Immunoglobulin Haplotype Frequencies in Anabaptist Population

Transcription

Western Washington University
Western CEDAR
Anthropology
Social and Behavioral Sciences
2-1996
Immunoglobulin Haplotype Frequencies in
Anabaptist Population Samples: Kansas and
Nebraska Mennonites and Indiana Amish
Joan C. Stevenson Dr.
Follow this and additional works at: http://cedar.wwu.edu/anthropology_facpubs
Recommended Citation
Stevenson, Joan C. Dr., "Immunoglobulin Haplotype Frequencies in Anabaptist Population Samples: Kansas and Nebraska
Mennonites and Indiana Amish" (1996). Anthropology. Paper 7.
http://cedar.wwu.edu/anthropology_facpubs/7
This Article is brought to you for free and open access by the Social and Behavioral Sciences at Western CEDAR. It has been accepted for inclusion in
Anthropology by an authorized administrator of Western CEDAR. For more information, please contact [email protected].
Immunoglobulin Haplotype Frequencies in Anabaptist Population Samples: Kansas and Nebraska
Mennonites and Indiana Amish
Author(s): K. MARTIN, J.C. STEVENSON, M.H. CRAWFORD, P.M. EVERSON and M.S.
SCHANFIELD
Source: Human Biology, Vol. 68, No. 1 (February 1996), pp. 45-62
Published by: Wayne State University Press
Stable URL: http://www.jstor.org/stable/41465452 .
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Immunoglobulin Haplotype Frequencies in Anabaptist
Population Samples: Kansas and Nebraska Mennonites
and Indiana Amish
M.H.CRAWFORD,2
P.M.EVERSON,1
K. MARTIN,'
J.C.STEVENSON,1
ANDM.S.SCHANFIELD3
fisis a chronicle
ofrepeated
Abstract Anabaptist
migrations,
history
The
effects
of
these
events
of
various
and
fusions
sions,
subgroups.
No
inthepopulation
shouldbe evident
groups.
biologyoftheAnabaptist
andhighlyinformpriorgeneticstudieshaveincludedthepolymorphic
1
markers.
ativeimmunoglobulin
Here,685 serumsamplesrepresenting
were
Amishand 3 Mennonite
community
samples(7 congregations)
IGHG*F B,
studiedfor immunoglobulin
allotypes.The haplotypes
from0.542 to
IGHG*A,ZG, and IGHG*A,X,ZG rangein frequency
IGK*1frequen0.765,0.123to0.290,and0.075to0.170,respectively.
are withinexpected
cies rangefrom0.035 to 0.077. All frequencies
andwestern
samples.Therewas
Europeanpopulation
rangesforcentral
considerable
samplesthat
variability
amongtheAnabaptist
intergroup
was statistically
(%l = 22.63,0.005 < p < 0.01). Principal
significant
theimmunoglobulin
frequencies
allotype
including
analyses,
component
demonstrate
dataonABO,MN,andRhesus(Dd) markers,
andpublished
tiesgroup
thattheMennonite
sampleswithclosehistorical
congregation
fromtheAmishand Meridiancongregation
and are distinct
together
samples.
to as Anabaptists,a label appliedto
The Amishand Mennonitesare referred
of each
butarose independently
shared
certain
sects
that
principles
religious
centuries(Hostetler1980; Smith
otherduringthe sixteenthand seventeenth
and Krahn 1981). Amishpeoples have receivedconsiderableattentionfrom
social scientists[e.g., Cong (1992), Hostetler(1980), Hostetlerand Huntington(1971), and Kraybill(1989)]. Theirdistinctive
way of life,isolationfrom
the outsideworld,and lengthygenealogiesfacilitateresearchby demographers(Cross and McKusick 1970; Ericksenet al. 1979; Espenshade 1971;
1
WA98225.
Western
ofAnthropology,
Bellingham,
Washington
University,
2Department
KS66045.
ofKansas,
ofAnthropology,
Lawrence,
University
3Department
CO80231.
S.Drive,
7808Cherry
Creek
Genetic
Center
#201,
Denver,
Inc.,
Testing
Analytical
v.68,no.1,pp.45-62.
Human
1996,
Biology,
February
48201-1309
State
© 1996Wayne
Press,
Detroit,
Michigan
University
Copyright
STRUCTURE
GENETIC
KEYWORDS:
ETHNOHISTORY,
GM,KM,MENNONITE
ET AL.
