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Amaphethini omhlaba achaza i-morphology yogebhezi lwekhanda lomuntu ngokuhlaziywa kwemodeli ye-homology yendawo enezinhlangothi ezintathu.

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Lolu cwaningo luhlole ukuhlukahluka kwesifunda ku-cranial morphology yomuntu kusetshenziswa imodeli ye-geometric homology esekelwe kudatha yokuskena evela kumaqembu ezizwe ayi-148 emhlabeni jikelele.Le ndlela isebenzisa ubuchwepheshe bokufaka isifanekiso ukuze kukhiqizwe ama-meshes afanayo ngokwenza uguquko olungaqinile kusetshenziswa i-algorithm yephoyinti ephindaphindayo.Ngokusebenzisa ukuhlaziya kwengxenye eyinhloko kumamodeli akhethiwe angama-homologous angu-342, ushintsho olukhulu kunawo wonke kusayizi lutholakele futhi lwaqinisekiswa ngokucacile kugebhezi oluncane lwekhanda oluvela eNingizimu Asia.Umehluko wesibili ngobukhulu isilinganiso sobude nobubanzi be-neurocranium, okubonisa umehluko phakathi kwezingebhezi ezinde zabantu base-Afrika kanye nezingebhezi zekhanda eziqondile zaseNyakatho-mpumalanga ye-Asia.Kuyaphawuleka ukuthi lesi sithako asihlobene kancane nokujika kobuso.Izici zobuso ezaziwayo njengezihlathi eziphumile eNyakatho-mpumalanga ye-Asia kanye namathambo ama-maxillary ahlangene kwabase-Europe aphinde aqinisekiswa.Lezi zinguquko zobuso zihlobene eduze ne-contour yogebhezi, ikakhulukazi izinga lokuthambekela kwamathambo angaphambili kanye ne-occipital.Amaphethini e-Allometric atholakala ngokulingana kobuso ngokuhlobene nosayizi wekhanda logebhezi;ogebhezini abakhudlwana izinhlaka zobuso zivame ukuba zinde futhi zibe mncane, njengoba kuye kwaboniswa kubantu abaningi boMdabu baseMelika kanye nabaseNyakatho-mpumalanga ye-Asia.Nakuba ucwaningo lwethu lungazange lufake idatha eziguquguqukayo zemvelo ezingase zibe nomthelela ku-cranial morphology, njengesimo sezulu noma izimo zokudla, isethi yedatha enkulu yamaphethini e-homologous cranial izoba usizo ekufuneni izincazelo ezihlukene zezici ze-phenotypic zamathambo.
Umehluko wendawo ekubunjweni kogebhezi lwekhanda lomuntu uye wacwaningwa isikhathi eside.Abacwaningi abaningi baye bahlola ukuhlukahluka kokujwayela kwemvelo kanye / noma ukukhethwa kwemvelo, ikakhulukazi izici zezulu1,2,3,4,5,6,7 noma umsebenzi wokukhukhumeza kuye ngezimo zokudla okunomsoco5,8,9,10,11,12.13..Ngaphezu kwalokho, ezinye izifundo zigxile emiphumeleni yebhodlela, ukukhukhuleka kofuzo, ukugeleza kwezakhi zofuzo, noma izinqubo zokuguquguquka kwe-stochastic ezibangelwa ukuguqulwa kwezakhi zofuzo ezingathathi hlangothi14,15,16,17,18,19,20,21,22,23.Isibonelo, umumo oyisiyingi we-cranial vault ebanzi nemfishane iye yachazwa njengokuzivumelanisa nengcindezi ekhethiwe ngokomthetho ka-Allen24, obeka ukuthi izilwane ezincelisayo zinciphisa ukulahlekelwa ukushisa ngokunciphisa indawo yomzimba uma kuqhathaniswa nevolumu2,4,16,17,25 .Ukwengeza, ezinye izifundo ezisebenzisa i-rule26 ka-Bergmann zichaze ubuhlobo phakathi kosayizi wogebhezi kanye nezinga lokushisa elingu-3,5,16,25,27, okuphakamisa ukuthi ubukhulu bubonke buvame ukuba bukhulu ezindaweni ezibandayo ukuvimbela ukulahlekelwa ukushisa.Umthelela wemishini wokucindezelwa kwe-masticatory iphethini yokukhula ye-cranial vault kanye namathambo obuso kuye kwaphikiswana ngakho mayelana nezimo zokudla okubangelwa isiko lokudla noma umehluko wokuziphilisa phakathi kwabalimi nabaqoqi abazingeli8,9,11,12,28.Incazelo evamile ukuthi ukuncipha kwengcindezi yokuhlafuna kunciphisa ubulukhuni bamathambo obuso nemisipha.Ucwaningo oluningana lomhlaba wonke luye lwaxhumanisa ukuhlukahluka kogebhezi lwekhanda ikakhulukazi emiphumeleni ye-phenotypic yebanga elingathathi hlangothi lezakhi zofuzo kunokujwayela imvelo21,29,30,31,32.Enye incazelo yezinguquko ekubunjweni kogebhezi isekelwe kumqondo wokukhula kwe-isometric noma i-allometric6,33,34,35.Isibonelo, ubuchopho obukhulu buvamise ukuba namalobe angaphambili abanzi uma kuqhathaniswa endaweni ebizwa ngokuthi “i-Broca's cap”, futhi ububanzi bamalobe angaphambili buyanda, inqubo yokuziphendukela kwemvelo ebhekwa ngokususelwa ekukhuleni kwe-allometric.Ukwengeza, ucwaningo oluhlola izinguquko zesikhathi eside esimweni sogebhezi lwathola ukuthambekela kwe-allometric ngase-brachycephaly (ukuthambekela kogebhezi ukuba lube yindilinga kakhudlwana) nokuphakama okukhulayo33.
Umlando omude wocwaningo lwe-cranial morphology uhlanganisa imizamo yokuhlonza izici eziyisisekelo ezibhekele izici ezihlukahlukene zokuhlukahluka kwezimo ze-cranial.Izindlela zendabuko ezisetshenziswe ezifundweni eziningi zokuqala zazisekelwe kudatha yokulinganisa yomugqa we-bivariate, ngokuvamile kusetshenziswa izincazelo zikaMartin noma zikaHowell36,37.Ngesikhathi esifanayo, izifundo eziningi ezibalwe ngenhla zasebenzisa izindlela ezithuthuke kakhulu ezisekelwe kubuchwepheshe be-3D geometric morphometry (GM) 5,7,10,11,12,13,17,20,27,34,35,38.39. Isibonelo, indlela eshelelayo ye-semilandmark, esekelwe ekunciphiseni amandla okugoba, kube yindlela esetshenziswa kakhulu ku-transgenic biology.Ikhiqiza izimpawu zendawo zesifanekiso kusampula ngayinye ngokuslayida eduze kwejika noma indawo38,40,41,42,43,44,45,46.Kuhlanganisa nezindlela ezinjalo zokubeka phezulu, izifundo eziningi ze-3D GM zisebenzisa ukuhlaziya okuvamile kwe-Procrustes, i-algorithm 47 yephoyinti eliseduze eliphindaphindayo (ICP) ukuze kuvunyelwe ukuqhathanisa okuqondile kobumo nokubanjwa kwezinguquko.Kungenjalo, indlela yepuleti elincane elincane (TPS)48,49 nayo isetshenziswa kabanzi njengendlela yoguquko engaqinile yokwenza imephu ukuqondanisa kwe-semilandmark kumajamo asuselwe enezikhala.
