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>Embryonic differentiation waves - 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":"cc8149f1-e780-4877-b520-38183a03ada9","wgCanonicalNamespace":"","wgCanonicalSpecialPageName":false,"wgNamespaceNumber":0,"wgPageName":"Embryonic_differentiation_waves","wgTitle":"Embryonic differentiation waves","wgCurRevisionId":987222691,"wgRevisionId": 987222691,"wgArticleId":55661946,"wgIsArticle":true,"wgIsRedirect":false,"wgAction":"view","wgUserName":null,"wgUserGroups":["*"],"wgPageViewLanguage":"en","wgPageContentLanguage":"en","wgPageContentModel":"wikitext","wgRelevantPageName":"Embryonic_differentiation_waves","wgRelevantArticleId":55661946,"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":7000,"wgRelatedArticlesCompat":[],"wgCentralAuthMobileDomain":true,"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":"ca","autonym":"català","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":"de","autonym":"Deutsch","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":"es","autonym":"español","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":"fi","autonym":"suomi","dir":"ltr"},{"lang":"fj","autonym":"Na Vosa Vakaviti","dir":"ltr"},{"lang":"fo","autonym":"føroyskt","dir":"ltr"},{"lang":"fon","autonym":"fɔ̀ngbè","dir":"ltr"},{"lang":"fr","autonym":"français","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":"he","autonym":"עברית","dir":"rtl"},{"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":"it","autonym":"italiano","dir":"ltr"},{"lang":"iu","autonym":"ᐃᓄᒃᑎᑐᑦ / inuktitut","dir":"ltr"},{"lang":"ja","autonym":"日本語","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":"pl","autonym":"polski","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":"sr","autonym":"српски / srpski","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":"sv","autonym":"svenska","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":"Q48998242","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.quicksurveys.init","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.4"> <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/5/5e/Schematic_of_Cell_State_Splitter_Organelle.jpg"> <meta property="og:image:width" content="1200"> <meta property="og:image:height" content="796"> <meta property="og:image" content="https://upload.wikimedia.org/wikipedia/commons/5/5e/Schematic_of_Cell_State_Splitter_Organelle.jpg"> <meta property="og:image:width" content="800"> <meta property="og:image:height" content="531"> <meta property="og:image:width" content="640"> <meta property="og:image:height" content="425"> <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="Embryonic differentiation waves - 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=Embryonic_differentiation_waves&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/Embryonic_differentiation_waves"> <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-Embryonic_differentiation_waves rootpage-Embryonic_differentiation_waves 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=Embryonic+differentiation+waves" 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=Embryonic+differentiation+waves" 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&utm_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">Embryonic differentiation waves</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/Embryonic_differentiation_waves" rel="" data-event-name="tabs.subject">Article</a> </li> <li class="minerva__tab "> <a class="minerva__tab-text" href="/wiki/Talk:Embryonic_differentiation_waves" 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="" 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 disabled"> <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=Embryonic+differentiation+waves" 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=Embryonic_differentiation_waves&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"><p>A <a href="/wiki/Mechanochemistry" title="Mechanochemistry">mechanochemical</a> based model for <a href="/wiki/Neurulation#Primary_neural_induction" title="Neurulation">primary neural induction</a> was first proposed in 1985 by Brodland and <a href="/wiki/Richard_Gordon_(theoretical_biologist)" title="Richard Gordon (theoretical biologist)">Gordon</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> They proposed that there is a mechanically sensitive bistable organelle made of <a href="/wiki/Microtubules" class="mw-redirect" title="Microtubules">microtubules</a> and <a href="/wiki/Microfilaments" class="mw-redirect" title="Microfilaments">microfilaments</a> in the apical ends of cells within cell sheets that are about to differentiate (that are competent) and these cells are under mechanical tension. The microtubules and microfilaments are in mechanical opposition in a proposed embryonic