Although lipid autoxidation in the boiling kettle is a key determinant of the cardboard flavor of aged beers, recent results show that mashing is another significant source of wort nonenal potential, the well-known indicator of how a beer will release (E)-2-nonenal during storage. Although unstable, deuterated (E)-2-nonenal nitrogen adducts created during mashing can in some cases partially persist in the pitching wort, to release deuterated (E)-2-nonenal during beer aging. In the experiment described here, the relative contributions of mashing and boiling were estimated at 30 and 70%, respectively. The presence of oxygen during mashing and, to a lesser extent, high lipoxygenase activity can intensify the stale cardboard flavor.
Liégeois, C., Meurens, N., Badot, C., & Collin, S. (2002). Release of deuterated (E)-2-nonenal during beer aging from labeled precursors synthesized before boiling. Journal of agricultural and food chemistry, 50(26), 7634-7638. https://doi.org/10.1021/jf020617v (Original work published 2002)