Non-structural Carbohydrate Storage Research Articles

Legacy effects of early cotyledon removal on the growth, carbon and nitrogen storage, and drought response of Quercus variabilis seedlings

Predation of cotyledons can reduce short-term survival and growth in emerging oak seedlings, potentially affecting species recruitment. However, the physiological basis of reduced seedling performance after early cotyledon predation and the mid-term consequences of cotyledon predation on seedling drought tolerance remain poorly understood. We performed an experiment where we simulated predation by systematically removing cotyledons from Quercus variabilis seedlings at seven growth stages during emergence in the first growing season. We assessed survival, growth, and nitrogen (N) and non-structural carbohydrate (NSC) storage. Subsequently, in the second growing season, we assessed drought tolerance in seedlings with cotyledons removed at four growth stages in the previous season, and subjected seedlings to two drought levels to assess overall performance. During the first growing season, seedling survival showed less sensitivity to cotyledon removal compared to growth, N and NSC storage, which were significantly reduced when cotyledons were removed within the first 4 days after emergence. Early cotyledon removal (4 days after emergence) in the first growing season enhanced drought tolerance in the second growing season by reducing leaf embolism vulnerability, but it did not affect leaf drought tolerance traits such as the osmotic potential or the modulus of elasticity. Early cotyledon removal had a lasting effect on seedling growth and N and NSC storage building in the second growing season. Additionally, it exacerbated the negative consequences of drought on seedlings growth and N storage building, but it did not influence survival or NSC storage. Our findings evidence that early cotyledon removal in emerging oak seedlings has significant short- and medium-term repercussions on growth and capacity to store N and NSC, and it hinders the capacity of seedlings to respond to drought despite early cotyledon removal increases leaf drought tolerance. These functional legacies can influence oak seedling recruitment.

Residual water losses mediate the trade-off between growth and drought-survival across saplings of 12 tropical rainforest tree species with contrasting hydraulic strategies.

Knowledge of the physiological mechanisms underlying species vulnerability to drought is critical to better understand patterns of tree mortality. Investigating plant adaptive strategies to drought should thus help to fill this knowledge gap, especially in tropical rainforests exhibiting high functional diversity. In a semi-controlled drought experiment on 12 rainforest tree species, we investigated the diversity in hydraulic strategies and whether they determined the ability of saplings to use stored non-structural carbohydrates during an extreme imposed drought. We further explored the importance of water- and carbon-use strategies in relation to drought-survival through a modelling approach. Hydraulic strategies varied considerably across species with a continuum between dehydration- tolerance and -avoidance. During dehydration leading to hydraulic failure and irrespective of hydraulic strategies, species showed strong declines in whole-plant starch concentrations and a maintenance or even an increase in soluble sugar concentrations potentially favouring osmotic adjustments. Residual water losses mediated the trade-off between time to hydraulic failure and growth, suggesting that it is linked to the 'fast-slow' continuum of plant performances and that dehydration avoidance is an effective drought-survival strategy at the sapling stage. Further investigations on residual water losses may be key to understanding the response of tropical rainforest tree communities to climate change.

Seasonal patterns of nonstructural carbohydrate storage and mobilization in two tree species with distinct life-history traits.

Nonstructural carbohydrates (NSC) are essential for tree growth and adaptation, yet our understanding of the seasonal storage and mobilization dynamics of whole-tree NSC is still limited, especially when tree functional types are involved. Here, Quercus acutissima Carruth. and Pinus massoniana Lamb. with distinct life-history traits (i.e., a deciduous broadleaf species vs. an evergreen coniferous species) were studied to assess the size and seasonal fluctuations of organ and whole-tree NSC pools with a focus on comparing differences in carbon resource mobilization patterns between the two species. We sampled the organs (leaf, branch, stem, and root) of the target trees repeatedly over four seasons of the year. Then, NSC concentrations in each organ were paired with biomass estimates from the allometric model to generate whole-tree NSC pools. The seasonal dynamics of the whole-tree NSC of Q. acutissima and P. massoniana reached the peak in autumn and summer, respectively. The starch pools of the two species were supplemented in the growing season while the soluble sugar pools were the largest in the dormant season. Seasonal dynamics of organ-level NSC concentrations and pools were affected by organ type and tree species, with above-ground organs generally increasing during the growing season and P. massoniana roots decreasing during the growing season. In addition, the whole-tree NSC pools of P. massoniana were larger but Q. acutissima showed larger seasonal fluctuations, indicating that larger storage was not associated with more pronounced seasonal fluctuations. We also found that the branch and root were the most dynamic organs of Q. acutissima and P. massoniana, respectively, and were the major suppliers of NSC to support tree growth activities. These results provide fundamental insights into the dynamics and mobilization patterns of NSC at the whole-tree level, and have important implications for investigating environmental adaptions of different tree functional types.

