Fitness costs associated with Cry1F resistance in the European corn borer

Crops producing insecticidal toxins derived from the bacterium Bacillus thuringiensis (Bt) are widely planted to manage insect pests. Bt crops can provide an effective tool for pest management; however, the evolution of Bt resistance can diminish this benefit. The European corn borer, Ostrinia nubilalis Hübner, is a significant pest of maize and is widely managed with Bt maize in the Midwest of the United States. When Bt crops are grown in conjunction with non‐Bt refuges, fitness costs of Bt resistance can delay the evolution of resistance. Importantly, fitness costs often vary with ecological factors, including host‐plant genotype and diapause. In this study, we examined fitness costs associated with Cry1F resistance in O. nubilalis when insects were reared on three maize lines. Fitness costs were tested in two experiments. One experiment assessed the fitness costs when Cry1F‐resistant and Cry1F‐susceptible insects were reared on plants as larvae and experienced diapause. The second experiment tested resistant, susceptible and F1 heterozygotes that were reared on plants but did not experience diapause. Despite some evidence of greater adult longevity for Cry1F‐resistant insects, these insects produced fewer fertile eggs than Cry1F‐susceptible insects, and this occurred independent of diapause. Reduced fecundity was not detected among heterozygous individuals, which indicated that this fitness cost was recessive. Additionally, maize lines did not affect the magnitude of this fitness cost. The lower fitness of Cry1F‐resistant O. nubilalis may contribute to the maintenance of Cry1F susceptibility in field populations more than a decade after Cry1F maize was commercialized.     

The impact of secondary pests on Bacillus thuringiensis (Bt) crops

The intensification of agriculture and the development of synthetic insecticides enabled worldwide grain production to more than double in the last third of the 20th century. However, the heavy dependence and, in some cases, overuse of insecticides has been responsible for negative environmental and ecological impacts across the globe, such as a reduction in biodiversity, insect resistance to insecticides, negative effects on nontarget species (e.g. natural enemies) and the development of secondary pests. The use of recombinant DNA technology to develop genetically engineered insect‐resistant crops could mitigate many of the negative side effects of insecticides. One such genetic alteration enables crops to express toxic crystalline (Cry) proteins from the soil bacteria Bacillus thuringiensis (Bt). Despite the widespread adoption of Bt crops, there are still a range of unanswered questions concerning longer term agro‐ecosystem interactions. For instance, insect species that are not susceptible to the expressed toxin can develop into secondary pests and cause significant damage to the crop. Here, we review the main causes surrounding secondary pest dynamics in Bt crops and the impact of such outbreaks. Regardless of the causes, if nonsusceptible secondary pest populations exceed economic thresholds, insecticide spraying could become the immediate solution at farmers’ disposal, and the sustainable use of this genetic modification technology may be in jeopardy. Based on the literature, recommendations for future research are outlined that will help to improve the knowledge of the possible long‐term ecological trophic interactions of employing this technology.     

Determining the major Bt refuge crops for cotton bollworm in North China

Evaluation of the effectiveness of refuge strategies involved in cotton bollworm Bt resistance management would be aided by technologies that allow monitoring and quantification of key factors that affect the process under field conditions. We hypothesized that characterization of stable carbon and nitrogen isotopes in adult bollworm, Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) moths may aid in determining the larval host that they developed upon. We found moths reared from larvae fed on peanut, soybean or cotton, respectively, could be differentiated using isotopic analyses that also corresponded to their respective larval host origins. These techniques were then used to classify feral second‐generation bollworm moths caught in Bt cotton (Gossypium hirsutum) fields into different populations based on their isotopic signatures. In 2006–2007 feral moths captured in Bt cotton fields predominantly correlated with the peanut (Arachis hypogea) having served as their larval host, indicating this is the most important refuge crop for Bt‐susceptible bollworm individuals (providing 58%?64% individuals) during independent moth peaks for the second generation in North China. The remaining feral moths correlated with soybean (Glycine max) (0?10%); other C₃ plant (20%?22%) and non‐C₃ plant (12%?14%) host types also provided some Bt‐sensitive moths. Field observations showed that peanut constitutes the primary refuge crop contributing to sustaining Bt‐susceptible moths dispersing into cotton in North China. These results suggest that peanut may be a more effective refuge to sustain Bt‐susceptible bollworm individuals and reduce the risk of development of a Bt‐resistant biotype.     

