于好強(qiáng)，孫福艾，馮文奇，路風(fēng)中，李晚忱，付鳳玲

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轉(zhuǎn)錄因子BES1/BZR1調(diào)控植物生長(zhǎng)發(fā)育及抗逆性

于好強(qiáng)，孫福艾，馮文奇，路風(fēng)中，李晚忱，付鳳玲

四川農(nóng)業(yè)大學(xué)玉米研究所，農(nóng)業(yè)部西南玉米生物學(xué)與遺傳育種重點(diǎn)實(shí)驗(yàn)室，溫江 611130

油菜素內(nèi)酯(brassinosteroid, BR)是植物特有的甾體激素，在植物生長(zhǎng)發(fā)育及逆境應(yīng)答過(guò)程中起重要作用。轉(zhuǎn)錄因子BES1/BZR1(BRI1 EMS SUPPRESSOR 1/BRASSINAZOLE RESISTANT 1)是BR信號(hào)轉(zhuǎn)導(dǎo)的核心成員，被BR信號(hào)激活后，結(jié)合到下游靶基因啟動(dòng)子區(qū)的E框(CANNTG)或BRRE元件(CGTGT/CG)，調(diào)節(jié)靶基因表達(dá)。除介導(dǎo)BR信號(hào)，BES1/BZR1還參與脫落酸、赤霉素及光等信號(hào)轉(zhuǎn)導(dǎo)途徑，協(xié)同調(diào)控植物的生長(zhǎng)發(fā)育。最新研究發(fā)現(xiàn)，BES1/BZR1還參與調(diào)控植物的抗逆性。本文對(duì)轉(zhuǎn)錄因子BES1/BZR1通過(guò)信號(hào)轉(zhuǎn)導(dǎo)調(diào)控植物生長(zhǎng)發(fā)育和抗逆性分子機(jī)制的新近研究進(jìn)展進(jìn)行了綜述，以期為相關(guān)研究提供參考。

油菜素內(nèi)酯；生長(zhǎng)發(fā)育；信號(hào)轉(zhuǎn)導(dǎo)；抗逆性；BES1/BZR1轉(zhuǎn)錄因子

油菜素內(nèi)酯(brassinosteroid, BR)是植物特有的甾體激素，在生長(zhǎng)發(fā)育及環(huán)境脅迫應(yīng)答中起重要作用，其生理活性遠(yuǎn)高于生長(zhǎng)素(auxin, IAA)、赤霉素(gibberellins, GA)、細(xì)胞分裂素(cytokinin, CTK)、脫落酸(abscisic acid, ABA)和乙烯(ethylene, ET)[1,2]。BR合成基因過(guò)量表達(dá)或缺失對(duì)植物生長(zhǎng)發(fā)育及產(chǎn)量、品質(zhì)等農(nóng)藝性狀育均產(chǎn)生嚴(yán)重影響[3~6]。BR信號(hào)轉(zhuǎn)導(dǎo)被阻斷的植物則顯現(xiàn)矮化、開(kāi)花延遲、早衰等缺陷表型[7,8]。

BR被細(xì)胞膜上BRASSINOSTEROID INSEN-SITIVE 1 (BRI1)及BRI1-ASSOCIATED RECEPTOR KINASE1 (BAK1)等激酶接受后，通過(guò)信號(hào)轉(zhuǎn)導(dǎo)激活轉(zhuǎn)錄因子BRI1 EMS SUPPRESSOR 1 (BES1)及其同源蛋白BRASSINAZOLE RESISTANT 1 (BZR1)的活性[9,10]。BES1與BZR1氨基酸序列相似性達(dá)88%，N端結(jié)構(gòu)域相似性高達(dá)97%[11]，編碼基因以家族形式存在，本課題組在前期研究中將其統(tǒng)一命名為BES1/BZR1[12]。被BR信號(hào)激活后，BES1/BZR1直接或與其他轉(zhuǎn)錄因子一起結(jié)合到生長(zhǎng)發(fā)育相關(guān)基因啟動(dòng)子的E框(CANNTG)或BRRE元件(CGTGT/ CG)，調(diào)節(jié)這些基因的表達(dá)[13~15]。例如，BES1/BZR1抑制葉腋分生組織發(fā)育基因表達(dá)，可促進(jìn)小穗發(fā)育，增加水稻產(chǎn)量[16]。BES1/BZR1調(diào)節(jié)根尖分生組織發(fā)育相關(guān)基因表達(dá)，進(jìn)而調(diào)控根發(fā)育[4,17~19]。除介導(dǎo)BR信號(hào)，BES1/BZR1還參與ABA、GA及光等信號(hào)轉(zhuǎn)導(dǎo)途徑，調(diào)控植物的生長(zhǎng)發(fā)育以及抗凍、耐旱、抗病等抗逆性。

