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daglas
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evolucia
daglas
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Evolution. Douglas J. Futuyama
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ISBN 978-9941-9117-2-9
3/5, 0162,
Ilia State University Press
3/5 K. Cholokashvili Ave, Tbilisi, 0162, Georgia
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guidetothecreation/evolutioncontroversy(JomnesandBartlett,Subdury,MA,2001)M.Pigliuccio:Denying
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. J. Felsenstein: Inferring phylogenies (SinauerAssociates,
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1990)
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pylchella)
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,photoResourceHawaii/AlamyImages,
AnthonyiMercieca/PhotoResearchersInc.
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56
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1996)..,
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57
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,
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()(Ambystoma
tigrinum),
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,photoPainet,Inc.,photobyHenkWallays.)
58
()
3
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.(),()Hemirhamphidae()
.
(fromJordanandEvermann,1973).
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(FromMillais1897)
60
.3.18.
Thorius
Pseudoeurycea
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61
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(
Eusthenopteron)
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Milleretta).
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(AfterRomer1966).
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).,
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62
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. , ,
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. ,
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olivacea
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inornata
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heliobates
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pallida
Camarhynchus
parvulus
Camarhynchus
pauper
Camarhynchus
psittacula
Geospiza
scandens
Geospiza
conirostris
Geospiza
magnirostris
.3.22.
(AfterSchluter2000
andBurnsetal.2000)
Geospiza fortis
Geospiza
fuliginoza
Geospiza
dicillis
()
63
Argyroxiphium sandwicense
()
()
Wilkesia hobdyi
Dubautia menziesii
,
. , 800
. ,
(CarsonandKaneshiro1976).
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140 , 500 , .
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tropheops
Petrotilapia
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polydon
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sandwicense,
.
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(,photobyElizabethN.Orians,photo
byGeraldD.Carr,photobyNobleProcter/
PhotoResearchersInc.)
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.
(AfterFryerandIles1972).
64
1. ()
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2. .
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,
() () ,
.
.
9. .
.
10.
,
.
()
()
.Bernhard Rensch: Evolution above the species level (Columbia University
Press,NewYork,1959)..
: D.R. Brooks and D.A.McLennan in The nature of diversity: An evolutionary voyage of
discovery(UniversityofChicagoPress,1959),D.J.Futuyma,Thefruitofthetreeoflife:Insights
intoevolutionandecology(pp.2539inAssemblingthetreeoflife,J.CracraftandM.J.Donoghue,eds.Oxford
UniversityPress,NewYork,2004).
:
Stephen Jay Gould: Ontogeny adn phylogeny (Harvard University Press, Cambridge, MA, 1977).
. M.L.McKinney and K.J.McNamara: Heterochrony:The evolution of
ontogeny(Plenum,NewYork,1991)andbyR.A.Raff:Theshapeoflife:Genes,development,andtheevolution
ofanimalform(UniversityofChicagoPress,1996).:G.P.Wagner:The
biologicalhomologyconcept(AnnualReviewofEcologyandSystematics205169,1989).
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: 1) GCTGATGAGT 2) ATCAATGAGT 3)
GTTGCAACGT4)GTCAATGACA
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.
(1974)
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(,
. 3.10) ) (, . 3.21) )
(,Amphignathodon).
7. ,
.(InternationalHumanGenomeSequencing
Consortium, 2001 Venter et al. 2001)
5.
8. ,
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) ,
65
66
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NaturalHistoryMuseum,London).
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()
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145
(J)
200
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251
299
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,,
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359
, lycopsid, sphenopsid
,
.
(D)
416
,,,,
(S)
444
,,.
(O)
488
()
542
(T)
Homo erectus
Homosapiens
2500
(19001700 )
(1000)
, Cnidaria (), Annelida (
),Arthropoda()
(Q)
3500
()
,
(K)
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.,
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(Jablonskietal.1986).,
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.
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4)
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2),.
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,,
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,
.
.(Levinton,2001).,
(.4.3Sheldon1987).
(1985) Gasterosteus doryssus samekala
,111000
.5000.
(.4.4).
.
(),
(.4.4).5000
,.
.
,,
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,
.
.
,
.,
71
72
Cnemidopyge
Nileids
Ogygiocarella
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Nobillasaphus
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0
0
.4.3.
.,
(After
Sheldon1987)
9 11
12
13
9 10 11 12 131415 16
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.,
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()
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),,(.
4.6).,
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4.6).
10.5
10.0
9.5
9.0
8.5
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.,
(),
(),
.5000110000
(AfterBelletal.1985photocourtesyofM.Bell)
8.0
()
3.0
2.5
2.0
Gasterosteus doryssus
1.5
1.0
0.5
1
10
20
30
40
50
60
70
80
90
100 110
()
73
.4.5. .()
Sphecomyrmafreyi,
.
.(),,
Sphecomyrma(courtesyofE.O.WilsonAfterWilsonetal.1967)
Sphecomyrma freyi
()
phecomyrma-
.
(.4.6).
.
..
.
.(Ichtiostega.4.6).,,
,.
,(4.64.6
).(
)
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4.6)..
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.,
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(Clack,2002).(,
).
(
)(
,).
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.,
Aves,.,
,
(ChiappeandDyke,2002Xuetal.2003).
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.Archaeopteryx(.4.7).,
,,
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(. 4.8), . ,
,
(Sinosauropteryx),,(
?)(Caudipteryx),(Microraptorgui),
(.4.7).ArchaeopteryxAves
..
74
.4.6. Eusthenopteron,
(Rhipidistia),
.()
.()
.()
.
.()
Icthyostega.
(),
,()
.
,
.
.
:jI
o()
ppmpo
,ppprf
qj
sq()st
t()
[(,)AfterAndrewsandWelstoll,1970()afterMoy
ThomasadnMiles1971()afterJarvik1955()after
CoatesandClack1990()afterJarvik1980)]
()
Eustenopteron ()
()
Ichtyostega-
()
()
() Ichthyostega ( )
()
()
,(),
(4.9),,
,,.Aves
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(),
.,
( ,
75
). ()
,
(.11).
.
(Kemp1982SidorandHopson1998).
,
,,
.
,.
(),
(
),. ()
(),
()
Archeopteryx
()
.4.7. .()Microraptorgui
(140
).,
.,
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.
.
.(fromX.Xuetal.2003photoTomStack/Painet,Inc.)
()
()
.4.8. ()Archaeopteryx,()()()(
)Deinonychus.Archaeopteryx
(1)(2)
,(3)
,(4),
(5)6)
.Archaeopteryx
(,afterColbert1980afterOstrom1976).
76
Therapoda
Allosauroidea
Tyrannosauroidea
Neotetanurae
Sinosauropteryx
3
Coelurosauria
Oviraptorosauria
(Caudipteryx-
)
4
Maniraptora
Microraptor gui
Paraves
Dromaeosauroidea
Archaeopteryx
6
Aves
Confuciusornis
Ornithurae
Ornithoraces
Euorithes
( )
Ornithothoraces
235
200
160
145
125
(. )
.4.9.
Archaeopteryx
.
(
).(1)
,,
(Theropoda)(2)
(Neotetanurae)(3)(Coelurosauriaa)
(4)Sinosauropteryx,
()(5)Maniraptora(6)
Paraves.
M.gui(7),
Archaeopteryx()(8)
Ornithurae
(9)Euornithes
.
(After
Sereno1998ChiappeandDyke2002Prum2003).
(,).
.,,
,.
,
(),
.
,,
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(.4.10).
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.
Morganucodon(.4.10).
/
.
,
.
Hadrocodium
Morganucodon
(Luoetal.
2001).Morganucodon
,
.4.10 ,
.()
Haptodus.:(f),
,/
(art/q).()Biarmosuchus.
.()
Procynosuchus.,
,(j)(sq).
.()
Thrinaxodon.,
,
.()
Probainognathus.
,
,(d)(art)
(sa).)Morganucodon.
(
),
(condyle)(dc)()
(sq)(AfterFutuyma1995basedonCarroll1998andvarious
sources).
77
() (Haptodus)
()
(Biarmosuchus
() (Procynosuchus)
f ( )
()
(Thrinaxodon)
()
(Probaompgnathus)
,
,
()
Morganucodon
Morganocodon-
.
.
78
..
(),
,45.
, (, ,
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), . . ,
.,
.
.
.
.
,.
(130
),.
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,(Gatesyetal.1999).
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,,,
,,
(),
.
().()()
,.
,...
,5035
(.4.11)(Gingerichetal.2001ThewissenandBajpai2001Thewissen
andWilliams2002).Pakicetus(5348)()
.
Ambulocetus (4847 ) ,
.,,
.
.Ambulocetus.
,.(,
Rodhocetus,4939).,
.
,
,.
Dorudon(35).
,,
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.,
.
.
,56
(.2)..
.67(
()
Elomeryx
Elomeryx
.
.
Ambulocetus
.
.
()
Ambulocetus
Rodhocetus .
.
()
Rodhocetus
Dorudon
.
()
Durodon
()
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,()).
,,,
(. 4.12). ,
.
.,
. ,
(e.g.,WoodandCollard1999).
,
.,
(),,
,
,.,,
,
(CiochonadnFleagle1993,Strait
etal.1997Begun2004).
79
.4.11. ,
(
),
.(),
Elomeryx,
.
()Ambulocetus.()
Rodhocetus
,
()Dorudon,
.
.(),
Phocoenaphocoena
.
,
,
(after
Gingerich2003anddeMuizon2001artby
NancyHaver)
80
()
( )
()
.
,(Brunetetal.2002).,Sahelanthropusthcadnsis,
67.Sahelanthropusthcadnsis
,
60
,,
..
40
Orrorin
tugenensis(6),Ardipithecuskadabba(5,25,8),Ardipithecusramidus(4,4
)Australopithecusanamensis(4,23,9).
20
,,,
.3,5Kenyanthropusplatyops.
() 0
3,5
Australopithecusafarensis(.).
2000
Homo sapiens
..
(platyops),,
1500
,(4003),
Homo erectus
().
1000
Homo habilis
,Anamensisafarensis
.,afarensis,
Australopithecus
.
500
afarensis,
( )
(.4.13).,
.4.12. ()
. (Parnthropus) (
().
),
,
..
2
,2,62,3
.
. ,
(AfterJones .
etal.1992).
Australopithecusafricanus,A.afarensis
(4503)(.4.14,).Africanus
,
.
Homo1,91,5(
80
Homo sapiens
( )
()
2
3
4
5
H. neandathalensis
H. heidelbergensis
H.antecessor
.4.13.
.
(AfterWood2002).
H. ergaster
P.aethiopicus
Au. rudolfensis
Au bahrelghazali
Au. anamensis
Kenyanthropus platypos
, ,
Au. africanus
Au. afarensis
Ardipithecus
ramidus
Ar. kadabba
Sahelanthropus
tvhadensis
, ,
P.robustus
Au. garhi
H.erectus
, ,
Paranthropus
boisei
Australopithecus
habilis
Orrorin tugenensis
). Homo habilis ,
H.habilis, H.ergaster H.rudolfensis,
.Homohabilis,(
)(.4.14).Australopithecus(Wood
and Collard 1999). Australopithecus africanus,
Homo erectus . Homo
habilis,
(6108003),,.U
,
,,
,
. Homo habilis (
)
,(Potts1988).
1,6200000
Homoerectus.,habilis,erectus
sapiens.
erectus,
.,.,
,
,10002
(.4.14).1
Erectus,
. erectus .
.A
,H.habilis.,
.
Homoerectus,Homosapiens
.400300.Sapiens,
11752,200
,14002.
,,
,
.homosapiens
,
Homoneanderthalensis.
, (. 4.14).
, .
,,M.
,(15003).
(
).
12030.
sapiens
.(170)
.,
.4.14. .
()Pantroglodytes.,,
.()Australopithecusafarensis
.()Australopithecusafricanus
.()Homohabilis.,
..()
Homo(sapiens)neanderthalensis.H.erectus,
[(,)afterJonesetal.1992)afterHowell1978)].
()
()
81
A.afarensis-
()
()
...
...
...
()
...
(
)...
()
...
82
4
,,
40000 .
6 , , sapiens sapiens (
),,
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40 .
.,
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11000.
.,,
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,,
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( ). , .
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,,,,.
.,
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(Lovejoy1981Fedigan1986).,
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.,
.
,.
,,
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,
.
,,
,,
.
83
.(Archaeognatha).
(.4.15).,
.,
,,
.
(Donoghueetal.1989).,
.
.,,
(),
.,(Equidae)
50
(.4.16).
.,
(.4.16)..
,Morganucodon,
.
, ,
.
,.,
,
.
,,
.
(1978),,,
.,
.,
.
.,
, .
.
.,
. ,
.,
,,
.,
(.4.17).,,
,
(Palmer1982).
,.
,,
.
.1972
,()
.
,.
.4.15.
(Archaoegnatha).
,,
(AfterCSIRO1991).
84
()
550
500
Equus
()
400
Merychippus
()
350
300
()
450
.4.16. Equidae.()40
.
,.()
,.
(),
()(AfterMacFadden1986).
.,
,
200
,
150
,
100
(.4.18).
50
(),
,(.4.18).
()
.
0.30
(Mimomysoccitanus)(
)(.4.19).
0.26
,,
0.22
,
0.18
,
,
(ChalineandLaurin1986).,(Ectoprocta),
0.14
Metrarabdotos(
)(.4.20,Cheetham1987).
0.10
.
,
,.
0.06
,
(.4.18).
0.02
,
60 55 50 45 40 35 30 25 20 15 10 5
.
( )
195416.
,,
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,.
,.
,,
,,
,,
.
(.913).,
,.,
,,
,.(.4.18).
(.4.21).
(GouldandEldredge1993)(Levintone2001).
