Non-structural Carbohydrate Reserves Research Articles

Canopy status modulates formation of wood rays in scots pine under hemiboreal conditions

Non-structural carbohydrates (NSC) reserves are crucial for trees to cope with weather extremes, thus to ensure their survival and ecological plasticity. The NSC reserves can depend on social status, suggesting uneven plasticity of trees at the stand level. In stemwood of Scots pine (Pinus sylvestris L.), which is a widespread and important species, NSC reserves are stored in parenchyma in wood rays (WR). The quantity of WRs is adjusted intra-annually, allowing retrospective analysis of factors affecting their formation. Accordingly, the differences in WR quantity in stemwood of dominant and intermediate (canopy trees with reduced and narrow crowns) maturing Scots pine were assessed by quantitative wood anatomy. Tangential cuts from the outermost 30 tree-rings were analysed. The relative ray area was intermediate, i.e. covering about 5% of the tangential cut, yet expressed high individuality among the trees. The size and amount of WR mainly differed between the earlywood and latewood; WRs in latewood were higher although narrower in comparison to earlywood, yet their total amount was higher in earlywood. Canopy status had only a slight effect, as quantity and height of WR tended to be higher for the intermediate trees, particularly in earlywood. The size and quantity of WR expressed inter-annual variation, which was mainly related to the meteorological conditions prior to the formation of the tree-ring (previous summer and autumn) indicating legacy effects of climatic factors on NSC and susceptibility of trees to cumulative effects of weather extremes. However, the climatic signals in the inter-annual variation of WR were weaker than observed before, likely due to location of the studied stand in the mid-part of the species range. Nevertheless, the observed differences in mean values and inter-annual variation of WR suggested a within-species diversity of carbon allocation patterns, supporting adaptability of the species.

Declining carbohydrate content of Sitka-spruce treesdying from seawater exposure.

Increasing sea levels associated with climate change threaten the survival of coastal forests, yet the mechanisms by which seawater exposure causes tree death remain poorly understood. Despite the potentially crucial role of nonstructural carbohydrate (NSC) reserves in tree survival, their dynamics in the process of death under seawater exposure are unknown. Here we monitored progressive tree mortality and associated NSC storage in Sitka-spruce (Picea sitchensis) trees dying under ecosystem-scale increases in seawater exposure in western Washington, USA. All trees exposed to seawater, because of monthly tidal intrusion, experienced declining crown foliage during the sampling period, and individuals with a lower percentage of live foliated crown (PLFC) died faster. Tree PLFC was strongly correlated with subsurface salinity and needle ion contents. Total NSC concentrations in trees declined remarkably with crown decline, and reached extremely low levels at tree death (2.4% and 1.6% in leaves and branches, respectively, and 0.4% in stems and roots). Starch in all tissues was almost completely consumed, while sugars remained at a homeostatic level in foliage. The decreasing NSC with closer proximity to death and near zero starch at death are evidences that carbon starvation occurred during Sitka-spruce mortality during seawater exposure. Our results highlight the importance of carbon storage as an indicator of tree mortality risks under seawater exposure.

Hydraulic dysfunction due to root-exposure-initiated water stress is responsible for the mortality of Salix gordejevii shrubs on the windward slopes of active sand dunes

Sand-fixing shrub Salix gordejevii plays an important role in sand dune stabilization but often shows severe decline and mortality on the windward slopes of active dunes. The objective of this study was to explore the environmental drivers and physiological mechanisms behind its decline and mortality on the windward slopes of active sand dunes. The xylem hydraulics, sap flow, water relations and non-structural carbohydrate reserves were measured for S. gordejevii plants that respectively inhabited interdune, leeward and windward positions of active dunes with contrasting soil water regimes. Root exposure caused by wind erosion, rather than low soil moisture, predisposed windward S. gordejevii plants to stronger water stress. Correspondingly, the windward plants showed reduced transpirational water use and exhibited a suite of acclimative adjustments in leaf traits favoring water conservation. Despite these adjustments, windward S. gordejevii plants still suffered from higher risks of hydraulic failure and branch mortality. No evidence of carbon depletion in windward S. gordejevii plants was found although the soluble sugar to starch ratios changed significantly. Our findings suggest that wind-erosion-induced root exposure triggers water stress and eventually causes the mortality of windward S. gordejevii plants, during which increased risk of hydraulic dysfunction is pronounced.

