54U Al\1EllICAN JOUItNAL OF PHYSlCAL ANTllltOPUL!
IGY
1\LL,\'o HU L I~ IN CHA:\01NU Tll E li'ACE ()l<' 'I'll E E,\I{TI1.
\Villialll L. 'rllOlIlas, Jr., Editor, \"itlt collahoration or Carl O.
o,lllcr, ::\Ial'slon BatcN, and Lc\,,;s ~\lLllllrOl'l1. 11:):) [ijl., 180 ill us.,
;;;) tallles. (c;l:2.,')(). I :lIi\'ersity of Chicago l'n's'i. l:J::Jli.
1Il'l'l~ an: ]!lllJlioiled ,'J:) ]1,lckg]'ound ]JilpCl"i and slUllmClries of
discllsoions at tlll: \Velllll'l··Gr('1l F'oullliatioll's symposiulll Oil ":\1<111'0
Hole ill ('lwngillg tlw 1''<lc<: or tilc Earth" held .Jlln(~ 1G-:2:2, JD.')CJ.
TlJis \"(~l'y bU'g't: VOllllllC coml,illl'S the points of "iew of 'i('vend (li.s­
~ijlli!1('s whell ~ollsidl'rilJg 111(~ efl'l'cls of lllllWllI <ldi\'ity, nll11 i!1('lllllc:,
('xtellsive bibliography, illlls!l'ntiolls ,lUd IlI111lcrOllS dnL!. .\0 Sllllllllill'y
of sll~h a \'OllllllC c<ln do jnsticp to it - only' <I rC<J(1iw~.
P.P.T.
t 1 Thi::;
I
._.1.
ON 'l'HE EVIDENCE OJ1' HANDO'J\I GENErl'IC DB lJ1''l'
IN HUMAN POPULA'l'lONS 1
DJ:;;?-;'['Ll': Y GI, Ai'JS
JJcparlllicilt of IJio!oqy, The .lolll!" llollkillo l/liversity
I;\TJWDUCTION
The name of Sewall 'Wright is invariably, inevitah1y asso­
ciated with the concept of gcnetic drift. Gencti.c drift is even
commonly rdened to as the "Sewull\Vright (oSTeeL" This
is the more surrlrising when a careful stuely Jl1nst indicate
that Sewall 'Wright himself has scarcely c~ver nsed thc~ phrase
and at no time has iudicated it io be an evolutiolHll'y factor
of dccisi.ve sigllificance. 'Wright ':,; OWll term of l)rcference is
"random genetic drift," \\' hich lllO re explicitly labels the
nature of ilw fador than do(~s Ute COllllllOll1y nscd phra:w.
'Wright ('4Da, b; '50, 51a) has dassificd the lllo(les of e1WIIg'C
in g(:ne frecluency into thn)e categories, (1) those f'(~snltillg
from systmuatic pressures (recurrent mntation, illtrapo[Ju­
lation sc~leetiull, and l'l~Cl1rrcnt immigration and crossbreed­
ing), (2) those I'("su1ting from Huctuatiolls in t he systematic
jlreSS11l'C'S or from ra]j(10m accic1ents of sampling, an(1 (:3)
thosc~ resulting from unicl11e events, S11C~h liS a ulutation
favoralJ1e at its first OCC~11l'I'ellCe, a nnicIlw se]()divc: event or
hyhric1izatioll, s\nmlping hy mal:is immigration, OJ' a unifjlw
recluction in lmmhl~rs. Amolll; thesc~,it is the lIue( nat ious ill
gcnc freCJ.uencies tllle to acci(1e1l1S of smnplillg' ill lneiosi"
and fertilization t1wt an' propcrly ca]]ed randolJ) genetic
drift whenevl~r they hec01l1c compollncled ovc~r sl1ceessin~
generatiolls into wic1e c1eparlul'es from (he origillnl eqlli·
ll~lI>el' ,yas gin'lI :l:-' a Jwrt or a sylllposillll1 orgallized in hOllor of 8t'\Y~lJl
\Yright lJy the Biumetric 8o(,)(·t.,", the GeneUcs of Alllcrieu, :Illa t1l1'
SOeLllty
AlIlf'r1CtlIl Socid:Y of 1I1l1Urln Ch-u('ties and hl'1<l at the A.T.H.S. lllce1illg-s ill };;lsl
Lan~l11g, 1\iidligan J on Se})t8Jnber G, 1955.
511
.\:,-rEr::rC.\K JOUlC"-_-\lJ OJ,, l'HYSWA1, .\N'l'llROPOLOGY, VI)L. 14 .N.s., KO. ·1
DDCE,)IllElt 1956
 ;J-+ :.l BENTLEY GLMiS
librium. rrlw alterations of gene fl'l'llllellcy Ilrodueed hy
uuique eVl~llb; Itave O!'tl'U hl~ell illclulll'd in tlte eoneept of
genetic ch'ifj l)()eallSe they too repJ"(~sellt aceillents of smnpJing,
ill a sC'n;,;e, ill particular such l~Vi'llts as a llniquc J'l'c!nctiolJ in
uUflllJl'],:S, wllich like gelll~tic lhift is e1o;,;ely rl'1ate<1 to the
l~volllti01I,ny d'fect of the i:ii~c' of popll]ntiolJ.
1\0 one \\'0111<1 qnl':stioll that aeeilleuts in sarnpJillg thl' gl'ne
pool of a lJarC'ldal genc~J'atioll lllU:st oeeUr in the [orlllatioll of
the gell(~ ]lool ill tlw offspring gml(,J'atioll; or tbnt thl' magni­
tUlle of tlwse dC'viatic)H;'; from expectation varies inn3rsely
witli the size of the gmw pool, or with X, tIll' effl'ctive ;,;iz(~
of tlw JloJiulatic)lI, TIll! ndller aerimonions l'OldJ'ovc~rsy that
has lllal'lwd discu;,;;,;ioll;,; of gell(~tic drift in the past ;';l'l'IllS to
