Leaf Stable Carbon Isotope Composition Research Articles

Updating the occurrence of crassulacean acid metabolism (CAM) in the genus Clusia through carbon isotope analysis of species from Colombia

Clusia is a widely distributed neotropical genus with 321 currently described species. This remarkable genus is the only one known to contain trees sensu stricto with CAM photosynthesis. To survey the occurrence of CAM in Clusia species from Colombia, we determined the leaf stable carbon isotope composition (δ13C) of 568 specimens from 114 species deposited in 12 Colombian herbaria. In the vast majority of specimens, δ13C values indicated that C3 photosynthesis was the principal contributor to carbon gain. δ13C values typical of strong CAM (less negative than -20‰) were observed in only five species, in four of them for the first time. All samples with CAM-type isotopic signatures were collected below 1,000 m a.s.l., whereas species with predominantly C3 occurred from sea level to 3,500 m a.s.l. Together with information already available in the literature, we conclude that CAM is present in 22% (35/156) of the species of Clusia investigated thus far.

Shade tree canopy cover affects coffee plant traits across elevations in coffee farms in southwest Ethiopia

Coffee is an important crop in the global south. However, ongoing changes in the climate system reinforce the need to quantify coffee plants' ecological and eco‐physiological traits to assure coffee production in the future. One way to assess how environmental changes affect coffee performance is via leaf traits, most notably leaf carbon and nitrogen concentrations (to reflect the nutrient status), leaf stable carbon isotope composition (δ13C) to determine intrinsic water use efficiency (WUEi), and specific leaf area (SLA) to describe carbon gain relative to water loss within a plant canopy as these traits are related to yields. Therefore, we sampled coffee plants growing at contrasting elevations using a space‐for‐time substitution approach for warming and superimposed a canopy cover gradient to assess whether increasing canopy cover could modulate responses to temperature. Three coffee shrubs were sampled in each of 59 coffee farms in southwest Ethiopia across elevations of 1500–2160 m a.s.l. and along canopy cover gradients from open to deep shade. Soil nutrient concentrations, light availability, soil temperature and moisture were quantified for each coffee shrub. Elevation and shade tree canopy cover significantly and interactively affected WUEi. Elevation was found to be the driving factor for microclimate and soil factors which indirectly influenced both SLA and WUEi. Both of these coffee leaf traits are moderately governed by soil temperature whereas leaf N and C:N are mainly controlled by soil temperature and soil chemical variables. As elevation increased, WUEi kept increasing at light (< 35%) to intermediate shade levels (35–65%), and the values decreased at dense shade levels (65–100%) at high elevations, suggesting that coffee plants growing at high elevations with light shade can assimilate more CO2with minimum evaporative water loss. SLA declined with elevation. Leaf N and leaf C:N responded negatively and positively to shade canopy cover, respectively. In sum, elevation and canopy cover interactively determined microclimate and coffee leaf traits. Our findings are useful to adjust the intensity of shade, along with other tree‐level management tools, to modulate climate‐change effects on coffee at the farm level.

The genetic architecture of leaf stable carbon isotope composition in Zea mays and the effect of transpiration efficiency on leaf elemental accumulation.

With increased demand on freshwater resources for agriculture, it is imperative that more water-use efficient crops are developed. Leaf stable carbon isotope composition, δ13C, is a proxy for transpiration efficiency and a possible tool for breeders, but the underlying mechanisms effecting δ13C in C4 plants are not known. It has been suggested that differences in specific leaf area (SLA), which potentially reflects variation in internal CO2 diffusion, can impact leaf δ13C. Furthermore, although it is known that water movement is important for elemental uptake, it is not clear how manipulation of transpiration for increased water-use efficiency may impact nutrient accumulation. Here, we characterize the genetic architecture of leaf δ13C and test its relationship to SLA and the ionome in five populations of maize. Five significant QTL for leaf δ13C were identified, including novel QTL as well as some that were identified previously in maize kernels. One of the QTL regions contains an Erecta-like gene, the ortholog of which has been shown to regulate transpiration efficiency and leaf δ13C in Arabidopsis. QTL for δ13C were located in the same general chromosome region, but slightly shifted, when comparing data from two different years. Our data does not support a relationship between δ13C and SLA, and of the 19 elements analyzed, only a weak correlation between molybdenum and δ13C was detected. Together these data add to the genetic understanding of leaf δ13C in maize and suggest that improvements to plant water use may be possible without significantly influencing elemental homeostasis.

