Insect Control: Biological and Synthetic Agents - Index of

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protein; Bole et al., 1986) play key roles in protein folding, transport, and quality control in a wide range of eukaryotic cells including insect cells (Ruddon and Bedows, 1997; Ailor and Betenbaugh, 1999; Trombetta and Parodi, 2003). The coexpression of BiP by a recombinant baculovirus has been shown to reduce aggregation and increase secretion of baculovirus expressed murine IgG in T. ni derived cells (Hsu et al., 1994). Taniai et al. (2002) have investigated whether coexpression of BiP can improve the secretion of AcAaIT expressed AaIT (rAaIT) in Sf-21 cells. They found that coexpression of BiP increased the amount of soluble rAaIT, however, the amount of active rAaIT that was secreted into the culture medium was not improved. Protein disulfide isomerase (PDI) is another factor within the ER that is known to help protein folding and secretion in insect cells by catalyzing the oxidization, reduction, and isomerization of disulfide bonds in vitro (Hsu et al., 1996; Ruddon and Bedows, 1997). Coexpression of a PDI expressing baculovirus with an AaITexpressing baculovirus was also ineffective in increasing the amount of active AaIT that was secreted into the culture medium (Taniai, Inceoglu, and Hammock, unpublished data). 10.3.4.3. Expression of multiple synergistic toxins At least six distinct receptor sites are found on the voltage gated sodium channels of mammals and insects that are the molecular targets of a broad range of neurotoxins. Two additional receptor sites are found on the insect sodium channel that are the molecular targets of insect selective excitatory and depressant scorpion toxins (Cestele and Catterall, 2000). On the basis of binding experiments using radiolabeled toxins, the depressant toxins bind to two noninteracting binding sites (one showing high affinity and the other low affinity) on the insect sodium channel (Gordon et al., 1992; Zlotkin et al., 1995; Cestele and Catterall, 2000). The excitatory toxins bind only to the high-affinity receptor site (Gordon et al., 1992). Herrmann et al. (1995) have shown that when excitatory and depressant toxins are simultaneously coinjected into larvae of the blowfly S. falculata or H. virescens, the amount of toxin required to give the same paralytic response is reduced five- to tenfold in comparison to the amount required when only one of the toxins is injected. On the basis of this synergism, they suggested that speed of kill of recombinant baculovirus(es) could be further reduced by: (1) the coinfection of two or more recombinant baculoviruses that each express toxin genes with synergistic properties; or (2) genetic modification such that two or more synergistic toxin genes are simultaneously expressed. 10: Genetically Modified Baculoviruses for Pest Insect Control 353 Regev et al. (2003) have generated a recombinant AcMNPV (vAcLqIT1-IT2) that expresses both the excitatory LqhIT1 and the depressant LqhIT2 toxins under the very late p10 and polh promoters, respectively. Time-response bioassays (at an LC95 dose) using neonate H. virescens showed that the ET 50 of vAcLqIT1-IT2 was reduced by 41% (46.9 versus 79.1 h) in comparison to AcMNPV. In comparison, the ET50s of recombinant AcMNPVs expressing only LqhIT1 (vAcLqIT1) or only LqhIT2 (vAcLqIT2) were 57.3 and 60.0 h, respectively. Thus, expression of both LqhIT1 and LqhIT2 by vAcLqIT1-IT2 resulted in a synergistic effect that reduced the ET 50 by 10.4 h (18%) and 13.1 h (22%), respectively. A synergistic effect (15–19%) was also observed following coinfection of neonate H. virescens with a mixture of both vAcLqIT1 and vAcLqIT2 (ET50 of 48.8 h). Time-response bioassays (at an LC95 dose) using neonate H. armigera showed that the ET50 of vAcLqIT1-IT2 was reduced by 24% (69.1 versus 90.8 h) in comparison to AcMNPV. However, a synergistic effect was not elicited in comparison to neonate H. armigera infected with vAcLqIT1 or vAcLqIT2 alone. Since H. armigera is only a semi-permissive host of AcMNPV, Regev et al. (2003) hypothesized that inefficient oral infection may have diminished any synergistic effect. By intrahemocoelic injection (at an LD95 dose) of third instar larvae of H. armigera, they found that the ET 50 of vAcLqIT1-IT2 (72 h) was reduced by 13 h (15%) and 70 h (49%) in comparison to larvae injected with only vAcLqIT1 or vAcLqIT2, respectively. In similar intrahemocoelic injection assays of third instar larvae of S. littoralis (nonpermissive by oral infection for AcMNPV), they found that the ET50 of vAcLqIT1-IT2 (54 h) was reduced by 43 h (44%) and 20 h (27%) in comparison to control larvae injected with only vAcLqIT1 or vAcLqIT2, respectively. Similar or decreased levels of synergism were found with a recombinant AcMNPV (vAcLqaIT- IT2) expressing both alpha and depressant scorpion toxins in orally and intrahemocoelically infected larvae of H. virescens, H. armigera, and S. littoralis. Using thoracic neurons of the grasshopper Schistocerca americana Drury, Prikhod’ko et al. (1998) have shown that the spider and sea anemone toxins, m-Aga-IV and As II, respectively, act at distinct sites on the insect sodium channel and synergistically promote channel opening. These toxins also showed synergism when injected into third instar blowfly Lucilia sericata and fourth instar fall armyworm S. frugiperda. Prikhod’ko et al. (1998) have generated recombinant AcMNPVs (vMAg4Sat2 and vhsMAg4SAt2) that coexpress both m-Aga-IV and
