CINXE.COM

<!DOCTYPE html> <html class="client-nojs skin-theme-clientpref-day mf-expand-sections-clientpref-0 mf-font-size-clientpref-small mw-mf-amc-clientpref-0" lang="en" dir="ltr"> <head> <meta charset="UTF-8"> <title>Morphogen - Wikipedia</title> <script>(function(){var className="client-js skin-theme-clientpref-day mf-expand-sections-clientpref-0 mf-font-size-clientpref-small mw-mf-amc-clientpref-0";var cookie=document.cookie.match(/(?:^|; )enwikimwclientpreferences=([^;]+)/);if(cookie){cookie[1].split('%2C').forEach(function(pref){className=className.replace(new RegExp('(^| )'+pref.replace(/-clientpref-\w+$|[^\w-]+/g,'')+'-clientpref-\\w+( |$)'),'$1'+pref+'$2');});}document.documentElement.className=className;}());RLCONF={"wgBreakFrames":false,"wgSeparatorTransformTable":["",""],"wgDigitTransformTable":["",""],"wgDefaultDateFormat":"dmy","wgMonthNames":["","January","February","March","April","May","June","July","August","September","October","November","December"],"wgRequestId":"b2525925-6bd4-43ef-ab8a-e7df3801838d","wgCanonicalNamespace":"","wgCanonicalSpecialPageName":false,"wgNamespaceNumber":0,"wgPageName":"Morphogen","wgTitle":"Morphogen","wgCurRevisionId":1260200925,"wgRevisionId":1260200925,"wgArticleId":1687906,"wgIsArticle" :true,"wgIsRedirect":false,"wgAction":"view","wgUserName":null,"wgUserGroups":["*"],"wgPageViewLanguage":"en","wgPageContentLanguage":"en","wgPageContentModel":"wikitext","wgRelevantPageName":"Morphogen","wgRelevantArticleId":1687906,"wgIsProbablyEditable":true,"wgRelevantPageIsProbablyEditable":true,"wgRestrictionEdit":[],"wgRestrictionMove":[],"wgNoticeProject":"wikipedia","wgCiteReferencePreviewsActive":false,"wgFlaggedRevsParams":{"tags":{"status":{"levels":1}}},"wgMediaViewerOnClick":true,"wgMediaViewerEnabledByDefault":true,"wgPopupsFlags":0,"wgVisualEditor":{"pageLanguageCode":"en","pageLanguageDir":"ltr","pageVariantFallbacks":"en"},"wgMFMode":"stable","wgMFAmc":false,"wgMFAmcOutreachActive":false,"wgMFAmcOutreachUserEligible":false,"wgMFLazyLoadImages":true,"wgMFEditNoticesFeatureConflict":false,"wgMFDisplayWikibaseDescriptions":{"search":true,"watchlist":true,"tagline":false,"nearby":true},"wgMFIsSupportedEditRequest":true,"wgMFScriptPath":"", "wgWMESchemaEditAttemptStepOversample":false,"wgWMEPageLength":20000,"wgRelatedArticlesCompat":[],"wgEditSubmitButtonLabelPublish":true,"wgSectionTranslationMissingLanguages":[{"lang":"ace","autonym":"Acèh","dir":"ltr"},{"lang":"ady","autonym":"адыгабзэ","dir":"ltr"},{"lang":"alt","autonym":"алтай тил","dir":"ltr"},{"lang":"am","autonym":"አማርኛ","dir":"ltr"},{"lang":"ami","autonym":"Pangcah","dir":"ltr"},{"lang":"an","autonym":"aragonés","dir":"ltr"},{"lang":"ang","autonym":"Ænglisc","dir":"ltr"},{"lang":"ann","autonym":"Obolo","dir":"ltr"},{"lang":"anp","autonym":"अंगिका","dir":"ltr"},{"lang":"ar","autonym":"العربية","dir":"rtl"},{"lang":"ary","autonym":"الدارجة","dir":"rtl"},{"lang":"arz","autonym":"مصرى","dir":"rtl"},{"lang":"as","autonym":"অসমীয়া","dir":"ltr"},{"lang":"ast","autonym":"asturianu","dir":"ltr"},{"lang":"av","autonym":"авар","dir":"ltr"},{"lang":"avk","autonym":"Kotava","dir":"ltr"},{"lang": "awa","autonym":"अवधी","dir":"ltr"},{"lang":"ay","autonym":"Aymar aru","dir":"ltr"},{"lang":"az","autonym":"azərbaycanca","dir":"ltr"},{"lang":"azb","autonym":"تۆرکجه","dir":"rtl"},{"lang":"ba","autonym":"башҡортса","dir":"ltr"},{"lang":"ban","autonym":"Basa Bali","dir":"ltr"},{"lang":"bar","autonym":"Boarisch","dir":"ltr"},{"lang":"bbc","autonym":"Batak Toba","dir":"ltr"},{"lang":"bcl","autonym":"Bikol Central","dir":"ltr"},{"lang":"bdr","autonym":"Bajau Sama","dir":"ltr"},{"lang":"be","autonym":"беларуская","dir":"ltr"},{"lang":"bew","autonym":"Betawi","dir":"ltr"},{"lang":"bg","autonym":"български","dir":"ltr"},{"lang":"bho","autonym":"भोजपुरी","dir":"ltr"},{"lang":"bi","autonym":"Bislama","dir":"ltr"},{"lang":"bjn","autonym":"Banjar","dir":"ltr"},{"lang":"blk","autonym":"ပအိုဝ်ႏဘာႏသာႏ","dir":"ltr"},{"lang":"bm","autonym":"bamanankan","dir":"ltr"},{"lang":"bn","autonym":"বাংলা","dir":"ltr"}, {"lang":"bo","autonym":"བོད་ཡིག","dir":"ltr"},{"lang":"bpy","autonym":"বিষ্ণুপ্রিয়া মণিপুরী","dir":"ltr"},{"lang":"br","autonym":"brezhoneg","dir":"ltr"},{"lang":"bs","autonym":"bosanski","dir":"ltr"},{"lang":"btm","autonym":"Batak Mandailing","dir":"ltr"},{"lang":"bug","autonym":"Basa Ugi","dir":"ltr"},{"lang":"cdo","autonym":"閩東語 / Mìng-dĕ̤ng-ngṳ̄","dir":"ltr"},{"lang":"ce","autonym":"нохчийн","dir":"ltr"},{"lang":"ceb","autonym":"Cebuano","dir":"ltr"},{"lang":"ch","autonym":"Chamoru","dir":"ltr"},{"lang":"chr","autonym":"ᏣᎳᎩ","dir":"ltr"},{"lang":"ckb","autonym":"کوردی","dir":"rtl"},{"lang":"co","autonym":"corsu","dir":"ltr"},{"lang":"cr","autonym":"Nēhiyawēwin / ᓀᐦᐃᔭᐍᐏᐣ","dir":"ltr"},{"lang":"crh","autonym":"qırımtatarca","dir":"ltr"},{"lang":"cs","autonym":"čeština","dir":"ltr"},{"lang":"cu","autonym":"словѣньскъ / ⰔⰎⰑⰂⰡⰐⰠⰔⰍⰟ","dir":"ltr"},{"lang": "cy","autonym":"Cymraeg","dir":"ltr"},{"lang":"da","autonym":"dansk","dir":"ltr"},{"lang":"dag","autonym":"dagbanli","dir":"ltr"},{"lang":"dga","autonym":"Dagaare","dir":"ltr"},{"lang":"din","autonym":"Thuɔŋjäŋ","dir":"ltr"},{"lang":"diq","autonym":"Zazaki","dir":"ltr"},{"lang":"dsb","autonym":"dolnoserbski","dir":"ltr"},{"lang":"dtp","autonym":"Kadazandusun","dir":"ltr"},{"lang":"dv","autonym":"ދިވެހިބަސް","dir":"rtl"},{"lang":"dz","autonym":"ཇོང་ཁ","dir":"ltr"},{"lang":"ee","autonym":"eʋegbe","dir":"ltr"},{"lang":"el","autonym":"Ελληνικά","dir":"ltr"},{"lang":"eml","autonym":"emiliàn e rumagnòl","dir":"ltr"},{"lang":"eo","autonym":"Esperanto","dir":"ltr"},{"lang":"et","autonym":"eesti","dir":"ltr"},{"lang":"eu","autonym":"euskara","dir":"ltr"},{"lang":"fa","autonym":"فارسی","dir":"rtl"},{"lang":"fat","autonym":"mfantse","dir":"ltr"},{"lang":"ff","autonym":"Fulfulde","dir":"ltr"},{"lang":"fj","autonym":"Na Vosa Vakaviti","dir":"ltr"},{"lang": "fo","autonym":"føroyskt","dir":"ltr"},{"lang":"fon","autonym":"fɔ̀ngbè","dir":"ltr"},{"lang":"frp","autonym":"arpetan","dir":"ltr"},{"lang":"frr","autonym":"Nordfriisk","dir":"ltr"},{"lang":"fur","autonym":"furlan","dir":"ltr"},{"lang":"fy","autonym":"Frysk","dir":"ltr"},{"lang":"gag","autonym":"Gagauz","dir":"ltr"},{"lang":"gan","autonym":"贛語","dir":"ltr"},{"lang":"gcr","autonym":"kriyòl gwiyannen","dir":"ltr"},{"lang":"gl","autonym":"galego","dir":"ltr"},{"lang":"glk","autonym":"گیلکی","dir":"rtl"},{"lang":"gn","autonym":"Avañe'ẽ","dir":"ltr"},{"lang":"gom","autonym":"गोंयची कोंकणी / Gõychi Konknni","dir":"ltr"},{"lang":"gor","autonym":"Bahasa Hulontalo","dir":"ltr"},{"lang":"gpe","autonym":"Ghanaian Pidgin","dir":"ltr"},{"lang":"gu","autonym":"ગુજરાતી","dir":"ltr"},{"lang":"guc","autonym":"wayuunaiki","dir":"ltr"},{"lang":"gur","autonym":"farefare","dir":"ltr"},{"lang":"guw","autonym":"gungbe","dir":"ltr"},{"lang":"gv", "autonym":"Gaelg","dir":"ltr"},{"lang":"ha","autonym":"Hausa","dir":"ltr"},{"lang":"hak","autonym":"客家語 / Hak-kâ-ngî","dir":"ltr"},{"lang":"haw","autonym":"Hawaiʻi","dir":"ltr"},{"lang":"hi","autonym":"हिन्दी","dir":"ltr"},{"lang":"hif","autonym":"Fiji Hindi","dir":"ltr"},{"lang":"hr","autonym":"hrvatski","dir":"ltr"},{"lang":"hsb","autonym":"hornjoserbsce","dir":"ltr"},{"lang":"ht","autonym":"Kreyòl ayisyen","dir":"ltr"},{"lang":"hu","autonym":"magyar","dir":"ltr"},{"lang":"hy","autonym":"հայերեն","dir":"ltr"},{"lang":"hyw","autonym":"Արեւմտահայերէն","dir":"ltr"},{"lang":"ia","autonym":"interlingua","dir":"ltr"},{"lang":"iba","autonym":"Jaku Iban","dir":"ltr"},{"lang":"ie","autonym":"Interlingue","dir":"ltr"},{"lang":"ig","autonym":"Igbo","dir":"ltr"},{"lang":"igl","autonym":"Igala","dir":"ltr"},{"lang":"ilo","autonym":"Ilokano","dir":"ltr"},{"lang":"io","autonym":"Ido","dir":"ltr"},{"lang":"is","autonym":"íslenska","dir":"ltr"},{"lang":"iu" ,"autonym":"ᐃᓄᒃᑎᑐᑦ / inuktitut","dir":"ltr"},{"lang":"jam","autonym":"Patois","dir":"ltr"},{"lang":"jv","autonym":"Jawa","dir":"ltr"},{"lang":"ka","autonym":"ქართული","dir":"ltr"},{"lang":"kaa","autonym":"Qaraqalpaqsha","dir":"ltr"},{"lang":"kab","autonym":"Taqbaylit","dir":"ltr"},{"lang":"kbd","autonym":"адыгэбзэ","dir":"ltr"},{"lang":"kbp","autonym":"Kabɩyɛ","dir":"ltr"},{"lang":"kcg","autonym":"Tyap","dir":"ltr"},{"lang":"kg","autonym":"Kongo","dir":"ltr"},{"lang":"kge","autonym":"Kumoring","dir":"ltr"},{"lang":"ki","autonym":"Gĩkũyũ","dir":"ltr"},{"lang":"kk","autonym":"қазақша","dir":"ltr"},{"lang":"kl","autonym":"kalaallisut","dir":"ltr"},{"lang":"km","autonym":"ភាសាខ្មែរ","dir":"ltr"},{"lang":"kn","autonym":"ಕನ್ನಡ","dir":"ltr"},{"lang":"ko","autonym":"한국어","dir":"ltr"},{"lang":"koi","autonym":"перем коми","dir":"ltr"},{"lang":"krc","autonym":"къарачай-малкъар","dir":"ltr"},{ "lang":"ks","autonym":"कॉशुर / کٲشُر","dir":"rtl"},{"lang":"ku","autonym":"kurdî","dir":"ltr"},{"lang":"kus","autonym":"Kʋsaal","dir":"ltr"},{"lang":"kv","autonym":"коми","dir":"ltr"},{"lang":"kw","autonym":"kernowek","dir":"ltr"},{"lang":"ky","autonym":"кыргызча","dir":"ltr"},{"lang":"lad","autonym":"Ladino","dir":"ltr"},{"lang":"lb","autonym":"Lëtzebuergesch","dir":"ltr"},{"lang":"lez","autonym":"лезги","dir":"ltr"},{"lang":"lg","autonym":"Luganda","dir":"ltr"},{"lang":"li","autonym":"Limburgs","dir":"ltr"},{"lang":"lij","autonym":"Ligure","dir":"ltr"},{"lang":"lld","autonym":"Ladin","dir":"ltr"},{"lang":"lmo","autonym":"lombard","dir":"ltr"},{"lang":"ln","autonym":"lingála","dir":"ltr"},{"lang":"lo","autonym":"ລາວ","dir":"ltr"},{"lang":"lt","autonym":"lietuvių","dir":"ltr"},{"lang":"ltg","autonym":"latgaļu","dir":"ltr"},{"lang":"lv","autonym":"latviešu","dir":"ltr"},{"lang":"mad","autonym":"Madhurâ","dir":"ltr"},{"lang":"mai","autonym": "मैथिली","dir":"ltr"},{"lang":"map-bms","autonym":"Basa Banyumasan","dir":"ltr"},{"lang":"mdf","autonym":"мокшень","dir":"ltr"},{"lang":"mg","autonym":"Malagasy","dir":"ltr"},{"lang":"mhr","autonym":"олык марий","dir":"ltr"},{"lang":"mi","autonym":"Māori","dir":"ltr"},{"lang":"min","autonym":"Minangkabau","dir":"ltr"},{"lang":"mk","autonym":"македонски","dir":"ltr"},{"lang":"ml","autonym":"മലയാളം","dir":"ltr"},{"lang":"mn","autonym":"монгол","dir":"ltr"},{"lang":"mni","autonym":"ꯃꯤꯇꯩ ꯂꯣꯟ","dir":"ltr"},{"lang":"mnw","autonym":"ဘာသာမန်","dir":"ltr"},{"lang":"mos","autonym":"moore","dir":"ltr"},{"lang":"mr","autonym":"मराठी","dir":"ltr"},{"lang":"mrj","autonym":"кырык мары","dir":"ltr"},{"lang":"ms","autonym":"Bahasa Melayu","dir":"ltr"},{"lang":"mt","autonym":"Malti","dir":"ltr"},{"lang":"mwl","autonym":"Mirandés","dir":"ltr"},{"lang":"my","autonym":"မြန်မာဘာသာ", "dir":"ltr"},{"lang":"myv","autonym":"эрзянь","dir":"ltr"},{"lang":"mzn","autonym":"مازِرونی","dir":"rtl"},{"lang":"nah","autonym":"Nāhuatl","dir":"ltr"},{"lang":"nan","autonym":"閩南語 / Bân-lâm-gú","dir":"ltr"},{"lang":"nap","autonym":"Napulitano","dir":"ltr"},{"lang":"nb","autonym":"norsk bokmål","dir":"ltr"},{"lang":"nds","autonym":"Plattdüütsch","dir":"ltr"},{"lang":"nds-nl","autonym":"Nedersaksies","dir":"ltr"},{"lang":"ne","autonym":"नेपाली","dir":"ltr"},{"lang":"new","autonym":"नेपाल भाषा","dir":"ltr"},{"lang":"nia","autonym":"Li Niha","dir":"ltr"},{"lang":"nl","autonym":"Nederlands","dir":"ltr"},{"lang":"nn","autonym":"norsk nynorsk","dir":"ltr"},{"lang":"nqo","autonym":"ߒߞߏ","dir":"rtl"},{"lang":"nr","autonym":"isiNdebele seSewula","dir":"ltr"},{"lang":"nso","autonym":"Sesotho sa Leboa","dir":"ltr"},{"lang":"ny","autonym":"Chi-Chewa","dir":"ltr"},{"lang":"oc","autonym":"occitan","dir":"ltr"},{"lang":"om","autonym": "Oromoo","dir":"ltr"},{"lang":"or","autonym":"ଓଡ଼ିଆ","dir":"ltr"},{"lang":"os","autonym":"ирон","dir":"ltr"},{"lang":"pa","autonym":"ਪੰਜਾਬੀ","dir":"ltr"},{"lang":"pag","autonym":"Pangasinan","dir":"ltr"},{"lang":"pam","autonym":"Kapampangan","dir":"ltr"},{"lang":"pap","autonym":"Papiamentu","dir":"ltr"},{"lang":"pcd","autonym":"Picard","dir":"ltr"},{"lang":"pcm","autonym":"Naijá","dir":"ltr"},{"lang":"pdc","autonym":"Deitsch","dir":"ltr"},{"lang":"pms","autonym":"Piemontèis","dir":"ltr"},{"lang":"pnb","autonym":"پنجابی","dir":"rtl"},{"lang":"ps","autonym":"پښتو","dir":"rtl"},{"lang":"pt","autonym":"português","dir":"ltr"},{"lang":"pwn","autonym":"pinayuanan","dir":"ltr"},{"lang":"qu","autonym":"Runa Simi","dir":"ltr"},{"lang":"rm","autonym":"rumantsch","dir":"ltr"},{"lang":"rn","autonym":"ikirundi","dir":"ltr"},{"lang":"ro","autonym":"română","dir":"ltr"},{"lang":"rsk","autonym":"руски","dir":"ltr"},{"lang":"rue","autonym": "русиньскый","dir":"ltr"},{"lang":"rup","autonym":"armãneashti","dir":"ltr"},{"lang":"rw","autonym":"Ikinyarwanda","dir":"ltr"},{"lang":"sa","autonym":"संस्कृतम्","dir":"ltr"},{"lang":"sah","autonym":"саха тыла","dir":"ltr"},{"lang":"sat","autonym":"ᱥᱟᱱᱛᱟᱲᱤ","dir":"ltr"},{"lang":"sc","autonym":"sardu","dir":"ltr"},{"lang":"scn","autonym":"sicilianu","dir":"ltr"},{"lang":"sco","autonym":"Scots","dir":"ltr"},{"lang":"sd","autonym":"سنڌي","dir":"rtl"},{"lang":"se","autonym":"davvisámegiella","dir":"ltr"},{"lang":"sg","autonym":"Sängö","dir":"ltr"},{"lang":"sgs","autonym":"žemaitėška","dir":"ltr"},{"lang":"sh","autonym":"srpskohrvatski / српскохрватски","dir":"ltr"},{"lang":"shi","autonym":"Taclḥit","dir":"ltr"},{"lang":"shn","autonym":"ၽႃႇသႃႇတႆး ","dir":"ltr"},{"lang":"si","autonym":"සිංහල","dir":"ltr"},{"lang":"sk","autonym":"slovenčina","dir":"ltr"},{"lang":"skr","autonym": "سرائیکی","dir":"rtl"},{"lang":"sl","autonym":"slovenščina","dir":"ltr"},{"lang":"sm","autonym":"Gagana Samoa","dir":"ltr"},{"lang":"smn","autonym":"anarâškielâ","dir":"ltr"},{"lang":"sn","autonym":"chiShona","dir":"ltr"},{"lang":"so","autonym":"Soomaaliga","dir":"ltr"},{"lang":"sq","autonym":"shqip","dir":"ltr"},{"lang":"srn","autonym":"Sranantongo","dir":"ltr"},{"lang":"ss","autonym":"SiSwati","dir":"ltr"},{"lang":"st","autonym":"Sesotho","dir":"ltr"},{"lang":"stq","autonym":"Seeltersk","dir":"ltr"},{"lang":"su","autonym":"Sunda","dir":"ltr"},{"lang":"sw","autonym":"Kiswahili","dir":"ltr"},{"lang":"szl","autonym":"ślůnski","dir":"ltr"},{"lang":"ta","autonym":"தமிழ்","dir":"ltr"},{"lang":"tay","autonym":"Tayal","dir":"ltr"},{"lang":"tcy","autonym":"ತುಳು","dir":"ltr"},{"lang":"tdd","autonym":"ᥖᥭᥰ ᥖᥬᥲ ᥑᥨᥒᥰ","dir":"ltr"},{"lang":"te","autonym":"తెలుగు","dir":"ltr"},{"lang":"tet","autonym":"tetun","dir":"ltr"},{"lang":"tg", "autonym":"тоҷикӣ","dir":"ltr"},{"lang":"th","autonym":"ไทย","dir":"ltr"},{"lang":"ti","autonym":"ትግርኛ","dir":"ltr"},{"lang":"tk","autonym":"Türkmençe","dir":"ltr"},{"lang":"tl","autonym":"Tagalog","dir":"ltr"},{"lang":"tly","autonym":"tolışi","dir":"ltr"},{"lang":"tn","autonym":"Setswana","dir":"ltr"},{"lang":"to","autonym":"lea faka-Tonga","dir":"ltr"},{"lang":"tpi","autonym":"Tok Pisin","dir":"ltr"},{"lang":"tr","autonym":"Türkçe","dir":"ltr"},{"lang":"trv","autonym":"Seediq","dir":"ltr"},{"lang":"ts","autonym":"Xitsonga","dir":"ltr"},{"lang":"tt","autonym":"татарча / tatarça","dir":"ltr"},{"lang":"tum","autonym":"chiTumbuka","dir":"ltr"},{"lang":"tw","autonym":"Twi","dir":"ltr"},{"lang":"ty","autonym":"reo tahiti","dir":"ltr"},{"lang":"tyv","autonym":"тыва дыл","dir":"ltr"},{"lang":"udm","autonym":"удмурт","dir":"ltr"},{"lang":"ur","autonym":"اردو","dir":"rtl"},{"lang":"uz","autonym":"oʻzbekcha / ўзбекча","dir":"ltr"},{ "lang":"ve","autonym":"Tshivenda","dir":"ltr"},{"lang":"vec","autonym":"vèneto","dir":"ltr"},{"lang":"vep","autonym":"vepsän kel’","dir":"ltr"},{"lang":"vi","autonym":"Tiếng