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Wheat plants (Triticum durum Desf., cv. Regallo) were grown in the field to study the effects of contrasting [CO(2)] conditions (700 versus 370 μmol mol(-1)) on growth, photosynthetic performance, and C management during the post-anthesis period. The aim was to test whether a restricted capacity of sink organs to utilize photosynthates drives a loss of photosynthetic capacity in elevated CO(2). The ambient (13)C/(12)C isotopic composition (δ(13)C) of air CO(2) was changed from -10.2‰ in ambient [CO(2)] to -23.6‰ under elevated [CO(2)] between the 7th and the 14th days after anthesis in order to study C assimilation and partitioning between leaves and ears. Elevated [CO(2)] had no significant effect on biomass production and grain filling, and caused an accumulation of C compounds in leaves. This was accompanied by up-regulation of phosphoglycerate mutase and ATP synthase protein content, together with down-regulation of adenosine diphosphate glucose pyrophosphatase protein. Growth in elevated [CO(2)] negatively affected Rubisco and Rubisco activase protein content and induced photosynthetic down-regulation. CO(2) enrichment caused a specific decrease in Rubisco content, together with decreases in the amino acid and total N content of leaves. The C labelling revealed that in flag leaves, part of the C fixed during grain filling was stored as starch and structural C compounds whereas the rest of the labelled C (mainly in the form of soluble sugars) was completely respired 48 h after the end of labelling. Although labelled C was not detected in the δ(13)C of ear total organic matter and respired CO(2), soluble sugar δ(13)C revealed that a small amount of labelled C reached the ear. The (12)CO(2) labelling suggests that during the beginning of post-anthesis the ear did not contribute towards overcoming flag leaf carbohydrate accumulation, and this had a consequent effect on protein expression and photosynthetic acclimation.

Despite the relevance of carbon (C) loss through respiration processes (with its consequent effect on the lower C availability for grain filling), little attention has been given to this topic. Literature data concerning the role of respiration in cereals are scarce and these have been produced using indirect methods based on gas-exchange estimations. We have developed a new method based on the capture of respired CO(2) samples and their analysis by gas chromatography-combustion-isotope ratio mass spectrometry (GC-C-IRMS). In order to analyse the main processes involved in the C balance during grain filling (photosynthesis, respiration, allocation and partitioning) the ambient isotopic (13)C/(12)C composition (delta(13)C) of the growth chamber was modified during this period (delta(13)C ca. -12.8 +/- 0.3 per thousand to ca. -20.0 +/- 0.2 per thousand). The physiological performance, together with the C allocation on total organic matter (TOM) and respiration of wheat (Triticum aestivum L., var. Califa sur) and two hybrids, tritordeum (X Tritordeum Asch. & Graebn line HT 621) and triticale (X Triticosecale Wittmack var. Imperioso), were compared during post-anthesis water stress. In spite of the larger ear DM/total ratio, especially under drought conditions, the grain filling of triticale and wheat was mainly carried out with pre-anthesis C, while the majority of C assimilated during post-anthesis was invested in respiration processes. In the case of wheat and tritordeum, the C balance data suggested a reallocation during grain filling of photoassimilates stored prior to anthesis from shoot to ear. Furthermore, the lower percentage of labeled C on respired CO(2) of droughted tritordeum plants, together with the lower plant biomass, explained the fact that those plants had more C available for grain filling., (Copyright (c) 2009 John Wiley & Sons, Ltd.)

Starch and soluble sugars are the major photosynthetic products, and their carbon isotope signatures reflect external versus internal limitations of CO(2) fixation. There has been recent renewed interest in the isotope composition of carbohydrates, mainly for use in CO(2) flux partitioning studies at the ecosystem level. The major obstacle to the use of carbohydrates in such studies has been the lack of an acknowledged method to isolate starch and soluble sugars for isotopic measurements. We here report on the comparison and evaluation of existing methods (acid and enzymatic hydrolysis for starch; ion-exchange purification and compound-specific analysis for sugars). The selectivity and reproducibility of the methods were tested using three approaches: (i) an artificial leaf composed of a mixture of isotopically defined compounds, (ii) a C(4) leaf spiked with C(3) starch, and (iii) two natural plant samples (root, leaf). Starch preparation methods based on enzymatic or acid hydrolysis did not yield similar results and exhibited contaminations by non-starch compounds. The specificity of the acidic hydrolysis method was especially low, and we therefore suggest terming these preparations as HCl-hydrolysable carbon, rather than starch. Despite being more specific, enzyme-based methods to isolate starch also need to be further optimized to increase specificity. The analysis of sugars by ion-exchange methods (bulk preparations) was fast but produced more variable isotope compositions than compound-specific methods. Compound-specific approaches did not in all cases correctly reproduce the target values, mainly due to unsatisfactory separation of sugars and background contamination. Our study demonstrates that, despite their wide application, methods for the preparation of starch and soluble sugars for the analysis of carbon isotope composition are not (yet) reliable enough to be routinely applied and further research is urgently needed to resolve the identified problems., (2009 John Wiley & Sons, Ltd.)

