34 Evolutionary

ARTICLES

Apportionment of Racial Diversity: A Review

RYAN A. BROWN AND GEORGE J. ARMELAGOS

ogists and 31% of cultural anthropol- It has become increasingly popular to theorize and assert significant genetic ogists accept the validity of biological differences between arbitrary regional, ethnic, and racial groupings of . races in Homo sapiens.40 There are Beginning with Livingstone, Brace, and Newman is the early 1960s, biological many reasons for the decline in accep- anthropologists have shown that variation in traits is non-concordant along tance of race as a means of under- racial lines, as they are products of overlapping, dynamic selective pressures. standing human variation. Some In 1972, Lewontin analyzed blood groups, serum protein, and red blood cell enzyme claim that anthropologists are under variants and found that only about 6% of total genetic variance was accounted for by pressure to maintain a politically cor- race, while the majority of variance is accounted for by differences between individ- rect position.41–43 However, one of the uals. Using similar assays, Latter obtained similar results in 1980. In 1982, Nei and underlying reasons for the decline is Roychoudhury analyzed 62 protein variants and 23 blood groups, finding that roughly the arbitrary nature of racial classifi- 10% of genetic variance was accounted for by race. Analyzing protein, blood group, cations: The boundaries between and HLA variants, Ryman and coworkers obtained similar figures in 1983. More races depend on the specific traits recently, Dean and coworkers (1994) and Barbujani and coworkers (1997) have used used and the classifier’s own cultural PCR techniques to analyze RFLP and microsattelite loci, again yielding estimates of norms.11,25 Other reasons include the around 10% for the amount of genetic variance accounted for by race. Furthermore, lack of correlation of traits used in recent research on regional and racial variance in mtDNA (Excoffier and coworkers, classification11,25 and the existence of 1992), a traditional marker for human racial groupings, shows a higher proportion of alternative methods for explaining hu- variance within than across racial categories. man variation. These studies used a variety of assays and analytical techniques, some of which are The success of racial classification designed to maximize the amount of variance accounted for by race. In light of this, the low depends on the number of traits used proportion of genetic variance across racial groupings strongly suggests a re-examination in ordering the races.25 A single trait of the race concept. It no longer makes sense to adhere to arbitrary racial categories, or such as skin color will result in a clas- to expect that the next genetic study will provide the key to racial classification. sification system that is easily deter- mined. Add another trait and classifi- cation becomes a more difficult task, In the last 30 years there has been have discarded it as a research and and there usually are groups that can- an assault on the race concept.1–35 teaching tool.36–39 Even use of the not be classified. As you increase the Many anthropologists question the race concept in textbooks is declin- number of traits, the problems in ra- usefulness of the race concept and ing.39 Only 50% of physical anthropol- cial classification become insur- mountable. Newman44 used genetic traits as a Ryan A. Brown is a graduate student in the Laboratory for Comparative Human at means of systematically testing the va- Emory University. His main interests lie in developmental psychiatric epidemiology, with a lidity of Garn and colleagues’45,46 clas- focus on the relevance of physiologic reactivity to mental health. Brown recently returned from a summer of research on the physiology, , ecology, and ethnography of diurnal sification of geographic and local sleep in Egypt. He is currently planning a one-year longitudinal study of male adolescents and races. He selected a number of traits mental health in Cairo. and systematically determined whether George J. Armelagos is Professor of Anthropology at Emory University in Atlanta, Georgia. His or not they clustered. Newman found research has focused on diet and in human adaptation. He has coauthored Demo- that three of the geographic races graphic Anthropology with Alan Swedlund and Consuming Passions: The Anthropology of Eating with Peter Farb. He has coedited Paleopathology at the Origins of Agriculture with Mark (Asian, Amerindian, and African) ap- Cohen and Disease in Populations in Transition: Anthropological and Epidemiological