The influence of water content and leaf anatomy on carbon isotope discrimination and photosynthesis in Sphagnum

The relative effect of diffusional resistance due to water films (r_wf) and leaf anatomy (r_p) on rates of net photosynthesis and on-line measures of carbon isotope discrimination (Δ=Δδ¹³C) was investigated in Sphagnum. Sphagnum species differ in the exposure of photosynthetic cells at the leaf surface. In S. affine, photosynthetic cells are widely exposed at the surface, whereas in S. magellanicum, photo-synthetic cells are enclosed within water-filled hyaline cells. This difference is expected to lead to variation in diffusive resistance within leaves (r_p). Net photosynthesis and on-line Δ were measured at two water contents: greenhouse water content (wet) and blotted dry (dry). Without correcting for respiration, on-line Δ values differed significantly between wet (23.7%_o) and dry (30.9%_o) plants. However, there was no significant difference between species means and no species × water content interaction. Corrections for respiration lowered Δ values by approximately 8.1%_o and reduced the mean difference to 3.1%_o, but did not alter the rank order of treatments. Net photosynthesis also decreased by 16% in wet plants, but there was no significant difference between the two species. In addition, five populations of S. affine and S. magellanicum grown in a common garden were analysed for their organic matter carbon isotope composition (δ¹³C). These values varied more within each species (0.9–1.2%_o) than between the two species (0.6%_o). Therefore, we conclude that variation in surface water films leads to a greater difference in resistance to CO₂ uptake and carbon isotope discrimination than that due to variation in leaf anatomical properties in Sphagnum.     

Short-term changes in leaf carbon isotope discrimination in salt- and water-stressed c(4) grasses

Online carbon isotope discrimination (Δ) and leaf gas exchange measurements were made with control and salt-stressed Zea mays and Andropogon glomeratus, two NADP-ME type C₄ grasses. Linear relationships between Δ and p_i/p_a (the ratio of intercellular to atmospheric CO₂ partial pressure) were found for control plants which agreed well with theoretical models describing carbon isotope discrimination in C₄ plants. These data provided estimates of , the proportion of CO₂ fixed by phosphoenolpyruvate carboxylase which leaks out of the bundle sheath and the component of fractionation due to diffusion in air. Salt-stressed plants had wider variation in Δ for the same or less range in p_i/p_a. Additional work indicated Δ changed independently of p_i/p_a in both water- and salt-stressed plants, suggesting a possible diurnal change in as plant water status changed linked to a decrease in the activity of the C₃ photosynthetic pathway relative to C₄ pathway activity. The possible effect of stress-induced changes in on organic matter δ¹³ C of C₄ plants is apt to be most apparent in chronically stressed environments.     

Effect of leaf structure and water status on carbon isotope discrimination in field-grown durum wheat

The relationships between leaf and kernel carbon isotope discriminations (Δ) and several leaf structural parameters that are indicators of photosynthetic capacity were studied in durum wheat grown in the field under three water regimens. A set of 144 genotypes were cultivated in two rain-fed trials, and 125 of these were grown under supplementary irrigation before heading. Total chlorophyll and nitrogen (N) contents, the dry mass per unit leaf area (LDM, the reciprocal of specific leaf area) and carbon isotope discrimination (Δ) were measured in penultimate leaves and Δ of mature kernels was also analysed. Both LDM and N per unit area showed significant (P≤ 0.001) negative correlation (r=–0.60 and r=–0.36, respectively) with leaf Δ in the wettest trial. Little or no correlation was found for any structural parameter and leaf Δ in the rain-fed trials. In contrast, in the two rain-fed trials LDM was the parameter with the strongest positive correlation (P≤ 0.001) with kernel Δ (r= 0.47 and 0.30) and grain yield (r= 0.43 and 0.29), whereas no correlation was found in the irrigation trial. These correlations, rather than representing a causal link between the amount of photosynthetic tissue and Δ, were probably indirect associations caused by a parallel effect of water status and phenology on leaf structure, grain Δ and yield. Correlations across trials (i.e. environments) between leaf structure and either Δ and yield were very high, although also spurious. Our results suggest that LDM should be used to cull segregating population differences in leaf Δ based on the internal photosynthetic capacity only in the absence of drought. Selecting for kernel Δ and grain yield on the basis of LDM is worthwhile for rain-fed trials.     

Early senescence induced by 2-3H-benzoxazolinone (BOA) in Arabidopsis thaliana

Measurements of chlorophyll a fluorescence, nutrient and trace elements, total protein content and malonyldialdehyde in leaves of Arabidopsis thaliana between 1 and 192 h after treatment with 0, 1 or 3 mM 2-3H-benzoxazolinone (BOA), together with imaging of chlorophyll a fluorescence and of the distributions of hydrogen peroxide and superoxide anion, suggested that the primary phytotoxic action of BOA is the induction of premature senescence, and that oxidative stress is a secondary effect that sets in a day or two later.     

