尖角突脐孢菌侵染稗草的机制及产生毒素的初步研究
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摘要
我们从中国农业大学杂草室获得了17株特异感染稗草(Echinochloa crus-galli)的病原菌，该菌是从13个不同生态区域的稗草植株的叶片病斑上分离得到的，经鉴定为尖角突脐孢菌(Exserohilum monoceras)。本论文旨在为其开发成新型生物农药—微生物除草剂作致病机理和应用前景的基础研究，主要研究了其侵染过程、致病性和致病毒素。我们通过对其侵染规律和无孢子悬液对稗草毒害的测定，提出尖角突脐孢产生毒素和有开发为毒素除草剂的可能；并选择生物测定的合适方法，从17个菌株中筛选出合适的产生毒素的菌株32、X27、MC9；进一步选用菌株32研究产生毒素的最佳条件，这些为毒素的制备和机理研究提供了基础。
研究表明尖角突脐孢是稗草的强致病菌。该菌不仅能从稗草叶片的气孔或伤口侵入，而且能直接穿透叶片表皮细胞侵入叶肉细胞，接种后24—26小时完成侵染过程。被侵染后的稗草植株叶肉细胞膜结构发生变形，叶绿体迅速失绿，后期膜被破坏，提示尖角突脐孢在侵染过程中有可能分泌毒素，并作用于细胞膜。
对17株菌株的生测结果表明，种子萌发抑制法和根生长抑制法是比较理想的生测方法，而离体叶片检测会得到比较直观的结果，但不能定量，根冠细胞测定法，喷雾法不适用于检测毒素的致病性。
用1×10~6／ml孢子浓度处理稗草植株，14天后调查各菌株对稗草植株的抑制情况，得出尖角突脐孢的强致病菌株是X27、MC9和32。对17株菌滤液的生物测定结果综合分析，这17株菌都或多或少的产生致病毒素，以MC9、32、X27、PX52、PX2最为有效，说明分生孢子的致病性和毒素的作用之间有正相关性，综合分析，我们选定进一步研究的适宜菌株为活力最高的MC9、32、X27。
选用尖角突脐孢菌32，研究了其产生毒素的最优条件为：在土豆液体培养基上，起始pH5.5—6，接种量10~4个孢子，培养温度28℃，摇床培养200rpm，培养时间7天。我们分析了毒素产生条件与菌和孢子培养
条件的相关性，说明毒素的产生是与菌株的生长活性相一致的，在一定
程度上有量的积累，但二者之间也有差异。
Seventeen strains of Exserohilum monoceras isolated from the diseased leaves of Echinochloa curs-galli occurring in thirteen different places were gained. The objects of this study were to determine its infection process, pathological characters and toxins. We observed the infection process and lesions on leaves of E. crus-galli damaged by filtrate of E. monoceras, and concluded it may produce toxins; we compared the bioassay methods to select the strains 32, X27 and MC9 which could produce vast toxins; We determined conditions of toxins production of strain 32.
The infection process showed that E. monoceras had high pathological action on E. crus-galli. The hypha began to form from the base point of the conidia 8 hours after inoculation. Penetration into E. crus-galli leaves by the hypha occurred through the stomata or randomized places 13 hours after inoculation. The whole infection process finished within 48 to 72 hours.
We compared the different bioassay methods and found that the root and shoot growth control experiment was most optional. Strain 32, MC9, X27 had the highest control rate.
For toxin production, the optimal conditions were: PGB liquid media, initial pH 5.5-6.0, initial inoculation 104 conidia/ml, temperature 28癈, 200rpm, seven days. The toxin productions was related to the conditions of mycelium growth.
