牛蒡低聚果糖诱导植物抗病的分子机制研究
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摘要
作物产量受到多种因素的影响,其中病害是造成作物减产的一个重要因素。目前,人们对植物病害的防治主要依赖化学合成农药。但化学合成农药的滥用对环境和农产品安全造成了严重的影响。因此,筛选、研制对人畜安全、环境友好的生物农药已成为人们关注的热点。
当遭受病原体侵染时,植物自身可产生多种抗性响应,包括活性氧爆发、植保素合成、抗病相关蛋白基因表达等。这种抗性响应同样可被一类称为激发子(elicitor)的物质所诱导。其中寡糖分子作为一种激发子可激发多种抗性响应。牛蒡低聚果糖(BFO)是本课题组从牛蒡根中分离得到的一种聚合度为13的果糖低聚糖。前期的研究表明,该糖可诱导植物提高对多种病害的抗性。然而,对BFO诱导提高植物系统抗性的分子机制仍不清楚。本文利用分子生物学以及生物化学的手段对BFO诱导提高植物抗性的分子机制进行了系统的研究,以期为BFO作为抗病激发子的应用提供科学依据。此外,本文还以烟草为植物材料,研究了不同叶龄叶片对烟草花叶病毒(TMV)和BFO的抗性响应,以期为研究植物抗TMV机制提供理论依据。主要结果如下:
1.壳寡糖(CO)作为一种激发子已经被广泛研究,结果表明其可诱导提高对多种植物病害的抗性。本研究以CO为阳性对照,以蒸馏水(DW)为阴性对照研究了BFO对采后番茄抗病性的影响及其影响机制。结果表明BFO具有与CO类似的激发子功能,可有效提高番茄果实对自然病害的抗性以及对接种灰霉病的抗性。在处理后5天时,自然感病率和感病指数分别是DW处理的53.9%和45.2%,并且在处理后的15天内其自然感病率和感病指数都小于DW处理。接种灰霉病4天时BFO处理番茄的感病率和感病指数分别是DW处理的62.4%和48.1%。对其抗病机制进行初步的研究表明,BFO可快速诱导酸性PR蛋白(PR-1a、PR-2a、PR-3a)、碱性PR蛋白(PR-2b、PR-3b)以及苯丙氨酸解氨酶(PAL)基因的表达。另外BFO还可诱导过氧化物酶(POD)活性提高以及酚类物质累积,但BFO对多酚氧化酶(PPO)活性无显著影响。本研究还发现BFO和CO都不能诱导POD和PPO出现新的同工酶谱带,但BFO可诱导提高POD和PPO同工酶酶谱的强度,而CO仅诱导提高POD同工酶酶谱的强度,对PPO同工酶酶谱无明显影响。这些结果表明BFO可有效提高采后番茄的抗病性,其机制可能与其诱导抗病相关基因的表达、提高POD的活性以及诱导酚类物质在体内的累积有关。
2.以烟草为植物材料,利用分子生物学以及生物化学的方法研究了BFO对烟草TMV抗性的影响及其影响机制。结果表明,BFO处理可显著抑制TMV在烟草中的增殖,接种TMV 24h时DW处理叶片的TMV含量是BFO处理的7.0倍。此外,接种TMV 5天时DW处理叶片已出现明显的花斑症状,而BFO处理叶片仍表现正常,从而表明BFO处理增强了烟草对TMV的抗性。对其抗病机制进行初步的研究表明,BFO不仅可快速诱导酸性PR蛋白(包括PR-1a、PR-2、PR-3、PR-4以及PR-5)基因,还可诱导碱性PR-1蛋白基因以及抗病相关酶PAL和倍半萜环化酶(EAS)基因在烟草局部叶和系统叶中转录表达。H_2O_2和水杨酸(SA)是植物体内重要的信号分子,参与调控系统获得抗性(SAR)的发生,在本研究中BFO同样可快速诱导H_2O_2在烟草体内累积以及诱导SA和水杨酸葡萄糖苷(SAG)在烟草局部叶和系统叶中的生物合成。由于PR基因的表达常被作为诱导SAR发生的标记。因此,综合以上结果可以看出BFO提高烟草对TMV抗性的机制可能在于BFO通过H_2O_2以及SA介导了SAR的发生。
3.以烟草为植物材料,研究了不同叶龄叶片对TMV的抗性反应。结果表明,接种TMV48h时,老叶中TMV含量最高,大约是成熟叶和幼叶的8,1倍和35.1倍,从而说明幼叶对TMV的抗性最强,其次为成熟叶,老叶的抗性最弱。但是通过检测抗病相关蛋白(PR-1a、PR-2、PR-3、PR-4、PR-5、PAL)基因表达对TMV的响应发现,上述抗病相关蛋白基因在老叶中表达量最高,幼叶中表达量最弱。此外,烟草不同叶龄叶片经BFO或者DW处理后再接种TMV,发现BFO处理可以抑制TMV在不同叶龄烟草叶片中的增殖,并且接种TMV 48h时DW处理幼叶、成熟叶和老叶的TMV含量分别是BFO处理的1.4倍、1.9倍和11.0倍,即BFO处理对老叶的抑制效果最好,幼叶的抑制效果最弱。但接种TMV后上述抗病相关蛋白基因在BFO处理烟草叶片中的表达量低于DW处理叶片,从而都表现出TMV含量越高抗病相关蛋白表达量越高的现象。另外,我们的研究发现,BFO与TMV对烟草具有不同的抗性诱导效应,即BFO处理后上述抗病相关蛋白基因在幼叶中的表达量最高,在老叶中的表达量最低;而TMV处理后上述抗病相关蛋白基因在老叶中的表达量最高,在幼叶中的表达量最低。因此,综合以上结果可以看出,不同叶龄叶片对TMV具有不同的抗性,即幼叶的TMV抗性最强,老叶的抗性最弱;BFO处理可以抑制不同叶龄烟草叶片中TMV的增殖,但BFO对老叶的TMV抑制效果最好,幼叶最弱,并且BFO对烟草的诱导效应不同于TMV对烟草的诱导效应。
Plant pathogens can cause considerable loss of crop yields.At present,the plant diseases mainly controlled by using of synthetic fungicides.But the use of fungicides to control plant diseases has been restricted due to their effects on food and possible side-effects on human health.Thus,it has become a worldwide trend to explore new alternatives to reduce the use of synthetic fungicides
