High-density populations of diamondback moth have broader host-plant diets

Although ecological specialists exploit a relatively limited resource base, it is unclear whether specialization limits local population density. Here, we focus on the relationship between diet specialization and local population density of a phytophagous insect, the diamondback moth, Plutella xylostella L. (Lepidoptera: Plutellidae). We compared key life history traits and food plant use among five pairs of high- and low-density populations from widely separate geographical areas. Moths from populations that persist at high densities were more generalized in food plant use than moths from low-density populations. In particular, the oviposition preference and larval performance of moths from some high-density populations were less variable across a suite of food plants, suggesting that moths from high- density populations had a broader diet. In addition, low- density populations were less similar to each other, exhibiting opposing preferences for particular plant species. Hence diet breadth may explain some of the persistent differences in the population density of diamondback moths in the field, consistent with the idea that ecological specialization may be generally associated with population density. Received: 19 December 1996 / Accepted: 2 May 1997     

Does the age of the host plant modulate migratory activity of Plutella xylostella?

Ephemeral plants and most tropical crops are available to herbivore insects as irregular and unpredictable patches of resources. Insects that exploit patched habitats usually have well‐developed migratory ability, and migration is a fundamental process in the cycle of colonization and extinction of the local populations. We ask here whether Plutella xylostella (Lepidoptera: Yponomeutidae) uses the aging process of the host plant to modulate its migratory activity. The insect was reared in the laboratory on the leaves from the middle estratum (fourth expanded leaves) of cabbage (Brassica oleraceae var. capitata) of various ages, and on leaves from three strata of the mature plant. Females that spent their larval phase in young and tender plants lived for a shorter period of time and laid eggs immediately after metamorphosis. In contrast, mature plants, particularly the new leaves from the superior stratum, favored the development of adult phenotypic traits that occur in migratory forms, such as reduced body size, increased longevity and delayed reproductive activity. We suggest that the lesser nutritional quality and the short temporal persistence of mature plants are selective forces favoring individuals that are better prepared to abandon their habitat soon after emergence. Plutella xylostella may use the predictable ontogenetic changes of the ephemeral host plant to modulate its physiological and behavioral migratory ability.     

Effects of adult‐derived carbohydrates and amino acids on the reproduction of Plutella xylostella

The use of floral resources to improve the efficiency of natural enemies has grown in recent years, although their effect on pest populations has been overlooked. To understand how access to food resources by adults of Plutella xylostella (L.) (Lepidoptera: Plutellidae) affects their reproduction, the effects of amino acids and carbohydrates (i.e. fructose, glucose and sucrose) on the longevity, lifetime fecundity, egg viability and population growth of this important pest of brassicas are evaluated. Carbohydrate intake increases longevity, oviposition period and lifetime fecundity of P. xylostella. Oviposition period and fecundity are six‐fold higher in females that consumed carbohydrates. By contrast, amino acid intake by adults does not affect reproductive parameters, even when mixed with carbohydrates. The estimated demographic parameters indicate that the adult diet is important for pest population growth because the net reproductive rate and the intrinsic rate of increase are higher when females have access to carbohydrates. These results are important from both physiological and ecological points of view. The increasing use of flowering plants as a microhabitat and food source for natural enemies may also result in benefits for herbivorous insects, compromising any success in pest control.     

Trichome production and variation in young plant resistance to the specialist insect herbivore Plutella xylostella among natural populations of Arabidopsis lyrata

The strength of plant‐herbivore interactions varies spatially and through plant ontogeny, which may result in variable selection on plant defense, both among populations and life‐history stages. To test whether populations have diverged in herbivore resistance at an early plant stage, we quantified oviposition preference and larval feeding by Plutella xylostella (L.) (Lepidoptera: Plutellidae) on young (5–6 weeks old) Arabidopsis lyrata (L.) O'Kane & Al‐Shehbaz (Brassicaceae) plants, originating from 12 natural populations, six from Sweden and six from Norway. Arabidopsis lyrata can be trichome‐producing or glabrous, with glabrous plants usually receiving more damage from insect herbivores in natural populations. We used the six populations polymorphic for trichome production to test whether resistance against P. xylostella differs between the glabrous and the trichome‐producing morph among young plants. There was considerable variation among populations in the number of eggs received and the proportion of leaf area consumed by P. xylostella, but not between regions (Sweden vs. Norway) or trichome morphs. Rosette size explained a significant portion of the variation in oviposition and larval feeding. The results demonstrate that among‐population variation in resistance to insect herbivory can be detected among very young individuals of the perennial herb A. lyrata. They further suggest that trichome densities are too low at this plant developmental stage to contribute to resistance, and that the observed among‐population variation in resistance is related to differences in other plant traits.     

