Testing Central and Inner Asian admixture among contemporary Hungarians

Full text

Testing
Central
and
Inner
Asian
admixture
among
contemporary
Hungarians
Andra
´s
Bı
´ro
´
a
,
Tibor
Fehe
´r
b
,
Guszta
´v
Ba
´ra
´ny
b
,
Horolma
Pamjav
b,
*
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
1.
Introduction
For
centuries,
great
efforts
were
made
by
Hungarian
historians
to
study
the
earliest
period
of
their
national
history.
While
the
academic
mainstream
was
clearly
in
favour
of
the
Hungarian
language
belonging
to
the
Uralic
family,
many
other
researchers
favour
the
theory
of
a
closer
relationship
with
the
Turkic
language
family
and
Turkic
peoples.
Anthropological
analysis
of
bones
originating
in
the
10th
century
showed
characteristics
of
Central
Asian
origin
[1–3].
Archaeological
remains
of
weapons,
haversacks,
belt
mountings,
and
ornaments
on
clothing
also
showed
similari-
ties
to
those
of
Central
and
Inner
Asia
[4–7].
Hungarian
archae-
ologists
and
ethnographers
showed
that
there
are
similarities
in
the
traditions
of
the
ancient
Hungarians
and
various
Central
and
Inner
Asian
cultures
[8–10].
These
were
in
the
areas
of
burial,
belief,
and
figurative
arts.
Therefore,
an
origin
of
the
Hungarian
language
and
early
culture
in
a
region
ranging
from
Asia
to
Siberia
is
suggested,
but
a
specific
origin
has
been
difficult
to
identify.
Finns
were
thought
to
be
close
genetic
relatives
of
Hungarians.
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
was
proven
between
two
Hungarian
ethnic
groups,
the
Csangos
and
Seklers.
Both
groups
showed
genetic
affiliations
with
certain
Central
Asian
and
European
populations.
These
findings
could
have
Forensic
Science
International:
Genetics
15
(2015)
121–126
A
R
T
I
C
L
E
I
N
F
O
Keywords:
Hungarian
speakers
Central
and
Inner
Asian
populations
Y-STRs
Y-SNPs
A
B
S
T
R
A
C
T
Historically,
the
Carpathian
Basin
was
the
final
destination
for
many
nomadic
peoples
who
migrated
westward
from
Inner
and
Central
Asia
towards
Europe.
Proto-Hungarians
(Steppe
Magyars)
were
among
those
who
came
from
the
East,
the
Eurasian
Steppe
in
the
early
middle
ages.
In
order
to
detect
the
paternal
genetic
contribution
from
nomadic
Steppe
tribes,
we
tested
966
samples
from
Central
Asian
(Uzbekistan,
Kazakhstan),
Inner
Asian
(Mongolians
and
Buryats
in
Mongolia)
and
Hungarian-speaking
European
(Hungarian,
Sekler
and
Csango)
populations.
We
constructed
median-joining
networks
of
certain
haplogroups
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
haplogroup
resolution,
including
ancient
DNA
data.
ß
2014
Elsevier
Ireland
Ltd.
All
rights
reserved.
*Corresponding
author
at:
Institute
of
Forensic
Medicine,
Network
of
Forensic
Science
Institutes,
Ministry
of
Justice,
PO
216,
1536
Budapest,
Hungary.
Tel.:
+36
1
457
01
83;
fax:
+36
1
457
0182.
E-mail
address:
phorolma@hotmail.com
(H.
Pamjav).
Contents
lists
available
at
ScienceDirect
Forensic
Science
International:
Genetics
jou
r
nal
h
o
mep
ag
e:
w
ww
.elsevier
.co
m
/loc
ate/fs
ig
http://dx.doi.org/10.1016/j.fsigen.2014.11.007
1872-4973/ß
2014
Elsevier
Ireland
Ltd.
All
rights
reserved.

supported
theories
about
a
partially
Asian
origin
of
Hungarian
population
[11].