organelle they called the cell state splitter. Depending on where the cell is within a sheet, the tension will be resolved by either the apical end contracting or the apical end expanding. The resolution will begin at one point and spread over the rest of the tissue limited by other mechanical forces at boundaries. An actual physical wave of contraction has been found which traverses the presumptive neural epithelium of the developing salamander, the <a href="/wiki/Axolotl" title="Axolotl">axolotl</a> (<i>Ambystoma mexicanum</i>). The contraction wave's trajectory was more complex than predicted in the original model however it did originate from the precise location of the <a href="/wiki/Hans_Spemann" title="Hans Spemann">Spemann organizer</a> and traversed only the presumptive neural epithelium.<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> Electron microscopy showed <a href="/wiki/Intermediate_filaments" class="mw-redirect" title="Intermediate filaments">intermediate filaments</a> are also present in the cell state splitter.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Additional waves of both contraction and expansion were also discovered by time lapse photography of axolotl <a href="/wiki/Gastrulation" title="Gastrulation">gastrulation</a>. Among them was a wave of expansion that occurs in ectoderm only in the presumptive epithelium. When the trajectories of the waves were superimposed on the <a href="/wiki/Fate_map" class="mw-redirect" title="Fate map">fate map</a> of the axolotl it was shown that there is a unique combination of expansion and contraction waves that correlates with the tissue types determined during gastrulation and that this set of wave trajectories could explain the shape of the fate map.<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p><figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Schematic_of_Cell_State_Splitter_Organelle.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5e/Schematic_of_Cell_State_Splitter_Organelle.jpg/220px-Schematic_of_Cell_State_Splitter_Organelle.jpg" decoding="async" width="220" height="146" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/5/5e/Schematic_of_Cell_State_Splitter_Organelle.jpg 1.5x" data-file-width="324" data-file-height="215"></a><figcaption>Schematic of cell state splitter organelle</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:How_Embryonic_Differentiation_Waves_Create_Gradients.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2b/How_Embryonic_Differentiation_Waves_Create_Gradients.jpg/220px-How_Embryonic_Differentiation_Waves_Create_Gradients.jpg" decoding="async" width="220" height="402" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/2/2b/How_Embryonic_Differentiation_Waves_Create_Gradients.jpg 1.5x" data-file-width="287" data-file-height="525"></a><figcaption>How embryonic differentiation waves create gradients</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Differentiation_Tree_of_the_Axolotl.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/86/Differentiation_Tree_of_the_Axolotl.jpg/220px-Differentiation_Tree_of_the_Axolotl.jpg" decoding="async" width="220" height="170" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/86/Differentiation_Tree_of_the_Axolotl.jpg/330px-Differentiation_Tree_of_the_Axolotl.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/86/Differentiation_Tree_of_the_Axolotl.jpg/440px-Differentiation_Tree_of_the_Axolotl.jpg 2x" data-file-width="1254" data-file-height="969"></a><figcaption>Differentiation tree of the <a href="/wiki/Axolotl" title="Axolotl">Axolotl</a></figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Signal_Transduction_Model_for_Embryonic_Differentiation_Waves.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5a/Signal_Transduction_Model_for_Embryonic_Differentiation_Waves.png/220px-Signal_Transduction_Model_for_Embryonic_Differentiation_Waves.png" decoding="async" width="220" height="158" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5a/Signal_Transduction_Model_for_Embryonic_Differentiation_Waves.png/330px-Signal_Transduction_Model_for_Embryonic_Differentiation_Waves.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5a/Signal_Transduction_Model_for_Embryonic_Differentiation_Waves.png/440px-Signal_Transduction_Model_for_Embryonic_Differentiation_Waves.png 2x" data-file-width="635" data-file-height="455"></a><figcaption>Signal transduction model for embryonic differentiation waves</figcaption></figure> <p>A biochemical basis for the signal transduction from the cytoskeleton to the nucleus resulting in changes in gene expression was first proposed by Björklund (now Gordon) and Gordon in 1993<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> This would result in a biochemical transduction of the biomechanical signal from the <a href="/wiki/Cytoskeleton" title="Cytoskeleton">cytoskeleton</a> that is thereby passed on to the nucleus. This then signals the changes in gene expression. If the cell has experienced contraction, one signal is sent and if the cell has experienced expansion then another signal is sent. The signal from the cytoskeleton is what causes determination of cell fate. The phenomena of gene gradients during development is dismissed as an epiphenomena resulting from the passage of the biomechanical wave initiating changes in gene expression in individual cells as the wave passes through a cell sheet.