46 /MARTIN
Hammanet al. 1981; Markle and Pasco 1977; Pollack 1978; Smith 1960),
geneticists(Jacksonet al. 1968; Juberget al. 1971; Kostyu et al. 1974;
McKusicket al. 1964; McKusick 1978; Morganet al. 1980; Wardet al. 1972),
as groupfissioning
and otherswho studysuchaspectsof populationstructure
and inbreeding[e.g., Hurd (1983, 1985a,b)]. Mennonitesare examinedfor
thesame reasonsand are also studiedby behavioralscientists[e.g., Boynton
(Allen and Redekop 1967; Ev(1986) and Redekop (1989)], demographers
ersonet al. 1995; Heaton 1986; Lin and Crawford1983; Stevensonand Everson 1989, 1990; Stevensonet al. 1989, 1994; Yoder 1985), and geneticists
(Allen 1988; Allen and Redekop 1987; Brown et al. 1974; Crawfordet al.
1989; Moore 1987; Rogers 1984, 1987; Stevensonet al. 1990).
relativeto the gestudiesthatexamine ethnohistory
Interdisciplinary
netic structure
of a populationare particularlyrevealingbecause cultural
eventscan be assessed in termsof theirbiological impact(Crawford1973).
are easily ascertained,have
The best geneticmarkersforsuch investigations
a simple Mendelian inheritance,and are highlypolymorphic(Schanfield
1980). However,only a few studieson Amish and Mennonitepopulations
have providedsuchdetail(Crawfordet al. 1989; McKusick et al. 1964; Rogand highly
ers 1984), and no priorgeneticstudiesincludedthepolymorphic
markers.The inheriteddifferenceson human
informative
immunoglobulin
(includingtheGM and KM markers)are anthropologically
immunoglobulins
usefulbecause manyof thehaplotypesare able to serveas uniquepopulation
markers(Schanfield1980; Schanfieldand Fudenberg1975; Steinbergand
are distinguished
Cook 1981). Populationsthatare in closerproximity
by less
in haplotypefrequencies.The objectivehereis to present
markeddifferences
the firstimmunoglobulin
haplotypefrequenciesfor Amish and Mennonite
context.
populationsand place themin ethnohistorical
Populations
sects are derivedfromtheAnaThe Amishand MennoniteProtestant
in sixteenth-century
baptistmovement,whicharose duringtheReformation
Switzerlandand spread eastwardinto what is now southernGermanyand
Austriaand northwardinto the Netherlandsand northern
Germany(Dyck
1981; Hostetier1980; Smithand Krahn1981). Anabaptisminitiallyconsisted
of manylocal factionsthatorganizedaroundcharismaticpersonalities,such
as JacobAmmann(Amish)andMennoSimons(Mennonites).All Anabaptists
rejectedthe impositionof a statechurch,infantbaptism,and militaryconscription,whichput themat odds withmostEuropeanleaders.As a result,
theirhistoriesare characterizedby a series of persecutionsand migrations.
Figure 1 representsthe migrationsof the Amish and Mennoniteswithin
Europe.
Ig HaplotypeFrequenciesin Anabaptists/47
inthisstudy.
1. Principal
Dutch
andnorthern
Mennonite
involved
Figure
Germany
migrations
and
TheAlsace
andPalatinate
areasaresources
oftheearliest
(Mennonites
Anabaptist
inthelateseventeenth
andearly
totheUnited
which
States,
Amish)
migrations
began
centuries.
from
Bender
andSmith
(1956,
p.257).]
eighteenth
[Adapted
WilliamPenn,granteda charterby King CharlesII of Englandto form
and invitedthemto
a colonyin America,knewof theAnabaptists'suffering
relocate fromthe Rhine area to Pennsylvania(Fisher 1908; Peare 1957;
Schwiederand Schwieder1975). The firstAnabaptistsettlersarrivedin 1683,
in two waves: 1727-1770 and 1815-1860 (Smith
buttheAmishimmigrated
and Krahn 1981). Eventually,the Amish moved west intoHolmes County,
Ohio, after1812 and intoLagrangeand othercountiesin Indianaduringthe
in theElkhartand La1840s. The Amishof thisstudyrepresentsettlements
grangecountiesof Indiana.
