氯化钾抑制玉米茎腐病发生的机理研究
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摘要
玉米茎腐病在我国玉米主产区发生严重。它作为一种典型的土传病害,在防治上难度较大,应用一般的化学农药、种子包衣等均难以有效控制该病的发生。有研究发现,钾肥能够显著降低玉米茎腐病的发生率,但是关于钾在抑制玉米茎腐病方面的作用机制尚不清楚。本研究采用12年氯化钾肥长期定位试验研究了氯化钾对茎腐病的抑制效果;结合室内生物培养试验研究了不同钾量级土壤浸提液对主要致病菌禾谷镰刀菌(Fusarium graminearum)生长的影响;利用平板计数法分析了长期施用氯化钾对玉米根际土壤微生物区系的变化。通过钾和氯不同量级的大田试验和氯化钾长期定位试验,进一步比较了氯化钾中钾和氯的抑病作用;结合室内生理生化分析方法研究了钾和氯对玉米各生育时期植株体内糖分分配的影响,并结合抽雄期接种处理探讨了钾对病原菌侵染前后酚类物质代谢的影响。本研究通过水培试验,利用高压液相色谱定量分析了钾对玉米苗期根系分泌物中各组分含量的影响;并结合室内生物培养试验,研究了根系分泌物中糖和酚酸对禾谷镰刀菌生长的影响。采用土培试验,利用透射电镜对接种后寄主与病原菌互作过程中根尖细胞超微结构的变化进行了观察。本文主要从施用氯化钾对土壤微生态和植物生理生化代谢影响的角度,对氯化钾抑制玉米茎腐病发生的机理进行系统研究和深入探讨,取得以下主要研究结果:
1.在缺钾土壤上,施用氯化钾和硝酸钾均能够显著抑制玉米茎腐病的发生,而氯素的抑病作用不明显。在缺钾条件下,抗病品种(吉单180)较感病品种(吉单327)具有更强的吸钾能力。玉米各生育时期的根、苇和叶部钾素含量均与玉米茎腐病的发生率存在显著负相关关系,茎腐病发生率随着植株体含钾量的提高而显著降低。
2.室内培养试验发现,一定浓度的氯化钾不能直接抑制禾谷镰刀菌(Fusarium graminearum)的生长,长期施用氯化钾的土壤浸提液对其具有明显的抑制作用。与不施钾处理相比,长期施用氯化钾对玉米生育前期根际土壤细菌数目无明显影响,适量的氯化钾可以显著增加根际土壤放线菌数目和真菌数目,且施钾处理的苗期根际土壤真菌以木霉菌占优势。而且施钾处理与不施钾处理的根际土壤真菌和放线菌数目差异显著时期正是玉米茎腐病病原菌的主要侵染期(生育前期)。说明长期施用氯化钾引起的根际土壤微生物区系的变化可能与病原菌对寄主根系的侵染存在一定的联系。
3.与缺钾处理相比,在以硝态氮为主要氮源的条件下,氯化钾和硝酸钾能够抑制根系糖的分泌,促进有机酸和酚酸的分泌。缺钾处理导致根系总糖尤其是还原糖的分泌量显著增加,有利于禾谷镰刀菌在体外的生长繁殖,可能由此提高了病原菌与寄主识别的几率,利于病原菌的侵染。阿魏酸是玉米根系分泌的主要酚酸组分,抗病品种吉单180阿魏酸分泌量明显高于感病品种吉单327,且随着钾的供应而大幅度提高。抗病品种绿原酸分泌量明显低于感病品种,供钾导致两个品种根系绿原酸分泌量下降。体外培养试验发现,阿魏酸和绿原酸均能够抑制Fusarium graminearum的生长,但阿魏酸的抑菌效果远远大于绿原酸。这说明钾主要通过阿魏酸分泌量的提高抑制根际禾谷镰刀菌的生长繁殖,从而可能降低其对寄主的侵染几率。
钾能够促进寄主细胞壁加厚,形成阻止病原菌侵染的第一道屏障;即使在病原菌侵染后,钾也能够刺激寄主细胞迅速分泌降解物质,抑制病原菌的进一步扩展。缺钾则导致寄主细胞内线粒
Stalk rot is one of the most widespread and destructive disease that reduces maize yield and quality. As a kind of soil-borne fungal disease, maize stalk rot is difficult to control by conventional strategies such as the use of synthetic fungicides and fungicidal seed treatment. Many studies confirm that application of potassium is an effective method to reduce the stalk rot incidence of maize. However, little attention has been paid to the mechanisms of suppression of corn stalk rot by potassium. A 12-year long-term field experiment with different rates of potassium chloride (KC1) application was used to investigate the suppression effect of KC1 on stalk rot, the effect of soil extracts on the growth of Fusarium graminearum, the most common stalk rot fungi in this area, and the changes of microflora in rhizosphere with plate counting method. A separate field trial with different rates of potassium and chloride application was conducted to valuate the suppression effects of potassium and chloride from KC1 and study the effects of potassium on the distribution of soluble carbohydrate in plant at the growth stages, and the phenolic metabolism in stalk pith tissue of the second internode above ground after inoculation affected by potassium. Hydroponic experiment was conducted to measure the influence of potassium supply on the root exudates using high performance liquid chromatography, and the effects of sugar and phenolic compounds on the growth of F. graminearum was analyzed with plate culture method. Pot trial under greenhouse condition was carried out to examine the alteration of ultrastructure of