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title="Morphogen – German" lang="de" hreflang="de" data-title="Morphogen" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Morf%C3%B3geno" title="Morfógeno – Spanish" lang="es" hreflang="es" data-title="Morfógeno" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Morphog%C3%A8ne" title="Morphogène – French" lang="fr" hreflang="fr" data-title="Morphogène" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Morfogeno" title="Morfogeno – Italian" lang="it" hreflang="it" data-title="Morfogeno" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%9E%D7%95%D7%A8%D7%A4%D7%95%D7%92%D7%9F" title="מורפוגן – Hebrew" lang="he" hreflang="he" data-title="מורפוגן" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E3%83%A2%E3%83%AB%E3%83%95%E3%82%A9%E3%82%B2%E3%83%B3" title="モルフォゲン – Japanese" lang="ja" hreflang="ja" data-title="モルフォゲン" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Morfogen" title="Morfogen – Norwegian Bokmål" lang="nb" hreflang="nb" 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Unsourced material may be challenged and removed.<br /><small><span class="plainlinks"><i>Find sources:</i> <a rel="nofollow" class="external text" href="https://www.google.com/search?as_eq=wikipedia&q=%22Morphogen%22">"Morphogen"</a> – <a rel="nofollow" class="external text" href="https://www.google.com/search?tbm=nws&q=%22Morphogen%22+-wikipedia&tbs=ar:1">news</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.google.com/search?&q=%22Morphogen%22&tbs=bkt:s&tbm=bks">newspapers</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.google.com/search?tbs=bks:1&q=%22Morphogen%22+-wikipedia">books</a> <b>·</b> <a rel="nofollow" class="external text" href="https://scholar.google.com/scholar?q=%22Morphogen%22">scholar</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.jstor.org/action/doBasicSearch?Query=%22Morphogen%22&acc=on&wc=on">JSTOR</a></span></small></span> <span class="date-container"><i>(<span class="date">May 2008</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1251242444"><table class="box-Expert_needed plainlinks metadata ambox ambox-content" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/b/b4/Ambox_important.svg/40px-Ambox_important.svg.png" decoding="async" width="40" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/b/b4/Ambox_important.svg/60px-Ambox_important.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/b/b4/Ambox_important.svg/80px-Ambox_important.svg.png 2x" data-file-width="40" data-file-height="40" /></span></span></div></td><td class="mbox-text"><div class="mbox-text-span">This article <b>needs attention from an expert in Biology</b>. The specific problem is: <b>Out of date, rightly flagged since 2008, needs references.</b> See the <a href="/wiki/Talk:Morphogen#Biology" title="Talk:Morphogen">talk page</a> for details.<span class="hide-when-compact"> <a href="/wiki/Wikipedia:WikiProject_Biology" title="Wikipedia:WikiProject Biology">WikiProject Biology</a> may be able to help recruit an expert.</span> <span class="date-container"><i>(<span class="date">August 2015</span>)</i></span></div></td></tr></tbody></table> </div> </div><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Drosophila_melanogaster_laboratory_culture-vial.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/16/Drosophila_melanogaster_laboratory_culture-vial.jpg/220px-Drosophila_melanogaster_laboratory_culture-vial.jpg" decoding="async" width="220" height="311" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/16/Drosophila_melanogaster_laboratory_culture-vial.jpg/330px-Drosophila_melanogaster_laboratory_culture-vial.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/16/Drosophila_melanogaster_laboratory_culture-vial.jpg/440px-Drosophila_melanogaster_laboratory_culture-vial.jpg 2x" data-file-width="2492" data-file-height="3520" /></a><figcaption>Morphogenesis of <i><a href="/wiki/Drosophila" title="Drosophila">Drosophila</a></i> fruit flies is intensively studied in the laboratory</figcaption></figure> <p>A <b>morphogen</b> is a substance whose non-uniform distribution governs the <a href="/wiki/Natural_patterns" class="mw-redirect" title="Natural patterns">pattern</a> of tissue development in the process of <a href="/wiki/Morphogenesis" title="Morphogenesis">morphogenesis</a> or <a href="/wiki/Pattern_formation" title="Pattern formation">pattern formation</a>, one of the core processes of <a href="/wiki/Developmental_biology" title="Developmental biology">developmental biology</a>, establishing positions of the various specialized cell types within a tissue. More specifically, a morphogen is a signaling molecule that acts directly on cells to produce specific cellular responses depending on its local concentration. </p><p>Typically, morphogens are produced by source cells and diffuse through surrounding tissues in an embryo during early development, such that concentration gradients are set up. These gradients drive the process of differentiation of unspecialised <a href="/wiki/Stem_cell" title="Stem