Changes In Non-structural Carbohydrates Research Articles

Facilitation drives tree seedling survival at alpine treelines

Abstract Alpine treelines are considered ecological monitors recording the impacts of climate change on trees and forests. To date, most treeline research has focused on how climate change drives treeline dynamics. However, little is known about how biotic interactions mediate treeline shifts, particularly in the case of tree recruitment, a bottleneck of treeline dynamics. We hypothesized that inter- and intraspecific facilitation determined the establishment and survival of tree seedlings at alpine treelines. To test this hypothesis, 630 Abies georgei var. smithii seedlings with different ages (4–6, 7–9 and 10–15 years old) were transplanted into three growth habitats (canopy-in, canopy-out and meadow) across the alpine treeline ecotone (4300–4500 m) in the Sygera Mountains, on the southeastern Qinghai-Tibetan Plateau. Microclimate, height growth, mortality rates and leaf functional traits of transplanted seedlings were measured over 3 years. We found that the variations in leaf functional traits were driven by microclimate. After the transplantation, the leaf concentrations of soluble sugars and starch and C:P ratio increased, whereas leaf size decreased. The resource use of seedlings gradually shifted to a more conservative strategy as indicated by changes in non-structural carbohydrates and nutrient concentrations. Radiation, temperature and moisture conditions, mediated by plant interactions, influenced seedling mortality and annual growth by affecting leaf morphological traits. Our findings illustrate how facilitation plays a crucial role in altering solar radiation and leaf trait functioning, determining seedling survival and growth at alpine treelines. We provide new insights into the underlying mechanisms for tree establishment and alpine treeline shifts in response to climate change.

Species-specific responses of C and N allocation to N addition: evidence from dual 13C and 15N labeling in three tree species

Soil nitrogen (N) availability affects plant carbon (C) utilization. However, it is unclear how various tree functional types respond to N addition in terms of C assimilation, allocation, and storage. Here, a microcosm experiment with dual 13C and 15N labeling was conducted to study the effects of N addition (i.e., control, 0 g N kg−1; moderate N addition, 1.68 g N kg−1; and high N addition, 3.36 g N kg−1 soil) on morphological traits, on changes in nonstructural carbohydrates (NSC) in different organs, as well as on C and N uptake and allocation in three European temperate forest tree species (i.e., Acer pseudoplatanus, Picea abies and Abies alba). Our results demonstrated that root N uptake rates of the three tree species increased by N addition. In A. pseudoplatanus, N uptake by roots, N allocation to aboveground organs, and aboveground biomass allocation significantly improved by moderate and high N addition. In A. alba, only the high N addition treatment considerably raised aboveground N and C allocation. In contrast, biomass as well as C and N allocation between above and belowground tissues were not altered by N addition in P. abies. Meanwhile, NSC content as well as C and N coupling (represented by the ratio of relative 13C and 15N allocation rates in organs) were affected by N addition in A. pseudoplantanus and P. abies but not in A. alba. Overall, A. pseudoplatanus displayed the highest sensitivity to N addition and the highest N requirement among the three species, while P. abies had a lower N demand than A. alba. Our findings highlight that the responses of C and N allocation to soil N availability are species-specific and vary with the amount of N addition.

The mobilization and transport of newly fixed carbon are driven by plant water use in an experimental rainforest under drought.

Non-structural carbohydrates (NSCs) are building blocks for biomass and fuel metabolic processes. However, it remains unclear how tropical forests mobilize, export, and transport NSCs to cope with extreme droughts. We combined drought manipulation and ecosystem 13CO2 pulse-labeling in an enclosed rainforest at Biosphere 2, assessed changes in NSCs, and traced newly assimilated carbohydrates in plant species with diverse hydraulic traits and canopy positions. We show that drought caused a depletion of leaf starch reserves and slowed export and transport of newly assimilated carbohydrates below ground. Drought effects were more pronounced in conservative canopy trees with limited supply of new photosynthates and relatively constant water status than in those with continual photosynthetic supply and deteriorated water status. We provide experimental evidence that local utilization, export, and transport of newly assimilated carbon are closely coupled with plant water use in canopy trees. We highlight that these processes are critical for understanding and predicting tree resistance and ecosystem fluxes in tropical forest under drought.