seedling root tissue 4 days after inoculation with transmission electron microscope. This paper discussed the possible mechanism of potassium chloride suppressing stalk rot of maize from the aspects of microecology of rhizosphere, and plant physiology and biochemistry affected by potassium chloride. The main results are as following:1. In K-deficiency soil, potassium applied as KC1 and KNO_3 decreased the incidence of maize stalk rot significantly, but chloride applied as CaCl_2 had no obvious effect on reducing the incidence of maize stalk rot compared to the control. Resistant cultivar Jidan180 had a higher potassium uptake capacity than susceptible cultivar Jidan327. The significant negative correlations were found between the stalk rot incidence and the K content in root, stem and ear-leaf of maize at the growth stages.2. KC1 addition to the PDA medium could not directly suppress F. graminearum development. Soil extracts from soil with long-term KC1 treatments suppressed F. graminearum development more significantly, compared with that from the treatment of no KC1 supply. Long-term application of KC1 increased the populations of rhizosphere fungi and actinomyces at the early growth stages. However, there was no significant difference existed in rhizosphere bacteria population among the treatments. The sensitive infection stages of pathogen to maize were consisted with the stages when the significant difference in rhizosphere fungi and actinomyces populations. And at seedling stage, Trichoderma was the fungi dominating in the rhizosphere of maize with long-term KC1 application. Therefore, the change in rhizosphere microflora (especially fungi) perhaps was one of the mechanisms of suppressing maize stalk rot by KC1 application.3. Potassium supply decreased the total amount of sugar exudates and stimulated the release of
organic acids and phenolic acids under nitrate as the main nitrogen source. Potassium significantly increased the ferulic acid content and decreased the chlorogenic acid content. The amount of ferulic acid exudates by root of resistance cultivar was higher than that of susceptible cultivar, while a reverse trend was found in chlorogenic acid. In vitro, the growth of Fusarium graminearum was stimulated by sugar and inhibited by ferulic acid and chlorogenic acid, and the suppression effect of ferulic acid was more effective than chlorogenic acid. The decrease of sugar content and the increase of the amount of ferulic acid in seedling root exudates with potassium supply maybe help reduce the infection probability by F. graminearum. The cell wall of maize root tip with potassium supply became thicker than that without K application. Once the cell of maize root tip with potassium supply was infected, it would secreted material for degrading fungi hypha, and inhibited the rapid development of pathology in host. Mitochondria accumulation which marked the abnormal of respiration was found in the cell of seedling root with K deficiency.4. Potassium stimulated carbohydrate translocation to root and stem, and increased the sugar