Allocation patterns of nonstructural carbohydrates in response to CO_(2)elevation and nitrogen deposition in Cunninghamia lanceolata saplings

Stored nonstructural carbohydrates(NSC)indicate a balance between photosynthetic carbon(C)assimilation and growth investment or loss through respiration and root exudation.They play an important role in plant function and whole-plant level C cycling.CO_(2)elevation and nitrogen(N)deposition,which are two major environmental issues worldwide,aff ect plant photosynthetic C assimilation and C release in forest ecosystems.However,information regarding the eff ect of CO_(2)elevation and N deposition on NSC storage in diff erent organs remains limited,especially regarding the trade-off between growth and NSC reserves.Therefore,here we analyzed the variations in the NSC storage in diff erent organs of Chinese fi r(Cunninghamia lanceolata)under CO_(2)elevation and N addition and found that NSC concentrations and contents in all organs of Chinese fi r saplings increased remarkably under CO_(2)elevation.However,N addition induced diff erential accumulation of NSC among various organs.Specifi cally,N addition decreased the NSC concentrations of needles,branches,stems,and fi ne roots,but increased the NSC contents of branches and coarse roots.The increase in the NSC contents of roots was more pronounced than that in the NSC content of aboveground organs under CO_(2)elevation.The role of N addition in the increase in the structural biomass of aboveground organs was greater than that in the increase in the structural biomass of roots.This result indicated that a diff erent tradeoff between growth and NSC storage occurred to alleviate resource limitations under CO_(2)elevation and N addition and highlights the importance of separating biomass into structural biomass and NSC reserves when investigating the eff ects of environmental change on biomass allocation.

Polyphosphate Accelerates Transformation of Nonstructural Carbohydrates to Improve Growth of ppk-Expressing Transgenic Rice in Phosphorus Deficiency Culture

Crop yield and quality are often limited by the amount of phosphate fertilizer added to infertile soils,a key limiting factor for sustainable development in modern agriculture.The polyphosphate kinase(ppk)gene-expressing transgenic rice with a single-copy line(ETRS)is constructed to improve phosphate fertilizer utilization efficiency for phosphorus resource conservation.To investigate the potential mechanisms of the increased biomass in ETRS in low phosphate culture,ETRS was cultivated in a low inorganic phosphate(Pi)culture medium(15μmol/L Pi,LP)and a normal Pi culture medium(300μmol/L Pi,CP),respectively.After 89 d of cultivation in different concentrations of phosphate culture media,the total phosphorus,polyphosphate(polyP),biomass,photosynthetic rate,nonstructural carbohydrate(NSC)contents,related enzyme activities,and related gene expression levels were analyzed.The results showed that ETRS had a high polyP amount to promote the photosynthetic rate in LP,and its biomass was almost the same as the wild type(WT)in CP.The NSC content of ETRS in LP was higher than that of WT in LP,but slightly lower than that of WT in CP.PolyP notably promoted the sucrose phosphate synthase activities of ETRS and significantly down-regulated the expression levels of sucrose transporter genes(OsSUT3 and OsSUT4),resulting in inhibiting the transport of sucrose from shoot to root in ETRS.It was concluded that polyP can stimulate the synthesis of NSCs in LP,which improved the growth of ETRS and triggered the biological activities of ETRS to save phosphate fertilizer.Our study provides a new way to improve the utilization rate of phosphate fertilizer in rice production.

Interannual dynamics of stemwood nonstructural carbohydrates in temperate forest trees surrounding drought

Interactions between water and carbon dynamics underlie drought-related tree mortality.While whole-tree water relations have been shown to play a key role in the response to and recovery from drought,the role of nonstructural carbohydrates(NSC) and how their storage and allocation changes surrounding drought events deserves further attention and is critical for understanding tree survival.Here,we quantified in situ NSC responses of temperate forest trees to the 2016 drought in the northeastern United States.Sugar and starch concentrations were measured in the stemwood of five tree species from 2014 to 2019,which allowed us to monitor NSCs in relation to climatic conditions before,during,and after the natural drought.We found that immediately following the drought,measured stemwood NSC concentrations decreased.However,NSC concentrations rebounded quickly within three years.Notably,trees allocated proportionally more to starch than to sugars following the 2016 drought.In winter 2017,starch comprised 45% of total stemwood stores,whereas starch made up only 1-2% in other years.Further,we modeled and assessed the climatic drivers of total NSC concentrations in the stem.Variation in total NSC concentrations was significantly predicted by the previous year’s temperature,precipitation,and standardized precipitation-evapotranspiration index(SPEI),with stemwood concentrations decreasing following hotter,drier periods and increasing following cooler,wetter periods.Overall,our work provides insight into the climatic drivers of NSC storage and highlights the important role that a tree’s carbon economy may play in its response and recovery to environmental stress.

