Involvement of abscisic acid and cytokinins in the senescence and remobilization of carbon reserves in wheat subjected to water stress during grain filling

This study investigated the possibility that abscisic acid (ABA) and cytokinins may mediate the effect of water deficit that enhances plant senescence and remobilization of pre‐stored carbon reserves. Two high lodging‐resistant wheat (Triticum aestivum L.) cultivars were field grown and treated with either a normal or high amount of nitrogen at heading. Well‐watered (WW) and water‐stressed (WS) treatments were imposed from 9 d post‐anthesis until maturity. Chlorophyll (Chl) and photosynthetic rate (Pr) of the flag leaves declined faster in WS plants than in WW plants, indicating that the water deficit enhanced senescence. Water stress facilitated the reduction of non‐structural carbohydrate in the stems and promoted the re‐allocation of prefixed ¹⁴C from the stems to grains, shortened the grain filling period and increased the grain filling rate. Water stress substantially increased ABA but reduced zeatin (Z) + zeatin riboside (ZR) concentrations in the stems and leaves. ABA correlated significantly and negatively, whereas Z + ZR correlated positively, with Pr and Chl of the flag leaves. ABA but not Z + ZR, was positively and significantly correlated with remobilization of pre‐stored carbon and grain filling rate. Exogenous ABA reduced Chl in the flag leaves, enhanced the remobilization, and increased grain filling rate. Spraying with kinetin had the opposite effect. The results suggest that both ABA and cytokinins are involved in controlling plant senescence, and an enhanced carbon remobilization and accelerated grain filling rate are attributed to an elevated ABA level in wheat plants when subjected to water stress.     

氮素形态对小麦花后不同器官内源激素含量的影响

采用盆栽方法,研究了酰胺态氮、铵态氮和硝态氮对小麦(Triticum aestivum)花后根系、旗叶和籽粒内源激素IAA、GA₃、ABA和ZR含量的影响。结果表明,小麦不同器官的内源激素含量对3种氮素形态的响应不同。氮素形态调节籽粒灌浆是通过根系、旗叶和籽粒中内源激素的协同作用而实现的。酰胺态氮与硝态氮处理相比,小麦花后5~15 d,旗叶GA₃含量、籽粒IAA和ABA含量较高,籽粒灌浆速率(Grain-filling rate, GFR)较高;花后15~25 d,根系GA₃含量、旗叶IAA和GA₃含量、籽粒ABA含量较高,籽粒IAA含量较低,GFR较低。铵态氮与硝态氮处理相比,小麦花后5 d,籽粒ZR含量较高;花后15 d前后,籽粒IAA、ABA含量较低,GFR较低;花后20~25 d,根系ZR、GA₃含量较低,旗叶IAA、GA₃含量较低,ABA含量较高,籽粒ABA、GA₃含量较低,IAA含量较高,GFR较高。铵态氮比硝态氮处理的小麦籽粒粒重显著增加。铵态氮和酰胺态氮处理比硝态氮处理增产显著。建议在‘豫麦49’施肥时,使用铵态氮或酰胺态氮并配施硝化抑制剂。     

Abscisic acid and cytokinins in the root exudates and leaves and their relationship to senescence and remobilization of carbon reserves in rice subjected to water stress during grain filling

The possible regulation of senescence-initiated remobilization of carbon reserves in rice (Oryza sativa L.) by abscisic acid (ABA) and cytokinins was studied using two rice cultivars with high lodging resistance and slow remobilization. The plants were grown in pots and either well-watered (WW, soil water potential = 0 MPa) or water-stressed (WS, soil water potential = -0.05 MPa) from 9 days after anthesis until they reached maturity. Leaf water potentials of both cultivars markedly decreased at midday as a result of water stress but completely recovered by early morning. Chlorophyll (Chl) and photosynthetic rate (Pr) of the flag leaves declined faster in WS plants than in WW plants, indicating that the water deficit enhanced senescence. Water stress accelerated starch remobilization in the stems, promoted the re-allocation of pre-fixed (14)C from the stems to grains, shortened the grain-filling period and increased the grain-filling rate. Sucrose phosphate synthase (SPS, EC 2.4.1.14) activity was enhanced by water stress and positively correlated with sucrose accumulation in both the stem and leaves. Water stress substantially increased ABA but reduced zeatin (Z) + zeatin riboside (ZR) concentrations in the root exudates and leaves. ABA significantly and negatively, while Z+ZR positively, correlated with Pr and Chl of the flag leaves. ABA, not Z+ZR, was positively and significantly correlated with SPS activity and remobilization of pre-stored carbon. Spraying ABA reduced Chl in the flag leaves, and enhanced SPS activity and remobilization of carbon reserves. Spraying kinetin had the opposite effect. The results suggest that both ABA and cytokinins are involved in controlling plant senescence, and an enhanced carbon remobilization is attributed to an elevated ABA level in rice plants subjected to water stress.     

