, - 03.01.03 – 2011 ! # " .$. % # : ! , $ % : & ! '(" )( , , / & ' ( " ) ! "-!" # !. 17 + # "# 2011 #. $. . , ____ - , * .501.001.76 # # , - .$. % # # # ' , . .! # " , # ,% : 119992, # - , , /&, 536. ! " /& .$. % # , & *"+ "# % ): )* # "# «__» # " 2011 , 0 2 . . / (.&! ) '. 0 1 +2 , , " # 1 . # # # ,# , # " , * , * , # ,# , 1 +* " + + 0 ", + , 2 + + # " * * # # (1.) – " 3!, ' + # +* # " , , + , 3!/1 # 1- # , Hordeivirus). ! , "+ ( # 0 ( +* # # , ( , # 3! 3!/1 $-$ 3!/3 1%$ Pomovirus) , 3!/3 ), 1. (Morozov & Solovyev, 2003, Verchot-Lubicz et al., 2010). $ * 3!/2 #, # + (3!/): 4 # (Harries et al., 2010). # # - 4 , ' 2 5 . + # . / – , #, , + ,# # +* " (3!), ' . , + , # , 3!/1 , 1. 3!/3 (Morozov & Solovyev, 2003, Verchot-Lubicz et al., 2010), , . 3 ' , 3!/3 , 1., # # (Schepetilnikov et al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«% # /& , /& # " ! # , .$. % # # ' , –2011». &1 "(!5"" 1 # . # . 4 - . &(.& ! !1 .2 . # », « , " 6! " «$ », «2 , », « ' », «$ », «- ». !1 .2 $, # 3!/1 $-$ 3 , # , * , " 3!/1 +* Nicotiana benthamiana $-$ , # " " # , 15–18 , $ * # + +* # . 1 , # GFP-3!/1, 24 4 # ' ' 3 ( benthamiana , . 1 , !). 1 GFP-3!/1 " * , 3- ' ). # *"+ # # , " 3( , , , # ,# '# , # ,# (Zamyatnin et al., 2004). $ GFP # , # 35S- (green fluorescent protein, GFP), +* + , " ' GFP-3!/1, # # ( , "'. 1. % ' , GFP-3!/1 Tal. – # # , , ' , ' ' 0 # #. 5 , ' ). GFP-3!/1 $-$ , " N. benthamiana. # *"+ " +* # . – GFP-3!/1. – , #-# # # # YFP, GFP-3!/1 #3 YFP-MAP4. . : GFP , YFP – , ' – . 7 YFP # # ' . , ' + , " # , ( , – ) + , ( ). : – 20 # #, – 5 # #, – 10 # 24 , , N. 8 " + #-# -4 # *"+ # " ", YFP) , , # ", YFP # ' 1$). , YFP "+ # -4 $-$ " ( # $3 # # 3 YFP 3- , +* # MAP4 " YFP-Tal, + , # # # 4 , " , ( # YFP-MAP4 . 1/). ( , 3. $, # ' # (YFP-Tal) (Kost et al., 1998). . GFP-3!/1 , - #- # # (yellow fluorescent protein, # GFP-3!/1 GFP-3!/1 # 0 # 0 (YFP-MAP4) (Marc et al., 1998). 1 GFP # +* # # * # , ## # +* 3 3- N. benthamiana. $ ' - , # GFP + , 3!/1 3 3!, # , ( . ) (Heinlein et al., 1995, Boyko et al., 2000a, 2000b, 2007, Ashby et al., 2006, Sambade et al., 2008). 1 3! $3 , # # # ' , # 4 1., ,# ,# # ((1 ) " + - " (11-) (Taliansky et al., 2008, Epel, 2009, Heinlein & Epel, 2004). 1 , 1. , #, 1- # 3!/1 $-$ " , 3! $3 " 3! $3 , +* ' # # " #. 3 ' 4 , , # 3 (Kahn et al., 1998, 3! $3 , (Boyko et , # , # « 3, 3! $3 , , # ' , # # , # in vitro (McLean et al., 1995, Ashby et al., 2006, Ferralli et al., 2006, Sambade et al., 2008). " # 3!