he 1>n;,;e<111ot on a gelJuine difference of' olliniclll iu Oli;,; l'l~spect,
\l\It ndlw]' U[l011 a ]Jel':sistent failIu'(' of tll(' 0ppOllents of
gc~nl'tic drift to rl'cogllize it as a ;,;ignificmd l']l'nll'llt in the
interplay of evolnti01Iary forees. Wrigbt's view:s, properly
I'epl'l'sentl'd, :stres;,; OIl' great sigllifieanel~ of intergronp selec­
tion in l'elativcoly largl' poplllation;,; snl)divided into locally
diffel'l'lltiatell ;';l1hgrollIls. Both the dfeetiveues:s of i;,;olatiou
of thc~ ;,;nhgrollps al](1 their dl"edive pOllnlatlo11 sizes are
irnportallt to thl' a1l10nnt of 10l',al difTen311tiatioll that call
O('Clll". Althollgh \\'\'iglit agrc'es tll11t "filletllatiolls in gene
heqlleneius ill sma1l eomplc:tely isolatell C'omrnnnitil'~ rarely
if l~ver COld rillllte to eYOllltiOlwry advanC'e, lillt mel'ply to
trivial <1ifferelltilJtioll, or ill ('xtn~1I1e ('uses to c!l'geuerntion
aJ}(l extinction" (\\'right, '51h), this j;,; not to dcny tlwt randolll
genetil~ chin teJj(ls to oppose the (lirectioll of sc,lection iu
;';llch pOTllllatioll", amI may l'itller oYl'r\\"hpllll it (\vl!C'lwver
s i;,; Ipss than f'i) or moclify it (\\"lwnevpr s i;,; greater
than 2~)' rrhe outcome is simply l1etl'rminel1 hy the relative
magnitul1l's of the interacting evolutionary factors. It is in
principl'1 1111so1111(l to ignore rallllom genetic l1rift as heing
negligihll' ill effed except when it can l)e l1emonstratC'c1 that
the magnitwle of selection and the size of population arc
such as to make it Tlegligible. Eve11 \"hen the size of population
is qnite acellrately known, aml when it seems olwions that
GE;<IETIC IJHIFT IN HUMAN POPULATIONS 343
selc;ction is favoring the sIll'eal1 of one allele and the reiOtrietioll
of its cOlllpetitor or cornpetitor:s, as ill the case of the medioni.
gra allele in thl~ moth ['({)loxia dOlUinula L. ;,;tudiec1 ;';0 fully hy
Jj'isher aud ]'onl ('47) aml by Sheppard ('5:i), it sti1l Joe;,;
not follow that nOlle of the change in gelt(~ frequellcies at tlw
locus studied i;,; attribntable to nlndom acC'idents of samplillg.
Nor does it follow that at oOwr loc~i ill the ;,;mne population,
whel'Cl the sl!lect ion preSSlue hetween com pet Lllg ulle les is
lower, genetic drift TIlay llOt he the predominant factor in
l'stahlishillg the local freqm'I1cies of the alleles.
Tile delllOJlstrutioll of i;CJletic drift bv
age-grou]J IlJlaL.lJsis
rrhe literature on lllood group freqnlmeies is full of pro­
llounced <1eviatlOlls occnrring' in small, genetically rathl~r
isolatell groups. A previous stndy (Glass et al., '50) dealt
with a group \\llere it may lw assllIned 1n01'8 plausihly than
in most situutions that the PIlvil'Olllllelltal, :splectiv\, conditions
acting npOlI the isolate Ul'e ielentleal with those in the main
population wit1lill which the isolate exi;,;ts. rl'llis was t1w stndy
of gelle t'l-equcneiEJs iLL a COllllllUllity of Dunkel'S living' ill
}'ranklill C01l\\ty, Peum;ylvl1llia, on farllls ,In<1 ill small towns
intermillgled with the gelwral population. rflIe l~vicleneu
presented indicated a striking dif[urenee ill gene frequencies
at the ABO lows, at the MN locus, and ill respect to cert.ain
other pllysical trait:s of no klLown or ohvious selective signif­
ieallce. It: may, howevl'l', 1)'; ohjeetecl that ill tlWS(~ im;tances on\~
cannot disting"llish the random genc~tic drift over a succession
of geneJ'ations from the effect of' a unique sampling that pro­
vided a very few genotypes in the progenitors of the isolate.
Probably the most satisfactory demonstration of genetic
drift that can he made is to show that in snceessive generatiolls
of an isolate which shares a common ellviromnent with a large
population within whieh the allalyz(Jll gClW frc>\lUencies arc
quite stahle, the gene frequencil's ill the isolate on the contrary
arc shifting in such a way that aHhough no two successive