Per- peared to “stand up well,” three spective with Alan Swedlund. He has been president of American Association of Physical Anthropologists, Northeast Anthropological Association, chairman of the Biological Anthro- (Melanesian, Polynesian and Micro- pology unit of the American Anthropological Association and Chair of the Anthropology nesian) fell within the categories of Section of the American Association for the Advancement of Science. Johnnetta Cole and he “suspense account” and “may be valid are collaborating on a book entitled Conversations on Race. but the critical data is lacking,” and Department of Anthropology, Emory University, Atlanta GA 31322. three (European, Indian and Austra- E-mail: [email protected]. lian) were labeled “unwarranted ab- straction.” He commented that Garn Evolutionary Anthropology 10:34–40 (2001) and coworkers’ list of local races “har- ARTICLES Evolutionary Anthropology 35 bor[s] many conceptual left-overs the first serious attempt to determine counted for 8.3%, and individual from the days of typological thought the extent to which racial groups ac- differences 85.4%. Thus, genetic dif- in racial anthropology” (p. 192). New- count for . ferences between individuals have lit- man was pleased with his results and Drawing on existing studies of the dis- tle to do with racial or ethnic bound- said that with more study and the dis- tribution of various biochemical aries. covery of new genetic traits racial markers in populations around the Lewontin’s analysis was met with classification would be refined. Unfor- world, Lewontin compiled data for disbelief. It was thought that if the tunately, he appears to have missed nine blood groups (represented by dif- “correct” genetic traits were used, the implication of his analysis and the ferences in immunologic response to race would be shown to be a major issues it raises for racial studies. a specific challenge), as well as eight source of human diversity. Critics ar- Newman unwittingly had discov- serum protein and red blood cell en- gued that if Lewontin had used the ered that racial traits are nonconcor- zyme variants. In the absence of poly- right genetic markers, his results dant; that is, there is no agreement merase chain reaction technology, would have been quite different. between traits used in racial classifi- these markers acted as proxies for ac- Lewontin’s results have been repli- cation.12 If there is concordance, ev- tual genotypes; each variant was pre- cated by many studies, which in turn ery trait will result in the same classi- sumed to represent one allelic variant. have used a variety of different types fication. For concordance to occur, Lewontin used two sublevels of of data and different analytical meth- each trait must be selected for at the population structure in his analyses: ods. In 1980 Latter59 used major geo- same rate and in the same direction. races (Caucasians, Black Africans, graphical contiguity to classify six In reality, genetic traits are evolving at Mongoloids, South Asian Aborigines, “major human subgroups,” subdi- different rates and in different direc- vided these major divisions into “re- tions, and consequently become non- gions” (northern, southern, and east- concordant. In practice, racial classi- Restriction enzyme and ern), and subdivided these regions fiers have to select the genetic traits polymerase chain into populations (Greek, Zulu, and so and morphological features that sup- on). Thus, Latter added an additional port their preconceived notions of reaction techniques layer of population substructure to race. Racial lines have been drawn make obsolete use of Lewontin’s analysis. Just as Lewontin along the axes of aggression,47 sexual had done, Latter used published data behavior,48 intelligence,49–53 athletic “genetic systems” as a on 18 “genetic systems” (ten blood ability,54–58 and just about every other proxy for genotype groups, three serum proteins, and five behavioral and psychological charac- because they allow enzymes). However, in addition to the teristic one can think of. Often, these Shannon information measure, he supposed racial differences are pre- researchers to gain a used two new analytical methods for sumed to be genetic. However, there is more direct measure of quantifying genetic diversity: the pro- a major line of evidence that makes portion of shared genes between two claims about racial differences in ag- the genetic code. To randomly selected individuals and the gression, sexual behavior, intelli- many who supported probability that two randomly se- gence, and almost any other charac- lected individuals