Multiple stressors and disturbance effects on eelgrass and epifaunal macroinvertebrate assemblage structure

Multiple forms of environmental change and anthropogenic pressure co-occur in coastal marine ecosystems. These external forces affect ecosystem structure, functioning, and, eventually, services to humans. Studies that include more than 2 simultaneous stressors are still needed to understand potential interactions among multiple stressors. We evaluated single and interactive effects of density reduction of Zostera marina L. (a habitat-forming species), shading, and sediment nutrient enrichment on the response of Z. marina and its associated epifauna over 10 wk. Shading had the greatest effect on reducing the eelgrass relative leaf elongation rate (RLE), non-structural carbohydrate reserves, and eelgrass shoot density. A reduced eelgrass density sustained higher epifaunal densities and increased the eelgrass RLE. Sediment nutrient enrichment increased eelgrass shoot density but decreased epifaunal richness, diversity, and total abundance. Our disturbance and pair of stressors differed in their influence on diversity measures, but all affected assemblage structure. Most of the changes to the epifaunal assemblage and diversity likely occurred due to altered habitat availability and epiphytic algae load. We observed additive, antagonistic, and negatively synergistic interactions among our treatments, while most of the cumulative effects showed dominance by one stressor over another. Our results highlight the importance of field experiments that are based on multiple disturbances and stressors to determine their interaction type on communities.

Water relations and non-structural carbohydrate responses to the combined effects of defoliation and progressive drought in a dioecious tree

Increases in drought frequency and insect defoliation are emerging threats that typically affect the plantation production of Populus yunnanensis, which is an endemic dioecious tree species in China. Nevertheless, it is difficult to predict which sex of P. yunnanensis will be more affected by the combined effects of water limitations and defoliation. In this study, we examined the combined effects of progressive drought and defoliation on water relations, growth, gas exchange and leaf carbohydrate reserves in P. yunnanensis females and males to test whether defoliation alters the effects of drought and and how this response varies between genders. The experimental design included two watering regimes (well-watered and drought-stressed) and two levels of defoliation (leaves untouched and 60% defoliated). Defoliation exacerbated drought stress in females by decreasing photosynthesis, dry mass accumulation and leaf non-structural carbohydrate (NSC) concentration. In contrast, the total biomass of males grown under drought combined with defoliation was not significantly altered when compared to those grown under drought alone. These results suggest that defoliation does not aggravate the drought effects on males. The males presented a high capacity to maintain foliar water status through stomatal control under drought conditions irrespective of defoliation treatment. The conservative water use strategy in males resulted in a gradual reduction in leaf water potential, gas exchange rates and NSC. Our data showed that the different responses of the females and males to drought and defoliation were influenced by plant water status and consequent effects on changes in carbon assimilation and leaf non-structural carbohydrate reserves.

Xeric Tree Populations Exhibit Delayed Summer Depletion of Root Starch Relative to Mesic Counterparts

Research linking soil moisture availability to nonstructural carbohydrate (NSC) storage suggests greater NSC reserves promote survival under acute water stress, but little is known about how NSC allocation responds to long-term differences in water availabilty. We hypothesized populations experiencing chronic or frequent water stress shift carbon allocation to build greater NSC reserves for increased survival probability during drought relative to populations rarely experiencing water stress. Over a year, we measured soluble sugar and starch concentrations from branches, stems, and coarse roots of mature Pinus palustris trees at two sites differing in long-term soil moisture availability. Xeric and mesic populations exhibited a cycle of summer depletion-winter accumulation in root starch. Xeric populations reached a maximum root starch concentration approximately 1–2 months later than mesic populations, indicating delayed summer depletion. Xeric and mesic populations reached the same minimum root starch at similar times, suggesting extended winter accumulation for xeric populations. These results suggest seasonal mobilization from root starch is compressed into a shorter interval for xeric populations instead of consistently greater reserves as hypothesized. Seasonal trends differed little between xeric and mesic populations for starch and sugars, suggesting the importance of roots in seasonal carbon dynamics and the primacy of starch for storage. If roots are the primary organ for longterm storage, then our results suggest that whole-plant mobilization and allocation respond to chronic differences in water availability.

