水稻FIS类多梳蛋白基因启动子特性及基因表达分析
|
摘要
基因组印迹是一种非孟德尔遗传现象。它是指在配子或合子发生期间,来自亲本的等位基因或染色体产生专一性的加工修饰,导致后代体细胞中两个亲本来源的等位基因有不同的表达活性。印迹基因的特点就是基因的表达与否决定于其亲本来源,导致两个不同亲源的等位基因具有不同的表型效应,即具有亲源效应。在植物中已经发现了一些对早期胚乳发育的启动有重要作用并与亲本印记有关的基因,如拟南芥中的基因FERTILIZATION-INDEPENDENT ENDOSPERM(FIE)、FERTILIZATION-INDEPENDENT SEED1/MEDEA(FIS1/MEA)和EMF2这些基因编码的蛋白属于多梳家族蛋白(polycomb group protein, PcG)。玉米中的PcG类基因(FIE1、FIE2、Mez1、Mez2)也参与胚乳发育的调控。而水稻是世界主要的粮食作物之一,也是单子叶模式植物,其主要食用部分胚乳在胚胎发育过程中起着至关重要的作用。对水稻中PcG类基因(EMF,FIE2a,FIE2b,EZ1,EZ3)的研究,将有助于认识禾谷类作物胚乳发育的机制,并有潜在的育种价值。
为了研究水稻胚乳中PcG类基因的印记模式,本试验通过两个方面的研究思路试图解决这个问题。
一、对FIS类基因的启动子片段,用生物信息学分析其功能区段,并将启动子片段分别连上带有GFP和GUS两种不同标记基因的表达载体上。分离相关的启动子并尝试将启动子片段装入含有报告基因的启动子表达载体可以为转基因植株研究提供载体平台,这对精细研究FIS基因及调控胚乳发育的分子机制大有帮助。而这部分的实验也分别获得了相应的启动子GFP和GUS两套表达载体,为以后的研究打下基础。
二、将粳稻品种日本晴和籼稻品种9311正反杂交,取授粉后一定时期内的花序材料剥离胚和胚乳,以此为材料分析胚和胚乳中多梳蛋白(polycomb group protein, PcG)的表达。通过这部分的实验发现大致可以判断FIE2b和EZ1两个基因在受精后一段时间内胚乳中的表达量是有增加的趋势。从某种程度上证明这两个基因与胚乳的发育相关,而与印迹也有很微妙的联系。
Genomic imprinting is a genetic phenomenon by which certain genes are expressed in a parent-of-origin-specific manner. It is an inheritance process independent of the classical Mendelian inheritance. Imprinted genes are either expressed only from the allele inherited from the mother, or in other instances from the allele inherited from the father. Forms of genomic imprinting have been demonstrated in insects, mammals and flowering plants(eg. FIE、FIS1/MEA and EMF2 in Arabidopsis thaliana; FIE1、FIE2、Mezl and Mez2 in Zea mays).As one of the most important crops of the world, Oryza sativa is a model of monocotyledon plant. The endoperm is the main edible part of rice which takes a crucial role in the embryo development. It is useful to carry out some necessary researches on PcG genes of rice, because it is helpful to understand the mechanism of endosperm development in cereal which has a potential value in breeding.
For the purpose of making clear the mechanism of PcG genes of rice. I try to make it clear in two different ways.
In one hand,bioimformation method was used to analyst the promoter of PcG genes in rice. After that I construct the expression vector use two vector which contain GFP and GUS.
In the other hand, positive and negative crossbreed was used to study the expressing of FIS class gene.After crossbreeding in 9311 and Nipponbare, I take inflorescences of rice after pollination, and separate the endsperm and embryo. Then by the tool of RT PCR, I can inspect the expression level of FIS class gene. And the result indicates that gene FIE2b and EZ1 maybe is related to genome imprinting.
为了研究水稻胚乳中PcG类基因的印记模式,本试验通过两个方面的研究思路试图解决这个问题。
一、对FIS类基因的启动子片段,用生物信息学分析其功能区段,并将启动子片段分别连上带有GFP和GUS两种不同标记基因的表达载体上。分离相关的启动子并尝试将启动子片段装入含有报告基因的启动子表达载体可以为转基因植株研究提供载体平台,这对精细研究FIS基因及调控胚乳发育的分子机制大有帮助。而这部分的实验也分别获得了相应的启动子GFP和GUS两套表达载体,为以后的研究打下基础。
二、将粳稻品种日本晴和籼稻品种9311正反杂交,取授粉后一定时期内的花序材料剥离胚和胚乳,以此为材料分析胚和胚乳中多梳蛋白(polycomb group protein, PcG)的表达。通过这部分的实验发现大致可以判断FIE2b和EZ1两个基因在受精后一段时间内胚乳中的表达量是有增加的趋势。从某种程度上证明这两个基因与胚乳的发育相关,而与印迹也有很微妙的联系。
Genomic imprinting is a genetic phenomenon by which certain genes are expressed in a parent-of-origin-specific manner. It is an inheritance process independent of the classical Mendelian inheritance. Imprinted genes are either expressed only from the allele inherited from the mother, or in other instances from the allele inherited from the father. Forms of genomic imprinting have been demonstrated in insects, mammals and flowering plants(eg. FIE、FIS1/MEA and EMF2 in Arabidopsis thaliana; FIE1、FIE2、Mezl and Mez2 in Zea mays).As one of the most important crops of the world, Oryza sativa is a model of monocotyledon plant. The endoperm is the main edible part of rice which takes a crucial role in the embryo development. It is useful to carry out some necessary researches on PcG genes of rice, because it is helpful to understand the mechanism of endosperm development in cereal which has a potential value in breeding.
For the purpose of making clear the mechanism of PcG genes of rice. I try to make it clear in two different ways.
In one hand,bioimformation method was used to analyst the promoter of PcG genes in rice. After that I construct the expression vector use two vector which contain GFP and GUS.
In the other hand, positive and negative crossbreed was used to study the expressing of FIS class gene.After crossbreeding in 9311 and Nipponbare, I take inflorescences of rice after pollination, and separate the endsperm and embryo. Then by the tool of RT PCR, I can inspect the expression level of FIS class gene. And the result indicates that gene FIE2b and EZ1 maybe is related to genome imprinting.
引文
1.张文伟,江玲,曹少先,万建民.拟南芥印记基因M E A.植物生理学通讯,2005a,41(6)：710-714
2.张文伟,曹少先,江玲,朱速松,万建民.基因组印记与种子发育.遗传,2005b,27(4)：665-670
3.Agger K, Cloos P A, Christensen J, Pasini D, Rose S, Rappsilber J, Issaeva I, Canaani E, Salcini A E and Helin K UTX and JMJD3 are histone H3K27 demethylases involved in HOX gene regulation and development. Nature,2007,449:731-734
4.Allis C D, Berger S L, Cote J, Dent S,Jenuwien T, Kouzarides T, Pillus L, Reinberg D, Shi Y, Shiekhattar R, Shilatifard A, Workman J and Zhang Y New nomenclature for chromatin-modifying enzymes.Cell,2007,131:633-636
5.Alvarez-Venegas R, Pien S, Sadder M, Witmer X, Grossniklaus U and Avramova Z ATX-1,an Arabidopsis homolog of trithorax, activates flower homeotic genes.Curr Biol,2003,13:627-637
6.Arnaud P, Feil R MEDEA takes control of its own imprinting Cell,2006,124: 468-470
7.Aubert D, Chen L, Moon Y-H, Martin D, Castle L A, Yang C-H, Sung Z R. EMF1,a novel protein involved in the control of shoot architecture and flowering in Arabidopsis. The Plant Cell,2001,13:1865-1875
8.Autran D, Mamani W H, Calzada J P V.Genomic imprinting in plants:the epigenetic version of an Oedipus complex. Current Opinion in Plant Biology,2005,8:19-25
9. Baroux C, Gagliardini V, Page D R, Grossniklaus U. Dynamic regulatory interactions of polycomb group genes:MEDEA autoregulation is required for imprinted gene expression in Arabidopsis. Genetics & Development,2006,20:1081-1086
10. Baroux C, Pien S, Grossniklaus U. Chromatin modification and remodeling during early seed development. Genetics & Development,2007,17:473-479
11.Baroux C, Spillane C, Grossniklaus U. Genomic imprinting during seed development. Adv Genet,2002,46:165-214
12.Baubec T, Scheid O M. Medea in full self-control. TRENDS in Plant Science,2006, 11:469-471
13.Benhamed M, Bertrand C,Servet C and Zhou D X Arabidopsis GCN5,HD1,and TAF1/HAF2 interact to regulate histone acetylation required for light-responsive gene expression. Plant Cell,2006,18:2893-2903
14. Berndsen C E and Denu J M Catalysis and substrate selection by histone/protein lysine acetyltransferases.Curr Opin Struct Biol,2008,18:682-689
15.Berger F Endosperm:the crossroad of seed development. Current Opinion in Plant Biology,2003,6:42-50
16.Berger F, Grini P E, Schnittger A. Endosperm:an integrator of seed growth and development. Current Opinion in Plant Biology,2006,9:664-670
17.Brink R A, Cooper D C.The endosperm in seed development The Botanical Review, 1947,13(9):479-541
18.Cao R, Wang L, Wang H, Xia L, Erdjument-Bromage H, Tempst P, Jones R S and Zhang Y Role of histone H3 lysine 27 methylation in Polycomb-group silencing. Science,2002,298:1039-1043
19.Cao Y, Dai Y, Cui S and Ma L Histone H2B monoubiquitination in the chromatin of FLOWERING LOCUS C regulates flowering time in Arabidopsis.Plant Cell,2008, 20:2586-2602
20.Cartagena J A, Matsunaga S,Seki M, Kurihara D, Yokoyama M, Shinozaki K, Fujimoto S, Azumi Y, Uchiyama S and Fukui K The Arabidopsis SDG4 contributes to the regulation of pollen tube growth by methylation of histone H3 lysines 4 and 36 in mature pollen Dev Biol,2008,315:355-368
