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Lolu cwaningo luhlole ukuhlukahluka kwesifunda ku-cranial morphology yabantu kusetshenziswa imodeli ye-geometric homology esekelwe kudatha yokuskena evela emaqenjini ezinhlanga angu-148 emhlabeni jikelele. Le ndlela isebenzisa ubuchwepheshe bokufaka ithempulethi ukukhiqiza ama-homologous meshes ngokwenza izinguquko ezingaqinile kusetshenziswa i-algorithm yephuzu eliseduze eliphindaphindayo. Ngokusebenzisa ukuhlaziywa kwengxenye eyinhloko kumamodeli angu-342 akhethiwe e-homologous, kwatholakala ushintsho olukhulu ngobukhulu obuphelele futhi lwaqinisekiswa ngokusobala ngekhanda elincane elivela eNingizimu Asia. Umehluko wesibili omkhulu yisilinganiso sobude nobubanzi be-neurocranium, okubonisa umehluko phakathi kwamakhanda amade ase-Afrika kanye namakhanda aqondile ase-Northeast Asians. Kubalulekile ukuqaphela ukuthi lesi sithako asihlobene kakhulu nokwakheka kobuso. Izici zobuso ezaziwayo njengezihlathi ezivelele kuma-Northeast Asians kanye namathambo amancane aqinile kuma-Europe aqinisekisiwe. Lezi zinguquko zobuso zihlobene eduze nokwakheka kobuso, ikakhulukazi izinga lokuthambekela kwamathambo angaphambili nawase-occipital. Amaphethini e-allometric atholakale ngokulingana kobuso uma kuqhathaniswa nosayizi wobuso bonke; Ezinkalweni ezinkulu, ubuso buvame ukuba bude futhi bube buncane, njengoba kuye kwabonakala kubantu abaningi baseMelika kanye naseNyakatho-mpumalanga ye-Asia. Nakuba ucwaningo lwethu lungazange lufake idatha ngezinto eziguquguqukayo zemvelo ezingase zithonye isimo sekhanda, njengezimo zesimo sezulu noma zokudla, isethi enkulu yedatha yamaphethini ekhanda afanayo izoba usizo ekufuneni izincazelo ezahlukene zezici ze-skeletal phenotypic.
Umehluko wezindawo ngesimo sogebhezi lomuntu uye wafundwa isikhathi eside. Abacwaningi abaningi baye bahlola ukuhlukahluka kokuzivumelanisa nemvelo kanye/noma ukukhethwa kwemvelo, ikakhulukazi izici zesimo sezulu1,2,3,4,5,6,7 noma umsebenzi wokuhlafuna kuye ngezimo zokudla okunempilo5,8,9,10, 11,12. 13. . Ngaphezu kwalokho, ezinye izifundo zigxile emiphumeleni ye-bottleneck, ukukhukhuleka kwezakhi zofuzo, ukugeleza kwezakhi zofuzo, noma izinqubo zokuziphendukela kwemvelo ezingapheli ezibangelwa ukushintshashintsha kwezakhi zofuzo okungathathi hlangothi14,15,16,17,18,19,20,21,22,23. Isibonelo, ukuma okuyindilinga kwesikhala esibanzi nesifushane se-cranial kuchazwe njengokuzivumelanisa nokucindezela okukhethayo ngokusho komthetho ka-Allen24, ophakamisa ukuthi izilwane ezincelisayo zinciphisa ukulahlekelwa ukushisa ngokunciphisa indawo yomzimba uma kuqhathaniswa nevolumu2,4,16,17,25. Ngaphezu kwalokho, ezinye izifundo ezisebenzisa umthetho kaBergmann26 zichaze ubudlelwano phakathi kobukhulu bekhanda kanye nokushisa3,5,16,25,27, okuphakamisa ukuthi usayizi jikelele uvame ukuba mkhulu ezindaweni ezibandayo ukuvimbela ukulahleka kokushisa. Ithonya lobuchwepheshe lokucindezeleka kokuhlafuna ephethini yokukhula kwe-cranial vault kanye namathambo obuso liye laxoxwa maqondana nezimo zokudla ezibangelwa ukungafani kwamasiko okupheka noma okuphilayo phakathi kwabalimi kanye nabazingeli-abaqoqi8,9,11,12,28. Incazelo evamile ukuthi ukuncipha kwengcindezi yokuhlafuna kunciphisa ubulukhuni bamathambo nemisipha yobuso. Izifundo eziningana zomhlaba wonke zixhumanise ukuhlukahluka kobume bekhanda ngokuyinhloko nemiphumela ye-phenotypic yebanga lezakhi zofuzo elingathathi hlangothi kunokuba kube nokuzivumelanisa nemvelo21,29,30,31,32. Enye incazelo yezinguquko ekubumeni kwekhanda isekelwe emqondweni wokukhula kwe-isometric noma kwe-allometric6,33,34,35. Isibonelo, ubuchopho obukhulu buvame ukuba nama-frontal lobes abanzi kakhulu endaweni ebizwa ngokuthi “i-Broca's cap”, futhi ububanzi bama-frontal lobes buyakhula, inqubo yokuziphendukela kwemvelo ecatshangelwa ngokusekelwe ekukhuleni kwe-allometric. Ngaphezu kwalokho, ucwaningo oluhlola izinguquko zesikhathi eside esimweni se-skull luthole ukuthambekela kwe-allometric ekubhekeni ku-brachycephaly (ukuthambekela kwe-skull kokuba yindilinga) ngokuphakama okwandayo33.
Umlando omude wocwaningo mayelana nokwakheka kwe-cranial uhlanganisa imizamo yokuhlonza izici eziyisisekelo ezibangela izici ezahlukene zokuhlukahluka kwezimo ze-cranial. Izindlela zendabuko ezisetshenziswe ezifundweni eziningi zokuqala zazisekelwe kudatha yokulinganisa imigqa emibili, ngokuvamile kusetshenziswa izincazelo zikaMartin noma uHowell36,37. Ngesikhathi esifanayo, izifundo eziningi ezishiwo ngenhla zisebenzise izindlela ezithuthukisiwe kakhulu ezisekelwe kubuchwepheshe be-spatial 3D geometric morphometry (GM)5,7,10,11,12,13,17,20,27,34,35,38. 39. Isibonelo, indlela ye-sliding semilandmark, esekelwe ekunciphiseni amandla okugoba, ibilokhu isetshenziswa kakhulu ku-transgenic biology. Iveza i-semi-landmark yethempulethi kusampula ngayinye ngokushelela eceleni kwejika noma ubuso38,40,41,42,43,44,45,46. Kubandakanya izindlela ezinjalo zokubeka izinto endaweni ethile, izifundo eziningi ze-3D GM zisebenzisa ukuhlaziywa kwe-Procrustes okujwayelekile, i-algorithm ye-iterative nearest point (ICP) 47 ukuvumela ukuqhathanisa okuqondile kwezimo kanye nokuthwebula izinguquko. Ngaphandle kwalokho, indlela ye-thin plate spline (TPS)48,49 nayo isetshenziswa kabanzi njengendlela yokuguqula engaqinile yokudweba ukuvumelanisa kwe-semilandmark kuya ezimeni ezisekelwe ku-mesh.