Climate warming could free cold-adapted trees from C-conservative allocation strategy of storage over growth.

Carbon allocation has been fundamental for long-lived trees to survive cold stress at their upper elevation range limit. Although carbon allocation between non-structural carbohydrate (NSC) storage and structural growth is well-documented, it still remains unclear how ongoing climate warming influences these processes, particularly whether these two processes will shift in parallel or respond divergently to warming. Using a combination of an insitu downward-transplant warming experiment and an ex situ chamber warming treatment, we investigated how subalpine fir trees at their upper elevation limit coordinated carbon allocation priority among different sinks (e.g., NSC storage and structural growth) at whole-tree level in response to elevated temperature. We found that transplanted individuals from the upper elevation limit to lower elevations generally induced an increase in specific leaf area, but there was no detected evidence of warming effect on leaf-level saturated photosynthetic rates. Additionally, our results challenged the expectation that climate warming will accelerate structural carbon accumulation while maintaining NSC constant. Instead, individuals favored allocating available carbon to NSC storage over structural growth after 1 year of warming, despite the amplification in total biomass encouraged by both insitu and ex situ experimental warming. Unexpectedly, continued warming drove a regime shift in carbon allocation priority, which was manifested in the increase of NSC storage in synchrony to structural growth enhancement. These findings imply that climate warming would release trees at their cold edge from C-conservative allocation strategy of storage over structural growth. Thus, understanding the strategical regulation of the carbon allocation priority and the distinctive function of carbon sink components is of great implication for predicting tree fate in the future climate warming.

Starch depletion in the xylem and phloem ray parenchyma of grapevine stems under drought.

While nonstructural carbohydrate (NSC) storage can support long-lived woody plants during abiotic stress, the timing and extent of their use are less understood, as are the thresholds for cell mortality as NSCs and water supplies are consumed. Here, we combine physiological and imaging tools to study the response of Vitis riparia to a 6-week experimental drought. We focused on the spatial and temporal dynamics of starch consumption and cell viability in the xylem and phloem of the stem. Starch dynamics were further corroborated with enzymatic starch digestion and X-ray microcomputed tomography imaging. Starch depletion in the stems of droughted plants was detected after 2 weeks and continued over time. We observed distinct differences in starch content and cell viability in the xylem and phloem. By the end of the drought, nearly all the starch was consumed in the phloem ray parenchyma (98 % decrease), and there were almost no metabolically active cells in the phloem. In contrast, less starch was consumed in the xylem ray parenchyma (30 % decrease), and metabolically active cells remained in the ray and vessel-associated parenchyma in the xylem. Our data suggest that the higher proportion of living cells in the phloem and cambium, combined with smaller potential NSC storage area, rapidly depleted starch, which led to cell death. In contrast, the larger cross-sectional area of the xylem ray parenchyma with higher NSC storage and lower metabolically active cell populations depleted starch at a slower pace. Why NSC source-sink relationships between xylem and phloem do not allow for a more uniform depletion of starch in ray parenchyma over time is unclear. Our data help to pinpoint the proximate and ultimate causes of plant death during prolonged drought exposure and highlight the need to consider the influence of within-organ starch dynamics and cell mortality on abiotic stress response.

Is the amount of parenchyma in storage organs a good predictor of non-structural carbohydrate storage in herbs?

Is the amount of parenchyma in storage organs a good predictor of non-structural carbohydrate storage in herbs?

