Functional compartmentalisation of nutrients and phenolics in the tissues of galls induced by Leptocybe invasa (Hymenoptera: Eulophidae) on Eucalyptus camaldulensis (Myrtaceae)

Galling herbivores induce structural and chemical alterations in their host plant's tissues. These insects have been the focus of little study in the case of Australian taxa. Leptocybe invasa, a native Australian galling hymenopteran associated with Eucalyptus species, causes economic damage to plantation eucalypts in many countries around the world. Leptocybe invasa oviposits in the midribs and petioles of expanding leaves thereby intercepting photosynthates and impairing normal expansion. We analysed the ultrastructural and chemical cellular changes in L. invasa galls on Eucalyptus camaldulensis (probably subspecies camaldulensis) to diagnose how the insect manipulates plant cells and tissues and the significance of these alterations for insect nutrition and protection. Galling stimuli induce the formation of two functionally compartmentalised types of tissue. Phenolic plant secondary metabolites and anthocyanins (plant pigments) accumulate in the outer compartment, while primary metabolites accumulate in the inner compartment. The nutritive cells (inner compartment) accumulate protein and lipids that provide food for the larvae. Total polyphenol concentrations did not differ significantly between outer and inner compartments. Nevertheless, the concentrations of quercetin and kaempferol derivatives were higher in the outer compartment than in the inner compartment. These differences could be related to the protection of plant tissues against ultraviolet rays and the maintenance of redox homeostasis. There were higher ratios of hexahydroxydiphenoyl-containing hydrolysable tannins rather than galloyl-containing hydrolysable tannins in the inner compartment. This shift in the oxidative capacity of the polyphenols in the inner compartment could represent a defensive plant response to the larvae.