本文對(duì)轉(zhuǎn)錄因子BES1/BZR1通過(guò)信號(hào)轉(zhuǎn)導(dǎo)調(diào)控植物生長(zhǎng)發(fā)育和抗逆性分子機(jī)制的新近研究進(jìn)展進(jìn)行了綜述，以期為相關(guān)研究提供參考。

1 BES1/BZR1介導(dǎo)BR信號(hào)轉(zhuǎn)導(dǎo)

2002年，Wang等[20]利用EMS誘變篩選到一個(gè)BR合成抑制突變體()，圖位克隆獲得基因，該基因編碼核蛋白且受BR誘導(dǎo)。同年，Yin等[21]利用EMS誘變篩選到BR受體抑制因子BES1，受BR誘導(dǎo)并在細(xì)胞核中積累。后經(jīng)證實(shí)BES1是一個(gè)BZR1類(lèi)蛋白(BZR1- like protein)，二者具有高度的序列相似性，N端均有一個(gè)核定位信號(hào)(NLS)，C端均有22~24個(gè)絲氨酸或蘇氨酸殘基(S/TXXXS/T)，該殘基是BIN2、GSK-3等激酶磷酸化位點(diǎn)，磷酸化后進(jìn)入細(xì)胞質(zhì)被14-3-4蛋白降解[16,20,21]。直至2005年，Yin等[10]進(jìn)一步證實(shí)BES1/BZR1是植物中特有的新一類(lèi)轉(zhuǎn)錄因子，也是BR信號(hào)轉(zhuǎn)導(dǎo)途徑的唯一轉(zhuǎn)錄因子。細(xì)胞膜上的BRI1、BKI1和BAK1等激酶接受BR信號(hào)后，自身磷酸化并催化BRASSINOSTEROID-SIGNALLING KINASE1 (BSK)和CONSTITUTIVE DIFFERENTIAL GROWTH1 (CDG1)磷酸化，BSK與CDG1進(jìn)一步磷酸化BRI1-SUPPRESSOR1 (BSU1)，BSU1催化BRASSINOSTEROID INSENSITIVE2 (BIN2)去磷酸化，導(dǎo)致其自身被蛋白酶體降解，削弱BIN2對(duì)BES1/BZR1的磷酸化從而使其活性增加[9,10,21~23]，BES1/BZR1通過(guò)調(diào)節(jié)下游靶基因的表達(dá)，調(diào)控植物的生長(zhǎng)發(fā)育(圖1A)。

BES1/BZR1成員N端均有一個(gè)bHLH結(jié)構(gòu)域，可特異性結(jié)合到靶基因啟動(dòng)子區(qū)的E框或BRRE元件[10,24~26]。此外，多數(shù)BES1/BZR1成員均含有能被BIN2等激酶磷酸化的絲氨酸(serine, S)富集位點(diǎn)，個(gè)別成員包含一個(gè)與蛋白穩(wěn)定性緊密相關(guān)的脯氨酸(proline, P)、谷氨酸(glutamic acid, E)、絲氨酸(serine, S)和蘇氨酸(threonine, T)富集區(qū)(PEST基序)[10]。目前，擬南芥()和水稻()BES1/BZR1基因家族已被全部鑒定：擬南芥AtBES1/BZR1基因家族有6個(gè)成員，且功能存在部分冗余[10,20,21]；水稻OsBES1/BZR1基因家族有4個(gè)成員[27]。玉米()ZmBES1/BZR1基因家族有11個(gè)成員[23,28]。此外，從白菜(ssp)、棉花()、油菜()和桉樹(shù)()中均鑒定出多個(gè)BES1/ BZR1基因家族成員[29~33](表1)。進(jìn)一步研究證實(shí)，BES1/BZR1基因家族成員通過(guò)不同信號(hào)途徑調(diào)控植物生理代謝過(guò)程，進(jìn)而調(diào)控植物生長(zhǎng)發(fā)育及逆境響應(yīng)。