Many Dutch, German, and other Anabaptistsfled east to Polandcontrolledareas around Danzig (Gdansk), and in 1699 eighteenfamilies
formedthePrzechovkachurchlocated 60 miles southof Danzig (Duerksen
1955; Rogers 1984). Prussiatookcontrolof thearea in 1772 and limitedthe
service(Frieavailabilityoflandand reexaminedtheexemptionfrommilitary
ET AL.
48 /MARTIN
sen 1989). CatherinetheGreatinvitedMennonitesto settlein southernRussia; migrations
began in 1788 and theChortitzaColonywas foundednearthe
Dnieper River. More Mennonitesmigratedin 1803 and foundedthe MolotschnaColony in Taurida Provinceon the MolochnayaRiver. Almostthe
entirePrzechovkacongregation(21 families)joined theMolotschnaColony
in 1824 and foundedthe Alexanderwohlchurchand settlement
(Duerksen
the fission-fusion
1955; Rogers 1984). Figure 2 reconstructs
processesexperiencedby theMennonitepopulations.
The communitiesflourisheduntil land shortagesand the exemption
frommilitaryservicewere again jeopardized (Dyck 1981; Duerksen 1955;
to AmerRogers1984; Smithand Krahn1981). Aftersendingrepresentatives
ica in 1873, mostof thevillageof Alexanderwohlemigratedon two separate
shipsthefollowingyear(Sawatzky 1971; Wedel 1974). One groupfounded
the AlexanderwohlChurchnear Goessel, Kansas. As the churchgrew and
expanded,two daughterchurcheswere established(Duerksen 1955): Tabor
MennoniteChurchin 1909, built5 miles southeastof theparentchurch;and
Goessel MennoniteChurch,formedin 1920 in Goessel, Kansas. All three
are represented
in thisstudy.
congregations
Anothercontingent
fromthe Russian Alexanderwohlcongregationarrivedin Lincoln,Nebraska,and settledabout40 mileswestofLincoln,founding the Bethesda MennoniteChurchat the presenttown of Henderson,a
sample of which is representedhere (Crawfordand Rogers 1982; Rogers
theHender1984; Voth 1975). A secondNebraskaMennonitecongregation,
son MennoniteBrethren
Church,tracesitsoriginsto 1835, whenMennonites
in theChortitzaand Molotschnacommunitieswerejoined by Prussianemi(Crawford
greswho had been influenced
spiritually
bytheMoravianBrethren
and Rogers1982; Rogers1984; Dueck 1989). Theyformeda separatechurch
in 1860 and migratedto Americain 1874. In 1876 theyfoundedthe HendersonMennoniteBrethren
Churchin YorkandHamiltoncounties,Nebraska;
theyare also sampledin thisstudy.
Finally,disputesoverdoctrinalissues continue,and new congregations
oftenresult.In 1858 JohnHoldeman of New Pittsburg,
Ohio, foundedthe
Churchof God in ChristMennonites(Dyck 1981; Smithand Krahn 1981).
He recruitedhis membersfromdescendantsof Swiss and southernGerman
Mennoniteswho had arrivedin theUnitedStatesduringthe eighteenth
and
nineteenth
centuries.This sectis represented
by theMeridianchurch,located
nearHesston,Kansas.
Methods
This studyincludesblood samplesfromone AmishandthreeMennonite
communities,
representedby at least seven congregations.Biological relawere
knownfor the participantsof the study,and all first-order
tionships
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ET AL.
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UsedforImmunoglobulin
Table1. Reagents
Specificity
Allotyping:
Amish
Specificity
G1MA
F
X
G3MBO
B1
B3
B4
B5
C3
C3 + C5
G
G5
S
T
KM1
Agglutinator
PAN
STA
ALX
R-60
KEK
LOG
KEK
FIE
HEN
HAW
R-68
BRO
YAR
CRA
SINorRUT
Mennonite
Coat
PET
DAN
PET
PUH
ADA
HUN
HUN
PUH
422
422
SUL
SUL
PUH
PUH
PET
Agglutinator
PAN
STA
ALX
TOL
TOL
LOG
GOEL
FIE
HEN
HAW
R-68
BRO
YAR
CRA
SIN
Coat
PET
DAN
PET
PUH
HUN
HUN
HUN
PUH
ADA
ADA
SUL
SUL
PUH
PUH
PET
relativeswere removed.The 599 blood specimensforthe Kansas and Nebraska Mennoniteswere collectedas partof a three-year
multidisciplinary
studyon aging,which is describedby Crawfordand Rogers (1982). This
almostone-third
of theMennonitecitizensof Henderson,
samplerepresents
almost
one-third
of
the
Alexanderwohl
the largest
Nebraska,
congregation,
of
the
town
of
and
about
of
the
Meridian
Goessel,
congregation
one-quarter
The
86
Amish
blood
were
collected
as
of
congregation.
samples
part a twoon
the
association
of
HLA
and
viral
yearstudy
haplotypes
antibodyresponse
in a sample of Amish living in Elkhartand Lagrange countiesin Indiana
(Hsia et al. 1977).
