植物组蛋白去甲基化酶研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2022-0165

摘要/Abstract

摘要：

组蛋白甲基化修饰在调控真核细胞的基因表达中起重要作用,组蛋白甲基化的水平受到组蛋白甲基转移酶（histone methyltransferases,HMTs）和组蛋白去甲基化酶（histone demethylases,HDMs）的动态调控,其中以在赖氨酸残基上的甲基化修饰最为常见。系统总结了植物组蛋白赖氨酸去甲基化酶的最新研究进展,阐述其在植物开花时间调控、昼夜节律调控等生长发育过程,以及植物对干旱、温度、病原菌等胁迫响应中的重要作用及其调控机制,为进一步利用植物组蛋白去甲基化酶在作物遗传改良中提供参考。

关键词: 组蛋白修饰, 组蛋白去甲基化酶, 生长发育, 胁迫响应

Abstract:

Histone methylation plays an important role in regulating gene expression in eukaryotic cells. Histone methylation is dynamically regulated by histone methyltransferases（HMTs）and demethylases（HDMs）,among which methylation on lysine residues is the most common. In this paper,the latest research progress of plant histone lysine demethylase is systematically summarized,and its important role and regulation mechanism in the growth and development process of plant flowering time regulation and circadian rhythm regulation,as well as the responses of plants to drought,temperature,pathogenic bacteria and other stresses are expounded. The paper provides reference for further utilization of plant histone lysine demethylase in crop genetic improvement.

Key words: histone modification, histone lysine demethylases, growth and development, stress response

汤茜茜, 林楚宇, 陶增. 植物组蛋白去甲基化酶研究进展[J]. 生物技术通报, 2022, 38(7): 13-22.

TANG Qian-qian, LIN Chu-yu, TAO Zeng. Research Progress in Histone Demethylase in Plant[J]. Biotechnology Bulletin, 2022, 38(7): 13-22.

