詹妮，謝耀堅(jiān)，吳志華，劉果，尚秀華

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ceRNA對(duì)植物纖維素形成的調(diào)控研究進(jìn)展

詹妮，謝耀堅(jiān)，吳志華＊，劉果，尚秀華

(國(guó)家林業(yè)和草原局桉樹(shù)研究開(kāi)發(fā)中心，廣東 湛江 524022)

植物纖維素的形成是由多個(gè)基因參與且呈網(wǎng)絡(luò)調(diào)控。通過(guò)對(duì)纖維素形成過(guò)程中的關(guān)鍵酶基因、轉(zhuǎn)錄因子以及ceRNA研究的闡述，深入了解纖維素生物合成調(diào)控機(jī)制。綜述植物纖維素形成過(guò)程中的纖維素合酶、蔗糖合成酶、MYB等重要基因以及l(fā)ncRNA、miRNA、circRNA類(lèi)ceRNA，闡述其復(fù)雜的分子調(diào)控網(wǎng)絡(luò)，以期解析植物纖維素形成過(guò)程中的分子調(diào)控機(jī)制，深入了解植物纖維素形成過(guò)程。

競(jìng)爭(zhēng)性內(nèi)源RNA；纖維素；轉(zhuǎn)錄因子；表達(dá)調(diào)控

纖維素作為植物細(xì)胞壁中必不可少的結(jié)構(gòu)成分，發(fā)揮著重要的作用。植物細(xì)胞初生壁中的纖維素微纖絲在植物細(xì)胞的擴(kuò)增階段調(diào)控植物形態(tài)建成，次生壁中的纖維素使植物細(xì)胞具有特定功能，纖維素對(duì)植物生長(zhǎng)的重要作用使得對(duì)其研究具有重要意義[1]。纖維素形成是個(gè)較復(fù)雜的過(guò)程，該過(guò)程涉及一系列重要生物學(xué)過(guò)程，其中每個(gè)過(guò)程均由多基因參與且呈網(wǎng)絡(luò)調(diào)控，研究植物纖維素形成過(guò)程中的關(guān)鍵基因及其生物合成調(diào)控機(jī)制，已成為當(dāng)前研究的熱點(diǎn)[2]。

ceRNA(Competing endogenous RNA，競(jìng)爭(zhēng)性內(nèi)源RNA)是指生物體內(nèi)復(fù)雜的轉(zhuǎn)錄調(diào)控網(wǎng)絡(luò)中的RNA，包括長(zhǎng)鏈非編碼RNA(Long noncoding RNA，lncRNA)、微小RNA(MicroRNA，miRNA)以及環(huán)狀RNA(CircleRNA，circRNA)等[3-4]。ceRNA通過(guò)miRNA應(yīng)答元件(MicroRNA response element，MRE)與靶mRNA競(jìng)爭(zhēng)性的結(jié)合同種的miRNA分子，使miRNA的表達(dá)水平及活性相對(duì)下降，從而抑制了miRNA對(duì)靶mRNA 的沉默效應(yīng)，發(fā)揮調(diào)控的作用[5-7]。RNA轉(zhuǎn)錄物通過(guò)它們所共有的 MREs位點(diǎn)來(lái)彼此調(diào)節(jié)，共有的MREs數(shù)量越多，它們的交流或共調(diào)節(jié)的程度也越大，因此ceRNA 能形成一種大規(guī)模轉(zhuǎn)錄調(diào)控網(wǎng)絡(luò)，實(shí)現(xiàn)lncRNAs，miRNAs及circRNAs等通過(guò)MREs進(jìn)行相互作用，通過(guò)競(jìng)爭(zhēng)MREs 構(gòu)成一個(gè)完整復(fù)雜的ceRNA分子調(diào)控網(wǎng)絡(luò)[6]。

1 植物纖維素形成過(guò)程中的關(guān)鍵基因

纖維素合酶(Cellulose synthase, CesA)組成纖維素合酶復(fù)合體(Cellulose synthase complex, CSC)，催化β-1,4糖苷鍵的形成，合成纖維素，在植物纖維素合成途徑中發(fā)揮主要調(diào)節(jié)作用[8]。在大青楊()、苧麻()、馬尾松()等木本植物都相繼克隆出CesA基因[9-11]。在擬南芥()中CesA1、CesA3、CesA6 負(fù)責(zé)初生壁的形成[12-14]，CesA4、CesA7、CesA8負(fù)責(zé)次生壁的合成[15-16]。