cell">stem cells</a> into different cell types, ultimately forming all the tissues and organs of the body. The control of morphogenesis is a central element in <a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">evolutionary developmental biology</a> (evo-devo). </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="History">History</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Morphogen&action=edit&section=1" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The term was coined by <a href="/wiki/Alan_Turing" title="Alan Turing">Alan Turing</a> in the paper "<a href="/wiki/The_Chemical_Basis_of_Morphogenesis" title="The Chemical Basis of Morphogenesis">The Chemical Basis of Morphogenesis</a>", where he predicted a chemical mechanism for biological <a href="/wiki/Pattern_formation" title="Pattern formation">pattern formation</a>,<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> decades before the formation of such patterns was demonstrated.<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><p>The concept of the morphogen has a long history in developmental biology, dating back to the work of the pioneering <i><a href="/wiki/Drosophila" title="Drosophila">Drosophila</a></i> (fruit fly) <a href="/wiki/Geneticist" title="Geneticist">geneticist</a>, <a href="/wiki/Thomas_Hunt_Morgan" title="Thomas Hunt Morgan">Thomas Hunt Morgan</a>, in the early 20th century. <a href="/wiki/Lewis_Wolpert" title="Lewis Wolpert">Lewis Wolpert</a> refined the morphogen concept in the 1960s with the <a href="/wiki/French_flag_model" title="French flag model">French flag model</a>, which described how a morphogen could subdivide a tissue into domains of different target <a href="/wiki/Gene_expression" title="Gene expression">gene expression</a> (corresponding to the colours of the French flag). This model was championed by the leading <i>Drosophila</i> biologist, <a href="/wiki/Peter_Lawrence_(biologist)" class="mw-redirect" title="Peter Lawrence (biologist)">Peter Lawrence</a>. <a href="/wiki/Christiane_N%C3%BCsslein-Volhard" title="Christiane Nüsslein-Volhard">Christiane Nüsslein-Volhard</a> was the first to identify a morphogen, <a href="/wiki/Bicoid" class="mw-redirect" title="Bicoid">Bicoid</a>, one of the <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factors</a> present in a gradient in the <i>Drosophila</i> <a href="/wiki/Syncitial" class="mw-redirect" title="Syncitial">syncitial</a> embryo. She was awarded the 1995 <a href="/wiki/Nobel_Prize_in_Physiology_and_Medicine" class="mw-redirect" title="Nobel Prize in Physiology and Medicine">Nobel Prize in Physiology and Medicine</a> for her work explaining the morphogenic <a href="/wiki/Embryology" title="Embryology">embryology</a> of the common fruit fly.<sup id="cite_ref-Nusslein_3-0" class="reference"><a href="#cite_note-Nusslein-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Arthur2002_4-0" class="reference"><a href="#cite_note-Arthur2002-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Winchester_5-0" class="reference"><a href="#cite_note-Winchester-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> Groups led by Gary Struhl and Stephen Cohen then demonstrated that a secreted signalling protein, <a href="/wiki/Decapentaplegic" title="Decapentaplegic">decapentaplegic</a> (the <i>Drosophila</i> homologue of <a href="/wiki/Transforming_growth_factor_beta" title="Transforming growth factor beta">transforming growth factor beta</a>), acted as a morphogen during the later stages of <i>Drosophila</i> development. </p> <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=Morphogen&action=edit&section=2" title="Edit section: Mechanism"><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">Further information: <a href="/wiki/Drosophila_embryogenesis" title="Drosophila embryogenesis">Drosophila embryogenesis</a></div> <p>During early development, morphogen gradients result in the differentiation of specific <a href="/wiki/Cell_type" title="Cell type">cell types</a> in a distinct spatial order. The morphogen provides spatial information by forming a <a href="/wiki/Concentration_gradient" class="mw-redirect" title="Concentration gradient">concentration gradient</a> that subdivides a field of cells by inducing or maintaining the <a href="/wiki/Gene_expression" title="Gene expression">expression</a> of different target <a href="/wiki/Genes" class="mw-redirect" title="Genes">genes</a> at distinct concentration thresholds. Thus, cells far from the source of the morphogen will receive low levels of morphogen and express only low-threshold target <a href="/wiki/Gene" title="Gene">genes</a>. In contrast, cells close to the source of morphogen will receive high levels of morphogen and will express both low- and high-threshold target genes. Distinct cell types emerge as a consequence of the different combination of target gene expression. In this way, the field of cells is subdivided into different types according to their position relative to the source of the morphogen. This model is assumed to be a general mechanism