植物表皮蜡质合成及调控因子WIN/SHN的研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2020-0572

摘要/Abstract

摘要：

植物表皮蜡质是植物与外界环境直接相接触的一层疏水保护层,由于它能够调节植物体内的水分应对干旱、盐和冷等非生物胁迫,因此对植物的生长和发育起着重要的作用。在不同的物种之间,植物表皮蜡质的结构和组成均有着很大差异,但都具有类似的蜡质合成途径,对该途径的研究已经取得了显著的成果。多种转录因子参与了植物的蜡质生物合成,其中隶属于AP2转录因子家族中的WIN/SHN通过与某些基因的启动子结合,调节蜡质合成相关基因的表达,从而影响蜡质的合成。在植物表皮蜡质合成的基础之上,重点对调节蜡质形成的转录因子WIN/SHN的结构特点、表达模式、转录调节作用及其对生理和表型的影响,以及对逆境的应答反应进行了综述,并对蜡质合成的复杂性问题进行了论述。鉴于蜡质对生命科学和农业生产实践的重要价值,深入开展包括WIN/SHN在内的遗传因子及环境因子对蜡质合成、调控与运转的机制研究将具有重要意义。

关键词: 蜡质, 生物合成, 蜡质诱导因子WIN/SHN, 转录调控

Abstract:

Plant cuticular wax is a hydrophobic protective layer by which plants directly contact with the external environment. Because it can regulate the water in plants to deal with abiotic stresses such as drought,salt and cold,it plays an important role in plant growth and development. Among different species,the structure and composition of the plant cuticular wax vary largely,but all have similar wax synthesis pathways,and remarkable research results on these pathways has achieved. Many transcription factors（TFs）are involved in the wax biosynthesis of plants. WIN/SHN,which belongs to the AP2 TF family,regulates the expressions of wax synthesis-related genes by binding to their promoters,thereby affecting the waxy synthesis. Based on the synthesis of plant cuticular wax,this article focuses on the review of WIN/SHN TFs,including their structural characteristics,expression patterns,transcriptional regulation and the effects on physiology and phenotype,as well as their responses to stresses. Furthermore,the complexity of wax synthesis is also discussed. In view of the important value of wax for life science and agricultural production practice,it is of great significance to carry out in-depth research on the mechanism of wax synthesis,regulation,transport and deposition regulated by the genetic and environmental factors including WIN/SHN TFs.

Key words: wax, biosynthesis, wax induction factor WIN/SHN, transcriptional regulation

李晓佩, 王思宁, 史晶晶, 高志民. 植物表皮蜡质合成及调控因子WIN/SHN的研究进展[J]. 生物技术通报, 2020, 36(12): 129-136.

LI Xiao-pei, WANG Si-ning, SHI Jing-jing, GAO Zhi-min. Progress of Plant Cuticular Wax Synthesis and Its Regulatory Factor WIN/SHN[J]. Biotechnology Bulletin, 2020, 36(12): 129-136.

