黄淮海夏玉米主产区茎腐病主要病原菌及优势种分析
|
摘要
【目的】明确黄淮海夏玉米主产区玉米茎腐病的主要病原菌组成及优势种,为病原菌致病机制、抗性育种及茎腐病防治提供依据。【方法】于2014—2017年采集黄淮海夏玉米主产区3个省(河北、河南、山东)的玉米茎腐病样本850份。通过形态学、通用引物和特异引物对病组织分离物进行鉴定,并采用上述引物直接检测病组织。整合分析分离物鉴定法和组织分子检测法的结果以确定玉米茎腐病的主要病原菌及优势种;分析主要病原菌在各省间以及同一省份不同年度的检出率,揭示主要病原菌的种群动态变化;分析单个样本中病原菌的检出率,探讨多种病原菌的共存模式。【结果】有667份样本检测出真菌或卵菌,占总样本的78.47%;年度间样本检出率存在差异,2014年的检出率不足50%,而2015—2017年的检出率相近,均高于90%。在所有样本中共检出20属46种真菌或卵菌,其中镰孢菌(Fusarium spp.)的检出率最高,为89.96%,包括禾谷镰孢复合种(F. graminearum species complex)、层出镰孢(F. proliferatum)、拟轮枝镰孢(F. verticillioides)、厚垣镰孢(F.chlamydosporum)、亚粘团镰孢(F. subglutinans)、尖镰孢(F. oxysporum)、黄色镰孢(F. culmorum)、藤仓镰孢(F. fujikuroi)、变红镰孢(F. incarnatum)、木贼镰孢(F. equiseti)和茄镰孢(F. solani)11个种;腐霉菌(Pythium spp.)的检出率为34.18%,包括芒孢腐霉(P. aristosporum)、禾生腐霉(P. graminicola)、棘腐霉(P. acanthicum)、孤雌腐霉(P. amasculinum)和寡雄腐霉(P. oligandrum)5个种。拟轮枝镰孢、禾谷镰孢复合种、芒孢腐霉和层出镰孢为4种主要病原菌,检出率依次为62.07%、46.93%、29.09%和28.04%,其中拟轮枝镰孢为优势种。各省之间,上述4种主要病原菌的检出率存在一定差异。拟轮枝镰孢在河北省的检出率为73.98%,明显高于该菌在河南省和山东省的检出率;禾谷镰孢复合种在3个省的检出率相近;层出镰孢和芒孢腐霉在山东省的检出率分别为35.78%和34.31%,均高于在其他两省的检出率。同一省份不同年度上述4种主要病原菌的检出率呈动态变化,其中任何一种病原菌均有可能上升为优势种。进一步分析表明,单个样本中可以检测出一种或多种病原菌,检测出1种菌的样本占38.38%,检出2种和3种病原菌的样本分别占29.24%和19.04%;2种及以上病原菌共存以镰孢菌和腐霉菌、镰孢菌和镰孢菌模式为主。【结论】黄淮海夏玉米主产区茎腐病主要病原菌为拟轮枝镰孢、禾谷镰孢复合种、芒孢腐霉和层出镰孢,其中拟轮枝镰孢为优势种;拟轮枝镰孢在河北省的检出率最高,层出镰孢和芒孢腐霉在山东省的检出率最高,禾谷镰孢复合种在3个省的检出率相近;单个样本中存在多种病原菌共存的模式。
【Objective】The objective of this study is to determine the main pathogens and dominant species of maize stalk rot in the main summer maize producing areas of Huang-Huai-Hai, and to provide a basis for understanding the pathogenic mechanism, resistant breeding, and ultimately maize stalk rot management.【Method】In 2014-2017, 850 samples of maize stalk rot were collected from three provinces(Hebei, Henan and Shandong) in the main summer maize producing areas of Huang-Huai-Hai. The isolated fungi or oomycetes were morphologically characterized and confirmed by using universal and specific primers, those primers were also used for detecting pathogens in diseased tissues. The combined results of isolate identification and tissue molecular detection were analyzed to determine the main pathogens and the dominant species. The detection rates of main pathogens in different provinces and different years of the same province were analyzed to reveal the population dynamics. The detection rates of pathogens in individual samples were also analyzed to determine the coexistence patterns of multiple pathogens.【Result】Fungi or oomycetes were detected in 667 samples, accounting for 78.47% of all samples. The detection rates were different among years, less than 50% in 2014 and more than 90% in 2015-2017. Detected fungi or oomycetes were classified to 46 species of 20 genera. Fusarium spp. had the highest detection rate of 89.96%, including F. graminearum species complex, F. proliferatum, F. verticillioides, F. chlamydosporum, F. subglutinans, F. oxysporum, F. culmorum, F. fujikuroi, F. incarnatum, F. equiseti and F. solani. In addition, Pythium spp. had the second highest detection rate of 34.18%, including P. aristosporum, P. graminicola, P. acanthicum, P. amasculinum and P. oligandrum. The detection rates of four main pathogens, F. verticillioides, F. graminearum species complex, P. aristosporum and F. proliferatum, were 62.07%, 46.93%, 29.09% and 28.04%, respectively. It indicated that the dominant species was F. verticillioides. There were some differences in the detection rates of the four main pathogens among provinces. The detection rate of F. verticillioides in Hebei Province was 73.98%, which was significantly higher than that in Henan and Shandong Provinces, and the detection rates of F. graminearum species complex in the three provinces were similar. The detection rates of F. proliferatum and P. aristosporum in Shandong Province were 35.78% and 34.31%, respectively, which were higher than those in other two provinces. The detection rates of the four main pathogens from the same province were dynamic in different years, and any one of them can be the dominant species. Furthermore, several pathogens could be detected in a single sample. While 38.38% samples were colonized by only one pathogen, 29.24% and 19.04% samples were colonized by two and three pathogens. The patterns of two or multiple pathogens in single samples were mainly coexistence of Fusarium spp. and Pythium spp., or coexistence of Fusarium spp. and Fusarium spp..【Conclusion】The main pathogens of maize stalk rot in the main summer maize producing areas of Huang-Huai-Hai are F. verticillioides, F. graminearum species complex, P. aristosporum and F. proliferatum, and the dominant species is F. verticillioides. The detection rate of F. verticillioides in Hebei Province and that of F. proliferatum and P. aristosporum in Shandong Province are the highest, and the detection rate of F. graminearum species complex in the three provinces is similar. There is a coexistence pattern of two or multiple pathogens in a single sample.