Mapping of QTL associated with nitrogen storage and remobilization in barley (Hordeum vulgare L.) leaves

Nitrogen uptake and metabolism are central for vegetative and reproductive plant growth. This is reflected by the fact that nitrogen can be remobilized and reused within a plant, and this process is crucial for yield in most annual crops. A population of 146 recombinant inbred barley lines (F(8) and F(9) plants, grown in 2000 and 2001), derived from a cross between two varieties differing markedly in grain protein concentration, was used to compare the location of QTL associated with nitrogen uptake, storage and remobilization in flag leaves relative to QTL controlling developmental parameters and grain protein accumulation. Overlaps of support intervals for such QTL were found on several chromosomes, with chromosomes 3 and 6 being especially important. For QTL on these chromosomes, alleles associated with inefficient N remobilization were associated with depressed yield and higher levels of total or soluble organic nitrogen during grain filling and vice versa; therefore, genes directly involved in N recycling or genes regulating N recycling may be located on these chromosomes. Interestingly, the most prominent QTL for grain protein concentration (on chromosome 6) did not co-localize with QTL for nitrogen remobilization. However, QTL peaks for nitrate and soluble organic nitrogen were detected at this locus for plants grown in 2001 (but not in 2000). For these, alleles associated with low grain protein concentration were associated with higher soluble nitrogen levels in leaves during grain filling; therefore, gene(s) found at this locus might influence the nitrogen sink strength of developing barley grains.     

Effect of abscisic acid on the transport of assimilates in barley

The effect of abscisic acid (ABA) on assimilate transport in barley was investigated in two parallel experiments. First, the effect upon [¹⁴C]sucrose transport from the flag leaf to the ear of a single ABA application made at different stages of growth of the fruits was investigated; the effect was measured 24 h after treatment. Second, the effect of a single application of ABA made at the same stages of growth as above on grain weight of the mature plant was investigated. In both types of experiments ABA was applied once to the ear of different plants as an aqueous solution (10^-3–10^-5 M), one to five weeks after anthesis. [¹⁴C] sucrose was applied by means of agar blocks. Parallel to these experiments, the endogenous content of ABA was investigated in the developing grains. When ears were treated with ABA two or four weeks after anthesis, an increase of up to 70% in the ¹⁴C-transport from the flag leaf to the ear was observed within a 24-h period after treatment (short duration experiments). At these growth stages the endogenous concentrations of ABA were low. In sharp contrast, ABA, especially in a concentration of 10^-3 M, decreased ¹⁴C-import from the flag leaf when applied three weeks after anthesis. At this stage the endogenous ABA content had reached its maximum. Long duration experiments with a single application of ABA to the car two weeks after anthesis resulted in a marked increase of weight per thousand kernels. ABA applications made earlier or later than two weeks after anthesis either reduced the grain weight or had no effect. It is concluded that ABA is involved in the regulation of assimilate transport from the leaves to the grains, possibly by influencing the unloading of sieve tubes in the ears. Promotion or inhibition of assimilate import by exogenously applied ABA may depend on the developmental stage of the grains and on the endogenous ABA level.Abbreviations ABA abscisic acid - TKW weight per thousand kernels     

Hormonal changes in the grains of rice subjected to water stress during grain filling

