evolucia

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daglas
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Tbilisi 2009

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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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)

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33

100

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,
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andFitch1974).

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75

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125

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8,3.
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1977).,(
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34

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35

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36

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200

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Datura
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Atropa
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Lycium
Juanulloa
Solandra
Mandragora
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tabacum
Nicotiana
acuminata
Anthocercis
Petunia
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Ipomoea

37

,


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38
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39

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)
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40

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B.G.
Hall:Phylogenetictreemadeeasy:Ahowtomanualformolecularbiologists(secondedition,SinauerAssociates,
Sunderland,MA,2004).
PAUP*
. J. Felsenstein: Inferring phylogenies (SinauerAssociates,
Sunderland,MA,2004).

W.P. Maddison and
D.R.MaddisonMacClade,version3.0(SinauerAssociates,Sunderland,MA.1992).
,

41

42

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.

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44

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45

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47

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48

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49

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50

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1995)

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U
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51

(
).
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.

,
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1981).
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Balanus(.3.6..4.17).
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,
).
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, ,
.(1997)
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()

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.,

,
.
(
)

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1990)

()

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52

Octomeris
(
8 )

Balanus

Chthamalus

b
a

( ; 6 )

.3.6. ,
ChthamalusBalanus
.

Branchiopoda

,

(. 3.7).

.

,
.,
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Amphignathodon
(Noble 1931). Amphignathodon
,

.
(
)
.

,
.
,
,
.
,

()

H1 H2 H3 T1 T2 T3 T4
Anostraca

Leptostraca

H1 H2 H3 T1 T2 T3 T4

Paranebalia ()
H1 H2 H3 T1 T2 T3 T4

Maxillopoda

Mysida

Copepoda

H1 H2 H3 T1 T2 T3 T4
Mesocyclops

Ubx/abdA

Mysidium ()

.3.7. .
()UbxabdA.
(Paranebalia,Laptostraca).H3
(,MxII),(EnEx)T1T2
.Mysidium(Mysida)T3(En
),T2T1En()
.Mesocyclops()
(AfterAveroffandPatel1997).

53

(.3.8).
, .
.
.19
,.
.
, () . ,
,
(..18.23).
.
,
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(),(
)(..18.24),.

( , 1999).
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()

()

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.()
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Thraupidae(Cyanerpescaruleus).
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vestiariacoccinea,Fringillidae.

.(),
Trochilidae.
(Campylopterushemileucurus).
(),Nectariniidae.
(Nectarinia
pylchella)
(,photofotolincs/AlamiImeges
,photoResourceHawaii/AlamyImages,
AnthonyiMercieca/PhotoResearchersInc.
PhotoResourceHawaii/AlamyImages.)

54

3
. .
.,
,()
.

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(,.,
).
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).

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(Scolopacidae).18
()
166(
).,
,
.

,
,

(AfterHaymanet
al.1986).

25

55

(, , )
,
.,()
4.

.,,
(.3.9.
3.21).

.,,
Hymenpotera
().,
,
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,.
1828,(
Vertebrata) ()
.
. ,
,,,,
(.3.11).

Passioraceae ()

Ranunculaceae-

Bignoniaceae-

Rubiaceae-

.3.10.

(AfterHutchinson1969)

56

3
,
: . ,
.
,,
.19
, ( )
(Gould 1977).
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, ,

. ,
. ,

.

,
.
.

(.20).,,
,(Rensch1959Muller1990Raff1996Wagner1996).

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(AdaptedfromRichardsonetal.1998photo
courtesyofM.Richardson.)

.
, ,
(Raff1996).
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. ,
,,
.
, (Wagner 1996 Muller and Wagner
1996)..,
, (
, , ) .

(.3.12).

()

. (Gould 1977 McKinney and McNamara

1991)
.
,
(Raff1996).

, (
)
( ).
,
,

(. 3.13). ,
. ( paedos
morphos ). ,
,
.
.,


.3.12.
.
.

.
(afterRomer,1966afterVaughan1986)

57

Kuchneosaurus

,

,

(McNamara1997).
.

Prozeuglodon

. ,
,
.


. ,

.

,

(. . 3.4).
Hemirhamphidae(.
3.14).
:
y=bxa
yx,
(x,,
). a
. a = 1,
.yx,
,
a>1().y,
, a<1 ( ) (. 3.15). y x
,
logy=logb+alogx
,x,y,

()

()

()

,
, ,

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()(Ambystoma
tigrinum),
,()
)
.()
(Ambystomamexicanum)

(,photobyTwanLeenders
,photoPainet,Inc.,photobyHenkWallays.)

58
()

3
.3.14.

.(),()Hemirhamphidae()
.

(fromJordanandEvermann,1973).

( )

() ( ) ( )

()

( )

log10y

( 2)

()()(Alberch
etal.1979).

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( +
. 3.16). ,
(Megaceros giganteus)
, .
,
(. 3.17).
(()) (
.3.15. yx
) y (

a
).
.y=bx
(. . 3.13)
.().

(a=1),yx(b)
, Thorius
.(a>1)

(a<1).()
.a
,.(.
.
3.18).
1.
.
.
()

()
()

3
300
. ,
3
y= bxa

(3) b = 0.32
250

a = 3/2
2
.
1
2
200
,
,
4
(2)
b
=
2a
=
1

150
(Raff1996).

1
100

.
(4) b = 1a = 1
,

50

,
(4) b =2.2a = 2/3

0
1
2
0
20
40
60
80
100

( 1)

log10x

()

()

()

y
, x

59

()

, x-
y ,
.

y-
x-

:
=1

(
)

.
(.3.19).,
.
. ,
,
, . (Muller 1990).
,(Ailuropodamelanoleuca)
,(..21.8).

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.
x
y
.x
,y
,.
1
.()

.
(>1).()
(+
)

y.()
()
,
.()y
x,
,

,.

.3.17.

(Megacerosgiganteus).
,.
(FromMillais1897)

60

.3.18.
Thorius
Pseudoeurycea
.Thorius

(AfterHanken1984)

()

(Thorius)

()

,

Thorius-

.
.
,
,
(21).,
,
.,,
, ,

/ .

( ),
( ).
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.

.
, Zygothrica ,
(.
3.21) .

,
, ,

.3.19. Philodendron,,
.Philodendron,
.(Photophotolibrary.com)


()

Eusthenopteron

()

()

Milleretta ( )

.
21.
antedispar

,


.
,
,
(Schluter 2000).
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dimidiata

dissimulata

61

Canis ( )

.3.20.

.
(
Eusthenopteron)
,
,
(
Milleretta).
,
(Canis),

(AfterRomer1966).

.3.21. Zygothrica().
,(
).,
,
(AfterGrimaldi1987).

Zygothrica

prodispar

r
ra
is
ta
s
r
a
pa
ce
nn
an
pa
ula tura
iat
dis
pe
ero
d
i
e
m
i
a
i
t
dis ispar uber
c
c
s pi
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s
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r
p
an
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a
dim
s
d
p
d
p
sx

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sxuberans

62

3
. , ,
,.

.
.
, .
,.Geospiza
. ,
Camarhynchus, , Certhidea ,
Cactospiza(.3.22).

Certhidea
olivacea

Platispiza
crassirostris

Pinaroloxias
inornata

Cactospiza
heliobates

Cactospiza
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).
, ,
. Asteraceae
, .
,
(.3.23).,
(Carlquistetal.2003).

(KornfieldandSmith2000).200,
140 , 500 , .
,,
(. 3.24) .
, , , ,
,,.
, ,
.,
.
.

Pseudotropheus
tropheops

Petrotilapia
tridentiger

Labidochromis
vellicans

Haplochromis
euchilus

Haplochromis
polydon

.3.23.
.()Argyroxiphium
sandwicense,

.
()Wilkesiahobdyiylortiani
.()Dubautiamenziesii
(,photobyElizabethN.Orians,photo
byGeraldD.Carr,photobyNobleProcter/
PhotoResearchersInc.)

.3.24.
.

(AfterFryerandIles1972).

64

1. ()
.
, ,
.
2. .
. ,

(),.
.
3. .
4.
. ,
().
5. .
.
.
.
6. .
,.
7. .
.
8. .
,
() () ,
.
.
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).

1. ,
: 1) GCTGATGAGT 2) ATCAATGAGT 3)
GTTGCAACGT4)GTCAATGACA
2. . ,
, .
, , , 14
,.
,.
3. , ,
( )
?
4. 3.14
?
?
5. (. 3.17)
,,
. ,
.,
.
(1974)
,,
.,
?,
?
6. ,,
. , : )
(,
. 3.10) ) (, . 3.21) )
(,Amphignathodon).
7. ,
.(InternationalHumanGenomeSequencing
Consortium, 2001 Venter et al. 2001)
5.
8. ,
,
.
: )
Amphignathodon )
) ,

65

66

3
) ,
)
().

().
,

,
.

,(
)
.
.

,
.(
).
.
.
,,
.

Australopithecusafarensis
.

(PhotoThe
NaturalHistoryMuseum,London).

68

4

..

().,
(..
4.1)..
,,,
.
.
,
,,
..,
,.

1915,
60,
.
.
, ,
.
,.
.,,
,
,,(.4.1).
510.
().
.

,
.

.4.1.
.

()
.
.

()

69

,
.
, ,

(.6).

,14

(1=1,000)

.
4,6,

3,8

.3,5
,
800(0,8)(.5).
(-)
.

,

1/2

,


,
26,
13 13
,13.
.4.2. .

.
. .
(,U235Pb207),. ,
,
.,U2350,7

,U235Pb207
., .
(.4.2). ,

(AfterEicher,1976).
.
,
(),,
.
.,
,
.
.
.,,
.,,
,.

(4.1)
..
.
.F
.
. , (
),
...

.(542),
,,(251),(,65,5
)(1,8).

70

4.1.

()

0,01

1,8

5,3

23,0

33,9

55,8

65,5

,
,,,,
.

145

(J)

200

(Tr)

251

299

,
,,

(C)

359

, lycopsid, sphenopsid
,
.

(D)

416

,,,,

(S)

444

,,.

(O)

488

()

542

(T)

Homo erectus
Homosapiens

2500

(19001700 )
(1000)
, Cnidaria (), Annelida (
),Arthropoda()

(Q)

3500
()
,

(K)

(P)

(International Commission on Stratigraphy, 2004)

.,
, . , , ,
(Jablonskietal.1986).,
.
.
:1)
,,
,2)
,3)

,.,
4)
,.,
,
..
2500001%.
,,,1)
2)
3)
4)
,,.
:1)
.
2),.
,
,,
,.

,
.
.(Levinton,2001).,

(.4.3Sheldon1987).
(1985) Gasterosteus doryssus samekala
,111000
.5000.
(.4.4).
.
(),
(.4.4).5000
,.

.
,,
,2.,
,
.
.
,
.,

71

72

Cnemidopyge

Nileids

Ogygiocarella

Platycalymene

Nobillasaphus

Bergamia

Whittardolithus

Ogyginus

0
0

.4.3.
.,
(After
Sheldon1987)

9 11

12

13

9 10 11 12 131415 16

. ,
,
,.,..,
....,
.

