摘要
小麦是人类重要的粮食作物之一,花器官是粮食产生的基础。小麦花器官发育研究对小麦育种有着潜在的应用前景。目前,已鉴定克隆了部分小麦花器官A,B,C,D,E 5类功能基因,加深了人们对小麦花器官发育机制的认识。A功能基因主要有WFUL1,2,3/WAP1/VRN1/TaVRT1/TaAP1-1,2,3,与小麦由营养生长向生殖生长转变以及内外稃和浆片的发育有关。B功能基因有WAP3/TaAP3/TaMADS 51、TaMADS 82、WPI1/TaPI-1、WPI2/TaPI-2/TaAGL26,决定浆片和雄蕊的发育。C功能基因参与雄蕊和雌蕊的发育,包括WAG-1/TaAG-1、WAG-2/TaAG-2/WM29A/WM29B/TaAGL39。D功能基因在胚珠发育中起重要作用,主要有TaAG-3/TaAGL2/TaAGL9/TaAGL31、TaAG-4和WSTK。E功能基因在各轮花器官中均有一定作用,主要有WSEP、WLHS1、TaMADS1、TaSEP-1,2,3,4,5,6,以及TaAGL37/TaAGL6-1A,TaMADS#12/TaAGL6-1B和TaAGL6-1C。相比于双子叶植物,小麦花器官发育仍有许多问题尚需阐明。本文综述了小麦花器官形成的A,B,C,D,E功能基因研究现状,总结小麦花器官形成的ABCDE模型,并提出目前研究存在的问题,为深入研究小麦花器官发育提供一定的理论依据。
Wheat is one of the most important food crops in the world. The wheat floral organs provide the basis for grain formation. Study of wheat floral organ development has a potential application prospect for seed plant breeding. At present, some A, B, C, D, E function genes concerning floral organ development in wheat have been cloned, which makes researchers further realize the development mechanism of wheat floral organ. The class A genes are WFUL1,2,3/WAP1/VRN1/TaVRT1/TaAP1-1,2,3, which are associated with the transition from vegetative to reproductive growth and development of lemma and lodicule. The class B genes are WAP3/TaAP3/TaMADS 51, TaMADS 82, WPI1/TaPI-1and WPI2/TaPI-2/TaAGL26, which play a central role in development of lodicule and stamen. WAG-1/TaAG-1 and WAG-2/TaAG-2/WM29A/WM29B/TaAGL39 are members of the class C genes and act in pistil and stamen formation. D-class genes TaAG-3/TaAGL2/TaAGL9/TaAGL31, TaAG-4 and WSTK promote ovule identity. Some class E genes(WSEP, WLHS1, TaMADS1, TaSEP-1,2,3,4,5,6, TaMADS#12/TaAGL6-1B, TaAGL37/TaAGL6-1A and TaAGL6-1C) have been found, which show partially redundant functions in identity determination of lemma, lodicule, stamens and carpels. However, compared to the dicots, the regulation and mechanism of specific genes involved in flower development of wheat are still unknown and need to further be elucidated. In the present review, the roles of A, B, C, D, E class genes in wheat are summarized. The current understanding of ABCDE model in wheat is described, and the problems existing in current research are presented, which would play important roles in further research of floral organ formation in wheat.