/1 $-$ #», ' ,# ' " # , % , # # ' , . 2 , - # « " , # # 3!/2 , 3!/3 " ", N. benthamiana " 4 # 6 3!/3 3!/1 $-$ # " # 4 # , , + 1 in vitro ( . . ). 3!/1 $-$ +* # , 3!/1 $-$ , 3!/2 # : » 3! $3 ", , $3 3 (Heinlein & Epel, " # +* " # ' al., 2000b). . ' , " * # *"+ # Boyko et al., 2000c). 1 # # , # 2004, Verchot-Lubicz et al., 2010). - ' (Taliansky et al., 2008). 3 # + " 3, , GFP-3!/1, 1 3!/2 – , # 4 , # +* 4 # , # 3!/2 3!/3 # " , " 1., GFP-3!/1 (Zamyatnin et al., 2004). 2 # " + # 0 3!/1 +* , * " 3!/2/3!/3 3!/1 " 10:10:1, 0 , .$ - , # 35S- 0 3!/1:3!/2:3!/3 +* , # 3!/2/3!/3 4 ' 10:1 (Zhou & Jackson, 1996). 0 ", 3!/2/3!/3, + , , # , , # # $ - . 4 # # , , 4 , , 1.) , , 1. # " GFP-3!/1 4 #, ' ' , # . 3!/3 # , GFP-3!/1 +* ( 3!/2 (Zhou & Jackson, 1996). $ 4 # " + +* 3!/3 " # # + # 3!/2 3!/3 (Zamyatnin et al., 2004). # 4 , + , 3!/1:3!/2:3!/3 100:10:1 (Jackson et al., 2009). . ' # , 0 4 ,# . 1 35S- # , # # 35S# # + 3!/2, # $-$ , , # ' * +, , * # , +* 3!/3 (Jackson et al., 2009, Verchot-Lubicz et al., 2010). 3 +* pLH-35S:GFP-3!/1-NOS:3!/2/3!/3 ( "'. 2. - # , , # ,, 0 3!/1, 3!/2 3!/3, , ' # # # # + + , # , 4 $-$ +* , , , - " # (Sanders et al., 1987). $ 4 , - # +* – 3!/3, '* , GFP-3!/1 , , , # 4 3!/2 # # # 7 3! . 2). pLH-35S:GFP-3!/1-NOS:3!/2/3!/3. # – # ./ , +* # . " " , #, # GFP, $ pLH-35S:GFP-3!/1-NOS:3!/2/3!/3 # *"+ " " . % , , " + +* # *"+ GFP-3!/1 # ( . . .). . . . ' , ). " + , 3!/3 ( . , # # . 1 , ! "0 ' "0 , + # # '* ' , . , " # * " # # +* ( ' # - ' , + " " , #, . . . 3, " # 3. GFP-3!/1 $-$ " 3, ' GFP-3!/1" . . . ,# , 3!/2 GFP-3!/1 ' " 3, . . . GFP # 0,3–0,8 # #, + GFP ' , 4 " " . . . 3 ). 3!, $). " GFP ( , ,# N. benthamiana , + ( ,# , # * # . 3/). # + , , # +( . 3/). "'. 3. % , GFP-3!/1 , 4 pLH35S:GFP-3!/1-NOS:3!/2/3!/3, " " N. benthamiana. , ' # *"+ " , +* # . –2 . . . – 3 . . . – 4 . . . 1 , ' , ( , , ) , ( ). 0 : , – 20 # #, – 5 # #. ( + 0 # # , " (Johnston et al., 1998) Harries et al., 2009a). 1 – + 3-, # # 3 1998, Harries et al., 2009a). $ ' " (Vogel et al., 2007, # , # ' , # + #, # N. benthamiana, # # , '* ", 8 # (Johnston et al., ' # " # 4 ' pLH-35S:GFP-3!/1-NOS:3!/2/3!/3, , . . . ( ). 3 # , #, #, 3!/1, 4 # # 3!/2 # . # # 4 4 " + , " , " 3!/1- 1., # # '* ' , # # # # 4 , " 4 3 * # , 3! # 2008). $ ' # " " # $3 . +* " , , ' $3 , 3! (Boyko et al., 2007, Sambade et al., ", 3! $3 , " , ' , - , 1- # 3 * , # 3!/3. ' # " , # "0 " , " 3!/2 / 1 " + , , " ' , 3!/2/3!/3 GFP-3!