New wind in the sails: improving the agronomic value of crop plants through RNAi‐mediated gene silencing

RNA interference (RNAi) has emerged as a powerful genetic tool for scientific research over the past several years. It has been utilized not only in fundamental research for the assessment of gene function, but also in various fields of applied research, such as human and veterinary medicine and agriculture. In plants, RNAi strategies have the potential to allow manipulation of various aspects of food quality and nutritional content. In addition, the demonstration that agricultural pests, such as insects and nematodes, can be killed by exogenously supplied RNAi targeting their essential genes has raised the possibility that plant predation can be controlled by lethal RNAi signals generated in planta. Indeed, recent evidence argues that this strategy, called host‐induced gene silencing (HIGS), is effective against sucking insects and nematodes; it also has been shown to compromise the growth and development of pathogenic fungi, as well as bacteria and viruses, on their plant hosts. Here, we review recent studies that reveal the enormous potential RNAi strategies hold not only for improving the nutritive value and safety of the food supply, but also for providing an environmentally friendly mechanism for plant protection.     

Effects of seed mixture sowing with transgenic Bt rice and its parental line on the population dynamics of target stemborers and leafrollers,and non‐target planthoppers

The widespread planting of insect‐resistant crops has caused a dramatic shift in agricultural landscapes, thus raising concerns about the potential impacts on both target and non‐target pests. In this study, we examined the potential effects of intra‐specific seed mixture sowing with transgenic Bt rice (Bt) and its parental non‐transgenic line (Nt) (100% Bt rice [Bt₁₀₀], 5% Nt+95% Bt [Nt₀₅Bt₉₅], 10% Nt+90% Bt [Nt₁₀Bt₉₀], 20% Nt+80% Bt [Nt₂₀Bt₈₀], 40% Nt+60% Bt [Nt₄₀Bt₆₀] and 100% Nt rice [Nt₁₀₀]) on target and non‐target pests in a 2‐year field trial in southern China. The occurrence of target pests, Sesamia inferens, Chilo suppressalis and Cnaphalocrocis medinalis, decreased with the increased ratio of Bt rice, and the mixture ratios with more than 90% Bt rice (Bt₁₀₀ and Nt₀₅Bt₉₅) significantly increased the pest suppression efficiency, with the lowest occurrences of non‐target planthoppers, Nilaparvata lugens and Sogatella furcifera in Nt₁₀₀ and Nt₀₅Bt₉₅. Furthermore, there were no significant differences in 1000‐grain dry weight and grain dry weight per 100 plants between Bt₁₀₀ and Nt₀₅Bt₉₅. Seed mixture sowing of Bt rice with ≤10% (especially 5%) of its parent line was sufficient to overcome potential compliance issues that exist with the use of block or structured refuge to provide most effective control of both target and non‐target pests without compromising the grain yield. It is also expected that the strategy of seed mixture sowing with transgenic Bt rice and the non‐transgenic parental line would provide rice yield stability while decreasing the insecticide use frequency in rice production.     

