Toll-like receptor - Knowledge (XXG)

570: 1545: 1624: 1585:; and also of the CpG islands found in the promoters of eukaryotic DNA; as well as certain other RNA and DNA molecules. As TLR ligands are present in most pathogens, they may also be present in pathogen-derived vaccines (e.g. MMR, influenza, polio vaccines) most commercially available vaccines have been assessed for their inherent TLR ligands' capacity to activate distinct subsets of immune cells. For most of the TLRs, 1910:) were shown in turn to provoke host responses that are normally protective. However, these responses can be detrimental if they are excessively prolonged or intense. It followed logically that there must be receptors for such molecules, capable of alerting the host to the presence of infection, but these remained elusive for many years. Toll-like receptors are now counted among the key molecules that alert the 2018: 42: 1615:, extracellular matrix components and self DNA (it is normally degraded by nucleases, but under inflammatory and autoimmune conditions it can form a complex with endogenous proteins, become resistant to these nucleases and gain access to endosomal TLRs as TLR7 or TLR9). These endogenous ligands are usually produced as a result of non-physiological cell death. 480: 1553:(i.e., cannot be mistaken for self molecules that are normally expressed under physiological conditions). Pathogen-associated molecules that meet this requirement are thought to be critical to the pathogen's function and difficult to change through mutation; they are said to be evolutionarily conserved. Somewhat conserved features in pathogens include 622:). Cleaved spätzle then binds to the toll receptor and crosslinks its ectodomains. This triggers conformational changes in the receptor resulting in signalling through toll. From this point forward, the signalling cascade is very similar to mammalian signalling through TLRs. The toll-induced signalling complex (TICS) is composed of 394: 1772:

TLR4 is the only TLR that uses all four adaptors. Complex consisting of TLR4, MD2 and LPS recruits TIR domain-containing adaptors TIRAP and MyD88 and thus initiates activation of NFκB (early phase) and MAPK. TLR4-MD2-LPS complex then undergoes endocytosis and in endosome it forms a signalling complex

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The "unnatural" enantiomers of opioid drugs such as (+)-morphine and (+)-naloxone lack affinity for opioid receptors, still produce the same activity at TLR4 as their "normal" enantiomers. So, "unnatural" entianomers of opioids such as (+)-naloxone, can be used to block the TLR4 activity of opioid

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Because of the specificity of toll-like receptors (and other innate immune receptors) they cannot easily be changed in the course of evolution, these receptors recognize molecules that are constantly associated with threats (i.e., pathogen or cell stress) and are highly specific to these threats

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When microbes were first recognized as the cause of infectious diseases, it was immediately clear that multicellular organisms must be capable of recognizing them when infected and, hence, capable of recognizing molecules unique to microbes. A large body of literature, spanning most of the last

586:. Fruit flies have only innate immune responses allowing studies to avoid interference of adaptive immune mechanisms on signal transduction. The fly response to fungal or bacterial infection occurs through two distinct signalling cascades, one of which is the toll pathway and the other is the 537:

It has been estimated that most mammalian species have between ten and fifteen types of toll-like receptors. Thirteen TLRs (named simply TLR1 to TLR13) have been identified in humans and mice together, and equivalent forms of many of these have been found in other mammalian species. However,

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dependent signalling pathway. To explore the role of this pathway in retroviral reprograming, knock down techniques of TLR3 or TRIF were prepared, and results showed that only the TLR3 pathway is required for full induction of target gene expression by the retrovirus expression vector. This

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knockdown of TLR3 or its adaptor protein TRIF. Taken together, stimulation of TLR3 causes great changes in chromatin remodeling and nuclear reprogramming, and activation of inflammatory pathways is required for these changes, induction of pluripotency genes and generation of human induced

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with TRAM and TRIF adaptors. This TRIF-dependent pathway again leads to IRF3 activation and production of type I interferons, but it also activates late-phase NFκB activation. Both late and early phase activation of NFκB is required for production of inflammatory cytokines.

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Toll-like receptors bind and become activated by different ligands, which, in turn, are located on different types of organisms or structures. They also have different adapters to respond to activation and are located sometimes at the cell surface and sometimes to internal

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Aleynick, Mark; Svensson-Arvelund, Judit; Pantsulaia, Gvantsa; Kim, Kristy; Rose, Samuel A.; Upadhyay, Ranjan; Yellin, Michael; Marsh, Henry; Oreper, Daniel; Jhunjhunwala, Suchit; Moussion, Christine Carine; Merad, Miriam; Brown, Brian D.; Brody, Joshua D. (July 2023).

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as an inactive dimeric precursor. The toll receptor shares the cytoplasmatic TIR domain with mammalian TLRs, but the ectodomain and intracytoplasmatic tail are different. This difference might reflect a function of these receptors as cytokine receptors rather than

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The stereotypic inflammatory response provoked by toll-like receptor activation has prompted speculation that endogenous activators of toll-like receptors might participate in autoimmune diseases. TLRs have been suspected of binding to host molecules including

546:. On the other hand, mice express TLRs 11, 12, and 13, none of which is represented in humans. Other mammals may express TLRs that are not found in humans. Other non-mammalian species may have TLRs distinct from mammals, as demonstrated by the anti-cell-wall 1856:

and hyperalgesia. Drugs that block the action of TNF-α or IL-1β have been shown to increase the analgesic effects of opioids and reduce the development of tolerance and other side-effects, and this has also been demonstrated with drugs that block TLR4 itself.

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has been used to decipher the TLR signaling pathways. When activated, TLRs recruit adapter molecules within the cytoplasm of cells to propagate a signal. Four adapter molecules are known to be involved in signaling. These proteins are known as

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Sameer AS, Nissar S. Toll-Like Receptors (TLRs): Structure, Functions, Signaling, and Role of Their Polymorphisms in Colorectal Cancer Susceptibility. Biomed Res Int. 2021 Sep 12;2021:1157023. doi: 10.1155/2021/1157023. PMID 34552981; PMCID:

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was not then known, it was assumed that TIL (now known as TLR1) might participate in mammalian development. However, in 1991 (prior to the discovery of TIL) it was observed that a molecule with a clear role in immune function in mammals, the

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recognition specificity has now been established by gene targeting (also known as "gene knockout"): a technique by which individual genes may be selectively deleted in mice. See the table above for a summary of known TLR ligands.

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TLR signaling ultimately leads to the induction or suppression of genes that orchestrate the inflammatory response. In all, thousands of genes are activated by TLR signaling, and collectively, the TLRs constitute one of the most

1729:. On binding, TAK1 phosphorylates IKK-β, which then phosphorylates IκB causing its degradation and allowing NFκB to diffuse into the cell nucleus and activate transcription and consequent induction of inflammatory cytokines. 4087:

Schreibelt, Gerty; Benitez-Ribas, Daniel; Schuurhuis, Danita; Lambeck, Annechien J. A.; van Hout-Kuijer, Maaike; Schaft, Niels; Punt, Cornelis J. A.; Figdor, Carl G.; Adema, Gosse J.; de Vries, I. Jolanda M. (29 July 2010).

5199:"Prediction of the coding sequences of unidentified human genes. I. The coding sequences of 40 new genes (KIAA0001-KIAA0040) deduced by analysis of randomly sampled cDNA clones from human immature myeloid cell line KG-1" 5640: 5238:

Taguchi T, Mitcham JL, Dower SK, Sims JE, Testa JR (March 1996). "Chromosomal localization of TIL, a gene encoding a protein related to the Drosophila transmembrane receptor Toll, to human chromosome 4p14".

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In recent years TLRs were identified also in the mammalian nervous system. Members of the TLR family were detected on glia, neurons and on neural progenitor cells in which they regulate cell-fate decision.

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in somatic cells. This is supported by study, which shows, that efficiency and amount of human iPSC generation, using retroviral vectors, is reduced by knockdown of the pathway with peptide inhibitors or

458:(Ig) domains. Proteins with subgroup 2 TIR domains are classical TLRs, and bind directly or indirectly to molecules of microbial origin. A third subgroup of proteins containing TIR domains consists of 1898:(LPS) produced by most Gram-negative bacteria. This lipopolysaccharide is an integral part of the gram-negative membrane and is released upon destruction of the bacterium. Other molecules (bacterial 4404:

Yamamoto M, Sato S, Hemmi H, Uematsu S, Hoshino K, Kaisho T, et al. (November 2003). "TRAM is specifically involved in the Toll-like receptor 4-mediated MyD88-independent signaling pathway".

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and colleagues. Each TLR is now believed to detect a discrete collection of molecules – some of microbial origin, and some products of cell damage – and to signal the presence of infections.

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The first reported human toll-like receptor was described by Nomura and colleagues in 1994, mapped to a chromosome by Taguchi and colleagues in 1996. Because the immune function of toll in

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with TLR1 or TLR6, each dimer having a different ligand specificity. TLRs may also depend on other co-receptors for full ligand sensitivity, such as in the case of TLR4's recognition of

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Oldenburg M, Krüger A, Ferstl R, Kaufmann A, Nees G, Sigmund A, et al. (August 2012). "TLR13 recognizes bacterial 23S rRNA devoid of erythromycin resistance-forming modification".

6475: 5107:"Possible involvement of toll-like receptor 4/myeloid differentiation factor-2 activity of opioid inactive isomers causes spinal proinflammation and related behavioral consequences" 5056:

Hutchinson MR, Coats BD, Lewis SS, Zhang Y, Sprunger DB, Rezvani N, Baker EM, Jekich BM, Wieseler JL, Somogyi AA, Martin D, Poole S, Judd CM, Maier SF, Watkins LR (November 2008).

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Yamamoto M, Sato S, Hemmi H, Sanjo H, Uematsu S, Kaisho T, et al. (November 2002). "Essential role for TIRAP in activation of the signalling cascade shared by TLR2 and TLR4".

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In 2011, Beutler and Hoffmann were awarded the Nobel Prize in Medicine or Physiology for their work. Hoffmann and Akira received the Canada Gairdner International Award in 2011.

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Seizer L, Rahimi S, Santos-Sierra S, Drexel M (2022) Expression of toll like receptor 8 (TLR8) in specific groups of mouse hippocampal interneurons. PLoS ONE 17(5): e0267860.

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to prove that mice that could not respond to LPS had mutations that abolished the function of TLR4. This identified TLR4 as one of the key components of the receptor for LPS.

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Hashimoto C, Hudson KL, Anderson KV (January 1988). "The Toll gene of Drosophila, required for dorsal-ventral embryonic polarity, appears to encode a transmembrane protein".

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Poltorak A, He X, Smirnova I, Liu MY, Van Huffel C, Du X, et al. (December 1998). "Defective LPS signaling in C3H/HeJ and C57BL/10ScCr mice: mutations in Tlr4 gene".

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Peana M, Zdyb K, Medici S, Pelucelli A, Simula G, Gumienna-Kontecka E, Zoroddu MA (December 2017). "Ni(II) interaction with a peptide model of the human TLR4 ectodomain".

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showed that a toll-like receptor now known as TLR4 could, when artificially ligated using antibodies, induce the activation of certain genes necessary for initiating an

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Ferrandon D, Imler JL, Hetru C, Hoffmann JA (November 2007). "The Drosophila systemic immune response: sensing and signalling during bacterial and fungal infections".

1792:, is a TLR7 and TLR8 agonist. Recently, resiquimod has been explored as an agent for cancer immunotherapy, acting through stimulation of tumor-associated macrophages. 3631:"Quantitative expression of toll-like receptor 1-10 mRNA in cellular subsets of human peripheral blood mononuclear cells and sensitivity to CpG oligodeoxynucleotides" 6253: 4205:

Hoebe K, Du X, Georgel P, Janssen E, Tabeta K, Kim SO, et al. (August 2003). "Identification of Lps2 as a key transducer of MyD88-independent TIR signalling".