研究表明尖角突脐孢是稗草的强致病菌。该菌不仅能从稗草叶片的气孔或伤口侵入，而且能直接穿透叶片表皮细胞侵入叶肉细胞，接种后24—26小时完成侵染过程。被侵染后的稗草植株叶肉细胞膜结构发生变形，叶绿体迅速失绿，后期膜被破坏，提示尖角突脐孢在侵染过程中有可能分泌毒素，并作用于细胞膜。
对17株菌株的生测结果表明，种子萌发抑制法和根生长抑制法是比较理想的生测方法，而离体叶片检测会得到比较直观的结果，但不能定量，根冠细胞测定法，喷雾法不适用于检测毒素的致病性。
用1×10~6／ml孢子浓度处理稗草植株，14天后调查各菌株对稗草植株的抑制情况，得出尖角突脐孢的强致病菌株是X27、MC9和32。对17株菌滤液的生物测定结果综合分析，这17株菌都或多或少的产生致病毒素，以MC9、32、X27、PX52、PX2最为有效，说明分生孢子的致病性和毒素的作用之间有正相关性，综合分析，我们选定进一步研究的适宜菌株为活力最高的MC9、32、X27。
选用尖角突脐孢菌32，研究了其产生毒素的最优条件为：在土豆液体培养基上，起始pH5.5—6，接种量10~4个孢子，培养温度28℃，摇床培养200rpm，培养时间7天。我们分析了毒素产生条件与菌和孢子培养
条件的相关性，说明毒素的产生是与菌株的生长活性相一致的，在一定
程度上有量的积累，但二者之间也有差异。
Seventeen strains of Exserohilum monoceras isolated from the diseased leaves of Echinochloa curs-galli occurring in thirteen different places were gained. The objects of this study were to determine its infection process, pathological characters and toxins. We observed the infection process and lesions on leaves of E. crus-galli damaged by filtrate of E. monoceras, and concluded it may produce toxins; we compared the bioassay methods to select the strains 32, X27 and MC9 which could produce vast toxins; We determined conditions of toxins production of strain 32.
The infection process showed that E. monoceras had high pathological action on E. crus-galli. The hypha began to form from the base point of the conidia 8 hours after inoculation. Penetration into E. crus-galli leaves by the hypha occurred through the stomata or randomized places 13 hours after inoculation. The whole infection process finished within 48 to 72 hours.
We compared the different bioassay methods and found that the root and shoot growth control experiment was most optional. Strain 32, MC9, X27 had the highest control rate.
For toxin production, the optimal conditions were: PGB liquid media, initial pH 5.5-6.0, initial inoculation 104 conidia/ml, temperature 28癈, 200rpm, seven days. The toxin productions was related to the conditions of mycelium growth.
引文
1．陈华林摘译，利用微生物制剂控制杂草，世界农业，1997，12，27-29，原载[英]Hort Science 1997，32(2)，201—205
2．陈涛主编，有害生物的微生物防治原理和技术，湖北科学技术发展出版社，1997，408-448
3．陈勇，尖角突脐孢菌作为真菌除草剂防除稗草潜力的研究，中国农业大学博士学位论文，2001
4．陈勇强等，国外微生物除草剂的研究及应用现状，天津农业科学，1998 4(20)，5-9
5．丁建清，农田杂草的生物防治，中国生物防治，1995，11(3)，129—123
6．董金皋、王江柱，植物病原真菌毒素活性测定方法，董金皋、李树正主编，植物病原菌毒素研究进展，1997，第一卷，中国科学技术出版社，北京，61—74
7．董金皋、黄梧芳，玉米叶斑病菌毒素的研究现状，董金皋、李树正主编，植物病原菌毒素研究进展，1997，第一卷，中国科学技术出版社，北京，102—126
8．高昭远等，我国利用植物病原微生物防除杂草研究的回顾与现状，1995，中国农学通报，11(1)，31-34
9．菅原二三男，利用植物病原微生物开发除草剂，农药译丛，1992，14(3)，40—45
10．郭凤根、李扬汉、邓福珍，园林菟丝子生防真菌的筛选研究，中国生物防治，1998，14(4)，159-162
11．黄世文、Alan K.Watson、段桂芳、余柳青，利用稗草病原菌开发微生物除草剂的研究，喻子牛等主编，微生物农药及其产业化，2000，科学出版社，260—266
12．黄世文、余柳青，放线菌及其代谢产物对稗草和稻纹枯病的抑(杀)效果研究初报，植物保护21世纪展望，1998，中国科学技术出版社，693-696
13．李扬汉，张宗俭、王建书、冀玲惠，有关真菌除草剂研究的进展，生物防治通报，1994，10(1)，35-39
14．强胜，生物除草剂的最新研究进展，杂草科学，1997，(4)，9—10
15．申继忠，微生物农药剂型加工研究进展，中国生物防治，1998，14(3)，129—133
16．万方浩，包建中，世界农业害虫及杂草生物防治的进展及发展趋势，农牧情报研究，1993，(6)，13-21
17．王明旭、罗宽、陈寅，稗叶枯菌及其毒素的研究Ⅰ，病原鉴定及产孢产毒条件，湖南农学院学报，1991，17(1)，34-41
18．王明旭、罗宽、陈寊，稗叶叶菌及其毒素的研究Ⅱ，病菌寄主范围和对稗的致病性，湖南农学院学报，1993，19(3)，248-251
19．吴畏、杨家书等，小麦长蠕胞属毒素，真菌学报，1989，8(1)，70—78
20．邢来君，真菌除草剂的研究现状与展望，真菌学报，1995，14(2)，153-157
21．余柳青、黄世文、徐正浩，微生物除草剂的研究和开发进展，喻子牛等主编，微生物农药及其产业化，科学出版社，2000，256-259
22．喻子牛等，微生物除草剂的研究和开发进展，喻子牛等主编，微生物农药及其产业化，科学出版社，2000，256-259
23．张朝贤、钱益新、胡详恩、张建中，我国农田杂草综合治理现状和任务，孙鼎昌主编，面向21世纪中国农田杂草可持续治理(第六次全国杂草科学学术研讨会论文集)，广西民族出版社，1999，10—15