Dynamics of nonstructural carbohydrates in seagrass Thalassia hemprichii and its response to shading

A field survey was performed to examine nonstructural carbohydrate （NSC） dynamics in seagrass Thalassia hemprichii at the Xincun Bay in southern China. An indoor experiment to investigate the response of NSC in T. hemprichiito shade was conducted. Belowground tissue of T. hemprichiiwas the dominant site of NSC reserves, and soluble sugar was the primary storage compound. The starch content of belowground tissue was lower in high intertidal areas than in low intertidal areas, indicating that the longer air exposure in high intertidal areas resulted in less NSC synthesis and less accumulation of NSC in T. hemprichii. The lowest level of soluble sugar and its proportion to NSC in belowground tissue were observed near the cage culture area, where the nutrient concentration in water and sediment was the highest; while the highest level of that was observed near the coastal shrimp farm, where salinity was the lowest. Soluble sugar in belowground tis- sue showed the following trend： summer〉spring〉winter〉autumn. This corresponded to seasonal changes in the intensity of light. Leaf sugar accumulated during the autumn-winter period, providing a carbon and energy source for flower bud formation and seed germination. Short-term shading decreased NSC accumu- lation. Collectively, these results suggest that nutrient enrichment, freshwater discharge and exposure to air affect NSC dynamics in T. hemprichii. Light intensity, flower bud formation, and seed germination were all found to induce seasonal variations in NSC in T. hemprichii.

Divergent allocations of nonstructural carbohydrates shape growth response to rainfall reduction in two subtropical plantations

Nonstructural carbohydrates(NSC)are indicators of tree carbon balance and play an important role in regulating plant growth and survival.However,our understanding of the mechanism underlying drought-induced response of NSC reserves remains limited.Here,we conducted a long-term throughfall exclusion(TFE)experiment to investigate the seasonal responses of NSC reserves to manipulative drought in two contrasting tree species(a broadleaved tree Castanopsis hystrix Miq.and a coniferous tree Pinus massoniana Lamb.)of the subtropical China.We found that in the dry season,the two tree species differed in their responses of NSC reserves to TFE at either the whole-tree level or by organs,with significantly depleted total NSC reserves in roots in both species.Under the TFE treatment,there were significant increases in the NSC pools of leaves and branches in C.hystrix,which were accompanied by significant decreases in fine root biomass and radial growth without significant changes in canopy photosynthesis;while P.massoniana exhibited significant increase in fine root biomass without significant changes in radial growth.Our results suggested that under prolonged water limitation,NSC usage for growth in C.hystrix is somewhat impaired,such that the TFE treatment resulted in NSC accumulation in aboveground organs(leaf and branch);whereas P.massoniana is capable of efficiently utilizing NSC reserves to maintain its growth under drought conditions.Our findings revealed divergent NSC allocations under experimental drought between the two contrasting tree species,which are important for better understanding the differential impacts of climate change on varying forest trees and plantation types in subtropical China.

Effects of pruning intensity on nonstructural carbohydrates of Populus alba×P.talassica in the arid desert region of Northwest China

