油菜素内酯和寡糖素提高杨梅对凋萎病抗性的机理初探

doi:10.16420/j.issn.0513-353x.2022-0873

摘要/Abstract

摘要：

为探究免疫诱导剂油菜素内酯和寡糖素对杨梅凋萎病的抗性诱导机理，以‘东魁’杨梅幼苗为试材，研究外源诱导剂对接种凋萎病菌后杨梅的抗性、生长指标、叶绿素含量、抗氧化酶活性及病程相关基因表达的影响。结果表明：2,4-表油菜素内酯和寡糖 · 链蛋白可显著降低杨梅凋萎病病情指数，抗性诱导效果分别达47.94%和39.01%；两者均能有效促进杨梅幼苗生长发育，处理后的杨梅株高、叶长和叶面积均高于对照；在接种后28 d，2,4-表油菜素内酯和寡糖 · 链蛋白处理显著提高了杨梅叶片叶绿素a、叶绿素b和类胡萝卜素含量，较对照提高13.9% ~ 26.8%。在2,4-表油菜素内酯和寡糖 · 链蛋白处理下，SOD、POD和CAT酶活性短时间内显著升高，28 d时2,4-表油菜素内酯处理分别比对照提高87.4%、106.5%和77.9%，而MDA含量显著低于对照。2,4-表油菜素内酯和寡糖 · 链蛋白均能显著诱导杨梅病程相关基因MrPR1、MrPR2和MrPR10上调表达，在接种后7 ~ 28 d仍维持在较高表达水平。综上，2,4-表油菜素内酯和寡糖 · 链蛋白对杨梅的凋萎病抗性诱导效果较好，其诱导机理可能与抗氧化酶活性的增强以及病程相关基因的上调表达密切相关。

关键词: 杨梅, 诱导剂, 油菜素内酯, 寡糖素, 凋萎病, 抗氧化酶活性, 病程相关蛋白

Abstract:

To explore the mechanism of induced resistance by immune inducers to Chinese bayberry （Myrica rubra）twig blight disease，young plants of‘Dongkui’were used as test materials to study the effects of exogenous inducers on the induction effect，growth index，chlorophyll content，antioxidant enzyme activity and expressions of pathogenesis-related genes in M. rubra after inoculation with twig blight strains. The results showed that 2,4-epibrassinolide（EBR）and oligosaccharins · plant activator protein（OPAP）could significantly reduce the twig blight disease index，with better effects of resistance induction（reaching 47.94% and 39.01%，respectively）. Both EBR and OPAP effectively promoted the growth of M. rubra，including plant height，leaf length and leaf area，which were higher than those of the control. Furthermore，EBR and OPAP significantly improved the contents of chlorophyll a，chlorophyll b and carotenoids on 28-day after inoculation，which were increased by 13.9%-26.8% compared with the control. Under the treatment of EBR and OPAP，the activities of SOD，POD and CAT significantly increased in a short time，and their activities were increased by 87.4%，106.5% and 77.9% compared with the control on 28-day after inoculation，while MDA content was lower than that of control. Both EBR and OPAP could significantly induce up-regulated expressions of M. rubra pathogenesis-related genes during 7-day to 28-day after inoculation，including MrPR1，MrPR2 and MrPR10，and their expressions were maintained at a high level. In conclusion，EBR and OPAP have better effects on the induced resistance to M. rubra twig blight disease，and their potential induction mechanisms may be closely related to the enhancement of antioxidant enzyme activity and the up-regulation of pathogenesis-related genes.

Key words: Chinese bayberry, inducer, brassinolide, oligosaccharin, twig blight disease, antioxidant enzyme activity, pathogenesis-related protein

俞浙萍, 严秀娟, 张淑文, 郑锡良, 孙鹂, 任海英, 梁森苗, 戚行江. 油菜素内酯和寡糖素提高杨梅对凋萎病抗性的机理初探[J]. 园艺学报, 2023, 50(10): 2229-2241.

