摘要
马铃薯黑痣病是内蒙古地区的主要病害之一,研究马铃薯抗黑痣病鉴定技术及抗病机制对马铃薯抗病育种及其防治具有重要的理论和实践意义。本试验以28份马铃薯为材料系统地研究了马铃薯抗黑痣病鉴定技术,同时对马铃薯材料进行了抗性评价,并从马铃薯形态、结构及生理生化方面探讨其抗黑痣病机制,结果如下：
1.从马铃薯地下茎黑痣病病样上分离到1个菌株,致病性测定结果显示该菌为致病菌。经鉴定该菌为立枯丝核菌AG2-1(Rhizoctonia solani AG2-1)。
2.立枯丝核菌AG2-1能产生致病毒素,为致病的关键因子,但低浓度的毒素对马铃薯幼苗生长有促进作用。
3.初步建立了马铃薯黑痣病抗性鉴定技术体系,首先利用薯片接种和毒素处理马铃薯脱毒组培苗的方法进行初步鉴定,再以麸皮培养物2～4g田间单株接种鉴定。
4.28份马铃薯材料的抗性鉴定结果显示,地下茎和块茎的抗性具有相关性,品种间抗性存在显著性差异,但没有免疫和高抗的品种,品种底西芮表现较高的抗病性,而夏坡蒂、大西洋、费乌瑞它等多数品种表现为感病。生育期长的品种较抗病,生育期短的品种感病。红皮等马铃薯表皮颜色较深的品种较抗病,白皮品种感病。
5.病原菌侵染过程的研究表明,侵染块茎、地下茎和地上茎的的最初侵染时间分别为4h、8h、8～12h。侵染过程中产生少量疏松的侵染垫及大量的附着胞,附着胞有二叉状、球状、三叉状、足状及不规则形状,在附着胞的前端有明显的侵染钉。地上茎产生的侵染结构最多,地下茎和块茎相对较少。病原菌侵染块茎的途径主要是角质层裂缝、皮孔、细胞间隙,也可从伤口及直接侵入；对地下茎的侵染途径主要是表皮裂缝、皮孔、细胞间隙；地上茎主要是通过细胞间隙或直接侵入,少量菌丝从气孔侵入。
6.感病品种大西洋比抗病品种底西芮侵染结构多,侵入组织内的菌丝也多,皮孔大而多、角质层薄、表皮较粗糙、蜡质含量较少。
7.黑痣病菌毒素可诱导马铃薯幼苗体内POD、PPO及SOD活性升高,在毒素处理24～96h,3种酶活性与马铃薯品种抗病性正相关。毒素也能诱导CAT、PAL、几丁质酶、β-1,3-葡聚糖酶活性升高,但几种酶活性的变化和品种的抗病性不相关。毒素处理后马铃薯幼苗体内木质素和游离脯氨酸含量增加,且抗病品种增加值明显多于感病品种。毒素处理后马铃薯体内MDA含量增加,96h以内抗病品种增加幅度大于感病品种。毒素伤害马铃薯幼苗,使其细胞膜透性增加,感病品种的细胞膜透性增加的较早且幅度较大。
8.对马铃薯体内的营养物质和抗黑痣病关系研究表明,可溶性淀粉含量和抗病性正相关,可溶性蛋白含量和抗病性负相关,可溶性糖及游离氨基酸和品种的抗病性没有明显的相关性。
The stem canker and black scurf in potato by Rhizotonia solani is one of the main diseases in Inner Mongolia. The studies on identification technology and resistance mechanism were important in theory and practice. The resistance and identification technology on28different potato varieties to R. solani were studied. The resistance mechanism were research from potato structure and physiological and biochemical. The results were as follows:
1. The pathogen of stem canker and black scurf by R. solani on potato was isolated from underground stems. The isolation had pathogenicity to potato. It's identification result showed that it was R. solani AG2-1.
2. The toxin of R. solani AG2-1had pathogenicity to potao seedling. It was a key pathogenic factor. But the low concentrations toxin can promote seedlings growth of potato.
3. The resistance identification technology system was established to stem canker and black scurf in potato. The ways of potato pieces inoculated with R. solani and seedlings treated with toxin can identify preliminaryly. Then the tests of inoculating with wheat bran inoculum2～4g per potato seed were conducted in field.
4. The resistance of the28potato cultivars was significantly different to R. solani AG2-1but none of these was completely resistant. Most of them were susceptible. The cultivar Diseree showed the less percentage of infection by R. solani, while the cultivars Atlantic, Favorita, Shepody showed a high percentage of infection on tubers.The potato varieties of long growing period and the deeper epidermis color were resistant than the ones of short growth period and white skin.
5. In the infection process of pathogen, the initial infection time were4h,8h and8～12h at tubers, underground stems and stems. The infection structure were a few loose infection cushion, most were appressorium, there were lobate, spherical, trident form, foot and irregular shape, infection pegs were formed from the appressorium. There were more infection structure at stems than the underground stems and tubers.The infection site were mainly cuticle cracks, lenticels, cell gap, wound and a small amount of mycelium direct invasion at tubers. At underground stem the infection site were mainly crack, lenticels, epidermal cell gap and a small amount of hyphae direct invasion. At stems the invasion site were mainly the intercellular and direct invasion breakthrough epidermal, a small amount of mycelium from the stoma invading.