【Objective】The objective of this study is to determine the main pathogens and dominant species of maize stalk rot in the main summer maize producing areas of Huang-Huai-Hai, and to provide a basis for understanding the pathogenic mechanism, resistant breeding, and ultimately maize stalk rot management.【Method】In 2014-2017, 850 samples of maize stalk rot were collected from three provinces(Hebei, Henan and Shandong) in the main summer maize producing areas of Huang-Huai-Hai. The isolated fungi or oomycetes were morphologically characterized and confirmed by using universal and specific primers, those primers were also used for detecting pathogens in diseased tissues. The combined results of isolate identification and tissue molecular detection were analyzed to determine the main pathogens and the dominant species. The detection rates of main pathogens in different provinces and different years of the same province were analyzed to reveal the population dynamics. The detection rates of pathogens in individual samples were also analyzed to determine the coexistence patterns of multiple pathogens.【Result】Fungi or oomycetes were detected in 667 samples, accounting for 78.47% of all samples. The detection rates were different among years, less than 50% in 2014 and more than 90% in 2015-2017. Detected fungi or oomycetes were classified to 46 species of 20 genera. Fusarium spp. had the highest detection rate of 89.96%, including F. graminearum species complex, F. proliferatum, F. verticillioides, F. chlamydosporum, F. subglutinans, F. oxysporum, F. culmorum, F. fujikuroi, F. incarnatum, F. equiseti and F. solani. In addition, Pythium spp. had the second highest detection rate of 34.18%, including P. aristosporum, P. graminicola, P. acanthicum, P. amasculinum and P. oligandrum. The detection rates of four main pathogens, F. verticillioides, F. graminearum species complex, P. aristosporum and F. proliferatum, were 62.07%, 46.93%, 29.09% and 28.04%, respectively. It indicated that the dominant species was F. verticillioides. There were some differences in the detection rates of the four main pathogens among provinces. The detection rate of F. verticillioides in Hebei Province was 73.98%, which was significantly higher than that in Henan and Shandong Provinces, and the detection rates of F. graminearum species complex in the three provinces were similar. The detection rates of F. proliferatum and P. aristosporum in Shandong Province were 35.78% and 34.31%, respectively, which were higher than those in other two provinces. The detection rates of the four main pathogens from the same province were dynamic in different years, and any one of them can be the dominant species. Furthermore, several pathogens could be detected in a single sample. While 38.38% samples were colonized by only one pathogen, 29.24% and 19.04% samples were colonized by two and three pathogens. The patterns of two or multiple pathogens in single samples were mainly coexistence of Fusarium spp. and Pythium spp., or coexistence of Fusarium spp. and Fusarium spp..【Conclusion】The main pathogens of maize stalk rot in the main summer maize producing areas of Huang-Huai-Hai are F. verticillioides, F. graminearum species complex, P. aristosporum and F. proliferatum, and the dominant species is F. verticillioides. The detection rate of F. verticillioides in Hebei Province and that of F. proliferatum and P. aristosporum in Shandong Province are the highest, and the detection rate of F. graminearum species complex in the three provinces is similar. There is a coexistence pattern of two or multiple pathogens in a single sample.
