自毒物质与病原真菌协同对连作地黄的致害作用研究
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摘要
怀地黄是我国著名“四大怀药”之一,但其药用植物(玄参科多年生草本植物)地黄(Rehmannia glutinosa Libosch.),却存在非常严重的连作障碍。本研究基于当前连作障碍的研究现状,构建了地黄无菌苗体系,分析评价了地黄须根自毒作用,从地黄连作土壤和发病植株中分离了地黄专化型病原真菌,并进一步探讨了须根自毒物质与病原菌在地黄连作障碍中的协同作用。实验结果如下：
道地产区与非道地产区地黄品质存在着差异,但正茬与重茬地黄的品质差异明显高于产区之间差异；两产区地黄的连作土壤均能抑制地黄幼苗生长；连作还造成根际土壤细菌数量减少,土壤真菌和放线菌数量增多,土壤微生物类型由“细菌型”向“真菌型”过渡；与正茬土壤相比,连作根际土壤中土壤脲酶、蔗糖酶和过氧化氢酶的活性显著下降,土壤多酚氧化酶、纤维素酶和蛋白酶的活性有所增加,但土壤磷酸酶活性在两个产区连作土壤中呈现相反的变化趋势。可见,地黄连作会引起土壤酶活性和土壤微生态区系的变化,影响土壤酶活性,降低了地黄对营养物质的吸收利用,进而造成地黄药用品质下降。
通过对以地黄块根、叶片、地黄顶芽及地黄新出根生芽等作为外殖体材料的无菌苗构建体系的比较性分析,认为新出根生芽可以作为地黄组织培养的良好外殖体材料。优化并建立了以MS+6-BA1mgL-1+NAA0. mgL-11为培养基的成苗诱导体系；确定了以MS+6-BA0.3mgL-1+NAA0.2mgL-1为培养基诱导的地黄丛生苗繁殖率高,苗体粗壮；经MS+PP3330.1mgL-1生根诱导的地黄无菌苗发根快而多,根部生长强壮,植株叶色浓绿,经炼苗后移栽可作为地黄连作障碍研究提供实验材料。
室内自毒作用潜力评价表明,相同浓度处理下的地黄须根提取物、地黄茬后土壤提取物、地黄根系分泌物均能抑制地黄生长,表现出自毒作用,且地黄须根提取物具有更强的化感抑制作用；95%乙醇进行地黄须根自毒物质浸提,经硅胶柱色谱分离,进行活性追踪实验确定部位Ⅲ为抑制活性最高的分离部位,硅烷化衍生化后进行气质联用分析得到32个化学物质；通过文献检阅后选择9个单体物质对地黄幼苗进行生物测试,结果表明所选的9个单体物质能较大程度上抑制地黄生长。进一步分析发现,地黄自毒物质不仅引起地黄体内膜脂过氧化物质的增高,导致细胞生物膜透性增强,保护酶活性持续升高,还影响地黄幼苗内源激素水平,造成植物伤害。
通过PDA平板培养法,从连作土壤中分离到病原真菌42株,从地黄病株中分离到病原真菌24株。这66株真菌型病原菌分属子囊菌门、结合菌门、担子菌门和半知菌门等4个门。采用回接发病实验鉴定到有4株菌地黄专化型病原菌,结合菌落形态、孢子形态和iTS区的PCR扩增后测序结果初步推断菌株CCS038为镰刀菌属(Fusaiunn)雪腐镰刀菌种(Fusarium nivale),菌株CCS043为镰刀菌属(Fusaiunn)尖孢镰刀菌种(Fusarium oxysporum),菌株CCS025为曲霉属(Aspergillus)黄曲霉菌(Aspergillus flavus),菌株CCS012为曲霉属(Aspergillus)烟曲霉菌(Aspergillus fumigatus)。
地黄须根自毒物质与病原菌存在协同作用,地黄须根自毒物质特别是阿魏酸和对羟基苯甲酸不仅能诱导地黄专化型病原菌的菌丝生长和孢子增殖和萌发,还通过增强真菌信号转导系统如SH3结构和F-box蛋白体系和分子伴侣等修饰原件加强机体内结构基因表达等过程的调控,还增强了对真菌营养代谢相关的调控,促进真菌对营养物质的利用,进而影响真菌形态建成,还能通过破坏宿主细胞壁和防御体系,增强病原菌对宿主的侵染能力,从而加剧对地黄的致害过程。
Rehmannia (Rehmannia glutinosa Libosch) is in the Scrophulariaceae family and is one of the most common and important medicinal herbal plants in China. However, the consecutively monocultured plants are prone to severe diseases resulting in reduced biomass, especially the tuberous products. In this study, the propagation system of Rehmannia was established, the autotoxicity of fibrous root was evaluated, and the host-specific pathogenic fungi were isolated from replanted soil and wilt plants of Rehmannia. Furthermore, the synergistic effect of autotoxins and phytopathogenic fungi on the consecutive monoculture of Rehmannia was explored. The main results were as follows:
In this study, the experiments were conducted to evaluate the quality of Rehmannia due to consecutive monoculture, and to explore the decline in the quality of Rehmannia from the sides of soil micro-ecology (soil microbes and enzyme activities). The results showed that, the quality of Rehmannia from the origin region (Jiaozuo, Henan province) and production region (Linfen, Shanxi Province) were different, but the differences between the newly planted soil and monoculture soil were significantly higher than the differences in regions. The monoculture soils from the two regions both inhibited the seedling growth. The continuous cropping caused the reduction in the number of rhizosphere soil bacteria, accompanied by the increase of fungi and actinomycetes, resulted in the transition of soil type from "bacteria" to "fungi". Compared to the rotational