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Elevated greenhouse gases (GHGs) induce adverse conditions directly and indirectly, causing decreases in plant productivity. To deal with climate change effects, plants have developed various mechanisms including the fine-tuning of metabolism. Plant respiratory metabolism is highly flexible due to the presence of various alternative pathways. The mitochondrial alternative oxidase (AOX) respiratory pathway is responsive to these changes, and several lines of evidence suggest it plays a role in reducing excesses of reactive oxygen species (ROS) and reactive nitrogen species (RNS) while providing metabolic flexibility under stress. Here we discuss the importance of the AOX pathway in dealing with elevated carbon dioxide (CO2), nitrogen oxides (NOx), ozone (O3), and the main abiotic stresses induced by climate change. ; This work is supported by the DST-DAAD exchange program between KJG and ARF. This work in the KJG lab is partly supported by a Ramalingaswami Fellowship and IYBA from the Department of Biotechnology, Government of India. IFS has received funding from the European Union's Horizon 2020 research and innovation programme under the Marie Skłodowska-Curie grant agreement no. 753301. ; Peer reviewed

This article belongs to the Special Issue Tackling Grapevine Water Relations in a Global Warming Scenario. ; Climate change forecasts suggest temperature increases and lower rainfall rates, both of which challenge viticulture, particularly in semi-arid areas where water availability is critical. In this scenario, the use of the genetic variability in grapevine varieties reported around the world represents an important strategy for the selection of climate-resilient cultivars. In this work, physiological and metabolomics analyses were conducted to compare the water deficit stress (WDS) responses of red and white, local and widespread grapevines cultivars. Leaf gas exchange, water use efficiency (WUE) and water relation parameters were determined in plants under well-watered and WDS conditions alongside assessment of the levels of foliar primary metabolites using gas-chromatography coupled to mass-spectrometry. Results denote that red and white local cultivars displayed more adapted physiological performance under WDS as compared to the widely-distributed ones. Multivariate analyses and specific changes in leaf primary metabolites indicate genotype-specific responses of local cultivars as compared to widespread ones. Differences in ascorbate-related and shikimate/phenylpropanoid metabolism could explain the better physiological performance under WDS in red local as compared to widespread cultivars. On the other hand, coordinated changes in respiratory- and stress-related sugars and amino acids could underlie the better WUE under WDS in the white local cultivar. All these results suggest several metabolic targets that could be useful as metabolic markers or for metabolic engineering in grapevine breeding programs to improve drought tolerance. V ; This research was funded by the project RTA2013-00068-C03-03 (INIA and FEDER funds) from Ministerio de Economía y Competitividad (MINECO, Spain). I.F.-S. has received funding from the European Union's Horizon 2020 research and innovation programme under the Marie Skłodowska-Curie grant agreement no. 753301'. M.J.C.-M. was financed with a Juan de la Cierva contract (FPDI-2013-18167). ; Peer reviewed

[EN] When growing in search for light, plants can experience continuous or occasional shading by other plants. Plant proximity causes a decrease in the ratio of R to far-red light (low R:FR) due to the preferential absorbance of R light and reflection of FR light by photosynthetic tissues of neighboring plants. This signal is often perceived before actual shading causes a reduction in photo-synthetically active radiation (low PAR). Here, we investigated how several Brassicaceae species from different habitats respond to low R:FR and low PAR in terms of elongation, photosynthesis, and photoacclimation. Shade-tolerant plants such as hairy bittercress (Cardamine hirsuta) displayed a good adaptation to low PAR but a poor or null response to low R:FR exposure. In contrast, shade-avoider species, such as Arabidopsis (Arabidopsis thaliana), showed a weak photosynthetic performance under low PAR but they strongly elongated when exposed to low R:FR. These responses could be genetically uncoupled. Most interestingly, exposure to low R:FR of shade-avoider (but not shade-tolerant) plants improved their photoacclimation to low PAR by triggering changes in photosynthesis-related gene expression, pigment accumulation, and chloroplast ultrastructure. These results indicate that low R:FR signaling unleashes molecular, metabolic, and developmental responses that allow shade-avoider plants (including most crops) to adjust their photosynthetic capacity in anticipation of eventual shading by nearby plants. ; L.M. received a predoctoral fellowships from La Caixa Foundation (INPhINIT fellowship LCF/BQ/IN18/11660004). W.Q. is a recipient of a predoctoral Chinese Scholarship Council (CSC) fellowship. A.I.-S. is supported by a predoctoral fellowship from MICINN (PRE2018-083610). I.F.-S. has received funding from the European Union's Horizon 2020 research and innovation programme under the Marie Sklodowska-Curie grant agreement no. 753301. Our research is supported by grants from MICINN-FEDER (BIO2017-85316-R, and BIO2017-84041-P) and ...

