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Links between the carbon and water economies of plants are coupled by combining the biochemical demand for atmospheric CO2 with gas transfer through stomates, liquid water transport in the soil-xylem hydraulic system and sucrose export in the phloem. We formulated a model to predict stomatal conductance (gs), consistent with the maximum energy circulation concept of Lotka and Odum, by maximizing the sucrose flux out of photosynthesizing leaves. The proposed modeling approach recovers all prior results derived from stomatal optimization theories and profit-maximization arguments for the xylem hydraulic system aimed at predicting gs. The novel features of this approach are its ability to 1) predict the price of losing water in carbon units using xylem and phloem properties (i.e., the marginal water use efficiency) and 2) explain why water molecules become more expensive to exchange for CO2 molecules when soil moisture becomes limiting or when plants acclimate to new elevated atmospheric CO2 concentration. On short time-scales (sub-daily), predicted gs under many environmental stimuli were consistent with measurements reported in the literature, including a general sensitivity of gs to vapor pressure deficit and leaf water potential. During progressive droughts, differences in the coordination among the leaf, xylem, and phloem functioning determine the isohydric-to-anisohydric behavior among plants. 

Abstract The genusPinushas wide geographical range and includes species that are the most economically valued among forest trees worldwide. Pine needle length varies greatly among species, but the effects of needle length on anatomy, function, and coordination and trade‐offs among traits are poorly understood. We examined variation in leaf morphological, anatomical, mechanical, chemical, and physiological characteristics among five southern pine species:Pinus echinata,Pinus elliottii,Pinus palustris,Pinus taeda, andPinus virginiana. We found that increasing needle length contributed to a trade‐off between the relative fractions of support versus photosynthetic tissue (mesophyll) across species. From the shortest (7 cm) to the longest (36 cm) needles, mechanical tissue fraction increased by 50%, whereas needle dry density decreased by 21%, revealing multiple adjustments to a greater need for mechanical support in longer needles. We also found a fourfold increase in leaf hydraulic conductance over the range of needle length across species, associated with weaker upward trends in stomatal conductance and photosynthetic capacity. Our results suggest that the leaf size strongly influences their anatomical traits, which, in turn, are reflected in leaf mechanical support and physiological capacity. 

Abstract The ability to transport water through tall stems hydraulically limits stomatal conductance (g_s), thereby constraining photosynthesis and growth. However, some plants are able to minimize this height‐related decrease ing_s, regardless of path length. We hypothesized that kudzu (Pueraria lobata) prevents strong declines ing_swith height through appreciable structural and hydraulic compensative alterations. We observed only a 12% decline in maximumg_salong 15‐m‐long stems and were able to model this empirical trend. Increasing resistance with transport distance was not compensated by increasing sapwood‐to‐leaf‐area ratio. Compensating for increasing leaf area by adjusting the driving force would require water potential reaching −1.9 MPa, far below the wilting point (−1.2 MPa). The negative effect of stem length was compensated for by decreasing petiole hydraulic resistance and by increasing stem sapwood area and water storage, with capacitive discharge representing 8–12% of the water flux. In addition, large lateral (petiole, leaves) relative to axial hydraulic resistance helped improve water flow distribution to top leaves. These results indicate thatg_sof distal leaves can be similar to that of basal leaves, provided that resistance is highest in petioles, and sufficient amounts of water storage can be used to subsidize the transpiration stream.