Insect Control: Biological and Synthetic Agents - Index of

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11: Entomopathogenic Fungi and their Role in Regulation of Insect Populations 415 when compared to other groups of organisms, but some interesting research has been done. The most advanced research has been on the hyphomycete fungi M. anisopliae and B. bassiana. Transformation systems have been developed for some entomopathogenic fungi. The first transformations of an entomopathogenic fungus were by Bernier et al. (1989) and Goettel et al. (1990b), who expressed a benomyl resistance gene in M. anisopliae. Electroporation and biolistics have also been used to transform M. anisopliae (St. Leger et al., 1995). Polyethylene glycol (PEG) mediated transformation of protoplasts is another method for transformation of entomopathogenic fungi, as done with P. fumosoroseus and P. lilacinus (Inglis et al., 1999b) using benomyl as the selective agent. Sandhu et al. (2001) developed a heterologous transformation system for B. bassiana and M. anisopliae based on the use of the Aspergillus nidulans nitrate reductase gene (niaD), after EMS treatment of protoplasts and regeneration on chlorate medium. Genetic modification of entomopathogenic fungi by direct manipulation has been led by St. Leger, and the area was recently reviewed by St. Leger and Screen (2001), covering both insect and plant pathogenic fungi. In the numerous publications, these researchers described the identification and cloning of protease genes, particularly the pr1 cuticle degrading protease gene from M. anisopliae (St. Leger et al., 1992a). Several other entomopathogenic Hyphomycetes (A. flavus, B. bassiana, Paecilomyces farinosus, Tolypocladium niveum, and L. lecanii) also have pr1-type enzymes (St. Leger et al., 1991, 1992a). Efforts to produce more effective strains of fungi have included the overexpression of pr1 by insertion of multiple copies, resulting in increased speed of kill of a host insect, although transformants had very poor sporulation ability (St. Leger, 2001). Modification of pr1 gene expression in M. anisopliae resulted in melanization and cessation of feeding 25–30 h earlier than the wildtype disease in caterpillars (St. Leger et al., 1996). The group has gone on to place the pr1 gene under control of a constitutive promoter and included a green fluorescent protein encoding gene for tracking the fungi after field release (Hu and St. Leger, 2002). Genes involved in regulation of PR1 expression have also been identified from M. anisopliae (Screen et al., 1997, 1998). Chitinase, produced by many insect pathogenic fungi and implicated in pathogenesis, has also been the target of molecular studies. Several chitinase gene sequences from Beauveria and Metarhizium have been submitted to GenBank and described in publications (e.g., Bogo et al., 1998). Overexpres- sion of extracellular chitinase has also been demonstrated for M. anisopliae var. acridum, but did not alter virulence to the caterpillar, M. sexta, compared to the wild-type fungus (Screen et al., 2001). 11.6.3.3. Formulation and application strategies There are a number of limitations that still affect the use of fungi. Formulation and application of fungal propagules has the potential to overcome some of these limitations. Jones and Burges (1998) defined four basic functions of formulation: (1) stabilization of the agent during production, distribution, and storage; (2) aiding the handling and application of the product to the target in the most appropriate manner and form; (3) protection of the agent from harmful environmental factors at the target site, thereby increasing persistence; and (4) enhancing efficacy of the agent at the target site by increasing activity, reproduction, contact, and interaction with the target pest. Progress has been made in formulation science, and formulations for entomopathogens have been comprehensively reviewed by Burges (1998). Early formulations of entomopathogenic fungi often used just a weak detergent to get the hydrophobic spores into solution, without any efforts to use the formulations to improve efficacy. More recently, the use of oil as a formulation ingredient has been successfully used to overcome several problems. In the LUBILOSA program developing M. anisopliae var. acridum for control of locusts in Africa, the combination of oil formulations with ultra low volume (ULV) spraying has led to successful development of a biopesticide (Lomer et al., 2001). Although it would seem impossible to use an entomopathogenic fungus requiring high humidity against locusts in the hot, dry conditions prevalent in much of their environment, studies have shown that ambient humidity may not be as critical as once thought (Fargues et al., 1997; see also Section 11.3.1). Formulating Metarhizium conidia in nonevaporative diluents such as oils allowed the conidia to attach, spread (Inglis et al., 1996a), and germinate on susceptible locusts, even at low ambient humidities (Bateman, 1997). Formulation can also assist in extending shelf stability of fungal propagules generally desired by commercial producers. Spore survival after mass production varies greatly, depending on the species, but few entomopathogenic species of fungi can achieve the 1–2 years of shelf stability at formulation. Formulation in clay material is known to improve the shelf life of many conidial preparations (e.g., Shi, 1998). This appears to be due to the ability of clay to reduce the moisture content of the spores,
416 11: Entomopathogenic Fungi and their Role in Regulation of Insect Populations as spores survive longer with less than 5% moisture (Wraight et al., 2001). Moore et al. (1996) have shown that survival of conidia of M. anisopliae was highest at low (
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INSECT CONTROL BIOLOGICAL AND SYNTH
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INSECT CONTROL BIOLOGICAL AND SYNTH
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CONTENTS Preface vii Contributors i
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PREFACE When Elsevier published the
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J T Andaloro E. I. Du Pont de Nemou
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1 Pyrethroids B P S Khambay and P J
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activity. In contrast to most other
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Figure 1 Commercial and novel pyret
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Figure 2 Pyrethroids referred to in
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4-position result in almost complet
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and their primary site of action is
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Figure 4 Folded and extended confor
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aromatic rings or methyl groups by
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pyrethroids) and consequently a sec
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1998), although the precise mechani
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polymorphisms in the protein. Of th
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observed resistance to pyrethroids,
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propoxur against Culex quinquefasci
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esistance in house fly. Insect Bioc
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Shan, G.M., Hammock, B.D., 2001. De
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A1.4. Resistance The increasing num
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B A IIS4-S5 linker, IIS5 helix and
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2 Indoxacarb and the Sodium Channel
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Figure 2 Structures of pyrazoline-l
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observed that both indoxacarb and i
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deflections resulting from stresses
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Figure 8 Dihydropyrazole block appe
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potential conduction in the CNS of
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known to affect blocker affinity, w
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Figure 13 Diagrammatic representati
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that the probable mechanism of ovil
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conventional chemistries and spinos
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Clare, J.J., Tate, S.N., Nobbs, M.,
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Tsurubuchi, Y., Karasawa, A., Nagat
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R1 N N O N H indoxacarb. Thus, whil
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3 Neonicotinoid Insecticides P Jesc
