狗牙根弯孢霉叶斑病的病原学研究
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摘要
叶斑病是发生在高尔夫球场草坪上的一种重要病害，主要引起多种草坪草的叶斑和叶枯，严重影响草坪景观，破坏草坪质量。目前对该病的研究仍仅限于病害的调查和初步的化学防治，国内外还未见有较系统的研究报道。本研究旨在确定引起狗牙根叶斑病的病原，了解病菌的寄主范围、生物学特性，以及进行杀菌剂的初步筛选和病菌对四种草坪草的侵入过程观察。研究结果如下：
从鄂州红莲湖高尔夫球场的狗牙根发病植株叶片上分离菌株Curvularia sp.，用菌丝块接种狗牙根叶片进行致病性测定，证实Curvularia sp.为致病菌。该菌在PDA培养基上培养5d后，菌落呈墨绿色，菌丝褐色，有隔，分枝，分生孢子平均大小为11.07×25.18μm，褐色至暗褐色，椭圆形，具有3个隔膜，主隔膜明显加粗，中间两细胞较大。rDNA-ITS测序结果与GeneBank中的序列进行同源性比较发现菌株与Curvularia verruculosus的同源性高达99％。根据形态学特征和分子鉴定结果，确认该病原菌为Curvularia verruculosus。寄主范围测定结果表明，该菌可侵染供试的四种草坪草(狗牙根、黑麦草、高羊茅、早熟禾)、水稻、小麦和玉米等禾本科植物。
病原菌生物学特性研究结果表明，在有无光照或连续光照的条件下病菌均可生长和产孢，光暗交替最有利于菌丝的生长，黑暗条件下促进病菌产孢。该菌株在15-35℃的范围内均能生长和产孢，30℃为菌丝生长最快温度，最适产孢温度为25℃。病菌在pH值为4-12的PDA培养基上菌丝均能生长，其中以pH7生长最快；病原菌在pH 5-12范围内均可产孢，产孢最适pH值为6。病菌能够有效利用多种碳源和氮源，甚至在无碳或无氮条件下都能生长，但不同碳源和氮源对弯孢霉病菌菌丝生长和产孢有极显著的影响。病原菌分别以淀粉为碳源以及以酵母膏为氮源时生长最好，半乳糖和硝酸钙有利于该菌分生孢子的产生。试验证明55℃的温水处理10min可杀死菌丝体、抑制孢子萌发。PDA平板抑菌实验结果表明，供试六种药剂在设置的不同浓度下对病原菌的生长都有一定的抑制作用，其中10％世高可分散粒剂和30％爱苗乳油对菌丝生长有很好的抑制效果，10％多抗霉素B粉剂抑制效果最差。
喷雾病原菌分生孢子悬浮液于狗牙根、黑麦草、高羊茅、早熟禾等几种草坪草活体叶片上并保湿培养，在接种后不同时间点分别从每个寄主上取样，对样品进行组织透明、染色，观察病原菌对寄主植物的侵染过程。结果表明，在四种寄主上接种后3h内均可以观察到分生孢子的萌发，分生孢子可以从一端或两端萌发，产生单根或2-4根芽管。接种后3h时分生孢子在不同寄主叶片上的萌发率没有差异性，而6h后分生孢子在不同寄主上的萌发率有微小差异，12h时，病原菌分生孢子在狗牙根和早熟禾叶片上的萌发率分别为96％和92％，略高于其在黑麦草和高羊茅上的萌发率。弯孢霉分生孢子在四种不同的草坪草上都较易萌发，24h时萌发率均超过90％，48h时在狗牙根叶片上萌发率达100％。
病原菌分生孢子在寄主表面萌发后，可以从叶片气孔或表皮直接侵入，其中以气孔侵入为主。接种后6h便可以观察到有病菌从气孔侵入，病原菌分生孢子在四种草坪草寄主上从气孔侵入的百分率有所差异，24h时萌发的分生孢子在狗牙根和早熟禾上侵入的百分率较在黑麦草和高羊茅上的侵入百分率高。菌丝直接侵入寄主组织的情况比较少，接种后12h可观察到部分菌丝分支产生类似侵染钉的结构从同一部位或不同部位直接侵入寄主细胞。
The Leaf spot is one of important diseases in golf courses. It causes leaf spot and leaf blight on many species of grasses, and can influence sights and destroy quality of grasses. At present there are only reports about the disease investigation and primary chemical control. The aim of this study is to indentify the pathogen of bermudagrass Leaf spot, to understand host range, biological characteristics, and the infection process of the pathogen on four species of grasses. The results showed as follow:
Curvularia sp. was isolated from the diseased leaf of Cynodon hybrid in Red Lotus Lake golf courses in E-zhou. Pathogenicity was tested by inoculating the leaf with mycelium plug of Curvularia sp., the result indicated that Curvularia sp. isolate was the pathogen. The morphological characteristic showed that the colonies were black green and the hyphae were septum, brown, Spore size was 11.07×25.18μm, mostly ellipse, with three septate , two central cells were larger than both terminal cells , brown to dark brown , terminal cells subhyaline to light Brown after cultured on PDA medium at 5d. The DNA sequence of ribosomal ITS of the isolates was found to match 99% with an ITS sequence of Curvularia verruculosus in GenBank. According to the morphological characteristics, and the DNA sequence of ribosomal ITS, the pathogen was identified as C.verruculosus. The pathogen can infected four turfgrass species (Cynodon hybrid, Lolium perenne, Agrostis plustris, Poa pratensis) and other gramineous plants ( Oryzae sativa, Blumeria tristic, Zea mays).
