T helper cell - Knowledge (XXG)

368:. Typically, the APC responsible is a dendritic cell. If the antigen expresses appropriate molecular patterns (sometimes known as signal 0), it can induce maturation of the dendritic cell which results in enhanced expression of costimulatory molecules needed to activate T cells (see signal 2) and MHC Class II. Once at the lymph nodes, the APCs begin to present antigen peptides that are bound to Class II MHC, allowing CD4 T cells that express the specific TCRs against the peptide/MHC complex to activate. 332:). MHC Class II binding pockets are flexible with respect to the length of the peptides they hold. Generally, there are 9 core amino acid residues with several flanking amino acids which form a length of about 12–16 amino acids total but have been known to hold as many as 25 amino acids. By comparison, MHC Class I proteins are usually 9-10 peptides long. The activation of naive T cells is commonly explained in terms of the 3-signal model, elaborated upon below. 204: 387:, a co-receptor of the TCR complex, also binds to a different section of the MHC molecule. It is estimated that approximately 50 of these interactions are required for the activation of a helper T cell and assemblies known as microclusters have been observed forming between the TCR-CD3-CD4 complexes of the T cell and the MHC Class II proteins of the dendritic cell at the zone of contact. When these all come together, the CD4 is able to recruit a kinase called 38: 341: 779: 411:. CD45 activates Lck by dephosphorylating a tyrosine in its C-terminal tail, while Csk phosphorylates Lck at that site. The loss of CD45 produces a form of SCID because failure to activate Lck prevents appropriate T cell signaling. Memory T cells also make use of this pathway and have higher levels of Lck expressed and the function of Csk is inhibited in these cells. 485:. As naïve CD8 T cells have no true bias towards foreign sources, these T cells must rely on the activation of CD28 for confirmation that they recognize a foreign antigen (as CD80/CD86 is only expressed by active APC's). CD28 plays an important role in decreasing the risk of T cell auto-immunity against host antigens. 641:) can be essential for a successful outcome from infection. In order to be effective, helper T cells must determine which cytokines will allow the immune system to be most useful or beneficial for the host. Understanding exactly how helper T cells respond to immune challenges is currently of major interest in 1202:(TGF-β) and IL-10. Both cytokines are inhibitory to helper T cells; TGF-β suppresses the activity of most of the immune system. There is evidence to suggest that TGF-β may not suppress activated Th2 cells as effectively as it might suppress naive cells, but it is not typically considered a Th2 cytokine. 1579:

that would normally elicit a helper T cell response to bypass the immune system. While these complete bypass situations only occur when the helper T cell response is absolutely necessary for infection clearance, most infections increase in severity and/or duration because the immune system's helper T

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that could potentially be detected. The depletion of CD4 T cells and the development of chronic inflammation are signature processes in HIV pathogenesis that propel progression to acquired immune deficiency syndrome (AIDS). CD4 T cell depleted to the cell count of less than 200cell/μL in blood during

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One major difference between regulatory T cells and effector T cells is that regulatory T cells typically serve to modulate and deactivate the immune response, while effector T cell groups usually begin with immune-promoting cytokines and then switch to inhibitory cytokines later in their life cycle.

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activation (CD45RA to CD45RO), but whether this change in length influences activation is unknown. It has been proposed that the larger CD45RA may decrease the accessibility of the T cell receptor for the antigen-MHC molecule, thereby necessitating an increase in the affinity (and specificity) of the

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It has been proposed that during the non-symptomatic phase of HIV infection, the virus has a relatively low affinity towards T cells (and has a higher affinity for macrophages), resulting in a slow kill rate of CD4 T cells by the immune system. This is initially compensated for via the production of

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Considering the diverse and important role helper T cells play in the immune system, it is not surprising that these cells often influence the immune response against disease. They also occasionally generate non-beneficial responses. Very rarely, the helper T cell response could lead to the death of

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Having received the first TCR/CD3 signal, the naïve T cell must activate a second independent biochemical pathway, known as Signal 2. This verification step is a protective measure to ensure that a T cell is responding to a foreign antigen. If this second signal is not present during initial antigen

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While we know about the types of cytokine patterns helper T cells tend to produce, we understand less about how the patterns themselves are decided. Various evidence suggests that the type of APC presenting the antigen to the T cell has a major influence on its profile. Other evidence suggests that

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Once the naïve T cell has both pathways activated, the biochemical changes induced by Signal 1 are altered, allowing the cell to activate instead of undergoing anergy. The second signal is then obsolete; only the first signal is necessary for future activation. This is also true for memory T cells,

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both involve complications from auto-immune or low affinity antibodies. In both of these reactions, T cells may play an accomplice role in generating these auto-specific antibodies, although some of these reactions under Type 2 hypersensitivity would be considered normal in a healthy immune system

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T cells are not stimulated as effectively during the AIDS stage of HIV infection, making AIDS patients very susceptible to most viruses, including HIV itself. This decline in killing of CD4 T cells results in the virus being produced for a longer period (the infected CD4 T cells are not killed as

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producing cells were initially described as a pathogenic population implicated in autoimmunity but are now thought to have their own distinct effector and regulatory functions. Of note, some evidence suggest that functional plasticity is an intrinsic capacity of T helper cells. Indeed, a study in

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Additional populations of memory T cells are now known to exist. These include tissue-resident memory T (Trm) cells and virtual memory T cells. The single unifying theme for all memory T cell subtypes is that they are long-lived and can expand quickly to large numbers of effector T cells upon

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it binds to during infection), and the immune system is overwhelmed. Studies suggest that only ~5% of the lymphoid-derived CD4 T cells targeted by HIV are permissive and become productively infected with the virus. More than 95% of the CD4 T cells that die are resting and are unable to support

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cells are not a monolithic immunological entity because they are diverse in terms of function and their interaction with partner cells. In general, mature naive T cells are stimulated by professional antigen presenting cells to acquire an effector module. These are defined by the presence of a

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of the immune system but rather have a non-hematopoietic origin, and in general lack MHC Class II, meaning they are not true professional antigen-presenting cells; however, follicular dendritic cells may acquire MHC Class II proteins via exosomes that become attached to them). T cells require

1393:, focus on suppressing mast cells or other allergic cells; T cells do not play a primary role during the actual inflammatory response. It's important to note that the numeral allocation of hypersensitivity "types" does not correlate (and is completely unrelated) to the "response" in the T 1019:

the concentration of antigen presented to the T cell during primary activation influences its choice. The presence of some cytokines (such as the ones mentioned above) will also influence the response that will eventually be generated, but our understanding is nowhere near complete.

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to apoptose virus-infected cells and to induce host as well as viral DNA fragmentation. IFN alpha/beta can suppress transcription to avoid virus replication and transmission. Overactivation of THαβ against autoantigen will cause type 2 antibody-dependent cytotoxic hypersensitivity.

455:(anergy is generated from the unprotected biochemical changes of Signal 1). Anergic cells will not respond to any antigen in the future, even if both signals are present later on. These cells are generally believed to circulate throughout the body with no value until they undergo 1498:

believes, by mistake, that a host antigen is foreign. As a result, the CD8 T cells treat the host cell presenting that antigen as infected, and go on to destroy all host cells (or in the case of transplant rejection, transplant organ) that express that antigen.

731:. They are triggered by the polarising cytokines IL-4 and IL-2, and their effector cytokines are IL-4, IL-5, IL-9, IL-10, IL-13 and IL-25. The main effector cells are eosinophils, basophils, and mast cells as well as B cells, and IL-4/IL-5 CD4 T cells. The key T 1011:

in helper T cells and IL-12 in dendritic cells and macrophages. The combined action of these two cytokines suggests that once the T cell has decided to produce these cytokines, that decision is preserved (and also encourages other T cells to do the same).

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Inhibition of CD4 T-cell expansion during HIV infection may occur due to microbial translocation in an IL-10-dependent way. Triggering PD-1 expressed on activated monocytes by its ligand PD-L1, induces IL-10 production which inhibits CD4 T-cell function.

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and cytokine release. Antibodies do not play a direct role in this allergy type. T cells play an important role in this hypersensitivity, as they activate against the stimulus itself and promote the activation of other cells; particularly macrophages via

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Historically, memory T cells were thought to belong to either the effector or central memory subtypes, each with their own distinguishing set of cell surface markers. Central memory T cells reside in the lymph nodes while effector memory T cells lack the

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The immune system must achieve a balance of sensitivity in order to respond to foreign antigens without responding to the antigens of the host itself. When the immune system responds to very low levels of antigen that it usually shouldn't respond to, a

195:, though the term has been criticized for being too reductive). The loss of function in a lineage specifying transcription factor results in the absence of the corresponding class of helper T cell which can be devastating for the health of the host. 1174:

subset of helper T cells. Terms such as "regulatory" and "suppression" have become ambiguous after the discovery that helper CD4 T cells are also capable of regulating (and suppressing) their own responses outside of dedicated regulatory T cells.