Việt","dir":"ltr"},{"lang":"vls","autonym":"West-Vlams","dir":"ltr"},{"lang":"vo","autonym":"Volapük","dir":"ltr"},{"lang":"vro","autonym":"võro","dir":"ltr"},{"lang":"wa","autonym":"walon","dir":"ltr"},{"lang":"war","autonym":"Winaray","dir":"ltr"},{"lang":"wo","autonym":"Wolof","dir":"ltr"},{"lang":"wuu","autonym":"吴语","dir":"ltr"},{"lang":"xal","autonym":"хальмг","dir":"ltr"},{"lang":"xh","autonym":"isiXhosa","dir":"ltr"},{"lang":"xmf","autonym":"მარგალური","dir":"ltr"},{"lang":"yi","autonym":"ייִדיש","dir":"rtl"},{"lang":"yo","autonym":"Yorùbá","dir":"ltr"},{"lang":"yue","autonym":"粵語","dir":"ltr"},{"lang":"za","autonym":"Vahcuengh","dir":"ltr"},{"lang":"zgh","autonym":"ⵜⴰⵎⴰⵣⵉⵖⵜ ⵜⴰⵏⴰⵡⴰⵢⵜ","dir":"ltr"},{"lang":"zh","autonym": "中文","dir":"ltr"},{"lang":"zu","autonym":"isiZulu","dir":"ltr"}],"wgSectionTranslationTargetLanguages":["ace","ady","alt","am","ami","an","ang","ann","anp","ar","ary","arz","as","ast","av","avk","awa","ay","az","azb","ba","ban","bar","bbc","bcl","bdr","be","bew","bg","bho","bi","bjn","blk","bm","bn","bo","bpy","br","bs","btm","bug","ca","cdo","ce","ceb","ch","chr","ckb","co","cr","crh","cs","cu","cy","da","dag","de","dga","din","diq","dsb","dtp","dv","dz","ee","el","eml","eo","es","et","eu","fa","fat","ff","fi","fj","fo","fon","fr","frp","frr","fur","fy","gag","gan","gcr","gl","glk","gn","gom","gor","gpe","gu","guc","gur","guw","gv","ha","hak","haw","he","hi","hif","hr","hsb","ht","hu","hy","hyw","ia","iba","ie","ig","igl","ilo","io","is","it","iu","ja","jam","jv","ka","kaa","kab","kbd","kbp","kcg","kg","kge","ki","kk","kl","km","kn","ko","koi","krc","ks","ku","kus","kv","kw","ky","lad","lb","lez","lg","li","lij","lld","lmo","ln","lo","lt","ltg","lv","mad","mai","map-bms","mdf", "mg","mhr","mi","min","mk","ml","mn","mni","mnw","mos","mr","mrj","ms","mt","mwl","my","myv","mzn","nah","nan","nap","nb","nds","nds-nl","ne","new","nia","nl","nn","nqo","nr","nso","ny","oc","om","or","os","pa","pag","pam","pap","pcd","pcm","pdc","pl","pms","pnb","ps","pt","pwn","qu","rm","rn","ro","rsk","rue","rup","rw","sa","sah","sat","sc","scn","sco","sd","se","sg","sgs","sh","shi","shn","si","sk","skr","sl","sm","smn","sn","so","sq","sr","srn","ss","st","stq","su","sv","sw","szl","ta","tay","tcy","tdd","te","tet","tg","th","ti","tk","tl","tly","tn","to","tpi","tr","trv","ts","tt","tum","tw","ty","tyv","udm","ur","uz","ve","vec","vep","vi","vls","vo","vro","wa","war","wo","wuu","xal","xh","xmf","yi","yo","yue","za","zgh","zh","zu"],"isLanguageSearcherCXEntrypointEnabled":true,"mintEntrypointLanguages":["ace","ast","azb","bcl","bjn","bh","crh","ff","fon","ig","is","ki","ks","lmo","min","sat","ss","tn","vec"],"wgWikibaseItemId":"Q417084","wgCheckUserClientHintsHeadersJsApi":["brands" ,"architecture","bitness","fullVersionList","mobile","model","platform","platformVersion"],"GEHomepageSuggestedEditsEnableTopics":true,"wgGETopicsMatchModeEnabled":false,"wgGEStructuredTaskRejectionReasonTextInputEnabled":false,"wgGELevelingUpEnabledForUser":false,"wgMinervaPermissions":{"watchable":true,"watch":false},"wgMinervaFeatures":{"beta":false,"donate":true,"mobileOptionsLink":true,"categories":false,"pageIssues":true,"talkAtTop":true,"historyInPageActions":false,"overflowSubmenu":false,"tabsOnSpecials":true,"personalMenu":false,"mainMenuExpanded":false,"echo":true,"nightMode":true},"wgMinervaDownloadNamespaces":[0]};RLSTATE={"ext.globalCssJs.user.styles":"ready","site.styles":"ready","user.styles":"ready","ext.globalCssJs.user":"ready","user":"ready","user.options":"loading","ext.cite.styles":"ready","skins.minerva.styles":"ready","skins.minerva.content.styles.images":"ready","mediawiki.hlist":"ready","skins.minerva.codex.styles":"ready","skins.minerva.icons":"ready", "skins.minerva.amc.styles":"ready","ext.wikimediamessages.styles":"ready","mobile.init.styles":"ready","ext.relatedArticles.styles":"ready","wikibase.client.init":"ready","ext.wikimediaBadges":"ready"};RLPAGEMODULES=["ext.cite.ux-enhancements","mediawiki.page.media","site","mediawiki.page.ready","skins.minerva.scripts","ext.centralNotice.geoIP","ext.centralNotice.startUp","ext.gadget.switcher","ext.urlShortener.toolbar","ext.centralauth.centralautologin","ext.popups","mobile.init","ext.echo.centralauth","ext.relatedArticles.readMore.bootstrap","ext.eventLogging","ext.wikimediaEvents","ext.navigationTiming","ext.cx.eventlogging.campaigns","ext.cx.entrypoints.mffrequentlanguages","ext.cx.entrypoints.languagesearcher.init","mw.externalguidance.init","ext.checkUser.clientHints","ext.growthExperiments.SuggestedEditSession","wikibase.sidebar.tracking"];</script> <script>(RLQ=window.RLQ||[]).push(function(){mw.loader.impl(function(){return["user.options@12s5i",function($,jQuery,require,module){mw.user.tokens.set({"patrolToken":"+\\","watchToken":"+\\","csrfToken":"+\\"}); }];});});</script> <link rel="stylesheet" href="/w/load.php?lang=en&modules=ext.cite.styles%7Cext.relatedArticles.styles%7Cext.wikimediaBadges%7Cext.wikimediamessages.styles%7Cmediawiki.hlist%7Cmobile.init.styles%7Cskins.minerva.amc.styles%7Cskins.minerva.codex.styles%7Cskins.minerva.content.styles.images%7Cskins.minerva.icons%2Cstyles%7Cwikibase.client.init&only=styles&skin=minerva"> <script async="" src="/w/load.php?lang=en&modules=startup&only=scripts&raw=1&skin=minerva"></script> <meta name="ResourceLoaderDynamicStyles" content=""> <link rel="stylesheet" href="/w/load.php?lang=en&modules=site.styles&only=styles&skin=minerva"> <meta name="generator" content="MediaWiki 1.44.0-wmf.5"> <meta name="referrer" content="origin"> <meta name="referrer" content="origin-when-cross-origin"> <meta name="robots" content="max-image-preview:standard"> <meta name="format-detection" content="telephone=no"> <meta name="theme-color" content="#eaecf0"> <meta property="og:image" content="https://upload.wikimedia.org/wikipedia/commons/thumb/1/16/Drosophila_melanogaster_laboratory_culture-vial.jpg/1200px-Drosophila_melanogaster_laboratory_culture-vial.jpg"> <meta property="og:image:width" content="1200"> <meta property="og:image:height" content="1695"> <meta property="og:image" content="https://upload.wikimedia.org/wikipedia/commons/thumb/1/16/Drosophila_melanogaster_laboratory_culture-vial.jpg/800px-Drosophila_melanogaster_laboratory_culture-vial.jpg"> <meta property="og:image:width" content="800"> <meta property="og:image:height" content="1130"> <meta property="og:image" content="https://upload.wikimedia.org/wikipedia/commons/thumb/1/16/Drosophila_melanogaster_laboratory_culture-vial.jpg/640px-Drosophila_melanogaster_laboratory_culture-vial.jpg"> <meta property="og:image:width" content="640"> <meta property="og:image:height" content="904"> <meta name="viewport" content="width=device-width, initial-scale=1.0, user-scalable=yes, minimum-scale=0.25, maximum-scale=5.0"> <meta property="og:title" content="Morphogen - Wikipedia"> <meta property="og:type" content="website"> <link rel="preconnect" href="//upload.wikimedia.org"> <link rel="manifest" href="/w/api.php?action=webapp-manifest"> <link rel="alternate" type="application/x-wiki" title="Edit this page" href="/w/index.php?title=Morphogen&action=edit"> <link rel="apple-touch-icon" href="/static/apple-touch/wikipedia.png"> <link rel="icon" href="/static/favicon/wikipedia.ico"> <link rel="search" type="application/opensearchdescription+xml" href="/w/rest.php/v1/search" title="Wikipedia (en)"> <link rel="EditURI" type="application/rsd+xml" href="//en.wikipedia.org/w/api.php?action=rsd"> <link rel="canonical" href="https://en.wikipedia.org/wiki/Morphogen"> <link rel="license" href="https://creativecommons.org/licenses/by-sa/4.0/deed.en"> <link rel="dns-prefetch" href="//meta.wikimedia.org" /> <link rel="dns-prefetch" href="//login.wikimedia.org"> </head> <body class="mediawiki ltr sitedir-ltr mw-hide-empty-elt ns-0 ns-subject mw-editable page-Morphogen rootpage-Morphogen stable issues-group-B skin-minerva