Leaf Intrinsic Water Use Efficiency Research Articles

Drivers of Natural Variation in Water-Use Efficiency Under Fluctuating Light Are Promising Targets for Improvement in Sorghum.

Improving leaf intrinsic water-use efficiency (iWUE), the ratio of photosynthetic CO2 assimilation to stomatal conductance, could decrease crop freshwater consumption. iWUE has primarily been studied under steady-state light, but light in crop stands rapidly fluctuates. Leaf responses to these fluctuations substantially affect overall plant performance. Notably, photosynthesis responds faster than stomata to decreases in light intensity: this desynchronization results in substantial loss of iWUE. Traits that could improve iWUE under fluctuating light, such as faster stomatal movement to better synchronize stomata with photosynthesis, show significant natural diversity in C3 species. However, C4 crops have been less closely investigated. Additionally, while modification of photosynthetic or stomatal traits independent of one another will theoretically have a proportionate effect on iWUE, in reality these traits are inter-dependent. It is unclear how interactions between photosynthesis and stomata affect natural diversity in iWUE, and whether some traits are more tractable drivers to improve iWUE. Here, measurements of photosynthesis, stomatal conductance and iWUE under steady-state and fluctuating light, along with stomatal patterning, were obtained in 18 field-grown accessions of the C4 crop sorghum. These traits showed significant natural diversity but were highly correlated, with important implications for improvement of iWUE. Some features, such as gradual responses of photosynthesis to decreases in light, appeared promising for improvement of iWUE. Other traits showed tradeoffs that negated benefits to iWUE, e.g., accessions with faster stomatal responses to decreases in light, expected to benefit iWUE, also displayed more abrupt losses in photosynthesis, resulting in overall lower iWUE. Genetic engineering might be needed to break these natural tradeoffs and achieve optimal trait combinations, e.g., leaves with fewer, smaller stomata, more sensitive to changes in photosynthesis. Traits describing iWUE at steady-state, and the change in iWUE following decreases in light, were important contributors to overall iWUE under fluctuating light.

Comparative photosynthesis physiology of cultivated and wild lettuce under control and low‐water stress

AbstractLettuce (Lactuca sativa L.) is an important vegetable crop; however, increased demand and reduced recharge to the aquifer systems in the major field growing regions are making irrigation requirements more difficult and costly to meet. If lettuce production is to be sustainable in these regions, there is a need to breed varieties with higher water use efficiency (WUE) that produce similar biomass under reduced irrigation. The wild progenitor of lettuce, Lactuca serriola L., is a common, ruderal weed that is generally considered more drought tolerant than its cultivated relative. We compared the photosynthetic physiology of two representative accessions of these species under progressive low‐water stress for 10 d and observed that leaf intrinsic WUE was higher under moderate low‐water stress in L. serriola acc. US96UC23 than lettuce cultivar ‘Salinas’. Carbon assimilation (A) and carboxylating efficiency was generally higher in US96UC23 under control water treatments, which we determine to be due to higher mesophyll conductance (gm). No differences were observed between the two species in concentrations of the carboxylating enzyme Rubisco. We find mesophyll limitations to be the most significant relative limiting factor in Salinas. The domestication of cultivated lettuce from this wild relative involved intensive morphological alteration of the leaf, which appears to have led to significantly reduced gm and, concomitantly, A. Breeding for improved gm in lettuce may lead to increases in A and WUE under moderate low‐water stress.

A comparative study on physiological responses to drought in walnut genotypes (RX1, Vlach, VX211) commercially available as rootstocks

