蜡蚧轮枝孢的生物学特性及防治温室桃蚜的研究
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摘要
蜡蚧轮枝孢[Verticillium lecanii (Zimm.) Viegas]是一种重要的丝孢类昆虫病原真菌，近年来在国外深受重视，已有多种商品制剂注册生产并用于温室白粉虱、蚜虫等刺吸式口器害虫的防治。本研究对该菌的生物学特性、生理生化因子、双相发酵技术、孢子粉干燥工艺及孢子粉在贮藏过程中内源营养物质代谢和对温室桃蚜防治应用进行了较系统的研究，旨在为利用蜡蚧轮枝孢控制刺吸式口器害虫提供依据。
用11株蜡蚧轮枝孢对桃蚜(Myzus persicae Sulzer)2～3龄若虫进行生物测定。以桃蚜的校正死亡率、感染率及致死中时(LT_(50))为指标综合评价不同菌株毒力。结果表明，11株蜡蚧轮枝孢菌株中，VL07菌株毒力最强，校正死亡率达100％，感染率达93％，致死中时为3.30d。其次是VL18菌株，致死中时为4.02d，校正死亡率和感染率分别为93.83％和84.44％。在所测试温度中，25℃是蜡蚧轮枝孢侵染桃蚜的最适温度，致死中剂量为533孢子／蚜。
温、湿度对蜡蚧轮枝孢生长和发育的影响为：蜡蚧轮枝孢在10～30℃的温度下均可生长，25℃为菌丝生长最适温度，其中VL07、VL18和VL09菌株菌丝生长量较高，14d累计生长量分别为4.69cm、4.98cm和4.72cm；不同菌株在不同温度下产孢量差异极显著，25℃为蜡蚧轮枝孢最适产孢温度，其中VL07和VL18菌株产孢量最高，14d分别达5.83×10~9和4.64×10~9孢子／cm~2；相对湿度对分生孢子萌发试验表明，在RH为100％时，VL07、VL10和VL18菌株分生孢子萌发快而整齐，24h萌发率均达100％，当相对湿度在75％以下时，除VL07和VL18外，其余各菌株的萌发率均降低到20％以下，而且生长缓慢。
测定4种生物源农药与蜡蚧轮枝孢的相容性结果表明：4种农药高浓度下对蜡蚧轮枝孢分生孢子的萌发均有一定的影响，而在低浓度下影响较小，如在2×10~4倍浓度下25％灭幼脲Ⅲ号悬浮剂对高毒力菌株VL07分生孢子的校正抑制率仅为1.66％。因此，可考虑亚致死剂量的灭幼脲Ⅲ号与蜡蚧轮枝孢混合使用以提高防治效果。
对蜡蚧轮枝孢生理生化因子的测定结果表明：蜡蚧轮枝孢的毒力与菌株酯酶总量及前沿带的酯酶量之间呈显著线性相关，相关系数分别为r=0.9198和r=0.8820；蜡蚧轮枝孢毒力与胞外蛋白酶产生水平以及草酸产生水平之间均无明显相关性。
通过对不同的碳、氮源、无机盐、起始pH值、接种量以及正交试验，以菌丝生物量为指标筛选出了适合于VL07菌株工业生产最佳发酵工艺配方，液相发酵培养基
配方为：黄豆粉 4％，白砂糖一％，KH。PO’ 0刀2％，MgSO厂 7HZO 0刀1％，FeSO厂 7H。O
0＊5％，VB。0＊5％，VC 0＊5％，pH值 6．89，初始接种量为 10％－15％；固相发酵培养
基配方为：鼓皮 42％，稻壳 28％，玉米粉 20％，黄豆粉 10％。当培养基初始含水率为
50％时，蜡蚊轮枝抱的产抱量最高，产抱量可达 1．05Xwi‘抱子哈。
将抱子粉进行不同干燥处理，结果表明：低热抽干24h和 30hffi子萌发率显著高
于烘干粉，萌发率分别为 88刀0％和 86．75％；含水率分别为 13．38％和 12刀0％，含抱量
分别达9．53X10‘’和 l．09X10”抱子伯。烘干粉虽然含水率可降到最低8．36O，含抱量
可提高到 1．15 X 101‘抱子哈，但其活抱率仅有的．33％。因此可以首选操作方便，成本
较低的低热抽干的干燥工艺。
在VL07菌株抱子粉贮存过程中发现：抱子粉的活抱率与其内源蛋白和内源总糖
之间均呈极显著的正相关性。高温下贮存的抱子粉较低温下贮存的内源物质代谢快，
冻干16h和24h及抽干30h的抱子粉在4℃下贮存Zld的活抱率分别降到63．5O、72．50
和61．5％。25C下贮存各抱子粉活抱率均降到50％以下。
蜡蚊轮枝抱 VL07菌株对温室桃蚜的防治效果表明：抱悬液门 抱子／ml）防治
区和混合悬液门‘抱子／ml与 25o灭幼腺Ill号 Zxlo‘混合液）防治区的控制效果较好，
喷药后第m天校正虫口下降率分别达乃斗2土1．92％和85．49土3．27％；防治效果分别
为 66。39土4．83％和 80．35土3．98％。
The entomopathogenic fungi, Verticillium lecanii(Zimm.) Viegas, is one of the most promising fungal species which is used to control greenhouse aphids and other sucking insect pests. In recent years, different commercial formulations have been registered in many countries. In order to develop mass production and application of this fungus in China, its biological characteristics, biochemical and physiological factors, diphasic fermentation, conidium drying techniques, nutrient metabolism during storage, and field trial against greenhouse aphids. were studied in this thesis.
Biological characters of 11 strains of V. lecanii isolated from different regions were investigated. At the same time, Bioassay was carried out against the 2~3th instar Myzus persicae larvae with the 11 strains. The most virulent strain was screened out after evaluated by the indices of mortality, infection rate and the median lethal time (LT50). The result showed that VL07 was the most virulent strain which caused mortality soon after 3 days, followed by VL18. The mortality reached 100% and 93.83%, with infection rate at 93.0% and 84.44%, LT50 at 3.30d and 4.02d, respectively. The optimum temperature for V. lecanii infection on M. persicae was 25℃. LD50 of VL07 reached 533 conidia per aphid.
The mycelial biomass and the production of conidia of V. lecanii were studied at different temperatures. The result showed that all strains grew at 10℃ to 30℃, but 25℃ was the optimum temperature for mycelium growth and conidium production. The mycelia biomass of VL07, VL09 and VL18 was compazatively high at 25℃ after 14d. These diameters of colonies reached 4.69cm, 4.72cm and 4.98cm, respectively, obviously higher than other strains. The differentce of conidia production was significant between different strains at different temperature. For VL07, VL18, conidia production reached 5.83 X 109/cm2 and 4.64 X 109/cm2. The result of germination test of conidia showed that the conidia germinated rapidly and uniformly. The germination rates all reached 100% at RH 100%. The germination rates of all strains except for VL07, VL18 declined to 20% as RH
decreased down to below 75%, and grew very slowly.
Some non-hazard insecticides were tested for their effection on conidial germina -tion of V. lecanii. The result showed that they made some effect at high concentration, but the effect was hight at low concentration. 25% flowable colloid of diflubenzuron induced little effect on the germination of the virulent strain VL07. The inhibition rate reached only 1.66% at a concentration of 2 ×104 times.
Some physiological and biochemical factors of V. lecanii were tested. The result showed that the total amount of esterases and the frontier band esterase number were significantly correlated to the virulence, with r=0.9198 and r=0.8820, respectively, and the virulence was poorly correlated to extracellular protease production level and oxalic acid production level.
A series experiments were conducted including screening of carbon and nitrogen resources, inorganic salt, initial value of pH, and initial inoculum by an orthogonal test. The optimum liquid medium for industrial fermentation production of the strain VL07 was determined by comparison of biomass of dry mycelia. Its main components were as follows: soybean powder 4%, white sugar 4%, KH2PO4 0.02%, MgSO4 . 7H2O 0.01%, FeSO4 . 7H2O 0.05%, Vb2 0.05%, Vc 0.05%, with pH value of 6.89. The initial inoculum ranged from 10% to 15%. 21.25mg/ml dry mycelial biomass was acquired after 72h cultivation with inoculum at 3.69-5.50mg/ml. The solid fermentation medium was screened for industrial production of strain VL07 by a L9 (34) orthogonal experiment. The optimum medium determined included bran 42%, rice shell 28%, maize powder 20% and soybean powder 10%. As the initial ratio of water content in the medium was 50%, conidiation reached the highest at 1.05 × 1010 conidia per gram.