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productive infection. These cells undergo abortive infection with HIV. Cell death is triggered when the host cell detects HIV foreign DNA intermediates and initiates a suicidal death pathway in an attempt to protect the host, leading to

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generated a murine monoclonal antibody, 5c8 that inhibited contact-dependent T cell helper function in human cells which characterized the 32 kDa surface protein transiently expressed on CD4 T cells. Richard Armitage at

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Toscano MA, Bianco GA, Ilarregui JM, Croci DO, Correale J, Hernandez JD, et al. (August 2007). "Differential glycosylation of TH1, TH2 and TH-17 effector cells selectively regulates susceptibility to cell death".

235:). Of note, only a very small minority of T cells egresses from the thymus (estimates commonly range from 1–5% but some experts feel even this is generous). Maturation of RTE in SLO results in the generation of mature 2985:

Harrington LE, Hatton RD, Mangan PR, Turner H, Murphy TL, Murphy KM, Weaver CT (November 2005). "Interleukin 17-producing CD4+ effector T cells develop via a lineage distinct from the T helper type 1 and 2 lineages".

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declines significantly once helper T cell function fails. The immune system loses its ability to improve the affinity of their antibodies, and are unable to generate B cells that can produce antibody groups such as

755:, and leukotriene to cause broncho-constriction, intestinal peristalsis, gastric fluid acidification to expel helminths. IL-5 from CD4 T cells will activate eosinophils to attack helminths. IL-10 suppresses T 1610:

2 cytokines (and less interactions by helper T cells). All of these complications result in an increased susceptibility to aggressive bacterial infections, especially in areas of the body not accessible by

1606:. These effects are primarily due to the loss of any helper T cell that can interact with the B lymphocyte correctly. Another symptom of AIDS is the reduction in antibody levels due to a decrease in T 1249:

2 model is enlightening and gives insight into the functions of helper T cells, it is far too simple to define its entire role or actions. Some immunologists question the model completely, as some

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complex. The T cell receptor (TCR) consists of both constant and variable regions. The variable region determines what antigen the T cell can respond to. CD4 T cells have TCRs with an affinity for

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1 transcription factors are STAT4 and T-bet. IFN-γ secreted by CD4 T cells can activate macrophages to phagocytose and digest intracellular bacteria and protozoa. In addition, IFN-γ can activate

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At this point chronic inflammation ensues, and functional CD4 T cell levels begin to decrease, eventually to a point where the CD4 T cell population is too small to recognize the full range of

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which is an allergic reaction mediated by IgE. Allergic rhinitis, atopic dermatitis, and asthma belong to this category of overactivation . In addition to expressing different cytokines, T

692:), typically against intracellular bacteria and protozoa. They are triggered by the polarising cytokine IL-12 and their effector cytokines are IFN-γ and IL-2. The main effector cells of T 481:

Although the verification stage is necessary for the activation of naïve helper T cells, the importance of this stage is best demonstrated during the similar activation mechanism of CD8

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during child-birth is a normal immune response against child antigens). The understanding of the role of helper T cells in these responses is limited but it is generally thought that T

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they will succumb usually to either cancers or infections; the immune system finally reaches a point where it is no longer coordinated or stimulated enough to deal with the disease.

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and then itself becomes phosphorylated, wherein it orchestrates the downstream signaling required for T cell activation. Lck activation is controlled by the opposing actions of

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model has still played an important part in developing our understanding of the roles and behaviour of helper T cells and the cytokines they produce during an immune response.

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It is unknown what role the relatively bulky extracellular region of CD45 plays during cell interactions, but CD45 has various isoforms that change in size depending on the T

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Armitage RJ, Fanslow WC, Strockbine L, Sato TA, Clifford KN, Macduff BM, et al. (May 1992). "Molecular and biological characterization of a murine ligand for CD40".

3276:"CD4 virtual memory: Antigen-inexperienced T cells reside in the naïve, regulatory, and memory T cell compartments at similar frequencies, implications for autoimmunity" 1241:

Many of the cytokines in this article are also expressed by other immune cells (see individual cytokines for details), and it is becoming clear that while the original T

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subsets in humans. Human IL-10 (hIL-10) suppresses the proliferation and cytokine production of all T cells and the activity of macrophages, but continues to stimulate

3080:"Multiparameter grouping delineates heterogeneous populations of human IL-17 and/or IL-22 T-cell producers that share antigen specificities with other T-cell subsets" 3223:

Sallusto F, Lenig D, Förster R, Lipp M, Lanzavecchia A (October 1999). "Two subsets of memory T lymphocytes with distinct homing potentials and effector functions".

1506:, where both antibodies and immune cells are known to play a role in the pathology. Generally the immunology of most auto-immune diseases is not well understood. 392: 1099: 132:

for those cytokines. These cells help polarize the immune response depending on the nature of the immunological insult (for example; virus vs. extracellular

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T cell for activation. However, once the activation has occurred, CD45 shortens, allowing easier interactions and activation as an effector T helper cell.

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cells contain and release cytokines to aid other immune cells. Cytokines are small protein mediators that alter the behavior of target cells that express

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cells that express the surface protein CD4. Genetic variation in regulatory elements expressed by CD4 cells determines susceptibility to a broad class of

2828:"Interleukin-10 production by Th1 cells requires interleukin-12-induced STAT4 transcription factor and ERK MAP kinase activation by high antigen dose" 207:

T-cell dependent B-cell activation, showing TH2-cell (left) B-cell (right) and several interaction molecules self-made according to Janeway et al,

4207: 493:. Faster responses occur upon reinfection because memory T cells have already undergone confirmation and can produce effector cells much sooner. 1691: 2080:"The Length Distribution of Class I-Restricted T Cell Epitopes Is Determined by Both Peptide Supply and MHC Allele-Specific Binding Preference" 3844: 2809: 2136: 1303:

encountering their cognate antigen. By this mechanism they provide the immune system with "memory" against previously encountered pathogens.

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or IL-2R), enabling a fully functional receptor that can bind with IL-2, which in turn activates the T cell's proliferation pathways.

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2 cells differentiation. Besides, IL-4 stimulates B-cells to produce IgE antibodies, which in turn stimulate mast cells to release

3758:"Transcriptional and epigenetic control of T helper cell specification: molecular mechanisms underlying commitment and plasticity" 2393:"Identification of a novel surface protein on activated CD4+ T cells that induces contact-dependent B cell differentiation (help)" 4546: 4099: 1992:

Unanue ER, Turk V, Neefjes J (May 2016). "Variations in MHC Class II Antigen Processing and Presentation in Health and Disease".

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17 cells are especially good at fighting extracellular pathogens and fungi, particularly during mucocutaneous immunity against

807: 704:(inducible nitric oxide synthase) to produce nitric oxide free radicals to directly kill intracellular bacteria and protozoa. T 427: 244: 4541: 2029:"Major Histocompatibility Complex (MHC) Class I and MHC Class II Proteins: Conformational Plasticity in Antigen Presentation" 81: 775:

1 cells in their cell surface glycans (oligosaccharides), which makes them less susceptible to some inducers of cell death.

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There are also other types of T cells that can influence the expression and activation of helper T cells, such as natural

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Proliferating helper T cells that develop into effector T cells differentiate into two major subtypes of cells known as T

4678: 4144: 4114: 4094: 3568:"Programmed death-1-induced interleukin-10 production by monocytes impairs CD4+ T cell activation during HIV infection" 4132: 4127: 3967: 1155:

2 response in humans, but acts to prevent over-stimulation of helper T cells while still maximising the production of

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Helper T cells are capable of influencing a variety of immune cells, and the T cell response generated (including the

408: 2493:"A 39-kDa protein on activated helper T cells binds CD40 and transduces the signal for cognate activation of B cells" 4370: 304: 4673: 3837: 988:, an important cytokine associated with the Type 2 response, and thus it also acts to preserve its own response. 622: 474:(B7.2) on the professional APCs. Both CD80 and CD86 activate the CD28 receptor. These proteins are also known as 224: 1636: 1594: 1491: 1400: 953: 709: 645:, because such knowledge may be very useful in the treatment of disease and in increasing the effectiveness of 1522:(HIV) infection. HIV mainly targets lymphoid CD4 T cells, but can infect other cells that express CD4 such as 1502:

Some of this section is a simplification. Many auto-immune diseases are more complex. A well-known example is

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THαβ helper cells provide the host immunity against viruses. Their differentiation is triggered by IFN α/β or

3930: 1238:. A subsequent study furthermore showed that extensive T helper cell plasticity is also prominent in humans. 4688: 4550: 4308: 4171: 1894:

van den Broek T, Delemarre EM, Janssen WJ, Nievelstein RA, Broen JC, Tesselaar K, et al. (March 2016).