Although pruning is important to obtain highquality,large-diameter timber,the effects of pruning on nonstructural carbohydrates(NSC)in aboveground organs of many timber species are not well understood.Three intensities of pruning(none,moderate and severe)were tested on poplars(Populus alba×P.talassica)in the arid desert region of northwest China to compare the concentrations of soluble sugar(SS),starch(ST)and total nonstructural carbohydrate(TNC)in leaves,branches and trunks during the growing season.The concentration of NSC components after different pruning intensities varied similarly in seasonal patterns,increasing slowly at the beginning of the growing season,continuously declining in the middle,then gradually recovering by the end of the growing season.The monthly mean NSC concentration in poplar differed significantly among the three pruning intensities(p<0.05).The SS concentration in pruned trees was higher than in unpruned trees(p<0.05).For moderately pruned trees,the concentrations of ST and TNC in trunks and branches were higher than in unpruned and in severely pruned trees(p<0.05).Compared with no pruning,pruning changed the seasonal variation in NSC concentration.The orders of SS and TNC concentrations in aboveground organs were leaf>branch>trunk,while the order of ST concentration was trunk>leaf>branch,which was related to functional differences of plant organs.The annual average growth in height of unpruned,moderately pruned,and severely pruned poplars was 0.21±0.06,0.45±0.09 and 0.24±0.05 m,respectively,and the annual average growth in DBH were 0.92±0.04,1.27±0.06 and 1.02±0.05 cm,respectively.Our results demonstrate that moderate pruning may effectively increase the annual growth in tree height and DBH while avoiding damage caused by excessive pruning to the tree body.Therefore,moderate pruning may increase the NSC storage and improve the growth of timber species.

Effects of AG1 and AG2 QTLs on Nonstructural Carbohydrate and Seed Management Options for Rice Seedling Growth and Establishment under Flooding Stress

Rice(Oryza sativa)plants acquired excess photosynthates in the form of nonstructural carbohydrates(NSCs)in their stems and grain.Despite keen interest in rice NSC,the dynamics of NSC accumulation,translocation and re-accumulation have not yet been well investigated.AG1 and AG2 QTLs associated with flooding tolerance through catalyzing starch into soluble sugar in germinating seeds.Here we conducted three experiments,greenhouse and field to lay the groundwork for large-scale diversity studies on grain NSC and some agronomic traits under direct-seeded rice(DSR)system,using elite lines incorporating AG1,AG2 and AG1-AG2 QTLs into the popular varieties PSB Rc82 and Ciherang-Sub1 along with the donors Kho Hlan On(AG1)and Ma-Zhan Red(AG2).In germinating seedlings,soluble sugars increased,while starch concentration decreased gradually especially in the tolerant checks and AG1-AG2 introgression lines under flooded soil.Soluble sugar accumulation in stem started to increase from the vegetative stage and peaked at the panicle initiation stage then gradually decreased towards the maturity stage.But Sub1-AG lines had higher sugar and starch concentrations at different growth stages than other genotypes in wet season 2016 and dry season 2017.Plant survival rate was positively correlated with the stem NSC at the early vegetative stage(21 days after sowing),and stem NSC was positively associated with plant height at different growth stages.Among the tested seeding rate,the most suitable seeding rate,4 g/m2 with shallow burial depth(0.5 cm),resulted in better seedling establishment,relatively higher seedling vigor index and higher leaf area index under flooding in DSR system.Introgression of AG1-AG2 QTLs had no any negative impact on nonstructural carbohydrate,germination rate,and growth and biomass production.

The storage and utilization of carbohydrates in response to elevation mediated by tree organs in subtropical evergreen broad-leaved forests

Global climate change can affect tree growth and carbon sink function by influencing plant carbohydrate synthesis and utilization,while elevation can be used as an ideal setting under natural conditions to simulate climate change effects.The effect of elevation on tree growth may depend on organ type.However,the allocation patterns of nonstructural and structural carbohydrates(NSCs and SCs,respectively)in different tree organs and their response to elevation remain unclear.We selected four dominant tree species,Schima superba,Castanopsis eyrei,Castanopsis fargesii and Michelia maudiae,along an elevation gradient from 609 to 1,207 m in subtropical evergreen broad-leaved forests and analyzed leaf,trunk,and fine root NSCs,carbon(C),nitrogen(N)and phosphorus(P)concentrations and the relative abundance of SCs.Leaf NSCs increased initially and then decreased,and trunk NSCs increased with increasing elevation.However,root NSCs decreased with increasing elevation.The relative abundance of SCs in leaves and trunks decreased,while the relative abundance of root SCs increased with increasing elevation.No significant correlations between SCs and NSCs in leaves were detected,while there were negative correlations between SCs and NSCs in trunks,roots,and all organs.Hierarchical partitioning analysis indicated that plant C/N and C/P were the main predictors of changes in SCs and NSCs.Our results suggest that tree organs have divergent responses to elevation and that increasing elevation will inhibit the aboveground part growth and enhance the root growth of trees.A tradeoff between the C distribution used for growth and storage was confirmed along the elevation gradient,which is mainly manifested in the"sink"organs of NSCs.Our results provide insight into tree growth in the context of global climate change scenarios in subtropical forest ecosystems.