soil, the activities of urease, sucrase and catalase in the continuous decreased significantly (P＜0.05), while the activities of polyphenoloxidase, cellulase and protease were slightly increased soil. But the phosphatase activity showed an opposite trend in the newly planted soil and monoculture soils from the two regions. Therefore, we speculated that autotoxicity changed the soil micro-ecology, affected soil enzyme activity, thereby reduced Rehmannia to absorb nutrients.
To establish the technical system of rapid propagation, the root tuber, leaves, apical buds, and radical buds of Rehmannia were cultured, and the results showed that using the radical buds as explants was best. Further, by optimizing the culture condition, the rapid propagation system of Rehmannia was established base on of radical buds inducing, which is MS+6-BA1+N AA0.1for induction, MS+6-BA0.3+NAA0.2for prolifera tion, and MS+PP3330.1for rooting. The cultivated aseptic seedlings grew in good condition and were used as material for the further study. The evaluation of potential autotoxicity showed that, fibrous root extract, replanted soil extracts and root exudates could inhibit the growth of Rehmannia glutinosa under the same concentration, and the fibrous root showed the strongest inhibitory effect. Four groups of autotoxic compounds from the aqueous extracts of the fibrous roots were isolated and characterized. The ethyl acetate extracts of these water-soluble compounds were further analyzed and separated into five fractions. Among them, the most autotoxic fraction (Fr3) was subjected to GC/MS analysis, resulting in32identified compounds. Based on literature, nine compounds were selected for testing their autotoxic effects on radicle growth. Seven out of the9compounds were phenolic, which significantly reduced radicle growth in a concentration-dependent manner. The other two were aliphatic compounds that showed a moderate inhibition effect at three concentrations. Concentration of these compounds in soil samples was determined by HPLC. Furthermore, the autotoxic compounds were also found in the top soil of the commercially cultivated Rehmannia fields. It appears that a close link exists between the autotoxic effects on the seedlings and the compounds extracted from fibrous roots of Rehmannia. Further analysis found that the autotoxic compounds not only resulted in membrane lipid peroxidation, the increasing of membrane permeability and antioxidant enzymes content, but also acted on plant indirectly by changing levels of endogenous phytohormones.
In this paper,66fungi strains were isolated from replanted soil and wilt plants of Rehmannia. Next, morphological observation and ITS sequence analysis were employed to identify the strains. The results showed that66fungi strains could be divided into phylums of Ascomycota, Zygomycota, Basidiomycota and Deuteromycotina. By the back infestation test, the strains of CCS038, CCS043, CCS025and CCS012were screened as host-specific pathogenic fungi of Rehmannia, they were grouped into Fusarium nivale, Fusarium oxysporum, Aspergillus flavus and A spergillus fum igatus re specti vely.
