Non-structural Carbohydrate Storage Research Articles

The role of stored carbohydrates and nitrogen in the growth and stress tolerance of planted forest trees

Plants store compounds that supplement external resources to maintain primary functions. We reviewed the role of stored non-structural carbohydrates (NSC) and nitrogen (N) in juvenile woody species for spring growth and cold and drought stress tolerance, which are crucial processes for early performance of forest plantations. Plant functional types differed in NSC and N partitioning and allocation to new growth. In general, however, new leaves/shoots were more enriched in remobilized resources than new fine roots. Conifers used less remobilized resources than broadleaf species for fine root growth. New shoots/leaves were mostly comprised of remobilized N (>60 %) in conifers and broadleaf deciduous species, while broadleaf evergreens relied more on soil N (<50 % remobilized N). In contrast, few differences among functional groups existed in the contribution of remobilized carbon (C) to new leaves/shoots, which comprised 28–45 % of stored C reflecting the importance of current photosynthesis and distinctions in C and N remobilization physiology. The amount of N remobilized by an organ was positively related to its contribution to seedling N content. However, leaves are priority N sources in evergreens, which remobilized more N than predicted by their contribution to seedling N content. In contrast, roots in broadleaf evergreens and conifers were poor contributors of remobilized N. Under low stress, spring growth has little effect on NSC reserves. However, prolonged and intense photosynthesis depression strongly reduces NSC. In contrast, N reserves usually decline after planting and their replenishment takes longer than for NSC reserves. Strong storage reduction can hinder seedling stress acclimation and survival capacity. Resource storage can be promoted in the nursery by arresting plant growth and supplying resources at a higher rate than seedling growth and maintenance rate. We conclude that the way in which woody plants manage stored resources drives their growth and stress tolerance. However, plant functional types differ in storage physiology, which should be considered in silvicultural management.

Allocation to carbon storage pools in Norway spruce saplings under drought and low CO2.

Non-structural carbohydrates (NSCs) are critical to maintain plant metabolism under stressful environmental conditions, but we do not fully understand how NSC allocation and utilization from storage varies with stress. While it has become established that storage allocation is unlikely to be a mere overflow process, very little empirical evidence has been produced to support this view, at least not for trees. Here we present the results of an intensively monitored experimental manipulation of whole-tree carbon (C) balance (young Picea abies (L.) H Karst.) using reduced atmospheric [CO2] and drought to reduce C sources. We measured specific C storage pools (glucose, fructose, sucrose, starch) over 21 weeks and converted concentration measurement into fluxes into and out of the storage pool. Continuous labeling ((13)C) allowed us to track C allocation to biomass and non-structural C pools. Net C fluxes into the storage pool occurred mainly when the C balance was positive. Storage pools increased during periods of positive C gain and were reduced under negative C gain. (13)C data showed that C was allocated to storage pools independent of the net flux and even under severe C limitation. Allocation to below-ground tissues was strongest in control trees followed by trees experiencing drought followed by those grown under low [CO2]. Our data suggest that NSC storage has, under the conditions of our experimental manipulation (e.g., strong progressive drought, no above-ground growth), a high allocation priority and cannot be considered an overflow process. While these results also suggest active storage allocation, definitive proof of active plant control of storage in woody plants requires studies involving molecular tools.

Measuring and modelling seasonal patterns of carbohydrate storage and mobilization in the trunks and root crowns of peach trees.

