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class="vector-toc-list"> <li id="toc-Ligand_binding" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Ligand_binding"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1</span> <span>Ligand binding</span> </div> </a> <ul id="toc-Ligand_binding-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Receptor_recruitment_and_phosphorylation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Receptor_recruitment_and_phosphorylation"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2</span> <span>Receptor recruitment and phosphorylation</span> </div> </a> <ul id="toc-Receptor_recruitment_and_phosphorylation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-SMAD_phosphorylation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#SMAD_phosphorylation"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.3</span> <span>SMAD phosphorylation</span> </div> </a> <ul id="toc-SMAD_phosphorylation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-CoSMAD_binding" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#CoSMAD_binding"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4</span> <span>CoSMAD binding</span> </div> </a> <ul id="toc-CoSMAD_binding-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Transcription" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Transcription"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.5</span> <span>Transcription</span> </div> </a> <ul id="toc-Transcription-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Pathway_regulation" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Pathway_regulation"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Pathway regulation</span> </div> </a> <button aria-controls="toc-Pathway_regulation-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Pathway regulation subsection</span> </button> <ul id="toc-Pathway_regulation-sublist" class="vector-toc-list"> <li id="toc-Ligand_agonists/antagonists" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Ligand_agonists/antagonists"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Ligand agonists/antagonists</span> </div> </a> <ul id="toc-Ligand_agonists/antagonists-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Receptor_regulation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Receptor_regulation"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Receptor regulation</span> </div> </a> <ul id="toc-Receptor_regulation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-R-SMAD_regulation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#R-SMAD_regulation"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>R-SMAD regulation</span> </div> </a> <ul id="toc-R-SMAD_regulation-sublist" class="vector-toc-list"> <li id="toc-Role_of_inhibitory_SMADs" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Role_of_inhibitory_SMADs"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3.1</span> <span>Role of inhibitory SMADs</span> </div> </a> <ul id="toc-Role_of_inhibitory_SMADs-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-R-SMAD_ubiquitination" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#R-SMAD_ubiquitination"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3.2</span> <span>R-SMAD ubiquitination</span> </div> </a> <ul id="toc-R-SMAD_ubiquitination-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </li> <li id="toc-Summary_table" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Summary_table"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Summary table</span> </div> </a> <ul id="toc-Summary_table-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> 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class="vector-pinnable-header-toggle-button vector-pinnable-header-pin-button" data-event-name="pinnable-header.vector-appearance.pin">move to sidebar</button> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-unpin-button" data-event-name="pinnable-header.vector-appearance.unpin">hide</button> </div> </div> </div> </nav> </div> </div> <div id="bodyContent" class="vector-body" aria-labelledby="firstHeading" data-mw-ve-target-container> <div class="vector-body-before-content"> <div class="mw-indicators"> </div> <div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Signaling pathway involving transforming growth factor beta proteins</div> <p>The <b>transforming growth factor beta (TGFβ) signaling pathway</b> is involved in many cellular processes in both the adult organism and the developing <a href="/wiki/Embryo" title="Embryo">embryo</a> including <a href="/wiki/Cell_growth" title="Cell growth">cell growth</a>, <a href="/wiki/Cell_differentiation" class="mw-redirect" title="Cell differentiation">cell differentiation</a>, cell migration, <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a>, cellular homeostasis and other cellular functions. The TGFβ signaling pathways are conserved.<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> In spite of the wide range of cellular processes that the TGFβ signaling pathway regulates, the process is relatively simple. <a href="/wiki/Transforming_growth_factor_beta_family" class="mw-redirect" title="Transforming growth factor beta family">TGFβ superfamily</a> ligands bind to a type II receptor, which recruits and <a href="/wiki/Phosphorylation" title="Phosphorylation">phosphorylates</a> a type I receptor. The type I receptor then phosphorylates receptor-regulated SMADs (<a href="/wiki/R-SMAD" title="R-SMAD">R-SMADs</a>) which can now bind the coSMAD <a href="/wiki/SMAD4" class="mw-redirect" title="SMAD4">SMAD4</a>. R-SMAD/coSMAD complexes accumulate in the nucleus where they act as <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factors</a> and participate in the regulation of target gene expression.<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Mechanism">Mechanism</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=TGF_beta_signaling_pathway&action=edit&section=1" title="Edit section: Mechanism"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Ligand_binding">Ligand binding</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=TGF_beta_signaling_pathway&action=edit&section=2" title="Edit section: Ligand binding"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Main articles: <a href="/wiki/Transforming_growth_factor_beta_family" class="mw-redirect" title="Transforming growth factor beta family">Transforming growth factor beta family</a> and <a href="/wiki/TGF-beta_receptor_family" title="TGF-beta receptor family">TGF-beta receptor family</a></div> <figure class="mw-halign-right" typeof="mw:File"><a href="/wiki/File:TGFbeta_Pathway_1.svg" class="mw-file-description" title="TGF Beta ligand binds to receptor"><img alt="TGF Beta ligand binds to receptor" src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c0/TGFbeta_Pathway_1.svg/150px-TGFbeta_Pathway_1.svg.png" decoding="async" width="150" height="294" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c0/TGFbeta_Pathway_1.svg/225px-TGFbeta_Pathway_1.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c0/TGFbeta_Pathway_1.svg/300px-TGFbeta_Pathway_1.svg.png 2x" data-file-width="153" data-file-height="300" /></a><figcaption>TGF Beta ligand binds to receptor</figcaption></figure> <p>The TGF beta superfamily of ligands includes: <a href="/wiki/Bone_morphogenetic_proteins" class="mw-redirect" title="Bone morphogenetic proteins">bone morphogenetic proteins (BMPs)</a>, <a href="/wiki/Growth_and_differentiation_factor" class="mw-redirect" title="Growth and differentiation factor">growth and differentiation factors (GDFs)</a>, <a href="/wiki/Anti-m%C3%BCllerian_hormone" class="mw-redirect" title="Anti-müllerian hormone">anti-Müllerian hormone (AMH)</a>, <a href="/wiki/Activin_and_inhibin" title="Activin and inhibin">Activin</a>, <a href="/wiki/Nodal_(protein)" class="mw-redirect" title="Nodal (protein)">Nodal</a> and <a href="/wiki/TGF_beta" class="mw-redirect" title="TGF beta">TGFβs</a>.<sup id="cite_ref-superfam_3-0" class="reference"><a href="#cite_note-superfam-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Signaling begins with the binding of a TGF beta superfamily ligand to a TGF beta type II receptor. The type II receptor is a serine/threonine receptor kinase, which <a href="/wiki/Catalyst" class="mw-redirect" title="Catalyst">catalyzes</a> the <a href="/wiki/Phosphorylation" title="Phosphorylation">phosphorylation</a> of the Type I receptor. Each class of ligand binds to a specific type II receptor.<sup id="cite_ref-molbiocell_4-0" class="reference"><a href="#cite_note-molbiocell-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> In mammals there are seven known type I receptors and five type II receptors.<sup id="cite_ref-Nodal_ALK7_5-0" class="reference"><a href="#cite_note-Nodal_ALK7-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>There are three activins: <a href="/wiki/Activin_and_inhibin" title="Activin and inhibin">Activin A</a>, <a href="/wiki/Activin_and_inhibin" title="Activin and inhibin">Activin B</a> and <a href="/wiki/Activin_and_inhibin" title="Activin and inhibin">Activin AB</a>. Activins are involved in embryogenesis and osteogenesis. They also regulate many <a href="/wiki/Hormones" class="mw-redirect" title="Hormones">hormones</a> including <a href="/wiki/Pituitary" class="mw-redirect" title="Pituitary">pituitary</a>, gonadal and <a href="/wiki/Hypothalamic" class="mw-redirect" title="Hypothalamic">hypothalamic</a> hormones as well as <a href="/wiki/Insulin" title="Insulin">insulin</a>. They are also <a href="/wiki/Nerve_cell" class="mw-redirect" title="Nerve cell">nerve cell</a> survival factors. </p><p>The BMPs bind to the <a href="/wiki/BMPR2" title="BMPR2">bone morphogenetic protein receptor type-2</a> (BMPR2). They are involved in a multitude of cellular functions including osteogenesis, <a href="/wiki/Cell_differentiation" class="mw-redirect" title="Cell differentiation">cell differentiation</a>, anterior/posterior axis specification, growth, and homeostasis. </p><p>The TGFβ family includes: <a href="/wiki/TGF%CE%B21" class="mw-redirect" title="TGFβ1">TGFβ1</a>, <a href="/wiki/TGF%CE%B22" class="mw-redirect" title="TGFβ2">TGFβ2</a>, <a href="/wiki/TGF%CE%B23" class="mw-redirect" title="TGFβ3">TGFβ3</a>. Like the BMPs, TGFβs are involved not only in embryogenesis and cell differentiation, but also in apoptosis and other functions. They bind to <a href="/wiki/TGF-beta_receptor_type-2" class="mw-redirect" title="TGF-beta receptor type-2">TGF-beta receptor type-2</a> (TGFβR2). </p><p><a href="/wiki/Nodal_(protein)" class="mw-redirect" title="Nodal (protein)">Nodal</a> binds to activin A receptor, type IIB <a href="/wiki/ACVR2B" title="ACVR2B">ACVR2B</a>. It can then either form a receptor complex with activin A receptor, type IB (<a href="/wiki/ACVR1B" title="ACVR1B">ACVR1B</a>) or with activin A receptor, type IC (<a href="/wiki/ACVR1C" title="ACVR1C">ACVR1C</a>).