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Testing Central and Inner Asian admixture among contemporary

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Forensic Science International: Genetics 15 (2015) 121–126

Contents lists available at ScienceDirect

Forensic Science International: Genetics

jou rnal homepage: www.elsevier.com/locate/fsig

Testing Central and Inner Asian admixture among contemporary Hungarians

a b b b,

Andra´s Bı´ro´ , Tibor Fehe´r , Guszta´v Ba´ra´ny , Horolma Pamjav *

a

Department of Anthropology, Hungarian Natural History Museum, Budapest H-1088, Hungary

b

Institute of Forensic Medicine, Network of Forensic Science Institutes, Ministry of Justice, Budapest, Hungary

A R T I C L E I N F O A B S T R A C T

Keywords: Historically, the Carpathian Basin was the final destination for many nomadic peoples who migrated

Hungarian speakers westward from Inner and towards Europe. Proto-Hungarians (Steppe Magyars) were among

Central and Inner Asian populations

those who came from the East, the Eurasian Steppe in the early middle ages. In order to detect the

Y-STRs

paternal genetic contribution from nomadic Steppe tribes, we tested 966 samples from Central Asian

Y-SNPs

(Uzbekistan, ), Inner Asian (Mongolians and Buryats in Mongolia) and Hungarian-speaking

European (Hungarian, Sekler and Csango) populations. We constructed median-joining networks of

certain in Hungarian-speaking European, and Altaic-speaking Central and Inner Asian

populations. We estimated that the possible paternal genetic contribution from the above described

populations among contemporary Hungarian speaking populations ranged between 5% and 7.4%. It is

lowest among Hungarians from Hungary (5.1%), while higher among Hungarian-speaking groups in

Romania, notably Sekler (7.4%) and Csango (6.3%). However, these results represent only an upper limit.

Actual Central/Inner Asian admixture might be somewhat lower as some of the related lineages may

have come from a common third source. The main haplogroups responsible for the Central/Inner Asian

admixture among Hungarians are J2*-M172 (xM47, M67, M12), J2-L24, R1a-Z93; Q-M242 and E-M78.

Earlier studies showed very limited Uralic genetic influence among Hungarians, and based on the

present study, Altaic/Turkic genetic contribution is also not significant, although significantly higher

than the Uralic one. The conclusion of this study is that present-day Hungarian speakers are genetically

very similar to neighbouring populations, isolated Hungarian speaking groups having relatively higher

presence of Central and Inner Asian genetic elements. At the same time, the reliable historical and

genetic conclusions require an extension of the study to a significantly larger database with deep

resolution, including ancient DNA data.

ß 2014 Elsevier Ireland Ltd. All rights reserved.

1. Introduction belt mountings, and ornaments on clothing also showed similari-

ties to those of Central and Inner Asia [4–7]. Hungarian archae-

For centuries, great efforts were made by Hungarian historians ologists and ethnographers showed that there are similarities in

to study the earliest period of their national history. While the the traditions of the ancient Hungarians and various Central and

academic mainstream was clearly in favour of the Hungarian Inner Asian cultures [8–10]. These were in the areas of burial,

language belonging to the Uralic family, many other researchers belief, and figurative arts. Therefore, an origin of the Hungarian

favour the theory of a closer relationship with the Turkic language language and early culture in a region ranging from Asia to

family and Turkic peoples. Anthropological analysis of bones is suggested, but a specific origin has been difficult to identify.

originating in the 10th century showed characteristics of Central were thought to be close genetic relatives of Hungarians.

Asian origin [1–3]. Archaeological remains of weapons, haversacks, However, based on studies done with mtDNA, Y chromosome STRs

and SNPs, they seem to have little genetically in common with

Hungarians [11,12]. And this is despite the fact that they also speak

a non-Indo-European Finno-Ugric language. A genetic relationship

* Corresponding author at: Institute of Forensic Medicine, Network of Forensic

was proven between two Hungarian ethnic groups, the Csangos

Science Institutes, Ministry of Justice, PO 216, 1536 Budapest, Hungary.

and Seklers. Both groups showed genetic affiliations with certain

Tel.: +36 1 457 01 83; fax: +36 1 457 0182.

