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The Nanai Clan Samar: The structure of gene pool based on Y-chromosome markers

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Abstract

Members of the Nanai clan Samar reside in the Gorin area of the Khabarovsk Territory. Their gene pool was studied using the SNP markers of the Y-chromosome. The major haplogroup, occurring in more than 83% of clansmen, is the northern Eurasian haplogroup N1c1-M178. Four other haplogroups are C∗-M130, I∗-M170, J2a1a-M47, and O2-P31. The most frequent haplogroup, N1c1-M178, indicates mostly Tungus origin of the Samar clan; other haplogroups detected by complete sequencing, such as the minor haplogroup C3∗-M130, reveal ties with native populations of the Amur basin. Genetic distances and their multidimensional scaling demonstrate marked affinities of Samar clansmen with Yakuts, Khakas, and certain groups of Buryats, suggesting a common origin. Nanai of other regions are much further from the Samar. Copyright © 2015, Siberian Branch of Russian Academy of Sciences, Institute of Archaeology and Ethnography of the Siberian Branch of the Russian Academy of Sciences. Published by Elsevier B.V. All rights reserved.
ARCHAEOLOGY,
ETHNOLOGY
& ANTHROPOLOGY
OF EURASIA
Archaeology Ethnology & Anthropology of Eurasia 43/2 (2015) 146–152
E-mail: Eurasia@archaeology.nsc.ru
Copyright © 2015, Siberian Branch of Russian Academy of Sciences, Institute of Archaeology and Ethnography of the Siberian Branch of the Russian
Academy of Sciences. Published by Elsevier B.V. All rights reserved
doi:10.1016/j.aeae.2015.09.015
146
Introduction
The position of the Lower Amur region, on the periphery
of eastern Siberia and the Paci¿ c coast, has become a
*Supported by the Russian Foundation for Basic Research
(Projects No. 13-04-10102, 13-06-90705, 14-06-00384,
14-06-10026, and 14-14-00827); full sequence analysis of
Y-chromosome was supported by the Russian Science
Foundation (Project No. 14-14-00827).
Y.V. Bogunov1, O.V. Maltseva2,
A.A. Bogunova3, and E.V. Balanovskaya4
1Vavilov Institute of General Genetics, Russian Academy of Sciences,
Gubkina 3, Moscow, 119991, Russia
ȿ-mail: forbogunov@inbox.ru
2Institute of Archaeology and Ethnography, Siberian Branch, Russian Academy of Sciences,
Pr. Akademika Lavrentieva 17, Novosibirsk, 630090, Russia
ȿ-mail: olymals@gmail.com
3Amur State University of Humanities and Pedagogy,
Kirova 17, Komsomolsk-on-Amur, 681000, Russia
ȿ-mail: annasyachina@mail.ru
4Medical Genetic Research Center,
Moskvorechye 1, Moscow, 115478, Russia
ȿ-mail: balanovska@mail.ru
THE NANAI CLAN SAMAR:
THE STRUCTURE OF GENE POOL BASED
ON Y-CHROMOSOME MARKERS*
Members of the Nanai clan Samar reside in the Gorin area of the Khabarovsk Territory. Their gene pool was studied
using the SNP markers of the Y-chromosome. The major haplogroup, occurring in more than 83 % of clansmen, is
the northern Eurasian haplogroup N1c1-M178. Four other haplogroups are ɋ*-Ɇ130, I*-M170, J2a1ɚ-M47, and
O2-P31. The most frequent haplogroup, N1c1-Ɇ178, indicates mostly Tungus origin of the Samar clan; other
haplogroups detected by complete sequencing, such as the minor haplogroup C3*-M130, reveal ties with native
populations of the Amur basin. Genetic distances and their multidimensional scaling demonstrate marked af¿ nities
of Samar clansmen with Yakuts, Khakas, and certain groups of Buryats, suggesting a common origin. Nanai of other
regions are much further from the Samar.
Keywords: Nanais, tribal structure, Samar clan, gene pool, Y-chromosome, haplogroups.
decisive factor in the formation of the population gene
pool, including populations of both the taiga and the
coastal areas. At least since the Neolithic, this region
was the migration zone for people from Siberia and
East Asia—small groups of Tungus, Turks, Mongols,
and Manchus mixed with the previous population. The
study of the gene pool of Nanais, the most numerous
ethnic group in the Lower Amur region (approximately
ten thousand people), relying on the knowledge of tribal
structures and methods of ethnic genomics, allows us
ANTHROPOLOGY AND PALEOGENETICS
Y.V. Bogunov et al. / Archaeology, Ethnology and Anthropology of Eurasia 43/2 (2015) 146–152 147
not only to clarify ethnogenesis, but also to identify the
different layers in the gene pool of the populations of
Amur, and to reconstruct historical processes on the
territory of the Far East.
