Characteristic of the Pepper CaRGA2 Gene in Defense Responses against Phytophthora capsici Leonian

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1 J o 2013, 14, ; o:103390/jm O o Jou o ou wwwmom/jou/jm h o h 2 o hyohho o - Zh 1,2,, - J 1,, - 1,2, Ju- 1,2, -Xu 1,2 Zh-u o 1,2, * 1 2 o o ouu, ohw & Uy, , hx, h; -: zy41982@126om (-Z); j@wuu (-J); x@wuu (-); wju1127@163om (J-); yyxu2008@wuu (-X) y booy o o Booy, ohw & Uy, , hx, h h uho obu uy o h wo * uho o whom oo hou b ; -: zho@wuu; : ; x: : 25 Juy 2013; om: / : / ubh: b: h mo h o ouo wow h hyohho bh, whh u by h oomy ho, hyohho o o o h oo h, w o hz, 2, om hh ( uum 334) yz uo by h mho o -m u-u () h 2 h u-h o 3,018 b wh 2,874 b o m (O) o 957- o h o h mou wh o 1086, h o o 8106 u -m h 2 xo w y u by h xo w bw h ub u 2 w uy x h u, 334, h o 24 h ouo wh, -o hh h h o ub u Ou u u h 2 h o xo y o o h h 2 w, h w y u, obo h w uo by m-u - h ouo y

2 J o 2013, h mo h u o Ou u uo h h h 2 my how h o h y w o ow b o bo ub oomy mo u ywo: um uum ; hyohho bh; hyohho o; 2 ; uo y; u-u () 1 ouo (um uum ), mmb o h my, o, o o h mo mo b o wow hyohho bh o u by h oomy, hyohho, o o h mo u o h wo [1 4] h oo, m, u o o ho o omo,, uumb, wmo, um, uh oo [5,6] h ho h y o om u m-o, bh, o bh w oow by h o o mu mz, y x, w-o o [7 9] y 60 - () h b o om mooo o o [10] b h o oz h o ho o [11] o o h h h b hz hu h m uu mo o bo o my h o o wh uo-b (B) u-h () om [12,13] h u homoo () o o (), u o h u my o ow bu om, o o u om mo, houh m b o o mo [14] u h b om h my o [15] h o o o b b : (1) u [16]; (2) o- wh [17,18]; (3) o uoy o h uy o h xo o u ho wou ow o m whh hy y o o hy my o [19] u-u () o hh hu om o h y o uo uo hu u o o-o-uo y o [20,21] h hu o o o ow o o h o omo [22 24] oby m-oum h b omo oh jo mhoooy u h ouo o o [25,26] h xo o h umb o ho o u, uh um uum [27], oum [28], omum [29], u u [30], hhoz o [31] bo [32,33] hyohho bh o umb o u by o ou h ou o h o u o oo (426259) -bb2(122125), u m mo o

3 J o 2013, () o my, h 1, 2, 3, 4 5 w o om 334 xo how 2 o y wh oh u h uy, w mu h o um uum by h hyo u () u [34], whh o zym oo boyh, u wh u u o hoobh [35] ommoy u m xm u-u () u -m o 2, y, w u o o- -o-uo o h o o h oomy ho B o h u, w oo h 2 y o ho 2 u 21 o o h ooy w wh (u 1) wh myum u h ho w uu o h oo xo () mum o h y h myum w oo, h o hmoom h u bh (u 1b) h o o h oum mohooy w o o hm w obu-o, yom o o, wh w w h o u, b hz o oo h oum h obou mo (u 1) o o hm h o mo, wh w h wo mo h zooo w om h oum w (u 1) h ho u h hyohho bh hx w hyohho o o o h myum mohooy, oum h, z, h ymom b ou, h h o h o u ho u 1 ohooy o ooy o () h ooy mohooy o ow o y o oo xo () mum; (b) mohooy o my oum omo; () zooo ( h mo); () zooo ; () omo o h o zooo