 54·1 BENTL]<JY GLASS

GENETIC DHIFT IN HUMAN POP ULATlONS 545

generatiolls manifest a statistically significant difference, the transcends the range betl\'l~en the ,V. German populatioll
cumulated shift over several generations possesses statistical from which the Dunkel'S originally came and ihe major
significance. rI'his Illay be doue by the method I have else­ population within whieh they now live, must he attributed
wllUre (Glass, '54) called "age-group analysis," which has to some shift in gene frequencies that occurred more than
now been applied to the data obtainecl from the Dunker three generations ago - perhaps to the unique sampling eHect
community in Ij'ranklin County, Pennsylvania.
that occurred when the '27 Dunker families first caIlle to
'1'ABLE I
TABLE 2
lIf(l!d,lin Couuly (Pa.) lJ"ithrs -- Age-group analysis
FranUin C('1(n ty (Pa.) Dunkers -- A iJe'-yrOl')J anal!!"i"
I. ABll 1100d group" -l'hellotYlH''i
- ----_._---­
(>
_ .. _--~ ---_.­ -----­ I. M~ J,lood groups --- p!Jcnotn'es
u AB TOTAl,
-­ ------­
"
.\1'..; Tt!TAL
Au. S{) .SII. (~ ~V ,i. /-.'
.vo. -,--- -- ---------­
/u
Gen. l No. 0/0 .,yo. s{) Sri. C,;)
Hi ~O 2 0 44
-,,-------/
nell. 1 12 28.6 ~~ :12.4 8 1:J.n 4:2
:;tiA ' fiD.O -J-.O ((pn. 2 34 44.8 ({2 42.1 10 l:U Hi
~
U\"ll. 3 48 5;'.8 30 :H.:J 8 ~U BI;
(;{'ll. 3-1: 47 86
_,--_----.J
x:l:! Gen. 1 and 2 = 2.DS .30> P > .~O ::-f =0 204
JD.G 54.G ,').~
x', Gen. 2 and 3 = 2.00 .50> P > .30
n('11. :1 ,;1 ('0
L 2 98
x', Gen. 1 [JIl,l 3 ~~ 9.50 .OJ > P> .00;;
,~--./
;1JJj 03.3 5.1 T r. )[ <.LlHl .N g'l..'lJC frt:lJ.llC'lIcies
-----._--- --- ---_._--­ -_._------ ---_._---­
------
X~::i GCll. ] and :] 0.27 .975> P > .:),') N= 228 "
K
------,---_._.~----
TO'\"\!.

----­

X 2.J Gl~ll. 2 and 3 1.40 .80 > p > .70 }". o. f Ao .

x', Gen. 1 alld 3 0.29 .975> P > .95 (J ("11. 1 4o(i ;~8 BJ
x 20 Gell. 1, 2, and 3 1.50 .975> P > .05 .5;;0 .4;;0

Gell. :3 Ilill ~J2 15~

II. A, B, and 0 gene frC(IUcnei,'s
.li85 .342
A B o TOTAL
------------- Gen. 3 J:2ti 46 172
;Yo. i ~Vu. f- .Yu. f· 0735 .205
Gen. 1 32.5 .:170 ".w
00
.036 52.3 .594 88 _._-------------_ .. _ - - - - ­
Gcn. 2 59,0 .347 2.G .015 10D.8 .G38 172 x\ Geu. ] nnd " ~.:JO .HI :> P > .0;; ~ = 408
GCll. 3 81.0 .413 4.0 .025 110.0 .5G2 196 x'\ Gen. ~ [\llcl 3 = 2.11 P > .10

x'. = 3.32 .70> P> .fiO

X', Gen. 1 ~n!l 3 = 8.55 .00;; > l' > .00]

N = 456 X2 Gen. ], 2, 3 = 8.83

;.l .O~ > P> .0]

Table 1 gives the analysis with respect to the ABO blood North America ill the early 18th ceutury, or to nIlother uniqne
group frequencies. It is obvious that there occurred no sampling effect that occurred when the Dunkel'S split away
significant shift in these frequencies over the period of the from the more numerous and less strict branches of the sect,
three now-living generations, aged 1-27 years, 28-55 years, in 1881; or possibly to genetic drift that occurred more gradu­
and 5G years of age or older. Consequently the highly sig­ ally during some portion of the intervl~lling time.
nificant difference between the ABO frequencies of the Table 2 presents the analysis for tIle ]\(N hlood group
Dunkel'S and the general U. S. ·White population, which frequencies. ,Vhether the analysis is carried out on the basis

 ;)4(;
DEKTl,};Y GLASe;
GENiOl'lU IJI:1F'f IN nUIVIAN POPULATIONS 5+7
of tho phenotypic fl'oqlwllcies (tahle 2, I) 01" (Ill the hasis of \yiclely from the seeond, and very signiiieant Iy frollI tlJ(~
the g'C1l8 f'requclIeiee; (ta])lo 2, 11), it i" e]Cl<W that although frr~qlllm~ies of the iirst aUll secoJllI gel\(,ratioli~ cOllIbined.
nei ther the shift hc!\\"eC!1l the Ii l"S t alld soconcl g'Cl w nltions, It is well at this 1ll001lC'nt to "ound a 1wte of \\'<1 riling. Con­
1101' the "hift lJet\\"eell the r-;econd ancl thi 1"(1 gellerat iOlls, is
ceivably the shifts ill thc~ frl'qllclICY of a gl'nctic 1\'ait ill llif­
significant, yet the continuatioll of the "hift ill tlw SHmo ferent generatiolls, if sl/iJIjl!n! (It r1iD'erent ((ges, lnight he due
direction over two g'elleratic)]ls ka<1,., to a r-;igllificanl <1iff'C'l'euce to differential 11I0rtaliiy, or to n latu, vnJ'ying' ag'e or on"et,
Iwt\\"een the first al111 thinl gem'rations.
or to lllOlliiicntioll of tlw trail ])y envil'c)\lnwlltnl 11l'e""llrus.
TAULI; :: In the pres(!nt lusLillcl', alt hough r \\'ou\cl \lot can' to "ng-gc'st
li'·""l.;li,, COil"!!! (1'<1.) ])nili;{rs~ FLh liloo!l t,ll(iCS that rig;htlw!lllucl people tend to outlive ll~nltalldc;d Olles, I
{{In:!
----_._--- - ' - . -" --- -- ----~--­
tltillk that there may well havo hCl'l1 a gn;a(l'l' d(!gT\;e of fmnily
H~ It,_, rb TUT c~L
TI"
So {,:; } ; 0, c'
,I ~'{(). (;{i Xo.
amI socinl l)r('ssun~ tu t nlllsL'Ol'lll lc!ftllnndeLl ehildren into
~:-~ .:\ n, ~.{;
U(\I1. J 5 1 J.D :2:3 5+.8 H) 23.1J 4 U.tj +~
right in 1'o['\\wl' gCl\(~ratiol1s thall no\\', Parellthl'ticnliy, it
(~l'll. ~ ] G 2 J. I :39 51A 1I IV, fj ]] .8 j
.,