250
HyracotheriumMesohippus
(
(
)
)
()
,,
85
Pyrgoma
milleporae
(1 )
Tetraclita
radiate
(4 )
Pyrogomatidae
Balanidae
Tetraclitidae
Chthamalinae
Bathylasmathidae
Balanus
balanoides
Archaeobalanidae
Chthamalus
intertextus
(6 )
Ca
to
ph
ra
gm
id
ae
Octomeris
brunnea
(8 )
da
ali
m
tha
Ch
Chthamoloidea
()
Balanoidea
()
.4.17.
.
,(..3.6).
(
).
(AfterPalmer1983drawingsbyG.Sowerby,from
Darwin1854).
Coronuloidea
( )
.4.18.
.()
.()
.
()
.()
,
.
,.(
Malmgrenetal.(1983)
).
()
,
,
.
()
86
4
.4.19. .Mimomys
.,
()
().1,5
.
.
.,
(AfterChalineandLaurin1986)
(. )
M. tenue
(. )
M. auriculatum
#10
#9
M. colligatum
10
.,
1 , 1
,5
50.,
.
...,
.2,718
().
,
,
.
(1993) (
..4.229B).
,
. ,
Hyracotheriumgrangeri650000
(.4.23).
0,0000057 (
1130002
).0,00030(6745)
(Gingerich1993).
.
.
.
,
,
(,
,,
).
(4.4
).,
.
(. 4.23). ,
.,
()HyracotheriumgrangeriH.aemulon
135000,00024
(.4.23).,H.grangeri
.4.20. .(Metrarabdotos)
.
.
(AfterCheetham1987photoscourtesybayAlanCheetham).
87
.4.21 ,
Globorotalia
.()
()M()
.
.
,
.(AfterMalmgrenetal.1983photoscourtesyofB.Malgren.)
...
(. )
(.
3
4.23).
...
.
,
/
4
,
.
, ,
,
5
,
().,100
,,
6
0,024
,
0,010
0,035(HendryandKonnison1993.
.13.3).(100)
10.4
,
-0.10
-0.05
0.00
+0.05
Hyracotherium
.
,
()
,,
,
, ,
.
,
.
+0.10
s
, 68% ( x )
1s-
.
x-3s
x-2s
x-1s
x+1s
x+2s
x+3s
()
( x ) 0,25s-
0,5s-,
, 0,25
0,5 .
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(s),
68%( x )s
,.95%2s
,99,7%3s
.()(
).
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0,5s.
,
0,250,5.
x-3s
x-2s
x-1s
x+1s
x+2s
x+3s
88
()
H. aemulor
H. gangeri
,
.
.
Loge
()
Log10 ()
.4.23. Hyracotherium,
().()
(
).
H.grangeri650000
(15201760),
H.aemulor.()
2.
,
.
(
).
,
(,
),
0,648log0,225(After
Gingerich1993).
H.grangeri- H.aemulor .
,
H,grangeri
.
Log10 ()
1. ,.
,.
2.
.,()
,,.
3. ,,,,
Homo,.
,,
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4. .
5. ,,
.
6. ,,
,.
7. ( )
.
,,
().
.
8. .,,
,,
.
(
,)
()
()
,
,Earthsystemhistory(W.H.Freeman,NewYork,2005)byS.M.Stanley.Secondeddition.
:R.L.Carrol:Vertebratepaleontologyandevolution(W.H.Freeman,NewYork,1988)
Paleobiology:Asybthesis,edittedbyD.E.G.BriggsandP.R.Crowther(BlackwellPublishing,Oxford,1990).
Humanevolution:AnillustratedintroductionbyR.Lewin(BlackwellPublishing,Oxford,2005)
.
Punctuatedequilibriumcomesofage(Nature366223227,1993)byS.J.GouldandN.Eldredge.
.
.Genetics,paleontologyandmacroevolution(Cambridge
UniversityPress,Cambridge,UK,2001)byJ.S.Levinton.Secondedition.
1.
.(.4.4).
,20000
,,5000
?
,7000030000?
89
90
4
2. ()
,(.8).
?
?
3. ,
(..4.18).
?
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,,
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5. ,
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6. ,
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? () 65
60(.4.4),()7025
()MorganucodonProcynosuchus(.4.10).
,,
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7. ,
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3500000000
,
.
.
,
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(
,
),
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5
,,()
.
,
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.
,500
.
(..
97).
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Pikaia.
Hallucigenia
Marrella(ArtJohnSibbik/
NHMPL).
92
5
5,,
.()
,.
,
:
1.
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)
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7.,
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9.
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,
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.
()
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.4.1)3,8.
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,,
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D,
.
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(),
.
,(Orgel1994
MaynardSmithandSzathmary1995Rasmussenetal.2004).
.,
(CH4),(NH3),(H2)(H2O)
(HCN)(H2CO)
,,,
(.5.1).
.(
,),
.,,
().,
.(),
,
.
,,,
.,,
.,
, ,
.
2
.
(1970),(,Qb
)()
H2, CH3 NH3-
3
.
(),
,
.
,
1
() .,,
H2O
.
,
,
.,
.
,
.,
.,,
,(450,150
).
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H 2O
N2
N H3
CH4
H2 CO2
`
94
1 R1
.
2 -
R1
R2 ,
.
AA
R1
R1
AA
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.,
()
,(AA)
.
(R1),
.
,
R2
(AfterMaynard
SmithandSzathmary,1995).
R2
AA
AA
.(1993),
,
,
(.5.2).
.,
.
,
,
.
,
.
2,52,5542
,.
(3,8 ) ,
.3,5
(
.5.3).(
).
,
.
. 2,2
,,
,
(Knoll,2003).
.
Eucarya()
Archaea Bacteria
(..2.1).2
.
Archaea
,
{(
(
) ,
PCR (
), )}.
.5.3. ,SharkBay().
.(
.
).
EUCARIA
Sulfolobus
ARCAEA
Thermoproteus
Thermoplasma
Halobacteria
BACTERIA
,EucaryaArchaea
,Bacteria(.5.4.).,
,
,,,
,()(Woese2000).
,
.
.
,
.,
.,
.,Giardias
,
, (
,).
,
(Margulis,1993MaynardSmithandSzathmary,
1995),(..5.4)..
(E.coli),
,,
69.,
.
,.
,
2,7
,1,5(.5.5).
,
95
.5.4.
.
.(After
Knoll2003andothersources.)
96
()
()
()
.5.5. .
()Eontophysalis1,5
.()
,Tappania(
)1,5
..()
(PhotographscourtesyofA.H.Knoll).
.,,
.
,Plantae,Fungi
Aminalia(.5.6).
,.
1,
.
.
.
:,
().
640.
.
,
(.5.7).
(
)
,
Cnidaria () Bilateria
().
Mycetozoa (
)
Lobosa ()
542
,.10
,1025
,
,
.
brachiopoda () (.
5.8), (. 5.8)
Plantae ( )
Chlorophyta ( )
Rhodophyta ( )
Phaeophyceae ( )
Oomycetes ( )
Ciliophora ()
Apicomplexa (
. )
Euglenozoa ( ..)
Parabasalia
Diplomonadidae (. Giardia)
.5.6.
.
.
Parabasalia
Diplomonadida(AfterBaldauf
etal.2000andMcGrathandKatz2004.)
()
()
.5.7. .()
Mawsonitesspriggi
()
,
.()Dickinsoniacostata
.(Thenatural
HistoryMuseum,LondonKenLucas/Visuals
Unlimited)
, , .
(.5.9
).
.
.Haikouichtys,,,
,(.5.10Shuetal.
2003).(.5.10).
(Bauplane)
,
.
(Erwin,1991LippsadmSignor,1992).
?(1996)
,
,1000.
.,
,.
.
(KnollandCarroll,1999Knoll,2003).
(.20)
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paleontologyandevolution(ChapmanandHall,London1993)P.KenrickandP.R.Crane,Theoriginandearlydiversificationoflandplants
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Science 268: 5258, 1995). : J. W. Valenine (ed.), Phanerozoic diversity
patterns:Profilesinmacroevolution(PrincetonUniversityPress,1985),D.Jablonskietal.(eds.),Evolutionary
paleobiology (University of Chicago Press, 1996).
: P.D.Taylor and G. P. Larwood (Eds.), Major evolutionary radiations (Clarendon
Press,Oxford,1990).:A.HallamandP.B.WignallsMass
exstintionsandtheiraftermath,OxfordUniversityPress,1996)M.J.BentonsWhenlifenearlydied:Thegreatest
massexstintionofalltime(Thames&Hudson,NewYork,2003)
:E.O.Wil
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2p1p2,2p1p32p2p3.
:
- :
,.
.
.:1)
.
,
2),,
.,,
.
,
.(
),.
,,,
.
:
1..,
,p2:2pq:q2
2. ( ,
)..,
H=1-(p12+p22+p32)
196
9
.,
,,
(),,100,,
50.,
.()
.
3..A1
A2.
.
.,
:
4..,
(8),,.
5..
,/
.,,
.
, , ,
,
.
.,
().,
,1:1.
.
,
.,
,!,
,
.,
,
.
,
,
(.9.5).,
.(,A1)
(A2),(A1A1A1A2)p2+2pq.
,
.,q=0,01,
(0,01)2=0,0001.,
.,
.
,.,
Cleorarepandata
,.
10%(Ford1971).
,.
?.
,,
,.
197
.
B.
,.
H=H0(1F),H0,
,F.
,
, ( ,
,).A1A1
A1 A1
A1A1.,A2A2A2A2
.A1A2.
.,
1/4A1A1,1/2A1A1,1/4A2A2(
).,
.,F
(.B).(
),F
.
,
.,Avenafatua
(.9.6).
.,
.
.9.5.
,A
,.
1/32/3
,
(AfterHartlandClarck1989).
(poly,morph)(
).,
0,01..
,
.
(q)
.
0.9 0.8 0.7 0.6 0.5 0.4 0.3 0.2
1.0
,
(. . 9.1) . , Drosophila
0.9
melanogaster ,
,
0.9
p ( A A )
q (A A )
.24
0.7
.
AsteraceaeLyiaglandulosa
0.6
(. 9.7).
2pq (A A )
.
0.5
, Asteraceae
0.4
(
).,
0.3
,
.
0.2
20,Drosophila
0.1
,
p=q=0,5.
0.1
0
0.1
0.2
0.3
0.3
0.5
0.6
(q)
0.7
0.8
0.9
1.0
198
9.
pA1.
,A1A1
Fp.FqA2A2.
(.2):
.
. A1
,A2.A1
A1*,
A1*0,52=0,25.
A1*,
A1*A1*.
,(A1),
(A1*)(.1).A1*A1*
,.
(
).
F
.
,F=0,
,F=1.
,F
,1F
.A1A2
pq,,
A1A1(1F)p2.
(1F)2pq,
A2A2(1F)q2.
F
,
,
.,A1
A* A
A A
A* A
A* A
.1.
.
,.A1A1*
()
.1A1*
(
).A1
,
(.)
A1A1
p2(1-F) +
pF
= p2+Fpq=D
A1A2
2pq(1-F)
A2A2
q2(1-F) +
= 2pq(1-F)=H
qF
= q2 +Fqp=R
,
,,
,.
H
(1F),
(2pq).
2pqH0:
H=H0(1-F) F=(H0-H)/H0
,
. H
2pq , p
q.
F
,
.
0.5
A*A*
(F=0)
(F=0.5)
() A1
(
).
0.48
0.48
0.4
0.36
0.3
0.28
0.24
0.2
0.16
0.1
A A
AA
A A
.2. ,
p=0,4
q=0,6,
(F=0),
(F=0,5).
199
.9.6. Avenafatua
,
.,
F(DatafromJainand
Marshall1967.)
0.7
0.6
H-W
0.5
F = 0.854
F = 0.862
0.4
, ,
,.
0.3
,
,
0.2
.
.
0.1
,
(. 9.8). ,
0.0
,
B B
,,,
.F3,
.
(
),,(
),.
,
.
Drosophilapseudoobscura
* . 10%
(.9.9).
,.
,,,
.,
.,(1956)
,:
35,
.
P1
.
.,
,
, . ,
,()
,
(. 9.9). , ,
.,
aaBB,AAbb,
B B
B B
P2
F1
F2
.9.7. .,
.P2,
Lyiaglandulosa.P1,
,.
F2,
12.
.Asteraceae
(FromClausenetal.1947)
H H H H
H H
200
9
1
+1 +2
, CyL/Pm .
, (,
+2) CyL/Pm .
+1
Cy L
+3
Pm
+2
Pm
+4
Cy L
Cy L
Cy L
Pm
+2
Pm
Pm
Cy L
+2
+3
Cy L
5
.
.
CyL/Pm
.
Cy L
P
3
2 +3 +4
+2
Cy L
Cy L
Pm
+2
Pm
+2
Cy L
+2
Pm
Cy L
,
(, +3)
CyL/Pm
.
Pm
+3
Cy L
Pm
+3
1/4 (+/+)
(Cy, L
Pm) . 1/4-
+/+, +2
( ) , +2/+2
.
+2
Cy L
Cy L
Pm
+3
+3
+2
7
Pm
+2
+3
+3
+3
.9.8. Drosophilamelanogaster
,.+
,.
(+1,+2+3,+4),
,
.Cy()L(),
Pm().
(.8).(,+2)F3
.F3,
.+/+CyL/+Pm/+.
F3.1:1:1:1
.,
,(+2+3).
1:1:1:1(AfterDobzhansky1970.)
AaBb,,
,.
(q<0,01)
.
,,
.,
,,,
..
(.9.10).,
2753%.
40
,
.
30
20
10
0.12
0.24
0.36
0.48
0.60
0.72
0.84
0.96
1.08
1.20
,,.
(Stern1973).
.
,.
(Frankhametal.
2002.9.11)..
(1995,1999)(Viperaberus)
,40.,
,(
).
20
. .
(.9.12),
.
.
,
.,,,
.
, .
,()
.,
.
1966Drosophilapseudoobscura.
