Toll-like receptor

559: 1534: 1613: 2007: 1574:; 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, 31: 1899:) 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 469: 1604:, 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. 383: 1542:(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 611:). 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 1761:

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

575:. 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 526:

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

535:. 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 1845:

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

1718:. 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. 4076:

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).

5188:"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" 5227:

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

447:(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 1887:(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 4393:

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.

5629: 1971:

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".

5096:"Possible involvement of toll-like receptor 4/myeloid differentiation factor-2 activity of opioid inactive isomers causes spinal proinflammation and related behavioral consequences" 5045:

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".

1781:, is a TLR7 and TLR8 agonist. Recently, resiquimod has been explored as an agent for cancer immunotherapy, acting through stimulation of tumor-associated macrophages. 3620:"Quantitative expression of toll-like receptor 1-10 mRNA in cellular subsets of human peripheral blood mononuclear cells and sensitivity to CpG oligodeoxynucleotides" 5622: 4194:

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".

179:. 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 6464: 424:, 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. 1813:, 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, 1750:. Meanwhile, activation of RIPK1 causes the polyubiquitination and activation of TAK1 and NFκB transcription in the same manner as the MyD88-dependent pathway. 579:. The toll pathway is similar to mammalian TLR signalling, but unlike mammalian TLRs, toll is not activated directly by pathogen-associated molecular patterns ( 527:

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

619:), phosphorylated Cactus is polyubiquitylated and degraded, allowing nuclear translocation of DIF (dorsal-related immunity factor; a homologue of mammalian 5615: 1651:

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

1953: 4354:"TLR2 is constitutively expressed within the kidney and participates in ischemic renal injury through both MyD88-dependent and -independent pathways" 2429:

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).

3678:"Identification of TLR10 as a key mediator of the inflammatory response to Listeria monocytogenes in intestinal epithelial cells and macrophages" 546:

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".

5359:"Cutting edge: Toll-like receptor 4 (TLR4)-deficient mice are hyporesponsive to lipopolysaccharide: evidence for TLR4 as the Lps gene product" 3199:(8th ed.). Philadelphia Baltimore New York London Buenos Aires Hong Kong Sydney Tokyo: Wolters Kluwer/Lippincott Williams & Wilkins. 6010: 4690:"Dual regulation of mu opioid receptors in SK-N-SH neuroblastoma cells by morphine and interleukin-1β: evidence for opioid-immune crosstalk" 6015: 3583:

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

1247: 6457: 4079:"Commonly used prophylactic vaccines as an alternative for synthetically produced TLR ligands to mature monocyte-derived dendritic cells" 3211:

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".

2884:"The Circulating Protease Persephone Is an Immune Sensor for Microbial Proteolytic Activities Upstream of the Drosophila Toll Pathway" 6141: 6121: 6111: 6106: 6076: 5521: 4613: 1662:

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

6184: 6063: 5597: 6151: 5094:

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

4780:"Evidence that intrathecal morphine-3-glucuronide may cause pain enhancement via toll-like receptor 4/MD-2 and interleukin-1beta" 2504: 2473: 536: 176: 172: 2384:

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

6485: 6450: 5937: 4514:"TLR7/8-agonist-loaded nanoparticles promote the polarization of tumour-associated macrophages to enhance cancer immunotherapy" 5592: 4848:"Etanercept restores the antinociceptive effect of morphine and suppresses spinal neuroinflammation in morphine-tolerant rats" 4614:"Toll like Receptor 7 Messenger Ribonucleic Acid Expression Levels in Dairy Animals in an Outbreak of Foot-and-mouth disease" 3407:"Lipopolysaccharide activates distinct signaling pathways in intestinal epithelial cell lines expressing Toll-like receptors" 3238: 3204: 405: 355:. In the case of a viral factor, the infected cell may shut off its protein synthesis and may undergo programmed cell death ( 198: 6523: 2835:"The dorsoventral regulatory gene cassette spätzle/Toll/cactus controls the potent antifungal response in Drosophila adults" 615:, Tube, and Pelle (the orthologue of mammalian IRAK). Signal from TICS is then transduced to Cactus (homologue of mammalian 1694:. Ligand binding and conformational change that occurs in the receptor recruits the adaptor protein MyD88, a member of the 343:, and survival, while others lead to greater adaptive immunity. If the ligand is a bacterial factor, the pathogen might be 1795:

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

6563: 6235: 4998:"Non-stereoselective reversal of neuropathic pain by naloxone and naltrexone: involvement of toll-like receptor 4 (TLR4)" 1860: 1537:

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

409: 6368: 6353: 5607: 3233:. Lippincott's Illustrated reviews. Philadelphia: Wolters Kluwer Health/Lippincott Williams & Wilkins. p. 17. 1641: 448: 284: 2536:"Toll-like receptors 9 and 3 as essential components of innate immune defense against mouse cytomegalovirus infection" 5755: 4754: 4496: 4029:"Bacteria evade immune recognition via TLR13 and binding of their 23S rRNA by MLS antibiotics by the same mechanisms" 5900: 4828: 1691: 4897:"An IL-1 receptor antagonist blocks a morphine-induced attenuation of locomotor recovery after spinal cord injury" 6613: 6508: 5463:"FLS2: an LRR receptor-like kinase involved in the perception of the bacterial elicitor flagellin in Arabidopsis" 421: 187: 1980:(IL-1) receptor, also had homology to drosophila toll; the cytoplasmic portions of both molecules were similar. 6228: 1682:

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

1923: 543:

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

5502: 4946:"The "toll" of opioid-induced glial activation: improving the clinical efficacy of opioids by targeting glia" 1022: 512: 6633: 6628: 6477: 3659: 3481:

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

589: 397: 94: 2882:

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.

6052: 1965: 1806: 1235: 583:). Its receptor ectodomain recognizes the cleaved form of the cytokine spätzle, which is secreted in the 296: 3876:"Expression and distribution of Toll-like receptors 11-13 in the brain during murine neurocysticercosis" 5642: 3827:"Recognition of profilin by Toll-like receptor 12 is critical for host resistance to Toxoplasma gondii" 3618:

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

3483:"Toxoplasma gondii- derived profilin triggers human toll-like receptor 5-dependent cytokine production" 1952:" ("That's amazing!"), in reference to the underdeveloped ventral portion of a fruit fly larva. It was 6442: 5573: 3259:"The role of Toll-like receptors in the regulation of neutrophil migration, activation, and apoptosis" 4778:

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

391: 3776:"UNC93B1 is essential for TLR11 activation and IL-12-dependent host resistance to Toxoplasma gondii" 6618: 5583: 5577: 5567: 4139:"Pattern recognition receptor agonists in pathogen vaccines mediate antitumor T-cell cross-priming" 2505:"Cloning and characterization of a sub-family of human toll-like receptors: hTLR7, hTLR8 and hTLR9" 1859:

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

1331: 3717:

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

3081:"Toll-Like Receptor 11 (TLR11) Interacts with Flagellin and Profilin through Disparate Mechanisms" 2014:

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

1742:, which creates a branch in the signaling pathway. The TRIF/TBK1 signaling complex phosphorylates 6503: 2740:"More Than Black or White: Melanization and Toll Share Regulatory Serine Proteases in Drosophila" 2101:"TLR10 and Its Unique Anti-Inflammatory Properties and Potential Use as a Target in Therapeutics" 1918: 571: 563: 483: 304: 272:

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

97: 5563: 1992: 1872: 1842: 1016: 928: 898: 624: 603:. First, the Spätzle processing enzyme (SPE) is activated in response to infection and cleaves 596: 417: 184: 5664: 6623: 6358: 6322: 6317: 5816: 5638: 5602: 4512:

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

4305:"The role of pattern-recognition receptors in innate immunity: update on Toll-like receptors" 3653: 3079:

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

2672: 1735: 1714:, which in turn polyubiquinates the protein TAK1, as well as itself to facilitate binding to 1575: 1048: 770:, on the other hand, are efficiently infected. As a result, this knockout mouse can act as a 427:

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

408:(PAMPs). In addition to the recognition of exogenous PAMPs, TLRs can also bind to endogenous 320: 5726: 5693: 2158:"Type B coxsackieviruses and their interactions with the innate and adaptive immune systems" 323:) that further amplify the signal and ultimately lead to the upregulation or suppression of 6473: 6312: 6307: 6302: 6297: 6292: 6287: 6282: 6277: 6272: 6220: 5716: 5587: 5546: 5419: 5323: 5272: 4445: 4352:

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

4254: 4203: 3985: 3730: 3092: 2609: 2547: 2345: 1883:. In the decades that followed, endotoxin was chemically characterized and identified as a 1785: 1616:

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

277: 90: 3825:

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).

