‘月月粉’月季PP2C家族基因鉴定及非生物胁迫响应分析

doi:10.16420/j.issn.0513-353x.2022-0752

摘要/Abstract

摘要：

利用生物信息方法从月季‘月月粉’（Rosa chinensis‘Old Blush’）基因组中共鉴定出69个PP2C家族成员，其编码氨基酸的长度在122 ~ 1 082 aa之间，等电点介于4.03 ~ 9.34，大部分蛋白呈亲水性。系统进化树分析将其分为12个亚族，同一亚族内基因结构和保守基序具有较高的相似性。基因片段复制事件是PP2C家族基因扩张的主要原因。启动子顺式作用元件分析表明约85%的月季PP2C含有逆境应答元件、激素应答元件等，推测其在响应逆境胁迫的过程中发挥重要作用。月季PP2C基因的表达具有组织和发育时期特异性，且部分基因参与干旱和高温胁迫响应过程，其中3个A亚族基因在干旱胁迫中响应明显，1个A亚族基因和2个G亚族基因在高温胁迫中响应明显。通过qRT-PCR分析11个不同亚族PP2C基因在月季失水胁迫和ABA处理下的表达模式，发现6个A亚族基因（RC0G0099200、RC1G0458700、RC1G0569200、RC2G0066800、RC2G0089100和RC5G0571100）和1个G亚族基因（RC3G0083700）的表达量显著升高，表明这些基因可能受ABA诱导并参与植物失水胁迫响应。

关键词: 月季, PP2C基因家族, 非生物胁迫, ABA, 基因表达

Abstract:

A total of 69 members of the PP2C family were identified from the genome of Rosa chinensis‘Old Blush’using bioinformatics techniques. The majority of the proteins were hydrophilic，and the amino acid lengths ranged from 122 to 1 082 amino acids，with isoelectric points ranging 4.03 and 9.34. Phylogenetic analysis indicated that PP2Cs were divided into 12 subfamily groups，and the genetic structure and conserved motif of the same subfamily had a high similarity. Segmental duplication events were the main reason for the expansion of PP2C family genes. A significant function for the PP2C family of genes in response to stress may be inferred from the prediction of the cis-elements of promoters，which revealed that nearly 85% of PP2Cs had stress response elements and phytohormone response elements. The expression of rose PP2Cs was specific in tissue and developmental stage，and some genes were involved in the response to drought and high-temperature stress. Among them，three members of subgroup A were significantly responsive to drought stress，one member of subgroup A and two members of subgroup G were significantly responsive to high-temperature stress. Using qRT-PCR，the expression patterns of 11 PP2Cs genes under ABA and dehydration treatment were examined. Six subgroup A genes （RC0G0099200，RC1G0458700，RC1G0569200，RC2G0066800，RC2G0089100 and RC5G0571100）and a subgroup G gene（RC3G0083700）showed significantly higher expression levels following two treatments，suggesting that these genes may be activated by ABA and take part in the dehydration response.

Key words: Rosa chinensis, PP2C gene family, abiotic stress, ABA, gene expression

申玉晓, 邹金玉, 罗平, 尚文倩, 李永华, 何松林, 王政, 石力匀. ‘月月粉’月季PP2C家族基因鉴定及非生物胁迫响应分析[J]. 园艺学报, 2023, 50(10): 2139-2156.

SHEN Yuxiao, ZOU Jinyu, LUO Ping, SHANG Wenqian, LI Yonghua, HE Songlin, WANG Zheng, SHI Liyun. Genome-wide Identification and Abiotic Stress Response Analysis of PP2C Family Genes in Rosa chinensis‘Old Blush’[J]. Acta Horticulturae Sinica, 2023, 50(10): 2139-2156.

