Biomechanics of the columnar cactus Pachycereus pringlei

Abstract

We report the longitudinal variations in stiffness and bulk density of tissue samples drawn from along the length of two Pachycereus pringlei plants measuring 3.69 and 5.9 m in height to determine how different tissues contribute to the mechanical stability of these massive vertical organs. Each of the two stems was cut into segments of uniform length and subsequently dissected to obtain and mechanically test portions of xylem strands, stem ribs, and a limited number of pith and cortex samples. In each case, morphometric measurements were taken to determine the geometric contribution each tissue likely made to the ability of whole stems to resist bending forces. The stiffness of each xylem strand increased basipetally toward the base of each plant where stiffness sharply decreased, reaching a magnitude comparable to that of strands 1 m beneath the stem apex. The xylem was anisotropic in behavior, i.e., its stiffness measured in the radial and in the tangential directions differed significantly. Despite the abrupt decrease in xylem strand stiffness at the stem base, the contribution made by this tissue to resist bending forces increased exponentially from the tip to the base of each plant due to the accumulation of wood. A basipetal increase in the stiffness of the pith (and, to limited extent, that of the cortex) was also observed. In contrast, the stiffness of stem rib tissues varied little as a function of stem length. These tissues were stiffer than the xylem in the corresponding portions of the stem along the upper two-fifths of the length of either plant. Tissue stiffness and bulk density were not significantly correlated within or across tissue types. However, a weak inverse relationship was observed for these properties in the case of the xylem and stem rib tissues. We present a simple formula that predicts when stem ribs rather than the xylem strands serve as the principal stiffening agents in stems. This formula successfully predicted the observed aspect ratio of the stem ribs (the average quotient of the radial and tangential dimensions of rib transections), and thus provided circumstantial evidence that the ribs are important for mechanical stability for the distal and younger regions of the stems examined.