will have different teristic highly unlikely, if not the racial model, this (nonidentical) genotypes. Using these completely untenable. was the scientific three measures, 7.5% to 10.4% of ge- This evidence comes from research netic diversity was accounted for by on how human variation is parti- breakthrough that would “major geographical groups” (races), tioned and distributed among individ- prove their case. while 83.8% to 87% of genetic di- uals and groups such as nations and versity was due to individual varia- races. The apportionment of human tion within populations. Interregional genetic diversity shows that only and interpopulation differences ac- about 5% to 10% of genetic variation counted for the remaining 5.5% to can be accounted for by traditional Amerinds, and Oceanians) and popu- 6.6% of genetic diversity. Variation racial classification. Research that lations (Chinese, Navaho, Portuguese, in percentages depended on the type supposedly classifies individuals by and so on). Employing the Shannon of biochemical marker used (blood their racial origins using genetic data information measure, a measure of the groups) versus enzymes. is methodologically flawed and ulti- frequencies of at particular In 1982, Nei and Roychoudhury60 mately is not very useful for under- loci, Lewontin analyzed the propor- analyzed data from 62 protein loci standing biological variability. tion of genetic variation accounted for and 23 blood group loci for “Cauca- A host of studies has concluded that by differences between races, between soid,” “Negroid,” and “Mongoloid” racial classification schemes can ac- populations within races, and be- populations. Assessing the proportion count for only a negligible proportion tween individuals within populations. of total genetic heterozygosity attrib- of human genetic diversity. In 1972, He found that racial classifications ac- utable to racial classifications, they Richard Lewontin24 published “The counted for about 6.3% of genetic di- found that this was roughly 9% to Apportionment of Human Diversity,” versity, population differences ac- 11% (8.8% for protein loci, 10.9% for 36 Evolutionary Anthropology ARTICLES blood group loci). Interestingly, de- blood groups attributable to race to lected and analyzed polymerase chain spite these very low figures, the au- 15.4%. This still represents only a mi- reaction data for 30 microsatellite loci thors went on to discuss “the pattern nor proportion of total human genetic and 79 restriction fragment length of of the three major races” variation. polymorphisms in 16 human “popula- (p. 11). This speaks to the logical dis- The salvation of those who believe tions.” When analyzed by major geo- connect shown by many researchers in the importance of race in under- graphic group, these combined DNA who simultaneously prove the irrele- standing human variability was a data yielded 10.8% of the total genetic vance of genetic race and then pro- more basic measure of genetic struc- diversity. Only a single locus was ceed to discuss the genetic evolution ture. Restriction enzyme and poly- found in which individual differences of races. The findings of Lewontin, merase chain reaction techniques within populations accounted for less Latter, and Nei and Roychoudhury make obsolete use of “genetic sys- than 50% of genetic variation. were so counter to conventional wis- tems” as a proxy for genotype because A recent study by Hartmann and dom that they stimulated more re- they allow researchers to gain a more colleagues64 claims to have found ev- searchers to seek the “right set of direct measure of the genetic code. To idence of greater genetic diversity be- genes” and “the right analytical tech- many who supported the racial tween than within races. This study niques.” model, this was the scientific break- sampled Hispanic, African American, 61 Ryman and coworkers replicated through that would prove their case. and East Asian populations in South- 24,59,60 the previous findings using two Not surprisingly, the use of restriction ern California, using polymerase human lymphocyte antigen (HLA) enzyme and polymerase chain reac- chain reaction techniques to examine types, nine blood groups, and 14 elec- tion to determine the portion of hu- variation due to genetic differences in trophoretically detectable protein man genetic variability that is attrib- restriction fragment length polymor- types. As in previous studies, these utable to race yields results that are phism sites. The authors claim to have data represented proxies for actual ge- found evidence counter to the conclu- netic loci. Ryman and