寡雄腐霉RCU1菌株及寡雄蛋白诱导番茄抗病性的研究
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摘要
寡雄腐霉Pythium oligandrum Drechsler是一种土壤习居型腐生菌,它能在多种农作物根围定殖,能拮抗或寄生20多种植物病原菌,拮抗机制主要包括真菌寄生、抗生作用和诱导抗性等。寡雄腐霉对植物无毒性,对环境安全。寡雄蛋白(oligandrin)是寡雄腐霉产生的一种拟激发素蛋白,对番茄植株本身无毒性,不会引起番茄产生过敏性反应坏死斑,能诱导番茄对多种蔬菜重要病害产生系统获得抗性。本文在温室生产环境下,在番茄根部施用寡雄蛋白,系统地研究了寡雄蛋白诱导番茄叶片对灰霉病的抗性及其机理,进一步应用荧光定量PCR的方法探明经寡雄蛋白诱导番茄植株产生系统获得抗性与病程相关蛋白的关系,以及诱导抗性的信号转导途径；并制备了寡雄蛋白的多克隆抗体,应用免疫胶体金标记技术对寡雄蛋白在植株中的传递途径、作用位点等进行了亚细胞定位,主要研究结果如下：
1、寡雄腐霉菌株RCUl及其分泌物的抑菌作用及对番茄灰霉病的防效
应用玻璃纸法和生长速率法测定了寡雄腐霉菌株RCUl分泌物对9种植物病原真菌菌丝生长的抑制作用。结果表明,该分泌物对供试的9种植物病原真菌菌丝生长均有抑制作用；分子量小于8KD的寡雄腐霉分泌物能强烈抑制森禾腐霉、灰葡萄孢、黄色镰刀菌的菌丝生长,抑菌率分别达到88.9%、86.7%和84.6%,对供试的四种腐霉以及尖孢镰刀菌的4个专化型的抑菌率均达显著性差异。在扫描电镜下可见,经寡雄腐霉分泌物处理的灰葡萄孢,菌丝细胞破裂、穿孔、干瘪,且菌丝分化出的分生孢子梗少,产孢量明显下降。温室盆栽试验表明,寡雄腐霉滤液对番茄灰霉病具有显著的预防保护作用和治疗作用,其预防保护作用与杀菌剂50%多菌灵可湿性粉剂1000倍液的防治效果达到同一显著水平。
2、寡雄蛋白诱导番茄苗对灰霉病的抗性及机理研究
温室盆栽实验表明,寡雄蛋白(10μg/mL)处理番茄苗根系能够诱导番茄叶片对灰霉病的抗性,诱抗效果对供试品种合作903和凯特2号无显著差异。寡雄蛋白番茄根部诱导24h后接种灰葡萄孢,保护作用达61.3%,治疗效果可达51.8%；应用荧光定量PCR的方法研究了寡雄蛋白对番茄叶片基因表达量的影响,结果表明,寡雄蛋白根部处理24h,诱导番茄叶片病程相关蛋白几丁质酶和β-1,3葡聚糖酶基因表达量分别比对照增强4.7倍和3.1倍。
寡雄蛋白处理番茄根系24h后接种灰葡萄孢,处理后3d及5d木质素含量比接种灰葡萄孢对照分别升高18.5%和34.4%(合作903),22.3%和27.6%(凯特2号)；番茄叶片POD、PPO、PAL活性均明显高于接种灰葡萄孢对照,峰值较对照高20.0%,32.6%,57.1%(合作903),43.9%、32.6%、41.5%(凯特2号)。诱导番茄植株防御酶活性的提高,调控体内次生物质代谢,诱导病程相关蛋白基因表达量的上调,可能是寡雄蛋白诱导番茄叶片抵御灰霉病的主要机理。
3、寡雄蛋白诱导番茄抗灰霉病的信号转导途径
应用荧光定量PCR的方法研究了寡雄蛋白对番茄叶片的基因表达量的影响,结果表明,寡雄蛋白根部处理可诱导番茄叶片乙烯反应因子ERF2和茉莉酸(JA)相关因子PR6基因表达量分别比对照增强6.6倍和3.6倍,寡雄蛋白诱导对番茄灰霉病的系统抗性依赖于ISR中的JA和ET信号转导途径；寡雄蛋白根部处理24h,诱导番茄叶片病程相关蛋白几丁质酶和β-1,3葡聚糖酶基因表达量分别比对照增强4.7倍和3.1倍,表明寡雄蛋白诱导番茄对灰霉病的抗性同时具有ISR和SAR的部分特征。寡雄蛋白可能是通过水杨酸(SA)协同JA／乙烯信号转导途径诱导番茄对灰霉病的抗性。
4、寡雄蛋白诱导番茄果实对灰霉病的抗性及机理研究