引文
[1]付晋峰,王璞.播期和种植密度对玉米子粒灌浆的影响.玉米科学,2016,24(3):117-122,130.FU J F,WANG P.Effects of sowing date and planting density on maize filling.Journal of Maize Sciences,2016,24(3):117-122,130.(in Chinese)
[2]马传禹,姚丽姗,杜腓利,徐明良.玉米抗茎腐病研究进展.玉米科学,2018,26(2):131-137.MA C Y,YAO L S,DU F L,XU M L.Advanced resistance and genetic research for maize stalk rot.Journal of Maize Sciences,2018,26(2):131-137.(in Chinese)
[3]徐书法,陈捷,高增贵,邹庆道,纪明山,刘海南.中国玉米茎基腐病和穗腐病研究进展.植物病理学报,2006,36(3):193-203.XU S F,CHEN J,GAO Z G,ZOU Q D,JI M S,LIU H N.Maize stalk rot and ear rot in China.Acta Phytopathologica Sinica,2006,36(3):193-203.(in Chinese)
[4]汤蒙蒙.黄淮地区玉米茎基腐病病原菌鉴定及防治技术研究[D].郑州:河南农业大学,2016.TANG M M.Identification of pathogens and the control technology of corn stalk rot in Huanghuai Area[D].Zhengzhou:Henan Agricultural University,2016.(in Chinese)
[5]白金铠,尹志,胡吉成.东北玉米茎腐病病原的研究.植物保护学报,1988,15(2):93-98.BAI J K,YIN Z,HU J C.Studies on pathogenic fungus of stalk rot of corn in the northeastern China.Acta Phytophylacica Snica,1988,15(2):93-98.(in Chinese)
[6]张明智,王守正,王振跃,周挺.河南省玉米青枯病病原菌初步研究.河南农业大学学报,1988,22(2):135-148.ZHANG M Z,WANG S Z,WANG Z Y,ZHOU T.Preliminary study on pathogenic fungi of the Fusarium wilt(Fusarium spp.)of corn in Henan Province.Acta Agriculturae Universitatis Henanensis,1988,22(2):135-148.(in Chinese)
[7]贺娟,何鹏飞,赵正龙,吴毅歆,康志钰,何月秋.云南省玉米茎基腐病病原镰孢菌的种群结构研究.玉米科学,2017,25(4):135-143.HE J,HE P F,ZHAO Z L,WU Y X,KANG Z Y,HE Y Q.Population structure of Fusarium pathogen of corn stalk rot in Yunnan Province.Journal of Maize Sciences,2017,25(4):135-143.(in Chinese)
[8]郭满库,王晓鸣,何苏琴,刘永刚,金社林,曹世勤,魏宏玉.2009年甘肃省玉米穗腐病、茎基腐病的发生危害.植物保护,2011,37(4):134-137.GUO M K,WANG X M,HE S Q,LIU Y G,JIN S L,CAO S Q,WEIH Y.Occurrence of maize kernel rot and corn stalk rot in Gansu in2009.Plant Protection,2011,37(4):134-137.(in Chinese)
[9]GAI X T,YANG R X,PAN X J,YUAN Y,WANG S N,LIANG B B,GAO Z G.First report of Fusarium incarnatum causing stalk rot on maize in China.Plant Disease,2016,100(5):1010.
[10]高卫东,鲍金草,赵晋荣.山西玉米茎腐病病原种类及其复合侵染的研究.山西农业大学学报,1987,7(2):199-207.GAO W D,BAO J C,ZHAO J R.Studies on pathogens and their complex infection of corn stalk rot in Shanxi Province.Journal of Shanxi Agricultural University,1987,7(2):199-207.(in Chinese)
[11]张超冲,贤振华,韦继光,陈超美,周筱兰.玉米青枯病菌的侵染及发病规律研究.广西农学院学报,1983(1):53-62.ZHANG C C,XIAN Z H,WEI J G,CHEN C M,ZHOU X L.Study on regularity of infection and occurrence of pathogenic fungi of corn stalk rot.Journal of Guangxi Agricultural College,1983(1):53-62.(in Chinese)
[12]王晓鸣,吴全安,刘晓娟,马国忠.寄生玉米的6种腐霉及其致病性研究.植物病理学报,1994,24(4):343-346.WANG X M,WU Q A,LIU X J,MA G Z.Identification and pathogenicity of Pythium spp.isolated from maize.Acta Phytopathologica Sinica,1994,24(4):343-346.(in Chinese)
[13]罗畔池,张成和,刘爱国,王月恋,张小青,孔令晓,霍志清,刘庆义.玉米茎腐病原及栽培与发病关系.华北农学报,1993,8(增刊):110-114.LUO P C,ZHANG C H,LIU A G,WANG Y L,ZHANG X Q,KONGL X,HUO Z Q,LIU Q Y.Studies on the pathogen of maize stalk rot and the relationship between maize cultivation and development of the disease.Acta Agriculturae Boreali-Sinica,1993,8(Suppl.):110-114.(in Chinese)
[14]王连生,刘克明,刘玉瑛,苏海,朱小阳,吴全安.河北省玉米青枯病病原菌的分离及致病性测定.河北农业科学,1992(1):1-4.WANG L S,LIU K M,LIU Y Y,SU H,ZHU X Y,WU Q A.Isolation and pathogenicity determination of pathogen for corn stalk rot in Hebei Province.Journal of Hebei Agricultural Sciences,1992(1):1-4.(in Chinese)
[15]石洁.玉米镰刀菌型茎腐、穗腐、苗期根腐病的相互关系及防治[D].保定:河北农业大学,2002.SHI J.Relationship and control of Fusarium stalk rot,ear rot and seedling root rot in maize[D].Baoding:Hebei Agricultural University,2002.(in Chinese)
[16]徐作珽,张传模.山东玉米茎基腐病病原菌的初步研究.植物病理学报,1985,15(2):103-108.XU Z T,ZHANG C M.Study on the causal organism of root and basal stalk rot of corn in Shandong Province.Acta Phytopathologica Sinica,1985,15(2):103-108.(in Chinese)
[17]张艺铭.山东省玉米青枯病病原鉴定及生防菌株和化学药剂的筛选[D].泰安:山东农业大学,2016.ZHANG Y M.Pathogen identification of corn stalk rot in Shandong and screening of antagonistic bacteria and effective agrochemicals[D].Taian:Shandong Agricultural University,2016.(in Chinese)
[18]袁虹霞,闵营辉,张丹丹,邢小萍,李洪连,雷彩艳.河南省玉米茎腐病病原菌分离与致病性测定.玉米科学,2011,19(6):122-124,128.YUAN H X,MIN Y H,ZHANG D D,XING X P,LI H L,LEI C Y.Isolation and pathogenicity determination of pathogen for corn stalk rot in Henan Province.Journal of Maize Sciences,2011,19(6):122-124,128.(in Chinese)
