外源酚酸诱导枳菌根共生相关基因筛选
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摘要
【目的】基于丛枝菌根(AM)真菌改变柑橘酚类现象,筛选外源酚酸诱导菌根共生差异基因,分析酚相关基因功能。【方法】以枳(Poncirus trifoliate L.)接种AM真菌,并施加外源酚酸,采用随机引物扩增,获得差异表达片段;继而克隆酚相关基因并进行生物信息学分析。【结果】试验采用20条随机引物和3条锚定引物扩增cDNA,共获得了154条差异显示的表达片段;对其中16条Northern杂交显示阳性的片段测序与比对,发现9条有相关功能注释,参与环境因子及内源信号的识别,调节共生体的形成;DD-11基因只有在外源酚酸添加和接种AM真菌的根系中表达,该片段cDNA全长序列长度为1382 bp,编码454个氨基酸,分子式为C_(2210)H_(3427)N_(603)O_(657)S_(31),其理论等电点和相对分子量分别为7.81和49950.03;磷酸化分析发现,该蛋白有22个丝氨酸残基、14个苏氨酸残基及7个酪氨酸残基可能成为蛋白激酶磷酸化位点。通过PredictProtein服务器对DD-11蛋白质的二级结构进行分析,该蛋白含有α螺旋22%、无规则卷曲58%、延伸链20%,不含有β折叠结构。【结论】本试验获得菌根共生过程酚相关基因,其功能与植物内源信号识别密切相关,为探讨酚类在菌根共生过程中的分子机制提供了新的视角。
[Objective] It has been reported that phnolics contents could be changed by the inoculation with arbuscular mycorrhizal(AM) fungi. In the present study, the differentially expressed genes were screened, and the genes function was analyzed with the exogenous phenolic addition. [Methods] Poncitrus trifoliata(L.) inoculated with AM fungi was treated by exogenous phenolic. We obtained the differentially expressed bands by differential-display RT-PCR(DDRT-PCR), and then cloned the related genes and analyzed the bioninformation.[Results] A total of 154 differentially accumulated transcript-derived fragments were identified. After the sequencing and blast of 16 fragments, 9 genes were found related to putative functions which participated into the recognition of environment factors and internal signals, and the adjust of mycorrhizal formation. The bioinformatic analysis showed that the cDNA sequences of DD-11 was 1382 bp in length, containing a complete 1080 bp open reading frame and 454 amino acids were encoded, which was only expressed under mycorrhizal inoculation and exogenous phenolic. The molecular weight, theoretical pI of the DD-11 gene deduced protein were 49950.03 Da,7.81, C_(2210)H_(3427)N_(603)O_(657)S_(31). The deduced DD-11 protein included 22 Ser, 14 Thr, 7 Tyr, which became the phosphorylation sites of protein kinase. The secondary structure of the deduced DD-11 protein mainly included Alpha helix, Random coil, Extended strand, Beta turn, which were 21.81%, 57.71%, 20.48% and have no Beta turn.[Conclusion] The genes related to phenolic expressed in mycorrhizal formation were obtained in our study, which were related closely to the plants signal recognition. Our study provide a new perspective of molecular mechanism of phnolics occurring in mycorrhizal formation.
[Objective] It has been reported that phnolics contents could be changed by the inoculation with arbuscular mycorrhizal(AM) fungi. In the present study, the differentially expressed genes were screened, and the genes function was analyzed with the exogenous phenolic addition. [Methods] Poncitrus trifoliata(L.) inoculated with AM fungi was treated by exogenous phenolic. We obtained the differentially expressed bands by differential-display RT-PCR(DDRT-PCR), and then cloned the related genes and analyzed the bioninformation.[Results] A total of 154 differentially accumulated transcript-derived fragments were identified. After the sequencing and blast of 16 fragments, 9 genes were found related to putative functions which participated into the recognition of environment factors and internal signals, and the adjust of mycorrhizal formation. The bioinformatic analysis showed that the cDNA sequences of DD-11 was 1382 bp in length, containing a complete 1080 bp open reading frame and 454 amino acids were encoded, which was only expressed under mycorrhizal inoculation and exogenous phenolic. The molecular weight, theoretical pI of the DD-11 gene deduced protein were 49950.03 Da,7.81, C_(2210)H_(3427)N_(603)O_(657)S_(31). The deduced DD-11 protein included 22 Ser, 14 Thr, 7 Tyr, which became the phosphorylation sites of protein kinase. The secondary structure of the deduced DD-11 protein mainly included Alpha helix, Random coil, Extended strand, Beta turn, which were 21.81%, 57.71%, 20.48% and have no Beta turn.[Conclusion] The genes related to phenolic expressed in mycorrhizal formation were obtained in our study, which were related closely to the plants signal recognition. Our study provide a new perspective of molecular mechanism of phnolics occurring in mycorrhizal formation.