.,
.
(.4.5).

()
. Sarcopterygii
,408.
,(
),,(.
4.6).,
.(.
4.6).

10.5

10.0
9.5
9.0
8.5

.4.4. (Gasterosteusdoryssus)
.,
(),
(),
.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
).(
)
,(4.6
4.6)..
,,
(Clack,2000).,
.,
.13
(Clack,2002).(,
).
(
)(
,).
. ,
.,
Aves,.,
,
(ChiappeandDyke,2002Xuetal.2003).
,
.Archaeopteryx(.4.7).,
,,
(. 4.8 ,). (theropod)
(. 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
,:,,
..
,Sinosauropteryx
(),
.,
( ,

75

). ()
,
(.11).
.

(Kemp1982SidorandHopson1998).
,
,,
.
,.
(),
(
),. ()
(),

()

Archeopteryx

()

.4.7. .()Microraptorgui
(140
).,

.,
.()Archaeopteryxlithographica
.
.
.(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).

(,).
.,,
,.
,
(),
.
,,
.
( ) ,
(.4.10).
(.4.10).
Therapsida(.4.10)
,,
.
,.


.
,(.4.10,).
,
,
,
.
.

(.4.10)
,
. ,
, .

.
/ ,

.
.

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.
, (, ,
, , ,
), . . ,
.,
.
.
.
.
,.
(130
),.
.
.
.
,(Gatesyetal.1999).
, . ,
,,.
,
.Y
,,,
,,
(),
.
().()()
,.
,...
,5035
(.4.11)(Gingerichetal.2001ThewissenandBajpai2001Thewissen
andWilliams2002).Pakicetus(5348)()
.
Ambulocetus (4847 ) ,
.,,
.
.Ambulocetus.
,.(,
Rodhocetus,4939).,
.

,
,.
Dorudon(35).
,,
,.
.,
.
.

,56
(.2)..
.67(


()

Elomeryx

Elomeryx
.

.

Ambulocetus
.

.

()

Ambulocetus

Rodhocetus .

.

()

Rodhocetus

Dorudon
.

()

Durodon

()

Ohocoena

,()).
,,,
(. 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)].

()

()

Australopithecus afarensis (3,5 .)

81

A.afarensis-

()

()

Australopithecus africanus (3,0-2,5 .)

Homo habilis (1,9-1,5 .)

...

...
...

()

Homo erectus (1,6-0,2 .)

...
(
)...

()

Homo sapiens neanderthalensis (120 -30 .)

...

82

4
,,
40000 .
6 , , sapiens sapiens (
),,
.12000,,

.
40 .
.,
.
,35,
.
11000.
.,,
,
,,
.

( ). , .
Afarensis Africanus, erectus sapiens
,,,,.

.,
,.

(Lovejoy1981Fedigan1986).,
,,
.
,
.,

..
,.
,,
.
. , ,
,,,
,
,,,
.

,
,.,
,
.,
.,
.
,.
,,
.
,
.
,,
,,
.

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.
,,
,
,.
,.
,,
,,
,,
.
(.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 .

.4.22. ,
(s),
68%( x )s
,.95%2s
,99,7%3s
.()(
).
0,25s,
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,.
,,


..
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).
?
?
4. ,
.,
,,
..
5. ,
.?
6. ,
..

.,
? () 65
60(.4.4),()7025
()MorganucodonProcynosuchus(.4.10).
,,
?
7. ,
?
?

3500000000
,

.

.

,

.
(
,
),
,
.
5
,,()
.
,
.

.
,500
.

(..
97).

Anomalocaris
Pikaia.
Hallucigenia
Marrella(ArtJohnSibbik/
NHMPL).

92

5
5,,

.()
,.
,
:
1.

2.,

3.(..
)
4.

5.

6.,
7.,

8. ,

9.
10..

, 14
( ) .
,
.10,
.
()
,.
,4,6
.,(.
.4.1)3,8.
,
.
,
.,4,5
.
4,
.

,,
. , , ,
,
.
,,
,,,
,.,
, , ,
,
().,
.
,

93

,,
.,
L,.L
D,
.

.
,
(),
.
,(Orgel1994
MaynardSmithandSzathmary1995Rasmussenetal.2004).

.,
(CH4),(NH3),(H2)(H2O)
(HCN)(H2CO)
,,,
(.5.1).
.(
,),
.,,
().,
.(),
,
.
,,,
.,,
.,
, ,
.
2

.
(1970),(,Qb
)()
H2, CH3 NH3-

3


.
(),

,
.
,

1
() .,,


H2O

.
,
,
.,
.
,
.,
.,,

,(450,150

).

.5.1.

H 2O
N2
N H3

CH4
H2 CO2
`

94

1 R1


.

2 -

R1
R2 ,

.

AA

R1

R1

AA

AA

.5.2.
.,
()
,(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
,
,
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102

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115

S.M.Stanley,Earthandlifethroughtime,secondedition(W.H.Freeman,NewYork,1993)
.W.I.AusichandN.G.Lane,Lifeofthepast(PrenticeHall,UpperSaddleRiver,N.J.,
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116

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A.K.Behrensmeyeretal.(eds.),Terrestrialecosystemsthroughtime:Evolutionarypaleoecologyofterrestrialplantsandanimals(Universityof
ChicagoPress,Chicago,1992)
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E. N. K. Clarkson, Invertebrate
paleontologyandevolution(ChapmanandHall,London1993)P.KenrickandP.R.Crane,Theoriginandearlydiversificationoflandplants
(SmithonianInstitutionPress,Wachington,1997R.L.Carooll,Vertebratepaleontologyandevolution,(W.H.Freeman,NewYork,1988)D.
B.Weishampel,P.Dodson,andH.Osmolska,TheDinosauria(UniversityofCaliforniaPress,Berkieley,1990).

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124

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128

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129

130

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.C.B.CoxandP.D.Moore:Biogeography:Anecologicalandevolutionary
approach(BlackwellScientificPublications,Oxford,1993).
R.E.RicklefsandD.Schluter:SpeciesdiversityinecologicalcommunitiesHistoricalandgeographicalperspectives(UniversityofChicagoPress,
Chicago,1993)..
.
J.C.Avise:Phylogeography(HarvardUniversityPress,Cambridge,Mass,2000).
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156

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158

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20046,4
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159

160

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, ,
:P.W.Signor(Thegeologicalhistoryofdiversity,AnnualReviewof
EcologyandSystematics21:509539,1990)M.J.Benton(Diversificationandexstintioninthehistoryoflife,
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
sonsThediversityoflife(W.W.W.Norton,NewYork,1999).
:Principlesofconservqtionbiology,edditedbyG.K.MeffeandC.R.Carroll(SinauerAssociates,Sunderland,
M.A.,1997).

1.
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173

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178

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179

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181

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185

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W.H.Li,MolecularEvolution(SinauerAssociates,Sunderland,MA,1997)
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190

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= 0.48
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.,
(,,
,).
(..9.4),
,
(0,36,0,480,16,
0,4, 0,4 0,2 ).
.MN
p=0,763q=0,237.
(320),
.
:

MM
p2
0,582

MN
2pq
0,362

NN
q2
0,056

(
)

186

116

18

187

114

19

195

9A. -
,A1A1,A1A2A2A2
D,HR,A1A2p
q..

.
,.
:

A1A2

A2A2

A1A1xA1A1

A1A1xA1A2

2DH

DH

A1A1xA2A2

2DR

A1A2xA1A2

H2

A1A2xA2A2

2HR

A2A2xA2A2

A1A1

, ,
.
A1p,A1A1
p2.
:

A1 (p)

A2 (p)
A1A2 (pq)

A2 (q)

A1A2 (pq)

A2 A2 (q2)

DH
2DR

H2/4

A1 (p)
A1A1 (p2)

H2/2

H2/4

HR

HR

R2

*,93
3.2
.

,p=D+H/2q=H/2+R.
:
A1A1:D2+DH+H2/4=(D+H/2)2=p2
A1A2:DH+2DR+H2/2+HR=2[(D+H/2)(H/2+R)]=2pq
A2A2:H2/4+HR+R2=(H/2+R)2=q2
,,

,.k
(A1,A2,...Ak),
Aipi2,
AiAj
2pipj,pipj.
(H)
2
H=1 p i .
,p1,p2p3A1,A2A3,
(A1A1,A2A2,A3A3)
p12, p22 p32, (A1A2,A1A3,A2A3)
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

Pm/+ CyL/+ , 1/4

+2 +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)
:

x = (f i X i ) = (0,16)(9) + (0,48)(10) + (0,36)(11) = 10,2

i =1

.
,
() .

.
?
( ) ,
.
,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=

16(9 10,2) 2 + 48(10 10,2) 2 + 36(11 10,2) 2

= 0,485
99

,
,
.,
(.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.
.
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217

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219

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,
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220

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221

222

6.
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()
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.

()

()

()
,
(
)

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.
.:TheodosiusDobzhanskysGeneticsofevolutionaryprocess
(ColumbiaUniversityPressNewYork,1970),Geneticsandtheoriginofspecies.
.ErnstMayrsAnimalspecies
andevolution(HarvardUniversityPress,Cambridge,MA1963).
.

:RichardC.LewontinsThegeneticbasisofevolutionarychange(ColumbiaUniversityPress,NewYork,
1974),
.
:PronciplesofpopulationgeneticsbyD.L.HartlandA.
G.Clark(ThirdEdition,SinauerAssociates,Sunderland,MA,1997),
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evolutionbyJ.C.Avise(SecondEdition,SinauerAssociates,Sunderland,MA,2004).,
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.

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(,A2A30,09).,
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287
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A1A1,86A1A228A2A2.
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.,
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0,23).A1A1A2A2

223

224

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44A2A2B1B2,48A1A1B2B2,97A1A2B2B2,49A2A2B2B2.()A1A2(pA,qA)B1B2
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.()
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226

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227

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 - :

229

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230

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.10.3. ()9
(2N=18)()50
(2N=100)

.
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 - :
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4. , .
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5.,Ne
.Ne.,
100,150,25,150125,Ne
70(*),110.

231

232

10

.10.5.
(Miroungaangustirostris)
,

(PhotoRichardHansen/Photo
Researchers).

.,
,(,,
,).
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,()
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,
.,
(.10.6).

.(1956)Drosophilamelanogaster
. 107 . 8
8.(bwbw75)
.,
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bw75,28.
.
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,
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),
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 - :

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15

()

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10

N0 = 10

r=1.0,

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r=1.0 ,

233

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.

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10 2

10

10

10

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(p=1)

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30

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20

10

32
0

20
16

.10.7.
Drosophilamelanogaster107
.
16bw75/bw
16(8
8).bw75

,
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,
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032
.
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,
bw75 , -6
.

10

15

19 0

24

75

19 (bw75
) 0- 1- (32 )

(0),
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234

10
.