The studies on biological characteristic revealed that this fungus could grow and sporulate under dark or continuous illumination, the optimal illumination condition for growth of mycelium was alternant illumination and darkness facilitated spotulation.The mycelium could grow under the temperature from 15-35℃, the optimal temperature for the mycelial growth and sporulation was 30℃and 25℃, respectively. pH value range for mycelium growth were 4-12 and pH7 was optimal. pH value range for sporulation were 5-12 and pH6 was optimal. The isolate could utilize various carbon and nitrogen sources, but different carbon and nitrogen sources had different influence for mycelial growth and sporulation. Among carbon and nitrogen sources, starch and yeast extract were best for mycelial growth, D-galctose and CaNO_3 were best for sporulation.The lethal temperature for mycelial growth and spores gemination were 55℃for 10 min, respectively.
The result of determining of the toxicity of fungicides to C.verruculosus showed that the six fundicides choosed could inhibit mycelial growth at different concentration. 10% Score GR and 30% Armure EC showed best inhibiting to mycelial growth, but the 10% polyoxin B DP was the worst.
Leaves of four turfgrass species (Cynodon hybrid, Lolium perenne, Agrostis plustris, Poa pratensis) were inoculated with conidial suspensions of C.verruculosus. Inoculated leaves were incubated at 25℃under high humidity.Samples were collected from inoculated leaves at different timing intervals,and then decolorated and stained. Observe the infection process on four species of grasses with fluorescence microscope under bright field.. The result indicated that the conidia began to germinate after 3 hours of inoculation, and formed one to four primary germtubes from one or two ends. The rate of conidial germination of C.verruculosus on leaves of four turfgrass species after 3h of inoculation showed no significant differences. After 12 hours of inoculation, the percentage of germination were significantly larger on the Cynodon hybrid and Poa pratensis than on the other two hosts.
The pathogen C.verruculosus was able to penetrate host through two paths such as leaf stomata and epidermis.The penetration from leaf stomata was considered as a major way, and can be observed after 6h of inoculation.The rate of pathogen infecting from leaf stomata of four turfgrass species showed significant differences. After 24 hours of inoculation, the percentage of infecting were significantly larger on the Cynodon hybrid and Poa pratensis than the other two hosts. Direct penetration into epidermis was uncommon observed.