4 J o 2013, oo u y o h 2 h 2 w o om ( uum 334) om u h 2 (B o o u116570) om 3,018 b uo, wh 2,874 b o m (O) o 957- o h o,, uo 22, h o oo,, uo 2,895 h o h mou wh o 1086, o o w ou w by 1% o ob h x z ( o how) 23 o u m hyo y u m o h homooy o u mo o ho u om B b w om u 60 u m how hh y wh oh o, u um uum bh o (o o B432551), 1 (535071), 4 (U515341), 5 (U515351) oum buboum B- o (o o ) 68%, 71%, 68%, 71% 67%, y (u 2) h oo o (mo ) u o 2 w o b o h B B (o) uo o y homooou o [36] h u o bo o -oo umy o o om, 16, B-, 8, 22, O (u 3) o o m h o h o o, u oh w m o hyo h hyo oh w b uo o uo u (u 4) h hyo oh mo 20 om how h h um uum, oum buboum, y mx, o uu, Oyz h y o oh 2 h hh y wh um uum bh o 1 (535071) h oh o ow, h 2 how o oh wh h om o h bo h h h o uu o hhy o ho, om h hh o oo o u h ou o ouo, whh h uo o 2-24 xo y o 2 u o h oo ouo mho, xo o h 2 w yz u wh u h xo o 2 w yz oo wh h u h h 2 xo w bw h ub u (u 5,b) h 2 w uu - oo o u h 12 h oow by, wh h o xo m ow h ub u (B27) wh om o oo (Ub3), wh hh o xo w 334 B602 u h m o xo w y oo o 334, h xo w uu (24 h) oo (48 h) h u o b owu h m

5 J o 2013, , wh h xo w hh h oo h hh xo o 2 x oo o B h 12 h ouo, y h hh xo o 2 x oo o B27 h h oh, boh 12 h ouo h xo h wo u ( oo) w by h m h o y mho how h m xo, x hh h (u 5) ow h oo (u 5) u 2 u u m o h u mo u o 2 wh oh o () o wh 60 how om o o boom u mo u o h 2 (um uum, B432551), 4 (um uum, U515341), 1 (um uum, ), bb- (oum buboum, ) 5 (um uum, U515351) o u h; o ou o mo h m bb ou m w y h m bb ou y h w bb ou y w w m bb ou w w h h y w bb ou w m h h y bb ou h y m bb ou m w y h bb ou bb ou

6 J o 2013, u 3 o om h o u o h 2 u 4 hyo oh o 2 o ou om oh u m w om u m 60 om, h hyo w uz u w o u u o m oow: um uum bh o (B432551), 1 (535071), 4 (U515341), 5 (U515351); oum buboum u o 3 (7X402), 4 (7X391), 4 (451741), 2 (451852), 2 (451642), 2 (451882); oum buboum bh o (290721), 3 (290711); oum buboum B- o (164801); y mx u o 3 (X ), 3 (X ), 2 (X ); o uu o (X ), u o 4 (X ); Oyz B--y o (BB440791) h w y hyom whh bh h ooo o umb o h bh boo u (o 1,000 )

7 J o 2013, u 5 u -o oym h o (-) o 2 wo ouo mho o hyohho bh, o h oo ouo mho h oh h o y mho o w o o y m oo ouo wh zooo o hyohho ( zooo/m) om () oo (b) h u h m y u o y mho o () oo () h u - w om o u - m o 2 h b h u o xo m o oow ouo o o - bw hh (b b, 334), mo (y b, B602) ub (wh b, B27) h xo o 2 w omz o h xo o ubu-oju o u w u o h 2 oow h o, o how 25 h uby o o 2- ob o 2 xo o o m o uh xm h o o-o-uo o h 2 o bo o, w h 2 u h obo u ()-b hu by my o w u oo omo u o o oo o h u h o ou w obum-o, 2- w u o xm wy y, - (-m hyo u o y

8 J o 2013, bh uz o o h ) xhb hoobh hoy, how u y o h ym u (u 6), ymom uu o by h u u 6 hoy o hyo u () w ob 334 hooh w om ou w oo () w oo; obum uu om wh obo u () o w mx 1:1 o; (b) w h o o y o ouo; () w w oo om o h: o- oo, (: 2) (: ) b 26 m-u - y y w moo by m-u - y o 2 my o oo (: 00) 2- (: 2) (u 7) o h 2 oo, m o h omb - wo ou w o o-ouo y om o h w-y - oo, wh h o Ub3 u h m m boh y h u om o h oou 2 h /2 u 7 omo o h 2 h -2- bo 2 :00 :2 :00 :2 :00 :2 ubu oo m 27 h ouo o whh h o 2 o u o o, h om o-, my o oo 2- w xo o 8 mm m myum u om hm h o mo oow by ubo h 25 o wo y hy o (w-o o) ou 2 o h 334 oy uo h, wh h

