Coping with multiple enemies: an integration of molecular and ecological perspectives.

How plants respond to attack by the range of herbivores and pathogens that confront them in the field is the subject of considerable research by both molecular biologists and ecologists. However, in spite of the shared focus of these two bodies of research, there has been little integration between them. We consider the scope for such integration, and how greater dialogue between molecular biologists and ecologists could advance understanding of plant responses to multiple enemies.

Beetle grazing reduces natural infection of Rumex obtusifolius by fungal pathogens.

In previously reported laboratory experiments, infection of Rumex obtusifolius by the rust fungus Uromyces rumicis was decreased on leaves which had prior herbivory by the beetle Gastrophysa viridula. In this paper we investigate whether this interaction is robust for natural infection by a variety of fungi in field experiments carried out in spring and autumn with plants given different levels of nitrogen fertilization. Grazing by G. viridula led to a decrease in lesion density of Ramularia rubella and Venturia rumicis in the spring and V. rumicis and U. rumicis in the autumn experiment. For V. rumicis and U. rumicis significant reductions in lesion density occurred on the undamaged leaves of damaged plants, compared with similar leaves on undamaged plants, suggesting systemic induced resistance. This induced resistance was usually independent of the amount of nitrogen fertilization, although the inhibitory effect of grazing on R. rubella in the spring and V. rumicis in the autumn experiment was enhanced by increasing nitrogen fertilization and was inhibited by increasing nitrogen fertilization for V. rumicis in the spring. In both experiments, the lesion density of V. rumicis was greater on leaves on which R. rubella was also present, and the presence of U. rumicis in the autumn experiment was linked to a similar but greater effect on V. rumicis lesion density. We found no evidence of induced resistance by fungi against fungi in these experiments. We highlight the complex interactions between inhibitory and facilitatory processes acting on leaf fungal infection. These results are compared with the proposed molecular mechanisms of induced resistance(s) and we consider the benefits of closer integration between molecular and ecological investigations of induced resistances that occur in the field.

The effect of an insect herbivore and a rust fungus individually, and combined in sequence, on the growth of two Rumex species.

The chrysomelid beetle Gastrophysa viridula and the rust fungus Urmnyces rumicis both occur on leaves of Rumex crispus and R. obtusifolius. We investigated the effect of beetle grazing or rust infection individually and when combined in sequence on the growth of their hosts in the field. Singly, beetle or rust reduced leaf area and plant biomass; the effect was greater on R. crispus, and rust caused greater damage than the beetle. Beetle grazing with subsequent rust infection caused damage no greater than that caused by rust alone, although on R. obtusifolius damage was greater than that from beetle grazing alone. Rust infection of R. obtusifolius with subsequent beetle grazing produced damage similar to that from other treatments; involving rust infection. In R. crispus this treatment produced the greatest reduction in biomass, The reductions in root and total plant weight from rust infection with subsequent beetle grazing were accurately predicted by a model including the damage produced by beetle and rust alone and the length of time each was present on the plant. This model also predicted accurately the damage to R. obtusifolius from the beetle followed by rust treatment, but over-estimated by up to 40% the damage to R. crispus. This can be explained mainly by an inhibition of rust infection by beetle grazing.