6. Susceptible cultivars Atlantic had more infection structures, larger and more lenticels, thin cuticle, rough epidermis, less wax than the resistant varieties Diseree.
7. The physiological and biochemical resistance mechanism showed that the toxin of R. solani AG2-1can increase the POD, PPO and SOD activity in potato seedlings, and the several enzyme activitys of resistance varietie were all higher than the susceptible one, at24-96h, The CAT, PAL, chitinase, β-1,3-glucanase activity were increased also, but the changes and resistance were not related. The content of proline and Lignin were all increased after inoculating and the resistance cultivar was greater than the susceptible one. The MDA content and cell membrane permeability were all increased also. O～96h, MDA content of susceptible cultivar was greater than the resistance one. The cell membrane permeability of susceptible varieties increased earlier and more.
8. The relations on nutrition of potato and resistance showed that the soluble starch content and resistance was positive correlation, on the other hand, the soluble protein content was negatively correlation with the resistance, soluble sugar and amino acid had no obvious correlation with the resistance.
引文
1 Hadi M R, Balali G, R. The effect of salicylic acid on the reduction of Rizoctonia solani damage in the tubers of marfona potato cultivar[J]. American-Eurasian J Agric & Environ Sci,2010,7 (4):492-496
2 屈冬玉,金黎平,谢开云,等.中国马铃薯产业现状、问题和趋势.陈伊里主编.马铃薯产业与西部开发[M].哈尔滨：哈尔滨工程大学出版社,2001,1-8
3 谢开云,屈冬玉,金黎平,等.中国马铃薯生产与世界先进国家的比较[J].世界农业,2008, 349(5)： 35-41
4 屈冬玉,金黎平,谢开云.中国马铃薯产业10年回顾[M].北京：中国农业科学技术出版社,2008
5 Kuhn J G. Die Krankheiten der kulturegewachse, ihre ursachen und ihre verhutung, gustav bosselmann[J]. Berlin,1858,312
6 Abe H, Tsuboki K. Anastomosis groups of isolates of Rhizoctonia solaoni Kuhn from potatoes [J]. Bulleting of Hokkaido Prefectural Agricultural Experiment Station, 1978,40:61-70
7 Bains P S, Bisht V S. Anastomosis group identity and virulence of Rhizoctonia solaoni isolates collected from potato plants in Alberta, Canada [J]. Plant Disease、1995, 79:241-242
8 Balali G R, Neate S M, Scott E S, et al. Anastomosis group and pathogenicity of isolates of Rhizoctonia solani from potato crops in South Australia [J]. Ptant Pathotogy,1995,44:1050-1057
9 Rolfs F M. Potato failures. A preliminary report[J]. Colo Agri Expt Sta Bull,1902, 70:1-12
10 Campion C, Chatot C, Perraton B, et al. Anastomosis groups, pathogenicity and sensitivity to fungicides of Rhizoctonia solani isolates collected on potato crops in France [J]. European Journal of Plant Pathology,2003,109:983-992
11 Chand T, Logan C. Cultural and pathogenic variation in ptato isolates of Rhizoctonia solani in Northern Ireland[J]. Transactions of British Mycological Society,1983, 81:585-589
12 Calleros V G, Portugal 0 V, Carling D E. Anastomosis groups of Rhizoctonia solani on potato in centrai Mexico and potential for biological and chemical control [J]. American Journal of Potato Research,2000,77:219-224
13 ELBakali A M, Martin M P. Black scurf of potato[J]. Mycologist,2006,20:130-132
14 Carling DE, Leiner R H,Westphale P C. Symptoms, signs and yield reduction associated with Rhizoctonia disease of potato induced by tuberborne inoculum of Rhizoctonia solani AG-3 [J]. American Potato Journal,1989.66:693-701
15 Weinhold A R, Bowman T, Hall D H. Rhizoctonia disease of potato:effect on yield and control by seed tuber treatment [J]. Plant Disease,1982,66(9):815-818
16 Banville G J. Yield losses and damage to potato plants caused by Rhizoctonia solani Ktihn[J]. American Potato Journal,1989,66:821-834
17 Scholte K. Effects of soil-borne Rhizoctonia solani Kiahn on yield and quality of ten potato cultivars [J]. Potato Research,1989,32:367-376
18 Simons S A, Gilligan C A. Relationships between stem canker, stolon canker, black scurf(Rhizoctonia solani) and yield of potato (Solanum tuberosum) under different agronomic conditions [J]. Plant Pathology,1997,46:651-658
19邱广伟,马铃薯黑痣病的发生与防治[J].粮食作物,2009,6：]33-134
20徐鸿皋.台湾马铃薯黑痣病之初步研究[J].植物病理学报(中兴大学),1978,7：33-48
21 Wu W S. Evaluation of the effectiveness of four fungicides on controlling potato black scruf disease[J]. Plant Prot Bull (Taiwan),1978,20:253-259
22张义璋,杜金池Rhizoctonia solani各菌丝融合群对马铃薯之侵害情形[J].中国农业研究,1980,29(1)：27-33