1576: 1427: 1404: 1351: 791: 764: 353: 284: 160: 129: 1546:(or T-tropic) however, it begins to infect CD4 T cells far more efficiently (likely due to a change in the 1253:

studies suggest that individual helper T cells usually do not match the specific cytokine profiles of the T

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clearance. Despite the reduced levels of CD4, COVID-19 patients with severe disease had higher levels of T

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In 1991, three groups reported discovering CD154, which is the molecular basis of T cell helper function.

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Two components of the immune system are particularly affected in AIDS, due to its CD4 T cell dependency:

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The mechanism that killer T cells use during auto-immunity is almost identical to their response against

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3 cells, which transform into a regulatory subset after its initial activation and cytokine production.

1151:, ensuring that antibody production still occurs. As such, hIL-10 is not believed to truly promote the T 945: 701: 324:

on MHC Class II (in the case of helper T cells because they express CD4) or MHC class I (in the case of

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Burren OS, Rubio García A, Javierre BM, Rainbow DB, Cairns J, Cooper NJ, et al. (September 2017).

1467:. Helper T cells are required to fuel the development of these diseases. In order to create sufficient 4653: 4648: 4536: 4469: 4380: 4304: 4077: 3894: 3830: 3442: 3385: 3232: 2504: 2449: 1591:

quickly), increasing the proliferation of the virus, and accelerating the development of the disease.

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must be produced, and this is supplied by CD4 T cells. CD4 T cells can also stimulate cells such as

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Microarray analysis of PBMC gene expression profiles after Plasmodium falciparum malarial infection

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Wieczorek M, Abualrous ET, Sticht J, Álvaro-Benito M, Stolzenberg S, Noé F, Freund C (2017-03-17).

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as well as CD8 T cells, IgG B cells, and IL-10 CD4 T cells. The key THαβ transcription factors are

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generated an antibody that bound a 39 kDa protein on murine T cells and inhibited helper function.

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proteins, a CD4 cell will aid those cells through a combination of cell to cell interactions (e.g.

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van den Broek T, Borghans JA, van Wijk F (June 2018). "The full spectrum of human naive T cells".

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by dendritic cells and macrophages, and via positive feedback, IL-12 stimulates the production of

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cloned a cDNA encoding CD154 by screening an expression library with CD40-Ig. Randolph Noelle at

4683: 4416: 4058: 4045: 3815: 3256: 3205: 3113: 3011: 2952: 2782: 2555:"Thymic and Postthymic Regulation of Naïve CD4(+) T-Cell Lineage Fates in Humans and Mice Models" 2473: 2185: 1876: 1494:. Cellular auto-immune disease occurs because the host antigen recognition systems fail, and the 1187: 1163: 361: 113: 85: 3325:"Abortive HIV infection mediates CD4 T cell depletion and inflammation in human lymphoid tissue" 2293:

Courtney AH, Shvets AA, Lu W, Griffante G, Mollenauer M, Horkova V, et al. (October 2019).

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1 immunity are macrophages as well as CD8 T cells, IgG B cells, and IFN-γ CD4 T cells. The key T

203: 2709:"Glycans in the immune system and The Altered Glycan Theory of Autoimmunity: a critical review" 1676: 383:

complex binds strongly to the peptide-MHC complex present on the surface of professional APCs.

303:(at all times). Some APCs also bind native (or unprocessed) antigens to their surface, such as 4464: 4230: 4109: 3990: 3909: 3889: 3787: 3744: 3690: 3649: 3597: 3519: 3468: 3411: 3354: 3305: 3248: 3197: 3162: 3105: 3060: 3003: 2944: 2909: 2857: 2805: 2774: 2738: 2689: 2640: 2586: 2532: 2465: 2422: 2373: 2324: 2275: 2234: 2177: 2142: 2132: 2109: 2078:

Trolle T, McMurtrey CP, Sidney J, Bardet W, Osborn SC, Kaever T, et al. (February 2016).

2060: 2009: 1974: 1925: 1868: 1833: 1798: 1769:"Master regulators or lineage-specifying? Changing views on CD4+ T cell transcription factors" 1749: 1359: 1104: 155:. CD4 T cells are generally treated as having a pre-defined role as helper T cells within the 37: 3431:"IFI16 DNA sensor is required for death of lymphoid CD4 T cells abortively infected with HIV" 1659:

admission, and CD4 cell count was the only parameter that predicted length of time for viral

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Doitsh G, Cavrois M, Lassen KG, Zepeda O, Yang Z, Santiago ML, et al. (November 2010).

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Tangye SG, Puel A (2023). "The Th17/IL-17 Axis and Host Defense Against Fungal Infections".

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Maverakis E, Kim K, Shimoda M, Gershwin ME, Patel F, Wilken R, et al. (February 2015).

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counts decline, but both CD4 and CD8 cells decline to a far greater extent. Indicating that

1612: 1603: 1599: 1587: 1480: 1456: 1431: 1333: 1206: 1074: 1037: 1032: 1008: 977: 969: 879: 856: 724: 689: 557: 482: 380: 345: 325: 192: 89: 1420:(SLE) and other auto-immune diseases of similar nature can be linked to the production of T 4479: 3974: 3773: 3618:

Huang W, Berube J, McNamara M, Saksena S, Hartman M, Arshad T, et al. (August 2020).

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Marusina AI, Ono Y, Merleev AA, Shimoda M, Ogawa H, Wang EA, et al. (February 2017).

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Larsen M, Arnaud L, Hié M, Parizot C, Dorgham K, Shoukry M, et al. (September 2011).

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exposure, the T cell presumes that it is auto-reactive. This results in the cell becoming

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Institut National de la Santé et de la Recherche Médicale (Inserm) UMR-S 945: 2596–2605.

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cell's activation and maturation status. For example, CD45 shortens in length following T

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The binding of the antigen-MHC to the TCR complex and CD4 may also help the APC and the T

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Doitsh G, Galloway NL, Geng X, Yang Z, Monroe KM, Zepeda O, et al. (January 2014).

3236: 2508: 2453: 223:, these cells (termed recent thymic emigrants (RTE)) egress from the thymus and home to 4506: 4451: 4392: 4312: 4285: 3782: 3757: 3739: 3714: 3644: 3619: 3592: 3567: 3514: 3487: 3463: 3430: 3406: 3373: 3349: 3324: 3300: 3275: 3157: 3132: 3055: 3030: 2904: 2879: 2852: 2827: 2733: 2708: 2684: 2659: 2635: 2610: 2581: 2554: 2417: 2392: 2368: 2343: 2319: 2294: 2229: 2204: 2104: 2079: 2055: 2028: 1969: 1944: 1920: 1896:"Neonatal thymectomy reveals differentiation and plasticity within human naive T cells" 1895: 1793: 1768: 1744: 1717: 1222: 1140: 1087: 1065: 1061: 1057: 1000: 973: 921: 917: 891: 581: 475: 321: 312: 172: 137: 3429:

Monroe KM, Yang Z, Johnson JR, Geng X, Doitsh G, Krogan NJ, Greene WC (January 2014).

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Saraiva M, Christensen JR, Veldhoen M, Murphy TL, Murphy KM, O'Garra A (August 2009).

1718:"Chromosome contacts in activated T cells identify autoimmune disease candidate genes" 778: 4637: 4474: 4363: 4181: 4163: 4001: 3884: 2956: 2527: 2492: 1495: 1472: 1409: 1285: 1269: 1004: 992: 985: 913: 909: 905: 901: 887: 819: 815: 634: 609: 553: 518: 510: 264: 156: 3566:

Said EA, Dupuy FP, Trautmann L, Zhang Y, Shi Y, El-Far M, et al. (April 2010).

3488:"NLRP3 inflammasome induces CD4+ T cell loss in chronically HIV-1-infected patients" 3117: 3029:

Hirota K, Duarte JH, Veldhoen M, Hornsby E, Li Y, Cua DJ, et al. (March 2011).

3015: 2786: 2205:"Molecular control of steady-state dendritic cell maturation and immune homeostasis" 2189: 340: 4441: 4426: 4421: 4358: 4256: 3996: 3899: 3260: 3209: 2477: 1880: 1556: 1468: 1443: 1416:

2 cytokines would promote such disorders. For example, studies have suggested that

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Noelle RJ, Roy M, Shepherd DM, Stamenkovic I, Ledbetter JA, Aruffo A (July 1992).

2895: 2843: 2626: 2270: 2253: 1483:, encouraging these cytotoxic cells to kill host cells in certain circumstances. 4176: 3879: 3874: 1655:

attacks the CD4 cells during infection. Low CD4 predicted greater likelihood of

1547: 1539: 1523: 1439: 1435: 1390: 1371: 935: 646: 597: 506: 357: 292: 283:. Class II MHC proteins are generally only found on the surface of professional 232: 101: 3730: 3685: 3668: 3486:

Zhang C, Song JW, Huang HH, Fan X, Huang L, Deng JN, et al. (March 2021).