21.Chanvivattana Y, Bishopp A, Schubert D, Stock C, Moon Y-H, Goodrich Z R S a J Interaction of Polycomb-group proteins controlling flowering in Arabidopsis. Development,2004,131:5263-5276
22. Chaudhury A M, Ming L, Miller C, Craig S, Dennis E, Peacock W J. Fertilization-independent seed development in Arabidopsis thaliana. Plant Biology, 1997,(94):4223-4228
23.Choi Y, Gehring M, Johnson L, Hannon M, Harada J J, Goldberg R B, Jacobsen S E, Fischer R L. DEMETER, a DNA glycosylase domain protein, is required for endosperm gene imprinting and seed viability in Arabidopsis. Cell,2002,110:33-42
24. Cho Y H, Yoo S D and Sheen J Regulatory functions of nuclear hexokinasel complex in glucose signaling. Cell,2006,127:579-589
25.Choi Y, Gehring M, Johnson L, Hannon M, Harada J J, Goldberg R B, Jacobsen S E and Fischer R. L.DEMETER, a DNA glycosylase domain protein, is required for endosperm gene imprinting and seed viability in Arabidopsis. Cell,2002,110:33-42
26.Collinge M A, Spillane C,Kohler C, Gheyselinck J, Grossniklaus U Genetic interaction of an origin recognition complex subunit and the polycomb group gene MEDEA during seed development. The Plant Cell,2004,16:1035-1046
27. Costa L M, Gutierrez-Marcos J F, Dickinson H G. More than a yolk:the short life and complex times of the plant endosperm. TRENDS in Plant Science,2004,9(10): 507-514
28.Curtis M D, Grossniklaus U Molecular control of autonomous embryo and endosperm development. Sex Plant Reprod,2008,21:79-88
29.Danilevskaya O N, Hermon P, Hantke S,Muszynski M G, Kollipara K, Ananiev E V. Duplicated fie genes in maize:expression pattern and imprinting suggest distinct functions. The Plant Cell,2003,15:425-438
30.De Crombrugghe B,Busby S, Buc H. Cyclic AMP Receptor Protein:Role in Transcription Activation.Science,1984,224:831-838
31.De-Pater S,Greco V, Pham K, Memelink J, Kijne J.Characterization of a zinc-dependent transcriptional activator from Arabidopsis. Nucleic Acids Res,1996, 24:4624-4631
32.Degenhardt J, Tobin E M. A DNA binding activity for one of two closely defined phytochrome regulatory elements in an Lhcb promoter is more abundant in etiolated than in green plants. Plant Cell,1996,8:31-41
33.Dilkes B P, Comai L. A differential dosage hypothesis for parental effects in seed development. The Plant Cell,2004,16:3174-3180
34. Dumas C, Rogowsky P.Fertilization and early seed formation. Comptes Rendus Biologies,2008,331:715-725
35.Ding Y,Wang X, Su L, Zhai J, Cao S,Zhang D, Liu C, Bi Y, Qian Q, Cheng Z, Chu C and Cao X SDG714, a histone H3K9 methyltransferase, is involved in Tos17 DNA methylation and transposition in rice. Plant Cell,2007,19:9-22
36.Dong G, Ma D P and Li J The histone methyltransferase SDG8 regulates shoot branching in Arabidopsis. Biochem Biophys Res Commun,2008,373:659-664
37.Earley K, Lawrence R J, Pontes O, Reuther R, Enciso A J, Silva M, Neves N, Gross M, Viegas W and Pikaard C S Erasure of histone acetylation by Arabidopsis HDA6 mediates large-scale gene silencing in nucleolar dominance. Genes Dev,2006, 20:1283-1293
38.Elmen J, Lindow M, Schutz S,Lawrence M, Petri A, Obad S,Lindholm M, Hedtjarn M, Hansen H F, Berger U, Gullans S,Kearney P, Sarnow P, Straarup E M, Kauppinen S.LNA-mediated microRNA silencing in non-human primates.Nature, 2008,452:896-899
39.Feng Q, Wang H, Ng H H, Erdjument-Bromage H, Tempst P, Struhl K and Zhang Y Methylation of H3-lysine 79 is mediated by a new family of HMTases without a SET domain Curr Biol,2002,12:1052-1058
40.Feil R, Berger F Convergent evolution of genomic imprinting in plants and mammals. TRENDS in Genetics,2007,23 (4):192-199
41.Finnegan E J, Dennis E S.Isolation and identification by sequence homology of a putative cytosine methyltransferase from Arabidopsis thaliana. Nucleic Acids Res, 1993,21(10):2383-2388
42.Friedman W E The evolution of double fertilization and endsperm:an'historical' perspective. Sex Plant Reprod,1998,11:6-16
43.Garcia D, Saingery V, Chambrier P, Mayer U, Jurgens G, Berger F. Arabidopsis haiku mutants reveal new controls of seed size by endosperm. Plant Physiology,2003, 131:1661-1670
44.Gary S.A gene product required for correct initiation of segmental determination in Drosophila. Nature,1981,293(3):36-41
45.Gehring M, Choi Y, Fischer R L.Imprinting and seed development. The Plant Cell, 2004,16:S203-S213
46.Gehring M, Huh J H, Hsieh T F, Penterman J, Choi Y, Harada J J, Goldberg R B and Fischer R L DEMETER DNA glycosylase establishes MEDEA polycomb gene self-imprinting by allele-specific demethylation. Cell,2006,124:495-506
47.Gendler K, Paulsen T and Napoli C ChromDB:the chromatin database. Nucleic Acids Res,2008,36:D298-302
48.Gong Z, Morales-Ruiz T, Ariza R R, Roldan-Arjona T, David L and Zhu J K ROS1,a repressor of transcriptional gene silencing in Arabidopsis, encodes a DNA glycosylase/lyase.Cell,2002,111:803-814
49.Gonzalez D, Bowen A J, Carroll T S and Conlan R S.The transcription corepressor LEUNIG interacts with the histone deacetylase HDA19 and mediator components MED14 (SWP) and CDK8 (HEN3) to repress transcription. Mol Cell Biol,2007, 27:5306-5315
50. Goodrich J, Puangsomlee P, Martin M, Long D, Meyerowitz E M, Coupland G. A Polycomb-group gene regulates homeotic gene expression in Arabidopsis. Nature, 1997,386:44-51
51.Grimanelli D, Perotti E, Ramirez J, Leblanc O.Timing of the maternal-to-zygotic transition during early seed development in maize. The Plant Cell,2005,17: 1061-1072
52.Grossniklaus U, Philippe J, Calzada V, Hoeppner M A, Gagliano W B.Maternal control of embryogenesis by MEDEA,a polycomb group gene in Arabidopsis. Science,1998a, (280):446-450
53.Grozinger C M, Chao E D, Blackwell H E, Moazed D and Schreiber S L Identification of a class of small molecule inhibitors of the sirtuin family of NAD-dependent deacetylases by phenotypic screening. J Biol Chem,2001, 276:38837-38843
54. Guitton A-E, Page D R, Chambrier P, Lionnet C, Faure J-E, Grossniklaus U, Berger F. Identification of new members of Fertilisation Independent Seed Polycomb Group pathway involved in the control of seed development in Arabidopsis thaliana. Development,2004,131:2971-2981
55.Gutierrez-Marcos J F,Costa L M,Pra M D,Scholten S,Kranz E,Perez P,Dickinson H G. Epigenetic asymmetry of imprinted genes in plant gametes.Nature genetics, 2006,38(8):876-878
56.Hennig L, Taranto P, Walser M, Schonrock N, Gruissem W. Arabidopsis MSI1 is required for epigenetic maintenance of reproductive development. Development, 2003,130:2555-2565
57.Hermon P, Srilunchang K, Zou J, Dresselhaus T, Danilevskaya O N.Activation of the imprinted Polycomb Group Fiel gene in maize endosperm requires demethylation of the maternal allele. Plant Mol Biol,2007,64:387-395
58. Hsieh T-F, Hakim O, Ohad N, Fischer R L. From flour to flower:how Polycomb group proteins influence multiple aspects of plant development. TRENDS in Plant Science, 2003,8(9):439-445
59.Huang L, Sun Q, Qin F, Li C, Zhao Y and Zhou D X Down-regulation of a SILENT INFORMATION REGULATOR2-related histone deacetylase gene, OsSRTl,induces DNA fragmentation and cell death in rice. Plant Physiol,2007,144:1508-1519
60. Husbands A, Bell E M, Shuai B,Smith H M and Springer P S LATERAL ORGAN BOUNDARIES defines a new family of DNA-binding transcription factors and can interact with specific bHLH proteins. Nucleic Acids Res,2007,35:6663-6671
61.Huh J H, Bauer M J, Hsieh T-F, Fischer R. Endosperm gene imprinting and seed development. Current Opinion in Genetics & Development,2007,17:480-485