23霍茂林.要注意防治马铃薯丝核菌病[J].现代农业,1988(4)：26
24 田晓燕,蒙美莲,张笑宇,等.马铃薯黑痣病菌菌丝融合群的鉴定[J].中国马铃薯,2011,25(5)：298-301
25 刘宝玉,胡俊,蒙美莲,等.马铃薯黑痣病病原菌分子鉴定及其生物学特性[J].植物保护学报,2011,38(4)：379-380
26 刘宝玉,蒙美莲,胡俊,等.5种杀菌剂对马铃薯黑痣病的病菌毒力及田间防效[J].中国马铃薯,2010,24(5)：306-310
27 曹春梅,张智芳,李文刚,等.新型杀菌剂对马铃薯黑痣病病菌的室内毒力测定和田间效果分析[J].中国马铃薯,2011,25(4)：246-250
28 Woodhall J W, Lees A K, Edwards S G, et al. Infection of potato by Rhizoctonia solani: effect of anastomosis group[J]. Plant Pathology,2008,57:897-905
29 Carl ing D E, Leiner H R. Isolation and characterization of Rhizoctonia solani and binucleate R. solani-like fungi from aerial stems and subterranean qrgans of potato plants [J]. Phytopathology,1986,76(7):725-729
30 Lehtonen M J, Wilson P S, Ahvenniemi P, et al. Formation of canker lesions on stems and black scurf on tubers in experimentally inoculated potato plants by isolates of AG2-1, AG-3 and AG-5 of Rhizoctonia solani:apilot study and literature review[J]. Agricultural and Food Science,2009,18:223-233
31 Anguiz R, Martin C. Anastomosis groups, pathogenicity, and other characteristics of Rhizoctonia solani isolated from potatoes in Peru[J]. Plant Disease,1989,77: 199-201
32 Carling D E, Brainard K A, Virgen C G, et al. First report of Rhizoctonia solani AG7 on potato in Mexico[J]. Plant Disease,1998,82:127
33 Hide G A, Firmager J P. Effects of an isolate of Rhizoctonia solani Kiihn AG8 from diseased barley on the growth and infection of potatoes (Solanum tuberosum) [J]. Potato Research,1990,33:229-234
34陈延熙,张敦华,段霞渝,等.关于Rhizoctoniasolani菌丝融合分类和有性世代的研究*[J].植物病理学报,1985,15(3)：39-43
35 Adams G C Jr, Butler e e. influence of nutrition on the formation of basidia and basidio spores in Thanatephorus cucumeris[J]. The American Phytopathological Society,1983,73 (2):147-151
36 Gerard C A, Adams J R, Bustler E E. Environmental on the formation of basidia and basidio spores in Thana tephorus cucumeris[J]. The American Phytopathologycal Society.1983,73 (2):152-155
37 Jeger M J, Hide G A, Van den Boogert P, et al. Pathology and control of soil-borne fungal pathogens of potato[J]. Potato Research,1996,39:437-469
38 Parmeter J R Jr, Whitney H S, Platt W D. Affinities of Rhizoctonia species that resemble mycelium of Thanatephorus cucumeris[J]. Phytopathology,1967,57:218-223
39 Parmeter J R Jr, Sherwood R T, Platt W D. Anastomosis grouping among isolates of Thanatephorus cucumeris[J]. Phytopathology.1969,59:1270-1277
40 Ogoshi A. Ecology and pathogenicity of anastomosis and intraspecific groups of Rhizoctonia solani kuhn[J]. Ann Rev Phytopathol,1987,25:125-143
41 李菊.中国东北地区玉米纹枯病菌融合群鉴定及遗传多样性研究[D].泰安：山东农业大学,2011
42 Jin M S, Korpradiskul V. Isozyme analysis of genetic diversity among Rhizoctonia solani anastomosis groupl-IA(AG1-IA) [J]. Mycosystem,1998,17(4):331-338
43王洪凯,刘开启,吴洵耻.丝核菌分类研究进展[J].山东农业大学学报,1997,28(3)：375-381
44 Kuninaga S, Carling D E, Takeuchi T, et al. Comparison of rDNA-ITS sequences between potato and tobacco strains of Rhizoctonia solani AG3[J]. Journal of General Plant Pathology,2000,66:2-11
45 Yanar Y, Yllmaz G, Cesmeli I, at al. Characterization of Rhizoctonia solani isolates from potatoes in Turkey and screening potato cultivars for resistance to AG-3 isolates [J]. Phytoparasitica,2005,33(4):370-376
46 Rauf C A, Ahmad I, Ashraf M. Anastomosis groups of Rhizoctonia solani K u hn isolates from potato in Pakistan[J]. Pak J Bot,2007,39(4):1335-1340
47 Carl ing D E, Leiner R H, Kebler K M. Characterization of Rhizoctonia solani and binucleate Rhizoctonia-like fungi collected from Alaskan soils with varied crop histories [J]. Canadian Journal of Plant Pathology,1986,8:305-310
48 Cedeno L, Carrero C. Quintero K, et al. Identificacion y virulencia de grupos deanastomosis de Rhizoctonia solani Kuhn asociados con papa en Merida, Venezuela [J]. Interciencia,2001,26:296-300
49 Woodhall J W, Lees A K, Edwards S G, et al. Characterization of Rhizoctonia solani from potato in Great Britain[J]. Plant Pathology,2007,56:286-295
50 Bandy B P, Leach S S, Tavantzis S M. Anastomosis group 3 is the major cause of Rhizoctonia disease of potato in Maine [J]. Plant Disease,1988,72(7):596-598
51 Justesen A F, Yohalem D, Bay A, et al. Genetic diversity in potato field populations of Thanatephorus cucumeris AG-3, revealed by ITS polymorphism and RAPD markers [J]. Mycological Research,2003,107:1323-1331
