: 4303: 4293: 4173: 4165: 4124: 4116: 4072: 4062: 3996: 3988: 3941: 3876: 3868: 3818: 3810: 3763: 3712: 3669: 3629: 3609: 3548: 3507: 3497: 3448: 3440: 3401: 3364: 3356: 3315: 3305: 3259: 3249: 3208: 3198: 3157: 3149: 3108: 3098: 3050: 3042: 3001: 2991: 2980:"Translating Unconventional T Cells and Their Roles in Leukemia Antitumor Immunity" 2944: 2907: 2858: 2817: 2809: 2767: 2759: 2718: 2710: 2669: 2659: 2610: 2558: 2531: 2511: 2458: 2425: 2417: 2373: 2363: 2322: 2314: 2275: 2213: 2205: 2157: 2105: 2066: 2029: 2019: 1945: 1937: 1896: 1855: 1845: 1732: 1430: 1270: 1017:. Unlike conventional T cells that recognize protein peptide antigens presented by 803: 772: 764: 751: 490:, also known as recent thymic emigrants. This process is an important component of 242: 200: 150: 5670:
Kong Y, Zhang J, Claxton DF, Ehmann WC, Rybka WB, Zhu L, et al. (July 2015).
5459: 3732: 2457:. Advances in Experimental Medicine and Biology. Vol. 841. pp. 209–230. 1361: 662:
Immunologically important against extracellular pathogens, such as worm infections
6508: 6003: 5303:"IL-10 Induces T Cell Exhaustion During Transplantation of Virus Infected Hearts" 4738: 4298: 4120: 3992: 3153: 1901: 1884: 1532: 1523: 1446: 1258: 1163: 1137: 1044: 961: 950: 353: 193: 166: 98: 5230: 3405: 2615: 2563: 2546: 2462: 2140: 287: 6535: 6480: 6421: 6341: 6314: 5750:
Proceedings of the National Academy of Sciences of the United States of America
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Williams MA, Bevan MJ (2007-01-01). "Effector and memory CTL differentiation".
3482:
Proceedings of the National Academy of Sciences of the United States of America
2846: 2813: 2004:
Proceedings of the National Academy of Sciences of the United States of America
1792: 1667: 1458: 1183: 1121: 760:
Superresolution image of a group of cytotoxic T cells surrounding a cancer cell
466: 283: 46: 5509: 4988: 4639: 4348: 4067: 3238:"Mucosal-associated invariant T-cells: new players in anti-bacterial immunity" 2421: 2209: 1998:
Ross JO, Melichar HJ, Au-Yeung BB, Herzmark P, Weiss A, Robey EA (June 2014).
1297:, which allows the aggregation of signalling complexes around these proteins. 6666: 6503: 6392: 6210: 6192: 6030: 5992: 5913: 5278: 5261: 5037: 4447: 4169: 3310: 3254: 3103: 2763: 2368: 1622: 1500: 1211: 835:(CD62L). Central memory T cells also have intermediate to high expression of 808: 797: 736:
Pathogenesis of allergic airway diseases and predominantly anti-inflammatory
563: 547: 534: 325: 228: 204: 154: 69: 5770: 4924:"Immunosuppression in patients who die of sepsis and multiple organ failure" 3857:"Glycans in the immune system and The Altered Glycan Theory of Autoimmunity" 3767: 3502: 3203: 3087:"The Role of Mucosal Associated Invariant T Cells in Antimicrobial Immunity" 2024: 1162:
Activation of CD4 T cells occurs through the simultaneous engagement of the
6470: 6455: 6450: 6387: 6285: 6025: 5933: 5928: 5789: 5705: 5656: 5620: 5585: 5528: 5477: 5427: 5370: 5328: 5287: 5238: 5196: 5178: 5147: 5129: 5090: 5055: 5006: 4957: 4939: 4905: 4854: 4805: 4756: 4707: 4658: 4599: 4564: 4466: 4415: 4366: 4317: 4187: 4138: 4086: 4010: 3953: 3890: 3832: 3775: 3724: 3681: 3621: 3562: 3521: 3462: 3413: 3378: 3329: 3273: 3222: 3171: 3138:"Mucosal associated invariant T cells and the immune response to infection" 3122: 3064: 3015: 2996: 2956: 2921: 2872: 2831: 2781: 2732: 2683: 2646:
Koch S, Larbi A, Derhovanessian E, Özcelik D, Naumova E, Pawelec G (2008).
2624: 2547:"Loss of CD45R and gain of UCHL1 reactivity is a feature of primed T cells" 2523: 2480: 2439: 2387: 2336: 2287: 2227: 2169: 2078: 2043: 1959: 1910: 1869: 1388: 1243: 590: 487: 462: 437: 310: 279: 250: 246: 216: 64: 5490: 4515: 2664: 2572: 2303:"APC-derived cytokines and T cell polarization in autoimmune inflammation" 2119: 1850: 320:
Double negative thymocytes can be identified by the surface expression of
6205: 5908: 5903: 4836: 4546: 3553: 3536: 3046: 1941: 1727: 1601: 1369: 1346: 1321: 1048: 784: 559: 405: 181: 162: 5687: 5210:
Steger U, Denecke C, Sawitzki B, Karim M, Jones ND, Wood KJ (May 2008).
4282:"Immunological exhaustion: How to make a disparate concept operational?" 3716: 3613: 539: 261: 207:
T cells. (These are named for the presence of the cell surface proteins
40: 6593: 6346: 6252: 5882: 4818: 3945: 1638: 1313: 1117: 1056: 1052: 969: 855: 840: 832: 780: 571: 5361: 5344: 5341: 5319: 5302: 4787: 3444: 3360: 2714: 1021:(MHC) molecules, NKT cells recognize glycolipid antigens presented by 135: 6651: 6356: 6251: 6178: 6133: 5981: 5259: 5114:"The role of peripheral T-cell deletion in transplantation tolerance" 4209: 2318: 1286: 689: 479: 291: 5720:"U.S. FDA Approved Immune-Checkpoint Inhibitors and Immunotherapies" 5542:
Lakins MA, Ghorani E, Munir H, Martins CP, Shields JD (March 2018).
4886: 4591: 2948: 2894:
Singh B, Schwartz JA, Sandrock C, Bellemore SM, Nikoopour E (2013).
2863: 2594:
Willinger T, Freeman T, Hasegawa H, McMichael AJ, Callan MF (2005).
2161: 1154: 6641: 6426: 6414: 6372: 6326: 6290: 5947: 5898: 4689: 4529:
Angelosanto JM, Blackburn SD, Crawford A, Wherry EJ (August 2012).
3814: 2977: 1810:
Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002).
1791:
Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002).
1506: 1434: 1227: 1060: 965: 627: 598: 360: 297:
The earliest cells which arrived in the thymus are commonly termed
220: 50: 4674:"Interleukin-10 determines viral clearance or persistence in vivo" 4528: 2847:"Regulatory T cells: recommendations to simplify the nomenclature" 2515: 1329: 946:
gene can prevent regulatory T cell development, causing the fatal
6646: 6460: 6309: 6280: 6166: 5807: 2593: 2545:
Akbar AN, Terry L, Timms A, Beverley PC, Janossy G (April 1988).
1885:"One Niche to Rule Both Maintenance and Loss of Stemness in HSCs" 1175: 1132:(DMPP). Many microbes produce the active compound hydroxy-DMAPP ( 1113: 586: 583: 409: 372: 188:
gland to develop (or mature). T cells derive their name from the
3583: 1174:) on the T cell by the major histocompatibility complex (MHCII) 632: 6618: 6351: 6302: 5890: 4970: 2934: 2453:
Jia L, Wu C (2014). "The Biology and Functions of Th22 Cells".
1659: 1528: 1465:. Causes of partial insufficiencies of T cell function include 1251: 1187: 1133: 1101: 802:
Antigen-naive T cells expand and differentiate into memory and
555: 189: 185: 5851: 5811:
Immunobiology 5 : the immune system in health and disease
3974: 3391: 2893: 2645: 916:
that escaped the process of negative selection in the thymus.
5724:
Medical Writer Agency | 香港醫學作家 | MediPR | MediPaper Hong Kong
5391: 4721:
Tinoco R, Alcalde V, Yang Y, Sauer K, Zuniga EI (July 2009).
4620: 3643: 3028: 2844: 2350:
Wang W, Sung N, Gilman-Sachs A, Kwak-Kim J (18 August 2020).
