Seasonal variation in the content of hydrolysable tannins in leaves of Betula pubescens

The contents of 13 hydrolysable tannins in the leaves of white birch (Betula pubescens L.) trees were analysed at twelve stages throughout the growing season. All individual galloylglucoses, from 1-O-galloyl-beta-D-glucopyranose to 1,2,3,4,6-penta-O-galloyl-beta-D-glucopyranose, accumulated in young leaves, while ellagitannins showed significantly variable seasonal trends. The major ellagitannin during the whole growing season was pedunculagin while 2,3-(S)-HHDP-glucopyranose. the end product of the proposed ellagitannin pathway, accumulated in mature leaves. Relationships between the characteristics of seasonal variation in the contents of individual ellagitannins and their chemical structures were used to unravel the biogenesis of ellagitannins in birch leaves. Evidence of degradation of ellagitannins through hydrolysis during leaf growth and development is presented and implications for herbivory are discussed.