苯并噻二唑诱导苹果抗斑点落叶病研究
|
摘要
苹果斑点落叶病是影响苹果产量和品质的重要病害。当前,有关对苹果斑点落叶病的研究,仅局限在对病原菌生物学特性以及利用化学手段防治病害方面的研究。鉴于诱导抗病性的研究进展与植物激活剂的特点,本文以苯并噻二唑(Benzothiadiazole,BTH)为诱导因子,以苹果树和斑点落叶病菌(Alternaira alternata f.sp.mali)为研究对象,研究诱导剂在不同浓度下对抗苹果斑点落叶病的诱导作用、诱导抗病的持效期、组织病理学特征以及生理生化机制。其中,生理生化机制主要研究诱导抗病性与苯丙氨酸解氨酶(PAL)活性、几丁质酶活性、β-1,3-葡聚糖酶活性、木质素和富含羟脯氨酸糖蛋白(HRGP)含量的关系。
采用常规组织分离法分离纯化苹果斑点落叶病菌,病原形态观察和人工回接试验鉴定病原菌。BTH诱导苹果主枝顶梢,整株接种病原菌后,采用病情指数统计发病程度。用整叶组织透明染色法进行组织病理学观察,用比色法测定PAL、几丁质酶、β-1,3-葡聚糖酶活性以及木质素和HRGP含量的变化。PDA培养基测定不同浓度BTH离体抑菌活性。
结果表明:BTH能诱导苹果树产生系统获得性抗性,当诱导浓度为150μg·mL-1时,可使病情指数由对照的26.34下降到16.87,抗病持效期可达15d,其中最佳诱导时间为处理后3～6d,而BTH本身对苹果斑点落叶病菌分生孢子萌发及菌丝生长无明显影响。组织病理学观察发现,BTH诱导抑制病菌侵染钉的形成,限制病菌向细胞内扩展;生理生化指标测定表明,BTH诱导能提高苹果叶片PAL、几丁质酶、β-1,3-葡聚糖酶活性以及木质素和HRGP的含量。BTH处理的果苗防卫反应启动早于对照果苗。
Alternaira alternata f.sp.mali (AAM) is a most important epidemic disease that threatens the yield and quality of apple production. Currently, the research about AAM is just around biological character of pathogen and the chemical prevention of the disease. Considering the advance of research on induced disease resistance and the characteristic of plant activator, the effects of resistance induced by benzothiadiazole (BTH) at different concentrations against AAM in apple were investigated. The duration of induced disease resistance were studied. Moreover the histopathological mechanism and physiological mechanism were researched. The physiological mechanism comprised the relationship of BTH induced disease resistence with the contents of hydroxyproline-rich glycoprotein (HRGP) and Lignin, with the activity of phenylalanine ammonialyase (PAL), chitinase andβ-1, 3-glucanase.
The AAM was isolated and purified from infected apple leaves by tissue isolating method. The conidiophores and mycelia structure of the pathogen were observed by microscope. The pathogenic power was investigated by leaf in vitro inoculation. Apples were inoculated with spores of AAM after applying BTH as a foliar spray to peak branch and disease index was investigated. The histopathological response was investigated with whole leaf clearing and staining technique. The activities of enzyme and the contents of Lignin and HRGP were measured with colorimetric method. The inhibitory effects of different concentration BTH against AAM on potato dextrose agar were studied.
The result showed that application of BTH at different concentrations on apple leaves had significant effect to restrain AAM. The effect of 150μg·mL-1 was distinctly better than others, and it could reduce the disease index from 26.34 to 16.87. In addition, the induced disease resistance could last more than 15 days, but the best induced times were 3～6 days after 150μg·mL-1 BTH treatment. However, conidiophores germination and mycelia growth were not restricted by BTH itself. The penetration pegs’formation of pathogen was inhibited, and then pathogen extending was restricted. The activities of PAL, chitinase andβ-1, 3-glucanase of apple leaf rised after BTH inducement. Moreover, the contents of Lignin and HRGP of apple leaf increased too. So the defendable reaction of induced apple trees was started earlier than control.
采用常规组织分离法分离纯化苹果斑点落叶病菌,病原形态观察和人工回接试验鉴定病原菌。BTH诱导苹果主枝顶梢,整株接种病原菌后,采用病情指数统计发病程度。用整叶组织透明染色法进行组织病理学观察,用比色法测定PAL、几丁质酶、β-1,3-葡聚糖酶活性以及木质素和HRGP含量的变化。PDA培养基测定不同浓度BTH离体抑菌活性。
结果表明:BTH能诱导苹果树产生系统获得性抗性,当诱导浓度为150μg·mL-1时,可使病情指数由对照的26.34下降到16.87,抗病持效期可达15d,其中最佳诱导时间为处理后3～6d,而BTH本身对苹果斑点落叶病菌分生孢子萌发及菌丝生长无明显影响。组织病理学观察发现,BTH诱导抑制病菌侵染钉的形成,限制病菌向细胞内扩展;生理生化指标测定表明,BTH诱导能提高苹果叶片PAL、几丁质酶、β-1,3-葡聚糖酶活性以及木质素和HRGP的含量。BTH处理的果苗防卫反应启动早于对照果苗。
Alternaira alternata f.sp.mali (AAM) is a most important epidemic disease that threatens the yield and quality of apple production. Currently, the research about AAM is just around biological character of pathogen and the chemical prevention of the disease. Considering the advance of research on induced disease resistance and the characteristic of plant activator, the effects of resistance induced by benzothiadiazole (BTH) at different concentrations against AAM in apple were investigated. The duration of induced disease resistance were studied. Moreover the histopathological mechanism and physiological mechanism were researched. The physiological mechanism comprised the relationship of BTH induced disease resistence with the contents of hydroxyproline-rich glycoprotein (HRGP) and Lignin, with the activity of phenylalanine ammonialyase (PAL), chitinase andβ-1, 3-glucanase.
