Endophytic fungal entomopathogens with activity against plant pathogens: Ecology and evolution

The Ecology of Fungal Entomopathogens

Volumn , Issue , 2010, Pages 113-128

  Ownley, Bonnie H ^a   Gwinn, Kimberly D ^a   Vega, Fernando E ^b  

^a UNIVERSITY OF TENNESSEE   (United States)

^b AGRICULTURAL RESEARCH SERVICE   (United States)

Author keywords

Beauveria bassiana; Fungal endophyte; Hypocreales; Induced systemic resistance; Lecanicillium; Mycoparasite; Trichoderma

Indexed keywords

EID: 84900290129     PISSN: None     EISSN: None     Source Type: Book    
DOI: 10.1007/978-90-481-3966-8_9     Document Type: Chapter

References (115)

1. Ahmed AS, Sanchez CP, Candela ME (2000) Evaluation of induction of systemic resistance in pepper plants (Capsicum annuum) to Phytophthora capsici using Trichoderma harzianum and its relation with capsidiol accumulation. Eur J Plant Pathol 106:817-824
2. Aidemark M, Andersson C-J, Rasmusson AG, Widell S (2009) Regulation of callose synthase activity in situ in alamethicin-permeabilized Arabidopsis and tobacco suspension cells. BMC Plant Biol 9:27 (in press)
3. Allen DJ (1982) Verticillium lecanii on the bean rust fungus, Uromyces appendiculatus. Trans Br Mycol Soc 79: 362-364
4. Arnold AE (2007) Understanding the diversity of foliar endophytic fungi: progress, challenges, and frontiers. Fungal Biol Rev 21:51-66
5. Arnold AE, Maynard Z, Gilbert GS, Coley PD, Kursar TD (2000) Are tropical fungal endophytes hyperdiverse? Ecol Lett 3:267-274
6. Askary H, Yarmand H (2007) Development of the entomopathogenic hyphomycete Lecanicillium muscarium (Hyphomycetes: Moniliales) on various hosts. Eur J Entomol 104:67-72
7. Askary H, Benhamou N, Brodeur J (1997) Ultrastructural and cytochemical investigations of the antagonistic effect of Verticillium lecanii on cucumber powdery mildew. Phytopathology 87:359-368
8. Askary H, Carriere Y, Belanger RR, Brodeur J (1998) Pathogenicity of the fungus Verticillium lecanii to aphids and powdery mildew. Biocontrol Sci Technol 8:23-32
9. Bark YG, Lee DG, Kim YH, Kang SC (1996) Antibiotic properties of an entomopathogenic fungus, Beauveria bassiana, on Fusarium oxysporum and Botrytis cinerea. Korean J Plant Pathol 12:245-250
10. Belanger RR, Labbe C (2002) Control of powdery mildews without chemicals: prophylactic and biological alternatives for horticultural crops. In: Belanger RR, Bushnell WR, Dik AJ, Carver TLW (eds) The powdery mildews: A comprehensive treatise. American Phytopathological Society, St. Paul, pp 256-267
11. Benhamou N (2004) Potential of the mycoparasite, Verticillium lecanii, to protect citrus fruit against Penicillium digitatum, the causal agent of green mold: A comparison with the effect of chitosan. Phytopathology 94:693-705
12. Benhamou N, Brodeur J (2000) Evidence of antibiosis and induced host defense reaction in the interaction between Verticillium lecanii and Penicillium digitatum, the causal agent of green mold. Phytopathology 90:932-943
13. Benhamou N, Brodeur J (2001) Pre-inoculation of Ri T-DNA transformed cucumber roots with the mycoparasite, Verticillium lecanii, induces host defense reactions against Pythium ultimum infection. Physiol Mol Plant Pathol 58:133-146
14. Berg G, Smalla K (2009) Plant species and soil type cooperatively shape the structure and function of microbial communities in the rhizosphere. FEMS Microbiol Ecol 68:1-13
15. Berg G, Zachow C, Lottmann J, Gotz M, Costa R, Smalla K (2005) Impact of plant species and site on rhizosphereassociated fungi antagonistic to Verticillium dahlia Kleb. Appl Environ Microbiol 71:4203-4213
