寡糖生物防治应用及机理研究进展

doi:10.16409/j.cnki.2095-039x.2015.05.016

摘要/Abstract

摘要： 植物诱导防御是当前国际生物防治学的热点内容,研究发现寡糖类物质是一类有效的诱导子,对于植物具有免疫调节作用,可以增强植物应对病虫害的能力。利用其进行植物病虫害防治是植物保护的新途径。本文将对目前应用较为广泛的寡糖诱导子来源和实际应用情况,以及寡糖诱抗机理研究中寡糖识别机制的研究进展进行综述。

关键词: 寡糖, 生物防治, 植物病害, 植物免疫, 受体

Abstract: Plant induced defense is a current hot topic of biocontrol research in the world. Oligosaccharides is a kind of elicitor, it has been shown a wide range of biological applications in plant immunity regulation and defense induction against pathogen. Employing oligosaccahrides induced resistance for sustainable plant protection against pathogens is a new strategy in crop protection. This paper summarizes the origin of oligosaccharides which are widely used at present and their practical application in different crops. Recent advances in oligosaccahrides signal perception are also presented in this review.

Key words: oligosacchartides, biocontrol, plant disease, plant immunity, receptor

中图分类号:

S476

王文霞, 赵小明, 杜昱光, 尹恒. 寡糖生物防治应用及机理研究进展[J]. 中国生物防治学报, 2015, 31(5): 757-769.

WANG Wenxia, ZHAO Xiaoming, DU Yuguang, YIN Heng. Progress on Application and Mechanism of Oligosaccahrides in Plant Biocontrol Response[J]. journal1, 2015, 31(5): 757-769.

参考文献

[1] Jones J D G, Dangl J L. The plant immune system[J]. Nature, 2006, 444(7117): 323-329.
[2] Ayers A R, Ebel J, Finelli F, et al. Host-pathogen interactions: Ⅸ. quantitative assays of elicitor activity and characterization of elicitor present in extracellular medium of cultures of Phytophthora Megasperma var. sojae[J]. Plant Physiology, 1976, 57(5): 751-759.
[3] Shibuya N, Minami E. Oligosaccharide signalling for defence responses in plant[J]. Physiological and Molecular Plant Pathology, 2001, 59(5): 223-233.
[4] Okinaka Y, Mimori K, Takeo K, et al. A structural model for the mechanisms of elicitor release from fungal cell-walls by plant beta-1,3-endoglucanase[J]. Plant Physiology, 1995, 109(3): 839-845.
[5] Waldmuller T, Cosio E G, Grisebach H, et al. Release of highly elicitor-active glucans by germinating zoospores of Phytophthora megasperma f. sp. glycinea[J]. Planta, 1992, 188(4): 498-505.
[6] Kurita K. Chitin and chitosan: functional biopolymers from marine crustaceans[J]. Marine Biotechnology, 2006, 8(3): 203-226.
[7] Sharp J K, Valent B, Albersheim P. Host-pathogen interactions: 26. purification and partial characterization of a beta-glucan fragment that elicits phytoalexin accumulation in soybean[J]. Journal of Biological Chemistry, 1984, 259(18): 1312-1320.
[8] Klarzynski O, Plesse B, Joubert J M, et al. Linear beta-1,3 glucans are elicitors of defense responses in tobacco[J]. Plant Physiology, 2000, 124(3): 1027-1037.
[9] Ridley B L, O'Neill M A, Mohnen D A. Pectins: structure, biosynthesis, and oligogalacturonide-related signaling[J]. Phytochemistry, 2001, 57(6): 929-967.
[10] Rodriguez A T, Ramirez M A, Cardenas R M, et al. Induction of defense response of Oryza sativa L. against Pyricularia grisea (Cooke) Sacc. by treating seeds with chitosan and hydrolyzed chitosan[J]. Pesticide Biochemistry and Physiology, 2007, 89(3): 206-215.