Lodging-resistant rice (Oryza sativa) cultivars usually show slow grain filling when nitrogen is applied in large amounts. This study investigated the possibility that a hormonal change may mediate the effect of water deficit that enhances whole plant senescence and speeds up grain filling. Two rice cultivars showing high lodging resistance and slow grain filling were field grown and applied with either normal or high amount nitrogen (HN) at heading. Well-watered and water-stressed (WS) treatments were imposed 9 days post anthesis to maturity. Results showed that WS increased partitioning of fixed (14)CO(2) into grains, accelerated the grain filling rate but shortened the grain filling period, whereas the HN did the opposite way. Cytokinin (zeatin + zeatin riboside) and indole-3-acetic acid contents in the grains transiently increased at early filling stage and WS treatments hastened their declines at the late grain filling stage. Gibberellins (GAs; GA(1) + GA(4)) in the grains were also high at early grain filling but HN enhanced, whereas WS substantially reduced, its accumulation. Opposite to GAs, abscisic acid (ABA) in the grains was low at early grain filling but WS remarkably enhanced its accumulation. The peak values of ABA were significantly correlated with the maximum grain filling rates (r = 0.92**, P < 0.01) and the partitioning of fixed (14)C into grains (r = 0.95**, P < 0.01). Exogenously applied ABA on pot-grown HN rice showed similar results as those by WS. Results suggest that an altered hormonal balance in rice grains by water stress during grain filling, especially a decrease in GAs and an increase in ABA, enhances the remobilization of prestored carbon to the grains and accelerates the grain filling rate.     

限水灌溉冬小麦冠层氮分布与转运特征及其对供氮的响应

以高产冬小麦品种周麦18为材料,在大田春灌1水条件下,设置不同供氮水平和氮肥运筹处理试验,研究并探讨了在华北地区限水灌溉条件下氮肥施用对冬小麦冠层叶片氮素时空分布与转运及氮肥利用的影响。结果表明,冬小麦适量施氮可显著增产,2008-2009年以施氮量180 kg/hm²时(N21)产量最高,为8749 kg/hm²;2009-2010年以施氮量270 kg/hm²时(N32)产量最高,但施氮量210 kg/hm²(N22)处理与N32处理产量无显著差异,分别为8340 kg/hm²和8558 kg/hm²。氮肥利用效率和氮肥偏生产力均随施氮量增加而降低;氮肥利用率与氮肥农学效率均随施氮量的增加呈先升后降的趋势,分别在N21和N22处理时最高。冠层叶片氮素含量和积累量随叶层层次自上而下降低而下降,垂直梯度分明,各时期冠层叶片氮素垂直梯度随施氮量的增加总体呈先增大后减小的趋势。冠层叶片氮素转运量、转运率和对籽粒的贡献率均呈现为:第1层>第2层>第3层>第4层。相关分析表明,冠层叶片氮素梯度与叶片氮素转运率呈显著正相关关系(R²=0.722^*),与贡献率呈极显著正相关关系(R²=0.975^**)。适量施氮(120-210 kg/hm²)增大了叶层间氮素垂直分布梯度,促进了氮素在植株内的运移分配,有利于叶片氮素向外转运,提高了叶片氮素转运量和对籽粒贡献率,保持了较高的氮素利用率。施氮过多(330 kg/hm²)减小了叶层间氮素垂直分布梯度,减弱了氮素在植株内的再利用,叶片氮素转运不畅,导致叶片氮素转运量和对籽粒贡献率下降,氮素利用率显著降低。连续两年试验结果显示,通过适量氮肥调控可以增大冠层叶片氮素垂直梯度,有利于叶片中的氮素输出,促进氮素的再分配、再利用,从而提高氮素利用率,并可获得较高的籽粒产量和蛋白质含量。     

Relationships of endogenous plant hormones to accumulation of grain protein and starch in winter wheat under different post-anthesis soil water statusses

