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Olu phononongo luvavanye umahluko wengingqi kwimo ye-cranial yomntu kusetyenziswa imodeli ye-homology yejiyometri esekwe kwidatha yeskeni evela kumaqela eentlanga ali-148 kwihlabathi liphela. Le ndlela isebenzisa itekhnoloji yokufakela itemplate ukuvelisa i-homologous meshes ngokwenza utshintsho olungenabugocigoci kusetyenziswa i-algorithm yenqaku elisondeleyo eliphindaphindayo. Ngokusebenzisa uhlalutyo lwecandelo eliphambili kwiimodeli ze-homologous ezikhethiweyo ezingama-342, utshintsho olukhulu kubukhulu ngokubanzi lufunyenwe kwaye lwaqinisekiswa ngokucacileyo kwi-skull encinci evela eMzantsi Asia. Umahluko wesibini omkhulu ngumlinganiselo wobude nobubanzi be-neurocranium, obonisa umahluko phakathi kwee-skulls ezinde zama-Afrika kunye nee-skulls ezingqukuva zama-Asia aseMpuma-mpuma. Kubalulekile ukuqaphela ukuba esi sithako asinanto yakwenza nokuqunjelwa kobuso. Iimpawu zobuso ezaziwayo ezifana nezidlele ezivele kuma-Asia aseMpuma-mpuma kunye namathambo aqinileyo emaxillary kumaYurophu aqinisekisiwe. Olu tshintsho lobuso lunxulumene kakhulu nokwakheka kwe-skull, ngakumbi inqanaba lokuthambekela kwamathambo angaphambili kunye nama-occipital. Iipateni ze-allometric zifunyenwe ngobungakanani bobuso xa kuthelekiswa nobukhulu be-skull iyonke; Kwii-skulls ezinkulu, ubuso budla ngokuba bude kwaye bunciphe, njengoko kubonisiwe kubantu abaninzi baseMelika nakuma-Asia aseMantla-mpuma. Nangona uphando lwethu lungakhange lubandakanye idatha ngezinto eziguquguqukayo ezingqongileyo ezinokuchaphazela imo ye-cranial, njengemozulu okanye iimeko zokutya, iseti enkulu yedatha yeepateni ze-cranial ezifanayo ziya kuba luncedo ekufuneni iingcaciso ezahlukeneyo zeempawu ze-skeletal phenotypic.
Umahluko wejografi kwimilo yentloko yomntu uye wafundwa ixesha elide. Abaphandi abaninzi baye bavavanya ukwahluka kokuguquguquka kwendalo kunye/okanye ukukhethwa kwendalo, ngakumbi izinto zemozulu1,2,3,4,5,6,7 okanye umsebenzi wokuhlafuna ngokuxhomekeke kwiimeko zesondlo5,8,9,10, 11,12. 13. Ukongeza, ezinye izifundo zigxile kwimiphumo ye-bottleneck, i-genetic drift, ukuhamba kwezakhi zofuzo, okanye iinkqubo zokuzivelela ezibangelwa kukuguquka kwezakhi zofuzo okungathathi cala14,15,16,17,18,19,20,21,22,23. Umzekelo, imilo engqukuva ye-cranial vault ebanzi nemfutshane ichazwe njengokulungelelaniswa noxinzelelo olukhethiweyo ngokomthetho ka-Allen24, ocinga ukuba izilwanyana ezincancisayo zinciphisa ukulahleka kobushushu ngokunciphisa indawo yomzimba xa kuthelekiswa nomthamo2,4,16,17,25. Ukongeza, ezinye izifundo ezisebenzisa umthetho kaBergmann26 zichaze ubudlelwane phakathi kobukhulu be-skull kunye nobushushu3,5,16,25,27, nto leyo ebonisa ukuba ubukhulu ngokubanzi buhlala bukhulu kwiindawo ezibandayo ukuthintela ukulahleka kobushushu. Impembelelo yoxinzelelo lokuhlafuna kwindlela yokukhula kwe-cranial vault kunye namathambo obuso iye yaxoxwa ngokunxulumene neemeko zokutya ezibangelwa yinkcubeko yokupheka okanye umahluko wokuphila phakathi kwabalimi kunye nabazingeli-abaqokeleli8,9,11,12,28. Ingcaciso ngokubanzi kukuba uxinzelelo oluphantsi lokuhlafuna lunciphisa ubunzima bamathambo kunye nezihlunu zobuso. Izifundo ezininzi zehlabathi zidibanise ukwahluka kwesimo se-skull ngokuyintloko kwimiphumo ye-phenotypic yomgama we-genetic ongathathi hlangothi endaweni yokulungelelaniswa kwendalo21,29,30,31,32. Enye ingcaciso yotshintsho kwisimo se-skull isekelwe kwingcamango yokukhula kwe-isometric okanye i-allometric6,33,34,35. Umzekelo, iingqondo ezinkulu zihlala zineendawo ezibanzi ngaphambili kwindawo ebizwa ngokuba yi-"Broca's cap", kwaye ububanzi beendawo ezingaphambili buyanda, inkqubo yokuzivelela ethathwa njengesekelwe ekukhuleni kwe-allometric. Ukongeza, uphando oluhlola utshintsho lwexesha elide kwimo ye-skull lufumene ukuthambekela kwe-allometric kwi-brachycephaly (ukuthambekela kwe-skull ukuba ibe yi-spherical) ngokuphakama okwandayo33.
Imbali ende yophando malunga nemo ye-cranial ibandakanya imizamo yokuchonga izinto ezisisiseko ezibangela iinkalo ezahlukeneyo zokwahluka kweemilo ze-cranial. Iindlela zemveli ezisetyenzisiweyo kwizifundo ezininzi zokuqala zazisekelwe kwidatha yokulinganisa umgca we-bivariate, rhoqo kusetyenziswa iinkcazo zikaMartin okanye uHowell36,37. Kwangaxeshanye, uninzi lwezifundo ezikhankanyiweyo apha ngasentla zisebenzise iindlela eziphucukileyo ezisekelwe kwiteknoloji ye-spatial 3D geometric morphometry (GM)5,7,10,11,12,13,17,20,27,34,35,38. 39. Umzekelo, indlela ye-sliding semilandmark, esekelwe ekunciphiseni amandla okugoba, iye yaba yeyona ndlela isetyenziswa kakhulu kwi-transgenic biology. Ibonisa ii-semi-landmarks zetemplate kwisampulu nganye ngokutyibiliza ecaleni kwegophe okanye kumphezulu38,40,41,42,43,44,45,46. Kuquka ezo ndlela zokubeka izinto phezulu, uninzi lwezifundo ze-3D GM zisebenzisa uhlalutyo lwe-Procrustes oluqhelekileyo, i-algorithm ye-iterative nearest point (ICP) 47 ukuvumela uthelekiso oluthe ngqo lweemilo kunye nokubamba utshintsho. Kungenjalo, indlela ye-thin plate spline (TPS)48,49 ikwasetyenziswa kakhulu njengendlela yokuguqula engaqinanga yokubonisa ulungelelwaniso lwe-semilandmark kwimilo esekelwe kwi-mesh.