Cabbage Seasonal Leaf Quality Mediating the Diamondback Moth Plutella xylostella (L.) (Lepidoptera: Yponomeutidae) Performance

Seasonal variation in plant quality may be intense enough to generate predictable patterns in insect herbivore populations. In order to explain seasonal oscillations in neotropical populations of the diamondback moth Plutella xylostella (L.), we tested the following: (1) if nutritional quality of cabbage (Brassica oleraceae var. capitata), a primary host plant of diamondback moth, adversely affects the performance of this insect in late spring and early summer, when populations decline and go extinct, and (2) if nutritional features of cabbage change with the seasons. We measured the performance of diamondback moth reared on leaves of cabbages grown during the four seasons of the year. Summer plants proved to be worse for the survival of the immature stages and subsequent adult fecundity, but there were no significant differences between the remaining seasons. Our results support the hypothesis that short-lived plants, grown in different seasons of the year in the tropics, have different nutritional and defensive attributes. We analyzed nutritional quality of cabbage leaves from the four seasons, but only total lipids were reduced in summer plants. Neotropical populations of diamondback moth collapse before plant quality decay in the summer. If the diamondback moth is well adapted to the seasonal deterioration of the habitat, including the reduction in the quality of host plants, it is expected that emigration happens before the mortality increases and natality decreases during the summer.     

Effect of plant patch shape on the distribution and abundance of three lepidopteran species associated with Brassica oleracea

• 1 Plant patch shape may affect the abundance of herbivorous insects. Patches of the same size but longer or irregular have a higher perimeter/area relationship (P/A) than square or regular ones, which may determine the immigration, emigration and abundance of individuals in the patch.
• 2 Only specialist species should be affected by plant patch shape. Those species that are more abundant in smaller patches should be more abundant in patches with higher P/A, whereas those that are more abundant in larger patches should be more abundant in patches with lower P/A.
• 3 We studied the density of eggs, larvae and pupae of Pieris brassicae, Plutella xylostella and Trichoplusia ni in square (low P/A) and I‐shaped (high P/A) patches of 144 plants of Brassica oleracea. We also estimated their immigration to these patches, and the final plant weight.
• 4 Plant patch shape affected the abundance, but not the distribution, of the two specialist species. Whereas P. brassicae was denser in I‐shaped patches, P. xylostella was more abundant in square patches. The generalist T. ni was not affected by patch shape. Immigration of P. brassicae was higher in I‐shaped patches, but immigration of P. xylostella and T. ni was not affected by patch shape. Plants were heavier in the centre of square patches.
• 5 Our results suggest that plant patch shape affects the density of herbivorous insects and should be considered independently from other plant patch variables when studying the population dynamics of these organisms.

     

Inhibitory effects of permethrin on flight responses,host‐searching,and foraging behaviour of Cotesia vestalis (Hymenoptera: Braconidae), a larval parasitoid of Plutella xylostella (Lepidoptera: Plutellidae)