2190: 1365: 1282: 1277:
motifs. The ITAM motifs on the CD3ζ can be phosphorylated by
1207: 1171: 1064: 939: 638: 554:
cells) assist other lymphocytes, including the maturation of
271: 81: 5160: 4102: 3587:"MR1 presents microbial vitamin B metabolites to MAIT cells" 2349: 1997: 1304:
recruits SLP-76 to the membrane, where it can then bring in
775:
rejection. These cells are defined by the expression of the
4671: 4382:"T-cell exhaustion: characteristics, causes and conversion" 4017: 3854: 3694: 2265: 1663: 1637:
and inhibitory receptors combined with lower expression of
1634: 1630: 1350: 1309: 1247: 1239: 1235: 1195: 1167: 1022: 859: 836: 813: 670: 483: 436:
This process does not filter for thymocytes that may cause
333: 5808:
Janeway Jr CA, Travers P, Walport M, Shlomchik MJ (2001).
5209: 4257:. The Leukemia & Lymphoma Society. May 2006. p. 2 1047:, defending against microbial activity and infection. The 942:
which can be used to identify the cells. Mutations of the
473:(APCs), allowing for presentation of self-antigens on MHC 5541: 4479: 4026:"SpaceX ready for CRS-3 Dragon launch and new milestones" 3085:
Napier RJ, Adams EJ, Gold MC, Lewinsohn DM (2015-07-06).
2794: 1488:
The main pathogens of concern in T cell deficiencies are
1477:(CBSs), and B cell and T cell combined disorders such as 1278: 935:
respectively. Both subsets require the expression of the
776: 651: 579: 337: 329: 321: 306: 302: 212: 208: 5440: 4769: 3537:"Bacteria, mucosal-associated invariant T cells and MR1" 3426: 3084: 2493: 1923: 1514:
are also more common and severe in T cell deficiencies.
597:(APCs). Once activated, they divide rapidly and secrete 340:
is expressed. Expression of both CD4 and CD8 makes them
5019: 4223: 3235: 2544: 1809: 1790: 1083:, although definitive evidence is yet to be published. 1028: 5669: 5300: 4867: 4770:
Veiga-Parga T, Sehrawat S, Rouse BT (September 2013).
4720: 4480:
Matloubian M, Concepcion RJ, Ahmed R (December 1994).
4154:"An induced rebinding model of antigen discrimination" 4048: 3184: 1831: 1648: 854:
cells) express CD45RO but lack expression of CCR7 and
5837:
National Institute of Allergy and Infectious Diseases
5743: 4330: 4279: 1055:, is responsible for presenting bacterially-produced 839:. This memory subpopulation is commonly found in the 681:
Defense against gut pathogens and at mucosal barriers
486:
cells. The remaining cells exit the thymus as mature
262:
Origin, early development and migration to the thymus
215:.) CD8+ T cells, also known as "killer T cells", are 4219: 4217: 3906: 3902: 3900: 3291: 2056: 1353:, which then translocates to the nucleus. NFAT is a 1039:
Mucosal associated invariant T (MAIT) cells display
858:. They also have intermediate to high expression of 5744:Bhadra R, Gigley JP, Weiss LM, Khan IA (May 2011). 4921: 4772:"Role of regulatory T cells during virus infection" 4151: 1977:(8th ed.). Garland Science. pp. 301–305. 1269:) genes. The other proteins in the complex are the 4623:"+ T cells during bovine leukemia virus infection" 4280:Kaminski H, Lemoine M, Pradeu T (September 2021). 4224:Jones J, Bannister BA, Gillespie SH, eds. (2006). 4049:Belikov AV, Schraven B, Simeoni L (October 2015). 3236:Ussher JE, Klenerman P, Willberg CB (2014-10-08). 2745: 2399: 2397: 5111: 4917: 4915: 4214: 4098: 4096: 3897: 2403: 2091: 593:molecules, which are expressed on the surface of 6664: 5112:Wells AD, Li XC, Strom TB, Turka LA (May 2001). 4973:"Anti-PD-L1 peptide improves survival in sepsis" 4103:Feinerman O, Germain RN, Altan-Bonnet G (2008). 3797:Sun JC, Williams MA, Bevan MJ (September 2004). 3796: 3475: 2455:T Helper Cell Differentiation and Their Function 523: 5633: 4199: 4197: 2394: 1832:Luckheeram RV, Zhou R, Verma AD, Xia B (2012). 1818:(4th ed.). Garland Science. p. 1367. 1535:, and accounts for perhaps one in ten cases of 1285:. Lck and/or ZAP-70 can also phosphorylate the 53:(center), and a T lymphocyte (right); colorized 4912: 4093: 3292:Howson LJ, Salio M, Cerundolo V (2015-06-16). 2748:"Alternative memory in the CD8 T cell lineage" 2245:(8th ed.). Garland Science. p. 297. 1786: 1784: 1186:, T cell receptor signalling alone results in 739:Crohn's Disease, Rheumatoid Arthritis, Tumors 344:, and matures into either CD4+ or CD8+ cells. 6237: 5867: 5167:Journal of the American Society of Nephrology 4428: 4244: 3659: 3135: 2300: 5068: 5062: 4578:Wherry EJ (June 2011). "T cell exhaustion". 4422: 4275: 4273: 4271: 4194: 4145: 3968: 3925: 3745: 2887: 2838: 2294: 2259: 2134: 1834:"CD4⁺T cells: differentiation and functions" 1666:(LAG3). Soluble molecules such as cytokines 5814:(5th ed.). New York: Garland Science. 5398:Biology of Blood and Marrow Transplantation 2696: 2538: 2092:Zerrahn J, Held W, Raulet DH (March 1997). 1781: 1590:It has been suggested that this section be 332:. Still during the double negative stages, 6244: 6230: 5874: 5860: 4379: 3534: 2404:Saravia J, Chapman NM, Chi H (July 2019). 1812:"Helper t Cells and Lymphocyte Activation" 1793:"Helper T Cells and Lymphocyte Activation" 1469:(AIDS), and hereditary conditions such as 894: 497: 161:. T cells can be distinguished from other 39: 27: 5779: 5769: 5695: 5575: 5518: 5508: 5467: 5417: 5360: 5318: 5277: 5186: 5137: 5045: 4996: 4947: 4895: 4885: 4844: 4795: 4746: 4697: 4648: 4638: 4554: 4505: 4456: 4446: 4405: 4356: 4307: 4297: 4268: 4177: 4128: 4076: 4066: 4000: 3907:Tatham P, Gomperts BD, Kramer IM (2003). 3880: 3822: 3552: 3511: 3501: 3452: 3368: 3319: 3309: 3263: 3253: 3212: 3202: 3161: 3112: 3102: 3054: 3005: 2995: 2911: 2862: 2821: 2771: 2722: 2673: 2663: 2614: 2562: 2429: 2377: 2367: 2326: 2217: 2109: 2033: 2023: 1949: 1900: 1859: 1849: 1682: 1539:. The main forms of T cell lymphoma are: 1401: 1289:on many other molecules, not least CD28, 1206:that are essential for the activation of 1140:, which upregulate endogenous IPP/DMAPP. 745: 184:. Developing T cells then migrate to the 5598: 3674:10.1146/annurev.immunol.25.022106.141548 2746:Lee YJ, Jameson SC, Hogquist KA (2011). 2071:10.1146/annurev.immunol.21.120601.141107 1662:, T cell membrane protein-3 (TIM3), and 1429: 1414: 1242:proteins, which together constitute the 1153: 1150:T-cell receptor § Signaling pathway 1009:of the innate immune system) bridge the 991: 880:is the intern αeβ7, also known as CD103. 