.,
(),
,.
.,
(,,),
,..
.
.
,,
21 , (%)
.9.9.
.
Drosophilapseudoobscura9.8
.
1.00
F3
.
(
)
195.
,
.,,
.
(AfterLewontin1974).
50
201
30
27.8
25
20
28.9
22.7
20.2
1
- -
.9.10.
.
.
,
,19031907
.
21
(AfterStern1973).
202
()
()
.9.11. ()
(Lentopithecusrosalia),
.140
.
,500
.
()
,
.
()
()
(Tom&PatLesson/Photo
Researchers,Inc.FromFrankhametal.2002.)
.,
,
.,
.,.
(.9.13).
,,
.,
. ,
.
,,
.
35
30
25
1992
20
.
20
15
10
.9.12.
(AfterMmadsenet
al.1999.)
5
0
1981
1983
1985
1987
1989
1991
1993
1995
1997
1999
1996 .
203
Drosophila pseudoobcura 18 . ,
.
,
.,
(..9.5A).
2
,(H) 1 p i ,pii
,pi2AiAi.18(
)0,12.
,12(
H ).
(1972),7128%
,( H )0,07.
.
.
(seealsoLewontin1974),
.,
(
300033000=101431,
).,
.,
.
.:
?
. 1966
,
(..9.1).,
.,
.,
.
(),
,.
.
.
(1983).272111.Drosophila
melanogaster..
(2/5, 1/2)
(2/2)
1
2
3
4
5
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18
.9.13. (Fundulus
zebrinus)13
.()
.1,
.,
.
2/2,2/2,2/2,2/5,2/5,2/3,3/3,2/4,1/2,2/52/2,1/2,2/2,2/2,1/2,2/2,2/2
(CourtesyofJ.B.Mitton)
204
9
.(AdhF)(AdhS).
11,
.
.
Adh , ,
(.9.14).1143
/.
(1,8%),(2,4%).,
14,,
.,AdhFAdhS
.
,
,,
.,
,
(). ,
().Adh=0,0065,
D.melanogaster(=0,05)(Li1997).
=0,0008.
.9.14.
DrosophilamelanogasterAdh.
()
()
.
.
43,
6()(
),.
11
.
27
,
()87
.
(AfterKreitman1983)
()
(()
,
)..,
,
.,A1B1
,B2,A1B1.
,.A1A1B1B1,
A2A2B2B2..
A1B1/A1B1,A1B1/A2B2A2B2/A2B2(
,).
.A1B1A2B2.,
().,
()(,
.
,,
).
. ,
A1B1A2B2,
AdhF AdhS
.
.
/ .
200
400
600
800
1000
1200
1400
1600
1800
2000
2200
2400
2600 bp
. (A1B1/A2B2) A1B2 A2B1
A1B1 A2B2 A1B1/A1B2 .
(A1B2,A2B1)
,
(.9.15).,
.
,,A
,B.
()
..A1A2
pAqA(pA+qA=1),B1B2pBqB(pB+qB=1),
()A1B1pApB,
A1A1B1B1(A1B1/A1B1)()(pApB)2
pA2pB2.,
.,
.
,
,.
,.,Primulavulgaris
, .
(),
()(.9.16).
,
..
,
.,,
.Gg,,,Aa
,.
GA/ga,ga/ga.GagA
.,
.,
.
(.17),
.
,(.17).
,
.
,..
.
. (
,),,
( Drosophila
). ,
(.9.17).
.
.
( ), :
205
0.500
0.5
0.5
0.375
0.125
AA BB
A A BB
AA BB
AA BB
AABB
AAB B
A A BB
AABB
AA BB
F
0.5
0.500
0.375
0.25
0.25
0.250
0.125
AA BB
A A BB
AA BB
AA BB
AABB
AAB B
A A BB
AABB
AA BB
0.125
F
0.500
...
.
0.375
0.31
0.14
0.375
0.375
0.125
0.125
F1
...
A2A1A1B1B1
,B2.
.
.
,
,
0.14
0.09
0.09
0.02
0.09
0.02
A A BB
AABB
AA BB
0.09
AA BB
A A BB
AA BB
AA BB
AABB
AAB B
0.125
.9.15.
.
(A1A1B1B1A2A2B2B2)
.
.(
),
.
(F1),
(A1B2,A2B1),
(A1B1,A2B2),
(A1B1/A2B2).
,
.,
.
0.500
0.375
0.250
0.5
0.250
0.125
0.250
0.5
0.375
0.250
0.500
A B
AB
A B
AB
A B
AB
A B
AB
206
a A .
g G
.
GA / ga
ga / ga
.9.16. (Primula
vulgaris).
.()
,
(AfterFord1971.)
gA ,
.
gA /ga
207
(,
).
,
,
.,
.
,,
.(1982)Drosophila
pseudoobscura 10 .
.,
,
(.9.18).(
),,
.
.
,
.
,(.9.19)
.
(
)..
.
(.C).
(Vp)(VG)
(VE),VP=VG+VE.,
(),
,(
).
VG,()
VE.
,,
h2.:
h2=VG/(VG+VE)
VGh2
*.,
,
(.9.20).,
.,VG/VP(h2
)1.0.
,,
160
140
120
N=85
100
80
60
40
20
0
20
60
100
140
180
220
260
300
.9.17.
.825.
(
).
.0,95(AfterHolt1955.)
208
35
.9.18. .
()Drosophilapseudoobscura10
.
.
,
.()510
(170C200C),
,
,
(170C200C
,
y,
).
,
.()230C250C
,,
(AfterGuptaand
Lewonntin1982).
Drosophila- 10
()
9
1
5
2
10
8
4
3
6
7
30
25
14
21
26
()
()
, 170C- 200C- . .
, 260C- 280C- .
.
g10
25
g5
g10
g5
30
32
25
14
21
26
14
21
26
,.
(1983)
(Geospizafortis).G.fortis
,.
.
(.9.20).,
0,9
90%,10%.
,.
.
,,h2,,
0,10,9(e.g.,MousseauandRoff1987).
(,)
,
,
.
1.0
0.75
0.75
p = 0.5
q = 0.5
.9.19. (),
.
()p=0,5,q=0,5()p=0,9,q=0,1.(
).
,
,(
B
).VG()
0,5,()0,472.
.
209
0.50
0.25
p = 0.9
q = 0.1
0.50
0.25
(
).
, ,
, .
9
9
10
11
10
11
,
AA
AA
AA
AA
AA
AA
()
()
,,
(0,840,94),
(0,96)
.9.20.
(LynchandWalsh1998).
.
.
, ,
()
.
().
,,
,
(),.
.,
(1969)Drosophilapseudoobscura20
.
,
.
.(
11.0
.
.
= 0
()
= ~ 1.0
,
.(),
.
1.00.(),
.
0.00.
()Geospizafortis1976
1978.1978,
.
0,9(afterGrant1986,
basedonBoag1983.)
10.0
G.fortis
= 0.9
1978
9.0
1976
8.0
8.0
9.0
10.0
()
11.0
210
9C. ,
,.
(,)(
) . Xi
i (,
X3=10).n,
X1+X2+...+Xn
k
X
i 1
Xi
).:
(,),
X1n1,X2n2..k
.:
n1X1 n 2 X 2 ... n k
n1 n 2 ... n k
nin:
9(X1=9)n1=16,10
(X2=10)n2=4811(X3=11)n3=36.
,(k=3)
:
.
,
() .
.
?
( ) ,
.
,515
,
.,
,
.()
V=
ni/n=fi,Xi
:
x (f i X i )
i 1
(X1 x ) 2 + (X 2 x ) 2 + ... + (X n x ) 2
1
=
n i (Xi x ) 2
n1 1
n 1 i =1
n
n1 .
:
, n=100
),(
).
,15.
16(.9.21).,15
,.,15
,16.
89.16
.,300
.2525,
(),(),
.20.
89
(.9.21).
.
.,,
.,
,0,09.
,Drosophila,
( ), , (
, ), (,
211
V=
,
,
.,
(.4.22):
s= V
0,485
s=
=0,696.
,91,10
181181
. x =10,8,V=0,149s=0,387.V
s,,
.
, ,
(. 1).
(
)
68%
,96%
,99,7%
3. ,
, x =100
s=5(,V=252),68%95105
,96%90110.
, s=10
(V=1002),68%90100
.
- 3
-2
68%
96%
99,7%
.1. ,
.,
(s)x.
,
,
.
()s.
ss.
),(,)
.,
,,.
.,
,.,
,,.
,.,
(Cyprinodondiabolis),.
.,
.
.
212
()
.9.21. ()Drosophilapseudoobscura.()
.,
.
..
153.
()20,
(AfterDobzhanskyandSpassky
1969).
( 3)
( 2)
( 1)
()
1
3
5
7
9
11
13
15
2
,
,
(synpatra).
,
.
.
(.9.22).
.1516.
, ).
,
(.9.22).,(Colaptesauratus)
, auratus cafer,
.,,
(Short1965MooreandPrice1993).,
(.9.23)
(,
8 10 12 14 16 18 20
).
(.9.22).,Elapheobsoluta(.9.24)
.
,
,(Burbrinketal.2000).
(..9.32).
,()
.
(.9.22).,(Odocoileusvirginianus)
.
.
,.
.
,
,.
Drosophila melanogaster (. . 9.14)
(),
AdhF()(.9.25).(
).A,
(Oakeshottetal.1982).
(.9.22)
.()
.
,,
.9.22. .()
,.
().()
(,
).().()
,(
).
213
()
()
.,
.
,(1940)
.(Potentillaglandulosa)
.,,
,
.
, ()
:,.
,.
,,.
.,
.,
,,
(.9.26).
()
,.
.,
.
.,,
(),
(
)..,
(cafer)
(auratus)
.9.23.
(Calaptesauratus):(
)(C.a.auratus)(C.a.cafer).
(AfterMooreandPrice1993.)
214
9
Elaphe obsoleta obsoleta
.9.24. Elapheobsoleta.
,
.
,
.,
(AfterConant1958)
E. o. quadrivittata
E. o. rossalleni
E. o. lindheimeri
.
E. o. spiloides
.,
,.
.,
(Ranaclamitans),
,.
(),,
,(Bervnetal.1979).
,
.
.
,..
,
(.16
18).
(Grant1986).,,
,Geospizafortis,G.fuliginosas.
.(),G.
fortisG.fuliginosa.
(.9.27).
()
()
AdhF
..
(),
().,
,.
(,)
().
.
..
()
215
()
Potentilla glandulosa
.9.25. Drosophilamelanogaster
AdhF
()
().
AdhF
.(AfterOakeshott
atal.1982)
90
hanseni
80
()
70
nevadensis
60
50
typica
40
30
20
10
P.g. typica
()
P.g. hanseni
( )
P. g. nevadensis
( )
30 . .
1400 . .
3050 . .
.9.26. (Potentillaglandulosa).(),
1400
.().
,
(AfterClausenetal.1940.)
216
()
G. fulginosa
0.40
0.30
0.20
0.10
()
G. fulginosa
0.40
0.30
,
.,,
(),A(.
1012).,
,,
(.9.28).
(m).
,
.
,.
(McCauley1993).
,
.,
,
.
.,,
,
.A.
(Danausplexippus),
G. fortis
,
fulginosa
. ,
.,
fortis
,
,.
, (. 10 ),
, , .
.
, (1943)
Drosophila pseudoobscura 4000 ,
3
4
5
6
7
8
. ,
(
()
),.(
),
.(
)250.
10000050.
,5001000.
,
.15
G. fortis
(Coyneetal.1982).
(1947)
fulginosa
( )
()
.,4050
1%,
fortis
0.20
0.10
()
.9.27.
.
..()
G.fortis,G.fuliginosa.()
(AfterGrant1986photoscourtesyofPeterR.Grant.)
(. 9.29). ,
(Levin1984).
0.8
5
4
0.6
0.4
0.2
() ,
.
1.0
(p)
.9.28.
.
..
m=0,1(AfterHartlandClark1989).
217
= p
3
2
1
10
20
30
.
,,:
Vq
FST =
( q )(1 q )
q ,Vq
.GST.OFSTGST
0()1(
).
(1998)GST.
Wyeomyiasmithii,
.
(,
)(,,).
.
,
(GST),(.
9.30). , ,
,
.,
,,
,.
(Nei1987).,
.*.
,()
()().,
,
(.9.30).,
,.
,(),
(.26).,
,(Oporornistolmiei)
b(A)(.9.31,).,
A,.
,
(.9.31).,
.
40
50
218
9
,,A
.,
..
A
(Miletal.2000).
,
,
90%,10%
.
.
0.7
0.6
0.5
0.4
.
0.3
0.2
0.1
10
20
30
40
50
60
()
70
80
.9.29. .y
,
,.
,
.(After
Bateman1947)
0.8
Gst
0.6
, ,
.
0.2
0.0
0
1000
2000
3000
()
FL1
FL2
A L1
FL3
AL2
AL3
AL4
AL5
MS
W .M . .
W .V. .
.9.30.
Wyeomyiasmithii.()
GST()(
)
,
.
.(),
.,
.(MB,ON),(ME),(NJ)
(PA),
(NC)
(Fl,AL,MS).W.M.W.V.Wyeomyia
(AfterArmbruster1998.)
MB 1
MB2
ON
ME5
M E1
ME2
ME3
ME 6
ME4
NJ 1
NJ2
NJ6
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.:TheodosiusDobzhanskysGeneticsofevolutionaryprocess
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.ErnstMayrsAnimalspecies
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:RichardC.LewontinsThegeneticbasisofevolutionarychange(ColumbiaUniversityPress,NewYork,
1974),
.
:PronciplesofpopulationgeneticsbyD.L.HartlandA.
G.Clark(ThirdEdition,SinauerAssociates,Sunderland,MA,1997),
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241
242
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byM.Kimura(CambridgeUniversityPress,Cambridge,1983)
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248
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300
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(ReprintedwithpermissionofCronicle
Features,SanFrancisco.)