Ngokuthuthukiswa kwezikena ze-3D zomzimba wonke ezingokoqobo kusukela ngasekupheleni kwekhulu lama-20, izifundo eziningi zisebenzise izikena ze-3D zomzimba wonke zokulinganisa usayizi50,51. Idatha yokuskena yasetshenziswa ukukhipha ubukhulu bomzimba, okudinga ukuchaza izimo zobuso njengezindawo kunokuba kube amafu akhombayo. Ukufakwa kwamaphethini kuyindlela ethuthukiswe ngale njongo emkhakheni wezithombe zekhompyutha, lapho isimo sobuso sichazwa khona ngemodeli ye-polygonal mesh. Isinyathelo sokuqala sokufakwa kwamaphethini ukulungiselela imodeli ye-mesh ezosetshenziswa njengethempulethi. Amanye ama-vertices akha iphethini ayizimpawu zomhlaba. Ithempulethi bese iguqulwa futhi ivumelane nobuso ukuze kuncishiswe ibanga phakathi kwethempulethi nefu lamaphuzu ngenkathi kulondolozwa izici zomumo wendawo yethempulethi. Izimpawu zomhlaba kuthempulethi zihambisana nezimpawu zomhlaba efwini lamaphuzu. Ngokusebenzisa ukufakelwa kwethempulethi, yonke idatha yokuskena ingachazwa njengemodeli ye-mesh enenani elifanayo lamaphuzu edatha kanye ne-topology efanayo. Nakuba ukuhlanganiswa okuqondile kukhona kuphela ezikhundleni zomhlaba, kungacatshangwa ukuthi kukhona ukuhlanganiswa okujwayelekile phakathi kwamamodeli akhiqizwe njengoba izinguquko ku-geometry yamathempulethi zincane. Ngakho-ke, amamodeli egridi adalwe ngokufakwa kwethempulethi ngezinye izikhathi abizwa ngokuthi amamodeli e-homology52. Inzuzo yokufakwa kwethempulethi ukuthi ithempulethi ingaguqulwa futhi ilungiswe ezingxenyeni ezahlukene zento eqondiwe eziseduze nendawo kodwa kude nayo (isibonelo, i-zygomatic arch kanye nesifunda sesikhashana sekhanda) ngaphandle kokuthintana. ukuguqulwa. Ngale ndlela, ithempulethi ingaboshwa ezintweni ezihlanganisa amagatsha njenge-torso noma ingalo, ihlombe lisesimweni sokuma. Ububi bokufakwa kwethempulethi yizindleko eziphezulu zokubala zokuphindaphinda okuphindaphindiwe, nokho, ngenxa yokuthuthuka okukhulu ekusebenzeni kwekhompyutha, lokhu akuseyona inkinga. Ngokuhlaziya amanani okuxhumanisa ama-vertices akha imodeli ye-mesh kusetshenziswa amasu okuhlaziya amaningi njengokuhlaziywa kwezingxenye eziyinhloko (i-PCA), kungenzeka ukuhlaziya izinguquko esimweni sobuso bonke kanye nesimo esibonakalayo kunoma yisiphi isikhundla ekusabalalisweni. Bala futhi ubone ngeso lengqondo53. Namuhla, amamodeli e-mesh akhiqizwa ngokufakwa kwethempulethi asetshenziswa kabanzi ekuhlaziyweni kwesimo emikhakheni ehlukahlukene52,54,55,56,57,58,59,60.
Intuthuko kwezobuchwepheshe bokuqopha i-mesh eguquguqukayo, kanye nokuthuthukiswa okusheshayo kwamadivayisi okuskena e-3D aphathekayo akwazi ukuskena ngesisombululo esiphezulu, isivinini, kanye nokuhamba kune-CT, kwenza kube lula ukuqopha idatha yendawo ye-3D kungakhathaliseki ukuthi ikuphi. Ngakho-ke, emkhakheni we-anthropology yezinto eziphilayo, ubuchwepheshe obusha obunjalo buthuthukisa ikhono lokulinganisa nokuhlaziya izibalo zamasampula abantu, kufaka phakathi amasampula ekhanda, okuyinhloso yalolu cwaningo.
Ngamafuphi, lolu cwaningo lusebenzisa ubuchwepheshe obuthuthukisiwe bokumodela i-3D homology obusekelwe ekufanisweni kwethempulethi (Isithombe 1) ukuhlola amasampula ekhanda angu-342 akhethwe kubantu abangu-148 emhlabeni jikelele ngokuqhathanisa indawo emhlabeni wonke. Ukuhlukahluka kokwakheka kwekhanda (Ithebula 1). Ukuze sibheke izinguquko ekubunjweni kwekhanda, sisebenzise ukuhlaziywa kwe-PCA kanye ne-receiver operating characteristic (ROC) kusethi yedatha yemodeli ye-homology esiyikhiqizile. Okutholakele kuzosiza ekuqondeni kangcono izinguquko zomhlaba wonke ekubunjweni kwekhanda, kufaka phakathi amaphethini esifunda kanye nokwehla kokuhlelwa koshintsho, izinguquko ezihlobene phakathi kwezingxenye zekhanda, kanye nokuba khona kwezitayela ze-allometric. Nakuba lolu cwaningo lungabheki idatha ngeziguquguquko zangaphandle ezimelelwa yizimo zesimo sezulu noma zokudla ezingase zithonye ukwakheka kwekhanda, amaphethini ezindawo okwakheka kwekhanda abhalwe ocwaningweni lwethu azosiza ukuhlola izici zemvelo, ze-biomechanical, kanye nezakhi zofuzo zokwehluka kwekhanda.
Ithebula 2 libonisa ama-eigenvalues kanye nama-coefficients omnikelo we-PCA asetshenziswe kusethi yedatha engajwayelekile yama-vertices angu-17,709 (ama-coordinate angu-53,127 XYZ) amamodeli e-skull angu-342 afanayo. Ngenxa yalokho, kwatholakala izingxenye eziyinhloko ezingu-14, umnikelo wazo ekuhlukeni okuphelele wawungaphezu kuka-1%, kanti isabelo esiphelele sokuhlukahluka sasingu-83.68%. Ama-vector okulayisha ezingxenye eziyinhloko ezingu-14 aqoshwe kuThebula Elingeziwe S1, kanti amaphuzu ezingxenye abalwe amasampula e-skull angu-342 anikezwe kuThebula Elingeziwe S2.
Lolu cwaningo luhlole izingxenye eziyisishiyagalolunye ezinkulu ezineminikelo engaphezu kuka-2%, ezinye zazo ezibonisa ukwehluka okukhulu nokuphawulekayo kwezindawo ekubunjweni kwe-cranial. Isibalo 2 ama-plot curves akhiqizwe ekuhlaziyweni kwe-ROC ukuze kuboniswe izingxenye ze-PCA ezisebenza kahle kakhulu zokuchaza noma ukuhlukanisa inhlanganisela ngayinye yamasampula kumayunithi amakhulu ezindawo (isb., phakathi kwamazwe ase-Afrika nawangewona ase-Afrika). Inhlanganisela yasePolynesia ayizange ihlolwe ngenxa yobukhulu obuncane besampula obusetshenziswe kulolu vivinyo. Idatha ephathelene nokubaluleka komehluko ku-AUC kanye nezinye izibalo eziyisisekelo ezibalwe kusetshenziswa ukuhlaziywa kwe-ROC iboniswe kuThebula Elingeziwe S3.
Ama-ROC curve asetshenziswe ekulinganisweni kwezingxenye eziyisishiyagalolunye eziyinhloko ngokusekelwe kusethi yedatha ye-vertex equkethe amamodeli angu-342 ekhanda labesilisa elifana. I-AUC: Indawo engaphansi kwe-curve ekubalulekeni okungu-0.01% esetshenziselwa ukuhlukanisa inhlanganisela ngayinye yezwe kwezinye izinhlanganisela eziphelele. I-TPF iyiqiniso futhi ilungile (ukubandlulula okuphumelelayo), i-FPF iyiqiniso futhi ilungile (ukubandlulula okungavumelekile).
Ukuchazwa kwejika le-ROC kufingqiwe ngezansi, kugxilwe kuphela ezingxenyeni ezingahlukanisa amaqembu okuqhathanisa ngokuba ne-AUC enkulu noma enkulu kakhulu kanye nezinga eliphakeme lokubaluleka elinamathuba angaphansi kuka-0.001. I-complex yaseNingizimu Asia (Isithombe 2a), equkethe ikakhulukazi amasampula avela eNdiya, ihluke kakhulu kwamanye amasampula axubene ngokwendawo ngoba ingxenye yokuqala (i-PC1) ine-AUC enkulu kakhulu (0.856) uma kuqhathaniswa nezinye izingxenye. Isici se-complex yase-Afrika (Isithombe 2b) yi-AUC enkulu kakhulu ye-PC2 (0.834). Ama-Austro-Melanesia (Isithombe 2c) abonise ukuthambekela okufanayo nama-Afrika ase-Sub-Saharan nge-PC2 ene-AUC enkulu kakhulu (0.759). Abantu baseYurophu (Umfanekiso 2d) bahluke ngokusobala ekuhlanganisweni kwe-PC2 (AUC = 0.801), i-PC4 (AUC = 0.719) kanye ne-PC6 (AUC = 0.671), isampula yaseNyakatho-mpumalanga ye-Asia (Umfanekiso 2e) ihluke kakhulu ku-PC4, eno-0.714 omkhulu, kanti umehluko kusuka ku-PC3 ubuthakathaka (AUC = 0.688). Amaqembu alandelayo nawo akhonjwe ngamanani aphansi e-AUC kanye namazinga aphezulu okubaluleka: Imiphumela ye-PC7 (AUC = 0.679), i-PC4 (AUC = 0.654) kanye ne-PC1 (AUC = 0.649) ibonise ukuthi amaNdiya aseMelika (Umfanekiso 2f) anezimpawu ezithile ezihlotshaniswa nalezi zingxenye, amaNdiya aseNingizimu-mpumalanga ye-Asia (Umfanekiso 2g) ahluke ku-PC3 (AUC = 0.660) kanye ne-PC9 (AUC = 0.663), kodwa iphethini yamasampula avela eMpumalanga Ephakathi (Umfanekiso 2h) (kufaka phakathi iNyakatho Afrika) iyafana. Uma kuqhathaniswa nezinye akukho mehluko omkhulu.
Esinyathelweni esilandelayo, ukuze kuhunyushwe ngokubonakalayo ama-vertices ahlobene kakhulu, izindawo zobuso ezinenani eliphezulu lomthwalo elingaphezu kuka-0.45 zifakwe imibala ngolwazi lwe-X, Y, kanye ne-Z coordinate, njengoba kuboniswe ku-Figure 3. Indawo ebomvu ikhombisa ukuhlangana okuphezulu nezixhumanisi ze-X-axis, okuhambisana nesiqondiso esivundlile esivundlile. Isifunda esiluhlaza sihlobene kakhulu nezixhumanisi eziqondile ze-Y axis, kanti isifunda esiluhlaza okwesibhakabhaka esimnyama sihlobene kakhulu nezixhumanisi ze-sagittal ze-Z axis. Isifunda esiluhlaza okwesibhakabhaka esikhanyayo sihlotshaniswa nama-axes e-Y coordinate kanye nama-axes e-Z coordinate; indawo ephinki - exubile ehlotshaniswa nama-axes e-X kanye ne-Z coordinate; indawo ephuzi - ehlotshaniswa nama-axes e-X kanye ne-Y coordinate; Indawo emhlophe iqukethe i-axis e-X, Y kanye ne-Z coordinate eboniswayo. Ngakho-ke, kulo mkhawulo wenani lomthwalo, i-PC 1 ihlotshaniswa kakhulu nobuso bonke bekhanda. Isimo sekhanda esibonakalayo esingu-3 SD ohlangothini oluphambene lwale axis yengxenye naso siboniswe kulesi sithombe, futhi izithombe ezisontekile zethulwa ku-Supplementary Video S1 ukuqinisekisa ngokubonakalayo ukuthi i-PC1 iqukethe izici zobukhulu bekhanda lonke.
Ukusatshalaliswa kwemvamisa yamaphuzu e-PC1 (ijika elijwayelekile lokulingana), imephu yombala wobuso bekhanda ihlobene kakhulu nama-vertices e-PC1 (incazelo yemibala ehlobene nobukhulu bezinhlangothi eziphambene zale axis ingu-3 SD. Isikali siyindilinga eluhlaza enobubanzi obungu-50 mm.
Isibalo 3 sibonisa igrafu yokusatshalaliswa kwemvamisa (ijika elijwayelekile lokulingana) lamaphuzu e-PC1 ngamanye abalwe ngokwehlukana kwamayunithi angu-9 ezindawo. Ngaphezu kokulinganiswa kwejika le-ROC (Isibalo 2), ukulinganiswa kwabantu baseNingizimu Asia ngezinga elithile kuthambekele kakhulu kwesobunxele ngoba amakhanda abo mancane kunalawo amanye amaqembu esifunda. Njengoba kuboniswe kuThebula 1, laba bantu baseNingizimu Asia bamele amaqembu ezinhlanga eNdiya okuhlanganisa iziQhingi zase-Andaman naseNicobar, eSri Lanka naseBangladesh.
I-dimensional coefficient itholakale ku-PC1. Ukutholakala kwezifunda ezixhumene kakhulu kanye nezimo ezibonakalayo kwaholela ekucacisweni kwezici zefomu zezingxenye ezingezona i-PC1; noma kunjalo, izici zobukhulu azisuswa ngokuphelele ngaso sonke isikhathi. Njengoba kuboniswe ngokuqhathanisa ama-ROC curve (Isithombe 2), i-PC2 ne-PC4 kwakuyizo ezihlukanisa kakhulu, kulandelwa i-PC6 ne-PC7. I-PC3 ne-PC9 zisebenza kahle kakhulu ekuhlukaniseni inani lesampula libe amayunithi ezindawo. Ngakho-ke, la ma-axes ezingxenye abonisa ngobuciko ama-scatterplot ezikolo ze-PC kanye nezindawo zombala ezihlobene kakhulu nengxenye ngayinye, kanye nokuguquguquka kwesimo okubonakalayo okunobukhulu bezinhlangothi eziphambene ze-3 SD (Izithombe 4, 5, 6). Ukumbozwa kwe-convex hull kwamasampula avela kuyunithi ngayinye yezindawo emelelwe kulezi zindawo cishe kungama-90%, yize kukhona izinga elithile lokufana ngaphakathi kwamaqoqo. Ithebula 3 linikeza incazelo yengxenye ngayinye ye-PCA.
Ama-scatterplot e-PC2 kanye nama-PC4 scores abantu abakhubazekile be-cranial abavela kumayunithi ayisishiyagalolunye ezindawo (phezulu) kanye namayunithi amane ezindawo (phansi), ama-plot ombala wobuso be-skull wama-vertice ahlobene kakhulu ne-PC ngayinye (uma kuqhathaniswa no-X, Y, Z). Incazelo yombala wama-axes: bheka umbhalo), kanye nokuguqulwa kwesimo esibonakalayo ezinhlangothini eziphambene zala ma-axes kungu-3 SD. Isikali siyindilinga eluhlaza enobubanzi obungu-50 mm.
Ama-scatterplot e-PC6 kanye nama-PC7 amaphuzu abantu abane-cranial abavela kumayunithi ayisishiyagalolunye e-geographic (phezulu) kanye namayunithi amabili e-geographic (phansi), ama-cranial surface color plots ama-vertice ahlobene kakhulu ne-PC ngayinye (uma kuqhathaniswa no-X, Y, Z). Incazelo yombala wama-axes: bheka umbhalo), kanye nokuguqulwa kwesimo esibonakalayo ezinhlangothini eziphambene zala ma-axes kungu-3 SD. Isikali siyindilinga eluhlaza enobubanzi obungu-50 mm.
Ama-scatterplot e-PC3 kanye nama-PC9 scores abantu abango-cranial abavela kumayunithi ayisishiyagalolunye ezindawo (phezulu) kanye namayunithi amathathu ezindawo (phansi), kanye nama-color plots ebusweni bekhanda (uma kuqhathaniswa nama-X, Y, Z axes) ama-vertice ahlobene kakhulu nokuchazwa kombala ngamunye we-PC: cm . text), kanye nokuguquguquka kwesimo okubonakalayo ezinhlangothini eziphambene zala ma-axes anobukhulu obungu-3 SD. Isikali siyindilinga eluhlaza enobubanzi obungu-50 mm.
Kugrafu ekhombisa amaphuzu e-PC2 kanye ne-PC4 (Isithombe 4, Amavidiyo Angeziwe S2, S3 abonisa izithombe ezikhubazekile), imephu yombala wobuso nayo iyaboniswa lapho umkhawulo wenani lomthwalo usethwe uphakeme kuno-0.4, okuphansi kune-PC1 ngoba inani le-PC2 umthwalo uphelele ungaphansi kune-PC1.