Ngophuhliso lwezikena ze-3D zomzimba wonke ezisebenzisekayo ukususela ngasekupheleni kwenkulungwane yama-20, izifundo ezininzi zisebenzise izikena ze-3D zomzimba wonke zokulinganisa ubungakanani50,51. Idatha yokuskena yasetyenziswa ukukhupha ubungakanani bomzimba, nto leyo efuna ukuchaza iimilo zomphezulu njengeendawo ezingaphezulu kuneengqimba zamafu. Ukufakelwa kweepateni yindlela eyenzelwe le njongo kwicandelo lemizobo yekhompyutha, apho imilo yomphezulu ichazwa yimodeli ye-polygonal mesh. Inyathelo lokuqala lokufakelwa kweepateni kukulungiselela imodeli ye-mesh ukuyisebenzisa njengetemplate. Ezinye zee-vertices ezenza ipateni ziimpawu zomhlaba. Emva koko itemplate iyaguqulwa kwaye ihambelane nomphezulu ukunciphisa umgama phakathi kwetemplate kunye nelifu le-point ngelixa kugcinwa iimpawu zemilo yendawo yetemplate. Iimpawu zomhlaba kwitemplate zihambelana neempawu zomhlaba kwilifu le-point. Ukusebenzisa ukufakelwa kwetemplate, yonke idatha yokuskena ingachazwa njengemodeli ye-mesh enenani elifanayo lamanqaku edatha kunye ne-topology efanayo. Nangona ukuhambelana okuchanekileyo kukho kuphela kwiindawo zomhlaba, kunokucingelwa ukuba kukho ukuhambelana ngokubanzi phakathi kweemodeli ezivelisiweyo kuba utshintsho kwi-geometry yeetemplate luncinci. Ke ngoko, iimodeli zegridi ezenziwe ngokufaka itemplate ngamanye amaxesha zibizwa ngokuba ziimodeli ze-homology52. Inzuzo yokufaka itemplate kukuba itemplate inokuguqulwa kwaye ilungiswe kwiindawo ezahlukeneyo zento ekujoliswe kuyo ezikufutshane nendawo kodwa kude nayo (umzekelo, i-zygomatic arch kunye nommandla wexesha lekhanda) ngaphandle kokuchaphazela omnye nomnye. ukuguqulwa. Ngale ndlela, itemplate inokuxhonywa kwizinto ezidibanisa amasebe ezifana ne-torso okanye ingalo, kunye negxalaba likwindawo yokuma. Ingxaki yokufaka itemplate yindleko ephezulu yokubala yokuphindaphinda okuphindaphindiweyo, nangona kunjalo, ngenxa yophuculo olukhulu ekusebenzeni kwekhompyutha, oku akusekho ngxaki. Ngokuhlalutya amaxabiso e-coordinate ee-vertices ezenza imodeli ye-mesh kusetyenziswa iindlela zohlalutyo lwe-multivariate ezifana nohlalutyo lwe-principal component (PCA), kunokwenzeka ukuhlalutya utshintsho kwimo yomphezulu wonke kunye nemo ebonakalayo kuyo nayiphi na indawo ekuhanjisweni. Bala kwaye ubonise53. Kule mihla, iimodeli ze-mesh ezenziwe ngokufaka itemplate zisetyenziswa kakhulu kuhlalutyo lwemilo kwiindawo ezahlukeneyo52,54,55,56,57,58,59,60.
Inkqubela phambili kubuchwepheshe bokurekhoda i-mesh eguquguqukayo, kunye nophuhliso olukhawulezileyo lwezixhobo zokuskena ze-3D eziphathwayo ezikwaziyo ukuskena ngesisombululo esiphezulu, isantya, kunye nokuhambahamba kune-CT, zenza kube lula ukurekhoda idatha yomphezulu we-3D nokuba ikweyiphi na indawo. Ke ngoko, kwicandelo le-anthropology yezinto eziphilayo, ezo teknoloji zintsha ziphucula amandla okulinganisa nokuhlalutya ngokwezibalo iisampulu zabantu, kuquka iisampulu ze-skull, nto leyo eyinjongo yolu phononongo.
Ngamafutshane, olu phononongo lusebenzisa iteknoloji yokumisela i-3D homology ephucukileyo esekwe kuthelekiso lweetemplate (Umfanekiso 1) ukuvavanya iisampuli ze-skull ezingama-342 ezikhethwe kubantu abayi-148 kwihlabathi liphela ngokuthelekiswa kwejografi. Ukwahluka kwemo ye-cranial (Itheyibhile 1). Ukuze siqwalasele utshintsho kwimo ye-skull, sisebenzise uhlalutyo lwe-PCA kunye ne-receiver operating characteristic (ROC) kwiseti yedatha yemodeli ye-homology esiyivelisileyo. Iziphumo ziya kuba negalelo ekuqondeni ngcono utshintsho lwehlabathi kwimo ye-cranial, kubandakanya iipateni zengingqi kunye nokulandelelana kotshintsho olunciphileyo, utshintsho oluhambelanayo phakathi kwamacandelo e-cranial, kunye nokuba khona kweendlela ze-allometric. Nangona olu phononongo lungajongani nedatha kwiinguqu zangaphandle ezimelwe yimozulu okanye iimeko zokutya ezinokuchaphazela imo ye-cranial, iipateni zejografi zemo ye-cranial ezibhalwe kwisifundo sethu ziya kunceda ukuphonononga izinto ezingqongileyo, ze-biomechanical, kunye neze-genetic ze-cranial variation.
Itheyibhile 2 ibonisa i-eigenvalues kunye ne-PCA contribution coefficients ezisetyenzisiweyo kwi-dataset engamiselwanga ye-17,709 vertices (53,127 XYZ coordinates) yeemodeli ze-skull ezingama-342 ezifanayo. Ngenxa yoko, kwachongwa izinto eziphambili ezili-14, igalelo lazo kwi-variance iyonke lalingaphezulu kwe-1%, kwaye isabelo sisonke se-variance yayiyi-83.68%. Iivektha zokulayisha zezinto eziphambili ezili-14 zirekhodwe kwi-Supplementary Table S1, kwaye amanqaku ezinto abalwe kwiisampuli ze-skull ezingama-342 aboniswe kwi-Supplementary Table S2.
Olu phononongo luvavanye izinto eziphambili ezilithoba ezinegalelo elikhulu kune-2%, ezinye zazo zibonisa umahluko omkhulu nobalulekileyo wejografi kwimo ye-cranial. Umfanekiso 2 ii-plot curves ezenziwe ngohlalutyo lwe-ROC ukubonisa izinto ezisebenzayo ze-PCA zokuchonga okanye ukwahlula indibaniselwano nganye yeesampulu kwiiyunithi ezinkulu zejografi (umz., phakathi kwamazwe ase-Afrika nangaphandle kwawo ase-Afrika). Indibaniselwano yasePolynesia ayizange ivavanywe ngenxa yobukhulu obuncinci besampulu esetyenzisiweyo kolu vavanyo. Idatha malunga nokubaluleka komahluko kwi-AUC kunye nezinye izibalo ezisisiseko ezibalwe kusetyenziswa uhlalutyo lwe-ROC iboniswe kwiTheyibhile eyoNgezelelweyo S3.
Ii-ROC curves zisetyenzisiwe kuqikelelo lwe-principal component ezisithoba ezisekelwe kwi-dataset ye-vertex equlathe iimodeli ze-homologous skull ezingama-342. I-AUC: Indawo engaphantsi kwe-curve kukubaluleka kwe-0.01% esetyenziselwa ukwahlula indibaniselwano nganye yejografi kwezinye iindibaniselwano zizonke. I-TPF iyinyani (ucalucalulo olusebenzayo), i-FPF iyinyani (ucalucalulo olungasebenziyo).
Ukuchazwa kwe-ROC curve kushwankathelwe apha ngezantsi, kugxilwe kuphela kwizinto ezinokwahlula amaqela okuthelekisa ngokuba ne-AUC enkulu okanye enkulu kunye nenqanaba eliphezulu lokubaluleka elinokubakho ngaphantsi kwe-0.001. I-complex yaseMzantsi Asia (Umzobo 2a), equlathe ikakhulu iisampuli ezivela eIndiya, yahluke kakhulu kwezinye iisampuli ezixutyiweyo ngokwejografi kuba icandelo lokuqala (PC1) line-AUC enkulu kakhulu (0.856) xa kuthelekiswa nezinye izinto. Uphawu lwe-complex yaseAfrika (Umzobo 2b) yi-AUC enkulu kakhulu ye-PC2 (0.834). Ama-Austro-Melanesia (Umzobo 2c) abonise umkhwa ofanayo nama-Afrika aseMzantsi Sahara nge-PC2 ene-AUC enkulu ngakumbi (0.759). AmaYurophu (Umzobo 2d) ahluke ngokucacileyo kwindibaniselwano ye-PC2 (AUC = 0.801), i-PC4 (AUC = 0.719) kunye ne-PC6 (AUC = 0.671), isampuli yaseMantla-mpuma ye-Asia (Umzobo 2e) yahluke kakhulu kwi-PC4, ene-0.714 enkulu, kwaye umahluko kwi-PC3 ubuthathaka (AUC = 0.688). La maqela alandelayo achongiweyo kunye namaxabiso aphantsi e-AUC kunye namanqanaba aphezulu okubaluleka: Iziphumo ze-PC7 (AUC = 0.679), i-PC4 (AUC = 0.654) kunye ne-PC1 (AUC = 0.649) zibonise ukuba amaMerika aseMelika (Umzobo 2f) aneempawu ezithile ezinxulumene nezi zinto, ama-Southeast Asians (Umzobo 2g) ahluke kwi-PC3 (AUC = 0.660) kunye ne-PC9 (AUC = 0.663), kodwa ipateni yeesampulu ezivela kuMbindi Mpuma (Umzobo 2h) (kuquka noMntla Afrika) iyafana. Xa kuthelekiswa nezinye akukho mahluko mkhulu.
