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id="p-views" class="page-actions-menu__list"> <li id="language-selector" class="page-actions-menu__list-item"> <a role="button" href="#p-lang" data-mw="interface" data-event-name="menu.languages" title="Language" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet language-selector"> Language </a> </li> <li id="page-actions-watch" class="page-actions-menu__list-item"> <a role="button" id="ca-watch" href="/w/index.php?title=Special:UserLogin&returnto=Hippo+signaling+pathway" data-event-name="menu.watch" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet menu__item--page-actions-watch"> Watch </a> </li> <li id="page-actions-edit" class="page-actions-menu__list-item"> <a role="button" id="ca-edit" href="/w/index.php?title=Hippo_signaling_pathway&action=edit" data-event-name="menu.edit" data-mw="interface" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet edit-page menu__item--page-actions-edit"> Edit </a> </li> </ul> </nav>  <div id="mw-content-subtitle"></div> </div> <div id="bodyContent" class="content"> <div id="mw-content-text" class="mw-body-content"><script>function mfTempOpenSection(id){var block=document.getElementById("mf-section-"+id);block.className+=" open-block";block.previousSibling.className+=" open-block";}</script><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><section class="mf-section-0" id="mf-section-0"> The Hippo signaling pathway, also known as the Salvador-Warts-Hippo (SWH) pathway, is a <a href="/wiki/Signaling_pathway" class="mw-redirect" title="Signaling pathway">signaling pathway</a> that controls <a href="/wiki/Organ_(anatomy)" class="mw-redirect" title="Organ (anatomy)">organ</a> size in <a href="/wiki/Animals" class="mw-redirect" title="Animals">animals</a> through the regulation of <a href="/wiki/Cell_proliferation" title="Cell proliferation">cell proliferation</a> and <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a>. The pathway takes its name from one of its key signaling components—the <a href="/wiki/Protein_kinase" title="Protein kinase">protein kinase</a> Hippo (Hpo). Mutations in this gene lead to <a href="/wiki/Tissue_(biology)" title="Tissue (biology)">tissue</a> overgrowth, or a "<a href="/wiki/Hippopotamus" title="Hippopotamus">hippopotamus</a>"-like <a href="/wiki/Phenotype" title="Phenotype">phenotype</a>. <figure typeof="mw:File/Thumb"><a href="/wiki/File:Protein_MST1_PDB_2asu.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/52/Protein_MST1_PDB_2asu.png/300px-Protein_MST1_PDB_2asu.png" decoding="async" width="300" height="254" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/52/Protein_MST1_PDB_2asu.png/450px-Protein_MST1_PDB_2asu.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/52/Protein_MST1_PDB_2asu.png/600px-Protein_MST1_PDB_2asu.png 2x" data-file-width="891" data-file-height="755"></a><figcaption><a href="/wiki/MST1" title="MST1">MST1</a>, the <a href="/wiki/Human" title="Human">human</a> homologue of the Hippo protein, is part of the Hippo signalling pathway in humans</figcaption></figure> A fundamental question in <a href="/wiki/Developmental_biology" title="Developmental biology">developmental biology</a> is how an organ knows to stop growing after reaching a particular size. Organ growth relies on several processes occurring at the cellular level, including <a href="/wiki/Cell_division" title="Cell division">cell division</a> and <a href="/wiki/Programmed_cell_death" title="Programmed cell death">programmed cell death</a> (or apoptosis). The Hippo signaling pathway is involved in restraining cell proliferation and promoting apoptosis. As many cancers are marked by unchecked cell division, this signaling pathway has become increasingly significant in the study of human <a href="/wiki/Cancer" title="Cancer">cancer</a>.<a href="#cite_note-1">[1]</a> The Hippo pathway also has a critical role in stem cell and tissue specific progenitor cell self-renewal and expansion.