190:. Humans lack genes for TLR11, TLR12 and TLR13 and mice lack a functional gene for TLR10. The receptors TLR1, TLR2, TLR4, TLR5, TLR6, and TLR10 are located on the 435:, known as the "interleukin-1 receptor / toll-like receptor superfamily"; all members of this family have in common a so-called TIR (toll-IL-1 receptor) domain. 6021: 1824:, and constant low-level release of these modulators is thought to reduce the efficacy of opioid drug treatment with time, and is involved in opioid tolerance, 1761:. Meanwhile, activation of RIPK1 causes the polyubiquitination and activation of TAK1 and NFκB transcription in the same manner as the MyD88-dependent pathway. 590:. The toll pathway is similar to mammalian TLR signalling, but unlike mammalian TLRs, toll is not activated directly by pathogen-associated molecular patterns ( 538:

equivalents of certain TLR found in humans are not present in all mammals. For example, a gene coding for a protein analogous to TLR10 in humans is present in

630:), phosphorylated Cactus is polyubiquitylated and degraded, allowing nuclear translocation of DIF (dorsal-related immunity factor; a homologue of mammalian 6468: 1662:

derived from viruses as well as endogenous nucleic acids in context of pathogenic events. Activation of these receptor leads to production of inflammatory

4365:"TLR2 is constitutively expressed within the kidney and participates in ischemic renal injury through both MyD88-dependent and -independent pathways" 6074: 2440:

Rolls A, Shechter R, London A, Ziv Y, Ronen A, Levy R, Schwartz M (September 2007). "Toll-like receptors modulate adult hippocampal neurogenesis".

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are well known to be required for host defence against infection. The TLRs thus appear to be one of the most ancient, conserved components of the

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Hutchinson MR, Zhang Y, Brown K, Coats BD, Shridhar M, Sholar PW, Patel SJ, Crysdale NY, Harrison JA, Maier SF, Rice KC, Watkins LR (July 2008).

3689:"Identification of TLR10 as a key mediator of the inflammatory response to Listeria monocytogenes in intestinal epithelial cells and macrophages" 557:

Vertebrate TLRs are divided by similarity into the families of TLR 1/2/6/10/14/15, TLR 3, TLR 4, TLR 5, TLR 7/8/9, and TLR 11/12/13/16/21/22/23.

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The MyD88-dependent response occurs on dimerization of TLRs, and is used by every TLR except TLR3. Its primary effect is activation of NFκB and

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Shavit Y, Wolf G, Goshen I, Livshits D, Yirmiya R (May 2005). "Interleukin-1 antagonizes morphine analgesia and underlies morphine tolerance".

5370:"Cutting edge: Toll-like receptor 4 (TLR4)-deficient mice are hyporesponsive to lipopolysaccharide: evidence for TLR4 as the Lps gene product" 3210:(8th ed.). Philadelphia Baltimore New York London Buenos Aires Hong Kong Sydney Tokyo: Wolters Kluwer/Lippincott Williams & Wilkins. 3249: 3215: 416: 4701:"Dual regulation of mu opioid receptors in SK-N-SH neuroblastoma cells by morphine and interleukin-1β: evidence for opioid-immune crosstalk" 3594:

Chuang T, Ulevitch RJ (March 2001). "Identification of hTLR10: a novel human Toll-like receptor preferentially expressed in immune cells".

4090:"Commonly used prophylactic vaccines as an alternative for synthetically produced TLR ligands to mature monocyte-derived dendritic cells" 6246: 3222:

Broadly, TLRs 1, 2, 4 to 6, and 10 are expressed on the plasma membrane, and TLRs 3, 7 to 9, and 11 to 13 are confined to the endosomes.

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Hemmi H, Takeuchi O, Kawai T, Kaisho T, Sato S, Sanjo H, et al. (December 2000). "A Toll-like receptor recognizes bacterial DNA".

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Flajnik, Martin F.; Singh, Nevil J.; Holland, Steven M., eds. (2023). "Chapter 12 Pattern Recognition Receptors and the IL-1 Family".

2895:"The Circulating Protease Persephone Is an Immune Sensor for Microbial Proteolytic Activities Upstream of the Drosophila Toll Pathway" 1652: 459: 295: 5532: 5766: 4765: 4624: 4507: 1673:

The adapter proteins and kinases that mediate TLR signaling have also been targeted. In addition, random germline mutagenesis with

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Hutchinson MR, Lewis SS, Coats BD, Rezvani N, Zhang Y, Wieseler JL, Somogyi AA, Yin H, Maier SF, Rice KC, Watkins LR (May 2010).

4791:"Evidence that intrathecal morphine-3-glucuronide may cause pain enhancement via toll-like receptor 4/MD-2 and interleukin-1beta" 2515: 2484: 1702: 2395:

Sharma N, Akhade AS, Qadri A (April 2013). "Sphingosine-1-phosphate suppresses TLR-induced CXCL8 secretion from human T cells".

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Toll-like receptors have also been shown to be an important link between innate and adaptive immunity through their presence in

6624: 6519: 4525:"TLR7/8-agonist-loaded nanoparticles promote the polarization of tumour-associated macrophages to enhance cancer immunotherapy" 432: 198: 5603: 4859:"Etanercept restores the antinociceptive effect of morphine and suppresses spinal neuroinflammation in morphine-tolerant rats" 6239: 4625:"Toll like Receptor 7 Messenger Ribonucleic Acid Expression Levels in Dairy Animals in an Outbreak of Foot-and-mouth disease" 3418:"Lipopolysaccharide activates distinct signaling pathways in intestinal epithelial cell lines expressing Toll-like receptors" 366:. In the case of a viral factor, the infected cell may shut off its protein synthesis and may undergo programmed cell death ( 2846:"The dorsoventral regulatory gene cassette spätzle/Toll/cactus controls the potent antifungal response in Drosophila adults" 1934: 626:, Tube, and Pelle (the orthologue of mammalian IRAK). Signal from TICS is then transduced to Cactus (homologue of mammalian 1705:. Ligand binding and conformational change that occurs in the receptor recruits the adaptor protein MyD88, a member of the 354:, and survival, while others lead to greater adaptive immunity. If the ligand is a bacterial factor, the pathogen might be 1806:

TLR7 messenger RNA expression levels in dairy animals in a natural outbreak of foot-and-mouth disease have been reported.

1033: 523: 5009:"Non-stereoselective reversal of neuropathic pain by naloxone and naltrexone: involvement of toll-like receptor 4 (TLR4)" 1548:

Toll-Like Receptor (TLR) ligands among RNA and DNA viruses, Gram-positive and Gram-negative bacteria, fungi, and protists

6644: 6639: 6574: 6534: 6496: 6488: 3670: 600: 408: 105: 5618: 3244:. Lippincott's Illustrated reviews. Philadelphia: Wolters Kluwer Health/Lippincott Williams & Wilkins. p. 17. 6063: 2547:"Toll-like receptors 9 and 3 as essential components of innate immune defense against mouse cytomegalovirus infection" 1976: 1817: 1246: 307: 4040:"Bacteria evade immune recognition via TLR13 and binding of their 23S rRNA by MLS antibiotics by the same mechanisms" 5653: 4839: 4908:"An IL-1 receptor antagonist blocks a morphine-induced attenuation of locomotor recovery after spinal cord injury" 5474:"FLS2: an LRR receptor-like kinase involved in the perception of the bacterial elicitor flagellin in Arabidopsis" 402: 1991:(IL-1) receptor, also had homology to drosophila toll; the cytoplasmic portions of both molecules were similar. 6629: 5588: 5578: 1693:

TLR signaling is divided into two distinct signaling pathways, the MyD88-dependent and TRIF-dependent pathway.

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pufferfish. This may complicate the process of using experimental animals as models of human innate immunity.

5513: 4957:"The "toll" of opioid-induced glial activation: improving the clinical efficacy of opioids by targeting glia" 6514: 1929: 582: 574: 494: 315: 209: 108: 3492:

Salazar Gonzalez RM, Shehata H, O'Connell MJ, Yang Y, Moreno-Fernandez ME, Chougnet CA, Aliberti J (2014).

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Issa N, Guillaumot N, Lauret E, Matt N, Schaeffer-Reiss C, Van Dorsselaer A, et al. (February 2018).

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TIR domain from TLR2. This is a signal transduction domain distinct from the LRR domain discussed earlier.

195: 594:). Its receptor ectodomain recognizes the cleaved form of the cytokine spätzle, which is secreted in the 6634: 6369: 6333: 6328: 5827: 5649: 3887:"Expression and distribution of Toll-like receptors 11-13 in the brain during murine neurocysticercosis" 3664: 1746: 1586: 1059: 331: 3838:"Recognition of profilin by Toll-like receptor 12 is critical for host resistance to Toxoplasma gondii" 3629:

Hornung V, Rothenfusser S, Britsch S, Krug A, Jahrsdörfer B, Giese T, Endres S, Hartmann G (May 2002).

3494:"Toxoplasma gondii- derived profilin triggers human toll-like receptor 5-dependent cytokine production" 1963:" ("That's amazing!"), in reference to the underdeveloped ventral portion of a fruit fly larva. It was 6453: 5584: 3270:"The role of Toll-like receptors in the regulation of neutrophil migration, activation, and apoptosis" 1964: 6484: 6323: 6318: 6313: 6308: 6303: 6298: 6293: 6288: 6283: 5727: 5598: 5557: 5430: 5334: 5283: 4789:

Lewis SS, Hutchinson MR, Rezvani N, Loram LC, Zhang Y, Maier SF, Rice KC, Watkins LR (January 2010).

4456: 4265: 4214: 3996: 3741: 3103: 2620: 2558: 2356: 1796: 288: 101: 3787:"UNC93B1 is essential for TLR11 activation and IL-12-dependent host resistance to Toxoplasma gondii" 6547: 6524: 6262: 5594: 5562: 4150:"Pattern recognition receptor agonists in pathogen vaccines mediate antitumor T-cell cross-priming" 2516:"Cloning and characterization of a sub-family of human toll-like receptors: hTLR7, hTLR8 and hTLR9" 2173: 1887: 1870:

century, attests to the search for the key molecules and their receptors. More than 100 years ago,

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and LPS, respectively. For TLR3, dsRNA leads to activation of the receptor, recruiting the adaptor

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Yarovinsky F, Zhang D, Andersen JF, Bannenberg GL, Serhan CN, Hayden MS, et al. (June 2005).

3092:"Toll-Like Receptor 11 (TLR11) Interacts with Flagellin and Profilin through Disparate Mechanisms" 2025:

In turn, the other TLR genes were ablated in mice by gene targeting, largely in the laboratory of

1753:, which creates a branch in the signaling pathway. The TRIF/TBK1 signaling complex phosphorylates 5454: 5399: 5307: 5179: 4888: 4681: 4480: 4429: 4345: 4289: 4238: 4020: 3767: 3398: 2969: 2875: 2751:"More Than Black or White: Melanization and Toll Share Regulatory Serine Proteases in Drosophila" 2465: 2112:"TLR10 and Its Unique Anti-Inflammatory Properties and Potential Use as a Target in Therapeutics" 2011: 1895: 1644: 1608: 1558: 1324: 1072: 1022: 424: 283:

or in the form of invasive moieties during times of natural infection - to the TLR marks the key

6026: 1320: 1258: 5574: 614:. First, the Spätzle processing enzyme (SPE) is activated in response to infection and cleaves 6542: 6107: 6102: 6053: 6034: 5552: 5495: 5446: 5391: 5350: 5299: 5256: 5220: 5171: 5136: 5087: 5038: 4986: 4937: 4880: 4820: 4771: 4761: 4730: 4673: 4605: 4554: 4503: 4472: 4421: 4386: 4337: 4281: 4230: 4187: 4169: 4129: 4121: 4069: 4012: 3969: 3918: 3867: 3818: 3759: 3710: 3652: 3611: 3576: 3523: 3474: 3439: 3390: 3349: 3291: 3245: 3211: 3188: 3139: 3121: 3069: 3020: 2961: 2926: 2867: 2823: 2772: 2728: 2687: 2648: 2586: 2527: 2496: 2457: 2412: 2374: 2320: 2285: 2244: 2200: 2149: 2131: 2092: 1972: 1758: 1667: 1623: 1566: 1481: 1414: 1180: 934: 739: 615: 611: 265: 6152: 6132: 6122: 6117: 6087: 5613: 4523:

Rodell CB, Arlauckas SP, Cuccarese MF, Garris CS, Li R, Ahmed MS, et al. (August 2018).