62.Ingouff M, Haseloff J,Berger F. Polycomb group genes control developmental timing of endosperm. The Plant Journal,2005,42:663-674
63.Jeong D-H, An S, Kang H-G, Moon S, Han J-J, Park S, Lee H S, An K, An G. T-DNA insertional mutagenesis for activation tagging in rice.Plant Physiology,2002,130: 1636-1644
64.Jeong D-H, An S,Park S, Kang H-G, Park G-G, Kim S-R, Sim J, Kim Y-O, Kim M-K, Kim S-R, Kim J, Shin M, Jung M, An G. Generation of a flanking sequence-tag database for activation-tagging lines in japonica rice. The Plant Journal,2006,45: 123-132
65.Jackson J P, Lindroth A M, Cao X and Jacobsen S E Control of CpNpG DNA methylation by the KRYPTONITE histone H3 methyltransferase. Nature,2002, 416:556-560
66.Jackson M D, Schmidt M T, Oppenheimer N J and Denu J M Mechanism of nicotinamide inhibition and transglycosidation by Sir2 histone/protein deacetylases.J Biol Chem,2003,278:50985-50998
67.Jiang D, Wang Y, Wang Y, He Y. Repression of FLOWERING LOCUS C and FLOWERING LOCUS T by the Arabidopsis Polycomb Repressive Complex 2 Components. PLoS ONE,2008,3(10):1-12
68.Jullien P E, Katz A, Oliva M, Ohad N, Berger F. Polycomb group complexes self-regulate imprinting of the polycomb group gene MEDEA in Arabidopsis. Current Biology,2006a,16:486-492
69.Jenuwein T, Laible G, Dorn R and Reuter G SET domain proteins modulate chromatin domains in eu-and heterochromatin. Cell Mol Life Sci,1998,54:80-93
70.Jeon J S,Lee S,Jung K H, Jun S H, Jeong D H, Lee J, Kim C, Jang S,Yang K, Nam J, An K, Han M J, Sung R J, Choi H S, Yu J H, Choi J H, Cho S Y, Cha S S, Kim S I and An G T-DNA insertional mutagenesis for functional genomics in rice.Plant J, 2000,22:561-570
71. Katz A, Oliva M, Mosquna A, Hakim O, Ohad N. FIE and CURLY LEAF polycomb proteins interact in the regulation of homeobox gene expression during sporophyte development. The Plant Journal,2004,37:707
72.Kim K N, Guiltinan M J. Identification of cis-acting elements important for expression of the starch-branching enzyme I gene in maize endosperm. Plant Physiol., 1999,121:225-236
73.Kim K C,Lai Z, Fan B and Chen Z Arabidopsis WRKY38 and WRKY62 transcription factors interact with histone deacetylase 19 in basal defense. Plant Cell, 2008,20:2357-2371
74.Klose R J, Kallin E M and Zhang Y JmjC-domain-containing proteins and histone demethylation. Nat Rev Genet,2006a,7:715-727
75.Kinoshita T, Harada J J, Goldberg R B, Fischer R L. Polycomb repression of flowering during early plant development. Proc Natl Acad Sci USA,2001,98: 14156-14161
76.Kinoshita T, Yadegari R, Harada J J, Goldberg R B, Fischer R L. Imprinting of the MEDEA polycomb gene in the Arabidopsis endosperm. The Plant Cell,1999,11: 1945-1952
77.Komiya R, Ikegami A, Tamaki S, Yokoi S and Shimamoto K Hd3a and RFT1 are essential for flowering in rice. Development,2008,135:767-774
78.Kouzarides T. Chromatin modifications and their function. Cell,2007,128:693-705
79.Kiyosue T, Ohad N, Yadegar R, Hannon M, Dinneny J, Wells D, Katz L M A, Harada J J, Goldberg R B,Fischer R L. Control of fertilization-independent endosperm development by the MEDEA polycomb gene in Arabidopsis. Proc Natl Acad Sci USA,1999,96:4186-4191
80.Kladnik A, Chamusco K, Dermastia M, Chourey P.Evidence of programmed cell death in post-phloem transport cells of the maternal pedicel tissue in developing Caryopsis of Maize. Plant Physiology,2004,136:3572-3581
81.Kohler C,Hennig L, Bouveret R, Gheyselinck J, Grossniklaus U, Gruissem W. Arabidopsis MSI1 is a component of the MEA/FIE Polycomb group complex and required for seed development. The EMBO Journal,2003a,22(18):4804-4814
82.Kohler C, Hennig L, Spillane C, Pien S,Gruissem W, Grossniklaus U.The Polycomb-group protein MEDEA regulates seed development by controlling expression of the MADS-box gene PHERES1.Genetics & Development,2003b: 1540-1553
83.Kohler C,Page D R, Gagliarliardini V, Grossniklaus U.The Arabidopsis thaliana MEDEA Polycomb group protein controls expression of PHERES1 by parental imprintng. Nature genetics,2005,37(1):28-30
84.Kollipara K P, Saab I N, Wych R D, Lauer M J, Singletary G W. Expression profiling of reciprocal maize hybrids divergent for cold germination and desiccation tolerance. Plant Physiology,2002,129:974-992
85.Kumar S, Tamura K and Nei M MEGA3:Integrated software for Molecular Evolutionary Genetics Analysis and sequence alignment. Brief Bioinform.,2004, 5:150-163
86. Koltunow A M, Grossniklaus U. Apomixis:A debelopmental perspective.In Annual Review of Plant Biology,2003,54:547-574
87. Leblanc O, Pointe C, Hernandez M. Cell cycle progression during endosperm development in Zea mays depends on parental dosage effects. The Plant Journal, 2002,32:1057-1066
88.Leroy O, Hennig L, Breuninger H, Laux T, Kohler C.Polycomb group proteins function in the female gametophyte to determine seed development in plants. Development,2007,134:3639-3648
89. Lee M G, Villa R, Trojer P, Norman J, Yan K P, Reinberg D, Croce L D and Shiekhattar R. Demethylation of H3K27 regulates polycomb recruitment and H2A ubiquitination. Science,2007b,318:447-450
90.Letunic I, Copley R R, Pils B, Pinkert S, Schultz J and Bork P SMART 5:domains in the context of genomes and networks. Nucleic Acids Res.,2006,34:D257-260
91.Lewis E B A gene complex controlling segmentation in Drosopihla. Nature,1978, 276(7):565-570
92.Li K, Yang J, Liu J, Du X, Wei C, Su W, He G,Zhang Q, Hong F, Qian X. Cloning, Characterization And Tissue-specific Expression Of A Cdna Encoding A Novel Embryonic Flower 2 Gene (Osemf2) In Oryza Sativa Mitochondrial DNA,2006,17: 74-78
93.Loidl P. A plant dialect of the histone language. Trends Plant Sci,2004,9:84-90
94. Lu C, Meyers B C and Green P J Construction of small RNA cDNA libraries for deep sequencing. Methods,2007,43:110-117
95.Lohe A, Chaudhury A. Demeter:on seeds and goddesses. The Plant Cell,2002,14: 2981-2983
96.Lopes M A, Larkins B A. Endosperm origin, development, and function. Plant Cell, 1993,5(10):1383-1399
97.Luo M, Bilodeau P, Dennis E S,Peacock W J, Chaudhury A. Expression and parent-of-origin effects for EMF2, MEA,and FIE in the endosperm and embryo of developing Arabidopsis seeds. Proc Natl Acad Sci USA,2000,97(19):10637-10642
98.Liu F, Quesada V, Crevillen P, Baurle I, Swiezewski S and Dean C The Arabidopsis RNA-binding protein FCA requires a lysine-specific demethylase 1 homolog to downregulate FLC.Mol Cell,2007a,28:398-407
99.Luo M, Bilodeau P, Koltunow A, Dennis E S,Peacock W J, Chaudhury A M. Genes controlling fertilization-independent seed development in Arabidopsis thaliana. Proc Natl Acad Sci USA,1999,96:296-301
100.Malagnac F, Bartee L and Bender J An Arabidopsis SET domain protein required for maintenance but not establishment of DNA methylation. Embo J,2002, 21:6842-6852
101.Mathieu O, Probst A V and Paszkowski J Distinct regulation of histone H3 methylation at lysines 27 and 9 by CpG methylation in Arabidopsis.Embo J,2005, 24:2783-2791
102.Makarevich G, Leroy O, Akinci U, Schubert D, Clarenz O, Goodrich J, Grossniklaus U, Kohler C.Different Polycomb group complexes regulate common target genes in Arabidopsis. EMBO report,2006,7 (9):947-952
103.Makarevich G, Villar C B R, Erilova A, Kohler C.Mechanism of PHERES1 imprinting in Arabidopsis. Journal of Cell Science,2008,121:906-912
104.Moazed D.Enzymatic activities of Sir2 and chromatin silencing. Curr Opin Cell Biol, 2001,13:232-238
105.Morales-Ruiz T. Ortega-Galisteo A P, Ponferrada-Marin M I, Martinez-Macias M I, Ariza R R and Roldan-Arjona T DEMETER and REPRESSOR OF SILENCING 1 encode 5-methylcytosine DNA glycosylases. Proc Natl Acad Sci U S A,2006, 103:6853-6858