E-mail address: [email protected] (H. Pamjav). Central Asian and European populations. These findings could have

http://dx.doi.org/10.1016/j.fsigen.2014.11.007

1872-4973/ß 2014 Elsevier Ireland Ltd. All rights reserved.

122 A. Bı´ro´ et al. / Forensic Science International: Genetics 15 (2015) 121–126

supported theories about a partially Asian origin of Hungarian manufacturer’s instructions. Fragment sizes and allele designa-

population [11]. However, most of the Central Asian-Hungarian tions were determined with a 3130 Genetic Analyzer (Life

Y-chromosomal relationship was based on the high frequency of Technologies, Foster City, CA) using GeneMapper IDX 1.2.1. soft-

-M198 among Kyrgyz and a small Hungarian ware.

sample, without knowing the deep structure of this haplogroup. When testing Y-SNP markers, amplifications of 1–2 ng

Since then, first Pamjav et al. [13], and then in a more genomic DNA were performed in an ABI 7500 Real-time PCR

comprehensive analysis by Underhill et al. [14] it was shown that instrument with Taqman Assay (Life Technologies, Foster City,

there is a clear SNP-based distinction between Eastern European CA) using the programmes designed by the manufacturer. The

(Z282, Z280, M458) and Central Asian (Z93) R1a-M198 males. relative fluorescence of the PCR products were analysed on an ABI

It was also noted that Hungarians show very limited or no 7500 with its’ SDS software, as described in the manufacturer’s

presence of Haplogroup N-M231 – including subclade N1c-Tat – manual (Life Technologies, Foster City, CA). Fifty-five Y-chromo-

which is frequent among other Uralic-speaking populations somal SNP markers were tested with Taqman Assays (Fig. 1). The

[15,16]. However, the potential genetic relationship with Turkic haplogroups tested and the markers used in the study originated

and Inner Asian peoples has been less researched, although this from YCC (Y-Chromosomal Consortium). The nomenclature of

relationship could shed light on the genetic basis of the alternative haplogroups followed the ISOGG 2014 Y-DNA haplogroup tree

Turkic (Turanian) theory. Different Turkic-speaking populations due to recent, new additions uncovered by YCC (Y-Chromosomal

have widely differing Y-chromosomal gene pools. They range from Consortium).

N1c-Tat dominated through C3-M47 dominated , A list of primers and Taqman probes for binary markers was

and Q-M25 dominated to genetically more diverse previously published [19], but we now updated the list with new

Uzbeks, Azeri and Anatolian Turks (Table S1). Therefore, we chose SNPs studied, as shown in Table S2. A new downstream SNP

not to focus only on haplogroup frequencies, but on analysing marker, L24, was tested for J2*-M172 (xM47, M67, M12) samples

haplotype structure. We have undertaken a survey of 966 samples to obtain more resolution within the haplogroup as suggested by

from Europe and Asia. This study is expected to provide insights van Oven et al. [20].

relevant to the Central and Inner Asian genetic contribution into Supplementary table related to this article can be found, in the

Hungarian speaking populations. It will also provide insight into online version, at doi:10.1016/j.fsigen.2014.11.007.

how the genetic variation is distributed in the contemporary

Hungarian, Central and Inner Asian population gene pool studied. 2.3. Data analysis

Supplementary table related to this article can be found, in the

online version, at doi:10.1016/j.fsigen.2014.11.007. To examine the STR variation within the haplogroups,

networks were constructed using the Network 4.6.1.2 programme

[21]. Repeats of the locus DYS389I were subtracted from the locus

2. Materials and methods

DYS389II and, as is common practice, the locus DYS385 was

excluded from the network. Within the network programme, the

2.1. DNA samples

rho statistic was used to estimate the time to the most recent

common ancestor (TMRCA) of haplotypes within the compared

To analyse the genetic relationship of present-day Hungarians

haplogroups.