by which cell type diversity can be generated in <a href="/wiki/Embryonic_development" class="mw-redirect" title="Embryonic development">embryonic development</a> in animals. </p><p>Some of the earliest and best-studied morphogens are <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factors</a> that <a href="/wiki/Diffusion" title="Diffusion">diffuse</a> within early <i><a href="/wiki/Drosophila_melanogaster" title="Drosophila melanogaster">Drosophila melanogaster</a></i> (fruit fly) embryos. However, most morphogens are <a href="/wiki/Secretion" title="Secretion">secreted</a> proteins that <a href="/wiki/Cell_signalling" class="mw-redirect" title="Cell signalling">signal between cells</a>. </p> <div class="mw-heading mw-heading2"><h2 id="Genes_and_signals">Genes and signals</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Morphogen&action=edit&section=3" title="Edit section: Genes and signals"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A morphogen spreads from a localized source and forms a concentration gradient across a developing tissue.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> In developmental biology, 'morphogen' is rigorously used to mean a signalling molecule that acts directly on cells (not through serial induction) to produce specific cellular responses that depend on morphogen concentration. This definition concerns the mechanism, not any specific chemical formula, so simple compounds such as <a href="/wiki/Retinoic_acid" title="Retinoic acid">retinoic acid</a> (the active metabolite of <a href="/wiki/Retinol" title="Retinol">retinol</a> or <a href="/wiki/Vitamin_A" title="Vitamin A">vitamin A</a>) may also act as morphogens. The model is not universally accepted due to specific issues with setting up a gradient in the tissue outlined in the <a href="/wiki/French_flag_model" title="French flag model">French flag model</a><sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> and subsequent work showing that the morphogen gradient of the Drosophila embryo is more complex than the simple gradient model would indicate.<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> </p> <div class="mw-heading mw-heading2"><h2 id="Examples">Examples</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Morphogen&action=edit&section=4" title="Edit section: Examples"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Proposed mammalian morphogens include <a href="/wiki/Retinoic_acid" title="Retinoic acid">retinoic acid</a>, sonic hedgehog (<a href="/wiki/Sonic_hedgehog" class="mw-redirect" title="Sonic hedgehog">SHH</a>), transforming growth factor beta (<a href="/wiki/TGF-%CE%B2" class="mw-redirect" title="TGF-β">TGF-β</a>)/bone morphogenic protein (<a href="/wiki/Bone_morphogenic_protein" class="mw-redirect" title="Bone morphogenic protein">BMP</a>), and <a href="/wiki/Wnt_signaling_pathway" title="Wnt signaling pathway">Wnt</a>/<a href="/wiki/Beta-catenin" class="mw-redirect" title="Beta-catenin">beta-catenin</a>.<sup id="cite_ref-Kam_2012_10-0" class="reference"><a href="#cite_note-Kam_2012-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Moore_2013_11-0" class="reference"><a href="#cite_note-Moore_2013-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Morphogens in <i><a href="/wiki/Drosophila" title="Drosophila">Drosophila</a></i> include <a href="/wiki/Decapentaplegic" title="Decapentaplegic">decapentaplegic</a> and <a href="/wiki/Hedgehog_(cell_signaling)" class="mw-redirect" title="Hedgehog (cell signaling)">hedgehog</a>.<sup id="cite_ref-Kam_2012_10-1" class="reference"><a href="#cite_note-Kam_2012-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p><p>During development, <a href="/wiki/Retinoic_acid" title="Retinoic acid">retinoic acid</a>, a metabolite of <a href="/wiki/Vitamin_A" title="Vitamin A">vitamin A</a>, is used to stimulate the growth of the <a href="/wiki/Posterior_(anatomy)" class="mw-redirect" title="Posterior (anatomy)">posterior</a> end of the organism.<sup id="cite_ref-Cunningham_and_Duester_12-0" class="reference"><a href="#cite_note-Cunningham_and_Duester-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> Retinoic acid binds to <a href="/wiki/Retinoic_acid_receptor" title="Retinoic acid receptor">retinoic acid receptors</a> that acts as transcription factors to regulate the expression of <a href="/wiki/Hox_gene" title="Hox gene">Hox genes</a>. Exposure of embryos to exogenous retinoids especially in the first trimester results in birth defects.<sup id="cite_ref-Moore_2013_11-1" class="reference"><a href="#cite_note-Moore_2013-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/TGF-%CE%B2" class="mw-redirect" title="TGF-β">TGF-β</a> family members are involved in <a href="/wiki/Neural_tube#Dorsal-ventral_patterning" title="Neural tube">dorsoventral patterning</a> and the formation of some organs. Binding to TGF-β to type II <a href="/wiki/TGF_beta_receptors" class="mw-redirect" title="TGF beta receptors">TGF beta receptors</a> recruits type I receptors causing the latter to be transphosphorylated. The type I receptors activate <a href="/wiki/SMAD_(protein)" title="SMAD (protein)">Smad</a> proteins that in turn act as transcription factors that regulate gene transcription.