755: 538: 378: 6114: 3931: 3648:"Understanding the Immune System (pdf)" 1838:Clinical & Developmental Immunology 412:. These self-antigens are expressed by 227:) cells function by further activating 6665: 4577: 4380:Yi JS, Cox MA, Zajac AJ (April 2010). 4227:Infection: Microbiology and Management 4023: 3911:. Amsterdam: Elsevier Academic Press. 3287: 3285: 3283: 3080: 3078: 3076: 3074: 2240: 1972: 1383:(TCAS) experiment was launched to the 1198:at the plasma membrane and recruiting 1086: 975: 528: 16:White blood cells of the immune system 6225: 5855: 4051:"T cells and reactive oxygen species" 3636: 3342: 3136:Gold MC, Lewinsohn DM (August 2011). 2452: 1882: 1594:out into another article titled 1005:(NKT cells – not to be confused with 520:gene or protein expression patterns. 443: 408:, where they are presented with self- 399: 266:All T cells originate from c-kitSca1 5497:Journal of Hematology & Oncology 5307:Cellular Physiology and Biochemistry 4825:The Journal of Experimental Medicine 3535:Chua WJ, Hansen TH (November 2010). 3433:Clinical and Experimental Immunology 2697:Shin H, Iwasaki A (September 2013). 1930:The Journal of Experimental Medicine 1664:lymphocyte activation gene 3 protein 1576: 1202:containing signaling molecules like 1178:and co-stimulatory molecules on the 1166:and a co-stimulatory molecule (like 1029:Mucosal associated invariant T cells 684:MS, Rheumatoid Arthritis, Psoriasis 6093:Mucosal associated invariant T cell 5349:American Journal of Transplantation 4152:Dushek O, van der Merwe PA (2014). 3280: 3191:The Journal of Biological Chemistry 3071: 2410:Cellular & Molecular Immunology 1649:During chronic infection and sepsis 1467:acquired immune deficiency syndrome 1035:Mucosal associated invariant T cell 908:are crucial for the maintenance of 722:Asthma and other allergic diseases 665:Asthma and other allergic diseases 13: 5800: 3385: 3336: 3229: 3178: 1567:Angioimmunoblastic T cell lymphoma 1394:T cell activation is modulated by 1349:. Calcineurin, in turn, activates 514: 347: 294:, the immature stage of a T cell. 149:are one of the important types of 45:Scanning electron micrograph of a 14: 6699: 1324:), inositol-1,4,5-trisphosphate ( 923:cells have been described—FOXP3 T 791: 451:medullary thymic epithelial cells 6129:Lymphokine-activated killer cell 5839:. September 2003. Archived from 4977:The Journal of Surgical Research 4498:10.1128/JVI.68.12.8056-8063.1994 4398:10.1111/j.1365-2567.2010.03255.x 4230:. Wiley-Blackwell. p. 435. 2699:"Tissue-resident memory T cells" 2280:10.1146/annurev.immunol.18.1.529 1799:(4th ed.). Garland Science. 1718:Chimeric antigen receptor T cell 1696: 1581: 1455:severe combined immunodeficiency 1019:major histocompatibility complex 574:. These cells are also known as 505: 414:thymic cortical epithelial cells 369:major histocompatibility complex 130:Anatomical terms of microanatomy 5881: 5737: 5712: 5663: 5627: 5592: 5535: 5484: 5434: 5385: 5335: 5294: 5253: 5203: 5154: 5105: 5013: 4964: 4861: 4812: 4763: 4714: 4665: 4614: 4571: 4522: 4473: 4373: 4324: 4042: 3848: 3839: 3790: 3739: 3688: 3653: 3577: 3528: 3469: 3420: 3129: 3022: 2971: 2928: 2788: 2739: 2690: 2639: 2587: 2487: 2446: 2406:"Helper T cell differentiation" 2343: 2234: 2184: 2085: 1656:programmed cell death protein 1 1340:, which induces the release of 1120:precursors, collectively named 719:Help B cells produce antibodies 290:. Henceforth they are known as 157:and play a central role in the 6571:Immunoglobulin class switching 4333:"Defining 'T cell exhaustion'" 2984:Journal of Immunology Research 2050: 1991: 1966: 1917: 1876: 1825: 1803: 1751: 1562:Anaplastic large cell lymphoma 1475:chromosomal breakage syndromes 972:have been added to this list. 870:Tissue-resident memory T cells 301:, as they express neither the 256: 1: 5460:10.1158/0008-5472.CAN-11-1620 4055:Journal of Biomedical Science 2111:10.1016/S0092-8674(00)81905-4 1816:Molecular Biology of the Cell 1797:Molecular Biology of the Cell 1759:"5. Hematopoietic Stem Cells" 1744: 1572: 1419: 1143: 829:C-C chemokine receptor type 7 524:Conventional adaptive T cells 6174:Type 3 innate lymphoid cells 6162:Type 2 innate lymphoid cells 6157:Type 1 innate lymphoid cells 6144:Uterine natural killer cells 6124:Cytokine-induced killer cell 5026:Journal of Leukocyte Biology 4739:10.1016/j.immuni.2009.06.015 4299:10.1371/journal.ppat.1009892 4121:10.1016/j.molimm.2007.03.028 3993:10.1182/blood-2010-05-283770 3154:10.1016/j.micinf.2011.03.007 2301:Gutcher I, Becher B (2007). 1902:10.1016/j.immuni.2016.12.003 1360:PLC-γ can also initiate the 7: 5231:10.1097/TP.0b013e31816dd64a 3662:Annual Review of Immunology 3541:Immunology and Cell Biology 3406:10.1016/j.imlet.2017.01.009 2616:10.4049/jimmunol.175.9.5895 2564:10.4049/jimmunol.140.7.2171 2463:10.1007/978-94-017-9487-9_8 2059:Annual Review of Immunology 1711: 1385:International Space Station 1130:dimethylallyl pyrophosphate 1106:intraepithelial lymphocytes 783:in length) associated with 605:CD4+ Helper T cell subsets 10: 6704: 6400:Polyclonal B cell response 5649:10.1016/j.clim.2018.02.009 5613:10.1016/j.clim.2011.11.009 5568:10.1038/s41467-018-03347-0 5410:10.1016/j.bbmt.2017.11.022 5083:10.1016/j.clim.2018.04.010 4337:Nature Reviews. Immunology 3873:10.1016/j.jaut.2014.12.002 2814:10.1016/j.jaut.2016.11.001 1700: 1549:Cutaneous T cell lymphomas 1544:Extranodal T cell lymphoma 1521: 1423: 1381:T-Cell Activation in Space 1147: 1090: 1081:inflammatory bowel disease 1032: 995: 919:Two major classes of CD4 T 898: 846:Effector memory T cells (T 795: 749: 532: 351: 6634: 6592: 6534: 6435: 6365: 6273: 6266: 6191: 6152: 6105: 6073: 5977: 5968: 5889: 5510:10.1186/s13045-016-0345-3 4989:10.1016/j.jss.2016.08.099 4640:10.1186/s13567-018-0543-9 4349:10.1038/s41577-019-0221-9 4068:10.1186/s12929-015-0194-3 2422:10.1038/s41423-019-0220-6 2210:10.1172/jci.insight.93739 1883:Kondo M (December 2016). 1517: 1463:cartilage–hair hypoplasia 1126:isopentenyl pyrophosphate 1104:, within a population of 823:Central memory T cells (T 461:peptides. Some mTECs are 268:haematopoietic stem cells 128: 116: 104: 92: 80: 75: 63: 58: 38: 26: 21: 5279:10.4049/jimmunol.1500440 5038:10.1189/jlb.4hi0616-255r 4448:10.3389/fimmu.2018.00219 4170:10.1016/j.it.2014.02.002 3311:10.3389/fimmu.2015.00303 3255:10.3389/fimmu.2014.00450 3104:10.3389/fimmu.2015.00344 2764:10.1016/j.it.2010.12.004 2369:10.3389/fimmu.2020.02025 2241:Murphy, Kenneth (2011). 