Determining the host‐plant resistance mechanisms for Mamestra brassicae (Lepidoptera: Noctuidae) pest in cabbage

Six cabbage (Brassica oleracea var. capitata) varieties with different levels of resistance to Mamestra brassicae (Lepidoptera: Noctuidae) were investigated in order to assess whether antibiosis and antixenosis mechanisms are involved in the resistance to this pest or not. Four experiments were conducted to determine the effect of variety and plant ontogeny on larval behaviour, adult oviposition and leaf damages in non‐choice and choice tests. Larval survival, time to development and larval weights differed depending on the varieties and plant stages that we tested. At the pre‐head stage, larval mortality was higher, larvae died faster, time to pupation was shorter, pupae were lighter and the percentage of viable pupae and growth index (GI) values were lower than larvae reared from plants at the head stage. The commercial hybrid ‘Corazón de buey’ and the local variety named ‘BRS0535’ exhibited antibiosis to M. brassicae as they reduced its survival and growth and delayed its development time. In addition, these varieties were the most resistant after artificial infestation in terms of head foliage consumption and number of larvae per plant. Oviposition tests demonstrated that resistance found in ‘Corazón de buey’ and BRS0535 could be also based on antixenosis mechanisms as they resulted in fewer egg batches on plants, whereas BRS0402 could be classified as resistant because M. brassicae larvae showed less preference for it. Thus, resistance to M. brassicae found in cabbage crops may be due to the joint action of several factors involving antibiosis and antixenosis. We found significant differences in the resistance of BRS0535 depending on the plant ontogeny as it loses its resistance while developing. Further studies are required to identify the mechanism of antibiotic resistance which is present in this variety at the pre‐head stage and the changes that occur in plant defence as it grows.     

Phloem‐specific resistance in Brassica oleracea against the whitefly Aleyrodes proletella

The cabbage whitefly [Aleyrodes proletella L. (Hemiptera: Aleyrodidae)] is becoming a serious pest in Brassica oleracea L. (Brassicaceae) crops. However, almost nothing is known about the interaction of this insect with its host plants. Previous studies have shown differences in the natural occurrence of adults, eggs, and nymphs on the closely related B. oleracea cultivars Christmas Drumhead and Riviera grown in the field. In this study, we aimed to identify the nature of these differences and to gain insight into the resistance mechanisms against A. proletella. We used no‐choice experiments on field‐ and greenhouse‐grown plants to show that the differences between the two cultivars are mainly based on antibiosis (traits that reduce herbivore performance) and not on antixenosis (traits that deter herbivory). This was further supported by laboratory choice experiments that indicated little or no discrimination between the two cultivars based on plant volatiles. We showed that resistance is dependent on plant age, that is, resistance increased during plant development, and is mainly independent of environmental factors. Analysis of probing behaviour revealed that the resistance trait affects A. proletella at the phloem level and that morphological differences between the two cultivars are most likely not involved. We suggest that compounds present in the phloem reduce sap ingestion by the whitefly and that this explains the observed resistance.     

Induced systemic resistance (ISR) against pathogens – a promising field for ecological research

Putative fitness costs provide an explanation for why ISR is induced instead of constitutive, and they might constrain the use of ISR as preventative protection of cultivated plants. Though ISR is mainly elicited by and effective against pathogens, further biotic agents such as leaf-chewing herbivores, leaf miners, aphids and even non-pathogenic root-colonising bacteria can induce systemic pathogen resistance, while some ISR traits can have a defensive effect against herbivores. ‘Cross-resistance’ elicited by and effective against non-microbial plant enemies thus might add significantly to the function of ISR. On the other hand, ‘trade-offs” have been reported, i.e. increased susceptibility to herbivores in ISR-expressing plants. Finally, ISR is a rather unspecific response, being active against different microbes. It thus might have effects on mutualistic bacteria and fungi, too. The question of how expression of ISR affects the large variety of mutualistic and antagonistic plant-microbe and plant-insect interactions cannot yet be answered. This knowledge is, however, needed to obtain a risk assessment for the use of chemically induced or genetically engineered ISR in crop protection. This review aims to provide an overview and to highlight some of the many open questions which require intensive ecological research.     

