Crown ratio influences allometric scaling in trees

Allometric theories suggest that the size and shape of organisms follow universal rules, with a tendency toward quarter-power scaling. In woody plants, however, structure is influenced by branch death and shedding, which leads to decreasing crown ratios, accumulation of heartwood, and stem and branch tapering. This paper examines the impacts on allometric scaling of these aspects, which so far have been largely ignored in the scaling theory. Tree structure is described in terms of active and disused pipes arranged as an infinite branching network in the crown, and as a tapering bundle of pipes below the crown. Importantly, crown ratio is allowed to vary independently of crown size, the size of the trunk relative to the crown deriving from empirical results that relate crown base diameter to breast height diameter through crown ratio. The model implies a scaling relationship in the crown which reduces to quarter-power scaling under restrictive assumptions but would generally yield a scaling exponent somewhat less than three-quarters. For the whole tree, the model predicts that scaling between woody mass and foliage depends on crown ratio. Measurements on three boreal tree species are consistent with the model predictions.