鎽樸€€瑕侊細锛凤疾锛脊 杞綍璋冩帶鍥犲瓙鐨勭敓鐗╁鍔熻兘娑夊強妞嶇墿鐢熼暱鍙戣偛銆佺墿璐ㄤ唬璋€佹姉鐥呰€愰€嗐€佹哀鍖栬“鑰佺瓑璇稿鏂归潰銆傛按绋诲叏鍩哄洜缁勬祴搴忓畬鎴愬悗锛屾按绋伙挤锛诧极锛?杞綍璋冩帶鍥犲瓙鍩哄洜鐨勫姛鑳界爺绌堕殢涔嬮€愭笎寮€灞曪紝鐩墠宸茬粡鍙戠幇瀹冨湪妞嶇墿鎶楃梾銆佽€愰€嗐€佽“鑰併€佺硸浠ｈ阿浠ュ強褰㈡€佸缓鎴愭柟闈㈠彂鎸ラ噸瑕佷綔鐢ㄣ€傞殢鐫€鐮旂┒鐨勬繁鍏ワ紝姘寸ɑ锛凤疾锛脊 鍩哄洜鐨勭紪鍙锋湭鑳界粺涓€锛屽緢瀹规槗璁╀汉娣锋穯锛屾湁蹇呰杩涜鏍℃銆傜粨鍚堜綔鑰呭疄楠屽鐨勪竴浜涚爺绌剁粨鏋滐紝瀵规按绋伙挤锛诧极锛?鍩哄洜瀹舵棌鐨勭爺绌剁幇鐘惰繘琛屼簡缁艰堪锛屼互鏈熶负杩涗竴姝ユ繁鍏ュ紑灞曪挤锛诧极锛?鍩哄洜瀹舵棌鐨勭爺绌舵彁渚涘府鍔┿€?/p>
涓浗姘寸ɑ绉戝 (Chinese Journal of Rice Science)锛?/span>2009锛?/span>23锛?/span>447-455銆?/span>
鍓嶈█
姘寸ɑ鍏ㄥ熀鍥犵粍娴嬪簭宸ヤ綔鐨勫畬鎴愪娇姘寸ɑ鎴愪负绮浣滅墿涓熀鍥犺儗鏅渶涓烘竻鏅扮殑鐗╃[1]锛屼篃涓哄叾浠栧崟瀛愬彾妞嶇墿鍩哄洜缁勬祴搴忓伐浣滄彁渚涗簡寰堝ソ鐨勫弬鑰冦€傚悓鍏朵粬鐢熺墿涓€鏍凤紝姘寸ɑ鍩哄洜缁勪腑涔熷寘鎷紪鐮佽浆褰曡皟鎺у洜瀛愮殑鍩哄洜锛屽叾涓竴浜涜浆褰曡皟鎺у洜瀛愬鏃忔嫢鏈変笂鐧句釜鎴愬憳鍩哄洜锛屾瘮濡?/span>MYB瀹舵棌锛?/span>WRKY瀹舵棌锛?/span>AP2/EREBP瀹舵棌锛?/span>bHLH瀹舵棌锛?/span>bZIP瀹舵棌锛?/span>C2H2瀹舵棌锛?/span>NAC瀹舵棌绛?/span>[2, 3]锛屽畠浠湪姘寸ɑ鐨勭敓闀垮彂鑲插拰鑰愰€嗘姉鐥呰繃绋嬩腑閮藉彂鎸ラ噸瑕佺殑鐢熺墿瀛﹀姛鑳姐€?/span>
鐪熸牳鐢熺墿涓殑杞綍璋冩帶鍥犲瓙鍙堢О鍙嶅紡浣滅敤鍥犲瓙锛屾槸涓€绫婚€氳繃涓庡叾闈跺熀鍥犲惎鍔ㄥ瓙鍖哄煙鐨勯『寮忎綔鐢ㄥ厓浠剁壒寮傛€х粨鍚堬紝浠庤€屽奖鍝嶉澏鍩哄洜杞綍姘村钩鏄捐憲鍙樺寲鐨?/span>DNA缁撳悎铔嬬櫧銆傝繖绉嶇粨鍚堢殑鐗瑰紓鎬у弽鏄犲湪杞綍璋冩帶铔嬬櫧鐗规畩鐨勭┖闂寸粨鏋勫拰椤哄紡浣滅敤鍏冧欢鍩烘湰瀵瑰簲鐨勬牳閰稿簭鍒椾笂銆?/span>WRKY杞綍璋冩帶鍥犲瓙瀹舵棌涓ぇ閮ㄥ垎鎴愬憳鍏锋湁鍏稿瀷鐨?/span>WRKYGQK缁撴瀯鍩燂紝閮ㄥ垎涓?/span>WRKYGKK[4]锛岃€屽湪姘寸ɑ涓張鍙戠幇浜嗘柊鐨?/span>WRKY缁撴瀯鍩?/span>WRKYGEK[5]銆?/span>WRKY杞綍璋冩帶鍥犲瓙铔嬬櫧缁撳悎鐨?/span>DNA搴忓垪鏄?/span>W鐩掞紙T锛?/span>TGACC锛?/span>A/T锛夛紝涓€鑸涓烘牳蹇冨簭鍒楁槸TGAC鐨勫惎鍔ㄥ瓙搴忓垪灏卞叿澶囦竴瀹氱殑W鐩掑姛鑳?/span>[6]銆?/span>
鍒扮洰鍓嶄负姝紝閫氳繃鐢熺墿淇℃伅瀛︾殑鏂规硶宸茬粡鍦ㄦ按绋荤殑涓や釜鏍藉煿绉嶇渤绋绘棩鏈櫞锛?/span>japonica锛夊拰绫肩ɑ93-11锛?/span>indica锛変腑鍒嗗埆棰勬祴鍒?/span>103涓拰102涓?/span>WRKY鍩哄洜[7, 8]銆傝嚜2000骞寸涓€涓按绋?/span>WRKY鍩哄洜OsWRKY4琚厠闅嗗悗[9]锛屽凡鏈夋暟绡囨姤閬撴彮绀烘按绋?/span>WRKY鍩哄洜鐨勫姛鑳藉箍娉涘湴娑夊強浜嗙敱鐥呮瘨[10]銆佺粏鑿屻€佺湡鑿?/span>[11~13]绛夊紩璧风殑鐥呭鍙嶅簲鍙?/span>SA鍜?/span>JA鐩稿叧鐨勪俊鍙烽€氳矾[11]銆傛垜浠巼鍏堝姘寸ɑWRKY鍩哄洜杩涜闈炵敓鐗╅€嗗鑳佽揩鍚庣殑琛ㄨ揪璋卞垎鏋愮粨鏋滆〃鏄庝竴浜?/span>WRKY鍩哄洜寰堝彲鑳芥秹鍙婁簡姘寸ɑ鐨勮€愰€嗗弽搴?/span>[5]銆傝繘涓€姝ュ姛鑳藉垎鏋愬彂鐜板婧愭€х殑楂樿〃杈?/span>OsWRKY45浼氬寮鸿浆鍩哄洜鎷熷崡鑺ュ骞叉棻鑳佽揩鐨勮€愭€?/span>[14]锛屽唴婧愭€х殑杩囪〃杈?/span>OsWRKY11鍙互澧炲己姘寸ɑ瀵归珮娓╁拰骞叉棻鑳佽揩鐨勮€愭€?