图/表 7

参考文献 43

[1]	Cao J, Jiang M, Li P, Chu Z. 2016. Genome-wide identification and evolutionary analyses of the PP2C gene family with their expression profiling in response to multiple stresses in Brachypodium distachyon. BMC Genomics, 17 (1):175. doi: 10.1186/s12864-016-2526-4 URL
[2]	Chen C J, Chen H, Zhang Y, Thomas H R, Frank M H, He Y H, Xia R. 2020. Tbtools:an integrative toolkit developed for interactive analyses of big biological data. Mol Plant, 13 (8):1194-1202. doi: 10.1016/j.molp.2020.06.009 URL
[3]	Chu M L, Chen P W, Meng S F, Xu P, Lan W Z. 2021. The Arabidopsis phosphatase PP2C49 negatively regulates salt tolerance through inhibition of AtHKT1;1. Journal of Integrative Plant Biology, 63(3):528-542.
[4]	Daccord N, Celton J M, Linsmith G, Becker C, Choisne N, Schijlen E, Geest H, Bianco L, Micheletti D, Velasco R, Di P E A, Gouzy J, Rees D, Guerif P, Muranty H, Durel C E, Laurens F, Lespinasse Y, Gaillard S, Aubourg S, Quesneville H, Weigel D, Weg E, Troggio M, Bucher E. 2017. High-quality denovo assembly of the apple genome and methylome dynamics of early fruit development. Nature Genetics, 49 (7):1099-1106. doi: 10.1038/ng.3886
[5]	Finkelstein R. 2013. Abscisic acid synthesis and response. The Arabidopsis Book, 11:e0166. doi: 10.1199/tab.0166 URL
[6]	Haider M S, Khan N, Pervaiz T, Zhongjie L, Nasim M, Jogaiah S, Mushtaq N, Jiu S T, Fang J G. 2019. Genome-wide identification,evolution,and molecular characterization of the PP2C gene family in woodland strawberry. Gene, 702:27-35. doi: 10.1016/j.gene.2019.03.025 URL
[7]	Haider M S, Zhang C, Kurjogi M M, Pervaiz T, Zheng T, Zhang C, Lide C, Shangguan L, Fang J. 2017. Insights into grapevine defense response against drought as revealed by biochemical,physiological and RNA-Seq analysis. Scientific Reports, 7 (1):13134. doi: 10.1038/s41598-017-13464-3
[8]	Han Y, Yu J, Zhao T, Cheng T, Wang J, Yang W, Pan H, Zhang Q. 2019. Dissecting the genome-wide evolution and function of R2R3-MYB transcription factor family in Rosa chinensis. Genes, 10:823. doi: 10.3390/genes10100823 URL
[9]	He Honghong, Lu Zhihao, Ma Zonghuan, Liang Guoping, Ma Lijuan, Wan Peng, Mao Juan. 2018. Genome-wide identification and expression analysis of the PP2C gene family in Vitis vinifera. Acta Horticulturae Sinica, 45 (7):1237-1250. (in Chinese)
	何红红, 路志浩, 马宗桓, 梁国平, 马丽娟, 万鹏, 毛娟. 2018. 葡萄PP2C家族基因的鉴定与表达分析. 园艺学报, 45 (7):1237-1250. doi: 10.16420/j.issn.0513-353x.2017-0824 URL
[10]	He Y J, Liu Y, Li M Z, Lamin-Samu A T, Yang D D, Yu X L, Izhar M, Jan I, Ali M, Lu G. 2021. The Arabidopsis SMALL AUXIN UP RNA 32 protein regulates ABA-mediated responses to drought stress. Frontiers in Plant Science, 12:625493. doi: 10.3389/fpls.2021.625493 URL
[11]	Huang X S, Luo T, Fu X Z, Fan Q J, Liu J H. 2011. Cloning and molecular characterization of a mitogen-activated protein kinase gene from Poncirus trifoliata whose ectopic expression confers dehydration/drought tolerance in transgenic tobacco. Journal of Experimental Botany, 62 (14):5191-5206. doi: 10.1093/jxb/err229 URL
[12]	Jia X, Feng H, Bu Y H, Ji N Z, Lyu Y M, Zhao S W. 2021. Comparative transcriptome and weighted gene co-expression network analysis identify key transcription factors of Rosa chinensis‘Old Blush’after exposure to a gradual drought stress followed by recovery. Front Genet, 12:690264. doi: 10.3389/fgene.2021.690264 URL
[13]	Jeong S, Lim C W, Lee S C. 2021. CaADIP1-dependent CaADIK1-kinase activation is required for ABA signaling and drought stress response in Capsicum annuum. The New Phytologis, 231 (6):2247-2261. doi: 10.1111/nph.v231.6 URL
[14]	Li W H, Yang J, Gu X. 2005. Expression divergence between duplicate genes. Trends in Genetics, 21 (11):602-607. doi: 10.1016/j.tig.2005.08.006 URL
[15]	Li Z Q, Xing W, Luo P, Zhang F J, Jin X L, Zhang M H. 2019. Comparative transcriptome analysis of Rosa chinensis ‘Slater's crimson China’ provides insights into the crucial factors and signaling pathways in heat stress response. Plant Physiol Biochem, 142:312-331. doi: 10.1016/j.plaphy.2019.07.002 URL
[16]	Liang B, Sun Y F, Wang J, Zheng Y, Zhang W B, Xu Y D, Li Q, Leng P. 2020. Tomato Protein Phosphatase 2C(SlPP2C3)influences fruit ripening onset and fruit glossiness. Journal of Experimental Botany, 72 (7):2403-2418. doi: 10.1093/jxb/eraa593 URL