[19]崔小伟.河南省玉米茎腐病研究[D].杨凌:西北农林科技大学,2013.CUI X W.Study on corn stalk rot in Henan Province[D].Yangling:Northwest A&F University,2013.(in Chinese)
[20]孙静,谢淑娜,刘佳中,刘京宝,郝俊杰,邓士政.河南省玉米茎基部镰刀菌的形态和分子鉴定.植物病理学报,2014,44(1):8-16.SUN J,XIE S N,LIU J Z,LIU J B,HAO J J,DENG S Z.Morphological and molecular identification of Fusarium isolated from basal stalks of maize in Henan Province.Acta Phytopathologica Sinica,2014,44(1):8-16.(in Chinese)
[21]范志业,崔小伟,施艳,陈琦,刘迪,侯艳红,李世民,闫海霞,袁刘正,孙虎.河南省玉米茎基腐病主要病原菌鉴定及主栽玉米品种的抗性分析.河南农业科学,2014,43(12):87-90.FAN Z Y,CUI X W,SHI Y,CHEN Q,LIU D,HOU Y H,LI S M,YAN H X,YUAN L Z,SUN H.Identification of pathogens causing corn stalk rot and resistance test of main cultivars in Henan Province.Journal of Henan Agricultural Sciences,2014,43(12):87-90.(in Chinese)
[22]何婧.玉米茎腐病病原菌分离鉴定与镰孢菌群体遗传多样性研究[D].乌鲁木齐:新疆农业大学,2010.HE J.Isolation,identification of causal agents of maize stalk rot and genetic diversity analysis of Fusarium spp.[D].Urumqi:Xinjiang Agricultural University,2010.(in Chinese)
[23]马红霞,张海剑,孙华,石洁,陈丹,郭宁.玉米茎腐病病原菌检测方法研究.植物保护,2017,43(3):149-153.MA H X,ZHANG H J,SUN H,SHI J,CHEN D,GUO N.Comparison of pathogen detection methods for corn stalk rot.Plant Protection,2017,43(3):149-153.(in Chinese)
[24]马秉元,李亚玲,段双科.陕西省关中地区玉米青枯病病原菌及其致病性的研究.植物病理学报,1985,15(3):150-152.MA B Y,LI Y L,DUAN S K.Study on pathogen of corn stalk rot and its pathogenicity in middle Shaanxi.Acta Phytopathologica Sinica,1985,15(3):150-152.(in Chinese)
[25]吴全安,梁克恭,朱小阳,王晓鸣,金加同,王桂跃.北京和浙江地区玉米青枯病病原菌的分离与鉴定.中国农业科学,1989,22(5):71-75.WU Q A,LIANG K G,ZHU X Y,WANG X M,JING J T,WANG G Y.Isolation and identification of the pathogen of maize stalk rot in Beijing and Zhejiang.Scientia Agricultura Sinica,1989,22(5):71-75.(in Chinese)
[26]魏景超.真菌鉴定手册.上海:上海科学技术出版社,1979.WEI J C.Handbook of Fungus Identification.Shanghai:Shanghai Scientific and Technical Publishers,1979.(in Chinese)
[27]孔华忠.中国真菌志(第35卷):青霉属及其相关有性型属.北京:科学出版社,2007.KONG H Z.Chinese Mycology(Volumn 35):The Penicillia and their Related Teleomorphs.Beijing:Science Press,2007.(in Chinese)
[28]孙华,郭宁,石洁,张海剑,马红霞,刘树森.海南玉米穗腐病病原菌分离鉴定及优势种的遗传多样性分析.植物病理学报,2017,47(5):577-583.SUN H,GUO N,SHI J,ZHANG H J,MA H X,LIU S S.Characterization of the maize ear rot pathogens and genetic diversity analysis of dominant species in Hainan.Acta Phytopathologica Sinica,2017,47(5):577-583.(in Chinese)
[29]NICHOLSON P,SIMPSON D R,WESTON G,REZANOOR H N,LEES A K,PARRY D W,JOYCE D.Detection and quantification of Fusarium culmorum and Fusarium graminearum in cereals using PCR assays.Physiological and Molecular Plant Pathology,1998,53(1):17-37.
[30]MULèG,SUSCA A,STEA G,MORETTI A.A species-specific PCR assay based on the calmodulin partial gene for identification of Fusarium verticillioides,F.proliferatum and F.subglutinans.European Journal of Plant Pathology,2004,110(5/6):495-502.
[31]RAHJOO V,ZAD J,JAVAN-NIKKHAH M,GOHARI A M,OKHOVVAT S M,BIHAMTA M R,RAZZAGHIAN J,KLEMSDALS S.Morphological and molecular identification of Fusarium isolated from maize ears in Iran.Journal of Plant Pathology,2008,90(3):463-468.
[32]MISHRA P K,FOX R T,CULHAM A.Development of a PCR-based assay for rapid and reliable identification of pathogenic Fusaria.FEMS Microbiology Letters,2003,218(2):329-332.
[33]BONANTS P,WEERDT M H,VAN GENT-PELZER M,LACOURTI,COOKE D,DUNCAN J.Detection and identification of Phytophthora fragariae Hickman by the polymerase chain reaction.European Journal of Plant Pathology,1997,103(4):345-355.
[34]O’DONNELL K,KISTLER H C,CIGELNIK E,PLOETZ R C.Multiple evolutionary origins of the fungus causing Panama disease of banana:concordant evidence from nuclear and mitochondrial gene genealogies.Proceedings of the National Academy of Sciences of the United States of America,1998,95(5):2044-2049.
[35]WHITE T J,BRUNS T,LEE S,TAYLOR J.Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics//INNISM A,GELFAND D H,SNINSKY J J,WHITE T J.PCR Protocols:a Guide to Methods and Applications.San Diego:Academic Press,1990:315-322.
[36]CARBONE I,KOHN L M.A method for designing primer sets for speciation studies in filamentous ascomycetes.Mycologia,1999,91(3):553-556.
