141 !"#$%& '(%), 2010, *+, 14, - 1 !"#$ %&'()*+-,-&$( ./ 011&!)( 1(-2*-( #'*& !("*3&,) &''./ +-*4'#!( 5 *'6-&$'./ * #5)6-&$'./ -#,)&'*7 . . ,89:;, &. . 2<=>9?@, $. . !:ABC9:; !" #$%$&'()* +$,-+.-)''/01.2$30/4'()* )+'3)3,3 '0/4'(.5. 6.78*'32$ )%. !.9. :,/$*(.2$ ;<#=!, >070+-,(, ;.'')8, 0-mail: [email protected] ?@'($7$+. ) .A.'+.2$+. B&01B./.C0+)0 .A DB)50+03)-0'(.% 6$&$(30&0 27$)%.10*'32)8 «50+.3)B–'&01$». E&) .F0+(0 50+03)-0'()6 2$&)$+' , &$7+.2.7&$'3+@6 B.3.%(.2 D3. 27$)%.10*'32)0 G)(')&,03'8 ($( DB)'3$3)-0'()0 DGG0(3@. E.($7$+$ +$ )+A&01+.% (8&.2$8 %85($8 BH0+)F$) ) $,3A&01+.% (5)A&)1+$8 (,(,&,7$) .AI0(3$6 7$(.+.%0&+.'34 A./00 +)7(.5. ,&.2+8 2(/$1$ 50+.3)B-'&01.2@6 DGG0(3.2 2 G.&%)&.2$+)0 (./)-0'320++@6 B&)7+$(.2 , 5)A&)1.2 B. '&$2+0+)J ' 5.%.7)5.3+@%) G.&%$%). "-)3@2$8 2 7+$-)30/4+.* '30B0+) DB)50+03)-0'()* 6$&$(30& 27$)%.10*'32)8 «50+.3)B–'&01$», D3, 7$(.+.%0&+.'34 %.C+. .AI8'+)34 ' B.7)F)* 5)B.307@ «DB)50+03)-0'(.5. A$/$+'$». !D8AB;EB FD:;<: 27$)%.10*'32)0 «50+.3)B–'&01$», DB)50+03)-0'($8 )7%0+-)2.'34, DB)'3$7, DB)50+03)-0'()* A$/$+', BH0+)F$, (,(,&,7$. ;B=B@?B ? '3&,(3,&0 G0+.3)B)-0'(.* )7%0+-)2.'3) (./)-0'320++@6 B&)7+$(.2, ' (.3.&.* )%003 10/. '0/0(F).+0&, .'.A.0 %0'3. 7$+)%$03 27$)%.10*'32)0 50+.3)B–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aligari, Mather, 1975; Connoly, Jinks, 1975; Hill, 1975; Jinks et al., 1977; Jinks, Pooni, 1982; O&$5$2F02 ) 1&., 1984; N)/4-02'()*, =.3@/02$, 1997; Dorn, Schmitt, 1997). C.R. Landry ' '.$23. (2006) +$ 6 H3$%%$6 1&.CC02@6 5&)A.2 2 3&06 D(./.5)-0'()6 ,'/.2)86 +$H/) 221 3&$+'(&)B3, .3203'320++@* 7$ 27$)%.10*'32)0 «50+.3)B–'&01$». E.7C0 +$ 3.% C0 .AI0(30 1/8 2 H3$%%.2 ) 109 &0(.%A)+$+3.2 .3 '(&0L)2$+)8 D3)6 H3$%%.2 +$ 12,6 G.+$6 ,5/0&.1)'3.5. B)3$+)8 2@82/0+. 1555 50+03)-0'()6 'F0B/0+)* 1/8 1382 3&$+'(&)B3.2, (.3.&@0 B.($7@2$/) ',L0'320++.0 50+.3)B-'&01.2.0 27$)%.10*'32)0 (Smith, Kruglyak, 2008). Y. Li ' '.$23. (2006) +$ Caenorhabditis elegans +$ G.+0 12,6 30%B0&$3,&+@6 &0C)%.2 2@82)/) 197 50+03)-0'()6 'F0B/0+)*, .3203'320++@6 7$ B.1.A+@0 27$)%.10*'32)8. E. M. Rutter ' '.$23. (2006), «genotypeenvironment interplay» – D3. .AL)* 30&%)+, (.3.&@* B.(&@2$03 +0'(./4(. &$'6.18L)6'8 B.+83)* ' &$7/)-+@% '.10&C$+)0%. ?.-B0&2@6, D3. 27$)%.10*'32)0 'B0F)G)-0'()6 )10+3)G)F)&.2$++@6 50+.2 ) .B&010/0++@6 '3&0''.2@6 ,'/.2)* '&01@, 3. 0. 'B0F)G)-0'($8 $1$B3)2- 142 +.'34, (.3.&$8 +0 82/803'8 .AI0(3.% +$H05. &$''%.3&0+)8. ?.-23.&@6, D3. 3$( +$7@2$0%@0 50+.3)B-'&01.2@0 (.&&0/8F)), 2'0 3&) 3)B$ (.3.&@6 (B$'')2+$8, &0$(3)2+$8 ) $(3)2+$8, Plomin et al., 1977) .3+.'83'8 ( B.2010+-0'()% G,+(F)8% -0/.20($ ) +0 '2.