中国贵州、云南、广西炭疽菌属真菌多基因分子系统学研究
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摘要
炭疽菌属(Colletotrichum Cda.)是分生孢子盘具有刚毛的一类全球性分布的植物病原菌,又名刺盘孢属。该属真菌引起多种作物采收前和采收后炭疽病,每年导致大量的经济损失。自Corda于1831年建立炭疽菌属以来,经历了三次较大的变更,但其分类仍然十分混乱。有必要对该属真菌进行全面研究,明确种的概念,理顺该属真菌的系统学关系。
本研究主要采集具有炭疽病病症的茄科、豆科、百合科、兰科、石蒜科、葫芦科植物的叶、茎、果实等为实验材料,以单孢分离法分离病原菌；同时从国际菌种保藏中心收集部分模式菌株进行比较研究。采用形态学结合多基因(肌动蛋白-ACT、钙调蛋白-CAL、几丁质合成酶A-CHSI、3-磷酸甘油醛脱氢酶-GPDH、核糖体内转录间区-ITS、微管蛋白基因基因-TUB2)分子系统学并辅以寄主范围的方法,确定分离物的分类地位,弄清以上3省的主要炭疽病病原,探讨形态指标和多基因系统学的分类价值。
从云南、贵州、广西3省14市(县)的茄科等7科35属38种寄主共采集323份具有炭疽病病症的标本,分离得到190株菌株,表明该地区炭疽病非常普遍。其中豆科47株、百合科26株、茄科72株、石蒜科17株、兰科22株、葫芦科4株；五加科2株；并从荷兰微生物菌种保存中心等国际菌种保藏中心收集了6株模式菌株或可信度高的菌株参与比较研究。
结合形态学、多基因系统学和寄主范围,将分离的190株菌株确定为22个分类单元。其中新种5个,分别为朱顶红炭疽菌(Colletotrichum cliviae Y. L. Yang, Zuo Y Liu, K.D. Hyde & L. Cai, sp. Nov.)、君子兰炭疽菌(C. hemerocallidis Y. L. Yang, Zuo Y. Liu, K.D. Hyde & L. Cai, sp. Nov.)、喀斯特炭疽菌(C. hippeastri Y. L. Yang, Zuo Y. Liu, K.D. Hyde & L. Cai, sp. Nov.)、蜘蛛兰炭疽菌(C. hymenocallidis Y. L. Yang, Zuo Y. Liu, K.D. Hyde & L. Cai, sp. Nov.)和萱草炭疽菌(C. karstii Y. L. Yang, Zuo Y. Liu, K.D. Hyde & L. Cai, sp. Nov.)。对朱顶红炭疽菌、君子兰炭疽菌、喀斯特炭疽菌、蜘蛛兰炭疽菌的致病性测试显示,该4种真菌没有严格的寄主专一性。中国新记录种5种,分别为C. brevispora Phoulivong, Noireung, L. Cai & K.D. Hyde、C. fructicola Prihastuti,L.Cai & K.D.Hyde.C.liriopes Damm,P.F.Cannon & Crous.C. siamense Prihastuti,L. Cai & K.D.Hyde和C. spaethianum(Allesch.)Damm,P.F.Cannon & Crous。已知种9种。全球近三十年来在科技论文中出现的炭疽菌属真菌仅为66个,在我国云南、广西和贵州3省的检出率就达到三分之一,表明该地区植物炭疽病比较普遍。
豆类炭疽菌在我国贵州、云南和广西非常严重,共分离到41株,占分离总数的21.6%,是引起豆科豇豆、大豆,茄科辣椒的主要炭疽病病原,分别占该3种作物炭疽病病原检出总数的84.2%、69.4%和17.2%；此外也感染茄科的西红柿、茄子和石蒜科的蜘蛛兰。君子兰炭疽菌的寄主较广泛,能感染石蒜科、兰科、豆科、茄科的多种植物,但不严重,共分离到13株菌株,占分离总数的6.8%。2008年引起贵州凤冈县局部爆发的辣椒炭疽病的病原真菌为壳皮炭疽菌。胶孢炭疽菌的寄主范围相对较窄,仅从兰科万代兰属的某一种、茄科枸杞、百合科萱草共分离到10株,占分离总数的5.3%。
本研究首次系统的对茄科、豆科、百合科、兰科、石蒜科植物进行炭疽病病原的分离和鉴定,弄清了该地区的炭疽病病原种类。在国内首次采用多基因系统学结合形态学方法识别炭疽菌属真菌,这一方法对该属及其它属的鉴定具有一定的借鉴价值。同时,本研究不仅在真菌学和植物病理学上具有重要的科学价值,而且在防治植物炭疽病、作物育种方面具有重要的实践意义,也有助于海关有效检疫。
Colletotrichum Cda. is one of the most important plant pathogens worldwide causing the economically important disease anthracnose in a wide range of hosts including cereals, legumes, vegetables, perennial crops and tree fruits. The current identifying Colletotrichum species is largely based on a combination of morphological and cultural characteristics. Due to limited numbers of morphological characters available in culture coupled with inherent phenotypic plasticity, correct identification of the species has always been difficult since introduced by Corda in 1831. it is urgent to elucidate species concept and clarify the systematics of the genus. Colletotrichum species have not been studyied thoroughly in China. The main objective of this study was to identify correctly and characterize the Colletotrichum species associated with anthracnose in Guizhou, Yunan and Guangxi, China, and then evaluated the role of morphology characters and multilocue phylogenics analysis through compairing those type or epitype and other isolates.