However,
most
of
the
Central
Asian-Hungarian
Y-chromosomal
relationship
was
based
on
the
high
frequency
of
haplogroup
R1a-M198
among
Kyrgyz
and
a
small
Hungarian
sample,
without
knowing
the
deep
structure
of
this
haplogroup.
Since
then,
first
Pamjav
et
al.
[13],
and
then
in
a
more
comprehensive
analysis
by
Underhill
et
al.
[14]
it
was
shown
that
there
is
a
clear
SNP-based
distinction
between
Eastern
European
(Z282,
Z280,
M458)
and
Central
Asian
(Z93)
R1a-M198
males.
It
was
also
noted
that
Hungarians
show
very
limited
or
no
presence
of
Haplogroup
N-M231
–
including
subclade
N1c-Tat
–
which
is
frequent
among
other
Uralic-speaking
populations
[15,16].
However,
the
potential
genetic
relationship
with
Turkic
and
Inner
Asian
peoples
has
been
less
researched,
although
this
relationship
could
shed
light
on
the
genetic
basis
of
the
alternative
Turkic
(Turanian)
theory.
Different
Turkic-speaking
populations
have
widely
differing
Y-chromosomal
gene
pools.
They
range
from
N1c-Tat
dominated
Yakuts
through
C3-M47
dominated
Kazakhs,
and
Q-M25
dominated
Turkmens
to
genetically
more
diverse
Uzbeks,
Azeri
and
Anatolian
Turks
(Table
S1).
Therefore,
we
chose
not
to
focus
only
on
haplogroup
frequencies,
but
on
analysing
haplotype
structure.
We
have
undertaken
a
survey
of
966
samples
from
Europe
and
Asia.
This
study
is
expected
to
provide
insights
relevant
to
the
Central
and
Inner
Asian
genetic
contribution
into
Hungarian
speaking
populations.
It
will
also
provide
insight
into
how
the
genetic
variation
is
distributed
in
the
contemporary
Hungarian,
Central
and
Inner
Asian
population
gene
pool
studied.
Supplementary
table
related
to
this
article
can
be
found,
in
the
online
version,
at
doi:10.1016/j.fsigen.2014.11.007.
2.
Materials
and
methods
2.1.
DNA
samples
To
analyse
the
genetic
relationship
of
present-day
Hungarians
with
present-day
Central
and
Inner
Asians,
we
tested
522
samples
from
Hungarian-speaking
populations
(332
Hungarians
from
Hungary,
95
Sekler
from
Romanian
Transylvania,
95
Csango
from
Romanian
Moldova),
115
Uzbek
samples
from
various
parts
of
Uzbekistan
(Ferghana
Valley,
Tashkent,
Khwarezm,
Samarqand,
Surkhodarya,
Karakalpakstan),
8
samples
from
Kazakhstan’s
Aqto
¨be
region,
127
Mongolian
and
88
Buryat
Mongolian
samples
from
Mongolia.
Archaic
Sekler
and
Csango
populations
were
included
to
increase
the
matching
potential,
and
we
also
collected
additional
samples
from
tribes
whose
self-designation
may
have
connection
to
the
ethnonym
Magyar,
i.e.
61
Madjars
from
Uzbekistan
and
45
Madjars
from
Kazakhstan.
Out
of
the
966
samples,
the
45
Kazakh
Madjars
[17],
and
215
Hungarian
samples
[12]
were
published
before,
but
tested
for
further
SNPs
and
samples
in
this
study.
The
new
samples
published
herein
were
sent
to
the
YHRD
and
the
accession
numbers
are
the
following:
Uzbekistan
[Uzbek]
YA003994,
Uzbekistan
[Madjar]
YA003995,
Mongolia
[Buryat]
YA003996,
Mongolia
[Mongolian]
YA003997
and
Kazakhstan
[Mad-
jar,
Aqto
¨be]
YA003998.