Pioneer and late stage tropical rainforest tree species (French Guiana) growing under common conditions differ in leaf gas exchange regulation,carbon isotope discrimination and leaf water potential

Leaf gas exchange rates, predawn _wp and daily minimum _wm leaf water potentials were measured during a wet-to-dry season transition in pioneer (Jacaranda copaia, Goupia glabra andCarapa guianensis) and late stage rainforest tree species (Dicorynia guianensis andEperua falcata) growing in common conditions in artificial stands in French Guiana. Carbon isotope discrimination () was assessed by measuring the stable carbon isotope composition of the cellulose fraction of wood cores. The values were 2.7 higher in the pioneer species than in the late stage species. The calculated time integratedC _i values derived from the values averaged 281 mol mol^–1 in the pioneers and 240 mol mol^–1 in the late stage species. The corresponding time-integrated values of intrinsinc water-use efficiency [ratio CO₂ assimilation rate (A)/leaf conductance (g)] ranged from 37 to 47 mmol mol^–1 in the pioneers and the values were 64 and 74 mmol mol^–1 for the two late stage species. The high values were associated—at least inJ. copaia—with high maximumg values and with high plant intrinsinc specific hydraulic conductance [Cg/(_wm–_wp], which could reflect a high competitive ability for water and nutrient uptake in the absence of soil drought in the pioneers. A further clear discriminating trait of the pioneer species was the very sensitive stomatal response to drought in the soil, which might be associated with a high vulnerability to cavitation in these species. From a methodological point of view, the results show the relevance of for distinguishing ecophysiological functional types among rainforest trees.     

Productivity, carbon isotope discrimination and leaf traits of trees of Eucalyptus globulus Labill. in relation to water availability

The stand basal area, carbon isotope discrimination (Δ) in tree rings and leaves, leaf area index and leaf traits of trees were measured in 6‐ to 8‐year‐old stands of Eucalyptus globulus Labill. across a gradient of rainfall of 600–1400 mm year^?1 in south‐western Australia to better understand the importance of leaf traits and gas‐exchange as determinants of stand productivity. Δ ranged from 17‰ to 21‰. Δ and basal area were highly, positively correlated with each other and the ratio of mean annual rainfall to potential evaporation (P/PE). Leaf area index, soil water holding capacity and leaf nitrogen content were only weakly correlated with basal area. Δ and P/PE were negatively correlated with leaf nitrogen content. Δ was negatively correlated with leaf density but positively correlated with specific leaf area. This is consistent with the theory that larger leaf nitrogen content and smaller specific leaf area are associated with increased photosynthetic capacity and increased leaf‐scale water‐use‐efficiency, and that Δ is influenced by mesophyll conductance. It is concluded that canopy conductance is a more important determinant of growth in water‐limited conditions than either leaf area index or leaf traits in fertilized stands of E. globulus. Water availability was dictated more by rainfall than soil type.     

Responses of photosynthesis,dry mass and carbon isotope discrimination in winter wheat to different irrigation depths

Photosynthetica - In order to test the effects of irrigation depth on winter wheat photosynthesis, four treatments were applied in a field experiment using PVC growth tubes (identical amounts of...     

Coordinated changes in photosynthesis,water relations and leaf nutritional traits of canopy trees along a precipitation gradient in lowland tropical forest

We investigated leaf physiological traits of dominant canopy trees in four lowland Panamanian forests with contrasting mean annual precipitation (1,800, 2,300, 3,100 and 3,500 mm). There was near complete turn-over of dominant canopy tree species among sites, resulting in greater dominance of evergreen species with long-lived leaves as precipitation increased. Mean structural and physiological traits changed along this gradient as predicted by cost–benefit theories of leaf life span. Nitrogen content per unit mass (N_mass) and light- and CO₂-saturated photosynthetic rates per unit mass (P_mass) of upper canopy leaves decreased with annual precipitation, and these changes were partially explained by increasing leaf thickness and decreasing specific leaf area (SLA). Comparison of 1,800 mm and 3,100 mm sites, where canopy access was available through the use of construction cranes, revealed an association among extended leaf longevity, greater structural defense, higher midday leaf water potential, and lower P_mass, N_mass, and SLA at wetter sites. Shorter leaf life spans and more enriched foliar ¹⁵N values in drier sites suggest greater resorption and re-metabolism of leaf N in drier forest. Greater dominance of short-lived leaves with relatively high P_mass in drier sites reflects a strategy to maximize photosynthesis when water is available and to minimize water loss and respiration costs during rainless periods. Overall, our study links coordinated change in leaf functional traits that affect productivity and nutrient cycling to seasonality in lowland tropical forests.     

