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== Genetics == {{Main|Genetic history of East Asians|Peopling of China}} === Internal genetic structure === The internal genetic structure of the Han Chinese is consistent with the vast geographical expanse of China and the recorded history of large [[Northern and Southern dynasties#Demographics|migratory waves]] over the past several millennia have engendered the emergence of the diverse [[Han Chinese subgroups|Han subgroups]] displaying slight but discernible physical and physiological differences. Although genetically similar, Han Chinese subgroups exhibit a north–south stratification in their genetics,<ref name="GS">{{cite journal |last1=Chen |first1=Jieming |last2=Zheng |first2=Houfeng |last3=Bei |first3=Jin-Xin |last4=Sun |first4=Liangdan |last5=Jia |first5=Wei-hua |last6=Li |first6=Tao |last7=Zhang |first7=Furen |last8=Seielstad |first8=Mark |last9=Zeng |first9=Yi-Xin |last10=Zhang |first10=X |last11=Liu |first11=J |display-authors=9 |year=2009 |title=Genetic Structure of the Han Chinese Population Revealed by Genome-wide SNP Variation |journal=The American Journal of Human Genetics |volume=85 |issue=6 |pages=775–85 |doi=10.1016/j.ajhg.2009.10.016 |pmc=2790583 |pmid=19944401}}</ref><ref name="HC">{{cite journal |last1=Zhao |first1=Yong-Bin |last2=Zhang |first2=Ye |last3=Zhang |first3=Quan-Chao |last4=Li |first4=Hong-Jie |last5=Cui |first5=Ying-Qiu |last6=Xu |first6=Zhi |last7=Jin |first7=Li |last8=Zhou |first8=Hui |last9=Zhu |first9=Hong |date=4 May 2015 |editor1-last=Hofreiter |editor1-first=Michael |title=Ancient DNA Reveals That the Genetic Structure of the Northern Han Chinese Was Shaped Prior to 3,000 Years Ago |journal=PLOS ONE |volume=10 |issue=5 |pages=e0125676 |bibcode=2015PLoSO..1025676Z |doi=10.1371/journal.pone.0125676 |pmc=4418768 |pmid=25938511 |doi-access=free}}</ref><ref name="Genetic evidence supports demic dif">{{cite journal |year=2004 |title=Genetic evidence supports demic diffusion of Han culture |journal=Nature |volume=431 |issue=7006 |pages=302–305 |bibcode=2004Natur.431..302W |doi=10.1038/nature02878 |pmid=15372031 |given1=Bo |surname1=Wen |given2=Hui |surname2=Li |given3=Daru |surname3=Lu |given4=Xiufeng |surname4=Song |given5=Feng |surname5=Zhang |given6=Yungang |surname6=He |given7=Feng |surname7=Li |given8=Yang |surname8=Gao |given9=Xianyun |surname9=Mao |given10=Liang |surname10=Zhang |given11=Ji |surname11=Qian |given12=Jingze |surname12=Tan |given13=Jianzhong |surname13=Jin |given14=Wei |surname14=Huang |given15=Ranjan |surname15=Deka |given16=Bing |surname16=Su |given17=Ranajit |surname17=Chakraborty |given18=Li |surname18=Jin}}</ref><ref name="PP">{{Cite journal |last1=Gan |first1=Rui-Jing |last2=Pan |first2=Shang-Ling |last3=Mustavich |first3=Laura F. |last4=Qin |first4=Zhen-Dong |last5=Cai |first5=Xiao-Yun |last6=Qian |first6=Ji |last7=Liu |first7=Cheng-Wu |last8=Peng |first8=Jun-Hua |last9=Li |first9=Shi-Lin |last10=Xu |first10=Jie-Shun |last11=Jin |first11=Li |last12=Li |first12=Hui |year=2008 |title=Pinghua population as an exception of Han Chinese's coherent genetic structure |journal=Journal of Human Genetics |volume=53 |issue=4 |pages=303–13 |doi=10.1007/s10038-008-0250-x |pmid=18270655 |doi-access=free}}</ref> with centrally placed populations acting as conduits for outlying ones.<ref name="GS" /> Despite there being no clear genetic divide between the north and south due to the Han Chinese being a clinal population, many studies simply categorize the Han Chinese into two subgroups out of convenience: northern and southern Han Chinese.<ref>{{cite journal |last1=Zhao |first1=Yong-Bin |last2=Zhang |first2=Ye |last3=Zhang |first3=Quan-Chao |last4=Li |first4=Hong-Jie |last5=Cui |first5=Ying-Qiu |last6=Xu |first6=Zhi |last7=Jin |first7=Li |last8=Zhou |first8=Hui |last9=Zhu |first9=Hong |date=2015 |title=Ancient DNA Reveals That the Genetic Structure of the Northern Han Chinese Was Shaped Prior to three-thousand Years Ago |journal=[[PLoS ONE]] |volume=10 |issue=5 |page=e0125676 |bibcode=2015PLoSO..1025676Z |doi=10.1371/journal.pone.0125676 |pmc=4418768 |pmid=25938511 |doi-access=free}}</ref> Several genetic studies show that both northern and southern Han Chinese share ancestry with [[Peopling of China#Archaeogenetic studies in the Central Plains|Neolithic Chinese populations from the Central Plains]], who were an admixture of local northern and southern populations.