A rapid hypersensitive response associated with resistance in the willow Salix viminalis against the gall midge Dasineura marginemtorquens

Great genotypic variation in resistance against the gall midge Dasineura marginemtorquens Bremi (Diptera: Cecidomyiidae) exists within its willow host Salix viminalis L. (Salicaceae). In some resistant genotypes larvae die within 40 h after attempting to initiate galls. The present study tested the hypothesis that the hypersensitive response (HR) is involved in incompatible interactions between D. marginemtorquens and S. viminalis. By means of UV/blue light and visible light microscopy, we verified a rapidly (within 12 h after egg hatch) spreading cell death of an HR-type due to larval attack in resistant willow genotypes. Twelve h after egg hatch, the cell death had spread up to six cell layers in resistant S. viminalis genotypes whereas in susceptible genotypes only up to two cell layers were affected. In the groups of dead cells on the resistant genotypes accumulation of phenolics was observed within 24 h after egg hatch. The rapidity of the cell death induction, the early local accumulation of phenolic compounds, and the strong association of the cell death with larval mortality suggest that the formation of dead cells in resistant willow genotypes should be interpreted as a true HR.     

Impact of non‐pathogenic bacteria on insect disease resistance: importance of ecological context

1. The aerial surface of plants is a habitat for large and diverse microbial communities; termed the phyllosphere. These microbes are unavoidably consumed by herbivores, and while the entomopathogens are well studied, the impact of non‐pathogenic bacteria on herbivore life history is less clear. 2. Previous work has suggested that consumption of non‐entomopathogenic bacteria induces a costly immune response that might decrease the risk of infection. However, we hypothesised that insect herbivores should be selective in how they respond to commonly encountered non‐pathogenic bacteria on their host plants to avoid unnecessary and costly immune responses. 3. An ecologically realistic scenario was used in which we fed cabbage looper, Trichoplusia ni Hübner, larvae on cabbage or cucumber leaves treated with the common non‐entomopathogenic phyllosphere bacteria, Pseudomonas fluorescens and P. syringae. Their constitutive immunity and resistance to a pathogenic bacterium (Bacillus thuringiensis; Bt) and a baculovirus (T. ni single nucleopolyhedrovirus) were then examined. 4. While feeding on bacteria‐treated leaves reduced the growth rate and condition of T. ni, there was no effect on immunity (haemolymph antibacterial and phenoloxidase activities and haemocyte numbers). Phyllosphere bacteria weakly affected the resistance of T. ni to Bt but the direction of this effect was concentration dependent; resistance to the virus was unaffected. Host plant had an impact, with cucumber‐fed larvae being more susceptible to Bt. 5. The lack of evidence for a costly immune response to non‐entomopathogenic bacteria suggests that T. ni are probably adapted to consuming common phyllosphere bacteria, and highlights the importance of the evolutionary history of participants in multi‐trophic interactions.     

Responses and adaptation by Nephotettix virescens to monogenic and pyramided rice lines with Grh‐resistance genes

The green leafhopper, Nephotettix virescens (Distant) (Hemiptera: Cicadellidae), occasionally damages rice in Asia either directly, by feeding on the host phloem, or indirectly by transmitting tungro virus. We assessed the nature of resistance against the leafhopper in monogenic and pyramided near‐isogenic rice lines containing the resistance genes Grh2 and Grh4. Only the pyramided line was resistant to leafhopper damage. Leafhopper nymphs and adults had high mortality and low weight gain when feeding on the pyramided line and adults laid few eggs. In contrast, although there was some minor resistance in 45‐day‐old plants that possessed either Grh2 or Grh4 genes, the monogenic lines were generally as susceptible to the leafhopper as the recurrent parent line Taichung65 (T65). Resistance in the pyramided line was stable as the plant aged and under high nitrogen, and affected each of five Philippine leafhopper populations equally. Furthermore, in a selection study, leafhoppers failed to adapt fully to the pyramided resistant line: nymph and adult survival did improve during the first five generations of selection and attained similar levels as on T65, but egg‐laying failed to improve over 10 generations. Our preliminary results suggested that resistance was associated with physiological costs to the plants in some experiments. The results of this study demonstrate the success of pyramiding resistance genes through marker‐assisted breeding, to achieve a strong and potentially durable resistance. We discuss the utility of gene pyramiding and the development of near‐isogenic lines for leafhopper management.     