In an attempt to develop a drying method of the conidia of V. lecanii produced in diphasic fermentation, several drying methods
用11株蜡蚧轮枝孢对桃蚜(Myzus persicae Sulzer)2～3龄若虫进行生物测定。以桃蚜的校正死亡率、感染率及致死中时(LT_(50))为指标综合评价不同菌株毒力。结果表明，11株蜡蚧轮枝孢菌株中，VL07菌株毒力最强，校正死亡率达100％，感染率达93％，致死中时为3.30d。其次是VL18菌株，致死中时为4.02d，校正死亡率和感染率分别为93.83％和84.44％。在所测试温度中，25℃是蜡蚧轮枝孢侵染桃蚜的最适温度，致死中剂量为533孢子／蚜。
温、湿度对蜡蚧轮枝孢生长和发育的影响为：蜡蚧轮枝孢在10～30℃的温度下均可生长，25℃为菌丝生长最适温度，其中VL07、VL18和VL09菌株菌丝生长量较高，14d累计生长量分别为4.69cm、4.98cm和4.72cm；不同菌株在不同温度下产孢量差异极显著，25℃为蜡蚧轮枝孢最适产孢温度，其中VL07和VL18菌株产孢量最高，14d分别达5.83×10~9和4.64×10~9孢子／cm~2；相对湿度对分生孢子萌发试验表明，在RH为100％时，VL07、VL10和VL18菌株分生孢子萌发快而整齐，24h萌发率均达100％，当相对湿度在75％以下时，除VL07和VL18外，其余各菌株的萌发率均降低到20％以下，而且生长缓慢。
测定4种生物源农药与蜡蚧轮枝孢的相容性结果表明：4种农药高浓度下对蜡蚧轮枝孢分生孢子的萌发均有一定的影响，而在低浓度下影响较小，如在2×10~4倍浓度下25％灭幼脲Ⅲ号悬浮剂对高毒力菌株VL07分生孢子的校正抑制率仅为1.66％。因此，可考虑亚致死剂量的灭幼脲Ⅲ号与蜡蚧轮枝孢混合使用以提高防治效果。
对蜡蚧轮枝孢生理生化因子的测定结果表明：蜡蚧轮枝孢的毒力与菌株酯酶总量及前沿带的酯酶量之间呈显著线性相关，相关系数分别为r=0.9198和r=0.8820；蜡蚧轮枝孢毒力与胞外蛋白酶产生水平以及草酸产生水平之间均无明显相关性。
通过对不同的碳、氮源、无机盐、起始pH值、接种量以及正交试验，以菌丝生物量为指标筛选出了适合于VL07菌株工业生产最佳发酵工艺配方，液相发酵培养基
配方为：黄豆粉 4％，白砂糖一％，KH。PO’ 0刀2％，MgSO厂 7HZO 0刀1％，FeSO厂 7H。O
0＊5％，VB。0＊5％，VC 0＊5％，pH值 6．89，初始接种量为 10％－15％；固相发酵培养
基配方为：鼓皮 42％，稻壳 28％，玉米粉 20％，黄豆粉 10％。当培养基初始含水率为
50％时，蜡蚊轮枝抱的产抱量最高，产抱量可达 1．05Xwi‘抱子哈。
将抱子粉进行不同干燥处理，结果表明：低热抽干24h和 30hffi子萌发率显著高
于烘干粉，萌发率分别为 88刀0％和 86．75％；含水率分别为 13．38％和 12刀0％，含抱量
分别达9．53X10‘’和 l．09X10”抱子伯。烘干粉虽然含水率可降到最低8．36O，含抱量
可提高到 1．15 X 101‘抱子哈，但其活抱率仅有的．33％。因此可以首选操作方便，成本
较低的低热抽干的干燥工艺。
在VL07菌株抱子粉贮存过程中发现：抱子粉的活抱率与其内源蛋白和内源总糖
之间均呈极显著的正相关性。高温下贮存的抱子粉较低温下贮存的内源物质代谢快，
冻干16h和24h及抽干30h的抱子粉在4℃下贮存Zld的活抱率分别降到63．5O、72．50
和61．5％。25C下贮存各抱子粉活抱率均降到50％以下。
蜡蚊轮枝抱 VL07菌株对温室桃蚜的防治效果表明：抱悬液门 抱子／ml）防治
区和混合悬液门‘抱子／ml与 25o灭幼腺Ill号 Zxlo‘混合液）防治区的控制效果较好，
喷药后第m天校正虫口下降率分别达乃斗2土1．92％和85．49土3．27％；防治效果分别
为 66。39土4．83％和 80．35土3．98％。
The entomopathogenic fungi, Verticillium lecanii(Zimm.) Viegas, is one of the most promising fungal species which is used to control greenhouse aphids and other sucking insect pests. In recent years, different commercial formulations have been registered in many countries. In order to develop mass production and application of this fungus in China, its biological characteristics, biochemical and physiological factors, diphasic fermentation, conidium drying techniques, nutrient metabolism during storage, and field trial against greenhouse aphids. were studied in this thesis.
Biological characters of 11 strains of V. lecanii isolated from different regions were investigated. At the same time, Bioassay was carried out against the 2~3th instar Myzus persicae larvae with the 11 strains. The most virulent strain was screened out after evaluated by the indices of mortality, infection rate and the median lethal time (LT50). The result showed that VL07 was the most virulent strain which caused mortality soon after 3 days, followed by VL18. The mortality reached 100% and 93.83%, with infection rate at 93.0% and 84.44%, LT50 at 3.30d and 4.02d, respectively. The optimum temperature for V. lecanii infection on M. persicae was 25℃. LD50 of VL07 reached 533 conidia per aphid.
The mycelial biomass and the production of conidia of V. lecanii were studied at different temperatures. The result showed that all strains grew at 10℃ to 30℃, but 25℃ was the optimum temperature for mycelium growth and conidium production. The mycelia biomass of VL07, VL09 and VL18 was compazatively high at 25℃ after 14d. These diameters of colonies reached 4.69cm, 4.72cm and 4.98cm, respectively, obviously higher than other strains. The differentce of conidia production was significant between different strains at different temperature. For VL07, VL18, conidia production reached 5.83 X 109/cm2 and 4.64 X 109/cm2. The result of germination test of conidia showed that the conidia germinated rapidly and uniformly. The germination rates all reached 100% at RH 100%. The germination rates of all strains except for VL07, VL18 declined to 20% as RH
decreased down to below 75%, and grew very slowly.
Some non-hazard insecticides were tested for their effection on conidial germina -tion of V. lecanii. The result showed that they made some effect at high concentration, but the effect was hight at low concentration. 25% flowable colloid of diflubenzuron induced little effect on the germination of the virulent strain VL07. The inhibition rate reached only 1.66% at a concentration of 2 ×104 times.
Some physiological and biochemical factors of V. lecanii were tested. The result showed that the total amount of esterases and the frontier band esterase number were significantly correlated to the virulence, with r=0.9198 and r=0.8820, respectively, and the virulence was poorly correlated to extracellular protease production level and oxalic acid production level.
A series experiments were conducted including screening of carbon and nitrogen resources, inorganic salt, initial value of pH, and initial inoculum by an orthogonal test. The optimum liquid medium for industrial fermentation production of the strain VL07 was determined by comparison of biomass of dry mycelia. Its main components were as follows: soybean powder 4%, white sugar 4%, KH2PO4 0.02%, MgSO4 . 7H2O 0.01%, FeSO4 . 7H2O 0.05%, Vb2 0.05%, Vc 0.05%, with pH value of 6.89. The initial inoculum ranged from 10% to 15%. 21.25mg/ml dry mycelial biomass was acquired after 72h cultivation with inoculum at 3.69-5.50mg/ml. The solid fermentation medium was screened for industrial production of strain VL07 by a L9 (34) orthogonal experiment. The optimum medium determined included bran 42%, rice shell 28%, maize powder 20% and soybean powder 10%. As the initial ratio of water content in the medium was 50%, conidiation reached the highest at 1.05 × 1010 conidia per gram.
In an attempt to develop a drying method of the conidia of V. lecanii produced in diphasic fermentation, several drying methods
引文
[1] Rombach MC, Gillespie AT. Entomogenous Hyphomycetes for insect and mite control on greenhouse crops. Biocontrol News and Information. 1988, 9: 7～18
[2] 李国霞，郭友中，茅洪新等．应用蜡蚧轮枝菌对大棚蔬菜蚜虫防治效果的研究．中国病毒学，2000，15：262
[3] 陈吉棣．蜡蚧轮枝孢及其在生防中的应用．生物防治通报，1985，1(4)：32-37
[4] Gams W. Cephalosporiurn artige Schimmelipilze (Hyphomycetes) Gustsv Fischer Verlag, Stuttgart, 1971, (5): 518-521
[5] Balazy S. Bull. Soc. Amis. Sic. Lett., poznan, D. 1973, 14: 101-137
[6] Hall R A. The fungus Verticillium lecanii as a microbial insecticide against aphids and scales. In Microbial Control of Pests and Plant Diseases. In:Burges H D,ed. London: Academic Press, 1981, 483～498.
