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Genetics and the Origin of Human "Races"

Article  in  Genetika · September 2001

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Russian Journal of Genetics, Vol. 37, No. 8, 2001, pp. 853–867. Translated from Genetika, Vol. 37, No. 8, 2001, pp. 1029–1045.
Original Russian Text Copyright © 2001 by Tetushkin.

THEORETICAL PAPERS
AND REVIEWS

Genetics and the Origin of Human “Races”

E. Ya. Tetushkin
Vavilov Institute of General Genetics, Russian Academy of Sciences, Moscow, 119991 Russia; e-mail: eugene@vigg.ru
Received February 7, 2001

Abstract—In the last decades, the concept of human races was considered scientifically unfounded as it was
not confirmed by genetic evidence. None of the racial classifications, which strongly differ in the number of
races and their composition, reflects actual genetic similarity and genealogy of human populations inferred
from variability of classical markers and DNA regions. Moreover, intercontinental (“interracial”) variability
was shown to be far lower than that within populations: the former constitutes 7 to 10% of the total genetic vari-
ation and the latter about 85% of it. It is believed that the low level of differentiation of regional population
groups contradicts their race status and suggests a recent origin of humans from one ancestral population. The
results of studies of various genetic systems are in agreement with the latter conclusion rejecting the hypothesis
of regional continuity. According to this hypothesis, the populations of continents regarded as large races have
developed during long evolution from local types of archaic humans, in particular, Neanderthals. Phenotypic
similarity of different, sometimes unrelated, populations united into one “race” is explained by strong selection
since race-diagnostic traits characterize body surface and thus are directly subjected to the influence of envi-
ronmental (primarily climatic) factors. It has been recently established that variability of the most important of
these traits, body and hair pigmentation, is largely controlled by one locus (MC1R), which accounts for its high
evolutionary lability. Other traits used for race identification are also likely to be labile and controlled by major
genes. However, the fact that the currently existing race classifications are groundless does not mean that such
classifications are impossible in principle. Commonly used argumentation (races do not exist because popula-
tions are not genetically separated) does not hold water. A polytypic species is characterized by genetic conti-
nuity of allopatric populations rather than the presence of narrow genetic boundaries between them. Borderlines
between races are usually conventional and arbitrary. As to intergroup variation in humans, it is indeed low but
comparable with that in a number of other species. There are no obstacles to the development of genetic sys-
tematics of human races.

INTRODUCTION dence obtained at the population, organismal, and molec-

In the last quarter of the 20th century, due to
advances in genetics, the concept of human races has
been regarded as not scientifically valid. As Tishkov
aptly pointed out, “American scientific community suc-
cessfully initiated the rejection of the notion of race as
a scientific category” [1].At present, at least in genetic
literature, this term is typically used in quotation marks.
At the same time, there is a special field, race science,
devoted to investigation of human races. This is a major
part of physical anthropology, which has long traditions
and ample accumulated material. It appeared that this
field has been losing its subject matter and thus was
destined to disappear.
This situation seemed disturbing to Russian anthro-
pologists. They responded (with a 20-year delay since
the ideas rejecting the race concept have been actively
circulating since the 1970s) by organizing in 1998 the
First International Conference “Race: Myth or Real-
ity”, which was held under the auspices of the Russian
Section of the European Anthropological Association
[2]. In addition to Russian researchers, the conference
hosted representatives of Belarus, Italy, Latvia, Mongolia,
Poland, and Estonia. The resolution adopted at the confer-
ence stated that the existence of races “is confirmed by evi-
ular levels of investigation of humans” [2].
Why then the most influential population and
molecular geneticists involved in this problem deny the
existence of races? The conference organizers believe
that “some foreign scientific schools…rejected the race
concept…because of the political situation” [2].
To be sure, in the developed countries political cor-
rectness in fact has an impact on various fields of activ-
ity including science. However, the refutation of the
race concept is based on specific population-genetic
arguments that absolutely lack political implications.
Moreover, race statistics has been conducted in the
United States until the present [1], which is at variance
with the “political” explanation of the “negative atti-
tude toward the race science” [2]. The conclusion on
the absence of races is based on scientific facts rather
than externally imposed. The existence of the official
race statistics testifies to the fact that the general com-
munity is not ready to accept this bold conclusion.
Surprisingly, the quoted resolution does not cite any
of the arguments against the race concept. In view of
the aims of the conference, it would seem logical to
present these arguments and try to invalidate them.
Without this, the position of race science’s supporters

1022-7954/01/3708-0853$25.00 © 2001 MAIK “Nauka /Interperiodica”

854 TETUSHKIN
looks unconvincing. Their key argument presented
above (the other arguments were not directly related to
the problem) does not stand elementary testing. It is
sufficient to compare schemes of genetic differentiation
of human populations presented in the comprehensive
review by Cavalli-Sforza and his students [3] with race
classifications proposed by Bunak [4], Cheboksarov
[5], Alekseev [6], and other authors. This comparison
demonstrates that these classifications do not reflect
actual genetic similarity of populations and give a dis-
torted picture of their phylogeny [7]. However, a more
or less reliable genealogy of the main human divisions
based on molecular markers was already established in
the 1980s when the cited monographs appeared.
To be fair, it should be noted that there are no studies
specifically concerned with the critique of the race con-
cept. The monographs by Cavalli-Sforza et al. [3, 8]
dealing with this problem do not consider all pros and
cons systematically and in detail. The latest book of this
outstanding geneticist (not published in Russia)
received very favorable reviews but also some criti-
cisms precisely for superficial and politicized discus-
sion of the race problems [9].
As the discussed problem has been generated by the
development in molecular evolutionary genetics, this
discipline should shed light on the situation. In what
follows, I consider the current state of the race concept
in the context of key advances in this field of genetics.
THE RACE CONCEPT
AND GENETIC DIVERSITY OF HUMANS
According to “genetic definition” of race proposed
by Vogel and Motulsky, “a race is a large population of
individuals who have a significant proportion of their
genes in common and can be distinguished from other
races by their common gene pool” [10]. But what is this
“significant proportion” and how could one isolate a
gene pool of a particular race from the spatial contin-
uum of gene frequencies encompassing the entire
humankind? It is clear that this definition is not opera-
tional: it cannot be used in practice.
As more adequate, the following definition can be
proposed: “human races are historically formed groups
of people connected by their common origin that man-
ifests in shared hereditary morphological and physio-
logical traits varying within certain limits.” This defini-
tion emphasizes that a race is a set of populations rather
than of individuals [11]. Similar definitions are given in
English-language explanatory dictionaries, which also
note that these traits are inherited (see, e.g., [12]).1
The idea of subdivision of the humankind into
groups of related population units (demes) seems quite
plausible, particularly in view of more than three centu-
1 Commenting on one of such definitions, Cavalli-Sforza [8]
remarked that “genetically” should be added to the phrase “can
be transmitted to progeny.”
RUSSIAN JOURNAL OF GENETICS
ries of experience in constructing race classifications.
What is the result of these investigations?
Modern anthropologists did not reach consensus
with regard to the number of “races” (varying from 3
to 200 in works of different authors) and their compo-
sition. Moreover, it is clear that their taxonomic and
phylogenetic constructions (overall and in details) are
at variance with more accurate pictures of genetic relat-
edness and differentiation of human populations based
on molecular markers.
To illustrate this point, compare a traditional classi-
fication of Caucasoids presented in a specialized ency-
clopedia [11] with a tree constructed on the basis of
gene frequencies of “classical” markers [3]. In the sys-
tem adopted by Russian authors, Caucasoids are classi-
fied into three race groups: southern, northern, and
intermediate. These groups differ mainly in pigmenta-
tion intensity. For this reason, these groups (secondary
races) are also called melanchroys (dark-haired), xan-
thochroys (blonde-haired), and brown-haired [5]. A com-
parison shown in Fig. 1 demonstrates that xanthochroys
are closely genetically related to both brown-haired
people and melanchroys inhabiting Western Europe.
These Western Europeans are more distant from East-
ern Europeans (mostly brown-haired) than from each
other. From the point of view of genetics, their division
into southern, intermediate, and northern cannot serve
even as the first approximation. Europeans also cannot
be divided into western and eastern, since some peoples
inhabiting Europe are not included in the cluster com-
prising most of the population of Eastern, Western, and
Central Europe. These outsiders include brown-haired
people, xanthochroys, and melanchroys. As to melano-
chroys (or, according to another terminology, members
of the Indo-Mediterranean race) from Asia and Africa,
which constitute majority in this group, they do not
group with Europeans and form separate clusters of
Caucasoids–non-Europeans. Thus, genetic classifica-
tion of Europeans, let alone Caucasoids in general, can-
not be confined to such simple schemes (see [13–16]).
One of a few results of the traditional systematics
that were seemingly confirmed is subdivision of the
humankind into large races. However, this issue is also
not completely settled. The authors of the aforemen-
tioned encyclopedic entry, following many other spe-
cialists, distinguish Caucasoid (Eurasian), Mongoloid
(Asian–American), Negroid (African), and Australoid
(Oceanic) races. They permit combining the latter two
into one Equatorial race but regard this view as less
substantiated. (Note that this viewpoint stated in
anthropology textbooks [17, 18] has been long refuted
by genetic studies [19–23]). The subdivision of the
humankind into four major groups does not contradict
results of Nei and Roychoudhury [21–23] (Fig. 2a).
However, there are some discrepancies as to the com-
position of these groups. For instance, anthropologists
generally assign Ainu to Australoids whereas based on
genetic distances they group together with Mongoloids
Vol. 37 No. 8 2001
 GENETICS AND THE ORIGIN OF HUMAN “RACES” 855