[11] 白春微, 蒋选利. 几种诱导因子对水稻纹枯病的诱导抗病性研究[J]. 江苏农业科学, 2012, 40(11): 116-118.
[12] Burkhanova G F, Yarullina L G, Maksimov I V. The control of wheat Defense responses during infection with Bipolaris sorokiniana by chitooligosaccharides[J]. Russian Journal of Plant Physiology, 2007, 54(1): 104-110.
[13] 王正明, 陈明, 王文刚. 几种植物诱抗剂对小麦纹枯病防治效果研究[J]. 湖北植保, 2012, 6: 18-19.
[14] Sharathchandra R G, Raj S N, Shetty N P, et al. A chitosan formulation elexa (TM) induces downy mildew disease resistance and growth promotion in pearl millet[J]. Crop Protection, 2004, 23(10): 881-888.
[15] Vasyukova N I, Zinov'eva S V, Il'inskaya L I, et al. Modulation of plant resistance to diseases by water-soluble chitosan[J]. Applied Biochemistry and Microbiology, 2001, 37(1): 103-109.
[16] 孔德英, 肖崇刚. 氨基寡糖素对番茄青枯病防治作用[J]. 西南农业大学学报(自然科学版), 2005, 3: 327-330.
[17] 乔康, 姬小雪, 邱士芬. 氨基寡糖素2%水剂防治番茄病毒病试验[J]. 农药科学与管理, 2012, 5: 59-60.
[18] 楚书丽. 0.5%氨基寡糖素水剂防治番茄晚疫病田间药效试验[J]. 河南农业, 2013, 5: 32, 37.
[19] 肖征军. 5%氨基寡糖素在番茄上的应用试验[J]. 南方园艺, 2014, 2: 19-21.
[20] 檀志全, 谭海文, 覃保荣, 等. 5%氨基寡糖素AS在番茄上的应用效果初探[J]. 中国植保导刊, 2013, 10: 65-66.
[21] Chirkov S N, Il'ina A V, Surgucheva N A, et al. Effect of chitosan on systemic viral infection and some defense responses in potato plants[J]. Russian Journal of Plant Physiology, 2001, 48(6): 774-779.
[22] Ben-Shalom N, Ardi R, Pinto R, et al. Controlling gray mould caused by Botrytis cinerea in cucumber plants by means of chitosan[J]. Crop Protection, 2003, 22(2): 285-290.
[23] Zinov'eva S V, Perekhod E A, Il'ina A V, et al. PR proteins in plants infested with the root-knot nematode Meloidogyne incognita (Kofoid et White, 1919) Chitwood 1949[J]. Doklady Biological Sciences, 2001, 379(1-6): 393-395.
[24] 郭志刚. 氨基寡糖素(0.5%水剂)对黄瓜根结线虫病的防治试验[J]. 农业科技通讯, 2014, 3: 123-124.
[25] 解婷婷, 吕淑霞, 马镝, 等. 壳寡糖诱导黄瓜抗黄瓜黑星病的初步研究[J]. 天然产物研究与开发, 2011, 3: 508-511.
[26] 杜昱光, 白雪芳. 甲壳素生物降解制备低聚氨基葡萄糖[J]. 精细与专用化学品, 2003, 7: 16-18.
[27] Molloy C, Cheah L H, Koolaard J P. Induced resistance against Sclerotinia sclerotiorum in carrots treated with enzymatically hydrolysed chitosan[J]. Postharvest Biology and Technology, 2004, 33(1): 61-65.
[28] Xu J, Zhao X, Han X, et al. Antifungal activity of oligochitosan against Phytophthora capsici and other plant pathogenic fungi in vitro[J]. Pesticide Biochemistry and Physiology, 2007, 87(3): 220-228.
[29] 赵小明, 杜昱光, 白雪芳. 氨基寡糖素诱导作物抗病毒病药效试验[J]. 中国农学通报, 2004, 4: 245-247.
[30] Hadwiger L A, Beckman J M. Chitosan as a component of pea-Fusarium solani interactions[J]. Plant Physiology, 1980, 66(2): 205-211.