4316:"The role of pattern-recognition receptors in innate immunity: update on Toll-like receptors" 3090:

Hatai, Hirotsugu; Lepelley, Alice; Zeng, Wangyong; Hayden, Matthew S.; Ghosh, Sankar (2016).

2683: 1725:, which in turn polyubiquinates the protein TAK1, as well as itself to facilitate binding to 781:, on the other hand, are efficiently infected. As a result, this knockout mouse can act as a 438:

Three subgroups of TIR domains exist. Proteins with subgroup 1 TIR domains are receptors for

419:(PAMPs). In addition to the recognition of exogenous PAMPs, TLRs can also bind to endogenous 6509: 6195: 6006: 5485: 5438: 5381: 5342: 5291: 5248: 5210: 5163: 5126: 5118: 5077: 5069: 5028: 5020: 4976: 4968: 4927: 4919: 4870: 4810: 4802: 4753: 4720: 4712: 4665: 4595: 4585: 4544: 4536: 4464: 4413: 4376: 4327: 4273: 4222: 4177: 4161: 4111: 4101: 4059: 4051: 4004: 3959: 3949: 3908: 3898: 3857: 3849: 3808: 3798: 3749: 3700: 3642: 3603: 3566: 3513: 3505: 3466: 3429: 3380: 3339: 3329: 3281: 3178: 3170: 3129: 3111: 3059: 3051: 3010: 3000: 2953: 2916: 2906: 2857: 2813: 2803: 2762: 2718: 2679: 2638: 2628: 2576: 2566: 2449: 2404: 2364: 2312: 2275: 2234: 2190: 2182: 2169:"Type B coxsackieviruses and their interactions with the innate and adaptive immune systems" 2139: 2123: 2082: 2072: 2007: 1999: 1946: 1544: 1227: 795: 767: 334:) that further amplify the signal and ultimately lead to the upregulation or suppression of 128: 6231: 4363:

Shigeoka AA, Holscher TD, King AJ, Hall FW, Kiosses WB, Tobias PS, et al. (May 2007).

1894:. In the decades that followed, endotoxin was chemically characterized and identified as a 1627:

Signaling pathway of toll-like receptors. Dashed grey lines represent unknown associations.

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Koblansky AA, Jankovic D, Oh H, Hieny S, Sungnak W, Mathur R, et al. (January 2013).

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Regan T, Nally K, Carmody R, Houston A, Shanahan F, Macsharry J, Brint E (December 2013).

2345:"A human homologue of the Drosophila Toll protein signals activation of adaptive immunity" 1995: 1442: 1103: 1092: 343: 319: 229: 81: 4639: 5434: 5338: 5287: 4460: 4269: 4218: 4182: 4149: 4000: 3745: 3555:"Human TLRs 10 and 1 share common mechanisms of innate immune sensing but not signaling" 3416:

Cario E, Rosenberg IM, Brandwein SL, Beck PL, Reinecker HC, Podolsky DK (January 2000).

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does not use the MyD88 dependent pathway. Its ligand is retroviral double-stranded RNA (

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The involvement of toll signalling in immunity was first demonstrated in the fruit fly,

522:. Molecular building blocks of the TLRs are represented in bacteria and in plants, and 493:

Please expand the section to include this information. Further details may exist on the

5712: 5131: 5122: 5106: 5082: 5057: 5033: 5008: 4981: 4956: 4932: 4907: 4815: 4806: 4790: 4725: 4700: 4600: 4573: 4549: 4524: 4064: 4039: 3964: 3937: 3913: 3886: 3862: 3837: 3813: 3786: 3518: 3493: 3344: 3317: 3183: 3158: 3134: 3091: 3064: 3039: 3015: 2988: 2921: 2894: 2818: 2792:"Immune genes and divergent antimicrobial peptides in flies of the subgenus Drosophila" 2791: 2643: 2608: 2239: 2222: 2195: 2168: 2144: 2087: 2060: 1938: 1891: 1841: 1800: 1604: 867: 455: 451: 385:, a TLR5 ligand, induces cytokine secretion on interacting with TLR5 on human T cells. 378: 370:). Immune cells that have detected a virus may also release anti-viral factors such as 241: 120: 5490: 5473: 5368:

Hoshino K, Takeuchi O, Kawai T, Sanjo H, Ogawa T, Takeda Y, et al. (April 1999).

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Nomura N, Miyajima N, Sazuka T, Tanaka A, Kawarabayasi Y, Sato S, et al. (1994).

4757: 3607: 2862: 2581: 2546: 2485:"Three novel mammalian toll-like receptors: gene structure, expression, and evolution" 6618: 6598: 5693: 5167: 5024: 4024: 2607:

Roach JC, Glusman G, Rowen L, Kaur A, Purcell MK, Smith KD, et al. (July 2005).

2061:"Sweeten PAMPs: Role of Sugar Complexed PAMPs in Innate Immunity and Vaccine Biology" 1988: 1942: 1911: 1632: 782: 778: 774: 527: 363: 233: 191: 124: 112: 5458: 5183: 4906:

Hook MA, Washburn SN, Moreno G, Woller SA, Puga D, Lee KH, Grau JW (February 2011).

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Several TLR ligands are in clinical development or being tested in animal models as

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small synthetic compounds; single-stranded Viral RNA, phagocytized bacterial RNA(24)

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Lee J, Sayed N, Hunter A, Au KF, Wong WH, Mocarski ES, et al. (October 2012).

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Tabeta K, Georgel P, Janssen E, Du X, Hoebe K, Crozat K, et al. (March 2004).

2026: 1825: 1659: 872: 710: 539: 519: 439: 355: 339: 202: 17: 5750: 5442: 5419:"A receptor kinase-like protein encoded by the rice disease resistance gene, Xa21" 5417:

Song WY, Wang GL, Chen LL, Kim HS, Pi LY, Holsten T, et al. (December 1995).

5346: 3157:

Mathur R, Oh H, Zhang D, Park SG, Seo J, Koblansky A, et al. (October 2012).

1816:. Its activation leads to downstream release of inflammatory modulators including 4875: 4858: 4381: 4364: 4106: 4089: 3853: 3116: 2911: 2767: 2750: 6602: 5968: 5916: 5812: 5737: 5704: 5675: 5666: 5386: 5369: 3647: 3630: 3470: 2036:

were discovered by Pamela Ronald in 1995 (rice XA21) and Thomas Boller in 2000 (

1899: 1875: 1785: 1766: 1640: 1562: 1270: 962: 924: 838: 587: 351: 303: 280: 269: 245: 135: 74: 5418: 4972: 4669: 3541: 3434: 3417: 3174: 3055: 3040:"Activation of innate immunity is required for efficient nuclear reprogramming" 2613:

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

2551:

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

2280: 2263: 5073: 4923: 4857:

Shen CH, Tsai RY, Shih MS, Lin SL, Tai YH, Chien CC, Wong CS (February 2011).

4540: 3936:

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2844:

Lemaitre B, Nicolas E, Michaut L, Reichhart JM, Hoffmann JA (September 1996).

2808: 2017: 1849: 1789: 1600: 1514:

bacterial ribosomal RNA sequence "CGGAAAGACC" (but not the methylated version)

1445: 1277: 1196: 1082: 914: 887: 861: 803: 799: 639: 595: 543: 515: 443: 371: 261: 116: 5058:"Proinflammatory cytokines oppose opioid-induced acute and chronic analgesia" 4173: 4165: 4125: 3705: 3688: 3571: 3554: 3125: 3005: 2723: 2706: 2135: 2077: 6582: 5980: 5215: 5198: 4590: 4008: 3954: 3803: 3754: 3553:

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Takeda K, Akira S (January 2005). "Toll-like receptors in innate immunity".

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Mechanism of allodynia evoked by intrathecal morphine-3-glucuronide in mice

4734: 4677: 4609: 4558: 4476: 4425: 4390: 4341: 4285: 4234: 4191: 4133: 4073: 4016: 3973: 3922: 3903: 3871: 3822: 3763: 3714: 3656: 3615: 3580: 3527: 3478: 3443: 3394: 3385: 3368: 3353: 3295: 3192: 3143: 3073: 3024: 2965: 2930: 2845: 2827: 2776: 2732: 2691: 2667: 2652: 2590: 2531: 2500: 2461: 2416: 2324: 2289: 2248: 2204: 2153: 2127: 2096: 1979:, which it achieved by activating the synthesis of antimicrobial peptides. 1957: 479: 208:

TLRs received their name from their similarity to the protein coded by the

5450: 5354: 5303: 5260: 5224: 5175: 4574:"Immune Adjuvant Effect of Molecularly-defined Toll-Like Receptor Ligands" 3730:"TLR11 activation of dendritic cells by a protozoan profilin-like protein" 2987:

Chowdhury M, Li CF, He Z, Lu Y, Liu XS, Wang YF, et al. (June 2019).

2871: 2378: 1953:

axis. It was named after Christiane Nüsslein-Volhard's 1985 exclamation, "

6030: 5984: 5683: 1923: 1737:

Both TLR3 and TLR4 use the TRIF-dependent pathway, which is triggered by

1578: 1570: 1475: 1408: 1174: 989: 857: 631: 551: 447: 412: 347: 311: 225: 221: 127:. Once these microbes have reached physical barriers such as the skin or 4468: 4226: 2408: 6147: 6142: 6137: 6127: 6112: 6097: 6092: 6082: 4752:. International Review of Neurobiology. Vol. 85. pp. 207–19. 4055: 3938:"A novel Toll-like receptor that recognizes vesicular stomatitis virus" 2186: 1950: 1658:

A set of endosomal TLRs comprising TLR3, TLR7, TLR8 and TLR9 recognize

958: 463: 359: 299: 276: 97: 4116: 3509: 3369:"Regulating B-cell activation and survival in response to TLR signals" 1975:

and his colleagues to have an essential role in the fly's immunity to

298:(proteins that mediate other protein-protein interactions) within the 6437: 6427: 6412: 6220: 6215: 6210: 6205: 6200: 6190: 6185: 6180: 6175: 6170: 5903: 5894: 5787: 5295: 4277: 2110:

Fore, Faith; Indriputri, Cut; Mamutse, Janet; Nugraha, Jusak (2020).

1837: 1813: 1437: 1403: 1372: 1124: 1114: 733: 415:

but distinguishable from host molecules, collectively referred to as

291:

responses and the development of antigen-specific acquired immunity.

257: 253: 131: 4332: 4315: 3729: 2957: 2749:

Dudzic JP, Hanson MA, Iatsenko I, Kondo S, Lemaitre B (April 2019).

2453: 673:

has also been designated as CD282 (cluster of differentiation 282).

627: 3596:

Biochimica et Biophysica Acta (BBA) - Gene Structure and Expression

3286: 3269: 731:, but mice cells do. Mouse-specific TLR11 recognizes uropathogenic 6552: 6442: 6432: 6417: 6402: 6397: 6392: 6387: 5875: 5871: 5274:

Gay NJ, Keith FJ (May 1991). "Drosophila Toll and IL-1 receptor".

4417: 3334: 2369: 2344: 2016: 1861:

analgesic drugs without having any affinity for μ-opioid receptor

1750: 1738: 1722: 1718: 1714: 1710: 1706: 1683: 1679: 1622: 1612: 1543: 1431: 1292: 1222: 1188: 1134: 1017: 980: 946: 881: 846: 832: 728: 715: 706: 685: 623: 591: 568: 542:, but appears to have been damaged at some point in the past by a 392: 335: 327: 323: 179: 175: 4623:

Audarya, S.D.; Pattnaik, B.; Sanyal, A.; Mohapatra, J.K. (2017).

1812:

has been shown to be important for the long-term side-effects of

656:

family genes that interact with each other to differing degrees.

6562: 6557: 6422: 6407: 6359: 6016: 6011: 5958: 5953: 5943: 5879: 5851: 5835: 5831: 5745: 5722: 5717: 5689: 4748:

Komatsu T, Sakurada S, Katsuyama S, Sanai K, Sakurada T (2009).

