Irregular Heartwood Formation in Maritime Pine (Pinus pinaster Ait):Consequences for Biomechanical and Hydraulic Tree Functioning

As the proportion of sapwood (SW) transformed into heartwood (HW) is irregular both radially and longitudinally in trunks of Maritime pine ( Pinus pinaster Ait.), it has been suggested that HW formation is a developmental process, regulated internally within the tree. In trees where stem growth is eccentric due to stem lean or wind action, the number of annual growth rings of SW transformed into HW is greater on the compressed side of the tree. To determine the contribution to bending stiffness, if any, of this prematurely formed HW, four point bending tests were carried out on fresh HW and SW samples taken from the same growth ring, or neighbouring growth rings, at different cross-sectional positions at a height of 2 m from six 52-year-old Maritime pines. The mean (±s.e.) modulus of elasticity ( E ) was 7.6 ± 0.3 GPa (longitudinal direction) for all samples. No significant differences in E were found between HW and SW; thus HW does not play a significant mechanical role in bending stiffness. To test a second hypothesis that early HW formation on the compressed side of trees may maintain a constant, optimal volume of SW around the tree, the Pipe Model Theory was applied to 12 52-year-old leaning Maritime pines (angle of lean varied from 0–22°). The surface area ( S ) of the SW was determined at different heights up the trunk and correlated with crown surface area ( S crown ). Regressions between S SW and S crown were highly significant, thereby supporting the theory that HW formation and extension is controlled internally in Maritime pine. HW formation in Maritime pine then serves to maintain an optimal proportion of functional SW which is an important criterion for survival in a species often subjected to severe drought for long periods. 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