with present-day Central and Inner Asians, we tested 522 samples

from Hungarian-speaking populations (332 Hungarians from

3. Results

Hungary, 95 Sekler from Romanian Transylvania, 95 Csango from

Romanian Moldova), 115 Uzbek samples from various parts of

Based on 10 Y-STR loci, networks were constructed within each

Uzbekistan (Ferghana Valley, Tashkent, Khwarezm, Samarqand,

of the haplogroups. These haplogroups overlapped among

Surkhodarya, Karakalpakstan), 8 samples from Kazakhstan’s

populations studied. All haplotype and haplogroup results can

Aqto¨be region, 127 Mongolian and 88 Buryat Mongolian samples

be found in Table S3. Haplogroup results are summarized in

from Mongolia. Archaic Sekler and Csango populations were

Table S4. For our analysis, we only considered those haplogroups

included to increase the matching potential, and we also collected

which occurred in more than one sample among both Hungarians

additional samples from tribes whose self-designation may have

and Central Asians (Uzbeks, Kazakhs, Madjars), or among

connection to the ethnonym Magyar, i.e. 61 Madjars from Uzbekistan

Hungarians and Inner Asians (Mongolians, Buryats). With this

and 45 Madjars from Kazakhstan. Out of the 966 samples, the

method, we identified nine haplogroups, which might indicate a

45 Kazakh Madjars [17], and 215 Hungarian samples [12] were

genetic relationship between contemporary Magyars and Altaic-

published before, but tested for further SNPs and samples in this

speaking populations. They are E-M78, G2a-P15, J2*(xM47, M67

study. The new samples published herein were sent to the YHRD and

and M12), N1c-L708, Q-M242, R1a-M458, R1a-Z280, R1a-Z93 and

the accession numbers are the following: Uzbekistan [Uzbek]

R1b*-P25(xM412). To verify or confute the relationships, we

YA003994, Uzbekistan [Madjar] YA003995, Mongolia [Buryat]

created median-joining networks on 10 loci for all ‘‘suspected’’

YA003996, Mongolia [Mongolian] YA003997 and Kazakhstan [Mad-

haplogroups. Results are discussed only in the context of the

jar, Aqto¨be] YA003998. The new populations, as well as the previously

potential matches between Hungarian and Altaic-speaking popu-

published populations were Hungarian [12], with accession number

lations, haplogroup by haplogroup. The haplogroups are described

YA003187, Csango [Romanian] YA002984 [18] and Sekler [Romanian]

as follows.

YA002983 [18]. These were used for comparison and can be

Supplementary tables related to this article can be found, in the

referenced at www.yhrd.org. The Y-SNP haplogroups for Sekler and

online version, at doi:10.1016/j.fsigen.2014.11.007.

Csango populations were tested by us and included in our data.

Each person gave their informed consent prior to their inclusion

3.1. Haplogroup E-M78

in the study.

2.2. Testing of Y-STR and Y-SNP markers The median joining network (MJ) of 31 E-M78 haplotypes is

shown in Fig. 2A. The network shows a star-like pattern. The

DNA was amplified with the PowerPlex Y (Promega, USA) biggest cluster (cluster 1 in Fig. 2A) was the modal haplotype

amplification kit including 12 Y-STR loci, according to the shared by three Hungarian speaking population groups, which

A. Bı´ro´ et al. / Forensic Science International: Genetics 15 (2015) 121–126 123

3.2. Haplogroup G2a-P15

MJ network of 38 G2a-P15 haplotypes is depicted in Fig. 2B. The

modal haplotype cluster (cluster 2 in Fig. 2B) is shared by four

populations including two Hungarians, two Seklers, one Csango

and one Mongolian male. One Csango is on a common branch with

three Uzbeks (on the top of the network). The three Uzbeks are

from Khwarezm subregion.

3.3. Haplogroup J2*-M172 and J2-L24

MJ network of 51 J2*-M172 haplotypes is seen in Fig. 2C.

There is no visible modal haplotype cluster and it yielded a non-

star like network. The lower part of the figure includes all Uzbek

Madjars, who are a homogenous population most likely affected

by a founder effect or genetic drift. The biggest cluster (cluster

2 in Fig. 2C) consists of 17 Uzbek Madjar males. There are some

other Uzbeks, one Mongolian and two Hungarians, which we

consider to be Central and Inner Asian. The upper part of the

figure is less clear, but we can see that one Hungarian and two

Seklers derive from the Kazakh Madjar in the centre (to the left),

and two Hungarians who come from Uzbek haplotypes in the

upper right part. So among J2*-M172 haplotypes, we consider

two Seklers and five Hungarians to be of Central Asian

admixture.