2334:"A human homologue of the Drosophila Toll protein signals activation of adaptive immunity" 558: 70: 8: 6536: 6513: 6251: 5551: 4628: 2162: 1876: 1128: 974: 238: 58: 36: 5637: 5423: 5327: 5276: 4449: 4258: 4207: 4171: 4138: 3989: 3734: 3544:"Human TLRs 10 and 1 share common mechanisms of innate immune sensing but not signaling" 3405:

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

3096: 2613: 2551: 2349: 673:

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

569:

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

511:. Molecular building blocks of the TLRs are represented in bacteria and in plants, and 482:

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

5443: 5388: 5296: 5168: 5120: 5111: 5095: 5071: 5046: 5022: 4997: 4970: 4945: 4921: 4896: 4877: 4804: 4795: 4779: 4714: 4689: 4670: 4589: 4562: 4538: 4513: 4469: 4418: 4334: 4278: 4227: 4053: 4028: 4009: 3953: 3926: 3902: 3875: 3851: 3826: 3802: 3775: 3756: 3507: 3482: 3387: 3333: 3306: 3172: 3147: 3123: 3080: 3053: 3028: 3004: 2977: 2958: 2910: 2883: 2864: 2807: 2781:"Immune genes and divergent antimicrobial peptides in flies of the subgenus Drosophila" 2780: 2632: 2597: 2454: 2228: 2211: 2184: 2157: 2133: 2076: 2049: 2000: 1884: 1633: 1597: 1547: 1313: 1309: 1061: 1011: 413: 374:, a TLR5 ligand, induces cytokine secretion on interacting with TLR5 on human T cells. 359:). Immune cells that have detected a virus may also release anti-viral factors such as 5479: 5462: 5357:

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

5186:

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

4746: 3596: 2851: 2570: 2535: 2474:"Three novel mammalian toll-like receptors: gene structure, expression, and evolution" 6531: 6042: 5541: 5484: 5435: 5380: 5339: 5288: 5245: 5209: 5160: 5156: 5125: 5076: 5027: 5013: 4975: 4926: 4869: 4809: 4760: 4750: 4719: 4662: 4594: 4543: 4492: 4461: 4410: 4375: 4326: 4270: 4219: 4176: 4158: 4118: 4110: 4058: 4013: 4001: 3958: 3907: 3856: 3807: 3748: 3699: 3641: 3600: 3565: 3512: 3463: 3428: 3379: 3338: 3280: 3234: 3200: 3177: 3128: 3110: 3058: 3009: 2950: 2915: 2856: 2812: 2761: 2717: 2676: 2637: 2596:

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

2575: 2516: 2485: 2446: 2401: 2363: 2309: 2274: 2233: 2189: 2138: 2120: 2081: 2050:"Sweeten PAMPs: Role of Sugar Complexed PAMPs in Innate Immunity and Vaccine Biology" 1961: 1747: 1656: 1555: 1470: 1403: 1169: 923: 728: 600: 254: 5447: 5172: 4895:

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

4881: 4473: 4422: 4338: 4231: 3760: 3391: 2962: 2868: 2458: 1784:

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)

604: 6498: 5995: 5474: 5427: 5392: 5370: 5331: 5300: 5280: 5237: 5199: 5152: 5115: 5107: 5066: 5058: 5017: 5009: 4965: 4957: 4916: 4908: 4859: 4799: 4791: 4742: 4709: 4705: 4701: 4674: 4654: 4584: 4574: 4533: 4525: 4453: 4402: 4365: 4316: 4282: 4262: 4211: 4166: 4150: 4100: 4090: 4048: 4040: 3993: 3948: 3938: 3897: 3887: 3846: 3838: 3797: 3787: 3738: 3689: 3631: 3592: 3555: 3502: 3494: 3455: 3418: 3369: 3328: 3318: 3270: 3167: 3159: 3118: 3100: 3048: 3040: 3027:

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

2999: 2989: 2942: 2905: 2895: 2846: 2802: 2792: 2751: 2707: 2668: 2627: 2617: 2565: 2555: 2534:

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

2438: 2393: 2353: 2301: 2264: 2223: 2179: 2171: 2128: 2112: 2071: 2061: 1996: 1988: 1935: 1216: 784: 756: 117: 5431: 5408:"A receptor kinase-like protein encoded by the rice disease resistance gene, Xa21" 5406:

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

5335: 3146:

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

1805:. Its activation leads to downstream release of inflammatory modulators including 5928: 5791: 5645: 4864: 4847: 4370: 4353: 4095: 4078: 3842: 3105: 2900: 2756: 2739: 1984: 1431: 1092: 1081: 332: 308: 218: 5375: 5358: 3636: 3619: 3459: 2025:

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

63: 5701: 5407: 4961: 4658: 3530: 3423: 3406: 3163: 3044: 3029:"Activation of innate immunity is required for efficient nuclear reprogramming" 2602:

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

2540:

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

2269: 2252: 1927: 1880: 1830: 1789: 1593: 856: 444: 440: 367: 230: 109: 5062: 4912: 4846:

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

4529: 3925:

Shi Z, Cai Z, Sanchez A, Zhang T, Wen S, Wang J, et al. (February 2011).

2833:

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

2797: 1612: 1503:

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

6607: 6587: 5682: 5047:"Proinflammatory cytokines oppose opioid-induced acute and chronic analgesia" 4162: 4154: 4114: 3694: 3677: 3560: 3543: 3114: 2994: 2712: 2695: 2124: 2066: 1977: 1931: 1900: 1621: 1533: 771: 767: 763: 516: 352: 222: 180: 113: 101: 5204: 5187: 4579: 3997: 3943: 3792: 3743: 3542:

Guan Y, Ranoa DR, Jiang S, Mutha SK, Li X, Baudry J, Tapping RI (May 2010).

2622: 2560: 2305: 2292:

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

2100: 5910: 5488: 5384: 5241: 5129: 5080: 5031: 4979: 4930: 4873: 4813: 4764: 4739:

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

4723: 4666: 4598: 4547: 4465: 4414: 4379: 4330: 4274: 4223: 4180: 4122: 4062: 4005: 3962: 3911: 3892: 3860: 3811: 3752: 3703: 3645: 3604: 3569: 3516: 3467: 3432: 3383: 3374: 3357: 3342: 3284: 3181: 3132: 3062: 3013: 2954: 2919: 2834: 2816: 2765: 2721: 2680: 2656: 2641: 2579: 2520: 2489: 2450: 2405: 2313: 2278: 2237: 2193: 2142: 2116: 2085: 2015: 1968:, which it achieved by activating the synthesis of antimicrobial peptides. 1946: 1814: 1648: 861: 699: 528: 508: 428: 344: 328: 197:

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

191: 5439: 5343: 5292: 5249: 5213: 5164: 4563:"Immune Adjuvant Effect of Molecularly-defined Toll-Like Receptor Ligands" 3719:"TLR11 activation of dendritic cells by a protozoan profilin-like protein" 2976:

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

2860: 2367: 1942:

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

6591: 5957: 5905: 5801: 5739: 5655: 1912: 1888: 1864: 1774: 1755: 1726:

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

1629: 1551: 1259: 951: 913: 827: 576: 340: 292: 269: 258: 234: 124: 116:. Once these microbes have reached physical barriers such as the skin or 4457: 4215: 4075: 3480: 2397: 6472: 4741:. International Review of Neurobiology. Vol. 85. pp. 207–19. 4044: 3927:"A novel Toll-like receptor that recognizes vesicular stomatitis virus" 2175: 1838: 1778: 1647:

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

1589: 1434: 1266: 1185: 1071: 903: 876: 850: 792: 788: 628: 584: 532: 504: 432: 360: 250: 105: 4135: 4105: 3498: 3358:"Regulating B-cell activation and survival in response to TLR signals" 1964:

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

287:(proteins that mediate other protein-protein interactions) within the 6571: 5969: 5284: 4266: 2099:

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

1892: 1868: 1834: 1818: 1770: 1652: 1625: 1543: 1443: 1251: 940: 741: 404:

but distinguishable from host molecules, collectively referred to as

371: 356: 280:

responses and the development of antigen-specific acquired immunity.

273: 226: 4321: 4304: 3718: 2946: 2738:

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

2442: 662:

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

6019: 5973: 3585:

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

3275: 3258: 1567: 1559: 1464: 1397: 1163: 978: 846: 720:, but mice cells do. Mouse-specific TLR11 recognizes uropathogenic 540: 436: 401: 336: 300: 214: 210: 5672: 5263:

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

4406: 3323: 2358: 2333: 2006: 1850:

analgesic drugs without having any affinity for μ-opioid receptor

620: 531:, but appears to have been damaged at some point in the past by a 6136: 6131: 6126: 6116: 6101: 6086: 6081: 6071: 4612:

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

3716: 1939: 1801:

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

947: 645:

family genes that interact with each other to differing degrees.

452: 348: 288: 265: 86: 30: 4737:

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

2331: 468: 6426: 6416: 6401: 6209: 6204: 6199: 6194: 6189: 6179: 6174: 6169: 6164: 6159: 5892: 5883: 5776: 2832: 2379: 2377: 1930:

and colleagues. It was known for its developmental function in

1826: 1802: 1426: 1392: 1361: 1113: 1103: 722: 416:(HSPs) or plasma membrane constituents. TLRs together with the 246: 242: 120: 112:, that recognize structurally conserved molecules derived from 5593:

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

4944:

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

4611: 4511: 3617: 1995:. TLR 4 function as an LPS sensing receptor was discovered by 1746:

allowing its translocation into the nucleus and production of

616: 6541: 6431: 6421: 6406: 6391: 6386: 6381: 6376: 5864: 5860: 4736: 3975: 3307:"The instructive role of dendritic cells on T-cell responses" 3228: 1739: 1727: 1711: 1707: 1703: 1699: 1695: 1672: 1668: 1601: 1420: 1281: 1211: 1177: 1123: 1006: 969: 935: 870: 835: 821: 717: 704: 695: 674: 612: 580: 324: 316: 312: 168: 164: 6250: 3774:

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

2374: 2098: 1792:

in 2017, which contains a monophosphoryl lipid A component.

595:

The toll pathway is activated by different stimuli, such as

303:

are then responsible for the subsequent activation of other

221:

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

6551: 6546: 6411: 6396: 6348: 6005: 6000: 5947: 5942: 5932: 5868: 5840: 5824: 5820: 5734: 5711: 5706: 5678: 5093: 4777: 3773: 3404: 2932: 2253:"[Innate immunity: structure and function of TLRs]" 1822: 1810: 1798: 1743: 1731: 1715: 1710:. IRAK kinases then phosphorylate and activate the protein 1676: 1644:) are known to facilitate the presentation of LPS to MD-2. 1637: 1325: 986: 691: 687: 683: 678: 670: 659: 654: 160: 156: 152: 148: 144: 140: 136: 132: 128: 5185: 4943: 2250: 455:

and mediate signaling from proteins of subgroups 1 and 2.

