松材线虫纤维素酶系及其cDNA基因的克隆与RNA干扰研究
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摘要
随着全球一体化和国际社会交往与贸易往来的与日俱增,外来有害生物随人口流动、动植物及其产品跨国扩散日益严重,给入侵地生态系统和国土安全造成巨大的灾害性损失。松材线虫(Bursaphelenchu xylophilus (Steiner & Buhrer) Nickle)已经成为世界上重要的检疫对象和我国有史以来危害最为严重的林业外来有害生物。基于经典与现代生物学的理论与方法,国内外众多学者对松材线虫的生物学、生态学和致病性等进行了深入研究,但是迄今尚未有有效控制措施。本研究基于前人对松材线虫致病过程中纤维素酶的认识,研究分析松材线虫和近似种拟松材线虫(B. mucronatus)分泌的纤维素酶系组成,纤维素酶系活力与线虫迁移扩散能力的关系,以及纤维素酶相关编码基因的克隆与功能鉴定,意图阐明纤维素酶在松材线虫致病过程中的分子机制,为松材线虫病的可持续控制提供理论基础。
采用真菌平板培养法研究了不同松材线虫和拟松材线虫株系的繁殖能力。在灰葡萄孢上培养10d,松材线虫和拟松材线虫繁殖倍数分别达到248倍和373倍,而在松球壳孢上培养10d,松材线虫和拟松材线虫繁殖倍数分别为133和98倍。结合性比及幼虫/成虫指标数据,结果表明,在本实验室条件下,线虫在灰葡萄孢(Botrytis cinerea)上繁殖速度最快,繁殖后得到合适数量的混合龄期线虫;以反映线虫繁殖能力的三个指标:繁殖倍数、性比和幼虫/成虫分析比较了供试的6个松材线虫株系和4个拟松材线虫株系,以松材线虫Bx02、Bx03和拟松材线虫Bm01表现较高的繁殖能力。
分别采用羧甲基纤维素钠(CMC)、微晶纤维素(MC)和水杨素(SC)三种底物,结合DNS定量分析法,研究了松材线虫和拟松材线虫体外分泌物中内切-β-1,4-葡聚糖酶、外切-β-1,4-葡聚糖酶和β-葡萄糖苷酶三类酶的活性。结果表明,在离体培养条件下,松材线虫和拟松材线虫均能产生包括三类酶的完整纤维素酶系。其中,松材线虫分泌物中三类酶活性明显高于拟松材线虫分泌物中这三类酶活性。在内切-β-1,4-葡聚糖酶中,松材线虫Bx01株系的酶活性最高为9.36μg·ml-1·min-1,拟松材线虫Bm03株系的酶活性最高是6.32μg·ml-1·min-1,前者酶活性是后者的1.48倍;外切-β-1,4-葡聚糖酶酶活性最高的松材线虫Bx01是6.67μg·ml-1·min-1,酶活性最高的拟松材线虫Bm03是4.22μg·ml-1·min-1,前者是后者的1.58倍;Β-葡萄糖苷酶活性最高的松材线虫Bx02株系为7.35μg·ml-1·min-1,最高的拟松材线虫Bm01株系为4.63μg·ml-1·min-1,前者是后者酶活性的1.59倍。方差分析和多重比较结果表明,松材线虫Bx01、Bx02、Bx04、Bx05和Bx06株系内切-β-1,4-葡聚糖酶活性同所有拟松材线虫株系相比,结果达到极显著差异。结合二者致病力的显著差异,研究结果为更好的解释松材线虫致病机理提供了科学数据。
通过人工接种方法,比较了供试松材线虫和拟松材线虫株系在黑松枝段内的迁移扩散和繁殖能力。结果表明,与拟松材线虫相比,松材线虫在黑松枝段内迁移能力更强,表现在以更快的速度和更大的种群数量穿过黑松枝段。多数松材线虫株系在黑松枝段内的繁殖力高于拟松材线虫,并且多数松材线虫株系繁殖数量同其中三个拟松材线虫株系繁殖数量差异显著。结合松材线虫株系分泌的纤维素酶系活性明显高于拟松材线虫株系的纤维素酶系,纤维素酶可能是线虫在寄主体内迁移扩散的主要驱动力之一。
以松材线虫总RNA为模板,运用RT-PCR技术分离和克隆了编码松材线虫纤维素酶系组分之一的内切-β-1,4-葡聚糖酶基因的cDNA片段(GenBank注册号为EU660207)。经T载体克隆、酶切鉴定和序列测定获得cDNA片段为一个长678bp的完整的开放阅读框,编码全长为225个氨基酸的蛋白,同GenBank数据库中已登录的日本松材线虫内切-β-1,4-葡聚糖酶基因表现出较高的同源性,相似性为95%。用生物信息学方法分析和预测了编码蛋白质的主要特性。同源性分析表明,该蛋白同真菌中糖基水解酶相似性比较高,同植物寄生线虫和细菌的糖基水解酶相似性较低,证明该基因是来自真菌的水平基因漂移(HGT)。
进一步运用RANi技术鉴定了松材线虫内切-β-1,4-葡聚糖酶基因的功能。荧光显微镜观察结果表明,溶液浸泡可以把siRNA成功导入松材线虫体内,产生基因沉默效应。采用荧光定量PCR技术检测松材线虫内切-β-1,4-葡聚糖酶基因mRNA水平的表达抑制效应发现,siRNA15和siRNA275的沉默效率最高,siRNA581沉默基因的效率最低。生物学效应分析结果表明,siRNA松材线虫处理后,松材线虫分泌物中纤维素酶活性显著下降,说明松材线虫β-1,4-葡聚糖酶基因的表达受到明显抑制。同时,松材线虫种群繁殖力明显降低,穿过黑松枝段的时间明显延长,说明松材线虫内切-β-1,4-葡聚糖酶可能对松材线虫生长发育和运动扩散能力也有一定的影响。
With the development of the global integration and international exchanging and trading, the invasive alien species dispersal causes a large damage to the ecology system of the country invaded. Pine wood nematode (Bursaphelenchus xylophilus), as an agent of pine wilt disease and the important quarantine target of many countries around the world, has been the most serious invasive pest in China. Based on the classic and modern biology theory and method, many scientists focused on the biology, ecology and pathogenicity and so on from all over the world, but until recently there were no effective controlling measures to the pine wilt disease. Based on the understanding of cellulase from pine wood nematode secretion, the component of cellulase from B. xylophilus and B. mucronatus was analyzed while the relationship between cellulase activity and dispersal ability of the nematode was also discussed. And cellulase encoding gene was isolated and identified with RNAi. This will try to demonstrate the molecular mechanism of cellulase in the pathogenicity, and give science data to provide an academic support for the sustainable management of pine wilt disease.