[31] 李萍, 张善学, 李国梁, 等. 氨基寡糖素在豇豆上的应用效果[J]. 中国植保导刊, 2013, 7: 48-51.
[32] Trotel-Aziz P, Couderchet M, Vernet G, et al. Chitosan stimulates defense reactions in grapevine leaves and inhibits development of Botrytis cinerea[J]. European Journal of Plant Pathology, 2006, 114(4): 405-413.
[33] 徐作珽, 李林, 李长松, 等. 中生菌素和氨基寡糖素对西瓜枯萎病防治试验[J]. 中国蔬菜, 2003, 3: 12-14.
[34] Falcon A B, Cabrera J C, Costales D, et al. The effect of size and acetylation degree of chitosan derivatives on tobacco plant protection against Phytophthora parasitica nicotianae[J]. World Journal of Microbiology and Biotechnology, 2008, 24(1): 103-112.
[35] 尹恒, 王文霞, 卢航, 等. 壳寡糖诱导油菜抗菌核病机理研究初探[J]. 西北农业学报, 2008, 5: 81-85.
[36] 王尽松, 徐荣燕. 5%氨基寡糖素增强棉花抗病性试验及增产效果[J]. 中国棉花, 2014, 8: 26-27.
[37] 蒋林, 周兴挺, 梁伟军, 等. 氨基寡糖素对芦荟炭疽病防治效果及安全性实验[J]. 现代中药研究与实践, 2005, 5: 14-16.
[38] Li S, Zhu T. Biochemical response and induced resistance against anthracnose (Colletotrichum camelliae) of camellia (Camellia pitardii) by chitosan oligosaccharide application[J]. Forest Pathology, 2013, 43(1): 67-76.
[39] Ma Z, Yang L, Yan H, et al. Chitosan and oligochitosan enhance the resistance of peach fruit to brown rot[J]. Carbohydrate Polymers, 2013, 94(1): 272-277.
[40] 赵小明, 李东鸿, 杜昱光, 等. 2%氨基寡糖防治苹果花叶病[J]. 植物保护, 2002, 5: 15-17.
[41] 卢学松, 张祥雄, 游泳, 等. 好普对西瓜三种主要病害的药效试验[J]. 福建农业科技, 2003, 1: 51-52.
[42] 贺春娟, 薛敏云. 5%氨基寡糖素AS在桃树上的应用效果[J]. 山西果树, 2014, 3: 3-5.
[43] 陆红霞, 周文静, 黄不谎, 等. 氨基寡糖素对贡柑抗病增产品质改善作用的研究[J]. 中国果菜, 2014, 6: 59-62.
[44] 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.
[45] 王正明, 陈明, 王文刚. 几种植物诱抗剂对小麦纹枯病防治效果研究[J]. 湖北植保, 2012, 6: 18-19.
[46] Ning W, Chen F, Mao B, et al. N-acetylchitooligosaccharides elicit rice defence responses including hypersensitive response-like cell death, oxidative burst and defence gene expression[J]. Physiological and Molecular Plant Pathology, 2004, 64(5): 263-271.
[47] Falcon-Rodriguez A B, Costales D, Cabrera J C, et al. Chitosan physico-chemical properties modulate defense responses and resistance in tobacco plants against the oomycete Phytophthora nicotianae[J]. Pesticide Biochemistry and Physiology, 2011, 100(3): 221-228.
[48] 高智谋, 沈奕, 王革, 等. 几丁寡糖对烟草赤星病的控制效应及其机制[J]. 热带作物学报, 2009, 8: 1132-1137.
[49] 王正明, 曾玲玲, 檀银忠, 等. 几种植物诱抗剂对棉花黄萎病的防治效果[J]. 棉花科学, 2012, 6: 24-25.
[50] 周锋, 李金丽, 梁宏杰, 等. 0.5%几丁聚糖FS对油菜菌核病的田间防效[J]. 中国植保导刊, 2014, 1: 68-70.
