@07

Assi
gnment
Assi
gnmentt
opi
c : Har
dy-
Wei
nber
gTheor
em

Submi
tt
edby : M.
Adi
lMehmood

Submi
tt
edt
o : Si
rNaeem Ahmad

Rol
lNo : 2207/
591605

Sessi
on : 2018-2022

Depar
tment : Zool
ogy
(Mor
ning)

Gov
tpostGr
aduat
ecol
legeofsci
enceFai
sal
abad

23/
10/
10
@07

Cont
ent
s
 I
ntr
oduct
ion

 Def
ini
ti
on

 Condi
ti
onsf
orHar
dy-
Wei
nber
gEqui
l
ibr
ium

 Har
dy-
Wei
nber
gequat
ion

 Si
gni
fi
canceofHar
dy-
Wei
nber
gPr
inci
ple

 Dev
iat
ionsf
rom Har
dyWei
nber
gTheor
em

 Random genet
icdr
if
t

 Nonr
andom mat
ing

 Recur
ri
ngmut
ati
on

 Mi
grat
ion

 Nat
ural
sel
ect
ion

 Ref
erences

23/
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10
@07

HARDY-
WEI
NBERGTHEOREM
I
ntr
oduct
ion

 I
n 1908 ,Bri
tish mathemat i
cian,Godf r
ey Hardy and t
he German physi
cian
,Wil
helm Weinber
gi ndependentlyderi
vedamat hemati
calequati
ontorepr
esent
thedi
stri
but
ionofal l
elesandgenot ypespert
ainingtoaspeci
ficgenel
ocusina
panmict
icMendelianpopulati
on.

 Later,i
n1929- 30,R.
A.FisherandJ.B.Hal
daneinEngl andandSewal lWri
ghtin
UnitedStat
eswor kedoutthatundercer
tai
ncondit
ions,thegenet
icstr
uctureofa
populati
onmaynotchangeov erti
me.Theyestabli
shedt heconceptof'Hardy
-
Weinbergprincipl
eofequil
ibr
ium.'

Def
ini
ti
on

 According tot hist heorem,the heredit

aryprocess alone does notpr oduce
evol
utionar
ychange.I nl ar
ge bipar
entalpopulati
ons,al l
eli
cf requenci
es and
genotypicrati
osat tainanequi li
bri
um inonegener at
ionandr emainconstant
ther
eafterunlessdi sturbedbyrecurr
ingmut ati
ons,naturalselecti
on,migrat
ion,
nonrandom mat ing,orgenet i
cdrif
t(random sort
ing)
.Suchdi sturbancesarethe
sourcesofmi croevoluti
onarychange.

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10
@07

Expl
anat
ion
Hardy Weinber
gl aw descr
ibes a t
heor
eti
calsi
tuat
ion i
n whi
ch a popul
ati
on i
s
under
goingnoev ol
uti
onar
ychange.

 I
texplai
nst hatiftheev ol
uti
onaryforcesareabsent,thepopulat
ionislarge,i
ts
i
ndivi
dualshaver andom mat i
ng,eachparentproducesroughlyequalnumberof
gametes,theal l
elef r
equenci
esar eidenti
calinmal esandf emales,andt he
gametespr oducedbyt hemat escombineatr andom andt hegenef requency
remai
nsconst ant,thent hegeneti cequili
bri
um oft hegenesi nquest ionis
maint
ainedandt hev ari
abi
li
typresenti
nthepopulati
onsispreserv
ed.

 Inmanyhumanpopul ati
ons,geneti
call
yr ecessiv
et rai
ts,i
ncl
udi
ngt heO bl ood
type,blondhair,andbl ueeyes,areverycommon.Whyhav enotthegeneticall
y
domi nantal t
ernati
ves graduall
y supplanted these recessi
vetrai
ts? Itisa
commonmi sconcepti
ont hatacharacteri
sticassociatedwithadomi nantallel
e
i
ncr easesinfrequencybecauseofi t
sgenet icdomi nance.Thi
smisconcepti
oni s
refuted by a princi
ple call
ed Hardy-
Wei nberg equili
bri
um,which f or
ms t he
foundationforpopulat
iongenetics.

 Ar are al
lel
e,accor ding tothis princi
ple,does notdi sappearf r
om a l arge
popul at
ionmer elybecausei tisr are.Certainr ar
et rai
ts,suchasal binism and
cy sticfi
brosis,persi
stf ormanygener ati
ons.Forexampl e,albi
nism inhumansi s
causedbyar arerecessiveal
lelea.Onl yoneper soni n20, 000isanal bino,and
thisi ndivi
dualmustbehomozy gous(a/ a)f orther ecessiveallele.Obv i
ouslyt he
popul at
ioncont ai
nsmanycar r
iers,peoplewi thnor malpi gment ati
onwhoar e
het erozygous(A/ a)foralbi
nism.Whati st heirfr
equency ?Aconv enientwayt o
cal culat
et hef requenciesofgenot ypesi napopul at
ioni swi t hthebi nomi al
expansi onof(p q). Wel etprepresenttheal lel
icfrequencyofAandqt healleli
c
frequencyofa.

