Non-structural carbohydrate concentrations in tree organs vary across biomes and leaf habits, but are independent of the fast-slow plant economic spectrum.

Carbohydrate reserves play a vital role in plant survival during periods of negative carbon balance. Under a carbon-limited scenario, we expect a trade-offs between carbon allocation to growth, reserves, and defense. A resulting hypothesis is that carbon allocation to reserves exhibits a coordinated variation with functional traits associated with the 'fast-slow' plant economics spectrum. We tested the relationship between non-structural carbohydrates (NSC) of tree organs and functional traits using 61 angiosperm tree species from temperate and tropical forests with phylogenetic hierarchical Bayesian models. Our results provide evidence that NSC concentrations in stems and branches are decoupled from plant functional traits. while those in roots are weakly coupled with plant functional traits. In contrast, we found that variation between NSC concentrations in leaves and the fast-slow trait spectrum was coordinated, as species with higher leaf NSC had trait values associated with resource conservative species, such as lower SLA, leaf N, and leaf P. We also detected a small effect of leaf habit on the variation of NSC concentrations in branches and roots. Efforts to predict the response of ecosystems to global change will need to integrate a suite of plant traits, such as NSC concentrations in woody organs, that are independent of the 'fast-slow' plant economics spectrum and that capture how species respond to a broad range of global change drivers.

Presence of inulin in sunflower (Helianthus annuus L.) grown under high irradiance.

Fructans are important polysaccharides synthesized from sucrose which are present in about 12-15% of angiosperms. Sunflower (Helianthus annuus L.) is considered a non-fructan bearing plant even though its close relative, Helianthus tuberosus, accumulates the inulin type of the polymer in large amounts. Previous work suggested that putative fructan-synthesizing enzymes may be expressed in sunflower, but only very limited amounts of the trisaccharide isokestose were found in stems of plants storing high levels of sucrose due to capitulum removal. The present work is aimed at investigating whether intact sunflower plants may indeed synthesize fructans in any of its parts when grown in conditions that favor sucrose availability. Plants were grown in the field at a low density, resulting in a high light availability and low competition for resources, in comparison with controls (usual crop planting density). Plants were harvested at anthesis. Thinned treatment led to an increase in carbohydrates level especially in the capitulum. Carbohydrates analysis of this tissue in thinned plants revealed, for the first time in this species, the presence of inulin-type fructans. The amount of each member of the series appeared to decline starting from isokestose, being DP = 15 the longest fructan detected. Results suggest that, in sunflower, fructans could be synthesized only when sucrose availability exceeds a high threshold, which may not be attained under usual growing conditions. Given the relationship between fructans and tolerance to abiotic stresses including drought, the present finding opens a new perspective for breeding and management of this crop.

Metabolic adjustments during starvation in Daphnia pulex.

Zooplankton organisms face a variable food supply in their habitat. Metabolic adjustments during periods of starvation were analysed from changes in metabolite level to gene expression in the microcrustacean Daphnia pulex during starvation. The animals exploited their carbohydrate stores first, but their lipid and protein reserves were also degraded, albeit more slowly. Glycogenolysis and probably gluconeogenesis led to hyperglycaemia after 16 h of starvation. The concentration of α-ketoglutarate and the rate of oxygen consumption also reached maxima during this period. Nuclear HIF-1α levels and α-ketoglutarate concentration showed inverse correlation. Effects of this 2-oxoacid on prolyl hydroxylase activity, HIF-1α stability and the role of this transcription factor in the changes of the expression level of several putatively HIF-1-mediated metabolic genes are discussed. Transcriptome profiling via RNA-Seq revealed a downregulation of genes for protein biosynthesis and an upregulation of genes for carbohydrate metabolism during starvation. Thus, the adjustments of energy metabolism in response to food deprivation were quantified from the level of metabolites, signal transduction and gene expression, and possible connections of the respective dynamics of observed changes were analysed.

Microplastic burden in Daphnia is aggravated by elevated temperatures.