引文
Adam H.,Jounannic S.,Morcillo F.,Verdeil J.L.,Duval Y.,and Tregear J.W.,2007,Determination of flower structure in Elaeis guineesis:Do palms use the same homoetic genes a other species?Annual of Botany,100(1):1-12
Airoldi C.A.,2010,Determination of sexual organ development Sexural Plant Reproduction,23(1):53-62
Ambrose B.A.,Lerner D.R.,Ciceri P.,Padilla C.M.,Yanofsk M.F.,and Schmidt R.J.,2000,Molecular and genetic analysi of the Silky1 gene reveals conservation in floral organ speciation between eudicots and monocots,Molecular Cell5(3):569-579
Angenent G.C.,Busscher M.,Franken J.,Mol J.N.M.,and va Tunen A.J.,1992,Differential expression of two MADS bo genes in wild-type and mutant petunia flowers,The Plan Cell,4(8):983-993
Becker A.,Kaufmann K.,Freialdenhoven A.,Vincent C.,Li MA.,Saedler H.,and Theiβen G.,2002,A novel MADS-box gene subfamily with a sister-group relationship to class Bfloral homeotic genes,Molecular Genetics and Genomics266(6):942-950
Bowman J.L.,Smyth D.R.,and Meyerowitz E.M.,1989,Gene directing flower development in Arabidopsis,The Plan Cell,1(1):37-52
Coen E.S.,and Meyerowitz E.M.,1991,The war of the whorls genetic interactions controlling flower development,Na ture,353(6339):31-37
Danyluk J.,Kane N.A.,Breton G.,Limin A.E.,Fowler D.B.,and Sarhan F.,2003,Ta VRT-1,a putative transcription facto associated with vegetative to reproductive transition in cere als,Plant Physiol.,132(4):1849-1860
Hama E.,Takumi S.,Ogihara Y.,and Murai K.,2004,Pistillod is caused by alterations to the class-B MADS-box gene expression pattern in alloplasmic wheats,Planta,218(5)712-720
Honma T.,and Goto K.,2001,Complexes of MADS-box protein are sufficient to convert leaves into floral organs,Nature409(6819):525-529
Kang H.G.,Jeon J.S.,Lee S.C.,and An G.H.,1998,Identificatio of class B and class Cfloral organ identity genes from ric plants,Plant Molecular Biology,38(6)1021-1029
Kanno A.,Nakasda M.,Akita Y.,and Hirai M.,2007,Class Bgene expression and the modified ABC model in nongras monocots,The Scientific World Journal,7(68):268-279
Kinjo H.,Shitsukawa N.,Takumi S.,and Murai K.,2012,Divers ification of three APETALA1/FRUITFULL-like genes in wheat,Molecular Genetics and Genomics,287(4):283-294
Kramer E.M.,Dorit R.L.,and Irish V.F.,1998,Molecular evolu tion of genes controlling petal and stamen development Duplication and divergence within the APETALA3 and PISTILLATA MADS-box gene lineages,Genetics,1998149(2):765-783
Kyozuka J.,and Shimamoto K.,2002,Ectopic expression of Os MADS3,a rice ortholog of AGAMOUS,caused a homeo tic transformation of lodicules to stamens in transgenic ric plants,Plant and Cell Physiology,43(1):130-135
Kyozuka J.,Kobayashi T.,Morita M.,and Shimamoto K.,2000Spatially and temporally regulated expression of rice MADSbox genes with similarity to Arabidopsis class A,B and Cgenes,Plant and Cell Physiology,41(6):710-718
Lee D.Y.,and An G.H.,2012,Two AP2 family genes,SUPERN-UMERARY BRACT(SNB)and Os INDETERMINATE SPIKELET 1(Os IDS1),synergistically control inflorescence ar chitecture and floral meristem establishment in rice,Plant J.69(3):445-461