[7] 李国霞，严毓骅，王丽英．温度和营养对北京地区蜡蚧轮枝孢生长发育的影响．生物防治通报，1991，7(3)：115～119
[8] 王丽英，严毓骅．北京温室白粉虱上蜡蚧轮枝孢的自然流行．植物保护学报，1988，15(4)：283
[9] Meyer SLF, Wergin WP. Colonization of soybean cyst nematode females cysts and gelatinous matrices by the fungus Verticillium lecanii. Journal of Nematology. 1998, 30 (4): 436～450.
[10] Meyer SLF, Roberts DP, Wergin WP. Association of the plant-beneficial fungus Verticillium lecanii with soybean roots and rhizosphere. Journal of Nematology, 1998, 30 (4): 451～460
[11] 阎峻．生理生化方法在蜡蚧轮枝菌及一些相关菌聚类分析上的应用．中国虫生真菌研究与应用，北京：中国农业科技出版社，1993，Vol(3)：179～185
[12] 蒲蛰龙(主编)．昆虫病理学．广东：科学技术出版社，1994，69～81
[13] Jackson CW, Heale JB, Hall R A. Traits associated with virulence to the aphid macro-siphoniella sanborni in eighteen isolates of Verticillium lecanii. Annals of Applied Bioliogy, 1985, 106:39-48
[14] Schserffenberg B. Biological and environmental conditions for the development of mycoses caused by Beauveria and Metarhiziurn. J Insect Pathol, 1964, 6:8～12
[15] Fargues J. Etude des condition d'infection des larves de Doryphora, Leptinotarsa decemlieata Say, par Beauvaria bassiana (Bals.) Vuill. Entompophaga, 1972, 17: 319～337
[16] 唐晓庆，樊美珍，李增智．球孢白僵菌继代培养中菌落局变现象及环境影响因素的研究．真菌学报，1996，15(3)：45～53
[17] 唐晓庆，唐燕平，李增智．球孢白僵菌菌种退化及其对马尾松毛虫防治的影响．安徽农业大学学报，1996，23(3)：246～253
[18] 蒲蛰龙，李增智．虫生真菌在有害虫物综合治理中的应用策略．蒲蛰龙，李增智(主编)，昆虫真菌学，合肥：安徽科学技术出版社，1996，369
[19] Gindin G, Geschtovt NU, Raccah B, Barash I. Pathogenicity of Verticillium lecanii to different developmental stagesofthe silver leaf whitefly, Bemisia argentifolii. Phyto-parasitica, 2000, 28 (3): 229～239
[20] 李国霞，严毓骅，王丽英．蜡蚧轮枝菌11个单孢菌株的生物学及其对温室白粉虱致病性的比较和筛选．中国农业大学学报，1996，1(1)：83～88
[21] 洪华珠，武觐文．致病病理学．昆虫真菌学(蒲蛰龙，李增智主编)，合肥：安徽科学技术出版社，1996，93～110
[22] Khachatourians GG. Biochemistry and molecular biology of entomopathoogenic fungi. In The Mycota VI. (Ed. Miler, J. D.). Springer-Verlag: Berlin Heidelberg, 1996, 331～363
[23] 冯明光，李增智．轮枝孢及其应用．有害生物的微生物防治原理与技术(陈涛主编)，武汉：湖北科学技术出版社，1995，256～263
[24] Helyer N et al. Control of chrysanthe mum pests with verticillium lecanii. Phytoparasitica, 1992, 20:5～9
[25] 张仙红，贺运春，王建明等．蜡蚧轮枝菌对害虫致病性的初步研究．植物保护，2001，27(1)：44-46
[26] Williams MED, Edmondson RN, Gill G. The potential of some adjuvants in promoting infection with Verticillium lecanii: laboratory bioassays with Myzus persicae. Annals of applied biology, 2000, 137 (3): 337～345
[27] 罗小刚．一种重要的昆虫病原真菌——蜡蚧轮枝孢V. lecanii. 贵州农业科学，1988，(1)：57～64
[28] Bidochka MJ, Khachatourians GG. Protein hydrolysis in grasshoppers by entomopathogenic fungal proteases. J. Invertbr. Pathol., 1994, 63:7～13
[29] Bidochka MJ, Khachatourians GG. Basic protease of entomopathogenic fungi differ in their adsorption properties to insect cuticle. J. Invertebr. Pathol., 1994, 64:26～32
[30] Askary H, Benhamou N, Brodeur J. Ultrastructural and cytochemical characterization of aphid invasion by the hyphomycete Verticillium lecanii. Journal of Invertebrate Pathology, 1999, 74 (1) 1～13
[31] 洪华珠，杨红．杀虫微生物学纲要．武汉：华中师范大学出版社，1997，139～193
[32] 王海川，尤民生．绿僵菌分生孢子对寄主附着机制的研究进展．昆虫知识，1999，36(3)：189～192
[33] 冯明光．胞外蛋白酶和脂酶活性作为球孢白僵菌毒力指标的可靠性分析．微生物学报，1998，38(6)：461～467
[34] 徐均焕，冯明光，许谦．昆虫病原真菌胞外蛋白酶的病理作用与测定技术．中国生物
防治，1998，14(4)：123～129
[35] 姚剑，李增智，樊美珍．虫生真菌入侵过程中酶的研究．安徽农业大学学报，1996，23(3)：303～308
[36] Khachatourians GG. Physiology and genetics of entomopathogenic fungi. In handbook of applied mycology, Vol 2:humans, animals, and insects.(Ed. Arora, D.K., Ajello L., and Mukerji K. G.). New York:Marcel Dekker, 1991, 613-661
[37] Charnley AK, St Leger RJ. The role of cuticle degrading enzymes in fungal pathogenesis in insects. In The fungal spore and diseade initiation in plants and animals (Ed. Cole, G. T., and Hoch, H. C.). New York: Plenum Press, 1991, 267～286
[38] Michael J, Bidochka Susan Burke Luna Ng. Extracellular hydrolytic enzymes in the fungal genus Verticillium: adaptations for pathogenesis. Canadian journal of microbiology, 1999, 45 (10): 856～864
[39] Gupta SC, Leathers TD, El-Sayed GN, Ignoffo CM. Insect degrading enzymes from entomogenous fungus Beauvaria bassiana Exp. Mycol., 1992, 16:132～137
[40] Bidochka MJ, Khachatourians GG. The implication of metabolic acids produced by Beauveria bassiana in pathogenicity toward the migratory grasshopper, Melanoplus sanguinipes. J. insect. Pathol., 1991, 58:362～370
[41] Havukkala I, Mitamuta C, Hara S, Hirayae K, Nishizawa Y, Hibi T. Induction and purification of Beauvaria bassiana chitinolytic enzymes. J. Invertebr. Pathol., 1993, 61:97～102
[42] Gupta SC, Leathers TD, El-Sayed GN, Ignoffo CM. Production of degradative enaymes by Metarhiziurn anisopliae during growth on defined media and insect cutile. Exp. Mycol., 1991, 15: 310～315
[43] El-Sayed GN, Ignoffo CM, Leathers TD, Gupta SC. Insects cuticle and yeast extract effects on germination growth and production of hydrolytic enzymes of Nomurea rileyi. Mycopathologia, 1993, 122:143～147
[44] El-Sayed GN, Ignoffo CM, Leathers TD, Gupta SC. Cuticular and non-cuticular substrate influence on expressesion of cuticle degrading enzymes from conidia of an entomopathogenic fungus Nornurea rileyi. Mycopathologia, 1993, 122: 79～87
[45] Hsiao WF, Bidochka MJ, Khachatourians GG. Effect of diphenols on the growth of three entomopathogenic fungi. Can. J. Microbiol., 1992, 38:1000-1003
[46] Gindin G, et al. Effect of endotoxin compounds isolated from V. lecanii on the sweet potato whitefly, Bemidia tabaci. Phytoparasitica, 1994, 22 (3): 189～496
[47] Whipps JM. A review of white rust (Puccinia horiana Henn.) disease on chryanthemum and the potential for its biological control with Verticillium lecanii (Zimm.) Viegas [J]. Ann. Appl. Biol. 1993, 122:173～187
[48] 李伟，许维岸，李照会．蜡蚧轮枝菌(Verticillium lecanii)的致病机理及其生防应用研究新进展．山东农业大学学报(自然科学版)，2001，32(4)：561～565
