摘要
开花期是牧草重要的农艺性状,对牧草产量、品质和利用价值具有重要影响。开花时间由微效多基因控制,调控网络复杂。开展牧草开花基因的研究,阐明牧草开花调控分子机理,有利于加快牧草开花基因挖掘,培育满足生产需求的不同成熟度的牧草新品种。牧草开花基因在模式植物拟南芥(Arabidopsis thaliana)、水稻(Oryza sativa)、小麦(Triticum aestivum)等农作物已开展了系统深入的研究,而牧草研究相对较少。本文综述了牧草开花基因研究最新进展,以期为牧草开花基因研究提供理论参考。
Flowering time is an important agronomic trait, and has an important influence on the yield, quality, and utilization of forage grasses. Flowering time is controlled by micro-effect polygenes and the regulatory network is complex.Research on flowering genes and the molecular mechanisms of flowering regulation will facilitate the exploration and utilization of flowering genes, and lead to the development of new forage varieties with different maturities to meet different production requirements. Although flowering genes have been extensively and systematically studied in model plants such as Arabidopsis thaliana, rice (Oryza sativa), wheat (Triticum aestivum), and other crops, there has been comparatively little study of these genes in forage grasses. In this paper, we review the latest advances in the study of flowering genes in order to provide a theoretical reference for the study of these genes in forage grasses.
引文
[1]WOODS D P,MCKEOWN M A,DONG Y,PRESTON J C,AMASINO R M.Evolution of VRN2/GhD7-like genes in vernalization-mediated repression of grass flowering.Plant Physiology,2016,170(4):2124-2135.
[2]DENG W,CASAO M C,WANG P,SATO K,HAYES P M,FINNEGAN E J,TREVASKIS B.Direct links between the vernalization response and other key traits of cereal crops.Nature Communication,2015,6(6):5882.
[3]VOGEL K P,BREJDA J J,WALTERS D T,BUXTON D R.Switchgrass biomass production in the midwest USA.Agronomy Journal,2002,94(3):413-420.
[4]MOURADOV A,CREMER F,COUPLAND G.Control of flowering time:Interacting pathways as a basis for diversity.Plant Cell,2002,14(Suppl):S111-S130.
[5]BOSS P K,BASTOW R M,MYLNE J S,DEAN C.Multiple pathways in the decision to flower:Enabling,promoting,and resetting.Plant Cell,2004,16(Suppl):S18-S31.
[6]PENG F Y,HU Z,YANG R C.Bioinformatic prediction of transcription factor binding sites at promoter regions of genes for photoperiod and vernalization responses in model and temperate cereal plants.BMC Genomics,2016,17(1):573.
[7]DORCA-FORNELL C,GREGIS V,GRANDI V,COUPLAND G,COLOMBO L,KATER M M.The Arabidopsis SOC1-like genes AGL42,AGL71 and AGL72 promote flowering in the shoot apical and axillary meristems.Plant Journal for Cell&Molecular Biology,2011,67(6):1006-1017.
[8]COCKRAM J,JONES H,LEIGH F J,O'SULLIVAN D,POWELL W,LAURIE D A,GREENLAND A J.Control of flowering time in temperate cereals:Genes,domestication,and sustainable productivity.Journal of Experimental Botany,2007,58(6):1231-1244.
[9]HIGGINS J A,BAILEY P C,LAURIE D A.Comparative genomics of flowering time pathways using Brachypodium distachyon as a model for the temperate grasses.PLoS One,2010,5(4):e10065.
[10]CHIRUTA C,FILIPOV F,CALIN M.Estimating the duration of daylight in a given time of the year depending on the latitude of the location.Research Journal of Agricultural Science,2010,42(3):71-76.
[11]WADA M,SHIMAZAKI K,IINO M.Light Sensing in Plants.2005.Berlin:Springer,2005:333-337.
