ceRNA對植物纖維素形成的調控研究進展

詹妮，謝耀堅，吳志華，劉果，尚秀華

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ceRNA對植物纖維素形成的調控研究進展

詹妮，謝耀堅，吳志華＊，劉果，尚秀華

(國家林業和草原局桉樹研究開發中心，廣東 湛江 524022)

植物纖維素的形成是由多個基因參與且呈網絡調控。通過對纖維素形成過程中的關鍵酶基因、轉錄因子以及ceRNA研究的闡述，深入了解纖維素生物合成調控機制。綜述植物纖維素形成過程中的纖維素合酶、蔗糖合成酶、MYB等重要基因以及lncRNA、miRNA、circRNA類ceRNA，闡述其復雜的分子調控網絡，以期解析植物纖維素形成過程中的分子調控機制，深入了解植物纖維素形成過程。

競爭性內源RNA；纖維素；轉錄因子；表達調控

纖維素作為植物細胞壁中必不可少的結構成分，發揮著重要的作用。植物細胞初生壁中的纖維素微纖絲在植物細胞的擴增階段調控植物形態建成，次生壁中的纖維素使植物細胞具有特定功能，纖維素對植物生長的重要作用使得對其研究具有重要意義[1]。纖維素形成是個較復雜的過程，該過程涉及一系列重要生物學過程，其中每個過程均由多基因參與且呈網絡調控，研究植物纖維素形成過程中的關鍵基因及其生物合成調控機制，已成為當前研究的熱點[2]。

ceRNA(Competing endogenous RNA，競爭性內源RNA)是指生物體內復雜的轉錄調控網絡中的RNA，包括長鏈非編碼RNA(Long noncoding RNA，lncRNA)、微小RNA(MicroRNA，miRNA)以及環狀RNA(CircleRNA，circRNA)等[3-4]。ceRNA通過miRNA應答元件(MicroRNA response element，MRE)與靶mRNA競爭性的結合同種的miRNA分子，使miRNA的表達水平及活性相對下降，從而抑制了miRNA對靶mRNA 的沉默效應，發揮調控的作用[5-7]。RNA轉錄物通過它們所共有的 MREs位點來彼此調節，共有的MREs數量越多，它們的交流或共調節的程度也越大，因此ceRNA 能形成一種大規模轉錄調控網絡，實現lncRNAs，miRNAs及circRNAs等通過MREs進行相互作用，通過競爭MREs 構成一個完整復雜的ceRNA分子調控網絡[6]。

1 植物纖維素形成過程中的關鍵基因

纖維素合酶(Cellulose synthase, CesA)組成纖維素合酶復合體(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揮主要調節作用[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 負責初生壁的形成[12-14]，CesA4、CesA7、CesA8負責次生壁的合成[15-16]。

蔗糖合成酶(Sucrose synthase, SuSy)影響植物細胞分化以及細胞壁的形成，能夠提供細胞壁合成的底物，SuSy的活性與纖維素合成有關[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基因對棉花、煙草()以及楊樹的纖維素含量、纖維長度以及纖維強度等至關重要[32-34]。

擴展蛋白(Expansin，EXP)是植物細胞壁重要的組成部分，調節細胞伸展性，通過打斷細胞壁纖維素和半纖維素之間的非共價鍵，從而改變細胞壁承重網絡，使其產生位移，導致細胞壁伸展，加速細胞生長，調節組織生長[35-36]。1989年COSGROVE[37]首次從黃瓜()根尖細胞壁中提取分離出EXP。EXP能夠塑造初生細胞壁中纖維素-半纖維素網絡，EXP的活動能夠影響細胞壁的結構和組分，進而影響纖維和導管的形態[38]。GRAY等[39]在楊樹中克隆了α-EXP基因和β-EXP基因，發現PttEXP1基因在成熟莖段的次生生長較為活躍。XU等[40]發現在棉花纖維細胞的伸長過程中，α-EXP蛋白發揮了重要的調控作用。SARA等[41]從牽牛花()中獲得了一條PhEXP1基因，反義轉化后，發現牽牛花表皮細胞面積也相應的減少，細胞壁發生了改變，導致細胞壁機械強度下降。

AGO蛋白(Argonaute protein)主要包含PAZ和PIWI結構域，是小RNA介導的RNA沉默通路中RNA誘導的沉默復合物(RNA-induced silencing complex，RISC)的核心成分。AGO蛋白通過與miRNAs(microRNAs)、siRNAs(small interfering RNAs)、piRNAs(Piwi-interacting RNAs)等不同類型的小非編碼RNA(small non-coding RNA)結合，AGO蛋白能夠特異地停留在與小RNA互補的靶基因mRNA上，其自身的內切酶可以對目標靶基因進行切割，從而引起靶基因沉默，在調控植物生長發育中起到重要的作用[42-43]。

