李魯華,王忠妮,王文新,王懷玉,任明見,徐如宏*

植物ABI5轉(zhuǎn)錄因子的研究進展

李魯華1,2,王忠妮3,王文新1,王懷玉1,任明見1,2,徐如宏1,2*

1. 貴州大學農(nóng)學院, 貴州 貴陽 550025 2. 貴州大學國家小麥改良中心貴州分中心, 貴州 貴陽 550025 3. 貴州省農(nóng)業(yè)科學院水稻研究所, 貴州 貴陽 550006

植物脫落酸不敏感蛋白5（Abscisic acid-insensitive 5, ABI5）為堿性亮氨酸拉鏈類型（Basic leucine zipper, bZIP）的轉(zhuǎn)錄因子，在ABA信號途徑中起著重要作用。目前已經(jīng)報道的ABI5功能研究主要是以模式植物擬南芥（）為研究對象，而在小麥（）、水稻（）等農(nóng)作物中的研究報道極少。本文概述了植物中ABI5在種子發(fā)育、植物生長（特別是蔗糖信號途徑和蛋白質(zhì)泛素化過程）、花青素積累等生物學過程以及植物對干旱、低氮等逆境脅迫響應方面的最新研究進展，闡明在農(nóng)作物中開展ABI5分子網(wǎng)絡(luò)調(diào)控解析工作的重要性，以期為選育具有種子萌發(fā)可控（如抗穗發(fā)芽）、抗逆性強（如低氮）以及高產(chǎn)量等優(yōu)良性狀的作物品種提供理論基礎(chǔ)，對作物分子育種具有指導意義。

ABI5轉(zhuǎn)錄因子; 研究進展

植物激素脫落酸（Abscisic acid, ABA）廣泛參與植物生長發(fā)育的調(diào)控以及植物對逆境脅迫的響應過程。Finkelstein借助遺傳學和分子生物學手段鑒定得到了一系列與ABA響應有關(guān)的組分如ABI1（Abscisic acid-insensitive 1）、ABI2、ABI3、ABI4、ABI5。其中，ABI5為堿性亮氨酸拉鏈（Basic leucine zipper, bZIP）類型的轉(zhuǎn)錄因子[1]。ABI5含有C1-C4 4個保守的結(jié)構(gòu)域和1個bZIP結(jié)構(gòu)域[2]（圖1），其中bZIP結(jié)構(gòu)域中含有分別與泛素化和蘇素化有關(guān)的賴氨酸殘基（分別為K344和K391），而C1-C4結(jié)構(gòu)域中含有與磷酸化有關(guān)的絲氨酸殘基（S42、S145和S439）和蘇氨酸殘基（T201）[3]。

圖 1 ABI5保守結(jié)構(gòu)域示意圖

近年來研究發(fā)現(xiàn)，ABI5在種子休眠和萌發(fā)[4]、植物的生長發(fā)育[5]、植物對逆境脅迫的響應[6]、花青素的積累[7]以及26S蛋白酶體降解途徑[8]等生物學過程起著重要的作用。本文對近年來ABI5的研究進展進行了梳理和歸納，以期為培育具有優(yōu)良性狀的農(nóng)作物提供一定的思路。

1 ABI5與種子發(fā)育

ABA參與種子發(fā)育過程的調(diào)控，而在該生物學過程中ABI5起著重要的作用[1]。

1.1 通過對ABI5的轉(zhuǎn)錄調(diào)控

擬南芥中的研究發(fā)現(xiàn)，轉(zhuǎn)錄中介體（Mediator 25, MED25）和MADS-box轉(zhuǎn)錄因子AGL21（MADS-box transcription factor AGL21）能夠結(jié)合到ABI5的啟動子區(qū)域，分別負調(diào)控/正調(diào)控ABI5的表達進而參與種子發(fā)育過程的調(diào)控[9,10]；ABI5也能夠直接結(jié)合到過氧化氫酶1（Catalase 1, CAT1）基因和SHB1（Short hypocotyl under blue 1, SHB1）基因的啟動子區(qū)域，分別通過激活CAT1進而影響活性氧簇的動態(tài)平衡[11]和負調(diào)控SHB1[12]的表達進而調(diào)控ABI5下游信號組分的表達參與種子發(fā)育過程的調(diào)控。ABI5還能夠直接靶向編碼晚期胚胎豐富蛋白（Late embryogenesis abundant, LEA）的Em1和Em6基因并負調(diào)控兩者的表達[13]；擬南芥核因子Y家族蛋白（Nuclear factor Y family protein, NF-YC9）能夠直接與ABI5結(jié)合，進而使其結(jié)合并激活靶基因EM6的表達，實現(xiàn)對ABI5轉(zhuǎn)錄活性的正調(diào)控作用，從而參與種子萌發(fā)對ABA的響應[14]。小麥中的研究發(fā)現(xiàn)，ABI5基因的表達量隨著小麥種子成熟度的增加而增加，表明其參與種子發(fā)育過程的調(diào)控[15]。