106.Mosquna A, Katz A, Shochat S,Grafi G, Ohad N. Interaction of FIE, a Polycomb protein, with pRb:a possible mechanism regulating endosperm development. Mol Gen Genomics,2004,271:651-657
107.Muller J, Kassis J A. Polycomb response elements and targeting of Polycomb group proteins in Drosophila. Current Opinion in Genetics & Development,2006,16: 476-484
108.Naumann K, Fischer A, Hofmann I, Krauss V, Phalke S,Irmler K, Hause G, Aurich A C, Dorn R, Jenuwein T and Reuter G Pivotal role of AtSUVH2 in heterochromatic histone methylation and gene silencing in Arabidopsis. Embo J,2005,24:1418-1429
109.Niu L, Lu F, Pei Y, Liu C and Cao X Regulation of flowering time by the protein arginine methyltransferase AtPRMT10. EMBO Rep,2007,8:1190-1195
110.Ng D W-K, Wang T, Chandrasekharan M B,Aramayo R, Kertbundit S,Hall T C. Plant SET domain-containing proteins:Structure, function and regulation. Biochimica et Biophysica Acta,2007,1769:316-329
111.Nicole Schonrock R B, Olivier Leroy, Lorenzo Borghi, Claudia Kohler, Wilhelm Gruissem and Lars Hennig. Polycomb-group proteins repressthe floral activator AGL19 in the FLC-independent vernalization pathway. Genetics & Development, 2006,20:1667-1678
112.Ohad N, Margossian L, Hsu D-C,Williams C, Repetti P, Fischer R L. A mutation that allows endosperm development without fertilization. Proc Natl Acad Sci USA,1996, 93:5319-5324
113.Ohad N, Ramin Yadegari L M, Mike Hannon. Mutations in FIE, a WD polycomb group gene, allow endosperm development without fertilization. The Plant Cell,1999, 11(3):407-415
114.Onyango P, Miller W, Lehoczky J, Leung C T, Birren B, Wheelan S,Dewar K, Feinberg A P. Sequence and comparative analysis of the mouse 1-Megabase region orthologous to the human 11p15 imprinted domain. Genome Res,2000,10(11): 1697-1710
115.Pagnussat G C, Yu H J, Ngo Q A, Rajani S,Mayalagu S, Johnson C S,Capron A, Xie L F, Ye D and Sundaresan V Genetic and molecular identification of genes required for female gametophyte development and function in Arabidopsis. Development, 2005,132:603-614
116.Pandey R, Muller A, Napoli C A, Selinger D A, Pikaard C S,Richards E J, Bender J, Mount D W and Jorgensen R A Analysis of histone acetyltransferase and histone deacetylase families of Arabidopsis thaliana suggests functional diversification of chromatin modification among multicellular eukaryotes. Nucleic Acids Res,2002, 30:5036-5055
117.Philippe J,Calzada V, Thomas J, Spillane C, Coluccio A, Hoeppner M A, Grossniklaus U. Maintenance of genomic imprinting at the Arabidopsis medea locus requires zygotic DDM1 activity. Genetics & Development,1999,13:2971-2982
118.Portereiko M F, Lloyd A, Steffen J G, Punwani J A, Otsuga D, Drews G N. AGL80 Is required for central cell and endosperm development in Arabidopsis. The Plant Cell, 2006,18:1862
119.Reiser L, Fische R L. The ovule and the embryo sac. The Plant Cell,1993,5: 1291-1301
120.Reyes J C, Grossniklaus U. Diverse functions of Polycomb group proteins during plant development. Seminars in Cell & Developmental Biology,2003,14:77-84
121.Rossi V, Varotto S,Locatelli S, Lanzanova C, Lauria M, Zanotti E, Hartings H, Motto M. The maize WD-repeat gene ZmRbApl encodes a member of the MSI/RbAp sub-family and is differentially expressed during endosperm development. Molecular Genetics and Genomics,2001,256(4):576-584
122.Rossi V, Locatelli S,Varotto S,Donn G, Pirona R, Henderson D A, Hartings H.and Motto M.Maize histone deacetylase hda101 is involved in plant development, gene transcription, and sequence-specific modulation of histone modification of genes and repeats.Plant Cell,2007,19:1145-1162
123.Rubin H. Etymology of epigenetics. Science,2001,294:2477-2478
124.Rubio V, Shen Y, Saijo Y, Liu Y, Gusmaroli G, Dinesh-Kumar S P and Deng X W An alternative tandem affinity purification strategy applied to Arabidopsis protein complex isolation. Plant J,2005,41:767-778
125.Saleh A, Al-Abdallat A, Ndamukong I, Venegas R A, Avramova Z. The Arabidopsis homologs of trithorax (ATX1)and enhancer of zeste (CLF) establish'bivalent chromatin marks'at the silent AGAMOUS locus. Nucleic Acids Research,2007,35 (18):6290-6296
126.Schwartz Y B, Pirrotta V;Polycomb silencing mechanisms and the management of genomic programmes. Nature genetics,2007,8:9-22
127.Scott R J, Spielman M. Deeper into the maize:new insights into genomic imprinting in plants. BioEssays,2006,28:1167-1171
128.Scebba F, De Bastiani M, Bernacchia G, Andreucci A, Galli A and Pitto L PRMT11:a new Arabidopsis MBD7 protein partner with arginine methyltransferase activity. Plant J,2007,52:210-222
129.Schmitz R J, Sung S and Amasino R M Histone arginine methylation is required for vernalization-induced epigenetic silencing of FLC in winter-annual Arabidopsis thaliana. Proc Natl Acad Sci U S A,2008,105:411-416
130.Spillane C, MacDougall C, Stock C, Kohler C, Calzada J P V, Nunes S M, Grossniklaus U, Goodrich J. Interaction of the Arabidopsis Polycomb group proteins FIE and MEA mediates their common phenotypes.Current Biology,2000,(10): 1535-1538
131.Spillane C, Schmid K J, Duprat S L, Pien S, Escobar-Restrepo J-M. Positive darwinian selection at the imprinted MEDEA locus in plants.Nature,2007,448(19): 349-354
132.Springer N M, Napoli C A, Selinger D A, Pandey R, Cone K C, Chandler V L, Kaeppler H F and Kaeppler S M Comparative analysis of SET domain proteins in maize and Arabidopsis reveals multiple duplications preceding the divergence of monocots and dicots.Plant Physiol,2003,132:907-925
133.Sridha S and Wu K Identification of AtHD2C as a novel regulator of abscisic acid responses in Arabidopsis. Plant J,2006,46:124-133
134.Sridhar V V, Kapoor A, Zhang K, Zhu J, Zhou T, Hasegawa P M, Bressan R A and Zhu J K Control of DNA methylation and heterochromatic silencing by histone H2B deubiquitination. Nature,2007,447:735-738
135.Strickert M, Sreenivasulu N, Usadel B,Seiffert U. Correlation-maximizing surrogate gene space for visual mining of gene expression patterns in developing barley endosperm tissue. BMC Bioinformatics,2007,8:165-175
136.Sung Z R, Belachew A, Shunong B, Bertrand-Garcia R. EMF, an Arabidopsis gene required for vegetative shoot development Science,1992,258 (5088):1645-1647
137.Theien G Secret life of genes. Nature,2002,415:741
138.Tsukada Y, Fang J, Erdjument-Bromage H, Warren M E, Borchers C H, Tempst P and Zhang Y Histone demethylation by a family of JmjC domain-containing proteins. Nature,2006,439:811-816
139.Turck F, Roudier F, Farrona S,Martin-Magniette M L, Guillaume E, Buisine N, Gagnot S, Martienssen R A, Coupland G and Colot V Arabidopsis TFL2/LHP1 specifically associates with genes marked by trimethylation of histone H3 lysine 27. PLoS Genet,2007,3:e86
140.Turner B M Cellular memory and the histone code. Cell,2002,111:285-291
141.Vinkenoog R, Scott R J Autonomous endosperm development in flowering plants: how to overcome the imprinting problem? Sex Plant Reprod,2001,14:189-194
142.Vinkenoog R, Spielman M, Adams S,Fischer R L, Dickinson H G, Scott R J. Hypomethylation promotes autonomous endosperm development and rescues postfertilization lethality in fie Mutants. The Plant Cell,2000b,12:2271-2282
143.Wang D F, Tyson M D, Jackson S S, Yadegari R. Partially redundant functions of two SET-domain polycomb-group proteins in controlling initiation of seed development in Arabidopsis. Proc Natl Acad Sci USA,2006,103:13244-13249
144.Wang X, Zhang Y, Ma Q, Zhang Z, Xue Y, Bao S and Chong K SKB1-mediated symmetric dimethylation of histone H4R3 controls flowering time in Arabidopsis. Embo J,2007,26:1934-1941
145.Warthmann N, Chen H, Ossowski S, Weigel D and Herve P Highly specific gene silencing by artificial miRNAs in rice.PLoS ONE,2008,3:e1829