*'320++$ (./)-0'320++@% B&)7+$($% &$'30+)*. N 3&0340%, 2)1, 50+.3)B-'&01.2@6 DGG0(3.2 .3+.'83'8 )7%0+0+)8 D('B&0'')) 50+.2 B.1 2/)8+)0% G$(3.&.2 '&01@, 2@72$++@0 DB)50+03)-0'()%) %06$+)7%$%). E&) D3.% )%0J3'8 2 2)1, 3$()0 DB)50+03)-0'()0 )7%0+0+)8, (.3.&@0 B0&01$J3'8 B. +$'/01'32,. ?B0&2@0 B.1.A+@0 G0+.%0+@ A@/) 2@82/0+@ A. Durrant (1958, 1962). P+ .A+$&,C)/, -3. +0(.3.&@0 '.&3$ /4+$ +$ G.+$6 %)+0&$/4+.5. B)3$+)8 'B.'.A+@ A@/) B&).A&03$34 B0&01$2$0%@0 B. B.3.%'32, +.2..A&$7.2$+)8, '6.1+@0 )/) )10+3)-+@0 &$+00 2@82/0++@% B$&$%,3$F)8% (Brink, 1956, 1960). :$()0 C0 )7%0+0+)8 A@/) .A+$&,C0+@ B.7C0 , 3$A$($ (Hill, 1967; Hill, Perkins, 1969). G.M. Evans (1968) .A+$&,C)/ '2874 «'&01.2@6» )7%0+0+)* ' )7%0+0+)8%) 2 '.10&C$+)) O!N. !@+0 2@82/0+@ A).6)%)-0'()0 %06$+)7%@ (%03)/)&.2$+)0 F)3.7)+.2@6 .'+.2$+)*, $F03)/)&.2$+)0 5)'3.+.2), .3+.'8L)0'8 ( 3&$+'(&)BF).++.%, 3)B, B.1$2/0+)8 D('B&0'')) 50+.2 (Chang, Whitelaw, 2004; N.&.-()+, 2006), $ 3$(C0 .A+$&,C0+@ %)(&.-;!N, .B&010/8JL)0 B.'33&$+'(&)BF).++.0 &05,/)&.2$+)0 $(3)2+.'3) 50+.2 (N$3.6)+ ) 1&., 2006). P1+$(. DB)50+03)-0'()* 6$&$(30& )%0J3 ) 50+.3)B-'&01.2@0 DGG0(3@, '287$++@0 ' )7%0+0+)0% D('B&0'')) 50+.2 2 B&.F0''0 )+1)2)1,$/4+.5. &$72)3)8 ) G.&%)&.2$+)8 F0+.7.2 A07 7$(&0B/0+)8 D3)6 DGG0(3.2 2 B.3.%'320. :$(, A.H. Paterson ' '.$23. (1991) +$ 3.%$3$6 2@82)/), -3. 1/8 .1+.5. ) 3.5. C0 B&)7+$($ 2 &$7+@6 D(./.5)-0'()6 ,'/.2)86 B&.82/8J3'8 &$7+@0 QTL (quantitative trait loci). <+$/.5)-+@0 &07,/43$3@ B./,-0+@ M.C. Ungerer ' '.$23. (2003) +$ $&$A)1.B')'0 ) A. Börner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«DB)'3$7». E0&2.0 B.+83)0 (A)./.5)-0'()* DB)'3$7) 22010+. >D3'.+.% (Batson, 1909) ) .B&010/0+. ($( B.1$2/0+)0 10*'32)8 .1+.* $//0/) 1&,5.*, +0$//0/4+.* B0&2.*. J.B. Holland (2001) .B&010/)/ %06$+)7%@ 10*'32)8 A)./.5)-0'(.5. DB)'3$7$ ($( 27$)%.10*'32)0 D+7)%.2 –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– '3$3)'3)-0'()* DB)'3$7 – ($( /JA.0 +0$//0/4+.0 27$)%.10*'32)0 50+.2 220/ R.A. Fisher (1918). M.J. Wade ' '.$23. (2001), '&$2+)2$8 B.+83)8 «DB)'3$7» 2 12,6 30.&03)-0'()6 3&$1)F)86 D2./JF).++.* 50+03)(), 50+03)() B.B,/8F)* ) (./)-0'320++.* 50+03)(), B.($7$/), ($( +07$2)')%.'34 10*'32)8 50+$ (B&01'3$2/0++$8 %,/43)B/)($3)2+.* %.10/4J B.B,/8F).++.* 50+03)()) %.C03 .3/)-$34'8 .3 .3',3'32)8 DB)'3$7$ 2 B&01'3$2/0+)) (./)-0'320++.* 50+03)(). P+) 10/$J3 [email protected] . B&8%.* $+$/.5)) '3$3)'3)-0'(.5. DB)'3$7$ 27$)%.10*'32)J 50+.3)B-'&01$. E&) )'B./47.2$+)) %./0(,/8&+@6 %$&(0&.2 +$ (,(,&,70 (Blanc et al., 2006) ) $&$A)1.B')'0 (Juengei et al., 2005; Malmberg et al., 2006) 2@82/0+. 7+$-)30/4+.0 (./)-0'32. QTL-QTL DB)'3$3)-0'()6 27$)%.10*'32)* B. '$%@% &$7+@% B&)7+$($%. E&)-0% 20/)-)+$ DB)'3$3)-0'()6 (3. 