Liao M.L.,Peng Z.S.,Yang Z.J.,Wei S.H.,and Martinek P.2015,Identification of differentially expressed genes in pistillody common wheat mutant using an annealing contro primer system,Genet.Mol.Res.,14(2):3995-4004
Lopez-Dee Z.P.,Wittich P.,Enrico PèM.,Rigola D.,Del Buon I.,Sari-Gorla M.,Kater M.M.,and Colombo L.,1999Os MADS13,a novel rice MADS-box gene expressed durin ovule development,Dev.Genet.,25(3):237-244
Meguro A.,Takumi S.Y.,Ogihara Y.,and Murai K.,2003WAG,a wheat AGAMOUS homolog,is associated with development of pistil-like stamens in alloplasmic wheats Sexural Plant Reproduction,15(5):221-230
Mena M.,Mandel M.A.,Lerner D.R.,Yanofsky M.F.,an Schimidt R.J.,1995,A characterization of the MADS-box gene family in maize,The Plant Journal,8(6):845-854
Mizumoto K.,Hatano H.,Hirabayashi C.,Murai K.,and Takum S.,2009,Altered expression of wheat AINTEGUMENTAhomolog,WANT-1,in pistil and pistil-like transformed stamen of an alloplasmic line with Aegilops crassa cytoplasm Development Genes and Evolution,219(4):175-187
Murai K.,2013,Homeotic Genes and the ABCDE model for flora organ formation in wheat,Plants,2(3):379-395
Murai K.,and Tsunewaki K.,1993,Photoperiod-sensitive cytopl asmic male sterility in wheat with Aegilops crassa cytoplasm Euphytica,67(1):41-48
Murai K.,Miyamae M.,Kato H.,Takumi S.,and Ogihara Y.2003,WAP1,a wheat APETALA1 homolog,plays a centra role in the phase transition from vegetative to reproductiv growth,Plant and Cell Physiology,44(12):1255-1265
Murai K.,Takumi S.,Koga H.,and Ogihara Y.,2002,Pistillody homeotic transformation of stamens into pistil-like struc tures,caused by nuclear-cytoplasm interaction in wheat,Th Plant Journal,29(2):169-181
Nair S.K.,Wang N.,Turuspekov Y.,Pourkheirandish M.,Sinsu wongwat S.,Chen G.,Sameri M.,Tagiri A.,Honda I.Watanabe Y.,Kanamori H.,Wicker T.,Stein N.,Nagamur Y.,Matsumoto T.,and Komatsuda T.,2010,Cleistogamou flowering in barley arises from the suppression of micro RNAguided Hv AP2 mR NA cleavage,Proc.Natl.Acad.Sci.USA107(1):490-495
Ning S.,Wang N.,Sakuma S.,Pourkheirandish M.,Wu J.Z.Matsumoto T.,Koba T.,and Komatsuda T.,2013,Structure transcription and post-transcriptional regulation of the bread wheat orthologs of the barley cleistogamy gene Cly1,Theor Appl.Genet.,126(5):1273-1283
Ogihara Y.,Mochida K.,Nemoto Y.,Murai K.,Yamazaki Y.Shin-I T.,and Kohara Y.,2003,Correlated clustering and virtual display of gene expression patterns in the wheat lif cycle by large-scale statistical analyses of expressed sequenc tags,The Plant Journal,33(6):1001-1011
Paolacci A.R.,Tanzarella O.A.,Porceddu E.,Varotto S.,and Ciaffi M.,2007,Molecular and phylogenetic analysis o MADS-box genes of MIKC type and chromosome location of SEP-like genes in wheat(Triticum aestivum L.),Molecula Genetics and Genomics,278(6):689-708
Pelaz S.,Ditta G.S,Baumann E.,Wisman E.,and Yanofsky MF.2000,B and C floral organ identity functions require SEPALLATA MADS-box genes,Nature,405(6783):200-203