/1 , , (Zamyatnin et al., 2004). 1 4 , # - + , – , +* 3!/3, ", 3!/2/3!/3 4 GFP-3!/1 (Zamyatnin et al., 2004), , + ' # 3. 1 4 3 , ' 1- # 1. (Sambade et al., 2008). $ 3 " , ' ". 3 # '* -3!/13 # ", 4 , 3 ,# , - , +* * 3-, 2, 3 6 " , 3 (Harries et al., 2009a, , # #, ", + # . , # # .3 , 3! (Curin et al., 2007), " 3! , 4 , # ,# ' # . # 0 . , + # Martiniere et al., 2009). $ ,# ' , " " 0 ' ' # , , # , , ' ", 3, + # 3!/1 $-$ , , # ' , , 1.. $ 3!/1 '* , , # # 3! # , ' , 3 – 4 , 3! $3 , 3-, 3 # 3! (Vogel et al., 2007). 1 "0 # , + 9 # " 3 # # , ' , ' # # (Vogel et al., 2007). 1 # # 4 3!/1 " , # #, + 1., # ' 0 # 0 '* , # ' # , , 1. (Ding, 1998, Kim & Zambryski, 2005, Maule, 2008, Faulkner & Maule, 2011, Tilsner et al., 2011). ! " #. ( 0 , +* + 0 , pLH-35S:GFP-3!/1-NOS:3!/2/3!/3 . . ., " 3!/3 1. ' 4 ' (Garcia-Mata et al., 2002). # 3!/2 8 " ' " ! # , ( ' , GFP-3!/1, , + - ' . 4 ), , + 3!/1 1.. "'. 4. % , GFP-3!/1, 4 NOS:3!/2/3!/3, 1. 4 " N. benthamiana # *"+ " , +* # .1 , , GFP-3!/1 , , 0 # , – , ' , ' – . 7 # # ' , ' ., + GFP , . – , GFP-3!/1 ,# , , ' * # – , ' , ' – , + 0 + ,# ## , 0 GFP-3!/1. 0 : 15 # ! , , + " #, , # #. - . .- , pLH-35S:GFP-3!/1' # " ,. – GFP , , , +* GFP , , , ' # , ' – #. , # , 1. # . . . # GFP-3!/1 1.. . 4 " N. benthamiana, # pLH-35S:GFP-3!/1-NOS:3!/2/3!/3, , " # GFP-3!/1 ,# ,. ! + ' + 10 ,# , , , # ,# ## , . 4!). ( ( " 3!/1 1.. 1 # 3! , " + 1.. $ # , # , al., 2007). 3! $3 ,# #, , 3! $3 1- # 1. , (Hofmann et # , " 11- 1. (Wolf et al., 1989, Hofmann et al., 2007, Waigmann & Heinlein, 2007). ,# 11- 1. # # # ' " , # " , # & Zambryski, 2001, Taliansky et al., 2008, Epel, 2009). % 1- 1. (Crawford , 1. , 3! (Erhardt et al., 2000, Haupt et al., 2005, Lim et al., 2009, Tilsner et al., 2010). - 3!/3 , 3!/2, , , 4 # , # 1. , 3!/1, , 3!/2 ,# ' , " ,# 11- 1. (Morozov & Solovyev, 2003, Jackson et al., 2009). - ' 4 3!/1 $-$ 1 " 3!/1 3 0 " , 3!/1 . # *"+ " 3 # ($ -) - 4 # # . +* TRV-TubAS, 4 * (Ratcliff et al., 2001), 1 N. benthamiana. - $ - N. benthamiana (Gillespie et al., 2002). " N. benthamiana " . # - . * # TRV-TubAS 8 , , ' # , , 20 , , " , ", # " $ - GFP-MAP4 , 35S:GFP-3!/1-NOS:3!/2/3!/3 # , " # TRV-TubAS $ - # . $ , , , " 3( GFP-MAP4 ( " 3 GFP-MAP4 " , * # . 5 ). # # #, ", , , ( " " " TRV- # , " # *"+ + ' $ - , " "# TRV-TubAS " " $ , " . 5$). $ , " ' , . 5$). $ - , 4 , 20 + pLH" 11 # TRV- , TubAS ' ( , ## , - $ # # MAP4 # " " # # GFP", #. ( # ", + $ - # "0 (Caplan & Dinesh-Kumar, 2006). 