Tropical forest lianas have greater non-structural carbohydrate concentrations in the stem xylem than trees.

Lianas (woody vines) are important components of tropical forests and are known to compete with host trees for resources, decrease tree growth and increase tree mortality. Given the observed increases in liana abundance in some forests and their impacts on forest function, an integrated understanding of carbon dynamics of lianas and liana-infested host trees is critical for improved prediction of tropical forest responses to climate change. Non-structural carbohydrates (NSC) are the main substrate for plant metabolism (e.g., growth, respiration), and have been implicated in enabling tree survival under environmental stress, but little is known of how they vary among life-forms or of how liana infestation impacts host tree NSC. We quantified stem total NSC (NSC) concentrations and its fractions (starch and soluble sugars) in trees without liana infestation, trees with more than 50% of the canopy covered by lianas, and the lianas infesting those trees. We hypothesized that i) liana infestation depletes NSC storage in host trees by reducing carbon assimilation due to competition for resources; ii) trees and lianas, which greatly differ in functional traits related to water transport and carbon uptake, would also have large differences in NSC storage, and that As water availability has a significant role in NSC dynamics of Amazonian tree species, we tested these hypotheses within a moist site in western Amazonia and a drier forest site in southern Amazonia. We did not find any difference in NSC, starch or soluble sugar concentrations between infested and non-infested trees, in either site. This result suggests that negative liana impact on trees may be mediated through mechanisms other than depletion of host tree NSC concentrations. We found lianas have higher stem NSC and starch than trees in both sites. The consistent differences in starch concentrations, a long term NSC reserve, between life forms across sites reflect differences in carbon gain and use of lianas and trees. Soluble sugar concentrations were higher in lianas than in trees in the moist site but indistinguishable between life forms in the dry site. The lack of difference in soluble sugars between trees and lianas in the dry site emphasize the importance of this NSC fraction for plant metabolism of plants occurring in water limited environments. Abstract in Portuguese and Spanish are available in the supplementary material.

Functional trade-offs are driven by coordinated changes among cell types in the wood of angiosperm trees from different climates.

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 (Kth ) 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 Kth 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.

Carbon starvation following a decade of experimental drought consumes old reserves in Pinus edulis.

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 (Δ14 C) 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.

Stem Carbohydrate Richness in Two Cycad Species

The pachycaulous stem of cycad plants enables the storage of abundant nonstructural carbohydrates. Cycas revoluta Thunb. and Zamia furfuracea L.f. stems were analyzed for starch and 15 sugars to determine carbohydrate richness. All 15 sugars were detected in both species. The tetrasaccharide stachyose and the disaccharide sucrose comprised most of the sugar content. Total sugar content of these cycad stems was 330 to 360 mg·g−1 and was similar to starch content. The stems of these two species were composed of 64% to 79% nonstructural carbohydrates. The cycad stem is ideally designed to store and mobilize nonstructural carbohydrates to support sink activity when needed, and stachyose may play a prominent functional role.

Correction to: Whole-Tree Storage of Non-structural Carbohydrates in Apple and Pear Trees on Size-Controlling Rootstocks

Correction to: Whole-Tree Storage of Non-structural Carbohydrates in Apple and Pear Trees on Size-Controlling Rootstocks

Moisture stress limits radial mixing of non-structural carbohydrates in sapwood of trembling aspen.

Dynamics in non-structural carbohydrate (NSC) pools may underlie observed drought legacies in tree growth. We assessed how aridity influences the dynamics of different-aged NSC pools in tree sapwood at two sites with differing climate conditions ('wet' vs. 'dry') that also experienced widespread regional drought five years earlier. We used an incubation method to measure radiocarbon (Δ14C) in CO2 respired from Populus tremuloides (aspen) tree rings to evaluate NSC storage and mixing patterns, coupled with measurements of NSC (soluble sugars, starch) concentrations and respired δ13C-CO2. At a wet site, CO2 respired from rings formed during 1962-1967 was only ~ 11years old, suggesting deep sapwood mixing of NSCs as starch. At a dry site, total NSC was about one-third of wet site totals, maximum ages in deep rings were lower, and ages more rapidly increased in shallow rings then plateaued. These results suggest historically shallower mixing and/or relatively higher consumption of NSCs under dry conditions. Both sites, however, had similar aged NSC (<1yr) in the most recent six rings, indicative of deep radial mixing following relatively wet conditions during the sampling year. We suggest significant differences in NSC mixing among sites are driven by moisture stress, where aridity reduces NSC reserves and restricts the depth of radial mixing. However, dynamic climate conditions in the southwestern US resulted in more complex radial patterns of sapwood NSC age than previously described. We suggest a novel conceptual framework to understand how moisture variability might influence the dynamics of NSC mixing in the sapwood.

