Spatial heterogeneity in stomatal features during leaf elongation: an analysis using Rosa hybrida.

Within-leaf heterogeneity in stomatal traits poses a key uncertainty in determining a representative value for the whole leaf. Accounting for this heterogeneity, we studied stomatal initiation on expanding leaves and estimated stomatal conductance (gs) of mature leaves. The entire lamina was evaluated at four percentages of full leaflet elongation (FLE; leaflet length relative to its final length) in Rosa hybrida L. plants grown at 60% relative air humidity (RH), and at 100% FLE following cultivation at elevated (95%) RH. Over 80% of the stomata were initiated between 33 and 67% FLE, whereas stomatal growth mostly occurred afterwards. At 100% FLE, the heterogeneity in stomatal density was the result of uneven stomatal differentiation, while an uneven differentiation of epidermal cells contributed to this variation only at elevated RH. Noticeable within-leaf differences (up to 40%) in gs were calculated at 100% FLE. Avoiding leaflet periphery decreased this heterogeneity. Despite the large promotive effect of elevated RH on stomatal and pore dimensions, the within-leaf variation remained unaffected in all characters, besides pore aperture (and, thus, gs). The noted level of within-leaf variation in stomatal features demands a sampling scheme tailored to the leaf developmental stage, the feature per se and the evaporative demand during growth.