9 J o 2013, wh my o ou y y hoy 3 h ou w o 334 o o ouo, mo umou o wh w-o w o om o h oo (u 8) w ob h o- my o oo hb om, how m o z om o h o 2- u 8 ymom o o h o- oo, my o oo (: 00) (: 2) h by () o- oo; (b) wh my o; (), () wh 2 ( y) wy y u-u () m, h w h by o ouo wh myum u u w 72 h h ouo o :00 : 2 : 2 b h u y o h o h h how m o uh 2 h o z o h om h : 2 ou w oxmy 40% h ho o om h o- : 00 oo (u 9) h xm u h 2 oow u uby o, hu, 2 u o 3 uo 31 o o hyohho Bh ho h h b w o o h oo o o hyohho bh ho h o o hx o u ow h obo o hyohho bh o hx, h, how h h m ymom o h h o muy o hyohho bh,, o bow hum, w, wh mo y o h u u h ho w o, uu hyohho o mohoo obo [37]

10 J o 2013, u 9 o h o o h ou wh o om w o h o o 72 h o ouo o b h o o h m o h w o ( 005) 32 oo Boom y o 2 h uy, 2 w o om h u ( uum 334) o om h omooy y h h 2 w hhy o oh wh, om oh homooou h hhy o mo h hyo b h how h 2 bo o h o my hh h y mo h B- om, hhy mho-h- () mo,, whh u h y o h o h oow uo m by h om mo wy om h o o h u h h 3' o 5'-o o b o o 2 o uo - (hy o) h 33 xo o 2 h ouo ho -m hooy o w oou o uy hyohoy [38] Bu o y, y ouby, -m ub o y ho o o mo h ho uby o oy u o o ho ow, u o ow xo, b u o u bo y [39] h o o h X1 o oow ho ouo [40] h h h o o h o h y o -ho o

11 J o 2013, h uy, h xo o 2 w o h o om o (u 5) o wh y h h xo o 2, u oo bw h o hu, 2 mh b o h -u o m xo h b ob o 23, b 1 o 1, owy mw uumb [41], b [42,43] mo- u b [44], y h ouo mho o h o h xo o 2 omo o ouo mho o hyohho bh how hhy oo ( = o h oo ouo mho = 08384, < 001 o o y mho, uubh) wo mho u h xo o 2 u ub w ou wh h oomy,, o u h xo o o o o m h hh xo o 2 h o 334 w ob 24 h ouo, wh h oo, ou 48 h h u mo h h xo o 2 w muh hh h h h oo (u 5) [45] o h h hh m w ou h, oow by h m oo h y o xo w by uw h oww h 2 xo w bw h ub u h u 334 how muh hh xo om o h ub u, B27, u h ho h xo bw h ub mh u h o, h b xo o ub oo u h um w [46] Ou u u h h bw uby o y o o h ho h hh o o u h y ob o o 34 u-u o 2 h u-u oh h o y uu h y o uo [47 49] b hh houhu oh, o o oo mu, o o h h, bu h y u h xo o h o o houh o o m oo mu u o uy h uoy o om o o h hoy, bu b u o uy h o uh [50] O o h o h h x u () h o u h b y u o, ho, h uoy o b u [24,48] o h uo o o 2 u h o o ho, ho-u 2 xo w omom u by h hu o y, - o ox, O, u mmb o, 1, w uoy hz u [51,52] ow [53] o h uo o h o