This article belongs to the Section Plant Physiology and Metabolism. ; Plant respiration provides metabolic flexibility under changing environmental conditions by modulating the activity of the nonphosphorylating alternative pathways from the mitochondrial electron transport chain, which bypass the main energy-producing components of the cytochrome oxidase pathway (COP). While adjustments in leaf primary metabolism induced by changes in day length are well studied, possible differences in the in vivo contribution of the COP and the alternative oxidase pathway (AOP) between different photoperiods remain unknown. In our study, in vivo electron partitioning between AOP and COP and expression analysis of respiratory components, photosynthesis, and the levels of primary metabolites were studied in leaves of wild-type (WT) plants and cytochrome c (CYTc) mutants, with reduced levels of COP components, under short- and long-day photoperiods. Our results clearly show that differences in AOP and COP in vivo activities between WT and cytc mutants depend on the photoperiod likely due to energy and stress signaling constraints. Parallel responses observed between in vivo respiratory activities, TCA cycle intermediates, amino acids, and stress signaling metabolites indicate the coordination of different pathways of primary metabolism to support growth adaptation under different photoperiods. ; This research was funded by FONDECYT No. 1191118 from the National Agency for Research and Development (ANID) and by the Agencia Nacional de Promoción Científica y Tecnológica, grant PICT-2018-01439. In addition, I.F.-S. has received funding from the European Union's Horizon 2020 research and innovation programme under the Marie Skłodowska-Curie grant agreement no. 753301, the 'Ramon y Cajal' contract RYC2019-027244-I/AEI/10.13039/501100011033 and the European Social Fund. ; Peer reviewed

In plants, there is a complex interaction between carbon (C) and nitrogen (N) metabolism, and its coordination is fundamental for plant growth and development. Here, we studied the influence of thioredoxin (Trx) m on C and N partitioning using tobacco plants overexpressing Trx m from the chloroplast genome. The transgenic plants showed altered metabolism of C (lower leaf starch and soluble sugar accumulation) and N (with higher amounts of amino acids and soluble protein), which pointed to an activation of N metabolism at the expense of carbohydrates. To further delineate the effect of Trx m overexpression, metabolomic and enzymatic analyses were performed on these plants. These results showed an up-regulation of the glutamine synthetase–glutamate synthase pathway; specifically tobacco plants overexpressing Trx m displayed increased activity and stability of glutamine synthetase. Moreover, higher photorespiration and nitrate accumulation were observed in these plants relative to untransformed control plants, indicating that overexpression of Trx m favors the photorespiratory N cycle rather than primary nitrate assimilation. Taken together, our results reveal the importance of Trx m as a molecular mediator of N metabolism in plant chloroplasts. ; This work was supported by the Spanish Ministry of Science and Innovation (AGL2016-79868) and Phenome-FPPN (ANR-11-INBS-0012). MA is a recipient of a PhD fellowship from the Spanish Ministry of Education (FPU13/01675). IFS has received funding from the European Union's Horizon 2020 research and innovation programme under the Marie Skłodowska-Curie grant agreement no. 753301, the 'Ramon y Cajal' contract RYC2019-027244-I/AEI/10.13039/501100011033, and the European Social Fund. ; Peer reviewed

[EN] Geranylgeranyl diphosphate (GGPP) produced by GGPP synthase (GGPPS) serves as a precursor for many plastidial isoprenoids, including carotenoids. Phytoene synthase (PSY) converts GGPP into phytoene, the first committed intermediate of the carotenoid pathway. Here we used biochemical, molecular, and genetic tools to characterise the plastidial members of the GGPPS family in tomato (Solanum lycopersicum) and their interaction with PSY isoforms. The three tomato GGPPS isoforms found to localise in plastids (SlG1, 2 and 3) exhibit similar kinetic parameters. Gene expression analyses showed a preferential association of individual GGPPS and PSY isoforms when carotenoid biosynthesis was induced during root mycorrhization, seedling de-etiolation and fruit ripening. SlG2, but not SlG3, physically interacts with PSY proteins. By contrast, CRISPR-Cas9 mutants defective in SlG3 showed a stronger impact on carotenoid levels and derived metabolic, physiological and developmental phenotypes compared with those impaired in SlG2. Double mutants defective in both genes could not be rescued. Our work demonstrates that the bulk of GGPP production in tomato chloroplasts and chromoplasts relies on two cooperating GGPPS paralogues, unlike other plant species such as Arabidopsis thaliana, rice or pepper, which produce their essential plastidial isoprenoids using a single GGPPS isoform. ; We greatly thank Juan Antonio Lopez-Raez for providing cDNA samples of nonmycorrhized and mycorrhized tomato roots; Ernesto Llamas for providing the pGWB417_AtPRK construct, and Albert Ferrer and Laura Gutierrez for the pDE-Cas9 (with kanamycin resistance) plasmid. The technical support of M. Rosa Rodriguez-Goberna and all CRAG services is also appreciated. This work was funded by the European Regional Development Fund (FEDER) and the Spanish Agencia Estatal de Investigacion (grants BIO2017-84041-P and BIO2017-90877-REDT) and Generalitat de Catalunya (2017SGR-710) to MRC. Support by the collaborative European Union's Horizon 2020 (EU-H2020) ...

