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Hervibors Hosts Influence on Insecticide Resistance: a Review

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Hervibors Hosts Influence on Insecticide Resistance: a Review Rev. Fac. Agron. (LUZ).1999,16: 127-140 Docurnent: Hervibors hosts influence on insecticide resistance: a review Documento: La influencia del hospedero en la resistencia de hervíboros a insecticidas: una revisión Abstract Resistance is a worldwide problem, which if ignored or improperly niana- ged, will significantly reduce worldwide agricultura1 production and public health. Resistance is influenced by genetic factors but also there is an environmental effect, which in the case of phytofagas diseases is partially represented by the chemicals found in the host plants. Species with an evolutionary history of feeding on heavily chemically defended plant structures shoiild have elwated levels of enzyrnes that detoxify defensive chemicals, and therefore an enhrinced ability to evolve resistance to synthetic toxins. The role of host plant cheniistry on the expression and evolution of pesticide resistance is reviewed from the per- spective of understanding the non-genetic factors influencing pesticide resist arice. This perspective is important since environmental factors may have re1ai;ively important effects influencing the activity of detoxification enzymes in anirnals, and hence, susceptibility to xenobiotics. Research on non-genetic factors influ- encing pesticide resistance must be undertaken if we are to iiicrease our confi- dence in proposed management strategies. Key words :detoxification enzymes, pesticide resistance, non-genetic factors, susceptibility to allelochemicals. Resumen La resistencia de hervi%oros a insecticidas es un problema a nivel mundial, que si es ignorado o manejado inadecuadamente, reduciría significativame-ntela producción agrícola mundial y la salud pública. La resistencia está influenciada por factores genéticos pero también existe un efecto del medio ambiente, que en el caso de las plagas fitófagas está en parte representado por sustancias químicas ~écibidoel 19-12-1996 Aceptado el 24-09-1997 1. Cument address: Departamento Fitosanitario, Facultad de Agronomía, 1,UZ.Apartado Postal 15378.Maracaibo Edo. Zulia Vcnczuela Fax N (061) 596183 E-mail :[email protected]. (To whom correspondcnce should be addressed) 2. Department of Entomology, 501ASI, The Pennsylvania State University, Universitj Park, PA 16802, USA. Fax N(814) 865-3048 E-mail : BAMQ PSUVM.PSU.EDU. Domínguez y McPheron presentes en las plantas hospederas. Las especies que se alimentan de estructv.ras de plantas muy bien defendidas químicamente deberían tener elevados nivele:; de enzimas que detoxifiquon las sustancias químicas usadas por las plantas para defenderse, y por lo tanto muestran una habilidad mejorada para desarrollar resistencia a las toxinas sintéticas. Se revisa el rol de la química de la planta hospedera, desde el punto de vista de entender el efecto de los factores no-genéticos que influyen en la resistencia a plaguicidas de los insectos herbívoros. Elsta perspectiva es importante ya que los factores del medio ambiente pueden llegar a tener un importante efecto en la actividad enzimática de detoxificación de los animales, y por lo tanto, la respectiva susceptibilidad a los xenobióticos. Investigación de los factores no-genéticos que influyen en la resistencia a plaguicidas debe ser llevada a cabo si queremos incrementar nuestra confianza en las estrategias de manejo propuestas. Palabras clave: enzimas, detoxificación, resistencia, plaguiciclas, factores no- genéticos, susceptibilidad, nleloquímicos. Introduction The evolution of resistance to pes- pest populations, the phenotype upon ticides is an example of the evolution- which selection operates is a func-;ion ary process. The pesticicle is the selec- of both the genotype and the envilson- tion pressure, which creates a very ment. Relatively little research has strong fitness differential between sus- focused on the influence of envil-on- ceptible and resistant genotypes. The mental factors on the evolution of ?es- survival and subsequent reproduction ticide resistance. In t.he case of p!ant of resistant individuals leads to a pests, the chemical constituents of change in the frequency over time of plants are a significant part of the en- alleles conferring resistance. Wide- vironment, a part that has been spread application of pesticides has led shown to affect the action of many re- to a global resistance problem (12,13). sistance mechanisms. We review tlie Resistance compromise crops, animal role of host plant chemistry on the ex- production and human health pression and evolution of pesticid~ire- (through pesticide resistance in vectors sistance and show that this interac- of animal and human diseases and, tion must be considered if we are to drug resistance in the pathogens and develop rational pest managerrient parasites). While selection pressure strategies for safe and efficient i:rop acts to change allele Frequencies within production. Relationship between detoxification (Enzymatic Mechanisms) and host plants (Allelochemicals) It became obvious relatively early els of insecticide resist.ance rather rap- in the history of insecticide use that idly. Gordon (15) suggested that the polyphagous species develop high lev- natural exposure of polyphagous spe- Rev. Fac. Agron. (LUZ). 