Plant could generate various defense responses to protect themselves from pathogens.Many studies have shown that these defense responses could also be induced by a class of molecules called elicitors.Oligosaccharides are a well characterized class of elicitors that trigger a large array of defense responses.Burdock fructooligosaccharide(BFO) is a fructosan oligomer that was first isolated from the root tissue of Arcitum lappa by our group.Previous studies have shown that BFO induces various defense responses in plants.However,the systemic defense responses and the underlying molecular mechanism that mediated by BFO in plants have not been elucidated.Therefore,we used biochemical and molecular approaches to investigate BFO-mediated non-specific defense activation in plants.In addition,we also studied the defense responses to TMV and BFO in tobacco leaves at different ages.The mainly results are as followed:
1.As an elicitor,chitosan oligosaccharide(CO) has been extensively researched, the results indicated that CO can induce resistance against multi-diseases in plants.In this study,the effects of BFO on the control of postharvest diseases in tomato fruit and the underlying mechanism were investigated with CO as a positive control and distilled water(DW) as a negative control.The results indicated that both BFO and CO could effectively inhibit natural postharvest diseases and reduce incidence of disease inoculated with Botrytis cinerea in tomato fruit.After treated 5 d by BFO the natural disease incidence and disease index were about 53.9%and 45.2%to DW treatment,and the natural disease incidence and disease index were still lower than DW treatment during the time after BFO treatment.In addition,inoculation with B. cinerea 4 d,the disease incidence and disease index were about 62.4%and 48.1%to DW treatment in BFO treated tomato fruit.BFO increased the mRNA level of genes encoding pathogenesis-related proteins(PRs),such as PR-1a,PR-2a(extracellularβ-1,3-glucanase),PR-2b(intracellularβ-1,3-glucanase),PR-3a(extracellular chitinase) and PR-3b(intracellular chitinase),and induced the mRNA accumulation of phenylalanine ammonia lyase(PAL) gene in tomato fruit.Furthermore,it evoked the activity of peroxides(POD) and enhanced the biosynthesis of phenolic compounds, but did not affect the activity of polyphenol oxidase(PPO).In this study,BFO and CO can not induce new isoenzyme bands of POD and PPO emerge,but the width of their isoenzyme bands were enhanced treated by BFO in tomato fruit.But interesting, treatment with CO could only strengthen the width the isoenzyme bands of POD,it has no effect on PPO in tomato fruit.These results suggest that controlling effects of BFO on both natural diseases and gray mould in tomato fruit might be attributed to enhance the transcript exprssion of defense related genes,increase the activity of POD and induce accumulation of phenolic compounds.