Kwinyathelo elilandelayo, ukuze kutolikwe ngokubonakalayo ii-vertices ezihambelanayo kakhulu, iindawo zomphezulu ezinexabiso eliphezulu lomthwalo elingaphezulu kwe-0.45 zifakwe imibala ngolwazi lwe-X, Y, kunye ne-Z coordinate, njengoko kubonisiwe kuMfanekiso 3. Indawo ebomvu ibonisa ulwalamano oluphezulu nee-X-axis coordinate, ezihambelana nolwalathiso oluthe tye olunqamlezileyo. Ummandla oluhlaza unxulumene kakhulu ne-vertical coordinate ye-Y axis, kwaye ummandla oluhlaza okwesibhakabhaka omnyama unxulumene kakhulu ne-sagittal coordinate ye-Z axis. Ummandla oluhlaza okwesibhakabhaka okhanyayo unxulumene nee-axes ze-Y coordinate kunye nee-axes ze-Z coordinate; indawo epinki - exutyiweyo enxulumene nee-axes ze-X kunye ne-Z coordinate; indawo etyheli - enxulumene nee-axes ze-X kunye ne-Y coordinate; Ummandla omhlophe uqulathe i-X, Y kunye ne-Z coordinate axis ebonakalisiweyo. Ke ngoko, kulo mgangatho wexabiso lomthwalo, i-PC 1 inxulunyaniswa kakhulu nomphezulu wonke we-skull. Ubume be-3 SD virtual skull kwicala elichaseneyo lale axis yecandelo nayo iboniswe kulo mfanekiso, kwaye imifanekiso egobileyo iboniswe kwiVidiyo eyoNgezelelweyo S1 ukuqinisekisa ngokubonakalayo ukuba i-PC1 iqulethe izinto ezinobukhulu be-skull iyonke.
Ukusasazwa rhoqo kwamanqaku e-PC1 (ijika eliqhelekileyo lokulingana), imephu yombala womphezulu wekhakhayi inxulumene kakhulu nee-vertices ze-PC1 (ingcaciso yemibala enxulumene nobukhulu bamacala achaseneyo ale axis yi-3 SD. Isikali sisphere esiluhlaza esinobubanzi obuyi-50 mm.
Umfanekiso 3 ubonisa iploti yokusasazwa rhoqo (i-normal fit curve) yamanqaku e-PC1 ngabanye abalwe ngokwahlukeneyo kwiiyunithi ezili-9 zejografi. Ukongeza kuqikelelo lwe-ROC curve (Umfanekiso 2), uqikelelo lwabantu baseMzantsi Asia luthe tye kakhulu ngasekhohlo kuba iintloko zabo zincinci kunezo zamanye amaqela engingqi. Njengoko kubonisiwe kwiTheyibhile 1, aba bantu baseMzantsi Asia bamele amaqela eentlanga eIndiya kuquka iziqithi zaseAndaman naseNicobar, eSri Lanka naseBangladesh.
I-dimensional coefficient ifunyenwe kwi-PC1. Ukufunyanwa kweendawo ezihambelanayo kakhulu kunye neemilo ezibonakalayo kubangele ukucaciswa kwezinto zefom kwiinxalenye ezingezizo i-PC1; nangona kunjalo, izinto zobukhulu azisuswa ngokupheleleyo rhoqo. Njengoko kubonisiwe ngokuthelekisa ii-ROC curves (Umfanekiso 2), i-PC2 kunye ne-PC4 zezona zicalucalulayo kakhulu, zilandelwa yi-PC6 kunye ne-PC7. I-PC3 kunye ne-PC9 zisebenza kakhulu ekwahluleni inani labantu abasampulu kwiiyunithi zejografi. Ke ngoko, ezi zimbini zee-component axes zibonisa ngokwesikem ii-scatterplots zamanqaku e-PC kunye nombala ohambelana kakhulu nenxalenye nganye, kunye nokuguquguquka kwesimo esibonakalayo kunye nobukhulu bamacala achaseneyo e-3 SD (IiFigs. 4, 5, 6). Ukugubungela kwe-convex hull yeesampuli ezivela kwiyunithi nganye yejografi emelwe kwezi plots malunga ne-90%, nangona kukho inqanaba elithile lokudityaniswa ngaphakathi kwamaqela. Itheyibhile 3 inika inkcazo yenxalenye nganye ye-PCA.
Iigrafu ze-PC2 kunye ne-PC4 zamanqaku e-cranial kubantu abakwiiyunithi ezilithoba zejografi (phezulu) kunye neeyunithi ezine zejografi (ezantsi), iigrafu zombala womphezulu we-skull wee-vertices zihambelana kakhulu ne-PC nganye (ngokuthelekiswa no-X, u-Y, u-Z). Inkcazo yombala wee-axes: jonga umbhalo), kwaye ukuguquguquka kwesimo esibonakalayo kumacala ahlukeneyo ezi axes yi-3 SD. Isikali sisphere eluhlaza enobubanzi obuyi-50 mm.
Iigrafu ze-PC6 kunye ne-PC7 zamanqaku e-cranial kubantu abakwiiyunithi ezilithoba zejografi (phezulu) kunye neeyunithi ezimbini zejografi (ezantsi), iigrafu zemibala yomphezulu we-cranial zee-vertices zihambelana kakhulu ne-PC nganye (ngokuthelekiswa ne-X, Y, Z). Inkcazo yombala wee-axes: jonga umbhalo), kwaye ukuguquguquka kwesimo esibonakalayo kumacala ahlukeneyo ezi axes yi-3 SD. Isikali sisphere eluhlaza enobubanzi obuyi-50 mm.
Iigrafu ze-PC3 kunye ne-PC9 zamanqaku e-cranial kubantu abakwiiyunithi ezilithoba zejografi (phezulu) kunye neeyunithi ezintathu zejografi (ezantsi), kunye neegrafu zemibala yomphezulu wekhakhayi (ngokuthelekiswa nee-axis ze-X, Y, Z) zee-vertices ezihambelana kakhulu nokuchazwa kombala ngamnye we-PC: cm . umbhalo), kunye nokuguquguquka kwemilo ebonakalayo kumacala ahlukeneyo ezi axis ezinobukhulu be-3 SD. Isikali sisphere eluhlaza enobubanzi be-50 mm.
Kwigrafu ebonisa amanqaku e-PC2 kunye ne-PC4 (Umzobo 4, Iividiyo ezongezelelweyo S2, S3 ezibonisa imifanekiso eguqukileyo), imephu yombala womphezulu ikwaboniswa xa umda wexabiso lomthwalo ubekwe ngaphezulu kwe-0.4, ophantsi kune-PC1 kuba ixabiso le-PC2 umthwalo uwonke ungaphantsi kune-PC1.