257
258
11
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260
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,
. : Natural selection: Domains, levels, and Challenges
(OxfordUniversityPress,NewYork1992)
R.Dawkins,Theselfishgene(OxfordUniversityPress,OxfordandSanFrancisco,1989)Theblindwatchmaker(Notron,
NewYork,1986).
.
:Thenatureofselection:EvolutionarytheoryinphilosophicalfocusbyE.Sober(MITPress,Cambridge,MA
1984)ThecomparativemethodinevolutionarybiologybyP.H.HarveyandM.D.Pagel(OxfordUniversityPress,
Oxford1991)..
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:
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.
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:
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:
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276
12
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100
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Nm
.
..
.
,,
.,
,
.12.12.
(.12.12StorferandSih,1998Storferetal.1999).
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(O),
,
.Nm
AE
FST(.10).
()
.
,
.
,
,
Daphnia,
.
,
,
,
,(AfterStorferandSih1998Storfer
etal.1999.)
24
80
40,()
.
,,.
,9,
.(1),
.,(2)
(
), (3) ( ) (4)
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(
).
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,.
,
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.
,
()(.12.13.
A.12.7).
281
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.
0.1
0,25 0.1
.
0
0
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:
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,
w = 0.90 1.0 0.95
w = 0.90 1.0 0.90
.
s = 0.10,t=0.5
s = 0.10,t= 0.10
,
()
(Aa)
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(AA,aa).
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,
0.7
(
AbaB,AB
),
0.6
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0.5
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)
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, ,
.
.
0.2
()
,
,
0.1
,
.
.
0
.
AA AS SS
(
).
w = 0.90 1.0 0.20
,
AA.AS
s = 0.10,t=0.80
.
,
SS
. Arabidopsis thaliana
.
, ,
(MitchellOlds 1995).
((CavalliSforzaandBodmer
1971).(S),
A
.S.
,
.(AS),
(SS)().
.(AA)
,(AS),
(Plasmodiumfalciparum).
,.,
(.12.13)S
Y,,(q=0,13)(
WAA=0,89,WAS=1,WSS=0,2).
..
,,AA
.,,S
0,05.
,
.2.
a(
.
.
.,
282
12
(CleggandWeatherall1999).G6PD
.
150
100
50
10
0.2
0.1
0.0
10
.,
.,(Curtsingeretal.1994).
,A1A1A1A20,5,
A2A20,4.
,0,60,9.
,A1A2,
.A1A1A1A2
(0,5)(0,6)=0,3,A2A2(0,4)(0,9)=0,36.
,A2A2A2.
(),
().
, , ,
(Felsenstein1976Hedrik1986).
,
,.
,
,(
).,
,.
.
,
,
.
,
,.,
(
).
.
,
(,
,
.
,,
12.8
15.7
18.5
,,
,
.
(Pyrenestesostrinus)(Smith1993).
(.12.14).
.
12.8
15.7
18.5
.
()
.
,
.2700,
,
.12.14. .()
,.()
.
(AfterSmith1993
photographscourtesyofThomasB.Smith.)
283
(.12.14).,
.
..
,.
,14.
.
,(.12.15).
,(A1A1A1A2)
,.,
.(A2A2).,A2
,,A2A2A1A1A1A2
.A1.,
,,(
q2=0,5).
.
.
.,
.S1,S2S3,S1S2S3
,S1S2S1S3
.S2 ()
S1S3,S3
1.0
S1S2(,
W11
W22
).
0.8
S
W12
, .
0.6
1/k,kS
. ,
0.4
.
.
0.2
(1993)
0
0.2
0.4
0.6
0.8
Perissodusmicrolepis.
A1 p
. () ()
() (. 12.16).
.12.15. ,
.()
.
,(pA1).
A1A1,A1A2A2A2W11=1sp2,W12=1spq
W22=1sq2.s=1.
p=q=0,5.()
,
.A1A1,A1A2A2A2
1+sp2,1+2spq1+sq2(AfterHartlandClark1989.)
1.0
0.8
p=0.
0.6
W22
p=1.
W11
W12
W12
0.4
0.2
0.2
0.4
0.6
A1 p
0.8
284
()
12
()
.12.16.
Perissodusmicrolepis.()
.()
.(after
Hori1993.)
1.0
0.5
1980
1982
1984
1986
1988
1990
,.
.0,5(.12.16).
.
,
.
.,,.
,
.,.,
P1P2,P1
,P2.
P1,P1P2
P2.
P1,P1
.P2
.P2.
Anthoxanthumodoratum.
,.
(1984),.
,
,,
.
(),
.,,
. ,
(,
).
.
,
.
.
(1:1)?,
,,
.
,,
,?
..(1930),
,,
.
285
,,,
,
,.
,
(S)
(s).,
(
).
.
.
,
.
, 0,25 (1 3 ),
.,
4.4.,
,12(
3).,
4,12.
(34)+(112)=24.,0,50(2
2). 24=8,
2x12=24.32.,
.
,,
.,
,
.,(0,5)
(.12.17)(0,5
(ESS).14).
(1994)
Xiphophorusmaculatus.W,XY.XX,WX
WY,XYYY.0,5
,XXYY.WXXY
0,25,
.
.,
.()
0,25%0,78%.0,5,
(.12.18).
,,
,.,
4
S=0.75
.12.17. .
(s
).
(S).
2.S=0,2525%,
s0,25,.
.S=0,75
.,s0,75(AfterCharnov1982.)
S=0.25
2
1
S =0.25
0
0.25
S =0.75
0.50
0.75
(s)
1.0
286
.
,
,.
(
)
0,5- .
1.0
0.8
12
0.78
0.6
0.4
0.2
0.25
.12.18. Xiphophorusmaculates
(
).
0,25%0,78%.
0,5
(AfterBasolo1994.)
, . ,,
(..12.15).
,Heliconiuserato
..
.
.
Heliconbius,
(.12.19).,
,
.
(1989),Heliconiuserato
()
:,
, ,
.,
.
H. erato ,
.
,.,
,
,.
. ,
.
.0,52
.,,
,s=0,17.
.
,
.
A1A1A2
, A2
(A1A2) . A1A1 ,
A2(q)0.,A2A2
A1,A2A2A1A2.
,A1A1A2A2
.,
Heliconius melpomene
Heliconius erato
.12.19. .
Heliconiuserato
.
Heliconiusmelpomene.
.H.erato
,.H.melpomene(FromCornell
UniversityInsectCollection,courtesyofAndrewBrower.)
.,
,
(..8.218.22).
,
,,.,
.,,
.
( w )
(p), w p(.
.12.7,).,,
( w ).
:
, w .
,p=0p=1
(..12.7).,p
, w .
12.7
.12.7,
.
. . (
).
()p2,2pqq2.p
x,
.17, w p
,(p=0p=1
)( w ),
.,
,
,
w p.
,.
.
( ),
(Ne)
(s).
,
,,s1/(4Ne).s1/(4Ne),
,:
.
:1)
,
:
2)
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.,
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,
.,
.
,
,,
287
12
()
.12.20. ,
.
,
,A1A1,A1A2A2A2
1,1s1.
( w )
p,A1
,(p=0
p=1)
.
p=0,5
.p
.()A1
0
,
,.()
,
p
(0,5).
A1
,.
A1- ...
... 0- ,
/ .
A1
0.5
0.5
()
288
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/
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... A1- ,
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0.5
0.5
P
0.5
0.5
(..12.7).
,,:,
12.20).
,
,(.
12.20).(BartonandCharlesworth1984)
(),.
,
,,
.
,,
.,
,(.8).
, Vandiemenella viatica
.
.
,,
.,
200300
(White1978).,
,
.
,
.10,
289
4N e u 0
4N e u 0 1
()
(5 )
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,,
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(.9).
,
,
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.,
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,
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,
.,
,,
,
, ()
(7 )
,
()
(12 )
()
(4 )
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,
(
10.15).
(12),
.
().
,
.
.()
.
7.()
.
,
5.()AA
.
()
.
,
.().
x.
290
12
,.
.
,(.12.21),
(. 12.21).
,,
().
.
,
,
,.
(,
).,
.(,
).
, ,
(. 12.21). ,
(Stribeck1983).
,,
,
.,
.()
,.
.(Charlesworthetal.1993Charlesworth1994)
. ,
,(.
12.21).,,
,.
,,,
.
(1994)Drosophilamelanogaster(
)
.19(
,.10)
.,
,.,
,.
,
(),(Andalfallo
andPrzeworski2001).
(..9.25),
Drosophilamelanogaster(Adh)
,1490
(.9.14).1490
Adh,(.12.22).
.
. Solanaceae
,
(30,.,)
,
.12.22.
DrosophilamelanogasterAdh.Adh
Adhdup.
.x
,0Adh
.Adh.
,
D.melanogaster
.
1490
,,
(AfterKreitmanandHudson1991.)
5
4
1490-
. .
2
1
0
-1500
-500
500
1500
2500
Adh
291
3500
Adh-dup
(.12.23).,(MHC)
,
(.12.23).(),
.MHC,
,
(NeiandHughes1991).
.12.23.
()
.()Solanaceae
.
Nicotianaalata
,
Nicotiana,PetuniaSolanum
.()(MHC)A
B
.
AB.
.,
,
.,
.
5(after
Loegeretal.1990afterNeiandHughes1991.)
()
Nicotiana- ()
, .
, Nicotiana-
.
()
H-A3
H-A11
C-A108 MHC
A
C-A126
H-A24
H-B13
C-B1
H-B44
C-B2
H-B27
0.08
0.06
0.04
0.02
MHC
B
(
),
.
292
12
Med A/A/
G6PD (AC)n (ATC)n (CTT)n 4.28
6.79 7.54
G6PD (5 ) X .
.G6PD
,G6PD
.,
20%:AMed,
.
(B).
(2001)G6PD
*(.12.24).
(),BA
Med.AMed
,B.AMed
().
,A,
Med(.12.24).
,AMed
.,
.,G6PD
.
,
,,,
,
.A
,A(s)0,0446357
.117603840
()
G6PD (18)
3 4
6 7
10 11 12
()
13
4.28kb
6.79kb 7.54kb
Med A
, .
60
40
20
.12.24. G6PD
.
()
AC,ATCTT
.G6PD13
.A45
,
Med6.
()
G6PD
,A
,Med
.
149
.
(After
Tishkoffetal.2001.)
Med
21
AC
/
41
AT
/C
61
TT
G6
81
PD
101
121
0
A- (- )
141
Med ( )
293
Med0,034,3330
(66401600).,Plasmodium
falciparum ,
10000(Volkmanetal.2001
Joyetal.2003).Plasmodium
.
10000.,
Plasmodium.
20 30
,.30
.
(Bistonbetularia)
.
.19
,
,()
1%,,90%.,(
50%)(Haldane
1932). ,
,
(
).(.12.25)..
(Majerus1998).
,,
(. 12.25 Grant and Wiseman 2002 Cook 2003).
(.A),
(s)0,05
0,2.
.,s,
.,
(.12.26).,,
,.,
.,.
.12.25.()
(Bistombetularia)
....
,
,
.()
.()
.
(,,photolibrary.com
afterCook2003.)
1.0
0.5
0.0
1950
Biston betularia-
3
50 .
1960
1970
1980
1990
2000
294
12
()
()
10
10
0.5
1.0
.12.26. ,
(s)
.
,
,.N
,
.()(
)(AfterEndler1986.)
0.5
1.0
1. ()
,.
,.
2. . :
,.
()
.
3.
.
.
4.
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,
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5. ,:
,
.
6. ,
.
.,
,.
7. ,
..
,
.
8. .
, ( )
()
.
,.
9. ,
,
.
(
)
()
()
Naturalselectioninthewild,byJ.A.Endler(PrincetonUniversitypress,Princeton,NJ,1986),
.
D.L.,HartlandA.G.ClarksPrinciplesofpopulationgenetics(thirdedition,SinauerAssociates,Sunderland,
MA,1997)P.W.HedricksGeneticsofpopulations(JonesandBartlett,Sudbury,MA,2000).
.
1. ,0,05
,
?(:q=0,048p2=0,9071,2pq=0,0907q2=0,0023).
.
106,?
2. ,A1A180%,A1A2,
A2A295%.A1(p)
?
?,,,
A1A1,A1A2A2A21,0,950,9
.A1?(:0,208).
3. A1A1,A1A2 A2A2
90%,85%75%,50,55
70.(R).
?,
,.
50,5570,70,6555.
,?
55,65,75,75,65,55?
295
296
12
4. ,,
,,
,
(MIttonandGrant1984Zouros1987).
(seereferencesinAvise2004).,
.
5. ,
:
??()
.
6. ()
()()
.
7. (e.g.,Wells2000)(Hooper2002),..
.(2002),(2003)
.:
?
.
13
,
.
,
,
.
:,
.
.
,
.
.
, ,
.
(
).
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,
),
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Painet,Inc.bassetLynnM.Stone/
naturepl.com.)
( )
298
13
15.5
Chilo ()
(
15.0
),.
Cydia ()
,
,(
Cydia-
14.5
Chilo-
).,
(),
.
Chilo-
14.0
(Endler 1986Taylor et al. 1991). ,
.
,
.
13.5
(
32
34
36
38
40
42
44
46
)(
(N)
).,
,
. 13.1.
.
.(Cydiapomonella).
. Chilo suppressalis
1750.
. Cydia pomonella .
12.
250 .
,
.
(After Tauber et al. 1986).
(.13.1).
50 ?
(Sylviaatricapilla)
.50.
.,.,
(
).
(1992),
()
,
..
.,
,
,,
.
,.
.
700100,
(.13.2).,.
. 13.2.
Mimulus guttatus- . () ,
. .
(From Macnair 1981; courtesy of M. Macnair.)
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.
..
.
()..