Insect defoliation is linked to a decrease in soil ectomycorrhizal biomass and shifts in needle endophytic communities.

Insect outbreaks of increasing frequency and severity in forests are predicted due to climate change. Insect herbivory is known to promote physiological changes in forest trees. However, little is known about whether these plant phenotypic adjustments have cascading effects on tree microbial symbionts such as fungi in roots and foliage. We studied the impact of defoliation by the pine processionary moth in two infested Pinus nigra forests through a multilevel sampling of defoliated and non-defoliated trees. We measured tree growth, nutritional status and carbon allocation to chemical defenses. Simultaneously, we analysed the putative impact of defoliation on the needle endophytes and on the soil fungal communities. Higher concentrations of chemical defenses were found in defoliated trees, likely as a response to defoliation; however, no differences in non-structural carbohydrate reserves were found. In parallel to the reductions in tree growth and changes in chemical defenses, we observed shifts in the composition of needle endophytic and soil fungal communities in defoliated trees. Defoliated trees consistently corresponded with a lower biomass of ectomycorrhizal fungi in both sites, and a higher alpha diversity and greater relative abundance of belowground saprotrophs and pathogens. However, ectomycorrhizal alpha diversity was similar between non-defoliated and defoliated trees. Specific needle endophytes in old needles were strongly associated with non-defoliated trees. The potential role of these endophytic fungi in pine resistance should be further investigated. Our study suggests that lower biomass of ectomycorrhizal fungi in defoliated trees might slow down tree recovery since fungal shifts might affect tree-mycorrhizal feedbacks and can potentially influence carbon and nitrogen cycling in forest soils.

Hydraulic and photosynthetic limitations prevail over root non-structural carbohydrate reserves as drivers of resprouting in two Mediterranean oaks.

Resprouting is an ancestral trait in angiosperms that confers resilience after perturbations. As climate change increases stress, resprouting vigor is declining in many forest regions, but the underlying mechanism is poorly understood. Resprouting in woody plants is thought to be primarily limited by the availability of non-structural carbohydrate reserves (NSC), but hydraulic limitations could also be important. We conducted a multifactorial experiment with two levels of light (ambient, 2-3% of ambient) and three levels of water stress (0, 50 and 80 percent losses of hydraulic conductivity, PLC) on two Mediterranean oaks (Quercus ilex and Q. faginea) under a rain-out shelter (n = 360). The proportion of resprouting individuals after canopy clipping declined markedly as PLC increased for both species. NSC concentrations affected the response of Q. ilex, the species with higher leaf construction costs, and its effect depended on the PLC. The growth of resprouting individuals was largely dependent on photosynthetic rates for both species, while stored NSC availability and hydraulic limitations played minor and non-significant roles, respectively. Contrary to conventional wisdom, our results indicate that resprouting in oaks may be primarily driven by complex interactions between hydraulics and carbon sources, whereas stored NSC play a significant but secondary role.

Non-structural carbohydrate concentrations of Fagus sylvatica and Pinus sylvestris fine roots are linked to ectomycorrhizal enzymatic activity during spring reactivation

We evaluated whether changes in fine root non-structural carbohydrate reserves of Fagus sylvatica and Pinus sylvestris trees influence potential enzymatic activities of their ectomycorrhizal symbionts from winter towards spring reactivation, and whether these changes influence potential soil enzymatic activities. We analyzed sugar and starch concentrations in the fine roots of Fagus sylvatica and Pinus sylvestris and potential activities of ß-glucosidase, ß-xylosidase, and cellobiohydrolase (as proxies for carbon-degrading enzymes) as well as leucine aminopeptidase and chitinase (as proxies for nitrogen-degrading enzymes) of their dominant ectomycorrhizal symbionts as well as in the soil. Sugar concentrations in the fine roots were significantly positively correlated with enzymatic activities of the ectomycorrhizal symbionts. In Pinus sylvestris, both carbon- and nitrogen-degrading enzyme activities showed significant positive correlations with fine root sugar concentrations. In Fagus sylvatica, fine root sugar concentrations were explicitly positively correlated with the activity of nitrogen-degrading enzymes. The chitinase activity in the soil was found to be strongly positively correlated with the enzymatic activity of the ectomycorrhizal symbionts as well as with fine root sugar concentrations. Fine root carbohydrate concentrations of Fagus sylvatica and Pinus sylvestris trees and enzymatic activities of their associated ectomycorrhizal fungi are connected. The specific nutrient demand of the tree species during spring reactivation may affect ectomycorrhizal enzymatic activity via carbon mobilization in the fine roots of Fagus sylvatica and Pinus sylvestris. Moreover, our results suggest that trees indirectly contribute to the degradation of fungal necromass by stimulating ectomycorrhizal chitinase activity in the soil.