Dutch Brown-haired
Danes Xanthochroys
English
Swiss
Germans
Belgians Brown-haired
Austrians
French
Swedes
Norwegians Xanthochroys
Czechs and Slovaks Brown-haired
Portuguese
Italians Melanochroys
Spaniards
Hungarians
Poles Brown-haired
Russians
Scots
Irish
Xanthochroys
Finns
Icelanders
Basques
Yugoslavians Melanochroys
Greeks
Sardinians
Lapps Brown-haired

0.04 0.03 0.02 0.01 0

Genetic distance

Fig. 1. Tree of genetic relatedness of a number of European populations [3] with their “race” attribution.

rather than Australian aborigines and Papuans. South-
ern Indians speaking Dravidian languages (Dravidian
or Southern Indian race) are attributed to the Negroid
branch [24] or consider a transitory form intermediate
between the Negroid and Caucasoid races [5]. In real-
ity, this group is as distant from Negroids as the other
(assigned to Caucasoids) peoples of India, to which it is
rather closely related [3]. And this list can be continued.
Comparison of traditional classifications with the
results of Cavalli-Sforza et al. [3, 25, 26] reveal more
significant and essential differences. These authors
divide non-Africans into indigenous inhabitants of
Europe, Middle East, Near, Northern, Central, and
Eastern Asia, Indostan, and America, on one hand, and
the population of Southeast Asia, Oceania, New
Guinea, and Australia, on the other (Fig. 2b). In other
words, their genetic tree does not have a single cluster
dures used in their construction. Further studies are
required to choose between these alternatives.
The fact that the traditional race classifications are
erroneous does not necessarily mean that such classifi-
cation is impossible in principle. Maybe the previous
systems merely should be revised in order to create a
new genetic systematics [27] of human races? How-
ever, at present most authors agree that the existence of
“relatively genetically homogeneous groups (“races”)
distinguished by major biological differences is not
consistent with genetic evidence” [28]. What facts and
rationale underlie this viewpoint?
Before the 1970s, it was believed that race identifi-
cation based on gene frequencies supplements and
(a) New Guinea and Australia

corresponding to Mongoloids. The inhabitants of the Southeast Asia

northern part of Asia (up to southern China) are united East Asia
Pacific Islands;
with Caucasoids (European and Asian) and native America
Americans. All other Asians that are classified as Caucasoids–non-Europeans
Southasian transitory group [11] are united with inhab- Europe
itants of Oceania and Australia. This subdivision of Africa
non-Africans into two major groups is consistent with (b) New Guinea and Australia
the idea on parallel genetic and linguistic divergence Pacific Islands
advocated by these authors. Almost all members of the Southeast Asia
first group except Amerindians speak languages of the East Asia
Nostratic or overlapping with it Eurasian linguistic Northeast Asia
superfamily2. Most peoples of the second group speak America
Europe
languages of the Austric superfamily. The principal dif- Caucasoids–non-Europeans
ference in topology of the trees presented in Fig. 2 is Africa
explained, in particular, by different clustering proce-
Fig. 2. Phylogenetic trees of major groups of human popu-
2 Some authors believe that Amerindian languages except na-dene lations: (a) after Nei and Roychoudhuri (the tree was con-
also belong to the Nostratic superfamily (see [3, 26] for refer- structed in [53] based on data from [23]); (b) after Cavalli-
ences). Sforza et al. [3].