<ref>{{cite journal |last1=Ning |first1=Chao |last2=Li |first2=Tianjiao |last3=Wang |first3=Ke |last4=Zhang |first4=Fan |last5=Li |first5=Tao |last6=Wu |first6=Xiyan |last7=Gao |first7=Shizhu |last8=Zhang |first8=Quanchao |last9=Zhang |first9=Hai |last10=Hudson |first10=Mark J. |last11=Dong |first11=Guanghui |last12=Wu |first12=Sihao |last13=Fang |first13=Yanming |last14=Liu |first14=Chen |last15=Feng |first15=Chunyan |last16=Li |first16=Wei |last17=Han |first17=Tao |last18=Li |first18=Ruo |last19=Wei |first19=Jian |last20=Zhu |first20=Yonggang |last21=Zhou |first21=Yawei |last22=Wang |first22=Chuan-Chao |last23=Fan |first23=Shengying |last24=Xiong |first24=Zenglong |last25=Sun |first25=Zhouyong |last26=Ye |first26=Maolin |last27=Sun |first27=Lei |last28=Wu |first28=Xiaohong |last29=Liang |first29=Fawei |last30=Cao |first30=Yanpeng |last31=Wei |first31=Xingtao |last32=Zhu |first32=Hong |last33=Zhou |first33=Hui |last34=Krause |first34=Johannes |last35=Robbeets |first35=Martine |last36=Jeong |first36=Choongwon |last37=Cui |first37=Yinqiu |title=Ancient genomes from northern China suggest links between subsistence changes and human migration |journal=Nature Communications |date=June 2020 |volume=11 |issue=1 |page=2700 |doi=10.1038/s41467-020-16557-2 |pmid=32483115 |pmc=7264253 }}</ref><ref>{{cite journal |last1=Gao |first1=Shizhu |last2=Cui |first2=Yinqiu |title=Ancient genomes reveal the origin and evolutionary history of Chinese populations |journal=Frontiers in Earth Science |date=10 January 2023 |volume=10 |doi=10.3389/feart.2022.1059196 |doi-access=free }}</ref><ref>{{cite journal |last1=Wu |first1=Xiyan |last2=Ding |first2=Baoxu |last3=Nie |first3=Linyi |last4=Zhong |first4=Canshuo |last5=Liu |first5=Pengxiang |last6=Liang |first6=Jingteng |last7=Wang |first7=Lin |last8=Gao |first8=Xiangping |last9=Wei |first9=Jiyin |last10=Zhou |first10=Yawei |title=Genomic insights into the complex demographic history and inbreeding phenomena during Zhou Dynasty on the Central Plains of China |journal=Frontiers in Microbiology |date=13 September 2024 |volume=15 |doi=10.3389/fmicb.2024.1471740 |doi-access=free |pmid=39345259 |pmc=11427373 }}</ref><ref>{{cite journal |last1=Gao |first1=Xing |title=Explore the ancient roots of the Huaxia people and Chinese civilization |journal=International Journal of Anthropology and Ethnology |date=17 June 2024 |volume=8 |issue=1 |doi=10.1186/s41257-024-00111-9 |doi-access=free }}</ref> A 2020 study models both the northern Han Chinese and southern Han Chinese as having Neolithic Yellow River ([[Sino-Tibetan peoples|Sino-Tibetan]]) and [[Kra–Dai-speaking peoples|Kra-Dai]] ancestries, although Kra-Dai ancestry is more common in southern Han.<ref name=":0">{{cite report |type=Preprint |doi=10.1101/2020.11.08.373225 |title=Genomic Insights into the Demographic History of Southern Chinese |date=2020 |last1=Huang |first1=Xiufeng |last2=Xia |first2=Zi-Yang |last3=Bin |first3=Xiaoyun |last4=He |first4=Guanglin |last5=Guo |first5=Jianxin |last6=Lin |first6=Chaowen |last7=Yin |first7=Lianfei |last8=Zhao |first8=Jing |last9=Ma |first9=Zhuofei |last10=Ma |first10=Fuwei |last11=Li |first11=Yingxiang |last12=Hu |first12=Rong |last13=Wei |first13=Lan-Hai |last14=Wang |first14=Chuan-Chao }}</ref> The massive Han sub-groups differ in some of their ancestral components despite a shared Neolithic Yellow River ancestry, reflective of their vast demographic history. They tend to share some maternal ancestry with geographically close minority groups. For example, southern Han show evidence of being admixed with populations of Tai-Kadai and Austronesian ancestry. Southwestern Han show admixture with [[Hmong people|Hmong-Mien]] speakers, whilst Northwestern Han have very minor West Eurasian ancestral components, dating 4,500–1,200/1,300 years ago. Northeastern Han have more Yellow River Basin and ancient Northeast Asian ancestry than Southern Han Chinese.<ref>{{Cite journal |last1=Zhou |first1=Jingbin |last2=Zhang |first2=Xianpeng |last3=Li |first3=Xin |last4=Sui |first4=Jie |date=2022 |title=Genetic structure and demographic history of Northern Han people in Liaoning Province inferred from genome-wide array data |journal=Frontiers in Ecology and Evolution |volume=10 |doi=10.3389/fevo.2022.1014024 |doi-access=free}}</ref> Other studies also show bidirectional gene flow between Hmong-Mien-related and Kra-Dai-related ancestries in southern China.<ref name=":0" /><ref>{{cite journal |last1=Wang |first1=Jiawen |last2=Wu |first2=Jun |last3=Sun |first3=Qiuxia |last4=Wu |first4=Qian |last5=Li |first5=Youjing |last6=Duan |first6=Shuhan |last7=Yang |first7=Lin |last8=Wu |first8=Wenxin |last9=Wang |first9=Zheng |last10=Liu |first10=Yan |last11=Tang |first11=Renkuan |last12=Yang |first12=Junbao |last13=Wang |first13=Chuanchao |last14=Liu |first14=Chao |last15=Xu |first15=Jianwei |last16=Wang |first16=Mengge |last17=He |first17=Guanglin |title=Extensive genetic admixture between Tai-Kadai-speaking people and their neighbours in the northeastern region of the Yungui Plateau inferred from genome-wide variations |journal=BMC Genomics |date=12 June 2023 |volume=24 |issue=1 |page=317 |doi=10.1186/s12864-023-09412-3 |doi-access=free |pmid=37308851 |pmc=10259048 }}</ref> Variation notwithstanding, Han Chinese subgroups are genetically closer to each other than they each are to their Korean and Yamato neighbors,<ref name="pmid 29636655">{{Cite journal |last1=Wang |first1=Yuchen |last2=Lu |first2=Dongsheng |last3=Chung |first3=Yeun-Jun |last4=Xu |first4=Shuhua |year=2018 |title=Genetic structure, divergence and admixture of Han Chinese, Japanese and Korean populations |journal=Hereditas |publication-date=6 April 2018 |volume=155 |page=19 |doi=10.1186/s41065-018-0057-5 |pmc=5889524 |pmid=29636655 |doi-access=free}}</ref> to whom they are also genetically close in general. The close genetic relationship between the Han across the entirety of China has led to their characterization as having a "coherent genetic structure".