llIay be poillted out that, taken at raee \'alm', tltl',.,e data won]cl
<kll. ;·l 9 JO.6 .0
'iG
55 G·l,O 15 17.J 7 8.1 IJG all!lear to cxeJulll' Eife's (';)1) hYJ)o(hesi,., that lllllhidC'xturity
X~tl =~ (iA:2
.50> p> .;10 is till' llCterozygous cOllclitioll of II ll1011ol'acto]'inl g'elldic
C\ = ~O+
(N" ll('gh.t't~~d).
lliffcrl'llce. I>nt of com'Sl' til(' lliagnostie critl'l'ia in !l\l' pr('Wllt
T.\BL1; .[ stndy lllay not he tho,se used hy Hife, a\ld ill fact are hasccl
J"rankUn COllulll (Po.) lJ/()tkcr", --lland(lr71/l':5S "olely on till' "tatmllcnls of tlte sllhject" !llL'lmwlve".
].
Lit \( 'T'i 1'1'.\] ,
1'ahll' ;) gives the Hg'l'-group allnlyse.'i for ;J otllel' ph)'sic;nl
(;l'll. I --- -- -. __ ._-- .. _~-----~-­
9'0 R.
traits, for which the lllolh,s of inlwritancp lmve llOt !ll;ell 1'1111y
0 +3 H
{;('fl. ~ 0) D7.7

[I 1Jl) 8:1
D7.6 worked out. A.Hhongh as a wltole, l1lidcligital IlHil' ]wttel'lls
nt'lt. :~ 1~

U,) IO!) <S7.2 fail to :ohow a statistically :oigniftcmlt cliffl'l'PllCC hc~h\'(,l'll

7.~%, H:3_S(/~, N '---= 235 generations, the]'(~ arp el'rtaillly 80me stJ'iking alteratio1l8
J·ll11.
(', J +" ..., I
, ' " \ ::i.• ,) , __ 7 C)7 of frequency, [,specially in Ch18:-il'S 1 and :;. '\'1Ip type of l,ar
X , 1 L + LH I'S.!{ S- .. .011;; > p> .oot
lobes was rt·markahly cOll"if;tl'llt fl'0l11 gC'lll'ratil)][ to gCll[~ra­
(Y,,!e, tOrr.)

tion; an(l tl\() sanH' r,tay hL' saill of thc~ [']'('ljl1PIICY Dr 11i,.;ta 1
hypcrextcnt3ihility of the 111111nl>". ]\nt 11'1I;2;tll of till' 0th digit
'rahle :l ~lIows that for tlw Hit ))looll typC'i'J there is no in C'OllllJarisoll with ille ±( b, and thl~ [>1,(,";1'11<:1' 01' <l lllt'dimi
stati'itically significant drift, altholl,glt tlw fl'cqllclIcios of R1R" uppel' llia8telila ])et\\'cl~1I tlw illc:i8or (ul'tlt sl!o\\'c'd more vari­
(eDID) ancl Hz (eD:B~) }lhenotypes ali!' I' so erratically that ation, although ill the lattl~1' ea"c' ouly at tIll' 1>onlc,l' of
lIad the population been slightly la rgcr they would be so. stati"ti~al sig·nificance. Til Ow ea"c' of t1le length of the 5tll
In this ease the trond in the seco1ll1 gCllcration reversed the digit, the saIlle kind of shift appc~ars as sho\\,c,d 11]l ill n,gard
direct ion of tl18 trnnd OccllT'ri ng in t he first genel'U tion.
to handedness; the first: Hl](l seeol1(l gC'lwrations are alike,
Table 4, the analysis of the frequencie" of left anel right but the third generation is qni te lli f'fel'C'nt.
hallCledness, offC'l'" yet unother type of shift in gene frequen­
Tn short, over the interval of three generatiom, some geJll,tic
(~iL'S. 'rhe first and second generatiolls are virtually identical loci in the Dunker populatioll exlli1)it l'alHlom gelletic llrift,
in their frequellcies, whereas the third gcneration departs
SOIIle remain stahle', some shift auel return, ami some shift
 'l'A TILE 5

]?nwl.;lin COIt/Lty (J>a.) lh('JlJ,;cn~/--A[)(; g)"OIl]) (lIw):;,,>.i.:; GENETIC DRIFT lK TILMAN l'OPULATlOl"S 54!)
J. Mid-digital hair pattf'l"llS

CJ.AS~W:;
at one time but not at another. The picture is much as would
--'--'-~~-------

4
lip i\xpecteu in a small gouetic isolate (N '---' 90) where the
~\r o.
"
---------------------­ 'fO'1.'AT,