YU Zheping, YAN Xiujuan, ZHANG Shuwen, ZHENG Xiliang, SUN Li, REN Haiying, LIANG Senmiao, QI Xingjiang. Preliminary Study on the Mechanism of Improvement of Resistance to Chinese Bayberry Twig Blight Disease by Brassinolide and Oligosaccharin[J]. Acta Horticulturae Sinica, 2023, 50(10): 2229-2241.

图/表 7

表1 用于qRT-PCR的引物序列

Table 1 Primer sequences for qRT-PCR

基因 Gene	上游引物（5′-3′） Forward primer	下游引物（5′-3′） Reverse primer
MrActin	AATGGAACTGGAATGGTCAAGGC	TGCCAGATCTTCTCCATGTCATCCCA
MrPR1	GCCTTGGTCCTTGATTCCGA	GGTCGAGCTCATCAACCCTT
MrPR2	GCTAACTGCCCTTCCGAGTT	AGCGTAACTATACGCGGTCG
MrPR10	GAACCGGCTGCAACTTTGAC	GACGACTAGTGGCACGTTGA

表2 诱导剂2,4-表油菜素内酯（EBR）和寡糖 · 链蛋白（OPAP）处理对杨梅凋萎病的影响

Table 2 Effects of 2,4-epibrassionolide（EBR）and oligosaccharins · plant activator protein（OPAP）treatments on twig blight disease in Chinese bayberry

诱导剂 Inducer	病情指数 Disease index	抗性诱导效果/% Resistance induction effect
对照 Control	54.42 ± 2.69 a	0
OPAP	33.19 ± 1.72 b	39.01 ± 2.27 b
EBR	28.33 ± 1.64 c	47.94 ± 2.56 a

表3 诱导剂2,4-表油菜素内酯（EBR）和寡糖 · 链蛋白（OPAP）处理对杨梅幼苗株高的影响

Table 3 Effects of 2,4-epibrassionolide（EBR）and oligosaccharins·plant activator protein（OPAP）treatment on plant height of Chinese bayberry cm

诱导剂 Inducer	接种凋萎病菌后天数/d Days after inoculation with pathogen of twig blight disease
诱导剂 Inducer	0	7	14	21	28
对照 Control	45.3 ± 2.8 a	47.7 ± 2.9 a	50.2 ± 3.7 b	55.8 ± 1.6 b	57.1 ± 3.6 b
OPAP	44.7 ± 2.2 ab	49.3 ± 3.7 a	52.5 ± 2.2 b	61.6 ± 1.7 ab	67.8 ± 2.1 ab
EBR	42.4 ± 1.7 b	49.8 ± 1.1 a	58.5 ± 0.9 a	64.7 ± 2.6 a	69.4 ± 2.8 a

图1 诱导剂2,4-表油菜素内酯（EBR）和寡糖 · 链蛋白（OPAP）处理对杨梅叶片生长的影响不同小写字母表示同一处理时间不同处理之间差异显著。下同。

Fig. 1 Effects of 2,4-epibrassionolide（EBR）and oligosaccharins · plant activator protein（OPAP）treatments on leaf growth of Chinese bayberry Different lowercase letters indicate significant differences among different treatments at same treatment time. The same below.

图2 诱导剂2,4-表油菜素内酯（EBR）和寡糖 · 链蛋白（OPAP）处理对杨梅叶片叶绿素和类胡萝卜素含量的影响

Fig. 2 Effects of 2,4-epibrassionolide（EBR）and oligosaccharins · plant activator protein（OPAP）treatments on chlorophyll content in Chinese bayberry leaves

图3 诱导剂2,4-表油菜素内酯（EBR）和寡糖 · 链蛋白（OPAP）处理对杨梅叶片SOD、POD、CAT活性和MDA含量的影响

Fig. 3 Effects of 2,4-epibrassionolide（EBR）and oligosaccharins · plant activator protein（OPAP）treatments on activity of SOD，POD，CAT and MDA content in Chinese bayberry leaves

图4 诱导剂2,4-表油菜素内酯（EBR）和寡糖 · 链蛋白（OPAP）处理对杨梅MrPR1、MrPR2和MrPR10基因表达的影响

Fig. 4 Effects of 2,4-epibrassionolide（EBR）and oligosaccharins · plant activator protein（OPAP）treatments on the expression of MrPR1，MrPR2 and MrPR10 in Chinese bayberry