16. Bidochka MJ, Burke S, Ng L (1999) Extracellular hydrolytic enzymes in the fungal genus Verticillium: Adaptations for pathogenesis. Can J Microbiol 45:856-864
17. Bills GF, Polishook JD (1991) Microfungi from Carpinus caroliniana. Can J Bot 69:1477-1482
18. Boucias DG, Pendland JC (1998) Principles of insect pathology. Kluwer Acad Pub, Boston
19. Boucias DG, Mazet I, Pendland J, Hung SY (1995) Comparative analysis of the in vivo and in vitro metabolites produced by the entomopathogen Beauveria bassiana. Can J Bot 73(Suppl 1):S1092-S1099
20. Bruinsma M, Pang B, Mumm R, van Loon JJA, Dicke M (2009) Comparing induction at an early and late step in signal transduction mediating indirect defence in Brassica oleracea. J Exp Bot 60:2589-2599
21. Clark MM, Gwinn KD, Ownley BH (2006) Biological control of Pythium myriotylum. Phytopathology 96:S25
22. Conrath U, Beckers GJM, Flors B, Garc?a-Agust?n P, Jakab G, Mauch F, Newman M-A, Pieterse CMJ, Poinssot B, Pozo MJ, Pugin A, Schaffrath Ton J, Wendehenne D, Zimmerli L, Mauch-Mani B (2006) Priming: getting ready for battle. Mol Plant Microbe Interact 19:1062-1071
23. Copping LG, Menn JJ (2000) Biopesticides: A review of their action, applications and efficacy. Pest Manag Sci 56: 651-676
24. Crespo R, Pedrini N, Juarez MP, Dal Bello GM (2008) Volatile compounds released by the entomopathogenic fungus Beauveria bassiana. Microbiol Res 163:148-151
25. Djonovic S, Pozo MJ, Dangott LJ, Howell CR, Kenerley CM (2006) Sm1, a proteinaceous elicitor secreted by the biocontrol fungus Trichoderma virens induces plant defense responses and systemic resistance. Mol Plant Microbe Interact 19:838-853
26. Djonovic S, Vargas WA, Kolomiets MV, Horndeski M, Wiest A, Kenerley CM (2007) A proteinaceous elicitor Sm1 from the beneficial fungus Trichoderma virens is required for induced systemic resistance in maize. Plant Physiol 145:875-889
27. Dubos B (1987) Fungal antagonism in aerial agrobiocenoses. In: Chet I (ed) Innovative approaches to plant disease control. Wiley-Liss, New York, pp 107-135
28. Elliot SL, Sabelis MW, Janssen A, van der Geest LPS, Beerling EAM, Fransen J (2000) Can plants use entomopathogens as bodyguards? Ecol Lett 3:228-235
29. Ernst M, Medgen KW, Wirsel SGR (2003) Endophytic fungal mutualists: seed borne Stagonospora spp. enhance reed biomass production in axenic microcosms. Mol Plant Microbe Interact 16:580-587
30. Eyal J, Mabud MDA, Fischbein KL, Walter JF, Osborne LS, Landa Z (1994) Assessment of Beauveria bassiana Nov. EO-1 strain, which produces a red pigment for microbial control. Appl Biochem Biotechnol 44:65-80
31. Fenice M, Selbmann L, Zucconi L, Onofri S (1997) Production of extracellular enzymes by Antarctic fungal strains. Polar Biol 17:275-280
32. Flori P, Roberti R (1993) Treatment of onion bulbs with antagonistic fungi for the control of Fusarium oxysporum f. sp. cepae. Difesa delle Piante 16:5-12
33. Fravel DR (1988) Role of antibiosis in the biocontrol of plant diseases. Annu Rev Phytopathology 26:75-91
34. Fry WE (1982) Principles of plant disease management. Academic Press, New York
35. Genthner FJ, Cripe GM, Crosby DJ (1994) Effect of Beauveria bassiana and its toxins on Mysidopsis bahia (Mysidacea). Arch Environ Contam Toxicol 26:90-94
36. Gimenez C, Cabrera R, Reina M, Gonzalez-Coloma M (2007) Fungal endophytes and their role in plant protection. Curr Org Chem 11:707-720
37. Goettel MS, Koike M, Kim JJ, Aiuchi D, Shinya R, Brodeur J (2008) Potential of Lecanicillium spp. for management of insects, nematodes and plant diseases. J Invertebr Pathol 98:256-261