Gen. ] 2r,
% ,Yo. C;{, 11-0, {:{ Xo (/0 ..Yo. (;,~)
- yectorial evolutionaT,Y forces, and especially selection, are
,:'jS.:! B' 7,0 4 ~l.3
n(~n. :J 42 .ll.D HI 2:L-l n '7"..1-
9 ' 20.9
" 'LG ]] ]:,,6 43 sometimes overwhelmed by the rUll(lom scattering effed of
Gen. 0 50 :37.1 ~l
3 3.7 SI
21.4 ]2 12.~.) 9' 9.2 [is the sampling process, It may be pointpd 0[11 i1lat even thongh
I.----..._-~ " the gene flow into ilw isolate was estimate(l as amounting to
CUlil hi 11('(1

------
X~6 (CI:1SSCS :-3 .tinct ,j co,nlJln('(l) = 11.18

.10>1'>.05. 10-15~{) pOl' gCl1ondion, that is still in absolnte llumbers So

Higllcst t',ontl'ibutions to X·l.
:l
:\ 'J'};}

small a supply of g'oncs from the main population that it could

H.E,"· 101"'8 scarcely be a rellI'csellt at iva sample of the slllTo1md ing gellC
-------- -

--------~--
I,OBEl)
---'--"~ -~--'-
NQX-],()l\.h:D
'.1'01'_\[, (,{ :;";0)\'1,
pool, but must itself be subject to the vagaries uf small
0(;11 • .1 1U
samples.
GPll. 2
3K 4S 7}l.1
., 57 79
(ic'n. ,')
",
"
~4 ]1\
7:2.1 That llO such exaggeratecl fluctnations a11pc[tr ill snccpssin
71.7
'\ =, 20S
geupratiolls of largc' llOpu]ations is delllonstrable for some of
x', =, O.S~
.70> l' > .50
the S31ne genetic elw.racteristics. Hoorman ('SO) has shown
] 1 r. Die'''] Lq>cHxtl'n'il,11lt.l' of th" tllUlll),
how very similar the ABO aud Hh frequencies aI'O in 2,000
--._---, -- ----._-
Torc\.],
';{, JJT1T
comecl]tive LonclLm mothers and UH'ir 2,000 infants; and
(i-('Il. -J

~
23 ;j 2(;
11.5
Beyce et n1. ('fiO) have found ill(> salJW cOllsistency ill tho
Ot'lt.
:.:!I ;)
(;('1:. ::
";, II
H
;:J,;
11,.1
);'.11
ABO frequencies of 7,856 Anstralian 1I1(dhc~r-infallt pairs,
23,583 blood donors, niH] 176,943 ll1cnnbers 0[' the defeJlse
X', «;t'll. ] +2 "S. :J) ~c \I.:W
.70> p > .50
: \ -'---:c: .123
forces. This constancy, in fact, existed ill spite of tlip slatis­
:1 V. LI-'lJgtll of [ith digit I
tical1y significant excosses or' dODoicllcies of c('J'fnin types of
infants born to mothers of a givell 1.)oocl group. It is,
1'1)'1'\ T
.1."\'u. --_._._--~_._- -"- ---- ­
cOJlsequently, clear that 111c ex tnml I; t1nctnatiollS of gene
l'{o. (,''0 ~'(J. f;
( Jell. 1 S ~1;J
GCH. 2 ;; (;2
IiLIi S 1G.7 ~II frcquencies in the Dunkel' cOIllllHmity o\'c'r th\' 11\]'(,,(' genera"
7<l.l;
(-~"ll. a \I 7G m.o
1I \ I.a 81 tions stu(lied arc aHrilmtallle to random gelHdic cleift.
:.3--1 :11.0 110
x', (C,'n. ] +
2 \'8. :1) = ]~.G5 .005> P > .Oi)]
x', (SaJlle iglloring ela,,) ~o 0.14 + .0:25> P> .111 ~= ~3D
TIts interplay of [Jenetir: drift (//l!l selection
l,t--. fIJdicah.'S GIll l11git lungt·r th:lll juillt 1Jtt'IVl~('U millel1!] 111Jd (lbtul seglllf:'llts of
4th l1ig'it (llistal joint).

-- IllIli('.atl'::::i 5th digit Hhol't('T' th~lll (lil"'\tiil jOlllt.

In the long rJlJl \\'8 must cxped eVOlution, insofar as it is
= bulitlltl's 5tll dig'it. eXlending to llist:J1 joint..
adaptive, to he deterrnined )y se1odion. H anclom g'enetic
V. Mcuian upper diastema drift can scarcely do more thall lteightCll the store of genetic
---------~~
+
'rOTAl, variability present in local clifferentiatc(l Iiop111ations upon
Gtm. 1
Gen, 2
41
G7
2 J:J [),).-±
which inh~rgroup selection may act. That such variability
7 74
U('ll, il ~(j
20 ]OG
91l.5 may likewise exist ~when a trait is ol)viollsly snhject to strong
81.J
N = 22:1
selection may be illustrated by the frequencies of the sickle­
x\ (Gen. 1 a!lel 3) = 3.77
(Yates torr.)

.10>P>.05 cell trai tin varions African trihe8. Fov et a1. ('54, 'fiB), among
X', (Gen. 1,2, 3) ~~ G.70
.05> p> .025 others, have publisllcc1 such data, shOl\'ing not only great
548