The
new
populations,
as
well
as
the
previously
published
populations
were
Hungarian
[12],
with
accession
number
YA003187,
Csango
[Romanian]
YA002984
[18]
and
Sekler
[Romanian]
YA002983
[18].
These
were
used
for
comparison
and
can
be
referenced
at
www.yhrd.org.
The
Y-SNP
haplogroups
for
Sekler
and
Csango
populations
were
tested
by
us
and
included
in
our
data.
Each
person
gave
their
informed
consent
prior
to
their
inclusion
in
the
study.
2.2.
Testing
of
Y-STR
and
Y-SNP
markers
DNA
was
amplified
with
the
PowerPlex
Y
(Promega,
USA)
amplification
kit
including
12
Y-STR
loci,
according
to
the
manufacturer’s
instructions.
Fragment
sizes
and
allele
designa-
tions
were
determined
with
a
3130
Genetic
Analyzer
(Life
Technologies,
Foster
City,
CA)
using
GeneMapper
IDX
1.2.1.
soft-
ware.
When
testing
Y-SNP
markers,
amplifications
of
1–2
ng
genomic
DNA
were
performed
in
an
ABI
7500
Real-time
PCR
instrument
with
Taqman
Assay
(Life
Technologies,
Foster
City,
CA)
using
the
programmes
designed
by
the
manufacturer.
The
relative
fluorescence
of
the
PCR
products
were
analysed
on
an
ABI
7500
with
its’
SDS
software,
as
described
in
the
manufacturer’s
manual
(Life
Technologies,
Foster
City,
CA).
Fifty-five
Y-chromo-
somal
SNP
markers
were
tested
with
Taqman
Assays
(Fig.
1).
The
haplogroups
tested
and
the
markers
used
in
the
study
originated
from
YCC
(Y-Chromosomal
Consortium).
The
nomenclature
of
haplogroups
followed
the
ISOGG
2014
Y-DNA
haplogroup
tree
due
to
recent,
new
additions
uncovered
by
YCC
(Y-Chromosomal
Consortium).
A
list
of
primers
and
Taqman
probes
for
binary
markers
was
previously
published
[19],
but
we
now
updated
the
list
with
new
SNPs
studied,
as
shown
in
Table
S2.
A
new
downstream
SNP
marker,
L24,
was
tested
for
J2*-M172
(xM47,
M67,
M12)
samples
to
obtain
more
resolution
within
the
haplogroup
as
suggested
by
van
Oven
et
al.
[20].
Supplementary
table
related
to
this
article
can
be
found,
in
the
online
version,
at
doi:10.1016/j.fsigen.2014.11.007.
2.3.
Data
analysis
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
DYS389II
and,
as
is
common
practice,
the
locus
DYS385
was
excluded
from
the
network.
Within
the
network
programme,
the
rho
statistic
was
used
to
estimate
the
time
to
the
most
recent
common
ancestor
(TMRCA)
of
haplotypes
within
the
compared
haplogroups.
3.
Results
Based
on
10
Y-STR
loci,
networks
were
constructed
within
each
of
the
haplogroups.
These
haplogroups
overlapped
among
populations
studied.
All
haplotype
and
haplogroup
results
can
be
found
in
Table
S3.
Haplogroup
results
are
summarized
in
Table
S4.
For
our
analysis,
we
only
considered
those
haplogroups
which
occurred
in
more
than
one
sample
among
both
Hungarians
and
Central
Asians
(Uzbeks,
Kazakhs,
Madjars),
or
among
Hungarians
and
Inner
Asians
(Mongolians,
Buryats).
With
this
method,
we
identified
nine
haplogroups,
which
might
indicate
a
genetic
relationship
between
contemporary
Magyars
and
Altaic-
speaking
populations.
They
are
E-M78,
G2a-P15,
J2*(xM47,
M67
and
M12),
N1c-L708,
Q-M242,
R1a-M458,
R1a-Z280,
R1a-Z93
and
R1b*-P25(xM412).
