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Olu phononongo luvavanye iyantlukwano yengingqi kwi-cranial morphology yabantu kusetyenziswa imodeli yejiyometri yehomology esekwe kwidatha yokuskena evela kumaqela eentlanga angama-148 kwihlabathi liphela.Le ndlela isebenzisa itekhnoloji yokufaka itekhnoloji ukuvelisa i-meshes e-homologous ngokwenza iinguqu ezingaguqukiyo usebenzisa i-algorithm yenqaku eliphindaphindayo.Ngokusebenzisa uhlalutyo lwecandelo eliphambili kwiimodeli ze-342 ezikhethiweyo ze-homologous, utshintsho olukhulu kubukhulu becala lufunyenwe kwaye luqinisekiswe ngokucacileyo kwi-skull encinci evela eMzantsi Asia.Umahluko omkhulu wesibini ngumlinganiselo wobude ukuya kububanzi be-neurocranium, ebonisa umahluko phakathi kokhakhayi olude lwama-Afrika kunye nokhakhayi oluyiconvex lwaseMntla-mpuma Asia.Kubalulekile ukuqaphela ukuba esi sithako asinanto yakwenza nokujika kobuso.Iimpawu zobuso ezaziwayo ezifana nezidlele eziphuma kuMntla-mpuma we-Asiya kunye namathambo e-compact maxillary kubantu baseYurophu aphinde aqinisekiswa.Olu tshintsho lobuso luhambelana ngokusondeleyo ne-contour ye-skull, ngokukodwa iqondo lokuthambekela kwamathambo angaphambili kunye ne-occipital.Iipateni ze-Allometric zifunyenwe ngokulingana kobuso ngokumalunga nobukhulu bokakayi jikelele;kukhakhayi ezinkulu imbonakalo yobuso ithande ukuba nde kwaye ibe mxinwa, njengoko kubonisiwe kubantu abaninzi bomthonyama baseMelika nakuMntla-mpuma Asia.Nangona uphononongo lwethu aluzange lubandakanye idatha kwiinguqu zendalo ezinokuchaphazela i-cranial morphology, njengemozulu okanye iimeko zokutya, isethi enkulu yedatha yeepatheni ze-homologous cranial iya kuba luncedo ekufuneni iinkcazo ezahlukeneyo kwiimpawu ze-phenotypic zamathambo.
Umahluko ngokwejografi kwimilo yokakayi bomntu sele ifundwe ixesha elide.Abaphandi abaninzi baye bavavanya ukuhlukahluka kokusingqongileyo kunye / okanye ukukhethwa kwendalo, ngokukodwa iimeko zemozulu1,2,3,4,5,6,7 okanye umsebenzi we-masticatory kuxhomekeke kwiimeko zesondlo5,8,9,10, 11,12.13. .Ukongezelela, ezinye izifundo zigxininise kwimiphumo ye-bottleneck, i-genetic drift, i-gene flow, okanye iinkqubo ze-stochastic evolutionary ezibangelwa ukuguqulwa kwemfuza okungathathi hlangothi14,15,16,17,18,19,20,21,22,23.Umzekelo, imilo engqukuva yevault ebanzi kunye nemfutshane ye-cranial vault icaciswe njengokulungelelaniswa koxinzelelo olukhethiweyo ngokomthetho ka-Allen24, ochaza ukuba izilwanyana ezincancisayo zinciphisa ukulahleka kobushushu ngokunciphisa indawo yomzimba ngokumalunga nomthamo2,4,16,17,25 .Ukongezelela, ezinye iiphononongo ezisebenzisa i-rule26 ye-Bergmann zichaze ubudlelwane phakathi kobukhulu bekhanda kunye neqondo lokushisa3,5,16,25,27, ebonisa ukuba ubungakanani ngokubanzi buvame ukuba bukhulu kwimimandla ebandayo ukukhusela ukulahleka kobushushu.Impembelelo yomatshini woxinzelelo lwe-masticatory kwipateni yokukhula kwe-cranial vault kunye namathambo obuso kuye kwaxoxwa ngayo ngokumalunga neemeko zokutya ezibangelwa yinkcubeko yokupheka okanye ukungafani kokuphila phakathi kwamafama kunye nabazingeli-abaqokeleli8,9,11,12,28.Inkcazo ngokubanzi kukuba ukunciphisa uxinzelelo lokuhlafuna kunciphisa ukuqina kwamathambo obuso kunye nezihlunu.Izifundo ezininzi zehlabathi ziye zanxulumanisa ukuhlukahluka kwemilo ye-skull ngokukodwa kwimiphumo ye-phenotypic yomgama ongathathi hlangothi wemfuza kunokuba ulungelelanise indalo21,29,30,31,32.Enye ingcaciso yotshintsho kwimilo yokakayi isekelwe kwingcamango ye-isometric okanye ukukhula kwe-alometric6,33,34,35.Umzekelo, ubuchopho obukhulu buthanda ukuba neelobe zangaphambili ezibanzi ngokwentelekiso kwindawo ebizwa ngokuba yi "Broca's cap" ummandla, kwaye ububanzi beelobes ezingaphambili buyanda, inkqubo yenguquko ethathelwa ingqalelo ngokusekwe ekukhuleni kweallometric.Ukongeza, uphononongo oluvavanya utshintsho lwexesha elide kwimilo yokakayi lufumene ukuthambekela kwe-allometric malunga ne-brachycephaly (utyekelo lokhakhayi ukuba lube ngqukuva ngakumbi) ngokunyuka kobude33.
Imbali ende yophando kwi-cranial morphology ibandakanya iinzame zokuchonga izinto ezisisiseko ezinoxanduva lwemiba eyahlukeneyo yokwahluka kweemilo zecranial.Iindlela zemveli ezisetyenzisiweyo kwizifundo ezininzi zangaphambili zazisekelwe kwidatha yokulinganisa umgca we-bivariate, ngokuphindaphindiweyo usebenzisa iinkcazo zikaMartin okanye zikaHowell36,37.Ngelo xesha, uninzi lwezifundo ezikhankanywe ngasentla zisebenzisa iindlela eziphambili ezisekelwe kwi-spatial 3D geometric morphometry (GM) iteknoloji5,7,10,11,12,13,17,20,27,34,35,38.39. Umzekelo, indlela ye-sliding semilandmark, esekelwe ekunciphiseni amandla okugoba, ibe yeyona ndlela isetyenziswa ngokuqhelekileyo kwi-biology ye-transgenic.Iprojekthi iprojekthi ye-semi-landmarks yethemplethi kwisampulu nganye ngokutyibilika ecaleni kwegophe okanye umphezulu38,40,41,42,43,44,45,46.Kubandakanya iindlela ezinjalo eziphezulu, ezininzi izifundo ze-3D GM zisebenzisa uhlalutyo oluqhelekileyo lwe-Procrustes, i-iterative pointest (ICP) i-algorithm 47 ukuvumela ukuthelekiswa ngokuthe ngqo kweemilo kunye nokubamba utshintsho.Kungenjalo, ipleyiti encinci spline (TPS)48,49 indlela ikwasetyenziswa ngokubanzi njengendlela yenguqu engangqongqo yokwenza imephu yolungelelwaniso lwe-semilandmark kwiimilo ezisekwe ngomnatha.