The AAM was isolated and purified from infected apple leaves by tissue isolating method. The conidiophores and mycelia structure of the pathogen were observed by microscope. The pathogenic power was investigated by leaf in vitro inoculation. Apples were inoculated with spores of AAM after applying BTH as a foliar spray to peak branch and disease index was investigated. The histopathological response was investigated with whole leaf clearing and staining technique. The activities of enzyme and the contents of Lignin and HRGP were measured with colorimetric method. The inhibitory effects of different concentration BTH against AAM on potato dextrose agar were studied.
The result showed that application of BTH at different concentrations on apple leaves had significant effect to restrain AAM. The effect of 150μg·mL-1 was distinctly better than others, and it could reduce the disease index from 26.34 to 16.87. In addition, the induced disease resistance could last more than 15 days, but the best induced times were 3～6 days after 150μg·mL-1 BTH treatment. However, conidiophores germination and mycelia growth were not restricted by BTH itself. The penetration pegs’formation of pathogen was inhibited, and then pathogen extending was restricted. The activities of PAL, chitinase andβ-1, 3-glucanase of apple leaf rised after BTH inducement. Moreover, the contents of Lignin and HRGP of apple leaf increased too. So the defendable reaction of induced apple trees was started earlier than control.
引文
[1] 胡同乐,曹克强,王树桐,等.生长季苹果斑点落叶病流行主导因素的确定[J].植物病理学报, 2005, 35(4):374-377
[2] 吴桂本,王英姿,王培松,等.苹果斑点落叶病菌的分化及生物学研究[J].中国果树,1999,(4):11-15
[3] 薛 军,孙继玲,王孝友.苹果斑点落叶病发生规律的研究[J].落叶果树,2006,(1):36-37
[4] 吴桂本,刘传德,王培松,等.苹果轮纹病和炭疽病发生规律的研究[J].莱阳农学院学报, 2000, 17(2): 86-92
[5] 吴桂本,刘传德,王继秋,等.苹果轮纹病和炭疽病病原菌生物学研究[J].中国果树,2001,(1):7-11
[6] 张 姝,张永杰,刘慧平,等.苹果斑点落叶病菌的分离及其对杀菌剂的敏感性[J].山西农业大学学报,2004,24(4):382-384
[7] Durrant W E, Dong X. Systemic acquired resistance [J].Annual Review of Phytopathology, 2004, 42: 185-209
[8] Vallad G E, Goodman R M. Systemic acquired resistance and induced systemic resistance in conventional agriculture [J]. Crop Science, 2004, 44: 1920-1934
[9] 张一宾.非杀菌农药对植物病害的防治机理—包括植物激活剂[J].世界农药,2003,25(2):19-23
[10] Kloepper J W, Tunzun S. Proposed definition related to induced disease resistance [J]. Biocontrol Sci Technology. 1992, 2:349-351
[11] Silue D, Pajot E, Cohen Y. Induction of resistance to downy mildew(Peronospora parasitica) in cauliflower by DL-β-amino-n-butanoic acid(BABA)[J]. Plant Pathol, 2002, 51:97-102
[12] Hewitt G. New modes of action of fungicides [J].Pesticide Outlook, 2000, l1:28-32
[13] Kessmann H, Staub T, Ligon J, et a1. Activation of systemic acquired disease resistance in plants[J]. Eur J Plant Pathol,1994,100:359-369
[14] Kessmann H, Staub T, Hofmann C, et a1. Induction of systemic acquired disease resistance in plants by chemicals [J]. Annual Review Phytopathol,1994,32:439-459
[15] 董汉松.植物诱导抗性原理和研究[M].北京:科学出版社,1995
[16] 陈 捷.植物病理生理学[M].沈阳:辽宁科学技术出版社,1994
[17] Kuchitsu K, YazakiY, Sakano K, et al. Transient cytoplasmic PH change and ion fluxes through the plasma membrane in suspension-cultured rice cells triggered by N-acetylchito oligosaccharide elicitor [J].Plant Cell Physiol,1997,38:1012-1018
[18] Ajlan A M, Potter D A. Lack of effect of tobacco mosaic virus induced systemic acquired resistance on arthropod herbivores in tobacco [J]. Phytopathology, 1992, 82:647-651