/span>[15]锛屽畠浠潎浠庨仐浼犲鐨勮搴﹁瘉瀹炰簡姘寸ɑWRKY杞綍鍥犲瓙鍙備笌妞嶇墿鑰愰€嗘€х殑鎺ㄦ祴銆傚叾娆¤繕鍙戠幇澶栨簮鎬х殑楂樿〃杈?/span>OsWRKY23浼氫績杩涜浆鍩哄洜鎷熷崡鑺ュ彾鐗囩殑琛拌€?/span>[16]锛岃€屼竴浜涙姤閬撲篃鎺ㄦ祴OsWRKY17[17]銆?/span>OsWRKY4鍜?/span>OsWRKY82 [18]寰堝彲鑳藉湪姘寸ɑ鐨勮“鑰佽繃绋嬩腑鍙戞尌涓€瀹氱殑浣滅敤銆傛澶栬繕鎶ラ亾浜嗗婧愭€х殑楂樿〃杈?/span>OsWRKY72鍙互褰卞搷杞熀鍥犳嫙鍗楄姤妞嶆牚鐨勫舰鎬佸缓鎴?/span>[19]锛屽唴婧愭€х殑杩囪〃杈?/span>OsWRKY31浼氬奖鍝嶆按绋绘牴鐨勫舰鎬佸缓鎴?/span>[20]銆傜敱姝ゅ彲瑙佹按绋?/span>WRKY鍩哄洜鐨勫姛鑳藉叿鏈夐噸瑕佹€у拰澶氭牱鎬с€?/span>
缁撳悎鏈疄楠屽鐨勭浉鍏崇粨鏋滐紝鎴戜滑瀵规按绋?/span>WRKY杞綍璋冩帶鍥犲瓙瀹舵棌鎴愬憳鐨勫垎绫绘儏鍐靛拰宸叉姤閬撴按绋?/span>WRKY杞綍璋冩帶鍥犲瓙鍩哄洜鐨勫姛鑳藉仛涓€缁艰堪锛屼互鏈熷鏈潵鐨勭爺绌舵湁鎵€甯姪銆?/span>
1. 姘寸ɑWRKY杞綍璋冩帶鍥犲瓙鐨勭郴缁熷垎缁勫拰缂栧彿
2004骞撮鍏堢敱Shen瀹為獙瀹わ紙77涓級[21, 22]鍜屾垜浠疄楠屽锛?/span>97涓級[5]鎶ラ亾浜嗘按绋?/span>WRKY杞綍璋冩帶鍥犲瓙鎴愬憳鐨勫熀鍥犳敞閲婂強鍚勬垚鍛樺垎缁勬儏鍐点€?/span>2005骞?/span>Guo绛?/span>[8]鍜?/span>Zhang绛?/span>[23]鍚勮嚜鍙戣〃浜嗗叧浜庢按绋?/span>WRKY杞綍璋冩帶鍥犲瓙瀹舵棌鎴愬憳鐨勫垎缁勬儏鍐点€傝€屾渶鏂板彂琛ㄥ湪銆婃鐗╁鎶ャ€嬩笂鐨勫垎缁勬儏鍐靛垯鏄敱Shen瀹為獙瀹ょ殑杩涗竴姝ュ畬鍠?/span>[24]銆備互涓婂洓涓皬缁勭殑鍒嗙粍鍩烘湰涓婂埄鐢ㄤ簡Eulgem绛夌殑鍒嗙粍鍘熷垯锛氱涓€绫诲瀷鐨勬垚鍛樿泲鐧藉簭鍒楀惈鏈変袱涓吀鍨嬬殑WRKY缁撴瀯鍩熷拰Cys2-His2閿屾寚鍨嬬粨鏋勶紝绗簩绫诲瀷鐨勬垚鍛樻嫢鏈変竴涓吀鍨嬬殑WRKY缁撴瀯鍩熷拰Cys2-His2閿屾寚鍨嬬粨鏋勶紝绗笁绫诲瀷鍒欎互鍏舵垚鍛樺惈鏈変竴涓吀鍨嬬殑WRKY缁撴瀯鍩熷拰Cys2-His/Cys閿屾寚鍨嬬粨鏋勪负鍩烘湰鐗瑰緛[4]銆傜敱浜庡湪鍒嗙被杩囩▼涓WRKY鎴愬憳鍛藉悕鐨勫樊寮傦紝闄や簡鏈疄楠屽鍜?/span>Shen瀹為獙瀹や繚鎸佷竴鑷翠互澶栵紝鍏跺畠涓ょ粍鐨勫懡鍚嶅簭鍙峰瓨鍦ㄨ緝澶у樊寮傘€傛瘮濡傜敱Xie绛夊拰鎴戜滑鍛藉悕鐨?/span>OsWRKY72锛屽湪Wu绛夌殑鏂囩珷涓鍛藉悕涓?/span>OsWRKY92锛屽湪Zhang and Wang鏂囩珷鐨勪腑琚懡鍚嶄负OsWRKY78銆傛绫婚棶棰橀€犳垚浜嗚鏂囧拰璇存槑鐨勪笉渚匡紝涔熷洶鎵扮潃鍥藉唴澶栧悓琛?/span>[7]銆傜粺璁＄洰鍓嶅凡鍙戣〃鏂囩珷鐨勪娇鐢ㄦ儏鍐碉紝澶ч儴鍒嗘牴鎹?/span>Xie绛夊拰鎴戜滑鐨勭紪鍙凤紝涓ょ瘒閲囩敤Wu绛夌殑缂栧彿锛岃€?/span>Zhang 绛夌殑缂栧彿鍩烘湰鏈閲囩撼銆傛牴鎹紭鍏堝師鍒欙紝鎴戜滑寤鸿浣跨敤鏈疄楠屽鍜?/span>Shen瀹為獙瀹や竴鑷寸殑缂栧彿锛屽苟璋冩暣OsWRKY55鍜?/span>OsWRKY100鐨勫懡鍚嶏紙鍚?/span>OsWRKY31涓?/span>OsWRKY89璋冩崲锛夈€傛垜浠富瑕侀噰鐢ㄩ殢鏈虹紪鍙风殑鏂瑰紡渚濇灏嗘柊鍙戠幇鐨勫熀鍥犻『娆＄紪鍙?/span>[5, 21, 22, 24]锛岃繖绉嶇紪鍙锋柟寮忓湪涓嶇牬鍧忓師鏈夌紪鍙锋搴忕殑鎯呭喌涓嬫湁鍒╀簬鏂伴壌瀹氬熀鍥犵殑鍛藉悕鍜屾帓搴忋€備负浜嗕究浜庢煡闃咃紝鎴戜滑灏嗕笁绉嶇紪鍙锋柟寮忕粺绛规垚琛紙琛?/span>1锛夈€?/span>
