摘要
本论文在研究板栗疫病的病原菌产毒条件的基础上,通过有机溶剂浸提获得了病菌的粗提毒素,用生物测定的方法明确了所提取的粗毒素的致病作用及致病范围,并用粗毒素进行了抗性鉴定:进一步采用柱层析、HPLC分析等从粗提毒素中获得了毒素纯品,用质谱、核磁共振、红外光谱对毒素的分子结构进行了鉴定:在板栗植株和细胞的水平,通过对其生理生化的测定及组织超微结构的观察进行致病机理的研究:并以Cp-Ⅱ毒素为诱导因子诱导具有抗性的板栗愈伤组织,获得如下主要结果:
1.寄生隐丛赤壳菌在供试的液体培养基中可以产生对板栗有毒性作用的毒素,为了得到寄生隐丛赤壳菌的最佳产毒条件,本实验对该病菌的产毒培养条件及致病范围进行了研究。结果表明:不同培养液的产毒能力有明显的差异,7种培养液中,PD+板栗煎汁培养液产毒能力最强；栗疫病病菌产毒能力以培养温度为26℃,培养基pH值为6,振荡培养(120 r/min)条件下培养18 d生物活性最强;最佳氮源为蛋白胨,较好的碳源为糊精。试验选择中国板栗当年生幼嫩枝条进行粗毒素生物检测,表明培养得到的毒素原液具有一定的毒性。通过对致病范围的测定结果显示,在供试的25科33种植物中除对板栗有致病作用外,对其它不同科属的多种植物也具有致病活性,表明该毒素是一种非专化型毒素。
2.寄生隐丛赤壳菌(Cryphonectria parasitica)经液体培养,石油醚萃取其发酵液获得对板栗带叶嫩枝具有致萎活性的粗提物,以氯仿:石油醚:甲醇(6:2:2)、正丁醇:水:甲醇(8:1:1.5)作洗脱剂,粗提物经硅胶色谱分离,共得到3种纯毒素,分别为Cp-Ⅰ、Cp-Ⅱ、Cp-Ⅲ,3组份都对板栗幼苗致萎活性较高。质谱、核磁共振和红外光谱测定表明:Cp-Ⅰ分子量为278,化学式为C_(16)H_(22)O_4,化学名称为:dibutyl phthalate;Cp-Ⅱ分子量为157,化学式为C_9H_(19)ON,化学名称为1-(4-ethylpiperidin-1-yl)pmpan-2-one;Cp-Ⅲ分子量为128,化学式为C_7H_(14)ON,3-propylmorpholine。其化学结构式分别为:
3.应用酶活力测定的方法,研究板栗Castanea mollissima Blume不同抗性品种叶组织经栗疫菌Cryphonectria parasitica毒素Cp-处理后对其过氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)、抗坏血酸过氧化物酶(APX)、多酚氧化酶(PPO)和苯丙氨酸解氨酶(PAL)活性的影响。结果表明:适宜浓度(25-50μg/mL)的Cp-Ⅱ毒素处理板栗叶片,能提高保护性酶SOD、POD、CAT、APX、PPO和PAL的活性,抗病品种(北裕2号)活性变化幅度最大；随着毒素处理浓度升高,活性氧清除系统开始受到破坏,各品种的酶活性相继下降:Cp-Ⅱ毒素处理浓度达200μg/mL时,破坏了板栗叶片中的抗氧化酶系,造成活性氧过量积累,APX、PPO和PAL的活性随之降低,最终导致植物受到伤害最终导致植物受到破害;可溶性蛋白和可溶性糖含量先升后降,电导率和MDA含量升高,保护性酶SOD和CAT的活性升高及POD活性下降。Cp-Ⅱ毒素引起寄主的这种生理生化规律性变化可作为抗性鉴定的一种评价指标。
4.用寄生隐丛赤壳菌产生的具致萎活性的Cp-Ⅱ毒素处理板栗带叶嫩枝,研究毒素对叶组织、茎组织超微结构的影响。结果表明:清水处理后的板栗叶片,其叶肉细胞超微结构比较清晰,细胞质致密,可见排列整齐的细胞结构。随着毒素浓度的增加,叶细胞的细胞壁变形,质壁分离,质膜断裂,叶绿体膜、线粒体膜、核膜膨胀、断裂,内部间质电子透明化,叶绿体片层排列紊乱,线粒体脊膨胀甚至消失,损害发生早和严重的是叶绿体片层和质膜,抗病品种的膜系统比感病品种的受害轻。茎组织经毒素处理后,感品种茎组织细胞均出现细胞壁变形甚至断裂,质膜断裂,叶绿体变形严重,类囊体片层解体,叶绿体膜膨胀、断裂;抗病品种变化较小。
5.本研究以栗疫病抗病品种北裕2号和感病品种红光愈伤组织为材料,用Cp-Ⅱ毒素处理板栗愈伤组织,对处理后的愈伤组织进行细胞超微结构观察,测定其体内β-1,3-葡聚糖酶与几丁质酶活性,筛选具有抗性的愈伤组织。结果显示:Cp-Ⅱ毒素处理后的红光细胞膜对毒素的作用最敏感,北裕2号的愈伤组织变化较红光出现的晚；对毒素处理的愈伤组织的几丁质酶与β-1,3-葡聚糖酶活性测定结果表明,抗病品种比感病品种酶的活性增加幅度大、时间早,且两种酶的活性在毒素处理后均不同程度地高于对照;Cp-Ⅱ毒素浓度为50μg/mL胁迫愈伤组织28 d以后,经抗性分析感病病品种更易诱导出具有抗性的愈伤组织。
On the base of production conditions of the toxin from Cryphonectria parasitica, the pathogenic activity and its host range of the toxin was determined by bioassay methods. The resistance of different chestnut varieties was identified by using the toxin. The toxin was purified by silica gel column chromatography and HPLC, and was identified by the analysis of mass spectrography (MS), nuclear magnetic resonance (NMR) and infrared spectrum (IR). The pathogenesis of C. parasitica toxin was studied by physiological and biochemical determination, ultrastructural observation at plant and cell level. And Cp-Ⅱtoxin was taken as inducible factor to induce resistant callus. The results were as follows.