Developing a conceptual and functional framework for simulating annual long-term carbohydrate storage and mobilization in trees has been a weak point for virtually all tree models. This paper provides a novel approach for solving this problem using empirical field data and details of structural components of simulated trees to estimate the total carbohydrate stored over a dormant season and available for mobilization during spring budbreak. The seasonal patterns of mobilization and storage of non-structural carbohydrates in bark and wood of the scion and rootstock crowns of the trunks of peach (Prunus persica) trees were analysed subsequent to treatments designed to maximize differences in source-sink behaviour during the growing season. Mature peach trees received one of three treatments (defruited and no pruning, severe pruning to 1·0 m, and unthinned with no pruning) in late winter, just prior to budbreak. Selected trees of each treatment were harvested at four times (March, June, August and November) and slices of trunk and root crown tissue above and below the graft union were removed for carbohydrate analysis. Inner bark and xylem tissues from the first to fifth rings were separated and analysed for non-structural carbohydrates. Data from these experiments were then used to estimate the amount of non-structural carbohydrates available for mobilization and to parameterize a carbohydrate storage sub-model in the functional-structural L-PEACH model. The mass fraction of carbohydrates in all sample tissues decreased from March to June, but the decrease was greatest in the severely pruned and unthinned treatments. November carbohydrate mass fractions in all tissues recovered to values similar to those in the previous March, except in the older xylem rings of the severely pruned and unthinned treatment. Carbohydrate storage sink capacity in trunks was empirically estimated from the mean maximum measured trunk non-structural carbohydrate mass fractions. The carbohydrate storage source available for mobilization was estimated from these maximum mass fractions and the early summer minimum mass fractions remaining in these tissues in the severe treatments that maximized mobilization of stored carbohydrates. The L-PEACH sink-source carbohydrate distribution framework was then used along with simulated tree structure to successfully simulate annual carbohydrate storage sink and source behaviour over years. The sink-source concept of carbohydrate distribution within a tree was extended to include winter carbohydrate storage and spring mobilization by considering the storage sink and source as a function of the collective capacity of active xylem and phloem tissue of the tree, and its annual behaviour was effectively simulated using the L-PEACH functional-structural plant model.

Reserves Accumulated in Non-Photosynthetic Organs during the Previous Growing Season Drive Plant Defenses and Growth in Aspen in the Subsequent Growing Season

Plants store non-structural carbohydrates (NSC), nitrogen (N), as well as other macro and micronutrients, in their stems and roots; the role of these stored reserves in plant growth and defense under herbivory pressure is poorly understood, particularly in trees. Trembling aspen (Populus tremuloides) seedlings with different NSC and N reserves accumulated during the previous growing season were generated in the greenhouse. Based on NSC and N contents, seedlings were assigned to one of three reserve statuses: Low N-Low NSC, High N-Medium NSC, or High N-High NSC. In the subsequent growing season, half of the seedlings in each reserve status was subjected to defoliation by forest tent caterpillar (Malacosoma disstria) while the other half was left untreated. Following defoliation, the effect of reserves was measured on foliar chemistry (N, NSC) and caterpillar performance (larval development). Due to their importance in herbivore feeding, we also quantified concentrations of phenolic glycoside compounds in foliage. Seedlings in Low N-Low NSC reserve status contained higher amounts of induced phenolic glycosides, grew little, and supported fewer caterpillars. In contrast, aspen seedlings in High N-Medium or High NSC reserve statuses contained lower amounts of induced phenolic glycosides, grew faster, and some of the caterpillars which fed on these seedlings developed up to their fourth instar. Furthermore, multiple regression analysis indicated that foliar phenolic glycoside concentration was related to reserve chemistry (NSC, N). Overall, these results demonstrate that reserves accumulated during the previous growing season can influence tree defense and growth in the subsequent growing season. Additionally, our study concluded that the NSC/N ratio of reserves in the previous growing season represents a better measure of resources available for use in defense and growth than the foliar NSC/N ratios.

An integrated model of environmental effects on growth, carbohydrate balance, and mortality of Pinus ponderosa forests in the southern Rocky Mountains.