 550 BENTLEY GLASS
\'Cniations in the fn'qllelll;y of sickling, as Wl'II as in the ABO
fn>tInCneit·", alllong diffcrent trihes, Imt (~"en "in subgroups
within the sallW tribc, althongh such :mhgronps speak the
same langnagc, ha"e tile salllC enltnl'al and soeial chal'actcl'­
i"ties, and now illtl~J'lnany fret'ly." 11 SCt~lllS that this case
affol'lls another l'xmnple of the iutcqllay of gcnetic dl'ift aud
sull~djon, although it lllay he impossible ill dealing with a
lJistoricaJ sitllation ('\'('L' to analyze it completely a 1\(1 satis­
fac10ri ly.
[\;0 Oue has lllOl'e dearly jlereeived thc neeessity of dealing
wit h the injnl'play hl'twecn g'enct ie drift awl :wleeti\'(~ forceR
in analyzillg adlwl gCllC !'rclineney "itnatiollS than Alice 11.
Bl'ut's ('5..J-).In a strikingly ul'iginal Illlaly"is she has ntilizetl
llw Vl,l'y exislellce oj' tht· randolllizing cfJl'ct of genetic dl'ift
to Ilostlliate the lIIagllitudt>s clud dil't,etions of the sl'n~l'al
sul(wtin~ fOl'ces which act llpUll the lllllllilU ABO hlood gronp
t'n'i}II(,llcies in slwh a Wily that, inst(~cld of all possible COlll­
llimd iOlls of fl'l'qnencie" Ileing ]'('alihl'll, only a limited, rat hl'1'
slllall I'ange of fn'i}lll'ncies of A, H, awl 0 ill adnality OCCnI'R.
Nlw llOtl~::; that "!Iw etrt;cts of gt,netie drift arl' ::;trongly
eviclent ill l'(~Sr)('et to tlw A-B-O gl'nc' f)'()(jlwllcies within
]JOlllllations (Jtht~rwise distinct awl relatin·ly hOlnogelwon:,,"
e.g., tlw North AlllPrican Inclians, PI'e-Dl'aI'illian" of Intlia,
l~skilllOS, PolyllC'sians, AlI"lralian, and Aillll, The range of
A frequencies wit hi II these othcl'wise quite homog('neous
groups is half or lLlorc than half of the total wol'1(l ral1g'e
- in fact, the North American ImlianR bracket the entire
world range - and it appears quite impossihle to fWCOllllt
for Rncli (lifferences on the lmsiR of gl'OllP intermixture and
hyhriclizatioIl, or of selpctioll 11asecl on differenc"R iu environ­
ment. Indced, "tlIe evidrnce for the "igor of geudic drift
uncleI' more primitive conditiOlls is so convincing a,s to be
disturbing. If it has acteel so strongly even after the formation
of certain raees and subl'aces as we now know t hl)ll1, why has
it not produced, dnring the wholl' perio ll of the cliffel'entiation
of the species, often untlcl' eonditions of isolation and small­
ness of numhers more marked than have existed subRequenily,
t;EXJ<;TIU IJfUFT IN HUMAN l'OI'ULAT10NS 5,")1
a range of variation nlllch grea tel' than we now setd" Only
tlIe differential ueiion of suleetion on ~1, H, aIld (j Sel'llIS
adequate to cxplain tlle n:stl'ictioll of the fl'l~quenci0s, sillce
the time in generations since llwse alleles originatod is eloady
far morc than enonglr to havl~ perllliUt;d eOllljlluln"nllldollli­
zation in the allsence of selection Pl't;sslu'es. 'l'ht( supposed
non-auaj)tive natIlre of the various blooll gTonjJ systc:llls ean
tlien he dismissod a" a fiction, even wpre tlwn: no inll'"I)()nllent
evidtmee of t1iH'erelltial fel'tility, materual-fc;tal ineolllpati­
bility, and the Ii lee.
[Possibly Ii ~ystelll of lllutation pl'CSSm'efl conld he devis(~t1
Owt would bl'illg' abont the sallle fretluellc:y distrilJllt ion of
the ABO alleles as aetnally exists, and Miss Bnws has ignored
this possibility. 11ow(:ver, at jll'esent we Imow nothillg allo11l
the mutatiou l"Hh~s of the ABO allelefl, allll eau do no hettt'r
than interpret the HTwlysis in terms of selection and/or
umtatiOll pressurc~s, I
The faet that tIlt: A-B-O freqneuey dislrilmtiou for all
populations is restrietetl and lIaR a single center implieR,
as T\Ji~s Bl'llCS points out, tlmt he1erozygot(,S are ill general
more favored than hOlllOhygOttlS. l1 Ill'O\'t~cl jlosRible to arri"e
at a satisfactory mathematical model to explain the existing
family of ABO geue frequencies hy postnlating a 2;)70 loss
of OA hcterozygotes tlIl'ongh maternal-ft>lal illcompatilJility,
and selt:etioll preSSUl'es of O.Oj~) for 00, + Iflro for OA,
+ 5% fOl' OR, - G% for AA, + 3% for riB, amI - 4% for
BB. This vectorial system foeuses upon tite point at whieh
the respective frhqncncies of A, B, ami 0 are 25%, 15/~), and
60jc, the vecU)rs being yury small neaL' the focus and larger
and larger as distauc(~ fruH! the focus increases. Miss Ernus
points out that "any ~nch vector system ... will he always
in (('rHct iug wit h gellU t ie dri f1. Small 0 r isolated popula t iOllS
will lw contilllwlly scattering as the result of drift, but ill
more copions populations . . . genetic drift will he quite
inutl'eetive, a]](l s()]eetive )lre~SUl'es will act with grlmt con­
,
,,-;jancy. All such populous groups should then tend inexorahly
to move toward the final equil ihriulIl point. . , . Most, ..

