건강과 대안 » 부계혈통 http://www.chsc.or.kr 연구공동체 Mon, 13 Apr 2026 01:34:28 +0000 ko-KR hourly 1 http://wordpress.org/?v=3.5.2 [진화] 유럽인 부계 혈통 대부분 신석기 시대 아시아(근동) 기원 http://www.chsc.or.kr/?post_type=reference&p=1701 http://www.chsc.or.kr/?post_type=reference&p=1701#comments Thu, 21 Jan 2010 19:29:34 +0000 건강과대안 http://www.chsc.or.kr/?post_type=reference&p=1701

유럽인 부계 혈통 대부분 신석기 시대 기원

그동안 현대 유럽 인구 중 구석기 시대 사냥꾼(수렵인)의 후손과 신석기 시대 근동(Near East, 현재의 메소포타미아 지역)에서 이주한 농부들의 상대적 기여도에 대한 격렬한 논쟁이 벌어졌습니다.

같은 미토콘드리아 유전자형을 가진 그룹을 Haplogroup이라고 하는데,
 R1b1b2 (R-M269)는 유럽 남성들에게 가장 일반적인 Y성염색체에 있는 유전자입니다. 이 유전자는 서쪽(유럽)에서 동쪽(근동, 중동지역)으로 갈수록  그 빈도가 높아집니다.

선행 연구에서는 현대 유럽 남성이 구석기 시대 사냥꾼(수렵인)의 후손일 것으로 추정했지만, 이번 레스터대학의 연구팀은 유전자 연구를 통해서 현대 유럽 남성이 근동지방(초승달 유역으로 불리는 메소포타미아 지방)에서 아나톨리아(현재의 터키)를 거쳐 유럽으로 퍼져나갔다는 점을 보여주고 있습니다.

논문 원문과 프레시안의 기사를 아래에 첨부합니다.
=============================


A Predominantly Neolithic Origin for European Paternal Lineages


Patricia Balaresque1, Georgina R. Bowden1, Susan M. Adams1, Ho-Yee Leung1, Turi E. King1, Zoë H. Rosser1, Jane Goodwin2, Jean-Paul Moisan3, Christelle Richard3, Ann Millward4, Andrew G. Demaine4, Guido Barbujani5, Carlo Previderè6, Ian J. Wilson7, Chris Tyler-Smith8, Mark A. Jobling1*


1 Department of Genetics, University of Leicester, Leicester, United Kingdom, 2 Ty Celyn, Maeshafod, Blaina, Gwent, United Kingdom, 3 Laboratoire d’Etude du Polymorphisme de l’ADN, Faculté de Médecine, Nantes, France, 4 Molecular Medicine Research Group, Peninsula Medical School, Universities of Exeter and Plymouth, Plymouth, United Kingdom, 5 Dipartimento di Biologia ed Evoluzione, Università di Ferrara, Ferrara, Italy, 6 Dipartimento di Medicina Legale e Sanità Pubblica, Università di Pavia, Pavia, Italy, 7 Institute of Human Genetics, Newcastle University, Newcastle upon Tyne, United Kingdom, 8 The Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton, United Kingdom



Abstract


The relative contributions to modern European populations of Paleolithic hunter-gatherers and Neolithic farmers from the Near East have been intensely debated. Haplogroup R1b1b2 (R-M269) is the commonest European Y-chromosomal lineage, increasing in frequency from east to west, and carried by 110 million European men. Previous studies suggested a Paleolithic origin, but here we show that the geographical distribution of its microsatellite diversity is best explained by spread from a single source in the Near East via Anatolia during the Neolithic. Taken with evidence on the origins of other haplogroups, this indicates that most European Y chromosomes originate in the Neolithic expansion. This reinterpretation makes Europe a prime example of how technological and cultural change is linked with the expansion of a Y-chromosomal lineage, and the contrast of this pattern with that shown by maternally inherited mitochondrial DNA suggests a unique role for males in the transition.



