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Summary Measurements of stable isotope ratios in organic compounds are widely used tools for plant ecophysiological studies. However, the complexity of the processes involved in shaping hydrogen isotope values (δ2H) in plant carbohydrates has limited its broader application. To investigate the underlying biochemical processes responsible for 2H fractionation among water, sugars, and cellulose in leaves, we studied the three main CO2 fixation pathways (C3, C4, and CAM) and their response to changes in temperature and vapor pressure deficit (VPD). We show significant differences in autotrophic 2H fractionation (εA) from water to sugar among the pathways and their response to changes in air temperature and VPD. The strong 2H depleting εA in C3 plants is likely driven by the photosynthetic H+ production within the thylakoids, a reaction that is spatially separated in C4 and strongly reduced in CAM plants, leading to the absence of 2H depletion in the latter two types. By contrast, we found that the heterotrophic 2H‐fractionation (εH) from sugar to cellulose was very similar among the three pathways and is likely driven by the plant's metabolism, rather than by isotopic exchange with leaf water. Our study offers new insights into the biochemical drivers of 2H fractionation in plant carbohydrates. [ABSTRACT FROM AUTHOR]

Summary: Non‐structural carbohydrates (NSCs), as the labile fraction and dominant carbon currency, are essential mediators of plant adaptation to environments. However, whether and how NSC coordinates with plant economic strategy frameworks, particularly the well‐recognized leaf economics spectrums (LES) and root economics space (RES), remains unclear.We examined the relationships between NSC and key plant economics traits in leaves and fine roots across 90 alpine coniferous populations on the Tibetan Plateau, China.We observed contrasting coordination of NSC with economics traits in leaves and roots. Leaf total NSC and soluble sugar aligned with the leaf economic spectrum, conveying a trade‐off between growth and storage in leaves. However, NSC in roots was independent of the root economic spectrum, but highly coordinated with root foraging, with more starch and less sugar in forage‐efficient, thinner roots. Further, NSC‐trait coordination in leaves and roots was, respectively, driven by local temperature and precipitation.These findings highlight distinct roles of NSC in shaping the above‐ and belowground multidimensional economics trait space, and NSC‐based carbon economics provides a mechanistic understanding of how plants adapt to heterogeneous habitats and respond to environmental changes. [ABSTRACT FROM AUTHOR]

Summary: Drought affects the complex interactions between Norway spruce, the bark beetle Ips typographus and associated microorganisms. We investigated the interplay of tree water status, defense and carbohydrate reserves with the incidence of bark beetle attack and infection of associated fungi in mature spruce trees.We installed roofs to induce a 2‐yr moderate drought in a managed spruce stand to examine a maximum of 10 roof and 10 control trees for resin flow (RF), predawn twig water potentials, terpene, phenolic and carbohydrate bark concentrations, and bark beetle borings in field bioassays before and after inoculation with Endoconidiophora polonica and Grosmannia penicillata.Drought‐stressed trees showed more attacks and significantly longer fungal lesions than controls, but maintained terpene resin defenses at predrought levels. Reduced RF and lower mono‐ and diterpene, but not phenolic concentrations were linked with increased host selection. Bark beetle attack and fungi stimulated chemical defenses, yet G. penicillata reduced phenolic and carbohydrate contents.Chemical defenses did not decrease under mild, prolonged drought in our simulated small‐scale biotic infestations. However, during natural mass attacks, reductions in carbon fixation under drought, in combination with fungal consumption of carbohydrates, may deplete tree defenses and facilitate colonization by I. typographus. [ABSTRACT FROM AUTHOR]

Switchgrass (Panicum virgatum) is a bioenergy model crop valued for its energy efficiency and drought tolerance. The related monocot species rice (Oryza sativa) and maize (Zea mays) deploy species-specific, specialized metabolites as core stress defenses. By contrast, specialized chemical defenses in switchgrass are largely unknown. To investigate specialized metabolic drought responses in switchgrass, we integrated tissue-specific transcriptome and metabolite analyses of the genotypes Alamo and Cave-in-Rock that feature different drought tolerance. The more drought-susceptible Cave-in-Rock featured an earlier onset of transcriptomic changes and significantly more differentially expressed genes in response to drought compared to Alamo. Specialized pathways showed moderate differential expression compared to pronounced transcriptomic alterations in carbohydrate and amino acid metabolism. However, diterpenoid-biosynthetic genes showed drought-inducible expression in Alamo roots, contrasting largely unaltered triterpenoid and phenylpropanoid pathways. Metabolomic analyses identified common and genotype-specific flavonoids and terpenoids. Consistent with transcriptomic alterations, several root diterpenoids showed significant drought-induced accumulation, whereas triterpenoid abundance remained predominantly unchanged. Structural analysis verified select drought-responsive diterpenoids as oxygenated furanoditerpenoids. Drought-dependent transcriptome and metabolite profiles provide the foundation to understand the molecular mechanisms underlying switchgrass drought responses. Accumulation of specialized root diterpenoids and corresponding pathway transcripts supports a role in drought stress tolerance.

Summary: Leaf‐out in temperate forests is a critical transition point each spring and advancing with global change. The mechanism linking phenological variation to external cues is poorly understood. Nonstructural carbohydrate (NSC) availability may be key.Here, we use branch cuttings from northern red oak (Quercus rubra) and measure NSCs throughout bud development in branch tissue. Given genes and environment influence phenology, we placed branches in an arrayed factorial experiment (three temperatures × two photoperiods, eight genotypes) to examine their impact on variation in leaf‐out timing and corresponding NSCs.Despite significant differences in leaf‐out timing between treatments, NSC patterns were much more consistent, with all treatments and genotypes displaying similar NSC concentrations across phenophases. Notably, the moderate and hot temperature treatments reached the same NSC concentrations and phenophases at the same growing degree days (GDD), but 20 calendar days apart, while the cold treatment achieved only half the GDD of the other two.Our results suggest that NSCs are coordinated with leaf‐out and could act as a molecular clock, signaling to cells the passage of time and triggering leaf development to begin. This link between NSCs and budburst is critical for improving predictions of phenological timing. [ABSTRACT FROM AUTHOR]