Multi-year throughfall reduction enhanced the growth and non-structural carbohydrate storage of roots at the expenses of above-ground growth in a warm-temperate natural oak forest

The more frequent occurrence and severer drought events resulting from climate change are increasingly affecting the physiological performance of trees and ecosystem carbon sequestration in many regions of the world.However,our understanding of the mechanisms underlying the responses and adaption of forest trees to prolonged and multi-year drought is still limited.To address this problem,we conducted a long-term manipulative throughfall reduction(TFR,reduction of natural throughfall by 50%–70%during growing seasons)experiment in a natural oriental white oak(Quercus aliena var.acuteserrata Maxim.)forest under warm-temperate climate.After seven years of continuous TFR treatment,the aboveground growth in Q.aliena var.acuteserrata started to decline.Compared with the control plots,trees in the TFR treatment significantly reduced growth increments of stems(14.2%)and leaf area index(6.8%).The rate of net photosynthesis appeared to be more susceptible to changes in soil water in trees subjected to the TFR than in the control.The TFR-treated trees allocated significantly more photosynthates to belowground,leading to enhanced growth and nonstructural carbohydrates(NSC)storage in roots.The 7-year continuous TFR treatment increased the biomass,the production and the NSC concentration in the fine roots by 53.6%,153.6%and 9.6%,respectively.There were clear trade-offs between the aboveground growth and the fine root biomass and NSC storage in Q.aliena var.acuteserrata trees in response to the multi-year TFR treatment.A negative correlation between the fine root NSC concentration and soil water suggested a strategy of preferential C storage over growth when soil water became deficient;the stored NSC during water limitation would then help promote root growth when drought stress is released.Our findings demonstrate the warm-temperate oak forest adopted a more conservative NSC use strategy in response to long-term drought stress,with enhanced root growth and NSC storage at the expenses of above-ground growth to mitigate climate changeinduced drought.

Understory seedlings of Quercus mongolica survive by phenological escape

Understanding understory seedling regeneration mechanisms is important for the sustainable development of temperate primary forests in the context of increasingly intense climate warming events.The poor regeneration of dominant tree species,however,is one of the biggest challenges it faces at the moment.Especially,the regeneration of the shade-intolerant Quercus mongolica seedling is difficult in primary forests,which contrasts with the extreme abundance of understory seedlings in secondary forests.The mechanism behind the interesting phenomenon is still unknown.This study used in-situ monitoring and nursery-controlled experiment to investigate the survival rate,growth performance,as well as nonstructural carbohydrate (NSC) concentrations and pools of various organ tissues of seedlings for two consecutive years,further analyze the understory light availability and simulate the foliage carbon (C) gain in the secondary and primary forest.Results suggested that seedlings in the secondary forest had greater biomass allocation aboveground,height and specific leaf area (SLA) in summer,which allowed the seedling to survive longer in the canopy closure period.High light availability and positive C gain in early spring and late autumn are key factors affecting the growth and survival of understory seedlings in the secondary forest,whereas seedlings in the primary forest had annual negative carbon gain.Through the growing season,the total NSC concentrations of seedlings gradually decreased,whereas those of seedlings in the secondary forest increased significantly in autumn,and were mainly stored in roots for winter consumption and the following year's summer shade period,which was verified by the nursery-controlled experiment that simulated autumn enhanced light availability improved seedling survival rate and NSC pools.In conclusion,our results revealed the survival trade-off strategies of Quercus mongolica seedlings and highlighted the necessity of high light availability during the spring and autumn phenological periods for shade-intolerant tree seedling recruitment.

