Leaf contents differ depending on the position in a rice leaf sheath during sink-source transition.

Carbohydrate contents varied with position in a leaf sheath, and differed between the flag leaf sheath and the second leaf sheath below the flag leaf (-2 leaf sheath) in rice (Oryza sativa L.). In the -2 leaf sheath before heading, light microscopy revealed differences in the distribution of starch granules depending on position. Leaf sheaths were divided into several parts, and the contents of carbohydrates (starch, sucrose, and hexoses) were measured in each part. Before heading, the content of accumulated starch increased linearly from the top to the bottom in -2 leaf sheaths (r2=0.99, P<0.001), as did the contents of accumulated sucrose and hexoses in flag leaf sheaths (r2=0.94, P<0.01). In flag leaf sheaths, the relative content of sucrose synthase (SuS), which plays a central role in the degradation of sucrose into hexoses, increased from the top to the bottom, consistent with hexose contents. After heading, the accumulated carbohydrates were dramatically decreased. In -2 leaf sheaths, the activity of alpha-amylase (EC 3.2.1.1), the rate-limiting step in starch degradation, was consistent with the degree of starch degradation, but in flag leaf sheaths with little starch before heading. These results show that carbohydrate contents differ, depending on the position in a leaf sheath. In addition, there were big differences in leaf contents between flag leaf sheaths and -2 leaf sheaths.     

Changes in Nonstructural Carbohydrates in Different Parts of Soybean (Glycine max [L.] Merr.) Plants during a Light/Dark Cycle and in Extended Darkness

Diurnal patterns of nonstructural carbohydrate (starch, sucrose, and hexose sugars) concentration were characterized in different parts (leaves, petioles, stems, and roots) of vegetative soybean (Glycine max [L.] Merr.) plants. Pronounced changes in all carbohydrate pools were observed in all plant parts during the normal photosynthetic period; however, starch accumulation within leaves accounted for more than 80% of the nonstructural carbohydrate accumulated by the plant during the light period. Efficiency of utilization of starch and sucrose during the normal dark period differed among organs, with leaves being most efficient in mobilizing starch reserves and roots being most efficient in utilizing sucrose reserves. The vast majority (about 85%) of the whole plant carbohydrate reserves present at the end of the photosynthetic period were utilized during the normal dark period. Sink leaf expansion ceased in plants transferred to extended darkness and the cessation in leaf expansion corresponded with carbohydrate depletion in the subtending source leaf and the remainder of the plant. Collectively, the results indicated that under the conditions employed, leaves are the whole plant's primary source of carbon at night as well as during the day.     

猕猴桃果实采后成熟过程中糖代谢及其调节

20℃下采后猕猴桃果实中淀粉酶活性快速上升于果实软化启动阶段,随着果实进入快速软化阶段,淀粉迅速水解,葡萄糖和果糖快速积累,SPS活性增加,酸性转化酶活性下降,蔗糖积累;至果实软化后期,SPS活性降低,蔗糖含量下降.AsA和低温可抑制淀粉酶活性、己糖积累、SPS活性上升和酸性转化酶活性下降,延缓蔗糖积累,相反,乙烯则可促进淀粉酶活性,加速淀粉降解和己糖积累进而直接或间接增加SPS活性,促使蔗糖积累.采后猕猴桃果实的SPS活性变化中有己糖激活效应和蔗糖反馈抑制效应.AsA、低温和乙烯等对糖代谢的调节主要是通过对SPS活性的影响而实现的.     