coworkers sions of Lewontin.24 However, they searched the literature for data from ...intrapopulational base their claim on an analysis of re- 18 populations, which they subse- striction fragment length polymor- quently grouped into six to seven differences still make up phism size diversity that compares the more inclusive groups (races). An im- the bulk of mtDNA genetic diversity between racial portant point is that they created groups to that between regional these broader groupings using two genetic diversity. This is groups, not between individuals. In distinct methods. First they used remarkable, given that doing so, Hartmann and coworkers64 traditional “anthropological races,” blatantly ignore the findings of Le- based on classical racial typology; mtDNA is said to be the wontin and all of the subsequent stud- next they used a computer program best marker of “racial ies on population substructuring and designed to create groups using den- origin.” the proportion of genetic diversity at- drograms of genetic relatedness. tributable to race. These studies Using the Shannon information showed that the majority of variance measure, Ryman and colleagues61 in genetic diversity is attributable to found that individual variation within individual variation within racial cat- populations accounted for 86% of all similar to those of past studies that egories, not differences between eth- genetic diversity. The numbers ranged used genetic traits. For example, Dean nic or regional groups. Hartmann and from 82.7% (blood group loci) to and coworkers62 conducted a study of colleagues64 do not perform the nec- 90.3% (electrophoretic protein sub- 257 restriction fragment length poly- essary analysis to determine what per- types). Interestingly, use of the com- morphism loci in American Cauca- centage of size diversity in restriction putational dendrogram slightly in- sian, African American, Asian, and fragment length is at- creased the proportion of genetic American Indian individuals. Using tributable to individual variation. diversity attributable to the broader Lewontin’s24 statistical techniques for Recent analyses of the genetic appor- groups (an average of 1.3%) as com- approximating heterozygosity, as well tionment of mitochondrial (mtDNA) pared to the use of classical racial ty- as a technique that describes gene di- and Y chromosome diversity comple- pology. This is expected, given that versity more generally, they found ment and support the results obtained the program was designed to maxi- that between 9.5 and 10.1% of the di- for autosomal genetic diversity. For mize genetic differences between versity in restriction fragment length example, Excoffier and coworkers65 groups. It does not counter the claim polymorphism loci was attributable to found that roughly 75% to 80% of that racial groupings account for a rel- racial categories.62 Interestingly, com- haplotypic diversity in mtDNA resided atively minor proportion of total ge- parisons of racial groups that allowed between individuals within popula- netic diversity, given the small in- the use of more restriction fragment tions, with 15% to 22% attributable to crease yielded by reorganizing the length polymorphism loci yielded regional (racial) origin. Seielstad and racial categorization scheme. Using a much lower estimates of interracial colleagues66 found that roughly 81% dendrogram to define races only in- variation, 4.4% to 4.75%. In a similar of mtDNA diversity was attributable creases the genetic diversity from study, Barbujani and colleagues63 col- to individual differences within popu- ARTICLES Evolutionary Anthropology 37 lations and that close to 16% was at- that this dynamic biases results in favor phism studies62,63 also produced tributable to regional differences. The of individual variation within popu- similar estimates. slightly higher proportion of mtDNA lations. However, localized frequency- Some evidence presented in genetic diversity attributable to continent of dependent selection does not preclude apportionment studies seems, on the origin is likely the result of a lack of broader geographical (racial) differ- surface, to support Miller’s67 “right significant selective pressure on mtDNA entiation. In fact, the differential geo- genes” argument. Notably, the per- (thus making convergent evolution on graphical distribution of infectious centage of genetic variation ac- different continents unlikely), as well disease types would likely favor ra- counted for by race differs for differ- as its haplotypic pattern of inheri- cially based genetic diversity in these ent loci. Both Lewontin24 