AMERICAN JOURNAL OF HUMAN BIOLOGY 63-1 1 (1994)

Ethnicity and Human Biology WILLIAM S . POLLITZER University of North. Carolina, Chapel Hill, North Curolina 27599

ABSTRACT Stanley Garn’s book, Human Races (1963, captured the essence of the new genetic concept of ethnic groups as populations which share the same gene pool, shaped by natural selection. Since then new genetic traits, especially DNA polymorphisms, have been employed to build upon his vision. They have proven useful in the forensic sciences, medicine, and the human biology of adaptation, migration, evolution, and phylogeny. Some diseases like sickle cell anemia are controlled by single genes concentrated in a large ethnic group; their alleles are valuable for understanding human biology. Other conditions with a marked genetic component, such as diabetes, heart disease, and osteoporosis,predominate in certain ethnic groups. The new molecular biology tends t o support the migration of farmers from west Asia into Europe in the Neolithic, the isolation of the Basques, and the early entrance of people into the New World. Some DNA data also favor the debatable hypothesis of the development of humanity in Africa and its subsequent dispersal. In many cases linguistic distance parallels genetic distance as language tends to influence gene flow. Apolipoproteins, with alleles unique to one major population group, are useful markers of admixture as well as significant for cardiovascular health. The Gullah-speaking Black people of coastal Carolina are distinctive and close to African ancestors in language, culture, and biology, including genetic markers. Modern research confirms that race (by any name) is quantitative rather than qualitative, relative rather than absolute, and dynamic rather than static. Q 1994 Wiley-Liss, Inc. The rise of population genetics provided a stimulus to physical anthropology, gave new insights into humanity’s diversity, and called into question the meaning of “race.” I t became increasingly clear that a discrete human population, occupying any given territory of the globe, whether labelled a race or ethnic group, shares a common gene pool. Moreover, in any given inherited trait, be it skin color o r the frequency of blood Group B, gradients exist between populations. In light of these clines some rejected race altogether. Others saw the utility of recognizing clusters of people of common heritage and distinctive genetic traits. If a giant hand could lift a thousand people from Nigeria, another handful from England, and a third from Japan, no one would have difficulty telling which people came from each country, even though there would be diversity within each group and endless gradations in all traits in individuals along the lines that connect these three centers. Some human biologists focussed on genetics and others

0 1994 Wtley-Ltss, Inc

stressed physiology, but all recognized populations a s groups adapted t o their environment and subject to the pressure of natural selection. Stanley Garn’s book, Human Races (19651,first published three decades ago, captured the message and meaning of the new concept. With his usual skill, he described not only visible differences, but also those hidden in the blood; he explained adaptations to heat and cold; and he recognized the place of disease in characterizing ethnic groups. While he identified several large geographical races and many smaller local races within them, he was careful to avoid drawing rigid boundaries around them. Ladinos and Neo-Hawaiians were just as much entitled to a n identity as older local races. Today we realize t h a t race must be

Received March 23,1992; accepted November 14,1992. Address repiint requests to the above.

4

W.S. POLLITZER

seen as quantitative rather than qualitative, relative rather than absolute, dynamic rather than static. It is more fruitful to investigate ongoing processes than to erect sterile classifications. In the picturesque phrase of Hartl (19851,the human gene pool is composed of a number of puddles. In the three decades since Garn published his volume, myriad new tools have become available to those who would document human biological diversity and ponder its causes. In addition t o morphology, pigmentation, and the well-known blood types and hemoglobin variants, new genetic polymorphisms have been discovered: chromosome structure, red cell enzymes, plasma proteins, complex immunoglobulins, and human lymphocyte antigens (HLA). When Kan and Dozy (1978) described a restriction fragment length polymorphism (RFLP) adjacent to the beta-globin gene in 1978,they opened up a new tool of molecular biology available to those who study human diversity. Restriction enzymes recognize small, specific DNA sequences of the human genome, including noncoding ones overlooked in the search for proteins. The DNA heterozygosity index of 1:270 is ten times higher than that for protein coding regions (Thompson et al., 1991). Thanks t o the polymerase chain reaction (PCR), millions of copies of DNA can be produced. Today more than 2,400 DNA marker loci are known and many have been mapped; they have been studied in Chinese, Pygmies, Melanesians, and American Indians as well as several Caucasian groups (Kidd, 1992). Moreover, the many alleles of the variable number of tandem repeated DNA sequences (VNTR)in the interval between two restriction sites are valuable for linkage and identification. Finally, mitochrondrial DNA (mtDNA), with maternal inheritance, no recombination and geographical heterogeneity, has proven useful for evolutionary studies. DNA can now also be determined from saliva and amplified by PCR (Von Dornum and Ruvolo, 1992). The identification of human variation has been extended backward in time by techniques that detect albumin (Cattaneo et al., 1992), immunoglobulins (Tuross and Owsley, 19921, and mtDNA (Stone, 1992) in ancient bones, The recognition of new molecular polymorphisms in the great apes has further enriched understanding of human phylogeny.