Accumulation of protein and starch in grain is a key process determining grain yield and quality in wheat. Under drought or waterlogging, endogenous plant hormone levels will change and may have an impact on the yield and quality of wheat. In a greenhouse experiment, four winter wheat (Triticum aestivum L.) varieties differing in grain protein content, Heimai 76, Wanmai 38, Yangmai 10 and Yangmai 9, were subjected to drought (SRWC = 4550%, DR), waterlogging (WL) and moderate water supply (SRWC = 7580%, CK), beginning from 4 days post-anthesis (DPA) to maturity. On the 10 (grain enlargement stage) and 20 (grain filling stage) DPA, endogenous abscisic acid (ABA), gibberellins (GA₁₊₃), indole-3-acetic acid (IAA) and zeatin riboside (ZR) were determined in sink and source organs of wheat plants by enzyme linked immunosorbent assay (ELISA). The patterns of hormonal changes were similar in four varieties. The ABA levels were much higher under DR and WL than under CK. Compared with CK, GA₁₊₃ levels in whole-plant under DR and WL changed a little at 10 DPA, but markedly decreased under DR and WL at 20 DPA. Changes of endogenous IAA level under DR and WL exhibited a complicated pattern, depending on organs and growth stages. Particularly at the 20 DPA, the mean levels of IAA in roots, leaves and grains decreased significantly under DR and WL. In comparison with CK, ZR levels in all organs significantly decreased under DR and WL at both stages. The correlation analyses between yields and contents of starch and protein in grains and levels and ratios of four hormones in source and sink organs indicated that the changes in yield and content of grain starch and protein under DR and WL were associated with the reduced IAA, ZR and GA₁₊₃ levels and elevated ABA level in plants, especially in grains. It was proposed that the changed levels of endogenous hormones under post-anthesis DR and WL might indirectly affect protein and starch accumulation in grains by influencing the regulatory enzymes and processes.     

Activities of fructan- and sucrose-metabolizing enzymes in wheat stems subjected to water stress during grain filling

This study investigated if a controlled water deficit during grain filling of wheat (Triticum aestivum L.) could accelerate grain filling by facilitating the remobilization of carbon reserves in the stem through regulating the enzymes involved in fructan and sucrose metabolism. Two high lodging-resistant wheat cultivars were grown in pots and treated with either a normal (NN) or high amount of nitrogen (HN) at heading time. Plants were either well-watered (WW) or water-stressed (WS) from 9 days post anthesis until maturity. Leaf water potentials markedly decreased at midday as a result of water stress but completely recovered by early morning. Photosynthetic rate and zeatin + zeatin riboside concentrations in the flag leaves declined faster in WS plants than in WW plants, and they decreased more slowly with HN than with NN when soil water potential was the same, indicating that the water deficit enhanced, whereas HN delayed, senescence. Water stress, both at NN and HN, facilitated the reduction in concentration of total nonstructural carbohydrates (NSC) and fructans in the stems but increased the sucrose level there, promoted the re-allocation of pre-fixed ¹⁴C from the stems to grains, shortened the grain-filling period, and accelerated the grain-filling rate. Grain weight and grain yield were increased under the controlled water deficit when HN was applied. Fructan exohydrolase (FEH; EC 3.2.1.80) and sucrose phosphate synthase (SPS; EC 2.4.1.14) activities were substantially enhanced by water stress and positively correlated with the total NSC and fructan remobilization from the stems. Acid invertase (EC 3.2.1.26) activity was also enhanced by the water stress and associated with the change in fructan concentration, but not correlated with the total NSC remobilization and ¹⁴C increase in the grains. Sucrose:sucrose fructosyltransferase (EC 2.4.1.99) activity was inhibited by the water stress and negatively correlated with the remobilization of carbon reserves. Sucrose synthase (EC 2.4.1.13) activity in the stems decreased sharply during grain filling and showed no significant difference between WW and WS treatments. Abscisic acid (ABA) concentration in the stem was remarkably enhanced by water stress and significantly correlated with SPS and FEH activities. Application of ABA to WW plants yielded similar results to those for WS plants. The results suggest that the increased remobilization of carbon reserves by water stress is attributable to the enhanced FEH and SPS activities in wheat stems, and that ABA plays a vital role in the regulation of the key enzymes involved in fructan and sucrose metabolism.Abbreviations ABA Abscisic acid - DAS Days after sowing - DPA Days post anthesis - ESC Ethanol-soluble carbohydrate - FEH Fructan exohydrolase - HN High amount of nitrogen - INV Invertase - NN Normal amount of nitrogen - NSC Nonstructural carbohydrate - _leaf Leaf water potential - _soil Soil water potential - Pr Photosynthetic rate - SPS Sucrose phosphate synthase - SS Sucrose synthase - SST Sucrose:sucrose fructosyltransferase - V_limit Limiting substrate - V_max Saturated substrate - WS Water stressed - WSC Water-soluble carbohydrate - WW Well watered - Z Zeatin - ZR Zeatin riboside     