[19] Mucharromah E, Kud J. Oxalate and phosphates induce systemic resistance against disease caused by fungi, bacteria and viruses in cucumber [J]. CropProt, 1991, 10: 265-270
[20] Andebrhan T, Woodr K S. The effect of ultraviolet radiation on the reaction of Phaseolus vulgaristo species of Colletotrichum [J].Physiol Plant Pathol, 1980, 17: 105-110
[21] 李玉红,程智慧,陈 鹏,等.苯并噻二唑(BTH)诱导黄瓜幼苗对霜霉病抗性的研究[J].园艺学报,2006, 33(2): 278-282
[22] Reinbothe S, Mollenhauer B, Reinbothe C. The regulation of plant gene expression by jasmonates in response to environmental cues and pathogens[J].The Plant Cell,1994,6:1197- 1209
[23] Heil M, Hilpert A,Kaiser W, et al. Reduced growth and seed set following chemical induction of pathogen defense: does systemic acquired resistance(SAR) incur allocation costs [J]. Journal of Ecology,2000,88:645-654
[24] Heil M, Baldwin I T. Fitness costs of induced resistance the emerging experimental support for a slippery concept[J].Trends in Plant Science, 2002,7: 61-67
[25] Ellingboe A H. Plant-pathogen interactions: genetic and comparative analysis [J].Euro. J. Plant Pathol, 2001, 107:79 -84
[26] Tuzun S, Kuc J. Persistence of induced systemic resistance to bule mold in tobacco plants desired via tissue culture [J].Phytopathology,1987,77: 1032-1035
[27] Kuc J. Protection of cucumber against Colletotrichum lagenarium by Colletotrichum lagenarium [J]. Physiol. Plant Pathol, 1975,7:195-199
[28] 王锐萍.新疆甜瓜对瓜类疫霉菌抗性的诱导[J].植物保护,2000, 26(3): 9-11
[29] Alstrom S. Induction of disease resistance in common bean susceptible to halo blight bacterial pathogen after seed bacterization with rhizosphere pseudomonas[J]. J. Gen. Appl.Microbiol, 1991,37:495- 501
[30] Wei G , Kloepper J W , Tuzun S .Introduction of systemic resistance of cucumber to colletotrichum orbiculare by select strains of plant growth promoting rhizobacter [J]. Phytopathol, 1991, 81:1508 -1512
[31] Van Buren A, Andre C, Ishimaru C A. Biological control of bacterial ring rot pathogen by entophytic bacterial solated from potato[J].(abstract)Phytopathol, 1993, 83:1406
[32] 杨海莲,孙晓璐,宋未.植物根际促生细菌和内生细菌的诱导抗病性的研究进展[J].植物病理学报, 2000,30(2):106-110
[33] Palva T K, Holmstrom KO, Heino P, et al. Induction of plant defense response by exoenzymes of Erwinia carotovora sub sp. Carotovora[J].Mot Plant Microbe Interact, 1993,6(2):190-196
[34] 彭霞薇,张洪勋,白志辉.草酸青霉菌果胶酶诱导黄瓜抗黑星病研究[J].植物病理学报, 2004,34(1): 69-74
[35] 胡东维,李振岐,康振生.小麦抗白粉菌初侵染过敏性反应的细胞学研究[J].浙江农业大学学报, 1997, 23(4):397-404
[36] 于汉寿,张益民, 陈永萱,等.水溶性壳聚糖对小麦和油菜几丁质酶的诱导作用[J].江苏农业学报, 1999,15(2):67-70
[37] 刘喜存,刘红彦,李洪连.植物诱导抗病性及其研究现状[J].河南农业科学, 2006 (4): 12-16
[38] Wildon D C, Thain J F, Minchin P H E, et al. Electrical signaling and systemic proteinase inhibitor induction in the wounded plant [J]. Nature,1992,360:62-65
[39] Uknes S, Mauch-Mani B, Moyer M ,et al. Acquired resistance in Arabidopsis [J]. Plant Cell,1992,4: 645- 655
[40] Sequeira L. Mechanisms of induced resistance in plants [J]. Annu Rev Microbiol.1986,37:51 -79
[41] 鲍丽丽,刘凤丽,范志金. 具有植物诱导抗病活性的先导化合物及其结构修饰[J]. 农药学学报,2005, 7(3):201-209
[42] Delaney T P, Ukness, Vernooij B, et al. A central role of salicylic acid in plant disease resistance [J]. Science, 1994,266(5188):1247-1250
[43] 侯 珲,朱建兰,周红平,等.BTH 和水杨酸(SA)对甜瓜抗白粉病的诱导作用[J].果树学报, 2006,23(5):736-739
[44] 李淑菊,马德华,庞金安,等.水杨酸对黄瓜几种酶活性及抗病性的诱导作用[J].华北农学报, 2000, 15(2):118-122