52 Truter M, Wehner F C. Anastomosis grouping of Rhizoctonia solani associated with black scurf and stem canker of potato in South Africa [J]. Plant Disease,2004,88(1): 83
53 Mohammadi R B, Salari M, Zafari D. Anastomosis groups of Rhizoctonia solani from potato and survey of somatic compatible groups by morphological characters and rapd marker[J]. Iran J Plant Path,2010,45(4):87-89
54 Richards B L. Pathogenicity of Corticium vagum on the potato as affected by soil temperature [J]. J Agric Res,1921,21:459-482
55 Ritchie F, Bain R A, McQuilken M P. Effects of nutrient status, temperature and pH on mycelial growth, sclerotial production and germination of Rhizoctonia Solani from potato [J]. Journal of Plant Pathology,2009,91(3):589-596
56 Car ling D E, Leiner R H. Effect of temperature on virulence of Rhizoctonia solani and other Rhizoctonia on potato [J]. Phytopathology,1990,80:930-934
57 Grosch R, Faltin F, Lottmann J, et al. Effectiveness of 3 antagonistic bacterial isolates to control Rhizoctonia solani Kuhn on lettuce and potato [J]. Canadian Journal of Microbiology,2005,51:345-353
58 Doornik AW. Some factors affecting the parasitic and saprophytic activity of Rhizoctonia solani[J]. Acta Horticulturae,1980,109:387-394
59 张天晓Rhizoctonia solani菌丝融合群的研究[J].湖南师院学报,1984,(2)：69-72
60 Maniruzzaman M K, Khair A, Khurshed M A B. Disease reaction of different crops against virulent potato isolates of Rhizoctonia Solani Kuhn [J]. Bangladesh J Bot,2008,37(1): 75-80
61 张爱萍.中国部分地区草坪丝核菌的融合群鉴定及生防木霉菌的筛选[D].泰安：山东农业大学,2010
62 Date H, Yagi S, Okamoke Y, et al. On the leaf blight of tomatoes by Thanatephorus cucumeris (Frank) Donk(Rhizoctonia Solani) [J]. Ann Phytopathol Soc Jpn,1984,50: 339
63 Kodama T, Horimota K, Ogoshi A. On the brown spot of egglants caused by Thanatephorus cucumeris (Frank) Donk(Rhizoctonia Solani) AG-3[J]. Ann Phytopathol Soc Jpn,1982, 48:356
64 Stevens J J, Jones R K, Shew H D, et al. Characterization of populations of Rhizoctonia solani AG-3 from potato and tobacco. Phytopathology[J].1993,83:854-858
65 James W C, McKenzie A R. The effect of tuber-borne sclerotia of Rhizoctonia solani Kuhn on the potato crop[J]. Am Potato J,1972,49:296-301
66 Hide G A, Hirst J M, Stedman O J. Effects of black scurf(Rhizoctonia solani) potatoes[J]. Ann Appl Biol,1973,74:139-148
67 Gudmestad N C, Zink R T, Huguelet J E. The effect of harvest date and tuber-borne sclerotia on the severity of Rhizoctonia disease of potato [J]. Am Potato J,1979, 56:35-41
68 Tsror L, Peretz A I. The influence of the inoculum source of Rhizoctonia solani on development of black scurf on potato[J].Phytopathology,2005,153:240-244
69 Frank J A, Leach S S. Comparison of tuberborne and soilborne inoculum in the Rhizoctonia disease, of potato[J]. Ecology and Epidemiology,1980,70(1):51-53
70 Atkinson D, Thornton M K, Miller j S. Development of Rhizoctonia solani on stems, stolons and tubers of potatoes I. Effect of inoculum sourcelJ]. Am J Pot Res,2010, 87:374-381
71 Simons S A. Gilligan C A. Factors affecting the temporal progress of stem canker (Rhizoctonia solani) on potatoes(Solanum tuberosum) [J]. Plant Pathology,1997, 46:642-650
72 Adams M J, Hide G A. Relationships between disease levels on seed tubers, on crops during growth and in stored potatoes.5. Seed stocks grown at Rothamsted [J]. Potato Research,1980,23:291-302
73 Hide G A, Read P J, SandisonJ. Stem canker(Rhizoctonia solani) of maincrop potatoes; I. Development of the disease[J]. Annals of Applied Biology,1985,106:413-422
74 Naz F, Rauf C A, Abbasi N A, et al. Influence of inoculum levels of Rhizoctonia solani and susceptibility on new potato germplasm[J], Pakistan Journal of Botany,2008, 40(5):2199-2209
75 Djebali N, Belhassen T. Field study of the relative susceptibility of eleven potato (Solahum tuberosum L.)varieties and the efficacy of two fungicides against Rhizoctonia solani attack [J]. Crop Protection,2010,29:998-1002
76 Bains P S, Bennypaul H S, Lynch D R, et al. Rhizoctonia disease of potatoes (Rhizocionia solani):fungicidal efficacy and cultivar susceptibility[J]. American Journal of Potato Research,2002,79:99-106