Author Summary


Arguably the most important cultural transition in the history of modern humans was the development of farming, since it heralded the population growth that culminated in our current massive population size. The genetic diversity of modern populations retains the traces of such past events, and can therefore be studied to illuminate the demographic processes involved in past events. Much debate has focused on the origins of agriculture in Europe some 10,000 years ago, and in particular whether its westerly spread from the Near East was driven by farmers themselves migrating, or by the transmission of ideas and technologies to indigenous hunter-gatherers. This study examines the diversity of the paternally inherited Y chromosome, focusing on the commonest lineage in Europe. The distribution of this lineage, the diversity within it, and estimates of its age all suggest that it spread with farming from the Near East. Taken with evidence on the origins of other lineages, this indicates that most European Y chromosomes descend from Near Eastern farmers. In contrast, most maternal lineages descend from hunter-gatherers, suggesting a reproductive advantage for farming males over indigenous hunter-gatherer males during the cultural transition from hunting-gathering to farming.



Citation: Balaresque P, Bowden GR, Adams SM, Leung H-Y, King TE, et al. (2010) A Predominantly Neolithic Origin for European Paternal Lineages. PLoS Biol 8(1): e1000285. doi:10.1371/journal.pbio.1000285


Academic Editor: David Penny, Massey University, New Zealand



Received: May 8, 2009; Accepted: December 10, 2009; Published: January 19, 2010


Copyright: © 2010 Balaresque et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.


Funding: MAJ was supported by a Wellcome Trust Senior Fellowship in Basic Biomedical Science (grant number 057559); PB, GRB, SMA, ZHR, and CTS were supported by the Wellcome Trust (www.wellcome.ac.uk). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.


Competing interests: The authors have declared that no competing interests exist.


Abbreviations: hg, haplogroup; KYA, thousand years ago; mtDNA, mitochondrial DNA; TMRCA, time to the most recent common ancestor


* E-mail: maj4@le.ac.uk



Introduction


Events underlying the distribution of genetic diversity among modern European populations have been the subject of intense debate since the first genetic data became available [1]. Anatomically modern humans, originating in East Africa, colonized Europe from the Near East ~40 thousand years ago (KYA), then during the last glacial maximum populations retreated into the peninsulas of Iberia, Italy, and the Balkans, followed by northward recolonization from these refugia ~14 KYA. The most important cultural transition was the adoption of agriculture originating in the Fertile Crescent in the Near East at the start of the Neolithic, ~10 KYA [2]. It spread rapidly westwards via Anatolia [3] (Figure 1A), reaching Ireland by 6 KYA, accompanied by the development of sedentary populations and demographic expansion. Debate has focused on whether this spread was due to the movement and expansion of Near-Eastern farmers (demic diffusion), or to the transmission of cultural innovation to existing populations (acculturation), who then themselves expanded.


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Figure 1. Maps showing dates of the spread of early farming in Europe, and the frequency and microsatellite variance of haplogroup R1b1b2.


(A) Isochron map representing dates of early Neolithic sites in Europe, based on data of Pinhasi et al. (2005) [3]. KYBP, thousand years before present. (B) Geographical distribution of haplogroup frequency of hgR1b1b2, shown as an interpolated spatial frequency surface. Filled circles indicate populations for which microsatellite data and TMRCA estimates are available. Unfilled circles indicate populations included to illustrate R1b1b2 frequency only. Population codes are defined in Table 1. (C) Geographical distribution of mean microsatellite variance within hgR1b1b2, shown as an interpolated spatial frequency surface. Samples shown are those used for the calculation of variance only.

doi:10.1371/journal.pbio.1000285.g001

The observation of southeast–northwest frequency clines for “classical” genetic markers [1],[4], autosomal DNA markers [5],[6], and Y-chromosomal markers [7],[8] (though not for mitochondrial DNA [mtDNA] [9]) has been used to support the demic diffusion model. No dates can be automatically attached to these clines, however, and some [1], detected by principal component analysis, may simply reflect isolation by distance [10]. The direction of movement underlying a cline can also be ambiguous: the high-frequency pole could indicate the area of preexisting substrate least affected by a migration originating far away, or the final destination of a wave of migration into thinly populated territory, where expansion and drift have had their greatest effects [11].