146.Wang X X, Ma L G. Polycomb-group (Pc-G) proteins control seed development in Arabidopsis thaliana L. Journal of Integrative Plant Biology,2007,47(1):52-59
147.Wood C C, Robertson M, Tanner G, Peacock W J, Dennis E S,Helliwell C A. The Arabidopsis thaliana vernalization response requires a polycomb-like protein complex that also includes VERNALIZATION INSENSITIVE 3.Proc Natl Acad Sci USA,2006,103:14631-14636
148.Yadegari R, Drews G N. Female gametophyte development. The Plant Cell,2004,16: S133-S141
149.Yadegari R, Kinoshita T, Lotan O, Cohen G, Katz A, Choi Y, Katz A, Nakashima K, Harada J J, Goldberg R B, Fischer R L, Ohad N, Mutations in the FIE and MEA genes that encode interacting polycomb proteins cause parent-of-origin effects on seed development by distinct mechanisms The Plant Cell,2000,12:2367-2382
150.Yoshida N, Yanai Y, Chen L, Kato Y, Hiratsuka J, Miwa T, Sung Z R, Takahashi S. EMBRYONIC FLOWER2, a novel polycomb group protein homolog, mediates shoot development and flowering in Arabidopsis. The Plant Cell,2001,13: 2471-2481
151.Yu X, Li L, Li L, Guo M, Chory J and Yin Y;Modulation of brassinosteroid-regulated gene expression by Jumonji domain-containing proteins ELF6 and REF6 in Arabidopsis. Proc Natl Acad Sci U S A,2008,105:7618-7623
152.Yu Y, Dong A and Shen W H. Molecular characterization of the tobacco SET domain protein NtSET1 unravels its role in histone methylation, chromatin binding, and segregation. Plant J,2004,40:699-711
153.Zhang D, Yoon H G and Wong J. JMJD2A is a novel N-CoR-interacting protein and is involved in repression of the human transcription factor achaete scute-like homologue 2(ASCL2/Hash2).Mol Cell Biol,2005,25:6404-6414
154.Zhang J, Guo D, Chang Y, You C, Li X, Dai X, Weng Q, Zhang J, Chen G, Li X, Liu H, Han B,Zhang Q, Wu C.Non-random distribution of T-DNA insertions at various levels of the genome hierarchy as revealed by analyzing 13 804 T-DNA flanking sequences from an enhancer-trap mutant library. The Plant Journal,2007,49: 947-959
155.Zhou C, Zhang L, Duan J, Miki B.and Wu K. HISTONE DEACETYLASE19 is involved in jasmonic acid and ethylene signaling of pathogen response in Arabidopsis.Plant Cell,2005,17:1196-1204
156.Zilberman D and Henikoff S Genome-wide analysis of DNA methylation patterns. Development,2007,134:3959-3965
2.张文伟,曹少先,江玲,朱速松,万建民.基因组印记与种子发育.遗传,2005b,27(4)：665-670
3.Agger K, Cloos P A, Christensen J, Pasini D, Rose S, Rappsilber J, Issaeva I, Canaani E, Salcini A E and Helin K UTX and JMJD3 are histone H3K27 demethylases involved in HOX gene regulation and development. Nature,2007,449:731-734
4.Allis C D, Berger S L, Cote J, Dent S,Jenuwien T, Kouzarides T, Pillus L, Reinberg D, Shi Y, Shiekhattar R, Shilatifard A, Workman J and Zhang Y New nomenclature for chromatin-modifying enzymes.Cell,2007,131:633-636
5.Alvarez-Venegas R, Pien S, Sadder M, Witmer X, Grossniklaus U and Avramova Z ATX-1,an Arabidopsis homolog of trithorax, activates flower homeotic genes.Curr Biol,2003,13:627-637
6.Arnaud P, Feil R MEDEA takes control of its own imprinting Cell,2006,124: 468-470
7.Aubert D, Chen L, Moon Y-H, Martin D, Castle L A, Yang C-H, Sung Z R. EMF1,a novel protein involved in the control of shoot architecture and flowering in Arabidopsis. The Plant Cell,2001,13:1865-1875
8.Autran D, Mamani W H, Calzada J P V.Genomic imprinting in plants:the epigenetic version of an Oedipus complex. Current Opinion in Plant Biology,2005,8:19-25
9. Baroux C, Gagliardini V, Page D R, Grossniklaus U. Dynamic regulatory interactions of polycomb group genes:MEDEA autoregulation is required for imprinted gene expression in Arabidopsis. Genetics & Development,2006,20:1081-1086
10. Baroux C, Pien S, Grossniklaus U. Chromatin modification and remodeling during early seed development. Genetics & Development,2007,17:473-479
11.Baroux C, Spillane C, Grossniklaus U. Genomic imprinting during seed development. Adv Genet,2002,46:165-214
12.Baubec T, Scheid O M. Medea in full self-control. TRENDS in Plant Science,2006, 11:469-471
13.Benhamed M, Bertrand C,Servet C and Zhou D X Arabidopsis GCN5,HD1,and TAF1/HAF2 interact to regulate histone acetylation required for light-responsive gene expression. Plant Cell,2006,18:2893-2903
14. Berndsen C E and Denu J M Catalysis and substrate selection by histone/protein lysine acetyltransferases.Curr Opin Struct Biol,2008,18:682-689
15.Berger F Endosperm:the crossroad of seed development. Current Opinion in Plant Biology,2003,6:42-50
16.Berger F, Grini P E, Schnittger A. Endosperm:an integrator of seed growth and development. Current Opinion in Plant Biology,2006,9:664-670
17.Brink R A, Cooper D C.The endosperm in seed development The Botanical Review, 1947,13(9):479-541
18.Cao R, Wang L, Wang H, Xia L, Erdjument-Bromage H, Tempst P, Jones R S and Zhang Y Role of histone H3 lysine 27 methylation in Polycomb-group silencing. Science,2002,298:1039-1043
19.Cao Y, Dai Y, Cui S and Ma L Histone H2B monoubiquitination in the chromatin of FLOWERING LOCUS C regulates flowering time in Arabidopsis.Plant Cell,2008, 20:2586-2602
20.Cartagena J A, Matsunaga S,Seki M, Kurihara D, Yokoyama M, Shinozaki K, Fujimoto S, Azumi Y, Uchiyama S and Fukui K The Arabidopsis SDG4 contributes to the regulation of pollen tube growth by methylation of histone H3 lysines 4 and 36 in mature pollen Dev Biol,2008,315:355-368
21.Chanvivattana Y, Bishopp A, Schubert D, Stock C, Moon Y-H, Goodrich Z R S a J Interaction of Polycomb-group proteins controlling flowering in Arabidopsis. Development,2004,131:5263-5276
22. Chaudhury A M, Ming L, Miller C, Craig S, Dennis E, Peacock W J. Fertilization-independent seed development in Arabidopsis thaliana. Plant Biology, 1997,(94):4223-4228
23.Choi Y, Gehring M, Johnson L, Hannon M, Harada J J, Goldberg R B, Jacobsen S E, Fischer R L. DEMETER, a DNA glycosylase domain protein, is required for endosperm gene imprinting and seed viability in Arabidopsis. Cell,2002,110:33-42
24. Cho Y H, Yoo S D and Sheen J Regulatory functions of nuclear hexokinasel complex in glucose signaling. Cell,2006,127:579-589
25.Choi Y, Gehring M, Johnson L, Hannon M, Harada J J, Goldberg R B, Jacobsen S E and Fischer R. L.DEMETER, a DNA glycosylase domain protein, is required for endosperm gene imprinting and seed viability in Arabidopsis. Cell,2002,110:33-42
26.Collinge M A, Spillane C,Kohler C, Gheyselinck J, Grossniklaus U Genetic interaction of an origin recognition complex subunit and the polycomb group gene MEDEA during seed development. The Plant Cell,2004,16:1035-1046
27. Costa L M, Gutierrez-Marcos J F, Dickinson H G. More than a yolk:the short life and complex times of the plant endosperm. TRENDS in Plant Science,2004,9(10): 507-514
28.Curtis M D, Grossniklaus U Molecular control of autonomous embryo and endosperm development. Sex Plant Reprod,2008,21:79-88
29.Danilevskaya O N, Hermon P, Hantke S,Muszynski M G, Kollipara K, Ananiev E V. Duplicated fie genes in maize:expression pattern and imprinting suggest distinct functions. The Plant Cell,2003,15:425-438
30.De Crombrugghe B,Busby S, Buc H. Cyclic AMP Receptor Protein:Role in Transcription Activation.Science,1984,224:831-838
31.De-Pater S,Greco V, Pham K, Memelink J, Kijne J.Characterization of a zinc-dependent transcriptional activator from Arabidopsis. Nucleic Acids Res,1996, 24:4624-4631
32.Degenhardt J, Tobin E M. A DNA binding activity for one of two closely defined phytochrome regulatory elements in an Lhcb promoter is more abundant in etiolated than in green plants. Plant Cell,1996,8:31-41