<ref name="HC" /><ref name="PP" /> The two notable exceptions to this structure are [[Pinghua]] and [[Tanka people]],<ref>{{cite journal |author=McFadzean A.J.S., Todd D. |year=1971 |title=Cooley's anaemia among the tanka of South China |journal=Transactions of the Royal Society of Tropical Medicine and Hygiene |volume=65 |issue=1 |pages=59–62 |doi=10.1016/0035-9203(71)90185-4 |pmid=5092429}}</ref> who on their patrilines, bear a closer genetic resemblance to aboriginal peoples, but have Han matrilines. The Tanka are a group of boat-dwellers who speak a Sinitic language and who claim Han ancestry, but who have traditionally faced severe discrimination from the other Southern Han subgroups. Unlike the Guangdong, Fujian and Hainan Han (whose dominant Y-chromosome haplotype is the Han patriline O2-M122), the Tanka have been shown instead to have a predominantly non-Han patriline similar to Daic peoples from Guizhou.<ref name="UG">{{Cite journal |last1=Luo |first1=Xiao-Qin |last2=Du |first2=Pan-Xin |last3=Wang |first3=Ling-Xiang |last4=Zhou |first4=Bo-Yan |last5=Li |first5=Yu-Chun |last6=Zheng |first6=Hong-Xiang |last7=Wei |first7=Lan-Hai |last8=Liu |first8=Jun-Jian |last9=Sun |first9=Chang |last10=Meng |first10=Hai-Liang |last11=Tan |first11=Jing-Ze |last12=Su |first12=Wen-Jing |last13=Wen |first13=Shao-Qing |last14=Li |first14=Hui |date=2019 |title=Uniparental Genetic Analyses Reveal the Major Origin of Fujian Tanka from Ancient Indigenous Daic Populations |journal=Human Biology |volume=91 |issue=4 |pages=257–277 |doi=10.13110/humanbiology.91.4.05 |pmid=32767896 }}</ref><ref name="GF">{{Cite journal |last=He |first=Guanglin |date=16 November 2021 |title=The genomic formation of the Tanka people, an isolated "gypsies in the water" in the coastal region of southeast China |journal=Wiley, American Journal of Biological Anthropology}}</ref> However, matrilineally, the Tanka are closely clustered with the Hakka Han and Teochew Han, rather than with Austronesian or Austroasiatic populations, thus supporting an admixture hypothesis and validating, even if only partially, their own claims to Han ancestry.<ref name="UG" /><ref name="GF" />[[File:Han chinese genetic 0.png|thumb|310x310px|A PCA graph illustrates the genetic differences among Han Chinese subgroups.<ref>{{cite journal |display-authors=6 |vauthors=Cao Y, Li L, Xu M, Feng Z, Sun X, Lu J, Xu Y, Du P, Wang T, Hu R, Ye Z, Shi L, Tang X, Yan L, Gao Z, Chen G, Zhang Y, Chen L, Ning G, Bi Y, Wang W |date=September 2020 |title=The ChinaMAP analytics of deep whole genome sequences in 10,588 individuals |journal=Cell Research |volume=30 |issue=9 |pages=717–731 |doi=10.1038/s41422-020-0322-9 |pmc=7609296 |pmid=32355288}}</ref>]] === Demic diffusion and north-south differences === The estimated genetic contribution of northern Han to southern Han is substantial in the ancestral patrilineage in addition to a geographic [[cline (biology)|cline]] that exists for the corresponding matrilineage. These genetic findings align with historical records outlining the continuous and large migratory waves of northern Han Chinese inhabitants settling in southern China, escaping dynastic changes, geopolitical upheavals, instability, warfare and famine.<ref>{{cite book |last=Gernet |first=Jacques |url=https://archive.org/details/historyofchinese00gern_0 |title=A History of Chinese Civilization |publisher=Cambridge University Press |year=1996 |isbn=978-0-521-49781-7 |edition=2nd}}</ref><ref name="Genetic evidence supports demic dif" /><ref>{{Cite book |last1=Crawford |first1=Dorothy H. |title=Cancer Virus: The story of Epstein-Barr Virus |last2=Rickinson |first2=Alan |last3=Johannessen |first3=Ingolfur |publisher=Oxford University Press |year=2014 |publication-date=March 14, 2014 |pages=98}}</ref><ref name="Springer" /><ref name="Six Dynasties" /><ref name="MS" /><ref name="QC" /><ref name="China: A New Cultural History" /><ref name="The History of China" /><ref name="books.google.com" /><ref name="Hunter Publishing, Inc" /><ref name="China: An Environmental History" /><ref name="Lexington Books" /><ref name="Cambridge University Press" /><ref name="Newspaper Enterprise Limited" /><ref name="Rowman Altamira" />{{Excessive citations inline|date=September 2024}} The subsequent intermarriages between northern Han migrants and southern aborigines over the past few thousand years gave rise to modern Chinese demographics - a Han Chinese super-majority and minority non-Han Chinese indigenous peoples.<ref name="Genetic evidence supports demic dif" /> Han Chinese from Fujian and Guangdong show excessive ancestries from Late Neolithic Fujianese sources (35.0–40.3%), which are more significant in modern [[Amis people|Ami]], [[Atayal people|Atayal]] and [[Kankanaey people|Kankanaey]] (66.9–74.3%), and less significant in Han Chinese from Zhejiang (22%), Jiangsu (17%) and Shandong (8%).