, , .
,(1970)
(Musmusculus).,
,,
.
,
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,(10.1).
,
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(Miroungaangustirostris..10.5).
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1974),,
(. 9 ). 30000 , 1890
20.
,
20%.
.
,,
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,
.
,9
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,(Linepithemahumile).
.
,.,
.
.,
.
,
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.
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,(Tsutsuietal
2000.10.8).

,
,.,
.
30
10.1.

60

.


Es-3b
Hbb
Es-3b
Hbb
,

( 10)
29
0,418
0,849
0,0506
0,1883

( 200
13
0,372
0,843
0,0125
0,0083
.
Source: After Selander 1970
,
.

 - :
()

.10.8. (Linepithemahumile)
.()

.(),
,
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2
...

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2.2

2.6

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()

100
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,
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80

60
40

20
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235

22

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2.6

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.,
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.(KingandJukes1996)
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236

10
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,,
:
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..,4Ne
.
.
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 - :
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t

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.10.9. .
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(n).()
,

.
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.
,
(AfterCrowandKimura
1970).

237

()
t
,,
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.
t

u0(
),
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.
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.
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(.10.10..2.15).
10.10,
.
510,.
40.



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(10.10
0,01,108).

(Li1997).
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(..10.8).
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,
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( t ) .

238

10
10-15
,
.

,
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0.4
0.3

4N e u 0
4N e u 0 1

(.10.11).,
106(Voelkeretal.1980),
Ne=1000,0,004
.Ne250000,0,5
.

0.2

,

0.1
/
u0.

20
0
60
80
40
,
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,

,
.10.10.

(1991)Adh()

(..9.14)
6120

Drosophila
.
(AfterBrownetal.1979).
()(
),,().
uR,
uS,,
uR:uS,
. ( 10.2) ,
5%(29%),
.,
,
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,,
.10.11.
,
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,
(u0)
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(AfterHartlandClark1989.)

?
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0.8
0.6
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10

,
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,()
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 - :

239

10.2.
(,
Drosophila- ),
(10.12).:,
a
,

(10.3).,

2
7
.

42
17
.
()

4,5
29,2
,(C)
Source: Data from McDonald and Kreitman 1991
,,
a
D. melanogaster, D. simulans D. yakuba
.
C6,
,(Kimura1983).,
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.
(
):,
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(,).
(. 10.13).
,.,
(.10.13).,,
,(uT)
.,

,.
.,

,
10.3. (Evaize et
. al. 1992).
.

FST
(. 9 ). ,


Ne(N)
.

(
) .
m
.
,
(FST)
:

135

0,000,00

4,520,87

102

0,000,00

3,940,81

S17

134

0,060,04

2,690,53

376

0,010,04

2,920,34

51

0,200,10

3,031,02

31

1,070,37

4,782,14

(-

141

0,560,11

4,380,77

(-

146

0,780,14

2,580,49

106

2,030,30

5,561,18

136

3,060,37

5,501,45

-3-

332

0,200,04

2,300,30

- A

331

0,190,04

4,060,49

Source: from Li 1997

a
, 109 .
80 (8107). , , .

.10.12.
.
.
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.,

(..8.2).(AfterLi1997).

10

109

240

FST =

Nm
. m=1/(N) (
),
Nm=1FST =0,2.,
.
:

Nm =

.

.

1
4 Nm + 1

()

,FST
. 9
,,,
,
(Slatkin1985).

()

()

(1 / FST ) 1
4

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17.6
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70.3

100
.

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,100
.()(28),
(12,3+17,6=29,9)(12,3+17,5=29,8)
.()(8,9)
,
(1,2+5,7=6,9).(5,7),(7,5).
.,
(AfterEastealandCollet1994).

 - :
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,
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FST).
,
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,.,
.
,()
.
.
(Thomomysbottae).
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.,
.
50(
)82521
(.10.14).50FST
0,412(Nm=0,36),0,198(Nm=1,01).
,
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(PattonandYang1977).

10,,
,
.
().,
(Ne)
(Hudson1990).,,Ne
,
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.
(Ne),
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(tCA)
2Ne,tCA=4Ne.
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,
tCA=Ne)
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u,
ut.,
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t=2Ne,=4Neu.,
,(10.15
).
(u)

,
(),
,:
Ne=/4u

241

242

10

.10.14.
Thomomysbottae
.
4Adh,
3Ldh1
.
,
(AfterPattonandYang1977.)

Adh

Ldh

4- Adh
,
3-
Ldh-1 .

50 100

200

(Ne=/u)

Homo sapiens-
,
.
,
(..6.17).,
,.6,

.,
Homosapiens
.
:
,,,
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Horaietal.1995Ingmanetal.2000).Y(
)
(Hammer1995Goldsteinetal.1995).,,
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Y
,.

.,
40000 143000 .

,

 - :
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()

N=6

6
.

()

N=12

24

12

()

Homosapiens
.
,Homosapiens100000
(.10.16).
()
(see
Hammer1995Rogers1995).,
460011200!
.
,
.,
,
(Rogers
andHarpendling1992)..

1. (
).
.
,(
).
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3. ,.
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,

243

.10.15. ()
()()
.

,
.

N
2N.
.
.
,
()
.

244

10

.10.16.
HomoerectusY1,8
()
H.neanderthalensisH.
sapiens.
,H.sapiens

100000
().

(12000)
,

Homo erectus 1,8

.

Homo
sapiens 100000
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1/(2N).
4. ,
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,.
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).
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().
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10000
,150000
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 - :

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(=
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FundamentalsofmolecularevolucionbyD.GraurandW.H.Li(SinauerAssociates,Sunderland,MA,2000).
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257

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260

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262

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. : Natural selection: Domains, levels, and Challenges


(OxfordUniversityPress,NewYork1992)
R.Dawkins,Theselfishgene(OxfordUniversityPress,OxfordandSanFrancisco,1989)Theblindwatchmaker(Notron,
NewYork,1986).
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276

12

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100
80
60
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0
1975 1976

1977 1978

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.
,1976
,

(AfterBishop1981.)

278

12

600

(Roushand
McKenzie1987).
500

(.).,

400

,
300

200
(RoushandMcKenzie1987).
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100
.R,,
,
0
1940
1950
1960
1970
1980
1990 ..23
,,
.
.12.9.
510%
.
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.
.
,.
:
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(After
MetcalfandLuckmann1994.)

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.
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()
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279

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.

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280

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.
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(

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,
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.
,

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.
, ,
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1971).(S),
A
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,
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,(AS),
(Plasmodiumfalciparum).
,.,
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Y,,(q=0,13)(
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,,AA
.,,S
0,05.
,
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.
.
.,

282

12
(CleggandWeatherall1999).G6PD
.

150

100

50

10
0.2

0.1

0.0

10

.,
.,(Curtsingeretal.1994).
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A2A20,4.
,0,60,9.
,A1A2,
.A1A1A1A2
(0,5)(0,6)=0,3,A2A2(0,4)(0,9)=0,36.
,A2A2A2.

(),
().
, , ,
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).,
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.

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.
12.8
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.
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photographscourtesyofThomasB.Smith.)

283

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.

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,,(
q2=0,5).
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.

.,

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,S1S2S1S3
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W11
W22
).
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W12
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0.6
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. ,
0.4

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.
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(1993)

0
0.2
0.4
0.6
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. () ()
() (. 12.16).
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.()
.
,(pA1).
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W22=1sq2.s=1.
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,
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p=0.

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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.
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(1984),.
,
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(,
).
.
,
.
.
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285

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).
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, 0,25 (1 3 ),
.,
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3).,
4,12.
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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).

,,
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4

S=0.75

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286

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(
).
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0,5
(AfterBasolo1994.)

, . ,,
(..12.15).
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..
.
.
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.
(1989),Heliconiuserato
()
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, ,
.,
.
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.
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,
,.

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.
.0,52
.,,
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.

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.
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(A1A2) . A1A1 ,
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,A1A1A2A2
.,

Heliconius melpomene

Heliconius erato

.12.19. .
Heliconiuserato
.
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UniversityInsectCollection,courtesyofAndrewBrower.)


.,
,
(..8.218.22).

,
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.,,
.

( w )
(p), w p(.
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:
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, w .
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).
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.


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287

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,

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12.20).
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,
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.
.
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289

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290

12
,.
.
,(.12.21),
(. 12.21).
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.
,
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).,

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).
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(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.
,
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.
1490

,,
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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.)

()

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, .
, Nicotiana-
.

()
H-A3
H-A11
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A
C-A126
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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
.,
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.
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().
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.,
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.
,
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()

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10 11 12

()

13
4.28kb

6.79kb 7.54kb

Med A
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60

40

20

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.A45
,
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()

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,Med
.
149
.
(After
Tishkoffetal.2001.)

Med

21
AC
/

41

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61
TT

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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).,,
,.,
.,.

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,
,
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.()

.
(,,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)
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,
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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)
?
?,,,
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70.(R).
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295

296

12
4. ,,
,,
,
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.
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.
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.(2002),(2003)
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298

13

15.5

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(
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,
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Cydia-
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Chilo-
).,
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.
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(Endler 1986Taylor et al. 1991). ,

.
,
.
13.5
(
32
34
36
38
40
42
44
46

)(
(N)
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.

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1750.
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12.
250 .
,

.
(After Tauber et al. 1986).
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(Sylviaatricapilla)
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.,.,
(
).
(1992),
()
,
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.,
,
,,
.
,.
.
700100,
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. .

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.13.3. (Jaderahaematoloma).
.
..

.



()..
(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%,
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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

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,
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VA=2pqa2
6

10

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.
,
.
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).
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,),.
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

,
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X1
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(,
L=6
L=4

X2
X1
Y1 Z1
),

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X2
X2
Y2 Z2
. QTL
(QTLQuantitativeTraitLocus

).QTL
X1
Y1 Z1
1 2 ,,
(1)
.
X2
Y2 Z2

(
X2
Y1 Z1
) 2
.,
.
(2)

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,
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Y2 Z2
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().L
,.
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,
.

(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

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X2

Y2 Z2

301

4.4

4.6

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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. ,
.
()
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()
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.
,
QTL.
QTL.
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QTL.QTL
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,
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.

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.
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303

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13
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R=0,5,


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R =0.001

0.9
0.8

.

.

0.7
R =0.01

0.6

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(R)

(D)
(D0).R=0,5
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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).
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,
(,,
),.,,
,
(..13.2),.
.
. ,
,
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(Macnair1981Bradshaw1991).
,
.

z(,
),z(
,
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,(.13.8).
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(13.8)
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305

R=h2NS
h2N=R/S.S(
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F1-
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()

10

12

14

16

18

10

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12

14

16

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12
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P

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z>12.

( P )
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(R).
z
,

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.
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(R).
()
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,
z>14.
,S
(R)
.

306

13
,
(Orr1998).
,
. ,
.Vm
103VE.
.,
VA/(VA+VE)=h2=0,51000
.(
,(HansenandHoule2004.)