Summary: Shifts in the age or turnover time of non‐structural carbohydrates (NSC) may underlie changes in tree growth under long‐term increases in drought stress associated with climate change. But NSC responses to drought are challenging to quantify, due in part to large NSC stores in trees and subsequently long response times of NSC to climate variation.We measured NSC age (Δ14C) along with a suite of ecophysiological metrics in Pinus edulis trees experiencing either extreme short‐term drought (−90% ambient precipitation plot, 2020–2021) or a decade of severe drought (−45% plot, 2010–2021). We tested the hypothesis that carbon starvation – consumption exceeding synthesis and storage – increases the age of sapwood NSC.One year of extreme drought had no impact on NSC pool size or age, despite significant reductions in predawn water potential, photosynthetic rates/capacity, and twig and needle growth. By contrast, long‐term drought halved the age of the sapwood NSC pool, coupled with reductions in sapwood starch concentrations (−75%), basal area increment (−39%), and bole respiration rates (−28%).Our results suggest carbon starvation takes time, as tree carbon reserves appear resilient to extreme disturbance in the short term. However, after a decade of drought, trees apparently consumed old stored NSC to support metabolism. [ABSTRACT FROM AUTHOR]

Summary: Recent methodological advancements in determining the nonexchangeable hydrogen isotopic composition (δ2Hne) of plant carbohydrates make it possible to disentangle the drivers of hydrogen isotope (2H) fractionation processes in plants.Here, we investigated the influence of phylogeny on the δ2Hne of twig xylem cellulose and xylem water, as well as leaf sugars and leaf water, across 73 Northern Hemisphere tree and shrub species growing in a common garden.2H fractionation in plant carbohydrates followed distinct phylogenetic patterns, with phylogeny reflected more in the δ2Hne of leaf sugars than in that of twig xylem cellulose. Phylogeny had no detectable influence on the δ2Hne of twig or leaf water, showing that biochemistry, not isotopic differences in plant water, caused the observed phylogenetic pattern in carbohydrates. Angiosperms were more 2H‐enriched than gymnosperms, but substantial δ2Hne variations also occurred at the order, family, and species levels within both clades. Differences in the strength of the phylogenetic signals in δ2Hne of leaf sugars and twig xylem cellulose suggest that the original phylogenetic signal of autotrophic processes was altered by subsequent species‐specific metabolism.Our results will help improve 2H fractionation models for plant carbohydrates and have important consequences for dendrochronological and ecophysiological studies. [ABSTRACT FROM AUTHOR]

This article is a Commentary on Potkay & Feng (2023), 238: 506–528. [ABSTRACT FROM AUTHOR]

Summary: During stem elongation, wheat (Triticum aestivum) increases its stem carbohydrate content before anthesis as a reserve for grain filling. Hydraulic functioning during this mobilization process is not well understood, and contradictory results exist on the direct effect of drought on carbohydrate mobilization.In a dedicated experiment, wheat plants were subjected to drought stress during carbohydrate mobilization. Measurements, important to better understand stem physiology, showed some unexpected patterns that could not be explained by our current knowledge on water transport. Traditional water flow and storage models failed to properly describe the drought response in wheat stems during carbohydrate mobilization. To explain the measured patterns, hypotheses were formulated and integrated in a dedicated model for wheat.The new mechanistic model simulates two hypothetical water storage compartments: one where water is quickly exchanged with the xylem and one that contains the carbohydrate storage. Water exchange between these compartments is turgor‐driven. The model was able to simulate the measured increase in stored carbohydrate concentrations with a decrease in water content and stem diameter.Calibration of the model showed the importance of turgor‐driven apoplastic water flow during carbohydrate mobilization. This resulted in an increase in stem hydraulic capacitance, which became more important under drought stress. [ABSTRACT FROM AUTHOR]

Summary: Trade‐offs among carbon sinks constrain how trees physiologically, ecologically, and evolutionarily respond to their environments. These trade‐offs typically fall along a productive growth to conservative, bet‐hedging continuum. How nonstructural carbohydrates (NSCs) stored in living tree cells (known as carbon stores) fit in this trade‐off framework is not well understood.We examined relationships between growth and storage using both within species genetic variation from a common garden, and across species phenotypic variation from a global database.We demonstrate that storage is actively accumulated, as part of a conservative, bet‐hedging life history strategy. Storage accumulates at the expense of growth both within and across species. Within the species Populus trichocarpa, genetic trade‐offs show that for each additional unit of wood area growth (in cm2 yr−1) that genotypes invest in, they lose 1.2 to 1.7 units (mg g−1 NSC) of storage. Across species, for each additional unit of area growth (in cm2 yr−1), trees, on average, reduce their storage by 9.5% in stems and 10.4% in roots.Our findings impact our understanding of basic plant biology, fit storage into a widely used growth‐survival trade‐off spectrum describing life history strategy, and challenges the assumptions of passive storage made in ecosystem models today. [ABSTRACT FROM AUTHOR]

Summary: Leaf habit is a major axis of plant diversity that has consequences for carbon balance since the leaf is the primary site of photosynthesis. Nonstructural carbohydrates (NSCs) produced by photosynthesis can be allocated to storage and serve as a resiliency mechanism to future abiotic and biotic stress. However, how leaf habit affects NSC storage in an evolutionary context has not been shown.Using a comparative physiological framework and an analysis of evolutionary model fitting, we examined if variation in NSC storage is explained by leaf habit. We measured sugar and starch concentrations in 51 oak species (Quercus spp.) growing in a common garden and representing multiple evolutions of three different leaf habits (deciduous, brevideciduous and evergreen).The best fitting evolutionary models indicated that deciduous oak species are evolving towards higher NSC concentrations than their brevideciduous and evergreen relatives. Notably, this was observed for starch (the primary storage molecule) in the stem (a long‐term C storage organ).Overall, our work provides insight into the evolutionary drivers of NSC storage and suggests that a deciduous strategy may confer an advantage against stress associated with a changing world. Future work should examine additional clades to further corroborate this idea. [ABSTRACT FROM AUTHOR]