1973:Murphy, Kenneth (2011). 1483:Wiskott–Aldrich syndrome 1220:antigen-presenting cells 819:Memory T cell subtypes: 616:Key Transcription Factor 595:antigen-presenting cells 471:antigen presenting cells 178:hematopoietic stem cells 159:adaptive immune response 6201:Hematopoietic stem cell 5960:Lymphoplasmacytoid cell 5771:10.1073/pnas.1015298108 4024:Graham W (2014-04-14). 3768:10.1126/science.1082305 3503:10.1073/pnas.0903196106 3298:Frontiers in Immunology 3242:Frontiers in Immunology 3204:10.1074/jbc.R115.685990 3091:Frontiers in Immunology 2802:Journal of Autoimmunity 2356:Frontiers in Immunology 2243:Janeway's Immunobiology 2025:10.1073/pnas.1408482111 1975:Janeway's Immunobiology 1490:intracellular pathogens 1396:reactive oxygen species 910:immunological tolerance 895:Regulatory CD4+ T cells 827:cells) express CD45RO, 498:TCR development summary 469:; this makes them AIRE 453:(mTECs). mTECs must be 6514:Tolerance in pregnancy 6256:adaptive immune system 5179:10.1681/ASN.2014060588 5130:10.1098/rstb.2001.0845 4940:10.1001/jama.2011.1829 3142:Microbes and Infection 3035:Infection and Immunity 1683:During transplantation 1438: 1402:Antigen discrimination 1159: 1011:adaptive immune system 1003:Natural killer T cells 985:unconventional T cells 884:Virtual memory T cells 761: 746:Cytotoxic CD8+ T cells 619:Role in immune defense 544: 467:thymic dendritic cells 176:T cells are born from 6549:Somatic hypermutation 6383:Polyclonal antibodies 6378:Monoclonal antibodies 6107:Innate lymphoid cells 6083:Natural killer T cell 5548:Nature Communications 5266:Journal of Immunology 4776:Immunological Reviews 3343:Hinks TS (May 2016). 2703:Immunological Reviews 2665:10.1186/1742-4933-5-6 2652:Immunity & Ageing 2603:Journal of Immunology 1763:Stem Cell Information 1531:of T cells is termed 1522:Further information: 1479:ataxia-telangiectasia 1451:hereditary conditions 1433: 1415:Clinical significance 1157: 1128:(IPP) and its isomer 998:Natural killer T cell 992:Natural killer T cell 759: 675:IL-17F, IL-17A, IL-22 542: 379:TCR β-chain selection 336:expression stops and 313:c-kit and are termed 276:embryonic development 165:by the presence of a 6566:Junctional diversity 6334:Antigen presentation 5676:Blood Cancer Journal 4837:10.1084/jem.20132577 4547:10.1128/JVI.00889-12 3554:10.1038/icb.2010.104 3047:10.1128/IAI.01178-06 2997:10.1155/2021/6633824 2752:Trends in Immunology 1942:10.1084/jem.20061871 1537:non-Hodgkin lymphoma 1495:Herpes simplex virus 1379:In spring 2014, the 1355:transcription factor 1281:and in turn recruit 1138:aminobisphosphonates 1015:innate immune system 1007:natural killer cells 937:transcription factor 914:autoreactive T cells 646:MS, Type 1 diabetes 578:as they express the 566:, and activation of 455:Autoimmune regulator 6561:V(D)J recombination 6544:Affinity maturation 6296:Antigenic variation 6075:Innate-like T cells 5955:Transitional B cell 5830:"The Immune System" 5762:2011PNAS..108.9196B 5688:10.1038/bcj.2015.58 5637:Clinical Immunology 5601:Clinical Immunology 5560:2018NatCo...9..948L 5071:Clinical Immunology 4627:Veterinary Research 4535:Journal of Virology 4486:Journal of Virology 4030:NASAspaceflight.com 3909:Signal transduction 3760:2003Sci...300..337S 3717:10.1038/nature01441 3709:2003Natur.421..852J 3614:10.1038/nature11605 3606:2012Natur.491..717K 3494:2009PNAS..106.8290H 2508:1999Natur.401..708S 2016:2014PNAS..111E2550R 2010:(25): E2550–E2558. 1851:10.1155/2012/925135 1738:Parafollicular cell 1723:Gut-specific homing 1232:heat shock proteins 1194:pathway generating 1098:Gamma delta T cells 1087:Gamma delta T cells 1073:autoimmune diseases 1063:and are capable of 981:Innate-like T cells 976:Innate-like T cells 703:Multiple Sclerosis 606: 529:Helper CD4+ T cells 389:V-D-J recombination 315:double-negative one 33:Animation of T cell 3946:10.1002/bies.10352 3394:Immunology Letters 2900:Indian J. Med. Res 2268:Annu. Rev. Immunol 1769:on 29 October 2016 1740:also called C cell 1439: 1345:can then activate 1160: 1093:Gamma delta T cell 1077:multiple sclerosis 948:autoimmune disease 906:Regulatory T cells 762: 613:Cytokines Produced 604: 545: 444:Negative selection 400:Positive selection 239:Regulatory T cells 111:H2.00.04.1.02007 6660: 6659: 6588: 6587: 6338:professional APCs 6219: 6218: 6187: 6186: 6101: 6100: 5821:978-0-8153-3642-6 5756:(22): 9196–9201. 5362:10.1111/ajt.13288 5320:10.1159/000443067 4788:10.1111/imr.12085 4580:Nature Immunology 4237:978-1-4051-2665-6 3918:978-0-12-289632-3 3803:Nature Immunology 3646:resource booklet 3445:10.1111/cei.12277 3361:10.1111/imm.12582 2943:(11): 1114–1123. 2937:Nature Immunology 2715:10.1111/imr.12087 2502:(6754): 708–712. 2472:978-94-017-9486-2 2150:Nature Immunology 1614: 1613: 1609: 1597:T cell exhaustion 1557:Mycosis fungoides 1512:fungal infections 1471:DiGeorge syndrome 1443:T cell deficiency 1426:T cell deficiency 927:cells and FOXP3 T 901:Regulatory T cell 765:Cytotoxic T cells 743: 742: 656:IL-4, IL-5, IL-13 622:Related diseases 568:cytotoxic T cells 492:central tolerance 387:locus, combining 151:white blood cells 144: 143: 139: 6695: 6554:Clonal selection 6526:Immune privilege 6521:Immunodeficiency 6476:Cross-reactivity 6466:Hypersensitivity 6271: 6270: 6246: 6239: 6232: 6223: 6222: 6139:Adaptive NK cell 6112: 6111: 5975: 5974: 5876: 5869: 5862: 5853: 5852: 5847: 5846:on 25 June 2009. 5845: 5834: 5825: 5794: 5793: 5783: 5773: 5741: 5735: 5734: 5732: 5731: 5716: 5710: 5709: 5699: 5667: 5661: 5660: 5631: 5625: 5624: 5596: 5590: 5589: 5579: 5539: 5533: 5532: 5522: 5512: 5488: 5482: 5481: 5471: 5438: 5432: 5431: 5421: 5389: 5383: 5382: 5364: 5339: 5333: 5332: 5322: 5298: 5292: 5291: 5281: 5257: 5251: 5250: 5216: 5207: 5201: 5200: 5190: 5158: 5152: 5151: 5141: 5124:(1409): 617–23. 5109: 5103: 5102: 5066: 5060: 5059: 5049: 5032:(6): 1239–1254. 5017: 5011: 5010: 5000: 4968: 4962: 4961: 4951: 4934:(23): 2594–605. 