姣旇緝涓ょ涓昏鐨勭紪鍙锋柟寮忓拰鍩哄洜娉ㄩ噴鍙互鍙戠幇閮藉瓨鍦ㄤ竴浜涢棶棰樸€傝櫧鐒?/span>Wu绛夊姘寸ɑWRKY鐨勭紪鍙锋€讳綋涓婇噰鐢ㄤ簡椤烘煋鑹蹭綋鐨勪綅缃懡鍚?/span>[8]锛屼絾鍏朵腑浜屽崄澶氫釜鎴愬憳鐨勯『搴忓苟涓嶇鍚堣瑙勫垯锛屽浠?/span>OsWRKY96锛?/span>Os12g02470锛夊埌OsWRKY100锛?/span>Os12g02400锛夐兘娌℃湁缁濆椤烘煋鑹蹭綋浣嶇疆鍛藉悕锛堣〃1锛夈€傚叾娆★紝Os03g21710銆?/span>Os11g02520銆?/span>Os12g02400銆?/span>Os12g02440鍥涗釜WRKY铔嬬櫧鐢?/span>Wu绛夊懡鍚嶄负OsWRKY32銆?/span>OsWRKY89銆?/span>OsWRKY100銆?/span>OsWRKY98锛岃€?/span>Xie绛夊垯鍛藉悕涓?/span>OsWRKY44銆?/span>OsWRKY100銆?/span>OsWRKY81銆?/span>OsWRKY56锛屼絾鏄袱缁?/span>WRKY铔嬬櫧鐨勫簭鍒椾笉涓€鑷?/span> [8, 22]銆傚彟澶栵紝鍊煎緱娉ㄦ剰鐨勬槸涓嶈兘鎺掗櫎鍦ㄦ按绋讳腑鏈夋柊鐨?/span>WRKY鍩哄洜琚壌瀹氾紝杩欎簺鏂扮殑鎴愬憳灏嗘棤娉曢噰鐢ㄩ『鏌撹壊浣撶殑浣嶇疆鍛藉悕鐨勬柟寮忥紝濡?/span>Os04g04300鍜?/span>Os08g09840[7]锛屾垜浠皢鍏惰ˉ鍏呭懡鍚嶄负OsWRKY101鍜?/span>OsWRKY102锛堣〃1锛夈€?/span>
2. 姘寸ɑWRKY杞綍璋冩帶鍥犲瓙鐨勭敓鐗╁鍔熻兘鍒濇鍒嗘瀽
琛ㄨ揪璋卞垎鏋愭湁鍔╀簬瀵瑰熀鍥犲姛鑳藉仛鍒濇鐨勬帹娴嬶紝涓昏閲囩敤Northern 鏉備氦銆?/span>RT-PCR銆?/span>Microarray绛夊垎瀛愮敓鐗╁鏂规硶銆傛ā寮忔鐗╂嫙鍗楄姤WRKY杞綍璋冩帶鍥犲瓙鐨勫姛鑳界爺绌朵竴寮€濮嬪氨杩涜浜嗚〃杈捐氨鐨勫垎鏋愶紝鎴愬姛鍦伴娴嬩簡鍦ㄦ鐗╄€愰€嗗拰鎶楃梾鍙嶅簲涓彂鎸ラ噸瑕佺敓鐗╁鍔熻兘鐨勬垚鍛樺熀鍥犮€傛洿杩涗竴姝ョ殑閬椾紶瀛︾爺绌跺垯璇佸疄WRKY杞綍璋冩帶鍥犲瓙鍦ㄨ皟鎺ф鐗╁厤鐤弽搴旀柟闈㈠叿鏈夋櫘閬嶆€?/span> [25]銆傝繎骞存潵鍦ㄦ按绋讳腑涔熻瘉瀹炰簡涓€浜涙垚鍛樺弬涓庢鐗╁厤鐤弽搴旓紝涓昏鏈?/span>OsWRKY12[26]銆?/span>OsWRKY13[27]銆?/span>OsWRKY45[13, 14]銆?/span>OsWRKY53[12]鍜?/span>OsWRKY71[28, 29]銆傞櫎鍙備笌鐢熺墿閫嗗鐨勫簲绛斿弽搴斿锛屼竴浜涙按绋?/span>WRKY杞綍璋冩帶鍥犲瓙杩樺彈鐩愬銆佸共鏃便€佷綆娓┿€侀珮娓?/span>[5, 7, 14, 15]鍜岃緪灏?/span>[30]绛夐潪鐢熺墿閫嗗鐨勮瀵笺€傛澶栵紝杩樻湁鐮旂┒琛ㄦ槑姘寸ɑWRKY鍩哄洜瀹舵棌鎴愬憳鍦ㄨ皟鎺ф鐗╃殑琛拌€佽繃绋嬪拰褰㈡€佸缓鎴愯繃绋嬩腑鍏锋湁閲嶈浣滅敤銆?/span>
2.1鍙備笌鎶楃梾 妞嶇墿涓ぇ澶氭暟WRKY杞綍璋冩帶鍥犲瓙鍙備笌鎶楃梾鍙嶅簲鐨勪俊鍙蜂紶閫掋€傜敱浜庡姘寸ɑWRKY鍩哄洜瀹舵棌鐨勭爺绌惰捣姝ヨ緝鏅氾紝鐩墠澶у鏁扮粨鏋滃彧鍋滅暀鍦ㄥ埄鐢ㄧ梾鍘熻弻澶勭悊鍚庣殑琛ㄨ揪璋卞垎鏋愪笂銆傜敤绋荤槦鑿岋紙Magnaporthe grisea锛夊拰鐧藉彾鏋弻锛?/span>Xanthomonas oryzae pv. oryzae锛夊鐞嗗悗鐨勫熀鍥犲垎鏋愭樉绀猴紝瓒呰繃1/3鐨?/span>WRKY鍩哄洜琛ㄨ揪姘村钩鍙戠敓鏄捐憲鍙樺寲[11]锛屽叾涓緝涓鸿灏界殑鐮旂┒涓昏闆嗕腑鍦?/span>OsWRKY12銆?/span>OsWRKY71銆?/span>OsWRKY45鍜?/span>OsWRKY13鍦ㄧ梾瀹充镜鏌撹繃绋嬩腑鐨勫垎鏋愩€?/span>OsWRKY12鍦ㄦ姉鐥呰繃绋嬩腑鐨勪綔鐢ㄩ鍏堟槸鍒╃敤鍏跺湪鐧藉彾鏋粏鑿屻€佹按鏉ㄩ吀銆佽寜鑾夐吀鐢查叝鍜屼箼鐑埄澶勭悊鍚庡彈璇卞鐨勮〃杈捐氨鎺ㄦ柇鐨勶紝杞熀鍥犳按绋荤殑鍔熻兘鐮旂┒鍙戠幇杩囪〃杈?/span>OsWRKY12鍩哄洜鍚庢按绋诲唴婧愮殑鎶楃梾鍩哄洜OsNPR1銆?/span>OsPR1b銆?/span>ZB8鍜?