外来植物南美蟛蜞菊非结构性抗病机制及对其成功入侵的贡献
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摘要
生物入侵为当下全球变化的主要内容之一,也是重要的国际环境生态问题,威胁着全球的环境生态和经济发展。虽然关于生物入侵的机制研究已有不少报道,却极少见系统研究入侵植物的抗病性机制的报道。本研究从群落学、次生代谢产物、植物抗病基因(R-gene)、植物内生菌等四个方面首次对入侵植物南美蟛蜞菊(Wedelia trilobata (L.) Hitchc)非结构性抗病机制进行了研究,并探讨了其对南美蟛蜞菊成功入侵的可能贡献,具体如下：
1.通过传统的群落学调查方法,确定了入侵植物南美蟛蜞菊在野外很少染病的情况,南美蟛蜞菊的入侵会显著降低其入侵植物群落物种多样性。并且,经济活动加剧了南美蟛蜞菊的入侵。
2.从生化水平探讨南美蟛蜞菊抗病的机制,确定了南美蟛蜞菊不同组织的次生代谢产物既能够抑制立枯丝核及香蕉枯萎病等植物病原真菌,亦能够促进丁香假单胞等植物病原细菌的能动性与细胞膜生成。
3.通过对南美蟛蜞菊抗病基因同源序列NBS-LRR RGAs的克隆及分析,获得了2类(tir-NBS-LRR和non-tir-NBS-LRR)3条NBS-LRR RGAs序列WTRGA1(基因库登录号JX566690), WTRGA2(基因库登录号JX566691); WTRGA3(基因库登录号JX566692);进一步计算这两类NBS-LRR R-gene在入侵植物与非入侵植物中的进化速率dN/dS,结果显示,对于tir-NBS-LRR R-gene,入侵植物的进化速率显著高于非入侵植物,而对于non-tir-NBS-LRR R-gene,入侵植物的进化速率也比非入侵植物的高。意味着入侵植物具有更强的产生遗传变异的能力,也表明与非入侵植物相比,入侵植物对病原的识别能力和抗病能力可能更强。这将促使入侵植物在应对新生境当地地病原选择压力上赢得与本地植物的竞争。
4.采用微生物学传统分离培养法及16S rDNA基因序列分析的方法,对采集自我国5个省区的南美蟛蜞菊样品进行内生菌的分离,共分离鉴定了21个属156株南美蟛蜞菊的内生细菌(WTB)。16S rDNA序列分析结果表明南美蟛蜞菊的内生细菌与己报道其他植物类似,一些常见的类群,如肠杆菌属(Enterobacter)、假单胞菌(Pseudomonas)等广泛分布在各个种群。选取其中3个比较典型地潜在具有抗病性的类群(Bacillus, Pseudomonas, Variovorax)各选取1株代表菌株(JS、Psp、Vp8)进行抗病性测试。结果表明,南美蟛蜞菊的内生细菌能通过分泌一些抗菌物质显著抑制立枯丝核及香蕉枯萎病等植物病原真菌及丁香假单胞病原细菌。并且在病原压力下,接种过该内生细菌的南美蟛蜞菊茎段比未接种内生细菌的茎段生长得更好。另外,将采集自上述5个地方(海南海口、广东佛山、福建厦门、广西南宁、云南西双版纳)的南美蟛蜞菊茎段及实验室培养的无菌苗,在灭菌土壤里进行扦插,每三天施以无菌蒸馏水,同时无菌苗作为对照,3个月后收获,结果表明,扦插苗总体比无菌苗长势好,各项生理指标也更佳。
总结以上几部分研究,南美蟛蜞菊极其可能存在如下3方面非结构性抗病机制使其在野外极少染病：1)南美蟛蜞菊本身能合成具有抗真菌活性的代谢产物,2)南美蟛蜞菊NBS-LRR抗病基因具有高的进化速率,潜在的增加了其抗病基因的丰富度,3)南美蟛蜞菊具有能抑制病原菌的内生细菌。这几方面的非结构性抗病机制都能增强南美蟛蜞菊的抗病性,增强其入侵力,促进其成功入侵。另外,南美蟛蜞菊可能可以合成分泌一些代谢产物吸引类似丁香假单胞的病原细菌,南美蟛蜞菊本身不仅对此类病原细菌不感病,还可抗此类病原,并以此为武器来抑制其他本地植物,这从反面相对增强其自身同其他植物的竞争力。加上南美蟛蜞菊内生细菌的促生作用,这些都可为南美蟛蜞菊定殖新生境时抵抗当地病原、利用当地病原对抗本地植物及快速生长做出贡献,均可能在一定程度上增强其入侵性。
Currently, as the main content of the current global changes and the important international environmental ecological problem, biological invasions continue to homogenize biota, and have already became another major threat to global biodiversity. It threats both global ecological security and economic development. There are many reports about biological invasion mechanisms, however, there is few reports about systematic disease resistance of invasive species. Non-constitutive disease resistance mechanisms of invasive weed Wedelia trilobata and their contribution to W. trilobata's invasion were first studied from four different aspects in this thesis: community, secondary metabolites, R-gene, and endophytic bacteria. The details are as following:
1. By traditional community survey, this study showed that few W. trilobata suffer from disease in the field and its invasion significantly decreased its invaded community's flora species diversity. And W. trilobata's invasion showed an positive correlation withlocal economic output,
2. Exploration disease-resistance mechanism of W. trilobata from biochemical level. The secondary metabolites from different issues of W. trilobata can suppress the growth of plant pathogen fungus Fusarium oxysporum f.spcuben (Fo) and Rhizoctonia solani(Rs), whereas, it can promote the motility and biofilm formation of plant pathogen bacteria Pseudomonas syringae pv. tomato DC3000(Pst).