Alteration in plant spacing improves submergence tolerance in Sub1 and non-Sub1 rice (cv. IR64) by better light interception and effective carbohydrate utilisation under stress.

Besides genetic improvement for developing stress-tolerant cultivars, agronomic management may also add considerable tolerance against different abiotic stresses in crop plants. In the present study, we evaluated the effect of six different spacing treatments (S1: 10 × 10 cm; S2: 15 × 10 cm; S3:15 × 15 cm; S4:20 × 10 cm; S5: 20 × 15 cm; S6: 20 × 20 cm (row-row × plant-plant)) for improving submergence tolerance in rice. A high yielding submergence intolerant rice cultivar IR64 was tested against its SUB1 QTL introgressed counterpart (IR64-Sub1) for 12 days of complete submergence for different spacing treatments in field tanks. Relatively wider spaced plants showed higher individual plant biomass and early seedling vigour, which was particularly helpful for IR64 in increasing plant survival (by 150% in S6 over S1) under 12 days of submergence, whereas the improvement was less in IR64-Sub1 (13%). Underwater radiation inside the plant canopy, particularly beyond 40 cm water depth, was significantly greater in wider spacing treatments. Leaf senescence pattern captured by SPAD chlorophyll meter reading and chlorophyll fluorescence imaging data (Fm, Fv/Fm) taken at different time intervals after stress imposition suggested that there was lesser light penetration inside the canopy of closer spaced plants, and that it might hasten leaf senescence and damage to the photosynthetic system. The initial content of total non-structural carbohydrate (NSC) was higher in wider spaced plants of IR64, and also the rate of depletion of NSC was lesser compared with closer spaced plants. In contrast, there was not much difference in NSC depletion rate under different spacing in IR64-Sub1. Further, higher antioxidant enzyme activities in wider spaced plants (both IR64 and IR64-Sub1) after de-submergence indicated better stress recovery and improved tolerance. Taken together we found that wider spacing (row-row: 20 cm and plant-plant: 15 cm and more) can significantly improve submergence tolerance ability in rice, particularly in submergence intolerant non-Sub1 cultivar like IR64, perhaps due to better underwater light penetration, delayed leaf senescence and slower depletion of NSC reserve.

Managing Grazing Intensity Linked to Forage Quantity and Quality Trade-off in Semiarid Rangelands

Knowledge of herbivory tolerance of dominant plant species and their responses to varying grazing intensity is required for sustainable grazing management in semiarid rangeland ecosystems. In a field experiment, we studied the effects of simulated grazing 0%, 30%, 50%, and 80% leaf removal (control, partial, intermediate, and heavy leaf removal) of three dominant perennial legumes: Astragalus cyclophyllon, Astragalus effuses, and Hedysarum criniferum in semiarid rangelands of southwestern Iran for 2 yr and tested 1) differential herbivory tolerance in terms of regrowth ability of belowground and aboveground biomass (AGB), fecundity, and nonstructural carbohydrate reserves; and 2) trade-off between forage quantity and quality—less defoliated plants produce larger amount of biomass with less forage quality and vice versa. The results showed a reduction in AGB and reproduction in each of the three plants. However, three forbs exhibited some sort of variations in shoot and root growth, reproduction and forage quality under intermediate and heavy leaf removal. Leaf removal in individuals of A. effusus and H. criniferum are more likely to compensate for tissue loss by allocating the available resources to the expansion of root biomass, at the cost of AGB and reproductive effort. Leaf removal also led to an increase in forage quality in each of the three forb species, depending on the level of height removal. The control plant had a higher amount of biomass production and lower amount of crude protein compared with heavy leaf removal treatments (i.e., 80%), by contrasting responses of acid detergent fiber and neutral detergent fiber, indicating a trade-off between forage quality and forage quantity. The results indicate that there is a balance among forage quantity, quality, and regrowth ability of belowground and aboveground tissues in intermediate leaf removal treatments of these forb species. This balance is used to develop principles for grazing management of steppe rangelands because such a regime meets different criteria including forage quantity, quality, and nonstructural carbohydrate reserves.