Associations between partitioning of carboxylase activity and bundle sheath leakiness to CO2, carbon isotope discrimination, photosynthesis, and growth in sugarcane

Genetic and environmental variation in carbon isotope discrimination(     

Mono-carboxylic acids and their salts inhibit wound-induced phenolic accumulation in excised lettuce (Lactuca sativa) leaf tissue

Wounding, as during excision and preparation of lettuce ( Lactuca sativa L.) leaf tissue for salads, induces the synthesis and accumulation of phenolic compounds that participate in subsequent reactions that cause tissue browning. Exposure of excised 5-mm mid-rib segments of romaine lettuce leaf tissue to vapors of mono-carboxylic acids or aqueous solutions of mono-carboxylic acids or their salts inhibited wound-induced phenolic accumulation (WIPA) and subsequent tissue browning. The decline in phenolic content followed a quadratic curve with increasing concentration, reaching a maximum inhibition after 60 min of 74 ± 8% for 50 m M sodium acetate (2 carbons, C2) and 91 ± 4% for 20 m M sodium decanoate (capric acid, C10). Respiration (i.e. carbon dioxide production) was unaffected by concentrations of formic, acetic, or propionic acids that reduced wound-induced phenolic content or that increase ion leakage from the tissue into an isotonic mannitol solution. However, WIPA was suppressed up to 70% at concentrations (20 m M acetate) that did not increase ion leakage over that of water controls. Various acetate salts (i.e. ammonium, calcium, magnesium, and sodium) all produced the same level of inhibition. The effectiveness of the compounds increased with increasing number of carbons in the molecule from 1 to 10, and was unaffected by whether the carbons were a straight chain or branched or whether the treatment was delayed by up to 6 h. The effectiveness of butyrate (C4) in reducing WIPA (27% reduction at 20 m M ) was less than that predicted from the response of the two adjacent mono-carboxylates similarly applied: propionate (C3) (62%) and valerate (C5) (73%). It appears that, unlike the n-alcohols, mono-carboxylates are not interfering with the synthesis or propagation of a wound signal but are interfering with subsequent steps in the production and accumulation of wound-induced phenolic compounds.     

Stomatal density of cowpea correlates with carbon isotope discrimination in different phosphorus, water and CO2 environments

* Stomatal formation is affected by a plant's external environment, with long-distance signaling from mature to young leaves seemingly involved. However, it is still unclear what is responsible for this signal. To address this question, the relationship between carbon isotope discrimination (Delta) and stomatal density was examined in cowpea (Vigna sinensis). * Plants were grown under various environments that combined different amounts of soil phosphorus (P), soil water, and atmospheric CO(2). At harvest, stomatal density was measured in the youngest fully expanded leaf. The (13)C : (12)C ratio was measured in a young leaf to determine the Delta in mature leaves. * Results indicated that stomatal density is affected by P as well as by amounts of water and CO(2). However, stomatal responses to water and CO(2) were complex because of strong interactions with P. This suggests that the responses are relative, depending on some internal factor being affected by each external variable. Despite such complicated responses, a linear correlation was found between stomatal density and Delta across all environments examined. * It is proposed that the Delta value is a good surrogate for the long-term mean of the intercellular (C(i)) to the atmospheric (C(a)) CO(2) concentration ratio (C(i) : C(a)) and may be useful in understanding stomatal formation beyond complicated interactions.     

Productivity, leaf traits and carbon isotope discrimination in 29 Populus deltoides x P. nigra clones

Here we tested whether some leaf traits could be used as predictors for productivity in a range of Populus deltoides x P. nigra clones. These traits were assessed in 3-yr-old rooted cuttings from 29 clones growing in an open field trial, in a five randomized complete block design, under optimal irrigation. Variables were assigned to four groups describing productivity (above-ground biomass, total leaf area), leaf growth (total number of leaves increment rate), leaf structure (area of the largest leaf, specific leaf area, carbon and nitrogen contents), and carbon isotope discrimination in the leaves (Delta). High-yielding clones displayed larger total leaf area and individual leaf area, while no correlation could be detected between productivity and either leaf structure or Delta. By contrast, Delta was negatively correlated with number of leaves increment rate and leaf N content. Our study shows that there is a potential to improve water-use efficiency in poplar without necessarily reducing the overall productivity.     