2.The effects of BFO on the control of TMV in tobacco leaves and the underlying molecular mechanism were investigated.The results indicated that treating cells with BFO 48 h before inoculation with TMV significantly inhibited TMV multiplication. The level of TMV-CP transcripts in DW treated leaves was about 7.0-fold higher than that in the BFO treated leaves 24 h after inoculation with TMV.In addition,about 5 d later,the leaves treated with DW had the typical mosaic pattern,while the leaves treated with BFO did not.These results suggested that BFO treatment improved the level of resistance to TMV.Our results also indicated that the levels of transcription of PR protein genes,including acidic PRs(PR-1a,PR-2,PR-3,PR-4,PR-5) and basic PR-1 and defense-related enzymes[PAL and 5-epi-aristolochene synthase,(EAS)] genes were increased both locally and systemically by BFO treatment,in correlation with the induction of resistance against tobacco mosaic virus(TMV).In plants,H_2O_2 and salicylic acid(SA) act as signal molecules to induce systemic acquired resistance (SAR).And interestingly,BFO also induced accumulation of H_2O_2,SA and SA 2-O-β-D-glucoside(SAG) in tobacco leaves.In addition,the expression of PR genes is usually taken as a marker for systemic acquired resistance(SAR).Therefore,these results strongly suggested that BFO activated the defense responses and the mode of action of BFO might have been induction of SAR through H_2O_2 and SA signal molecules.
3.The defense responses to TMV in tobacco leaves at different ages(young,adult and old) were studied.The results indicated that the content of TMV in old leaves was the most at 48 h after inoculation with TMV among of them,and about 8.1 and 35.1-fold higher than that in adult leaves and in young leaves at that time,respectively. Therefore,there is the strongest resistance against TMV in young leaves,followed by adult leaves,and old leaves have the weakest resistace.Followed,we investigated the response of defense related genes(PR-1a,PR-2,PR-3,PR-4,PR-5,PAL )to TMV in tobacco leaves at different ages.The results shown that the mRNA levels of above defense related genes all could be induced by TMV,but the responses intensity to TMV of the above genes was the weakest in young leaves,while it was the strongest in old leaves.In addition,we further studied the defense responses to BFO in tobacco leaves at different ages,and further investigated the effects on the resistance against TMV treatment with BFO in tobacco leaves at different ages.The results indicated that the response of the above defense relate genes was the strongest in young leaves, followed by adult leaves,and the old leaves is the weakest.But the effects of BFO on the resistance against TMV were the strongest in old leaves,followed by adult leaves, and the young leaves is the weakest.Inoculation with TMV after treated by BFO 48 h, the content of TMV was smaller than DW treated.At 48h inoculation with TMV,the content of TMV in DW treated tobacco young leaves,adult leaves and old leaves, were about 1.4,1.9 and 11.0-fold higher than BFO treated leaves,respectively.This result suggests,the effect of BFO treatment on the resistace against TMV was the strongest in old leaves,followed by adult leaves,and the young leaves is the weakest. In oppositing leaves at different ages,the transcripts level of the above defense related genes and the content of TMV in leaves treated by BFO both were smaller than the leaves treated by DW.Together above results,we come to the conclusion that there are different resistance to TMV in tobacco leaves at different ages,and the resistace against TMV was the strongest in young leaves,followed by adult leaves,and the old leaves is the weakest.In addition,the results indicated that treatment with BFO could inhibit the multiplication of TMV in tobacco leaves at different ages,and there is the best effect in old leaves,the young leaves are the worst.The results also indicated that there is different defense response between BFO and TMV in tobacco leaves.