从鄂州红莲湖高尔夫球场的狗牙根发病植株叶片上分离菌株Curvularia sp.，用菌丝块接种狗牙根叶片进行致病性测定，证实Curvularia sp.为致病菌。该菌在PDA培养基上培养5d后，菌落呈墨绿色，菌丝褐色，有隔，分枝，分生孢子平均大小为11.07×25.18μm，褐色至暗褐色，椭圆形，具有3个隔膜，主隔膜明显加粗，中间两细胞较大。rDNA-ITS测序结果与GeneBank中的序列进行同源性比较发现菌株与Curvularia verruculosus的同源性高达99％。根据形态学特征和分子鉴定结果，确认该病原菌为Curvularia verruculosus。寄主范围测定结果表明，该菌可侵染供试的四种草坪草(狗牙根、黑麦草、高羊茅、早熟禾)、水稻、小麦和玉米等禾本科植物。
病原菌生物学特性研究结果表明，在有无光照或连续光照的条件下病菌均可生长和产孢，光暗交替最有利于菌丝的生长，黑暗条件下促进病菌产孢。该菌株在15-35℃的范围内均能生长和产孢，30℃为菌丝生长最快温度，最适产孢温度为25℃。病菌在pH值为4-12的PDA培养基上菌丝均能生长，其中以pH7生长最快；病原菌在pH 5-12范围内均可产孢，产孢最适pH值为6。病菌能够有效利用多种碳源和氮源，甚至在无碳或无氮条件下都能生长，但不同碳源和氮源对弯孢霉病菌菌丝生长和产孢有极显著的影响。病原菌分别以淀粉为碳源以及以酵母膏为氮源时生长最好，半乳糖和硝酸钙有利于该菌分生孢子的产生。试验证明55℃的温水处理10min可杀死菌丝体、抑制孢子萌发。PDA平板抑菌实验结果表明，供试六种药剂在设置的不同浓度下对病原菌的生长都有一定的抑制作用，其中10％世高可分散粒剂和30％爱苗乳油对菌丝生长有很好的抑制效果，10％多抗霉素B粉剂抑制效果最差。
喷雾病原菌分生孢子悬浮液于狗牙根、黑麦草、高羊茅、早熟禾等几种草坪草活体叶片上并保湿培养，在接种后不同时间点分别从每个寄主上取样，对样品进行组织透明、染色，观察病原菌对寄主植物的侵染过程。结果表明，在四种寄主上接种后3h内均可以观察到分生孢子的萌发，分生孢子可以从一端或两端萌发，产生单根或2-4根芽管。接种后3h时分生孢子在不同寄主叶片上的萌发率没有差异性，而6h后分生孢子在不同寄主上的萌发率有微小差异，12h时，病原菌分生孢子在狗牙根和早熟禾叶片上的萌发率分别为96％和92％，略高于其在黑麦草和高羊茅上的萌发率。弯孢霉分生孢子在四种不同的草坪草上都较易萌发，24h时萌发率均超过90％，48h时在狗牙根叶片上萌发率达100％。
病原菌分生孢子在寄主表面萌发后，可以从叶片气孔或表皮直接侵入，其中以气孔侵入为主。接种后6h便可以观察到有病菌从气孔侵入，病原菌分生孢子在四种草坪草寄主上从气孔侵入的百分率有所差异，24h时萌发的分生孢子在狗牙根和早熟禾上侵入的百分率较在黑麦草和高羊茅上的侵入百分率高。菌丝直接侵入寄主组织的情况比较少，接种后12h可观察到部分菌丝分支产生类似侵染钉的结构从同一部位或不同部位直接侵入寄主细胞。
The Leaf spot is one of important diseases in golf courses. It causes leaf spot and leaf blight on many species of grasses, and can influence sights and destroy quality of grasses. At present there are only reports about the disease investigation and primary chemical control. The aim of this study is to indentify the pathogen of bermudagrass Leaf spot, to understand host range, biological characteristics, and the infection process of the pathogen on four species of grasses. The results showed as follow:
Curvularia sp. was isolated from the diseased leaf of Cynodon hybrid in Red Lotus Lake golf courses in E-zhou. Pathogenicity was tested by inoculating the leaf with mycelium plug of Curvularia sp., the result indicated that Curvularia sp. isolate was the pathogen. The morphological characteristic showed that the colonies were black green and the hyphae were septum, brown, Spore size was 11.07×25.18μm, mostly ellipse, with three septate , two central cells were larger than both terminal cells , brown to dark brown , terminal cells subhyaline to light Brown after cultured on PDA medium at 5d. The DNA sequence of ribosomal ITS of the isolates was found to match 99% with an ITS sequence of Curvularia verruculosus in GenBank. According to the morphological characteristics, and the DNA sequence of ribosomal ITS, the pathogen was identified as C.verruculosus. The pathogen can infected four turfgrass species (Cynodon hybrid, Lolium perenne, Agrostis plustris, Poa pratensis) and other gramineous plants ( Oryzae sativa, Blumeria tristic, Zea mays).