In the works of many researchers of late 19–
20th centuries, the Nanai community appears as a
conglomerate of disparate groups with an appropriating
type of economy, and without a clear social structure and
clear boundaries separating it from other communities
of the Amur River valley (Schrenk, 1883; Lopatin,
1922; Smolyak, 1975).
Until the 19th century, a period of consolidation
of local tribes and clans into the Nanai ethnic
group, all populations of Amur were divided into
several large groups, according to the area of their
residence and cultural and linguistic influence:
Manchu (Natki, Duchers), Mongolian (Solons, Daurs),
and Tungus (Manegirs, Birars, Kile) (Schrenk, 1883:
11–93; Dolgikh, 1960). In the 19th century, the
abovementioned tribal formations disappeared from
the ethnographic map of the Amur region, initiating
the Nanai clan organization. In 20th century, the clan-
names became surnames while maintaining previous
territorial binding. However, V.A. Tugolukov, on
examining the overall picture of ethnic relations in the
Amur region during the last three centuries, concluded
that the term “clan”, in relation to the local community,
is not entirely correct. In the background of the Tungus
expansion in the region in the 17th century, which led
to the assimilation of the indigenous Amur population
and variations of the clan names, the appropriate term
is “family” (Tugolukov, 1972: 105). However, most
scientists, when considering the social organization
of ethnic communities in the Lower Amur region,
including Nanai, still use the concept of “clan” that
we ¿ nd in earlier references.
The infusion into the foreign cultural environment
was performed through the institution of doha,
whereby small clans formed alliances between
themselves in order to survive. Relationships in a
formed patronymic community were based on labor
and legal partnership between its members, and
involved joint action in religious ceremonies (Karger,
1929: 4–8; Smolyak, 1970: 266, 288–289; 2001:
14). While studying the Udege and the Orochi clans,
Tugolukov assumed that doha in the Lower Amur
region served as connection between the ¿ lial clans
which had a common ancestor. Its origin is associated
with the Tungus expansion, which reshaped the
local matrilineal kinship system that can be traced
from Ainu and Nivkh. So, the maternal (Nivkh) clan
entered into a relationship with the paternal clan
(Tungus). Children of such marriages, according to
the Nivkh rules, cannot marry within a father’s clan,
and, according to the Tungus rules, neither can they
within a mother’s clan. A compromise option could
be bilinearity (patrilineal or matrilineal combination)
of a kinship system, or searching for a third clan. The
paternal clan organization of the Tungus-Manchu
peoples as an assimilated community coexisted with
the principles of matrilineality, which had local roots.
Marrying a widow into a neighboring clan that could
open the way to exogamous relationships in doha
marked a transition of her children into a new father’s
clan, with a further prohibition for them on marital
relationships within the stepfather’s clan. Even the
groups living widely separated joined in doha, which
indicates the process of fragmentation of large local
communities into small ones, their intermixing, and
movement caused by the migration of Tungus. As a
result, the neighbors could be kinship groups pulled
together through the institution of doha. It is also
assumed that the emerged Samar tribe could have been
based on the original genetic kinship of its members
(Tugolukov, 1972).
In this article, we investigate the modern spread
of the Samar clan in Khabarovsk Territory, its gene
pool by using SNP markers of the Y-chromosome,
and the degree of the genetic kinship with other Nanai
populations and peoples in the adjacent territories.
Features of genogeographic research
In recent decades, studying the genetic diversity
in human populations has become one of the most
important research areas. This is partly because of
the discovery of the so-called non-recombining uni-
parental markers, localized in mitochondrial DNA and
the Y-chromosome of humans. Since mitochondrial
DNA is maternally inherited, and the Y-chromosome
is paternally inherited, they engender separate ideas
of the contributions of different sexes to the history of
the gene pool’s formation. At present, phylogeographic
research on Y-chromosome markers is conducted
more intensely than that on mtDNA, because the
Y-chromosome revealed a greater genetic diversity
and a clear correlation with the geographical position
and ethnic history of the population (Balanovskaya,
Balanovsky, 2007). Since this article is based on the use
of Y-chromosome markers, we will focus further on the
features of its genome and characteristic of its types of
polymorphism.