action-view skin--responsive mw-mf-amc-disabled mw-mf"><div id="mw-mf-viewport"> <div id="mw-mf-page-center"> <a class="mw-mf-page-center__mask" href="#"></a> <header class="header-container header-chrome"> <div class="minerva-header"> <nav class="navigation-drawer toggle-list view-border-box"> <input type="checkbox" id="main-menu-input" class="toggle-list__checkbox" role="button" aria-haspopup="true" aria-expanded="false" aria-labelledby="mw-mf-main-menu-button"> <label role="button" for="main-menu-input" id="mw-mf-main-menu-button" aria-hidden="true" data-event-name="ui.mainmenu" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet toggle-list__toggle"> <span class="minerva-icon minerva-icon--menu"></span> <span></span> </label> <div id="mw-mf-page-left" class="menu view-border-box"> <ul id="p-navigation" class="toggle-list__list"> <li class="toggle-list-item "> <a class="toggle-list-item__anchor menu__item--home" href="/wiki/Main_Page" data-mw="interface"> <span class="minerva-icon minerva-icon--home"></span> <span class="toggle-list-item__label">Home</span> </a> </li> <li class="toggle-list-item "> <a class="toggle-list-item__anchor menu__item--random" href="/wiki/Special:Random" data-mw="interface"> <span class="minerva-icon minerva-icon--die"></span> <span class="toggle-list-item__label">Random</span> </a> </li> <li class="toggle-list-item skin-minerva-list-item-jsonly"> <a class="toggle-list-item__anchor menu__item--nearby" href="/wiki/Special:Nearby" data-event-name="menu.nearby" data-mw="interface"> <span class="minerva-icon minerva-icon--mapPin"></span> <span class="toggle-list-item__label">Nearby</span> </a> </li> </ul> <ul id="p-personal" class="toggle-list__list"> <li class="toggle-list-item "> <a class="toggle-list-item__anchor menu__item--login" href="/w/index.php?title=Special:UserLogin&returnto=Morphogen" data-event-name="menu.login" data-mw="interface"> <span class="minerva-icon minerva-icon--logIn"></span> <span class="toggle-list-item__label">Log in</span> </a> </li> </ul> <ul id="pt-preferences" class="toggle-list__list"> <li class="toggle-list-item skin-minerva-list-item-jsonly"> <a class="toggle-list-item__anchor menu__item--settings" href="/w/index.php?title=Special:MobileOptions&returnto=Morphogen" data-event-name="menu.settings" data-mw="interface"> <span class="minerva-icon minerva-icon--settings"></span> <span class="toggle-list-item__label">Settings</span> </a> </li> </ul> <ul id="p-donation" class="toggle-list__list"> <li class="toggle-list-item "> <a class="toggle-list-item__anchor menu__item--donate" href="https://donate.wikimedia.org/wiki/Special:FundraiserRedirector?utm_source=donate&utm_medium=sidebar&utm_campaign=C13_en.wikipedia.org&uselang=en&wmf_key=minerva" data-event-name="menu.donate" data-mw="interface"> <span class="minerva-icon minerva-icon--heart"></span> <span class="toggle-list-item__label">Donate</span> </a> </li> </ul> <ul class="hlist"> <li class="toggle-list-item "> <a class="toggle-list-item__anchor menu__item--about" href="/wiki/Wikipedia:About" data-mw="interface"> <span class="toggle-list-item__label">About Wikipedia</span> </a> </li> <li class="toggle-list-item "> <a class="toggle-list-item__anchor menu__item--disclaimers" href="/wiki/Wikipedia:General_disclaimer" data-mw="interface"> <span class="toggle-list-item__label">Disclaimers</span> </a> </li> </ul> </div> <label class="main-menu-mask" for="main-menu-input"></label> </nav> <div class="branding-box"> <a href="/wiki/Main_Page"> <span><img src="/static/images/mobile/copyright/wikipedia-wordmark-en.svg" alt="Wikipedia" width="120" height="18" style="width: 7.5em; height: 1.125em;"/> </span> </a> </div> <form action="/w/index.php" method="get" class="minerva-search-form"> <div class="search-box"> <input type="hidden" name="title" value="Special:Search"/> <input class="search skin-minerva-search-trigger" id="searchInput" type="search" name="search" placeholder="Search Wikipedia" aria-label="Search Wikipedia" autocapitalize="sentences" title="Search Wikipedia [f]" accesskey="f"> <span class="search-box-icon-overlay"><span class="minerva-icon minerva-icon--search"></span> </span> </div> <button id="searchIcon" class="cdx-button cdx-button--size-large cdx-button--icon-only cdx-button--weight-quiet skin-minerva-search-trigger"> <span class="minerva-icon minerva-icon--search"></span> <span>Search</span> </button> </form> <nav class="minerva-user-navigation" aria-label="User navigation"> </nav> </div> </header> <main id="content" class="mw-body"> <div class="banner-container"> <div id="siteNotice"></div> </div> <div class="pre-content heading-holder"> <div class="page-heading"> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Morphogen</span></h1> <div class="tagline"></div> </div> <ul id="p-associated-pages" class="minerva__tab-container"> <li class="minerva__tab selected"> <a class="minerva__tab-text" href="/wiki/Morphogen" rel="" data-event-name="tabs.subject">Article</a> </li> <li class="minerva__tab "> <a class="minerva__tab-text" href="/wiki/Talk:Morphogen" rel="discussion" data-event-name="tabs.talk">Talk</a> </li> </ul> <nav class="page-actions-menu"> <ul 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"> <span class="minerva-icon minerva-icon--language"></span> <span>Language</span> </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=Morphogen" 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"> <span class="minerva-icon minerva-icon--star"></span> <span>Watch</span> </a> </li> <li id="page-actions-edit" class="page-actions-menu__list-item"> <a role="button" id="ca-edit" href="/w/index.php?title=Morphogen&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"> <span class="minerva-icon minerva-icon--edit"></span> <span>Edit</span> </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"> <style data-mw-deduplicate="TemplateStyles:r1251242444">.mw-parser-output .ambox{border:1px solid #a2a9b1;border-left:10px solid #36c;background-color:#fbfbfb;box-sizing:border-box}.mw-parser-output .ambox+link+.ambox,.mw-parser-output .ambox+link+style+.ambox,.mw-parser-output .ambox+link+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+style+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+link+.ambox{margin-top:-1px}html body.mediawiki .mw-parser-output .ambox.mbox-small-left{margin:4px 1em 4px 0;overflow:hidden;width:238px;border-collapse:collapse;font-size:88%;line-height:1.25em}.mw-parser-output .ambox-speedy{border-left:10px solid #b32424;background-color:#fee7e6}.mw-parser-output .ambox-delete{border-left:10px solid #b32424}.mw-parser-output .ambox-content{border-left:10px solid #f28500}.mw-parser-output .ambox-style{border-left:10px solid #fc3}.mw-parser-output .ambox-move{border-left:10px solid #9932cc}.mw-parser-output .ambox-protection{border-left:10px solid #a2a9b1}.mw-parser-output .ambox .mbox-text{border:none;padding:0.25em 0.5em;width:100%}.mw-parser-output .ambox .mbox-image{border:none;padding:2px 0 2px 0.5em;text-align:center}.mw-parser-output .ambox .mbox-imageright{border:none;padding:2px 0.5em 2px 0;text-align:center}.mw-parser-output .ambox .mbox-empty-cell{border:none;padding:0;width:1px}.mw-parser-output .ambox .mbox-image-div{width:52px}@media(min-width:720px){.mw-parser-output .ambox{margin:0 10%}}@media print{body.ns-0 .mw-parser-output .ambox{display:none!important}}</style><style data-mw-deduplicate="TemplateStyles:r1248332772">.mw-parser-output .multiple-issues-text{width:95%;margin:0.2em 0}.mw-parser-output .multiple-issues-text>.mw-collapsible-content{margin-top:0.3em}.mw-parser-output .compact-ambox .ambox{border:none;border-collapse:collapse;background-color:transparent;margin:0 0 0 1.6em!important;padding:0!important;width:auto;display:block}body.mediawiki .mw-parser-output .compact-ambox .ambox.mbox-small-left{font-size:100%;width:auto;margin:0}.mw-parser-output .compact-ambox .ambox .mbox-text{padding:0!important;margin:0!important}.mw-parser-output .compact-ambox .ambox .mbox-text-span{display:list-item;line-height:1.5em;list-style-type:disc}body.skin-minerva .mw-parser-output .multiple-issues-text>.mw-collapsible-toggle,.mw-parser-output .compact-ambox .ambox .mbox-image,.mw-parser-output .compact-ambox .ambox .mbox-imageright,.mw-parser-output .compact-ambox .ambox .mbox-empty-cell,.mw-parser-output .compact-ambox .hide-when-compact{display:none}</style><table class="box-Multiple_issues plainlinks metadata ambox ambox-content ambox-multiple_issues compact-ambox" role="presentation"><tbody><tr><td class="mbox-text"><div class="mbox-text-span"><div class="multiple-issues-text mw-collapsible"><b>This article has multiple issues.