From division to ‘divergence’: to understand wood growth across timescales, we need to (learn to) manipulate it

Wood formation is the Rosetta stone of tree physiology: a traceable, integrated record of physiological and morphological status. It also produces a large and persistent annual sink for terrestrial carbon, motivating predictive understanding. Xylogenesis studies have greatly expanded our knowledge of the intra-annual controls on wood formation, while dendroecology has quantified the environmental drivers of multi-annual variability. But these fields operate on different timescales, making it challenging to predict how short (e.g. turgor) and long timescale processes (e.g. disturbance) interactively influence wood formation. Toward this challenge, wood growth responses to natural climate events provide useful but incomplete explanations of tree growth variability. By contrast, direct manipulations of the tree vascular system have yielded unexpected insights, particularly outside of model species like boreal conifers, but they remain underutilized. To improve prediction of global wood formation, we argue for a new generation of experimental manipulations of wood growth across seasons, species, and ecosystems. Such manipulations should expand inference to diverse forests and capture inter- and intra-specific differences in wood growth. We summarize the endogenous and exogenous factors influencing wood formation to guide future experimental design and hypotheses. We highlight key opportunities for manipulative studies integrating measurements from xylogenesis, dendroanatomy, dendroecology, and ecophysiology.