Ngokuthuthukiswa kwezikena ezisebenzayo ze-3D zomzimba wonke kusukela ngasekupheleni kwekhulunyaka lama-20, izifundo eziningi zisebenzise izikena ze-3D zomzimba wonke ukuze kukale usayizi50,51.Idatha yokuskena isetshenziswe ukuze kukhishwe ubukhulu bomzimba, okudinga ukuchaza umumo ongaphezulu njengendawo engaphezulu kunamafu akhomba.Ukufakwa kwephethini kuyindlela eyakhelwe le njongo emkhakheni wezithombe zekhompiyutha, lapho ukwakheka kwendawo kuchazwa ngemodeli ye-polygonal mesh.Isinyathelo sokuqala sokufakwa kwephethini ukulungisa imodeli yemeshi ezosetshenziswa njengesifanekiso.Amanye ama-vertices akha iphethini yizimpawu zendawo.Isifanekiso sibe sesigogekile futhi sihambisane nendawo ukuze kuncishiswe ibanga phakathi kwesifanekiso nefu lephoyinti kuyilapho kulondolozwa izici zomumo wendawo zesifanekiso.Izimpawu zendawo kusifanekiso zihambisana nezimpawu zendawo efwini lephoyinti.Ngokusebenzisa ukufakwa kwesifanekiso, yonke idatha yokuskena ingachazwa njengemodeli yemeshi enenani elifanayo lamaphoyinti edatha kanye ne-topology efanayo.Nakuba i-homology enembile ikhona kuphela ezindaweni eziyingqopha-mlando, kungacatshangwa ukuthi kukhona i-homology evamile phakathi kwamamodeli akhiqiziwe njengoba izinguquko ku-geometry yezifanekiso zincane.Ngakho-ke, amamodeli egridi adalwe ngokufakwa kwesifanekiso kwesinye isikhathi abizwa ngamamodeli we-homology52.Ubuhle bokufakwa kwesifanekiso ukuthi ithempulethi ingakhubazeka futhi ilungiswe ezingxenyeni ezahlukene zento eqondiwe eziseduze nendawo kodwa ezikude nayo (isibonelo, i-zygomatic arch kanye nesifunda sesikhashana sogebhezi lwekhanda) ngaphandle kokuthinta indawo ngayinye. okunye.ukuguquguquka.Ngale ndlela, ithempulethi ingavikelwa ezintweni ezihlanganisayo ezifana ne-torso noma ingalo, ihlombe lisesimweni sokuma.Ububi bokufakwa kwesifanekiso yizindleko eziphezulu zokubala zokuphindaphinda, nokho, ngenxa yentuthuko enkulu ekusebenzeni kwekhompyutha, lokhu akuseyona inkinga.Ngokuhlaziya amanani wokuxhumanisa ama-vertices akha imodeli ye-mesh esebenzisa amasu okuhlaziya ama-multivariate afana nokuhlaziywa kwengxenye eyinhloko (PCA), kungenzeka ukuhlaziya izinguquko kuwo wonke umumo ongaphezulu kanye nokuma okubonakalayo kunoma yisiphi isikhundla ekusabalaliseni.ingatholwa.Bala futhi ubone ngeso lengqondo53.Namuhla, amamodeli anezikhala akhiqizwe ukufakwa kwesifanekiso asetshenziswa kakhulu ekuhlaziyweni komumo emikhakheni ehlukahlukene52,54,55,56,57,58,59,60.
Intuthuko kubuchwepheshe bokurekhoda obunemeshi obuvumelana nezimo, obuhambisana nokuthuthuka okusheshayo kwamadivayisi aphathekayo okuskena e-3D akwazi ukuskena ngokulungiswa okuphezulu, isivinini, nokuhamba kune-CT, kwenza kube lula ukurekhoda idatha yendawo ye-3D kungakhathaliseki indawo.Ngakho-ke, emkhakheni we-biological anthropology, ubuchwepheshe obusha obunjalo buthuthukisa ikhono lokulinganisa nokuhlaziya ngokwezibalo izibonelo zabantu, okuhlanganisa nezibonelo zogebhezi, okuyinhloso yalolu cwaningo.
Kafushane, lolu cwaningo lusebenzisa ubuchwepheshe bokumodela be-homology be-3D obuthuthukisiwe obususelwe ekufanisweni kwezifanekiso (Umfanekiso 1) ukuze kuhlolwe izifanekiso zogebhezi lwekhanda ezingama-342 ezikhethwe kubantu abayi-148 emhlabeni wonke ngokuqhathanisa kwendawo emhlabeni wonke.Ukuhlukahluka kwe-cranial morphology (Ithebula 1).Ukuze siphendule ngezinguquko ku-morphology yogebhezi, sisebenzise i-PCA kanye nesici sokusebenza somamukeli (ROC) sihlaziya kusethi yedatha yemodeli ye-homology esiyikhiqizile.Okutholakele kuzoba nomthelela ekuqondeni kangcono izinguquko zomhlaba jikelele ku-cranial morphology, okuhlanganisa amaphethini wesifunda kanye nokwehla kokuhlelwa koshintsho, izinguquko ezihlotshaniswayo phakathi kwamasegimenti e-cranial, kanye nokuba khona kwamathrendi e-allometric.Nakuba lolu cwaningo lungabheki idatha ekuguquguqukeni kwangaphandle okumelelwe isimo sezulu noma izimo zokudla ezingase zibe nomthelela ku-cranial morphology, amaphethini ezindawo ze-cranial morphology abhalwe ocwaningweni lwethu azosiza ukuhlola izici zemvelo, ze-biomechanical, kanye nezakhi zofuzo zokuhlukahluka kwe-cranial.
Ithebula lesi-2 libonisa ama-eigenvalues ​​kanye nama-coefficients omnikelo we-PCA asetshenziswe kudathasethi engalinganiselwe yama-vertices angu-17,709 (izixhumanisi ezingu-53,127 XYZ) zamamodeli we-skull angama-homologous angu-342.Ngenxa yalokho, kuhlonzwe izingxenye eziyinhloko eziyi-14, umnikelo wawo ekuhlukeni usuwonke ube ngaphezu kwe-1%, kanti isamba sesamba sokuhluka sasingama-83.68%.Ama-vector alayishwayo ezingxenye eziyinhloko eziyi-14 arekhodwa kuThebula Lokwengeza S1, futhi amaphuzu engxenye abalwe amasampula ogebhezi lwama-342 ethulwa kuThebula Lokwengeza S2.
Lolu cwaningo luhlole izingxenye eziyisishiyagalolunye ezinkulu ngeminikelo emikhulu kuno-2%, ezinye zazo ezibonisa ukuhluka okukhulu nokubalulekile kwezindawo ku-cranial morphology.Umfanekiso wesi-2 wamajika akhiqizwe ekuhlaziyweni kwe-ROC ukukhombisa izingxenye ze-PCA ezisebenza kahle kakhulu zokulinganisa noma ukuhlukanisa inhlanganisela ngayinye yamasampuli kuwo wonke amayunithi amakhulu ezindawo (isb, phakathi kwamazwe ase-Afrika nalawo angewona awase-Afrika).Inhlanganisela yasePolynesia ayizange ihlolwe ngenxa yosayizi omncane wesampula osetshenziswe kulokhu kuhlolwa.Idatha ephathelene nokubaluleka komehluko ku-AUC nezinye izibalo eziyisisekelo ezibalwe kusetshenziswa ukuhlaziywa kwe-ROC ziboniswa Kuthebula Elingeziwe S3.
Amajika e-ROC asetshenziswe ekulinganisweni kwezingxenye eziyinhloko eziyisishiyagalolunye ngokusekelwe kudathasethi ye-vertex ehlanganisa amamodeli we-skull wesilisa angama-342.I-AUC: Indawo engaphansi kwejika ngo-0.01% wokubaluleka esetshenziselwa ukuhlukanisa inhlanganisela yendawo ngayinye kwezinye izinhlanganisela eziphelele.I-TPF inethemba langempela (ukucwasa okusebenzayo), i-FPF inemibono engelona iqiniso (ukucwasa okungavumelekile).
Ukuchazwa kwejika le-ROC kufingqiwe ngezansi, kugxile kuphela ezingxenyeni ezingahlukanisa amaqembu okuqhathanisa ngokuba ne-AUC enkulu noma enkulu uma kuqhathaniswa nezinga eliphezulu lokubaluleka okungenzeka libe ngaphansi kuka-0.001.I-South Asian complex (Fig. 2a), ehlanganisa ikakhulukazi amasampula avela e-India, ihluke kakhulu kwamanye amasampula axutshwe ngokwendawo ngokuthi ingxenye yokuqala (PC1) ine-AUC enkulu kakhulu (0.856) uma iqhathaniswa nezinye izingxenye.Isici se-African complex (Fig. 2b) yi-AUC enkulu kakhulu ye-PC2 (0.834).Ama-Austro-Melanesians (Fig. 2c) abonise ukuthambekela okufanayo kubantu base-Afrika base-Sub-Saharan nge-PC2 ene-AUC enkulu ngokuqhathaniswa (0.759).Abantu baseYurophu (Fig. 2d) bahluke ngokucacile ekuhlanganisweni kwe-PC2 (AUC = 0.801), i-PC4 (AUC = 0.719) ne-PC6 (AUC = 0.671), isampula yaseNyakatho-mpumalanga ye-Asia (Fig. 2e) ihluke kakhulu ku-PC4, ngokuqhathaniswa okukhulu ku-0.714, futhi umehluko kusuka ku-PC3 ubuthakathaka (AUC = 0.688).Amaqembu alandelayo aphinde akhonjwa ngamavelu e-AUC aphansi kanye namazinga okubaluleka aphezulu: Imiphumela ye-PC7 (AUC = 0.679), i-PC4 (AUC = 0.654) ne-PC1 (AUC = 0.649) ibonise ukuthi aboMdabu baseMelika (Fig. 2f) banokuthize izici ezihlotshaniswa nalezi zingxenye, ama-Southeast Asian (Fig. 2g) ahlukaniswe ku-PC3 yonkana (AUC = 0.660) ne-PC9 (AUC = 0.663), kodwa iphethini yamasampuli asuka eMpumalanga Ephakathi (Fig. 2h) (kuhlanganise neNyakatho Afrika) ihambisane.Uma kuqhathaniswa nabanye awukho umehluko omningi.