12 J o 2013, o h uby o b ho o h 2- w hy mo ub o om o : 00 oo o 2 xo u h omb o o h uo o ub ymom o, om by oo h u h h uo o h 2 h ub o u owm o h bo-um o, hby ow h uby o ho 35 m-u - y o ow, h h o ou, xo o h Ou m-u - u o mo h h 2 xo w o ow, h h o ou, h 2-, whh o wh h u o [50] hh o w ob by -, whh how muh ow o bo1 om o h oo h u o h h ox y o h o, umb o w-o o h h oo [54] m u w ob h h h h h b ob h u o, m oo by m-u - h xo o 2 how hh m h oo h u o m-u - y o how h h o o 2 m y u h oo o o, omo o h my o (u 7) oo ou b bh bw xo o 2 h o owh o h bh,, h hh h xo, h ow h om h obo u h o uo o 2 my b o o h ouo o by h o h o owh o h, whh y om o h u [55,56] 36 uo o o 2 om om u o h h bh h om, h umb o y o oo, my o h - o u-,, hy oy om u b om oh wh h o u- o uum mo w-y u ( o how) ou h 2 u h uby m 334 (u 8 9) ou u o h h bo m4 oou how h uby o bo [57] Ou uo y mo h h 2 y o h uo by ou obo h 2 w ub ouo h 2 uby o, o ymom o owh ob h h h om h 2 how o owh om o h oo, whh my b h ou o u h

13 J o 2013, h u h h xo o h 2 my u h o 4 xm 41, ho ouo uum 334 u,, (hh ), B602 (mo ) B27 (ub), w u o o by w ow owh hmb 25 u 16 8 h o h, y h x, w ou wh [58,59] w o om h xm o ohw & Uy (, h) h ho w o om by m u om o h ow h u w o 75% oho o 30 wh w o 30 h 10% bh o 2 3 m h, h u w wh 2 3 m wh w by bob h u w o h o (oo-xo ) hyohho- mum (oo 200 / bo 30 m, 17 /, uo 20 /, 50 m/m, m 100 m/m) om h how my owh, 4 mm u w y o h o w mum h o o y bo w wh m (xb, B, h, U) o omo o h o w m o 4 w ow o h h hyohho o w om by obo o oo h o h ooy 3 y o uu o h u ho o h h, w ub u ouou h o 5 7 y h u ho w o o h uu h (9 m 9 m) ou umb o oum h myum h h oum w o h wh w o ob h mohooy o h oum u h moo Zooo w u by h uu 4 o 30 m h ub oom mu o m Zooo oo w ou by u hmoyom h oo ju o zooo/m h y w o w o mo bo ou h m o o uo w ou h 3 m ho 3 m wy om h oo o h ouo, h o w mo ub 28 w ou wh h o uo ( zooo/m) u o y mho h mo hmb 28 om h y w u wh 3 h m o oo w o u xo yh h oo w o 0, 6, 12, 24, 48, h ouo wh, o w o u zo (o, b,, U), o o h muu uo o w ou u m ommy () yh (,, h) h oo

14 J o 2013, o o w mu ohoomy u oo um (hmo oo 2000 hoo, mo,, U), h uy w u h 260/ /230 o o by oo 43 o o 2 o w x om o ou-w-o 334 o om 5' 3' o o h u 2 oo, h mo o () mho w u - yh w om u m mo (oh) wo m w o o h m (B o umb: 7XB91), my, 5'--3' (3'--1) 5'--3' (3'--2) wo 5'-- m w, my, 5'--3' (5'--1) 5'--3' (5'--2) h u m o 5' 3' h u-h w ob by mo wh - m (ow m: 5'--3'; m: 5'--3') h hm o w oow: uo 1 y 95 o 5 m; 30 y 95 o 30, 53 3 m 72 o 1 m; oow by 1 y o xo 72 o 10 m h ou w o o 19- o u 44 u y hyo y h y o 2 u w o u boom oo h uo u, u mo u o m (O) w yz, h u omo w ou houh b h u h B om (B, o o Bohooy ) [36] u u m w o u 60 (yo Boo, uu,, ) ow o om h B o o 2 wh oh o hz o hyo w ou u hbo-jo mho wh m u - o xo y u - w om wh y uoo o ym (Bo-, u,, U) w ou wh 96-w u B mx x (,, h) h o (20 μ) o o 10 μ B mx x, 2 μ u 04 μ ow m - y oo w oow: 95 o 1 m 45 y o 95 o 15, 52 o o 30 uo w o u h 52, xo o Ub3 w u o [60] h o m u w: 2 (U116570; ow, 5'- -3';, 5'--3'); Ub3 ( ;