One practical application of the enormous human diversity seen at the molecular level is in the forensic sciences, where controversy over DNA fingerprinting based on VNTRs has spilled over into the pages of the journal Science. Two distinguished population geneticists, Chakraborty and Kidd (19911, support the reliability of such evidence which utilizes allele frequencies from large ethnic groups, while two other distinguished population geneticists, Lewontin and Hartl (19911, citing the variability within such gene pools, deplore the practice. While data from more populations and more loci, and judicious restraint in the interpretation of results are in order, new work (Chakraborty et al., 1991a; Chakraborty and Jin, 1992) appears t o put the proponents of fingerprinting ahcad. The relationship between disease and ethnicity is clear and growing rapidly in importance. Garn (1965) recognized the interaction of G-6-PD deficiency with diet in Mediterranean people with his description of “Favism: When Gene Meets Bean.” Unravelling the complexities of hemoglobin variants underlying sickle cell disease, thalassemias, and allied hemoglobinopathies is a triumph of molecular biology too extensive and well known to recount here. The beta-globin haplotypes, Senegal, Benin, and Bantu, now figure in population dynamics alongside electrophoretic variants (Livingstone, 1989).A host of diseases caused by a single gene are known t o predominate in specific ethnic groups, such as Tay-Sachs in Ashkenazic Jews. HLA antigens with striking disease associations vary in frequency in populations and are medically significant in organ transplantation. One polymorphic system of recent discovery stands out for its clinical significance, for the application of molecular techniques to the study of human physiology, and for its role in understanding population dynamics, the apolipoproteins. Responsible for lipid transport, for the transfer of lipoproteins across membranes, and for an effect on key enzymes involved in lipid metabolism, they are of greatest importance for cardiovascular disease (Ferrell, 1989; Chan and Dressell, 1990). Controlled by a dozen genes, they have been studied in many populations, including American Whites and Blacks, Nigerians (Sepehrnia et al., 19891, Australian Aborigines (Kamboh et al., 1991c),Samoans (Crews et al., 19411,Papua

ETHNlClTY AND HUMAN BIOLOGY

New Guineans (Kamboh et al., 199Oj, and Yanomami Indians (Crews et al., 1992). Of all loci, Apo E exhibits wide polymorphism (Sepehrnia et al., 1989); its alleles, working in a uniform way in different normal populations (Hallman et al., 1991; Kamboh et al., 1992), have great effect on factors influencing cardiovascular health (Kaprio et al., 1991). Apo J and Apo C-11, monomorphic in Whites but polymorphic in Blacks, and certain unique alleles of Apo A-IV (Kamboh et al., 1991a,bj are useful markers in admixture studies. The most recent analysis, employing 15 polymorphic loci with 18 variants unique to Blacks and 5 unique to Caucasians, confirms 25% admixture of White genes in American Blacks of Pittsburgh (Chakraborty et al., 1991b). Many conditions prominent in one race where single genes are not identified have a high hereditary component such as osteoporosis common in elderly White females. Mildred Trotter first established the greater density of bone in Blacks than Whites (Trotter et al., 1959). (Bone mass and growth is another important field to which Garn has made notable contributions.) While the cause of this illness is not fully understood, genetic factors are increasingly identified (Pollitzer and Anderson, 1989); different alleles of alpha 2 HS-glycoproteins affect hormones that influence bone density (Eichner et al., 1990). We now have more than familiarity with ethnicity and disease, with its focus on diabetes (Ferrell, 1992) in American Indians (Young, 1992) and Japanese Americans (Leonetti and Fujimoto, 1992; Newell-Morris et al., 1992), and on obesity in the Navaho (Hall et al., 1992) and Afi-ican Americans, significant for cardiovascular disease (Adams-Campbell, 1992). The physical anthropologist is interested in the processes of evolution that gave rise t o the human species and its differentiation into ethnic groups. Their studies of adaptation have made major contributions to the understanding of the formation of such populations, as molecular biology has helped to chart the course of the migrations, admixture, relationships, and distances among them. Fossil data indicating that human beings arose in Africa and then dispersed found debatable support from at least some molecular studies. Haplotypes of the Apo B locus of chimp, gorilla, and human popula-

