Do tall tree species have higher relative stiffness than shorter species?

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.

Toward an improved model of maple sap exudation: the location and role of osmotic barriers in sugar maple, butternut and white birch.

Two theories have been proposed to explain how high positive pressures are developed in sugar maple stems when temperatures fluctuate around freezing. The Milburn-O'Malley theory proposes that pressure development is purely physical and does not require living cells or sucrose. The osmotic theory invokes the involvement of living cells and sucrose to generate an osmotic pressure difference between fibers and vessels, which are assumed to be separated by an osmotic barrier. We analyzed wood of Acer saccharum Marsh., Juglans cinerea L. and Betula papyrifera Marsh. (all generate positive pressures) examining three critical components of the osmotic model: pits in cell walls, selectivity of the osmotic barrier and stability of air bubbles under positive xylem pressure. We examined the distribution and type of pits directly by light and scanning electron microscopy (SEM), and indirectly by perfusion of branch segments with fluorescent dyes with molecular masses similar to sucrose. The latter approach allowed us to use osmotic surrogates for sucrose that could be tracked by epifluorescence. Infusion experiments were used to assess the compartmentalization of sucrose and to determine the behavior of gas bubbles as predicted by Fick's and Henry's laws. The SEM images of sugar maple revealed a lack of pitting between fibers and vessels but connections between fiber-tracheids and vessels were present. Fluorescein-perfusion experiments demonstrated that large molecules do not diffuse into libriform fibers but are confined within the domain of vessels, parenchyma and fiber-tracheids. Results of the infusion experiments were in agreement with those of the fluorescein perfusions and further indicated the necessity of a compartmentalized osmolyte to drive stem pressure, as well as the inability of air bubbles to maintain such pressure because of instability. These results support the osmotic model and demonstrate that the secondary cell wall is an effective osmotic barrier for molecules larger than 300 g mol(-1).

Populus species from diverse habitats maintain high night-time conductance under drought.

We investigated the interspecific variability in nocturnal whole-plant stomatal conductance under well-watered and drought conditions in seedlings of four species of Populus from habitats characterized by abundant water supply (mesic and riparian) or from drier upland sites. The study was carried out to determine whether (i) nocturnal conductance varies across different species of Populus according to their natural habitat, (ii) nocturnal conductance is affected by water stress similarly to daytime conductance based on species habitat and (iii) differences in conductance among species could be explained partly by differences in stomatal traits. We measured whole-plant transpiration and conductance (G) of greenhouse-grown seedlings using an automated high-resolution gravimetric technique. No relationship was found between habitat preference and daytime G (GD), but night-time G (GN) was on average 1.5 times higher in riparian and mesic species (P. deltoides Bartr. ex Marsh. and P. trichocarpa Torr. & Gray) than in those from drier environments (P. tremuloides Michx. and P. × petrowskyana Schr.). GN was not significantly reduced under drought in riparian species. Upland species restricted GN significantly in response to drought, but it was still at least one order of magnitude greater that the cuticular conductance until leaf death was imminent. Under both well-watered and drought conditions, GN declined with increasing vapour pressure deficit (D). Also, a small increase in GN towards the end of the night period was observed in P. deltoides and P. × petrowskyana, suggesting the involvement of endogenous regulation. The anatomical analyses indicated a positive correlation between G and variable stomatal pore index among species and revealed that stomata are not likely to be leaky but instead seem capable of complete occlusion, which raises the question of the possible physiological role of the significant GN observed under drought. Further comparisons among closely related species that occupy ecologically diverse habitats may provide a better understanding of the genetic versus environmental regulations of nocturnal water loss.