1833: 1821: 1809: 1754: 1742: 1726: 1687: 1648: 1336: 997: 702: 698: 694: 689: 681: 670: 665: 171: 167: 163: 159: 155: 151: 147: 143: 139: 6457: 6235: 5622: 2390: 2388: 1941:

and colleagues. It was known for its developmental function in

427:(HSPs) or plasma membrane constituents. TLRs together with the 393: 123:, that recognize structurally conserved molecules derived from 5990: 5976: 5604:

The Toll-Like Receptor Family of Innate Immune Receptors (pdf)

4955:

Watkins LR, Hutchinson MR, Rice KC, Maier SF (November 2009).

2006:. TLR 4 function as an LPS sensing receptor was discovered by 1907: 1757:

allowing its translocation into the nucleus and production of

1674: 1582: 1574: 473: 3318:"The instructive role of dendritic cells on T-cell responses" 3785:

Pifer R, Benson A, Sturge CR, Yarovinsky F (February 2011).

1803:

in 2017, which contains a monophosphoryl lipid A component.

606:

The toll pathway is activated by different stimuli, such as

314:

are then responsible for the subsequent activation of other

232:

called toll-like receptors (TLRs) that are expressed on the

2264:"[Innate immunity: structure and function of TLRs]" 1721:. IRAK kinases then phosphorylate and activate the protein 1655:) are known to facilitate the presentation of LPS to MD-2. 466:

and mediate signaling from proteins of subgroups 1 and 2.

4699:

Mohan S, Davis RL, DeSilva U, Stevens CW (October 2010).

4500:

Dermatologie Venerologie : Grundlagen. Klinik. Atlas

2343:

Medzhitov R, Preston-Hurlburt P, Janeway CA (July 1997).

2059:

Mahla RS, Reddy MC, Prasad DV, Kumar H (September 2013).

1799:, with the first clinical use in humans in a recombinant 411:(PRR) and recognize molecules that are broadly shared by 1967:

by the laboratory of Kathryn Anderson in 1988. In 1996,

798:. Furthermore, they are expressed by different types of 773:, which causes food- and waterborne gastroenteritis and 3263: 3261: 693:

retroviral expression of four transcriptional factors (

50:

region of toll-like receptors, represented here by TLR3

747:

its ligand is the protein profilin and the ligand for

3234: 3232: 3230: 2483:

Du X, Poltorak A, Wei Y, Beutler B (September 2000).

2262:

Delneste Y, Beauvillain C, Jeannin P (January 2007).

766:

effectively, normal mice do not get infected by oral

3240:

Waltenbaugh C, Doan T, Melvold R, Viselli S (2008).

1769:

yet tightly regulated gateways for gene modulation.