0. 50+.3)B-'&01.2@6) 27$)%.10*'32)* B&)%0&+. 2 12$ &$7$ B&02@H$03 $11)3)2+@0 DGG0(3@. E&) D3.% '/01,03 7$%03)34, -3. 0L0 2 (.+F0 1950-6– +$-$/0 1960-6 55. A@/. 2@82/0+. +$ 1&.7.G)/0 (Parsons, 1959) ) 1.%$H+0* B3)F0 (Hull et al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– B0&2@* G$(3.&, – 23.&.* G$(3.&, A/B – 27$)%.10*'32)0, / – B.23.&0+)8, Z – .H)A($) 2 12,6G$(3.&+.% 1)'B0&').++.% (.%B/0('0 (E/.6)+'()*, 1978). P1+$(., ,-)3@2$8, -3. %0&.* )7%0+-)2.'3) (./)-0'320++.5. B&)7+$($ 2 B.B,/8F)86 82/803'8 1)'B0&')8 (U2ph = U2g + U2e + U2g/e; Falconer, 1981), F0/0'..A&$7+00 )'B./47.2$34 B$&$%03&@ V2, B&01'3$2/8JL)0 )7 '0A8 %$30%$3)-0'()0 .C)1$+)8 '..3203'32,JL)6 1)'B0&')*, &$''-)3@2$0%@6 +$ .'+.20 '3&,(3,&@ '&01+)6 msA – msZ 2 (2$1&$3.2 .3(/.+0+)* (V2g = n·c ; V e = msA/B – msZ 2 msB – msZ 2 = ; V z = msZ, 510 n – n·r n·r ; V g/e = (./)-0'32. B.23.&0+)*, c – (./)-0'32. D(./.5)-0'()6 3.-0(, r – (./)-0'32. 50+.3)B.2; ;.()F()*, 1973). "-)3@2$8, -3. 2.72010+)0 .3(/.+0+)* 2 (2$1&$3 82/803'8 'B.'.A.% )6 %.1,/)7$F)) ) -3. B.1.A+$8 F0/4 1.'3)5$03'8 2.72010+)0% 2..AL0 2 /JA,J -03+,J '30B0+4, 1/8 2.72&$L0+)8 (&)30&)8 )7%0+-)2.'3) 2 )'6.1+,J 01)+)-+,J '30B0+4 ' F0/4J )'B./47.2$+)8 2 ($-0'320 %0&@ 2(/$1$ .310/4+@6 DGG0(3.2 2 G0+.3)B)-0'(,J )7%0+-)2.'34 %@ B&)%0+8/) B.($7$30/4 V. ? ($-0'320 )+A&01+.5. .AI0(3$ )''/01.2$+)* A@/) 2783@ '.&3$ 8&.2.* %85(.* BH0+)F@ E&.6.&.2($, :,/$*(.2'($8 5, :,/$*(.2'($8 10, :,/$*(.2'($8 100, :,/$*(.2'($8 7./.3)'3$8, :,/$*(.2'($8 .'3)'3$8 ) Q5.-?.'3.-+$8 2, (.3.&@0 )'B@3@2$/)'4 2 8 D(./.5)-0'()6 '&01$6. PB@3@ 7$(/$1@2$/) +$ B./86 #$%$&'(.5. !MM#= )%. !.9. :,/$*(.2$ 2 2006–2008 55. +$ 12,6 G.+$6 (B. B$&.2.%, ) 70&+.2.%, B&01H0'320++)($%). E/.L$14 10/8+() 25 %2, B.23.&+.'34 – 3&06(&$3+$8. E.'02 B&.2.1)/'8 '08/(.* #!-10W, ,A.&($ – '0/0(F).++@% (.%A$*+.% Sampo 130. # ($C1.* 10/8+() &0+1.%)7)&.2$++. .3A)&$/'8 B&.A+@* '+.B ' B/.L$1) 1 %2. <+$/)7, B.120&5$/.'4 B. 50 B&.1,(3)2+@6 '30A/0*. E. 2'0% B&)7+$($% B&.2.1)/'8 .1+.G$(3.&+@* 1)'B0&').++@* $+$/)7. E. B&)7+$($%, &$7/)-)8 %0C1, 2$&)$+3$%) B. (.3.&@% A@/) 1.'3.20&+@, B&.2.1)/'8 12,6G$(3.&+@* 1)'B0&').++@* $+$/)7 (G$(3.& < – '.&3$, G$(3.& ? – 5.1@ × G.+@). ? ($-0'320 $,3A&01+.5. .AI0(3$ A@/ 2783 +$A.& 16 5)A&)1.2 (,(,&,7@ 5&,BB@ 'B0/.'3) X<P 200–250 '0/0(F)) &$7/)-+@6 ,-&0C10+)* ;.'')). PB@3 7$(/$1@2$/'8 2 2.'4%) D(./.5)-0'()6 3.-($6 (#$%$&'()* !MM#= – #$%$&'($8 .A/$'34, ?!MM (,(,&,7@ ) PPP ME< <5&.G)&%$ «P3A.&» – N$A$&1)+.->$/($&)8, X)/)$/ «E.2./C'($8 '0/0(F).++.-.B@3+$8 '3$+F)8» !" ?!MMPY – ?./5.5&$1'($8 .A/$'34, X !" «;.''.&5.» – #$&$3.2'($8 .A/$'34, N&$'+.1$&'()* !MM#= ) !E#X «!MN<#» – N&$'+.1$&'()* (&$*, ?.&.