77 Olanya O M, Lambert D H, Reeves A F, et al. Evaluation of potato clones for resistance to stem canker and tuber black scurf in field studies following artificial inoculation with Rhizoctonia solani AG-3 in Maine [J]. Archives of Phytopathology and Plant Protection,2009,42(5):409-418
78 Leach S S, Webb R E. Evaluation of potato cultivars, clones and a true seed population for resistance to Rhizoctonia solani[J], American Potato Journal,1993,70:317-328
79 Otrysko B E, Banville G J. Effect of infection by Rhizoctania solani on the quality of tubers for processing[J]. Am Potato J,1992,69:645-652
80 Errampalli D, Johnston H W. Control of tuber-borne black scurf(Rhizoctonia solani)and common scab(Streptomyces scabies) of potatoes with a combination of sodium hypochlorite and thiophanate-metnyl preplanting seed tuber treatment [J]. Can J Plant Pathol,2001,23:68-77
81 Secor G A, Gudmestad N C. Managing fungal diseases of potato [J]. Can J Plant Pathol, 1999,21:213-221
82 Wilson P S, Ketola E 0, Ahvenniemi P M, et al. Dynamics of soilborne Rhizoctonia solani in the presence of Trichoderma harzianum:effects on stem canker, black scurf and progeny tubers of potato[J]. Plant Pathology,2008,57:152-161
83 Gajda I, Kurzawi ska H. Biological protection of potato against Helminthosporium solani and Rhizoctonia solani[1]. Phytopathol Pol,2004,34:51-58
84 Beagle-Ristaino J E, Papavizas G C. Biological control of Rhizoctcnia stem canker and black scurf of potato[J].Phytopathology,1985,75(5):560-564
85 Lahlali R, Hi jri M. Screening, identification and evaluation of potential biocontrol fungal endophytes against Rhizoctonia solani AG3 on potato plants [J]. Ferns Microbiol Lett,2010,311:152-159
86 Demirci E, Eken C, Dane E. Biological control of Rhizoctonia solani on potato by Verticillium biguttatum[J]. African Journal of Biotechnology,2009,8 (11): 2503-2507
87 Tomilova 0 G, Shternshis M V. The effect of a preparation from Chaetomium fungi on the growth of phytopathogenic fungi [J]. Applied Biochemistry and Microbiology,2006, 42(1):67-71
88 Kurzawinska H, Mazur S. Biological control of potato against Rhizoctonia solani (K u hn) [J]. Sodininkyste Ir Dar ininkyste,2008,27(2):419-425
89 Bautista G, Mendoza H, Uribe D. Biocontrol of Rhizoctonia solani in native poiato (Solanum phureja) plants using native Pseudomonas fluorescens[1]. Acta Biol Colomb, 2007,12(1):19-32
90 Brewer M T, Larkin R P. Efficacy of several potential biocontrol organisms against Rhizoctonia solani on potato[J]. Crop Protection,2005,24:939-950
91 Shahrokhi S, Shahidi B G H, Saadoun I. Biological control of potato isolate of Rhizoctonia solani by Streptomyces olivaceus Strain 115 [J]. Biotechnology,2005, 4(2):132-138
92 Tsror(Lahkim)L, Barak R, Sneh B. Biological control of black scurf on potato under organic management [J]. Crop Protection,2001,20:145-150
93 Bandy B P, Tavantzis S M. Effect of hypovirulent Rhizoctonia solani on Rhizoctonia disease, growth and development of potato plants [J]. American Potato Journal,1990, 67:189-199
94 Larkin R P, Honeycutt CW. Effects of different 3-year cropping systems on soil microbial communities and Rhizoctonia disease of potato[J]. Phytopathology,2006, 96(1):68-79
95许志刚主编.普通植物病理学(第三版)[M].北京：中国农业出版社,2003,297-299
96 Abbas H K, Boyette C D. Bioherbicidal patential of Fusarium monoliforme and itsphytotoxin fumonisin[J].1991, Weed Science,39(4):673-677
97 Aoki H, Sassa T, Tamura T. Phytotoxic metabolites of Rhizoctonia solani[J]. Nature, 1963,200:575
98 Brooks S A. Sensitivity to a phytotoxin from Rhizoctonia solani correlates with sheath blight susceptibility in rice[J]. Phytopathology,2007,10(97):1027-1212
99 Lakpale N, Rajiv K, Kumar R, et al. Studies on toxin produced by Rhizoctonia solani causing sheath blight of rice[J]. Indian Journal of Mycology and Plant Pathology, 1996,26(3):263-265
100 Mandava N B, Orellana R C, Warthen D J, et al. Phytotoxins in Rhizoctonia solani:Isolation and biological activity of m-hydroxy and m-methoxypheny lacetic acids [J]. Joural of Agricultural Food Chemistry,1980,28:71-75
101 Paranidharan V, Palaniswami A, Vidhyasekaran P, et al. Host-specific toxin of Rhizoctonia solani triggers superoxide dismutase (SOD) activity in rice [J]. Archives of Phytopathology and Plain Protection,2005,38(2):151-156
102 Ramalingam P. Involve of Rhizoctonia solani toxins in phenylalanine ammonia lyase and tyrosine ammonia lyase activities of rice tissues [J]. Current Science, India, 1986,55(3):157-159