194:, whereas TLR3, TLR7, TLR8, and TLR9 are located in 6591: 6573: 6533: 6495: 6378: 6352: 6274: 6161: 6073: 6052: 5999: 5967: 5932: 5902: 5893: 5864: 5844: 5820: 5811: 5795: 5786: 5779: 5759: 5736: 5703: 5674: 5663: 2790:Hanson MA, Hamilton PT, Perlman SJ (October 2016). 2223:"Toll to be paid at the gateway to the vessel wall" 68: 60: 55: 34: 2227:Arteriosclerosis, Thrombosis, and Vascular Biology 3459:Journal of Trace Elements in Medicine and Biology 2609:"The evolution of vertebrate Toll-like receptors" 1933:, was discovered in 1985 by 1995 Nobel Laureates 5595:TollML: Toll-like receptors and ligands database 5514:"The Nobel Prize in Physiology or Medicine 2011" 3885:Mishra BB, Gundra UM, Teale JM (December 2008). 2839: 2837: 5002: 5000: 3085: 3083: 3268:Sabroe I, Dower SK, Whyte MK (November 2005). 2216: 2214: 1832:. Morphine induced TLR4 activation attenuates 634:) and induction of transcription of genes for 220:The ability of the immune system to recognize 134:, they are recognized by TLRs, which activate 6469: 6247: 5634: 4309: 4307: 4305: 4303: 3682: 3680: 3367:Gerondakis S, Grumont RJ, Banerjee A (2007). 3159:"A mouse model of Salmonella typhi infection" 2744: 2742: 2668:"The host defense of Drosophila melanogaster" 2602: 2600: 2338: 2336: 2334: 8: 5533:"B.C. doctor wins prestigious medical prize" 4840:"Small Molecules Target Toll-Like Receptors" 3542:https://doi.org/10.1371/journal.pone.0267860 3311: 3309: 3307: 3305: 3238:Unless else specified in boxes then ref is: 2705:Valanne S, Wang JH, Rämet M (January 2011). 2167:Kemball CC, Alirezaei M, Whitton JL (2010). 1954: 1690:, and TRAM (TRIF-related adaptor molecule). 1569:; proteins such as flagellin from bacterial 1319:Intestinal epithelial cells (IECs) *only in 727:As noted above, human cells do not express 491:about choanoflagellate TLR (pmid29848444) . 6476: 6462: 6454: 6254: 6240: 6232: 5899: 5817: 5792: 5783: 5671: 5641: 5627: 5619: 1852:, and other negative side effects such as 573:The Toll immunity pathway as found in the 40: 5587:at the U.S. National Library of Medicine 5577:at the U.S. National Library of Medicine 5489: 5385: 5214: 5130: 5081: 5032: 4980: 4931: 4874: 4814: 4724: 4599: 4589: 4548: 4380: 4331: 4181: 4115: 4105: 4063: 3963: 3953: 3912: 3902: 3861: 3812: 3802: 3753: 3704: 3646: 3641:(9). Baltimore, Md. : 1950: 4531–7. 3570: 3517: 3433: 3384: 3343: 3333: 3285: 3182: 3133: 3115: 3063: 3014: 3004: 2920: 2910: 2861: 2817: 2807: 2766: 2722: 2642: 2632: 2580: 2570: 2514:Chuang TH, Ulevitch RJ (September 2000). 2368: 2279: 2238: 2194: 2143: 2086: 2076: 1917:The prototypic member of the family, the 1914:to the presence of microbial infections. 1635:. Though most TLRs appear to function as 279:- either in the form of adjuvant used in 4038:Hochrein H, Kirschning CJ (March 2013). 2684:10.1146/annurev.immunol.25.022106.141615 808: 755:. The flagellin from the enteropathogen 719:pluripotent stem cells (iPSC) colonies. 2707:"The Drosophila Toll signaling pathway" 2051: 1840:and enhances the development of opioid 1788:) is a TLR7 agonist, and its successor 3662: 1882:" to describe a substance produced by 417:pathogen-associated molecular patterns 346:events. Some of these events lead to 31: 5472:Gómez-Gómez L, Boller T (June 2000). 7: 5013:The European Journal of Neuroscience 762:As mouse TLR11 is able to recognize 421:damage-associated molecular patterns 228:is, in part, due to the presence of 4572:Toussi DN, Massari P (April 2014). 4154:Journal for Immunotherapy of Cancer 3942:The Journal of Biological Chemistry 3791:The Journal of Biological Chemistry 3316:Sallusto F, Lanzavecchia A (2002). 2993:The Journal of Biological Chemistry 2221:Hansson GK, Edfeldt K (June 2005). 2010:and colleagues. These workers used 524:plant pattern recognition receptors 5123:10.1016/j.neuroscience.2010.02.011 4961:Trends in Pharmacological Sciences 4807:10.1016/j.neuroscience.2009.10.011 2240:10.1161/01.ATV.0000168894.43759.47 2021:The history of Toll-like receptors 25: 5767:Polymeric immunoglobulin receptor 1670:) to help fight viral infection. 1631:TLRs are believed to function as 306:to propagate the antigen-induced 252:, cells of the adaptive immunity 5025:10.1111/j.1460-9568.2008.06321.x 1703:Mitogen-activated protein kinase 1577:of viruses; or the unmethylated 478: 4844:Chemical & Engineering News 4502:(in German). Berlin: Springer. 2666:Lemaitre B, Hoffmann J (2007). 1666:as well as type I interferons ( 1581:islands of bacterial and viral 789:Summary of known mammalian TLRs 287:events that ultimately lead to 27:Class of immune system proteins 5979:(with two glycoprotein chains 4717:10.1016/j.jneuroim.2010.06.007 737:and the apicomplexan parasite 294:Upon activation, TLRs recruit 1: 6489:Pattern recognition receptors 6075:Killer-cell IG-like receptors 5491:10.1016/S1097-2765(00)80265-8 5443:10.1126/science.270.5243.1804 5347:10.1126/science.282.5396.2085 5062:Brain, Behavior, and Immunity 4912:Brain, Behavior, and Immunity 4758:10.1016/S0074-7742(09)85016-2 4529:Nature Biomedical Engineering 4314:Kawai T, Akira S (May 2010). 3608:10.1016/s0167-4781(00)00289-x 3208:Paul's fundamental immunology 2863:10.1016/s0092-8674(00)80172-5 1886:that could provoke fever and 1745:. TRIF activates the kinases 759:is also recognized by TLR11. 201:(because they are sensors of 5531:Mitchell B (23 March 2011). 5168:10.1016/0092-8674(88)90516-8 4876:10.1213/ANE.0b013e3182025b15 4382:10.4049/jimmunol.178.10.6252 4107:10.1182/blood-2009-11-251884 3891:Journal of Neuroinflammation 3854:10.1016/j.immuni.2012.09.016 3274:Clinical Infectious Diseases 3117:10.1371/journal.pone.0148987 2912:10.1016/j.molcel.2018.01.029 2768:10.1016/j.celrep.2019.03.101 2397:Journal of Leukocyte Biology 1709:family. MyD88 then recruits 1531:conventional dendritic cells 1501:conventional dendritic cells 1498:plasmacytoid dendritic cells 1362:Plasmacytoid dendritic cells 1247:Plasmacytoid dendritic cells 1193:multiple diacyl lipopeptides 1056:B lymphocytes (only in mice) 409:pattern recognition receptor 138:responses. The TLRs include 100:that play a key role in the 6163:Leukocyte IG-like receptors 5387:10.4049/jimmunol.162.7.3749 3648:10.4049/jimmunol.168.9.4531 3471:10.1016/j.jtemb.2017.07.006 3373:Immunology and Cell Biology 2672:Annual Review of Immunology 1935:Christiane Nüsslein-Volhard 1313:a subset of dendritic cells 1158:a subset of dendritic cells 777:in humans. TLR11 deficient 454:and all have extracellular 308:signal transduction pathway 224:that are broadly shared by 6661: 5654:immunoglobulin superfamily 5609:Toll-Like receptor Pathway 4973:10.1016/j.tips.2009.08.002 4838:Drahl C (22 August 2012). 4705:Journal of Neuroimmunology 4670:10.1016/j.pain.2005.02.003 3498:Journal of Innate Immunity 3435:10.4049/jimmunol.164.2.966 3175:10.1016/j.cell.2012.08.042 3056:10.1016/j.cell.2012.09.034 2946:Nature Reviews. Immunology 1393:Intestinal epitelial cells 923:multiple lipopeptides and 663: 400: 5074:10.1016/j.bbi.2008.05.004 4924:10.1016/j.bbi.2010.10.018 4541:10.1038/s41551-018-0236-8 2809:10.1186/s12862-016-0805-y 2520:European Cytokine Network 2489:European Cytokine Network 1651:and LPS-Binding Protein ( 1425: 1422: 1419: 1402: 1329:hippocampal interneurons 1240: 1237: 1234: 1232:small synthetic compounds 1231: 1221: 1187: 1152: 1149: 1146: 1133: 1041: 1038: 1031: 1016: 929:Bacterial peptidoglycans 900: 897: 894: 880: 588:immune deficiency pathway 403:Toll-interleukin receptor 80: 39: 5589:Medical Subject Headings 5585:Toll+protein,+Drosophila 5579:Medical Subject Headings 4863:Anesthesia and Analgesia 4166:10.1136/jitc-2023-007198 3706:10.4049/jimmunol.1203245 3669:: CS1 maint: location ( 3572:10.4049/jimmunol.0901888 3006:10.1074/jbc.RA118.006804 2796:BMC Evolutionary Biology 2724:10.4049/jimmunol.1002302 2305:International Immunology 2281:10.1051/medsci/200723167 2078:10.3389/fimmu.2013.00248 2004:adaptive immune response 1377:triacylated lipopeptides 1343:CpG Oligodeoxynucleotide 1077:Bacteria and host cells 892:Bacterial peptidoglycans 785:of human typhoid fever. 550:, which is found in the 260:, and non-immune cells ( 6515:Formyl peptide receptor 5650:Transmembrane receptors 4591:10.3390/vaccines2020323 4009:10.1126/science.1220363 3955:10.1074/jbc.M110.159590 3804:10.1074/jbc.M110.171025 3755:10.1126/science.1109893 2634:10.1073/pnas.0502272102 2572:10.1073/pnas.0400525101 2065:Frontiers in Immunology 1930:Drosophila melanogaster 1927:; Tl) in the fruit fly 1697:MyD88-dependent pathway 1647:, which requires MD-2. 1050:Myeloid dendritic cells 910:Myeloid dendritic cells 688:), which activates the 583:Drosophila melanogaster 429:Interleukin-1 receptors 6625:Developmental genetics 5253:10.1006/geno.1996.0150 4498:Peter Fritsch (2004). 3904:10.1186/1742-2094-5-53 3386:10.1038/sj.icb.7100097 2128:10.4110/in.2020.20.e21 2022: 1955: 1884:Gram-negative bacteria 1854:respiratory depression 1733:TRIF-dependent pathway 1628: 1549: 1028:Gram-negative bacteria 940:Gram-positive bacteria 636:antimicrobial peptides 608:gram-positive bacteria 577: 489:is missing information 398: 358:and digested, and its 6380:Downstream signalling 5614:BioScience Animations 5374:Journal of Immunology 5216:10.1093/dnares/1.1.27 4369:Journal of Immunology 3693:Journal of Immunology 3635:Journal of Immunology 3559:Journal of Immunology 3422:Journal of Immunology 2711:Journal of Immunology 2317:10.1093/intimm/dxh186 2020: 1801:herpes zoster vaccine 1626: 1547: 1528:monocytes/macrophages 1428:monocytes/macrophages 1396:monocytes/macrophages 1359:monocytes/macrophages 1348:Bacteria, DNA viruses 1310:monocytes/macrophages 1243:monocytes/macrophages 1209:monocytes/macrophages 1161:Intestinal epithelium 1060:Intestinal epithelium 1044:monocytes/macrophages 903:monocytes/macrophages 843:Bacterial lipoprotein 572: 462:that are exclusively 442:that are produced by 396: 111:usually expressed on 6485:Innate immune system 5599:University of Munich 5558:Immunologic adjuvant 3328:(Suppl 3): S127-32. 