Ngophuhliso lweskena esisebenzayo se-3D somzimba uphela ukusukela ekupheleni kwenkulungwane yama-20, izifundo ezininzi zisebenzise izikena zomzimba wonke we-3D kwimilinganiselo yobungakanani50,51.Idatha yokuskena isetyenziselwe ukukhupha imilinganiselo yomzimba, efuna ukuchazwa kweemilo zomphezulu njengemigangatho kuneengqimba zamafu.Ukufakwa kwepateni bubuchule obuphuhliselwe le njongo kwicandelo lemizobo yekhompyuter, apho ubume bomphezulu buchazwa yimodeli ye-polygonal mesh.Inyathelo lokuqala ekufakeni ipateni kukulungisa imodeli yemesh ukuze isetyenziswe njenge template.Ezinye zeentsika ezenza ipateni ziimpawu zomhlaba.Ithemplate iye ikhubazeke kwaye ihambelane nomphezulu ukunciphisa umgama phakathi kwetemplate kunye nelifu lenqaku ngelixa ugcina iimpawu zemilo yendawo yetemplate.Iimpawu zomhlaba kwitemplate zihambelana neempawu zomhlaba kwilifu lenqaku.Ukusebenzisa i-template fitting, yonke idatha yokuskena inokuchazwa njengemodeli ye-mesh enenani elifanayo lamanqaku edatha kunye ne-topology efanayo.Nangona i-homology echanekileyo ikhona kuphela kwiindawo eziphawulekayo, kunokucingelwa ukuba kukho i-homology ngokubanzi phakathi kweemodeli ezenziweyo ukususela ekubeni utshintsho kwijometri yeetemplates zincinci.Ke ngoko, iimodeli zegridi ezenziwe ngokufakwa kwetemplate ngamanye amaxesha zibizwa ngokuba yimodeli ye-homology52.I-advanteji yokufakela itemplate kukuba itemplate inokonakala kwaye ilungelelaniswe kwiindawo ezahlukeneyo zento ekujoliswe kuyo ekufutshane nendawo kodwa kude kuyo (umzekelo, i-zygomatic arch kunye nommandla wexeshana wokakayi) ngaphandle kokuchaphazela nganye. enye.deformation.Ngale ndlela, ithemplate inokukhuselwa kwizinto ze-branching ezifana ne-torso okanye ingalo, kunye nehlombe kwindawo yokuma.Ukungalungi kokufakwa kwetemplate yindleko ephezulu yokubala yokuphindaphinda ngokuphindaphindiweyo, nangona kunjalo, ngenxa yokuphuculwa okubonakalayo ekusebenzeni kwekhompyuter, oku akuseyona nto.Ngokuhlalutya ulungelelwaniso lwamaxabiso ee-vertices ezenza imodeli yemesh usebenzisa iindlela zokuhlalutya izinto ezininzi ezinje ngohlalutyo lwecandelo eliphambili (PCA), kunokwenzeka ukuhlalutya utshintsho kuyo yonke imilo yomphezulu kunye nemilo ebonakalayo nakweyiphi na indawo ekusasazeni.inokufunyanwa.Bala uze ubone umfanekiso53.Namhlanje, iimodeli zemesh eziveliswe ngokufakela itemplate zisetyenziswa ngokubanzi kuhlalutyo lwemilo kwiinkalo ezahlukeneyo52,54,55,56,57,58,59,60.
Ukuqhubela phambili kwitekhnoloji yokurekhoda ye-mesh eguquguqukayo, kunye nophuhliso olukhawulezayo lwezixhobo zokuskena ze-3D eziphathwayo ezikwazi ukuskena kwisisombululo esiphezulu, isantya, kunye nokuhamba kune-CT, zenza kube lula ukurekhoda idatha yomphezulu we-3D kungakhathaliseki indawo.Ke ngoko, kwinkalo ye-anthropology yebhayoloji, ezo teknoloji zintsha zonyusa amandla okulinganisa kunye nokuhlalutya ngokweenkcukacha iisampulu zabantu, kubandakanywa imizekelo yokakayi, okuyinjongo yolu phononongo.
Isishwankathelo, olu phononongo lusebenzisa itekhnoloji ye-3D ye-homology ephuculweyo esekwe kuthelekiso lwetemplate (Umfanekiso 1) ukuvavanya imizekelo yokakayi engama-342 ekhethwe kuluntu lwe-148 kwihlabathi liphela ngokuthelekisa ngokwejografi kwihlabathi liphela.Ukwahlukana kwe-cranial morphology (Itheyibhile 1).Ukuphendula ngotshintsho kwi-skull morphology, sasebenzisa i-PCA kunye ne-receiver operating character (ROC) ihlalutya kwiseti yedatha yemodeli ye-homology esiyivelisayo.Iziphumo ziya kuba negalelo ekuqondeni ngcono utshintsho lwehlabathi kwi-cranial morphology, kubandakanywa iipateni zengingqi kunye nokunciphisa umyalelo wokutshintsha, utshintsho oluhambelanayo phakathi kwamacandelo e-cranial, kunye nobukho beendlela ze-allometric.Nangona olu pho nonongo alujonganga idatha kwiinguqu zangaphandle ezimelwe yimozulu okanye iimeko zokutya ezinokuchaphazela i-cranial morphology, iipatheni zejografi ze-cranial morphology ezibhalwe kwisifundo sethu ziya kunceda ukuphonononga imeko yendalo, i-biomechanical, kunye ne-genetic factor of cranial variation.
Itheyibhile 2 ibonisa ixabiso le-eigenvalues kunye ne-PCA yegalelo lee-coefficients ezisetyenzisiweyo kwi-dataset engafanelekanga ye-17,709 vertices (53,127 XYZ coordinates) ye-342 yemodeli ye-skull e-homologous.Ngenxa yoko, amacandelo aphambili ali-14 achongiwe, igalelo lawo kumahluko xa lilonke lalingaphezu kwe-1%, kwaye isabelo sisonke somahluko sasingama-83.68%.Iivektha zokulayisha zamacandelo aziinqununu ali-14 zirekhodwa kwiTheyibhile eyoNgezelelweyo ye-S1, kwaye amanqaku ecandelo abalwe kwiisampulu zokakayi ezingama-342 anikwe kwiTheyibhile eyoNgezelelweyo ye-S2.
Olu phononongo luvavanye amacandelo angundoqo alithoba ngegalelo elikhulu kune-2%, amanye awo abonisa umahluko omkhulu nobalulekileyo wejografi kwi-cranial morphology.Umzobo we-2 wezicwangciso ze-curves eziveliswe kuhlalutyo lwe-ROC ukubonisa eyona nxalenye ye-PCA esebenzayo yokubonakalisa okanye ukwahlula indibaniselwano nganye yeesampulu kwiiyunithi ezinkulu zejografi (umzekelo, phakathi kwamazwe ase-Afrika kunye namazwe angewona ase-Afrika).Ukudibanisa kwePolynesia akuzange kuhlolwe ngenxa yesayizi encinci yesampuli esetyenziswe kolu vavanyo.Idatha malunga nokubaluleka kweyantlukwano kwi-AUC kunye nezinye izibalo ezisisiseko ezibalwe kusetyenziswa uhlalutyo lwe-ROC ziboniswa kwi-Supplementary Table S3.
Iigophe ze-ROC zisetyenziswe kuqikelelo lwecandelo elisithoba eliphambili elisekelwe kwidathasethi ye-vertex ebandakanya i-342 yeemodeli ze-skull zamadoda angama-homologous.I-AUC: Indawo ephantsi kwegophe kwi-0.01% yokubaluleka esetyenziselwa ukwahlula indibaniselwano nganye yejografi ukusuka kwezinye iindibaniselwano zizonke.I-TPF iyinyani yokwenene (ucalucalulo olusebenzayo), i-FPF inobuxoki (ucalulo olungekho mthethweni).
Ukuchazwa kwe-curve ye-ROC kushwankathelwa ngezantsi, kugxininise kuphela kumacandelo angakwazi ukwahlula amaqela okuthelekisa ngokuba ne-AUC enkulu okanye enkulu kunye nenqanaba eliphezulu lokubaluleka okunokwenzeka ngaphantsi kwe-0.001.I-complex yase-Asia yaseMzantsi (Umfanekiso we-2a), equka ubukhulu becala iisampulu ezivela e-Indiya, yahluke kakhulu kwezinye iisampuli ezixutywe ngokwejografi ukuba icandelo lokuqala (PC1) line-AUC enkulu kakhulu (0.856) xa kuthelekiswa namanye amacandelo.Isici se-African complex (Umfanekiso 2b) yi-AUC enkulu ye-PC2 (0.834).I-Austro-Melanesians (Umfanekiso we-2c) ubonise indlela efanayo kubantu base-Afrika base-Sub-Saharan nge-PC2 kunye ne-AUC enkulu (0.759).Abantu baseYurophu (umzobo 2d) bahluke ngokucacileyo ekudityanisweni kwe-PC2 (AUC = 0.801), i-PC4 (AUC = 0.719) kunye ne-PC6 (AUC = 0.671), isampuli ye-Northeast Asia (Fig. 2e) ihluke kakhulu kwi-PC4, kunye enkulu 0.714, kwaye umahluko ukusuka PC3 buthathaka (AUC = 0.688).La maqela alandelayo achongiwe ngexabiso eliphantsi le-AUC kunye namanqanaba aphezulu okubaluleka: Iziphumo ze-PC7 (AUC = 0.679), i-PC4 (AUC = 0.654) kunye ne-PC1 (AUC = 0.649) ibonise ukuba amaNative Americans (Fig. 2f) aneenkcukacha ezithile. iimpawu ezinxulumene nala macandelo, ama-Southeast Asia (Umfanekiso 2g) ahluke kwi-PC3 (AUC = 0.660) kunye ne-PC9 (AUC = 0.663), kodwa iphethini yeesampuli ezivela kuMbindi Mpuma (umzobo 2h) (kuquka iNyakatho Afrika) ihambelana.Xa kuthelekiswa nabanye akukho mahluko ungako.