Ukwandiswa kwee-frontal kunye ne-occipital lobes kwicala le-sagittal ecaleni kwe-Z-axis (uluhlaza okwesibhakabhaka obumnyama) kunye ne-parietal lobe kwicala le-coronal (obomvu) kwi-pink), i-Y-axis ye-occiput (eluhlaza okwesibhakabhaka) kunye ne-Z-axis yebunzi (uluhlaza okwesibhakabhaka obumnyama). Le grafu ibonisa amanqaku abo bonke abantu kwihlabathi liphela; nangona kunjalo, xa zonke iisampuli eziqulathe inani elikhulu lamaqela ziboniswa kunye ngaxeshanye, ukutolikwa kweepateni zokusasazeka kunzima kakhulu ngenxa yobuninzi bokugqithana; ke ngoko, kwiiyunithi ezine ezinkulu zejografi (oko kukuthi, iAfrika, i-Australasia-Melanesia, iYurophu, kunye ne-Northeast Asia), iisampuli zisasazeke ngaphantsi kwegrafu ene-3 SD virtual cranial deformation ngaphakathi kolu luhlu lwamanqaku e-PC. Kumfanekiso, i-PC2 kunye ne-PC4 ziimbini zamanqaku. Ama-Afrika kunye nama-Austro-Melanesia adibanisana ngakumbi kwaye asasazeke kwicala lasekunene, ngelixa amaYurophu esasazeke kwicala eliphezulu lasekhohlo kwaye ama-Northeast Asia adla ngokuqokelelana kwicala elisezantsi lasekhohlo. I-axis ethe tye ye-PC2 ibonisa ukuba ama-Afrika/ama-Australia aseMelanesia ane-neurocranium ende kunabanye abantu. I-PC4, apho iindibaniselwano zaseYurophu nakumntla-mpuma we-Asia zahlulwe ngokukhululekileyo, inxulunyaniswa nobukhulu kunye nokubonakaliswa kwamathambo e-zygomatic kunye ne-lateral contour ye-calvarium. Isicwangciso sokulinganisa sibonisa ukuba amaYurophu anamathambo amancinci kakhulu e-maxillary kunye ne-zygomatic, indawo encinci ye-temporal fossa ethintelwe yi-zygomatic arch, ithambo elingaphambili eliphakanyisiweyo ngokuthe nkqo kunye nethambo elisezantsi le-occipital, ngelixa ama-Northeast Asian edla ngokuba namathambo e-zygomatic abanzi nabonakalayo. I-frontal lobe ithambekile, isiseko sethambo le-occipital siphakanyisiwe.
Xa kugxilwe kwi-PC6 kunye ne-PC7 (Umzobo 5) (Iividiyo ezongezelelweyo S4, S5 zibonisa imifanekiso eguqukileyo), umbala ubonisa umda wexabiso lomthwalo ongaphezulu kwe-0.3, nto leyo ebonisa ukuba i-PC6 inxulunyaniswa nokwakheka kwe-maxillary okanye i-alveolar (ubomvu: i-X axis kunye nohlaza). I-Y axis), imo yethambo lexesha (uluhlaza: ii-axes ze-Y kunye ne-Z) kunye nemo yethambo le-occipital (pink: ii-axes ze-X kunye ne-Z). Ukongeza kububanzi bebunzi (ubomvu: i-X-axis), i-PC7 ikwahambelana nokuphakama kwe-alveoli ye-anterior maxillary (uluhlaza: i-Y-axis) kunye nemo yentloko ye-Z-axis ejikeleze ummandla we-parietotemporal (uluhlaza okwesibhakabhaka obumnyama). Kwiphaneli ephezulu yoMfanekiso 5, zonke iisampulu zejografi zisasazwa ngokweenqaku ze-PC6 kunye ne-PC7. Ngenxa yokuba i-ROC ibonisa ukuba i-PC6 iqulethe iimpawu ezikhethekileyo eYurophu kwaye i-PC7 imele iimpawu ze-Native American kolu hlalutyo, ezi sampuli zimbini zengingqi zidwetshwe ngokukhetha kule sibini see-axes zecandelo. AmaMerika aseMelika, nangona ebandakanyiwe kakhulu kwisampulu, asasazeke kwikona ephezulu ekhohlo; Ngokwahlukileyo koko, iisampulu ezininzi zaseYurophu zihlala kwikona esezantsi ngasekunene. Isibini se-PC6 kunye ne-PC7 zimele inkqubo encinci ye-alveolar kunye ne-neurocranium ebanzi kakhulu yamaYurophu, ngelixa amaMerika ebonakala ngebunzi elimxinwa, i-maxilla enkulu, kunye nenkqubo ebanzi nende ye-alveolar.
Uhlalutyo lwe-ROC lubonise ukuba i-PC3 kunye/okanye i-PC9 zazixhaphakile kubemi baseMzantsi-mpuma nakumantla-mpuma aseAsia. Ngoko ke, amanqaku adibanisa i-PC3 (ubuso obuluhlaza obuphezulu kwi-y-axis) kunye ne-PC9 (ubuso obuluhlaza obuphantsi kwi-y-axis) (Umzobo 6; Iividiyo ezongezelelweyo S6, S7 zibonelela ngemifanekiso eguquliweyo) zibonisa ukwahluka kwama-Asia aseMpuma. , nto leyo eyahluke kakhulu kumlinganiselo ophezulu wobuso bama-Asia aseMpuma-mpuma kunye nesimo sobuso esiphantsi sama-Asia aseMpuma-mpuma. Ngaphandle kwezi mpawu zobuso, olunye uphawu lwabanye abantu baseMpuma-mpuma aseAsia kukuthambeka kwethambo le-occipital, ngelixa abanye abantu baseMpuma-mpuma aseAsia benesiseko esincinci sekhanda.
Inkcazo engasentla yezinto eziphambili kunye nenkcazo ye-PC5 kunye ne-PC8 ayifakwanga kuba akukho mpawu zithile zengingqi ezifunyenweyo phakathi kweeyunithi eziphambili zejografi ezilithoba. I-PC5 ibhekisa kubungakanani benkqubo ye-mastoid yethambo lexesha, kwaye i-PC8 ibonisa ukungalingani kobume be-skull iyonke, zombini zibonisa umahluko ohambelanayo phakathi kweendibaniselwano zeesampuli zejografi ezilithoba.
Ukongeza kwii-scatterplots zamanqaku e-PCA akwinqanaba lomntu ngamnye, sikwabonelela ngee-scatterplots zeendlela zeqela ukuze kuthelekiswe ngokubanzi. Ngenxa yoku, imodeli ye-avareji ye-cranial homology yenziwe kwiseti yedatha ye-vertex yeemodeli ze-homology zomntu ngamnye ezivela kumaqela eentlanga ali-148. Ii-bivariate plots zeeseti zamanqaku ze-PC2 kunye ne-PC4, i-PC6 kunye ne-PC7, kunye ne-PC3 kunye ne-PC9 ziboniswe kwi-Supplementary Figure S1, zonke zibalwe njengemodeli ye-avareji ye-skull yesampulu yabantu abali-148. Ngale ndlela, ii-scatterplots zifihla umahluko womntu ngamnye kwiqela ngalinye, zivumela ukutolikwa okucacileyo kokufana kwe-skull ngenxa yokusasazwa kwengingqi okusisiseko, apho iipateni zihambelana nezo ziboniswe kwiiploti zomntu ngamnye ezinokugqithisana okuncinci. I-Supplementary Figure S2 ibonisa imodeli ye-avareji iyonke yeyunithi nganye yejografi.
Ukongeza kwi-PC1, eyayinxulunyaniswa nobukhulu obupheleleyo (Itheyibhile Eyongezelelekileyo S2), ubudlelwane be-allometric phakathi kobukhulu obupheleleyo kunye nemilo ye-skull buhlolwe kusetyenziswa ubukhulu be-centroid kunye neeseti zoqikelelo lwe-PCA ezivela kwidatha engaqhelekanga. Ii-coefficients ze-allometric, amaxabiso angaguqukiyo, amaxabiso e-t, kunye namaxabiso e-P kuvavanyo lokubaluleka ziboniswe kwiTheyibhile 4. Akukho zixhobo zepatheni ye-allometric ezibalulekileyo ezinxulumene nobukhulu be-skull zifunyenwe kuyo nayiphi na i-morphology ye-cranial kwinqanaba le-P < 0.05.