Climate-induced tree mortality is an increasing concern for forest managers around the world. We used a coupled hydrologic and ecosystem carbon cycling model to assess temperature and precipitation impacts on productivity and survival of ponderosa pine (Pinus ponderosa). Model predictions were evaluated using observations of productivity and survival for three ponderosa pine stands located across an 800 m elevation gradient in the southern Rocky Mountains, USA, during a 10-year period that ended in a severe drought and extensive tree mortality at the lowest elevation site. We demonstrate the utility of a relatively simple representation of declines in non-structural carbohydrate (NSC) as an approach for estimating patterns of ponderosa pine vulnerability to drought and the likelihood of survival along an elevation gradient. We assess the sensitivity of simulated net primary production, NSC storage dynamics, and mortality to site climate and soil characteristics as well as uncertainty in the allocation of carbon to the NSC pool. For a fairly wide set of assumptions, the model estimates captured elevational gradients and temporal patterns in growth and biomass. Model results that best predict mortality risk also yield productivity, leaf area, and biomass estimates that are qualitatively consistent with observations across the sites. Using this constrained set of parameters, we found that productivity and likelihood of survival were equally dependent on elevation-driven variation in temperature and precipitation. Our results demonstrate the potential for a coupled hydrology-ecosystem carbon cycling model that includes a simple model of NSC dynamics to predict drought-related mortality. Given that increases in temperature and in the frequency and severity of drought are predicted for a broad range of ponderosa pine and other western North America conifer forest habitats, the model potentially has broad utility for assessing ecosystem vulnerabilities.

Spatial variations in non-structural carbohydrates in stems of twelve temperate tree species

The radial, axial and inter-specific variations in concentrations and contents of non-structural carbohydrates (NSC) in stems were investigated for 12 Chinese temperate tree species. These species had contrasting leaf phenology (evergreen and deciduous) and wood types (non-, ring- and diffuse-porous wood). For each species, we sampled bark (periderm and phloem), outer wood (light-colored) and inner wood (dark-colored) at four heights along the stem (stump, breast height, crown base and mid-crown). Concentrations of total NSC (TNC, sum of sugars and starch), sugars and starch were much higher in bark than those in wood. On average, contents of sugars and starch accounted for 48 and 52 % of the TNC, respectively, and contents of TNC in bark, outer wood, and inner wood accounted for 34, 38, and 28 % of the stem total, respectively. Bark was the major pool of sugars in the stem (accounting for 50 % of the stem total on average), while outer wood was the major pool of starch (41 %). The concentration of sugars varied axially for all the conifers but did not for the broadleaved species. Mean concentrations of TNC, sugars and starch in stem varied by more than twofold among the species. However, there were no significant differences in these values for the species groups with different leaf phenology or wood types. Ignoring the radial, axial and inter-specific variations in NSC in stem would introduce large bias in estimating NSC storage at tree or ecosystem levels.

Tracking carbon within the trees

Tracking carbon within the trees

Comparing the intra-annual wood formation of three European species (Fagus sylvatica, Quercus petraea and Pinus sylvestris) as related to leaf phenology and non-structural carbohydrate dynamics

Monitoring cambial phenology and intra-annual growth dynamics is a useful approach for characterizing the tree growth response to climate change. However, there have been few reports concerning intra-annual wood formation in lowland temperate forests with high time resolution, especially for the comparison between deciduous and coniferous species. The main objective of this study was to determine how the timing, duration and rate of radial growth change between species as related to leaf phenology and the dynamics of non-structural carbohydrates (NSC) under the same climatic conditions. We studied two deciduous species, Fagus sylvatica L. and Quercus petraea (Matt.) Liebl., and an evergreen conifer, Pinus sylvestris L. During the 2009 growing season, we weekly monitored (i) the stem radial increment using dendrometers, (ii) the xylem growth using microcoring and (iii) the leaf phenology from direct observations of the tree crowns. The NSC content was also measured in the eight last rings of the stem cores in April, June and August 2009. The leaf phenology, NSC storage and intra-annual growth were clearly different between species, highlighting their contrasting carbon allocation. Beech growth began just after budburst, with a maximal growth rate when the leaves were mature and variations in the NSC content were low. Thus, beech radial growth seemed highly dependent on leaf photosynthesis. For oak, earlywood quickly developed before budburst, which probably led to the starch decrease quantified in the stem from April to June. For pine, growth began before the needles unfolding and the lack of NSC decrease during the growing season suggested that the substrates for radial growth were new assimilates of the needles from the previous year. Only for oak, the pattern determined from the intra-annual growth measured using microcoring differed from the pattern determined from dendrometer data. For all species, the ring width was significantly influenced by growth duration and not by growth rate, which differs from previous studies. The observed between-species difference at the intra-annual scale is key information for anticipating suitability of future species in temperate forests.