+0C'()* G)/)$/ ?!MMN – ?.&.+0C'($8 .A/$'34). E/.L$14 10/8+() 10 %2, B.23.&+.'34 3&06(&$3+$8. E.'02 B&.2.1)/'8 '08/(.* «Hege-95». ?@-/0+0+)0 50+.3)B)-0'()6, '&01.2@6 ) 50+.3)B-'&01.2@6 DGG0(3.2 B&.2.1)/'8 B. B&)2010++.%, 2@H0 $/5.&)3%,. E.1'-03@ B&.2.1)/)'4 +$ B0&'.+$/4+.% (.%B4J30&0 ' )'B./47.2$+)0% B$(03$ B&.5&$%% «Agros-2.09». -BGHDIC<CE ? :JFHK=B@?B N$( 2)1+. )7 3$A/. 1, B. 2'0% )7,-$2H)%'8 B&)7+$($% 1./8 50+.3)B$ 2 G.&%)&.2$+)) G0+.3)B$ 8&.2.* BH0+)F@ B&)%0&+. .1)+$(.2$ ) &$2+$ 10,0–15,5 %. E. 12,% B&)7+$($% ((./)-0'32. 70&0+ 2 (./.'0 ) 2@'.3$ &$'30+)8) 1./8 DGG0(3.2 27$)%.10*'32)8 «50+.3)B/'&01$» 7+$-)30/4+. +)C0, -0% 50+.3)B)-0'()0 ) .'.A0++. '&01.2@0 2(/$1@. E. D3)% B&)7+$($% .3A.& %.C03 A@34 DGG0(3)2+@% 2 /.($/4+@6 ,'/.2)86 B&) +$/)-)) (.+3&$'3+@6 B. G0+.3)B, &0(.%A)+$+3.2 2 5)A&)1+@6 B.B,/8F)86. PA$ D3) B&)7+$($ 2 (.+(&03+.% 'B0(3&0 G/J(3,- 144 !"#$%& '(%), 2010, *+, 14, - 1 )<JD?L< 1 SGG0(3@ (V) ) 1./8 2/)8+)8 (%) 50+.3)B$, '&01@ ) 27$)%.10*'32)8 «50+.3)B/'&01$» +$ G.&%)&.2$+)0 (./)-0'320++@6 B&)7+$(.2 , 8&.2.* %85(.* BH0+)F@ M'3.-+)( 2$&4)&.2$+)8 #.&3$ (<) "'/.2)8 '&01@ (?) ?7$)%.10*'32)0 (< × ?) PH)A($ (Z) 9$''$ 70&+$ ?@'.3$ O/)+$ 20&6+05. "&.C$* 70&+$, N./)-0'32. 70&0+ 2 (./.'0 ' (./.'$, 5 &$'30+)8, '% %0C1.,7/)8, '% 5/%2 V % V % V % V % V % 8,24 30,7 24,8 18,7 10,0 37,1 30,0 22,9 0,51 1,67 0,15 2,31 11,0 35,9 3,3 49,8 0,026 0,059 0,050 0,094 11,5 25,8 21,7 41,0 1,97 5,88 0,74 4,09 15,5 46,4 5,8 32,3 1,21 3,60 3,42 2,52 11,3 33,5 31,8 23,4 )&,JL)6 G$(3.&.2 '&01@ B. H($/0 +$B&8C0++.'3) '0/0(3)&,0%@6 B&)7+$(.2 1./C+@ A@34 .3+0'0+@ ( '&01+0+$B&8C0++@%. :&) B&)7+$($ (,&.C$* 70&+$, %$''$ 70&+$ ' (./.'$ ) 1/)+$ 20&6+05. %0C1.,7/)8) )%0J3 ',L0'320++@0 +$5&,7() DGG0(3.2 50+.3)B'&01.2.5. 27$)%.10*'32)8 (2 2–3 &$7$ B&02@H$JL)6 2(/$1 50+.3)B$) ) B.D3.%, .3+.'83'8 ( .3+.')30/4+. +$B&8C0++@% B&)7+$($%, .3A.& B. (.3.&@% %.C03 A@34 DGG0(3)2+@% /)H4 21./4 'B0F)$/4+.5. D(./.5)-0'(.5. 20(3.&$ (#J(.2 ) 1&., 2008). ? .3/)-)0 .3 BH0+)F@ 2 D('B0&)%0+3$6 ' 5)A&)1+.* (,(,&,7.* B. A./4H)+'32, B&)7+$(.2 1.'3.20&+@6 DGG0(3.2 50+.3)B-'&01.2@6 27$)%.10*'32)* .3%0-0+. +0 A@/.. E. B&)7+$($% «,&.C$* 70&+$» ) «(./)-0'32. 1+0* 1. 2@+.'$ +)30*» %0C'.&3.2@0 &$7/)-)8 2+.')/) 7+$-)30/4+. A./4H)* 2(/$1 2 G.&%)&.2$+)0 G0+.3)B$, +0C0/) 2'0 )7,-$2H)0'8 (./)-0'320++@0 B&)7+$() , )+A&01+.5. .AI0(3$ (3$A/. 2). 0+.3)B-'&01.2$8 '.'3$2/8JL$8 , (,(,&,7@ B. D3)% B&)7+$($% A@/$ %0+00 7+$-)%$, -0% %0C- '.&3.2$8. E. B&)7+$(, «,A.&.-+$8 2/$C+.'34 70&+$» '..3+.H0+)0 50+.3)B)-0'(.* ) 50+.3)B'&01.2.* '.'3$2/8JL0* A@/. $+$/.5)-+@% 3$(.2.%, , A./4H)+'32$ (./)-0'320++@6 B&)7+$(.2, +$A/J1$0%@6 , )+A&01+.* (,/43,&@. P1+$(. 