<sup id="cite_ref-Moore_2013_11-2" class="reference"><a href="#cite_note-Moore_2013-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p><p>Sonic hedgehog (SHH) are morphogens that are essential to early patterning in the developing embryo. SHH binds to the <a href="/wiki/Patched" title="Patched">Patched</a> receptor which in the absence of SHH inhibits the <a href="/wiki/Smoothened" title="Smoothened">Smoothened</a> receptor. Activated smoothened in turn causes <a href="/wiki/Gli1" class="mw-redirect" title="Gli1">Gli1</a>, <a href="/wiki/Gli2" class="mw-redirect" title="Gli2">Gli2</a>, and <a href="/wiki/Gli3" class="mw-redirect" title="Gli3">Gli3</a> to be translocated into the nucleus where they activate target genes such at <a href="/wiki/PTCH1" title="PTCH1">PTCH1</a> and <a href="/wiki/Engrailed_(gene)" title="Engrailed (gene)">Engrailed</a>.<sup id="cite_ref-Moore_2013_11-3" class="reference"><a href="#cite_note-Moore_2013-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Fruit_fly">Fruit fly</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Morphogen&action=edit&section=5" title="Edit section: Fruit fly"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><i><a href="/wiki/Drosophila_melanogaster" title="Drosophila melanogaster">Drosophila melanogaster</a></i> has an unusual developmental system, in which the first thirteen cell divisions of the embryo occur within a <a href="/wiki/Syncytium" title="Syncytium">syncytium</a> prior to cellularization. Essentially the embryo remains a single cell with over 8000 nuclei evenly spaced near the membrane until the fourteenth cell division, when independent membranes furrow between the nuclei, separating them into independent cells. As a result, in fly embryos <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factors</a> such as <a href="/wiki/Bicoid" class="mw-redirect" title="Bicoid">Bicoid</a> or Hunchback can act as morphogens because they can freely diffuse between nuclei to produce smooth gradients of concentration without relying on specialized intercellular signalling mechanisms. Although there is some evidence that <a href="/wiki/Homeobox" title="Homeobox">homeobox</a> <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factors</a> similar to these can pass directly through cell membranes,<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> this mechanism is not believed to contribute greatly to morphogenesis in cellularized<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. (November 2012)">clarification needed</span></a></i>]</sup> systems. </p><p>In most developmental systems, such as human embryos or later <i>Drosophila</i> development, syncytia occur only rarely (such as in skeletal muscle), and morphogens are generally secreted signalling proteins. These proteins bind to the extracellular domains of transmembrane <a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">receptor</a> proteins, which use an elaborate process of <a href="/wiki/Signal_transduction" title="Signal transduction">signal transduction</a> to communicate the level of morphogen to the nucleus. The nuclear targets of signal transduction pathways are usually transcription factors, whose activity is regulated in a manner that reflects the level of morphogen received at the cell surface. Thus, secreted morphogens act to generate gradients of transcription factor activity just like those that are generated in the syncitial <i>Drosophila</i> embryo. </p><p>Discrete target genes respond to different thresholds of morphogen activity. The expression of target genes is controlled by segments of DNA called '<a href="/wiki/Enhancer_(genetics)" title="Enhancer (genetics)">enhancers</a>' to which <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factors</a> bind directly. Once bound, the transcription factor then stimulates or inhibits the transcription of the gene and thus controls the level of expression of the gene product (usually a protein). 'Low-threshold' target genes require only low levels of morphogen activity to be regulated and feature enhancers that contain many high-affinity binding sites for the transcription factor. 'High-threshold' target genes have relatively fewer binding sites or low-affinity binding sites that require much greater levels of transcription factor activity to be regulated. </p><p>The general mechanism by which the morphogen model works, can explain the subdivision of tissues into patterns of distinct cell types, assuming it is possible to create and maintain a gradient. However, the morphogen model is often invoked for additional activities such as controlling the growth of the tissue or orienting the polarity of cells within it (for example, the hairs on your forearm point in one direction) which cannot be explained by model. </p> <div class="mw-heading mw-heading2"><h2 id="Eponyms">Eponyms</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Morphogen&action=edit&section=6" title="Edit section: Eponyms"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The organizing role that morphogens play during animal development was acknowledged in the 2014 naming of a new beetle genus, <i>Morphogenia</i>. The type species, <i><a href="/w/index.php?title=Morphogenia_struhli&action=edit&redlink=1" class="new" title="Morphogenia struhli (page does not exist)">Morphogenia struhli</a></i>, was named in honour of Gary Struhl, the US developmental biologist who was instrumental in demonstrating that the <i>decapentaplegic</i> and <i>wingless</i> genes encode proteins that function as morphogens during <i>Drosophila</i> development.