In the present study, we collected the tissues with anthracnose from Amaryllidaceae, Leguminoseae, Liliaceae, Orchideaceae, Solanaceae plants in Guizhou, Yunnan and Guangxi, China, and obtained the type or epitype stains from the international culture centres for compairing study. We used the single spore isolation method to obtain the pathogen from the tissues with anthracnose and recognized those isolates based on morphological characters and multilocus (ACT, CAL, CHS I, GPDH, ITS, TUB2) sequences phylogenetic analysis, as well as their host range.
In this study, total 190 isolates were obtained from 38 hosts belonging to 35 genera of 7 families in Guizhou, Guangxi and Yunnan, China, and six type (epitypes) or well known strains were collected from some international centres. Combination morphological, melocular and host rangs, the 190 isolates represent 22 taxa. Among of them,5 are novel to science. They are Colletotrichum cliviae Y. L. Yang, Zuo Y. Liu, K.D. Hyde & L. Cai, sp. Nov., C. hemerocallidis Y. L. Yang, Zuo Y. Liu, K.D. Hyde & L. Cai, sp. Nov., C. hippeastri Y. L. Yang, Zuo Y. Liu, K.D. Hyde & L. Cai, sp. Nov., C. hymenocallidis Y. L. Yang, Zuo Y. Liu, K.D. Hyde & L. Cai, sp. Nov. and C. karstii Y. L. Yang, Zuo Y. Liu, K.D. Hyde & L. Cai, sp. Nov. The pathogenecity for Colletotrichum cliviae, C. hippeastri, C. hymenocallidis, C. karstii showed that they are not specific-host, and following isolation confirmed this. In addition,5 species, Colletotrichum brevispora Phoulivong, Noireung, L. Cai & K.D. Hyde, C. fructicola Prihastuti, L. Cai & K.D. Hyde, C. liriopes Damm, P.F. Cannon & Crous, C. siamense Prihastuti, L. Cai & K.D. Hyde and C. spaethianum (Allesch.) Damm, P. F. Cannon & Crous are new records in China. There are 66 species recorded in scientific literatures or formally accepted in the recent three decades. Our isolates in this study from the three provinces is one-third. This shows that the plant anthracnose is severe in the area.
Colletotrichum truncatum is a mainly agent causing anthracnose in the three provinces, China. Total 41 strains of this species were isolated in this study. This species causes severe anthracnose on longbean, common bean and chili (84.2%,69.4 and 17.2% to the total anthracnose pathogen of the three crops, respectively). In addition, this species also, infects tomato, eggplant (Solanaceae) and spider lily (Amaryllidaceae). Colletotrichum cliviae has an abroad host range, it occurres on plants in Amaryllidaceae, Leguminoseae, Orchideaceae and Solanaceae. But this species is not the mainly anthracnose cause agent on these plants.13 isolates were obtained from above motioned families,6.8% to the total 186 isolates. Colletotrithum crassipes was responsibility for chili anthracnose in Fenggang, Guizhou in 2008.10 isolates (5.3% to total 190 isolates) is Colletotrichum gloeosporioides, this indicated that this species is not abroad host range. C. gloeosporioides caused anthracnose on Lycium chinense, Vanda sp., Capsicum annuum, Hemerocallis fulva and H. citrina.
