Field cage performance of two tachinid parasitoids of the tomato fruitworm on insect resistant and susceptible tomato lines

Rates of parasitism ofHelicoverpa (=Heliothis) zea (Boddie) (Lepidoptera: Noctuidae) by two tachinid parasitoid species differing in larviposition habits were measured in field cages on tomato plant lines with or without methyl-ketone (2-tridecanone and 2-undecanone)-mediated insect resistance. Hosts were placed on resistant or susceptible plants, exposed to parasitoids for 24–48 h, then held on artificial diet for parasitoid emergence. Rates of parasitism byArchytas marmoratus (Townsend) (Diptera: Tachinidae), which larviposits on its host's food plant, were significantly reduced on the resistant plants, relative to those on the susceptible plants. Parastism by another Tachinid,Eucelatoria bryani Sabrosky, which larviposits directly into its host and does not directly contact the foliage of its hosts' food plant as a small larva, was not affected by the methyl-ketone mediated resistance.     

Indirect plant-mediated effects on insect immunity and disease resistance in a tritrophic system

Host plant quality can significantly influence the growth and condition of phytophagous insects, and consequently their susceptibility to pathogens. This study examined the relationship between host plant quality, insect condition, immune responsiveness and resistance to pathogens in the cabbage looper, Trichoplusia ni. Two baseline and induced immune parameters were estimated, haemocyte numbers and haemolymph phenoloxidase (PO) activity, for larvae on two host plants, broccoli and cucumber. Haemolymph protein concentration was assessed as an indication of insect condition, and the susceptibility of larvae to T. ni single nucleopolyhedrovirus (SNPV) was used as a measure of disease resistance. T. ni growth, survival and condition was much higher on broccoli than cucumber. Haemocyte numbers were significantly higher in broccoli-reared larvae, whereas PO activity was not. An immune challenge induced significantly elevated numbers of haemocytes for larvae reared on both host plants, but did not affect PO activity or protein concentrations. Susceptibility to T. ni SNPV was markedly higher in larvae reared on cucumber than on broccoli. These results clearly indicate that host plant quality can affect both immune response and disease resistance of T. ni larvae and that bottom-up effects could be important in interactions between insects and entomopathogens.     

Cascading effects of variation in plant vigour on the relative performance of insect herbivores and their parasitoids

Abstract 1. Consequences of variation in food plant quality were estimated for a system consisting of two monophagous noctuid herbivores and three ichneumonid parasitoids.
2. In a natural population, pupal weights of the herbivores in this system, Nonagria typhae and Archanara sparganii , were found to be highly variable. Pupal weights increased strongly and consistently with the increase in the vigour of the host plant, Typha latifolia , providing support for the plant vigour hypothesis. Correspondingly, as the moths do not feed as adults, a strong, positive correlation between host vigour and fecundity of the herbivores would be expected.
3. There were strong and positive relationships between adult body sizes of the parasitoids and the sizes of their lepidopteran hosts. Moreover, a direct, positive link between plant quality and parasitoid size was documented.
4. For all three parasitoids, cascading effects of plant quality on body size were weaker than for the herbivores. Differences in the importance of adult feeding and oviposition behaviour suggest that dependence of fitness on body size is also weaker in the parasitoids than in the moths. It is therefore concluded that the numerical response of the herbivore population to a change in plant quality should exceed the corresponding response in the parasitoids.
5. The results of this work imply that variation in plant variables may affect performance of different trophic levels to a different extent. It is suggested that the importance of adult feeding for the reproductive success (capital vs. income breeding strategies) in both herbivores and parasitoids is an essential aspect to consider when predicting responses of such a system to changes in plant quality.     

Field tests with insecticides and insect growth regulators to control insect pests of cowpea and its effects on certain beneficial insects