<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p> <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=Morphogen&action=edit&section=7" 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" style="column-width: 33em;"> <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 .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFTuring,_A._M.1952" class="citation journal cs1">Turing, A. 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(2015). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4636111">"Mechanisms of retinoic acid signalling and its roles in organ and limb development"</a>. <i>Nat. Rev. Mol. Cell Biol</i>. <b>16</b> (2): 110–123. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fnrm3932">10.1038/nrm3932</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/PMC4636111">4636111</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/25560970">25560970</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nat.+Rev.+Mol.+Cell+Biol.&rft.atitle=Mechanisms+of+retinoic+acid+signalling+and+its+roles+in+organ+and+limb+development&rft.volume=16&rft.issue=2&rft.pages=110-123&rft.date=2015&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4636111%23id-name%3DPMC&rft_id=info%3Apmid%2F25560970&rft_id=info%3Adoi%2F10.1038%2Fnrm3932&rft.aulast=Cunningham&rft.aufirst=T.J.&rft.au=Duester%2C+G.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4636111&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMorphogen" class="Z3988"></span></span> </li> <li id="cite_note-13"><span class="mw-cite-backlink"><b><a href="#cite_ref-13">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDerossiJoliotChassaingProchiantz1994" class="citation journal cs1">Derossi D, Joliot AH, Chassaing G, Prochiantz A (April 1994). <a rel="nofollow" class="external text" href="http://www.jbc.org/cgi/pmidlookup?view=long&pmid=8144628">"The third helix of the Antennapedia homeodomain translocates through biological membranes"</a>. <i>J. Biol. Chem</i>. <b>269</b> (14): 10444–50. <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/10.1016%2FS0021-9258%2817%2934080-2">10.1016/S0021-9258(17)34080-2</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/8144628">8144628</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=J.+Biol.+Chem.&rft.atitle=The+third+helix+of+the+Antennapedia+homeodomain+translocates+through+biological+membranes&rft.volume=269&rft.issue=14&rft.pages=10444-50&rft.date=1994-04&rft_id=info%3Adoi%2F10.1016%2FS0021-9258%2817%2934080-2&rft_id=info%3Apmid%2F8144628&rft.aulast=Derossi&rft.aufirst=D&rft.au=Joliot%2C+AH&rft.au=Chassaing%2C+G&rft.au=Prochiantz%2C+A&rft_id=http%3A%2F%2Fwww.jbc.org%2Fcgi%2Fpmidlookup%3Fview%3Dlong%26pmid%3D8144628&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMorphogen" class="Z3988"></span><sup class="noprint Inline-Template"><span style="white-space: nowrap;">[<i><a href="/wiki/Wikipedia:Link_rot" title="Wikipedia:Link rot"><span title=" Dead link tagged August 2024">permanent dead link</span></a></i><span style="visibility:hidden; color:transparent; padding-left:2px">‍</span>]</span></sup></span> </li> <li id="cite_note-14"><span class="mw-cite-backlink"><b><a href="#cite_ref-14">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFParker_J2014" class="citation journal cs1">Parker J (23 January 2014). <a rel="nofollow" class="external text" href="https://www.pensoft.net/journals/zookeys/article/6788/abstract/morphogenia-a-new-genus-of-the-neotropical-tribe-jubini-coleoptera-staphylinidae-pselaphinae-from">"Morphogenia: a new genus of the Neotropical tribe Jubini (Coleoptera, Staphylinidae, Pselaphinae) from the Brazilian Amazon"</a>. <i>ZooKeys</i> (373): 57–66. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2014ZooK..373...57P">2014ZooK..373...57P</a>. <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/10.3897%2Fzookeys.373.6788">10.3897/zookeys.373.6788</a></span>. <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/PMC3909807">3909807</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/24493960">24493960</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=ZooKeys&rft.atitle=Morphogenia%3A+a+new+genus+of+the+Neotropical+tribe+Jubini+%28Coleoptera%2C+Staphylinidae%2C+Pselaphinae%29+from+the+Brazilian+Amazon&rft.issue=373&rft.pages=57-66&rft.date=2014-01-23&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3909807%23id-name%3DPMC&rft_id=info%3Apmid%2F24493960&rft_id=info%3Adoi%2F10.3897%2Fzookeys.373.6788&rft_id=info%3Abibcode%2F2014ZooK..373...57P&rft.au=Parker+J&rft_id=https%3A%2F%2Fwww.pensoft.net%2Fjournals%2Fzookeys%2Farticle%2F6788%2Fabstract%2Fmorphogenia-a-new-genus-of-the-neotropical-tribe-jubini-coleoptera-staphylinidae-pselaphinae-from&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMorphogen" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Morphogen&action=edit&section=8" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <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="CITEREFBriscoeLawrenceVincent2010" class="citation book cs1">Briscoe B, Lawrence P, Vincent JP, eds. (2010). <i>Generation and Interpretation of Morphogen Gradients: A Subject Collection from Cold Spring Harbor Perspectives in Biology</i>. Cold Spring Harbor, N.Y.: Cold Spring Harbor Laboratory Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-87969-881-2" title="Special:BookSources/978-0-87969-881-2"><bdi>978-0-87969-881-2</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Generation+and+Interpretation+of+Morphogen+Gradients%3A+A+Subject+Collection+from+Cold+Spring+Harbor+Perspectives+in+Biology&rft.place=Cold+Spring+Harbor%2C+N.Y.&rft.pub=Cold+Spring+Harbor+Laboratory+Press&rft.date=2010&rft.isbn=978-0-87969-881-2&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMorphogen" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRothLynch2012" class="citation journal cs1">Roth S, Lynch J (2012). <a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.cell.2012.04.007">"Does the Bicoid gradient matter?"</a>. <i>Cell</i>. <b>149</b> (3): 511–2. <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/10.1016%2Fj.cell.2012.04.007">10.1016/j.cell.2012.04.007</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/22541424">22541424</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Cell&rft.atitle=Does+the+Bicoid+gradient+matter%3F&rft.volume=149&rft.issue=3&rft.pages=511-2&rft.date=2012&rft_id=info%3Adoi%2F10.1016%2Fj.cell.2012.04.007&rft_id=info%3Apmid%2F22541424&rft.aulast=Roth&rft.aufirst=S&rft.au=Lynch%2C+J&rft_id=https%3A%2F%2Fdoi.org%2F10.1016%252Fj.cell.2012.04.007&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMorphogen" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGrove_EA_Monuki_ES2013" class="citation book cs1">Grove EA Monuki ES (2013). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=QvjurYuh-jMC&q=Morphogen&pg=PA26">"Chapter 2 - Morphogens, Patterning Centers, and their Mechanisms of Action"</a>. In Rubenstein J, Rakic P (eds.). <i>Patterning and Cell Type Specification in the Developing CNS and PNS: Comprehensive Developmental Neuroscience</i> (First ed.). San Diego: Academic Press. pp. 25–44. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0123972651" title="Special:BookSources/978-0123972651"><bdi>978-0123972651</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Chapter+2+-+Morphogens%2C+Patterning+Centers%2C+and+their+Mechanisms+of+Action&rft.btitle=Patterning+and+Cell+Type+Specification+in+the+Developing+CNS+and+PNS%3A+Comprehensive+Developmental+Neuroscience&rft.place=San+Diego&rft.pages=25-44&rft.edition=First&rft.pub=Academic+Press&rft.date=2013&rft.isbn=978-0123972651&rft.au=Grove+EA+Monuki+ES&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DQvjurYuh-jMC%26q%3DMorphogen%26pg%3DPA26&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMorphogen" class="Z3988"></span></li></ul> </div> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output 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abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Genarch" title="Template:Genarch"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Genarch" title="Template talk:Genarch"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Genarch" title="Special:EditPage/Template:Genarch"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="The_development_of_phenotype" style="font-size:114%;margin:0 4em">The <a href="/wiki/Morphogenesis" title="Morphogenesis">development</a> of <a href="/wiki/Phenotype" title="Phenotype">phenotype</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Key concepts</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/Genotype%E2%80%93phenotype_distinction" title="Genotype–phenotype distinction">Genotype–phenotype distinction</a></li> <li><a href="/wiki/Reaction_norm" title="Reaction norm">Reaction norm</a></li> <li><a href="/wiki/Gene%E2%80%93environment_interaction" title="Gene–environment interaction">Gene–environment interaction</a></li> <li><a href="/wiki/Gene%E2%80%93environment_correlation" title="Gene–environment correlation">Gene–environment correlation</a></li> <li><a href="/wiki/Operon" title="Operon">Operon</a></li> <li><a href="/wiki/Heritability" title="Heritability">Heritability</a></li> <li><a href="/wiki/Quantitative_genetics" title="Quantitative genetics">Quantitative genetics</a></li> <li><a href="/wiki/Heterochrony" title="Heterochrony">Heterochrony</a> <ul><li><a href="/wiki/Neoteny" title="Neoteny">Neoteny</a></li></ul></li> <li><a