Esinyathelweni esilandelayo, ukuze utolike ngokubonakalayo ama-vertices ahlobane kakhulu, izindawo ezingaphezulu ezinamavelu omthwalo ophezulu angaphezu kuka-0.45 zinemibala ehambisana no-X, Y, kanye no-Z wolwazi, njengoba kuboniswe kuMfanekiso 3. Indawo ebomvu ibonisa ukuhlobana okuphezulu Izixhumanisi ze-X-eksisi, ezihambisana nesiqondiso esivundlile.Isifunda esiluhlaza sihlotshaniswa kakhulu nokuxhumanisa okuqondile kwe-eksisi ye-Y, futhi indawo eluhlaza okwesibhakabhaka emnyama ihlotshaniswa kakhulu ne-sagittal coordinate ye-eksisi Z.Isifunda esiluhlaza okwesibhakabhaka esikhanyayo sihlotshaniswa nezimbazo zokuxhumanisa ezingu-Y kanye nezimbazo zokuxhumanisa u-Z;pink - indawo exubile ehlotshaniswa nezimbazo zokuxhumanisa u-X no-Z;ophuzi - indawo ehlotshaniswa nezimbazo zokuxhumanisa u-X kanye no-Y;Indawo emhlophe iqukethe i-eksisi yokuxhumanisa u-X, Y kanye no-Z eboniswayo.Ngakho-ke, kulo mkhawulo wenani lomthwalo, i-PC 1 ihlotshaniswa kakhulu nendawo yonke yogebhezi.Ukwakheka kogebhezi olubonakalayo lwe-3 SD ngakolunye uhlangothi lwale axis yengxenye nakho kuyavezwa kulo mfanekiso, futhi izithombe ezisontekile zethulwa ku-Supplementary Video S1 ukuze kuqinisekiswe ukuthi i-PC1 iqukethe izici zosayizi wogebhezi lulonke.
Ukusatshalaliswa kwemvamisa kwezikolo ze-PC1 (ijika elifanele elivamile), imephu yombala yendawo yogebhezi ihlotshaniswa kakhulu nama-PC1 vertices (incazelo yemibala ehlobene ne-Ubukhulu bezinhlangothi eziphambene zale eksisi ngu-3 SD. Isikali siyindilinga eluhlaza enobubanzi kwe 50 mm.
Umfanekiso wesi-3 ubonisa isiqephu sokusabalalisa imvamisa (ijika elilinganayo elivamile) lamaphuzu e-PC1 angawodwana abalwe ngokuhlukene kumayunithi angu-9 ezindawo.Ngokungeziwe ezilinganisweni zejika le-ROC (Umfanekiso 2), izilinganiso zabantu baseNingizimu Asia ngokwezinga elithile zitshekele kwesokunxele ngenxa yokuthi ukhakhayi lwabo luncane kunelamanye amaqembu esifunda.Njengoba kukhonjisiwe kuThebula 1, laba bantu baseNingizimu Asia bamele amaqembu ezizwe e-India okuhlanganisa i-Andaman ne-Nicobar Islands, i-Sri Lanka ne-Bangladesh.
I-dimensional coefficient itholwe ku-PC1.Ukutholwa kwezifunda ezixhumene kakhulu kanye nokuma okubonakalayo kuholele ekucacisweni kwezinto zamafomu zezingxenye ngaphandle kwe-PC1;noma kunjalo, izici zesayizi aziqedi ngokuphelele ngaso sonke isikhathi.Njengoba kuboniswe ngokuqhathanisa amajika e-ROC (Umfanekiso 2), i-PC2 ne-PC4 yayibandlulula kakhulu, ilandelwa yi-PC6 ne-PC7.I-PC3 ne-PC9 zisebenza kahle kakhulu ekuhlukaniseni isampula labantu libe ngamayunithi ezindawo.Ngakho-ke, lawa mapheya ama-axes engxenye abonisa ngohlelo ama-scatterplots wezikolo ze-PC kanye nezindawo zombala ezihlotshaniswa kakhulu nengxenye ngayinye, kanye nokuguquguquka komumo okubonakalayo okunobukhulu bezinhlangothi eziphambene ze-3 SD (Fig. 4, 5, 6).Ukumbozwa kwesakhiwo se-convex samasampula avela endaweni ngayinye emelwe kulezi ziqephu cishe kungama-90%, nakuba kukhona izinga elithile lokugqagqana ngaphakathi kwamaqoqo.Ithebula lesi-3 linikeza incazelo yengxenye ngayinye ye-PCA.
Ama-scatterplots we-PC2 kanye nezikolo ze-PC4 zabantu abahlanyayo abavela ezindaweni eziyisishiyagalolunye (phezulu) kanye namayunithi ezindawo ezine (phansi), iziqephu zombala wogebhezi lwama-vertices ahlotshaniswa kakhulu ne-PC ngayinye (okuhlobene no-X, Y, Z).Incazelo yombala wama-ax: bona umbhalo), futhi ukuguqulwa kwefomu elibonakalayo ezinhlangothini eziphambene zalezi zimbazo kungu-3 SD.Isikali siyindilinga eluhlaza enobubanzi obungama-50 mm.
Ama-scatterplots we-PC6 kanye nezikolo ze-PC7 zabantu abahlanyayo abavela ezindaweni eziyisishiyagalolunye (phezulu) kanye namayunithi amabili ezindawo (phansi), iziza zombala we-cranial surface for vertices ezihlotshaniswa kakhulu ne-PC ngayinye (okuhlobene no-X, Y, Z).Incazelo yombala wama-ax: bona umbhalo), futhi ukuguqulwa kwefomu elibonakalayo ezinhlangothini eziphambene zalezi zimbazo kungu-3 SD.Isikali siyindilinga eluhlaza enobubanzi obungama-50 mm.
Ama-scatterplots ezikolo ze-PC3 ne-PC9 zabantu abahlanyayo abavela ezindaweni eziyisishiyagalolunye (phezulu) kanye namayunithi amathathu ezindawo (phansi), kanye nezindawo zombala zogebhezi (ezihlobene nezimbazo ezingu-X, Y, Z) ezihlotshaniswa kakhulu nokutolikwa kombala kwe-PC ngayinye. :cm.umbhalo), kanye nokuguqulwa komumo obonakalayo ezinhlangothini eziphambene zalezi zimbazo ezinobukhulu obungu-3 SD.Isikali siyindilinga eluhlaza enobubanzi obungama-50 mm.
Kugrafu ebonisa amaphuzu e-PC2 kanye ne-PC4 (I-Fig. 4, Supplementary Videos S2, S3 ebonisa izithombe ezikhubazekile), ibalazwe lombala ongaphezulu liyaboniswa uma umkhawulo wenani lomthwalo usethwe ngaphezulu kuka-0.4, ongaphansi kune-PC1 ngoba Inani le-PC2 inani lomthwalo eliphelele lingaphansi kwe-PC1.