The 685 serumsamplesweretypedforimmunoglobulin
allotypesat the
of
the
American
Red
Cross
Blood
Services
Immunohematology
Laboratory
NationalHeadquartersin Washington,
DC, in 1981 and 1982. Minimally,all
samples were testedforG1M (allotypesA, F, and X), G3M (allotypesBO,
B1 , B3, B4, B59C, G, S , and 7), and KM1 usingpreviouslydescribedmethods
(SchanfieldPoleskyet al. 1975). The reagentsused are presentedin Table 1.
In addition,some G1M (allotypeA) and G3M (allotypeB) positiveserawere
testedforG1M (allotypeZ) (see Table 2).
Haplotypefrequenciesfor the GM systemwere estimatedusing the
allocationmethodof Andersson(1985). The IGK*1 frequencieswere determinedfromthesquarerootof theKM (7 - ) frequency.
A heterogeneity
chi-squarewas used to measurepopulationdifferences
and divergence.
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ET AL.
52 /MARTIN
Principalcomponentsanalysesfacilitatedthe comparisonof ethnohisand althoughGM is a highlyinformative
torywithgeneticstructure,
system,
the GM data were supplementedby alreadypublisheddata on additional
geneticmarkers.The Mennonitesamplesof thisstudyhave been typedfor
additionalgeneticmarkers(Crawfordet al. 1989), butABO, MN, and Rhesus
(Dd) data fromanothersampleof IndianaAmishwere used in theprincipal
componentsanalyses.Bothprincipalcomponentsanalysestookintoaccount
populationsample sizes.
Principalcomponentsanalyses were performedusing antana (Harpendingand Rogers 1984). The programproducedorthogonalsynthetic
gene
frequenciesthataccountforthegreatestamountof populationvariation.Two
analyseswererun,includinggene frequenciesand samplesizes for7 and 15
populations,respectively.The 7-populationanalysisincludedthe following
systems:ABO, MN, and Rhesus (Dd) fromCrawfordet al. (1989) exceptfor
theAmish(Juberget al. 1971) and GM and KM data fromthisstudyforall
the Amish and Mennonitecongregations.The 15-populationanalysis includedGM and KM dataforthefourAmishand Mennonitecommunities
and
ABO, MN, and Rhesus (Dd) fromCrawfordet al. (1989) fortheMennonites
and fromJuberget al. (1971) forthe Amish. See Mourantet al. (1976) for
theABO, MN, and Rhesus (Dd) frequencydata forthe additional11 European populationsexcludingtheNetherlands(Fraseret al. 1974) and Table 3
fortheimmunoglobulin
haplotypefrequenciesand referencesforall theEuropeanpopulations.
Results
and
Immunoglobulinallotypeand phenotype(GM-KM) distributions
for
the
studied
Amish
and
Mennonite
haplotypefrequencies
populationsare
in
Tables
2
and
KM
1
and
4.
distributions
presented
frequenciesare givenin
Table 5. Table 3 presentspreviouslypublishedhaplotypefrequenciesof centralEuropeanpopulationsforcomparison.
The numberof phenotypesand thenumberof haplotypesrangedfrom
fiveto sevenand threeto five,respectively.
deviations
Statistically
significant
fromHardy-Weinberg
expectationswerefoundin theAlexanderwohlsample,
whichled to significant
deviationsin the Kansas Community(see Table 4).
In theAlexanderwohland Kansas Communitysamples(notrepresenting
the
Meridiancongregation)
occurred
theGM A G andGM A,X,FB,G phenotypes
more than expected and the GM A,X G phenotypeoccurredless than
expected.