Walnut production in California depends on the performance of three clonal hybrid rootstocks, and this study provides evidence that roots of genotype RX1 exhibit unique properties to cope with water stress induced by drought. Three hybrid genotypes RX1 (Juglans microcarpa × J. regia), Vlach and VX211 (both J. hindsii × J. regia) are the most commonly used rootstocks for walnut production in California. These rootstocks provide various levels of disease resistance, but their performance under drought is unknown. Based on our findings on xylem stress physiology of native walnut species, we hypothesize that hybrid genotypes originating from wild species native to drier habitats will exhibit superior root performance under drought stress. Using a whole-plant experimental approach, we evaluated root and canopy physiological characteristics of 2-year-old tree saplings of RX1, Vlach and VX211 under two soil moisture treatments (‘control’ 70–90% and ‘drought’ 20–40% w/w soil moisture). In control saplings, root biomass was twofold smaller in RX1 as compared to VX211, but root system hydraulic conductance (Kro, predominantly cell-to-cell pathway) was more than 50% greater in RX1 and Vlach as compared to VX211. The relatively low Kro of VX211 was related to a larger number of root cortical cell layers and endodermis development. In drought-stressed saplings, root biomass was reduced by 27% (P < 0.05) in VX211 but no significant reduction in root biomass was detected in RX1 and Vlach. Maintenance of root biomass under drought in RX1 and Vlach was associated with an 80% decrease in Kro, a threefold increase in leaf intrinsic water-use efficiency, and maintenance of leaf turgor as compared to control conditions. Drought-induced reductions in Kro were linked to the formation of a multiseriate root endodermis in all three genotypes.

The response of mesophyll conductance to nitrogen and water availability differs between wheat genotypes

Increased mesophyll conductance (gm) has been suggested as a target for selection for high productivity and high water-use efficiency in crop plants, and genotypic variability in gm has been reported in several important crop species. However, effective selection requires an understanding of how gm varies with growth conditions, to ensure that the ranking of genotypes is consistent across environments. We assessed the genotypic variability in gm and other leaf gas exchange traits, as well as growth and biomass allocation for six wheat genotypes under different water and nitrogen availabilities. The wheat genotypes differed in their response of gm to growth conditions, resulting in genotypic differences in the mesophyll limitation to photosynthesis and a significant increase in the mesophyll limitation to photosynthesis under drought. In this experiment, leaf intrinsic water-use efficiency was more closely related to stomatal conductance than to mesophyll conductance, and stomatal limitation to photosynthesis increased more in some genotypes than in others in response to drought. Screening for gm should be carried out under a range of growth conditions.

Variation in mesophyll conductance among Australian wheat genotypes

CO2 diffusion from substomatal intercellular cavities to sites of carboxylation in chloroplasts (mesophyll conductance; gm) limits photosynthetic rate and influences leaf intrinsic water-use efficiency (A/gsw). We investigated genotypic variability of gm and effects of gm on A/gsw among eleven wheat (Triticum aestivum L.) genotypes under light-saturated conditions and at either 2 or 21% O2. Significant variation in gm and A/gsw was found between genotypes at both O2 concentrations, but there was no significant effect of O2 concentration on gm. Further, gm was correlated with photosynthetic rate among the 11 genotypes, but was unrelated to stomatal conductance. The effect of leaf age differed between genotypes, with gm being lower in older leaves for one genotype but not another. This study demonstrates a high level of variation in gm between wheat genotypes; 0.5 to 1.0μmolm-2s-1 bar-1. Further, leaf age effects indicate that great care must be taken to choose suitable leaves in studies of genotypic variation in gm and water-use efficiency.

δ13C of leaf‐respired CO2 reflects intrinsic water‐use efficiency in barley

Leaf intrinsic water-use efficiency (WUE), the ratio of photosynthetic rate to stomatal conductance (A/g(s) ), is a key plant trait linking terrestrial carbon and water cycles. A rapid, integrative proxy for A/g(s) is of benefit to crop breeding programmes aiming to improve WUE, but also for ecologists interested in plant carbon-water balance in natural systems. We hypothesize that the carbon isotope composition of leaf-respired CO(2) (δ(13) C(Rl) ), two hours after leaves are transferred to the dark, records photosynthetic carbon isotope discrimination and so provides a proxy for A/g(s) . To test this hypothesis, δ(13) C(Rl) was measured in four barley cultivars grown in the field at two levels of water availability and compared to leaf-level gas exchange (the ratio of leaf intercellular to ambient CO(2) partial pressure, C(i) /C(a) , and A/g(s) ). Leaf-respired CO(2) was more (13) C-depleted in plants grown at higher water availability, varied between days as environmental conditions changed, and was significantly different between cultivars. A strong relationship between δ(13) C(Rl) and δ(13) C of sucrose was observed. δ(13) C(Rl) was converted into apparent photosynthetic discrimination (Δ(13) C(Rl) ) revealing strong relationships between Δ(13) C(Rl) and C(i) /C(a) and A/g(s) during the vegetative stage of growth. We therefore conclude that δ(13) C(Rl) may provide a rapid, integrative proxy for A/g(s) in barley.

