Lewontin concluded ‘‘Since
. . .
racial classification is nowseentobeofvirtuallynogeneticor taxonomicsignificance
. . .
,no justification can be offered for its continuance’’ (fullquotation given above).Lewontin included similar remarks in his 1974 book
The Genetic Basis of Evolutionary Change
‘‘The taxonomicdivision of the human species into races places a completelydisproportionateemphasisonaverysmallfraction ofthetotalof human diversity. That scientists as well as nonscientistsnevertheless continue to emphasize these genetically minordifferences and find new ‘scientific’ justifications for doing sois an indication of the power of socioeconomically basedideology over the supposed objectivity of knowledge.’’
These conclusions are based on the old statistical fallacyof analysing data on the assumption that it contains noinformationbeyondthatrevealedonalocus-by-locusanalysis,and then drawing conclusions solely on the results of such ananalysis. The ‘taxonomic significance’ of genetic data in factoftenarisesfromcorrelationsamongstthedifferentloci,foritisthesethatmaycontaintheinformationwhichenablesastableclassification to be uncovered.Cavalli-Sforza and Piazza
coined the word ‘treeness’ todescribe the extent to which a tree-like structure was hiddenamongst the correlations in gene-frequency data. Lewontin’ssuperficial analysis ignores this aspect of the structure of thedataandleadsinevitablytotheconclusionthatthedatadonotpossess such structure. The argument is circular. A contrast-ing analysis to Lewontin’s, using very similar data, waspresented by Cavalli-Sforza and Edwards at the 1963International Congress of Genetics.
Making no priorassumptions about the form of the tree, they derived aconvincing evolutionary tree for the 15 populations that theystudied.Lewontin,
thoughheparticipatedintheCongress,did not refer to this analysis.The statistical problem has been understood at least sincethe discussions surrounding Pearson’s ‘coefficient of raciallikeness’
in the 1920s. It is mentioned in all editions ofFisher’s
Statistical Methods for Research Workers
from1925 (quoted above). A useful review is that by Gower
in a1972 conference volume
The Assessment of Population Affinities in Man
. As he pointed out, ‘‘
. . .
the human minddistinguishes between different groups
there arecorrelated characters within the postulated groups.’’The original discussions involved anthropometric data, butthe fallacy may equally be exposed using modern geneticterminology. Consider two haploid populations each of size
’, at a single diallelic locus is
and in population 2 it is
1. (The symmetry is deliberate.) Each popula-tion manifests simple binomial variability, and the overallvariability is augmented by the difference in the means. Thenatural way to analyse this variability is the analysis ofvariance, from which it will be found that the ratio of thewithin-populationsumofsquarestothetotalsumofsquaresissimply4
0.7,thisratiois0.84;84%ofthe variability is within-population, corresponding closely toLewontin’s figure. The probability of misclassifying an indivi-dual based on his gene is
, in this case 0.3. The genes at asingle locus are hardly informative about the population towhich their bearer belongs.Now suppose there are
similar loci, all with genefrequency
in population 1 and
in population 2. The ratioof the within-to-total variability is still 84% at each locus. Thetotal number of ‘
’ genes in an individual will be binomial withmean
in population 1 and
in population 2, with variance
in both cases. Continuing with the former gene frequen-cies and taking
100 loci (say), the mean numbers are30 and 70 respectively, with variances 21 and thus standarddeviations of 4.58. With a difference between the means of40 and a common standard deviation of less than 4.6, thereis virtually no overlap between the distributions, and theprobability of misclassification is infinitesimal, simply on thebasis of counting the number of ‘
’ genes. Fig. 1 shows howthe probability falls off for up to 20 loci.One way of looking at this result is to appreciate that thetotalnumberof‘
’genesislikethefirstprincipalcomponentina principal component analysis (Box 1). For this componentthe between-population sum of squares is very much greaterthan the within-population sum of squares. For the othercomponentsthereversewillhold,sothatoverallthebetween-population sum of squares is only a small proportion (in thisexample 16%) of the total. But this must not beguile one intothinkingthatthetwopopulationsarenotseparable,whichtheyclearly are.Each additional locus contributes equally to the within-population and between-population sums of squares, whose
Graph showing how the probability of misclassifi-cationfallsoffasthenumberofgenelociincreases,for thefirstexamplegiveninthetext.Theproportionofthevariabilitywithingroupsremainsat84%asinLewontin’sdata,buttheprobabilityof misclassification rapidly becomes negligible.
Problems and paradigmsBioEssays 25.8