content after the tasselling stage. Sugar contents in root at the filling stage and those in stem at the dough stage were positively correlated with the resistance to stalk rot. The total soluble sugar content in root or stem could be used as a better indicator of resistance to maize stalk rot compared to the sucrose content and the reducing sugar content. Enhancement of sugar metabolism by potassium was responsible for the resistance to maize stalk rot.5. The initial level of lignin in the susceptible cultivar was higher than that in the resistant cultivar, while the content of induced lignin by inoculation in susceptible cultivar was lower than that in resistant cultivar. Therefore, it was not the initial level of lignin but the induced level of lignin could indicate the resistance to stalk rot. Infection of F. graminearum increased the accumulation of induce lignin and triggered rapid increase in activities of phenylalanine ammonia-lyase (PAL), tyrosine ammonia-lyase (TAL), peroxidase (POD) and cinnamyl alcohol dehydrogenase (CAD) in the stalk pith tissue of maize with potassium application, compared with the K-deficiency plant. These results implied that after inoculation, potassium could stimulate the rapid expression of defence genes, increase the activities of defense enzymes, promote the synthesis of inducing soluble phenolic compounds and then enhance the resistance of maize plants against stalk rot.6. The background level of phenolics and the induced phenolics by inoculation in the susceptible cultivar was higher than that in the resistant cultivar. Potassium supply increased the rapid accumulation of induced phenolics and increased the activity of polyphenoloxidase (PPO), stimulated the translation of phenolics compounds to quinone Regardless of inoculation, the concentration of chlorogenic acid increased with the increase of potassium application rates, which was the predominant phenolic compound in maize stalk pith tissue, and chlorogenic acid proved to inhibit the growth of F. graminearum in vitro. The results suggested that chlorogenic acid may play a role in inhibiting the development of F. graminearum in the stalk pith tissue of maizeIn conclusion, the possible mechanisms of potassium chloride suppressing stalk rot of maize could be summarized as follows: 1) KC1 impeded directly the infection of pathology by making the cell wall
thicker, and decreased indirectly the probability of infection of F. graminearum by changing the root exudates and microflora in rhizosphere. 2) KCl maintained plant physiology vigor higher at the later growth stage by accelerating the transportation of carbohydrate and increasing sugar available to the root and stem tissue after tasselling stage. 3) KCl enhanced the resistance of maize against stalk rot through stimulating the rapid expression of defence genes, furthermore increasing the activities of defense enzymes, and then promoting the synthesis of inducing soluble phenolic compounds after infection by pathology. All of those may contribute to the decrease in incidence of maize stalk rot.