24．张泽溥，移栽稻田杂草治理，植物生长物质和除草剂研究及应用进展(第一辑)，中国农业科技出版社，1998，100—103
25．张宗俭、李扬汉，野燕麦生防真菌燕麦叶枯菌的分离培养及致病性研究，中国生物防治，1996，12(4)，171—173
26．赵永贵、郭超群、韩恒芬，微生物农药研究的现状及展望，河南农业大学学报，1995，29(3)，304-310
27．章元寿，关于植物病原真菌毒素研究中的几个问题的商榷，董金皋、李树正主编，植物病原菌毒素研究进展，1997，第一卷，中国科学技术出版社，北京，3—7
28．朱杰华，王江柱等，玉米大斑菌毒素产生及几种生物学特性与氮源的关系，董金皋、李树正主编，植物病原菌毒素研究进展，1997，第一卷，中国科学技术出版社，北京，75—80
29.Aldridge D. C., Turner W. B., Metabolites of Helminthosporium monoceras:Structures of monocerin and related benzopyrans, J. Chem. Soc. (C), 1970,2598-2600
30.Auld B. A., Morin L., Constraints in the development of bioherbicides, Weed
Technology, 1995, Vol. 9, 638-652
31 . Bashan. B., Levy, Y., Differential response of sweet corn cultivars to phytotoxic water-soluble compounds from culture filtrates of Exserohilum turcicum, Plant Disease, 1992, Vol. 76, No. 5, 451-454
32． Bashan. B., Levy, R. S., Cojocaru, M., Levy, Y., Purification and structural determination of a phytotoxic substance from Exserohilum turcicum, Physiological and Molecular Plant Pathology, 1995, Vol. 47, 225-235
33． Bilgrami, K. S., Sinha, K. K., Ansari, A. A., Rahman, M. F., Widespread occurrence of toxigenic Alternarias in cereals and oilseeds, Indian Phytopathology, 1995, Vol. 48, No. 2, 150-153
34． Bowers R. C., Commercialization of Collego-an industrialist's view, Weed Science, 1986, Vol. 34, 24-25
35． Bronson, C. R., Yoder, O. C., The genetics of phytotoxin production by plant pathogenic fungi, Experientia, 1991, Vol.47, 771-776
36． Claydon, N., Frederick Grove John, Pople, M., Insecticidal secondary metabolic products from the entomogenous fungus fusarium larbarum, Journal of Invertebrate Pathology, 1979, Vol. 33, 364-367
37． Ciotola, M., Tommaso, DI. A., Watson, A. K., Chlamydospore production, inoculation methods and pathogenicity of Fusarium oxysporum M12-4A, a biocontrol for Striga hermonthica, Biocontrol and Technology, 2000, Vol. 10, 129-145
38． Chung, Y. R., Kim, B. S., Kim, H. T., and Cho,K.Y. Identification of Exserohilum species, a fungi pathogen causing leaf blight of barnyard grass (Echinochloa crusglli), Korean J. Plant Pathol, 1990, Vol. 6, 429-433
39． Cullen, J. M., Kable, P. F., Catt, M., Epidemic spread of a rust imported for biological control, Nature, 1973, Vol. 244, 462-464
40． Cuq, F., Herrmann-Gorline, S., Klaebe, A., Monocerin in Exserohilum turcicum isolates from maize and a study of its phytotoxicity, Phytochemistry, 1993, Vol. 34, No. 5, 1265-1270
41． Cuq, F., Brown, S. C., Petitprez, M., Alibert, G., Effects of monocerin on cell cycle progression in maize root meristems synchronized with aphidicolin, Plant Cell Reports, 1995, Vol 15, 138-142
42． Dillon Michael P., Simpson Thomas J. and Sweeney J. B., Enantioselective synthesis of monocerin and fusarentin ethers: antifungal and insecticidal fungal metabolites, Tetrahedron Letters, 1992, Vol.33, No.49, 7569-7572
43． Evidente Antonio, Capasso Renato, Cutignano Adele, Taglialatela-Scafati Orazio, Vurro Maurizio, Zonno Maria Chiara, Motta Andrea, Ascaulitoxin, a phytotoxic bis-amino acid N-glucoside from Ascochyta caulina, Phytochemistry, Vol. 48, No. 7, 1131-1137
44． George E.Templeton 1988 阿肯色大学的真菌除草剂研究一过去、现在与将来 农药译丛 10(1) 23-25原载 Weed Science, 1986, Vol. 34, No. 1, 35-37
45． Gilchrist, D. G., Grogan, R. G., Production and nature of a host-specific toxin from Alternaria alternata f.sp lycopersiei, Phytopathology, 1976, Vol. 66, 165-171