[49] Askary H, Benhamou N, Brodeur J. Ultrastructural and cytochemical Investigations of the Antagonistic Effect of Verticillium lecanii on cucumber powdery mildew [J]. Phytopathology, 1997, 87 (3): 359～368
[50] Gregor, Langen, et al. A β-1, 3-D-endo-mannanase from culture fitrates of the hyperparasites Verticillium lecanii and Aphanocladium album that specifically lyses the germ pore plug from uredospores of Puccinia graminis f. sp. Tritici. Can. J.Bot, 1992, 70: 853～860
[51] 殷凤鸣，陈权才，叶燕华等．蜡蚧轮枝菌防治湿地松粉蚧的技术研究．林业科技通讯，1996(5)：15～18
[52] 邓春生，吴正铠，叶正楚．蜡蚧轮枝孢原生质体形成和再生的研究．中国虫生真菌研究与应用，北京：中国农业科技出版社，1991，Vol(2)：176～179
[53] 农向群，吴正铠，包建中．金龟子绿僵菌原生质体形成和再生的研究[J]．微生物学通报，1990，17(2)：76～79
[54] 邓春生，张爱文，农向群等．紫外线对蜡蚧轮菌原生质体的诱变效果．生物防治通报，1991，7：120～123
[55] 张道海，王未名，陆文华．生防真菌的分子生物学研究．中国虫生真菌研究与应用，北京：中国农业科技出版社，1997，Vol(4)：240～244
[56] Yeo H Pell JK, Aiderson PG, Clark S J, Pye BJ. Laboratory evaluation of temperature effects on the germination and growth of entomopathogenic fungi and on their pathogenicity to two aphid species. Pest management science, 2003, 59 (2): 156～165
[57] 李国霞，严毓骅，王丽英．温度和营养对北京地区蜡蚧轮枝孢生长发育的影响．生物防治通报，1991，7(3)：115～119
[58] 张仙红，贺运春，于桂风等．昆虫病源真菌蜡蚧轮枝菌生物学特性初步研究．山西农业大学学报，2000，20(3)：221～224
[59] Terry Mabbett. Greehouses kept pest free with hot and cold fogging. International Pest Control, 2002, 44 (6):.328～329
[60] Braga GUL, Rangel DEN, Flint SD, Miller CD, Anderson AJ, Roberts DW. Damage and recovery from UV-B exposure in conidia of the entomopathogens Verticillium lecanii and Aphanocladium album. Mycologia, 2002, 94(6): 912～920
[61] Leger R. Changes in translatable mRNA species associated with nutrient depriration and protease synthesis in Metarhizium anisopliae. Journal General Microbioliogy, 1990, 137: 807～815.
[62] Kouvelis VN, Zare R, Typas MA. Differentiation of mitochondrialsubgroups in the Verticillium lecanii species complex. Letters in Applied Microbiology, 1999, 28(4): 263～268.
[63] Zare R ,Typas MA, Bridge PD, Kouvelis VN. Presence of a 20 bp insertion/deletion in the ITS1 region of Verticillium lecanii. Letters in Applied Micro-biology, 1999 (28) 4:258～262
[64] Henia M, Gindin G, Raccah B. Diversity among isolates of Verticillium lecanii expressed by DNA polymorphism and virulence towards Bemisia tabaci Phytoparasitica, 1996, 24(2): 111～118.
[65] Kanaoka M et al. Bassianolide, a new insecticidal cyclodepsipep-tide from Beauveria bassiana and Verticillium lecanii. Agric Biol Chem, 1978, 42(3): 629～635
[66] Claydon N, Grove JF. Insecticidal secondary metabolic products from the entomoenous fungus Verticillium lecanii. J Invert Pathol, 1982, 40:413～418
[67] Gindin G,et al. Effect of endotoxin compounds isolated from V. lecanii on the sweet potato whitefly, Bemisia tabaci. Phytoparasitica, 1994, 22 (3):189～196.
[68] 李国霞，高希武，刘青春等．蜡蚧轮枝孢的发酵培养及其代谢产物对害虫毒杀作用的初步研究．北京农业大学学报，1995，21(4)：409～415
[69] 李国霞，周夏娣，郭友中．两种虫生真菌的次生代谢物质对害虫的毒力．中国昆虫学会2000年学术年会论文集，北京：中国科学技术出版社，2000，428～432
[70] 王克勤，李新民，刘春来等．蜡蚧轮枝孢毒素防治温室白粉虱初步研究．植物保护，2000，26(4)：44～45
[71] 罗科，张青文，郑应华．一株拟青霉代谢产物的杀虫效果及生理活性研究．生物防治通报，1992，8(1)：22～25
[72] 方祺霞．808制剂在茶叶上应用试验．中国虫生真菌研究与应用，北京：学术期刊出版社，1988，Vol(1)：182
[73] 梁宗琦．昆虫病原真菌的综合开发研究．中国虫生真菌研究与应用，北京：学术期刊出版社，1988，Vol(1)：26～35
[74] 吴正铠，李增智．防治农业害虫．昆虫真菌学(蒲蛰龙，李增智主编)，合肥：安徽科学技术出版社，1996，387～403
[75] Majchrowicz L, Poprawski TJ. Effects in vitro of nine fungicides on growth of entomopathgenic fungi. Biocontrol Sci. Technol., 1993, 3:321～336
[76] Poprawski RJ, Majchrowicz I. Effects of herbicides on in vitro vegetative growth and sporulaion of entomopathogenic fungi. Crop Protection, 1995, 14:81～87
[77] 李振，叶琪明，包环玉等．28种农药对蜡蚧轮枝菌菌丝生长的影响．浙江农业学报，1996，8(5)：319～320
[78] Goettel MS, Poprawski TJ, Vandenberg JD et al. Safety to nontarget inver-tebrates of fungal bioaontrol agents. In Safety of microbial insectidides (M. Laird, L. A. Lacey, and E. W. Davidson, EDS). CRC, Boca Raton., 1990, 209～231
[79] James RR, Lightrart B, Susceptibility of the convergent lady beetle (Coleoptera: Coccinellidae) to four entomogenous fungi. Environ. Entomol. 1994, 23:190～192
[80] Poprawski T J, Legaspi JC, Parker P. Influence of entomopathogenic fungi on Serangium parcesetosurn (Coleoptera: Coccinellidae), an important predator of whiteflies (Homoptera: Aleyrodidae). Environ. Entomol.,1998, 27:785～795
[81] Hajek AE, Butler L, Walsh SRA. et al. Host range of the gypsy moth (Lepidoptera:
Lymantriidae) pathogen Entomophaga maimaiga (Zygomycetes: Entomophthorales) in the field versus laboratory. Environ. Entomol. 1996, 25:709～721
[82] Wilding N. Pest control by entomophthorales. In Microbial Control of Pests and Plant Diseases (D. W. Burges Ed). London and New York: Academic Press Inc., 1981, 539～554
[83] Rana JS, Singh D. Entomopathogenic fungi, Verticillium lecanii (Zimm.) as a potential bio-control agent against mustard aphid, Lipaphis Erysimi (Kait.) on Rapeseed-mustard. Cruciferae Newsletter, 2002, 24:97～98
[84] 李国霞，严毓骅，王丽英．蜡蚧轮枝菌11个单孢菌株的生物学及其对温室白粉虱致病性的比较和筛选．中国农业大学学报，1996，1(1)：83～88
[85] Puterka GL, Humber RA, Poprawsk TJ. Virulence of fungal parhogens (imperfect fungi Hyphomycetes) to pear psylla (Homoptera: Psyllidae) [J]. J. Invertbr. P atoll. 1994.23(2): 514～520
[86] 肖燕，任顺祥，邱宝利．应用虫生真菌烟粉虱的现状．昆虫天敌，2001，23(1)：30～34
[87] Meyer LF. Effects of a mutant strain and a wild type strain of Verticillium lecanii on Heterodera glycines populations in the greenhouse [J]. J. Nematol. 1995, 27(3): 409～417
[88] Grabski GC, Mindgem K. The use of Verticillium lecanii as a biological agent against the bean rust fungus Uromyces appendiculatus var. appendiculatus in field and the glasshouse. Phytopath. J, 1985, 113: 243～251
[89] Yohalem DS. Research in antagonistic microorganisms with potential for control of pathogenic foliar fungi. 17th Danish Plant Protection Conference, Horticulture. DJF. Raport Havebrug, 2000, 12: 31～35.