[37]邱法展.玉米单倍体育种及苗期耐渍性研究[D].武汉:华中农业大学,2007.QIU F Z.Study on waterlogging tolerance during seedling stage and haploid breeding in maize[D].Wuhan:Huazhong Agricultural University,2007.(in Chinese)
[38]全国农业技术推广服务中心.中国玉米新品种动态2014年国家级玉米品种区试报告.北京:中国农业科学技术出版社,2015:106.The National Agro-tech Extension and Service Center.China’s Corn Varieties Dynamic-2014 National Corn Variety Regional Test Report.Beijing:China Agricultural Science and Technology Press,2015:106.(in Chinese)
[39]张瑞英,张坪.黑龙江省玉米茎基腐病病原菌研究初报.植物保护学报,1993,20(3):287-288.ZHANG R Y,ZHANG P.The preliminary study on corn stalk base rot in Heilongjiang Province.Acta Phytophylacica Sinica,1993,20(3):287-288.(in Chinese)
[40]许明学,冯芬芬,柳迎春.玉米抗茎腐病育种的研究.玉米科学,1993,1(3):67-70.XU M X,FENG F F,LIU Y C.Research progress on maize breeding for resistance to stalk rot.Maize Science,1993,1(3):67-70.(in Chinese)
[41]吕国忠,赵志慧,孙晓东,姜子德,杨红.串珠镰孢菌种名的废弃及其与腾仓赤霉复合种的关系.菌物学报,2010,29(1):143-151.LüG Z,ZHAO Z H,SUN X D,JIANG Z D,YANG H.The abandonment of the name Fusarium moniliforme and the concept of Gibberella fujikuroi species complex.Mycosystema,2010,29(1):143-151.(in Chinese)
[42]任旭,朱振东,李洪杰,段灿星,王晓鸣.轮枝镰孢SSR标记开发及在玉米分离群体遗传多样性分析中的应用.中国农业科学,2012,45(1):52-66.REN X,ZHU Z D,LI H J,DUAN C X,WANG X M.SSR marker development and analysis of genetic diversity of Fusarium verticillioides isolated from maize in china.Scientia Agricultura Sinica,2012,45(1):52-66.(in Chinese)
[43]朱华,梁继农,王彰明,陈厚德,奚京平,仇宝华,王全领.江苏省玉米茎腐病菌种类鉴定.植物保护学报,1997,24(1):49-54.ZHU H,LIANG J N,WANG Z M,CHEN H D,XI J P,QIU B H,WANG Q L.Identification of pathogens causing maize stalk rot in Jiangsu Province.Acta Phytophylacica Sinica,1997,24(1):49-54.(in Chinese)
[44]王富荣,傅玉红,陈永欣.山西省玉米茎腐病病原菌的分离及致病性测定.山西农业科学,1992(9):20-21.WANG F R,FU Y H,CHEN Y X.Isolation and pathogenicity determination of pathogen for corn stalk rot in Shanxi Province Journal of Shanxi Agricultural Sciences,1992(9):20-21.(in Chinese)
[45]宋佐衡,梁景颐,白金铠.辽宁省玉米茎腐病病原菌的研究.沈阳农业大学学报,1990,21(3):214-218.SONG Z H,LIANG J Y,BAI J K.Preliminary studies on the etiology of stalk rot of corn in Liaoning Province.Journal of Shenyang Agricultural University,1990,21(3):214-218.(in Chinese)
[46]杨山山,郝彦俊,邱荣芳,王晓鸣,吴全安.新疆玉米青枯病病原菌分离和鉴定.新疆农业大学学报,1997,20(2):29-36.YANG S S,HAO Y J,QIU R F,WANG X M,WU Q A.Etiology of corn stalk rot in Xinjiang.Journal of Xinjiang Agricultural University,1997,20(2):29-36.(in Chinese)
[47]吴小龙.玉米青枯病菌的鉴定、生物学特性及生物防治研究[D].雅安:四川农业大学,2010.WU X L.Isolation and identification and biological characteristics of pathogen of corn stalk rot and primary study on diseases biological control[D].Yaan:Sichuan Agricultural University,2010.(in Chinese)
[48]孙秀华,孙亚杰,张春山,白金铠,宋佐衡,陈捷.玉米茎腐病病原菌相互作用研究.沈阳农业大学学报,1992,23(2):93-96.SUN X H,SUN Y J,ZHANG C S,BAI J K,SONG Z H,CHEN J.Studies on the relationship among pathogens of corn stalk rot.Journal of Shenyang Agricultural University,1992,23(2):93-96.(in Chinese)
[49]陈绍江,宋同明,吴全安.玉米青枯病病原腐霉对其伴生镰刀菌的影响.植物病理学报,1997,27(3):251-256.CHEN S J,SONG T M,WU A Q.Effect of Pythium on the growth of Fusarium in maize stalk rot.Acta Phytopathologica Sinica,1997,27(3):251-256.(in Chinese)
[2]马传禹,姚丽姗,杜腓利,徐明良.玉米抗茎腐病研究进展.玉米科学,2018,26(2):131-137.MA C Y,YAO L S,DU F L,XU M L.Advanced resistance and genetic research for maize stalk rot.Journal of Maize Sciences,2018,26(2):131-137.(in Chinese)
[3]徐书法,陈捷,高增贵,邹庆道,纪明山,刘海南.中国玉米茎基腐病和穗腐病研究进展.植物病理学报,2006,36(3):193-203.XU S F,CHEN J,GAO Z G,ZOU Q D,JI M S,LIU H N.Maize stalk rot and ear rot in China.Acta Phytopathologica Sinica,2006,36(3):193-203.(in Chinese)