[17]	Liu X, Zhu Y, Zhai H, Cai H, Ji W, Luo X, Li J, Bai X. 2012. AtPP2CG1,a protein phosphatase 2C,positively regulates salt tolerance of Arabidopsis in abscisic acid-dependent manner. Biochem Biophys Res Commun, 422 (4):710-715. doi: 10.1016/j.bbrc.2012.05.064 URL
[18]	Luan S. 2003. Protein phosphatases in plants. Annu Rev Plant Biol, 54:63-92. pmid: 14502985
[19]	Lv J, Liu J Y, Ming Y H, Shi Y T, Song C P, Gong Z Z, Yang S H, Ding Y L. 2021. Reciprocal regulation between the negative regulator PP2CG 1 phosphatase and the positive regulator OST1 kinase confers cold response in Arabidopsis. Journal of Integrative Plant Biology, 63 (8):1568-1587. doi: 10.1111/jipb.v63.8 URL
[20]	Mahajan A, Yuan C, Lee H, Chen E S, Wu P Y, Tsai M D. 2008. Structure andfunction of the phosphothreonine-specific FHA domain. Science Signaling, 1 (51):re12.
[21]	Park S Y, Fung P, Nishimura N, Jensen D R, Fujii H, Zhao Y, Lumba S, Santiago J, Rodrigues A, Chow T F, Alfred S E, Bonetta D, Finkelstein R, Provart N J, Desveaux D, Rodriguez P L, McCourt P, Zhu J K, Schroeder J I, Volkman B F, Cutler S R. 2009. Abscisic acid inhibits type 2C protein phosphatases via the PYR/PYL family of START proteins. Science, 324 (5930):1068-1071. doi: 10.1126/science.1173041 URL
[22]	Perkin L C, Bell A, Hinze L L, Suh C P, Arick M A, Peterson D G, Udall J A. 2021. Genome assembly of two nematode-resistant cotton lines(Gossypium hirsutum L.). G3 Genes/Genomes/Genetics, 11 (11):276.
[23]	Pertea M, Kim D, Pertea G M, Leek J T, Salzberg S L. 2016. Transcript-level expression analysis of RNA-seq experiments with HISAT,StringTie and Ballgown. Nature Protocols, 11 (9):1650-1667. doi: 10.1038/nprot.2016.095
[24]	Raymond O, Gouzy J, Just J, Badouin H, Verdenaud M, Lemainque A, Vergne P, Moja S, Choisne N, Pont C, Carrère S, Caissard J C, Couloux A, Cottret L, Aury J M, Szécsi J, Latrasse D, Madoui M A, François L, Fu X, Yang S H, Dubois A, Piola F, Larrieu A, Perez M, Labadie K, Perrier L, Govetto B, Labrousse Y, Villand P, Bardoux C, Boltz V, Lopez-Roques C, Heitzler P, Vernoux T, Vandenbussche M, Quesneville H, Boualem A, Bendahmane A, Liu C, Le Bris M, Salse J, Baudino S, Benhamed M, Wincker P, Bendahmane M. 2018. The Rosa genome provides new insights into the domestication of modern roses. Nature Genetics, 50 (6):772-777. doi: 10.1038/s41588-018-0110-3
[25]	Saint-Oyant L H, Ruttink T, Hamama L, Kirov I, Lakhwani D, Zhou N N, Bourke P M, Daccord N, Leus L, Schulz D, Geest H V D, Hesselink T, Laere K V, Debray K,Bal. zergue S, Thouroude T, Chastellier A, Jeauffre J, Voisine L, Gaillard S, Borm T J A, Arens P, Voorrips R E,Mal. iepaard C, Neu E, Linde M, Le Paslier M C, Bérard A, Bounon R, Clotault J, Choisne N, Quesneville H, Kawamura K, Aubourg S, Sakr S, Smulders M J M, Schijlen E, Bucher E, Debener T, de Riek J, Foucher F. 2018. A high-quality genome sequence of Rosa chinensis to elucidate ornamental traits. Nature Plants, 4 (7):473-484. doi: 10.1038/s41477-018-0166-1 pmid: 29892093
[26]	Schweighofer A, Hirt H, Meskiene I. 2004. Plant PP2C phosphatases:emerging functions in stress signaling. Trends in Plant Science, 9 (5):236-243. doi: 10.1016/j.tplants.2004.03.007 pmid: 15130549
[27]	Schweighofer A, Kazanaviciute V, Scheikl E, Teige M, Doczi R, Hirt H, Schwanninger M, Kant M, Schuurink R, Mauch F, Buchala A, Cardinale F, Meskiene I. 2007. The PP2C-type phosphatase AP2C1,which negatively regulates MPK4 and MPK6,modulates innate immunity,jasmonic acid,and ethylene levels in Arabidopsis. Plant Cell, 19 (7):2213-2224. pmid: 17630279
[28]	Servet C, Benhamed M, Latrasse D, Kim W, Delarue M, Zhou D X. 2008. Characterization of a phosphatase 2C protein as an interacting partner of the histone acetyltransferase GCN5 in Arabidopsis. Biochim Biophys Acta, 1779 (6-7):376-382.
[29]	Shazadee H, Khan N, Wang J, Wang C, Zeng J, Huang Z, Wang X. 2019. Identification and expression profiling of protein phosphatases 2C(PP2C) gene family in Gossypium hirsutum L. International Journal of Molecular Sciences, 20 (6):1395. doi: 10.3390/ijms20061395 URL