[90] Verhaar MA, Hijwegen A, Zadoks JC. Improvement of the efficacy of Verticillium lecanii use in biocontrol of Sphaerotheca fuliginea by addition of oil formulations. Biocontrol, 1999, 44(1): 73～87
[91] Verhaar MA, Hijwegen A, Zadoks JC. Selection of Verticillium lecanii isolates with high potential for biocontrol powdery mildew by means of components analysis at different humidity regimes [J]. Biocontrol. Sci. Technol, 1998, 8(4): 465～477
[92] Shskira Q, Naheed A, Maryam M, et al. Effects of emtomopathogens on albino mice, Pakistan Journal of Scientific and Industuial Research, 2000, 43 (1): 42～45
[93] 朱国仁，李宝聚．设施蔬菜产业可持续发展的病虫防治对策．中国蔬菜，(增刊)：2000，20～25
[94] Eastop VF. In: Harris K F and M arm orosch K. Aphids as Virus Vectors, 1977, 4～62
[95] 姚建仁，郑永权，董丰收．浅谈农药残留污染、中毒与控制策略．植物保护，2001，27(3)：31～35
[96] 冯明光，黄世文，胡国成．昆虫病原真菌与害虫生物防治：机会与挑战．中国微生
物学会和中国昆虫学会第六届全国杀虫微生物学术讨论会论文摘要集，1995，6
[97] 李增智，黄耀坚．接合菌亚门．昆虫真菌学(蒲蛰龙，李增智主编)，合肥：安徽科学技术出版社，1996，189～223
[98] Humber RA. USDA-ARS Collection of Entomopathogenic Fungal Cultures: Arsef Catalog of strains. Ithaca, New Your, 1992, 504
[99] Milner RJ. Prospects for biopesticides for aphid control. Entomophaga, 1997, 42 (1-2): 227～239
[100] 王未名，赵瑾，陆文华．蚜虫寄生真菌—诺氏虫疫霉．中国虫生真菌研究—应用(第一卷)，学术期刊出版社，1988，153～155
[101] 李增智．虫霉在有害生物综合治理中的应用策略．杀虫微生物(第二卷)，北京：中国农业科技出版社，1989，36～42
[102] 李增智．菌物在害虫、植病和杂草治理中的现状和未来．中国生物防治，1999，15(1)：35～40
[103] Rombach MC, Gillespie AT. Entomogenous Hyphomycetes for insect and mite control on greenhouse crops. Biocontrol News and Information 1988, Vol.(9): 7～18
[104] Hincaie VU et al. Evahuation of the entomopathogen Verticillium lecanii in the control of the aphid Myzus persicae on chrysanthemum. Revista Colombiana de Entomologia. 1990, 16(2): 21～27
[105] Pinna M. Use of Verticillium lecanii (zimm.) for the biological control of Aphis gossypii (Glover) on cucumber in protectedcultivation Informatore Fitopatologico. 1992, 42(10): 56～58
[106] Helyer N. Verticillium lecanii for control of aphids and thrips on cucumber. Bulletin OILB SROP. 1993, 16(2): 63～66
[107] 刘银泉，冯明光，刘树生等．不同温度下球孢白僵菌对桃蚜的毒力．中国生物防治，2000，16(2)：56～60
[108] Adams JB, Emden HF. The biological properties of aphids and teir host plant relationships. In Aphid Technology (H. F. van Emden, Ed).London; academic Press, pp. 1972, 47～104
[109] 方仲达．植病研究方法．第3版，北京：中国农业出版社，1998，343～344
[110] 深见顺一，上杉康彦等编(李树正，王笃祜等译)．农药实验法—杀菌剂篇．第1版，北京：中国农业出版社，1991，32～33
[111] Bidochka MJ, Khachatourians GG. Purification and properties of an extracellular protease produced by Beauveria bassiana. Appl Environ Microbiol, 1987, 53:1679～1684
[112] 唐晓庆，李增智．球孢白僵菌胞外蛋白酶产生水平及其与毒力关系的研究．安徽农业大学学报，1996，23(3)：329～335
[113] 丁德贵．硕士论文，安徽农业大学，1999，19～21
[114] 李建武，萧能庚，余瑞元等．生物化学实验原理和方法(第二版)，北京：北京大
学出版社，1994，189～193
[115] 汪玉树，杜鸣．分光光度法测定菠菜中草酸盐总量．营养学报，1989，11(7)：71～75
[116] Jackson M A. Optimizing nutritional conditions for the liquid cultures production of effective fungal biological control agents. Journal of Industrial Microbiology and Biotechnology, 1997, 19: 180～187
[117] 李连德．硕士论文，安徽农业大学，1998，31
[118] Cliquet S, Jackson MA. Influence of culture condition on production and freeze-drying tolerance of Paecilomyces fumosoroseus blastospores. Journal of Industrial Micro-biology and Biotechnology 1999, 23: 97～102.
[119] Aives SB, Pereira RM, Stimac JL, et al. Delayed germination of Beauveria bassiana conidia after prolonged storage at low, above-freezing temperature. Bicontrol Science and Technology 1996, 6: 575～582.
[120] 李建武，萧能庚，余瑞元等．生物化学实验原理和方法(第二版)北京：北京大学出版社，1994，131～132，402
[121] Lowry OH, Rosbrough NJ, Farr AL, Randdall RJ. Protein measurement wigh folin phenol reagent. J. Biol. Chem., 1995, 193: 265.
[122] 陈其瑚，俞水炎．蚜虫及其防治，上海：上海科学技术出版社，1988，352～358
[123] Feng MG, Poprawski TJ, Khachatoourians GG. Production of formulation and application of the entomopathogenic fungus Beauveria bassiana for insect control: Current status. Biocontr Sci Technol, 1994, 4: 3～34
[124] 许寿涛，冯明光，应盛华．球孢白僵菌对桃蚜接种后特定时间内的侵染率．应用生态学报，2002，13(6)：701～704
[125] 樊美珍，胡锦江，吴克刚．球孢白僵菌对马尾松毛虫的毒力与有关系生化指标的关系．中国虫生真菌研究与应用，北京：中国农业科技出版社，1993，Vol(3)：107～113
[126] 樊美珍，胡锦江．白僵菌酯酶同工酶、脂肪酸与其毒力的关系．安徽农业大学学报，1996，23(3)：260～266
[127] 汤坚，丁珊，孙继美等．47珠球孢白僵菌酯酶同工酶和初步研究．中国虫生真菌研究与应用，北京：中国农业科技出版社，1991，Vol(2)：104～107
[128] Bidochka MJ, Khachatourians GG. Identification of Beauveria bassiana, extracellular protease as virulence factor in pathogenicity toward the migratory grasshopper, Melanoplus sunguinipes. J Invert Patyol, 1990, 56: 362～370
[129] Bidochka MJ, Khachatourians GG. The implicationof metabolic acids produced by B. bassiana. In pathogensis of the migratory grasshopper Melanoplus sanguiipes. J. Invertebr. Patho. 1991, 58: 106～117
[130] Bidochka MJ, Khachatourians GG. Oxalic acid hyperproduction in B. bassiana. Mutants is trlated to a utilizable carbon sources but not to virulence. J. Invertebr. Patho. 1993, 62: 53～57
[131] 唐晓庆，陈东来，黄勃，李增智．球孢白僵菌草酸钙结晶体的提取及其与毒力的关系．安徽农业大学学报，1996，23(3)：293～296
[132] 周德庆．微生物遗传学(第1版)．1998，北京：高等教育出版社，99～106
[133] 中国医学科学院卫生研究所编著，食物成分表，人民卫生出版社，1983，19
[134] 叶满红．关于白砂糖灰分问题的探讨．甘蔗糖业，1998，(5)：49～62
[135] 陶玉贵．生物农药固态发酵条件的研究．安徽师范大学学报(自然科学版)，2001，24(1)：31～33
[136] 殷凤鸣，潘务耀，李增智．白僵菌生产企业标准．安徽农业大学学报，1996，23(3)：321～325
[137] 冯明光，应盛华．不同含水量和温度下贮存中球孢白僵菌分生孢子活力与内贮营养的衰变．应用生态学报，2002，13(4)：439～443
[138] 朱国仁，李宝聚．2000．设施蔬菜产业可持续发展的病虫防治对策，中国蔬菜，(增刊)：20～25
[139] 谢明，邱卫亮，何伟志等．常用杀菌剂对蜡蚧轮枝菌孢子萌发的影响．安徽农业大学学报，1999，26(增刊)：187
[2] 李国霞，郭友中，茅洪新等．应用蜡蚧轮枝菌对大棚蔬菜蚜虫防治效果的研究．中国病毒学，2000，15：262
[3] 陈吉棣．蜡蚧轮枝孢及其在生防中的应用．生物防治通报，1985，1(4)：32-37
[4] Gams W. Cephalosporiurn artige Schimmelipilze (Hyphomycetes) Gustsv Fischer Verlag, Stuttgart, 1971, (5): 518-521
[5] Balazy S. Bull. Soc. Amis. Sic. Lett., poznan, D. 1973, 14: 101-137
[6] Hall R A. The fungus Verticillium lecanii as a microbial insecticide against aphids and scales. In Microbial Control of Pests and Plant Diseases. In:Burges H D,ed. London: Academic Press, 1981, 483～498.
