中国春小麦肌醇磷脂依赖的磷脂酶C基因的全基因组鉴定及表达分析

doi:10.3864/j.issn.0578-1752.2020.24.001

摘要/Abstract

摘要：

【目的】探知小麦基因组序列中肌醇磷脂依赖的磷脂酶C（PLC）的编码基因,解析小麦中PLC基因的结构与进化特征,揭示TaPLC基因在小麦各组织中的表达模式及其响应盐胁迫和干旱胁迫过程中的表达规律,以便深入分析小麦TaPLC基因调节小麦应答盐或干旱胁迫中的生理作用。【方法】基于Ensembl Plants全基因组数据库,以水稻和拟南芥PLC基因为参考序列,检索小麦TaPLC基因家族（http://plants.ensembl.org/index.html）,并利用在线蛋白结构分析工具（Pfam、CDD和SMART）对TaPLC基因进行结构分析和鉴定;运用ExPASy（http://cn.expasy.org/tools）分析TaPLC基因的分子和生化特征;使用WoLF PSORT（https://www.genscript.com/wolf-psort.html）预测TaPLC基因的亚细胞定位;运用MEGA 7.0软件进行系统进化树分析;并利用实时荧光定量PCR分析TaPLC基因在不同组织以及幼苗期在盐或干旱胁迫下的表达特征。【结果】在小麦基因组中鉴定了11个TaPLC基因序列,其编码蛋白与其他植物PLC结构相似,均具有X和Y保守催化结构域以及C2结构域;系统进化分析表明11个TaPLC基因可分为4组,即TaPLC1A/TaPLC1D、TaPLC2A/TaPLC2B/TaPLC2D、TaPLC3A/TaPLC3B/TaPLC3D、TaPLC4A/TaPLC4B/TaPLC4D,均为植物PLC基因家族的直系同源基因,在进化方面具有保守性,但也存在一定的分歧;通过与小麦祖先种比对分析,发现B亚基因组中的TaPLC1在异源六倍体形成前就已经缺失,进一步表明小麦亚基因组间进化的不对称性;亚细胞定位预测TaPLC基因定位于叶绿体、线粒体或细胞质中;TaPLC基因家族各成员的启动子普遍存在植物激素响应和应激响应相关的顺式调控元件;实时定量结果表明,不同的TaPLC基因在不同的组织中有不同的表达水平,其主要在根、叶、穗和籽粒中表达;且多数基因参与小麦对盐和干旱胁迫的响应,无论是在耐旱和耐盐品种中,还是在对照品种中,TaPLC基因均受到胁迫的诱导表达,并在12 h内迅速达到峰值。【结论】在小麦基因组中共鉴定11个TaPLC基因,编码典型的植物PLC蛋白,在进化上具有保守性,但亚基因组间也存在进化的不对称性。不同的TaPLC基因有各自的组织和细胞表达特征,在小麦生长发育过程中起调节作用;TaPLC基因在应答盐或干旱胁迫中起重要作用。

关键词: 小麦, 肌醇磷脂依赖的磷脂酶C, 盐胁迫, 干旱胁迫

Abstract:

【Objective】To characterize the gene family encoding phosphatidylinositol-specific phospholipase-C (PI-PLC), the gene number, structure, phylogenetic relationship and expression pattern of TaPLC genes were analyzed. In addition, we explored their relative transcription levels in various tissue and the expression pattern under drought and salt stress to further analysis the physiological role of TaPLC genes in response to abiotic stress.【Method】Based on the genome database (Ensembl Plants, http://plants.ensembl.org/index.html), TaPLC genes were identified from Triticum aestivum, and the gene structures were analyzed by some bioinformatics tools, such as Pfam, CDD and SMART. The protein sequence characteristics of the members in TaPLC family were analyzed with the online server ExPASy (http://cn.expasy.org/tools). The subcellular localization of TaPLC genes were predicted using the WoLF PSORT (https://www.genscript.com/wolf-psort.html). Phylogenic tree analysis was performed by software MEGA 7.0. The real-time PCR was used to detect the gene expression levels in different tissues under abiotic stress. 【Result】A total of 11 TaPLC genes members within the heterologous hexaploid wheat genome were identified and analyzed. All TaPLC genes have conserved X-Y catalytic domains and C2 domains, just like other plant PLC genes. Subcellular localization showed that these proteins were located in the chloroplast or mitochondria. Phylogenetic analysis revealed that all of the TaPLC genes were unevenly classified into four numbered, TaPLC1A/TaPLC1D, TaPLC2A/TaPLC2B/TaPLC2D, TaPLC3A/TaPLC3B/TaPLC3D, and TaPLC4A/TaPLC4B/TaPLC4D, which were all orthologous genes of plant PLC genes in plants, and highly conserved during evolution. However, the deletion of TaPLC1 in B subgenome indicates asymmetry evolution in different subgenomes. Further analysis of cis-regulatory elements elucidated that these members of TaPLC family shared similar elements, such as the phytohormone and the stress response cis-elements. Each PLC gene has a unique tissue expression pattern, and they are expressed mainly in roots, leaves, spikes, and gains. In addition, PLC genes mediate the response of wheat to salt and drought stresses, which are induced rapidly by salt or drought stress stimulation. 【Conclusion】The TaPLC family in Triticum aestivum contains 11 members which have conserved X-Y catalytic domains and C2 domains. Although the TaPLC genes were highly conserved, they were asymmetrically evolved in different subgenomes. They had specific expression pattern in different tissues, and were induced by salt or drought stress, which suggested TaPLC genes play a role in response to salt or drought in wheat.