[12]FERNáNDEZ V,TAKAHASHI Y,LEGOURRIEREC J,COUPLAND G.Photoperiodic and thermosensory pathways interact through CONSTANS to promote flowering at high temperature under short days.The Plant Journal,2016,86(5):426-440.
[13]ONOUCHI H,IGE?O M I,PéRILLEUX C,GRAVES K,COUPLAND G.Mutagenesis of plants overexpressing CONSTANSdemonstrates novel interactions among Arabidopsis flowering-time genes.Plant Cell,2000,12(6):885-900.
[14]IZAWA T,MIHARA M,SUZUKI Y,GUPTA M,ITOH H,ATSUSHI J,NAGANO,MOTOYAMA R,SAWADA Y,YANO M,MASAMI YOKOTA HIRAI,MAKINO A,NAGAMURA Y.Os-GIGANTEA confers robust diurnal rhythms on the global transcriptome of rice in the field.Plant Cell,2011,23(5):1741-1755.
[15]BENDIX C,MENDOZA J M,STANLEY D N,MEELEY R,HARMON F G.The circadian clock-associated gene gigantea1,affects maize developmental transitions.Plant Cell&Environment,2013,36(7):1379-1390.
[16]MARTIN J,STORGAARD M,ANDERSEN C H,NIELSEN K K.Photoperiodic regulation of flowering in perennial ryegrass involving a CONSTANS-like homolog.Plant Molecular Biology,2004,56(2):159-169.
[17]GAGIC M,FAVILLE M,KARDAILSKY I,PUTTERILL J.Comparative genomics and functional characterisation of the GIGANTEA,gene from the temperate forage perennial ryegrass Lolium perenne.Plant Molecular Biology Reporter,2015,33(4):1-9.
[18]HECHT V,LAURIE R E,WELLER J.Isolation and functional analysis of CONSTANS-LIKE genes suggests that a central role for CONSTANS in flowering time control is not evolutionarily conserved in Medicago truncatula.Frontiers in Plant Science,2014,5(17):486.
[19]王鹏,张春庆,陈化榜,吴承来.小麦冬性强弱评价体系的建立.生态学报,2012,32(4):1230-1240.WANG P,ZHANG C Q,CHEN H B,WU C L.The evaluation system of strength of winterness in wheat.Acta Ecologica Sinica,2012,32(4):1230-1240.
[20]ZHANG X K,XIAO Y G,ZHANG Y,XIA X C,DUBCOVSKY J,HE Z H.Allelic variation at the vernalization genes Vrn-A1,Vrn-B1,Vrn-D1,and Vrn-B3 in Chinese wheat cultivars and their association with growth habit.Crop Science,2008,42(2):1690-1694.
[21]安艳荣.二穗短柄草VILs基因对开花时间的调控机理研究.青岛:山东农业大学博士学位论文,2015.AN Y R.Mechanism of regulation on flowering time by VIL genes in Brachypodium distachyon.PhD Thesis.Qingdao:Shandong Agricultural University,2015.
[22]YOSHIDA T,NISHIDA H,ZHU J,NITCHER R,DISTELFELD A,AKASHI Y,KATO K,DUBCOVSKY J.Vrn-D4,is a vernalization gene located on the centromeric region of chromosome 5D in hexaploid wheat.Theoretical&Applied Genetics,2010,120(3):543-552.
[23]PIDAL B,YAN L,FU D,ZHANG,TRANQUILLI G,DUBCOVSKY J.The CArG-box located upstream from the transcriptional start of wheat vernalization gene VRN1 is not necessary for the vernalization response.Journal of Heredity,2009,100(3):355-364.
[24]DUBCOVSKY J,CHEN C,YAN L.Molecular characterization of the allelic variation at the VRN-H2 vernalization locus in barley.Molecular Breeding,2005,15(4):395-407.
