Seasonal changes in birch leaf chemistry: are there trade-offs between leaf growth and accumulation of phenolics?

Several plant-herbivore hypotheses are based on the assumption that plants cannot simultaneously allocate resources to growth and defence. We studied seasonal patterns in allocation to growth and putatively defensive compounds by monitoring several chemical and physical traits in the leaves of mountain birch from early June (budburst) to late September (leaf senescence). We found significant seasonal changes in all measured characteristics, both in terms of concentrations (mg g-1) and amounts (mg leaf-1). Changes were very rapid in the spring, slow in the middle of the season, and there was another period of fast changes in the senescing leaves. Co-occurring changes in physical leaf traits and concentrations of several compounds indicated a seasonal decline in foliage suitability for herbivores. Concentrations of protein and free amino acids declined through the growing season whereas individual sugars showed variable seasonal patterns. The seasonal trends of phenolic groups differed drastically: concentrations of soluble proanthocyanidins increased through the season, whereas cell wall-bound proanthocyanidins, gallotannins and flavonoid glycosides declined after an initial increase in young leaves. We failed to find proof that the seasonal accumulation of phenolics would have been seriously compromised by leaf or shoot growth, as assumed by the growth/differentiation balance hypothesis and the protein competition model hypothesis. On the contrary, there was a steady increase in the total amount of phenolics per leaf even during the most active leaf growth.