参考文献 53

[1]	Breen S, Williams S J, Outram M, Kobe B, Solomon P S. 2017. Emerging insights into the functions of pathogenesis-related protein 1. Trends in Plant Science, 22 (10):871-879. doi: S1360-1385(17)30137-1 pmid: 28743380
[2]	Chen Li-min, Wu Qian-qian, He Tian-jun, Pan Yi-ming, Chen Ting-ting, Wu Quan-cong. 2021. The control effects of seed soaking with oligosaccharins·plant activator protein to bean fusarium root rot. Journal of Zhejiang Agricultural Sciences, 62 (10):2030-2033. (in Chinese)
	陈利民, 吴倩倩, 何天骏, 潘逸明, 陈婷婷, 吴全聪. 2021. 寡糖 · 链蛋白浸种对菜豆根腐病的防控效果. 浙江农业科学, 62 (10):2030-2033.
[3]	Chen Nai-qi, Long Chuan, Wang Jian-li. 2019. Effects of plant immune inducer 6% oligosaccharins · plant activator protein(atailin)on grape disease resistance,stress resistance,high quality and yield increase test. Yantai Fruits, 145 (1):19-21. (in Chinese)
	陈乃琦, 龙川, 王健立. 2019. 植物免疫诱抗剂6%寡糖 · 链蛋白(阿泰灵)对葡萄抗病抗逆优质增产试验. 烟台果树, 145 (1):19-21.
[4]	Chen X R, Xing Y P, Zhang T X, Zheng J T, Xu J Y, Wang Z R, Tong Y H, Xu Y J. 2012. Identification and characterization of Pestalotiopsis spp. causing leaf necrosis of red bayberry in China. Acta Phytopathologica Sinica, 42 (6):626-632.
[5]	Chen Z F, Wang Z, Yang Y G, Li M, Xu B C. 2018. Abscisic acid and brassinolide combined application synergistically enhances drought tolerance and photosynthesis of tall fescue under water stress. Scientia Horticulturae, 228:1-9. doi: 10.1016/j.scienta.2017.10.004 URL
[6]	Cheng G X, Li R J, Wang M, Huang L J, Khan A, Ali M, Gong Z H. 2018. Variation in leaf color and combine effect of pigments on physiology and resistance to whitefly of pepper(Capsicum annuum L.). Scientia Horticulturae, 229:215-225. doi: 10.1016/j.scienta.2017.11.014 URL
[7]	Ding Hai-dong, Liu Hui, Chen Yi, Wang Dan, Liang Jian-sheng. 2011. Involvements of H₂O₂ and mitogen-activated protein kinase signals in brassinosteroid-activated tomato antioxidant defense enzymes superoxide dismutase and catalase. Plant Physiology Journal, 47 (10):1010-1016. (in Chinese)
	丁海东, 刘慧, 陈一, 王丹, 梁建生. 2011. H₂O₂和MAPK介导油菜素内酯诱导番茄抗氧化防护酶SOD和CAT的信号途径. 植物生理学报, 47 (10):1010-1016.
[8]	Dong X Y, Hong Z L, Chatterjee J, Kim S H, Verma D S. 2008. Expression of callose synthase genes and its connection with Npr 1 signaling pathway during pathogen infection. Planta, 229 (1):87-98. doi: 10.1007/s00425-008-0812-3 URL
[9]	Grove M D, Spencer G F, Rohwedder W K, Mandava N, Worley J F, Warthen J D, Steffens G L, Flippen-Anderson J L, Cook J L C. 1979. Brassinolide,a plant growth-promoting steroid isolated from Brassica napus pollen. Nature, 281 (5728):216-217. doi: 10.1038/281216a0