Although it is well known that the application of broad‐spectrum synthetic insecticides reduces the effectiveness of natural enemies, the details of the actual mechanisms, including the lethal and sublethal effects of this reduction, are not fully understood. The inhibitory effects of a pyrethroid insecticide (permethrin), Adion 20% EC on the flight responses, host‐searching behaviour and foraging behaviour of Cotesia vestalis (Hymenoptera: Braconidae), a larval parasitoid of the diamondback moth, Plutella xylostella (Lepidoptera: Plutellidae), were investigated under laboratory conditions. In choice trials, the wasps showed significant preference for P. xylostella‐infested Komatsuna plants over insecticide‐treated plants, suggesting an inhibitory effect of the insecticide on the flight response of C. vestalis. When offered a pair of plants, the wasps showed a significant preference for P. xylostella‐infested plants compared to uninfested plants. However, significantly more wasps were attracted to infested permethrin‐treated plants than to uninfested plants, suggesting that the wasps are attracted to the volatile infochemicals from the infested plants, even if treated with permethrin. The searching time was significantly shorter and the mortality of C. vestalis adults on the insecticide‐treated plants significantly higher than in the control plants treated with distilled water. These results suggest that the application of the insecticide had an inhibitory effect on the wasps’‐searching behaviour and consequently reduced the effectiveness of C. vestalis as a biological control agent against P. xylostella. In addition, the strength of the inhibitory effect of permethrin on the attraction of the wasps to the plants is critical to the survival of C. vestalis. Our results suggest that the attraction of the wasps to the permethrin‐treated infested plants increases the risk of their exposure to this insecticide.     

Indirect effects of an unspecialized endophytic fungus on specialized plant – herbivorous insect interactions

The effects of Acremonium alternatum Gams (Ascomycotina, Clavicipitacea) on the development and nutrition of diamondback moth larvae Plutella xylostella L. (Lepidoptera, Plutellidae) were studied in the laboratory. All experiments were conducted before the endophyte reached the green parts of the plants; thus P. xylostella, a folivore, was not in direct contact with the endophyte. Larvae feeding on leaves of previously inoculated plants suffered from increased mortality, especially during the first 10 days of development. Likewise, during early development surviving larvae had a reduced relative growth rate (RGR), which, however, did not result in reduced pupal or adult weight. We found sexual differences in the food utilization efficiency; female P. xylostella progeny reacted more sensitively to endophytic infection of cabbage than male larvae. Female larvae feeding on leaves of endophyte-infested plants responded to reduced efficiency of conversion of ingested food (ECI) by increasing their relative consumption rate (RCR). The underlying mechanisms for these results are discussed in relation to changes in plant phytosterol metabolism which could account for reduced larval growth on inoculated cabbage plants. Our data suggest that unspecialized, soil-borne endophytic fungi, even when their association with the host plant is weak, can influence aboveground herbivore development and should be considered when investigating plant-insect interactions. Received: 3 November 1997 / Accepted 29 December 1997     

Fitness of the parasitoid Diadegma insulare is affected by its host's food plants

The diamondback moth, Plutella xylostella (L.) is considered a specialist on Brassicaceae, but it is capable of expanding its food range by incorporation of non-brassicaceous plants into its diet. The use of a new food plant may change food availability and vulnerability of P. xylostella to its natural enemies, especially specialist parasitoids. In this laboratory study, we evaluated the bottom-up effects of two Brassicaceae viz. Brassica napus L. and Descurainia sophia (L.) Webb ex Prantl and two non-Brassicaceae viz. Tropaeolum majus L. and Cleome hassleriana Chodat on several fitness correlates of the specialist parasitoid, Diadegma insulare (Cresson) reared on P. xylostella larvae. Percentage of parasitism of host larvae by D. insulare varied among the plant species and was highest on T. majus and lowest on D. sophia. Values of several fitness correlates for D. insulare differed when the host was feeding on various plant species. Egg to adult development was fastest on B. napus followed by C. hassleriana, D. sophia and T. majus. Female wasps reared on C. hasseleriana lived the longest in absence of food. The response of D. insulare to potential food-plant expansion by its host P. xylostella is discussed.     

Engineering of benzylglucosinolate in tobacco provides proof-of-concept for dead-end trap crops genetically modified to attract Plutella xylostella (diamondback moth)

Glucosinolates are biologically active natural products characteristic of crucifers, including oilseed rape, cabbage vegetables and the model plant Arabidopsis thaliana. Crucifer‐specialist insect herbivores, like the economically important pest Plutella xylostella (diamondback moth), frequently use glucosinolates as oviposition stimuli. This suggests that the transfer of a glucosinolate biosynthetic pathway to a non‐crucifer would stimulate oviposition on an otherwise non‐attractive plant. Here, we demonstrate that stable genetic transfer of the six‐step benzylglucosinolate pathway from A. thaliana to Nicotiana tabacum (tobacco) results in the production of benzylglucosinolate without causing morphological alterations. Benzylglucosinolate‐producing tobacco plants were more attractive for oviposition by female P. xylostella moths than wild‐type tobacco plants. As newly hatched P. xylostella larvae were unable to survive on tobacco, these results represent a proof‐of‐concept strategy for rendering non‐host plants attractive for oviposition by specialist herbivores with the long‐term goal of generating efficient dead‐end trap crops for agriculturally important pests.     