第3類亮氨酸拉鏈蛋白(ClassⅢ homeodomain leucine zipper, HD-ZipⅢ)轉錄因子、MYB(V-myb avian myeloblastosis viral oncogene homo)轉錄因子以及NAC(No apical meristem/Arabidopsis thaliana transcription activator factor/CUP- -shaped cotyledon)轉錄因子等在植物細胞生長發育過程中具重要調控作用，參與植物的生長代謝調控[44]。

研究表明，在擬南芥和水稻HD-ZipⅢI家族各有5個成員[45]，毛果楊()基因組中則含有8個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HD-Zip III基因[47]。MYB類轉錄因子參與植物苯丙烷類代謝途徑的調節，調控次生細胞壁的形成[48-49]。目前，MYB轉錄因子已在擬南芥、金魚草()、大豆()、煙草、蘋果()、白樺(、毛白楊()等物種中分離并鑒定[50]。HAI 等[51]對玉米和擬南芥MYB轉錄因子分析發現，有4個亞組的MYB轉錄因子參與調控次生壁的增厚。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控形成層的發育。葉勝龍[53]研究發現，毛白楊MYB055轉錄因子參與調控次生壁合成，影響苯丙氨酸代謝途徑，從而調控纖維素合成等相關基因的表達。

2 植物纖維素形成過程中的ceRNA

測序技術日益發展使基因數據庫與轉錄組數據庫日益充實，為ceRNA的挖掘和功能研究提供了有利的數據支持。ceRNA在生物發育和基因表達中發揮著復雜的精確調控功能，對其深入研究有助于揭示基因表達調控網絡對于生命體的復雜性[54]。

lncRNA指長度大于200個核苷酸，但含有1個少于100個氨基酸開放閱讀框(Open reading frame，ORF)的RNA，可分為長鏈非編碼自然反義轉錄本(Long noncoding natural antisense transcripts，lincNATs)、內含子 lncRNAs(Intronic lncRNAs)、啟動子 lncRNAs(Promoter lncRNAs)和長鏈基因間 ncRNAs(Long intergenic ncRNAs，lincRNAs)。lncRNAs 可作為與其互作分子的招募者、系結者、引導者、誘捕者和信號分子，從而發揮調控作用[55-57]。lncRNAs通過與miRNA結合，從而隔離miRNA，調控miRNA的表達水平，降低miRNA對mRNA的調控，最終促進了mRNA的表達。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是一類內生的且長度約為20 ~ 24個核苷酸的小RNA，在轉錄以及轉錄后的過程中調控基因表達[68]。miRNA在細胞內具有多種重要的調節作用，參與了植物器官發育、代謝調節，與細胞的增殖、分化、凋亡等一系列生理過程密切相關[69]。miRNA是通過切割目標靶基因mRNA或抑制其翻譯來實現對目標靶基因的調控，這種調控既能夠通過一個miRNA調控多個基因的表達，亦可通過幾個miRNA共同調控某個基因的表達，從而形成復雜的調控網絡[70]。MCNAIR[71]研究發現12個miRNA在正常生長和快速生長桉樹()中的表達模式，miRNA在正常桉樹和應拉木的發育過程中起重要作用。利用高通量測序技術挖掘了包括2個藍桉()基因型的木質部，獲得了大量的miRNA信息[72]。李崇奇等[73]研究表明，有41個miRNA與巨桉()木質形成相關，主要調控ARF、HD-ZIPIII、KAN、MYB 和NAC轉錄因子。circRNA是一類線性閉合環狀內源性的非編碼RNA分子，circRNA通過吸附miRNA并參與其表達調控過程，circRNA 能夠特異性結合miRNA，使其失去調控mRNA 的功能，調控基因表達等生物過程[74-77]。circRNA分為外顯子circRNA、基因間circRNA和內含子circRNA[78-79]。circRNA廣泛表達于不同的植物中，表達具有時空組織特異性，circRNA作為內源性非編碼RNA在真核生物的生長發育過程中發揮著重要作用，引起人們廣泛的關注[80]。2014年在擬南芥根部發現circRNA后[81]，2015年ANDREEVA和COOPER研究報道了circRNA廣泛存在于動植物細胞組織中，且具有很多特殊的生物學特性之后，引起國內外科學家的高度重視[82]。在水稻[83-84]、大麥()[85]、番茄()[86]以及小麥[87]中發現存在大量的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circRNAs。

表1 與木質相關的miRNA[73]

通過對參與植物纖維素形成過程中的關鍵基因、ceRNA的進一步研究，能夠更好的解析植物纖維素形成過程中的分子調控機制，以期獲得對植物纖維素形成過程的深入了解，從而為育種工作服務。現今在植物中已經鑒定出許多ceRNA，但只有少數做了功能驗證，今后植物ceRNA的研究方向可能會趨向于搜索基因的功能，剖析功能冗余以及其應用等方面。

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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

國家自然科學基金面上項目“桉樹抗風特性及其主要影響因子研究”(31570615);國家重點研發計劃課題“桉樹、云南松(思茅松）、華山松豐產增效技術集成與示范”(2017YFD0601202）

詹妮(1990― )，女，博士研究生，主要從事桉樹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