()
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(.13.9).
.
.
,,
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.
.
.
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(HillandCaballero1992)....Drosophila
melanogaster
.86250
.50.
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().86
3545(.13.10).,316%,12
19,,
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.
.
.
(Mackayetal.1994).
:.
,86
(

),,
.
,
.
.
()
.,,
,
.,
(KearseyandBarnes1970
Mackeyetal.1992).
,
(e.g.,WeberandDiggins1990LopezandLopezFanjul1993).
.
.,


()

307

.13.9.
.

.()

.()
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,
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32.
,
(
)12
.

(1868).

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()

(12,

).

60

50

40

30

20

10
10

20

30

40

50

60

70

80

90

100

110

120

.13.10. Drosophilamelanogaster

.
86
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(AfterYoo1980.)

308

13


.
.
,( z )(V)
( z b ,Vb)( z a ,Va)(,
,
).,:

za zb
VP

VP.
,
:

Va Vb
Vb

(Endler1986).,,,
.
:(z)
(w)b*.
, .
(z1)(z2).(,z2)
,(
):

w=a+b1z1+b2z2
(LandeandArnold1983).,b1b2(
),
.
, , ,
.

. (1986,1989)
.
, ( )
,.
,.1977.
.(Geospizafortis)
.
,
(.13.11).( z a )(
z b )ib
:

i
0,28
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b
0,23
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0,43

i,0,20,3,
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.
,
.,
,
,,,
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(Travis1989).(1992),
( (Eurosa solidaginis) )
.
().
,
, .
EurytomagiganteaE.obtusiventris
.
.
.
,E.gigantea
(
),.Eurytoma
obtusiventris,

.,
(j=0,3), ,
,
(i=0,34)(.13.13).

G. fortis

1.0
0.8

,
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,
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0.6

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J S N J MM J S N J MM J S N J MM J S N J
1975
1976
1977
1978
.13.11.
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.
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.13.12.
.
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.(AfterCavalliSforzaandBodmer1971.)

100

20

.
,.,

,
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309

-70
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15

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10 11

310

13
.13.13. ()(Erostasolidaginis).()Esolidaginis
.
,
.
(ScottCamazine/PhotoResearchers,Inc.afterWeisetal.
1992).

()

()

500

...

...

400

300
200
100

10

15

20

()

25

30

.,
,
(Fongetal.1995).
Gammarus minus ,
,
. .
12,
.(1989)
,
.2
.,
(b) 0,3
.
,
,
,
.

.
(Kingsoleretal.2001).,
,(b0,25)(.13.14).
(.13.14).

,.

,
( )
().Vm
,
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1988).
,,
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,.,
.,
, .

,
,(.10).,
,
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.10
).,.


0.4

, ,
.
(1990) (
)
,
(
,),

.,
(Vm)
.,
,

.Homosapiens
,
.
,
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, ,
,(.
.4.21).,
,
.

()

0.3

0.2

,
.

0.1

0.25

0.5

0.75

()

0.6

.13.14.

(),

.()
(
)(
).

(AfterKingsolveretal.2001.)

311

0.4
...
.

...

0.2

-1

-0.5

(BartonandTurelli1989).
,(,
,,)h2N,
(Houle1989).,
,.,
.,

,Drosophila.,,
,
,(Dobzhansky1956).
.
.
,
.

0.5

()

312

13
.,,
,
,
.
(Goldstein and Holsinger 1992). ,

.,
(Brger1989).
,,
(Vm)
(Lande1976Houleetal.1996).,
,Vm,
,(Turelli1984).,,
(,),,
(Laietal.1994).,
,
, Vm
(VA).,VA
,,
(Houleetal.1996).

.,
:,
.:
.

,.
,.
(1992)
Thamnophisordinoides(.13.15).
,.,
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,
.,,
.
.
.
126646
.(
)..
,,
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13.15).,,.
,.
.

: ,
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(r),+1(,

313

)1(
...
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rp,,,

...
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.
.
.
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,
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3.3

,rG.
2.2
.
,
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.
0.1
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-1.1

,
1.2

-1.1
3.5

().,

(+,+,),
(+,,+),.

.
.13.15. Thamnophisordinoides
(Turelli1988).,,

Primula(..9.16),
.
,.

,,
.
,,,
()(After
,,.
Brodie1992.)
.
,.
, (Lucilia cuprina) ,
R,
.(
),(
).R.
.R
,R,
.,
,R
(McKenzieandClarke1988).

N
,.,
(1993),,
,
,.,
(rG=0,17):
,,,
.
.,
,.
,.
.
(1992)

314

13

()

1.0
0.9

0.8

(9 )

0.7
0.6
0.5
0.4
0.3
0.2
0.1

()

1.0
0.9
0.8
0.7

.13.16. ()()
.

,,
9,
.,
.
(PetLen,PetWid),(TubeLen),
(LongSt,ShortSt)
(Pistil).
(AfterConner2002).

,

(TubeLen)
(ShortSt)
(LongSt)
.
.

0.6
0.5
0.4
0.3
0.2
0.1

.(Raphanusraphanistrum)
, , 9
.
.
(.13.16).,
,,9
.

.
,.
.
z1z2,z1
,(
z1),z2.z2z1
,z1z2.
.,
(1981)
(z1) (z2) 0,89
(.13.17).
,
(z1),
(z2),
.
(z2),
()(z1)(.13.17)(
,
).,,
.
.
(.13.17).
(,13.17),
(z1,).
z1.

.
.
(,Geospizafortis
,).
,.,()
,
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()
()

()

,

,
,

.

()

,

,


.

,


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,
,(.17).
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.
,
(Wagner1998).,,
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,(,,).
,
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.
(,8).

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,
, ,

315

.13.17.

.()

.
.()
(rG)(z1)
(z2).

.
.
(,),

,
.
,

.
(*),
.

,
(*).

(PhotocourtesyofS.J.Arnold.)

316

13
(Stebbins1974Schluter1996).
.,
(
),,
,,
.,
,
(Futuymaetal.1995).
,
,
.(.13.18).(2000)(gmax)
,
,
.gmax,
.Gasterosteusaculeatus
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gmax
(. 13.18). ,
,
.
,
.,
4.
,
.
.,

,
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(Steppanetal.2002).

.
(.13.19.
.9.18).
.

,
(GE)(VGxE)(.
13.19,).(.13.19),GE
(VGxE=0).

:
(WestEberhard 2003).
,
(.13.20).,
:,(.
13.20).
,
(SchlichtingandPigliucci1998).


()

X3

X2

gmax-
.

2


.

X1
1

X0

()

1.6

gmax

1.4
1.2
1.0
0.8
0.6
0.4
0.2
0.0 0.05 0.10 0.15 0.20 0.25 0.30

(
)

( )
.
(.13.19).,
,,
.
,(Scharloo1991).
,,
.
(1953)
.,Drosophila
,.
,.
.
.
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.

317

.13.18.

.()

.
(gmax)

.
x 0
(*),
,

x 1
.()
gmax1.

, x 0 x 2
gmax
,(2).
(3),

.()

(,)
.
gmax.
(A
)gmax,

9.
9

(
)gmax,gmax

.
,
gmax
.
(AfterSchluter1996.)

 13

Z1

Z2

()

E1

Z1

()
Z2

E1

E2

Z1

Z2

. E1 E2
.

( ).

()

E1

. GE
.

E2

.13.19.
.,E1E2
.z1z2.

.().GE
.
,.()
,.GE
.E1E2
().
,
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.
().

318

E2

()


, .

( )

.,
.
,
.
,

.,
,
,
(Scharloo1991).

()

.13.20. .
()NemoriaArizonaria(
)()
..
()
.()Ranunculus
aquatilis,
(,photos
courtesyofErickGreenfromCook1968.)



,
(Wagneretal.1997).,(,
),()
.
,
.,,
.
,?
,,
.
.
,,
.
.,
.
,
.
,(
).,Drosophilamelanogaster
( ) (4) .
,scute(sc),
(.13.21).
, , scut .
.
scute,()
(1966)
.(
2).30
(.13.21).,
.
?(1997)
(2000).,
,
.
.
,,
,
. , ,
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,
.,
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.




,
,.,
,89(.4).
,
.,,
.
.21
.
(Olson and Miller 1958),
(Pigliucci and Preston 2004) ,
,,
.(1996)

319

320

13

.13.21.
.()Drosophila
melanogaster
.

().()

scute

.
1,31,81121
(BasedondatafromRendeletal.1966.)

Drosophila

()

scute

,
scute ,
.

()

31

81

121

1.00

0.50

0.00

0 1 2 3 4

0 1 2 3 4

0 1 2 3 4

0 1 2 3 4

,
,
(13.18
).,,
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,
.
,

(Armbrusteretal.2000,2004).
,,
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,(Stylidium),
,
(.13.22).
(0,81).
,.
,
.,
(Armbrusteretal.1999Herreraetal.2002).
.

1.
()(VP)(VG)
(VE).
(VA)
.
().,


VA.
2. V
A/VP(h2N).,

.()
.
3. ,
.
,
.
4. (QTL)
.
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.
5. .
.
6. ,
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.
7. . ,
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,
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9. ,
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, .
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(
),,
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),,
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().
10.
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11.




,
.,
,
.
.
.

321

.13.22. Stylidiumbicolor
.
.

,
.,
(
)

.
(0,81)(From
Armbrusteretal.2004.)

322

13

(QTL)

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)
.,
:C.D.Schlichtingand
M.PigliuccisPhenotypicevolution:Areactionnormperspective(SinauerAssociates,Sunderland,MA,1998).

1.
,,(R)
,(VP),
(VA),(VE)
(S):()VP=22,VA=1,252,VE=0,752,S=1,33
()VP=22,VA=0,952,VE=1,052,S=1,33()VP=22,VA=1,252,VE=0,752,S=2,67(
:0,83).
10,
?
2.
,
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,,
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()(Houleetal.1996).
,
?
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,.(Lande
1980)..
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5. ,
(
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.
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323

14

.,


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,
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. (Polistesgallica)
,

(PhotoBartomeuBorrel/AGEFotostock)

326

14
.
,
.,
.,
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.,,
.
11,,(
),.
,

.,
().
.

.14.1.

,
()
()
.

:
.
: .
:
.

,
.,
,,,,
.,
,.(1999),
,

(.14.1).
,
,.
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.
,

.
.,

..,
.
.,

(, ,
, ,.
).
()
(
).
. .

(, ,
, .
).

(Hamilton1964Michod,1982).,
,

() .
(,

,,
).


..
,(b),
(r)(c)

rb>c
r(),
(Grafen,1991)(.9).
,
,r=0,5.,r=0,5,
,0,25,
.

. A
,,
,A,
.c=1()b=1()
Ar=0,5.,
,,.
() .
,r=0,25,

,,,.
,
,.

. A
,,
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,
.

(.12).AA
..AA,
(),A,
.A
,A.
,
.

.(ESS),
(1982),:
,().
,,
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,.ESS
,()
.
.
,.
ESS,
.

327

328

14
.14.2. (Oryxbeisa)
,.()()
.()
,
()
.
(AfterWalter1984).