(AfterCarrollandBoyd1992.)
299
: 6.05
: 6.680.82
.
..
,
,
(Antonovicsetal.1971Macnair1981.)
, (Jadera
: 7.09
7.230.47
haematoloma)(CarrollandBoyd1992Carrolletal.1997)
.
(Sapindaceae)
(.13.3).,
.2050
.
(Koelreuteriapaniculata).
.
.,
.,
.K.paniculata
8%,
.K.elegans
25%,
..
,.
9
.(V).
.
(s= V ).,.
(),68%
,96%
,99.7%(.C,9).
(VP)(
VG)(
VE).VP=VG+VE.
,
(.13.4).A1A1
+a,A2A2a.a
..
,
.
.VA.(VG
,
).
: 11.92
9.320.86
: 2.82
6.930.48
300
13
VE
. s
A2 A2
A1 A 2
A1 A 1
,
.,
,,
,,
.13.4,(pq)
,VA:
VA=2pqa2
6
10
.13.4.
.
,
.
A1A2
a(
a=2).a
VA.
VE.
,
.VA
.
,
.,
*.
.,VA.
.
,h2N.
(VA),
(VE),,
.:
h2N=VA/VP
VP=VG+VE,VGVA.
,
,.,
,0,75,3/4,1/4
.,,
..
.
(Houle1992).
.
,
.
.
(h2N) (y)
(b)(x)(.
13.8..9.20).,y
.
,(13.8).
,,(
,),.
VAVA,
().
,
,,
.
,
.
.:,
.
(.13.5).
X1Y1,
X2Y2.F1(X1X2Y1Y2xX2X2Y2Y2)
.,X1X2Y2Y2X2X2Y2Y2,
.,
,
X.(34213.5).Y
,X2X2Y1Y2X2X2Y2Y2,
,.(13.5
243).
,(YZ)
.Y,Z,
F1.Y
Z,
,
.,(13.5
4546.,
).
,
,,
,(,
1
L=8
L=4
).
X2
X1
Y1 Z1
,
X2
X1
Y1 Z1
(,
L=6
L=4
X2
X1
Y1 Z1
),
F1
X2
X2
Y2 Z2
. QTL
(QTLQuantitativeTraitLocus
).QTL
X1
Y1 Z1
1 2 ,,
(1)
.
X2
Y2 Z2
(
X2
Y1 Z1
) 2
.,
.
(2)
Y2 Z2
X2
,
X1
Y2 Z2
.
(3)
.13.5. ,
.
(L).
(X,Y,Z).F116
().L
,.
.
,23,
,
.
(4)
(5)
(6)
Y2 Z2
Y2 Z2
Y2 Z2
Y2 Z2
L
6
5
X2
Y2 Z2
X2
Y2 Z2
X2
Y2 Z2
X2
Y1 Z2
X2
Y2 Z2
X2
Y2 Z1
X2
Y2 Z2
301
4.4
4.6
X ,
X
.
Y Z
,
.
302
13
,
.
QTL
.
Drosophilamelanogaster
(e.g.,Nuzhdinetal.1999DildaandMackay2002).
().,
.
.
(Longetal.1995).
X3
.
53QTL,33,31
, 11 (. 13.6). QTLi
,
..QTL
:,
.,
.
QTL ,
QTL(
QTL,
9..9.18).
QTL,
:
,,
.,
Drosophila
(Lai et al. 1994).
.,
(12).
(Mackay2001DildaandMackay2002).
QTL
.13.6. ,
.
()
.3QTL
,Drosophilamelanogaster
.
()
.,
()()
.,
()()QTL
.
,
QTL.
QTL.
,.
,
.QTL
,,
.
QTL.QTL
,(FromDildaand
Mackay2002).
,
.
.,
.
, :
,.
,(
),.
,,
,,
(.9).
.
.
,A1A2pAqA,B1B2pB
qB.(A1B1,A1B2,A2B1,A2B2).g11,g12,g21
g22.AB,
(,
A1B2g12=pAqB).,
,,A1
B1pB,B1.
,
,
.
,:D=(g11xg22)(g12xg21).D>0,A1B1A2B2
(,A1A1B1B1).
A1A1B1B1A2A2B2B2
(),
,
().D<0,
.
,D.
,(D>0)
.
(.13.7..9.15),
.
?:
1..
2.,
.
.
,
3.
,
4. .
()
5..
,
.
,
303
304
13
(Dt/D0)
1.0
R=0,5,
.
R =0.001
0.9
0.8
.
.
0.7
R =0.01
0.6
.13.7.
(R)
(D)
(D0).R=0,5
.(After
HartlandClark1989.)
0.5
0.4
6.,
,
0.3
R =0.1
0.2
0.1
0
R =0.50
10
20
30
40
50
60
70
80
(t)
90
100
(LynchandWalsh1998).
.,
(),
(MousseauandRoff1987).h2N
,
VE,h2N=VA/VP.
(VA),
(Houleetal.1996).
.
,
(,,
),.,,
,
(..13.2),.
.
. ,
,
,,,
(Macnair1981Bradshaw1991).
,
.
z(,
),z(
,
).,
.
.( z )S
,(.13.8).
( z )( z )R
(.13.8,).R
(13.8)
S*(13.8).
R,S,
305
R=h2NS
h2N=R/S.S(
) R ,
.h2N
.Drosophilapseudoobscura
0,09(..9.21).
, ,
.,
,(
),.,.,
,.
F1-
( ).
()
10
12
14
16
18
10
S
P
R .
12
14
16
18
R{
S
P
Z s
()
, R-
.
10
12
S
14
16
18
()
S
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320
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Introductiononquantitativegenetics,byD.S.FalconerandT.F.C.Mackay(fourthedition,LongmanGroupLtd.,Harlow,
U.K.,1996).D.A.Roff,Evolutionaryquantitativegenetics
(Chapman and Hall, New Yourk, 1997), .
Geneticsandanalysisofquantitativetraits,byM.LynchandJ.B.Walsh(SinauerAssociates,Sunderland,MA,
1998).
T.F.C.MackaysThegeneticarchitectureofquantitativetraits(2001,AnnualReviewofGenetics35:303339)
.,
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M.PigliuccisPhenotypicevolution:Areactionnormperspective(SinauerAssociates,Sunderland,MA,1998).
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UniversityPress,Princeton,N1999).:L.D.Hurst,
A.Atlan,andB.O.Bengtsson,Geneticconflicts(1996,Q.Rev.Biol.71:317364).
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).
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15
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15
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,
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simulansY().
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1
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3
4
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5
6
7
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8
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simulans D. melanogaster
9
,
200 (Presgraves
10
2003).
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. ,
12
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13
(,D.
14
simulans ).
0
,
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D. mauritiana
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.,
.
(.15.14).
,,
(.15.14).
()
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0.30
()
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.
368
15
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X
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,,.
X,X
(
X).,
,,,
,.
,
,(RitchieandPhillips1998).
DrosophilasimulansD.mauritiana
(..15.8).19
(Zengetal.2000).,
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()
(EZ11).
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E:Z,
,().,
,
,
E:Z,(Roelofsetal.1987).
,,
,.
F2QTL,(1998)
MimuluslewisiiM.cardinalis
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16.
, 25%.
F2,
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,(,,
),.
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).,,
369
(R/S):
D.yakuda
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16/21
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.
/
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(AfterPresgravesetal.2003).
2/8
49+ 1/ 120
,
.
(.15.15).
,.
(.8)
(,.16).
(King1993Rieseberg
2001CoyneandOrr2004).,
()
,.
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3.4,1234.
1.43.2.F1
(1.2,3.4,1.4,3.2).,(1.2
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()(..8.22).(,1.23.2
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3
4
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12
3
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10
9
56
10
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15
27
26
30
30
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370
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annuus H. petiolaris ,
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15.18,2).
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().
NasoniavitripennisN.giraulti.
(. 15.18, 3 4 ).
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).,Wolbachia.A
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,(Werren1998).
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1940
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,
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giraulti
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372
15
.(,.15.11),
.,
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).,
,,
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1
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,,
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b).
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e
a
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b
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373
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p
1
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1
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andAvise1986PamilioandNei1988).
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30
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,
.,
(..2.19).
,
(MoranandKornfield1993).
,
.,
(MHC)
.,
5.
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c
b
,
(,1990).
,
.
(Arnold1997).
(Harrison1990,1993).
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).,,
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374
15
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)
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.
(Szymura1993).
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.
,,,
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.
,,
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()
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()
-20
-10
10
20
30
40
-20
-10
10
20
30
40
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(BombinabombinaB.variegata).
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().()
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(AfterSzymura1993).
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0.99
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)
40
:A.CoyneandH.A.Orr,Speciation(SinauerAssociates,Sunderland,MA,2004).
,
.
:ErnstMayrAnimalspecies
and evolution (Harvard University Press, Cambridge, MA, 1963), : Populations,
speciesandevolution(HarvardUniversityPress,Cambridge,1970):Systematicsand
theoriginofspecies(ColumbiaUniversityPress,NewYork,1942).
:Variationandevolutionintheplants,byG.L.Stebbins(ColumbiaUniversityPress,New
York1950).:V.Grant,Plantspeciation(ColumbiaUniversityPress,NewYork,1981).
:Endlessforms:
Speciesandspeciation,editedbyD.J.HowardandS.H.Berlocher(OxfordUniversityPress,OxfordandNewYork,
1998).,
:Hybridzonesandtheevolutionaryprocess,editedbyR.G.Harrison(OxfordUniversityPress,New
York,1993).
1. ,
.,,,
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(),
(),()
()
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,(1976)(1979).
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377
378
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H. anomalus - H.
annuus- H. petiolaris fallax-
(. . 16.20).
,
(Photo bu Jason Rock, courtesy
of Loren Rieseberg.)
380
16
16.1.
I.
.
1.
2.
.
.
II.
a
. (
)
1.
2.(
)
3.
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ii.
iii.
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.16.1).
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)..
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,(Mayer1963CoyneandOrr2004).
,
(,6.15).
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.
,(.15,
).
,()
.
.
,,
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/.,
(1990)
(Desmognathusochrophaeus).
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381
382
16
()
2.00
()
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ochrophaeus
()
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(
D,
)(AfterTilleyet
al.1990.)
1.0
100
200
2.0
1.0
300
()
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0.4
0.6
D).
.,
. .
1%
(Knowlton1993).
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(1986Avise1994)
.
.
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,
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,
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(Hills1998).
,(.16.4).,
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.,
,
,
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,
.,
.,
.:
Centrarchidae(),Amiidae(,)
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Lepomis microlophus
Lepomis gulosus
Lepomis macrochirus
()
()
pipiens
blairi
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)
.
()
,
.
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pipiens()
Ranasphenocephalasphenocephala
(afterHills1998RodPlanck/Photo
ResearchersInc.SuzanneandJoseph
Collins/PhotoResearchersInc.)
()
sphenocephala
sheri
onca
yavapaiensis
magnaocularis
forreri
sp. a 8 b
neovolcanica
spectabilis
omiltemana
383
berlandeiri
tlaloci
miadis
sp. c
sp. d
sp. e
taylori
,
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,
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.
,,
.
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(1940).
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.
(
).
()
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,
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,
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,
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,
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. 16.5.
,
.
. (Malurus )
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,
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and Volger 2000.)
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(Xiphophorus)
100
50
100
(Ellipsoptera)
(Rhagoletis)
50
16
.16.6.
,
,
A2A2B2B2
.()
(
,
),
.
()
(A1B1).A1A2B1B2
(
),A1B1
()
A1
384
()
B1
A2A2B2B2
A2A2 B2B2
A1A1
A1A2 B2B2
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B1B1
A2A2 B1B2
B2B2
A1A1B2B2
A2A2B1B1
A1 B1
.
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A1A1B2B2A2A2B1B1.A1A2A1A1
A2A2,,
B2B2 .B1B2B1B1B2B2,
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)().A1B1
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).,
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.
, C1 A1A1B1B1 ,
A2 B2 , .
D1,A2,B2C2.
,,
.,
(Orr1995).
,,
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) (. 15 ).
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,
.
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,
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,,,
,,,
(Mayr1963).
,,,
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( )
, (
).
.
,,
385
,
( )
,
.
.
Drosophila
.
SI=0.86
. ,
SI=0.64
.(RiceandHostert1994).
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(
)
SI=0.15
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,,
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.
)
,
(CoyneandOrr2004).,
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bebbianae
.,
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Christie1983).(1998),
.
,,
, (SI),
,,.
.
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(.16.7).
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photobyChristopherBrown,courtesyofDaniel
.,,
Funk.)
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)(,).
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.
,,
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()
()
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0.4
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0.2
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.
,(
afterSchluterandNagel1995afterRundleetal.2000.)
386
16
(Rundeletal.2000).
.,.
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.
,
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,
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:
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.(,
TonyTiford/photolibrary.com,W.Peckover,
VIREO,RobertTyrell/OSF/photolibrary.com
,NationalTrustPhotolibrary/AlamyImages).
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,
.(Lande1981PomiankowskiandIwassa
1998Turellietal.2001).
,(..15.6).
.
.,
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,
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(Mitraetal.1996).,,,
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()
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()
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()
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103
5
387
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(Dobzhansky1951,208).
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.(1963).,
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.
.:
.,
,,
,
.,
,
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,
!(:Grant1966
()
()
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.:
(MllerandCuervo1998).
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).,(Physalaemuspustulosus)
(.14).
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.(RyanandRand1993).
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,
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0
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0
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.
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()
,
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.
.(AfterRitchie2000.)
388
16
andCoyne1974).
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.,
.,
.
.,(Sorexaraneus)
,
.,
,(Searle1993).
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.,
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1
(1989).,
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,
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h.R(Howard1993).
389
()
1.5
1
0.5
0
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0.5
1.5
()
100
(%)
,
,,
(
,
).