<a href="#cite_note-2">[2]</a> The Hippo signaling pathway appears to be <a href="/wiki/Conserved_sequence" title="Conserved sequence">highly conserved</a>. While most of the Hippo pathway components were identified in the fruit fly (<a href="/wiki/Drosophila_melanogaster" title="Drosophila melanogaster">Drosophila melanogaster</a>) using mosaic <a href="/wiki/Genetic_screen" title="Genetic screen">genetic screens</a>, <a href="/wiki/Homology_(biology)#Orthology" title="Homology (biology)">orthologs</a> to these components (genes that are related through speciation events and thus tend to retain the same function in different <a href="/wiki/Species" title="Species">species</a>) have subsequently been found in <a href="/wiki/Mammals" class="mw-redirect" title="Mammals">mammals</a>. Thus, the delineation of the pathway in Drosophila has helped to identify many genes that function as <a href="/wiki/Oncogene" title="Oncogene">oncogenes</a> or <a href="/wiki/Tumor_suppressor" class="mw-redirect" title="Tumor suppressor">tumor suppressors</a> in mammals. <div id="toc" class="toc" role="navigation" aria-labelledby="mw-toc-heading"><input type="checkbox" role="button" id="toctogglecheckbox" class="toctogglecheckbox" style="display:none"><div class="toctitle" lang="en" dir="ltr"><h2 id="mw-toc-heading">Contents</h2><label class="toctogglelabel" for="toctogglecheckbox"></label></div> <ul> <li class="toclevel-1 tocsection-1"><a href="#Mechanism">1 Mechanism</a></li> <li class="toclevel-1 tocsection-2"><a href="#In_cancer">2 In cancer</a> <ul> <li class="toclevel-2 tocsection-3"><a href="#As_a_drug_target">2.1 As a drug target</a></li> </ul> </li> <li class="toclevel-1 tocsection-4"><a href="#Regulation_of_human_organ_size">3 Regulation of human organ size</a></li> <li class="toclevel-1 tocsection-5"><a href="#Gene_name_confusion">4 Gene name confusion</a></li> <li class="toclevel-1 tocsection-6"><a href="#Summary_table">5 Summary table</a></li> <li class="toclevel-1 tocsection-7"><a href="#References">6 References</a></li> <li class="toclevel-1 tocsection-8"><a href="#Further_reading">7 Further reading</a></li> </ul> </div> </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(1)"><h2 id="Mechanism">Mechanism</h2> <a role="button" href="/w/index.php?title=Hippo_signaling_pathway&action=edit&section=1" title="Edit section: Mechanism" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div><section class="mf-section-1 collapsible-block" id="mf-section-1"> The Hippo pathway consists of a core <a href="/wiki/Kinase" title="Kinase">kinase</a> cascade in which Hpo <a href="/wiki/Phosphorylation" title="Phosphorylation">phosphorylates</a> (Drosophila) the protein kinase Warts (Wts).<a href="#cite_note-3">[3]</a><a href="#cite_note-4">[4]</a> Hpo (MST1/2 in mammals) is a member of the Ste-20 family of protein kinases. This highly conserved group of <a href="/wiki/Serine-threonine_kinase" class="mw-redirect" title="Serine-threonine kinase">serine/threonine kinases</a> regulates several cellular processes, including cell proliferation, apoptosis, and various stress responses.<a href="#cite_note-5">[5]</a> Once phosphorylated, Wts (<a href="/wiki/LATS1" title="LATS1">LATS1</a>/2 in mammals) becomes active. Misshapen (Msn, MAP4K4/6/7 in mammals) and Happyhour (Hppy, MAP4K1/2/3/5 in mammals) act in parallel to Hpo to activate Wts.<a href="#cite_note-6">[6]</a><a href="#cite_note-7">[7]</a><a href="#cite_note-8">[8]</a> Wts is a nuclear DBF-2-related kinase. These kinases are known regulators of cell cycle progression, growth, and development.<a href="#cite_note-9">[9]</a> Two proteins are known to facilitate the activation of Wts: Salvador (Sav) and Mob as tumor suppressor (Mats). Sav (<a href="/wiki/SAV1" title="SAV1">SAV1</a> in mammals) is a <a href="/wiki/WW_domain" title="WW domain">WW domain</a>-containing protein, meaning that this protein contains a sequence of <a href="/wiki/Amino_acids" class="mw-redirect" title="Amino acids">amino acids</a> in which a <a href="/wiki/Tryptophan" title="Tryptophan">tryptophan</a> and an invariant <a href="/wiki/Proline" title="Proline">proline</a> are highly conserved.<a href="#cite_note-10">[10]</a> Hpo can bind to and phosphorylate Sav, which may function as a <a href="/wiki/Scaffold_protein" title="Scaffold protein">scaffold protein</a> because this Hpo-Sav interaction promotes phosphorylation of Wts.