/span>POX22.3鐨勮〃杈炬按骞冲潎鏄捐憲涓婅皟[26]銆?/span>OsWRKY71鍦ㄦ姉鐥呰繃绋嬩腑鐨勪綔鐢ㄩ鍏堟槸鍒╃敤SA鍜?/span>MeJA澶勭悊鍚庣殑琛ㄨ揪璋卞垎鏋愬彂鐜扮殑锛屽畠鐨勮〃杈鹃噺鍦ㄥ鐞嗗悗0.5灏忔椂灏辨湁鏄庢樉澧炲己锛岃繃琛ㄨ揪OsWRKY71鐨勮浆鍩哄洜姘寸ɑ瀵圭粏鑿屾€х櫧鍙舵灟鐥呯殑鎶楁€у緱鍒颁簡澧炲己锛岃€屼笖鎶楃梾鐩稿叧鍩哄洜OsNH1鍜?/span>OsPR1鐨勮〃杈鹃噺涔熷湪杞熀鍥犳按绋讳腑鏄捐憲澧炲己[28]銆備笌姝ゅ悓鏃讹紝鍙︿竴鐮旂┒灏忕粍杩樺彂鐜?/span>OsWRKY71鍙楃湡鑿岃泲鐧?/span>Chitin oligosaccharide 鍜?/span>Fugal cerebroside鐨勮瀵硷紝瀹冪殑琛ㄨ揪閲忎篃鍦ㄥ鐞嗗悗0.5灏忔椂灏辨槑鏄惧鍔狅紝鍩哄洜鑺墖鍒嗘瀽鍙戠幇杩囪〃杈?/span>OsWRKY71鐨勮浆鍩哄洜姘寸ɑ缂栫爜鍑犱竵璐ㄩ叾锛?/span>Chitinase锛夌殑6涓熀鍥犲拰PR5鍩哄洜鐨勮〃杈鹃噺鏄捐憲澧炲己[29, 31, 32, 33]锛岃繖浜涢兘涓鸿繃琛ㄨ揪OsWRKY71鐨勮浆鍩哄洜姘寸ɑ澧炲己瀵圭粏鑿屾€х櫧鍙舵灟鐥呮姉鎬х殑鍒嗗瓙鏈虹悊鎻愪緵浜嗚瘉鎹紱鐩哥被浼肩殑鏄?/span>OsWRKY53鍩哄洜锛屽悓OsWRKY71鐨勮〃杈捐氨鐩镐技锛屽畠涔熷彈涓婅堪涓や釜鐪熻弻铔嬬櫧鍜岀ɑ鐦熻弻璇卞[12, 31, 32]锛屼笖杩囪〃杈捐浆鍩哄洜妞嶆牚鍐?/span>PBZ1銆?/span>PR14銆?/span>Chitinase1銆?/span>PR5 鍜?/span> Chitinase2鐨勮〃杈鹃噺涔熸樉钁楀寮猴紝杩涜€岃〃鐜板嚭浜嗗鐪熻弻鎬хɑ鐦熺梾鐨勬姉鎬?/span>[12]銆傚拰OsWRKY71涓嶅悓鐨勬槸OsWRKY45锛?/span>OsWRKY71杩囪〃杈炬按绋讳腑NH1鐨勮〃杈鹃噺鏄樉钁楀寮虹殑锛?/span>OsWRKY45鐨勬按绋荤獊鍙樹綋鍜?/span>NH1鐨勬按绋荤獊鍙樹綋閮戒笉浼氬奖鍝嶅鏂圭殑琛ㄨ揪姘村钩锛岃€?/span>OsWRKY45鐨勭獊鍙樹綋鍜岃繃琛ㄨ揪姘寸ɑ閮戒笉浼氭樉钁楀奖鍝嶆姉鐥呯浉鍏冲熀鍥?/span>PR1a鍜?/span>PR1b鐨勮〃杈撅紝浣嗗嵈鑳芥樉钁楁姂鍒跺拰璇卞GST鍜?/span>P450鐨勮〃杈撅紝鏇翠负娣卞叆鐨勭粨鏋滆繕琛ㄦ槑BTH璇卞鐨?/span>OsWRKY45鍦ㄦ按绋绘姷鎶楃ɑ鐦熺梾杩囩▼涓彂鎸ラ噸瑕佺殑浣滅敤[13, 34]锛涘彟澶栵紝鍦ㄧ粏鑿屾€х櫧鍙舵灟鑿屽拰鐪熻弻鎬хɑ鐦熻弻渚垫煋姘寸ɑ鐨勮繃绋嬩腑鍙戠幇琚瀵肩殑OsWRKY13[35]鍙互璋冩帶閮ㄥ垎姘存潹閰镐俊鍙烽€斿緞涓婃父鎴栬€呬笅娓哥殑鍩哄洜[36]锛岃繃琛ㄨ揪OsWRKY13鐨勮浆鍩哄洜姘寸ɑ杩樿〃鐜板嚭瀵圭粏鑿屾€х櫧鍙舵灟鑿屽拰鐪熻弻鎬хɑ鐦熻弻鐨勬槑鏄炬姉鎬?/span>[27]锛?/span>OsWRKY13琛ㄨ揪鍙楁姂鍒剁殑杞熀鍥犳按绋诲垯瀵逛笂杩扮梾瀹虫洿鍔犳晱鎰燂紝杩欎簺缁撴灉寮烘湁鍔涘湴璇存槑鍙互閫氳繃璋冩帶浣滅敤婵€娲绘按绋荤殑鎶楃梾鍙嶅簲锛屾洿涓虹粏鑷寸殑宸ヤ綔杩樻壘鍒颁簡鍦ㄦ鎶楃梾鍙嶅簲淇″彿閫斿緞涓皟鎺?/span>OsWRKY13鐨勪笂娓稿厓浠?/span>[37]銆?/span>
鍙﹀寰堝瀹為獙瀹ゅ湪瀵规按绋昏繘琛屾姉鐥呭疄楠岀殑鍚屾椂涔熺瓫閫夊埌涓€浜涜〃杈鹃噺鍙楀奖鍝嶇殑WRKY鍩哄洜銆傚湪浜插拰鎬хɑ鐦熺湡鑿屼镜鏌撴按绋荤殑杩囩▼涓紝 OsWRKY28銆?/span>OsWRKY56銆?/span>OsWRKY45銆?/span>OsWRKY72銆?/span>OsWRKY26 [38]鍜?/span>OsWRKY52鍩哄洜鏄捐憲涓婅皟5鍊嶄互涓?/span>[39]锛涘湪姘寸ɑ鐭缉鐥呮瘨渚垫煋姘寸ɑ9澶╁悗锛?/span>OsWRKY77銆?/span>OsWRKY55銆?/span>OsWRKY8鍩哄洜鏄捐憲涓婅皟2鍊嶄互涓?/span>[10]锛涘湪鍗婂瘎鐢熸鐗╄彑涓濆瓙锛?/span>Striga hermonthica锛変笌姘寸ɑ鐩镐簰浣滅敤鐨勮繃绋嬩腑锛?/span>OsWRKY19銆?/span>OsWRKY45銆?/span>OsWRKY62銆?/span>OsWRKY76銆?/span>OsWRKY77绛夊熀鍥犳樉钁椾笂璋?/span>5鍊嶄互涓?/span>[40]銆?/span>