Kwinqanaba elilandelayo, ukutolika ngokubonakalayo ii-vertices ezinxibelelene kakhulu, iindawo zomhlaba ezinamaxabiso aphezulu omthwalo omkhulu kuno-0.45 zinemibala kunye noX, Y, kunye no-Z onxibelelanisa ulwazi, njengoko kubonisiwe kuMfanekiso 3. Indawo ebomvu ibonisa ulungelelwaniso oluphezulu Ulungelelwaniso lwe-X-axis, oluhambelana necala elinqamlezileyo elinqamlezileyo.Ummandla oluhlaza unxulunyaniswa kakhulu nolungelelwaniso oluthe nkqo lwe-Y axis, kwaye ummandla oluhlaza okwesibhakabhaka unxulunyaniswa kakhulu nolungelelwaniso lwe-sagittal lwe-axis ye-Z.Ummandla oluhlaza okwesibhakabhaka unxulunyaniswa nee-axes zolungelelwaniso lwe-Y kunye nee-axes zokulungelelanisa u-Z;ipinki - indawo exubileyo ehambelana ne-X kunye no-Z yokulungelelanisa;tyheli - indawo ehambelana ne-X kunye ne-Y yokulungelelanisa i-axes;Indawo emhlophe iqulathe i-X, Y kunye no-Z yolungelelwaniso oluhamba phambili olubonakalisiweyo.Ke ngoko, kulo mda wexabiso lomthwalo, iPC 1 idityaniswa kakhulu nomhlaba wonke wokakayi.Imilo yeskull ye-3 ye-SD yenyani kwelinye icala le-axis yecandelo nayo ibonisiwe kulo mzobo, kwaye imifanekiso ephothiweyo iboniswe kwiVidiyo eyoNgezelelweyo ye-S1 ukuze iqinisekise ngokubonakalayo ukuba i-PC1 iqulethe izinto zobukhulu bokakayi ngokubanzi.
Ukuhanjiswa okuphindaphindwayo kwamanqaku ePC1 (igophe elilungelelanisiweyo eliqhelekileyo), imephu yombala webala lokakayi idityaniswe kakhulu nee nkqo zePC1 (ingcaciso yemibala enxulumene nobukhulu bamacala achaseneyo ale axis yi-3 SD. Isikali yingqukuva eluhlaza enobubanzi ye 50 mm.
Umzobo wesi-3 ubonisa iploti yokusasazwa kwamaza (i-curve fit curve) yamanqaku e-PC1 ngamnye abalwe ngokwahlukeneyo kwiiyunithi zejografi ezisi-9.Ukongeza kuqikelelo lwegophe lwe-ROC (Umfanekiso 2), uqikelelo lwabantu base-Asiya baseMzantsi Afrika luye lwajika ngokuphawulekayo ukuya ekhohlo ngenxa yokuba iintloko zabo zincinci kunezo zamanye amaqela ommandla.Njengoko kuboniswe kwiThebhile 1, aba baseMzantsi Asia bamele amaqela eentlanga e-Indiya kuquka i-Andaman kunye ne-Nicobar Islands, iSri Lanka kunye ne-Bangladesh.
I-dimensional coefficient ifunyenwe kwi-PC1.Ukufunyaniswa kwemimandla enxibelelene kakhulu kunye neemilo ezibonakalayo kukhokelele ekucacisweni kweefom zemiba yamacandelo ngaphandle kwePC1;nangona kunjalo, izinto zobungakanani azisoloko zipheliswa ngokupheleleyo.Njengoko kuboniswe ngokuthelekisa i-ROC curves (Figure 2), i-PC2 kunye ne-PC4 zezona zicalucalulo, zilandelwa yi-PC6 kunye ne-PC7.I-PC3 kunye ne-PC9 zisebenza kakhulu ekwahluleni isampuli yabemi ngokweeyunithi zejografi.Ke, ezi zibini zecandelo lamazembe abonisa ngokucwangcisiweyo i-scatterplots yamanqaku ePC kunye nemiphezulu yombala enxibelelene kakhulu necandelo ngalinye, kunye nokuguqulwa kwemilo ebonakalayo enemilinganiselo yamacala achaseneyo we-3 SD (Imifanekiso 4, 5, 6).Ukugqunywa kwe-convex hull yeesampulu kwiyunithi nganye yejografi emelwe kwezi ploti imalunga nama-90%, nangona kukho iqondo elithile lokuxinana phakathi kwamaqela.Itheyibhile 3 inika inkcazo yecandelo ngalinye lePCA.
I-Scatterplots ye-PC2 kunye ne-PC4 amanqaku abantu abane-cranial abasuka kwiiyunithi ezilithoba zejografi (phezulu) kunye neeyunithi ezine zejografi (ezantsi), iziza zombala wokakayi womphezulu weenkqo ezihambelana kakhulu nePC nganye (ngokunxulumene no-X, Y, Z).Inkcazo yombala wee-axes: bona isicatshulwa), kunye nokuguqulwa kwefom ebonakalayo kumacala ahlukeneyo ala ma-axes yi-3 SD.Umlinganiselo yingqukuva eluhlaza kunye nobubanzi obuyi-50 mm.
I-Scatterplots ye-PC6 kunye ne-PC7 amanqaku abantu abane-cranial abasuka kwiiyunithi ezilithoba zejografi (phezulu) kunye neeyunithi ezimbini zejografi (ezantsi), i-cranial surface color plots for vertices ehambelana kakhulu ne-PC nganye (ngokunxulumene no-X, Y, Z).Inkcazo yombala wee-axes: bona isicatshulwa), kunye nokuguqulwa kwefom ebonakalayo kumacala ahlukeneyo ala ma-axes yi-3 SD.Umlinganiselo yingqukuva eluhlaza kunye nobubanzi obuyi-50 mm.
I-Scatterplots ye-PC3 kunye ne-PC9 amanqaku abantu abane-cranial abasuka kwiiyunithi ezilithoba zejografi (phezulu) kunye neeyunithi ezintathu zejografi (ezantsi), kunye nemibala yemibala yendawo yokakayi (enxulumene ne-X, Y, i-axes Z) yeenkqongo ezihambelana kakhulu nokutolikwa kombala kwePC nganye. : i-cm .okubhaliweyo), kunye nogudo lwemilo yenyani kumacala ahlukeneyo ezi axes ezinobukhulu be-3 SD.Umlinganiselo yingqukuva eluhlaza kunye nobubanzi obuyi-50 mm.
Kwigrafu ebonisa amanqaku e-PC2 kunye ne-PC4 (umzobo 4, iividiyo ezongezelelweyo ze-S2, i-S3 ebonisa imifanekiso ephosakeleyo), imephu yombala ongaphezulu iboniswa xa ixabiso lomthwalo libekwe phezulu kune-0.4, ephantsi kune-PC1 kuba Ixabiso lePC2 umthwalo uwonke ungaphantsi kwePC1.