[4]汤蒙蒙.黄淮地区玉米茎基腐病病原菌鉴定及防治技术研究[D].郑州:河南农业大学,2016.TANG M M.Identification of pathogens and the control technology of corn stalk rot in Huanghuai Area[D].Zhengzhou:Henan Agricultural University,2016.(in Chinese)
[5]白金铠,尹志,胡吉成.东北玉米茎腐病病原的研究.植物保护学报,1988,15(2):93-98.BAI J K,YIN Z,HU J C.Studies on pathogenic fungus of stalk rot of corn in the northeastern China.Acta Phytophylacica Snica,1988,15(2):93-98.(in Chinese)
[6]张明智,王守正,王振跃,周挺.河南省玉米青枯病病原菌初步研究.河南农业大学学报,1988,22(2):135-148.ZHANG M Z,WANG S Z,WANG Z Y,ZHOU T.Preliminary study on pathogenic fungi of the Fusarium wilt(Fusarium spp.)of corn in Henan Province.Acta Agriculturae Universitatis Henanensis,1988,22(2):135-148.(in Chinese)
[7]贺娟,何鹏飞,赵正龙,吴毅歆,康志钰,何月秋.云南省玉米茎基腐病病原镰孢菌的种群结构研究.玉米科学,2017,25(4):135-143.HE J,HE P F,ZHAO Z L,WU Y X,KANG Z Y,HE Y Q.Population structure of Fusarium pathogen of corn stalk rot in Yunnan Province.Journal of Maize Sciences,2017,25(4):135-143.(in Chinese)
[8]郭满库,王晓鸣,何苏琴,刘永刚,金社林,曹世勤,魏宏玉.2009年甘肃省玉米穗腐病、茎基腐病的发生危害.植物保护,2011,37(4):134-137.GUO M K,WANG X M,HE S Q,LIU Y G,JIN S L,CAO S Q,WEIH Y.Occurrence of maize kernel rot and corn stalk rot in Gansu in2009.Plant Protection,2011,37(4):134-137.(in Chinese)
[9]GAI X T,YANG R X,PAN X J,YUAN Y,WANG S N,LIANG B B,GAO Z G.First report of Fusarium incarnatum causing stalk rot on maize in China.Plant Disease,2016,100(5):1010.
[10]高卫东,鲍金草,赵晋荣.山西玉米茎腐病病原种类及其复合侵染的研究.山西农业大学学报,1987,7(2):199-207.GAO W D,BAO J C,ZHAO J R.Studies on pathogens and their complex infection of corn stalk rot in Shanxi Province.Journal of Shanxi Agricultural University,1987,7(2):199-207.(in Chinese)
[11]张超冲,贤振华,韦继光,陈超美,周筱兰.玉米青枯病菌的侵染及发病规律研究.广西农学院学报,1983(1):53-62.ZHANG C C,XIAN Z H,WEI J G,CHEN C M,ZHOU X L.Study on regularity of infection and occurrence of pathogenic fungi of corn stalk rot.Journal of Guangxi Agricultural College,1983(1):53-62.(in Chinese)
[12]王晓鸣,吴全安,刘晓娟,马国忠.寄生玉米的6种腐霉及其致病性研究.植物病理学报,1994,24(4):343-346.WANG X M,WU Q A,LIU X J,MA G Z.Identification and pathogenicity of Pythium spp.isolated from maize.Acta Phytopathologica Sinica,1994,24(4):343-346.(in Chinese)
[13]罗畔池,张成和,刘爱国,王月恋,张小青,孔令晓,霍志清,刘庆义.玉米茎腐病原及栽培与发病关系.华北农学报,1993,8(增刊):110-114.LUO P C,ZHANG C H,LIU A G,WANG Y L,ZHANG X Q,KONGL X,HUO Z Q,LIU Q Y.Studies on the pathogen of maize stalk rot and the relationship between maize cultivation and development of the disease.Acta Agriculturae Boreali-Sinica,1993,8(Suppl.):110-114.(in Chinese)
[14]王连生,刘克明,刘玉瑛,苏海,朱小阳,吴全安.河北省玉米青枯病病原菌的分离及致病性测定.河北农业科学,1992(1):1-4.WANG L S,LIU K M,LIU Y Y,SU H,ZHU X Y,WU Q A.Isolation and pathogenicity determination of pathogen for corn stalk rot in Hebei Province.Journal of Hebei Agricultural Sciences,1992(1):1-4.(in Chinese)
[15]石洁.玉米镰刀菌型茎腐、穗腐、苗期根腐病的相互关系及防治[D].保定:河北农业大学,2002.SHI J.Relationship and control of Fusarium stalk rot,ear rot and seedling root rot in maize[D].Baoding:Hebei Agricultural University,2002.(in Chinese)
[16]徐作珽,张传模.山东玉米茎基腐病病原菌的初步研究.植物病理学报,1985,15(2):103-108.XU Z T,ZHANG C M.Study on the causal organism of root and basal stalk rot of corn in Shandong Province.Acta Phytopathologica Sinica,1985,15(2):103-108.(in Chinese)
[17]张艺铭.山东省玉米青枯病病原鉴定及生防菌株和化学药剂的筛选[D].泰安:山东农业大学,2016.ZHANG Y M.Pathogen identification of corn stalk rot in Shandong and screening of antagonistic bacteria and effective agrochemicals[D].Taian:Shandong Agricultural University,2016.(in Chinese)
[18]袁虹霞,闵营辉,张丹丹,邢小萍,李洪连,雷彩艳.河南省玉米茎腐病病原菌分离与致病性测定.玉米科学,2011,19(6):122-124,128.YUAN H X,MIN Y H,ZHANG D D,XING X P,LI H L,LEI C Y.Isolation and pathogenicity determination of pathogen for corn stalk rot in Henan Province.Journal of Maize Sciences,2011,19(6):122-124,128.(in Chinese)
