Richard Jagels1, Maria A Equiza2, Douglas A Maguire3, Damian Cirelli4. 1. University of Maine, School of Forest Resources, Orono, Maine, 04469-5755, USA. 2. University of Alberta, Department of Renewable Resources, Edmonton, Alberta, T6G2R3, Canada. 3. Oregon State University, Department of Forest Engineering, Resources, and Management, Corvallis, Oregon, 97331, USA. 4. University of Alberta, Department of Biological Sciences, Edmonton, Alberta, T6G2R3, Canada.
Abstract
PREMISE OF THE STUDY: In 1757 Leonhard Euler demonstrated that to avoid bending tall columns needed to be stiffer but not stronger than shorter columns of equal diameter and material density. Many researchers have concluded that trees have a fixed stiffness to basic density ratio, and therefore, trees adjust for increasing height by adding mass to adjust stem form. But the wood science literature points to considerable variance in stiffness with respect to green wood density. METHODS: Using the vast global repository of green wood mechanical properties, we compared relative stiffness and relative strength between taller and shorter species. For North American trees, we examined stem moisture distribution. KEY RESULTS: For all regions of the world, taller species on average possessed greater stiffness, but not strength, than shorter species of equal basic specific gravity. We looked for a possible universal mechanism that might allow taller tree species to adjust stiffness without affecting xylem specific gravity and concluded that the evidence points to a decrease in cellulose microfibril angle in structural cell walls combined with possible increases in holocellulose percentage. The evidence is strongest for conifers. We also showed that tall conifers have the ability to adjust the distribution of xylem moisture to maximize conduction while minimizing column load. CONCLUSIONS: Our research reveals that taller trees have developed internal stem adjustments to minimize diameter increase while attaining ever-greater heights, thus enabling these taller species to reduce energy expended on biomass accumulation while gaining greater access to solar radiation.
PREMISE OF THE STUDY: In 1757 Leonhard Euler demonstrated that to avoid bending tall columns needed to be stiffer but not stronger than shorter columns of equal diameter and material density. Many researchers have concluded that trees have a fixed stiffness to basic density ratio, and therefore, trees adjust for increasing height by adding mass to adjust stem form. But the wood science literature points to considerable variance in stiffness with respect to green wood density. METHODS: Using the vast global repository of green wood mechanical properties, we compared relative stiffness and relative strength between taller and shorter species. For North American trees, we examined stem moisture distribution. KEY RESULTS: For all regions of the world, taller species on average possessed greater stiffness, but not strength, than shorter species of equal basic specific gravity. We looked for a possible universal mechanism that might allow taller tree species to adjust stiffness without affecting xylem specific gravity and concluded that the evidence points to a decrease in cellulose microfibril angle in structural cell walls combined with possible increases in holocellulose percentage. The evidence is strongest for conifers. We also showed that tall conifers have the ability to adjust the distribution of xylem moisture to maximize conduction while minimizing column load. CONCLUSIONS: Our research reveals that taller trees have developed internal stem adjustments to minimize diameter increase while attaining ever-greater heights, thus enabling these taller species to reduce energy expended on biomass accumulation while gaining greater access to solar radiation.
Keywords:
global forests; holocellulose; materials science; mechanical properties of wood; microfibril angle; modulus of elasticity; modulus of rupture; moisture load in trees; specific gravity; wood anatomy