Ukwelulwa kwe-lobes yangaphambili ne-occipital kuhlangothi lwe-sagittal eduze kwe-axis engu-Z (luhlaza okwesibhakabhaka) kanye ne-parietal lobe endaweni ye-coronal (bomvu) ngokubomvana), i-Y-axis ye-occiput (eluhlaza okotshani) kanye ne-Z-eksisi. ebunzini (blue blue).Le grafu ikhombisa amaphuzu abo bonke abantu emhlabeni jikelele;nokho, lapho wonke amasampula ahlanganisa inani elikhulu lamaqembu eboniswa ndawonye ngesikhathi esisodwa, ukuchazwa kwamaphethini okuhlakazeka kunzima kakhulu ngenxa yenani elikhulu lokugqagqana;ngakho-ke, kusukela kumayunithi amane amakhulu ezindawo (okungukuthi, i-Afrika, i-Australasia-Melanesia, i-Europe, kanye ne-Northeast Asia), amasampula ahlakazekile ngezansi kwegrafu nge-3 SD virtual cranial deformation ngaphakathi kwalolu hlu lwezikolo ze-PC.Emfanekisweni, i-PC2 ne-PC4 zingamapheya ezikolo.Ama-Afrika kanye nama-Austro-Melanesians agqagqana kakhulu futhi asakazwa ohlangothini lwesokudla, kuyilapho ama-Europe ahlakazekile ngasenhla kwesokunxele kanti abaseNyakatho-mpumalanga ye-Asia bavame ukuhlangana kwesokunxele.I-axis evundlile ye-PC2 ibonisa ukuthi ama-Afrika/Australian Melanesians ane-neurocranium ende kunabanye abantu.I-PC4, lapho inhlanganisela yaseYurophu kanye nenyakatho-mpumalanga ye-Asia ihlukaniswa ngokukhululekile, ihlotshaniswa nobukhulu obuhlobene nokuqagela kwamathambo e-zygomatic kanye ne-lateral contour ye-calvarium.Uhlelo lokushaya amaphuzu lubonisa ukuthi abantu baseYurophu banamathambo amancane e-maxillary kanye ne-zygomatic, isikhala esincane se-fossa yesikhashana esinqunyelwe i-zygomatic arch, ithambo langaphambili eliphakame phezulu kanye nethambo eliyisicaba, eliphansi le-occipital, kuyilapho abantu baseNyakatho-mpumalanga ye-Asia bevame ukuba namathambo e-zygomatic abanzi futhi avelele kakhudlwana. .I-lobe yangaphambili ithambekele, isisekelo sethambo le-occipital siphakanyiswa.
Uma ugxile ku-PC6 ne-PC7 (Fig. 5) (Amavidiyo Engeziwe S4, i-S5 ebonisa izithombe ezikhubazekile), isakhiwo sombala sibonisa umkhawulo wenani lomthwalo omkhulu kuno-0.3, okubonisa ukuthi i-PC6 ihlotshaniswa ne-maxillary noma i-alveolar morphology (obomvu : i-X axis kanye green).I-axis engu-Y), ukwakheka kwethambo lesikhashana (okuluhlaza okwesibhakabhaka: izimbazo zika-Y kanye no-Z) kanye ne-occipital bone shape (pink: X kanye nezimbazo Z).Ngaphezu kobubanzi besiphongo (obomvu: i-X-axis), i-PC7 iphinde ihlobane nokuphakama kwe-alveoli ye-maxillary yangaphambili (eluhlaza: i-Y-axis) kanye nokuma kwekhanda le-Z-axis ezungeze isifunda se-parietotemporal (blue blue).Kuphaneli ephezulu yoMfanekiso 5, wonke amasampula ezindawo asatshalaliswa ngokuya ngezingxenye ze-PC6 ne-PC7.Ngenxa yokuthi i-ROC ikhombisa ukuthi i-PC6 iqukethe izici ezihlukile kwe-Europe futhi i-PC7 imele izici zoMdabu waseMelika kulokhu kuhlaziywa, lawa masampuli amabili esifunda ahlelwe ngokukhetha kulokhu kupheya kwezimbazo zengxenye.Abomdabu baseMelika, nakuba befakwe kabanzi kusampula, bahlakazekile ekhoneni elingenhla kwesokunxele;ngokuphambene, amasampula amaningi aseYurophu avame ukutholakala ekhoneni elingezansi kwesokudla.Ipheya i-PC6 ne-PC7 imele inqubo encane ye-alveolar kanye ne-neurocranium ebanzi ngokuqhathaniswa yabantu baseYurophu, kuyilapho amaMelika ebonakala ngebunzi eliwumngcingo, i-maxilla enkulu, kanye nenqubo ye-alveolar ebanzi futhi ende.
Ukuhlaziywa kwe-ROC kubonise ukuthi i-PC3 kanye/noma i-PC9 zazivamile kubantu baseNingizimu-mpumalanga nasenyakatho-mpumalanga ye-Asia.Ngokuvumelana nalokho, amaphuzu amapheya i-PC3 (ubuso obuphezulu obuluhlaza ku-axis engu-y) kanye ne-PC9 (ubuso obuluhlaza obuphansi obuphansi ku-y-eksisi) (Umfanekiso 6; Amavidiyo Angeziwe S6, S7 ahlinzeka ngezithombe eziguquguqukayo) abonisa ukuhlukahluka kwabantu baseMpumalanga Asia., ehluke kakhulu ngokulingana okuphezulu kobuso baseNyakatho-mpumalanga ye-Asia kanye nesimo esiphansi sobuso sabase-Southeast Asia.Ngaphandle kwalezi zici zobuso, esinye isici sabanye abantu baseNyakatho-mpumalanga ye-Asia ukutsheka kwe-lambda kwethambo le-occipital, kuyilapho abanye abantu baseNingizimu-mpumalanga ye-Asia benesisekelo sogebhezi oluncane.
Incazelo engenhla yezingxenye eziyinhloko kanye nencazelo ye-PC5 ne-PC8 ayifakiwe ngoba azikho izici ezithile zesifunda ezitholakele phakathi kwamayunithi ayisishiyagalolunye ezindawo.I-PC5 isho usayizi wenqubo ye-mastoid yethambo lesikhashana, futhi i-PC8 ikhombisa ukungafani kokuma kogebhezi lulonke, womabili abonisa ukuhluka okuhambisanayo phakathi kwezinhlanganisela zamasampula ezindawo eziyisishiyagalolunye.
Ngokungeziwe kuma-scatterplots wezinga ngalinye lezikolo ze-PCA, siphinde futhi sinikeze ama-scatterplots ezindlela zeqembu ukuze kuqhathaniswe sekukonke.Kuze kube manje, imodeli ye-cranial homology emaphakathi yadalwa kusukela kusethi yedatha ye-vertex yamamodeli e-homology ngayinye evela kumaqembu ezizwe eziyi-148.Iziqephu ze-Bivariate zamasethi wamaphuzu we-PC2 ne-PC4, i-PC6 ne-PC7, ne-PC3 ne-PC9 ziboniswa ku-Supplementary Figure S1, konke kubalwa njengemodeli yogebhezi olumaphakathi lwesampula labantu abangabodwana abangu-148.Ngale ndlela, ama-scatterplots afihla umehluko ngamunye eqenjini ngalinye, okuvumela ukuchazwa okucacile kokufana kogebhezi ngenxa yokusatshalaliswa kwesifunda okuyisisekelo, lapho amaphethini afana nalawo avezwe ezakhiweni ezingazodwana ezinokugqagqana okuncane.I-Supplementary Figure S2 ibonisa imodeli emaphakathi yeyunithi ngayinye yendawo.
Ngokungeziwe ku-PC1, eyayihlotshaniswa nosayizi jikelele (Ithebula Le-Supplementary S2), ubudlelwano be-allometric phakathi kobukhulu obuphelele kanye nokuma kogebhezi buhlolwe kusetshenziswa ubukhulu obuyi-centroid namasethi wezilinganiso ze-PCA kusuka kudatha engajwayelekile.Ama-coefficients e-Allometric, amanani angashintshi, amanani angu-t, namanani angu-P ekuhlolweni kokubaluleka aboniswa kuThebula 4. Azikho izingxenye ezibalulekile zephethini ye-allometric ezihlotshaniswa nosayizi wekhanda logebhezi olutholakele kunoma iyiphi i-cranial morphology ezingeni le-P <0.05.
