Sperry’s packing rule affects the spatial proximity but not clustering of xylem conduits: the case of Fagus sylvatica L.

Sperry’s packing rule predicts the optimum packing of xylem conduits in woody plants, where the frequency of xylem conduits varies approximately inversely with the square of the conduit radius. However, it is well established that such anatomical disposition does not remain fixed but is subject to a suite of adaptations induced by physiological constraints driven by both ontogenetic development and environmental characteristics. Here we challenge the hypothesis that increasing frequency of xylem conduits, concomitant with the decrease in their lumen area along the xylem pathway, would affect the spatial distribution of vessels inside tree-rings and their aggregation. To this end, we measured the vessels’ anatomical characteristics inside each tree-ring along with a complete radial series taken at different stem heights of Fagus sylvatica L. trees. Point pattern analysis indicated a significant effect of the distance from the tree base and a weak effect of cambial age on the nearest neighbour distance among xylem vessels, suggesting that vessels were closer to each other near the apex, and became progressively more distant toward the base. The spatial pattern of xylem vessels violated the assumption of complete spatial randomness, vessel spatial arrangement followed a uniform distribution at different distances from the tree base. Although there was an increase in the intensity and proximity among vessels, we demonstrated that no patterns of aggregation between vessels were found in sampled F. sylvatica trees. Rather, point pattern profiles clearly highlighted a lack of aggregation of vessels in the face of a regular spatial distribution in the annual growth rings along the stems.