Dynamics Of Non-structural Carbohydrates Research Articles

Effects of climate and soil conditions on the productivity and defence capacity of Picea abies in Sweden—An ecosystem model assessment

Climate change can lead to an increased frequency of extreme weather events, which may induce a decline in tree vitality, rendering the forest more vulnerable to other stress factors. In this study we used the ecosystem model LPJ-GUESS to compare the NPP of boreal conifers on till soil and sandy soil, simulating the growth of Norway spruce during the 21st century using climate model projections corresponding to RCP 8.5. Model runs with and without nitrogen cycling enabled, in combination with two different approaches to implement the dynamics of non-structural carbohydrates (NSC) were compared. The simulations showed that the forest productivity in Sweden is influenced by an interplay between site-specific temperature conditions and precipitation. Local differences in soil conditions can have an impact on the productivity, mediated by soil water and nutrient content. A warmer climate will lead to a longer growing season, but the productivity increase will partly be offset by a higher frequency of drought stress in summer and higher autotrophic respiration in winter. The ecosystem model runs indicated that periods with drought stress can cause low NSC levels, leading to time periods with decreased defence capacity against attacks by secondary agents such as bark beetles. The risks may however not become apparent until the forest stands are middle-aged. The model results are discussed in relation to current forestry practice, with Norway spruce (Picea abies) commonly being planted on dry soils better suited for Scots pine (Pinus sylvestris) in order to reduce moose browsing damage of seedlings, which in the longer term may create a forest landscape more sensitive to spruce bark beetle attacks.

Non-Structural Carbohydrate Dynamics in Leaves and Branches of Pinus massoniana (Lamb.) Following 3-Year Rainfall Exclusion

Drought-induced tree mortality is an increasing and global ecological problem. Stored non-structural carbohydrates (NSCs) may be a key determinant of drought resistance, but most existing studies are temporally limited. In this study, a 3-year 100% rainfall exclusion manipulation experiment was conducted to evaluate the response of NSC dynamics to drought stress in 25-year-old Pinus massoniana leaves and branches. The results showed: (1) compared with the control condition, leaf NSC concentration in the drought treatment increased 90% in the early stage (days 115–542) (p < 0.05), and then decreased 15% in the late stage (days 542–1032), which was attributed to water limitation instead of phenology; (2) the response of leaf NSCs to drought was more significant than branch NSCs, demonstrating a time lag effect; and (3) the response of P. massoniana to mild drought stress was to increase the soluble sugars and starch in the early stage, followed by an increase in soluble sugars caused by decreasing starch in the later stress period. Considering these results, mid-term drought stress had no significant effect on the total NSC concentration in P. massoniana, removing carbon storage as a potential adaptation to drought stress.

Non-structural carbohydrate dynamics associated with drought-induced die-off in woody species of a shrubland community.

The relationship between plant carbon economy and drought responses of co-occurring woody species can be assessed by comparing carbohydrate (C) dynamics following drought and rain periods, relating these dynamics to species' functional traits. We studied nine woody species coexisting in a continental Mediterranean shrubland that experienced severe drought effects followed by rain. We measured total non-structural carbohydrates (NSC) and soluble sugars (SS) in roots and stems during drought and after an autumn rain pulse in plants exhibiting leaf loss and in undefoliated ones. We explored whether their dynamics were related to foliage recovery and functional traits (height [H], specific leaf area [SLA], wood density [WD]). During drought, NSC concentrations were overall lower in stems and roots of plants experiencing leaf loss, while SS decreases were smaller. Roots had higher NSC concentrations than stems. After the rain, NSC concentrations continued to decrease, while SS increased. Green foliage recovered after rain, particularly in plants previously experiencing higher leaf loss, independently of NSC concentrations during drought. Species with lower WD tended to have more SS during drought and lower SS increases after rain. In low-WD species, plants with severe leaf loss had lower NSC relative to undefoliated ones. No significant relationship was found between H or SLA and C content or dynamics. Our community-level study reveals that, while responses were species-specific, C stocks overall diminished in plants affected by prolonged drought and did not increase after a pulse of seasonal rain. Dynamics were faster for SS than NSC. We found limited depletion of SS, consistent with their role in basal metabolic, transport and signalling functions. In a scenario of increased drought under climate change, NSC stocks in woody plants are expected to decrease differentially in coexisting species, with potential implications for their adaptive abilities and community dynamics.

