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Red spruce (Picea rubens Sarg.) and American beech (Fagus grandifolia Ehrh.) nutritional imbalances observed during 1998–2000 in response to nitrogen additions beginning in 1989 at Bear Brook Watershed in Maine, USA, were reversed by 2013. However, nitrogen-containing metabolites continued to accumulate to detoxify ammonia. While sugar maple (Acer saccharum Marsh.) was N-limited and benefitted from N+S additions, spruce and birch established new homeostatic status via adjusting cellular metabolism.

Dynamic species-specific metabolic changes in the trees exposed to chronic N+S additions at the Bear Brook Watershed in Maine, USA