<ref>{{Cite journal |last1=Huang |first1=Xiufeng |last2=Xia |first2=Zi-Yang |last3=Bin |first3=Xiaoyun |last4=He |first4=Guanglin |display-authors=3 |date=2022 |title=Genomic Insights Into the Demographic History of the Southern Chinese |journal=Frontiers in Ecology and Evolution |volume=10 |doi=10.3389/fevo.2022.853391 |doi-access=free}}</ref> Ancient population admixture with Ami and Atayal also exists for Han Chinese from Guangdong and [[Sichuan]]<ref>{{cite journal |last1=Chiang |first1=Charleston W K |last2=Mangul |first2=Serghei |last3=Robles |first3=Christopher |last4=Sankararaman |first4=Sriram |title=A Comprehensive Map of Genetic Variation in the World's Largest Ethnic Group—Han Chinese |journal=Molecular Biology and Evolution |date=November 2018 |volume=35 |issue=11 |pages=2736–2750 |doi=10.1093/molbev/msy170 |pmid=30169787 |pmc=6693441 }}</ref><ref>{{cite journal |last1=Lo |first1=Yun-Hua |last2=Cheng |first2=Hsueh-Chien |last3=Hsiung |first3=Chia-Ni |last4=Yang |first4=Show-Ling |last5=Wang |first5=Han-Yu |last6=Peng |first6=Chia-Wei |last7=Chen |first7=Chun-Yu |last8=Lin |first8=Kung-Ping |last9=Kang |first9=Mei-Ling |last10=Chen |first10=Chien-Hsiun |last11=Chu |first11=Hou-Wei |last12=Lin |first12=Chiao-Feng |last13=Lee |first13=Mei-Hsuan |last14=Liu |first14=Quintin |last15=Satta |first15=Yoko |last16=Lin |first16=Cheng-Jui |last17=Lin |first17=Marie |last18=Chaw |first18=Shu-Miaw |last19=Loo |first19=Jun-Hun |last20=Shen |first20=Chen-Yang |last21=Ko |first21=Wen-Ya |title=Detecting Genetic Ancestry and Adaptation in the Taiwanese Han People |journal=Molecular Biology and Evolution |date=27 September 2021 |volume=38 |issue=10 |pages=4149–4165 |doi=10.1093/molbev/msaa276 |pmid=33170928 |pmc=8476137 }}</ref> and the ancestors of [[Taiwanese Han]].<ref>{{cite journal |last1=Lo |first1=Yun-Hua |last2=Cheng |first2=Hsueh-Chien |last3=Hsiung |first3=Chia-Ni |last4=Yang |first4=Show-Ling |last5=Wang |first5=Han-Yu |last6=Peng |first6=Chia-Wei |last7=Chen |first7=Chun-Yu |last8=Lin |first8=Kung-Ping |last9=Kang |first9=Mei-Ling |last10=Chen |first10=Chien-Hsiun |last11=Chu |first11=Hou-Wei |last12=Lin |first12=Chiao-Feng |last13=Lee |first13=Mei-Hsuan |last14=Liu |first14=Quintin |last15=Satta |first15=Yoko |last16=Lin |first16=Cheng-Jui |last17=Lin |first17=Marie |last18=Chaw |first18=Shu-Miaw |last19=Loo |first19=Jun-Hun |last20=Shen |first20=Chen-Yang |last21=Ko |first21=Wen-Ya |title=Detecting Genetic Ancestry and Adaptation in the Taiwanese Han People |journal=Molecular Biology and Evolution |date=27 September 2021 |volume=38 |issue=10 |pages=4149–4165 |doi=10.1093/molbev/msaa276 |pmid=33170928 |pmc=8476137 }}</ref> Overall, Cantonese people represent the southernmost Han sub-group, splitting from northern Han between the Qin and Tang dynasties,<ref>{{cite journal |last1=Chen |first1=Jieming |last2=Zheng |first2=Houfeng |last3=Bei |first3=Jin-Xin |last4=Sun |first4=Liangdan |last5=Jia |first5=Wei-hua |last6=Li |first6=Tao |last7=Zhang |first7=Furen |last8=Seielstad |first8=Mark |last9=Zeng |first9=Yi-Xin |last10=Zhang |first10=Xuejun |last11=Liu |first11=Jianjun |title=Genetic Structure of the Han Chinese Population Revealed by Genome-wide SNP Variation |journal=The American Journal of Human Genetics |date=December 2009 |volume=85 |issue=6 |pages=775–785 |doi=10.1016/j.ajhg.2009.10.016 |pmid=19944401 |pmc=2790583 }}</ref><ref>{{cite report |type=Preprint |doi=10.1101/2020.07.03.166413 |title=Fine-scale Population Structure and Demographic History of Han Chinese Inferred from Haplotype Network of 111,000 Genomes |date=2020 |last1=Lan |first1=Ao |last2=Kang |first2=Kang |last3=Tang |first3=Senwei |last4=Wu |first4=Xiaoli |last5=Wang |first5=Lizhong |last6=Li |first6=Teng |last7=Weng |first7=Haoyi |last8=Deng |first8=Junjie |last9=Zheng |first9=Qiang |last10=Yao |first10=Xiaotian |last11=Chen |first11=Gang }}</ref> although there is evidence of additional admixture with Teochew and Hakka people.<ref>{{cite journal |last1=Xiao |first1=Cheng |last2=Yang |first2=Xingyi |last3=Yu |first3=Zhonghao |last4=Wu |first4=Weibin |last5=Wang |first5=Yuan |last6=Xu |first6=Quyi |last7=Chen |first7=Ling |title=Genetic polymorphism and variability in the Guangdong Hakka, Teochew, and Cantonese groups: A comprehensive analysis of 19 X-STRs |journal=Annals of Human Genetics |date=September 2023 |volume=87 |issue=5 |pages=232–240 |doi=10.1111/ahg.12518 |pmid=37337755 }}</ref> Other studies suggest that Han Chinese from [[Fujian]] and Taiwan are equally as southern.