Faba bean forisomes can function in defence against generalist aphids

Phloem sieve elements have shut‐off mechanisms that prevent loss of nutrient‐rich phloem sap when the phloem is damaged. Some phloem proteins such as the proteins that form forisomes in legume sieve elements are one such mechanism and in response to damage, they instantly form occlusions that stop the flow of sap. It has long been hypothesized that one function of phloem proteins is defence against phloem sap‐feeding insects such as aphids. This study provides the first experimental evidence that aphid feeding can induce phloem protein occlusion and that the aphid‐induced occlusions inhibit phloem sap ingestion. The great majority of phloem penetrations in Vicia faba by the generalist aphids Myzus persicae and Macrosiphum euphorbiae triggered forisome occlusion and the aphids eventually withdrew their stylets without ingesting phloem sap. This contrasts starkly with a previous study on the legume‐specialist aphid, Acyrthosiphon pisum, where penetration of faba bean sieve elements did not trigger forisome occlusion and the aphids readily ingested phloem sap. Next, forisome occlusion was demonstrated to be the cause of failed phloem ingestion attempts by M. persicae: when occlusion was inhibited by the calcium channel blocker lanthanum, M. persicae readily ingested faba bean phloem sap.     

Reevaluating the conceptual framework for applied research on host‐plant resistance

Applied research on host‐plant resistance to arthropod pests has been guided over the past 60 years by a framework originally developed by Reginald Painter in his 1951 book, Insect Resistance in Crop Plants. Painter divided the “phenomena” of resistance into three “mechanisms,” nonpreference (later renamed antixenosis), antibiosis, and tolerance. The weaknesses of this framework are discussed. In particular, this trichotomous framework does not encompass all known mechanisms of resistance, and the antixenosis and antibiosis categories are ambiguous and inseparable in practice. These features have perhaps led to a simplistic approach to understanding arthropod resistance in crop plants. A dichotomous scheme is proposed as a replacement, with a major division between resistance (plant traits that limit injury to the plant) and tolerance (plant traits that reduce amount of yield loss per unit injury), and the resistance category subdivided into constitutive/inducible and direct/indirect subcategories. The most important benefits of adopting this dichotomous scheme are to more closely align the basic and applied literatures on plant resistance and to encourage a more mechanistic approach to studying plant resistance in crop plants. A more mechanistic approach will be needed to develop novel approaches for integrating plant resistance into pest management programs.     

Expression of Caenorhabditis elegans PCS in the AtPCS1‐deficient Arabidopsis thaliana cad1‐3 mutant separates the metal tolerance and non‐host resistance functions of phytochelatin synthases

Phytochelatin synthases (PCS) play key roles in plant metal tolerance. They synthesize small metal‐binding peptides, phytochelatins, under conditions of metal excess. Respective mutants are strongly cadmium and arsenic hypersensitive. However, their ubiquitous presence and constitutive expression had long suggested a more general function of PCS besides metal detoxification. Indeed, phytochelatin synthase1 from Arabidopsis thaliana (AtPCS1) was later implicated in non‐host resistance. The two different physiological functions may be attributable to the two distinct catalytic activities demonstrated for AtPCS1, that is the dipeptidyl transfer onto an acceptor molecule in phytochelatin synthesis, and the proteolytic deglycylation of glutathione conjugates. In order to test this hypothesis and to possibly separate the two biological roles, we expressed a phylogenetically distant PCS from Caenorhabditis elegans in an AtPCS1 mutant. We confirmed the involvement of AtPCS1 in non‐host resistance by showing that plants lacking the functional gene develop a strong cell death phenotype when inoculated with the potato pathogen Phytophthora infestans. Furthermore, we found that the C. elegans gene rescues phytochelatin synthesis and cadmium tolerance, but not the defect in non‐host resistance. This strongly suggests that the second enzymatic function of AtPCS1, which remains to be defined in detail, is underlying the plant immunity function.     