46． Gohbara, M., and Yamaguchi, K., Biological control agents for rice paddy weed management in Japan, Integrated Management of Paddy and Aquatic Weeds in Asia, 1994, 184-194
47． Goto, m., The relationship between Emmalocera sp, and barnyard grass and its potential as a biological control, Integrated Management of Paddy and Aquatic Weeds in Asia. 113-121
48． Grove, Fredirick John, Pople, Michael, Metabolic products of Fusarium larvarum Fuckel. the fusarentins and the absolute configuration of monocerin, J. C. S. Perkin I, 1979,2048-2051
49． Harris, P., Host Specificity in Weed Biocontrol, The Proceedings. Of the 18th Asian-Pacific Weed Science Society Conference, 2001, Standards Press of China, Beijing, China, 341-347
50． Hawes, M.C. Technique for using isolated corn root cap cells in a simple, quantitative assay for the pathotoxin produced by Helminthosporium maydis Race T, Phytopathology, 1983, Vol. 73, No. 8, 1184-1187
51． Hetherington, S. D., Tuat, N. V, Auld, B. A., Thanh, H. M. The development of a bioherbicide for Echinochloa crusgalli based on the fungal pathogen Exserohilum monoceras in Vietnam, The Proceedings Of the 18th Asian-Pacific Weed Science Society Conference, 2001, Standards Press of China, Beijing, China, 387-392
52． Holm, L. G., Plucknett, D. L., Pancho, J. V, and Herberger, J. P., The world's
worst weeds. Distribution and Biology, The University Press of Hawaii, 1977, HI: The University Press of Hawaii, Honolulu, 32-46
53． Holmstrom-ruddick B., Mortensen, K., Factors affecting pathogenicity of a benomyl-resistant strain of Colletotrichum gloeosporioides f. Sp. malvae, Mycol. Res., 1995, Vol. 99, No. 9, 1108-1112
54． Kohmoto, K., Otani, H., Host recognition by toxigenic plant pathogens, Experientia, 1991, Vol. 47, 755-764
55． Michael, P. Dillon, Thomas, J. Simpson, Sweeney, J. B., Enantioselective systhesis of monocerin and fusarentin ethers: antifungal and insecticidal fungal metabolites, Tetrahedron Letters, Vol. 33, No. 49, 7569-7572
56． Mori Kenji and Takaishi Rideo, Synthesis of monocerin, an antifungal, insecticidal and phytotoxic heptaketide metabolite of Exserohilum monoceras, Tetrahedron, 1989, Vol.45, No. 6, 1639-1646
57． Norman, David J., Development of Colletotrichum gloeosporioides f. Sp. Clidemiae and Septoria passiflorae into two mycoherbicides with extended viability, Plant Disease, 1995, Vol. 79, No. 10, 1029-1032
58． Pringle, R. B., Scheffer, R. P., Isolation of the host-specific toxin and a related substance with nonspecific toxicity from Helminthosporium carbonum, Phytopathology, 1967, Vol. 57, 1169-1172
59． Quimby P. C., Jr Zidack N. K. A simple method for stabilizing and granulating fungi, Biocontrol Science and Technology, 1999, Vol. 9, 5-8
60． Richard L. Duble, Bipolaris & Exserohilum (Helminthosporium) Diseases, Turfgrass Specialist Texas Agricultural Extension Service, download from Internet
61． Robeson, D. J., Strobel, G. A., Monocerin, a phytotoxin from Exserohilum trucicum (Drechslera turcica). Agric. Biol. Chem. 1982, Vol. 46, 2681-83
62． Shibyama H., Kiritani K., and Bay-Petersen J., eds., FFTC Book series No.45． Food and Fertilizer Technology Center for the Asian and Pacific Region, 1992, Taipei Press, Taiwan.