Competition for water between walnut seedlings (Juglans regia) and rye grass (Lolium perenne) assessed by carbon isotope discrimination and delta18O enrichment.

Container-grown walnut seedlings (Juglans regia L.) were subjected to competition with rye grass (Lolium perenne L.) and to a 2-week soil drying cycle. One and 2 weeks after the beginning of the drought treatment, H2 18O (delta approximately equals +100%) was added to the bottom layer of soil in the plant containers to create a vertical H2 18O gradient. Rye grass competition reduced aboveground and belowground biomass of the walnut seedlings by 60%, whereas drought had no effect. The presence of rye grass reduced the dry weight of walnut roots in the upper soil layer and caused a 50% reduction in lateral root length. Rye grass competition combined with the drought treatment reduced walnut leaf CO2 assimilation rate (A) and leaf conductance (gw) by 20 and 39%, respectively. Transpiration rates in rye grass, both at the leaf level and at the plant or tiller level, were higher than in walnut seedlings. Leaf intrinsic water-use efficiency (A/gw) of walnut seedlings increased in response to drought and no differences were observed between the single-species and mixed-species treatments, as confirmed by leaf carbon isotope discrimination measurements. Measurement of delta18O in soil and in plant xylem sap indicated that the presence of rye grass did not affect the vertical profile of soil water uptake by walnut seedlings. Walnut seedlings and rye grass withdrew water from the top and middle soil layers in well-watered conditions, whereas during the drought treatment, walnut seedlings obtained water from all soil layers, but rye grass took up water from the bottom soil layer only.

Interspecific variability of δ13C among trees in rainforests of French Guiana: functional groups and canopy integration.

The interspecific variability of sunlit leaf carbon isotope composition (δ13C), an indicator of leaf intrinsic water-use efficiency (WUE, CO2 assimilation rate/leaf conductance for water vapour), was investigated in canopy trees of three lowland rainforest stands in French Guiana, differing in floristic composition and in soil drainage characteristics, but subjected to similar climatic conditions. We sampled leaves with a rifle from 406 trees in total, representing 102 species. Eighteen species were common to the three stands. Mean species δ13C varied over a 6.0‰ range within each stand, corresponding to WUE varying over about a threefold range. Species occurring in at least two stands displayed remarkably stable δ13C values, suggesting a close genetic control of species δ13C. Marked differences in species δ13C values were found with respect to: (1) the leaf phenology pattern (average δ13C=-29.7‰ and -31.0‰ in deciduous-leaved and evergreen-leaved species, respectively), and (2) different types of shade tolerance defined by features reflecting the plasticity of growth dynamics with respect to contrasting light conditions. Heliophilic species exhibited more negative δ13C values (average δ13C=-30.5‰) (i.e. lower WUE) than hemitolerant species (-29.3‰). However, tolerant species (-31.4‰) displayed even more negative δ13C values than heliophilic ones. We could not provide a straightforward ecophysiological interpretation of this result. The negative relationship found between species δ13C and midday leaf water potential (Ψwm) suggests that low δ13C is associated with high whole tree leaf specific hydraulic conductance. Canopy carbon isotope discrimination (Δ A ) calculated from the basal area-weighed integral of the species δ13C values was similar in the three stands (average Δ A =23.1‰), despite differences in stand species composition and soil drainage type, reflecting the similar proportions of the three different shade-tolerance types among stands.

Functional diversity in an Amazonian rainforest of French Guyana: a dual isotope approach (δ15N and δ13C).