Summary: Nonstructural carbohydrates (NSCs) have been suggested to affect xylem transport under fluctuating water availability, but conclusive evidence is still lacking. We tested the effect of shade‐induced NSC depletion on xylem vulnerability to embolism and hydraulic recovery on Populus nigra saplings.Vulnerability was assessed in light‐exposed (L) and shaded (S) plants with the hydraulic method, and in vivo with the optical method and X‐ray micro‐computed tomography. Plants were stressed to 80% loss of hydraulic conductance (PLC) and re‐irrigated to check for possible recovery. We measured PLC, bark and wood NSC content, as well as xylem sap pH, surface tension (γsap) and sugar concentration, before, during and after drought.Shading induced depletion of stem NSC (mainly starch) reserves. All methods converged in indicating higher xylem vulnerability in S than in L plants. This difference was not explained by xylem vessel and pit anatomy or by γsap. Shading impeded sap acidification and sugar accumulation during drought in S plants and prevented hydraulic recovery, which was observed in L plants.Our results highlight the importance of stem NSCs to sustain xylem hydraulic functioning during drought and suggest that light and/or adequate stem NSC thresholds are required to trigger xylem sap chemical changes involved in embolism recovery. [ABSTRACT FROM AUTHOR]

We know surprisingly little about whole-tree nonstructural carbon (NSC; primarily sugars and starch) budgets. Even less well understood is the mixing between recent photosynthetic assimilates (new NSC) and previously stored reserves. And, NSC turnover times are poorly constrained. We characterized the distribution of NSC in the stemwood, branches, and roots of two temperate trees, and we used the continuous label offered by the radiocarbon (carbon-14, 14C) bomb spike to estimate the mean age of NSC in different tissues. NSC in branches and the outermost stemwood growth rings had the 14C signature of the current growing season. However, NSC in older aboveground and belowground tissues was enriched in 14C, indicating that it was produced from older assimilates. Radial patterns of 14C in stemwood NSC showed strong mixing of NSC across the youngest growth rings, with limited 'mixing in' of younger NSC to older rings. Sugars in the outermost five growth rings, accounting for two-thirds of the stemwood pool, had a mean age < 1 yr, whereas sugars in older growth rings had a mean age > 5 yr. Our results are thus consistent with a previously-hypothesized two-pool ('fast' and 'slow' cycling NSC) model structure. These pools appear to be physically distinct.

Large amounts of carbon are required for plant growth, but young, growing tissues often also have high concentrations of defensive secondary metabolites. Plants' capacity to allocate resources to growth and defense is addressed by the growth-differentiation balance hypothesis and the optimal defense hypothesis, which make contrasting predictions. Isotope labeling can demonstrate whether defense compounds are synthesized from stored or newly fixed carbon, allowing a detailed examination of these hypotheses. Populus trichocarpa saplings were pulse-labeled with 13CO2 at the beginning and end of a growing season, and the 13C signatures of phenolic glycosides (salicinoids), sugars, bulk tissue, and respired CO2 were traced over time. Half of the saplings were also subjected to mechanical damage. Populus trichocarpa followed an optimal defense strategy, investing 13C in salicinoids in expanding leaves directly after labeling. Salicinoids turned over quickly, and their production continued throughout the season. Salicin was induced by early-season damage, further demonstrating optimal defense. Salicinoids appear to be of great value to P. trichocarpa, as they command new C both early and late in the growing season, but their fitness benefits require further study. Export of salicinoids between tissues and biochemical pathways enabling induction also needs research. Nonetheless, the investigation of defense production afforded by isotope labeling lends new insights into plants' ability to grow and defend simultaneously.

The allocation of nonstructural carbon (NSC) to growth, metabolism and storage remains poorly understood, but is critical for the prediction of stress tolerance and mortality. We used the radiocarbon ((14) C) 'bomb spike' as a tracer of substrate and age of carbon in stemwood NSC, CO2 emitted by stems, tree ring cellulose and stump sprouts regenerated following harvesting in mature red maple trees. We addressed the following questions: which factors influence the age of stemwood NSC?; to what extent is stored vs new NSC used for metabolism and growth?; and, is older, stored NSC available for use? The mean age of extracted stemwood NSC was 10 yr. More vigorous trees had both larger and younger stemwood NSC pools. NSC used to support metabolism (stem CO2 ) was 1-2 yr old in spring before leaves emerged, but reflected current-year photosynthetic products in late summer. The tree ring cellulose (14) C age was 0.9 yr older than direct ring counts. Stump sprouts were formed from NSC up to 17 yr old. Thus, younger NSC is preferentially used for growth and day-to-day metabolic demands. More recently stored NSC contributes to annual ring growth and metabolism in the dormant season, yet decade-old and older NSC is accessible for regrowth.

Summary: Under prolonged drought and reduced photosynthesis, plants consume stored nonstructural carbohydrates (NSCs). Stored NSC depletion may impair the regulation of plant water balance, but the underlying mechanisms are poorly understood, and whether such mechanisms are independent of plant water deficit is not known. If so, carbon costs of fungal symbionts could indirectly influence plant drought tolerance through stored NSC depletion.We connected well‐watered Pinus ponderosa seedling pairs via ectomycorrhizal (EM) networks where one seedling was shaded (D) and the other kept illuminated (LD) and compared responses to seedling pairs in full light (L). We measured plant NSCs, osmotic and water potential, and transfer of 13CO2 through EM to explore mechanisms linking stored NSCs to plant water balance regulation and identify potential tradeoffs between plant water retention and EM fungi under carbon‐limiting conditions.NSCs decreased from L to LD to D seedlings. Even without drought, NSC depletion impaired osmoregulation and turgor maintenance, both of which are critical for drought tolerance. Importantly, EM networks propagated NSC depletion and its negative effects on water retention from carbon stressed to nonstressed hosts.We demonstrate that NSC storage depletion influences turgor maintenance independently of plant water deficit and reveal carbon allocation tradeoffs between supporting fungal symbionts and retaining water. [ABSTRACT FROM AUTHOR]

Summary: Nutrient imbalances cause the deterioration of tree health in European forests, but the underlying physiological mechanisms are unknown. Here, we investigated the consequences of decreasing root carbohydrate reserves for phosphorus (P) mobilisation and uptake by forest trees.In P‐rich and P‐poor beech (Fagus sylvatica) forests, naturally grown, young trees were girdled and used to determine root, ectomycorrhizal and microbial activities related to P mobilisation in the organic layer and mineral topsoil in comparison with those in nongirdled trees.After girdling, root carbohydrate reserves decreased. Root phosphoenolpyruvate carboxylase activities linking carbon and P metabolism increased. Root and ectomycorrhizal phosphatase activities and the abundances of bacterial genes catalysing major steps in P turnover increased, but soil enzymes involved in P mobilisation were unaffected. The physiological responses to girdling were stronger in P‐poor than in P‐rich forests. P uptake was decreased after girdling. The soluble and total P concentrations in roots were stable, but fine root biomass declined after girdling.Our results support that carbohydrate depletion results in reduced P uptake, enhanced internal P remobilisation and root biomass trade‐off to compensate for the P shortage. As reductions in root biomass render trees more susceptible to drought, our results link tree deterioration with disturbances in the P supply as a consequence of decreased belowground carbohydrate allocation. [ABSTRACT FROM AUTHOR]