此外，擬南芥SAG（A protein containing the midasin homologue 1 domain）[16]、萌發(fā)延遲基因1（Delay of Germination 1, DOG1）[17]、DELLA蛋白RGL2[4]以及轉(zhuǎn)錄因子AtMyb7[18]和RAV1[19]等都通過調(diào)控ABI5，進而參與ABA對種子萌發(fā)的調(diào)控。

1.2 通過對ABI5的穩(wěn)定性調(diào)控

研究發(fā)現(xiàn)擬南芥BIN2（Brassinazole insensitive 2）[20]、SOS2類似的蛋白激酶5（SOS2-like protein kinase 5, PSK5）[21]和蛋白磷酸化2A相關(guān)蛋白（Protein phosphatase2A (PP2A)-associated protein, TAP46）[22]通過磷酸化作用穩(wěn)定ABI5，維持種子中較高含量的ABI5水平從而抑制種子萌發(fā)。相反地，擬南芥光敏色素相關(guān)的絲氨酸/蘇氨酸蛋白磷酸酶（Phytochrome associated serine/threonine protein phosphatase, FyPP）與SnRK2激酶拮抗作用調(diào)控ABI5的磷酸化，通過去磷酸化降低ABI5蛋白的穩(wěn)定性，降低種子中ABI5的含量進而促進種子的萌發(fā)[23]。

擬南芥小泛素相關(guān)修飾物（Small ubiquitin-related modifier, SUMO）E3連接酶SIZ1通過SUMO修飾保護ABI5免受蛋白酶的降解，進而負調(diào)控ABA對種子萌發(fā)的調(diào)控[24]。相反，研究發(fā)現(xiàn)擬南芥CRWN（Crowded nuclei）蛋白家族通過調(diào)控ABI5的降解參與ABA控制的種子萌發(fā)，CRWN突變體對ABA超敏感并積累更高水平的ABI5蛋白[25]。此外，NO對ABI5蛋白153位半胱氨酸的S-亞硝基化有利于ABI5通過KEG（Keep on going）E3連接酶的生物學降解，從而促進種子的萌發(fā)。而ABI5蛋白153位半胱氨酸的突變能夠引起NO失去對ABI5蛋白穩(wěn)定性的調(diào)控，進而表現(xiàn)出抑制種子萌發(fā)的表型[26]。

綜上可知，ABI5在種子發(fā)育過程中有著極其重要的作用，ABI5基因的表達水平及ABI5蛋白的穩(wěn)定性直接影響種子的萌發(fā)和/或休眠等生物學過程。因此，在小麥、大豆、高粱等作物開展ABI5基因的研究是必要的，進而為種業(yè)技術(shù)的發(fā)展提供一定的理論基礎(chǔ)。

2 ABI5與植物的生長

蔗糖具有類生長素信號分子的功能，在擬南芥生長發(fā)育過程中起著重要的調(diào)控作用[27]。研究發(fā)現(xiàn)，ABI5能夠通過抑制生長素運輸載體PIN1的積累引起根中生長素水平的降低，通過調(diào)節(jié)根中生長素的水平進而參與蔗糖介導的對根分生組織區(qū)的抑制過程[28]。過表達ABI5能夠造成開花轉(zhuǎn)變過程的延遲，染色質(zhì)免疫沉淀發(fā)現(xiàn)ABI5通過結(jié)合到花發(fā)育基因FLC（Flowering locus c）的啟動子元件參與植物的花發(fā)育過程，進一步研究發(fā)現(xiàn)蔗糖非發(fā)酵1型相關(guān)蛋白激酶2（Sucrose nonfermenting 1-related protein kinase2, SnRK2）介導的ABI5的磷酸化在該過程中起著重要的作用，SnRK2通過調(diào)控ABI5的磷酸化能夠促進FLC基因的表達，進而實現(xiàn)擬南芥中ABA對開花轉(zhuǎn)變過程的抑制作用[29]。