Relationship of Nitrogen Deficiency-Induced Leaf Senescence with ROS Generation and ABA Concentration in Rice Flag Leaves

Nitrogen (N) deficiency is one of the critical environmental factors that induce leaf senescence, and its occurrence may cause the shorten leaf photosynthetic period and markedly lowered grain yield. However, the physiological metabolism underlying N deficiency-induced leaf senescence and its relationship with the abscisic acid (ABA) concentration and reactive oxygen species (ROS) burst in leaf tissues are not well understood. In this paper, the effect of N supply on several senescence-related physiological parameters and its relation to the temporal patterns of ABA concentration and ROS accumulation during leaf senescence were investigated using the premature senescence of flag leaf mutant rice (psf) and its wild type under three N treatments. The results showed that N deficiency hastened the initiation and progression of leaf senescence, and this occurrence was closely associated with the upregulated expression of 9-cis-epoxycarotenoiddioxygenase genes (NCEDs) and with the downregulated expression of two ABA 8′-hydroxylase isoform genes (ABA8ox2 and ABA8ox3) under LN treatment. Contrarily, HN supply delayed the initiation and progression of leaf senescence, concurrently with the suppressed ABA biosynthesis and relatively lower level of ABA concentration in leaf tissues. Exogenous ABA incubation enhanced ROS generation and MDA accumulation in a dose-dependent manner, but it decreased the activities of glutamine synthetase (GS) and glutamate dehydrogenase (GDH) in detached leaf. These results suggested that the participation of ABA in the regulation of ROS generation and N assimilating/remobilizing metabolism in rice leaves was strongly responsible for induction of leaf senescence by N deficiency.

     

Regulation of Nitrogen Metabolism,Photosynthetic Activity,and Yield Attributes of Spring Wheat by Nitrogen Fertilizer in the Semi-arid Loess Plateau Region

It is critical for spring wheat (Triticum aestivum L.) production in the semi-arid Loess Plateau to understand the impact of nitrogen (N) fertilizer on changes in N metabolism, photosynthetic parameters, and their relationship with grain yield and quality. The photosynthetic capacity of flag leaves, dry matter accumulation, and N metabolite enzyme activities from anthesis to maturity were studied on a long-term fertilization trial under different N rates [0 kg ha^?1(N1), 52.5 kg ha^?1 (N2), 105 kg ha^?1 (N3), 157.5 kg ha^?1 (N4), and 210 kg ha^?1 (N5)]. It was observed that N3 produced optimum total dry matter (5407 kg ha^?1), 1000 grain weight (39.7 g), grain yield (2.64 t ha^?1), and protein content (13.97%). Our results showed that N fertilization significantly increased photosynthetic parameters and N metabolite enzymes at all growth stages. Nitrogen harvest index, partial productivity factor, agronomic recovery efficiency, and nitrogen agronomic efficiency were decreased with increased N. Higher N rates (N3–N5) maintained higher photosynthetic capacity and dry matter accumulation and lower intercellular CO₂ content. The N supply influenced NUE by improving photosynthetic properties. The N3 produced highest chlorophyll content, photosynthetic rate, stomatal conductance and transpiration rate, grain yield, grain protein, dry matter, grains weight, and N metabolite enzyme activities compared to the other rates (N1, N2, N4, and N5). Therefore, increasing N rates beyond the optimum quantity only promotes vegetative development and results in lower yields.