当遭受病原体侵染时,植物自身可产生多种抗性响应,包括活性氧爆发、植保素合成、抗病相关蛋白基因表达等。这种抗性响应同样可被一类称为激发子(elicitor)的物质所诱导。其中寡糖分子作为一种激发子可激发多种抗性响应。牛蒡低聚果糖(BFO)是本课题组从牛蒡根中分离得到的一种聚合度为13的果糖低聚糖。前期的研究表明,该糖可诱导植物提高对多种病害的抗性。然而,对BFO诱导提高植物系统抗性的分子机制仍不清楚。本文利用分子生物学以及生物化学的手段对BFO诱导提高植物抗性的分子机制进行了系统的研究,以期为BFO作为抗病激发子的应用提供科学依据。此外,本文还以烟草为植物材料,研究了不同叶龄叶片对烟草花叶病毒(TMV)和BFO的抗性响应,以期为研究植物抗TMV机制提供理论依据。主要结果如下:
1.壳寡糖(CO)作为一种激发子已经被广泛研究,结果表明其可诱导提高对多种植物病害的抗性。本研究以CO为阳性对照,以蒸馏水(DW)为阴性对照研究了BFO对采后番茄抗病性的影响及其影响机制。结果表明BFO具有与CO类似的激发子功能,可有效提高番茄果实对自然病害的抗性以及对接种灰霉病的抗性。在处理后5天时,自然感病率和感病指数分别是DW处理的53.9%和45.2%,并且在处理后的15天内其自然感病率和感病指数都小于DW处理。接种灰霉病4天时BFO处理番茄的感病率和感病指数分别是DW处理的62.4%和48.1%。对其抗病机制进行初步的研究表明,BFO可快速诱导酸性PR蛋白(PR-1a、PR-2a、PR-3a)、碱性PR蛋白(PR-2b、PR-3b)以及苯丙氨酸解氨酶(PAL)基因的表达。另外BFO还可诱导过氧化物酶(POD)活性提高以及酚类物质累积,但BFO对多酚氧化酶(PPO)活性无显著影响。本研究还发现BFO和CO都不能诱导POD和PPO出现新的同工酶谱带,但BFO可诱导提高POD和PPO同工酶酶谱的强度,而CO仅诱导提高POD同工酶酶谱的强度,对PPO同工酶酶谱无明显影响。这些结果表明BFO可有效提高采后番茄的抗病性,其机制可能与其诱导抗病相关基因的表达、提高POD的活性以及诱导酚类物质在体内的累积有关。
2.以烟草为植物材料,利用分子生物学以及生物化学的方法研究了BFO对烟草TMV抗性的影响及其影响机制。结果表明,BFO处理可显著抑制TMV在烟草中的增殖,接种TMV 24h时DW处理叶片的TMV含量是BFO处理的7.0倍。此外,接种TMV 5天时DW处理叶片已出现明显的花斑症状,而BFO处理叶片仍表现正常,从而表明BFO处理增强了烟草对TMV的抗性。对其抗病机制进行初步的研究表明,BFO不仅可快速诱导酸性PR蛋白(包括PR-1a、PR-2、PR-3、PR-4以及PR-5)基因,还可诱导碱性PR-1蛋白基因以及抗病相关酶PAL和倍半萜环化酶(EAS)基因在烟草局部叶和系统叶中转录表达。H_2O_2和水杨酸(SA)是植物体内重要的信号分子,参与调控系统获得抗性(SAR)的发生,在本研究中BFO同样可快速诱导H_2O_2在烟草体内累积以及诱导SA和水杨酸葡萄糖苷(SAG)在烟草局部叶和系统叶中的生物合成。由于PR基因的表达常被作为诱导SAR发生的标记。因此,综合以上结果可以看出BFO提高烟草对TMV抗性的机制可能在于BFO通过H_2O_2以及SA介导了SAR的发生。
3.以烟草为植物材料,研究了不同叶龄叶片对TMV的抗性反应。结果表明,接种TMV48h时,老叶中TMV含量最高,大约是成熟叶和幼叶的8,1倍和35.1倍,从而说明幼叶对TMV的抗性最强,其次为成熟叶,老叶的抗性最弱。但是通过检测抗病相关蛋白(PR-1a、PR-2、PR-3、PR-4、PR-5、PAL)基因表达对TMV的响应发现,上述抗病相关蛋白基因在老叶中表达量最高,幼叶中表达量最弱。此外,烟草不同叶龄叶片经BFO或者DW处理后再接种TMV,发现BFO处理可以抑制TMV在不同叶龄烟草叶片中的增殖,并且接种TMV 48h时DW处理幼叶、成熟叶和老叶的TMV含量分别是BFO处理的1.4倍、1.9倍和11.0倍,即BFO处理对老叶的抑制效果最好,幼叶的抑制效果最弱。但接种TMV后上述抗病相关蛋白基因在BFO处理烟草叶片中的表达量低于DW处理叶片,从而都表现出TMV含量越高抗病相关蛋白表达量越高的现象。另外,我们的研究发现,BFO与TMV对烟草具有不同的抗性诱导效应,即BFO处理后上述抗病相关蛋白基因在幼叶中的表达量最高,在老叶中的表达量最低;而TMV处理后上述抗病相关蛋白基因在老叶中的表达量最高,在幼叶中的表达量最低。因此,综合以上结果可以看出,不同叶龄叶片对TMV具有不同的抗性,即幼叶的TMV抗性最强,老叶的抗性最弱;BFO处理可以抑制不同叶龄烟草叶片中TMV的增殖,但BFO对老叶的TMV抑制效果最好,幼叶最弱,并且BFO对烟草的诱导效应不同于TMV对烟草的诱导效应。
Plant pathogens can cause considerable loss of crop yields.At present,the plant diseases mainly controlled by using of synthetic fungicides.But the use of fungicides to control plant diseases has been restricted due to their effects on food and possible side-effects on human health.Thus,it has become a worldwide trend to explore new alternatives to reduce the use of synthetic fungicides