5

tions support the African origin of our species and subsequent migrations to Europe and the Pacific (Rapacz et al., 1991). DNA markers in Pygmies, Melanesians, Chinese, and Caucasoids favor a primary split between Eurasia and Africa (Bowcock et al., 1987). Analysis of beta-globin haplotypes from many populations supports this separation and the relatively recent evolution of human races (Long et al., 1990); DNA fragments from both X and Y chromosomes also tend to confirm this distinctiveness of African people (Papiha et al., 1991; Torroni et al., 1990). One tree that grows out of molecular biology indicates that a mixture of modern people of Afi-ica and East Asia gave rise t o West Asians who then replaced Neanderthals in Europe 30-40 thousand years ago; the migration of people from the Near East some 9,000 years ago, fueled by the population explosion of the Neolithic, rather than cultural diffusion, pushed farming into Europe (Ammerman and Cavalli-Sforza, 1984; Bowcock et al., 1991; Sokal et al., 1991). As one person expressed it, farming is in the blood (Jones, 1991). A rash of genetic markers now join archeology, language, and cranial morphology to declare the Basques descendants of Paleolithic and Mesolithic people, with some later admixture with Neolithic migrants (Bertranpetit and Cavalli-Sforza, 1991; Garcia et al., 1990; Aguirre et al., 1991a,b). Recent genetic studies continue t o shed light on the affinities of many other populations. A vast array of alleles characterize Sardinians and distinguish them from Italians (Walter et al., 1991; Floris-Masala et al., 1990; Barbujani and Sokal, 1991a). Algerians are said, from blood types, to be intermediate between sub-Saharan Africans and Caucasoids (Aireche and Benabadji, 1990). Similarly, mtDNA and beta-globin haplotypes as well as classical markers reflect Caucasoid and sub-Saharan components in Ethiopian Jews (Zoossman-Diskin et al., 1991). New Y-linked haplotypes also indicate such an intermediate position for people of India (Lucotte et al., 1990). DNA polymorphisms for the X-linked coagulation factor, Malmo B, show a decline from Europeans through Asians and Africans (Wallmark et al., 1991). We cannot grant human biology a divorce from language and culture. In sub-Saharan Africa language-family relationship, proba-

6

W.S. POLLITZER

114,788 6,676

121,464

Africa Total West Indies Total Grand Total

D

im m 0

3m 400 m 6m Too wnrlrs 2M

K.3

m

mion

SOURCE OF SLAVES IMPORTED INTO SOUTH CAROLINA, 1716-1807

N NAMIBIA

-1

I (ItOCRAFlnCB

phnPl

Fig. 1. Source of slaves imported into South Carolina, 1716-1807. Estimated from advertisements in Charleston newspapers, Naval Office Shipping Lists, and Records of the Public Treasurer of South Carolina.

bly reflecting migrations, is the best predictor of genetic relationship based on Rh and Gm haplotypes (Excoffier et al., 3991). In Italy, linguistic rather than geographical factors parallel gene flow (Barbujani and Sokal, 1991b). Among American Indians of the Southwest mtDNA is closely associated with linguistic groups (Lorenz, 1992). However, data on tribes in the vast Amazon basin show their distinctiveness and a weak correlation of their genetic distance with either linguistic or geographical distance (Salzano et al., 1991). However, the scrutiny of markers in the blood does not mean that older measures of diversity are dead. Quantification of skin pigmentation is still a useful technique; it is closely correlated with ancestry among Mexican Americans (Mitchell et al., 1992). But, judging from skin color in one-half of a dozen endogamous groups of eastern Nepal,