Finally, the synergistic effect of autotoxins and pathogenic fungi was discussed. The autotoxins especially ferulic acid and p-hydroxy benzoic acid not only promoted the hypha growth, spores proliferation directly, but also upgraded the expression of signal transduction system and nutrition metabolization related genes. In addition, the expression of some pathogenic genes were enhanced, which caused more sever damage to rehmannia.
道地产区与非道地产区地黄品质存在着差异,但正茬与重茬地黄的品质差异明显高于产区之间差异；两产区地黄的连作土壤均能抑制地黄幼苗生长；连作还造成根际土壤细菌数量减少,土壤真菌和放线菌数量增多,土壤微生物类型由“细菌型”向“真菌型”过渡；与正茬土壤相比,连作根际土壤中土壤脲酶、蔗糖酶和过氧化氢酶的活性显著下降,土壤多酚氧化酶、纤维素酶和蛋白酶的活性有所增加,但土壤磷酸酶活性在两个产区连作土壤中呈现相反的变化趋势。可见,地黄连作会引起土壤酶活性和土壤微生态区系的变化,影响土壤酶活性,降低了地黄对营养物质的吸收利用,进而造成地黄药用品质下降。
通过对以地黄块根、叶片、地黄顶芽及地黄新出根生芽等作为外殖体材料的无菌苗构建体系的比较性分析,认为新出根生芽可以作为地黄组织培养的良好外殖体材料。优化并建立了以MS+6-BA1mgL-1+NAA0. mgL-11为培养基的成苗诱导体系；确定了以MS+6-BA0.3mgL-1+NAA0.2mgL-1为培养基诱导的地黄丛生苗繁殖率高,苗体粗壮；经MS+PP3330.1mgL-1生根诱导的地黄无菌苗发根快而多,根部生长强壮,植株叶色浓绿,经炼苗后移栽可作为地黄连作障碍研究提供实验材料。
室内自毒作用潜力评价表明,相同浓度处理下的地黄须根提取物、地黄茬后土壤提取物、地黄根系分泌物均能抑制地黄生长,表现出自毒作用,且地黄须根提取物具有更强的化感抑制作用；95%乙醇进行地黄须根自毒物质浸提,经硅胶柱色谱分离,进行活性追踪实验确定部位Ⅲ为抑制活性最高的分离部位,硅烷化衍生化后进行气质联用分析得到32个化学物质；通过文献检阅后选择9个单体物质对地黄幼苗进行生物测试,结果表明所选的9个单体物质能较大程度上抑制地黄生长。进一步分析发现,地黄自毒物质不仅引起地黄体内膜脂过氧化物质的增高,导致细胞生物膜透性增强,保护酶活性持续升高,还影响地黄幼苗内源激素水平,造成植物伤害。
通过PDA平板培养法,从连作土壤中分离到病原真菌42株,从地黄病株中分离到病原真菌24株。这66株真菌型病原菌分属子囊菌门、结合菌门、担子菌门和半知菌门等4个门。采用回接发病实验鉴定到有4株菌地黄专化型病原菌,结合菌落形态、孢子形态和iTS区的PCR扩增后测序结果初步推断菌株CCS038为镰刀菌属(Fusaiunn)雪腐镰刀菌种(Fusarium nivale),菌株CCS043为镰刀菌属(Fusaiunn)尖孢镰刀菌种(Fusarium oxysporum),菌株CCS025为曲霉属(Aspergillus)黄曲霉菌(Aspergillus flavus),菌株CCS012为曲霉属(Aspergillus)烟曲霉菌(Aspergillus fumigatus)。
地黄须根自毒物质与病原菌存在协同作用,地黄须根自毒物质特别是阿魏酸和对羟基苯甲酸不仅能诱导地黄专化型病原菌的菌丝生长和孢子增殖和萌发,还通过增强真菌信号转导系统如SH3结构和F-box蛋白体系和分子伴侣等修饰原件加强机体内结构基因表达等过程的调控,还增强了对真菌营养代谢相关的调控,促进真菌对营养物质的利用,进而影响真菌形态建成,还能通过破坏宿主细胞壁和防御体系,增强病原菌对宿主的侵染能力,从而加剧对地黄的致害过程。
Rehmannia (Rehmannia glutinosa Libosch) is in the Scrophulariaceae family and is one of the most common and important medicinal herbal plants in China. However, the consecutively monocultured plants are prone to severe diseases resulting in reduced biomass, especially the tuberous products. In this study, the propagation system of Rehmannia was established, the autotoxicity of fibrous root was evaluated, and the host-specific pathogenic fungi were isolated from replanted soil and wilt plants of Rehmannia. Furthermore, the synergistic effect of autotoxins and phytopathogenic fungi on the consecutive monoculture of Rehmannia was explored. The main results were as follows:
In this study, the experiments were conducted to evaluate the quality of Rehmannia due to consecutive monoculture, and to explore the decline in the quality of Rehmannia from the sides of soil micro-ecology (soil microbes and enzyme activities). The results showed that, the quality of Rehmannia from the origin region (Jiaozuo, Henan province) and production region (Linfen, Shanxi Province) were different, but the differences between the newly planted soil and monoculture soil were significantly higher than the differences in regions. The monoculture soils from the two regions both inhibited the seedling growth. The continuous cropping caused the reduction in the number of rhizosphere soil bacteria, accompanied by the increase of fungi and actinomycetes, resulted in the transition of soil type from "bacteria" to "fungi". Compared to the rotational soil, the activities of urease, sucrase and catalase in the continuous decreased significantly (P＜0.05), while the activities of polyphenoloxidase, cellulase and protease were slightly increased soil. But the phosphatase activity showed an opposite trend in the newly planted soil and monoculture soils from the two regions. Therefore, we speculated that autotoxicity changed the soil micro-ecology, affected soil enzyme activity, thereby reduced Rehmannia to absorb nutrients.
