Photosynthetic Carbon Isotope Research Articles

Why leaves become isotopically lighter than photosynthetic carbon isotope discrimination explains: on the importance of post-photosynthetic fractionation.

This article comments on: Yu YZ, Liu HT, Yang F, Li L, Schäufele R, Tcherkez G, Schnyder H, Gong XY. 2024. δ13C of bulk organic matter and cellulose reveal post-photosynthetic fractionation during ontogeny in C4 grass leaves. Journal of Experimental Botany 75, 1451–1464

Variable carbon isotope fractionation of photosynthetic communities over depth in an open-ocean euphotic zone

Marine particulate organic carbon (POC) contributes to carbon export, food webs, and sediments, but uncertainties remain in its origins. Globally, variations in stable carbon isotope ratios (δ13C values) of POC between the upper and lower euphotic zones (LEZ) indicate either varying aspects of photosynthetic communities or degradative alteration of POC. During summertime in the subtropical north Atlantic Ocean, we find that δ13C values of the photosynthetic product phytol decreased by 6.3‰ and photosynthetic carbon isotope fractionation (εp) increased by 5.6‰ between the surface and the LEZ-variation as large as that found in the geologic record during major carbon cycle perturbations, but here reflecting vertical variation in δ13C values of photosynthetic communities. We find that simultaneous variations in light intensity and phytoplankton community composition over depth may be important factors not fully accounted for in common models of photosynthetic carbon isotope fractionation. Using additional isotopic and cell count data, we estimate that photosynthetic and non-photosynthetic material (heterotrophs or detritus) contribute relatively constant proportions of POC throughout the euphotic zone but are isotopically more distinct in the LEZ. As a result, the large vertical differences in εp result in significant, but smaller, differences in the δ13C values of total POC across the same depths (2.7‰). Vertical structuring of photosynthetic communities and export potential from the LEZ may vary across current and past ocean ecosystems; thus, LEZ photosynthesis may influence the exported and/or sedimentary δ13C values of both phytol and total organic carbon and affect interpretations of εp over geologic time.

Phenotypic variation in leaf photosynthetic traits, leaf area index, and carbon discrimination of field-grown wheat genotypes and their relationship with yield performance in Mediterranean environments.

Leaf water potential, gas exchange, and chlorophyll fluorescence exhibited significant differences among genotypes, high environmental effects, but low heritability. The highest-yielding and drought-tolerant genotypes presented superior harvest index and grain weight, compared to drought-susceptible ones. Physiological phenotyping can help identify useful traits related to crop performance under water-limited conditions. A set of fourteen bread wheat genotypes with contrasting grain yield (GY) was studied in eight Mediterranean environments in Chile, resulting from the combination of two sites (Cauquenes and Santa Rosa), two water conditions (rainfed-WL and irrigated-WW), and four growing seasons (2015-2018). The objectives were to (i) evaluate the phenotypic variation of leaf photosynthetic traits after heading (anthesis and grain filling) in different environments; (ii) analyze the relationship between GY and leaf photosynthetic traits and carbon isotope discrimination (Δ13C); and (iii) identify those traits that could have a greater impact in the determination of tolerant genotypes under field conditions. Agronomic traits exhibited significant genotypic differences and genotype × environment (GxE) interaction. The average GY under the WW condition at Santa Rosa was 9.2Mgha-1 (range 8.2-9.9Mgha-1) and under the WL condition at Cauquenes was 6.2Mgha-1 (range 3.7-8.3Mgha-1). The GY was closely related to the harvest index (HI) in 14 out of 16 environments, a trait exhibiting a relatively high heritability. In general terms, the leaf photosynthetic traits presented low GxE interaction, but high environmental effects and low heritability, except for the chlorophyll content. The relationships between GY and leaf photosynthetic traits were weaker when performed across genotypes in each environment, indicating low genotypic effects, and stronger when performed across environments for each genotype. The leaf area index and Δ13C also presented high environmental effects and low heritability, and their correlations with GY were influenced by environmental effects. The highest-yielding and drought-tolerant genotypes presented superior HI and grain weight, but no clear differences in leaf photosynthetic traits or Δ13C, compared to drought-susceptible ones. It seems that the phenotypic plasticity of agronomic and leaf photosynthetic traits is very important for crop adaptation to Mediterranean environments.