Twenty-four J2-L24 haplotypes resulted in a non-star like

network split into two parts, primarily based on DYS437 and

DYS391 loci (Fig. 2D). In the network on the upper right side, four

Hungarians derive from an Uzbek (Ferghana) haplotype, so we

considered them to be of a Central Asian admixture.

The other part of the network (on the left side) included

Hungarian speaking males, except for one Uzbek male and no

shared haplotype was seen.

3.4. Haplogroup N1c-Tat

An MJ network of 54 N1c-Tat haplotypes is shown in Fig. 2E. The

modal haplotype cluster (cluster 1 in Fig. 2E) is shared by five

population groups including 31 Buryat, two Mongolian, one Sekler,

one Uzbek and one Kazakh chromosome. One Sekler matches the

Buryat-Mongolian modal haplotype and thus can be considered

Inner Asian admixture in Hungary. Other Hungarian and Sekler

haplotypes are very far from Altaic N1c haplotypes and are

therefore more likely of Uralic or Baltic origin. It has been noted

that the most Buryat males share the same haplotype, which is due

to genetic drift.

3.5. Haplogroup Q-M242

The network of Q-M242 haplotypes shows a non-star like and

more diverse pattern, which makes it rather difficult to analyse

the relationship (Fig. 2F). However, due to the pre-eminence of

Q-M242 among Altaic Turkmens [22,23] and its’ general absence

in Europe and Finno-Ugric speaking populations, we assume

Central Asian admixture for all the five Hungarian speaking Q

individuals.

3.6. Haplogroup R1a-M458

Fig. 1. A phylogenetic tree of the tested 55 Y-SNP loci.

The network of 49 R1a-M458 haplotypes breaks down into two

easily identifiable star-like subclusters (Fig. 2G). The biggest

cluster (cluster 3 in Fig. 2G) includes three Hungarians, one Sekler

consisted of two Hungarian, one Sekler and three Csango males. and one Csango male. The second largest cluster consists of four

There is a potential that two Seklers and two Hungarians have a Hungarians (Fig. 2G, cluster 2). An interesting picture is noted in that

common origin with Uzbeks (one from Tashkent, one from one Uzbek from Khwarezm and one Madjar from Kazakhstan are in

Khwarezm) on the bottom of the network. the middle of the network connecting the two separate clusters as

124 A. Bı´ro´ et al. / Forensic Science International: Genetics 15 (2015) 121–126

Fig. 2. Median joining (MJ) networks of the Hungarian speaking, Central and Inner Asian populations compared. (A) MJ network of Y-STRs within E-M78 haplogroup for the

populations compared. (B) MJ network of Y-STRs within G2a-P15 haplogroup for the populations compared. (C) MJ network of Y-STRs within J2*-M172 haplogroup for the

populations compared. (D) MJ network of Y-STRs within J2-L24 haplogroup for the populations compared. (E) MJ network of Y-STRs within N1c-Tat haplogroup for the

populations compared. (F) MJ network of Y-STRs within Q-M242 haplogroup for the populations compared. (G) MJ network of Y-STRs within R1a-M458 haplogroup for the

populations compared. (H) Median-joining network of Y-STRs within R1a-Z93 haplogroup for the populations compared. (I) MJ network of Y-STRs within R1b-P25

haplogroup for the populations compared. The circle sizes are proportional to the haplotype frequencies. The smallest area is equivalent to one individual.

the median haplotype (Fig. 2G, cluster 1). One Csango descends from 3.9. -P25

this central haplotype and thus can be designated as Central Asian

admixture. While R1a-M458 is generally considered as an Eastern The network of 44 R1b-P25 (xM412) haplotypes clearly consists

European haplogroup, being especially frequent among Western of two clusters (Fig. 2I). On the left side, we find the star-like

Slavs and to a lesser extent, Eastern [24], based on our result network of M269 haplotypes, while on the right side, the typically

we cannot exclude the possibility that R1a-M458 originates from Central Asian subgroup M73 is visible (Note: M269 and M73

Central Asia. These Khwarezm and Madjar haplotypes may be the markers were not tested in this study, but a comparison with

remnants of the ancestral population. Attributing one Kazakh, one Myres et al. [25] STR-data suggest the connection). Among R1b-