()

(.14.2)
.

.
, .
,
,
,
(A).,
,,
.
?
,
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(V),(C).C>V,
ESS,,,
,P=V/C.,,
,,
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,,

()

14A. ESS
, (H)
,
( C )

( V ).
(D) H

.
,


=V/2.

1/2

V/2C/2.
():

1/2(VC)

V/2

x y
E(x,y),
IJ
(w):
w(I)=w0+(1p)E(I,I)+pE(I,J)
w(J)=w0+(1p)+pE(J,J)
p1pJI
,w0
.JIESS,
w(I)>w(J).
,,
E(I,I)>E(J,I)E(I,I)=E(J,I)E(I,J)>E(J,J).
(D)ESS,I=D,J=H,
E(D,D)<E(H,D)(V/2<V),D
ESS.(H)ESS,V>C,
E(H,H)=(VC)/2E(D,H)=0
,V>C.V<C,H
D(ESS)
.
,V<CF,
H P,
D 1P.

P ,
,

.
, H D
P1P
P(VC)/2+(1P)V P(0)+(1P)(V/2).
P,P=V/C
. ,
P
(MaynardSmith,1982).


()

329

()
.14.3. ()(Ucasp.)

.()Unaannulipes
,
,.

,
(Photoscourtesyof
PatriciaBackwellfromBackwelletal.,2000.)

,,
.ESS.,
.,
(BufoBufo).
.
.

(DaviesandHalliday1978).
,
.
,.,
,,
.
.,
(Grafen1990JohnstoneandNorris1993).,(Bradburyand
Vehrencamp1998).,.
.Ucaannulipes
,,
.
,(Backwelletal.
2000.14.3).

,,,
,.
,
.,
.

.,
,,
.
:
().
(14.1).
,
(),().
.
,

330

14
14.1.

,(
,,,,).

,
,

(,
)

Source: After Andersson 1994; Andersson and Iawasa 1996

.14.4.

(..10.5).140

.
,
(AfterGouldandGould1989,basedondataofB.
J.LeBandJ.Reiter.)

.,
(Trivers1972).,
,,
.,
,
.,
(.14.4)(WadeandArnold1980).
.,Phalaropus
(.14.5)(..14.19)
,..

.
,,.
.
(.14.6).
.,(Agelaiuspheniceus)
(Andersson1994).
.
100
,

,.,
80

10%-

.,
...
60
.
,
...
40
,

,,,
20
.

10
,(
0
10 20
40
60
80
100
120
140
)(WestEberhard1983).

,
140

331

.14.5. (Phalaropusfulicarius)
().,(
),.
,(PhotoK.Karlson.)

.,
,
.
,
(ThornhillandAlcock,1983Birkheadand
Mller1992).
(,,
).,
,
.Drosophila,

, (Partridge
andHurst1998).
.

(.14.7).,
(Parker1970Birkhead2000).

.,
(..11.20).
,
.Drosophilamelanogaster
,
(Clarketal.1994ClarkandBegun1998).

(,),
.

()

()

.14.6.

.()(Cervus
elaphus).()
(Babyrousababyrussa)
.

(1869)(Aphoto
byJ.H.Robinson/PhotoResearchers,Inc.)

332

14
.14.7. .
Calopteryx.()
.()(CourtesyofJ.
Waage.)

(Andersson1994Johnstone1995AnderssonandIwasa1996Houleand
Kondrashov2002).
,
.
,,.
,,(Euplectes),
(..11.9).
,,
.,(Physalaemusputulosus)
,,
,(Trachops
cirrhosus)(.14.8)..
,,
(Andresson1994).

.,,
,

(..15.6)
,
,
.:,
?
.1)2)
,3)4).
.

()

()

,
, ,
,
.
,
.
, ,
. ,
(Carpodacusmexicanus)
.
,
(Hill
1991).

.14.8. ()(Physalaemuspustulosus).()
().
..()(Trachopscirrhosus)
,(afterRyan1985MerlinTuttle/BCI/
PhotoResearchersInc.)


(Ryan1998).,
.

..
.,
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,
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333

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335

 14
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336

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337

 14
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339

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14

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341

342

14

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343

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14

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344

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345

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346

14
,
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347

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348

14

( )

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(AfterAtlanetal.1997.)

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15

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1997).,
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V
V

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349

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V

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350

14

1.
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:AnevolutionaryapproachJ.Alock(sevenedition,SinauerAssociates,Sunderland,MA,2001)
: M.Andersson Sexual selection
(Princeton University Press, Princeton, N 1994).
: Levels of selection, edited by L. Keller (Princeton
UniversityPress,Princeton,N1999).:L.D.Hurst,
A.Atlan,andB.O.Bengtsson,Geneticconflicts(1996,Q.Rev.Biol.71:317364).

:J.MaynardSmithandE.SzathmryThemajortransitionsinevolution(OxfordUniversityPress,1995).

1. F
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351

352

14

5.

6.

7.

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(Seeforexample,DalyandWilson1983versusKitcher1985.)
:
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15

.

.

(
).

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(..3.3).

,
. (Bombina
Bombina)(B.
variegata)
.

(Photographs
BorisI.Timofeev/PensoftPublishers.)

354

15

,
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1978).,
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).

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15.1.
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),,,
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Corvus corax ()

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Corvus corone ( )

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355

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,
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356

15
(,)
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15A.

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357

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,

.
.

(PhotoscourtesyofM.Cain).

Qercus grisea
(
)

Quercus gambelii
( )

358

15
.15.3. ()(Pipilomaculatus,),(P.ocai,)
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Pipio maculatus
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obscura
punctiferella
piperella()
piperella()
piperella()

G.piperella
G.mitellae

piperella()

359

.15.4. Greya()
.
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G.mitellae.,G.mitellae
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mitellae(,)

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).
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.,
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Hosogai1994)..Iris
fulvaI.hexagona(),,
()(Nasonetal.1992).
. (
).

360

15
15.2.

.15.5. (Chrysoperla)

.
C.adamsiC.johnsoni

,
,
(AfterMartinezWells
andHenry1992).

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.:()
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)
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.
)
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.
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C. plorabunda

C. adamsi

C. johnsoni

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361

.15.6.
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),
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).

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()

()

()

()

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.
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(From
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362

15

()

, .
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Cardinalis,,(Schemskeand
Bradshaw 1999 . 15.7).
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(Nilsson1983).

,
()

()

.15.8. Drosophila,

.
()D.simulans,()D.sechellia()D.
mauritiana.
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,
(Photoscourtesy
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.,
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,
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(Eberhard1996.15.8).
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,
,
(,).
,

363

(King1993Reiseberg2001).
,
(,
..
,
).
(CoyneandOrr1989).
F1,F2
.


,
F2F.,Drosophila
pseudoobscuraF2,
.,F1
,.,
,
.,
,(Dobzhansky1955).

()
,,
,
. , ,
(.15.9),
. ( )

.,
.
,
?(.9),,
,,.
,,

Phrynosoma

cornutum
solare

modestum

coronatum
mcalli

platyrhinos

douglassi

.15.9.

.
(Phrynosoma)

,,
,

(FromStebbins1954)

364

15

15B.
Opharella
, ,

.,O.notulata
Iva
.
Opharella

.

O. notulata ,
Ambrosia
artemisiifolia .
,
.
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(Futuyma 1991).
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.
,
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0,968 , Iva

0,989.
,
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,

.
Y
AmbrosiaIva

,
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.,

, ,

(Ambrosia Iva),

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.
,,,Iva
Ambrosia,
.

,Ambrosia,

Ophraellaslobodkini,O.
notulata.
Ophraella slobodkini

. ()
,,
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.,
,,
,
,
(.B).
,
.

.
,
.
.
,,
.
.,
Drosophilawillistoni,
,(.15.10).
(D)
,
.
(1989,1997)


()

.15.10. ,
Drosophilawillistoni()
,(),(),.
(I)()1
().I(D)
()(AfterAyalaetal.1974).

90
80

()

70
60

I=0,970

50

()

()

40

365

I=0,795

30

I=0,517

20

I=0,352

10
0

0.2 0.4 0.6 0.8 1.0

0.2 0.4 0.6 0.8 1.0

0.2 0.4 0.6 0.8 1.0

(I)

0.2 0.4 0.6 0.8 1.0

,

.60
,Drosophila
..
1.,
(.15.11).,
().
2.,
,D0,30,53,(
,Drosophilla),1,53,5
.,1
.
3. ,
,(
).
.,
(. 15.12). ,
.
4.

1.0

0.8
0.6
0.4

0.2

0.5

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1.5

() (D)

2.0

.15.11. Drosophila
(D)
..

.
(AfterCoyneandOrr1997.)

366

15

,

1.0

.15.12. Drosophila
(D).
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(AfterCoyneandOrr1997.)

0.8
0.6
0.4
0.2
0

0.5

.
.,

,.

,,
,
1.0
1.5
2.0
(D)
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(
),.
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.

()

,
,,
,.
,,
,,
.

Drosophila
,
.
(1936,1937).
,QTL()(.13)
.
(1984)D.simulansD.mauritiana
.F1,
.D.simulans,,
X().
D.simulansD.maurutiana,F1
D.simulans.,
,D.simulans.
(
).,
,(.15.13.).
,,D.simulans
D.maurutiana,.

.
DrosophilamauritianaD.sechelliaX
,,D.simulans
.,.
,X40


.15.13.
Drosophilasimulans
D.simulansD.mauritiana
,
.D.
simulansY().
,D.mauritiana
D.simulans
(1)(After
Coyne1984).

X
1

3
L R L R

0.10

0.30

0.20

2
3
4

-5 -6

, -3

.

5
6
7

120
8
(Wu and Hollocher 1998). Drosophila
simulans D. melanogaster
9
,
200 (Presgraves
10
2003).
11

. ,
12
D. pseudoobscura
200000
13
(,D.
14
simulans ).

0
,
D. simulans

D. mauritiana
(OrrandIrving2001).,

.

.,
.
(.15.14).
,,

(.15.14).

()

A'

0.20

0.30

()

C'

A B

C'

A B

A' B'

C'

0.10

367

C
C'

A
C

.15.14.
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.()AC
.().
X(AB)
,.

(A,B)(C).
.

C'

B'

C'

A'

C'
C'

A B

C'
C'
C'

,
, A B
, C
.

368

15
,..(1936)(1963).,
, ,
,.
,X
.X,
X
(. 15.14). Drosophila
,,.
X,X
(
X).,
,,,
,.
,
,(RitchieandPhillips1998).
DrosophilasimulansD.mauritiana
(..15.8).19
(Zengetal.2000).,
.,(Ostrinianubilalis)
()
(EZ11).
,E:Z.
E:Z,
,().,
,
,
E:Z,(Roelofsetal.1987).
,,
,.
F2QTL,(1998)
MimuluslewisiiM.cardinalis
(..15.7),,12,
16.
, 25%.
F2,
.,124
.QTL,
,.,
.

,(,,
),.