5979: 5965: 5356: 5044: 4995: 4688:

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

4489:

Dermatologie Venerologie : Grundlagen. Klinik. Atlas

3355: 2737: 2332:

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

2048:

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

1896: 1788:, with the first clinical use in humans in a recombinant 1663: 1571: 1563: 400:(PRR) and recognize molecules that are broadly shared by 4894: 4687: 4644: 4392: 4351: 4244: 3824: 3675: 3445: 2881: 1956:

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

787:. Furthermore, they are expressed by different types of 762:, which causes food- and waterborne gastroenteritis and 5226: 5142: 3252: 3250: 3078: 2155: 682:

retroviral expression of four transcriptional factors (

382: 39:

region of toll-like receptors, represented here by TLR3

4435: 2533: 2428: 736:

its ligand is the protein profilin and the ligand for

5313: 4193: 3223: 3221: 3219: 2778: 2595: 2472:

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

2251:

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

2047: 755:

effectively, normal mice do not get infected by oral

3304: 3247: 3229:

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

2471: 1758:

yet tightly regulated gateways for gene modulation.

183:, whereas TLR3, TLR7, TLR8, and TLR9 are located in 3541: 3194: 2779:Hanson MA, Hamilton PT, Perlman SJ (October 2016). 2212:"Toll to be paid at the gateway to the vessel wall" 4026: 3924: 3216: 2216:Arteriosclerosis, Thrombosis, and Vascular Biology 777: 5460: 3873: 3448:Journal of Trace Elements in Medicine and Biology 3145: 2598:"The evolution of vertebrate Toll-like receptors" 2383: 1922:, was discovered in 1985 by 1995 Nobel Laureates 6605: 5584:TollML: Toll-like receptors and ligands database 5503:"The Nobel Prize in Physiology or Medicine 2011" 3874:Mishra BB, Gundra UM, Teale JM (December 2008). 3256: 3026: 2828: 2826: 2693: 2654: 5405: 4991: 4989: 4845: 3074: 3072: 2975: 4820: 3582: 3257:Sabroe I, Dower SK, Whyte MK (November 2005). 2502: 2209: 2205: 2203: 1821:. Morphine induced TLR4 activation attenuates 623:) and induction of transcription of genes for 549: 209:The ability of the immune system to recognize 123:, they are recognized by TLRs, which activate 6458: 6236: 5623: 4560: 4486: 4298: 4296: 4294: 4292: 3671: 3669: 3356:Gerondakis S, Grumont RJ, Banerjee A (2007). 3148:"A mouse model of Salmonella typhi infection" 2823: 2733: 2731: 2657:"The host defense of Drosophila melanogaster" 2591: 2589: 2327: 2325: 2323: 5522:"B.C. doctor wins prestigious medical prize" 5087: 5038: 4986: 4937: 4888: 4839: 4829:"Small Molecules Target Toll-Like Receptors" 4771: 4730: 4681: 4638: 4480: 4020: 3767: 3611: 3576: 3535: 3531:https://doi.org/10.1371/journal.pone.0267860 3300: 3298: 3296: 3294: 3227:Unless else specified in boxes then ref is: 3139: 3069: 3020: 2926: 2694:Valanne S, Wang JH, Rämet M (January 2011). 2156:Kemball CC, Alirezaei M, Whitton JL (2010). 1943: 1679:, and TRAM (TRIF-related adaptor molecule). 1558:; proteins such as flagellin from bacterial 1308:Intestinal epithelial cells (IECs) *only in 5136: 4554: 2291: 2200: 2149: 716:As noted above, human cells do not express 480:about choanoflagellate TLR (pmid29848444) . 6465: 6451: 6243: 6229: 5630: 5616: 4302: 4289: 3666: 2728: 2586: 2320: 1841:, and other negative side effects such as 1685: 562:The Toll immunity pathway as found in the 29: 5576:at the U.S. National Library of Medicine 5566:at the U.S. National Library of Medicine 5478: 5374: 5203: 5119: 5070: 5021: 4969: 4920: 4863: 4803: 4713: 4588: 4578: 4537: 4369: 4320: 4170: 4104: 4094: 4052: 3952: 3942: 3901: 3891: 3850: 3801: 3791: 3742: 3693: 3635: 3630:(9). Baltimore, Md. : 1950: 4531–7. 3559: 3506: 3422: 3373: 3332: 3322: 3291: 3274: 3171: 3122: 3104: 3052: 3003: 2993: 2909: 2899: 2850: 2806: 2796: 2755: 2711: 2631: 2621: 2569: 2559: 2503:Chuang TH, Ulevitch RJ (September 2000). 2357: 2268: 2227: 2183: 2132: 2075: 2065: 1906:The prototypic member of the family, the 1903:to the presence of microbial infections. 1721: 1624:. Though most TLRs appear to function as 268:- either in the form of adjuvant used in 5519: 5262: 4027:Hochrein H, Kirschning CJ (March 2013). 3867: 2673:10.1146/annurev.immunol.25.022106.141615 2005: 1611: 1532: 744:. The flagellin from the enteropathogen 708:pluripotent stem cells (iPSC) colonies. 557: 381: 4826: 2696:"The Drosophila Toll signaling pathway" 2035: 1829:and enhances the development of opioid 1777:) is a TLR7 agonist, and its successor 6606: 1871:" to describe a substance produced by 406:pathogen-associated molecular patterns 335:events. Some of these events lead to 6446: 6224: 5611: 5461:Gómez-Gómez L, Boller T (June 2000). 1582: 5002:The European Journal of Neuroscience 1765: 751:As mouse TLR11 is able to recognize 462: 410:damage-associated molecular patterns 217:is, in part, due to the presence of 4561:Toussi DN, Massari P (April 2014). 4143:Journal for Immunotherapy of Cancer 3931:The Journal of Biological Chemistry 3780:The Journal of Biological Chemistry 3305:Sallusto F, Lanzavecchia A (2002). 2982:The Journal of Biological Chemistry 2210:Hansson GK, Edfeldt K (June 2005). 1999:and colleagues. These workers used 513:plant pattern recognition receptors 13: 5112:10.1016/j.neuroscience.2010.02.011 4950:Trends in Pharmacological Sciences 4796:10.1016/j.neuroscience.2009.10.011 2229:10.1161/01.ATV.0000168894.43759.47 2010:The history of Toll-like receptors 458: 14: 6645: 5756:Polymeric immunoglobulin receptor 5557: 1659:) to help fight viral infection. 1620:TLRs are believed to function as 295:to propagate the antigen-induced 241:, cells of the adaptive immunity 5014:10.1111/j.1460-9568.2008.06321.x 1692:Mitogen-activated protein kinase 1566:of viruses; or the unmethylated 467: 5513: 5495: 5454: 5399: 5350: 5307: 5256: 5220: 5179: 4833:Chemical & Engineering News 4605: 4505: 4491:(in German). Berlin: Springer. 4429: 4386: 4345: 4238: 4187: 4129: 4069: 3969: 3918: 3818: 3710: 3523: 3474: 3439: 3398: 3349: 3188: 2969: 2875: 2772: 2687: 2655:Lemaitre B, Hoffmann J (2007). 2648: 2527: 2496: 2465: 2422: 1655:as well as type I interferons ( 1570:islands of bacterial and viral 778:Summary of known mammalian TLRs 276:events that ultimately lead to 16:Class of immune system proteins 5968:(with two glycoprotein chains 4706:10.1016/j.jneuroim.2010.06.007 2412: 2285: 2244: 2092: 2041: 726:and the apicomplexan parasite 377: 283:Upon activation, TLRs recruit 1: 6478:Pattern recognition receptors 6064:Killer-cell IG-like receptors 5480:10.1016/S1097-2765(00)80265-8 5432:10.1126/science.270.5243.1804 5336:10.1126/science.282.5396.2085 5051:Brain, Behavior, and Immunity 4901:Brain, Behavior, and Immunity 4747:10.1016/S0074-7742(09)85016-2 4518:Nature Biomedical Engineering 4303:Kawai T, Akira S (May 2010). 3597:10.1016/s0167-4781(00)00289-x 3197:Paul's fundamental immunology 2852:10.1016/s0092-8674(00)80172-5 1875:that could provoke fever and 1734:. TRIF activates the kinases 748:is also recognized by TLR11. 