健康与染溃疡病107杨、中林46杨树皮真菌和细菌群落研究
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摘要
杨树是杨柳科(Salicacae)杨属(Populus L.)植物的统称,是世界上分布最广、适应性最强的树种,近年来华北平原大面积推广栽培优良品种107杨(Populus×euramericana cv.'74/76'),仅在河南濮阳就有数十万亩。但自2005年以来欧美杨品种107杨和中林46杨(P.×euramericana cv.'Zhonglin46')发生一种新的溃疡病,对欧美杨的发展造成威胁。如何有效地控制病害已成为生产上迫切需要解决的问题,利用植物内生菌控制病害可能成为防治的一个新途径。
本研究以欧美杨品种107杨和中林46杨为研究对象,采用组织分离方法,对欧美杨健康树皮内生真菌、细菌进行了分离鉴定、物种多样性及优势种群季节性变化动态分析。对近年发生的欧美杨溃疡病病斑中的真菌和细菌群落组成进行了分析,并对分离到的不动杆菌属(Acinetobacter Brisou and Prevot1954)和Lonsdalea Brady et al.2012属的细菌进行多相分类鉴定。通过比较两品种健康植株与染病植株干部树皮内微生物菌群的组成和变化,分析讨论病害发生过程中可能的协同菌群,明确溃疡病发生对杨树树皮中微生物群落结构和组成的影响。研究工作可为理解微生物环境对欧美杨溃疡病发生的影响,发掘可能的生防菌资源,进而有效防治欧美杨溃疡病提供理论依据。主要研究结果如下：
107杨和中林46杨树皮具有丰富的内生真菌、细菌物种多样性,其内生真菌多样性指数分别为2.38和2.36,分离得到17属、30种及未定种子囊菌内生真菌以及32属58种的内生细菌。该结果揭示了107杨和中林46杨树皮中蕴藏着大量的待开发的微生物资源。内生真菌优势种群包括链格孢(Alternaria alternata (Fr.) Keissl.1912)、葡萄座腔菌(Botryosphaeria dothidea (Moug.) Ces.&De Not.1863)、桑砖红镰孢(Fusarium lateritium Nees1816)、间座壳菌(Diaporthe sp.)和Fusarium incarnatum-equiseti,其中链格孢、葡萄座腔菌是最为常见的优势种群。
温度湿度气候条件对内生菌数量、组成及其多样性指数具有重要的影响,内生细菌数量季节变化规率为夏季>春季和秋季>冬季；多样性指数季节变化趋势为春季、秋季>夏季、冬季。
健康107杨和中林46杨树皮内生菌组成中,有很多种类是对植物生长或微生态环境有利的菌群,如Pseudomonas、Pantoea等能产生IAA促进植物生长并且能起到解磷作用的属,Rhizobium能够结瘤固氮的属,Trichoderma等用于植物病害生物防治的属,该结果揭示了两个品种树皮内具有丰富的待开发的偍生、生防菌资源。107杨和中林46杨健康树皮中分离到了三种植物潜在致病菌,如葡葡座腔菌、茄镰孢菌(Fusarium solani (Mart.) Sacc.1881)和茄科黄单胞(Xanthomonas vesicatoria (ex Doidge1920)Vauterin et al.1995),而且葡葡座腔菌还足优势种群之一,说明本研究试验林的杨树存在较大的发病风险。
溃疡病的发生改变了树皮的生理生化组成与结构,也改变了其中的真菌与细菌群落结构,使得健康和染病107杨和中林46杨树皮真菌细菌组成差异非常大。新增加的微生物种类可能来自土壤、空气和树皮表面。健康植株树皮微生态系统中以对植物生长或微生态环境有利的菌群为主,如Pseudomonas)属、Pantoea属、Rhizobium属、Microbacterium属和Bacillus属。而感染溃疡病的树皮发病部位微生态系统则以各类病原菌、肠道细菌和能发酵细菌为主,真菌优势种群为茄镰孢菌,细菌优势种群为"Lonsdalea quercina subsp. populi'Timea Toth et al.2012,另外还包括柳树水纹病病菌(Brenncria salicis Hauben et al.1999)、马铃薯茎腐病病菌(Pectobacterium wasabiae (Goto and Matsumoto1987) Gardan et al.2003)、软腐果胶杆菌软腐亚种(Pectobacterium carotovorum subsp. carotovorum (Jones1901) Hauben et al.1999)等植物病原菌,这些病原菌与该病害病原菌的关系有待进一步研究。另外还分离到乳杆菌(Lactobacillus plantarum (Orla-Jensen1919) Bergey et al.1923)、微黄棒状杆菌(Corynebacterium flavescens Barksdale et al.1979)和变异棒杆菌(Corynebacterium variabile (Miiller1961) Collins1987)等发酵型菌株,这些菌种的作用有待进一步研究。
欧美杨主要病原菌Lonsdalea属多相分类研究表明,菌株N-5-1、HZ1031和4-4等5个菌株鉴定为‘L. quercina subsp. populi")亚种,是我国的新纪录。本研究将7个不动杆菌的菌株鉴定为3个新种,菌株BQ4-1T和NH13-2命名为Acinetobactcr puyangensis sp. nov、菌株PBJ7T和PZ6拟命名为Acinetobactcr populi sp. nov、菌株BJl、2C-3-1和HF5-2拟命名为Acinetobacter qingfengensis sp. nov.。