Key words: Triticum aestivum, PI-PLC, salt stress, drought stress

司旭阳,贾哓玮,张洪艳,贾羊羊,田士军,张科,潘延云. 中国春小麦肌醇磷脂依赖的磷脂酶C基因的全基因组鉴定及表达分析[J]. 中国农业科学, 2020, 53(24): 4969-4981.

SI XuYang,JIA XiaoWei,ZHANG HongYan,JIA YangYang,TIAN ShiJun,ZHANG Ke,PAN YanYun. Genomic Profiling and Expression Analysis of Phosphatidylinositol- specific PLC Gene Families Among Chinese Spring Wheat[J]. Scientia Agricultura Sinica, 2020, 53(24): 4969-4981.

图/表 9

表1

表2

图1

图2

图3

图4

图5

图6

图7

参考文献 43

[1]	GONG Z, XIONG L, SHI H, YANG S, HERRERA-ESTRELLA L R, XU G, CHAO D Y, LI J, WANG P Y, QIN F, LI J, DING Y, SHI Y, WANG Y, YANG Y, GAO Y, ZHU J K. Plant abiotic stress response and nutrient use efficiency. Science China Life Sciences, 2020,63(5):635-674. doi: 10.1007/s11427-020-1683-x pmid: 32246404
[2]	江成, 周厚君, 赵岩秋, 何辉, 楚立威, 宋学勤, 卢孟柱. 干旱和高盐胁迫下钙离子依赖核酸酶基因CDD对银腺杨84K生长发育的影响. 林业科学, 2019,55(2):33-40.
	JIANG C, ZHOU H J, ZHAO Y Q, HE H, CHU L W, SONG X Q, LU M Z. Effects of CDD gene on the growth and development of Populus alba× P. glandulosa ‘84K’ in response to drought and salt stresses. Scientia Silvae Sinicae, 2019,55(2):33-40. (in Chinese)
[3]	HOU Q, UFER G, BARTELS D. Lipid signalling in plant responses to abiotic stress. Plant, Cell and Environment, 2016,39(5):1029-1048. doi: 10.1111/pce.12666 pmid: 26510494
[4]	MUNNIK T, VERMEER J E. Osmotic stress-induced phosphoinositide and inositol phosphate signalling in plants. Plant, Cell and Environment, 2010,33(4):655-669. doi: 10.1111/j.1365-3040.2009.02097.x pmid: 20429089
[5]	TASMA I M, BRENDEL V, WHITHAM S A, BHATTACHARYYA M K. Expression and evolution of the phosphoinositide-specific phospholipase C gene family inArabidopsis thaliana. Plant Physiology and Biochemistry, 46(7):627-637. doi: 10.1016/j.plaphy.2008.04.015 pmid: 18534862
[6]	SHI J, GONZALES R A, BHATTACHARYYA M K. Characterization of a plasma membrane-associated phosphoinositide-specific phospholipase C from soybean. The Plant Journal, 1995,8(3):381-390. doi: 10.1046/j.1365-313x.1995.08030381.x pmid: 7550376
[7]	PICAL C, KOPKA J, MULLER R B, HETHERINGTON A M, GRAY J E. Isolation of 2 cDNA clones for phosphoinositide-specific phospholipase C from epidermal peels (accession No. X95877) and guard cells (accession No. Y11931) of Nicotiana rustica (PGR 97-086). Plant Physiology, 1997,114:747-749. doi: 10.1104/pp.114.2.747 pmid: 9235602
[8]	TRIPATHY M K, TYAGI W, GOSWAMI M, KAUL T, SINGLA-PAREEK S L, DESWAL R, REDDY M K, SOPORY S K. Characterization and functional validation of tobacco PLC delta for abiotic stress tolerance. Plant Molecular Biology Reporter, 2012,30(2):488-497. doi: 10.1007/s11105-011-0360-z
[9]	LI L, WANG F, YAN P, JING W, ZHANG C, KUDLA J, ZHANG W. A phosphoinositide-specific phospholipase C pathway elicits stress-induced Ca²⁺ signals and confers salt tolerance to rice . New Phytologist, 2017,214(3):1172-1187. doi: 10.1111/nph.14426
[10]	KOPKA J, PICAL C, GRAY J E, MULLER-ROBER B. Molecular and enzymatic characterization of three phosphoinositide-specific phospholipase C isoforms from potato. Plant Physiology, 1998,116:239-250. doi: 10.1104/pp.116.1.239 pmid: 9449844
[11]	WANG C R, YANG A F, YUE G D, GAO Q, YIN H Y, ZHANG J R. Enhanced expression of phospholipase C 1 (ZmPLC1) improves drought tolerance in transgenic maize. Planta, 2008,227(5):1127-1140. doi: 10.1007/s00425-007-0686-9
[12]	PAN Y Y, WANG X, MA L G, SUN D Y. Characterization of phosphatidylinositol-specific phospholipase C (PI-PLC) from Lilium daviddi pollen. Plant and Cell Physiology, 2005,46(10):1657-1665. doi: 10.1093/pcp/pci181 pmid: 16085656
[13]	KIM Y J, KIM J E, LEE J H, LEE M H, JUNG H W, BAHK Y Y, HWANG B K, HWANG I, KIM W T. The Vr-PLC3 gene encodes a putative plasma membrane-localized phosphoinositide-specific phospholipase C whose expression is induced by abiotic stress in mung bean (Vigna radiata L.). FEBS Letters, 2004,556(1):127-136. doi: 10.1016/S0014-5793(03)01388-7
[14]	GNANARAJ M, UDHAYAKUMAR N, RAJIV GR, MANOHARAN K. Isolation and gene expression analysis of Phospholipase C in response to abiotic stresses from Vigna radiata (L.) Wilczek. Indian Journal of Experimental Biology, 2015,53(6):335-341. pmid: 26155672