RUSSIAN JOURNAL OF GENETICS Vol. 37 No. 8 2001

856
refines race classifications based on morphological
characters (see [29, 30] for review). This view started to
change after the studies by Lewontin [31] and Nei and
Roychoudhury [32, 33]. These authors showed that
interracial and interpopulation genetic variation in pro-
teins and blood groups is lower than intrapopulation
variation. According to Lewontin, intrapopulation,
within-race interpopulation, and interracial diversity
respectively accounts for about 85.4, 8.3, and 6.3% of
the total variation of human populations. On the
strength of this close similarity of human races, a con-
clusion was made that “race classification is now seen
to be of no genetic or taxonomic significance either, no
justification can be offered for its continuance” [31].
Using different methodical approaches, Nei and Roy-
chouldhury also have found that the codon differences
among populations belonging to the three major
races—Caucasoid, Negroid, and Mongoloid3—consti-
tute a small part of the codon differences among two
genomes randomly selected from the same population.
For instance, the interracial codon differences between
Caucasoids and Negroids constitute only 7–10% of the
intraracial differences.
In the late 1970s, Mitton criticized the methods
employed by the above-cited authors and made an
attempt to refute their results [34]. However, the ensu-
ing discussion revealed that his approach to the prob-
lem of genetic differentiation was erroneous [35–38].
Interestingly, Nei [39], commenting on the discussion
later, challenged Lewontin’s view on futility and incon-
sistency of all racial classifications. Nei suggested that
this position reflected humanistic views of Lewontin
and might have been provoked by then-popular specu-
lations on racial differences in IQ rather than being
based on Lewontin’s statistical computations. Although
relative interracial gene differences are small, they can
be used for constructing a racial classification, if based
on high number of loci. According to Nei, investigation
of systematics of human races is crucial for understand-
ing human evolution. In an analogous comment, Neel
[40] emphasized distinguishing between two mutually
supplementing questions: (1) what portions of the total
genetic variation within a large group are explained by
differences among subgroups and among individuals?
and (2) are the interpopulation levels of allele fre-
quency variation sufficient for constructing a taxonomy
of human populations? Giving the positive answer to
the second question, Neel avoided using the term race
and referred to the largest subdivisions of the human-
kind as major ethnic groups. As to Nei, he excluded
from his vocabulary this term, together with generally
accepted race names, only at the turn of the 1990s (see,
e.g., [41–43]).
3 Lewontin
analyzed a wider range of populations that he classified
into Caucasoids, black Africans, Mongoloids, aborigines of
South Asia, Amerindians, inhabitants of Oceania, and Australian
aborigines.
TETUSHKIN
Following Lewontin, other authors carried out parti-
tioning of genetic variation of the humankind into com-
ponents [44–47]. The estimates of the corresponding
components obtained by them are similar. However, all
these studies are based on data on protein markers and
blood groups covering no more than 25 loci. It was
questionable whether polymorphism of these loci ade-
quately reflects the structure of genetic variation in
H. sapiens.
To shed light on this issue, in the late 1990s Barbu-
jani et al. [48] examined the diversity of 16 populations
from all continents for 109 DNA markers including
30 microsatellite loci and 79 polymorphic restriction
sites. These authors estimated differences between
individuals from the same populations, between popu-
lations from the same continent, and between geo-
graphical populations groups from different continents.
On average, the components of the total variation con-
stituted 84.4 (within populations), 4.7 (between popu-
lations within a continent) and 10.8% (between conti-
nents). At that, high variability was characteristic of all
populations, from the largest, encompassing whole
countries and subcontinents (e.g., China or Northern
Europe) to the smallest, localized in separate villages or
settlements of hunters–gatherers (although large popu-
lations were on average more variable than small ones).
These values of intergroup components reflect variation
in extremely small part of the genome because only
0.08% [49] of nucleotide sites of human DNA are poly-
morphic (10–15% of 0.08%!)
The relative homogeneity of the human gene pool
indicates short differentiation time and migration
between populations. As was shown in studies of mito-
chondrial DNA (mtDNA) and Y-chromosomal markers,
the rate of intercontinental migration was eight times
higher for women than for men [50, 51]. The intrapop-
ulation variation component of mtDNA is approxi-
mately equal to that of autosomal loci (81%) [52] but in
Y-chromosomal markers this value is much lower (only
36%); the greatest part of their total diversity 53% is
explained by the intercontinental component [50].
However, it should be kept in mind that gene flow inten-
sity was probably lower at the earlier stages of human
evolution as compared to the later stages.
The possibility of grouping populations on the basis
of small but significant genetic differences does not
remove doubts in the reality of human races. The mul-
tilevel, without marked chiatuses, hierarchic structure
of the groups hinders objective identification of the lev-
els corresponding to the beginning of race differentia-
tion. The choice of these levels is arbitrary and subjec-
tive, and the taxa determined by this procedure are arti-
ficial. It is generally believed [3] that convincing
between-cluster boundaries that could justify such clas-
sifications do not exist. In addition, these groups are
unstable. Any changes in the locus choice or clustering
procedure lead to moving populations from one cluster
to another. Only groups that form, according to the apt
RUSSIAN JOURNAL OF GENETICS Vol. 37 No. 8 2001
 GENETICS AND THE ORIGIN OF HUMAN “RACES”
description of Lewontin [31], “nodes in the taxonomic
space”, are more or less compact and stable. These
groups consist of populations that probably were not
subjected to strong migration pressure. There are no
genetic boundaries between continents and large
regions; those found in gene-geographic studies of
European populations [14–16] are weak and lack taxo-
nomic significance. They are related to geographic and
linguistic barriers hindering gene exchange.
How then are explained substantial phenotypic dif-
ferences underlying race classifications? Answering
this question, one should recall that race identification
was typically based on a small number of visual traits,
e.g., skin, hair, and eye pigmentation, hair structure,
facial features, etc. These traits characterize the body
surface and are subjected to the direct environmental,
and primarily climatic, impact. Hence, it can be
assumed that they were under strong selection, which
ultimately resulted in the formation of the adaptive
types known as races. The same differential selection
eliminated variants deviating from the regional types
and having lower fitness under the conditions of partic-
ular regions. This resulted in homogenization of the
“racial appearance” of the inhabitants of climatic
zones. It is also possible that Darwin’s sexual selection
played a significant role in evolution of the “race-diag-
nostic” traits [3]. The rate of evolution of these traits
was probably high. Because of this, their differences
often do not reflect genetic difference. A similarity in
these traits primarily suggests similar environmental
conditions at the localities of residence.
DNA MARKERS, “RACE” DIFFERENTIATION,
AND THE ORIGIN OF HUMANS
The present structure of genetic variation in humans
can be explained only on the basis of reconstruction and
analysis of evolution of the humankind. Since most of
this variation is inherited from ancestral species, this
analysis should be started at least with H. erectus, who
is the most probable direct predecessor of H. sapiens.
Homo erectus appeared 1.7–1.8 Mya. The oldest
fossils of this species having this age were found in East
and South Africa (see [53] for review). Its brain volume
varied from 800 to 1300 (on average 1100) cc, and the
body proportions, excluding the head, are close to those
of the modern humans. Homo erectus is the first wide-
spread hominid species that colonized both Africa and
Eurasia, starting to disperse throughout the latter conti-
nent not later than 1 Mya. In particular, H. erectus
includes synanthropus or Beijing (Peking) man, and
pithecanthropus, or Java man. The accepted dating of
its latest fossils is 250 000 years. However, fossils of
H. erectus dated at only 27 000–53 000 years ago were
reported to be found in Java [54].
Many paleontologists believe that the first represen-
tatives of our species H. sapiens appeared earlier than
0.5 Mya [55–60]. Their average brain volume, approx-
RUSSIAN JOURNAL OF GENETICS Vol. 37 No. 8
857
imately 1 400 cc, was the same as in the modern
humans but the skull shape was different. Low and long
skulls of this early H. sapiens had such primitive traits
as strong and undivided superciliary arches, thick
bones, and somewhat prominent, “apelike” face. This
type of human was termed archaic H. sapiens (some
authors recognize it as a separate species, H. heidelber-
gensis). The place and time of its origin are still unclear.
Cavally-Sforza et al. [3]conjecture that it originated in
Eastern Europe or West Asia and then migrated to other
regions of Asia, Europe, and Africa suitable for living.
Paleoanthropologists also assume a European origin of
H. sapiens [59]. However, no direct evidence in favor of
this hypothesis was presented. An assumption was put
forward [59] that the transformation of erectus into
sapiens was triggered by cooling of European climate
during the Pleistocene glaciation. The severe climatic
conditions in fact could accelerate evolution of our
ancient ancestors via increasing selection pressure. The
most ancient remains of sapiens were found in Africa
and Europe and the more recent ones, in Asia (see [53]).
For a long period of time, archaic sapiens was contem-
poraneous to erectus.
About 200 000 years ago, local types of H. sapiens
appeared on these continents. The most known of these
is the European type, or the Neanderthal man. It is gen-
erally assigned to a separate subspecies, H. sapiens
neanderthalensis. The set of traits characteristic of this
subspecies was completely formed about 150 000 years
ago. The evolution of its predecessors apparently took
place in Europe where some intermediate (pre-Nean-
derthal) forms linking Neanderthals to the earliest
H. sapiens were discovered. About 120 000 years ago
during the interglacial period, Neanderthals colonized