Whereas the effects of water and nitrogen (N) on plant Delta(13)C have been reported previously, these factors have scarcely been studied for Delta(18)O. Here the combined effect of different water and N regimes on Delta(13)C, Delta(18)O, gas exchange, water-use efficiency (WUE), and growth of four genotypes of durum wheat [Triticum turgidum L. ssp. durum (Desf.) Husn.] cultured in pots was studied. Water and N supply significantly increased plant growth. However, a reduction in water supply did not lead to a significant decrease in gas exchange parameters, and consequently Delta(13)C was only slightly modified by water input. Conversely, N fertilizer significantly decreased Delta(13)C. On the other hand, water supply decreased Delta(18)O values, whereas N did not affect this parameter. Delta(18)O variation was mainly determined by the amount of transpired water throughout plant growth (T(cum)), whereas Delta(13)C variation was explained in part by a combination of leaf N and stomatal conductance (g(s)). Even though the four genotypes showed significant differences in cumulative transpiration rates and biomass, this was not translated into significant differences in Delta(18)O(s). However, genotypic differences in Delta(13)C were observed. Moreover, approximately 80% of the variation in biomass across growing conditions and genotypes was explained by a combination of both isotopes, with Delta(18)O alone accounting for approximately 50%. This illustrates the usefulness of combining Delta(18)O and Delta(13)C in order to assess differences in plant growth and total transpiration, and also to provide a time-integrated record of the photosynthetic and evaporative performance of the plant during the course of crop growth.

Intense efforts are currently devoted to disentangling the relationships between plant carbon (C) allocation patterns and soil nitrogen (N) availability because of their consequences for growth and more generally for C sequestration. In cold ecosystems, only a few studies have addressed whole-plant C and/or N allocation along natural elevational or topographical gradients. (12)C/(13)C and (14)N/(15)N isotope techniques have been used to elucidate C and N partitioning in two alpine graminoids characterized by contrasted nutrient economies: a slow-growing species, Kobresia myosuroides (KM), and a fast-growing species, Carex foetida (CF), located in early and late snowmelt habitats, respectively, within the alpine tundra (French Alps). CF allocated higher labelling-related (13)C content belowground and produced more root biomass. Furthermore, assimilates transferred to the roots were preferentially used for growth rather than respiration and tended to favour N reduction in this compartment. Accordingly, this species had higher (15)N uptake efficiency than KM and a higher translocation of reduced (15)N to aboveground organs. These results suggest that at the whole-plant level, there is a compromise between N acquisition/reduction and C allocation patterns for optimized growth.

Most of the literature focused on internal CO(2) (Ci) determinations in plants has used indirect methods based on gas-exchange estimations. We have developed a new method based on the capture of internal air gas samples and their analysis by gas chromatography-combustion-isotope ratio mass spectrometry (GC-C-IRMS). This method provided a direct measure of intercellular CO(2) concentrations combined with stable carbon isotopic composition in O. ficus-indica plants. Plants were grown at both ambient and elevated CO(2) concentration. During the day period, when the stomata are closed, the Ci was high and was very (13)C-enriched in both ambient and elevated CO(2)-grown plants, reflecting Rubisco's fractionation (this plant enzyme has been shown to discriminate by 29 per thousand, in vitro, against (13)CO(2)). Other enzyme fractionations involved in C metabolism in plants, such as carbonic anhydrase, could also be playing an important role in the diurnal delta(13)C enrichment of the Ci. During the night, when stomata are open, Ci concentrations were higher in elevated (and the corresponding delta(13)C values were more (13)C-depleted) than in ambient CO(2)-grown plants., ((c) 2008 John Wiley & Sons, Ltd.)