The population propagation ability of B. xylophilus and B. mucronatus was analyzed. The results showed that both B. xylophilus and B. mucronatus reproduced very fast on the Botrytis cinerea culture which produced large number of mixed stage nematodes. Three index including propagation rate, sex ratio and larvae/adult were used to comparing the propagation ability. B. xylophilus strain Bx02, Bx03 and B. mucronatus Bm01 exhibitted the potential high propagating ability.
The secretion cellulase of B. xylophilus and B. mucronatus was analyzed while CMC, MC and SC were used to be the celllose resouces for the nematodes secretion degrading. The results showed that in vitro endo-β-1,4-glucanase, exo-β-1,4-glucanase andβ-glycosidase activity could be detected in the secretion of both B. xylophilus and B. mucronatus. And the three cellulase activity from B. xylophilus secretion was higher than that from B. mucronatus. It is inferred that the pathogenicity of B. xylophilus was more serious than B. mucronatus for the reason of the cellulase activity.
The dispersal and propagation ability of B. xylophilus and B. mucronatus in Pinus thunbergii shoot section were analyzed with artificial inoculation. Compared with B. mucronatus, B. xylophilus passed through the shoot section of P. thunbergii with a higher speed and less time. The numbers of nematode passing through the section at first were not significantly different among the most strains of nematodes including B. mucronatus, although the numbers of most strains of B. xylophilus were much more than that of B. mucronatus. The reproduction numbers of B. xylophilus were more than that of B. mucronatus in the section for 20 days and all the numbers of B. xylophilus strains were significantly different with the numbers of three strains of B. mucronatus. Considering the same phenomena in the investigation of the cellulase activity of both species nematodes above, the cellulase probably was one of the driving for the nematode dispersal inside the host tree.
The cDNA fragment coding endo-β-1,4-glucanase of B. xylophilus was isolated and identified from total RNA with RT-PCR techniques. The cDNA fragment was 678 bp in length and contained an open reading frame of 225 amino acids (GenBank accession number EU660207). The cDNA could eccode a protein similar to glycosyl hydrolase family 45. The Blast result showed that the cDNA shared 95% identity with endo-β-1,4-glucanase from Japanese strain of B. xylophilus. The characteristic of the putative protein were analyzed with biological information method. The alignment of several selected sequences similar to this cDNA in GenBank suggested that the cellulase gene from Chinese strain Bx02 of B. xylophilus was acquired by horizontal gene transfer from fungi.