：未知過(guò)程；：相互作用；：促進(jìn)作用；：抑制作用

A：BES1/BZR1介導(dǎo)的BR信號(hào)轉(zhuǎn)導(dǎo)；B：BES1/BZR1參與的ABA信號(hào)途徑；C：BES1/BZR1參與的GA信號(hào)途徑；D：BES1/BZR1參與的光信號(hào)途徑；E：BES1/BZR1調(diào)控逆境應(yīng)答途徑；F：BES1/BZR1參與的生長(zhǎng)素、乙烯及其他信號(hào)途徑。BR：油菜素內(nèi)酯；BES1/BZR1：轉(zhuǎn)錄因子；BKI1、BRI1、BAK1、BSK1及CDG1：蛋白激酶；BSU1：BRI1抑制因子；ABA：脫落酸；GA：赤霉素；PP2C：2C型絲氨酸蘇//氨酸蛋白激酶；PP2A：2A型絲氨酸/蘇氨酸蛋白激酶；PYL：ABA受體；BIN2：磷酸激酶；ABI3與ABI5：ABA響應(yīng)的bZIP轉(zhuǎn)錄因子；DELLA：赤霉素負(fù)調(diào)控轉(zhuǎn)錄因子；SINAT與COP1：E3泛素連接酶；GATA2與HY5：光形態(tài)建成相關(guān)轉(zhuǎn)錄因子；UVR8：紫外光受體；PIF4：光敏色素互作因子；CRY：隱花色素。RD26與WRKY26：干旱相關(guān)轉(zhuǎn)錄因子；REF：乙烯應(yīng)答因子；MEK6：促細(xì)胞分裂原活化蛋白激酶；P：磷。

2 BES1/BZR1參與ABA信號(hào)途徑

ABA是植物體內(nèi)重要激素之一，通過(guò)其直接受體PYL (pyrabactin resistance 1-like protein)、第二信使2C型蛋白磷酸酶(PP2C)及第三信使蔗糖非酵解型蛋白激酶(SnRK)向下游進(jìn)行信號(hào)傳遞，在植物生長(zhǎng)發(fā)育及抗逆過(guò)程中扮演重要角色，如衰老、抗旱、耐鹽等[34~36]。研究發(fā)現(xiàn)，在突變體中，BZR1結(jié)合到ABA誘導(dǎo)型轉(zhuǎn)錄因子ABA INSENSITIVE 5 (ABI5)編碼基因的啟動(dòng)子，抑制其表達(dá)，因而抑制突變體對(duì)ABA誘導(dǎo)的應(yīng)答[37]。同時(shí)，BES1抑制ABA調(diào)節(jié)的轉(zhuǎn)錄因子ABI3編碼基因的表達(dá)，進(jìn)而抑制ABI3對(duì)下游ABI5轉(zhuǎn)錄因子的激活，致使ABA信號(hào)轉(zhuǎn)導(dǎo)受阻，表現(xiàn)為苗期發(fā)育遲緩[38,39]。

表1 已鑒定的不同植物BES1/BZR1基因家族成員

此外，外源ABA不僅誘導(dǎo)基因表達(dá)，而且誘導(dǎo)BES1蛋白磷酸化，使其穩(wěn)定性降低，從而抑制BR信號(hào)轉(zhuǎn)導(dǎo)，此過(guò)程依賴(lài)于ABA第二信使PP2C成員ABI1和ABI2[12,29,40,41]。最新研究表明，ABI1、ABI2與BIN2激酶互作后催化BIN2去磷酸化，從而調(diào)控BES1活性。ABA還可促進(jìn)BIN2磷酸化并抑制ABI2的活性[42]。在ABA存在時(shí)，BIN2磷酸化ABI5使其穩(wěn)定性增強(qiáng)，調(diào)控種子發(fā)育過(guò)程[43]。在大豆()中，PP2C-1與GmBZR1直接互作，催化GmBZR1去磷酸化以增強(qiáng)GmBZR1活性，促進(jìn)種子大小相關(guān)基因()、()和()等表達(dá)，調(diào)控種子的大小與重量[44,45]。BZR1也可結(jié)合到ABA受體PYL6編碼基因的啟動(dòng)子區(qū)，上調(diào)表達(dá)，從而參與PYL6介導(dǎo)的ABA信號(hào)轉(zhuǎn)導(dǎo)[46]。研究還發(fā)現(xiàn)，BZR1的PEST結(jié)構(gòu)域與蛋白磷酸酶2A(PP2A)的B亞基直接互作，使BZR1被PP2A去磷酸化，激活BZR1介導(dǎo)的BR信號(hào)途徑，調(diào)控植物的生長(zhǎng)發(fā)育[47](圖1B)。