[30]	Singh A, Pandey A, Srivastava A K, Tran L S, Pandey G K. 2016. Plant protein phosphatases 2C:from genomic diversity to functional multiplicity and importance in stress management. Critical Reviews in Biotechnology, 36 (6):1023-1035. doi: 10.3109/07388551.2015.1083941 URL
[31]	Song S K, Lee M M, Clark S E. 2006. POL and PLL 1 phosphatases are CLAVATA1 signaling intermediates required for Arabidopsis shoot and floral stem cells. Development, 133 (23):4691-4698. doi: 10.1242/dev.02652 URL
[32]	Umbrasaite J, Schweighofer A, Kazanaviciute V, Magyar Z, Ayatollahi Z, Unterwurzacher V, Choopayak C, Boniecka J, Murray J A, Bogre L, Meskiene I. 2010. Substrate analysis of Arabidopsis PP2C-type protein phosphatases. PLoS ONE, 5 (12):e15357. doi: 10.1371/journal.pone.0015357 URL
[33]	Wang G M, Sun X, Guo Z H, Joldersma D, Guo L, Qiao X, Qi K J, Gu C, Zhang S L. 2021. Genome-wide identification and evolution of the PP2C gene family in eight rosaceae species and expression analysis under stress in Pyrus bretschneideri. Frontiers in Genetics, 12:770014. doi: 10.3389/fgene.2021.770014 URL
[34]	Wang J, Zhang S H, Fu Y P, He T T, Wang X W. 2020. Analysis of dynamic global transcriptional atlas reveals common regulatory networks of hormones and photosynthesis across nicotiana varieties in response to long-term drought. Frontiers in Plant Science, 11:672. doi: 10.3389/fpls.2020.00672 URL
[35]	Wong J H, Klejchová M, Snipes S A, Nagpal P, Bak G, Wang B, Dunlap S, Park M Y, Kunkel E N, Trinidad B, Reed J W, Blatt M R, Gray W M. 2021. SAUR proteins and PP2C.D phosphatases regulate H⁺-ATPases and K⁺ channels to control stomatal movements. Plant Physio, 185 (1):256-273. doi: 10.1093/plphys/kiaa023 URL
[36]	Xue T T, Wang D, Zhang S Z, Ehlting J, Ni F, Jakab S, Zheng C C, Zhong Y. 2008. Genome-wide and expression analysis of protein phosphatase 2C in rice and Arabidopsis. BMC Genomics, 9 (1):550. doi: 10.1186/1471-2164-9-550
[37]	Yang Jie, Chen Rong,Hu Wen-juan,Wu Qiao-ling,Tong Xiao-nan,Li Xing-tao. 2022. ldentification and expression analysis of PP2C gene family in Poncirus trifoliata. Journal of Fruit Science, 39 (4):532-547. (in Chinese)
	杨杰, 陈蓉, 胡文娟, 吴巧玲, 佟晓楠, 李兴涛. 2022. 枳PP2C基因家族的鉴定及表达分析. 果树学报, 39 (4):532-547.
[38]	Yu X F, Han J P, Wang E F, Xiao J, Hu R, Yang G X, He G Y. 2019. Genome-wide identification and homoeologous expression analysis of PP2C genes in wheat(Triticum aestivum L.). Frontiers in Genetics, 10:561. doi: 10.3389/fgene.2019.00561 URL
[39]	Zhang Chunling, Zhou Lijie, Wang Guiluan, Li Yuanyuan, Wang Xiaofei, Hao Yujin. 2018. Molecular cloning of protein phosphatase gene MdPP2C-Like in apple and the response analysis of ABA. Acta Horticulturae Sinica, 45 (7):1225-1236. (in Chinese) doi: 10.16420/j.issn.0513-353x.2017-0726
	张春玲, 周李杰, 王桂栾, 李媛媛, 王小非, 郝玉金. 2018. 苹果蛋白磷酸酶基因MdPP2C-Like 的克隆及其对ABA 的响应分析. 园艺学报, 45 (7):1225-1236. doi: 10.16420/j.issn.0513-353x.2017-0726
[40]	Zhang Tingting, Li Yuxin, Zhang Deyao, Kang Yuqian, Wang Jian, Song Xiqiang, Zhou Yang. 2021. Genome-wide identification and expression analyses of PP2C gene family in Dendrobium catenatum. Acta Horticulturae Sinica, 48 (12):2458-2470. (in Chinese) doi: 10.16420/j.issn.0513-353x.2021-0382 URL
	张婷婷, 李雨欣, 张德遥, 康宇乾, 王健, 宋希强, 周杨. 2021. 铁皮石斛蛋白磷酸酶PP2C家族基因鉴定及其表达分析. 园艺学报, 48 (12):2458-2470. doi: 10.16420/j.issn.0513-353x.2021-0382 URL
[41]	Zhang Z H, Ali S, Zhang T X, Wang W P, Xie L N. 2020. Identification,evolutionary and expression analysis of PYL-PP2C-SnRK2s gene families in soybean. Plants, 9 (10):1356. doi: 10.3390/plants9101356 URL
[42]	Zhao Shu-ping, Tan Hong-bin, Lu Dan, Pei Li-li, Cui Xi-yan, Ma You-zhi, Chen Ming, Xu Zhao-shi, Zhang Xiao-hong. 2017. Research progress of plant protein kinase mediated abiotic stress and hormone signal transduction pathwa. Journal of Plant Genetic Resources, 18 (2):358-366. (in Chinese)
	赵书平, 谈宏斌, 鹿丹, 裴丽丽, 崔喜艳, 马有志, 陈明, 徐兆师, 张小红. 2017. 植物蛋白激酶介导的非生物胁迫和激素信号转导途径的研究进展. 植物遗传资源学报, 18 (2):358-366. doi: 10.13430/j.cnki.jpgr.2017.02.022
[43]	Zhou Y, Peng Q, Zhang L, Cheng S, Zeng L, Dong F, Yang Z. 2018. Characterization of enzymes specifically producing chiral flavor compounds (R)- and (S)-1-phenylethanol from tea(Camellia sinensis)flowers. Food Chemistry, 280:27-33. doi: 10.1016/j.foodchem.2018.12.035 URL