4919: 4910: 4909: 4899: 4889: 4865: 4859: 4858: 4848: 4816: 4810: 4809: 4799: 4767: 4761: 4760: 4750: 4718: 4712: 4711: 4701: 4669: 4663: 4662: 4652: 4642: 4618: 4612: 4611: 4575: 4569: 4568: 4558: 4526: 4520: 4519: 4509: 4477: 4471: 4470: 4460: 4450: 4426: 4420: 4419: 4409: 4377: 4371: 4370: 4360: 4328: 4322: 4321: 4311: 4301: 4277: 4266: 4265: 4263: 4262: 4256: 4248: 4242: 4241: 4221: 4212: 4205:T-Cell Disorders 4201: 4192: 4191: 4181: 4149: 4143: 4142: 4132: 4100: 4091: 4090: 4080: 4070: 4046: 4040: 4039: 4037: 4036: 4021: 4015: 4014: 4004: 3972: 3966: 3965: 3940:(11): 1096–105. 3929: 3923: 3922: 3904: 3895: 3894: 3884: 3852: 3846: 3843: 3837: 3836: 3826: 3794: 3788: 3787: 3743: 3737: 3736: 3692: 3686: 3685: 3657: 3651: 3640: 3634: 3633: 3600:(7426): 717–23. 3591: 3581: 3575: 3574: 3556: 3532: 3526: 3525: 3515: 3505: 3473: 3467: 3466: 3456: 3424: 3418: 3417: 3389: 3383: 3382: 3372: 3340: 3334: 3333: 3323: 3313: 3289: 3278: 3277: 3267: 3257: 3233: 3227: 3226: 3216: 3206: 3197:(51): 30204–11. 3182: 3176: 3175: 3165: 3133: 3127: 3126: 3116: 3106: 3082: 3069: 3068: 3058: 3026: 3020: 3019: 3009: 2999: 2975: 2969: 2968: 2932: 2926: 2925: 2915: 2891: 2885: 2884: 2866: 2842: 2836: 2835: 2825: 2792: 2786: 2785: 2775: 2743: 2737: 2736: 2726: 2694: 2688: 2687: 2677: 2667: 2643: 2637: 2636: 2618: 2600: 2591: 2585: 2584: 2566: 2542: 2536: 2535: 2491: 2485: 2484: 2450: 2444: 2443: 2433: 2401: 2392: 2391: 2381: 2371: 2347: 2341: 2340: 2330: 2319:10.1172/JCI31720 2298: 2292: 2291: 2263: 2257: 2256: 2238: 2232: 2231: 2221: 2188: 2182: 2181: 2147: 2138: 2132: 2131: 2113: 2089: 2083: 2082: 2054: 2048: 2047: 2037: 2027: 1995: 1989: 1988: 1970: 1964: 1963: 1953: 1921: 1915: 1914: 1904: 1895:(6): 1177–1179. 1880: 1874: 1873: 1863: 1853: 1829: 1823: 1822: 1807: 1801: 1800: 1788: 1779: 1778: 1776: 1774: 1755: 1733:Immunosenescence 1703:Immunosenescence 1605: 1585: 1584: 1577: 1437:-infected T cell 1316:and potentially 1079:, arthritis and 804:effector T cells 752:Cytotoxic T cell 607: 603: 203:(cytotoxic) and 136:edit on Wikidata 133: 43: 31: 19: 18: 6703: 6702: 6698: 6697: 6696: 6694: 6693: 6692: 6663: 6662: 6661: 6656: 6630: 6584: 6530: 6509:Clonal deletion 6437: 6431: 6361: 6262: 6250: 6220: 6215: 6183: 6148: 6097: 6069: 6063: 6055: 6047: 6039: 6015: 6007: 6000: 5964: 5942: 5885: 5880: 5850: 5843: 5832: 5828: 5822: 5803: 5801:Further reading 5798: 5797: 5742: 5738: 5729: 5727: 5718: 5717: 5713: 5668: 5664: 5632: 5628: 5597: 5593: 5540: 5536: 5489: 5485: 5448:Cancer Research 5439: 5435: 5390: 5386: 5340: 5336: 5299: 5295: 5258: 5254: 5219:Transplantation 5214: 5208: 5204: 5159: 5155: 5110: 5106: 5067: 5063: 5018: 5014: 4969: 4965: 4920: 4913: 4887:10.1186/cc10332 4866: 4862: 4817: 4813: 4768: 4764: 4719: 4715: 4678:Nature Medicine 4670: 4666: 4619: 4615: 4592:10.1038/ni.2035 4576: 4572: 4541:(15): 8161–70. 4527: 4523: 4492:(12): 8056–63. 4478: 4474: 4427: 4423: 4378: 4374: 4343:(11): 665–674. 4329: 4325: 4292:(9): e1009892. 4278: 4269: 4260: 4258: 4254: 4252:"The Lymphomas" 4250: 4249: 4245: 4238: 4222: 4215: 4202: 4195: 4150: 4146: 4101: 4094: 4047: 4043: 4034: 4032: 4022: 4018: 3973: 3969: 3930: 3926: 3919: 3905: 3898: 3853: 3849: 3844: 3840: 3795: 3791: 3754:(5617): 337–9. 3744: 3740: 3703:(6925): 852–6. 3693: 3689: 3658: 3654: 3641: 3637: 3589: 3582: 3578: 3533: 3529: 3474: 3470: 3425: 3421: 3390: 3386: 3341: 3337: 3290: 3281: 3234: 3230: 3183: 3179: 3134: 3130: 3083: 3072: 3027: 3023: 2976: 2972: 2949:10.1038/ni.3298 2933: 2929: 2892: 2888: 2864:10.1038/ni.2554 2843: 2839: 2793: 2789: 2744: 2740: 2695: 2691: 2644: 2640: 2609:(9): 5895–903. 2598: 2592: 2588: 2543: 2539: 2492: 2488: 2473: 2451: 2447: 2402: 2395: 2348: 2344: 2307:J. Clin. Invest 2299: 2295: 2264: 2260: 2253: 2239: 2235: 2189: 2185: 2162:10.1038/ni.1874 2145: 2139: 2135: 2090: 2086: 2055: 2051: 1996: 1992: 1985: 1971: 1967: 1922: 1918: 1881: 1877: 1830: 1826: 1808: 1804: 1789: 1782: 1772: 1770: 1757: 1756: 1752: 1747: 1714: 1705: 1699: 1685: 1651: 1610: 1586: 1582: 1575: 1553:Sézary syndrome 1533:T-cell lymphoma 1526: 1524:T-cell lymphoma 1520: 1447:lymphocytopenia 1428: 1422: 1417: 1404: 1300:Phosphorylated 1259:T cell receptor 1210:, and eventual 1164:T-cell receptor 1152: 1146: 1122:phosphoantigens 1095: 1089: 1051:-like protein, 1037: 1031: 1000: 994: 978: 930: 926: 922: 903: 897: 889: 879: 875: 865: 853: 849: 826: 800: 794: 770: 754: 748: 553: 537: 531: 526: 517: 515:Types of T cell 508: 500: 449:MHC complex of 446: 431:single positive 402: 394:double-positive 381: 356: 354:T-cell receptor 350: 348:TCR development 342:double positive 299:double-negative 264: 259: 234: 226: 194:immune response 180:, found in the 169:(TCR) on their 167:T-cell receptor 140: 54: 34: 17: 12: 11: 5: 6701: 6691: 6690: 6685: 6680: 6675: 6658: 6657: 6655: 6654: 6649: 6644: 6638: 6636: 6632: 6631: 6629: 6628: 6623: 6622: 6621: 6611: 6610: 6609: 6598: 6596: 6590: 6589: 6586: 6585: 6583: 6582: 6573: 6568: 6563: 6558: 6557: 6556: 6551: 6540: 6538: 6536:Immunogenetics 6532: 6531: 6529: 6528: 6523: 6518: 6517: 6516: 6511: 6506: 6501: 6496: 6484: 6483: 6481:Co-stimulation 6478: 6473: 6468: 6463: 6458: 6453: 6448: 6441: 6439: 6433: 6432: 6430: 6429: 6424: 6422:Immune complex 6418: 6417: 6412: 6407: 6402: 6397: 6396: 6395: 6390: 6385: 6380: 6369: 6367: 6363: 6362: 6360: 6359: 6354: 6349: 6344: 6342:Dendritic cell 6330: 6329: 6324: 6323: 6322: 6320:Conformational 6317: 6306: 6305: 6300: 6299: 6298: 6293: 6288: 6277: 6275: 6268: 6264: 6263: 6249: 6248: 6241: 6234: 6226: 6217: 6216: 6214: 6213: 6208: 6203: 6197: 6195: 6189: 6188: 6185: 6184: 6182: 6181: 6176: 6171: 6170: 6169: 6159: 6153: 6150: 6149: 6147: 6146: 6141: 6136: 6131: 6126: 6120: 6118: 6109: 6103: 6102: 6099: 6098: 6096: 6095: 6090: 6085: 6079: 6077: 6071: 6070: 6068: 6067: 6066: 6065: 6061: 6057: 6053: 6049: 6045: 6041: 6037: 6028: 6023: 6013: 6005: 5998: 5990: 5984: 5978: 5972: 5966: 5965: 5963: 5962: 5957: 5952: 5951: 5950: 5940: 5936: 5931: 5926: 5921: 5916: 5911: 5906: 5901: 5895: 5893: 5887: 5886: 5879: 5878: 5871: 5864: 5856: 5849: 5848: 5826: 5820: 5804: 5802: 5799: 5796: 5795: 5736: 5711: 5662: 5626: 5607:(2): 105–106. 5591: 5534: 5483: 5433: 5404:(3): 618–622. 5384: 5355:(9): 2431–42. 5334: 5313:(3): 1171–81. 5293: 5272:(4): 1838–48. 5252: 5225:(9): 1339–47. 5202: 5173:(7): 1711–20. 5153: 5104: 5061: 5012: 4963: 4911: 4860: 4831:(9): 1905–18. 