The length of the Y-chromosome is approximately
60 million pairs of nucleotides, its end (telomeric)
sections cross over with homologous sections of
148 Y.V. Bogunov et al. / Archaeology, Ethnology and Anthropology of Eurasia 43/2 (2015) 146–152
X-chromosome. All the remaining space (more than
50 million pairs of nucleotides) is a combination of
linked genes, which is not destroyed by recombination
and is transmitted from generation to generation
as a single haplotype. New haplotypes arise only
through mutagenesis. This mechanism of variability
allows the analysis of individual kinship between the
similar sequences of the non-recombining part of the
Y-chromosome, which is impossible for recombining
autosomal markers.
In the study of Y-chromosome diversity,
two types of genetic polymorphism (two types
of markers, respectively) are investigated. Single
nucleotide polymorphisms (SNP) quite often occur in
Y-chromosomes. These point-mutations very rarely
occur in the same position twice. SNP study can
allow attribution of the analyzed sample to one or
another branch of the family tree of the Y-chromosome
(haplogroup) or its sub-branches. Thus, the mutation
is a “marker” for all subsequent generations, and can
reliably distinguish one “clan” from another. Generally,
the designation of a haplogroup (e.g., N1c1) indicates
the name of the marker (index), which determines the
haplogroup (e.g., for N1c1, the index is M172). This
is because the Y-chromosomal tree of haplogroups is
constantly improving, which leads to changes in their
names; however, markers of specific mutations do
not change.
In the second type of polymorphism, the differences
between individuals are determined by the number
of alleles—short tandem repeats (STR)—in certain
Y-chromosome loci. Over several generations, this
indicator is maintained, i.e., a boy, his father, his
grandfather, etc. will have the same haplotype of the
Y-chromosome. However, there are changes in the
number of alleles—new mutations occur. They occur
signi¿ cantly more often than SNP, approximately once
in 22 generations (500 years).
Let us pay attention to the fact that if we do not know
what haplogroup the sample of interest belongs to, we
cannot accurately say whether or not the individual
is a relative of another individual with a similar set
of STR markers. To check this, one needs to analyze
SNP markers. Therefore, parallel study of both genetic
systems (SNP and STR) is the most effective way. They
are ¿ guratively comparable to hour and minute hands:
the ¿ rst one (SNP) shows the position of the sample in
the family tree in a “global” scale, and the second (STR)
shows the position that is more accurate. The study
of STRs while also having SNP data allows dating of
the emergence of separate clusters of haplotypes (i.e.,
branches of the family tree of the Y-chromosome). There
are two approaches to the analysis of SNP markers of
the Y-chromosome. In the ¿ rst (traditional) one, a set
of separate, most informative markers is analyzed.
In the second approach (which became technically
possible only in recent years) the full nucleotide
sequence of Y-chromosome is deciphered, which allows
identi¿ cation of all Y-chromosome SNP markers that
are present in the sample. Owing to the high cost of
full genome sequencing, this approach in our work
was implemented for one sample (a representative of
haplogroup C3*-M130), and the ¿ rst approach was used
for all samples.
Thus, an individual possesses a set of genes that
contains information about his ancestors. The entire set
of changes in the gene pool of the population reÀ ects
the sequence of genetic transformations in a number
of generations. When we know the average frequency
of STR marker mutations in the Y-chromosome, we
can compare spectra of haplotypes in different human
populations, identify the degree of their proximity, and
trace the history of changes in Y-haplotypes. Therefore,
certain mutations in the Y-chromosome serve as a
“witness” to historical processes: they can reÀ ect time,
migrations, assimilation, and crossbreeding.
The general structure of genogeographic research is
as follows: collecting a set of biological samples from
men unrelated by kinship for at least three generations,
but belonging to the same population; genotyping STR-
and SNP-markers of the Y-chromosome; analyzing the
frequencies of haplogroups and calculating genetic
distances; and analyzing the diversity of haplogroups
with the subsequent phylogeographical analysis of STR
haplotypes within each haplogroup.
The history of the Samar clan is stored in its gene
pool. Over many generations, different branches of
ethnicity assimilated. However, from the perspective
of the abovementioned methodological mechanism
of modern population genetics, the studied gene
pool is not a mixture of historical gene pools. It
is well structured in the paternal lineages of the
Y-chromosome, which gives hope of identi¿ cation
of traces not only of the Tungus expansion, but the
subsequent layers as well.