Life at 0 °C: the biology of the alpine snowbed plant Soldanella pusilla

All plant species reach a low temperature range limit when either low temperature extremes exceed their freezing tolerance or when their metabolism becomes too restricted. In this study, we explore the ultimate thermal limit of plant tissue formation exemplified by a plant species that seemingly grows through snow. By a combination of studies in alpine snowbeds and under controlled environmental conditions, we demonstrate and quantify that the clonal herb Soldanella pusilla (Primulaceae) does indeed grow its entire flowering shoot at 0 °C. We show that plants resume growth under 2–3 m of snow in mid-winter, following an internal clock, with the remaining period under snow until snow melt (mostly in July) sufficient to produce a flowering shoot that is ready for pollination. When snow pack gets thin, the flowering shoot intercepts and re-radiates long-wave solar radiation, so that snow and ice gently melt around the fragile shoot and the flowers emerge without any mechanical interaction. We evidence bud preformation in the previous season and enormous non-structural carbohydrate reserves in tissues (mainly below ground) in the form of soluble sugars (largely stachyose) that would support basic metabolism for more than 2 entire years under snow. However, cell-wall formation at 0 °C appears to lack unknown strengthening factors, including lignification (assessed by confocal Raman spectroscopy imaging) that require between a few hours or a day of warmth after snow melt to complete tissue strengthening. Complemented with a suite of anatomical data, the work opens a window towards understanding low temperature limits of plant growth in general, with potential relevance for winter crops and trees at the natural climatic treeline.

Climate and disturbance influence self-sustaining stand dynamics of aspen (Populus tremuloides) near its range margin.

Species that are primarily seral may form stable (self-sustaining) communities under certain disturbance regimes or environmental conditions, yet such populations may also be particularly vulnerable to ecological change. Aspen (Populus spp.) are generally considered seral throughout the Northern Hemisphere, including P.tremuloides, the most widely distributed tree species in North America. Recent declines in aspen populations have occurred, especially along drought-sensitive margins of its range and where fire exclusion and herbivory have promoted community transition. However, aspen also forms stable stands, and examination of the mechanisms that influence persistence can offer conservation insights, especially where populations are vulnerable to changing climate or altered disturbance dynamics. We sampled tree age and stand characteristics of isolated aspen forests in the arid Great Basin (USA) to determine if (1) aspen communities are more fire-dependent and seral or fire-independent and stable; (2) ungulate browsing inhibits aspen stability; and (3) temporal patterns of vegetative reproduction (i.e., ramet establishment or "suckering") are correlated with climate. Aspen size and age class densities strongly fit negative exponential distributions, whether grouped geographically or by functional type, suggesting landscape-scale persistence. Continuous age distributions and high proportions of recruitment-sized to overstory trees suggest stability at stand scales, with exceptions including stands with higher browsing pressure. Few stands had evidence of fire, and relationships between dead tree size and variability in live tree size suggest a lack of fire dependency. Several 5-yr averaged climate variables and one sea surface temperature index were correlated with aspen ramet establishment densities over time, with strongest relationships occurring ~5yr prior to establishment year, often followed by inverse relationships ~1yr after. Indeed, aspen establishment density for a recent 41-yr period was reliably reconstructed using antecedent climate conditions derived from a single drought index. Temporally synchronized aspen ramet establishment across the study region may be due to climate-driven storage of nonstructural carbohydrate reserves in clonal root systems later used for regeneration. Complex regeneration dynamics of these self-sustaining aspen stands, especially sensitivity to climate variability, suggest they may serve as harbingers of ecological change in the arid Great Basin and in other aspen populations near their range margin.