Middle East and Central Asia. The physical features of
Neanderthals are related to their adaptation to the harsh
climate of the glacial period. During glaciation that
began about 110 000 years ago, the cold northern
regions of Western and Central Europe were populated
by more typical (“classical”) Neanderthals than the
southern and eastern European regions. The average
brain volume of the subspecies was 1500 cc. Its mem-
bers had elongated, rather low skulls with the promi-
nent occupital part (bone “chignon”), large faces, teeth,
and superciliary arches. Neanderthals are thought to die
out about 30 000–35 000 years ago [61] although the
recent evidence indicates that the most recent of them
existed 28 000–29 000 years ago.
The East Asian and African Neanderthal contempo-
raries were markedly different from it and from one
another. The late African archaic H. sapiens (the so-
called Rodesian man) were most similar to the modern
humans. Most paleoanthropologists believe that exactly
this form was the predecessor of our subspecies,
H. sapiens sapiens, which originated, as judged by
osseous remains, about 100 000 years ago. Originally it
inhabited only Africa and Middle East and then dis-
persed on the global scale.
2001
858
The problem of the place of origin of the anatomi-
cally modern human has long been (and still is) under
dispute4. There are two alternative hypothesis to this
effect. According to the first of them, all modern
humans share common origin; according to the second,
different races have different origins [56, 62–71]. The first
hypothesis, which was formerly known as the monocen-
trism theory, suggests that this subspecies appeared on one
continent and then replaced other H. sapiens populations
via migration. At present, this assumption is reduced to
the African origin of modern humans. The second
hypothesis (polycentrism theory) is based on the con-
cept of parallel continuous evolution of local continen-
tal variants of H. erectus that ultimately led, via the
archaic H. sapiens, to the appearance of H. s. sapiens
races. According to the modern version of this hypoth-
esis (currently known as the multiregional model or the
regional continuity hypothesis), the ancestral popula-
tions constituted one population system united by gene
exchange. At present, most paleoanthropologists are
inclined to accept the first hypothesis. However, they
could not adequately resolve this discussion that lasted
five decades. Their opponents persisted until advances
in molecular genetics brought about a radical turn in the
discussion.
An early and most known genetic evidence for the
African origin of the anatomically modern human was
obtained in comparative studies of mitochondrial DNA
(mtDNA) of different races. It is based on the “mito-
chondrial Eve” hypothesis advanced by Wilson and
colleagues in the late 1980s [73–77].
MtDNA is characterized by maternal inheritance.
This small (16 569 bp), circular molecule is very help-
ful in studies on reconstruction of evolutionary history
of populations and closely related species. It evolves
five to ten times faster than nuclear DNA and appar-
ently lack recombination although the latter statement
was recently questioned (see [78–82] and a substanti-
ated answer to these criticisms [83]). Mitochondria of
each particular individual almost always (except
extremely rare cases) contain completely identical mol-
ecules of mtDNA.
Studies based on restriction analysis and later DNA
sequencing have demonstrated that all mtDNA variants
detected descended from a single ancestral molecule
and hence from one female ancestor who lived before
the division of the humankind into major “races.” The
maternal transmission facilitates random fixation of
variants. Each of the variants, like any other polymor-
phic marker, can be eventually lost. The mean time of
such loss is directly proportional to the effective popu-
lation size of the species Ne, which depends on the
number and population history of the species and on
other factors. All other conditions being equal, the
smaller the species number, the lower Ne and the faster
the mtDNA variation decrease. In evolution, the
4 The state of this problem in the “pre-molecular era” is discussed
in detail in Roginsky [72].
TETUSHKIN
humankind has lost all mtDNA lineages except the one
existing today. This retained variant gave rise to a mul-
titude of others via mutation.
The female ancestor whose mitochondria were
inherited by the modern humans was dubbed “mito-
chondrial Eve.” She is the last common ancestor trans-
mitting her mtDNA to us. In this connection, the above
studies were mistakenly interpreted to mean that all
humankind descended from one female ancestor, a
counterpart of the biblical Eve. However, authorities
understand that the contribution of mitochondrial Eve
to the nuclear gene pool of the future humankind was
approximately the same as that of thousands of her con-
temporaries. This does not pertain to mtDNA: “Eve’s”
variant persisted whereas all other variants were elimi-
nated via random processes. A major achievement of
the molecular geneticists studying this problem was
establishing when and where this woman existed.
The residence of mitochondrial Eve was established
by analyses of (1) mtDNA genealogy and (2) variation
of mtDNA in various “races.” The genealogical tree has
two branches stemming from mitochondrial Eve. One
of the branches has only African nucleotide sequences
whereas the other, both African and non-African ones.
The simplest assumption is that the first, most ancient
tree branching (as all others, caused by a mutation) split
Africans into two groups. The ancestors that founded
one of these major branches produced descendants
spreading to other continents. Thus, the mitochondrial
ancestor most probably lived in Africa and is hence
referred to as “African Eve.”
However, this conclusion was not universally
accepted. The criticisms of its opponents (see, e.g.,
[84–86]) are related to the methodological aspects of
the relevant studies. The genealogical trees were con-
structed using the method of maximum parsimony,
which is often used in such studies. The essence of this
method is choosing out of numerous possible phyloge-
netic schemes the one that requires the fewest muta-
tions for explanation of the existing differences. If as in
the discussed case, the number of species or popula-
tions analyzed is high, it is very difficult to find the
“best,” i.e., most parsimonious, of them. The existing
computer software does not guarantee this solution.
The critics have found more than 10 000 trees that were
more parsimonious than the original one. They argue
that most of these trees fail to support an African origin
of mitochondrial Eve.
The second proof of an African origin of “Eve” is
not related to the first one. It is not connected with phy-
logenetic trees but is based on comparing mtDNA vari-
ability in populations of different races. As a measure of
this variability, intrapopulation divergence of
sequences (the number of mutations determining dif-
ferences between mtDNA types in individuals from the
same population) was used. The sequence diversity is
positively associated with the age of the population
RUSSIAN JOURNAL OF GENETICS Vol. 37 No. 8 2001
 GENETICS AND THE ORIGIN OF HUMAN “RACES”
examined. MtDNA of Africans exhibited the highest
heterogeneity. Hence, mitochondrial Eve was African.
The age of the mitochondrial ancestor was deter-
mined using molecular clock calibrated on the basis of
archeological dating of the time of peopling New
Guinea, Australia, and North America. Also the rates of
nucleotide sequence evolution obtained in large-scale
comparative studies of vertebrate mtDNA were taken
into account. The authors concluded that the best esti-
mate of the mean DNA divergence rate in human pop-
ulations is 3% in million years and the values of these rates
vary from 2 to 4% per million years. Since the divergence
depth of the two major mtDNA branches is about 0.6%,
the probably age of the ancestor is 200 000 years with the
lower and upper limits of approximately 140 000 and
290 000 years, respectively. Thus, mitochondrial Eve
most probably belonged to archaic H. sapiens and
existed before the modern humans.
In the 1990s, a team of Japanese researchers [87]
sequenced the mitochondrial genome of humans (Afri-
cans, Europeans, Japanese), chimpanzee, and gorilla,
which convincingly confirmed an African origin of
human mtDNA. Their estimate of the age of our last
common mtDNA ancestor corresponds to the lower
limit of the value given above (see also [88]).
Thus, our mitochondrial genome is of an African
origin. Can it be inferred from this fact that Africa is the
sole place of origin of the present-day humans? The
advocates of the multiregional hypothesis give a nega-
tive answer to this question referring to the fact that
phylogenetic gene trees of species or populations often
do not coincide with the corresponding evolutionary
trees. However, if H. s. sapiens evolved independently
on different continents, its local populations must have
inherited mtDNA from regional variants of H. erectus,
but the latter species, as noted above, appeared in Eur-
asia not later than 1 Mya. Therefore, the divergence of
the Eurasian and African mtDNA types must have
occurred during at least 1 million years. No mtDNA
variants whose differences would correspond to this
dates have been found yet although the samples of non-
Africans studied were quite large. It seems likely that
such mtDNAs do not exist, which contradicts the
hypothesis of multiregional evolution.
The hypothesis of a recent African origin of the
modern humankind is also called the recent (or global)
replacement model. The attribute “recent” is not redun-
dant since the both alternative models are based on the
assumption of an African origin. However, according to
the multiregional model, our ancestors migrated out of
Africa very long ago (1–2 Mya), and these migrants
were H. erectus rather than H. sapiens.
The replacement of aboriginal archaic populations
of Asia and Europe by African migrants is confirmed by
evidence from various genetic systems. For instance,
molecular clock based on “classical” genetic markers
(i.e., polymorphic proteins and blood groups) shows
that Africans and non-Africans diverged 100 000–
RUSSIAN JOURNAL OF GENETICS Vol. 37 No. 8
859
110 000 years ago, whereas Europeans and Asians
diverged 40 000–50 000 years ago [3, 32, 33]. The
depth of the earliest splitting of the arising humankind
corresponds to the paleontological dating of the appear-
ance of our subspecies (100 000–130 000 years ago).
The similar, in view of the approximate nature of such
estimates, dating of the deepest divergence in the
human history was obtained from data on polymor-
phism of autosomal microsatellites (DNA regions con-
sisting of short tandem repeats whose length typically
does not exceed 2 to 5 bp). This estimate is 156 000 years