,
(Liou and Price 1994
KirkpatrickandServedo1999Ceinetal.1999).,,
(B),
, (P)
,(T),,
.
(P1,P2),
,
()P
T(.15).
,
T1T2
,
,,
.
(Howard1993
Noor 1999). Drosophila persimilis
D. pseudoobscura .
.
.D.pseudoobscura
D.persimilis,
,
(Noor1995).(Hylacinerea)(H.gratiosa)
. ,
.
H.cinereaH.gratiosa
,H.cinerea
(.16.11),H.cinerea
(.16.11)
80
60
40
20
0
()
.
(1954).
(1982).
.,
,,
.,
Utastansburiana
,
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.(Soule1966).
,,,
,.
,,
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,
(
.10).
,
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),
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,
,
.
,
,
(AfterHbel
andGerhardt2001.)
390
16
()
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,
,
.
,.
(1954),,
,
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,,
,
.,
.,
(.421),
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B 1(1972).
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. (1980) ,
()
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.
,,.
(..12.20).,
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(),
(.16.12).,
,
.
A1
,
.,
.
(),
,
.,
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,(,A1A1B1B1A2A2B2B2)
,
.
( w ).
AB
.()
()
.
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,
.()
,
F1
(AfterGavriletsandHastings1996).
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()(CharlesworthandRouhani1988Turelli
etal.2001).,
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),
,.,
,
.,,
,
.,
,,
.
,(Priceetal.1993
Wagneretal.1994Gavrilets2004).,
(
)(.16.12).
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391
,
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,
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,
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15.4..
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.
-
Sim-13
Sim-1
Sim-5
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Sim-7
Sim-8
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Sec-3
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Sec
Sec
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Sec
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()
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Sim-7
Mau-3
Sim-9
Sim-11
Mau-9
Mau-1
Mau-6
Mau-2
Mau-5
Mau-4
Mau-7
Mau-8
Sim-11
Mau-1
D. melanogaster
()
.16.13 (
),
Drosophilamauritiana(Mau)
D.sechellia(Sec)
.
.D.
simulans(Sim)
Mau
SecSim
.,Simulans
(AfterKlimanetal.2000).
392
16
()
4
3
2
1
.16.14.
.()(Muscadomestica)
.
,1,416.
.
().
,
.
.()
Drosophilamelanogaster50,
.
()
():
.
.
(afterMeffertandBriant1991afterRundleetal.1998)
()
10
,
.
.
,
.
8
6
4
2
Drosophilasechellia,
.,
(Kliman et al. 2000).
D.mauritiana
D.simulans.
D.simulans,
. D. sechellia D.
simulans,,
(.
16.13.).D.sechelliaD.
mauritianaD.simulans.,
D. sechellia
.
.Drosophilasechellia,
.
.
,(summarizedby
RiceandHostert1993CoyneandOrr2004).
(1991)(Muscadomestica).,
,
(.16.14).,,
, . (2003 Rundle et al.
1998)DrosophilapseudooscuraD.melanogaster40
(.16.14).
,
.
,
.
.
.
393
(
)
,. ()
.,
,/
2
3 45
.
6
7
8
1
,
65 20 10 35
60
50
60
,.
,.
()
,
, ()
0
..
,
(McCuneandLovejoy1998).
10
()
,,
20
,
2
3 45
6
7
8
1
,
(Endler1977). ()
.,
1.0
,
1964
.,(,)
0.8
,
0.6
.,
1965
0.4
(Endler1977BartonandHewitt1985).,
.
0.2
1
2
3 45
6
7
8
. ,
,
.,
.(1982),
.
.16.15. Anthoxanthumodoratum
.
.,
,,
(CoyneandOrr2004).
..Anthoxanthumodoratum .()
, ()
.
.(.13).
,
.,
,,
,., .()
.,(.16.15).
,. .()
.
,
. .
,
,,.,
.
,.
(1942,1963)
(AfterMcNeillyand
Antonovics1968Antonovisc1968).
394
16
.,
..
(DiehlandBush1989dieckmannandDoebeli1999seeTurellietal.2001).
.
()(.12).
(
),()
..
.
,
..
,.
.(1981),
,.
A()B().A1A1A2A2
,
,.
(B)(,
).B1B1,
B2B2.B2B2
.B2.(
),B2A2(
), B2 A1A2 ,
B1.,A2B2.
B2.
(
).
,(Felsenstein1981).
,,,
,.,
.,,A1A1A2A2
.B
.,.
.(1990)(Tephritidae)
.,
,.
,
(.15.2).
B1,
B2.
A1A1B1B1()A2A2B2B2(
).(,
A2A2B1B1),
.,
,(
).(2003),A2B2
(
) (. 16.16).
.,,
.,,
,(.16.16).
,,
,(DieckmannandDoebelin1999KondrashovandKondrashov1999).
(Gavrilets2004CoyneandOrr2004).
()
1.00
0.75
,
0,5- .
0.50
0.25
0.00
(s)
()
0.50
395
,
,
.
0.45
0.40
0.35
0.30
0.25
0.20
0.15
0.10
0.05
0.00
10
12
14
16
,
.
.16.16.
.(),
,+.
.+(
),(),
,+,
.,
,.()
,
.
s(.12)..
(AfterFry2003).
.16.17. Drosophilamelanogaster,
.
.
.30().
.
(AfterRiceandSalt1990).
100
50
. ,
,
.,
.
,.,
,.
Drosophila,
(RiceandHostert1993)..
,
.,
.
30.
.
(.16.17).
, ,
,
. (Rhagoletis
pomonella)(Bush1969Feder1998).
0
100
10
20
30
( )
50
10
20
30
396
16
.16.18. Rhalogetispomonella,
.
.
(AfterFederetal.1990.)
0.55
0.50
0.45
0.40
0.35
0.30
0.25
39
40
41
42
43
44
45
46
(N)
.
,.,
(Crataegus).
150R.pomonella
(Malus) .
.
,,
(.16.18).(2).
(3
).,.
..
,.
,,
.(Feder
etal.2003).,.
,
(..3.24).,
,
(,..)
..
,,
(Ricoetal.2003).,
(Schliewenetal.1994).
,,
..
.
(Salvelinusalpinus)
,.
(.8).,
.
()
..
.
.
,,
..
35%,
(Goldblatt1979).,
75%,
(Grant1981).
. :
.
,
.,.,
(Solanumtuberosum)
..
,,
.
(..8.20).,
.,
.,
().
,..
.,
,
,.,(Zea
mays),,
(GillesandRandolph1951).
.
(RamseyandSchemske1998).
,(2n)(n)(3n)
.3n(n)
.(,
).
, .
(
).
1930
..
,2n=32Galeopsistetrahit(2n=16)G.pubescensG.speciosa
.
F1.
G.pubescens.
G.tetrahit,
,G.tetrahit
.
.
,.
.,
.
,.
(Dawrlington1939Stebbing1950).
.
, , (Tragopogon dubius, T. porrifolius T. pratensis)
.F1.1950
,T.mirusT.micellus,T.mirus
T.dubiusT.porrifolius,T.micellusT.dubiusT.
pratensis(.16.19)
(1991),
,.
397
398
16
()
()
T. mirus- T. dubius-
-11.2 , T. porrifolius-
-5.9 -5.3 .
T. mirus-
.
2
1.2
6.2
5.9
5.3
T. mirus-
6.2 3.8
, T. porrifolius-
. T. mirus
.
3.8
porrifolius
(2N)
.16.19. (Thagopogon)
.()
(1)T.porrifolius,(2)T.dubius,(3)T.pratensis
T.
mirus(4,12)T.micellus(5,23)
.
,
.O()T.dubius(
)T.porrifollius(
)T.mirus
.
,
,
.
.
(fromOwnbey1950
afterSoltisandSoltis1991).
mirus
(4N)
dubius
(2N)
,
(.16.19).,
.
.
.,
,,
,
..
(Chamerion angustifolium)
,,
,(Husband2000).
,,
(FowlerandLevin1984Rodrigues1996).
.,,
(Levin1983).
.
,
,
.,
(Stebbins1950).
,
,.F1
.
,.
(Rieseberg1997).
(Grant1981).
.
(Rieseberg and Wendel 1993 Riesegerg 1997).
, .
atrorubens
.16.20.
.,
,Helianthus
anomalus,H.paradoxusH.deserticolaH.annuus
H.petiolarisfallax
.
.
,
(RiesebergandWendel1993.)
maximiliani
porter
niveus niveus
praecox
6
5
debilis
petiolaris petiolaris
petiolaris
fallax (1)
2
2
399
petiolaris
fallax (2)
neglectus
niveus
canescens
paradoxus
anomalus
niveus
tephrodes
deserticola
bolanderi
exillis
annuus
agrophyllus
,Helianthus
, Helianthus annuus H. petiolaris
(H.anomalus,H.paradoxus,H.deserticola)
(.16.20).F1,
.
,
.
,
.
,H.anomalus60
(Ungereretal.1998).
()
,.,H.
anomalus,(.
).
.(2003)H.annuusH.petiolaris
..
, annuus petiolaris
()
()
BSI
TFS
6
5
4
3
2
1
0
BSI
.16.21.
.()
(TFS)
.
(BSI)
.()BSI
,
,
.
BSI
.
400
16
,.
. H. anomalus
,,
,.
.
, .
.
5
4
3
2
(Coyneand
Orr 2004). (Time For
1
2
3
4
5
6
7
8
9
10
11
Speciation).,
(%)
(. 16.21).
(BSI),
2%-
,
. BSI
1 .
TFS,
.,,
,(),
.16.22.
TFS,
.
.,Drosophila.
,
800000.
4%.
. ,0,355
,2,6(KlickaandZink1997.16.22).
2
(1997)
,TFS
(..15.1115.12),Drosophila
(AfterKlickaandZink1997).
1,12,7,
0,080,2(,
0
.16.23.
.
,
.I
TFS
.II
(BSI)
.IIIBSI
.
,BSI
(DatafromCoyneandOrr2004)
1. TFS
Drosophila ()
Drosophila ()
.
2. BSI (
)
3. BSI
( )
Drosophila
11
14
25
33
).
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.16.23,I).
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(S)(E).R
,(t),
(N):
Nt=eRt
(7,
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(.16.21).,(E=0),
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),10(
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,(611
.16.23,II).BSI.
(25)(33).
,.
.
,.
,
,.,
, ,
(),,
(BSI).,RhagoletispomonellaHeliathus
anomalus,
,,
(Alpheus)
(3,5)(Knowltonetal.1993).
?
,,
,
(..7.816.9).
, :
,.
,,,
()
(CoyneandOrr2004).,
(
).,
(Schluter2000).
. ,
,,
.
.
(1972)(seealsoStanley1979GouldandEldredge1993)
,
401
402
16
()
,
,
.
()
()
.16.24.
,
.
.
()
,(,)
.
.
,
(.7).,
,
,
. (1954) ,
.
.
,
()
,
(Charlesworthetal.1982).,
.
(2002),,
( )
.,
,
(Futuyma1987).
,
:
.
.,(.16.24).
.
.
,,
(
,),
(,
).(JanssonandDynesius2002).
,
.,
,.(1979),
.,,
,
(Sheldon1990).(1963,.621):
.,
..
1.
.
./
.
2.
3.
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5.
6.
7.
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).
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.
.
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.
,
.
403
404
16
()
15 ,
:Speciation,byJ.A.CoyneandH.A.Orr.
,.
:SergeiGavrilets.Fitnesslandscapesandtheoriginofspecies(PrincetonUniversityPress,Pinceton,
NJ2004).
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2. (1997) ,
,.
.
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).
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,
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()?
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)
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.
.
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.
17
.,
, (Pinus aristata)
4600,
(Poplus tremuloides)
10000.
,
, ,
,
.
.
,
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musculus),
.(Apteryx),
25%(.17.1).
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4600.
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406
17
.17.1. (Apteryx)
,
25%(Photocourtesy
OtorohangaZoologicalSociety.)
.
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,
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,,(.17.2).
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,.
.
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.
,
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.
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,
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()(Aphisnerii).
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:
.
.,
,
.,,,,11
.,
.
,
,.
,(),
(),
.
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,,
(
)(Williams,1966).
.
().,
.,
,,.
,
,
(Stearns,1992
Charlesworth,1994bRolf,2002)..
12,
().
R.
(L)
(M):
R=LM
.x,lxx
(x),mx
x()(17.3lxmx
Drosophila).,1,2,34
0,4,80.
75%,50%,25%10%.:
x
0
1
2
3
4
5
= R
lx
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0,50
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0
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407
408
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17
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0.2
0
10
20
30
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50
60
70
80
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100
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0
)
30
20
10
0
40
50
60
70
(mx=0),
()
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.
,
()
.,
lx()mx
.,
.
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,,3,
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,
2.
.
,,,2,
(),3.
.
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(r).
.
,
.
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al.1963.)
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,
.
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()
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.
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)
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,
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1x.()A
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.
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:
(Physalaemuspustulosus)
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.,
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.
.
,
.
,
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105
,
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(Reznick1985)
1.
.,
104
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.(.17.5).
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103
,
102
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104
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.
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,
.
(.13).,
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(Lawatal.1979).,
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.
1000
.
3.
(Reznick1985Stearns1992).
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10.
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. 19
,
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0
300
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.
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.
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0
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10
20
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()
40
50
60
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8
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2000
()
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-2.5
-3.5
-4.5
-5.5
-6.5
10
20
30
40
50
60
70 80
()
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10,
,,()
,.()
,
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al.1999andSgrandPartridge1999).
411
:
(.17.7).
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,
(FowlerandPartridge1989BellandKoufopanou1986).
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(
(r)),,
(Charlesworth1994).17.1
,(l1)
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.
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a
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, ,
.
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(1957).
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,(,,
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).
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, (Charlesworth 1994).
,
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:
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(
)(
).
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10
,
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,
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?