Variation and inheritance of carbon isotope discrimination in wheat (Triticum aestivum)

Significant small variations exist in the proportion of the heavy isotope (¹³C) in organic and inorganic materials. The amounts of variations depend on the discrimination against or (in favor of) the heavy isotope through equilibrium reactions or kinetic processes particularly during CO₂ fixation in plants. These variations have also paved the way to measure carbon isotope discrimination in plant materials as a surrogate for water-use efficiency. Thus looking for variation and understanding the inheritance of this character is a prerequisite for improving wheat genotypes showing high water use efficiency and tolerance to drought. The main objectives of this review is to provide an overview of various aspects in which inheritance of ? is documented and of variation for carbon isotope discrimination reported in wheat under different circumstances. The relationship between carbon isotope discrimination and drought tolerance is also discussed exclusively in wheat.     

Variation in leaf carbon isotope discrimination in Encelia farinosa: implications for growth,competition, and drought survival

Population-level variation in the leaf carbon isotope discrimination () values was examined in Encelia farinosa, a common Sonoran Desert shrub. There was approximately a 2 range in values among different plants. These differences in values among neighboring plants were maintained through time, both under conditions when neighbors were present and after neighbors had been removed. Individuals with high values were found to have an accelerated growth rate when these plants were released from competition for water. Individuals with low values were better able to persist through long-term drought. These data suggest possible tradeoffs between conditions favoring high- and low--value plants within a natural population. Given the temporal variability in precipitation between years and spatial variability in microhabitat quality in the Sonoran Desert, variation in values among E. farinosa plants will be maintained within a population.     

The effect of transient and continuous drought on yield, photosynthesis and carbon isotope discrimination in sugar beet (Beta vulgaris L.)

Stable carbon isotope discrimination (delta13C), photosynthetic performance (A), dry matter accumulation (DW), and sucrose yield (Y(s)) of sugar beet were evaluated in a glasshouse experiment under transient (TS) and permanent (PS) water stress. A was significantly reduced under drought, to an extent depending on stress duration. The reduced A was strictly associated with a low DW and Y(s), the later being 42% lower in PS than control plants (C). Restoring water steeply increased A and the associated leaf traits (RWC, leaf water potential etc.), but the increase of Y(s) was negligible. Therefore, the negative effects of severe water stress in the early growth period, though reversible on gas-exchange and most leaf traits, can drastically reduce Y(s) of sugar beet. Furthermore, A seems not to be effective in predicting sucrose accumulation, although it was very effective in detecting the occurrence of plant water stress. The A/C(i) model was used to assess the photosynthetic adjustments to continuous or transient drought by calculating the photosynthetic parameters Vcmax and Jmax and then compared with delta13C. Mesophyll conductance (g(m)) was estimated by comparing delta13C measured on soluble sugars and gas-exchange data. This approach confirmed the expectation that g(m) was limiting A and that there was a significant drop in [CO2] from the substomatal cavities and the chloroplast stroma both in favourable and drought conditions. Therefore, the carbon concentration at the carboxylation site was overestimated by 25-35% by conventional gas-exchange measurements, and Vcmax was consistently underestimated when g(m) was not taken into account, especially under severe drought. Root delta13C was found to be strictly related to sucrose content (brix%), Y(s) and root dry weight, and this was especially clear when delta13C was measured on bulk dry matter. By contrast, leaf delta13C measured in soluble sugars (delta(s)) and bulk dry matter (delta(dm)) were found to correlate weakly to brix% and yield, and this was not surprising as the integration time-scale of leaf delta(s) and delta(dm) were found to be shorter than that of root delta13C in bulk dry matter. The effect of water stress on diffusive and biochemical limitations with different integration times ranged from 1 d (leaf delta(s)) to more than 1 month (root delta(dm)).     

Development of 2-aryl substituted quinazolin-4(3H)-one,pyrido[4,3-d]pyrimidin-4(3H)-one and pyrido[2,3-d]pyrimidin-4(3H)-one derivatives as microsomal prostaglandin E2 synthase-1 inhibitors

mPGES-1 is inducible terminal synthase acting downstream of COX enzymes in arachidonic acid pathway, regulates the biosynthesis of pro-inflammatory prostaglandin PGE₂. Cardiovascular side effect of coxibs and NSAIDs, selective for COX-2 inhibition, stimulated interest in mPGES-1, a therapeutic target with potential to deliver safe and effective anti-inflammatory drugs. The synthesis and structure activity relationship of a series of compounds from 2-aryl substituted quinazolin-4(3H)-one, pyrido[4,3-d]pyrimidin-4(3H)-one and pyrido[2,3-d]pyrimidin-4(3H)-one scaffolds as mPGES-1 inhibitor are discussed. A set of analogs (28, 48, 49) were identified with <10 nM potencies in the recombinant human mPGES-1 enzyme and in the A549 cellular assays. These analogs were also found to be potent in the human whole blood assay (<400 nM). Furthermore, the representative compound 48 was shown to be selective with other prostanoid synthases and was able to effectively regulate PGE₂ biosynthesis in clinically relevant inflammatory settings, in comparison with celecoxib.     

