湖北省灰霉病病菌区系和灰葡萄孢菌多样性研究
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摘要
灰霉病(grey mould)是葡萄孢属真菌(Botrytis spp.)引起的一类世界范围分布的重要的植物病害。对该属病原菌的分类鉴定工作具有非常重要的理论研究和实际应用价值。许多真菌分类学家多年来一直致力于不断提高葡萄孢属真菌的分类水平,目前已经有28种葡萄孢菌的纪录。2008年已完成了该属模式菌-灰葡萄孢菌(B. cinerea)的全基因组测序。同时,Staats et al.(2005)分子水平上证明葡萄孢属真菌与寄主的进化的关系。本论文利用形态学和分子系统学分析方法对湖北省重要经济作物上的葡萄孢属真菌种类进行了分离和鉴定。对一些重要种类进行了分布调查、生物学特性、致病性和遗传多样性分析。取得的研究结果如下：
1、对采自湖北省不同寄主作物样品进行了葡萄孢属真菌的分离与培养,共获得了818株葡萄孢属真菌和类似葡萄孢属真菌分离物。共鉴定出14个种,包括两个新种：拟蚕豆葡萄孢(B. fabiopsis)和中国葱葡萄孢(B. sinoallii),分别引起蚕豆赤斑病、葱属蔬菜叶斑病。命名了一种新病害：大蒜盲种葡萄孢菌(B. porri)引起大蒜叶斑病。确定了一种病原的分类属性,即引起洋葱茎腐病的病原为B. aclada,而非B. alli。分离两个疑似新种：葡萄孢种1(Botrytis sp.l)和葡萄孢种2(Botrytis sp.2)。鉴定出3种葡萄孢属类似真菌,它们是：Amphyobotrys ricini、Streptobotrys caulophylli和Verrucobotrys geranii。对拟蚕豆葡萄孢和中国葱葡萄孢菌的致病力进行了测定。结果表明：拟蚕豆葡萄孢菌引起典型的蚕豆赤斑病,其致病力与蚕豆葡萄孢菌相似。但拟蚕豆葡萄孢菌的在湖北省的自然分布较蚕豆葡萄孢菌广泛。可见,拟蚕豆葡萄孢菌对蚕豆的危害较蚕豆葡萄孢菌严重。中国葱葡萄孢菌在接种的条件下,能够感染小葱、大蒜和韭菜。在发病部位产生大量分生孢子,其潜在重要性不能忽视。
2、利用ITS和G3PDH基因序列,对葡萄孢属及其近缘属真菌(Amphyobotrys, Streptobotrys, Verrucobotrys, Sclerotinia, Monilinia)的系统发育关系进行了分析。与ITS相比,G3PDH基因序列可以很好的解决葡萄孢属真菌及其近缘属真菌的系统发育问题。Verrucobotrys与Botrytis的亲缘关系最近,虽呈独立分支,但Verrucobotrys属真菌是属于葡萄孢属真菌一大类群里的亚支,将其作为一个独立的属还有待研究。
3、研究确定了两个新种拟蚕豆葡萄孢(B. fabiopsis)和中国葱葡萄孢(B.sinoallii)的生物学特性。B. fabiopsis菌丝生长的温度范围为5～30℃,最适生长温度为20-25℃。菌丝pH值范围为2～10,pH值为4.0-5.0时利于菌丝生长和菌核形成,最适宜的碳氮源分别为可溶性淀粉,丙氨酸。B.fabiopsis、B.fabae和B. cinerea之间对温度、pH值和碳源／氮源的反应存在明显差异。B.fabiopsis和B. fabae比较而言具有更广泛的生长温度范围和pH值范围。B. sinoallii菌丝生长的温度范围为5-30℃,最适生长温度为20℃。菌丝pH值范围为2-10,最适pH值为5.0,pH值为6.0时有利于菌核形成,最适宜的碳氮源分别为葡萄糖,蛋白胨和丙氨酸。B. sinoallii和B. squamosa之间对温度、pH值和碳源／氮源的反应存在一定的差异,相对而言,B. sinoallii较B. squamosa更适应偏高的生长温度(25℃和28℃)。
4、探讨了灰葡萄孢菌的多样性规律。灰葡萄孢菌菌株在培养特征表现为菌丝型、孢子型和菌核型等3类,以菌核型为主。在转座子方面,可分为transposa和vacuma两类,以transposa为主。抗药性分析结果表明：大部分菌株对多菌灵、菌核净、农利灵和环酰菌胺为敏感菌株,并从分子水平分析了部分抗性菌株抗药的可能机制。不同来源的灰葡萄孢菌株都有对油菜的致病能力,但是菌株之间存在差异。即使是相同田块来源的菌株在特养特征、致病性、转座子和抗药性上同样也存在丰富的多样性。
Botrytis spp., the causal agents of grey mould diseases, can infect many fruits, vegetables and ornamental crops. These fungi are world-wide plant pathogens. It is necessary to keep continuous research on Botrytis taxonomy and identification. The fungal taxonomists all over the world have been making efferts to promote Botrytis taxonomy. Nowadays, there are 28 species in this genus. The genome sequencing of B. cinerea has been completed in 2008. The relationship between Botrytis and host plants has been reported and many new species and diversity of Botrytis have been elucidated.