[10]	Guo Fang, Wang Ya-jun, Xie Zhong-kui, Gao Yong-ping, Guo Zhi-hong, Zhang Yu-bao, Tong Xun-zhang. 2010. The effects of inducers on growth,disease resistance and related enzymes activities of lily cut flower. Acta Horticulturae Sinica, 37 (5):763-768. (in Chinese)
	郭芳, 王亚军, 谢忠奎, 高永平, 郭志鸿, 张玉宝, 童勋章. 2010. 诱抗剂对切花百合生长、抗病性及相关酶活性的影响. 园艺学报, 37 (5):763-768.
[11]	Hammes T T S. 2004. Response of potato grown under non-inductive condition paclobutrazol: shoot growth,chlorophyll content,net photosynthesis,assimilate partitioning,tuber yield,quality,and dormancy. Plant Growth Regulation, 43 (3):227-236. doi: 10.1023/B:GROW.0000045992.98746.8d URL
[12]	Hameed A, Goher M, Iqbal N. 2013. Drought induced programmed cell death and associated changes in antioxidants proteases and lipid peroxidation in wheat leaves. Biologia Plantarum, 57 (2):370-374. doi: 10.1007/s10535-012-0286-9 URL
[13]	He Yong-ming, Ding Xiu-ying, Cong Lin-ye, Guo Guang, Chen Shu-jun, Zhang Jun. 2007. Induction of PR protein and inhibition of TMV in tobacco by β-amino-butyric acid. Acta Phytopathologica Sinica, 37 (3):271-277.
	何永明, 丁秀英, 丛林晔, 郭光, 陈淑君, 张军. 2007. β-氨基丁酸诱导烟草产生PR蛋白及对TMV的抑制作用. 植物病理学报, 37 (3):271-277.
[14]	Huang Xue-mei, Zhang Chan, Pang Xue-qun, Zhang Zhao-qi. 2011. Early changes of reactive oxygen species in 2,6-dichloroisonicotianic acid inducing tolerance in postharvest banana fruits. Acta Horticulturae Sinica, 38 (2):265-272. (in Chinese)
	黄雪梅, 张灿, 庞学群, 张昭其. 2011. INA 诱导的香蕉果实抗病性与早期活性氧积累的关系. 园艺学报, 38 (2):265-272.
[15]	José G E, Briceño E X, Keith L M, Latorre B A. 2008. Canker and twig dieback of blueberry caused by Pestalotiopsis spp. and a Truncatella sp. in Chile. Plant Disease, 92 (10):1407-1414. doi: 10.1094/PDIS-92-10-1407 URL
[16]	Khripach V A, Zhabinskii V N, Groot A E. 1999. Brassinosteroids-a new class of plant hormones. Academic Press:7-23.
[17]	Li Sheng-hui. 2012. Mechanisms of OsNDPK1 improving disease-resistance and promoting cell division of plant[Ph. D. Dissertation]. Baoding: Hebei Agricultural University. (in Chinese)
	李生辉. 2012. OsNDPK1提高植物抗病性及促进细胞分裂的机理研究[博士论文]. 保定: 河北农业大学.
[18]	Lin Rui, Ren Hai-ying,An Xiao-xiao,Zheng Xi-liang,Liang Sen-miao,Zhang Shu-wen,Qi Xing-jiang. 2019. Effects of bio-organic fertilizer on twig blight disease control and recovery of tree vigor in bayberry. Acta Agriculturae Zhejiangensis, 31 (7):1096-1104. (in Chinese) doi: 10.3969/j.issn.1004-1524.2019.07.09
	林瑞, 任海英, 安笑笑, 郑锡良, 梁森苗, 张淑文, 戚行江. 2019. 生物有机肥对杨梅凋萎病防控及其树势恢复的影响. 浙江农业学报, 31 (7):1096-1104. doi: 10.3969/j.issn.1004-1524.2019.07.09
[19]	Liu Gao-feng. 2005. Studies on the selection of resistance elicitor against Marssonina coronaria(Ell.et Daris)Daris and the mechanisms of resistance induced[M. D. Dissertation]. Tai’an: Shandong Agricultural University. (in Chinese)