;):):2 BK1\TLEY GLAS:::;
major popnlaiion groups, ~White, .l\fongoloid, and Negro alike
[[tan] vcry low vectors .... '1'hc primitive condition of wide
random nl1lge of IJlooJ group frequencie8 is not too far in
the 11ast to aCcoullt for the diversities of the now more
1JOl11110us groups, even though constant selective faefors may
have he en standal'<lizing blood group frequencies since popu­
la Iion levels increased." This is imleed a refreshi~lg approach
ill contrast to that ov(~remphasis either on genetic drift or
on selection which has so characterized the "genetic drift
cOllh'oversy," and which cannot fail to remind one of the
sterile "heredity 01' environment" controversies of an earlier
period. Miss Brues provides a very apt analogy in likening
the interaciiOJ] of genetic drift and selection to the respective
efTects of Brownian movement and gravity on Suslwllded
1Ja rticles of different sizes. ~\V ould anyo]lf; suppose tll11 t
because gravity aets on suspended particles tll(! cffects of
Browlllall rnoven](~nt are inconseqnential? 'rhat depends on
the size of tlw particle.
Genetic drift as an C /'01 utionary fact or
It remains to consider whether the va riety of gelle fre­
quencies established in small populations hy random genetic
drift may subsequently have any significant role in evolu­
tionary processes. I have previonsly snggested (Glass, '54)
that in the human species "explosive increases in population
size, such as took place with the advent of agriculture, might
allow the particular genetic characteristics of one isolate to
become stamped upon a population too large to be swamped
when absorbing smaller groups." That idea has been attrac­
tive to many anthropologists. Brues ('54) likewise agrees,
saying, "Later increase in size will not reverSe any
peculiarities acquired while a group is small." But evirlence
to support this view is not forthcoming at present from the
field of human population genetics. Instead, we must appeal
to tlIe Drosophila geneticist. In studying the selective action
of exposure to DDT' upon fruitflies, King ('55) found that
~ - ')
GENETIC Dl\IFT IN HUMAN 1'0PULATlONS ~)~)t-)
each of two selected lines developed its o\\'n uniquo integrated
polygenic system. I am permitted IJy his kindness to quote
from a paper still in press and to discuss the implications of
these oxperim(mts in his own words. "'1'he faet that each
of two selected lines developed it~ own integrat ell polygenic
system witllin a dozen generatioIl~ leads oTle to snspect that
there arc more than two snch systems ]Jossihle, prohably
many. ~Wallace and King ('51, '52) discovf>.red in their work
with irradiated and control ]Jopulations that a closed popu­
lation very ra]Ji(lly develops a uniquely integrated genetic
system wlIidl thon tends to persist. Y cry prohahly this is
whal lJUppene(l in each selected line. '1'he manner in which
a line coul(l rosIJond to selection was to sortle extent lloter­
mined IJy Ow gcmetic nature of the sample frorn which it was
started, and, having started alOllg Olle c(~rtain road, it kept
on. '1'110 inevi table sampling error which occurs when a line
is taken from a larger ]Jopulation is vory likely the anlage
of the genetic individuality of the line. 1'his is, of conrse,
Hn example of the principle of random gendic drift, and its
operation can easily he envis:lg'e<l without insisting that
sampling error CHn ontweigh selection eitlIeI' in general or
in a single generatioJl in a going pOlJlllation. All Umt is
necessary is for the new population to be deuied access to
the larger gene pool of which it was formerly a part.
",Ye obtain here, 1 heliove, some insight into the nature
of the creative aspect of evolution. Every slwcies has within
it potentialities for adaptive radiation so vast that no fillite
popula tion erm \actnalize them all. As long a s the populatioll
remains a substantially panmictic unit, the random elJb and
flow of genes will result in a variability maintained within
limits determined by the adaptive norm, ~When a sample from
the larg'er population is shut off from the whole, the degree to
which and the manner in which the frequencies within its
gene pool depart from those prevailing- in HJP larger Ullit
exclude some potentialities entirely and hy this very fact
increase the probability that others will be actualized. ,Vh(,n­
ever a new elosed population is set up, a gamhle is taken