Eco-physiological response mechanism of Tamarix chinensis to soil water changes in coastal wetlands of the Yellow River Delta

Elucidating the effect of soil moisture on the adaptation of dominant plants in coastal wetlands is important for predicting the evolution of vegetation in the region. In this paper, Tamarix chinensis, a dominant species in the Yellow River Delta, was used as the object to study the changes of its growth and physiological parameters with increasing soil salinity under different moisture conditions (normal watering, persistent drought and persistent waterlogging). Different salt stress (2‰, 5‰, 8‰, 12‰, 16‰, and 20‰) using pot experiments was also used to reveal the mechanism of soil moisture on its salt tolerance. The results showed that the relative growth rate between 5‰-8‰ soil salinity was the largest, and growth was significantly inhibited above 20‰. Among different moisture conditions, the difference in relative growth rate under normal watering and persistent drought were nonsignificant, while both were significantly lower than those under persistent waterlogging. With increasing soil salinity, relative water content and total chlorophyll content significantly decreased, and cell membrane permeability (malondialdehyde), sodium ion, osmoregulatory substances (proline, soluble protein), and protective enzyme activity (SOD) significantly increased, while changes in non-structural carbohydrates (NSC) were not significant. Compared with normal watering and persistent waterlogging, persistent drought had the lowest leaf relative water content, total chlorophyll content, and sodium ions, and the highest cell membrane permeability, osmoregulatory substances and protective enzyme activity. With increasing treatment time, the relative leaf water content and total chlorophyll content significantly decreased, and cell membrane permeability, osmoregulatory substances and protective enzyme activity increased more significantly than normal watering and persistent waterlogging. NSC increased under normal watering and persistent waterlogging, while significantly decreased under persistent drought. Correlation analysis showed that the relationships between sodium ions, total chlorophyll content and malondialdehyde were various under different moisture conditions. Under persistent drough, malondialdehyde was significantly positively correlated with relative conductivity, superoxide dismutase, proline, soluble protein and soluble sugar. Total chlorophyll content was the key indicator reflecting the salt and waterlogging tolerance of T. chinensis under normal watering and persistent waterlogging, while cell membrane damage was under persistent drought. In summary, T. chinensis has strong salt and waterlogging tolerance, but persistent drought with salt stress can have serious impacts on its growth and survival.

Unveiling resilience mechanisms of Quercus ilex seedlings to severe water stress: Changes in non-structural carbohydrates, xylem hydraulic functionality and wood anatomy

Over the last few decades, extensive dieback and mortality episodes of Quercus ilex L. have been documented after severe drought events in many Mediterranean forests. However, the underlying physiological, anatomical, and biochemical mechanisms remain poorly understood. We investigated the physiological and biochemical processes linked to embolism formation and non-structural carbohydrates (NSCs) dynamics in Q. ilex seedlings exposed to severe water stress and rewatering. Measurements of leaf gas exchange, water relations, non-structural carbohydrates, drought-related gene expression, and anatomical changes in wood parenchyma were assessed. Under water stress, the midday stem water potential dropped below - 4.5 MPa corresponding to a ~ 50 % loss of hydraulic conductivity. A 70 % reduction in stomatal conductance led to a strong depletion of wood NSCs. Starch consumption, resulting from the upregulation of the β-amylase gene BAM3, together with the downregulation of glucose (GPT1) and sucrose (SUC27) transport genes, suggests glucose utilization to sustain cellular metabolism in the wood parenchyma. After rewatering, the presence of residual xylem embolism led to an incomplete recovery of leaf gas exchanges. However, the partial restoration of photosynthesis allowed the accumulation of new starch reserves in the wood parenchyma and the production of new narrower vessels. In addition, changes in the cell wall composition of the wood parenchyma fibers were observed. Our findings indicate that thirty days of rewatering were sufficient to restore the NSCs reserves and growth rates of Q. ilex seedlings and that the carryover effects of water stress were primarily caused by hydraulic dysfunction.