The influence of (photo)respiration on carbon isotope discrimination in plants

The contribution which (photo)respiration makes to carbon isotope discrimination (Δ¹³C) was examined by conducting simultaneous gas exchange measurements and isotopic analysis of carbon dioxide passing over leaves of Triticum aestivum and Phaseolus vulgaris, via manipulations of the carbon isotope composition (δ¹³C) of source CO₂ during growth and measurement. Dark respiration only altered net Δ¹³C (Δ_obs) at low CO₂ assimilation, and was sensitive to source CO₂δ¹³C during measurement. Photorespiration reduced Δ_obs relative to Δ¹³C predicted from p_i/p_a (Δ_i) over the full range of CO₂ assimilation, to a greater degree under elevated oxygen partial pressure (pO₂), indicating fractionation during photorespiration (f) in T. aestivum. For P. vulgaris, Δ_obs was insensitive to elevated pO₂ at higher assimilation rates, suggesting that f was minimal. A model was developed to calculate gross discrimination (Δ_ps), independent of (photo)respiration, from which estimates of f were obtained for T. aestivum (3.3‰) and P. vulgaris (0.5‰). Because photorespiratory fractionation varies interspecifically, and influences net Δ¹³C which is directly reflected in leaf δ¹³C, consideration of (photo)respiratory fractionation is necessary when interpreting δ¹³C of leaf material, especially under conditions where (photo)respiratory CO₂ losses make a large relative contribution to total plant carbon budgets.     

Involvement of components of the phospholipid-signaling pathway in wound-induced phenylpropanoid metabolism in lettuce (Lactuca sativa) leaf tissue

In plant tissue, a wound signal is produced at the site of injury and propagates or migrates into adjacent tissue where it induces increased phenylalanine ammonia lyase (PAL, EC 4.3.1.5) activity and phenylpropanoid metabolism. We used excised mid-rib leaf tissue from Romaine lettuce (Lactuca sativa L., Longifolia) as a model system to examine the involvement of components of the phospholipid-signaling pathway in wound-induced phenolic metabolism. Exposure to 1-butanol vapors or solutions inhibited wound-induced increase in PAL activity and phenolic metabolism. Phospholipases D (EC 3.1.4.4), an enzyme involved in the phospholipid-signaling pathway is specifically inhibited by 1-butanol. Re-wounding tissue, in which an effective 1-butanol concentration had declined below active levels by evaporation, did not elicit the normal wound response. It appears the 1-butanol-treated tissue developed resistance to wound-induced increases in phenylpropanoid metabolism that persisted even when active levels of 1-butanol were no longer present. However, a metabolic product of 1-butanol, rather than 1-butanol itself, may be the active compound eliciting persistence resistance. Inhibiting a subsequent enzyme in the phospholipid-signaling pathway, lipoxygenase (LOX; EC 1.13.11.12) with 1-phenyl-3-pyrazolidinone (1P3P) or reducing the product of LOX activity with diethyldithio-carbamic acid (DIECA) also inhibited wound-induced PAL activity and phenolic accumulation. The effectiveness of 1-butanol, DIECA, and 1P3P declined as the beginning of the 1-h immersion period was delayed from 0 to 4 h after excision. This decline in effectiveness is consistent with involvement of the inhibitors in the production or propagation of a wound signal. The wound signal in lettuce moves into adjacent tissue at 0.5 cm h⁻¹, so delaying application would allow the signal to move into and induce the wound response in adjacent tissue before the delayed application inhibited synthesis of the signal. Salicylic acid (SA) inhibits allene oxide synthase (AOS, EC 4.2.1.92), another enzyme in the phospholipid-signaling pathway. Exposure to 1 or 10 mM SA for 60 min reduced wound-induced phenolic accumulation by 26 or 56%, respectively. However, 1 mM SA lost its effectiveness if applied 3 h after excision, while 10 mM SA remained effective even when applied 4 h after excision. At 1 mM, SA may be perturbing the wound signal through inhibition of AOS, while at 10 mM it appears to have some generally inhibitory effect on subsequent phenolic metabolism. These data further implicate the phospholipid-signaling pathway in the generation of a wound signal that induces phenolic metabolism in wounded leaf tissue.