<ref>{{Cite journal |last=He |first=Guanglin |last2=Li |first2=Yinxiang |last3=Zou |first3=Xing |last4=Yeh |first4=Hui-Yuan |display-authors=3 |date=2021 |title=The northern gene flow into southeastern East Asians inferred from genome wide array genotyping |url=https://www.biorxiv.org/content/10.1101/2021.07.25.453681v1.full |journal=bioRxiv |quote=Fujian and Taiwan populations were localized at the southmost end of the Han Chinese cline and showed a close relationship with modern Tai-Kadai-speaking populations and ancient southern East Asians (Liangdao, Hanben, Tanshishan, Xitoucun and so on). |via=bioRxiv}}</ref> Han Chinese from Guangdong also show affinities with local Kra-Dai and Hmong-Mien groups<ref>{{cite journal |last1=Du |first1=Weian |last2=Feng |first2=Chunlei |last3=Yao |first3=Ting |last4=Xiao |first4=Cheng |last5=Huang |first5=Hongyan |last6=Wu |first6=Weibin |last7=Zhu |first7=Linnan |last8=Qiao |first8=Honghua |last9=Liu |first9=Chao |last10=Chen |first10=Ling |title=Genetic variation and forensic efficiency of 30 indels for three ethnic groups in Guangxi: relationships with other populations |journal=PeerJ |date=3 May 2019 |volume=7 |pages=e6861 |doi=10.7717/peerj.6861 |doi-access=free |pmid=31110924 |pmc=6501771 }}</ref><ref>{{cite journal |last1=Wang |first1=Fei |last2=He |first2=Guanglin |last3=Wang |first3=Zheng |last4=Wang |first4=Mengge |last5=Liu |first5=Jing |last6=Zou |first6=Xing |last7=Wang |first7=Shouyu |last8=Song |first8=Mengyuan |last9=Ye |first9=Ziwei |last10=Xie |first10=Mingkun |last11=Hou |first11=Yiping |title=Population Genetics and Forensic Efficiency of 30 InDel Markers in Four Chinese Ethnic Groups Residing in Sichuan |journal=Forensic Sciences Research |date=3 July 2022 |volume=7 |issue=3 |pages=498–502 |doi=10.1080/20961790.2020.1737470 |pmid=36353334 |pmc=9639512 }}</ref><ref>{{cite report |type=Preprint |doi=10.1101/2021.06.19.449013 |title=New insights from the combined discrimination of modern/Ancient genome-wide shared alleles and haplotypes: Differentiated demographic history reconstruction of Tai-Kadai and Sinitic people in South China |date=2021 |last1=Wang |first1=Mengge |last2=He |first2=Guanglin |last3=Zou |first3=Xing |last4=Chen |first4=Pengyu |last5=Wang |first5=Zheng |last6=Tang |first6=Renkuan |last7=Yang |first7=Xiaomin |last8=Chen |first8=Jing |last9=Yang |first9=Meiqing |last10=Li |first10=Yingxiang |last11=Liu |first11=Jing |last12=Wang |first12=Fei |last13=Zhao |first13=Jing |last14=Guo |first14=Jianxin |last15=Hu |first15=Rong |last16=Wei |first16=Lan-Hai |last17=Chen |first17=Gang |last18=Yeh |first18=Hui-Yuan |last19=Wang |first19=Chuan-Chao }}</ref><ref>{{cite journal |last1=Wang |first1=Fei |last2=Song |first2=Feng |last3=Song |first3=Mengyuan |last4=Luo |first4=Haibo |last5=Hou |first5=Yiping |title=Genetic structure and paternal admixture of the modern Chinese Zhuang population based on 37 Y-STRs and 233 Y-SNPs |journal=Forensic Science International: Genetics |date=May 2022 |volume=58 |pages=102681 |doi=10.1016/j.fsigen.2022.102681 |pmid=35263703 }}</ref> and cluster with Han Chinese from Guangxi and Hainan, who themselves mixed with the minority peoples of their respective provinces.<ref>{{cite journal |last1=Cong |first1=Pei-Kuan |last2=Bai |first2=Wei-Yang |last3=Li |first3=Jin-Chen |last4=Yang |first4=Meng-Yuan |last5=Khederzadeh |first5=Saber |last6=Gai |first6=Si-Rui |last7=Li |first7=Nan |last8=Liu |first8=Yu-Heng |last9=Yu |first9=Shi-Hui |last10=Zhao |first10=Wei-Wei |last11=Liu |first11=Jun-Quan |last12=Sun |first12=Yi |last13=Zhu |first13=Xiao-Wei |last14=Zhao |first14=Pian-Pian |last15=Xia |first15=Jiang-Wei |last16=Guan |first16=Peng-Lin |last17=Qian |first17=Yu |last18=Tao |first18=Jian-Guo |last19=Xu |first19=Lin |last20=Tian |first20=Geng |last21=Wang |first21=Ping-Yu |last22=Xie |first22=Shu-Yang |last23=Qiu |first23=Mo-Chang |last24=Liu |first24=Ke-Qi |last25=Tang |first25=Bei-Sha |last26=Zheng |first26=Hou-Feng |title=Genomic analyses of 10,376 individuals in the Westlake BioBank for Chinese (WBBC) pilot project |journal=Nature Communications |date=26 May 2022 |volume=13 |issue=1 |page=2939 |doi=10.1038/s41467-022-30526-x |pmid=35618720 |pmc=9135724 }}</ref><ref>{{cite journal |last1=Zou |first1=Xing |last2=Nie |first2=Qianyun |last3=Li |first3=Wenhui |last4=Chen |first4=Yinyu |last5=Song |first5=Tao |last6=Zhang |first6=Peng |title=Genetic variation and phylogenetic analysis of 23 STR in Chinese Han population from Hainan, Southern China |journal=Medicine |date=31 May 2024 |volume=103 |issue=22 |pages=e38428 |doi=10.1097/MD.0000000000038428 |pmid=39259071 |pmc=11142786 }}</ref><ref name=":1">{{cite journal |last1=Sun |first1=Qiuxia |last2=Wang |first2=Mengge |last3=Lu |first3=Tao |last4=Duan |first4=Shuhan |last5=Liu |first5=Yan |last6=Chen |first6=Jing |last7=Wang |first7=Zhiyong |last8=Sun |first8=Yuntao |last9=Li |first9=Xiangping |last10=Wang |first10=Shaomei |last11=Lu |first11=Liuyi |last12=Hu |first12=Liping |last13=Yun |first13=Libing |last14=Yang |first14=Junbao |last15=Yan |first15=Jiangwei |last16=Nie |first16=Shengjie |last17=Zhu |first17=Yanfeng |last18=Chen |first18=Gang |last19=Wang |first19=Chuan-Chao |last20=Liu |first20=Chao |last21=He |first21=Guanglin |last22=Tang |first22=Renkuan |title=Differentiated adaptative genetic architecture and language-related demographical history in South China inferred from 619 genomes from 56 populations |journal=BMC Biology |date=6 March 2024 |volume=22 |issue=1 |page=55 |doi=10.1186/s12915-024-01854-9 |doi-access=free |pmid=38448908 |pmc=10918984 }}</ref> One study shows that Han Chinese from northern Guangxi have more ancestry related to Austronesian, Kra-Dai and Austroasiatic groups, than those from southern Guangxi.