2.2鍙備笌鑰愰€?/span> 鍦ㄥ緢澶氭鐗╀腑閮藉彂鐜?/span>WRKY杞綍璋冩帶鍥犲瓙鍙備笌閫嗗鑰愬彈鐨勪俊鍙疯浆瀵艰繃绋嬶紝濡傚ぇ楹︼紙Hordeum vulgare锛夌殑HvWRKY38鍙楀喎瀹冲拰骞叉棻鐨勮瀵?/span>[41]锛屾矙婕犺嵂鐢ㄥ父缁跨亴鏈ㄤ笁榻挎媺鐟為樋锛?/span>Larrea tridentata锛夌殑LtWRKY21鍩哄洜鍙互婵€娲昏劚钀介吀淇″彿杞閫斿緞骞跺弬涓庤劚钀介吀浠嬪鐨勬姉鏃辨€х殑寤虹珛[42]銆傚湪绛涢€夋按绋诲彈閫嗗璇卞鐨?/span>WRKY杞綍璋冩帶鍥犲瓙鐨勭爺绌跺伐浣滀腑锛屾垜浠彂鐜?/span>OsWRKY8銆?/span>OsWRKY9銆?/span>OsWRKY12銆?/span>OsWRKY13銆?/span>OsWRKY14銆?/span>OsWRKY16銆?/span>OsWRKY17銆?/span>OsWRKY21銆?/span>OsWRKY23銆?/span>OsWRKY24銆?/span>OsWRKY26銆?/span>OsWRKY30鍜?/span>OsWRKY45鍦ㄩ珮鐩愩€佸共鏃便€佸喎瀹炽€侀珮娓╃瓑鍥涚閫嗗鍥犲瓙鑳佽揩涓笉鍚岀▼搴﹀湴琚瀵?/span>[5]銆傚婧愰珮琛ㄨ揪杞熀鍥犳嫙鍗楄姤閬椾紶瀛﹀垎鏋愯瘉瀹?/span>OsWRKY45鍙備笌妞嶇墿鑰愬共鏃卞拰鐩愬鐨勮儊杩弽搴旓紝杩欑杞熀鍥犳嫙鍗楄姤鑰愭棻鎬у拰鑰愮洂鎬х殑鏀瑰彉鏄拰ABA渚濊禆鎬х殑澶氫釜鑰愰€嗗熀鍥犵浉鍏崇殑[14]銆傝繘涓€姝ュ緱鍒扮殑OsWRKY45鍐呮簮杩囪〃杈捐浆鍩哄洜姘寸ɑ涔熻〃鐜板嚭浜嗚緝濂界殑鑰愭棻鎬э紙http://www.xtbg.ac.cn/Upload/Show_News.aspx?id=2794锛夛紝鑰屾柊杩戠殑瀹為獙杩樺彂鐜?/span>OsWRKY11鍐呮簮杩囪〃杈捐浆鍩哄洜姘寸ɑ涔熻〃鐜板嚭浜嗚€愭棻鎬?/span>[15]銆傛澶栵紝Wang绛夊彂鐜?/span>OsWRKY89鐨勮繃琛ㄨ揪澧炲己浜嗚浆鍩哄洜姘寸ɑ瀵圭传澶栬緪灏勭殑鑰愬彈鑳藉姏锛岃繖绉嶈€愬彈鑳藉姏鏄€氳繃鏀瑰杽姘寸ɑ鍙惰〃闈㈢殑铚¤川浠ｈ阿鍜屽彾缁跨礌鍒嗗竷鑰屽疄鐜扮殑[43]銆?/span>
鐩稿叧鐨勮〃杈捐氨鍒嗘瀽缁撴灉涔熸樉绀猴紝鍦ㄥ喎瀹冲鐞嗕笅15涓?/span>WRKY鍩哄洜鐨勮〃杈鹃噺鏄捐憲涓嬮檷锛岃€?/span>OsWRKY79鍩哄洜鐨勮〃杈鹃噺鏄捐憲涓婅皟锛涘湪骞叉棻澶勭悊涓?/span>19涓?/span>WRKY鍩哄洜鐨勮〃杈鹃噺鏄捐憲涓婅皟锛涘湪鐩愬澶勭悊涓?/span>25涓?/span>WRKY鍩哄洜鐨勮〃杈鹃噺鏄捐憲涓婅皟锛岃€?/span>OsWRKY74鍩哄洜鐨勮〃杈鹃噺鏄捐憲涓嬮檷锛涘彟澶栧湪婵€绱?/span>ABA鐨勫鐞嗕笅10涓?/span>WRKY鍩哄洜鐨勮〃杈鹃噺鏄捐憲涓婅皟锛?/span>6涓?/span>WRKY鍩哄洜鐨勮〃杈鹃噺鏄捐憲涓嬮檷[7]銆?/span>
2.3鍙備笌琛拌€?/span> 鍦ㄦ嫙鍗楄姤WRKY铔嬬櫧鐨勭爺绌朵腑锛岀洿鎺ユ姤閬撲笌琛拌€佺浉鍏崇殑WRKY杞綍璋冩帶鍥犲瓙鏈?/span>AtWRKY6[44]銆?/span>AtWRKY53[45]鍜?/span>AtWRKY70[46]銆傚彟澶栬繕鏈変竴浜涢棿鎺ョ殑缁撴灉鏆楃ずAtWRKY3銆?/span>AtWRKY4銆?/span>AtWRKY7鍜?/span>AtWRKY11鍙兘鍙備笌琛拌€佽皟鎺?/span>[44]銆傚埌鐩墠涓烘杩樻病鏈夌洿鎺ョ殑璇佹嵁璇佸疄姘寸ɑWRKY杞綍璋冩帶鍥犲瓙璋冩帶姘寸ɑ鐨勮“鑰佽繘绋嬶紝浣嗘槸鏈変竴浜涜瘉鎹殫绀轰簡杩欑鐩稿叧鎬с€傚湪姘寸ɑ鏃楀彾琛拌€佹棭鏈熻繘灞曚腑琚瀵艰〃杈剧殑WRKY鍩哄洜鏈?/span>OsWRKY4鍜?/span>OsWRKY82[18]锛岃€屾按绋诲湪缂洪搧鎯呭喌涓嬪彂鐢熺殑琛拌€佽繃绋嬩腑OsWRKY17琛ㄨ揪姘村钩琚笂璋?/span>[17]銆傛澶栵紝鎴戜滑瀹為獙瀹ゅ彂鐜?/span>OsWRKY23涓昏鍦ㄦ按绋昏“鑰佸彾鐗囦腑琛ㄨ揪锛屽苟淇冭繘楂樿〃杈捐浆鍩哄洜鎷熷崡鑺ョ浣撳彾鐗囧湪榛戞殫澶勭悊涓殑琛拌€佽繘灞?/span>[16]銆?/span>