2(/$1 27$)%.10*'32)8 «50+.3)B–'&01$» 2'0 C0 212.0 +)C0, -0% , BH0+)F@. :. 0'34 2 F0/.% B.1320&C1$03'8 .3%0-0++$8 &$+00 +$ C)2.3+@6 .AI0(3$6 (Parsons, 1959; Hull et al., 1963) 30+10+F)8 ( +0)7%0++. A./00 +)7(.%, ,&.2+J 2(/$1$ 50+.3)B-'&01.2@6 DGG0(3.2 2 G.&%)&.2$+)0 (./)-0'320++@6 B&)7+$(.2 , 5)A&)1.2 B. '&$2+0+)J ' 5.%.7)5.3+@%) G.&%$%). "-)3@2$8 2 7+$-)30/4+.* '30B0+) DB)50+03)-0'()* 6$&$(30& 27$)%.10*'32)8 «50+.3)B–'&01$», D3, 7$(.+.%0&+.'34 %.C+. .AI8'+)34 ' B.7)F)* 5)B.307@ «MN?OB@BC?ABF9:O: J<D<@F<». #,34 D3.* 5)B.307@ 7$(/J-$03'8 2 3.%, -3. 2 /JA.* (.+(&03+.* '.2.(,B+.'3) G$(3.&.2 '&01@ 50+.G.+1 B.B,/8F)) &0$/)7,03 .B&010/0++,J '.2.(,B+.'34 3&$+'(&)A)&,0%@6 B.'/01.2$30/4+.'30* ) &05,/83.&+@6 50+.2, .A0'- )<JD?L< 2 SGG0(3@ (V) ) 1./8 2/)8+)8 (%) 50+.3)B$, '&01@ ) 27$)%.10*'32)8 «50+.3)B/'&01$» +$ G.&%)&.2$+)0 (./)-0'320++@6 B&)7+$(.2 , (,(,&,7@ M'3.-+)( 2$&4)&.2$+)8 #.&3$ (<) "'/.2)8 '&01@ (?) ?7$)%.10*'32)0 (< × ?) PH)A($ (Z) "&.C$* 70&+$, 5/%2 V 0,2927 0,3320 0,2500 0,3887 % 23,2 26,3 19,8 30,7 "A.&.-+$8 2/$C+.'34 70&+$, % V % 0,6437 8,4 5,0904 66,0 1,1420 14,8 0,8342 10,8 N./)-0'32. 1+0* 1. 2@+.'$ +)30* V % 0,8533 16,4 3,0680 58,5 0,5309 10,1 0,7884 15,0 !"#$%& '(%), 2010, *+, 14, - 1 B0-)2$JL)6 .B3)%$/4+,J G)7)./.5)-0'(,J F0/.'3+.'34 F0+.7$ 2. 2&0%0+) ) B&.'3&$+'320 ' +$)%0+4H)%) D+0&503)-0'()%) 7$3&$3$%). :$(.0 '.'3.8+)0 ')'30%@ «50+.3)B–'&01$» %@ +$7@2$0% DB)50+03)-0'()% A$/$+'.%. E&) )+)F)$F)) )7%0+0+)8 /JA.5. )7 D/0%0+3.2 DB)50+.3)B$ ('.2.(,B+.'3) G$(3.&.2 2+0H+0* '&01@ )/) 50+03)-0'(.* '&01@) B&.)'6.1)3 )+1,(F)8 D('B&0'')) 1.B./+)30/4+@6 3&$+'(&)B3.+.2, .A0'B0-)2$JL)6 G.&%)&.2$+)0 +.2.5. DB)50+03)-0'(.5. A$/$+'$. E.'(./4(, 2 5030&.7)5.3+@6 5)A&)1$6 ,C0 &0$/)7.2$+$ -$'34 DB)50+03)-0'(.* )7%0+-)2.'3), 3. D3$ -$'34 +0 A,103 7$G)(')&.2$+$ 2 '3$3)'3)(0 V ($( %0&0 27$)%.10*'32)8 «50+.3)B–'&01$». Z'+., -3. +0 B. 2'0% (./)-0'320++@% B&)7+$($% D3$ 7$(.+.%0&+.'34 A,103 &$A.3$34 .1)+$(.2. ,'B0H+.. E&.82/0+)0 D3.* 30+10+F)) A,103 7$2)'034 ($( .3 +$B&$2/0+)8 ) H)&.3@ )7%0+-)2.'3) D(./.5)-0'(.5. G$(3.&$, 3$( ) .3 'B0F)G)() 50+.3)B)-0'(.* 2$&)$A0/4+.'3) )7,-$0%.5. .AI0(3$. !$B&)%0&, +)7(.0 7+$-0+)0 '3$3)'3)() V B. B&)7+$(, «2@'.3$ &$'30+)8» , 8&.2.* BH0+)F@ %.C03 .B&010/834'8 2@'.(.* 1./0* $11)3)2+.* 2$&)$A0/4+.'3) 2 /.(,'0 RhtB1. :&) '.&3$ (:,/$*(.2'($8 5, :,/$*(.2'($8 100 ) E&.6.&.2($) +0',3 2@'.(.D('B&0'')2+,J $//0/4 RhtB1b, .'3$/4+@0 – RhtB1a. :$()% .A&$7.%, B./02@0 D('B0&)%0+3@ +$ 5030&.7)5.3+.% $,3A&01+.% &$'3)30/4+.% .AI0(30 (5)A&)1@ (,(,&,7@) 2 '&$2+0+)) ' )+A&01+@% 5.%.7)5.3+@% .AI0(3.% B.1320&C1$J3 7$(.+.%0&+.'34 A./00 +)7(.5. ,&.2+8 2(/$1$ 50+.3)B-'&01.2@6 DGG0(3.2 2 G.