 Assumingthatmatingi
srandom,thedi
str
ibut
ionofgenot
ypi
cfr
equenci
esi
sp2
A/A,2pq A/a,andq2 a/a.Onl ythef
requencyofgenot
ypea/
aisknownwit
h
cer
tai
nty,
1/20000:

23/
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10
@07

 Oneper soni nev ery70i sacar ri

er!Althought her ecessivet r
aitisr are,the
recessi v
eal lel
ei ssur prisinglycommoni nt hepopul ation.Therei samessage
heref orany onepr oposingt oel i
mi natea“ bad”recessiv eallel
efrom apopul ati
on
by cont r oll
ing r eproduct i
on.I ti s practicall
yi mpossi ble.Because onl yt he
homozy gousr ecessivei ndividualsr evealthephenot y peagai nstwhi char ti
fi
cial
selectioncoul dact( byst eril
ization,forexampl e),theal l
elewouldper sistthrough
heterozy gouscar r
iers.Forar ecessiveal l
elepr esentin2ofev ery100per sons
(buthomozy gousi nonly1i n10, 000per sons) ,i
twoul dr equir
e50gener ati
onsof
compl etesel ectionagai nstt hehomozy gotesjustt oreducei tsfrequencyt oone
i
n100per sons.

Condi
ti
onsf
orHar
dy-
Wei
nber
gEqui
l
ibr
ium
Accordi
ngtoHardy-
Weinber
gpr inci
ple,thegeneandgenot ypefrequenci
esofeach
al
lel
einapopul
ati
onremainatanequi l
ibr
ium (
stat
ic)gener
ati
onaf
tergenerat
ion,
ift
hat
popul
ati
onexhi
bit
sfol
lowi
ngattr
ibutes:

23/
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@07

1.Ext
remel
yLargePopul ati
onSize:Theequi
l
ibr
ium i
ngeneandgenot ypefrequencies
occur
sonlyinlargesi
zedpopul at
ions.I
nsmallpopulati
ons,ther
ewillbesignif
icant
sampli
ngerr
orsandrandom fluct
uati
onsi
nthegenefrequencybychance,t
hesocal led
genet
icdr
if
torrandom dr
ift
.

2.Random Mat ing:Thepopul at

ionshal
lbepanmi ct
icwhereev
erygametehasanequal
opportunit
yoff usi
ngwi thanyot hergameteoftheoppositesex.Thi
snonpreferenti
al
fusi
onofgamet esiscalledrandom mating.Thenatur
alpopul
ati
onsmaynotf ulf
il
lthis
equali
ty,becauseindi
vidualsshowpr ef
erenti
almati
ngwithi
nasubsetofthepopulati
on
andrandom mi xingofgamet esfail
stooccur.

3.Bi
parentalModeofRepr oduct
ion:Hardy-
Weinbergprinci
pleisappl
i
cabl
eonlyfor
bi
parentsexual
l
yreproducingspecies.Uni
sexualorasexuall
yrepr
oduci
ngpopul
ati
ons
donotfol
lowHardy-
Wei nberglaw.

4.Homogeneous Age Str

ucture:A populat
ion nor
mall
yconsist
s ofindi
vidual
s at
di
ffer
entst
ages ofreproducti
ve matur
it
y.Accor di
ng t
o Har
dy and Weinberg,the
popul
ati
onmustbehomogeneousi nt
hisregard.

Summar
yofAssumpt
ionsofHar
dy-
Wei
nber
gEqui
l
ibr
ium

 Parent
srepresentarandom sampl
eofgenesandgenef
requenci
esf
rom t
he
genepoolofapopulat
ion.

 Heter
ozy got
es for any gene pai
r pr oduce t
wo kinds of gamet
es i
n
equal
frequencyshowi
ngcomplet
esegregati
onofal
lel
esofagenepai
r.

 Bot
hthepar
ent
sar
eequal
l
yfer
ti
le.

 Allthegametesareequal
l
yfer
ti
leand hav
eequalchancesofsur
viv
aland
for
mi ngazy
gote.

 Allthezygot
esf
ormed hav
eequalchancesofsur
viv
aland dev
elopi
ng i
nto
anadul
t.

 Genef
requenci
esar
ethesamei
nbot
hmal
eandf
emal
epar
ent
s.