Pelaz S.,Tapia-Lopez R.,Alvarez-Buylla E.R.,and Yanofsky MF.,2001,Conversion of leaves into petals in Arabidopsis Current Biology,11(6):182-184
Pelucchi N.,Fornara F.,Favalli C.,Masiero S.,Lago C.,Colomb L.,and Kater M.M.,2002,Comparative analysis of ric MADS-box genes expressed during flower development Sexural Plant Reproduction,15(3):113-122
Peng Z.S.,2003,A new mutaion in wheat producing three pistil in a floret,Journal of Agromony&Crop Science,189(4)270-272
Peng Z.S.,Martinek P.,Kosuge K.,Kuboyama T.,and Watanab N.,2008,Genetic mapping of a mutant gene producing thre pistils per floret in common wheat,Journal of Applied Genetics,49(2):135-139
Peng Z.S.,Yang J.,Wei S.H.,and Zeng J.H,2004,Characterization of the common wheat(Triticum aestivum L.)mutation line producing three pistils in a floret,Hereditas(Sweden)141(1):15-18
Peng Z.S.,Yang Z.J.,Ouyang Z.M.,and Yang H.,2013,Charac terization of a novel pistillody mutation in common wheat Australian Journal of Crop Science,7(1):159-164
Ren D.Y.,Li Y.F.,Zhao F.M.,San X.C.,Shi J.Q.,Wang N.,Guo S.,Ling Y.H.,Zhang C.W.,Yang Z.L.,and He G.H.,2013MULTI-FLORET SPIKELET1,which encodes and AP2/ERFprotein,determines spikelet meristem gate and sterile lemm identity in rice,Plant Physiol.,162(2):872-884
Shitsukawa N.,Tahira C.,Kassai K.I.,Hirabayashi C.,Shimizu T.,Takumi S.,Mochida K.,Kawaura K.,Ogihara Y.,and Murai K.,2007,Genetic and epigenetic alteration among three homoeologous genes of a classs E MADS box gene in hexaploid wheat,The Plant Cell,19(6):1723-1737
Smaczniak C.,Immink R.G.H.,Muino J.M.,Blanvillain R.,Buss cher M.,Busscher-Lange J.,Dinh Q.D.P.,Liu S.J.,Westpha A.H.,Boeren S.,Parcy F.,Xu L.,Carles C.C.,Angenen G.C.,and Kaufmann K.,2012,Characterization of MADSdomain transcription factor complexes in Arabidopsis flowe development,Proc.Natl.Acad.Sci.USA,109(5):1560-1565
Thei覻en G.,Strater T.,Fischer A.,and Saedler H.,1995,Structura characterization,chromosomal localization and phylogeneti evaluation of two pairs of AGAMOUS-like MADS-box genes from maize,Gene,156(2):155-166
Wang Q.H.,Yang Z.J.,Wei S.H.,Jiang Z.Y.,Yang Y.F.,Hu Z.SSun Q.X.,and Peng Z.S.,2015,Molecular cloning,characte rization and expression analysis of WAG-1 in the pistillody line of common wheat,Genetics and Molecular Research14(4):12455-12465
Wang Y.W.,Peng Z.S.,Yang Z.J.,Wei S.H.,Liao M.l.,Zhao H.Yang Y.F.,and Wang Q.H.,2014,Cloning and Expression Analysis of Ta AP1-3 Genes of Three Pistils Mutation in Wheat,Huabei Nongxue Bao(Acta Agriculturae Boreali Sinica),29(6):28-35(王育伟,彭正松,杨在君,魏淑红,廖明莉,赵欢,杨会,杨宇凤,王清海,2014,小麦三雌蕊突变体Ta AP1-3基因的克隆及表达分析,华北农学报,29(6)28-35)
Wei S.H.,2013,Cloning,Chromosomal location and differentia expression of a flower development gene WAG-2 in commo wheat and its relatives,Dissertation for Ph.D.,Sichuan Agriculture University,Supervisor:Peng Z.S.,and Zhou Y.H.pp.100-114(魏淑红,2013,普通小麦及其近缘种花发育基因WAG-2的克隆、定位及差异表达分析,博士学位论文四川农业大学,导师:彭正松,周永红,pp.100-114)
Wei S.H.,Peng Z.S.,and Yang Z.J.,2015,A simple approach based on intron length polymorphism(ILP)for chromosoma localization of Gene WAG-2,Indian Journal of Genetics and Plant Breeding,75(3):314-317