190:(because they are sensors of 5520:Mitchell B (23 March 2011). 5157:10.1016/0092-8674(88)90516-8 4865:10.1213/ANE.0b013e3182025b15 4371:10.4049/jimmunol.178.10.6252 4096:10.1182/blood-2009-11-251884 3880:Journal of Neuroinflammation 3843:10.1016/j.immuni.2012.09.016 3263:Clinical Infectious Diseases 3106:10.1371/journal.pone.0148987 2901:10.1016/j.molcel.2018.01.029 2757:10.1016/j.celrep.2019.03.101 2386:Journal of Leukocyte Biology 1853: 1698:family. MyD88 then recruits 1607: 1520:conventional dendritic cells 1490:conventional dendritic cells 1487:plasmacytoid dendritic cells 1351:Plasmacytoid dendritic cells 1236:Plasmacytoid dendritic cells 1182:multiple diacyl lipopeptides 1045:B lymphocytes (only in mice) 398:pattern recognition receptor 127:responses. The TLRs include 89:that play a key role in the 7: 6152:Leukocyte IG-like receptors 5535: 5376:10.4049/jimmunol.162.7.3749 3637:10.4049/jimmunol.168.9.4531 3460:10.1016/j.jtemb.2017.07.006 3362:Immunology and Cell Biology 2661:Annual Review of Immunology 1924:Christiane Nüsslein-Volhard 1302:a subset of dendritic cells 1147:a subset of dendritic cells 766:in humans. TLR11 deficient 443:and all have extracellular 297:signal transduction pathway 213:that are broadly shared by 204: 10: 6650: 5643:immunoglobulin superfamily 5598:Toll-Like receptor Pathway 4962:10.1016/j.tips.2009.08.002 4827:Drahl C (22 August 2012). 4694:Journal of Neuroimmunology 4659:10.1016/j.pain.2005.02.003 3487:Journal of Innate Immunity 3424:10.4049/jimmunol.164.2.966 3164:10.1016/j.cell.2012.08.042 3045:10.1016/j.cell.2012.09.034 2935:Nature Reviews. Immunology 1528: 1382:Intestinal epitelial cells 912:multiple lipopeptides and 652: 389: 6580: 6562: 6522: 6484: 6367: 6341: 6263: 6150: 6062: 6041: 5988: 5956: 5921: 5891: 5882: 5853: 5833: 5809: 5800: 5784: 5775: 5768: 5748: 5725: 5692: 5663: 5652: 5063:10.1016/j.bbi.2008.05.004 4913:10.1016/j.bbi.2010.10.018 4530:10.1038/s41551-018-0236-8 2798:10.1186/s12862-016-0805-y 2509:European Cytokine Network 2478:European Cytokine Network 1640:and LPS-Binding Protein ( 1414: 1411: 1408: 1391: 1318:hippocampal interneurons 1229: 1226: 1223: 1221:small synthetic compounds 1220: 1210: 1176: 1141: 1138: 1135: 1122: 1030: 1027: 1020: 1005: 918:Bacterial peptidoglycans 889: 886: 883: 869: 577:immune deficiency pathway 392:Toll-interleukin receptor 69: 57: 49: 44: 28: 23: 5578:Medical Subject Headings 5574:Toll+protein,+Drosophila 5568:Medical Subject Headings 4852:Anesthesia and Analgesia 4155:10.1136/jitc-2023-007198 3695:10.4049/jimmunol.1203245 3658:: CS1 maint: location ( 3561:10.4049/jimmunol.0901888 2995:10.1074/jbc.RA118.006804 2785:BMC Evolutionary Biology 2713:10.4049/jimmunol.1002302 2294:International Immunology 2270:10.1051/medsci/200723167 2067:10.3389/fimmu.2013.00248 1993:adaptive immune response 1366:triacylated lipopeptides 1332:CpG Oligodeoxynucleotide 1066:Bacteria and host cells 881:Bacterial peptidoglycans 774:of human typhoid fever. 711: 539:, which is found in the 249:, and non-immune cells ( 6504:Formyl peptide receptor 5639:Transmembrane receptors 4580:10.3390/vaccines2020323 3998:10.1126/science.1220363 3944:10.1074/jbc.M110.159590 3793:10.1074/jbc.M110.171025 3744:10.1126/science.1109893 2623:10.1073/pnas.0502272102 2561:10.1073/pnas.0400525101 2054:Frontiers in Immunology 1919:Drosophila melanogaster 1916:; Tl) in the fruit fly 1686:MyD88-dependent pathway 1636:, which requires MD-2. 1039:Myeloid dendritic cells 899:Myeloid dendritic cells 677:), which activates the 665: 648: 572:Drosophila melanogaster 418:Interleukin-1 receptors 6614:Developmental genetics 5242:10.1006/geno.1996.0150 4487:Peter Fritsch (2004). 3893:10.1186/1742-2094-5-53 3375:10.1038/sj.icb.7100097 2117:10.4110/in.2020.20.e21 2011: 1944: 1873:Gram-negative bacteria 1843:respiratory depression 1722:TRIF-dependent pathway 1617: 1538: 1017:Gram-negative bacteria 929:Gram-positive bacteria 625:antimicrobial peptides 597:gram-positive bacteria 566: 478:is missing information 387: 347:and digested, and its 6369:Downstream signalling 5603:BioScience Animations 5363:Journal of Immunology 5205:10.1093/dnares/1.1.27 4358:Journal of Immunology 3682:Journal of Immunology 3624:Journal of Immunology 3548:Journal of Immunology 3411:Journal of Immunology 2700:Journal of Immunology 2306:10.1093/intimm/dxh186 2009: 1790:herpes zoster vaccine 1615: 1536: 1517:monocytes/macrophages 1417:monocytes/macrophages 1385:monocytes/macrophages 1348:monocytes/macrophages 1337:Bacteria, DNA viruses 1299:monocytes/macrophages 1232:monocytes/macrophages 1198:monocytes/macrophages 1150:Intestinal epithelium 1049:Intestinal epithelium 1033:monocytes/macrophages 892:monocytes/macrophages 832:Bacterial lipoprotein 561: 451:that are exclusively 431:that are produced by 385: 100:usually expressed on 6474:Innate immune system 5588:University of Munich 5547:Immunologic adjuvant 3317:(Suppl 3): S127-32. 2036:Notes and references 1934:by establishing the 1881:experimental animals 1773:(cardinally used in 1554:, lipopeptides, and 1144:monocyte/macrophages 503:TLRs are present in 331:responses and other 307:proteins, including 299:. These recruited 239:natural killer cells 91:innate immune system 6634:Toll-like receptors 6629:Signal transduction 6537:RIG-I-like receptor 6514:Complement receptor 6486:Membrane-bound_PRRs 5989:Accessory molecules 5854:Accessory molecules 5564:Toll-Like+Receptors 5552:RIG-I-like receptor 5424:1995Sci...270.1804S 5328:1998Sci...282.2085P 5277:1991Natur.351..355G 4627:(3). Archived from 4458:10.1038/nature01182 4450:2002Natur.420..324Y 4259:2000Natur.408..740H 4216:10.1038/nature01889 4208:2003Natur.424..743H 3990:2012Sci...337.1111O 3735:2005Sci...308.1626Y 3269:(Suppl 7): S421-6. 3097:2016PLoSO..1148987H 2988:(26): 10172–10181. 2750:(4): 1050–1061.e3. 2614:2005PNAS..102.9577R 2552:2004PNAS..101.3516T 2431:Nature Cell Biology 2398:10.1189/jlb.0712328 2350:1997Natur.388..394M 2163:Future Microbiology 1895:, and unmethylated 1867:, coined the term " 1675:(also called Mal), 1598:heat shock proteins 1548:lipopolysaccharides 1273:single-stranded RNA 1153:Breast cancer cells 1129:Bacterial flagellin 1062:heat shock proteins 1053:Breast cancer cells 975:double-stranded RNA 414:heat shock proteins 396:TLRs are a type of 79:Toll-like receptors 37:leucine-rich repeat 6509:Scavenger receptor 6494:Toll-like receptor 6053:cytokine receptors 5653:Antibody receptor: 4045:10.4161/onci.23141 3311:Arthritis Research 2176:10.2217/fmb.10.101 2021:Plant homologs of 2012: 2001:positional cloning 1885:lipopolysaccharide 1618: 1583:Endogenous ligands 1562:; double-stranded 1539: 1314:ulcerative colitis 1012:lipopolysaccharide 637:have a total of 9 567: 388: 53:Toll-like receptor 24:Toll-like receptor 6601: 6600: 6532:NOD-like receptor 6440: 6439: 6258:signaling pathway 6252:Signaling pathway 6218: 6217: 6043:Cytokine receptor 6037: 6036: 6033: 6032: 5878: 5877: 5849: 5848: 5764: 5763: 5542:NOD-like receptor 5509:. 