Seasonal and diurnal leaf orientation, bifacial sunlight incidence, and leaf structure in the sand dune herb Hydrocotyle bonariensis

A conceptual model has been proposed whereby leaf orientation and resulting sunlight exposure dictate functional leaf structure. Specifically, the model states that this relationship is driven by the absolute amount and ratio of incident sunlight on adaxial and abaxial leaf surfaces. To test this model, the relationships between corresponding values of leaf orientation and incident sunlight on both leaf surfaces were measured for the sand dune herb Hydrocotyle bonariensis over a growth season, along with examination of leaf structure. For mature leaves, leaf angle from horizontal and azimuth angle significantly increased over the growing season, indicating diurnal midday avoidance and seasonal maximization of incident sunlight. Consequently, seasonal changes in leaf orientation resulted in an overall decrease in midday sunlight incidence on the adaxial surface and a slight shift in the daily occurrence of peak abaxial incidence. Adaxial surfaces received three to four times more sunlight than abaxial surfaces, and leaf cross-sections revealed relatively thick (564 μm) leaves with multiple adaxial palisade layers and stomata on both leaf surfaces, as predicted by the conceptual model and measured ratio of incident sunlight on adaxial and abaxial leaf surfaces. These data provide further evidence of the relationship between leaf orientation and resulting absolute levels of sunlight incidence on both leaf surfaces, as well as their ratio, and corresponding differences in internal and external leaf structure.