To
verify
or
confute
the
relationships,
we
created
median-joining
networks
on
10
loci
for
all
‘‘suspected’’
haplogroups.
Results
are
discussed
only
in
the
context
of
the
potential
matches
between
Hungarian
and
Altaic-speaking
popu-
lations,
haplogroup
by
haplogroup.
The
haplogroups
are
described
as
follows.
Supplementary
tables
related
to
this
article
can
be
found,
in
the
online
version,
at
doi:10.1016/j.fsigen.2014.11.007.
3.1.
Haplogroup
E-M78
The
median
joining
network
(MJ)
of
31
E-M78
haplotypes
is
shown
in
Fig.
2A.
The
network
shows
a
star-like
pattern.
The
biggest
cluster
(cluster
1
in
Fig.
2A)
was
the
modal
haplotype
shared
by
three
Hungarian
speaking
population
groups,
which
A.
Bı
´ro
´et
al.
/
Forensic
Science
International:
Genetics
15
(2015)
121–126
122

consisted
of
two
Hungarian,
one
Sekler
and
three
Csango
males.
There
is
a
potential
that
two
Seklers
and
two
Hungarians
have
a
common
origin
with
Uzbeks
(one
from
Tashkent,
one
from
Khwarezm)
on
the
bottom
of
the
network.
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
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
and
one
Csango
male.
The
second
largest
cluster
consists
of
four
Hungarians
(Fig.
2G,
cluster
2).
An
interesting
picture
is
noted
in
that
one
Uzbek
from
Khwarezm
and
one
Madjar
from
Kazakhstan
are
in
the
middle
of
the
network
connecting
the
two
separate
clusters
as
Fig.
1.
A
phylogenetic
tree
of
the
tested
55
Y-SNP
loci.
A.
Bı
´ro
´et
al.
/
Forensic
Science
International:
Genetics
15
(2015)
121–126
123

the
median
haplotype
(Fig.
2G,
cluster
1).
One
Csango
descends
from
this
central
haplotype
and
thus
can
be
designated
as
Central
Asian
admixture.
While
R1a-M458
is
generally
considered
as
an
Eastern
European
haplogroup,
being
especially
frequent
among
Western
Slavs
and
to
a
lesser
extent,
Eastern
Slavs
[24],
based
on
our
result
we
cannot
exclude
the
possibility
that
R1a-M458
originates
from
Central
Asia.
These
Khwarezm
and
Madjar
haplotypes
may
be
the
remnants
of
the
ancestral
population.
Attributing
one
Kazakh,
one
Kazakh
Madjar,
and
all
three
Uzbek
R1a-M458
haplogroups
to
Slavic
admixture
seems
unlikely,
especially
given
the
nearly
complete
lack
of
other
typically
European
haplogroups
I-M170
and
R1b-M412
among
our
Central
and
Inner
Asian
samples
(Table
S3).
3.7.
Haplogroup
R1a-Z280
The
R1a-Z280
haplotypes
produced
a
star-like
network
(figure
not
shown),
with
all
the
Central
Asians
exactly
matching
Hungarian
haplotypes
(Table
S4).
Therefore,
we
assume
that
a
genetic
link
was
from
Finno-Ugric
or
Slavic
peoples
to
Central
Asians
[13].
3.8.
Haplogroup
R1a-Z93
Thirty-six
R1a-Z93
haplotypes
produced
a
non-star
like
and
very
diverse
network
(Fig.
2H).
These
included
mostly
Uzbek
haplotypes
found
in
the
central
area,
who
were
Hungarian-
speaking,
Uzbek
Madjar,
Buryat
and
Mongolian
populations
were
branching
off
towards
the
edges.
While
some
R1a-Z93
haplotypes
might
be
a
result
of
Roma
admixture
[13],
for
the
purpose
of
this
study
we
assumed
that
all
haplotypes
of
Hungarian-speaking
population
groups
to
be
Central/Inner
Asian
admixture.