<sup id="cite_ref-Nodal_ALK7_5-1" class="reference"><a href="#cite_note-Nodal_ALK7-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>When the receptor-ligand binding occurs via local action, this is classified as <a href="/wiki/Paracrine_signalling" class="mw-redirect" title="Paracrine signalling">paracrine signalling</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Receptor_recruitment_and_phosphorylation">Receptor recruitment and phosphorylation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=TGF_beta_signaling_pathway&action=edit&section=3" title="Edit section: Receptor recruitment and phosphorylation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/TGF-beta_receptor_family#Type_I" title="TGF-beta receptor family">TGF-beta receptor family § Type I</a></div> <figure class="mw-halign-left" typeof="mw:File"><a href="/wiki/File:TGF_beta_pathway_2.svg" class="mw-file-description" title="Type II receptor recruits type I receptor and phosphorylates"><img alt="Type II receptor recruits type I receptor and phosphorylates" src="//upload.wikimedia.org/wikipedia/commons/thumb/5/52/TGF_beta_pathway_2.svg/150px-TGF_beta_pathway_2.svg.png" decoding="async" width="150" height="296" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/52/TGF_beta_pathway_2.svg/225px-TGF_beta_pathway_2.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/52/TGF_beta_pathway_2.svg/300px-TGF_beta_pathway_2.svg.png 2x" data-file-width="152" data-file-height="300" /></a><figcaption>Type II receptor recruits type I receptor and phosphorylates</figcaption></figure> <p>The TGF beta ligand binds to a type II receptor dimer, which recruits a type I receptor dimer forming a hetero-tetrameric complex with the ligand.<sup id="cite_ref-dimer_6-0" class="reference"><a href="#cite_note-dimer-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> These receptors are <a href="/wiki/Protein_kinase#Serine/threonine-specific_protein_kinases" title="Protein kinase">serine/threonine kinase receptors</a>. They have a <a href="/wiki/Cysteine" title="Cysteine">cysteine</a> rich <a href="/wiki/Transmembrane_receptor#Extracellular_domains" class="mw-redirect" title="Transmembrane receptor">extracellular domain</a>, a <a href="/wiki/Transmembrane_receptor#Transmembrane_domains" class="mw-redirect" title="Transmembrane receptor">transmembrane domain</a>, and a <a href="/wiki/Membrane_receptor#top" class="mw-redirect" title="Membrane receptor">cytoplasmic</a> serine/threonine rich domain. The <a href="/w/index.php?title=GS_domain&action=edit&redlink=1" class="new" title="GS domain (page does not exist)">GS domain</a> of the type I receptor consists of a series of about thirty <a href="/wiki/Serine" title="Serine">serine</a>-<a href="/wiki/Glycine" title="Glycine">glycine</a> repeats.<sup id="cite_ref-GS_domain_7-0" class="reference"><a href="#cite_note-GS_domain-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> The binding of a TGFβ family ligand causes the rotation of the receptors so that their cytoplasmic kinase domains are arranged in a catalytically favorable orientation. The Type II receptor <a href="/wiki/Phosphorylation" title="Phosphorylation">phosphorylates</a> <a href="/wiki/Serine" title="Serine">serine</a> residues of the Type I receptor, which activates the protein. </p> <div class="mw-heading mw-heading3"><h3 id="SMAD_phosphorylation">SMAD phosphorylation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=TGF_beta_signaling_pathway&action=edit&section=4" title="Edit section: SMAD phosphorylation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-halign-right" typeof="mw:File"><a href="/wiki/File:TGF_beta_pathway_step3.svg" class="mw-file-description" title="Type I receptor phosphorylates R-SMAD"><img alt="Type I receptor phosphorylates R-SMAD" src="//upload.wikimedia.org/wikipedia/commons/thumb/6/69/TGF_beta_pathway_step3.svg/150px-TGF_beta_pathway_step3.svg.png" decoding="async" width="150" height="296" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/69/TGF_beta_pathway_step3.svg/225px-TGF_beta_pathway_step3.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/69/TGF_beta_pathway_step3.svg/300px-TGF_beta_pathway_step3.svg.png 2x" data-file-width="152" data-file-height="300" /></a><figcaption>Type I receptor phosphorylates R-SMAD</figcaption></figure> <p>There are five receptor regulated SMADs: <a href="/wiki/SMAD1" class="mw-redirect" title="SMAD1">SMAD1</a>, <a href="/wiki/SMAD2" class="mw-redirect" title="SMAD2">SMAD2</a>, <a href="/wiki/SMAD3" class="mw-redirect" title="SMAD3">SMAD3</a>, <a href="/wiki/SMAD5" class="mw-redirect" title="SMAD5">SMAD5</a>, and <a href="/wiki/SMAD9" class="mw-redirect" title="SMAD9">SMAD9</a> (sometimes referred to as SMAD8). There are essentially two intracellular pathways involving these <a href="/wiki/R-SMAD" title="R-SMAD">R-SMADs</a>. TGFβs, Activins, Nodals and some GDFs are mediated by SMAD2 and SMAD3, while BMPs, AMH and a few GDFs are mediated by <a href="/wiki/SMAD1" class="mw-redirect" title="SMAD1">SMAD1</a>, <a href="/wiki/SMAD5" class="mw-redirect" title="SMAD5">SMAD5</a> and <a href="/wiki/SMAD9" class="mw-redirect" title="SMAD9">SMAD9</a>. The binding of the R-SMAD to the type I receptor is mediated by a zinc double finger <a href="/wiki/FYVE_domain" title="FYVE domain">FYVE domain</a> containing protein. Two such proteins that mediate the TGFβ pathway include SARA (the <a href="/wiki/SMAD_anchor_for_receptor_activation" class="mw-redirect" title="SMAD anchor for receptor activation">SMAD anchor for receptor activation</a>) and HGS (Hepatocyte growth factor-regulated tyrosine kinase substrate). </p><p>SARA is present in an early <a href="/wiki/Endosome" title="Endosome">endosome</a> which, by <a href="/wiki/Clathrin-mediated_endocytosis" class="mw-redirect" title="Clathrin-mediated endocytosis">clathrin-mediated endocytosis</a>, internalizes the receptor complex.<sup id="cite_ref-SMADendosome_8-0" class="reference"><a href="#cite_note-SMADendosome-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> <a href="/wiki/SMAD_anchor_for_receptor_activation" class="mw-redirect" title="SMAD anchor for receptor activation">SARA</a> recruits an <a href="/wiki/R-SMAD" title="R-SMAD">R-SMAD</a>. SARA permits the binding of the R-SMAD to the L45 region of the Type I receptor.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> SARA orients the R-SMAD such that serine residue on its <a href="/wiki/C-terminus" title="C-terminus">C-terminus</a> faces the catalytic region of the Type I receptor. The Type I receptor <a href="/wiki/Phosphorylation" title="Phosphorylation">phosphorylates</a> the serine residue of the R-SMAD. Phosphorylation induces a conformational change in the <a href="/w/index.php?title=MH2_(domain)&action=edit&redlink=1" class="new" title="MH2 (domain) (page does not exist)">MH2 domain</a> of the R-SMAD and its subsequent dissociation from the receptor complex and SARA.<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="CoSMAD_binding">CoSMAD binding</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=TGF_beta_signaling_pathway&action=edit&section=5" title="Edit section: CoSMAD binding"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-halign-left" typeof="mw:File"><a href="/wiki/File:TGF_beta_Pathway_step4.svg" class="mw-file-description" title="R-SMAD binds coSMAD"><img alt="R-SMAD binds coSMAD" src="//upload.wikimedia.org/wikipedia/commons/thumb/5/50/TGF_beta_Pathway_step4.svg/150px-TGF_beta_Pathway_step4.svg.png" decoding="async" width="150" height="296" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/50/TGF_beta_Pathway_step4.svg/225px-TGF_beta_Pathway_step4.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/50/TGF_beta_Pathway_step4.svg/300px-TGF_beta_Pathway_step4.svg.png 2x" data-file-width="152" data-file-height="300" /></a><figcaption>R-SMAD binds coSMAD</figcaption></figure> <p>The now phosphorylated RSMAD has high affinity for coSMAD (e.g. <a href="/wiki/SMAD4" class="mw-redirect" title="SMAD4">SMAD4</a>) and forms a complex with one. The phosphate group does not act as a docking site for coSMAD, but rather the phosphorylation opens up an amino acid stretch allowing interaction. </p> <div class="mw-heading mw-heading3"><h3 id="Transcription">Transcription</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=TGF_beta_signaling_pathway&action=edit&section=6" title="Edit section: Transcription"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-halign-right" typeof="mw:File"><a href="/wiki/File:Pathwaystep5.svg" class="mw-file-description" title="R-SMAD-coSMAD complex enters nucleus"><img alt="R-SMAD-coSMAD complex enters nucleus" src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b1/Pathwaystep5.svg/150px-Pathwaystep5.svg.png" decoding="async" width="150" height="296" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b1/Pathwaystep5.svg/225px-Pathwaystep5.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b1/Pathwaystep5.svg/300px-Pathwaystep5.svg.png 2x" data-file-width="152" data-file-height="300" /></a><figcaption>R-SMAD-coSMAD complex enters nucleus</figcaption></figure> <p>The phosphorylated RSMAD/coSMAD complex enters the nucleus where it binds transcription promoters/cofactors and causes the transcription of DNA. </p><p>Bone morphogenetic proteins cause the transcription of <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNAs</a> involved in <a href="/wiki/Osteogenesis" class="mw-redirect" title="Osteogenesis">osteogenesis</a>, <a href="/wiki/Neurogenesis" title="Neurogenesis">neurogenesis</a>, and <a href="/wiki/Mesoderm" title="Mesoderm">ventral mesoderm</a> specification. </p><p>TGFβs cause the transcription of mRNAs involved in <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a>, <a href="/wiki/Extracellular_matrix" title="Extracellular matrix">extracellular matrix</a> neogenesis and <a href="/wiki/Immunosuppression" title="Immunosuppression">immunosuppression</a>. They are also involved in <a href="/wiki/G1_phase" title="G1 phase">G1</a> arrest in the <a href="/wiki/Cell_cycle" title="Cell cycle">cell cycle</a>. </p><p>Activin causes the transcription of mRNAs involved in <a href="/wiki/Gonad" title="Gonad">gonadal</a> growth, embryo differentiation and placenta formation. </p><p>Nodal causes the transcription of mRNAs involved in left and right axis specification, <a href="/wiki/Mesoderm" title="Mesoderm">mesoderm</a> and <a href="/wiki/Endoderm" title="Endoderm">endoderm</a> induction. </p> <div class="mw-heading mw-heading2"><h2 id="Pathway_regulation">Pathway regulation</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=TGF_beta_signaling_pathway&action=edit&section=7" title="Edit section: Pathway regulation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The TGF beta signaling pathway is involved in a wide range of cellular process and subsequently is very heavily regulated. There are a variety of mechanisms where the pathway is modulated either positively or negatively, including the agonists for ligands and R-SMADs, the decoy receptors, and the <a href="/wiki/Ubiquitinated" class="mw-redirect" title="Ubiquitinated">ubiquitination</a> of R-SMADs and receptors. </p> <div class="mw-heading mw-heading3"><h3 id="Ligand_agonists/antagonists"><span id="Ligand_agonists.2Fantagonists"></span>Ligand agonists/antagonists</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=TGF_beta_signaling_pathway&action=edit&section=8" title="Edit section: Ligand agonists/antagonists"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Both <a href="/wiki/Chordin" title="Chordin">chordin</a> and <a href="/wiki/Noggin_(protein)" title="Noggin (protein)">noggin</a> are <a href="/wiki/Receptor_antagonist" title="Receptor antagonist">antagonists</a> of BMPs. They bind BMPs preventing the binding of the ligand to the receptor.