     

施氮量和花后土壤含水量对小麦旗叶衰老及粒重的影响

在防雨池栽培条件下,研究了施氮量和花后土壤含水量对小麦旗叶衰老和粒重的影响.结果表明：各氮肥处理下,小麦旗叶SPAD值、可溶性蛋白质含量、超氧化物歧化酶（SOD）活性、过氧化氢酶（CAT）活性和光合速率（P_n）均表现为：花后土壤含水量60%～70%处理>80%～90%处理>40%～50%处理,小麦旗叶丙二醛（MDA）含量表现为：花后土壤含水量40%～50%处理>80%～90%处理>60%～70%处理,表明花后土壤含水量过高或过低均可导致小麦旗叶早衰,影响籽粒灌浆,降低粒重.在花后相同土壤含水量条件下,旗叶SPAD值、可溶性蛋白质含量、SOD活性、CAT活性和P_n均随施氮量的增加而升高,MDA含量随施氮量的增加而降低,表明增施氮肥可以延缓小麦旗叶衰老,但过量施用氮肥则不利于小麦粒重的提高,尤其是在花后土壤缺水的情况下,施用过多氮肥可导致小麦粒重下降.在小麦生产中可以将施用氮肥和控制花后土壤水分含量相结合,延缓小麦植株衰老,提高粒重.     

施氮量对花铃期短期渍水棉花叶片抗氧化酶活性和内源激素含量的影响

于2005-2006年在江苏南京（32°02′ N,118°50′ E）南京农业大学卫岗试验站进行盆栽试验,设置正常灌水和棉花花铃期短期渍水处理（渍水8 d,恢复15 d）,每个水分处理设置3个施氮水平（0、240、480 kg N·hm^-2）,研究施氮量对渍水棉花叶片抗氧化酶活性和内源激素含量的影响.结果表明：在渍水结束时,与正常灌水相比,渍水棉花叶片可溶性蛋白含量、超氧化物歧化酶和过氧化氢酶活性降低,过氧化物酶活性升高,丙二醛（MDA）含量升高;与此同时,内源激素含量发生变化,脱落酸（ABA）含量升高,玉米素核苷（ZR）、赤霉素（GA）、生长素（IAA）含量及ZR/ABA、GA/ABA、IAA/ABA降低;其中以240 kg N·hm^-2渍水棉花MDA和ABA含量最低,而ZR、GA和IAA含量及ZR/ABA、GA/ABA和IAA/ABA最高,净光合速率最高.到停止渍水15 d时,渍水棉花叶片的抗氧化酶活性、MDA含量、内源激素含量与正常灌水处理的差异较小;施氮可提高渍水处理棉花叶片抗氧化酶活性,降低MDA含量,使ABA水平降低,ZR、GA、IAA水平及ZR/ABA、GA/ABA、IAA/ABA升高.本试验中,渍水条件下以240 kg N·hm^-2处理的效果最好,生物量和籽棉产量最高.     

NEMA, a functional-structural model of nitrogen economy within wheat culms after flowering. II. Evaluation and sensitivity analysis

Background and Aims

Simulating nitrogen economy in crop plants requires formalizing the interactions between soil nitrogen availability, root nitrogen acquisition, distribution between vegetative organs and remobilization towards grains. This study evaluates and analyses the functional–structural and mechanistic model of nitrogen economy, NEMA (Nitrogen Economy Model within plant Architecture), developed for winter wheat (Triticum aestivum) after flowering.

Methods

NEMA was calibrated for field plants under three nitrogen fertilization treatments at flowering. Model behaviour was investigated and sensitivity to parameter values was analysed.

Key Results

Nitrogen content of all photosynthetic organs and in particular nitrogen vertical distribution along the stem and remobilization patterns in response to fertilization were simulated accurately by the model, from Rubisco turnover modulated by light intercepted by the organ and a mobile nitrogen pool. This pool proved to be a reliable indicator of plant nitrogen status, allowing efficient regulation of nitrogen acquisition by roots, remobilization from vegetative organs and accumulation in grains in response to nitrogen treatments. In our simulations, root capacity to import carbon, rather than carbon availability, limited nitrogen acquisition and ultimately nitrogen accumulation in grains, while Rubisco turnover intensity mostly affected dry matter accumulation in grains.

Conclusions

NEMA enabled interpretation of several key patterns usually observed in field conditions and the identification of plausible processes limiting for grain yield, protein content and root nitrogen acquisition that could be targets for plant breeding; however, further understanding requires more mechanistic formalization of carbon metabolism. Its strong physiological basis and its realistic behaviour support its use to gain insights into nitrogen economy after flowering.     