Ngenxa yokuthi ezinye izici zosayizi zingase zifakwe ezilinganisweni ze-PC ezisekelwe kumasethi edatha angajwayelekile, siqhubekile sahlola inkambiso ye-allometric phakathi kosayizi we-centroid kanye nezikolo ze-PC ezibalwe kusetshenziswa amasethi edatha ajwayeleke ngosayizi we-centroid (imiphumela ye-PCA namasethi wamaphuzu ethulwa kokuthi Supplementary Tables S6 ).,C7).Ithebula lesi-4 libonisa imiphumela yokuhlaziywa kwe-allometric.Ngakho-ke, izitayela eziphawulekayo ze-allometric zitholwe ezingeni le-1% ku-PC6 kanye nezinga le-5% ku-PC10.Umfanekiso 7 ubonisa ukuhlehla kwemithambeka yalobu budlelwano bomugqa welogi phakathi kwezikolo ze-PC kanye nosayizi we-centroid onama-dummies (±3 SD) kunoma ikuphi ukuphela kosayizi welogi we-log.Isikolo se-PC6 isilinganiso sobude obuhlobene nobubanzi bogebhezi.Njengoba ubukhulu bogebhezi lukhula, ugebhezi nobuso kuba phezulu, kanti isiphongo, izisekelo zamehlo namakhala kuvame ukusondelana eduze.Iphethini yokusatshalaliswa kwesampula iphakamisa ukuthi le ngxenye ivame ukutholakala eNyakatho-mpumalanga ye-Asia kanye naboMdabu baseMelika.Ngaphezu kwalokho, i-PC10 ibonisa ukuthambekela ekuncishisweni ngokulinganayo kobubanzi obuphakathi nendawo ngaphandle kokunaka isifunda sendawo.
Ngobudlelwano obubalulekile be-allometric obusohlwini lwethebula, umthambeko wokuhlehla komugqa welogi phakathi kwengxenye ye-PC yengxenye yomumo (etholakala kudatha evamile) nosayizi we-centroid, ukwakheka komumo obonakalayo kunosayizi ongu-3 SD ohlangothini oluphambene lomugqa wesi-4.
Iphethini elandelayo yezinguquko ku-cranial morphology iye yaboniswa ngokuhlaziywa kwamasethi edatha wamamodeli angaphezulu kwe-3D angama-homologous.Ingxenye yokuqala ye-PCA ihlobene nosayizi wekhanda logebhezi.Sekuyisikhathi eside kucatshangwa ukuthi izingebhezi ezincane zaseNingizimu Asia, kuhlanganise nezibonelo ezivela eNdiya, eSri Lanka nase-Andaman Islands, Bangladesh, ngenxa yobukhulu bemizimba yazo emincane, ngokuhambisana nomthetho we-ecogeographic kaBergmann noma umthetho wesiqhingi613,5,16,25, 27,62 .Eyokuqala ihlobene nezinga lokushisa, kanti eyesibili incike endaweni etholakalayo kanye nezinsiza zokudla ze-niche yemvelo.Phakathi kwezingxenye zomumo, ushintsho olukhulu kakhulu isilinganiso sobude nobubanzi be-cranial vault.Lesi sici, esiqokiwe i-PC2, sichaza ubudlelwano obuseduze phakathi kwezingebhezi ezinde ngokulinganayo zama-Austro-Melanesians kanye nama-Afrika, kanye nokwehluka kogebhezi oluyindilinga lwabathile baseYurophu nabaseNyakatho-mpumalanga ye-Asia.Lezi zici zibikwe ezifundweni eziningi ezedlule ngokusekelwe ezilinganisweni ezilula zomugqa37,63,64.Ngaphezu kwalokho, lesi sici sihlotshaniswa ne-brachycephaly kubantu abangewona ama-Afrika, osekuyisikhathi eside kuxoxwa ngayo ezifundweni ze-anthropometric kanye ne-osteometric.I-hypothesis eyinhloko ngemuva kwalesi ncazelo yukuthi ukuncipha kwe-mastication, njengokunciphisa imisipha ye-temporalis, kunciphisa ukucindezela ku-scalp yangaphandle5,8,9,10,11,12,13.Enye inkolelo-mbono ihilela ukuzivumelanisa nezimo zezulu ezibandayo ngokunciphisa indawo engaphezulu kwekhanda, okuphakamisa ukuthi ugebhezi oluyindilinga kakhudlwana lunciphisa indawo engcono kunokwakheka kwendilinga, ngokwemithetho ka-Allen16,17,25.Ngokusekelwe emiphumeleni yocwaningo lwamanje, le mibono ingahlolwa kuphela ngokusekelwe ekuxhumaneni okuphambene kwamasegimenti e-cranial.Kafushane, imiphumela yethu ye-PCA ayisekeli ngokugcwele i-hypothesis yokuthi isilinganiso sobude bobubanzi be-cranial sithonywa kakhulu izimo zokuhlafuna, njengoba ukulayisha kwe-PC2 (ingxenye ende/ye-brachycephalic) bekungahlobene kakhulu nezilinganiso zobuso (kuhlanganise nobukhulu obuhlobene nobukhulu be-maxillary).kanye nesikhala esihlobene se-fossa yesikhashana (ebonisa umthamo we-temporalis muscle).Ucwaningo lwethu lwamanje aluzange luhlaziye ubudlelwano phakathi kokuma kogebhezi nezimo zemvelo zemvelo ezifana nezinga lokushisa;Nokho, incazelo esekelwe kumthetho ka-Allen ingase ifanele ukucatshangelwa njenge-hypothesis yekhandidethi ukuchaza i-brachycephalon ezindaweni ezibandayo zesimo sezulu.
Ukuhluka okuphawulekayo kwabe sekutholakala ku-PC4, okuphakamisa ukuthi abantu baseNyakatho-mpumalanga ye-Asia banamathambo amakhulu, avelele e-zygomatic ku-maxilla namathambo e-zygomatic.Lokhu kutholakala kuhambisana nesici esikhethekile esaziwayo sabantu baseSiberia, okucatshangwa ukuthi bajwayele izimo zezulu ezibandayo kakhulu ngokuhamba phambili kwamathambo e-zygomatic, okuholela ekukhuleni kwevolumu yezono kanye nobuso obuthambile 65.Okusha okutholakele kumodeli yethu ye-homologous ukuthi ukugoba kwesihlathi kubantu baseYurophu kuhlotshaniswa nomthambeka ongaphambili oncishisiwe, kanye namathambo e-occipital ayisicaba futhi amancane kanye ne-nuchal concavity.Ngokuphambene, abantu baseNyakatho-mpumalanga ye-Asia bavame ukuba namabunzi atshekile futhi baphakamise izifunda ze-occipital.Izifundo zethambo le-occipital zisebenzisa izindlela ze-geometric morphometric35 zibonise ukuthi ama-skulls ase-Asia naseYurophu ane-flatter nuchal curve kanye nesimo esiphansi se-occiput uma kuqhathaniswa nama-Afrika.Kodwa-ke, ama-scatterplots ethu e-PC2 kanye ne-PC4 kanye ne-PC3 kanye ne-PC9 ngamapheya abonise ukuhluka okukhulu kwabase-Asia, kuyilapho abaseYurophu bebonakala ngesisekelo esiyisicaba se-occiput kanye ne-occiput ephansi.Ukungezwani ezicini zase-Asia phakathi kwezifundo kungase kube ngenxa yomehluko kumasampuli ohlanga asetshenzisiwe, njengoba senza isampula inombolo enkulu yamaqembu ezizwe avela ku-spectrum ebanzi yaseNyakatho-mpumalanga naseNingizimu-mpumalanga ye-Asia.Izinguquko ekubunjweni kwethambo le-occipital zivame ukuhlotshaniswa nokuthuthukiswa kwemisipha.Kodwa-ke, le ncazelo eguquguqukayo ayibali ukuhlobana phakathi kwebunzi kanye nesimo se-occiput, okuboniswe kulolu cwaningo kodwa akunakwenzeka ukuthi kuboniswe ngokugcwele.Mayelana nalokhu, kufanelekile ukucabangela ubudlelwano phakathi kwebhalansi yesisindo somzimba kanye nendawo ye-gravity noma ukuhlangana komlomo wesibeletho (i-foramen magnum) noma ezinye izici.