Poplar trees, deciduous plants, are part of the Salicaceae family, widespread in Europe, Asia, North America, and is one of the world's most widely distributed and most adaptable tree species. Populus x euramericana cv.'74/76'was wildly cultivated in the north China plain in recent years, and was planted more than hundreds of thousands of acres in Puyang, Henan province. A new canker desease was observed from P. xeuramericana cv.'74/76'and P. x euramericana cv.'zhonglin46', which threats the cultivation of those cultivars. How to effectively control the diseases has become urgent problem needed to be solved, the disease prevention and control using plant endophytes has become a possible approach.
In order to understand predominant species dynamic and diversity of fungal and bacterial endophytes in barks of Populus x euramericana, fungal and bacterial endophytes were isolated from barks of P. x euramericana cv.'74/76'and P. x euramericana cv.'Zhonglin46,and identified by sequence analysis of the internal transcribed spacer (ITS) and morphological characterization by microscopic observations. The seasonal dynamic analysis of fungal and bacterial endophytes in both cultivars was performed. The cultivable bacteria and fungi from poplar canker bark in Puyang city were isolated, and the dominant population of fungi and bacteria hosted in P. x euroamericana canker was analysed. And the isolates belong to the genus Acinetobacter and Lonsdalea were identified by polyphasic taxonomy. The main results and conclusions are as follows:
Fungal endophytes isolated from barks of P. x euramericana cv.'74/76'and P. x euramericana cv.'zhonglin46'were classified into30different fungal taxa belonging to17genera of Ascomycota. The results revealed that there are high phylogenetic diversity of endophytic fungus in the bark of P. x euramericana cv.'74/76'and P.×euramericana cv.'zhonglin46'.Endophytic bacteria strains isolated from barks of P. x euramericana cv.'74/76'and P. x euramericana cv.'zhonglin46' belong to58species of32genera, including32species of20genera from the barks of P. x euramericana cv.'74/76' and36taxon of19genera from the barks of P×euramericana cv.'zhonglin46'. Alternaria alternata, Botryosphaeria dothidea, Fusarium spp., Diaporthe spp., were predominant species of fungal endophytes in barks of both cultivars of P.×euramericana while Alternaria alternata and Botryosphaeria dothidea were most common predominant species.