3. Further research also being conducted to detect the resistance gene analogs (RGAs) which involved two NBS-LRR of RGAs:tir-NBS-LRR (WTRGA1(Genbank Accession No. JX566690) and WTRGA2(Genbank Accession No. JX566691)) and non-tir-NBS-LRR (WTRGA3(Genbank Accession No. JX566692)). Then these two classes'dN/dS of NBS-LRR R-gene/RGAs were calculated. It was found that average dN/dS of invasive plants is significantly higher than that of non-invasive plants for tir-NBS-LRR R-gene/RGAs. While the invasive plants also showed higher dN/dS than that of non-invasion species for non-tir-NBS-LRR R-gene/RGAs.. The higher evolutionary rate of the RGAs of invasive plants compared to that of non-invasion species indicated the stronger ability to produce genetic variation and to identify and resistance to pathogens. And all these ability of the invasive plants will lead to that they win the competition with native plants under the local pathogens.
4.156endophytic bacterias, which were isolated from W. trilobata samples, were collected from five provinces by traditional microbiological separation culture methods and16Sr DNA gene sequence analysis. These156strains belonged to21genu. Similar to previous reports, common endophyte bacterias were indentified and existed in each population of W. trilobata, such as Enterobacter. and Pseudomonas. In addition, one strain from three typical potential resistance group(Bacillus, Pseudomonas, and Variovorax), which were termed as JS、Psp、Vp8accordingly, were selected to test resistance to pathogens. The results showed that W. trilobata endophytic bacteria can secrete some antibacterial substances to suppress the plant pathogen fungus Rs, Fo and plant pathogen bacteria Pst. And under pathogen pressure, the growth of the stem segments of W. trilobata inoculated with endophytic bacteria was better than that of without endophytic bacteria inoculation. Meanwhile, stem segments of W. trilobata collected from those above five provinces were cultivated in the sterile soil, watering with sterile water every three days and harvested after three months. Sterile seedlings of W. trilobata were conducted as control treatment. The results indicated that, the growth of stem segments was better than that of sterile seedlings. Each physiological indicator of stem segments was better than that of sterile seedlings.
The above researches indicated that W. trilobata might have three non-structural disease-resistance mechanisms, which then strengthened its disease-resistance ability:1) W. trilobata could synthesize metabolites with disease-resistance activity;2) the NBS-LRR R-gene of W. trilobata has a higher evolutionary rate, which potentially increased the richness of its R-gene;3) W. trilobata possesses endophytic bacterias which can depress the growth of indigenous bacteria. Additionally, W. trilobata may be able to attract pathogen bacteria (e.g. Pst) by secreting some metabolites, whereas W. trilobata could not be susceptible to these pathogen bacteria, it even could resistence to these pathogens, which could be used as weapons to depress local plants then to increase its competition with these local plants. And endophytic bacterias of W. trilobata possess promoting effects. All the above mechanisms, which could prevent the infection from local pathogens and use local pathogens to infect local plants and promote the rapid growth of W. trilobata, contributed great to strengthen W. trilobata's invasiveness.