图/表 2

参考文献 87

[1]	Luger K, Mäder AW, Richmond RK, et al. Crystal structure of the nucleosome core particle at 2. 8 A resolution[J]. Nature, 1997, 389(6648):251-260. doi: 10.1038/38444 URL
[2]	Kouzarides T. Chromatin modifications and their function[J]. Cell, 2007, 128(4):693-705. doi: 10.1016/j.cell.2007.02.005 pmid: 17320507
[3]	Bannister AJ, Kouzarides T. Regulation of chromatin by histone modifications[J]. Cell Res, 2011, 21(3):381-395. doi: 10.1038/cr.2011.22 pmid: 21321607
[4]	Strahl BD, Allis CD. The language of covalent histone modifications[J]. Nature, 2000, 403(6765):41-45. doi: 10.1038/47412 URL
[5]	Jenuwein T, Allis CD. Translating the histone code[J]. Science, 2001, 293(5532):1074-1080. pmid: 11498575
[6]	Chen XS, Hu YF, Zhou DX. Epigenetic gene regulation by plant Jumonji group of histone demethylase[J]. Biochim Biophys Acta, 2011, 1809(8):421-426.
[7]	Zhang YJ, Sun ZX, Jia JQ, et al. Overview of histone modification[J]. Adv Exp Med Biol, 2021, 1283:1-16.
[8]	Martin C, Zhang Y. The diverse functions of histone lysine methylation[J]. Nat Rev Mol Cell Biol, 2005, 6(11):838-849.
[9]	Zhang Y, Reinberg D. Transcription regulation by histone methylation:interplay between different covalent modifications of the core histone tails[J]. Genes Dev, 2001, 15(18):2343-2360. doi: 10.1101/gad.927301 URL
[10]	Bedford MT, Clarke SG. Protein arginine methylation in mammals:who, what, and why[J]. Mol Cell, 2009, 33(1):1-13. doi: 10.1016/j.molcel.2008.12.013 pmid: 19150423
[11]	Strahl BD, Ohba R, Cook RG, et al. Methylation of histone H3 at lysine 4 is highly conserved and correlates with transcriptionally active nuclei in Tetrahymena[J]. PNAS, 1999, 96(26):14967-14972. pmid: 10611321
[12]	Black JC, van Rechem C, Whetstine JR. Histone lysine methylation dynamics:establishment, regulation, and biological impact[J]. Mol Cell, 2012, 48(4):491-507. doi: 10.1016/j.molcel.2012.11.006 URL
[13]	Klose RJ, Zhang Y. Regulation of histone methylation by demethylimination and demethylation[J]. Nat Rev Mol Cell Biol, 2007, 8(4):307-318. doi: 10.1038/nrm2143 URL
[14]	Lu FL, Li GL, Cui X, et al. Comparative analysis of JmjC domain-containing proteins reveals the potential histone demethylases in Arabidopsis and rice[J]. J Integr Plant Biol, 2008, 50(7):886-896. doi: 10.1111/j.1744-7909.2008.00692.x URL
[15]	Jiang DH, Yang WN, He YH, et al. Arabidopsis relatives of the human lysine-specific demethylase1 repress the expression of FWA and FLOWERING LOCUS C and thus promote the floral transition[J]. Plant Cell, 2007, 19(10):2975-2987. doi: 10.1105/tpc.107.052373 URL
[16]	Henderson IR, Dean C. Control of Arabidopsis flowering:the chill before the bloom[J]. Development, 2004, 131(16):3829-3838. pmid: 15289433
[17]	Simpson GG, Dean C. Arabidopsis, the Rosetta stone of flowering time?[J]. Science, 2002, 296(5566):285-289. pmid: 11951029
[18]	Kardailsky I, Shukla VK, Ahn JH, et al. Activation tagging of the floral inducer FT[J]. Science, 1999, 286(5446):1962-1965. pmid: 10583961
[19]	Michaels SD, Amasino RM. FLOWERING LOCUS C encodes a novel MADS domain protein that acts as a repressor of flowering[J]. Plant Cell, 1999, 11(5):949-956. pmid: 10330478
[20]	Suárez-López P, Wheatley K, Robson F, et al. CONSTANS mediates between the circadian clock and the control of flowering in Arabidopsis[J]. Nature, 2001, 410(6832):1116-1120. doi: 10.1038/35074138 URL
[21]	He YH, Michaels SD, Amasino RM. Regulation of flowering time by histone acetylation in Arabidopsis[J]. Science, 2003, 302(5651):1751-1754. doi: 10.1126/science.1091109 URL
[22]	Noh B, Lee SH, Kim HJ, et al. Divergent roles of a pair of homologous jumonji/zinc-finger-class transcription factor proteins in the regulation of Arabidopsis flowering time[J]. Plant Cell, 2004, 16(10):2601-2613. doi: 10.1105/tpc.104.025353 URL
[23]	Yang HC, Howard M, Dean C. Physical coupling of activation and derepression activities to maintain an active transcriptional state at FLC[J]. Proc Natl Acad Sci USA, 2016, 113(33):9369-9374. doi: 10.1073/pnas.1605733113 URL
[24]	Zheng SZ, Hu HM, Ren HM, et al. The Arabidopsis H3K27me3 demethylase JUMONJI 13 is a temperature and photoperiod dependent flowering repressor[J]. Nat Commun, 2019, 10(1):1303. doi: 10.1038/s41467-019-09310-x URL
[25]	Gan ES, Xu YF, Wong JY, et al. Jumonji demethylases moderate precocious flowering at elevated temperature via regulation of FLC in Arabidopsis[J]. Nat Commun, 2014, 5:5098. doi: 10.1038/ncomms6098 URL
[26]	Lu FL, Cui X, Zhang SB, et al. JMJ 14 is an H3K4 demethylase regulating flowering time in Arabidopsis[J]. Cell Res, 2010, 20(3):387-390. doi: 10.1038/cr.2010.27 URL
[27]	Yang WN, Jiang DH, Jiang JF, et al. A plant-specific histone H3 lysine 4 demethylase represses the floral transition in Arabidopsis[J]. Plant J, 2010, 62(4):663-673. doi: 10.1111/j.1365-313X.2010.04182.x URL
[28]	Ning YQ, Ma ZY, Huang HW, et al. Two novel NAC transcription factors regulate gene expression and flowering time by associating with the histone demethylase JMJ14[J]. Nucleic Acids Res, 2015, 43(3):1469-1484. doi: 10.1093/nar/gku1382 URL
[29]	Rodrigues VL, Dolde U, Sun B, et al. A microProtein repressor complex in the shoot meristem controls the transition to flowering[J]. Plant Physiol, 2021, 187(1):187-202. doi: 10.1093/plphys/kiab235 pmid: 34015131
[30]	Yang HC, Han ZF, Cao Y, et al. A companion cell-dominant and developmentally regulated H3K4 demethylase controls flowering time in Arabidopsis via the repression of FLCexpression[J]. PLoS Genet, 2012, 8(4):e1002664.
[31]	Yang HC, Mo HX, Fan D, et al. Overexpression of a histone H3K4 demethylase, JMJ15, accelerates flowering time in Arabidopsis[J]. Plant Cell Rep, 2012, 31(7):1297-1308. doi: 10.1007/s00299-012-1249-5 URL