Mi‐mediated aphid resistance in tomato: tissue localization and impact on the feeding behavior of two potato aphid clones with differing levels of virulence

The Mi‐1.2 gene in tomato, Solanum lycopersicum L. (Solanaceae), confers resistance against several herbivores, including the potato aphid, Macrosiphum euphorbiae (Thomas) (Hemiptera: Sternorrhyncha: Aphididae) and the sweetpotato whitefly, Bemisia tabaci (Gennadius) (Hemiptera: Sternorrhyncha: Aleyrodidae). Previous studies on the tissue localization of resistance have given varying results; whitefly resistance was attributed to factors localized in the mesophyll or epidermis, whereas aphid resistance was attributed to factors localized in the phloem. Our study utilizes the direct current electrical penetration graph (DC‐EPG) technique to compare aphid feeding behavior on resistant (Mi‐1.2+) and susceptible (Mi‐1.2?) tomato plants. This study also compares the impact of resistance on the feeding behavior of two aphid clones that vary in their virulence, or their ability to survive and reproduce on resistant plants. Previous work had shown that the avirulent WU11 clone is almost completely inhibited by resistance, whereas the semi‐virulent WU12 clone can colonize resistant hosts. Here, DC‐EPG analysis shows that both aphid clones take longer to initiate cell sampling and to establish a confirmed sieve element phase on resistant plants than on susceptible hosts, and have shorter ingestion periods on resistant plants. However, the magnitude of these deterrent effects is far less for the semi‐virulent clone than for the avirulent aphids. In particular, the WU12 clone is less sensitive to factors that limit sieve element ingestion, showing shorter non‐probe duration and rapidly establishing sustained phloem ingestion on resistant plants when compared to the WU11 clone. We conclude that, in addition to previously described factors in the phloem that inhibit ingestion, Mi‐mediated aphid resistance also involves factors (possibly in the mesophyll and/or epidermis) that delay initiation of phloem salivation, and that act in the intercellular spaces to deter the first cell sampling. Furthermore, the relative effectiveness of these components of resistance differs among insect populations.     

Differences in parasite susceptibility and costs of resistance between naturally exposed and unexposed host populations

It is generally assumed that resistance to parasitism entails costs. Consequently, hosts evolving in the absence of parasites are predicted to invest less in costly resistance mechanisms than hosts consistently exposed to parasites. This prediction has, however, rarely been tested in natural populations. We studied the susceptibility of three naïve, three parasitized and one recently isolated Asellus aquaticus isopod populations to an acanthocephalan parasite. We found that parasitized populations, with the exception of the isopod population sympatric with the parasite strain used, were less susceptible to the parasite than the naïve populations. Exposed but uninfected (resistant) isopods from naïve populations, but not from parasitized populations, exhibited greater mortality than controls, implying that resistance entails survival costs primarily for naïve isopods. These results suggest that parasites can drive the evolution of host resistance in the wild, and that co‐existence with parasites may increase the cost‐effectiveness of defence mechanisms.     

Role of host‐plant selection in resistance of wild Solanum species to Macrosiphum euphorbiae and Myzus persicae

