植物DNA甲基化研究进展

doi:10.11923/j.issn.2095-4050.cjas2020-0152

摘要/Abstract

摘要：

DNA甲基化是一种重要的表观遗传修饰,能够有效调控基因组稳定性。为了了解DNA甲基化对植物生长发育的影响,本文归纳了近年来植物DNA甲基化的模式,总结了植物DNA甲基化的生物学功能,概括了DNA甲基化的研究方法,最后总结了植物DNA甲基化研究中存在的问题,并指明了研究方向,为后续植物基因组研究提供理论依据。

关键词: 植物, DNA甲基化, 表观遗传, 修饰, 生长发育, 逆境胁迫, 基因组, 稳定性

Abstract:

DNA methylation is an important epigenetic modification that can effectively regulate genome stability. In order to understand the impact of DNA methylation on plant growth and development, this article summarizes plant DNA methylation patterns, concludes the physiological functions of plant DNA methylation, and reviews the research methods of DNA methylation. At last, this article sums up the problems in the study of plant DNA methylation and points out the research directions in the future, providing a theoretical basis for subsequent plant genome research.

Key words: Plants, DNA methylation, Epigenetic, Modification, Growth and Development, Adversity Stress, genome, stability

中图分类号:

S184

陈子涵, 任建国, 王俊丽. 植物DNA甲基化研究进展[J]. 农学学报, 2021, 11(11): 88-97.

Chen Zihan, Ren Jianguo, Wang Junli. Research Advances on Plant DNA Methylation[J]. Journal of Agriculture, 2021, 11(11): 88-97.