Background and aims: The tracing of C assimilation and the subsequent partitioning among plant organs has been a central focus of studies utilising Free Air CO2 Enrichment (FACE) facilities. The approach makes use of the fossil origin of this carbon, which is depleted in 13C. However, there is little data for desert environments. The Nevada Desert FACE Facility (NDFF), located in the Mojave Desert, has been one of the main facilities for the study of C dynamics in arid ecosystems and how they respond to rising atmospheric CO2 concentrations. In this experiment, we studied the incorporation of fixed CO2 during the previous two years (detectable by its lower δ13C) in the soil fraction surrounding roots. Methods: The soil was collected monthly in direct vicinity to the roots during a complete growth season, at two depths (5 and 15 cm). Soil samples were dried and fractionated by size (> 50 μm and < 50 μm) by wet sieving, and both size fractions were then analysed for the δ13C of their organic matter and their carbonates. Results: In the coarse fraction (> 50 μm), δ13C values ranged between −1 and − 2‰ for carbonates and between −23 and − 25‰ for soil organic matter. These values did not significantly change throughout the experiment and were not affected by depth (5 or 15 cm). In contrast, δ13C values for both organic and inorganic carbon in the fine fraction (< 50 μm) were much more variable than in the coarse fraction (> 50 μm). The δ13C values for organic C ranged mostly between −20‰ and − 27‰, and were roughly maintained throughout the sampling period. For inorganic C, the δ13C values were mostly between 0‰ and − 15‰, and tended to become less negative during the course of the sampling period. Overall the effect of [CO2] on δ13C values of either organic or inorganic carbon was not significant for any experimental condition (plant species, soil depth, soil fraction). Conclusion: Little or no signs of recently fixed CO2 (13C-depleted) were detected in the soils close to the roots, in the coarse fraction (> 50 μm), the fine fraction (< 50 μm), the organic matter, or in carbonates. This indicates a slow C turnover in the studied soils, which can result from a highly conservative use of photoassimilates by plants, including a very low release of organic matter into the soil in the form of dead roots or root exudates, and from a conservative use of available C reserves. [ABSTRACT FROM AUTHOR]

Legumes such as alfalfa (Medicago sativa L.) are vital N[sub 2]-fixing crops accounting for a global N[sub 2] fixation of ~35 Mt N year[sup -1]. Although enzymatic and molecular mechanisms of nodule N[sub 2] fixation are now well documented, some uncertainty remains as to whether N[sub 2] fixation is strictly coupled with photosynthetic carbon fixation. That is, the metabolic origin and redistribution of carbon skeletons used to incorporate nitrogen are still relatively undefined. Here, we conducted isotopic labelling with both [sup 15]N[sub 2] and [sup 13]C-depleted CO[sub 2] on alfalfa plants grown under controlled conditions and took advantage of isotope ratio mass spectrometry to investigate the relationship between carbon and nitrogen turn-over in respired CO[sub 2], total organic matter and amino acids. Our results indicate that CO[sub 2] evolved by respiration had an isotopic composition similar to that in organic matter regardless of the organ considered, suggesting that the turn-over of respiratory pools strictly followed photosynthetic input. However, carbon turn-over was nearly three times greater than N turn-over in total organic matter, suggesting that new organic material synthesised was less N-rich than pre-existing organic material (due to progressive nitrogen elemental dilution) or that N remobilisation occurred to sustain growth. This pattern was not consistent with the total commitment into free amino acids where the input of new C and N appeared to be stoichiometric. The labelling pattern in Asn was complex, with contrasted C and N commitments in different organs, suggesting that neosynthesis and redistribution of new Asn molecules required metabolic remobilisation. We conclude that the production of new organic material during alfalfa growth depends on both C and N remobilisation in different organs. At the plant level, this remobilisation is complicated by allocation and metabolism in the different organs. The combination of double C[sup 13] and N[sup 15] labelling and subsequent compound-specific analysis in amino acids showed the imbalance observed between CO[sub 2] and N[sub 2] fixation in nodulated alfalfa plants. It provided a novel conclusion about the use and importance of pre-existing reserves in the production of new organic matter during growth. [ABSTRACT FROM AUTHOR]