Nonstructural carbohydrate reserves support tree metabolism and growth when current photosynthates are insufficient, offering resilience in times of stress. We monitored stemwood nonstructural carbohydrate (starch and sugars) concentrations of the dominant tree species at three sites in the northeastern United States. We estimated the mean age of the starch and sugars in a subset of trees using the radiocarbon ((14) C) bomb spike. With these data, we then tested different carbon (C) allocation schemes in a process-based model of forest C cycling. We found that the nonstructural carbohydrates are both highly dynamic and about a decade old. Seasonal dynamics in starch (two to four times higher in the growing season, lower in the dormant season) mirrored those of sugars. Radiocarbon-based estimates indicated that the mean age of the starch and sugars in red maple (Acer rubrum) was 7-14 yr. A two-pool (fast and slow cycling reserves) model structure gave reasonable estimates of the size and mean residence time of the total NSC pool, and greatly improved model predictions of interannual variability in woody biomass increment, compared with zero- or one-pool structures used in the majority of existing models. This highlights the importance of nonstructural carbohydrates in the context of forest ecosystem carbon cycling.

Some essential components of fleshy fruits are dependent on photosynthetic activity and carbohydrate metabolism. Nevertheless, the regulatory mechanisms linking chlorophyll and carbohydrate metabolism remain partially understood. Here, we uncovered the role of SlGRAS9 and SlZHD17 transcription factors in controlling chlorophyll and carbohydrate accumulation in tomato fruit. Knockout or knockdown of SlGRAS9 or SlZHD17 resulted in marked increase in chlorophyll content, reprogrammed chloroplast biogenesis and enhanced accumulation of starch and soluble sugars. Combined genome-wide transcriptomic profiling and promoter-binding experiments unveiled a complex mechanism in which the SlGRAS9/SlZHD17 regulatory module modulates the expression of chloroplast and sugar metabolism either via a sequential transcriptional cascade or through binding of both TFs to the same gene promoters, or, alternatively, via parallel pathways where each of the TFs act on different target genes. For instance, the regulation of SlAGPaseS1 and SlSUS1 is mediated by SlZHD17 whereas that of SlVI and SlGLK1 occurs only through SlGRAS9 without the intervention of SlZHD17. Both SlGRAS9 and SlZHD17 can also directly bind the promoter of SlPOR-B to regulate its expression. Taken together, our findings uncover two important regulators acting synergistically to manipulate chlorophyll and carbohydrate accumulation and provide new potential breeding targets for improving fruit quality in fleshy fruits., (© 2024 The Authors. New Phytologist © 2024 New Phytologist Foundation.)

Summary: Parenchyma cells in the xylem store nonstructural carbohydrates (NSC), providing reserves of energy that fuel woody perennials through periods of stress and/or limitations to photosynthesis. If the capacity for storage is subject to selection, then the fraction of wood occupied by living parenchyma should increase towards stressful environments.Ray parenchyma fraction (RPF) and seasonal NSC dynamics were quantified for 12 conifers and three oaks along a transect spanning warm dry foothills (500 m above sea level) to cold wet treeline (3250 m asl) in California's central Sierra Nevada.Mean RPF was lower for both conifer and oak species with warmer dryer ranges. RPF variability increased with elevation or in relation to associated climatic variables in conifers – treeline‐dominant Pinus albicaulis had the lowest mean RPF measured (c. 3.7%), but the highest environmentally standardized variability index. Conifer RPF variability was explained by environment, increasing predominantly towards cooler wetter range edges. In oaks, NSC was explained by environment – values increasing for evergreen and decreasing for deciduous oaks with elevation. Lastly, all species surveyed appear to prioritize filling available RPF with sugar to achieve molarities that balance reasonable tensions over starch to maximize stored carbon.RPF responds to environment but is unlikely to spatially constrain NSC storage. [ABSTRACT FROM AUTHOR]

Wood performs several functions to ensure tree survival and carbon allocation to a finite stem volume leads to trade-offs among cell types. It is not known to what extent these trade-offs modify functional trade-offs and if they are consistent across climates and evolutionary lineages. Twelve wood traits were measured in stems and coarse roots across 60 adult angiosperm tree species from temperate, Mediterranean and tropical climates. Regardless of climate, clear trade-offs occurred among cellular fractions, but did not translate into specific functional trade-offs. Wood density was negatively related to hydraulic conductivity (K th ) in stems and roots, but was not linked to nonstructural carbohydrates (NSC), implying a functional trade-off between mechanical integrity and transport but not with storage. NSC storage capacity was positively associated with K th in stems and negatively in roots, reflecting a potential role for NSC in the maintenance of hydraulic integrity in stems but not in roots. Results of phylogenetic analyses suggest that evolutionary histories cannot explain covariations among traits. Trade-offs occur among cellular fractions, without necessarily modifying trade-offs in function. However, functional trade-offs are driven by coordinated changes among xylem cell types depending on the dominant role of each cell type in stems and roots., (© 2023 The Authors. New Phytologist © 2023 New Phytologist Foundation.)

Summary: While photosynthetic isotope discrimination is well understood, the postphotosynthetic and transport‐related fractionation mechanisms that influence phloem and subsequently tree ring δ13C are less investigated and may vary among species.We studied the seasonal and diel courses of leaf‐to‐phloem δ13C differences of water‐soluble organic matter (WSOM) in vertical crown gradients and followed the assimilate transport via the branches to the trunk phloem at breast height in European beech (Fagus sylvatica) and Douglas fir (Pseudotsuga menziesii). δ13C of individual sugars and cyclitols from a subsample was determined by compound‐specific isotope analysis.In beech, leaf‐to‐phloem δ13C differences in WSOM increased with height and were partly caused by biochemical isotope fractionation between leaf compounds. 13C‐Enrichment of phloem sugars relative to leaf sucrose implies an additional isotope fractionation mechanism related to leaf assimilate export. In Douglas fir, leaf‐to‐phloem δ13C differences were much smaller and isotopically invariant pinitol strongly influenced leaf and phloem WSOM. Trunk phloem WSOM at breast height reflected canopy‐integrated δ13C in beech but not in Douglas fir.Our results demonstrate that leaf‐to‐phloem isotope fractionation and δ13C mixing patterns along vertical gradients can differ between tree species. These effects have to be considered for functional interpretations of trunk phloem and tree ring δ13C. [ABSTRACT FROM AUTHOR]