Analysis of the Relationships between Growth, Photosynthesis and Carbohydrate Metabolism Using Quantitative Trait Loci (QTLs) in Young Maize Plants Subjected to Water Deprivation

One hundred to 120 maize recombinant inbred lines at the mature fourth leaf stage derived from F-2 and Io parental lines were grown in a glasshouse and were deprived of water for 9 days in order to detect pertinent markers of the physiological response to water stress which may be used for breeding. Carbohydrate metabolism QTLs were compared to photosynthesis gas exchange QTLs. The locations of these QTLs were further compared with those of morphological trait QTLs when water availability varied. The traits ranged from three enzyme activities (invertase, sucrose-P synthase, ADP glucose pyrophosphorylase) and hexose, sucrose, starch content to CO₂ uptake and stomatal conductance, water status, leaf size, root/shoot ratio, and ABA (leaf, root and xylem sap). Four main results were obtained (1) only 14 % of QTLs were common to both drought and watered treatments, confirming the existence of stress specific chromosome regions, (2) the QTLs tended to form clusters, frequently consisting of QTLs from different classes (growth, photosynthesis, water status, carbohydrate metabolism and ABA), (3) carbohydrate metabolism trait QTLs were more frequently co-located with growth trait QTLs than photosynthesis related ones, especially in control conditions, (4) one co-location was observed between the three enzyme activities implied in sucrose and starch metabolism and a corresponding structural gene, which can be considered as a candidate gene for explaining part of the variability of each enzymatic trait (invertase, sucrose-P synthase, ADPglucose pyrophosphorylase). It is concluded that, carbohydrate metabolism provides valuable traits for understanding and improving maize responses to water stress.     

Decreased sucrose content triggers starch breakdown and respiration in stored potato tubers (Solanum tuberosum)

To change the hexose-to-sucrose ratio within phloem cells, yeast-derived cytosolic invertase was expressed in transgenic potato (Solanum tuberosum cv. Desirée) plants under control of the rolC promoter. Vascular tissue specific expression of the transgene was verified by histochemical detection of invertase activity in tuber cross-sections. Vegetative growth and tuber yield of transgenic plants was unaltered as compared to wild-type plants. However, the sprout growth of stored tubers was much delayed, indicating impaired phloem-transport of sucrose towards the developing bud. Biochemical analysis of growing tubers revealed that, in contrast to sucrose levels, which rapidly declined in growing invertase-expressing tubers, hexose and starch levels remained unchanged as compared to wild-type controls. During storage, sucrose and starch content declined in wild-type tubers, whereas glucose and fructose levels remained unchanged. A similar response was found in transgenic tubers with the exception that starch degradation was accelerated and fructose levels increased slightly. Furthermore, changes in carbohydrate metabolism were accompanied by an elevated level of phosphorylated intermediates, and a stimulated rate of respiration. Considering that sucrose breakdown was restricted to phloem cells it is concluded that, in response to phloem-associated sucrose depletion or hexose elevation, starch degradation and respiration is triggered in parenchyma cells. To study further whether elevated hexose and/or hexose-phosphates or decreased sucrose levels are responsible for the metabolic changes observed, sucrose content was decreased by tuber-specific expression of a bacterial sucrose isomerase. Sucrose isomerase catalyses the reversible conversion of sucrose into palatinose, which is not further metabolizable by plant cells. Tubers harvested from these plants were found to accumulate high levels of palatinose at the expense of sucrose. In addition, starch content decreased slightly, while hexose levels remained unaltered, compared with the wild-type controls. Similar to low sucrose-containing invertase tubers, respiration and starch breakdown were found to be accelerated during storage in palatinose-accumulating potato tubers. In contrast to invertase transgenics, however, no accumulation of phosphorylated intermediates was observed. Therefore, it is concluded that sucrose depletion rather than increased hexose metabolism triggers reserve mobilization and respiration in stored potato tubers.     