[25]YAN L,LOUKOIANOV A,BLECHL A,TRANQUILLI G,RAMAKRISHNA W,SANMIGUEL P,BENNETZEN J L,ECHENIQUE V,DUBCOVSKY J.The wheat VRN2 gene is a flowering repressor downregulated by vernalization.Science,2004,303(5664):1640-1644.
[26]KIPPES N,ZHU J,CHEN A,VANZETTI L,LUKASZEWSKI A,NISHIDA H,KATO K,DVORAK J,DUBCOVSKY J.Fine mapping and epistatic interactions of the vernalization gene VRN-D4 in hexaploid wheat.Molecular Genetics and Genomics,2014,289(1):47-62.
[27]MARQUARDT S,BOSS P K,HADFIELD J,DEAN C.Additional targets of the Arabidopsis autonomous pathway members,FCAand FY.Journal of Experimental Botany,2006,57(13):3379-3386.
[28]CHENG J Z,ZHOU Y P,LV T X,XIE C P,TIAN C E.Research progress on the autonomous flowering time pathway in Arabidopsis.Physiology&Molecular Biology of Plants,2017,23(3):1-9.
[29]SIMPSON G G.The autonomous pathway:Epigenetic and post-transcriptional gene regulation in the control of Arabidopsis flowering time.Current Opinion in Plant Biology,2004,7(5):570-574.
[30]EOM H,PARK S J,KIM M K,KIM H,KANG H,LEE I.TAF15b,involved in the autonomous pathway for flowering,represses transcription of FLOWERING LOCUS C.Plant Journal for Cell&Molecular Biology,2017,93(1):79-91.
[31]YABUTA T,HAYASI T.Biochemical studies of'bakanae'fungus of rice.Journal of the Imperial Agricultural Experimental Station Nisigahara Tokyo,1940,25(3):365-400.
[32]黄桃鹏,李媚娟,王睿,李玲.赤霉素生物合成及信号转导途径研究进展.植物生理学报,2015(8):1241-1247.HUANG T P,LI M J,WANG R,LI L.Progress in study of gibberellins biosynthesis and signaling transduction pathway.Plant Physiology Journal,2015(8):1241-1247.
[33]BLAZQUEZ M A,GREEN R,NILSSON O,SUSSMAN M R,WEIGEL D.Gibberellins promote flowering of Arabidopsis by activating the LEAFY promoter.Plant Cell,1998,10(5):791-800.
[34]XUE L J,ZHANG J J,XUE H W.Genome-wide analysis of the complex transcriptional networks of rice developing seeds.PloSOne,2012,7(2):e31081.
[35]黄磊玉.植物开花基因的研究进展.农家科技,2017(3):257.HUANG L Y.Research progress of flowering genes in plants.Nongjia Keji,2017(3):257.
[36]李昱,罗志鹏,赵淑清.拟南芥开花时间调控的整合途径.植物生理学报,2007,43(5):799-804.LI Y,LUO Z P,ZHAO S Q.Integration pathway of flowering time control in Arabidopsis.Plant Physiology Journal,2007,43(5):799-804.
[37]袁敏,邢朝斌,葛伟娜,王莉,郭棣.拟南芥开花诱导基因FT的蛋白表达及纯化.基因组学与应用生物学,2017,36(8):3053-3056.YUAN M,XING C B,GE W N,WANG L,GUO D.Protein expression and purification of flowering inducer gene FT in Arabidopsis thaliana.Genomics and Applied Biology,2017,36(8):3053-3056.
[38]兰树斌,李建国.植物LFY基因的研究进展.基因组学与应用生物学,2007,26(s1):132-137.LAN S B,LI J G.Advance of studies on LFY gene in plants.Genomics and Applied Biology,2007,26(s1):132-137.
[39]王利琳,梁海曼,庞基良,朱睦元.拟南芥LEAFY基因在花发育中的网络调控及其生物学功能.遗传,2004,26(1):137-142.WANG L L,LIANG H M,PANG J L,ZHU M Y.Regulation network and biological roles of LEAFY in Arabidopsis thaliana in floral development.Hereditas,2004,26(1):137-142.