Contamination of freshwater habitats with microplastic is threatening particularly filter-feeders within the aquatic community. Using Daphnia magna and Daphnia pulex as models, the effects of food supply and temperature on the ingestion of polystyrene spheres (diameter 1 µm, concentration of 200 ng*ml-1) was analysed. The ingestion rates of microplastic beads were increased in conditions of low food and high temperatures, reflecting the complex regulation patterns of the water current generated by the animals' thoracic limbs. Maximal enrichment of 1160 times the concentration in the ambient medium was observed within one hour. Analyses of the impact of microplastic ingestion on physiological parameters used the carbohydrate concentration as an indicator for the animals' metabolic state. Exposure to the microplastic beads for three days in the presence or absence of Desmodesmus subspicatus did not affect the animals' glycogen reserves beyond the response to the prevailing food and temperature conditions. Projecting the insights from laboratory experiments to the habitat situation, increased burdens of microplastic particles can be expected in filtering zooplankton organisms in warm water and scarce supply of food, like the clear-water phase of lakes in the summer.

Phosphorus and potassium effects on taproot C and N reserve pools and long-term persistence of alfalfa (Medicago sativa L.).

Improved P and K nutrition can enhance yield and persistence of alfalfa (Medicago sativa L.) grown on low fertility soils, but it is unknown if the improved agronomic performance is associated with greater taproot N and C reserves. Our objective was to use cluster analysis to determine how alfalfa plant persistence is altered by P and K fertilization, and determine if changes in specific taproot C and/or N reserves were associated with alfalfa plant death. Taproots were dug and plants counted in May and December of each year and taproots analyzed for P, K, starch, sugar, amino-N, and soluble protein. K-means clustering was used to create six clusters that were subsequently compared using two-sample t-tests. Low K in herbage and taproots was associated with low yield and poor persistence of the Low and Very Low clusters and taproots of these plants generally had low starch, protein, and amino-N concentrations. Plants died primarily between May and December. Plant persistence of the low yielding, P-deficient Medium cluster was high and associated with high starch concentrations. Low amino-N concentrations in taproots may provide an early indication of potential plant death because these were evident in poor-persisting Low and Very Low clusters early in the study.

Potential effect of atmospheric warming on grapevine phenology and post-harvest heat accumulation across a range of climates.

Carbohydrates are accumulated within the perennial structure of grapevines when their production exceeds the requirements of reproduction and growth. The period between harvest and leaf-fall (the post-harvest period) is a key period for carbohydrate accumulation in relatively warmer grape-growing regions. The level of carbohydrate reserves available for utilisation in the following season has an important effect on canopy growth and yield potential and is therefore an important consideration in vineyard management. In a warming climate, the post-harvest period is lengthening and becoming warmer, evidenced through studies in wine regions worldwide that have correlated recent air temperature increases with changing grapevine phenology. Budbreak, flowering, veraison, and harvest have all been observed to be occurring earlier than in previous decades. Additionally, the final stage of the grapevine phenological cycle, leaf-fall, occurs later. This study explored the potential for increased post-harvest carbohydrate accumulation by modelling heat accumulation following harvest dates for the recent climate (1975-2004) and two warmer climate projections with mean temperature anomalies of +1.26 and +2.61 °C. Summaries of post-harvest heat accumulation between harvest and leaf-fall were produced for each of Australia's Geographical Indications (wine regions) to provide comparisons from the base temperatures to projected warmer conditions across a range of climates. The results indicate that for warmer conditions, all regions observe earlier occurring budbreak and harvest as well as increasing post-harvest growing degree days accumulation before leaf-fall. The level of increase varies depending upon starting climatic condition, with cooler regions experiencing the greatest change.

Endogenous hormone concentrations correlate with fructan metabolism throughout the phenological cycle in Chrysolaena obovata.

BACKGROUND AND AIMS: Chrysolaena obovata, an Asteraceae of the Brazilian Cerrado, presents seasonal growth, marked by senescence of aerial organs in winter and subsequent regrowth at the end of this season. The underground reserve organs, the rhizophores, accumulate inulin-type fructans, which are known to confer tolerance to drought and low temperature. Fructans and fructan-metabolizing enzymes show a characteristic spatial and temporal distribution in the rhizophores during the developmental cycle. Previous studies have shown correlations between abscisic acid (ABA) or indole acetic acid (IAA), fructans, dormancy and tolerance to drought and cold, but the signalling mechanism for the beginning of dormancy and sprouting in this species is still unknown. METHODS: Adult plants were sampled from the field across phenological phases including dormancy, sprouting and vegetative growth. Endogenous concentrations of ABA and IAA were determined by GC-MS-SIM (gas chromatography-mass spectrometry-selected ion monitoring), and measurements were made of fructan content and composition, and enzyme activities. The relative expression of corresponding genes during dormancy and sprouting were also determined. KEY RESULTS: Plants showed a high fructan 1-exohydrolase (EC 3.2.1.153) activity and expression during sprouting in proximal segments of the rhizophores, indicating mobilization of fructan reserves, when ABA concentrations were relatively low and precipitation and temperature were at their minimum values. Concomitantly, higher IAA concentrations were consistent with the role of this regulator in promoting cell elongation and plant growth. With high rates of precipitation and high temperatures in summer, the fructan-synthesizing enzyme sucrose:sucrose 1-fructosyltransferase (EC 2.4.1.99) showed higher activity and expression in distal segments of the rhizophores, which decreased over the course of the vegetative stage when ABA concentrations were higher, possibly signalling the entry into dormancy. CONCLUSIONS: The results show that fructan metabolism correlates well with endogenous hormone concentrations and environmental changes, suggesting that the co-ordinated action of carbohydrate metabolism and hormone synthesis enables C. obovata to survive unfavourable field conditions. Endogenous hormone concentrations seem to be related to regulation of fructan metabolism and to the transition between phenophases, signalling for energy storage, reserve mobilization and accumulation of oligosaccharides as osmolytes.