此外，ABI5還參與植物子葉變綠[30]以及黑暗誘導葉片衰老[31]的生物學過程，但具體的作用機理仍有待于進一步的研究。

3 ABI5與逆境脅迫的響應

ABA與植物對逆境脅迫的響應過程緊密相關(guān)。對擬南芥突變體hls1（HOOKLESS1）進行研究，發(fā)現(xiàn)突變體中ABA介導的抗真菌侵染的現(xiàn)象消失，進一步研究發(fā)現(xiàn)HSL1介導的ABI5表達水平降低與該過程具有相關(guān)性[6]。低氮脅迫處理發(fā)現(xiàn)，擬南芥ABI5的表達水平受到顯著誘導，上調(diào)表達超過100倍[32]。在棉花中共表達擬南芥ABI3/Viviparous1（命名為AtRAV2）和ABI5能夠通過調(diào)節(jié)活性氧簇清除以及滲透調(diào)節(jié)marker基因的表達提高棉花對干旱脅迫的抗性[33]。此外，水稻[34]、水曲柳[35]等中的研究也發(fā)現(xiàn)ABI5與植物對逆境脅迫的響應有關(guān)。此外，ABI5還參與逆境脅迫下三酰基甘油（triacylglycerol, TAG）的積累。研究發(fā)現(xiàn)逆境脅迫條件下，擬南芥abi5中TAG合成途徑的限速酶二酰基甘油轉(zhuǎn)移酶（diacylglycerol acyltransferase 1, DGAT1）基因的表達水平以及TAG的積累量均降低；相反地，ABI5過表達轉(zhuǎn)基因株系中DGAT1的表達水平以及TAG的積累量均升高[36]。ABI5參與植物對逆境脅迫響應的生物學過程，并在植物對逆境脅迫的響應過程中起著重要作用。

4 ABI5與花青素的積累

對擬南芥abi5-4進行研究，發(fā)現(xiàn)3%蔗糖處理使得突變體中花青素的積累量高于野生型[7]。對擬南芥絲氨酸/精氨酸豐富（serine/arginine-rich, SR）蛋白SR45的研究發(fā)現(xiàn)，3%蔗糖處理能夠顯著誘導sr45中花青素合成基因APL3（ADP-Glc pyrophosphorylase, large subunit）、CHS以及ABI5的表達[37]。此外，擬南芥中的研究發(fā)現(xiàn)外源蔗糖和葡萄糖處理能夠誘導調(diào)控花青素合成途徑的花色苷色素產(chǎn)生基因（Production of anthocyanin pigment, PAP）PAP1和PAP2的表達，而突變體abi5-1能夠減弱該處理對PAP1和PAP2表達的誘導作用[38]，進而調(diào)節(jié)花青素的積累?？芍?，ABI5能夠通過影響花青素合成相關(guān)基因的表達實現(xiàn)對花青素的生物學調(diào)控，但其作用機制仍需要進一步闡明。

5 ABI5與26S蛋白酶體降解途徑

蛋白的泛素化過程在植物生長過程中具有廣泛而重要的生物學功能，該過程一般由泛素激活酶（Ubiquitin-activating enzymes, E1）、泛素結(jié)合酶（Ubiquitin-conjugating enzymes, E2）和E3催化的級聯(lián)反應完成[39]。26S蛋白酶體能夠識別泛素化的蛋白，經(jīng)過去泛素化酶（Deubiquitinating enzymes, DUBs）去泛素化后被26S蛋白酶體中的蛋白酶降解[40]。擬南芥中研究發(fā)現(xiàn)定位于細胞質(zhì)和內(nèi)膜系統(tǒng)的泛素化E3連接酶KOG能夠通過泛素化作用抑制ABI5的積累水平[8]，進一步研究發(fā)現(xiàn)泛素化的ABI5通過26S蛋白酶體進行降解，從而實現(xiàn)KOG對ABI5蛋白表達水平的調(diào)控[3]。此外，Cullin4 E3復合體底物受體DWA1/DWA2（DWD hypersensitive to ABA1/2）和ABD1（ABA-hypersensitive DCAF1）也能夠通過26S蛋白酶體途徑調(diào)控細胞核中ABI5的水平[41]。總之，植物可以通過26S蛋白酶體降解途徑實現(xiàn)對細胞核和細胞質(zhì)中ABI5表達水平的調(diào)節(jié)，進而對ABA信號通路進行調(diào)控。