103 Sriram S, Raguchander T, Babu S, et al. Inactivation of phytotoxin produced by the rice sheath blight pathogen Rhizoctonia solani[J]. Canadia Journal Microbiology, 2000,46(6):520-524
104 Vidhyasekaran P, Ponmalar T R, Samiyappan R, et al. Host-specific toxin production by Rhizoctonia solani, the rice sheath blight pathogen [J].1997,87(12):1258-1263
105 Vidhyasekaran P, Ponmalar T R, Samiyappan, et al. Rice sheath blight produces host-specific toxin[J]. Rice-Biotechnology-Quarterly,1998,35:21
106 Wang Y Z, Miller J D. Effects of fusarison graminearaan metabolites on wheat tissue in relation to fusarium head blight resistance[J]. Phytopathology,1988,122: 118-125
107暴增海.植物病原菌毒素的利用现状[J].张家口农专学报,2000,16(1)：37-38
108陈怀谷.禾谷丝核菌的胞外产物及其致病机理的研究[D].南京：南京农大学,1999
109陈捷,唐朝荣,邹庆道,等.玉米纹枯病菌致病因子的研究[J].沈阳农业大学学报,1999,30(3)： 189-194
110董金皋.寄主选择性真菌毒素与植物病害特异性[A].董金皋,李树正.植物病原真菌毒素研究(第一卷)[C].北京：中国科学技术出版社,1997
111康绍兰,董金皋,张汀,诱发小麦抗根腐病突变体的研究,Ⅱ影响小麦根腐菌致病毒素作用的因素与生物测定初报[J].河北农业大学学报,1991,14(2)54-59
112刘国胜,黄梧芳.玉米小斑病菌致病毒素研究近况[J].1991,17(5)：32-34
113刘新琼.小麦赤霉病菌毒素研究进展[J].1997,湖北植保,3：23-24
114马力功营养及培养条件对大豆根腐病主要病原菌及其产毒的作用[J].2008,东北农业大学
115孟令军.水稻纹枯病菌毒素的提纯、组分分析及钝化[J].扬州：扬州大学,2009
116王裕中,陈怀谷,杨新宁.禾谷镰刀菌粗毒素的生物活性及其在小麦品种抗赤霉病性鉴定中的应用[J].中国农业科学,1989,22(4)：54-57
117徐艳.水稻纹枯病菌毒素的致病机理及对寄主防御酶活性的影响[J].扬州：扬州大学,2006
118叶建仁,杨斌,包宏.松针褐斑病菌毒素活性的生物检测[J].南京林业大学学报(自然科学版),2002,26(1)：27-31
119赵小明,杨家荣,吕金殿.应用棉花黄萎病菌致萎毒素鉴定品种抗病性初探[J].陕西农业科学,1998,6：14-15
120 Adams M J, Hide G. A, Lapwood D H. Sampling potatoes for the incidence of tuber diseases and levels of inoculum[J]. Annals of Applied Biology,1985,107:189-203
121 Adams M J, Hide G A, Lapwood D H. Relationships between disease levels on seed tubers, on crops during growth and in stored potatoes:1. Introduction and black scurf [J]. Potato Research,1980,23,201-204
122 Johnk J S, Jones R K, Shew H D, et al. Characterization of populations of Rhizoctonia solani AG-3 from potato and tobacco [J]. Phytopathology,1993,83:854-858
123 Rosenbaum G T, Sando J J. Binary pathway for early infection process by Fusarium in Piants [J]. Can J Bot,1993,62:1232-1244
124 Steinkamp M P, Martin S S, Hoefert L L. infrastructure of leisons produced by Cercospora bsticola in leaves of BetaVulgaris[J]. Physiological Plant Pathology, 1979,15(1):13-16
125康立功,齐凤坤,许向阳.番茄叶片蜡质和角质层与芝麻斑病菌侵染的关系[J].中国蔬菜,2010,(18)47-50
126 Seedbold K, Datnoff L, Correa F, et al. Effects of silicon and host resistance on leaf and neck blast in upland rice[J]. (Annual Meeting of the American Phytopathological Society, Las veges, Nevada, USA, November 8-12,1993). Phytopathol, 1998,88(9):80
127 Belanger R R, Bowen P A, Ehret D L, et al. Soluble silicon:its role in crop and disease management of greenhouse crops [J]. Plant Disease,1995,79(4):329-336
128柯玉诗,黄小红,张壮塔,等.硅肥对水稻氮磷钾营养的影响及增产原因分析[J].广东农业科学,1997,(5)：25-27
129 Marschner H. Mineral nutrition of higher plant[J]. San Diego:Academic Press Inc, 1995,289-306,417-427
130 Kim S G, Kim K W, Park E W et al. Silicon-induced cell wall fortification of rice leaves:a possible cellular mechanism of enhanced host resistance to blast [J]. Phytopathology,2002,92(10):1095-1103
131孙万春,薛高峰,张杰.硅对水稻防御性关键酶活性的影响及其与抗稻瘟病的关系[J].植物营养与肥料学报,2009,15(5)：1023-1028
132李广旭,沈永波,高艳敏.皮孔组织结构及密度与苹果枝干粗皮病发生的关系[J].果树学报,2004,214：350-353
133李淼,檀根甲,李瑶,等.猕猴桃品种枝条组织结构与抗溃疡病关系的初步研究[J].安徽农业大学学报,2003,30(3)：240-245
134陶家凤.立枯丝核菌对水稻侵染过程的研究[J].四川农业大学学报,1992,10(3)：471-477
135 Matsuura K. Scanning electron microscopy of the infection process of Rhizoctonia solani in leaf sheath of rice plains [J]. Phytopathology,1986,76:811-814
136 Marshall D S, Rush M C. Infection cushion formation on rice sheaths by Rhizoctonia solanil[J]. Phytopathology,1980:70:947-950
137张红生,朱立宏,沙学延,等.水稻纹枯病抗性机理的初步研究[A].朱立宏.主要农作物抗病性遗传研究进展[C].南京：江苏科学技术出版社,1990.153-164
138 Demirci E, Doken M T. Host penetration and infection by the anastomosis groups of Rhizoctonia solani Kuhn isolated from potatoes[J]. Tr. J. of Agriculture and Forestry,1998,22:609-613
139张国良,闫大伟,何祖华.水稻纹枯病菌侵染过程的细胞学特征[J].细胞生物学杂志,2010,32(3)：451-455
140刘雪梅,肖建国.小麦纹枯病菌侵染过程的组织学研究[J].菌物系统,1999,18(3)：288--293
141陶家凤,谭方河.立枯丝核菌侵染玉米的研究[J].植物病理学报,1995,25(3)：253-257
142 Marshall D S, Rush M C. Relation between infection by Rhizoctonia solani and R. oryzae and disease severity in rice[J]. Phytopathology,1980,70:941-946.