<sup id="cite_ref-Control_11-0" class="reference"><a href="#cite_note-Control-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> It has been demonstrated that Chordin and Noggin dorsalize <a href="/wiki/Mesoderm" title="Mesoderm">mesoderm</a>. They are both found in the dorsal lip of <i><a href="/wiki/Xenopus" title="Xenopus">Xenopus</a></i> and convert otherwise <a href="/wiki/Epidermis_(skin)" class="mw-redirect" title="Epidermis (skin)">epidermis</a> specified tissue into neural tissue (see <a href="/wiki/Neurulation" title="Neurulation">neurulation</a>). Noggin plays a key role in cartilage and bone patterning. Mice Noggin-/- have excess cartilage and lacked joint formation.<sup id="cite_ref-Control_11-1" class="reference"><a href="#cite_note-Control-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p><p>Members of the DAN family of proteins also antagonize TGF beta family members. They include <a href="/wiki/Cerberus_(protein)" title="Cerberus (protein)">Cerberus</a>, <a href="/wiki/DAN_(protein)" class="mw-redirect" title="DAN (protein)">DAN</a>, and <a href="/wiki/Gremlin_(protein)" title="Gremlin (protein)">Gremlin</a>. These proteins contain nine conserved <a href="/wiki/Cysteine" title="Cysteine">cysteines</a> which can form disulfide bridges. It is believed that DAN antagonizes <a href="/wiki/GDF5" title="GDF5">GDF5</a>, <a href="/wiki/GDF6" title="GDF6">GDF6</a> and <a href="/wiki/GDF7" title="GDF7">GDF7</a>. </p><p><a href="/wiki/Follistatin" title="Follistatin">Follistatin</a> inhibits Activin, which it binds. It directly affects <a href="/wiki/Follicle-stimulating_hormone" title="Follicle-stimulating hormone">follicle-stimulating hormone</a> (FSH) secretion. Follistatin also is implicated in prostate cancers where mutations in its gene may preventing it from acting on activin which has anti-proliferative properties.<sup id="cite_ref-Control_11-2" class="reference"><a href="#cite_note-Control-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Lefty_(protein)" title="Lefty (protein)">Lefty</a> is a regulator of TGFβ and is involved in the axis patterning during embryogenesis. It is also a member of the TGF superfamily of proteins. It is asymmetrically expressed in the left side of murine embryos and subsequently plays a role in left-right specification. Lefty acts by preventing the phosphorylation of R-SMADs. It does so through a constitutively active TGFβ type I receptor and through a process downstream of its activation.<sup id="cite_ref-Lefty_12-0" class="reference"><a href="#cite_note-Lefty-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p><p>Drug-based antagonists have also been identified, such as SB431542,<sup id="cite_ref-SB431542_13-0" class="reference"><a href="#cite_note-SB431542-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> which selectively inhibits ALK4, ALK5, and ALK7. </p> <div class="mw-heading mw-heading3"><h3 id="Receptor_regulation">Receptor regulation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=TGF_beta_signaling_pathway&action=edit&section=9" title="Edit section: Receptor regulation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The <a href="/wiki/TGFBR3" title="TGFBR3">transforming growth factor receptor 3</a> (TGFβR3) is the most abundant of the TGF-β receptors yet,<sup id="cite_ref-TGFBR3_14-0" class="reference"><a href="#cite_note-TGFBR3-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> it has no known signaling domain.<sup id="cite_ref-TGF3_mim_15-0" class="reference"><a href="#cite_note-TGF3_mim-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> It however may serve to enhance the binding of TGFβ ligands to TGFβ type II receptors by binding TGFβ and presenting it to TGFβR2. One of the downstream targets of TGF β signaling, <i>GIPC</i>, binds to its PDZ domain, which prevents its proteosomal degradation, which subsequently increases TGFβ activity. It may also serve as an <a href="/wiki/Inhibin" class="mw-redirect" title="Inhibin">inhibin</a> coreceptor to <a href="/wiki/Activin_type_2_receptors" title="Activin type 2 receptors">ActivinRII</a>.<sup id="cite_ref-Control_11-3" class="reference"><a href="#cite_note-Control-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/BAMBI" title="BAMBI">BMP and activin membrane bound inhibitor</a> (BAMBI), has a similar extracellular domain as type I receptors. It lacks an intracellular serine/threonine protein kinase domain and hence is a pseudoreceptor. It binds to the type I receptor preventing it from being activated. It serves as a negative regulator of TGFβ signaling and may limit TGFβ expression during embryogeneis. It requires BMP signaling for its expression </p><p>FKBP12 binds the GS region of the type I receptor preventing phosphorylation of the receptor by the type II receptors. It is believed that FKBP12 and its homologs help to prevent type I receptor activation in the absence of a ligands, since ligand binding causes its dissociation. </p> <div class="mw-heading mw-heading3"><h3 id="R-SMAD_regulation">R-SMAD regulation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=TGF_beta_signaling_pathway&action=edit&section=10" title="Edit section: R-SMAD regulation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading4"><h4 id="Role_of_inhibitory_SMADs">Role of inhibitory SMADs</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=TGF_beta_signaling_pathway&action=edit&section=11" title="Edit section: Role of inhibitory SMADs"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There are two other SMADs which complete the SMAD family, the <a href="/wiki/I-SMAD" title="I-SMAD">inhibitory SMADs</a> (I-SMADS), <a href="/wiki/SMAD6" class="mw-redirect" title="SMAD6">SMAD6</a> and <a href="/wiki/SMAD7" class="mw-redirect" title="SMAD7">SMAD7</a>. They play a key role in the regulation of TGF beta signaling and are involved in negative feedback. Like other SMADs they have an MH1 and an MH2 domain. SMAD7 competes with other R-SMADs with the Type I receptor and prevents their phosphorylation.<sup id="cite_ref-Control_11-4" class="reference"><a href="#cite_note-Control-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-BMP_ISMAD_16-0" class="reference"><a href="#cite_note-BMP_ISMAD-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> It resides in the nucleus and upon TGFβ receptor activation translocates to the cytoplasm where it binds the type I receptor. <a href="/wiki/SMAD6" class="mw-redirect" title="SMAD6">SMAD6</a> binds SMAD4 preventing the binding of other R-SMADs with the coSMAD. The levels of I-SMAD increase with TGFβ signaling suggesting that they are downstream targets of TGFβ signaling. </p> <div class="mw-heading mw-heading4"><h4 id="R-SMAD_ubiquitination">R-SMAD ubiquitination</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=TGF_beta_signaling_pathway&action=edit&section=12" title="Edit section: R-SMAD ubiquitination"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The E3 ubiquitin-protein <a href="/wiki/Ligase" title="Ligase">ligases</a> <a href="/wiki/SMURF1" title="SMURF1">SMURF1</a> and <a href="/wiki/SMURF2" title="SMURF2">SMURF2</a> regulate the levels of SMADs. They accept <a href="/wiki/Ubiquitin" title="Ubiquitin">ubiquitin</a> from an E2 conjugating enzyme where they transfer ubiquitin to the RSMADs which causes their ubiquitination and subsequent <a href="/wiki/Proteosome" class="mw-redirect" title="Proteosome">proteosomal</a> degradation. SMURF1 binds to <a href="/wiki/SMAD1" class="mw-redirect" title="SMAD1">SMAD1</a> and <a href="/wiki/SMAD5" class="mw-redirect" title="SMAD5">SMAD5</a> while SMURF2 binds <a href="/wiki/SMAD1" class="mw-redirect" title="SMAD1">SMAD1</a>, <a href="/wiki/SMAD2" class="mw-redirect" title="SMAD2">SMAD2</a>, <a href="/wiki/SMAD3" class="mw-redirect" title="SMAD3">SMAD3</a>, <a href="/wiki/SMAD6" class="mw-redirect" title="SMAD6">SMAD6</a> and <a href="/wiki/SMAD7" class="mw-redirect" title="SMAD7">SMAD7</a>. It <sup class="noprint Inline-Template" style="margin-left:0.1em; white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Please_clarify" title="Wikipedia:Please clarify"><span title="The text near this tag may need clarification or removal of jargon. (March 2019)">clarification needed</span></a></i>]</sup> enhances the inhibitory action of SMAD7 while reducing the transcriptional activities of SMAD2. </p> <div class="mw-heading mw-heading2"><h2 id="Summary_table">Summary table</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=TGF_beta_signaling_pathway&action=edit&section=13" title="Edit section: Summary table"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>TGF-β ligands of <i>H.sapiens</i> highlighted in grey, of <i>D.melanogaster</i> ligands in pink, of <i>C.elegans</i> in yellow. </p> <table class="wikitable"> <tbody><tr> <th style="background:Green; color:white">TGF-β superfamily ligand</th> <th>Ligand inhibitors</th> <th style="background:#0054A6; color:white">Type II Receptor</th> <th style="background:#ED1C24; color:white">Type I receptor</th> <th style="background:#CADB2A; color:white"><a href="/wiki/R-SMAD" title="R-SMAD">R-SMADs</a></th> <th style="background:#F7931D; color:white">coSMAD</th> <th><a href="/wiki/I-SMAD" title="I-SMAD">I-SMADs</a> </th></tr> <tr> <th><a href="/wiki/Activin_and_inhibin" title="Activin and inhibin">Activin A</a> </th> <td><a href="/wiki/Follistatin" title="Follistatin">Follistatin</a> </td> <td><a href="/wiki/ACVR2A" title="ACVR2A">ACVR2A</a> </td> <td><a href="/wiki/ACVR1B" title="ACVR1B">ACVR1B</a> (ALK4) </td> <td><a href="/wiki/SMAD2" class="mw-redirect" title="SMAD2">SMAD2</a>, <a href="/wiki/SMAD3" class="mw-redirect" title="SMAD3">SMAD3</a> </td> <td><a href="/wiki/SMAD4" class="mw-redirect" title="SMAD4">SMAD4</a> </td> <td><a href="/wiki/SMAD7" class="mw-redirect" title="SMAD7">SMAD7</a> </td></tr> <tr> <th><a