4811: 4762: 4713: 4690:10.1038/nm1492 4684:(11): 1301–9. 4664: 4613: 4570: 4521: 4472: 4421: 4372: 4323: 4286:PLOS Pathogens 4267: 4243: 4236: 4213: 4193: 4158:Trends Immunol 4144: 4092: 4041: 4016: 3987:(3): 1042–52. 3967: 3924: 3917: 3896: 3847: 3838: 3815:10.1038/ni1105 3789: 3738: 3687: 3652: 3635: 3576: 3527: 3488:(20): 8290–5. 3468: 3419: 3384: 3335: 3279: 3228: 3177: 3148:(8–9): 742–8. 3128: 3070: 3041:(5): 2171–80. 3021: 2970: 2927: 2886: 2837: 2787: 2738: 2689: 2638: 2586: 2537: 2486: 2471: 2445: 2416:(7): 634–643. 2393: 2342: 2313:(5): 1119–27. 2293: 2258: 2251: 2233: 2183: 2156:(6): 512–519. 2133: 2104:(5): 627–636. 2084: 2065:(1): 139–176. 2049: 1990: 1983: 1965: 1936:(3): 619–631. 1916: 1875: 1824: 1802: 1780: 1749: 1748: 1746: 1743: 1742: 1741: 1735: 1730: 1725: 1720: 1713: 1710: 1698: 1695: 1684: 1681: 1650: 1647: 1612: 1611: 1589: 1587: 1580: 1574: 1571: 1570: 1569: 1564: 1559: 1546: 1519: 1516: 1459:Omenn syndrome 1424:Main article: 1421: 1418: 1416: 1413: 1403: 1400: 1184:co-stimulation 1145: 1142: 1091:Main article: 1088: 1085: 1033:Main article: 1030: 1027: 996:Main article: 993: 990: 977: 974: 928: 924: 920: 899:Main article: 896: 893: 892: 891: 887: 881: 877: 873: 867: 863: 851: 847: 844: 824: 796:Main article: 793: 792:Memory T cells 790: 768: 750:Main article: 747: 744: 741: 740: 737: 734: 731: 728: 724: 723: 720: 717: 714: 711: 705: 704: 701: 698: 695: 692: 686: 685: 682: 679: 676: 673: 667: 666: 663: 660: 657: 654: 648: 647: 644: 641: 636: 630: 624: 623: 620: 617: 614: 611: 564:memory B cells 551: 548:T helper cells 533:Main article: 530: 527: 525: 522: 516: 513: 507: 504: 499: 496: 445: 442: 401: 398: 380: 377: 352:Main article: 349: 346: 284:lymphoid cells 263: 260: 258: 255: 232: 229:memory B cells 224: 142: 141: 132: 126: 125: 120: 114: 113: 108: 102: 101: 96: 90: 89: 84: 78: 77: 73: 72: 67: 61: 60: 56: 55: 47:red blood cell 44: 36: 35: 32: 24: 23: 15: 9: 6: 4: 3: 2: 6700: 6689: 6686: 6684: 6683:Immune system 6681: 6679: 6676: 6674: 6671: 6670: 6668: 6653: 6650: 6648: 6645: 6643: 6640: 6639: 6637: 6633: 6627: 6624: 6620: 6617: 6616: 6615: 6612: 6608: 6605: 6604: 6603: 6600: 6599: 6597: 6595: 6591: 6581: 6577: 6574: 6572: 6569: 6567: 6564: 6562: 6559: 6555: 6552: 6550: 6547: 6546: 6545: 6542: 6541: 6539: 6537: 6533: 6527: 6524: 6522: 6519: 6515: 6512: 6510: 6507: 6505: 6504:Clonal anergy 6502: 6500: 6497: 6495: 6492: 6491: 6490: 6486: 6485: 6482: 6479: 6477: 6474: 6472: 6469: 6467: 6464: 6462: 6459: 6457: 6454: 6452: 6449: 6447: 6443: 6442: 6440: 6434: 6428: 6425: 6423: 6420: 6419: 6416: 6413: 6411: 6408: 6406: 6403: 6401: 6398: 6394: 6393:Microantibody 6391: 6389: 6386: 6384: 6381: 6379: 6376: 6375: 6374: 6371: 6370: 6368: 6364: 6358: 6355: 6353: 6350: 6348: 6345: 6343: 6339: 6335: 6332: 6331: 6328: 6325: 6321: 6318: 6316: 6313: 6312: 6311: 6308: 6307: 6304: 6301: 6297: 6294: 6292: 6289: 6287: 6284: 6283: 6282: 6279: 6278: 6276: 6272: 6269: 6265: 6261: 6257: 6254: 6247: 6242: 6240: 6235: 6233: 6228: 6227: 6224: 6212: 6211:Prolymphocyte 6209: 6207: 6204: 6202: 6199: 6198: 6196: 6194: 6193:Lymphopoiesis 6190: 6180: 6177: 6175: 6172: 6168: 6165: 6164: 6163: 6160: 6158: 6155: 6154: 6151: 6145: 6142: 6140: 6137: 6135: 6132: 6130: 6127: 6125: 6122: 6121: 6119: 6117: 6113: 6110: 6108: 6104: 6094: 6091: 6089: 6086: 6084: 6081: 6080: 6078: 6076: 6072: 6064: 6058: 6056: 6050: 6048: 6042: 6040: 6034: 6033: 6032: 6031:Memory T cell 6029: 6027: 6024: 6021: 6017: 6009: 6001: 5994: 5991: 5989: 5988:Cytotoxic CD8 5985: 5983: 5980: 5979: 5976: 5973: 5971: 5967: 5961: 5958: 5956: 5953: 5949: 5946: 5945: 5944: 5937: 5935: 5932: 5930: 5927: 5925: 5924:Marginal zone 5922: 5920: 5917: 5915: 5912: 5910: 5907: 5905: 5902: 5900: 5897: 5896: 5894: 5892: 5888: 5884: 5877: 5872: 5870: 5865: 5863: 5858: 5857: 5854: 5842: 5838: 5831: 5827: 5823: 5817: 5813: 5812: 5806: 5805: 5791: 5787: 5782: 5777: 5772: 5767: 5763: 5759: 5755: 5751: 5747: 5740: 5725: 5721: 5715: 5707: 5703: 5698: 5693: 5689: 5685: 5681: 5677: 5673: 5666: 5658: 5654: 5650: 5646: 5642: 5638: 5630: 5622: 5618: 5614: 5610: 5606: 5602: 5595: 5587: 5583: 5578: 5573: 5569: 5565: 5561: 5557: 5553: 5549: 5545: 5538: 5530: 5526: 5521: 5516: 5511: 5506: 5502: 5498: 5494: 5487: 5479: 5475: 5470: 5465: 5461: 5457: 5454:(4): 917–27. 5453: 5449: 5445: 5437: 5429: 5425: 5420: 5415: 5411: 5407: 5403: 5399: 5395: 5388: 5380: 5376: 5372: 5368: 5363: 5358: 5354: 5350: 5346: 5338: 5330: 5326: 5321: 5316: 5312: 5308: 5304: 5297: 5289: 5285: 5280: 5275: 5271: 5267: 5263: 5256: 5248: 5244: 5240: 5236: 5232: 5228: 5224: 5220: 5213: 5206: 5198: 5194: 5189: 5184: 5180: 5176: 5172: 5168: 5164: 5157: 5149: 5145: 5140: 5135: 5131: 5127: 5123: 5119: 5115: 5108: 5100: 5096: 5092: 5088: 5084: 5080: 5076: 5072: 5065: 5057: 5053: 5048: 5043: 5039: 5035: 5031: 5027: 5023: 5016: 5008: 5004: 4999: 4994: 4990: 4986: 4982: 4978: 4974: 4967: 4959: 4955: 4950: 4945: 4941: 4937: 4933: 4929: 4925: 4918: 4916: 4907: 4903: 4898: 4893: 4888: 4883: 4879: 4875: 4874:Critical Care 4871: 4864: 4856: 4852: 4847: 4842: 4838: 4834: 4830: 4826: 4822: 4815: 4807: 4803: 4798: 4793: 4789: 4785: 4782:(1): 182–96. 4781: 4777: 4773: 4766: 4758: 4754: 4749: 4744: 4740: 4736: 4733:(1): 145–57. 4732: 4728: 4724: 4717: 4709: 4705: 4700: 4695: 4691: 4687: 4683: 4679: 4675: 4668: 4660: 4656: 4651: 4646: 4641: 4636: 4632: 4628: 4624: 4617: 4609: 4605: 4601: 4597: 4593: 4589: 4585: 4581: 4574: 4566: 4562: 4557: 4552: 4548: 4544: 4540: 4536: 4532: 4525: 4517: 4513: 4508: 4503: 4499: 4495: 4491: 4487: 4483: 4476: 4468: 4464: 4459: 4454: 4449: 4444: 4440: 4436: 4435:Front Immunol 4432: 4425: 4417: 4413: 4408: 4403: 4399: 4395: 4392:(4): 474–81. 4391: 4387: 4383: 4376: 4368: 4364: 4359: 4354: 4350: 4346: 4342: 4338: 4334: 4327: 4319: 4315: 4310: 4305: 4300: 4295: 4291: 4287: 4283: 4276: 4274: 4272: 4253: 4247: 4239: 4233: 4229: 4228: 4220: 4218: 4211: 4207: 4206: 4200: 4198: 4189: 4185: 4180: 4175: 4171: 4167: 4163: 4159: 4155: 4148: 4140: 4136: 4131: 4126: 4122: 4118: 4115:(3): 619–31. 4114: 4110: 4106: 4099: 4097: 4088: 4084: 4079: 4074: 4069: 4064: 4060: 4056: 4052: 4045: 4031: 4027: 4020: 4012: 4008: 4003: 3998: 3994: 3990: 3986: 3982: 3978: 3971: 3963: 3959: 3955: 3951: 3947: 3943: 3939: 3935: 3928: 3920: 3914: 3910: 3903: 3901: 3892: 3888: 3883: 3878: 3874: 3870: 3866: 3862: 3858: 3851: 3842: 3834: 3830: 3825: 3820: 3816: 3812: 3809:(9): 927–33. 