Some researches on morphology, histology, molecular biology and proteomics of Botrytis have made considerable progress. However, it has been short of taxonomic work on Botrytis in China. The morphological and molecular character of Botrytis were identified in this paper. The genetic diversity of Botrytis cinerea isolated from different host in Hubei province was studied based on mophyological characteristics, pathogenicity, resistance and molecular biology techniques. The aim of this study is to determine Botrytis species and diversity of B. cinerea. The results are summarized below:
1. The Botrytis cultures were isolated from different hosts and 818 isolates were identied based on morphological and molecular characteristics, including 11 species of Botrytis and 3 Botrytis-like genera. Two new species of Botrytis were described, B. sinoallii causing leaf blight of Allium and B. fabiopsis causing chocolate spot of broad bean. A new disease of garlic caused by B. porri was identified. The taxonomic status of the causal agent of onion bulb rot was determined to be B. aclada, rather than B. allii. Two suspected new species, Botrytis spl. and Botrytis sp2 were identified. Three Botrytis-like species were Amphyobotrys ricini, Streptobotrys caulophylli and Verrucobotrys geranii isolated and identified. In addition, the distribution and the pathogenicity of some species of Botrytis spp. were investigated.
2. Phylogenetic relationships were analyzed using the sequence of ITS and G3PDH gene of Botrytis and related genera(Amphyobotrys, Streptobotrys, Verrucobotrys, Sclerotinia, Monilinia). The partial sequence of G3PDH gene can be used for solving the phylogeny problem, compared to ITS. Verrucobotrys is closely related to Botrytis. Verrucobotrys was clustered in an independent branch of a clade of Botrytis in the phylogenetic tree. So, it needs to further investigation that Verrucobotrys was a separate genus needs.
3. Two new species, B. fabiopsis and B. sinoallii, were characterized for basic biological properties (the optimum temperature, pH and the optimum carbon and nitrogen sources). The result showed B. fabiopsis could grow in 5-30℃, pH values of 2.0-10.0. The most optimum temperature is 20-25℃for mycelium growth and the most optimum pH value is 4.0-5.0. The most suitable carbon and nitrogen sources were soluble starch and alanine, respectively. B. fabiopsis, B. fabae and B. cinerea showed a dramatic difference in response to temperature, pH value and nutritent sources. B. sinoallii could grow in 5-30℃, pH values of 2.0-10.0 with the optimum temperature of 20℃and ph value of 4.0-5.0. The most suitable carbon and nitrogen sources were glucose and peptone/alanine, respectively. B. sinoallii and B. squamosa also showed a dramatic difference in response to temperature, pH value and nutrient sources. B. sinoallii showed better growth at 25℃and 28℃than B. squamosa.