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Proceedings of the National Academy of Sciences of the United States of America

1945:"The ins and outs of MHC class II-mediated antigen processing and presentation" 548:

cells receiving both signals of activation and proliferation will then become T

395:(ITAMs) present on the CD3 gamma, delta, epsilon, and zeta chains. The protein 299:, although dendritic cells are the only cell group that expresses MHC Class II 4564: 4317: 4223: 3853: 3148: 2359: 2310: 2146: 1864: 1734: 1652: 1648: 1560: 1296: 1265: 1264:

Studies by Stockinger et al. revealed that another T helper subset may exist.

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lineage-determining (or lineage-specifying) transcription factor (also called

105: 97: 2675: 2095: 2045: 1490:, and some viruses have been accused of causing auto-immune diseases such as 344:

Antigen presentation stimulates naïve CD8+ and CD4+ T cells to become mature

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Lederman S, Yellin MJ, Krichevsky A, Belko J, Lee JJ, Chess L (April 1992).

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Guy B (July 2007). "The perfect mix: recent progress in adjuvant research".

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2 response during helper T cell development. Preventive treatments, such as

1367: 1299:(CD62L) receptors, which prevents them from trafficking to the lymph nodes. 1257:

model, and many cells express cytokines from both profiles. That said, the T

991:

The Type 2 response promotes its own profile using two different cytokines.

869: 752: 748: 728: 533: 529: 514: 456: 243:

that they are programmed to respond to), but naive T cells now lack or have

180: 74: 17: 3807: 3791: 3748: 3694: 3653: 3601: 3523: 3472: 3415: 3358: 3309: 3252: 3201: 3166: 3109: 3064: 3007: 2948: 2913: 2861: 2778: 2742: 2693: 2644: 2590: 2571: 2408: 2377: 2328: 2279: 2238: 2181: 2113: 2064: 2013: 1978: 1929: 1872: 1837: 1802: 1753: 430:

on the APC are the primary molecules of adhesion in this cell interaction.

279:, and CD4 is involved in determining MHC affinity during maturation in the 3100: 3079: 2536: 2469: 2426: 4612: 4397: 4385: 4343: 4297: 4261: 3918: 3869: 3635: 3374:"Cell death by pyroptosis drives CD4 T-cell depletion in HIV-1 infection" 1378: 1273: 1199: 1156: 957: 883: 638: 573: 133: 3397: 3193: 2173: 673:

2 cells (also known as Type 1 and Type 2 helper T cells, respectively).

4617: 4431: 4280: 4251: 4137: 1567: 1487: 1337: 949: 790:

2 Model for helper T cells. An antigen is ingested and processed by an

544:'s via the IL-2R thus driving proliferation and clonal expansion. The T 296: 240: 164: 3504: 3180:

Tato CM, Cua DJ (December 2008). "Alternative lifestyles of T cells".

1090:. Their key effector cytokine is IL-10. Their main effector cells are 942:

T cells. Also promotes the production of IgG, an opsonizing antibody.

4589: 4577: 4322: 4273: 3940: 3904: 3861: 2461: 1911: 1640: 1535: 1363: 1355: 1098:

and STAT3 as well as IRFs. IL-10 from CD4 T cells activate NK cells'

861: 509:. It achieves this by releasing a potent T cell growth factor called 452: 280: 228: 220: 216: 78: 66: 3583: 3046: 1960: 1784: 3031:"Fate mapping of IL-17-producing T cells in inflammatory responses" 2999: 2770: 1135:

2 model can be more complicated in some animals. For example, the T

30:"Th2" redirects here. For the professional wrestling tag team, see 3244: 1095: 984:

1 profile. IFNγ also inhibits the production of cytokines such as

777: 740: 736: 618: 517:

fashion. Activated T cells also produce the alpha sub-unit of the

423: 404: 396: 202: 176: 36: 1336:

response occurs. Hypersensitivity is believed to be the cause of

247:(reduced) expression of the RTE-related surface markers, such as 3620:"Lymphocyte Subset Counts in COVID-19 Patients: A Meta-Analysis" 1816:

Fink PJ (2013-03-21). "The biology of recent thymic emigrants".

1434:, are caused via the over-stimulation of immune cells, commonly 522: 471: 467: 463: 252: 248: 4196: 3826: 501:

Once the two-signal activation is complete the T helper cell (T

3822: 3756:

Kanno Y, Vahedi G, Hirahara K, Singleton K, O'Shea JJ (2012).

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are a subset of T helper cells developmentally distinct from T

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2 cytokines (including itself; it is auto-regulatory), while

716:

and Type 1 diabetes belong to this category of autoimmunity.

287:(APCs). Professional antigen-presenting cells are primarily 3131:

Nakayamada S, Takahashi H, Kanno Y, O'Shea JJ (June 2012).

1347:

Hypersensitivity reactions can be divided into four types:

759:

1 cells differentiation and function of dendritic cells. T

600:

and carry out many other functions of the immune response.

73:. They aid the activity of other immune cells by releasing 3715:"Dissecting How CD4 T Cells Are Lost During HIV Infection" 798:

0, is a T helper cell. The fragment is presented to it by

375:

cell encounters and recognizes the antigen on an APC, the

1619:

If the patient does not respond to (or does not receive)

960:

production (IgG, IgM and IgA as well as IgE antibodies).

1205:

The novel characterisation of another T helper subtype,

794:. It presents fragments from it to T cells. The upper, T 360:

antigens (typically bacteria or viruses), which undergo

2929:

Journal of Allergy and Clinical Immunology: In Practice

1563:(a highly inflammatory form of programmed cell death). 1166:, along with less common cytokine profiles such as the 2880:"Th17 cytokines in mucosal immunity and inflammation" 2660:"GATA3: a master of many trades in immune regulation" 2553:

Belizário JE, Brandão W, Rossato C, Peron JP (2016).

2295:"CD45 functions as a signaling gatekeeper in T cells" 995:

acts on helper T cells to promote the production of T

2873: 2871: 727:, typically against extracellular parasites such as 88:

in dendritic cells, in the activation and growth of

4605: 4563: 4505: 4406: 4336: 4244: 4237: 4162: 4085: 4076: 4044: 3939: 3860: 1697:

List of distinct cell types in the adult human body

239:(naïve meaning they have never been exposed to the 41:

Activation of macrophage or B cell by T helper cell

1060:(IL-17), a pro-inflammatory substance, as well as 1003:(IL-10) inhibits a variety of cytokines including 708:1 overactivation against autoantigens will cause 462:The second signal involves an interaction between 1667:1 CD4 cells than patients with moderate disease. 3669:"COVID-19 poses a riddle for the immune system" 3613: 3611: 2548: 2546: 1272:)–producing T cell subset focused on defending 393:immunoreceptor tyrosine-based activation motifs 2344:"Apoptosis: a review of programmed cell death" 1580:cells provide less efficient immune response. 1514:Perhaps the best example of the importance of 743:. IL-4 is the positive feedback cytokine for T 4208: 3838: 2800:Rang HP, Dale MM, Riter JM, Moore PK (2003). 1127:The interactions between cytokines from the T 629:Determination of the effector T cell response 364:, then travel from the infection site to the 8: 1575:to escape T cell recognition, thus allowing 763:2 overactivation against antigen will cause 712:or delayed-type hypersensitivity reaction. 588:1 cell production. Conversely, IL-4 drives T 399:can bind these phosphorylated ITAMs via its 2611:"CD4 T cells: fates, functions, and faults" 1767:Oestreich KJ, Weinmann AS (November 2012). 894:production has been reported in activated T 4241: 4215: 4201: 4193: 4082: 3945: 3845: 3831: 3823: 2604: 2602: 2600: 1455:Other cellular hypersensitivities include 980:in helper T cells, thereby promoting the T 422:cell activation, but the integrin protein 3781: 3738: 3684: 3643: 3591: 3513: 3503: 3462: 3405: 3348: 3299: 3156: 3099: 3054: 2974:(Ph.D. thesis). Johns Hopkins University. 2903: 2851: 2732: 2683: 2634: 2580: 2570: 2526: 2516: 2416: 2367: 2318: 2269: 2228: 2103: 2054: 2044: 1968: 1919: 1792: 1743: 1733: 1530:(both groups express CD4 at low levels). 1354:includes common immune disorders such as 1213:17) has cast further doubt on the basic T 536:secretion of IL-2 can bind to that same T 3133:"Helper T cell diversity and plasticity" 934:. Maximizes the killing efficacy of the 824: 552:0 (T helper 0) cells that secrete IL-2, 339: 2254:"Why is there so much CD45 on T cells?" 1708: 1479:and macrophages via cytokines such as 1143:inhibits cytokine production of both T 592:2 cell production and IFN-γ inhibits T 564:0 cells will then differentiate into T 348:and "helper" CD4+ cells respectively . 3774:10.1146/annurev-immunol-020711-075058 3492:The Journal of Clinical Investigation 2221:10.1146/annurev-immunol-020711-074929 2006:10.1146/annurev-immunol-041015-055420 1900:The Journal of Clinical Investigation 1830:10.1146/annurev-immunol-032712-100010 952:into proliferation, to induce B-cell 354:professional antigen-presenting cells 267:. Like all T cells, they express the 7: 2804:. Edinburgh: Churchill Livingstone. 2397:The Journal of Experimental Medicine 1692:Cancer vaccine targeting CD4 T cells 680:1 helper cells lead to an increased 513:(IL-2) which acts upon itself in an 4064:Mucosal associated invariant T cell 1322:CD4+ T cells and antitumor immunity 938:and the proliferation of cytotoxic 466:on the CD4 T cell and the proteins 77:. They are considered essential in 69:that play an important role in the 3713:Doitsh G, Greene WC (March 2016). 1682:CD4 T cells and antitumor immunity 199:Activation of naive helper T cells 147:cells express the surface protein 25: 2609:Zhu J, Paul WE (September 2008). 1943:Roche PA, Furuta K (April 2015). 4100:Lymphokine-activated killer cell 320:into short fragments which form 136:vs. intracellular bacterium vs. 2884:Current Opinion in HIV and AIDS 1200:transforming growth factor-beta 956:, and to increase neutralizing 255:, Complement Receptor 1 and 2 ( 4542:Immunoglobulin class switching 3084:European Journal of Immunology 2203:Hammer GE, Ma A (2013-03-21). 1374:. These reactions all involve 311:the same type of cells as the 1: 3137:Current Opinion in Immunology 2878:Guglani L, Khader SA (2010). 2252:Zamoyska R (September 2007). 1518:T cells is demonstrated with 1432:delayed type hypersensitivity 1293:C-C chemokine receptor type 7 596:2 cells. These cytokines are 4145:Type 3 innate lymphoid cells 4133:Type 2 innate lymphoid cells 4128:Type 1 innate lymphoid cells 4115:Uterine natural killer cells 4095:Cytokine-induced killer cell 2896:10.1097/COH.0b013e328335c2f6 2844:10.1016/j.immuni.2009.05.012 2627:10.1182/blood-2008-05-078154 2271:10.1016/j.immuni.2007.08.009 2162:Nature Reviews. Microbiology 1534:new helper T cells from the 1520:human immunodeficiency virus 1463:, and a similar phenomenon; 1179:The latter is a feature of T 972:increases the production of 808:transforming growth factor β 735:2 transcription factors are 3762:Annual Review of Immunology 2209:Annual Review of Immunology 1994:Annual Review of Immunology 1818:Annual Review of Immunology 928:Immune stimulation promoted 576:environment. IFN-γ drives T 352:During an immune response, 4705: 4371:Polyclonal B cell response 3731:10.1016/j.chom.2016.02.012 3686:10.1038/d41586-020-02379-1 3341:10.1016/j.cell.2010.11.001 3292:10.1016/j.jaut.2016.11.001 2941:10.1016/j.jaip.2023.04.015 2725:10.1016/j.jaut.2014.12.002 1949:Nature Reviews. Immunology 1853:Nature Reviews. Immunology 1773:Nature Reviews. Immunology 1542:). Once the virus becomes 1319: 1283: 1268:are claimed to be an IL9 ( 1030: 771:2 cells also differ from T 497:Differentiation (signal 3) 305:follicular dendritic cells 29: 4123: 3948: 3667:Perlman S (August 2020). 3149:10.1016/j.coi.2012.01.014 2559:Mediators of Inflammation 2360:10.1080/01926230701320337 2311:10.1126/scisignal.aaw8151 1865:10.1038/s41577-018-0001-y 1735:10.1186/s13059-017-1285-0 1230:17 cells transform into T 1111:belong to this category. 723:2 helper cells lead to a 225:secondary lymphoid organs 140:vs. fungus vs. protist). 2676:10.1016/j.it.2014.04.002 2096:10.4049/jimmunol.1501721 2046:10.3389/fimmu.2017.00292 1637:coronavirus disease 2019 1595:Antibody class switching 1577:opportunistic infections 1492:Type 1 diabetes mellitus 1226:mice demonstrated that T 954:antibody class switching 623:Dartmouth Medical School 580:1 cell production while 505:) then allows itself to 489:which is one example of 476:co-stimulatory molecules 318:antigens to be processed 285:antigen-presenting cells 263:) and the production of 108:. CD4 cells are mature T 82:antibody class switching 4172:Hematopoietic stem cell 3931:Lymphoplasmacytoid cell 3719:Cell Host & Microbe 3455:10.1126/science.1243640 3280:Journal of Autoimmunity 2713:Journal of Autoimmunity 2518:10.1073/pnas.89.14.6550 2129:Janeway's immunobiology 2033:Frontiers in Immunology 1442:, resulting in chronic 1428:Type 4 hypersensitivity 1410:Rhesus factor reactions 1405:Type 3 hypersensitivity 1352:Type 1 hypersensitivity 765:Type I hypersensitivity 725:humoral immune response 161:antigen-presenting cell 159:. For example, when an 151:and are referred to as 4485:Tolerance in pregnancy 4227:adaptive immune system 2409:10.1084/jem.175.4.1091 2342:Elmore S (June 2007). 