Functional aspects of biodiversity were investigated in a lowland tropical rainforest in French Guyana (5°2'N, annual precipitation 2200 mm). We assessed leaf δ15N as a presumptive indicator of symbiotic N2 fixation, and leaf and wood cellulose δ13C as an indicator of leaf intrinsic water-use efficiency (CO2 assimilation rate/leaf conductance for water vapour) in dominant trees of 21 species selected for their representativeness in the forest cover, their ecological strategy (pioneers or late successional stage species, shade tolerance) or their potential ability for N2 fixation. Similar measurements were made in trees of native species growing in a nearby plantation after severe perturbation (clear cutting, mechanical soil disturbance). Bulk soil δ15N was spatially quite uniform in the forest (range 3-5‰), whereas average leaf δ15N ranged from -0.3‰ to 3.5‰ in the different species. Three species only, Diplotropis purpurea, Recordoxylon speciosum (Fabaceae), and Sclerolobium melinonii (Caesalpiniaceae), had root bacterial nodules, which was also associated with leaf N concentrations higher than 20 mg g-1. Although nodulated trees displayed significantly lower leaf δ15N values than non-nodulated trees, leaf δ15N did not prove a straightforward indicator of symbiotic fixation, since there was a clear overlap of δ15N values for nodulated and non-nodulated species at the lower end of the δ15N range. Perturbation did not markedly affect the difference δ15Nsoil - δ15Nleaf, and thus the isotopic data provide no evidence of an alteration in the different N acquisition patterns. Extremely large interspecific differences in sunlit leaf δ13C were observed in the forest (average values from -31.4 to -26.7‰), corresponding to intrinsic water-use efficiencies (ratio CO2 assimilation rate/leaf conductance for water vapour) varying over a threefold range. Wood cellulose δ13C was positively related to total leaf δ13C, the former values being 2-3‰ higher than the latter ones. Leaf δ13C was not related to leaf δ15N at either intraspecific or interspecific levels. δ13C of sunlit leaves was highest in shade hemitolerant emergent species and was lower in heliophilic, but also in shade-tolerant species. For a given species, leaf δ13C did not differ between the pristine forest and the disturbed plantation conditions. Our results are not in accord with the concept of existence of functional types of species characterized by common suites of traits underlying niche differentiation; rather, they support the hypothesis that each trait leads to a separate grouping of species.

Divergent selection for carbon isotope discrimination in crested wheatgrass

Because plant growth on semiarid rangelands is frequently water-limited, breeding for enhanced water-use efficiency (WUE, kg dry matter gained per kg water transpired) should improve forage production on these areas. In crested wheatgrass [Agropyron desertorum (Fischer ex Link) Schultes], variation for carbon isotope discrimination (Δ) has been negatively associated with WUE, suggesting that selection for lowered Δ would increase WUE. To determine the potential of altering Δ through breeding, we selected nine clones from a crested wheatgrass breeding population based on their Δ values, equally subdivided them into three groups (low, medium, and high Δ), and made a series of diallel crosses within each group. The parental clones and single-cross progenies were established in the field as replicated spaced plants on 1-m centers in spring 1989. Forage dry matter yield and Δ were determined in 1990 and 1991; leaf gas exchange traits were determined for low and high Δ classes in 1990, and for low, medium, and high Δ classes in 1991. The previous ranking of Δ classes was confirmed in the present studies. Combined across years, values for Δ, leaf CO2 exchange rate (CER), and stomatal conductance (gs) were significantly (P &lt; 0.05) lower in the low than high Δ class. When data were combined across clones and progenies, Δ was negatively correlated with leaf intrinsic water-use efficiency (WUEi, expressed as the CER/gs ratio) in 1990 (r = −0.87**, df = 14) and in 1991 (r = −0.83**, df = 23). Forage yield and Δ were not correlated, suggesting that the two traits may be under separate genetic control. Progenies from crosses among the low-Δ clones had significantly lower Δ values than progenies from either the medium-Δ or high-Δ clones during each year and when the data were combined across years. Divergent selection for low Δ also was reflected by improved WUEi in the subsequent generation. Moreover, narrow-sense heritability values for Δ were in excess of 0.75 and correlations between progeny means for Δ and means of the corresponding midparents were significant in each of the three analyses (r = 0.87** in 1990, 0.91** in 1991, and 0.92** in the combined analysis). These results complement earlier findings and confirm that genetic improvement for WUE can be effectively achieved in crested wheatgrass through indirect selection for Δ. Key words: Agropyron desertorum, water-use efficiency, parent-progeny correlations, drought, carbon isotope ratio, grass breeding, forage yield