Ukwandiswa kwe-lobes yangaphambili kunye ne-occipital kwicala le-sagittal ecaleni kwe-Z-axis (bhlowu emnyama) kunye ne-parietal lobe kwicala le-coronal (bomvu) ngopinki), i-Y-axis ye-occiput (eluhlaza) kunye ne-Z-axis ebunzi (blue blue).Le grafu ibonisa amanqaku abo bonke abantu kwihlabathi jikelele;nangona kunjalo, xa zonke iisampuli ezibandakanya inani elikhulu lamaqela ziboniswa kunye ngaxeshanye, ukutolikwa kweepateni zokuchithakala kunzima kakhulu ngenxa yobuninzi bokuxinana;ngoko ke, ukusuka kwiiyunithi ezine kuphela ezinkulu zejografi (okt, i-Afrika, i-Australasia-Melanesia, iYurophu, kunye noMntla-mpuma Asia), iisampulu zithe saa ngaphantsi kwegrafu nge-3 SD virtual deformation cranial ngaphakathi koluhlu lwamanqaku ePC.Kumzobo, i-PC2 kunye ne-PC4 ngamaqela amanqaku.Ama-Afrika kunye ne-Austro-Melanesians ziyadibana ngakumbi kwaye zisasazwe kwicala lasekunene, ngelixa ama-Europe esasazeke kwicala eliphezulu lasekhohlo kwaye ama-Asiya akuMntla-mpuma athande ukuhlangana ukuya ngasezantsi ngasekhohlo.I-axis ethe tyaba ye-PC2 ibonisa ukuba abantu base-Afrika/ base-Australia base-Melanesia bane-neurocranium ende kunabanye abantu.I-PC4, apho indibaniselwano yaseYurophu kunye nempuma-mpuma ye-Asia ihlukaniswe ngokukhululekileyo, idibaniswa nobukhulu obuhambelanayo kunye nokuqikelelwa kwamathambo e-zygomatic kunye ne-lateral contour ye-calvarium.Iskimu sokukora sibonisa ukuba abantu baseYurophu banamathambo amaxillary amancinci kunye ne-zygomatic, isithuba esincinci se-fossa esinqunyelwe yi-zygomatic arch, ithambo langaphambili eliphakamileyo kunye nethambo eliphantsi, eliphantsi, ngelixa i-Northeast Asia ithande ukuba namathambo e-zygomatic abanzi kwaye avelele ngakumbi. .I-lobe yangaphambili ithambekele, isiseko sethambo le-occipital siphakanyiswa.
Xa ugxininise kwi-PC6 kunye ne-PC7 (umzobo 5) (Iividiyo ezongezelelweyo ze-S4, i-S5 ebonisa imifanekiso ephosakeleyo), umbala webala ubonisa ixabiso lomthwalo wexabiso elikhulu kune-0.3, ebonisa ukuba i-PC6 idibene ne-maxillary okanye i-alveolar morphology (obomvu: i-X axis kunye luhlaza).I-axis ye-Y), imilo yethambo lexeshana (eluhlaza okwesibhakabhaka: i-Y kunye ne-Z axs) kunye ne-occipital bone shape (pinki: X kunye no-Z).Ukongeza kububanzi bebunzi (obomvu: i-X-axis), i-PC7 iphinda inxibelelane nobude be-alveoli ye-maxillary yangaphambili (eluhlaza: i-Y-axis) kunye ne-Z-axis yentloko imilo ejikeleze ingingqi ye-parietotemporal (blue blue).Kwiphaneli ephezulu yoMfanekiso 5, zonke iisampuli zejografi zisasazwa ngokwePC6 kunye nePC7 amanqaku ecandelo.Ngenxa yokuba i-ROC ibonisa ukuba i-PC6 iqulethe iimpawu ezizodwa eYurophu kwaye i-PC7 imele iimpawu zeMveli yaseMelika kolu hlalutyo, ezi sampuli zimbini zengingqi ziye zacwangciswa ngokukhethayo kwesi sibini seezembe zecandelo.Abantu bomthonyama baseMelika, nangona bebandakanyiwe ngokubanzi kwisampulu, bahlakazekile kwikona ephezulu ngasekhohlo;ngokuchaseneyo, iisampulu ezininzi zaseYurophu zikholisa ukuba zibekwe kwikona esezantsi ekunene.I-PC6 kunye ne-PC7 ezibini zimele inkqubo ye-alveolar emxinwa kunye ne-neurocranium ebanzi yaseYurophu, ngelixa abantu baseMelika bebonakala ngebunzi elimxinwa, i-maxilla enkulu, kunye nenkqubo ye-alveolar ebanzi kunye nede.
Uhlalutyo lwe-ROC lubonise ukuba i-PC3 kunye / okanye i-PC9 yayixhaphake kwi-Southeast and Northeast Asian populations.Ngokuhambelanayo, izibini zamanqaku e-PC3 (ubuso obuphezulu obuluhlaza kwi-y-axis) kunye ne-PC9 (ubuso obuhlaza obusezantsi kwi-y-axis) (Umfanekiso 6; Iividiyo ezongezelelweyo ze-S6, i-S7 zibonelela ngemifanekiso ye-morphed) zibonisa ukuhlukahluka kwe-East Asia., ephikisana ngokukhawuleza kunye nobukhulu obuphezulu bobuso baseMntla-mpuma ye-Asia kunye nesimo esisezantsi sobuso bama-Asia-mpuma ase-Asia.Ngaphandle kwezi mpawu zobuso, olunye uphawu lwabanye abantu baseMntla-mpuma Asia kukuthambeka kwe-lambda yethambo le-occipital, ngelixa abanye abantu base-Asia-mpuma ye-Asiya benesiseko sokakayi esimxinwa.
Le nkcazo ingentla yamacandelo aphambili kunye nenkcazo ye-PC5 kunye ne-PC8 ayifakwanga ngenxa yokuba akukho mpawu zikhethekileyo zengingqi zifunyenweyo phakathi kweeyunithi ezilithoba eziphambili zejografi.I-PC5 ibhekisa kubungakanani benkqubo ye-mastoid yethambo lexeshana, kwaye i-PC8 ibonisa i-asymmetry yesheyiphu ye-skull yonke, zombini ibonisa ukuhluka okuhambelanayo phakathi kwendibaniselwano yesampulu ezilithoba zejografi.
Ukongeza kwi-scatterplots yamanqaku e-PCA kwinqanaba lomntu ngamnye, sikwabonelela ngee-scatterplots zeendlela zeqela ukuthelekisa ngokubanzi.Ukuza kuthi ga ngoku, imodeli ye-cranial homology yenziwe ukusuka kwiseti yedatha ye-vertex yeemodeli zehomology nganye ukusuka kumaqela ezizwe ezili-148.Iiplani zeBivariate zeeseti zamanqaku e-PC2 kunye ne-PC4, i-PC6 kunye ne-PC7, kunye ne-PC3 kunye ne-PC9 ziboniswa kwi-Supplementary Figure S1, zonke zibalwe njengemodeli ye-skull eqhelekileyo kwisampulu yabantu abayi-148.Ngale ndlela, i-scatterplots ifihla ukungafani komntu ngaphakathi kweqela ngalinye, ivumela ukutolikwa okucacileyo kokufana kokakayi ngenxa yokusasazwa kommandla ophantsi, apho iipateni zihambelana nezo zichazwe kwiiplani zomntu ngamnye kunye nokudibana okuncinci.Umzobo owoNgezelelweyo S2 ubonisa imodeli yentsingiselo iyonke kwiyunithi nganye yejografi.
Ukongeza kwi-PC1, eyayanyaniswa nobukhulu obupheleleyo (i-Supplementary Table S2), ubudlelwane be-allometric phakathi kobukhulu obupheleleyo kunye nokuma kwe-skull bahlolwe kusetyenziswa imilinganiselo ye-centroid kunye neeseti ze-PCA eziqikelelwayo ezivela kwiidatha ezingezona eziqhelekileyo.I-Allometric coefficients, amaxabiso angaguqukiyo, amaxabiso e-t, kunye namaxabiso e-P kuvavanyo lokubaluleka abonisiwe kwiThebhile 4. Akukho malungu abalulekileyo epateni yeallometric adityaniswe nobungakanani bentloko yonke efunyenwe kuyo nayiphi na i-cranial morphology kwinqanaba le-P <0.05.