932:Cellular immune system 849:Main partner cell type 822: 682:cell-mediated response 349: 346:"cytotoxic" CD8+ cells 212: 120:Structure and function 71:adaptive immune system 42: 4520:Somatic hypermutation 4354:Polyclonal antibodies 4349:Monoclonal antibodies 4078:Innate lymphoid cells 4054:Natural killer T cell 3101:10.1002/eji.201041131 2348:Toxicologic Pathology 2084:Journal of Immunology 1538:(originally from the 1198:produce the cytokine 946:Humoral immune system 781: 572:2 cells depending on 540:cell or neighboring T 391:which phosphorylates 343: 336:Activation (signal 1) 206: 40: 4537:Junctional diversity 4305:Antigen presentation 3636:10.1002/cyto.a.24172 2664:Trends in Immunology 2658:Wan YY (June 2014). 2572:10.1155/2016/9523628 1571:AIDS allows various 1504:rheumatoid arthritis 1477:natural killer cells 1465:transplant rejection 1115:Limitations to the T 968:The Type 1 cytokine 418:cell adhere during T 265:interleukin 8 (IL-8) 92:, and in maximizing 4679:Infectious diseases 4532:V(D)J recombination 4515:Affinity maturation 4267:Antigenic variation 4046:Innate-like T cells 3926:Transitional B cell 3447:2014Sci...343..428M 3398:10.1038/nature12940 3390:2014Natur.505..509D 3237:1999Natur.401..708S 3194:10.1038/ni1208-1323 2509:1992PNAS...89.6550N 2454:1992Natur.357...80A 2174:10.1038/nrmicro1681 2131:. Garland Science. 1657:intensive care unit 1645:natural killer cell 1461:auto-immune disease 1381:, which require a T 1342:auto-immune disease 1068:. This means that T 827: 714:Tuberculin reaction 614:Columbia University 446:Survival (signal 2) 114:autoimmune diseases 27:Type of immune cell 3816:Cardiff University 1316:Antitumor immunity 1188:regulatory T cells 1164:regulatory T cells 876:Cytokines produced 826:Th1/Th2 dichotomy 825: 823: 584:and IL-4 inhibit T 426:on the T cell and 350: 213: 63:CD4-positive cells 43: 4631: 4630: 4559: 4558: 4309:professional APCs 4190: 4189: 4158: 4157: 4072: 4071: 3679:(7821): 345–346. 3624:Cytometry. Part A 3505:10.1172/JCI138861 3441:(6169): 428–432. 3384:(7484): 509–514. 3231:(6754): 708–712. 3035:Nature Immunology 2994:(11): 1123–1132. 2988:Nature Immunology 2811:978-0-443-07145-4 2759:Nature Immunology 2503:(14): 6550–6554. 2305:(604): eaaw8151. 2299:Science Signaling 2138:978-0-8153-4551-0 2127:Murphy K (2017). 1360:allergic rhinitis 1207:T helper 17 cells 1105:Myasthenia gravis 1082:THαβ helper cells 1056:17 cells produce 1016: 1015: 690:cytotoxic T cells 604:Effector function 483:cytotoxic T cells 326:cytotoxic T cells 90:cytotoxic T cells 57:), also known as 16:(Redirected from 4696: 4674:Lymphatic system 4525:Clonal selection 4497:Immune privilege 4492:Immunodeficiency 4447:Cross-reactivity 4437:Hypersensitivity 4242: 4217: 4210: 4203: 4194: 4110:Adaptive NK cell 4083: 3946: 3847: 3840: 3833: 3824: 3819: 3795: 3785: 3752: 3742: 3699: 3698: 3688: 3664: 3658: 3657: 3647: 3615: 3606: 3605: 3595: 3563: 3557: 3556: 3554: 3553: 3544:. Archived from 3534: 3528: 3527: 3517: 3507: 3483: 3477: 3476: 3466: 3426: 3420: 3419: 3409: 3369: 3363: 3362: 3352: 3320: 3314: 3313: 3303: 3271: 3265: 3264: 3220: 3214: 3213: 3177: 3171: 3170: 3160: 3128: 3122: 3121: 3103: 3075: 3069: 3068: 3058: 3026: 3020: 3019: 2982: 2976: 2975: 2967: 2961: 2960: 2935:(6): 1624–1634. 2924: 2918: 2917: 2907: 2875: 2866: 2865: 2855: 2823: 2817: 2815: 2797: 2791: 2790: 2753: 2747: 2746: 2736: 2704: 2698: 2697: 2687: 2655: 2649: 2648: 2638: 2621:(5): 1557–1569. 2606: 2595: 2594: 2584: 2574: 2550: 2541: 2540: 2530: 2520: 2488: 2482: 2481: 2462:10.1038/357080a0 2437: 2431: 2430: 2420: 2403:(4): 1091–1101. 2388: 2382: 2381: 2371: 2339: 2333: 2332: 2322: 2290: 2284: 2283: 2273: 2249: 2243: 2242: 2232: 2200: 2194: 2193: 2157: 2151: 2150: 2124: 2118: 2117: 2107: 2090:(4): 1480–1487. 2075: 2069: 2068: 2058: 2048: 2024: 2018: 2017: 1989: 1983: 1982: 1972: 1940: 1934: 1933: 1923: 1912:10.1172/JCI84997 1906:(3): 1126–1136. 1891: 1885: 1884: 1848: 1842: 1841: 1813: 1807: 1806: 1796: 1764: 1758: 1757: 1747: 1737: 1713: 1481:interferon-gamma 1471:killer T cells, 1457:cytotoxic T cell 1430:, also known as 1334:hypersensitivity 1327:Hypersensitivity 1033:T helper 17 cell 880:Interferon gamma 828: 637:signals such as 558:interferon gamma 491:learned immunity 193:master regulator 65:, are a type of 21: 4704: 4703: 4699: 4698: 4697: 4695: 4694: 4693: 4634: 4633: 4632: 4627: 4601: 4555: 4501: 4480:Clonal deletion 4408: 4402: 4332: 4233: 4221: 4191: 4186: 4154: 4119: 4068: 4040: 4034: 4026: 4018: 4010: 3986: 3978: 3971: 3935: 3913: 3856: 3851: 3806: 3803: 3798: 3755: 3712: 3708: 3706:Further reading 3703: 3702: 3666: 3665: 3661: 3617: 3616: 3609: 3584:10.1038/nm.2106 3572:Nature Medicine 3565: 3564: 3560: 3551: 3549: 3536: 3535: 3531: 3485: 3484: 3480: 3428: 3427: 3423: 3371: 3370: 3366: 3322: 3321: 3317: 3273: 3272: 3268: 3222: 3221: 3217: 3179: 3178: 3174: 3130: 3129: 3125: 3077: 3076: 3072: 3047:10.1038/ni.1993 3028: 3027: 3023: 2984: 2983: 2979: 2969: 2968: 2964: 2926: 2925: 2921: 2877: 2876: 2869: 2825: 2824: 2820: 2812: 2799: 2798: 2794: 2755: 2754: 2750: 2706: 2705: 2701: 2657: 2656: 2652: 2608: 2607: 2598: 2552: 2551: 2544: 2490: 2489: 2485: 2448:(6373): 80–82. 2439: 2438: 2434: 2390: 2389: 2385: 2341: 2340: 2336: 2292: 2291: 2287: 2251: 2250: 2246: 2202: 2201: 2197: 2159: 2158: 2154: 2139: 2126: 2125: 2121: 2077: 2076: 2072: 2026: 2025: 2021: 1991: 1990: 1986: 1961:10.1038/nri3818 1942: 1941: 1937: 1893: 1892: 1888: 1850: 1849: 1845: 1815: 1814: 1810: 1785:10.1038/nri3321 1779:(11): 799–804. 1766: 1765: 1761: 1715: 1714: 1710: 1705: 1673: 1666: 1633: 1609: 1559:, thus causing 1528:dendritic cells 1512: 1450: 1423: 1415: 1396: 1387:corticosteroids 1384: 1329: 1324: 1318: 1309: 1307:Role in disease 1288: 1282: 1260: 1256: 1248: 1244: 1233: 1229: 1221:2 model. These 1220: 1216: 1212: 1195: 1182: 1171: 1154: 1146: 1138: 1134: 1130: 1125: 1122: 1118: 1109:Graves' disease 1084: 1071: 1062:interleukins 21 1055: 1051: 1047: 1042:17 helper cells 1041: 1035: 1029: 1027:17 helper cells 1026: 998: 983: 965:Other functions 897: 843: 836: 797: 789: 785: 774: 770: 762: 758: 746: 734: 722: 707: 699: 695: 679: 672: 668: 663: 660: 656: 631: 606: 595: 591: 587: 579: 571: 567: 563: 551: 547: 543: 539: 504: 499: 448: 440: 436: 421: 417: 374: 338: 322:linear epitopes 313:dendritic cells 289:dendritic cells 269:T cell receptor 201: 189: 146: 127: 122: 111: 86:cross-tolerance 54: 35: 28: 23: 22: 15: 12: 11: 5: 4702: 4700: 4692: 4691: 4689:Cell signaling 4686: 4681: 4676: 4671: 4666: 4661: 4656: 4651: 4646: 4636: 4635: 4629: 4628: 4626: 4625: 4620: 4615: 4609: 4607: 4603: 4602: 4600: 4599: 4594: 4593: 