The origins of a frequency cline of a lineage can be illuminated by analysing the diversity within it. For Y-chromosomal lineages defined by binary markers (haplogroups), this can be done using multiple microsatellites. This approach has been applied to haplogroups E, J [12], and I [13] within Europe, but the major western European lineage has not yet been focused upon. The frequency of the major western European lineage, haplogroup (hg) R1b1b2, follows a cline from 12% in Eastern Turkey to 85% in Ireland (Figure 1B), and is currently carried by some 110 million European men. Previous studies of lineages approximately equivalent to hgR1b1b2 [7],[8] suggested that it has a Paleolithic origin, based simply on its high frequency in the west. Here, in contrast, we show that the geographical distribution of diversity within the haplogroup is best explained by its spread from a single source from the Near East via Anatolia during the Neolithic. Taken together with the evidence on the origins of many other European haplogroups, this indicates that the great majority of the Y chromosomes of Europeans have their origins in the Neolithic expansion.



Results


To investigate the origins of hgR1b1b2, we assembled a dataset of 840 chromosomes from this haplogroup with associated nine-locus microsatellite haplotypes (Table 1; Table S1). The diversity of the lineage within each population (measured by mean microsatellite variance) should reflect its age: under a hypothesis of recolonization from southern refugia, we expect a gradient of diversity correlating with latitude, whereas Neolithic expansion from Anatolia predicts a correlation primarily with longitude. Figure 1C shows the geographical distribution of mean microsatellite variance, and Figure 2 shows that although there is no evidence for correlation with latitude (R2 = 0.06; p = 0.268), the correlation with longitude is significant (R2 = 0.358; p = 0.004), with greatest diversity in the east (strongly influenced by highly diverse samples within Turkey), thus providing support for the Neolithic colonization hypothesis.


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Figure 2. Relationship of diversity among 840 R1b1b2 chromosomes with (A) longitude and (B) latitude.


Population codes are defined in Table 1.

doi:10.1371/journal.pbio.1000285.g002

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Table 1. Frequency of haplogroup R1b1b2 in European populations, with geographical coordinates for sampled populations.

doi:10.1371/journal.pbio.1000285.t001

The two hypotheses also make different predictions for the number of sources of diversity within hgR1b1b2: under the postglacial recolonization model, we expect multiple sources, whereas under the Neolithic expansion model, we expect only one. We can test this by examining the phylogenetic relationships among microsatellite haplotypes. A reduced median network of 859 haplotypes (Figure 3) shows a simple star-like structure indicative of expansion from one source: 74 haplotypes (8.6%) lie in its central node, and this node plus its single-step mutational neighbours together comprise 214 haplotypes (24.9%). Haplotypes belonging to populations from all three refugia are present in the core of the network. This pattern seems incompatible with recolonization from differentiated refugial populations, and in terms of the history of hgR1b1b2, the refugia possess no special status. The core of the network also contains haplotypes from Turkey (Anatolia), which is compatible with a subpopulation from this region acting as a source for the westwards-expanding lineage.


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Figure 3. Reduced median network of microsatellite haplotypes within haplogroup R1b1b2.


Molecular relationships between the nine-locus microsatellite haplotypes of 849 hgR1b1b2 chromosomes, including seven Serbian and two Greek haplotypes not included in the other analyses because population sample sizes were too small. Circles represent haplotypes, with area proportional to frequency and coloured according to population. Lines between circles represent microsatellite mutational steps.

doi:10.1371/journal.pbio.1000285.g003

Does the time to the most recent common ancestor (TMRCA) of the hgR1b1b2 chromosomes support a Paleolithic origin? Mean estimates for individual populations vary (Table 2), but the oldest value is in Central Turkey (7,989 y [95% confidence interval (CI): 5,661–11,014]), and the youngest in Cornwall (5,460 y [3,764–7,777]). The mean estimate for the entire dataset is 6,512 y (95% CI: 4,577–9,063 years), with a growth rate of 1.95% (1.02%–3.30%). Thus, we see clear evidence of rapid expansion, which cannot have begun before the Neolithic period.