Plant could generate various defense responses to protect themselves from pathogens.Many studies have shown that these defense responses could also be induced by a class of molecules called elicitors.Oligosaccharides are a well characterized class of elicitors that trigger a large array of defense responses.Burdock fructooligosaccharide(BFO) is a fructosan oligomer that was first isolated from the root tissue of Arcitum lappa by our group.Previous studies have shown that BFO induces various defense responses in plants.However,the systemic defense responses and the underlying molecular mechanism that mediated by BFO in plants have not been elucidated.Therefore,we used biochemical and molecular approaches to investigate BFO-mediated non-specific defense activation in plants.In addition,we also studied the defense responses to TMV and BFO in tobacco leaves at different ages.The mainly results are as followed:
1.As an elicitor,chitosan oligosaccharide(CO) has been extensively researched, the results indicated that CO can induce resistance against multi-diseases in plants.In this study,the effects of BFO on the control of postharvest diseases in tomato fruit and the underlying mechanism were investigated with CO as a positive control and distilled water(DW) as a negative control.The results indicated that both BFO and CO could effectively inhibit natural postharvest diseases and reduce incidence of disease inoculated with Botrytis cinerea in tomato fruit.After treated 5 d by BFO the natural disease incidence and disease index were about 53.9%and 45.2%to DW treatment,and the natural disease incidence and disease index were still lower than DW treatment during the time after BFO treatment.In addition,inoculation with B. cinerea 4 d,the disease incidence and disease index were about 62.4%and 48.1%to DW treatment in BFO treated tomato fruit.BFO increased the mRNA level of genes encoding pathogenesis-related proteins(PRs),such as PR-1a,PR-2a(extracellularβ-1,3-glucanase),PR-2b(intracellularβ-1,3-glucanase),PR-3a(extracellular chitinase) and PR-3b(intracellular chitinase),and induced the mRNA accumulation of phenylalanine ammonia lyase(PAL) gene in tomato fruit.Furthermore,it evoked the activity of peroxides(POD) and enhanced the biosynthesis of phenolic compounds, but did not affect the activity of polyphenol oxidase(PPO).In this study,BFO and CO can not induce new isoenzyme bands of POD and PPO emerge,but the width of their isoenzyme bands were enhanced treated by BFO in tomato fruit.But interesting, treatment with CO could only strengthen the width the isoenzyme bands of POD,it has no effect on PPO in tomato fruit.These results suggest that controlling effects of BFO on both natural diseases and gray mould in tomato fruit might be attributed to enhance the transcript exprssion of defense related genes,increase the activity of POD and induce accumulation of phenolic compounds.