引文
[1]Bonfante P,Genre A.Arbuscular mycorrhizal dialogues:do you speak‘plantish’or‘fungish’?Trends in Plant Science,2015,20:150-154.
[2]Bonfante P,Requena N.Dating in the dark:how roots respond to fungal signals to establish arbuscular mycorrhizal symbiosis.Current Opinion in Plant Biology,2011,14(4):451-457.
[3]Nagahashi G,Jr DD.The effects of hydroxy fatty acids on the hyphal branching of germinated spores of AM fungi.Fungal Biology,2011,15(4):351-358.
[4]Oldroyd GED.Speak,friend,and enter:signalling systems that promote beneficial symbiotic associations in plants.Nature Reviews Microbiology,2013,11(4):252-263.
[5]Lanfranco L,Fiorilli V,Venice F,Bonfante P.Strigolactones cross the kingdoms:plants,fungi,and bacteria in the arbuscular mycorrhizal symbiosis.Journal of Experimental Botany,2018,69(9):2175-2188.
[6]Zhang RQ,Zhao HQ,Zhu HH,Yao Q.Research progress in the biosynthesis of phenols in plants induced by arbuscular mycorrhizal fungi.Microbiology China,2010,37(8):1216-1221.(in Chinese)张瑞芹,赵海泉,朱红惠,姚青.丛枝菌根真菌诱导植物产生酚类物质的研究进展.微生物学通报,2010,37(8):1216-1221.
[7]Perner H,Rohn S,Driemel G,Batt N,Schwarz D,Kroh LW,George E.Effect of nitrogen species supply and mycorrhizal colonization on organosulfur and phenolic compounds in onions.Journal of Agricultural and Food Chemistry,2008,56(10):3538-3545.
[8]王森.药用植物丛枝菌根真菌资源及对麦冬的接种效应.西北农林科技大学硕士论文,2008
[9]He XL,Liu T,Zhao LL.Effects of inoculating AM fungi on physiological characters and nutritional components of Astragalus membranaceus under different N application levels.Chinese Journal of Applied Ecology,2009,20(9):2118-2122.(in Chinese)贺学礼,刘媞,赵丽莉.接种丛枝菌根对不同施氮水平下黄芪生理特性和营养成分的影响.应用生态学报,2009,20(9):2118-2122.
[10]He XL,Li J,Meng JJ,Zhao LL.Effects of Glomus mosseae on the growth and nutritional contents of Salvia miltiorrhiza Bge.under different P levels.Journal of Hebei University(Natural Science Edition),2011,31(4):418-423.(in Chinese)贺学礼,李君,孟静静,赵丽莉.摩西球囊霉和施P量对丹参生长和养分含量的影响.河北大学学报(自然科学版),2011,31(4):418-423.
[11]Zhang RQ,Zhu HH,Zhao HQ,Yao Q.Arbuscular mycorrhizal fungal inoculation increases phenolic synthesis in clover roots via hydrogen peroxide,salicylic acid and nitric oxide signaling pathways.Journal of Plant Physiology,2013,170(1):74-79.
[12]Ceccarelli N,Curadi M,Martelloni L,Sbrana C,Picciarelli P,Giovannetti M.Mycorrhizal colonization impacts on phenolic content and antioxidant properties of artichoke leaves and flower heads two years after field transplant.Plant and Soil,2010,335(1-2):311-323.
[13]Jurkiewicz A,Ryszka P,Anielska T,Waligorski P,Bialonska D,Goralska K,Tsimilli-Michael M,Turnau K.Optimization of culture conditions of Arnica montana L.:effects of mycorrhizal fungi and competing plants.Mycorrhiza,2010,20(5):293-306.
[14]Chen SC,Jin WJ,Liu AR,Zhang SJ,Liu DL,Wang FH,Lin XM,He CX.Arbuscular mycorrhizal fungi(AMF)increase growth and secondary metabolism in cucumber subjected to low temperature stress.Scientia Horticulturae,2013,160:222-229.
[15]Lopez-Raez JA,Charnikhova T,Fernandez I,Bouwmeester H,Pozo MJ.Arbuscular mycorrhizal symbiosis decreases strigolactone production in tomato.Journal of Plant Physiology,2011,168(3):294-297.