143 Kousik C S, Snow J P, Berggren G T. Factors affecting infection cushion development by Rhizoctonia solani AG-1 IA and IB on soybean leaves [J]. Plant Pathology,1994, 43:237-244
144童蕴慧,徐敬友,潘学彪,等.水稻植株对纹枯病菌侵染反应及其机理的初步研究[J].江苏农业研究,2000,21(4)：45-47
145陈志宜,.王玉环,殷尚智.水稻纹枯病抗性机制的研究[J].中国农业科学,1992,25(4)：41-46
146 Massaquoi R C, Rush M C. Relationship of quantity of epiciticular wax to resistance of rice to sheath blight[J]. Phytopathology,1987,77(12):17-23
147 Selitrennikoff C P. Antifungal proteins[J]. Appl Environ Microbiol,2001,67: 2883-2894
148王勇刚,曾富华,吴志华,等.植物诱导抗病与病程相关蛋白[J].湖南农业大学学报(自然科学版),2002,28(2)：177-182
149左豫虎,康振生,杨传平,等.β-1,3-葡聚糖酶和几丁质酶活性与大豆对疫霉根腐病抗性的关系[J].植物病理学报,2009,39(6)：600-607
150 Chinnusaray V, Jagendorf A, Zhu J K. Understanding and improving salt tolerance in plants [J]. Crop Scienee,2005,45:437-448
151郝建军.植物生理学实验技术[M].北京：化学工业出版社,2007,175-177
152夏平.马铃薯植株感染Y病毒后生理指标变化与抗病性的关系[J].中国马铃薯,2007,21(3)：129-133
153赵小虎,陈翠莲,焦春香,等.不同油菜品种对油菜菌核病敏感性差异的生理生化特性研究[J].华中农业大学学报,2006,25(5)：488-492
154王力先.节瓜枯萎病抗性机理的研究[D].济南：暨南大学,2007
155赵秀娟,张衍荣,胡开林.酶活性、丙二醛含量变化与豇豆抗枯萎病的关系[J].湖南农业大学学报(自然科学版),2009,35(3)：245-248
156薛莲.采后苹果与炭疽菌互作的生理生化机制的研究[D].合肥：安徽农业大学,2005
157石振亚.提高黄瓜植株含糖量预防黄瓜霜霉病[J].河北农学报,1981,6(4)：60-64
158云兴福.黄瓜组织中氨基酸、糖和叶绿素含量与其对霜霉病抗性的关系[J].华北农学报,1993,8(4)： 52-58
159 Hwang B K. Sugar composition and acid invertase activity in spring barley plants in relation to adult-plant resistance to powdery mildew[J]. Phytopathology,1986, 76:365-369
160夏正俊,顾本康,李经仪.棉花品种抗黄萎病性与生化成分相关分析[J].中国农业科学,1991,24(1)： 92
161李淼,檀根甲,李瑶.猕猴桃品种中糖分及木质素含量与抗溃疡病的关系[J].植物保护学报,2005,32(2)：138-142
162陈捷.植物病理生理学[M].辽宁科学技术出版社,1994,1-247
163 Cottrell M T, Suttle C A. Dynamics of a lytic virus infecting the photosynthetic marine picoflagellate[J]. Micromonas Pusilla Limnol Oceanoger,1995,40:730-739
164 Berry J A, Downton W J S. Environmental regulation of photosynthesis. In:Govindjeeed, Photosynthesis(Vol.Ⅱ)[M]. New York:AcademicPress,1982:294-306
165李赤,于莉,刘付东标,等.富贵竹中可溶性糖、蛋白质含量与细菌性茎腐病的关系[J].吉林农业大学学报,2007,29(6)：620-622
166黄世文,王玲,王全永.纹枯病菌对不同水稻品种叶片中抗病性相关酶活性的影响[J].中国水稻科学,2008,22(2)：219-222
167李荣花,陈捷,高增贵,等.玉米纹枯病抗性与防御酶关系的研究[J].天津师范大学学报(自然科学版),2005,25(4)：32-36
168史俊卿,张浩,于红威,等.立枯丝核菌毒素对人参防御酶活性及丙二醛含量的影响[J].湖南农业大学学报(自然科学版),2010,36(4)：434-436
169张穗,刘卫群,陈汝梅,等.不同小麦品种对纹枯病的抗性机理的初步研究[J].中国农学通报,1994,10(6)：6-12
170 Hashiba T, Yamaguchi T, Mogi S. Quantitative changes in nitrogen and starch contents in rice sheaths during vertical disease development of sheath blight caused by Rhizoctonia solani Kuhn [J]. Ann Phytopathol Soc Jpn,1977,43:1
171郑达.玉米纹枯病菌致病性及玉米阶段生理生化因素研究[D].雅安：四川农业大学,1999
172唐莉.玉米纹枯病田间发生动态、阶段抗性的生化因素及种质资源抗性评价研究[J].雅安：四川农业大学,2001
173 Frank J A, Francis S K. The effect of a Rhizoctonia solani phytotoxin on potatoes'[J]. Canadian Journal of Botany,1976,54:2536-2540
174敖世恩,杨媚,周而勋,等.水稻抗纹枯病突变体的离体筛选[J].华南农业大学学报,2006,27(1)：47-50
175潘学彪,陈宗祥,徐敬友.不同接种调查方法对抗水稻纹枯病遗传研究的影响[J].江苏农学院学报,1997,18(3)：27-32
176史建荣,王裕中,陈怀古,等.小麦纹枯病品种抗性鉴定技术及抗病资源的筛选与分析[J].植物保护学报,2000,27(2)：107-112