The function of gene encoding endo-β-1,4-glucanase was identified with RNAi. The siRNA was introduced into the body of pine wood nematodes with the fluorescence microscope investigation. The effect of inhibition on the expression of gene encoding endo-β-1,4-glucanase was detected with fluorescence quantity real time PCR. siRNA15 and siRNA275 showed more higher effective inhibition than other siRNA. The biology characteristic of pine wood nematodes after RNAi was demonstrated and analyzed. The cellulase activity decreased to a low level and the reproduction rate was also reduced. Furthermore, it took some more time to pass through the shoot section of P. thunbergii for pine wood nematodes after RNAi. This indicated that some functions related to the development or dispersal ability of pine wood nematodes were likely to be influenced while siRNAs interferenced the gene encoding endo-β-1,4-glucanase.
采用真菌平板培养法研究了不同松材线虫和拟松材线虫株系的繁殖能力。在灰葡萄孢上培养10d,松材线虫和拟松材线虫繁殖倍数分别达到248倍和373倍,而在松球壳孢上培养10d,松材线虫和拟松材线虫繁殖倍数分别为133和98倍。结合性比及幼虫/成虫指标数据,结果表明,在本实验室条件下,线虫在灰葡萄孢(Botrytis cinerea)上繁殖速度最快,繁殖后得到合适数量的混合龄期线虫;以反映线虫繁殖能力的三个指标:繁殖倍数、性比和幼虫/成虫分析比较了供试的6个松材线虫株系和4个拟松材线虫株系,以松材线虫Bx02、Bx03和拟松材线虫Bm01表现较高的繁殖能力。
分别采用羧甲基纤维素钠(CMC)、微晶纤维素(MC)和水杨素(SC)三种底物,结合DNS定量分析法,研究了松材线虫和拟松材线虫体外分泌物中内切-β-1,4-葡聚糖酶、外切-β-1,4-葡聚糖酶和β-葡萄糖苷酶三类酶的活性。结果表明,在离体培养条件下,松材线虫和拟松材线虫均能产生包括三类酶的完整纤维素酶系。其中,松材线虫分泌物中三类酶活性明显高于拟松材线虫分泌物中这三类酶活性。在内切-β-1,4-葡聚糖酶中,松材线虫Bx01株系的酶活性最高为9.36μg·ml-1·min-1,拟松材线虫Bm03株系的酶活性最高是6.32μg·ml-1·min-1,前者酶活性是后者的1.48倍;外切-β-1,4-葡聚糖酶酶活性最高的松材线虫Bx01是6.67μg·ml-1·min-1,酶活性最高的拟松材线虫Bm03是4.22μg·ml-1·min-1,前者是后者的1.58倍;Β-葡萄糖苷酶活性最高的松材线虫Bx02株系为7.35μg·ml-1·min-1,最高的拟松材线虫Bm01株系为4.63μg·ml-1·min-1,前者是后者酶活性的1.59倍。方差分析和多重比较结果表明,松材线虫Bx01、Bx02、Bx04、Bx05和Bx06株系内切-β-1,4-葡聚糖酶活性同所有拟松材线虫株系相比,结果达到极显著差异。结合二者致病力的显著差异,研究结果为更好的解释松材线虫致病机理提供了科学数据。
通过人工接种方法,比较了供试松材线虫和拟松材线虫株系在黑松枝段内的迁移扩散和繁殖能力。结果表明,与拟松材线虫相比,松材线虫在黑松枝段内迁移能力更强,表现在以更快的速度和更大的种群数量穿过黑松枝段。多数松材线虫株系在黑松枝段内的繁殖力高于拟松材线虫,并且多数松材线虫株系繁殖数量同其中三个拟松材线虫株系繁殖数量差异显著。结合松材线虫株系分泌的纤维素酶系活性明显高于拟松材线虫株系的纤维素酶系,纤维素酶可能是线虫在寄主体内迁移扩散的主要驱动力之一。
以松材线虫总RNA为模板,运用RT-PCR技术分离和克隆了编码松材线虫纤维素酶系组分之一的内切-β-1,4-葡聚糖酶基因的cDNA片段(GenBank注册号为EU660207)。经T载体克隆、酶切鉴定和序列测定获得cDNA片段为一个长678bp的完整的开放阅读框,编码全长为225个氨基酸的蛋白,同GenBank数据库中已登录的日本松材线虫内切-β-1,4-葡聚糖酶基因表现出较高的同源性,相似性为95%。用生物信息学方法分析和预测了编码蛋白质的主要特性。同源性分析表明,该蛋白同真菌中糖基水解酶相似性比较高,同植物寄生线虫和细菌的糖基水解酶相似性较低,证明该基因是来自真菌的水平基因漂移(HGT)。
进一步运用RANi技术鉴定了松材线虫内切-β-1,4-葡聚糖酶基因的功能。荧光显微镜观察结果表明,溶液浸泡可以把siRNA成功导入松材线虫体内,产生基因沉默效应。采用荧光定量PCR技术检测松材线虫内切-β-1,4-葡聚糖酶基因mRNA水平的表达抑制效应发现,siRNA15和siRNA275的沉默效率最高,siRNA581沉默基因的效率最低。生物学效应分析结果表明,siRNA松材线虫处理后,松材线虫分泌物中纤维素酶活性显著下降,说明松材线虫β-1,4-葡聚糖酶基因的表达受到明显抑制。同时,松材线虫种群繁殖力明显降低,穿过黑松枝段的时间明显延长,说明松材线虫内切-β-1,4-葡聚糖酶可能对松材线虫生长发育和运动扩散能力也有一定的影响。
With the development of the global integration and international exchanging and trading, the invasive alien species dispersal causes a large damage to the ecology system of the country invaded. Pine wood nematode (Bursaphelenchus xylophilus), as an agent of pine wilt disease and the important quarantine target of many countries around the world, has been the most serious invasive pest in China. Based on the classic and modern biology theory and method, many scientists focused on the biology, ecology and pathogenicity and so on from all over the world, but until recently there were no effective controlling measures to the pine wilt disease. Based on the understanding of cellulase from pine wood nematode secretion, the component of cellulase from B. xylophilus and B. mucronatus was analyzed while the relationship between cellulase activity and dispersal ability of the nematode was also discussed. And cellulase encoding gene was isolated and identified with RNAi. This will try to demonstrate the molecular mechanism of cellulase in the pathogenicity, and give science data to provide an academic support for the sustainable management of pine wilt disease.