3.9.
Haplogroup
R1b-P25
The
network
of
44
R1b-P25
(xM412)
haplotypes
clearly
consists
of
two
clusters
(Fig.
2I).
On
the
left
side,
we
find
the
star-like
network
of
M269
haplotypes,
while
on
the
right
side,
the
typically
Central
Asian
subgroup
M73
is
visible
(Note:
M269
and
M73
markers
were
not
tested
in
this
study,
but
a
comparison
with
Myres
et
al.
[25]
STR-data
suggest
the
connection).
Among
R1b-
P25
haplotypes,
no
connection
can
be
made,
as
Hungarian-
speakers
dominantly
belong
to
the
M269
branch,
while
Uzbeks
belong
to
the
M73
part.
The
limited
number
of
Central
Asians
(M269)
is
situated
on
the
edges
of
the
network,
thus
representing
external
admixture
rather
than
source.
Cluster
2
(Fig.
2I)
in
the
part
of
M269
consists
of
four
Hungarians
and
one
Sekler
male.
4.
Discussion
On
examination
of
haplogroups
with
an
N
>
1
frequency
among
both
Hungarian-speaking
European,
and
Altaic-speaking
Central
and
Inner
Asian
populations,
we
showed
that
the
possible
maximum
Central/Inner
Asian
admixture
among
contemporary
Hungarian
populations
ranges
around
5–7.4%.
We
took
into
account
only
those
haplotypes
which
could
derive
from
Central/
Inner
Asian
haplotypes
according
to
the
MJ-networks.
The
admixture
was
lowest
among
Hungarians
from
Hungary
(5.1%),
while
somewhat
higher
among
Hungarian-speaking
populations
in
Romania,
notably
Sekler
(7.4%)
and
Csango
(6.3%).
The
average
of
these
results
was
5.7%
among
522
Hungarian-speaking
males
(see
Table
S4).
The
reason
of
the
difference
might
be
the
long-time
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.
A.
Bı
´ro
´et
al.
/
Forensic
Science
International:
Genetics
15
(2015)
121–126
124

isolation
of
Sekler
and
Csango
groups,
resulting
in
lower
admixture
from
neighbouring
populations.
However,
we
also
must
acknowl-
edge
that
these
numbers
represent
an
upper
limit
and
that
actual
Central
and
Inner
Asian
admixture
might
be
somewhat
lower.
In
these
admixture
cases,
the
genetic
links
are
not
necessarily
directly
from
Altaic
populations
to
Hungarians,
as
both
populations
may
have
received
these
genetic
markers
from
a
common
third
unidentified
source
(e.g.
Middle
East,
Caucasus,
and
East
Slavs).
Because
of
this
possibility,
further
research
is
needed.
We
also
have
to
note
that
Central
Asian
admixture
among
Hungarians
does
not
necessarily
come
from
Altaic-speakers.
It
may
also
come
from
ancient
Iranian
tribes
who
were
later
Turkicized
by
Altaic
conquerors.
The
main
haplogroups
responsible
for
the
Central/
Inner
Asian
admixture
among
Hungarians
are
J2-M172
(xM47,
M67,
L24,
M12),
J2-L24,
R1a-Z93,
Q-M242
and
E-M78.
Earlier
studies
reported
that
Haplogroup
E,
J
and
their
main
subgroups
spread
from
the
Middle
East
with
the
Neolithic
agricultural
revolution
[26–28].
It
spread
towards
both
Europe
and
Central
Asia,
thus
some
of
the
common
haplotypes
such
as
E-
M78,
J2-M172*
and
J2-L24
may
indicate
a
common
Middle
Eastern
origin
for
both
Hungarian-speaking
and
Central/Inner
Asian
samples.
This
is
in
contrast
to
the
idea
of
a
male
migration
from
Asia
towards
the
Carpathian
Basin.