	刘高峰. 2005. 苹果褐斑病抗性诱导剂筛选及诱抗机理初探[硕士论文]. 泰安: 山东农业大学.
[20]	Liu Jian-ping, Tang Tao, Zhao Ming-ping. 2015. Control efficacy on Southern rice black-streaked dwarf virus of oligosaccharins · plant activator protein and its effects on growth promotion and yield increase of rice. Agrochemicals, 54 (8):606-609. (in Chinese)
	刘见平, 唐涛, 赵明平. 2015. 寡糖 · 链蛋白对南方水稻黑条矮缩病的防治效果及其对水稻的促长增产作用. 农药, 54 (8):606-609.
[21]	Liu Qing, Luan Xue-tao, Xu Shi-yan, Meng Ying, Gao Jiang-man, Xi Zhu-mei. 2016. Effect of 2,4-epibrassinolide treatment on grapevine leaf against Plasmopara viticola. Scientia Agricultura Sinica, 49 (15):3010-3018. doi: 10.3864/j.issn.0578-1752.2016.15.015
	刘庆, 栾雪涛, 徐世彦, 孟莹, 高江曼, 惠竹梅. 2016. 2,4-表油菜素内酯对葡萄叶片抵御霜霉菌侵染的影响. 中国农业科学, 49 (15):3010-3018. doi: 10.3864/j.issn.0578-1752.2016.15.015
[22]	Liu W X, Zhang F C, Zhang W Z, Song L F, Wu W H, Chen Y F. 2013. Arabidopsis Di 19 functions as a transcription factor and modulates PR1,PR2,and PR5 expression in response to drought stress. Molecular Plant, 6 (5):1487-1502. doi: 10.1093/mp/sst031 URL
[23]	Livak K J, Schmittgen T D. 2001. Analysis of relative gene expression data using real-time quantitative PCR and the 2^-ΔΔCT method. Methods, 25 (4):402-408. doi: 10.1006/meth.2001.1262 pmid: 11846609
[24]	Luan Xue-tao. 2017. Study of 2,4-epibrassinolide induced the resistance of grape detached leaves against downy mildew[Ph. D. Dissertation]. Yangling: Northwest A & F University. (in Chinese)
	栾雪涛. 2017. 2,4-表油菜素内酯处理提高葡萄离体叶片对霜霉病抗性的研究[博士论文]. 杨凌: 西北农林科技大学.
[25]	Miao Li-xiang, Yang Xiao-fang, Zhang Yu-chao, Jiang Gui-hua. 2021. Application situation of plant immune inducers in strawberry. Vegetables,(1):24-28. (in Chinese)
	苗立祥, 杨肖芳, 张豫超, 蒋桂华. 2021. 植物免疫诱导剂在草莓中的应用情况. 蔬菜,(1):24-28.
[26]	Niu Ji-shan, Liu Jing, Ni Yong-jing, Yin Jun. 2011. Induction of PR-1,PR-2,PR-5,Ta-JA2 and wheat powdery mildew resistance in response to MeJA treatment. Acta Phytopathologica Sinica, 41 (3):270-277. (in Chinese)
	牛吉山, 刘靖, 倪永静, 尹钧. 2011. 茉莉酸对PR-1、PR-2、PR-5和Ta-JA2基因表达以及小麦白粉病抗性的诱导. 植物病理学报, 41 (3):270-277.
[27]	Qiu De-wen. 2016. Research status and trend analysis of plant immune induction technology in China. Plant Proction, 42 (5):10-14. (in Chinese)
	邱德文. 2016. 我国植物免疫诱导技术的研究现状与趋势分析. 植物保护, 42 (5):10-14.
[28]	Ren H Y, Li G, Qi X J, Li F, Wang H R, Wei J G, Zhong S B. 2013. Identification and characterization of Pestalotiopsis spp. causing twig blight disease of bayberry(Myrica rubra Sieb. & Zucc)in China. European Journal of Plant Pathology, 137 (3):451-461. doi: 10.1007/s10658-013-0255-y URL