[16]Araim G,Saleem A,Arnason JT,Charest C.Root colonization by an arbuscular mycorrhizal(AM)fungus increases growth and secondary metabolism of purple coneflower,Echinacea purpurea(L.)Moench.Journal of Agricultural and Food Chemistry,2009,57(6):2255-2258.
[17]Toussaint JP,Smith FA,Smith SE.Arbuscular mycorrhizal fungi can induce the production of phytochemicals in sweet basil irrespective of phosphorus nutrition.Mycorrhiza,2007,17(4):291-297.
[18]Liu JY,Maldonado-Mendoza I,Lopez-Meyer M,Cheung F,Town,CD,Harrison MJ.Arbuscular mycorrhizal symbiosis is accompanied by local and systemic alterations in gene expression and an increase in disease resistance in the shoots.Plant Journal,2007,50(3):529-544.
[19]Li JF,He XH,Li H,Wang MY.Arbuscular mycorrhizal fungi increase growth and phenolics synthesis in Poncirus trifoliata under iron deficiency.Scientia Horticulturae,2015,183:87-92.
[20]Zhang XW,Dong WT,Sun JH,Feng F,Deng YW,He ZH,Oldroyd GED,Wang ET.The receptor kinase CERK1 has dual functions in symbiosis and immunity signaling.Plant Journal,2015,81(2):258-267.
[21]李建福.缺铁胁迫下柑橘菌根共生体酚类物质及其关键基因研究.华侨大学硕士论文,2015
[22]López-Ráez JA,Flors V,García JM,Pozo,M.J.AM symbiosis alters phenolic acid content in tomato roots.Plant Signaling&Behavior,2010,5(9):1138-1140.
[23]Schaarschmidt S,Kopka J,Ludwig-Muller J,Hause B.Regulation of arbuscular mycorrhization by apoplastic invertses:a enhanced invertase activity in the leaf apoplast affects the symbiotic interaction.Plant Journal,2007,51(3):390-405.
[24]Akiyama K,Matsuzaki K,Hayashi H.Plant sesquiterpenes induce hyphal branching in arbuscular mycorrhizal fungi.Nature,2005,435(7043):824-827.
[25]Tamasloukht M,Sejalon-Delmas N,Kluever A,Jauneau A,Roux C,Becard G.Root factors induce mitochondrial related gene expression and fungal respiration during the developmental switch from asymbiosis to presymbiosis in the arbuscular mycorrhizal fungus Gigaspora rosea.Plant Physiology,2003,131(3):1468-1478.
[26]Cutler S R,Rodriguez PL,Finkelstein RR,Abrams SR.Abscisic acid:emergence of a core signaling network.Annual Review of Plant Biology,2010,61(1):651-679.
[27]Charpentier M,Sun J,Wen JQ,Mysore KS,Oldroyd GED.Abscisic acid promotion of arbuscular mycorrhizal colonization requires a component of the PROTEINPHOSPHATASE 2A complex.Plant Physiology,2014,166(4):2077-2090.
[28]Chen MY,Liao JH,Li HR,Cai ZX,Guo ZJ,Wang ZS.Analysis of differentially expressed proteins in later developing stage fruitbody of Agaricus bisporus.Mycosystema,2013,32(5):855-861.(in Chinese)陈美元,廖剑华,李洪荣,蔡志欣,郭仲杰,王泽生.双孢蘑菇子实体发育后期差异表达蛋白质分析.菌物学报,2013,32(5):855-861.
[29]Aharon S,Gelli A,Senedden WA,Blumwald E.Activation of a plant plasma membrane Ca2+channel by TGα1,a heterotrimeric G proteinα-subunit homologue.Febs Letters,1998,424(1-2):17-21.
[30]Millner PA,Groarke DA,White IR.Synthetic peptides as probes of plant cell signaling.Plant Growth Regulation,1996,18(1-2):143-147.
[31]Liu SS,Chen J,Li SC,Zeng X,Meng ZX,Guo SX.Comparative transcriptome analysis of genes involved in GA-GID1-DELLA regulatory module in symbiotic and asymbiotic seed germination of Anoectochilus roxburghii(Wall.)Lindl.(Orchidaceae).International Journal of Molecular Sciences,2015,16(12):30190-30203.
[32]Gallou A,Declerck S,Cranenbrouck S.Transcriptional regulation of defence genes and involvement of the WRKYtranscription factor in arbuscular mycorrhizal potato root colonization.Functional&Integrative Genomics,2011,12(1):183-198.