with the confidence interval of 95 000–290 000 years
[89–91]. Even the upper limit of this estimate is sub-
stantially lower than the values admissible in the multi-
regional model.
The assumption on recent African roots of the mod-
ern humans was confirmed by evidence pertaining to
most DNA markers, autosomal, Y-chromosomal, and
mitochondrial (see [96-92] for review). The data on
geographical variation of the CD4 locus located on
chromosome 12 permitted to develop a detailed sce-
nario of migration out of Africa [97]. This marker con-
sists of two parts, conserved and variable, which makes
it very informative. The first region, an Alu sequence, is
represented by two variants, short and complete. The
second region includes short tandem repeats and has
12 variants. Almost all of the marker forms were found
south of Sahara, some of them, in Ethiopia and Somali,
and only one, outside Africa. This shows that spreading
of people that was terminated by an exit from Africa
included two stages. At the first stage, during which a
considerable part of the variation was lost, the popula-
tion that colonized northeastern Africa stemmed out. At
the second stage, a small population separated that had
only one type of the marker (with the incomplete Alu);
this population migrated from Africa. The migration
route was established in more detail in another impor-
tant study [98]. The standing hypotheses propose two
routes: through Isthmus of Suez and via a land bridge
on the place of Bab el Mandeb (see [99]). This study of
geographical variation of mitochondrial haplogroup M
variants showed that the initial dispersal most probably
followed only the second route: through the territory of the
present-day Ethiopia and further along the southern border
of Asia. Certain variants are absent in Lebanon but occurs
at high frequencies in India, South Arabia, and Ethiopia. It
was shown that the Asian haplogroup M separated from
the east African one earlier than 50 000 years ago. Thus,
almost all, with a very few exceptions [100, 101], non-
African markers originated from African ones. The
gene pool of humans living outside Africa is by origin
a sample from the African gene pool.
Impressive evidence against the multiregional
hypothesis was obtained by comparing mtDNA of
Neanderthals and modern humans [102–104]. MtDNA
was isolated from a shoulder bone of a fossil specimen
from the Neander Valley [102, 103] in the vicinity of
Dusseldorf and from a rib of the Neanderthal child
found in the Mezmaiskaya Cave (North Caucasus)
2001
860 TETUSHKIN
[104]. The differences between the examined hyper-
variable mtDNA regions of Neanderthals and modern
humans were almost three times greater that the corre-
sponding differences between the most diverged mtDNAs
of the modern humans. Hence, the ancestral mitochon-
drial genome shared by these subspecies (or, as some
authors maintain, separate species) existed about
500 000 ago. No regional continuity was found. Judg-
ing by the sequences examined, the present-day Euro-
peans differ from Neanderthals no less than inhabitants
of other continents. Apparently, Neanderthals did not
make contribution to the modern human mitochondrial
gene pool.
Of course, these results do not completely exclude
Neanderthals from participating in our genetic history.
The samples of three (with [105]) individuals is too
small to provide grounds for definite conclusions.
Moreover, in the case of possible outbreeding, the
Neanderthal mtDNA could be eliminated via genetic
drift during this long time interval. For instance, if the
contribution of Neanderthals to the present-day gene
pool was 25%, the probability of losing their mtDNA
would be higher than 50%. Mathematical models based
on the results of the above studies shows that the com-
pletely random crossing of H. s. sapiens and H. s. nean-
derthalensis is impossible but does not refute suggested
partial admixture [106, 107].
The current interest to the interbreeding of H. sapi-
ens forms was heated by a study [108] that reported
finding in Portugal a skeleton of a four-year old child of
a contemporary type presumably possessing a number
of Neanderthal traits. The age of this Gravettian (by
culture) fossils is 24 500 years. Since Neanderthals
were extinct more than 30 000 years ago, this finding
means that some of their traits persisted for more than
200 generations, which is dubious. During this time,
quantitative genes of Neanderthals must have “dis-
solved” in the immigrant gene pool. The interpretation
of the found remains as possessing a mosaic of the
modern and archaic characteristics was refuted by other
paleoanthropologists [109]. They attributed the skele-
ton in question to a common, somewhat sturdy Gravet-
tian child having nothing in common with Neander-
thals.
A Neanderthal admixture in fossil or existing repre-
sentatives of H. s. sapiens can be detected only via phy-
logenetic analysis of DNA sequences. Some authors
recommend for such analysis mitochondrial or Y chro-
mosomal markers [96] as more informative than con-
served autosomal ones. However, mathematical models
showed that unlinked neutral nuclear loci are more
expedient for this purpose. Estimation of human popu-
lation variability at 50–100 of such loci can solve the
question of Neanderthal or other archaic contribution to
the modern human gene pool [107]. However, if the
archaic admixture was small, it cannot be detected even
using a multitude of loci.
RUSSIAN JOURNAL OF GENETICS
Paleodemography also provides evidence in favor of
the replacement hypothesis. Data from various genetic
systems (autosomal, Y-chromosomal, and mitochon-
drial) demonstrate that in the remote past, the H. s. sapi-
ens population was small for a long period of time,
although this long “bottleneck” never became very nar-
row. Studies of DNA polymorphism indicate that the
effective size of the early human population was about
10 000 ([110, 111] and others). A population with such
low, almost critical, Ne could not inhabit three conti-
nents at the same time maintaining genetic integrity
through migration. The conclusion on the small size of
this ancestor population disagrees with the multire-
gional hypothesis. The low number of our ancestors
prior to their global expansion probably accounts for
the relatively low (compared with our closest relative,
chimpanzee [112, 113]) genetic diversity of modern
humans (based on DNA rather than proteins [114]).
The replacement hypothesis convincingly explains
the structure and amount of genetic variation of the
humankind, whereas the high proportion of intragroup
genetic variation is explained by the recent origin of
humans: the time of its existence was not sufficient for
genetic drift to significantly reduce this variability. The
same reason applies to the low intergroup variation: the
populations merely did not have time to differentiate. In
addition, their explosive growth rapidly eliminated iso-
lation by distance that hindered gene exchange.
GENETICS AND EVOLUTION
OF “RACE-DIAGNOSTIC” TRAITS
Why then people from different regions that repre-
sent populations with complex genetic history have con-
sistently distinctive traits? Above, I briefly answered this
question associating these traits with the impact of envi-
ronmental factors. However, this strong selection
response assumed in this case requires explanation based
on genetics of these “race-diagnostic” characteristics.
Until recently, the genetic basis of the “interracial”
differences was virtually unknown. The first results
shedding light on the mechanisms of underlying
genetic control of the main “race-diagnostic” trait—
skin and hair pigmentation—have appeared only very
recently ([115–117] and others). The greatest part of
variation in skin and hair coloration was shown to be
determined by alleles of only one gene encoding mel-
anocortin 1 receptor (MC1R). The critical influence of
this locus is explained by the fact that MC1R acts as the
key mediator in the melanocite production of two major
melanin pigments, eumelanin (brown and black mela-
nins) and phaeomelanin (red and yellow melanins).
Variation in relative amounts of eumelanin and phae-
omelanin underlie all skin and hair color spectrum in
mammals, including humans.
What is the mechanism of regulation of the amount,
proportions, and distribution of different melanins? The
answer to this question was obtained largely from stud-
Vol. 37 No. 8 2001
 GENETICS AND THE ORIGIN OF HUMAN “RACES”
ies of melanogenesis genetics in the mouse. In this
organism, this process is controlled primarily by two
genes: extension and agouti. The first gene is expressed
in melanocites and codes for MC1R. Upon binding the
α-melanocite-stimulating hormone or adrenocortico-
tropic hormone, this receptor affects adenilylcyclase
and increases cellular concentration of cAMP. This
leads to the increased production of eumelanin but not
phaeomelanin. The product of the second gene, agouti,
which is expressed outside melanocites, is antagonistic
to the α-melanocite-stimulating hormone. However,
the role of its already identified and cloned human
homolog is still unclear.
In humans as in mice, mutations of the MC1R gene
(homologous to extension) reducing its function
increase the phaeomelanin /eumelanin ration and lead
to lighter skin and hair coloration. These gene variants
occur at highest (above 80%) frequency in people hav-
ing red hair and/or light, slightly tanned skin. In brown-
haired and black-haired people, the frequency of these
mutations is below 20%, and in individuals with good
ability to acquire suntan it is as low as 4%. In Africans,
mutant alleles of MC1R were not found. At the same
time, the set of common MC1R amino-acid substitu-
tions in Asians is different from that in Europeans.
Interestingly, such mutants have yellow or red coat
color in all examined animals (mice, cattle, foxes, and
horses).
Interspecific comparisons of humans and anthro-
poid apes demonstrated that MC1R is the most rapidly
evolving member of the protein subfamily of melano-
cortin receptors. In human populations, nucleotide
diversity of MC1R was shown to be several times as
high as the corresponding mean estimated for the total
genome. The combination of the high variability and
the prevalence of synonymous substitutions over the
nonsynonymous ones favors the supposition that
MC1R is subject to strong diversifying selection.
What are the mechanisms of this selection? In the
majority of the proposed hypotheses, the intensity of
melanin pigmentation is related to solar illumination,
specifically its UV component. It is suggested that dark
skin color inherent to indigenous residents of tropical
regions, protects its owners from the adverse conse-
quences of UV irradiation, i.e., burning, skin cancer,
photolysis, etc. The light-colored skin of inhabitants of
higher latitude regions is regarded as adaptation pro-
moting UV-induced biosynthesis of vitamin D3 upon