Tritrophic Choice Experiments with Bt Plants,the Diamondback Moth (Plutella xylostella) and the parasitoid Cotesia plutellae

Parasitoids are important natural enemies of many pest species and are used extensively in biological and integrated control programmes. Crop plants transformed to express toxin genes derived from Bacillus thuringiensis (Bt) provide high levels of resistance to certain pest species, which is likely to have consequent effects on parasitoids specialising on such pests. A better understanding of the interaction between transgenic plants, pests and parasitoids is important to limit disruption of biological control and to provide background knowledge essential for implementing measures for the conservation of parasitoid populations. It is also essential for investigations into the potential role of parasitoids in delaying the build-up of Bt-resistant pest populations. The diamondback moth (Plutella xylostella), a major pest of brassica crops, is normally highly susceptible to a range of Bt toxins. However, extensive use of microbial Bt sprays has led to the selection of resistance to Bt toxins in P. xylostella. Cotesia plutellae is an important endoparasitoid of P. xylostella larvae. Although unable to survive in Bt-susceptible P. xylostella larvae on highly resistant Bt oilseed rape plants due to premature host mortality, C. plutellae is able to complete its larval development in Bt-resistant P. xylostella larvae. Experiments of parasitoid flight and foraging behaviour presented in this paper showed that adult C. plutellae females do not distinguish between Bt and wildtype oilseed rape plants, and are more attracted to Bt plants damaged by Bt-resistant hosts than by susceptible hosts. This stronger attraction to Bt plants damaged by resistant hosts was due to more extensive feeding damage. Population scale experiments with mixtures of Bt and wildtype plants demonstrated that the parasitoid is as effective in controlling Bt-resistant P. xylostella larvae on Bt plants as on wildtype plants. In these experiments equal or higher numbers of parasitoid adults emerged per transgenic as per wildtype plant. The implications for integrated pest management and the evolution of resistance to Bt in P. xylostella are discussed.     

Assessing risks and benefits of floral supplements in conservation biological control

The use of flowering field margins is often proposed as a method to support biological control in agro-ecosystems. In addition to beneficial insects, many herbivores depend on floral food as well. The indiscriminate use of flowering species in field margins can therefore lead to higher pest numbers. Based on results from field observations and laboratory experiments we assessed risks as well as benefits associated with the provision of nectar plants in field margins, using Brussels sprouts as a model system. Results show that Brussels sprouts bordered by nectar plants suitable for the cabbage white Pieris rapae L., suffered higher infestation levels by this herbivore. In contrast, nectar plants providing accessible nectar for the diamondback moth Plutella xylostella L., did not raise densities of P. xylostella larvae in the Brassica crop. Margins with Anethum graveolens L., selected on the basis of its suitability as nectar plant for parasitoids, significantly increased the number of adult Diadegma semiclausum Hellén in the crop. This didn’t translate into enhanced parasitism rates, as parasitism of P. xylostella by D. semiclausum exceeded 65 % in all treatments, irrespective of the plants in the field margin. Our findings emphasize the importance of taking a multitrophic approach when choosing flowering field margin plants for biocontrol or other ecosystem services.     

Genetic control of Plutella xylostella in omics era

Diamondback moth, Plutella xylostella (L.), is a specialist pest on cruciferous crops of economic importance. The large‐scale use of chemical insecticides for the control of this insect pest has caused a number of challenges to agro‐ecosystems. With the advent of the omics era, genetic pest management strategies are becoming increasingly feasible and show a powerful potential for pest control. Here, we review strategies for using transgenic plants and sterile insect techniques for genetic pest management and introduce the major advances in the control of P. xylostella using a female‐specific RIDL (release of insects carrying a dominant lethal gene) strategy. Further, the advantages of gene drive developed in combination with sex determination and CRISPR/Cas9 systems are addressed, and the corresponding prospects and implementation issues are discussed. It is predictable that under the policy and regulation of professional committees, the genetic pest control strategy, especially for gene drive, will open a new avenue to sustainable pest management not only for P. xylostella but also for other insect pests.     