 ;).).J
BENTLEY GLASS
GENETIC DRIFT IN H1HfA N POPULATIONS 5;15
which mny result ill llothing of consequence or which may Foy, 11., ATIH;NA KmmI, t;, L. '1'L\L\IS, W. HHASS AND F. HC;SllRA 1%1 The
l'ngemlre]' a strikingly new creation." variability of ,i,,!de-ccll rates in tile trillUs of l\CIIY:1 :Jnd th e wnth"rJ[
In thii::i way, then, we may pieture the cn~ative l'OJe of Sou,]all. Brit. Me<1, .r" .lfJ54-/,' ~94-~n7.
ULAS~, H. 10:)-1 (h~1Il'tie C'hnng(~s in lllUUfW pupnlntlons, e::\llc('irdly those (luc to
1'l1nrlom genetic drift. Liktl the randOlnllcss of mutation, which g(~nl' iiow aud g(,llL'tic drift. A(lvml('('~ ill GeIl(~t., (j: 9;')-1;1D.
IJrodUCCi::i a h1l1l<lrcrl or 11 thousa11(1 deleterious g'elws 10 one GLA~8, D.) ::,r. ~. SACKS, r---';r,SA }\ .LJ\.II),...r~~~D C. HESS In~)~ Gl'lldie drift, in <l
that aids the orgallism in its struggle i'0!' survival, genetic fvlig-ioHs ls(l!Hh': nIl anrdysis of tIl(' ('al1~(':-l of Y:ni:dioll in 11100(1 g'l'CJujl
drift may estahlish a lltl1lrlrcrl 01' 11 tl10llsallrl delcirel'ious <llld On\(~r gelle fl'eqllt'Jl('iC':-l ill ;t slllillI population. Alli. ;\;11., sa:
J.lJ-IUII. HV['l'illlel1 in Y"il]'I",,,1t of l'll}s, '\111\"'0]'" 1!l;)~: ]·14-1,)H,
lllntations, fated cvelltnally to be eliminaterl, heforl' it ai(ls l\lKG, .T. C. Hl;l;) E\'i<1cllce for 111e i1ltpgl':ttion of Ole gl'lliJ pool from ~t\ldil'~
llIaterially in tlte initial estahlii::ilnl1cllt of' a lI111tatiOll of value. ot' IJDT l'L'si:::;tmH'c ill !ho>iopltiln. Cohl kpriJlg H<trlJUr t),YHl!l0:sia U1I
And, like l'eeollllJinat ion -- illdce(1, through reeomhiw1tion­ QU;Jlllibtiw Dio]ogy, .?o:
H1F'E, D. C. 19~I "''['edity :1l11] h'l!I(I"IIIlC". Hcil'llt, ~loJltltl'y, 7;1: ISH-IDL
genetic drift flj'(lVir1(~s those lIniqlle gCllOtYP(~i::i that clwrac­ t;lIEPI-'_\RV, P.!'I1. 195:; l'OlyllUIl'pllt1:l111 .'lnd POIJu1ntioll htu(livl:-i. S~'ll]PO~. ~()l'. Exp.
terize small (~Joserl popnlations, and UpOIl which seleetioll Hicl1., /: ~7 --!-~Hj).
(:a1l a(~t \\'ith so 1111lch rid\(~r a variety of outCOlll(' lhu1I when '\'-U,L-\(;E 1 B., A~'D .1. (1. KI~G 19,")1 Ot'lletie ('lJ:l]lgC~ ill Jlopulnti()n~ uIllter
irr:llljatioll. ~\lll. S"L, Sf): :2(\~>-~'2.~.
il is limited ill malol'ial to the IJlwnotypic llllifonnity of the
}0;)2 .A genctil' alli\l'y~i:-:i of tIle <\\lnptiv(' \'nlul'~ ut' vopnlatiol\:-I.
large, ]Hllilnietic pOIJulatioll. 1n Ne\vall \\'right's own pic­ Pl'{)('. Xilt. Acad. i-·h·i., \\';\1:,;]1., SS: TOG-I];).
turesqne imugcry, the way from onr' adaptiv(~ peak to anolher \VR1GH'I', N. H1451;1 AdHptatltlll :tlId ::-:c']edioll. Tn (if II ('iir's, l'alrnll(o{n!I!J, lO/(!
Rl'olutLon (C;. L. ,1 C' pi:'; l'll , E. ~l;l.yr, awl C;. C1:. 8illlllS0I1, ed~.:), IIp.
lear],; tllro ugh the vaHcy. The shadow of I'wl('et iOll hro(J(ls
:iti::i-;-1SU. Prlli\'l;1011 Un]\'. l'l'('~~l Prl1h'l'tfJH.
dl11'kly ove1' the evollltiollHI'Y \l'uy, tll\d ollly the stnlllg or 1H--!9h Poprdntion ,...,trudul'l' ill C'n!lutioll. Pl'(l(~. Am. Pllilos. Ho('.,
tltr' fortnnatr~ e,,;caT)(', ollly those forearHle<1 against the ,~:i: 4,.1-178.
exigencies they meet. f:jtalJilizing sel('ct iOlI adapts mOre lD;")O P\.lputltioJ\ ~trlldur(' H~ n 1'n('tor ill eVU1J1UuIl. III J/udcrl/c
lJiolo,Oic: Ftstsc1u'i/t Zl/fi~ DO. UdJIU'18!O[f lJmis ~'laeht.'dwim, pp.
T)(~rfectly ollly to those cOllditiolls that lwve cxisted up to
27:,)-:':S7. P. \\~. Pdl'l'~, Berlin.
HOW. It may ho that solely among the lebs llighly adapted, 19,11n 1'}lc g\.!llL'tieHI ~tl'\lC'tlll'e (If po}.lul:1tiOlI~. Alll!. EUgf'll'j 15:
i-;lI]u]J isolated ]lopnlntions there will oceur the genotypic 3~:j-:J:)4.

19[)11, Ji'islIL']' "1Ii] l"or.j on "The 8c,,",,11 Wl'igl,tf;ffeet." AIll,

system that fits the lIew cOllc1itiolls. [11 helping to crcate a
Beirut., 8,1): 452--4;')8, 479.
(1iversity of such sysh'11Is, l'al\(lo111 gc'n(·tie (hift lilays its part.

LI'rEHA'I'HR1~ ell'Ell

Boom,IAN, KA'I'lJLEEN E, 1950 .\n antilysis of th,' lil 0 0<] types anel clinical
eondition of 2000 consecutive ItlOthrn; Hu(l tlH!ir infants. Ann. El1gl'11.,
15: 120-134.
HI\UEB, ALICI~.r..r. 1954 8elcc-tioll :tlHl P(JlYIllorphisUl in tIlt, ...:\ D-O 1)}00<1 gTonps.
Am .•J. Pltys. Alltltrojl., 12 n.s.: 559~597.
BRY('E, Lvcy M., HAeIIEL JAKOllOWIC~, NORMA ~lcAR'rlTUR AND L. S. PENROSE
1!150 Blood-groujJ frequeneie" in a mot1",]' all'] inf:mt SIlIllP1c of the
Au~trnli<ln poplllatioll. Anll. Engen.! 15: ~7]-27G.
FISHEl{, H. A" AND E. D, Jo'rJ]{]) 1947 Tlte '['read of a gcne in n,\turn) (,ol,,]ilions
ill n, ('01011)' of tIle moth Pana:da llo}f,inula. l[C'n~aity, 1: ]'1:~-174.
1'0\, n" \Y, BRAS~, G. I" 'J'nnrs AND H. A. ~IoORE 195~ Nj('kling ancl malaria.
Brit. MUl1. ,J., 1fJ55-[,' :!8;)-~8G.