weak UV irradiation. It is thought that eumelanin func-
tions as an optic philter whereas phaeomelanin is
responsible for UV-induced skin damage including
melanoma (due to its ability to generate UV-radiation-
induced free radicals).
Recently, Yablonski and Chaplin published a com-
prehensive review paper [118] presenting new evidence
in favor of the still-disputed idea of the adaptive signif-
icance of skin color variation. These authors for the first
time ever quantitatively tested the hypothesis on asso-
RUSSIAN JOURNAL OF GENETICS Vol. 37 No. 8
861
ciation of this characteristic with parameters of UV
light. Skin color was shown to strongly correlate with
both geographic latitude and the latitude-dependent UV
radiation level. The results obtained indicate that
human skin coloration is highly adaptive.
The authors demonstrated that the primary selective
agent controlling skin pigmentation of residents of low-
latitude regions acts to reduce photolysis of folate, a
derivative of folic acid necessary for nucleotide biosyn-
thesis and thus that of DNA. Folate deficiency leads to
a number of anomalies in the embryo including various
defects of the nervous tube. This is explained by the
participation of this compound in purine and pyrimi-
dine biosynthesis. Due to UV-rays, natural sunlight
decreases the level of folates in fair-skinned women,
which reduces the probability of embryonic survival.
(As shown for chronic alcoholics, ethanol has the same
effect). In addition, experiments on animals have
shown that folate deficiency arrests spermatogenesis
and results in male sterility. Because of this, the content
of melanin photoprotectors directly affects individual
reproductive success under conditions of high UV
exposure. Another significant contribution of melaniza-
tion to enhancing individual fitness is connected to the
protection of sweat glands from UV damage, which can
cause impaired thermal regulation.
However, in high latitudes, especially in regions sit-
uated north of 40° N, dark skin is not adaptive. The
dense melanin philter stops UV photons necessary for
the formation of vitamin D3, the deficit of which leads
to skeletal defects. Consequently, in these latitudes and
upon low UV exposure, high fitness requires skin
depigmentation rather than melanization. In their com-
prehensive monograph, Vogel and Motulsky [10] state
that ancient humans were dark-skinned because most
contemporary primates are characterized by dark pig-
mentation. However, this thesis is questionable. Pri-
mates do not have dark pigmentation. Most of them,
like our closest phylogenetic relative, the chimpanzee,
possess white or light-colored skin covered with dark
hair (see [118] for references). Noticeable pigmenta-
tion is observed only on hairless parts, such as the face,
eyelids, lips, etc. Hence, it may well be that the imme-
diate hominid predecessors also were fair-skinned.
When the hair of our ancestors thinned (which was
caused by specific reasons [118]), their skin darkened
as they lived in the tropical regions. According to
Yablonski and Chaplin [118], the intensity of skin pig-
mentation is labile and changed many times during
human evolution. Note that this statement is in conflict
with classical estimations by Stern (cited in [41]),
according to which the skin-color differences between
Caucasoids and Negroids are determined by fixation of
different alleles of four to six loci. Mathematical mod-
els developed by Nei [41] showed that in this case, the
formation of the observed differences would require at
least 100 000 years of divergent evolution. This time is
comparable with the age of the modern humankind. At
the same time, the above statement is in good agree-
2001
862
ment with the evidence on MC1R gene variability. In
view of these results, one cannot but agree that “skin
color is not related to phylogenetic relationships
between groups of humans” [118].
It is very likely that other traits used for “race” identifi-
cation are also under relatively simple genetic and physio-
logical control. Hence, all these “race-taxonomic” charac-
ters apparently cannot be used for constructing phyloge-
netic classifications of human populations.
DO HUMAN RACES EXIST?
Note the point important for our discussion. Among
other things, a race is a taxonomic category occupying
a certain position in the hierarchic system of interspe-
cies categories (species–subspecies–race–tribe), which
was proposed by Semenov-Tyan’-Shanskii in 1910
(see [119]). Race is a general biological concept used
not only in anthropology but also in zoology.
However, in practice the term race is often used syn-
onymously with subspecies. According to Mayr [120],
“subspecies may be in many cases called ecological and
geographical races.” Dobzhansky [121, 122] inferred
that this applied to human races, which he attributed to
subspecies of the humankind. This author argued that
races, including human ones, are biological reality.
Apparently, the most substantial argument against
reality of human races is the aforementioned conclu-
sion on the absence of genetic boundaries between any
adjacent populations of our species. “Races” merge and
continuously grade into one another, which makes their
separation impossible. In a relatively recent study [48]
dealing with this issue, Barbujani et al. point out that
the intrapopulation variance component was lower than
50% only in one out of 109 microsatellite loci and
restriction sites. The authors urge the remaining advo-
cates of the reality of human races to search for systems
having intercontinental breaks in the distribution of
genetic variation. Reinforcing their point, they remind
that frequencies of human genes almost always and
everywhere form smooth clines. Some discontinuities
exist but the local gradients are so small that can be
detected only by combined analysis of many loci using
complex statistical methods. Moreover, these breaking
lines do not contour large population groups inhabiting
whole continents. On the contrary, these boundaries
divide populations within continents or even countries
and typically coincide with geographic or linguistic
barriers. Genetic enclaves (isolates) are mainly located
on islands.
The opponents of the human race concept agree that
comparisons by sufficiently large number of loci may
be able to reveal differences between any two popula-
tions and even shed light on geographic origins of each
particular individual [123]. However, this does not
mean that the humankind can be divided into few dis-
tinct racial or continental groups. Analysis of genetic
data does not confirm the existence of such groups.
TETUSHKIN
In justification of their view on the absence of
human races, Barbujani et al. [48] cite Mayr and argue
that a species can be divided into races if represented by
“essentially discontinuous set of individuals.” How-
ever, in his cited fundamental work, Mayr [124] states
that a polytypic species (and H. sapiens is considered
polytypic since it is classified into races—E.T.) is char-
acterized by actual or potential genetic continuity of
allopatric populations (p. 274). In Mayr’s opinion, the
better known geographic variability of a species, the
more difficult to determine the (often arbitrarily set)
boundaries of its subspecies. The traditional concept of
human races is consistent with Mayr’s views. This
author distinguishes primary and secondary intergrada-
tion of subspecies and/or races. Primary intergradation
zones are gradually developed in the process of con-
stant contact between all participating populations.
Secondary intergradation zones result from contact of
once separated and significantly diverged populations.
The human history apparently involved intergradation
of both types but the primary intergradation prevailed,
which is confirmed by our knowledge of human dis-
persal. In this case, geographical variation of specific
traits must be gradual and continuous.
The weakness of the above argument against the
existence of human races is evident even without invok-
ing hybrid zones. Reasoning of the advocates of this
view implies that the absence of transient intermediate
populations would mean the existence of races. Their
existence depends on the width of the intergradation
zones rather than the amount of differentiation of pop-
ulation groups. This flawed logic was evident to
Dobzhansky even in the early 1960s [121]. (It turns out
that the existence of human races was debated involv-
ing a similar rationale long before the appearance of
Lewontin’s key study!). As Dobzhansky said, “this is
about as logical as it would be to argue that youth is not
different from old age because the gradient between the
two is almost completely smooth.”
Another argument used to refute the human race
concept seems more substantial. It concerns the rank of
these intraspecific taxa. Since any group of related pop-
ulations cannot be regarded as a race, a question logi-
cally arises whether “races” of H. sapiens” correspond
to races (or subspecies) of other species?
To answer this question, we should compare the
structure of genetic variation in different species
including our own. This structure can be inferred from
Wright’s statistics FST (fixation index). This parameter
measures subdivision of metapopulations and estimates
the proportion of intergroup variability (variance) in the
total variation of population systems. Its analog for
multiallele loci was denoted GST [125]. The tables of
FST (GST) estimates show that interracial variability of
humans is comparable to interpopulation variability of
other polytypic species [126, 127]. The mean FST of
protein loci is about 20% for vertebrates [127] (over
24% for mammals) whereas for humans, as noted
RUSSIAN JOURNAL OF GENETICS Vol. 37 No. 8 2001
 GENETICS AND THE ORIGIN OF HUMAN “RACES”
above, this estimate is approximately 15%; the inter-
continental (interracial proper) component accounts for
7 to 10% of the variation. Thus, intergroup variation of
our species is lower than the mean variation in animals
(including invertebrates) but not essentially so.
However, in spite of these close differentiation lev-
els of intraspecific geographical subdivisions, the struc-
ture of H. sapiens is more uniform than that of other
polytypic species [128]. Genetic differences (including
those in quantitative traits) among subspecies and races
are generally more distinct and marked than differences
between human “races.” For instance, no cases of fixa-
tion of alternative alleles, which are rather usual for ani-
mal subspecies and races, were recorded for humans.
(Allele Fy – of the Duffy system, which provides stabil-
ity towards an exciter of malaria Plasmodium vivax, is
close to fixation in Africa, but this and similar cases are
readily explained by constant selection pressure).
Homogeneity of the humankind is also indicated by the
comparable levels of genetic diversity in populations of
extremely different sizes: these levels are high in both