Changes of non-structural carbohydrates and nitrogen contents of needles and twigs in Pinus sylvestris var. mongolica plantations along an aridity gradient

With six Pinus sylvestris var. mongolica plantations (Huinan, Xifeng, Fujia, Zhanggutai, Naiman and Wulanaodu) along an aridity gradient in the Horqin sandy land, we examined the changes in non-structural carbohydrates (NSCs) and nitrogen (N) contents of current and one-year-old needles and twigs, to explore the carbon supply and demand status as well as the nutrient accumulation strategies of P. sylvestris var. mongolica under drought. The results showed that the contents of NSCs and soluble sugars in needles and twigs of P. sylvestris var. mongolica plantations significantly decreased with increasing aridity. From the most humid site (Huinan) to the most aridity site (Wulanaodu), the soluble sugar contents in current and one-year old needles of P. sylvestris var. mongolica decreased from 12.8% and 12.5% to 9.0% and 9.5%, respectively. The soluble sugar contents in current-year old twigs decreased from 15.6% to 9.2%. With increasing aridity, the starch contents in needles and twigs remained relatively stable, soluble sugars/starch ratio in current and one-year old needles decreased, the N contents in current and one-year old twigs significantly increased. The P. sylvestris var. mongolica plantations in the Horqin sandy land consumed soluble sugar storage under drought, resulting in a risk of mortality from 'carbon starvation'. P. sylvestris var. mongolica tended to maintain stable starch storage and accumulate N in twigs to cope with long-term drought stress.

Changes in Non-Structural Carbohydrates, Wood Properties and Essential Oil During Chemically-Induced Heartwood Formation in Dalbergia odorifera.

The highly valuable heartwood of Dalbergia odorifera T. Chen, known as Jiang Xiang in traditional Chinese medicine, is formed very slowly, and there is a need to better understand the process and promote heartwood formation. Chemical induction is considered to be one of the promising methods to induce heartwood formation. However, to date no method has been proved effective for D. odorifera as little is known about biochemical and physiological changes during heartwood development. Three potential heartwood induction substances viz. acetic acid, sodium chloride, and hydrogen peroxide solutions were injected into the trunk of D. odorifera to determine the effect on heartwood formation and physiological activity. Non-structural carbohydrates, lipids, wood properties, and essential oil were assessed in the post-treatment period. As also observed in the formation of natural heartwood, chemical-induced Jiang Xiang production was accompanied by sapwood dehydration, non-structural carbohydrates consumption, and synthesis of heartwood substances. As the heartwood substances accumulated, basic density and essential oil content increased gradually, thereby Jiang Xiang was finally produced. In this process, physiological parameters of discolored sapwood gradually evolved to resemble those of natural heartwood. Hydrogen peroxide-induced Jiang Xiang was closest to natural heartwood, and the essential oil components met the standards for high-quality Jiang Xiang, while the induction effects of acetic acid and sodium chloride were unsatisfactory. Thus, this study indicates that hydrogen peroxide has the potential to induce Jiang Xiang production in Dalbergia odorifera.

The Effects of Different Degrees of Defoliation on ‘Seolhyang’ Strawberry’s Fruit Characteristics, Plant Growth and Changes in Nonstructural Carbohydrates

The Effects of Different Degrees of Defoliation on ‘Seolhyang’ Strawberry’s Fruit Characteristics, Plant Growth and Changes in Nonstructural Carbohydrates

Diurnal vertical and seasonal changes in non-structural carbohydrates in Marandu palisade grass

AbstractForage is the primary feed source for livestock in tropical regions and energy is one of the most important nutrients for ruminant nutrition. The effects of harvest management of Marandu palisade grass (Brachiaria brizantha cv. Marandu Syn. Urochloa brizantha cv. Marandu) on non-structural carbohydrate (NSC) concentrations were evaluated. A plot (Experiment 1) and a greenhouse study (Experiment 2) were conducted in 2013–14. In Experiment 1, treatments were the factorial arrangement of two harvest times and two vertical canopy layers (upper and intermediate), distributed in a completely randomized design with five replicates. In Experiment 2, treatments were the factorial arrangement of six harvest times and two morphological fractions (leaf blade and pseudostem). In both experiments, NSC concentration increased during the day. Upper and intermediate canopy layers had greater NSC concentration at 15.00 than 06.00 h during spring and summer. In addition, the magnitude of NSC increase was greater in the upper than intermediate canopy layer and in spring than summer. Marandu palisade grass shows greater digestibility in the afternoon than morning, representing an opportunity to optimize energy concentration through harvest management.