目前，對農(nóng)作物中ABI5的功能報道較少，然而由于該蛋白是ABA信號途徑中重要的轉(zhuǎn)錄因子，闡明農(nóng)作物中ABI5的調(diào)控網(wǎng)絡(luò)為最終應用于分子育種具有指導性的意義。

6 總結(jié)與展望

植物ABI5是ABA信號通路中重要的轉(zhuǎn)錄因子，參與種子發(fā)育、植物生長、花青素積累等生物學過程，并在植物對逆境脅迫如干旱、低氮等的響應方面起著重要的作用。目前擬南芥中ABI5的功能研究已有較多的報道，而對小麥、水稻等農(nóng)作物中的研究報道極少。綜上可知，ABI5在種子發(fā)育、植物生長（特別是蔗糖信號途徑和蛋白質(zhì)泛素化過程）、植物對逆境脅迫的響應以及花青素積累等生物學過程中起重要作用，在農(nóng)作物中可以重點開展該蛋白在上述方面的研究。然而，農(nóng)作物中ABI5具有的生物學功能是否與擬南芥中的功能相同？哪些蛋白（靶向或者非靶向）參與調(diào)控ABI5的生物學功能？ABI5又能夠通過調(diào)控哪些蛋白實現(xiàn)上游信號的傳遞？在某一特定生物學過程中ABI5分子調(diào)控網(wǎng)絡(luò)是如何協(xié)同的？等問題仍需要進行系統(tǒng)深入的研究和探討。對小麥、水稻、玉米等重要農(nóng)作物中ABI5進行深入的研究，有助于人們深入了解ABI5的分子調(diào)控網(wǎng)絡(luò)，為培育具有抗穗發(fā)芽、抗逆性強、延緩衰老等優(yōu)良性狀的品種提供一定的理論基礎(chǔ)。綜上所述，在農(nóng)作物中開展闡明ABI5分子調(diào)控網(wǎng)絡(luò)的相關(guān)研究是必要可行的。

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Research Advances of Plant ABI5 Transcription Factors

LI Lu-hua1,2, WANG Zhong-ni3, WANG Wen-xin1, WANG Huai-yu1, REN Ming-jian1,2, XU Ru-hong1,2*

1.550025,2.550025,3.550006,

Plant abscisic acid insensitive 5 (ABI5) protein is transcription factors of basic leucine zipper (bZIP) type and plays an important role in ABA signaling pathway. Currently, the functional research of ABI5 in plant mainly focused on, while few studies have been reported on crops such as wheat () and rice (). In this paper, we summarized the advanced researches of ABI5 in biological process such as seed development, plant growth (especially the sucrose signaling pathway and protein ubiquitination process), anthocyanin accumulation and in stress responses such as drought and low nitrogen, and elucidated the importance of carrying out the molecular regulation research of ABI5 in crops, which would provide theoretical basis for crop varieties with excellent traits, such as controllable seed germination (e.g. preharvest sprouting resistance), strong stress resistance (e.g. low nitrogen) and high yield, and has instructive significance of molecular breeding in crops.

ABI5 transcription factors; research advance

Q7

A

1000-2324(2019)04-0537-05

2018-08-22

2018-10-10

貴州省科技計劃項目(黔科合基礎(chǔ)[2019]1073號);國家自然科學基金項目(31660390); 貴州省農(nóng)業(yè)成果轉(zhuǎn)化計劃項目(黔科合成果(2016)4022號);貴州大學引進人才科研項目(貴大人基合字2017(49號))

李魯華(1985-),男,博士,講師,主要從事作物遺傳育種工作. E-mail:luhua_li@163.com

Author for correspondence. E-mail:xrhgz@163.com