3 October 2011. 4634:on 28 April 2021. 4395:Nature Immunology 4309:Nature Immunology 3499:10.1159/000362367 3240:978-0-7817-9543-2 3206:978-1-9751-4251-3 2894:(4): 539–550.e6. 2257:Médecine/Sciences 1962:Jules A. Hoffmann 1786:vaccine adjuvants 1766:Medical relevance 1748:Interferon type I 1657:interferon type I 1556:lipoarabinomannan 1526: 1525: 1471:Toxoplasma gondii 1404:Toxoplasma gondii 1170:Toxoplasma gondii 991:cell compartment 924:lipoteichoic acid 826:multiple triacyl 785:cell compartments 729:Toxoplasma gondii 601:virulence factors 501: 500: 327:that orchestrate 255:endothelial cells 85:) are a class of 76: 75: 6641: 6524:Cytoplasmic_PRRs 6499:Mannose receptor 6467: 6460: 6453: 6444: 6443: 6245: 6238: 6231: 6222: 6221: 5922:Antigen receptor 5889: 5888: 5807: 5806: 5785:Antigen receptor 5782: 5781: 5773: 5772: 5769:Antigen receptor 5727:Alpha (α)/mu (μ) 5661: 5660: 5646:immune receptors 5632: 5625: 5618: 5609: 5608: 5530: 5529: 5517: 5511: 5510: 5499: 5493: 5492: 5482: 5458: 5452: 5451: 5418:(5243): 1804–6. 5403: 5397: 5396: 5378: 5354: 5348: 5347: 5322:(5396): 2085–8. 5311: 5305: 5304: 5285:10.1038/351355b0 5260: 5254: 5253: 5224: 5218: 5217: 5207: 5183: 5177: 5176: 5140: 5134: 5133: 5123: 5091: 5085: 5084: 5074: 5042: 5036: 5035: 5025: 4993: 4984: 4983: 4973: 4941: 4935: 4934: 4924: 4892: 4886: 4885: 4867: 4843: 4837: 4836: 4824: 4818: 4817: 4807: 4775: 4769: 4768: 4734: 4728: 4727: 4717: 4685: 4679: 4678: 4642: 4636: 4635: 4633: 4621:Buffalo Bulletin 4618: 4609: 4603: 4602: 4592: 4582: 4558: 4552: 4551: 4541: 4509: 4503: 4502: 4484: 4478: 4477: 4433: 4427: 4426: 4390: 4384: 4383: 4373: 4349: 4343: 4342: 4324: 4300: 4287: 4286: 4267:10.1038/35047123 4242: 4236: 4235: 4191: 4185: 4184: 4174: 4133: 4127: 4126: 4108: 4098: 4073: 4067: 4066: 4056: 4024: 4018: 4017: 3984:(6098): 1111–5. 3973: 3967: 3966: 3956: 3946: 3922: 3916: 3915: 3905: 3895: 3871: 3865: 3864: 3854: 3822: 3816: 3815: 3805: 3795: 3771: 3765: 3764: 3746: 3729:(5728): 1626–9. 3714: 3708: 3707: 3697: 3673: 3664: 3663: 3657: 3649: 3639: 3615: 3609: 3608: 3580: 3574: 3573: 3563: 3539: 3533: 3527: 3521: 3520: 3510: 3478: 3472: 3471: 3443: 3437: 3436: 3426: 3402: 3396: 3395: 3377: 3353: 3347: 3346: 3336: 3326: 3302: 3289: 3288: 3278: 3254: 3245: 3244: 3225: 3214: 3213: 3192: 3186: 3185: 3175: 3143: 3137: 3136: 3126: 3108: 3076: 3067: 3066: 3056: 3024: 3018: 3017: 3007: 2997: 2973: 2967: 2966: 2930: 2924: 2923: 2913: 2903: 2879: 2873: 2872: 2854: 2830: 2821: 2820: 2810: 2800: 2776: 2770: 2769: 2759: 2735: 2726: 2725: 2715: 2691: 2685: 2684: 2652: 2646: 2645: 2635: 2625: 2593: 2584: 2583: 2573: 2563: 2531: 2525: 2524: 2500: 2494: 2493: 2469: 2463: 2462: 2426: 2420: 2416: 2410: 2409: 2381: 2372: 2371: 2361: 2329: 2318: 2317: 2289: 2283: 2282: 2272: 2248: 2242: 2241: 2231: 2207: 2198: 2197: 2187: 2170:(9): 1329–1347. 2153: 2147: 2146: 2136: 2096: 2090: 2089: 2079: 2069: 2045: 1997:Bruce A. Beutler 1989:Ruslan Medzhitov 1966:fungal infection 1951: 1915: 1861:Richard Pfeiffer 1512:cell compartment 1479:cell compartment 1412:cell compartment 1343:cell compartment 1294:cell compartment 1227:cell compartment 1217:imidazoquinoline 798: 797: 698:; OSKM) induces 496: 493: 487: 471: 463: 449:adaptor proteins 420:form a receptor 412:(DAMPs) such as 285:adaptor proteins 219:immune receptors 118:intestinal tract 33: 21: 20: 6649: 6648: 6644: 6643: 6642: 6640: 6639: 6638: 6619:Insect immunity 6604: 6603: 6602: 6597: 6581:Other/ungrouped 6576: 6558: 6518: 6480: 6471: 6441: 6436: 6363: 6342:Other receptors 6337: 6259: 6249: 6219: 6214: 6146: 6058: 6029: 5984: 5952: 5917: 5874: 5845: 5829: 5796: 5760: 5744: 5721: 5688: 5654: 5648: 5636: 5560: 5538: 5533: 5518: 5514: 5501: 5500: 5496: 5459: 5455: 5404: 5400: 5355: 5351: 5312: 5308: 5271:(6325): 355–6. 5261: 5257: 5225: 5221: 5184: 5180: 5141: 5137: 5092: 5088: 5043: 5039: 4994: 4987: 4942: 4938: 4893: 4889: 4844: 4840: 4825: 4821: 4776: 4772: 4757: 4735: 4731: 4686: 4682: 4643: 4639: 4631: 4616: 4610: 4606: 4559: 4555: 4510: 4506: 4499: 4485: 4481: 4444:(6913): 324–9. 4434: 4430: 4401:(11): 1144–50. 4391: 4387: 4350: 4346: 4322:10.1038/ni.1863 4301: 4290: 4253:(6813): 740–5. 4243: 4239: 4202:(6950): 743–8. 4192: 4188: 4134: 4130: 4074: 4070: 4025: 4021: 3974: 3970: 3923: 3919: 3872: 3868: 3823: 3819: 3772: 3768: 3715: 3711: 3688:(12): 6084–92. 3674: 3667: 3651: 3650: 3616: 3612: 3591:(1–2): 157–61. 3581: 3577: 3554:(9): 5094–103. 3540: 3536: 3528: 3524: 3479: 3475: 3444: 3440: 3403: 3399: 3354: 3350: 3303: 3292: 3255: 3248: 3241: 3226: 3217: 3207: 3193: 3189: 3144: 3140: 3091:(2): e0148987. 3077: 3070: 3025: 3021: 2974: 2970: 2947:10.1038/nri2194 2931: 2927: 2880: 2876: 2831: 2824: 2777: 2773: 2736: 2729: 2692: 2688: 2653: 2649: 2608:(27): 9577–82. 2594: 2587: 2546:(10): 3516–21. 2532: 2528: 2501: 2497: 2470: 2466: 2443:10.1038/ncb1629 2427: 2423: 2417: 2413: 2382: 2375: 2344:(6640): 394–7. 2330: 2321: 2290: 2286: 2249: 2245: 2208: 2201: 2154: 2150: 2097: 2093: 2046: 2042: 2038: 1985:Charles Janeway 1911: 1863:, a student of 1856: 1825:suppression by 1768: 1724: 1688: 1610: 1585: 1531: 1506:Virus, bacteria 1432:urinary bladder 1093:hyaluronic acid 1082:heparan sulfate 996:Dendritic cells 962:Numerous others 857:dendritic cells 807:Ligand location 780: 714: 668: 657: 651: 627:(AMPs) such as 556: 497: 491: 488: 481: 472: 461: 459:Extended family 441:dendritic cells 394: 380: 368:dendritic cells 333:transcriptional 309:protein kinases 264:The binding of 231:dendritic cells 207: 110:dendritic cells 95:single-spanning 40: 17: 12: 11: 5: 6647: 6637: 6636: 6631: 6626: 6621: 6616: 6599: 6598: 6596: 6595: 6584: 6582: 6578: 6577: 6575: 6574: 6568: 6566: 6560: 6559: 6557: 6556: 6555: 6554: 6549: 6544: 6534: 6528: 6526: 6520: 6519: 6517: 6516: 6511: 6506: 6501: 6496: 6490: 6488: 6482: 6481: 6470: 6469: 6462: 6455: 6447: 6438: 6437: 6435: 6434: 6429: 6424: 6419: 6414: 6409: 6404: 6399: 6394: 6389: 6384: 6379: 6373: 6371: 6365: 6364: 6362: 6361: 6356: 6351: 6345: 6343: 6339: 6338: 6336: 6335: 6330: 6325: 6320: 6315: 6310: 6305: 6300: 6295: 6290: 6285: 6280: 6275: 6269: 6267: 6261: 6260: 6248: 6247: 6240: 6233: 6225: 6216: 6215: 6213: 6212: 6207: 6202: 6197: 6192: 6187: 6182: 6177: 6172: 6167: 6162: 6156: 6154: 6148: 6147: 6145: 6144: 6139: 6134: 6129: 6124: 6119: 6114: 6109: 6104: 6099: 6094: 6089: 6084: 6079: 6074: 6068: 6066: 6060: 6059: 6057: 6056: 6047: 6045: 6039: 6038: 6035: 6034: 6031: 6030: 6028: 6027: 6013: 6008: 6003: 5998: 5992: 5990: 5986: 5985: 5983: 5982: 5977: 5962: 5960: 5954: 5953: 5951: 5950: 5945: 5940: 5935: 5925: 5923: 5919: 5918: 5916: 5915: 5914: 5913: 5908: 5897: 5895: 5886: 5880: 5879: 5876: 5875: 5873: 5872: 5857: 5855: 5851: 5850: 5847: 5846: 5844: 5843: 5837: 5835: 5831: 5830: 5828: 5827: 5813: 5811: 5804: 5798: 5797: 5795: 5794: 5788: 5786: 5779: 5770: 5766: 5765: 5762: 5761: 5759: 5758: 5752: 5750: 5746: 5745: 5743: 5742: 5737: 5731: 5729: 5723: 5722: 5720: 5719: 5714: 5709: 5704: 5698: 5696: 5690: 5689: 5687: 5686: 5675: 5669: 5667: 5658: 5650: 5649: 5635: 5634: 5627: 5620: 5612: 5606: 5605: 5600: 5595: 5590: 5581: 5571: 5559: 5558:External links 5556: 5555: 5554: 5549: 5544: 5537: 5534: 5532: 5531: 5512: 5507:Nobel Media AB 5494: 5473:(6): 1003–11. 