38. Gomez-Vidal S, Lopez-Llorca LV, Jansson H-B, Salinas J (2006) Endophytic colonization of date palm (Phoenix dactylifera L) leaves by entomopathogenic fungi. Micron 37:624-632
39. Gomez-Vidal S, Salinas J, Tena M, Lopez-Llorca LV (2009) Proteomic analysis of date palm (Phoenix dactylifera L) responses to endophytic colonization by entomopathogenic fungi. Electrophoresis 30:2996-3005
40. Griffin MR (2007) Beauveria bassiana, a cotton endophyte with biocontrol activity against seedling disease. Ph.D. Dissertation, The University of Tennessee, Knoxville, TN, USA
41. Grove J, PopleM(1980) The insecticidal activity of beauvericin and the enniatin complex. Mycopathologia 70:103-105
42. Gupta S, Montllor C, Hwang Y-S (1995) Isolation of novel beauvericin analogues from the fungus Beauveria bassiana. J Nat Prod 58:733-738
43. Gutjahr C, Paszkowski U (2009) Weights in the balance: jasmonic acid and salicylic acid signaling in root-biotroph interactions. Mol Plant Microbe Interact 22:763-772
44. Hajek AE (1997) Ecology of terrestrial fungal entomopathogens. Adv Microb Ecol 15:193-249
45. Hall RA (1980) Laboratory infection of insects by Verticillium lecanii strains isolated from phytopathogenic fungi. Trans Br Mycol Soc 74:445-446
46. Hall RA (1981) The fungus Verticillium lecanii as a microbial insecticide against aphids and scales. In: Burges HD (ed) Microbial control of pests and plant diseases 1970-1980. Academic Press, London, pp 483-498
47. Hanson LE, Howell CR (2004) Elicitors of plant defense responses from biocontrol strains of Trichoderma virens. Phytopathology 94:171-176
48. Harmon GE, Howell CR, Viterbo A, Chet I, Lorito M (2004) Trichoderma species-opportunistic, avirulent plant symbionts. Nature Rev Microbiol 2:43-56
49. Hartley SE, Gange AC (2009) Impacts on plant symbiotic fungi on insect herbivores: mutualism in a multitrophic context. Annu Rev Entomol 54:323-342
50. Hirano E, Koike M, Aiuchi D, Tani M (2008) Pre-inoculation of cucumber roots with Verticillium lecanii (Lecanicillium muscarium) induces resistance to powdery mildew. Res Bull Obihiro Univ 29:82-94
51. Holder DJ, Keyhani NO (2005) Adhesion of the entomopathogenic fungus Beauveria (Cordyceps) bassiana to substrata. Appl Environ Microbiol 71:5260-5266
52. Hyde KD, Soytong K (2008) The fungal endophyte dilemma. Fungal Divers 33:163-173
53. Inglis GD, Goettel MS, Butt TM, Strasser H (2001) Use of hyphomycetous fungi for managing insect pests. In: Butt TM, Jackson C, Magan N (eds) Fungi as biocontrol agents. Progress, problems and potential. CABI Publishing, Oxfordshire, pp 23-69
54. Jeger MJ, Jeffries P, Elad Y, Xu X-M (2009) A generic theoretical model for biological control of foliar plant diseases. J Theor Biol 256:201-214
55. Kamp AM (2002) Pleiomorphic deterioration in entomopathogenic fungi. MS thesis, Brock University. St. Catherines, Ontario, Canada
56. Kanaoka M, Isogai A, Murakoshi S, Ichinoe M, Suzuki A, Tamura S (1978) Bassianolide, a new insecticidal cyclodepsipeptide from Beauveria bassiana and Verticillium lecanii. Agr Biol Chem 42:629-635
57. Keller S, Zimmermann G (1989) Mycopathogens of soil insects. In: Wilding N, Collins NM, Hammond PM, Webber JF (eds) Insect-fungus interactions. Acad Press, London, pp 239-270
58. Kim JJ, Goettel MS, Gillespie DR (2007) Potential of Lecanicillium species for dual microbial control of aphids and the cucumber powdery mildew fungus, Sphaerotheca fuliginea. Biol Control 40:327-332
59. Kim JJ, Goettel MS, Gillespie DR (2008) Evaluation of Lecanicillium longisporum, Vertalec for simultaneous suppression of cotton aphid, Sphaerotheca fuliginea, on potted cucumbers. Biol Control 45:404-409