15 J o 2013, ow, 5'--3';, 5'--3') uo o xo w u wh h - mho 46 m ouo h obo u ()- by o 1 (192) 2 (156) h b b ouy [34] m o 300 b w m om h u m o o 2 o u m 2-- (5'- -3') 2--(5'- -3'), y 2-2 h ou w h ou o h obum um, 3101, by h z-hw mho 47 u-u () o 2 (obo u)-b w ou oow h ou omz ouy o h y [61,62] om wh 1, 2-, 2- (hyo u, o oo) 2-my o ( oo) o um 3101 by h z-hw mho B w ow 28 o u-b (B) mum h o 1 2 w om y h u by ooy y ooy w ou 10 m u uu o B (y x 10, o 10, 5 /) wh o bo (50 m/m o my 50 m/m o m), h 200 m 28 o h h O600 h u bw h, 2 m o uu w ou o 50 m uo mum () o bo, 10 m 200 μ oyo h uu w ub wh h 200 m 28 o h h O600 h u bw 05 1 h b w h by uo u uy o o bu wh 10 m 2, 10 m, 200 μ oyo, 55, ju o O o 10 oom mu o 3 5 h bo o o h obum o 1 2 o o (2 o 2-2) w mx 1:1 o o o, mxu o h obum o 1 2 o w o h 334 oyo o 2 o 3-w-o wh 1 m - y h obum- w o owh hmb w m 17 o 1 60% humy h, hy w oh owh oom 25 wh 16 h h 8 h hooo y u 3 4 w o m o h xm, 60 w u o h xm xm w w 48 m-u - y o w x om oo, m o w-y 334, o- ( wh my o 1 2) u zo (o, b,, U), o o h muu ooo o m-u -, m w

16 J o 2013, o ou h o o o u h oy h oou wou b, xo o Ub3 w u o [60] 49 y o h y w u o yz h o h oomy ho, h h (9 m 9 m) w o om hhy uy-x o 4-w-o wh h o-, my o oo (: 00) h ou (: 2) u o w ow o x y o oo xo (, o) 28 bo b u o h ouo by 8 mm m myum u om oo h bx u o wo oo (h h ouh u ) w y m, h xm w h m h w mmy wh m o m hh humy h 1 ub omy oo owh hmb (28 ; uo h 60 μmo m 2 o 12-h hooo) hoy o z o h ou h w hooh 72 h o ouo (h) 410 y h u o w ubj o y u y ym ow ( u, o 82) oow h o-wy y o (O) () 005 w u o om h mo m w x h m ± o h ( = 3) 5 ouo h 2 w om o o w hz y o h uo o 2 by u - o xo y 334, 2 xo w y u by, u o ho o o h ou ou uo o o 2 b o o b o oh u h hu h w uo by obo h 2 h h ombo o h y o h b o o b owu oo o y h my y o o uh u o h uo w h u y h mhm o h o b h ou wo w h u b u h mou mhm o oo owm h h w uo by h o u ouo o h (o ), h wh - o o hooy u (o 2011B12B03) h hx o hooy oo o

17 J o 2013, oj ( ) h hx o hooy om oj ( ) o o h uho o o o 1 w, B; m, hyohho bh o b oo o 1995, 79, , B; m, B; h, ; w, B o um u o o o hyohho om um b 2001, 85, o, JB; Joho, ooy b oh o mm o hyohho bh o b 1999, 83, ub, ; mou, hyohho o b o: h o mm h 2004, 88, B, ; Buo, B;, ;, hyohho zooo o o u ou hy om o O 1995, 75, O, ; Bo, o o o hyohho oo o o bh um uum J m o o 2003, 128, By, O; uu, ; o, bzo--mhy u o hyohho Bo 2005, 49, o, B; o, J; o, ;, h o o hoom b o o o hyohho 2012, 3, ou, ; ohoo, ;, wmo ooo o ow o u by hyohho o o 2012, 39, , B; Boo, J;, U h uo o o u Bo 2003, 54, , J; Jo, J ho o o o u 2001, 411, B, J;, ;, J;, ; mm, J; bb, ; Zho, B; u, ; o, J; h, J; y uo b -u-h om 2002, 12, o, B; Zhu, ; Bum, ;, ; y, ; oo, ; ou, y, buo, xoom oh wh h o- B- ubm J o o 2002, 54, ow, J; mo, h o ouo o ho 2006, 7, , X; Xo, B;, ; u, XJ;, ; u, X; X, J;, ;, o o B-y homoo obo om o Bo 2010, 28,