This is first detailed investigation the anthracnose cause agent on the Amaryllida-ceae, Leguminoseae, Liliaceae, Orchideaceae, Solanaceae plants, and the above outcome will have important practical implications for the plant pathology, plant breeding and quarantine communities both internationally and in China.
本研究主要采集具有炭疽病病症的茄科、豆科、百合科、兰科、石蒜科、葫芦科植物的叶、茎、果实等为实验材料,以单孢分离法分离病原菌；同时从国际菌种保藏中心收集部分模式菌株进行比较研究。采用形态学结合多基因(肌动蛋白-ACT、钙调蛋白-CAL、几丁质合成酶A-CHSI、3-磷酸甘油醛脱氢酶-GPDH、核糖体内转录间区-ITS、微管蛋白基因基因-TUB2)分子系统学并辅以寄主范围的方法,确定分离物的分类地位,弄清以上3省的主要炭疽病病原,探讨形态指标和多基因系统学的分类价值。
从云南、贵州、广西3省14市(县)的茄科等7科35属38种寄主共采集323份具有炭疽病病症的标本,分离得到190株菌株,表明该地区炭疽病非常普遍。其中豆科47株、百合科26株、茄科72株、石蒜科17株、兰科22株、葫芦科4株；五加科2株；并从荷兰微生物菌种保存中心等国际菌种保藏中心收集了6株模式菌株或可信度高的菌株参与比较研究。
结合形态学、多基因系统学和寄主范围,将分离的190株菌株确定为22个分类单元。其中新种5个,分别为朱顶红炭疽菌(Colletotrichum cliviae Y. L. Yang, Zuo Y Liu, K.D. Hyde & L. Cai, sp. Nov.)、君子兰炭疽菌(C. hemerocallidis Y. L. Yang, Zuo Y. Liu, K.D. Hyde & L. Cai, sp. Nov.)、喀斯特炭疽菌(C. hippeastri Y. L. Yang, Zuo Y. Liu, K.D. Hyde & L. Cai, sp. Nov.)、蜘蛛兰炭疽菌(C. hymenocallidis Y. L. Yang, Zuo Y. Liu, K.D. Hyde & L. Cai, sp. Nov.)和萱草炭疽菌(C. karstii Y. L. Yang, Zuo Y. Liu, K.D. Hyde & L. Cai, sp. Nov.)。对朱顶红炭疽菌、君子兰炭疽菌、喀斯特炭疽菌、蜘蛛兰炭疽菌的致病性测试显示,该4种真菌没有严格的寄主专一性。中国新记录种5种,分别为C. brevispora Phoulivong, Noireung, L. Cai & K.D. Hyde、C. fructicola Prihastuti,L.Cai & K.D.Hyde.C.liriopes Damm,P.F.Cannon & Crous.C. siamense Prihastuti,L. Cai & K.D.Hyde和C. spaethianum(Allesch.)Damm,P.F.Cannon & Crous。已知种9种。全球近三十年来在科技论文中出现的炭疽菌属真菌仅为66个,在我国云南、广西和贵州3省的检出率就达到三分之一,表明该地区植物炭疽病比较普遍。
豆类炭疽菌在我国贵州、云南和广西非常严重,共分离到41株,占分离总数的21.6%,是引起豆科豇豆、大豆,茄科辣椒的主要炭疽病病原,分别占该3种作物炭疽病病原检出总数的84.2%、69.4%和17.2%；此外也感染茄科的西红柿、茄子和石蒜科的蜘蛛兰。君子兰炭疽菌的寄主较广泛,能感染石蒜科、兰科、豆科、茄科的多种植物,但不严重,共分离到13株菌株,占分离总数的6.8%。2008年引起贵州凤冈县局部爆发的辣椒炭疽病的病原真菌为壳皮炭疽菌。胶孢炭疽菌的寄主范围相对较窄,仅从兰科万代兰属的某一种、茄科枸杞、百合科萱草共分离到10株,占分离总数的5.3%。
本研究首次系统的对茄科、豆科、百合科、兰科、石蒜科植物进行炭疽病病原的分离和鉴定,弄清了该地区的炭疽病病原种类。在国内首次采用多基因系统学结合形态学方法识别炭疽菌属真菌,这一方法对该属及其它属的鉴定具有一定的借鉴价值。同时,本研究不仅在真菌学和植物病理学上具有重要的科学价值,而且在防治植物炭疽病、作物育种方面具有重要的实践意义,也有助于海关有效检疫。
Colletotrichum Cda. is one of the most important plant pathogens worldwide causing the economically important disease anthracnose in a wide range of hosts including cereals, legumes, vegetables, perennial crops and tree fruits. The current identifying Colletotrichum species is largely based on a combination of morphological and cultural characteristics. Due to limited numbers of morphological characters available in culture coupled with inherent phenotypic plasticity, correct identification of the species has always been difficult since introduced by Corda in 1831. it is urgent to elucidate species concept and clarify the systematics of the genus. Colletotrichum species have not been studyied thoroughly in China. The main objective of this study was to identify correctly and characterize the Colletotrichum species associated with anthracnose in Guizhou, Yunan and Guangxi, China, and then evaluated the role of morphology characters and multilocue phylogenics analysis through compairing those type or epitype and other isolates.