2.4鍙備笌绯栦唬璋?/span> 鏈€鏃╁湪鐢滆柉涓厠闅嗗埌鐨?/span>WRKY鍩哄洜SPF1鍩哄洜鐨勫姛鑳芥秹鍙婅皟鎺х硸浠ｈ阿淇″彿閫斿緞[47]銆傚ぇ楹︿腑鐨?/span>WRKY铔嬬櫧HvWRKY2锛?/span>SUSIBA2锛夊彲浠ラ€氳繃鐗瑰紓鍦扮粨鍚堝埌寮傛瀯娣€绮夐叾1锛?/span>iso1: isoamylase1锛夊熀鍥犲惎鍔ㄥ瓙鍖哄煙鐨?/span>SURE鍏冧欢鍜?/span>W鐩掑簭鍒楋紝鍦ㄨ浆褰曟按骞充笂璋冩帶iso1鍩哄洜鐨勮〃杈?/span>[48]銆傚彈姘存潹閰歌瀵肩殑HvWRKY38铔嬬櫧璐ㄩ€氳繃鐗瑰紓鍦扮粨鍚堝埌a-娣€绮夐叾锛?/span>Amy32b锛夊熀鍥犲惎鍔ㄥ瓙鍖虹殑W鐩掑簭鍒楄瀵?/span>a-娣€绮夐叾鍩哄洜鐨勮〃杈撅紝骞堕樆鏂丹闇夌礌淇″彿閫斿緞[49]锛涜€屼笌HvWRKY38楂樺害鍚屾簮鐨?/span>OsWRKY71铔嬬櫧涔熷彲浠ラ€氳繃涓?/span>a-娣€绮夐叾鍩哄洜鍚姩瀛愬尯鐨?/span>W鐩掑簭鍒楃粨鍚堣繘鑰屾姂鍒惰丹闇夌礌璇卞鐨?/span>Amy32b鍩哄洜鐨勮〃杈?/span>[21]銆傝繘涓€姝ョ爺绌跺彂鐜版按绋荤硦绮夊眰缁嗚優涓?/span>OsWRKY24銆?/span>OsWRKY51銆?/span>OsWRKY71銆?/span>OsWRKY72绛夊熀鍥犲彈鑴辫惤閰歌瀵艰〃杈俱€傞€氳繃鐬椂琛ㄨ揪浣撶郴璇佸疄OsWRKY24鍜?/span>OsWRKY45铔嬬櫧璐ㄨ兘鏈夋晥鍦版姂鍒?/span>ABA淇″彿杞閫斿緞锛岃€?/span>OsWRKY72鍜?/span>OsWRKY77鍒欐縺娲?/span>ABA淇″彿杞閫斿緞[22, 50]銆傛洿杩涗竴姝ョ殑鐮旂┒鍒欐樉绀鸿劚钀介吀璇卞鐨勪袱涓熀鍥?/span>OsWRKY51鍜?/span>OsWRKY71鍦ㄦ按绋荤瀛愮殑绯婄矇灞傜粏鑳炰腑鑳芥湁鏁堝湴鎶戝埗璧ら湁绱犱俊鍙疯浆瀵奸€斿緞[51]銆傚彟澶栬繕鍙戠幇姘寸ɑ鎮诞缁嗚優鍦ㄧ硸楗ラタ鏉′欢涓嬩細璇卞OsWRKY62銆?/span>OsWRKY67銆?/span>OsWRKY45绛夊涓?/span>WRKY鍩哄洜鐨勮〃杈鹃噺鍙戠敓鍙樺寲[52]銆?/span>
2.5 鍙備笌褰㈡€佸缓鎴?/span> 鏈夊叧WRKY杞綍璋冩帶鍥犲瓙璋冩帶妞嶇墿褰㈡€佸缓鎴愭渶缁忓吀鐨勪緥璇佹槸AtWRKY44锛?/span>TTG2锛夊彲浠ラ€氳繃鍜屽叾瀹冪浉鍏冲熀鍥犵殑浣滅敤璋冩帶姣涚姸浣撶殑鍙戠敓涓庡惁銆佹暟鐩灏戝拰鍒嗗弶鎯呭喌锛屾殫绀?/span>AtWRKY44瀵硅〃鐨瘺鐘朵綋鐨勫舰鎬佸缓鎴愭湁璐＄尞[53]銆傝緝涓烘繁鍏ョ殑鐮旂┒杩樺彂鐜?/span>AtWRKY44杩樺弬涓庤皟鎺ф嫙鍗楄姤鏍规瘺缁嗚優鐨勫垎鍖?/span>[54]锛岃€?/span>RNAi 鎶戝埗AtWRKY75鐨勮〃杈惧悗澧炲己浜嗘嫙鍗楄姤涓嶅畾鏍瑰拰鏍规瘺鐨勭敓闀匡紝杩欑璋冩帶浣滅敤鐙珛浜庡叾鍙備笌纾蜂唬璋㈢殑鍔熻兘[55]銆傚湪姘寸ɑ褰㈡€佸缓鎴愯繃绋嬩腑鍙備笌杩涙潵鐨?/span>WRKY杞綍璋冩帶鍥犲瓙鐮旂┒鐩墠鍙湁涓や緥銆傝繃琛ㄨ揪OsWRKY31鎶戝埗浜嗘按绋讳笉瀹氭牴鐨勫舰鎴愶紝杩涜€屽奖鍝嶆按绋绘牴鐨勫舰鎬佸缓鎴?/span>[20]锛屽悓鏃惰繕鍙戠幇杩欑褰卞搷浼撮殢鐫€瀵圭敓闀跨礌鍝嶅簲鍙嶅簲鐨勫共棰勩€傛湁鍏崇敓闀跨礌鍝嶅簲鐢熺悊鍙嶅簲鐨勭爺绌跺彂鐜帮紝鐢熼暱绱犱俊鍙烽€斿緞鐨勭磰涔变細瀵艰嚧鏋佹€ц繍杈撶殑姝ｅ父杩涜锛屼細琛ㄧ幇鍑洪《绔紭鍔夸抚澶辩殑琛ㄥ瀷鍑虹幇[56]锛岃€屾垜浠殑寮傛簮楂樿〃杈?