图/表 3

参考文献 98

[1]	牛亚丹, 林伊荷, 张汉清, 等. DNA甲基化与骨代谢调节及骨质疏松症研究进展[J]. 生命科学, 2020, 32(2):162-169.
[2]	汪炳良, 李水凤, 曾广文, 等. 5-azaC对萝卜茎尖DNA甲基化和开花的影响[J]. 核农学报, 2005, 19(4):265-268.
[3]	陈芳, 王子成. 5-氮杂胞苷对小麦生长发育及DNA甲基化的影响[J]. 河南大学学报:自然科学版, 2011, 41(01):61-66.
[4]	袁媛, 魏渊, 于军, 等. 表观遗传与药材道地性研究探讨[J]. 中国中药杂志, 2015, 40(13):2679-2683.
[5]	申燕红. 表观遗传改良棉——纤维培育新突破[J]. 中国纤检, 2017(7):140-141.
[6]	Zhang X Y, Junshi Y, Ambka S, et al. Genome-wide High-Resolution Mapping and Functional Analysis of DNA Methylation in Arabidopsis[J]. Cell, 2006, 126(6):1189-1201. doi: 10.1016/j.cell.2006.08.003 URL
[7]	Ryan L, Ronan C. O'M>, Julian T F, et al. Highly Integrated Single-Base Resolution Maps of the Epigenome in Arabidopsis[J]. Cell, 2008, 133(3):523-526. doi: 10.1016/j.cell.2008.03.029 URL
[8]	Marjori A M, Rebecca A M. RNA-directed DNA methylation: An epigenetic pathway of increasing complexity[J]. Nature Reviews.Genetics, 2014, 15(6):394-408.
[9]	Julie A L, Jiamu D, Christopher J, et al. Polymerase IV occupancy at RNA-directed DNA methylation sites requires SHH1[J]. Nature: International weekly journal of science, 2013, 498(7454):385-389.
[10]	Zhang H, Ma Z Y, Zeng L, et al. DTF1 is a core component of RNA-directed DNA methylation and may assist in the recruitment of Pol IV[J]. Proceedings of the National Academy of Sciences, 2013, 110(20):8290-8295. doi: 10.1073/pnas.1300585110 URL
[11]	Zheng B L, Wang Z M, Li S B, et al. Intergenic transcription by RNA polymerase II coordinates Pol IV and Pol V in siRNA-directed transcriptional gene silencing in Arabidopsis[J]. Genes and Development: a Journal Devoted to the Molecular Analysis of Gene Expression in Eukaryotes, Prokaryotes, and Viruses, 2009, 23(24):2850-2860.
[12]	Saicageethi N, Andrea D M, Kaushik P, et al. The initiation of epigenetic silencing of active transposable elements is triggered by RDR6 and 21-22 nucleotide small interfering RNAs[J]. Plant Physiology, 2013, 162:116-31. doi: 10.1104/pp.113.216481 URL
[13]	Zhong X, Hale C J, Law J A, et al. DDR complex facilitates global association of RNA polymerase V to promoters and evolutionarily young transposons[J]. Nature Structural & Molecular Biology, 2012, 19(9):870-875. doi: 10.1038/nsmb.2354 URL
[14]	Law J A, Ausin I, Johnson L M, et al. A protein complex required for polymerase V transcripts and RNA- directed DNA methylation in Arabidopsis[J]. Current Biology, 2010, 20(10):951-956. doi: 10.1016/j.cub.2010.03.062 URL
[15]	Ausin I, Todd C M, Joanne C, et al. IDN1 and IDN2 are required for de novo DNA methylation in Arabidopsis thaliana[J]. Nature Structural& Molecular Biology, 2009, 16(12):1325-1327.
[16]	Ausin I, Maxin V.C.G, Dhirendra K. S, et al. INVOLVED IN DE NOVO 2-containing complex involved in RNA-directed DNA methylation in Arabidopsis[J]. Proceedings of the National Academy of Sciences of the United States of America, 2012, 109(22):8374-8381.
[17]	Zhu Y M, Jordan R, Gudrun B, et al. A SWI/SNF chromatin-remodeling complex acts in noncoding RNA-mediated transcriptional silencing[J]. Molecular Cell, 2013, 49(2):298-309. doi: 10.1016/j.molcel.2012.11.011 URL
[18]	Zhang H, Tang K, Qian W, et al. An Rrp6-like protein positively regulates noncoding RNA levels and DNA methylation in Arabidopsis[J]. Molecular Cell, 2014, 54(3):418-430.
[19]	Henikoff S, Comai L. A DNA methyltransferase homolog with a chromodomain exists in multiple polymorphic forms in Arabidopsis[J]. Genetics, 1998, 149:307-318. pmid: 9584105
