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1. Decades‐old carbon reserves are widespread among tree species, constrained only by sapwood longevity

   https://doi.org/10.1111/nph.20310  

   Peltier, Drew_M P; Carbone, Mariah S; Ogle, Kiona; Koch, George W; Richardson, Andrew D (February 2025, New Phytologist)

   Summary Carbon reserves are distributed throughout plant cells allowing past photosynthesis to fuel current metabolism. In trees, comparing the radiocarbon (Δ¹⁴C) of reserves to the atmospheric bomb spike can trace reserve ages.We synthesized Δ¹⁴C observations of stem reserves in nine tree species, fitting a new process model of reserve building. We asked how the distribution, mixing, and turnover of reserves vary across trees and species. We also explored how stress (drought and aridity) and disturbance (fire and bark beetles) perturb reserves.Given sufficient sapwood, young (< 1 yr) and old (20–60+ yr) reserves were simultaneously present in single trees, including ‘prebomb’ reserves in two conifers. The process model suggested that most reserves are deeply mixed (30.2 ± 21.7 rings) and then respired (2.7 ± 3.5‐yr turnover time). Disturbance strongly increased Δ¹⁴C mean ages of reserves (+15–35 yr), while drought and aridity effects on mixing and turnover were species‐dependent. Fire recovery inSequoia sempervirensalso appears to involve previously unobserved outward mixing of old reserves.Deep mixing and rapid turnover indicate most photosynthate is rapidly metabolized. Yet ecological variation in reserve ages is enormous, perhaps driven by stress and disturbance. Across species, maximum reserve ages appear primarily constrained by sapwood longevity, and thus old reserves are probably widespread. 

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2. Temporal controls on crown nonstructural carbohydrates in southwestern US tree species

   https://doi.org/10.1093/treephys/tpaa149  

   Peltier, Drew M; Guo, Jessica; Nguyen, Phiyen; Bangs, Michael; Gear, Linnea; Wilson, Michelle; Jefferys, Stacy; Samuels-Crow, Kimberly; Yocom, Larissa L; Liu, Yao; et al (November 2020, Tree Physiology)

   Meinzer, Frederick (Ed.)

   Abstract In trees, large uncertainties remain in how nonstructural carbohydrates (NSCs) respond to variation in water availability in natural, intact ecosystems. Variation in NSC pools reflects temporal fluctuations in supply and demand, as well as physiological coordination across tree organs in ways that differ across species and NSC fractions (e.g., soluble sugars vs starch). Using landscape-scale crown (leaves and twigs) NSC concentration measurements in three foundation tree species (Populus tremuloides, Pinus edulis, Juniperus osteosperma), we evaluated in situ, seasonal variation in NSC responses to moisture stress on three timescales: short-term (via predawn water potential), seasonal (via leaf δ13C) and annual (via current year’s ring width index). Crown NSC responses to moisture stress appeared to depend on hydraulic strategy, where J. osteosperma appears to regulate osmotic potentials (via higher sugar concentrations), P. edulis NSC responses suggest respiratory depletion and P. tremuloides responses were consistent with direct sink limitations. We also show that overly simplistic models can mask seasonal and tissue variation in NSC responses, as well as strong interactions among moisture stress at different timescales. In general, our results suggest large seasonal variation in crown NSC concentrations reflecting the multiple cofunctions of NSCs in plant tissues, including storage, growth and osmotic regulation of hydraulically vulnerable leaves. We emphasize that crown NSC pool size cannot be viewed as a simple physiological metric of stress; in situ NSC dynamics are complex, varying temporally, across species, among NSC fractions and among tissue types. 

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3. Interannual dynamics of stemwood nonstructural carbohydrates in temperate forest trees surrounding drought

   https://doi.org/10.1007/s11676-022-01566-2  

   Blumstein, Meghan J.; Furze, Morgan E. (February 2023, Journal of Forestry Research)

   NA (Ed.)

   Abstract Interactions between water and carbon dynamics underlie drought-related tree mortality. While whole-tree water relations have been shown to play a key role in the response to and recovery from drought, the role of nonstructural carbohydrates (NSC) and how their storage and allocation changes surrounding drought events deserves further attention and is critical for understanding tree survival. Here, we quantified in situ NSC responses of temperate forest trees to the 2016 drought in the northeastern United States. Sugar and starch concentrations were measured in the stemwood of five tree species from 2014 to 2019, which allowed us to monitor NSCs in relation to climatic conditions before, during, and after the natural drought. We found that immediately following the drought, measured stemwood NSC concentrations decreased. However, NSC concentrations rebounded quickly within three years. Notably, trees allocated proportionally more to starch than to sugars following the 2016 drought. In winter 2017, starch comprised 45% of total stemwood stores, whereas starch made up only 1–2% in other years. Further, we modeled and assessed the climatic drivers of total NSC concentrations in the stem. Variation in total NSC concentrations was significantly predicted by the previous year’s temperature, precipitation, and standardized precipitation-evapotranspiration index (SPEI), with stemwood concentrations decreasing following hotter, drier periods and increasing following cooler, wetter periods. Overall, our work provides insight into the climatic drivers of NSC storage and highlights the important role that a tree’s carbon economy may play in its response and recovery to environmental stress. 

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4. Global variation in nonstructural carbohydrate stores in response to climate

   https://doi.org/10.1111/gcb.16573  

   Blumstein, Meghan; Gersony, Jessica; Martínez‐Vilalta, Jordi; Sala, Anna (April 2023, Global Change Biology)

   NA (Ed.)

   Abstract 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. 

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5. Seasonal fluctuation of nonstructural carbohydrates reveals the metabolic availability of stemwood reserves in temperate trees with contrasting wood anatomy

   https://doi.org/10.1093/treephys/tpaa080  

   Furze, Morgan E; Huggett, Brett A; Chamberlain, Catherine J; Wieringa, Molly M; Aubrecht, Donald M; Carbone, Mariah S; Walker, Jennifer C; Xu, Xiaomei; Czimczik, Claudia I; Richardson, Andrew D (June 2020, Tree Physiology)

   Cernusak, Lucas (Ed.)

   Abstract Nonstructural carbohydrates (NSCs) play a critical role in plant physiology and metabolism, yet we know little about their distribution within individual organs such as the stem. This leaves many open questions about whether reserves deep in the stem are metabolically active and available to support functional processes. To gain insight into the availability of reserves, we measured radial patterns of NSCs over the course of a year in the stemwood of temperate trees with contrasting wood anatomy (ring porous vs diffuse porous). In a subset of trees, we estimated the mean age of soluble sugars within and between different organs using the radiocarbon (14C) bomb spike approach. First, we found that NSC concentrations were the highest and most seasonally dynamic in the outermost stemwood segments for both ring-porous and diffuse-porous trees. However, while the seasonal fluctuation of NSCs was dampened in deeper stemwood segments for ring-porous trees, it remained high for diffuse-porous trees. These NSC dynamics align with differences in the proportion of functional sapwood and the arrangement of vessels between ring-porous and diffuse-porous trees. Second, radial patterns of 14C in the stemwood showed that sugars became older when moving toward the pith. The same pattern was found in the coarse roots. Finally, when taken together, our results highlight how the radial distribution and age of NSCs relate to wood anatomy and suggest that while deeper, and likely older, reserves in the stemwood fluctuated across the seasons, the deepest reserves at the center of the stem were not used to support tree metabolism under usual environmental conditions. 

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