Whole-Tree Storage of Non-structural Carbohydrates in Apple and Pear Trees on Size-Controlling Rootstocks

Whole-Tree Storage of Non-structural Carbohydrates in Apple and Pear Trees on Size-Controlling Rootstocks

Effects of NSC in different organs and at different growth stages on the yield of oil peony Fengdan with different ages.

Non-structural carbohydrates (NSC) as resource reserves of plants play important roles in energy supply for normal growth and reproduction under environmental stress. The yield of perennial crops is mainly determined by the carbohydrate production and allocation in the current growth season, as well as the re-allocation of NSC reserves. However, the contribution of NSC to crop yield has not been fully determined. Fengdan (Paeonia ostii) is a variety of oil Peony that is newly developed in China. The effects of tree age and NSC on yield were examined by investigated the variations of biomass, soluble sugars, starch, and NSC in the organ and whole tree levels in the dormant and ripening stages of Fengdan populations with 4-, 6-, and 8-year-old in 2020 and 5-, 7-, and 9- year old in 2021. Results showed that the biomass, yield (seed biomass), soluble sugars, starch, and NSC reserve of Fengdan at the whole tree level increased with the increase in age. Although consistent correlations were observed between soluble sugars, starch and NSC storage, and yield among the plants with different ages, Fengdan showed allometric growth relationships between the accumulation of soluble sugars, starch, and NSC and yield and biomass (standardized major axis analyses slope b ≠ 1). Tree age significantly affected biomass and its allocation and NSC levels, especially the yield of Fengdan plants. The results of the investigation of the variations in the relationships between the yield and seasonal fluctuations of NSC and biomass indicate that roots is the key storage structure, whereas stems both serve as sink and/or source functions for the adult (7-9a) plants. NSC level, particularly the concentration of soluble sugars, in stems mainly influences Fengdan yield. These findings together provide new insights into the mechanisms underlying the yield formation of Fengdan and have implications for manipulating sink-source relationship to achieve high yield.

Effect of Ectomycorrhizal Fungi on the Drought Resistance of Pinus massoniana Seedlings.

Studies on the dynamics of non-structural carbohydrates (NSCs) play an important role in understanding the mechanisms of plant responses to drought stress. The objective of this study was to assess the influence of ectomycorrhizal fungi (ECMF) on the content and distribution of NSCs in Pinus massoniana seedlings under different drought intensities and to further explore the possible mechanism by which ECMF enhances the stress resistance of host plants. We conducted a pot experiment using P. massoniana seedlings that were inoculated (M) or non-inoculated (NM) with Suillus luteus (Sl) under well-watered, moderate, and severe drought stress conditions. The results showed that drought significantly reduced the photosynthetic capacity of P. massoniana seedlings and inhibited their growth rate. P. massoniana could respond to different degrees of drought stress by increasing the accumulation of NSCs and increasing WUE. However, compared with well-watered treatment, NSCs consumption began to appear in the roots of NM due to the decrease in starch content under severe drought, whereas NSCs content in M seedlings was higher than that in the well-watered treatment, showing that the ability to maintain C balance was higher in M seedlings. Compared with NM, inoculation with Sl increased the growth rate and biomass of roots, stems, and leaves under moderate and severe drought. In addition, Sl can also improve the gas exchange parameters (net photosynthetic rate, transpiration rate, intercellular CO2 concentration and stomatal conductance) of P. massoniana seedlings compared with NM seedlings, which was conducive to the hydraulic regulation of seedlings and improved their C fixation capacity. Meanwhile, the content of NSCs in M seedlings was higher. Moreover, the soluble sugar content and SS/St ratio of leaves, roots, and whole plants were higher under drought stress after Sl inoculation, indicating that Sl could also change the C distribution mode, regulate more soluble sugar to respond to drought stress, which was conducive to improving the osmotic adjustment ability of seedlings, and providing more available C sources for plant growth and defense. Overall, inoculation with Sl could enhance the drought resistance of seedlings and promote their growth under drought stress by improving NSCs storage, increasing soluble sugar distribution, and improving the plant water balance of P. massoniana seedlings.