[15]	CAO Z, ZHANG J, LI Y, XU X, LIU G, BHATTACHARRYA M K, YANG H, REN D. Preparation of polyclonal antibody specific for AtPLC4, an Arabidopsis phosphatidylinositol-specific phospholipase C in rabbits. Protein Expression and Purification, 2007,52(2):306-312. doi: 10.1016/j.pep.2006.10.007
[16]	XIA K, WANG B, ZHANG J, LI Y, YANG H, REN D. Arabidopsis phosphoinositide-specific phospholipase C 4 negatively regulates seedling salt tolerance. Plant, Cell and Environment, 2017,40(8):1317-1331. doi: 10.1111/pce.12918 pmid: 28102910
[17]	KANEHARA K, YU C Y, CHO Y, CHEONG W F, TORTA F, SHUI G, WENK M R, NAKAMURA Y. Arabidopsis AtPLC2 is a primary phosphoinositide-specific phospholipase C in phosphoinositide metabolism and the endoplasmic reticulum stress response. PLoS Genetics, 2015,11(9):e1005511. doi: 10.1371/journal.pgen.1005511 pmid: 26401841
[18]	LI L, HE Y, WANG Y, ZHAO S, CHEN X, YE T, WU Y, WU Y. Arabidopsis PLC2 is involved in auxin-modulated reproductive development. The Plant Journal: for Cell and Molecular Biology, 2015,84(3):504-215. doi: 10.1111/tpj.2015.84.issue-3
[19]	ZHANG Q, VAN WIJK R, ZARZA X, SHAHBAZ M, VAN HOOREN M, GUARDIA A, SCUFFI D, GARCÍA-MATA C, VAN DEN ENDE W, HOFFMANN-BENNING S, HARING M A, LAXALT A M, MUNNIK T. Knock-down of Arabidopsis PLC5 reduces primary root growth and secondary root formation while overexpression improves drought tolerance and causes stunted root hair growth. Plant and Cell Physiology, 2018,59(10):2004-2019. doi: 10.1093/pcp/pcy120 pmid: 30107538
[20]	ZHANG Q, VAN WIJK R, SHAHBAZ M, ROELS W, SCHOOTEN B V, VERMEER J E M, ZARZA X, GUARDIA A, SCUFFI D, GARCÍA-MATA C, LAHA D, WILLIAMS P, WILLEMS L A J, LIGTERINK W, HOFFMANN-BENNING S, GILLASPY G, SCHAAF G, HARING M A, LAXALT A M, MUNNIK T. Arabidopsis phospholipase C3 is involved in lateral root initiation and ABA responses in seed germination and stomatal closure. Plant and Cell Physiology, 2018,59(3):469-486. doi: 10.1093/pcp/pcx194 pmid: 29309666
[21]	VAN WIJK R, ZHANG Q, ZARZA X, LAMERS M, MARQUEZ F R, GUARDIA A, SCUFFI D, GARCÍA-MATA C, LIGTERINK W, HARING M A, LAXALT A M, MUNNIK T. Role for Arabidopsis PLC7 in stomatal movement, seed mucilage attachment, and leaf serration. Frontiers in Plant Science, 2018,9:1721. doi: 10.3389/fpls.2018.01721 pmid: 30542361
[22]	SINGH A, KANWAR P, PANDEY A, TYAGI A K, SOPORY S K, KAPOOR S, PANDEY G K. Comprehensive genomic analysis and expression profiling of phospholipase C gene family during abiotic stresses and development in rice. PLoS ONE, 2013,8(4):e62494. doi: 10.1371/journal.pone.0062494 pmid: 23638098
[23]	SINGH A, BHATNAGAR N, PANDEY A, PANDEY G K. Plant phospholipase C family: Regulation and functional role in lipid signaling. Cell Calcium, 2015,58(2):139-146. doi: 10.1016/j.ceca.2015.04.003 pmid: 25933832
[24]	POKOTYLO I, KOLESNIKOV Y, KRAVETS V, ZACHOWSKI A, RUELLAND E. Plant phosphoinositide-dependent phospholipases C: Variations around a canonical theme. Biochimie, 2014,96:144-157. doi: 10.1016/j.biochi.2013.07.004 pmid: 23856562
[25]	MELIN P M, SOMMARIN M, SANDELIUS A S, JERGIL B. Identification of Ca²⁺-stimulated polyphosphoinositide phospholipase C in isolated plant plasma membranes . FEBS Letters, 1987,223(1):87-91. doi: 10.1016/0014-5793(87)80515-x pmid: 2822482
[26]	MELIN P M, PICAL C, JERGIL B, SOMMARIN M. Polyphosphoinositide phospholipase C in wheat root plasma membranes, partial purification and characterization. Biochimica et Biophysica Acta, 1992,1123(2):163-169. doi: 10.1016/0005-2760(92)90107-7 pmid: 1310875
[27]	PICAL C, SANDELIUS A S, MELIN P M, SOMMARIN M. Polyphosphoinositide phospholipase C in plasma membranes of wheat (Triticum aestivum L.): Orientation of active site and activation by Ca and Mg. Plant Physiology, 1992,100(3):1296-1303. doi: 10.1104/pp.100.3.1296 pmid: 16653120
[28]	ZHANG K, JIN C, WU L, HOU M, DOU S, PAN Y. Expression analysis of a stress-related phosphoinositide-specific phospholipase C gene in wheat (Triticum aestivum L.). PLoS ONE, 2014,9(8):e105061. doi: 10.1371/journal.pone.0105061 pmid: 25121594
[29]	吴少辉, 张学品, 杨洪强, 冯伟森. GS旱地小麦新品种-洛旱7号. 中国农业科技导报, 2009,11(S2):118-120.
	WU S H, ZHANG X P, YANG H Q, FENG W S. GS dryland wheat new variety-luohan 7. Journal of Agricultural Science and Technology, 2009,11(S2):118-120. (in Chinese)