The alternative oxidase pathway (AOP) is associated with excess energy dissipation in leaves of terrestrial plants. To address whether this association is less important in palustrine plants, we compared the role of AOP in balancing energy and carbon metabolism in palustrine and terrestrial environments by identifying metabolic relationships between primary carbon metabolites and AOP in each habitat. We measured oxygen isotope discrimination during respiration, gas exchange, and metabolite profiles in aerial leaves of ten fern and angiosperm species belonging to five families organized as pairs of palustrine and terrestrial species. We performed a partial least square model combined with variable importance for projection to reveal relationships between the electron partitioning to the AOP (τa) and metabolite levels. Terrestrial plants showed higher values of net photosynthesis (AN) and τa, together with stronger metabolic relationships between τa and sugars, important for water conservation. Palustrine plants showed relationships between τa and metabolites related to the shikimate pathway and the GABA shunt, to be important for heterophylly. Excess energy dissipation via AOX is less crucial in palustrine environments than on land. The basis of this difference resides in the contrasting photosynthetic performance observed in each environment, thus reinforcing the importance of AOP for photosynthesis. ; This work was mainly supported by Ministerio de Economía y Competitividad (MINECO, Spain) (project PGC2018-093824-B-C41) and the ERDF (FEDER). Others funding were provided by FONDECYT No. 1191118 from National Agency for Research and Development (ANID) and the Chilean Scholarship Program/Becas de doctorado nacional/2017–21180329, the European Union's Horizon 2020 Research and Innovation Program under the Marie Skłodowska-Curie grant agreement no. 753301, the 'Ramon y Cajal' contract RYC2019-027244-I/AEI/10.13039/501100011033 and the European Social Fund. ; Peer reviewed

This article belongs to the Special Issue Plant Respiration. ; The interaction of the alternative oxidase (AOX) pathway with nutrient metabolism is important for understanding how respiration modulates ATP synthesis and carbon economy in plants under nutrient deficiency. Although AOX activity reduces the energy yield of respiration, this enzymatic activity is upregulated under stress conditions to maintain the functioning of primary metabolism. The in vivo metabolic regulation of AOX activity by phosphorus (P) and nitrogen (N) and during plant symbioses with Arbuscular mycorrhizal fungi (AMF) and Rhizobium bacteria is still not fully understood. We highlight several findings and open questions concerning the in vivo regulation of AOX activity and its impact on plant metabolism during P deficiency and symbiosis with AMF. We also highlight the need for the identification of which metabolic regulatory factors of AOX activity are related to N availability and nitrogen-fixing legume-rhizobia symbiosis in order to improve our understanding of N assimilation and biological nitrogen fixation. ; This work was supported by FONDECYT No. 1191118 from National Agency for Research and Development (ANID) and the Chilean Scholarship Program/Becas de doctorado nacional/2017–21180329. I.F.-S. has received funding from the European Union's Horizon 2020 research and innovation program under the Marie Skłodowska-Curie grant agreement no. 753301. ; Peer reviewed

In plants, there is a complex interaction between carbon (C) and nitrogen (N) metabolism, and its coordination is fundamental for plant growth and development. Here, we studied the influence of thioredoxin (Trx) m on C and N partitioning using tobacco plants overexpressing Trx m from the chloroplast genome. The transgenic plants showed altered metabolism of C (lower leaf starch and soluble sugar accumulation) and N (with higher amounts of amino acids and soluble protein), which pointed to an activation of N metabolism at the expense of carbohydrates. To further delineate the effect of Trx m overexpression, metabolomic and enzymatic analyses were performed on these plants. These results showed an up-regulation of the glutamine synthetase-glutamate synthase pathway; specifically tobacco plants overexpressing Trx m displayed increased activity and stability of glutamine synthetase. Moreover, higher photorespiration and nitrate accumulation were observed in these plants relative to untransformed control plants, indicating that overexpression of Trx m favors the photorespiratory N cycle rather than primary nitrate assimilation. Taken together, our results reveal the importance of Trx m as a molecular mediator of N metabolism in plant chloroplasts. ; This work was supported by the Spanish Ministry of Science and Innovation (AGL2016-79868) and Phenome-FPPN (ANR-11-INBS-0012). MA is a recipient of a PhD fellowship from the Spanish Ministry of Education (FPU13/01675). IFS has received funding from the European Union's Horizon 2020 research and innovation programme under the Marie Skłodowska-Curie grant agreement no. 753301, the 'Ramon y Cajal' contract RYC2019-027244-I/AEI/10.13039/501100011033, and the European Social Fund.