[45] OkunoT, Nakyama M, Okajima N, et al. Systemic resistance to down mildew and appearance of acid soluble proteins in cucumber leaves treated with biotic and abiotic inducers[J].Ann Phytopath Soc Japan,1991, 57: 202- 211
[46] Bol J F, van kan JA. The Synthesis and possible functions of virus –induced proteins in plants[J].Microbiol. Sci1, 1988,5(2):47
[47] Perumal V, Jay S, Levespue C, et al. A role for jasmonate in pathogen defense of Arabidopsis[J]. Proceedings of the National Academy of Sciences of the USA, 1998, 95(12): 7209-7214
[48] Neto G C, Kono Y, Hyakutake H, et al. Isolation and identification of jasmonic acid from wild rice as an antifungal substance[J].Agricultural and Biological Chemistry, 1991, 55: 3097-3098
[49] Thaler J S, Owen B, Higgins V J. The Role of the Jasmonate response in plant susceptibility to diverse pathogens with a range of lifestyles [J]. Plant Physiology, 2004, 135(1): 530-538
[50] Kogel K H, Beckhove U, Dreschers J. et al .Acquired resistance in barley-the resistance mechanism induced by 2,6-dichloroisonicotinic acid is a phenocopy of a genetically based mechanism governing race specific powdery mildew resistance[J]. Plant Physiol, 1994, 106:1269-1277
[51] Dann E, Diers B, Byrum J, et al. Effect of treating soybean with 2,6-dichloroisonicotinic acid(INA) and benzothiadiazole(BTH) on seed yields and the level of disease caused by Sclerotinia sclerotiorum in field and greenhouse studies[J].Euro J Plant Pathol, 1998, 104: 271-278
[52] 程智慧,李玉红,孟焕文,等.BTH 诱导黄瓜幼苗对霜霉病的抗性与细胞壁HRGP和木质素含量的关系[J].中国农业科学,2006,39(5):935-940
[53] Buonaurio R, Scarponi L, Ferrara M, et al. Induction of systemic acquired resistance in pepper plants by acibenzolar-S-methyl against bacterial spot disease[J]. European Journal of Plant Pathology,2002, 108 (1): 41-49
[54] Eikemo H, Stensvand A, Tronsmo A M. Induced resistance as a possible means to control diseases of strawberry caused by Phytophthora spp[J]. Plant Disease, 2003, 87(4): 345-350
[55] Perez L, Rodriguez M E, Rodriguez F, et al. Efficacy of acibenzolar-S-methyl, an inducer of systemic acquired resistance against tobacco blue mould caused by Peronospora hyoscyami f. sp. tabacina[J]. Crop Protection, 2003, 22: 405-413
[56] 王晓军,徐尚成,肖国民.植物激活剂的研究开发进展[J].化工纵横,2002,(1):22-26
[57] Hammerschmidt R, Lamport D T A, Muldoon E P. Cell wall hydroxyproline enhancement and lignin deposition as an early event in the resistance of cucumber to Cladosporium cucumerinum [J]. Physiol Plant Pathol,1984,24:43-47
[58] Kovats K, Binder A, Hohl H R. Cytology of induced systemic resistance of cucumber to Colletotrichum lagenarium [J].Planta,1991,183:484-490
[59] Bradley D J, Kjellbom P, Lamb C J. Elicitor-induced and wound-induced oxidative cross-linking of a proline-rich plant cell wall protein-a novel, rapid defense response[J]. Cell, 1992,70:21-30
[60] Van’t Slot K A E, Knogge W. A dual role for microbial pathogen-derived effector proteins in plant disease and resistance [J]. Critical Reviews in Plant Sciences, 2002, 21(3): 229-271
[61] 李振歧主编.植物免疫学[M].北京:中国农业出版社, 1995
[62] Kosuge T. The role of phenolics in host response to infection [J]. Annu Rev Phytopathol, 1969,7:105-122
[63] Cramer C L, Ryder T B, Bell I N, et al. Rapid switching of plant gene expression induced by fungus elicitor [J]. Science, 1985,227:1240-1243