1.在缺钾土壤上,施用氯化钾和硝酸钾均能够显著抑制玉米茎腐病的发生,而氯素的抑病作用不明显。在缺钾条件下,抗病品种(吉单180)较感病品种(吉单327)具有更强的吸钾能力。玉米各生育时期的根、苇和叶部钾素含量均与玉米茎腐病的发生率存在显著负相关关系,茎腐病发生率随着植株体含钾量的提高而显著降低。
2.室内培养试验发现,一定浓度的氯化钾不能直接抑制禾谷镰刀菌(Fusarium graminearum)的生长,长期施用氯化钾的土壤浸提液对其具有明显的抑制作用。与不施钾处理相比,长期施用氯化钾对玉米生育前期根际土壤细菌数目无明显影响,适量的氯化钾可以显著增加根际土壤放线菌数目和真菌数目,且施钾处理的苗期根际土壤真菌以木霉菌占优势。而且施钾处理与不施钾处理的根际土壤真菌和放线菌数目差异显著时期正是玉米茎腐病病原菌的主要侵染期(生育前期)。说明长期施用氯化钾引起的根际土壤微生物区系的变化可能与病原菌对寄主根系的侵染存在一定的联系。
3.与缺钾处理相比,在以硝态氮为主要氮源的条件下,氯化钾和硝酸钾能够抑制根系糖的分泌,促进有机酸和酚酸的分泌。缺钾处理导致根系总糖尤其是还原糖的分泌量显著增加,有利于禾谷镰刀菌在体外的生长繁殖,可能由此提高了病原菌与寄主识别的几率,利于病原菌的侵染。阿魏酸是玉米根系分泌的主要酚酸组分,抗病品种吉单180阿魏酸分泌量明显高于感病品种吉单327,且随着钾的供应而大幅度提高。抗病品种绿原酸分泌量明显低于感病品种,供钾导致两个品种根系绿原酸分泌量下降。体外培养试验发现,阿魏酸和绿原酸均能够抑制Fusarium graminearum的生长,但阿魏酸的抑菌效果远远大于绿原酸。这说明钾主要通过阿魏酸分泌量的提高抑制根际禾谷镰刀菌的生长繁殖,从而可能降低其对寄主的侵染几率。
钾能够促进寄主细胞壁加厚,形成阻止病原菌侵染的第一道屏障;即使在病原菌侵染后,钾也能够刺激寄主细胞迅速分泌降解物质,抑制病原菌的进一步扩展。缺钾则导致寄主细胞内线粒
Stalk rot is one of the most widespread and destructive disease that reduces maize yield and quality. As a kind of soil-borne fungal disease, maize stalk rot is difficult to control by conventional strategies such as the use of synthetic fungicides and fungicidal seed treatment. Many studies confirm that application of potassium is an effective method to reduce the stalk rot incidence of maize. However, little attention has been paid to the mechanisms of suppression of corn stalk rot by potassium. A 12-year long-term field experiment with different rates of potassium chloride (KC1) application was used to investigate the suppression effect of KC1 on stalk rot, the effect of soil extracts on the growth of Fusarium graminearum, the most common stalk rot fungi in this area, and the changes of microflora in rhizosphere with plate counting method. A separate field trial with different rates of potassium and chloride application was conducted to valuate the suppression effects of potassium and chloride from KC1 and study the effects of potassium on the distribution of soluble carbohydrate in plant at the growth stages, and the phenolic metabolism in stalk pith tissue of the second internode above ground after inoculation affected by potassium. Hydroponic experiment was conducted to measure the influence of potassium supply on the root exudates using high performance liquid chromatography, and the effects of sugar and phenolic compounds on the growth of F. graminearum was analyzed with plate culture method. Pot trial under greenhouse condition was carried out to examine the alteration of ultrastructure of seedling root tissue 4 days after inoculation with transmission electron microscope. This paper discussed the possible mechanism of potassium chloride suppressing stalk rot of maize from the aspects of microecology of rhizosphere, and plant physiology and biochemistry affected by potassium chloride. The main results are as following:1. In K-deficiency soil, potassium applied as KC1 and KNO_3 decreased the incidence of maize stalk rot significantly, but chloride applied as CaCl_2 had no obvious effect on reducing the incidence of maize stalk rot compared to the control. Resistant cultivar Jidan180 had a higher potassium uptake capacity than susceptible cultivar Jidan327. The significant negative correlations were found between the stalk rot incidence and the K content in root, stem and ear-leaf of maize at the growth stages.2. KC1 addition to the PDA medium could not directly suppress F. graminearum development. Soil extracts from soil with long-term KC1 treatments suppressed F. graminearum development more significantly, compared with that from the treatment of no KC1 supply. Long-term application of KC1 increased the populations of rhizosphere fungi and actinomyces at the early growth stages. However, there was no significant difference existed in rhizosphere bacteria population among the treatments. The sensitive infection stages of pathogen to maize were consisted with the stages when the significant difference in rhizosphere fungi and actinomyces populations. And at seedling stage, Trichoderma was the fungi dominating in the rhizosphere of maize with long-term KC1 application. Therefore, the change in rhizosphere microflora (especially fungi) perhaps was one of the mechanisms of suppressing maize stalk rot by KC1 application.3. Potassium supply decreased the total amount of sugar exudates and stimulated the release of