1.通过传统的群落学调查方法,确定了入侵植物南美蟛蜞菊在野外很少染病的情况,南美蟛蜞菊的入侵会显著降低其入侵植物群落物种多样性。并且,经济活动加剧了南美蟛蜞菊的入侵。
2.从生化水平探讨南美蟛蜞菊抗病的机制,确定了南美蟛蜞菊不同组织的次生代谢产物既能够抑制立枯丝核及香蕉枯萎病等植物病原真菌,亦能够促进丁香假单胞等植物病原细菌的能动性与细胞膜生成。
3.通过对南美蟛蜞菊抗病基因同源序列NBS-LRR RGAs的克隆及分析,获得了2类(tir-NBS-LRR和non-tir-NBS-LRR)3条NBS-LRR RGAs序列WTRGA1(基因库登录号JX566690), WTRGA2(基因库登录号JX566691); WTRGA3(基因库登录号JX566692);进一步计算这两类NBS-LRR R-gene在入侵植物与非入侵植物中的进化速率dN/dS,结果显示,对于tir-NBS-LRR R-gene,入侵植物的进化速率显著高于非入侵植物,而对于non-tir-NBS-LRR R-gene,入侵植物的进化速率也比非入侵植物的高。意味着入侵植物具有更强的产生遗传变异的能力,也表明与非入侵植物相比,入侵植物对病原的识别能力和抗病能力可能更强。这将促使入侵植物在应对新生境当地地病原选择压力上赢得与本地植物的竞争。
4.采用微生物学传统分离培养法及16S rDNA基因序列分析的方法,对采集自我国5个省区的南美蟛蜞菊样品进行内生菌的分离,共分离鉴定了21个属156株南美蟛蜞菊的内生细菌(WTB)。16S rDNA序列分析结果表明南美蟛蜞菊的内生细菌与己报道其他植物类似,一些常见的类群,如肠杆菌属(Enterobacter)、假单胞菌(Pseudomonas)等广泛分布在各个种群。选取其中3个比较典型地潜在具有抗病性的类群(Bacillus, Pseudomonas, Variovorax)各选取1株代表菌株(JS、Psp、Vp8)进行抗病性测试。结果表明,南美蟛蜞菊的内生细菌能通过分泌一些抗菌物质显著抑制立枯丝核及香蕉枯萎病等植物病原真菌及丁香假单胞病原细菌。并且在病原压力下,接种过该内生细菌的南美蟛蜞菊茎段比未接种内生细菌的茎段生长得更好。另外,将采集自上述5个地方(海南海口、广东佛山、福建厦门、广西南宁、云南西双版纳)的南美蟛蜞菊茎段及实验室培养的无菌苗,在灭菌土壤里进行扦插,每三天施以无菌蒸馏水,同时无菌苗作为对照,3个月后收获,结果表明,扦插苗总体比无菌苗长势好,各项生理指标也更佳。
总结以上几部分研究,南美蟛蜞菊极其可能存在如下3方面非结构性抗病机制使其在野外极少染病：1)南美蟛蜞菊本身能合成具有抗真菌活性的代谢产物,2)南美蟛蜞菊NBS-LRR抗病基因具有高的进化速率,潜在的增加了其抗病基因的丰富度,3)南美蟛蜞菊具有能抑制病原菌的内生细菌。这几方面的非结构性抗病机制都能增强南美蟛蜞菊的抗病性,增强其入侵力,促进其成功入侵。另外,南美蟛蜞菊可能可以合成分泌一些代谢产物吸引类似丁香假单胞的病原细菌,南美蟛蜞菊本身不仅对此类病原细菌不感病,还可抗此类病原,并以此为武器来抑制其他本地植物,这从反面相对增强其自身同其他植物的竞争力。加上南美蟛蜞菊内生细菌的促生作用,这些都可为南美蟛蜞菊定殖新生境时抵抗当地病原、利用当地病原对抗本地植物及快速生长做出贡献,均可能在一定程度上增强其入侵性。
Currently, as the main content of the current global changes and the important international environmental ecological problem, biological invasions continue to homogenize biota, and have already became another major threat to global biodiversity. It threats both global ecological security and economic development. There are many reports about biological invasion mechanisms, however, there is few reports about systematic disease resistance of invasive species. Non-constitutive disease resistance mechanisms of invasive weed Wedelia trilobata and their contribution to W. trilobata's invasion were first studied from four different aspects in this thesis: community, secondary metabolites, R-gene, and endophytic bacteria. The details are as following:
1. By traditional community survey, this study showed that few W. trilobata suffer from disease in the field and its invasion significantly decreased its invaded community's flora species diversity. And W. trilobata's invasion showed an positive correlation withlocal economic output,
2. Exploration disease-resistance mechanism of W. trilobata from biochemical level. The secondary metabolites from different issues of W. trilobata can suppress the growth of plant pathogen fungus Fusarium oxysporum f.spcuben (Fo) and Rhizoctonia solani(Rs), whereas, it can promote the motility and biofilm formation of plant pathogen bacteria Pseudomonas syringae pv. tomato DC3000(Pst).