<ref name=":1" /> ==== Patrilineal DNA ==== Typical Y-DNA haplogroups of present-day Han Chinese include Haplogroup O-M95, [[Haplogroup O-M122]], [[Haplogroup O-M175]], C,[[Haplogroup N-M231]] and [[Haplogroup Q-M120]].<ref name="AD">{{cite journal |last1=Zhao |first1=Yong-Bin |last2=Zhang |first2=Ye |last3=Zhang |first3=Quan-Chao |last4=Li |first4=Hong-Jie |last5=Cui |first5=Ying-Qiu |last6=Xu |first6=Zhi |last7=Jin |first7=Li |last8=Zhou |first8=Hui |last9=Zhu |first9=Hong |date=4 May 2015 |title=Ancient DNA Reveals That the Genetic Structure of the Northern Han Chinese Was Shaped Prior to 3,000 Years Ago |journal=PLOS ONE |volume=10 |issue=5 |pages=e0125676 |bibcode=2015PLoSO..1025676Z |doi=10.1371/journal.pone.0125676 |pmc=4418768 |pmid=25938511 |doi-access=free}}</ref> The Y-chromosome haplogroup distribution between Southern Han Chinese and Northern Han Chinese populations and principal core component analysis indicates that almost all modern Han Chinese populations form a tight cluster in their Y chromosome: * Haplogroups prevalent in non-Han southern natives such as O1b-M110, O2a1-M88 and O3d-M7, which are prevalent in non-Han southern natives, were observed in 4% of Southern Han Chinese and not at all in the Northern Han.<ref name="table">{{cite journal |last1=Wen |first1=Bo |last2=Li |first2=Hui |last3=Lu |first3=Daru |last4=Song |first4=Xiufeng |last5=Zhang |first5=Feng |last6=He |first6=Yungang |last7=Li |first7=Feng |last8=Gao |first8=Yang |last9=Mao |first9=Xianyun |last10=Zhang |first10=Liang |last11=Qian |first11=Ji |last12=Tan |first12=Jingze |last13=Jin |first13=Jianzhong |last14=Huang |first14=Wei |last15=Deka |first15=Ranjan |last16=Su |first16=Bing |last17=Chakraborty |first17=Ranajit |last18=Jin |first18=Li |title=Genetic evidence supports demic diffusion of Han culture |journal=Nature |date=September 2004 |volume=431 |issue=7006 |pages=302–305 |bibcode=2004Natur.431..302W |doi=10.1038/nature02878 |pmid=15372031 }}</ref><ref name="EJH">{{cite journal |last1=Xue |first1=Fuzhong |last2=Wang |first2=Yi |last3=Xu |first3=Shuhua |last4=Zhang |first4=Feng |last5=Wen |first5=Bo |last6=Wu |first6=Xuesen |last7=Lu |first7=Ming |last8=Deka |first8=Ranjan |last9=Qian |first9=Ji |last10=Jin |first10=L |display-authors=9 |year=2008 |title=A spatial analysis of genetic structure of human populations in China reveals distinct difference between maternal and paternal lineages |journal=European Journal of Human Genetics |volume=16 |issue=6 |pages=705–17 |doi=10.1038/sj.ejhg.5201998 |pmid=18212820 |doi-access=free}}</ref> * Biological research findings have also demonstrated that the paternal lineages Y-DNA O-M119,<ref name="Li et al (2008)">{{cite journal |last1=Li |first1=Hui |year=2008 |title=Paternal genetic affinity between western Austronesians and Daic populations |journal=BMC Evolutionary Biology |volume=8 |issue=1 |page=146 |bibcode=2008BMCEE...8..146L |doi=10.1186/1471-2148-8-146 |pmc=2408594 |pmid=18482451 |doi-access=free}}</ref> O-P201,<ref name="Karafet et al (2010)">{{cite journal |last1=Karafet |first1=Tatiana |last2=Hallmark |first2=B |last3=Cox |first3=M.P. |last4=Sudoyo |first4=H |last5=Downey |first5=S |last6=Lansing |first6=J.S. |last7=Hammer |first7=M.F. |date=August 2010 |title=Major East–West Division Underlies Y Chromosome Stratification across Indonesia |journal=Molecular Biology and Evolution |volume=27 |issue=8 |pages=1833–44 |doi=10.1093/molbev/msq063 |pmid=20207712 |doi-access=free}}</ref> O-P203<ref name="Karafet et al (2010)" /> and O-M95<ref name="Karafet et al 2005">{{cite book |last1=Karafet |first1=Tatiana |last2=Hagberg |first2=L |last3=Hanson |first3=L. A. |last4=Korhonen |first4=T |last5=Leffler |first5=H |last6=Olling |first6=S |year=1981 |chapter=Balinese Y-chromosome perspective on the peopling of Indonesia: genetic contributions from pre-Neolithic hunter-gatherers, Austronesian farmers, and Indian traders |title=Ciba Foundation Symposium 80 - Adhesion and Microorganism Pathogenicity |series=Novartis Foundation Symposia |volume=80 |pages=161–87 |doi=10.1002/9780470720639.ch11 |pmid=6114819|isbn=978-0-470-72063-9 }}</ref> are found in commonly Southern non-Han minorities, less commonly in Southern Han, and even less frequently in Northern Han Chinese.<ref name="Yan et al (2011)">{{cite journal |last1=Yan |first1=Shi |last2=Wang |first2=C.C. |last3=Li |first3=H |last4=Li |first4=S.L. |last5=Jin |first5=L |year=2011 |title=An updated tree of Y-chromosome Haplogroup O and revised phylogenetic positions of mutations P164 and PK4 |journal=European Journal of Human Genetics |volume=19 |issue=9 |pages=1013–15 |doi=10.1038/ejhg.2011.64 |pmc=3179364 |pmid=21505448}}</ref> * Haplogroups O1 and O2 significantly peak in the southeastern coastlines and eastern regions of China respectively, according to one study.