Kazakh Madjar, and all three Uzbek R1a-M458 haplogroups to Slavic P25 haplotypes, no connection can be made, as Hungarian-

admixture seems unlikely, especially given the nearly complete lack speakers dominantly belong to the M269 branch, while Uzbeks

of other typically European haplogroups I-M170 and R1b-M412 belong to the M73 part. The limited number of Central Asians

among our Central and Inner Asian samples (Table S3). (M269) is situated on the edges of the network, thus representing

external admixture rather than source. Cluster 2 (Fig. 2I) in the part

3.7. Haplogroup R1a-Z280 of M269 consists of four Hungarians and one Sekler male.

The R1a-Z280 haplotypes produced a star-like network (figure

not shown), with all the Central Asians exactly matching Hungarian 4. Discussion

haplotypes (Table S4). Therefore, we assume that a genetic link was

from Finno-Ugric or Slavic peoples to Central Asians [13]. On examination of haplogroups with an N > 1 frequency among

both Hungarian-speaking European, and Altaic-speaking Central

3.8. Haplogroup R1a-Z93 and Inner Asian populations, we showed that the possible

maximum Central/Inner Asian admixture among contemporary

Thirty-six R1a-Z93 haplotypes produced a non-star like and Hungarian populations ranges around 5–7.4%. We took into

very diverse network (Fig. 2H). These included mostly Uzbek account only those haplotypes which could derive from Central/

haplotypes found in the central area, who were Hungarian- Inner Asian haplotypes according to the MJ-networks. The

speaking, Uzbek Madjar, Buryat and Mongolian populations were admixture was lowest among Hungarians from Hungary (5.1%),

branching off towards the edges. While some R1a-Z93 haplotypes while somewhat higher among Hungarian-speaking populations

might be a result of Roma admixture [13], for the purpose of this in Romania, notably Sekler (7.4%) and Csango (6.3%). The average of

study we assumed that all haplotypes of Hungarian-speaking these results was 5.7% among 522 Hungarian-speaking males (see

population groups to be Central/Inner Asian admixture. Table S4). The reason of the difference might be the long-time

A. Bı´ro´ et al. / Forensic Science International: Genetics 15 (2015) 121–126 125

isolation of Sekler and Csango groups, resulting in lower admixture The conclusion of this study is that present-day Hungarian

from neighbouring populations. However, we also must acknowl- speakers are genetically very similar to neighbouring populations,

edge that these numbers represent an upper limit and that actual isolated Hungarian speaking groups having relatively higher

Central and Inner Asian admixture might be somewhat lower. In presence of Central and Inner Asian genetic elements. However,

these admixture cases, the genetic links are not necessarily directly we could not show any significant genetic correlation between

from Altaic populations to Hungarians, as both populations may Hungarian-speaking and Central/Inner Asian samples which

have received these genetic markers from a common third would explain the linguistic difference among Hungarians and

unidentified source (e.g. , , and East Slavs). neighbouring populations. At the same time, the reliable historical

Because of this possibility, further research is needed. We also have and genetic conclusions require an extension of the study to a

to note that Central Asian admixture among Hungarians does not significantly larger database with deep haplogroup resolution,

necessarily come from Altaic-speakers. It may also come from including ancient DNA data.

ancient Iranian tribes who were later Turkicized by Altaic

conquerors. The main haplogroups responsible for the Central/ Conflict of interest

Inner Asian admixture among Hungarians are J2-M172 (xM47,

M67, L24, M12), J2-L24, R1a-Z93, Q-M242 and E-M78. The authors declare no conflict of interest.

Earlier studies reported that Haplogroup E, J and their main

subgroups spread from the Middle East with the Neolithic Acknowledgements

agricultural revolution [26–28]. It spread towards both Europe

and Central Asia, thus some of the common haplotypes such as E-

We would like to say special thanks to Dr. Eva Susa (General

M78, J2-M172* and J2-L24 may indicate a common Middle Eastern

Director of the Network of Forensic Science Institutes) for her

origin for both Hungarian-speaking and Central/Inner Asian

financial support. We thank sample donors and Betty-Jean Sigethy

samples. This is in contrast to the idea of a male migration from

and Rayn Hoyt for the English editing. We say special thanks to two

Asia towards the Carpathian Basin.

unknown reviewers for their constructive comments and sugges-

Based on the results of the Central and Inner Asian samples tions.

analysed in this study, we could not find a strong genetic

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