<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> They have outlined their research and their theory of differentiation waves in detail in their book <i>Embryogenesis Explained</i>. For example, the first differentiation that takes place during mammalian compaction is explained in terms of their differentiation waves model thus; Cells on the outside of the <a href="/wiki/Morula" class="mw-redirect" title="Morula">morula</a> expand due to the effect of their position on the outside of the early ball of cells and they become determined to be <a href="/wiki/Trophoblast" title="Trophoblast">trophoblast</a>. Cells on the inside of the ball contract instead due to the mechanical force of being on the inside and they become determined to be the <a href="/wiki/Inner_cell_mass" title="Inner cell mass">inner cell mass</a>. All the other activity, such as changes in gene expression, signalling proteins, release of <a href="/wiki/Morphogen" title="Morphogen">morphogens</a>, and <a href="/wiki/Epigenetics" title="Epigenetics">epigenetic changes</a>, are considered the result of differentiation after the response of the cytoskeleton to mechanical signals which then determines cell fate using purely mechanical signals.<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><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> Failure of <a href="/wiki/Neural_tube" title="Neural tube">neural tube</a> closure is explained as a failure of methylation of cytoskeleton of developing neural tissue for those <a href="/wiki/Neural_tube_defects" class="mw-redirect" title="Neural tube defects">neural tube defects</a> which are <a href="/wiki/Folate" title="Folate">folate</a> sensitive and prevented by <a href="/wiki/Folic_acid" class="mw-redirect" title="Folic acid">folic acid</a> supplementation.<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(1)"><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=Embryonic_differentiation_waves&action=edit&section=1" 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-1 collapsible-block" id="mf-section-1"> <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"> <div class="mw-references-wrap"><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">Gordon, R. Brodland, GW. <a rel="nofollow" class="external text" href="https://link.springer.com/article/10.1007/BF02797122">The cytoskeletal mechanics of brain morphogenesis: cell state splitters cause primary neural induction.</a> Gell Biophys. 11: 177-238. (1987)</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">Brodland, GW” Gordon, R, Scott MJ, Bjorklund NK, Luchka KB, Martin, CC, Matuga, C., Globus, M., Vethamany-Globus S. and Shu, D. <a rel="nofollow" class="external text" href="http://onlinelibrary.wiley.com/doi/10.1002/jmor.1052190203/abstract">Furrowing surface contraction wave coincident with primary neural induction in amphibian embryos.</a> J Morphol. 219: 131-142. 1994</span> </li> <li id="cite_note-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text">Martin, C.C. & R. Gordon. Ultrastructural analysis of the cell state splitter in <a href="/wiki/Ectoderm" title="Ectoderm">ectoderm</a> cells differentiating to neural plate and epidermis during gastrulation in embryos of the axolotl Ambystoma mexicanum. Russian J. Dev. Biol. 28(2), 71-80 1997</span> </li> <li id="cite_note-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-4">^</a></b></span> <span class="reference-text">Björklund, N.K. & R. Gordon (1994). Surface contraction and expansion waves correlated with differentiation in axolotl embryos. I. Prolegomenon and differentiation during the plunge through the blastopore, as shown by the fate map. Computers & Chemistry 18(3), 333- 345 [Appendix VI].</span> </li> <li id="cite_note-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-5">^</a></b></span> <span class="reference-text">Björklund, N.K. & R. Gordon. [Nuclear state splitting: a working model for the mechanochemical coupling of differentiation "waves" with the controlling genes (master genes)] [Russian]. Ontogenez 24(2), 5-23 1993</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">Gordon, R., Björklund, N, Nieuwkoop, PD, International Review of Cytology, <a rel="nofollow" class="external text" href="http://www.sciencedirect.com/science/article/pii/S0074769608615454">Appendix: Dialogue on Embryonic Induction and Differentiation Waves</a>, Vol 150 1994/02/01, Pg 373-420</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">Gordon, N. Gordon, R.