δ13C as a tool for iron and phosphorus deficiency prediction in crops.

Many studies proposed the use of stable carbon isotope ratio (δ13C) as a predictor of abiotic stresses in plants, considering only drought and nitrogen deficiency without further investigating the impact of other nutrient deficiencies, that is, phosphorus (P) and/or iron (Fe) deficiencies. To fill this knowledge gap, we assessed the δ13C of barley (Hordeum vulgare L.), cucumber (Cucumis sativus L.), maize (Zea mays L.), and tomato (Solanum lycopersicon L.) plants suffering from P, Fe, and combined P/Fe deficiencies during a two-week period using an isotope-ratio mass spectrometer. Simultaneously, plant physiological status was monitored with an infra-red gas analyzer. Results show clear contrasting time-, treatment-, species-, and tissue-specific variations. Furthermore, physiological parameters showed limited correlation with δ13C shifts, highlighting that the plants' δ13C, does not depend solely on photosynthetic carbon isotope fractionation/discrimination (Δ). Hence, the use of δ13C as a predictor is highly discouraged due to its inability to detect and discern different nutrient stresses, especially when combined stresses are present.

Carbon assimilation and distribution in cotton photosynthetic organs is a limiting factor affecting boll weight formation under drought.

Previous studies have documented cotton boll weight reductions under drought, but the relative importance of the subtending leaf, bracts and capsule wall in driving drought-induced reductions in boll mass has received limited attention. To investigate the role of carbon metabolism in driving organ-specific differences in contribution to boll weight formation, under drought conditions. Controlled experiments were carried out under soil relative water content (SRWC) (75 ± 5)% (well-watered conditions, control), (60 ± 5)% (moderate drought) and (45 ± 5)% (severe drought) in 2018 and 2019 with two cultivars Yuzaomian 9110 and Dexiamian 1. Under severe drought, the decreases of photosynthetic rate (Pn) and carbon isotope composition (δ13C) were observed in the subtending leaf, bract and capsule wall, suggesting that carbon assimilation of three organs was restricted and the limitation was most pronounced in the subtending leaf. Changes in the activities of sucrose phosphate synthase (SPS), sucrose synthase (SuSy), invertases as well as the reduction in expression of sucrose transporter (GhSUT1) led to variabilities in the sucrose content of three organs. Moreover, photosynthate distribution from subtending leaf to seeds plus fibers (the components of boll weight) was significantly restricted and the photosynthetic contribution rate of subtending leaf to boll weight was decreased, while contributions of bracts and capsule wall were increased by drought. This, in conjunction with the observed decreases in boll weight, indicated that the subtending leaf was the most sensitive photosynthetic organ to drought and was a dominant driver of boll weight loss under drought. Therefore, the subtending leaf governs boll weight loss under drought due to limitations in carbon assimilation, perturbations in sucrose metabolism and inhibition of sucrose transport.

Effects of Light Spectral Quality on Photosynthetic Activity, Biomass Production, and Carbon Isotope Fractionation in Lettuce, Lactuca sativa L., Plants.

The optimization of plant-specific LED lighting protocols for indoor plant growing systems needs both basic and applied research. Experiments with lettuce, Lactuca sativa L., plants using artificial lighting based on narrow-band LEDs were carried out in a controlled environment. We investigated plant responses to the exclusion of certain spectral ranges of light in the region of photosynthetically active radiation (PAR); in comparison, the responses to quasimonochromatic radiation in the red and blue regions were studied separately. The data on plant phenotyping, photosynthetic activity determination, and PAM fluorometry, indicating plant functional activity and stress responses to anomalous light environments, are presented. The study on carbon isotopic composition of photoassimilates in the diel cycle made it possible to characterize the balance of carboxylation and photorespiration processes in the leaves, using a previously developed oscillatory model of photosynthesis. Thus, the share of plant photorespiration (related to plant biomass enrichment with 13C) increased in response to red-light action, while blue light accelerated carboxylation (related to 12C enrichment). Blue light also reduced water use efficiency. These data are supported by the observations from the light environments missing distinct PAR spectrum regions. The fact that light of different wavelengths affects the isotopic composition of total carbon allowed us to elucidate the nature of its action on the organization of plant metabolism.