[33]Wang W,Zhao P,Zhou XM,Xiong HX,Sun MX.Genome-wide identification and characterization of cystatin family genes in rice(Oryzasativa L.).Plant Cell Reports,2015,34(9):1579-1592.
[34]Song FQ,Kong XS,Li JZ,Chang W.Screening the related genes in the AM fungi and Amorpha fruticosa symbiosis with the suppression subtractive hybridization technique.Scientia Silvae Sinicae,2014,50(11):64-74.(in Chinese)宋福强,孔祥仕,李季泽,常伟.基于抑制消减杂交技术筛选AM真菌与紫穗槐共生相关基因.林业科学,2014,50(11):64-74.
[35]Yatzkan E,Szoor B,Feher Z,Dombradi V,Yarden O.Protein phosphatase 2A is involved in hyphal growth of Neurospora crassa.Molecular&general genetics:MGG,1998,259(5):523-531.
[36]Kosmidou E,Lunness P,Doonan JH.A type 2A protein phosphatase gene from Aspergillus nidulans is involved in hyphal morphogenesis.Current Genetics,2001,39(1):25-34.
[37]Du Y,Shi Y,Yang J,Chen XL,Xue MF,Zhou W,Peng YL.Aserine/threonine-protein phosphatase PP2A catalytic subunit is essential for asexual development and plant infection in Magnaporthe oryzae.Current Genetics,2013,59(1):33-41.
[38]Hildmann T,Ebneth M,Pe?acortés H,Sánchez-Serrano JJ,Willmitzer L,Prat S.General roles of abscisic and jasmonic acids in gene activation as a result of mechanical wounding.Plant Cell,1992,4(9):1157-1170.
[39]Chen HJ,Su CT,Lin CH,Huang GJ,Lin YH.Expression of sweet potato cysteine protease SPCP2 altered developmental characteristics and stress responses in transgenic Arabidopsis plants.Journal of Plant Physiology,2010,167(10):838-847.
[40]Ma YY,Zhang J,Chen J,Zhang LY,Zhu SP,Yan ST,Zhong GY.Isolation,subcellular localization and expression analysis of a citrus cysteine protease gene,CsCysP.Acta Horticulturae Sinica,2014,41(4):621-630.(in Chinese)马岩岩,张军,陈娇,张凌云,朱世平,闫树堂,钟广炎.柑橘半胱氨酸蛋白酶基因CsCysP的分离、亚细胞定位及表达分析.园艺学报,2014,41(4):621-630.
[2]Bonfante P,Requena N.Dating in the dark:how roots respond to fungal signals to establish arbuscular mycorrhizal symbiosis.Current Opinion in Plant Biology,2011,14(4):451-457.
[3]Nagahashi G,Jr DD.The effects of hydroxy fatty acids on the hyphal branching of germinated spores of AM fungi.Fungal Biology,2011,15(4):351-358.
[4]Oldroyd GED.Speak,friend,and enter:signalling systems that promote beneficial symbiotic associations in plants.Nature Reviews Microbiology,2013,11(4):252-263.
[5]Lanfranco L,Fiorilli V,Venice F,Bonfante P.Strigolactones cross the kingdoms:plants,fungi,and bacteria in the arbuscular mycorrhizal symbiosis.Journal of Experimental Botany,2018,69(9):2175-2188.
[6]Zhang RQ,Zhao HQ,Zhu HH,Yao Q.Research progress in the biosynthesis of phenols in plants induced by arbuscular mycorrhizal fungi.Microbiology China,2010,37(8):1216-1221.(in Chinese)张瑞芹,赵海泉,朱红惠,姚青.丛枝菌根真菌诱导植物产生酚类物质的研究进展.微生物学通报,2010,37(8):1216-1221.
[7]Perner H,Rohn S,Driemel G,Batt N,Schwarz D,Kroh LW,George E.Effect of nitrogen species supply and mycorrhizal colonization on organosulfur and phenolic compounds in onions.Journal of Agricultural and Food Chemistry,2008,56(10):3538-3545.
[8]王森.药用植物丛枝菌根真菌资源及对麦冬的接种效应.西北农林科技大学硕士论文,2008
[9]He XL,Liu T,Zhao LL.Effects of inoculating AM fungi on physiological characters and nutritional components of Astragalus membranaceus under different N application levels.Chinese Journal of Applied Ecology,2009,20(9):2118-2122.(in Chinese)贺学礼,刘媞,赵丽莉.接种丛枝菌根对不同施氮水平下黄芪生理特性和营养成分的影响.应用生态学报,2009,20(9):2118-2122.
