Nonstructural carbon (NSC) provides the carbon and energy for plant growth and survival. In woody plants, fundamental questions about NSC remain unresolved: Is NSC storage an active or passive process? Do older NSC reserves remain accessible to the plant? How is NSC depletion related to mortality risk? Herein we review conceptual and mathematical models of NSC dynamics, recent observations and experiments at the organismal scale, and advances in plant physiology that have provided a better understanding of the dynamics of woody plant NSC. Plants preferentially use new carbon but can access decade-old carbon when the plant is stressed or physically damaged. In addition to serving as a carbon and energy source, NSC plays important roles in phloem transport, osmoregulation, and cold tolerance, but how plants regulate these competing roles and NSC depletion remains elusive. Moving forward requires greater synthesis of models and data and integration across scales from -omics to ecology. 667

SummaryNonstructural 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.

SummaryThe 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, CO 2 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 CO 2 ) 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 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.

Decomposition of O horizon organic matter made up 20% or less of soil respiration in the younger (o40 years since fire) stands, increasing to $ 50% in mature stands. This is a minimum for total heterotrophic contribution, since mineral soil CO 2 had D 14 C close to or less than those we have assigned to autotrophic respiration. Decomposition of old organic matter in mineral soils clearly contributed to soil respiration in younger stands in 2003, a very dry year, when D 14C of soil respiration in younger successional stands dropped below those of the atmospheric CO 2 .