href="/wiki/GDF1" title="GDF1">GDF1</a> </th> <td> </td> <td><a href="/wiki/ACVR2A" title="ACVR2A">ACVR2A</a> </td> <td><a href="/wiki/ACVR1B" title="ACVR1B">ACVR1B</a> (ALK4) </td> <td><a href="/wiki/SMAD2" class="mw-redirect" title="SMAD2">SMAD2</a>, <a href="/wiki/SMAD3" class="mw-redirect" title="SMAD3">SMAD3</a> </td> <td><a href="/wiki/SMAD4" class="mw-redirect" title="SMAD4">SMAD4</a> </td> <td><a href="/wiki/SMAD7" class="mw-redirect" title="SMAD7">SMAD7</a> </td></tr> <tr> <th><a href="/wiki/GDF11" title="GDF11">GDF11</a> </th> <td> </td> <td><a href="/wiki/ACVR2B" title="ACVR2B">ACVR2B</a> </td> <td><a href="/wiki/ACVR1B" title="ACVR1B">ACVR1B</a> (ALK4), <a href="/wiki/TGF%CE%B2RI" class="mw-redirect" title="TGFβRI">TGFβRI</a> (ALK5) </td> <td><a href="/wiki/SMAD2" class="mw-redirect" title="SMAD2">SMAD2</a>, <a href="/wiki/SMAD3" class="mw-redirect" title="SMAD3">SMAD3</a> </td> <td><a href="/wiki/SMAD4" class="mw-redirect" title="SMAD4">SMAD4</a> </td> <td><a href="/wiki/SMAD7" class="mw-redirect" title="SMAD7">SMAD7</a> </td></tr> <tr> <th><a href="/wiki/Bone_morphogenetic_proteins" class="mw-redirect" title="Bone morphogenetic proteins">BMP2-8</a> </th> <td><a href="/wiki/Noggin_(protein)" title="Noggin (protein)">Noggin</a>, <a href="/wiki/Chordin" title="Chordin">Chordin</a>, <a href="/wiki/DAN_(protein)" class="mw-redirect" title="DAN (protein)">DAN</a> </td> <td><a href="/wiki/BMPR2" title="BMPR2">BMPR2</a> </td> <td><a href="/wiki/BMPR1A" title="BMPR1A">BMPR1A</a> (ALK3), <a href="/wiki/BMPR1B" title="BMPR1B">BMPR1B</a> (ALK6) </td> <td><a href="/wiki/SMAD1" class="mw-redirect" title="SMAD1">SMAD1</a> <a href="/wiki/SMAD5" class="mw-redirect" title="SMAD5">SMAD5</a>, <a href="/wiki/SMAD8" class="mw-redirect" title="SMAD8">SMAD8</a> </td> <td><a href="/wiki/SMAD4" class="mw-redirect" title="SMAD4">SMAD4</a> </td> <td><a href="/wiki/SMAD6" class="mw-redirect" title="SMAD6">SMAD6</a>, <a href="/wiki/SMAD7" class="mw-redirect" title="SMAD7">SMAD7</a> </td></tr> <tr> <th><a href="/wiki/Nodal_(protein)" class="mw-redirect" title="Nodal (protein)">Nodal</a> </th> <td><a href="/wiki/Lefty_(protein)" title="Lefty (protein)">Lefty</a> </td> <td><a href="/wiki/ACVR2B" title="ACVR2B">ACVR2B</a> </td> <td><a href="/wiki/ACVR1B" title="ACVR1B">ACVR1B</a> (ALK4), <a href="/wiki/ACVR1C" title="ACVR1C">ACVR1C</a> (ALK7) </td> <td><a href="/wiki/SMAD2" class="mw-redirect" title="SMAD2">SMAD2</a>, <a href="/wiki/SMAD3" class="mw-redirect" title="SMAD3">SMAD3</a> </td> <td><a href="/wiki/SMAD4" class="mw-redirect" title="SMAD4">SMAD4</a> </td> <td><a href="/wiki/SMAD7" class="mw-redirect" title="SMAD7">SMAD7</a> </td></tr> <tr> <th><a href="/wiki/Transforming_growth_factor_beta" title="Transforming growth factor beta">TGFβs</a> </th> <td><a href="/wiki/LTBP1" class="mw-redirect" title="LTBP1">LTBP1</a>, <a href="/wiki/THBS1" class="mw-redirect" title="THBS1">THBS1</a>, <a href="/wiki/Decorin" title="Decorin">Decorin</a> </td> <td><a href="/wiki/TGF%CE%B2RII" class="mw-redirect" title="TGFβRII">TGFβRII</a> </td> <td><a href="/wiki/ACVRL1" title="ACVRL1">ACVRL1</a> (ALK1), <a href="/wiki/TGF%CE%B2RI" class="mw-redirect" title="TGFβRI">TGFβRI</a> (ALK5) </td> <td><a href="/wiki/SMAD2" class="mw-redirect" title="SMAD2">SMAD2</a>, <a href="/wiki/SMAD3" class="mw-redirect" title="SMAD3">SMAD3</a> </td> <td><a href="/wiki/SMAD4" class="mw-redirect" title="SMAD4">SMAD4</a> </td> <td><a href="/wiki/SMAD7" class="mw-redirect" title="SMAD7">SMAD7</a> </td></tr> <tr> <th style="background: pink;"><a href="/wiki/Decapentaplegic" title="Decapentaplegic">Dpp</a> </th> <td> </td> <td><a href="/w/index.php?title=Punt_(gene)&action=edit&redlink=1" class="new" title="Punt (gene) (page does not exist)">Punt</a> </td> <td><a href="/w/index.php?title=Tkv&action=edit&redlink=1" class="new" title="Tkv (page does not exist)">Tkv</a> </td> <td><a href="/wiki/Mothers_against_decapentaplegic" title="Mothers against decapentaplegic">Mad</a> </td> <td><a href="/wiki/Med_(gene)" class="mw-redirect" title="Med (gene)">Medea</a> </td> <td> </td></tr> <tr> <th style="background: pink;"><a href="/wiki/Screw" title="Screw">Screw</a> </th> <td> </td> <td><a href="/w/index.php?title=Punt_(gene)&action=edit&redlink=1" class="new" title="Punt (gene) (page does not exist)">Punt</a> </td> <td><a href="/w/index.php?title=Sax_(gene)&action=edit&redlink=1" class="new" title="Sax (gene) (page does not exist)">Sax</a> </td> <td><a href="/wiki/Mothers_against_decapentaplegic" title="Mothers against decapentaplegic">Mad</a> </td> <td><a href="/wiki/Med_(gene)" class="mw-redirect" title="Med (gene)">Medea</a> </td> <td> </td></tr> <tr> <th style="background: pink;"><a href="/w/index.php?title=Myoglianin&action=edit&redlink=1" class="new" title="Myoglianin (page does not exist)">myoglianin</a> </th> <td> </td> <td><a href="/w/index.php?title=Wit_(gene)&action=edit&redlink=1" class="new" title="Wit (gene) (page does not exist)">Wit</a> </td> <td><a href="/w/index.php?title=Baboon_(gene)&action=edit&redlink=1" class="new" title="Baboon (gene) (page does not exist)">Baboon</a> </td> <td><a href="/w/index.php?title=DSmad2&action=edit&redlink=1" class="new" title="DSmad2 (page does not exist)">dSmad2</a> </td> <td><a href="/wiki/Med_(gene)" class="mw-redirect" title="Med (gene)">Medea</a> </td> <td> </td></tr> <tr> <th style="background: pink;"><a href="/w/index.php?title=DActivin&action=edit&redlink=1" class="new" title="DActivin (page does not exist)">dActivin</a> </th> <td> </td> <td><a href="/w/index.php?title=Wit_(gene)&action=edit&redlink=1" class="new" title="Wit (gene) (page does not exist)">Wit</a>, <a href="/w/index.php?title=Punt_(gene)&action=edit&redlink=1" class="new" title="Punt (gene) (page does not exist)">Punt</a> </td> <td><a href="/w/index.php?title=Baboon_(gene)&action=edit&redlink=1" class="new" title="Baboon (gene) (page does not exist)">Baboon</a> </td> <td><a href="/w/index.php?title=DSmad2&action=edit&redlink=1" class="new" title="DSmad2 (page does not exist)">dSmad2</a> </td> <td><a href="/wiki/Med_(gene)" class="mw-redirect" title="Med (gene)">Medea</a> </td> <td> </td></tr> <tr> <th style="background: pink;"><a href="/wiki/Glass_bottom_boat" class="mw-redirect" title="Glass bottom boat">Gbb</a> </th> <td> </td> <td><a href="/w/index.php?title=Wit_(gene)&action=edit&redlink=1" class="new" title="Wit (gene) (page does not exist)">Wit</a>, <a href="/w/index.php?title=Punt_(gene)&action=edit&redlink=1" class="new" title="Punt (gene) (page does not exist)">Punt</a> </td> <td><a href="/w/index.php?title=Tkv&action=edit&redlink=1" class="new" title="Tkv (page does not exist)">Tkv</a>, <a href="/w/index.php?title=Sax_(gene)&action=edit&redlink=1" class="new" title="Sax (gene) (page does not exist)">Sax</a> </td> <td><a href="/wiki/Mothers_against_decapentaplegic" title="Mothers against decapentaplegic">Mad</a> </td> <td><a href="/wiki/Med_(gene)" class="mw-redirect" title="Med (gene)">Medea</a> </td> <td> </td></tr> <tr> <th style="background: #FFF8DC;"><a href="/wiki/Daf-7" title="Daf-7">Daf-7</a> </th> <td> </td> <td><a href="/wiki/Daf-4" title="Daf-4">Daf-4</a> </td> <td><a href="/wiki/Daf-1" title="Daf-1">Daf-1</a> </td> <td><a href="/wiki/Daf-8" title="Daf-8">Daf-8</a>, <a href="/wiki/Daf-14" title="Daf-14">Daf-14</a> </td> <td><a href="/wiki/Daf-3" title="Daf-3">Daf-3</a> </td> <td> </td></tr> <tr> <th style="background: #FFF8DC;"><a href="/w/index.php?title=Dbl-1&action=edit&redlink=1" class="new" title="Dbl-1 (page does not exist)">Dbl-1</a> </th> <td> </td> <td><a href="/wiki/Daf-4" title="Daf-4">Daf-4</a> </td> <td><a href="/w/index.php?title=Sma-6&action=edit&redlink=1" class="new" title="Sma-6 (page does not exist)">Sma-6</a> </td> <td><a href="/w/index.php?title=Sma-2&action=edit&redlink=1" class="new" title="Sma-2 (page does not exist)">Sma-2</a>, <a href="/w/index.php?title=Sma-3&action=edit&redlink=1" class="new" title="Sma-3 (page does not exist)">Sma-3</a>, <a href="/w/index.php?title=Sma-4&action=edit&redlink=1" class="new" title="Sma-4 (page does not exist)">Sma-4</a> </td> <td><a href="/w/index.php?title=Sma-4&action=edit&redlink=1" class="new" title="Sma-4 (page does not exist)">Sma-4</a> </td> <td> </td></tr></tbody></table> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=TGF_beta_signaling_pathway&action=edit&section=14" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/KEGG_PATHWAY_Database" class="mw-redirect" title="KEGG PATHWAY Database">Kyoto Encyclopedia of Genes and Genomes</a> -<a rel="nofollow" class="external text" href="http://www.genome.ad.jp/dbget-bin/show_pathway?hsa04350+7040">TGF beta signaling pathway</a> map</li> <li><a href="/wiki/Netpath" class="mw-redirect" title="Netpath">Netpath</a> - A curated resource of signal transduction pathways in humans</li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=TGF_beta_signaling_pathway&action=edit&section=15" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width reflist-columns-2"> <ol class="references"> <li id="cite_note-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output 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"Inhibition of transforming growth factor (TGF)-beta1-induced extracellular matrix with a novel inhibitor of the TGF-beta type I receptor kinase activity: SB-431542". <i>Molecular Pharmacology</i>. <b>62</b> (1): 58–64. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1124%2Fmol.62.1.58">10.1124/mol.62.1.58</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/12065755">12065755</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:792324">792324</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Molecular+Pharmacology&rft.atitle=Inhibition+of+transforming+growth+factor+%28TGF%29-beta1-induced+extracellular+matrix+with+a+novel+inhibitor+of+the+TGF-beta+type+I+receptor+kinase+activity%3A+SB-431542&rft.volume=62&rft.issue=1&rft.pages=58-64&rft.date=2002-07&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A792324%23id-name%3DS2CID&rft_id=info%3Apmid%2F12065755&rft_id=info%3Adoi%2F10.1124%2Fmol.62.1.58&rft.aulast=Laping&rft.aufirst=NJ&rft.au=Grygielko%2C+E&rft.au=Mathur%2C+A&rft.au=Butter%2C+S&rft.au=Bomberger%2C+J&rft.au=Tweed%2C+C&rft.au=Martin%2C+W&rft.au=Fornwald%2C+J&rft.au=Lehr%2C+R&rft.au=Harling%2C+J&rft.au=Gaster%2C+L&rft.au=Callahan%2C+JF&rft.au=Olson%2C+BA&rfr_id=info%3Asid%2Fen.wikipedia.org%3ATGF+beta+signaling+pathway" class="Z3988"></span></span> </li> <li id="cite_note-TGFBR3-14"><span class="mw-cite-backlink"><b><a href="#cite_ref-TGFBR3_14-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBlobeLiuFangHow2001" class="citation journal cs1">Blobe GC, Liu X, Fang SJ, How T, Lodish HF (October 2001). <a rel="nofollow" class="external text" href="https://doi.org/10.1074%2Fjbc.M106831200">"A novel mechanism for regulating transforming growth factor beta (TGF-beta) signaling. Functional modulation of type III TGF-beta receptor expression through interaction with the PDZ domain protein, GIPC"</a>. <i>The Journal of Biological Chemistry</i>. <b>276</b> (43): 39608–17. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1074%2Fjbc.M106831200">10.1074/jbc.M106831200</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11546783">11546783</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Journal+of+Biological+Chemistry&rft.atitle=A+novel+mechanism+for+regulating+transforming+growth+factor+beta+%28TGF-beta%29+signaling.+Functional+modulation+of+type+III+TGF-beta+receptor+expression+through+interaction+with+the+PDZ+domain+protein%2C+GIPC&rft.volume=276&rft.issue=43&rft.pages=39608-17&rft.date=2001-10&rft_id=info%3Adoi%2F10.1074%2Fjbc.M106831200&rft_id=info%3Apmid%2F11546783&rft.aulast=Blobe&rft.aufirst=GC&rft.au=Liu%2C+X&rft.au=Fang%2C+SJ&rft.au=How%2C+T&rft.au=Lodish%2C+HF&rft_id=https%3A%2F%2Fdoi.org%2F10.1074%252Fjbc.M106831200&rfr_id=info%3Asid%2Fen.wikipedia.org%3ATGF+beta+signaling+pathway" class="Z3988"></span></span> </li> <li id="cite_note-TGF3_mim-15"><span class="mw-cite-backlink"><b><a href="#cite_ref-TGF3_mim_15-0">^</a></b></span> <span class="reference-text"><a href="/wiki/Online_Mendelian_Inheritance_in_Man" title="Online Mendelian Inheritance in Man">Online Mendelian Inheritance in Man</a> (OMIM): <a rel="nofollow" class="external text" href="https://omim.org/entry/600742">TRANSFORMING GROWTH FACTOR-BETA RECEPTOR, TYPE III; TGFBR3 - 600742</a></span> </li> <li id="cite_note-BMP_ISMAD-16"><span class="mw-cite-backlink"><b><a href="#cite_ref-BMP_ISMAD_16-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFItohAsaoSugamuraHeldin2001" class="citation journal cs1">Itoh F, Asao H, Sugamura K, Heldin CH, ten Dijke P, Itoh S (August 2001). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC149146">"Promoting bone morphogenetic protein signaling through negative regulation of inhibitory Smads"</a>. <i>The EMBO Journal</i>. <b>20</b> (15): 4132–42. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Femboj%2F20.15.4132">10.1093/emboj/20.15.4132</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC149146">149146</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11483516">11483516</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+EMBO+Journal&rft.atitle=Promoting+bone+morphogenetic+protein+signaling+through+negative+regulation+of+inhibitory+Smads&rft.volume=20&rft.issue=15&rft.pages=4132-42&rft.date=2001-08&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC149146%23id-name%3DPMC&rft_id=info%3Apmid%2F11483516&rft_id=info%3Adoi%2F10.1093%2Femboj%2F20.15.4132&rft.aulast=Itoh&rft.aufirst=F&rft.au=Asao%2C+H&rft.au=Sugamura%2C+K&rft.au=Heldin%2C+CH&rft.au=ten+Dijke%2C+P&rft.au=Itoh%2C+S&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC149146&rfr_id=info%3Asid%2Fen.wikipedia.org%3ATGF+beta+signaling+pathway" class="Z3988"></span></span> </li> </ol></div> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist 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href="/wiki/G_protein-coupled_receptor" title="G protein-coupled receptor">GPCR</a></li> <li class="mw-empty-elt"></li> <li><a href="/wiki/Wnt_signaling_pathway" title="Wnt signaling pathway">Wnt</a></li> <li><a href="/wiki/Receptor_tyrosine_kinase" title="Receptor tyrosine kinase">RTK</a> <ul><li><a class="mw-selflink selflink">TGF beta</a></li> <li><a href="/wiki/MAPK/ERK_pathway" title="MAPK/ERK pathway">MAPK/ERK</a></li></ul></li> <li><a href="/wiki/Notch_signaling_pathway" title="Notch signaling pathway">Notch</a></li> <li><a href="/wiki/JAK-STAT_signaling_pathway" title="JAK-STAT signaling pathway">JAK-STAT</a></li> <li><a href="/wiki/Akt/PKB_signaling_pathway" title="Akt/PKB signaling pathway">Akt/PKB</a></li> <li><a href="/wiki/Apoptosis" title="Apoptosis">Fas apoptosis</a></li> <li><a href="/wiki/Hippo_signaling_pathway" title="Hippo signaling pathway">Hippo</a></li> <li><a href="/wiki/PI3K/AKT/mTOR_pathway" title="PI3K/AKT/mTOR pathway">PI3K/AKT/mTOR pathway</a></li> <li><a href="/wiki/Integrin#Signal_transduction" title="Integrin">Integrin receptors</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Agents</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Ligand_(biochemistry)" title="Ligand (biochemistry)">Receptor ligands</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Hormone" title="Hormone">Hormones</a></li> <li><a href="/wiki/Neurotransmitter" title="Neurotransmitter">Neurotransmitters</a>/<a href="/wiki/Neuropeptide" title="Neuropeptide">Neuropeptides</a>/<a href="/wiki/Neurohormone" title="Neurohormone">Neurohormones</a></li> <li><a href="/wiki/Cytokine" title="Cytokine">Cytokines</a></li> <li><a href="/wiki/Growth_factor" title="Growth factor">Growth factors</a></li> <li><a href="/wiki/Cell_signaling" title="Cell signaling">Signaling molecules</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">Receptors</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cell_surface_receptor" title="Cell surface receptor">Cell surface</a></li> <li><a href="/wiki/Intracellular_receptor" title="Intracellular receptor">Intracellular</a></li> <li><a href="/wiki/Co-receptor" title="Co-receptor">Co-receptor</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Second_messenger_system" title="Second messenger system">Second messenger</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/CAMP-dependent_pathway" title="CAMP-dependent pathway">cAMP-dependent pathway</a></li> <li><a href="/wiki/Calcium_signaling" title="Calcium signaling">Ca<sup>2+</sup> signaling</a></li> <li><a href="/wiki/Lipid_signaling" title="Lipid signaling">Lipid signaling</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Assistants:</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Signal_transducing_adaptor_protein" title="Signal transducing adaptor protein">Signal transducing adaptor protein</a></li> <li><a href="/wiki/Scaffold_protein" title="Scaffold protein">Scaffold protein</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Transcription_factor" title="Transcription factor">Transcription factors</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/General_transcription_factor" title="General transcription factor">General</a></li> <li><a href="/wiki/Transcription_preinitiation_complex" title="Transcription preinitiation complex">Transcription preinitiation complex</a></li> <li><a href="/wiki/Transcription_factor_II_D" title="Transcription factor II D">TFIID</a></li> <li><a href="/wiki/Transcription_factor_II_H" title="Transcription factor II H">TFIIH</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">By distance</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Juxtacrine_signalling" title="Juxtacrine signalling">Juxtacrine</a></li> <li><a href="/wiki/Autocrine_signalling" class="mw-redirect" title="Autocrine signalling">Autocrine</a> / <a href="/wiki/Paracrine_signalling" class="mw-redirect" title="Paracrine signalling">Paracrine</a></li> <li><a href="/wiki/Endocrine_system" title="Endocrine system">Endocrine</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other concepts</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Intracrine" title="Intracrine">Intracrine action</a></li> <li>Neurocrine signaling <ul><li><a href="/wiki/Synaptic_transmission" class="mw-redirect" title="Synaptic transmission">Synaptic transmission</a></li> <li><a href="/wiki/Chemical_synapse" title="Chemical synapse">Chemical synapse</a></li></ul></li> <li><a href="/wiki/Neuroendocrine_cell" title="Neuroendocrine cell">Neuroendocrine signaling</a></li> <li><a href="/wiki/Exocrine_gland" title="Exocrine gland">Exocrine signalling</a> <ul><li><a href="/wiki/Pheromone" title="Pheromone">Pheromones</a></li></ul></li> <li><a href="/wiki/Mechanotransduction" title="Mechanotransduction">Mechanotransduction</a></li> <li><a href="/wiki/Visual_phototransduction" title="Visual phototransduction">Phototransduction</a></li> <li><a href="/wiki/Ion_channel" title="Ion channel">Ion channel gating</a></li> <li><a href="/wiki/Gap_junction" title="Gap junction">Gap junction</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Cell_signaling:_TGFβ_signaling_pathway" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2" style="background:#e7dcc3"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:TGF_beta_signaling" title="Template:TGF beta signaling"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:TGF_beta_signaling" title="Template talk:TGF beta signaling"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:TGF_beta_signaling" title="Special:EditPage/Template:TGF beta signaling"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Cell_signaling:_TGFβ_signaling_pathway" style="font-size:114%;margin:0 4em"><a href="/wiki/Cell_signaling" title="Cell signaling">Cell signaling</a>: <a href="/wiki/TGF%CE%B2_signaling_pathway" class="mw-redirect" title="TGFβ signaling pathway">TGFβ signaling pathway</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/Transforming_growth_factor_beta_superfamily" title="Transforming growth factor beta superfamily">TGF beta superfamily of ligands</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite">Ligand of <a href="/wiki/Activin_receptor" title="Activin receptor">ACVR</a> or <a href="/wiki/TGF_beta_receptor" title="TGF beta receptor">TGFBR</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Transforming_growth_factor_beta" title="Transforming growth factor beta">TGF beta family</a> <ul><li><a href="/wiki/TGF_beta_1" title="TGF beta 1">TGF-β1</a></li> <li><a href="/wiki/TGF_beta_2" title="TGF beta 2">TGF-β2</a></li> <li><a href="/wiki/TGF_beta_3" class="mw-redirect" title="TGF beta 3">TGF-β3</a></li></ul></li> <li><a href="/wiki/Activin_and_inhibin" title="Activin and inhibin">Activin and inhibin</a> <ul><li><a href="/wiki/INHA" title="INHA">INHA</a></li> <li><a href="/wiki/INHBA" title="INHBA">INHBA</a></li> <li><a href="/wiki/INHBB" title="INHBB">INHBB</a></li> <li><a href="/wiki/INHBC" title="INHBC">INHBC</a></li></ul></li> <li><a href="/wiki/NODAL" class="mw-redirect" title="NODAL">Nodal</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite">Ligand of <a href="/wiki/Bone_morphogenetic_protein_receptor" title="Bone morphogenetic protein receptor">BMPR</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bone_morphogenetic_protein" title="Bone morphogenetic protein">Bone morphogenetic proteins</a> <ul><li><a href="/wiki/Bone_morphogenetic_protein_2" title="Bone morphogenetic protein 2">BMP2</a></li> <li><a href="/wiki/Bone_morphogenetic_protein_3" title="Bone morphogenetic protein 3">BMP3</a></li> <li><a href="/wiki/Bone_morphogenetic_protein_4" title="Bone morphogenetic protein 4">BMP4</a></li> <li><a href="/wiki/Bone_morphogenetic_protein_5" title="Bone morphogenetic