<a href="#cite_note-11">[11]</a> Hpo can also phosphorylate and activate Mats (MOBKL1A/B in mammals), which allows Mats to associate with and strengthen the kinase activity of Wts.<a href="#cite_note-12">[12]</a> Activated Wts can then go on to phosphorylate and inactivate the <a href="/wiki/Coactivator_(genetics)" title="Coactivator (genetics)">transcriptional coactivator</a> Yorkie (Yki). Yki is unable to bind DNA by itself. In its active state, Yki binds to the transcription factor Scalloped (Sd), and the Yki-Sd complex becomes localized to the nucleus. This allows for the expression of several genes that promote organ growth, such as <a href="/wiki/Cyclin_E" title="Cyclin E">cyclin E</a>, which promotes cell cycle progression, and diap1 (Drosophila inhibitor of apoptosis protein-1), which, as its name suggests, prevents apoptosis.<a href="#cite_note-13">[13]</a> Yki also activates expression of the bantam <a href="/wiki/MicroRNA" title="MicroRNA">microRNA</a>, a positive growth regulator that specifically affects cell number.<a href="#cite_note-14">[14]</a><a href="#cite_note-15">[15]</a> Thus, the inactivation of Yki by Wts inhibits growth through the transcriptional repression of these pro-growth regulators. By phosphorylating Yki at serine 168, Wts promotes the association of Yki with <a href="/wiki/14-3-3_proteins" class="mw-redirect" title="14-3-3 proteins">14-3-3 proteins</a>, which help to anchor Yki in the <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a> and prevent its transport to the nucleus. In mammals, the two Yki orthologs are Yes-associated protein (YAP) and transcriptional coactivator with PDZ-binding motif (<a href="/wiki/WWTR1" title="WWTR1">WWTR1, also known as TAZ</a>).<a href="#cite_note-16">[16]</a> When activated, YAP and TAZ can bind to several transcription factors including <a href="/wiki/P73" title="P73">p73</a>, <a href="/wiki/Runx2" class="mw-redirect" title="Runx2">Runx2</a> and several TEADs.<a href="#cite_note-17">[17]</a> YAP regulates the expression of Hoxa1 and Hoxc13 in mouse and human epithelial cells in vivo and in vitro.<a href="#cite_note-18">[18]</a> The upstream regulators of the core Hpo/Wts kinase cascade include the <a href="/wiki/Transmembrane_protein" title="Transmembrane protein">transmembrane protein</a> <a href="/wiki/FAT_(gene)" class="mw-redirect" title="FAT (gene)">Fat</a> and several membrane-associated proteins. As an atypical <a href="/wiki/Cadherin" title="Cadherin">cadherin</a>, Fat (FAT1-4 in mammals) may function as a receptor, though an extracellular <a href="/wiki/Ligand" title="Ligand">ligand</a> has not been positively identified. The <a href="/wiki/Lipid-anchored_protein" title="Lipid-anchored protein">GPI</a>-anchored cell surface protein glypican-3 (GPC3) is known to interact with Fat1 in human liver cancer.<a href="#cite_note-19">[19]</a> GPC3 is also shown to modulate Yap signaling in liver cancer.<a href="#cite_note-20">[20]</a> While Fat is known to bind to another atypical cadherin, <a href="/wiki/DCHS1" title="DCHS1">Dachsous</a> (Ds), during tissue patterning,<a href="#cite_note-21">[21]</a> it is unclear what role Ds has in regulating tissue growth. Nevertheless, Fat is recognized as an upstream regulator of the Hpo pathway. Fat activates Hpo through the apical protein Expanded (Ex; FRMD6/Willin in mammals). Ex interacts with two other apically-localized proteins, Kibra (<a href="/wiki/WWC1" title="WWC1">KIBRA</a> in mammals) and <a href="/wiki/Merlin_(protein)" title="Merlin (protein)">Merlin</a> (Mer; NF2 in mammals), to form the Kibra-Ex-Mer (KEM) complex. Both Ex and Mer are <a href="/wiki/FERM_domain" title="FERM domain">FERM domain</a>-containing proteins, while Kibra, like Sav, is a WW domain-containing protein.<a href="#cite_note-22">[22]</a> The KEM complex physically interacts with the Hpo kinase cascade, thereby localizing the core kinase cascade to the plasma membrane for activation.