Ngenxa yokuba ezinye izinto zobungakanani zinokuqukwa kuqikelelo lwe-PC olusekwe kwiiseti zedatha ezingaqhelekiyo, siye saphinda savavanya intsingiselo ye-allometric phakathi kobungakanani becentroid kunye namanqaku ePC abalwa kusetyenziswa iiseti zedatha eziqhelekileyo ngokwesayizi ye-centroid (iziphumo zePCA kunye neeseti zamanqaku zithiwe thaca kwiiTheyibhuli ezongezelelweyo ze-S6. ).,C7).Itheyibhile 4 ibonisa iziphumo zohlalutyo lwe-allometric.Ngaloo ndlela, iindlela ezibalulekileyo ze-alometric zifunyenwe kwinqanaba le-1% kwi-PC6 kunye ne-5% kwinqanaba le-PC10.Umzobo 7 ubonisa ukuhlehla kwethambeka kolu budlelwane belog-linear phakathi kwamanqaku ePC kunye nobungakanani becentroid enedummies (± 3 SD) nakwesiphi na isiphelo selog yesayizi ye-centroid.Amanqaku ePC6 ngumlinganiselo wobude obunxulumene nobubanzi bokakayi.Njengoko ubungakanani bentloko bukhula, ukhakhayi kunye nobuso buba phezulu, kwaye ibunzi, iisokethi zamehlo kunye neempumlo zivame ukusondelana ecaleni.Ipateni yokusasazwa kwesampulu icebisa ukuba lo mlinganiso udla ngokufumaneka kumaMntla-mpuma aseAsia kunye nabemi bomthonyama baseMelika.Ngaphaya koko, i-PC10 ibonisa intsingiselo ebhekiselele ekuncitshisweni okulinganayo kububanzi obuphakathi kungakhathaliseki ukuba ummandla wejografi.
Kubudlelwane obubalulekileyo be-allometric obudweliswe kwitheyibhile, ithambeka lokuhlehla komgca welog phakathi komlinganiselo wePC wecandelo lemilo (efunyenwe kwidatha eqhelekileyo) kunye nobukhulu becentroid, ukuguqulwa kwemilo ebonakalayo kunobungakanani be-3 SD icala elichaseneyo lomgca wesi-4.
Le pateni ilandelayo yotshintsho kwi-cranial morphology ibonakaliswe ngohlalutyo lweeseti zedatha yeemodeli zomgangatho we-3D we-homologous.Icandelo lokuqala le-PCA linxulumene nobukhulu bokakayi ngokubanzi.Sekude kucingelwa ukuba ama-skulls amancinci aseMzantsi Asia, kuquka imizekelo evela eIndiya, eSri Lanka kunye neziqithi zaseAndaman, eBangladesh, ngenxa yobukhulu bomzimba wabo omncinci, ngokuhambelana nomthetho we-ecogeographic kaBergmann okanye isiqithi 613,5,16,25, 27,62 .Eyokuqala inxulumene neqondo lokushisa, kwaye okwesibini kuxhomekeke kwindawo ekhoyo kunye nemithombo yokutya ye-ecological niche.Phakathi kwamacandelo emilo, olona tshintsho lukhulu lumlinganiselo wobude kunye nobubanzi be-cranial vault.Eli nqaku, elichongiweyo i-PC2, lichaza unxulumano olusondeleyo phakathi kokhakhayi olude ngokulinganayo lwe-Austro-Melanesians kunye nama-Afrika, kunye nomahluko kukhakhayi olungqukuva lwabanye abantu baseYurophu nabaseMntla-mpuma Asia.Ezi mpawu zichazwe kwizifundo ezininzi zangaphambili ezisekelwe kwimilinganiselo elula yomgca37,63,64.Ngaphezu koko, olu phawu ludibene ne-brachycephaly kubantu abangewona ama-Afrika, ekudala kuxoxwa ngayo kwizifundo ze-anthropometric kunye ne-osteometric.I-hypothesis ephambili emva kwale nkcazo kukuba ukunciphisa i-mastication, njengokunciphisa i-muscle ye-temporalis, kunciphisa uxinzelelo kwi-scalp yangaphandle5,8,9,10,11,12,13.Enye i-hypothesis ibandakanya ukulungelelaniswa kwemozulu ebandayo ngokunciphisa indawo yentloko, iphakamisa ukuba i-skull engaphezulu ye-spherical inciphisa indawo engcono kunokwakheka kwe-spherical, ngokwemigaqo ka-Allen16,17,25.Ngokusekelwe kwiziphumo zophando lwangoku, ezi ngcamango zinokuvavanywa kuphela ngokusekelwe kwi-cross-correlation yamacandelo e-cranial.Isishwankathelo, iziphumo zethu ze-PCA aziyixhasi ngokupheleleyo i-hypothesis yokuba umlinganiselo we-cranial ubude-width ratio uphenjelelwa kakhulu ziimeko zokuhlafuna, njengoko i-PC2 (inxalenye ende / ye-brachycephalic) ukulayishwa kwakungahambelani kakhulu nobungakanani bobuso (kubandakanywa nobukhulu be-maxillary dimensions).kunye nesithuba esihambelanayo se-fossa yesikhashana (ebonisa umthamo we-temporalis muscle).Uphononongo lwethu lwangoku aluzange luhlalutye ubudlelwane phakathi kobume bekhanda kunye neemeko zendalo zendalo ezifana nobushushu;nangona kunjalo, inkcazo esekelwe kumgaqo ka-Allen inokufaneleka ukuba ithathelwe ingqalelo njengomviwa we-hypothesis ukuchaza i-brachycephalon kwimimandla yemozulu ebandayo.
Ukwahluka okuphawulekayo kwafunyanwa ke kwi-PC4, ebonisa ukuba i-Northeast Asia inamathambo amakhulu, avelele amathambo e-zygomatic kwi-maxilla kunye namathambo e-zygomatic.Oku kufunyaniswayo kuhambelana neempawu ezicacileyo ezaziwayo zaseSiberia, ezicatshangelwa ukuba zilungelelanise kwiindawo ezibandayo kakhulu ngokuhamba phambili kwamathambo e-zygomatic, okubangelwa ukwanda kwenani lezono kunye nobuso obuthambileyo be65.Uphando olutsha oluvela kwimodeli yethu ye-homologous kukuba i-cheek drooping kubantu baseYurophu inxulunyaniswa ne-slope yangaphambili encitshisiweyo, kunye namathambo e-occipital athambileyo kunye ne-nuchal concavity.Ngokwahlukileyo koko, abantu baseMntla-mpuma Asia badla ngokuba namabunzi athambekileyo kwaye baphakamise imimandla ye-occipital.Izifundo zethambo le-occipital usebenzisa iindlela zejometri ze-morphometric35 zibonise ukuba i-skulls yase-Asia kunye neYurophu ine-curve nuchal curve kunye nesikhundla esisezantsi se-occiput xa kuthelekiswa nama-Afrika.Nangona kunjalo, i-scatterplots yethu ye-PC2 kunye ne-PC4 kunye ne-PC3 kunye ne-PC9 ngababini babonise ukuhluka okukhulu kuma-Asiya, ngelixa amaYurophu ayebonakaliswe ngesiseko esisicaba se-occiput kunye ne-occiput ephantsi.Ukungangqinelani kwiimpawu zase-Asiya phakathi kwezifundo kusenokuba kungenxa yomahluko kwiisampulu zobuhlanga ezisetyenzisiweyo, njengoko siye sathatha iisampulu ezininzi zamaqela ezizwe ezisuka kuluhlu olubanzi lwaseMntla-mpuma nakuMzantsi-mpuma Asia.Utshintsho kwimilo yethambo le-occipital lihlala lihambelana nokuphuhliswa kwemisipha.Nangona kunjalo, le nkcazo eguquguqukayo ayibandakanyi ukulungelelaniswa phakathi kwebunzi kunye ne-occiput shape, eboniswe kolu cwaningo kodwa akunakwenzeka ukuba iboniswe ngokupheleleyo.Kule nkalo, kuyafaneleka ukuqwalasela ubudlelwane phakathi kokulinganisela kobunzima bomzimba kunye neziko lomxhuzulane okanye ukuhlangana komlomo wesibeleko (foramen magnum) okanye ezinye izinto.