The studies on biological characteristic revealed that this fungus could grow and sporulate under dark or continuous illumination, the optimal illumination condition for growth of mycelium was alternant illumination and darkness facilitated spotulation.The mycelium could grow under the temperature from 15-35℃, the optimal temperature for the mycelial growth and sporulation was 30℃and 25℃, respectively. pH value range for mycelium growth were 4-12 and pH7 was optimal. pH value range for sporulation were 5-12 and pH6 was optimal. The isolate could utilize various carbon and nitrogen sources, but different carbon and nitrogen sources had different influence for mycelial growth and sporulation. Among carbon and nitrogen sources, starch and yeast extract were best for mycelial growth, D-galctose and CaNO_3 were best for sporulation.The lethal temperature for mycelial growth and spores gemination were 55℃for 10 min, respectively.
The result of determining of the toxicity of fungicides to C.verruculosus showed that the six fundicides choosed could inhibit mycelial growth at different concentration. 10% Score GR and 30% Armure EC showed best inhibiting to mycelial growth, but the 10% polyoxin B DP was the worst.
Leaves of four turfgrass species (Cynodon hybrid, Lolium perenne, Agrostis plustris, Poa pratensis) were inoculated with conidial suspensions of C.verruculosus. Inoculated leaves were incubated at 25℃under high humidity.Samples were collected from inoculated leaves at different timing intervals,and then decolorated and stained. Observe the infection process on four species of grasses with fluorescence microscope under bright field.. The result indicated that the conidia began to germinate after 3 hours of inoculation, and formed one to four primary germtubes from one or two ends. The rate of conidial germination of C.verruculosus on leaves of four turfgrass species after 3h of inoculation showed no significant differences. After 12 hours of inoculation, the percentage of germination were significantly larger on the Cynodon hybrid and Poa pratensis than on the other two hosts.
The pathogen C.verruculosus was able to penetrate host through two paths such as leaf stomata and epidermis.The penetration from leaf stomata was considered as a major way, and can be observed after 6h of inoculation.The rate of pathogen infecting from leaf stomata of four turfgrass species showed significant differences. After 24 hours of inoculation, the percentage of infecting were significantly larger on the Cynodon hybrid and Poa pratensis than the other two hosts. Direct penetration into epidermis was uncommon observed.
引文
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35．赵杰．ITS序列分析及其在植物真菌病害分子检测中的应用．陕西农业科学，2004(4)：35-37
36．赵美琦，孙明，王慧敏．草坪病害(草坪全景)．北京：中国林业出版社，1999， 131-196
37．赵美琦，孙彦，张青文．草坪养护技术．北京：中国林业出版社，2001，93-120
38．朱灿星．稻瘟病菌与水稻品种相互关系中附着胞行为的观察．浙江大学学报(农业与生命科学版)，1984，03
39．朱云枝，强胜．马唐生防真菌弯孢霉(Curvularia sp.)菌株QZ-200的生物学特性研究．植物病理学报，2003，33(6)：524-529
40. Boedijn K B. Ueber einige phragmosporen Dematiazeen. Bull. Jard Bot Buitenzorg (Ser 3) 1933, 13: 120-134
41. Brown G E, Cole H J, and Nelson R R. Pathogenicity of Curvularia spp. to turfgrass. Plant Disease, 1972, 56: 59-63
42. Busch L V, Walker J C. Studies of cucumber anthracnose. Phytopathology, 1958, 48: 302-304
43. Carpita N, Gibeaut D. Structural models of primary cell walls in flowering plants: consistency of molecular structure with the physical properties of the walls during growth. Plant J, 1993, 3:1-3