[20]	Stroud H, Do T, Du J, et al. Non-CG methylation patterns shape the epigenetic landscape in Arabidopsis.[J]. Nature structural & molecular biology, 2013, 21(1):64-72. doi: 10.1038/nsmb.2735 URL
[21]	Jiamu D, Zhong X H, Yana V, et al. Dual Binding of Chromomethylase Domains to H3K9me2-Containing Nucleosomes Directs DNA Methylation in Plants[J]. Cell, 2012, 151(1):167-180. doi: 10.1016/j.cell.2012.07.034 pmid: 23021223
[22]	James P J, Lianna J, Zuzana J, et al. Dimethylation of histone H3 lysine 9 is a critical mark for DNA methylation and gene silencing in Arabidopsis thaliana[J]. Chromosoma, 2004, 112:308-315. doi: 10.1007/s00412-004-0275-7 URL
[23]	Bartee L, Bende J, Ebbs M L. H3 lysine 9 methylation is maintained on a transcribed inverted repeat by combined action of SUVH6 and SUVH4 methyltransferases[J]. Molecular and Cellular Biology, 2005, 25(23):10507-10515. doi: 10.1128/MCB.25.23.10507-10515.2005 URL
[24]	Finnegan, E J, Dennis, E S. Isolation and identification by sequence homology of a putative cytosine methyltransferase from Arabidopsis thaliana[J]. Nucleic Acids Research, 1993, 21(10):2383-2388. pmid: 8389441
[25]	陈欣, 王子成. 植物DNA甲基转移酶[J]. 生命的化学, 2009, 29(4):534-538.
[26]	Mark W K, Douglas E R, Trevor L S, et al. Arabidopsis MET1 cytosine methyltransferase mutants[J]. Genetics, 2003, 163(3):1109-1122. doi: 10.1093/genetics/163.3.1109 URL
[27]	Lang Z B, Zhang H M, Zhu J K. Dynamics and function of DNA methylation in plants[J]. Nature reviews: molecular cell biology, 2018, 19(8):489-506. doi: 10.1038/s41580-018-0016-z URL
[28]	Caszymo X, Soringer N M, Muszynsk M G, et al. Conserved plant genes with similarity to mammalian de novo DNA methyltransferases[J]. Proceedings of the National Academy of Sciences of the United States of America, 2000, 97(9):4979-4984.
[29]	Liu Z W, Shao C R, Zhang C J, et al. The SET domain proteins SUVH2 and SUVH9 are required for Pol V occupancy at RNA-directed DNA methylation loci[J]. PLoS genetics, 2014, 10(1):e1003948. doi: 10.1371/journal.pgen.1003948 URL
[30]	Bruno H, Tatsuo K, Lucia D, et al. Endogenous targets of RNA-directed DNA methylation and Pol IV in Arabidopsis[J]. The EMBO journal, 2006, 25(12):2828-2836. doi: 10.1038/sj.emboj.7601150 URL
[31]	Assaf Z, M. Yvonne K, Hsieh P H, et al. The Arabidopsis Nucleosome Remodeler DDM1 Allows DNA Methyltransferases to Access H1-Containing Heterochromatin[J]. Cell, 2013, 153(1):193-205. doi: 10.1016/j.cell.2013.02.033 URL
[32]	李向男, 蔡宇良. 马哈利樱桃矮化砧的DNA甲基化水平及模式分析[J]. 西北植物学报, 2017, 37(5):864-871.
[33]	刘琼瑶, 黄华宏, 冯惠平, 等. 矮生观赏杉木DNA甲基化的水平与模式分析[J]. 园艺学报, 2015, 42(10):2015-2022.
[34]	李际红, 邢世岩, 张倩, 等. 叶籽银杏DNA甲基化水平与模式变异的研究[J]. 园艺学报, 2014, 41(8):1535-1544.
[35]	朱芹芹, 李忠爱, 何艳霞, 等. 表观遗传与花期调控研究进展[J]. 园艺学报, 2013, 40(9):1787-1794.
[36]	高乐旋, 陈家宽, 杨继. 表型可塑性变异的生态-发育机制及其进化意义[J]. 植物分类学报, 2008, 46(4):441-451.
[37]	Filomena D L, Pedro C, Alexandra M E J, et al. A PHD-polycomb repressive complex 2 triggers the epigenetic silencing of FLC during vernalization[J]. Proceedings of the National Academy of Sciences, 2008, 105(44):16831-16836. doi: 10.1073/pnas.0808687105 URL
[38]	James W, Gao H B, Zhang J Y, et al. Sexual-lineage-specific DNA methylation regulates meiosis in Arabidopsis[J]. Nature genetics, 2018, 50(1):130-137. doi: 10.1038/s41588-017-0008-5 URL
[39]	Kit A H, Bernacchia G, Rolin D, et al. Tissue dependent variations of DNA methylation and endoreduplication levels during tomato fruit development and ripening[J]. Planta: An International Journal of Plant Biology, 2008, 228(3):391-399.
[40]	Cheng J F, Niu Q F, Zhang B, et al. Downregulation of RdDM during strawberry fruit ripening[J]. Genome biology, 2018, 19(1):212. doi: 10.1186/s13059-018-1587-x URL