[30]	张明明, 董宝娣, 乔匀周, 赵欢, 刘孟雨, 陈骎骎, 杨红, 郑鑫. 播期、播量对旱作小麦‘小偃60’生长发育、产量及水分利用的影响. 中国生态农业学报, 2016,24(8):1095-1102.
	ZHANG M M, DONG B D, QIAO Y Z, ZHAO H, LIU M Y, CHEN Q Q, YANG H, ZHEN X. Effects of sowing date and seeding density on growth, yield and water use efficiency of ‘Xiaoyan 60’ wheat under rainfed condition. Chinese Journal of Eco-Agriculture, 2016,24(8):1095-1102. (in Chinese)
[31]	TASMA I M, BRENDEL V, WHITHAM S A, BHATTACHARYYA M K. Expression and evolution of the phosphoinositide-specific phospholipase C gene family in Arabidopsis thaliana. Plant Physiology and Biochemistry, 2008,46(7):627-637. doi: 10.1016/j.plaphy.2008.04.015
[32]	SINGH A, KANWAR P, PANDEY A, TYAGI A K, SOPORY S K, KAPOOR S, PANDEY G K. Comprehensive genomic analysis and expression profiling of phospholipase C gene family during abiotic stresses and development in rice. PLoS ONE, 2013,8(4):e62494. doi: 10.1371/journal.pone.0062494 pmid: 23638098
[33]	VOSSEN J H, ABD-EL-HALIEM A, FRADIN E F, VAN DEN BERG G C M, EKENGREN S K, MEIJER H J G, SEIFI A, BAI Y, TEN HAVE A, MUNNIK T, THOMMA B P H J, JOOSTEN M H A J. Identification of tomato phosphatidylinositol-specific phospholipase- C (PI-PLC) family members and the role of PLC4 and PLC6 in HR and disease resistance. The Plant Journal: for Cell and Molecular Biology, 2010,62(2):224-239. doi: 10.1111/j.1365-313X.2010.04136.x
[34]	KHALIL H B, WANG Z, WRIGHT J A, RALEVSKI A, DONAYO A O, GULICK P J. Heterotrimeric Gα subunit from wheat (Triticum aestivum), GA3, interacts with the calcium-binding protein, Clo3, and the phosphoinositide-specific phospholipase C, PI-PLC1. Plant Molecular Biology, 2011,77(1/2):145-158. doi: 10.1007/s11103-011-9801-1
[35]	MORRAN S, EINI O, PYVOVARENKO T, PARENT B, SINGH R, ISMAGUL A, ELIBY S, SHIRLEY N, LANGRIDGE P, LOPATO S. Improvement of stress tolerance of wheat and barley by modulation of expression of DREB/CBF factors. Plant Biotechnology Journal, 2011,9(2):230-249. doi: 10.1111/j.1467-7652.2010.00547.x
[36]	HIRAYAMA T, OHTO C, MIZOGUCHI T, SHINOZAKI K. A gene encoding a phosphatidylinositol-specific phospholipase C is induced by dehydration and salt stress in Arabidopsis thaliana. Proceedings of the National Academy of Sciences of the United States of America, 1995,92(9):3903-3907. doi: 10.1073/pnas.92.9.3903 pmid: 7732004
[37]	LIN W H, YE R, MA H, XU Z H, XUE H W. DNA chip-based expression profile analysis indicates involvement of the phosphatidylinositol signaling pathway in multiple plant responses to hormone and abiotic treatments. Cell Research, 2004,14(1):34-45. doi: 10.1038/sj.cr.7290200 pmid: 15040888
[38]	TASMA I M, BRENDEL V, WHITHAM S A, BHATTACHARYYA M K. Expression and evolution of the phosphoinsitide-specific phospholipase C gene family in Arabidopsis thaliana. Plant Physiology and Biochemistry, 2008,46(7):627-637. doi: 10.1016/j.plaphy.2008.04.015
[39]	ZHAI S M, SUI Z H, YANG A F, ZHANG J R. Characterization of a novel phosphoinositide-speciWc phospholipase C from Zea mays and its expression in Escherichia coli. Biotechnology Letters, 2005,27(11):799-804. doi: 10.1007/s10529-005-5802-y
[40]	DARWISH E, TESTERINK C, KHALIL M, EL-SHIHY O, MUNNIK T. Phospholipid signaling responses in salt-stressed rice leaves. Plant Cell Physiology, 2009,50(5):986-997. doi: 10.1093/pcp/pcp051 pmid: 19369274
[41]	邓先俊. 水稻磷脂酰肌醇特异性磷脂酶C4(OsPLC4)调节水稻渗透胁迫响应及生长发育[D]. 武汉: 华中农业大学, 2018.
	DENG X J. Phosphatidylinositol-specific phosphatidylase C 4 (OsPLC4) regulates osmotic stress response and growth and development in rice[D]. Wuhan: Huazhong Agricultural University, 2018. (in Chinese)
[42]	GEORGES F, DAS S, RAY H, BOCK C, NOKHRINA K, KOLLA A, KELLER W. Over-expression of Brassica napus phosphatidylinositol- phospholipase C2 in canola induces significant changes in gene expression and phytohormone distribution patterns, enhances drought tolerance and promotes early flowering and maturation. Plant, Cell and Environment, 2009,32(12):1664-1681. doi: 10.1111/j.1365-3040.2009.02027.x pmid: 19671099
[43]	杜康兮, 沈文辉, 董爱武. 表观遗传调控植物响应非生物胁迫的研究进展. 植物学报, 2018,53(5):581-593. doi: 10.11983/CBB17143
	DU K X, SHEN W H, DONG A W. Advances in epigenetic regulation of abiotic stress response in plants. Chinese Bulletin of Botany, 2018,53(5):581-593. (in Chinese) doi: 10.11983/CBB17143