[32]	Yokoo T, Saito H, Yoshitake Y, et al. Se14, encoding a JmjC domain-containing protein, plays key roles in long-day suppression of rice flowering through the demethylation of H3K4me3 of RFT1[J]. PLoS One, 2014, 9(4):e96064.
[33]	Dutta A, Choudhary P, Caruana J, et al. JMJ27, an Arabidopsis H3K9 histone demethylase, modulates defense against Pseudomonas syringae and flowering time[J]. Plant J, 2017, 91(6):1015-1028. doi: 10.1111/tpj.13623 URL
[34]	Hung FY, Lai YC, Wang JH, et al. The Arabidopsis histone demethylase JMJ28 regulates CONSTANS by interacting with FBH transcription factors[J]. Plant Cell, 2021, 33(4):1196-1211. doi: 10.1093/plcell/koab014 URL
[35]	Wang ZY, Tobin EM. Constitutive expression of the CIRCADIAN CLOCK ASSOCIATED 1(CCA1)gene disrupts circadian rhythms and suppresses its own expression[J]. Cell, 1998, 93(7):1207-1217. pmid: 9657153
[36]	Schaffer R, Ramsay N, Samach A, et al. The late elongated hypocotyl mutation of Arabidopsis disrupts circadian rhythms and the photoperiodic control of flowering[J]. Cell, 1998, 93(7):1219-1229. pmid: 9657154
[37]	Strayer C, Oyama T, Schultz TF, et al. Cloning of the Arabidopsis clock gene TOC1, an autoregulatory response regulator homolog[J]. Science, 2000, 289(5480):768-771. pmid: 10926537
[38]	Alabadí D, Oyama T, Yanovsky MJ, et al. Reciprocal regulation between TOC1 and LHY/CCA1 within the Arabidopsis circadian clock[J]. Science, 2001, 293(5531):880-883. pmid: 11486091
[39]	Harmer SL, Kay SA. Positive and negative factors confer phase-specific circadian regulation of transcription in Arabidopsis[J]. Plant Cell, 2005, 17(7):1926-1940. doi: 10.1105/tpc.105.033035 URL
[40]	Gendron JM, Pruneda-Paz JL, Doherty CJ, et al. Arabidopsis circadian clock protein, TOC1, is a DNA-binding transcription factor[J]. Proc Natl Acad Sci USA, 2012, 109(8):3167-3172. doi: 10.1073/pnas.1200355109 URL
[41]	Jones MA, Covington MF, DiTacchio L, et al. Jumonji domain protein JMJD5 functions in both the plant and human circadian systems[J]. Proc Natl Acad Sci USA, 2010, 107(50):21623-21628. doi: 10.1073/pnas.1014204108 URL
[42]	Lu SX, Knowles SM, Webb CJ, et al. The Jumonji C domain-containing protein JMJ30 regulates period length in the Arabidopsis circadian clock[J]. Plant Physiol, 2011, 155(2):906-915. doi: 10.1104/pp.110.167015 URL
[43]	Mizuno T, Nomoto Y, Oka H, et al. Ambient temperature signal feeds into the circadian clock transcriptional circuitry through the EC night-time repressor in Arabidopsis thaliana[J]. Plant Cell Physiol, 2014, 55(5):958-976. doi: 10.1093/pcp/pcu030 URL
[44]	Jones MA, Morohashi K, Grotewold E, et al. Arabidopsis JMJD5/JMJ30 acts independently of LUX ARRHYTHMO within the plant circadian clock to enable temperature compensation[J]. Front Plant Sci, 2019, 10:57. doi: 10.3389/fpls.2019.00057 URL
[45]	Hung FY, Chen FF, Li CL, et al. The LDL1/2-HDA6 histone modification complex interacts with TOC1 and regulates the core circadian clock components in Arabidopsis[J]. Front Plant Sci, 2019, 10:233. doi: 10.3389/fpls.2019.00233 URL
[46]	Song QX, Huang T, Yu HH, et al. Diurnal regulation of SDG2 and JMJ14 by circadian clock oscillators orchestrates histone modification rhythms in Arabidopsis[J]. Genome Biol, 2019, 20(1):170. doi: 10.1186/s13059-019-1777-1 URL
[47]	Lee HG, Seo PJ. The Arabidopsis JMJ29 protein controls circadian oscillation through diurnal histone demethylation at the CCA1 and PRR9 loci[J]. Genes, 2021, 12(4):529. doi: 10.3390/genes12040529 URL
[48]	Bentsink L, Koornneef M. Seed dormancy and germination[J]. Arabidopsis Book, 2008, 6:e0119.
[49]	Bouyer D, Roudier F, Heese M, et al. Polycomb repressive complex 2 controls the embryo-to-seedling phase transition[J]. PLoS Genet, 2011, 7(3):e1002014.
[50]	Cho JN, Ryu JY, Jeong YM, et al. Control of seed germination by light-induced histone arginine demethylation activity[J]. Dev Cell, 2012, 22(4):736-748. doi: 10.1016/j.devcel.2012.01.024 URL
[51]	van Zanten M, Zöll C, Wang Z, et al. HISTONE DEACETYLASE 9 represses seedling traits in Arabidopsis thaliana dry seeds[J]. Plant J, 2014, 80(3):475-488. doi: 10.1111/tpj.12646 URL
[52]	Zhao ML, Yang SG, Liu XC, et al. Arabidopsis histone demethylases LDL1 and LDL2 control primary seed dormancy by regulating DELAY OF GERMINATION 1 and ABA signaling-related genes[J]. Front Plant Sci, 2015, 6:159.
[53]	Wu JF, Ichihashi Y, Suzuki T, et al. Abscisic acid-dependent histone demethylation during postgermination growth arrest in Arabidopsis[J]. Plant Cell Environ, 2019, 42(7):2198-2214. doi: 10.1111/pce.13547 URL
[54]	Wu JF, Yan ML, Zhang DW, et al. Histone demethylases coordinate the antagonistic interaction between abscisic acid and brassinosteroid signaling in Arabidopsis[J]. Front Plant Sci, 2020, 11:596835. doi: 10.3389/fpls.2020.596835 URL
[55]	Wang TJ, Huang SZ, Zhang A, et al. JMJ17-WRKY40 and HY5-ABI5 modules regulate the expression of ABA-responsive genes in Arabidopsis[J]. New Phytol, 2021, 230(2):567-584. doi: 10.1111/nph.17177 URL
[56]	Lim PO, Kim HJ, Nam HG. Leaf senescence[J]. Annu Rev Plant Biol, 2007, 58:115-136. doi: 10.1146/annurev.arplant.57.032905.105316 URL
[57]	Liu P, Zhang SB, Zhou B, et al. The histone H3K4 demethylase JMJ16 represses leaf senescence in Arabidopsis[J]. Plant Cell, 2019, 31(2):430-443. doi: 10.1105/tpc.18.00693 URL
[58]	Wang XL, Gao J, Gao S, et al. The H3K27me3 demethylase REF6 promotes leaf senescence through directly activating major senescence regulatory and functional genes in Arabidopsis[J]. PLoS Genet, 2019, 15(4):e1008068.
[59]	Chinnusamy V, Zhu JK. Epigenetic regulation of stress responses in plants[J]. Curr Opin Plant Biol, 2009, 12(2):133-139. doi: 10.1016/j.pbi.2008.12.006 URL