Ngenxa yokuba ezinye izinto zobukhulu zinokufakwa kuqikelelo lwe-PC olusekelwe kwiiseti zedatha ezingacwangciswanga, sihlolisise ngakumbi i-allometric trend phakathi kobukhulu be-centroid kunye namanqaku e-PC abalwe kusetyenziswa iiseti zedatha ezicwangciswe ngokobukhulu be-centroid (iziphumo ze-PCA kunye neeseti zamanqaku ziboniswe kwiiTheyibhile ezongezelelweyo S6). , C7). Itheyibhile 4 ibonisa iziphumo zohlalutyo lwe-allometric. Ke ngoko, ii-allometric trends ezibalulekileyo zifunyenwe kwinqanaba le-1% kwi-PC6 nakwinqanaba le-5% kwi-PC10. Umfanekiso 7 ubonisa imiqolo yokubuyela umva kwezi budlelwane ze-log-linear phakathi kwamanqaku e-PC kunye nobukhulu be-centroid kunye nee-dummies (±3 SD) nakweyiphi na isiphelo sobukhulu be-log centroid. I-PC6 score yi-ratio yokuphakama kunye nobubanzi be-skull. Njengoko ubukhulu be-skull bukhula, i-skull kunye nobuso ziya ziphakama, kwaye ibunzi, amasokethi eeso kunye neempumlo zidla ngokusondelana ecaleni. Ipateni yokusasazwa kwesampulu ibonisa ukuba eli nani lihlala lifumaneka kuma-Asia aseMpuma-mpuma kunye namaMerika aseMelika. Ngaphezu koko, i-PC10 ibonisa i-trend eya ekunciphiseni ngokulinganayo ububanzi be-midface kungakhathaliseki ummandla wejografi.
Kwimiba ebalulekileyo ye-allometric edweliswe kwitheyibhile, i-slope ye-log-linear regression phakathi kwe-PC proportion ye-shape component (efunyenwe kwi-normalized data) kunye nobukhulu be-centroid, i-virtual shape deformation inobukhulu be-3 SD kwelinye icala lomgca we-4.
Olu hlobo lulandelayo lotshintsho kwimo ye-cranial luye lwabonakaliswa ngohlalutyo lwedatha yeemodeli zomphezulu we-3D ezifanayo. Icandelo lokuqala le-PCA linxulumene nobukhulu be-skull iyonke. Kudala kucingwa ukuba ii-skull ezincinci zaseMzantsi Asia, kuquka iisampulu ezivela eIndiya, eSri Lanka nakwiZiqithi zaseAndaman, eBangladesh, zibangelwa bubukhulu bazo bomzimba obuncinci, obuhambelana nomthetho we-ecogeographic kaBergmann okanye umthetho wesiqithi613,5,16,25,27,62. Eyokuqala inxulumene nobushushu, kwaye eyesibini ixhomekeke kwindawo ekhoyo kunye nezixhobo zokutya ze-niche yendalo. Phakathi kwezinto zemilo, utshintsho olukhulu lumlinganiselo wobude nobubanzi be-cranial vault. Olu phawu, olubizwa ngokuba yi-PC2, luchaza ubudlelwane obusondeleyo phakathi kwee-skull ezinde ngokulinganayo zama-Austro-Melanesia nama-Afrika, kunye nomahluko kwii-skull ezingqukuva zabanye abantu baseYurophu nabase-Asia basemantla-mpuma. Ezi mpawu zixeliwe kwizifundo ezininzi zangaphambili ngokusekelwe kumlinganiselo olula we-linear37,63,64. Ngaphezu koko, olu phawu lunxulunyaniswa ne-brachycephaly kubantu abangengabo amaAfrika, nto leyo ekudala ixutyushwa kwizifundo ze-anthropometric kunye ne-osteometric. Ingcamango ephambili emva kwale ngcaciso kukuba ukuncipha kokuhlafuna, njengokuncitshiswa kwemisipha ye-temporalis, kunciphisa uxinzelelo kwintloko yangaphandle5,8,9,10,11,12,13. Enye ingcamango ibandakanya ukuziqhelanisa nemozulu ebandayo ngokunciphisa indawo yomphezulu wentloko, nto leyo ebonisa ukuba ukhakhayi oluyi-spherical lunciphisa indawo yomphezulu ngcono kunemilo eyi-spherical, ngokwemithetho ka-Allen16,17,25. Ngokusekelwe kwiziphumo zophando lwangoku, ezi ngcamango zinokuvavanywa kuphela ngokusekelwe kulwalamano olunqamlezileyo lweziqendu ze-cranial. Ngamafutshane, iziphumo zethu ze-PCA azixhasi ngokupheleleyo ingcamango yokuba umlinganiselo wobude nobubanzi be-cranial uchatshazelwa kakhulu ziimeko zokuhlafuna, njengoko umthwalo we-PC2 (icandelo elide/le-brachycephalic) wawunganxulumananga kakhulu nobukhulu bobuso (kubandakanya ubukhulu be-maxillary). kunye nendawo enxulumeneyo ye-temporal fossa (ebonakalisa ubungakanani bomsipha we-temporalis). Uphononongo lwethu lwangoku aluzange luhlalutye ubudlelwane phakathi kobume be-spherical kunye neemeko zejoloji ezifana nobushushu; nangona kunjalo, ingcaciso esekelwe kumthetho ka-Allen ingafaneleka ukuqwalaselwa njengengcamango efunekayo yokuchaza i-brachycephalon kwimimandla yemozulu ebandayo.
Umahluko omkhulu emva koko wafunyanwa kwi-PC4, nto leyo ebonisa ukuba ama-Asia aseMpuma-mpuma anamathambo amakhulu, abonakalayo e-zygomatic kumathambo e-maxilla kunye ne-zygomatic. Olu phando luhambelana nophawu oluthile olwaziwayo lwamaSiberia, ekucingelwa ukuba aqhelane nemozulu ebanda kakhulu ngokuhamba phambili kwamathambo e-zygomatic, nto leyo ebangela ukwanda komthamo wee-sinuses kunye nobuso obuthe tyaba 65. Uphando olutsha oluvela kwimodeli yethu efana nalo kukuba ukugoba kwezidlele kumaYurophu kunxulunyaniswa nokuthambeka okungaphantsi, kunye namathambo e-occipital athambileyo nancinci kunye ne-nuchal concavity. Ngokwahlukileyo koko, ama-Asia aseMpuma-mpuma adla ngokuba namabunzi athambileyo kunye nemimandla ephakamileyo ye-occipital. Izifundo zethambo le-occipital zisebenzisa iindlela ze-geometric morphometric35 zibonise ukuba ii-skulls zase-Asia nezaseYurophu zine-flatter nuchal curve kunye nendawo ephantsi ye-occiput xa kuthelekiswa nama-Afrika. Nangona kunjalo, ii-scatterplots zethu ze-PC2 kunye ne-PC4 kunye ne-PC3 kunye ne-PC9 pairs zibonise umahluko omkhulu kuma-Asia, ngelixa amaYurophu ayephawulwa ngesiseko esithe tyaba se-occiput kunye ne-occiput esezantsi. Ukungangqinelani kwiimpawu zase-Asia phakathi kwezifundo kusenokuba ngenxa yokwahluka kwiisampulu zohlanga ezisetyenzisiweyo, njengoko sithathe iisampulu kumaqela amaninzi eentlanga ezivela kuluhlu olubanzi lwaseMantla-mpuma naseMazantsi-mpuma e-Asia. Utshintsho kwimo yethambo le-occipital ludla ngokunxulunyaniswa nophuhliso lwemisipha. Nangona kunjalo, le nkcazo eguquguqukayo ayibalulekanga ulwalamano phakathi kwebunzi kunye nesimo se-occiput, esibonakaliswe kolu phononongo kodwa akunakwenzeka ukuba sibonakaliswe ngokupheleleyo. Kule nkalo, kufanelekile ukuqwalasela ubudlelwane phakathi kokulinganisela ubunzima bomzimba kunye neziko lomxhuzulane okanye i-cervical junction (foramen magnum) okanye ezinye izinto.