Non-structural Carbohydrates in Dormant Woody Perennials; The Tale of Winter Survival and Spring Arrival

Woody perennials’ reliance on nonstructural carbohydrates (NSC) reserves for the resumption of spring growth necessitates an accumulation of NSC prior to dormancy. It is assumed that during dormancy temperature-regulated biological activities gauge the progression of winter and affect the metabolic rates and physiology of NSC reserves. Thus, changes in temperature signal the arrival of spring and determine the amount of reserves available for growth resumption. As woody perennials are dependent on dispersed storage of NSC during spring, they need an integrated remobilization and redistribution for synchronous and effective development of photosynthetic and reproductive organs. However, it is not known how storage compartments interact at the whole plant level, when NSC reserves are mobilized, or how local and distal storage compartments influence the biology of spring growth resumption. The goal of this mini-review is to shift the focus of winter biology from bud-centric to the whole plant. We discuss winter NSC management in the context of climate change with a special emphasis on how projected mild winters may affect the carbon budget, transport, and allocation during winter. We look at three aspects of NSC regulation underlying dormancy (I) the molecular regulation of dormancy (II) temperature dependent winter NSC metabolism, and (III) spring NSC remobilization and redistribution processes.

Characteristics of non-structural carbohydrate reserves of three dominant tree species in broadleaved Korean pine forest in Changbai Mountain, China.

Non-structural carbohydrates (NSC, including soluble sugars and starch) are key meta-bolites in tree, the storage characteristics of which in tree organs have received extensive attention. It is still unclear how NSC are allocated in the tissues (phloem and xylem) that have different function. In this study, we analyzed the concentration and allocation of NSC in the roots, and in phloem and xylem of the trunk in three dominant species of broadleaved Korean pine forest in the Changbai Mountain, Pinus koraiensis, Fraxinus mandschurica, and Tilia amurensis. The results showed that there was a significant difference in the concentration of NSC between the phloem and xylem. The soluble sugar dominated in the phloem, while starch dominated in the xylem. The concentration of NSC in trunk outside (divided by annual rings, 0-20 years), intermediate (20-40 years) and inner (>40 years) of different tree species was significantly different, but with no difference in the roots. The total soluble sugar concentration in the phloem of P. koraiensis and F. mandschurica was significantly higher than that of T. amurensis, while the difference in xylem was not significant. The results indicated that NSC allocation in the phloem and xylem of the tree had clear tissue differentiation, which might be related to the succession stage of the tree species or the functional evolution of the tissue. These findings would improve our understanding of the carbon storage characteristics and allocation mechanism in temperate trees.

The interaction between nonstructural carbohydrate reserves and xylem hydraulics in Korean pine trees across an altitudinal gradient.

Nonstructural carbohydrates (NSC) have been proposed to play an important role in maintaining the hydraulic integrity of trees, particularly in environments with high risks of embolism formation, but knowledge about the interaction between NSC reserves and xylem hydraulics is still very limited. We studied the variation of NSC reserves and hydraulic traits in Pinus koraiensis Sieb. et Zucc. (Korean pine) in March and June across a relatively large altitudinal gradient in Changbai Mountain of Northeast China. One of the major aims was to investigate the potential role NSC plays in maintaining hydraulic integrity of overwintering stems in facing freezing-induced embolism. Consistent with our hypotheses, substantial variations in both NSC contents and hydraulic traits were observed across altitudes and between the two seasons. In March, when relatively high degrees of winter embolism exist, the percentage loss of conductivity (PLC) showed an exponential increase with altitude. Most notably, positive correlations between branch and trunk soluble sugar content and PLC (P = 0.053 and 0.006) were observed across altitudes during this period. These correlations could indicate that more soluble sugars are required for maintaining stem hydraulic integrity over the winter by resisting or refilling freezing-induced embolism in harsher environments, although more work is needed to establish a direct causal relationship between NSC dynamics and xylem hydraulics. If the correlation is indeed directly associated with varying demands for maintaining hydraulic integrity across environmental gradients, greater carbon demands may compromise tree growth under conditions of higher risk of winter embolism leading to a trade-off between competitiveness and stress resistance, which may be at least partially responsible for the lower dominance of Korean pine trees at higher altitudes.

Whole-tree nonstructural carbohydrate storage and seasonal dynamics in five temperate species.

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.

Trees use more non-structural carbohydrate reserves during epicormic than basal resprouting.