[64] Friend J, Reynolds S B, Aveyard M A. Phenylalanine ammonia-lyase、Chlorogenic acid and lignin in potato tuber tissue inoculated with Phytophthora infestans [J]. Physiol Mal Plant Pathol,1973,3: 495- 507
[65] 王雅平.小麦对赤霉病抗性不同品种SOD活性[J].植物生理学报,1993, 19 (4):353-358
[66] 陈利锋,宋玉立,徐雍皋.抗感赤霉病小麦品种超氧化物歧化酶和过氧化氢酶的活性比较[J].植物病理学报,1997,27(3):209-213
[67] 朱有林,刘纪麟.受玉米大斑病菌侵染后玉米抗感近位等基因系SOD动态变化的研究[J]植物病理学报,1996,26(2): 133-137
[68] 曾永三, 王振中.活性氧和超氧化物岐化酶在植物抗病反应中的作用[J]. 仲恺农业技术学院学报, 1999,12(4):55-63
[69] 秦引萍,杨依军,彭友良等.稻瘟菌侵染诱导性水稻脂氧合酶的同工酶鉴定及性状分析[J].农业生物技术学报,1996,4(2):180-189
[70] 杨依军,秦引萍,黄俊斌等.稻瘟菌诱导性稻叶脂氧合酶的进一步纯化及抗体制备[J].植物病理学报,1997,27(3):243-249
[71] 方仲达.植病研究法[M].北京:中国农业出版社,1998
[72] 陈荣丽.BION 对小麦抗叶锈病性的诱导及生化机制研究[D].学位论文,2005,四川农业大学:20
[73] 史益敏.几丁质酶酶活性的测定[A].现代植物生理学实验指南[C],中国科学院上海植物生理研究所主编,北京:科学出版社,1999. 129-130
[74] 史益敏. β-1,3-葡聚糖酶活性的测定[A].现代植物生理学实验指南[C],中国科学院上海植物生理研究所主编.北京:科学出版社.1999. 128-129
[75] X.H. 波钦诺克 , 植物生物化学分析方法 ( 荆家海 , 丁钟荣 译 ) [M]. 北京 : 科学出版社,1981,178-181
[76] 胡景江, 朱 玮, 文建雷.杨树细胞壁HRGP和木质素的诱导积累与其对溃疡病抗性的关系[J].植物病理学报, 1999, 29(2): 151-156
[77] 李建华,李雄彪.细胞壁的制备及其羟脯氨酸含量的测定[J].植物生物学通讯, 1993,29 (5): 363-365
[78] 李伟,熊谨,陈晓阳.木质素代谢的生理意义及其遗传控制研究进展[[J].西北植物学报,2003,234:675-681
[79] Nicole Benhamou, Richard R Belanger. Benzothiadiazole-mediated induced resistance to Fusarium oxysporum f. sp. Radicis-lycopersici in tomato[J].Plant Physiology, 1998, 118(4): 1203-1213
[80] 王晨芳.化学诱导剂在诱导黄瓜抗病性中的作用[D].学位论文,2005,西北农林科技大学
[81] Nicole Benhamou, Richard R Belanger. Induction of systemic resistance to Pythium damping-off in cucumber plants by benzothiadiazole: Ultrastructure and cytochemistry of the host response [J].Plant J,1998,14:13-21
[82] Xuei X.L., Jarlfors U, Kuc J. Ultrastructural changes associated with induced systemic resistance of cucumber to disease: host response and development of colletotrichum lagenarium in systemically protected leaves[J]. Can.J.Bot.,1988, 66: 1028-1038
[83] 王颖, 景耀. 植物诱导抗病机理的研究进展[J].西北林学院学报,1997,12(3):52-57
[84] 程智慧,李玉红,孟焕文,等.BTH 诱导黄瓜幼苗对霜霉病的抗性与细胞壁HRGP和木质素含量的关系[J].中国农业科学,2006,39(5):935-940
[85] 葛秀春,宋凤鸣,陈永叶,等.苯并噻二唑诱发水稻对稻瘟病抗性中防卫相关酶活性的变化.中国水稻科学[[J].2002, 16(2):171-175
[86] Selitrennikoff C P. Antifungal proteins[J]. Appl Environ Microbiol, 2001,67:2883-2894
[87] Ye X S, Pan S Q, Kuc K J. Activity, isozyme pattern, and cellular localization of peroxidase as related to systemic resistance of tobacco to blue mold (Peronospora tabacina) and to tobacco moasaic virus [J]. Phytopathology, 1990, 80:1295-1299
[88]宾金华,姜 胜,黄胜琴,潘瑞炽.茉莉酸甲酯诱导烟草幼苗抗炭疽病与PAL活性及细胞壁 物质的关系[J].植物生理学报,2000,26(1):1-6
[2] 吴桂本,王英姿,王培松,等.苹果斑点落叶病菌的分化及生物学研究[J].中国果树,1999,(4):11-15
[3] 薛 军,孙继玲,王孝友.苹果斑点落叶病发生规律的研究[J].落叶果树,2006,(1):36-37
[4] 吴桂本,刘传德,王培松,等.苹果轮纹病和炭疽病发生规律的研究[J].莱阳农学院学报, 2000, 17(2): 86-92
[5] 吴桂本,刘传德,王继秋,等.苹果轮纹病和炭疽病病原菌生物学研究[J].中国果树,2001,(1):7-11
[6] 张 姝,张永杰,刘慧平,等.苹果斑点落叶病菌的分离及其对杀菌剂的敏感性[J].山西农业大学学报,2004,24(4):382-384
[7] Durrant W E, Dong X. Systemic acquired resistance [J].Annual Review of Phytopathology, 2004, 42: 185-209
[8] Vallad G E, Goodman R M. Systemic acquired resistance and induced systemic resistance in conventional agriculture [J]. Crop Science, 2004, 44: 1920-1934
[9] 张一宾.非杀菌农药对植物病害的防治机理—包括植物激活剂[J].世界农药,2003,25(2):19-23
[10] Kloepper J W, Tunzun S. Proposed definition related to induced disease resistance [J]. Biocontrol Sci Technology. 1992, 2:349-351
[11] Silue D, Pajot E, Cohen Y. Induction of resistance to downy mildew(Peronospora parasitica) in cauliflower by DL-β-amino-n-butanoic acid(BABA)[J]. Plant Pathol, 2002, 51:97-102