3808: 3804: 3800: 3793: 3785: 3781: 3777: 3773: 3769: 3765: 3761: 3757: 3753: 3749: 3742: 3734: 3730: 3726: 3722: 3718: 3714: 3710: 3706: 3702: 3698: 3691: 3683: 3679: 3675: 3671: 3668:(1): 171–92. 3667: 3663: 3656: 3649: 3645: 3639: 3631: 3627: 3623: 3619: 3615: 3611: 3607: 3603: 3599: 3595: 3588: 3580: 3572: 3568: 3564: 3560: 3555: 3550: 3546: 3542: 3538: 3531: 3523: 3519: 3514: 3509: 3504: 3499: 3495: 3491: 3487: 3483: 3479: 3472: 3464: 3460: 3455: 3450: 3446: 3442: 3439:(2): 266–74. 3438: 3434: 3430: 3423: 3415: 3411: 3407: 3403: 3399: 3395: 3388: 3380: 3376: 3371: 3366: 3362: 3358: 3354: 3350: 3346: 3339: 3331: 3327: 3322: 3317: 3312: 3307: 3303: 3299: 3295: 3288: 3286: 3284: 3275: 3271: 3266: 3261: 3256: 3251: 3247: 3243: 3239: 3232: 3224: 3220: 3215: 3210: 3205: 3200: 3196: 3192: 3188: 3181: 3173: 3169: 3164: 3159: 3155: 3151: 3147: 3143: 3139: 3132: 3124: 3120: 3115: 3110: 3105: 3100: 3096: 3092: 3088: 3081: 3079: 3077: 3075: 3066: 3062: 3057: 3052: 3048: 3044: 3040: 3036: 3032: 3025: 3017: 3013: 3008: 3003: 2998: 2993: 2989: 2985: 2981: 2974: 2966: 2962: 2958: 2954: 2950: 2946: 2942: 2938: 2931: 2923: 2919: 2914: 2909: 2905: 2901: 2897: 2890: 2882: 2878: 2874: 2870: 2865: 2860: 2856: 2852: 2848: 2841: 2833: 2829: 2824: 2819: 2815: 2811: 2807: 2803: 2799: 2791: 2783: 2779: 2774: 2769: 2765: 2761: 2757: 2753: 2749: 2742: 2734: 2730: 2725: 2720: 2716: 2712: 2709:(1): 165–81. 2708: 2704: 2700: 2693: 2685: 2681: 2676: 2671: 2666: 2661: 2657: 2653: 2649: 2642: 2634: 2630: 2626: 2622: 2617: 2612: 2608: 2604: 2597: 2590: 2582: 2578: 2574: 2570: 2565: 2560: 2557:(7): 2171–8. 2556: 2552: 2548: 2541: 2533: 2529: 2525: 2521: 2517: 2516:10.1038/44385 2513: 2509: 2505: 2501: 2497: 2490: 2482: 2478: 2474: 2468: 2464: 2460: 2456: 2449: 2441: 2437: 2432: 2427: 2423: 2419: 2415: 2411: 2407: 2400: 2398: 2389: 2385: 2380: 2375: 2370: 2365: 2361: 2357: 2353: 2346: 2338: 2334: 2329: 2324: 2320: 2316: 2312: 2308: 2304: 2297: 2289: 2285: 2281: 2277: 2273: 2269: 2262: 2254: 2252:9780815342434 2248: 2244: 2237: 2229: 2225: 2220: 2215: 2211: 2207: 2203: 2199: 2195: 2187: 2179: 2175: 2171: 2167: 2163: 2159: 2155: 2151: 2144: 2137: 2129: 2125: 2121: 2117: 2112: 2107: 2103: 2099: 2095: 2088: 2080: 2076: 2072: 2068: 2064: 2060: 2053: 2045: 2041: 2036: 2031: 2026: 2021: 2017: 2013: 2009: 2005: 2001: 1994: 1986: 1984:9780815342434 1980: 1976: 1969: 1961: 1957: 1952: 1947: 1943: 1939: 1935: 1931: 1927: 1920: 1912: 1908: 1903: 1898: 1894: 1890: 1886: 1879: 1871: 1867: 1862: 1857: 1852: 1847: 1843: 1839: 1835: 1828: 1821: 1817: 1813: 1806: 1798: 1794: 1787: 1785: 1768: 1764: 1760: 1754: 1750: 1739: 1736: 1734: 1731: 1729: 1726: 1724: 1721: 1719: 1716: 1715: 1709: 1704: 1697:During cancer 1694: 1691: 1680: 1677: 1673: 1669: 1665: 1661: 1657: 1646: 1644: 1640: 1636: 1632: 1628: 1624: 1618: 1608: 1603: 1599: 1598: 1593: 1588: 1579: 1578: 1568: 1565: 1563: 1560: 1558: 1554: 1550: 1547: 1545: 1542: 1541: 1540: 1538: 1534: 1530: 1525: 1515: 1513: 1509: 1508: 1503: 1502: 1501:Mycobacterium 1497: 1496: 1491: 1486: 1484: 1480: 1476: 1472: 1468: 1464: 1460: 1456: 1452: 1448: 1444: 1436: 1432: 1427: 1412: 1409: 1399: 1397: 1392: 1390: 1386: 1382: 1377: 1373: 1371: 1367: 1363: 1362:NF-κB pathway 1358: 1356: 1352: 1348: 1343: 1339: 1335: 1331: 1327: 1323: 1319: 1315: 1311: 1307: 1303: 1298: 1296: 1292: 1288: 1284: 1280: 1276: 1272: 1268: 1264: 1260: 1255: 1253: 1249: 1245: 1241: 1237: 1233: 1229: 1223: 1221: 1216: 1213: 1209: 1205: 1201: 1197: 1193: 1189: 1185: 1181: 1177: 1173: 1169: 1165: 1156: 1151: 1141: 1139: 1135: 1131: 1127: 1123: 1119: 1115: 1111: 1107: 1103: 1099: 1094: 1084: 1082: 1078: 1074: 1070: 1066: 1062: 1058: 1054: 1050: 1046: 1042: 1036: 1026: 1024: 1020: 1016: 1012: 1008: 1004: 999: 989: 986: 982: 973: 971: 967: 963: 959: 954: 952: 949: 945: 941: 938: 932: 917: 915: 911: 907: 902: 885: 882: 871: 868: 861: 857: 845: 842: 838: 834: 830: 822: 821: 820: 817: 815: 810: 809:memory T cell 805: 799: 798:Memory T cell 789: 786: 782: 778: 774: 766: 758: 753: 738: 735: 732: 729: 726: 725: 721: 718: 715: 712: 710: 707: 706: 702: 699: 696: 693: 691: 688: 687: 683: 680: 677: 674: 672: 669: 668: 664: 661: 658: 655: 653: 650: 649: 645: 642: 640: 637: 634: 631: 629: 626: 625: 621: 618: 615: 612: 609: 608: 602: 600: 596: 592: 588: 585: 581: 577: 573: 569: 565: 561: 557: 549: 541: 536: 535:T helper cell 521: 512: 506:Thymic output 503: 495: 493: 489: 488:naive T cells 485: 481: 476: 472: 468: 464: 460: 456: 452: 441: 439: 434: 432: 427: 423: 418: 415: 411: 407: 406:thymic cortex 397: 395: 390: 386: 376: 374: 370: 364: 362: 355: 345: 343: 339: 335: 331: 327: 323: 318: 316: 312: 308: 304: 300: 295: 293: 289: 285: 281: 277: 273: 269: 254: 252: 248: 244: 240: 236: 230: 222: 218: 214: 210: 206: 205:CD4+ "helper" 202: 201:CD8+ "killer" 197: 195: 191: 187: 183: 179: 174: 172: 168: 164: 160: 156: 155:immune system 152: 148: 137: 131: 127: 124: 121: 119: 115: 112: 109: 107: 103: 100: 97: 95: 91: 88: 87:lymphocytus T 85: 83: 79: 74: 71: 70:Immune system 68: 66: 62: 57: 52: 48: 42: 37: 30: 25: 20: 6606: 6471:Inflammation 6456:Alloimmunity 6451:Autoimmunity 6436:Immunity vs. 6388:Autoantibody 6286:Superantigen 5969: 5841:the original 5836: 5810: 5753: 5749: 5739: 5728:. Retrieved 5726:. 2018-08-21 5723: 5714: 5679: 5675: 5665: 5640: 5636: 5629: 5604: 5600: 5594: 5551: 5547: 5537: 5500: 5496: 5486: 5451: 5447: 5436: 5401: 5397: 5387: 5352: 5348: 5337: 5310: 5306: 5296: 5269: 5265: 5255: 5222: 5218: 5205: 5170: 5166: 5156: 5121: 5117: 5107: 5074: 5070: 5064: 5029: 5025: 5015: 4980: 4976: 4966: 4931: 4927: 4877: 4873: 4863: 4828: 4824: 4814: 4779: 4775: 4765: 4730: 4726: 4716: 4681: 4677: 4667: 4630: 4626: 4616: 4586:(6): 492–9. 4583: 4579: 4573: 4538: 4534: 4524: 4489: 4485: 4475: 4438: 4434: 4424: 4389: 4385: 4375: 4340: 4336: 4326: 4289: 4285: 4259:. Retrieved 4246: 4226: 4203: 4164:(4): 153–8. 4161: 4157: 4147: 4112: 4109:Mol. Immunol 4108: 4058: 4054: 4044: 4033:. Retrieved 4029: 4019: 3984: 3980: 3970: 3937: 3933: 3927: 3908: 3864: 3860: 3850: 3841: 3806: 3802: 3792: 3751: 3747: 3741: 3700: 3696: 3690: 3665: 3661: 3655: 3638: 3597: 3593: 3579: 3547:(8): 767–9. 