</b> Please help <b><a href="/wiki/Special:EditPage/Morphogen" title="Special:EditPage/Morphogen">improve it</a></b> or discuss these issues on the <b><a href="/wiki/Talk:Morphogen" title="Talk:Morphogen">talk page</a></b>. <small><i>(<a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove these messages</a>)</i></small> <div class="mw-collapsible-content"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1251242444"><table class="box-More_citations_needed plainlinks metadata ambox ambox-content ambox-Refimprove" role="presentation"><tbody><tr><td class="mbox-text"><div class="mbox-text-span">This article <b>needs additional citations for <a href="/wiki/Wikipedia:Verifiability" title="Wikipedia:Verifiability">verification</a></b>.<span class="hide-when-compact"> Please help <a href="/wiki/Special:EditPage/Morphogen" title="Special:EditPage/Morphogen">improve this article</a> by <a href="/wiki/Help:Referencing_for_beginners" title="Help:Referencing for beginners">adding citations to reliable sources</a>. 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-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> <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><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>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> <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><span class="toctogglespan"><label class="toctogglelabel" for="toctogglecheckbox"></label></span></div> <ul> <li class="toclevel-1 tocsection-1"><a href="#History"><span class="tocnumber">1</span> <span class="toctext">History</span></a></li> <li class="toclevel-1 tocsection-2"><a href="#Mechanism"><span class="tocnumber">2</span> <span class="toctext">Mechanism</span></a></li> <li class="toclevel-1 tocsection-3"><a href="#Genes_and_signals"><span class="tocnumber">3</span> <span class="toctext">Genes and signals</span></a></li> <li class="toclevel-1 tocsection-4"><a href="#Examples"><span class="tocnumber">4</span> <span class="toctext">Examples</span></a></li> <li class="toclevel-1 tocsection-5"><a href="#Fruit_fly"><span class="tocnumber">5</span> <span class="toctext">Fruit fly</span></a></li> <li class="toclevel-1 tocsection-6"><a href="#Eponyms"><span class="tocnumber">6</span> <span class="toctext">Eponyms</span></a></li> <li class="toclevel-1 tocsection-7"><a href="#References"><span class="tocnumber">7</span> <span class="toctext">References</span></a></li> <li class="toclevel-1 tocsection-8"><a href="#Further_reading"><span class="tocnumber">8</span> <span class="toctext">Further reading</span></a></li> </ul> </div> </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(1)"><span class="indicator mf-icon mf-icon-expand mf-icon--small"></span><h2 id="History">History</h2><span class="mw-editsection"> <a role="button" href="/w/index.php?title=Morphogen&action=edit&section=1" title="Edit section: History" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> <span class="minerva-icon minerva-icon--edit"></span> <span>edit</span> </a> </span> </div><section class="mf-section-1 collapsible-block" id="mf-section-1"> <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> </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(2)"><span class="indicator mf-icon mf-icon-expand mf-icon--small"></span><h2 id="Mechanism">Mechanism</h2><span class="mw-editsection"> <a role="button" href="/w/index.php?title=Morphogen&action=edit&section=2" 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 "> <span class="minerva-icon minerva-icon--edit"></span> <span>edit</span> </a> </span> </div><section class="mf-section-2 collapsible-block" id="mf-section-2"> <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> </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(3)"><span class="indicator mf-icon mf-icon-expand mf-icon--small"></span><h2 id="Genes_and_signals">Genes and signals</h2><span class="mw-editsection"> <a role="button" href="/w/index.php?title=Morphogen&action=edit&section=3" title="Edit section: Genes and signals" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> <span class="minerva-icon minerva-icon--edit"></span> <span>edit</span> </a> </span> </div><section class="mf-section-3 collapsible-block" id="mf-section-3"> <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> </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(4)"><span class="indicator mf-icon mf-icon-expand mf-icon--small"></span><h2 id="Examples">Examples</h2><span class="mw-editsection"> <a role="button" href="/w/index.php?title=Morphogen&action=edit&section=4" title="Edit section: Examples" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> <span class="minerva-icon minerva-icon--edit"></span> <span>edit</span> </a> </span> </div><section class="mf-section-4 collapsible-block" id="mf-section-4"> <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> </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(5)"><span class="indicator mf-icon mf-icon-expand mf-icon--small"></span><h2 id="Fruit_fly">Fruit fly</h2><span class="mw-editsection"> <a role="button" href="/w/index.php?title=Morphogen&action=edit&section=5" title="Edit section: Fruit fly" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> <span class="minerva-icon minerva-icon--edit"></span> <span>edit</span> </a> </span> </div><section class="mf-section-5 collapsible-block" id="mf-section-5"> <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> </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(6)"><span class="indicator mf-icon mf-icon-expand mf-icon--small"></span><h2 id="Eponyms">Eponyms</h2><span class="mw-editsection"> <a role="button" href="/w/index.php?title=Morphogen&action=edit&section=6" title="Edit section: Eponyms" class="cdx-button cdx-button--size-large cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--icon-only cdx-button--weight-quiet "> <span class="minerva-icon minerva-icon--edit"></span> <span>edit</span> </a> </span> </div><section class="mf-section-6 collapsible-block" id="mf-section-6"> <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> </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(7)"><span class="indicator mf-icon mf-icon-expand mf-icon--small"></span><h2 id="References">References</h2><span class="mw-editsection"> <a role="button" href="/w/index.php?title=Morphogen&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 "> <span class="minerva-icon minerva-icon--edit"></span> <span>edit</span> </a> </span> </div><section class="mf-section-7 collapsible-block" id="mf-section-7"> <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. M. (1952). "The chemical basis of morphogenesis". <i>Philosophical Transactions of the Royal Society of London B</i>. <b>237</b> (641): 37–72. <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/1952RSPTB.237...37T">1952RSPTB.237...37T</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1098%2Frstb.1952.0012">10.1098/rstb.1952.0012</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Philosophical+Transactions+of+the+Royal+Society+of+London+B&rft.atitle=The+chemical+basis+of+morphogenesis&rft.volume=237&rft.issue=641&rft.pages=37-72&rft.date=1952&rft_id=info%3Adoi%2F10.1098%2Frstb.1952.0012&rft_id=info%3Abibcode%2F1952RSPTB.237...37T&rft.au=Turing%2C+A.