基因 ID	上游引物（5′-3′）	下游引物（5′-3′）
Gene ID	Forward primer	Reverse primer
RC0G0099200	TGGGAAATACTGGCAAGGGTC	CACTTCGGATGGCTGCTTTACT
RC1G0458700	GGTCTAGTTGGAGAAAGCG	ATCGGCGGCAATACACTTA
RC1G0478300	AACTCTCCGATCTTCTCACCGT	ATGCCTCTGACTCTGAAACCTTGC
RC1G0569200	AGGATGCTGGCAATTCTGT	TACTCCCGTCCCTCGTTTCA
RC2G0066800	AAGTGATGGACTGTGGGATGT	CTGACGTTTTCTTTTCGTTTTGA
RC2G0089100	ATGATGAGGGTGAGATTTGC	CCTTCCCATTGGAGCTTGTC
RC3G0083700	CTGATGTTAAACCAGCGGA	GTCCAATGTCAGCGAAGC
RC3G0340800	CTCCGAACTTCTGGAACTGT	ATCATATCTTGTTGGCACCC
RC5G0571100	AAGAGAAGGGTAATCTGGTCAAA	TCCGAACTTGGCGTCAGG
RC6G0593000	GCGGTGCCATTTAGAGTAGGT	CCCTGAATGACTACAATTACAATCC
RC7G0167900	CCGAGATTATGATGTCCCAAGAA	GTCGTAGACGGCGAAGAAGTG
RcGAPDH	TGAAGGGTGGTGCCAAGAA	AAGGGGAGCAAGACAGTTGG