/span>OsWRKY72杞熀鍥犳嫙鍗楄姤灏卞嚭鐜颁簡杩欑被琛ㄥ瀷[19]銆傚彟澶栨垜浠繕姣旇緝浜嗗唴婧愭€ц繃琛ㄨ揪OsWRKY31姘寸ɑ鍜屽婧愭€ч珮琛ㄨ揪OsWRKY72鎷熷崡鑺ユ牴鐨勭敓闀挎儏鍐碉紝鍙戠幇杞熀鍥犳鏍潎鍑虹幇涓嶅畾鏍圭敓闀垮彈鎶戝埗鐨勫吀鍨嬭〃鍨嬶紙Fig 1锛夈€傛垜浠涓猴紝姘寸ɑ鐨勪竴浜?/span>WRKY杞綍璋冩帶鍥犲瓙寰堝彲鑳介€氳繃褰卞搷鐢熼暱绱犱俊鍙烽€斿緞杩涜€屽奖鍝嶆鐗╃殑褰㈡€佸缓鎴愶紝鑰屾绫荤爺绌剁殑娣卞叆寰堝彲鑳戒細涓板瘜鐢熼暱绱犲搷搴斿熀鍥犮€?/span>WRKY杞綍璋冩帶鍥犲瓙鍜屾鐗╂姉鐥呭熀鍥犳瀯鎴愮殑璋冩帶缃戠粶銆?/span>
3. 姘寸ɑWRKY杞綍鍥犲瓙鍙備笌骞挎硾鐨勮浆褰曡皟鎺?/span>
鍦ㄦ嫙鍗楄姤銆佺儫鑽夊拰澶ч害绛夊涓墿绉嶄腑宸茬粡鏈夌粨鏋滄樉绀?/span>WRKY铔嬬櫧鍙互鐩存帴缁撳悎鍒颁竴浜涢澏鍩哄洜鍚姩瀛愬尯鐨?/span>W鐩掍笂鑰屽鐩稿叧闈跺熀鍥犺繘琛岃浆褰曟按骞崇殑璋冩帶銆傚鎷熷崡鑺ョ殑WRKY铔嬬櫧鍙互鐩存帴缁撳悎鍦ㄦ姉鐥呯浉鍏冲熀鍥?/span>PR1鍜?/span>NPR1鐨勫惎鍔ㄥ瓙鍖哄鍏惰繘琛岃皟鎺ц繘鑰屽奖鍝嶆鐗╃殑鍏嶇柅鍙嶅簲[25]锛屽ぇ楹︾殑WRKY铔嬬櫧鍙互鐩存帴閫氳繃缁撳悎鍚姩瀛愬尯璋冩帶ISO1鍩哄洜鍜?/span>Amy32b鍩哄洜鐨勮〃杈炬按骞筹紝浠庤€屽奖鍝嶇硸浠ｈ阿閫斿緞[48, 57]銆傛湁鍏虫按绋?/span>WRKY鍩哄洜璋冩帶鐩稿叧鎶楃梾鍩哄洜鐨勭洿鎺ヨ瘉鎹洰鍓嶈繕娌℃湁锛屼絾鏄敱浜?/span>OsNH1鍜?/span>AtNPR1鐨勯珮搴﹀悓婧愭€?/span>[58]锛屾湁鍏?/span>OsWRKY45鍜?/span>OsWRKY71鐨勭爺绌跺凡缁忔殫绀哄OsNH1鍩哄洜琛ㄨ揪姘村钩璋冩帶鐨勪袱绉嶄綔鐢ㄦā寮?/span>[13, 28]銆傛瘮杈?/span>OsWRKY71鍜?/span>OsWRKY45鐨勭浉鍏崇粨鏋滃悗锛屾垜浠彂鐜拌繖涓や釜姘寸ɑ鍩哄洜寰堝彲鑳介兘澶勪簬SA鐩稿叧鐨勪俊鍙烽€氳矾涓紝浣嗗墠鑰呭鎶楃梾淇″彿鐨勮皟鎺т緷璧栦簬NH1鑰屽悗鑰呬笉渚濊禆銆傞櫎浜嗗湪楂樿〃杈?/span>OsWRKY71鐨勮浆鍩哄洜姘寸ɑ涓娴嬪埌NH1鍩哄洜鍜?/span>PR1鍩哄洜鏄捐憲澧炲己浠ュ锛屽湪OsWRKY12鍜?/span>OsWRKY13涓ょ杞熀鍥犳按绋讳腑涔熷彂鐜?/span>NH1鍩哄洜鍜?/span>PR1鍩哄洜鏄捐憲澧炲己[26, 27]锛岃繖浜涚粨鏋滈兘璇存槑杩欎笁涓按绋?/span>WRKY鍩哄洜寰堝彲鑳藉浜庢姉鐥呬俊鍙烽€斿緞涓?/span>NH1鐨勪笂娓搞€傛澶栵紝杩樻湁鎶ラ亾鍦ㄧ爺绌跺姛鑳藉熀鍥犵殑杩囩▼涓彂鐜板叾鍚姩瀛愬尯鏈夎緝澶?/span>W鐩掞紝寰堝彲鑳藉彈鍒?/span>WRKY鍩哄洜鐨勮皟鎺с€傚OsWRKY71鍙互缁撳悎鍦?/span>Amy32b鍩哄洜鍚姩瀛愬尯鐨?/span>W鐩掍笂[51]銆傛按绋诲€嶅崐钀滅幆鍖栭叾鍩哄洜OsDTC2鐨勫惎鍔ㄥ瓙鍖哄氨鏈?/span>6涓?/span>W鐩掞紙−1709 bp 鑷?/span> −1450 bp锛夛紝浣滆€呮帹娴嬫按绋?/span>WRKY杞綍璋冩帶鍥犲瓙瀵瑰叾鍏锋湁璋冩帶浣滅敤[59]銆傛渶鍚庯紝涓€浜涚爺绌跺垯鍏虫敞浜嗚皟鎺?/span>WRKY鍩哄洜鐨勪笂娓稿熀鍥狅紝鍙戠幇OsWRKY13寰堝彲鑳藉彈PRE2鍜?/span>PRE4铔嬬櫧鐨勮皟鎺э紝杩欑璋冩帶涔熷缓绔嬪湪OsWRKY13鍚姩瀛愬尯鐨勭壒寮傚簭鍒椾笂[37]銆?/span>
4. 姘寸ɑWRKY杞綍璋冩帶鍥犲瓙鍜屾嫙鍗楄姤WRKY杞綍璋冩帶鍥犲瓙鍔熻兘鐨勭浉浜掑嵃璇佸強鐮旂┒灞曟湜