[7] 李国霞，严毓骅，王丽英．温度和营养对北京地区蜡蚧轮枝孢生长发育的影响．生物防治通报，1991，7(3)：115～119
[8] 王丽英，严毓骅．北京温室白粉虱上蜡蚧轮枝孢的自然流行．植物保护学报，1988，15(4)：283
[9] Meyer SLF, Wergin WP. Colonization of soybean cyst nematode females cysts and gelatinous matrices by the fungus Verticillium lecanii. Journal of Nematology. 1998, 30 (4): 436～450.
[10] Meyer SLF, Roberts DP, Wergin WP. Association of the plant-beneficial fungus Verticillium lecanii with soybean roots and rhizosphere. Journal of Nematology, 1998, 30 (4): 451～460
[11] 阎峻．生理生化方法在蜡蚧轮枝菌及一些相关菌聚类分析上的应用．中国虫生真菌研究与应用，北京：中国农业科技出版社，1993，Vol(3)：179～185
[12] 蒲蛰龙(主编)．昆虫病理学．广东：科学技术出版社，1994，69～81
[13] Jackson CW, Heale JB, Hall R A. Traits associated with virulence to the aphid macro-siphoniella sanborni in eighteen isolates of Verticillium lecanii. Annals of Applied Bioliogy, 1985, 106:39-48
[14] Schserffenberg B. Biological and environmental conditions for the development of mycoses caused by Beauveria and Metarhiziurn. J Insect Pathol, 1964, 6:8～12
[15] Fargues J. Etude des condition d'infection des larves de Doryphora, Leptinotarsa decemlieata Say, par Beauvaria bassiana (Bals.) Vuill. Entompophaga, 1972, 17: 319～337
[16] 唐晓庆，樊美珍，李增智．球孢白僵菌继代培养中菌落局变现象及环境影响因素的研究．真菌学报，1996，15(3)：45～53
[17] 唐晓庆，唐燕平，李增智．球孢白僵菌菌种退化及其对马尾松毛虫防治的影响．安徽农业大学学报，1996，23(3)：246～253
[18] 蒲蛰龙，李增智．虫生真菌在有害虫物综合治理中的应用策略．蒲蛰龙，李增智(主编)，昆虫真菌学，合肥：安徽科学技术出版社，1996，369
[19] Gindin G, Geschtovt NU, Raccah B, Barash I. Pathogenicity of Verticillium lecanii to different developmental stagesofthe silver leaf whitefly, Bemisia argentifolii. Phyto-parasitica, 2000, 28 (3): 229～239
[20] 李国霞，严毓骅，王丽英．蜡蚧轮枝菌11个单孢菌株的生物学及其对温室白粉虱致病性的比较和筛选．中国农业大学学报，1996，1(1)：83～88
[21] 洪华珠，武觐文．致病病理学．昆虫真菌学(蒲蛰龙，李增智主编)，合肥：安徽科学技术出版社，1996，93～110
[22] Khachatourians GG. Biochemistry and molecular biology of entomopathoogenic fungi. In The Mycota VI. (Ed. Miler, J. D.). Springer-Verlag: Berlin Heidelberg, 1996, 331～363
[23] 冯明光，李增智．轮枝孢及其应用．有害生物的微生物防治原理与技术(陈涛主编)，武汉：湖北科学技术出版社，1995，256～263
[24] Helyer N et al. Control of chrysanthe mum pests with verticillium lecanii. Phytoparasitica, 1992, 20:5～9
[25] 张仙红，贺运春，王建明等．蜡蚧轮枝菌对害虫致病性的初步研究．植物保护，2001，27(1)：44-46
[26] Williams MED, Edmondson RN, Gill G. The potential of some adjuvants in promoting infection with Verticillium lecanii: laboratory bioassays with Myzus persicae. Annals of applied biology, 2000, 137 (3): 337～345
[27] 罗小刚．一种重要的昆虫病原真菌——蜡蚧轮枝孢V. lecanii. 贵州农业科学，1988，(1)：57～64
[28] Bidochka MJ, Khachatourians GG. Protein hydrolysis in grasshoppers by entomopathogenic fungal proteases. J. Invertbr. Pathol., 1994, 63:7～13
[29] Bidochka MJ, Khachatourians GG. Basic protease of entomopathogenic fungi differ in their adsorption properties to insect cuticle. J. Invertebr. Pathol., 1994, 64:26～32
[30] Askary H, Benhamou N, Brodeur J. Ultrastructural and cytochemical characterization of aphid invasion by the hyphomycete Verticillium lecanii. Journal of Invertebrate Pathology, 1999, 74 (1) 1～13
[31] 洪华珠，杨红．杀虫微生物学纲要．武汉：华中师范大学出版社，1997，139～193
[32] 王海川，尤民生．绿僵菌分生孢子对寄主附着机制的研究进展．昆虫知识，1999，36(3)：189～192
[33] 冯明光．胞外蛋白酶和脂酶活性作为球孢白僵菌毒力指标的可靠性分析．微生物学报，1998，38(6)：461～467
[34] 徐均焕，冯明光，许谦．昆虫病原真菌胞外蛋白酶的病理作用与测定技术．中国生物
防治，1998，14(4)：123～129
[35] 姚剑，李增智，樊美珍．虫生真菌入侵过程中酶的研究．安徽农业大学学报，1996，23(3)：303～308
[36] Khachatourians GG. Physiology and genetics of entomopathogenic fungi. In handbook of applied mycology, Vol 2:humans, animals, and insects.(Ed. Arora, D.K., Ajello L., and Mukerji K. G.). New York:Marcel Dekker, 1991, 613-661
[37] Charnley AK, St Leger RJ. The role of cuticle degrading enzymes in fungal pathogenesis in insects. In The fungal spore and diseade initiation in plants and animals (Ed. Cole, G. T., and Hoch, H. C.). New York: Plenum Press, 1991, 267～286
[38] Michael J, Bidochka Susan Burke Luna Ng. Extracellular hydrolytic enzymes in the fungal genus Verticillium: adaptations for pathogenesis. Canadian journal of microbiology, 1999, 45 (10): 856～864
[39] Gupta SC, Leathers TD, El-Sayed GN, Ignoffo CM. Insect degrading enzymes from entomogenous fungus Beauvaria bassiana Exp. Mycol., 1992, 16:132～137
[40] Bidochka MJ, Khachatourians GG. The implication of metabolic acids produced by Beauveria bassiana in pathogenicity toward the migratory grasshopper, Melanoplus sanguinipes. J. insect. Pathol., 1991, 58:362～370
[41] Havukkala I, Mitamuta C, Hara S, Hirayae K, Nishizawa Y, Hibi T. Induction and purification of Beauvaria bassiana chitinolytic enzymes. J. Invertebr. Pathol., 1993, 61:97～102
[42] Gupta SC, Leathers TD, El-Sayed GN, Ignoffo CM. Production of degradative enaymes by Metarhiziurn anisopliae during growth on defined media and insect cutile. Exp. Mycol., 1991, 15: 310～315
[43] El-Sayed GN, Ignoffo CM, Leathers TD, Gupta SC. Insects cuticle and yeast extract effects on germination growth and production of hydrolytic enzymes of Nomurea rileyi. Mycopathologia, 1993, 122:143～147
[44] El-Sayed GN, Ignoffo CM, Leathers TD, Gupta SC. Cuticular and non-cuticular substrate influence on expressesion of cuticle degrading enzymes from conidia of an entomopathogenic fungus Nornurea rileyi. Mycopathologia, 1993, 122: 79～87
[45] Hsiao WF, Bidochka MJ, Khachatourians GG. Effect of diphenols on the growth of three entomopathogenic fungi. Can. J. Microbiol., 1992, 38:1000-1003
[46] Gindin G, et al. Effect of endotoxin compounds isolated from V. lecanii on the sweet potato whitefly, Bemidia tabaci. Phytoparasitica, 1994, 22 (3): 189～496
[47] Whipps JM. A review of white rust (Puccinia horiana Henn.) disease on chryanthemum and the potential for its biological control with Verticillium lecanii (Zimm.) Viegas [J]. Ann. Appl. Biol. 1993, 122:173～187
[48] 李伟，许维岸，李照会．蜡蚧轮枝菌(Verticillium lecanii)的致病机理及其生防应用研究新进展．山东农业大学学报(自然科学版)，2001，32(4)：561～565