Elinye icandelo elibalulekileyo elinoguquko olukhulu linxulumene nokuphuhliswa kwezixhobo zokuphulula, ezimelwe yi-fossae ye-maxillary kunye neyexeshana, echazwe ngokudibanisa amanqaku e-PC6, i-PC7 kunye ne-PC4.Oku kuncitshiswa kuphawulwe kumacandelo ecranial kuphawula abantu baseYurophu ngaphezu kwalo naliphi na elinye iqela lejografi.Olu phawu luye lwatolikwa ngenxa yokuncipha kozinzo kwi-morphology yobuso ngenxa yokuphuhliswa kwangaphambili kwezolimo kunye neendlela zokulungiselela ukutya, okwathi kwanciphisa umthwalo womatshini kwi-masticatory apparatus ngaphandle kwesixhobo esinamandla se-masticatory9,12,28,66.Ngokutsho kwe-masticatory function hypothesis, 28 oku kukhatshwa yinguqu kwisiseko sokakayi ukuya kwi-angle ebukhali yecranial kunye nophahla olungqukuva ngakumbi.Ngokwale mbono, abemi bezolimo bakholisa ukuba nobuso obubambeneyo, ukuphuma kancinci kwe-mandible, kunye ne-globular meninges.Ngoko ke, le deformation inokuchazwa ngoluhlu olubanzi lwemilo esecaleni yokhakhayi lwaseYurophu kunye nezitho ezincitshisiweyo zokukrala.Nangona kunjalo, ngokutsho kolu phononongo, le ngcaciso iyinkimbinkimbi kuba ukubaluleka kokusebenza kobudlelwane be-morphological phakathi kwe-globose neurocranium kunye nokuphuhliswa kwezixhobo zokuphulukisa akwamkelekanga, njengoko kuqwalaselwe kwiinkcazo zangaphambili ze-PC2.
Umahluko phakathi kwama-Asiya aseMntla-mpuma kunye nama-Asia-mpuma ase-Asia aboniswa ngumahluko phakathi kobuso obude kunye nethambo le-occipital elithambekileyo kunye nobuso obufutshane kunye nesiseko sokakayi olumxinwa, njengoko kubonisiwe kwi-PC3 kunye ne-PC9.Ngenxa yokunqongophala kwedatha ye-geoecological, uphononongo lwethu lubonelela kuphela ngengcaciso elinganiselweyo yesi siphumo.Inkcazo enokwenzeka kukuziqhelanisa nemozulu eyahlukileyo okanye iimeko zesondlo.Ukongeza kwi-ecological adaptation, iyantlukwano yendawo kwimbali yabantu baseMntla-mpuma kunye ne-Southeast Asia nayo yathathelwa ingqalelo.Ngokomzekelo, kwimpuma ye-Eurasia, imodeli yemigangatho emibini iye yacatshangelwa ukuqonda ukusasazeka kwabantu be-anatomically namhlanje (AMH) ngokusekelwe kwidatha ye-cranial morphometric data67,68.Ngokwalo mzekelo, "inqanaba lokuqala", oko kukuthi, amaqela okuqala e-Late Pleistocene AMH colonizers, ayenenzala ethe ngqo okanye engaphantsi ukusuka kubemi bomthonyama bommandla, njengama-Austro-Melanesians anamhlanje (iphe. Okokuqala stratum)., kwaye kamva bafumana umxube omkhulu wabantu bezolimo basemantla kunye neempawu zaseMntla-mpuma Asia (umaleko wesibini) kummandla (malunga ne-4,000 yeminyaka eyadlulayo).Ukuhamba kweGene okufakwe kwimaphu kusetyenziswa imodeli “yomaleko-mbini” kuya kufuneka ukuze kuqondwe imilo yecranial yaseMazantsi-mpuma yeAsia, njengoko imilo yecranial yaseMazantsi-mpuma yeAsia inokuxhomekeka ngokuyinxenye kwilifa lemveli lendawo yokuqala.
Ngokuvavanya ukufana kwecranial kusetyenziswa iiyunithi zejografi ezizotywe kusetyenziswa imifuziselo efanayo, sinokuthelekelela isiseko sembali yabemi ye-AMF kwiimeko ezingaphandle kwe-Afrika.Iimodeli ezininzi ezahlukeneyo "eziphuma e-Afrika" ziye zacetywa ukuba zichaze ukuhanjiswa kwe-AMF ngokusekelwe kwidatha ye-skeletal kunye ne-genomic.Kwezi, uphando olutshanje lubonisa ukuba i-AMH ye-colonization yeendawo ezingaphandle kwe-Afrika yaqala malunga ne-177,000 kwiminyaka edlulileyo69,70.Nangona kunjalo, ukuhanjiswa komgama omde we-AMF e-Eurasia ngeli xesha akuqinisekanga, kuba iindawo zokuhlala zala ma-fossils okuqala zilinganiselwe kuMbindi Mpuma kunye neMeditera kufuphi neAfrika.Esona siganeko silula kukuhlala kwindawo enye ecaleni kwendlela yokufuduka isuka eAfrika isiya e-Eurasia, idlula imiqobo yejografi efana neeHimalaya.Enye imodeli icebisa amaza amaninzi okufuduka, eyokuqala eyasasazeka ukusuka e-Afrika ecaleni konxweme loLwandlekazi lwaseIndiya ukuya kumazantsi-mpuma e-Asia nase-Australia, emva koko yasasazeka kumantla e-Eurasia.Uninzi lwezi zifundo ziqinisekisa ukuba i-AMF yanwenwela ngaphaya kweAfrika kwiminyaka engama-60,000 eyadlulayo.Ngokubhekiselele koku, iisampulu zeAustralasian-Melanesia (kubandakanywa iPapua) zibonisa ukufana okukhulu kwiisampulu zaseAfrika kunalo naluphi na olunye uthotho lwejografi kucazululo lwamacandelo aphambili kwiimodeli zehomology.Oku kufunyaniswayo kuxhasa i-hypothesis yokuba amaqela okuqala okusasazwa kwe-AMF ecaleni komda osezantsi we-Eurasia avela ngokuthe ngqo kwi-Afrika22,68 ngaphandle kotshintsho olubalulekileyo lwe-morphological ekuphenduleni kwimozulu ethile okanye ezinye iimeko ezibalulekileyo.
Ngokumalunga nokukhula kwe-allometric, uhlalutyo kusetyenziswa amacandelo emilo athathwe kwiseti yedatha eyahlukileyo eqheleke ngokwesayizi ye-centroid ibonise umzila obalulekileyo we-allometric kwi-PC6 kunye ne-PC10.Omabini amacandelo anxulumene nokumila kwebunzi kunye neenxalenye zobuso, eziye zibe zincinci njengoko ubungakanani bentloko bukhula.Abantu baseMntla-mpuma ye-Asiya kunye naseMelika badla ngokuba nalo olu phawu kwaye banokakayi obukhulu.Oku kufunyanisiweyo kuyaphikisana neepateni ze-allometric ezixeliweyo ngaphambili apho ubuchopho obukhulu bunelobes zangaphambili eziphangaleleyo kwindawo ebizwa ngokuba yi "Broca's cap", okukhokelela ekwandeni kobubanzi belobe yangaphambili34.Lo mahluko uchazwa ngomahluko kwiiseti zesampulu;Uphononongo lwethu luhlalutye iipateni ze-allometric zobukhulu be-cranial ngokubanzi kusetyenziswa abantu banamhlanje, kunye nezifundo ezithelekisayo zijongana nentsingiselo yexesha elide kwindaleko yomntu ehambelana nobungakanani bobuchopho.
Ngokumalunga ne-allometry yobuso, olunye uphononongo olusebenzisa idatha yebhayometriki78 lufumanise ukuba imilo yobuso kunye nobukhulu bunokuthi bunxibelelene kancinane, kanti uphononongo lwethu lufumanise ukuba ukhakhayi olukhulu ludla ngokunxulunyaniswa nobuso obude, obumxinwa.Nangona kunjalo, ukuhambelana kwedatha ye-biometric ayicacanga;Iimvavanyo zokubuyisela umva ngokuthelekisa i-ontogenetic allometry kunye ne-static allometry ibonisa iziphumo ezahlukeneyo.Utyekelo lwe-allometric olubhekiselele kwimilo yokakayi obungqukuva ngenxa yokuphakama okwandisiweyo nayo ixeliwe;nangona kunjalo, asizange sihlalutye idatha yobude.Uphononongo lwethu lubonisa ukuba akukho datha ye-allometric ebonisa ulungelelwaniso phakathi kobungakanani beglobula ye-cranial kunye nobukhulu be-cranial jikelele ngeseshini nganye.
Nangona uphononongo lwethu lwangoku alujongani nedatha kwizinto eziguquguqukayo zangaphandle ezimelwe yimozulu okanye iimeko zokutya ezinokuthi ziphembelele i-cranial morphology, isethi enkulu yedatha yeemodeli ze-homologous ze-3D ze-cranial surface ezisetyenziswe kolu phononongo ziya kunceda ukuvavanya i-phenotypic morphological variation ehambelanayo.Izinto zokusingqongileyo ezifana nokutya, imozulu kunye neemeko zesondlo, kunye namandla angathathi hlangothi afana nokufuduka, ukuhamba kofuzo kunye nokuhamba kwemfuzo.