Carbohydrates and carbohydrate enzymes in developing cotton ovules

Patterns of carbohydrates and carbohydrate enzymes were investigated in developing cotton ovules to establish which of these might be related to sink strength in developing bolls. Enzymatic analysis of extracted tissue indicated that beginning 1 week following anthesis, immature cotton seeds (Gossypium hirsutum L. cv. Coker 100A glandless) accumulated starch in the tissues which surround the embryo. Starting at 15 days post anthesis (DPA), this starch was depleted and starch simultaneously appeared in the embryo. Sucrose entering the tissues surrounding the embryo was rapidly degraded, apparently by sucrose synthase; the free hexose content of these tissues reached a peak at about 20 DPA. During the first few weeks of development these tissues contained substantial amounts of hexose but little sucrose; the reverse was true for cotton embryos. Embryo sucrose content rose sharply from the end of the first week until about 20 DPA; it then remained roughly constant during seed maturation. Galactinol synthase (EC 2.4.1.x) appeared in the embryos approximately 25 days after flowering. Subsequently, starch disappeared and the galactosides raffinose and stachyose appeared in the embryo. Except near maturity, sucrose synthase (EC 2.4.1.13) activity in the embryos predominated over that of both sucrose phosphate synthase (EC 2.4.1.14) and acid invertase (EC 3.2.1.26). Activities of the latter enzymes increased during the final stages of embryo maturation. The ratio of sucrose synthase to sucrose phosphate synthase was found to be high in young cotton embryos but the ratio reversed about 45 DPA, when developing ovules cease being assimilate sinks. Insoluble acid invertase was present in developing cotton embryos, but at very low activities; soluble acid invertase was present at significant activities only in nearly mature embryos. From these data it appears that sucrose synthase plays an important role in young cotton ovule carbohydrate partitioning and that sucrose phosphate synthase and the galactoside synthesizing enzymes assume the dominant roles in carbohydrate partitioning in nearly mature cotton seeds. Starch was found to be an important carbohydrate intermediate during the middle stages of cotton ovule development and raffinose and stachyose were found to be important carbohydrate pools in mature cotton seeds.     

Carbohydrate Content and Enzyme Metabolism in Developing Canola Siliques

Little biochemical information is available on carbohydrate metabolism in developing canola (Brassica napus L.) silique (pod) wall and seed tissues. This research examines the carbohydrate contents and sucrose (Suc) metabolic enzyme activities in different aged silique wall and seed tissues during oil filling. The silique wall partitioned photosynthate into Suc over starch and predominantly accumulated hexose. The silique wall hexose content and soluble acid invertase activity rapidly fell as embryos progressed from the early- to late-cotyledon developmental stages. A similar trend was not evident for alkaline invertase, Suc synthase (SuSy), and Suc-phosphate synthase. Silique wall SuSy activities were much higher than source leaves at all times and may serve to supply the substrate for secondary cell wall thickening. In young seeds starch was the predominant accumulated carbohydrate over the sampled developmental range. Seed hexose levels dropped as embryos developed from the early- to midcotyledon stage. Hexose and starch were localized to the testa or liquid endosperm, whereas Suc was evenly distributed among seed components. With the switch to oil accumulation, seed SuSy activity increased by 3.6-fold and soluble acid invertase activity decreased by 76%. These data provide valuable baseline knowledge for the genetic manipulation of canola seed carbon partitioning.     

Soluble acid invertase activity in leaves is independent of species differences in leaf carbohydrates, diurnal sugar profiles and paths of phloem loading

Leaf sucrose, starch, hexose and maximum extractable soluble acid invertase activity were compared throughout the day in source leaves of 13 plant species chosen for their putative phloem-loading type (apoplastic or symplastic). Four species which represent the different phloem-loading types (tomato, barley, maize and Fuchsia ) were studied in detail. Using this information we wished to determine whether a positive correlation between foliar carbohydrates and acid invertase activity exists in leaves from different species and, furthermore, whether this relationship is determined by phloem-loading type. Acid invertase activity was relatively constant throughout the day in all species. The extent of sucrose, hexose and starch accumulation and the sucrose: starch ratio measured at a given time were species-dependent. No correlations were found between foliar soluble acid invertase activity and the hexose, sucrose or starch content of the leaves in any of the species, regardless of phloem-loading type. The species examined could be divided into three distinct groups: (1) high sucrose, low invertase; (2) low sucrose, low invertase; and (3) low sucrose, high invertase. The absence of an inverse relationship between leaf sucrose, hexose or starch contents and endogenous soluble acid invertase suggests that this enzyme is not directly involved in carbon partitioning in leaves but serves an auxiliary function.     

