Spatial and Temporal Relationships of Stomatal Development and Function in a Temperate Forest Canopy

Mechanisms that control stomatal development ultimately constrain leaf physiology by determining the anatomical maximum rate for gas-exchange (gsmax). However, we know comparatively less about how these regulatory processes define stomatal conductance (gs) and photosynthesis or how this information translates between model systems and important crop or native plant species. Here, we test relationships between stomatal development and leaf physiology that have been established in model systems by sampling trees in a mature forest ecosystem. We found that plasticity in gsmax was limited throughout the canopy, despite other changes in leaf structure and function that are driven by environmental gradients in the canopy. However, the ratio between gs and gsmax was predictive of gas flux in the canopy and species-independent. Variation in the gs : gsmax ratio appeared to minimize the energy required to control aperture size via guard cell turgor pressure, thus compensating for the initial over-investment in stomatal production. gsmax also remained a strong predictor of photosynthetic potential and intrinsic water-use efficiency. The temporal relationship between gsmax and these functional leaf traits might depend on long-term adjustments in stomatal development, which was sensitive to increases in atmospheric CO2 in our study. The absence of a spatial response and the presence of a temporal response in stomatal development infers that multiple mechanisms may integrate environmental signaling in the developmental pathway. Collectively, this research helps to define the larger significance of the stomatal mechanisms being identified in model systems.