4592: 4582: 4581: 4580: 4569: 4567: 4561: 4560: 4557: 4556: 4554: 4553: 4544: 4539: 4534: 4529: 4528: 4527: 4522: 4511: 4509: 4507:Immunogenetics 4503: 4502: 4500: 4499: 4494: 4489: 4488: 4487: 4482: 4477: 4472: 4467: 4455: 4454: 4452:Co-stimulation 4449: 4444: 4439: 4434: 4429: 4424: 4419: 4412: 4410: 4404: 4403: 4401: 4400: 4395: 4393:Immune complex 4389: 4388: 4383: 4378: 4373: 4368: 4367: 4366: 4361: 4356: 4351: 4340: 4338: 4334: 4333: 4331: 4330: 4325: 4320: 4315: 4313:Dendritic cell 4301: 4300: 4295: 4294: 4293: 4291:Conformational 4288: 4277: 4276: 4271: 4270: 4269: 4264: 4259: 4248: 4246: 4239: 4235: 4234: 4222: 4220: 4219: 4212: 4205: 4197: 4188: 4187: 4185: 4184: 4179: 4174: 4168: 4166: 4160: 4159: 4156: 4155: 4153: 4152: 4147: 4142: 4141: 4140: 4130: 4124: 4121: 4120: 4118: 4117: 4112: 4107: 4102: 4097: 4091: 4089: 4080: 4074: 4073: 4070: 4069: 4067: 4066: 4061: 4056: 4050: 4048: 4042: 4041: 4039: 4038: 4037: 4036: 4032: 4028: 4024: 4020: 4016: 4012: 4008: 3999: 3994: 3984: 3976: 3969: 3961: 3955: 3949: 3943: 3937: 3936: 3934: 3933: 3928: 3923: 3922: 3921: 3911: 3907: 3902: 3897: 3892: 3887: 3882: 3877: 3872: 3866: 3864: 3858: 3857: 3852: 3850: 3849: 3842: 3835: 3827: 3821: 3820: 3808:"T-cell Group" 3802: 3801:External links 3799: 3797: 3796: 3753: 3725:(3): 280–291. 3709: 3707: 3704: 3701: 3700: 3659: 3630:(8): 772–776. 3607: 3578:(4): 452–459. 3558: 3529: 3478: 3421: 3364: 3335:(5): 789–801. 3315: 3266: 3215: 3188:(12): 1323–5. 3172: 3143:(3): 297–302. 3123: 3070: 3041:(3): 255–263. 3021: 3000:10.1038/ni1254 2977: 2962: 2919: 2890:(2): 120–127. 2867: 2838:(2): 209–219. 2818: 2810: 2792: 2771:10.1038/ni1482 2765:(8): 825–834. 2748: 2699: 2670:(6): 233–242. 2650: 2596: 2542: 2483: 2432: 2383: 2354:(4): 495–516. 2334: 2285: 2264:(3): 421–423. 2244: 2215:(1): 743–791. 2195: 2168:(7): 505–517. 2152: 2137: 2119: 2070: 2019: 2000:(1): 265–297. 1984: 1955:(4): 203–216. 1935: 1886: 1859:(6): 363–373. 1843: 1808: 1759: 1722:Genome Biology 1707: 1706: 1704: 1701: 1700: 1699: 1694: 1689: 1684: 1679: 1672: 1669: 1664: 1632: 1629: 1617: 1616: 1607: 1592: 1555:activation in 1511: 1508: 1453: 1452: 1448: 1425: 1421: 1413: 1408:(for example, 1398: 1394: 1382: 1328: 1325: 1320:Main article: 1317: 1314: 1308: 1305: 1284:Main article: 1281: 1278: 1258: 1254: 1246: 1242: 1231: 1227: 1218: 1214: 1210: 1193: 1180: 1169: 1152: 1144: 1136: 1132: 1128: 1124: 1120: 1116: 1113: 1083: 1080: 1069: 1058:interleukin 17 1053: 1049: 1045: 1039: 1031:Main article: 1028: 1024: 1021: 1014: 1013: 1001:interleukin 10 996: 989: 981: 974:interleukin 12 966: 962: 961: 943: 929: 925: 924: 922:interleukin 13 918:interleukin 10 899: 895: 892:interleukin 10 877: 873: 872: 859: 850: 846: 845: 841: 838: 834: 831: 795: 787: 783: 772: 768: 760: 756: 744: 732: 720: 705: 697: 693: 684:(primarily by 677: 670: 666: 662: 658: 654: 651: 630: 627: 605: 602: 593: 589: 585: 577: 569: 565: 561: 560:(IFN-γ). The T 549: 545: 541: 537: 502: 498: 495: 447: 444: 438: 434: 419: 415: 372: 337: 334: 328:which express 301:constitutively 211:(Berlin, 2002) 200: 197: 187: 179:) and through 173:CD40 (protein) 144: 125: 121: 118: 109: 52: 47:T helper cells 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 4701: 4690: 4687: 4685: 4682: 4680: 4677: 4675: 4672: 4670: 4669:Immune system 4667: 4665: 4662: 4660: 4657: 4655: 4652: 4650: 4647: 4645: 4642: 4641: 4639: 4624: 4621: 4619: 4616: 4614: 4611: 4610: 4608: 4604: 4598: 4595: 4591: 4588: 4587: 4586: 4583: 4579: 4576: 4575: 4574: 4571: 4570: 4568: 4566: 4562: 4552: 4548: 4545: 4543: 4540: 4538: 4535: 4533: 4530: 4526: 4523: 4521: 4518: 4517: 4516: 4513: 4512: 4510: 4508: 4504: 4498: 4495: 4493: 4490: 4486: 4483: 4481: 4478: 4476: 4475:Clonal anergy 4473: 4471: 4468: 4466: 4463: 4462: 4461: 4457: 4456: 4453: 4450: 4448: 4445: 4443: 4440: 4438: 4435: 4433: 4430: 4428: 4425: 4423: 4420: 4418: 4414: 4413: 4411: 4405: 4399: 4396: 4394: 4391: 4390: 4387: 4384: 4382: 4379: 4377: 4374: 4372: 4369: 4365: 4364:Microantibody 4362: 4360: 4357: 4355: 4352: 4350: 4347: 4346: 4345: 4342: 4341: 4339: 4335: 4329: 4326: 4324: 4321: 4319: 4316: 4314: 4310: 4306: 4303: 4302: 4299: 4296: 4292: 4289: 4287: 4284: 4283: 4282: 4279: 4278: 4275: 4272: 4268: 4265: 4263: 4260: 4258: 4255: 4254: 4253: 4250: 4249: 4247: 4243: 4240: 4236: 4232: 4228: 4225: 4218: 4213: 4211: 4206: 4204: 4199: 4198: 4195: 4183: 4182:Prolymphocyte 4180: 4178: 4175: 4173: 4170: 4169: 4167: 4165: 4164:Lymphopoiesis 4161: 4151: 4148: 4146: 4143: 4139: 4136: 4135: 4134: 4131: 4129: 4126: 4125: 4122: 4116: 4113: 4111: 4108: 4106: 4103: 4101: 4098: 4096: 4093: 4092: 4090: 4088: 4084: 4081: 4079: 4075: 4065: 4062: 4060: 4057: 4055: 4052: 4051: 4049: 4047: 4043: 4035: 4029: 4027: 4021: 4019: 4013: 4011: 4005: 4004: 4003: 4002:Memory T cell 4000: 3998: 3995: 3992: 3988: 3980: 3972: 3965: 3962: 3960: 3959:Cytotoxic CD8 3956: 3954: 3951: 3950: 3947: 3944: 3942: 3938: 3932: 3929: 3927: 3924: 3920: 3917: 3916: 3915: 3908: 3906: 3903: 3901: 3898: 3896: 3895:Marginal zone 3893: 3891: 3888: 3886: 3883: 3881: 3878: 3876: 3873: 3871: 3868: 3867: 3865: 3863: 3859: 3855: 3848: 3843: 3841: 3836: 3834: 3829: 3828: 3825: 3817: 3813: 3809: 3805: 3804: 3800: 3793: 3789: 3784: 3779: 3775: 3771: 3767: 3763: 3759: 3754: 3750: 3746: 3741: 3736: 3732: 3728: 3724: 3720: 3716: 3711: 3710: 3705: 3696: 3692: 3687: 3682: 3678: 3674: 3670: 3663: 3660: 3655: 3651: 3646: 3641: 3637: 3633: 3629: 3625: 3621: 3614: 3612: 3608: 3603: 3599: 3594: 3589: 3585: 3581: 3577: 3573: 3569: 3562: 3559: 3548:on 2015-04-15 3547: 3543: 3539: 3533: 3530: 3525: 3521: 3516: 3511: 3506: 3501: 3497: 3493: 3489: 3482: 3479: 3474: 3470: 3465: 3460: 3456: 3452: 3448: 3444: 3440: 3436: 3432: 3425: 3422: 3417: 3413: 3408: 3403: 3399: 3395: 3391: 3387: 3383: 3379: 3375: 3368: 3365: 3360: 3356: 3351: 3346: 3342: 3338: 3334: 3330: 3326: 3319: 3316: 3311: 3307: 3302: 3297: 3293: 3289: 3285: 3281: 3277: 3270: 3267: 3262: 3258: 3254: 3250: 3246: 3245:10.1038/44385 3242: 3238: 3234: 3230: 3226: 3219: 3216: 3211: 3207: 3203: 3199: 3195: 3191: 3187: 3183: 3176: 3173: 3168: 3164: 3159: 3154: 3150: 3146: 3142: 3138: 3134: 3127: 3124: 3119: 3115: 3111: 3107: 3102: 3097: 3093: 3092:UPMC Paris 06 3089: 3085: 3081: 3074: 3071: 3066: 3062: 3057: 3052: 3048: 3044: 3040: 3036: 3032: 3025: 3022: 3017: 3013: 3009: 3005: 3001: 2997: 2993: 2989: 2981: 2978: 2973: 2970:Hu W (2007). 