基因 ID	上游引物（5′-3′）	下游引物（5′-3′）
Gene ID	Forward primer	Reverse primer
RC0G0099200	TGGGAAATACTGGCAAGGGTC	CACTTCGGATGGCTGCTTTACT
RC1G0458700	GGTCTAGTTGGAGAAAGCG	ATCGGCGGCAATACACTTA
RC1G0478300	AACTCTCCGATCTTCTCACCGT	ATGCCTCTGACTCTGAAACCTTGC
RC1G0569200	AGGATGCTGGCAATTCTGT	TACTCCCGTCCCTCGTTTCA
RC2G0066800	AAGTGATGGACTGTGGGATGT	CTGACGTTTTCTTTTCGTTTTGA
RC2G0089100	ATGATGAGGGTGAGATTTGC	CCTTCCCATTGGAGCTTGTC
RC3G0083700	CTGATGTTAAACCAGCGGA	GTCCAATGTCAGCGAAGC
RC3G0340800	CTCCGAACTTCTGGAACTGT	ATCATATCTTGTTGGCACCC
RC5G0571100	AAGAGAAGGGTAATCTGGTCAAA	TCCGAACTTGGCGTCAGG
RC6G0593000	GCGGTGCCATTTAGAGTAGGT	CCCTGAATGACTACAATTACAATCC
RC7G0167900	CCGAGATTATGATGTCCCAAGAA	GTCGTAGACGGCGAAGAAGTG
RcGAPDH	TGAAGGGTGGTGCCAAGAA	AAGGGGAGCAAGACAGTTGG