Non-structural carbohydrates characteristics of different forest layers in monsoon broad-leaved evergreen forest in Pu'er, Yunnan Province, China

Based on cluster analysis of species vertical stratification, the variations in concentration, allocation and seasonal dynamics of non-structural carbohydrates were investigated in a monsoon broad-leaved evergreen forest in Pu'er, Yunnan Province. The results showed that the concentrations of soluble sugar (3.9%) and nonstructural carbohydrates (NSC, 13.3%) were the highest in the sub-canopy. In contrast, the sugar/starch (0.76) of understory was the lowest. There was no significant difference in starch content among three forest layers. For all forest stories, the soluble sugar was mainly allocated to leaves and the roots had more starch and NSC. The soluble sugar contents in leaves and trunks of sub-canopy species were higher than canopy and understory species. There were no significant differences in soluble sugar content of twigs and roots among three different forest layers. The starch content of leaves increased with decreasing height of forest layers. In contrast, the lowest starch content (10.7%) of roots was in understory. There was no significant difference in starch content of twigs and trunks among three different forest layers. The NSC content of leaves was lower in canopy (10.7%) than that in sub-canopy (12.3%) and understory (12.0%). The lowest NSC content of roots was in understory (14.2%). The lowest sugar/starch of leaves, twigs, and trunks presented in understory, while the lowest sugar/starch of roots presented in canopy (0.79). There were dramatic seasonal variations in concentrations and components of NSC. The concentration of soluble sugar and the sugar/starch were higher in the rainy season than that in the dry season. However, the concentration of starch and NSC were higher in the dry season than in the rainy season. The variations in concentration of NSC and its components among forest layers indicated that species with different heights varied in their utilization strategies of carbon, which partly explained species coexistence.

Carbohydrate dynamics of three dominant species in a Chinese savanna under precipitation exclusion.

The potential impact of drought on the carbon balance in plants has gained great attention. Non-structural carbohydrate (NSC) dynamics have been suggested as an important trait reflecting carbon balance under drought conditions. However, NSC dynamics under drought and the response mechanisms of NSC to drought remain unclear, especially in water-limited savanna ecosystems. A precipitation exclusion experiment was performed to simulate different drought intensities in a savanna ecosystem in Yuanjiang valley in southwestern China. Growth, total NSC concentration and diurnal change of NSC were determined for the leaves and non-photosynthetic organs of three dominant species (Lannea coromandelica, Polyalthia cerasoides and Heteropogon contortus) throughout the growing season. Drought significantly reduced the growth of all the three species. Total NSC concentration averaged ~8.1%, varying with species, organ and sampling period, and did not significantly decrease under drought stress. By contrast, the diurnal change of NSC in these three species increased under drought stress. These results indicate that these three dominant species did not undergo carbon limitation. Thus, relative change in NSC is a more sensitive and effective indicator than carbon reserves in evaluation of plant carbon balance. These findings provide new insights for the understanding of carbon balance and the mechanisms of carbon starvation.

Are storage and tree growth related? Seasonal nutrient and carbohydrate dynamics in evergreen and deciduous Mediterranean oaks

Seasonal dynamics of branch carbohydrates differed sharply between coexisting evergreen and deciduous Mediterranean oaks. Branch carbon storage was crucial in the evergreen, while it played a minor role in the deciduous oaks. The aim of this study was to describe the seasonal dynamics of nutrients and NSC in relation to the aboveground phenology of coexisting winter-deciduous (Quercus faginea) and evergreen (Quercus ilex subsp. ballota) oak species, and to analyse the relationship between the resource budget of branches and shoot growth. Monthly concentrations of nitrogen (N), phosphorus (P), potassium (K) and non-structural carbohydrates (NSC) plus the aboveground phenology of branches were measured over 2 years. We also analysed the correlation between the resource budget of branches prior to the growing season and the subsequent shoot and stem growth. Seasonal branch nutrient dynamics could be explained by shoot growth phenology, showing similar patterns across species. However, NSC dynamics varied between the two species, owing to the differences in leaf phenology and the contrasting role of branches as storage sites. NSC and N branch storage were crucial for the early stages of shoot growth in the evergreen trees. Accordingly, branch N and NSC storage pools in late winter correlated positively with spring growth, and NSC concentrations dropped during bud burst in all branch organs of Q. ilex. Contrastingly, branch NSC concentrations of the deciduous Q. faginea were only marginally affected by spring growth and no relationship was observed between branch N and NSC stores prior to bud burst. These results challenge previous assumptions on the lower relevance of branch NSC storage for the spring growth of evergreen trees and call for further studies where closely related pairs of coexisting evergreen and deciduous tree species are compared.