The combined use of stable carbon and oxygen isotopes in plant matter is a tool of growing interest in cereal crop management and breeding, owing to its relevance for assessing the photosynthetic and transpirative performance under different growing conditions including water and N regimes. However, this method has not been applied to wheat grown under real field conditions. Here, plant growth, grain yield (GY) and the associated agronomic components, carbon isotope discrimination (ΔC) plus oxygen isotope composition ( δO) as well as leaf and canopy gas exchange were measured in field-grown wheat subjected to different water and N availabilities. Water limitation was the main factor affecting yield, leaf and canopy gas exchange and ΔC and δO, whereas N had a smaller effect on such traits. The combination of ΔC and δO gave a clear advantage compared with gas exchange measurements, as it provides information on the instantaneous and the long-term plant photosynthetic and transpirative performance and are less labour intensive than gas exchange measurements. In addition, the combination of plant ΔC and δO predicted differences in GY and related agronomical parameters, providing agronomists and breeders with integrative traits for selecting crop management practices and/or genotypes with better performance under water-limiting and N-limiting conditions. [ABSTRACT FROM AUTHOR]

Two slow-growing plant species ( Chamaerops humilis, L. and Cycas revoluta Thunb.) were exposed to elevated CO2 conditions over a 20-month period in order to study the CO2 effect on growth, photosynthetic capacity and leaf carbon (C) management. The ambient isotopic 13C/12C composition ( δ13C) of the greenhouse module corresponding to elevated CO2 (800 μmol mol−1 CO2) conditions was changed from δ13C ca. −12.8±0.3‰ to ca. −19.2±0.2‰. Exposure of these plants to elevated CO2 enhanced dry mass (DM) by 82% and 152% in Chamerops and Cycas, respectively, mainly as a consequence of increases in plant level photosynthetic rates. However, analyses of A– Ci curve parameters revealed that elevated CO2 diminished leaf photosynthetic rates of Chamaerops whereas in Cycas, no photosynthetic acclimation was detected. The fact that Chamaerops plants had a lower DM increase, together with a longer leaf C residence time and a diminished capacity to respire recently fixed C, suggests that this species was unable to increase C sink strength. Furthermore, the consequent C source/sink imbalance in Chamaerops might have induced the downregulation of Rubisco. Cycas plants were capable of avoiding photosynthetic downregulation due to a greater ability to increase C sink strength, as was confirmed by DM values, and 12C-enriched CO2 labeling data. Cycas developed the ability to respire a larger proportion of recently fixed C and to reallocate the recently fixed C away from leaves to other plant tissues. These findings suggest that leaf C management is a key factor in the responsiveness of slow-growing plants to future CO2 scenarios. [ABSTRACT FROM AUTHOR]

Investigates the carbon isotope composition of carbon dioxide (CO[sub 2]) produced in darkness by intact French bean. Decrease of the isotope composition as temperature increases; Enrichment of the evolved CO [sub 2] in carbon isotope; Contribution of two major decarboxylation processes that occur in darkness to the determination of carbon isotope composition of dark inspired CO[sub 2].

Two slow-growing plant species ( Chamaerops humilis, L. and Cycas revoluta Thunb.) were exposed to elevated CO2 conditions over a 20-month period in order to study the CO2 effect on growth, photosynthetic capacity and leaf carbon (C) management. The ambient isotopic 13C/12C composition ( δ13C) of the greenhouse module corresponding to elevated CO2 (800 μmol mol−1 CO2) conditions was changed from δ13C ca. −12.8±0.3‰ to ca. −19.2±0.2‰. Exposure of these plants to elevated CO2 enhanced dry mass (DM) by 82% and 152% in Chamerops and Cycas, respectively, mainly as a consequence of increases in plant level photosynthetic rates. However, analyses of A– Ci curve parameters revealed that elevated CO2 diminished leaf photosynthetic rates of Chamaerops whereas in Cycas, no photosynthetic acclimation was detected. The fact that Chamaerops plants had a lower DM increase, together with a longer leaf C residence time and a diminished capacity to respire recently fixed C, suggests that this species was unable to increase C sink strength. Furthermore, the consequent C source/sink imbalance in Chamaerops might have induced the downregulation of Rubisco. Cycas plants were capable of avoiding photosynthetic downregulation due to a greater ability to increase C sink strength, as was confirmed by DM values, and 12C-enriched CO2 labeling data. Cycas developed the ability to respire a larger proportion of recently fixed C and to reallocate the recently fixed C away from leaves to other plant tissues. These findings suggest that leaf C management is a key factor in the responsiveness of slow-growing plants to future CO2 scenarios. [ABSTRACT FROM AUTHOR]