&%)&.2$+)0 (./)-0'320++@6 B&)7+$(.2 , 5)A&)1.2 B. '&$2+0+)J ' 5.%.7)5.3+@%) G.&%$%). "-)3@2$8 2 7+$-)30/4+.* '30B0+) DB)50+03)-0'()* 6$&$(30& 27$)%.10*'32)8 «50+.3)B–'&01$», D3, 7$(.+.%0&+.'34 %.C+. .AI8'+)34 ' B.7)F)* 5)B.307@ «DB)50+03)-0'(.5. A$/$+'$». "?CBP<CHP< ?$2)/.2 !.M. !$,-+@0 .'+.2@ '0/0(F)) BH0+)F@ // :0.&03)-0'()0 .'+.2@ '0/0(F)) &$'30+)*. :. 2. 9.; T., 1935. #. 3–214. O&$5$2F02 ?.<., <20&48+.2$ <.X. 906$+)7%@ 27$)%.10*'32)8 50+.3)B–'&01$ ) 5.%0.'3$7 (./)-0'320++@6 B&)7+$(.2 &$'30+)* // 0+03)($. 1983$. :. 19. [ 11. #. 1806–1810. O&$5$2F02 ?.<., <20&48+.2$ <.X. E0&0.B&010/0+)0 145 50+03)-0'()6 G.&%,/ (./)-0'320++@6 B&)7+$(.2 2 &$7+@6 ,'/.2)86 '&01@ // 0+03)($. 1983A. :. 19. [ 11. #. 1811. O&$5$2F02 ?.<., W)/4(0 ;.<., ;0*30& >. . ) 1&. 0+03)($ B&)7+$(.2 B&.1,(3)2+.'3) 8&.2@6 BH0+)F 2 Y$B$1+.* #)A)&). !.2.')A)&'(, 1984. 230 '. N$3.6)+ <.?., N,7+0F.2$ :.!., P%0/48+-,( !.<. 9)-;!N – +.2@0 &05,/83.&@ $(3)2+.'3) 50+.2 , D,($&).3 // M+G.&%. 20'3+)( ?P )#. 2006. :. 10. [ 2. #. 241–272. N)/4-02'()* <.?., =.3@/02$ T.?. S(./.5)-0'($8 '0/0(F)8 &$'30+)*. 9)+'(: :D6+$/.5i8, 1997. 372 '. N.&.-()+ T.M. R3. 3$(.0 DB)50+03)($ // 0+03)($. 2006. :. 42. [ 9. #. 1156–1164. E/.6)+'()* !.<. 9$30%$3)-0'()0 %03.1@ 2 A)./.5)). 9.: M71-2. 9 ", 1978. 265 '. ;.()F()* E.X. >)./.5)-0'($8 '3$3)'3)($. 9)+'(: ?@HD*H. H(., 1973. 320 '. #J(.2 ?.?. 0+03)-0'(.0 .A.'+.2$+)0 2@A.&$ G.+.2 B&) .3A.&0 B. (./)-0'320++@% B&)7+$($% // E&.A/0%@ )+30+')G)($F)) ) D(./.5)7$F)) 70%/010/)8 ;.'')): 9$30&. +$,-.-B&$(3. (.+G. 14–15 )J+8 2006 5. ;$''203, 2006. #. 458–462. #J(.2 ?.?., Y$6$&.2 ?. ., N&)2.A.-0( ?. . ) 1&. 903.1 .3A.&$ 50+.3)B.2 8&.2.* %85(.* BH0+)F@ +$ 5.%0.$1$B3)2+.'34: 903.1. ,($7$+)8. #$%$&$, 2008. 18 '. R0'+.(.2 Q.?., E.-0B+8 !.?., >0&+0& <. ) 1&. S(./.5.-50+03)-0'($8 .&5$+)7$F)8 (./)-0'320++@6 B&)7+$(.2 &$'30+)* ) ($&3)&.2$+)0 /.(,'.2, .B&010/8JL)6 $5&.+.%)-0'() 2$C+@0 B&)7+$() , %85(.* BH0+)F@ // O.(/. ;<!. 2008. :. 418. [ 5. #. 693–696. Batson W. Mendel’s principles of heredity / Ed. W. Batson. Cambridge, 1909. Blanc G., Charcosset A., Mangin B. et al. Connected populations for detecting quantitative trait loci and testing for epistasis: an application in maize // Theor. Appl. Genet. 2006. V. 113. [ 2. ;. 206–224. Börner A., Schumann E., Fürste A. et al. Mapping of quantitative trait loci determining agronomic important characters in hexaploid wheat (Triticum aestivum L.) // Theor. Appl. Genet. 2002. V. 105. ;. 921–936. Brink R.A. A genetic change associated with the R locus in maize which is directed and potentially reversible // Genetics. 1956. V. 41. ;. 872–889. Brink R.A. Paramutation and chromosome organization // Quart. Rev. Biol. 1960. V. 35. ;. 