2047:Notes and references 1945:by establishing the 1892:experimental animals 1784:(cardinally used in 1565:, lipopeptides, and 1155:monocyte/macrophages 514:TLRs are present in 342:responses and other 318:proteins, including 310:. These recruited 250:natural killer cells 102:innate immune system 6645:Toll-like receptors 6640:Signal transduction 6548:RIG-I-like receptor 6525:Complement receptor 6497:Membrane-bound_PRRs 6000:Accessory molecules 5865:Accessory molecules 5575:Toll-Like+Receptors 5563:RIG-I-like receptor 5435:1995Sci...270.1804S 5339:1998Sci...282.2085P 5288:1991Natur.351..355G 4638:(3). Archived from 4469:10.1038/nature01182 4461:2002Natur.420..324Y 4270:2000Natur.408..740H 4227:10.1038/nature01889 4219:2003Natur.424..743H 4001:2012Sci...337.1111O 3746:2005Sci...308.1626Y 3280:(Suppl 7): S421-6. 3108:2016PLoSO..1148987H 2999:(26): 10172–10181. 2761:(4): 1050–1061.e3. 2625:2005PNAS..102.9577R 2563:2004PNAS..101.3516T 2442:Nature Cell Biology 2409:10.1189/jlb.0712328 2361:1997Natur.388..394M 2174:Future Microbiology 1906:, and unmethylated 1878:, coined the term " 1686:(also called Mal), 1609:heat shock proteins 1559:lipopolysaccharides 1284:single-stranded RNA 1164:Breast cancer cells 1140:Bacterial flagellin 1073:heat shock proteins 1064:Breast cancer cells 986:double-stranded RNA 425:heat shock proteins 407:TLRs are a type of 90:Toll-like receptors 48:leucine-rich repeat 18:Toll-like receptors 6520:Scavenger receptor 6505:Toll-like receptor 6064:cytokine receptors 5664:Antibody receptor: 4056:10.4161/onci.23141 3322:Arthritis Research 2187:10.2217/fmb.10.101 2032:Plant homologs of 2023: 2012:positional cloning 1896:lipopolysaccharide 1629: 1594:Endogenous ligands 1573:; double-stranded 1550: 1325:ulcerative colitis 1023:lipopolysaccharide 648:have a total of 9 578: 399: 64:Toll-like receptor 35:Toll-like receptor 6612: 6611: 6543:NOD-like receptor 6451: 6450: 6269:signaling pathway 6263:Signaling pathway 6229: 6228: 6054:Cytokine receptor 6048: 6047: 6044: 6043: 5889: 5888: 5860: 5859: 5775: 5774: 5553:NOD-like receptor 5520:. 3 October 2011. 4645:on 28 April 2021. 4406:Nature Immunology 4320:Nature Immunology 3510:10.1159/000362367 3251:978-0-7817-9543-2 3217:978-1-9751-4251-3 2905:(4): 539–550.e6. 2268:Médecine/Sciences 1973:Jules A. Hoffmann 1797:vaccine adjuvants 1777:Medical relevance 1759:Interferon type I 1668:interferon type I 1567:lipoarabinomannan 1537: 1536: 1482:Toxoplasma gondii 1415:Toxoplasma gondii 1181:Toxoplasma gondii 1002:cell compartment 935:lipoteichoic acid 837:multiple triacyl 796:cell compartments 740:Toxoplasma gondii 612:virulence factors 512: 511: 338:that orchestrate 266:endothelial cells 96:) are a class of 87: 86: 16:(Redirected from 6652: 6535:Cytoplasmic_PRRs 6510:Mannose receptor 6478: 6471: 6464: 6455: 6256: 6249: 6242: 6233: 5933:Antigen receptor 5900: 5818: 5796:Antigen receptor 5793: 5784: 5780:Antigen receptor 5738:Alpha (α)/mu (μ) 5672: 5657:immune receptors 5643: 5636: 5629: 5620: 5541: 5540: 5528: 5522: 5521: 5510: 5504: 5503: 5493: 5469: 5463: 5462: 5429:(5243): 1804–6. 5414: 5408: 5407: 5389: 5365: 5359: 5358: 5333:(5396): 2085–8. 5322: 5316: 5315: 5296:10.1038/351355b0 5271: 5265: 5264: 5235: 5229: 5228: 5218: 5194: 5188: 5187: 5151: 5145: 5144: 5134: 5102: 5096: 5095: 5085: 5053: 5047: 5046: 5036: 5004: 4995: 4994: 4984: 4952: 4946: 4945: 4935: 4903: 4897: 4896: 4878: 4854: 4848: 4847: 4835: 4829: 4828: 4818: 4786: 4780: 4779: 4745: 4739: 4738: 4728: 4696: 4690: 4689: 4653: 4647: 4646: 4644: 4632:Buffalo Bulletin 4629: 4620: 4614: 4613: 4603: 4593: 4569: 4563: 4562: 4552: 4520: 4514: 4513: 4495: 4489: 4488: 4444: 4438: 4437: 4401: 4395: 4394: 4384: 4360: 4354: 4353: 4335: 4311: 4298: 4297: 4278:10.1038/35047123 4253: 4247: 4246: 4202: 4196: 4195: 4185: 4144: 4138: 4137: 4119: 4109: 4084: 4078: 4077: 4067: 4035: 4029: 4028: 3995:(6098): 1111–5. 3984: 3978: 3977: 3967: 3957: 3933: 3927: 3926: 3916: 3906: 3882: 3876: 3875: 3865: 3833: 3827: 3826: 3816: 3806: 3782: 3776: 3775: 3757: 3740:(5728): 1626–9. 3725: 3719: 3718: 3708: 3684: 3675: 3674: 3668: 3660: 3650: 3626: 3620: 3619: 3591: 3585: 3584: 3574: 3550: 3544: 3538: 3532: 3531: 3521: 3489: 3483: 3482: 3454: 3448: 3447: 3437: 3413: 3407: 3406: 3388: 3364: 3358: 3357: 3347: 3337: 3313: 3300: 3299: 3289: 3265: 3256: 3255: 3236: 3225: 3224: 3203: 3197: 3196: 3186: 3154: 3148: 3147: 3137: 3119: 3087: 3078: 3077: 3067: 3035: 3029: 3028: 3018: 3008: 2984: 2978: 2977: 2941: 2935: 2934: 2924: 2914: 2890: 2884: 2883: 2865: 2841: 2832: 2831: 2821: 2811: 2787: 2781: 2780: 2770: 2746: 2737: 2736: 2726: 2702: 2696: 2695: 2663: 2657: 2656: 2646: 2636: 2604: 2595: 2594: 2584: 2574: 2542: 2536: 2535: 2511: 2505: 2504: 2480: 2474: 2473: 2437: 2431: 2427: 2421: 2420: 2392: 2383: 2382: 2372: 2340: 2329: 2328: 2300: 2294: 2293: 2283: 2259: 2253: 2252: 2242: 2218: 2209: 2208: 2198: 2181:(9): 1329–1347. 2164: 2158: 2157: 2147: 2107: 2101: 2100: 2090: 2080: 2056: 2008:Bruce A. Beutler 2000:Ruslan Medzhitov 1977:fungal infection 1962: 1926: 1872:Richard Pfeiffer 1523:cell compartment 1490:cell compartment 1423:cell compartment 1354:cell compartment 1305:cell compartment 1238:cell compartment 1228:imidazoquinoline 809: 709:; OSKM) induces 507: 504: 498: 482: 474: 460:adaptor proteins 431:form a receptor 423:(DAMPs) such as 296:adaptor proteins 230:immune receptors 129:intestinal tract 44: 32: 21: 6660: 6659: 6655: 6654: 6653: 6651: 6650: 6649: 6630:Insect immunity 6615: 6614: 6613: 6608: 6592:Other/ungrouped 6587: 6569: 6529: 6491: 6482: 6452: 6447: 6374: 6353:Other receptors 6348: 6270: 6260: 6230: 6225: 6157: 6069: 6040: 5995: 5963: 5928: 5885: 5856: 5840: 5807: 5771: 5755: 5732: 5699: 5665: 5659: 5647: 5571: 5549: 5544: 5530: 5529: 5525: 5512: 5511: 5507: 5471: 5470: 5466: 5416: 5415: 5411: 5367: 5366: 5362: 5324: 5323: 5319: 5282:(6325): 355–6. 5273: 5272: 5268: 5237: 5236: 5232: 5196: 5195: 5191: 5153: 5152: 5148: 5104: 5103: 5099: 5055: 5054: 5050: 5006: 5005: 4998: 4954: 4953: 4949: 4905: 4904: 4900: 4856: 4855: 4851: 4837: 4836: 4832: 4788: 4787: 4783: 4768: 4747: 4746: 4742: 4698: 4697: 4693: 4655: 4654: 4650: 4642: 4627: 4622: 4621: 4617: 4571: 4570: 4566: 4522: 4521: 4517: 4510: 4497: 4496: 4492: 4455:(6913): 324–9. 4446: 4445: 4441: 4412:(11): 1144–50. 4403: 4402: 4398: 4362: 4361: 4357: 4333:10.1038/ni.1863 4313: 4312: 4301: 4264:(6813): 740–5. 4255: 4254: 4250: 4213:(6950): 743–8. 4204: 4203: 4199: 4146: 4145: 4141: 4086: 4085: 4081: 4037: 4036: 4032: 3986: 3985: 3981: 3935: 3934: 3930: 3884: 3883: 3879: 3835: 3834: 3830: 3784: 3783: 3779: 3727: 3726: 3722: 3699:(12): 6084–92. 3686: 3685: 3678: 3661: 3628: 3627: 3623: 3602:(1–2): 157–61. 3593: 3592: 3588: 3565:(9): 5094–103. 3552: 3551: 3547: 3539: 3535: 3491: 3490: 3486: 3456: 3455: 3451: 3415: 3414: 3410: 3366: 3365: 3361: 3315: 3314: 3303: 3267: 3266: 3259: 3252: 3239: 3237: 3228: 3218: 3205: 3204: 3200: 3156: 3155: 3151: 3102:(2): e0148987. 3089: 3088: 3081: 3037: 3036: 3032: 2986: 2985: 2981: 2958:10.1038/nri2194 2943: 2942: 2938: 2892: 2891: 2887: 2843: 2842: 2835: 2789: 2788: 2784: 2748: 2747: 2740: 2704: 2703: 2699: 2665: 2664: 2660: 2619:(27): 9577–82. 2606: 2605: 2598: 2557:(10): 3516–21. 2544: 2543: 2539: 2513: 2512: 2508: 2482: 2481: 2477: 2454:10.1038/ncb1629 2439: 2438: 2434: 2428: 2424: 2394: 2393: 2386: 2355:(6640): 394–7. 2342: 2341: 2332: 2302: 2301: 2297: 2261: 2260: 2256: 2220: 2219: 2212: 2166: 2165: 2161: 2109: 2108: 2104: 2058: 2057: 2053: 2049: 1996:Charles Janeway 1922: 1874:, a student of 1867: 1836:suppression by 1779: 1735: 1699: 1621: 1596: 1542: 1517:Virus, bacteria 1443:urinary bladder 1104:hyaluronic acid 1093:heparan sulfate 1007:Dendritic cells 973:Numerous others 868:dendritic cells 818:Ligand location 791: 725: 679: 668: 662: 638:(AMPs) such as 567: 508: 502: 499: 492: 483: 472: 470:Extended family 452:dendritic cells 405: 391: 379:dendritic cells 344:transcriptional 320:protein kinases 275:The binding of 242:dendritic cells 218: 121:dendritic cells 106:single-spanning 51: 28: 23: 22: 15: 12: 11: 5: 6658: 6656: 6648: 6647: 6642: 6637: 6632: 6627: 6617: 6616: 6610: 6609: 6607: 6606: 6595: 6593: 6589: 6588: 6586: 6585: 6579: 6577: 6571: 6570: 6568: 6567: 6566: 6565: 6560: 6555: 6545: 6539: 6537: 6531: 6530: 6528: 6527: 6522: 6517: 6512: 6507: 6501: 6499: 6493: 6492: 6483: 6481: 6480: 6473: 6466: 6458: 6449: 6448: 6446: 6445: 6440: 6435: 6430: 6425: 6420: 6415: 6410: 6405: 6400: 6395: 6390: 6384: 6382: 6376: 6375: 6373: 6372: 6367: 6362: 6356: 6354: 6350: 6349: 6347: 6346: 6341: 6336: 6331: 6326: 6321: 6316: 6311: 6306: 6301: 6296: 6291: 6286: 6280: 6278: 6272: 6271: 6261: 6259: 6258: 6251: 6244: 6236: 6227: 6226: 6224: 6223: 6218: 6213: 6208: 6203: 6198: 6193: 6188: 6183: 6178: 6173: 6167: 6165: 6159: 6158: 6156: 6155: 6150: 6145: 6140: 6135: 6130: 6125: 6120: 6115: 6110: 6105: 6100: 6095: 6090: 6085: 6079: 6077: 6071: 6070: 6068: 6067: 6058: 6056: 6050: 6049: 6046: 6045: 6042: 6041: 6039: 6038: 6024: 6019: 6014: 6009: 6003: 6001: 5997: 5996: 5994: 5993: 5988: 5973: 5971: 5965: 5964: 5962: 5961: 5956: 5951: 5946: 5936: 5934: 5930: 5929: 5927: 5926: 5925: 5924: 5919: 5908: 5906: 5897: 5891: 5890: 5887: 5886: 5884: 5883: 5868: 5866: 5862: 5861: 5858: 5857: 5855: 5854: 5848: 5846: 5842: 5841: 5839: 5838: 5824: 5822: 5815: 5809: 5808: 5806: 5805: 5799: 5797: 5790: 5781: 5777: 5776: 5773: 5772: 5770: 5769: 5763: 5761: 5757: 5756: 5754: 5753: 5748: 5742: 5740: 5734: 5733: 5731: 5730: 5725: 5720: 5715: 5709: 5707: 5701: 5700: 5698: 5697: 5686: 5680: 5678: 5669: 5661: 5660: 5648: 5646: 5645: 5638: 5631: 5623: 5617: 5616: 5611: 5606: 5601: 5592: 5582: 5570: 5569:External links 5567: 5566: 5565: 5560: 5555: 5548: 5545: 5543: 5542: 5523: 5518:Nobel Media AB 5505: 5484:(6): 1003–11. 5478:Molecular Cell 5464: 5409: 5380:(7): 3749–52. 5360: 5317: 5266: 5230: 5189: 5146: 5097: 5068:(8): 1178–89. 5048: 4996: 4967:(11): 581–91. 4947: 4898: 4849: 4830: 4781: 4766: 4740: 4711:(1–2): 26–34. 4691: 4648: 4615: 4564: 4535:(8): 578–588. 4515: 4508: 4490: 4439: 4396: 4375:(10): 6252–8. 4355: 4299: 4248: 4197: 4160:(7): e007198. 4139: 4100:(4): 564–574. 