60. Koike M, Higashio T, Komori A, Akiyama K, Kishimoto N, Masuda E, Sasaki M, Yoshida S, Tani M, Kuramoti K, Sugimoto M, Nagao H (2004) Verticillium lecanii (Lecanicillium spp) as epiphyte and its application to biological control of arthropod pests and diseases. IOBC/ WPRS Bull 27:41-44
61. Korolev N, David DR, Elad Y (2008) The role of phytohormones in basal resistance and Trichoderma-induced systemic resistance to Botrytis cinerea in Arabidopsis thaliana. BioControl 53:667-683
62. Lacey LA, Horton DR, Jones DC, Headrick HL, Neven LG (2009) Efficacy of biofumigant fungus Muscodor albus (Ascomycota: Xylariales) for control of codling moth (Lepidoptera: Tortricidae) in stimulated storage conditions. J Econ Entomol 102:43-49
63. Leckie BM, Ownley BH, Pereira RM, Klingeman WE, Jones CJ, Gwinn KD (2008) Mycelia and spent fermentation broth of Beauveria bassiana incorporated into synthetic diets affect mortality, growth and development of larval Helicoverpa zea (Lepidoptera: Noctuidae). Biocontrol Sci Technol 18:697-710
64. Lee S-M, Yeo W-H, Jee H-J, Shin S-C, Moon Y-S (1999) Effect of entomopathogenic fungi on growth of cucumber and Rhizoctonia solani. FRI J For Sci 62:118-125
65. Linder MB (2009) Hydrophobins: proteins that self assemble at interfaces. Curr Opin Colloid Interface Sci 14:356-363
66. Mercier J, Jimenez JI (2004) Control of fungal decay of apples and peaches by the biofumigant fungus Muscodor albus. Postharvest Biol Technol 31:1-8
67. Mercier J, Smilanick JL (2005) Control of green mold and sour rot of stored lemon by biofumigation with Muscodor albus. Biol Control 32:401-407
68. Meyer SLF, Huettel RN, Sayre RM (1990) Isolation of fungi from Heterodera glycines and in vitro bioassays for their antagonism to eggs. J Nematol 22:532-537
69. Meyling N, Eilenberg J (2007) Ecology of the entomopathogenic fungi Beauveria bassiana and Metarhizium anisopliae in temperate agroecosystems: potential for conservation biological control. Biol Control 43:145-155
70. Moran-Diez E, Hermosa R, Ambrosino P, Cardoza RE, Gutie rrez S, Lorito M, Monte E (2009) The ThPG1 endopolygalacturonase is required for the Trichoderma harzianum-plant beneficial interaction. Mol Plant Microbe Interact 22:1021-1031
71. Oller-Lopez JL, Iranzo M, Mormeneo S, Oliver E, Cuerva JM, Oltra JE (2005) Bassianolone: An antimicrobial precursor of cephalosporolides E and F from the entomoparasitic fungus Beauveria bassiana. Org Biomol Chem 3:1172-1173
72. Ownley BH, Windham MT (2007) Biological control of plant pathogens. In: Trigiano RN, Windham MT, Windham AS (eds) Plant pathology concepts and laboratory exercises, 2nd edn. CRC Press, New York, pp 423-436
73. Ownley BH, Bishop DG, Pereira RM (2000) Biocontrol of Rhizoctonia damping-off of tomato with Beauveria bassiana. Phytopathology 90:S58
74. Ownley BH, Pereira RM, Klingeman WE, Quigley NB, Leckie BM (2004) Beauveria bassiana, a dual purpose biocontrol organism, with activity against insect pests and plant pathogens. In: Lartey RT, Caesar A (eds) Emerging concepts in plant health management. Research Signpost, Kerala, pp 256-269
75. Ownley BH, Griffin MR, Klingeman WE, Gwinn KD, Moulton JK, Pereira RM (2008) Beauveria bassiana: endophytic colonization and plant disease control. J Invertebr Pathol 3:267-270
76. Perazzolli M, Dagostin S, Ferrar A, Elad Y, Pertot I (2008) Induction of systemic resistance against Plasmopara viticola in grapevine by Trichoderma harzianum T39 and benzothiadiazole. Biol Control 47:228-234
77. Petrini O (1981) Endophytische pilze in epiphytischen Araceae, Bromeliaceae und Orchidiaceae. Sydowia 34: 135-148