63． Simpson, T. J., Prof Tom Simpson's research Interests: Bio-organic and natural productchemistry, from http: // www.tlchm.bris.ac.uk/org/research/simpres.htm
64． Simpson, Thomas J., Polyketide biosynthesis, Chemistry & Industry, 1995, June, 407-411
65． Simpson, Thomas J., Michael P. Dillon, Donovan, Tessa M., Synthetic and mechanistic studies on fungal metabolic pathways: a guide to fungicide design, Pestic. Sci. 1991, Vol. 31, 539-554
66． Stephen O. Duke, John Lydon,许学胜译，天然化合物源除草剂，农药译丛， 1989, 11 (3) , 14-18, 原载 Weed Technology, 1987, Vol. 1,No.2, 122-128
67． Steiner, G. W., Strobel, G. A., Helminthosporoside, a host specific toxin from Helminthosporium sacchari, J. Biol. Chem. 1971, Vol. 246, 4350-4357
68． Stierle, A., Sgrobel, G., Stierle, D., Sugawara, F., Analytical methods for phytotoxins. Pp. 1-31 ln H. F. Linskens and J. F. Jackson, Modern Methods of Plant Analysis New Series, Vol. 13, Plant Toxin Analysis, Springer-Verlag Berhn Heidelberg
69． Wan, F.H., Guo, J. Y., Wang, R., Biological weed control in China: an update report, The Proceedings Of the 18th Asian-Pacific Weed Science Society Conference, 2001, Standards Press of China, Beijing, China, 366-376
70． Watanabe, H., Uchino, A., Tachibana, M., Application period of Drechslera monoceras, plant pathogen for the mycoherbicide, to control Echinochloa oryzicola vasing in rice fields of Tohoku, Japan, The Proceedings Of the 18th Asian-Pacific Weed Science Society Conference, 2001, Standards Press of China, Beijing, China, 416-421
71 . Yoder, O.C., Payne, G.A., Gracen, V. E., Bioassays for detection and quantification of Helminthosporium maysis race T-toxin: a comparison, Physiol. Plant Pathol. Vol. 10, 237-245
72． Yang Young-Kook, Kim Soon-Ok, Chung Hoo-Sup, Lee Yong-Hwan, Use of Colletotrichum graminicola KA001 to Control Barnyard Grass, Plant Disease, 2000, Vol. 84, No. 1,55-59
73． Zhang W. M., Moody K., Watson A. K., Responses of Echinochloa Species and rice (Oryza saliva) to indigenous pathogenic fungi, Plant Disease, 1996, Vol. 80, No. 9, 1053-1058
74． Zhang W. M., Watson A. K., Characterization of growth and conidia production of Exserohilun monoceras on different substrates, Biocontrol Science and Technology, 1997, Vol. 7, 75-86
75． Zhang Wenming, Watson Alan K., Host range of Exserohilum monoceras, a
potential bioherbicide for the control pf Echinochloa species, Canadian Journal of Botany, 1997, Vol. 75, 685-692
76． Zhang W. M., Sulz M., Myketiak, T., Potential of using microbes and microbial natural products for control of weeds, The Proceedings. Of the 18th Asian-Pacific Weed Science Society Conference, 2001, Standards Press of China, Beijing, China, 358-365
2．陈涛主编，有害生物的微生物防治原理和技术，湖北科学技术发展出版社，1997，408-448
3．陈勇，尖角突脐孢菌作为真菌除草剂防除稗草潜力的研究，中国农业大学博士学位论文，2001
4．陈勇强等，国外微生物除草剂的研究及应用现状，天津农业科学，1998 4(20)，5-9
5．丁建清，农田杂草的生物防治，中国生物防治，1995，11(3)，129—123
6．董金皋、王江柱，植物病原真菌毒素活性测定方法，董金皋、李树正主编，植物病原菌毒素研究进展，1997，第一卷，中国科学技术出版社，北京，61—74
7．董金皋、黄梧芳，玉米叶斑病菌毒素的研究现状，董金皋、李树正主编，植物病原菌毒素研究进展，1997，第一卷，中国科学技术出版社，北京，102—126
8．高昭远等，我国利用植物病原微生物防除杂草研究的回顾与现状，1995，中国农学通报，11(1)，31-34
9．菅原二三男，利用植物病原微生物开发除草剂，农药译丛，1992，14(3)，40—45
10．郭凤根、李扬汉、邓福珍，园林菟丝子生防真菌的筛选研究，中国生物防治，1998，14(4)，159-162
11．黄世文、Alan K.Watson、段桂芳、余柳青，利用稗草病原菌开发微生物除草剂的研究，喻子牛等主编，微生物农药及其产业化，2000，科学出版社，260—266
12．黄世文、余柳青，放线菌及其代谢产物对稗草和稻纹枯病的抑(杀)效果研究初报，植物保护21世纪展望，1998，中国科学技术出版社，693-696
13．李扬汉，张宗俭、王建书、冀玲惠，有关真菌除草剂研究的进展，生物防治通报，1994，10(1)，35-39
14．强胜，生物除草剂的最新研究进展，杂草科学，1997，(4)，9—10
15．申继忠，微生物农药剂型加工研究进展，中国生物防治，1998，14(3)，129—133
16．万方浩，包建中，世界农业害虫及杂草生物防治的进展及发展趋势，农牧情报研究，1993，(6)，13-21