Living on next to nothing: tree seedlings can survive weeks with very low carbohydrate concentrations.

The usage of nonstructural carbohydrates (NSCs) to indicate carbon (C) limitation in trees requires knowledge of the minimum tissue NSC concentrations at lethal C starvation, and the NSC dynamics during and after severe C limitation. We completely darkened and subsequently released seedlings of two deciduous and two evergreen temperate tree species for varying periods. NSCs were measured in all major organs, allowing assessment of whole-seedling NSC balances. NSCs decreased fast in darkness, but seedlings survived species-specific whole-seedling starch concentrations as low as 0.4-0.8% per dry matter (DM), and sugar (sucrose, glucose and fructose) concentrations as low as 0.5-2.0% DM. After re-illumination, the refilling of NSC pools began within 3wk, while the resumption of growth was delayed or restricted. All seedlings had died after 12wk of darkness, and starch and sugar concentrations in most tissues were lower than 1% DM. We conclude that under the applied conditions, tree seedlings can survive several weeks with very low NSC reserves probably also using alternative C sources like lipids, proteins or hemicelluloses; lethal C starvation cannot be assumed, if NSC concentrations are higher than the minimum concentrations found in surviving seedlings; and NSC reformation after re-illumination occurs preferentially over growth.

Unfolding Non-structural Carbohydrates from Sapling to Dying Black Locust on China’s Loess Plateau

Non-structural carbohydrates (NSC) play important roles in metabolic processes of plants and represent important functional traits in plant adaptation to the external environment. However, there are few reports concerning intra-annual NSC distribution in temperate deciduous forests, especially for the purpose of comparison among different stand ages on China’s Loess Plateau. Here, NSC allocation dynamics with age ring-porous black locust tree was determined for the growing (May) and dormant (November) seasons over the period from sapling to dying trees—defined as a completely defoliated tree with dried branches in the growing season. It was noted that regardless of tree age, NSC concentration was highest in coarse roots [16.4 g per 100 g of dry mass (16.4% DM)] and stems (15.1% DM). At the tree level, NSC concentration was highest (14.3% DM) in a 30-year-old stand in November and lowest (4.1% DM) in dying stands in May. The pool of NSC at tree level was highest (25.2 kg DM per tree) in 30-year-old stands in November and lowest (0.13 kg DM per tree) in sapling stands in May. The concentration of NSC was significantly lower in May than in November for all tree ages, organs, and biochemical components. The results underscored the importance of NSC in plant growth on China’s Loess Plateau. It also provided a useful insight into the dynamics of NSC from sapling to dying broadleaved tree species.

Teresa Rosas.

Teresa Rosas.

Tree-killing aphid dramatically reduces bark contents in carbohydrates and nitrogen compounds

Abstract Aphids can affect the growth or architecture of their host-trees, but rarely challenge their survival. Nonetheless, the woolly poplar aphid, Phloeomyzus passerinii, can kill healthy, mature poplars during outbreaks. Trees fail to open their buds and flush their leaves in the spring following a severe infestation. This insect develops on poplar trunks, where it induces galls which accumulate amino acids and deplete starch content in the bark tissues. Here, we hypothesized that an infestation may then reduce non-structural carbohydrates (NSC) and non-structural nitrogen compounds (NSNC) contents in the bark, and that the reduction would be positively correlated with the infestation duration. This would in turn compromise the spring reactivation of trees. Consequently we also hypothesized that the bark contents of NSC and NSNC could be positively correlated with the amount of branches exhibiting bud break in spring, and the subsequent radial growth. The bark contents of NSC and NSNC were quantified in autumn for poplars exhibiting different levels of infestation and located in two distant stands. In one stand, NSC and NSNC contents in the bark were also quantified in the following spring and the damage level in the crowns were evaluated. In both stands, the infestation dramatically reduced the NSC and NSNC contents in autumn, especially starch and protein contents. While the drop in protein content was not related to infestation duration, the starch content gradually decreased when infestation duration increased. The infestation effect on starch content was still detectible in spring, after reserves had likely been mobilized for the spring reactivation. Starch content in autumn and spring was strongly positively correlated with crown health and radial increment. These results suggest that poplars infested by P. passerinii may die from carbon reserves deficiency, which would be an unprecedented situation for tree-aphid and tree-gall inducing insects interactions. Further experiments monitoring seasonal dynamics of NSC and NSNC reserves in different organs of infested trees should be conducted to validate this hypothesis. The results also suggest that the starch content in the bark of infested trees, in autumn, might be used as a predictor of tree decay by managers.