[29]	Ren Hai-ying, Qi Xing-jiang, Chen An-liang, Zheng Xi-liang, Liang Sen-miao, Yan Li-ju, Wang Ji-can, Li Gang. 2013. Effects of fungicides on twig blight disease of bayberry. Journal of Fruit Science, 30 (5):848-853. (in Chinese)
	任海英, 戚行江, 陈安良, 郑锡良, 梁森苗, 颜丽菊, 王继灿, 李岗. 2013. 十种杀菌剂对杨梅凋萎病的药效评价. 果树学报, 30 (5):848-853.
[30]	Shah K, Kumar R G, Verma S, Dubey R S. 2001. Effect of cadmium on lipid peroxidation,superoxide anion generation and activities of antioxidant enzymes in growing rice seedlings. Plant Science, 161 (6):1135-1144. doi: 10.1016/S0168-9452(01)00517-9 URL
[31]	Shang Qing-mao, Zhang Zhi-gang, Dong Tao, Song Shi-qing, Li Xiao-fen. 2007. Resistance of cucumber seedlings to Botrytis cinerea induced by exogenous brassinolide treatment. Chinese Journal of Applied & Environmental Biology, 13 (5):630-634. (in Chinese)
	尚庆茂, 张志刚, 董涛, 宋士清, 李晓芬. 2007. 油菜素内酯诱导黄瓜幼苗抗灰霉病研究. 应用与环境生物学报, 13 (5):630-634.
[32]	van Loon L C, Rep M, Pieterse C M. 2006. Significance of inducible defense-related proteins in infected plants. Annual Review of Phytopathology, 44:135-162. pmid: 16602946
[33]	Volynets A P, Pschenichnaya L A, Manzhelesova N E, Morozik G V, Khripach V A. 1997. The nature of protective action of 2,4-epibrassinolide on barley plants. Plant Growth Regulation, 24:133-137.
[34]	Wang Fang. 2014. Induced resistance and pathogenic mechanism of the shrink disease on jujube[Ph. D. Dissertation]. Yangling: Northwest A & F University. (in Chinese)
	王方. 2014. 枣缩果病抗病性诱导及致病机理初探[博士论文]. 杨凌: 西北农林科技大学.
[35]	Wang Fang, Wei Ting-ting, Mao Zhi-qiang, Jiao Jiao, Cui Shun-yan, Zhou Fan. 2020. Lactulose:control effect on cucumber Fusarium wilt and its induced resistance in cucumber. Chinese Agricultural Science Bulletin, 36 (13):154-159. (in Chinese) doi: 10.11924/j.issn.1000-6850.casb20190400052
	王芳, 魏婷婷, 毛志强, 焦姣, 崔顺艳, 周繁. 2020. 乳果糖对黄瓜枯萎病防效及其诱导抗性的作用. 中国农学通报, 36 (13):154-159. doi: 10.11924/j.issn.1000-6850.casb20190400052
[36]	Wang Kai-tuo, Liao Yun-xia, Di Hua-tao, Han Lin, Zheng Yong-hua. 2015. An investigation on the mechanism involved in defense response induced by methyl jasmonate in grape cell suspensions. Acta Horticulturae Sinica, 42 (7):1225-1232. (in Chinese) doi: 10.16420/j.issn.0513-353x.2015-0001 URL
	汪开拓, 廖云霞, 狄华涛, 韩林, 郑永华. 2015. 茉莉酸甲酯诱导葡萄悬浮细胞防卫反应机制的研究. 园艺学报, 42 (7):1225-1232. doi: 10.16420/j.issn.0513-353x.2015-0001 URL
[37]	Wang Xue-kui, Huang Jian-liang. 2015. Experimental principle and technology of plant physiology and biochemistry. Beijing: Higher Education Press. (in Chinese)
	王学奎, 黄见良. 2015. 植物生理生化实验原理与技术. 北京: 高等教育出版社.
[38]	Wees S V, Ent S V D, Pieterse C. 2008. Plant immune responses triggered by beneficial microbes. Current Opinion in Plant Biology, 11 (4):443-448. doi: 10.1016/j.pbi.2008.05.005 pmid: 18585955