A small‐plot tests were conducted on cowpea, Vigna ungiculata (L.) Walp, to determine the effectiveness of 14 selected insecticides representing four insecticide classes (organophosphates, carbamates, pyrethroids, and insect growth regulators, IGRs), and four insecticide/IGR mixtures on cowpea insect pests and its effects on certain beneficial insects. By day 3 after treatement, the insecticides phenthoate, isoxathion, cyanophos, carbaryl, and cypermethrin used at sub‐recommended rates reduced the leafhopper, Empoasca spp., populations by >83% than those in the control. On that day, all insecticide treatments significantly reduced the cowpea aphid, Aphis craccivora, numbers below that of the control. The prothiofos, isoxathion, pirimicarb, and fenpropathrin treatments provided continuing control to both leafhoppers and aphids through day 21 after spraying. It seems that most insecticide treatments were not effective in whitefly, Bemisia tabaci, control. The best control of the whitefly immatures was obtained by day 3 after spraying in plots received thiodicarb (76%) and fenpropathrin (60%). All selected insecticides and rates used had very low residual effect against B. tabaci immatures. By day 3 posttreatment, the carbaryl (2.02 kg/ha) caused completely protection for cowpea pods frMw blue pea butterfly, Lampides boeticus, larvae. The IGRs, flufenoxuron and Dowco‐439, applied alone and those applied in mixtures with insecticides, carbaryl/teflubenzuron, chlorpyrifos/XRD‐473, and methomyl/ flufenoxuron, exhibited satisfactory control (>81%) to butterfly larvae on day 7 posttreatment. All treatments did not exhibit a considerable residual activity against L. boeticus larvae through 2–3 weeks followed spray. Most insecticides applied at the higher rates used significantly reduced the numbers of limabean pod borer, Etiella zinckenella, larvae found in cowpea pods collected on day 7 after application. The IGRs, flufenoxuron and Dowco‐439, applied alone, at 0.119 kg/ha, or in mixtures, methomyl/flufenoxuron (0.167 kg/ha) and chlorpyrifos/Dowco‐439 (0.911 kg/ha) caused >73% control of E. zinckenella larvae through day 21 posttreatment. The best control of cowpea weevil, Callosobru‐chus spp., adults (>83%), on day 3 after spraying, was obtained in treatments of cyanophos (1.19 kg/ha), prothiofos (1.43 kg/ha), flufenoxuron (0.119 kg/ha), and chlorpyrifos/Dowco‐439 (0.911 kg/ha). The IGR flufenoxuron applied alone or in mixture, methomyl/flufenoxuron (0.164 kg/ ha) exhibited satisfactory residual activity against Callosobruchus adults through day 21 posttreatment. Percentage seed damage by the larvae of cowpea weevils was significantly lower in most treatments than that of control. Populations of insect predators found in all treated plots were extremely reduced than those found in untreated plots. By day 21 after spraying, the IGRs flufenoxuron, XRD‐473, and Dowco‐439, applied at the low rate of 0.071 kg/ha, seemed to be less effect against insect predators than other insecticides used.     

Mechanisms and diversity of resistance to insect pests in wild relatives of groundnut

The levels of resistance to insect pests in cultivated groundnut (Arachis hypogaea) germplasm are quite low, and therefore, we screened 30 accessions of Arachis spp. and 12 derived lines for resistance to insect pests under field and greenhouse conditions. Accessions belonging to Arachis cardenasii, Arachis duranensis, Arachis kempff-mercadoi, Arachis monticola, Arachis stenosperma, Arachis paraguariensis, Arachis pusilla, and Arachis triseminata showed multiple resistance to the leaf miner Aproaerema modicella, Helicoverpa armigera, Empoasca kerri, and to rust, Puccnia arachidis Speg., and late leaf spot, Cercosporidium personatum (Berk. et Curt.). Arachis cardenasii (ICG 8216), Arachis ipaensis (ICG 8206), A. paraguariensis (ICG 8130), and Arachis appressipila (ICG 8946) showed resistance to leaf feeding and antibiosis to Spodoptera litura under no-choice conditions. Six lines, derived from wild relatives, showed resistance to H. armigera and S. litura, and/or leaf miner. Plant morphological characteristics such as main stem thickness, hypanthium length, leaflet shape and length, leaf hairiness, standard petal length and petal markings, basal leaflet width, main stem thickness and hairiness, stipule adnation length and width, and peg length showed significant correlation and/or regression coefficients with damage by H. armigera, S. litura, and leafhoppers, and these traits can possibly be used as markers to select for resistance to these insect pests. Principal component analysis placed the Arachis spp. accessions into five groups, and these differences can be exploited to diversify resistance to the target insect pests in groundnut.     