1999,16: 127-140 cies to a wide variety of plant in the esterases are acetylcholinest- allelochemicals had resulted in high erase, important in the proper t rans- capacities for their detoxification, mission of nerve siglials, and juveriile which would now enable the insects to horrnone esterase, wluch helps to regu- develop resistance to synthetic insec- late the process of metamorpkiosis. ticides. This idea directly implicated These enzymes work, in general, by plant allelochemicals as the natural breaking carboxylester and substrates for insecticide detoxifying phosphorotriester bonds. The:~are enzymes for the first time. active against many types of in:;ecti- Plant allelochemicals modify lev- cides, especially organophosphates and els of detoxifjmg enzymes in herbivores pyrethroids. Much of the evidence for and, therefore, their susceptibility to a role of esterases in insecticide iaesis- insecticides (4, 6, 26, 29, 34, 41, 45). tance comes from nssays of general Insects have detoxification mecha- esterase activity using model sub- nisms to deal with plant chemicals and strates. However, this is only an indi- also often the same mechanisms are rect measure of the role of esterases. involved in pesticide resistance. It is Some studies with synergists havc?also important to understand the interac- implicated esterases. Esterase genes tion of plant allelochemicals with the associated with insecticide resistance detoxification system. have been identified in mosquitoes 'and Three systems OS detoxification aphids. enzymes e., polysubstrate Mullin and Croft (33) fo-r ex- monooxygenases (PSMO),general es- ample, found large differences in gen- terases (GE),and glutathione S-trans- eral esterase activity relative to ferases (GST) are commonly regarded snapbean (ranging from 0.4-fold on a as the most important biochemical mint to 2.4-fold on umbellifers) for mechanism for the metabolism of Tetranychus urticae fed 13 difforent xenobiotics (7, 45) including host-adapted strains. allelochemicais (53)and pesticides (17). Lindroth (25) studied the ef'fects Xenobiotics may act as inducers by of food plant on larval performance and stimulating enzyme synthesis (53). midgut detoxification enzymes iii lar- Insects induced by dietary vae of the luna moth, Actias luna. He allelochemicals or host plants appar- found that larva1 food plants (1)lack ently increase metabolism of severa1 cherry, cottonwood, quaking aspen, synthetic pesticides, as demonstrated white willow, red oak, white oal.;, tu- by their increased tolerance to these lip tree, paper birch, black walnut, compounds (17). Insecticide resistant butternut, shagbark hickory) affected strains of insects often have greater the activities of solul>leesterases and detoxikng enzyme activities (43),and were 1.8-fold higher in larvae fed wal- in at least one example, enzyme induc- nut, than in larvae fed birch. Microso- ibility was greater than in a suscep- mal esterases exhibited an opposite tible strain (37). trend in activity, with lowest v:ilues Esterases. This is a very large in larvae fed walnuf;, and highest in family of related enzymes. Included those fed birch. Domínguez y McPheron Activities of microsomal cis- and actions and many substrates. Each trans-epoxide hydrolase in northern particular enzyme hris a broad, but corn rootworm, Diabrotica barberi unique, pattern of substrate spekfic- Smith& Lawrence, were significantly ity. Most of our knowledge of the increased by diet shifts from corn ear monooxygenase system comes fisom to squash blossom and sunflower in- studies on mammalian liver. How- florescence, while levels of these en- ever, some recent gerietic studie:; in zymes in the western corn rootworm, insects are beginning to add to our D. virgifera uirgifera LeConte \vere understanding. It is now clear tha3ta unaffected (42). specific cytochrome P,,,-dependent Susceptible larvae from artificial monooxygenase is responsible for the diet had significantly higher nonspe- ability of black swallowtail butterfly cific esterase activity than susceptible caterpillars to deal with certain ch~mi- larvae fed apple, gorse, broom, and cals in their diet. In the case of m 3ny blackberry. Furthermore, activity of organophosphate insecticides, cerí ain nonspecific esterases of resistant lar- monooxygenase enzymes actu:tlly vae fed blackberry
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