寡雄蛋白(10μg/mL)处理番茄果蒂处24h后在番茄果肩处接种灰葡萄孢,常温下寡雄蛋白处理的番茄果实腐烂率比对照降低了49.5%。常温贮藏结束时,寡雄蛋白处理番茄的细胞质膜相对透性比对照低63%；MDA产生量比对照降低31%。寡雄蛋白处理番茄果实PAL、PPO和POD活性峰值比接种处理高18.8%,20.0%和22.7%。应用荧光定量PCR的方法研究了寡雄蛋白对番茄果实基因表达量的影响,结果表明,寡雄蛋白处理的番茄果实PR-2a和PR-3a基因表达量比对照升高2.7倍和3.6倍。诱导番茄防御酶活性的提高,维持细胞结构的完整性,诱导PR-2a和PR-3a基因表达量的升高,是寡雄蛋白诱导番茄果实抵御灰霉病的关键。
5、寡雄蛋白与寄主植物互作的细胞化学定位
制备了寡雄蛋白的多克隆抗体,应用免疫胶体金标记技术对寡雄蛋白在寡雄腐霉菌丝中、寡雄腐霉在番茄根部定殖以及寡雄蛋白在番茄叶片中的移动进行亚细胞定位,结果表明,PDA平板或液体培养基中培养的寡雄腐霉菌丝中,寡雄蛋白主要积累在菌丝细胞壁,可以分泌到寡雄腐霉细胞外,表明寡雄蛋白是一种外泌蛋白。
免疫胶体金标记结果表明,寡雄腐霉在番茄根部定殖4h即可在番茄根部细胞中检测到寡雄蛋白,随着时间的推移,寡雄蛋白的量逐渐增加,在根组织细胞的细胞壁、叶绿体等处聚集。菌丝中合成的寡雄蛋白主要束缚在菌丝壁上,而菌丝内部较少,说明寡雄蛋白合成后很快分泌出去,而不是在成熟的菌丝中积累。寡雄腐霉接种24h即可在番茄根组织细胞内检测到大量寡雄蛋白,表明番茄是寡雄腐霉的适宜寄主,寡雄腐霉可以大量在番茄根部定殖,分泌寡雄蛋白诱导番茄系统获得抗性。
寡雄蛋白处理番茄中部叶片,免疫胶体金标记结果表明,4h后番茄上、中、下部叶片细胞壁和细胞核中均有大量寡雄蛋白存在,表明寡雄蛋白可在植物体内进行上下转运。寡雄蛋白可能是从侵入部位传导到植物其他部位并激发系统获得抗性反应的信号,其信号受体可能定位在细胞膜上,在细胞核中的特异性积累可能对植物防卫反应具有调节效应,从而提高植物的抗病能力。
Pythium oligandrum Drechsler is one kind of saprophyte living in soil. It has been shown to colonize the rhizosphere of a wide range of agriculturally important crops. It is antagonistic to and parasitic on more than twenty important pathogenic fungi or other oomycetes. It is Non-pathogenic, harmless to plant and environment. Its antagonistic mechanisms include mycoparasitism, antibiosis and induce resistance. Oligandrin, the elicitin-like protein produced by P. oligandrum, induces systemic resistance in tomato plants, but cause no hypersensitive response (HR). In this study, we applied oligandin to roots to study the effects and mechanism of anti-stress by oligandrin to induce resistance of tomato seedling against B. cinerea in greenhouse. We also researched the mechanisms of the interactions between oligandrin and pathogens or host plant.
The main results were obtained as followings:
1. Inhibitory of the secretion of Pythium oligandrum strain RCU1 on plant pathogenic fungi and the control effect against tomato gray mould in greenhouse.