5467:Molecular Cell 5453: 5398: 5369:(7): 3749–52. 5349: 5306: 5255: 5219: 5178: 5135: 5086: 5057:(8): 1178–89. 5037: 4985: 4956:(11): 581–91. 4936: 4887: 4838: 4819: 4770: 4755: 4729: 4700:(1–2): 26–34. 4680: 4637: 4604: 4553: 4524:(8): 578–588. 4504: 4497: 4479: 4428: 4385: 4364:(10): 6252–8. 4344: 4288: 4237: 4186: 4149:(7): e007198. 4128: 4089:(4): 564–574. 4068: 4033:Oncoimmunology 4019: 3968: 3937:(6): 4517–24. 3917: 3866: 3817: 3786:(5): 3307–14. 3766: 3709: 3665: 3610: 3575: 3534: 3522: 3473: 3438: 3397: 3348: 3290: 3276:10.1086/431992 3246: 3239: 3215: 3205: 3187: 3158:(3): 590–602. 3138: 3068: 3019: 2968: 2941:(11): 862–74. 2925: 2888:Molecular Cell 2874: 2822: 2771: 2727: 2686: 2647: 2585: 2526: 2495: 2464: 2421: 2411: 2373: 2319: 2284: 2243: 2199: 2148: 2105:Immune Network 2091: 2039: 2037: 2034: 1928:Eric Wieschaus 1855: 1852: 1767: 1764: 1723: 1720: 1687: 1684: 1609: 1606: 1594:blood clotting 1584: 1581: 1530: 1527: 1524: 1523: 1522: 1521: 1518: 1513: 1510: 1507: 1504: 1501: 1497: 1496: 1495: 1494: 1491: 1488: 1485: 1480: 1477: 1474: 1467: 1462: 1458: 1457: 1446: 1440: 1439: 1438: 1437: 1429: 1424: 1418: 1413: 1410: 1407: 1400: 1395: 1389: 1388: 1387: 1386: 1383: 1380: 1375: 1372: 1369: 1367: 1364: 1358: 1357: 1356: 1355: 1352: 1349: 1344: 1341: 1338: 1335: 1328: 1322: 1321: 1320: 1319: 1316: 1306: 1303: 1300: 1295: 1292: 1289: 1287: 1284: 1278: 1277: 1274: 1270: 1269: 1263: 1262: 1256: 1255: 1244: 1243: 1242: 1241: 1238: 1233: 1228: 1225: 1222: 1219: 1214: 1208: 1207: 1206: 1205: 1202: 1199: 1194: 1191: 1188: 1183: 1180: 1174: 1173: 1166: 1160: 1159: 1158: 1157: 1154: 1151: 1148: 1145: 1140: 1137: 1134: 1131: 1126: 1120: 1119: 1117: 1109: 1108: 1106: 1100: 1099: 1096: 1089: 1088: 1085: 1078: 1077: 1074: 1068: 1067: 1064: 1057: 1056: 1055: 1054: 1051: 1046: 1043: 1040: 1037: 1034: 1029: 1026: 1019: 1014: 1009: 1003: 1002: 1001: 1000: 997: 992: 989: 984: 981: 972: 966: 965: 963: 959: 958: 955: 944: 943: 938: 932: 931: 926: 920: 919: 916: 909: 908: 907: 906: 901: 896: 893: 888: 885: 882: 879: 873: 867: 866: 865: 864: 859: 853: 842: 839: 833: 830: 824: 818: 817: 814: 811: 808: 805: 802: 779: 776: 713: 710: 667: 664: 653:Main article: 650: 647: 555: 548: 499: 498: 475: 473: 466: 460: 457: 445:Immunoglobulin 390:Main article: 379: 376: 206: 203: 102:sentinel cells 74: 73: 67: 66: 61: 55: 54: 51: 47: 46: 42: 41: 34: 26: 25: 15: 9: 6: 4: 3: 2: 6646: 6635: 6632: 6630: 6627: 6625: 6622: 6620: 6617: 6615: 6612: 6611: 6609: 6593: 6589: 6588:Immunophilins 6586: 6585: 6583: 6579: 6573: 6570: 6569: 6567: 6565: 6564:Secreted_PRRs 6561: 6553: 6550: 6548: 6545: 6543: 6540: 6539: 6538: 6535: 6533: 6530: 6529: 6527: 6525: 6521: 6515: 6512: 6510: 6507: 6505: 6502: 6500: 6497: 6495: 6492: 6491: 6489: 6487: 6483: 6479: 6475: 6468: 6463: 6461: 6456: 6454: 6449: 6448: 6445: 6433: 6430: 6428: 6425: 6423: 6420: 6418: 6415: 6413: 6410: 6408: 6405: 6403: 6400: 6398: 6395: 6393: 6390: 6388: 6385: 6383: 6380: 6378: 6375: 6374: 6372: 6370: 6366: 6360: 6357: 6355: 6352: 6350: 6347: 6346: 6344: 6340: 6334: 6331: 6329: 6326: 6324: 6321: 6319: 6316: 6314: 6311: 6309: 6306: 6304: 6301: 6299: 6296: 6294: 6291: 6289: 6286: 6284: 6281: 6279: 6276: 6274: 6271: 6270: 6268: 6266: 6262: 6257: 6253: 6246: 6241: 6239: 6234: 6232: 6227: 6226: 6223: 6211: 6208: 6206: 6203: 6201: 6198: 6196: 6193: 6191: 6188: 6186: 6183: 6181: 6178: 6176: 6173: 6171: 6168: 6166: 6163: 6161: 6158: 6157: 6155: 6153: 6149: 6143: 6140: 6138: 6135: 6133: 6130: 6128: 6125: 6123: 6120: 6118: 6115: 6113: 6110: 6108: 6105: 6103: 6100: 6098: 6095: 6093: 6090: 6088: 6085: 6083: 6080: 6078: 6075: 6073: 6070: 6069: 6067: 6065: 6061: 6055: 6054: 6049: 6048: 6046: 6044: 6040: 6025: 6021: 6018:(also called 6017: 6014: 6012: 6009: 6007: 6004: 6002: 5999: 5997: 5994: 5993: 5991: 5987: 5981: 5978: 5975: 5971: 5967: 5964: 5963: 5961: 5959: 5955: 5949: 5946: 5944: 5941: 5939: 5936: 5934: 5930: 5927: 5926: 5924: 5920: 5912: 5909: 5907: 5904: 5903: 5902: 5899: 5898: 5896: 5894: 5890: 5887: 5885: 5881: 5870: 5866: 5862: 5859: 5858: 5856: 5852: 5842: 5839: 5838: 5836: 5832: 5826: 5822: 5818: 5815: 5814: 5812: 5808: 5805: 5803: 5799: 5793: 5790: 5789: 5787: 5783: 5780: 5778: 5774: 5771: 5767: 5757: 5754: 5753: 5751: 5747: 5741: 5738: 5736: 5733: 5732: 5730: 5728: 5724: 5718: 5715: 5713: 5710: 5708: 5705: 5703: 5700: 5699: 5697: 5695: 5691: 5684: 5683:C-type lectin 5680: 5676: 5674: 5671: 5670: 5668: 5666: 5662: 5659: 5657: 5651: 5647: 5644: 5640: 5633: 5628: 5626: 5621: 5619: 5614: 5613: 5610: 5604: 5601: 5599: 5596: 5594: 5591: 5589: 5585: 5582: 5579: 5575: 5572: 5569: 5565: 5562: 5561: 5553: 5550: 5548: 5545: 5543: 5540: 5539: 5527: 5523: 5516: 5508: 5504: 5498: 5490: 5486: 5481: 5476: 5472: 5468: 5464: 5457: 5449: 5445: 5441: 5437: 5433: 5429: 5425: 5421: 5417: 5413: 5409: 5402: 5394: 5390: 5386: 5382: 5377: 5372: 5368: 5364: 5360: 5353: 5345: 5341: 5337: 5333: 5329: 5325: 5321: 5317: 5310: 5302: 5298: 5294: 5290: 5286: 5282: 5278: 5274: 5270: 5266: 5259: 5251: 5247: 5243: 5239: 5235: 5231: 5223: 5215: 5211: 5206: 5201: 5197: 5193: 5189: 5182: 5174: 5170: 5166: 5162: 5158: 5154: 5151:(2): 269–79. 5150: 5146: 5139: 5131: 5127: 5122: 5117: 5113: 5109: 5106:(3): 880–93. 5105: 5101: 5097: 5090: 5082: 5078: 5073: 5068: 5064: 5060: 5056: 5052: 5048: 5041: 5033: 5029: 5024: 5019: 5015: 5011: 5007: 5003: 4999: 4992: 4990: 4981: 4977: 4972: 4967: 4963: 4959: 4955: 4951: 4947: 4940: 4932: 4928: 4923: 4918: 4914: 4910: 4907:(2): 349–59. 4906: 4902: 4898: 4891: 4883: 4879: 4875: 4871: 4866: 4861: 4857: 4853: 4849: 4842: 4834: 4830: 4823: 4815: 4811: 4806: 4801: 4797: 4793: 4790:(2): 569–83. 4789: 4785: 4781: 4774: 4766: 4762: 4758: 4756:9780123748935 4752: 4748: 4744: 4740: 4733: 4725: 4721: 4716: 4711: 4707: 4703: 4699: 4695: 4691: 4684: 4676: 4672: 4668: 4664: 4660: 4656: 4653:(1–2): 50–9. 4652: 4648: 4641: 4630: 4626: 4622: 4615: 4608: 4600: 4596: 4591: 4586: 4581: 4576: 4573:(2): 323–53. 4572: 4568: 4564: 4557: 4549: 4545: 4540: 4535: 4531: 4527: 4523: 4519: 4515: 4508: 4500: 4498:3-540-00332-0 4494: 4490: 4483: 4475: 4471: 4467: 4463: 4459: 4455: 4451: 4447: 4443: 4439: 4432: 4424: 4420: 4416: 4412: 4408: 4407:10.1038/ni986 4404: 4400: 4396: 4389: 4381: 4377: 4372: 4367: 4363: 4359: 4355: 4348: 4340: 4336: 4332: 4328: 4323: 4318: 4315:(5): 373–84. 4314: 4310: 4306: 4299: 4297: 4295: 4293: 4284: 4280: 4276: 4272: 4268: 4264: 4260: 4256: 4252: 4248: 4241: 4233: 4229: 4225: 4221: 4217: 4213: 4209: 4205: 4201: 4197: 4190: 4182: 4178: 4173: 4168: 4164: 4160: 4156: 4152: 4148: 4144: 4140: 4132: 4124: 4120: 4116: 4112: 4107: 4102: 4097: 4092: 4088: 4084: 4080: 4072: 4064: 4060: 4055: 4050: 4046: 4042: 4039:(3): e23141. 4038: 4034: 4030: 4023: 4015: 4011: 4007: 4003: 3999: 3995: 3991: 3987: 3983: 3979: 3972: 3964: 3960: 3955: 3950: 3945: 3940: 3936: 3932: 3928: 3921: 3913: 3909: 3904: 3899: 3894: 3889: 3885: 3881: 3877: 3870: 3862: 3858: 3853: 3848: 3844: 3840: 3837:(1): 119–30. 