17．王明旭、罗宽、陈寅，稗叶枯菌及其毒素的研究Ⅰ，病原鉴定及产孢产毒条件，湖南农学院学报，1991，17(1)，34-41
18．王明旭、罗宽、陈寊，稗叶叶菌及其毒素的研究Ⅱ，病菌寄主范围和对稗的致病性，湖南农学院学报，1993，19(3)，248-251
19．吴畏、杨家书等，小麦长蠕胞属毒素，真菌学报，1989，8(1)，70—78
20．邢来君，真菌除草剂的研究现状与展望，真菌学报，1995，14(2)，153-157
21．余柳青、黄世文、徐正浩，微生物除草剂的研究和开发进展，喻子牛等主编，微生物农药及其产业化，科学出版社，2000，256-259
22．喻子牛等，微生物除草剂的研究和开发进展，喻子牛等主编，微生物农药及其产业化，科学出版社，2000，256-259
23．张朝贤、钱益新、胡详恩、张建中，我国农田杂草综合治理现状和任务，孙鼎昌主编，面向21世纪中国农田杂草可持续治理(第六次全国杂草科学学术研讨会论文集)，广西民族出版社，1999，10—15
24．张泽溥，移栽稻田杂草治理，植物生长物质和除草剂研究及应用进展(第一辑)，中国农业科技出版社，1998，100—103
25．张宗俭、李扬汉，野燕麦生防真菌燕麦叶枯菌的分离培养及致病性研究，中国生物防治，1996，12(4)，171—173
26．赵永贵、郭超群、韩恒芬，微生物农药研究的现状及展望，河南农业大学学报，1995，29(3)，304-310
27．章元寿，关于植物病原真菌毒素研究中的几个问题的商榷，董金皋、李树正主编，植物病原菌毒素研究进展，1997，第一卷，中国科学技术出版社，北京，3—7
28．朱杰华，王江柱等，玉米大斑菌毒素产生及几种生物学特性与氮源的关系，董金皋、李树正主编，植物病原菌毒素研究进展，1997，第一卷，中国科学技术出版社，北京，75—80
29.Aldridge D. C., Turner W. B., Metabolites of Helminthosporium monoceras:Structures of monocerin and related benzopyrans, J. Chem. Soc. (C), 1970,2598-2600
30.Auld B. A., Morin L., Constraints in the development of bioherbicides, Weed
Technology, 1995, Vol. 9, 638-652
31 . Bashan. B., Levy, Y., Differential response of sweet corn cultivars to phytotoxic water-soluble compounds from culture filtrates of Exserohilum turcicum, Plant Disease, 1992, Vol. 76, No. 5, 451-454
32． Bashan. B., Levy, R. S., Cojocaru, M., Levy, Y., Purification and structural determination of a phytotoxic substance from Exserohilum turcicum, Physiological and Molecular Plant Pathology, 1995, Vol. 47, 225-235
33． Bilgrami, K. S., Sinha, K. K., Ansari, A. A., Rahman, M. F., Widespread occurrence of toxigenic Alternarias in cereals and oilseeds, Indian Phytopathology, 1995, Vol. 48, No. 2, 150-153
34． Bowers R. C., Commercialization of Collego-an industrialist's view, Weed Science, 1986, Vol. 34, 24-25
35． Bronson, C. R., Yoder, O. C., The genetics of phytotoxin production by plant pathogenic fungi, Experientia, 1991, Vol.47, 771-776
36． Claydon, N., Frederick Grove John, Pople, M., Insecticidal secondary metabolic products from the entomogenous fungus fusarium larbarum, Journal of Invertebrate Pathology, 1979, Vol. 33, 364-367
37． Ciotola, M., Tommaso, DI. A., Watson, A. K., Chlamydospore production, inoculation methods and pathogenicity of Fusarium oxysporum M12-4A, a biocontrol for Striga hermonthica, Biocontrol and Technology, 2000, Vol. 10, 129-145
38． Chung, Y. R., Kim, B. S., Kim, H. T., and Cho,K.Y. Identification of Exserohilum species, a fungi pathogen causing leaf blight of barnyard grass (Echinochloa crusglli), Korean J. Plant Pathol, 1990, Vol. 6, 429-433
39． Cullen, J. M., Kable, P. F., Catt, M., Epidemic spread of a rust imported for biological control, Nature, 1973, Vol. 244, 462-464
40． Cuq, F., Herrmann-Gorline, S., Klaebe, A., Monocerin in Exserohilum turcicum isolates from maize and a study of its phytotoxicity, Phytochemistry, 1993, Vol. 34, No. 5, 1265-1270
41． Cuq, F., Brown, S. C., Petitprez, M., Alibert, G., Effects of monocerin on cell cycle progression in maize root meristems synchronized with aphidicolin, Plant Cell Reports, 1995, Vol 15, 138-142
42． Dillon Michael P., Simpson Thomas J. and Sweeney J. B., Enantioselective synthesis of monocerin and fusarentin ethers: antifungal and insecticidal fungal metabolites, Tetrahedron Letters, 1992, Vol.33, No.49, 7569-7572
43． Evidente Antonio, Capasso Renato, Cutignano Adele, Taglialatela-Scafati Orazio, Vurro Maurizio, Zonno Maria Chiara, Motta Andrea, Ascaulitoxin, a phytotoxic bis-amino acid N-glucoside from Ascochyta caulina, Phytochemistry, Vol. 48, No. 7, 1131-1137