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Table 2. Estimates of TMRCA for individual populations, arranged from west to east.

doi:10.1371/journal.pbio.1000285.t002

The similarity between the isochron map of Neolithic sites (Figure 1A; [3]) and those of hgR1b1b2 frequency (Figure 1B) and diversity (Figure 1C) is striking. Further support for the association of the expansion of hgR1b1b2 with that of farming comes from a statistical comparison of the variables. The frequency of hgR1b1b2 at different points in Europe is significantly negatively correlated (R2 = 0.390; p = 0.0005) with the dates of local Neolithic sites (Figure 4A). For the local variance of the microsatellite haplotypes within hgR1b1b2, the correlation with Neolithic dates is significantly positive (R2 = 0.331; p = 0.0124; Figure 4B).


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Figure 4. Correlation of dates of Neolithic sites with hgR1b1b2 (A) frequency and (B) variance.


Population codes are defined in Table 1. YBP, years before present.

doi:10.1371/journal.pbio.1000285.g004


Discussion


Previous observations of the east–west clinal distribution of the common Western European hgR1b1b2 (or its equivalent) [7],[8] considered it to be part of a Paleolithic substrate into which farmers from the Near East had diffused. Later analyses have also considered variance, and have conformed to the Paleolithic explanation [14],[15]. Here, we concur that the cline results from demic diffusion, but our evidence supports a different interpretation: that R1b1b2 was carried as a rapidly expanding lineage from the Near East via Anatolia to the western fringe of Europe during the Neolithic. Such mutations arising at the front of a wave of expansion have a high probability of surviving and being propagated, and can reach high frequencies far from their source [11]. Successive founder effects at the edge of the expansion wave can lead to a reduction in microsatellite diversity, even as the lineage increases in frequency.


The innovations in the Near East also spread along the southern shore of the Mediterranean, reflected in the expansion of hgE1b1b1b (E-M81) [16], which increases in frequency and reduces in diversity from east to west. In sub-Saharan Africa, hgE1b1a (E-M2) underwent a massive expansion associated with the Bantu expansion [17],[18]. In India, the spread of agriculture has been associated with the introduction of several Y lineages [19], and in Japan, lineages within hgO spread with the Yayoi migration [20], which brought wet rice agriculture to the archipelago. On a more recent timescale, the expansion of the Han culture in China has been linked to demic diffusion [21]. In this context, the apparently low contribution of incoming Y chromosomes to the European Neolithic, despite its antiquity and impact, has appeared anomalous. Our interpretation of the history of hgR1b1b2 now makes Europe a prime example of how expansion of a Y-chromosomal lineage tends to accompany technological and cultural change.


Other lineages also show evidence of European Neolithic expansion, hgE1b1b1 (E-M35) and hgJ, in particular [12]. Indeed, hgI is the only major lineage for which a Paleolithic origin is generally accepted, but it comprises only 18% of European Y chromosomes [13]. The Basques contain only 8%–20% of this lineage, but 75%–87% hgR1b1b2 (Table S1); our findings therefore challenge their traditional “Mesolithic relict” status, and in particular, their use as a proxy for a Paleolithic parental population in admixture modelling of European Y-chromosomal prehistory [22].


Is the predominance of Neolithic-expansion lineages among Y chromosomes reflected in other parts of the genome? Mitochondrial DNA diversity certainly presents a different picture: no east–west cline is discernible, most lineages have a Paleolithic TMRCA [23], and hgH [24] and hgV [25] show signatures of postglacial expansion from the Iberian peninsula. Demic diffusion involves both females and males, but the disparity between mtDNA and Y-chromosomal patterns could arise from an increased and transmitted reproductive success for male farmers compared to indigenous hunter-gatherers, without a corresponding difference between females from the two groups. This would lead to the expansion of incoming Y lineages—as suggested by the high growth rate observed for hgR1b1b2. Similar conclusions have been reached for the Bantu expansion (in which the current Bantu-speaking populations carry many mtDNA lineages originating from hunter-gatherers [26]), the introduction of agriculture to India [19] and the Han expansion [21].