The Importance of Initial Seedling Characteristics in Controlling Allocation to Growth and Reserves under Different Soil Moisture Conditions

After disturbance, forest regeneration and resiliency depend on the ability of seedlings to respond, survive, and grow under a variety of stress conditions, including drought. Despite recent efforts to improve our fundamental knowledge surrounding plant response mechanisms to stress and their application in seedling quality research, initial seedling characteristics are often ignored when exploring seedling responses to stress in field plantings or ecophysiological studies. Here, we explore how initial differences in size, biomass allocation, and non-structural carbohydrate (NSC) storage affect the subsequent partitioning of new biomass, growth potential, and drought response in seedlings of a deciduous broad-leaved (Populus tremuloides) and an evergreen coniferous species (Pinus banksiana). We exposed seedlings of both species to different growing conditions in their first growing season in order to manipulate the aforementioned seedling characteristics. In a second growing season, we exposed these different seedling types to a subsequent drought stress. While drought reduced both structural growth and NSC storage in all seedling types, the expected shift in allocation favoring roots was only observed in seedling types with initially low root:shoot or root:stem ratios. Overall, we also found that the traits associated with greater growth were quite different between pine and aspen. While larger seedlings led to greater growth in pine, it was the smallest seedling type in aspen with the largest root:stem ratio that produced the most new growth. In aspen, this smaller seedling type was the only one that did not undergo a shift in biomass relative to its initial allometry, suggesting that adjustments in biomass allocation made by other, larger seedling types must have come at the cost of lower growth. In contrast, adjustments in allocation did not appear to negatively impact pine, possibly because reduced root:shoot ratios of larger seedlings did not reduce NSC storage, as it did in aspen. Our results highlight (1) the complexity of how differences in biomass allocation and changes in seedling size may alter storage and the response of species to drought, and (2) the importance of accounting for initial seedling characteristics (both morphological and physiological) when predicting seedling growth and the impacts of environmental stressors.

Plastic and adaptive response of carbon allocation to temperature change in alpine treeline trees

Understanding how tree individuals allocate nonstructural carbohydrate (NSC) between storage and growth is key to reveal the mechanism of alpine treeline formation as well as its response to climate warming. Yet, little is known about whether carbon allocation of treeline trees responds to temperature change by means of phenotypic plasticity and/or ecotypic adaptation. Here, we employed a reciprocal transplant experiment and a growth chamber warming treatment to explore the effects of temperature change on functional traits and carbon allocation between NSC storage and growth in treeline species Larix chinensis. In the reciprocal transplant experiment, we observed that the individuals grown at high elevation always displayed relatively lower specific leaf area (SLA) than those grown at lower elevations irrespective of species provenance, demonstrating the phenotypic plasticity of functional traits in relation to elevation. Higher growth rate and late-season NSC concentration in the individuals at their provenance elevation than at the elevations away from home, indicated that trees were locally adapted to their home environments in terms of NSC storage and growth. Additionally, individuals from high-elevation provenance showed considerably less phenotypic plasticity in carbon allocation (i.e., the ratio of NSC storage to growth) in response to temperature change compared to those from low-elevation provenance. In the chamber experiment, warming greatly accelerated shoot growth but not accompanied by a down-regulation in NSC storage in cold-edge trees, resulting in a lower ratio of NSC storage to growth. It was noteworthy that there was a time lag in variations of carbon allocation, especially in variations of NSC storage when cold-edge trees were grown in warmed climates. Our findings demonstrate that the responses of NSC storage and growth to temperature change might be a combination of short-term plastic and long-term adaptive reactions, and suggest a limited response of cold-edge populations in carbon allocation to climate warming.