3 , ", TRV 4 , ' #, $ - , α, # TRV-TubAS, * , " 1. # $ -, 4 , 4 , # . 5!). $# ( 3 N. benthamiana " , , . . . ! # , 20 " , GFP-3!/1 3!/1 , # , . 5/). + PDS, CTR1 " # 80% (Liu et al., 2002). 1 , , ' , CTR2 # , # 4 # $ # " (Caplan & Dinesh-Kumar, 2006). "'. 5. % MAP4 , -# 3 GFPGFP-3!/1, 4 pLH-35S:GFP-3!/1NOS:3!/2/3!/3, 20$ " N. benthamiana. – , ' N. benthamiana " # TRV2-TubAS ( ) TRV2 , ( ). – ## # # : ' 1 – 4 , " , " TRV2 , , ' 2 – 4 , " , " TRV2-TubAS. , # # # , . . – , # 3 GFP-MAP4 " , " TRV2 , ( ) TRV2-TubAS ( ). – , GFP3!/1 " , " TRV2 , ( ) TRV2-TubAS ( ). : , ' # *"+ " , +* # . 1 , ' , . 0 : 10 # #. 3 # , #, # " 4 #, 0 , " , +* # # " 0 # .1 ' 3!/1 # ' 4 1., # ' ", # 3 3!/1. 12 + 2 # 3!/1 1. , # $ " 0 4 , # + 3!/1 $-$ " , , #– , # 3!/1 # 3. $ GFP- 3 # ", " , ,4 +* " 4. # benthamiana . . . $ 4 +* , ,# , , 0 3!/1 ' 3 GFP-3!/1 +* # " (1 # . 6 ), +* ' , " 0 3 ,# . 6!). 3 ( " #, " 3!/1 , ( ,4 1., + # + " , # . $ 4 # , 0 " #, , " , GFP-MAP4, 3. $ , " pLH-GFP-MAP4. " # " " N. pLH-35S:GFP-3!/1-NOS:3!/2/3!/3 , + # 1. # ,# , #, , # #, ' #. "'. 6. % , 4 3!/1-NOS:3!/2/3!/3, 1 ( # #. 1 " # # # # (Barton 3 & , +* (1 # Overall, 1- # # 2010), 3, # # # (Boevink et al., 1998), 3! , # GFP-3!/1, pLH-35S:GFP-# 3 GFP-MAP4 N. benthamiana # . . . , ' # *"+ " , +* # . , ' , . – , GFP-MAP4 " ( ) ( ) # . – , GFP-3!/1 " ( ) ) # . 0 : 20 * + 1.. - # , 4 # " "+ " , ' , # 3! " " (Sambade et al., 2008). 1 13 1- # 3 + " , " # (1 -, # ' + + # " $-$ . , (1 . 1 3!/2 " , +* # # " 3!/1 , # # # (1 +* 3!/1 $-$ ' ", " # *"+ # , 3!/3 (1 , # ' (1 3 GFP-3!/1 " , , 0 3!/2 , (1 # 3!/3, / # (Morozov & Solovyev, 2003, Verchot-Lubicz et al., 2010). , # 3!/1 3 ' , # ," # ' # 1. . 3!/1, ", # # ( . # ,# # HEL # # , # # ' # / , 3!/1. 3 , # , # 0 " # * 0 # # 0 # 3!/1-C2 3!/2 ' " , , pLH-35S:GFP-3!/1", NOS:3!/2/3!/3, " , # # 3!/3 (Zamyatnin et al., 2004). 3!/1 , # * N. benthaminana. % 3!/1- # , + + # 3!/1. ( 3!/1 # # .7//.7, # , GSGKT 37 (Feig, 1999, Nagy et al., 2009). # + 3!/1- # . 7). # # 37//37 # " , NTP , NTP , # ( . 7). $ # ( , / 34 37–84 4 , # C3 3!/1-dN , # NTP , 3!/1-Mlu1 (Zamyatnin et al., # # 188 # 3!/1-GKN # 3!/1-Mlu2 # 3, # 37–85, # # , # # 7). # 2004), , " " # *"+ " . . . ! ' # , " " 3 1., # (3 3!/1-dN, 4 # - ' # dN, Mlu1, GKN, C2 , " + , # # # , C3 # " # # . 8). 3 , . 1, # . . . ' 3( , # ,4 +* , , . 8 ). 3, 14 4 . . . ' " + , " "'. 7. - # " " ' NTP/ ' " , ' , ' # # , ' # , # , # " # # " "'. 8. % , # " N. benthamiana +* # . , # # ' – 3 . . . – , # Mlu2. # -2. 