The fate of recently fixed carbon after drought release: towards unravelling C storage regulation in Tilia platyphyllos and Pinus sylvestris.

Carbon reserves are important for maintaining tree function during and after stress. Increasing tree mortality driven by drought globally has renewed the interest in how plants regulate allocation of recently fixed C to reserve formation. Three-year-old seedlings of two species (Tilia platyphyllos and Pinus sylvestris) were exposed to two intensities of experimental drought during ~10weeks, and 13 C pulse labelling was subsequently applied with rewetting. Tracking the 13 C label across different organs and C compounds (soluble sugars, starch, myo-inositol, lipids and cellulose), together with the monitoring of gas exchange and C mass balances over time, allowed for the identification of variations in C allocation priorities and tree C balances that are associated with drought effects and subsequent drought release. The results demonstrate that soluble sugars accumulated in P.sylvestris under drought conditions independently of growth trends; thus, non-structural carbohydrates (NSC) formation cannot be simply considered a passive overflow process in this species. Once drought ceased, C allocation to storage was still prioritized at the expense of growth, which suggested the presence of 'drought memory effects', possibly to ensure future growth and survival. On the contrary, NSC and growth dynamics in T.platyphyllos were consistent with a passive (overflow) view of NSC formation.

Seasonal changes in above- and below-ground non-structural carbohydrates (NSC) in Spartina alterniflora in a marsh in Georgia, USA

Spartina alterniflora is the dominant grass appearing in salt marshes along the east coast of the USA. The development of predictive, mechanistic models of Spartina has been hindered by the lack of information on below-ground biomass and its dynamics, and in particular the storage of resources that can be used for spring re-growth. We studied the dynamics of non-structural carbohydrates (glucose, fructose, sucrose, and starch) and biomass in 8 different above- and below-ground tissues in S. alterniflora over the course of a year in a salt marsh on Sapelo Island, Georgia, USA. We found greater seasonal variability in non-structural carbohydrates in S. alterniflora than had been previously reported, with concentrations varying between 3.3% through 17.3% of the total biomass and between 0% and 19.5% of dry weight depending on the type of tissue, with statistical differences between the different tissues. We found that sucrose was the dominant non-structural carbohydrate in above- and below-ground tissue, and that this sugar was likely used for long-term storage during winter months and as a resource for early spring growth. Glucose, fructose, and starch showed less variability, with glucose following changes in above-ground biomass more closely indicating their use as short-term storage. We were unable to develop a coherent carbon budget for the plants largely because of uncertainties in modeled net primary production and heterogeneity in below-ground biomass.

Dynamic allocation and transfer of non-structural carbohydrates, a possible mechanism for the explosive growth of Moso bamboo (Phyllostachys heterocycla).

Moso bamboo can rapidly complete its growth in both height and diameter within only 35–40 days after shoot emergence. However, the underlying mechanism for this “explosive growth” remains poorly understood. We investigated the dynamics of non-structural carbohydrates (NSCs) in shoots and attached mature bamboos over a 20-month period. The results showed that Moso bamboos rapidly completed their height and diameter growth within 38 days. At the same time, attached mature bamboos transferred almost all the NSCs of their leaves, branches, and especially trunks and rhizomes to the “explosively growing” shoots via underground rhizomes for the structural growth and metabolism of shoots. Approximately 4 months after shoot emergence, this transfer stopped when the leaves of the young bamboos could independently provide enough photoassimilates to meet the carbon demands of the young bamboos. During this period, the NSC content of the leaves, branches, trunks and rhizomes of mature bamboos declined by 1.5, 23, 28 and 5 fold, respectively. The trunk contributed the most NSCs to the shoots. Our findings provide new insight and a possible rational mechanism explaining the “explosive growth” of Moso bamboo and shed new light on understanding the role of NSCs in the rapid growth of Moso bamboo.

