Associations between leaf structure, orientation, and sunlight exposure in five Western Australian communities

Five plant communities in Western Australia, as well as selected desert and Rocky Mountain species of the western USA, were surveyed to evaluate associations among leaf structure, orientational properties, and the sunlight exposure and precipitation characteristic of each community. Selected leaf structural features have been associated previously with photosynthetic function and included shape, thickness, the ratio of thickness to width, stomatal distribution, leaf surface coloration, and the number and distribution of palisade cell layers. Decreases in annual precipitation (<4 to over 15 cm/yr) and increases in total daily sunlight (4.2 to 29.2 mol photons/m1) corresponded strongly to an increase in the percentage of species in a given community with more inclined (more inclined than +/- 45 degrees from horizontal) or thicker leaf mesophyll (>0.4 mm) leaves. Also, the percentage of species with a leaf thickness to width ratio >0.1, which were amphistomatous, or which had palisade cell layers beneath both leaf surfaces, increased from >20% in the highest rainfall and lowest sunlight community to >80% in the community with least rainfall but greatest sunlight exposure. Over 70% of the species in the most mesic, shaded community had lighter abaxial than adaxial leaf surfaces (leaf bicoloration). All of the above structural features were positively associated with a more inclined leaf orientation (r1 = 0.79), except for leaf bicoloration, which was negatively associated (r1 = 0.75). The ratio of adaxial to abaxial light was more strongly associated with leaf bicoloration (r1 = 0.83) and the presence of multiple adaxial and isobilateral palisade cell layers(r1 = 0.80) than with total incident sunlight on just the adaxial leaf surface (r1 = 0.69 and 0.73, respectively). These results provide field evidence that leaf orientation and structure may have evolved in concert to produce a photosynthetic symmetry in leaf structure in response to the amount of sunlight and other limiting factors of the community. This structural symmetry may serve fundamentally to regulate the distribution of both light and CO2 levels inside the leaf and, thus, increase photosynthetic CO2 uptake per unit leaf biomass.