Role of Extractable and Non-Extractable Polyphenols in the Formation of Beech (Fagus sylvatica L.) Red Heartwood Chromophores

Despite the long history of beech (Fagus sylvatica L.) red heartwood research, there has been no experimental proof on the structure of the chromophores yet. For the first time, using high-performance liquid chromatography/diode array detection/multistage electrospray ionization mass spectrometry, it was evidenced that red heartwood chromophores are water/methanol solvent extractable high molecular weight (400–2200 Da) compounds, which are polymerized, transformed, and oxidized products of (epi)catechin and taxifolin. Acid soluble non-extractable polyphenols (flavonoids, tannins) were not evidenced in the cell wall structure, while alkaline soluble compounds (ferulic acid, dehydrodiferulic acid, p-coumaric acid) have been identified for the first time from the sapwood/red heartwood boundary tissues: these supposedly play a role in the structural reinforcement of the cell wall structure and in the antioxidant protection and have a lesser role in color formation. Results on the structure of chromophores and on cell wall composition may enhance color homogenization technologies and contribute to a better utilization of red-heartwooded timber in the future.