<a rel="nofollow" class="external text" href="http://www.worldscientific.com/worldscibooks/10.1142/8152">Embryogenesis Explained</a> <a href="/wiki/World_Scientific_Publishing" class="mw-redirect" title="World Scientific Publishing">World Scientific Publishing</a>, Singapore, 2016</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">Gordon, NK, Gordon R <a rel="nofollow" class="external text" href="https://link.springer.com/article/10.1186/s12976-016-0037-2">The organelle of differentiation in embryos: the cell state splitter</a> Theor Biol Med Model (2016) 13: 11. <a rel="nofollow" class="external free" href="https://doi.org/10.1186/s12976-016-0037-2">https://doi.org/10.1186/s12976-016-0037-2</a></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">Björklund NK, Gordon R <a rel="nofollow" class="external text" href="http://www.ijdb.ehu.es/web/paper.php?doi=052102nb">A hypothesis linking low folate intake to neural tube defects due to failure of post-translation methylations of the cytoskeleton</a> Int. J. Dev. Biol. 50: 135 - 141 2006</span> </li> </ol></div></div> <ul><li><i>As of <a class="external text" href="https://en.wikipedia.org/w/index.php?title=Embryonic_differentiation_waves&oldid=%7B%7B%7Brevision%7D%7D%7D">this edit</a>, this article uses content from </i><a rel="nofollow" class="external text" href="https://biology.stackexchange.com/questions/9276/which-exact-mechanism-triggers-the-first-cell-differentiation-after-n-divisions">"which exact mechanism triggers the first cell differentiation after n divisions?"</a><i>, which is licensed in a way that permits reuse under the <a href="/wiki/Wikipedia:Text_of_Creative_Commons_Attribution-ShareAlike_3.0_Unported_License" class="mw-redirect" title="Wikipedia:Text of Creative Commons Attribution-ShareAlike 3.0 Unported License">Creative Commons Attribution-ShareAlike 3.0 Unported License</a>, but not under the <a href="/wiki/Wikipedia:Text_of_the_GNU_Free_Documentation_License" title="Wikipedia:Text of the GNU Free Documentation License">GFDL</a>. All relevant terms must be followed.</i></li></ul>    </section></div> <noscript><img src="https://login.m.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1&mobile=1" 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=Embryonic_differentiation_waves&oldid=987222691">https://en.wikipedia.org/w/index.php?title=Embryonic_differentiation_waves&oldid=987222691</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=Embryonic_differentiation_waves&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="Akrasia25" data-user-gender="unknown" data-timestamp="1604599736"> <span>Last edited on 5 November 2020, at 18:08</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"></ul> <p>This page is not available in other languages.</p> </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 5 November 2020, at 18:08<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=Embryonic_differentiation_waves&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-5cd4cd96d5-btbm7","wgBackendResponseTime":189,"wgPageParseReport":{"limitreport":{"cputime":"0.042","walltime":"0.055","ppvisitednodes":{"value":229,"limit":1000000},"postexpandincludesize":{"value":1442,"limit":2097152},"templateargumentsize":{"value":387,"limit":2097152},"expansiondepth":{"value":8,"limit":100},"expensivefunctioncount":{"value":0,"limit":500},"unstrip-depth":{"value":0,"limit":20},"unstrip-size":{"value":6163,"limit":5000000},"entityaccesscount":{"value":0,"limit":400},"timingprofile":["100.00% 24.933 1 -total"," 93.78% 23.383 1 Template:Reflist"," 5.96% 1.486 1 Template:CCBYSASource"," 5.65% 1.408 2 Template:Main_other"]},"scribunto":{"limitreport-timeusage":{"value":"0.002","limit":"10.000"},"limitreport-memusage":{"value":589522,"limit":52428800}},"cachereport":{"origin":"mw-api-int.codfw.main-6bcd787d7f-fq4lm","timestamp":"20241126040533","ttl":2592000,"transientcontent":false}}});});</script> <script type="application/ld+json">{"@context":"https:\/\/schema.org","@type":"Article","name":"Embryonic differentiation waves","url":"https:\/\/en.wikipedia.org\/wiki\/Embryonic_differentiation_waves","sameAs":"http:\/\/www.wikidata.org\/entity\/Q48998242","mainEntity":"http:\/\/www.wikidata.org\/entity\/Q48998242","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":"2017-10-29T23:28:07Z","dateModified":"2020-11-05T18:08:56Z","image":"https:\/\/upload.wikimedia.org\/wikipedia\/commons\/5\/5e\/Schematic_of_Cell_State_Splitter_Organelle.jpg"}</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>