and tance. Nevertheless, intrapopulational systems under Miller’s67 scenario of Barbujani and coworkers63 list sum- differences still make up the bulk of disease-based selection. mary for individual loci. Le- mtDNA genetic diversity. This is re- Lewontin24 and Latter59 present wontin24 found two loci in which ra- markable, given that mtDNA is said to evidence that is even more damaging cial differences accounted for roughly be the best marker of “racial origin.” to Miller’s67 argument. They used a 25% of total genetic diversity (Rh and A recent investigations of Y chro- variety of nonantigenic systems, Duffy blood groups, 25.9% and 25.3%, mosome genetic diversity claims that which generally yield even more respectively). Meanwhile, Barbujani individual differences within popula- conservative estimates of the per- and coworkers63 found four restric- tions may account for as little as 35% centage of genetic variation attribut- tion fragment length polymorphism of diversity in the Y chromosome,66 able to race. For example, Ryman, sites in which racial (“geographic”) with between-continent variation ac- Chakraborty, and Nei’s61 results grouping accounted for 22%, 23.9%, counting for almost 53%. However, show that nonantigenic systems, in 35.5%, and 42.7% of total genetic vari- these figures hold only for single nu- ance. However, these loci are ex- cleotide polymorphisms on a nonre- tremely rare (two of 17 and four of combining segment of the Y chromo- ...it seems that 109). The high proportion of genetic some. When microsatellite diversity is the morphological diversity explained by race in these considered on parts of the Y chromo- examples could easily be the result of some that do recombine, individual characteristics that coincidences between genetic drift variation accounts for more than 83% racialist scientists and and culturally transmitted racial ty- of Y chromosome genetic diversity. It pologies. is important to note that these appor- others do find useful are Nevertheless, it is likely that those tionment figures were obtained from determined by simple such as Miller67 will seize upon these the analysis of a noncoding segment variations in very examples as giving hope to the theory of the Y chromosome. Thus, relaxed that “those important genes” actually selection pressures could lead to in- specific regions, and are do differ significantly along racial creased variation by continent, for the quite direct results of lines. For example, commenting in ra- reasons described earlier. Likewise, cial apportionment studies, Miller67 the haplotypic nature of Y chromo- climactic selective declared, “None of these genes af- some inheritance might also lead to pressures. fected skin color, nose shape, body increased intercontinental variation. build, size, etc. to mention [sic] char- Most importantly, neither mtDNA nor acteristics that differ between races” the Y chromosome DNA discussed (p. 97). Counter to Miller’s argument, previously code for anything function- it seems that the morphological char- ally significant. Thus, these studies do this case electrophoretically detect- acteristics that racialist scientists and not provide evidence of racial varia- able proteins, exhibit the least ra- others do find useful are determined tion in behavior, intelligence, or mor- cially contingent diversity. Further- by simple variations in very specific phology. more, Ryman, Chakraborty, and regions, and are quite direct results of Not surprisingly, these results have Nei61 suggest that these data are climactic selective pressures.68 Owens not been well received by racial typol- most likely to be representative of and King68 argue that variation in ogists and others who dislike the po- total genomic diversity due to closer “stereotypic features of ‘race’” such as litical implications of the lack of ap- mapping between codon differences skin color, hair form, and facial fea- preciable genetic divergence between and electrophoretically detectable tures are “literally superficial” and races. Some have argued that Le- differences. Given that these au- does not support the utility of the race wontin and others simply have not thors’ figures for electrophoretic concept. used “the right genes” in their analy- “loci” estimate racially contingent A number of studies have emerged ses. For example, Miller67 has argued genetic diversity to be 5.4% to 7.3%, that use genetic data to classify indi- that local frequency-dependent selec- depending on the classification viduals based on race or “geographi- tion should increase the local diversity scheme, it appears that Lewontin’s cal origin,” perhaps the most notable of HLA and other antigenic blood original figure of 6.3% is close to the recent attempt being Cavalli-Sforza group types due to evolutionary arms mark. Notably, microsatellite and and coauthors’ The History and Geog- races with infectious , and restriction fragment length polymor- raphy of the Human Genes.69 Such 38 Evolutionary Anthropology ARTICLES