基因名称 Gene name	基因ID Gene ID	染色体定位^a Chromosomal location	开放阅读框 Open reading frame (bp)	蛋白质^b Protein
基因名称 Gene name	基因ID Gene ID	染色体定位^a Chromosomal location	开放阅读框 Open reading frame (bp)	大小 Size (aa)	分子量 MW (kD)	等电点 pI
TaPLC1A	TraesCS1A02G069300	1A:51700021—51707185	1758	585	66.01	6.07
TaPLC1D	TraesCS1D02G071800	1D:52338417—52345805	1761	586	66.19	6.03
TaPLC2A	TraesCS2A02G084000	2A:38534860—38538480	1830	609	68.5	5.91
TaPLC2B	TraesCS2B02G098500	2B:58321242—58325230	1821	606	68.22	6.06
TaPLC2D	TraesCS2D02G082000	2D:35259471—35263395	1824	607	68.45	5.88
TaPLC3A	TraesCS4A02G109000	4A:129086595—129090166	1902	633	71.11	5.54
TaPLC3B	TraesCS4B02G195200	4B:420197892—420201623	1902	633	71.00	5.75
TaPLC3D	TraesCS4D02G195800	4D:340085748—340090648	1902	633	71.05	5.84
TaPLC4A	TraesCS5A02G155300	5A:333407514—333413175	1773	590	65.73	6.05
TaPLC4B	TraesCS5B02G153600	5B:283008744—283014326	1770	589	65.65	6.06
TaPLC4D	TraesCS5D02G160300	5D:250061407—250066699	1770	589	65.65	6.06