,.
(2003),Drosophilasimulans
,,D.melanogasteris
,(Nup96).30
,
.D.melanogaster
,.,
,,,
Nup96,(,
).,,

369

(R/S):

D.yakuda

5/43
D.melanogaster

22/69
D.simulans

D.mauritiana

0/0

(R/S)

3/10

16/21

.15.15. Nup96.
DrosophilasimulansD.melanogaster
.
/
.12

(AfterPresgravesetal.2003).

2/8

49+ 1/ 120

,

.

(.15.15).
,.

(.8)
(,.16).
(King1993Rieseberg
2001CoyneandOrr2004).,
()
,.
.,1.2
3.4,1234.
1.43.2.F1
(1.2,3.4,1.4,3.2).,(1.2
3.4,1.43.2),
()(..8.22).(,1.23.2
,3.41.4)(),
.
.(Dobzhansly1951),
DaturastramoniumD.discolor12.F1
7.5
.D.stramonium1.2,3.4,5.6,7.89.10,D.discolor1.3,2.7,
4.10,5.96.8.,Stramonium
,Discolor(.15.16).
.
,, () D. stramonium
D. discolor
1
, () .
2
1
3

2
3
4
7
, ,
5
6
4
10
.,
8
7
Spalaxehrenbergi
9
5
,.
9
10
6
,0,3
2,8(.15.17)
,
.
,,
.,
,
,,

.15.16. ()Daturastramonium
D.discolor
.

.()
F1.

()

F1

1
8

3
4

10

12
3

78
10
9

56

10
, Stramonium-
Discolor-
.

 15

27

26

30

30

.15.17. Spalaxehrenbergi.
.

.(AfterNevo1991.)

29

370

(.11).,
:,
. ,
. , Helianthus
annuus H. petiolaris ,
,(Riesebergetal.1999).

,.
,
,.(2004),
,
,,
.

,
,Wolbachia.
,
(Werren1998).Wolbachia
.Wolbachia

.,Wolbachia,
,Wolbachia.
,Drosophila,.
,,()
,,,
.,,,,
,(.
15.18,2).
Wolbachia,,,
.,Wolbachia
.,Wolbachia,
,.,Wolbachia
,
.()
.
().
NasoniavitripennisN.giraulti.

(. 15.18, 3 4 ).
,Wolbachia,
(.15.18,5
).,Wolbachia.A
,,
Wolbachia.
,(Werren1998).

1. G G (g)

g
G

3. G V (V)

g
G

v
g
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GG

2. G G

GG

, () V

v
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, ,

4. V G (g)

v
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g
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g
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5. V* G (g)

371

GV

Wolbachia-

. ,
,G,.A,
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,,,
.,A1A1B1B1A1A1B2B2A2A2B1B1
,A2B2
.
,1934
1940
.
,
,.

.
,,,
, ,
.,(1940),,

,.,,
,
.

.15.18.
Nasoniavitripennis(V)N.giraulti
(G).,

,
Wolbachia,

(v,g).Wolbachia
modvmod
g.(1)()N.giraultis
,
Wolbachiag.(2)N.
giraulti

()
.(3)

N.giraulti
.4)

.5),
Wolbachia
,,
F1.

372

15


.(,.15.11),
.,
.
,

.()
(McCuneandLovejoy1998).
( )
.,
,
(,),
().
(),
(.15.19).
(
).,
,,
(.15.19,t1).
10()
,
,,:
(coalescence)(
).
.
(15.19)

t3
f
1

g
t2

.15.19. ,,
.

,t1
.
,,
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()b.

,.t1
t2
,
,,
a,
,
(,
b).
(g)
t2,2,
.t2t3
,
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,
,e
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f
(AfterAviseandBall1990.)

t1
e
a
c
b
d

373

()
p

1
a

()
1

,,(15.19
,t2).,

,
,.
,
.
(.15.19,
t3).

,(Ne)(Neigel
andAvise1986PamilioandNei1988).

:.,
30
.11
,.
,
.,
(..2.19).
,
(MoranandKornfield1993).
,
.,
(MHC)
.,
5.
,MHC.

c
b

,
(,1990).
,
.
(Arnold1997).

(Harrison1990,1993).
(
).,,
,,
(..15.1).
.
,
.
. ,
, , ,
().,
(Endler1977).,,

,
(. 15.20). , ,
,

.15.20.
(p)
,()

()()
.
.()

,(
)
.d
()
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(
),,
d,

.
,
,

.
d
(),.

374

15
(
).(
)
()(.15.21).
,,
.
(Bombinabombina)
(B.variegata)6(.
).()
(. 15.21),
.
,
.
(Szymura1993).

,
.
,,,
(BartonandGale1993).,
,,,
.
,,
,.
,12(),
,A1A2,A1A2.

()

(p )
(log )

0.99

0.9

0.5

0.1

(p=0.5)

0.01

()

-20

-10

10

20

30

40

-20

-10

10

20

30

40

.15.21.
(BombinabombinaB.variegata).
60
().()
.()
.
,
,
(AfterSzymura1993).

(log )

0.99

0.9

0.5

0.1

0.01
0


,
.,,
.,F1
().,A1
A2,
()(.12).
.
F1,A1A2
.,
.A
.
,.
,B1B2,12
.B1B2,A
.,F1
.,B1B2.A
,A
(), A ()
.,(A),
. , B
,B1B2
,,
,()(..15.21).
,F1,A1A2B1B2A1A1B1B1
(F1)A1A1B1B2.
,B2,(),
A1A2,.,
B1.
(
),.

(),,
,
.,,
,,
,,.
,
,,
.(Rieseberg2001).(1999),
.
,
,().,
,
,
,.(
).,Bombina
,
55(Szymura1993BartonandGale1993).

8000
.,
(,)(Mayr1963).
:,.
:
1.

375

376

15
,.
,.

,.
,.
2. ,
.
3.,.
,,
.
4.
.
.

1. .
.
,
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,,,
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,
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3.
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, (, )
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4.
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.
,,.
5. ,
,
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,,.
.
.
6. . (
)()
..
7.
,.
.

,.
.
8. ().
,.
(
)
.,
.

(
)(BSC)

(
)

F2

(,
)

(
)

(PSC)
(
)
(
)

(
)

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. ,
.,,,
?
2. ,.
(),
(),()
()
?
3. .
,(1976)(1979).
4. .,
()()
.
5. ,

377

378

15
,,
(Avise1994pp..284290).,
,,
,()
.?
6. , ,
.

,.
.
?,
,.
??
7. ,
,2.19.
:
()?

16

.,

,
:

(. 1013 ).
,

,
,

,.
,
,,.
A1A1A2A2,A1A2
,
?(
,A1),A1A2(A2)
.
,.

. Helianthus
anomalus .
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.
a.
i.
ii.
iii.
b.
.
1.
2.
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-
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- ,
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.
a

,
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.
,.
:
.
,
.

(16.1).
,.
,
(.16.1).,
,
.
:(
.16.1),(
.16.1).
,
,
(.16.1).
,
(.16.1).,
.,

(),

() - () .

( ).

() .

() .

.16.1.

.()
.()

.()
.()
.

,

,
.

.

.

(
).,
.
, :
(),
,(
)..
, ,
.
,
,
,.
,.
,(11
).
,
(Muller1940Mayr1963).
(,
,),
.

.
,:,()
.,
.,,
.(..16.1).
.,
,
(, ,
).,
,(Mayer1963CoyneandOrr2004).
,
(,6.15).
,
.
,(.15,
).
,()
.

.
,,
.
/.,
(1990)
(Desmognathusochrophaeus).
()()
.
.
(.16.2).
(

381

382

16
()
2.00

()
.16.2. Demognathus
ochrophaeus
()
()
(
D,
)(AfterTilleyet
al.1990.)

1.0

100

200

2.0

1.0

300

()

0.2

0.4

0.6

D).
.,

. .
1%
(Knowlton1993).
,.,
(1986Avise1994)
.
.
(.16.3).
,
,.
,34.,
,.
,
. ,
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(Hills1998).
,(.16.4).,
:,,
/.
.,
,
,
(CoyneandPrice2000).

.16.3.
,
.,

.,
.:
Centrarchidae(),Amiidae(,)
Poeciliidae()(AfterAvise1994.)

Lepomis punctatus

Lepomis microlophus

Lepomis gulosus

Lepomis macrochirus


()

()

pipiens
blairi

.16.4. ()Ranapipiens(
)
.
()
,
.

.()Ranapipiens
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

,
.
,
.
.
,,
.
,,.(1936)
(1940).
.
.
(
).

()

,
.,.
,
..
,,
.,
(WernerandSherry1987..3.2218).
(2000),

,
.,
,

.
.,
,(.16.5,),
,
(.16.5,).

. 16.5.
,

.
. (Malurus )
, .

Rhagoletis-
,
(After Barraclough
and Volger 2000.)

(Malurus)

(Xiphophorus)

100

50

100

(Ellipsoptera)

(Rhagoletis)

50

 16

.16.6.
,

,
A2A2B2B2
.()
(
,

),

.
()

(A1B1).A1A2B1B2
(

),A1B1

()
A1

384

()

B1

A2A2B2B2

A2A2 B2B2

A1A1
A1A2 B2B2
A2A2

B1B1
A2A2 B1B2
B2B2

A1A1B2B2

A2A2B1B1

A1 B1

.

,A2A2B2B2(.16.6).
A1A2,B2B1.
A1A1B2B2A2A2B1B1.A1A2A1A1
A2A2,,
B2B2 .B1B2B1B1B2B2,
A2A2.,(
)().A1B1
:A1A2B1B2
,A1A1B2B2A2A2B1B1(
).,
,.
.
, C1 A1A1B1B1 ,
A2 B2 , .
D1,A2,B2C2.
,,
.,
(Orr1995).
,,
(
) (. 15 ).
. ,
,
.
. ,
,
,/.
,,,
,,,
(Mayr1963).
,,,
..
( )
, (
).
.

,,

385

,
( )
,
.
.
Drosophila
.
SI=0.86

. ,
SI=0.64

.(RiceandHostert1994).

, ,
(

)
SI=0.15
,.,

,,

(
.
)

,
(CoyneandOrr2004).,
.16.7. Neochlamisus
Mimulusguttatus
bebbianae
.,
(,)(Mcnairand ,
Christie1983).(1998),
.
,,
, (SI),

,,.
.
,Neochlamisusbebianae
, ,

(.16.7).
(AfterFunk1998

photobyChristopherBrown,courtesyofDaniel
.,,
Funk.)
,.
15(Mimulus)
(..15.7).(Gasterosteus),
.(
)(,).
.
, (. 16.8).
,
.
,,
(.16.8).,
.

()

()
0.5
0.4
0.3
0.2
0.1

.16.8.
(Gasterosteus).()

.()
.
.
,(
afterSchluterandNagel1995afterRundleetal.2000.)

386

16
(Rundeletal.2000).

.,.
,DrosophilaNup96(..15.15).
,.

.16.9.
.
,
,
(,),
,

(,).