Large-sized rare tree species contribute disproportionately to functional diversity in resource acquisition in African tropical forest.

Increasing evidence is available for a positive effect of biodiversity on ecosystem productivity and standing biomass, also in highly diverse systems as tropical forests. Biodiversity conservation could therefore be a critical aspect of climate mitigation policies. There is, however, limited understanding of the role of individual species for this relationship, which could aid in focusing conservation efforts and forest management planning. This study characterizes the functional specialization and redundancy for 95% of all tree species (basal area weighted percentage) in a diverse tropical forest in the central Congo Basin and relates this to species' abundance, contribution to aboveground carbon, and maximum size. Functional characterization is based on a set of traits related to resource acquisition (wood density, specific leaf area, leaf carbon, nitrogen and phosphorus content, and leaf stable carbon isotope composition). We show that within both mixed and monodominant tropical forest ecosystems, the highest functional specialization and lowest functional redundancy are solely found in rare tree species and significantly more in rare species holding large‐sized individuals. Rare species cover the entire range of low and high functional redundancy, contributing both unique and redundant functions. Loss of species supporting functional redundancy could be buffered by other species in the community, including more abundant species. This is not the case for species supporting high functional specialization and low functional redundancy, which would need specific conservation attention. In terms of tropical forest management planning, we argue that specific conservation of large‐sized trees is imperative for long‐term maintenance of ecosystem functioning.

Leaf stable carbon isotope composition reflects transpiration efficiency in Zea mays.

The increasing demand for food production and predicted climate change scenarios highlight the need for improvements in crop sustainability. The efficient use of water will become increasingly important for rain-fed agricultural crops even in fertile regions that have historically received ample precipitation. Improvements in water-use efficiency in Zea mays have been limited, and warrant a renewed effort aided by molecular breeding approaches. Progress has been constrained by the difficulty of measuring water-use in a field environment. The stable carbon isotope composition (δ13 C) of the leaf has been proposed as an integrated signature of carbon fixation with a link to stomatal conductance. However, additional factors affecting leaf δ13 C exist, and a limited number of studies have explored this trait in Z.mays. Here we present an extensive characterization of leaf δ13 C in Z.mays. Significant variation in leaf δ13 C exists across diverse lines of Z.mays, which we show to be heritable across several environments. Furthermore, we examine temporal and spatial variation in leaf δ13 C to determine the optimum sampling time to maximize the use of leaf δ13 C as a trait. Finally, our results demonstrate the relationship between transpiration and leaf δ13 C in the field and the greenhouse. Decreasing transpiration and soil moisture are associated with decreasing leaf δ13 C. Taken together these results outline a strategy for using leaf δ13 C and reveal its usefulness as a measure of transpiration efficiency under well-watered conditions rather than a predictor of performance under drought.

Spatial Variation in Leaf Stable Carbon Isotope Composition of Three Caragana Species in Northern China

Leaf stable carbon isotope (δ13C) composition provides comprehensive insight into plant carbon cycles and water use efficiency and has also been widely used to evaluate the response of plants to environmental change. In the present study, leaf δ13C was analyzed in samples of Caragana microphylla Lam., C. liouana Zhao, and C. korshinskii Kom. from 38 populations. These species provide great environmental benefits and economic value and are distributed east to west continuously across northern China. We studied the relationship of δ13C to altitude, mean annual precipitation (MAP), mean annual temperature (MAT), mean annual relative humidity (RH), leaf nitrogen (N), and phosphorus (P) concentrations to examine the patterns and controls of leaf δ13C variation in each species. Results indicated that, across the three species, leaf δ13C significantly decreased with MAP, RH, and leaf N and P concentrations, while it increased with altitude and MAT. However, patterns and environmental controls of leaf δ13C varied proportionally with species. C. korshinskii was mainly controlled by MAP and leaf N concentration, C. liouana was controlled by both MAT and MAP, and C. microphylla was mainly controlled by MAT. Further analysis indicated significant differences in leaf δ13C between species, which tended to increase from C. microphylla to C. korshinskii. Overall, these results suggest that the three Caragana species may respond differently to future climate change due to different controlling factors on leaf δ13C variation, as well as differentiation in water use efficiency, which likely contributes to the geographical distribution of these species.