The frequencyof haplotypeIGHG*F B rangesfrom0.542 in theMeridiansampleto 0.765 in theGoessel sample(see Table 4). The low frequency
is lower thanwhat is typicalfornorthwestern
Europe (see Switzerlandin
Table 3), whereasthe highestfrequencyof 0.765 is not unusualrelativeto
Table3. Comparative
Population
Haplotype
Frequencies
Haplotype
B
IGHG3*: G
G
N IGHG1
*:
A
IGK*I
Source
A,X F
Population
andMickerts
Austria
0.185 0.074 0.741 0.083 Mayr
(Vienna) 1602
(1970)
0.134 0.035 0.831 0.053 Schanfield,
Croatia
452
Herzog
etal.(1975)
etal.(1969)
CzechRepublic 684
0.135 0.079 0.776 0.060 Fraser
Bohemia)
(central
etal.(1964)
772
0.137 0.061 0.728 0.061 Ritter
Germany
(Freiburg)
0.169 0.091 0.740 0.059 Ropartz
etal.
(Munich) 131
Germany
(1963)
etal.
386
0.204 0.099 0.697 0.055 Ropartz
Germany
(1964)
(Rheinland-Pfalz)
182
0.145 0.051 0.777 0.073 Schanfield,
Gergely
Hungary
etal.(1975)
(Budapest)
etal.(1974)
Netherlands
792
0.189 0.102 0.709 0.094 Fraser
Poland
0.138 0.056 0.806 0.055 SochaandKaczera
(Krakow) 600
(1968)
Poland
300
0.168 0.065 0.767 0.070 Schiesingerand
(Lower
LuczkiewiczSilesia)
Mulczykowa
(1971)
andCook
0.253 0.119 0.628 0.080 Steinberg
Switzerland
98
(1981,
p.43)
typicalfrequenciesof thishaplotypein easternand southernEurope (Steinberg and Cook 1981; Stevensonand Schanfield1981). Croatia and Poland
exhibitslightlyhigherfrequencies(Table 3).
The IGHG*A,Z G haplotypefrequency
rangesfrom0. 123 in theGoessel
G haplotype
sample to 0.290 in the Meridiansample,and the/G//G*A,X,Z
frequencyrangesfrom0.075 in theHendersonsampleto 0.170 in theAmish
sample. The highestfrequenciesof these haplotypesare typicallyfoundin
northern
and westernEurope (Steinbergand Cook 1981), and therangeshere
for
theAmishand Meridiansamples) are a bit lowerand morecon(except
sistentwithwhatis typicalof easternand southernEuropeanpopulations.
Seventeenindividualsexhibitthe phenotypeslisted in the last three
columnsofTable 2. These phenotypes
cannotbe explainedsolelybythemost
common European haplotypes[e.g., IGHG*A,Z G, IGHG*A,X,Z G, and
referred
to as IGHG*F 5)] (Steinbergand Cook
IGHG*F B0,1,3,5 (hereafter
is
1981). Pedigreeanalysis necessaryto establishwithcertaintythe haplotypesinvolved.However,some of thephenotypesare probablytheresultof
ET AL.
54 /MARTIN
of Haplotype
in SeveralAnabaptist
Table 4. Distribution
Frequencies
Congregations
andCommunities
N
Population
Amish
(Indiana) 86
Kansas
Mennonites
Alexanderwohl
166
Goessei
49
Tabor
83
Kansas
266
Community
Meridian
60
Nebraska
Mennonites
Bethesda
189
(Standard
Error)
Haplotype
Frequency
Bb
Bc
IGHG3*: Ga
Ga
*:
A
F
IGHG1
A,X
Z,A
0.196 0.170 0.628 0.006
(0.030) (0.029) (0.037) (0.006)
0.131
(0.019)
0.123
(0.033)
0.143
(0.027)
0.131
(0.015)
0.290
(0.041)
0.110
(0.017)
0.092
(0.029)
0.092
(0.022)
0.104
(0.013)
0.151
(0.033)
0.750
(0.024)
0.765
(0.043)
0.753
(0.033)
0.752
(0.019)
0.542
(0.045)
0.009
(0.005)
0.020
(0.014)
0.012
(0.008)
0.013
(0.005)
0.0085
(0.0084)
B,^
Z,A
0.000
0.000
0.000
0.000
0.000
0.0085
(0.0084)
0.000
0.200 0.080 0.717 0.003
(0.021) (0.014) (0.023) (0.003)
Henderson
41
0.254 0.075 0.671 0.000
0.000
(0.048) (0.029) (0.052)
Nebraska
234
0.210 0.079 0.708 0.003
0.000
(0.019) (0.012) (0.021) (0.003)
Community6
a. GMZ nottested
forbutalways
associated
with
GMG.
b. Represents
thehaplotype
IGHG*F
B0,1,3,
4,5.
c. Represents
thehaplotype
IGHG*A,Z
B0,1,3,
4,5.
d. Represents
thehaplotype
IGHG*A,Z
B0,3,5,S.
e. Includes
a fewunrelated
ofanEvangelical
Mennonite
Brethren
representatives
congregation.