Mechanism associated with nonstructural carbohydrate accumulation in submergence tolerant rice (Oryza sativa L.) cultivars

Nonstructural carbohydrate (NSC) accumulation in submergence tolerant rice cultivars (cv) was studied in six Indica rice [Oryza sativa (L.)] cv under control and simulated submerged conditions. Tolerant cultivars accumulated greater contents of NSC compared to the susceptible cultivars. Starch and total NSC content showed significant positive association with survival percentage. On the other hand, elongation due to submergence was significantly a negative association with survival. The CO₂ photosynthetic rate, chlorophyll content, maximum photochemical efficiency of PS II (Fv/Fm), and activities of Rubisco were not significantly different between tolerant and susceptible cv under control condition. The ADP glucose pyrophosphorylase (AGPPase) activity was significantly higher in the tolerant cv and was a positive association with starch/NSC, whereas Fructose 1,6-diphosphatase (FDPase) activity was significantly higher in susceptible cv compared to tolerant cv and was a negative association with starch/NSC. Greater activities of AGPPase along with lower activities of FDPase might facilitate greater accumulation of NSC in tolerant rice cultivars.     

河套蜜瓜果实发育过程中糖积累与蔗糖代谢相关酶的关系

以河套蜜瓜为试材,采用外部形态观测与内部生理指标测定相结合的方法,对其果实发育过程中果实生长模式以及果实中蔗糖、果糖、葡萄糖和淀粉含量以及蔗糖代谢相关酶活性进行测定,以揭示河套蜜瓜果实生长发育过程中糖的代谢积累与相关酶的关系.结果显示:(1)河套蜜瓜果实生长速率呈单"S"曲线,果实发育早期以积累葡萄糖为主,进入成熟期后蔗糖积累量迅速增加,最终由蔗糖和己糖共同构成果实品质.(2)在河套蜜瓜果实成熟期前,蔗糖磷酸合成酶(SPS)活性维持较低水平,进入成熟期后,SPS活性迅速升高;蔗糖合成酶(SS)活性在成熟期前为分解活性大于合成活性,成熟期后表现为合成活性大于分解活性;在整个果实发育期,酸性转化酶(AI)活性较低,中性转化酶(NI)活性始终高于AI.(3)在果实整个发育期,蔗糖含量与蔗糖代谢酶的净活力呈极显著正相关,蔗糖代谢相关酶共同作用决定果实中蔗糖含量.研究表明,在河套蜜瓜果实发育前期,以蔗糖分解代谢为主,且蔗糖合成酶和中性转化酶是催化蔗糖分解的关键酶;果实成熟期间,蔗糖代谢转为合成方向为主,蔗糖合成酶和蔗糖磷酸合成酶在蔗糖积累中起主导作用.     

开放式空气CO2增高对水稻物质生产与分配的影响

在大田栽培条件下,研究开放式空气CO2增加(FACE)200μmol·mol^-1的处理对水稻物质生产与分配的影响．结果表明,FACE处理使移栽至抽穗后20d的干物质积累量显著增加,使抽穗后20d至成熟期的干物质生产量显著减少,生物产量显著提高．移栽至抽穗期的干物质积累量增加是由于叶面积系数和净同化率共同提高所致;抽穗期至抽穗后20d的干物质积累量增加主要是由于叶面积系数的增加所致;抽穗后20d至成熟期的干物质生产量减少主要是由于净同化率的下降所造成．提高茎鞘占全株干物重的比例,降低叶片占全株干物重的比例,对穗占全株干物重的比例无显著影响,能显著提高水稻抽穗期茎鞘中可溶性糖、淀粉的含有率和含量,提高FACE处理的生物产量能极显著提高水稻产量(r＝0．7825)．     

Identification and physiological analyses of a locus for rice yield potential across the genetic background