Some studies have found evidence of east–west clines for autosomal loci [6],[27]. By contrast, recent genome-wide SNP typing surveys [28]–[30] find a basic south–north division or gradient, including greater diversity in the south, but they provide no indication of the time-depth of the underlying events, which could in principle involve contributions from the original colonization, postglacial Paleolithic recolonization, Neolithic expansion, and later contact between Africa and southern Europe [31].


The distinction between the geographical patterns of variation of the Y chromosome and those of mtDNA suggest sex-specific factors in patterning European diversity, but the rest of the genome has yet to reveal definitive information. Detailed studies of X-chromosomal and autosomal haplotypes promise to further illuminate the roles of males and females in prehistory.



Materials and Methods 


Ethics Statement


Males were recruited with informed consent, following ethical approval by the Leicestershire Research Ethics Committee and the ethics committees of the Universities of Ferrara, Pavia, and Exeter and Plymouth.


DNA Samples and Haplotyping


A total of 2,574 DNA samples from European males, assigned to populations based on two generations of residence, were typed for the SNP M269 [17], defining hgR1b1b2. Following PCR amplification using the primers 5′-CTAAAGATCAGAGTATCTCCCTTTG-3′ and 5′-ATTTCTAGGAGTTCACTGTATTAC-3′, the T to C transition was analysed by digestion with BstNI, which cleaves M269-C-allele chromosomes only. Samples from the Iberian peninsula were typed using the SNaPshot (ABI) procedure [31]. Haplotype data were obtained for up to 20 Y-specific microsatellites [32],[33]. Data from the Ysearch database (http://www.ysearch.org) for Germany (GE) and Ireland (IR) were added, together with published data for Turkey, subdivided into East, West, and Central subpopulations based on published sampling information [14]. To avoid a bias from very large samples of hgR1b1b2 (GE and IR), these were randomly subsampled to give sample sizes of 75. This allowed a comparison of nine-locus haplotypes (DYS19, DYS388, DYS389I, DYS389II, DYS390, DYS391, DYS392, DYS393, and DYS439) for 849 hgR1b1b2 chromosomes, subdivided into 23 populations. Greek and Serbian samples were too small for population-based analyses, but were included in Network analysis.


Analysis


Neolithic dates, frequencies of hgR1b1b2, and local microsatellite variances were displayed using Surfer 8.02 (Golden Software) by the gridding method. Latitudes and longitudes were based on sampling centres.


Intrahaplogroup diversity was assessed for populations with hgR1b1b2 sample size ≥15 as the mean of the individual microsatellite variances [34], as has been done elsewhere (e.g., [35]); this measure is highly correlated (R2 = 0.871; p = 6.72×10−10) with a more conventional measure, average squared distance (ASD) [36]. Regression analyses were carried out in the R statistical package [37] to compare these two measures, and also to compare mean of variance with latitude and longitude.


A reduced median network [38] of microsatellite haplotypes was constructed using Network 4.5 and Network Publisher, using weighting based on the inverse of the microsatellite variances.


TMRCA and population growth rates were estimated using BATWING [39], under a model of exponential population growth and splitting. Whereas standard use of BATWING assumes a random sample from a population, we validated its use to analyse single haplogroups. Justification of this, together with other details, is given in Text S1.


To assess the correlation between the dates of Neolithic sites and the local hgR1b1b2 frequency and variance, we considered 765 sites and their associated calibrated radiocarbon dates [3]. We identified sites lying within a buffer-zone of 150-km radius around each location for which we had frequency or variance data (Figure 1B and 1C). When more than one site was identified in a given buffer-zone, we considered the mean of the dates. Regression analyses were carried out as described above.



Supporting Information


Table S1.


Haplotype data. Population abbreviations are as in Table 1; for each microsatellite (DYS19–DYS439), repeat unit numbers are given.


(0.14 MB PDF)


Text S1.


Details of application of BATWING.