[41]	许静静, 杨凯, 王文和, 等. 病毒侵染对西伯利亚百合DNA甲基化的影响[J]. 西北植物学报, 2011, 31(5):935-941.
[42]	Wada Y, Kusano T, Sano H, et al. Association between up-regulation of stress-responsive genes and hypomethylation of genomic DNA in tobacco plants[J]. Molecular biology reports, 2004, 271(6):658-666.
[43]	Satge C, Moreau S, Sallet E, et al. Reprogramming of DNA methylation is critical for nodule development in Medicago truncatula[J]. Nature plants, 2016, 2(11):16166. doi: 10.1038/nplants.2016.166 URL
[44]	Marianna N, AlagurajL V, et al. Ploidy-dependent changes in the epigenome of symbiotic cells correlate with specific patterns of gene expression[J]. Proceedings of the National Academy of Sciences of the United States of America, 2017, 114(17):4543-4548.
[45]	Rambani A, Rice J H, Liu J Y, et al. The Methylome of Soybean Roots during the Compatible Interaction with the Soybean Cyst Nematode[J]. Plant physiology, 2015, 168(4):1364-1377. doi: 10.1104/pp.15.00826 pmid: 26099268
[46]	Hewezi T, Lane T, Piya S, et al. Cyst Nematode parasitism induces dynamic changes in the root epigenome[J]. Plant Physiol, 2017, 174(1):405-420. doi: 10.1104/pp.16.01948 pmid: 28298479
[47]	徐妍. 灰斑病菌胁迫对大豆生理生化和DNA甲基化的影响[D]. 哈尔滨:哈尔滨师范大学, 2015.
[48]	Robert H.D, Mattia P J.S, et al. Widespread dynamic DNA methylation in response to biotic stress[J]. Proceedings of the National Academy of Sciences of the United States of America, 2012, 109(32):2183-2191. doi: 10.1073/pnas.1209329109 pmid: 22733782
[49]	Tiwari S, Taylor J M, Wang M B, et al. DNA demethylases target promoter transposable elements to positively regulate stress responsive genes in Arabidopsis[J]. Genome Biology, 2014, 15(9):458. doi: 10.1186/s13059-014-0458-3 URL
[50]	郝梦真. 低温胁迫下西洋参DNA甲基化与其品质形成的关系[D]. 青岛:青岛大学, 2019.
[51]	Zhang B, Tieman D M, Jiao C, et al. Chilling-induced tomato flavor loss is associated with altered volatile synthesis and transient changes in DNA methylation[J]. Proceedings of the National Academy of Sciences of the United States of America, 2016, 113(44):12580-12585. pmid: 27791156
[52]	晁秋杰, 刘晓, 韩磊, 等. 高温胁迫诱导半夏基因组甲基化的变异分析[J]. 中国中药杂志, 2020, 45(02):341-346.
[53]	任茂, 李博, 徐延浩, 等. 高温胁迫诱导棉花甲基化变化分析[J]. 分子植物育种, 2017, 15(3):1069-1076.
[54]	曾子入, 贺从安, 张小康, 等. 高温胁迫诱导萝卜基因组甲基化变异分析[J]. 分子植物育种, 2018, 16(07):2094-2098.
[55]	Michael S, Navdeep S.C. ROS Function in Redox Signaling and Oxidative Stress[J]. Current Biology: CB, 2014, 24(10):R453-R462. doi: 10.1016/j.cub.2014.03.034 URL
[56]	施江, 熊雨婕, 张晗, 等. 遮荫影响半夏DNA甲基化的MSAP分析[J]. 中国中药杂志, 2020, 45(06):1311-1315.
[57]	韩雅楠, 赵犇鹏, 崔向军. 非生物胁迫蒙古黄芪基因组的MSAP分析[J]. 基因组学与应用生物学, 2020, 39(7):3119-3125.
[58]	潘雅姣, 傅彬英, 王迪, 等. 水稻干旱胁迫诱导DNA甲基化时空变化特征分析[J]. 中国农业科学, 2009, 42(9):3009-3018.
[59]	Xu J D, Zhou S S, Gong X Q, et al. Single-base methylome analysis reveals dynamic epigenomic differences associated with water deficit in apple[J]. Plant biotechnology journal, 2018, 16(2):672-687. doi: 10.1111/pbi.2018.16.issue-2 URL
[60]	Rafael R A, Maria I P M, Aan P O G, et al. DEMETER and REPRESSOR OF SILENCING 1 encode 5-methylcytosine DNA glycosylases[J]. Proceedings of the National Academy of Sciences of the United States of America, 2006, 103(18):6853-6858.
[61]	杜驰, 张冀, 张丽丽, 张富春. 盐胁迫下盐穗木DNA甲基化程度与去甲基化酶基因(Ros1)表达的相关性研究[J]. 新疆农业科学, 2017, 54(5):878-885.
[62]	董蔚, 刘锡江, 高天雪, 等. 盐胁迫下蒺藜苜蓿MtMYBS1的DNA甲基化和组蛋白修饰状态分析[J]. 植物生理学报, 2018, 54(10):1596-1604.