The inhibition effect of the molecular weight less than 8 kD secretion of P. oligandrum on total 16 isolates of plant pathogenic fungi belonging to 9 species was tested on Petri dishes with cellophane paper and growth rate methods. The secretion of P. oligandrum distinctly depressed the mycelial growth of P. sylvaticum, Botrytis cinerea, and Fusarium culmorum with inhibition rates of 88.9%,86.7% and 84.6%, respectively. The inhibition rates of four Pythium species and four formae speciales of F. oxysporum were significantly differet (P=0.05), respectively. The morphology of B. cinerea treated with the secretion of P. oligandrum was abnormal under scanning electron microscope with mycelia shriveled, small spores appeared in the cell wall and reduced spore output significantly. In the greenhouse trial, the filtrate of P. oligandrum had significantly protective and curative effect against tomato gray mould, and the protective effect of the filtrate was equivalent strong with that of the trearment by using carbendazim.
2. Defense responses in tomato leaves against Botrytis cinerea induced by application of oligandrin to roots
In the greenhouse trial, the root treatment with oligandrin (10μg/mL) had significantly protective and curative effect against tomato gray mould in tomato leaves with efficiency of 61.3% and 51.8%, respectively. Furthermore, the expression of the pathogenesis-related proteins (PRs) genes,β-1,3-glucanase and chitinase were upregulated in the leaves of treated tomato plant 4.7 fold and 3.1 fold, respectively.
The oligandrin treatment increased contents of lignin with 18.5% and 34.4% on cultivar Hezuo-903,22.3% and 27.6% on cultivar Kaite-2 than that in control at the peak stage, respectively, 3d and 5d after inoculation. The results from bio-assay indicated that the oligandrin treatment stimulated the activity of the defense related enzymes, peroxidase (POD), polyphenol oxidase (PPO) and phenylalanine ammonia-lyase (PAL). Their activity in the oligandrin-treated plant was about 20.0%,32.6%,57.1% (Hezuo-903),43.9%、32.6%、41.5%(Kaite-2) higher than that in control at the peak stage after inoculation, respectively. We conclude that antioxidant enzyme activities and pathogenesis-related proteins (PRs) genes upregulated were two crucial targets of oligandrin in tomato leaves during the establishment of defense responses.
3. Signal pathway induced by oligandrin against Botrytis cinerea in tomato plant
Oligandrin (10μg/mL) treatment of tomato root induced the expression of ethylene (ET) -related transcription factor (ERF2) genes and the jasmonic acid (JA)-responsive gene for the basic PR-6 proein up-regulated 6.6 fold and 3.6 fold in tomato plants, respectively. These results indicated that the JA/ET signaling pathway is required for oligandrin induced resistance against B. cinerea in tomato. Furthermore, the expression of the pathogenesis-related proteins (PRs) genes, β-1,3-glucanase and chitinase are upregulated in the leaves of treated tomato plant 4.7 fold and 3.1 fold, respectively. Defense responses in tomato leaves against B. cinerea induced by oligandrin have the features both ISR and SAR. Oligandrin perhaps induced SA and JA/ET Signal pathway together.
4. Defense responses in tomato fruit induced by oligandrin against Botrytis cinerea
Oligandrin (10μg/mL) treated tomoto fruit tampuk (20μl) significantly reduced incidence and severity of gray mould (Botrytis cinerea). After artificial inoculation the severity of body rots was decreased by 49.5%. Vc contants was increased 36.1%. The results from bio-assay indicated that the treatment stimulated the activity of the defense related enzymes, phenylalanine ammonia-lyase (PAL), polyphenol oxidase (PPO), peroxidase (POD) activity in the oligandrin-treated fruit was about 18.8%,20.0%,22.7% higher than that in control on the 3rd day after the inoculation, respectively. Furthermore, increased expression of genes encoding pathogenesis -related proteins (PRs), such as PR-2a (extracellularβ-1,3 -glucanase) and PR-3a (extracellular chitinase) at mRNA level in tomato fruit was about 2.7-fold and 3.6-fold above that in the control plant at the peak stage. These results suggest that the defense responses of tomato fruit induced by oligandrin against gray mould might be attributed to systemic acquired resistance (SAR). Antioxidant enzyme activities and pathogenesis-related proteins (PRs) genes upregulated are two crucial targets of oligandrin in tomato leaves during the establishment of defense responses.