The population propagation ability of B. xylophilus and B. mucronatus was analyzed. The results showed that both B. xylophilus and B. mucronatus reproduced very fast on the Botrytis cinerea culture which produced large number of mixed stage nematodes. Three index including propagation rate, sex ratio and larvae/adult were used to comparing the propagation ability. B. xylophilus strain Bx02, Bx03 and B. mucronatus Bm01 exhibitted the potential high propagating ability.
The secretion cellulase of B. xylophilus and B. mucronatus was analyzed while CMC, MC and SC were used to be the celllose resouces for the nematodes secretion degrading. The results showed that in vitro endo-β-1,4-glucanase, exo-β-1,4-glucanase andβ-glycosidase activity could be detected in the secretion of both B. xylophilus and B. mucronatus. And the three cellulase activity from B. xylophilus secretion was higher than that from B. mucronatus. It is inferred that the pathogenicity of B. xylophilus was more serious than B. mucronatus for the reason of the cellulase activity.
The dispersal and propagation ability of B. xylophilus and B. mucronatus in Pinus thunbergii shoot section were analyzed with artificial inoculation. Compared with B. mucronatus, B. xylophilus passed through the shoot section of P. thunbergii with a higher speed and less time. The numbers of nematode passing through the section at first were not significantly different among the most strains of nematodes including B. mucronatus, although the numbers of most strains of B. xylophilus were much more than that of B. mucronatus. The reproduction numbers of B. xylophilus were more than that of B. mucronatus in the section for 20 days and all the numbers of B. xylophilus strains were significantly different with the numbers of three strains of B. mucronatus. Considering the same phenomena in the investigation of the cellulase activity of both species nematodes above, the cellulase probably was one of the driving for the nematode dispersal inside the host tree.
The cDNA fragment coding endo-β-1,4-glucanase of B. xylophilus was isolated and identified from total RNA with RT-PCR techniques. The cDNA fragment was 678 bp in length and contained an open reading frame of 225 amino acids (GenBank accession number EU660207). The cDNA could eccode a protein similar to glycosyl hydrolase family 45. The Blast result showed that the cDNA shared 95% identity with endo-β-1,4-glucanase from Japanese strain of B. xylophilus. The characteristic of the putative protein were analyzed with biological information method. The alignment of several selected sequences similar to this cDNA in GenBank suggested that the cellulase gene from Chinese strain Bx02 of B. xylophilus was acquired by horizontal gene transfer from fungi.
The function of gene encoding endo-β-1,4-glucanase was identified with RNAi. The siRNA was introduced into the body of pine wood nematodes with the fluorescence microscope investigation. The effect of inhibition on the expression of gene encoding endo-β-1,4-glucanase was detected with fluorescence quantity real time PCR. siRNA15 and siRNA275 showed more higher effective inhibition than other siRNA. The biology characteristic of pine wood nematodes after RNAi was demonstrated and analyzed. The cellulase activity decreased to a low level and the reproduction rate was also reduced. Furthermore, it took some more time to pass through the shoot section of P. thunbergii for pine wood nematodes after RNAi. This indicated that some functions related to the development or dispersal ability of pine wood nematodes were likely to be influenced while siRNAs interferenced the gene encoding endo-β-1,4-glucanase.
引文
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