33.Dilkes B P, Comai L. A differential dosage hypothesis for parental effects in seed development. The Plant Cell,2004,16:3174-3180
34. Dumas C, Rogowsky P.Fertilization and early seed formation. Comptes Rendus Biologies,2008,331:715-725
35.Ding Y,Wang X, Su L, Zhai J, Cao S,Zhang D, Liu C, Bi Y, Qian Q, Cheng Z, Chu C and Cao X SDG714, a histone H3K9 methyltransferase, is involved in Tos17 DNA methylation and transposition in rice. Plant Cell,2007,19:9-22
36.Dong G, Ma D P and Li J The histone methyltransferase SDG8 regulates shoot branching in Arabidopsis. Biochem Biophys Res Commun,2008,373:659-664
37.Earley K, Lawrence R J, Pontes O, Reuther R, Enciso A J, Silva M, Neves N, Gross M, Viegas W and Pikaard C S Erasure of histone acetylation by Arabidopsis HDA6 mediates large-scale gene silencing in nucleolar dominance. Genes Dev,2006, 20:1283-1293
38.Elmen J, Lindow M, Schutz S,Lawrence M, Petri A, Obad S,Lindholm M, Hedtjarn M, Hansen H F, Berger U, Gullans S,Kearney P, Sarnow P, Straarup E M, Kauppinen S.LNA-mediated microRNA silencing in non-human primates.Nature, 2008,452:896-899
39.Feng Q, Wang H, Ng H H, Erdjument-Bromage H, Tempst P, Struhl K and Zhang Y Methylation of H3-lysine 79 is mediated by a new family of HMTases without a SET domain Curr Biol,2002,12:1052-1058
40.Feil R, Berger F Convergent evolution of genomic imprinting in plants and mammals. TRENDS in Genetics,2007,23 (4):192-199
41.Finnegan E J, Dennis E S.Isolation and identification by sequence homology of a putative cytosine methyltransferase from Arabidopsis thaliana. Nucleic Acids Res, 1993,21(10):2383-2388
42.Friedman W E The evolution of double fertilization and endsperm:an'historical' perspective. Sex Plant Reprod,1998,11:6-16
43.Garcia D, Saingery V, Chambrier P, Mayer U, Jurgens G, Berger F. Arabidopsis haiku mutants reveal new controls of seed size by endosperm. Plant Physiology,2003, 131:1661-1670
44.Gary S.A gene product required for correct initiation of segmental determination in Drosophila. Nature,1981,293(3):36-41
45.Gehring M, Choi Y, Fischer R L.Imprinting and seed development. The Plant Cell, 2004,16:S203-S213
46.Gehring M, Huh J H, Hsieh T F, Penterman J, Choi Y, Harada J J, Goldberg R B and Fischer R L DEMETER DNA glycosylase establishes MEDEA polycomb gene self-imprinting by allele-specific demethylation. Cell,2006,124:495-506
47.Gendler K, Paulsen T and Napoli C ChromDB:the chromatin database. Nucleic Acids Res,2008,36:D298-302
48.Gong Z, Morales-Ruiz T, Ariza R R, Roldan-Arjona T, David L and Zhu J K ROS1,a repressor of transcriptional gene silencing in Arabidopsis, encodes a DNA glycosylase/lyase.Cell,2002,111:803-814
49.Gonzalez D, Bowen A J, Carroll T S and Conlan R S.The transcription corepressor LEUNIG interacts with the histone deacetylase HDA19 and mediator components MED14 (SWP) and CDK8 (HEN3) to repress transcription. Mol Cell Biol,2007, 27:5306-5315
50. Goodrich J, Puangsomlee P, Martin M, Long D, Meyerowitz E M, Coupland G. A Polycomb-group gene regulates homeotic gene expression in Arabidopsis. Nature, 1997,386:44-51
51.Grimanelli D, Perotti E, Ramirez J, Leblanc O.Timing of the maternal-to-zygotic transition during early seed development in maize. The Plant Cell,2005,17: 1061-1072
52.Grossniklaus U, Philippe J, Calzada V, Hoeppner M A, Gagliano W B.Maternal control of embryogenesis by MEDEA,a polycomb group gene in Arabidopsis. Science,1998a, (280):446-450
53.Grozinger C M, Chao E D, Blackwell H E, Moazed D and Schreiber S L Identification of a class of small molecule inhibitors of the sirtuin family of NAD-dependent deacetylases by phenotypic screening. J Biol Chem,2001, 276:38837-38843
54. Guitton A-E, Page D R, Chambrier P, Lionnet C, Faure J-E, Grossniklaus U, Berger F. Identification of new members of Fertilisation Independent Seed Polycomb Group pathway involved in the control of seed development in Arabidopsis thaliana. Development,2004,131:2971-2981
55.Gutierrez-Marcos J F,Costa L M,Pra M D,Scholten S,Kranz E,Perez P,Dickinson H G. Epigenetic asymmetry of imprinted genes in plant gametes.Nature genetics, 2006,38(8):876-878
56.Hennig L, Taranto P, Walser M, Schonrock N, Gruissem W. Arabidopsis MSI1 is required for epigenetic maintenance of reproductive development. Development, 2003,130:2555-2565
57.Hermon P, Srilunchang K, Zou J, Dresselhaus T, Danilevskaya O N.Activation of the imprinted Polycomb Group Fiel gene in maize endosperm requires demethylation of the maternal allele. Plant Mol Biol,2007,64:387-395
58. Hsieh T-F, Hakim O, Ohad N, Fischer R L. From flour to flower:how Polycomb group proteins influence multiple aspects of plant development. TRENDS in Plant Science, 2003,8(9):439-445
59.Huang L, Sun Q, Qin F, Li C, Zhao Y and Zhou D X Down-regulation of a SILENT INFORMATION REGULATOR2-related histone deacetylase gene, OsSRTl,induces DNA fragmentation and cell death in rice. Plant Physiol,2007,144:1508-1519
60. Husbands A, Bell E M, Shuai B,Smith H M and Springer P S LATERAL ORGAN BOUNDARIES defines a new family of DNA-binding transcription factors and can interact with specific bHLH proteins. Nucleic Acids Res,2007,35:6663-6671
61.Huh J H, Bauer M J, Hsieh T-F, Fischer R. Endosperm gene imprinting and seed development. Current Opinion in Genetics & Development,2007,17:480-485
62.Ingouff M, Haseloff J,Berger F. Polycomb group genes control developmental timing of endosperm. The Plant Journal,2005,42:663-674
63.Jeong D-H, An S, Kang H-G, Moon S, Han J-J, Park S, Lee H S, An K, An G. T-DNA insertional mutagenesis for activation tagging in rice.Plant Physiology,2002,130: 1636-1644
64.Jeong D-H, An S,Park S, Kang H-G, Park G-G, Kim S-R, Sim J, Kim Y-O, Kim M-K, Kim S-R, Kim J, Shin M, Jung M, An G. Generation of a flanking sequence-tag database for activation-tagging lines in japonica rice. The Plant Journal,2006,45: 123-132
65.Jackson J P, Lindroth A M, Cao X and Jacobsen S E Control of CpNpG DNA methylation by the KRYPTONITE histone H3 methyltransferase. Nature,2002, 416:556-560
66.Jackson M D, Schmidt M T, Oppenheimer N J and Denu J M Mechanism of nicotinamide inhibition and transglycosidation by Sir2 histone/protein deacetylases.J Biol Chem,2003,278:50985-50998
67.Jiang D, Wang Y, Wang Y, He Y. Repression of FLOWERING LOCUS C and FLOWERING LOCUS T by the Arabidopsis Polycomb Repressive Complex 2 Components. PLoS ONE,2008,3(10):1-12
68.Jullien P E, Katz A, Oliva M, Ohad N, Berger F. Polycomb group complexes self-regulate imprinting of the polycomb group gene MEDEA in Arabidopsis. Current Biology,2006a,16:486-492
69.Jenuwein T, Laible G, Dorn R and Reuter G SET domain proteins modulate chromatin domains in eu-and heterochromatin. Cell Mol Life Sci,1998,54:80-93
70.Jeon J S,Lee S,Jung K H, Jun S H, Jeong D H, Lee J, Kim C, Jang S,Yang K, Nam J, An K, Han M J, Sung R J, Choi H S, Yu J H, Choi J H, Cho S Y, Cha S S, Kim S I and An G T-DNA insertional mutagenesis for functional genomics in rice.Plant J, 2000,22:561-570
71. Katz A, Oliva M, Mosquna A, Hakim O, Ohad N. FIE and CURLY LEAF polycomb proteins interact in the regulation of homeobox gene expression during sporophyte development. The Plant Journal,2004,37:707
72.Kim K N, Guiltinan M J. Identification of cis-acting elements important for expression of the starch-branching enzyme I gene in maize endosperm. Plant Physiol., 1999,121:225-236
73.Kim K C,Lai Z, Fan B and Chen Z Arabidopsis WRKY38 and WRKY62 transcription factors interact with histone deacetylase 19 in basal defense. Plant Cell, 2008,20:2357-2371
74.Klose R J, Kallin E M and Zhang Y JmjC-domain-containing proteins and histone demethylation. Nat Rev Genet,2006a,7:715-727