44． George E.Templeton 1988 阿肯色大学的真菌除草剂研究一过去、现在与将来 农药译丛 10(1) 23-25原载 Weed Science, 1986, Vol. 34, No. 1, 35-37
45． Gilchrist, D. G., Grogan, R. G., Production and nature of a host-specific toxin from Alternaria alternata f.sp lycopersiei, Phytopathology, 1976, Vol. 66, 165-171
46． Gohbara, M., and Yamaguchi, K., Biological control agents for rice paddy weed management in Japan, Integrated Management of Paddy and Aquatic Weeds in Asia, 1994, 184-194
47． Goto, m., The relationship between Emmalocera sp, and barnyard grass and its potential as a biological control, Integrated Management of Paddy and Aquatic Weeds in Asia. 113-121
48． Grove, Fredirick John, Pople, Michael, Metabolic products of Fusarium larvarum Fuckel. the fusarentins and the absolute configuration of monocerin, J. C. S. Perkin I, 1979,2048-2051
49． Harris, P., Host Specificity in Weed Biocontrol, The Proceedings. Of the 18th Asian-Pacific Weed Science Society Conference, 2001, Standards Press of China, Beijing, China, 341-347
50． Hawes, M.C. Technique for using isolated corn root cap cells in a simple, quantitative assay for the pathotoxin produced by Helminthosporium maydis Race T, Phytopathology, 1983, Vol. 73, No. 8, 1184-1187
51． Hetherington, S. D., Tuat, N. V, Auld, B. A., Thanh, H. M. The development of a bioherbicide for Echinochloa crusgalli based on the fungal pathogen Exserohilum monoceras in Vietnam, The Proceedings Of the 18th Asian-Pacific Weed Science Society Conference, 2001, Standards Press of China, Beijing, China, 387-392
52． Holm, L. G., Plucknett, D. L., Pancho, J. V, and Herberger, J. P., The world's
worst weeds. Distribution and Biology, The University Press of Hawaii, 1977, HI: The University Press of Hawaii, Honolulu, 32-46
53． Holmstrom-ruddick B., Mortensen, K., Factors affecting pathogenicity of a benomyl-resistant strain of Colletotrichum gloeosporioides f. Sp. malvae, Mycol. Res., 1995, Vol. 99, No. 9, 1108-1112
54． Kohmoto, K., Otani, H., Host recognition by toxigenic plant pathogens, Experientia, 1991, Vol. 47, 755-764
55． Michael, P. Dillon, Thomas, J. Simpson, Sweeney, J. B., Enantioselective systhesis of monocerin and fusarentin ethers: antifungal and insecticidal fungal metabolites, Tetrahedron Letters, Vol. 33, No. 49, 7569-7572
56． Mori Kenji and Takaishi Rideo, Synthesis of monocerin, an antifungal, insecticidal and phytotoxic heptaketide metabolite of Exserohilum monoceras, Tetrahedron, 1989, Vol.45, No. 6, 1639-1646
57． Norman, David J., Development of Colletotrichum gloeosporioides f. Sp. Clidemiae and Septoria passiflorae into two mycoherbicides with extended viability, Plant Disease, 1995, Vol. 79, No. 10, 1029-1032
58． Pringle, R. B., Scheffer, R. P., Isolation of the host-specific toxin and a related substance with nonspecific toxicity from Helminthosporium carbonum, Phytopathology, 1967, Vol. 57, 1169-1172
59． Quimby P. C., Jr Zidack N. K. A simple method for stabilizing and granulating fungi, Biocontrol Science and Technology, 1999, Vol. 9, 5-8
60． Richard L. Duble, Bipolaris & Exserohilum (Helminthosporium) Diseases, Turfgrass Specialist Texas Agricultural Extension Service, download from Internet
61． Robeson, D. J., Strobel, G. A., Monocerin, a phytotoxin from Exserohilum trucicum (Drechslera turcica). Agric. Biol. Chem. 1982, Vol. 46, 2681-83
62． Shibyama H., Kiritani K., and Bay-Petersen J., eds., FFTC Book series No.45． Food and Fertilizer Technology Center for the Asian and Pacific Region, 1992, Taipei Press, Taiwan.