Enye ingxenye ebalulekile enokuhlukahluka okukhulu ihlobene nokuthuthukiswa kwemishini yokukhukhumeza, emelwe yi-maxillary kanye ne-temporal fossae, echazwa ngenhlanganisela yamaphuzu PC6, PC7 kanye ne-PC4.Lokhu kwehliswa okuphawuliwe kumasegimenti e-cranial kuveza abantu base-Europe ngaphezu kwanoma yiliphi elinye iqembu lendawo.Lesi sici sihunyushwe njengomphumela wokuncipha kokuqina kokuma kobuso ngenxa yokuthuthukiswa kwangaphambi kwesikhathi kwamasu okulungisa ezolimo nokudla, okwabuye kwanciphisa umthwalo wemishini emshinini wokuzikhukhumeza ngaphandle kwemishini enamandla yokukhukhumala9,12,28,66.Ngokuvumelana nenkolelo-mbono yokusebenza kwe-masticatory, 28 lokhu kuhambisana noshintsho ekugwetshweni kwesisekelo sogebhezi lube i-engeli ebukhali ye-cranial kanye nophahla oluyindilinga lwe-cranial.Kusukela kulo mbono, abantu bezolimo bavame ukuba nobuso obuhlangene, ukuphuma kancane kwe-mandible, kanye nama-meninges azungeza umhlaba.Ngakho-ke, lokhu kuguqulwa kungachazwa ngohlaka olujwayelekile lokuma kwe-lateral ye-skull yabaseYurophu abanezitho ezincishisiwe zokuxubha.Kodwa-ke, ngokwalolu cwaningo, le ncazelo iyinkimbinkimbi ngoba ukubaluleka kokusebenza kobudlelwane be-morphological phakathi kwe-globose neurocranium kanye nokuthuthukiswa kwe-masticatory apparatus akwamukeleki kancane, njengoba kucatshangelwe ekuchazeni kwangaphambili kwe-PC2.
Umehluko phakathi kwabaseNyakatho-mpumalanga ye-Asia kanye nabase-Asia-mpumalanga uboniswa umehluko phakathi kobuso obude obunethambo le-occipital elitshekile kanye nobuso obufushane obunesisekelo sogebhezi oluncane, njengoba kukhonjisiwe ku-PC3 naku-PC9.Ngenxa yokuntuleka kwedatha ye-geoecological, ucwaningo lwethu lunikeza kuphela incazelo elinganiselwe yalokhu kutholakele.Incazelo engenzeka ukujwayela isimo sezulu esihlukile noma izimo zokudla okunomsoco.Ngaphezu kokujwayela imvelo, umehluko wendawo emlandweni wabantu baseNyakatho-mpumalanga naseNingizimu-mpumalanga ye-Asia nawo wacatshangelwa.Isibonelo, empumalanga ye-Eurasia, imodeli enezingqimba ezimbili iye yacatshangelwa ukuze kuqondwe ukuhlakazwa kwabantu besimanje be-anatomically (AMH) ngokusekelwe kudatha ye-cranial morphometric67,68.Ngokwale modeli, “isigaba sokuqala”, okungukuthi, amaqembu okuqala ama-Late Pleistocene AMH colonizers, ayenozalo oluqondile noma oluncane oluvela kubantu bomdabu bakulesi sifunda, njengama-Austro-Melanesians anamuhla (p. First stratum)., futhi kamuva sathola ingxubevange enkulu yabantu bezolimo basenyakatho abanezici zaseNyakatho-mpumalanga ye-Asia (ungqimba lwesibili) esifundeni (eminyakeni engaba ngu-4,000 edlule).Ukugeleza kofuzo okufakwe kumephu kusetshenziswa imodeli “yezendlalelo ezimbili” kuzodingeka ukuze kuqondwe umumo we-cranial waseNingizimu-mpumalanga Asia, uma kubhekwa ukuthi ukwakheka kwe-cranial yaseNingizimu-mpumalanga Asia kungase kuncike ngokwengxenye efeni lofuzo lezinga lokuqala lendawo.
Ngokuhlola ukufana kwe-cranial kusetshenziswa amayunithi ezwe adwetshwe kusetshenziswa amamodeli ayafana, singafinyelela umlando wabantu oyisisekelo we-AMF ezimweni ezingaphandle kwe-Afrika.Kuphakanyiswe amamodeli amaningi ahlukene "angaphandle kwe-Afrika" ukuze achaze ukusatshalaliswa kwe-AMF ngokusekelwe kudatha yamathambo kanye ne-genomic.Kulezi, ucwaningo lwakamuva luphakamisa ukuthi ukwenziwa kwamakoloni kwe-AMH ezindaweni ezingaphandle kwe-Afrika kwaqala cishe eminyakeni eyi-177,000 edlule69,70.Kodwa-ke, ukusatshalaliswa okude kwe-AMF e-Eurasia phakathi nalesi sikhathi akuqinisekile, njengoba izindawo zokuhlala zalezi zinsalela zakuqala zilinganiselwe eMpumalanga Ephakathi naseMedithera eduze ne-Afrika.Icala elilula indawo yokuhlala eyodwa egudle umzila wokufuduka usuka e-Afrika uya e-Eurasia, udlula imingcele yendawo efana nezintaba ze-Himalaya.Enye imodeli iphakamisa amagagasi amaningi okufuduka, eyokuqala yawo esabalalisa esuka e-Afrika ngasogwini lolwandle i-Indian Ocean eya eNingizimu-mpumalanga ye-Asia nase-Australia, abese esakazekela enyakatho ye-Eurasia.Iningi lalezi zifundo liqinisekisa ukuthi i-AMF yasabalala yaze yadlula e-Afrika eminyakeni engu-60,000 edlule.Kulokhu, amasampula e-Australasian-Melanesia (kuhlanganise ne-Papua) abonisa ukufana okukhulu namasampuli ase-Afrika kunanoma yiluphi olunye uchungechunge lwezindawo ekuhlaziyeni kwezingxenye eziyinhloko zamamodeli e-homology.Lokhu kutholakala kusekela umbono wokuthi amaqembu okuqala okusabalalisa i-AMF ngasemaphethelweni aseningizimu ye-Eurasia avela ngqo e-Afrika22,68 ngaphandle kwezinguquko ezibalulekile ze-morphological ukuphendula izimo zezulu ezithile noma ezinye izimo ezibalulekile.
Mayelana nokukhula kwe-allometric, ukuhlaziya kusetshenziswa izingxenye zomumo ezithathwe kusethi yedatha ehlukile evamile ngosayizi we-centroid kubonise ukuthambekela okubalulekile kwe-allometric ku-PC6 naku-PC10.Zombili lezi zingxenye zihlobene nokuma kwebunzi kanye nezingxenye zobuso, eziba zincane njengoba ubukhulu bogebhezi bukhula.Abantu baseNyakatho-mpumalanga ye-Asia nabaseMelika bavame ukuba nalesi sici futhi banogebhezi olukhulu ngokuqhathaniswa.Lokhu okutholakele kungqubuzana namaphethini e-allometric abikwe ngaphambilini lapho ubuchopho obukhulu bunamalobe angaphambili abanzi ngokuqhathaniswa endaweni ebizwa ngokuthi “i-Broca's cap”, okuholela ekwenyukeni kobubanzi belobe yangaphambili34.Lo mehluko uchazwa ngokuhluka kwamasethi amasampula;Ucwaningo lwethu luhlaziye amaphethini e-allometric osayizi we-cranial uwonke kusetshenziswa isibalo sabantu besimanje, futhi izifundo eziqhathanisayo zikhuluma ngamathrendi esikhathi eside ekuziphendukeleni kwemvelo komuntu okuhlobene nosayizi wobuchopho.
Mayelana ne-allometry yobuso, ucwaningo olulodwa olusebenzisa idatha ye-biometric78 lwathola ukuthi ukuma kobuso nosayizi kungase kuhlotshaniswe kancane, kuyilapho ucwaningo lwethu lwathola ukuthi izingebhezi ezinkulu zivame ukuhlotshaniswa nobuso obude, obuncane.Nokho, ukuvumelana kwedatha ye-biometric akucacile;Ukuhlolwa kokuhlehla okuqhathaniswa ne-ontogenetic allometry kanye ne-static allometry kubonisa imiphumela ehlukene.Ukuthambekela kwe-allometric ekuqondeni kogebhezi oluyisiyingi ngenxa yokuphakama okwandayo nakho kubikiwe;Nokho, asizange sihlaziye idatha yobude.Ucwaningo lwethu luyabonisa ukuthi ayikho idatha ye-allometric ebonisa ukuhlobana phakathi kwezilinganiso ze-cranial globular kanye nosayizi we-cranial jikelele ngeseshini ngayinye.
Nakuba ucwaningo lwethu lwamanje lungabhekani nedatha yeziguquko zangaphandle ezimelelwe isimo sezulu noma izimo zokudla okungenzeka zibe nomthelela ku-cranial morphology, isethi yedatha enkulu yamamodeli e-homologous 3D cranial surface asetshenziswe kulolu cwaningo azosiza ukuhlola ukuhlukahluka okuhlobene kwe-phenotypic morphological.Izici zemvelo ezifana nokudla, isimo sezulu kanye nezimo zokudla okunomsoco, kanye namandla angathathi hlangothi njengokufuduka, ukugeleza kwezakhi zofuzo kanye nokukhukhuleka kofuzo.