Understanding the roles of nonstructural carbohydrates in forest trees - from what we can measure to what we want to know.

Contents 386 I. 386 II. 388 III. 392 IV. 392 V. 396 VI. 399 399 References 399 SUMMARY: Carbohydrates provide the building blocks for plant structures as well as versatile resources for metabolic processes. The nonstructural carbohydrates (NSC), mainly sugars and starch, fulfil distinct functional roles, including transport, energy metabolism and osmoregulation, and provide substrates for the synthesis of defence compounds or exchange with symbionts involved in nutrient acquisition or defence. At the whole-plant level, NSC storage buffers the asynchrony of supply and demand on diel, seasonal or decadal temporal scales and across plant organs. Despite its central role in plant function and in stand-level carbon cycling, our understanding of storage dynamics, its controls and response to environmental stresses is very limited, even after a century of research. This reflects the fact that often storage is defined by what we can measure, that is, NSC concentrations, and the interpretation of these as a proxy for a single function, storage, rather than the outcome of a range of NSC source and sink functions. New isotopic tools allow direct quantification of timescales involved in NSC dynamics, and show that NSC-C fixed years to decades previously is used to support tree functions. Here we review recent advances, with emphasis on the context of the interactions between NSC, drought and tree mortality.

Non-structural carbohydrate dynamics in Robinia pseudoacacia saplings under three levels of continuous drought stress

Only severe drought stress induced lower non-structural carbohydrate concentrations in Robinia pseudoacacia saplings, and non-structural carbohydrate reduction occurs solely in branches and stems at the end of the growing season. A better understanding of non-structural carbohydrate (NSC) dynamics in trees under drought stress is critical to elucidate the mechanisms underlying forest decline and tree mortality that result from extended periods of drought. We performed a continuous controlled drought pot experiment from May 14 to September 7, 2013 using Robinia pseudoacacia saplings under 14, 10, 8, and 6 % soil water content that represented the absence of drought (AD), moderate drought (MD), severe drought (SD), and very severe drought (VSD) stress, respectively. Growth, leaf gas exchange, and NSC concentrations in different sapling tissues were measured once a month. The results showed that net photosynthetic rates and the relative increments of basal diameter and height decreased with the intensification of drought stress. Saplings showed lower starch and NSC concentrations and higher soluble sugar concentrations and ratios of sugars to starch only under SD and VSD. The concentrations of starch and NSC were significantly decreased under SD and VSD solely in branches and stems at the end of the growing season. Our results indicated that the timescale of physiological processes plays an important role in NSC dynamics, and that only severe drought stress significantly decreases NSC concentrations, and NSC reduction occurs solely in branches and stems at the end of the growing season.

Desde el cámbium hasta el bosque: evaluando las respuestas de Juniperus thurifera L. a los condicionantes ambientales y antrópicos a múltiples escalas

Olano, J.M., Rozas, V. 2015. From cambium to woodland: analyzing Juniperus thurifera L. response to environmental constraints at multiple scales. Ecosistemas 24(2): 60-68. Doi.: 10.7818/ECOS.2015.24-2.09. Juniperus thurifera L. (Cupressaceae) is a dioecious tree inhabiting the Western Mediterranean Basin in areas under Mediterranean continental climate. Spain (95.1%) and Morocco (4.8%) host the largest populations. Diverse dendroecological research developed during the last years has improved our knowledge about this species functioning and its forest dynamics. This review approaches the advances produced on several topics including xylogenesis, climate forcing of secondary growth and xylem anatomy, regional variability on climate control, water-use efficiency, non-structural carbohydrates dynamics and present and future dynamics of its Iberian forests. A special point is paid on sex-related differences between male and female junipers, including non-structural carbohydrate accumulation patterns, secondary growth and response to summer drought, the main climatic factor limiting growth. This review aims to synthesize the available knowledge about this species at multiple spatio-temporal scales, being particularly useful for researchers and managers interested in the investigation, management and conservation of Iberian Juniperus thurifera woodlands.