Foliar carbohydrate differs between Picea crassifolia and Sabina przewalskii with the altitudinal variation of Qilian Mountains, China

Nonstructural carbohydrates （NSC） and nitrogen metabolism strongly influence growth and development in plants. The biosynthesis of cellulose and lignin （structural carbohydrates, SC） depends largely on the supply of NSC. We desire to examine which hypothesis, carbon limitation or growth limitation, best fits the alpine plant response between NSC, SC, carbon （C）, nitrogen （N） and altitude. We compared the foliar concentrations of carbohydrates, C and N between the leaves of Picea crassifolia （lower-elevation tree-line species） and Sabina przewalskii （high-elevation tree-line species） in their response to changing elevation. Our site was located in the mid-northern area of Qilian Mountains, China. We found that foliar soluble sugar （SG） concentrations were significantly higher in P. crassifolia than in S. przewalskii at the 2,700-3,400 m level. Foliar NSC levels in R crassifolia increased at the 2,700-3,100 m level, indicating that growth was limited gradually resulting in a surplus of NSC （to conform to GLH）, subsequently decreasing at the 3,100-3,400 m level, the assimilation declined leading to C deficit （to conform to CLH）. SC （SC metabolism disorders at 3,100-3,400 m）, C, N and starch were significantly lower in R crassifolia than in S. przewalskii. Conversely, foliar SG concentration shows a fall-rise trend with increasing elevation for S. przewalskii. SC concentration in S. przewalskii leaves decreased with an increase of elevation and has a significantly positive correlation to N concentration marking the assimilation of plants. Therefore, the high-elevation tree-line species （S. przewalskii） utilize or store more foliar SG leading to a decrease of SG concentration for survival and growth/regrowth in an adverse environment, higher total C, N, SC, starch contents and lower NSC level. Also, their change trends along the elevational gradient in leaves orS. przewalskii indicate better assimilation strategies for SG use under environmental stress compared to P. crassifolia. This indicates that foliar C metabolism along the elevation follows the principle of the growth-limitation hypothesis （GLH） or carbon limitation hypothesis （CLH）, which depends on the acclimation of different alpine life-forms to the environment.

QTL Analysis for Traits Associated with Feeding Value of Straw in Rice (Oryza sativa L.)

Genetic improvement of the digestibility of rice straw for increasing the utilization of the whole rice plant as feedstuffs is an important way to solve the feedstuffs shortage in southeastern China. To elucidate the genetic basis of the traits affecting the digestibility of rice straw, a rice population of 111 recombinant inbred lines （RILs） derived from a cross of Nekken 2xGaya was used to map the quantitative traits loci （QTLs） for in vitro dry matter digestion （IVDMD）, the percentages of nonstructural carbohydrate （NSC） and acid detergent fiber （exclusive of residual ash, ADFom） in 2005 and 2006. IVDMD was positively correlated with NSC, and negatively correlated with ADFom. A total of 16 QTLs were detected in the two years, and the amounts of variation explained by individual QTLs ranged from 6.9% to 15.5%. Some QTLs for IVDMD and ADFom on chromosome 2 were detected in the two years. On chromosome 2, the interval RM475-RM3515 contained QTLs for IVDMD in the two years and for NSC in 2006 only, while the interval RM3874-RM5305 influenced ADFom in the two years and NSC in 2005 only. At those loci, Gaya contributed favorable alleles to IVDMD and NSC, while Nekken 2 contributed positive alleles to ADFom.

Carbon and nutrient physiology in shrubs at the upper limits:a multispecies study

Aims Carbon and nutrient physiology of trees at their upper limits have been extensively studied,but those of shrubs at their upper limits have received much less attention.The aim of this study is to examine the general patterns of nonstructural carbohydrates(NSCs),nitrogen(N)and phosphorous(P)in shrubs at the upper limits,and to assess whether such patterns are similar to those in trees at the upper limits.Methods Across Eurasia,we measured the concentrations of soluble sugars,starch,total NSCs,N and P in leaves,branches and fine roots(<0.5 cm in diameter)of five shrub species growing at both the upper limits and lower elevations in both summer(peak growing season)and winter(dormancy season).Important Findings Neither elevation nor season had significant effects on tissue N and P concentrations,except for lower P concentrations in fine roots in winter than in summer.Total NSCs and soluble sugars in branches were significantly higher in winter than in summer.There were significant interactive effects between elevation and season for total NSCs,starch,soluble sugars and the ratio of soluble sugar to starch in fine roots,showing lower soluble sugars and starch in fine roots at the upper limits than at the lower elevations in winter but not in summer.These results suggest that the carbon physiology of roots in winter may play an important role in determining the upward distribution of shrubs,like that in the alpine tree-line trees.