SAFETY FACTORS IN VERTICAL STEMS: EVIDENCE FROM EQUISETUM HYEMALE

Predictions from a mechanical model for hollow vertical stems are tested against morphometric and mechanical studies of the vertical stems of Equisetum hyemale. The model predicts 1) that the wall thickness of hollow internodes must be at least 15% of the external radius of shoots, 2) that the elastic modulus of stems is quantitatively related to the ratio of apoplast (cell walls) to symplast (cytoplasm) areas in transverse sections through stems, and that (3) hollow stems are designed to sustain an additional and significant proportion of their own weight. The "safety factors" predicted for a hollow vertical stem are used to examine two adaptationist explanations for hollow stems: 1) "economy in design," which argues that natural selection will favor a reduction in the metabolic cost in constructing an organ, and 2) "mechanical design," which argues that stems are designed to maximize their mechanical stability during vertical growth. Evidence from E. hyemale indicates that 1) there is a developmental limit to the maximum allotment of biomass invested in the construction of stems, 2) as stem height increases, morphometric adjustments in internodal wall thickness occur which converge on predicted safety limits, and 3) the elastic modulus of stems changes as a function of the ratio of apoplast to symplast areas seen in transverse sections through shoots. Biomechanical and developmental evidence and the allometry of E. hyemale stems are consistent with the view that stems are designed for safety and are inconsistent with some predictions based on the economy in design.