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Summary In trunks of Juglans nigra and the hybrid J. major × J. regia, the presence of non-structural carbohydrates, sucrose synthesizing and degrading enzymes, and their correlation with heartwood formation was investigated. Contents of starch and sucrose were highest in the youngest sapwood, decreased with increasing age of the tissue, and were absent in the heartwood. Pools of the monosaccharides glucose and fructose were low in the sapwood, and fructose was absent from the heartwood. Glucose transiently increased at the sapwood heartwood boundary in trunks of the hybrid. In black walnut stems, however, glucose started to accumulate within the transition zone and reached considerable amounts in the heartwood. Cold-adaptation in walnut wood was characterized by accumulation of soluble sugars. Sucrose formation was enabled by enhanced rates of sucrose-phosphate synthase (SPS, EC 2.4.1.14). Mid-winter starch-sugar interconversion was accompanied by increases in the activity of sucrose synthase (SuSy, EC 2.4.1.13; black walnut), or acid invertases (EC 3.2.1.26; hybrid). In the tissues undergoing heartwood formation, sucrose breakdown was enhanced from late summer until early winter. Sucrolysis was dominated by acid invertases with minor contribution of sucrose synthase. The catalytic activity of UDP-glucose pyrophosphorylase (EC 2.7.7.9), involved in the metabolization of the sucrose cleavage products, followed this seasonal trend and showed elevated activities from late summer until early winter. These data are further proof for the earlier made hypothesis (Hauch and Magel 1998) that the in situ synthesis of heartwood flavonoids relies on an interaction between primary (sucrose) and secondary metabolism. Flavonoids, however, constitute only a minor fraction in the heartwood of walnut and the bulk of heartwood phenolics seem to derive from transformation of phenolic precursors. Therefore, these recent findings together with earlier data are taken as evidence that more than one type of heartwood formation exists. |