Material and methods
Material for demographic studies included household
books of the localities (places of traditional Nanai
residence) of three administrative districts of Khabarovsk
Territory (Solnechny, Komsomolsky, and Nanaisky);
for genetic studies, we used samples of venous blood
of 37 men of the Samar clan from rural settlements of
Kondon (n = 19), Nizhniye Khalby (n = 10), Belgo
Y.V. Bogunov et al. / Archaeology, Ethnology and Anthropology of Eurasia 43/2 (2015) 146–152 149
(n = 5), Verkhnyaya Ekon (n = 1), Cherny Mys
(n = 1), Nizhnyaya Tambovka (n = 1). The sample
included individuals who were unrelated to a depth of
at least three generations.
DNA was isolated by phenol-chloroform extraction.
The concentration of DNA was determined by real-time
PCR using the Quanti¿ ler® Kit (Applied Biosystems).
Using a Y-filer™ PCR Amplification Kit (Applied
Biosystems) and 3130×l genetic analyzer (Applied
Biosystems), 17 STR loci of Y-chromosome were
analyzed. Based on the identi¿ ed haplotypes, a prediction
of Y-chromosome haplogroups was conducted, with
subsequent confirmation by genotyping of SNP
markers. Such an approach allowed us to determine
the haplogroups reliably. Additionally, for the sample
with the haplogroup C3*-M130 (×M48) we performed
full sequencing of the Y-chromosome using the BigY
technology developed by the genetic and genealogical
company FamilyTreeDNA, and determined the position
of the sample in the overall phylogenetic structure of
the haplogroup.
Comparison of the Samar clan gene pool with
other populations of Nanai, and ethnic groups from
other regions, was performed via the Y-base database
(www.genofond.ru). On the basis of the frequencies
of haplogroups using the DJgenetic software, the
genetic distances were calculated (Nei, 1975). Using
this matrix of genetic distances and Statistica software,
a multidimensional scaling was performed, which
graphically visualizes the degree of similarity between
the populations of interest.
Research results
Modern expansion of the Samar clan in the Khabarovsk
Territory. The largest number is registered in the
historic district of residence—the village of Kondon
in Solnechny District: in 2012, there were 93 men
and 111 women comprising 38.3 % of the village
population. Over the past century, the members of the
Samar clan also dispersed to the Amur River valley,
mainly in the territory of the Komsomolsky District:
Verkhnyaya Ekon – 3 %, Belgo – 8.76 %, and Nizhniye
Khalby – 18.9 %. In the localities of the Nanaisky
District, individuals with a Samar surname were absent,
or their share in the ancestral (family) structure of the
population did not exceed 1 %.
The “genetic portrait” of the Samar clan. As a
result of Y-chromosome genotyping, it was found
that the “genetic portrait” of today’s clan members
is characterized by the presence of ¿ ve haplogroups:
N1c1-M178, C*-M130, I*-M170, J2a1ɚ-M47, and
O2-P31. The basis of the gene pool is haplogroup N1ɫ1
(over 83 %), the signi¿ cant predominance of which
can be seen as a generic characteristic feature that
distinguishes this population from other Nanai clans.
The North Eurasian haplogroup N1ɫ1 is found
in the gene pools of the peoples of Siberia with high
frequency: Yakuts (approx. 90 %), Buryats (in some
populations up to 78 %), Evenki (34 %), and Far-Eastern
Chukchas (61 %) and Koryaks (24 %) (Stepanov et al.,
2001; Kharkov, Stepanov, 2005; Kharkov et al., 2014).
The accumulation of this haplogroup in the Samar
clan indicates the presence of genetic connection of
the studied population with northern populations and
affirms the historical reconstructions of the Samar
migration-À ows to Amur.
The high frequency of the haplogroup J2a1a-M47
(8.1 %) is the second characteristic feature of the gene
pool of the Samar clan. According to our data, the
Middle Eastern haplogroup J2 is not found in the gene
pools of other Nanai clans in the Khabarovsk Territory
(unpublished data). It is found in Evenki and Evens
with a frequency of less than 2 %, and is not typical for
other peoples of Siberia; while in the ethnic groups of
the Central Asia it reaches up to 12 %.
The haplogroup C3*-M130, being one of the minor
components in the gene pool of the Samar clan, can
mark links with a very wide range of peoples who have
it. In order to clarify, we performed a full sequencing
of the sample with this haplogroup. We discovered a
great similarity with its carriers in other populations
of Nanai and Nivkh, and a signi¿ cant difference from
those in other peoples of Siberia and Central Asia. Thus,
the presence of the haplogroup C3*-M130 marks low-
frequency but clearly traceable genetic links of the
Samar clan to other populations of Amur.
