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Background and Aims Heartwood plays an important role in maintaining the structural integrity of trees. Although its formation has long been thought to be driven solely by internal ageing processes, more recent hypotheses suggest that heartwood formation acts as a regulator of the tree water balance by modulating the quantity of sapwood. Testing both hypotheses would shed light on the potential ecophysiological nature of heartwood formation, a very common process in trees. Methods We measured quantities of heartwood and sapwood, xylem conduits and the width and number of growth rings on 406 stems of Pericopsis elata with ages ranging from 2 to 237 years. A subset of 17 trees with similar ages but varying growth rate were sampled in a shaded (slower-growth) site and a sun-exposed (faster-growth) site. We used regression analysis and structural equation modelling to investigate the dynamics and drivers of heartwood formation. Key Results We found a positive effect of growth rate on the probability of heartwood occurrence, suggesting an earlier heartwood onset in faster-growing stems. After this onset age, heartwood area increased with stem diameter and age. Despite the similar heartwood production per unit stem diameter increment, shaded trees produced heartwood faster than sun-exposed trees. Tree age and hydraulics showed similar direct effects on heartwood and sapwood area of sun-exposed trees, suggesting their mutual role in driving the heartwood dynamics of sun-exposed trees. However, for shaded trees, only tree hydraulics showed a direct effect, suggesting its prominent role over age in driving the heartwood dynamics in limited growing conditions. The positive relationship between growth rate and maximum stomatal conductance supported this conclusion. Conclusions Heartwood area increases as the tree ages, but at a slower rate in trees where water demand is balanced by a sufficient water supply. Our findings suggest that heartwood formation is not only a structural process but also functional. [ABSTRACT FROM AUTHOR]

The transition from the living water-transporting sapwood to heartwood involves in many tree species impregnation with extractives. These differ in amount and composition, and enhance resistance against bacteria, insects or fungi. To understand the synthesis, transport and impregnation processes new insights into the biochemical processes are needed by in-situ methods. Here we show the extractive distribution in pine (Pinus sylvestris) microsections with a high lateral resolution sampled in a non-destructive manner using Confocal Raman Microscopy. Integrating marker bands of stilbenes and lipids enables to clearly track the rapid change from sapwood to heartwood within one tree ring. The higher impregnation of the cell corner, compound middle lamella, the S3 layer and pits reveals the optimization of decay resistance on the micron-level. Furthermore, deposits with changing chemical composition are elucidated in the rays and lumen of the tracheids. The spectral signature of these deposits shows the co-location of lipids and pinosylvins with changing ratios from the living to the dead tissue. The results demonstrate that the extractive impregnation on the micro- and nano-level is optimized by a symbiotic relationship of lipids and pinosylvins to enhance the tree's resistance and lifetime. [ABSTRACT FROM AUTHOR]