[19]崔小伟.河南省玉米茎腐病研究[D].杨凌:西北农林科技大学,2013.CUI X W.Study on corn stalk rot in Henan Province[D].Yangling:Northwest A&F University,2013.(in Chinese)
[20]孙静,谢淑娜,刘佳中,刘京宝,郝俊杰,邓士政.河南省玉米茎基部镰刀菌的形态和分子鉴定.植物病理学报,2014,44(1):8-16.SUN J,XIE S N,LIU J Z,LIU J B,HAO J J,DENG S Z.Morphological and molecular identification of Fusarium isolated from basal stalks of maize in Henan Province.Acta Phytopathologica Sinica,2014,44(1):8-16.(in Chinese)
[21]范志业,崔小伟,施艳,陈琦,刘迪,侯艳红,李世民,闫海霞,袁刘正,孙虎.河南省玉米茎基腐病主要病原菌鉴定及主栽玉米品种的抗性分析.河南农业科学,2014,43(12):87-90.FAN Z Y,CUI X W,SHI Y,CHEN Q,LIU D,HOU Y H,LI S M,YAN H X,YUAN L Z,SUN H.Identification of pathogens causing corn stalk rot and resistance test of main cultivars in Henan Province.Journal of Henan Agricultural Sciences,2014,43(12):87-90.(in Chinese)
[22]何婧.玉米茎腐病病原菌分离鉴定与镰孢菌群体遗传多样性研究[D].乌鲁木齐:新疆农业大学,2010.HE J.Isolation,identification of causal agents of maize stalk rot and genetic diversity analysis of Fusarium spp.[D].Urumqi:Xinjiang Agricultural University,2010.(in Chinese)
[23]马红霞,张海剑,孙华,石洁,陈丹,郭宁.玉米茎腐病病原菌检测方法研究.植物保护,2017,43(3):149-153.MA H X,ZHANG H J,SUN H,SHI J,CHEN D,GUO N.Comparison of pathogen detection methods for corn stalk rot.Plant Protection,2017,43(3):149-153.(in Chinese)
[24]马秉元,李亚玲,段双科.陕西省关中地区玉米青枯病病原菌及其致病性的研究.植物病理学报,1985,15(3):150-152.MA B Y,LI Y L,DUAN S K.Study on pathogen of corn stalk rot and its pathogenicity in middle Shaanxi.Acta Phytopathologica Sinica,1985,15(3):150-152.(in Chinese)
[25]吴全安,梁克恭,朱小阳,王晓鸣,金加同,王桂跃.北京和浙江地区玉米青枯病病原菌的分离与鉴定.中国农业科学,1989,22(5):71-75.WU Q A,LIANG K G,ZHU X Y,WANG X M,JING J T,WANG G Y.Isolation and identification of the pathogen of maize stalk rot in Beijing and Zhejiang.Scientia Agricultura Sinica,1989,22(5):71-75.(in Chinese)
[26]魏景超.真菌鉴定手册.上海:上海科学技术出版社,1979.WEI J C.Handbook of Fungus Identification.Shanghai:Shanghai Scientific and Technical Publishers,1979.(in Chinese)
[27]孔华忠.中国真菌志(第35卷):青霉属及其相关有性型属.北京:科学出版社,2007.KONG H Z.Chinese Mycology(Volumn 35):The Penicillia and their Related Teleomorphs.Beijing:Science Press,2007.(in Chinese)
[28]孙华,郭宁,石洁,张海剑,马红霞,刘树森.海南玉米穗腐病病原菌分离鉴定及优势种的遗传多样性分析.植物病理学报,2017,47(5):577-583.SUN H,GUO N,SHI J,ZHANG H J,MA H X,LIU S S.Characterization of the maize ear rot pathogens and genetic diversity analysis of dominant species in Hainan.Acta Phytopathologica Sinica,2017,47(5):577-583.(in Chinese)
[29]NICHOLSON P,SIMPSON D R,WESTON G,REZANOOR H N,LEES A K,PARRY D W,JOYCE D.Detection and quantification of Fusarium culmorum and Fusarium graminearum in cereals using PCR assays.Physiological and Molecular Plant Pathology,1998,53(1):17-37.
[30]MULèG,SUSCA A,STEA G,MORETTI A.A species-specific PCR assay based on the calmodulin partial gene for identification of Fusarium verticillioides,F.proliferatum and F.subglutinans.European Journal of Plant Pathology,2004,110(5/6):495-502.
[31]RAHJOO V,ZAD J,JAVAN-NIKKHAH M,GOHARI A M,OKHOVVAT S M,BIHAMTA M R,RAZZAGHIAN J,KLEMSDALS S.Morphological and molecular identification of Fusarium isolated from maize ears in Iran.Journal of Plant Pathology,2008,90(3):463-468.
[32]MISHRA P K,FOX R T,CULHAM A.Development of a PCR-based assay for rapid and reliable identification of pathogenic Fusaria.FEMS Microbiology Letters,2003,218(2):329-332.
[33]BONANTS P,WEERDT M H,VAN GENT-PELZER M,LACOURTI,COOKE D,DUNCAN J.Detection and identification of Phytophthora fragariae Hickman by the polymerase chain reaction.European Journal of Plant Pathology,1997,103(4):345-355.
