The paper presents the results of studying a series of mitochondrial DNA samples obtained from bone materials of representatives of the eastern variant of the Pakhomovo culture from the Stary Sad burial ground in the Barabinsk forest-steppe. Their comparison with similar data on previous populations of the region allowed us to reconstruct the processes of formation of the genetic composition of the Pakhomovsky population, taking into account the archaeological context. The components of the gene pool, whose origin is associated with the autochthonous pre-Andronov population of the West Siberian forest-steppe and migrant Andronov (and post-Andronov) groups, are identified. The article presents a preliminary reconstruction of ethno-cultural interactions in the region during the migration of the Andronovo (Fedorovsky) population and the formation of Late Bronze Age ethno-cultural groups on its territory, based on archaeological and paleogenetic data.

Key words: archeology, paleogenetics, ancient DNA, mitochondrial DNA, Pakhom culture, ethnocultural interactions, Andronovo (Fedorovskaya) culture, migrations.

Introduction

Long-term studies of archaeological sites of the Bronze Age in the Ob-Irtysh forest-steppe have made it possible to develop a scheme of historical and cultural development [Molodin, 1983, 1985], which is being improved as new sources appear [Molodin, 2010]. Today, the picture of the existence and interaction of cultures in the late Bronze Age in this area looks mosaic, and in the transition from bronze to iron time-even more complex.

It is obvious that the dominant place here was occupied by the carriers of the Irmen culture, the most powerful education for its time. They lived in vast areas of the West Siberian forest-steppe practically from the Achinsk-Mariinskaya basin to the left bank of the Irtysh (Bobrov, 1992; Molodin, 1985). Carriers of the Irmen culture were an autochthonous population for the region, which had a great influence on its neighbors and actively interacted with migrants. The northwestern part of the Ob-Irtysh region was occupied by representatives of the Baraba variant of the Suzguneka culture (Molodin and Chemyakina, 1984), a population that came from the north - west and was influenced by the Irmens (Molodin and Zakh, 1985).

Back in the 1980s, one of the authors of this work drew attention to separate burial complexes on the territory of Baraba, which differ in inventory (primarily ceramics) and funeral rites from both Irma and Suzgun (Molodin, 1981). Further analysis revealed the anthropological specifics of the people who left the data

Fig. 1. Map-layout of archaeological sites (necropolises) with manifestations of the eastern version of the Pakhom culture.

1-Tartas-1; 2-Old Garden; 3-Preobrazhenka-3; 4-Grishkina Zaimka.

2. Burial plans (1 - 4) of the eastern variant of the Pakhomovo culture and bronze tools found in them (5-12).

1, 2, 4 - 6, 12 - Old Garden; 3, 7-11-Grishkina Zaimka.

complexes (Molodin and Chikisheva, 1988). Even at the time of discovery of these objects, their connection with the Begazy-Dandybayev culture monuments in Central Kazakhstan was obvious, which confirmed the assumption of M. F. Kosarev and V. I. Matyushchenko about the Begazy-Dandybayev substrate among the bearers of the Elov culture [Kosarev, 1981, p. 28; Matyushchenko, 1974, p. 163]. Later, these connections for the Kulunda districts more south of Baraba were traced in the work of V. S. Udodov [1994].

Particularly important materials were obtained during the excavation of the Old Garden burial ground in Baraba* (Fig. They suggested the existence of a special cultural formation in Western Baraba during the Late Bronze Age (Molodin and Neskorov, 1992). 2, 5), the studied monument was assigned to the final stage of the Bronze Age-the transition from bronze to iron time. Consequently, the Stary Sad burial ground represents the final stage of the Pakhomov culture (but not the initial stage, as some researchers believe [Kostomarov, 2010, p. 22]).

Already in the 1980s, it was obvious that we were dealing with a special cultural entity, although no special work was carried out to identify it. In 1987, O. N. Korochkova identified the Pakhomovo culture [1987], whose range was within the limits of the Tobolsk-Irtysh region [Korochkova, 2010]. To date, it is clear that the series of monuments in the Barabinsk forest-steppe, which was discussed above, represents the eastern periphery of the area of the Pakhomov culture (see Figure 1). We can definitely say that this culture is multicomponent. Apparently, it was formed on the basis of the Pakhomovtsy (western) proper, where the southern begazy-Dandybaevsky component is extremely noticeable, and the Andronovo (Fedorovsky) component is also noticeable in the materials from the Baraba monuments (Fig. 3, 2 - 4, 7, 10). In addition, the influence of the autochthonous Irmen culture is evident (fig. 3,5, 6, 8, 9, 11, 12), as well as the constant influence of the Suzgun culture carriers from the north.