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esidues, like the pyrid-3-ylmethyl
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containing neonicotinoids, and neon
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Studies were also done using compar
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Becke, 1988; Klamt, 1995) level of
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sampling using forcefield methods (
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Figure 9 Stable conformations and p
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Figure 13 Binding to putative catio
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Figure 14 Systemicity of neonicotin
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site (Kayser et al., 2002). Clothia
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Figure 20 Important pest insects ta
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Barley yellow dwarf virus vectors s
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investigate the mode of action of n
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within the desensitized state over
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the accumulation of this acid in th
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3.8.1. Safety Profile The introduct
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Table 7 Environmental profile of co
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substance is bound irreversibly to
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imidacloprid conferring a high leve
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nAChR are comparable to the efficac
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Figure 25 Flea control achieved wit
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and the resolution of FAD was teste
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Brown, J.K., Perring, T.M., Cooper,
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In: Ishaaya, I. (Ed.), Biochemical
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Léna, C., Changeux, J.-P., 1993. A
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patterns in Colorado potato beetle
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nach Saatgutbeizung. Pflanzenschutz
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Today, photoaffinity labeling (e.g.
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for control of stinkbugs in soybean
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Biochem. Physiol., doi: 10.1016/j.p
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4 Insect Growth- and Development-Di
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The molting process is initiated by
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Table 1 Bisacylhydrazine insecticid
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4.2.2. Bisacylhydrazines as Tools o
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to the three EcRs with either muris
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of action of ecdysone agonists was
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thus inducing effects and symptomol
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and microsomes. Subsequently, Willi
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dipteran insects, like the midge, C
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eetle (Exomala orientalis) when app
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metabolic fate studies showed the i
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y specific proteins in signaling pa
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their reduced risk for the environm
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can be reversed by applying JH. JH
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door for a more rational approach t
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apparent that it was a bHLH-PAS and
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Table 9 Continued Bemisia tabaci (s
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Table 11 Insect control with some a
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Table 13 Ecotoxicological profile o
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Flucycloxuron Nonsystemic acaricide
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The discovery of diflubenzuron spaw
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Table 15 Environmental effects of s
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ecessive (R male S female) manner,
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esistance management programs due t
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IPS Paraconfusus lanier (Coleoptera
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larvae parasitized by Microplitis r
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Kostyukovsky, M., Chen, B., Atsmi,
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Three-dimensional quantitative stru
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Riddiford, L.M., 1994. Cellular and
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characterization of resistant clone
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Biological Approaches to Pest Contr
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M389 L308 M272 T304 T393 V404 L420
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5 Azadirachtin, a Natural Product i
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and Hemiptera and was related to ef
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eported, but this is rather nonspec
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important source of pest control at
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effects with physiological effects
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eing associated with an accumulatio
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et al., 1992). Salehzadeh et al. (2
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ehavior of Spodoptera littoralis (p
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indica A. Juss. IBH Publishing Comp
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Smith, S.L., Mitchell, M.J., 1988.
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which interact directly with azadir
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6 The Spinosyns: Chemistry, Biochem
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Table 1 Structures of the spinosyns
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Table 2 Biological activity of spin
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A large synthetic effort has gone i
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216 6: The Spinosyns: Chemistry, Bi
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Figure 4 Spinosad metabolism in avi
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A6 Addendum: The Spinosyns T C Spar
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246 A6: Addendum Scott, 2008) and i
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248 7: Bacillus thuringiensis: Mech
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A7 Addendum: Bacillus thuringiensis
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280 A7: Addendum toxins is magnitud
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Table 1 Comparative properties of s
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286 8: Mosquitocidal B. sphaericus:
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Table 4 Characteristics of various
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A8 Addendum: Bacillus sphaericus Ta
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310 A8: Addendum essential to devel
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314 9: Insecticidal Toxins from Pho
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A9 Addendum: Recent Advances in Pho
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330 A9: Addendum Lee, S.C., Stoilov
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332 10: Genetically Modified Baculo
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Page 375 and 376: 364 10: Genetically Modified Baculo
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Page 463 and 464: 452 Subject Index Aphis gossypii (c
Page 465 and 466: 454 Subject Index Bisacylhydrazine
Page 467 and 468: 456 Subject Index Cry toxins (conti
Page 469 and 470: 458 Subject Index Entomopathogenic
Page 471 and 472: 460 Subject Index Homoptera molting
Page 473 and 474: 462 Subject Index Kinoprene (ENSTAR
Page 475 and 476: 464 Subject Index Muscle(s) sodium
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466 Subject Index Photorhabdus (con
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468 Subject Index Sodium channel bl
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470 Subject Index Toxocara cati, im
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