(0.14 MB DOC)



Acknowledgments


We thank all DNA donors, Agnar Helgason for discussions, Nicolas Poulet for help with spatial analysis, and Jean-Michel Dugoujon and Evelyne Guitard for assistance with sample collection.



Author Contributions


The author(s) have made the following declarations about their contributions: Conceived and designed the experiments: CTS MAJ. Performed the experiments: PB GB SA HYL ZHR. Analyzed the data: PB IJW MAJ. Contributed reagents/materials/analysis tools: TEK ZHR JG JPM AM AGD GB CP. Wrote the paper: PB CTS MAJ.


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  “유럽남자 80%, 중동 출신 농부 자손”

레스터대 연구팀 “유럽 여자는 원주민이 조상”


출처 : 프레시안 기사입력 2010-01-21 오후 2:17:04


     오늘날 유럽에 살고 있는 남자들의 80% 이상, 그리고 많은 미국인들이 토착민이 아닌 중동인을 조상으로 하는 Y염색체를 갖고 있다는 충격적인 연구 결과가 발표됐다. 이 연구는 영국의 레스터대학 연구팀이 수행해 과학저널 <공공과학도서관-생물학(PLoS-Biology)>에 논문으로 실렸다.


20일 영국의 방송과 미국의 등 영미권 언론들이 요약한 보도 내용에 따르면, Y염색체는 남성에 고유한 ‘유전자군’으로 아들에게 거의 손상되지 않은 채 대대로 이어지는 특성을 이용해 유럽 남자들의 조상을 추적했다.


1만년 전 중동 농부들로 ‘씨앗 물갈이’


오늘날 1억 1000만 명의 유럽 남자들과 많은 미국인들이 갖고 있는 특정 Y염색체를 보유한 2574명의 유럽 남자들을 대상으로 조사한 결과, 이 염색체는 오늘날 이라크와 시리아에 걸친 이른바 ‘비옥한 초승달’ 지역에서 신석기 초기인 1만년 전 유럽으로 이동하기 시작한 중동인에서 유래됐다는 것이다.


이 중동인들이 유럽의 남동쪽에서 북서쪽으로 계속 확산돼 아일랜드에 도달한 것은 6000년 전이다. 현재 아일랜드 남자는 거의 100%가 이들로부터 유래된 Y염색체를 가졌으며, 웨일스 지방과 스페인도 90% 이상이다.


유럽 여자들에 대한 조사도 이뤄졌다. 여자들의 경우는 모계로만 대대로 변질되지 않고 전달되는 미토콘트리아 DNA를 추적하면 조상을 알 수 있다. 이 조사에서는 남자들과 반대로 수렵.채집 수준에 머물러 있던 토착민들을 조상으로 하고 있다는 결과가 나왔다.


유럽 여자들, 수렵.채집 남자보다 중동 농부 선호


이 논문의 제1저자 패트리샤 밸러레스크 박사는 “유럽인의 Y 염색체 80% 이상은 중동 출신의 농부에게서 왔고, 반대로 대부분의 여자들은 수렵.채집인들로부터 유래됐다는 것을 보여준다”고 말했다.


한마디로 ‘평화적인 종자 물갈이’가 이뤄졌다는 것이다. 이에 대해 논문 저자들은 ‘당시에 농부들이 여자들에게 훨씬 매력적이었다”고 표현했다.수렵.채집에 머물던 시대에 농사기술은 당시 세계를 뒤바꿀 혁명적인 기술이었다는 점에서 여자들이 농부들을 선호했다는 것이다.


밸러레스크 박사는 “수렵.채집 시대에서 농경 시대로 전환하는 동안, 농사 짓는 남자가 수렵.채집하는 토착 남자에게 번식에 유리했다는 것을 보여준다”고 말했다.


이 연구로 그동안 농사기술만 유럽으로 전파된 것인지, 아니면 중동 출신의 농부들의 이주에 의해 농사기술이 유럽에 전파된 것인지 계속된 논란에 종지부를 찍을지 주목된다.
 


/이승선 기자 

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