177 Dowley L J. Varietal susceptibility of potato tubers to Rhizoctonia solani in Ireland[J]. Iranian Journal of Agricultural Research,1972,11:281-285
178 Hide B G A, Read P J, Firmager J P, et al. Stem canker (Rhizoctonia solani) on five early and seven maincrop potato cultivars I. Infection of shoots, stolons and tubers [J]. Ann. appl. Biol,1989,114,255-265
179 Dennis Atkinson, Michael K, Thornton Jeffery S. Miller. Development of Rhizoctonia solani on stems, stolons and tubers of potatoes I. Effect of inoculum source [J]. Am J Pot Res,2010,87:374-381
180 Armentrout V N, Downer A J, Nameth S T. A simplified branching assay for Rhizoctonia solani[J]. Mycologia,1986,78:657-663
181 Di jst G. Formation of sclerotia by Rhizoctonia solani on artificial media and potato tubers[J]. Neth J PI Path,1988,94:233-242
182 Dijst G, Investigations on the effect of haulm destruction and additional root cutting on black scurf on potato tubers [J]. Netherlands Journal of Plant Pathology 1985,91:153-162
183 Allington W B. Sclerotial formation in Rhizoctonia solani as affected by nitritional and other factors[J]. Phytopathology,1936,26:831-844
184左示敏,王子斌,陈夕军,等.水稻纹枯病改良新抗源YSBR1的抗性评价[J].作物学报,2009,35(4)：608-614
185张华东.立枯丝核菌毒素的产生及与致病力的关系[J].扬州,扬州大学,2003
186 Almasia N I, Bazzini A A, Hopp H E, et al. Overexpression of snakin-1 gene enhances resistance to Rhizoctonia solani and Erwinia carotovora in transgenic potato plants [J]. Molecular Plant Pathology,2008,9(3):329-338
187戴伟民,张克勤,段彬伍.测定水稻硅含量的一种简易方法[J].中国水稻科学,2005,19(5)：460-462
188陈捷,唐朝荣,高增贵.玉米纹枯病病菌侵染过程研究[J].沈阳农业大学学报,2000,31(5)：503-506
189 Dodman R L, Flent je N T. The mechanism and physiology by Rhizoctonia solani:Biology and pathology pp.1970,149-160, Uni. California Press, Berkeley.
190 Downer A J, Armentrout V N. The effct of host exudates on infection cushion morphogensis of Rhizoctonia solani (Abstr.) [J]. Phytopathology,1983,73:958
191 Paranidharan V, Palaniswami A, Vidhyasekaran P, et al. Induction of enzymatic scavengers of active oxygen species in rice in response to infection by Rhizoctoni soiani[J]. Acta Physiologiae Plantarum,2003,25(1):91-96
192刘丽,马永毅,张志明,等.玉米不同防卫酶系对纹枯病作用的研究[J].玉米科学,2009,17(3)： 99-102
193 Sancho M A, Forchett M S, Pliego F, et al. Peroxidase activity and isoenzymes in the culture medium of NaCl adapted tomato to suspension cell[J]. Tissue and Organ Culture,1996,44:161-167
194金庆超,叶华智,张敏.苯丙氨酸解氨酶活性与玉米对纹枯病抗性的关系[J].四川农业大学学报,2003,21(2)：116-118
195 Punja Z K. Genetic engineering of plants to enhance resistance to fungal pathogens-e-review of progress and future prospects [J]. Can J Plant Pathol,2001, 23:216-235
196 Esfahani K, Motallebi M, Zamani M R, et al. Transformation of potato(Solatium tuberosum cv.Savalan) by chitinase and β-1,3-glucanase genes of raycoparasitic fungi towards improving resistance to Rhizoctonia solani AG-3[J]. Iranian Journal Of Biotechnology,2010,8(2):73-81
197史俊卿,许永华,张秋菊,等.立枯丝核菌毒素对人参PAL活性及细胞膜透性的影响[J].人参研究,2010,(2)：19-21