[40]SAMACH A,ONOUCHI H,GOLD S E,DITTA G S,SCHWARZ-SOMMER Z,YANOFSKY M F,COUPLAND G.Distinct roles of CONSTANS target genes in reproductive development of Arabidopsis.Science,2000,288:1613-1616.
[41]HEPWORTH S R,VALVERDE F,RAVENSCROFT D,MOURADOV A,COUPLAND G.Antagonistic regulation of floweringtime gene SOC1 by CONSTANS and FLC via separate promoter motifs.Embo Journal,2002,21(16):4327-4337.
[42]SEARLE I,HE Y,TURCK F,VINCENT C,FORNARA F,KR?BER S,AMASINO R A,COUPLAND G.The transcription factor FLC confers a flowering response to vernalization by repressing meristem competence and systemic signaling in Arabidopsis.Genes&Development,2006,20(7):898-912.
[43]SCHULTZ E A,HAUGHN G W.LEAFY,a homeotic gene that regulates inflorescence development in Arabidopsis.Plant Cell,1991,3(8):771-781.
[44]LOHMANN J U,HONG R L,HOBE M,BUSCH M A,PARCY F,SIMON R,WEIGEL D.A molecular link between stem cell regulation and floral patterning in Arabidopsis.Cell,2001,105(6):793-803.
[45]GUSTAFSON-BROWN C,SAVIDGE B,YANOFSKY M F.Regulation of the arabidopsis floral homeotic gene APETALA1.Cell,1994,76(1):131-143.
[46]SHELDON C C,ROUSE D T,FINNEGAN E J,PEACOCK W J,DENNIS E S.The molecular basis of vernalization:The central role of FLOWERING LOCUS C(FLC).Proceedings of the National Academy of Sciences of the United States of America,2000,97(7):3753-3758.
[47]MICHAELS S D,AMASINO R M.FLOWERING LOCUS C encodes a novel MADS domain protein that acts as a repressor of flowering.Plant Cell,1999,11(5):949-956.
[48]PUTTERILL J,ROBSON F,LEE K,SIMON R,COUPLAND G.The CONSTANS gene of Arabidopsis promotes flowering and encodes a protein showing similarities to zinc finger transcription factors.Cell,1995,80(6):847-857.
[49]SK?T L,SANDERSON R,THOMAS A,SK?T K,THOROGOOD D,LATYPOVA G,ASP T,ARMSTEAD I.Allelic variation in the perennial ryegrass FLOWERING LOCUS T gene is associated with changes in flowering time across a range of populations.Plant Physiology,2011,155(2):1013-1022.
[50]FIIL A,LENK I,PETERSEN K,JENSEN C S,NIELSEN K K,SCHEJBEL B,ANDERSEN J R,LüBBERSTEDT T.Nucleotide diversity and linkage disequilibrium of nine genes with putative effects on flowering time in perennial ryegrass(Lolium perenne L.).Plant Science,2011,180(2):228-237.
[51]WANG J,FORSTER J W.Flowering time regulation in perennial ryegrass.Euphytica,2017,213(5):106.
[52]XIE W G,ROBINS J G,BUSHMAN B S.A genetic linkage map of tetraploid orchardgrass(Dactylis glomerata L.)and quantitative trait loci for heading date.Genome,2012,55(5):360-369.
[53]谢文刚.鸭茅分子遗传连锁图谱构建及开花基因定位.成都:四川农业大学博士学位论文,2013.XIE W G.Genetic linkage map and flowering time gene mapping in orchardgrass(Dactylis glomerata L.).PhD Thesis.Chengdu:Sichuan Agricultural University,2013.
[54]ZHAO X X,HUANG L K,ZHANG X Q,WANG J P,YAN D F,LI J,TANG L,LI X L,SHI T W.Construction of high-density genetic linkage map and identification of flowering-time QTLs in orchardgrass using SSRs and SLAF-seq.Scientific Reports,2016,6:29345.