[60]	Kim JM, To TK, Ishida J, et al. Alterations of lysine modifications on the histone H3 N-tail under drought stress conditions in Arabidopsis thaliana[J]. Plant Cell Physiol, 2008, 49(10):1580-1588. doi: 10.1093/pcp/pcn133 URL
[61]	Huang SZ, Zhang A, Jin JB, et al. Arabidopsis histone H3K4 demethylase JMJ17 functions in dehydration stress response[J]. New Phytol, 2019, 223(3):1372-1387. doi: 10.1111/nph.15874 URL
[62]	Song T, Zhang Q, Wang HQ, et al. OsJMJ703, a rice histone demethylase gene, plays key roles in plant development and responds to drought stress[J]. Plant Physiol Biochem, 2018, 132:183-188. doi: 10.1016/j.plaphy.2018.09.007 URL
[63]	Wang QL, Liu P, Jing H, et al. JMJ27-mediated histone H3K9 demethylation positively regulates drought-stress responses in Arabidopsis[J]. New Phytol, 2021, 232(1):221-236. doi: 10.1111/nph.17593 URL
[64]	Charng YY, Liu HC, Liu NY, et al. A heat-inducible transcription factor, HsfA2, is required for extension of acquired thermotolerance in Arabidopsis[J]. Plant Physiol, 2007, 143(1):251-262. doi: 10.1104/pp.106.091322 URL
[65]	Lämke J, Brzezinka K, Altmann S, et al. A hit-and-Run heat shock factor governs sustained histone methylation and transcriptional stress memory[J]. EMBO J, 2016, 35(2):162-175. doi: 10.15252/embj.201592593 pmid: 26657708
[66]	Cui XY, Zheng Y, Lu Y, et al. Metabolic control of histone demethylase activity involved in plant response to high temperature[J]. Plant Physiol, 2021, 185(4):1813-1828. doi: 10.1093/plphys/kiab020 URL
[67]	Liu JZ, Feng LL, Gu XT, et al.An H3K27me3 demethylase-HSFA2 regulatory loop orchestrates transgenerational thermomemory in Arabidopsis[J]. Cell Res, 2019, 29(5):379-390.
[68]	Yamaguchi N, Matsubara S, Yoshimizu K, et al. H3K27me3 demethylases alter HSP22 and HSP17. 6C expression in response to recurring heat in Arabidopsis[J]. Nat Commun, 2021, 12(1):3480. doi: 10.1038/s41467-021-23766-w URL
[69]	Yamaguchi N, Ito T. JMJ histone demethylases balance H3K27me3 and H3K4me3 levels at the HSP21 locus during heat acclimation in Arabidopsis[J]. Biomolecules, 2021, 11(6):852. doi: 10.3390/biom11060852 URL
[70]	Gómez-Gómez L, Boller T. Flagellin perception:a paradigm for innate immunity[J]. Trends Plant Sci, 2002, 7(6):251-256. pmid: 12049921
[71]	Zipfel C, Felix G. Plants and animals:a different taste for microbes?[J]. Curr Opin Plant Biol, 2005, 8(4):353-360. pmid: 15922649
[72]	Coll NS, Epple P, Dangl JL. Programmed cell death in the plant immune system[J]. Cell Death Differ, 2011, 18(8):1247-1256. doi: 10.1038/cdd.2011.37 pmid: 21475301
[73]	Koornneef A, Pieterse CMJ. Cross talk in defense signaling[J]. Plant Physiol, 2008, 146(3):839-844. doi: 10.1104/pp.107.112029 pmid: 18316638
[74]	Pieterse CMJ, Leon-Reyes A, van der Ent S, et al. Networking by small-molecule hormones in plant immunity[J]. Nat Chem Biol, 2009, 5(5):308-316. doi: 10.1038/nchembio.164 pmid: 19377457
[75]	Thilmony R, Underwood W, He SY. Genome-wide transcriptional analysis of the Arabidopsis thaliana interaction with the plant pathogen Pseudomonas syringae pv. tomato DC3000 and the human pathogen Escherichia coli O157:H7[J]. Plant J, 2006, 46(1):34-53. pmid: 16553894
[76]	Alvarez-Venegas R, Abdallat AA, Guo M, et al. Epigenetic control of a transcription factor at the cross section of two antagonistic pathways[J]. Epigenetics, 2007, 2(2):106-113. pmid: 17965588
[77]	Ding B, Wang GL. Chromatin versus pathogens:the function of epigenetics in plant immunity[J]. Front Plant Sci, 2015, 6:675. doi: 10.3389/fpls.2015.00675 pmid: 26388882
[78]	Deleris A, Greenberg MVC, Ausin I, et al. Involvement of a Jumonji-C domain-containing histone demethylase in DRM2-mediated maintenance of DNA methylation[J]. EMBO Rep, 2010, 11(12):950-955. doi: 10.1038/embor.2010.158 URL
[79]	le Masson I, Jauvion V, Bouteiller N, et al. Mutations in the Arabidopsis H3K4me2/3 demethylase JMJ14 suppress posttranscriptional gene silencing by decreasing transgene transcription[J]. Plant Cell, 2012, 24(9):3603-3612. doi: 10.1105/tpc.112.103119 URL
[80]	Li D, Liu RY, Singh D, et al. JMJ14 encoded H3K4 demethylase modulates immune responses by regulating defence gene expression and pipecolic acid levels[J]. New Phytol, 2020, 225(5):2108-2121. doi: 10.1111/nph.16270 URL
[81]	Noh SW, Seo RR, Park HJ, et al. Two Arabidopsis homologs of human lysine-specific demethylase function in epigenetic regulation of plant defense responses[J]. Front Plant Sci, 2021, 12:688003. doi: 10.3389/fpls.2021.688003 URL
[82]	Chan C, Zimmerli L. The histone demethylase IBM1 positively regulates Arabidopsis immunity by control of defense gene expression[J]. Front Plant Sci, 2019, 10:1587. doi: 10.3389/fpls.2019.01587 URL
[83]	Li TT, Chen XS, Zhong XC, et al. Jumonji C domain protein JMJ705-mediated removal of histone H3 lysine 27 trimethylation is involved in defense-related gene activation in rice[J]. Plant Cell, 2013, 25(11):4725-4736. doi: 10.1105/tpc.113.118802 URL
[84]	Hou YX, Wang LY, Wang L, et al. 704 positively regulates rice defense response against Xanthomonas oryzae pv. oryzae infection via reducing H3K4me2/3 associated with negative disease resistance regulators[J]. BMC Plant Biol, 2015, 15:286. doi: 10.1186/s12870-015-0674-3 URL
[85]	Saze H, Shiraishi A, Miura A, et al. Control of genic DNA methylation by a jmjC domain-containing protein in Arabidopsis thaliana[J]. Science, 2008, 319(5862):462-465. doi: 10.1126/science.1150987 URL
[86]	Audonnet L, Shen Y, Zhou DX. JMJ24 antagonizes histone H3K9 demethylase IBM1/JMJ25 function and interacts with RNAi pathways for gene silencing[J]. Gene Expr Patterns, 2017, 25/26:1-7. doi: 10.1016/j.gep.2017.04.001 URL
[87]	Antunez-Sanchez J, Naish M, Ramirez-Prado JS, et al. A new role for histone demethylases in the maintenance of plant genome integrity[J]. eLife, 2020, 9:e58533.