Climate factors including temperature and humidity have important effects on the endogenous bacteria composition and dominant population of endophytic microorganism of P×euramericana cv. '74/76'and P.×euramericana cv."zhonglin46. The results showed that bacterial endophytes diversity index in spring and autuin was higher than that in summer and winter. In the cold season, genus Rhizobium and other beneficial strains become a dominant population, which maybe increase resistance of the tree to the bad weather.
In the endophytic bacterial communities of P.×euramericana cv.'74/76'and P.×euramericana cv.'Zhonglin46, there are many kinds of beneficial strains to plant, such as Pseudomonas, Pantoea, Rhizobium, Trichoclerma, which revealed that there are a lot of beneficial and bio-control microbial resources. While, two potential plant pathogens, B. doth idea and F. solani, were isolated from the bark of P.×euramericana cv.'74/76'and P.×euramericana cv."zhonglin46, which are important pathogens to some plants. So, the plants tested in this study have a big risk of the desease occurrence.
There are big difference in the microbial communities from healthy and canker-infected P.×euramericana cv.'74/76'and P.×euramericana cv.'Zhonglin46'. The beneficial strains were predominant in the healthy plants bark micro-ecosystem, including Pseudomonas spp., Pantoea spp., Rhizobium spp., Microbacterium spp. and Bacillus spp. While, pathogenic strains, intestinal bacteria and fermentation bacteria arc common species in the descased bark sites. Bacterial dominant population is Lonsdalea quercina subsp. populi in the sites of the bark infected canker, and Fusarium solani is fungal dominant population in the micro-ecosystem of the canker-infected bark. Furthermore, Brenneria salicis, Pectobacterium wasabiae, Pectohacterium carolovorum subsp. carotovorum, Alcaligenes faecalis, Lactobacillus plantarum, Corynehaelerium flavescens and Corynebacterium variubile were also isolated from the diseased bark parts, the relationship between those strains and the causal pathogen of the canker disease needs further study. According to the physiological and biochemical characteristics, genetic type features, five strains isolated from diseased barks, including N-5-1, HZ1031and4-4, were identified as Lonsdalea quercina subsp. populi, a new record species in China. Seven strains belong to genus Acinetobacter were identified as three new species, and strains BQ4-1T and NII13were named Acinetobacter puyangensis sp. nov; strains PBJ7and PZ6named Acinetobactei popli sp. nov, strains2E3J1T, HI'5-2and2C-3-1named Acinetobacter qingfengensis sp. nov