[55]ZHAO X X,BUSHMAN B S,ZHANG X Q,ROBBINS M D,LARSON S R,ROBINS J G,THOMAS A.Association of candidate genes with heading date in a diverse Dactylis glomerata population.Plant Science,2017,265:146-153.
[56]FENG G,HUANG L,LI J,WANG J,XU L,PAN L,ZHAO X,WANG X,HUANG T,ZHANG X.Comprehensive transcriptome analysis reveals distinct regulatory programs during vernalization and floral bud development of orchardgrass(Dactylis glomerata L.).BMC Plant Biology,2017,17(1):216.
[57]陈锡,赵德刚,陈莹,李小冬,吴佳海,王小利.高羊茅FaFT2基因克隆及表达分析.植物生理学报,2017(8):1523-1531.CHEN X,ZHAO D G,CHEN Y,LI X D,WU J H,WANG X L.Cloning and expression analysis of FaFT2 gene in tall fescue.Plant Physiology Journal,2017(8):1523-1531.
[58]王小利,陈伟,李晚忱,吴佳海,刘晓霞,杨义成.高羊茅春化基因FaVRN1的克隆与分析.核农学报,2009,23(5):778-784.WANG X L,CHEN W,LI W C,WU J H,LIU X X,YANG Y C.Cloning and characterization of vernalizational gene FaVRN1from tall fescue.Journal of Nuclear Agricultural Sciences,2009,23(5):778-784.
[59]SHINOZUKA H,HAND M L,COGAN N O,SPANGENBERG G C,FORSTER J W.Nucleotide diversity of vernalization and flowering-time-related genes in a germplasm collection of meadow fescue(Festuca pratensis Huds.)Darbysh.).Ecology&Evolution,2013,3(13):4415-4426.
[60]ZENG F,BILIGETU B,COULMAN B,SCHELLENBERG M P,FU Y B.RNA-Seq analysis of gene expression for floral development in crested wheatgrass(Agropyron cristatum L.).PLoS One,2017,12(5):e0177417.
[61]LOMAX A,WOODS D P,DONG Y,BOUCHéF,RONG Y,MAYER K S,ZHONG X,AMASINO R M.An ortholog of CURLYLEAF/ENHANCER OF ZESTE like-1 is required for proper flowering in Brachypodium distachyon.Plant Journal for Cell&Molecular Biology,2018,93(5):871-882.
[62]GAO R,GRUBER M Y,AMYOT L,HANNOUFA A.SPL13 regulates shoot branching and flowering time in Medicago sativa.Plant Molecular Biology,2018,96(1/2):1-15.
[63]AUNG B,GRUBER M Y,HANNOUFA A.The MicroRNA156 system:A tool in plant biotechnology.Biocatalysis&Agricultural Biotechnology,2015,4(4):432-442.
[64]LIU M,LEI L,MIAO F,POWERS C,ZHANG X,DENG J,TADEGE M,CARVER B F,YAN L.The STENOFOLIA gene from Medicago alters leaf width,flowering time and chlorophyll content in transgenic wheat.Plant Biotechnology Journal,2018,16(1):186-196.
[65]KOVI M R,AMDAHL H,ALSHEIKH M,ROGNLI O A.De novo and reference transcriptome assembly of transcripts expressed during flowering provide insight into seed setting in tetraploid red clover.Scientific Reports,2017,7:44383.
[66]NAVARRO M P,RIBALTA F M,HURGOBIN B,CROSER J S,KAUR P.Gene networks underlying faster flowering induction in response to far-red light.BioRxiv,2017.
[67]ADHIKARI K,BUIRCHELL B,YAN G,SWEETINGHAM M.Two complementary dominant genes control flowering time in albus lupin(Lupinus albus L.).Plant Breeding,2011,130(4):496-499.