This article is a Commentary on Drake et al. (2019a), 222: 1298–1312 and Drake et al. (2019b), 222: 1313–1324. [ABSTRACT FROM AUTHOR]

Summary: The concentrations of nonstructural carbohydrates (NSCs) in plant tissues are commonly used as an indicator of total plant carbon (C) supply; but some evidence suggests the possibility for high NSC concentrations during periods of C limitation. Despite this uncertainty, NSC dynamics have not been investigated experimentally under long‐term C limitation.We exposed saplings of 10 temperate tree species differing in shade tolerance to 6% of ambient sunlight for 3 yr to induce C limitation, and also defoliated one species, Carpinus betulus, in the third season. Growth and NSC concentrations were monitored to determine C allocation.Shade strongly reduced growth, but after an initial two‐fold decrease, NSC concentrations of shaded saplings recovered to the level of unshaded saplings by the third season. NSC concentrations were generally more depleted under shade after leaf flush, and following herbivore attacks. Only under shade did artificial defoliation lead to mortality and depleted NSC concentrations in surviving individuals.We conclude that, irrespective of shade tolerance, C storage is maintained under prolonged shading, and thus high NSC concentrations can occur during C limitation. Yet, our results also suggest that decreased NSC concentrations are indicative of C limitation, and that additional leaf loss can lead to lethal C shortage in deep shade. [ABSTRACT FROM AUTHOR]

Summary: Despite the importance of nonstructural carbohydrates (NSC) for growth and survival in woody plants, we know little about whole‐tree NSC storage. The conventional theory suggests that NSC reserves will increase over the growing season and decrease over the dormant season. Here, we compare storage in five temperate tree species to determine the size and seasonal fluctuation of whole‐tree total NSC pools as well as the contribution of individual organs.NSC concentrations in the branches, stemwood, and roots of 24 trees were measured across 12 months. We then scaled up concentrations to the whole‐tree and ecosystem levels using allometric equations and forest stand inventory data.While whole‐tree total NSC pools followed the conventional theory, sugar pools peaked in the dormant season and starch pools in the growing season. Seasonal depletion of total NSCs was minimal at the whole‐tree level, but substantial at the organ level, particularly in branches. Surprisingly, roots were not the major storage organ as branches stored comparable amounts of starch throughout the year, and root reserves were not used to support springtime growth.Scaling up NSC concentrations to the ecosystem level, we find that commonly used, process‐based ecosystem and land surface models all overpredict NSC storage. [ABSTRACT FROM AUTHOR]

The kinetics of wood formation in angiosperms are largely unknown because their complex xylem anatomy precludes using the radial position of vessels and fibers to infer their time of differentiation. We analyzed xylogenesis in ring-porous ash (Fraxinus angustifolia) and diffuse-porous beech (Fagus sylvatica) over 1 yr and proposed a novel procedure to assess the period of vessel and fiber enlargement using a referential radial file (RRF). Our approach captured the dynamics of wood formation and provided a robust estimation of the kinetics of vessel and fiber enlargement. In beech, fibers and vessels had a similar duration of enlargement, decreasing from 14 to 5 d between April and July. In ash, wide vessels formed in April enlarged at a rate of 27 × 10 3  μm 2 d -1 , requiring half the time of contemporary fibers (6 vs 12 d), and less time than the narrower vessels (14 d) formed in May. These findings reveal distinct cell-type-dependent mechanisms for differentiation in diffuse-porous and ring-porous trees, enhancing our understanding of angiosperm wood cell kinetics. Our approach presents an effective method for investigating angiosperm wood formation and provides a more accurate representation of vessel and fiber morphogenesis in wood formation models., (© 2023 The Authors New Phytologist © 2023 New Phytologist Foundation.)

Summary: Nonstructural carbohydrates (NSCs) are important for the growth and survival of trees. Drought may lead to a decrease in tree growth and to NSC depletion, whereas increased soil moisture in otherwise dry ecosystems may increase growth and NSC concentrations. A long‐term (13 yr) irrigation experiment was conducted in a Pinus sylvestris‐dominated forest located at the dry margin of the species in southern Switzerland. We measured the relative leaf area, growth, NSCs, needle δ13C, [N] and [P] in trees on control and irrigated plots. Irrigation resulted in higher growth rates and carbon isotope discrimination, but did not alter NSC levels. Growth and NSC decreased with decreasing leaf area in both treatments, but NSC did not correlate with leaf‐level gas exchange indices, such as foliar δ13C, [N] or [P]. A legacy effect was shown, as trees with initially low leaf area had limited ability to respond to prolonged irrigation. The NSC constancy across treatments provides evidence that carbohydrate storage may stay constant when climate changes are sufficiently slow to allow acclimation. Moreover, we speculate that total leaf area, rather than leaf gas exchange per unit leaf area, drives the variation in whole‐tree carbohydrate dynamics in this system. [ABSTRACT FROM AUTHOR]

Our understanding of how temporal variations of atmospheric water vapour and its isotopic composition (δ18 OV) influence water and assimilates in plants remains limited, restricting our ability to use δ18O as a tracer of ecophysiological processes., We exposed oak ( Quercus robur) saplings under wet and dry soil moisture conditions to 18O-depleted water vapour ( c. − 200‰) at high relative humidity ( c. 93%) for 5 h, simulating a fog event. We then traced the step change in δ18 OV into water and assimilates (e.g. sucrose, hexoses, quercitol and starch) in the leaf lamina, main veins and twigs over 24 h., The immediate δ18 OV effect was highest for δ18O of leaf lamina water, but 40% lower on δ18O of main vein water. To a smaller extent, we also observed changes in δ18O of twig xylem water. Depending on the individual assimilation rate of each plant, the 18O-label was partitioned among different assimilates, with highest changes in δ18O of starch/sucrose and lowest in δ18O of quercitol. Additionally, 18O-label partitioning and allocation towards leaf starch and twig phloem sugars was influenced by the plant water status., Our results have important implications for water isotope heterogeneity in plants and for our understanding of how the δ18O signal is incorporated into biomarkers. [ABSTRACT FROM AUTHOR]