78. Posada F, Vega FE (2005) Establishment of the fungal entomopathogen Beauveria bassiana (Ascomycota: Hypocreales) as an endophyte in cocoa seedlings (Theobroma cacao). Mycologia 97:1195-1200
79. Powell WA, Klingeman WE, Ownley BH, Gwinn KD (2009) Evidence of endophytic Beauveria bassiana in seedtreated tomato plants acting as a systemic entomopathogen to larval Helicoverpa zea (Lepidoptera: Noctuidae). J Entomol Sci 44:391-396
80. Reisenzein H, Tiefenbrunner W (1997) Growth inhibiting effect of different isolates of the entomopathogenic fungus Beauveria bassiana (Bals) Vuill to the plant parasitic fungi of the genera Fusarium, Armillaria and Rosellinia. Pflanzenschutz Berichte 57:15-24
81. Reithner B, Schuhmacher R, Stoppacher N, Pucher M, Brunner K, Zeilinger S (2007) Signaling via the Trichoderma atroviride mitogen-Activated protein kinase Tmk1 differentially affects mycoparasitism and plant protection. Fungal Genet Biol 44:1123-1133
82. Renwick A, Campbell R, Coe S (1991) Assessment of in vivo screening systems for potential biocontrol agents of Gaeumannomyces graminis. Plant Pathol 40:524-532
83. Riga K, Lacey LA, Guerra N (2008) The potential of the endophytic fungus, Muscodor albus, as a biocontrol agent against economically important plant parasitic nematodes of vegetable crops in Washington State. Biol Control 45:380-385
84. Ruocco M, Lanzuise S, Vinale F, Marra R, Turra D, Woo SL, Lorito M (2009) Identification of a new biocontrol gene in Trichoderma atroviride: the role of an ABC transporter membrane pump in the interaction with different plant pathogenic fungi. Mol Plant Microbe Interact 22:291-301
85. Saikkonen K, Faeth SH, Helander M, Sullivan TJ (1998) Fungal endophytes: A continuum of interactions with host plants. Annu Rev Ecol Systemat 29:319-343
86. Saksirirat W, Hoppe H-H (1991) Degradation of uredospores of the soybean rust fungus (Phakopsora pachyrhizi Syd) by cell-free culture filtrates of the mycoparasite Verticillium psalliotae Treschow. J Phytopathology 132:33-45
87. Schulz B, Boyle C (2005) The endophytic continuum. Mycol Res 109:661-686
88. Segarra G, Casanova E, Bellido D, Odena MA, Oliveira E, Trillas I (2007) Proteome, salicylic acid, and jasmonic acid changes in cucumber plants inoculated with Trichoderma asperellum strain T34. Proteomics 7:3943-3952
89. Shakeri J, Foster HA (2007) Proteolytic activity and antibiotic production by Trichoderma harzianum in relation to pathogenicity to insects. Enzym Microb Tech 40:961-968
90. Shinya R, Aiuchi D, Kushida A, Tani M, Kuramochi K, Koike M (2008) Effects of fungal culture filtrates of Verticillium lecanii (Lecanicillium spp) hybrid strains on Heterodera glycines eggs and juveniles. J Invertebr Pathol 97:291-297
91. Shoresh M, Harman GE (2008) The molecular basis of shoot responses of maize seedlings to Trichoderma harzianum T22 inoculation of the root: A proteomic approach. Plant Physiol 147:2147-2163
92. Spencer DM, Atkey PT (1981) Parasitic effects of Verticillium lecanii on two rust fungi. Trans Br Mycol Soc 77:535-542
93. St. Leger RJ, Joshi L, Roberts DW (1997) Adaptation of proteases and carbohydrases of saprophytic, phytopathogenic, and entomopathogenic fungi to the requirements of their ecological niches. Microbiology 143:1983-1992
94. Strobel GA (2006) Muscodor albus and its biological promise. J Ind Microbiol Biotechnol 33:514-522
95. Strobel GA, Dirkse E, Sears J, Markworth C (2001) Volatile antimicrobials from Muscodor albus, a novel endophytic fungus. Microbiology 147:2943-2950