44. Clemens B, Karl F G, Martin S, Paul L, Holger B D, Martin. Optical Measurements of Invasive Forces Exerted by Appressoria of a Plant Pathogenic Fungus. Science, 1999:1896-1899
45. Cooper R M. The mechanisms and significance of enzymatic degradation of host cell walls by parasites. Callow J A. Biochemical Plant Pathology. New York: John Wiley &. Sons Ltd, 1983:101-135
46. Deising H B, Werner S, Wemitz M. The role of fungal appressoria in plant infection. Microbes Infect, 2000, 2:1631-1641
47. Emmett R W, Parbery D G. Appressoria. Ann Rev Phytopathol, 1975, 13:147-167
48. Ellis M B . Dematiaceous hyphomycetes. Commonwealth Mycological Institute, Kew, Surrey, England, 1971:432-459
49. Hartmann H A, Kahmann R, Biker M. The pheromone response factor coordinates filamentous growth and pathogenic development in Ustilago maydis. EMBO J, 1996, 15:1632-1641
50. Howard F L. Helminthosporium-Curvularia Blights of Turf and Their Cure. Golf Course Rep. 1951, 21(2): 5-9
51. Hosokawa M, Tanaka C , Tsuda M .Conidium morphology of Curvularia geniculata and allied species. Mycoscience, 2003,44: 227-237
52. Huang J B , Zheng L and Hsiang T . First report of leaf spot caused by Curvularia affinis on Festuca arundinacea in Hubei, China. Plant Dis, 2004, 88: 1048
53. Kang Z , Huang L, Buchenauer H . Cytochemistry of cell wall component alterations in wheat roots infected by Gaeumannomyces graminis var.tritici. Journal of Plant Diseases and Protection, 2000,107(4): 337-351
54. Kauraw L P . Effect of Fungicides on the germination, RootlShoot Growth and Incidence of Seed-borne Pathogens in Rice. Central Indian Phytopathology, 1986, (39): 609-610
55. Kawanabe Y . The occurrence and control of turgrass diseases in Japan. Pesticide Information, 1991(59): 5-9
56. Khan A, Hsiang T .The infection process of Colletotrichum graminicola and relative aggressiveness on four turfgrass species. Can.J.Microbiol. 2003,49:433-442
57. Li G Q , Huang H C , Laroche A L ,and Acharya S N . Occurrence and characterization of hypovirulence in the tansclerotial isolate S10 of Sclerotinia sclerotiorum. Mycol. Res. 2003,107:1350-1360
58. Mandokhot R. Chemical changes in maize leaves in response to leaf spot pathogens. Indian Phytopathology, 1979, 32(4):658-660
59. Muchovej J J . Definition of Agrostis palustris Leaf Health at the Time of Infection and Colonization by Curvulatia lunata. Ann. Appl. Biol. 1986, 109: 249-258
60. Pietro D A , Roncero M I G. Cloning, expression, and role in pathogenicity of pg1 encoding the major extracellular endopolygalacturonase of the vascular wilt pathogen Fusarium oxysporum. Mol Plant-Microbe Interact, 1998, 11: 91-98
61. Shaw-ming yang. Isolation and effect of temperature on spore germination, radial growth, and pathogenicity of Curvularia senegalensis. Phytopathological notes, 1973,63:1541-1545
62. Sherwood R T, Vance C P . Resistance to Fungal Penetration in Gramineae. Cytology and Histology, 1980(70)4: 273-278
63. Smiley W R , Dernoeden P H , Clarke B B . Compendium of turfgrass diseases. Second Edition. The American Phytopathological Society, Minneapolis, Minnesota, USA, 1992: 1-296
64. Smith J D , Jackson N , Woolhouse A R . Fungal diseases of amenity turf grasses. Third Edition. New York, USA: E&F.N. Spon, 1989: 1-38
65. Talbot N J . Fungal biology-coming up for air and sporulation. Nature, 1999, 398: 295-296
66. Tonukari N J, Scott-Craig J S , Walton J D . The Cochliobolus carbonum SNF1 gene is required for cell wall-degrading enzyme expression and virulence on maize. Plant Cell, 2000,12:237-248