Agronomic and physiological performance of high-yielding wheat and rice in the lower reaches of Yangtze River of China

Understanding agronomic and physiological performance of crop high yield is essential to make strategies for breeding and crop management. This study aimed to identify major agronomic and physiological traits associated with high grain yields of rice and wheat in the wheat–rice rotation system in the lower reaches of Yangtze River of China. Two high-yielding cultivars, each for winter wheat and rice, were field-grown from 2008 to 2010 for 3 years. Two crop management treatments, the local farmer's practice (LFP) and improved high-yielding cultivation (IHC), were used. The IHC adopted two new techniques, i.e., site-specific nitrogen management in both rice and wheat and alternate wetting and moderate drying irrigation in rice and controlled soil drying irrigation in wheat. Across the 3 years, the IHC yielded an average 9.67tha−1 of wheat and 12.55tha−1 of rice with an annual total of 22.2tha−1, an increase of 26.8% when compared with that (17.5tha−1) under the LFP. The increased grain yield under the IHC was mainly attributed to expanded sink size as a result of more kernels per spike or more spikelets per panicle, increased kernel weight of wheat and increased percentage of filled grains of rice. When compared with LFP, IHC showed a greater percentage of productive tillers, more pre-anthesis nonstructural carbohydrate (NSC) storage in the stem, greater leaf photosynthetic rate and concentration of cytokinins in root exudates, higher activities of sucrose synthase and adenosine diphosphoglucose pyrophosphorylase in grains, more dry mater production during the grain filling period, and higher harvest index. We conclude that (1) expanding sink capacity through an increase in kernels per spike or spikelets per panicle, (2) enhancing grain filling efficiency by an increase in pre-anthesis NSC in the stem, and (3) increasing post-anthesis dry mater production by an enhancement in root activity during grain filling should be considered as three major strategies for further increases in grain yields of wheat and rice in the lower reaches of Yangtze River of China.

Increase in size and nitrogen concentration enhances seedling survival in Mediterranean plantations. Insights from an ecophysiological conceptual model of plant survival

Reduction in size and tissue nutrient concentration is widely considered to increase seedling drought resistance in dry and oligotrophic plantation sites. However, much evidence indicates that increase in size and tissue nutrient concentration improves seedling survival in Mediterranean forest plantations. This suggests that the ecophysiological processes and functional attributes relevant for early seedling survival in Mediterranean climate must be reconsidered. We propose a ecophysiological conceptual model for seedling survival in Mediterranean-climate plantations to provide a physiological explanation of the frequent positive relationship between outplanting performance and seedling size and nutrient concentration. The model considers the physiological processes outlined in the plantation establishment model of Burdett (Can J For Res 20:415–427, 1990), but incorporates other physiological processes that drive seedling survival, such as N remobilization, carbohydrate storage and plant hydraulics. The model considers that seedling survival in Mediterranean climates is linked to high growth capacity during the wet season. The model is for container plants and is based on three main principles, (1) Mediterranean climates are not dry the entire year but usually have two seasons of contrasting water availability; (2) summer drought is the main cause of seedling mortality; in this context, deep and large roots is a key trait for avoiding lethal water stress; (3) attainment of large root systems in the dry season is promoted when seedlings have high growth during the wet season. High growth is achieved when seedlings can divert large amount of resources to support new root and shoot growth. Functional traits that confer high photosynthesis, nutrient remobilization capacity, and non-structural carbohydrate storage promote high growth. Increases in seedling size and nutrient concentration strongly affect these physiological processes. Traits that confer high drought resistance are of low value during the wet season because hinder growth capacity. We provide specific evidence to support the model and finally we discuss its implications and the factors that may alter the frequent increase in performance with increase in seedling size and tissue nutrient concentration.

