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Herbivory is one of the most globally distributed disturbances affecting carbon (C)-cycling in trees, yet our understanding of how it alters tree C-allocation to different functions such as storage, growth or rhizodeposition is still limited. Prioritized C-allocation to storage replenishment vs growth could explain the fast recovery of C-storage pools frequently observed in growth-reduced defoliated trees. We performed continuous 13C-labeling coupled to clipping to quantify the effects of simulated browsing on the growth, leaf morphology and relative allocation of stored vs recently assimilated C to the growth (bulk biomass) and non-structural carbohydrate (NSC) stores (soluble sugars and starch) of the different organs of two tree species: diffuse-porous (Betula pubescens Ehrh.) and ring-porous (Quercus petraea [Matt.] Liebl.). Carbon-transfers from plants to bulk and rhizosphere soil were also evaluated. Clipped birch and oak trees shifted their C-allocation patterns above-ground as a means to recover from defoliation. However, such increased allocation to current-year stems and leaves did not entail reductions in the allocation to the rhizosphere, which remained unchanged between clipped and control trees of both species. Betula pubescens and Q. petraea showed differences in their vulnerability and recovery strategies to clipping, the ring-porous species being less affected in terms of growth and architecture by clipping than the diffuse-porous. These contrasting patterns could be partly explained by differences in their C cycling after clipping. Defoliated oaks showed a faster recovery of their canopy biomass, which was supported by increased allocation of new C, but associated with large decreases in their fine root biomass. Following clipping, both species recovered NSC pools to a larger extent than growth, but the allocation of 13C-labeled photo-assimilates into storage compounds was not increased as compared with controls. Despite their different response to clipping, our results indicate no preventative allocation into storage occurred during the first year after clipping in either of the species., (© The Author(s) 2020. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Although the mechanisms of nodule N(2) fixation in legumes are now well documented, some uncertainty remains on the metabolic consequences of water deficit. In most cases, little consideration is given to other organs and, therefore, the coordinated changes in metabolism in leaves, roots, and nodules are not well known. Here, the effect of water restriction on exclusively N(2)-fixing alfalfa (Medicago sativa L.) plants was investigated, and proteomic, metabolomic, and physiological analyses were carried out. It is shown that the inhibition of nitrogenase activity caused by water restriction was accompanied by concerted alterations in metabolic pathways in nodules, leaves, and roots. The data suggest that nodule metabolism and metabolic exchange between plant organs nearly reached homeostasis in asparagine synthesis and partitioning, as well as the N demand from leaves. Typically, there was (i) a stimulation of the anaplerotic pathway to sustain the provision of C skeletons for amino acid (e.g. glutamate and proline) synthesis; (ii) re-allocation of glycolytic products to alanine and serine/glycine; and (iii) subtle changes in redox metabolites suggesting the implication of a slight oxidative stress. Furthermore, water restriction caused little change in both photosynthetic efficiency and respiratory cost of N(2) fixation by nodules. In other words, the results suggest that under water stress, nodule metabolism follows a compromise between physiological imperatives (N demand, oxidative stress) and the lower input to sustain catabolism.

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 its relevance, protein regulation, metabolic adjustment, and the physiological status of plants under drought is not well understood in relation to the role of nitrogen fixation in nodules. In this study, nodulated alfalfa plants were exposed to drought conditions. The study determined the physiological, metabolic, and proteomic processes involved in photosynthetic inhibition in relation to the decrease in nitrogenase (N(ase)) activity. The deleterious effect of drought on alfalfa performance was targeted towards photosynthesis and N(ase) activity. At the leaf level, photosynthetic inhibition was mainly caused by the inhibition of Rubisco. The proteomic profile and physiological measurements revealed that the reduced carboxylation capacity of droughted plants was related to limitations in Rubisco protein content, activation state, and RuBP regeneration. Drought also decreased amino acid content such as asparagine, and glutamic acid, and Rubisco protein content indicating that N availability limitations were caused by N(ase) activity inhibition. In this context, drought induced the decrease in Rubisco binding protein content at the leaf level and proteases were up-regulated so as to degrade Rubisco protein. This degradation enabled the reallocation of the Rubisco-derived N to the synthesis of amino acids with osmoregulant capacity. Rubisco degradation under drought conditions was induced so as to remobilize Rubisco-derived N to compensate for the decrease in N associated with N(ase) inhibition. Metabolic analyses showed that droughted plants increased amino acid (proline, a major compound involved in osmotic regulation) and soluble sugar (D-pinitol) levels to contribute towards the decrease in osmotic potential (Ψ(s)). At the nodule level, drought had an inhibitory effect on N(ase) activity. This decrease in N(ase) activity was not induced by substrate shortage, as reflected by an increase in total soluble sugars (TSS) in the nodules. Proline accumulation in the nodule could also be associated with an osmoregulatory response to drought and might function as a protective agent against ROS. In droughted nodules, the decrease in N(2) fixation was caused by an increase in oxygen resistance that was induced in the nodule. This was a mechanism to avoid oxidative damage associated with reduced respiration activity and the consequent increase in oxygen content. This study highlighted that even though drought had a direct effect on leaves, the deleterious effects of drought on nodules also conditioned leaf responsiveness.

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.