 Mat
ingbet
weent
het
wosex
esi
snonpr
efer
ent
ial
andr
andom.

 Allgenot
ypeshaveequalr
eproduct
iveabi
li
tyandequalchancesofcont
ri
but
ing
thegenes/al
l
elest
othegenepoolofthepopul
ati
on.

23/
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10
@07

5.AbsenceofEvolut
ionaryForces:Thegenef r
equencywillr
emainstat
iconlyinthe
absenceofev
olut
ionaryforcesl
ikemutat
ions,sel
ect
ion,genet
icdr
if
tandmigrati
on,as
fol
lows:

 Ther
eshal
lbenogenemut
ati
ons,
becausemut
ati
onsaf
fectal
l
elef
requenci
es.

 Ther
e shal
lbe no gene f
low eit
herbyi mmigr
ation oremi
grat
ion,because
migr
antsal
ternumberofall
elesbydelet
ingf
rom thegenepooloraddi
ngt oit
.
Thi
sshowsabsenceofgeneflowinoroutoft
hepopulati
on.

 Allgenotypesi
napopulat
ionshal
lreproduceequallysuccessfull
y .Ther
eshallbe
nosel ect
ionandnopr ef
erenceforspecifi
cgenot ypeorphenot ypeformati
ng.
Theabsenceofselecti
onmeansev erygamet eisv i
able,allgamet eshaveequal
chancesofbecomingazy gote,everygamet i
cunion( zygote)alsosurviv
esand
gamet esorzygot
esarenotsegr egatedintoclassesofv aryi
ngsur vi
valvalue.
Undersuch conditi
ons,the geneti
c composi t
ion ofpopul at
ion willremain
unchanged.

 Genef
requenci
esar
esamei
nbot
hmal
eandf
emal
epar
ent
s.

From the above discussi

on itbecomes clearthatpopulat
ions exhibi
ti
ng genet
ic
equil
i
bri
um orfol
lowingHar dy
-Weinber
g'spri
nci
plear
estat
icwithzer
oev ol
uti
onar
yrat
e.
Thismeanssuchpopul ati
onsdonotev ol
veandanypopulati
onevolvingisnotstat
ic
anddoesnotexhibitgeneti
cequil
i
brium.

Har
dy-
Wei
nber
gequat
ion

 TheHardy-Wei
nbergequati
onisamat hemat
icalequati
ont hatcanbeusedt o
cal
cul
atethegeneti
cvar
iat
ionofapopul
ati
onatequil
ibr
ium.Toexploret
heHardy
-Wei
nbergequati
on,wecanexamineasi mpl
egeneticlocusatwhi chther
eare
twoal
lel
es,Aanda.

23/
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10
@07

 TheHar
dy-
Wei
nber
gequat
ioni
sexpr
essedas:

p2+2pq+q2=1

wher epi sthef r

equencyoft he" A"all
eleandqi sthefrequencyoft he"a"all
eleint he
populat i
on.Int heequat i
on,p2r epresentsthef r
equencyoft hehomozy gousgenot ype
AA,q2r epresentst hef r
equencyoft hehomozy gousgenot y
peaa,and2pqr epresents
thef requencyoft hehet erozygousgenot ypeAa.I nadditi
on,t hesum oft heal l
ele
frequenciesf oralltheallelesatt helocusmustbe1,sop+q=1.I fthepandqal l
ele
frequenciesar eknown,t hent hefrequenciesoft hethr
eegenot ypesmaybecal culated
usingt heHar dy-Wei nber
gequat i
on.Inpopul ati
ongeneti
csst udi
es, t
heHar dy-
Wei nberg
equationcanbeusedt omeasur ewhet hert heobservedgenot ypef r
equenciesi na
populat i
ondifferfrom thef requenciespredictedbytheequation.

Si
gni
fi
canceofHar
dy-
Wei
nber
gPr
inci
ple
Populationsi
nnat ur
erarelymeett hestri
ngentconditi
onsnecessar ytomai ntainthem
atHar dy -
Weinber
g equil
ibri
um.I tmeans Har dy-
Weinber
g Pr inci
plei s essenti
alt o
determine whetherthe popu lati
on is evol
ving ornot?I n orderto ascer tai
nt hat
evolut
ionaryagentsareoper ati
ngornot ,theal l
eleandgenot ypef r
equenciesoft he
populationaredetermi
nedgener ati
onaf t
ergenerat
ionandthencompar ed.Thepat tern
ofdev iati
on fr
om Har dy-
Weinber gr at
io wil
lt el
ltheagent/ agentsr esponsibl
ef or
evolut
ionarychange.Thus,Hardy-
Wei nbergpri
nciple:

 pr
ovi
desat
heor
eti
cal
basel
i
nef
ormeasur
ingev
olut
ionar
ychange

 t
endst
oconserv
egai
nswhi
chhav
ebeenmadei
nthepastandal
sot
oav
oidt
oo
r
api
dchanges

 mai
ntai
nshet
erozy
gosi
tyi
nthepopul
ati
on

 Pr
event
sev
olut
ionar
ypr
ogr
ess

23/
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10
@07

Dev
iat
ionsf
rom Har
dy–Wei
nber
gequi
li
bri
um
Genet
icequi
l
ibr
ium i
sdi
stur
bedi
nnat
ural
popul
ati
onsby