[12] Hewitt G. New modes of action of fungicides [J].Pesticide Outlook, 2000, l1:28-32
[13] Kessmann H, Staub T, Ligon J, et a1. Activation of systemic acquired disease resistance in plants[J]. Eur J Plant Pathol,1994,100:359-369
[14] Kessmann H, Staub T, Hofmann C, et a1. Induction of systemic acquired disease resistance in plants by chemicals [J]. Annual Review Phytopathol,1994,32:439-459
[15] 董汉松.植物诱导抗性原理和研究[M].北京:科学出版社,1995
[16] 陈 捷.植物病理生理学[M].沈阳:辽宁科学技术出版社,1994
[17] Kuchitsu K, YazakiY, Sakano K, et al. Transient cytoplasmic PH change and ion fluxes through the plasma membrane in suspension-cultured rice cells triggered by N-acetylchito oligosaccharide elicitor [J].Plant Cell Physiol,1997,38:1012-1018
[18] Ajlan A M, Potter D A. Lack of effect of tobacco mosaic virus induced systemic acquired resistance on arthropod herbivores in tobacco [J]. Phytopathology, 1992, 82:647-651
[19] Mucharromah E, Kud J. Oxalate and phosphates induce systemic resistance against disease caused by fungi, bacteria and viruses in cucumber [J]. CropProt, 1991, 10: 265-270
[20] Andebrhan T, Woodr K S. The effect of ultraviolet radiation on the reaction of Phaseolus vulgaristo species of Colletotrichum [J].Physiol Plant Pathol, 1980, 17: 105-110
[21] 李玉红,程智慧,陈 鹏,等.苯并噻二唑(BTH)诱导黄瓜幼苗对霜霉病抗性的研究[J].园艺学报,2006, 33(2): 278-282
[22] Reinbothe S, Mollenhauer B, Reinbothe C. The regulation of plant gene expression by jasmonates in response to environmental cues and pathogens[J].The Plant Cell,1994,6:1197- 1209
[23] Heil M, Hilpert A,Kaiser W, et al. Reduced growth and seed set following chemical induction of pathogen defense: does systemic acquired resistance(SAR) incur allocation costs [J]. Journal of Ecology,2000,88:645-654
[24] Heil M, Baldwin I T. Fitness costs of induced resistance the emerging experimental support for a slippery concept[J].Trends in Plant Science, 2002,7: 61-67
[25] Ellingboe A H. Plant-pathogen interactions: genetic and comparative analysis [J].Euro. J. Plant Pathol, 2001, 107:79 -84
[26] Tuzun S, Kuc J. Persistence of induced systemic resistance to bule mold in tobacco plants desired via tissue culture [J].Phytopathology,1987,77: 1032-1035
[27] Kuc J. Protection of cucumber against Colletotrichum lagenarium by Colletotrichum lagenarium [J]. Physiol. Plant Pathol, 1975,7:195-199
[28] 王锐萍.新疆甜瓜对瓜类疫霉菌抗性的诱导[J].植物保护,2000, 26(3): 9-11
[29] Alstrom S. Induction of disease resistance in common bean susceptible to halo blight bacterial pathogen after seed bacterization with rhizosphere pseudomonas[J]. J. Gen. Appl.Microbiol, 1991,37:495- 501
[30] Wei G , Kloepper J W , Tuzun S .Introduction of systemic resistance of cucumber to colletotrichum orbiculare by select strains of plant growth promoting rhizobacter [J]. Phytopathol, 1991, 81:1508 -1512
[31] Van Buren A, Andre C, Ishimaru C A. Biological control of bacterial ring rot pathogen by entophytic bacterial solated from potato[J].(abstract)Phytopathol, 1993, 83:1406
[32] 杨海莲,孙晓璐,宋未.植物根际促生细菌和内生细菌的诱导抗病性的研究进展[J].植物病理学报, 2000,30(2):106-110
[33] Palva T K, Holmstrom KO, Heino P, et al. Induction of plant defense response by exoenzymes of Erwinia carotovora sub sp. Carotovora[J].Mot Plant Microbe Interact, 1993,6(2):190-196
[34] 彭霞薇,张洪勋,白志辉.草酸青霉菌果胶酶诱导黄瓜抗黑星病研究[J].植物病理学报, 2004,34(1): 69-74