Olu phononongo lubandakanya imizekelo ye-342 yeekhakhayi zamadoda eziqokelelwe kubantu be-148 kwiiyunithi zejografi ze-9 (Itheyibhile 1).Amaqela amaninzi ayimizekelo yezelizwe, ngelixa amanye amaqela e-Afrika, uMntla-mpuma/uMzantsi-mpuma we-Asia kunye naseMelika (adweliswe ngoonobumba abakekeleyo) echazwa ngokobuhlanga.Iisampulu ezininzi zecranial ziye zakhethwa kuvimba womlinganiselo wecranial ngokwenkcazo yomlinganiselo we Martin cranial enikezwe ngu Tsunehiko Hanihara.Sikhethe ummeli wamadoda okakayi kuzo zonke izizwe zehlabathi.Ukuchonga amalungu eqela ngalinye, sibale imigama ye-Euclidean ngokusekwe kwimilinganiselo engama-37 yecranial esuka kwiqela ithetha kubo bonke abantu abakwelo qela.Kwiimeko ezininzi, sikhethe iisampuli ze-1-4 kunye nomgama omncinci ukusuka kwintsingiselo (iTheyibhile eyoNgezelelweyo ye-S4).Kula maqela, ezinye iisampulu zikhethwe ngokungenamkhethe ukuba azibhalwanga kwi-database yokulinganisa iHahara.
Uthelekiso lweenkcukacha-manani, iisampulu zabemi abali-148 zaye zadityaniswa zaba ziiyunithi ezinkulu zejografi, njengoko kubonisiwe kwiThebhile 1. Iqela “labaseAfrika” libandakanya kuphela iisampulu ezisuka kummandla we-sub-Sahara.Iisampuli ezivela kuMntla Afrika zifakwe "kuMbindi Mpuma" kunye neesampuli ezivela eNtshona Asia ezineemeko ezifanayo.Iqela laseMntla-mpuma Asia libandakanya kuphela abantu abangengowaseYurophu, kwaye iqela laseMelika libandakanya kuphela amaMerika Omthonyama.Ngokukodwa, eli qela lisasazwe kwindawo enkulu yamazwekazi aseMntla naseMzantsi Melika, kwiindawo ngeendawo ezahlukeneyo.Nangona kunjalo, siqwalasela isampula yase-US ngaphakathi kweli candelo elilodwa lejografi, linikezelwa kwimbali yabantu baseMerika abathathwa njengemvelaphi yaseMntla-mpuma ye-Asia, kungakhathaliseki ukufuduka okuphindaphindiweyo kwe80.
Sirekhode idatha yomphezulu we-3D yale mizekelo yokakayi yahlukileyo sisebenzisa iskena se-3D esinesisombululo esiphezulu (EinScan Pro by Shining 3D Co Ltd, isisombululo esincinci: 0.5 mm, https://www.shining3d.com/) emva koko senza i-mesh.Imodeli yemesh iqulathe malunga ne-200,000-400,000 yee-vertices, kwaye isoftware ebandakanyiweyo isetyenziselwa ukugcwalisa imingxuma kunye nemiphetho egudileyo.
Kwinqanaba lokuqala, sisebenzise idatha yokuskena kulo naliphi na ikhakhayi ukwenza imodeli yokhakhayi yemesh enetemplate enye equka i4485 vertices (8728 face polygonal faces).Isiseko sommandla wokakayi, obandakanya ithambo le-sphenoid, ithambo lexeshana le-petrous, inkalakahla, i-maxillary alveoli, namazinyo, yasuswa kwimodeli yemesh yetemplate.Isizathu kukuba ezi zakhiwo ngamanye amaxesha aziphelelanga okanye kunzima ukuzigqiba ngenxa yamacandelo amancinci okanye amancinci abukhali afana nomphezulu we-pterygoid kunye neenkqubo ze-styloid, ukuguga kwamazinyo kunye / okanye isethi engaguqukiyo yamazinyo.Isiseko seskull esijikeleze i-foramen magnum, kubandakanywa nesiseko, asizange siphinde sihlaziywe ngenxa yokuba le yindawo ebalulekileyo ye-anatomically yendawo yamalungu omlomo wesibeleko kunye nokuphakama kokakayi kufuneka kuhlolwe.Sebenzisa amakhonkco esipili ukwenza itemplate elinganayo kumacala omabini.Yenza i-isotropic meshing ukuguqula iimilo zepolygonal zibe zilinganayo kangangoko.
Emva koko, ii-landmarks ze-56 zabelwa kwii-vertices ezihambelana ne-anatomically ye-template model usebenzisa i-software ye-HBM-Rugle.Iisetingi zeempawu zomhlaba ziqinisekisa ukuchaneka kunye nokuzinza kokuma kophawu kunye nokuqinisekisa i-homology yezi ndawo kwimodeli ye-homology eyenziwe.Ziyakwazi ukuchongwa ngokusekelwe kwiimpawu zabo ezithile, njengoko kuboniswe kwiTheyibhile eyoNgezelelweyo ye-S5 kunye ne-Supplementary Figure S3.Ngokwenkcazelo ka-Bookstein81, uninzi lwala maphawu omhlaba ziimpawu zomhlaba zoHlobo lwe-I ezibekwe ekudibaneni kwezakhiwo ezithathu, kwaye ezinye ziimpawu zomhlaba zoHlobo lwe-II ezineendawo zokugoba okuphezulu.Iimpawu zomhlaba ezininzi ziye zakhutshelwa ukusuka kumanqaku achazwe kwimilinganiselo yomgca yecranial kwinkcazelo kaMartin 36. Sichaze iimpawu zomhlaba ezifanayo ezingama-56 zemizekelo eskeniweyo yesampulu zokakayi ezingama-342, ezabelwa ngesandla kwiinkqo ezihambelana ne-anatomically ukuvelisa imodeli yehomology echanekileyo kwicandelo elilandelayo.
Inkqubo yokulungelelanisa intloko-centric ichazwe ukuchaza idatha yokuskena kunye nethemplate, njengoko kuboniswe kwi-Supplementary Figure S4.Inqwelomoya ye-XZ yinqwelomoya ethe tyaba yaseFrankfurt edlula kwindawo ephezulu (inkcazo kaMartin: inxalenye) yomgca ophezulu wecala lasekhohlo nasekunene langaphandle lokuphicotha kunye neyona ndawo iphantsi (inkcazo kaMartin: i-orbit) yecala elisezantsi le-orbit yasekhohlo. ..Umgca we-X ngumgca odibanisa amacala asekhohlo nasekunene, kwaye u-X+ licala lasekunene.Inqwelomoya ye-YZ idlula phakathi kwendawo ekhohlo nasekunene kunye neengcambu zempumlo: Y + phezulu, Z + phambili.Inqaku lesalathiso (imvelaphi: ulungelelwaniso lwe-zero) lusekwe kwi-intersection ye-YZ plane (i-midplane), indiza ye-XZ (indiza yaseFrankfort) kunye ne-XY (indiza ye-coronal).
Sasebenzisa isoftware ye-HBM-Rugle (i-Medic Engineering, Kyoto, http://www.rugle.co.jp/) ukwenza imodeli ye-mesh ye-homologous ngokwenza i-template fitting usebenzisa i-56 yendawo ephawulekayo (icala lasekhohlo loMzobo 1).Elona candelo lesoftware, elaphuhliswa kuqala liZiko loPhando lweDijithali loLuntu kwiZiko leSayensi yoShishino oluPhezulu kunye neTekhnoloji eJapan, libizwa ngokuba yi-HBM kwaye linemisebenzi yokulungelelanisa iitemplates kusetyenziswa iimpawu zomhlaba kunye nokudala imifuziselo emihle yemesh usebenzisa umphezulu wokwahlula82.Inguqulelo yesoftware elandelayo (mHBM) 83 yongeze inqaku lokulinganisa ipateni ngaphandle kweempawu zomhlaba ukuphucula ukusebenza okufanelekileyo.I-HBM-Rugle idibanisa isoftware ye-mHBM kunye neempawu ezongezelelweyo ezisebenzisekayo ezibandakanya ukulungelelanisa iinkqubo kunye nokutshintsha ubungakanani bedatha yegalelo.Ukuthembeka kokuchaneka kokufakwa kwesoftware kuye kwaqinisekiswa kwizifundo ezininzi52,54,55,56,57,58,59,60.