蔗糖合成酶(Sucrose synthase, SuSy)影響植物細(xì)胞分化以及細(xì)胞壁的形成，能夠提供細(xì)胞壁合成的底物，SuSy的活性與纖維素合成有關(guān)[17-23]。SuSy基因廣泛存在于植物中，CARDINI等[24]首次從小麥()中克隆了SuSy基因，此后在擬南芥、棉花(.)、馬鈴薯()、胡蘿卜()、玉米()、柑橘()、水稻()、棗()、甘蔗()等植物中獲得SuSy基因[25-31]。SuSy基因?qū)γ藁?、煙?)以及楊樹(shù)的纖維素含量、纖維長(zhǎng)度以及纖維強(qiáng)度等至關(guān)重要[32-34]。

擴(kuò)展蛋白(Expansin，EXP)是植物細(xì)胞壁重要的組成部分，調(diào)節(jié)細(xì)胞伸展性，通過(guò)打斷細(xì)胞壁纖維素和半纖維素之間的非共價(jià)鍵，從而改變細(xì)胞壁承重網(wǎng)絡(luò)，使其產(chǎn)生位移，導(dǎo)致細(xì)胞壁伸展，加速細(xì)胞生長(zhǎng)，調(diào)節(jié)組織生長(zhǎng)[35-36]。1989年COSGROVE[37]首次從黃瓜()根尖細(xì)胞壁中提取分離出EXP。EXP能夠塑造初生細(xì)胞壁中纖維素-半纖維素網(wǎng)絡(luò)，EXP的活動(dòng)能夠影響細(xì)胞壁的結(jié)構(gòu)和組分，進(jìn)而影響纖維和導(dǎo)管的形態(tài)[38]。GRAY等[39]在楊樹(shù)中克隆了α-EXP基因和β-EXP基因，發(fā)現(xiàn)PttEXP1基因在成熟莖段的次生生長(zhǎng)較為活躍。XU等[40]發(fā)現(xiàn)在棉花纖維細(xì)胞的伸長(zhǎng)過(guò)程中，α-EXP蛋白發(fā)揮了重要的調(diào)控作用。SARA等[41]從牽牛花()中獲得了一條PhEXP1基因，反義轉(zhuǎn)化后，發(fā)現(xiàn)牽?；ū砥ぜ?xì)胞面積也相應(yīng)的減少，細(xì)胞壁發(fā)生了改變，導(dǎo)致細(xì)胞壁機(jī)械強(qiáng)度下降。

AGO蛋白(Argonaute protein)主要包含PAZ和PIWI結(jié)構(gòu)域，是小RNA介導(dǎo)的RNA沉默通路中RNA誘導(dǎo)的沉默復(fù)合物(RNA-induced silencing complex，RISC)的核心成分。AGO蛋白通過(guò)與miRNAs(microRNAs)、siRNAs(small interfering RNAs)、piRNAs(Piwi-interacting RNAs)等不同類(lèi)型的小非編碼RNA(small non-coding RNA)結(jié)合，AGO蛋白能夠特異地停留在與小RNA互補(bǔ)的靶基因mRNA上，其自身的內(nèi)切酶可以對(duì)目標(biāo)靶基因進(jìn)行切割，從而引起靶基因沉默，在調(diào)控植物生長(zhǎng)發(fā)育中起到重要的作用[42-43]。

第3類(lèi)亮氨酸拉鏈蛋白(ClassⅢ homeodomain leucine zipper, HD-ZipⅢ)轉(zhuǎn)錄因子、MYB(V-myb avian myeloblastosis viral oncogene homo)轉(zhuǎn)錄因子以及NAC(No apical meristem/Arabidopsis thaliana transcription activator factor/CUP- -shaped cotyledon)轉(zhuǎn)錄因子等在植物細(xì)胞生長(zhǎng)發(fā)育過(guò)程中具重要調(diào)控作用，參與植物的生長(zhǎng)代謝調(diào)控[44]。