populations of large countries and very small villages
(see above). This is a consequence of our recent origin
from the common ancestral population. The greatest
part of genetic variation in present-day humans is
inherited from this population.
Thus, as we see, doubts in reality of human races are
rather well grounded. At the same time, the reasons
invoked in this connection are clearly not sufficient for
definitely rejection the possibility of their existence.
The boundaries between subspecies and races are usu-
ally conventional and sometimes, in Mayr’s words,
“absolutely arbitrary” [124]. The same applies to taxo-
nomic ranks. The necessary level of difference between
intraspecific taxonomic units can be established only
on consensus between taxonomists [124]. In addition,
classification of divisions of related species should be
taken into account: their subspecies and races should
correspond to each other, although the differentiation
levels of these taxa are very different in different species,
which considerably hinders such comparisons. The sub-
species of the closest relatives of humans (chimpanzee,
gorilla, orangutan) diverged more than 1 Mya [129, 130].
Their age is far greater than that of our species.
In my view, the problem of reality of human races is
still unresolved. The molecular studies proved that the
available classifications are erroneous. However, this
does not mean that the humankind cannot be divided
into races on the basis of another classification princi-
ple. Why not identify races with relatively conventional
boundaries? For instance, humans can be grouped into
two races, conventionally denoted Africans and non-
Africans (which can be regarded as first-order races),
with a diffuse arbitrary border along southern Sahara
frontiers. Race is not a purely cultural category. The
social phenomenon of it reflects biological reality. Due
to genomics, we are coming close to understanding the
genetic basis of this phenomenon.
RUSSIAN JOURNAL OF GENETICS Vol. 37 No. 8
863
EPILOGUE
December 15, 2000, an Interdisciplinary Workshop
“Molecular Genetics and Anthropology: Union or Con-
frontation?” was held in the Research Institute and
Museum of Anthropology. The Workshop was orga-
nized by the Russian Division of European Anthropo-
logical Association and Moscow Association of Genet-
icists and Breeders; Academician of the Russian Acad-
emy of Sciences T.I. Alekseeva and Corresponding
Member of the Russian Academy of Medical Sciences
E.K. Ginter were chairpersons. Geneticists and anthro-
pologists from many scientific institutions participated
in its work. The invited speakers, in particular I.V. Per-
evozchikov and E.V. Balanovskaya, discussed the issue
of reality of human races but regrettably did not present
its conceptual analysis. The present paper is my contri-
bution to the discussion started at the workshop.
ACKNOWLEDGMENTS
I am grateful to A.A. Prozorov for his proposal to
present this work as a seminar at the Vavilov Institute of
General Genetics whose chairperson he was. Special
thanks are due to O.L. Kurbatova and E.V. Bala-
novskaya who invited me to the workshop, participat-
ing in which motivated my work on the paper.
REFERENCES
1. Tishkov, V.A., Anthropology of Russian Transforma-
tions, EO, 2000, no. 1, pp. 3–19.
2. Korotkevich, G.V., Race: A Myth or a Real Phenome-
non, Priroda, 1998, no. 12, pp. 102–103.
3. Cavalli-Sforza, L.L., Menozzi, P., and Piazza, A., The
History and Geography of Human Genes, Princeton:
Princeton Univ. Press, 1994.
4. Bunak, V.V., Rod Homo, ego vozniknovenie i posleduy-
ushchaya evolyutsiya (The Genus Homo, Its Origin and
Further Evolution), Moscow: Nauka, 1980.
5. Cheboksarov, N.N. and Cheboksarova, I.A., Narody,
rasy, kul’tury (Ethnoses, Races, Cultures), Moscow:
Nauka, 1985.
6. Alekseev, V.P., Stanovlenie chelovechestva (Formation
of Mankind), Moscow: Politizdat, 1984.
7. Tetushkin, E.Ya., Genetic and Morphological Distances
between Great Human Races, Genetika (Moscow),
1989, vol. 25, no. 5, pp. 918–922.
8. Cavalli-Sforza, L.L. and Cavalli-Sforza, F., The Great
Human Diasporas: The History of Diversity and Evolu-
tion, Reading: Addison–Wesley, 1995.
9. Stoneking, M., How It All Happened, Nat. Genet.,
2000, vol. 25, p. 379.
10. Vogel, F. and Motulsky, A.G., Human Genetics: Prob-
lems and Approaches, Berlin: Springer-Verlag, 1986.
11. Bruk, S.I. and Cheboksarov, N.N., Human Races, in
Narody mira (World Ethnoses), Moscow: Sov. Entsik-
lopediya, 1988, pp. 16–25.
2001
864 TETUSHKIN
12. Webster’s Seventh New Collegiate Dictionary, Spring-
field: G&C Merriam, 1969.
13. Ammerman, A.J. and Cavalli-Sforza, L.L., The
Neolithic Transition and the Genetics of Populations in
Europe, Princeton: Princeton Univ. Press, 1984.
14. Derish, P.A. and Sokal, R.R., A Classification of Euro-
pean Populations Based on Gene Frequencies and Cra-
nial Measurements: A Map-Quadrate Approach, Hum.
Biol., 1988, vol. 60, pp. 801–824.
15. Sokal, R.R., Genetic, Geographic, and Linguistic Dis-
tances in Europe, Proc. Natl. Acad. Sci. USA, 1988,
vol. 85, pp. 1722–1726.
16. Sokal, R.R., Oden, N.L., Legendre, P., et al., Genetics
and Language in European Populations, Am. Nat., 1990,
vol. 135, pp. 157–175.
17. Roginskii, Ya.Ya. and Levin, M.G., Antropologiya
(Anthropology), Moscow: Vysshaya Shkola, 1978.
18. Tegako, L.I. and Salivon, I.I., Osnovy sovremennoi
antropologii (Basics of Modern Anthropology), Minsk:
Universitetskoe, 1989.
19. Cavalli-Sforza, L.L. and Edwards, A.W.F., Analysis of
Human Evolution, Genetics Today: Proc. XI Int. Congr.
of Genetics (The Hague, Netherlands, September, 1963),
Oxford: Pergamon, 1964, pp. 923–933.
20. Cavalli-Sforza, L.L., Origin and Differentiation of
Human Races, Proc. R.A.I., 1972, pp. 15–25.
21. Nei, M., Evolution of Human Races at the Gene Level,
Human Genetics, part A: The Unfolding Genome, New
York: Alan Liss, 1982, pp. 167–181.
22. Nei, M. and Roychoudhury, A.K., Genetic Relationship
and Evolution of Human Races, Evol. Biol., New York:
Plenum, 1982, vol. 14, pp. 1–59.
23. Nei, M. and Roychoudhury, A.K., Evolutionary Rela-
tionships of Human Populations on a Global Scale, Mol.
Biol. Evol., 1993, vol. 10, pp. 927–943.
24. Alekseev, V.P., Geografiya chelovecheskikh ras (Geog-
raphy of Human Races), Moscow: Mysl’, 1974.
25. Cavalli-Sforza, L.L., Piazza, A., and Menozzi, P.,
Reconstruction of Human Evolution: Bringing together
Genetic, Archaeological, and Linguistic Data, Proc.
Natl. Acad. Sci. USA, 1988, vol. 85, pp. 6002–6006.
26. Cavalli-Sforza, L.L., Genes, Peoples and Languages,
Proc. Natl. Acad. Sci. USA, 1997, vol. 94, pp. 7719–
7724.
27. Tetushkin, E.Ya., Molecular Paleogenetics of Primates,
Genetika (Moscow), 1997, vol. 33, no. 3, pp. 293–307.
28. Owens, K. and King, M.-C., Genomic Views of Human
History, Science, 1999, vol. 286, pp. 451–453.
29. Boyde, W.C., Genetics and the Human Race, Science,
1963, vol. 140, pp. 1057–1064.
30. Dubinin, N.P., The Race Problem and Modern Genetics,
Usp. Sovrem. Biol., 1972, vol. 73, no. 2, pp. 209–230.
31. Lewontin, R.C., The Apportionment of Human Diver-
sity, Evolutionary Biology, New York: Appleton-Cen-
tury-Grofts, 1972, vol. 6, pp. 381–398.
RUSSIAN JOURNAL OF GENETICS
32. Nei, M. and Roychoudhury, A.K., Gene Differences
between Caucasian, Negro, and Japanese Populations,
Science, 1972, vol. 177, pp. 434–436.
33. Nei, M. and Roychoudhury, A.K., Genic Variation
within and between the Three Major Races of Man,
Caucasoids, Negroids, and Mongoloids, Am. J. Hum.
Genet., 1974, vol. 26, no. 4, pp. 421–443.
34. Mitton, J.B., Genetic Differentiation of Races of Man as
Judged by Single-Locus and Multilocus Analyses, Am.
Nat., 1977, vol. 111, pp. 203–212.
35. Chakraborty, R., Single-Locus and Multilocus Analyses
of Genetic Differentiation of Races of Man: A Critique,
Am. Nat., 1978, vol. 112, pp. 1134–1138.
36. Lewontin, R.C., Single-Locus and Multilocus Measures
of Genetic Distance between Groups, Am. Nat., 1978,
vol. 112, pp. 1138–1139.
37. Mitton, J.B., Measurement of Differentiation: Reply to
Lewontin, Powell, and Taylor, Am. Nat., 1978, vol. 112,
pp. 1142–1144.
38. Powell, J.R. and Taylor, C.E., Are Human Races “Sub-
stantially” Different Genetically?, Am. Nat., 1978,
vol. 112, pp. 1139–1142.
39. Nei, M., Genetic Differences between Human Races,
Am. Nat., 1981, vol. 117, pp. 88–89.
40. Neel, J.V., The Major Ethnic Groups: Diversity in the
Midst of Similarity, Am. Nat., 1981, vol. 117, pp. 83–
87.
41. Nei, M., Human Evolution at the Molecular Level, Pop-
ulation Genetics and Molecular Evolution, Tokyo:
Japan Sci. Soc., 1985, pp. 41–64.
42. Nei, M. and Livshits, G., Evolutionary Relationships of
Europeans, Asians, and Africans at the Molecular
Level, Population Biology of Genes and Molecules,
Tokyo: Baifukan, 1990, pp. 251–265.
43. Nei, M. and Livshits, G., Genetic Relationships of
Europeans, Asians and Africans and the Origin of Mod-
ern Homo sapiens, Hum. Hered., 1989, vol. 39, pp. 276–
281.
44. Latter, B.D.H., Genetic Differentiation within and
between Populations of the Major Human Subgroups,
Am. Nat., 1980, vol. 116, pp. 220–237.
45. Ryman, N., Chakraborty, R., and Nei, M., Differences
in the Relative Distribution of Human Gene Diversity
between Electrophoretic and Red and White Cell Anti-
gen Loci, Hum. Hered., 1983, vol. 33, pp. 93–102.
46. Balanovskaya, E.V. and Rychkov, Yu.G., Ethnical
Genetics: Ethnogeographical Diversity of the Gene
Pool in World Ethnoses, Genetika (Moscow), 1990,
vol. 26, no. 1, pp. 114–121.
47. Spitsyn, V.A., Biokhimicheskii polimorfizm cheloveka
(Biochemical Polymorphism of Humans), Moscow:
Mosk. Gos. Univ., 1985.
48. Barbujani, G., Magagni, A., Minch, E., and Cavalli-
Sforza, L.L., The Apportionment of Human DNA
Diversity, Proc. Natl. Acad. Sci. USA, 1997, vol. 94,
pp. 4516–4519.
Vol. 37 No. 8 2001
 GENETICS AND THE ORIGIN OF HUMAN “RACES”
49. Przeworski, M., Hudson, R.R., and Rienzo, A.Di.,
Adjusting the Focus on Human Variation, Trends.
Genet., 2000, vol. 16, no. 7, pp. 296–302.
50. Seielstad, M.T., Minch, E., and Cavalli-Sforza, L.L.,
Genetic Evidence for a Higher Female Migration Rate
in Humans, Nat. Genet., 1998, vol. 20, pp. 278–280.
51. Stoneking, M., Women on the Move, Nat. Genet., 1998,
vol. 20, pp. 219–220.
52. Excoffier, L., Smouse, P.E., and Quattro, J.M., Analysis
of Molecular Variance Inferred from Metric Distances
among DNA Haplotypes: Application to Human Mito-
chondrial DNA Data, Genetics, 1992, vol. 131, pp. 479–
491.
53. Tetushkin, E.Ya., Chronology of the Human Evolution-
ary History, Usp. Sovrem. Biol., 2000, vol. 120, no. 3,
pp. 227–239.
54. Swisher, C.C. III, Rink, W.J., Anton, S.C., et al., Latest
Homo erectus of Java: Potential Contemporaneity with
Homo sapiens in Southeast Asia, Science, 1996, vol. 274,
pp. 1870–1874.
55. Stringer, C., Human Evolution and Biological Adapta-
tion in the Pleistocene, in Hominid Evolution, London:
Academic, 1984, pp. 55–83.
56. Smith, F.H., Continuity and Change in the Origin of
Modern Homo sapiens, Z. Morph. Anthrop., 1985,