A new locus responsible for increased yield potential across the genetic background in rice (Oryza sativa L.) was identified and evaluated. Quantitative trait loci (QTLs) were analysed for the ratio of filled grains, a yield component, in backcrossed inbred lines of a japonica 'Nipponbare'xindica 'Kasalath' cross for 3 years. Only one QTL (rg5), with a positive Kasalath allele, was detected across environments (years). The physiological functions of rg5 were clarified in a near-isogenic line (NILrg5) with a Kasalath chromosome segment containing rg5 in a Nipponbare genetic background. In NILrg5, carbohydrate storage capacity before heading or sink activity in the first or last stages of the reproductive phase was significantly higher than in Nipponbare (control). The ratio of filled grains and yield per plant were significantly higher in NILrg5 than in Nipponbare, by 5% (P <0.01) and 15% (P <0.05), respectively. These results suggest that rg5 improves carbohydrate storage capacity and keeps sink activity higher in the reproductive stage, and consequently increases yield potential. Greater capacity to accumulate carbohydrate is the main target for increasing rice yield potential; therefore, rg5 might function under other genetic backgrounds. Substitution of the rice cv. Kasalath chromosome segment containing rg5 gave higher yield potential in the top premium rice cv. Koshihikari. These results suggest that rg5 might be able to affect yield under different genetic backgrounds, and physiological analyses of the targeted locus might reveal these effects.     

Are contents of Rubisco, soluble protein and nitrogen in flag leaves of rice controlled by the same genetics?

Genetic relations among the contents of Rubisco, soluble protein and total leaf nitrogen (N) in leaves of rice (Oryza sativa L.) were studied by quantitative trait loci (QTL) analysis with a population of backcross inbred lines (BILs) of japonica Nipponbarexindica Kasalath. The ratio of Rubisco to total leaf N in leaves is the main target in improving photosynthetic N-use efficiency in plants. QTLs controlling Rubisco content were not detected near QTLs for total leaf N content. These results indicate that contents of Rubisco and total leaf N are controlled by different genetics. QTLs that controlled the ratio of Rubisco to total leaf N (CORNs) were detected. These results suggest that some mechanism(s) may be involved in determining this ratio, while the contents of Rubisco and total leaf N are controlled in other ways. In elite BILs, the ratios of Rubisco to total leaf N were higher than those of both parents. These results suggest a good possibility of improving N-use efficiency by CORNs in cultivated rice. A QTL controlling Rubisco content was mapped near a QTL for soluble protein content on chromosome 8 at 5 d after heading and on chromosome 9 at 25 d. In each chromosome region, the peaks of both QTLs overlapped accurately, giving a high possibility of pleiotropic effects by the same genes. Different QTLs controlling soluble protein or Rubisco were detected from those detected at 5 d or 25 d after heading. This suggests that these traits are genetically controlled depending on the growth stages of leaves.     

Genetic analysis of sugar-related traits in rice grain

Sugar is the primary product of photosynthesis in plant and plays a critical role in regulating plant growth and development. In this study, quantitative trait loci (QTLs) for total soluble sugar, sucrose, and fructose contents in rice grain were identified using a double haploid population derived from a cross between japonica CJ06 and indica TN1. A total of 17 QTLs, including four QTLs for total soluble sugar content, seven QTLs for sucrose content, and six QTLs for fructose content, were detected on chromosome 1, 3, 4, 5, 6, and 8, with the LOD ranges from 2.61 to 3.85. Furthermore, among the determined varieties, we found that the total soluble sugar content in japonica showed higher than that in indica. Comparative genetic analysis showed that starch synthesis related gene is presumably involved in sugar-related metabolic activity in rice grain.     

Carbon partitioning and sucrose metabolism in tomato plants growing under salinity