7 , # 4 # , # ( # 3!/1-Mlu1, ' + ; 3, . . . # , # " 3!/1# – +* 2 . . ., Mlu1. , . 8!). 3 # "0 + 15 8 ). $ , 0" 3 . . . ( , + . . " , " . . . , , . . . , +* # 3!/1 $-$ . # 0 I – VI. ) # «+» , , , , GFP-3!/1, 4 # *"+ pLH-35S:GFP-3!/1# -NOS:3!/2/3!/3, ' # *"+ " , , . , , 4 . . . – , # dN. – , # , # HEL. , # GKN. -3. 0 : 10 # #. " , . ) # # # " # # .$ # GKN, C2 -3: , ' – #"+ ./ , " " # 3!/1-GKN # " ; # # 1. ( . . . 3!/1--2: ' 3, , " # , # , * # ", # 3 # , 3!/1--3 ' # - "# 3, !1 "5!1. % , " . "( # 2 , % , 2 . . . . , 1., 4 . . . 3 . . . +++ +++ +++ dN + + + Mlu1 + + + Mlu2 - - - HEL - - - GKN + ++ + C2 ++ +++ ++ C3 + ++ + . , , # – 3!/1-Mlu2 " ,# 3, # # (3 , 3!/1-HEL – + , # " 3, + . . ., # ' " "+ ' , . 8$, /). - . 1, . 89). 3!/1 $-$ 3, 3!/1 , # " ! , , . + , ' 3, " - ' ,# . 2 " + . . . , ' , " 3!/1--3 . 8 ). * 3!/1 - "0 . . . " . . . ,# , 3, . . . ' 3. , ( # . . ., , # 3, # , . . . , . 8.). ", , # ' # , # 3!/1-Mlu2 3!/1-HEL ( " #, , # # * ). 3 # , #, 3!/1 3 , " 0 1. .! 3 ,# ' " " + ' ", 86 -" + + # , , # 3!/1 184 # 3!/1. ( " # +* 16 # N' + + (Makarov et al., 2009). 1 3!/1 , , # ), , , # ,4 # , , "# – ' ", " ( # # ' " 3. 2 " ,# # 3!/1 $-$ . 3 +* # (1 , # " , , + 3!/2 + 1. 3!/1 , . " ' , - 3!/2, ' # # (1 , . 3!/1, , , +* . 3 , +* - + # #, 3 , +* " . 3 # # , 4 ", # 1, . * 3 , , GFP-3!/1 $-$ , 4 # # (Wright et al., 2010). $ 4 3 , +* # # ' , ' , ' * ' # ' 5 ' ", # , , 0 (Wright et al., 2010). 2 17 , , , # , N- " #4 , ' #, N- # # Wright et al. (2010) 4 # . 3 . ! 3, # , , , " # 4 , # 3!/1, # , , # , , #, # # , 3!/1 $-$ 3!/1 ' " , 3!/1 , " " 3!/1 # ' , $-$ # # "+ , 3, # # ' , $ ' , "+ # , 0 # 3. +* 3!/3 # , # (1 , " (Verchot-Lubicz et al., 2010). & # # ; # # 3!/1 3!/1, 3!/2, # 3!/1 # # 3!/1, # # , # " , # " " . ) # # $ ,# ' , " 3!/1 , # , +* + # , # 3 . # 4 , " # " 4 , # ' # # ## ' 5 # , ' # ' # # " #, '* # . . ($ 4 1999). 2 +* # # 3!/ $ , # # # , 3!/ 1%$ # (Solovyev et al., , # " , " " , * ). ! # + ,, " , #, , + " + # # # # # ( 3!/3 1%$ . $ ' # , . 9). ", # 4 (Gorshkova et " " # # 3!/3 + , . 9). $ ( , # # ' al., 2003), ' " # " # # , , # ' # * # ' ' " ( #. # " # ' " " "0 # (Gorshkova et al., 2003), , # # (Gorshkova et al., 2003). 1 # # +# # 3!/3 1%$ ", (Zhou & Jackson, , ' # , # 1996, Jackson et al., 2009) # ## 3!/3 1%$ 3!/3 4 # , 95°- , ". - , # # 3!/3 # # #, 4 # , # '' # ' 0 21 3 # N. benthamiana , 684, , + * , 3!/3 1%$ , 4 4 # ), 3!/3 1%$ , ' # # , # " # 3!/3 1%$ , 3!