Water stress, shoot growth and storage of non‐structural carbohydrates along a tree height gradient in a tall conifer

We analysed concentrations of starch, sucrose, glucose and fructose in upper branch wood, foliage and trunk sapwood of Douglas-fir trees in height classes ranging from ~2 to ~57 m. Mean concentrations of non-structural carbohydrates (NSC) for all tissues were highest in the tallest height class and lowest in the lowest height class, and height-related trends in NSC were most pronounced in branches. Throughout a 17-month sampling period, mean values of branch NSC from the 57 m trees ranged between 30 and 377% greater than the 2 m trees. Branch NSC was inversely correlated with midday shoot water potential (Ψ(l)), shoot osmotic potential at full turgor (Ψ) and shoot extension. Temporal fluctuation in branch NSC was inversely correlated with height, and positively correlated with midday Ψ(l) , Ψ and shoot extension. The positive correlation between height and storage of NSC, and the negative correlation between NSC storage and shoot extension provide evidence that size-related growth decline in trees is not strongly associated with constraints on photosynthesis. The negative correlation between height and fluctuation in NSC suggests that mobilization of photosynthate in taller trees is constrained by some factor such as reductions in turgor-driven cell expansion or constraints on phloem transport.

Size-related auto-fragment production and carbohydrate storage in auto-fragment of Myriophyllum spicatum L. in response to sediment nutrient and plant density

Size-related asexual reproduction of submersed macrophytes is still poorly understood. Here, we investigate how size-related auto-fragmentation in Myriophyllum spicatum L. responds to sediment nutrients and plant density. An experiment was carried out with sediments containing two different nutrient levels and with two levels of plant density. The results show that sediment nutrients and plant density brought about a strong dependency of auto-fragment production and the amount of total non-structural carbohydrate (TNC) storage in auto-fragments on individual plant size (total plant biomass). However, these two factors acted differently on size dependency. Sediment nutrients positively affected auto-fragment production and the amount of TNC in auto-fragments of M. spicatum. High concentrations of sediment nutrients significantly increased these two traits in absolute value and the value relative to plant size. Although the auto-fragment biomass and the amount of TNC in auto-fragments did not differ between density treatments when plant size was considered, the absolute values of these two traits were much larger in the low plant density treatment than in the high plant density treatment, which suggested an indirect negative effect of plant density on the auto-fragmentation of M. spicatum. In addition, higher percentages of large auto-fragments (>100 mg) were produced by plants that grew in nutrient poor sediment and low plant density environment than plants in nutrient rich sediment and high plant density environment. These results do not solely highlight a size-dependent effect, but also a size-independent effect of auto-fragment production and the amount of TNC in auto-fragments of M. spicatum. Furthermore, such size-independent effects can be explained by the significant biomass partitioning differences and the similar TNC-concentrations in auto-fragments under different environmental conditions.

Unique responses of respiration, growth, and non-structural carbohydrate storage in sink tissue of conifer seedlings to an elevation gradient at timberline

The role of carbon balance, and particularly carbon sinks, to forest boundaries and climate responses is a major question in forest ecophysiology. At timberline, low-temperature limitations on carbon-sink processes of stem and especially root tissue have been implicated in hypotheses of the upper range limits to tree distributions. Studies on carbon sinks in root and stem tissue of trees at timberline typically report variation in only one carbon sink, such as either growth, respiration, or non-structural carbohydrates (NSCs). However, these carbon sinks may differ in their response to elevation. We asked how three carbon-sink processes in root and stem tissue (i.e. all tissue below the crown of needles) change in conifer seedlings growing from the lower (2450 m) to the upper (3000 m) edges of the timberline ecotone throughout their first growth season. We repeatedly measured respiration (mg −1 and individual −1), growth (relative growth rates [RGR] and biomass), and NSCs in root and stem tissue of Abies lasiocarpa and Pseudotsuga menziesii. RGR of root and stem tissue were less at the upper compared to lower elevation, but only for the first few weeks of the growing season. Total biomass of root and stem tissue was generally less at the upper site, apparently due to low early season RGR, but ultimately did not significantly differ between sites by the end of the growing season. Unlike growth, respiration rates (mg −1) did not differ between elevations during any period of the growing season. Nevertheless, total respiration of CO 2 from root and stem tissue (individual −1) was 22% less at the upper site, which was attributable to less biomass. NSCs of root and stem tissue, specifically starch, were overall greater at the upper site, particularly for A. lasiocarpa at the end of the season, which did not parallel spatiotemporal trends in growth or respiration. The differences in seasonal trends and the effects of elevation on carbon sinks indicate a degree of independence or uncoupling of growth, respiration, and NSCs of root and stem tissue, which is not commonly appreciated in hypotheses about physiological limitations for trees at timberline.