Leaf 13C data constrain the uncertainty of the carbon dynamics of temperate forest ecosystems

AbstractStable carbon isotope discrimination occurred in plant biophysical and biogeochemical processes can help understand plant physiology and soil biogeochemistry with respect to carbon cycling. Here, we incorporated the photosynthetic carbon isotope discrimination into a process‐based land surface model (iTEM) to test if stable carbon isotope composition (δ13C) can impose additional constraint on model parameters. Sequential data assimilation was implemented at six eddy covariance flux tower sites using carbon flux observations including gross primary productivity (GPP) and net ecosystem exchange (NEE) with and without considering foliar δ13C (δ13Cf) measurement constraints, respectively. Our model‐data fusion showed that δ13Cf can provide useful constraint on photosynthetic (Vcmax25, the maximum rate of carboxylation at 25°C) and stomatal (g1, the slope of stomatal function) parameters as well as the posterior carbon fluxes. When including δ13Cf measurement, g1 spatially varies among the six sites and is significantly correlated with annual precipitation. We incorporated the statistical relationship between g1 and annual precipitation into iTEM, which is then used to quantify the regional carbon dynamic in temperate forest ecosystems of the Northern Hemisphere. Compared with the simulation only conditioned on carbon flux observations, regional carbon flux estimations performed slightly better against the FLUXCOM products and the uncertainties of modeled carbon fluxes were reduced by 27%. Our study demonstrated that δ13Cf data constrains carbon flux uncertainties across space.

A general model for carbon isotopes in red-lineage phytoplankton: Interplay between unidirectional processes and fractionation by RubisCO

The carbon isotopic composition of organic matter preserved in marine sediments provides a window into the global carbon cycle through geologic time, including variations in atmospheric CO2 levels. Traditional models for interpreting isotope records of marine phytoplankton assume that these archives primarily reflect kinetic isotope discrimination by the carbon-fixing enzyme RubisCO. However, some in vivo and in vitro measurements appear to contradict this assumption, indicating that significant questions remain about the mechanistic underpinning of algal isotopic signatures, including the role of carbon concentrating mechanisms (CCMs). Here, we present a general model to explain photosynthetic carbon isotope fractionation (εP) in marine red-lineage phytoplankton groups; the model reproduces existing chemostat and batch culture datasets with a normalized root mean squared error (nRMSE) of 6.8%. Our framework proposes that a nutrient- and light-dependent step upstream of RubisCO is a kinetic barrier to carbon acquisition and therefore represents a significant source of isotopic fractionation. We suggest this step represents a carbon concentrating strategy that becomes favorable to cells under conditions of excess photon flux. The primary implications are that RubisCO is predicted to exert minimal isotopic control in photon-rich, nutrient-limited regimes but becomes influential as growth becomes light-limited. This framework enables both environment-specific and taxon-specific isotopic predictions. By refining the mechanistic understanding of marine photosynthetic carbon isotope fractionation, we may begin to reconcile existing datasets and reexamine Phanerozoic isotope records—including the resulting CO2 reconstructions—by emphasizing the influence of different types of resource limitation on photosynthetic carbon acquisition.

Seasonal gas exchange and resource-use efficiency in evergreen versus deciduous species from a tropical dry forest.

Tropical dry forests (TDFs) experience a long dry season in which plant species are subject to several months of water deficit. However, TDFs maintain a diverse group of plant life forms, growth forms and leaf phenology, and it is not clear how they vary in their mechanisms for coping with seasonal drought. We studied seasonal changes in leaf water potential (Ψleaf), gas exchange, photochemical activity and functional traits in evergreen and drought-deciduous species from a TDF to determine if leaf phenology mediates plant responses to drought. We found seasonal decreases in Ψleaf, stomatal conductance (gs) and transpiration rate (E), and increases in both intrinsic and instantaneous water-use efficiency. We did not find seasonal differences in photosynthetic rate (Aarea) and carbon isotope composition (δ13C); however, these traits differed between leaf phenology groups, with drought-deciduous plants having higher Aarea and δ13C than evergreen plants. We also found that plants with high leaf nitrogen concentration (Narea) also had low mass-based photosynthetic rate (Amass), photosynthetic-nitrogen-use efficiency and specific leaf area, contrary to the expected relationships given by the leaf economics spectrum. Despite higher Narea, sclerophyllous leaves maintained lower Amass, and this increased structural toughness of leaves may be imposing a stronger limitation for CO2 diffusion and hence photosynthesis. Overall, we found more water-conservative traits in deciduous than in evergreen plants, contrary to what is known about these two leaf phenology groups in other seasonal sites both at tropical and temperate latitudes.