[51] Bell A A, Hubbard J C, Liu L, et al. Effects of chitin and chitosan on the incidence and severity of Fusarium yellows of celery[J]. Plant Disease, 1998, 82(3): 322-328.
[52] Renard-Merlier D, Randoux B, Nowak E, et al. Iodus 40, salicyclic acid, heptanoyl salicylic acid and trehalose exhibit different efficacies and defence targets during a wheat/powdery mildew interaction[J]. Phytochemistry, 2007, 68(8): 1156-1164.
[53] Li J, Zhu L, Lu G, et al. Curdlan beta-1,3-glucooligosaccharides induce the defense responses against Phytophthora infestans infection of potato (Solanum tuberosum L. cv. McCain G1) leaf cells[J]. PLoS ONE, 2014, 9(5): e97197.
[54] Wolski E A, Maldonado S, Daleo G R, et al. A novel alpha-1,3-glucan elicits plant defense responses in potato and induces protection against Rhizoctonia solani AG-3 and Fusarium solani f. sp. eumartii[J]. Physiological and Molecular Plant Pathology, 2006, 69(1-3): 93-103.
[55] Fu Y B, Yin H, Wang W X, et al. Beta-1,3-glucan with different degree of polymerization induced different defense responses in tobacco[J]. Carbohydrate Polymers, 2011, 86(2): 774-782.
[56] 景岚, 李凌欣, 裴旭, 等. 寡糖诱导向日葵抗锈病的信号转导途径[J]. 中国油料作物学报, 2012, 5: 523-527.
[57] Hao Y H, Wu C F, Zhao D W, et al. Proteomic analysis of glucohexaose induced resistance to downy mildew in cucumis sativus[J]. Australian Journal of Crop Science, 2013, 7(9): 1242-1251.
[58] Aziz A, Poinssot B, Daire X, et al. Laminarin elicits defense responses in grapevine and induces protection against Botrytis cinerea and Plasmopara viticola[J]. Molecular Plant-Microbe Interactions, 2003, 16(12): 1118-1128.
[59] Randoux B, Renard-Merlier D, Mulard G, et al. Distinct defenses induced in wheat against powdery mildew by acetylated and nonacetylated oligogalacturonides[J]. Phytopathology, 2010, 100(12): 1352-1363.
[60] Moerschbacher B M, Mierau M, Graessner B, et al. Small oligomers of galacturonic acid are endogenous suppressors of disease resistance reactions in wheat leaves[J]. Journal of Experimental Botany, 1999, 50(334): 605-612.
[61] 赵小明, 杜昱光, 白雪芳, 等. 中科2号防治辣椒病毒病药效试验//新世纪（首届）全国绿色环保农药技术论坛暨产品展示会[C]. 苏州, 2002, 2.
[62] 卢航, 赵小明, 白雪芳, 等. 寡聚半乳糖醛酸诱导烟草抗烟草花叶病毒研究初探[J]. 植物保护, 2008, 34(3): 38-41.
[63] Aziz A, Heyraud A, Lambert B. Oligogalacturonide signal transduction, induction of defense-related responses and protection of grapevine against Botrytis cinerea[J]. Planta, 2004, 218(5): 767-774.
[64] 赵小明, 杜昱光, 白雪芳. 寡聚半乳糖醛酸防治苹果花叶病田间药效试验[J]. 中国农学通报, 2004, 6: 262-264.
[65] 李瑞环, 周一万, 冯俊涛, 等. 4.5%丙·壳寡糖可溶性液剂对番茄灰霉病的控制效果[J]. 西北农业学报, 2012, 10: 169-173.
[66] 刘晓帆, 亓晓光, 郭小珍, 等. “寡糖.吗啉胍”防治辣椒病毒病田间药效试验[J]. 农业开发与装备, 2013, 2: 91.
[67] 黄晓妹, 卢行尚. 5%氨基寡糖素对香蕉黑星病的抗病增产效果[J]. 广西植保, 2014, 2: 7-9.