[63]	Dong W, Song Y G, Zhao Z, et al. The Medicago truncatula R2R3-MYB transcription factor gene MtMYBS1 enhances salinity tolerance when constitutively expressed in Arabidopsis thaliana[J]. Biochemical and biophysical research communications, 2017, 490(2):225-230. doi: S0006-291X(17)31144-0 pmid: 28602696
[64]	赵晓辉, 徐正进, 刘宛, 等. 镉、铜胁迫下拟南芥基因组DNA甲基化的MSAP分析[J]. 安徽农业科学, 2013, 41(5):1911-1915.
[65]	何玲莉, 沈虹, 王燕, 等. 铅胁迫下萝卜基因组DNA甲基化分析[J]. 核农学报, 2015, 29(7):1278-1284.
[66]	张凯凯, 陈兴银, 杨鹏, 等. 不同浓度镉胁迫对孔雀草DNA甲基化的影响[J]. 草业科学, 2016, 33(9):1673-1680.
[67]	丁国华, 郭丹蒂, 关旸, 等. 重金属铅镉对濒危植物中华水韭(Isoetes sinensis)DNA甲基化的影响[J]. 农业环境科学学报, 2017, 36(2):246-249.
[68]	殷欣. 镉胁迫下大豆生理生化特性及DNA甲基化变异的研究[D]. 哈尔滨:哈尔滨师范大学, 2016.
[69]	黑淑梅. 重金属铬对小麦根系DNA甲基化水平的影响[J]. 吉林农业科学, 2012, 37(2):14-15,26.
[70]	Kuo K C, Mccune R A, Gehrke C W, et al. Quantitative reversed-phase high performance liquid chromatographic determination of major and modified deoxyribonucleosides in DNA[J]. Nucleic acids research, 1980, 8(20):4763-4778. pmid: 7003544
[71]	Wagner I, Capesius I. Determination of 5-methylcytosine from plant DNA by high-performance liquid chromatography[J]. Biochimica et Biophysica Acta, 1981, 654(1):52-56. pmid: 7272310
[72]	Xiong L Z, XU C G, Maroof S M A. Patterns of cytosine methylation in an elite rice hybrid and its parental lines,detected by a methylation-sensitive amplification polymorphism technique. Molecular and General Genetics, 1999, 261(3):439-446. pmid: 10323223
[73]	Hayatsu H. The bisulfite genomic sequencing used in the analysis of epigenetic states,a technique in the emerging environmental genotoxicology research[J]. Mutation Research: International Journal on Mutagenesis, Chromosome Breakage and Related Subjects, 2008, 659(1/2):77-82.
[74]	Costello J F, Fouse S D, Nagarajan R P. Genome-scale DNA methylation analysis[J]. Epigenomics, 2010, 2(1):105-117. doi: 10.2217/epi.09.35 URL
[75]	Thorne N P, Rakyan V K, Johnson N, et al. A Bayesian deconvolution strategy for immunoprecipitation-based DNA methylome analysis[J]. Nature biotechnology, 2008, 26(7):779-785. doi: 10.1038/nbt1414 URL
[76]	Osabe K, Jenny D, Bedon, et al. Genetic and DNA methylation changes in cotton(Gossypium)genotypes and tissues[J]. PloS one, 2014, 9(1):e86049. doi: 10.1371/journal.pone.0086049 URL
[77]	Li L X, Jie Y, Dong Y S, et al. Optimization of an HPLC Method for Determining the Genomic Methylation Levels of Taxus Cells[J]. Journal of chromatographic science, 2016, 54(2):1-6.
[78]	Gao Y, Hao J L, Wang Z, et al. DNA methylation levels in different tissues in tea plant via an optimized HPLC method[J]. Horticulture, Environment, and Biotechnology, 2019, 60(6):967-974. doi: 10.1007/s13580-019-00180-2 URL
[79]	Baurens F C, Nicolleau J, Legavre T, et al. Genomic DNA methylation of juvenile and mature Acacia mangium micropropagated in vitro with reference to leaf morphology as a phase change marker[J]. Tree physiology, 2004, 24(4):401-407. doi: 10.1093/treephys/24.4.401 URL
[80]	Gao Y, Hao J L, Wang Z, Song K J, et al. DNA methylation levels in different tissues in tea plant via an optimized HPLC method[J]. Horticulture, Environment, and Biotechnology, 2019, 60(6):967-974. doi: 10.1007/s13580-019-00180-2 URL
[81]	Finke A, Rozhon W, Pecinka A. Analysis of DNA Methylation Content and Patterns in Plants[J]. Methods in molecular biology (Clifton, N.J.), 2018, 1694(24):277-298.
[82]	Vos P, Hogers R, Bleeker M, et al. AFLP a new technique for DNA fingerprinting[J]. Nucleic Acids Research, 1995, 23(21):4407-4414. pmid: 7501463
[83]	Roberts R J, Vincze T, Posfai J, et al. REBASE--a database for DNA restriction and modification: enzymes, genes and genomes[J]. Nucleic Acids Research, 2015, 43(Database issue):D298-299. doi: 10.1093/nar/gku1046 URL