Elinye icandelo elibalulekileyo elinomahluko omkhulu linxulumene nophuhliso lwezixhobo zokuhlafuna, ezimelwe yi-maxillary kunye ne-temporal fossae, echazwa yindibaniselwano yamanqaku e-PC6, PC7 kunye ne-PC4. Oku kuncipha okuphawulekayo kwiinxalenye ze-cranial kubonisa abantu baseYurophu ngaphezu kwalo naliphi na elinye iqela leendawo. Olu phawu lutolikwe njengesiphumo sokuncipha kokuzinza kobume bobuso ngenxa yophuhliso lwasekuqaleni lweendlela zolimo nokulungiselela ukutya, nto leyo eyanciphisa umthwalo woomatshini kwizixhobo zokuhlafuna ngaphandle kwesixhobo esinamandla sokuhlafuna9,12,28,66. Ngokwe-hypothesis yomsebenzi wokuhlafuna, 28 oku kuhamba notshintsho ekujikeni kwesiseko se-skull ukuya kwi-angle ye-cranial ebukhali ngakumbi kunye nophahla lwe-cranial olujikelezayo ngakumbi. Ukusuka kule mbono, abemi bezolimo badla ngokuba nobuso obuncinci, ukuvela okuncinci kwe-mandible, kunye ne-globular meninges. Ke ngoko, olu tshintsho lunokuchazwa ngumda ngokubanzi wesimo se-lateral se-skull yabantu baseYurophu abanezitho zokuhlafuna ezinciphileyo. Nangona kunjalo, ngokutsho kolu phononongo, olu hlalutyo luyinkimbinkimbi kuba ukubaluleka kokusebenza kobudlelwane bemo phakathi kwe-globose neurocranium kunye nophuhliso lwesixhobo sokuhlafuna akwamkelekanga kangako, njengoko bekucingelwa kwiingcaciso zangaphambili ze-PC2.
Umahluko phakathi kwama-Asia aseMpuma-mpuma kunye nama-Asia aseMzantsi-mpuma uboniswa ngumahluko phakathi kobuso obude obunethambo le-occipital elithambekileyo kunye nobuso obufutshane obunesiseko esincinci sekhanda, njengoko kubonisiwe kwi-PC3 kunye ne-PC9. Ngenxa yokunqongophala kwedatha ye-geoecological, uphando lwethu lubonelela ngengcaciso encinci kuphela kolu fumaniso. Ingcaciso enokwenzeka kukuziqhelanisa nemozulu eyahlukileyo okanye iimeko zesondlo. Ukongeza ekuziqhelaniseni nendalo, umahluko wendawo kwimbali yabemi baseMpuma-mpuma naseMzantsi-mpuma Asia nawo wathathelwa ingqalelo. Umzekelo, empuma ye-Eurasia, imodeli enamaleko amabini iye yacingelwa ukuba iyaqonda ukusasazeka kwabantu banamhlanje (AMH) ngokusekelwe kwidatha ye-cranial morphometric67,68. Ngokwale modeli, "inqanaba lokuqala", oko kukuthi, amaqela okuqala e-Late Pleistocene AMH colonizers, ayenemvelaphi ethe ngqo evela kubemi bomthonyama balo mmandla, njengabase-Austro-Melanesia banamhlanje (p. First stratum). , kwaye kamva wadibana nomxube omkhulu wabantu bezolimo basemantla abaneempawu zasempuma-mpuma Asia (umaleko wesibini) kulo mmandla (malunga neminyaka engama-4,000 eyadlulayo). Ukuhamba kwezakhi zofuzo okumaphu kusetyenziswa imodeli "yeengqimba ezimbini" kuya kufuneka ukuze kuqondwe imo yengqimba yaseMzantsi-mpuma Asia, njengoko imo yengqimba yaseMzantsi-mpuma Asia inokuxhomekeka ngokuyinxenye kwilifa lezakhi zofuzo lendawo yokuqala.
Ngokuvavanya ukufana kwe-cranial sisebenzisa iiyunithi zejografi ezidwetshwe kusetyenziswa iimodeli ezifanayo, singaqikelela imbali yoluntu esisiseko ye-AMF kwiimeko ezingaphandle kwe-Afrika. Iimodeli ezininzi ezahlukeneyo "eziphuma e-Afrika" ziye zacetyiswa ukuchaza ukusasazwa kwe-AMF ngokusekelwe kwidatha yamathambo kunye ne-genomic. Kwezi, izifundo zakutshanje zibonisa ukuba i-AMH colonization yeendawo ezingaphandle kwe-Afrika yaqala malunga neminyaka eyi-177,000 eyadlulayo69,70. Nangona kunjalo, ukusasazwa kwe-AMF kwi-Eurasia ngeli xesha akukacaci, kuba iindawo zokuhlala zezi fossils zokuqala zilinganiselwe kuMbindi Mpuma nakwiMeditera kufutshane ne-Afrika. Ityala elilula yindawo yokuhlala enye ecaleni kwendlela yokufuduka esuka e-Afrika ukuya e-Eurasia, idlula imiqobo yejografi efana neHimalayas. Enye imodeli ibonisa amaza amaninzi okufuduka, eyokuqala eyasasazeka ukusuka e-Afrika ecaleni konxweme loLwandlekazi lwaseIndiya ukuya kuMzantsi-mpuma we-Asia nase-Australia, yaze yasasazeka kumantla e-Eurasia. Uninzi lwezi zifundo ziqinisekisa ukuba i-AMF yasasazeka ngaphaya kwe-Afrika malunga neminyaka engama-60,000 eyadlulayo. Kule meko, iisampulu zase-Australasian-Melanesian (kuquka nePapua) zibonisa ukufana okukhulu neesampulu zase-Afrika kunanoma yiluphi olunye uthotho lwejografi kuhlalutyo lwezinto eziphambili zeemodeli ze-homology. Olu fumaniso luxhasa ingcamango yokuba amaqela okuqala okusasazwa kwe-AMF kumda osemazantsi we-Eurasia avela ngqo e-Afrika22,68 ngaphandle kotshintsho olukhulu lwemo ephendula kwimozulu ethile okanye kwezinye iimeko ezibalulekileyo.
Ngokuphathelele ukukhula kwe-allometric, uhlalutyo olusebenzisa izinto ezikwimo ezithathwe kwiseti yedatha eyahlukileyo ehlengahlengiswe ngobukhulu be-centroid lubonise umkhwa obalulekileyo we-allometric kwi-PC6 kunye ne-PC10. Zombini ezi zinto zinxulumene nemilo yebunzi kunye neendawo zobuso, eziba ncinci njengoko ubukhulu bekhaka bukhula. Ama-Asia aseMpuma-mpuma namaMerika adla ngokuba nale mpawu kwaye aneekhaka ezinkulu. Olu phando luphikisana neepateni ze-allometric ezixeliweyo ngaphambili apho iingqondo ezinkulu zinee-frontal lobes ezibanzi kwindawo ebizwa ngokuba yi-"Broca's cap", nto leyo ebangela ukwanda kobubanzi be-frontal lobe34. Olu mahluko luchazwa ngokwahluka kwiiseti zeesampuli; Uphononongo lwethu luhlalutye iipateni ze-allometric zobukhulu be-cranial iyonke kusetyenziswa abantu banamhlanje, kwaye izifundo zokuthelekisa zijongana neendlela zexesha elide zokuguquka kwabantu ezinxulumene nobukhulu bobuchopho.
Ngokuphathelele i-allometric yobuso, olunye uphando olusebenzisa i-biometric data78 lufumanise ukuba imo yobuso kunye nobukhulu bunokuhambelana kancinci, ngelixa uphando lwethu lufumanise ukuba ii-skulls ezinkulu zihlala zinxulunyaniswa nobuso obude nobuncinci. Nangona kunjalo, ukuhambelana kwedatha ye-biometric akucaci; Uvavanyo lwe-Regression oluthelekisa i-ontogenetic allometry kunye ne-static allometry lubonisa iziphumo ezahlukeneyo. Ukuthambekela kwe-allometric kwimo ye-skull engqukuva ngenxa yokuphakama okuphezulu nako kuye kwaxelwa; nangona kunjalo, asizange sihlalutye idatha yokuphakama. Uphononongo lwethu lubonisa ukuba akukho datha ye-allometric ebonisa ulwalamano phakathi kwe-cranial globular proportions kunye nobukhulu be-cranial iyonke.
