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Genetické Vazby V Okolí Rudého Moře Hodnocené Pomocí Mtdna“ Zpracovala Samostatně a Že Jsem Uvedla Všechny Použité Infromační Zdroje a Literaturu

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Genetické Vazby V Okolí Rudého Moře Hodnocené Pomocí Mtdna“ Zpracovala Samostatně a Že Jsem Uvedla Všechny Použité Infromační Zdroje a Literaturu Genetické vazby v okolí Rudého moře hodnocené pomocí mtDNA The genetic links around the Red Sea as revealed by the mtDNA DIPLOMOVÁ PRÁCE Bc. Martina Čížková Vedoucí práce: doc. Mgr. Viktor Černý, Dr. KATEDRA ANTROPOLOGIE A GENETIKY ČLOVĚKA Praha 2014 Prohlášení Prohlašuji, že jsem diplomovou práci na téma ,,Genetické vazby v okolí Rudého moře hodnocené pomocí mtDNA“ zpracovala samostatně a že jsem uvedla všechny použité infromační zdroje a literaturu. Tato práce ani její podstatná část nebyla předložena k získání jiného nebo stejného akademického titulu. V Praze dne…………….. Podpis…………………… Martina Čížková Poděkování V první řadě patří mé poděkování vedoucímu této diplomové práce doc. Mgr. Viktoru Černému, Dr. za cenné rady, odborné vedení a především trpělivost v průběhu celého zpracovávání. Dále bych ráda poděkovala všem členům archeogenetické laboratoře Archeologického ústavu AVČR, Praha, v.v.i., jmenovitě Mgr. Elišce Podgorné za ochotu a pomoc při zaškolování v laboratoři a také Mgr. Editě Priehodové a Mgr. Janě Nováčkové za rady. V neposlední řadě děkuji své rodině za podporu a pomoc v průběhu celého studia. ABSTRAKT Okolí Rudého moře je jedním z důležitých regionů, které nám umožňují poodkrýt stopy evoluce anatomicky moderního člověka. Kromě otázek souvisejících s jeho šířením z Afriky je tato oblast významná i z hlediska vzájemného vlivu afrických a arabských populací, které se po delším období vzájemné izolace a genetické diferenciace související s klimatickými změnami v pleistocénu a následným rozvojem námořních i pozemních tras v holocénu začaly dostávat do stále častějšího kontaktu. Ačkoliv byla v rámci této oblasti provedena již řada archeogenetických analýz, některé otázky týkající se pozdějšího vývoje zůstávají dosud nedostatečně zodpovězené, především z důvodu nedostatečného materiálového zastoupení. Tato práce je zaměřena na analýzu 200 sekvencí mtDNA čtyř súdánských populací – dvou populací kočovných pastevců Rašajda a Bedža žijící v těsném sousedství v okolí města Kassala a hovořící odlišnými jazyky a dvou populací z údolí Nilu žijící usedlým způsobem života. Prostřednictvím intrapopulační analýzy byla zjištěna mnohem vyšší diverzita u usedlých populací (v této práci jsou reprezentovány Núbijci a Araby). Při hodnocení interpopulační variability a genetických vzdáleností v rámci dalších 46 populací z oblasti Rudého moře bylo dále zjištěno, že ačkoliv žijí Rašajdové a Bedžové v současné době v těsném sousedství, geneticky jsou lokalizovány v naprosto odlišných kontextech. Rašajdové disponují typicky arabským mtDNA genofondem, což je vzhledem k jejich relativně nedávné migraci na africké území očekávané. Naopak u populace Bedža je zcela zřejmé africké haplotypové zastoupení, ačkoliv některé starší práce předpokládají její původ na Arabském poloostrově. KLÍČOVÁ SLOVA mtDNA, archeogenetika, subsistenční strategie, Rudé moře, genetická diverzita, haploskupina, fylogenetická síť ABSTRACT The Red Sea region is one of the important places that allow us to uncover traces of the evolution of anatomically modern humans. Besides the questions related to its expansion out of Africa, this region is also important in terms of the mutual influence between populations of Africa and Arabia that after a long period of isolation and genetic differentiation related to climate change in the Pleistocene and subsequent development of seaways and land routes in the Holocene began to contact with each other more frequently. Number of genetic analyzes has been done but some issues concerning on the later development still remain inadequately answered, mainly because of insufficient material. This work is focused on the analysis of 200 mtDNA sequences of four Sudanese populations - two populations of nomadic herdsmen Rashaida and Beja living in the close neighborhood around the city of Kassala and speaking different languages and two populations of the Nile Valley with settled way of life. Analysis of the intrapopulation level revealed much higher diversity of the sedentary populations (in this work the sedentary populations are represented by the Nubians and Arabs). Interpopulation variability and genetic distances within other 46 populations of the Red Sea showed that although Rashaida and Beja people live currently close to each other, they are genetically located in completely different contexts. Rashaida people have typically Arabian mtDNA gene pool, which is expected due to their relatively recent migration to Africa. In contrast, the population of Beja presents typical African mtDNA portrait of haplotypes, although some earlier works place its origin on the Arabian Peninsula. KEY WORDS mtDNA, archaeogenetics, subsistent strategy, Red Sea, genetic diversity, haplogroup, network OBSAH 1 Úvod ......................................................................................................................................9 2 Cíle práce ............................................................................................................................ 11 3 Literární přehled ................................................................................................................ 13 3.1 Oblast Rudého moře – geografie ................................................................................... 13 3.1.1 Afrika ................................................................................................................... 13 3.1.2 Rudé moře ............................................................................................................ 14 3.1.3 Arabský poloostrov ............................................................................................... 15 3.2 Africké jazyky .............................................................................................................. 16 3.2.1 Afroasijské jazyky ................................................................................................ 16 3.2.1.a Čadská větev ..................................................................................................... 17 3.2.1.b Kušitská větev ................................................................................................... 17 3.2.1.c Severní afroasijská větev ................................................................................... 18 3.2.1.d Berberská větev ................................................................................................. 18 3.2.1.e Semitská větev .................................................................................................. 18 3.2.1.f Egyptská větev .................................................................................................. 19 3.2.2 Nilosaharské jazyky .............................................................................................. 19 3.2.3 Nigerokonžské jazyky ........................................................................................... 20 3.2.4 Khoisanské jazyky ................................................................................................ 21 3.3 Historie Súdánu ............................................................................................................ 22 3.4 Historie súdánských populací ........................................................................................ 27 3.4.1 Bedžové ................................................................................................................ 27 3.4.1.a Medjay .............................................................................................................. 27 3.4.1.b Blemmyové ....................................................................................................... 28 3.4.1.c Sociokulturní kontext ........................................................................................ 28 3.4.2 Rašajdové ............................................................................................................. 29 3.4.2.a Sociokulturní kontext ........................................................................................ 30 3.4.3 Núbijci .................................................................................................................. 31 6 3.4.4 Súdánští Arabové .................................................................................................. 32 3.5 MtDNA ........................................................................................................................ 33 3.5.1 Metody studia populační genetiky ......................................................................... 35 3.5.1.a Populační vyhodnocení ..................................................................................... 35 3.5.1.b Fylogenetické vyhodnocení ............................................................................... 36 3.5.2 Původ afrických mtDNA haploskupin ................................................................... 38 3.5.3 Původ arabských mtDNA haploskupin .................................................................. 40 4 Materiál .............................................................................................................................. 42 5 Metody ................................................................................................................................ 43 5.1 Izolace DNA................................................................................................................. 43 5.2 Amplifikace úseků HVS-1 – reakce PCR ...................................................................... 43 5.3 Gelová elektroforéza
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