[84]	Li A, Hu B Q, Xue Z Y, et al. DNA Methylation in Genomes of Several Annual Herbaceous and Woody Perennial Plants of Varying Ploidy as Detected by MSAP[J]. Plant Molecular Biology Reporter, 2011, 29(4):784-793. doi: 10.1007/s11105-010-0280-3 URL
[85]	Herrera C M, Bazaga P. Untangling individual variation in natural populations: ecological, genetic and epigenetic correlates of long-term inequality in herbivory[J]. Molecular ecology, 2011, 20(8):1675-1688. doi: 10.1111/j.1365-294X.2011.05026.x pmid: 21466603
[86]	陈瑞娟, 何蕾, 孙梨宗, 等. 铜胁迫对拟南芥幼苗生长和基因组DNA甲基化的影响[J]. 生态学杂志, 2015, 34(9):2650-2657.
[87]	Xin C H, Hou R K, Wu F, et al. Analysis of cytosine methylation status in potato by methylation-sensitive amplified polymorphisms under low-temperature stress[J]. Journal of Plant Biology, 2015, 58(6):383-390. doi: 10.1007/s12374-015-0316-1 URL
[88]	龚治, 张亚楠, 周世豪, 等. 甲基化敏感扩增多态性技术(MSAP)在生态学中的应用[J]. 生物安全学报, 2016, 25(1):7-12.
[89]	Baubec T, Akalin A. Genome-wide analysis of DNA methylation patterns by high-throughput sequencing. In: Aransay A, Lavín Trueba J, eds. Field Guidelines for Genetic Experimental Designs in High-throughput Sequencing[M]. Heidelberg: Springer, 2016, 9:197-221.
[90]	Sun Y Y, Fan M, He Y J. DNA Methylation Analysis of the Citrullus lanatus Response to Cucumber Green Mottle Mosaic Virus Infection by Whole-Genome Bisulfite Sequencing[J]. Genes, 2019, 10(5):344. doi: 10.3390/genes10050344 URL
[91]	Li Q, Peter J H, Nathan M.S. Detection of DNA Methylation by Whole-Genome Bisulfite Sequencing[J]. Methods in molecular biology(Clifton, N.J.), 2018, 1676:185-196.
[92]	Martin S, Michiel V B, Magdalena W, et al. Plant-RRBS, a bisulfite and next-generation sequencing-based methylome profiling method enriching for coverage of cytosine positions[J]. BMC Plant Biology, 2017, 17(1):115. doi: 10.1186/s12870-017-1070-y URL
[93]	Aniruddha C J R, Lan M M, Morison I M, et al. Tools and Strategies for Analysis of Genome-Wide and Gene-Specific DNA Methylation Patterns[J]. Methods in Molecular Biology, 2017(1537):249-277.
[94]	Xing X Y, Zhang B, Li D F, et al. Comprehensive Whole DNA Methylome Analysis by Integrating MeDIP-seq and MRE-seq[J]. Methods in molecular biology(Clifton, N. J.), 2018, 1708(12):209-246.
[95]	Vinning K J, Pomraning K R, Wilhelm L J, et al. Dynamic DNA cytosine methylation in the Populus trichocarpa genome: tissue-level variation and relationship to gene expression[J]. BMC genomics, 2012, 13:27. doi: 10.1186/1471-2164-13-27 URL
[96]	房媛媛, 肖亮, 卢文杰, 等. DNA甲基化研究进展及其在木本植物中的发展趋势[J]. 中国科学:生命科学, 2020, 50(2):154-166.
[97]	Daniel Z, Mary G, Robert T K, et al. Genome-wide analysis of Arabidopsis thaliana DNA methylation uncovers an interdependence between methylation and transcription[J]. Nature genetics, 2007, 39(1):61-69. doi: 10.1038/ng1929 URL
[98]	Li X Y, Wang X F, He K, et al. High-resolution mapping of epigenetic modifications of the rice genome uncovers interplay between DNA methylation, histone methylation, and gene expression[J]. The Plant cell, 2008, 20(2):259-276. doi: 10.1105/tpc.107.056879 URL

方法	基本原理	主要覆盖范围	是否需要参考基因	参考文献
高效液相色谱	水解、紫外检测	全基因组	否	[71]
甲基化敏感扩增多态性	限制性内切酶	高CG区域	否	[72]
全基因组亚硫酸盐测序	亚硫酸盐处理	全基因组	是	[73]
简并代表性亚硫酸氢盐测序技术	亚硫酸盐处理+限制性内切酶	启动子和CpG岛	是	[74]
甲基化DNA免疫共沉淀测序技术	抗体富集	高CG区域	是	[75]

方法	基本原理	主要覆盖范围	是否需要参考基因	参考文献
高效液相色谱	水解、紫外检测	全基因组	否	[71]
甲基化敏感扩增多态性	限制性内切酶	高CG区域	否	[72]
全基因组亚硫酸盐测序	亚硫酸盐处理	全基因组	是	[73]
简并代表性亚硫酸氢盐测序技术	亚硫酸盐处理+限制性内切酶	启动子和CpG岛	是	[74]
甲基化DNA免疫共沉淀测序技术	抗体富集	高CG区域	是	[75]