Nangona uphando lwethu lwangoku lungajongani nedatha kwiinguqu zangaphandle ezimelwe yimozulu okanye iimeko zokutya ezinokuchaphazela imo yentloko, iseti enkulu yedatha yeemodeli zomphezulu wentloko ze-3D ezisetyenziswa kolu phononongo ziya kunceda ukuvavanya umahluko wemo yentloko ohambelanayo. Izinto ezingqongileyo ezifana nokutya, imeko yemozulu kunye nesondlo, kunye namandla angathathi cala afana nokufuduka, ukuhamba kwezakhi zofuzo kunye nokuhamba kwezakhi zofuzo.
Olu phononongo luquke iisampulu ezingama-342 zeentloko zeengxilimbela zamadoda eziqokelelwe kwiindawo ezili-148 kwiiyunithi ezili-9 zejografi (Itheyibhile 1). Uninzi lwamaqela ziisampuli zejografi, ngelixa amanye amaqela aseAfrika, eMntla-mpuma/Mzantsi-mpuma weAsia naseMelika (adweliswe ngoonobumba abakekileyo) achazwe ngokweenhlanga. Iisampulu ezininzi zentloko zeengxilimbela zikhethwe kwisiseko sedatha yokulinganisa intloko ngokwenkcazo yokulinganisa intloko kaMartin ebonelelwe nguTsunehiko Hanihara. Sikhethe iintloko zeengxilimbela zamadoda ezimele kuzo zonke iintlanga ehlabathini. Ukuze sichonge amalungu eqela ngalinye, sibale imigama ye-Euclidean ngokusekelwe kwimilinganiselo yentloko engama-37 kwiqela eliphakathi kwabo bonke abantu abakwelo qela. Kwiimeko ezininzi, sikhethe iisampulu ezili-1–4 ezinomgama omncinci ukusuka kwi-avareji (Itheyibhile eyongezelelweyo S4). Kula maqela, ezinye iisampulu zikhethwe ngokungacwangciswanga ukuba azidweliswanga kwisiseko sedatha yokulinganisa iHahara.
Ukuze kuthelekiswe izibalo, iisampulu zabemi ezili-148 zahlulwe zaya kwiiyunithi ezinkulu zejografi, njengoko kubonisiwe kwiTheyibhile 1. Iqela "lama-Afrika" liqulathe kuphela iisampulu ezivela kummandla ongaphantsi kweSahara. Iisampulu ezivela eMntla Afrika zaqukwa kwi "Mbindi Mpuma" kunye neisampulu ezivela eNtshona Asia ezineemeko ezifanayo. Iqela laseMntla-mpuma Asia liquka kuphela abantu abangengobaseYurophu, kwaye iqela laseMelika liquka amaMelika aseMelika kuphela. Ngokukodwa, eli qela lisasazwe kwindawo enkulu yelizwekazi laseMntla naseMzantsi Melika, kwiindawo ezahlukeneyo. Nangona kunjalo, siqwalasela isampulu yase-US kule yunithi inye yejografi, ngenxa yembali yabantu baseMelika abathathwa njengabavela eMntla-mpuma Asia, nokuba bafudukele phi na 80.
Sirekhode idatha yomphezulu we-3D yezi sampuli ze-skull ezichaseneyo sisebenzisa iskena se-3D esinesisombululo esiphezulu (i-EinScan Pro yiShining 3D Co Ltd, isisombululo esincinci: 0.5 mm, https://www.shining3d.com/) saza savelisa i-mesh. Imodeli ye-mesh inamacala angama-200,000–400,000, kwaye isoftware ebandakanyiweyo isetyenziselwa ukugcwalisa imingxunya kunye nemiphetho egudileyo.
Kwinyathelo lokuqala, sisebenzise idatha yokuskena kuyo nayiphi na i-skull ukwenza imodeli ye-skull ene-template enye equlathe ii-vertices ezingama-4485 (ubuso be-polygonal obungama-8728). Isiseko sommandla we-skull, esakhiwe yi-sphenoid bone, i-petrous temporal bone, i-palate, i-maxillary alveoli, kunye namazinyo, sisusiwe kwimodeli ye-template mesh. Isizathu kukuba ezi zakhiwo ngamanye amaxesha aziphelelanga okanye kunzima ukuzigqiba ngenxa yeendawo ezibhityileyo okanye ezibhityileyo ezibukhali ezifana neendawo ze-pterygoid kunye neenkqubo ze-styloid, ukuguguleka kwamazinyo kunye/okanye iseti yamazinyo engahambelaniyo. Isiseko se-skull esijikeleze i-foramen magnum, kubandakanya nesiseko, asizange sisuswe kuba le yindawo ebalulekileyo ngokwe-anatomiki yendawo yamalungu omlomo wesibeleko kwaye ukuphakama kwe-skull kufuneka kuhlolwe. Sebenzisa amaringi esipili ukwenza itemplate elinganayo kumacala omabini. Yenza i-isotropic meshing ukuguqula iimilo ze-polygonal zibe zilingana kangangoko kunokwenzeka.
Okulandelayo, iindawo eziphawulweyo ezingama-56 zabelwa kwii-vertices ezihambelana nokwakheka komzimba zemodeli yetemplate kusetyenziswa isoftware ye-HBM-Rugle. Useto lweempawu eziphawulekayo luqinisekisa ukuchaneka kunye nokuzinza kwendawo ephawulweyo kwaye luqinisekisa ukuhambelana kwezi ndawo kwimodeli ye-homology evelisiweyo. Zinokuchongwa ngokusekelwe kwiimpawu zazo ezithile, njengoko kuboniswe kwiTheyibhile eyoNgezelelweyo S5 kunye noMfanekiso oNgezelelweyo S3. Ngokwenkcazo kaBookstein81, uninzi lwezi ndawo ziphawulweyo zii-Type I ezikwindawo edibana kuyo izakhiwo ezintathu, kwaye ezinye zii-Type II markmarks ezinamachaphaza aphezulu. Iindawo ezininzi eziphawulweyo zadluliselwa ukusuka kumanqaku achazwe kwiimilinganiselo ze-cranial ezithe ngqo kwinkcazo kaMartin 36. Sichaze iindawo ezifanayo ezingama-56 zeemodeli eziskeniweyo zeesampuli ze-skull ezingama-342, ezabelwa ngesandla kwii-vertices ezihambelana nokwakheka komzimba ukuvelisa iimodeli ze-homology ezichanekileyo ngakumbi kwicandelo elilandelayo.
Inkqubo yokulungelelanisa esekwe kwintloko ichazwe ukuchaza idatha yokuskena kunye netemplate, njengoko kuboniswe kwiSupplementary Figure S4. I-XZ plane yi-Frankfurt horizontal plane edlula kwindawo ephezulu (inkcazo kaMartin: inxalenye) yomphetho ophezulu wemijelo yangaphandle yasekhohlo nasekunene kunye nendawo ephantsi (inkcazo kaMartin: i-orbit) yomphetho ongezantsi we-orbit yasekhohlo. . I-X axis ngumgca odibanisa amacala asekhohlo nasekunene, kwaye i-X+ licala lasekunene. I-YZ plane idlula embindini weendawo zasekhohlo nasekunene kwaye ingcambu yempumlo: Y+ phezulu, Z+ phambili. Indawo yokubhekisa (imvelaphi: zero coordinate) ibekwe kwindawo yokudibana kwe-YZ plane (midplane), i-XZ plane (Frankfort plane) kunye ne-XY plane (coronal plane).
Sisebenzise isoftware ye-HBM-Rugle (Medic Engineering, Kyoto, http://www.rugle.co.jp/) ukwenza imodeli ye-homologous mesh ngokwenza i-template fitting sisebenzisa amanqaku angama-56 (icala lasekhohlo loMfanekiso 1). Icandelo lesoftware eliphambili, elaphuhliswa okokuqala yiZiko loPhando lweDijithali lwaBantu kwi-Institute of Advanced Industrial Science and Technology eJapan, libizwa ngokuba yi-HBM kwaye linemisebenzi yokufakela iitemplate kusetyenziswa iimpawu zomhlaba kunye nokudala iimodeli ze-fine mesh kusetyenziswa iindawo zokwahlulahlula82. Inguqulelo yesoftware elandelayo (mHBM) 83 yongeze uphawu lokufakela ipateni ngaphandle kweempawu zomhlaba ukuphucula ukusebenza kokufakela. I-HBM-Rugle idibanisa isoftware ye-mHBM kunye neempawu ezongezelelweyo ezisebenziseka lula kubandakanya ukwenza ngokwezifiso iinkqubo zokulungelelanisa kunye nokutshintsha ubungakanani bedatha yokufaka. Ukuthembeka kokuchaneka kokufakela isoftware kuqinisekisiwe kwizifundo ezininzi52,54,55,56,57,58,59,60.
