Xylem Architecture in Giant Cacti Stems Follows Universal Scaling Theory

Xylem conduits, a primary feature of most terrestrial plant taxa, deliver water to photosynthetic tissues and play a critical role in plant water relations and drought tolerance. Non-succulent woody taxa generally follow a universal rate of tip-to-base conduit widening such that hydraulic resistance remains constant throughout the plant stem. Giant cacti inhabit arid regions throughout the Americas and thrive in water-limited environments by complimenting water-storing succulent tissues with resource-efficient Crassulacean Acid Metabolism. Considering these adaptations, the objectives of this study were threefold: 1) determine whether xylem conduits in columnar cacti follow universal scaling theory as observed in woody taxa; 2) evaluate whether xylem hydraulic diameter is inversely correlated with xylem vessel density; and 3) determine whether xylem double-wall thickness-to-span ratio and other hydraulic architectural traits are convergent among phylogenetically diverse cactus species. This thesis investigates the xylem anatomy of nine cactus species native to the Sonoran Desert of Arizona and Mexico, the tropical dry forests of southern Mexico, and the Alto Plano region of Argentina. Soft xylem tissues closest to the stem apex underwent a modified polyethylene glycol treatment to stabilize for sectioning with a sledge microtome. Across all species: hydraulic diameter followed a basipetal widening rate of 0.21 (p < 0.001), closely matching the universal rate of 0.20 for woody taxa; and xylem vessel density was inversely correlated with both length from stem apex (p < 0.001) and hydraulic diameter (p < 0.001). Double-wall thickness-to-span ratio had little to no significant correlation with either length from stem apex or hydraulic diameter. There was no significant difference in hydraulic architectural trait patterns between phylogenetically diverse species with various stem morphologies, nor was there a significant correlation between conduit widening rates and volume-to-surface-area ratios. This study demonstrates that giant cacti follow similar internal anatomical constraints as non-succulent woody taxa, yet stem succulence and water storage behavior in cacti remain separate from internal hydraulic architecture, allowing cacti to thrive in arid environments. Understanding how cacti cope with severe water limitations provides new insights on evolutionary constraints of stem succulents as they functionally diverged from other life forms.