Carbonyl sulphide ( COS) is a potential tracer of gross primary productivity ( GPP), assuming a unidirectional COS flux into the vegetation that scales with GPP. However, carbonic anhydrase ( CA), the enzyme that hydrolyses COS, is expected to be light independent, and thus plants without stomata should continue to take up COS in the dark., We measured net CO2 ( AC) and COS ( AS) uptake rates from two astomatous bryophytes at different relative water contents ( RWCs), COS concentrations, temperatures and light intensities., We found large AS in the dark, indicating that CA activity continues without photosynthesis. More surprisingly, we found a nonzero COS compensation point in light and dark conditions, indicating a temperature-driven COS source with a Q10 (fractional change for a 10°C temperature increase) of 3.7. This resulted in greater AS in the dark than in the light at similar RWC. The processes underlying such COS emissions remain unknown., Our results suggest that ecosystems dominated by bryophytes might be strong atmospheric sinks of COS at night and weaker sinks or even sources of COS during daytime. Biotic COS production in bryophytes could result from symbiotic fungal and bacterial partners that could also be found on vascular plants. [ABSTRACT FROM AUTHOR]

Plants respond to herbivory by reconfiguring hormonal networks, increasing secondary metabolite production and decreasing growth. Furthermore, some plants display a decrease in leaf energy reserves in the form of soluble sugars and starch, leading to the hypothesis that herbivory-induced secondary metabolite production and growth reduction may be linked through a carbohydrate-based resource trade-off., In order to test the above hypothesis, we measured leaf carbohydrates and plant growth in seven genetically engineered Nicotiana attenuata genotypes that are deficient in one or several major herbivore-induced, jasmonate-dependent defensive secondary metabolites and proteins. Furthermore, we manipulated gibberellin and jasmonate signaling, and quantified the impact of these phytohormones on secondary metabolite production, sugar accumulation and growth., Simulated herbivore attack by Manduca sexta specifically reduced leaf sugar concentrations and growth in a jasmonate-dependent manner. These effects were similar or even stronger in defenseless genotypes with intact jasmonate signaling. Gibberellin complementation rescued carbohydrate accumulation and growth in induced plants without impairing the induction of defensive secondary metabolites., These results are consistent with a hormonal antagonism model rather than a resource-cost model to explain the negative relationship between herbivory-induced defenses, leaf energy reserves and growth. [ABSTRACT FROM AUTHOR]

Average responses of forest foliar traits to elevation are well understood, but far less is known about trait distributional responses to elevation at multiple ecological scales. This limits our understanding of the ecological scales at which trait variation occurs in response to environmental drivers and change., We analyzed and compared multiple canopy foliar trait distributions using field sampling and airborne imaging spectroscopy along an Andes-to-Amazon elevation gradient. Field-estimated traits were generated from three community-weighting methods, and remotely sensed estimates of traits were made at three scales defined by sampling grain size and ecological extent., Field and remote sensing approaches revealed increases in average leaf mass per unit area ( LMA), water, nonstructural carbohydrates ( NSCs) and polyphenols with increasing elevation. Foliar nutrients and photosynthetic pigments displayed little to no elevation trend. Sample weighting approaches had little impact on field-estimated trait responses to elevation. Plot representativeness of trait distributions at landscape scales decreased with increasing elevation. Remote sensing indicated elevation-dependent increases in trait variance and distributional skew., Multiscale invariance of LMA, leaf water and NSC mark these traits as candidates for tracking forest responses to changing climate. Trait-based ecological studies can be greatly enhanced with multiscale studies made possible by imaging spectroscopy. [ABSTRACT FROM AUTHOR]

Every existing optimal stomatal model uses photosynthetic carbon assimilation as a proxy for plant evolutionary fitness. However, assimilation and growth are often decoupled, making assimilation less ideal for representing fitness when optimizing stomatal conductance to water vapor and carbon dioxide. Instead, growth should be considered a closer proxy for fitness. We hypothesize stomata have evolved to maximize turgor-driven growth, instead of assimilation, over entire plants' lifetimes, improving their abilities to compete and reproduce. We develop a stomata model that dynamically maximizes whole-stem growth following principles from turgor-driven growth models. Stomata open to assimilate carbohydrates that supply growth and osmotically generate turgor, while stomata close to prevent losses of turgor and growth due to negative water potentials. In steady state, the growth optimization model captures realistic stomatal, growth, and carbohydrate responses to environmental cues, reconciles conflicting interpretations within existing stomatal optimization theories, and explains patterns of carbohydrate storage and xylem conductance observed during and after drought. Our growth optimization hypothesis introduces a new paradigm for stomatal optimization models, elevates the role of whole-plant carbon use and carbon storage in stomatal functioning, and has the potential to simultaneously predict gross productivity, net productivity, and plant mortality through a single, consistent modeling framework., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

Mast seeding is a widespread and widely studied phenomenon. However, the physiological mechanisms that mediate masting events and link them to weather and plant resources are still debated. Here, we explore how masting is affected by plant resource budgets, fruit maturation success, and hormonal coordination of cues including weather and resources. There is little empirical support for the commonly stated hypothesis that plants store carbohydrates over several years to expend in a high-seed year. Plants can switch carbohydrates away from growth in high-seed years, and seed crops are more probably limited by nitrogen or phosphorus. Resources are clearly involved in the proximate mechanisms driving masting, but resource budget (RB) models cannot create masting in the absence of selection because some underlying selective benefit is required to set the level of a 'full’ seed crop at greater than the annual resource increment. Economies of scale (EOSs) provide the ultimate factor selecting for masting, but EOSs probably always interact with resources, which modify the relationship between weather cues and reproduction. Thus, RB and EOS models are not alternative explanations for masting - both are required. Experiments manipulating processes that affect mast seeding will help clarify the physiological mechanisms that underlie mast seeding. [ABSTRACT FROM AUTHOR]