[68] 朱英波, 史凤玉, 张瑞敬, 等. 壳寡糖和钕复合处理诱导黄瓜对枯萎病的抗性[J]. 中国生物防治学报, 2014, 4: 528-533.
[69] van Aubel G, Buonatesta R, van Cutsem P. COS-OGA: A novel oligosaccharidic elicitor that protects grapes and cucumbers against powdery mildew[J]. Crop Protection, 2014, 65: 129-137.
[70] Rabea E I, Badawy M E T, Stevens C V, et al. Chitosan as antimicrobial agent: applications and mode of action[J]. Biomacromolecules, 2003, 4(6): 1457-1465.
[71] Ryan C A. Oligosaccharide signaling in plants[J]. Glycoconjugate Journal, 1988, 5(3): 359.
[72] Prome J C. Signalling events elicited in plants by defined oligosaccharide structures[J]. Current Opinion in Structural Biology, 1996, 6(5): 671-678.
[73] Ryan C A, Farmer E E. Oligosaccharide signals in plants: a current assessment[J]. Annual Review of Plant Physiology and Plant Molecular Biology, 1991, 42: 651-674.
[74] Yin H, Zhao X, Du Y. Oligochitosan: a plant diseases vaccine-A review[J]. Carbohydrate Polymers, 2010, 82(1): 1-8.
[75] Trouvelot S, Heloir M C, Poinssot B, et al. Carbohydrates in plant immunity and plant protection: roles and potential application as foliar sprays[J]. Frontiers in Plant Science, 2014, 5: 592.
[76] Shibuya N, Kaku H, Kuchitsu K, et al. Identification of a novel high-affinity binding site for N-acetylchitooligosaccharide elicitor in the membrane fraction from suspension-cultured rice cells[J]. Febs Letters, 1993, 329(1-2): 75-78.
[77] Shibuya N, Ebisu N, Kamada Y, et al. Localization and binding characteristics of a high-affinity binding site for N-acetylchitooligosaccharide elicitor in the plasma membrane from suspension-cultured rice cells suggest a role as a receptor for the elicitor signal at the cell surface[J]. Plant and Cell Physiology, 1996, 37(6): 894-898.
[78] Yamaguchi T, Ito Y, Shibuya N. Oligosaccharide elicitors and their receptors for plant defense responses[J]. Trends in Glycoscience and Glycotechnology, 2000, 12(64): 113-120.
[79] Ito Y, Kaku H, Shibuya N. Identification of a high-affinity binding protein for N-acetylchitooligosaccharide elicitor in the plasma membrane of suspension-cultured rice cells by affinity labeling[J]. Plant Journal, 1997, 12(2): 347-356.
[80] Okada M, Matsumura M, Shibuya N. Identification of a high-affinity binding protein for N-acetylchitooligosaccharide elicitor in the plasma membrane from rice leaf and root cells[J]. Journal of Plant Physiology, 2001, 158(1): 121-124.
[81] Okada M, Matsumura M, Ito Y, et al. High-affinity binding proteins for N-acetylchitooligosaccharide elicitor in the plasma membranes from wheat, barley and carrot cells: conserved presence and correlation with the responsiveness to the elicitor[J]. Plant and Cell Physiology, 2002, 43(5): 505-512.
[82] Kaku H, Nishizawa Y, Ishii-Minami N, et al. Plant cells recognize chitin fragments for defense signaling through a plasma membrane receptor[J]. Proceedings of the National Academy of Sciences of the United States of America, 2006, 103(29): 11086-11091.
[83] Lohmann G V, Shimoda Y, Nielsen M W, et al. Evolution and regulation of the Lotus japonicus LysM receptor gene family[J]. Molecular Plant-Microbe Interactions, 2010, 23(4): 510-521.
[84] Hayafune M, Berisio R, Marchetti R, et al. Chitin-induced activation of immune signaling by the rice receptor CEBiP relies on a unique sandwich-type dimerization[J]. Proceedings of the National Academy of Sciences of the United States of America, 2014, 111(3): 404-413.