[10]He XL,Li J,Meng JJ,Zhao LL.Effects of Glomus mosseae on the growth and nutritional contents of Salvia miltiorrhiza Bge.under different P levels.Journal of Hebei University(Natural Science Edition),2011,31(4):418-423.(in Chinese)贺学礼,李君,孟静静,赵丽莉.摩西球囊霉和施P量对丹参生长和养分含量的影响.河北大学学报(自然科学版),2011,31(4):418-423.
[11]Zhang RQ,Zhu HH,Zhao HQ,Yao Q.Arbuscular mycorrhizal fungal inoculation increases phenolic synthesis in clover roots via hydrogen peroxide,salicylic acid and nitric oxide signaling pathways.Journal of Plant Physiology,2013,170(1):74-79.
[12]Ceccarelli N,Curadi M,Martelloni L,Sbrana C,Picciarelli P,Giovannetti M.Mycorrhizal colonization impacts on phenolic content and antioxidant properties of artichoke leaves and flower heads two years after field transplant.Plant and Soil,2010,335(1-2):311-323.
[13]Jurkiewicz A,Ryszka P,Anielska T,Waligorski P,Bialonska D,Goralska K,Tsimilli-Michael M,Turnau K.Optimization of culture conditions of Arnica montana L.:effects of mycorrhizal fungi and competing plants.Mycorrhiza,2010,20(5):293-306.
[14]Chen SC,Jin WJ,Liu AR,Zhang SJ,Liu DL,Wang FH,Lin XM,He CX.Arbuscular mycorrhizal fungi(AMF)increase growth and secondary metabolism in cucumber subjected to low temperature stress.Scientia Horticulturae,2013,160:222-229.
[15]Lopez-Raez JA,Charnikhova T,Fernandez I,Bouwmeester H,Pozo MJ.Arbuscular mycorrhizal symbiosis decreases strigolactone production in tomato.Journal of Plant Physiology,2011,168(3):294-297.
[16]Araim G,Saleem A,Arnason JT,Charest C.Root colonization by an arbuscular mycorrhizal(AM)fungus increases growth and secondary metabolism of purple coneflower,Echinacea purpurea(L.)Moench.Journal of Agricultural and Food Chemistry,2009,57(6):2255-2258.
[17]Toussaint JP,Smith FA,Smith SE.Arbuscular mycorrhizal fungi can induce the production of phytochemicals in sweet basil irrespective of phosphorus nutrition.Mycorrhiza,2007,17(4):291-297.
[18]Liu JY,Maldonado-Mendoza I,Lopez-Meyer M,Cheung F,Town,CD,Harrison MJ.Arbuscular mycorrhizal symbiosis is accompanied by local and systemic alterations in gene expression and an increase in disease resistance in the shoots.Plant Journal,2007,50(3):529-544.
[19]Li JF,He XH,Li H,Wang MY.Arbuscular mycorrhizal fungi increase growth and phenolics synthesis in Poncirus trifoliata under iron deficiency.Scientia Horticulturae,2015,183:87-92.
[20]Zhang XW,Dong WT,Sun JH,Feng F,Deng YW,He ZH,Oldroyd GED,Wang ET.The receptor kinase CERK1 has dual functions in symbiosis and immunity signaling.Plant Journal,2015,81(2):258-267.
[21]李建福.缺铁胁迫下柑橘菌根共生体酚类物质及其关键基因研究.华侨大学硕士论文,2015
[22]López-Ráez JA,Flors V,García JM,Pozo,M.J.AM symbiosis alters phenolic acid content in tomato roots.Plant Signaling&Behavior,2010,5(9):1138-1140.
[23]Schaarschmidt S,Kopka J,Ludwig-Muller J,Hause B.Regulation of arbuscular mycorrhization by apoplastic invertses:a enhanced invertase activity in the leaf apoplast affects the symbiotic interaction.Plant Journal,2007,51(3):390-405.
[24]Akiyama K,Matsuzaki K,Hayashi H.Plant sesquiterpenes induce hyphal branching in arbuscular mycorrhizal fungi.Nature,2005,435(7043):824-827.
[25]Tamasloukht M,Sejalon-Delmas N,Kluever A,Jauneau A,Roux C,Becard G.Root factors induce mitochondrial related gene expression and fungal respiration during the developmental switch from asymbiosis to presymbiosis in the arbuscular mycorrhizal fungus Gigaspora rosea.Plant Physiology,2003,131(3):1468-1478.