Genetic relationships between populations. We
calculated the genetic distances between the gene
pools of peoples of Siberia and East Asia. For this
analysis we used the panel of 13 major haplogroups,
which are both most typical for the populations in
interest and well covered in the references. The
analysis included three Nanai populations: the Samar
clan (NAN-SAM), the total remaining population of
other Nanais of Amur (NAN-AMUR), and Nanais
of China (NAN-CHI). In the genetic space on the
multidimensional scaling plot, three clusters were
revealed (see Figure); each of them included one
of the Nanai populations. This indicates signi¿ cant
genetic differences between them. Nanais of China
joined Evens (genetic distance d = 0.09), Chinese
(0.10 < d < 0.12), and Vietnamese (d = 0.11). However,
their gene pool ¿ nds less signi¿ cant but still marked
similarity to the gene pools of Mongols (d = 0.21) and
150 Y.V. Bogunov et al. / Archaeology, Ethnology and Anthropology of Eurasia 43/2 (2015) 146–152
Orochi (d = 0.22). By contrast, Nanais of Amur cluster
together with Evenki (d = 0.05), Mongols (d = 0.09),
Buryats of the Transbaikal Territory (0.04 < d < 0.13),
and Orochi (d = 0.15).
The gene pool of the Samar clan is the most
peculiar: the genetic distance between the clan and
other Nanais of Amur (d = 0.60) is almost twice more
than between Nanais of Amur and Nanais of China
(d = 0.36). Such signi¿ cant differences between the
gene pools of Nanais in the Khabarovsk Territory are
associated with the prevalence of the North Eurasian
haplogroup N1c1-M217, and the low frequency of the
haplogroup C3c-M48 in the Samar clan. Therefore, its
gene pool is similar to that of Buryats in the Transbaikal
Territory (d = 0.01), Yakuts (d = 0.01), and Khakas
(0.01 < d < 0.15), despite the huge geographical
distances. The similarity of the “genetic portraits” of
these ethnic groups may indicate the commonality of
certain stages of their ethnogenesis. Surprising is the
relative proximity of the gene pools of the Samar clan
and the Eskimo people (d = 0.18). These populations
formed a single cluster in the genetic space.
Features of the gene pool of the Samar clan generally
con¿ rm the hypothesis of the historians regarding the
origin of the Samar people from more northern peoples
of Siberia. According to this hypothesis, they are
descendants of Evenki, who experienced the ¿ rst impact
of Yakuts, and then (in the 17th century), Buryats.
Despite the fact that the proportion of the haplogroup
N1c1 in the gene pool of modern Evenki does not exceed
34 %, Yakuts and Buryats have a more significant
accumulation—90 % and 47 %; accordingly ( “eastern”
Buryats can have the value of up to 78 %) (Kharkov,
Stepanov, Medvedeva et al., 2008; Kharkov, Khamina,
Medvedeva et al., 2014). In the studied population of
Evenki (Transbaikal Territory), the frequency of this
haplogroup is also low (23 %). However, the classical
genetic markers previously identified an extremely
high level of genetic diversity of this ethnic group,
which is well explained by considering its huge area
and extremely small population. This leads to genetic
drift, in which signi¿ cant differences between the gene
pools of the Evenki populations emerge. Therefore,
we can assume that the source of the gene pool of the
Samar clan was the Evenki subpopulation characterized
by higher frequencies of the haplogroup N1c1, even
though this is not necessarily true: the founder of a clan
could have been one of the carriers of this haplogroup.
The relative contribution of Evenki, Yakuts, and Buryats
can be assessed later, after the analysis of variants of the
haplogroup N1c1—STR haplotypes speci¿ c to the gene
pools of each of these ethnic groups.
The position of the Nanai populations within the genetic space of the peoples of Siberia and East
Asia: multidimensional scaling.
ALT – Altaians; BUR-1 – Buryats (Transbaikal Territory); BUR-2 – Buryats (Selenginsky and Khorinsky Districts,
Republic of Buryatia); BUR-3 – Buryats (Zakamensky District); VIE – Vietnamese; CHI – Chinese (Zhong et al.,
2010; Cai et al., 2011); NAN-SAM – Nanais of the Samar clan; NAN-AMUR – Nanais of Amur (Khabarovsk
Territory); NAN-CHI – Nanais of China (Chen et al., 2006); MON – Mongols (Kim et al., 2011); ORO – Orochi
(Xue, 2006); KH-SUM – Khakas, the total population; KH-SAG – Sagai; KH-KACH – Kacha; EWE – Evens;
EVE – Evenki (Zhong et al., 2010); ESK – Eskimo (Lell et al., 2002); YAK – Yakuts.