The different growth responses under salinity in relation to the carbon partitioning and sucrose metabolism in both sink and source organs have been studied in a salt-tolerant (cv. Pera) and in a salt-sensitive (cv. Volgogradskij) tomato genotype ( Lycopersicon esculentum Mill.). After 3 weeks of salinization, the plant dry weight was reduced by 12–34% in cv. Pera and by 45–58% in cv. Volgogradskij. Photosynthesis was positively correlated to plant growth in cv. Pera but not in cv. Volgogradskij. In this salt-sensitive genotype, both photosynthesis and growth were negatively correlated with fructose, glucose and sucrose accumulation in both mature and young leaves, suggesting a blockage in their use for growth. The transient accumulation of sucrose and hexoses in the young leaves of cv. Pera was linked to increases in all soluble sucrolytic activities, mainly acid invertase (EC 3.2.1.25) and sucrose synthase (EC 2.4.1.13), which was related to sink activity and growth capacity. The sucrose-phosphate synthase activity (EC 2.4.1.14) was related to the ability of mature leaves to regulate assimilate production, accumulation and export. The salt-tolerant cv. Pera accumulated a higher amount of total carbohydrates, but cv. Volgogradskij showed the highest soluble fraction under salinity. The carbohydrate availability and the photosynthetic rate do not seem to be the first limiting factors for plant growth under saline conditions, but the different behavior observed in both genotypes concerning the distribution and use of photoassimilates could help to explain their different salt-tolerance degrees.     

The base of the leaf acts as a localized sink for photosynthate in mature barley leaves

The gradients in photosynthetic and carbohydrate metabolism which persist within the fully expanded second leaf of barley ( Hordeum vulgare ) were examined. Although all regions of the leaf blade were green and photosynthetically active, the basal 5 cm, representing approximately 20% of the leaf area, retained some characteristics of sink tissue. The leaf blade distal from the leaf sheath exhibited characteristics typical of source tissue; the activities of sucrolytic enzymes (invertase and sucrose synthase) were relatively low, whilst that of sucrose phosphate synthase was high. These regions of the leaf accumulated sucrose throughout the photoperiod and starch only in the second half of the photoperiod whilst hexose sugars remained low. By contrast the leaf blade proximal to the leaf sheath retained relatively high activities of sucrolytic enzymes (especially soluble, acid invertase) whilst sucrose phosphate synthase activity was low. Glucose, as well as sucrose, accumulated throughout the photoperiod. Although starch accumulated in the second half of the photoperiod, a basal level of starch was present throughout the photoperiod, by contrast with the rest of the leaf. The ¹⁴CO₂ feeding experiments indicated that a constant amount of photosynthate was partitioned towards starch in this region of the leaf irrespective of irradiance. These findings are interpreted as the base of the leaf blade acting as a localized sink for carbohydrate as a result of sucrose hydrolysis by acid invertase.     

Development of Desiccation Tolerance during Embryogenesis in Rice (Oryza sativa) and Wild Rice (Zizania palustris) (Dehydrin Expression,Abscisic Acid Content,and Sucrose Accumulation)

The ability of seeds to withstand desiccation develops during embryogenesis and differs considerably among species. Paddy rice (Oryza sativa L.) grains readily survive dehydration to as low as 2% water content, whereas North American wild rice (Zizania palustris var interior [Fasset] Dore) grains are not tolerant of water contents below 6% and are sensitive to drying and imbibition conditions. During embryogenesis, dehydrin proteins, abscisic acid (ABA), and saccharides are synthesized, and all have been implicated in the development of desiccation tolerance. We examined the accumulation patterns of dehydrin protein, ABA, and soluble saccharides (sucrose and oligosaccharides) of rice embryos and wild rice axes in relation to the development of desiccation tolerance during embryogenesis. Dehydrin protein was detected immunologically with an antibody raised against a conserved dehydrin amino acid sequence. Both rice and wild rice embryos accumulated a 21-kD dehydrin protein during development, and an immunologically related 38-kD protein accumulated similarly in rice. Dehydrin protein synthesis was detected before desiccation tolerance had developed in both rice embryos and wild rice axes. However, the major accumulation of dehydrin occurred after most seeds of both species had become desiccation tolerant. ABA accumulated in wild rice axes to about twice the amount present in rice embryos. There were no obvious relationships between ABA and the temporal expression patterns of dehydrin protein in either rice or wild rice. Wild rice axes accumulated about twice as much sucrose as rice embryos. Oligosaccharides were present at only about one-tenth of the maximum sucrose concentrations in both rice and wild rice. We conclude that the desiccation sensitivity displayed by wild rice grains is not due to an inability to synthesize dehydrin proteins, ABA, or soluble carbohydrates.