* The research was started in the 1980s by V. I. Molodin and N. V. Polosmak, continued by A. V. Neskorov and completed in 2011 by V. I. Molodin and L. N. Mylnikova.

3. Vessels from burial grounds of the eastern variant of the Pakhomovo culture. 1 - 4, 6, 7, 10, 14 - Old Garden; 5, 8, 11-13-Grishkina Zaimka; 9-Preobrazhenka-3.

On the territory of Baraba there is a small part of the currently studied Pakhomovsky monuments (see Figure 1). However, they (primarily the Stary Sad necropolis) contain paleoanthropological materials that are probably the most representative of the Pakhomov population at present. In the western part of the area of the Pakhomovo culture, few funerary complexes have been discovered and studied so far [Ibid., pp. 68-70].

The cultural and anthropological specifics of the representatives of the eastern variant of the Pakhomov culture, the representativeness of paleoanthropological material make it extremely relevant to study the biological characteristics of the carriers of the Pakhomov culture in the Barabinsk forest-steppe, both by traditional methods of physical anthropology and paleogenetics. This paper presents the first results of the study of mtDNA variants of representatives of the Pakhomovskaya region.

cultures buried in the Old Garden burial ground in the Barabinsk forest-steppe, as well as their analysis from the point of view of the mechanisms of formation of the genetic composition of the inhabitants of the West Siberian forest-steppe of the final Bronze Age. In addition, the article presents ideas about some population-genetic processes that led to the formation of distinctive cultures of the region in this period.

Materials and methods

Paleoanthropological materials. A sample of bone tissue samples from the Pakhomovo culture carriers for the study was obtained from the paleoanthropological collection of the Institute of Electrotechnical Engineering SB RAS, formed from the materials of the excavations of the Stary Sad burial ground. Long limb bones (femur, tibia, or humerus) of 11 individuals with the best macroscopic preservation of their skeletons were selected for paleogenetic studies.

Pretreatment of paleoanthropological material and DNA extraction. The methods described in [Pilipenko et al., 2010; Pilipenko, Molodin, and Romashchenko, 2011] were used. The surface of the samples was treated with a 7% sodium hypochlorite solution to destroy possible contamination of modern DNA, then a layer ∼1 - 3 mm thick was mechanically removed and the sample was irradiated with ultraviolet light for at least 1 h on each side. Bone powder was drilled from the compact substance of long limb bones. To isolate DNA, bone powder was soaked in 5 M guanidinisothiocyanate buffer for 48 h [Pilipenko et al., 2010]. DNA extraction was performed by phenol-chloroform extraction followed by precipitation with isopropanol.

mtDNA sequence analysis. Amplification of GVSI mtDNA was performed by two different methods: amplification of four short overlapping fragments was performed by one-round PCR [Haak et al., 2005], and amplification of one long fragment was performed by nested PCR (including two rounds of reaction) [Pilipenko et al., 2010]. Amplification products of DHW I fragments were cloned in a bacterial vector using the pGEM-T Easy Vector System Kit (Promega, USA) and 6-15 clones were sequenced for each fragment.

The nucleotide sequence was determined using the ABI Prism BigDye Terminator Cycle Sequencing Ready Reaction Kit (Applied Biosystems, USA) in accordance with the manufacturer's recommendations. The products of the sequencing reaction were analyzed using an ABI Prism 3100 Genetic Analyzer (Applied Biosystems, USA) at the Center for Collective Use of Automatic DNA Sequencing SB RAS (Novosibirsk). The obtained sequences were compared with the refined Cambridge mtDNA reference sequence [Andrews et al., 1999]. Phylogenetic interpretation of the sequences was performed in accordance with the existing classification of mtDNA structural variants [van Oven and Kayser, 2009]. For phylogeographic analysis, we used our own database on the structure of GWS I mtDNA, which includes more than 20 thousand samples from modern populations of Eurasia, which was formed from literature sources.