Understanding physiological acclimation of photosynthesis and respiration is important in elucidating the metabolic performance of trees in a changing climate. Does physiological acclimation to climate warming mirror acclimation to seasonal temperature changes?, We grew Eucalyptus tereticornis trees in the field for 14 months inside 9-m tall whole-tree chambers tracking ambient air temperature ( Tair) or ambient Tair + 3°C (i.e. 'warmed'). We measured light- and CO2-saturated net photosynthesis ( Amax) and night-time dark respiration ( R) each month at 25°C to quantify acclimation. Tree growth was measured, and leaf nitrogen (N) and total nonstructural carbohydrate (TNC) concentrations were determined to investigate mechanisms of acclimation., Warming reduced Amax and R measured at 25°C compared to ambient-grown trees. Both traits also declined as mean daily Tair increased, and did so in a similar way across temperature treatments. Amax and R (at 25°C) both increased as TNC concentrations increased seasonally; these relationships appeared to arise from source-sink imbalances, suggesting potential substrate regulation of thermal acclimation., We found that photosynthesis and respiration each acclimated equivalently to experimental warming and seasonal temperature change of a similar magnitude, reflecting a common, nearly homeostatic constraint on leaf carbon exchange that will be important in governing tree responses to climate warming. [ABSTRACT FROM AUTHOR]

Disruption of photosynthesis and carbon transport due to damage to the tree crown and stem cambial cells, respectively, can cause tree mortality. It has recently been proposed that fire-induced dysfunction of xylem plays an important role in tree mortality. Here, we simultaneously tested the impact of a lethal fire dose on nonstructural carbohydrates (NSCs) and xylem hydraulics in Pinus ponderosa saplings. Saplings were burned with a known lethal fire dose. Nonstructural carbohydrates were assessed in needles, main stems, roots and whole plants, and xylem hydraulic conductivity was measured in the main stems up to 29 d postfire. Photosynthesis and whole plant NSCs declined postfire. Additionally, all burned saplings showed 100% phloem/cambium necrosis, and roots of burned saplings had reduced NSCs compared to unburned and defoliated saplings. We further show that, contrary to patterns observed with NSCs, water transport was unchanged by fire and there was no evidence of xylem deformation in saplings that experienced a lethal dose of heat from fire. We conclude that phloem and cambium mortality, and not hydraulic failure, were probably the causes of death in these saplings. These findings advance our understanding of the physiological response to fire-induced injuries in conifer trees., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

• Bark beetle outbreaks are an important cause of tree death, but the process by which trees die remains poorly understood. The effect of beetle attack on whole-tree nonstructural carbohydrate (NSC) dynamics is particularly unclear, despite the potential role of carbohydrates in plant defense and survival. • We monitored NSC dynamics of all organs in attacked and protected lodgepole pines (Pinus contorta) during a mountain pine beetle (Dendroctonus ponderosae) outbreak in British Columbia, starting before beetle flight in June 2011 through October 2012, when most attacked trees had died. • Following attack, NSC concentrations were first reduced in the attacked region of the bole. The first NSC reduction in a distant organ appeared in the needles at the end of 2011, while branch and root NSC did not decline until much later in 2012. Attacked trees that were still alive in October 2012 had less beetle damage, which was negatively correlated with initial bark sugar concentrations in the attack region. • The NSC dynamics of dying trees indicate that trees were killed by a loss of water conduction and not girdling. Further, our results identify locally reduced carbohydrate availability as an important mechanism by which stressors like drought may increase tree susceptibility to biotic attack. [ABSTRACT FROM AUTHOR]

This growth chamber experiment evaluates how temperature and humidity regimes shape soluble carbohydrate pools and growth rates in lichens with different photobionts., We assessed soluble carbohydrates, relative growth rates ( RGRs) and relative thallus area growth rates ( RTAGRs) in Parmelia sulcata (chlorolichen), Peltigera canina (cyanolichen) and Peltigera aphthosa (cephalolichen) cultivated for 14 d (150 μmol m−2 s−1; 12-h photoperiod) at four day : night temperatures (28 : 23°C, 20 : 15°C, 13 : 8°C, 6 : 1°C) and two hydration regimes (hydration during the day, dry at night; hydration day : night)., The major carbohydrates were mannitol (cephalolichen), glucose (cyanolichen) and arabitol (chlorolichen). Mannitol occurred in all species. During cultivation, total carbohydrate pools decreased in cephalo-/cyanolichens, but increased in the chlorolichen. Carbohydrates varied less than growth with temperature and humidity. All lichens grew rapidly, particularly at 13 : 8°C. RGRs and RTAGRs were significantly higher in lichens hydrated for 24 h than for 12 h. Strong photoinhibition occurred in cephalo- and cyanolichens kept in cool dry nights, resulting in positive relationships between RGR and dark-adapted photosystem II ( PSII) efficiency ( Fv/ Fm)., RGR increased significantly with the photobiont-specific carbohydrate pools within all species. Average RGR peaked in the chlorolichen lowest in total and photobiont carbohydrates. Nocturnal hydration improved recovery from photoinhibition and/or enhanced conversion rates of photosynthates into growth. [ABSTRACT FROM AUTHOR]

Reduced carbon (C) assimilation during prolonged drought forces trees to rely on stored C to maintain vital processes like respiration. It has been shown, however, that the use of carbohydrates, a major C storage pool and apparently the main respiratory substrate in plants, strongly declines with decreasing plant hydration. Yet no empirical evidence has been produced to what degree other C storage compounds like lipids and proteins may fuel respiration during drought., We exposed young scots pine trees to C limitation using either drought or shading and assessed respiratory substrate use by monitoring the respiratory quotient, δ13C of respired CO2 and concentrations of the major storage compounds, that is, carbohydrates, lipids and amino acids., Only shaded trees shifted from carbohydrate-dominated to lipid-dominated respiration and showed progressive carbohydrate depletion. In drought trees, the fraction of carbohydrates used in respiration did not decline but respiration rates were strongly reduced. The lower consumption and potentially allocation from other organs may have caused initial carbohydrate content to remain constant during the experiment., Our results suggest that respiratory substrates other than carbohydrates are used under carbohydrate limitation but not during drought. Thus, respiratory substrate shift cannot provide an efficient means to counterbalance C limitation under natural drought. [ABSTRACT FROM AUTHOR]