Wei S.H.,Peng Z.S.,Zhou Y.H.,Yang Z.J.,Wu K.,and Ouyang Z.M.,2011,Nucleotide diversity and molecular evolution o the WAG-2 gene in common wheat(Triticum aestivum L)and its relatives,Genetics and Molecular Biology,34(4):605-615
Wendel J.F.,2000,Genome evolution in polyploids,Plant Molec ular Biology,42(1):225-249
Wu K.,Peng Z.S.,Yang Z.J.,Wei S.H.,Zeng Q.L.,Liao M.L.and Ouyang Z.M.,2012,Differential Expression of WAGfrom the Common Wheat Line Three Pistils and China Spring,Xihua Shifan Daxue Xuebao(Journal of China Wes Normal University(Natural Science Edition)),33(2):5-11(吴凯,彭正松,杨在君,魏淑红,曾巧玲,廖明莉,欧阳钟鸣,2012,WAG1基因在小麦三雌蕊品系和中国春中的差异表达分析,西华师范大学学报(自然科学版),33(1):5-11)
Yamada K.,Saraike T.,Shitsukawa N.,Hirabayashi C.,Takum S.,and Murai K.,2009,Class D and Bsister MADS-box genes are associated with ectopic ovule formation in the pistil-like stamens of alloplasmic wheat(Triticum aestivum L.),Plant Molecular Biology,71(1):1-14
Yamaguchi T.,Lee D.Y.,Miyao A.,Hirochika H.,An G.,and Hirano H.Y.,Functional diversification of the two C-Clas MADS-box genes OSMADS3 and OSMADS58 in Oryza sativa The Plant Cell,2006,18(1):15-28
Yan L.,Loukoianov A.,Tranquilli G.,Helguera M.,Fahima T.and Dubcovsky J.,2003,Positional cloning of wheat vernalization gene VRN1,Proc.Natl.Sci.Acad.USA,100(10)6263-6268
Yang Y.F.,Peng Z.S.,Yang Z.J.,Wei S.H.,Liao M.L.,Wang YW.,Wang Q.H.,and Yang H.,2015,Cloning and expression analysis of class-B MADS-box genes WPI1 and WPI2 in wheat,Henong Xuebao(Journal of Nuclear Agricultural Sci ences),29(2):221-228(杨宇凤,彭正松,杨在君,魏淑红廖明莉,王育伟,王清海,杨会,2015,小麦B类MADS-bo基因WPI1和WPI2的克隆及表达分析,核农学报,29(2)221-228)
Yang Z.J.,Peng Z.S.,Wei S.H.,Yu Y.,and Cai P.,2011a,Identification of differentially expressed genes in three-pistil mu tation in wheat using annealing control primer system,Gene485(2):81-84
Yang Z.J.,Peng Z.S.,Wei S.H.,Liao M.l.,Yu Y.,and Jang Z.Y.2015,Pistillody mutant reveals key insights into stamen and pistil development in wheat(Triticum aestivum L.),BMCGenomics,16(1):211-220
Yang Z.J.,Peng Z.S.,Yang H.,Yang J.,and Wei S.H.,2011b Suppression subtractive hybridization tdentified differen tially expressed genes in pstil mutations in wheat,Plan Molecular Biology Reporter,29(2):431-439
Zhang Z.C.,Belcram H.,Gornicki P.,Charles M.,Just J.,Huneau C.,Magdelenat G.,Couloux A.,Samain S.,Gill B.S.Rasmussen J.B.,Barbe V.,Faris J.D.,and Chalhoub B.2011,Duplication and partitioning in evolution and function of homoeologous Q loci governing domestication character in polyploidy wheat,Proc.Natl.Acad.Sci.USA,108(46)18737-18742
Zhao T.,Ni Z.F.,Dai Y.,Yao Y.Y.,Nie X.L.,and Sun Q.X.2006b,Characterization and expression of 42 MADS-box genes in wheat(Triticum aestivum L.),Molecular Genetic and Genomics,276(4):334-356
Zhao X.Y.,Cheng Z.J.,and Zhang X.S.,2006a,Overexpression of Ta MADS1,a SEPALLATA-like gene in wheat,cause early flowering and the abnormal development of flora organs in Arabidopsis,Planta,223(4):698-707