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===Genetics and genomics=== {{see also|Genetic history of Europe}} [[File:Map of Early Neolithic migrations.jpg|thumb|Neolithic migrations {{circa|5000}}–4000 BC. The people of the Proto-Indo-European [[Sredny Stog culture]] were the result of a genetic admixture between the [[Eastern Hunter-Gatherer|Eastern European hunter-gatherers]] and [[Caucasus hunter-gatherer]]s.]] Ukrainians, like most Europeans, largely descend from three distinct lineages:<ref name="Indo-European"/> Mesolithic [[hunter-gatherer]]s, descended from populations associated with the Paleolithic [[Epigravettian]] culture;<ref>{{cite journal |author1=Posth, C. |author2=Yu, H. |author3=Ghalichi, A. |title=Palaeogenomics of Upper Palaeolithic to Neolithic European hunter-gatherers |journal=[[Nature (journal)|Nature]] |date=2023 |volume=615 |issue=2 March 2023 |pages=117–126 |doi=10.1038/s41586-023-05726-0 |pmid=36859578 |pmc=9977688 |bibcode=2023Natur.615..117P }}</ref> Neolithic [[Early European Farmers]] who migrated from Anatolia during the [[Neolithic Revolution]] 9,000 years ago;<ref>{{cite journal |last1=Gibbons |first1=Ann |title=Thousands of horsemen may have swept into Bronze Age Europe, transforming the local population |journal=Science |date=21 February 2017 |url=https://www.science.org/content/article/thousands-horsemen-may-have-swept-bronze-age-europe-transforming-local-population |access-date=25 March 2023 |archive-date=25 September 2022 |archive-url=https://web.archive.org/web/20220925154535/https://www.science.org/content/article/thousands-horsemen-may-have-swept-bronze-age-europe-transforming-local-population |url-status=live }}</ref> and [[Yamnaya culture|Yamnaya]] [[Western Steppe Herders|Steppe pastoralists]] who expanded into Europe from the [[Pontic–Caspian steppe]] of Ukraine and southern Russia in the context of [[Indo-European migrations]] 5,000 years ago.<ref name="Indo-European">{{Cite journal|last1=Haak |first1=Wolfgang |last2=Lazaridis |first2=Iosif |last3=Patterson |first3=Nick |last4=Rohland |first4=Nadin |last5=Mallick |first5=Swapan |last6=Llamas |first6=Bastien |last7=Brandt |first7=Guido |last8=Nordenfelt |first8=Susanne |last9=Harney |first9=Eadaoin |last10=Stewardson |first10=Kristin |last11=Fu |first11=Qiaomei |date=11 June 2015 |title=Massive migration from the steppe was a source for Indo-European languages in Europe |journal=[[Nature (journal)|Nature]] |volume=522 |issue=7555 |pages=207–211 |doi=10.1038/nature14317 |issn=0028-0836 |pmc=5048219 |pmid=25731166 |bibcode=2015Natur.522..207H |arxiv=1502.02783}}</ref> In a survey of 97 genomes for diversity in full genome sequences among self-identified Ukrainians from Ukraine, a study identified more than 13 million genetic variants, representing about a quarter of the total genetic diversity discovered in Europe.<ref>{{cite journal |last1= Oleksyk |first1= Taras K |last2= Wolfsberger |first2= Walter W |last3= Weber |first3= Alexandra M |last4= Shchubelka |first4= Khrystyna |last5= Oleksyk |first5= Olga |last6= Levchuk |first6= Olga |last7= Patrus |first7= Alla |last8= Lazar |first8= Nelya |last9= Castro-Marquez |first9= Stephanie O |last10= Hasynets |first10= Yaroslava |last11= Boldyzhar |first11= Patricia |last12= Neymet |first12= Mikhailo |last13= Urbanovych |first13= Alina |last14= Stakhovska |first14= Viktoriya |last15= Malyar |first15= Kateryna |last16= Chervyakova |first16= Svitlana |last17= Podoroha |first17= Olena |last18= Kovalchuk |first18= Natalia |last19= Rodriguez-Flores |first19= Juan L |last20= Zhou |first20= Weichen |last21= Medley |first21= Sarah |last22= Battistuzzi |first22= Fabia |last23= Liu |first23= Ryan |last24= Hou |first24= Yong |last25= Chen |first25= Siru |last26= Yang |first26= Huanming |last27= Yeager |first27= Meredith |last28= Dean |first28= Michael |last29= Mills |first29= Ryan |last30= Smolanka |first30= Volodymyr |date= 2021 |title= Genome diversity in Ukraine |url= https://academic.oup.com/gigascience/article/10/1/giaa159/6079618 |journal= GigaScience |volume= 10 |issue= 1 |pages= |doi= 10.1093/gigascience/giaa159 |pmid= 33438729 |pmc= 7804371 |access-date= 17 January 2021 |doi-access= free |archive-date= 12 March 2022 |archive-url= https://web.archive.org/web/20220312102545/https://academic.oup.com/gigascience/article/10/1/giaa159/6079618 |url-status= live }}</ref> Among these nearly 500,000 are previously undocumented and likely to be unique for this population. Medically relevant mutations whose prevalence in the Ukrainian genomes differed significantly compared to other European genome sequences, particularly from Western Europe and Russia.<ref>{{Cite web |date=2021-01-13 |title=Large study provides new understanding of genome diversity in Ukrainian population |url=https://www.news-medical.net/news/20210113/Large-study-provides-new-understanding-genome-diversity-in-Ukrainian-population.aspx |access-date=2024-02-04 |website=News-Medical |language=en}}</ref> Ukrainian genomes form a single cluster positioned between the Northern on one side, and Western European populations on the other.[https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7804371/][[File:Principal Component Analysis of European populations from the Genome Ukraine Project.png|thumb|Principal Component Analysis of European populations from the Genome Ukraine Project]] There was a significant overlap with Central European populations as well as with people from the [[Balkans]]. [[File:Structure plot of European populations from the Genome Ukraine Project.jpg|thumb|Structure plot of European populations from the Genome Ukraine Project]] In addition to the close geographic distance between these populations, this may also reflect the insufficient representation of samples from the surrounding populations.{{citation needed|date=February 2024}} The Ukrainian gene-pool includes the following Y-[[haplogroup]]s, in order from the most prevalent:<ref>Kushniarevich A, Utevska O (2015) "Genetic Heritage of the Balto-Slavic Speaking Populations: A Synthesis of Autosomal, Mitochondrial and Y-Chromosomal Data"</ref> * [[R1a]] (43%) * [[Haplogroup I-M438|I2a]] (23%) * [[R1b]] (8%) * [[E1b1b]] (7%) * [[Haplogroup I-M253|I1]] (5%) * [[Haplogroup N-M231|N1]] (5%) * [[Haplogroup J-M172|J2]] (4%) * [[Haplogroup G-M201|G]] (3%) * [[Haplogroup T-M184|T]] (1%) Roughly all R1a Ukrainians carry [[R1a]]-Z282; R1a-Z282 has been found significantly only in Eastern Europe.<ref name="Luca Giacomo Benincasa Popa 2006 pp. 132–139">{{cite journal |format= PDF |last1= Di Luca | first1= F. | last2= Giacomo | first2= F. | last3= Benincasa | first3= T. | last4= Popa | first4= L.O. | last5= Banyko |first5= J. |last6= Kracmarova |first6= A. |last7= Malaspina |first7= P. |last8= Novelletto |first8= A. |last9= Brdicka |first9= R. |title= Y-chromosomal variation in the Czech Republic |journal= American Journal of Physical Anthropology |volume= 132 |issue= 1 |year= 2006 |pages= 132–139 |url= https://www.researchgate.net/publication/6718583 | doi= 10.1002/ajpa.20500 |access-date= 16 March 2016 |pmid= 17078035 |hdl= 2108/35058 |hdl-access= free}}</ref> [[Chernivtsi Oblast]] is the only region in Ukraine where Haplogroup I2a occurs more frequently than R1a, much less frequent even in [[Ivano-Frankivsk Oblast]].