3 BES1/BZR1參與GA信號(hào)途徑

作為植物體內(nèi)重要激素之一，GA在種子萌發(fā)、細(xì)胞分裂、胚珠形成等生長(zhǎng)發(fā)育過(guò)程中起關(guān)鍵作用[48,49]。研究發(fā)現(xiàn)，在BR缺失的突變體中，GA合成關(guān)鍵基因表達(dá)顯著下調(diào)，而在突變體中，基因表達(dá)顯著上調(diào)。同時(shí)，在和突變體中，基因均受BR誘導(dǎo)，表達(dá)顯著上調(diào)。有研究表明，基因啟動(dòng)子區(qū)不含BES1/BZR1轉(zhuǎn)錄因子的結(jié)合位點(diǎn)。研究人員通過(guò)電泳遷移實(shí)驗(yàn)(electroph-oretic mobility shift assays, EMSA)和染色質(zhì)免疫共沉淀(chromatin immunoprecipitation, Chip)實(shí)驗(yàn)證實(shí)，BES1/BZR1可結(jié)合基因啟動(dòng)子區(qū)一個(gè)非E-Box且長(zhǎng)度為12 bp的基序(Motif)[50,51]。此外，GA信號(hào)負(fù)調(diào)控因子DELLA家族蛋白(RGA、GAI、RGL1、RGL2和RGL3)可以和BES1/BZR1結(jié)合，阻止BES1/BZR1與靶基因的結(jié)合[50,52~55]。這些研究結(jié)果證實(shí)，DELLA蛋白降解可促使BES1/BZR1活性增強(qiáng)，BES1/BZR1結(jié)合GA合成相關(guān)基因啟動(dòng)子，使其表達(dá)上調(diào)，促進(jìn)GA積累。此外，GA可通過(guò)PP2A促進(jìn)BES1/BZR1的去磷酸化[54]。

在水稻中，BR誘導(dǎo)GA合成基因表達(dá)，促使GA積累。外源GA又抑制BR合成及其信號(hào)轉(zhuǎn)導(dǎo)。進(jìn)一步研究表明，GA合成關(guān)鍵基因、、和的啟動(dòng)子均包含CATGTG、BRRE或G-box元件。BES1/BZR1與這類(lèi)元件直接結(jié)合，調(diào)節(jié)下游基因的表達(dá)，進(jìn)而影響GA合成[56]。在番茄()中過(guò)表達(dá)，GA合成關(guān)鍵酶之一的酮戊二酸脫氫酶2 (2-ODD2)蛋白水平在果實(shí)成熟期顯著增加，致使GA顯著積累促進(jìn)果實(shí)成熟[57]。水稻OsBZR1能夠促進(jìn)miR396d的積累，調(diào)控其靶基因()的表達(dá)，通過(guò)參與的GA合成及信號(hào)轉(zhuǎn)導(dǎo)途徑，調(diào)控水稻株高及葉夾角等形態(tài)建成[58](圖1C)。

4 BES1/BZR1參與光信號(hào)途經(jīng)

光是植物光合作用的能量之源，在調(diào)控植物生長(zhǎng)發(fā)育中起關(guān)鍵作用，如光信號(hào)參與調(diào)控種子萌發(fā)、光形態(tài)建成和開(kāi)花等[59]。轉(zhuǎn)錄因子GATA2、HY5正向調(diào)控植物光形態(tài)建成并受光誘導(dǎo)積累，黑暗促使其降解。研究發(fā)現(xiàn)，被BR激活的BZR1直接與GATA2互作，抑制轉(zhuǎn)錄，調(diào)控?cái)M南芥幼苗下胚軸伸長(zhǎng)[60,61]。黑暗條件下，HY5能特異地結(jié)合BZR1，抑制BZR1與子葉開(kāi)閉相關(guān)基因的結(jié)合能力，調(diào)控光形態(tài)建成[61]。光敏色素互作因子(phytochrome interacting factor，PIF)是一類(lèi)bHLH轉(zhuǎn)錄因子，在黑暗條件下，PIF大量積累，促進(jìn)植物暗形態(tài)建成，但在光照條件下，PIF發(fā)生磷酸化后降解，促進(jìn)植物光形態(tài)建成[62,63]。研究發(fā)現(xiàn)，BES1/BZR1與PIF 4相互作用，形成異源二聚體后作用于共同靶基因，其中80%靶基因受光誘導(dǎo)參與光形態(tài)建成[11]。此外，BZR1與PIF4共同作用的靶基因還受GA誘導(dǎo)，GA促進(jìn)細(xì)胞伸長(zhǎng)的過(guò)程依賴(lài)于BZR1和PIF4。DELLA- BZR1-PIF4復(fù)合體調(diào)控下游靶基因paclobutrazol resistance家族(PREs)表達(dá)，促進(jìn)細(xì)胞伸長(zhǎng)，調(diào)控光形態(tài)建成[11,53]。在高溫條件下，BZR1和PIF4相互作用，調(diào)控植物熱形態(tài)建成[64]。