鐢熺墿淇℃伅瀛︾殑鐮旂┒杩囩▼涓紝鏃犺鏄涓墿绉嶉棿WRKY搴忓垪鐨勬瘮杈冿紝杩樻槸鍗曚釜鐗╃鍐?/span>WRKY搴忓垪鐨勮繘鍖栨爲鏋勫缓锛岄兘浠?/span>WRKY鍩哄洜鐨勫垎瀛愯繘鍖栨€濇兂涓哄熀纭€銆備粠浣庣瓑妞嶇墿鑻旇棑锛?/span>Physcomitrella patens锛夊拰钑ㄧ被锛?/span>Ceratopteris richardii锛夊埌楂樼瓑妞嶇墿鎷熷崡鑺ュ拰姘寸ɑ閮藉彂鐜?/span>WRKY鍩哄洜[60]锛屽彟澶栧湪鍘熺敓鐢熺墿钃濇皬璐剧闉瘺铏紙Giardia lamblia锛夊拰缁嗚優绮樿川闇夎弻鐩樺熀缃戞焺鑿岋紙Dictyostelium discoideum锛夊熀鍥犵粍涓篃鍙戠幇WRKY鍩哄洜[23]锛屽洜鑰?/span>WRKY铔嬬櫧涔熷氨涓嶈兘绉颁箣涓烘鐗╃壒鏈夌殑浜嗐€傛湁鎰忔€濈殑鏄紝鏃犺鏄湪钃濇皬璐剧闉瘺铏拰鐩樺熀缃戞焺鑿屼腑杩樻槸鍦ㄨ嫈钘撳拰钑ㄧ被妞嶇墿涓紝鍙戠幇鐨?/span>WRKY铔嬬櫧鍩烘湰涓婇兘鍚湁涓や釜WRKY缁撴瀯鍩燂紝鑰屽湪鎷熷崡鑺ワ紙66/72锛夊拰姘寸ɑ锛?/span>89/103锛変腑鐨勫ぇ澶氭暟WRKY铔嬬櫧鍒欏彧鍚湁涓€涓?/span>WRKY缁撴瀯鍩熴€傝繖涓€鍙戠幇鏆楃ずWRKY铔嬬櫧鐨勮繘鍖栧叧绯绘槸鐢卞惈鏈変袱涓?/span>WRKY缁撴瀯鍩熺殑铔嬬櫧鍚戝惈鏈変竴涓粨鏋勫煙鐨勮泲鐧借繘鍖栥€傚熀浜庤繖涓€鎬濇兂锛屾垜浠彲浠ュ湪澶氫釜鐗╃闂村缓绔?/span>WRKY铔嬬櫧鐨勮繘鍖栧叧绯汇€傛垜浠涓洪€氳繃搴忓垪鐨勬瘮瀵瑰彲浠ラ儴鍒嗗湴棰勭ず涓や釜鐗╃闂村瓨鍦ㄨ繎缂樺叧绯荤殑WRKY铔嬬櫧鍏锋湁鐩镐技鐨勭敓鐗╁鍔熻兘锛岃€岃繖绉嶅姛鑳界浉浼兼€х殑纭畾闇€瑕佸垎瀛愮敓鐗╁鍜岄仐浼犲鐨勮繘涓€姝ヨ瘉鏄庛€傝€岀洰鍓嶅凡缁忔湁鎶ラ亾鎻愬強鎷熷崡鑺?/span>WRKY铔嬬櫧銆佹按绋?/span>WRKY铔嬬櫧浠ュ強鍏朵粬鐗╃鐨?/span>WRKY铔嬬櫧鐨勫姛鑳藉瓨鍦ㄤ竴瀹氱殑鐩镐技鎬с€傜粨鍚堟湰瀹為獙瀹ょ殑鍙戠幇锛屾垜浠妸鐩稿叧鐨?/span>WRKY铔嬬櫧鍦ㄨ〃2涓垪涓俱€?/span>
灏辩洰鍓嶇殑鐮旂┒鑰岃█锛屽凡缁忓彂鐜版鐗╃殑WRKY鍩哄洜鍙楀绉嶇敓鐗╅€嗗鍥犲瓙鍖呮嫭鐥呮瘨銆佺粏鑿屻€佺湡鑿屻€佸嵉鑿屻€佺嚎铏€佷互鍙婃槅铏瓑[60, 61, 62]鍜屽绉嶉潪鐢熺墿閫嗗鍥犲瓙濡備綆娓┿€侀珮娓┿€佸共鏃便€侀珮鐩愩€佸厜锛堣緪灏勶級銆侀珮婀裤€佷互鍙婃崯浼ょ瓑璇卞[5, 63]銆傚悓鏃跺彂鐜版鐗╃殑WRKY鍩哄洜鍦ㄤ笉鍚屽彂鑲查樁娈靛拰鐢熺悊鐘跺喌濡備紤鐪犮€佸紑鑺便€佽儦鑳庡彂鑲层€佽“鑰併€侀ゥ楗跨瓑鎯呭喌琛ㄨ揪[4, 64]銆傚彟澶栵紝妞嶇墿鐨?/span>WRKY鍩哄洜琛ㄨ揪姘村钩鍙椾笉鍚岀殑婵€绱犲拰淇″彿鍒嗗瓙鍖呮嫭鑴辫惤閰搞€佽丹闇夌礌銆佺敓闀跨礌銆佹按鏉ㄩ吀銆佽寜鑾夐吀銆佷互鍙婁竴姘у寲姘拰鍙屾哀姘村鐞嗚€屽彉鍖?/span>[51, 61, 63, 65]锛屽苟骞挎硾鍙備笌浜嗘按鏉ㄩ吀銆佽寜鑾夐吀銆佽劚钀介吀銆佽丹闇夌礌銆佺敓闀跨礌绛夌浉鍏崇殑淇″彿缃戠粶[11, 22, 65]銆傜洰鍓嶈澶氱爺绌剁粨鏋滆瘉瀹烇紝涓嶅悓鐨勬縺绱犱俊鍙烽€斿緞瀛樺湪骞挎硾鐨勪氦鍙?/span>[66]锛岀壒鍒槸妞嶇墿鎶楃梾鍙嶅簲锛?/span>SA閫斿緞锛夈€佹鐗╄€愰€嗗弽搴旓紙ABA鍜?/span>ROS锛変笌鍙戣偛锛?/span>GA鍜?/span>Auxin锛変俊鍙烽€斿緞鐨勪氦鍙夈€傚鎴戜滑鍏嬮殕鐨?/span>WRKY鍩哄洜OsWRKY45鍜?/span>OsWRKY72[14, 19]涓庢姤閬撶殑OsWRKY71[33, 51 ]銆?/span>OsWRKY31[20]鍜?/span>OsWRKY89[30]锛屽畠浠殑琛ㄨ揪鍙樺寲涓嶄粎褰卞搷杞熀鍥犳鐗╃殑鑰愰€嗘€э紝鑰屼笖涔熸敼鍙樹簡杞熀鍥犳鐗╃殑鎶楃梾鎬ф垨鐢熼暱绱犵浉鍏崇殑鐢熼暱鍙戣偛銆傜敱姝ゆ垜浠涓猴紝瀵规按绋?/span>WRKY杞綍璋冩帶鍥犲瓙鐨勭爺绌跺簲璇ヨ繍鐢ㄩ珮琛ㄨ揪鍜岄珮鎶戝埗涓ゅ绯荤粺鍏ㄩ潰鐨勮€冨療杞熀鍥犳鐗╃殑鐢熺悊鍙戣偛銆佽€愰€嗘€у彉鍖栧拰鎶楃梾鎬у彉鍖栵紝杩涜€屾洿鍔犵‘鍒囩殑璇犻噴璇ュ熀鍥犵殑鍔熻兘銆?/span>
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