[39]	Wu S W, Wang H W, Yang Z D, Kong L R. 2014. Expression Comparisons of pathogenesis-related(PR)genes in wheat in response to infection/infestation by Fusarium,Yellow dwarf virus(YDV)aphid-transmitted and hessian fly. Journal of Integrative Agriculture, 13 (5):926-936.
[40]	Xiang Miao-lian, Zhao Xian-yang, Chen Ming, Fu Yong-qi, Zeng Xiao-chun. 2017. Involvement of induced resistance by methyl jasmonate to bacterial wilt and reactive oxygen species metabolism in pepper seedlings. Acta Horticulturae Sinica, 44 (10):1985-1992. (in Chinese) doi: 10.16420/j.issn.0513-353x.2017-0004 URL
	向妙莲, 赵显阳, 陈明, 付永琦, 曾晓春. 2017. 茉莉酸甲酯诱导辣椒抗青枯病与活性氧代谢的关系. 园艺学报, 44 (10):1985-1992. doi: 10.16420/j.issn.0513-353x.2017-0004 URL
[41]	Xiao Rui-xue, Guo Li-li, Jia Qi-shi, Guo Da-long, Hou Xiao-gai. 2019. Research advances of brassinolide in the regulation of plant growth,development,yield and quality. Jiangsu Agricultural Sciences, 47 (10):16-21. (in Chinese)
	肖瑞雪, 郭丽丽, 贾琦石, 郭大龙, 侯小改. 2019. 油菜素内酯调控植物生长发育及产量品质研究进展. 江苏农业科学, 47 (10):16-21.
[42]	Xin Ya, Yu Hong,Xiao Wen-fei,Chai Wei-guo,Ruan Song-lin. 2020. Application of immune inducers in green prevention and control technology of strawberry diseases and insect pests. Journal of Zhejiang Agricultural Sciences, 61 (3):437-439. (in Chinese)
	忻雅, 余红, 肖文斐, 柴伟国, 阮松林. 2020. 免疫诱导剂在草莓病虫害绿色防控技术中的应用. 浙江农业科学, 61 (3):437-439.
[43]	Xu Run-dong, Sheng Shi-ying, Yang Xiu-fen, Liu Yong. 2016. Effect of induced resistance of oligosaccharins · plant activator protein on wheat to WYMV. Scientia Agricultura Sinica, 49 (18):3561-3568. (in Chinese) doi: 10.3864/j.issn.0578-1752.2016.18.010
	徐润东, 盛世英, 杨秀芬, 刘勇. 2016. 寡糖 · 链蛋白对小麦抗黄花叶病毒的免疫诱抗作用. 中国农业科学, 49 (18):3561-3568. doi: 10.3864/j.issn.0578-1752.2016.18.010
[44]	Yang Yi-lin, Zhang Zheng-min, Li Mei-lin, Zhao Li-yan, Jin Peng, Zheng Yong-hua. 2019. Modes of action of 2,4-epibrassionolide against postharvest gray mold decay of grapes. Food Science, 40 (15):231-238. (in Chinese) doi: 10.7506/spkx1002-6630-20180821-222
	杨艺琳, 张正敏, 李美琳, 赵立艳, 金鹏, 郑永华. 2019. 2,4-表油菜素内酯对葡萄果实采后灰霉病的抑制作用机理. 食品科学, 40 (15):231-238. doi: 10.7506/spkx1002-6630-20180821-222
[45]	Ye Xing-yuan, Tong Sheng-li, Liu Jian-jun. 2011. Effects of brassinolide on physiological character and yield and quality of strawberry in solar greenhouse. Guangdong Agricultural Sciences, 38 (8):39-41. (in Chinese)
	耶兴元, 仝胜利, 刘建军. 2011. 油菜素内酯对大棚温室草莓生理特性、产量及品质的影响. 广东农业科学, 38 (8):39-41.
[46]	Yuan Lian-lian, Li Wei, Xiao Zhi-xin, Zhang Jing, Li Yong-liang, Wang Feng-long, Yang Jin-guang. 2017. Effects of several kinds of plant inducer agents on disease resitance,agronomic traits,yield and quality in tobacco. Plant Protection, 43 (5):215-220. (in Chinese)