Photosynthetic Carbon Isotope Discrimination and Effects on Daytime NEE Partitioning in a Subtropical Mixed Conifer Plantation

Isoflux-based isotopic flux partitioning (isoflux-based IFP) separates net ecosystem exchange flux (NEE) into net photosynthesis (FA) and nonfoliar respiration (FNR), and avoids the spurious correlation of component fluxes found in statistical flux partitioning. Canopy photosynthetic discrimination (Δcanopy) is a key variable in isoflux-based IFP, but it ignores diffusional and respiratory fractionations, with unclear consequences for isoflux-based IFP. We used three versions of the Farquhar model (Δclassical, Δdiffusion, and Δsimple) to estimate Δcanopy based on in-situ measurements of CO2 isotope ratios and flux during a dry period (I; DOY 205–219, 2015) and a humid period (II; DOY 285–299, 2015) in a subtropical mixed conifer plantation. We found no significant differences in Δcanopy between Δclassical and Δdiffusion which ignored respiratory fractionations during photorespiration and dark respiration. The average differences between Δclassical and Δsimple, which ignored respiratory and diffusional fractionations across the leaf boundary layer and mesophyll, were 2.88 ± 2.34‰ and 2.16 ± 2.18‰ for the two periods. Similarly, there were no significant differences in FA with isoflux-based IFP using Δclassical and Δdiffusion to obtain Δcanopy; FA using Δsimple was 102 and 117% higher than that using Δclassical for the dry and humid periods. Moreover, using Δclassical and Δdiffusion captured the reduction in daytime respiration during the dry period; this may be associated with the Kok effect and seasonal drought. Sensitivity analysis showed that changes in mesophyll and stomatal conductances of ±50% resulted in > 15% change in FA. Diffusional fractionations in mesophyll and stoma are critical limitations to isoflux-based IFP, while photorespiratory fractionation is an unignorable component of Δcanopy and isoflux-based IFP.

Influence of starch deficiency on photosynthetic and post-photosynthetic carbon isotope fractionations.

Carbon isotope (13C) fractionations occurring during and after photosynthetic CO2 fixation shape the carbon isotope composition (δ13C) of plant material and respired CO2. However, responses of 13C fractionations to diel variation in starch metabolism in the leaf are not fully understood. Here we measured δ13C of organic matter (δ13COM), concentrations and δ13C of potential respiratory substrates, δ13C of dark-respired CO2 (δ13CR), and gas exchange in leaves of starch-deficient plastidial phosphoglucomutase (pgm) mutants and wild-type plants of four species (Arabidopsis thaliana, Mesembryanthemum crystallinum, Nicotiana sylvestris, and Pisum sativum). The strongest δ13C response to the pgm-induced starch deficiency was observed in N. sylvestris, with more negative δ13COM, δ13CR, and δ13C values for assimilates (i.e. sugars and starch) and organic acids (i.e. malate and citrate) in pgm mutants than in wild-type plants during a diel cycle. The genotype differences in δ13C values could be largely explained by differences in leaf gas exchange. In contrast, the PGM-knockout effect on post-photosynthetic 13C fractionations via the plastidic fructose-1,6-bisphosphate aldolase reaction or during respiration was small. Taken together, our results show that the δ13C variations in starch-deficient mutants are primarily explained by photosynthetic 13C fractionations and that the combination of knockout mutants and isotope analyses allows additional insights into plant metabolism.