<a href="#cite_note-23">[23]</a> Fat may also regulate Wts independently of Ex/Hpo, through the inhibition of the unconventional <a href="/wiki/Myosin" title="Myosin">myosin</a> Dachs. Normally, Dachs can bind to and promote the degradation of Wts.<a href="#cite_note-24">[24]</a> </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(2)"><h2 id="In_cancer">In cancer</h2> <a role="button" href="/w/index.php?title=Hippo_signaling_pathway&action=edit&section=2" title="Edit section: In cancer" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div><section class="mf-section-2 collapsible-block" id="mf-section-2"> In fruitfly, the Hippo signaling pathway involves a kinase cascade involving the Salvador (Sav), Warts (Wts) and Hippo (Hpo) <a href="/wiki/Protein_kinases" class="mw-redirect" title="Protein kinases">protein kinases</a>.<a href="#cite_note-25">[25]</a> Many of the genes involved in the Hippo signaling pathway are recognized as <a href="/wiki/Tumor_suppressor_gene" title="Tumor suppressor gene">tumor suppressors</a>, while Yki/YAP/TAZ is identified as an <a href="/wiki/Oncogene" title="Oncogene">oncogene</a>. YAP/TAZ can reprogram cancer cells into <a href="/wiki/Cancer_stem_cell" title="Cancer stem cell">cancer stem cells</a>.<a href="#cite_note-pmid25287865-26">[26]</a> YAP has been found to be elevated in some human cancers, including <a href="/wiki/Breast_cancer" title="Breast cancer">breast cancer</a>, <a href="/wiki/Colorectal_cancer" title="Colorectal cancer">colorectal cancer</a>, and <a href="/wiki/Liver_cancer" title="Liver cancer">liver cancer</a>.<a href="#cite_note-27">[27]</a><a href="#cite_note-28">[28]</a><a href="#cite_note-29">[29]</a> This may be explained by YAP’s recently defined role in overcoming <a href="/wiki/Contact_inhibition" title="Contact inhibition">contact inhibition</a>, a fundamental growth control property of normal cells in vitro and in vivo, in which proliferation stops after cells reach <a href="/wiki/Confluency" title="Confluency">confluence</a><a href="#cite_note-30">[30]</a> (in culture) or occupy maximum available space inside the body and touch one another. This property is typically lost in cancerous cells, allowing them to proliferate in an uncontrolled manner.<a href="#cite_note-31">[31]</a> In fact, YAP overexpression antagonizes contact inhibition.<a href="#cite_note-32">[32]</a> Many of the pathway components recognized as tumor suppressor genes are mutated in human cancers. For example, mutations in Fat4 have been found in breast cancer,<a href="#cite_note-33">[33]</a> while NF2 is mutated in familial and sporadic <a href="/wiki/Schwannoma" title="Schwannoma">schwannomas</a>.<a href="#cite_note-34">[34]</a> Additionally, several human cancer cell lines invoke mutations of the SAV1 and MOBK1B proteins.<a href="#cite_note-35">[35]</a><a href="#cite_note-36">[36]</a> However, recent research by <a href="/wiki/Marc_Kirschner" title="Marc Kirschner">Marc Kirschner</a> and Taran Gujral has demonstrated that Hippo pathway components may play a more nuanced role in cancer than previously thought. Hippo pathway inactivation enhanced the effect of 15 FDA-approved oncology drugs by promoting chemo-retention.<a href="#cite_note-37">[37]</a> In another study, the Hippo pathway kinases LATS1/2 were found to suppress cancer immunity in mice.<a href="#cite_note-38">[38]</a> Not all studies, however, support a role for Hippo signaling in promoting carcinogenesis. In <a href="/wiki/Hepatocellular_carcinoma" title="Hepatocellular carcinoma">hepatocellular carcinoma</a>, for instance, it was suggesting that <a href="/wiki/AXIN1" title="AXIN1">AXIN1</a> mutations would provoke Hippo signaling pathway activation, fostering the cancer development, but a recent study demonstrated that such an effect cannot be detected.<a href="#cite_note-39">[39]</a> Thus the exact role of Hippo signaling in the cancer process awaits further elucidation. <div class="mw-heading mw-heading3"><h3 id="As_a_drug_target">As a drug target</h3> <a role="button" href="/w/index.php?title=Hippo_signaling_pathway&action=edit&section=3" title="Edit section: As a drug target" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div> Two venture-backed oncology startups, Vivace Therapeutics and the General Biotechnologies subsidiary Nivien Therapeutics, are actively developing <a href="/wiki/Kinase_inhibitor" class="mw-redirect" title="Kinase inhibitor">kinase inhibitors</a> targeting the Hippo pathway.