vol. 75, pp. 197–222.
57. Andrews, P., Fossil Evidence on Human Origins and
Dispersal, Cold Spring Harbor Symp. Quant. Biol.,
1986, vol. LI, pp. 419–428.
58. Simons, E.L., Human Origins, Science, 1989, vol. 245,
pp. 1343–1350.
59. Delson, E., Fossil Human, New York: McGraw–Hill,
1997, vol. 7, pp. 446–458.
60. Wood, B. and Collard, M., The Human Genus, Science,
1999, vol. 284, pp. 65–71.
61. Stringer, C.B. and Burleigh, R., The Neanderthal Prob-
lem and the Prospects for Direct Dating of Neanderthal
Remains, Bull. Br. Mus. Nat. Hist. (Geol.), 1981,
vol. 35, pp. 225–241.
62. Zubov, A.A., Questionable Issues in Anthropogenesis,
Chelovek, 1997, no. 1, pp. 5–16.
63. Zubov, A.A., Natural History of the Ancient Mankind,
Priroda, 1998, no. 1, pp. 76–87.
64. Stringer, C.B., The Origin of Modern Humans, V Mire
Nauki, 1991, no. 2, pp. 54–61.
65. Stringer, C.B., Homo sapiens: Single or Multiple Ori-
gin?, in Human Origins, Oxford: Clarendon, 1989, pp. 63–
80.
66. Stringer, C.B., The Asian Connection, New Sci., 1990,
vol. 128, pp. 33–37.
67. Stringer, C.B., The Emergence of Modern Humans, Sci.
Am., 1990, vol. 262, pp. 98–104.
68. Stringer, C.B. and Andrews, P., Genetic and Fossil Evi-
dence for the Origin of Modern Humans, Science, 1988,
vol. 239, pp. 1263–1268.
69. Rightmire, G.P., Human Origin(s), TREE, 1996, vol. 11,
pp. 520–521.
RUSSIAN JOURNAL OF GENETICS Vol. 37 No. 8
865
70. Wolpoff, M.H., Human Evolution at the Peripheries:
The Pattern at the Eastern Edge, in Hominid Evolution:
Past, Present and Future, New York: Alan Liss, 1985,
pp. 355–365.
71. Wolpoff, M.H. and Zhi Wu Xin Thorne, A.G., Modern
Homo sapiens Origins: A General Theory of Hominid
Evolution Involving the Fossil Evidence from East
Asia, in The Origins of Modern Humans: A World Sur-
vey of the Fossil Evidence, New York: Alan Liss, 1984,
pp. 411–483.
72. Roginskii, Ya.Y., Problemy antropogeneza (Problems
of Anthropogenesis), Moscow: Vysshaya Shkola, 1977.
73. Cann, R.L., Stoneking, M., and Wilson, A.C., Mito-
chondrial DNA and Human Evolution, Nature, 1987,
vol. 325, pp. 31–36.
74. Wilson, A.C., Stoneking, M., Cann, R.L., et al., Mito-
chondrial Clans and the Age of Our Common Mother,
Human Genetics: Proc. Int. Congr., Berlin: Springer-
Verlag, 1987, pp. 158–164.
75. Stoneking, M. and Cann, R.L., African Origin of
Human Mitochondrial DNA, in The Human Revolu-
tion: Behavioral and Biological Perspectives on the
Origins of Modern Humans, Edinburgh: Edinburgh
Univ. Press, 1989, pp. 17–30.
76. Stoneking, M., Sherry, S.T., Redd, A.J., and Vigilant, L.,
New Approaches to Dating Suggest a Recent Age for
the Human mtDNA Ancestor, Phil. Trans. R. Soc. Lon-
don, B, 1992, vol. 337, pp. 167–175.
77. Stoneking, M., Recent African Origin of Human Mito-
chondrial DNA: Review of the Evidence and Current
Status of the Hypothesis, in Progress in Population
Genetics and Human Evolution, Berlin: Springer-Ver-
lag, 1997, pp. 1–13.
78. Eyre-Walker, A., Smith, N.H., and Smith, M., How
Clonal Are Human Mitochondria?, Proc. R. Soc. Lon-
don, B, 1999, vol. 266, pp. 477–483.
79. Awadalla, P., Eyre-Walker, A., and Smith, J.M., Link-
age Disequilibrium and Recombination in Hominid
Mitochondrial DNA, Science, 1999, vol. 286, pp. 2524–
2525.
80. Arctander, P., Mitochondrial Recombination?, Science,
1999, vol. 284, pp. 2090–2091.
81. Strauss, E., Can Mitochondrial Clocks Keep Time?,
Science, 1999, vol. 283, pp. 1435–1438.
82. Merriweather, D.A. and Kaestle, F.A., Mitochondrial
Recombination?, Science, 1999, vol. 285, p. 837.
83. Macaulay, V., Richards, M., and Sykes, B., Mitochon-
drial DNA Recombination—No Need to Panic, Proc. R.
Soc. London, B, 1999, vol. 266, pp. 2037–2039.
84. Darlu, P. and Tassy, P., Roots (A Comment on the Evo-
lution of Mitochondrial DNA and the Origins of Mod-
ern Humans), Hum. Evol., 1987, vol. 2, pp. 407–412.
85. Excoffier, L. and Langaney, A., Origin and Differentia-
tion of Human Mitochondrial DNA, Am. J. Hum.
Genet., 1989, vol. 44, pp. 73–85.
86. Templeton, A.R., Human Origins and Analysis of Mito-
chondrial DNA Sequences, Science, 1992, vol. 255,
p. 737.
2001
866 TETUSHKIN
87. Horai, S., Hayasaka, K., Kondo, R., et al., Man’s Place
in Hominoidea Revealed by Mitochondrial DNA Gene-
alogy, J. Mol. Evol., 1992, vol. 35, pp. 32–43.
88. Hasegawa, M., Rienzo, A.Di., Kocher, T.D., and Wil-
son, A.C., Toward a More Accurate Time Scale for the
Human Mitochondrial DNA Tree, J. Mol. Evol., 1993,
vol. 37, pp. 347–354.
89. Goldstein, D.B., Linares, A.R., Cavalli-Sforza, L.L.,
and Feldman, M.W., Genetic Absolute Dating Based on
Microsatellites and the Origin of Modern Humans,
Proc. Natl. Acad. Sci. USA, 1995, vol. 92, pp. 6723–
6727.
90. Nei, M., Genetic Support for the Out-of-Africa Theory
of Human Evolution, Proc. Natl. Acad. Sci. USA, 1995,
vol. 92, pp. 6720–6722.
91. Cavalli-Sforza, L.L., The DNA Revolution in Popula-
tion Genetics, Trends. Genet., 1998, vol. 14, no. 2,
pp. 60–65.
92. Jorde, L.B., Watkins, W.S., Bamshad, M.J., et al., The
Distribution of Human Genetic Diversity: A Compari-
son of Mitochondrial, Autosomal, and Y-Chromosome
Data, Am. J. Hum. Genet., 2000, vol. 66, pp. 979–988.
93. Fischman, J., Evidence Mounts for Our African Ori-
gins—An Alternatives, Science, 1996, vol. 271, p. 1364.
94. Disotell, T.R., Human Evolution: Sex-Specific Contri-
bution to Genome Variation, Curr. Biol., 1999, vol. 9,
pp. R29–R31.
95. Disotell, T.R., Human Evolution: Origins of Modern
Humans Still Look Recent, Curr. Biol., 1999, vol. 9,
pp. R647–R650.
96. Pääbo, S., Human Evolution, Trends Genet., 1999,
vol. 15, no. 12, pp. M13–M16.
97. Tishkoff, S.A., Dietzsch, E., Speed, W., et al., Global
Patterns of Linkage Disequilibrium at the CD4 Locus
and Modern Human Origins, Science, 1996, vol. 271,
pp. 1380–1387.
98. Quintana-Murci, L., Semino, O., Bandelt, H.-J., et al.,
Genetic Evidence of an Early Exit of Homo sapiens
from Africa through Eastern Africa, Nat. Genet., 1999,
vol. 23, pp. 437–441.
99. Stringer, C., Coasting out of Africa, Nature, 2000,
vol. 405, pp. 24–27.
100. Harris, E.E. and Hey, J., X Chromosome Evidence for
Ancient Human Histories, Proc. Natl. Acad. Sci. USA,
1999, vol. 96, pp. 3320–3324.
101. Pennisi, E., Genetic Study Shakes up Out-of-Africa
Theory, Science, 1999, vol. 283, p. 1828.
102. Krings, M., Stone, A., Schmitz, R.W., et al., Neander-
thal DNA Sequences and the Origin of Modern
Humans, Cell (Cambridge, Mass.), 1997, vol. 90,
pp. 19–30.
103. Krings, M., Geisert, H., Schmitz, R.W., et al., DNA
Sequence of the Mitochondrial Hypervariable Region II
from the Neanderthal Type Specimen, Proc. Natl. Acad.
Sci. USA, 1999, vol. 96, pp. 5581–5585.
104. Ovchinnikov, I.V., Götherström, A., Romanova, G.P.,
et al., Molecular Analysis of Neanderthal DNA from
RUSSIAN JOURNAL OF GENETICS
the Northern Caucasus, Nature, 2000, vol. 404,
pp. 490–493.
105. Krings, M., Capelli, C., Tshentscher, F., et al., A View
of Neanderthal Genetic Diversity, Nat. Genet., 2000,
vol. 26, pp. 144–146.
106. Nordborg, M., On the Probability of Neanderthal
Ancestry, Am. J. Hum. Genet., 1998, vol. 63, pp. 1237–
1240.
107. Wall, J.D., Detecting Ancient Admixture in Humans
Using Sequence Polymorphism Data, Genetics, 2000,
vol. 154, pp. 1271–1279.
108. Duarte, C., Mauricio, J., Pettitt, P.B., et al., The Early
Upper Paleolithic Human Skeleton from the Abrigo Do
Lagar Velho (Portugal) and Modern Human Emergence
in Iberia, Proc. Natl. Acad. Sci. USA, 1999, vol. 96,
pp. 7604–7609.
109. Tattersall, I. and Schwartz, J.H., Hominids and Hybrids:
The Place of Neanderthals in Human Evolution, Proc.
Natl. Acad. Sci. USA, 1999, vol. 96, pp. 7117–7119.
110. Harpending, H.C., Batzer, M.A., Gurven, M., et al.,
Genetic Traces of Ancient Demography, Proc. Natl.
Acad. Sci. USA, 1998, vol. 95, pp. 1961–1967.
111. Jorde, L.B., Bamshad, M.J., and Rogers, A.R., Using
Mitochondrial and Nuclear DNA Markers to Recon-
struct Human Evolution, BioEssays, 1998, vol. 20,
pp. 126–136.
112. Morin, P.A., Moore, J.J., Chakraborty, R., et al., Kin
Selection, Social Structure, Gene Flow, and the Evolu-
tion of Chimpanzees, Science, 1994, vol. 265, pp. 1193–
1201.
113. Crouau-Roy, B., Service, S., Slatkin, M., and Freimer, N.,
A Fine-Scale Comparison of the Human and Chimpan-
zee Genomes: Linkage, Linkage Disequilibrium and
Sequence Analysis, Hum. Mol. Genet., 1996, vol. 5,
pp. 1131–1137.
114. Takahata, N., Relaxed Natural Selection in Human Pop-
ulations during the Pleistocene, Jpn. J. Genet., 1993,
vol. 68, pp. 539–547.
115. Valverde, P., Healy, E., Jackson, I., et al., Variants of the
Melanocyte-Stimulating Hormone Receptor Gene Are
Associated with Red Hair and Fair Skin in Humans,
Nat. Genet., 1995, vol. 11, pp. 328–330.
116. Schiöth, H.B., Phillips, S.R., Rudzish, R., et al., Loss of
Function Mutations of the Human Melanocortin 1
Receptor Are Common and Are Associated with Red
Hair, Biochem. Biophys. Res. Commun., 1999, vol. 260,
pp. 488–491.
117. Rana, B.K., Hewett-Emmett, D., Jin, L., et al., High
Polymorphism at the Human Melanocortin 1 Receptor
Locus, Genetics, 1999, vol. 151, pp. 1547–1557.
118. Jablonski, N. and Chaplin, G., The Evolution of Human
Skin Coloration, J. Hum. Evol., 2000, vol. 39, pp. 57–
106.
119. Vorontsov, N.N., Razvitie evolyutsionnykh idei v
biologii (Development of Evolutionary Concepts in
Biology), Moscow: UNTs DO Mosk. Gos. Univ., 1999.
120. Mayr, E., Populations, Species, and Evolution, Cam-
bridge (Massachusetts): Harvard Univ. Press, 1970.
Vol. 37 No. 8 2001
 GENETICS AND THE ORIGIN OF HUMAN “RACES”
121. Dobzhansky, T., Genetic Entities in Hominid Evolution,
in Classification and Human Evolution, Chicago:
Aldine, 1963, pp. 347–362.
122. Dobzhansky, T., Evolution of Mankind in the Light of
Population Genetics, Proc. XII Int. Congr. on Genetics,
1969, vol. 3, pp. 281–292.
123. Bowcock, A.M., Ruiz-Linares, A., Tomfohrde, J., et al.,
High Resolution of Human Evolutionary Trees with
Polymorphic Microsatellites, Nature, 1994, vol. 368,
pp. 455–457.
124. Mayr, E., Animal Species and Evolution, Cambridge:
Belknap, 1963.
125. Nei, M., Analysis of Gene Diversity in Subdivided Pop-
ulations, Proc. Natl. Acad. Sci. USA, 1973, vol. 70,
pp. 3321–3323.
RUSSIAN JOURNAL OF GENETICS Vol. 37 No. 8
View publication stats
867
126. Nei, M., Molecular Evolutionary Genetics, New York:
Columbia Univ. Press, 1987.
127. Ward, R.D., Skibinski, D.O.F., and Woodwark, M., Pro-
tein Heterozygosity, Protein Structure, and Taxonomic
Differentiation, Evol. Biol., 1992, vol. 26, pp. 73–159.
128. Barton, N.H., Population Genetics: A New Apportion-
ment of Human Diversity, Curr. Biol., 1997, vol. 7,
pp. R757–R758.
129. Arnason, U., Xu, X., Gullberg, A., and Graur, D., The
“Phoca Standart”: An External Molecular Reference for
Calibrating Recent Evolutionary Divergences, J. Mol.
Evol., 1996, vol. 43, pp. 41–45.
130. Xu, X. and Arnason, U., The Mitochondrial DNA Mol-
ecule of Sumatran Orangutan and a Molecular Proposal
for Two (Bornean and Sumatran) Species of Orangutan,
J. Mol. Evol., 1996, vol. 43, pp. 431–437.
2001