Differential susceptibility and suitability of domestic and wild apple species for a florivorous weevil and its parasitoids

Crop plant domestication can change plant resistance to herbivores leading to differences in pest pressure experienced by crop plants and their wild relatives. To compare resistance to herbivores between domesticated and wild fruit trees, we quantified direct resistance and indirect resistance to a pest insect, the florivorous apple blossom weevil Anthonomus pomorum (Coleoptera: Curculionidae), for the cultivated apple Malus domestica and two wild apple species, the European crab apple M. sylvestris and the exotic M. kirghisorum. We measured weevil infestation and performance (weight, sex ratio), and weevil parasitism by parasitoid wasps for different cultivars of M. domestica and for the two wild apple species. To explain weevil and parasitoid responses to different apple species, we quantified tree characteristics including nitrogen content, size of flower buds, bark roughness, tree size, tree phenology and tree position. We found significant differences in susceptibility to weevil infestation between apple species, with lowest infestation (highest apple resistance) in M. domestica and highest infestation in M. kirghisorum. The suitability of apple species also varied significantly: weevils emerging from M. sylvestris were significantly lighter than those from M. kirghisorum. Parasitism of A. pomorum by different parasitoid species was significantly higher in M. sylvestris than in M. domestica. Infestation, weevil weight and parasitism were positively related to tree characteristics: infestation to bud nitrogen content and bark roughness, weevil size to nitrogen content and bud size, and parasitism to tree height and bud density. Our study revealed marked differences between apple species in susceptibility and suitability for the pest herbivore, but also for antagonistic parasitoids. Whereas direct resistance appeared to be higher in cultivated apple, indirect resistance via parasitoids was apparently higher in wild apple trees. Our findings suggest that wild and cultivated apple trees possess different resistance traits that may be combined to optimize resistance in commercial apple cultivars.     

Compound effects of induced plant responses on insect herbivores and parasitoids: implications for tritrophic interactions

1. Induced plant responses can affect herbivores either directly, by reducing herbivore development, or indirectly, by affecting the performance of natural enemies. Both the direct and indirect impacts of induction on herbivore and parasitoid success were evaluated in a common experimental system, using clonal poplar trees Populus nigra (Salicales: Salicaceae), the gypsy moth Lymantria dispar (L.) (Lepidoptera: Lymantriidae), and the gregarious parasitoid Glyptapanteles flavicoxis (Marsh) (Hymenoptera: Braconidae). 2. Female parasitoids were attracted to leaf odours from both damaged and undamaged trees, however herbivore‐damaged leaves were three times more attractive to wasps than undamaged leaves. Parasitoids were also attracted to herbivore larvae reared on foliage and to larval frass, but they were not attracted to larvae reared on artificial diet. 3. Prior gypsy moth feeding elicited a systemic plant response that retarded the growth rate, feeding, and survival of gypsy moth larvae, however induction also reduced the developmental success of the parasitoid. 4. The mean number of parasitoid progeny emerging from hosts fed foliage from induced trees was 40% less than from uninduced trees. In addition, the proportion of parasitised larvae that survived long enough to issue any parasitoids was lower on foliage from induced trees. 5. A conceptual and analytical model is provided to describe the net impacts of induced plant responses on parasitoids, and implications for tritrophic interactions and biological control of insect pests are discussed.     

Egg distribution of insect parasitoids: a survey of models

A number of (insect) parasitoids have been found to avoid superparasitism, i.e., these parasitoids distribute their eggs more evenly over the available hosts than might be expected from chance only. By doing so each parasitoid individual ensures a greater probability of survival for its offspring as a result of a reduced within-host-competition.Recently a number of mathematical models have been developed, describing the distribution of the parasitoid eggs in the hosts. This paper gives a survey of these models, placing them within one and the same mathematical framework. An essential conceptual distinction, neglected up to now, emerges: parasitoids can either react to the number of previous visits to a particular host, or they can react to the number of eggs already present in that host.For each model the probability-generating function, the mean, the variance, and the probability distribution of the number of eggs are given, as well as a discussion of estimating and testing procedures. A few possibilities for generalizations of these models are discussed too.     

Induction of streptomycin resistance in the wild tomato Lycopersicon peruvianum

Summary A protoplast mutagenesis and cell selection system was used for the isolation of streptomycin resistant Lycopersicon peruvianum colonies. Protoplasts were treated with the mutagen N-nitroso-methylurea and could be regenerated into fertile plants, carrying the streptomycin resistant character. Several classes of streptomycin resistance could be distinguished. Reciprocal crosses between streptomycin resistant and sensitive plants showed a non-Mendelian transmission of the resistance trait. Streptomycin resistance is the first selectable and maternally inherited cell organelle marker described in tomato.     