Clipping frequency but not nutrients affect the architecture and non-structural carbohydrates of a browsing lawn

“Grazing lawns” are well-known effects of large mammalian herbivores on a short grass sward that is maintained through a positive feedback effect; plants are re-grazed due to their increased palatability. However, there are few studies that have addressed the notion of “browsing lawns.” I simulated browsing and added nutrients to Acacia nigrescens, a palatable and dominant African tree, to examine the effects on the architecture of these trees in a greenhouse. I recorded investments in non-structural carbohydrates and crude protein between roots and stems as a possible mechanism behind such architectural changes. There was a significant decrease in mean branch internode length and an increase in the branching ratio with clipping; the last mentioned indicated that there was more extensive tree branching. There were no significant effects on plant architecture of nutrient addition. There were changes in non-structural carbohydrates and crude protein between the roots and stems due to clipping but not to nutrient addition. Non-structural carbohydrate concentrations decreased with clipping in the roots and increased in the stems. Crude protein decreased in the stems with clipping frequency but not in the roots. There was an increase in root:shoot ratio of crude protein with increasing clipping frequency. A. nigrescens performs like a browsing lawn because of the changes in tree architecture (mean branch internode length and branching ratio) that are consistent with clipping. Clipping but not nutrient addition was important for these trees. Clipping frequency was important for non-structural carbohydrate redistribution from roots to stems. A positive feedback could have long-term effects on the palatability of these trees which should cause large mammalian herbivores to consistently favor A. nigrescens.

Seasonal variation in formation, structure, and chemical properties of phloem in Picea abies as studied by novel microtechniques.

Phloem production and structural development were interlinked with seasonal variation in the primary and secondary metabolites of phloem. Novel microtechniques provided new perspectives on understanding phloem structure and chemistry. To gain new insights into phloem formation in Norway spruce (Picea abies), we monitored phloem cell production and seasonal variation in the primary and secondary metabolites of inner bark (non-structural carbohydrates and phenolic stilbene glucosides) during the 2012 growing season in southern and northern Finland. The structure of developing phloem was visualised in 3D by synchrotron X-ray microtomography. The chemical features of developing phloem tissues isolated by laser microdissection were analysed by chemical microanalysis. Within-year phloem formation was associated with seasonal changes in non-structural carbohydrates and phenolic extractive contents of inner bark. The onset of phloem cell production occurred in early and mid-May in southern and northern Finland, respectively. The maximal rate of phloem production and formation of a tangential band of axial phloem parenchyma occurred in mid-June, when total non-structural carbohydrates peaked (due to the high amount of starch). In contrast, soluble sugar content dropped during the most active growth period and increased in late summer and winter. The 3D visualisation showed that the new axial parenchyma clearly enlarged from June to August. Sub-cellular changes appeared to be associated with accumulation of stilbene glucosides and soluble sugars in the newest phloem. Stilbene glucosides also increased in inner bark during late summer and winter. Our findings may indicate that stilbene biosynthesis in older phloem predominantly occurs after the formation of the new band(s) of axial parenchyma. The complementary use of novel microtechniques provides new perspectives on the formation, structure, and chemistry of phloem.