(
1)r
andom genet
icdr
if
t

(
2)nonr
andom mat
ing

(
3)r
ecur
ri
ngmut
ati
on

(
4)mi
grat
ion

(
5)nat
ural
sel
ect
ion

Anyi nter
acti
onsamongt hesefactors.Recurr
ingmut ationistheul t
imat
esourceof
vari
abil
ityinal
lpopul
ati
ons,butitusuall
yrequiresint
eracti
onwi t
honeormor eofthe
otherfactor
stoupsetgenet
icequil
i
bri
um.Weconsi dertheseotherfact
orsi
ndi
vi
dual
ly

Genet
icDr
if
t
Ify ouf l
ipacoin1, 000times,ar esul
tof700headsand300t ail
smi ghtmakey ou
suspiciousaboutthatcoin.Buti fyoufli
pacoi nonly10times,anout comeof7heads
and3t ai
lswouldnotbesur pri
sing.Thesmal l
erthenumberofcoi nfl
ips,themor eli
kel
y
i
ti sthatchancealonewillcauseadev i
ati
onfrom thepredi
ctedresul
t.(Inthiscase,t
he
predicti
oni sanequalnumberofheadsandt ail
s.)Chanceevent
scanal socauseallel
e
frequencies tofl
uctuate unpredictabl
yf r
om
onegener at
iont
ot henext,especiall
yinsmall
populations—aprocesscalledgenet i
cdrif
t.

 Some speci es, such as cheet ahs

(Fi
gure6.29),cont ainv eryli
tt
legeneti
c
vari
ati
on, pr obably because t hei
r
ancestr
al lineages passed t hrough
peri
ods when t he t otalnumber of
i
ndivi
dualsi nt hepopul at
ionwasv er
y
small. A smal l popul ati
on clearl
y
cannot cont ain l arge amount s of
geneti
cvariati
on.

 Eachi
ndivi
dualorgani
sm hasatmost
twodi
ffer
entall
eli
cformsofeachgene,andasi
ngl
ebreedi
ngpai
rcont
ainsat
mostfourdif
fer
entall
eli
cfor
msofeachgene.Supposethatwehavesucha

23/
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@07

breedingpair.Weknow from Mendel

iangeneti
csthatchancedecideswhi chof
thedi ff
erental
lel
i
cformsofageneget spassedt ooffspri
ng.Itistheref
ore
possiblebychancealone.thatoneortwooftheparent
alallel
esinthi
sex ample
willnotbepassedtoanyof f
spri
ng.

 I
ti s highly unl
ikelyt hatthe diff
erentall
eles pr
esentin a smallancest
ral
populati
onar eal
l passedt odescendantswit
houtanychangeofall
eli
cfr
equency.
Thi
schancef l
uctuationi nall
eli
cf r
equencyf r
om onegenerati
ont othenext,
i
ncludinglossofal l
elesfrom t
hepopul at
ion,
iscall
edgenet
icdr
if
t.

 Geneti
cdri
ftoccurst osomedegr eei nallpopul
ati
onsofifi
nit
esize.Per
fect
const
ancyofallel
i
cf r
equenci
es,aspr edi
ctedbyHar dy
-Wei
nbergequil
ibr
ium,
occur
sonlyininfi
nitel
ylargepopulat
ions,andsuchpopulat
ionsoccuronlyin
mathemat
ical
model s.

 Allpopulat
ionsofani malsaref ini
teandt hereforeexper iencesomeef fectof
geneti
cdr i
ft,whichbecomesgr eater,onav erage,aspopul ati
onsizedeclines.
Geneti
cdr i
fterodesgeneticv
ari
abi l
i
tyofapopul at
ion.Ifpopulati
onsizeremai ns
smallformanygener at
ionsi
nar ow,geneticv ari
ati
oncanbegr eatl
ydepleted.
Thislossishar mfult oaspeci es’
sev ol
utionarysuccessbecausei trestr
icts
potent
ialgenet i
cresponsest o envir
onment alchange.I ndeed,biol
ogistsar e
concernedthatcheetahpopulat
ionsmayhav einsuffi
cientv ari
ati
onforconti
nued
survi
val.