Response of leaf stable carbon isotope composition to temporal and spatial variabilities of aridity index on two opposite hillslopes in a native vegetated catchment

The stable carbon isotope composition (δ13C) has been demonstrated to be a useful indicator of environmental conditions occurring during plant growth. Previous studies suggest that tree leaf δ13C is correlated with mean annual precipitation (MAP) over a broad range of climates with precipitation between 100 and 2000mm/year. However, this relationship confirmed at the large scale may not be present at the local scale with complex terrain where factors other than precipitation may lead to additional variability in plant water stress.In this study, we investigated δ13C of tree leaves in a native vegetation catchment over a local gradient of hydro-climatic conditions induced by two hillslopes with opposite aspects. Significant seasonal variations, calculated as a difference between the maximum and minimum δ13C values for each tree, were observed for two species, up to 1.9‰ for Eucalyptus (E.) paniculata, and up to 2.7‰ for Acacia (A.) pycnantha on the north-facing slope (NFS). Also the mean δ13C values calculated from all investigated trees of each hillslope were significantly different and leaf δ13C on the NFS was higher by 1.4±0.5‰ than that on the south-facing slope (SFS). These results cannot be explained by the negligible difference in precipitation between the two hillslopes located just 200m apart. The correlation coefficients between the δ13C of E. tree leaves and the integrated aridity index (AI) were statistically significant for temporal observations on the NFS (R2 0.18–0.44, p-value 0.00–0.06), and spatial observations (R2=0.35, p-value 0.05) at the end of the dry season. These results suggest that AI as a measure of plant water stress is better associated with leaf δ13C than precipitation. Therefore, leaf δ13C value can be used as a valuable proxy for plant water stress across the landscape in both time and space.

Water-use responses of 'living fossil' conifers to CO2 enrichment in a simulated Cretaceous polar environment.

During the Mesozoic, the polar regions supported coniferous forests that experienced warm climates, a CO(2)-rich atmosphere and extreme seasonal variations in daylight. How the interaction between the last two factors might have influenced water use of these conifers was investigated. An experimental approach was used to test the following hypotheses: (1) the expected beneficial effects of elevated [CO(2)] on water-use efficiency (WUE) are reduced or lost during the 24-h light of the high-latitude summer; and (2) elevated [CO(2)] reduces plant water use over the growing season. Measurements of leaf and whole-plant gas exchange, and leaf-stable carbon isotope composition were made on one evergreen (Sequoia sempervirens) and two deciduous (Metasequoia glyptostroboides and Taxodium distichum) 'living fossil' coniferous species after 3 years' growth in controlled-environment simulated Cretaceous Arctic (69 degrees N) conditions at either ambient (400 micromol mol(-1)) or elevated (800 micromol mol(-1)) [CO(2)]. Stimulation of whole-plant WUE (WUE(P)) by CO(2) enrichment was maintained over the growing season for the three studied species but this pattern was not reflected in patterns of WUE inferred from leaf-scale gas exchange measurements (iWUE(L)) and delta(13)C of foliage (tWUE(L)). This response was driven largely by increased rates of carbon uptake, because there was no overall CO(2) effect on daily whole-plant transpiration or whole-plant water loss integrated over the study period. Seasonal patterns of tWUE(L) differed from those measured for iWUE(L). The results suggest caution against over simplistic interpretations of WUE(P) based on leaf isotopic composition. The data suggest that the efficiency of whole-tree water use may be improved by CO(2) enrichment in a simulated high-latitude environment, but that transpiration is relatively insensitive to atmospheric CO(2) in the living fossil species investigated.