Table5. Distribution
andFrequencies
ofKM 1 andIGK*1
N
KM1
IGK*1
Population
N.Indiana
2
0.011
88
Amish,
Kansas
Mennonites
Meridian
0.027
56
3
congregation
Alexanderwohl
14
101
0.072
Goessei
7
0.054
47
Tabor
2
0.077
19
Kansas
17
0.059
148
Communitya
Nebraska
Mennonites
Bethesda
16
0.043
189
Henderson
41
0.037
3
Nebraska
22
0.048
234
Community3
a. Numbers
reduced
incommunity
from
sumiffirst-order
relatives
areincluded
pool.In
simple
individuals.
Mennonite
Brethren
theNebraska
includes
a fewEvangelical
addition,
Community
a combinationof one of the threecommonEuropeanhaplotypeswithhaplotypestypicallyfoundin otherpopulations,such as the centralAsian or
referred
to as IGHG*A,Z
AfricanhaplotypeIGHG*AyZBO,1,3,4,5 (hereafter
B ) and the AfricanhaplotypeIGHG*A,Z BO,3,5,S (hereafterreferredto as
IGHG*A,Z B,S).
The six individualswithphenotypesGM A G,B and thetenindividuals
withGM A,Z,F B resultmostlikelyfromthepresenceof the typicallyEucombinedwith
ropeanhaplotypesIGHG*A,Z G andIGHG*F B, respectively,
thecentralAsian or AfricanhaplotypeIGHG*A,Z B. The sampleswere not
testedfortheA2M markers,so thecentralAsian haplotype,associatedwith
fromtheAfricanhaplotype,associatedwith
A2M 1, cannotbe distinguished
A2M 2 (Stevensonet al. 1985). The IGHG*A,Z B haplotypeis foundin
centralEuropeans(Stevensonand Schanfield1981).
contemporary
The one GM A,Z B,G,S phenotypefoundin theMeridiancongregation
is probablydue to theEuropeanhaplotypeIGHG*A,Z G combinedwiththe
AfricanhaplotypeIGHG*A,Z B,S.
predominantly
IGK*1 frequenciesare variablein Europe withno apparentcline and
rangein centralEuropean populationsfrom0.035 in Croatiansto 0.253 in
theSwiss (Steinbergand Cook 1981; Stevensonand Schanfield1981). Some
of theKansas Mennonitescould notbe testedforIGK*1 because of insufficientblood samples. The gene frequenciesrangefrom0.011 in the Amish
sampleto 0.077 in theTabor sample.Overall,frequenciesare relativelylow
comparedwithfrequenciesobservedin Europe.Thereis no apparentpattern
of thisallele in Europe.
in thedistribution
Contingencychi-squaretestswere performedon the Kansas (xf6) =
and Nebraska(xf4)= 0.44, notsignificant)
2.59, notsignificant)
Community
bothcomwithin
differences
of
The
absence
significant
statistically
samples.
munitiesprecludedtheneedto subdivideeitherforthecontingency
chi-square
varanalysisof theAnabaptistpopulationsamples.Considerableintergroup
was
demonstrated
was
that
Amish,
(e.g.,
statistically
significant
iability
Kansas, Meridian,Nebraskasamples) (x%} = 22.63, 0.005 < p < 0.01).
The principalcomponentsanalysesare presentedgraphicallyin Figures
Five geneticsystems
3 and 4 forseven and fifteen
populations,respectively.
are used in theseanalyses,and populations,takingintoaccountsample size,
of the
are plottedby theireigenvectors,each weightedby themultiplication
eigenvalue(Harpendingand Jenkins1973,
squarerootof thecorresponding
p. 187). The firstthreefactorsaccountfor51% of thevariance.