<ref>{{cite journal |last1= Utevska |first1= O. M. |last2= Chukhraeva |first2= M. I. |last3= Agdzhoyan |first3= A. T. |last4= Atramentova |first4= L. A. |last5= Balanovska |first5= E. V. |last6= Balanovsky |first6= O. P. |title= Populations of Transcarpathia and Bukovina on the genetic landscape of surrounding regions |journal= Visnyk of Dnipropetrovsk University. Biology, Medicine |date= 21 September 2015 |volume= 6 |issue= 2 |pages= 133–140 |doi= 10.15421/021524 |url= http://oaji.net/articles/2015/922-1450176147.pdf |doi-access= free |access-date= 22 October 2016 |archive-date= 24 February 2021 |archive-url= https://web.archive.org/web/20210224025357/http://oaji.net/articles/2015/922-1450176147.pdf |url-status= live }}</ref> In comparison to their northern and eastern neighbors, Ukrainians have a similar percentage of [[Haplogroup R1a| Haplogroup R1a-Z280]] (43%) in their population—compare [[Belarusians]], [[Russians]], and [[Lithuanians]] and (55%, 46%, and 42% respectively). Populations in Eastern Europe which have never been Slavic do as well. Ukrainians in Chernivtsi Oblast (near the Romanian border) have a higher percentage of [[Haplogroup I (Y-DNA)|I2a]] as opposed to R1a, which is typical of the Balkan region, but a smaller percentage than Russians of the N1c1 lineage found among Finno-Ugric, Baltic, and Siberian populations, and also less [[Haplogroup R1b|R1b]] than [[West Slavs]].<ref name="Semino2000">{{cite journal | author1= Semino O. | author2= Passarino G. |author3= Oefner P.J. |author4= Lin A.A. |author5= Arbuzova S. |author6= Beckman L.E. |author7= De Benedictis G. |author8= Francalacci P. |author9= Kouvatsi A. |author10= Limborska S. |author11= Marcikiae M. |author12= Mika A. |author13= Mika B. |author14= Primorac D. |author15= Santachiara-Benerecetti A.S. |author16= Cavalli-Sforza L.L. |author17= Underhill P.A. |title= The Genetic Legacy of Paleolithic ''Homo sapiens sapiens'' in Extant Europeans: A Y Chromosome Perspective |journal= Science | volume= 290 | issue= 5494 | year= 2000 | pages= 1155–1159 |doi= 10.1126/science.290.5494.1155 |pmid= 11073453| bibcode= 2000Sci...290.1155S }}</ref><ref name="Varzari2006">Alexander Varzari, "Population History of the Dniester-Carpathians: Evidence from Alu Insertion and Y-Chromosome Polymorphisms" (2006)</ref><ref name="Pericic2005" > Marijana Peričić et al. 2005, [https://web.archive.org/web/20110624174942/http://mbe.oxfordjournals.org/content/22/10/1964.full High-Resolution Phylogenetic Analysis of Southeastern Europe Traces Major Episodes of Paternal Gene Flow Among Slavic Populations.] </ref> In terms of haplogroup distribution, the genetic pattern of Ukrainians most closely resembles that of Belarusians. The presence of the N1c lineage is explained by a contribution of the assimilated [[Finno-Ugric peoples|Finno-Ugric]] tribes.<ref> {{cite journal | title= Gene Pool Structure of Eastern Ukrainians as Inferred from the Y-Chromosome Haplogroups | journal= Russian Journal of Genetics | volume= 40 | issue= 3 | pages= 326–331 | date= 1 March 2004 | doi= 10.1023/B:RUGE.0000021635.80528.2f | last1 = Kharkov |first1 = V. N. |last2= Stepanov | first2= V. A. | last3= Borinskaya | first3= S. A. | last4= Kozhekbaeva | first4= Zh. M. | last5= Gusar | first5= V. A. | last6= Grechanina |first6= E. Ya. |last7= Puzyrev |first7= V. P. |last8= Khusnutdinova |first8= E. K. |last9= Yankovsky | first9= N. K. | s2cid= 25907265 }} </ref>
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