The effects of enhanced ultraviolet-B radiation and soil drought on water use efficiency of spring wheat

The effect of enhanced ultraviolet-B radiation (280–315 nm) and water stress on water consumption, instantaneous water use efficiency (WUEi), season-long water use efficiency (WUEs) and leaf stable carbon isotope composition (δ 13C) of three spring wheat cultivars ( Triticum aestivum L.) was investigated under field conditions. The relationship between WUEi and WUEs with δ 13C was analyzed. Compared with the control, enhanced UV-B or water stress alone or in combination led to lower water use, and soil drought had a stronger influence on water use than supplementary UV-B irradiance. Soil drought increased the instantaneous water use efficiency (WUEi) and UV-B radiation decreased it significantly in comparison to the control. The combination of UV-B and water stress resulted in increased/reduced or no changed WUEi, different with change. Season-long water use efficiency (WUEs) showed the same trend as observed with WUEi under the conditions of UV-B radiation and water stress, except that no significant difference between control and drought in cv. Heshangtou. WUEs under the combined conditions of UV-B and water stress, was clearly increased in every cultivar. Enhanced UV-B radiation and the combination with drought led to negative foliar stable carbon isotope composition (δ 13C) and drought alone resulted in a positive value for δ 13C. The relationship between foliar stable carbon isotope composition and instantaneous water use efficiency was not significant. Nevertheless, a positive correlation with δ 13C against season-long water use efficiency was observed. The results indicated that δ 13C can be a useable parameter for selecting a crop genotype having higher water use efficiency.

A preliminary study of the responsiveness to seasonal atmospheric and rainfall patterns of wash woodland species in the arid Richtersveld

Seasonal carbon and water relations were compared among seven tree or shrub wash woodland species in the winter rainfall desert of the Richtersveld National Park, South Africa. Plants were generally aseasonal with respect to gas exchange, but responsive to rainfall events with respect to water relations and phenology. Relatively narrow annual ranges in potential evapotranspiration due to the maritime influence could explain why these plants respond more to fluctuations in water acquisition potential than to evaporative demand. Two species were summer-deciduous, but one of them (Ozoroa concolor) responded to aseasonal summer rainfall by leafing out and flowering. These two species had high shoot xylem water potentials when in leaf. All other species were sclerophyllous evergreens with low water potentials, particularly the shallow-rooted shrub Zygophyllum prismatocarpum, and Boscia albitrunca which may have a different rooting pattern to the other phreatophytes. The latter species was also unique due to its high leaf nitrogen contents, photosynthetic rates and stomatal conductances, despite very low leaf water potentials. Leaf stable carbon isotope composition C13δC) varied between species (−22 to −27‰), but was lower than the mean for arid regions worldwide. The values indicated moderately high levels of water use efficiency, but a less conservative strategy in two species, including Boscia albitrunca. The affinities of these species to summer rainfall biomes, their apparent decline in the western arid regions in recent geological history following aridification, and their absence southwards in the winter rainfall regions, suggest that these wash species rely on sporadic summer rainfall events to some extent. They may be at risk if predicted increases in temperature and changes in rainfall patterns alter their effective moisture availability.

Leaf Stable Carbon Isotope Composition Records Increased Water-Use Efficiency of C 3 Plants in Response to Atmospheric CO 2 Enrichment

A total of 17 temperate C 3 grass and herb species were grown for 5 weeks at three mole fraction treatments of atmospheric CO 2 (350, 525 and 700 μmol mol -1 ). Leaf stable carbon isotope compositions (δ 13 C) were determined to record long-term exchange responses together with instantaneous gas exchange measurements. The isotopic composition of the atmospheric CO 2 (δ 13 C a ) integrated over the course of the CO 2 treatments was recorded biologically using the C 4 species Zea mays. We found that increases in the mole fraction of atmospheric CO 2 above current levels resulted in a sustained increase in instantaneous (photosynthesis, A/conductance, g s ) leaf water-use efficiency (IWUE), as calculated from carbon isotope-derived p i /p a ratios. Grass species showed a marked decline in the magnitude of WUE increase as the CO 2 mole fraction was increased from 525 to 700 μmol mol -1 , a response which was absent in herb species. Isotopic derivation of the ratio of intercellular CO 2 mole fraction (p i ) to that in the surrounding atmosphere (p a ), considered as a set point of leaf metabolism, showed no significant (P = 0.06) changes in response to increases in the mole fraction of CO 2 , for herb and grass species. Measurements of p i /p a determined from measurements of leaf gas exchange differed significantly (P<0.01) from those derived from stable isotope ratios. These differences are attributed to contrasting stomatal behaviour between herb and grass species. Leaf intercellular CO 2 mole fraction and previously reported above-ground biomass responses to CO 2 increases for the same species were positively correlated (P<0.05). This suggests that as atmospheric CO 2 levels continue to rise species showing sustained higher rates of leaf photosynthesis, may be translated into increased productivity depending on soil water and nutrient status.