亚族 Sub- group	基因ID Gene ID	染色体位置 Chromosome location	编码区长/bp CDS length	氨基酸数 Amino acid	外显子数 No. of exon	分子量 Molecular weight	等电点 pI	亲水系数 GRAVY	亚细胞定位 Subcellular localization
A	RC0G0047200	Chr00：4828630 ~ 4831238	699	232	4	25 695.51	5.53	-0.031	线粒体内膜Mitochondrial inner membrane
	RC0G0099200	Chr00：11758820 ~ 11762242	1 659	552	4	59 394.42	4.96	-0.063	质膜Plasma membrane
	RC1G0458700	Chr01：55057837 ~ 55059388	1 236	411	3	44 724.48	5.59	-0.377	细胞核Nucleus
	RC1G0569200	Chr01：62821746 ~ 62824036	1 161	386	3	42 500.89	6.98	-0.380	细胞核Nucleus
	RC2G0066800	Chr02：5299187 ~ 5304192	1 167	388	4	42 372.51	5.11	-0.344	质膜Plasma membrane
	RC2G0089100	Chr02：6920939 ~ 6922931	1 431	476	4	52 094.86	5.55	-0.336	质膜Plasma membrane
	RC5G0377600	Chr05：37177437 ~ 37179960	1 236	411	5	46 446.77	7.89	-0.403	质膜Plasma membrane
	RC5G0571100	Chr05：66869860 ~ 66872143	1 203	400	4	43 414.04	5.46	-0.310	细胞核Nucleus
	RC6G0593000	Chr06：66284925 ~ 66287701	1 656	551	4	59 602.25	4.60	-0.138	质膜Plasma membrane
	RC7G0167900	Chr07：13015997 ~ 13016725	465	154	1	16 745.85	6.06	-0.321	细胞质Cytoplasm
	RC7G0168100	Chr07：13018177 ~ 13019011	450	149	2	15 981.51	6.08	0.281	细胞质Cytoplasm
B	RC1G0478300	Chr01：56571375 ~ 56574342	1 143	380	4	41 067.86	8.18	-0.255	细胞核Nucleus
	RC4G0375700	Chr04：49184893 ~ 49188051	1 740	579	5	63 097.54	8.81	-0.546	细胞核Nucleus
	RC5G0606100	Chr05：70444924 ~ 70446964	1 149	382	4	41 507.95	7.53	-0.358	细胞核Nucleus
	RC6G0015800	Chr06：1590043 ~ 1592204	1 323	440	5	49 069.04	6.99	-0.331	细胞核Nucleus
C	RC3G0258700	Chr03：34531274 ~ 34531798	525	174	1	19 636.09	4.42	-0.137	细胞质Cytoplasm
	RC3G0259000	Chr03：34572348 ~ 34575352	2 046	681	4	76 180.77	5.67	-0.499	细胞核Nucleus
	RC4G0275900	Chr04：39562228 ~ 39565705	1 785	594	4	64 995.55	4.92	-0.125	质膜Plasma membrane
	RC5G0553600	Chr05：64738021 ~ 64741007	2 208	735	4	81 714.31	5.43	-0.486	细胞质Cytoplasm
	RC7G0290700	Chr07：26235143 ~ 26240816	2 301	766	5	83 960.48	5.27	-0.542	细胞核Nucleus
D	RC1G0426500	Chr01：52335182 ~ 52338968	1 155	384	4	42 742.27	8.82	-0.172	质膜Plasma membrane
	RC2G0128300	Chr02：10555792 ~ 10559454	1 182	393	4	43 613.79	8.56	-0.223	质膜Plasma membrane
	RC2G0207300	Chr02：18414833 ~ 18417905	1 119	372	4	41 544.15	5.91	-0.258	细胞核Nucleus
	RC3G0144700	Chr03：22626525 ~ 22629894	1 149	382	4	43 154.29	6.43	-0.222	线粒体基质Mitochondrial matrix space
	RC3G0245900	Chr03：33482641 ~ 33485442	1 143	380	4	42 790.86	9.34	-0.362	细胞核Nucleus
	RC4G0305900	Chr04：43009598 ~ 43013252	1 155	384	4	42 277.11	6.31	-0.182	细胞核Nucleus
	RC6G0083700	Chr06：8901670 ~ 8903762	1 149	382	4	42 629.68	8.78	-0.288	细胞核Nucleus
	RC6G0295500	Chr06：39208249 ~ 39213326	1 188	395	4	43 907.85	8.45	-0.291	细胞核Nucleus
E	RC2G0570600	Chr02：65118745 ~ 65121601	1 413	470	5	52 006.45	5.45	-0.488	细胞核Nucleus
	RC3G0300800	Chr03：37901846 ~ 37903908	1 359	452	5	50 180.95	4.92	-0.529	细胞质Cytoplasm
	RC3G0340800	Chr03：41156572 ~ 41159167	1 125	374	5	41 586.10	5.84	-0.343	细胞核Nucleus
	RC3G0415800	Chr03：46406403 ~ 46410803	1 500	499	5	54 901.47	5.28	-0.451	叶绿体基质Chloroplast stroma
	RC4G0174800	Chr04：25897436 ~ 25902101	1 233	410	5	44 205.25	7.96	-0.160	细胞核Nucleus
	RC4G0290900	Chr04：41153285 ~ 41155207	1 164	387	5	42 900.06	8.19	-0.217	内质网Endoplasmic reticulum
	RC5G0637500	Chr05：74500818-74504788	1 602	533	5	58 727.27	5.24	-0.404	叶绿体基质Chloroplast stroma
	RC6G0585500	Chr06：65787874 ~ 65789999	1 014	337	4	36 815.15	4.67	-0.238	细胞质Cytoplasm
F	RC1G0561000	Chr01：62166916 ~ 62170959	1 050	349	8	38 095.69	5.30	-0.349	线粒体基质Mitochondrial matrix space
	RC5G0161600	Chr05：12241573 ~ 12242147	384	127	2	13 699.74	6.26	0.189	细胞质Cytoplasm
	RC5G0287900	Chr05：25199030 ~ 25202031	849	282	5	31 280.45	6.07	-0.424	细胞质Cytoplasm
	RC6G0448200	Chr06：55396868 ~ 55401178	879	292	8	31 419.82	5.04	-0.334	细胞质Cytoplasm
	RC7G0553900	Chr07：63513147 ~ 63518291	849	282	5	31 005.94	7.73	-0.491	线粒体基质Mitochondrial matrix space
G	RC2G0115900	Chr02：9573649 ~ 9576363	1 260	419	4	45 563.01	6.43	-0.214	细胞核Nucleus
	RC2G0023300	Chr02：1683170 ~ 1685525	1 038	345	4	38 186.16	5.23	-0.292	细胞质Cytoplasm
	RC3G0083700	Chr03：14633161 ~ 14635520	1 176	391	4	43 297.18	4.93	-0.175	过氧化物酶体Peroxisome
	RC4G0312700	Chr04：43671766 ~ 43674255	1 140	379	4	42 030.04	5.33	-0.015	内质网Endoplasmic reticulum
	RC6G0313800	Chr06：41654079 ~ 41656687	1 158	385	4	41 854.12	5.25	-0.280	细胞质Cytoplasm
	RC6G0447100	Chr06：55271914 ~ 55274566	1 134	377	4	41 279.55	5.40	-0.315	细胞质Cytoplasm
	RC7G0347300	Chr07：33066169 ~ 33069400	1 089	362	4	39 359.71	5.58	-0.226	细胞质Cytoplasm
H	RC3G0339900	Chr03：41087943 ~ 41091127	1 308	435	7	46 914.75	6.20	-0.114	内质网Endoplasmic reticulum
	RC4G0389000	Chr04：50185684 ~ 50188564	1 293	430	9	46 157.67	6.23	-0.137	细胞质Cytoplasm
	RC6G0560400	Chr06：63913434 ~ 63917402	1 293	430	9	46 572.74	5.06	-0.319	质膜Plasma membrane
I	RC2G0409900	Chr02：47147013 ~ 47154068	1 212	403	11	44 511.31	5.32	-0.302	内质网Endoplasmic reticulum
	RC6G0451300	Chr06：55557811 ~ 55562884	1 152	383	10	42 008.38	4.59	-0.225	质膜Plasma membrane
J	RC3G0144900	Chr03：22635737 ~ 22638365	1 434	477	7	52 805.38	5.43	-0.232	内质网Endoplasmic reticulum
	RC6G0396700	Chr06：50768308 ~ 50778537	3 144	1 047	22	118 157.21	5.74	-0.345	细胞质Cytoplasm
K	RC2G0161300	Chr02：13346491 ~ 13350340	1 320	439	4	46 451.30	7.64	-0.095	细胞质Cytoplasm
	RC5G0146400	Chr05：10980909 ~ 10981772	864	287	1	31 988.92	5.48	-0.538	细胞核Nucleus
	RC5G0168100	Chr05：12965254 ~ 12966114	861	286	1	31 063.42	5.33	-0.106	细胞质Cytoplasm
	RC5G0616200	Chr05：71710833 ~ 71713396	915	304	11	32 791.37	5.51	0.045	细胞核Nucleus
	RC6G0481800	Chr06：58085355 ~ 58088427	1 512	503	4	53 601.63	6.07	-0.277	线粒体基质Mitochondrial matrix space
	RC7G0438400	Chr07：47207142 ~ 47214206	1 866	621	7	66 851.18	4.77	-0.624	细胞质Cytoplasm
L	RC2G0063500	Chr02：4938066 ~ 4945505	3 249	1 082	15	119 968.55	5.10	-0.268	叶绿体类囊体膜Chloroplast thylakoid membrane
	RC4G0257100	Chr04：37263358 ~ 37266748	1 281	426	3	46 285.35	5.08	-0.413	叶绿体基质Chloroplast stroma
Others	RC1G0573600	Chr01：63257965 ~ 63261116	1 200	399	11	43 032.47	5.24	-0.136	过氧化物酶体Peroxisome
	RC4G0054600	Chr04：5904928 ~ 5907032	840	279	5	31 486.20	9.00	-0.428	过氧化物酶体Peroxisome
	RC4G0234300	Chr04：34375420 ~ 34381498	1 707	568	13	62 714.19	5.46	-0.268	内质网Endoplasmic reticulum
	RC5G0408300	Chr05：4088239 ~ 40883657	369	122	3	13 395.99	4.03	0.011	细胞质Cytoplasm
	RC6G0347300	Chr06：45436974 ~ 45440004	1 977	658	1	72 871.19	5.71	-0.167	过氧化物酶体Peroxisome
	RC7G0024700	Chr07：1516095 ~ 1519053	999	332	10	36 056.93	7.69	-0.269	细胞核Nucleus