2.The effects of BFO on the control of TMV in tobacco leaves and the underlying molecular mechanism were investigated.The results indicated that treating cells with BFO 48 h before inoculation with TMV significantly inhibited TMV multiplication. The level of TMV-CP transcripts in DW treated leaves was about 7.0-fold higher than that in the BFO treated leaves 24 h after inoculation with TMV.In addition,about 5 d later,the leaves treated with DW had the typical mosaic pattern,while the leaves treated with BFO did not.These results suggested that BFO treatment improved the level of resistance to TMV.Our results also indicated that the levels of transcription of PR protein genes,including acidic PRs(PR-1a,PR-2,PR-3,PR-4,PR-5) and basic PR-1 and defense-related enzymes[PAL and 5-epi-aristolochene synthase,(EAS)] genes were increased both locally and systemically by BFO treatment,in correlation with the induction of resistance against tobacco mosaic virus(TMV).In plants,H_2O_2 and salicylic acid(SA) act as signal molecules to induce systemic acquired resistance (SAR).And interestingly,BFO also induced accumulation of H_2O_2,SA and SA 2-O-β-D-glucoside(SAG) in tobacco leaves.In addition,the expression of PR genes is usually taken as a marker for systemic acquired resistance(SAR).Therefore,these results strongly suggested that BFO activated the defense responses and the mode of action of BFO might have been induction of SAR through H_2O_2 and SA signal molecules.
3.The defense responses to TMV in tobacco leaves at different ages(young,adult and old) were studied.The results indicated that the content of TMV in old leaves was the most at 48 h after inoculation with TMV among of them,and about 8.1 and 35.1-fold higher than that in adult leaves and in young leaves at that time,respectively. Therefore,there is the strongest resistance against TMV in young leaves,followed by adult leaves,and old leaves have the weakest resistace.Followed,we investigated the response of defense related genes(PR-1a,PR-2,PR-3,PR-4,PR-5,PAL )to TMV in tobacco leaves at different ages.The results shown that the mRNA levels of above defense related genes all could be induced by TMV,but the responses intensity to TMV of the above genes was the weakest in young leaves,while it was the strongest in old leaves.In addition,we further studied the defense responses to BFO in tobacco leaves at different ages,and further investigated the effects on the resistance against TMV treatment with BFO in tobacco leaves at different ages.The results indicated that the response of the above defense relate genes was the strongest in young leaves, followed by adult leaves,and the old leaves is the weakest.But the effects of BFO on the resistance against TMV were the strongest in old leaves,followed by adult leaves, and the young leaves is the weakest.Inoculation with TMV after treated by BFO 48 h, the content of TMV was smaller than DW treated.At 48h inoculation with TMV,the content of TMV in DW treated tobacco young leaves,adult leaves and old leaves, were about 1.4,1.9 and 11.0-fold higher than BFO treated leaves,respectively.This result suggests,the effect of BFO treatment on the resistace against TMV was the strongest in old leaves,followed by adult leaves,and the young leaves is the weakest. In oppositing leaves at different ages,the transcripts level of the above defense related genes and the content of TMV in leaves treated by BFO both were smaller than the leaves treated by DW.Together above results,we come to the conclusion that there are different resistance to TMV in tobacco leaves at different ages,and the resistace against TMV was the strongest in young leaves,followed by adult leaves,and the old leaves is the weakest.In addition,the results indicated that treatment with BFO could inhibit the multiplication of TMV in tobacco leaves at different ages,and there is the best effect in old leaves,the young leaves are the worst.The results also indicated that there is different defense response between BFO and TMV in tobacco leaves.
引文
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