75.Kinoshita T, Harada J J, Goldberg R B, Fischer R L. Polycomb repression of flowering during early plant development. Proc Natl Acad Sci USA,2001,98: 14156-14161
76.Kinoshita T, Yadegari R, Harada J J, Goldberg R B, Fischer R L. Imprinting of the MEDEA polycomb gene in the Arabidopsis endosperm. The Plant Cell,1999,11: 1945-1952
77.Komiya R, Ikegami A, Tamaki S, Yokoi S and Shimamoto K Hd3a and RFT1 are essential for flowering in rice. Development,2008,135:767-774
78.Kouzarides T. Chromatin modifications and their function. Cell,2007,128:693-705
79.Kiyosue T, Ohad N, Yadegar R, Hannon M, Dinneny J, Wells D, Katz L M A, Harada J J, Goldberg R B,Fischer R L. Control of fertilization-independent endosperm development by the MEDEA polycomb gene in Arabidopsis. Proc Natl Acad Sci USA,1999,96:4186-4191
80.Kladnik A, Chamusco K, Dermastia M, Chourey P.Evidence of programmed cell death in post-phloem transport cells of the maternal pedicel tissue in developing Caryopsis of Maize. Plant Physiology,2004,136:3572-3581
81.Kohler C,Hennig L, Bouveret R, Gheyselinck J, Grossniklaus U, Gruissem W. Arabidopsis MSI1 is a component of the MEA/FIE Polycomb group complex and required for seed development. The EMBO Journal,2003a,22(18):4804-4814
82.Kohler C, Hennig L, Spillane C, Pien S,Gruissem W, Grossniklaus U.The Polycomb-group protein MEDEA regulates seed development by controlling expression of the MADS-box gene PHERES1.Genetics & Development,2003b: 1540-1553
83.Kohler C,Page D R, Gagliarliardini V, Grossniklaus U.The Arabidopsis thaliana MEDEA Polycomb group protein controls expression of PHERES1 by parental imprintng. Nature genetics,2005,37(1):28-30
84.Kollipara K P, Saab I N, Wych R D, Lauer M J, Singletary G W. Expression profiling of reciprocal maize hybrids divergent for cold germination and desiccation tolerance. Plant Physiology,2002,129:974-992
85.Kumar S, Tamura K and Nei M MEGA3:Integrated software for Molecular Evolutionary Genetics Analysis and sequence alignment. Brief Bioinform.,2004, 5:150-163
86. Koltunow A M, Grossniklaus U. Apomixis:A debelopmental perspective.In Annual Review of Plant Biology,2003,54:547-574
87. Leblanc O, Pointe C, Hernandez M. Cell cycle progression during endosperm development in Zea mays depends on parental dosage effects. The Plant Journal, 2002,32:1057-1066
88.Leroy O, Hennig L, Breuninger H, Laux T, Kohler C.Polycomb group proteins function in the female gametophyte to determine seed development in plants. Development,2007,134:3639-3648
89. Lee M G, Villa R, Trojer P, Norman J, Yan K P, Reinberg D, Croce L D and Shiekhattar R. Demethylation of H3K27 regulates polycomb recruitment and H2A ubiquitination. Science,2007b,318:447-450
90.Letunic I, Copley R R, Pils B, Pinkert S, Schultz J and Bork P SMART 5:domains in the context of genomes and networks. Nucleic Acids Res.,2006,34:D257-260
91.Lewis E B A gene complex controlling segmentation in Drosopihla. Nature,1978, 276(7):565-570
92.Li K, Yang J, Liu J, Du X, Wei C, Su W, He G,Zhang Q, Hong F, Qian X. Cloning, Characterization And Tissue-specific Expression Of A Cdna Encoding A Novel Embryonic Flower 2 Gene (Osemf2) In Oryza Sativa Mitochondrial DNA,2006,17: 74-78
93.Loidl P. A plant dialect of the histone language. Trends Plant Sci,2004,9:84-90
94. Lu C, Meyers B C and Green P J Construction of small RNA cDNA libraries for deep sequencing. Methods,2007,43:110-117
95.Lohe A, Chaudhury A. Demeter:on seeds and goddesses. The Plant Cell,2002,14: 2981-2983
96.Lopes M A, Larkins B A. Endosperm origin, development, and function. Plant Cell, 1993,5(10):1383-1399
97.Luo M, Bilodeau P, Dennis E S,Peacock W J, Chaudhury A. Expression and parent-of-origin effects for EMF2, MEA,and FIE in the endosperm and embryo of developing Arabidopsis seeds. Proc Natl Acad Sci USA,2000,97(19):10637-10642
98.Liu F, Quesada V, Crevillen P, Baurle I, Swiezewski S and Dean C The Arabidopsis RNA-binding protein FCA requires a lysine-specific demethylase 1 homolog to downregulate FLC.Mol Cell,2007a,28:398-407
99.Luo M, Bilodeau P, Koltunow A, Dennis E S,Peacock W J, Chaudhury A M. Genes controlling fertilization-independent seed development in Arabidopsis thaliana. Proc Natl Acad Sci USA,1999,96:296-301
100.Malagnac F, Bartee L and Bender J An Arabidopsis SET domain protein required for maintenance but not establishment of DNA methylation. Embo J,2002, 21:6842-6852
101.Mathieu O, Probst A V and Paszkowski J Distinct regulation of histone H3 methylation at lysines 27 and 9 by CpG methylation in Arabidopsis.Embo J,2005, 24:2783-2791
102.Makarevich G, Leroy O, Akinci U, Schubert D, Clarenz O, Goodrich J, Grossniklaus U, Kohler C.Different Polycomb group complexes regulate common target genes in Arabidopsis. EMBO report,2006,7 (9):947-952
103.Makarevich G, Villar C B R, Erilova A, Kohler C.Mechanism of PHERES1 imprinting in Arabidopsis. Journal of Cell Science,2008,121:906-912
104.Moazed D.Enzymatic activities of Sir2 and chromatin silencing. Curr Opin Cell Biol, 2001,13:232-238
105.Morales-Ruiz T. Ortega-Galisteo A P, Ponferrada-Marin M I, Martinez-Macias M I, Ariza R R and Roldan-Arjona T DEMETER and REPRESSOR OF SILENCING 1 encode 5-methylcytosine DNA glycosylases. Proc Natl Acad Sci U S A,2006, 103:6853-6858
106.Mosquna A, Katz A, Shochat S,Grafi G, Ohad N. Interaction of FIE, a Polycomb protein, with pRb:a possible mechanism regulating endosperm development. Mol Gen Genomics,2004,271:651-657
107.Muller J, Kassis J A. Polycomb response elements and targeting of Polycomb group proteins in Drosophila. Current Opinion in Genetics & Development,2006,16: 476-484
108.Naumann K, Fischer A, Hofmann I, Krauss V, Phalke S,Irmler K, Hause G, Aurich A C, Dorn R, Jenuwein T and Reuter G Pivotal role of AtSUVH2 in heterochromatic histone methylation and gene silencing in Arabidopsis. Embo J,2005,24:1418-1429
109.Niu L, Lu F, Pei Y, Liu C and Cao X Regulation of flowering time by the protein arginine methyltransferase AtPRMT10. EMBO Rep,2007,8:1190-1195
110.Ng D W-K, Wang T, Chandrasekharan M B,Aramayo R, Kertbundit S,Hall T C. Plant SET domain-containing proteins:Structure, function and regulation. Biochimica et Biophysica Acta,2007,1769:316-329
111.Nicole Schonrock R B, Olivier Leroy, Lorenzo Borghi, Claudia Kohler, Wilhelm Gruissem and Lars Hennig. Polycomb-group proteins repressthe floral activator AGL19 in the FLC-independent vernalization pathway. Genetics & Development, 2006,20:1667-1678
112.Ohad N, Margossian L, Hsu D-C,Williams C, Repetti P, Fischer R L. A mutation that allows endosperm development without fertilization. Proc Natl Acad Sci USA,1996, 93:5319-5324
113.Ohad N, Ramin Yadegari L M, Mike Hannon. Mutations in FIE, a WD polycomb group gene, allow endosperm development without fertilization. The Plant Cell,1999, 11(3):407-415
114.Onyango P, Miller W, Lehoczky J, Leung C T, Birren B, Wheelan S,Dewar K, Feinberg A P. Sequence and comparative analysis of the mouse 1-Megabase region orthologous to the human 11p15 imprinted domain. Genome Res,2000,10(11): 1697-1710
115.Pagnussat G C, Yu H J, Ngo Q A, Rajani S,Mayalagu S, Johnson C S,Capron A, Xie L F, Ye D and Sundaresan V Genetic and molecular identification of genes required for female gametophyte development and function in Arabidopsis. Development, 2005,132:603-614
116.Pandey R, Muller A, Napoli C A, Selinger D A, Pikaard C S,Richards E J, Bender J, Mount D W and Jorgensen R A Analysis of histone acetyltransferase and histone deacetylase families of Arabidopsis thaliana suggests functional diversification of chromatin modification among multicellular eukaryotes. Nucleic Acids Res,2002, 30:5036-5055