.()
:
(Paradisaeaminor),()
:
(Manucodiacomrii),()
(Trochiluspolytmus)()
(Apusapus),
.(,
TonyTiford/photolibrary.com,W.Peckover,
VIREO,RobertTyrell/OSF/photolibrary.com
,NationalTrustPhotolibrary/AlamyImages).

( )
,
.(Lande1981PomiankowskiandIwassa
1998Turellietal.2001).
,(..15.6).
.
.,
,Drosophila,.
(Panhuisetal.2001).
,
(. 16.9). ,
,.
,,
,
(Mitraetal.1996).,,,
(

()

()

()

33

()

319

103
5

387

,
,.,,
,
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:
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B,



,.,,
,
.
(Dobzhansky1951,208).
,
.(1963).,
,
.
.:
.,
,,
,
.,
,
(Howard1993Noor1999Turellietal.2001).

,
,
!(:Grant1966

()

()

),.

.:
(MllerandCuervo1998).
(
).,(Physalaemuspustulosus)
(.14).
Physallaemus
.(RyanandRand1993).
,
.(McKayeetal.
1984Seehausenetal.1999).,,
.
(2000),
.,Ephippigerephippiger
,
.,
.,

(.16.10).

,,.,
,
.?,
,
(HollandandRice1998.14).

1.00
0.75
0.50

0.25
0

1.00
0.75
0.50

0.25
0

.16.10. Ephippiger
ephippiger
.

.
()

,
()
.

.(AfterRitchie2000.)

388

16
andCoyne1974).
,
.,
.,
.
.,(Sorexaraneus)
,
.,
,(Searle1993).
,
.,
,,(Brown
andWilson1956)1*.
,,
.(Felsenstein1981Sanderson1989).,A1
A2,(),
F1 A1A2 (s). B
,,B1
B1,B2
(B2
,).
B1.,
B2(,),A1
A2,
.
A1A2(s=1),.
B2,A2.
,B2,,A2A2,
A1B1 A2B1
.B2B1,,B2
:,.
,B2.
,,,
.,B2,
.,
B1,,B2.
,.
B2
,A2,
A1A2.,F1(A1A2)
A1A2
A1A1,A2A2.B2A2
.A1B2,A2B2
,A1A2.
,B2,,
,A1A2.B2
,,
.,,,
1
(1989).,
,
, , , .
( ),
,,,
().
,
, , w
h.R(Howard1993).

389

()

1.5

1
0.5
0
-0.5

-1
-1.5
-2
-2.5

-2

-1.5

-1

-0.5

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
,
,,
.(Soule1966).
,,,
,.
,,
, ,
,
(
.10).
,

.16.11. (Hylacinerea)

.()
(
),

.()
Hylacinerea
,

,
.

,
,
(AfterHbel
andGerhardt2001.)

390

16

()

(w )

(w )

,.
,
,
.
,.
(1954),,
,
,.
F1
,,
,
.,

.,

(.421),

A1 -

B 1(1972).
(1975).,

. (1980) ,
()

,,,
.
,,.

(..12.20).,
( ),
F1
(),
(.16.12).,
,
.
A1
,

.,
.
(),
,
.,
.16.12.
,(,A1A1B1B1A2A2B2B2)
,
.

( w ).
AB
.()

()
.
F1
,

.()
,

F1
(AfterGavriletsandHastings1996).

.
()(CharlesworthandRouhani1988Turelli
etal.2001).,
.(
),
,.,
,
.,,
,
.,
,,
.
,(Priceetal.1993
Wagneretal.1994Gavrilets2004).,




(
)(.16.12).
,(..16.6).
,,

391

,
.
(Gavrilets2004).
, . ,
,
. ( )
.
,
.
,
(Avise1994).GreyapiperellaGreyamitellae
15.4..

, (, ),
. ,
.,
,,
.

-
Sim-13

Sim-1

Sim-5

Sim-8
Sim-7

Sim-8
Sim-1
Sim-2

Sec-2

Mau-4
Mau-9
Mau-8
Mau-2

Sec-8
Sec-2
Sec-5
Sec-6
Sec
Sec-3
Sec-4

Sec
Sec

Sec

Mau-5
Mau-3
Sim-3

Sec-7

Sec

Sec
Sim-3
Sim-4

Sim-10
Sim-6
Sim-13
Sim-12
Sim-3
Sim-2
Sim-6
Sim-8
Sec-1
Sec-7

Sim-12
Sim-8

D. melanogaster
()

Sim-10
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

-0,3 -0,2 -0,1 0

0,1 0,2 0,3

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

100 200 300

400 500 600 700 800 900 1000

(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


).
,1,5(Sasaetal.1998
.16.23,I).
R,
(S)(E).R
,(t),
(N):
Nt=eRt
(7,
.141).1/R
(.16.21).,(E=0),
BSI.
,BSI0,3(
),10(
).E,BSI3
,(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.

4.

5.

6.

7.

8.

9.

10.

11.

.
.

,,
.
( ). ,
.
,
,,
, .
.
.
,,
,.

.,
.
,
,,
.
.
.
,
.
,
,
. ,
,,,
,.
,.
,,
,.
.
, , .
.

,
..
()
.
().
,,.
(, )
,(

).
23
.
..
.

.
.
(),
.
,
.

403

404

16

()

15 ,
:Speciation,byJ.A.CoyneandH.A.Orr.
,.
:SergeiGavrilets.Fitnesslandscapesandtheoriginofspecies(PrincetonUniversityPress,Pinceton,
NJ2004).

1. ?
2. (1997) ,
,.
.

,.,

.?(
).
3. ,.
,
??
4. (Gasterosteus),
()?

?
5.
.(
)
?
6.
.
,().
?
7.
.
.
?
8. ,
.

17

.,

, (Pinus aristata)
4600,
(Poplus tremuloides)
10000.
,
, ,
,
.

.
,
,.(Balaenoptera
musculus),
.(Apteryx),
25%(.17.1).

(Pinusaristata),
4600.
(PhotoD.Cavagnaro/VisualsUnlimited.)

406

17

.17.1. (Apteryx)
,

25%(Photocourtesy
OtorohangaZoologicalSociety.)

.
:Drosophilamelanogaster10
,
.(Magicicada)1317,
,,(.17.2).
,,
,,(Agave)
,.
.
,.
.
,
.
.
.,:
,,
,.
.(
),,.
,
..
,
?
.
.

,?
??
??
,?,
?
,,
.,
(),,

()

()

.17.2. ,,
.()(Magicadaseptendecim).17
,.
,..
()(Aphisnerii).
.
.(JimmZipp/PhotoResearchers,Inc.photolibrary.com.)

 :
.

.,
,
.,,,,11
.,
.
,
,.
,(),
(),
.
,.
,,
(
)(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
1,00
0,75
0,50
0,25
0,10
0,00

mx
0
0
4
8
0
0

lxmx
0
0
2
2
0
0
4

407

408
()

17

R:

R lx mx

(Lx)

D. serrata-

1.0

0.8

,R=4(
,
).
:N0,
g:

0.6

0.4

N=N0Rg

0.2
0

10

20

30

40

50

60

70

80

90

()

()

4 (mx)

100
80

R
.
,

,.
r,R
R=er.N0,t
:

N=N0ert

60

R,r
.r
40
.
,xlx
20
R ( r) .
(,
0
)
30
20
10
0
40
50
60
70
(mx=0),
()
R.,
R.
.17.3.
,mx(x).
.
,
()
.,
lx()mx
.,
.
.,2
Drosophilaserrata
,,3,

.3,2,
,
2.
.
,,,2,

(),3.
.
,,

(r).

.
,
.

(AfterBirchet
al.1963.)

,
,
,.
,.
.
,.
,
,
.,
..

 :
()
x 1-x
,
.

A,A

B,B
1-x

.
, ,

()

.17.5. (Solidago)()
.
,
,
(AfterWernerandPlatt1976.)

.17.4. ,

(
)
.()A

.B

,
,x
1x.()A

,B
.

,Procellariiformes(,)
.
. ,
.
(tradeoff),
.,
:
(Physalaemuspustulosus)
(.14).
.
.,

,,
.
.
,
.
,
(.17.4)(vanNoordwijkanddeJong1986BellandKoufopanou1986).
105
,
.
(Reznick1985)
1.
.,
104
.,,
,,:

.(.17.5).
2.
103
,

102

409

102

103

(g)

104

410

17
.17.6. Poaannua
.
.
,,,
(Afterlaw1979.)

,


.

(.13).,
(Poaannua)
(Lawatal.1979).,
,,
(.17.6),
.
1000
.
3.
(Reznick1985Stearns1992).
(1999) Drosophila melanogaster
10.

,34
. 19

,
,
3,(.
0
300
600
17.7).1

(.
17.7).,
.,
.
4.
.,Drosophila
(SgrandPartridge1999).

.
,,
1
0
-1
-2
-3
-4
-5
-6

10

20

30

()

40

50

60

12
8
4
0

Loge

16

Loge

2000

()

-1.5
-2.5
-3.5
-4.5
-5.5
-6.5

10

20

30

40

50

60

70 80

()

.17.7. Drosophila.()
10,
,,()
,.()
,
., (AfterPartridgeet
al.1999andSgrandPartridge1999).

411

:
(.17.7).
Drosophila,,
,
(FowlerandPartridge1989BellandKoufopanou1986).

, ,
(
(r)),,
(Charlesworth1994).17.1
,(l1)
.,56(l5),
5(m6=0).,,
,(
,,
).
,,
(r),,
(17.1
St(x)Sm(x)).,
.
,,
,.

, ,
..?
(Rose1991).
,
.
(1952),,

17.1.(r),
a

lx

mx

r
mx

r
lx

lxmx

erx

erxlxmx

Sm(x)

Ss(x)

1000

1.000

0.00

0.00

1.000

0.000

0.335

0.334

750

0.750

0.00

0.000

0.796

0.000

0.200

0.334

600

0.600

1.20

0.720

0.634

0.456

0.128

0.182

480

0.480

1.40

0.672

0.505

0.339

0.081

0.068

360

0.360

1.03

0.396

0.402

0.159

0.049

0.018

180

0.180

0.96

0.144

0.320

0.046

0.019

0.018

100

0.100

0.00

0.000

0.255

0.000

0.011

1.932=R

1.000

Source: after Stearns 1992.

ar,

x z

1 e rx l m m x 0,228.
x a

Sm(x)Ss(x)x(mx)(lx)r.

y z ry
rx
e lymy
: Sm ( x ) e l m Ss ( x )

yx

412

17
, ,
.
,.
,(,
),.
(1957).
.,,
,(,,
),,
(,,
).
.

.Drosophila
, (Charlesworth 1994).
,
(..17.7).
:
..

.17.8.

(
)(
).

(AfterWilsonandThompson1989.)