[49] Askary H, Benhamou N, Brodeur J. Ultrastructural and cytochemical Investigations of the Antagonistic Effect of Verticillium lecanii on cucumber powdery mildew [J]. Phytopathology, 1997, 87 (3): 359～368
[50] Gregor, Langen, et al. A β-1, 3-D-endo-mannanase from culture fitrates of the hyperparasites Verticillium lecanii and Aphanocladium album that specifically lyses the germ pore plug from uredospores of Puccinia graminis f. sp. Tritici. Can. J.Bot, 1992, 70: 853～860
[51] 殷凤鸣，陈权才，叶燕华等．蜡蚧轮枝菌防治湿地松粉蚧的技术研究．林业科技通讯，1996(5)：15～18
[52] 邓春生，吴正铠，叶正楚．蜡蚧轮枝孢原生质体形成和再生的研究．中国虫生真菌研究与应用，北京：中国农业科技出版社，1991，Vol(2)：176～179
[53] 农向群，吴正铠，包建中．金龟子绿僵菌原生质体形成和再生的研究[J]．微生物学通报，1990，17(2)：76～79
[54] 邓春生，张爱文，农向群等．紫外线对蜡蚧轮菌原生质体的诱变效果．生物防治通报，1991，7：120～123
[55] 张道海，王未名，陆文华．生防真菌的分子生物学研究．中国虫生真菌研究与应用，北京：中国农业科技出版社，1997，Vol(4)：240～244
[56] Yeo H Pell JK, Aiderson PG, Clark S J, Pye BJ. Laboratory evaluation of temperature effects on the germination and growth of entomopathogenic fungi and on their pathogenicity to two aphid species. Pest management science, 2003, 59 (2): 156～165
[57] 李国霞，严毓骅，王丽英．温度和营养对北京地区蜡蚧轮枝孢生长发育的影响．生物防治通报，1991，7(3)：115～119
[58] 张仙红，贺运春，于桂风等．昆虫病源真菌蜡蚧轮枝菌生物学特性初步研究．山西农业大学学报，2000，20(3)：221～224
[59] Terry Mabbett. Greehouses kept pest free with hot and cold fogging. International Pest Control, 2002, 44 (6):.328～329
[60] Braga GUL, Rangel DEN, Flint SD, Miller CD, Anderson AJ, Roberts DW. Damage and recovery from UV-B exposure in conidia of the entomopathogens Verticillium lecanii and Aphanocladium album. Mycologia, 2002, 94(6): 912～920
[61] Leger R. Changes in translatable mRNA species associated with nutrient depriration and protease synthesis in Metarhizium anisopliae. Journal General Microbioliogy, 1990, 137: 807～815.
[62] Kouvelis VN, Zare R, Typas MA. Differentiation of mitochondrialsubgroups in the Verticillium lecanii species complex. Letters in Applied Microbiology, 1999, 28(4): 263～268.
[63] Zare R ,Typas MA, Bridge PD, Kouvelis VN. Presence of a 20 bp insertion/deletion in the ITS1 region of Verticillium lecanii. Letters in Applied Micro-biology, 1999 (28) 4:258～262
[64] Henia M, Gindin G, Raccah B. Diversity among isolates of Verticillium lecanii expressed by DNA polymorphism and virulence towards Bemisia tabaci Phytoparasitica, 1996, 24(2): 111～118.
[65] Kanaoka M et al. Bassianolide, a new insecticidal cyclodepsipep-tide from Beauveria bassiana and Verticillium lecanii. Agric Biol Chem, 1978, 42(3): 629～635
[66] Claydon N, Grove JF. Insecticidal secondary metabolic products from the entomoenous fungus Verticillium lecanii. J Invert Pathol, 1982, 40:413～418
[67] Gindin G,et al. Effect of endotoxin compounds isolated from V. lecanii on the sweet potato whitefly, Bemisia tabaci. Phytoparasitica, 1994, 22 (3):189～196.
[68] 李国霞，高希武，刘青春等．蜡蚧轮枝孢的发酵培养及其代谢产物对害虫毒杀作用的初步研究．北京农业大学学报，1995，21(4)：409～415
[69] 李国霞，周夏娣，郭友中．两种虫生真菌的次生代谢物质对害虫的毒力．中国昆虫学会2000年学术年会论文集，北京：中国科学技术出版社，2000，428～432
[70] 王克勤，李新民，刘春来等．蜡蚧轮枝孢毒素防治温室白粉虱初步研究．植物保护，2000，26(4)：44～45
[71] 罗科，张青文，郑应华．一株拟青霉代谢产物的杀虫效果及生理活性研究．生物防治通报，1992，8(1)：22～25
[72] 方祺霞．808制剂在茶叶上应用试验．中国虫生真菌研究与应用，北京：学术期刊出版社，1988，Vol(1)：182
[73] 梁宗琦．昆虫病原真菌的综合开发研究．中国虫生真菌研究与应用，北京：学术期刊出版社，1988，Vol(1)：26～35
[74] 吴正铠，李增智．防治农业害虫．昆虫真菌学(蒲蛰龙，李增智主编)，合肥：安徽科学技术出版社，1996，387～403
[75] Majchrowicz L, Poprawski TJ. Effects in vitro of nine fungicides on growth of entomopathgenic fungi. Biocontrol Sci. Technol., 1993, 3:321～336
[76] Poprawski RJ, Majchrowicz I. Effects of herbicides on in vitro vegetative growth and sporulaion of entomopathogenic fungi. Crop Protection, 1995, 14:81～87
[77] 李振，叶琪明，包环玉等．28种农药对蜡蚧轮枝菌菌丝生长的影响．浙江农业学报，1996，8(5)：319～320
[78] Goettel MS, Poprawski TJ, Vandenberg JD et al. Safety to nontarget inver-tebrates of fungal bioaontrol agents. In Safety of microbial insectidides (M. Laird, L. A. Lacey, and E. W. Davidson, EDS). CRC, Boca Raton., 1990, 209～231
[79] James RR, Lightrart B, Susceptibility of the convergent lady beetle (Coleoptera: Coccinellidae) to four entomogenous fungi. Environ. Entomol. 1994, 23:190～192
[80] Poprawski T J, Legaspi JC, Parker P. Influence of entomopathogenic fungi on Serangium parcesetosurn (Coleoptera: Coccinellidae), an important predator of whiteflies (Homoptera: Aleyrodidae). Environ. Entomol.,1998, 27:785～795
[81] Hajek AE, Butler L, Walsh SRA. et al. Host range of the gypsy moth (Lepidoptera:
Lymantriidae) pathogen Entomophaga maimaiga (Zygomycetes: Entomophthorales) in the field versus laboratory. Environ. Entomol. 1996, 25:709～721
[82] Wilding N. Pest control by entomophthorales. In Microbial Control of Pests and Plant Diseases (D. W. Burges Ed). London and New York: Academic Press Inc., 1981, 539～554
[83] Rana JS, Singh D. Entomopathogenic fungi, Verticillium lecanii (Zimm.) as a potential bio-control agent against mustard aphid, Lipaphis Erysimi (Kait.) on Rapeseed-mustard. Cruciferae Newsletter, 2002, 24:97～98
[84] 李国霞，严毓骅，王丽英．蜡蚧轮枝菌11个单孢菌株的生物学及其对温室白粉虱致病性的比较和筛选．中国农业大学学报，1996，1(1)：83～88
[85] Puterka GL, Humber RA, Poprawsk TJ. Virulence of fungal parhogens (imperfect fungi Hyphomycetes) to pear psylla (Homoptera: Psyllidae) [J]. J. Invertbr. P atoll. 1994.23(2): 514～520
[86] 肖燕，任顺祥，邱宝利．应用虫生真菌烟粉虱的现状．昆虫天敌，2001，23(1)：30～34
[87] Meyer LF. Effects of a mutant strain and a wild type strain of Verticillium lecanii on Heterodera glycines populations in the greenhouse [J]. J. Nematol. 1995, 27(3): 409～417
[88] Grabski GC, Mindgem K. The use of Verticillium lecanii as a biological agent against the bean rust fungus Uromyces appendiculatus var. appendiculatus in field and the glasshouse. Phytopath. J, 1985, 113: 243～251
[89] Yohalem DS. Research in antagonistic microorganisms with potential for control of pathogenic foliar fungi. 17th Danish Plant Protection Conference, Horticulture. DJF. Raport Havebrug, 2000, 12: 31～35.