skin reflectance values may be more useful for macro- than for micro-differentiation (Williams-Blangero and Blangero, 1991). Anthropometry remains significant not so much for classification as for understanding adaptation of humanity to climate, especially when used in conjunction with other data. A cluster analysis of cranial variables among Europeans reflects patterns comparable t o those for blood polymorphisms (Harding, 1990). New discoveries push back the time of entrance of humanity into the New World. mtDNA tends to support the greater antiquity of American Indians and the more recent arrival of NaDene and Inuit speakers (Ward et al., 1992). Immunoglobulins of Eskimos and other people of the far north imply that four different migrant groups may have crossed Beringia a t varying times (Schanfield et al., 1990). Craniometric anal-

ETHNlClTY AND HUMAN BIOLOGY

7

Fig. 2. Coastal South Carolina and Georgia. Original map drawn by P.H. Neumann

ysis of Paleoindian remains dating to 10,000 B.P. suggests that populations may have entered this continent before cranial features of modern northern Asians and North American Indians were fully developed (Steele and Powell, 1992). Investigations of growth among populations living in vastly different environ-

ments, such as differential altitudes, are also relevant to ethnic groups and the factors that produce them. Considerable differences in stature at the same age are found in different mountain-dwelling people. Sherpas and Lepchas inhabiting similar altitudes in the Darjeeling hills are anthropometrically closer to each other than Sherpas

8

W.S. POLLITZER

TABLE 1 . African linguistic sources o f Gullah uocabularv 3,595 Personal names

+

25 1 Words, conversation

Laneuaee

No.

%

No.

c/c

Kongo Yoruba Mende Ewe Bambara Twi Vai Hausa Fon Umbundu Other Total

706 775 433 405 323 281 218 251 229 211 1,040 4,872

14.5 15.9 8.9 8.3 6.6 5.8 4.5 5.2 4.7 4.3 21.3 100.0

99 13 31 15 21 14 30 21 10 18 128 400

24.8 3.2 7.8 3.8 5.2 3.5 7.5 5.2 2.5 4.5 32.0 100.0

+

92 Words, stories

No.

%,

3,938 Total

-

No. 805

64

68.8

1

1.1

27

29.0

1 93

1.1

100.0

788 528 420 345 295 275 272 239 229 1,169 5,365

w 15.0 14.7 9.8 7.8 6.4 5.5 5.1 5.1 4.4

4.3 21.9 100.0

'As one word i s often assigned meaning in several languages, the total cnnsiderahly exceeds the numhpr ofwords. (Calculated from Turner, 1949. I

at different altitudes (Gupta et al., 1989; Carolina and Georgia are viewed with most Majumder et al., 1986; Basu, personal com- affection. They are descendants of Africans munication). Current studies on hematocrit brought into Charleston. From 1670 to the (Shen et al., 19921, chest size, and pulmo- end of the legal slave trade in 1808, at least naryfunction (Weitz et al., 1992) in Han and 122,000 were imported, some 39% from Tibetan boys at different ages bear out this Congo-Angola, 20% from Senegambia, a complex genetic-environmental interaction. similar number from Sierra Leone and the Studies on seven Irish populations show Windward Coast, 13% from the Gold Coast, that genetic drift, in addition to migration and a few from the neighboring Bights of and admixture, can affect anthropometrics Benin and Biafra (Fig. 1). Notably in the and surnames as it does genetic markers earliest period as in the latest period, the (Relethford, 1991). Such random changes in Bantu-speaking Central Africans predomismall populations may well have been a pri- nated. In relative isolation for more than mary factor in human evolution. The role of two centuries, with little admixture, these culture and history in gene flow is well illus- coastal people retained much of their Afritrated in the Ards Peninsular of northeast can biology, culture, and language (Fig. 2). Ireland where a population divide, based Vocabulary from West Africa is found in upon seventeenth century migration of their common speech and in the personal or Scots Presbyterians into Catholic territory, basket names given to babies at birth is revealed in blood types, language, and (Turner, 1949). Kongo (Table 1) especially has contributed many words to our own lansurnames (Bittles and Smith, 1991). Surnames, inherited like the Y chromo- guage, including gumbo, goober, yam, and some, can delineate inbreeding as well as cooter (for turtle), as well as place names ethnic composition. My own investigation of (Vass, 1979). But, much of the deep strucmarriage isonymy from the 300 year old ture of grammar as well as vocabulary of records of Robin Hoods Bay on the north Gullah derives from languages of Upper Yorkshire coast of England, with the collab- Guinea. Religion and magic, music and oration of Malcolm Smith and Bob Williams dance, baskets and quilts, woodcarving and of the University of Durham, revealed the ironwork, and pottery and cemetery decoraethnic composition of the inhabitants and tions all bespeak the African heritage of the the role of parish boundaries and occupa- sea islanders. The biological person, in morphology, tions in channeling gene flow (Pollitzer et al., 1988). Even given names can be useful in blood types, and high frequency of sickle cell the analysis of population structure by dif- trait, shows only about 5% Caucasian adferentiating a cultural from a genetic compo- mixture (Pollitzer, 1958). Ilyas Kamboh and nent (Lasker, 1991; Guglielmino et al., Robert Ferrell, by analyzing blood samples in their lab at Pittsburgh, are providing bet1991). Of the populations that I have studied, the ter estimates of cdmixture based upon deGullah-speaking people of coastal South termination of genes, especially of apoli-