基因Gene	亚细胞定位Subcellular localization
TaPLC1A	线粒体：8,叶绿体：4,细胞核：1 Mito: 8, chlo: 4, nucl: 1
TaPLC1D	线粒体：8,叶绿体：4,细胞核：1 Mito: 8, chlo: 4, nucl: 1
TaPLC2A	叶绿体：7,细胞核：3,线粒体：2.5,线粒体基质：2 Chlo: 7, nucl: 3, mito: 2.5, cyto_mito: 2
TaPLC2B	叶绿体：6,线粒体：5.5,线粒体基质：3.5,细胞核：1 Chlo: 6, mito: 5.5, cyto_mito: 3.5, nucl: 1
TaPLC2D	叶绿体：7,线粒体：4.5,线粒体基质：3,细胞核：1 Chlo: 7, mito: 4.5, cyto_mito: 3, nucl: 1
TaPLC3A	细胞质：8,细胞核：4,叶绿体：1 Cyto: 8, nucl: 4, chlo: 1
TaPLC3B	细胞质：8,细胞核：3,叶绿体：1,线粒体：1 Cyto: 8, nucl: 3, chlo: 1, mito: 1
TaPLC3D	细胞质：6,细胞核：3,线粒体：2,过氧化物：2 Cyto: 6, nucl: 3, mito: 2, pero: 2
TaPLC4A	线粒体：10,叶绿体和线粒体：6.83333,线粒体基质：5.83333,叶绿体：2.5 Mito: 10, chlo_mito: 6.83333, cyto_mito: 5.83333, chlo: 2.5
TaPLC4B	线粒体：10,叶绿体和线粒体：6.83333,线粒体基质：5.83333,叶绿体：2.5 Mito: 10, chlo_mito: 6.83333, cyto_mito: 5.83333, chlo: 2.5
TaPLC4D	线粒体：10,叶绿体和线粒体：6.83333,线粒体基质：5.83333,叶绿体：2.5 Mito: 10, chlo_mito: 6.83333, cyto_mito: 5.83333, chlo: 2.5