title: Delayed induced changes in the biochemical composition of host plant leaves during an insect outbreak.
abstract: In birch, Betula pubescens, herbivore-induced delayed induced resistance (DIR) of defoliated trees may cause a strong reduction in the potential fecundity of a geometrid folivore Epirrita autumnata. In this study, we examined the biochemical basis of DIR in birch leaves during a natural outbreak of E. autumnata. A set of experimental trees was defoliated at four sites by wild larvae in the peak year of the outbreak, whereas control trees were protected from defoliation by spraying with an insecticide. The biochemical composition of leaves was analysed in the following year and, although the DIR response was weak during this outbreak, causing less than a 20% reduction in the potential fecundity of E. autumnata, some consistent relationships between defoliation, biochemistry and pupal mass of E. autumnata suggested a general biochemical basis for the defoliation-induced responses in birch leaves. Total concentrations of nitrogen, sugars and acetone-insoluble residue (e.g. cell wall polysaccharides, cell-wall-bound phenolics, protein, starch, lignin and hemicellulose) were consistently lower, and total concentrations of phenolics, especially of gallotannins and soluble proanthocyanidins, were higher in the leaves of trees defoliated in the previous year than in those protected from defoliation. The capacity of tannins to precipitate proteins correlated with contents of gallotannins, and was highest in defoliated trees. The pupal mass of E. autumnata showed a strong, positive correlation with concentrations of nitrogen and sugars, and a negative correlation with the acetone-insoluble residue and gallotannins in foliage. Correlations with other measured biochemical traits were weak. The correlation coefficients between biochemical traits and pupal mass consistently had similar signs for both defoliated and insecticide-sprayed trees, suggesting that variation in leaf quality due to defoliation in the previous year was based on similar biochemical traits as variation for other reasons. We suggest that DIR is associated with reduced growth activity of leaves, and may be seen as a delay in the biochemical maturation of leaves in defoliated trees. This explains the high concentration of gallotannins in defoliated trees, a characteristic feature of young leaves. However, the lower content of nitrogen and the higher content of soluble proanthocyanidins in defoliated trees are traits usually characterising mature, not young, leaves, indicating defoliation-induced changes in chemistry in addition to modified leaf age. Our results emphasise the importance of understanding the natural changes in chemistry during leaf maturation when interpreting defoliation-induced changes in leaf biochemistry.
authors.affiliation: Section of Ecology, Department of Biology, and Kevo Subarctic Research Institute, University of Turku, FIN-20014 Turku, Finland e-mail: Pekka.Kaitaniemi@utu.fi; Fax: +358 2 333 6550, , , , , , FI.
authors.forename: Pekka
authors.initials: P
authors.lastname: Kaitaniemi
authors.affiliation: Section of Ecology, Department of Biology, and Kevo Subarctic Research Institute, University of Turku, FIN-20014 Turku, Finland e-mail: Pekka.Kaitaniemi@utu.fi; Fax: +358 2 333 6550, , , , , , FI.
authors.forename: Kai
authors.initials: K
authors.lastname: Ruohomäki
authors.affiliation: Section of Ecology, Department of Biology, and Kevo Subarctic Research Institute, University of Turku, FIN-20014 Turku, Finland e-mail: Pekka.Kaitaniemi@utu.fi; Fax: +358 2 333 6550, , , , , , FI.
authors.forename: Vladimir
authors.initials: V
authors.lastname: Ossipov
authors.affiliation: Section of Ecology, Department of Biology, and Kevo Subarctic Research Institute, University of Turku, FIN-20014 Turku, Finland e-mail: Pekka.Kaitaniemi@utu.fi; Fax: +358 2 333 6550, , , , , , FI.
authors.forename: Erkki
authors.initials: E
authors.lastname: Haukioja
authors.affiliation: Laboratory of Physical Chemistry, Department of Chemistry, FIN-20014 Turku, Finland, , , , , , FI.
authors.forename: Kalevi
authors.initials: K
authors.lastname: Pihlaja
keywords: Gallotannins; Induced resistance; Key wordsBetula pubescens; Leaf biochemistry; Phenology