Measures against contamination. All work with the ancient material was carried out in laboratory rooms designed for working with ancient DNA: they are equipped with an independent supply ventilation system with an air filtration function that creates a pressure gradient from cleaner to less clean rooms, as well as an ultraviolet radiation system, laminar flow cabinets of the second class of protection, etc. The laboratory staff used sets of workwear for clean rooms, and frequently changed sterile gloves. All work surfaces and appliances were regularly treated with a solution of sodium hypochlorite (5%) and irradiated with ultraviolet light. The complete extraction and amplification procedure was performed in parallel with ancient samples and system purity checks (without adding paleomaterial) to detect possible contamination of the reagents and equipment used. For all employees of the paleogenetic laboratory who have access to clean rooms, as well as for specialists in anthropology who were engaged in the analysis of paleoanthropological remains and the selection of materials for genetic research, the sequence of GWS I mtDNA was determined in order to identify possible contamination of materials.

Results and discussion

Reliable results for determining the GWS I mtDNA sequence were obtained for 8 out of 11 samples. The high rate of success in obtaining DNA samples suitable for analysis is due both to the relatively favorable climatic conditions of Central Baraba (and the location of the Old Garden burial ground) for the preservation of DNA in bone remains, and to the strict selection of samples included in the sample according to the degree of preservation of bone material.

All the mtDNA samples studied differ in the GWS I sequence (see table). The structure-

Structure and phylogenetic position of mtDNA samples from the eastern variant of the Pakhomovo culture from the Old Garden burial ground in the Barabinsk forest steppe

* The numbering of positions in mtDNA corresponds to the Cambridge mtDNA reference sequence [Andrews et al., 1999] without 16000 (i.e., position 223 in this table corresponds to position 16223 in the reference sequence). The letters after the position number indicate base replacements only in the case of transversion.

Haplotype analysis makes it possible to unambiguously determine whether mtDNA variants belong to five haplogroups: West Eurasian U2e, U5a, T, and East Eurasian C and A10. The mixed structure of the gene pool, which combines mtDNA groups that are typical of populations in the western and eastern parts of Eurasia, is characteristic of all the ancient populations of the West Siberian forest-steppe belt that we have studied so far [Pilipenko, 2010].

The availability of an extensive sample of mtDNA samples from representatives of the Baraba forest-steppe population and adjacent territories of various periods of the Bronze Age makes it possible to conduct a comparative analysis of them. Therefore, the obtained data on the mtDNA gene pool of the Pakhomovsky population, taking into account the archaeological context, can be used to elucidate the mechanisms of formation of the genetic composition of this group.

According to available data, two groups of variants are distinguished in the mtDNA gene pool of the population of the eastern periphery of the Pakhomovo culture area. The first group, represented by the lines of the West Eurasian haplogroups U2e, U5a and East Eurasian A10 and C, brings the population under consideration closer to the autochthonous Neolithic and Pre-Iron Age populations of the region-carriers of the Ust-Tartass, Odin, and Krotovo cultures. Thus, haplogroups U2e and U5a formed the basis of the Western Eurasian component of the mtDNA gene pool of the listed groups of the Baraba population of the Early Metal, Early Bronze, and Early Bronze Age periods, and haplogroup C is most represented in the Eastern Eurasian component of their gene pool [Ibid.].

It should be emphasized that the similarity of these components of the gene pool of the Pakhomovsky population represented in the anthropological materials of the Stary Sad burial ground to the early Baraba populations is manifested not only at the level of haplogroups or mtDNA subgroups. Most of the specific structural variants identified in the Pakhomov sample are also present in the gene pool of the early groups. This is a variant of the West Eurasian haplogroup U2e with haplotype 051-183AC-189-256, which is traced in all the studied groups of the Baraba population from the Neolithic to the beginning of the developed Bronze Age, as well as a variant of the haplogroup U5a with haplotype 256-270, found in the gene pool of the Ust-Tartass population of the Baraba of the Early Metal age ([Ibid.], unpublished data of authors). At the level of specific haplotypes, it is more difficult to interpret haplogroup C variants in the gene pool of Pakhomov culture carriers. Root haplotype 223 - 298 - 327 It is a typical component of the gene pool of early groups of the Baraba population, especially the Ust-Tartass population ([Ibid.], unpublished data of the authors). However, according to phylogeography, this variant is widely represented in Siberia at the present time, as it probably was in the Bronze Age. Therefore, it is difficult to use it as a marker of genetic succession between groups of different times on a local territory. Haplotype 223 - 298 - 325 - 327 - 362, According to our data, it appears in the gene pool of the Baraba population only during the migration of the Andronovo population against the background of a general increase in the diversity of haplogroup C lineages. 223 - 298 - 325 - 327 recorded in the gene pool