Non-structural carbohydrates (NSCs) are crucial to support tree resprouting after disturbances that damage the crown or stem. Epicormic resprouting (from stem) could demand more from NSC reserves than basal resprouting (following top-kill), since epicormically resprouting trees need to maintain a greater living biomass. Yet, little is known about NSC use during epicormic resprouting, particularly the relative importance of stem and below-ground NSC reserves. We compared the distribution and magnitude of NSC decreases during epicormic and basal resprouting by experimentally removing crowns or stems of 14-year-old Eucalyptus obliqua L'Hér. trees in native forest, then harvesting these trees over a 10-month period (start, sprouts emerged, sprouts expanded) to measure changes in NSC concentration and mass by organ (stem, lignotuber, roots). We hypothesized that (i) NSC depletion during resprouting is primarily due to decreases in starch rather than soluble sugars concentrations; (ii) during epicormic resprouting, stem NSC concentrations are decreased irrespective of any decreases in roots; and (iii) absolute decreases in NSC mass are greater for epicormic than basal resprouting during the leafless period due to the carbon demands associated with maintaining greater living biomass. Results confirmed our hypotheses; starch was the primary storage carbohydrate, stems were an important source of starch during epicormic resprouting and carbon demands of maintenance functions were greater for epicormic resprouting, leading to greater decreases in NSC reserves. Roots were a more important starch storage organ than the lignotuber for both epicormic and basal resprouting. The proportional decrease in starch was severe for both modes of resprouting due to a long leafless period, after which trees resprouting epicormically relied on starch reserves for longer than those resprouting basally. It remains to be seen how the timing of disturbance affects the timing and vigour of resprouting, and how long-term NSC recovery differs for epicormic and basal resprouting.

Seasonal patterns of nonstructural carbohydrate reserves in four woody boreal species1

Plants store nonstructural carbohydrates (NSCs), such as sugars and starch, to use as carbon and energy sources for daily maintenance and growth needs as well as during times of stress. Allocation of NSCs to storage provides an important physiological strategy associated with future growth and survival, and thus understanding the seasonal patterns of NSC reserves provides insight into how species with different traits (e.g., growth form, leaf habit, wood anatomy) may respond to stress. We characterized the seasonal patterns of NSCs in four woody boreal plant species in Minnesota, USA. Sugar and starch concentrations were measured across the year in the roots and branches of two conifer trees, black spruce (Picea mariana (Mill.) B.S.P.) and eastern tamarack (Larix laricina (Du Roi) K. Koch), as well as in the leaves and branches of two evergreen broadleaf shrubs, bog Labrador tea (Rhododendron groenlandicum (Oeder) Kron & Judd) and leatherleaf (Chamaedaphne calyculata (L.) Moench). In general, seasonal variation was dominated by changes in starch across all organs and species. While similar seasonal patterns of NSCs were observed in the shrubs, different seasonal patterns were observed between the trees, particularly in the roots. Our results suggest that species-specific traits likely have consequences for organ-level storage dynamics, which may influence whole-plant growth and survival under global change.

Growth traits of juvenile American chestnut and red oak as adaptations to disturbance

American chestnut (Castanea dentata) was a dominant species in eastern North America prior to the importation of chestnut blight. In light of recent efforts to restore viable populations of chestnut in eastern forests, an increased understanding of its association with other co‐occurring, disturbance‐adapted oak species is necessary. We evaluated crown architecture and leaf morphology in juvenile chestnut and red oak (Quercus rubra) to assess potential differences in establishment strategies of both species. We also investigated differences in nonstructural carbohydrate reserves and whole tree biomass partitioning between species. Seedlings of both species were planted in forest stands treated either with midstory removal or small patch cuts, simulating potential restoration plantings. After 5–7 years, chestnut's allocation to its root system was lower than red oak's, with chestnut saplings instead diverting resources to branches and foliage. Chestnut had lower leaf area index, greater crown projection area, and higher specific leaf area than red oak, indicating the species may have an advantage in shaded understories. There were only minor differences in nonstructural root carbohydrate reserves, between red oak and American chestnut, indicating that chestnut may respond similarly to oak by resprouting after disturbances topkill young saplings. We suggest that American chestnut has morphological and physiological attributes that allow it to function as an opportunistic and plastic species that can utilize gaps to facilitate its canopy recruitment, yet still persist after occasional surface fire. This knowledge can guide restoration strategies for this iconic species of the eastern temperate forest region.