TABLE 1. STUDIES OF RACIAL APPORTIONMENT % Study Data Statistic % race % region population % individual Lewontin24 (1972) 17 blood groups Shannon information 6.3 8.3 85.4 measure Latter59 (1980) 10 blood groups, 3 Shannon information 2.8–14.0a 0.4–6.1 1.7–5.4 80.2–90.7 proteins, 5 measure, Hedrick enzymes genic similarity, Latter paired comparison Nei and Roychoudhury60 62 protein, 23 blood Nei’s standard genetic 15.0–19.0c 81.0–85.0 (1982) group difference,b but parsing of variance unclear Ryman, Chakraborty, and 14 protein, 9 blood Chakraborty hierarchical 6.4–15.4e 2.0–6.6 82.7–90.3 Nei61 (1983) group, 2 HLAd gene diversity Excoffier, Smouse, and 34 mtDNA loci Bootstrapping, null 15.7–22.0g 3.3–3.6 74.7–80.7 Quattro65 (1992) distribution assumptionf Dean and coworkers62 115 RFLP loci Unweighted averaging 9.5–10.1h 89.9–90.5 (1994) of frequencies Barbujani and coworkers63 79 RFLP, 30 Bootstrap multiple allelic 10.0–11.7i 3.9–5.5 84.5 (1997) microsattelite loci frequency comparison Seielstad, Minch, and 34 mtDNA loci,j 22 Bootstrapping (Y and 12.5 (mtDNA), 6.1 81.4 (mtDNA), Cavalli-Sforza66 (1998): biallelic mtDNA), Unclear 52.7 (Y)l (mtDNA), 35.5 (Y) single polymorphisms (autosomal) 11.8 (Y) (Y)k Seielstad, Minch and 29 autosomal loci, Bootstrappingm 2.0 0.2(autosm) 97.8 Cavalli-Sforza66 (1998): 10 Y chromosome (autosm) 14.0 (Y) (autosm), microsattelite loci 2.5 (Y) 83.5 (Y) a Depending on data type (blood group, protein, or enzyme) and statistic used. b Approximates number of codon differences. c Parsing of variance unclear, range of percentages unexplained. d From Nei and Roychoudhury.60 e Depending on data type and classification (a priori or post hoc to maximize group differences). f For analysis at each level of population substructuring. Assumes that other levels are artificial and random. g Depending on analytic technique (Euclidean distances between haplotypes, assumption of equal distance between haplotypes, network-based with racial assumptions, network-based with allowance for nonlinearity). h Depending upon method of estimating heterozygosity (Lewontin,24 Nei).77 i Depending on assay type (microsattelite or RFLP). j From Excoffier et al.65 k Autosomal statistics taken from Barbujani et al.63. l On nonrecombining part of Y chromosome (from Underhill et al.).76 m Similar methods to Excoffier et al.65

studies select the small proportion of flawed in design and easily misinter- workers70 entered their genetic infor- genetic variability that is roughly ap- preted. An example of this can be mation into a program that already portionable by race to plot out den- found in Bowcock and coworkers.70 used the a priori racial categories they drograms of essentially false categori- Despite a research design that should were trying to replicate. zations of human variability. To have maximized the degree to which It is important to note that these accomplish this, these studies use a the researchers were able to classify researchers’ results do not in any way priori categorizations of human vari- individuals by racial category, the re- locate diagnostic genetic markers for ability that are based on the inaccu- sults are something less than “high racial origin. Rather, they are based rate belief that classical racial catego- resolution” with respect to this goal. on pairwise comparisons of relative rization schemes delineate a series of For example, 88% of individuals were genetic distances. Thus, individuals isolated breeding populations. This classified as coming from the right were clustered by distance and then pursuit is not only futile; it leads to continent, while only 46% were clas- labeled based on a priori information faulty and misleading conclusions sified as coming from the right region about their origin. These labels were about the nature and origin of human within each continent. Notably, 0% assigned in a way that characterized differences. success was achieved in classifying the largest proportion of individuals Thus, studies that attempt to define East Asian populations to their region in each sorted group, a tactic that individuals on the basis of racial or or origin. These results occurred de- again gives Bowcock and colleagues70 “geographical” location