小麦花器官形成的A,B,C,D,E功能基因研究进展
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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.
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
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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
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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
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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
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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
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Bowman J.L.,Smyth D.R.,and Meyerowitz E.M.,1989,Gene directing flower development in Arabidopsis,The Plan Cell,1(1):37-52
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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
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