研究表明，在擬南芥和水稻HD-ZipⅢI家族各有5個(gè)成員[45]，毛果楊()基因組中則含有8個(gè)HD-ZipⅢ基因家族成員[46]。在白云杉()和火炬松()中已分別被克隆到4和5個(gè)HD-Zip III基因[47]。MYB類(lèi)轉(zhuǎn)錄因子參與植物苯丙烷類(lèi)代謝途徑的調(diào)節(jié)，調(diào)控次生細(xì)胞壁的形成[48-49]。目前，MYB轉(zhuǎn)錄因子已在擬南芥、金魚(yú)草()、大豆()、煙草、蘋(píng)果()、白樺(、毛白楊()等物種中分離并鑒定[50]。HAI 等[51]對(duì)玉米和擬南芥MYB轉(zhuǎn)錄因子分析發(fā)現(xiàn)，有4個(gè)亞組的MYB轉(zhuǎn)錄因子參與調(diào)控次生壁的增厚。劉慧子等[52]研究表明，白樺MYB家族中17條MYB家族基因中的絕大部分參與調(diào)控形成層的發(fā)育。葉勝龍[53]研究發(fā)現(xiàn)，毛白楊MYB055轉(zhuǎn)錄因子參與調(diào)控次生壁合成，影響苯丙氨酸代謝途徑，從而調(diào)控纖維素合成等相關(guān)基因的表達(dá)。

2 植物纖維素形成過(guò)程中的ceRNA

測(cè)序技術(shù)日益發(fā)展使基因數(shù)據(jù)庫(kù)與轉(zhuǎn)錄組數(shù)據(jù)庫(kù)日益充實(shí)，為ceRNA的挖掘和功能研究提供了有利的數(shù)據(jù)支持。ceRNA在生物發(fā)育和基因表達(dá)中發(fā)揮著復(fù)雜的精確調(diào)控功能，對(duì)其深入研究有助于揭示基因表達(dá)調(diào)控網(wǎng)絡(luò)對(duì)于生命體的復(fù)雜性[54]。

lncRNA指長(zhǎng)度大于200個(gè)核苷酸，但含有1個(gè)少于100個(gè)氨基酸開(kāi)放閱讀框(Open reading frame，ORF)的RNA，可分為長(zhǎng)鏈非編碼自然反義轉(zhuǎn)錄本(Long noncoding natural antisense transcripts，lincNATs)、內(nèi)含子 lncRNAs(Intronic lncRNAs)、啟動(dòng)子 lncRNAs(Promoter lncRNAs)和長(zhǎng)鏈基因間 ncRNAs(Long intergenic ncRNAs，lincRNAs)。lncRNAs 可作為與其互作分子的招募者、系結(jié)者、引導(dǎo)者、誘捕者和信號(hào)分子，從而發(fā)揮調(diào)控作用[55-57]。lncRNAs通過(guò)與miRNA結(jié)合，從而隔離miRNA，調(diào)控miRNA的表達(dá)水平，降低miRNA對(duì)mRNA的調(diào)控，最終促進(jìn)了mRNA的表達(dá)。在植物中鑒定出大量lncRNAs，如擬南芥[58]、小麥[59]、玉米[60]、谷子()[61]、棉花[62]、江南卷柏()[63]、沙棘()[64]、芒草()[65]以及毛果楊、毛白楊[66-67]。