4079: 4044:Oncoimmunology 4030: 3979: 3948:(6): 4517–24. 3928: 3877: 3828: 3797:(5): 3307–14. 3777: 3720: 3676: 3621: 3586: 3545: 3533: 3484: 3449: 3408: 3359: 3301: 3287:10.1086/431992 3257: 3250: 3226: 3216: 3198: 3169:(3): 590–602. 3149: 3079: 3030: 2979: 2952:(11): 862–74. 2936: 2899:Molecular Cell 2885: 2833: 2782: 2738: 2697: 2658: 2596: 2537: 2506: 2475: 2432: 2422: 2384: 2330: 2295: 2254: 2210: 2159: 2116:Immune Network 2102: 2050: 2048: 2045: 1939:Eric Wieschaus 1866: 1863: 1778: 1775: 1734: 1731: 1698: 1695: 1620: 1617: 1605:blood clotting 1595: 1592: 1541: 1538: 1535: 1534: 1533: 1532: 1529: 1524: 1521: 1518: 1515: 1512: 1508: 1507: 1506: 1505: 1502: 1499: 1496: 1491: 1488: 1485: 1478: 1473: 1469: 1468: 1457: 1451: 1450: 1449: 1448: 1440: 1435: 1429: 1424: 1421: 1418: 1411: 1406: 1400: 1399: 1398: 1397: 1394: 1391: 1386: 1383: 1380: 1378: 1375: 1369: 1368: 1367: 1366: 1363: 1360: 1355: 1352: 1349: 1346: 1339: 1333: 1332: 1331: 1330: 1327: 1317: 1314: 1311: 1306: 1303: 1300: 1298: 1295: 1289: 1288: 1285: 1281: 1280: 1274: 1273: 1267: 1266: 1255: 1254: 1253: 1252: 1249: 1244: 1239: 1236: 1233: 1230: 1225: 1219: 1218: 1217: 1216: 1213: 1210: 1205: 1202: 1199: 1194: 1191: 1185: 1184: 1177: 1171: 1170: 1169: 1168: 1165: 1162: 1159: 1156: 1151: 1148: 1145: 1142: 1137: 1131: 1130: 1128: 1120: 1119: 1117: 1111: 1110: 1107: 1100: 1099: 1096: 1089: 1088: 1085: 1079: 1078: 1075: 1068: 1067: 1066: 1065: 1062: 1057: 1054: 1051: 1048: 1045: 1040: 1037: 1030: 1025: 1020: 1014: 1013: 1012: 1011: 1008: 1003: 1000: 995: 992: 983: 977: 976: 974: 970: 969: 966: 955: 954: 949: 943: 942: 937: 931: 930: 927: 920: 919: 918: 917: 912: 907: 904: 899: 896: 893: 890: 884: 878: 877: 876: 875: 870: 864: 853: 850: 844: 841: 835: 829: 828: 825: 822: 819: 816: 813: 790: 787: 724: 721: 678: 675: 664:Main article: 661: 658: 566: 559: 510: 509: 486: 484: 477: 471: 468: 456:Immunoglobulin 401:Main article: 390: 387: 217: 214: 113:sentinel cells 85: 84: 78: 77: 72: 66: 65: 62: 58: 57: 53: 52: 45: 37: 36: 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 6657: 6646: 6643: 6641: 6638: 6636: 6633: 6631: 6628: 6626: 6623: 6622: 6620: 6604: 6600: 6599:Immunophilins 6597: 6596: 6594: 6590: 6584: 6581: 6580: 6578: 6576: 6575:Secreted_PRRs 6572: 6564: 6561: 6559: 6556: 6554: 6551: 6550: 6549: 6546: 6544: 6541: 6540: 6538: 6536: 6532: 6526: 6523: 6521: 6518: 6516: 6513: 6511: 6508: 6506: 6503: 6502: 6500: 6498: 6494: 6490: 6486: 6479: 6474: 6472: 6467: 6465: 6460: 6459: 6456: 6444: 6441: 6439: 6436: 6434: 6431: 6429: 6426: 6424: 6421: 6419: 6416: 6414: 6411: 6409: 6406: 6404: 6401: 6399: 6396: 6394: 6391: 6389: 6386: 6385: 6383: 6381: 6377: 6371: 6368: 6366: 6363: 6361: 6358: 6357: 6355: 6351: 6345: 6342: 6340: 6337: 6335: 6332: 6330: 6327: 6325: 6322: 6320: 6317: 6315: 6312: 6310: 6307: 6305: 6302: 6300: 6297: 6295: 6292: 6290: 6287: 6285: 6282: 6281: 6279: 6277: 6273: 6268: 6264: 6257: 6252: 6250: 6245: 6243: 6238: 6237: 6234: 6222: 6219: 6217: 6214: 6212: 6209: 6207: 6204: 6202: 6199: 6197: 6194: 6192: 6189: 6187: 6184: 6182: 6179: 6177: 6174: 6172: 6169: 6168: 6166: 6164: 6160: 6154: 6151: 6149: 6146: 6144: 6141: 6139: 6136: 6134: 6131: 6129: 6126: 6124: 6121: 6119: 6116: 6114: 6111: 6109: 6106: 6104: 6101: 6099: 6096: 6094: 6091: 6089: 6086: 6084: 6081: 6080: 6078: 6076: 6072: 6066: 6065: 6060: 6059: 6057: 6055: 6051: 6036: 6032: 6029:(also called 6028: 6025: 6023: 6020: 6018: 6015: 6013: 6010: 6008: 6005: 6004: 6002: 5998: 5992: 5989: 5986: 5982: 5978: 5975: 5974: 5972: 5970: 5966: 5960: 5957: 5955: 5952: 5950: 5947: 5945: 5941: 5938: 5937: 5935: 5931: 5923: 5920: 5918: 5915: 5914: 5913: 5910: 5909: 5907: 5905: 5901: 5898: 5896: 5892: 5881: 5877: 5873: 5870: 5869: 5867: 5863: 5853: 5850: 5849: 5847: 5843: 5837: 5833: 5829: 5826: 5825: 5823: 5819: 5816: 5814: 5810: 5804: 5801: 5800: 5798: 5794: 5791: 5789: 5785: 5782: 5778: 5768: 5765: 5764: 5762: 5758: 5752: 5749: 5747: 5744: 5743: 5741: 5739: 5735: 5729: 5726: 5724: 5721: 5719: 5716: 5714: 5711: 5710: 5708: 5706: 5702: 5695: 5694:C-type lectin 5691: 5687: 5685: 5682: 5681: 5679: 5677: 5673: 5670: 5668: 5662: 5658: 5655: 5651: 5644: 5639: 5637: 5632: 5630: 5625: 5624: 5621: 5615: 5612: 5610: 5607: 5605: 5602: 5600: 5596: 5593: 5590: 5586: 5583: 5580: 5576: 5573: 5572: 5568: 5564: 5561: 5559: 5556: 5554: 5551: 5550: 5546: 5538: 5534: 5527: 5524: 5519: 5515: 5509: 5506: 5501: 5497: 5492: 5487: 5483: 5479: 5475: 5468: 5465: 5460: 5456: 5452: 5448: 5444: 5440: 5436: 5432: 5428: 5424: 5420: 5413: 5410: 5405: 5401: 5397: 5393: 5388: 5383: 5379: 5375: 5371: 5364: 5361: 5356: 5352: 5348: 5344: 5340: 5336: 5332: 5328: 5321: 5318: 5313: 5309: 5305: 5301: 5297: 5293: 5289: 5285: 5281: 5277: 5270: 5267: 5262: 5258: 5254: 5250: 5246: 5242: 5234: 5231: 5226: 5222: 5217: 5212: 5208: 5204: 5200: 5193: 5190: 5185: 5181: 5177: 5173: 5169: 5165: 5162:(2): 269–79. 5161: 5157: 5150: 5147: 5142: 5138: 5133: 5128: 5124: 5120: 5117:(3): 880–93. 5116: 5112: 5108: 5101: 5098: 5093: 5089: 5084: 5079: 5075: 5071: 5067: 5063: 5059: 5052: 5049: 5044: 5040: 5035: 5030: 5026: 5022: 5018: 5014: 5010: 5003: 5001: 4997: 4992: 4988: 4983: 4978: 4974: 4970: 4966: 4962: 4958: 4951: 4948: 4943: 4939: 4934: 4929: 4925: 4921: 4918:(2): 349–59. 4917: 4913: 4909: 4902: 4899: 4894: 4890: 4886: 4882: 4877: 4872: 4868: 4864: 4860: 4853: 4850: 4845: 4841: 4834: 4831: 4826: 4822: 4817: 4812: 4808: 4804: 4801:(2): 569–83. 4800: 4796: 4792: 4785: 4782: 4777: 4773: 4769: 4767:9780123748935 4763: 4759: 4755: 4751: 4744: 4741: 4736: 4732: 4727: 4722: 4718: 4714: 4710: 4706: 4702: 4695: 4692: 4687: 4683: 4679: 4675: 4671: 4667: 4664:(1–2): 50–9. 4663: 4659: 4652: 4649: 4641: 4637: 4633: 4626: 4619: 4616: 4611: 4607: 4602: 4597: 4592: 4587: 4584:(2): 323–53. 4583: 4579: 4575: 4568: 4565: 4560: 4556: 4551: 4546: 4542: 4538: 4534: 4530: 4526: 4519: 4516: 4511: 4509:3-540-00332-0 4505: 4501: 4494: 4491: 4486: 4482: 4478: 4474: 4470: 4466: 4462: 4458: 4454: 4450: 4443: 4440: 4435: 4431: 4427: 4423: 4419: 4418:10.1038/ni986 4415: 4411: 4407: 4400: 4397: 4392: 4388: 4383: 4378: 4374: 4370: 4366: 4359: 4356: 4351: 4347: 4343: 4339: 4334: 4329: 4326:(5): 373–84. 4325: 4321: 4317: 4310: 4308: 4306: 4304: 4300: 4295: 4291: 4287: 4283: 4279: 4275: 4271: 4267: 4263: 4259: 4252: 4249: 4244: 4240: 4236: 4232: 4228: 4224: 4220: 4216: 4212: 4208: 4201: 4198: 4193: 4189: 4184: 4179: 4175: 4171: 4167: 4163: 4159: 4155: 4151: 4143: 4140: 4135: 4131: 4127: 4123: 4118: 4113: 4108: 4103: 4099: 4095: 4091: 4083: 4080: 4075: 4071: 4066: 4061: 4057: 4053: 4050:(3): e23141. 4049: 4045: 4041: 4034: 4031: 4026: 4022: 4018: 4014: 4010: 4006: 4002: 3998: 3994: 3990: 3983: 3980: 3975: 3971: 3966: 3961: 3956: 3951: 3947: 3943: 3939: 3932: 3929: 3924: 3920: 3915: 3910: 3905: 3900: 3896: 3892: 3888: 3881: 3878: 3873: 3869: 3864: 3859: 3855: 3851: 3848:(1): 119–30. 3847: 3843: 3839: 3832: 3829: 3824: 3820: 3815: 3810: 3805: 3800: 3796: 3792: 3788: 3781: 3778: 3773: 3769: 3765: 3761: 3756: 3751: 3747: 3743: 3739: 3735: 3731: 3724: 3721: 3716: 3712: 3707: 3702: 3698: 3694: 3690: 3683: 3681: 3677: 3672: 3666: 3658: 3654: 3649: 3644: 3640: 3636: 3632: 3625: 3622: 3617: 3613: 3609: 3605: 3601: 3597: 3590: 3587: 3582: 3578: 3573: 3568: 3564: 3560: 3556: 3549: 3546: 3543: 3537: 3534: 3529: 3525: 3520: 3515: 3511: 3507: 3504:(5): 685–94. 3503: 3499: 3495: 3488: 3485: 3480: 3476: 3472: 3468: 3464: 3460: 3453: 3450: 3445: 3441: 3436: 3431: 3428:(2): 966–72. 3427: 3423: 3419: 3412: 3409: 3404: 3400: 3396: 3392: 3387: 3382: 3378: 3374: 3370: 3363: 3360: 3355: 3351: 3346: 3341: 3336: 3335:10.1186/ar567 3331: 3327: 3323: 3319: 3312: 3310: 3308: 3306: 3302: 3297: 3293: 3288: 3283: 3279: 3275: 3271: 3264: 3262: 3258: 3253: 3247: 3243: 3235: 3233: 3231: 3227: 3223: 3219: 3213: 3209: 3202: 3199: 3194: 3190: 3185: 3180: 3176: 3172: 3168: 3164: 3160: 3153: 3150: 3145: 3141: 3136: 3131: 3127: 3123: 3118: 3113: 3109: 3105: 3101: 3097: 3093: 3086: 3084: 3080: 3075: 3071: 3066: 3061: 3057: 3053: 3050:(3): 547–58. 3049: 3045: 3041: 3034: 3031: 3026: 3022: 3017: 3012: 3007: 3002: 2998: 2994: 2990: 2983: 2980: 2975: 2971: 2967: 2963: 2959: 2955: 2951: 2947: 2940: 2937: 2932: 2928: 2923: 2918: 2913: 2908: 2904: 2900: 2896: 2889: 2886: 2881: 2877: 2873: 2869: 2864: 2859: 2856:(6): 973–83. 2855: 2851: 2847: 2840: 2838: 2834: 2829: 2825: 2820: 2815: 2810: 2805: 2801: 2797: 2793: 2786: 2783: 2778: 2774: 2769: 2764: 2760: 2756: 2752: 2745: 2743: 2739: 2734: 2730: 2725: 2720: 2717:(2): 649–56. 2716: 2712: 2708: 2701: 2698: 2693: 2689: 2685: 2681: 2677: 2673: 2669: 2662: 2659: 2654: 2650: 2645: 2640: 2635: 2630: 2626: 2622: 2618: 2614: 2610: 2603: 2601: 2597: 2592: 2588: 2583: 2578: 2573: 2568: 2564: 2560: 2556: 2552: 2548: 2541: 2538: 2533: 2529: 2525: 2521: 2517: 2510: 2507: 2502: 2498: 2495:(3): 362–71. 2494: 2490: 2486: 2479: 2476: 2471: 2467: 2463: 2459: 2455: 2451: 2448:(9): 1081–8. 2447: 2443: 2436: 2433: 2426: 2423: 2418: 2414: 2410: 2406: 2402: 2398: 2391: 2389: 2385: 2380: 2376: 2371: 2370:10.1038/41131 2366: 2362: 2358: 2354: 2350: 2346: 2339: 2337: 2335: 2331: 2326: 2322: 2318: 2314: 2310: 2306: 2299: 2296: 2291: 2287: 2282: 2277: 2273: 2269: 2265: 2258: 2255: 2250: 2246: 2241: 2236: 2233:(6): 1085–7. 