 Cert
ainci
rcumst
ancescanresulti
ngeneticdri
fthav
ingasignifi
canti
mpactona
popul
ati
on.Twoexamplesarethefounderef
fectandthebott
leneckef
fect

TheFounderEf
fect

"
Abottl
eneckassoci
atedwi
ththefoundi
ngofanewgeographi
cpopulat
ioni
scal
l
eda
f
ounderef
fectandmaybeassoci
atedwit
hfor
mati
onofanewspecies.
"

23/
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10
@07

 Whenaf ew i
ndivi
dual sbecomei solatedf rom al ar
gerpopul ati
on,thi
ssmal l
er
groupmayest abl
ishanewpopul at
ionwhosegenepooldi ff
ersfrom thesour ce
populati
on;t
hisiscal ledt hefounderef fect
.Thef ounderef f
ectmi ghtoccur,for
example,whenaf ew member sofapopul ati
onar eblownbyast ormt oanew
i
sland.Geneti
cdrift,inwhi chchanceev entsalt
eral l
elefr
equencies,willoccurin
suchacasei fthest ormi ndi
scri
minat el
yt r
ansportssomei ndiv
iduals(andt hei
r
all
eles)
,butnotother s,fr
om thesour cepopul at
ion.Thef oundereffectprobably
accountsfortherel ativel
yhighf r
equencyofcer taininheri
teddisordersamong
i
solatedhumanpopul ations.

 Forexampl e,i n1814,15Br i

ti
shcol onistsf oundedaset t
lementonTr i
standa
Cunha, agr oupofsmal lislandsi nt
heAt lanti
cOceanmi dwaybet weenAfri
caand
SouthAmer ica.Appar ent l
y,oneoft hecol onist
scar r
iedar ecessiveall
elef or
reti
nit
ispi gment osa,apr ogressiveform ofbl indnessthataffl
ictshomoz ygous
i
ndi v
iduals.Oft hef oundi ngcol oni
sts’240descendant sont hei sl
andinthelat e
1960s,4hadr etini
tispi gment osa.Thef requencyoft heallel
et hatcausest his
diseaseist ent imeshi gheronTr istandaCunhat haninthepopul ati
onsf r
om
whicht hef ounder scame.

TheBot
tl
eneckEf
fect

"Alar
gereducti
oni
nthesizeofapopulati
ont
hati
ncr
easesev
olut
ionar
ychangeby
genet
icdr
if
tiscommonl
ycall
edabot
tl
eneck.
"

 A sudden change i nt he
env ir
onment ,such as a
fi
re or f lood, may
drast i
call
yr educet hesi ze
ofapopul ation.A sev ere
drop i n popul ati
on si ze
cancauset hebot tl
eneck
effect,sonamedbecause
thepopul at i
onhaspassedt hrougha“ bottl
eneck”thatreducesi tssi
ze( Fi
gure
23. 10).Bychanceal one,cer tainall
elesmaybeov er
representedamongt he
sur vi
vors,other smaybeunder repr
esented,andsomemaybeabsent al
together
.
Ongoi nggenet icdri
ftisl i
kelytohav esubstanti
aleffect
sont hegenepoolunt i
l
thepopul ationbecomesl argeenought hatchanceev entshav elessimpact.But
ev enifapopul ati
ont hathaspassedt hroughabot t
leneckul t
imatelyr
ecoversin
size,itmayhav elow lev elsofgenet i
cv ar
iati
onforal ongper iodoftime—a
l
egacyoft hegenet i
cdr iftthatoccurr
edwhent hepopulationwassmal l.Human

23/
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10
@07

act
ionssomet
imescr
eat
esev
erebot
tl
enecksf
orot
herspeci
es.

I
mpactofGenet
icDr
if
tont
heGr
eat
erPr
air
ieChi
cken

 Mill
i
onsofgr eaterpr
airi
echi ckens(Tympanuchuscupi
do)onceli
vedont he
prai
ri
esofIl
l
inois.Asthesepr ai
rieswer
econvertedt
ofarml
andandotheruses
duri
ng t
he19t h and 20th centuri
es,t
henumberofgr eat
erprai
ri
echickens
pl
ummet ed(
Figure23.11a).

 By1993fewerthan50birdsremained.Thesefewsurvi
vi
ngbir
dshadlowlevel
s
ofgenet
icvari
ati
on,andlessthan50% oft heireggshat
ched,compar
edwi t
h
muchhigherhatchi
ngrat
esoft helargerpopulat
ionsi
nKansasandNebraska
(Fi
gur
e23.11b)
.