protein 5">BMP5</a></li> <li><a href="/wiki/Bone_morphogenetic_protein_6" title="Bone morphogenetic protein 6">BMP6</a></li> <li><a href="/wiki/Bone_morphogenetic_protein_7" title="Bone morphogenetic protein 7">BMP7</a></li> <li><a href="/wiki/Bone_morphogenetic_protein_8A" title="Bone morphogenetic protein 8A">BMP8a</a></li> <li><a href="/wiki/Bone_morphogenetic_protein_8b" class="mw-redirect" title="Bone morphogenetic protein 8b">BMP8b</a></li> <li><a href="/wiki/Bone_morphogenetic_protein_10" title="Bone morphogenetic protein 10">BMP10</a></li> <li><a href="/wiki/Bone_morphogenetic_protein_15" title="Bone morphogenetic protein 15">BMP15</a></li></ul></li> <li><a href="/wiki/Growth_differentiation_factor" title="Growth differentiation factor">Growth differentiation factors</a> <ul><li><a href="/wiki/GDF1" title="GDF1">GDF1</a></li> <li><a href="/wiki/GDF2" title="GDF2">GDF2</a></li> <li><a href="/wiki/GDF3" title="GDF3">GDF3</a></li> <li><a href="/wiki/GDF5" title="GDF5">GDF5</a></li> <li><a href="/wiki/GDF6" title="GDF6">GDF6</a></li> <li><a href="/wiki/GDF7" title="GDF7">GDF7</a></li> <li><a href="/wiki/Myostatin" title="Myostatin">Myostatin/GDF8</a></li> <li><a href="/wiki/Growth_differentiation_factor-9" title="Growth differentiation factor-9">GDF9</a></li> <li><a href="/wiki/GDF10" title="GDF10">GDF10</a></li> <li><a href="/wiki/GDF11" title="GDF11">GDF11</a></li> <li><a href="/wiki/GDF15" title="GDF15">GDF15</a></li></ul></li> <li><a href="/wiki/Anti-M%C3%BCllerian_hormone" title="Anti-Müllerian hormone">Anti-müllerian hormone</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/TGF_beta_receptors" class="mw-redirect" title="TGF beta receptors">TGF beta receptors</a><br />(<a href="/wiki/Activin_receptor" title="Activin receptor">Activin</a>, <a href="/wiki/Bone_morphogenetic_protein_receptors" class="mw-redirect" title="Bone morphogenetic protein receptors">BMP</a>, <a href="/wiki/TGF-beta_receptor_family" title="TGF-beta receptor family">family</a>)</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite">TGFBR1:</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Activin_type_1_receptors" title="Activin type 1 receptors">Activin type 1 receptors</a> <ul><li><a href="/wiki/ACVR1" title="ACVR1">ACVR1</a></li> <li><a href="/wiki/ACVR1B" title="ACVR1B">ACVR1B</a></li> <li><a href="/wiki/ACVR1C" title="ACVR1C">ACVR1C</a></li></ul></li> <li><a href="/wiki/ACVRL1" title="ACVRL1">ACVRL1</a></li> <li><a href="/wiki/Bone_morphogenetic_protein_receptor,_type_1" title="Bone morphogenetic protein receptor, type 1">BMPR1</a> <ul><li><a href="/wiki/BMPR1A" title="BMPR1A">BMPR1A</a></li> <li><a href="/wiki/BMPR1B" title="BMPR1B">BMPR1B</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite">TGFBR2:</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Activin_type_2_receptors" title="Activin type 2 receptors">Activin type 2 receptors</a> <ul><li><a href="/wiki/ACVR2A" title="ACVR2A">ACVR2A</a></li> <li><a href="/wiki/ACVR2B" title="ACVR2B">ACVR2B</a></li></ul></li> <li><a href="/wiki/Anti-M%C3%BCllerian_hormone_receptor" title="Anti-Müllerian hormone receptor">AMHR2</a></li> <li><a href="/wiki/BMPR2" title="BMPR2">BMPR2</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite">TGFBR3:</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Betaglycan" class="mw-redirect" title="Betaglycan">betaglycan</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/Signal_transduction" title="Signal transduction">Transducers</a>/<a href="/wiki/SMAD_(protein)" title="SMAD (protein)">SMAD</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/R-SMAD" title="R-SMAD">R-SMAD</a> (<a href="/wiki/Mothers_against_decapentaplegic_homolog_1" title="Mothers against decapentaplegic homolog 1">SMAD1</a></li> <li><a href="/wiki/Mothers_against_decapentaplegic_homolog_2" title="Mothers against decapentaplegic homolog 2">SMAD2</a></li> <li><a href="/wiki/Mothers_against_decapentaplegic_homolog_3" title="Mothers against decapentaplegic homolog 3">SMAD3</a></li> <li><a href="/wiki/Mothers_against_decapentaplegic_homolog_5" title="Mothers against decapentaplegic homolog 5">SMAD5</a></li> <li><a href="/wiki/Mothers_against_decapentaplegic_homolog_9" title="Mothers against decapentaplegic homolog 9">SMAD9</a>)</li> <li><a href="/wiki/I-SMAD" title="I-SMAD">I-SMAD</a> (<a href="/wiki/Mothers_against_decapentaplegic_homolog_6" title="Mothers against decapentaplegic homolog 6">SMAD6</a></li> <li><a href="/wiki/Mothers_against_decapentaplegic_homolog_7" title="Mothers against decapentaplegic homolog 7">SMAD7</a>)</li> <li><a href="/wiki/Mothers_against_decapentaplegic_homolog_4" title="Mothers against decapentaplegic homolog 4">SMAD4</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a class="mw-selflink-fragment" href="#Ligand_agonists.2Fantagonists">Ligand inhibitors</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cerberus_(protein)" title="Cerberus (protein)">Cerberus</a></li> <li><a href="/wiki/Chordin" title="Chordin">Chordin</a></li> <li><a href="/wiki/Decorin" title="Decorin">Decorin</a></li> <li><a href="/wiki/Follistatin" title="Follistatin">Follistatin</a></li> <li><a href="/wiki/Gremlin_(protein)" title="Gremlin (protein)">Gremlin</a></li> <li><a href="/wiki/Lefty_(protein)" title="Lefty (protein)">Lefty</a></li> <li><a href="/wiki/Latent_TGF-beta_binding_protein" title="Latent TGF-beta binding protein">LTBP1</a></li> <li><a href="/wiki/Noggin_(protein)" title="Noggin (protein)">Noggin</a></li> <li><a href="/wiki/Poly(A)-specific_ribonuclease" title="Poly(A)-specific ribonuclease">PARN</a></li> <li><a href="/wiki/Thrombospondin_1" title="Thrombospondin 1">THBS1</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/Co-receptor" title="Co-receptor">Coreceptors</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/BAMBI" title="BAMBI">BAMBI</a></li> <li><a href="/wiki/Cripto" title="Cripto">Cripto</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite">Other</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Zinc_finger_FYVE_domain-containing_protein_9" title="Zinc finger FYVE domain-containing protein 9">SARA</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="TGFβ_receptor_superfamily_modulators" style="padding:3px"><table class="nowraplinks hlist mw-collapsible mw-collapsed navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:TGF%CE%B2_receptor_superfamily_modulators" title="Template:TGFβ receptor superfamily modulators"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:TGF%CE%B2_receptor_superfamily_modulators" title="Template talk:TGFβ receptor superfamily modulators"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:TGF%CE%B2_receptor_superfamily_modulators" title="Special:EditPage/Template:TGFβ receptor superfamily modulators"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="TGFβ_receptor_superfamily_modulators" style="font-size:114%;margin:0 4em"><a href="/wiki/TGF%CE%B2_receptor_superfamily" class="mw-redirect" title="TGFβ receptor superfamily"><abbr title="Transforming growth factor beta">TGFβ</abbr> receptor superfamily</a> <a href="/wiki/Receptor_modulator" title="Receptor modulator">modulators</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:center;"><a href="/wiki/TGF%CE%B2_superfamily_receptor#Type_I" class="mw-redirect" title="TGFβ superfamily receptor">Type I</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:9em;;text-align:center;"><a href="/wiki/ALK1" class="mw-redirect" title="ALK1"><abbr title="Activin receptor-like kinase 1">ALK1</abbr> (<abbr title="Activin A receptor-like type 1">ACVRL1</abbr>)</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i>Agonists:</i> <a href="/wiki/Activin" class="mw-redirect" title="Activin">Activin</a> (<a href="/wiki/Activin_A" class="mw-redirect" title="Activin A">A</a>, <a href="/wiki/Activin_B" class="mw-redirect" title="Activin B">B</a>, <a href="/wiki/Activin_AB" class="mw-redirect" title="Activin AB">AB</a>)</li> <li><a href="/wiki/Avotermin" class="mw-redirect" title="Avotermin">Avotermin</a></li> <li><a href="/wiki/Bone_morphogenetic_protein" title="Bone morphogenetic protein">BMP</a> (<a href="/wiki/BMP10" class="mw-redirect" title="BMP10">10</a>)</li> <li><a href="/w/index.php?title=Cetermin&action=edit&redlink=1" class="new" title="Cetermin (page does not exist)">Cetermin</a></li> <li><a href="/wiki/Growth_differentiation_factor" title="Growth differentiation factor">GDF</a> (<a href="/wiki/GDF2" title="GDF2">2 (BMP9)</a>)</li> <li><a href="/wiki/Transforming_growth_factor_beta" title="Transforming growth factor beta">TGFβ</a> (<a href="/wiki/TGF_beta_1" title="TGF beta 1">1</a>, <a href="/wiki/TGF_beta_2" title="TGF beta 2">2</a>, <a href="/wiki/TGF_beta_3" class="mw-redirect" title="TGF beta 3">3</a>)</li></ul> <ul><li><i>Antibodies:</i> <a href="/wiki/Ascrinvacumab" title="Ascrinvacumab">Ascrinvacumab</a></li></ul> <ul><li><i>Kinase inhibitors:</i> <a href="/w/index.php?title=K-02288&action=edit&redlink=1" class="new" title="K-02288 (page does not exist)">K-02288</a></li> <li><a href="/w/index.php?title=ML-347&action=edit&redlink=1" class="new" title="ML-347 (page does not exist)">ML-347 (LDN-193719, VU0469381)</a></li></ul> <ul><li><i>Decoy receptors:</i> <a href="/wiki/Dalantercept" class="mw-redirect" title="Dalantercept">Dalantercept</a></li></ul> <ul><li><i>Other inhibitors:</i> <a href="/w/index.php?title=Disitertide&action=edit&redlink=1" class="new" title="Disitertide (page does not exist)">Disitertide</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:9em;;text-align:center;"><a href="/wiki/ALK2" class="mw-redirect" title="ALK2"><abbr title="Activin receptor-like kinase 2">ALK2</abbr> (<abbr title="Activin A receptor type 1A">ACVR1A</abbr>)</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i>Agonists:</i> <a href="/wiki/Activin" class="mw-redirect" title="Activin">Activin</a> (<a href="/wiki/Activin_A" class="mw-redirect" title="Activin A">A</a>, <a href="/wiki/Activin_B" class="mw-redirect" title="Activin B">B</a>, <a href="/wiki/Activin_AB" class="mw-redirect" title="Activin AB">AB</a>)</li> <li><a href="/wiki/Anti-M%C3%BCllerian_hormone" title="Anti-Müllerian hormone">AMH (MIS)</a></li> <li><a href="/wiki/Avotermin" class="mw-redirect" title="Avotermin">Avotermin</a></li> <li><a href="/wiki/Bone_morphogenetic_protein" title="Bone morphogenetic protein">BMP</a> (<a href="/wiki/BMP5" class="mw-redirect" title="BMP5">5</a>, <a href="/wiki/BMP6" class="mw-redirect" title="BMP6">6</a>, <a href="/wiki/BMP7" class="mw-redirect" title="BMP7">7</a>, <a href="/wiki/BMP8A" class="mw-redirect" title="BMP8A">8A</a>, <a href="/wiki/BMP8B" class="mw-redirect" title="BMP8B">8B</a>)</li> <li><a href="/wiki/Eptotermin_alfa" class="mw-redirect" title="Eptotermin alfa">Eptotermin alfa</a></li> <li><a href="/wiki/Transforming_growth_factor_beta" title="Transforming growth