Xa ufaka i-template ye-HBM-Rugle usebenzisa i-landmarks, imodeli ye-mesh ye-template ibekwe phezulu kwidatha yokuskena ekujoliswe kuyo ngobhaliso oluqinileyo olusekelwe kwi-teknoloji ye-ICP (ukunciphisa inani lemigama phakathi kweempawu zomhlaba ezihambelana netemplate kunye nedatha yokuskena ekujoliswe kuyo), kunye ke ngoguquko olungeluqilima lomnatha lulungelelanisa itemplate kwidatha yokuskena ekujoliswe kuyo.Le nkqubo yokufaneleka iphindwe kathathu kusetyenziswa amaxabiso ahlukeneyo eeparamitha ezifanelekileyo zokuphucula ukuchaneka kokufaneleka.Enye yezi parameters inciphisa umgama phakathi kwemodeli yegridi yetemplate kunye nedatha yokuskena ekujoliswe kuyo, kwaye enye ihlawulisa umgama phakathi kweempawu zomhlaba zetemplate kunye neendawo ekujoliswe kuzo.Imodeli yemesh yethempleyithi eyonakeleyo yahlulwa kusetyenziswa i-cyclic surface subdivision algorithm 82 ukwenza imodeli yemesh esulungekisiweyo equka i-17,709 vertices (34,928 polygons).Okokugqibela, imodeli yegridi yetemplate eyahluliweyo ifanelekile kwidatha yokuskena ekujoliswe kuyo ukuvelisa imodeli yehomology.Ekubeni iindawo eziphawulweyo zahluke kancinane kwezo zikwidatha yokuskena ekujoliswe kuyo, imodeli ye-homology yalungiswa kakuhle ukuzichaza kusetyenziswa inkqubo yolungelelwaniso lwentloko echazwe kwicandelo langaphambili.Umyinge womgama phakathi kweempawu zomhlaba zemodeli ehambelanayo kunye nedatha yokuskena ekujoliswe kuyo kuzo zonke iisampuli yayingu-<0.01 mm.Ukubalwa kusetyenziswa umsebenzi we-HBM-Rugle, umgama ophakathi phakathi kwamanqaku edatha yemodeli ye-homology kunye nedatha yokuskena ekujoliswe kuyo yi-0.322 mm (i-Supplementary Table S2).
Ukucacisa utshintsho kwi-cranial morphology, i-17,709 vertices (53,127 XYZ coordinates) yazo zonke iimodeli ze-homologous zahlalutywa nguhlalutyo lwecandelo eliphambili (PCA) lisebenzisa isoftware ye-HBS eyenziwe yiZiko leSayensi yoLuntu kwiZiko leSayensi yezoShishino oluPhezulu kunye neTekhnoloji., Japan (umthengisi wokusasaza: Ubunjineli beMedic, Kyoto, http://www.rugle.co.jp/).Siye sazama ukusebenzisa i-PCA kwiseti yedatha engaqhelekanga kunye neseti yedatha iqhelekile ngokwesayizi ye-centroid.Ngaloo ndlela, i-PCA esekelwe kwidatha engabonakaliyo ingabonakalisa ngokucacileyo imilo ye-cranial yeeyunithi ezilithoba zejografi kwaye iququzelele ukutolika kwecandelo kune-PCA usebenzisa idatha esemgangathweni.
Eli nqaku libonisa inani lamacandelo enqununu afunyenweyo kunye negalelo elingaphezu kwe-1% ye-variance iyonke.Ukumisela amacandelo aphambili asebenzayo kumaqela ahlukeneyo kwiiyunithi ezinkulu zejografi, uhlalutyo lwe-receiver operating character (ROC) lusetyenziswe kumanqaku enqununu (PC) kunye negalelo elikhulu kune-2% 84.Olu hlalutyo luvelisa i-curve enokwenzeka kwicandelo ngalinye le-PCA ukuphucula ukusebenza kokuhlelwa kunye nokuthelekisa ngokuchanekileyo izicwangciso phakathi kwamaqela ejografi.Umgangatho wamandla ocalucalulo unokuvavanywa ngummandla ophantsi kwegophe (AUC), apho amacandelo e-PCA anamaxabiso amakhulu akwazi ngcono ukucalula phakathi kwamaqela.Uvavanyo lwe-chi-square ke lwenziwa ukuvavanya umgangatho wokubaluleka.Uhlalutyo lwe-ROC lwenziwa kwiMicrosoft Excel kusetyenziswa iBell Curve yesoftware ye-Excel (uguqulelo 3.21).
Ukubona umahluko wejografi kwi-cranial morphology, ii-scatterplots zenziwe kusetyenziswa amanqaku ePC awahlula ngokufanelekileyo amaqela asuka kwiiyunithi ezinkulu zejografi.Ukutolika amacandelo aphambili, sebenzisa imephu yombala ukubona imodeli yeevertices ezinxibelelene kakhulu nezinto eziphambili.Ukongeza, ukuboniswa okubonakalayo kweziphelo zee-axes zecandelo eliphambili eziku-±3 ezitenxileyo eziqhelekileyo (SD) zamanqaku ecandelo eliphambili zibalwe zaza zaboniswa kwividiyo eyongezelelweyo.
I-Allometry yayisetyenziselwa ukumisela ubudlelwane phakathi kobume bekhanda kunye nobukhulu bezinto ezihlolwe kuhlalutyo lwe-PCA.Uhlalutyo luyasebenza kumacandelo aphambili kunye namagalelo> 1%.Unyino omnye wale PCA kukuba iinxalenye zeemilo azikwazi ukubonisa imilo ngokuzimeleyo kuba iseti yedatha engeyiyo eqhelekileyo ayisusi zonke izinto ezinobungakanani.Ukongeza ekusebenziseni iiseti zedatha ezingaqhelekanga, siye sahlalutya iintsingiselo ze-allometric sisebenzisa iiseti zamaqhezu e-PC ngokusekwe kwidatha yesiqhelo ye-centroid yesayizi esetyenziswa kumacandelo aphambili ngegalelo> 1%.
Iintsingiselo ze-Allometric zavavanywa kusetyenziswa i-equation Y = aXb 85 apho u-Y iyimilo okanye umlinganiselo wecandelo lemilo, u-X ubungakanani becentroid (Itheyibhuli eyoNgezelelweyo ye-S2), a lixabiso elingaguqukiyo, kwaye u-b yi-alometric coefficient.Le ndlela ngokusisiseko yazisa izifundo zokukhula kwe-allometric kwimophometry yejometri78,86.Inguqu yelogarithmic yale fomula yile: log Y = b × log X + log a.Uhlalutyo lokuhlehla kusetyenziswa indlela yesikwere esincinane esetyenziswe ukubala u-a kunye no-b.Xa u-Y (ubukhulu obuphakathi) kunye no-X (amanqaku ePC) ziguqulwa ngokwe-logarithmically, la maxabiso kufuneka abe positive;nangona kunjalo, uluhlu loqikelelo lwe X luqulathe amaxabiso angalunganga.Njengesisombululo, songeze ukurhangqa kwixabiso elipheleleyo lelona qhezu lincinci kunye no-1 kwiqhezu ngalinye kwicandelo ngalinye kwaye sisebenzise inguqu yelogarithmic kuwo onke amaqhezu aguquliweyo alungileyo.Ukubaluleka kwee-alometric coefficients kwavavanywa kusetyenziswa uvavanyo lwe-t olunemisila emibini yoMfundi.Ezi zibalo zokubala zokuvavanya ukukhula kwe-allometric zenziwa kusetyenziswa iBell Curves kwisoftware ye-Excel (uguqulelo 3.21).
Wolpoff, MH Iziphumo zemozulu kwiimpumlo ze-skeleton.Ewe.J. Phys.Ubuntu.29, 405–423.https://doi.org/10.1002/ajpa.1330290315 (1968).
Iibhili, imilo yeNtloko ye-KL kunye noxinzelelo lwemozulu.Ewe.J. Phys.Ubuntu.37, 85–92.https://doi.org/10.1002/ajpa.1330370111 (1972).
Ixesha lokuposa: Apr-02-2024