96. Taniguchi M, Kawaguchi T, Tanaka T, Oi S (1984) Antimicrobial and respiration inhibitory activities of oosporein. Agr Biol Chem 48:1065-1067
97. Tefera T, Vidal S (2009) Effect of inoculation method and plant growth medium on endophytic colonization of sorghum by the entomopathogenic fungus Beauveria bassiana. BioControl 54:663-669
98. Termorshuizen AJ, Jeger MJ (2009) Strategies of soilborne plant pathogenic fungi in relation to disease suppression. Fung Ecol 1:108-114
99. Van Loon LC, Bakker PAHM, Pieterse CMJ (1998) Systemic resistance induced by rhizosphere bacteria. Annu Rev Phytopathology 36:453-483
100. Van Wees SCM, Van der Ent S, Pieterse CMJ (2008) Plant immune responses triggered by beneficial microbes. Curr Opin Plant Biol 11:443-448
101. Vandermeer J, Perfecto I, Liere H (2009) Evidence for hyperparasitism of coffee rust (Hemileia vastatrix) by the entomogenous fungus, Lecanicillium lecanii, through a complex ecological web. Plant Pathol 58:636-641
102. Vargas WA, Djonovic S, Sukno SA, Kenerley CM (2008) Dimerization controls the activity of fungal elicitors that trigger systemic resistance in plants. J Biol Chem 283:19804-19815
103. Vega FE (2008) Insect pathology and fungal endophytes. J Invertebr Pathol 98:277-279
104. Vega FE, Posada F, Aime MC, Pava-Ripoll M, Infante F, Rehner SA (2008) Entomopathogenic fungal endophytes. Biol Control 46:72-82
105. Vega FE, Goettel MS, Blackwell M, Chandler D, Jackson MA, Keller S, Koike M, Maniania M, Monzon A, Ownley BH, Pell JK, Rangel DEN, Roy HE (2009a) Fungal entomopathogens: new insights on their ecology. Fungal Ecol 2:149-159
106. Vega FE, Simpkins A, Aime MC, Posada F, Peterson SW, Rehner SA, Infante F, Castillo A, Arnold AE (2009b) Fungal endophyte diversity in coffee plants from Colombia, Hawaii, Mexico, and Puerto Rico. Fungal Ecol (in press)
107. Verhaar MA, Hijwegen T, Zadoks JC (1996) Glasshouse experiments on biocontrol of cucumber powdery mildew (Sphaerotheca fuliginea) by the mycoparasites Verticillium lecanii and Sporothrix rugulosa. Biol Control 6: 353-360
108. Vesely D, Koubova D (1994) In vitro effect of the entomopathogenic fungi Beauveria bassiana (Bals.-Criv) Vuill and B. brongniartii (Sacc) Petch on phytopathogenic fungi. Ochr Rostl 30:113-120
109. Vinale F, Sivasithamparam K, Ghisalberti DL, Marra R, Woo SL, Lorito M (2008) Trichoderma-plant-pathogen interactions. Soil Biol Biochem 40:1-10
110. Viterbo A, Chet I (2006) TasHyd1, a new hydrophobin gene from the biocontrol agent Trichoderma asperellum, is involved in plant root colonization. Mol Plant Pathol 7:249-258
111. Whipps JM (1993) A review of white rust (Puccinia horiana Henn) disease on chrysanthemum and the potential for its biological control with Verticillium lecanii (Zimm) Vie-gas. Ann Appl Biol 122:173-187
112. White JF, Belanger F, Meyer W, Sullivan RF, Bischoff JF, Lewis EA (2002) Clavicipitalean fungal epibionts and endophytes-development of symbiotic interactions with plants. Symbiosis 33:201-213
113. Widler B, Muller E (1984) Untursuchungen uber endophytische Pilze von Arctostaphylos uva-ursi (L) Sprenger (Ericaceae). Bot Helv 94:307-337
114. Ying S-H, Feng M-G (2004) Relationship between thermotolerance and hydrophobin-like proteins in aerial conidia of Beauveria bassiana and Paecilomyces fumosoroseus as fungal biocontrol agents. J Appl Microbiol 97:323-331
115. Zhang Y, Zhao J, Fang W, Zhang J, Luo Z, Zhang M, Fan Y, Pei Y (2009) Mitogen-Activated protein kinase hog1 in the entomopathogenic fungus Beauveria bassiana regulates environmental stress responses and virulence to insects. Appl Environ Microbiol 75:3787-3795