18 J o 2013, , ;, ; Zh, J; Zh, Z;, J; h, oo hzo o u-h homoo om oyb ho 2003, 106, w, O; Bouz, ;, ; ho, J; ou bouh, ; o, ; ouzy, o o o -B- m o h 5/8 ou o o owy mw uow ho 2003, 106, , ; h, X;, ;, o oymohm m o- wh h 5 o o wh u ho 2003, 106, uz, ; méz, J; o, ; z, ;, ; omo, U-uo o o () h h y o o uum w uhy 2012, 186, ozb, ; Boo, ; o, ; ou, By mo u u mooo J 2002, 30, , XZ; Xu, ;, ; Zh, Z om o u-u - ym o uo y o h 2- hwy bo o Bo 2006, 62, Buomb, ow b o u-u u O Bo 1999, 2, h-um, ; hm, ; h, ; h, ; u, u-u ho o Bo 2003, 236, um, ; o, ;, ; yo, ; Uyum, uo hzo o o bhm homoo o u w -u by u-u 2007, 225, u, ; h, ; h, ; h-um, obo 1, 1 1/1 u o -m o obo mo u J 2002, 30, , ; -z, ; Buomb, h : obo u o o y o uo by J 2001, 25, hu, ; o, ; m, ; hu, J; Oh, ;, ;, J; Jou, ; ho, mho o hh uy u-u h (um uum Bu) o 2004, 17, B, ; -z, ; J, ; h, J; Buomb, ; B, B; Jo, J u-u oum J 2004, 39, m, ;, ; o, ;, ; h, u-u oo o y uo h b uo omum (oum oy) J 2005, 44, ou, B; m, u-u oo o uo y h m mo u (oumb, uu) ho 2007, o:101186/ , ; hoz, ; b, ; B, hy u u () o oy (hhoz o): uo o y o ob uo om o-mo Bo (o) 2007, 100,

19 J o 2013, Buh-mh, ; h, ; u, ; h-um, u u bo hyo 2006, 142, u, ;, J Omz b o u-u () u obo u ho 2008, o:101186/ u, ; h, ; h-um, u-u omo J 2002, 31, um, ; oo, J; -o, ; y, ; y, ;, yom hbo o oo boyh wh u- o U 1995, 92, Boom oo b o: h://wwwbmho/b/ ( o 23 mb 2012) 37 Zh, XB hyohho h hu; uu : Bj, h, 1997; , ; íz, J; o, -m oym h o uo o hyohho oy hyohooy 2005, 95, oé, ; bh, ;, ;, ; mz, wou-u -xo omo bo jmo boyh 1993, 191, ohmu, ; mouh, U; yo, ; o, xo o X1, b bh-, u by b ouo o U 1998, 95, , J; Zho, ZZ;, ;, ; h, J o hzo o o B-o uo o u o wh owy mw uum B Bo 2010, 10, , ZX; mouh, U; yo, ;, ; o, xo o h b -b- my u-u by om oo o o by hm h oy o Bo 2001, 47, , ZX; o, ; mouh, U; wmoo, ; o, ; y, ; yo, ;, h b o b bo o h uo b u-h o J 1999, 19, huu, ;, ; Ju, ;, h omo o h mo 1 o-1 mo-o - xo u b (B u ) bo h o Bo 2003, 52, , ; o, ; íz, J o o - u wh hyohho J hyo 2008, 165, Zh, ;, X;, ; h, J; Zh, ;, Z o hzo o um w om oyum bb uo y bo h 2011, 30, u, ; u, ; o, ; uz, ;, J; u, J; hj, J; Bhm, ; y, ; Buomb, h houhu u-u m h ho o 90 BO J 2003, 22, Buh-mh, ; o, J;, B; h-um, o o u u o uo u J 2004, 39,

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