最近研究發(fā)現(xiàn)，去磷酸化的BES1可與紫外光受體UVR8 (UV RESISTANCE LOCUS 8)互作，二者的復(fù)合體受紫外光(UV-B)誘導(dǎo)，并在細(xì)胞核大量積累。同時(shí)，UV-B不僅抑制BES1靶基因表達(dá)，其受體UVR8又抑制BES1與DNA的結(jié)合作用，最終控制植物光形態(tài)建成過(guò)程[65]。在藍(lán)光條件下，其受體隱花色素(cryptochrome, CRY) CRY1和CRY2特異性與去磷酸化的BES1互作，抑制BES1與DNA結(jié)合活性及其靶基因表達(dá)，最終抑制下胚軸伸長(zhǎng)[66]。

綜上所述，BES1/BZR1參與光信號(hào)途徑調(diào)控植物的形態(tài)建成過(guò)程。此外，還有研究發(fā)現(xiàn)，植物體內(nèi)BES1/BZR1的磷酸化狀態(tài)及穩(wěn)定性也受光信號(hào)調(diào)控。黑暗條件促進(jìn)BES1/BZR1去磷酸化以增強(qiáng)活性，而光照條件下，大多數(shù)BZR1被BIN2磷酸化以保持失活狀態(tài)[61,67,68]。Kim等[68]研究發(fā)現(xiàn)，黑暗條件下，E3泛素連接酶COP1催化磷酸化后的BZR1降解，去磷酸化的BZR1積累。同時(shí)，光照條件可誘導(dǎo)E3泛素連接酶SINAT積累，SINAT泛素化BES1促使其降解。相反，黑暗條件抑制SINAT的積累，從而阻止BES1降解[69](圖1D)。

5 BES1/BZR1調(diào)控植物抗逆性

BES1/BZR1除調(diào)控植物生長(zhǎng)發(fā)育外，在響應(yīng)生物和非生物逆境脅迫過(guò)程中也起重要作用。Guo等[70]研究發(fā)現(xiàn)，BES1/BZR1調(diào)控硫代糖苷合酶基因的表達(dá)，促進(jìn)硫代糖苷合成，而硫代糖苷在植物與食草動(dòng)物或與微生物互作中起重要作用。隨后，Miyaji等[71]發(fā)現(xiàn)BZR1可能參與茉莉酸信號(hào)途徑，增強(qiáng)植物抗蟲(chóng)能力。此外，研究表明，病原物相關(guān)分子模式(pathogen-associated molecular pattern, PAMP)感知可促進(jìn)BES1磷酸化，在PAMP誘導(dǎo)的免疫反應(yīng)(PAMP-triggered immunity, PTI)過(guò)程中，BES1作為病原菌誘導(dǎo)的MITOGEN-ACTIVATED PROTEIN KINASE 6 (MPK6)的直接底物被其磷酸化，參與調(diào)控植物對(duì)病菌的免疫反應(yīng)[72]。

Singh等[73]研究發(fā)現(xiàn)，低磷脅迫促進(jìn)BES1/ BZR1由細(xì)胞核向細(xì)胞質(zhì)轉(zhuǎn)移，低磷條件下，BES1/ BZR1顯著積累，維持根系正常生長(zhǎng)，賦予擬南芥對(duì)低磷脅迫的耐受性。研究表明，BES1/BZR1可促進(jìn)轉(zhuǎn)錄因子CBF (C-repeat binding factor)、WRKY6以及ABA受體PYL6等編碼基因的表達(dá)，并與WRKY54轉(zhuǎn)錄因子直接互作，正調(diào)控?cái)M南芥耐寒性，但負(fù)調(diào)控其耐旱性[46,74]。研究還發(fā)現(xiàn)，BES1/BZR1與NAC轉(zhuǎn)錄因子家族的RD26存在拮抗關(guān)系，BES1/ BZR1結(jié)合到基因啟動(dòng)子，抑制表達(dá)，而RD26蛋白又與BES1/BZR1蛋白結(jié)合，抑制RD26干旱應(yīng)答調(diào)節(jié)功能[75]。同時(shí)，在干旱和低碳脅迫下，BES1與泛素受體DSK2互作而被降解，參與脅迫誘導(dǎo)的自噬反應(yīng)過(guò)程，調(diào)控植物適應(yīng)逆境[76]。