Effects of Summer Pruning and Cropload on Summer and Winter Bud Carbohydrates in Sweet Cherry

Orchard management practices have the potential to influence carbohydrate supply to storage organs, including buds. This study was designed to assess if bud carbohydrates could be manipulated by orchard practices in sweet cherry (Prunus avium). Additionally, we investigated the impact of any such changes on subsequent bud burst and fruit quality the next season. We examined the effect of pruning at different fruit growth stages and cropload on summer and winter bud non-structural carbohydrates (NSCs) and on fruit quality at harvest the next summer in two cultivars. Buds were collected postharvest in summer and after the onset of dormancy in winter, and individual soluble sugars and starch were extracted. The next growing season, flower number and fruit set were recorded. When fruit reached full maturity, fruit were harvested for quality assessment. We observed qualitative changes in NSCs from buds collected in summer and winter. Pruning significantly reduced available NSCs in late summer buds but made little difference by winter; only early pruning showed slight changes in sucrose and glucose. However, early pruning positively influenced the next season’s fruit grade. High cropload resulted in higher NSCs in winter buds than the low cropload. Significant changes in sucrose from summer to winter were observed, and levels of sucrose in the buds differed between cultivars. Different levels of winter bud sucrose between cultivars corresponded to different rates of bud burst. Although pruning was able to manipulate NSCs in buds within a few weeks, these changes were not sufficient to influence the rate of bud burst within a cultivar, but pruning did influence fruit quality in the next season.

Seasonal changes in non-structural carbohydrates and sucrose metabolism enzymes in two Sabina species

Evergreen plants need to store reserves to allow for their survival during the winter months and for new leaf growth in the following spring. In many of the tree species, these reserve functions are mainly carried out by starch, which is degraded to soluble carbohydrates during the dormant season to maintain active respiration and provide protection against freezing. In the present study, two evergreen woody plants, S. przewalskii Kom. (SP) and S. chinensis (Lin.) Ant.(SC), were used to investigate the patterns of seasonal variation in the concentrations of soluble sugars, sucrose, fructose and starch, and the activities of sucrose-phosphate synthase (SPS), sucrose synthase (SUSY), neutral invertase (NI) and soluble acid invertase (SAI). Foliar soluble sugar, sucrose, fructose, and starch concentrations were markedly higher in SC than in SP; moreover, the activities of SPS, SUSY, NI and SAI were also higher in SC than in SP. There were generally higher concentrations of soluble sugars, sucrose and fructose and SPS activities in the winter than in the summer for both Sabina trees, which was consistent with the necessity for protection against freezing; however, opposite results were found with regards to starch concentrations and the activities of SUSY, NI and SAI. In contrast with the activities of SUSY, NI and SAI, the negative correlation observed between SPS and air temperature was likely a reflection of its crucial role in the acquisition of freezing tolerance by sucrose metabolism in the winter. These results suggest that higher carbon reserves do not give S. chinensis (Lin.) Ant. a competitive advantage in tolerating cold temperatures and that only SPS, but not SUSY, NI and SAI, may play a positive role in freezing tolerance by increasing soluble sugar.

Effect of phenology on susceptibility of Norway spruce (Picea abies) to fungal pathogens

Ceratocystis polonica and Heterobasidion parviporum are important fungal pathogens in Norway spruce (Picea abies). Tree susceptibility to these pathogens with respect to phenology was studied using artificial fungal inoculations at six stages of bud development, and assessed by measuring phloem necroses in the stems of 2‐ and 8‐year‐old trees. Tree capacity for resistance was assessed by measuring phloem nonstructural carbohydrates at each stage. Phloem necroses were significantly larger in trees with fungal versus control inoculations and increased significantly over time. Changes in nonstructural carbohydrates occurred in the trees; a significant decline in starch and a slight but significant increase in total sugars occurred over time. These results suggest that susceptibility to fungal pathogens and carbohydrate levels in the stems of the trees were related to fine‐scale changes in bud development. A trade‐off may occur between allocation of starch (the major fraction of the stem carbohydrate pool) to bud development/shoot growth versus defence of the stem. Previous tests of plant defence hypotheses have focused on herbivory on plants growing under different environmental conditions, but the role of phenology and the effect of pathogens are also important to consider in understanding plant resource allocation patterns.