+M.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMorphogen" class="Z3988"></span></span> </li> <li id="cite_note-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHiscockMegason2015" class="citation journal cs1">Hiscock, Tom W.; Megason, Sean G. (2015). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4707970">"Orientation of Turing-like Patterns by Morphogen Gradients and Tissue Anisotropies"</a>. <i>Cell Systems</i>. <b>1</b> (6): 408–416. <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.cels.2015.12.001">10.1016/j.cels.2015.12.001</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/PMC4707970">4707970</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/26771020">26771020</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Cell+Systems&rft.atitle=Orientation+of+Turing-like+Patterns+by+Morphogen+Gradients+and+Tissue+Anisotropies&rft.volume=1&rft.issue=6&rft.pages=408-416&rft.date=2015&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4707970%23id-name%3DPMC&rft_id=info%3Apmid%2F26771020&rft_id=info%3Adoi%2F10.1016%2Fj.cels.2015.12.001&rft.aulast=Hiscock&rft.aufirst=Tom+W.&rft.au=Megason%2C+Sean+G.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4707970&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMorphogen" class="Z3988"></span></span> </li> <li id="cite_note-Nusslein-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-Nusslein_3-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFNüsslein-Volhard,_C.Wieschaus,_E.1980" class="citation journal cs1">Nüsslein-Volhard, C.; Wieschaus, E. (October 1980). "Mutations affecting segment number and polarity in <i>Drosophila</i>". <i>Nature</i>. <b>287</b> (5785): 795–801. <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/1980Natur.287..795N">1980Natur.287..795N</a>. <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%2F287795a0">10.1038/287795a0</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/6776413">6776413</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:4337658">4337658</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=Mutations+affecting+segment+number+and+polarity+in+Drosophila&rft.volume=287&rft.issue=5785&rft.pages=795-801&rft.date=1980-10&rft_id=info%3Adoi%2F10.1038%2F287795a0&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A4337658%23id-name%3DS2CID&rft_id=info%3Apmid%2F6776413&rft_id=info%3Abibcode%2F1980Natur.287..795N&rft.au=N%C3%BCsslein-Volhard%2C+C.&rft.au=Wieschaus%2C+E.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMorphogen" class="Z3988"></span></span> </li> <li id="cite_note-Arthur2002-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-Arthur2002_4-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFArthur2002" class="citation journal cs1">Arthur, Wallace (14 February 2002). "The emerging conceptual framework of evolutionary developmental biology". <i>Nature</i>. <b>415</b> (6873): 757–764. <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/2002Natur.415..757A">2002Natur.415..757A</a>. <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%2F415757a">10.1038/415757a</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/11845200">11845200</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:4432164">4432164</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=The+emerging+conceptual+framework+of+evolutionary+developmental+biology&rft.volume=415&rft.issue=6873&rft.pages=757-764&rft.date=2002-02-14&rft_id=info%3Adoi%2F10.1038%2F415757a&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A4432164%23id-name%3DS2CID&rft_id=info%3Apmid%2F11845200&rft_id=info%3Abibcode%2F2002Natur.415..757A&rft.aulast=Arthur&rft.aufirst=Wallace&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMorphogen" class="Z3988"></span></span> </li> <li id="cite_note-Winchester-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-Winchester_5-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWinchester2004" class="citation journal cs1">Winchester, Guil (2004). <a rel="nofollow" class="external text" href="http://www.cell.com/current-biology/pdf/S0960-9822(04)00683-9.pdf">"Edward B. Lewis 1918-2004"</a> <span class="cs1-format">(PDF)</span>. <i>Current Biology</i>. <b>14</b> (18) (published Sep 21, 2004): R740–742. <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/2004CBio...14.R740W">2004CBio...14.R740W</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/10.1016%2Fj.cub.2004.09.007">10.1016/j.cub.2004.09.007</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/15380080">15380080</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:32648995">32648995</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Current+Biology&rft.atitle=Edward+B.+Lewis+1918-2004&rft.volume=14&rft.issue=18&rft.pages=R740-742&rft.date=2004&rft_id=info%3Adoi%2F10.1016%2Fj.cub.2004.09.007&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A32648995%23id-name%3DS2CID&rft_id=info%3Apmid%2F15380080&rft_id=info%3Abibcode%2F2004CBio...14.R740W&rft.aulast=Winchester&rft.aufirst=Guil&rft_id=http%3A%2F%2Fwww.cell.com%2Fcurrent-biology%2Fpdf%2FS0960-9822%2804%2900683-9.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMorphogen" class="Z3988"></span></span> </li> <li id="cite_note-6"><span class="mw-cite-backlink"><b><a href="#cite_ref-6">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20161013223611/https://www.ibiology.org/ibiomagazine/eric-wieschaus-and-christiane-nusselin-volhard.html">"Eric Wieschaus and Christiane Nüsslein-Volhard: Collaborating to Find Developmental Genes"</a>. iBiology. Archived from <a rel="nofollow" class="external text" href="https://www.ibiology.org/ibiomagazine/eric-wieschaus-and-christiane-nusselin-volhard.html">the original</a> on 13 October 2016<span class="reference-accessdate">. Retrieved <span class="nowrap">13 October</span> 2016</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Eric+Wieschaus+and+Christiane+N%C3%BCsslein-Volhard%3A+Collaborating+to+Find+Developmental+Genes&rft.pub=iBiology&rft_id=https%3A%2F%2Fwww.ibiology.org%2Fibiomagazine%2Feric-wieschaus-and-christiane-nusselin-volhard.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMorphogen" class="Z3988"></span></span> </li> <li id="cite_note-7"><span class="mw-cite-backlink"><b><a href="#cite_ref-7">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRussell2010" class="citation book cs1">Russell, Peter (2010). <i>iGenetics : a molecular approach</i>. San Francisco, CA: Pearson Benjamin Cummings. p. 566. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-321-56976-9" title="Special:BookSources/978-0-321-56976-9"><bdi>978-0-321-56976-9</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=iGenetics+%3A+a+molecular+approach&rft.place=San+Francisco%2C+CA&rft.pages=566&rft.pub=Pearson+Benjamin+Cummings&rft.date=2010&rft.isbn=978-0-321-56976-9&rft.aulast=Russell&rft.aufirst=Peter&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMorphogen" class="Z3988"></span></span> </li> <li id="cite_note-8"><span class="mw-cite-backlink"><b><a href="#cite_ref-8">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGordonGordon2016" class="citation journal cs1">Gordon, Natalie K.; Gordon, Richard (2016). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4785624">"The organelle of differentiation in embryos: The cell state splitter"</a>. <i>Theoretical Biology and Medical Modelling</i>. <b>13</b>: 11. <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/10.1186%2Fs12976-016-0037-2">10.1186/s12976-016-0037-2</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/PMC4785624">4785624</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/26965444">26965444</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Theoretical+Biology+and+Medical+Modelling&rft.atitle=The+organelle+of+differentiation+in+embryos%3A+The+cell+state+splitter&rft.volume=13&rft.pages=11&rft.date=2016&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4785624%23id-name%3DPMC&rft_id=info%3Apmid%2F26965444&rft_id=info%3Adoi%2F10.1186%2Fs12976-016-0037-2&rft.aulast=Gordon&rft.aufirst=Natalie+K.&rft.au=Gordon%2C+Richard&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4785624&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMorphogen" class="Z3988"></span></span> </li> <li id="cite_note-9"><span class="mw-cite-backlink"><b><a href="#cite_ref-9">^</a></b></span> <span class="reference-text">Roth S., Lynch J <a rel="nofollow" class="external text" href="http://www.cell.com/cell/fulltext/S0092-8674(12)00466-7">Does the Bicoid Gradient Matter?</a> Cell, Volume 149, Issue 3, p511–512, 27 April 2012.</span> </li> <li id="cite_note-Kam_2012-10"><span class="mw-cite-backlink">^ <a href="#cite_ref-Kam_2012_10-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Kam_2012_10-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKamDengChenZhao2012" class="citation journal cs1">Kam RK, Deng Y, Chen Y, Zhao H (2012). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3325842">"Retinoic acid synthesis and functions in early embryonic development"</a>. <i>Cell & Bioscience</i>. <b>2</b> (1): 11. <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/10.1186%2F2045-3701-2-11">10.1186/2045-3701-2-11</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/PMC3325842">3325842</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/22439772">22439772</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=Retinoic+acid+synthesis+and+functions+in+early+embryonic+development.&rft.volume=2&rft.issue=1&rft.pages=11&rft.date=2012&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3325842%23id-name%3DPMC&rft_id=info%3Apmid%2F22439772&rft_id=info%3Adoi%2F10.1186%2F2045-3701-2-11&rft.aulast=Kam&rft.aufirst=RK&rft.au=Deng%2C+Y&rft.au=Chen%2C+Y&rft.au=Zhao%2C+H&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3325842&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMorphogen" class="Z3988"></span></span> </li> <li id="cite_note-Moore_2013-11"><span class="mw-cite-backlink">^ <a href="#cite_ref-Moore_2013_11-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Moore_2013_11-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Moore_2013_11-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Moore_2013_11-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMoorePersaudTorchia2013" class="citation book cs1">Moore KL, Persaud TV, Torchia MG (2013). <a rel="nofollow" class="external text" href="https://www.inkling.com/read/the-developing-human-moore-9th/chapter-21/morphogens">"Common signaling pathways used during development: morphogens"</a>. <i>The developing human: clinically oriented embryology</i> (9th ed.). Philadelphia, PA: Saunders/Elsevier. pp. 506–509. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1437720020" title="Special:BookSources/978-1437720020"><bdi>978-1437720020</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Common+signaling+pathways+used+during+development%3A+morphogens&rft.btitle=The+developing+human%3A+clinically+oriented+embryology&rft.place=Philadelphia%2C+PA&rft.pages=506-509&rft.edition=9th&rft.pub=Saunders%2FElsevier&rft.date=2013&rft.isbn=978-1437720020&rft.aulast=Moore&rft.aufirst=KL&rft.au=Persaud%2C+TV&rft.au=Torchia%2C+MG&rft_id=https%3A%2F%2Fwww.inkling.com%2Fread%2Fthe-developing-human-moore-9th%2Fchapter-21%2Fmorphogens&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMorphogen" class="Z3988"></span></span> </li> <li id="cite_note-Cunningham_and_Duester-12"><span class="mw-cite-backlink"><b><a href="#cite_ref-Cunningham_and_Duester_12-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCunninghamDuester2015" class="citation journal cs1">Cunningham, T.J.; Duester, G. (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> </section><div class="mw-heading mw-heading2 section-heading" onclick="mfTempOpenSection(8)"><span class="indicator mf-icon mf-icon-expand mf-icon--small"></span><h2 id="Further_reading">Further reading</h2><span class="mw-editsection"> <a role="button" href="/w/index.php?title=Morphogen&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 "> <span class="minerva-icon minerva-icon--edit"></span> <span>edit</span> </a> </span> </div><section class="mf-section-8 collapsible-block" id="mf-section-8"> <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 .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:": "}.mw-parser-output .hlist dd::after,.mw-parser-output .hlist li::after{content:" · ";font-weight:bold}.mw-parser-output .hlist dd:last-child::after,.mw-parser-output .hlist dt:last-child::after,.mw-parser-output .hlist li:last-child::after{content:none}.mw-parser-output .hlist dd dd:first-child::before,.mw-parser-output .hlist dd dt:first-child::before,.mw-parser-output .hlist dd li:first-child::before,.mw-parser-output .hlist dt dd:first-child::before,.mw-parser-output .hlist dt dt:first-child::before,.mw-parser-output .hlist dt li:first-child::before,.mw-parser-output .hlist li dd:first-child::before,.mw-parser-output .hlist li dt:first-child::before,.mw-parser-output .hlist li li:first-child::before{content:" (";font-weight:normal}.mw-parser-output .hlist dd dd:last-child::after,.mw-parser-output .hlist dd dt:last-child::after,.mw-parser-output .hlist dd li:last-child::after,.mw-parser-output .hlist dt dd:last-child::after,.mw-parser-output .hlist dt dt:last-child::after,.mw-parser-output .hlist dt li:last-child::after,.mw-parser-output .hlist li dd:last-child::after,.mw-parser-output .hlist li dt:last-child::after,.mw-parser-output .hlist li li:last-child::after{content:")";font-weight:normal}.mw-parser-output .hlist ol{counter-reset:listitem}.mw-parser-output .hlist ol>li{counter-increment:listitem}.mw-parser-output .hlist ol>li::before{content:" "counter(listitem)"\a0 "}.mw-parser-output .hlist dd ol>li:first-child::before,.mw-parser-output .hlist dt ol>li:first-child::before,.mw-parser-output .hlist li ol>li:first-child::before{content:" ("counter(listitem)"\a0 "}</style><style data-mw-deduplicate="TemplateStyles:r1236075235">.mw-parser-output .navbox{box-sizing:border-box;border:1px solid #a2a9b1;width:100%;clear:both;font-size:88%;text-align:center;padding:1px;margin:1em auto 0}.mw-parser-output .navbox .navbox{margin-top:0}.mw-parser-output .navbox+.navbox,.mw-parser-output .navbox+.navbox-styles+.navbox{margin-top:-1px}.mw-parser-output .navbox-inner,.mw-parser-output .navbox-subgroup{width:100%}.mw-parser-output .navbox-group,.mw-parser-output .navbox-title,.mw-parser-output .navbox-abovebelow{padding:0.25em 1em;line-height:1.5em;text-align:center}.mw-parser-output .navbox-group{white-space:nowrap;text-align:right}.mw-parser-output .navbox,.mw-parser-output .navbox-subgroup{background-color:#fdfdfd}.mw-parser-output .navbox-list{line-height:1.5em;border-color:#fdfdfd}.mw-parser-output .navbox-list-with-group{text-align:left;border-left-width:2px;border-left-style:solid}.mw-parser-output tr+tr>.navbox-abovebelow,.mw-parser-output tr+tr>.navbox-group,.mw-parser-output tr+tr>.navbox-image,.mw-parser-output tr+tr>.navbox-list{border-top:2px solid #fdfdfd}.mw-parser-output .navbox-title{background-color:#ccf}.mw-parser-output .navbox-abovebelow,.mw-parser-output .navbox-group,.mw-parser-output .navbox-subgroup .navbox-title{background-color:#ddf}.mw-parser-output .navbox-subgroup .navbox-group,.mw-parser-output .navbox-subgroup .navbox-abovebelow{background-color:#e6e6ff}.mw-parser-output .navbox-even{background-color:#f7f7f7}.mw-parser-output .navbox-odd{background-color:transparent}.mw-parser-output .navbox .hlist td dl,.mw-parser-output .navbox .hlist td ol,.mw-parser-output .navbox .hlist td ul,.mw-parser-output .navbox td.hlist dl,.mw-parser-output .navbox td.hlist ol,.mw-parser-output .navbox td.hlist ul{padding:0.125em 0}.mw-parser-output .navbox .navbar{display:block;font-size:100%}.mw-parser-output .navbox-title .navbar{float:left;text-align:left;margin-right:0.5em}body.skin--responsive .mw-parser-output .navbox-image img{max-width:none!important}@media print{body.ns-0 .mw-parser-output .navbox{display:none!important}}</style></div>    </section></div> <noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1&useformat=mobile" 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=1260200925">https://en.wikipedia.org/w/index.php?title=Morphogen&oldid=1260200925</a>"</div></div> </div> <div class="post-content" id="page-secondary-actions"> </div> </main> <footer class="mw-footer minerva-footer" role="contentinfo"> <a class="last-modified-bar" href="/w/index.php?title=Morphogen&action=history"> <div class="post-content last-modified-bar__content"> <span class="minerva-icon minerva-icon-size-medium minerva-icon--modified-history"></span> <span class="last-modified-bar__text modified-enhancement" data-user-name="Citation bot" data-user-gender="unknown" data-timestamp="1732886105"> <span>Last edited on 29 November 2024, at 13:15</span> </span> <span class="minerva-icon minerva-icon-size-small minerva-icon--expand"></span> </div> </a> <div class="post-content footer-content"> <div id='mw-data-after-content'> <div class="read-more-container"></div> </div> <div id="p-lang"> <h4>Languages</h4> <section> <ul id="p-variants" class="minerva-languages"></ul> <ul class="minerva-languages"><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Morfogen" title="Morfogen – Catalan" lang="ca" hreflang="ca" data-title="Morfogen" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Morphogen" 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" data-title="Morfogen" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target"><span>Norsk bokmål</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Morfogen" title="Morfogen – Polish" lang="pl" hreflang="pl" data-title="Morfogen" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%9C%D0%BE%D1%80%D1%84%D0%BE%D0%B3%D0%B5%D0%BD" title="Морфоген – Serbian" lang="sr" hreflang="sr" data-title="Морфоген" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target"><span>Српски / srpski</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Morfogeeni" title="Morfogeeni – Finnish" lang="fi" hreflang="fi" data-title="Morfogeeni" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv badge-Q70893996 mw-list-item" title=""><a href="https://sv.wikipedia.org/wiki/Morfogen" title="Morfogen – Swedish" lang="sv" hreflang="sv" data-title="Morfogen" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></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 29 November 2024, at 13:15<span class="anonymous-show"> (UTC)</span>.</li> <li id="footer-info-copyright">Content is available under <a class="external" rel="nofollow" href="https://creativecommons.org/licenses/by-sa/4.0/deed.en">CC BY-SA 4.0</a> unless otherwise noted.</li> </ul> <ul id="footer-places" class="footer-places hlist hlist-separated"> <li id="footer-places-privacy"><a href="https://foundation.wikimedia.org/wiki/Special:MyLanguage/Policy:Privacy_policy">Privacy policy</a></li> <li id="footer-places-about"><a href="/wiki/Wikipedia:About">About Wikipedia</a></li> <li id="footer-places-disclaimers"><a href="/wiki/Wikipedia:General_disclaimer">Disclaimers</a></li> <li id="footer-places-contact"><a href="//en.wikipedia.org/wiki/Wikipedia:Contact_us">Contact Wikipedia</a></li> <li id="footer-places-wm-codeofconduct"><a href="https://foundation.wikimedia.org/wiki/Special:MyLanguage/Policy:Universal_Code_of_Conduct">Code of Conduct</a></li> <li id="footer-places-developers"><a href="https://developer.wikimedia.org">Developers</a></li> <li id="footer-places-statslink"><a href="https://stats.wikimedia.org/#/en.wikipedia.org">Statistics</a></li> <li id="footer-places-cookiestatement"><a href="https://foundation.wikimedia.org/wiki/Special:MyLanguage/Policy:Cookie_statement">Cookie statement</a></li> <li id="footer-places-terms-use"><a href="https://foundation.m.wikimedia.org/wiki/Special:MyLanguage/Policy:Terms_of_Use">Terms of Use</a></li> <li id="footer-places-desktop-toggle"><a id="mw-mf-display-toggle" href="//en.wikipedia.org/w/index.php?title=Morphogen&mobileaction=toggle_view_desktop" data-event-name="switch_to_desktop">Desktop</a></li> </ul> </div> </footer> </div> </div> <div class="mw-notification-area" data-mw="interface"></div>  <script>(RLQ=window.RLQ||[]).push(function(){mw.config.set({"wgHostname":"mw-web.codfw.main-5c59558b9d-777x8","wgBackendResponseTime":181,"wgPageParseReport":{"limitreport":{"cputime":"0.427","walltime":"0.504","ppvisitednodes":{"value":1855,"limit":1000000},"postexpandincludesize":{"value":92144,"limit":2097152},"templateargumentsize":{"value":10031,"limit":2097152},"expansiondepth":{"value":19,"limit":100},"expensivefunctioncount":{"value":7,"limit":500},"unstrip-depth":{"value":1,"limit":20},"unstrip-size":{"value":76119,"limit":5000000},"entityaccesscount":{"value":0,"limit":400},"timingprofile":["100.00% 444.048 1 -total"," 33.93% 150.656 1 Template:Reflist"," 24.34% 108.081 11 Template:Cite_journal"," 19.72% 87.548 1 Template:Multiple_issues"," 17.20% 76.394 1 Template:Genarch"," 17.09% 75.896 2 Template:Navbox"," 13.54% 60.117 1 Template:Short_description"," 11.82% 52.492 2 Template:Ambox"," 11.56% 51.322 1 Template:More_citations_needed"," 8.42% 37.384 2 Template:Pagetype"]},"scribunto":{"limitreport-timeusage":{"value":"0.274","limit":"10.000"},"limitreport-memusage":{"value":6974694,"limit":52428800}},"cachereport":{"origin":"mw-api-int.codfw.main-6fdd9f9b88-xkrbx","timestamp":"20241129131509","ttl":2592000,"transientcontent":false}}});});</script> <script type="application/ld+json">{"@context":"https:\/\/schema.org","@type":"Article","name":"Morphogen","url":"https:\/\/en.wikipedia.org\/wiki\/Morphogen","sameAs":"http:\/\/www.wikidata.org\/entity\/Q417084","mainEntity":"http:\/\/www.wikidata.org\/entity\/Q417084","author":{"@type":"Organization","name":"Contributors to Wikimedia projects"},"publisher":{"@type":"Organization","name":"Wikimedia Foundation, Inc.","logo":{"@type":"ImageObject","url":"https:\/\/www.wikimedia.org\/static\/images\/wmf-hor-googpub.png"}},"datePublished":"2005-04-03T17:44:50Z","dateModified":"2024-11-29T13:15:05Z","image":"https:\/\/upload.wikimedia.org\/wikipedia\/commons\/1\/16\/Drosophila_melanogaster_laboratory_culture-vial.jpg","headline":"biological substance that guides development by non-uniform distribution"}</script><script>(window.NORLQ=window.NORLQ||[]).push(function(){var ns,i,p,img;ns=document.getElementsByTagName('noscript');for(i=0;i<ns.length;i++){p=ns[i].nextSibling;if(p&&p.className&&p.className.indexOf('lazy-image-placeholder')>-1){img=document.createElement('img');img.setAttribute('src',p.getAttribute('data-src'));img.setAttribute('width',p.getAttribute('data-width'));img.setAttribute('height',p.getAttribute('data-height'));img.setAttribute('alt',p.getAttribute('data-alt'));p.parentNode.replaceChild(img,p);}}});</script> </body> </html>