Optimal partitioning theory revisited: Nonstructural carbohydrates dominate root mass responses to nitrogen

Under optimal partitioning theory (OPT), plants preferentially allocate biomass to acquire the resource that most limits growth. Within this framework, higher root mass under low nutrients is often assumed to reflect an allocation response to build more absorptive surface. However, higher root mass also could result from increased storage of total nonstructural carbohydrates (TNC) without an increase in non-storage mass or root surface area. To test the relative contributions of TNC and non-storage mass as components of root mass responses to resources, we grew seedlings of seven northern hardwood tree species (black, red, and white oak, sugar and red maple, American beech, and black cherry) in a factorial light x nitrogen (N) greenhouse experiment. Because root mass is a coarse metric of absorptive surface, we also examined treatment effects on fine-root surface area (FRSA). Consistent with OPT, total root mass as a proportion of whole-plant mass generally was greater in low vs. high N. However, changes in root mass were influenced by TNC mass in all seven species and were especially strong in the three oak species. In contrast, non-storage mass contributed to increased total root mass under low N in three of the seven species. Root morphology also responded, with higher fine-root surface area (normalized to root mass) under low vs. high N in four species. Although biomass partitioning responses to resources were consistent with OPT, our results challenge the implicit assumption that increases in root mass under low nutrient levels primarily reflect allocation shifts to build more root surface area. Rather, root responses to low N included increases in: TNC, non-storage mass and fine-root surface area, with increases in TNC being the largest and most consistent of these responses. The greatest TNC accumulation occurred when C was abundant relative to N. Total nonstructural carbohydrates storage could provide seedlings a carbon buffer when respiratory or growth demands are not synchronized with photosynthesis, flexibility in responding to uncertain and fluctuating abiotic and biotic conditions, and increased access to soil resources by providing an energy source for mycorrhizae, decomposers in the rhizosphere, or root uptake of nutrients.

Phenology, growth and physiological adjustments of oil palm (Elaeis guineensis) to sink limitation induced by fruit pruning.

Despite its simple architecture and small phenotypic plasticity, oil palm has complex phenology and source-sink interactions. Phytomers appear in regular succession but their development takes years, involving long lag periods between environmental influences and their effects on sinks. Plant adjustments to resulting source-sink imbalances are poorly understood. This study investigated oil palm adjustments to imbalances caused by severe fruit pruning. An experiment with two treatments (control and complete fruit pruning) during 22 months in 2006-2008) and six replications per treatment was conducted in Indonesia. Phenology, growth of above-ground vegetative and reproductive organs, leaf morphology, inflorescence sex differentiation, dynamics of non-structural carbohydrate reserves and light-saturated net photosynthesis (A(max)) were monitored. Artificial sink limitation by complete fruit pruning accelerated development rate, resulting in higher phytomer, leaf and inflorescence numbers. Leaf size and morphology remained unchanged. Complete fruit pruning also suppressed the abortion of male inflorescences, estimated to be triggered at about 16 months before bunch maturity. The number of female inflorescences increased after an estimated lag of 24-26 months, corresponding to time from sex differentiation to bunch maturity. The most important adjustment process was increased assimilate storage in the stem, attaining nearly 50 % of dry weight in the stem top, mainly as starch, whereas glucose, which in controls was the most abundant non-structural carbohydrate stored in oil palm, decreased. The development rate of oil palm is in part controlled by source-sink relationships. Although increased rate of development and proportion of female inflorescences constituted observed adjustments to sink limitation, the low plasticity of plant architecture (constant leaf size, absence of branching) limited compensatory growth. Non-structural carbohydrate storage was thus the main adjustment process.