4. The genetic diversity of Botrytis cinerea isolated from different hosts in Hubei Province was studied based on mophyological characteristics, pathogenicity, fungicide resistance and some molecular features. The results showed that:isolates of B. cinerea have three types:mycelium type, conidial type and sclerotial type. A PCR method was developed to identify the transposable elements Boty and Flipper that form two groups, transposa and vacuma. Most of isolates are sensitive to four fungicides.
1、对采自湖北省不同寄主作物样品进行了葡萄孢属真菌的分离与培养,共获得了818株葡萄孢属真菌和类似葡萄孢属真菌分离物。共鉴定出14个种,包括两个新种：拟蚕豆葡萄孢(B. fabiopsis)和中国葱葡萄孢(B. sinoallii),分别引起蚕豆赤斑病、葱属蔬菜叶斑病。命名了一种新病害：大蒜盲种葡萄孢菌(B. porri)引起大蒜叶斑病。确定了一种病原的分类属性,即引起洋葱茎腐病的病原为B. aclada,而非B. alli。分离两个疑似新种：葡萄孢种1(Botrytis sp.l)和葡萄孢种2(Botrytis sp.2)。鉴定出3种葡萄孢属类似真菌,它们是：Amphyobotrys ricini、Streptobotrys caulophylli和Verrucobotrys geranii。对拟蚕豆葡萄孢和中国葱葡萄孢菌的致病力进行了测定。结果表明：拟蚕豆葡萄孢菌引起典型的蚕豆赤斑病,其致病力与蚕豆葡萄孢菌相似。但拟蚕豆葡萄孢菌的在湖北省的自然分布较蚕豆葡萄孢菌广泛。可见,拟蚕豆葡萄孢菌对蚕豆的危害较蚕豆葡萄孢菌严重。中国葱葡萄孢菌在接种的条件下,能够感染小葱、大蒜和韭菜。在发病部位产生大量分生孢子,其潜在重要性不能忽视。
2、利用ITS和G3PDH基因序列,对葡萄孢属及其近缘属真菌(Amphyobotrys, Streptobotrys, Verrucobotrys, Sclerotinia, Monilinia)的系统发育关系进行了分析。与ITS相比,G3PDH基因序列可以很好的解决葡萄孢属真菌及其近缘属真菌的系统发育问题。Verrucobotrys与Botrytis的亲缘关系最近,虽呈独立分支,但Verrucobotrys属真菌是属于葡萄孢属真菌一大类群里的亚支,将其作为一个独立的属还有待研究。
3、研究确定了两个新种拟蚕豆葡萄孢(B. fabiopsis)和中国葱葡萄孢(B.sinoallii)的生物学特性。B. fabiopsis菌丝生长的温度范围为5～30℃,最适生长温度为20-25℃。菌丝pH值范围为2～10,pH值为4.0-5.0时利于菌丝生长和菌核形成,最适宜的碳氮源分别为可溶性淀粉,丙氨酸。B.fabiopsis、B.fabae和B. cinerea之间对温度、pH值和碳源／氮源的反应存在明显差异。B.fabiopsis和B. fabae比较而言具有更广泛的生长温度范围和pH值范围。B. sinoallii菌丝生长的温度范围为5-30℃,最适生长温度为20℃。菌丝pH值范围为2-10,最适pH值为5.0,pH值为6.0时有利于菌核形成,最适宜的碳氮源分别为葡萄糖,蛋白胨和丙氨酸。B. sinoallii和B. squamosa之间对温度、pH值和碳源／氮源的反应存在一定的差异,相对而言,B. sinoallii较B. squamosa更适应偏高的生长温度(25℃和28℃)。
4、探讨了灰葡萄孢菌的多样性规律。灰葡萄孢菌菌株在培养特征表现为菌丝型、孢子型和菌核型等3类,以菌核型为主。在转座子方面,可分为transposa和vacuma两类,以transposa为主。抗药性分析结果表明：大部分菌株对多菌灵、菌核净、农利灵和环酰菌胺为敏感菌株,并从分子水平分析了部分抗性菌株抗药的可能机制。不同来源的灰葡萄孢菌株都有对油菜的致病能力,但是菌株之间存在差异。即使是相同田块来源的菌株在特养特征、致病性、转座子和抗药性上同样也存在丰富的多样性。
Botrytis spp., the causal agents of grey mould diseases, can infect many fruits, vegetables and ornamental crops. These fungi are world-wide plant pathogens. It is necessary to keep continuous research on Botrytis taxonomy and identification. The fungal taxonomists all over the world have been making efferts to promote Botrytis taxonomy. Nowadays, there are 28 species in this genus. The genome sequencing of B. cinerea has been completed in 2008. The relationship between Botrytis and host plants has been reported and many new species and diversity of Botrytis have been elucidated.