3836: 3832: 3828: 3821: 3813: 3809: 3804: 3799: 3794: 3789: 3785: 3781: 3777: 3770: 3762: 3758: 3754: 3750: 3745: 3740: 3736: 3732: 3728: 3724: 3720: 3713: 3705: 3701: 3696: 3691: 3687: 3683: 3679: 3672: 3670: 3661: 3655: 3647: 3643: 3638: 3633: 3629: 3625: 3621: 3614: 3606: 3602: 3598: 3594: 3590: 3586: 3579: 3571: 3567: 3562: 3557: 3553: 3549: 3545: 3538: 3532: 3526: 3518: 3514: 3509: 3504: 3500: 3496: 3493:(5): 685–94. 3492: 3488: 3484: 3477: 3469: 3465: 3461: 3457: 3453: 3449: 3442: 3434: 3430: 3425: 3420: 3417:(2): 966–72. 3416: 3412: 3408: 3401: 3393: 3389: 3385: 3381: 3376: 3371: 3367: 3363: 3359: 3352: 3344: 3340: 3335: 3330: 3325: 3324:10.1186/ar567 3320: 3316: 3312: 3308: 3301: 3299: 3297: 3295: 3286: 3282: 3277: 3272: 3268: 3264: 3260: 3253: 3251: 3242: 3236: 3232: 3224: 3222: 3220: 3212: 3208: 3202: 3198: 3191: 3183: 3179: 3174: 3169: 3165: 3161: 3157: 3153: 3149: 3142: 3134: 3130: 3125: 3120: 3116: 3112: 3107: 3102: 3098: 3094: 3090: 3086: 3082: 3075: 3073: 3064: 3060: 3055: 3050: 3046: 3042: 3039:(3): 547–58. 3038: 3034: 3030: 3023: 3015: 3011: 3006: 3001: 2996: 2991: 2987: 2983: 2979: 2972: 2964: 2960: 2956: 2952: 2948: 2944: 2940: 2936: 2929: 2921: 2917: 2912: 2907: 2902: 2897: 2893: 2889: 2885: 2878: 2870: 2866: 2862: 2858: 2853: 2848: 2845:(6): 973–83. 2844: 2840: 2836: 2829: 2827: 2818: 2814: 2809: 2804: 2799: 2794: 2790: 2786: 2782: 2775: 2767: 2763: 2758: 2753: 2749: 2745: 2741: 2734: 2732: 2723: 2719: 2714: 2709: 2706:(2): 649–56. 2705: 2701: 2697: 2690: 2682: 2678: 2674: 2670: 2666: 2662: 2658: 2651: 2643: 2639: 2634: 2629: 2624: 2619: 2615: 2611: 2607: 2603: 2599: 2592: 2590: 2581: 2577: 2572: 2567: 2562: 2557: 2553: 2549: 2545: 2541: 2537: 2530: 2522: 2518: 2514: 2510: 2506: 2499: 2491: 2487: 2484:(3): 362–71. 2483: 2479: 2475: 2468: 2460: 2456: 2452: 2448: 2444: 2440: 2437:(9): 1081–8. 2436: 2432: 2425: 2415: 2407: 2403: 2399: 2395: 2391: 2387: 2380: 2378: 2369: 2365: 2360: 2359:10.1038/41131 2355: 2351: 2347: 2343: 2339: 2335: 2328: 2326: 2324: 2315: 2311: 2307: 2303: 2299: 2295: 2288: 2280: 2276: 2271: 2266: 2262: 2258: 2254: 2247: 2239: 2235: 2230: 2225: 2222:(6): 1085–7. 2221: 2217: 2213: 2206: 2204: 2195: 2191: 2186: 2181: 2177: 2173: 2169: 2165: 2164: 2159: 2152: 2144: 2140: 2135: 2130: 2126: 2122: 2118: 2114: 2110: 2106: 2102: 2095: 2087: 2083: 2078: 2073: 2068: 2063: 2059: 2055: 2051: 2044: 2040: 2033: 2030: 2028: 2024: 2019: 2017: 2008: 2004: 2002: 1998: 1994: 1990: 1986: 1981: 1979: 1978:interleukin-1 1974: 1969: 1967: 1963: 1960:was found by 1959: 1955: 1950: 1948: 1941: 1937: 1933: 1932:embryogenesis 1929: 1925: 1921: 1920: 1914: 1909: 1904: 1902: 1901:immune system 1898: 1894: 1890: 1886: 1882: 1878: 1874: 1870: 1866: 1862: 1851: 1847: 1844: 1840: 1836: 1832: 1828: 1824: 1820: 1816: 1812: 1808: 1804: 1800: 1796: 1793: 1791: 1787: 1782: 1780: 1776: 1772: 1763: 1759: 1757: 1751: 1749: 1745: 1741: 1737: 1733: 1729: 1719: 1717: 1713: 1709: 1705: 1701: 1697: 1693: 1683: 1680: 1678: 1674: 1670: 1665: 1660: 1658: 1654: 1650: 1645: 1643: 1639: 1635: 1631: 1628:, TLR2 forms 1627: 1623: 1614: 1605: 1603: 1599: 1595: 1592:(involved in 1591: 1580: 1577: 1573: 1569: 1565: 1561: 1557: 1553: 1549: 1546:cell-surface 1545: 1535: 1519: 1516: 1515: 1514: 1511: 1508: 1505: 1502: 1499: 1498: 1492: 1489: 1486: 1483: 1482: 1481: 1478: 1475: 1473: 1472: 1468: 1466: 1463: 1460: 1459: 1455: 1451: 1447: 1445: 1442: 1441: 1436: 1433: 1430: 1428: 1425: 1422: 1419: 1416: 1415: 1406: 1405: 1401: 1399: 1396: 1394: 1390: 1384: 1381: 1378: 1377: 1376: 1373: 1370: 1368: 1365: 1363: 1360: 1359: 1354:B lymphocytes 1353: 1350: 1347: 1346: 1345: 1342: 1339: 1336: 1333: 1330:unmethylated 1329: 1327: 1324: 1323: 1317: 1315: 1311: 1307: 1304: 1301: 1298: 1297: 1296: 1293: 1290: 1288: 1285: 1283: 1280: 1279: 1275: 1272: 1271: 1268: 1265: 1264: 1261: 1258: 1257: 1253: 1249: 1246: 1245: 1240:B lymphocytes 1239: 1237: 1234: 1231: 1230: 1218: 1215: 1213: 1209: 1204:B lymphocytes 1203: 1200: 1197: 1196: 1195: 1192: 1189: 1187: 1184: 1181: 1179: 1175: 1172: 1171: 1167: 1165: 1162: 1161: 1156:B lymphocytes 1155: 1152: 1149: 1146: 1143: 1142: 1132: 1130: 1127: 1125: 1121: 1118: 1115: 1111: 1110: 1107: 1105: 1102: 1101: 1097: 1094: 1091: 1090: 1086: 1083: 1080: 1079: 1075: 1073: 1070: 1069: 1065: 1063: 1059: 1058: 1052: 1050: 1047: 1044: 1041: 1038: 1035: 1032: 1031: 1024: 1018: 1015: 1013: 1010: 1008: 1004: 999:B lymphocytes 998: 995: 994: 993: 990: 988: 985: 982: 980: 976: 973: 971: 968: 967: 964: 961: 960: 956: 953: 949: 946: 945: 942: 939: 937: 934: 933: 930: 927: 925: 922: 921: 917: 915: 911: 910: 905: 902: 900: 897: 894: 891: 890: 880: 878: 874: 872: 868: 863: 862:B lymphocytes 860: 858: 854: 852: 848: 845: 844: 843: 840: 837: 834: 831: 829: 825: 823: 820: 819: 815: 812: 809: 806: 803: 800: 799: 796: 794: 790: 786: 775: 773: 772:disease model 769: 768:knockout mice 765: 764:typhoid fever 761: 759: 754: 749: 747: 743: 739: 735: 731: 730: 725: 724: 719: 709: 706: 701: 697: 693: 689: 685: 680: 676: 672: 663: 661: 656: 646: 644: 641:family and 6 640: 636: 632: 630: 626: 622: 618: 614: 610: 606: 602: 599:, fungi, and 598: 593: 591: 586: 582: 578: 574: 573: 565: 560: 553: 547: 544: 542: 538: 534: 530: 524: 520: 518: 517:immune system 514: 510: 509:invertebrates 506: 495: 492:December 2021 485: 479: 476:This section 474: 470: 465: 464: 456: 454: 450: 446: 442: 438: 434: 430: 425: 423: 419: 415: 411: 407: 403: 399: 393: 384: 375: 373: 369: 364: 362: 358: 354: 351:presented to 350: 346: 342: 341:proliferation 338: 334: 330: 326: 322: 318: 314: 310: 306: 302: 298: 294: 290: 286: 281: 279: 278:innate immune 275: 271: 267: 262: 260: 256: 252: 248: 244: 240: 236: 232: 228: 224: 220: 216: 212: 202: 200: 195: 193: 192:nucleic acids 189: 186: 185:intracellular 182: 181:cell membrane 178: 174: 170: 166: 162: 158: 154: 150: 146: 142: 138: 134: 130: 126: 122: 119: 115: 111: 107: 103: 99: 96: 92: 88: 84: 80: 72: 68: 65: 62: 60: 56: 52: 48: 43: 38: 32: 27: 22: 19: 6624:LRR proteins 6493: 6332: 6327: 6264: 6255: 6050: 5958:Co-receptors 5911:MHC class II 5525: 5515: 5506: 5497: 5470: 5466: 5456: 5415: 5411: 5401: 5366: 5362: 5352: 5319: 5315: 5309: 5268: 5264: 5258: 5236:(3): 486–8. 5233: 5229: 5222: 5198:(1): 27–35. 5195: 5192:DNA Research 5191: 5181: 5148: 5144: 5138: 5103: 5100:Neuroscience 5099: 5089: 5054: 5050: 5040: 5005: 5001: 4953: 4949: 4939: 4904: 4900: 4890: 4858:(2): 454–9. 4855: 4851: 4841: 4832: 4822: 4787: 4784:Neuroscience 4783: 4773: 4738: 4732: 4697: 4693: 4683: 4650: 4646: 4640: 4629:the original 4624: 4620: 4607: 4570: 4566: 4556: 4521: 4517: 4507: 4488: 4482: 4441: 4437: 4431: 4398: 4394: 4388: 4361: 4357: 4347: 4312: 4308: 4250: 4246: 4240: 4199: 4195: 4189: 4146: 4142: 4131: 4086: 4082: 4071: 4036: 4032: 4022: 3981: 3977: 3971: 3934: 3930: 3920: 3883: 3879: 3869: 3834: 3830: 3820: 3783: 3779: 3769: 3726: 3722: 3712: 3685: 3681: 3654:cite journal 3627: 3623: 3613: 3588: 3584: 3578: 3551: 3547: 3537: 3525: 3490: 3486: 3476: 3451: 3447: 3441: 3414: 3410: 3400: 3368:(6): 471–5. 