10

0 0.3 0.6 0.9 1.2 1.5 1.8 2.1

,
?,,
,
(.17.6).,
.,
.,,
?
,

,r ,
.,
,..
,,
,
.,,
,
,.
.
,,
,
,,
.
,
.,,

(Williams1966Charlesworth1994).
.
,
(Roff2002).
,
,(.17.8WilsonandThompson1989).,,

,,,
,:

413

Loge

2
(PeomislowandHarvey1991ShineandCharnov1992).

1.75


(Poeciliareticulata)(e.g.,Reznicketal.1990ReznickandTravis2002).
1.5
Crenicichliaalta()(.
1.25
11).Crenicichlia
1
.Crenicichlia
.75

,
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,(
.25
)
0
,.
-.25
Crenicichlia
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,.
-2.5
-2
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-1
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.
,
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,
,
,
.

(.17.10).

,,
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(1954),
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,
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,,

,
.
,

(AfterShineandCharnov1992.)

60

50

90

80

70

.
,
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()

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300

200

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0.90

0.80

100

.17.10.
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.()
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()(fromReznickandTravis2002.)

414

17
()

1.2

23

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1.6

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0.6

0.4

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12

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16

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)(
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,,

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rm,A
rm,B

,
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,
9(.17.11).
,
, . ,
,()
,
.,
(,
,.
).(1990)Sceloporusoccidentalis
,.
,.

.
.,
,.

(r),
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,,
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.,
,r
,()(rm),
,,
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), . (K) ,

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(N)

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415

,
(r).
, (. 7 )
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.,
.(.14)
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.
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,,.,
.
,,.
,(Oncorhynchuskisutch),
. ,
,,
,.
,
(Gross1984).
,,(
).,
,,
(.17.14).,(Thalassomabifasciatum)

()

(/;
)

()

.17.13.
.()
()
.,
,
.
,
.()

(PeterArnold,Inc./Alamy
ImagesafterVehrencampetal.1989.)

2600
2400
2200
2000
1800
1600
1400
1200
1000
800

100 200 300 400 500 600 700 800 900 1000

 17

.17.14.
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,
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(),
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,
,(AfterWarner1984.)

416

()

()

()

,,.
,(.17.15).
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.,
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,
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.

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,,
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,,
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,
.

,
.:,
.,
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()
()

,
,

.

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.()()()
..(GregoryG.Dimijian/Photo
Researchers,Inc.)

 :
,
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,,
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.,.
(),
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(LynchandLande1993).
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.

,
.,,,



.,.
,
.,
.,.
.

.,,,
.,

.
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,
,(Leigh1973Sniegowskietal.2000).
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(.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

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1.6
.
1.5
,
1.4
, .
1.3

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1.2

1.1
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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).
.,
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, .

()

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,.
,
,
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.

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,

(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.
,
.
,
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2/3,
1/3,
.S
,
aabb

420

17

,(,
).,,
.,,
,,,.
..
,,,
.
, , (Lynch et al. 1993).
,
,
.
,
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.
.

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(010)
.

(01)
.
,
.
0
,

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0
3

. ,
,,.
.(1988),
,
.,
( w ).
,,
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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

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AC

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B

AC

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AB

4: ,

AB

0.6
0.4
0.2
0

.17.22.
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.
(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

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..

A1A1
A1A2
A2A2

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1

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1:1 .

..

A1- ,
A1-
.

.17.23. ,,
.A1,,
..
.A1.A1
,,A1(,
,,).,.
A1(AfterWilsonandColwell1981.)

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.,,
.(,
,
)
.
,

.
,
,
.
,(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. (,
),
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9. ,

10.

11.

12.

13.

14.

15.

,.,,
,
.
,
.
.
,
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,

,
.,
.
:()
()
,
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,()

()
, , ()
,,
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,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)

, .

.
,

.

. ,
.
,

,.
,

,
.
,
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,
.

(,).
, , (
),(,),(,
)(,,
). (
).,
.,,
,.,,
(..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

Source: After Frank 1992.

: (
) () () . + ,

( ). , .

()

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).

Thamnophis sirtalis TTX

,

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
.
.,
,
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43
,40
(AfterDybdahlandLively1998.)

0.3
0.2

12

22

19

63

439

(. . 17.21).
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.(.18.14)(17
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).

,.

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, , ,
.
Angraecumsesquipidale30,
,,
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.(.18.15Nilsson
etal.1985).
,
,
..
..
.
.,(Yucca)
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(.18.16)..Tageticula

440

18

()

.18.15. .()Angraecum
sesquipedale.()
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GeraldandBuffCorsi/CaliforniaAcademyofSciencesTheNaturalHistoryMuseum,London.)

()

.
.
,
(.18.16).

,,
, .
. ,

(Bull and
Rice1991).,
,
(Herreetal.1999).,
,
,
,

()

.18.16.
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.()
.
,Greya,
.Tegeticula
Parategeticula
,Tegeticula
(courtesyofO.Pellmyrafter
PellmyrandLeebensMack1999.)

Tegeticula
yucasella

Tegeticula synthetica
Tegeticula maculata

Yucca

()

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Prodoxus

Mesepiola
Lampronia
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441

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,
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,(5001500
).,
15%,.
.,,,
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,.
,,.
,
.
.
.,.
:..
(..11.2).
,
,(..18.16).
,.
(Pellmyrand
LeebensMac1999).

Bombus

100

appositus

50
0

100

12

12

12

12

12

12

12

kirbiellus

50
0

100

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50
0
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, , .
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()(
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,
(Schluter2000).
,
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(,),
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,
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)
,,.

100

sylvicola

50
0

100

bifarius

50
0

100
frigidus

50
0

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.
().
.,
.B.appositusB.kirbyellus,B.frigidus,B.bifariusB.sylvicola
(AfterPyke1982.)

100

occidentalis

50
0

()

442

18
x, (, )

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.18.18. .x
(z),,.
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.,
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(,).

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Melanerpes striatus

()

Melanerpes aurifrons

.18.19. .

Melanerpesstriatus().

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()
.

.
.
(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)

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(.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

(CG)
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()
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.18.20 ()
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,
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)()
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.
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Lososetal.1998.)

 18

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,
.
.
,

,,
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Schoener1984.)

,
Accipiter 47 .

(
)
.

0.15

444

0.10

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1.05

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1.25

1.35

1.45

1.55

1.65

1.75

1.85

1.95

( )



.
. (1999)
,,
,
.
,,

.
(Benkman1999Benkmanetal,2003)
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contorta)(Tamiasciurus
hudsonicus)(Loxiacurvirostra)
(.18.22).

.
.,
,
,.
.
B
.
, .

.18.22. .(Pinuscontorta)
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,(Loxiacurvirostra)
().()
.(After
Benkmanetal.2003.)


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arthemis)(Battusphilenor).
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(LimentisMichaelGadomski/PhotoResearchers,Inc.BattusS.McKeever/PhotoResearchers,Inc.)

,,,
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,
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.

445

()

()

. ,
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(MalletandJoron1999JoronandMallet1998Turner1977).,
(.3).
(.18.23).
.
(

,).
().
.
.
.
.
()(.18.24).
. (
,Papiliodardanus..9.2).

H. melpomene

H. etaro

.18.24. .HelioconiusmelpomeneH.
erato,

,,
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Helioconius,
(
)()
(CourtesyofJ.Mallet.)

446

18
: ,

()

1.0
0.9
0.8
0.7
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()

0.60
0.59
0.58
0.57
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0.75
0.70
0.65
0.60

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.(AfterMalletand
Joron1999.)

,
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. ,
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,
.(.18.25)(
,,
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:,,
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,(.
18.25).
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( )

1. ,
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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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2.
,?
3.
50100
.,

447

448

18
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.
5. ,
(e.g.,BernaysandGraham1998).
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450

19

(A).200
,(,,,
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).
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,
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19A.
,
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451

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452

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453

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25

454

19
()
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(w>1),.
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(SwansonandVacquier2002).
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),
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()VERL
(.19.4).w,
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()
.,
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),,
(SwansonandVacquier2002).

,,
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.19.4. VERL

.(
)
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.
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,
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455

,
(.19.5).(1997)

,(
Colobinae)
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.
.
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/
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,(Stedmanetal.2004).MYH
.w0,11
,,.
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.
(OR) , (
)
. 1200 OR,
550.,OR
,
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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
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Glu126

.19.5.
,

.
,
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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).
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1,5%.
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10%,
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.(50000)
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(RobertsandSmith2002),
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:
(.17),
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EUCARIA

Plasmodium
Dictostelium
Entamoeba
Naegleria

Trichomonas
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.19.6.
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(AfterPalmerandLogsdon1991.)

Giardia

ARCHAEA

Thermotoga
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BACTERIA

Oscillatoria
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457

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458

19
(..18.3).
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,(.19.8)
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)(DeiningerandBatzer2002).
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,Y
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ALU
L1
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L2
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1.0

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3 5

7 9 11 13 15 17 19 21 23 25 27 29 31 33

460

19
.(V,DJ),
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.,

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(LGT).

.,BacteriaArchaea
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50.
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.(.19.10),
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4050(Salzbergetal.2001).
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,
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461

462

19
N. meningitides

E. coli-2

P. gingivalis

C. jeijuni

Synechocystis sp.

V. cholerae

P. furiosus

T. maritime
H.sapiens

P. prowazekii
H.sapiens

C. dicile
Entamoeba histolytica- .
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.19.10. Archaea
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.
400.
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,
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N. frontalis

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A. fulgidus

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A. pernix

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.,
,(
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,

.19.11. .
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.

463

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.
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D.yakuba.jingwei.Drosophila
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D.teissieriD.yakubaYmp
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464

19

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2,5
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(Adh
),Adh
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(Zhangetal.2004).
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(AfterLongetal.2003)

(GPCR),GPCR
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2003Zhangetal.2003).
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Thr Ala Ala

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.
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()
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(Guetal.2002)1739749
.,
(500),
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,(COL),6
(MHC)
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(. 19.18).
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19.19,(
Dujonetal.2004Guetal.2002)800(,
)1000(,).
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465

.19.15.
AFGP.()

(,
AFGP2(E2))
.()
2

,
16(
),AFGP
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2.
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,
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(AfterChengandChen1999.)

466

19

1
1

2
2

.19.16. .(a)
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1000,
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.
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,

.19.17.

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.,
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100
I

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50

50

150

250

350

450

550

650

750

850

950

1,050

Amphioxus-

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1,150 >1,200

467

.19.18. .
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6p (MHC)

19p

1q

9q

RXRB

RXRG

RXRA

COL11A2

COL11A1

OL5A1

RING3

RING3-

LMP2/LMP7

PSMB7

TAP1/TAP2

ABC2

NOTCH4

NOTCH3

NOTCH2

NOTCH1

PBX2

PBX1

PBX3

TNX

TNR

HXB

CYP21

CYP2

CAA/C4B

C3

C5

HSPA1

(HSPA6/7)

GRP78

HlA-A-A,-B,-C

CDI

VAV1

VAV2

LMNB2

LMNA

SPTA

SPTAN1

ABL2

ABL1

Notch

(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


()

()

()

(Zhangetal.1998),ECP,EDN,
.,EDN,ECPa,
,