href="/wiki/Heterotopy" title="Heterotopy">Heterotopy</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Genetic_architecture" title="Genetic architecture">Genetic architecture</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/Canalisation_(genetics)" title="Canalisation (genetics)">Canalisation</a></li> <li><a href="/wiki/Genetic_assimilation" title="Genetic assimilation">Genetic assimilation</a></li> <li><a href="/wiki/Dominance_(genetics)" title="Dominance (genetics)">Dominance</a></li> <li><a href="/wiki/Epistasis" title="Epistasis">Epistasis</a></li> <li><a href="/wiki/Fitness_landscape" title="Fitness landscape">Fitness landscape</a>/<a href="/wiki/Evolutionary_landscape" title="Evolutionary landscape">evolutionary landscape</a></li> <li><a href="/wiki/Pleiotropy" title="Pleiotropy">Pleiotropy</a></li> <li><a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">Plasticity</a></li> <li><a href="/wiki/Polygenic_inheritance" class="mw-redirect" title="Polygenic inheritance">Polygenic inheritance</a></li> <li><a href="/wiki/Transgressive_segregation" title="Transgressive segregation">Transgressive segregation</a></li> <li><a href="/wiki/Sequence_space_(evolution)" title="Sequence space (evolution)">Sequence space</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Non-genetic influences</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/Epigenetics" title="Epigenetics">Epigenetics</a></li> <li><a href="/wiki/Maternal_effect" title="Maternal effect">Maternal effect</a></li> <li><a href="/wiki/Genomic_imprinting" title="Genomic imprinting">Genomic imprinting</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li> <li><a href="/wiki/Polyphenism" title="Polyphenism">Polyphenism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Developmental architecture</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/Developmental_biology" title="Developmental biology">Developmental biology</a></li> <li><a href="/wiki/Morphogenesis" title="Morphogenesis">Morphogenesis</a> <ul><li><a href="/wiki/Eyespot_(mimicry)" title="Eyespot (mimicry)">Eyespot</a></li> <li><a href="/wiki/Pattern_formation" title="Pattern formation">Pattern formation</a></li></ul></li> <li><a href="/wiki/Segmentation_(biology)" title="Segmentation (biology)">Segmentation</a></li> <li><a href="/wiki/Metamerism_(biology)" title="Metamerism (biology)">Metamerism</a></li> <li><a href="/wiki/Modularity_(biology)" title="Modularity (biology)">Modularity</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Evolution of genetic systems</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/Evolvability" title="Evolvability">Evolvability</a></li> <li><a href="/wiki/Robustness_(evolution)" title="Robustness (evolution)">Robustness</a></li> <li><a href="/wiki/Neutral_network_(evolution)" title="Neutral network (evolution)">Neutral networks</a></li> <li><a href="/wiki/Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">Evolution of sexual reproduction</a></li> <li><a href="/wiki/Evolutionary_tinkering" title="Evolutionary tinkering">Tinkering</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Control of development</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%">Systems</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/Regulation_of_gene_expression" title="Regulation of gene expression">Regulation of gene expression</a></li> <li><a href="/wiki/Gene_regulatory_network" title="Gene regulatory network">Gene regulatory network</a></li> <li><a href="/wiki/Evo-devo_gene_toolkit" title="Evo-devo gene toolkit">Evo-devo gene toolkit</a></li> <li><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a></li> <li><a href="/wiki/Homeobox" title="Homeobox">Homeobox</a></li> <li><a href="/wiki/Hedgehog_signaling_pathway" title="Hedgehog signaling pathway">Hedgehog signaling pathway</a></li> <li><a href="/wiki/Notch_signaling_pathway" title="Notch signaling pathway">Notch signaling pathway</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Elements</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/Homeotic_gene" title="Homeotic gene">Homeotic gene</a></li> <li><a href="/wiki/Hox_gene" title="Hox gene">Hox gene</a></li> <li><a href="/wiki/Pax_genes" title="Pax genes">Pax genes</a> <ul><li><a href="/wiki/PAX6" title="PAX6">eyeless gene</a></li></ul></li> <li><a href="/wiki/DLX_gene_family" title="DLX gene family">Distal-less</a></li> <li><a href="/wiki/Engrailed_(gene)" title="Engrailed (gene)">Engrailed</a></li> <li><a href="/wiki/Cis-regulatory_element" title="Cis-regulatory element">cis-regulatory element</a></li> <li><a href="/wiki/Ligand_(biochemistry)" title="Ligand (biochemistry)">Ligand</a></li> <li><a class="mw-selflink selflink">Morphogen</a></li> <li><a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">Cell surface receptor</a></li> <li><a href="/wiki/Transcription_factor" title="Transcription factor">Transcription factor</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Influential figures</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/C._H._Waddington" title="C. H. Waddington">C. H. Waddington</a></li> <li><a