23/
10/
10
@07

Ef
fect
sofGenet
icDr
if
t:ASummar
y

Theexampl
eswe’
vedescr
ibedhi
ghl
i
ghtf
ourkeypoi
nts:

1.Genet
icdr
if
tissi
gni
fi
canti
nsmal
lpopul
ati
ons.

2.Genet
icdr
if
tcancauseal
l
elef
requenci
est
ochangeatr
andom.

3.Genet
icdr
if
tcanl
eadt
oal
ossofgenet
icv
ari
ati
onwi
thi
npopul
ati
ons.

4.Genet
icdr
if
tcancausehar
mful
all
elest
obecomef
ixed.

Nonr
andom Mat
ing

 Ifmatingisnonr andom,genotypi
cfrequencieswi l
ldevi
atef r
om Har dy-
Wei nberg
expectati
ons.Forexampl e,i
ftwodifferentall
elesofagenear eequall
yf requent
(p=q=. 5),weexpecthal fofthegenoty pestobehet erozygous( 2pq=2[ .5][.
5]
=.5)andone- quartertobehomozy gousf oreachoft her especti
veall
eles(p2=
q2= [.5]*
.5 =.25).Ifwe hav e posit
ive assortat
ive mat i
ng,indiv
iduals mat e
prefer
ential
l
ywi t
hot her
soft hesamegenot ype,suchasal binosmat ingwi th
otheralbi
nos.

 Mati
ngsamongindi
vi
dual
shomozygousf
ort
hesameal
l
elegener
ateof
fspr
ing
t
hatarehomozy
gousli
ket
hemsel
ves.

 Mat i
ngsamongi ndiv
idualsheterozygousf orthesamepai rofall
elespr oduceon
av erage50% het erozygousof fspri
ngand50% homozy gousof fspr i
ng( 25% of
eachal ternat i
vetype)eachgener ation.Positi
veassor tat
ivemat ingi ncreasest he
frequency of homozy gous genot ypes and decr eases t he f requency of
het erozy gousgenot ypesi napopul ati
onbutdoesnotchangeal l
eli
cf requenci es.
Pr eferent i
almat i
ngamongcl oser elati
vesalsoi ncreaseshomozy gosi tyandi s
cal l
edi nbr eeding.Wher easposi t
iveassor t
ati
v emat i
ngusual l
yaf fectsoneora
fewt rai
ts,i nbreedi
ngsi mult
aneousl yaffectsallv
ariablet r
aits.Str
ongi nbreeding
great l
yi ncr easeschancest hatr arer ecessi
vealleleswi llbecomehomoz ygous
andbeexpr essed.

 Becausei
nbreedi
ngandgenet
icdr
if
tarebothpr
omotedbysmal
lpopul
ati
onsi
ze,
theyar
eoft
enconfusedwit
heachother
.Thei
reff
ect
sarever
ydi
ff
erent
,howev
er.

23/
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10
@07

Inbreedingalonecannotchangeal lel
icfrequenci
esi nthepopul ati
on,onlythe
way st hatallel
esar ecombinedi ntogenot ypes.Geneticdr i
ftchangesal l
eli
c
frequenciesandconsequent lyalsochangesgenot ypicfrequencies.Evenv ery
l
ar gepopulationshavethepotent
ialforbeinghighl
yinbredifthereisabehav i
oral
preferenceformat i
ngwi t
hcloserelati
ves,alt
houghthissit
uationrarel
yoccursin
animal s.Geneti
cdrif
t,howev
er,willberelati
vel
yweaki nverylargepopulat
ions.

Mi
grat
ion

 Migrati
onpr event
sdi f
f erentpopulationsofaspeci esf r
om diver
ging.Ifalarge
speciesisdivi
dedintomanysmal lpopul at
ions,geneti
cdrif
tandselecti
onacting
separatelyint he di
fferentpopul ations can produce evol
uti
onarydiver
gence
amongt hem.Asmal lamountofmi grati
onineachgener ati
onkeepsthediff
erent
populati
onsf r
om becomi ngt oodistinctgeneti
call
y.

 Forexample,theFrenchandRussi
anpopulat
ionswhoseABOal l
elefrequencies
werediscussed previ
ousl
yshow somegenet icdi
vergence,butt heirgenet i
c
connecti
ont hr
oughi nt
erv
eni
ngpopulati
onsbycontinuingmi gr
ationpr events
them f
rom becomingcomplet
elydi
sti
nct
.