carriers of the Krotovo Baraba culture (burial ground Sopka-2 / 4B) [Ibid.]. Thus, this variant cannot yet be unambiguously considered as a genetic marker of association with the pre-Alexandron population of Baraba.

A particularly significant sign of mediated genetic continuity between representatives of the Pakhomov culture and the pre-Andronovo autochthonous population of the region is the presence in the sample of variants of haplogroup A10, which, apparently, was formed locally in the gene pools of the population of the forest-steppe zone of Western Siberia and underwent a long evolution in this territory*. Thus, according to most of the mtDNA lines studied, the Pakhomov population, whose remains were found in the Stary Sad burial ground, is close to the populations of the Central Baraba of the Pre-Iron Age of the Bronze Age, i.e. a significant part of it comes from the autochthonous pre-Iron Age population of Western Siberia.

It should be emphasized that based on the data obtained by us, it is premature to talk about the participation of specific groups of the pre-Andronovo population (for example, carriers of the Krotovo culture) in the formation of the gene pool of the Baraba Pakhomovites. We do not yet know the composition of mtDNA variants in the gene pools of representatives of other cultures of the Developed and Late Bronze Age of the forest-steppe zone of Western Siberia, except for the Krotovskaya one. It is possible that mtDNA variants that bring together the Pakhomov and Krotov (as well as Odinovek) groups of the Baraba population were also characteristic of other pre-Andron populations in Western Siberia. To clarify this issue, it is necessary to conduct a large-scale study of the gene pool of early population groups, as well as carriers of Late Bronze Age cultures (Irmen, Karasuk, etc.) in various territories of Western Siberia.

The second component in the studied series of samples is two variants of haplogroup T. Lines of haplogroup T, in our opinion, can be considered as genetic markers of migration of the Andronovo (Fedorovsky) population to Western Siberia. Haplogroup T variants are not represented in the sample of mtDNA samples from the Preandron population. At the same time, the lines of this haplogroup are characteristic of the Western Eurasian component of the mtDNA gene pool of the Andronovo population of the region (Molodin et al., 2011).

Thus, the series of mtDNA samples of eastern Pakhomovites is at least two-component and reflects the participation of two genetic substrates in the formation of the andronoid population of Western Siberia - the autochthonous pre-Andronovsky and the alien Andronovsky. It is possible that the Andronovo (migrant) component is to a certain extent a relict component that was dominant in the population of carriers of the Begaza-Dandybay culture, and genetically and culturally continues the line of development of the Andronovo (Fedorovsky) population of the central part of its range.

In general, our data correspond to the generally accepted ideas about the formation of andronoid cultures in Western Siberia (Kosarev, 1981; Korochkova, 2010, 2011). It is obvious that at the level of material and spiritual culture, this process was multi-stage (multi-stage), and its character and intensity in different parts of the forest-steppe belt of Western Siberia had specific features, which led to the emergence of so-called andronoid cultural groups that were diverse in their features.

Changes in the genetic composition of the population to some extent correlate with the transformations of material culture that occurred during large-scale ethno-cultural interaction. There are at least two stages in the formation of the gene pool structure of the Western Siberian andronoid population. The first stage is associated with the mass penetration of representatives of the Andronovo (Fedorovskaya) ethno-cultural community into the region and their interaction with autochthonous groups of the population. It seems to correspond to the monuments of the Andronovo culture proper, as well as the complexes left by the autochthonous (Late Krotovian for Baraba) There are also monuments that reflect the processes of mutual cultural influence of migrants and aborigines, i.e. they contain complexes with signs of cultural syncretism (Molodin, 2011). A striking example is the Tartas-1 burial ground, which reflects the processes of interaction between the alien Andronovo (Fedorovsky) and aboriginal Late Krotov populations in Central Baraba [Ibid.]. At the population-genetic level, during this period, genetic material was exchanged (in different territories, the degree of its intensity was different) between groups of migrants and aborigines, i.e. the gene pool of the local population of the region was enriched with new genetic components. At the same time, the genetic influence of the autochthonous population on migrants, at least in the female line, which reflects the mtDNA gene pool, may have been very significant, as can be traced from the materials of the Tartas-1 burial ground [Pilipenko, 2010; Molodin et al., 2010]. In material culture, the role of Andronovo traditions gradually increased [Molodin et al., 2010, 2011]. This stage is formed by-

* A special paper on this issue is being prepared for publication.