本研究以欧美杨品种107杨和中林46杨为研究对象,采用组织分离方法,对欧美杨健康树皮内生真菌、细菌进行了分离鉴定、物种多样性及优势种群季节性变化动态分析。对近年发生的欧美杨溃疡病病斑中的真菌和细菌群落组成进行了分析,并对分离到的不动杆菌属(Acinetobacter Brisou and Prevot1954)和Lonsdalea Brady et al.2012属的细菌进行多相分类鉴定。通过比较两品种健康植株与染病植株干部树皮内微生物菌群的组成和变化,分析讨论病害发生过程中可能的协同菌群,明确溃疡病发生对杨树树皮中微生物群落结构和组成的影响。研究工作可为理解微生物环境对欧美杨溃疡病发生的影响,发掘可能的生防菌资源,进而有效防治欧美杨溃疡病提供理论依据。主要研究结果如下：
107杨和中林46杨树皮具有丰富的内生真菌、细菌物种多样性,其内生真菌多样性指数分别为2.38和2.36,分离得到17属、30种及未定种子囊菌内生真菌以及32属58种的内生细菌。该结果揭示了107杨和中林46杨树皮中蕴藏着大量的待开发的微生物资源。内生真菌优势种群包括链格孢(Alternaria alternata (Fr.) Keissl.1912)、葡萄座腔菌(Botryosphaeria dothidea (Moug.) Ces.&De Not.1863)、桑砖红镰孢(Fusarium lateritium Nees1816)、间座壳菌(Diaporthe sp.)和Fusarium incarnatum-equiseti,其中链格孢、葡萄座腔菌是最为常见的优势种群。
温度湿度气候条件对内生菌数量、组成及其多样性指数具有重要的影响,内生细菌数量季节变化规率为夏季>春季和秋季>冬季；多样性指数季节变化趋势为春季、秋季>夏季、冬季。
健康107杨和中林46杨树皮内生菌组成中,有很多种类是对植物生长或微生态环境有利的菌群,如Pseudomonas、Pantoea等能产生IAA促进植物生长并且能起到解磷作用的属,Rhizobium能够结瘤固氮的属,Trichoderma等用于植物病害生物防治的属,该结果揭示了两个品种树皮内具有丰富的待开发的偍生、生防菌资源。107杨和中林46杨健康树皮中分离到了三种植物潜在致病菌,如葡葡座腔菌、茄镰孢菌(Fusarium solani (Mart.) Sacc.1881)和茄科黄单胞(Xanthomonas vesicatoria (ex Doidge1920)Vauterin et al.1995),而且葡葡座腔菌还足优势种群之一,说明本研究试验林的杨树存在较大的发病风险。
溃疡病的发生改变了树皮的生理生化组成与结构,也改变了其中的真菌与细菌群落结构,使得健康和染病107杨和中林46杨树皮真菌细菌组成差异非常大。新增加的微生物种类可能来自土壤、空气和树皮表面。健康植株树皮微生态系统中以对植物生长或微生态环境有利的菌群为主,如Pseudomonas)属、Pantoea属、Rhizobium属、Microbacterium属和Bacillus属。而感染溃疡病的树皮发病部位微生态系统则以各类病原菌、肠道细菌和能发酵细菌为主,真菌优势种群为茄镰孢菌,细菌优势种群为"Lonsdalea quercina subsp. populi'Timea Toth et al.2012,另外还包括柳树水纹病病菌(Brenncria salicis Hauben et al.1999)、马铃薯茎腐病病菌(Pectobacterium wasabiae (Goto and Matsumoto1987) Gardan et al.2003)、软腐果胶杆菌软腐亚种(Pectobacterium carotovorum subsp. carotovorum (Jones1901) Hauben et al.1999)等植物病原菌,这些病原菌与该病害病原菌的关系有待进一步研究。另外还分离到乳杆菌(Lactobacillus plantarum (Orla-Jensen1919) Bergey et al.1923)、微黄棒状杆菌(Corynebacterium flavescens Barksdale et al.1979)和变异棒杆菌(Corynebacterium variabile (Miiller1961) Collins1987)等发酵型菌株,这些菌种的作用有待进一步研究。
欧美杨主要病原菌Lonsdalea属多相分类研究表明,菌株N-5-1、HZ1031和4-4等5个菌株鉴定为‘L. quercina subsp. populi")亚种,是我国的新纪录。本研究将7个不动杆菌的菌株鉴定为3个新种,菌株BQ4-1T和NH13-2命名为Acinetobactcr puyangensis sp. nov、菌株PBJ7T和PZ6拟命名为Acinetobactcr populi sp. nov、菌株BJl、2C-3-1和HF5-2拟命名为Acinetobacter qingfengensis sp. nov.。
Poplar trees, deciduous plants, are part of the Salicaceae family, widespread in Europe, Asia, North America, and is one of the world's most widely distributed and most adaptable tree species. Populus x euramericana cv.'74/76'was wildly cultivated in the north China plain in recent years, and was planted more than hundreds of thousands of acres in Puyang, Henan province. A new canker desease was observed from P. xeuramericana cv.'74/76'and P. x euramericana cv.'zhonglin46', which threats the cultivation of those cultivars. How to effectively control the diseases has become urgent problem needed to be solved, the disease prevention and control using plant endophytes has become a possible approach.