Background and Aims: Chilling results in a significant decrease in Rubisco content and increased generation of reactive oxygen species (ROS) in cucumber (Cucumis sativus), a chilling-sensitive species. The role of roots in the regulation of the tolerance is unknown. Here, cucumber plants grafted onto figleaf gourd (Cucurbita ficifolia), a chilling-tolerant species were used to study the role of roots in the regulation of shoot functioning and the associated root-to-shoot communication., Methods: Gas exchange and chlorophyll fluorescence were measured using an infrared gas analyser combined with a pulse amplitude fluorimeter during chilling at 14 degrees C or 7 degrees C and subsequent recovery. At the same time, Rubisco content and activity and ROS generation were spectrophotometrically assayed. Abscisic acid and cytokinin concentrations in xylem sap were also determined by enzyme-linked immunosorbent assay., Key Results and Conclusions: Grafted plants showed a significantly higher light-saturated rate of CO(2) assimilation (A(sat)) than own-rooted plants when roots were gradually cooled, but no differences were detected when shoots were cooled. Chill at 7 degrees C irreversibly reduced A(sat), and significantly decreased maximum carboxylation activity, Rubisco content and initial Rubisco activity. However, grafted plants showed weaker inhibition, together with decreased electron flux in the water-water cycle. Higher activity of antioxidant enzymes with less ROS production was found in grafted plants. In addition, ABA concentration increased by 48.4-fold whilst cytokinin concentration decreased by 91.5% in the xylem sap of own-rooted plants after exposure to a 7 degrees C chill. In comparison, ABA and cytokinin concentrations increased by 10.5-fold and 36.9%, respectively, for the grafted plants. Improved plant growth was also observed in grafted plants after the chill. These results suggest that some signals coming from chilling-resistant roots (i.e. ABA and cytokinins) protect leaf photosynthesis in shoots of chilling-sensitive plants.

Genetic improvement of agronomic crops is necessary to cope with chilling stress. To identify the physiological factors responsible for this genotypic difference in chill-induced inhibition of photosynthesis, leaf CO2 assimilation, the electron flux in the chloroplast and the antioxidant metabolism in isolated chloroplasts were examined in two genotypes of cucumber (Cucumis sativus) plants with distinct chilling tolerance. Cucumber plants were exposed to 100 micromol m(-2) s(-1) at 9/7 degrees C (day/night) for 10 d and were then returned to optimal conditions for 2 d. Chilling resulted in more significant reductions in rbcL and rbcS transcripts, ribulose-1,5-bisphosphate carboxylase-oxygenase (Rubisco) content and initial Rubisco activity, leading to higher electron flux to O2 in the chilling-sensitive genotype than in the chilling-tolerant genotype. The chilling-tolerant genotype showed lower H2O2 contents in the chloroplasts by maintaining higher H2O2-scavenging activity in the chloroplasts than in the chilling-sensitive genotype. H2O2 accumulation in chloroplast was negatively correlated with the initial Rubisco activity and photosynthetic rate.

Very little is known about the primary carbon metabolism of the high mountain plant Ranunculus glacialis. It is a species with C3 photosynthesis, but with exceptionally high malate content in its leaves, the biological significance of which remains unclear. 13C/12C-isotope ratio mass spectrometry (IRMS) and 13C-nuclear magnetic resonance (NMR) labelling were used to study the carbon metabolism of R. glacialis, paying special attention to respiration. Although leaf dark respiration was high, the temperature response had a Q10 of 2, and the respiratory quotient (CO2 produced divided by O2 consumed) was nearly 1, indicating that the respiratory pool is comprised of carbohydrates. Malate, which may be a large carbon substrate, was not respired. However, when CO2 fixed by photosynthesis was labelled, little labelling of the CO2 subsequently respired in the dark was detected, indicating that: (i) most of the carbon recently assimilated during photosynthesis is not respired in the dark; and (ii) the carbon used for respiration originates from (unlabelled) reserves. This is the first demonstration of such a low metabolic coupling of assimilated and respired carbon in leaves. The biological significance of the uncoupling between assimilation and respiration is discussed.

The effects of 24-epibrassinolide (EBR) spray application on gas-exchange, chlorophyll fluorescence characteristics, Rubisco activity, and carbohydrate metabolism were investigated in cucumber (Cucumis sativus L. cv. Jinchun No. 3) plants grown in a greenhouse. EBR significantly increased the light-saturated net CO(2) assimilation rate (A(sat)) from 3 h to 7d after spraying, with 0.1 mg l(-1) EBR proving most effective. Increased A(sat) in EBR-treated leaves was accompanied by increases in the maximum carboxylation rate of Rubisco (V(c,max)) and in the maximum rate of RuBP regeneration (J(max)). EBR-treated leaves also had a higher quantum yield of PSII electron transport (phi(PSII)) than the controls, which was mainly due to a significant increase in the photochemical quenching (q(P)), with no change in the efficiency of energy capture by open PSII reaction centres (F'(v)/F'(m)). EBR did not influence photorespiration. In addition, significant increases in the initial activity of Rubisco and in the sucrose, soluble sugars, and starch contents were observed followed by substantial increases in sucrose phosphate synthase (SPS), sucrose synthase (SS), and acid invertase (AI) activities after EBR treatment. It was concluded that EBR increases the capacity of CO(2) assimilation in the Calvin cycle, which was mainly attributed to an increase in the initial activity of Rubisco.