Green light drives photosynthesis in mosses

Temperate forests are characterised by variable light quality (i.e. spectral composition of light) at or near the forest floor. These understory environments have a high concentration of green light, as red and blue light are preferentially absorbed by upper canopy leaves. Understory species may be well-adapted for using green light to drive photosynthesis. Angiosperms have been shown to use green light for photosynthesis, but this ability has not been demonstrated in shade-dwelling bryophytes. In this study, net photosynthetic rate (PN) of three temperate understory species of moss (Dichodontium pellucidum (Hedw.) Schimp., Leucobryum albidum (Brid. ex P.Beauv) Lindb. and Amblystegium serpens (Hedw.) Schimp.) was measured under green, red + blue, and red + blue + green light to assess green light use efficiency. All three species were capable of photosynthesising beyond their respiratory demands using solely green light, with higher green light use efficiency measured in plants collected from areas with greater canopy cover, suggesting growth in a green light concentrated environment increases green light use efficiency. Each species was also collected from sites differing in their degree of canopy cover and grown under three light treatments (high light, low light, and green light). Photosynthetic efficiency (chlorophyll fluorescence), tissue nitrogen and carbon isotope concentrations were assessed after a short growth period. Growth conditions had little effect on leaf chemistry and monochromatic green light did not significantly degrade photosynthetic efficiency. This study provides the first evidence to date of positive net ‘green light photosynthesis’ in mosses.

Climate and soils together regulate photosynthetic carbon isotope discrimination within C3 plants worldwide

AbstractAimWithin C3 plants, photosynthesis is a balance between CO2 supply from the atmosphere via stomata and demand by enzymes within chloroplasts. This process is dynamic and a complex but crucial aspect of photosynthesis. We sought to understand the spatial pattern in CO2 supply–demand balance on a global scale, via analysis of stable isotopes of carbon within leaves (Δ13C), which provide an integrative record of CO2 drawdown during photosynthesis.LocationGlobalTime period1951–2011.Major taxa studiedVascular plants.MethodsWe assembled a database of leaf carbon isotope ratios containing 3,979 species–site combinations from across the globe, including 3,645 for C3 species. We examined a wide array of potential climate and soil drivers of variation in Δ13C.ResultsThe strongest drivers of carbon isotope discrimination at the global scale included atmospheric pressure, potential evapotranspiration and soil pH, which explained 44% of the variation in Δ13C. Addition of eight more climate and soil variables (each explaining small but highly significant amounts of variation) increased the explained variation to 60%. On top of this, the largest plant trait effect was leaf nitrogen per area, which explained 11% of Δ13C variation.Main conclusions: By considering variation in Δ13C at a considerably larger scale than previously, we were able to identify and quantify key drivers in CO2 supply–demand balance previously unacknowledged. Of special note is the key role of soil properties, with greater discrimination on low‐pH and high‐silt soils. Unlike other plant traits, which show typically wide variation within sets of coexisting species, the global pattern in carbon stable isotope ratios is much more conservative; there is relatively narrow variation in time‐integrated CO2 concentrations at the site of carboxylation among plants in a given soil and climate.

Profligate and conservative: water use strategies in grain legumes

Yields of grain legumes are constrained by available water. Thus, it is crucial to understand traits influencing water uptake and the efficiency of using water to produce biomass. Global comparisons and comparisons at specific locations reveal that water use of different grain legumes is very similar, which indicates that water use efficiency varies over a wide range due to differences in biomass and yield. Moreover, yield increases more per millimetre of water used in cool season grain legumes than warm season species. Although greater contrasts have been observed across species and genotypes at the pot and lysimeter level, agronomic factors need to be taken into account when scaling those studies to field-level responses. Conservative water use strategies in grain legumes such as low stomatal conductance as approximated by low photosynthetic carbon isotope discrimination reduces yield potential, whereas temporal adjustments of stomatal conductance within the growing season and in response to environmental factors (such as vapour pressure deficit) helps to optimize the trade-off between carbon gain and water loss. Furthermore, improved photosynthetic capacity, reduced mesophyll conductance, reduced boundary layer, and re-fixation of respired CO2 were identified as traits that are beneficial without water deficit, but also under terminal and transient drought. Genotypic variability in some grain legume species has been observed for several traits that influence water use, water use efficiency, and yield, including root length and the temporal pattern of water use, but even more variation is expected from wild relatives. Albeit that N2 fixation decreases under drought, its impact on water use is still largely unknown, but the nitrogen source influences gas exchange and, thus, transpiration efficiency. This review concludes that conservative traits are needed under conditions of terminal drought to help maintain soil moisture until the pod-filling period, but profligate traits, if tightly regulated, are important under conditions of transient drought in order to profit from short intermittent periods of available soil moisture.