此外，在擬南芥、油菜和桉樹(shù)中，BES1/BZR1基因的表達(dá)受鹽、干旱、熱和冷等脅迫的誘導(dǎo)或抑制[29,32,33]，表明該基因家族參與這些逆境脅迫響應(yīng)過(guò)程(圖1E)。

6 BES1/BZR1參與的信號(hào)轉(zhuǎn)導(dǎo)網(wǎng)絡(luò)

BES1/BZR1是BR信號(hào)轉(zhuǎn)導(dǎo)途徑特異的轉(zhuǎn)錄因子，通過(guò)介導(dǎo)BR信號(hào)調(diào)控植物生長(zhǎng)發(fā)育。但近年來(lái)研究發(fā)現(xiàn)，BES1/BZR1在ABA、GA、光及逆境信號(hào)中也發(fā)揮著重要作用，并且還參與IAA、ET等信號(hào)途徑。如BZR1直接與生長(zhǎng)素誘導(dǎo)基因、的啟動(dòng)子結(jié)合而抑制其表達(dá)，從而影響生長(zhǎng)素的合成，調(diào)控植物生長(zhǎng)發(fā)育[77]。BES1/BZR1通過(guò)下調(diào)乙烯合成關(guān)鍵酶基因、和表達(dá)，抑制乙烯合成，而且與乙烯響應(yīng)因子ERF72互作調(diào)控其下游基因表達(dá)，最終影響植物生長(zhǎng)發(fā)育過(guò)程[4,78](圖1F)。綜上所述，BES1/BZR1參與多種信號(hào)途徑，調(diào)控植物生長(zhǎng)發(fā)育及逆境應(yīng)答，其功能和作用機(jī)制表現(xiàn)出多樣性。但是，BES1/BZR1調(diào)控植物響應(yīng)逆境脅迫方面的研究還不夠深入，除擬南芥外，在作物及其他植物中BES1/BZR1抗逆功能研究尚未見(jiàn)報(bào)道。因此，本文將BES1/BZR1參與的信號(hào)轉(zhuǎn)導(dǎo)網(wǎng)絡(luò)進(jìn)行了歸納總結(jié)(圖1)，以期為后續(xù)相關(guān)研究提供參考。

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The BES1/BZR1 transcription factors regulate growth, development and stress resistance in plants

Haoqiang Yu, Fuai Sun, Wenqi Feng, Fengzhong Lu, Wanchen Li, Fengling Fu

Brassinosteroid (BR) is a class of plant-specific steroidal hormone and plays vital roles in plant growth, developmental and stress response. As the core component of BR signaling, the BES1/BZR1 transcription factors are activated by the BR signal, bind to the E-box (CANNTG) or BRRE element (CGTGT/CG) enriched in the promoter of downstream target genes and regulate their expression. Besides BR signal transduction, BES1/BZR1s are also involved in other signaling pathways such as abscisic acid, gibberellin and light to co-regulate plant growth and development. Recently, BES1/BZR1s were found to be related to stress resistance. In this review, we summarize recent advances of molecular mechanism of the BES1/BZR1 transcription factors regulating plant growth, development and stress resistance through signal transduction to provide a reference for related researches.

brassinosteroid; growth and development; signal transduction; stress resistance; BES1/BZR1 transcription factors

2018-09-07;

2019-02-20

四川省科技計(jì)劃應(yīng)用基礎(chǔ)項(xiàng)目(編號(hào)：2018JY0470)資助[Supported by the Sichuan Science and Technology Program (No.2018JY0470)]

于好強(qiáng)，博士，講師，研究方向：植物分子生物學(xué)。E-mail: yhq1801@sicau.edu.cn

付鳳玲，博士，教授，研究方向：玉米遺傳育種與生物技術(shù)。E-mail: ffl@sicau.edu.cn

10.16288/j.yczz.18-253

2019/2/25 15:23:43

URI: http://kns.cnki.net/kcms/detail/11.1913.R.20190225.1523.004.html

(責(zé)任編委: 張憲省)