Seed germination and seedling development in response to submergence in tree species of the Central Amazonian floodplains.

Successful germination and seedling establishment are crucial steps for maintenance and expansion of plant populations and recovery from perturbations. Every year the Amazon River and its tributaries overflow and flood the adjacent forest, exerting a strong selective pressure on traits related to seedling recruitment. We examined seed characteristics, stored reserves, germination, seedling development and survival under water of eight representative tree species from the lower portions of the flood-level gradient to identify adaptive strategies that contribute to their regeneration in this extreme ecosystem. Submerged seedlings were assessed for longevity and survival until they showed symptoms of injury. At this point, the remaining healthy seedlings were planted in unsaturated soil to monitor recovery after re-exposure to air over 30 days. All small (seed mass ≤0.17 g) seeds had epigeal phanerocotylar-type germination, a trait that would allow plants to acquire light and CO2 in the shortest time. Cell wall storage polysaccharide was a major component of all seeds, suggesting plant investment in structural reserves. Seven of the eight species germinated and formed healthy seedlings under water that endured submersion without any apparent injury for periods of 20-115 days, depending on the species. Seedlings of some species changed the direction of root growth and grew towards the surface of the water, which might have increased the uptake of oxygen to the tissues. Only one of the seven species did not survive re-exposure to air. Species able to germinate and produce seedlings under submersion, which subsequently are able to establish in aerated soils, would have more time available for terrestrial growth. This is critical for colonization of lower portions of the flood-level gradient where establishment is constrained by the short terrestrial phase that precedes the next flood.

The tolerance of grain amaranth (Amaranthus cruentus L.) to defoliation during vegetative growth is compromised during flowering.

The biochemical processes underlying variations of tolerance are often accompanied by source-sink transitions affecting carbon (C) metabolism. We investigated the tolerance of Amaranthus cruentus L. to total mechanical defoliation through development and in different growing seasons. Defoliated A. cruentus recovered ∼80% of their above-ground biomass and ∼100% of grain yield compared to intact plants if defoliation occurred early during ontogeny, but could not compensate when defoliation occurred during flowering. Tolerance index was higher in the summer season (-0.3) than in the winter season (-0.7). Overall, defoliation tolerance was closely related to phosphoenolpyruvate carboxylase (PEPC) activity in leaves and the subsequent accumulation of starch (∼500 µmol/gDW) and sucrose (∼140 µmol/gDW) in stems and roots. Thus, A. cruentus accumulated sufficient C in roots and stem to allow branching and shoot re-growth after defoliation, but it only possessed sufficient C reserves to maintain <19% seed yield in the absence of new vegetative tissue. Seed size was larger during the warm season but it was not affected by foliar damage. Seed chemical composition was altered by defoliation at flowering. We conclude that A. cruentus defoliation tolerance depends on both, the re-allocation of starch from stem and roots, and the activation of dormant meristems before flowering to generate new photosynthetic capacity to sustain seed filling.

Distribution of above-ground and below-ground carbohydrate reserves in adult trees of two contrasting broad-leaved species (Quercus petraea and Fagus sylvatica).