5. Cytology of the interactions between oligandrin and host plant
The analysis of hyphae of P. oligandrum, tomato roots and leaves by electron microscopy immunogold labeling with polyclonal antibodies against oligandrin confirmed that oligandrin was distributed on the cell wall of the mycelia and secreted outside. During the infection and colonization process P. oligandrum secreted oligandrin, the antigen sites were found in the tomato plant cell wall and the cell nucleus which maybe contacted the induced resistance. Tomato is a good host for P. oligandrum. Spraying oligandrin to tomato leaves, the antigen sites were found in the plant cell wall, chloroplast, and the cell nucleus. Oligandrin can translocated throughout the plant. Even though the exact mechanisms by which oligandrin operates to trigger resistance in tomato are not fully elucidated, the present results demonstrate that oligandrin maybe a signal for triggering synthesis and accumulation of defense gene products, thus increased the disease resistance in tomato.
1、寡雄腐霉菌株RCUl及其分泌物的抑菌作用及对番茄灰霉病的防效
应用玻璃纸法和生长速率法测定了寡雄腐霉菌株RCUl分泌物对9种植物病原真菌菌丝生长的抑制作用。结果表明,该分泌物对供试的9种植物病原真菌菌丝生长均有抑制作用；分子量小于8KD的寡雄腐霉分泌物能强烈抑制森禾腐霉、灰葡萄孢、黄色镰刀菌的菌丝生长,抑菌率分别达到88.9%、86.7%和84.6%,对供试的四种腐霉以及尖孢镰刀菌的4个专化型的抑菌率均达显著性差异。在扫描电镜下可见,经寡雄腐霉分泌物处理的灰葡萄孢,菌丝细胞破裂、穿孔、干瘪,且菌丝分化出的分生孢子梗少,产孢量明显下降。温室盆栽试验表明,寡雄腐霉滤液对番茄灰霉病具有显著的预防保护作用和治疗作用,其预防保护作用与杀菌剂50%多菌灵可湿性粉剂1000倍液的防治效果达到同一显著水平。
2、寡雄蛋白诱导番茄苗对灰霉病的抗性及机理研究
温室盆栽实验表明,寡雄蛋白(10μg/mL)处理番茄苗根系能够诱导番茄叶片对灰霉病的抗性,诱抗效果对供试品种合作903和凯特2号无显著差异。寡雄蛋白番茄根部诱导24h后接种灰葡萄孢,保护作用达61.3%,治疗效果可达51.8%；应用荧光定量PCR的方法研究了寡雄蛋白对番茄叶片基因表达量的影响,结果表明,寡雄蛋白根部处理24h,诱导番茄叶片病程相关蛋白几丁质酶和β-1,3葡聚糖酶基因表达量分别比对照增强4.7倍和3.1倍。
寡雄蛋白处理番茄根系24h后接种灰葡萄孢,处理后3d及5d木质素含量比接种灰葡萄孢对照分别升高18.5%和34.4%(合作903),22.3%和27.6%(凯特2号)；番茄叶片POD、PPO、PAL活性均明显高于接种灰葡萄孢对照,峰值较对照高20.0%,32.6%,57.1%(合作903),43.9%、32.6%、41.5%(凯特2号)。诱导番茄植株防御酶活性的提高,调控体内次生物质代谢,诱导病程相关蛋白基因表达量的上调,可能是寡雄蛋白诱导番茄叶片抵御灰霉病的主要机理。
3、寡雄蛋白诱导番茄抗灰霉病的信号转导途径
应用荧光定量PCR的方法研究了寡雄蛋白对番茄叶片的基因表达量的影响,结果表明,寡雄蛋白根部处理可诱导番茄叶片乙烯反应因子ERF2和茉莉酸(JA)相关因子PR6基因表达量分别比对照增强6.6倍和3.6倍,寡雄蛋白诱导对番茄灰霉病的系统抗性依赖于ISR中的JA和ET信号转导途径；寡雄蛋白根部处理24h,诱导番茄叶片病程相关蛋白几丁质酶和β-1,3葡聚糖酶基因表达量分别比对照增强4.7倍和3.1倍,表明寡雄蛋白诱导番茄对灰霉病的抗性同时具有ISR和SAR的部分特征。寡雄蛋白可能是通过水杨酸(SA)协同JA／乙烯信号转导途径诱导番茄对灰霉病的抗性。
4、寡雄蛋白诱导番茄果实对灰霉病的抗性及机理研究
寡雄蛋白(10μg/mL)处理番茄果蒂处24h后在番茄果肩处接种灰葡萄孢,常温下寡雄蛋白处理的番茄果实腐烂率比对照降低了49.5%。常温贮藏结束时,寡雄蛋白处理番茄的细胞质膜相对透性比对照低63%；MDA产生量比对照降低31%。寡雄蛋白处理番茄果实PAL、PPO和POD活性峰值比接种处理高18.8%,20.0%和22.7%。应用荧光定量PCR的方法研究了寡雄蛋白对番茄果实基因表达量的影响,结果表明,寡雄蛋白处理的番茄果实PR-2a和PR-3a基因表达量比对照升高2.7倍和3.6倍。诱导番茄防御酶活性的提高,维持细胞结构的完整性,诱导PR-2a和PR-3a基因表达量的升高,是寡雄蛋白诱导番茄果实抵御灰霉病的关键。
5、寡雄蛋白与寄主植物互作的细胞化学定位