href="/wiki/Richard_Lewontin" title="Richard Lewontin">Richard Lewontin</a></li> <li><a href="/wiki/Fran%C3%A7ois_Jacob" title="François Jacob">François Jacob</a> + <a href="/wiki/Jacques_Monod" title="Jacques Monod">Jacques Monod</a> <ul><li><a href="/wiki/Lac_operon" title="Lac operon">Lac operon</a></li></ul></li> <li><a href="/wiki/Eric_F._Wieschaus" title="Eric F. Wieschaus">Eric F. Wieschaus</a></li> <li><a href="/wiki/Christiane_N%C3%BCsslein-Volhard" title="Christiane Nüsslein-Volhard">Christiane Nüsslein-Volhard</a></li> <li><a href="/wiki/William_McGinnis" title="William McGinnis">William McGinnis</a></li> <li><a href="/wiki/Michael_Levine_(biologist)" title="Michael Levine (biologist)">Mike Levine</a></li> <li><a href="/wiki/Sean_B._Carroll" title="Sean B. Carroll">Sean B. Carroll</a> <ul><li><i><a href="/wiki/Endless_Forms_Most_Beautiful_(book)" title="Endless Forms Most Beautiful (book)">Endless Forms Most Beautiful</a></i></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Debates</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/Nature_versus_nurture" title="Nature versus nurture">Nature versus nurture</a></li> <li><a href="/wiki/Morphogenetic_field" title="Morphogenetic field">Morphogenetic field</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div><a href="/wiki/Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index of evolutionary biology articles</a></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: absolute;"></noscript> <div class="printfooter" data-nosnippet="">Retrieved from "<a dir="ltr" href="https://en.wikipedia.org/w/index.php?title=Morphogen&oldid=1240853646">https://en.wikipedia.org/w/index.php?title=Morphogen&oldid=1240853646</a>"</div></div> <div id="catlinks" class="catlinks" data-mw="interface"><div id="mw-normal-catlinks" class="mw-normal-catlinks"><a href="/wiki/Help:Category" title="Help:Category">Category</a>: <ul><li><a href="/wiki/Category:Morphogens" title="Category:Morphogens">Morphogens</a></li></ul></div><div id="mw-hidden-catlinks" class="mw-hidden-catlinks mw-hidden-cats-hidden">Hidden categories: <ul><li><a href="/wiki/Category:All_articles_with_dead_external_links" title="Category:All articles with dead external links">All articles with dead external links</a></li><li><a href="/wiki/Category:Articles_with_dead_external_links_from_August_2024" title="Category:Articles with dead external links from August 2024">Articles with dead external links from August 2024</a></li><li><a href="/wiki/Category:Articles_with_permanently_dead_external_links" title="Category:Articles with permanently dead external links">Articles with permanently dead external links</a></li><li><a href="/wiki/Category:Articles_with_short_description" title="Category:Articles with short description">Articles with short description</a></li><li><a href="/wiki/Category:Short_description_matches_Wikidata" title="Category:Short description matches Wikidata">Short description matches Wikidata</a></li><li><a href="/wiki/Category:Articles_needing_additional_references_from_May_2008" title="Category:Articles needing additional references from May 2008">Articles needing additional references from May 2008</a></li><li><a href="/wiki/Category:All_articles_needing_additional_references" title="Category:All articles needing additional references">All articles needing additional references</a></li><li><a href="/wiki/Category:Articles_needing_expert_attention_from_August_2015" title="Category:Articles needing expert attention from August 2015">Articles needing expert attention from August 2015</a></li><li><a href="/wiki/Category:All_articles_needing_expert_attention" title="Category:All articles needing expert attention">All articles needing expert attention</a></li><li><a href="/wiki/Category:Biology_articles_needing_expert_attention" title="Category:Biology articles needing expert attention">Biology articles needing expert attention</a></li><li><a href="/wiki/Category:Articles_with_multiple_maintenance_issues" title="Category:Articles with multiple maintenance issues">Articles with multiple maintenance issues</a></li><li><a href="/wiki/Category:Wikipedia_articles_needing_clarification_from_November_2012" title="Category:Wikipedia articles needing clarification from November 2012">Wikipedia articles needing clarification from November 2012</a></li></ul></div></div> </div> </main> </div> <div class="mw-footer-container"> <footer id="footer" class="mw-footer" > <ul id="footer-info"> <li id="footer-info-lastmod"> This page was last edited on 17 August 2024, at 20:52<span class="anonymous-show"> (UTC)</span>.</li> <li id="footer-info-copyright">Text is available under the <a href="/wiki/Wikipedia:Text_of_the_Creative_Commons_Attribution-ShareAlike_4.0_International_License" title="Wikipedia:Text of the Creative Commons Attribution-ShareAlike 4.0 International License">Creative Commons Attribution-ShareAlike 4.0 License</a>; additional terms may apply. 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