miRNA是一類(lèi)內(nèi)生的且長(zhǎng)度約為20 ~ 24個(gè)核苷酸的小RNA，在轉(zhuǎn)錄以及轉(zhuǎn)錄后的過(guò)程中調(diào)控基因表達(dá)[68]。miRNA在細(xì)胞內(nèi)具有多種重要的調(diào)節(jié)作用，參與了植物器官發(fā)育、代謝調(diào)節(jié)，與細(xì)胞的增殖、分化、凋亡等一系列生理過(guò)程密切相關(guān)[69]。miRNA是通過(guò)切割目標(biāo)靶基因mRNA或抑制其翻譯來(lái)實(shí)現(xiàn)對(duì)目標(biāo)靶基因的調(diào)控，這種調(diào)控既能夠通過(guò)一個(gè)miRNA調(diào)控多個(gè)基因的表達(dá)，亦可通過(guò)幾個(gè)miRNA共同調(diào)控某個(gè)基因的表達(dá)，從而形成復(fù)雜的調(diào)控網(wǎng)絡(luò)[70]。MCNAIR[71]研究發(fā)現(xiàn)12個(gè)miRNA在正常生長(zhǎng)和快速生長(zhǎng)桉樹(shù)()中的表達(dá)模式，miRNA在正常桉樹(shù)和應(yīng)拉木的發(fā)育過(guò)程中起重要作用。利用高通量測(cè)序技術(shù)挖掘了包括2個(gè)藍(lán)桉()基因型的木質(zhì)部，獲得了大量的miRNA信息[72]。李崇奇等[73]研究表明，有41個(gè)miRNA與巨桉()木質(zhì)形成相關(guān)，主要調(diào)控ARF、HD-ZIPIII、KAN、MYB 和NAC轉(zhuǎn)錄因子。circRNA是一類(lèi)線性閉合環(huán)狀內(nèi)源性的非編碼RNA分子，circRNA通過(guò)吸附miRNA并參與其表達(dá)調(diào)控過(guò)程，circRNA 能夠特異性結(jié)合miRNA，使其失去調(diào)控mRNA 的功能，調(diào)控基因表達(dá)等生物過(guò)程[74-77]。circRNA分為外顯子circRNA、基因間circRNA和內(nèi)含子circRNA[78-79]。circRNA廣泛表達(dá)于不同的植物中，表達(dá)具有時(shí)空組織特異性，circRNA作為內(nèi)源性非編碼RNA在真核生物的生長(zhǎng)發(fā)育過(guò)程中發(fā)揮著重要作用，引起人們廣泛的關(guān)注[80]。2014年在擬南芥根部發(fā)現(xiàn)circRNA后[81]，2015年ANDREEVA和COOPER研究報(bào)道了circRNA廣泛存在于動(dòng)植物細(xì)胞組織中，且具有很多特殊的生物學(xué)特性之后，引起國(guó)內(nèi)外科學(xué)家的高度重視[82]。在水稻[83-84]、大麥()[85]、番茄()[86]以及小麥[87]中發(fā)現(xiàn)存在大量的circRNAs。YE等[84]在水稻的根和擬南芥的葉中分別鑒定了12 037和6 012個(gè)circRNAs。

表1 與木質(zhì)相關(guān)的miRNA[73]

通過(guò)對(duì)參與植物纖維素形成過(guò)程中的關(guān)鍵基因、ceRNA的進(jìn)一步研究，能夠更好的解析植物纖維素形成過(guò)程中的分子調(diào)控機(jī)制，以期獲得對(duì)植物纖維素形成過(guò)程的深入了解，從而為育種工作服務(wù)?，F(xiàn)今在植物中已經(jīng)鑒定出許多ceRNA，但只有少數(shù)做了功能驗(yàn)證，今后植物ceRNA的研究方向可能會(huì)趨向于搜索基因的功能，剖析功能冗余以及其應(yīng)用等方面。

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Research Progress in the Regulation of ceRNA on Plant Cellulose Formation

ZHAN Ni, XIE Yaojian, WU Zhihua, LIU Guo, SHANG Xiuhua

(,)

The formation of plant cellulose is regulated by multiple genes and pathways. In this paper, the key enzyme genes, transcription factors and ceRNA in the process of cellulose formation are elaborated to further understand the regulation mechanism of cellulose biosynthesis. The key genes including cellulose synthase, sucrose synthase, MYB and ceRNA including lncRNA, miRNA and circRNA in the process of plant cellulose formation are reviewed. The complex molecular control network is expounded in order to analyze the molecular control mechanism of plant cellulose formation, and to understand the process of plant cellulose formation.

ceRNA; cellulose; transcription factors; expression regulation

Q74

A

國(guó)家自然科學(xué)基金面上項(xiàng)目“桉樹(shù)抗風(fēng)特性及其主要影響因子研究”(31570615);國(guó)家重點(diǎn)研發(fā)計(jì)劃課題“桉樹(shù)、云南松(思茅松）、華山松豐產(chǎn)增效技術(shù)集成與示范”(2017YFD0601202）

詹妮(1990― )，女，博士研究生，主要從事桉樹(shù)林木遺傳育種方面的研究，E-mail: jennyzn1122@163.com

吳志華(1974― )，男，副研究員，主要從事林木逆境生理研究，E-mail: wzhua2889@163.com