CO2-dependent carbon isotope fractionation in the dinoflagellate Alexandrium tamarense

The carbon isotopic composition of marine sedimentary organic matter is used to resolve long-term histories of pCO2 based on studies indicating a CO2-dependence of photosynthetic carbon isotope fractionation (εP). It recently was proposed that the δ13C values of dinoflagellates, as recorded in fossil dinocysts, might be used as a proxy for pCO2. However, significant questions remain regarding carbon isotope fractionation in dinoflagellates and how such fractionation may impact sedimentary records throughout the Phanerozoic. Here we investigate εP as a function of CO2 concentration and growth rate in the dinoflagellate Alexandrium tamarense. Experiments were conducted in nitrate-limited chemostat cultures. Values of εP were measured on cells having growth rates (μ) of 0.14–0.35d−1 and aqueous carbon dioxide concentrations of 10.2–63μmolkg−1 and were found to correlate linearly with μ/[CO2(aq)] (r2=0.94) in accord with prior, analogous chemostat investigations with eukaryotic phytoplankton. A maximum fractionation (εf) value of 27‰ was characterized from the intercept of the experiments, representing the first value of εf determined for an algal species employing Form II RubisCO—a structurally and catalytically distinct form of the carbon-fixing enzyme. This value is larger than theoretical predictions for Form II RubisCO and not significantly different from the ∼25‰ εf values observed for taxa employing Form ID RubisCO. We also measured the carbon isotope contents of dinosterol, hexadecanoic acid, and phytol from each experiment, finding that each class of biomarker exhibits different isotopic behavior. The apparent CO2-dependence of εP values in our experiments strengthens the proposal to use dinocyst δ13C values as a pCO2 proxy. Moreover, the similarity between the εf value for A. tamarense and the consensus value of ∼25‰ indicates that the CO2-sensitivity of carbon isotope fractionation saturates at similar CO2 levels across all three ecologically prominent clades of eukaryotic phytoplankton. This continuity of εf across taxa may help to explain why there is no coherent signature of phytoplankton evolutionary succession in Phanerozoic carbon isotope records.

Evaluation of photosynthetic performance and carbon isotope discrimination in perennial ryegrass (Lolium perenne L.) under allelochemicals stress

Ferulic (FA) and p-hydroxybenzoic acid (pHBA) are commonly found as phenolic compounds (PHC) in many forage and cereal crops. Although the effects of these PHC on seedling growth are relatively explored, not many information is available regarding the phytotoxicity on ecophysiological processes of perennial ryegrass adult plants. The experiment was conducted with the aim to evaluate the phytotoxic potential of PHC on the seedling growth, leaf water relation, chlorophyll fluorescence attributes and carbon isotope discrimination adult plants of perennial ryegrass (Lolium perenne L.). The results clearly indicated that PHC behaved as potent inhibitors of chlorophyll fluorescence yield (Fv/Fm) in leaves of L. perenne and plants showed poor tolerance against allelochemicals stress. Quantum yield (ΦPSII), chlorophyll fluorescence quenching (qP) and non-photochemical quenching (NPQ) were decreased following exposure to FA and pHBA. The portion of absorbed photon energy that was thermally dissipated (D) in L. perenne was decreased. Exposure of the L. perenne seedlings to FA and pHBA stress led to a decrease in fresh/dry weight, relative water content and leaf osmotic potential. Carbon isotope composition ratio (δ13C) was significantly less negative than the control following treatment with FA or pHBA. The results suggested that PHC uptake was a key step for the effectiveness of these secondary metabolites and their phytotoxicity on L. perenne adult plants was mainly due to the alteration of leaf water status accompanied by photosystem II damage. Acquisition of such knowledge may ultimately provide a better understanding about the mode of action of the tested compounds.

Sex-specific differences in functional traits and resource acquisition in five cycad species.