Nat
uralSel
ect
ion
Somei ndi
vi
dualsleavebehindmoreprogenythanother s,andtherat
eatcharacteri
sti
cs.
Thisiscalled scl
ection.Sel
ecti
oncanbear ti
fi
cialselecti
onornat ur
alselecti
on.I n
art
if
ici
alselect
ion,thebreeder
sselectforthedesiredchar acter
s.I
nnat ur
alselecti
on,
theenvir
onmentplay sthi
srol
e,t
husaffect
ingthepropor ti
onsofgeneinapopul at
ion.

 Naturalsel ecti
oncanchangebot hall
elicfrequenci
esandgenot ypi
cfrequenci es
i
napopul at i
on.Althought heef f
ectsofsel ecti
onareof t
enreportedforpar ti
cular
poly
mor phicgenes,wemustst resst hatnat ur
alselecti
onact sont hewhol e
ani
mal ,not on i solated tr
ai t
s.An or gani
sm t hat possesses a super ior
combi nationoft r
aitswi l
lbef avored.Anani malmayhav et r
aitsthatconf erno
advantageorev enadi sadvantage,buti tissuccessfuloveral
lifitscombi nation
oftrai
tsi sf avorable.Whenwecl aimthatagenot y
peatapar ti
culargenehasa
hi
gherr elat i
vefit
nesst hanothers,west atethatonav er
agethatgenot ypeconf ers
an adv ant age in survivaland r eproduction inthe populati
on.I fal t
ernative
genotypes hav e unequalpr obabili
ti
es ofsur vi
valand r eproducti
on,Har dy-
Weinber gequi l
ibri
um isupset.

 Usi
ngt
hegenet
ict
heor
yofnat
uralsel
ect
ion,onecanmeasur
erel
ati
vef
it
ness

23/
10/
10
@07

val
uesassoci
atedwithdiff
erentgenotypesi
napopulati
on.Geneti
cist
softenuse
Wt odenot
etheexpect edaveragefi
tnessofagenot
ypei napopulati
on,wit
hthe
genoty
peofhighestfi
tnessgivenavalueofoneandfi
tnessesofothergenot
ypes
i
ndicat
edasfracti
ons.

 Wei llustratemeasur ementoff i

tnessusi nggenet i
cv ariati
onassoci at edwitht he
diseasesi ckl
e- cel lanemi ainhumanpopul ations.Consi deringonl yt heallel
esf or
normalhemogl obi n(A) andsi ckle-cellhemogl obin(S)f ort hebet a-hemogl obin
gene i n human popul ati
ons,t he possi ble genot ypes ar e AA,AS ,and SS.
Measur ement sofv iabili
tyofindiv idualsoft heset hreegenot y pesi nnonmal arial
env ironment sgi veaf i
tnessv alueof1t ogenot ypesAAandASandaf i
tnessof
0.2t ogenot y peSS.Peopl ehav ingt heSSgenot ype,whoar esuscept i
blet o
sev ereanemi a, ar eexpect edtocont ri
buteonl y20%asmanyof fspringt othenext
gener ationonav erageasar ei ndi vi
dualshav ingt heAA orASgenot ypes.I n
mal ar ialenv i
ronment s,genot ypeAShast hehi ghestf itness( 1) ; genot ypeAAhas
a sl ightlydecr eased f it
ness ( 0.9)because t hese i ndividual s hav e a greater
i
nci denceofmal ariat hanASi ndividuals,andSShasal owf itness( 0.2)because
of anemi a.Fr om t hese measur ed f i
tness v al
ues and knowl edge of t he
frequenci esofal lelesi napopul ationandi tssy stem ofmat i
ng, onecancal culate
theav erageef f ectt hatanal l
elehasont hephenot ypeofr elat ivefitnessint hat
popul ation.Int heexampl eofsi ckle-
cellanemi a,theav erageef fectofal l
ele
Sof tnessi namal arialenv i
ronmenti sabal ancebet weent hest rongl ynegative
effecti thas when homozy gous and t he posi ti
ve ef fectt hati thas when
heter ozy gouswi thal l
eleA.

 Theav er ageef f
ectofanal leleonf it
nessi sexpr essednotonl ybyi t
sdirect
contri
but i
ont othef i
tnessofi t
spossessor sbutbyai dthatitspossessor sgiv
et o
cl
oser elatives,whoar el i
kelyal sot ocont aincopi esoft heallel
e.Thet erm
“i
nclusivef itness”per tainstocaseswher etheav erageef fectofanal l
elewoul d
becal culatedi ncorrect l
yifonl yi tsdirecteffect sonf i
tnesswer emeasur ed.
Somet raitsandcombi nat i
onsoft rait
sar eadvant ageousf orcert
ainaspectsof
anor gani sm’ ssurv i
valorr epr oductionanddi sadv antageousf orothers.Darwin
used t he t erm sexualsel ection to denotet he selection oft r
ait
st hatar e
advantageous f orobt aini
ng mat es butnotf orsur viv
al .Br i
ghtcol or
s and
el
abor atef eat herscanenhanceamal ebird’scompet it
iveabi l
ityinobt ai
ning
mateswhi l
esi multaneousl yincreasinghi svi
sibi
litytopredat ors.