Ukwelulwa kwama-lobes angaphambili nawase-occipital ohlangothini lwe-sagittal eceleni kwe-Z-axis (oluhlaza okwesibhakabhaka okumnyama) kanye ne-parietal lobe ohlangothini lwe-coronal (obomvu) ku-pink), i-Y-axis ye-occiput (oluhlaza okwesibhakabhaka) kanye ne-Z-axis yebunzi (oluhlaza okwesibhakabhaka okumnyama). Le grafu ikhombisa amaphuzu abo bonke abantu emhlabeni jikelele; noma kunjalo, lapho wonke amasampula aqukethe inani elikhulu lamaqembu eboniswa ndawonye ngesikhathi esisodwa, ukuchazwa kwamaphethini okuhlakazeka kunzima kakhulu ngenxa yenani elikhulu lokugqagqana; ngakho-ke, kusukela kumayunithi amane amakhulu ezwekazi (okungukuthi, i-Afrika, i-Australasia-Melanesia, iYurophu, kanye ne-Northeast Asia), amasampula ahlakazekile ngaphansi kwegrafu ene-3 SD virtual cranial deformation ngaphakathi kwalolu hlu lwamaphuzu e-PC. Esifanekisweni, i-PC2 kanye ne-PC4 zingama-pair of scores. Abantu base-Afrika kanye nama-Austro-Melanesia badlulana kakhulu futhi basatshalaliswa ohlangothini lwesokudla, kuyilapho abantu baseYurophu besakazeke ngakwesobunxele esiphezulu kanti abantu baseNyakatho-mpumalanga base-Asia bavame ukuqoqana ngakwesobunxele esingezansi. I-axis evundlile ye-PC2 ikhombisa ukuthi abantu base-Afrika/base-Australia baseMelanesia bane-neurocranium ende kakhulu kunabanye abantu. I-PC4, lapho inhlanganisela yaseYurophu neyasenyakatho-mpumalanga ye-Asia ihlukaniswe khona ngokukhululekile, ihlotshaniswa nosayizi ohlobene kanye nokubonakala kwamathambo e-zygomatic kanye nokwakheka kwe-lateral kwe-calvarium. Uhlelo lokulinganisa lubonisa ukuthi abaseYurophu banamathambo amancane kakhulu e-maxillary kanye ne-zygomatic, isikhala esincane se-temporal fossa esinqunyelwe yi-arch ye-zygomatic, ithambo elingaphambili eliphakanyisiwe ngokuqondile kanye nethambo le-occipital eliyisicaba, eliphansi, kuyilapho abantu baseNyakatho-mpumalanga ye-Asia bevame ukuba namathambo e-zygomatic abanzi futhi avelele kakhulu. I-lobe engaphambili ithambekele, isisekelo sethambo le-occipital siphakanyisiwe.
Uma kugxilwe ku-PC6 kanye ne-PC7 (Isithombe 5) (Amavidiyo Angeziwe S4, S5 abonisa izithombe ezikhubazekile), umdwebo wombala ukhombisa umkhawulo wenani lomthwalo ongaphezu kuka-0.3, okubonisa ukuthi i-PC6 ihlotshaniswa nesimo se-maxillary noma se-alveolar (obomvu: X axis kanye nohlaza). I-Y axis), ukuma kwethambo lesikhashana (oluhlaza okwesibhakabhaka: Y kanye ne-Z axes) kanye nesimo sethambo le-occipital (opinki: X kanye ne-Z axes). Ngaphezu kobubanzi bebunzi (obomvu: X-axis), i-PC7 iphinda ihlobane nokuphakama kwe-alveoli yangaphambili ye-maxillary (oluhlaza: Y-axis) kanye nesimo sekhanda le-Z-axis nxazonke kwesifunda se-parietotemporal (oluhlaza okwesibhakabhaka olumnyama). Ephanelini eliphezulu le-Figure 5, wonke amasampula ezwe asatshalaliswa ngokuya ngamaphuzu ezingxenye ze-PC6 kanye ne-PC7. Ngenxa yokuthi i-ROC ikhombisa ukuthi i-PC6 iqukethe izici ezihlukile eYurophu kanti i-PC7 imelela izici zamaMelika Omdabu kulokhu kuhlaziywa, lawa masampula amabili esifunda adwetshwe ngokukhetha kulezi zingqimba ezimbili zezingxenye. AmaMelika Omdabu, nakuba efakwe kabanzi kusampula, ahlakazekile ekhoneni eliphezulu kwesobunxele; Ngakolunye uhlangothi, amasampula amaningi aseYurophu avame ukutholakala ekhoneni elingezansi kwesokudla. I-PC6 kanye ne-PC7 ezimbili zimelela inqubo encane ye-alveolar kanye ne-neurocranium ebanzi kakhulu yabantu baseYurophu, kuyilapho abaseMelika bebonakala ngebunzi elincane, i-maxilla enkulu, kanye nenqubo ebanzi nende ye-alveolar.
Ukuhlaziywa kwe-ROC kubonise ukuthi i-PC3 kanye/noma i-PC9 kwakuvame kakhulu kubantu baseNingizimu-mpumalanga naseNyakatho-mpumalanga ye-Asia. Ngakho-ke, amaphuzu ahlanganisa i-PC3 (ubuso obuluhlaza obuphezulu ku-y-axis) kanye ne-PC9 (ubuso obuluhlaza obuphansi ku-y-axis) (Isithombe 6; Amavidiyo Angeziwe S6, S7 anikeza izithombe eziguquguqukayo) abonisa ukuhlukahluka kwama-East Asia. , okungafani kakhulu nobukhulu bobuso bama-Northeast Asia kanye nesimo sobuso esiphansi sama-Southeast Asia. Ngaphandle kwalezi zici zobuso, esinye isici sabanye ama-Northeast Asia ukuthambekela kwe-lambda kwethambo le-occipital, kuyilapho abanye ama-Southeast Asia benesisekelo esincane sekhanda.
Incazelo engenhla yezingxenye eziyinhloko kanye nencazelo ye-PC5 kanye ne-PC8 ayifakiwe ngoba azikho izici ezithile zesifunda ezitholakale phakathi kwamayunithi ayisishiyagalolunye ayinhloko ezindawo. I-PC5 ibhekisela kusayizi wenqubo ye-mastoid yethambo lesikhashana, kanti i-PC8 ikhombisa ukungalingani kwesimo sekhanda lonke, kokubili okubonisa ukwehluka okuhambisanayo phakathi kwezinhlanganisela zesampula eziyisishiyagalolunye zezindawo.
Ngaphezu kwama-scatterplot wamaphuzu e-PCA ezingeni lomuntu ngamunye, siphinde sinikeze ama-scatterplot wezindlela zeqembu ukuze kuqhathaniswe konke. Ngenxa yalokhu, imodeli ejwayelekile ye-cranial homology yadalwa kusukela kusethi yedatha ye-vertex yamamodeli e-homology ngamanye avela emaqenjini ezinhlanga ezingu-148. Ama-Bivariate plot wamasethi wamaphuzu e-PC2 kanye ne-PC4, i-PC6 kanye ne-PC7, kanye ne-PC3 kanye ne-PC9 aboniswe ku-Supplementary Figure S1, wonke abalwa njengemodeli ye-skull evamile yesampula yabantu abangu-148. Ngale ndlela, ama-scatterplot afihla umehluko womuntu ngamunye ngaphakathi kweqembu ngalinye, okuvumela ukuchazwa okucacile kokufana kwe-skull ngenxa yokusatshalaliswa kwesifunda okuyisisekelo, lapho amaphethini afana nalawo aboniswe kuma-skull ngamanye anokugqagqana okuncane. I-Supplementary Figure S2 ibonisa imodeli ejwayelekile yeyunithi ngayinye yezwe.
Ngaphezu kwe-PC1, eyayihlotshaniswa nosayizi ojwayelekile (Ithebula Elingeziwe S2), ubudlelwano be-allometric phakathi kosayizi ojwayelekile kanye nesimo sekhanda buhlolwe kusetshenziswa ubukhulu be-centroid kanye namasethi ezilinganiso ze-PCA ezivela kudatha engeyona ejwayelekile. Ama-coefficient e-allometric, amanani angaguquki, amanani e-t, kanye namanani e-P ekuhlolweni kokubaluleka kuboniswe kuThebula 4. Azikho izingxenye ezibalulekile zephethini ye-allometric ezihlobene nosayizi wekhanda ojwayelekile ezitholakale kunoma iyiphi i-morphology ye-cranial ezingeni le-P < 0.05.