3544: 3540: 3530: 3485: 3481: 3471: 3436: 3432: 3422: 3397: 3393: 3387: 3352: 3348: 3338: 3301: 3297: 3245: 3241: 3231: 3194: 3190: 3180: 3145: 3141: 3131: 3094: 3090: 3038: 3034: 3024: 2987: 2983: 2973: 2940: 2936: 2930: 2906:(5): 591–4. 2903: 2899: 2889: 2857:(4): 307–8. 2854: 2851:Nat. Immunol 2850: 2840: 2805: 2801: 2790: 2758:(2): 50–56. 2755: 2751: 2741: 2706: 2702: 2692: 2655: 2651: 2641: 2606: 2602: 2589: 2554: 2550: 2540: 2499: 2495: 2489: 2454: 2448: 2413: 2409: 2359: 2355: 2345: 2310: 2306: 2296: 2271: 2267: 2261: 2242: 2236: 2201: 2197: 2186: 2153: 2149: 2136: 2101: 2097: 2087: 2062: 2058: 2052: 2007: 2003: 1993: 1974: 1968: 1933: 1929: 1919: 1892: 1888: 1878: 1841: 1837: 1827: 1819: 1815: 1805: 1796: 1771:. Retrieved 1767:the original 1762: 1753: 1706: 1686: 1652: 1619: 1615: 1606: 1595: 1527: 1505: 1499: 1493: 1492:, including 1487: 1440: 1405: 1393: 1389:SpaceX CRS-3 1378: 1374: 1359: 1299: 1266: 1262: 1256: 1224: 1217: 1161: 1096: 1038: 1001: 984: 980: 979: 968:. Recently, 955: 943: 933: 918: 904: 831:(CCR7), and 818: 801: 763: 591:MHC class II 575: 560:plasma cells 546: 518: 509: 501: 474: 463:phagocytosed 458: 447: 438:autoimmunity 435: 430: 425: 421: 419: 403: 393: 382: 365: 357: 341: 319: 314: 311:downregulate 298: 296: 265: 247:immune cells 237: 198: 175: 171:cell surface 146: 145: 86: 6678:Human cells 6594:Lymphocytes 6253:Lymphocytic 6206:Lymphoblast 5904:Plasmablast 5883:Lymphocytes 5682:(7): e330. 4880:(4): R183. 3867:(6): 1–13. 3861:J Autoimmun 3355:(1): 1–12. 2990:: 6633824. 2274:: 529–560. 2198:JCI Insight 1773:21 December 1728:Immunoblast 1347:calcineurin 1230:-bodies or 1049:MHC class I 850:cells and T 841:lymph nodes 785:MHC class I 781:amino acids 713:IL-21, IL-4 576:CD4 T cells 572:macrophages 257:Development 182:bone marrow 163:lymphocytes 76:Identifiers 6688:Immunology 6667:Categories 6635:Substances 6499:Peripheral 6487:Inaction: 6366:Antibodies 6347:Macrophage 6260:complement 6020:Regulatory 5993:Helper CD4 5919:Follicular 5730:2018-09-22 5554:(1): 948. 5503:(1): 116. 4386:Immunology 4261:2008-04-07 4035:2014-04-14 3349:Immunology 2551:J. Immunol 1844:: 925135. 1745:References 1701:See also: 1607:(May 2023) 1573:Exhaustion 1441:Causes of 1420:Deficiency 1332:and AP-1. 1148:See also: 1144:Activation 1118:isoprenoid 1075:, such as 970:Th17 cells 856:L-selectin 833:L-selectin 773:transplant 697:IRF4, PU.1 292:thymocytes 251:autoimmune 245:, whereby 49:(left), a 6652:Cytolysin 6642:Cytokines 6489:Tolerance 6438:tolerance 6357:Immunogen 6179:LTi cells 6134:Null cell 5982:Thymocyte 5643:: 32–40. 5077:: 78–84. 4983:: 33–39. 4633:(1): 50. 4210:eMedicine 3934:BioEssays 2808:: 76–88. 1481:(AT) and 1287:tyrosines 1200:PH domain 1061:cytokines 1057:vitamin B 1013:with the 610:Cell type 599:cytokines 480:apoptotic 361:pathogens 243:tolerance 221:cytokines 217:cytotoxic 6602:Cellular 6446:Immunity 6444:Action: 6427:Paratope 6415:Idiotype 6405:Allotype 6373:Antibody 6327:Mimotope 6291:Allergen 6274:Antigens 6267:Lymphoid 6167:Nuocytes 6116:NK cells 5948:B10 cell 5790:21576466 5706:26230954 5657:29477343 5621:22189042 5586:29507342 5529:27809856 5478:22186141 5428:29197679 5371:25943855 5329:26963287 5288:26170387 5247:33409478 5239:18475193 5197:25377077 5148:11375065 5099:21657071 5091:29689313 5056:27671246 5007:27993215 4958:22187279 4906:21798063 4855:25113973 4806:23947355 4757:19604493 4727:Immunity 4708:17041596 4659:29914540 4608:11052693 4600:21739672 4565:22623779 4467:29483916 4416:20201977 4367:31570879 4318:34555119 4188:24636916 4139:17825415 4087:26471060 4011:21045195 3962:28521713 3954:14579250 3891:25578468 3833:15300249 3784:38040377 3776:12690201 3725:12594515 3682:17129182 3622:23051753 3571:27717815 3563:20733595 3522:19416870 3463:24450998 3414:28119072 3379:26778581 3330:26136743 3274:25339949 3223:26468291 3172:21458588 3123:26217338 3065:17353286 3016:33506055 2965:30992456 2957:26482978 2922:24434314 2881:11294516 2873:23507634 2832:27894837 2782:21288770 2733:23947354 2684:18657274 2658:(6): 6. 2633:16412760 2625:16237082 2581:22340282 2524:10537110 2481:25261209 2440:30867582 2388:32973809 2362:: 2025. 2337:17476341 2288:10837068 2228:28814669 2178:33154019 2170:20431619 2128:15983629 2079:12414722 2044:24927565 1960:17325202 1911:28002722 1889:Immunity 1870:22474485 1712:See also 1510:. Also, 1507:Listeria 1457:(SCID), 1453:such as 1445:include 1228:necrotic 1069:adaptive 966:TGF-beta 812:express 587:antigens 475:class II 422:class II 410:antigens 373:antigens 288:engraft: 51:platelet 6673:T cells 6647:Opsonin 6626:NK cell 6614:Humoral 6494:Central 6461:Allergy 6410:Isotype 6310:Epitope 6281:Antigen 5970:T cells 5899:B1 cell 5891:B cells 5781:3107287 5758:Bibcode 5697:4526784 5577:5838096 5556:Bibcode 5520:5093947 5469:3288154 5419:5826878 5379:5348557 5188:4483591 5139:1088449 5047:5110001 4998:5535083 4949:3361243 4897:3387626 4846:4144726 4797:3748387 4748:3039716 4699:2535582 4650:6006750 4556:3421680 4516:7966595 4458:5816053 4441:: 219. 4407:2842494 4358:7286441 4309:8460019 4179:3989030 4130:2131735 4078:4608155 4002:3035066 3882:4340844 3824:2776074 3756:Bibcode 3748:Science 3705:Bibcode 3630:4419703 3602:Bibcode 3513:2688861 3490:Bibcode 3454:3992039 3400:: 1–7. 3370:4819138 3321:4468870 3304:: 303. 3265:4189401 3248:: 450. 3214:4683245 3163:3130845 3114:4492155 3097:: 344. 3056:1865739 3007:7808823 2913:3928692 2823:6066671 2773:3039080 2724:3748618 2675:2515281 2573:2965180 2532:4378970 2504:Bibcode 2431:6804569 2379:7461801 2328:1857272 2219:5621870 2120:9054502 2035:4078834 2012:Bibcode 1951:2137912 1861:3312336 1690:NK cell 1658:(PD1), 1602:Discuss 1485:(WAS). 1473:(DGS), 1387:on the 1342:calcium 1252:anergic 1215:cells. 1176:peptide 1112:. 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Index



red blood cell
platelet
System
Immune system
Latin
MeSH
D013601
TH
H2.00.04.1.02007
FMA
62870
Anatomical terms of microanatomy
edit on Wikidata
white blood cells
immune system
adaptive immune response
lymphocytes
T-cell receptor
cell surface
hematopoietic stem cells
bone marrow
thymus
thymus
immune response
CD8+ "killer"
CD4+ "helper"
CD8
CD4

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