[35] 胡东维,李振岐,康振生.小麦抗白粉菌初侵染过敏性反应的细胞学研究[J].浙江农业大学学报, 1997, 23(4):397-404
[36] 于汉寿,张益民, 陈永萱,等.水溶性壳聚糖对小麦和油菜几丁质酶的诱导作用[J].江苏农业学报, 1999,15(2):67-70
[37] 刘喜存,刘红彦,李洪连.植物诱导抗病性及其研究现状[J].河南农业科学, 2006 (4): 12-16
[38] Wildon D C, Thain J F, Minchin P H E, et al. Electrical signaling and systemic proteinase inhibitor induction in the wounded plant [J]. Nature,1992,360:62-65
[39] Uknes S, Mauch-Mani B, Moyer M ,et al. Acquired resistance in Arabidopsis [J]. Plant Cell,1992,4: 645- 655
[40] Sequeira L. Mechanisms of induced resistance in plants [J]. Annu Rev Microbiol.1986,37:51 -79
[41] 鲍丽丽,刘凤丽,范志金. 具有植物诱导抗病活性的先导化合物及其结构修饰[J]. 农药学学报,2005, 7(3):201-209
[42] Delaney T P, Ukness, Vernooij B, et al. A central role of salicylic acid in plant disease resistance [J]. Science, 1994,266(5188):1247-1250
[43] 侯 珲,朱建兰,周红平,等.BTH 和水杨酸(SA)对甜瓜抗白粉病的诱导作用[J].果树学报, 2006,23(5):736-739
[44] 李淑菊,马德华,庞金安,等.水杨酸对黄瓜几种酶活性及抗病性的诱导作用[J].华北农学报, 2000, 15(2):118-122
[45] OkunoT, Nakyama M, Okajima N, et al. Systemic resistance to down mildew and appearance of acid soluble proteins in cucumber leaves treated with biotic and abiotic inducers[J].Ann Phytopath Soc Japan,1991, 57: 202- 211
[46] Bol J F, van kan JA. The Synthesis and possible functions of virus –induced proteins in plants[J].Microbiol. Sci1, 1988,5(2):47
[47] Perumal V, Jay S, Levespue C, et al. A role for jasmonate in pathogen defense of Arabidopsis[J]. Proceedings of the National Academy of Sciences of the USA, 1998, 95(12): 7209-7214
[48] Neto G C, Kono Y, Hyakutake H, et al. Isolation and identification of jasmonic acid from wild rice as an antifungal substance[J].Agricultural and Biological Chemistry, 1991, 55: 3097-3098
[49] Thaler J S, Owen B, Higgins V J. The Role of the Jasmonate response in plant susceptibility to diverse pathogens with a range of lifestyles [J]. Plant Physiology, 2004, 135(1): 530-538
[50] Kogel K H, Beckhove U, Dreschers J. et al .Acquired resistance in barley-the resistance mechanism induced by 2,6-dichloroisonicotinic acid is a phenocopy of a genetically based mechanism governing race specific powdery mildew resistance[J]. Plant Physiol, 1994, 106:1269-1277
[51] Dann E, Diers B, Byrum J, et al. Effect of treating soybean with 2,6-dichloroisonicotinic acid(INA) and benzothiadiazole(BTH) on seed yields and the level of disease caused by Sclerotinia sclerotiorum in field and greenhouse studies[J].Euro J Plant Pathol, 1998, 104: 271-278
[52] 程智慧,李玉红,孟焕文,等.BTH 诱导黄瓜幼苗对霜霉病的抗性与细胞壁HRGP和木质素含量的关系[J].中国农业科学,2006,39(5):935-940
[53] Buonaurio R, Scarponi L, Ferrara M, et al. Induction of systemic acquired resistance in pepper plants by acibenzolar-S-methyl against bacterial spot disease[J]. European Journal of Plant Pathology,2002, 108 (1): 41-49
[54] Eikemo H, Stensvand A, Tronsmo A M. Induced resistance as a possible means to control diseases of strawberry caused by Phytophthora spp[J]. Plant Disease, 2003, 87(4): 345-350
[55] Perez L, Rodriguez M E, Rodriguez F, et al. Efficacy of acibenzolar-S-methyl, an inducer of systemic acquired resistance against tobacco blue mould caused by Peronospora hyoscyami f. sp. tabacina[J]. Crop Protection, 2003, 22: 405-413
[56] 王晓军,徐尚成,肖国民.植物激活剂的研究开发进展[J].化工纵横,2002,(1):22-26
[57] Hammerschmidt R, Lamport D T A, Muldoon E P. Cell wall hydroxyproline enhancement and lignin deposition as an early event in the resistance of cucumber to Cladosporium cucumerinum [J]. Physiol Plant Pathol,1984,24:43-47