[90] Verhaar MA, Hijwegen A, Zadoks JC. Improvement of the efficacy of Verticillium lecanii use in biocontrol of Sphaerotheca fuliginea by addition of oil formulations. Biocontrol, 1999, 44(1): 73～87
[91] Verhaar MA, Hijwegen A, Zadoks JC. Selection of Verticillium lecanii isolates with high potential for biocontrol powdery mildew by means of components analysis at different humidity regimes [J]. Biocontrol. Sci. Technol, 1998, 8(4): 465～477
[92] Shskira Q, Naheed A, Maryam M, et al. Effects of emtomopathogens on albino mice, Pakistan Journal of Scientific and Industuial Research, 2000, 43 (1): 42～45
[93] 朱国仁，李宝聚．设施蔬菜产业可持续发展的病虫防治对策．中国蔬菜，(增刊)：2000，20～25
[94] Eastop VF. In: Harris K F and M arm orosch K. Aphids as Virus Vectors, 1977, 4～62
[95] 姚建仁，郑永权，董丰收．浅谈农药残留污染、中毒与控制策略．植物保护，2001，27(3)：31～35
[96] 冯明光，黄世文，胡国成．昆虫病原真菌与害虫生物防治：机会与挑战．中国微生
物学会和中国昆虫学会第六届全国杀虫微生物学术讨论会论文摘要集，1995，6
[97] 李增智，黄耀坚．接合菌亚门．昆虫真菌学(蒲蛰龙，李增智主编)，合肥：安徽科学技术出版社，1996，189～223
[98] Humber RA. USDA-ARS Collection of Entomopathogenic Fungal Cultures: Arsef Catalog of strains. Ithaca, New Your, 1992, 504
[99] Milner RJ. Prospects for biopesticides for aphid control. Entomophaga, 1997, 42 (1-2): 227～239
[100] 王未名，赵瑾，陆文华．蚜虫寄生真菌—诺氏虫疫霉．中国虫生真菌研究—应用(第一卷)，学术期刊出版社，1988，153～155
[101] 李增智．虫霉在有害生物综合治理中的应用策略．杀虫微生物(第二卷)，北京：中国农业科技出版社，1989，36～42
[102] 李增智．菌物在害虫、植病和杂草治理中的现状和未来．中国生物防治，1999，15(1)：35～40
[103] Rombach MC, Gillespie AT. Entomogenous Hyphomycetes for insect and mite control on greenhouse crops. Biocontrol News and Information 1988, Vol.(9): 7～18
[104] Hincaie VU et al. Evahuation of the entomopathogen Verticillium lecanii in the control of the aphid Myzus persicae on chrysanthemum. Revista Colombiana de Entomologia. 1990, 16(2): 21～27
[105] Pinna M. Use of Verticillium lecanii (zimm.) for the biological control of Aphis gossypii (Glover) on cucumber in protectedcultivation Informatore Fitopatologico. 1992, 42(10): 56～58
[106] Helyer N. Verticillium lecanii for control of aphids and thrips on cucumber. Bulletin OILB SROP. 1993, 16(2): 63～66
[107] 刘银泉，冯明光，刘树生等．不同温度下球孢白僵菌对桃蚜的毒力．中国生物防治，2000，16(2)：56～60
[108] Adams JB, Emden HF. The biological properties of aphids and teir host plant relationships. In Aphid Technology (H. F. van Emden, Ed).London; academic Press, pp. 1972, 47～104
[109] 方仲达．植病研究方法．第3版，北京：中国农业出版社，1998，343～344
[110] 深见顺一，上杉康彦等编(李树正，王笃祜等译)．农药实验法—杀菌剂篇．第1版，北京：中国农业出版社，1991，32～33
[111] Bidochka MJ, Khachatourians GG. Purification and properties of an extracellular protease produced by Beauveria bassiana. Appl Environ Microbiol, 1987, 53:1679～1684
[112] 唐晓庆，李增智．球孢白僵菌胞外蛋白酶产生水平及其与毒力关系的研究．安徽农业大学学报，1996，23(3)：329～335
[113] 丁德贵．硕士论文，安徽农业大学，1999，19～21
[114] 李建武，萧能庚，余瑞元等．生物化学实验原理和方法(第二版)，北京：北京大
学出版社，1994，189～193
[115] 汪玉树，杜鸣．分光光度法测定菠菜中草酸盐总量．营养学报，1989，11(7)：71～75
[116] Jackson M A. Optimizing nutritional conditions for the liquid cultures production of effective fungal biological control agents. Journal of Industrial Microbiology and Biotechnology, 1997, 19: 180～187
[117] 李连德．硕士论文，安徽农业大学，1998，31
[118] Cliquet S, Jackson MA. Influence of culture condition on production and freeze-drying tolerance of Paecilomyces fumosoroseus blastospores. Journal of Industrial Micro-biology and Biotechnology 1999, 23: 97～102.
[119] Aives SB, Pereira RM, Stimac JL, et al. Delayed germination of Beauveria bassiana conidia after prolonged storage at low, above-freezing temperature. Bicontrol Science and Technology 1996, 6: 575～582.
[120] 李建武，萧能庚，余瑞元等．生物化学实验原理和方法(第二版)北京：北京大学出版社，1994，131～132，402
[121] Lowry OH, Rosbrough NJ, Farr AL, Randdall RJ. Protein measurement wigh folin phenol reagent. J. Biol. Chem., 1995, 193: 265.
[122] 陈其瑚，俞水炎．蚜虫及其防治，上海：上海科学技术出版社，1988，352～358
[123] Feng MG, Poprawski TJ, Khachatoourians GG. Production of formulation and application of the entomopathogenic fungus Beauveria bassiana for insect control: Current status. Biocontr Sci Technol, 1994, 4: 3～34
[124] 许寿涛，冯明光，应盛华．球孢白僵菌对桃蚜接种后特定时间内的侵染率．应用生态学报，2002，13(6)：701～704
[125] 樊美珍，胡锦江，吴克刚．球孢白僵菌对马尾松毛虫的毒力与有关系生化指标的关系．中国虫生真菌研究与应用，北京：中国农业科技出版社，1993，Vol(3)：107～113
[126] 樊美珍，胡锦江．白僵菌酯酶同工酶、脂肪酸与其毒力的关系．安徽农业大学学报，1996，23(3)：260～266
[127] 汤坚，丁珊，孙继美等．47珠球孢白僵菌酯酶同工酶和初步研究．中国虫生真菌研究与应用，北京：中国农业科技出版社，1991，Vol(2)：104～107
[128] Bidochka MJ, Khachatourians GG. Identification of Beauveria bassiana, extracellular protease as virulence factor in pathogenicity toward the migratory grasshopper, Melanoplus sunguinipes. J Invert Patyol, 1990, 56: 362～370
[129] Bidochka MJ, Khachatourians GG. The implicationof metabolic acids produced by B. bassiana. In pathogensis of the migratory grasshopper Melanoplus sanguiipes. J. Invertebr. Patho. 1991, 58: 106～117
[130] Bidochka MJ, Khachatourians GG. Oxalic acid hyperproduction in B. bassiana. Mutants is trlated to a utilizable carbon sources but not to virulence. J. Invertebr. Patho. 1993, 62: 53～57
[131] 唐晓庆，陈东来，黄勃，李增智．球孢白僵菌草酸钙结晶体的提取及其与毒力的关系．安徽农业大学学报，1996，23(3)：293～296
[132] 周德庆．微生物遗传学(第1版)．1998，北京：高等教育出版社，99～106
[133] 中国医学科学院卫生研究所编著，食物成分表，人民卫生出版社，1983，19
[134] 叶满红．关于白砂糖灰分问题的探讨．甘蔗糖业，1998，(5)：49～62
[135] 陶玉贵．生物农药固态发酵条件的研究．安徽师范大学学报(自然科学版)，2001，24(1)：31～33
[136] 殷凤鸣，潘务耀，李增智．白僵菌生产企业标准．安徽农业大学学报，1996，23(3)：321～325
[137] 冯明光，应盛华．不同含水量和温度下贮存中球孢白僵菌分生孢子活力与内贮营养的衰变．应用生态学报，2002，13(4)：439～443
[138] 朱国仁，李宝聚．2000．设施蔬菜产业可持续发展的病虫防治对策，中国蔬菜，(增刊)：20～25
[139] 谢明，邱卫亮，何伟志等．常用杀菌剂对蜡蚧轮枝菌孢子萌发的影响．安徽农业大学学报，1999，26(增刊)：187