亚族 Sub- group	基因ID Gene ID	染色体位置 Chromosome location	编码区长/bp CDS length	氨基酸数 Amino acid	外显子数 No. of exon	分子量 Molecular weight	等电点 pI	亲水系数 GRAVY	亚细胞定位 Subcellular localization
A	RC0G0047200	Chr00：4828630 ~ 4831238	699	232	4	25 695.51	5.53	-0.031	线粒体内膜Mitochondrial inner membrane
	RC0G0099200	Chr00：11758820 ~ 11762242	1 659	552	4	59 394.42	4.96	-0.063	质膜Plasma membrane
	RC1G0458700	Chr01：55057837 ~ 55059388	1 236	411	3	44 724.48	5.59	-0.377	细胞核Nucleus
	RC1G0569200	Chr01：62821746 ~ 62824036	1 161	386	3	42 500.89	6.98	-0.380	细胞核Nucleus
	RC2G0066800	Chr02：5299187 ~ 5304192	1 167	388	4	42 372.51	5.11	-0.344	质膜Plasma membrane
	RC2G0089100	Chr02：6920939 ~ 6922931	1 431	476	4	52 094.86	5.55	-0.336	质膜Plasma membrane
	RC5G0377600	Chr05：37177437 ~ 37179960	1 236	411	5	46 446.77	7.89	-0.403	质膜Plasma membrane
	RC5G0571100	Chr05：66869860 ~ 66872143	1 203	400	4	43 414.04	5.46	-0.310	细胞核Nucleus
	RC6G0593000	Chr06：66284925 ~ 66287701	1 656	551	4	59 602.25	4.60	-0.138	质膜Plasma membrane
	RC7G0167900	Chr07：13015997 ~ 13016725	465	154	1	16 745.85	6.06	-0.321	细胞质Cytoplasm
	RC7G0168100	Chr07：13018177 ~ 13019011	450	149	2	15 981.51	6.08	0.281	细胞质Cytoplasm
B	RC1G0478300	Chr01：56571375 ~ 56574342	1 143	380	4	41 067.86	8.18	-0.255	细胞核Nucleus
	RC4G0375700	Chr04：49184893 ~ 49188051	1 740	579	5	63 097.54	8.81	-0.546	细胞核Nucleus
	RC5G0606100	Chr05：70444924 ~ 70446964	1 149	382	4	41 507.95	7.53	-0.358	细胞核Nucleus
	RC6G0015800	Chr06：1590043 ~ 1592204	1 323	440	5	49 069.04	6.99	-0.331	细胞核Nucleus
C	RC3G0258700	Chr03：34531274 ~ 34531798	525	174	1	19 636.09	4.42	-0.137	细胞质Cytoplasm
	RC3G0259000	Chr03：34572348 ~ 34575352	2 046	681	4	76 180.77	5.67	-0.499	细胞核Nucleus
	RC4G0275900	Chr04：39562228 ~ 39565705	1 785	594	4	64 995.55	4.92	-0.125	质膜Plasma membrane
	RC5G0553600	Chr05：64738021 ~ 64741007	2 208	735	4	81 714.31	5.43	-0.486	细胞质Cytoplasm
	RC7G0290700	Chr07：26235143 ~ 26240816	2 301	766	5	83 960.48	5.27	-0.542	细胞核Nucleus
D	RC1G0426500	Chr01：52335182 ~ 52338968	1 155	384	4	42 742.27	8.82	-0.172	质膜Plasma membrane
	RC2G0128300	Chr02：10555792 ~ 10559454	1 182	393	4	43 613.79	8.56	-0.223	质膜Plasma membrane
	RC2G0207300	Chr02：18414833 ~ 18417905	1 119	372	4	41 544.15	5.91	-0.258	细胞核Nucleus
	RC3G0144700	Chr03：22626525 ~ 22629894	1 149	382	4	43 154.29	6.43	-0.222	线粒体基质Mitochondrial matrix space
	RC3G0245900	Chr03：33482641 ~ 33485442	1 143	380	4	42 790.86	9.34	-0.362	细胞核Nucleus
	RC4G0305900	Chr04：43009598 ~ 43013252	1 155	384	4	42 277.11	6.31	-0.182	细胞核Nucleus
	RC6G0083700	Chr06：8901670 ~ 8903762	1 149	382	4	42 629.68	8.78	-0.288	细胞核Nucleus
	RC6G0295500	Chr06：39208249 ~ 39213326	1 188	395	4	43 907.85	8.45	-0.291	细胞核Nucleus
E	RC2G0570600	Chr02：65118745 ~ 65121601	1 413	470	5	52 006.45	5.45	-0.488	细胞核Nucleus
	RC3G0300800	Chr03：37901846 ~ 37903908	1 359	452	5	50 180.95	4.92	-0.529	细胞质Cytoplasm
	RC3G0340800	Chr03：41156572 ~ 41159167	1 125	374	5	41 586.10	5.84	-0.343	细胞核Nucleus
	RC3G0415800	Chr03：46406403 ~ 46410803	1 500	499	5	54 901.47	5.28	-0.451	叶绿体基质Chloroplast stroma
	RC4G0174800	Chr04：25897436 ~ 25902101	1 233	410	5	44 205.25	7.96	-0.160	细胞核Nucleus
	RC4G0290900	Chr04：41153285 ~ 41155207	1 164	387	5	42 900.06	8.19	-0.217	内质网Endoplasmic reticulum
	RC5G0637500	Chr05：74500818-74504788	1 602	533	5	58 727.27	5.24	-0.404	叶绿体基质Chloroplast stroma
	RC6G0585500	Chr06：65787874 ~ 65789999	1 014	337	4	36 815.15	4.67	-0.238	细胞质Cytoplasm
F	RC1G0561000	Chr01：62166916 ~ 62170959	1 050	349	8	38 095.69	5.30	-0.349	线粒体基质Mitochondrial matrix space
	RC5G0161600	Chr05：12241573 ~ 12242147	384	127	2	13 699.74	6.26	0.189	细胞质Cytoplasm
	RC5G0287900	Chr05：25199030 ~ 25202031	849	282	5	31 280.45	6.07	-0.424	细胞质Cytoplasm
	RC6G0448200	Chr06：55396868 ~ 55401178	879	292	8	31 419.82	5.04	-0.334	细胞质Cytoplasm
	RC7G0553900	Chr07：63513147 ~ 63518291	849	282	5	31 005.94	7.73	-0.491	线粒体基质Mitochondrial matrix space
G	RC2G0115900	Chr02：9573649 ~ 9576363	1 260	419	4	45 563.01	6.43	-0.214	细胞核Nucleus
	RC2G0023300	Chr02：1683170 ~ 1685525	1 038	345	4	38 186.16	5.23	-0.292	细胞质Cytoplasm
	RC3G0083700	Chr03：14633161 ~ 14635520	1 176	391	4	43 297.18	4.93	-0.175	过氧化物酶体Peroxisome
	RC4G0312700	Chr04：43671766 ~ 43674255	1 140	379	4	42 030.04	5.33	-0.015	内质网Endoplasmic reticulum
	RC6G0313800	Chr06：41654079 ~ 41656687	1 158	385	4	41 854.12	5.25	-0.280	细胞质Cytoplasm
	RC6G0447100	Chr06：55271914 ~ 55274566	1 134	377	4	41 279.55	5.40	-0.315	细胞质Cytoplasm
	RC7G0347300	Chr07：33066169 ~ 33069400	1 089	362	4	39 359.71	5.58	-0.226	细胞质Cytoplasm
H	RC3G0339900	Chr03：41087943 ~ 41091127	1 308	435	7	46 914.75	6.20	-0.114	内质网Endoplasmic reticulum
	RC4G0389000	Chr04：50185684 ~ 50188564	1 293	430	9	46 157.67	6.23	-0.137	细胞质Cytoplasm
	RC6G0560400	Chr06：63913434 ~ 63917402	1 293	430	9	46 572.74	5.06	-0.319	质膜Plasma membrane
I	RC2G0409900	Chr02：47147013 ~ 47154068	1 212	403	11	44 511.31	5.32	-0.302	内质网Endoplasmic reticulum
	RC6G0451300	Chr06：55557811 ~ 55562884	1 152	383	10	42 008.38	4.59	-0.225	质膜Plasma membrane
J	RC3G0144900	Chr03：22635737 ~ 22638365	1 434	477	7	52 805.38	5.43	-0.232	内质网Endoplasmic reticulum
	RC6G0396700	Chr06：50768308 ~ 50778537	3 144	1 047	22	118 157.21	5.74	-0.345	细胞质Cytoplasm
K	RC2G0161300	Chr02：13346491 ~ 13350340	1 320	439	4	46 451.30	7.64	-0.095	细胞质Cytoplasm
	RC5G0146400	Chr05：10980909 ~ 10981772	864	287	1	31 988.92	5.48	-0.538	细胞核Nucleus
	RC5G0168100	Chr05：12965254 ~ 12966114	861	286	1	31 063.42	5.33	-0.106	细胞质Cytoplasm
	RC5G0616200	Chr05：71710833 ~ 71713396	915	304	11	32 791.37	5.51	0.045	细胞核Nucleus
	RC6G0481800	Chr06：58085355 ~ 58088427	1 512	503	4	53 601.63	6.07	-0.277	线粒体基质Mitochondrial matrix space
	RC7G0438400	Chr07：47207142 ~ 47214206	1 866	621	7	66 851.18	4.77	-0.624	细胞质Cytoplasm
L	RC2G0063500	Chr02：4938066 ~ 4945505	3 249	1 082	15	119 968.55	5.10	-0.268	叶绿体类囊体膜Chloroplast thylakoid membrane
	RC4G0257100	Chr04：37263358 ~ 37266748	1 281	426	3	46 285.35	5.08	-0.413	叶绿体基质Chloroplast stroma
Others	RC1G0573600	Chr01：63257965 ~ 63261116	1 200	399	11	43 032.47	5.24	-0.136	过氧化物酶体Peroxisome
	RC4G0054600	Chr04：5904928 ~ 5907032	840	279	5	31 486.20	9.00	-0.428	过氧化物酶体Peroxisome
	RC4G0234300	Chr04：34375420 ~ 34381498	1 707	568	13	62 714.19	5.46	-0.268	内质网Endoplasmic reticulum
	RC5G0408300	Chr05：4088239 ~ 40883657	369	122	3	13 395.99	4.03	0.011	细胞质Cytoplasm
	RC6G0347300	Chr06：45436974 ~ 45440004	1 977	658	1	72 871.19	5.71	-0.167	过氧化物酶体Peroxisome
	RC7G0024700	Chr07：1516095 ~ 1519053	999	332	10	36 056.93	7.69	-0.269	细胞核Nucleus