2232: 2228: 2224: 2217: 2215: 2211: 2206: 2202: 2197: 2192: 2188: 2184: 2180: 2176: 2175: 2170: 2163: 2160: 2155: 2151: 2146: 2141: 2137: 2133: 2129: 2125: 2121: 2117: 2113: 2106: 2103: 2098: 2094: 2089: 2084: 2079: 2074: 2070: 2066: 2062: 2055: 2052: 2046: 2044: 2041: 2039: 2035: 2030: 2028: 2019: 2015: 2013: 2009: 2005: 2001: 1997: 1992: 1990: 1989:interleukin-1 1985: 1980: 1978: 1974: 1971:was found by 1970: 1966: 1961: 1959: 1952: 1948: 1944: 1943:embryogenesis 1940: 1936: 1932: 1931: 1925: 1920: 1915: 1913: 1912:immune system 1909: 1905: 1901: 1897: 1893: 1889: 1885: 1881: 1877: 1873: 1864: 1862: 1858: 1855: 1851: 1847: 1843: 1839: 1835: 1831: 1827: 1823: 1819: 1815: 1811: 1807: 1804: 1802: 1798: 1793: 1791: 1787: 1783: 1776: 1774: 1770: 1768: 1762: 1760: 1756: 1752: 1748: 1744: 1740: 1732: 1730: 1728: 1724: 1720: 1716: 1712: 1708: 1704: 1696: 1694: 1691: 1689: 1685: 1681: 1676: 1671: 1669: 1665: 1661: 1656: 1654: 1650: 1646: 1642: 1639:, TLR2 forms 1638: 1634: 1625: 1618: 1616: 1614: 1610: 1606: 1603:(involved in 1602: 1593: 1591: 1588: 1584: 1580: 1576: 1572: 1568: 1564: 1560: 1557:cell-surface 1556: 1546: 1539: 1530: 1527: 1526: 1525: 1522: 1519: 1516: 1513: 1510: 1509: 1503: 1500: 1497: 1494: 1493: 1492: 1489: 1486: 1484: 1483: 1479: 1477: 1474: 1471: 1470: 1466: 1462: 1458: 1456: 1453: 1452: 1447: 1444: 1441: 1439: 1436: 1433: 1430: 1427: 1426: 1417: 1416: 1412: 1410: 1407: 1405: 1401: 1395: 1392: 1389: 1388: 1387: 1384: 1381: 1379: 1376: 1374: 1371: 1370: 1365:B lymphocytes 1364: 1361: 1358: 1357: 1356: 1353: 1350: 1347: 1344: 1341:unmethylated 1340: 1338: 1335: 1334: 1328: 1326: 1322: 1318: 1315: 1312: 1309: 1308: 1307: 1304: 1301: 1299: 1296: 1294: 1291: 1290: 1286: 1283: 1282: 1279: 1276: 1275: 1272: 1269: 1268: 1264: 1260: 1257: 1256: 1251:B lymphocytes 1250: 1248: 1245: 1242: 1241: 1229: 1226: 1224: 1220: 1215:B lymphocytes 1214: 1211: 1208: 1207: 1206: 1203: 1200: 1198: 1195: 1192: 1190: 1186: 1183: 1182: 1178: 1176: 1173: 1172: 1167:B lymphocytes 1166: 1163: 1160: 1157: 1154: 1153: 1143: 1141: 1138: 1136: 1132: 1129: 1126: 1122: 1121: 1118: 1116: 1113: 1112: 1108: 1105: 1102: 1101: 1097: 1094: 1091: 1090: 1086: 1084: 1081: 1080: 1076: 1074: 1070: 1069: 1063: 1061: 1058: 1055: 1052: 1049: 1046: 1043: 1042: 1035: 1029: 1026: 1024: 1021: 1019: 1015: 1010:B lymphocytes 1009: 1006: 1005: 1004: 1001: 999: 996: 993: 991: 987: 984: 982: 979: 978: 975: 972: 971: 967: 964: 960: 957: 956: 953: 950: 948: 945: 944: 941: 938: 936: 933: 932: 928: 926: 922: 921: 916: 913: 911: 908: 905: 902: 901: 891: 889: 885: 883: 879: 874: 873:B lymphocytes 871: 869: 865: 863: 859: 856: 855: 854: 851: 848: 845: 842: 840: 836: 834: 831: 830: 826: 823: 820: 817: 814: 811: 810: 807: 805: 801: 797: 788: 786: 784: 783:disease model 780: 779:knockout mice 776: 775:typhoid fever 772: 770: 765: 760: 758: 754: 750: 746: 742: 741: 736: 735: 730: 722: 720: 717: 712: 708: 704: 700: 696: 691: 687: 683: 676: 674: 672: 667: 659: 657: 655: 652:family and 6 651: 647: 643: 641: 637: 633: 629: 625: 621: 617: 613: 610:, fungi, and 609: 604: 602: 597: 593: 589: 585: 584: 576: 571: 564: 560: 558: 555: 553: 549: 545: 541: 535: 531: 529: 528:immune system 525: 521: 520:invertebrates 517: 506: 503:December 2021 496: 490: 487:This section 485: 481: 476: 475: 469: 467: 465: 461: 457: 453: 449: 445: 441: 436: 434: 430: 426: 422: 418: 414: 410: 404: 395: 388: 386: 384: 380: 375: 373: 369: 365: 362:presented to 361: 357: 353: 352:proliferation 349: 345: 341: 337: 333: 329: 325: 321: 317: 313: 309: 305: 301: 297: 292: 290: 289:innate immune 286: 282: 278: 273: 271: 267: 263: 259: 255: 251: 247: 243: 239: 235: 231: 227: 223: 215: 213: 211: 206: 204: 203:nucleic acids 200: 197: 196:intracellular 193: 192:cell membrane 189: 185: 181: 177: 173: 169: 165: 161: 157: 153: 149: 145: 141: 137: 133: 130: 126: 122: 118: 114: 110: 107: 103: 99: 95: 91: 83: 79: 76: 73: 71: 67: 63: 59: 54: 49: 43: 38: 33: 30: 19: 6635:LRR proteins 6504: 6343: 6338: 6275: 6266: 6061: 5969:Co-receptors 5922:MHC class II 5536: 5526: 5517: 5508: 5481: 5477: 5467: 5426: 5422: 5412: 5377: 5373: 5363: 5330: 5326: 5320: 5279: 5275: 5269: 5247:(3): 486–8. 5244: 5240: 5233: 5209:(1): 27–35. 5206: 5203:DNA Research 5202: 5192: 5159: 5155: 5149: 5114: 5111:Neuroscience 5110: 5100: 5065: 5061: 5051: 5016: 5012: 4964: 4960: 4950: 4915: 4911: 4901: 4869:(2): 454–9. 4866: 4862: 4852: 4843: 4833: 4798: 4795:Neuroscience 4794: 4784: 4749: 4743: 4708: 4704: 4694: 4661: 4657: 4651: 4640:the original 4635: 4631: 4618: 4581: 4577: 4567: 4532: 4528: 4518: 4499: 4493: 4452: 4448: 4442: 4409: 4405: 4399: 4372: 4368: 4358: 4323: 4319: 4261: 4257: 4251: 4210: 4206: 4200: 4157: 4153: 4142: 4097: 4093: 4082: 4047: 4043: 4033: 3992: 3988: 3982: 3945: 3941: 3931: 3894: 3890: 3880: 3845: 3841: 3831: 3794: 3790: 3780: 3737: 3733: 3723: 3696: 3692: 3665:cite journal 3638: 3634: 3624: 3599: 3595: 3589: 3562: 3558: 3548: 3536: 3501: 3497: 3487: 3462: 3458: 3452: 3425: 3421: 3411: 3379:(6): 471–5. 3376: 3372: 3362: 3325: 3321: 3277: 3273: 3241: 3221: 3207: 3201: 3166: 3162: 3152: 3099: 3095: 3047: 3043: 3033: 2996: 2992: 2989:"Drosophila" 2982: 2949: 2945: 2939: 2902: 2898: 2888: 2853: 2849: 2799: 2795: 2785: 2758: 2755:Cell Reports 2754: 2714: 2710: 2700: 2675: 2671: 2661: 2616: 2612: 2554: 2550: 2540: 2526:(3): 372–8. 2523: 2519: 2509: 2492: 2488: 2478: 2445: 2441: 2435: 2425: 2403:(4): 521–8. 2400: 2396: 2352: 2348: 2308: 2304: 2298: 2274:(1): 67–73. 2271: 2267: 2257: 2230: 2226: 2178: 2172: 2162: 2119: 2115: 2105: 2068: 2064: 2054: 2042: 2037: 2033: 2031: 2027:Shizuo Akira 2024: 1993: 1983: 1981: 1968: 1928: 1918: 1916: 1900:lipopeptides 1868: 1859: 1826:hyperalgesia 1808: 1805: 1794: 1780: 1771: 1763: 1736: 1700: 1692: 1672: 1660:nucleic acid 1657: 1641:heterodimers 1630: 1597: 1563:lipoproteins 1551: 1520:MyD88, TAK-1 1480: 1464: 1460: 1413: 1385:cell surface 1287:RNA viruses 1204:cell surface 1179: 1150:cell surface 1039:cell surface 925:proteolipids 898:cell surface 866:a subset of 852:cell surface 839:lipopeptides 792: 768: 763: 761: 756: 748: 744: 738: 732: 726: 711:pluripotency 680: 669: 653: 649: 645: 644: 619: 605: 581: 579: 562: 556: 536: 532: 513: 500: 488: 440:interleukins 437: 406: 376: 364:CD4+ T cells 356:phagocytosed 350:production, 340:inflammatory 293: 281:vaccinations 274: 219: 207: 93: 89: 88: 29: 6603:Cyclophilin 5917:MHC class I 5813:Co-receptor 5676:Epsilon (ε) 5667:Fc receptor 5019:(1): 20–9. 3465:: 151–160. 2678:: 697–743. 2430:PMC8452412. 2311:(1): 1–14. 2038:Arabidopsis 1956:Das ist ja 1876:Robert Koch 1786:dermatology 1767:pleiotropic 1504:macrophages 1271:bropirimine 1109:host cells 1098:host cells 1087:host cells 1047:neutrophils 963:Beta-glucan 906:neutrophils 888:glycolipids 862:macrophages 827:Cell types 518:as well as 516:vertebrates 444:macrophages 433:superfamily 389:Superfamily 372:interferons 304:immune cell 270:fibroblasts 246:macrophages 136:immune cell 117:macrophages 104:. They are 82:PIRSF037595 56:Identifiers 46:The curved 6619:Categories 5821:stimulate: 4117:2066/89493 3242:Immunology 2802:(1): 228. 2122:(3): e21. 1984:Drosophila 1921:receptor ( 1850:drug abuse 1790:resiquimod 1637:homodimers 1601:fibrinogen 1465:Salmonella 1459:Bacteria ( 1446:epithelium 1316:Mast cells 1278:resiquimod 1265:analogue) 1259:loxoribine 1212:Mast cells 1197:Mycoplasma 1083:fibrinogen 1053:Mast cells 1032:MyD88/MAL/ 952:Host cells 915:Mast cells 821:Adapter(s) 804:cell types 800:leucocytes 769:Salmonella 764:Salmonella 757:Salmonella 745:Toxoplasma 646:Drosophila 640:drosomycin 596:haemolymph 563:Drosophila 544:retrovirus 316:downstream 262:epithelial 240:including 238:leukocytes 70:Membranome 6583:Collectin 5760:Secretory 5705:Gamma (γ) 4174:2051-1426 4126:1528-0020 4025:206540638 3126:1932-6203 2136:1598-2629 1994:In 1997, 1904:flagellin 1880:endotoxin 1865:Discovery 1846:addiction 1842:tolerance 1830:allodynia 1782:Imiquimod 1664:cytokines 1619:Signaling 1555:bacterial 1455:Flagellin 1263:guanosine 1201:MyD88/MAL 1106:fragments 1095:fragments 895:MyD88/MAL 886:multiple 858:monocytes 815:Ligand(s) 802:or other 753:flagellin 575:fruit fly 495:talk page 464:cytosolic 448:monocytes 413:pathogens 383:Flagellin 368:apoptosis 285:molecular 234:membranes 226:pathogens 222:molecules 210:toll gene 109:receptors 6108:KIR2DL5B 6103:KIR2DL5A 5845:inhibit: 5728:Neonatal 5547:See also 5537:The Star 5500:10911994 5459:10548988 5396:10201887 5241:Genomics 5184:19439405 5141:20178837 5092:18599265 5043:18662331 4991:19762094 4942:20974246 4893:12295407 4885:21081778 4825:19833175 4776:19607972 4735:20615556 4678:15836969 4610:26344622 4578:Vaccines 4559:31015631 4485:16163262 4477:12447441 4434:13016860 4426:14556004 4391:17475853 4350:39414949 4342:20404851 4286:11130078 4243:15608748 4235:12872135 4192:37487664 4183:10373699 4134:20424184 4074:23802068 4017:22821982 3974:21131352 3923:19077284 3872:23246311 3842:Immunity 3823:21097503 3772:34165967 3764:15860593 3715:24198280 3657:11970999 3616:11267672 3581:20348427 3528:24861338 3479:28965571 3444:10623846 3403:30443009 3395:17637697 3354:12110131 3296:16237641 3193:23101627 3144:26859749 3096:PLOS ONE 3074:23101625 3025:31088910 2974:11104900 2966:17948019 2931:29452635 2880:10736743 2828:27776480 2777:31018123 2733:21209287 2692:17201680 2653:15976025 2591:14993594 2532:11022120 2501:11022119 2470:12517461 2462:17704767 2417:23345392 2325:15585605 2290:17212934 2249:15923538 2205:20860480 2154:32655969 2097:24032031 1611:(HSPs), 1571:flagella 1476:Profilin 1409:Profilin 1175:Profilin 1144:Bacteria 1123:Various 1071:several 994:viruses 990:poly I:C 824:Location 812:Receptor 565:immunity 561:TLRs in 552:Takifugu 360:antigens 348:cytokine 312:proteins 216:Function 199:vesicles 125:microbes 115:such as 98:proteins 6153:KIR3DS1 6148:KIR3DL3 6143:KIR3DL2 6138:KIR3DL1 6133:KIR2DS5 6128:KIR2DS4 6123:KIR2DS3 6118:KIR2DS2 6113:KIR2DS1 6098:KIR2DL4 6093:KIR2DL3 6088:KIR2DL2 6083:KIR2DL1 6027:ζ-chain 5904:Ligands 5895:T cells 5788:B cells 5723:FcγRIII 5451:8525370 5431:Bibcode 5423:Science 5404:7419784 5355:9851930 5335:Bibcode 5327:Science 5312:1700458 5304:1851964 5284:Bibcode 5261:8838819 5225:7584026 5176:2449285 5132:2854318 5083:2783238 5034:2588470 4982:2783351 4933:3025088 4816:2795035 4726:2942958 4686:7286123 4601:4494261 4550:6192054 4457:Bibcode 4294:4405163 4266:Bibcode 4215:Bibcode 4065:3661153 3997:Bibcode 3989:Science 3965:3039399 3914:2631477 3863:3601573 3814:3030336 3742:Bibcode 3734:Science 3519:4141014 3345:3240143 3184:3500584 3135:4747465 3104:Bibcode 3065:3506423 3016:6664172 2922:5823974 2872:8808632 2819:5078906 2644:1172252 2621:Bibcode 2559:Bibcode 2379:9237759 2357:Bibcode 2196:3045535 2145:7327153 2088:3759294 2071:: 248. 2040:FLS2). 1951:ventral 1838:opioids 1814:opioids 1561:(LPS), 1540:Ligands 1495:Neurons 1461:E. coli 1390:B cells 1382:unknown 1321:Crohn's 959:zymosan 749:E. coli 743:. 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