In Figure 3 the seven populationsamples are plottedagainstthe first
two factors,whichaccountfor45.6% of the variance.The Mennonitecongregationsamples are relativelyclose to each other,withthe Meridianand
withtheotherMennonitesamples.
Amishsamplesshowingtheleast affinity
Tabor and Goessel lie closest
Alexanderwohland its offshootcongregations
to each other.Factor 1 separatesthe Amish fromthe othersamples and is
weightedmost heavilyon ABO*A and IGHG*F B. Factor 2 separatesthe
ET AL.
56 /MARTIN
1.0 ?
ez * ' z
-n
/0
?
i
1
0.5 -
o
Meridian!
?
I
j Henderson! °
o.o -
I
-
0 Amish !
i
-0.5
-1.0 °
-1.0
1
'
oBethesd^
i
Goesöel o
o
Alexanderwohl
Tabjor°
;
j
1
'
-0.5
1
0.0
v
1
0.5
'
®
1.0
C2
3. Least-squares
reduction
Figure
genetic
mapofthesevenAnabaptist
congregation
samples.
from
alleles
andfive
lociwere
usedtoconstruct
theRmatrix.
Frequencies
eight
genetic
Meridiansampleand to a lesserextenttheHendersonsamplefromtheAmish
and otherMennonitesamplesand is weightedmostheavilyon IGHG*A,Z G
andABO*A. Factor3 accountsforonlyanother5.1% of thevarianceand thus
was notincludedin thegeneticmap.
theresultsof theprincipalcomponentsanalysisof
Figure4 represents
the Amish and Meridiancongregations,
the Kansas MennoniteCommunity
rel(Alexanderwohl,Goessei, and Tabor adjustedforremovalof first-order
atives) and theNebraskaMennoniteCommunity(Bethesda,Henderson,and
a few unrelatedmembersfroman Evangelical MennoniteBrethrencongregation),and 11 centralEuropeanpopulationsampleswiththe same fivegeneticsystems.The first
threefactorsaccountfor88.5% ofthevariance.Factor
1 separatestheMeridianand Amishsamplesfromall otherEuropeansamples
and fromthe two Communitysamples and is weightedon IGHG*F B and
IGHG*A,X,Z G. Factor 2 also separatesthe Amish and Meridiansamples
fromtheotherEuropeanand Anabaptistpopulationsamples and fromeach
otherand is weightedon ABO*A. Factor3 is weightedmostlyon ABO*B and
to a lesser extenton Rh D and KM and explainsonly an additional9% of
thevariance;therefore
it was notincludedin thegeneticmap. The Nebraska
and Kansas communitiesare centrallyclustered with other European
populations.
Discussion
The low numbersof GM phenotypes(5-7) and haplotypes(3-5) for
thesesamplesreflecttheirhistoryof relativeisolation.Five to 7 phenotypes
1.0 ?
05
c *
e* X
~~
:
Meridián
Nebraska
Î
i 11
- -°
1/2
_q g _^ Q O
-1.0
Amishj
o
1
1
-0.5
<>
i
;
7j
°ji4 .10
• !«° °«7
65Ff ,V"Z
3°
!4
Kjansas
1
i
1
0.0
!
!
~
!
j
i
0.5
1
Ô
1.0
+ ^ 1/2
4. Least-squares
reduction
with
Figure
genetic
mapof4 Anabaptist
population
samples
compared
11 European
from
alleles
andfivegenetic
loci
population
samples.
Frequencies
eight
were
usedtoconstruct
theRmatrix.
(1)Austria
(Vienna),
(2)Croatia,
(3)Czech
Republic
(central
Bohemia),
(4) Germany
(5) Germany
(Munich),
(6) Germany
(Freiburg),
(Rheinland-Pfalz),
(7)Hungary
(8)Netherlands,
(9)Poland
(Krakow),
(10)
(Budapest),
Poland
Forreferences
refer
toTable3.
(Lower
Silesia),
(11)Switzerland.
is fewrelativeto the20 or morefoundin a tri-ethnic
hybridpopulationsuch
as the Black Caribs of CentralAmerica (Crawford1983; Schanfieldet al.
in European
1984). At least thatmanyphenotypesare usuallydemonstrated
populationsamples [e.g., Stevensonand Schanfield(1981)]. However,Anabaptisthistoryis a chronicleof repeatedmigrationsand fissionsand in some
cases fusionsof varioussubgroups.
