Drought stress causes changes in vein and stomatal density. The objectives of this study were to determine (1) if the changes in vein and stomatal density are coordinated in cotton (Gossypium hirsutum L.) and (2) how these changes affect water-use efficiency (WUE). The results showed significant positive correlations between vein density and stomatal density when cotton was grown under different degrees of drought stress. WUE was significantly positively correlated with the densities of both veins and stomata. Stomatal pore area and stomatal density on the abaxial leaf side, but not the adaxial side, were significantly correlated with WUE, stomatal conductance, leaf net photosynthetic rate, and transpiration rate. In conclusion, coordinated changes in vein and stomatal density improve the WUE of cotton under drought stress. The abaxial leaf side plays a more important role than the adaxial side in WUE and gas exchange.
Mesophyll conductance (gm) has been shown to affect photosynthetic capacity and thus the estimates of terrestrial carbon balance. While there have been some attempts to model gm at the leaf and larger scales, the potential contribution of gm to the photosynthesis of non-leaf green organs has not been studied. Here, we investigated the influence of gm on photosynthesis of cotton bracts and how it in turn is influenced by anatomical structures, by comparing leaf palisade and spongy mesophyll with bract tissue. Our results showed that photosynthetic capacity in bracts is much lower than in leaves, and that gm is a limiting factor for bract photosynthesis to a similar extent to stomatal conductance. Bract and the spongy tissue of leaves have lower mesophyll conductance than leaf palisade tissue due to the greater volume fraction of intercellular air spaces, smaller chloroplasts, lower surface area of mesophyll cells and chloroplasts exposed to leaf intercellular air spaces and, perhaps, lower membrane permeability. Comparing bracts with leaf spongy tissue, although bracts have a larger cell wall thickness, they have a similar gm estimated from anatomical characteristics, likely due to the cumulative compensatory effects of subtle differences in each subcellular component, especially chloroplast traits. These results provide the first evidence for anatomical constraints on gm and photosynthesis in non-leaf green organs.
Mesophyll conductance (gm) is one of the major determinants of photosynthetic rate, for which it has an impact on crop yield. However, the regulatory mechanisms behind the decline in gm of cotton (Gossypium. spp) by drought are unclear. An upland cotton (Gossypium hirsutum) genotype and a pima cotton (Gossypium barbadense) genotype were used to determine the gas exchange parameters, leaf anatomical structure as well as aquaporin and carbonic anhydrase gene expression under well‐watered and drought treatment conditions. In this study, the decrease of net photosynthetic rate (AN) under drought conditions was related to a decline in gm and in stomatal conductance (gs). gm and gs coordinate with each other to ensure optimum state of CO2 diffusion and achieve the balance of water and CO2 demand in the process of photosynthesis. Meanwhile, mesophyll limitations to photosynthesis are equally important to the stomatal limitations. Considering gm, its decline in cotton leaves under drought was mostly regulated by the chloroplast surface area exposed to leaf intercellular air spaces per leaf area (Sc/S) and might also be regulated by the expression of leaf CARBONIC ANHYDRASE (CA1). Meanwhile, cotton leaves can minimize the decrease in gm under drought by maintaining cell wall thickness (Tcw). Our results indicated that modification of chloroplasts might be a target trait in future attempts to improve cotton drought tolerance.
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