An Intrinsic Geometric Constraint on Morphological Stomatal Traits

A strong negative non-linear relationship exists between stomatal density (SD) and size (SS) or length (SL), which is of high importance in gas exchange and plant evolution. However, the cause of this relationship has not been clarified. In geometry, SD has an intrinsic relationship with SS^-1 or SL^-2, which is defined as a geometric constraint here. We compiled global data to clarify the influence of this geometric constraint on the SD-SS relationship. The log-log scaling slope of the relationship between SD and SS and between SD and SL was not significantly different from -1 and -2, respectively. Although the non-geometric effect drove the SD-SS curve away from the power function with -1, a larger influence of the geometric constraint on SD was found. Therefore, the higher geometric constraint possibly causes the SD-SS relationship to be inevitably non-linear and negative. Compared to pteridophyta and gymnosperms, the geometric constraint was lower for angiosperm species, possibly due to most of them having smaller stomata. The relaxation of the geometric constraint seems to extend the upper range of SD in angiosperm species and hence enable them to exploit a wide range of environments.