/3 (Gorshkova et al., 2003). 8 , 3!/3, 4 , GFP , " # , # , +* # , . , # ' # ' 0 4 " , 4 # " , ", " (Wright et al., 2010), 2 " , . 2 3!/3 $-$ , # Cowan et al (2002), # + + # , # ' # 18 " ' #, 3!/3 1%$ . $ + " 3!/2, + , , # ' " , # # " 3!/3 (Cowan et al., 2002, # 3!/3 Lim et al., 2008) (Solovyev et al., 2000, Zamyatnin et al., 2002, Tilsner et al., 2010). "'. # # . 9. 3!/3 1%$ N. benthamiana, # # " # 684. , " # *"+ # " 3!/3. $ 4 ## ", ", " benthamiana, 1%$ , . , , . . , # , N. 3!/3 +* , , ( . . .). # # «<<» , + ' # 3!/3 # # .1 ' # ($ ) , # «<» , # # ($ !), ! 3!/3 " # 3!/2 3!/3, '' , , , + # # , – ", ' # + # # #, . 9). 3!/3 1%$ , # # ' # # # # # # , ' 1.: # # ", # +* # 3!/3 , al., 2008, Tilsner et al., 2010). 4 , # , , , ( # 4 -- # # " , # 3!/2 , +* # # # # " , # # , '' . , # ' " +* # , " (Lee et al., 2010). ! # # , , 3!/3 " (Lee et al., 2010). $ ,# ' , ' . ' (1 # # , , $ ! 4 3!/2 * # # (1 + +* , # , # ' # # " # # # # , (Solovyev et al., 2000, Schepetilnikov et , , + , + +* 3!/3 1%$ . . 3!/3 1%$ , # RSTD 19 , 4 3!/3-# 71, # " YQDLN "+ # -- # # , " ## ", " , . ( , # # 4 # # . 11), # # ' ' # , " # # , # ' " , ' 3!/3 # #. & , " N" ", " " , ./ # 0 ", #. . # 0 , ,# # 3!/3 , " , 1%$ # # # " # . 3!/3 " # .. # 3!/3-# 71 0 +* # 3!/3 , ' " , 3!/3-# 62 # , # # 20 . . / # "'. 11. 2 , # # # # 71 # 62. ! , N. benthamiana # *"+ " 2 . . . 10%- # 1 / ", 3!/3. " #, * # . # . . # " , 3!/3 1%$ - # " " # - " + YQDLN " , , . 11). 3 ( " ' ", "'. 10. #, # GFP-3!/3-# 71 # # (Gorshkova et al., 2003). GFP-3!/3-# 71 # (Gorshkova , GFP-3!/3-# 62 # " 3!/3 1%$ # # # *"+ " # ' 0 117 # (Solovyev et al., 2000, # . . . et al., 2003). ! , #, +* # 4 GFP, . # # # , 3!/3-# 62 , # # . 10). / Schepetilnikov et al., 2008) ( # " # RSTD YQDLN. -- # , # ' , # GFP, 4 , # . # 3!/3 1%$ " , ## # # " # " " , # # , , GFP-3!/3-# 71 # " , 4 # 3!/3-# 62 , , +* , 4 # .! ' , ' . 12/), # # 3!/3 Zamyatnin et al., 2002). , # , -4 GFP " , # (1 ( GFP , " # , #-# , " , ' + + # , " DsRed-3!/3-# 71 DsRed # 1 , + # " " (1 # / # # # " # " , /. 3 , 4 # 71 # ' # # ( # , # , ,4 DsRed-3!/3-# 71, + . 13). 3 " , , , DsRed-3!/3- GFP-3!/3-# 71 GFP-3!/3-# 71, " #. ( 4 +* DsRed-3!/3-# 71, # # # # , 0 100 . , " , " # . 13). , # ' 0 , # # ( " # GFP-3!/3-# 71 5 GFP-3!/3-# 71, 95°-, " (Gross et al., 2000). 2 " , 95°- # " , - /), # " - N. benthamiana ## + +* # # ## , , , 4 # " # " . $ # # , " .3 DsRed-3!/3-# 71, 4 " m-GFP5-ER " # , DsRed " # (1 , # # # *"+ # # . 