Evaluation of the impact of natural enemies on Plutella xylostella L. (Lepidoptera: Yponomeutidae) populations on commercial Brassica farms

Abstract 1 Accurate assessment of the impact of natural enemies on pest populations is fundamental to the design of robust integrated pest management programmes. In most situations, diseases, predators and parasitoids act contemporaneously on insect pest populations and the impact of individual natural enemies, or specific groups of natural enemies, is difficult to interpret. These problems are exacerbated in agro‐ecosystems that are frequently disrupted by the application of insecticides. 2 A combination of life‐table and natural enemy exclusion techniques was utilized to develop a method for the assessment of the impact of endemic natural enemies on Plutella xylostella populations on commercial Brassica farms. 3 At two of the experimental sites, natural enemies had no impact on P. xylostella survival, at two other sites, natural enemy impact was low but, at a fifth site, natural enemies drastically reduced the P. xylostella population. 4 The calculation of marginal death rates and associated k‐values allowed the comparison of mortality factors between experimental sites, and indicated that larval disappearance was consistently the most important mortality factor, followed by egg disappearance, larval parasitism and pupal parasitism. The appropriateness of the methods and assumptions made to calculate the marginal death rates are discussed. 5 The technique represents a robust and easily repeatable method for the analysis of the activity of natural enemies of P. xylostella, which could be adapted for the study of other phytophagous pests.     

Finish line plant–insect interactions mediated by insect feeding mode and plant interference: a case study of Brassica interactions with diamondback moth and turnip aphid

There are gaps in our understanding of plant responses under different insect phytophagy modes and their subsequent effects on the insect herbivores’ performance at late season. Here we compared different types of insect feeding by an aphid, Lipaphis erysimi, and a lepidopteran, Plutella xylostella, and how this affected defensive metabolites in leaves of 2 Brassica species when plants gain maturity. Thiocyanate concentrations after P. xylostella and L. erysimi feeding activities were the same. Total phenolics was higher after the phloem feeder feeding than the folivore activity. The plants compensatory responses (i.e., tolerance) to L. erysimi feeding was significantly higher than the responses to P. xylostella. This study showed that L. erysimi had higher carbon than P. xylostella whereas nitrogen in P. xylostella was 1.42 times that in L. erysimi. Population size of the phloem feeder was not affected by plant species or insect coexistence. However, there was no correlation between plant defensive metabolites and both insects’ population size and biomass. This suggests that plant root biomass and tolerance index after different insect herbivory modes are not necessarily unidirectional. Importantly, the interaction between the folivore and the phloem feeder insects is asymmetric and the phloem feeder might be a trickier problem for plants than the folivore. Moreover, as both plants’ common and special defenses decreased under interspecific interference, we suggest that specialist insect herbivores can be more challenged in ecosystems in which plants are not involved in interspecific interference.     

Host plant nutritional quality affects the performance of the parasitoid Diadegma insulare

The well documented biochemical profile of Brassicaceae, oligophagy of the herbivore Plutella xylostella (L.) (Lepidoptera: Plutellidae), and host specialization of the parasitoid Diadegma insulare (Cresson) (Hymenoptera: Ichneumonidae) provide an ideal system for investigating tritrophic interactions mediated by nutritional quality of plants. We evaluated the bottom-up effects of five soil fertility regimes on nutritional quality of canola (Brassica napus L.) and then on several fitness correlates of female and male D. insulare as mediated through P. xylostella. Variation in soil fertility influenced the nutritional quality of host plants and this in turn affected the performance of D. insulare. In general, D. insulare performed best on plants grown with 3.0 g fertilizer pot⁻¹; these plants had 2.06-, 3.77-, and 1.02-fold more nitrogen, phosphorous and potassium, respectively than ones grown without any added fertilizer. P. xylostella escape from D. insulare was highest (32%) on plants grown at 1.0 g fertilizer, and this could be attributed to both physical and physiological defense mechanisms mediated by host plant nutritional quality. Plant stress and plant vigor are competing paradigms pertaining to the performance of herbivorous insects on their host plants. These hypotheses were originally proposed to predict responses of herbivores, but may also explain the effects of plant quality on koinobiont parasitoids, such as D. insulare.     