制备了寡雄蛋白的多克隆抗体,应用免疫胶体金标记技术对寡雄蛋白在寡雄腐霉菌丝中、寡雄腐霉在番茄根部定殖以及寡雄蛋白在番茄叶片中的移动进行亚细胞定位,结果表明,PDA平板或液体培养基中培养的寡雄腐霉菌丝中,寡雄蛋白主要积累在菌丝细胞壁,可以分泌到寡雄腐霉细胞外,表明寡雄蛋白是一种外泌蛋白。
免疫胶体金标记结果表明,寡雄腐霉在番茄根部定殖4h即可在番茄根部细胞中检测到寡雄蛋白,随着时间的推移,寡雄蛋白的量逐渐增加,在根组织细胞的细胞壁、叶绿体等处聚集。菌丝中合成的寡雄蛋白主要束缚在菌丝壁上,而菌丝内部较少,说明寡雄蛋白合成后很快分泌出去,而不是在成熟的菌丝中积累。寡雄腐霉接种24h即可在番茄根组织细胞内检测到大量寡雄蛋白,表明番茄是寡雄腐霉的适宜寄主,寡雄腐霉可以大量在番茄根部定殖,分泌寡雄蛋白诱导番茄系统获得抗性。
寡雄蛋白处理番茄中部叶片,免疫胶体金标记结果表明,4h后番茄上、中、下部叶片细胞壁和细胞核中均有大量寡雄蛋白存在,表明寡雄蛋白可在植物体内进行上下转运。寡雄蛋白可能是从侵入部位传导到植物其他部位并激发系统获得抗性反应的信号,其信号受体可能定位在细胞膜上,在细胞核中的特异性积累可能对植物防卫反应具有调节效应,从而提高植物的抗病能力。
Pythium oligandrum Drechsler is one kind of saprophyte living in soil. It has been shown to colonize the rhizosphere of a wide range of agriculturally important crops. It is antagonistic to and parasitic on more than twenty important pathogenic fungi or other oomycetes. It is Non-pathogenic, harmless to plant and environment. Its antagonistic mechanisms include mycoparasitism, antibiosis and induce resistance. Oligandrin, the elicitin-like protein produced by P. oligandrum, induces systemic resistance in tomato plants, but cause no hypersensitive response (HR). In this study, we applied oligandin to roots to study the effects and mechanism of anti-stress by oligandrin to induce resistance of tomato seedling against B. cinerea in greenhouse. We also researched the mechanisms of the interactions between oligandrin and pathogens or host plant.
The main results were obtained as followings:
1. Inhibitory of the secretion of Pythium oligandrum strain RCU1 on plant pathogenic fungi and the control effect against tomato gray mould in greenhouse.
The inhibition effect of the molecular weight less than 8 kD secretion of P. oligandrum on total 16 isolates of plant pathogenic fungi belonging to 9 species was tested on Petri dishes with cellophane paper and growth rate methods. The secretion of P. oligandrum distinctly depressed the mycelial growth of P. sylvaticum, Botrytis cinerea, and Fusarium culmorum with inhibition rates of 88.9%,86.7% and 84.6%, respectively. The inhibition rates of four Pythium species and four formae speciales of F. oxysporum were significantly differet (P=0.05), respectively. The morphology of B. cinerea treated with the secretion of P. oligandrum was abnormal under scanning electron microscope with mycelia shriveled, small spores appeared in the cell wall and reduced spore output significantly. In the greenhouse trial, the filtrate of P. oligandrum had significantly protective and curative effect against tomato gray mould, and the protective effect of the filtrate was equivalent strong with that of the trearment by using carbendazim.