KDM类型 Type of KDM		底物 Substrate	拟南芥/水稻Arab-idopsis/ Oryza sativa
LSD1		H3K4	AtFLD
			AtLDL1
			AtLDL2
			AtLDL3
			OsHDM701
			OsHDM702
			OsHDM703
			OsHDM704
JHDMs	KDM5/JARID1	H3K4	AtJMJ14
			AtJMJ15
			AtJMJ16
			AtJMJ17
			AtJMJ18
			AtJMJ19
			OsJMJ703
			OsJMJ704
			OsJMJ708
	KDM4	H3K9	AtEFL6/AtJMJ11
			AtREF6/AtJMJ12
			AtJMJ13
			OsJMJ701
			OsJMJ702
			OsJMJ705
			OsJMJ706
			OsJMJ707
	KDM3/JHDM2	H3K9	AtJMJ24
			AtIBM1/AtJMJ25
			AtJMJ26
			AtJMJ27
			AtJMJ28
			AtJMJ29
			OsJMJ715
			OsJMJ716
			OsJMJ718
			OsJMJ719
			OsJMJ720
	JmjCdomain-only	H3K36、H3K27	AtJMJ30
			AtJMJ31
			AtJMJ32
			OsJMJ709
			OsJMJ710
			OsJMJ711
			OsJMJ712
			OsJMJ713
			OsJMJ714
			OsJMJ717