Field evaluation of the effects of the insect growth regulator tebufenozide on entomophagous arthropods and pests of apples

Organophosphate insecticides are very widely used in commercial apple production to control fruit-attacking pests but their broad-spectrum activity constrains biological control of other pests. Compounds with narrow-spectrum activity are therefore desirable. The insect growth regulator (IGR) tebufenozide was compared with another IGR, fenoxycarb, and the organophosphate, azinphos-methyl, in a replicated field trial in the 1994/1995 apple-production season. Vacuum sampling of the tree foliage on five occasions during the growing season showed significantly lower populations of various natural enemies (spiders, lacewings and the specialist mite predator Stethorus spp. adults and larvae) in the azinphos-methyl treatment than in either of the two IGR treatments. The two-spotted mite ( Tetranychus urticae Koch) was most numerous in the azinphos-methyl treatment. In 1995/1996, the entire trial area was placed under a tebufenozide treatment program to determine the extent to which natural enemy populations would recover when broad-spectrum insecticide (azinphos-methyl) use was halted. Populations of polyphagous natural enemies assumed levels broadly equivalent to those observed under IGR treatments the previous year. Numbers of Stethorus spp. were lower than in the 1994/1995 season, possibly because T. urticae (prey) populations were much reduced from the previous season's densities. All three insecticide treatments were equally effective in controlling the lepidopteran pests, codling moth ( Cydia pomonella (L.)), lightbrown apple moth ( Epiphyas postvittana (Walker)) and early season caterpillars (predominantly Helicoverpa punctigera (Wallengren)). Results indicate that tebufenozide provides good control of lepidopteran pests, while allowing the rapid build-up of natural enemies which contribute to control of other pests.     

Patterns of parasitism by insect parasitoids in patchy environments

Abstract. 1. This paper shows how the different spatial patterns of per cent parasitism in patches of different host density can be explained within a single model framework that takes into account the parasitoid's aggregative response, and the factors limiting the degree of host exploitation within patches.
2. Two contrasting laboratory examples are presented in which the distribution of searching parasitoids and the resulting levels of parasitism in different patches are both known for a range of parasitoid densities.
3. A model is described predicting the number of hosts parasitized per patch, in which the number of parasitoids searching is determined from a simple expression allowing different degrees of aggregation.
4. The model generates patterns of parasitism encompassing the two laboratory examples and a wide range of examples from the field.
5. The importance of density dependent spatial distributions of parasitism to population stability is briefly discussed.     

Life history of insect parasitoids involved in successful multiparasitism

Summary The occurrence of multiple parasitoid emergence from individual hosts was observed on numerous occasions. Characteristically such a phenomenon is extremely rare. In the observed instances of successful multiple parasitism of Autographa californica the parasitoid life history attributes of partial host consumption, gregariousness and timing of host parasitism are suggested as traits which contributed to certain species being able to coexist within individual hosts.Oregon State University, Agricultural Experiment Station Technical Paper No. 6239     

Selection on insect immunity in the wild

The strength of selection on immune function in wild populations has only been examined in a few vertebrate species. We report the results from a study measuring selection on a key insect immune enzyme, phenoloxidase (PO), in a wild population of the damselfly Calopteryx xanthostoma. We followed individually marked males from the pre-reproductive adult phase and recorded their lifetime mating success. We found positive selection on PO activity in response to an immune insult, but no selection on wing-spot quality, a trait actively displayed to females during courtship. We suggest that positive selection on PO activity in the year of study may be explained by annual fluctuations in parasite loads.     

Marker-assisted introgression of blackmold resistance QTL alleles from wild Lycopersicon cheesmanii to cultivated tomato (L. esculentum) and evaluation of QTL phenotypic effects