Effects of Environmental Factors on the Nonstructural Carbohydrates in Larix principis-rupprechtii

Nonstructural carbohydrates (NSCs) reflect the carbon balance in trees and play a central role in their responses to environmental stresses. However, our understanding of tree NSC storage dynamics under environmental stress is unclear. Here, we quantified the deciduous conifer Larix principis-rupprechtii at five altitudes on Luya Mountain, North-Central China. We measured NSCs in different organs and simultaneously monitored environmental variables during the growing seasons in 2018 and 2019. The results showed that temperature was the main environmental factor affecting tree NSCs. Furthermore, as the soil–atmosphere temperature gradient decreased, the concentrations of soluble sugars in the needles increased while their starch decreased, as well as the concentrations of starch in the shoots, stems, and roots increased. These results may indicate that temperature gradients may assist carbohydrate allocation within individual trees. The NSCs, especially starch, in the needles increased with lower soil moisture, and the soluble sugars and starch in the needles and shoots increased with a higher vapor pressure deficit. The soluble sugars in the stems were significantly affected by soil moisture, and the NSCs in the roots were mainly affected by temperature. These results provide new insights to improve our understanding of the environmental stress effects on future forest C balances.

Tree-ring isotopes from the Swiss Alps reveal non-climatic fingerprints of cyclic insect population outbreaks over the past 700years.

Recent experiments have underlined the potential of δ2H in tree-ring cellulose as a physiological indicator of shifts in autotrophic versus heterotrophic processes (i.e., the use of fresh versus stored non-structural carbohydrates). However, the impact of these processes has not yet been quantified under natural conditions. Defoliator outbreaks disrupt tree functioning and carbon assimilation, stimulating remobilization, therefore providing a unique opportunity to improve our understanding of changes in δ2H. By exploring a 700-year tree-ring isotope chronology from Switzerland, we assessed the impact of 79 larch budmoth (LBM, Zeiraphera griseana [Hübner]) outbreaks on the growth of its host tree species, Larix decidua[Mill]. The LBM outbreaks significantly altered the tree-ring isotopic signature, creating a 2H-enrichment and an 18O- and 13C-depletion. Changes in tree physiological functioning in outbreak years are shown by the decoupling of δ2H and δ18O (O-H relationship), in contrast to the positive correlation in non-outbreak years. Across the centuries, the O-H relationship in outbreak years was not significantly affected by temperature, indicating that non-climatic physiological processes dominate over climate in determining δ2H. We conclude that the combination of these isotopic parameters can serve as a metric for assessing changes in physiological mechanisms over time.

Global variation in nonstructural carbohydrate stores in response to climate.

Woody plant species store nonstructural carbohydrates (NSCs) for many functions. While known to buffer against fluctuations in photosynthetic supply, such as at night, NSC stores are also thought to buffer against environmental extremes, such as drought or freezing temperatures by serving as either back-up energy reserves or osmolytes. However, a clear picture of how NSCs are shaped by climate is still lacking. Here, we update and leverage a unique global database of seasonal NSC storage measurements to examine whether maximum total NSC stores and the amount of soluble sugars are associated with clinal patterns in low temperatures or aridity, indicating they may confer a benefit under freezing or drought conditions. We examine patterns using the average climate at each study site and the unique climatic conditions at the time and place in which the sample was taken. Altogether, our results support the idea that NSC stores act as critical osmolytes. Soluble Sugars increase with both colder and drier conditions in aboveground tissues, indicating they can plastically increase a plants' tolerance of cold or arid conditions. However, maximum total NSCs increased, rather than decreased, with average site temperature and had no relationship to average site aridity. This result suggests that the total amount of NSC a plant stores may be more strongly determined by its capacity to assimilate carbon than by environmental stress. Thus, NSCs are unlikely to serve as reservoir of energy. This study is the most comprehensive synthesis to date of global NSC variation in relation to climate and supports the idea that NSC stores likely serve as buffers against environmental stress. By clarifying their role in cold and drought tolerance, we improve our ability to predict plant response to environment.