Lolu cwaningo lwaluhlanganisa izibonelo ezingu-342 zezingebhezi zabesilisa eziqoqwe kubantu abangu-148 ezindaweni ezingu-9 zendawo (Ithebula 1).Amaqembu amaningi ayizibonelo zendawo, kuyilapho amanye amaqembu e-Afrika, eNyakatho-mpumalanga/eningizimu-mpumalanga ye-Asia naseMelika (abasohlwini olumalukeke) achazwa ngokobuhlanga.Izibonelo eziningi ze-cranial zikhethwe kusizindalwazi sokulinganisa i-cranial ngokuya ngencazelo ye-Martin cranial measurement ehlinzekwa ngu-Tsunehiko Hanihara.Sikhethe ogebhezi bekhanda besilisa abavela kuzo zonke izinhlanga emhlabeni.Ukuze sihlonze amalungu eqembu ngalinye, sibale amabanga e-Euclidean ngokusekelwe ezilinganisweni ezingu-37 ze-cranial kusuka eqenjini okusho kubo bonke abantu balelo qembu.Ezimweni eziningi, sikhethe amasampula angu-1–4 anebanga elincane ukusuka kuncazelo (Ithebula Elingeziwe S4).Kulawa maqembu, amanye amasampuli akhethwe ngokungahleliwe uma ayengekho ohlwini lwesizindalwazi sokulinganisa i-Hahara.
Ukuze kuqhathaniswe izibalo, amasampula abantu abayi-148 aqoqwe aba izingxenye ezinkulu zezwe, njengoba kukhonjisiwe kuThebula 1. Iqembu "labase-Afrika" liqukethe kuphela amasampula avela esifundeni esiseningizimu yeSahara.Izibonelo ezivela eNyakatho Afrika zafakwa "eMpumalanga Ephakathi" kanye nezibonelo ezivela eNtshonalanga ye-Asia ezinezimo ezifanayo.Iqembu laseNyakatho-mpumalanga ye-Asia lihlanganisa kuphela abantu abangebona abaseYurophu, futhi iqembu laseMelika lihlanganisa aboMdabu baseMelika kuphela.Ikakhulukazi, leli qembu lisatshalaliswa endaweni enkulu yamazwekazi aseNyakatho naseNingizimu Melika, ezindaweni ezihlukahlukene.Kodwa-ke, sicabangela isampula yase-US ngaphakathi kwale yunithi eyodwa yendawo, uma kubhekwa umlando wezibalo zabantu boMdabu baseMelika abathathwa njengabadabuka eNyakatho-mpumalanga ye-Asia, kungakhathaliseki ukufuduka okuningi okungu-80.
Sirekhode idatha yendawo engu-3D yalezi zibonelo zogebhezi oluhlukile sisebenzisa iskena se-3D sokucaca okuphezulu (i-EinScan Pro by Shining 3D Co Ltd, ukulungiswa okuphansi: 0.5 mm, https://www.shining3d.com/) sase sikhiqiza i-mesh.Imodeli ye-mesh iqukethe cishe ama-vertices angu-200,000–400,000, futhi isofthiwe efakiwe isetshenziselwa ukugcwalisa izimbobo namaphethelo abushelelezi.
Esinyathelweni sokuqala, sisebenzise idatha yokuskena kunoma yiluphi ugebhezi ukuze sakhe imodeli yogebhezi lwe-mesh yesifanekiso esisodwa ehlanganisa ama-vertices angu-4485 (8728 polygonal faces).Isisekelo sesifunda sogebhezi, esihlanganisa ithambo le-sphenoid, ithambo lesikhashana elithambile, ulwanga, i-maxillary alveoli, namazinyo, sikhishiwe kumodeli yemeshi yesifanekiso.Isizathu siwukuthi lezi zakhiwo ngezinye izikhathi aziphelele noma kunzima ukuziqedela ngenxa yezingxenye ezicijile ezincane noma ezincane ezifana nezindawo ze-pterygoid nezinqubo ze-styloid, ukuguga kwamazinyo kanye/noma isethi yamazinyo engahambisani.Isisekelo sogebhezi oluzungeze i-foramen magnum, okuhlanganisa nesisekelo, asizange sikhishwe kabusha ngoba lena indawo ebalulekile ye-anatomically yendawo yamalunga omlomo wesibeletho futhi ukuphakama kogebhezi kufanele kuhlolwe.Sebenzisa izindandatho zesibuko ukuze wenze isifanekiso esinokulinganisa kuzo zombili izinhlangothi.Yenza i-isotropic meshing ukuze uguqule umumo we-polygonal ukuze ulingane ngangokunokwenzeka.
Okulandelayo, izimpawu zendawo ezingama-56 zabelwa kuma-vertices ahambelana ngokwemvelo wemodeli yesifanekiso kusetshenziswa isofthiwe ye-HBM-Rugle.Izilungiselelo zophawu lomhlaba ziqinisekisa ukunemba nokuzinza kokuma kwendawo eyingqopha-mlando futhi ziqinisekisa i-homology yalezi zindawo kumodeli ye-homology ekhiqiziwe.Angabonakala ngokusekelwe ezicini zawo ezithile, njengoba kukhonjisiwe kuThebula Le-Supplementary S5 kanye Nomfanekiso Owengeziwe S3.Ngokwencazelo ka-Bookstein81, iningi lalezi zimpawu zomhlaba ziyizimpawu zomhlaba zohlobo I ezitholakala empambanweni yezakhiwo ezintathu, kanti ezinye ziyizimpawu zendawo zohlobo lwe-II ezinamaphoyinti okugoba okukhulu.Izimpawu zendawo eziningi zidluliswe zisuka ezindaweni ezichazwe ukuze kukale umugqa wohlaka oluqondile encazelweni ka-Martin 36. Sichaze izimpawu zendawo ezifanayo ezingu-56 zamamodeli askeniwe wezibonelo zogebhezi lwekhanda elingu-342, ezabelwa mathupha ezihlalweni ezihambisanayo ukuze kukhiqizwe amamodeli e-homology anembe kakhulu esigabeni esilandelayo.
Isistimu yokuxhumanisa i-head-centric coordinate yachazwa ukuchaza idatha yokuskena nesifanekiso, njengoba kuboniswe ku-Supplementary Figure S4.Indiza ye-XZ iyindiza evundlile yaseFrankfurt edlula endaweni ephakeme kakhulu (incazelo kaMartin: ingxenye) yonqenqema oluphakeme lwemisele yokuzwa yangaphandle kwesokunxele nesokudla kanye nephuzu eliphansi kakhulu (incazelo kaMartin: i-orbit) yonqenqema olungezansi lwe-orbit kwesokunxele. ..I-axis engu-X umugqa oxhuma izinhlangothi zesobunxele nesokudla, futhi u-X+ ohlangothini lwesokudla.Indiza ye-YZ idlula maphakathi nengxenye yesokunxele nesokudla kanye nempande yekhala: Y+ phezulu, Z+ phambili.Iphoyinti eliyinkomba (umsuka: i-zero coordinate) isethwe ezimpambanweni zendiza ye-YZ (indiza emaphakathi), indiza ye-XZ (indiza yaseFrankfort) kanye nendiza ye-XY (indiza yamakhorali).
Sisebenzise isofthiwe ye-HBM-Rugle (I-Medic Engineering, Kyoto, http://www.rugle.co.jp/) ukuze sakhe imodeli ye-mesh ye-homologous ngokwenza ukufaka isifanekiso sisebenzisa amaphuzu anomlando angu-56 (uhlangothi lwesobunxele loMfanekiso 1).Ingxenye yesofthiwe ewumgogodla, eyasungulwa ekuqaleni i-Center for Digital Human Research e-Institute of Advanced Industrial Science and Technology e-Japan, ibizwa nge-HBM futhi inemisebenzi yokufaka izifanekiso zisebenzisa izimpawu zendawo nokudala amamodeli anezikhala amahle kusetshenziswa indawo ehlukanisayo82.Inguqulo yesofthiwe eyalandela (mHBM) 83 yengeze isici sokulinganisa iphethini ngaphandle kwezimpawu zendawo ukuze kuthuthukiswe ukusebenza okufanelekile.I-HBM-Rugle ihlanganisa isofthiwe ye-mHBM nezici ezengeziwe ezisebenziseka kalula ezihlanganisa ukwenza ngendlela oyifisayo amasistimu wokuxhumanisa nokushintsha usayizi wedatha yokufaka.Ukuthembeka kokunemba kokufakwa kwesoftware kuqinisekisiwe ezifundweni eziningi52,54,55,56,57,58,59,60.