Nonstructural carbohydrate dynamics of lodgepole pine dying from mountain pine beetle attack.

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.

How do drought and warming influence survival and wood traits of Picea mariana saplings?

Warming and drought will occur with increased frequency and intensity at high latitudes in the future. How heat and water stress can influence tree mortality is incompletely understood. The aim of this study was to evaluate how carbon resources, stem hydraulics, and wood anatomy and density determine the ability of black spruce saplings to survive daytime or night-time warming (+ 6 °C in comparison with control) in combination with a drought period. Plant water relations, the dynamics of non-structural carbohydrates and starch, mortality rate, and wood anatomy and density of saplings were monitored. Warming, in conjunction with 25 d of water deficit, increased sapling mortality (10% and 20% in night-time and daytime warming, respectively) compared with the control conditions (0.8%). Drought substantially decreased gas exchange, and also pre-dawn and mid-day leaf water potential to values close to -3MPa which probably induced xylem embolism (xylem air entry point, P₁₂, being on average around -3MPa for this species). In addition, the recovery of gas exchange never reached the initial pre-stress levels, suggesting a possible loss of xylem hydraulic conductivity associated with cavitation. Consequently, mortality may be due to xylem hydraulic failure. Warmer temperatures limited the replenishment of starch reserves after their seasonal minimum. Lighter wood was formed during the drought period, reflecting a lower carbon allocation to cell wall formation, preventing the adaptation of the hydraulic system to drought. Saplings of black spruce experienced difficulty in adapting under climate change conditions, which might compromise their survival in the future.

Carbohydrate dynamics and mortality in a piñon-juniper woodland under three future precipitation scenarios.

Drought-induced forest mortality is an increasing global problem with wide-ranging consequences, yet mortality mechanisms remain poorly understood. Depletion of non-structural carbohydrate (NSC) stores has been implicated as an important mechanism in drought-induced mortality, but experimental field tests are rare. We used an ecosystem-scale precipitation manipulation experiment to evaluate leaf and twig NSC dynamics of two co-occurring conifers that differ in patterns of stomatal regulation of water loss and recent mortality: the relatively desiccation-avoiding piñon pine (Pinus edulis) and the relatively desiccation-tolerant one-seed juniper (Juniperus monosperma). Piñon pine experienced 72% mortality after 13-25 months of experimental drought and juniper experienced 20% mortality after 32-47 months. Juniper maintained three times more NSC in the foliage than twigs, and converted NSC to glucose and fructose under drought, consistent with osmoregulation requirements to maintain higher stomatal conductance during drought than piñon. Despite these species differences, experimental drought caused decreased leaf starch content in dying trees of both species (P < 0.001). Average dry-season leaf starch content was also a good predictor of drought-survival time for both species (R(2) = 0.93). These results, along with observations of drought-induced reductions to photosynthesis and growth, support carbon limitation as an important process during mortality of these two conifer species.

Drought stress, growth and nonstructural carbohydrate dynamics of pine trees in a semi-arid forest.

In trees exposed to prolonged drought, both carbon uptake (C source) and growth (C sink) typically decrease. This correlation raises two important questions: (i) to what degree is tree growth limited by C availability; and (ii) is growth limited by concurrent C storage (e.g., as nonstructural carbohydrates, NSC)? To test the relationships between drought, growth and C reserves, we monitored the changes in NSC levels and constructed stem growth chronologies of mature Pinus halepensis Miller trees of three drought stress levels growing in Yatir forest, Israel, at the dry distribution limit of forests. Moderately stressed and stressed trees showed 34 and 14% of the stem growth, 71 and 31% of the sap flux density, and 79 and 66% of the final needle length of healthy trees in 2012. In spite of these large reductions in growth and sap flow, both starch and soluble sugar concentrations in the branches of these trees were similar in all trees throughout the dry season (2-4% dry mass). At the same time, the root starch concentrations of moderately stressed and stressed trees were 47 and 58% of those of healthy trees, but never <2% dry mass. Our results show that all the studied trees maintain a fairly good coordination between C supply and demand, and even during prolonged drought there is more than one way for a tree to maintain a positive C balance.