Nitrogen assimilation in opaque and normal sorghum during grain development

Activity of key nitrogen assimilating enzymes was studied in developing grains of high-lysine opaque sorghum P-721 and normal sorghum CSV-5. The higher percentage of protein in opaque sorghum was mainly due to lower starch content since protein per grain was less than in CSV-5. During grain development, albufn and globulin decreased while prolafne and glutelin increased. Prolafne content in CSV-5 was higher than in opaque sorghum. Average nitrate reductase activity in flag and long leaf were similar in both the varieties. The nitrate reductase activity decreased during grain development. Glutamate dehydrogenase activity was higher during early development and lower at later stages in opaque sorghum than in CSV-5. Glutamate oxaloacetate transaminase activity was higher and glutamine synthetase lower in opaque sorghum than in CSV-5 grains during development. Glutamate synthase activity was higher in opaque sorghum up to day 20 and lower thereafter than in CSV-5. It is suggested that reduced activities of glutamine synthetase as well as glutamate synthase in opaque sorghum as compared to CSV-5 during later stages of development may restrict protein accumulation in the former.     

Effect of Subsoiling in Fallow Period on Soil Water Storage and Grain Protein Accumulation of Dryland Wheat and Its Regulatory Effect by Nitrogen Application

To provide a new way to increase water storage and retention of dryland wheat, a field study was conducted at Wenxi experimental site of Shanxi Agricultural University. The effect of subsoiling in fallow period on soil water storage, accumulation of proline, and formation of grain protein after anthesis were determined. Our results showed that subsoiling in fallow period could increase water storage in the 0–300 cm soil at pre-sowing stage and at anthesis stage with low or medium N application, especially for the 60–160 cm soil. However, the proline content, glutamine synthetase (GS) activity, glutamate dehydrogenase (GDH) activity in flag leaves and grains were all decreased by subsoiling in fallow period. In addition, the content of albumin, gliadin, and total protein in grains were also decreased while globulin content, Glu/Gli, protein yield, and glutelin content were increased. With N application increasing, water storage of soil layers from 20 to 200 cm was decreased at anthesis stage. High N application resulted in the increment of proline content and GS activity in grains. Besides, correlation analysis showed that soil storage in 40–160 cm soil was negatively correlated with proline content in grains; proline content in grains was positively correlated with GS and GDH activity in flag leaves. Contents of albumin, globulin and total protein in grains were positively correlated with proline content in grains and GDH activity in flag leaves. In conclusion, subsoiling in fallow period, together with N application at 150 kg·hm⁻², was beneficial to increase the protein yield and Glu/Gli in grains which improve the quality of wheat.     

水分胁迫对水稻籽粒蛋白质积累及营养品质的影响

以生产上广泛使用的水稻(Oryza sativa)品种‘汕优63’、‘扬稻6号’和‘武育粳3号’为材料,研究了水分胁迫对结实期水稻籽粒蛋白质积累及营养品质的影响。结果表明:正常施氮水平下,花后10~20 d的水分胁迫提高了谷氨酰胺合成酶(Glutamine synthetase,GS)和谷氨酸合酶(Glutamate synthase,GOGAT)活性,提高了籽粒自身利用无机氮合成氨基酸的能力,从而利于籽粒内蛋白质的积累,而高氮水平下,水分胁迫降低了籽粒自身合成氨基酸的能力。以重量为基数的蛋白质含有率在整个灌浆过程中呈“V”型消长,正常施氮水平下,水分胁迫明显提高了花后15 d至成熟期蛋白质含有率,而高氮水平下,水分胁迫处理的蛋白质含有率明显低于水层灌溉。与水层灌溉相比,水分胁迫提高了正常施氮水平下精米中醇溶蛋白和谷蛋白含量,但却明显降低了高氮水平下精米中醇溶蛋白和谷蛋白含量。水分胁迫对稻米中赖氨酸含量的影响因品种、植株的氮营养水平的不同而不同,水分胁迫显著降低了两种氮肥水平下‘汕优63’中赖氨酸含量,但却明显提高‘扬稻6号’中赖氨酸含量;而‘武育粳3号’于两种氮肥水平下表现恰好相反,正常施氮水平下赖氨酸含量略有升高;而高氮水平下赖氨酸含量明显降低。     