12$–.). ( (Baird et al., 2000, Wall et al., 2000). ( , # # # # # DsRed # 1 " # # 3!/3-# 71 # , DsRed (Solovyev et al., 2000, (Zamyatnin et al., 2002), # " , ' DsRed-3!/3 (Zamyatnin et al., 2002), ' ", DsRed- , DsRed-3!/3-# 71, , , DsRed-3!/3-# 71 # m-GFP5-ER, , # " ( DsRed-3!/3-# 71 ( +* # , " , , # # # "0 , ). ! , 4 # *"+ N. benthamiana * , # DsRed (Matz et al., 1999), " ,# , . 12 , !). $ (Schepetilnikov et al., 2008, , + GFP-3!/3-# 62 # , - , #, GFP-3!/3, # + # 3!/3 1%$ ", # # # 21 # " # # . ( , +* # , '' '* 3!/2 # # " # , , 3!/3 $ !, # , 0 (Lee et al., 2010). "'. 12. % , GFP-3!/3-# 71, GFP-3!/3# 62 DsRed-3!/3-# 71, 4 N. benthamiana # *"+ # " . , ' # *"+ " , +* # .1 , ' , . – , GFP-3!/3-# 71. – , GFP-3!/3-# 62. , , - -4 DsRed-3!/3-# 71 # (1 m-GFP5-ER. – , m-GFP5ER. – , DsRed-3!/3-# 71 # # " . – ' , ' . 0 : – 20 # #, , , – 5 # #. "'. 13. 2 GFP-3!/3-# " N. , ", 3!/3. * # " " * # # 1 * , * , 3!/3 . 2 , # 3!/3 3!/3 0 # #, " " ( , ). #, # # " , 4 " 4 " # # " .. # # / , YQDLN # # # # " " # ' + + + , " 12%- # 1 . (Schepetilnikov et al., 2008, Lim et al., – , # , " # 3!/3, # # , # 4 , # # 2009, Tilsner et al., 2010), # , # 71 DsRed-3!/3-# 71. ! benthamiana # *"+ ## # 2 . . # # " " . 1 95°- ( ) " " . 6 # , . . -- , # , .( # ' YQDLN 22 3!/3 # #, ' " , " " # *"+ # # # 3!/3 " ' (Schepetilnikov et al., 2005). " # # # # ", # # # # ' " DsRed. - , 1 , (1 , " , , , + , # 3!/3, # " (1 (Zamyatnin et al., 2002; Verchot-Lubicz et al., 2010). 3!/3 $ ! '' + # # . ! + " Yop1 (Lee et al., 2010). (Rtn1) + + " # , 4 , * * # - ' (1 , , # ' # # Rtn1 # # , DP1/Yop1, , +* # # # (1 , '' , (Voeltz et al., 2006, Voeltz & Prinz, 2007, English et al., 2009, Nziengui & Schoefs, 2009, Shibata et al., 2009). DP1/Yop1 # + Rtn1, # # , # (RHD, reticulon homology domain), , # # # # # , (1 (Shibata et al., 2008, English et al., 2009). 3 Rtn1 # # DP1/Yop1 " # # " Rtn1 , # et al., 2011). ! ' " 3!/3 ' ,# 3!/3 ## # ' # " # , , (1 ,# ' , , 1., 1. (Tilsner et al., 2011). 23 + # "0 , (Tilsner 1., , 3!/3 $ ,# " ,# # ' " , , (Lim et al., 2009, Tilsner et al., 2010). 3 # , (1 . " # " ,# # # # $-$ , ' ", # # , +* 3!/3 # # , , # , ' DP1/Yop1, , # # , (Morozov & Solovyev, 2003). ,, # " (Nziengui & Schoefs, 2009); , # # 3!/3 , RHD " 3!/3 RHD , . . # , # # " ,# , #, , 3!/1 1. # $ + 3!/3 -2 II- , # , # # , , (1 , , 2005). # # /37 , # # 3 , # COPII- , ", COPII- # # - +* Sar1[T34N], , 3!/3 1%$ # COPII- , :- , (:$ ) (Schepetilnikov et al., 2005, 2008). $ , , , COPII- , + # 3!