<ref>{{cite journal |last1=Li |first1=Jienan |last2=Song |first2=Feng |last3=Lang |first3=Min |last4=Xie |first4=Mingkun |title=Comprehensive insights into the genetic background of Chinese populations using Y chromosome markers |journal=Royal Society Open Science |date=September 2023 |volume=10 |issue=9 |doi=10.1098/rsos.230814 |pmid=37736526 |pmc=10509572 |bibcode=2023RSOS...1030814L }}</ref> Patrilineal DNA indicates the northern Han Chinese were the primary contributors to the paternal gene pool of modern southern Han Chinese.<ref name="AD" /><ref name="table" /><ref name="EJH" /><ref name="Yan et al (2011)" /> The data also indicates that the contribution of southern non-Han aboriginals to the southern Han Chinese genetics is limited. In short, male Han Chinese were the primary drivers of Han Chinese expansion in successive migratory waves from the north into what is now modern southern China as is shown by a greater contribution to the Y-chromosome than the mtDNA from northern to Southern Han.<ref name="Genetic evidence supports demic dif" /> During the Zhou dynasty, or earlier, peoples with haplogroup Q-M120 also contributed to the ethnogenesis of Han Chinese people. This haplogroup is implied to be spread across in the Eurasian steppe and north Asia since it is found among [[Cimmerians]] in [[Moldova]] and Bronze Age natives of [[Khövsgöl Province|Khövsgöl]]. But it is currently near-absent in these regions except for East Asia. In modern China, haplogroup Q-M120 can be found in the northern and eastern regions.<ref>{{cite journal |last1=Sun |first1=Na |last2=Ma |first2=Peng-Cheng |last3=Yan |first3=Shi |last4=Wen |first4=Shao-Qing |last5=Sun |first5=Chang |last6=Du |first6=Pan-Xin |last7=Cheng |first7=Hui-Zhen |last8=Deng |first8=Xiao-Hua |last9=Wang |first9=Chuan-Chao |last10=Wei |first10=Lan-Hai |title=Phylogeography of Y-chromosome haplogroup Q1a1a-M120, a paternal lineage connecting populations in Siberia and East Asia |journal=Annals of Human Biology |date=3 April 2019 |volume=46 |issue=3 |pages=261–266 |doi=10.1080/03014460.2019.1632930 |pmid=31208219 }}</ref> '''Matrilineal DNA''' MtDNA of Han Chinese increases in diversity as one looks from northern to Southern China, which suggests that the influx of male Han Chinese migrants intermarried with the local female non-Han aborigines after arriving in what is now modern-day Fujian, Guangdong, Guangxi, Hainan and other regions of Southern China.<ref name="table" /><ref name="EJH" /> In these populations, the contribution to mtDNA from Han Chinese and indigenous tribes is evenly matched, representing a substantial mtDNA contribution from non-Han groups, collectively known as the Bai Yue or Hundred Yue.<ref name="table" /><ref name="EJH" /> A study by the Chinese Academy of Sciences into the gene frequency data of Han sub-populations and ethnic minorities in China, showed that Han sub-populations in different regions are also genetically quite close to the local ethnic non-Han minorities, meaning that in many cases, the blood of ethnic minorities had mixed into Han genetic substrate through varying degrees of intermarriage, while at the same time, the blood of the Han had also mixed into the genetic substrates of the local ethnic non-Han minorities.<ref>{{cite journal |last1=Du |first1=R |last2=Xiao |first2=C |last3=Cavalli-Sforza |first3=LL |year=1997 |title=Genetic distances between Chinese populations calculated on gene frequencies of 38 loci |journal=Science China Life Sciences |volume=40 |issue=6 |pages=613–21 |doi=10.1007/BF02882691 |pmid=18726285 }}</ref> === Genetic continuity between ancient and modern Han Chinese === The Hengbei archaeological site in Jiang County, southern Shanxi was part of the suburbs of the capital during the Zhou dynasty. Genetic material from human remains in Hengbei have been used to examine the genetic continuity between ancient and modern Han Chinese.<ref name="HC" /> Comparisons of Y chromosome [[single-nucleotide polymorphism]]s (SNPs) and [[mitochondrial DNA]] (mtDNA) between modern northern Han Chinese and 3000-year-old Hengbei samples reveal extreme similarity, confirming genetic continuity between ancient Hengbei inhabitants to present-day northern Han Chinese. This shows that the core genetic structure of Northern Han Chinese was established more than three thousand years ago in the Central Plains Area.<ref name="HC" /><ref>[[Genographic Project#Geno 2.0 Next Generation|Reference Populations - Geno 2.0 Next Generation]]. (2017). [[Genographic Project|The Genographic Project]]. Retrieved 15 May 2017, from [https://genographic.nationalgeographic.com/reference-populations-next-gen/ link.] {{Webarchive|url=https://web.archive.org/web/20170207031612/https://genographic.nationalgeographic.com/reference-populations-next-gen//|date=7 February 2017}}</ref> Additionally, these studies indicate that contemporary northern and southern Han Chinese populations exhibit an almost identical Y-DNA genetic structure, indicating a common paternal descent, corroborating the historical record of Han Chinese migration to the south.