Effect of developmental stage on carbohydrate accumulation patterns during winter of timothy and perennial ryegrass

Changes in non-structural carbohydrates before and during winter are one of the major plant responses to winter stress. However, the observed pattern of changes is variable, not only between grass species but also between different experiments. This study examined the effect of developmental stage on carbohydrate reserves during winter in two grass species differing in winter hardiness. Timothy (Phleum pratense L.) and perennial ryegrass (Lolium perenne L.), sown early or late in the growing season, were sampled for total non-structural carbohydrates (TNC) in late October (end of main growing season), and then in late January, early March and late April. Quantitative and qualitative patterns of carbohydrate accumulation in crown segments were influenced both by plant genotype and developmental stage of the plants prior to winter. TNC increased from October to March, demonstrating considerable cell activity during winter, and declined to their lowest level in April. Sucrose proved to be the major reserve carbohydrate, followed by fructans. The highest content of TNC was found in the most winter-hardy cultivars, particularly in January and March. Early-sown plants accumulated higher levels of fructans than of sucrose, whereas late-sown plants mainly accumulated sucrose. These differences persisted during the winter. It can be concluded that the pattern of carbohydrate accumulation during winter differed between the two species studied and it was strongly affected by both cultivar and plant-developmental stage.

Non-structural carbohydrates in processed soft fried onion ( Allium cepa L.)

The effects of curing, removal of top and root, peeling, storage of raw and peeled onion on the content of non-structural carbohydrates and the ability of onion to develop a brown colour were studied. Glucose, fructose, sucrose, 1-kestose, neokestose, nystose, 1F-β-d-fructofuranosylnystose and four unknown oligofructosides were quantified using high performance anion exchange chromatography coupled with pulsed amperometric detection (HPAEC-PAD). Field drying affected the concentration of fructose only. Glucose, fructose and an unknown fructan decreased in concentration while the concentration of, e.g. 1-kestose, neokestose and nystose increased by removal of root and top of the onions when stored at 15 °C for one week. Peeling had minor influence on changes in non-structural carbohydrates. Storage of non-peeled (raw) onions at 5 °C and storage of mechanically damaged and peeled onion at 20 °C promoted fructan hydrolysis and browning of soft fried onion. Similar changes were produced by long-term storage of cultivars of common onion. Dry matter and fructose increased during storage of peeled onions at both 5 and 20 °C. The increase in fructose content during storage was due to a significantly decrease in the content of 1-kestose, neokestose, 1F-β-d-fructofuranosylnystose, an unknown fructan and total fructans. It was concluded that the missing ability of industrially processed onions to develop a brown colour may be overcome by storage of the peeled onions for a few hours or even overnight at 5 °C.

Adaptation to Winter Stress in Nine White Clover Populations: Changes in Non-structural Carbohydrates During Exposure to Simulated Winter Conditions and ‘Spring’ Regrowth Potential

Nine white clover populations: Grasslands Huia (H0, New Zealand), AberHerald (AH0, bred in the UK from material of Swiss origin) and Sandra (S0, Sweden); sub-populations of these (survival populations) collected after exposure to two (H2 and AH2) and four (H4, AH4 and S4) years of winter stress, respectively, at Uppsala (59°49′N, 17°39′E); and a wild ecotype (WE) from the area were subjected to two simulated winter environments [+2 °C, no light (W1) and +2/−2 °C, 4 h photoperiod (W2) for 3 weeks] after hardening. After hardening, concentrations of total non-structural carbohydrates (TNC) differed little among the populations. However, the changes in these carbohydrates during the winter treatments differed between genetic backgrounds and populations selected for winter survival. Populations S0, S4 and WE showed lower reductions of TNC at low constant temperatures compared with H and AH populations. These populations were also able to maintain the TNC concentration in both stolons and roots despite frequent exposure to sub-lethal frosts, whereas H populations and AH0 and AH2 lost both starch and water soluble carbohydrates from the stolons in treatment W2. The ability to cope with frequent sub-lethal frosts thus appears to play a crucial role in adaptation to northern climates. There appears to have been sufficient genetic variation in AH0 for selection for this trait in the field. The amount of biomass accumulated and the rate of node production during regrowth were generally greatest after treatment W2, when the plants tended to have the highest concentrations of TNC.