Ngenxa yokuthi ezinye izici zosayizi zingafakwa ezilinganisweni ze-PC ngokusekelwe kumasethi wedatha angajwayelekile, siphinde sahlola umkhuba we-allometric phakathi kosayizi we-centroid kanye namaphuzu e-PC abalwe kusetshenziswa amasethi wedatha ajwayelekile ngobukhulu be-centroid (imiphumela ye-PCA kanye namasethi wamaphuzu avezwe kumaThebula Angeziwe S6). , C7). Ithebula 4 likhombisa imiphumela yokuhlaziywa kwe-allometric. Ngakho-ke, kutholakale izitayela ezibalulekile ze-allometric ezingeni le-1% ku-PC6 kanye nasezingeni le-5% ku-PC10. Isithombe 7 sibonisa imithambeka yokubuyela emuva yalobu budlelwano be-log-linear phakathi kwamaphuzu e-PC kanye nosayizi we-centroid nama-dummies (±3 SD) kunoma yikuphi umkhawulo wosayizi we-log centroid. Isilinganiso se-PC6 yisilinganiso sokuphakama okuhlobene nobubanzi bekhanda. Njengoba usayizi wekhanda ukhula, ikhanda nobuso kuyaphakama, futhi ibunzi, amasokhethi amehlo kanye namakhala kuvame ukusondelana eceleni. Iphethini yokusabalala kwesampula iphakamisa ukuthi lesi silinganiso sivame ukutholakala kubantu baseNyakatho-mpumalanga base-Asia kanye naseMelika Yomdabu. Ngaphezu kwalokho, i-PC10 ikhombisa umkhuba wokunciphisa ngokulingana ububanzi be-midface kungakhathaliseki indawo.
Ngobudlelwano obubalulekile be-allometric obubhalwe kuthebula, ukuthambekela kokuhlehla kwe-log-linear phakathi kwesilinganiso se-PC sengxenye yesimo (esitholwe kudatha ejwayelekile) kanye nosayizi we-centroid, ukuguqulwa kwesimo esibonakalayo kunosayizi ongu-3 SD ngakolunye uhlangothi lomugqa ongu-4.
Iphethini elandelayo yezinguquko ekubunjweni kwe-cranial iboniswe ngokuhlaziywa kwemininingwane yamamodeli obuso be-3D afanayo. Ingxenye yokuqala ye-PCA ihlobene nosayizi we-skull iyonke. Sekuyisikhathi eside kucatshangwa ukuthi ama-skull amancane abantu baseNingizimu Asia, okuhlanganisa amasampula avela eNdiya, eSri Lanka naseziQhingini zase-Andaman, eBangladesh, abangelwa usayizi wawo omncane womzimba, okuhambisana nomthetho we-ecogeographic kaBergmann noma umthetho wesiqhingi613,5,16,25,27,62. Eyokuqala ihlobene nokushisa, kanti eyesibili incike endaweni etholakalayo kanye nezinsiza zokudla ze-niche yezemvelo. Phakathi kwezingxenye zesimo, ushintsho olukhulu kakhulu isilinganiso sobude nobubanzi be-cranial vault. Lesi sici, esibizwa ngokuthi i-PC2, sichaza ubudlelwano obuseduze phakathi kwama-skull amade ngokulinganayo ama-Austro-Melanesia nama-Afrika, kanye nomehluko ovela kuma-skull ayindilinga abanye abantu baseYurophu nabase-Northeast Asia. Lezi zici zibikwe ezifundweni eziningi zangaphambilini ngokusekelwe ekulinganisweni okulula kwemigqa37,63,64. Ngaphezu kwalokho, lesi sici sihlotshaniswa ne-brachycephaly kubantu abangewona ama-Afrika, okuxoxwe ngakho isikhathi eside ezifundweni ze-anthropometric kanye ne-osteometric. Umbono oyinhloko ngemuva kwale ncazelo ukuthi ukuncipha kokuhlafuna, njengokuncipha kwemisipha ye-temporalis, kunciphisa ingcindezi ekhanda elingaphandle5,8,9,10,11,12,13. Omunye umbono uhilela ukuzivumelanisa nezimo ezibandayo ngokunciphisa indawo yobuso bekhanda, okuphakamisa ukuthi ugebhezi oluyindilinga kakhulu lunciphisa indawo yobuso kangcono kunesimo esiyindilinga, ngokusho kwemithetho ka-Allen16,17,25. Ngokusekelwe emiphumeleni yocwaningo lwamanje, lawa ma-hypotheses angahlolwa kuphela ngokusekelwe ekuxhumaneni kwezingxenye ze-cranial. Ngamafuphi, imiphumela yethu ye-PCA ayisekeli ngokugcwele umbono wokuthi isilinganiso sobude nobubanzi be-cranial sithonywa kakhulu yizimo zokuhlafuna, njengoba umthwalo we-PC2 (ingxenye ende/ye-brachycephalic) wawungahlobene kakhulu nobukhulu bobuso (kufaka phakathi ubukhulu be-maxillary obuhlobene). kanye nendawo ehlobene ye-temporal fossa (ebonisa ivolumu yemisipha ye-temporalis). Ucwaningo lwethu lwamanje aluzange luhlaziye ubudlelwano phakathi kwesimo se-skull kanye nezimo zemvelo ze-geological ezifana nokushisa; noma kunjalo, incazelo esekelwe emthethweni ka-Allen ingase ifaneleke ukucatshangelwa njengombono ofanele ukuchaza i-brachycephalon ezindaweni ezibandayo.
Ukwehluka okuphawulekayo kwabe sekutholwa ku-PC4, okuphakamisa ukuthi abantu baseNyakatho-mpumalanga base-Asia banamathambo amakhulu, avelele e-zygomatic emathanjeni e-maxilla kanye ne-zygomatic. Lokhu okutholakele kuhambisana nesici esithile esaziwayo samaSiberia, okucatshangwa ukuthi ajwayele izimo ezibandayo kakhulu ngokuhamba phambili kwamathambo e-zygomatic, okuholela ekwandeni kwevolumu yama-sinuses kanye nobuso obuthambile 65. Okutholakele okusha okuvela kumodeli yethu efanayo ukuthi ukugoba kwezihlathi kubantu baseYurophu kuhlotshaniswa nokuthambekela okuphansi kwangaphambili, kanye namathambo e-occipital athambile futhi amancane kanye nokugoba kwe-nuchal. Ngokuphambene nalokho, abantu baseNyakatho-mpumalanga base-Asia bavame ukuba namabunzi athambile kanye nezindawo eziphakeme ze-occipital. Izifundo zethambo le-occipital zisebenzisa izindlela ze-geometric morphometric35 zibonise ukuthi izingebhezi zase-Asia nezaseYurophu zine-nuchal curve ethambile kanye nesikhundla esiphansi se-occiput uma kuqhathaniswa nabantu base-Afrika. Kodwa-ke, ama-scatterplot ethu e-PC2 kanye ne-PC4 kanye ne-PC3 kanye ne-PC9 pair abonise ukwehluka okukhulu kubantu base-Asia, kanti abaseYurophu babebonakala ngesisekelo esiyisicaba se-occiput kanye ne-occiput ephansi. Ukungahambisani kwezici zase-Asia phakathi kwezifundo kungase kube ngenxa yokwehluka kwamasampula ezizwe asetshenzisiwe, njengoba sihlole inani elikhulu lamaqembu ezizwe avela ezinhlobonhlobo eziningi zaseNyakatho-mpumalanga naseNingizimu-mpumalanga ye-Asia. Izinguquko esimweni sethambo le-occipital zivame ukuhlotshaniswa nokukhula kwemisipha. Kodwa-ke, le ncazelo eguquguqukayo ayibheki ukuhlobana phakathi kwebunzi nesimo se-occiput, okwaboniswa kulolu cwaningo kodwa akunakwenzeka ukuthi kwaboniswa ngokugcwele. Kulokhu, kufanelekile ukucabangela ubudlelwano phakathi kokulinganisela kwesisindo somzimba kanye nendawo ephakathi kwamandla adonsela phansi noma i-cervical junction (foramen magnum) noma ezinye izici.
Esinye isici esibalulekile esinokuhlukahluka okukhulu sihlobene nokuthuthukiswa kwedivayisi yokuhlafuna, emelelwa yi-maxillary kanye ne-temporal fossae, echazwa yinhlanganisela yamaphuzu e-PC6, PC7 kanye ne-PC4. Lokhu kuncipha okuphawulekayo kwezingxenye ze-cranial kuveza abantu baseYurophu ngaphezu kwanoma yiliphi elinye iqembu lendawo. Lesi sici sihunyushwe njengomphumela wokuncipha kokuzinza kokwakheka kobuso ngenxa yokuthuthukiswa kwasekuqaleni kwamasu ezolimo nokulungiselela ukudla, okwanciphisa umthwalo womshini kudivayisi yokuhlafuna ngaphandle kwedivayisi enamandla yokuhlafuna9,12,28,66. Ngokusho kombono womsebenzi wokuhlafuna, 28 lokhu kuhambisana noshintsho ekugobekeni kwesisekelo se-skull kuya ekhoneni le-cranial elibukhali kakhulu kanye nophahla lwe-cranial oluyindilinga kakhulu. Kusukela kulo mbono, abantu bezolimo bavame ukuba nobuso obuncane, ukuvela okuncane kwe-mandible, kanye ne-globular meninges. Ngakho-ke, lokhu kuguquguquka kungachazwa ngohlaka olujwayelekile lwesimo se-lateral se-skull yabantu baseYurophu abanezitho zokuhlafuna ezincishisiwe. Kodwa-ke, ngokusho kwalolu cwaningo, lokhu kuchazwa kuyinkimbinkimbi ngoba ukubaluleka kokusebenza kobudlelwano besimo phakathi kwe-globose neurocranium kanye nokuthuthukiswa kwedivayisi yokuhlafuna akwamukeleki kangako, njengoba kucatshangelwe ekuchazeni kwangaphambilini kwe-PC2.