• The exhaustive distribution of total carbohydrate reserves was investigated in oak and beech trees that were approx. 40 yr old and felled at two dates (October 1999 and June 2000) to estimate variations in reserve amounts at the tree level. • The total nonstructural carbohydrate (TNC) content was highest in the twigs and coarse roots, reaching 10 g 100 g-1 dry matter and 12 g 100 g-1 dry matter for beech and oak twigs, and 13 g 100 g-1 dry matter and 16 g 100 g-1 dry matter for beech and oak roots, respectively. Similar distribution in tree carbohydrates was observed for both species and date, but with contrasting starch/sugar sharing. • Scaling-up to reserve amounts at tree level was performed with extensive organ biomass measurements. Based on the respective biomass of the organs, stem and roots contained the highest quantity of reserves. Between October (before leaf fall) and June (after bud-burst and leaf area index expansion) oaks used double the reserves of beeches. • These differences in the allocation of carbohydrate reserves could arise from differential needs for spring growth and winter maintenance respiration between the two species.

Interspecific and intraspecific variation in tree seedling survival: effects of allocation to roots versus carbohydrate reserves.

We examined interspecific and intraspecific variation in tree seedling survival as a function of allocation to carbohydrate reserves and structural root biomass. We predicted that allocation to carbohydrate reserves would vary as a function of the phenology of shoot growth, because of a hypothesized tradeoff between aboveground growth and carbohydrate storage. Intraspecific variation in levels of carbohydrate reserves was induced through experimental defoliation of naturally occurring, 2-year-old seedlings of four northeastern tree species -Acer rubrum, A. saccharum, Quercus rubra, and Prunus serotina- with shoot growth strategies that ranged from highly determinate to indeterminate. Allocation to root structural biomass varied among species and as a function of light, but did not respond to the defoliation treatments. Allocation to carbohydrate reserves varied among species, and the two species with the most determinate shoot growth patterns had the highest total mass of carbohydrate reserves, but not the highest concentrations. Both the total mass and concentrations of carbohydrate reserves were significantly reduced by defoliation. Seedling survival during the year following the defoliation treatments did not vary among species, but did vary dramatically in response to defoliation. In general, there was an approximately linear relationship between carbohydrate reserves and subsequent survival, but no clear relationship between allocation to root structural biomass and subsequent survival. Because of the disproportionate amounts of reserves stored in roots, we would have erroneously concluded that allocation to roots was significantly and positively related to seedling survival if we had failed to distinguish between reserves and structural biomass in roots.

Winter starch reserves of white oak as a predictor of attack by the twolined chestnut borer, Agrilus bilineatus (Weber) (Coleoptera: Buprestidae).

The twolined chestnut borer, Agrilus bilineatus (Weber) (Coleoptera: Buprestidae), attacks oaks (Quercus spp.) and is associated with extensive mortality of trees in the eastern deciduous forests of North America. We tested the hypothesis that winter starch reserves of oak roots are an indicator of tree vigor and that only trees low in stored starch would be attacked by A. bilineatus. We measured the levels of stored starch in the roots of 200 non-infested healthy white oaks during the dormant season and determined their correlation with A. bilineatus attacks the following spring. There was a significant increase in A. bilineatus captures on sticky traps with a decrease in winter starch reserves. Trees low in stored starch that were also stressed by phloem-girdling attracted 3.7 times as many beetles as did non-girdled trees that were low in starch. However, non-girdled trees that had low winter starch reserves were also attacked. Only oaks that had had extremely low winter root starch reserves (<5mg/g dry weight of root sapwood tissue) were heavily attacked by A. bilineatus and subsequently died. One third of non-girdled low starch trees and 67% of phloem-girdled low starch trees died, whereas none of the trees with root starch >5 mg/g dry wt died. These results indicate that winter starch reserves are a good predictor of A. bilineatus attack.

Effect of Ditylenchus dipsaci on Alfalfa Mortality, Winterkill, and Yield.

Ditylenchus dipsaci-infected and noninfected alfalfa plants in a naturally infested field were studied from July 1980 to September 1982. Forty-one percent of the plants died during the study. Ninety-seven percent of the plants that died were infected with D. dipsaci. Sixty-nine percent of the observed mortality occurred during winter. Forage yield of infected plants was significantly lower than yield of noninfected plants at each harvest. Stored carbohydrates in infected plants were significantly lower than in noninfected plants. In a controlled environment test, significantly greater mortality occurred in frozen severely infected plants than in frozen noninfected plants, while no mortality occurred in severely infected or noninfected plants that were not frozen. Both forage yield and stored carbohydrates were significantly lower in severely infected than noninfected, non-frozen plants. Mortality in greenhouse-grown plants that were transplanted to field plots was significantly greater in D. dipsaci-infected plants than in noninfected plants after one winter.