Most likely, the growth of the Andronoid population was saturated with several migration events, which caused the appearance in the region of various genetic elements from the Andronoid environment, which was not genetically homogeneous. The results of ethno-cultural interaction are recorded not only by archaeological materials, but also by physical anthropological data: in the Andronovo series of Western Siberia, components are identified whose origin is associated with the influence of the aboriginal population (Chikisheva and Pozdnyakov, 2003).

The second stage of the formation of late andronoid groups, in particular the Pakhomov population, is characterized by the interaction of mixed groups formed in the region at the first stage of genetic contacts between migrants and aborigines, with cultural formations of the Begazy-Dandyba type (with already modified but clearly recognizable Andronov traits) and with a population that is genetically close (or identical) to the autochthonous, which existed in the forest-steppe before the arrival of the Andronovo groups, which to some extent experienced the Andronovo influence. From a genetic point of view, this means preserving the "balance" of the alien Andronovo origin and autochthonous components in the population's gene pool, which is confirmed by the first results of a study of a series of mtDNA samples of carriers of the eastern variant of the Pakhomovo culture from the Stary Sad burial ground.

Of course, the lack of paleogenetic data on the carriers of Begaza-Dandybaevskaya, Pakhomovskaya proper, Irmenskaya, and other Late Bronze Age cultures in the region significantly complicates the interpretation of our results. However, an analysis of the archaeological materials of the monuments of the eastern area of the Pakhomovo culture (and a number of monuments of its metropolis (Korochkova, 2010)) revealed a Late Bronze Age component of the "Karasuk type" cultures (as defined by N. L. Chlenova (1981, p. 17)), which includes both its own and relict andronoid elements.

The data obtained suggest that the role of elements associated with autochthonous Preandronic groups has increased in the gene pool of Late Bronze Age culture carriers. In the studied series, among the two components identified above - the native and "Andronovsky" - mtDNA variants native to the West Siberian forest-steppe somewhat prevail. The predominance of autochthonous components in the female part of the gene pool of populations was previously traced using materials from the Tartas-1 monument, which reflect direct contacts between aboriginal and alien populations.

The formation of each culture of the Late Bronze Age in the region in reality, of course, was a more complex and multi-factorial process than in the model proposed by us. Each of the groups that interacted was characterized by different directions of genetic contacts not only with northern populations, but also with other neighboring cultural groups, including andronoid ones, representing the southern direction (Begazy-Dandybaevtsy), which is recorded according to archaeological data (Molodin and Neskorov, 1992). Reconstruction of these genetic interactions is complicated by the similarity of the structure of the gene pool of cultural groups that were influenced by the Andronovites or their descendants. It should be noted that the paleogenetic results obtained by us (both for carriers of the Pakhomov culture and for representatives of previous ethno-cultural groups) include only data on the composition of mtDNA lines, which is inherited according to the maternal type and, therefore, primarily reflects the processes of formation of the genetic composition of the female part of the populations under consideration. It should be borne in mind that these data are still fragmentary and characterize local population groups. To create a complete and objective picture of the processes that took place and to test experimentally the assumptions made in this paper, it is necessary to conduct large - scale studies involving paleoanthropological groups of all the main groups participating in the processes under consideration, as well as to study materials on other genetic markers, primarily Y-chromosome markers, reflecting the genetic history of the male part of populations. Only after obtaining sufficient genetic material in terms of the volume and composition of markers is it possible to conduct a more detailed comparison of the accumulated extensive archaeological data on the formation of Late Bronze Age cultures (including Pakhomovskaya) with the corresponding population-genetic ones. This article is one of the first examples of joint searches of geneticists and archaeologists in this direction.

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The article was submitted to the Editorial Board on 15.06.12.

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