Y.V. Bogunov et al. / Archaeology, Ethnology and Anthropology of Eurasia 43/2 (2015) 146–152 151
Conclusions
Increasingly in the 20th century, the collapse of
the clan organization and the establishment of
territorially neighboring relations of the peoples of
Amur have contributed to blurring the boundaries of
intra-ethnic structures, and inter-ethnic contacts only
accelerated this assimilation. In this situation, and also
in view of the low number of Amur ethnic groups,
it is extremely dif¿ cult to conduct research on their
ethnogenesis. Modern genetics, based on markers of
the Y-chromosome and an intra-ethnic level of gene
pool structuring, opens up new sources of information,
and, therefore, new opportunities for studying the
ethnic groups.
A signi¿ cant proportion of modern Nanais currently
live in the vicinity of other ethnic groups and have close
contacts with them, mainly with Russians. However,
the Nanai clan Samar (area around the current Kondon
settlement) is unique in this respect. Considerable
remoteness from the main range of Nanais and dif¿ cult
access to the territory have contributed to a long
period of almost isolated existence of the population
and, hence, the preservation of the original gene pool.
This is directly evidenced by the low diversity of the
haplogroups and the preferential accumulation of only
one Y-chromosomal lineage.
The “genetic portrait” of the Nanai clan Samar
established herein indicates its Tungus origins. Over
a long period of Aldan-Lena-Amur migration to the
Devyatka River, the original gene pool has experienced
multiple external inÀ uences, as demonstrated by the
similarity to the gene pools of modern Yakuts and
Buryats. Archaeological and ethnographic materials, as
well as written records, also indicate the participation of
the peoples of northeastern Siberia in the ethnogenesis
of the Samar clan (Malyavin, 1998; Medvedev, 2005;
Medvedev, Filatova, 2014: 148–149). The degree of
such influence will be assessed in the future, after
performing a phylogeographic analysis of the identi¿ ed
haplotypes of the Y-chromosome.
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... This questionnaire, aimed mainly at the implementation of the "rule of three generations," also helps to identify family ties. The fact that the pedigree profile contains information about belonging to the clan not only reveals the kinship but also opens up a new direction in the genogeography, making it possible to more accurately study the structure of the population system and monitor migration flows [38][39][40][41][42][43][44][45]. ...
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Population biobanks are collections of thoroughly annotated biological material stored for many years. Population biobanks are a valuable resource for both basic science and applied research and are essential for extensive analysis of gene pools. Population biobanks make it possible to carry out fundamental studies of the genetic structure of populations, explore their genetic processes, and reconstruct their genetic history. The importance of biobanks for applied research is no less significant: they are essential for development of personalized medicine and genetic ecological monitoring of populations and are in high demand in forensic science. Establishment of an efficient and representative biobank requires strict observance of the principles of sample selection in populations, protocols of DNA extraction, quality control, and storage and documentation of biological materials. We reviewed regional biobanks and presented the organizational model of population biobank establishment based on the Biobank of Indigenous Population of Northern Eurasia created under supervision of E.V. Balanovska and O.P. Balanovsky. The results obtained using the biobanks in transdisciplinary research and prospective applications for the purposes of genogeography, genomic medicine, and forensic science are presented.
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In Rossiiskii Dalnii Vostok v drevnosti i srednevekovye: Otkrytiya, problemy, gipotezy. Vladivostok: Dalnauka
  • Neoliticheskiye Kultury
  • Nizhnego Priamurya
Neoliticheskiye kultury Nizhnego Priamurya. In Rossiiskii Dalnii Vostok v drevnosti i srednevekovye: Otkrytiya, problemy, gipotezy. Vladivostok: Dalnauka, pp. 234–267.
Clinal variability and territorial subdivision
  • Gene Pool
  • Buryats
Gene pool of Buryats: Clinal variability and territorial subdivision. Russian Journal of Genetics, vol. 50 (2): 203–213.
Yavleniye arkheologicheskoi lokalii
  • Drevniye Pamyatniki R
  • Devyatki
Drevniye pamyatniki r. Devyatki: Yavleniye arkheologicheskoi lokalii. In Project Amur. Tsubkuba: University of Tsubkuba, pp. 53–68.