Phytohormone-mediated transformation of sugars to starch in relation to the activities of amylases, sucrose-metabolising enzymes in sorghum grain

Indole-acetic acid (IAA) and abscisic acid (ABA) were fed throughcomplete liquid medium (containing 2, 4, 8% sucrose) to detached earheads of sorghum. The effect of these phytohormones on interconvertion ofsugarsand their transformation to starch in relation to the activities of-, -amylases, sucrose-synthase (synthesis), sucrose-phosphatesynthase and soluble invertases was studied in the grain. This effect on theuptake of (U-¹⁴C) sucrose by detached ear heads and incorporation of¹⁴C into free sugars and starch of grain and into free sugars ofinflorescence parts was also studied. At concentrations of up to 4%sucrose in the culture medium, IAA increased the content of total free sugarsinthe grain. However, accumulation of starch and activities of - and-amylases increased when lAA was present even beyond the 4%sucroseconcentration in the culture medium. At all sucrose concentrations, the effectsof ABA and IAA were opposite. With 4% sucrose, both phytohormones causedmaximum accumulation of starch in the grain. ABA enhanced the relativeproportion of sucrose in the sugar pool with a concomitant reduction in theactivities of soluble acid (pH 4.8) and neutral (pH 7.5) invertases. Incontrast, IAA decreased the sucrose proportion of grain sugars with asimultaneous elevation and reduction in the activities of invertases andsucrose-phosphate synthase, respectively. Irrespective of sucrose concentrationin the culture medium, the activity of sucrose synthase (synthesis) wasenhancedwith IAA as well as ABA at their 10 M concentration. IAA alsoenhanced incorporation of ¹⁴C from (U-¹⁴C) sucrose intothe EtOH extract (principally constituted by free sugars) and starch of thegrain, but ABA caused the reverse effect. Based on the results, it is suggestedthat IAA and ABA have contrasting effects on the transformation of sucrose tostarch in sorghum grain where its capacity to synthesise starch is modulatedpositively by IAA and negatively by ABA.     

小麦旗叶Rubisco周转与籽粒含氮量的关系

随着旗叶的衰老,Rubisco含量逐渐减少.延缓小麦旗叶的衰老进程(抽穗期施氮肥),可增加旗叶Rubisco的含量,提高籽粒的全氮含量.在小麦旗叶全展后28d内,Rubisco的15N丰度处于较高水平,表明仍有Rubisco的重新合成;而在28d以后,Rubisco的15N丰度处于低水平,表明无Rubisco的重新合成.但这时籽粒的15N丰度却上升.旗叶全展后14d内Rubisco的15N丰度高于旗叶中全氮的丰度,说明此时期Rubisco重新合成的速率高于其它蛋白质;旗叶衰老过程中Rubisco的15N丰度的净转移高于全氮,Rubi-sco净N转移也高于全氮,表明Rubisco向籽粒中转移的氮素多于其它蛋白质,对籽粒含氮量的影响最大.     

Wheat ear carbon assimilation and nitrogen remobilization contribute significantly to grain yield

The role of wheat ears as a source of nitrogen (N) and carbon (C) in the grain filling process has barely been studied. To resolve this question, five wheat genotypes were labeled with ¹⁵N‐enriched nutrient solution. N remobilization and absorption were estimated via the nitrogen isotope composition of total organic matter and Rubisco. Gas exchange analyses showed that ear photosynthesis contributed substantially to grain filling in spite of the great loss of C due to respiration. Of the total kernel N, 64.7% was derived from the N acquired between sowing and anthesis, while the remaining 35.3% was derived from the N acquired between anthesis and maturity. In addition, 1.87 times more N was remobilized to the developing kernel from the ear than from the flag leaf. The higher yielding genotypes showed an increased N remobilization to the kernel compared to the lower yielding genotypes. In addition, the higher yielding genotypes remobilized more N from the ears to the kernel than the lower yielding genotypes, while the lower yielding genotypes remobilized more N from the flag leaf to the kernel. Therefore, the ears contribute significantly toward fulfilling C and N demands during grain filling.