63． Simpson, T. J., Prof Tom Simpson's research Interests: Bio-organic and natural productchemistry, from http: // www.tlchm.bris.ac.uk/org/research/simpres.htm
64． Simpson, Thomas J., Polyketide biosynthesis, Chemistry & Industry, 1995, June, 407-411
65． Simpson, Thomas J., Michael P. Dillon, Donovan, Tessa M., Synthetic and mechanistic studies on fungal metabolic pathways: a guide to fungicide design, Pestic. Sci. 1991, Vol. 31, 539-554
66． Stephen O. Duke, John Lydon,许学胜译，天然化合物源除草剂，农药译丛， 1989, 11 (3) , 14-18, 原载 Weed Technology, 1987, Vol. 1,No.2, 122-128
67． Steiner, G. W., Strobel, G. A., Helminthosporoside, a host specific toxin from Helminthosporium sacchari, J. Biol. Chem. 1971, Vol. 246, 4350-4357
68． Stierle, A., Sgrobel, G., Stierle, D., Sugawara, F., Analytical methods for phytotoxins. Pp. 1-31 ln H. F. Linskens and J. F. Jackson, Modern Methods of Plant Analysis New Series, Vol. 13, Plant Toxin Analysis, Springer-Verlag Berhn Heidelberg
69． Wan, F.H., Guo, J. Y., Wang, R., Biological weed control in China: an update report, The Proceedings Of the 18th Asian-Pacific Weed Science Society Conference, 2001, Standards Press of China, Beijing, China, 366-376
70． Watanabe, H., Uchino, A., Tachibana, M., Application period of Drechslera monoceras, plant pathogen for the mycoherbicide, to control Echinochloa oryzicola vasing in rice fields of Tohoku, Japan, The Proceedings Of the 18th Asian-Pacific Weed Science Society Conference, 2001, Standards Press of China, Beijing, China, 416-421
71 . Yoder, O.C., Payne, G.A., Gracen, V. E., Bioassays for detection and quantification of Helminthosporium maysis race T-toxin: a comparison, Physiol. Plant Pathol. Vol. 10, 237-245
72． Yang Young-Kook, Kim Soon-Ok, Chung Hoo-Sup, Lee Yong-Hwan, Use of Colletotrichum graminicola KA001 to Control Barnyard Grass, Plant Disease, 2000, Vol. 84, No. 1,55-59
73． Zhang W. M., Moody K., Watson A. K., Responses of Echinochloa Species and rice (Oryza saliva) to indigenous pathogenic fungi, Plant Disease, 1996, Vol. 80, No. 9, 1053-1058
74． Zhang W. M., Watson A. K., Characterization of growth and conidia production of Exserohilun monoceras on different substrates, Biocontrol Science and Technology, 1997, Vol. 7, 75-86
75． Zhang Wenming, Watson Alan K., Host range of Exserohilum monoceras, a
potential bioherbicide for the control pf Echinochloa species, Canadian Journal of Botany, 1997, Vol. 75, 685-692
76． Zhang W. M., Sulz M., Myketiak, T., Potential of using microbes and microbial natural products for control of weeds, The Proceedings. Of the 18th Asian-Pacific Weed Science Society Conference, 2001, Standards Press of China, Beijing, China, 358-365