3365: 3361: 3351: 3314: 3310: 3266: 3262: 3230: 3210: 3196: 3190: 3155: 3151: 3141: 3088: 3084: 3036: 3032: 3022: 2985: 2981: 2978:"Drosophila" 2971: 2938: 2934: 2928: 2891: 2887: 2877: 2842: 2838: 2788: 2784: 2774: 2747: 2744:Cell Reports 2743: 2703: 2699: 2689: 2664: 2660: 2650: 2605: 2601: 2543: 2539: 2529: 2515:(3): 372–8. 2512: 2508: 2498: 2481: 2477: 2467: 2434: 2430: 2424: 2414: 2392:(4): 521–8. 2389: 2385: 2341: 2337: 2297: 2293: 2287: 2263:(1): 67–73. 2260: 2256: 2246: 2219: 2215: 2167: 2161: 2151: 2108: 2104: 2094: 2057: 2053: 2043: 2031: 2026: 2022: 2020: 2016:Shizuo Akira 2013: 1982: 1972: 1970: 1957: 1917: 1907: 1905: 1889:lipopeptides 1857: 1848: 1815:hyperalgesia 1797: 1794: 1783: 1769: 1760: 1752: 1725: 1689: 1681: 1661: 1649:nucleic acid 1646: 1630:heterodimers 1619: 1586: 1552:lipoproteins 1540: 1509:MyD88, TAK-1 1469: 1453: 1449: 1402: 1374:cell surface 1276:RNA viruses 1193:cell surface 1168: 1139:cell surface 1028:cell surface 914:proteolipids 887:cell surface 855:a subset of 841:cell surface 828:lipopeptides 781: 757: 752: 750: 745: 737: 733: 727: 721: 715: 700:pluripotency 669: 658: 642: 638: 634: 633: 608: 594: 570: 568: 551: 545: 525: 521: 502: 489: 477: 429:interleukins 426: 395: 365: 353:CD4+ T cells 345:phagocytosed 339:production, 329:inflammatory 282: 270:vaccinations 263: 208: 196: 82: 78: 77: 18: 6592:Cyclophilin 5906:MHC class I 5802:Co-receptor 5665:Epsilon (ε) 5656:Fc receptor 5008:(1): 20–9. 3454:: 151–160. 2667:: 697–743. 2419:PMC8452412. 2300:(1): 1–14. 2027:Arabidopsis 1945:Das ist ja 1865:Robert Koch 1775:dermatology 1756:pleiotropic 1493:macrophages 1260:bropirimine 1098:host cells 1087:host cells 1076:host cells 1036:neutrophils 952:Beta-glucan 895:neutrophils 877:glycolipids 851:macrophages 816:Cell types 507:as well as 505:vertebrates 433:macrophages 422:superfamily 378:Superfamily 361:interferons 293:immune cell 259:fibroblasts 235:macrophages 125:immune cell 106:macrophages 93:. They are 71:PIRSF037595 45:Identifiers 35:The curved 6608:Categories 5810:stimulate: 4106:2066/89493 3231:Immunology 2791:(1): 228. 2111:(3): e21. 1973:Drosophila 1910:receptor ( 1839:drug abuse 1779:resiquimod 1626:homodimers 1590:fibrinogen 1454:Salmonella 1448:Bacteria ( 1435:epithelium 1305:Mast cells 1267:resiquimod 1254:analogue) 1248:loxoribine 1201:Mast cells 1186:Mycoplasma 1072:fibrinogen 1042:Mast cells 1021:MyD88/MAL/ 941:Host cells 904:Mast cells 810:Adapter(s) 793:cell types 789:leucocytes 758:Salmonella 753:Salmonella 746:Salmonella 734:Toxoplasma 635:Drosophila 629:drosomycin 585:haemolymph 552:Drosophila 533:retrovirus 305:downstream 251:epithelial 229:including 227:leukocytes 59:Membranome 6572:Collectin 5749:Secretory 5694:Gamma (γ) 4163:2051-1426 4115:1528-0020 4014:206540638 3115:1932-6203 2125:1598-2629 1983:In 1997, 1893:flagellin 1869:endotoxin 1854:Discovery 1835:addiction 1831:tolerance 1819:allodynia 1771:Imiquimod 1653:cytokines 1608:Signaling 1544:bacterial 1444:Flagellin 1252:guanosine 1190:MyD88/MAL 1095:fragments 1084:fragments 884:MyD88/MAL 875:multiple 847:monocytes 804:Ligand(s) 791:or other 742:flagellin 564:fruit fly 484:talk page 453:cytosolic 437:monocytes 402:pathogens 372:Flagellin 357:apoptosis 274:molecular 223:membranes 215:pathogens 211:molecules 199:toll gene 98:receptors 6097:KIR2DL5B 6092:KIR2DL5A 5834:inhibit: 5717:Neonatal 5536:See also 5526:The Star 5489:10911994 5448:10548988 5385:10201887 5230:Genomics 5173:19439405 5130:20178837 5081:18599265 5032:18662331 4980:19762094 4931:20974246 4882:12295407 4874:21081778 4814:19833175 4765:19607972 4724:20615556 4667:15836969 4599:26344622 4567:Vaccines 4548:31015631 4474:16163262 4466:12447441 4423:13016860 4415:14556004 4380:17475853 4339:39414949 4331:20404851 4275:11130078 4232:15608748 4224:12872135 4181:37487664 4172:10373699 4123:20424184 4063:23802068 4006:22821982 3963:21131352 3912:19077284 3861:23246311 3831:Immunity 3812:21097503 3761:34165967 3753:15860593 3704:24198280 3646:11970999 3605:11267672 3570:20348427 3517:24861338 3468:28965571 3433:10623846 3392:30443009 3384:17637697 3343:12110131 3285:16237641 3182:23101627 3133:26859749 3085:PLOS ONE 3063:23101625 3014:31088910 2963:11104900 2955:17948019 2920:29452635 2869:10736743 2817:27776480 2766:31018123 2722:21209287 2681:17201680 2642:15976025 2580:14993594 2521:11022120 2490:11022119 2459:12517461 2451:17704767 2406:23345392 2314:15585605 2279:17212934 2238:15923538 2194:20860480 2143:32655969 2086:24032031 1600:(HSPs), 1560:flagella 1465:Profilin 1398:Profilin 1164:Profilin 1133:Bacteria 1112:Various 1060:several 983:viruses 979:poly I:C 813:Location 801:Receptor 554:immunity 550:TLRs in 541:Takifugu 349:antigens 337:cytokine 301:proteins 205:Function 188:vesicles 114:microbes 104:such as 87:proteins 6142:KIR3DS1 6137:KIR3DL3 6132:KIR3DL2 6127:KIR3DL1 6122:KIR2DS5 6117:KIR2DS4 6112:KIR2DS3 6107:KIR2DS2 6102:KIR2DS1 6087:KIR2DL4 6082:KIR2DL3 6077:KIR2DL2 6072:KIR2DL1 6016:ζ-chain 5893:Ligands 5884:T cells 5777:B cells 5712:FcγRIII 5440:8525370 5420:Bibcode 5412:Science 5393:7419784 5344:9851930 5324:Bibcode 5316:Science 5301:1700458 5293:1851964 5273:Bibcode 5250:8838819 5214:7584026 5165:2449285 5121:2854318 5072:2783238 5023:2588470 4971:2783351 4922:3025088 4805:2795035 4715:2942958 4675:7286123 4590:4494261 4539:6192054 4446:Bibcode 4283:4405163 4255:Bibcode 4204:Bibcode 4054:3661153 3986:Bibcode 3978:Science 3954:3039399 3903:2631477 3852:3601573 3803:3030336 3731:Bibcode 3723:Science 3508:4141014 3334:3240143 3173:3500584 3124:4747465 3093:Bibcode 3054:3506423 3005:6664172 2911:5823974 2861:8808632 2808:5078906 2633:1172252 2610:Bibcode 2548:Bibcode 2368:9237759 2346:Bibcode 2185:3045535 2134:7327153 2077:3759294 2060:: 248. 2029:FLS2). 1940:ventral 1827:opioids 1803:opioids 1550:(LPS), 1529:Ligands 1484:Neurons 1450:E. coli 1379:B cells 1371:unknown 1310:Crohn's 948:zymosan 738:E. coli 732:. With 605:spätzle 311:(IKKi, 291:of the 289:cytosol 266:ligands 247:B cells 243:T cells 6427:TOLLIP 6417:TICAM1 6402:MAP3K7 6210:LILRB5 6205:LILRB4 6200:LILRB3 6195:LILRB2 6190:LILRB1 6185:LILRA6 6180:LILRA5 6175:LILRA4 6170:LILRA3 6165:LILRA2 6160:LILRA1 5740:Fcα/μR 5707:FcγRII 5679:FcεRII 5580:(MeSH) 5570:(MeSH) 5487: 5446: 5438: 5391: 5383: 5342: 5299: 5291: 5265:Nature 5248: 5212: 5171: 5163: 5128: 5118: 5079: 5069: 5030: 5020: 4978: 4968: 4929: 4919: 4880: 4872: 4812: 4802: 4763: 4753: 4722: 4712: 4673: 4665: 4597: 4587: 4546: 4536: 4495: 4472: 4464: 4438:Nature 4421: 4413: 4378: 4337: 4329: 4281: 4273: 4247:Nature 4230: 4222: 4196:Nature 4179: 4169: 4161: 4121: 4113: 4061: 4051: 4012: 4004: 3961: 3951: 3910: 3900: 3886:: 53. 3859: 3849: 3810: 3800: 3759: 3751: 3702: 3644: 3603: 3568: 3515: 3505: 3466: 3431: 3390: 3382: 3341: 3331: 3283: 3237: 3203: 3180: 3170: 3131: 3121: 3113: 3061: 3051: 3012: 3002: 2961: 2953: 2918: 2908: 2867: 2859: 2815: 2805: 2764: 2720: 2679: 2640: 2630: 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