117.Philippe J,Calzada V, Thomas J, Spillane C, Coluccio A, Hoeppner M A, Grossniklaus U. Maintenance of genomic imprinting at the Arabidopsis medea locus requires zygotic DDM1 activity. Genetics & Development,1999,13:2971-2982
118.Portereiko M F, Lloyd A, Steffen J G, Punwani J A, Otsuga D, Drews G N. AGL80 Is required for central cell and endosperm development in Arabidopsis. The Plant Cell, 2006,18:1862
119.Reiser L, Fische R L. The ovule and the embryo sac. The Plant Cell,1993,5: 1291-1301
120.Reyes J C, Grossniklaus U. Diverse functions of Polycomb group proteins during plant development. Seminars in Cell & Developmental Biology,2003,14:77-84
121.Rossi V, Varotto S,Locatelli S, Lanzanova C, Lauria M, Zanotti E, Hartings H, Motto M. The maize WD-repeat gene ZmRbApl encodes a member of the MSI/RbAp sub-family and is differentially expressed during endosperm development. Molecular Genetics and Genomics,2001,256(4):576-584
122.Rossi V, Locatelli S,Varotto S,Donn G, Pirona R, Henderson D A, Hartings H.and Motto M.Maize histone deacetylase hda101 is involved in plant development, gene transcription, and sequence-specific modulation of histone modification of genes and repeats.Plant Cell,2007,19:1145-1162
123.Rubin H. Etymology of epigenetics. Science,2001,294:2477-2478
124.Rubio V, Shen Y, Saijo Y, Liu Y, Gusmaroli G, Dinesh-Kumar S P and Deng X W An alternative tandem affinity purification strategy applied to Arabidopsis protein complex isolation. Plant J,2005,41:767-778
125.Saleh A, Al-Abdallat A, Ndamukong I, Venegas R A, Avramova Z. The Arabidopsis homologs of trithorax (ATX1)and enhancer of zeste (CLF) establish'bivalent chromatin marks'at the silent AGAMOUS locus. Nucleic Acids Research,2007,35 (18):6290-6296
126.Schwartz Y B, Pirrotta V;Polycomb silencing mechanisms and the management of genomic programmes. Nature genetics,2007,8:9-22
127.Scott R J, Spielman M. Deeper into the maize:new insights into genomic imprinting in plants. BioEssays,2006,28:1167-1171
128.Scebba F, De Bastiani M, Bernacchia G, Andreucci A, Galli A and Pitto L PRMT11:a new Arabidopsis MBD7 protein partner with arginine methyltransferase activity. Plant J,2007,52:210-222
129.Schmitz R J, Sung S and Amasino R M Histone arginine methylation is required for vernalization-induced epigenetic silencing of FLC in winter-annual Arabidopsis thaliana. Proc Natl Acad Sci U S A,2008,105:411-416
130.Spillane C, MacDougall C, Stock C, Kohler C, Calzada J P V, Nunes S M, Grossniklaus U, Goodrich J. Interaction of the Arabidopsis Polycomb group proteins FIE and MEA mediates their common phenotypes.Current Biology,2000,(10): 1535-1538
131.Spillane C, Schmid K J, Duprat S L, Pien S, Escobar-Restrepo J-M. Positive darwinian selection at the imprinted MEDEA locus in plants.Nature,2007,448(19): 349-354
132.Springer N M, Napoli C A, Selinger D A, Pandey R, Cone K C, Chandler V L, Kaeppler H F and Kaeppler S M Comparative analysis of SET domain proteins in maize and Arabidopsis reveals multiple duplications preceding the divergence of monocots and dicots.Plant Physiol,2003,132:907-925
133.Sridha S and Wu K Identification of AtHD2C as a novel regulator of abscisic acid responses in Arabidopsis. Plant J,2006,46:124-133
134.Sridhar V V, Kapoor A, Zhang K, Zhu J, Zhou T, Hasegawa P M, Bressan R A and Zhu J K Control of DNA methylation and heterochromatic silencing by histone H2B deubiquitination. Nature,2007,447:735-738
135.Strickert M, Sreenivasulu N, Usadel B,Seiffert U. Correlation-maximizing surrogate gene space for visual mining of gene expression patterns in developing barley endosperm tissue. BMC Bioinformatics,2007,8:165-175
136.Sung Z R, Belachew A, Shunong B, Bertrand-Garcia R. EMF, an Arabidopsis gene required for vegetative shoot development Science,1992,258 (5088):1645-1647
137.Theien G Secret life of genes. Nature,2002,415:741
138.Tsukada Y, Fang J, Erdjument-Bromage H, Warren M E, Borchers C H, Tempst P and Zhang Y Histone demethylation by a family of JmjC domain-containing proteins. Nature,2006,439:811-816
139.Turck F, Roudier F, Farrona S,Martin-Magniette M L, Guillaume E, Buisine N, Gagnot S, Martienssen R A, Coupland G and Colot V Arabidopsis TFL2/LHP1 specifically associates with genes marked by trimethylation of histone H3 lysine 27. PLoS Genet,2007,3:e86
140.Turner B M Cellular memory and the histone code. Cell,2002,111:285-291
141.Vinkenoog R, Scott R J Autonomous endosperm development in flowering plants: how to overcome the imprinting problem? Sex Plant Reprod,2001,14:189-194
142.Vinkenoog R, Spielman M, Adams S,Fischer R L, Dickinson H G, Scott R J. Hypomethylation promotes autonomous endosperm development and rescues postfertilization lethality in fie Mutants. The Plant Cell,2000b,12:2271-2282
143.Wang D F, Tyson M D, Jackson S S, Yadegari R. Partially redundant functions of two SET-domain polycomb-group proteins in controlling initiation of seed development in Arabidopsis. Proc Natl Acad Sci USA,2006,103:13244-13249
144.Wang X, Zhang Y, Ma Q, Zhang Z, Xue Y, Bao S and Chong K SKB1-mediated symmetric dimethylation of histone H4R3 controls flowering time in Arabidopsis. Embo J,2007,26:1934-1941
145.Warthmann N, Chen H, Ossowski S, Weigel D and Herve P Highly specific gene silencing by artificial miRNAs in rice.PLoS ONE,2008,3:e1829
146.Wang X X, Ma L G. Polycomb-group (Pc-G) proteins control seed development in Arabidopsis thaliana L. Journal of Integrative Plant Biology,2007,47(1):52-59
147.Wood C C, Robertson M, Tanner G, Peacock W J, Dennis E S,Helliwell C A. The Arabidopsis thaliana vernalization response requires a polycomb-like protein complex that also includes VERNALIZATION INSENSITIVE 3.Proc Natl Acad Sci USA,2006,103:14631-14636
148.Yadegari R, Drews G N. Female gametophyte development. The Plant Cell,2004,16: S133-S141
149.Yadegari R, Kinoshita T, Lotan O, Cohen G, Katz A, Choi Y, Katz A, Nakashima K, Harada J J, Goldberg R B, Fischer R L, Ohad N, Mutations in the FIE and MEA genes that encode interacting polycomb proteins cause parent-of-origin effects on seed development by distinct mechanisms The Plant Cell,2000,12:2367-2382
150.Yoshida N, Yanai Y, Chen L, Kato Y, Hiratsuka J, Miwa T, Sung Z R, Takahashi S. EMBRYONIC FLOWER2, a novel polycomb group protein homolog, mediates shoot development and flowering in Arabidopsis. The Plant Cell,2001,13: 2471-2481
151.Yu X, Li L, Li L, Guo M, Chory J and Yin Y;Modulation of brassinosteroid-regulated gene expression by Jumonji domain-containing proteins ELF6 and REF6 in Arabidopsis. Proc Natl Acad Sci U S A,2008,105:7618-7623
152.Yu Y, Dong A and Shen W H. Molecular characterization of the tobacco SET domain protein NtSET1 unravels its role in histone methylation, chromatin binding, and segregation. Plant J,2004,40:699-711
153.Zhang D, Yoon H G and Wong J. JMJD2A is a novel N-CoR-interacting protein and is involved in repression of the human transcription factor achaete scute-like homologue 2(ASCL2/Hash2).Mol Cell Biol,2005,25:6404-6414
154.Zhang J, Guo D, Chang Y, You C, Li X, Dai X, Weng Q, Zhang J, Chen G, Li X, Liu H, Han B,Zhang Q, Wu C.Non-random distribution of T-DNA insertions at various levels of the genome hierarchy as revealed by analyzing 13 804 T-DNA flanking sequences from an enhancer-trap mutant library. The Plant Journal,2007,49: 947-959
155.Zhou C, Zhang L, Duan J, Miki B.and Wu K. HISTONE DEACETYLASE19 is involved in jasmonic acid and ethylene signaling of pathogen response in Arabidopsis.Plant Cell,2005,17:1196-1204
156.Zilberman D and Henikoff S Genome-wide analysis of DNA methylation patterns. Development,2007,134:3959-3965