[58] Kovats K, Binder A, Hohl H R. Cytology of induced systemic resistance of cucumber to Colletotrichum lagenarium [J].Planta,1991,183:484-490
[59] Bradley D J, Kjellbom P, Lamb C J. Elicitor-induced and wound-induced oxidative cross-linking of a proline-rich plant cell wall protein-a novel, rapid defense response[J]. Cell, 1992,70:21-30
[60] Van’t Slot K A E, Knogge W. A dual role for microbial pathogen-derived effector proteins in plant disease and resistance [J]. Critical Reviews in Plant Sciences, 2002, 21(3): 229-271
[61] 李振歧主编.植物免疫学[M].北京:中国农业出版社, 1995
[62] Kosuge T. The role of phenolics in host response to infection [J]. Annu Rev Phytopathol, 1969,7:105-122
[63] Cramer C L, Ryder T B, Bell I N, et al. Rapid switching of plant gene expression induced by fungus elicitor [J]. Science, 1985,227:1240-1243
[64] Friend J, Reynolds S B, Aveyard M A. Phenylalanine ammonia-lyase、Chlorogenic acid and lignin in potato tuber tissue inoculated with Phytophthora infestans [J]. Physiol Mal Plant Pathol,1973,3: 495- 507
[65] 王雅平.小麦对赤霉病抗性不同品种SOD活性[J].植物生理学报,1993, 19 (4):353-358
[66] 陈利锋,宋玉立,徐雍皋.抗感赤霉病小麦品种超氧化物歧化酶和过氧化氢酶的活性比较[J].植物病理学报,1997,27(3):209-213
[67] 朱有林,刘纪麟.受玉米大斑病菌侵染后玉米抗感近位等基因系SOD动态变化的研究[J]植物病理学报,1996,26(2): 133-137
[68] 曾永三, 王振中.活性氧和超氧化物岐化酶在植物抗病反应中的作用[J]. 仲恺农业技术学院学报, 1999,12(4):55-63
[69] 秦引萍,杨依军,彭友良等.稻瘟菌侵染诱导性水稻脂氧合酶的同工酶鉴定及性状分析[J].农业生物技术学报,1996,4(2):180-189
[70] 杨依军,秦引萍,黄俊斌等.稻瘟菌诱导性稻叶脂氧合酶的进一步纯化及抗体制备[J].植物病理学报,1997,27(3):243-249
[71] 方仲达.植病研究法[M].北京:中国农业出版社,1998
[72] 陈荣丽.BION 对小麦抗叶锈病性的诱导及生化机制研究[D].学位论文,2005,四川农业大学:20
[73] 史益敏.几丁质酶酶活性的测定[A].现代植物生理学实验指南[C],中国科学院上海植物生理研究所主编,北京:科学出版社,1999. 129-130
[74] 史益敏. β-1,3-葡聚糖酶活性的测定[A].现代植物生理学实验指南[C],中国科学院上海植物生理研究所主编.北京:科学出版社.1999. 128-129
[75] X.H. 波钦诺克 , 植物生物化学分析方法 ( 荆家海 , 丁钟荣 译 ) [M]. 北京 : 科学出版社,1981,178-181
[76] 胡景江, 朱 玮, 文建雷.杨树细胞壁HRGP和木质素的诱导积累与其对溃疡病抗性的关系[J].植物病理学报, 1999, 29(2): 151-156
[77] 李建华,李雄彪.细胞壁的制备及其羟脯氨酸含量的测定[J].植物生物学通讯, 1993,29 (5): 363-365
[78] 李伟,熊谨,陈晓阳.木质素代谢的生理意义及其遗传控制研究进展[[J].西北植物学报,2003,234:675-681
[79] Nicole Benhamou, Richard R Belanger. Benzothiadiazole-mediated induced resistance to Fusarium oxysporum f. sp. Radicis-lycopersici in tomato[J].Plant Physiology, 1998, 118(4): 1203-1213
[80] 王晨芳.化学诱导剂在诱导黄瓜抗病性中的作用[D].学位论文,2005,西北农林科技大学
[81] Nicole Benhamou, Richard R Belanger. Induction of systemic resistance to Pythium damping-off in cucumber plants by benzothiadiazole: Ultrastructure and cytochemistry of the host response [J].Plant J,1998,14:13-21
[82] Xuei X.L., Jarlfors U, Kuc J. Ultrastructural changes associated with induced systemic resistance of cucumber to disease: host response and development of colletotrichum lagenarium in systemically protected leaves[J]. Can.J.Bot.,1988, 66: 1028-1038
[83] 王颖, 景耀. 植物诱导抗病机理的研究进展[J].西北林学院学报,1997,12(3):52-57
[84] 程智慧,李玉红,孟焕文,等.BTH 诱导黄瓜幼苗对霜霉病的抗性与细胞壁HRGP和木质素含量的关系[J].中国农业科学,2006,39(5):935-940
[85] 葛秀春,宋凤鸣,陈永叶,等.苯并噻二唑诱发水稻对稻瘟病抗性中防卫相关酶活性的变化.中国水稻科学[[J].2002, 16(2):171-175
[86] Selitrennikoff C P. Antifungal proteins[J]. Appl Environ Microbiol, 2001,67:2883-2894
[87] Ye X S, Pan S Q, Kuc K J. Activity, isozyme pattern, and cellular localization of peroxidase as related to systemic resistance of tobacco to blue mold (Peronospora tabacina) and to tobacco moasaic virus [J]. Phytopathology, 1990, 80:1295-1299
[88]宾金华,姜 胜,黄胜琴,潘瑞炽.茉莉酸甲酯诱导烟草幼苗抗炭疽病与PAL活性及细胞壁 物质的关系[J].植物生理学报,2000,26(1):1-6