参考文献 56

[1]	Yeats TH, Rose JKC. The formation and function of plant cuticles[J]. Plant Physiology, 2013,163(1):5-20. doi: 10.1104/pp.113.222737 URL pmid: 23893170
[2]	Bernard A, Joubes J. Arabidopsis cuticular waxes:advances in synjournal, export and regulation[J]. Progress in Lipid Research, 2013,52(1):110-129. doi: 10.1016/j.plipres.2012.10.002 URL pmid: 23103356
[3]	Borisjuk N, Hrmova M, Lopato S. Transcriptional regulation of cuticle biosynjournal[J]. Biotechnology Advances, 2014,32(2):526-540. URL pmid: 24486292
[4]	Nawrath C. Unraveling the complex network of cuticular structure and function[J]. Current Opinion in Plant Biology, 2006,9(3):281-287. doi: 10.1016/j.pbi.2006.03.001 URL
[5]	李灵之, 马杰, 向建华, 等. 植物角质层内外蜡质的差异及其与抗逆性的关系[J]. 植物生理学报, 2011,47(7):680-684.
	Li LZ, Ma J, Xiang JH, et al. Composition differences of epicuticular and intracuticular wax layers and the relationship between cuticle and plant stress tolerance[J]. Plant Physiology Journal, 2011,47(7):680-684.
[6]	Eglinton G, Hamilton RJ. Leaf epicuticular waxes[J]. Science, 1967,156(3780):1322-1335. doi: 10.1126/science.156.3780.1322 URL pmid: 4975474
[7]	Barthlott W, Neinhuis C, Cutler D, et al. Classification and terminology of plant epicuticular waxes[J]. Botanical Journal of the Linnean Society, 1998,126(3):237-260. doi: 10.1111/boj.1998.126.issue-3 URL
[8]	Kunst L, Samuels AL. Biosynjournal and secretion of plant cuticular wax[J]. Progress in Lipid Research, 2003,42(1):51-80. doi: 10.1016/s0163-7827(02)00045-0 URL pmid: 12467640
[9]	Zeng Q, Liu DC, Liu Y. The overview and prospect of chemical composition of plant cuticular wax[J]. Acta Ecologica Sinica, 2013,33(17):5133-5140.
[10]	Lee , SH , Stephens JL, Englund PT. A fatty-acid synjournal mechanism specialized for parasitism[J]. Nature Reviews Microbiology, 2007,5(4):287-297. doi: 10.1038/nrmicro1617 URL pmid: 17363967
[11]	Li-Beisson Y, Shorrosh B, Beisson F, et al. Acyl-lipid metabolism[J]. The Arabidopsis Book, 2010,8:e0133. URL pmid: 22303259
[12]	Nawrath C, Schreiber L, Franke RB, et al. Apoplastic diffusion barriers in Arabidopsis[J]. The Arabidopsis Book, 2013,11:e0167. URL pmid: 24465172
[13]	Cheesbrough TM, Kolattukudy PE. Alkane biosynthesis by decarbonylation of aldehydes catalyzed by a particulate preparation from Pisum sativum[J]. Proceedings of the National Academy of Science of the United States of America, 1984,81(21):6613-6617.
[14]	Metz JG, Pollard MR, Anderson L, et al. Purification of a jojoba embryo fatty acyl-coenzyme A reductase and expression of its cDNA in high erucic acid rapeseed[J]. Plant Physiology, 2000,122(3):635-644. doi: 10.1104/pp.122.3.635 URL pmid: 10712526
[15]	Li F, Wu X, Lam P, et al. Identification of the wax ester synthase/acyl-coenzyme A:diacylglycerol acyltransferase WSD1 required for stem wax ester biosynjournal in Arabidopsis[J]. Plant Physiology, 2008,148(1):97-107. doi: 10.1104/pp.108.123471 URL pmid: 18621978
[16]	Kunst L, Samuels L. Plant cuticles shine:advances in wax biosynjournal and export[J]. Current Opinion in Plant Biology, 2009,12(6):721-727. doi: 10.1016/j.pbi.2009.09.009 URL pmid: 19864175
[17]	Lü SY, Zhao HY, Marais DL, et al. Arabidopsis ECERIFERUM9 involvement in cuticle formation and maintenance of plant water status[J]. Plant Physiology, 2012,159(3):930-944. doi: 10.1104/pp.112.198697 URL pmid: 22635115
[18]	Jenks MA, Rashotte AM, Tuttle HA, et al. Mutants in Arabidopsis thaliana altered in epicuticular wax and leaf morphology[J]. Plant Physiology, 1996,110(2):377-385. doi: 10.1104/pp.110.2.377 URL pmid: 12226189
[19]	Zhang D, Yang H, Wang X, et al. Cytochrome P450 family member CYP96B5 hydroxylates alkanes to primary alcohols and is involved in rice leaf cuticular wax synjournal[J]. New Phytologist, 2020,225(5):2094-2107.
[20]	Todaka D, Nakashima K, Shinozaki K, et al. Toward understanding transcriptional regulatory networks in abiotic stress responses and tolerance in rice[J]. Rice, 2012,5(1):6-13. URL pmid: 24764506
[21]	Licausi F, Ohme-Takagi M, Perata P. APETALA2/Ethylene Responsive Factor(AP2/ERF)transcription factors:mediators of stress responses and developmental programs[J]. New Phytologist, 2013,199(3):639-649. doi: 10.1111/nph.12291 URL