Adaptation to Winter Stress in Nine White Clover Populations: Changes in Non-structural Carbohydrates During Exposure to Simulated Winter Conditions and ‘Spring’ Regrowth Potential

Nine white clover populations: Grasslands Huia (H0, New Zealand), AberHerald (AH0, bred in the UK from material of Swiss origin) and Sandra (S0, Sweden); sub-populations of these (survival populations) collected after exposure to two (H2 and AH2) and four (H4, AH4 and S4) years of winter stress, respectively, at Uppsala (59°49'N, 17°39'E); and a wild ecotype (WE) from the area were subjected to two simulated winter environments [+2 °C, no light (W1) and +2/-2 °C, 4 h photoperiod (W2) for 3 weeks] after hardening. After hardening, concentrations of total non-structural carbohydrates (TNC) differed little among the populations. However, the changes in these carbohydrates during the winter treatments differed between genetic backgrounds and populations selected for winter survival. Populations S0, S4 and WE showed lower reductions of TNC at low constant temperatures compared with H and AH populations. These populations were also able to maintain the TNC concentration in both stolons and roots despite frequent exposure to sub-lethal frosts, whereas H populations and AH0 and AH2 lost both starch and water soluble carbohydrates from the stolons in treatment W2. The ability to cope with frequent sub-lethal frosts thus appears to play a crucial role in adaptation to northern climates. There appears to have been sufficient genetic variation in AH0 for selection for this trait in the field. The amount of biomass accumulated and the rate of node production during regrowth were generally greatest after treatment W2, when the plants tended to have the highest concentrations of TNC.

Diurnal changes in non-structural carbohydrates in leaves, phloem exudate and fruit in ‘Braeburn’ apple

Diurnal changes in carbohydrates in leaves, phloem exudate and within fruitwere determined in apple trees grown under high, low or no crop load. Inleaves, diurnal patterns of carbohydrate concentrations were most distinct inhigh-cropping trees and least so in non-cropping trees. In phloem exudate,sorbitol comprised about 65–70% of sugars and sucrose30–35%, with clear diurnal patterns in the amounts. Phloemexudates from stalks of sun and shade fruit were similar. Fruit carbohydratesshowed little diurnal variation. Crude fruit extracts showed little differencein sucrose synthase (SS, EC 2.4.1.13) activity between crop loads, whilesucrose phosphate synthase (SPS, EC 2.4.1.14) activity in high-cropping treespeaked at night and in low-cropping trees in the morning. SPS activity inshade fruit from high-cropping trees was only half of that in sun fruit (atmidday). Results indicate that the greater weight of individual fruit from lowcrop loads is primarily due to greater supply from the source, rather thanhigher metabolic activity in the sink, compared to fruit from high crop loads.In contrast, fruit size differences in sun and shade fruit of high-croppingtrees appear to be due to differences in sink metabolism rather than supplyfrom the source.

Changes in Non-structural Carbohydrates in Tulip Bulb Scales during Cold Treatment and Greenhouse Forcing

The energy and carbon needs for early shoot growth in tulips are mainly provided by reserve carbohydrates in bulbs. The cold-treatment of bulbs before greenhouse forcing enhances the breakdown and remobilization of reserve carbohydrates in bulb scales, and is necessary for proper shoot growth and flowering in tulips. Tulip bulbs are known to contain both starch and fructans as reserve carbohydrates. We evaluated several extraction solvents, including ethanol and distilled water, and several extraction temperatures to accurately determine the amounts of different types of non-structural carbohydrates in tulip bulb scales. Extraction with distilled water resulted in excessive solubilization of starch. For example, extraction at 70 °C solubilized more than 80% of starch to glucan polymers. On the other hand, 80% ethanol at 70 °C extracted all soluble sugars including fructans with no apparent solubilization of starch. The changes in non-structural carbohydrates in the outermost bulb scale of tulip (Tulipa gesneriana L. `Frankfurt') during 12 weeks of cooling at 8.8 °C followed by 5 weeks of greenhouse forcing were determined. Starch was the major carbohydrate in bulb scales consisting of ≈70% of the dry weight at the beginning of cold treatment. Starch content per scale decreased slightly during cold treatment, but rapidly after transferring to greenhouse. Sucrose and soluble fructan content per scale increased during cold treatment, then decreased after transferring to greenhouse. Glucose content per scale remained fairly constant during cooling and greenhouse forcing, while fructose content increased in the greenhouse.