/3-# 71 , # *"+ Sar1[T34N] + 3!/3-# 62. $ ", , (1 ST-YFP (Schepetilnikov et al., 2005), / " ' ( /) COPII-, # # # # # (Saint-Jore et al., 2002). $ # , +* # # (Mancias & Goldberg, 3!/3 , #: # , ST-YFP / ( # . 14 ), # Sar1[T34N] , " -4 , ST-YFP # # ST-YFP # COPII-, # (1 (Schepetilnikov et al., 2005, 2008, . 14!). "'. 14. $ 4 Sar1[T34N] , + GFP-3!/3-# 62, GFP-3!/3-# 71 DsRed-3!/3-# 71. ! 4 " N. benthamiana # *"+ # . , ' # *"+ " , +* # # ' , . – , ST-YFP. – , ST-YFP, -4 # Sar1[T34N]. – , GFP-3!/3-# 62, -4 # Sar1[T34N]. – , GFP-3!/3-# 71, 4 # Sar1[T34N]. – , DsRed-3!/3-# 71, -4 # Sar1[T34N]. 0 : 20 # #. , # , # GFP-3!/3-# 71 " , +* , (Solovyev et al., 2000, Schepetilnikov et al., 2008, , # , ,# + . 12 , !). ! +# GFP-3!/3-# 62 ( 24 . 14$), " COPII- # *"+ Sar1[T34N] , , + GFP-3!/3-# 62 , # , # # GFP- 3!/3-# 71 -4 . 14/). 3 ( # # , #, # , GFP-3!/3-# 71, # # COPII-, COPII- , ' , # ' ", " # 3!/3 # # . . , + Sar1[T34N]. ! -4 . 14.). 3 ( # , #, # ' # # # 1 1- # 1. , », + +* , , et al., 2011). 1 # # # 3!/2 , # , 3!/3 +* # , 3!/1 #, + , , 3!/3 , # " # # #, "+ (1 (Epel, 2009, Harries et al., 2010, Tilsner # , + # # « " ", #. , +* + # # (1 , # * # # # DsRed-3!/3-# 71 " COPII- , . # # 3!/3 1%$ # # , Sar1[T34N] , , 3!/3 DsRed, # # , + , DsRed-3!/3-# 71, , (1 , , , , ' " , COPII- , 4 # * ,# ' # # , # , Sar1[T34N] * # -- # # # # # # # # 3!/3. 2 1. 1 2 , 3!/1 $-$ , , . +* 2. GFP, , # # *"+ # , # , 3!/1, # , , # 3, (2) , , 3, ' + , , , 3!/3, 25 " + 3 , # # . , +, , # * (3) 3!/2 , 3 , + 3!/2 + , # , 3!/1 , 3!/1 $-$ 3!/3. 1 3. 2 3!/1 3. , (1) , 3 + ,# #. . 3!/2/3!/3-, 4. 1 , , , # , # ,# # # # . . $ + 3!/3 , " , , , # # 3!/3 , # # # . 3!/3 # # ' . ( -- # + # # # # , ## # # '* COPII- , # # " YQDLN 3!/3- # # 1 # # # . . 5. 3 3!/3 1%$ , # 3!/1 1%$ # # # 3!/3 . 0"' ( !1 ., 0&1 "( ! 2-0 . + "'' .!5"": 1. Shemyakina, E.A., Solovyev, A.G., Leonova, O.G., Popenko, V.I., Schiemann, J. and Morozov, S.Yu. (2011). The role of microtubule association in plasmodesmal targeting of potato mop-top virus movement protein TGBp1. The Open Virology Journal 5, 1-11. 2. Shemyakina, E.A., Erokhina, T.N., Gorshkova, E.N., Schiemann, J., Solovyev, A.G. and Morozov, S.Yu. (2011). Formation of protein complexes containing plant virus movement protein TGBp3 is necessary for its intracellular trafficking. Biochimie 93, 742-748. 3. # . . 0 # //XVIII «% # " # 3!/ 1 ' , -2011» (11-15 ., 2011. – -. 69-70. 26 2011): . ,. – - .–