[22]	Nakashima K, Yamaguchi-Shinozaki K, Shinozaki K. The transcriptional regulatory network in the drought response and its crosstalk in abiotic stress responses including drought, cold, and heat[J]. Frontiers in Plant Science, 2014,5:170. doi: 10.3389/fpls.2014.00170 URL pmid: 24904597
[23]	Aharoni A, Dixit S, Jetter R, et al. The SHINE clade of AP2 domain transcription factors activates wax biosynjournal, alters cuticle properties, and confers drought tolerance when overexpressed in Arabidopsis[J]. The Plant Cell, 2004,16(9):2463-2480.
[24]	Broun P, Poindexter P, Osborne E, et al. WIN1, a transcriptional activator of epidermal wax accumulation in Arabidopsis[J]. Proceedings of the National Academy of Sciences of the United States of America, 2004,101(13):4706-4711. URL pmid: 15070782
[25]	Sakuma Y, Liu Q, Dubouzet JG, et al. DNA-binding specificity of the ERF/AP2 domain of Arabidopsis DREBs, transcription factors involved in dehydration- and cold inducible gene expression[J]. Biochemical and Biophysical Research Communication, 2002,290(3):998-1009.
[26]	Nakano T, Suzuki K, Fujimura T, et al. Genome-wide analysis of the ERF gene family in Arabidopsis and rice[J]. Plant Physiology, 2006,140(2):411-432. URL pmid: 16407444
[27]	Wang Y, Wan L, Zhang L, et al. An ethylene response factor OsWR1 responsive to drought stress transcriptionally activates wax synjournal related genes and increases wax production in rice[J]. Plant Molecular Biology, 2012,78(3):275-288. doi: 10.1007/s11103-011-9861-2 URL pmid: 22130861
[28]	Xu Y, Wu H, Zhao M, et al. Overexpression of the transcription factors GmSHN1 and GmSHN9 differentially regulates wax and cutin biosynjournal, alters cuticle properties, and changes leaf phenotypes in Arabidopsis[J]. International Journal of Molecular Sciences, 2016,17(4):587.
[29]	Al-Abdallat A, Al-Debei H, Ayad J, et al. Over expression of SlSHN1 gene improves drought tolerance by increasing cuticular wax accumulation in tomato[J]. International Journal of Molecular Sciences, 2014,15(11):19499-19515. doi: 10.3390/ijms151119499 URL pmid: 25350113
[30]	Shi JX, Adato A, Alkan N, et al. The tomato SlSHINE3 transcription factor regulates fruit cuticle formation and epidermal patterning[J]. New Phytologist, 2013,197(2):468-480. doi: 10.1111/nph.12032 URL
[31]	Craft J, Samalova M, Baroux C, et al. New pOp/LhG4 vectors for stringent glucocorticoid-dependent transgene expression in Arabidopsis[J]. The Plant Journal, 2005,41(6):899-918. URL pmid: 15743453
[32]	Taketa S, Amano S, Tsujino Y, et al. Barley grain with adhering hulls is controlled by an ERF family transcription factor gene regulating a lipid biosynthesis pathway[J]. Proceedings of the National Academy of Sciences of the United States of America, 2008,105(10):4062-4067. doi: 10.1073/pnas.0711034105 URL pmid: 18316719
[33]	李铖, 潘健, 连红莉, 等. 黄瓜蜡质合成调控基因CsWIN1的克隆与功能初步分析[J]. 园艺学报, 2018,45(2):359-370.
	Li C, Pan J, Lian HL, et al. Cloning and functional analysis of CsWIN1, a transcription factor regulated the wax synjournal in cucumber[J]. Acta Horticulturae Sinica, 2018,45(2):359-370.
[34]	Zhou X, Jenks MA, Liu J, et al. Overexpression of transcription factor OsWR2 regulates wax and cutin biosynjournal in rice and enhances its tolerance to water deficit[J]. Plant Molecular Biology Reporter, 2014,32(3):719-731. doi: 10.1007/s11105-013-0687-8 URL
[35]	Isaacson T, Kosma DK, Matas AJ, et al. Cutin deficiency in the tomato fruit cuticle consistently affects resistance to microbial infection and biomechanical properties, but not transpirational water loss[J]. The Plant Journal, 2009,60(2):363-377.
[36]	Lashbrooke J, Aharoni A, Costa F. Genome investigation suggests MdSHN3, an APETALA2-domain transcription factor gene, to be a positive regulator of apple fruit cuticle formation and an inhibitor of russet development[J]. Journal of Experimental Botany, 2015,66(21):6579-6589. URL pmid: 26220084
[37]	Zhang YL, Zhang CL, Wang GL, et al. Apple AP2/EREBP transcription factor MdSHINE2 confers drought resistance by regulating wax biosynjournal[J]. Planta, 2019,249(5):1627-1643. URL pmid: 30826884
[38]	Djemal R, Mila I, Bouzayen M, et al. Molecular cloning and characterization of novel WIN1/SHN1 ethylene responsive transcription factor HvSHN1 in barley(Hordeum vulgare L.)[J]. Journal of Plant Physiology, 2018,228:39-46. URL pmid: 29852333