<a href="#cite_note-40">[40]</a> </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(3)"><h2 id="Regulation_of_human_organ_size">Regulation of human organ size</h2> <a role="button" href="/w/index.php?title=Hippo_signaling_pathway&action=edit&section=4" title="Edit section: Regulation of human organ size" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div><section class="mf-section-3 collapsible-block" id="mf-section-3"> The heart is the first organ formed during mammalian development. A properly sized and functional heart is vital throughout the entire lifespan. Loss of cardiomyocytes because of injury or diseases leads to heart failure, which is a major cause of human morbidity and mortality. Unfortunately, regenerative potential of the adult heart is limited. The Hippo pathway is a recently identified signaling cascade that plays an evolutionarily conserved role in organ size control by inhibiting cell proliferation, promoting apoptosis, regulating fates of stem/progenitor cells, and in some circumstances, limiting cell size. Research indicates a key role of this pathway in regulation of <a href="/wiki/Cardiomyocyte_proliferation" title="Cardiomyocyte proliferation">cardiomyocyte proliferation</a> and heart size. Inactivation of the Hippo pathway or activation of its downstream effector, the Yes-associated protein transcription coactivator, improves cardiac regeneration. Several known upstream signals of the Hippo pathway such as mechanical stress, G-protein-coupled receptor signaling, and oxidative stress are known to play critical roles in cardiac physiology. In addition, Yes-associated protein has been shown to regulate cardiomyocyte fate through multiple transcriptional mechanisms.<a href="#cite_note-41">[41]</a><a href="#cite_note-42">[42]</a><a href="#cite_note-43">[43]</a> </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(4)"><h2 id="Gene_name_confusion">Gene name confusion</h2> <a role="button" href="/w/index.php?title=Hippo_signaling_pathway&action=edit&section=5" title="Edit section: Gene name confusion" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div><section class="mf-section-4 collapsible-block" id="mf-section-4"> Note that Hippo TAZ protein is often confused with the gene TAZ, which is unrelated to the Hippo pathway. The gene TAZ produces the protein tafazzin. The official gene name for the Hippo TAZ protein is WWTR1. Also, the official names for MST1 and MST2 are STK4 and STK3, respectively. All databases for bioinformatics use the official gene symbols, and commercial sources for <a href="/wiki/PCR_primer" class="mw-redirect" title="PCR primer">PCR primers</a> or <a href="/wiki/SiRNA" class="mw-redirect" title="SiRNA">siRNA</a> also go by the official gene names. </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(5)"><h2 id="Summary_table">Summary table</h2> <a role="button" href="/w/index.php?title=Hippo_signaling_pathway&action=edit&section=6" title="Edit section: Summary table" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div><section class="mf-section-5 collapsible-block" id="mf-section-5"> <table class="wikitable"> <tbody><tr> <th>Drosophila melanogaster </th> <th>Human ortholog(s) </th> <th>Protein description and role in Hippo signaling pathway </th></tr> <tr> <td>Dachsous (Ds) </td> <td><a href="/wiki/DCHS1" title="DCHS1">DCHS1</a>, <a href="/wiki/DCHS2" title="DCHS2">DCHS2</a> </td> <td>Atypical cadherin that may act as a ligand for the Fat receptor </td></tr> <tr> <td>Fat (Ft) </td> <td><a href="/wiki/FAT_(gene)" class="mw-redirect" title="FAT (gene)">FAT1</a>, <a href="/w/index.php?title=FAT2&action=edit&redlink=1" class="new" title="FAT2 (page does not exist)">FAT2</a>, <a href="/w/index.php?title=FAT3&action=edit&redlink=1" class="new" title="FAT3 (page does not exist)">FAT3</a>, <a href="/wiki/FAT4" title="FAT4">FAT4</a> </td> <td>Atypical cadherin that may act as a