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,
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,
,(.17.8WilsonandThompson1989).,,
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,:
413
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2
(PeomislowandHarvey1991ShineandCharnov1992).
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(Poeciliareticulata)(e.g.,Reznicketal.1990ReznickandTravis2002).
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1.25
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.
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60
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80
70
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,(Oncorhynchuskisutch),
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()
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()
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1200
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,(.17.15).
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,:
.,
.,,,
.,.
,
.,
.,.
.
.,,,
.,
.
,.
,
,(Leigh1973Sniegowskietal.2000).
Escheria coli .
E.coli,
(.17.16).,
,(Shaveretal.2002).
417
.17.16. Escherichiacoli
()
.
()
.,
()
.
(After
Shaveretal.2002.)
1
0.9
0.8
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0
(),,,
..,
,,
.
,.
,
().,Chlamydomonas
,.
.,
,.,
,.
.
.
.,
1.8
.,
1.7
..
1.6
.
1.5
,
1.4
, .
1.3
.
1.2
1.1
.Drosophila
0
6500 7000
7500
8000
8500
9000
418
17
(Carson1967Broks1988).
,,.
,
.(Normarketal.2003).
,,
.,:
,
.Taraxacumofficinale.
, Taraxacum
.
,.
(.17.17).,
.
.,Bdelloida
(.17.17Normarketal.2003).,
.
,
,.
,,
.
..,Primula
vulgaris GA
,
ga.,
,Ga.
.
,
.,
,
,(.17.18).
.,
..,
,,.
, .
()
.17.17. ()Taraxacumofficinale
,.
,
,
,
.
.()Bdelloida
.
,
(photoPainet,Inc.
EricV.Grav.)
()
:
(
).
,
(.17.19).,
(
).,
,
.
,,
,,,
.
, ,
,(Maynard
Smith1978Charlesworth1989Kondrashov1993).
(kondrashov1993),
.
419
RAB
RAB
RAB
Rab
Rab
Rab
r .
rAB
rab
rAB
rab
R r ...
rAB
rAB
raB
rab
... r-
, .
.17.18.
(r),
AbaBABab
.Rr
.R,
,,
ABab
.
,.
.
,
.
- .
,
(Bernstein and
Bernstein1991).,
.
.,
,,
(MaynardSmith1988BartonandCharlesworth1998).
. ,
.
,,
,.(Williams1975Bell1982).
(BartonandCharlesworth,1998).
(ratchet).
.(1964)
.,
(.17.20).
0,1,2,...m.
,,.,
,.,
(s )
ssAABb
ssAABb
(S )
SSAa Bb
SSAa BB
SSAABB , SSAa BB
SSaabb , etc.
.17.19.
.
S
s.
,
.
,
,S
2/3,
1/3,
.S
,
aabb
420
17
,(,
).,,
.,,
,,,.
..
,,,
.
, , (Lynch et al. 1993).
,
,
.
,
,,
.
.
.17.20. .
(010)
.
(01)
.
,
.
0
,
(AfterMaynardSmith1988.)
0
3
. ,
,,.
.(1988),
,
.,
( w ).
,,
,.
,.,Escherichia
coli(ElenaandLenski1997).
. ,
,
(MaynardSmith1980).,A,B,C,D...,a,b,c,d...
.
,,AbCdaBcD
(.13).,
,ABCD
,.
(
),,
,
.,
, ,
(
AABBccdd)(.13).
,
7
8
9
10
.
(,ABCD),
,.,
(
,abCD),,
7
8
9
10
.
,
.
(.7),
,
7
8
9
10
:
()
1.0
20
10
0
-10
-20
-30
-3
-2
-1
AB
ABC
AC
ABC
B
AC
3: ,
AB
4: ,
AB
0.6
0.4
0.2
0
.17.22.
.A,BC
.
(13)AB(ABC)
,(A)
(B)
.,
(2),
,
.,
(4)
(Crowand
Kimura1965.)
2: ,
0.8
.17.21.
,
.()
,
.()
,
,
(afterJokelaandLively1995afterLivelyandDybdahl2000.)
(Log )
.
,
.
AB
,,A
, (B) ( ).
AB(.17.22).,
.
,(A)
.
.,.
.,
Chlamydomonasreinhardtii
.
,,
(Colegrave2002).
,
.,
,
,.
,
,(seeWestetal.1999).
,
(),.
1: ,
()
30
(Log )
.
,
(OttoandNuismer
2004).(Jokela
andLively1995LyvelyandDybdahl2000).,
Potamopyrgus antipodarum
,
(. 17.21),
,
(.17.21).,
,
.
421
AB
422
17
,
()().
? , ,
,,
,.
.,
(Charlesworth1978).
.,
,()0,5
(1:1).(Charnov1982Frank1990).
.(
)
(),12.12,
,
,
,,
.0,5
(ESS),
(..12.18).
,
.
,
. ,
..
(1967)
.
,,
,.,
, .
.,
.,,
(),.
,
(.17.23WilsonandColwell1981).
,.
,Nasoniavitripennis,
.,
.,
,,
.,
,,
,,,.
(1980)
,,
Nasonia.
(.17.24)
423
:
N .
..
..
A1
.
A2
A2
A1
A1
..
A1A1
A1A2
A2A2
1
1
A1-
1:1 .
..
A1- ,
A1-
.
.17.23. ,,
.A1,,
..
.A1.A1
,,A1(,
,,).,.
A1(AfterWilsonandColwell1981.)
.17.24.
.,,
.(,
,
)
.
,
.
,
,
.
,(AfterWerren1980.)
(,,)(,,
).,
,
.
.
(17.25),()
( ) .
(.17.25),,
.17.25
1.0
, .
0.75
0.50
0.25
0.2
0.4
0.6
0.8
1.0
1.2
1.4
1.6
4.6
424
17
()
1.0
1
2
1.0
.17.25. .(),
.
(1)(2)(3).()
.
,,
.
1.
(1.),
(=1.0
0).(3
),,
(AfterThomsonandBrunet1990.)
()
1.0
,.,
,,
,
.
,.
1.0
,,(
) . :
().
(),
(,
,)(Mattonet.al1994).
,
,,
.,
.,
(,,)
40%(,
),
,36
(..12.23).,,
(Clark1993.12).,,
, .
,,
..
.
,,.
(Thornhill1993).
,.
,,
(MHC).
(Pottsetal.1991).
.
.
(Rallsetal.1986).,.
.
:
,,
().,
(),
,
.
(.9).
,,,,
,
(Uyenoyamaetal.1993).,
, ,
, . ,
,,
(LandeandSchemske1985).
(1991).
Eischhorniapaniculata
.
,
.
(.17.26),
.,
:
,(.17.26).
425
.17.26. Eichhorniapaniculata()
()
(O1),
(S1S5)
(O5).()
(O1S5,
O5),(AfterBarrettand
Charlesworth1991photocourtesyofS.C.H.Barret).
()
500
400
300
200
100
O1
()
S1
S2
S3
S4
S5
O5
S1
S2
S3
S4
S5
O5
250
..(1941),,
( ),
,
.
:(),
.
.
3:2,
50% .
,50%
.,,
50%,.
(lloyd1992CharlesworthandCharlesworth1978).
.
.
.:
,
200
150
100
50
O1
426
17
.
(JarneandCharlesworth1993).
, , ,
,.
(Holsinger1991).,
.,
(1989)Ipomopsisaggregata
,
.,
,,
,(
) . ,
,
.
.
, :
.,
,.
(JarneandCharlesworth1993Wyatt1988).,
,
.,
.
1. (,r)
.
.
,.
2. ,(tradeoffs),
,
.
3. (lx)(mx),
.,
.
4. ().
.
5. (),
/.
.
,.
,
.
6.
,.
7. ,.
,
.(
).
8. (,
),
,.
9. ,
10.
11.
12.
13.
14.
15.
,.,,
,
.
,
.
.
,
. ,
,
,
.,
.
:()
()
,
(),
,()
()
, , ()
,,
,.
,1:1
,,1:1,
.
,
,,
.
( )
.,
.
,
50%.
,.,
:,
,
.
(outcrossing)
Tradeoffs()
427
428
17
:J.Roughgardens
Evolutionsrainbow:Diversity,gendr,andsexualityinnatureandpeople(UniversityofCaliforniaPress,Berkeley,
2004):TheevolutionoflifehistoryS.C.Stearns(OxfordUniversityPress,Oxford,1992)
Lifehistoryevolution,byD.A.Roff(SinauerAssociates,Sunderland,MA2002).
:Theevolutionofsex,editedbyR.E.MicholdandB.R.Levin(SinauerAssociates,Sunderland,
MA,1988):Kondrashov(1993),BartonandChalrlesworth(1998),JarneandCharlesworth
(1993),Uenoyamaetal.(1993).
1. ,
.
,
. ,
?
?
2. , , ,
,
().
,?
3.
, .
(see
referencesinStearns1992).?
4.
,.
.?
?(SeeOlivierietal.1995).
5. ,
.
??
,?
6. ,(pseudoarrhenotoky
,).()
,
..
?
7.
.,
,
.
8. ()
().(Lysandra)
24220(Wite1978).
??
18
Ficus
750 .
,(
) .
(.18.1).
.
,
.
Agaonidae
. .
,
,
.,
.
(PhotoGregoryDimijian/Photo
Researchers,Inc.)
430
18
.18.1. .
,
.
,
(PhotoOSF
/photolibrary.com)
, .
.
,
.
. ,
.
,
,.
,
,
.
,
.,
,
.
(,).
, , (
),(,),(,
)(,,
). (
).,
.,,
,.,,
(..12.1918.24).
,,,.,
.
,,
,.,
.,
,(Thompson1999).
, ,
,
.
.
(Futuyma and Slatkin 1983Thomson 1993).
(
).,.
.,
(. 18.2). ,
(.18.2),
.
()
()
()
()
.18.2. .,
.().(),
.()
.,
.
.
(HougenEitzamandRausher
1994).(.18.2)
.,
(1964),,
,
.
.
.
,
.
,(Buchnera).
(..
14.24). .
(.18.3).,Buchnera
.,
. .
.,
,,
.
()
(.18.3Hafneretal.2003).
,(Page2003).
(
),
.,.
, ,
.
,
(Farrell1998).,
.
431
432
18
.18.3.
()
,
()
.
Ruminobacter amylophilus
Proteus vulgaris
,
Escherichia coli
.()Buchneraaphidicola
Schlectendalia chinensis
Melaphis rhois
Pemphigus betae
.
Mindarus victoriae
(
Chaitophorus viminalis
).Buchnera
Diuraphis noxia
Acyrhosiphon pisum
.
Buchnera
Uroleucon sonchi
aphidicola
/
Myzus persicae
.()
Rhopalosiphum padi
.
Rhopalosiphum maidis
(,
Schizaphis graminum
),
(,C.merriami
G.perotensis)(afterMoranand
() ()
Bauman1994afterHafneretal.2003.)
()
48-70 ..
80-160 ..
80-120
..
30-80
..
()
Geomydoecus chapini
Orthogeomys hispidus
Geomydoecus setzeri
Orthogeomys underwoodi
Geomydoecus panamensis
Geomydoecus cherrieri
Geomydoecus costaricensis
Orthogeomys cavator
Orthogeomys cherrieri
Orthogeomys heterodus
Geomydoecus trichopi
Zygogeomys trichopus
Geomydoecus nadleri
Pappogeomys bulleri
Geomydoecus expansus
Cratogeomys castanops
Geomydoecus geomydis
Ctratogeomys merriami
Geomydoecus oklachomensis
Geomys bursarius
Geomydoecus ewingi
Geomys bursarius
Geomydoecus texanus
Geomys breviceps
Geomydoecus actuosi
Geomydoecus perotnsis
Geomydoecus thomomyus
Geomys personatus
Thomomys bottae
Thomomus talpoides
Thomomydoecus minor
Thomomydoecus barbarae
.,
,Hispinae(Labandeira2002).
:
,,
.
,(.18.4).
:
(.18.5..12.5)
(.18.5).
.18.4.
,
.()
(Himantolophus)
.()
(Leucochloridium)
.
,
(DavidShale/naturepl.com,photobyP.Lewis,courtesyofJ.Moore)
()
,(.
. 19.9).
.
,.
()
,
(Abrams 2000):
(DawkinsandKrebs1979),
,
()
,.
,,
.
.
,.,
,,
,10%
.,,
(Coleyetal.1985).
(Pastinacasativa),
,,,
.,,
(BerenbaumandZangerl1988).,
,.
,
.,,(Cucumis
sativus),
(DacostaandJones1971).
()
433
()
.18.5.
.()
(Uroplatus
phantasticus)
.()Dendtrobatus
azureus
.
(NickGarbutt/naturepl.
comBarryMansell/naturepl.com)
434
()
0
(6 )
()
()
20.000
.18.6. ()
()
.
.
(P1H1H1
).
(AfterSeger1992.)
.,
.
(Linumusitatissimum)
(Melampsora lini) .
, (Triticum)
(Mayetioladestructor).
,(R)
. (v) ,
6
R
(18.1).,
Ri,vi
, Ri .
R(Rj),vi
.,
21.000
(.18.6).
(.18.7).
,
,,
(ThrallandBurdon2003).
. (y)
(x)
(Abrams2000).,
(xy)(
)(
).,
()
.,xy
,x(y)
18.1.
R1-R2-
R1-r1r2
r1r1R2-
r1r1r2r2
V1-V2-
V1-v2v2
v1v1V2-
v1v1v2v2
()
18
1.0
0.8
0.6
0.4
0.2
1987
1988
1989 1990
1991 1992
.18.7.
6.,
,,
.
,80%
,(AfterThralland
Burdon2003.)
()
6
4
.18.8. ,
(,)
(,).()
.,
.
.
()
,
(AfterAbramsandMatsuda1997).
435
2
0
-2
-4
-6
x y .y
.
. ,
,
.
.
,
,,
.
(.18.9Vermeij1987).