Blackmold, caused by the fungus Alternaria alternata, is a major ripe fruit disease of processing tomatoes. Previously, we found blackmold resistance in a wild tomato (Lycopersicon cheesmanii) and quantitative trait loci (QTL) for resistance were mapped in an interspecific population. Five QTLs were selected for introgression from L. cheesmanii into cultivated tomato using marker-assisted selection (MAS). Restriction fragment length polymorphism and PCR-based markers flanking, and within, the chromosomal regions containing QTLs were used for MAS during backcross and selfing generations. BC₁ plants heterozygous at the QTLs, and subsequent BC₁S₁ and BC₁S₂ lines possessing different homozygous combinations of alleles at the target QTLs, were identified using DNA markers. Field experiments were conducted in 1998 (with 80 marker-selected BC₁S₂ lines) and 1999 (with 151 marker-selected BC₁S₂ and BC₁S₃ lines) at three California locations. Blackmold resistance was assessed during both years, and horticultural traits were evaluated in 1999. The BC₁S₂ and BC₁S₃ lines containing L. cheesmanii alleles at the QTLs were associated with a large genetic variance for resistance to blackmold and moderate heritability, suggesting that significant genetic gain may be achieved by selection in this genetic material. L. cheesmanii alleles at three of the five introgressed QTLs showed a significant, positive effect on blackmold resistance. A QTL on chromosome 2 had the largest positive effect on blackmold resistance, alone and in combination with other QTLs, and was also associated with earliness, a positive horticultural trait. The other four QTLs were associated primarily with negative horticultural traits. Fine mapping QTLs using near isogenic lines could help determine if such trait associations are due to linkage drag or pleiotropy.     

Multiple insect resistance in transgenic tomato plants over-expressing two families of plant proteinase inhibitors

Protease inhibitors have been proposed as potential defense molecules for increased insect resistance in crop plants. Compensatory over-production of insensitive proteases in the insect, however, has limited suitability of these proteins in plant protection, with very high levels of inhibitor required for increased plant resistance. In this study we have examined whether combined used of two inhibitors is effective to prevent this compensatory response. We show that leaf-specific over-expression of the potato PI-II and carboxypeptidase inhibitors (PCI) results in increased resistance to Heliothis obsoleta and Liriomyza trifolii larvae in homozygote tomato lines expressing high levels (#62;1 the total soluble proteins) of the transgenes. Leaf damage in hemizygous lines for these transformants was, however, more severe than in the controls, thus evidencing a compensation response of the larvae to the lower PI concentrations in these plants. Development of comparable adaptive responses in both insects suggests that insect adaptation does not entail specific recognition of the transgene, but rather represents a general adaptive mechanism triggered in response to the nutritional stress imposed by sub-lethal concentrations of the inhibitors. Combined expression of defense genes with different mechanisms of action rather than combinations of inhibitors may then offer a better strategy in pest management as it should be more effective in overcoming this general adaptive response in the insect.     

Detrimental and neutral effects of wild barley–Neotyphodium fungal endophyte associations on insect survival

Neotyphodium (Clavicipitaceae: Balansieae) fungal endophyte infection does not always confer temperate grass resistance to insect herbivores, although reports indicate that over 40 species are adversely affected by its infection. Laboratory and glasshouse experiments were conducted to improve our knowledge of the anti‐insect properties of Neotyphodium‐infected (E+) non‐commercial grasses, and E+ wild barley (Hordeum) specifically. Neotyphodium infection of four plant inventory (PI) lines of wild barley conferred resistance to Mayetiola destructor (Say) (Diptera: Cecidomyiidae), whereas none of the E+ wild barley accessions reduced the survival of Rhopalosiphum padi (L.) (Homoptera: Aphididae). Metopolophium dirhodum (Walker) (Homoptera: Aphididae) densities were significantly lower on the E+ clones of Hordeum brevisubulatum ssp. violaceum (Boissier and Hohenacker) (PI 440420), compared with densities on endophyte‐free (E–) plants of this species in population growth experiments. Neotyphodium infection of three Hordeum bogdanii (Wilensky) PI lines did not confer resistance to M. dirhodum; however, one of these E+ lines (PI 314696) was resistant to this aphid in a second population growth experiment. Our results provide additional evidence that the outcome of a grass–endophyte–insect interaction is influenced by the host grass species or genotype, Neotyphodium species or genotype, and the insect species involved. They also reinforce this phenomenon for non‐commercial grass–endophyte–insect interactions and underscore the potential role of endophytes in mediating wild barley–insect interactions and their potential to act as defensive mutualists.     

昆虫抗药性机制及抗性治理研究进展

化学杀虫剂的广泛使用,使昆虫产生抗药性的现象日渐突出。抗药性是一种潜在的、强大的、普遍的自然现象。研究昆虫的抗药性对于农林害虫防控、新型杀虫剂研发、生态环境的多样性等具有重要的意义。近年来随着昆虫基因组学的发展,昆虫抗药性研究取得突破性进展,本文介绍了基因组学在昆虫抗药性研究中的应用,着重概述了昆虫抗药性形成的分子机制以及其治理策略,并对今后如何延缓抗性的发展进行了探讨和展望。