Carbohydrate storage in wood and bark of rubber trees submitted to different level of C demand induced by latex tapping

When the current level of carbohydrates produced by photosynthesis is not enough to meet the C demand for maintenance, growth or metabolism, trees use stored carbohydrates. In rubber trees (Hevea brasiliensis Muell. Arg.), however, a previous study (Silpi U., A. Lacointe, P. Kasemsap, S. Thanisawanyangkura, P. Chantuma, E. Gohet, N. Musigamart, A. Clement, T. Améglio and P. Thaler. 2007. Carbohydrate reserves as a competing sink: evidence from tapping the rubber tree. Tree Physiol. 27:881-889) showed that the additional sink created by latex tapping results not in a decrease, but in an increase in the non-structural carbohydrate (NSC) storage in trunk wood. In this study, the response of NSC storage to latex tapping was further investigated to better understand the trade-off between latex regeneration, biomass and storage. Three tapping systems were compared to the untapped Control for 2 years. Soluble sugars and starch were analyzed in bark and wood on both sides of the trunk, from 50 to 200 cm from the ground. The results confirmed over the 2 years that tapped trees stored more NSC, mainly starch, than untapped Control. Moreover, a double cut alternative tapping system, which produced a higher latex yield than conventional systems, led to even higher NSC concentrations. In all tapped trees, the increase in storage occurred together with a reduction in trunk radial growth. This was interpreted as a shift in carbon allocation toward the creation of reserves, at the expense of growth, to cover the increased risk induced by tapping (repeated wounding and loss of C in latex). Starch was lower in bark than in wood, whereas it was the contrary for soluble sugars. The resulting NSC was twice as low and less variable in bark than in wood. Although latex regeneration occurs in the bark, changes related to latex tapping were more marked in wood than in bark. From seasonal dynamics and differences between the two sides of the trunk in response to tapping, we concluded that starch in wood behaved as the long-term reserve compartment at the whole trunk level, whereas starch in bark was a local buffer. Soluble sugars behaved like an intermediate, ready-to-use compartment in both wood and bark. Finally, the dynamics of carbohydrate reserves appears a relevant parameter to assess the long-term performance of latex tapping systems.

Carbohydrate storage, survival, and growth of two evergreen Nothofagus species in two contrasting light environments

AbstractA number of traits have been attributed important roles in tolerance of shade by plants. Some explanations emphasize traits enhancing net carbon gain; others emphasize energy conservation traits such as storage of non‐structural carbohydrates (NSC). To date, cross‐species studies have provided mixed support for the role of NSC storage in low‐light survival. We examined NSC status, survival, biomass, and growth of large seedlings of two evergreen species of differing shade tolerance (Nothofagus nitida and N. dombeyi) grown in deep shade and 50% light for two growing seasons. We expected to find higher NSC concentration in the more shade‐tolerant N. nitida and since allocation to storage involves sacrificing growth, higher growth rate in the shade‐intolerant N. dombeyi. NSC concentration of both species was &gt;twofold higher in 50% light than in deep shade, and in roots and stems did not differ significantly between species in either environment. NSC contents per plant were also similar between dead and living plants in deep shade. N. dombeyi outgrew N. nitida in 50% light, while this pattern was reversed in deep shade. Survival in deep shade was not correlated with NSC concentration. Leaf mass fraction was similar between species in 50% light, but lower in N. dombeyi in deep shade. Results provide little evidence of a link between carbohydrate storage and low‐light survival in Nothofagus species, and support the view that understorey survival is primarily a function of net carbon gain. Patterns of variation in NSC concentration of the temperate species we studied are likely dominated by more important influences than adaptation to shade, such as limitation of growth or adaptation to cold stress.