1. C. parasitica could produce phytotoxin in fluid media. The conditions of toxin production of C. parasitica were optimized. Results showed the liquid medium had the strongest toxin-producing capacity among 7 media. The optimal cultural conditions for producing toxin were cultured period at temperature 26℃, 18 d, and pH 6. Peptone and dextrin were the best nitrogenous source and carbonic sources respectively. The twigs of Chinese chestnut were selected to test the toxicity of crude toxin, which showed the cultured crude toxin had definite toxicity. C. parasitica toxin was a non-host-specific toxin. It had toxic activity to many plants from different families (33 species of 22 families). It could cause leaf blight and withered.
2. The strain of C. parasitica was cultivated by liquid fermentation. The crude material with wilt activity to chestnut browse was obtained by extracting antagonistic components with petroleum ether from the fermentation broth. Three compounds were obtained by silica gel column chromatography by using V(chloroform ):V(petroleum ether ):V(alcohol)=6:2:2 as eluent. The results showed the first compound (Cp-Ⅰ) had higher wilt activity than others. The chemical formula of the pathogenic fraction was identified by MS, NMR and IR. Cp-Ⅰwas C_(16)H_(22)O_4 (M=278), Cp-Ⅱwas C_9H_(19)ON (M=157), Cp-Ⅲwas C_7H_(14)ON (M=128). The structure of Cp-Ⅰ,Cp-Ⅱ,Cp-Ⅲwere as follows:
3. The method of excised leaf test was used to study the effect of Cp-Ⅱtoxin from C. parasitica on Castanea mollissima Blume. including four varieties with different resistance, specially on its activities of the enzymes superoxide dismutase (SOD), peroxidase (POD), ascorbate peroxidase (APX), catalase (CAT), polyphenol oxidase (PPO), phenylalanine pmmonia lyase (PAL). The results suggested: the activities of the protective enzymes, including SOD, POD, CAT, APX, PPO and PAL, increased notably and the change extent of the enzymes activities of the resistant variety was the largest after treated with the Cp-Ⅱtoxin concentration from 25μg/mL to 50μg/mL. With increasing concentration of the Cp-Ⅱtoxin, the activate oxygen metabolism system was destroyed, the balance of the protective enzyme system was broken. The enzymes activities were observed to decrease in all varieties. Finally, the antioxidant enzyme system was damaged, and the excessive accumulation of the active oxygen free radicals was enhanced when the treatment concentration of Cp-Ⅱtoxin was 200 ug/mL. The APX, PPO and PAL activities decreased with its accumulation, and eventually caused plant damage. The regularity change of physiological and biochemical of host caused by Cp-Ⅱtoxin could be used a marker to assess the resistant.
4. To study the ultra-structure of leaf and stem tissue of resistant and susceptible Chinese chestnut, the toxin of C. parasitica with wilt activity to Chinese chestnut browse was obtained by isolation and purification from its fermentation broth. The Results indicated that the damage on leaf tissue included the transformation of cell wall, the plasmolysis, the swollen and disruption of chloroplast, mitochondria and nuclei the transparency of in ternalmatrix, the disordering of chloroplast lamellae, and the expanding and the disappearing of mitochondria ridges. The lamellae and membrane of chloroplasts are more sensitive to the toxin and change earlier. The damages on membrane systems of the cells are lighter and occur later in the R-type than the S-type one. The stem was treated with Cp-Ⅱtoxin, the results showed that plasmolysis and chloroplast deformation only took place, the ultra-structure of Hongguang stem tissue was badly damaged, including the deformed and broken cell wall, the broken plasmalemma, the misshaped chloroplast, decomposed chloroplast lamellae and broken envolopen of chloroplast. Beiyu 2.
5. The callus of two chestnut cultivars, Beiyu 2 and Hongguang, which were susceptive and resistant to C. parasitica respectively, were compared at ultra-structural level after callus were treated with Cp-Ⅱtoxin. The callus were used to determine the activities of chitinase andβ-1,3-glucanase. The results indicated that plasma membrane was more sensitive to Cp-Ⅱtoxin than other organs. The changes of Hongguang exhibited earlier than Beiyu 2. The activities of chitinase andβ-1,3-glucanase in the callus cells treated with the Cp-Ⅱtoxin were increased to the higher level at earlier time point in resistant cultivars than these in the susceptible cultivars. Chitinase andβ-1,3-glucanase activities of the resistant varieties and the susceptible ones were higher than the control, suggested that treated with Cp-Ⅱtoxin could induce resistance to C. parasitica through activating chitinase andβ-1,3-glucanase. The resistant callus was obtained by taking Cp-Ⅱtoxin (50μg/mL) as inducible factor, and susceptible variety was induced easier by analysis of resistance.
引文
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