Features shared by host-specific phytophagous insects and biotrophic plant pathogens include gene-for-gene interactions and the ability to induce susceptibility in plants. The Hessian fly shows both. To protect against Hessian fly, grasses have H genes. Avirulent larvae die on H-gene-containing resistant plants but the cause of death is not known. Imaging techniques were used to examine epidermal cells at larval attack sites, comparing four resistant wheat genotypes (H6, H9, H13, and H26) to a susceptible genotype. Present in both resistant and susceptible plants attacked by larvae were small holes in the tangential cell wall, with the size of the holes (0.1 microm in diameter) matching that of the larval mandible. Absent from attacked resistant plants were signs of induced susceptibility, including nutritive tissue and ruptured cell walls. Present in attacked resistant plants were signs of induced resistance, including cell death and fortification of the cell wall. Both presumably limit larval access to food, because the larva feeds on the leaf surface by sucking up liquids released from ruptured cells. Resistance was associated with several subcellular responses, including elaboration of the endoplasmic reticulum-Golgi complex and associated vesicles. Similar responses are observed in plant resistance to fungi, suggesting that "vesicle-associated penetration resistance" also functions against insects.

Resistance (R) genes have a proven record for protecting plants against biotic stress. A problem is parasite adaptation via Avirulence (Avr) mutations, which allows the parasite to colonize the R gene plant. Scientists hope to make R genes more durable by stacking them in a single cultivar. However, stacking assumes that R gene-mediated resistance has no fitness cost for the plant. We tested this assumption for wheat's resistance to Hessian fly, Mayetiola destructor (Say) (Diptera: Cecidomyiidae). Our study included ten plant fitness measures and four wheat genotypes, one susceptible, and three expressing either the H6, H9, or H13 resistance gene. Because R gene-mediated resistance has two components, we measured two types of costs: the cost of the constitutively-expressed H gene, which functions in plant surveillance, and the cost of the downstream induced responses, which were triggered by Hessian fly larvae rather than a chemical elicitor. For the constitutively expressed Hgene, some measures indicated costs, but a greater number of measures indicated benefits of simply expressing the H gene. For the induced resistance, instead of costs, resistant plants showed benefits of being attacked. Resistant plants were more likely to survive attack than susceptible plants, and surviving resistant plants produced higher yield and quality. We discuss why resistance to the Hessian fly has little or no cost and propose that tolerance is important, with compensatory growth occurring after H gene-mediated resistance kills the larva. We end with a caution: Given that plants were given good growing conditions, fitness costs may be found under conditions of greater biotic or abiotic stress.

Plant pathogen effectors encoded by Avirulence (Avr) genes benefit the pathogen by promoting colonization and benefit plants that have a matching resistance (R) gene by constituting a signal that triggers resistance. The Hessian fly, Mayetiola destructor (Say) (Diptera: Cecidomyiidae), resembles a plant pathogen in showing R/Avr interactions. Because of these interactions, a wheat plant with the H13 resistance gene can be resistant or susceptible depending on the genotype of the larva that attacks the plant, being resistant if attack comes from a larva with a functional vH13 gene, but susceptible if attack comes from a larva with a non‐functional vH13 gene. In this study we asked: does this susceptible interaction involving plants with H13 look like susceptible interactions with plants lacking H13? Possibly, the H13 plant attacked by a larva with a non‐functional vH13 is induced to partial rather than complete resistance. Or the larva, lacking its vH13‐encoded effector, is compromised in its ability to induce susceptibility, which includes forcing the plant to create a gall nutritive tissue. Responses of epidermal cells to larval attack were explored using imaging techniques (light microscopy, scanning and transmission electron microscopy). Whole‐organism responses were investigated by measuring the growth of plants and larvae. No evidence was found for partial resistance responses by H13 plants or for a compromise in the ability of vH13 loss‐of‐function larvae to induce susceptibility. It appears that disrupting vH13 function is sufficient for overcoming the induced resistance mediated by the H13 gene.

Before embarking on the 5-10 yr effort it can take to transfer plant resistance (R) genes to adapted crop cultivars, a question must be asked: is the pest a sufficient threat to warrant this effort? We used the recently discovered female-produced sex pheromone of the Hessian fly, Mayetiola destructor (Say) (Diptera: Cecidomyiidae),to explore this question for populations in the Upper Great Plains. Methods for pheromone trapping were established and trapping data were used to explore geographic distribution, phenology, and density. The pheromone lure remained attractive for up to 10 d and only attracted male Hessian flies. Traps placed within the crop canopy caught flies but traps placed above the crop canopy did not. Hessian flies were trapped throughout North Dakota starting in the spring and continuing through the summer and autumn. Densities were low in the spring but increased greatly during the early part of the summer, with peak adult emergence taking place at a time (July/August) when spring wheat was being harvested and winter wheat had not yet been planted. In the autumn, adults were found at a time when winter wheat seedlings are growing. The discovery of flies on Conservation Reserve Program land supports the idea that pasture grasses serve as alternate hosts. We conclude that the Hessian fly is a risk to wheat in the Upper Great Plains and predict that global warming and the increasing cultivation of winter wheat will add to this risk.

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