To establish the technical system of rapid propagation, the root tuber, leaves, apical buds, and radical buds of Rehmannia were cultured, and the results showed that using the radical buds as explants was best. Further, by optimizing the culture condition, the rapid propagation system of Rehmannia was established base on of radical buds inducing, which is MS+6-BA1+N AA0.1for induction, MS+6-BA0.3+NAA0.2for prolifera tion, and MS+PP3330.1for rooting. The cultivated aseptic seedlings grew in good condition and were used as material for the further study. The evaluation of potential autotoxicity showed that, fibrous root extract, replanted soil extracts and root exudates could inhibit the growth of Rehmannia glutinosa under the same concentration, and the fibrous root showed the strongest inhibitory effect. Four groups of autotoxic compounds from the aqueous extracts of the fibrous roots were isolated and characterized. The ethyl acetate extracts of these water-soluble compounds were further analyzed and separated into five fractions. Among them, the most autotoxic fraction (Fr3) was subjected to GC/MS analysis, resulting in32identified compounds. Based on literature, nine compounds were selected for testing their autotoxic effects on radicle growth. Seven out of the9compounds were phenolic, which significantly reduced radicle growth in a concentration-dependent manner. The other two were aliphatic compounds that showed a moderate inhibition effect at three concentrations. Concentration of these compounds in soil samples was determined by HPLC. Furthermore, the autotoxic compounds were also found in the top soil of the commercially cultivated Rehmannia fields. It appears that a close link exists between the autotoxic effects on the seedlings and the compounds extracted from fibrous roots of Rehmannia. Further analysis found that the autotoxic compounds not only resulted in membrane lipid peroxidation, the increasing of membrane permeability and antioxidant enzymes content, but also acted on plant indirectly by changing levels of endogenous phytohormones.
In this paper,66fungi strains were isolated from replanted soil and wilt plants of Rehmannia. Next, morphological observation and ITS sequence analysis were employed to identify the strains. The results showed that66fungi strains could be divided into phylums of Ascomycota, Zygomycota, Basidiomycota and Deuteromycotina. By the back infestation test, the strains of CCS038, CCS043, CCS025and CCS012were screened as host-specific pathogenic fungi of Rehmannia, they were grouped into Fusarium nivale, Fusarium oxysporum, Aspergillus flavus and A spergillus fum igatus re specti vely.
Finally, the synergistic effect of autotoxins and pathogenic fungi was discussed. The autotoxins especially ferulic acid and p-hydroxy benzoic acid not only promoted the hypha growth, spores proliferation directly, but also upgraded the expression of signal transduction system and nutrition metabolization related genes. In addition, the expression of some pathogenic genes were enhanced, which caused more sever damage to rehmannia.
引文
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