Umehluko phakathi kwabantu baseNyakatho-mpumalanga ye-Asia kanye nabantu baseNingizimu-mpumalanga ye-Asia uboniswa umehluko phakathi kobuso obude obunethambo le-occipital elithambekile kanye nobuso obufushane obunesisekelo esincane sogebhezi, njengoba kuboniswe ku-PC3 kanye ne-PC9. Ngenxa yokuntuleka kwedatha ye-geoecological, ucwaningo lwethu lunikeza incazelo elinganiselwe kuphela yalokhu okutholakele. Incazelo engaba khona ukuzivumelanisa nesimo sezulu noma izimo zokudla ezihlukile. Ngaphezu kokuzivumelanisa nemvelo, umehluko wendawo emlandweni wabantu baseNyakatho-mpumalanga naseNingizimu-mpumalanga ye-Asia nawo wacatshangelwa. Isibonelo, empumalanga ye-Eurasia, imodeli enezingqimba ezimbili iye yacatshangelwa ukuthi iyaqonda ukusabalala kwabantu besimanje ngokwe-anatomically (AMH) ngokusekelwe kudatha ye-cranial morphometric67,68. Ngokusho kwale modeli, "isigaba sokuqala", okungukuthi, amaqembu okuqala amakoloni e-Late Pleistocene AMH, ayenokwehla okuqondile okuvela kubantu bomdabu balesi sifunda, njengabase-Austro-Melanesia banamuhla (ikhasi lokuqala). , futhi kamuva wabhekana nokuhlanganiswa okukhulu kwabantu bezolimo basenyakatho abanezimpawu zasenyakatho-mpumalanga ye-Asia (ungqimba lwesibili) esifundeni (cishe eminyakeni engu-4,000 eyedlule). Ukugeleza kwezakhi zofuzo okuhlelwe kusetshenziswa imodeli "yezingqimba ezimbili" kuzodingeka ukuze kuqondwe ukuma kwe-cranial yaseNingizimu-mpumalanga ye-Asia, njengoba isimo se-cranial yaseNingizimu-mpumalanga ye-Asia singase sincike ngokwengxenye efeni lezakhi zofuzo lendawo lezinga lokuqala.
Ngokuhlola ukufana kwe-cranial sisebenzisa amayunithi endawo ahlelwe kusetshenziswa amamodeli afanayo, singaphetha umlando wabantu abangaphansi kwe-AMF ezimweni ezingaphandle kwe-Afrika. Kuye kwaphakanyiswa amamodeli amaningi ahlukene "angaphandle kwe-Afrika" ukuchaza ukusatshalaliswa kwe-AMF ngokusekelwe kudatha yamathambo kanye ne-genomic. Kulawa, izifundo zakamuva ziphakamisa ukuthi ukwakhiwa kwezindawo ezingaphandle kwe-Afrika yi-AMH kwaqala cishe eminyakeni eyi-177,000 eyedlule69,70. Kodwa-ke, ukusatshalaliswa kwe-AMF ebangeni elide e-Eurasia phakathi nalesi sikhathi kusalokhu kungaqinisekile, njengoba izindawo zokuhlala zalezi zinsalela zokuqala zikhawulelwe eMpumalanga Ephakathi naseMedithera eduze ne-Afrika. Icala elilula kakhulu ukuhlala ndawonye endleleni yokufuduka esuka e-Afrika iye e-Eurasia, idlula izithiyo zendawo njengeHimalayas. Enye imodeli iphakamisa amagagasi amaningi okufuduka, elokuqala lawo lasakazeka lisuka e-Afrika ogwini lolwandle i-Indian Ocean liya eNingizimu-mpumalanga ye-Asia nase-Australia, bese lisakazeka enyakatho ye-Eurasia. Iningi lalezi zifundo liqinisekisa ukuthi i-AMF yasakazeka kude kakhulu ne-Afrika eminyakeni engaba ngu-60,000 eyedlule. Kulokhu, amasampula ase-Australasian-Melanesia (kufaka phakathi iPapua) abonisa ukufana okukhulu namasampula ase-Afrika kunanoma yiluphi olunye uchungechunge lwezindawo ekuhlaziyweni kwezingxenye eziyinhloko zamamodeli e-homology. Lokhu okutholakele kusekela umbono wokuthi amaqembu okuqala okusatshalaliswa kwe-AMF eceleni komphetho oseningizimu ye-Eurasia avela ngqo e-Afrika22,68 ngaphandle kwezinguquko ezibalulekile zesimo sengqondo ekuphenduleni isimo sezulu esithile noma ezinye izimo ezibalulekile.
Ngokuphathelene nokukhula kwe-allometric, ukuhlaziywa kusetshenziswa izingxenye zesimo ezithathwe kusethi yedatha ehlukile ejwayeleke ngobukhulu be-centroid kubonise ukuthambekela okuphawulekayo kwe-allometric ku-PC6 kanye ne-PC10. Zombili izingxenye zihlobene nokuma kwebunzi kanye nezingxenye zobuso, okuba mncane njengoba usayizi wekhanda ukhula. Abantu baseNyakatho-mpumalanga base-Asia kanye nabaseMelika bavame ukuba nalesi sici futhi banezikhumba ezinkulu kakhulu. Lokhu okutholakele kuphikisana namaphethini e-allometric abikwe ngaphambilini lapho ubuchopho obukhulu bunezikhumba zangaphambili ezibanzi kakhulu esifundeni esibizwa ngokuthi "i-Broca's cap", okuholela ekwandeni kobubanzi be-lobe yangaphambili34. Lokhu kungafani kuchazwa ngomehluko kumasethi esampula; Ucwaningo lwethu luhlaziye amaphethini e-allometric osayizi we-cranial jikelele kusetshenziswa abantu banamuhla, futhi izifundo zokuqhathanisa zibhekana nezitayela zesikhathi eside ekuziphendukeleni kwabantu okuhlobene nosayizi wobuchopho.
Ngokuphathelene ne-allometric yobuso, olunye ucwaningo olusebenzisa idatha ye-biometric78 luthole ukuthi ukuma kobuso nosayizi kungase kuhlobane kancane, kanti ucwaningo lwethu luthole ukuthi ama-skulls amakhulu avame ukuhlotshaniswa nobuso obude nobuncane. Kodwa-ke, ukuvumelana kwedatha ye-biometric akucaci; Ukuhlolwa kokuhlehla okuqhathanisa i-ontogenetic allometry kanye ne-static allometry kukhombisa imiphumela ehlukile. Ukuthambekela kwe-allometric ekubunjweni kwe-skull eyindilinga ngenxa yokuphakama okuphezulu nakho kubikiwe; noma kunjalo, asizange sihlaziye idatha yokuphakama. Ucwaningo lwethu lukhombisa ukuthi ayikho idatha ye-allometric ekhombisa ubudlelwano phakathi kwezilinganiso ze-cranial globular kanye nosayizi we-cranial iyonke.
Nakuba ucwaningo lwethu lwamanje lungabhekani nedatha ngeziguquguquko zangaphandle ezimelelwa yizimo zesimo sezulu noma zokudla okungenzeka zithonye ukwakheka kwe-cranial, isethi enkulu yedatha yamamodeli obuso be-cranial obufanayo obusetshenziswe kulolu cwaningo izosiza ekuhloleni ukuguquguquka kokwakheka kwe-phenotypic okuhlobene. Izici zemvelo ezifana nokudla, isimo sezulu kanye nezimo zokudla okunempilo, kanye namandla angathathi hlangothi njengokufuduka, ukugeleza kwezakhi zofuzo kanye nokushintshashintsha kwezakhi zofuzo.