[85] Miya A, Albert P, Shinya T, et al. CERK1, a LysM receptor kinase, is essential for chitin elicitor signaling in Arabidopsis[J]. Proceedings of the National Academy of Sciences of the United States of America, 2007, 104(49): 19613-19618.
[86] Wan J, Zhang X, Neece D, et al. A LysM receptor-like kinase plays a critical role in chitin signaling and fungal resistance in Arabidopsis[J]. Plant Cell, 2008, 20(2): 471-481.
[87] Shimizu T, Nakano T, Takamizawa D, et al. Two LysM receptor molecules, CEBiP and OsCERK1, cooperatively regulate chitin elicitor signaling in rice[J]. Plant Journal, 2010, 64(2): 204-214.
[88] Iizasa E, Mitsutomi M, Nagano Y. Direct binding of a plant LysM receptor-like kinase, LysM RLK1/CERK1, to chitin in vitro[J]. Journal of Biological Chemistry, 2010, 285(5): 2996-3004.
[89] Petutschnig E K, Jones A M E, Serazetdinova L, et al. The Lysin motif receptor-like kinase (LysM-RLK) CERK1 is a major chitin-binding protein in Arabidopsis thaliana and subject to chitin-induced phosphorylation[J]. Journal of Biological Chemistry, 2010, 285(37): 28902-28911.
[90] Shinya T, Motoyama N, Ikeda A, et al. Functional characterization of CEBiP and CERK1 homologs in Arabidopsis and rice reveals the presence of different chitin receptor systems in plants[J]. Plant and Cell Physiology, 2012, 53(10): 1696-1706.
[91] Liu T, Liu Z, Song C, et al. Chitin-induced dimerization activates a plant immune receptor[J]. Science, 2012, 336(6085): 1160-1164.
[92] Wan J, Tanaka K, Zhang X, et al. LYK4, a Lysin motif receptor-like kinase, is important for chitin signaling and plant innate immunity in Arabidopsis[J]. Plant Physiology, 2012, 160(1): 396-406.
[93] Cao Y, Liang Y, Tanaka K, et al. The kinase LYK5 is a major chitin receptor in Arabidopsis and forms a chitin-induced complex with related kinase CERK1[J]. Elife, 2014, 3: e03766.
[94] Yoshikawa M, Keen N T, Wang M C. A receptor on soybean membranes for a fungal elicitor of phytoalexin accumulation[J]. Plant Physiology, 1983, 73(2): 497-506.
[95] Cosio E G, Frey T, Ebel J. Solubilization of soybean membrane-binding sites for fungal beta-glucans that elicit phytoalexin accumulation[J]. Febs Letters, 1990, 264(2): 235-238.
[96] Cheong J J, Hahn M G. A specific, high-affinity binding site for the hepta-beta-glucoside elicitor exists in soybean membranes[J]. Plant Cell, 1991, 3(2): 137-147.
[97] Cosio E G, Frey T, Ebel J. Identification of a high-affinity binding protein for a hepta-beta-glucoside phytoalexin elicitor in soybean[J]. European Journal of Biochemistry, 1992, 204(3): 1115-1123.
[98] Cheong J J, Alba R, Cote F, et al. Solubilization of functional plasma membrane-localized hepta-beta-glucoside elicitor-binding proteins from soybean[J]. Plant Physiology, 1993, 103(4): 1173-1182.
[99] Mithofer A, Lottspeich F, Ebel J. One-step purification of the beta-glucan elicitor-binding protein from soybean (Glycine max L.) roots and characterization of an anti-peptide antiserum[J]. Febs Letters, 1996, 381(3): 203-207.
[100] Umemoto N, Kakitani M, Iwamatsu A, et al. The structure and function of a soybean beta-glucan-elicitor-binding protein[J]. Proceedings of the National Academy of Sciences of the United States of America, 1997, 94(3): 1029-1034.
[101] Fliegmann J, Mithofer A, Wanner G, et al. An ancient enzyme domain hidden in the putative beta-glucan elicitor receptor of soybean may play an active part in the perception of pathogen-associated molecular patterns during broad host resistance[J]. Journal of Biological Chemistry, 2004, 279(2): 1132-1140.