Microsatellites isolated from diamondback moth,Plutella xylostella (L.), for studies of dispersal in Australian populations

Diamondback moth, Plutella xylostella (L.), is a worldwide agricultural pest that has developed resistance to many insecticides used for its control. Population structure and gene flow are yet to be determined for P. xylostella in Australia, but are important factors for the design of effective control strategies. We have isolated six polymorphic microsatellite markers: three from a partial genomic library, two from an Expressed Sequence Tagged library and one from an aminopeptidase intron of P. xylostella. These microsatellites will be used to determine population structure and gene flow in Australian populations of P. xylostella to improve insecticide resistance management.     

Suitability of Arabidopsis thaliana as a model for host plant–Plutella xylostella–Cotesia plutellae interactions

Successful pest management is often hindered by the inherent complexity of the interactions of a pest with its environment. The use of genetically characterized model plants can allow investigation of chosen aspects of these interactions by limiting the number of variables during experimentation. However, it is important to study the generic nature of these model systems if the data generated are to be assessed in a wider context, for instance, with those systems of commercial significance. This study assesses the suitability of Arabidopsis thaliana (L.) Heynh. (Brassicaceae) as a model host plant to investigate plant–herbivore–natural enemy interactions, with Plutella xylostella (L.) (Lepidoptera: Plutellidae), the diamondback moth, and Cotesia plutellae (Kurdjumov) (Hymenoptera: Braconidae), a parasitoid of P. xylostella. The growth and development of P. xylostella and C. plutellae on an A. thaliana host plant (Columbia type) were compared to that on Brassica rapa var. pekinensis (L.) (Brassicaceae), a host crop that is widely cultivated and also commonly used as a laboratory host for P. xylostella rearing. The second part of the study investigated the potential effect of the different A. thaliana background lines, Columbia and Landsberg (used in wider scientific studies), on growth and development of P. xylostella and C. plutellae. Plutella xylostella life history parameters were found generally to be similar between the host plants investigated. However, C. plutellae were more affected by the differences in host plant. Fewer adult parasitoids resulted from development on A. thaliana compared to B. rapa, and those that did emerge were significantly smaller. Adult male C. plutellae developing on Columbia were also significantly smaller than those on Landsberg A. thaliana.     

Plant‐mediated interactions between two herbivores differentially affect a subsequently arriving third herbivore in populations of wild cabbage

• Plants are part of biodiverse communities and frequently suffer from attack by multiple herbivorous insects. Plant responses to these herbivores are specific for insect feeding guilds: aphids and caterpillars induce different plant phenotypes. Moreover, plants respond differentially to single or dual herbivory, which may cascade into a chain of interactions in terms of resistance to other community members. Whether differential responses to single or dual herbivory have consequences for plant resistance to yet a third herbivore is unknown.
• We assessed the effects of single or dual herbivory by Brevicoryne brassicae aphids and/or Plutella xylostella caterpillars on resistance of plants from three natural populations of wild cabbage to feeding by caterpillars of Mamestra brassicae. We measured plant gene expression and phytohormone concentrations to illustrate mechanisms involved in induced responses.
• Performance of both B. brassicae and P. xylostella was reduced when feeding simultaneously with the other herbivore, compared to feeding alone. Gene expression and phytohormone concentrations in plants exposed to dual herbivory were different from those found in plants exposed to herbivory by either insect alone. Plants previously induced by both P. xylostella and B. brassicae negatively affected growth of the subsequently arriving M. brassicae. Furthermore, induced responses varied between wild cabbage populations.
• Feeding by multiple herbivores differentially activates plant defences, which has plant‐mediated negative consequences for a subsequently arriving herbivore. Plant population‐specific responses suggest that plant populations adapt to the specific communities of insect herbivores. Our study contributes to the understanding of plant defence plasticity in response to multiple insect attacks.