9

ETHNlClTY AND HUMAN BIOLOGY

TABLE' 2. Frequency

of

unique alleles of plasma proteins i n four populations'

Locus

Alleles

U S . Blacks (Pittsburgh)

APO A-IV

A-IVyl A-N*2' A-N*5'

95.5% 3.5% 1.0%

APO C-I1

C-II"l C-II*43

97.5% 2.5% 0.0% 0.0%

D*l D*2

99.0% 1.0%

E"2 E"3 E"4"

9.0% 66.0%

Populatiodallele frequencies U S . Blacks African is. Carolina) Blacks 96.3% 0.5% 3.2% (N 308) 97.3%

us. Whites

97.0% 0.0% 3.0%

91.2% 8.8% 0.0%

94.1% 5.3%

lO0W 0.0% 0.0% 0.0%

-

C-11x23

C-II^33 APOD AF'OE

2.OW

0.5% 0.2%' (N = 221) 99.0% 1.0% (N = 162) 9.0% 66.0%

25.0% 149) l.l%, 93.26 4.3% 1.4% (N = 139) 70.0% 30.0% (N = 143) 26.2% 66.3% 3.64, 2.1% 1.8% (N = 141)

25.0%

0.4%

0.1% 98.0% 2.0%

100% 0.0%

8.0$% 62.0% 30.0%

8.0% 78.0% 14.0%

1.1% 89.5% 7.3% 2.195

5.9% 88.2'10 5.9% 0.0%

72.0% 28.0%

100.0% 0.0%

21.8% 72.3% 0.0% 3.6% 2.3%

69.0% 11.0%

iN APOH

H*I" H*2 H*3 H"4'

1.7% 90.2% 6.8%' 1.3%

APOJ

J*I

76.0%

593

24.0%

~ ~ * 1 4 FB*24 FR*32 FB^63 FB"233

35.5% 54.26 6.2% 2.85% 1.3%)

FXIIIB

20.0%

0.0% 0.0%

'Data from laboralory of M. Ilyas Kambuh, Pilttiburgh. *Ilnique White alleles. %nique Black alleles. 'Cornmun alleles ghuwiny significant ditkrence between Whites and Blacks

poproteins, as shown in these preliminary results (Table 2). In several alleles, such as those of APO A-IV and APO J, the Carolina Blacks are closer to Nigerians than are Blacks of Pittsburgh. The special value of these investigations is that admixture can be estimated in the individual, as well as the population, providing data of clinical importance. In addition to its contribution to history, anthropology and medicine, such research should enhance the pride that people take in their heritage. Increasing knowledge of human diversity enriches us all. Beyond the awareness of differences is the appreciation of our common humanity. We are blood brothers and sisters. Human biology must also be humane biology. And that is in keeping with the life and the legacy of Stanley Garn. LITERATURE CITED Adams-Campbell LL (1992) Epidemiology of obesity: A focus on African-Americans and Africans. Am. J. Hum. Biol. 4t140 (abstract).

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Ethnicity and human biology.

Stanley Garn's book, Human Races (1965), captured the essence of the new genetic concept of ethnic groups as populations which share the same gene poo...
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