67. Tucker S L , Talbot N J . Surface attachment and pre-penetration stage development by plant pathogenic fungi. Ann Rev Phytopathol, 2001, 39: 385-417
68. Vloutoglou I, Kalogerakis S N. Effects of inoculum concentration, wetness duration and plant age on development of early blight and on shedding of leaves in tomato plants. Plant Pathology, 2000,49:339-345
69. Wernham C C , Kirby R S . A New Turf Disease. Phytopathology, 1941, 31: 24
70. Walton J D . Deconstructing the cell wall. Pl. Physiol, 1994,104(4): 1113-1118
71. Yildiz F . The preliminary sdudies on turgrass diseases in Turkey. Journal of Turkish Phytopathology, 1990,19(1): 21-29
2．陈捷，蔺瑞明，高增贵等．玉米弯孢叶斑病菌毒素对寄主防御酶系活性的影响及诱导抗性效应．植物病理学报，2002，32(1)：43-48
3．鄂洪海．玉米弯孢霉叶斑病研究进展．沈阳农业大学报，1999，30(3)：369-371
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5．方中达．植病研究方法．中国农业出版社，1998
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13．蒋家珍，吴学民，何凤奇．几种杀菌剂对4种草坪致病菌的毒力测定．植物保护，2004，6：78-80
14．康振生，黄丽丽，韩青梅，蒋选利．禾谷镰刀菌在小麦穗部侵染过程的细胞学研究．植物病理学报，2004，34(4)：329-335
15．李晓宇，石洁，董金皋．几种玉米弯孢霉叶斑病菌生物学特性的比较．洞北农业大学学报，2002，3：61-64
16．刘悦秋，江幸福，赵和文．园林植物病虫害．北京气象出版社，2001，1-167
17．饶开晴，王晋峰．我国草坪业研究概况及现状分析．四川草原，2004，7：53-56
18．商鸿生等．草坪病虫害及其防治．北京：中国农业出版社，1996
19．商鸿生，吕学农．草坪早熟禾叶枯病的病原真菌鉴定．中国草地，1996，(4)：36-39
20．沈大刚．草坪草种的选择．林业实用技术，2006，2：38-39
21．孙吉雄．草坪绿地规划与建设管理．北京：科学技术文献出版社，2002
22．唐德志，孙毓彬，何苏琴．德雷疫霉侵染甜瓜叶片过程的研究．西北农业学报，1992，01
23．王璧生，戚佩坤．两种草坪真菌性病害的鉴定与防治．广东农业科学，1991，2：28-29
24．王春梅．草坪病虫害防治．延吉：延边大学出版社，2002，4：200-207
25．翁启勇，王青松，何玉仙，林敏，余德亿．福建草坪草病害初报．草业学报，1997，2：70-73
26．翁启勇，余德亿．草坪病虫害．福建科学技术出版社，2000：168-183
27．吴纯仁，刘后利．油菜菌核病菌致病机理的研究Ⅱ．病菌侵染过程的组织病理学．中国油料作物学报，1990，(1)：44-46
28．习平根，李梅辉，吴仕豪，曹俐，薛硕，姜子德．狗牙根草坪草真菌病害的病原菌鉴定．华南农业大学学报，2005，02：31-34
29．习平根，姜子德，戚佩坤．龙舌兰科二种观赏植物的病原真菌鉴定．广西农业生物科学，2005，12：314-318
30．杨旺．草坪病虫害．中国林业出版社，2001，1
31．张定法．玉米弯孢霉叶斑病菌生物学特性的研究．植物病理学报，1997，27(4)：307-308
32．张定法．玉米弯孢霉叶斑病初侵染菌源的研究．河南农业科学，1998，8：24-25
33．张敬泽，胡东维．影响柿树炭疽菌孢子萌发、附着胞形成及致病性的环境因子．植物病理学报，2004，02
34．张天宇．中国真菌志(第十六卷)链格孢属．北京：科学出版社，2003
35．赵杰．ITS序列分析及其在植物真菌病害分子检测中的应用．陕西农业科学，2004(4)：35-37
36．赵美琦，孙明，王慧敏．草坪病害(草坪全景)．北京：中国林业出版社，1999， 131-196
37．赵美琦，孙彦，张青文．草坪养护技术．北京：中国林业出版社，2001，93-120
38．朱灿星．稻瘟病菌与水稻品种相互关系中附着胞行为的观察．浙江大学学报(农业与生命科学版)，1984，03
39．朱云枝，强胜．马唐生防真菌弯孢霉(Curvularia sp.)菌株QZ-200的生物学特性研究．植物病理学报，2003，33(6)：524-529
40. Boedijn K B. Ueber einige phragmosporen Dematiazeen. Bull. Jard Bot Buitenzorg (Ser 3) 1933, 13: 120-134
41. Brown G E, Cole H J, and Nelson R R. Pathogenicity of Curvularia spp. to turfgrass. Plant Disease, 1972, 56: 59-63
42. Busch L V, Walker J C. Studies of cucumber anthracnose. Phytopathology, 1958, 48: 302-304
43. Carpita N, Gibeaut D. Structural models of primary cell walls in flowering plants: consistency of molecular structure with the physical properties of the walls during growth. Plant J, 1993, 3:1-3
44. Clemens B, Karl F G, Martin S, Paul L, Holger B D, Martin. Optical Measurements of Invasive Forces Exerted by Appressoria of a Plant Pathogenic Fungus. Science, 1999:1896-1899
45. Cooper R M. The mechanisms and significance of enzymatic degradation of host cell walls by parasites. Callow J A. Biochemical Plant Pathology. New York: John Wiley &. Sons Ltd, 1983:101-135
46. Deising H B, Werner S, Wemitz M. The role of fungal appressoria in plant infection. Microbes Infect, 2000, 2:1631-1641
47. Emmett R W, Parbery D G. Appressoria. Ann Rev Phytopathol, 1975, 13:147-167
48. Ellis M B . Dematiaceous hyphomycetes. Commonwealth Mycological Institute, Kew, Surrey, England, 1971:432-459