Lolu cwaningo luhlanganise amasampula angu-342 ezinkakha zamadoda aqoqwe kubantu abangu-148 kumayunithi angu-9 ezindawo (Ithebula 1). Amaqembu amaningi ayizibonelo zezindawo, kanti amanye amaqembu e-Afrika, eNyakatho-mpumalanga/eNingizimu-mpumalanga ye-Asia naseMelika (abhalwe ngo-italics) achazwe ngokohlanga. Amasampula amaningi ezinkakha zakhethwa kudathabheyisi yokulinganisa i-cranial ngokwencazelo yokulinganisa i-cranial kaMartin enikezwe nguTsunehiko Hanihara. Sikhethe amakhanda abesilisa amele wonke amaqembu ezinhlanga emhlabeni. Ukuze sithole amalungu eqembu ngalinye, sibale amabanga e-Euclidean ngokusekelwe ezilinganisweni ze-cranial ezingu-37 ezivela eqenjini eliphakathi kwabo bonke abantu abakulelo qembu. Ezimweni eziningi, sikhethe amasampula angu-1-4 anebanga elincane kakhulu ukusuka ku-average (Ithebula Elingeziwe S4). Kula maqembu, amanye amasampula akhethwe ngokungahleliwe uma engafakwanga ohlwini lwedathabheyisi yokulinganisa i-Hahara.
Ukuze kuqhathaniswe izibalo, amasampula abantu abangu-148 ahlanganiswe ngamayunithi amakhulu ezindawo, njengoba kuboniswe kuThebula 1. Iqembu "lase-Afrika" lakhiwe ngamasampula avela esifundeni sase-sub-Sahara kuphela. Amasampula avela eNyakatho Afrika afakwe "eMpumalanga Ephakathi" kanye namasampula avela eNtshonalanga Asia anezimo ezifanayo. Iqembu laseNyakatho-mpumalanga ye-Asia lihlanganisa kuphela abantu abangebona abaseYurophu, kanti iqembu laseMelika lihlanganisa amaMelika Omdabu kuphela. Ikakhulukazi, leli qembu lisatshalaliswa endaweni enkulu yamazwekazi aseNyakatho naseNingizimu Melika, ezindaweni ezahlukahlukene. Kodwa-ke, sicabangela isampula yase-US ngaphakathi kwale yunithi eyodwa yezindawo, uma sibheka umlando wabantu baseMelika Bomdabu ababhekwa njengabavela eNyakatho-mpumalanga ye-Asia, kungakhathaliseki ukufuduka okuningi 80.
Siqophe idatha yobuso be-3D yalezi zibonelo zekhanda ezihlukile sisebenzisa isithwebuli se-3D esinesinqumo esiphezulu (i-EinScan Pro yi-Shining 3D Co Ltd, isinqumo esincane: 0.5 mm, https://www.shining3d.com/) sabe sesikhiqiza i-mesh. Imodeli ye-mesh iqukethe ama-vertices angaba ngu-200,000–400,000, futhi isofthiwe efakiwe isetshenziselwa ukugcwalisa imigodi kanye nemiphetho ebushelelezi.
Esigabeni sokuqala, sisebenzise idatha yokuskena kunoma yiliphi i-skull ukudala imodeli ye-skull ene-template eyodwa equkethe ama-vertices angu-4485 (ubuso obunama-polygonal angu-8728). Isisekelo sesifunda se-skull, esakhiwe yithambo le-sphenoid, ithambo le-petrous temporal, i-palate, i-maxillary alveoli, namazinyo, sasuswa kumodeli ye-template mesh. Isizathu ukuthi lezi zakhiwo ngezinye izikhathi aziphelele noma kunzima ukuziqeda ngenxa yezingxenye ezincane noma ezincane ezibukhali njengezindawo ze-pterygoid kanye nezinqubo ze-styloid, ukuguguleka kwamazinyo kanye/noma isethi yamazinyo engaguquki. Isisekelo se-skull esizungeze i-foramen magnum, kufaka phakathi isisekelo, asizange sisuswe ngoba lena indawo ebalulekile ngokwesimo sendawo yamalunga omlomo wesibeletho futhi ukuphakama kwe-skull kumele kuhlolwe. Sebenzisa izindandatho zesibuko ukwakha ithempulethi elilingana nhlangothi zombili. Yenza i-isotropic meshing ukuguqula izimo ze-polygonal zibe ngokulinganayo ngangokunokwenzeka.
Okulandelayo, izindawo ezingu-56 zabelwa ama-vertices ahambisana nokwakheka kwemodeli yethempulethi kusetshenziswa isofthiwe ye-HBM-Rugle. Izilungiselelo zezindawo eziphawulekayo ziqinisekisa ukunemba nokuzinza kokuma kwezindawo eziphawulekayo futhi ziqinisekisa ukulingana kwalezi zindawo kumodeli yezindawo ezihlanganisiwe. Zingabonakala ngokusekelwe ezicini zazo ezithile, njengoba kuboniswe kuThebula Elingeziwe S5 kanye noMfanekiso Ongeziwe S3. Ngokusho kwencazelo kaBookstein81, iningi lalezi zindawo eziphawulekayo ziyizindawo eziphawulekayo zohlobo I ezitholakala ekuxhumaneni kwezakhiwo ezintathu, kanti ezinye ziyizindawo eziphawulekayo zohlobo II ezinamaphuzu aphezulu okugoba. Izindawo eziningi eziphawulekayo zadluliselwa kusuka kumaphuzu achazwe ngezilinganiso ze-cranial eziqondile encazelweni kaMartin 36. Sichaze izindawo ezifanayo ezingu-56 zamamodeli askeniwe amasampula ekhanda angu-342, anikezwe ngesandla ama-vertices ahambisana nokwakheka kwezindawo ukuze akhiqize amamodeli ezindawo ezinembile kakhulu esigabeni esilandelayo.
Uhlelo lokuxhumanisa olugxile ekhanda luchazwe ukuchaza idatha yokuskena kanye nethempulethi, njengoba kuboniswe ku-Supplementary Figure S4. Indiza ye-XZ iyindiza evundlile yaseFrankfurt edlula endaweni ephakeme kakhulu (incazelo kaMartin: ingxenye) yomphetho ophakeme wemisele yangaphandle yesobunxele nesokudla kanye nendawo ephansi kakhulu (incazelo kaMartin: i-orbit) yomphetho ongezansi womjikelezo wesobunxele. . I-X axis ingumugqa oxhumanisa izinhlangothi zesobunxele nesokudla, kanti i-X+ uhlangothi lwesokudla. Indiza ye-YZ idlula phakathi kwezingxenye zesobunxele nesokudla kanye nomsuka wekhala: Y+ phezulu, Z+ phambili. Iphuzu lokubhekisela (umsuka: i-zero coordinate) libekwe endaweni yokuhlangana kwendiza ye-YZ (midplane), indiza ye-XZ (Frankfort plane) kanye nendiza ye-XY (coronal plane).
Sisebenzise isofthiwe ye-HBM-Rugle (Medic Engineering, Kyoto, http://www.rugle.co.jp/) ukudala imodeli ye-homologous mesh ngokwenza ukufakwa kwethempulethi sisebenzisa amaphuzu angu-56 omlando (ohlangothini lwesobunxele lwesithombe 1). Ingxenye eyinhloko yesofthiwe, eyasungulwa ekuqaleni yi-Center for Digital Human Research e-Institute of Advanced Industrial Science and Technology eJapane, ibizwa ngokuthi i-HBM futhi inemisebenzi yokufaka amathempulethi kusetshenziswa izimpawu zendawo kanye nokudala amamodeli amahle omlando kusetshenziswa izindawo zokuhlukanisa82. Inguqulo yesofthiwe elandelayo (mHBM) 83 yengeze isici sokufakwa kwephethini ngaphandle kwezimpawu zendawo ukuze kuthuthukiswe ukusebenza kokufakwa. I-HBM-Rugle ihlanganisa isofthiwe ye-mHBM nezici ezengeziwe ezisebenziseka kalula ezifaka phakathi ukwenza ngezifiso izinhlelo zokuxhumanisa kanye nokushintsha usayizi wedatha yokufaka. Ukuthembeka kokunemba kokufakwa kwesofthiwe kuqinisekiswe ezifundweni eziningi52,54,55,56,57,58,59,60.