427 PB 162 INTERACTIONS BETWEEN POTASSIUM AND OTHER MINERAL ELEMENTS IN DROUGHT STRESSED AND NON-DROUGHT STRESSED HIBISCUS SINENSIS L. CV. LEPRECAUN

Hibiscus plants, were irrigated with full strength Hoagland's nutrient solution containing either 0,2,5, or 10 mM potasium(K). After 72 days of K treatment, half of the plants at each K level were subjected to a 21-day slowly developing drought stress cycle and the other half were non-drought stressed (ND). Mid-day leaf water potentials at day 21 was-1.5 to-1.6 MPa (DS), and -0.5 MPa (ND). Leaf K concentration increased with increasing K in nutrient solution for both DS and ND plants, but K was higher in DS than ND plants at 2.5 and 10 mM K. Of the macronutrient cations, only (Ca) was inversely correlated with nutrient solution K, in both DS and ND plants. Leaf concentrations of all the micronutrient cations increased with increasing K supply, regardless of drought stress. Potassium hadt significant positive correlation with total plant and leaf dry weight of DS, but not ND plants. Leaf stable carbon isotope composition (δ13 C,an estimate of long term water-use efficiency), was positively correlated with N, Mg and Ca, and negatively correlated with K, iron (Fe), and K:total cation ratio regardless of drought stress. Both net photosynthesis and stomatal conductance were negatively correlated with N and Ca, but positively correlated with K, Fe and manganese in ND plants.

Shifts in the δ 13 C composition of Salix herbacea L. leaves in response to spatial and temporal gradients of atmospheric CO 2 concentration

We investigated the response of three carbon isotope related parameters ($\delta ^{13}$C, discrimination and p$\_{i}$/p$\_{a}$ ratio) determined from Salix herbacea leaves to an altitudinal gradient in Austria and to short- (200 years) and long-term (> 140 000 years) increases in CO$\_{2}$ concentration. The altitudinal results showed leaf $\delta ^{13}$C declined with increasing altitude. This supports the view that local trends in leaf $\delta ^{13}$C may follow quite different patterns to those observed globally. The effects of short-term CO$\_{2}$ increases were investigated by analysis of dated herbarium material collected over the past two centuries. Leaf $\delta ^{13}$C declined in response to CO$\_{2}$ increases of the past 200 years, which probably reflects the anthropogenic increase of atmospheric $^{12}$C as a result of fossil fuel burning and deforestation. Quaternary leaf macrofossils were analysed to examine the response of leaf $\delta ^{13}$C to long-term alterations in CO$\_{2}$ and climate represented by a glacial cycle. The results suggest leaf $\delta ^{13}$C reflects the palaeotemperature under which the leaves formed, low temperatures being associated with high $\delta ^{13}$C. This is consistent both with observations from a global latitudinal survey of leaf $\delta ^{13}$C ranging from equatorial to polar regions and with experimental measurements made on plants grown at low temperatures. Comparisons of the general temperature trends of the Devensian late-glacial inferred from fossil leaf $\delta ^{13}$C values with palaeotemperature reconstructions from beetle remains and pollen show a similar overall pattern of events. Quaternary fossil leaves may therefore provide evidence for atmospheric CO$_{2}$ change (from the stomatal density record) and palaeotemperatures (from leaf $\delta ^{13}$C content) which together offer the exciting possibility of determining the comparative timing of both events.