Damage to potatoes by Macrosiphum euphorbiae (Thomas) and Myzus persicae (Sulzer) (both Hemiptera: Aphididae) can be controlled through plant resistance. We used ethological experiments and electric penetration graph (EPG) analysis to evaluate the role of host selection in the previously assessed resistance levels of Solanum accessions: Solanum circaeifolium Bitter subsp. capsicibaccatum (Cárdenas) (PI210036), S. chomatophilum Bitter (PI243340 and PI310990), S. okadae Hawkes & Hjert. (PI458367), S. oplocense Hawkes (PI473368), S. pinnatisectum Dunal (PI186553), S. polyadenium Greenm. (PI230463), S. tarijense Hawkes (PI414150), and S. trifidum Correll (PI255538), to M. euphorbiae and M. persicae. Through multivariate analysis, we grouped behavioural variables into factors, which we related to host selection behaviours, and then evaluated whether factors varied between each accession and the susceptible S. tuberosum. None of the factors obtained by ethological experiments differed among accessions. Four of six and three of five factors obtained through EPG varied among accessions for M. euphorbiae and M. persicae, respectively, and were used to suggest resistance characteristics. The resistance to M. persicae of both S. chomatophilum accessions was associated with pathway activity disturbance. Solanum tarijense and S. polyadenium resistance to M. persicae resulted from leaf surface characteristics, which may be trichomes. Solanum oplocense and S. trifidum resistance to M. euphorbiae resulted from the wound response system, whereas S. pinnatisectum resistance may stem from nutritionally unbalanced or toxic phloem sap. Solanum polyadenium resistance to M. euphorbiae was phloem‐based. Solanum circaeifolium ssp. capsicibaccatum resistance to M. persicae, and the resistance of PI243340 S. chomatophilum and S. tarijense to M. euphorbiae were not related to host selection and therefore were presumably due to physiologically active compounds.     

The inhibitory effects of rose powdery mildew infection on the oviposition behaviour and performance of beet armyworms

In this study, we investigated whether the oviposition behaviour and performance of the beet armyworm, Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae), on the rose cultivar Rosa chinensis Jacq. (Rosaceae) were affected when the plants were infected by rose powdery mildew, Podosphaera pannosa (Wallr.: Fr.) de Bary (Erysiphales). The bioassays revealed that the moths significantly avoided ovipositing on mildew‐infected rose leaves when compared to healthy leaves. Pupal weights, emergence rates, and fecundity decreased when the caterpillars were fed mildewed rose leaves. Further laboratory bioassays aimed to elucidate the effects of two volatile headspace extracts (separately collected from healthy and mildewed rose plants) on the oviposition behaviour and performance of the moths. The moths clearly preferred to oviposit on healthy rose leaves that were not sprayed with additional volatiles rather than on healthy leaves sprayed with the volatile extracts from mildewed plants. The mean number of eggs laid on the former leaves was more than six times higher than that laid on the latter leaves. Olfactory bioassays demonstrated that ovipositing moths were significantly more attracted to volatiles emitted by healthy rose leaves than to those emitted by mildew‐infected leaves. Similar results were obtained when comparisons were made between the volatile extracts collected from healthy and mildewed rose plants. Thus, volatiles from mildew‐infected roses have a strong inhibitory effect against the moths. These results indicated that rose volatiles play a role in the oviposition behaviour of the moths, and that the volatiles induced by powdery mildew might be used for insect control.     

Leaf biomechanical properties as mechanisms of resistance to black cutworm (Agrotis ipsilon) among Poa species

Biomechanical properties can be important parameters in resistance of plants to herbivorous insects. As plants age, however, there can be dramatic changes in physical defenses that can then influence their susceptibility to insect herbivores. We measured changes in leaf biomechanical properties during ontogeny of Poa species and the relationship of these changes to the development of a generalist herbivore, the black cutworm, Agrotis ipsilon Hufnagel (Lepidoptera: Noctuidae), was investigated. Larvae were reared on two representative age classes, i.e., young (<60 days after planting) and old (>1 year after planting), of foliage in laboratory assays. Foliage generally reaches a peak fracture force between 80 and 109 days after planting depending on grass type. Foliage from old plants was significantly tougher than that of young plants, and black cutworm larvae reared on foliage from young plants gained significantly (ca. four times) more weight than those fed on foliage from old Poa plants. In addition, fracture force has a negative relationship with black cutworm development. Plant fiber, particularly neutral detergent fiber accounted for 65 and 46% of the variation in fracture force and larval development, respectively. These results provide additional insight into how plant ontogeny influences physical defenses to an insect herbivore in a grass system. Likewise, this is seemingly the first study to suggest a mechanism for host plant resistance to black cutworm. Plant fiber may be a useful trait to explore in plant improvement programs in which black cutworm is a primary pest (e.g., managed turfgrass).