KDM类型 Type of KDM		底物 Substrate	拟南芥/水稻Arab-idopsis/ Oryza sativa
LSD1		H3K4	AtFLD
			AtLDL1
			AtLDL2
			AtLDL3
			OsHDM701
			OsHDM702
			OsHDM703
			OsHDM704
JHDMs	KDM5/JARID1	H3K4	AtJMJ14
			AtJMJ15
			AtJMJ16
			AtJMJ17
			AtJMJ18
			AtJMJ19
			OsJMJ703
			OsJMJ704
			OsJMJ708
	KDM4	H3K9	AtEFL6/AtJMJ11
			AtREF6/AtJMJ12
			AtJMJ13
			OsJMJ701
			OsJMJ702
			OsJMJ705
			OsJMJ706
			OsJMJ707
	KDM3/JHDM2	H3K9	AtJMJ24
			AtIBM1/AtJMJ25
			AtJMJ26
			AtJMJ27
			AtJMJ28
			AtJMJ29
			OsJMJ715
			OsJMJ716
			OsJMJ718
			OsJMJ719
			OsJMJ720
	JmjCdomain-only	H3K36、H3K27	AtJMJ30
			AtJMJ31
			AtJMJ32
			OsJMJ709
			OsJMJ710
			OsJMJ711
			OsJMJ712
			OsJMJ713
			OsJMJ714
			OsJMJ717