In order to understand predominant species dynamic and diversity of fungal and bacterial endophytes in barks of Populus x euramericana, fungal and bacterial endophytes were isolated from barks of P. x euramericana cv.'74/76'and P. x euramericana cv.'Zhonglin46,and identified by sequence analysis of the internal transcribed spacer (ITS) and morphological characterization by microscopic observations. The seasonal dynamic analysis of fungal and bacterial endophytes in both cultivars was performed. The cultivable bacteria and fungi from poplar canker bark in Puyang city were isolated, and the dominant population of fungi and bacteria hosted in P. x euroamericana canker was analysed. And the isolates belong to the genus Acinetobacter and Lonsdalea were identified by polyphasic taxonomy. The main results and conclusions are as follows:
Fungal endophytes isolated from barks of P. x euramericana cv.'74/76'and P. x euramericana cv.'zhonglin46'were classified into30different fungal taxa belonging to17genera of Ascomycota. The results revealed that there are high phylogenetic diversity of endophytic fungus in the bark of P. x euramericana cv.'74/76'and P.×euramericana cv.'zhonglin46'.Endophytic bacteria strains isolated from barks of P. x euramericana cv.'74/76'and P. x euramericana cv.'zhonglin46' belong to58species of32genera, including32species of20genera from the barks of P. x euramericana cv.'74/76' and36taxon of19genera from the barks of P×euramericana cv.'zhonglin46'. Alternaria alternata, Botryosphaeria dothidea, Fusarium spp., Diaporthe spp., were predominant species of fungal endophytes in barks of both cultivars of P.×euramericana while Alternaria alternata and Botryosphaeria dothidea were most common predominant species.
Climate factors including temperature and humidity have important effects on the endogenous bacteria composition and dominant population of endophytic microorganism of P×euramericana cv. '74/76'and P.×euramericana cv."zhonglin46. The results showed that bacterial endophytes diversity index in spring and autuin was higher than that in summer and winter. In the cold season, genus Rhizobium and other beneficial strains become a dominant population, which maybe increase resistance of the tree to the bad weather.
In the endophytic bacterial communities of P.×euramericana cv.'74/76'and P.×euramericana cv.'Zhonglin46, there are many kinds of beneficial strains to plant, such as Pseudomonas, Pantoea, Rhizobium, Trichoclerma, which revealed that there are a lot of beneficial and bio-control microbial resources. While, two potential plant pathogens, B. doth idea and F. solani, were isolated from the bark of P.×euramericana cv.'74/76'and P.×euramericana cv."zhonglin46, which are important pathogens to some plants. So, the plants tested in this study have a big risk of the desease occurrence.
There are big difference in the microbial communities from healthy and canker-infected P.×euramericana cv.'74/76'and P.×euramericana cv.'Zhonglin46'. The beneficial strains were predominant in the healthy plants bark micro-ecosystem, including Pseudomonas spp., Pantoea spp., Rhizobium spp., Microbacterium spp. and Bacillus spp. While, pathogenic strains, intestinal bacteria and fermentation bacteria arc common species in the descased bark sites. Bacterial dominant population is Lonsdalea quercina subsp. populi in the sites of the bark infected canker, and Fusarium solani is fungal dominant population in the micro-ecosystem of the canker-infected bark. Furthermore, Brenneria salicis, Pectobacterium wasabiae, Pectohacterium carolovorum subsp. carotovorum, Alcaligenes faecalis, Lactobacillus plantarum, Corynehaelerium flavescens and Corynebacterium variubile were also isolated from the diseased bark parts, the relationship between those strains and the causal pathogen of the canker disease needs further study. According to the physiological and biochemical characteristics, genetic type features, five strains isolated from diseased barks, including N-5-1, HZ1031and4-4, were identified as Lonsdalea quercina subsp. populi, a new record species in China. Seven strains belong to genus Acinetobacter were identified as three new species, and strains BQ4-1T and NII13were named Acinetobacter puyangensis sp. nov; strains PBJ7and PZ6named Acinetobactei popli sp. nov, strains2E3J1T, HI'5-2and2C-3-1named Acinetobacter qingfengensis sp. nov
引文
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