Lapho kufakwa isifanekiso se-HBM-Rugle kusetshenziswa izimpawu zendawo, imodeli yemeshi yesifanekiso ibekwa phezulu kudatha yokuskena okuqondiwe ngokubhaliswa okuqinile okusekelwe kubuchwepheshe be-ICP (ukunciphisa isamba samabanga phakathi kwezimpawu zendawo ezihambisana nesifanekiso kanye nedatha yokuskena okuqondiwe), futhi bese ngokuguqulwa okungaqinile kwe-mesh ivumelanisa isifanekiso kudatha yokuskena okuqondiwe.Le nqubo yokufaka iphindwe kathathu kusetshenziswa amanani ahlukene wamapharamitha amabili afanelekile ukuze kuthuthukiswe ukunemba kokufaka.Enye yalawa mapharamitha ikhawulela ibanga phakathi kwemodeli yegridi yesifanekiso kanye nedatha yokuskena okuqondiwe, futhi enye ijezisa ibanga phakathi kwezimpawu zendawo zesifanekiso nezimpawu zendawo eziqondiwe.Imodeli yemeshi yesifanekiso ekhubazekile yabe isihlukaniswa kusetshenziswa i-cyclic surface subdivision algorithm 82 ukuze kudalwe imodeli yemeshi ecwengisiswe kakhudlwana ehlanganisa amavertices angu-17,709 (34,928 polygons).Okokugcina, imodeli yegridi yesifanekiso esihlukanisiwe ilingana nedatha yokuskena okuqondiwe ukuze ikhiqize imodeli ye-homology.Njengoba izindawo eziyingqopha-mlando zihluke kancane kulezo zedatha yokuskena okuqondiwe, imodeli ye-homology yahlelwa kahle ukuze izichaze kusetshenziswa isistimu yokuhlanganisa yokuma kwekhanda echazwe esigabeni sangaphambilini.Ibanga elimaphakathi phakathi kwezimpawu zendawo zemodeli efanayo nedatha yeskeni eliqondiwe kuwo wonke amasampuli lalingu-<0.01 mm.Kubalwe kusetshenziswa umsebenzi we-HBM-Rugle, ibanga elimaphakathi phakathi kwamaphoyinti edatha yemodeli ye-homology kanye nedatha yokuskena okuqondiwe kwaba ngu-0.322 mm (Ithebula Elingeziwe S2).
Ukuze uchaze izinguquko ku-cranial morphology, ama-vertices angu-17,709 (izixhumanisi ze-XYZ ezingu-53,127) zawo wonke amamodeli afanayo ahlaziywa ngokuhlaziywa kwengxenye eyinhloko (PCA) kusetshenziswa isofthiwe ye-HBS edalwe i-Center for Digital Human Science ku-Institute of Advanced Industrial Science and Technology., Japan (umthengisi wokusabalalisa: I-Medic Engineering, Kyoto, http://www.rugle.co.jp/).Sibe sesizama ukusebenzisa i-PCA kusethi yedatha engajwayelekile kanye nesethi yedatha eyenziwa ngokwejwayelekile ngosayizi we-centroid.Ngakho-ke, i-PCA esekelwe kudatha engajwayelekile ingabonakalisa ngokucacile ukuma kwe-cranial yamayunithi ayisishiyagalolunye ezindawo futhi yenze kube lula ukutolika kwengxenye kune-PCA isebenzisa idatha ejwayelekile.
Lesi sihloko sethula inani lezingxenye eziyinhloko ezitholiwe ngomnikelo ongaphezu kuka-1% wesamba esihlukile.Ukuze kunqunywe izingxenye eziyinhloko ezisebenza kahle kakhulu emaqenjini ahlukanisayo kuwo wonke amayunithi amakhulu ezindawo, ukuhlaziywa kwesici somamukeli (ROC) kusetshenziswe kumaphuzu engxenye eyinhloko (PC) ngomnikelo omkhulu kuno-2% 84.Lokhu kuhlaziya kudala ijika lamathuba engxenye ngayinye ye-PCA ukuze kuthuthukiswe ukusebenza kwezigaba futhi kuqhathaniswe kahle iziza phakathi kwamaqembu ezindawo.Izinga lamandla okubandlulula lingahlolwa indawo engaphansi kwejika (AUC), lapho izingxenye ze-PCA ezinamavelu amakhulu zikwazi kangcono ukubandlulula phakathi kwamaqembu.Ukuhlolwa kwe-chi-square kwase kwenziwa ukuze kuhlolwe izinga lokubaluleka.Ukuhlaziywa kwe-ROC kwenziwa ku-Microsoft Excel kusetshenziswa i-Bell Curve yesofthiwe ye-Excel (inguqulo 3.21).
Ukuze ubone ngeso lengqondo umehluko wezindawo ku-cranial morphology, ama-scatterplots adalwe kusetshenziswa amaphuzu e-PC ahlukanise ngempumelelo amaqembu kumayunithi amakhulu ezindawo.Ukuze utolike izingxenye eziyinhloko, sebenzisa imephu yombala ukuze ubone ngeso lengqondo amamodeli aqondile ahlotshaniswa kakhulu nezingxenye eziyinhloko.Ngaphezu kwalokho, izethulo ezibonakalayo zokuphela kwengxenye eyinhloko yama-axes abekwe kokuthi ±3 ukuchezuka okujwayelekile (SD) kwamaphuzu engxenye eyinhloko kubalwe futhi kwethulwa kuvidiyo yokwengeza.
I-Allometry yasetshenziselwa ukunquma ubudlelwano phakathi kokuma kogebhezi nezici zesayizi ezihlolwe ekuhlaziyweni kwe-PCA.Ukuhlaziywa kuvumeleke ezingxenyeni eziyinhloko ezineminikelo >1%.Umkhawulo owodwa wale PCA ukuthi izingxenye zomumo azikwazi ukubonisa umumo ngazodwana ngoba isethi yedatha engajwayelekile ayisusi zonke izici zobukhulu.Ngokungeziwe ekusebenziseni amasethi edatha angajwayelekile, siphinde sahlaziya amathrendi e-allometric sisebenzisa amasethi amafrakshini e-PC ngokusekelwe kudatha yosayizi we-centroid evamile esetshenziswa ezingxenyeni eziyinhloko ngeminikelo > 1%.
Amathrendi e-Allometric ahlolwe kusetshenziswa i-equation Y = aXb 85 lapho u-Y ewumumo noma ingxenye yengxenye yomumo, u-X uwusayizi oyisentimitha (Ithebula Le-Supplementary S2), a uyivelu engaguquki, futhi u-b uyi-coefficient ye-allometric.Le ndlela ngokuyisisekelo yethula izifundo zokukhula kwe-allometric ku-morphometry yejometri78,86.Ukuguqulwa kwe-logarithmic yale fomula ithi: log Y = b × log X + log a.Ukuhlaziywa kokuhlehla kusetshenziswa indlela yezikwele ezincane kakhulu kusetshenziswe ukubala u-a no-b.Uma u-Y (usayizi oyi-centroid) kanye no-X (izikolo ze-PC) eguqulwa ngokwe-logarithmically, lawa manani kufanele abe phozithivu;kodwa, isethi yezilinganiso zika-X iqukethe amanani angemuhle.Njengesixazululo, sengeze ukufinyezwa kunani eliphelele lengxenye encane kanye no-1 engxenyeni ngayinye engxenyeni ngayinye futhi sasebenzisa ukuguqulwa kwe-logarithmic kuzo zonke izingxenyana ezinhle eziguquliwe.Ukubaluleka kwama-coefficients e-allometric kwahlolwa kusetshenziswa ukuhlolwa kwe-t yoMfundi enemisila emibili.Lezi zibalo zezibalo zokuhlola ukukhula kwe-allometric zenziwe kusetshenziswa i-Bell Curves kusofthiwe ye-Excel (inguqulo 3.21).
Wolpoff, MH Imiphumela yesimo sezulu emakhaleni ohlaka lwamathambo.Yebo.J. Phys.Ubuntu.29, 405–423.https://doi.org/10.1002/ajpa.1330290315 (1968).
Ama-Beals, i-KL Head shape kanye nokucindezeleka kwesimo sezulu.Yebo.J. Phys.Ubuntu.37, 85–92.https://doi.org/10.1002/ajpa.1330370111 (1972).


Isikhathi sokuthumela: Apr-02-2024