	袁莲莲, 李伟, 肖志新, 张静, 李永亮, 王凤龙, 杨金广. 2017. 几种植物诱导剂对烤烟抗病性、农艺性状及产量品质的影响. 植物保护, 43 (5):215-220.
[47]	Yuan Xin-lin, Li Mei-hua, Yu Si-si, Cui Yan-hua, Zhang Yin-bao, Wang Lan. 2016. Cotton resistance induced by 5% amino-oligosaccharin to Fusarium wilt and Verticillium wilt. China Cotton, 43 (3):15-21. (in Chinese)
	袁新琳, 李美华, 于丝丝, 崔燕华, 张银宝, 王兰. 2016. 5%氨基寡糖素诱导棉花抗枯黄萎病研究. 中国棉花, 43 (3):15-21.
[48]	Zeng Xiao-feng, Zeng Zhi-hong, Zeng Shun-de, Diao Yuan, Yin Xu-min, Shang Sang, Duan Wen-tao, Gao Lun-jiang. 2019. Effects of different resistance inducers on the antioxidant enzyme system in Flammulina velutipes. Journal of Southwest University(Natural Science Ddition), 41 (10):22-27. (in Chinese)
	曾小峰, 曾志红, 曾顺德, 刁源, 尹旭敏, 商桑, 段文韬, 高伦江. 2019. 不同抗性诱导剂对金针菇抗氧化酶系统的影响. 西南大学学报(自然科学版), 41 (10):22-27.
[49]	Zhang S W, Yu Z P, Qi X J, Wang Z, Zheng Y Y, Ren H Y, Liang S M, Zheng X L. 2021. Construction of a high-density genetic map and identification of leaf trait-related QTLs in Chinese bayberry(Myrica rubra). Frontiers in Plant Science,12:DOI:10.3389/fpls.2021.675855.
[50]	Zhang Xu, Li Jie, Feng Li-dan, He Jing, Li Yong-hui, Sun Tao, Zhang Sheng-dong. 2022. Effects of three disease resistance inducers on leaf growth,disease resistance and related enzyme activities of Lycium barbarum. Journal of Yunnan Agricultural University(Natural Science), 37 (3):413-421. (in Chinese)
	张煦, 李捷, 冯丽丹, 何静, 李永晖, 孙涛, 张生懂. 2022. 3种抗病诱导剂对宁夏枸杞生长、抗病性及相关酶活性的影响. 云南农业大学学报(自然科学), 37 (3):413-421.
[51]	Zhao Xiao-ming, Du Yu-guang, Bai Xue-fang. 2004. The field experiment of oligochitosan controlling plant virus disease. Chinese Agricultural Science Bulletin, 20 (4):245-247. (in Chinese)
	赵小明, 杜昱光, 白雪芳. 2004. 氨基寡糖素诱导作物抗病毒病药效试验. 中国农学通报, 20 (4):245-247.
[52]	Zheng Xiaodong, Xi Xiangli, Li Yuqi, Sun Zhijuan, Ma Changqing, Han Mingsan, Li Shaoxuan, Tian Yike, Wang Caihong. 2022. Effects and regulating mechanism of exogenous brassinosteroids on the growth of Malus hupehensis under saline-alkali stress. Acta Horticulturae Sinica, 49 (7):1401-1414. (in Chinese) doi: 10.16420/j.issn.0513-353x.2021-0499 URL
	郑晓东, 袭祥利, 李玉琪, 孙志娟, 马长青, 韩明三, 李少旋, 田义轲, 王彩虹. 2022. 油菜素内酯对盐碱胁迫下平邑甜茶幼苗生长的影响及调控机理研究. 园艺学报, 49 (7):1401-1414. doi: 10.16420/j.issn.0513-353x.2021-0499 URL
[53]	Zou Hui, Zhao Liang, Yuan Ting, Guo Qigao, Wu Di, Liang Guolu. 2021. Cloning and expression of a pathogenesis related protein gene EjPR1 from loquat induced by Colletotrichum gloeosporioides. Acta Horticulturae Sinica, 48 (2):367-376. (in Chinese) doi: 10.16420/j.issn.0513-353x.2020-0141 URL
	邹晖, 赵亮, 袁婷, 郭启高, 吴頔, 梁国鲁. 2021. 炭疽菌诱导的枇杷病程相关基因EjPR1的克隆及其表达分析. 园艺学报, 48 (2):367-376. doi: 10.16420/j.issn.0513-353x.2020-0141 URL