• Jasmonates regulate plant secondary metabolism and herbivore resistance. How they influence primary metabolites and how this may affect herbivore growth and performance are not well understood. • We profiled sugars and starch of jasmonate biosynthesis-deficient and jasmonate-insensitive Nicotiana attenuata plants and manipulated leaf carbohydrates through genetic engineering and in vitro complementation to assess how jasmonate-dependent sugar accumulation affects the growth of Manduca sexta caterpillars. • We found that jasmonates reduce the constitutive and herbivore-induced concentration of glucose and fructose in the leaves across different developmental stages. Diurnal, jasmonatedependent inhibition of invertase activity was identified as a likely mechanism for this phenomenon. Contrary to our expectation, both in planta and in vitro approaches showed that the lower sugar concentrations led to increased M. sexta growth. As a consequence, jasmonate- dependent depletion of sugars rendered N. attenuata plants more susceptible to M. sexta attack. • In conclusion, jasmonates are important regulators of leaf carbohydrate accumulation and this determines herbivore growth. Jasmonate-dependent resistance is reduced rather than enhanced through the suppression of glucose and fructose concentrations, which may contribute to the evolution of divergent resistance strategies of plants in nature. [ABSTRACT FROM AUTHOR]

Leaf hydraulics, gas exchange and carbon storage in Pinus edulis and Juniperus monosperma, two tree species on opposite ends of the isohydry-anisohydry spectrum, were analyzed to examine relationships between hydraulic function and carbohydrate dynamics., Leaf hydraulic vulnerability, leaf water potential ( Ψl), leaf hydraulic conductance ( Kleaf), photosynthesis ( A), stomatal conductance ( gs) and nonstructural carbohydrate ( NSC) content were analyzed throughout the growing season., Leaf hydraulic vulnerability was significantly lower in the relatively anisohydric J. monosperma than in the more isohydric P. edulis. In P. edulis, Ψl dropped and stayed below 50% loss of leaf hydraulic conductance ( P50) early in the day during May, August and around midday in September, leading to sustained reductions in Kleaf. In J. monosperma, Ψl dropped below P50 only during August, resulting in the maintenance of Kleaf during much of the growing season. Mean A and gs during September were significantly lower in P. edulis than in J. monosperma. Foliar total NSC was two to three times greater in J. monosperma than in P. edulis in June, August and September., Consistently lower levels of total NSC in P. edulis suggest that its isohydric strategy pushes it towards the exhaustion of carbon reserves during much of the growing season. [ABSTRACT FROM AUTHOR]

Drought affects the carbon (C) source and sink activities of plant organs, with potential consequences for belowground C allocation, a key process of the terrestrial C cycle. The responses of belowground C allocation dynamics to drought are so far poorly understood., We combined experimental rain exclusion with 13C pulse labelling in a mountain meadow to analyse the effects of summer drought on the dynamics of belowground allocation of recently assimilated C and how it is partitioned among different carbohydrate pools and root respiration., Severe soil moisture deficit decreased the ecosystem C uptake and the amounts and velocity of C allocated from shoots to roots. However, the proportion of recently assimilated C translocated belowground remained unaffected by drought. Reduced root respiration, reflecting reduced C demand under drought, was increasingly sustained by C reserves, whilst recent assimilates were preferentially allocated to root storage and an enlarged pool of osmotically active compounds., Our results indicate that under drought conditions the usage of recent photosynthates is shifted from metabolic activity to osmotic adjustment and storage compounds. [ABSTRACT FROM AUTHOR]

In wheat stems, the levels of fructan-dominated water-soluble carbohydrates ( WSC) do not always correlate well with grain yield., Field drought experiments were carried out to further explain this lack of correlation. Wheat ( Triticum aestivum) varieties, Westonia, Kauz and c. 20 genetically diverse double haploid ( DH) lines derived from them were investigated., Substantial genotypic differences in fructan remobilization were found and the 1- FEH w3 gene was shown to be the major contributor in the stem fructan remobilization process based on enzyme activity and gene expression results. A single nucleotide polymorphism ( SNP) was detected in an auxin response element in the 1- FEH w3 promoter region, therefore we speculated that the mutated Westonia allele might affect gene expression and enzyme activity levels. A cleaved amplified polymorphic ( CAP) marker was generated from the SNP. The harvested results showed that the mutated Westonia 1- FEH w3 allele was associated with a higher thousand grain weight ( TGW) under drought conditions in 2011 and 2012., These results indicated that higher gene expression of 1- FEH w3 and 1- FEH w3 mediated enzyme activities that favoured stem WSC remobilization to the grains. The CAP marker residing in the 1- FEH w3 promoter region may facilitate wheat breeding by selecting lines with high stem fructan remobilization capacity under terminal drought. [ABSTRACT FROM AUTHOR]

We explore needle sugar isotopic compositions (δ 18 O and δ 13 C) in boreal Scots pine (Pinus sylvestris) over two growing seasons. A leaf-level dynamic model driven by environmental conditions and based on current understanding of isotope fractionation processes was built to predict δ 18 O and δ 13 C of two hierarchical needle carbohydrate pools, accounting for the needle sugar pool size and the presence of an invariant pinitol pool. Model results agreed well with observed needle water δ 18 O, δ 18 O and δ 13 C of needle water-soluble carbohydrates (sugars + pinitol), and needle sugar δ 13 C (R 2  = 0.95, 0.84, 0.60, 0.73, respectively). Relative humidity (RH) and intercellular to ambient CO 2 concentration ratio (C i /C a ) were the dominant drivers of δ 18 O and δ 13 C variability, respectively. However, the variability of needle sugar δ 18 O and δ 13 C was reduced on diel and intra-seasonal timescales, compared to predictions based on instantaneous RH and C i /C a , due to the large needle sugar pool, which caused the signal formation period to vary seasonally from 2 d to more than 5 d. Furthermore, accounting for a temperature-sensitive biochemical 18 O-fractionation factor and mesophyll resistance in 13 C-discrimination were critical. Interpreting leaf-level isotopic signals requires understanding on time integration caused by mixing in the needle sugar pool., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

The target of rapamycin (TOR) protein kinase is a master regulator of cell growth in all eukaryotes, from unicellular yeast and algae to multicellular animals and plants. Target of rapamycin balances the synthesis and degradation of proteins, lipids, carbohydrates and nucleic acids in response to nutrients, growth factors and cellular energy to promote cell growth. Among nutrients, amino acids (AAs) and glucose are central regulators of TOR activity in evolutionary distant eukaryotes such as mammals, plants and algae. However, these organisms obtain the nutrients through totally different metabolic processes. Although photosynthetic eukaryotes can use atmospheric CO 2 as the sole carbon (C) source for all reactions in the cell, heterotrophic organisms get nutrients from other sources of organic C including glucose. Here, we discuss the impact of autotrophic and heterotrophic metabolism on the nutrient regulation of TOR, focusing on the role of AAs and C sources upstream of this signaling pathway., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)