120–137. Caligari P.D.S., Mather K. Genotype × environment interaction. III. Interactions in Drosophila melanogaster // Proc. R. Soc. Lond. 1975. B. 191. ;. 387–411. Connoly V., Jinks I.L. The genetical architecture of general and speciÞc environmental sensitivity // 146 Heredity. 1975. V. 35. P. 2. ;. 249–259. Chang S., Whitelaw E. Epigenetic germline inheritance // Curr. Opin. Genet. and Developm. 2004. V. 14. ;. 692–696. Dorn L.A., Schmitt J. Do plasticity genes exist or not? // Progr. Abstr. of 6th Congr. Eur. Soc. Evol. Biol. Arnehm, 24–28 Aug., 1997. Wageningen, 1997. Durrant A. The environmental induction of heritable changes in Linum // Heredity. 1962. V. 17. ;. 27–61. Durrant A. Environmental conditioning of ßax // Nature. 1958. V. 181. ;. 928–929. Evans G.M. Nuclear changes in ßax // Heredity. 1968. V. 23. ;. 25–28. Falconer D.S. Introduction to quantitative genetics. 2nd ed. N.Y.: Longman Sci. Techn., 1981. 438 p. Fisher R.A. The correlations between relatives on the supposition of Mendelian inheritance // Trans. Roy. Soc. Edinbourg, 1918. V. 52. ;. 399–433. Hill J. The environmental induction of heritable changes in Nicotiana rustica: Parental and selection lines // Genetics. 1967. V. 55. ;. 735–754. Hill J., Perkins J.M. The environmental induction of heritable changes in Nicotiana rustica: 0ffect of genotype-environment interactions // Genetics. 1969. V. 61. ;. 661–675. Hill J. Genotype–environment interactions – a challenge for plant breeding // J. Agric. Sci. 1975. V. 85. P. 3. ;. 477–498. Holland J.B. Epistasis and plant breeding // Plant Breed. Rev. 2001. V. 21. ;. 27–92. Hull P., Gove R.S., Slen S.B., Crawford R.D. A comparison of the interaction, with two types of environment, of pure strains or strain crosses of poultry // Genet. Res. 1963. V. 4. ;. 370–381. Jinks J.L., Jayasekara N.E.M., Boughey H. Joint selection for both extremes of mean performance and of sensitivity to a macroenvironmental variable. I. Single seed descent // Heredity. 1977. V. 39. P. 3. ;. 345–355. Jinks J.L., Pooni H.S. Determination of the environmental sensitivity of selection lines of Nicotiana rustica by the selection environment // Heredity. 1982. V. 49. P. 3. ;. 291–294. Juengei T.E., Son S., Stowe K.A., Simms E.L. Epistasis !"#$%& '(%), 2010, *+, 14, - 1 and genotype-environment interaction for quantitative trait loci affecting ßowering time in Arabidopsis thaliana // Genetica. 2005. V. 123. [ 1/2. ;. 87–105. Landry C.R., Oh J., Hartl D.L., Cavalieri D. Genome-wide scan reveals that genetic variation for transcriptional plasticity in yeast is biased towards multi-copy and dispensable genes // Gene. 2006. V. 366. ;. 