Uma kufakwa ithempulethi ye-HBM-Rugle kusetshenziswa izimpawu zomlando, imodeli ye-mesh yethempulethi ifakwa phezu kwedatha yokuskena okuqondiwe ngokubhaliswa okuqinile okusekelwe kubuchwepheshe be-ICP (ukunciphisa inani lamabanga phakathi kwezimpawu zomlando ezihambisana nethempulethi kanye nedatha yokuskena okuqondiwe), bese ngokuguqulwa okungaguquki kwe-mesh kuvumelanisa ithempulethi nedatha yokuskena okuqondiwe. Le nqubo yokufakela iphindwe kathathu kusetshenziswa amanani ahlukene amapharamitha amabili okufaka ukuze kuthuthukiswe ukunemba kokufakwa. Enye yalezi pharamitha ikhawulela ibanga phakathi kwemodeli yegridi yethempulethi kanye nedatha yokuskena okuqondiwe, kanti enye ijezisa ibanga phakathi kwezimpawu zomlando zethempulethi kanye nezimpawu zomlando eziqondiwe. Imodeli ye-mesh yethempulethi ekhubazekile yabe isihlukaniswa kusetshenziswa i-algorithm yokuhlukaniswa kwendawo ejikelezayo engu-82 ukudala imodeli ye-mesh ecwengekile kakhulu equkethe ama-vertices angu-17,709 (ama-polygon angu-34,928). Okokugcina, imodeli yegridi yethempulethi ehlukanisiwe ifanelana nedatha yokuskena okuqondiwe ukuze kukhiqizwe imodeli ye-homology. Njengoba izindawo eziphawulekayo zihlukile kancane kulezo ezikudatha yokuskena okuqondiwe, imodeli ye-homology yalungiswa kahle ukuze ichazwe kusetshenziswa uhlelo lokuqondisa ikhanda oluchazwe esigabeni esedlule. Ibanga elimaphakathi phakathi kwezimpawu eziphawulekayo zemodeli efanayo kanye nedatha yokuskena okuqondiwe kuzo zonke izibonelo laliyi-<0.01 mm. Kubalwa kusetshenziswa umsebenzi we-HBM-Rugle, ibanga elimaphakathi phakathi kwamaphuzu edatha yemodeli ye-homology kanye nedatha yokuskena okuqondiwe laliyi-0.322 mm (Ithebula Elingeziwe S2).
Ukuze sichaze izinguquko ku-cranial morphology, ama-vertices angu-17,709 (ama-coordinate angu-53,127 XYZ) awo wonke amamodeli afanayo ahlaziywe ngokuhlaziywa kwe-principal component (PCA) kusetshenziswa isofthiwe ye-HBS eyenziwe yi-Center for Digital Human Science e-Institute of Advanced Industrial Science and Technology. , eJapane (umthengisi wokusabalalisa: Medic Engineering, Kyoto, http://www.rugle.co.jp/). Sabe sesizama ukusebenzisa i-PCA kusethi yedatha engajwayelekile kanye nesethi yedatha ejwayelekile ngobukhulu be-centroid. Ngakho-ke, i-PCA esekelwe kudatha engajwayelekile ingachaza ngokucacile ukuma kwe-cranial kwamayunithi ayisishiyagalolunye ezindawo futhi ilula ukuhumusha kwengxenye kune-PCA kusetshenziswa idatha ejwayelekile.
Lesi sihloko sinikeza inani lezingxenye eziyinhloko ezitholiwe ngomnikelo ongaphezu kuka-1% womehluko ophelele. Ukuze kutholakale izingxenye eziyinhloko ezisebenza kahle kakhulu ekuhlukaniseni amaqembu kuwo wonke amayunithi amakhulu ezindawo, ukuhlaziywa kwesici sokusebenza kwesamukeli (i-ROC) kusetshenziswe kumaphuzu ezingxenye eziyinhloko (i-PC) ngomnikelo omkhulu kuno-2% 84. Lokhu kuhlaziywa kudala ijika elinokwenzeka lengxenye ngayinye ye-PCA ukuze kuthuthukiswe ukusebenza kokuhlukaniswa futhi kuqhathaniswe kahle ama-plots phakathi kwamaqembu ezindawo. Izinga lamandla okubandlulula lingahlolwa ngendawo engaphansi kwejika (i-AUC), lapho izingxenye ze-PCA ezinamanani amakhulu zikwazi kangcono ukuhlukanisa phakathi kwamaqembu. Ukuhlolwa kwe-chi-square kwenziwa ukuze kuhlolwe izinga lokubaluleka. Ukuhlaziywa kwe-ROC kwenziwa ku-Microsoft Excel kusetshenziswa isofthiwe ye-Bell Curve ye-Excel (inguqulo 3.21).
Ukuze ubone umehluko wezindawo ku-cranial morphology, ama-scatterplot adalwe kusetshenziswa amaphuzu e-PC ahlukanisa amaqembu kahle kakhulu kumayunithi amakhulu ezindawo. Ukuze uhumushe izingxenye eziyinhloko, sebenzisa imephu yombala ukuze ubone ama-vertices emodeli ahlobene kakhulu nezingxenye eziyinhloko. Ngaphezu kwalokho, ukumelwa okubonakalayo kweziphetho zama-axes ezingxenye eziyinhloko ezitholakala ku-±3 standard deviation (SD) yamaphuzu ezingxenye eziyinhloko kubalwe futhi kwethulwa kuvidiyo eyengeziwe.
I-Allometry yasetshenziswa ukunquma ubudlelwano phakathi kwezici zokwakheka kwekhanda nosayizi ezihlolwe ekuhlaziyweni kwe-PCA. Ukuhlaziywa kusebenza ezingxenyeni eziyinhloko ezineminikelo >1%. Omunye umkhawulo wale PCA ukuthi izingxenye zokwakheka azikwazi ukukhombisa isimo ngasinye ngoba isethi yedatha engajwayelekile ayisusi zonke izici zobukhulu. Ngaphezu kokusebenzisa amasethi edatha angajwayelekile, siphinde sahlaziya izitayela ze-allometric sisebenzisa amasethi e-PC fraction ngokusekelwe kudatha yobukhulu be-centroid ejwayelekile esetshenziswa ezingxenyeni eziyinhloko ezineminikelo >1%.
Amathrendi e-allometric ahlolwe kusetshenziswa i-equation Y = aXb 85 lapho u-Y eyisimo noma ingxenye yengxenye yesimo, u-X usayizi we-centroid (Ithebula Elingeziwe S2), u-a uyinani elingaguquki, kanti u-b uyi-coefficient ye-allometric. Le ndlela ngokuyisisekelo yethula izifundo zokukhula kwe-allometric ku-geometric morphometry78,86. Ukuguqulwa kwe-logarithmic kwale fomula yilokhu: i-log Y = b × i-log X + i-log a. Ukuhlaziywa kokuhlehla kusetshenziswa indlela yezikwele ezincane kusetshenziswe ukubala u-a no-b. Lapho u-Y (usayizi we-centroid) no-X (amaphuzu e-PC) beguqulwa ngokwe-logarithmic, lawa manani kumele abe mahle; noma kunjalo, isethi yezilinganiso ze-X iqukethe amanani amabi. Njengesixazululo, sengeze ukuzungeza enanini eliphelele lengxenye encane kakhulu kanye no-1 yengxenye ngayinye engxenyeni ngayinye futhi sasebenzisa ukuguqulwa kwe-logarithmic kuzo zonke izingxenyana ezinhle eziguquliwe. Ukubaluleka kwama-coefficients e-allometric kwahlolwa kusetshenziswa ukuhlolwa kwe-t komfundi okunemisila emibili. Lezi zibalo zezibalo zokuhlola ukukhula kwe-allometric zenziwa kusetshenziswa ama-Bell Curves kusofthiwe ye-Excel (inguqulo 3.21).
Wolpoff, MH Imiphumela yesimo sezulu emakhaleni ethambo. Yebo. J. Phys. Humanity. 29, 405–423. https://doi.org/10.1002/ajpa.1330290315 (1968).
UBeals, KL Ukuma kwekhanda kanye nokucindezeleka kwesimo sezulu. Yebo. J. Phys. Humanity. 37, 85–92. https://doi.org/10.1002/ajpa.1330370111 (1972).
Isikhathi sokuthunyelwe: Ephreli-02-2024