3. Further research also being conducted to detect the resistance gene analogs (RGAs) which involved two NBS-LRR of RGAs:tir-NBS-LRR (WTRGA1(Genbank Accession No. JX566690) and WTRGA2(Genbank Accession No. JX566691)) and non-tir-NBS-LRR (WTRGA3(Genbank Accession No. JX566692)). Then these two classes'dN/dS of NBS-LRR R-gene/RGAs were calculated. It was found that average dN/dS of invasive plants is significantly higher than that of non-invasive plants for tir-NBS-LRR R-gene/RGAs. While the invasive plants also showed higher dN/dS than that of non-invasion species for non-tir-NBS-LRR R-gene/RGAs.. The higher evolutionary rate of the RGAs of invasive plants compared to that of non-invasion species indicated the stronger ability to produce genetic variation and to identify and resistance to pathogens. And all these ability of the invasive plants will lead to that they win the competition with native plants under the local pathogens.
4.156endophytic bacterias, which were isolated from W. trilobata samples, were collected from five provinces by traditional microbiological separation culture methods and16Sr DNA gene sequence analysis. These156strains belonged to21genu. Similar to previous reports, common endophyte bacterias were indentified and existed in each population of W. trilobata, such as Enterobacter. and Pseudomonas. In addition, one strain from three typical potential resistance group(Bacillus, Pseudomonas, and Variovorax), which were termed as JS、Psp、Vp8accordingly, were selected to test resistance to pathogens. The results showed that W. trilobata endophytic bacteria can secrete some antibacterial substances to suppress the plant pathogen fungus Rs, Fo and plant pathogen bacteria Pst. And under pathogen pressure, the growth of the stem segments of W. trilobata inoculated with endophytic bacteria was better than that of without endophytic bacteria inoculation. Meanwhile, stem segments of W. trilobata collected from those above five provinces were cultivated in the sterile soil, watering with sterile water every three days and harvested after three months. Sterile seedlings of W. trilobata were conducted as control treatment. The results indicated that, the growth of stem segments was better than that of sterile seedlings. Each physiological indicator of stem segments was better than that of sterile seedlings.
The above researches indicated that W. trilobata might have three non-structural disease-resistance mechanisms, which then strengthened its disease-resistance ability:1) W. trilobata could synthesize metabolites with disease-resistance activity;2) the NBS-LRR R-gene of W. trilobata has a higher evolutionary rate, which potentially increased the richness of its R-gene;3) W. trilobata possesses endophytic bacterias which can depress the growth of indigenous bacteria. Additionally, W. trilobata may be able to attract pathogen bacteria (e.g. Pst) by secreting some metabolites, whereas W. trilobata could not be susceptible to these pathogen bacteria, it even could resistence to these pathogens, which could be used as weapons to depress local plants then to increase its competition with these local plants. And endophytic bacterias of W. trilobata possess promoting effects. All the above mechanisms, which could prevent the infection from local pathogens and use local pathogens to infect local plants and promote the rapid growth of W. trilobata, contributed great to strengthen W. trilobata's invasiveness.
引文
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