343–351. Li Y., Alvarez O.A., Gutteling F.W. et al. Mapping determinants of gene expression plasticity by genetical genomics in C. elegans // PLOS Genet. 2006. V. 2. e222. Malmberg R.L., Held S., Waits A., Mauricio R. Epistasis for Þtness-related quantitative traits in Arabidopsis thaliana grown in the Þeld and the greenhouse // Genetics. 2006. V. 171. [ 4. ;. 2013–2027. Parsons P.A. Genotypic-environmental interactions for various temperatures in Drosophila melanogaster // Genetics. 1959. V. 44. [ 6. ;. 1325–1333. Paterson A.H., Damon S., Hewitt J.D. et al. Mendelian factors ungerlying quantitative traits in tomato: Comparison across species, generations, and environments // Genetics. 1991. V. 127. ;. 181–197. Plomin R., DeFries J.C., Loehlin J.C. Genotypic-environmental interactions and correlation in the analysis of human behavior // Psychol. Bull. 1977. V. 84. [ 2. ;. 309–322. Rutter M., MofÞtt T.E., Caspi A. Gene-environment interplay and psychopathology: multiple varieties but real effects // J. Child Psychol. Psychiatry. 2006. V. 47. [ 3/4. ;. 226–261. Smith E.N., Kruglyak L. Gene-environment interaction in yeast gene expression // PLOS Biol. 2008. V. 6. [ 4. e83. Ungerer M.C., Halldorsdottir S.S., Purugganon M.D., Mackay T.F. Genotype-environmental interactions at quantitative trait loci affecting inßorescence development in Arabidopsis thaliana // Genetics. 2003. V. 165. ;. 353–365. Wade M.J., Winther R.G., Agrawal A.F., Goodnight C.J. Alternative deÞnitions of epistasis: dependence and interaction // Trends Ecol. Evol. 201. V. 16. [ 9. ;. 498–504. 147 !"#$%& '(%), 2010, *+, 14, - 1 THE CONTRIBUTION OF GENOTYPE-ENVIRONMENTAL EFFECTS TO THE FORMATION OF QUANTITATIVE TRAITS IN INBRED AND OUTBRED PLANTS V.V. Syukov, E.V. Madyakin, D.V. Kochetkov Samara ScientiÞc Research Institute of Agriculture, Bezenchuk, Russia, 0-mail: [email protected] Summary The assumption about the epigenetic nature of the genotype–environment interaction is made and proven. When genetic variances in descendants of various ages are analyzed, this interaction manifests itself as epistasis. Study of inbred (common spring wheat) and outbred (hybrid maize) forms shows that ther contribution of genotype–environmental factors to quantitative traits in hybrids is less than in homozygous forms. With regard to the substantially epigenetic nature of the genotype–environment interaction, this trend can be explained in terms of the epigenetic balance hypothesis. Key words: «genotype–environment» interaction, epigenetical variability, epistasis, epigenetical balance, wheat, corn.