receptor for the Hippo pathway </td></tr> <tr> <td>Expanded (Ex) </td> <td><a href="/wiki/FRMD6" title="FRMD6">FRMD6</a> </td> <td>FERM domain-containing apical protein that associates with Kibra and Mer as an upstream regulator of the core kinase cascade </td></tr> <tr> <td>Dachs (Dachs) </td> <td> </td> <td>Unconventional myosin that can bind Wts, promoting its degradation </td></tr> <tr> <td>Kibra (Kibra) </td> <td><a href="/wiki/WWC1" title="WWC1">WWC1</a> </td> <td>WW domain-containing apical protein that associates with Ex and Mer as an upstream regulator of the core kinase cascade </td></tr> <tr> <td>Merlin (Mer) </td> <td><a href="/wiki/Merlin_(protein)" title="Merlin (protein)">NF2</a> </td> <td>FERM domain-containing apical protein that associates with Ex and Kibra as an upstream regulator of the core kinase cascade </td></tr> <tr> <td>Hippo (Hpo) </td> <td><a href="/wiki/MST1" title="MST1">MST1</a>, <a href="/wiki/MST2" class="mw-redirect" title="MST2">MST2</a> – officially STK4/3 </td> <td>Sterile-20-type kinase that phosphorylates and activates Wts </td></tr> <tr> <td>Salvador (Sav) </td> <td><a href="/wiki/SAV1" title="SAV1">SAV1</a> </td> <td>WW domain-containing protein that may act as a scaffold protein, facilitating Warts phosphorylation by Hippo </td></tr> <tr> <td>Warts (Wts) </td> <td><a href="/wiki/LATS1" title="LATS1">LATS1</a>, <a href="/wiki/LATS2" title="LATS2">LATS2</a> </td> <td>Nuclear DBF-2-related kinase that phosphorylates and inactivates Yki </td></tr> <tr> <td>Mob as tumor suppressor (Mats) </td> <td><a href="/wiki/MOBKL1A" title="MOBKL1A">MOBKL1A</a>, <a href="/wiki/MOBKL1B" class="mw-redirect" title="MOBKL1B">MOBKL1B</a> </td> <td>Kinase that associates with Wts to potentiate its catalytic activity </td></tr> <tr> <td>Yorkie (Yki) </td> <td><a href="/wiki/YAP1" title="YAP1">YAP</a>, TAZ – officially <a href="/wiki/WWTR1" title="WWTR1">WWTR1</a> </td> <td>Transcriptional coactivator that binds to Sd in its active, unphosphorylated form to activate expression of transcriptional targets that promote cell growth, cell proliferation, and prevent apoptosis </td></tr> <tr> <td>Scalloped (Sd) </td> <td><a href="/wiki/TEAD1" title="TEAD1">TEAD1</a>, <a href="/wiki/TEAD2" title="TEAD2">TEAD2</a>, <a href="/wiki/TEAD3" title="TEAD3">TEAD3</a>, <a href="/wiki/TEAD4" title="TEAD4">TEAD4</a> </td> <td>Transcription factor that binds Yki to regulate target gene expression </td></tr></tbody></table> </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(6)"><h2 id="References">References</h2> <a role="button" href="/w/index.php?title=Hippo_signaling_pathway&action=edit&section=7" title="Edit section: References" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div><section class="mf-section-6 collapsible-block" id="mf-section-6"> <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"><a href="#cite_ref-1">^</a> <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 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Cell & Bioscience. 3 (1): 31. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1186%2F2045-3701-3-31">10.1186/2045-3701-3-31</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3849747">3849747</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/23985272">23985272</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Cell+%26+Bioscience&rft.atitle=Mst1+and+Mst2+kinases%3A+regulations+and+diseases&rft.volume=3&rft.issue=1&rft.pages=31&rft.date=2013-08&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3849747%23id-name%3DPMC&rft_id=info%3Apmid%2F23985272&rft_id=info%3Adoi%2F10.1186%2F2045-3701-3-31&rft.aulast=Qin&rft.aufirst=F&rft.au=Tian%2C+J&rft.au=Zhou%2C+D&rft.au=Chen%2C+L&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3849747&rfr_id=info%3Asid%2Fen.wikipedia.org%3AHippo+signaling+pathway" class="Z3988"> </li> <li id="cite_note-42"><a href="#cite_ref-42">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHilmanGat2011" class="citation journal cs1">Hilman D, Gat U (August 2011). <a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Fmolbev%2Fmsr065">"The evolutionary history of YAP and the hippo/YAP pathway"</a>. Molecular Biology and Evolution. 