The Amishbecamereproductively
isolatedfrommostMennonitesearly
in Anabaptisthistory.This is forcefully
demonstrated
by theprincipalcomponentsanalyses,in whichthe Amish are clearlyisolated relativeto other
populationsin thethree-dimensional
plots.Theyare distinguished
by thefirst
and second factorsin Figures 3 and 4, respectively.The extremesin gene
frequencies[includingrelativelyhigh frequenciesof ABO*A, MNS*N, and
RH d and comparatively
low frequenciesof AB0*0, ABO*B> MNS*N, and
RH D notedby Juberget al. (1971)] do not obscuretheirorigins,however.
Most oftheancestorsoftheAmisharederivedfromSwitzerlandand southern
Germany.The haplotypefrequencyforIGHG*F B is second lowestforthe
Anabaptistpopulationsof this studyand the haplotypeIGHG*A,Z G frequencyis relativelyhigh,but theselevels are similarto frequenciesin conSwiss and close to frequenciesfoundin contemporary
residentsof
temporary
southernGermanyand theNetherlands.
the
Amish
IGHG*A,X,Z
By contrast,
ET AL.
58 /MARTIN
residentsofSwitzerland,
G frequenciesarehighevenrelativeto contemporary
whichmaypartlybe due to thesmalland probablyunrepresentative
founding
population.
of the Kansas and NebraskaMennonite
Studyof the geneticstructure
communitiesand congregationsalso suggestsparallels with history.The
from
foundinggroupforthe Nebraskacommunityincludedrepresentatives
boththeRussianMolotschnaand Chortitzacolonies,butin generaltheethnic
originsof bothgroupsare essentiallythe same. As expected,the GM haplotypefrequenciesforthe Kansas and Nebraskacommunitiesare similarto
fromeach
thefrequenciesforcentralEurope and are only slightlydifferent
other.
The Meridiancongregationis a melangeof Russian and Pennsylvania
to be
GermanMennonites,althoughone would expectits geneticstructure
intermediate
relativeto theAmishand otherKansas Mennonites.This is not
in thehaplotypefrequenciesfortheGM system.The Meridianconreflected
gregationdisplaysthethelowestIGHG*F B, thehighestIGHG*AyZG, and
the second highestIGHG*A,X,ZG frequencies.This patternmay be due to
samplebias butis also suggestiveof a uniquehistory.
Overall,theseresultsare consistentwiththeanalysesof Crawfordet al.
noted
(1989) usingredcell antigensand blood proteins.The maindifference
is theincreaseddistinctiveness
of theMeridiancongregation'sgeneticstructurewhenGM is includedin theanalysis.
Conclusions
The presentation
of immunoglobulin
phenotypeand haplotypedistributionsforthe Anabaptistpopulationsamples do not reveal any surprises.
The patternsin thephenotypicand genotypicfrequenciesare consistentwith
expectationsbased on thecentraland westernEuropeanoriginsof thepopulationsinvolved.Theirrelativeisolationfromoutsidersis reflectedin the
low numberof haplotypesand phenotypes.
within
Contingency
chi-squareanalysesalso reveallittleheterogeneity
the Kansas and NebraskaMennonitecommunities.By contrast,significant
is evidentwhenthetwocommunities
are partof a contingency
heterogeneity
chi-squareanalysisthatalso includestheAmishand Meridiansamples.
Except forthe Meridiancongregation(whichhas a unique originand
composition),the Kansas and NebraskaMennonitesgrouptogetherand are
close to othercentralEuropeanpopulationsin graphicrepresentations
of the
resultssummarizing
theprincipalcomponentsanalyses(Figures3 and4). The
Meridianand Amishsamplesare outliersin bothfigures.
in partbytheNationalInstitutes
was supported
AcknowledgmentsThisresearch
to
thankDave Dentonof Western
We
wish
AGO
1646-03.
of Healththrough
grant
ofFigures1 and2, S. Hsia
for
media
services
preparation
Washington
University's
Red Crossstaffat theNationalHeadfortheAmishbloodsamples,theAmerican
thesamples,andtheAmishandMennonite
forimmunoglobulin
allotyping
quarters
withthe
We
thank
RectorAryaforhisassistance
in
this
who
took
study.
part
people
for
their
our
reviewers
and
helpful
suggestions.
analyses
components
principal
1995.
received27 February
Received22 December1993; revision
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Immunoglobulin Haplotype Frequencies in Anabaptist Population

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