The stomata on leaf surfaces control gas exchange and water loss, closing during dry periods to conserve water. The distribution and size of stomatal complexes is determined by epidermal cell differentiation and expansion during leaf growth. Regulation of these processes in response to water deficit may result in stomatal anatomical plasticity as part of the plant acclimation to drought. We quantified the leaf anatomical plasticity under water-deficit conditions in maize and soybean over two experiments. Both species produced smaller leaves in response to the water deficit, partly due to the reductions in the stomata and pavement cell size, although this response was greater in soybean, which also produced thicker leaves under severe stress, whereas the maize leaf thickness did not change. The stomata and pavement cells were smaller with the reduced water availability in both species, resulting in higher stomatal densities. Stomatal development (measured as stomatal index, SI) was suppressed in both species at the lowest water availability, but to a greater extent in maize than in soybean. The result of these responses is that in maize leaves, the stomatal area fraction (fgc) was consistently reduced in the plants grown under severe but not moderate water deficit, whereas the fgc did not decrease in the water-stressed soybean leaves. The water deficit resulted in the reduced expression of one of two (maize) or three (soybean) SPEECHLESS orthologs, and the expression patterns were correlated with SI. The vein density (VD) increased in both species in response to the water deficit, although the effect was greater in soybean. This study establishes a mechanism of stomatal development plasticity that can be applied to other species and genotypes to develop or investigate stomatal development plasticity.
Stomata on leaf surfaces control gas exchange and water loss, closing during dry periods to conserve water. The distribution and size of guard cells is determined by the balance between epidermal cell differentiation and expansion during leaf growth. Under water-deficit conditions, regulation of these processes could result in stomatal anatomical plasticity as part of plant acclimation to drought. We investigated stomatal anatomy plasticity under water deficit conditions in maize and soybean. In maize leaves, stomatal area fraction ( f gc ) was consistently reduced in plants grown at 50% and 30% media water content (MWC), whereas this was not the case in soybean leaves. This is because although stomatal size decreased with reduced MWC in both species, there was a large increase in stomatal density (SD) in soybean leaves due to large reductions in leaf area and pavement cell size. Stomatal index (SI) decreased in both maize and soybean leaves, but in soybean this reduction was too small relative to the decrease in pavement cell size to impact f gc . Despite reduced SI, few stomatal precursor cells were observed in waterstressed leaves of either species. Expression of one ortholog of two (maize) or three (soybean) SPEECHLESS genes was reduced in response to water deficit. This study shows that f gc reduction is enabled by stomatal development arrest being proportional to leaf size reduction in maize, whereas in soybean, when leaf size decrease is greater than the reduction in SI, SD and SS plasticity are antagonistic, resulting in maintenance of f gc .