factor beta">TGFβ</a> (<a href="/wiki/TGF_beta_1" title="TGF beta 1">1</a>, <a href="/wiki/TGF_beta_2" title="TGF beta 2">2</a>, <a href="/wiki/TGF_beta_3" class="mw-redirect" title="TGF beta 3">3</a>)</li></ul> <ul><li><i>Kinase inhibitors:</i> <a href="/w/index.php?title=DMH-1&action=edit&redlink=1" class="new" title="DMH-1 (page does not exist)">DMH-1</a></li> <li><a href="/w/index.php?title=DMH-2&action=edit&redlink=1" class="new" title="DMH-2 (page does not exist)">DMH-2</a></li> <li><a href="/w/index.php?title=Dorsomorphin&action=edit&redlink=1" class="new" title="Dorsomorphin (page does not exist)">Dorsomorphin (BML-275)</a></li> <li><a href="/w/index.php?title=K-02288&action=edit&redlink=1" class="new" title="K-02288 (page does not exist)">K-02288</a></li> <li><a href="/w/index.php?title=ML-347&action=edit&redlink=1" class="new" title="ML-347 (page does not exist)">ML-347 (LDN-193719, VU0469381)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:9em;;text-align:center;"><a href="/wiki/ALK3" class="mw-redirect" title="ALK3"><abbr title="Activin receptor-like kinase 3">ALK3</abbr> (<abbr title="Bone morphogenetic protein receptor type 1A">BMPR1A</abbr>)</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i>Agonists:</i> <a href="/wiki/Anti-M%C3%BCllerian_hormone" title="Anti-Müllerian hormone">AMH (MIS)</a></li> <li><a href="/wiki/Bone_morphogenetic_protein" title="Bone morphogenetic protein">BMP</a> (<a href="/wiki/Bone_morphogenetic_protein_2" title="Bone morphogenetic protein 2">2</a>, <a href="/wiki/BMP4" class="mw-redirect" title="BMP4">4</a>, <a href="/wiki/BMP5" class="mw-redirect" title="BMP5">5</a>, <a href="/wiki/BMP6" class="mw-redirect" title="BMP6">6</a>, <a href="/wiki/BMP7" class="mw-redirect" title="BMP7">7</a>, <a href="/wiki/BMP8A" class="mw-redirect" title="BMP8A">8A</a>, <a href="/wiki/BMP8B" class="mw-redirect" title="BMP8B">8B</a>)</li> <li><a href="/wiki/Dibotermin_alfa" class="mw-redirect" title="Dibotermin alfa">Dibotermin alfa</a></li> <li><a href="/wiki/Eptotermin_alfa" class="mw-redirect" title="Eptotermin alfa">Eptotermin alfa</a></li></ul> <ul><li><i>Kinase inhibitors:</i> <a href="/w/index.php?title=DMH-2&action=edit&redlink=1" class="new" title="DMH-2 (page does not exist)">DMH-2</a></li> <li><a href="/w/index.php?title=Dorsomorphin&action=edit&redlink=1" class="new" title="Dorsomorphin (page does not exist)">Dorsomorphin (BML-275)</a></li> <li><a href="/w/index.php?title=K-02288&action=edit&redlink=1" class="new" title="K-02288 (page does not exist)">K-02288</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:9em;;text-align:center;"><a href="/wiki/ALK4" class="mw-redirect" title="ALK4"><abbr title="Activin receptor-like kinase 4">ALK4</abbr> (<abbr title="Activin A receptor type 1B">ACVR1B</abbr>)</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i>Agonists:</i> <a href="/wiki/Activin" class="mw-redirect" title="Activin">Activin</a> (<a href="/wiki/Activin_A" class="mw-redirect" title="Activin A">A</a>, <a href="/wiki/Activin_B" class="mw-redirect" title="Activin B">B</a>, <a href="/wiki/Activin_AB" class="mw-redirect" title="Activin AB">AB</a>)</li> <li><a href="/wiki/Growth_differentiation_factor" title="Growth differentiation factor">GDF</a> (<a href="/wiki/GDF1" title="GDF1">1</a>, <a href="/wiki/GDF3" title="GDF3">3</a>, <a href="/wiki/GDF11" title="GDF11">11 (BMP11)</a>)</li> <li><a href="/wiki/Myostatin" title="Myostatin">Myostatin (GDF8)</a></li> <li><a href="/wiki/NODAL" class="mw-redirect" title="NODAL">Nodal</a></li></ul> <ul><li><i>Antagonists:</i> <a href="/wiki/Inhibin" class="mw-redirect" title="Inhibin">Inhibin</a> (<a href="/wiki/Inhibin_A" class="mw-redirect" title="Inhibin A">A</a>, <a href="/wiki/Inhibin_B" class="mw-redirect" title="Inhibin B">B</a>)</li> <li><a href="/wiki/Lefty_(protein)" title="Lefty (protein)">Lefty</a> (<a href="/wiki/LEFTY1" class="mw-redirect" title="LEFTY1">1</a>, <a href="/wiki/LEFTY2" title="LEFTY2">2</a>)</li></ul> <ul><li><i>Kinase inhibitors:</i> <a href="/w/index.php?title=A_83-01&action=edit&redlink=1" class="new" title="A 83-01 (page does not exist)">A 83-01</a></li> <li><a href="/wiki/SB-431542" title="SB-431542">SB-431542</a></li> <li><a href="/w/index.php?title=SB-505124&action=edit&redlink=1" class="new" title="SB-505124 (page does not exist)">SB-505124</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:9em;;text-align:center;"><a href="/wiki/ALK5" class="mw-redirect" title="ALK5"><abbr title="Activin receptor-like kinase 5">ALK5</abbr> (<abbr title="Transforming growth factor beta receptor 1">TGFβR1</abbr>)</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i>Agonists:</i> <a href="/wiki/Avotermin" class="mw-redirect" title="Avotermin">Avotermin</a></li> <li><a href="/wiki/Growth_differentiation_factor" title="Growth differentiation factor">GDF</a> (<a href="/wiki/GDF10" title="GDF10">10 (BMP3B)</a>, <a href="/wiki/GDF11" title="GDF11">11 (BMP11)</a>)</li> <li><a href="/wiki/Transforming_growth_factor_beta" title="Transforming growth factor beta">TGFβ</a> (<a href="/wiki/TGF_beta_1" title="TGF beta 1">1</a>, <a href="/wiki/TGF_beta_2" title="TGF beta 2">2</a>, <a href="/wiki/TGF_beta_3" class="mw-redirect" title="TGF beta 3">3</a>)</li></ul> <ul><li><i>Antibodies:</i> <a href="/wiki/Fresolimumab" title="Fresolimumab">Fresolimumab</a></li> <li><a href="/wiki/Lerdelimumab" title="Lerdelimumab">Lerdelimumab</a></li> <li><a href="/wiki/Metelimumab" title="Metelimumab">Metelimumab</a></li></ul> <ul><li><i>Kinase inhibitors:</i> <a href="/w/index.php?title=A_83-01&action=edit&redlink=1" class="new" title="A 83-01 (page does not exist)">A 83-01</a></li> <li><a href="/w/index.php?title=D-4476&action=edit&redlink=1" class="new" title="D-4476 (page does not exist)">D-4476</a></li> <li><a href="/w/index.php?title=GW-788388&action=edit&redlink=1" class="new" title="GW-788388 (page does not exist)">GW-788388</a></li> <li><a href="/w/index.php?title=LY-364947&action=edit&redlink=1" class="new" title="LY-364947 (page does not exist)">LY-364947</a></li> <li><a href="/wiki/LY-2109761" title="LY-2109761">LY-2109761</a></li> <li><a href="/wiki/Galunisertib" title="Galunisertib">Galunisertib (LY-2157299)</a></li> <li><a href="/w/index.php?title=R-268712&action=edit&redlink=1" class="new" title="R-268712 (page does not exist)">R-268712</a></li> <li><a href="/wiki/RepSox" title="RepSox">RepSox (E-616452, SJN-2511)</a></li> <li><a href="/wiki/SB-431542" title="SB-431542">SB-431542</a></li> <li><a href="/w/index.php?title=SB-505124&action=edit&redlink=1" class="new" title="SB-505124 (page does not exist)">SB-505124</a></li> <li><a href="/w/index.php?title=SB-525334&action=edit&redlink=1" class="new" title="SB-525334 (page does not exist)">SB-525334</a></li> <li><a href="/w/index.php?title=SD-208&action=edit&redlink=1" class="new" title="SD-208 (page does not exist)">SD-208</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:9em;;text-align:center;"><a href="/wiki/ALK6" class="mw-redirect" title="ALK6"><abbr title="Activin receptor-like kinase 6">ALK6</abbr> (<abbr title="Bone morphogenetic protein receptor type 1B">BMPR1B</abbr>)</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i>Agonists:</i> <a href="/wiki/Bone_morphogenetic_protein" title="Bone morphogenetic protein">BMP</a> (<a href="/wiki/Bone_morphogenetic_protein_2" title="Bone morphogenetic protein 2">2</a>, <a href="/wiki/BMP4" class="mw-redirect" title="BMP4">4</a>, <a href="/wiki/BMP5" class="mw-redirect" title="BMP5">5</a>, <a href="/wiki/BMP6" class="mw-redirect" title="BMP6">6</a>, <a href="/wiki/BMP7" class="mw-redirect" title="BMP7">7</a>, <a href="/wiki/BMP8A" class="mw-redirect" title="BMP8A">8A</a>, <a href="/wiki/BMP8B" class="mw-redirect" title="BMP8B">8B</a>, <a href="/wiki/BMP15" class="mw-redirect" title="BMP15">15 (GDF9B)</a>)</li> <li><a href="/wiki/Dibotermin_alfa" class="mw-redirect" title="Dibotermin alfa">Dibotermin alfa</a></li> <li><a href="/wiki/Eptotermin_alfa" class="mw-redirect" title="Eptotermin alfa">Eptotermin alfa</a></li> <li><a href="/wiki/Growth_differentiation_factor" title="Growth differentiation factor">GDF</a> (<a href="/wiki/GDF5" title="GDF5">5 (BMP14)</a>, <a href="/wiki/GDF6" title="GDF6">6 (BMP13)</a>, <a href="/wiki/GDF7" title="GDF7">7 (BMP12)</a>, <a href="/wiki/GDF9" class="mw-redirect" title="GDF9">9</a>, <a href="/wiki/GDF15" title="GDF15">15</a>)</li> <li><a href="/w/index.php?title=Radotermin&action=edit&redlink=1" class="new" title="Radotermin (page does not exist)">Radotermin</a></li></ul> <ul><li><i>Kinase inhibitors:</i> <a href="/w/index.php?title=DMH-2&action=edit&redlink=1" class="new" title="DMH-2 (page does not exist)">DMH-2</a></li> <li><a href="/w/index.php?title=Dorsomorphin&action=edit&redlink=1" class="new" title="Dorsomorphin (page does not exist)">Dorsomorphin (BML-275)</a></li> <li><a href="/w/index.php?title=K-02288&action=edit&redlink=1" class="new" title="K-02288 (page does not exist)">K-02288</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:9em;;text-align:center;"><a href="/wiki/ALK7" class="mw-redirect" title="ALK7"><abbr title="Activin receptor-like kinase 7">ALK7</abbr> (<abbr title="Activin A receptor type 1C">ACVR1C</abbr>)</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i>Agonists:</i> <a href="/wiki/Growth_differentiation_factor" title="Growth differentiation factor">GDF</a> (<a href="/wiki/GDF1" title="GDF1">1</a>, <a href="/wiki/GDF3" title="GDF3">3</a>, <a href="/wiki/GDF11" title="GDF11">11 (BMP11)</a>)</li> <li><a href="/wiki/NODAL" class="mw-redirect" title="NODAL">Nodal</a></li></ul> <ul><li><i>Antagonists:</i> <a href="/wiki/Lefty_(protein)" title="Lefty (protein)">Lefty</a> (<a href="/wiki/LEFTY1" class="mw-redirect" title="LEFTY1">1</a>, <a href="/wiki/LEFTY2" title="LEFTY2">2</a>)</li></ul> <ul><li><i>Kinase inhibitors:</i> <a href="/w/index.php?title=A_83-01&action=edit&redlink=1" class="new" title="A 83-01 (page does not exist)">A 83-01</a></li> <li><a href="/wiki/SB-431542" title="SB-431542">SB-431542</a></li> <li><a href="/w/index.php?title=SB-505124&action=edit&redlink=1" class="new" title="SB-505124 (page does not exist)">SB-505124</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:center;"><a href="/wiki/TGF%CE%B2_superfamily_receptor#Type_II" class="mw-redirect" title="TGFβ superfamily receptor">Type II</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:9em;;text-align:center;"><a href="/wiki/TGF%CE%B2R2" class="mw-redirect" title="TGFβR2"><abbr title="Transforming growth factor beta receptor 2">TGFβR2</abbr></a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i>Agonists:</i> <a href="/wiki/Avotermin" class="mw-redirect" title="Avotermin">Avotermin</a></li> <li><a href="/wiki/Growth_differentiation_factor" title="Growth differentiation factor">GDF</a> (<a href="/wiki/GDF10" title="GDF10">10 (BMP3B)</a>)</li> <li><a href="/wiki/Transforming_growth_factor_beta" title="Transforming growth factor beta">TGFβ</a> (<a href="/wiki/TGF_beta_1" title="TGF beta 1">1</a>, <a href="/wiki/TGF_beta_2" title="TGF beta 2">2</a>, <a href="/wiki/TGF_beta_3" class="mw-redirect" title="TGF beta 3">3</a>)</li></ul> <ul><li><i>Antibodies:</i> <a href="/wiki/Fresolimumab" title="Fresolimumab">Fresolimumab</a></li> <li><a href="/wiki/Lerdelimumab" title="Lerdelimumab">Lerdelimumab</a></li> <li><a href="/wiki/Metelimumab" title="Metelimumab">Metelimumab</a></li></ul> <ul><li><i>Kinase inhibitors:</i> <a href="/w/index.php?title=DMH-2&action=edit&redlink=1" class="new" title="DMH-2 (page does not exist)">DMH-2</a></li> <li><a href="/w/index.php?title=LY-364947&action=edit&redlink=1" class="new" title="LY-364947 (page does not exist)">LY-364947</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:9em;;text-align:center;"><a href="/wiki/BMPR2" title="BMPR2"><abbr title="Bone morphogenetic protein receptor type 2">BMPR2</abbr></a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i>Agonists:</i> <a href="/wiki/Bone_morphogenetic_protein" title="Bone morphogenetic protein">BMP</a> (<a href="/wiki/Bone_morphogenetic_protein_2" title="Bone morphogenetic protein 2">2</a>, <a href="/wiki/BMP4" class="mw-redirect" title="BMP4">4</a>, <a href="/wiki/BMP6" class="mw-redirect" title="BMP6">6</a>, <a href="/wiki/BMP7" class="mw-redirect" title="BMP7">7</a>, <a href="/wiki/BMP10" class="mw-redirect" title="BMP10">10</a>)</li> <li><a href="/wiki/Dibotermin_alfa" class="mw-redirect" title="Dibotermin alfa">Dibotermin alfa</a></li> <li><a href="/wiki/Eptotermin_alfa" class="mw-redirect" title="Eptotermin alfa">Eptotermin alfa</a></li> <li><a href="/wiki/Growth_differentiation_factor" title="Growth differentiation factor">GDF</a> (<a href="/wiki/GDF2" title="GDF2">2 (BMP9)</a>, <a href="/wiki/GDF5" title="GDF5">5 (BMP14)</a>, <a href="/wiki/GDF6" title="GDF6">6 (BMP13)</a>, <a href="/wiki/GDF7" title="GDF7">7 (BMP12)</a>)</li> <li><a href="/w/index.php?title=Radotermin&action=edit&redlink=1" class="new" title="Radotermin (page does not exist)">Radotermin</a></li></ul> <ul><li><i>Antagonists:</i> <a href="/wiki/Inhibin" class="mw-redirect" title="Inhibin">Inhibin</a> (<a href="/wiki/Inhibin_A" class="mw-redirect" title="Inhibin A">A</a>, <a href="/wiki/Inhibin_B" class="mw-redirect" title="Inhibin B">B</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:9em;;text-align:center;"><a href="/wiki/ACVR2A" title="ACVR2A"><abbr title="Activin receptor type 2A">ACVR2A</abbr> (<abbr title="Activin receptor type 2">ACVR2</abbr>)</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i>Agonists:</i> <a href="/wiki/Activin" class="mw-redirect" title="Activin">Activin</a> (<a href="/wiki/Activin_A" class="mw-redirect" title="Activin A">A</a>, <a href="/wiki/Activin_B" class="mw-redirect" title="Activin B">B</a>, <a href="/wiki/Activin_AB" class="mw-redirect" title="Activin AB">AB</a>)</li> <li><a href="/wiki/Bone_morphogenetic_protein" title="Bone morphogenetic protein">BMP</a> (<a href="/wiki/Bone_morphogenetic_protein_2" title="Bone morphogenetic protein 2">2</a>, <a href="/wiki/BMP4" class="mw-redirect" title="BMP4">4</a>, <a href="/wiki/BMP5" class="mw-redirect" title="BMP5">5</a>, <a href="/wiki/BMP6" class="mw-redirect" title="BMP6">6</a>, <a href="/wiki/BMP7" class="mw-redirect" title="BMP7">7</a>, <a href="/wiki/BMP8A" class="mw-redirect" title="BMP8A">8A</a>, <a href="/wiki/BMP8B" class="mw-redirect" title="BMP8B">8B</a>, <a href="/wiki/BMP15" class="mw-redirect" title="BMP15">15 (GDF9B)</a>)</li> <li><a href="/wiki/Dibotermin_alfa" class="mw-redirect" title="Dibotermin alfa">Dibotermin alfa</a></li> <li><a href="/wiki/Eptotermin_alfa" class="mw-redirect" title="Eptotermin alfa">Eptotermin alfa</a></li> <li><a href="/wiki/Growth_differentiation_factor" title="Growth differentiation factor">GDF</a> (<a href="/wiki/GDF1" title="GDF1">1</a>, <a href="/wiki/GDF3" title="GDF3">3</a>, <a href="/wiki/GDF5" title="GDF5">5 (BMP14)</a>, <a href="/wiki/GDF6" title="GDF6">6 (BMP13)</a>, <a href="/wiki/GDF7" title="GDF7">7 (BMP12)</a>, <a href="/wiki/GDF9" class="mw-redirect" title="GDF9">9</a>, <a href="/wiki/GDF11" title="GDF11">11 (BMP11)</a>, <a href="/wiki/GDF15" title="GDF15">15</a>)</li> <li><a href="/wiki/Myostatin" title="Myostatin">Myostatin (GDF8)</a></li> <li><a href="/wiki/NODAL" class="mw-redirect" title="NODAL">Nodal</a></li> <li><a href="/w/index.php?title=Radotermin&action=edit&redlink=1" class="new" title="Radotermin (page does not exist)">Radotermin</a></li></ul> <ul><li><i>Antagonists:</i> <a href="/wiki/Inhibin" class="mw-redirect" title="Inhibin">Inhibin</a> (<a href="/wiki/Inhibin_A" class="mw-redirect" title="Inhibin A">A</a>, <a href="/wiki/Inhibin_B" class="mw-redirect" title="Inhibin B">B</a>)</li> <li><a href="/wiki/Lefty_(protein)" title="Lefty (protein)">Lefty</a> (<a href="/wiki/LEFTY1" class="mw-redirect" title="LEFTY1">1</a>, <a href="/wiki/LEFTY2" title="LEFTY2">2</a>)</li></ul> <ul><li><i>Decoy receptors:</i> <a href="/wiki/Sotatercept" title="Sotatercept">Sotatercept</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:9em;;text-align:center;"><a href="/wiki/ACVR2A" title="ACVR2A"><abbr title="Activin receptor type 2B">ACVR2B</abbr></a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i>Agonists:</i> <a href="/wiki/Activin" class="mw-redirect" title="Activin">Activin</a> (<a href="/wiki/Activin_A" class="mw-redirect" title="Activin A">A</a>, <a href="/wiki/Activin_B" class="mw-redirect" title="Activin B">B</a>, <a href="/wiki/Activin_AB" class="mw-redirect" title="Activin AB">AB</a>)</li> <li><a href="/wiki/Bone_morphogenetic_protein" title="Bone morphogenetic protein">BMP</a> (<a href="/wiki/Bone_morphogenetic_protein_2" title="Bone morphogenetic protein 2">2</a>, <a href="/wiki/BMP4" class="mw-redirect" title="BMP4">4</a>, <a href="/wiki/BMP6" class="mw-redirect" title="BMP6">6</a>, <a href="/wiki/BMP7" class="mw-redirect" title="BMP7">7</a>)</li> <li><a href="/wiki/Dibotermin_alfa" class="mw-redirect" title="Dibotermin alfa">Dibotermin alfa</a></li> <li><a href="/wiki/Eptotermin_alfa" class="mw-redirect" title="Eptotermin alfa">Eptotermin alfa</a></li> <li><a href="/wiki/Growth_differentiation_factor" title="Growth differentiation factor">GDF</a> (<a href="/wiki/GDF1" title="GDF1">1</a>, <a href="/wiki/GDF3" title="GDF3">3</a>, <a href="/wiki/GDF5" title="GDF5">5 (BMP14)</a>, <a href="/wiki/GDF6" title="GDF6">6 (BMP13)</a>, <a href="/wiki/GDF7" title="GDF7">7 (BMP12)</a>)</li> <li><a href="/wiki/Myostatin" title="Myostatin">Myostatin (GDF8)</a></li> <li><a href="/wiki/NODAL" class="mw-redirect" title="NODAL">Nodal</a></li> <li><a href="/wiki/Bone_morphogenetic_protein_3" title="Bone morphogenetic protein 3">Osteogenin (BMP3, BMP3A)</a></li> <li><a href="/w/index.php?title=Radotermin&action=edit&redlink=1" class="new" title="Radotermin (page does not exist)">Radotermin</a></li></ul> <ul><li><i>Antagonists:</i> <a href="/wiki/Inhibin" class="mw-redirect" title="Inhibin">Inhibin</a> (<a href="/wiki/Inhibin_A" class="mw-redirect" title="Inhibin A">A</a>, <a href="/wiki/Inhibin_B" class="mw-redirect" title="Inhibin B">B</a>)</li> <li><a href="/wiki/Lefty_(protein)" title="Lefty (protein)">Lefty</a> (<a href="/wiki/LEFTY1" class="mw-redirect" title="LEFTY1">1</a>, <a href="/wiki/LEFTY2" title="LEFTY2">2</a>)</li></ul> <ul><li><i>Decoy receptors:</i> <a href="/w/index.php?title=Ramatercept&action=edit&redlink=1" class="new" title="Ramatercept (page does not exist)">Ramatercept</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:9em;;text-align:center;"><a href="/wiki/AMHR2" class="mw-redirect" title="AMHR2"><abbr title="Anti-Müllerian hormone receptor type 2">AMHR2</abbr> (<abbr title="Anti-Müllerian hormone receptor">AMHR</abbr>)</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i>Agonists:</i> <a href="/wiki/Anti-M%C3%BCllerian_hormone" title="Anti-Müllerian hormone">AMH (MIS)</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:center;"><a href="/wiki/TGF%CE%B2_superfamily_receptor#Type_III" class="mw-redirect" title="TGFβ superfamily receptor">Type III</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="TGFβR3_(β-glycan)" scope="row" class="navbox-group" style="width:9em;;text-align:center;"><a href="/wiki/TGF%CE%B2R3" class="mw-redirect" title="TGFβR3"><abbr title="Transforming growth factor beta receptor 3">TGFβR3</abbr> (β-glycan)</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i>Ligands:</i> <a href="/wiki/Avotermin" class="mw-redirect" title="Avotermin">Avotermin</a></li> <li><a href="/wiki/Inhibin" class="mw-redirect" title="Inhibin">Inhibin</a> (<a href="/wiki/Inhibin_A" class="mw-redirect" title="Inhibin A">A</a>, <a href="/wiki/Inhibin_B" class="mw-redirect" title="Inhibin B">B</a>)</li> <li><a href="/wiki/Transforming_growth_factor_beta" title="Transforming growth factor beta">TGFβ</a> (<a href="/wiki/TGF_beta_1" title="TGF beta 1">1</a>, <a href="/wiki/TGF_beta_2" title="TGF beta 2">2</a>, <a href="/wiki/TGF_beta_3" class="mw-redirect" title="TGF beta 3">3</a>)</li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:center;">Unsorted</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i>Endogenous antagonists/inhibitors:</i> <a href="/wiki/BAMBI" title="BAMBI">BAMBI</a></li> <li><a href="/wiki/Cerberus_(protein)" title="Cerberus (protein)">Cerberus (CER1)</a></li> <li><a href="/wiki/Chordin" title="Chordin">Chordin</a></li> <li><a href="/wiki/DAN_(protein)" class="mw-redirect" title="DAN (protein)">DAN (PARN)</a></li> <li><a href="/wiki/Decorin" title="Decorin">Decorin</a></li> <li><a href="/wiki/Follistatin" title="Follistatin">Follistatin</a></li> <li><a href="/wiki/Gremlin_(protein)" title="Gremlin (protein)">Gremlin (Drm)</a></li> <li><a href="/wiki/LTBP1_(gene)" title="LTBP1 (gene)">LTBP1</a></li> <li><a href="/wiki/Noggin_(protein)" title="Noggin (protein)">Noggin</a></li> <li><a href="/wiki/Transforming_growth_interacting_factor" title="Transforming growth interacting factor">TGIF</a></li> <li><a href="/wiki/Thrombospondin_1" title="Thrombospondin 1">Thrombospondin 1 (THBS1)</a></li> <li><a href="/w/index.php?title=Tomoregulin_1&action=edit&redlink=1" class="new" title="Tomoregulin 1 (page does not exist)">Tomoregulin 1</a></li></ul> <ul><li><i>Antibodies:</i> <a href="/wiki/Stamulumab" title="Stamulumab">Stamulumab</a> (against myostatin)</li> <li><a href="/wiki/TRC105" class="mw-redirect" title="TRC105">TRC105</a> (against endoglin)</li></ul> <ul><li><i>Others:</i> <a href="/wiki/Endoglin" title="Endoglin">Endoglin</a></li></ul> </div></td></tr></tbody></table></div>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1" alt="" width="1" height="1" style="border: none; position: 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