49. Hartmann H A, Kahmann R, Biker M. The pheromone response factor coordinates filamentous growth and pathogenic development in Ustilago maydis. EMBO J, 1996, 15:1632-1641
50. Howard F L. Helminthosporium-Curvularia Blights of Turf and Their Cure. Golf Course Rep. 1951, 21(2): 5-9
51. Hosokawa M, Tanaka C , Tsuda M .Conidium morphology of Curvularia geniculata and allied species. Mycoscience, 2003,44: 227-237
52. Huang J B , Zheng L and Hsiang T . First report of leaf spot caused by Curvularia affinis on Festuca arundinacea in Hubei, China. Plant Dis, 2004, 88: 1048
53. Kang Z , Huang L, Buchenauer H . Cytochemistry of cell wall component alterations in wheat roots infected by Gaeumannomyces graminis var.tritici. Journal of Plant Diseases and Protection, 2000,107(4): 337-351
54. Kauraw L P . Effect of Fungicides on the germination, RootlShoot Growth and Incidence of Seed-borne Pathogens in Rice. Central Indian Phytopathology, 1986, (39): 609-610
55. Kawanabe Y . The occurrence and control of turgrass diseases in Japan. Pesticide Information, 1991(59): 5-9
56. Khan A, Hsiang T .The infection process of Colletotrichum graminicola and relative aggressiveness on four turfgrass species. Can.J.Microbiol. 2003,49:433-442
57. Li G Q , Huang H C , Laroche A L ,and Acharya S N . Occurrence and characterization of hypovirulence in the tansclerotial isolate S10 of Sclerotinia sclerotiorum. Mycol. Res. 2003,107:1350-1360
58. Mandokhot R. Chemical changes in maize leaves in response to leaf spot pathogens. Indian Phytopathology, 1979, 32(4):658-660
59. Muchovej J J . Definition of Agrostis palustris Leaf Health at the Time of Infection and Colonization by Curvulatia lunata. Ann. Appl. Biol. 1986, 109: 249-258
60. Pietro D A , Roncero M I G. Cloning, expression, and role in pathogenicity of pg1 encoding the major extracellular endopolygalacturonase of the vascular wilt pathogen Fusarium oxysporum. Mol Plant-Microbe Interact, 1998, 11: 91-98
61. Shaw-ming yang. Isolation and effect of temperature on spore germination, radial growth, and pathogenicity of Curvularia senegalensis. Phytopathological notes, 1973,63:1541-1545
62. Sherwood R T, Vance C P . Resistance to Fungal Penetration in Gramineae. Cytology and Histology, 1980(70)4: 273-278
63. Smiley W R , Dernoeden P H , Clarke B B . Compendium of turfgrass diseases. Second Edition. The American Phytopathological Society, Minneapolis, Minnesota, USA, 1992: 1-296
64. Smith J D , Jackson N , Woolhouse A R . Fungal diseases of amenity turf grasses. Third Edition. New York, USA: E&F.N. Spon, 1989: 1-38
65. Talbot N J . Fungal biology-coming up for air and sporulation. Nature, 1999, 398: 295-296
66. Tonukari N J, Scott-Craig J S , Walton J D . The Cochliobolus carbonum SNF1 gene is required for cell wall-degrading enzyme expression and virulence on maize. Plant Cell, 2000,12:237-248
67. Tucker S L , Talbot N J . Surface attachment and pre-penetration stage development by plant pathogenic fungi. Ann Rev Phytopathol, 2001, 39: 385-417
68. Vloutoglou I, Kalogerakis S N. Effects of inoculum concentration, wetness duration and plant age on development of early blight and on shedding of leaves in tomato plants. Plant Pathology, 2000,49:339-345
69. Wernham C C , Kirby R S . A New Turf Disease. Phytopathology, 1941, 31: 24
70. Walton J D . Deconstructing the cell wall. Pl. Physiol, 1994,104(4): 1113-1118
71. Yildiz F . The preliminary sdudies on turgrass diseases in Turkey. Journal of Turkish Phytopathology, 1990,19(1): 21-29