are both spite the fact that Bowcock and co- the advantage. This makes the fact ARTICLES Evolutionary Anthropology 39 that only 88% of individuals were clas- tems in racial typology studies yield 8 Blakey M. 1987. Skull doctors: intrinsic social different classificatory schemes.23 and political bias in the history of American sified correctly by race and 46% by physical anthropology with special reference to region even less impressive. This is strong evidence of the clinical the work of Ales Hrdlicka. Critique Anthropol Other studies that classify individu- distribution of traits due to drift and 7:7–35. als into racial groups produce results differential selective pressure, a pat- 9 Blakey ML. 1994. Passing the buck: natural- ism and individualism as anthropological expres- that argue against racial typology. For tern that anthropologists have known sions of Euro-American denial. In: Gregory S, example, Excoffier, Smouse, and about for decades. Sanjek R, editors. New Brunswick: Rutgers Uni- versity Press. p 270–284. Quattro65 used a computer to create The evidence against genetically 10 Boas F. 1910. The real race problem. Crisis 7 racial groupings that maximized the mediated differences in behavior (Dec); amount of genetic diversity between along racial lines is overwhelming 11 Brace CL. 1964. On the race concept. Curr groups. Thus, while defining regional (Table 1). First of all, a host of studies, Anthrop 4:313–314. groups a priori, they allowed genetic beginning with those by Lewontin24 in 12 Brace CL. 1964. A nonracial approach to- wards the understanding of human diversity. In: distance to define racial groups. As a 1972 and most recently by Barbujani Montagu A, editor. The concept of race. New result, these authors observed that the and colleagues63 in 1997, have shown York: Free Press. p 103–52. ideal racial typology using mtDNA that the amount of human genetic di- 13 Centers for Disease Control and Prevention. versity that is attributable to race is 1993. Use of race and ethnicity in public health grouped South American Pima Indi- surveillance: summary for the Cdc/Atsdr work- ans with Finns and Mayans with Sicil- only about 5% to 10%. Following this, shop. MMWR 42:1–17. ians! any particular “population” includes 14 Du Bois WEB. 1898. The study of the Negro roughly 85% or more of the total hu- problems. Ann Am Acad Political Soc Sci 11:1– The inadequacy of thinking about 19. man genetic diversity.68 Also, racial human diversity in terms of isolated 15 Goodman AH. 1994. Problematics of “race” in breeding populations is well illus- genetics research that categorizes in- contemporary biological anthropology. In: Boaz trated by the work of Keita and dividuals by genetic profile, as does NT, Woke LD, editors. Biological anthropology: 70 The state of the science. Berd, OR: International Kittles,23 who have demonstrated how that of Bowcock and coworkers or Institute for Human Evolutionary Research. p the pairwise genetic differences found that defines genetic differences be- 221–243. by Bowcock and coworkers70 and tween hypothetical human “popula- 16 Goodman AH, Armelagos GJ. 1997. The res- tions”60,71–75 is plagued by a host of urrection of race: the concept of race in physical other researchers most likely are not anthropology in the 1990s. In: Reynolds LT, the result of separate breeding popu- methodological flaws and faulty as- Lieberman L, editors. Race and other misadven- sumptions about human evolution. tures: essays in honor of Ashley Montagu in His lations evolving in different direc- ninetieth Year. Dix Hills, NY: General Hall. p tions. Rather, this variation (which of Thus, results that on the surface seem 174–186. course is very small, as illustrated by to demonstrate genetic differences be- 17 Goodman AH, Leatherman TL. 1998. Travers- ing the chasm between biology and : an Lewontin and others) is more proba- tween the human races are actually quite meaningless underneath. We introduction. In Goodman AH, Leatherman TL, bly the result of selection pressures editors. Building a new biocultural synthesis: po- and genetic drift. Likewise, similari- support Owens and King’s conclu- litical-economic perspective on . sion:68 “The possibility that human Ann Arbor: University of Michigan Press. p 3–41. ties between population samples are history has been characterized by ge- 18 Gould SJ. 1993. American polygeny and not necessarily a result of isolated craniometry before Darwin: blacks and indians netically relatively homogeneous cases of population admixture be- as separate, inferior species. 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