organic acids and phenolic acids under nitrate as the main nitrogen source. Potassium significantly increased the ferulic acid content and decreased the chlorogenic acid content. The amount of ferulic acid exudates by root of resistance cultivar was higher than that of susceptible cultivar, while a reverse trend was found in chlorogenic acid. In vitro, the growth of Fusarium graminearum was stimulated by sugar and inhibited by ferulic acid and chlorogenic acid, and the suppression effect of ferulic acid was more effective than chlorogenic acid. The decrease of sugar content and the increase of the amount of ferulic acid in seedling root exudates with potassium supply maybe help reduce the infection probability by F. graminearum. The cell wall of maize root tip with potassium supply became thicker than that without K application. Once the cell of maize root tip with potassium supply was infected, it would secreted material for degrading fungi hypha, and inhibited the rapid development of pathology in host. Mitochondria accumulation which marked the abnormal of respiration was found in the cell of seedling root with K deficiency.4. Potassium stimulated carbohydrate translocation to root and stem, and increased the sugar content after the tasselling stage. Sugar contents in root at the filling stage and those in stem at the dough stage were positively correlated with the resistance to stalk rot. The total soluble sugar content in root or stem could be used as a better indicator of resistance to maize stalk rot compared to the sucrose content and the reducing sugar content. Enhancement of sugar metabolism by potassium was responsible for the resistance to maize stalk rot.5. The initial level of lignin in the susceptible cultivar was higher than that in the resistant cultivar, while the content of induced lignin by inoculation in susceptible cultivar was lower than that in resistant cultivar. Therefore, it was not the initial level of lignin but the induced level of lignin could indicate the resistance to stalk rot. Infection of F. graminearum increased the accumulation of induce lignin and triggered rapid increase in activities of phenylalanine ammonia-lyase (PAL), tyrosine ammonia-lyase (TAL), peroxidase (POD) and cinnamyl alcohol dehydrogenase (CAD) in the stalk pith tissue of maize with potassium application, compared with the K-deficiency plant. These results implied that after inoculation, potassium could stimulate the rapid expression of defence genes, increase the activities of defense enzymes, promote the synthesis of inducing soluble phenolic compounds and then enhance the resistance of maize plants against stalk rot.6. The background level of phenolics and the induced phenolics by inoculation in the susceptible cultivar was higher than that in the resistant cultivar. Potassium supply increased the rapid accumulation of induced phenolics and increased the activity of polyphenoloxidase (PPO), stimulated the translation of phenolics compounds to quinone Regardless of inoculation, the concentration of chlorogenic acid increased with the increase of potassium application rates, which was the predominant phenolic compound in maize stalk pith tissue, and chlorogenic acid proved to inhibit the growth of F. graminearum in vitro. The results suggested that chlorogenic acid may play a role in inhibiting the development of F. graminearum in the stalk pith tissue of maizeIn conclusion, the possible mechanisms of potassium chloride suppressing stalk rot of maize could be summarized as follows: 1) KC1 impeded directly the infection of pathology by making the cell wall
thicker, and decreased indirectly the probability of infection of F. graminearum by changing the root exudates and microflora in rhizosphere. 2) KCl maintained plant physiology vigor higher at the later growth stage by accelerating the transportation of carbohydrate and increasing sugar available to the root and stem tissue after tasselling stage. 3) KCl enhanced the resistance of maize against stalk rot through stimulating the rapid expression of defence genes, furthermore increasing the activities of defense enzymes, and then promoting the synthesis of inducing soluble phenolic compounds after infection by pathology. All of those may contribute to the decrease in incidence of maize stalk rot.
引文
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