354 10: Genetically Modified Baculoviruses for Pest Insect Control As II toxins under PSynXIV or hsp70 promoters, respectively. Time-response bioassay (at an LC95 dose) using neonate T. ni or S. frugiperda showed that the ET50 of vMAg4Sat2 was reduced by 39% (60.6 versus 99.7 h) and 45% (56.8 versus 104.0 h), respectively, in comparison to AcMNPV. A synergistic effect, however, was not found in comparison to larvae infected with recombinant AcMNPVs expressing only m-Aga-IV (vMAg4pþ) or As II (vDASAt2). In similar bioassays, the ET50 of vhsMAg4SAt2 was reduced by 47% (52.3 versus 99.2 h) and 57% (48.3 versus 111.1 h), respectively, in comparison to AcMNPV. A synergistic effect, however, was again not found in comparison to larvae infected with vhsMAg4pþ or vDAhsSAt2. They hypothesized that synergism was not detected in the case of coexpression of these toxins under the very late PSynXIV promoter (by vMAg4Sat2) because it takes time to activate this promoter during infection and, once activated, high enough levels of expression of either one of the toxins is sufficient to effectively debilitate the host insect. In the case of coexpression under the hsp70 promoter by vhsMAg4SAt2, they hypothesized that lower ET50s (i.e., a synergistic effect) may not be possible since it takes a minimum period of time (i.e., approximately 50 h) for the virus to initiate both primary and secondary infections within the host. Although synergism was not observed in terms of the ET 50 (mean time at which 50% of the test larvae cease to respond to stimulus), synergism was observed in terms of the FT50 (mean time at which 50% of the test larvae cease feeding) of neonate T. ni infected with vhsMAg4SAt2 (at an LC95 dose by droplet feeding). The FT50 of vhsMAg4SAt2 infected neonates was lower than that of vMAg4pþ, vDASAt2, and AcMNPV by approximately 25% (24.3 versus 32.3 h), 29% (24.3 versus 34.3 h), and 63% (24.3 versus 65.8 h). Prikhod’ko et al. (1998) suggested that 24 h may be the fastest time at which a recombinant baculovirus could stop feeding because of the time (approximately 12 h) that it takes to complete the primary infection cycle within the midgut. Furthermore, they suggested that an FT50 of 24 h is sufficiently rapid to make recombinant baculovirus pesticides competitive with chemical pesticides. 10.3.5. Toxin Interactions with Chemical Pesticides Pyrethroids, carbamates, and organophosphates are common synthetic chemical pesticides that are used for protection against pest insects worldwide. Pyrethroids currently account for about 20% of the total market for insecticides (Khambay, 2002). Pyrethroids are the most commonly used chemical insecticides for the control of the tobacco budworm H. virescens and cotton bollworm H. zea in North America. Pyrethroids are divided into two groups, type I and type II, on the basis of the absence or presence, respectively, of a cyano (-CN) group in the alpha position of the carboxyl moiety. Both type I and type II pyrethroids target receptors in the sodium channel causing them to remain open and resulting in prolonged sodium influx that causes hyper-excitation of the nervous system of both insects and mammals. Insects, however, are more sensitive to pyrethroid action than mammals due to several factors including the relative concentration of pyrethroid metabolizing enzymes, binding kinetics of the pyrethroid to the receptor, and differential sensitivity of the insect and mammalian sodium channels to pyrethroid action (Soderlund et al., 2002). Although the precise location of the pyrethroid receptor within the insect sodium channel is unknown, the binding site is apparently different from that of peptide toxins produced by spiders, sea anemones, and scorpions (Ghiasuddin and Soderlund, 1985; Herrmann et al., 1995). The differential binding of pyrethroid and toxin molecules indicate that when they are both present within the sodium channel they can act independently, synergistically, or antagonistically. McCutchen et al. (1997) have characterized the effect of low concentrations of six chemical pesticides (allethrin, cypermethrin, DDT, endosulfan, methomyl, and profenofos) on the efficacy of wildtype and recombinant baculoviruses. They found synergistic interactions when neonate H. virescens were exposed to AcAaIT and low concentrations (LC10–LC20 at 24 h) of the type II pyrethroid (cypermethrin) and a carbamate (methomyl). The interactions of AcAaIT with allethrin, DDT, endosulfan, or profenofos were not synergistic but additive. Specifically, neonate H. virescens that were inoculated with 2000 polyhedra per larva (a greater than LC99 dose) of AcAaIT and immediately exposed to an LC10–LC20 dose of cypermethrin or methomyl showed an LT50 of 30 h, a reduction of roughly 58% compared to control larvae infected with the virus alone. Neither synergistic nor antagonistic effects were observed in control experiments in which AcJHE.KK (a recombinant AcMNPV expressing a modified juvenile hormone esterase) was used in place of AcAaIT. Since the mode and site of action of JHE is putatively different from that of AaIT, this was not surprising. In contrast, the interactions of wild-type AcMNPV and the chemical pesticides (except for methomyl) were slightly antagonistic. McCutchen et al. (1997) have also examined the efficacy of AcAaIT in combination with a low dose
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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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Page 325 and 326: 314 9: Insecticidal Toxins from Pho
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404 11: Entomopathogenic Fungi and
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Zare, R., Gams, W., Culham, A., 200
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434 A11: Addendum although a number
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436 A11: Addendum Examination of ge
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440 12: Insect Transformation for U
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448 A12: Addendum genetic transform
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452 Subject Index Aphis gossypii (c
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454 Subject Index Bisacylhydrazine
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456 Subject Index Cry toxins (conti
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458 Subject Index Entomopathogenic
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460 Subject Index Homoptera molting
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462 Subject Index Kinoprene (ENSTAR
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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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