Xa kufakwa itemplate ye-HBM-Rugle kusetyenziswa iimpawu zomhlaba, imodeli yemesh yetemplate ifakwa kwidatha yokuskena ekujoliswe kuyo ngokubhalisa okuqinileyo okusekelwe kwitekhnoloji ye-ICP (ukunciphisa isixa semigama phakathi kweempawu zomhlaba ezihambelana netemplate kunye nedatha yokuskena ekujoliswe kuyo), kwaye ngokuguqulwa okungagungqiyo kwemesh kulungelelanisa itemplate kwidatha yokuskena ekujoliswe kuyo. Le nkqubo yokufakela iphindwe kathathu kusetyenziswa amaxabiso ahlukeneyo eeparameter ezimbini zokufakela ukuphucula ukuchaneka kokufakela. Enye yezi parameter ithintela umgama phakathi kwemodeli yegridi yetemplate kunye nedatha yokuskena ekujoliswe kuyo, kwaye enye ihlawulisa umgama phakathi kweempawu zomhlaba zetemplate kunye neempawu zomhlaba ezijoliswe kuyo. Imodeli yemesh yetemplate eguquliweyo yahlulwahlulwa kusetyenziswa i-algorithm yokuhlulwahlulwa komhlaba ejikelezayo 82 ukwenza imodeli yemesh ecociweyo equlathe ii-vertices ezili-17,709 (ii-polygon ezingama-34,928). Okokugqibela, imodeli yegridi yetemplate ehlukanisiweyo ifanelekile kwidatha yokuskena ekujoliswe kuyo ukuvelisa imodeli ye-homology. Ekubeni iindawo eziphawulweyo zahlukile kancinci kwezo zikwidatha yokuskena ekujoliswe kuyo, imodeli ye-homology yalungiswa kakuhle ukuze ichazwe kusetyenziswa inkqubo yokuhlela intloko echazwe kwicandelo elidlulileyo. Umgama ophakathi phakathi kweempawu ezihambelanayo zemodeli ye-homologous kunye nedatha yokuskena ekujoliswe kuyo kuzo zonke iisampuli yayiyi-<0.01 mm. Kubalwe kusetyenziswa umsebenzi we-HBM-Rugle, umgama ophakathi phakathi kwamanqaku edatha yemodeli ye-homology kunye nedatha yokuskena ekujoliswe kuyo yayiyi-0.322 mm (Itheyibhile eyongezelelweyo S2).
Ukuchaza utshintsho kwimo ye-cranial, ii-vertices ezili-17,709 (ii-53,127 XYZ coordinates) zazo zonke iimodeli ze-homologous zihlalutywe yi-principal component analysis (PCA) kusetyenziswa isoftware ye-HBS eyenziwe yiZiko leSayensi yeDijithali kwi-Institute of Advanced Industrial Science and Technology. , eJapan (umthengisi wosasazo: Medic Engineering, Kyoto, http://www.rugle.co.jp/). Emva koko sazama ukusebenzisa i-PCA kwiseti yedatha engacwangciswanga kwaye iseti yedatha icwangciswe ngobukhulu be-centroid. Ke ngoko, i-PCA esekelwe kwidatha engacwangciswanga inokuyichaza ngokucacileyo imo ye-cranial yeeyunithi ezilithoba zejografi kwaye iququzelele ukutolikwa kwecandelo kune-PCA kusetyenziswa idatha ecwangcisiweyo.
Eli nqaku libonisa inani leenxalenye eziphambili ezifunyenweyo ezinegalelo elingaphezulu kwe-1% yomahluko uwonke. Ukufumanisa ezona nxalenye ziphambili zisebenza kakhulu ekwahlukaniseni amaqela kwiiyunithi ezinkulu zejografi, uhlalutyo lweempawu zokusebenza kwereceiver (ROC) lusetyenziswe kumanqaku eenxalenye eziphambili (PC) ezinegalelo elingaphezulu kwe-2% 84. Olu hlalutyo luvelisa i-probability curve yecandelo ngalinye le-PCA ukuphucula ukusebenza kohlulo kunye nokuthelekisa ngokuchanekileyo iiploti phakathi kwamaqela ejografi. Ubungakanani bamandla ocalucalulo bunokuvavanywa yindawo engaphantsi kwe-curve (AUC), apho iinxalenye ze-PCA ezinamaxabiso amakhulu zikwazi ngcono ukwahlula phakathi kwamaqela. Uvavanyo lwe-chi-square lwenziwa emva koko ukuvavanya inqanaba lokubaluleka. Uhlalutyo lwe-ROC lwenziwa kwiMicrosoft Excel kusetyenziswa isoftware yeBell Curve ye-Excel (inguqulelo 3.21).
Ukuze ubone umahluko wejografi kwimo ye-cranial morphology, ii-scatterplots zenziwe kusetyenziswa amanqaku e-PC awahlula amaqela kwiiyunithi ezinkulu zejografi. Ukuze utolike iinxalenye eziphambili, sebenzisa imephu yemibala ukuze ubone ii-vertices zemodeli ezihambelana kakhulu neenxalenye eziphambili. Ukongeza, ukubonakaliswa okubonakalayo kweziphelo zee-axes ze-principal component ezikwi-±3 standard deviations (SD) zamanqaku e-principal component kubalwe kwaye kwaboniswa kwividiyo eyongezelelweyo.
I-Allometry isetyenzisiwe ukumisela ulwalamano phakathi kwemilo ye-skull kunye nezinto zobungakanani ezivavanyiweyo kuhlalutyo lwe-PCA. Uhlalutyo lusebenza kwiinxalenye eziphambili ezineminikelo >1%. Omnye umda wale PCA kukuba iinxalenye zemilo azinakubonisa imilo nganye nganye kuba iseti yedatha engaqhelekanga ayisusi zonke izinto ezinobukhulu. Ukongeza ekusebenziseni iiseti zedatha ezingaqhelekanga, sikwahlalutye iindlela ze-allometric sisebenzisa iiseti ze-PC fraction ngokusekelwe kwidatha yobukhulu be-centroid eqhelekileyo esetyenziswa kwiinxalenye eziphambili ezineminikelo >1%.
Iindlela ze-allometric zivavanyiwe kusetyenziswa i-equation Y = aXb 85 apho u-Y eyimilo okanye i-proportion ye-shape component, u-X bukhulu be-centroid (Itheyibhile Eyongezelelekileyo S2), u-a lixabiso elingaguqukiyo, kwaye u-b yi-allometric coefficient. Le ndlela ngokusisiseko yazisa izifundo zokukhula kwe-allometric kwi-geometric morphometry78,86. Utshintsho lwe-logarithmic lwale fomula luthi: i-log Y = b × i-log X + i-log a. Uhlalutyo lwe-regression kusetyenziswa indlela ye-least squares lusetyenzisiwe ukubala u-a kunye no-b. Xa u-Y (ubukhulu be-centroid) kunye no-X (amanqaku e-PC) ziguqulwa ngokwe-logarithmic, la maxabiso kufuneka abe mahle; nangona kunjalo, iseti yoqikelelo lwe-X iqulethe amaxabiso angalunganga. Njengesisombululo, songeze i-rounding kwixabiso elipheleleyo le-least fraction kunye no-1 kwi-fraction nganye kwi-component nganye kwaye sasebenzisa i-logarithmic transformation kuzo zonke ii-converted positive fractions. Ukubaluleka kwee-allometric coefficients kuhlolwe kusetyenziswa uvavanyo lwe-t lwabafundi olunemisila emibini. Ezi zibalo zezibalo zokuvavanya ukukhula kwe-allometric zenziwe kusetyenziswa iiBell Curves kwisoftware ye-Excel (inguqulelo 3.21).
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Ixesha lokuthumela: Epreli-02-2024