Based
on
the
results
of
the
Central
and
Inner
Asian
samples
analysed
in
this
study,
we
could
not
find
a
strong
genetic
relationship
with
contemporary
Hungarian-speaking
populations
which
would
imply
a
common
origin
for
these
populations
in
the
past.
Hungarians
were
heavily
affected
by
neighbouring
popula-
tions
and
also
had
effect
on
them.
We
should
also
take
into
account
that
the
East
Central
European
region
received
Central
and
Inner
Asian
genetic
influence
both
before
(Sarmatians,
Huns,
Avars,
Onogur-Bulgars)
and
after
(Pechenegs,
Yassic
people,
Cumans)
the
Hungarian
settlement
in
the
Carpathian
Basin
[3].
It
is
impossible
to
separate
the
genetic
effects
of
these
different
migrations
based
on
DNA
results
from
contemporary
populations.
The
Central
Asian
gene
pool
also
underwent
significant
changes
due
to
mediaeval
Turkic
and
Mongolian
invasions
as
well
[3].
Assuming
a
Central
Asian
origin
for
Eastern
European
subgroups
of
haplogroup
R1a-M198,
the
share
of
Central
Asian
ancestry
would
significantly
increase,
up
to
almost
30%.
But
then
neighbouring
Western
and
Eastern
Slavic
peoples
would
have
an
even
higher
Central
Asian
admixture
(50–60%)
than
that
of
Hungarians.
Despite
the
similarity
of
tribal
names
among
Kazakh
and
Uzbek
Madjars,
a
significant
genetic
connection
could
not
be
established;
this
based
on
our
meticulously
selected
samples,
which
included
pedigree
analyses.
Kazakh
Madjars
(dominated
by
Hg
G1)
differ
significantly
from
other
Kazakhs
and
do
not
exhibit
a
relationship
with
Caucasian
peoples
(Hg
G2).
Uzbek
Madjars
are
more
heterogeneous,
although
still
dominated
by
Hgs
C3
and
D,
which
are
entirely
absent
from
present-day
Hungarian
speakers.
Comprehensive
surveys
of
more
Central/Inner
Asian
and
less-
admixed
population
groups
in
Hungary,
including
pedigree
analyses,
for
deep-resolution
haplogroups
need
to
be
conducted
in
future
studies
to
be
able
to
draw
more
robust
conclusions
regarding
the
origins,
spread
and
genetic
affiliations
of
contempo-
rary
populations.
Earlier
studies
have
highlighted
the
very
low
genetic
affinity
of
present-day
Hungarians
with
linguistically
related
Uralic
peoples
[16].
A
recently
published
study
showed
a
limited
genetic
link
between
Hungarians
and
Ugric-speaking,
Western
Siberian
Mansi
based
on
the
new
subhaplogroup
N-L1034
defined
by
L1034
SNP
mutation
[29].
The
typically
Uralic
haplogroup
N
accounted
for
only
1.7%
among
the
Hungarian
samples,
this
being
even
lower
than
a
potential
Turkic
admixture.
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.
However,
we
could
not
show
any
significant
genetic
correlation
between
Hungarian-speaking
and
Central/Inner
Asian
samples
which
would
explain
the
linguistic
difference
among
Hungarians
and
neighbouring
populations.
At
the
same
time,
the
reliable
historical
and
genetic
conclusions
require
an
extension
of
the
study
to
a
significantly
larger
database
with
deep
haplogroup
resolution,
including
ancient
DNA
data.
Conflict
of
interest
The
authors
declare
no
conflict
of
interest.
Acknowledgements
We
would
like
to
say
special
thanks
to
Dr.
Eva
Susa
(General
Director
of
the
Network
of
Forensic
Science
Institutes)
for
her
financial
support.
We
thank
sample
donors
and
Betty-Jean
Sigethy
and
Rayn
Hoyt
for
the
English
editing.
We
say
special
thanks
to
two
unknown
reviewers
for
their
constructive
comments
and
sugges-
tions.
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