Roots of Australian alpine plant species contain high levels of stored carbohydrates independent of post-fire regeneration strategy

The relationships between root morphology, level of stored non-structural carbohydrates and post-fire regeneration strategy were investigated in 37 Australian alpine plant species: 6 reseeders, 14 resprouters and 17 capable of both reseeding and resprouting. High concentrations of stored carbohydrate (up to 61.8% DW) were a feature of most species, with more than half of the 37 species containing non-structural carbohydrate concentrations of more than 10%. Fructan was the major reserve polysaccharide in 32 of the 37 species, with particularly high concentrations in the Asteraceae (up to 43.1%). Herbaceous species stored higher concentrations of carbohydrates and had fleshier roots than did shrub species, but swollen underground storage organs were found in only one species (Microseris scapigera sensu Willis1, Asteraceae). There was no significant relationship between post-fire regeneration strategy and either root morphology or level of carbohydrate storage, contrasting with results from drier, Mediterranean environments. Root storage of high levels of carbohydrate, and especially fructan, in Australian alpine species could therefore result from an adaptation to the alpine environment, such as a need for annual regeneration of leaf tissue in alpine conditions.

Growth of Dactylis glomerata along a light gradient in the central Appalachian region of the eastern USA: I. Dry matter production and partitioning

Microsite influences on resource allocation related to silvopastoral swards undergoing repeated defoliation are unclear. Defoliating traditional pasture species according to canopy development criteria may not be appropriate when the plants grow as understory crops in woodlots. An experiment using container-grown plants was conducted under field conditions to determine how open (O), shaded woodland (W) and open-to-shaded woodland transition zones (E O, E W), representing the partial shade conditions found in silvopastoral settings, influenced productivity and dry matter allocation in Dactylis glomerata L. Plants established in spring (SP) and late summer (LS) were clipped each time mean sward height reached 20 cm. Dry matter production differed for SP and LS plantings, and was greater in O than W sites. Dry matter allocation to shoots of SP plants indicated some agronomic benefit associated with the E O and E W sites. Leaf mass did not vary with irradiance, but stembase mass did. The SP plants had numerous small tillers whereas LS plants were composed of comparatively fewer but more massive tillers. Plants growing at O, E O and E W were similar, whereas plants in the W site were smaller and had the fewest tillers regardless of planting time. Results suggest that defoliation management for shaded components of silvopastoral systems should be based on some index other than sward height, and that management criteria might change during the growing season. Less intensive removal (e.g., 10-cm rather that 5-cm residue) should be considered for defoliated, shade grown plants, since this could allow more stembase and nonstructural carbohydrate storage tissues to remain.

Using L‐systems for modeling source–sink interactions, architecture and physiology of growing trees: the L‐PEACH model

Functional-structural plant models simulate the development of plant structure, taking into account plant physiology and environmental factors. The L-PEACH model is based on the development of peach trees. It demonstrates the usefulness of L-systems in constructing functional-structural models. L-PEACH uses L-systems both to simulate the development of tree structure and to solve differential equations for carbohydrate flow and allocation. New L-system-based algorithms are devised for simulating the behavior of dynamically changing structures made of hundreds of interacting, time-varying, nonlinear components. L-PEACH incorporates a carbon-allocation model driven by source-sink interactions between tree components. Storage and mobilization of carbohydrates during the annual life cycle of a tree are taken into account. Carbohydrate production in the leaves is simulated based on the availability of water and light. Apices, internodes, leaves and fruit grow according to the resulting local carbohydrate supply. L-PEACH outputs an animated three-dimensional visual representation of the growing tree and user-specified statistics that characterize selected stages of plant development. The model is applied to simulate a tree's response to fruit thinning and changes in water stress. L-PEACH may be used to assist in horticultural decision-making processes after being calibrated to specific trees.