<ref name="HC" /> However, a study of [[mitochondrial DNA]] from Yinxu commoner graves in the [[Shang dynasty]] showed similarity with modern northern Han Chinese, but significant differences from southern Han Chinese - indicating admixture on the matriline.<ref>{{Cite conference |last1=Zeng |first1=Wen |last2=Li |first2=Jiawei |last3=Yue |first3=Hongbin |last4=Zhou |first4=Hui |last5=Zhu |first5=Hong |year=2013 |title=Poster: Preliminary Research on Hereditary Features of Yinxu Population |url=https://www.academia.edu/5297877 |conference=82nd Annual Meeting of the American Association of Physical Anthropologists}}</ref> About 2,000 years ago, between the Warring States period and Eastern Han dynasty, the northeast coastlines of China faced an eastward migration from the Central Plains, shaping the genetic structure of local populations to the present. These populations also have more southern East Asian ancestry compared to their predecessors.<ref>{{Cite journal |last1=Wang |first1=Baitong |last2=Hao |first2=Daohua |last3=Xu |first3=Yu |last4=Jin |first4=Li |date=2024 |title=Population expansion from central plain to northern coastal China inferred from ancient human genomes |journal=iScience |volume=27 |issue=12 |doi=10.1016/j.isci.2024.111405 |pmid=39697594 |pmc=11652891 }}</ref> === Closely related East Asian groups === The Han Chinese show a close yet distinguishable genetic relationship with other East Asian populations such as the [[Koreans]] and [[Yamato people|Yamato]].<ref>{{Cite journal |last1=Horai |first1=Satoshi |last2=Murayama |first2=Kumiko |date=1996 |title=mtDNA Polymorphism in East Asian Populations, with Special Reference to the Peopling of Japan |journal=[[American Journal of Human Genetics]] |location=Cambridge, Massachusetts |publisher=[[Cell Press]] |publication-date=1996 |volume=59 |issue=3 |pages=579–590 |pmc=1914908 |pmid=8751859}}</ref><ref>{{cite journal |last1=Shi |first1=Cheng-Min |last2=Liu |first2=Qi |last3=Zhao |first3=Shilei |last4=Chen |first4=Hua |title=Ancestry informative SNP panels for discriminating the major East Asian populations: Han Chinese, Japanese and Korean |journal=Annals of Human Genetics |date=September 2019 |volume=83 |issue=5 |pages=348–354 |doi=10.1111/ahg.12320 |pmid=31025319 }}</ref><ref>{{Cite journal |last1=Pan |first1=Ziqing |last2=Xu |first2=Shuhua |date=2019 |title=Population genomics of East Asian ethnic groups |journal=[[Hereditas]] |location=Berlin |publisher=[[BioMed Central]] |publication-date=2020 |volume=157 |issue=49 |page=49 |doi=10.1186/s41065-020-00162-w |pmc=7724877 |pmid=33292737 |doi-access=free}}</ref><ref>{{cite journal |last1=Shi |first1=Cheng-Min |last2=Liu |first2=Qi |last3=Zhao |first3=Shilei |last4=Chen |first4=Hua |title=Ancestry informative SNP panels for discriminating the major East Asian populations: Han Chinese, Japanese and Korean |journal=Annals of Human Genetics |date=September 2019 |volume=83 |issue=5 |pages=348–354 |doi=10.1111/ahg.12320 |pmid=31025319 |doi-access=free }}</ref><ref name="sciencemag2017">{{cite journal |last1=Siska |first1=Veronika |last2=Jones |first2=Eppie Ruth |last3=Jeon |first3=Sungwon |last4=Bhak |first4=Youngjune |last5=Kim |first5=Hak-Min |last6=Cho |first6=Yun Sung |last7=Kim |first7=Hyunho |last8=Lee |first8=Kyusang |last9=Veselovskaya |first9=Elizaveta |last10=Balueva |first10=Tatiana |last11=Gallego-Llorente |first11=Marcos |last12=Hofreiter |first12=Michael |last13=Bradley |first13=Daniel G. |last14=Eriksson |first14=Anders |last15=Pinhasi |first15=Ron |date=1 February 2017 |title=Genome-wide data from two early Neolithic East Asian individuals dating to 7700 years ago |journal=Science Advances |volume=3 |issue=2 |page=e1601877 |bibcode=2017SciA....3E1877S |doi=10.1126/sciadv.1601877 |pmc=5287702 |pmid=28164156 |last16=Bhak |first16=Jong |last17=Manica |first17=Andrea}}</ref><ref>{{cite journal |author1=Wang, Yuchen |author2=Lu Dongsheng |author3=Chung Yeun-Jun |author4=Xu Shuhua |year=2018 |title=Genetic structure, divergence and admixture of Han Chinese, Japanese and Korean populations |journal=Hereditas |volume=155 |page=19 |doi=10.1186/s41065-018-0057-5 |pmc=5889524 |pmid=29636655 |doi-access=free}}</ref><ref name="pmid 29636655" />{{Excessive citations inline|date=September 2024}} Although the genetic relationship is close, the various Han Chinese subgroups are genetically closer to each other than to their Korean and Yamato counterparts.<ref name="pmid 29636655" /> Other research suggests a significant overlap between Yamato Japanese and the Northern Han Chinese in particular.<ref>{{cite journal |last1=Cao |first1=Yanan |last2=Li |first2=Lin |last3=Xu |first3=Min |display-authors=etal |date=2020 |title=The ChinaMAP analytics of deep whole genome sequences in 10,588 individuals |journal=Cell Research |volume=30 |issue=9 |pages=717–731 |doi=10.1038/s41422-020-0322-9 |pmc=7609296 |pmid=32355288 |doi-access=free}}</ref>
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