2. Defense responses in tomato leaves against Botrytis cinerea induced by application of oligandrin to roots
In the greenhouse trial, the root treatment with oligandrin (10μg/mL) had significantly protective and curative effect against tomato gray mould in tomato leaves with efficiency of 61.3% and 51.8%, respectively. Furthermore, the expression of the pathogenesis-related proteins (PRs) genes,β-1,3-glucanase and chitinase were upregulated in the leaves of treated tomato plant 4.7 fold and 3.1 fold, respectively.
The oligandrin treatment increased contents of lignin with 18.5% and 34.4% on cultivar Hezuo-903,22.3% and 27.6% on cultivar Kaite-2 than that in control at the peak stage, respectively, 3d and 5d after inoculation. The results from bio-assay indicated that the oligandrin treatment stimulated the activity of the defense related enzymes, peroxidase (POD), polyphenol oxidase (PPO) and phenylalanine ammonia-lyase (PAL). Their activity in the oligandrin-treated plant was about 20.0%,32.6%,57.1% (Hezuo-903),43.9%、32.6%、41.5%(Kaite-2) higher than that in control at the peak stage after inoculation, respectively. We conclude that antioxidant enzyme activities and pathogenesis-related proteins (PRs) genes upregulated were two crucial targets of oligandrin in tomato leaves during the establishment of defense responses.
3. Signal pathway induced by oligandrin against Botrytis cinerea in tomato plant
Oligandrin (10μg/mL) treatment of tomato root induced the expression of ethylene (ET) -related transcription factor (ERF2) genes and the jasmonic acid (JA)-responsive gene for the basic PR-6 proein up-regulated 6.6 fold and 3.6 fold in tomato plants, respectively. These results indicated that the JA/ET signaling pathway is required for oligandrin induced resistance against B. cinerea in tomato. Furthermore, the expression of the pathogenesis-related proteins (PRs) genes, β-1,3-glucanase and chitinase are upregulated in the leaves of treated tomato plant 4.7 fold and 3.1 fold, respectively. Defense responses in tomato leaves against B. cinerea induced by oligandrin have the features both ISR and SAR. Oligandrin perhaps induced SA and JA/ET Signal pathway together.
4. Defense responses in tomato fruit induced by oligandrin against Botrytis cinerea
Oligandrin (10μg/mL) treated tomoto fruit tampuk (20μl) significantly reduced incidence and severity of gray mould (Botrytis cinerea). After artificial inoculation the severity of body rots was decreased by 49.5%. Vc contants was increased 36.1%. The results from bio-assay indicated that the treatment stimulated the activity of the defense related enzymes, phenylalanine ammonia-lyase (PAL), polyphenol oxidase (PPO), peroxidase (POD) activity in the oligandrin-treated fruit was about 18.8%,20.0%,22.7% higher than that in control on the 3rd day after the inoculation, respectively. Furthermore, increased expression of genes encoding pathogenesis -related proteins (PRs), such as PR-2a (extracellularβ-1,3 -glucanase) and PR-3a (extracellular chitinase) at mRNA level in tomato fruit was about 2.7-fold and 3.6-fold above that in the control plant at the peak stage. These results suggest that the defense responses of tomato fruit induced by oligandrin against gray mould might be attributed to systemic acquired resistance (SAR). Antioxidant enzyme activities and pathogenesis-related proteins (PRs) genes upregulated are two crucial targets of oligandrin in tomato leaves during the establishment of defense responses.
5. Cytology of the interactions between oligandrin and host plant
The analysis of hyphae of P. oligandrum, tomato roots and leaves by electron microscopy immunogold labeling with polyclonal antibodies against oligandrin confirmed that oligandrin was distributed on the cell wall of the mycelia and secreted outside. During the infection and colonization process P. oligandrum secreted oligandrin, the antigen sites were found in the tomato plant cell wall and the cell nucleus which maybe contacted the induced resistance. Tomato is a good host for P. oligandrum. Spraying oligandrin to tomato leaves, the antigen sites were found in the plant cell wall, chloroplast, and the cell nucleus. Oligandrin can translocated throughout the plant. Even though the exact mechanisms by which oligandrin operates to trigger resistance in tomato are not fully elucidated, the present results demonstrate that oligandrin maybe a signal for triggering synthesis and accumulation of defense gene products, thus increased the disease resistance in tomato.
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