[34]O’DONNELL K,KISTLER H C,CIGELNIK E,PLOETZ R C.Multiple evolutionary origins of the fungus causing Panama disease of banana:concordant evidence from nuclear and mitochondrial gene genealogies.Proceedings of the National Academy of Sciences of the United States of America,1998,95(5):2044-2049.
[35]WHITE T J,BRUNS T,LEE S,TAYLOR J.Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics//INNISM A,GELFAND D H,SNINSKY J J,WHITE T J.PCR Protocols:a Guide to Methods and Applications.San Diego:Academic Press,1990:315-322.
[36]CARBONE I,KOHN L M.A method for designing primer sets for speciation studies in filamentous ascomycetes.Mycologia,1999,91(3):553-556.
[37]邱法展.玉米单倍体育种及苗期耐渍性研究[D].武汉:华中农业大学,2007.QIU F Z.Study on waterlogging tolerance during seedling stage and haploid breeding in maize[D].Wuhan:Huazhong Agricultural University,2007.(in Chinese)
[38]全国农业技术推广服务中心.中国玉米新品种动态2014年国家级玉米品种区试报告.北京:中国农业科学技术出版社,2015:106.The National Agro-tech Extension and Service Center.China’s Corn Varieties Dynamic-2014 National Corn Variety Regional Test Report.Beijing:China Agricultural Science and Technology Press,2015:106.(in Chinese)
[39]张瑞英,张坪.黑龙江省玉米茎基腐病病原菌研究初报.植物保护学报,1993,20(3):287-288.ZHANG R Y,ZHANG P.The preliminary study on corn stalk base rot in Heilongjiang Province.Acta Phytophylacica Sinica,1993,20(3):287-288.(in Chinese)
[40]许明学,冯芬芬,柳迎春.玉米抗茎腐病育种的研究.玉米科学,1993,1(3):67-70.XU M X,FENG F F,LIU Y C.Research progress on maize breeding for resistance to stalk rot.Maize Science,1993,1(3):67-70.(in Chinese)
[41]吕国忠,赵志慧,孙晓东,姜子德,杨红.串珠镰孢菌种名的废弃及其与腾仓赤霉复合种的关系.菌物学报,2010,29(1):143-151.LüG Z,ZHAO Z H,SUN X D,JIANG Z D,YANG H.The abandonment of the name Fusarium moniliforme and the concept of Gibberella fujikuroi species complex.Mycosystema,2010,29(1):143-151.(in Chinese)
[42]任旭,朱振东,李洪杰,段灿星,王晓鸣.轮枝镰孢SSR标记开发及在玉米分离群体遗传多样性分析中的应用.中国农业科学,2012,45(1):52-66.REN X,ZHU Z D,LI H J,DUAN C X,WANG X M.SSR marker development and analysis of genetic diversity of Fusarium verticillioides isolated from maize in china.Scientia Agricultura Sinica,2012,45(1):52-66.(in Chinese)
[43]朱华,梁继农,王彰明,陈厚德,奚京平,仇宝华,王全领.江苏省玉米茎腐病菌种类鉴定.植物保护学报,1997,24(1):49-54.ZHU H,LIANG J N,WANG Z M,CHEN H D,XI J P,QIU B H,WANG Q L.Identification of pathogens causing maize stalk rot in Jiangsu Province.Acta Phytophylacica Sinica,1997,24(1):49-54.(in Chinese)
[44]王富荣,傅玉红,陈永欣.山西省玉米茎腐病病原菌的分离及致病性测定.山西农业科学,1992(9):20-21.WANG F R,FU Y H,CHEN Y X.Isolation and pathogenicity determination of pathogen for corn stalk rot in Shanxi Province Journal of Shanxi Agricultural Sciences,1992(9):20-21.(in Chinese)
[45]宋佐衡,梁景颐,白金铠.辽宁省玉米茎腐病病原菌的研究.沈阳农业大学学报,1990,21(3):214-218.SONG Z H,LIANG J Y,BAI J K.Preliminary studies on the etiology of stalk rot of corn in Liaoning Province.Journal of Shenyang Agricultural University,1990,21(3):214-218.(in Chinese)
[46]杨山山,郝彦俊,邱荣芳,王晓鸣,吴全安.新疆玉米青枯病病原菌分离和鉴定.新疆农业大学学报,1997,20(2):29-36.YANG S S,HAO Y J,QIU R F,WANG X M,WU Q A.Etiology of corn stalk rot in Xinjiang.Journal of Xinjiang Agricultural University,1997,20(2):29-36.(in Chinese)
[47]吴小龙.玉米青枯病菌的鉴定、生物学特性及生物防治研究[D].雅安:四川农业大学,2010.WU X L.Isolation and identification and biological characteristics of pathogen of corn stalk rot and primary study on diseases biological control[D].Yaan:Sichuan Agricultural University,2010.(in Chinese)
[48]孙秀华,孙亚杰,张春山,白金铠,宋佐衡,陈捷.玉米茎腐病病原菌相互作用研究.沈阳农业大学学报,1992,23(2):93-96.SUN X H,SUN Y J,ZHANG C S,BAI J K,SONG Z H,CHEN J.Studies on the relationship among pathogens of corn stalk rot.Journal of Shenyang Agricultural University,1992,23(2):93-96.(in Chinese)
[49]陈绍江,宋同明,吴全安.玉米青枯病病原腐霉对其伴生镰刀菌的影响.植物病理学报,1997,27(3):251-256.CHEN S J,SONG T M,WU A Q.Effect of Pythium on the growth of Fusarium in maize stalk rot.Acta Phytopathologica Sinica,1997,27(3):251-256.(in Chinese)