(Taricha
granulosa)(TTX).
TTX(
25000
25000
),
(BrodieandBrodie1999Brodieetal.2002).
TTX
(.18.10).
(.18.10).
25
50
75
100
125
150
175
125
150
175
()
30
.y
x.y
,x.
.
(.18.8)
25
20
15
10
5
25
50
75
100
()
()
()
.18.9. ,
,
.()Arcinella
()Murex
,
.()Cypraeamauritiana
(PhotosbyD.McIntyre.)
436
18
TTX-
( )
.18.10. TTX
.
,
(Tarichagranulosa)
.
TTX
.
(AfterBrodieandBrodie1999
photoHenkWallays.)
100
75
50
Taricha
Taricha , TTX
Tarcha
25
0.1
10
100
1000
10000
TTX- ( )
. ,
, (Papilio polyxenes)
(Berenbaum1983).
,,,
,.
.(.18.11).
,
,.
(Rotstein
andRobinson1998).(Cuculuscanorus)
,
. (. 18.11). .
,.
()
.18.11. ()(Cuculus
canorus)
(Acrocephalusscirpaceus).()
.
(:,
,,
,
Anthuspratensis,
).
.
,
.
(DavidKjaer/naturepl.com
photobyM.Brooke,courtesyofN.B.Davies.)
()
437
(1998)
.,,
(),
,,
.,
.,,,
,,,
,.,
,,
.,,
,.
(%)
:
,.
, , Wolbachia
(.15).
()
(,
100
).(,
90
)
80
(,
).
70
.,
60
.
50
,.
,
40
,,
30
(Ewald1994Bull1994).
.(FennerandRatcliffe1965),
20
(Oryctolagus cuniculus) ,
1
0
.,
()
(.18.12),
1950-51
100
(.12.18).,
,
0
.,
1952-55
.
.18.12.
,.()
.()
.(1)
(5).
(afterFennerandRatcliffe1965afterMayandAnderson1983.)
1955-58
(%)
. ,
,
. ,
,.
(MayandAnderson1983Bull1994Frank1996).
R0.R0
1959-63
1964-66
1967-69
1970-74
100
1975-81
0
Myxoma
438
18
:
.18.13.
.
Daphnia
magna.
.
(
)()
.()
.
()
,
,
.
,
(AfterEbert1994.)
( )
()
3.0
2.0
1.0
0.0
42 (%)
()
100
75
50
25
0
R0 =
bN
v+d+r
N,,b
,,v
(),d
r
.,
( ).
b,
v,
(,..).,bv.,
.v,,
.
.
.
:()
.,,,
,.
().
,,
.
. ,
,
..
,(Frank1996).
:(
r),
.
(,)
.
,
.(
),
. ,
.b
.,
.
.,
,,
(Bulletal.1991).
.
Daphnamagnas(Pleistophoraintestinalis).
0.0
1.0
2.0
3.0
4.0
.Daphnia,
,
.,
,
,(.18.13).
.,
.
(1998)
0.0
1.0
2.0
3.0
4.0
(Potamopyrgus antipodarum)
(Microphallus) . ,
()
0.2
0.1
0
0.4
0.3
0.2
.01
1992 1993 1994 1995 1996
()
()
1
0.9
0.8
0.7
0.6
0.5
0.4
.18.14.
.()
12
.
.,
,
.()
43
,40
(AfterDybdahlandLively1998.)
0.3
0.2
12
22
19
63
439
(. . 17.21).
,.
,.
Daphnia,
, . , ,
,(
)5
.,
(. 18.14). ,
,
(..18.6).404
.(.18.14)(17
,,,,
).
,.
..,
, , ,
.
Angraecumsesquipidale30,
,,
.24
.(.18.15Nilsson
etal.1985).
,
,
..
..
.
.,(Yucca)
(TageticulaParategeticula).,
(.18.16)..Tageticula
440
18
()
.18.15. .()Angraecum
sesquipedale.()
Xanthopammorganiipraedicta.
,(
GeraldandBuffCorsi/CaliforniaAcademyofSciencesTheNaturalHistoryMuseum,London.)
()
.
.
,
(.18.16).
,,
, .
. ,
(Bull and
Rice1991).,
,
(Herreetal.1999).,
,
,
,
()
.18.16.
.()Tegeticula
.()
.
,Greya,
.Tegeticula
Parategeticula
,Tegeticula
(courtesyofO.Pellmyrafter
PellmyrandLeebensMack1999.)
Tegeticula
yucasella
Tegeticula synthetica
Tegeticula maculata
Yucca
()
Parategeticula
Prodoxus
Mesepiola
Lampronia
Tetragma
Greya
441
..,
Buchnera,(..18.3),
..
.
,
,,(PellmyrandHuth1994).,
,,
.,
,?,
,(5001500
).,
15%,.
.,,,
,,.
,.
,,.
,
.
.
.,.
:..
(..11.2).
,
,(..18.16).
,.
(Pellmyrand
LeebensMac1999).
Bombus
100
appositus
50
0
100
12
12
12
12
12
12
12
kirbiellus
50
0
100
avifrons
50
0
Bobmus-
, , .
,,.
()(
).,
,.
,
(Schluter2000).
,
(.18.17).,,
.,(,)
(,),
,
,
(.18.18).,,(,
)
,,.
100
sylvicola
50
0
100
bifarius
50
0
100
frigidus
50
0
.18.17. (Bombus)
.
().
.,
.B.appositusB.kirbyellus,B.frigidus,B.bifariusB.sylvicola
(AfterPyke1982.)
100
occidentalis
50
0
()
442
18
x, (, )
K(z)
.18.18. .x
(z),,.
,,
.K(z).
()z.
.,
,.
.
(,).
z (, )
()
Melanerpes striatus
()
Melanerpes aurifrons
.18.19. .
Melanerpesstriatus().
.M.aurifrons
()
.
.
.
(After
Selander1966).
,
,.,
.
,
(Slatkin1980TaperandCase1992).
,(.).
,
(. 13 ).
.
,.
,.
,,.
,18.17
,,
.
(1956)
,
,.
,,
,
( ).
(TaperandCase1992Schluter2000).
,GeospizafortisG.fuliginosa
,(..9.27),,
.
.,
(Grant1986).Gasterosteus
aculeatus.
.,
(..16.8).,
.
(SchluterandMcPhail1992).
,
,
.
.,
,
(WernerandSherry1987).
,,
(.18.19Selander1966).
443
,.,
.
(.7).
,
,.
.,,
,.
,
.
,,
(SchluterandRicklefs1993).
, .
,6Anolis
.
,(..6.21).,,
.,,
,(.18.20).,
,,
.
(Accipiter)
.Accipiter
,47
(.18.21Schoener1984).,
()
P C C H H H H H P H C C C
J J
J H H P P P H H C C C C H C C
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TC
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Anolis
,
(C,H,hJ,P,
)()
.
.
(After
Lososetal.1998.)
18
.18.21.
(Accipiater)
,
,
.
.
,
,,
,
(After
Schoener1984.)
,
Accipiter 47 .
(
)
.
0.15
444
0.10
0.05
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1.25
1.35
1.45
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1.65
1.75
1.85
1.95
( )
.
. (1999)
,,
,
.
,,
.
(Benkman1999Benkmanetal,2003)
(PInus
contorta)(Tamiasciurus
hudsonicus)(Loxiacurvirostra)
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.
.,
,
,.
.
B
.
, .
.18.22. .(Pinuscontorta)
.().
,(Loxiacurvirostra)
().()
.(After
Benkmanetal.2003.)
.18.23. .,,(Limentis
arthemis)(Battusphilenor).
.
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,,,
. ,
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,
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.,
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.
445
()
()
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,
,
(MalletandJoron1999JoronandMallet1998Turner1977).,
(.3).
(.18.23).
.
(
,).
().
.
.
.
.
()(.18.24).
. (
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H. etaro
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erato,
,,
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Helioconius,
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446
18
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0.60
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,
.(.18.25)(
,,
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Papilodardanus,
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().
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,
.
J.N.Thompson,Thecoevolutionaryprocess(UniversityofChicagoPress,Chicago1994)
,
.:Thegeographicmosaicof
coevolution(UniversityofChicagoPress,2004).
: Plantanimal interactions: An evolutionary
approach,editedbyC.M.HerreraandO.Pellmyr(BlackwellScience,Oxford,2002).
:ModelsofparasitevirulencebyS.A.Frank(1996,QuarterlyReviewofBiology71:3778).
:Theecologyofadaptiveradiation,
byD.Schluter(OxfordUniversityPress,Oxford,2000)
1. ,
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3.
50100
.,
447
448
18
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.
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Anophelesgambinae).
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80
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1977
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452
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2002). ,
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20405
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Drosophila,
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Giotetal.2003AAAS)
0.9-1.0
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453
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25
454
19
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(SwansonandVacquier2002).
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,
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SwansonandVacquier2002.)
VERL-
VERL-
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455
,
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Colobinae)
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,
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,
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.
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etal.2002),
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/
,46
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28,100
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.
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)
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550.,OR
,
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,.
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(),
.,23,
.
,
,
.,
(e.g.,Stephensetal.2001).(Clarketal.2003)
7645w(dN/ds),
Glu14
Glu21
Asp75
Asn87
Lys126
Lys14
Lys21
Asp75
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Lys126
Lys14
Lys21
Asp75
Asn87
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.19.5.
,
.
,
,14,21,75,
87126.
.,
(Opisthocomus
hoazin).
.,
75
.
(.8.1)(AfterKornegayetal.1994
photophotolibrary.com.)
456
19
.:
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873
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(Pooleetal.1998),
.,,
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,(PalmerandLogsdon1991).,
.,
(.19.6).
.,
1,5%.
95%.,
,,
.,,
10%,
(Shabalinaetal.2001).,,,
,.,22bp()
.(50000)
(Bartel2004).
, ,
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(..8.1).,CD44,,
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(RobertsandSmith2002),
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,
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:
(.17),
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EUCARIA
Plasmodium
Dictostelium
Entamoeba
Naegleria
Trichomonas
Vairimorpha
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.
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.,
(AfterPalmerandLogsdon1991.)
Giardia
ARCHAEA
Thermotoga
,
.
.
Agrobacterium
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BACTERIA
Oscillatoria
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457
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.
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10009().
Mycoplasmagenitalium580
kb468,
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(..14.24),
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458
19
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,(.19.8)
,
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,,
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..
..
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.
,
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0,3% (
)(DeiningerandBatzer2002).
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Alu4050
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DNA
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1.0
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7 9 11 13 15 17 19 21 23 25 27 29 31 33
460
19
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,,EucaryaBacteria.
2(..2.21),,
(LGT).
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(Doolittle1999).
()
.,Entamoebahistolytica
50.
.,
.(.19.10),
Archaea(Fieldetal.2000).
4050(Salzbergetal.2001).
,
.,
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,
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.
.()(~100
),
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,
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461
462
19
N. meningitides
E. coli-2
P. gingivalis
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Synechocystis sp.
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H.sapiens
P. prowazekii
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Entamoeba histolytica- .
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.
,
,
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P. horikoshii
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A. fulgidus
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A. pernix
E. coli -1
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S. cerevisiae
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L. lactis
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.,
,(
)(.19.13).(2003),
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,
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.
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.
463
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.
jingwei,Drosophilateissieri
D.yakuba.jingwei.Drosophila
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,jingwei.,Adh
D.teissieriD.yakubaYmp
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jingwei,YYmp).
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464
19
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(Adh
),Adh
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(Zhangetal.2004).
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(AfterLongetal.2003)
(GPCR),GPCR
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.
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*.
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,(
).
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2003Zhangetal.2003).
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().
(,).
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(.19.16).
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.16).,
.
.
()
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,
,(PrinceandPickett2002).
(Guetal.2002)1739749
.,
(500),
(,
)(.19.17).,,
,.
().,
,(COL),6
(MHC)
.()1,919
(. 19.18).
.C.elegans.MHC
,.
19.19,(
Dujonetal.2004Guetal.2002)800(,
)1000(,).
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465
.19.15.
AFGP.()
(,
AFGP2(E2))
.()
2
,
16(
),AFGP
.()(Dissostichus
mawsoni)AFGP.,
2.
AFGP
,
2(E2),
(AfterChengandChen1999.)
466
19
1
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().,a
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,
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()
150
,
.19.17.
.
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,
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,
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.,
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II
100
I
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50
50
150
250
350
450
550
650
750
850
950
1,050
Amphioxus-
( )
1,150 >1,200
467
.19.18. .
.
,
(AfterKasahara1997.)
6p (MHC)
19p
1q
9q
RXRB
RXRG
RXRA
COL11A2
COL11A1
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RING3
RING3-
LMP2/LMP7
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ABC2
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PBX2
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CYP21
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HSPA1
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GRP78
HlA-A-A,-B,-C
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SPTAN1
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(PBX)
..,(
,(.
,(,
)(.(Tagleetal.1991)
d ( ,
.,,
,d(
,(.19.20).d(
600
500
400
300
200
100
2
3
4
5
9
10
>10
Yarrowia lipolytica
Debaromyces hansenii
Kluyveromyces lactis
Candida glabrata
Saccharomyces cerevisiae
.19.19.
.
,
,,
.
,
(AfterDujonetal.2004)
468
19
,,
1/3,b(.
,1824.
.,5
,d.
()
,
(OtaandNei1994):
1.
,
.
(.19.21).
2.,,
,
.,
,(.19.21).
3.,,
,,
(.19.21)
:,
. ,
.
. (SusumoOhno1970).
,,
,,.
()
.,.
.
(ECP)
(EDN),.
()
.19.20.
.()d(
.
5
,.
()d
b,db.
,,
.
.d
()
(AfterTagleetal.1991.)
14.8
23.0
12.4
0.8
5.0
0.0
18.0
()
1 2
1
2
1
2
()
()
()
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.,EDN,ECPa,
,