28 (8): 2403–17. <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%2Fmolbev%2Fmsr065">10.1093/molbev/msr065</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/21415026">21415026</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Molecular+Biology+and+Evolution&rft.atitle=The+evolutionary+history+of+YAP+and+the+hippo%2FYAP+pathway&rft.volume=28&rft.issue=8&rft.pages=2403-17&rft.date=2011-08&rft_id=info%3Adoi%2F10.1093%2Fmolbev%2Fmsr065&rft_id=info%3Apmid%2F21415026&rft.aulast=Hilman&rft.aufirst=D&rft.au=Gat%2C+U&rft_id=https%3A%2F%2Fdoi.org%2F10.1093%252Fmolbev%252Fmsr065&rfr_id=info%3Asid%2Fen.wikipedia.org%3AHippo+signaling+pathway" class="Z3988"> </li> <li id="cite_note-43"><a href="#cite_ref-43">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBoopathyHong2019" class="citation journal cs1">Boopathy GT, Hong W (2019). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6468149">"Role of Hippo Pathway-YAP/TAZ Signaling in Angiogenesis"</a>. Frontiers in Cell and Developmental Biology. 7: 49. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.3389%2Ffcell.2019.00049">10.3389/fcell.2019.00049</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/2296-634X">2296-634X</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6468149">6468149</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/31024911">31024911</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Frontiers+in+Cell+and+Developmental+Biology&rft.atitle=Role+of+Hippo+Pathway-YAP%2FTAZ+Signaling+in+Angiogenesis&rft.volume=7&rft.pages=49&rft.date=2019&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC6468149%23id-name%3DPMC&rft.issn=2296-634X&rft_id=info%3Apmid%2F31024911&rft_id=info%3Adoi%2F10.3389%2Ffcell.2019.00049&rft.aulast=Boopathy&rft.aufirst=Gandhi+T.+K.&rft.au=Hong%2C+Wanjin&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC6468149&rfr_id=info%3Asid%2Fen.wikipedia.org%3AHippo+signaling+pathway" class="Z3988"> </li> </ol></div> </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(7)"><h2 id="Further_reading">Further reading</h2> <a role="button" href="/w/index.php?title=Hippo_signaling_pathway&action=edit&section=8" title="Edit section: Further reading" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> edit </a> </div><section class="mf-section-7 collapsible-block" id="mf-section-7"> <style data-mw-deduplicate="TemplateStyles:r1239549316">.mw-parser-output .refbegin{margin-bottom:0.5em}.mw-parser-output .refbegin-hanging-indents>ul{margin-left:0}.mw-parser-output .refbegin-hanging-indents>ul>li{margin-left:0;padding-left:3.2em;text-indent:-3.2em}.mw-parser-output .refbegin-hanging-indents ul,.mw-parser-output .refbegin-hanging-indents ul li{list-style:none}@media(max-width:720px){.mw-parser-output .refbegin-hanging-indents>ul>li{padding-left:1.6em;text-indent:-1.6em}}.mw-parser-output .refbegin-columns{margin-top:0.3em}.mw-parser-output .refbegin-columns ul{margin-top:0}.mw-parser-output .refbegin-columns li{page-break-inside:avoid;break-inside:avoid-column}@media screen{.mw-parser-output .refbegin{font-size:90%}}</style><div class="refbegin" style=""> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPatelCamargoYimlamai2017" class="citation journal cs1">Patel SH, Camargo FD, Yimlamai D (February 2017). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5285449">"Hippo Signaling in the Liver Regulates Organ Size, Cell Fate, and Carcinogenesis"</a>. Gastroenterology. 152 (3): 533–545. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1053%2Fj.gastro.2016.10.047">10.1053/j.gastro.2016.10.047</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5285449">5285449</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/28003097">28003097</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Gastroenterology&rft.atitle=Hippo+Signaling+in+the+Liver+Regulates+Organ+Size%2C+Cell+Fate%2C+and+Carcinogenesis&rft.volume=152&rft.issue=3&rft.pages=533-545&rft.date=2017-02&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5285449%23id-name%3DPMC&rft_id=info%3Apmid%2F28003097&rft_id=info%3Adoi%2F10.1053%2Fj.gastro.2016.10.047&rft.aulast=Patel&rft.aufirst=SH&rft.au=Camargo%2C+FD&rft.au=Yimlamai%2C+D&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5285449&rfr_id=info%3Asid%2Fen.wikipedia.org%3AHippo+signaling+pathway" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGongZhangZouTian2019" class="citation journal cs1">Gong P, Zhang Z, Zou C, Tian Q, Chen X, Hong M, et al. (January 2019). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6193462">"Hippo/YAP signaling pathway mitigates blood-brain barrier disruption after cerebral ischemia/reperfusion injury"</a>. Behavioural Brain Research. 356: 8–17. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.bbr.2018.08.003">10.1016/j.bbr.2018.08.003</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6193462">6193462</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/30092249">30092249</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Behavioural+Brain+Research&rft.atitle=Hippo%2FYAP+signaling+pathway+mitigates+blood-brain+barrier+disruption+after+cerebral+ischemia%2Freperfusion+injury&rft.volume=356&rft.pages=8-17&rft.date=2019-01&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC6193462%23id-name%3DPMC&rft_id=info%3Apmid%2F30092249&rft_id=info%3Adoi%2F10.1016%2Fj.bbr.2018.08.003&rft.aulast=Gong&rft.aufirst=P&rft.au=Zhang%2C+Z&rft.au=Zou%2C+C&rft.au=Tian%2C+Q&rft.au=Chen%2C+X&rft.au=Hong%2C+M&rft.au=Liu%2C+X&rft.au=Chen%2C+Q&rft.au=Xu%2C+Z&rft.au=Li%2C+M&rft.au=Wang%2C+J&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC6193462&rfr_id=info%3Asid%2Fen.wikipedia.org%3AHippo+signaling+pathway" class="Z3988"></li> <li>Valentina Rausch, Carsten G. Hansen (2020). <a rel="nofollow" class="external text" href="https://www.cell.com/trends/cell-biology/fulltext/S0962-8924(19)30168-0">The Hippo Pathway, YAP/TAZ, and the Plasma Membrane</a>. Trends in Cell Biology <a rel="nofollow" class="external free" href="https://doi.org/10.1016/j.tcb.2019.10.005">https://doi.org/10.1016/j.tcb.2019.10.005</a></li></ul> </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 dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl ol,.mw-parser-output .hlist dl ul,.mw-parser-output .hlist ol dl,.mw-parser-output .hlist ol ol,.mw-parser-output .hlist ol ul,.mw-parser-output .hlist ul dl,.mw-parser-output .hlist ul ol,.mw-parser-output .hlist ul ul{display:inline}.mw-parser-output .hlist .mw-empty-li{display:none}.mw-parser-output .hlist dt::after{content:": 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<h4>Languages</h4> <section> <ul id="p-variants" class="minerva-languages"></ul> <ul class="minerva-languages"><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Voie_de_signalisation_Hippo" title="Voie de signalisation Hippo – French" lang="fr" hreflang="fr" data-title="Voie de signalisation Hippo" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target">Français</a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Via_di_segnalazione_Hippo" title="Via di segnalazione Hippo – Italian" lang="it" hreflang="it" data-title="Via di segnalazione Hippo" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target">Italiano</a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/Hippo%E3%82%B7%E3%82%B0%E3%83%8A%E3%83%AB%E4%BC%9D%E9%81%94%E7%B5%8C%E8%B7%AF" title="Hippoシグナル伝達経路 – Japanese" lang="ja" hreflang="ja" data-title="Hippoシグナル伝達経路" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target">日本語</a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/%C5%9Acie%C5%BCka_sygna%C5%82owa_Hippo" title="Ścieżka sygnałowa Hippo – Polish" lang="pl" hreflang="pl" data-title="Ścieżka sygnałowa Hippo" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target">Polski</a></li></ul> </section> </div> <div class="minerva-footer-logo"><img src="/static/images/mobile/copyright/wikipedia-wordmark-en.svg" alt="Wikipedia" width="120" height="18" style="width: 7.5em; height: 1.125em;"/> </div> <ul id="footer-info" class="footer-info hlist hlist-separated"> <li id="footer-info-lastmod"> This page was last edited on 10 October 2024, at 14:19 (UTC).</li> <li id="footer-info-copyright">Content is available under <a class="external" 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