[26]Cutler S R,Rodriguez PL,Finkelstein RR,Abrams SR.Abscisic acid:emergence of a core signaling network.Annual Review of Plant Biology,2010,61(1):651-679.
[27]Charpentier M,Sun J,Wen JQ,Mysore KS,Oldroyd GED.Abscisic acid promotion of arbuscular mycorrhizal colonization requires a component of the PROTEINPHOSPHATASE 2A complex.Plant Physiology,2014,166(4):2077-2090.
[28]Chen MY,Liao JH,Li HR,Cai ZX,Guo ZJ,Wang ZS.Analysis of differentially expressed proteins in later developing stage fruitbody of Agaricus bisporus.Mycosystema,2013,32(5):855-861.(in Chinese)陈美元,廖剑华,李洪荣,蔡志欣,郭仲杰,王泽生.双孢蘑菇子实体发育后期差异表达蛋白质分析.菌物学报,2013,32(5):855-861.
[29]Aharon S,Gelli A,Senedden WA,Blumwald E.Activation of a plant plasma membrane Ca2+channel by TGα1,a heterotrimeric G proteinα-subunit homologue.Febs Letters,1998,424(1-2):17-21.
[30]Millner PA,Groarke DA,White IR.Synthetic peptides as probes of plant cell signaling.Plant Growth Regulation,1996,18(1-2):143-147.
[31]Liu SS,Chen J,Li SC,Zeng X,Meng ZX,Guo SX.Comparative transcriptome analysis of genes involved in GA-GID1-DELLA regulatory module in symbiotic and asymbiotic seed germination of Anoectochilus roxburghii(Wall.)Lindl.(Orchidaceae).International Journal of Molecular Sciences,2015,16(12):30190-30203.
[32]Gallou A,Declerck S,Cranenbrouck S.Transcriptional regulation of defence genes and involvement of the WRKYtranscription factor in arbuscular mycorrhizal potato root colonization.Functional&Integrative Genomics,2011,12(1):183-198.
[33]Wang W,Zhao P,Zhou XM,Xiong HX,Sun MX.Genome-wide identification and characterization of cystatin family genes in rice(Oryzasativa L.).Plant Cell Reports,2015,34(9):1579-1592.
[34]Song FQ,Kong XS,Li JZ,Chang W.Screening the related genes in the AM fungi and Amorpha fruticosa symbiosis with the suppression subtractive hybridization technique.Scientia Silvae Sinicae,2014,50(11):64-74.(in Chinese)宋福强,孔祥仕,李季泽,常伟.基于抑制消减杂交技术筛选AM真菌与紫穗槐共生相关基因.林业科学,2014,50(11):64-74.
[35]Yatzkan E,Szoor B,Feher Z,Dombradi V,Yarden O.Protein phosphatase 2A is involved in hyphal growth of Neurospora crassa.Molecular&general genetics:MGG,1998,259(5):523-531.
[36]Kosmidou E,Lunness P,Doonan JH.A type 2A protein phosphatase gene from Aspergillus nidulans is involved in hyphal morphogenesis.Current Genetics,2001,39(1):25-34.
[37]Du Y,Shi Y,Yang J,Chen XL,Xue MF,Zhou W,Peng YL.Aserine/threonine-protein phosphatase PP2A catalytic subunit is essential for asexual development and plant infection in Magnaporthe oryzae.Current Genetics,2013,59(1):33-41.
[38]Hildmann T,Ebneth M,Pe?acortés H,Sánchez-Serrano JJ,Willmitzer L,Prat S.General roles of abscisic and jasmonic acids in gene activation as a result of mechanical wounding.Plant Cell,1992,4(9):1157-1170.
[39]Chen HJ,Su CT,Lin CH,Huang GJ,Lin YH.Expression of sweet potato cysteine protease SPCP2 altered developmental characteristics and stress responses in transgenic Arabidopsis plants.Journal of Plant Physiology,2010,167(10):838-847.
[40]Ma YY,Zhang J,Chen J,Zhang LY,Zhu SP,Yan ST,Zhong GY.Isolation,subcellular localization and expression analysis of a citrus cysteine protease gene,CsCysP.Acta Horticulturae Sinica,2014,41(4):621-630.(in Chinese)马岩岩,张军,陈娇,张凌云,朱世平,闫树堂,钟广炎.柑橘半胱氨酸蛋白酶基因CsCysP的分离、亚细胞定位及表达分析.园艺学报,2014,41(4):621-630.