[102] Fliegmann J, Montel E, Djulic A, et al. Catalytic properties of the bifunctional soybean beta-glucan-binding protein, a member of family 81 glycoside hydrolases[J]. Febs Letters, 2005, 579(29): 6647-6652.
[103] Leclercq J, Fliegmann J, Tellstrom V, et al. Identification of a multigene family encoding putative beta-glucan-binding proteins in Medicago truncatula[J]. Journal of Plant Physiology, 2008, 165(7): 766-776.
[104] Horn M A, Heinstein P F, Low P S. Receptor mediated endocytosis in plant cells[J]. Plant Cell, 1989, 1(10): 1003-1009.
[105] He Z H, Fujiki M, Kohorn B D. A cell wall-associated, receptor-like protein kinase[J]. Journal of Biological Chemistry, 1996, 271(33): 19789-19793.
[106] Decreux A, Messiaen J. Wall-associated kinase WAK1 interacts with cell wall pectins in a calcium-induced conformation[J]. Plant and Cell Physiology, 2005, 46(2): 268-278.
[107] Brutus A, Sicilia F, Macone A, et al. A domain swap approach reveals a role of the plant wall associated kinase 1 (WAK1) as a receptor of oligogalacturonides[J]. Proceedings of the National Academy of Sciences of the United States of America, 2010, 107(20): 9452-9457.
[108] Anderson C M, Wagner T A, Perret M, et al. WAKs: cell wall-associated kinases linking the cytoplasm to the extracellular matrix[J]. Plant Molecular Biology, 2001, 47(1-2): 197-206.
[109] Wagner T A, Kohorn B D. Wall-associated kinases are expressed throughout plant development and are required for cell expansion[J]. Plant Cell, 2001, 13(2): 303-318.
[110] Kohorn B D, Kobayashi M, Johansen S, et al. Wall-associated kinase 1 (WAK1) is crosslinked in endomembranes, and transport to the cell surface requires correct cell-wall synthesis[J]. Journal of Cell Science, 2006, 119(11): 2282-2290.
[111] Cabrera J C, Boland A, Messiaen J, et al. Egg box conformation of oligogalacturonides: The time-dependent stabilization of the elicitor-active conformation increases its biological activity[J]. Glycobiology, 2008, 18(6): 473-482.
[112] Kohorn B D, Johansen S, Shishido A, et al. Pectin activation of MAP kinase and gene expression is WAK2 dependent[J].Plant Journal, 2009, 60(6): 974-982.
[113] Kohorn B D, Kohorn S L, Saba N J, et al. Requirement for pectin methyl esterase and preference for fragmented over native pectins for wall associated kinase activated, EDS1/PAD4-dependent stress response in Arabidopsis[J]. Journal of Biological Chemistry, 2014, 289(27): 18978-18986.
[114] Zhao X M, She X P, Yu W, et al. Effects of oligochitosans on tobacco cells and role of endogenous nitric oxide burst in the resistance of tobacco to Tobacco mosaic virus[J]. Journal of Plant Pathology, 2007, 89(1): 55-65.
[115] 赵小明. 壳寡糖诱导植物抗病性及其诱抗机理的初步研究[D]. 大连: 中国科学院大连化学物理研究所, 2006.
[116] Lienart Y, Gautier C, Domard A. Isolation from rubus cell suspension cultures of a lectin specific for glucosamine oligomers[J]. Planta, 1991, 184(1): 8-13.
[117] Chen H P, Xu L L. Isolation and characterization of a novel chitosan-binding protein from non-heading Chinese cabbage leaves[J]. Journal of Integrative Plant Biology, 2005, 47(4): 452-456.
[118] Cabrera J C, Boland A, Cambier P, et al. Chitosan oligosaccharides modulate the supramolecular conformation and the biological activity of oligogalacturonides in Arabidopsis[J]. Glycobiology, 2010, 20(6): 775-786.