Selective pressures acting on plant life histories can drive extreme specialization. One example of such specialization is the evolution of dioecious breeding systems. Evolutionary and ecological theory posits that dioecy may subject male and female individuals to different selective pressures and result in unique sex-mediated adaptive traits related to resource allocation and ecophysiology. Cycads are the earliest diverging lineage of seed plants with strict dioecy, yet we know almost nothing about the ecology and physiology of this group. Especially limited is our understanding of potential sex-specific differences and how such differences may influence species ecology. Here we examine the ecophysiology of male and female cycads to understand better, the role that dioecy plays in this group. We evaluated sex-specific differences in ecophysiological traits and resource acquisition in five species. Specifically, we compared photosynthetic physiology, nitrogen and carbon content, isotope discrimination (δ15N and δ13C), and stomatal density. In some cycads, (i) males and females have similar investments in leaf nitrogen but females exhibit greater incorporation of nitrogen from nitrogen-fixing soil bacteria, (ii) males display higher photosynthetic capacity but females show decreased [corrected] water-use efficiency, and (iii) males have higher stomatal conductance but similar stomatal density to females. This study is the first to examine the ecophysiological differences that have evolved in the oldest dioecious lineage of seed-bearing plants. Our results show unexpected differences in photosynthetic physiology and highlight the co-evolution with nitrogen fixing soil bacteria as a potential new key player in an old lineage.

Strong Coupling of Shoot Assimilation and Soil Respiration during Drought and Recovery Periods in Beech As Indicated by Natural Abundance δ13C Measurements.

Drought down-regulates above- and belowground carbon fluxes, however, the resilience of trees to drought will also depend on the speed and magnitude of recovery of these above- and belowground fluxes after re-wetting. Carbon isotope composition of above- and belowground carbon fluxes at natural abundance provides a methodological approach to study the coupling between photosynthesis and soil respiration (SR) under conditions (such as drought) that influence photosynthetic carbon isotope discrimination. In turn, the direct supply of root respiration with recent photoassimilates will impact on the carbon isotope composition of soil-respired CO2. We independently measured shoot and soil CO2 fluxes of beech saplings (Fagus sylvatica L.) and their respective δ13C continuously with laser spectroscopy at natural abundance. We quantified the speed of recovery of drought stressed trees after re-watering and traced photosynthetic carbon isotope signal in the carbon isotope composition of soil-respired CO2. Stomatal conductance responded strongly to the moderate drought (-65%), induced by reduced soil moisture content as well as increased vapor pressure deficit. Simultaneously, carbon isotope discrimination decreased by 8‰, which in turn caused a significant increase in δ13C of recent metabolites (1.5–2.5‰) and in δ13C of SR (1–1.5‰). Generally, shoot and soil CO2 fluxes and their δ13C were in alignment during drought and subsequent stress release, clearly demonstrating a permanent dependence of root respiration on recently fixed photoassimilates, rather than on older reserves. After re-watering, the drought signal persisted longer in δ13C of the water soluble fraction that integrates multiple metabolites (soluble sugars, amino acids, organic acids) than in the neutral fraction which represents most recently assimilated sugars or in the δ13C of SR. Nevertheless, full recovery of all aboveground physiological variables was reached within 4 days – and within 7 days for SR – indicating high resilience of (young) beech against moderate drought.

Stable isotopes in tree rings: towards a mechanistic understanding of isotope fractionation and mixing processes from the leaves to the wood.

The mechanistic understanding of isotope fractionation processes is increasing but we still lack detailed knowledge of the processes that determine the isotopic composition of the tree-ring archive over the long term. Especially with regard to the path from leaf photosynthate production to wood formation, post-assimilation fractionations/processes might cause at least a partial decoupling between the leaf isotope signals that record processes such as stomatal conductance, transpiration and photosynthesis, and the wood or cellulose signals that are stored in the paleophysiological record. In this review, we start from the rather well understood processes at the leaf level such as photosynthetic carbon isotope fractionation, leaf water evaporative isotope enrichment and the issue of the isotopic composition of inorganic sources (CO2 and H2O), though we focus on the less explored 'downstream' processes related to metabolism and transport. We further summarize the roles of cellulose and lignin as important chemical constituents of wood, and the processes that determine the transfer of photosynthate (sucrose) and associated isotopic signals to wood production. We cover the broad topics of post-carboxylation carbon isotope fractionation and of the exchange of organic oxygen with water within the tree. In two case studies, we assess the transfer of carbon and oxygen isotopic signals from leaves to tree rings. Finally we address the issue of different temporal scales and link isotope fractionation at the shorter time scale for processes in the leaf to the isotopic ratio as recorded across longer time scales of the tree-ring archive.

Advances in measurements and models of photosynthetic carbon isotope discrimination in C3 plants

Advances in measurements and models of photosynthetic carbon isotope discrimination in <scp>C</scp>₃ plants