In the present study, we collected the tissues with anthracnose from Amaryllidaceae, Leguminoseae, Liliaceae, Orchideaceae, Solanaceae plants in Guizhou, Yunnan and Guangxi, China, and obtained the type or epitype stains from the international culture centres for compairing study. We used the single spore isolation method to obtain the pathogen from the tissues with anthracnose and recognized those isolates based on morphological characters and multilocus (ACT, CAL, CHS I, GPDH, ITS, TUB2) sequences phylogenetic analysis, as well as their host range.
In this study, total 190 isolates were obtained from 38 hosts belonging to 35 genera of 7 families in Guizhou, Guangxi and Yunnan, China, and six type (epitypes) or well known strains were collected from some international centres. Combination morphological, melocular and host rangs, the 190 isolates represent 22 taxa. Among of them,5 are novel to science. They are Colletotrichum cliviae Y. L. Yang, Zuo Y. Liu, K.D. Hyde & L. Cai, sp. Nov., C. hemerocallidis Y. L. Yang, Zuo Y. Liu, K.D. Hyde & L. Cai, sp. Nov., C. hippeastri Y. L. Yang, Zuo Y. Liu, K.D. Hyde & L. Cai, sp. Nov., C. hymenocallidis Y. L. Yang, Zuo Y. Liu, K.D. Hyde & L. Cai, sp. Nov. and C. karstii Y. L. Yang, Zuo Y. Liu, K.D. Hyde & L. Cai, sp. Nov. The pathogenecity for Colletotrichum cliviae, C. hippeastri, C. hymenocallidis, C. karstii showed that they are not specific-host, and following isolation confirmed this. In addition,5 species, Colletotrichum brevispora Phoulivong, Noireung, L. Cai & K.D. Hyde, C. fructicola Prihastuti, L. Cai & K.D. Hyde, C. liriopes Damm, P.F. Cannon & Crous, C. siamense Prihastuti, L. Cai & K.D. Hyde and C. spaethianum (Allesch.) Damm, P. F. Cannon & Crous are new records in China. There are 66 species recorded in scientific literatures or formally accepted in the recent three decades. Our isolates in this study from the three provinces is one-third. This shows that the plant anthracnose is severe in the area.
Colletotrichum truncatum is a mainly agent causing anthracnose in the three provinces, China. Total 41 strains of this species were isolated in this study. This species causes severe anthracnose on longbean, common bean and chili (84.2%,69.4 and 17.2% to the total anthracnose pathogen of the three crops, respectively). In addition, this species also, infects tomato, eggplant (Solanaceae) and spider lily (Amaryllidaceae). Colletotrichum cliviae has an abroad host range, it occurres on plants in Amaryllidaceae, Leguminoseae, Orchideaceae and Solanaceae. But this species is not the mainly anthracnose cause agent on these plants.13 isolates were obtained from above motioned families,6.8% to the total 186 isolates. Colletotrithum crassipes was responsibility for chili anthracnose in Fenggang, Guizhou in 2008.10 isolates (5.3% to total 190 isolates) is Colletotrichum gloeosporioides, this indicated that this species is not abroad host range. C. gloeosporioides caused anthracnose on Lycium chinense, Vanda sp., Capsicum annuum, Hemerocallis fulva and H. citrina.
This is first detailed investigation the anthracnose cause agent on the Amaryllida-ceae, Leguminoseae, Liliaceae, Orchideaceae, Solanaceae plants, and the above outcome will have important practical implications for the plant pathology, plant breeding and quarantine communities both internationally and in China.
引文
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