Some researches on morphology, histology, molecular biology and proteomics of Botrytis have made considerable progress. However, it has been short of taxonomic work on Botrytis in China. The morphological and molecular character of Botrytis were identified in this paper. The genetic diversity of Botrytis cinerea isolated from different host in Hubei province was studied based on mophyological characteristics, pathogenicity, resistance and molecular biology techniques. The aim of this study is to determine Botrytis species and diversity of B. cinerea. The results are summarized below:
1. The Botrytis cultures were isolated from different hosts and 818 isolates were identied based on morphological and molecular characteristics, including 11 species of Botrytis and 3 Botrytis-like genera. Two new species of Botrytis were described, B. sinoallii causing leaf blight of Allium and B. fabiopsis causing chocolate spot of broad bean. A new disease of garlic caused by B. porri was identified. The taxonomic status of the causal agent of onion bulb rot was determined to be B. aclada, rather than B. allii. Two suspected new species, Botrytis spl. and Botrytis sp2 were identified. Three Botrytis-like species were Amphyobotrys ricini, Streptobotrys caulophylli and Verrucobotrys geranii isolated and identified. In addition, the distribution and the pathogenicity of some species of Botrytis spp. were investigated.
2. Phylogenetic relationships were analyzed using the sequence of ITS and G3PDH gene of Botrytis and related genera(Amphyobotrys, Streptobotrys, Verrucobotrys, Sclerotinia, Monilinia). The partial sequence of G3PDH gene can be used for solving the phylogeny problem, compared to ITS. Verrucobotrys is closely related to Botrytis. Verrucobotrys was clustered in an independent branch of a clade of Botrytis in the phylogenetic tree. So, it needs to further investigation that Verrucobotrys was a separate genus needs.
3. Two new species, B. fabiopsis and B. sinoallii, were characterized for basic biological properties (the optimum temperature, pH and the optimum carbon and nitrogen sources). The result showed B. fabiopsis could grow in 5-30℃, pH values of 2.0-10.0. The most optimum temperature is 20-25℃for mycelium growth and the most optimum pH value is 4.0-5.0. The most suitable carbon and nitrogen sources were soluble starch and alanine, respectively. B. fabiopsis, B. fabae and B. cinerea showed a dramatic difference in response to temperature, pH value and nutritent sources. B. sinoallii could grow in 5-30℃, pH values of 2.0-10.0 with the optimum temperature of 20℃and ph value of 4.0-5.0. The most suitable carbon and nitrogen sources were glucose and peptone/alanine, respectively. B. sinoallii and B. squamosa also showed a dramatic difference in response to temperature, pH value and nutrient sources. B. sinoallii showed better growth at 25℃and 28℃than B. squamosa.
4. The genetic diversity of Botrytis cinerea isolated from different hosts in Hubei Province was studied based on mophyological characteristics, pathogenicity, fungicide resistance and some molecular features. The results showed that:isolates of B. cinerea have three types:mycelium type, conidial type and sclerotial type. A PCR method was developed to identify the transposable elements Boty and Flipper that form two groups, transposa and vacuma. Most of isolates are sensitive to four fungicides.
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