 Envi
ronment
alchanges,suchasexti
ncti
onofapredat
orpopul
ati
on,canalt
erthe
sel
ecti
veval
ueofdiff
erenttr
ait
s.Theacti
onofsel
ecti
ononcharact
ervar
iati
onis

23/
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10
@07

t
her
efor
ever
ycompl
ex.

Mut
ati
on
Mutati
onsar echangesint
hestruct
ureofgenesandchromosomes. TheHardy
–Wei
nber
g
t
heorem assumesthatnomut
ati
onsoccurort
hatmut
ati
onal
equi
li
bri
um exist
s.

 Mutat i
ons,however ,
areaf actoflif
e.Mosti mport
ant,mutat
ionsaretheori
ginofallnew
all
elesandasour ceofv ari
ationthatmaybeadapt i
veforananimal.Mutati
oncount ers
thelossofgenet i
cmat er
ialfrom naturalsel
ect
ionandgeneticdr
if
t,anditi
ncreasest he
l
ikel
ihoodt hatvariati
onswi llbepr esentthatall
ow someindivi
dualstosurvi
vef uture
environmentalshocks.

 Mutat
ionsmakeexti
nctionlessli
kel
y.
Mut at
ionsarerandom events,
andt
heli
keli
hoodof
amutati
onisnotaff
ectedbyt hemutati
on’susef
ulness.Organi
smscannotf
il
terharmf
ul
genet
icchangesf
rom advantageouschangesbeforetheyoccur.

 Theef fectsofmut ati

onsv ar
yenormously.Neutralmut ati
onsar eneitherharmfulnor
helpful totheorgani
sm.Neut r
almutati
onsmayoccuri nr egi
onsofDNAt hatdonotcode
forpr ot ei
ns.Ot herneut r
almut at
ionsmaychange a pr otein’
s str
ucture,butsome
proteinst ol
erat
emi norchangesi nstr
uctur
ewithoutaffecti
ngt hefuncti
onoft hepr
otein.
Mut ations thatdo af fectproteinfuncti
on are morel i
kel
yt o be detr
imentalthan
benef i
cial.Thisi str
uebecausear andom changei nanest abli
shedpr otei
nupset s
milli
onsofy ear
sofnat uralsel
ecti
onthatoccurr
edduringt hepr ot
ein’
sevoluti
on.

 Mutati
onsinDNAt hatisincorporatedint
oagamet ehav ethe potentialtoaff
ectthe
f
uncti
onofev er
ycellinanindividuali
nthenextgener
ation.Thesemut ationsarel
ikel
yto
i
nfl
uencetheev olut
ionofagr oupofor ganismsMutationalequi
li
brium existswhena
mutati
onfrom thewild-
typealleletoamut antfor
misbal ancedbyamut at
ionfr
om the
mutantbackt othewi l
dt ype.Thishast hesameef fectonal l
eli
cf requencyasi fno
mutati
onhadoccur r
ed.

 Mutat
ionalequi
li
bri
um rar
elyexi
sts,howev
er.Mut
ationpressur
eisameasur
eoft
he
t
endencyforgenefr
equenci
estochangethr
oughmutati
on.

23/
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10
@07

Ref
erences
Books

 I
ntegr
atedpr
inci
plesofZool
ogybyHi
ckman-
14E

 Campbel
lZool
ogy
-10E

 Zool
ogybySt
ephenMi
l
lerandJohnHar
ley
-10E

 Or
gani
cev
olut
ion byVi
rBal
aRast
ogi
-13E

 htt
ps:/
/bio.l
ibr
etexts.
org/
Bookshelves/I
ntr
oduct
ory_
and_General
_Bi
ology/
Bo
ok%3A_Introductor
y _
Biol
ogy
_(CK-12)/05%3A_Ev
olut
ion/
5.02%3A_Hardy-
Weinberg_Theorem

 ht
tps:
//en.
m.wi
ki
pedi
a.or
g/wi
ki
/Har
dy%E2%80%93Wei
nber
g_pr
inci
ple

 ht
tps:
//www.
nat
ure.
com/
sci
tabl
e/def
ini
ti
on/
har
dy-
wei
nber
g-equat
ion-
299/

 ht
tps:
//www.
sci
encedi
rect
.com/
topi
cs/
neur
osci
ence/
har
dy-
wei
nber
g-
pr
inci
ple

23/
10/
10