[39]	Kumar A, Yogendra KN, Karre S, et al. WAX INDUCER1(HvWIN1)transcription factor regulates free fatty acid biosynthetic genes to reinforce cuticle to resist Fusarium head blight in barley spikelets[J]. Journal of Experimental Botany, 2016,67(14):4127-4139. URL pmid: 27194736
[40]	陈伟, 刘德春, 杨莉, 等. 植物表皮蜡质及相关基因研究进展[J]. 植物生理学报, 2016,52(8):1117-1127.
	Chen W, Liu DC, Yang L, et al. Research progress of plant cuticular wax and related genes[J]. Plant Physiology Journal, 2016,52(8):1117-1127.
[41]	Lisso J, Schröder F, Schippers JHM, et al. NFXL2 modifies cuticle properties in Arabidopsis[J]. Plant Signaling & Behavior. 2012,7(5):551-555. URL pmid: 22516817
[42]	Qin F, Kodaira KS, Maruyama K, et al. SPINDLY, a negative regulator of gibberellic acid signaling, is involved in the plant abiotic stress response[J]. Plant Physiology, 2011,157(4):1900-1913. URL pmid: 22013217
[43]	Kannangara R, Branigan C, Liu Y, et al. The transcription factor WIN1/SHN1 regulates cutin biosynjournal in Arabidopsis thaliana[J]. The Plant Cell, 2007,19(4):1278-1294. URL pmid: 17449808
[44]	Karaba A. Improvement of water use efficiency in rice and tomato using Arabidopsis wax biosynthetic genes and transcription factors[D]. Wageningen:Wageningen University, 2007.
[45]	Djemal R, Khoudi H. Isolation and molecular characterization of a novel WIN1/SHN1 ethylene-responsive transcription factor TdSHN1 from durum wheat(Triticum turgidum. L. subsp. durum)[J]. Protoplasma, 2015,252(6):1461-1473. doi: 10.1007/s00709-015-0775-8 URL pmid: 25687296
[46]	王立山, 丁兵, 李玉花, 等. 植物表皮蜡质合成转运调控相关基因与干旱响应的研究进展[J]. 园艺学报, 2018,45(9):1831-1843.
	Wang LS, Ding B, Li YH, et al. Reaserch progress of plant cuticular wax biosynjournal, export and regulation related genes responsed to drought[J]. Acta Horticulturae Sinica, 2018,45(9):1831-1843.
[47]	Oshima Y, Shikata M, Koyama T, et al. MIXTA-like transcription factors and WAX INDUCER1/SHINE1 coordinately regulate cuticle development in Arabidopsis and Torenia fournieri[J]. The Plant Cell, 2013,25(5):1609-1624. URL pmid: 23709630
[48]	Ambavaram MMR, Krishnan A, Trijatmiko KR, et al. Coordinated activation of cellulose and repression of lignin biosynjournal pathways in rice[J]. Plant Physiology, 2011,155(2):916-931. URL pmid: 21205614
[49]	Bondada BR, Oosterhuis DM, Murphy JB, et al. Effect of water stress on the epicuticular wax composition and ultrastructure of cotton(Gossypium hirsutum L.)leaf, bract, and boll[J]. Environmental and Experimental Botany, 1996,36(1):61-69.
[50]	Samdur MY, Manivel P, Jain VK, et al. Genotypic differences and water-deficit induced enhancement in epicuticular wax load in peanut[J]. Crop Science, 2003,43(4):1294-1299.
[51]	Cameron KD, Teece MA, Smart LB. Increased accumulation of cuticular wax and expression of lipid transfer protein in response to periodic drying events in leaves of tree tobacco[J]. Plant Physiology, 2006,140(1):176-183. URL pmid: 16361524
[52]	Kosma DK, Bourdenx B, Bernard A, et al. The impact of water deficiency on leaf cuticle lipids of Arabidopsis[J]. Plant Physiology, 2009,151(4):1918-1929. doi: 10.1104/pp.109.141911 URL pmid: 19819982
[53]	Seo PJ, Lee SB, Suh MC, et al. The MYB96 transcription factor regulates cuticular wax biosynjournal under drought conditions in Arabidopsis[J]. The Plant Cell, 2011,23(3):1138-1152. URL pmid: 21398568
[54]	Yang J, Zhao X, Liang L, et al. Overexpression of a cuticle-degrading protease Ver112 increases the nematicidal activity of Paecilomyces lilacinus[J]. Applied Microbiology and Biotechnology, 2011,89(6):1895-1903. URL pmid: 21110018
[55]	Amid A, Lytovchenko A, Fernie AR, et al. The sensitive to freezing3 mutation of Arabidopsis thaliana is a cold-sensitive allele of homomeric acetyl-CoA carboxylase that results in cold-induced cuticle deficiencies[J]. Journal of Experimental Botany, 2012,63(14):5289-5299. URL pmid: 22791831
[56]	Uppalapati SR, Ishiga Y, Doraiswamy V, et al. Loss of abaxial leaf epicuticular wax in Medicago truncatula irg1/palm1 mutants results in reduced spore differentiation of anthracnose and nonhost rust pathogens[J]. The Plant Cell, 2012,24(1):353-370. doi: 10.1105/tpc.111.093104 URL pmid: 22294617