2966: 2963: 2958: 2954: 2950: 2946: 2942: 2938: 2934: 2930: 2923: 2920: 2915: 2911: 2906: 2901: 2897: 2893: 2889: 2885: 2881: 2874: 2872: 2868: 2863: 2859: 2854: 2849: 2845: 2841: 2837: 2833: 2829: 2822: 2819: 2813: 2807: 2803: 2796: 2793: 2788: 2784: 2780: 2776: 2772: 2768: 2764: 2760: 2752: 2749: 2744: 2740: 2735: 2730: 2726: 2722: 2718: 2714: 2710: 2703: 2700: 2695: 2691: 2686: 2681: 2677: 2673: 2669: 2665: 2661: 2654: 2651: 2646: 2642: 2637: 2632: 2628: 2624: 2620: 2616: 2612: 2605: 2603: 2601: 2597: 2592: 2588: 2583: 2578: 2573: 2568: 2564: 2560: 2556: 2549: 2547: 2543: 2538: 2534: 2529: 2524: 2519: 2514: 2510: 2506: 2502: 2498: 2494: 2487: 2484: 2479: 2475: 2471: 2467: 2463: 2459: 2455: 2451: 2447: 2443: 2436: 2433: 2428: 2424: 2419: 2414: 2410: 2406: 2402: 2398: 2394: 2387: 2384: 2379: 2375: 2370: 2365: 2361: 2357: 2353: 2349: 2345: 2338: 2335: 2330: 2326: 2321: 2316: 2312: 2308: 2304: 2300: 2296: 2289: 2286: 2281: 2277: 2272: 2267: 2263: 2259: 2255: 2248: 2245: 2240: 2236: 2231: 2226: 2222: 2218: 2214: 2210: 2206: 2199: 2196: 2191: 2187: 2183: 2179: 2175: 2171: 2167: 2163: 2156: 2153: 2148: 2144: 2140: 2134: 2130: 2123: 2120: 2115: 2111: 2106: 2101: 2097: 2093: 2089: 2085: 2081: 2074: 2071: 2066: 2062: 2057: 2052: 2047: 2042: 2038: 2034: 2030: 2023: 2020: 2015: 2011: 2007: 2003: 1999: 1995: 1988: 1985: 1980: 1976: 1971: 1966: 1962: 1958: 1954: 1950: 1946: 1939: 1936: 1931: 1927: 1922: 1917: 1913: 1909: 1905: 1901: 1897: 1890: 1887: 1882: 1878: 1874: 1870: 1866: 1862: 1858: 1854: 1847: 1844: 1839: 1835: 1831: 1827: 1823: 1819: 1812: 1809: 1804: 1800: 1795: 1790: 1786: 1782: 1778: 1774: 1770: 1763: 1760: 1755: 1751: 1746: 1741: 1736: 1731: 1727: 1723: 1719: 1712: 1709: 1702: 1698: 1695: 1693: 1690: 1688: 1685: 1683: 1680: 1678: 1677:CD4/CD8 ratio 1675: 1674: 1670: 1668: 1662: 1658: 1654: 1650: 1646: 1642: 1638: 1630: 1628: 1624: 1622: 1621:HIV treatment 1614: 1605: 1601: 1596: 1593: 1589: 1586: 1585: 1584: 1581: 1578: 1574: 1569: 1564: 1562: 1558: 1557:inflammasomes 1554: 1549: 1545: 1541: 1537: 1531: 1529: 1525: 1521: 1517: 1510:HIV infection 1509: 1507: 1505: 1500: 1497: 1496:immune system 1493: 1489: 1484: 1482: 1478: 1474: 1473:interleukin-2 1470: 1469:auto-reactive 1466: 1462: 1458: 1445: 1441: 1437: 1433: 1429: 1426: 1419: 1411: 1406: 1402: 1399: 1392: 1388: 1380: 1377: 1373: 1369: 1365: 1362:(hay fever), 1361: 1357: 1353: 1350: 1349: 1348: 1345: 1343: 1339: 1335: 1326: 1323: 1315: 1313: 1306: 1304: 1300: 1298: 1294: 1287: 1286:Memory T cell 1280:Memory T cell 1279: 1277: 1275: 1271: 1270:interleukin 9 1267: 1262: 1252: 1239: 1237: 1224: 1208: 1203: 1201: 1197: 1189: 1184: 1176: 1173: 1165: 1160: 1158: 1150: 1142: 1114: 1112: 1110: 1106: 1101: 1097: 1093: 1089: 1081: 1079: 1077: 1076: 1067: 1063: 1059: 1052:2 lineages. T 1043: 1034: 1022: 1020: 1010: 1006: 1005:interleukin 2 1002: 994: 993:Interleukin 4 990: 987: 986:interleukin 4 979: 975: 971: 967: 964: 963: 959: 955: 951: 948:. Stimulates 947: 944: 941: 937: 933: 930: 927: 926: 923: 919: 915: 914:interleukin 9 911: 910:interleukin 6 907: 906:interleukin 5 903: 902:Interleukin 4 900: 893: 889: 888:Interleukin 2 885: 881: 878: 875: 874: 871: 867: 863: 860: 858: 854: 851: 848: 847: 839: 832: 830: 829: 821: 820:interleukin 4 817: 816:interleukin 2 813: 809: 805: 801: 793: 780: 776: 766: 754: 750: 742: 738: 730: 726: 717: 715: 711: 703: 691: 687: 683: 674: 652: 650: 648: 644: 640: 636: 635:extracellular 628: 626: 624: 620: 615: 611: 610:Seth Lederman 603: 601: 599: 583: 575: 559: 555: 535: 531: 526: 524: 520: 519:IL-2 receptor 516: 512: 511:interleukin 2 508: 496: 494: 492: 486: 484: 479: 477: 473: 469: 465: 460: 458: 454: 445: 443: 431: 429: 425: 412: 410: 406: 402: 398: 394: 390: 386: 382: 378: 369: 367: 363: 359: 355: 347: 342: 335: 333: 331: 327: 323: 319: 314: 310: 306: 302: 298: 294: 290: 286: 282: 278: 274: 270: 266: 262: 258: 254: 250: 246: 245:downregulated 242: 238: 237:naive T cells 234: 230: 226: 222: 218: 210: 205: 198: 196: 194: 184: 182: 178: 174: 170: 166: 162: 158: 157:immune system 154: 150: 141: 139: 135: 131: 119: 117: 115: 107: 103: 99: 95: 91: 87: 83: 80: 76: 72: 68: 64: 60: 56: 48: 39: 33: 19: 4664:Cell biology 4442:Inflammation 4427:Alloimmunity 4422:Autoimmunity 4407:Immunity vs. 4359:Autoantibody 4257:Superantigen 3963: 3811: 3765: 3761: 3722: 3718: 3676: 3672: 3662: 3627: 3623: 3575: 3571: 3561: 3550:. Retrieved 3546:the original 3542:www.aids.gov 3541: 3532: 3495: 3491: 3481: 3438: 3434: 3424: 3381: 3377: 3367: 3332: 3328: 3318: 3286:(2): 76–88. 3283: 3279: 3269: 3228: 3224: 3218: 3185: 3181: 3175: 3140: 3136: 3126: 3087: 3083: 3073: 3038: 3034: 3024: 2991: 2987: 2980: 2971: 2965: 2932: 2928: 2922: 2887: 2883: 2835: 2831: 2821: 2802:Pharmacology 2801: 2795: 2762: 2758: 2751: 2716: 2712: 2702: 2667: 2663: 2653: 2618: 2614: 2562: 2558: 2500: 2496: 2486: 2445: 2441: 2435: 2400: 2396: 2386: 2351: 2347: 2337: 2302: 2298: 2288: 2261: 2257: 2247: 2212: 2208: 2198: 2165: 2161: 2155: 2128: 2122: 2087: 2083: 2073: 2036: 2032: 2022: 1997: 1993: 1987: 1952: 1948: 1938: 1903: 1899: 1889: 1856: 1852: 1846: 1824:(1): 31–50. 1821: 1817: 1811: 1776: 1772: 1762: 1725: 1721: 1711: 1647:, and total 1634: 1625: 1618: 1582: 1565: 1548:co-receptors 1544:lymphotropic 1532: 1513: 1501: 1485: 1454: 1451:1 cytokines. 1444:inflammation 1424:2 cytokines. 1370:(hives) and 1346: 1330: 1310: 1301: 1289: 1276:infections. 1263: 1250: 1240: 1235: 1204: 1185: 1177: 1161: 1149:plasma cells 1126: 1085: 1073: 1036: 1017: 804:interferon γ 718: 675: 664: 632: 607: 527: 500: 487: 480: 461: 449: 432: 413: 370: 351: 308: 277:Class II MHC 214: 208: 185: 169:MHC class II 152: 142: 123: 96:activity of 94:bactericidal 62: 58: 50: 46: 44: 32:The Hybrid 2 18:CD4+ T cells 4654:Macrophages 4649:Human cells 4565:Lymphocytes 4224:Lymphocytic 4177:Lymphoblast 3875:Plasmablast 3854:Lymphocytes 3768:: 707–731. 3538:"CD4 Count" 3182:Nat Immunol 2719:(6): 1–13. 2565:: 9523628. 1687:CD8 T cells 1639:(COVID-19) 1615:antibodies. 1540:bone marrow 1524:macrophages 1440:macrophages 1436:lymphocytes 1391:montelukast 1372:anaphylaxis 1295:(CCR7) and 1139:2 cytokine 936:macrophages 882:(IFNγ) and 686:macrophages 647:vaccination 598:pleiotropic 507:proliferate 366:lymph nodes 307:(these are 293:macrophages 233:lymph nodes 217:development 209:Immunologie 167:antigen on 163:displays a 153:CD4 T cells 106:neutrophils 102:macrophages 84:, breaking 4659:Phagocytes 4638:Categories 4606:Substances 4470:Peripheral 4458:Inaction: 4337:Antibodies 4318:Macrophage 4231:complement 3991:Regulatory 3964:Helper CD4 3890:Follicular 3552:2015-04-30 2147:1020120603 1728:(1): 165. 1703:References 1653:SARS-Cov-2 1649:lymphocyte 1561:pyroptosis 1379:antibodies 1312:the host. 1297:L-selectin 1157:antibodies 866:eosinophil 857:CD8 T cell 853:Macrophage 812:macrophage 643:immunology 470:(B7.1) or 401:SH2 domain 362:processing 358:endocytose 215:Following 98:phagocytes 4684:Apoptosis 4623:Cytolysin 4613:Cytokines 4460:Tolerance 4409:tolerance 4328:Immunogen 4150:LTi cells 4105:Null cell 3953:Thymocyte 2957:258380150 1573:pathogens 1553:caspase-1 1459:mediated 1368:urticaria 1340:and some 1266:Th9 cells 870:mast cell 840:Type 2/ T 833:Type 1/ T 806:; TGF-β, 802:. 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