Optimal nitrogen distribution within a leaf canopy under direct and diffuse light

Nitrogen distribution within a leaf canopy is an important determinant of canopy carbon gain. Previous theoretical studies have predicted that canopy photosynthesis is maximized when the amount of photosynthetic nitrogen is proportionally allocated to the absorbed light. However, most of such studies used a simple Beer's law for light extinction to calculate optimal distribution, and it is not known whether this holds true when direct and diffuse light are considered together. Here, using an analytical solution and model simulations, optimal nitrogen distribution is shown to be very different between models using Beer's law and direct–diffuse light. The presented results demonstrate that optimal nitrogen distribution under direct–diffuse light is steeper than that under diffuse light only. The whole‐canopy carbon gain is considerably increased by optimizing nitrogen distribution compared with that in actual canopies in which nitrogen distribution is not optimized. This suggests that optimization of nitrogen distribution can be an effective target trait for improving plant productivity.     

Decorrelating source and sink determinism of nitrogen remobilization during grain filling in wheat

Background and Aims

Nitrogen (N) remobilization is the major source of N for grain filling in wheat, the other being N uptake after anthesis (N_up); however, variations in remobilization efficiency are not fully understood. It is hard to tell whether the source or the sink effects predominate, because N in the culm at anthesis (N_ant) correlates strongly with both N remobilization (N_rem) and grain number (G_n), respectively the main source and the main sink.

Methods

A pot experiment was thus designed to assess the relative contributions of the source and sink to N_rem regulation. Using two cultivars of winter wheat (Triticum aestivum, ‘Apache’ and ‘Autan’), three pre-anthesis and two post-anthesis N fertilization levels were applied in order to vary the N sources, while ear trimming at anthesis reduced sink size.

Key Results

Unlike results observed at a scale of m², the equation binding N_ant to N_rem exhibited a negative intercept, challenging the concept of nitrogen remobilization efficiency. Before ear trimming, G_n fitted well to N_ant, with a slope dependent on genotype. To obtain a sink variable that was less correlated with N_ant, the difference δG_n was calculated between actual grain number and that which could be predicted from culm N before trimming. A multiple regression then predicted N_rem (r² = 0·95) from N_ant, N_up and δG_n, with fitting unbiased by fertilization treatment, trimming or genotype.

Conclusions

In untrimmed culms, δG_n had a negligible effect, so that N_rem could be fitted to N_ant and N_up only: grain N filling appeared to be determined by sources only (N_ant and N_up), not by sink, and the reduction of N_rem by N_up was quantified. In these ‘normal’ cases, the regulation of N_rem should thus be located within the N sources themselves. In contrast, ear-trimming needs to be considered with caution as it introduced a sink limitation on N_rem; moreover one with an important genotype effect.Key words: Triticum aestivum, winter wheat, source/sink, grain filling, nitrogen uptake, grain number, nitrogen harvest index, nitrogen remobilization efficiency, genotype × environment     

Plastid DNA during grain filling in wheat

During grain filling in wheat (Triticum aestivum L.) there is a progressive increase in the number of amyloplasts in the endosperm, as well as in cell number, DNA content and nuclear ploidy as the grain increases in size. The plastid DNA content also rises initially, and then there is a levelling off in the amount, with the percentage plastid DNA finally making up approximately 0.9% of the total endosperm DNA.     

旱地冬小麦灌浆期冠层温度与产量和水分利用效率的关系

利用红外测温仪,于2005～2006年在甘肃陇东旱原研究了我国北方冬麦区域的23个小麦品种(系)灌浆不同时期冠层温度的差异及其与产量和水分利用效率的关系。结果表明,不同基因型小麦在籽粒灌浆结实期存在着冠层温度高度分异的现象,其分异程度随灌浆过程的进行明显加大,到灌浆中后期达到最大。无论灌浆初期还是中期或中后期,旱地冬小麦产量、水分利用效率与冠层温度均呈极显著的负相关(R2=0.445-0.812),并且随着灌浆期推移,相关性增大,灌浆中后期冠层温度每升高1℃,旱地冬小麦产量减少近280 kg hm-2,水分利用效率下降约0.6 kg hm-2mm-1。灌浆中期以后不同基因型小麦冠层温度保持较高的一致性,冠层温度偏低的品种具有较高的产量和水分利用效率。灌浆中后期的冠层温度在评价小麦产量和水分利用效率上具有较高的可靠性,可作为一个田间选择指标应用。     

果树树冠遮阴对间作冬小麦籽粒灌浆的影响

果树-小麦间作模式是新疆南疆地区的主要农业种植模式.试验以扁桃-冬小麦(新冬20)间作模式为对象,通过主干分层形(DC)、小冠形(SC)、开心形(OC)和高干形(HS)4个树形和3个间作距离(分别距树干距离1.5、2.5和3.5 m)共同营造不同果树树冠遮光处理,以单作小麦为对照,对不同处理下冬小麦灌浆期环境因子、籽粒...     

Spatial distribution of proteins and metabolites in developing wheat grain and their differential regulatory response during the grain filling process

Grain filling and grain development are essential biological processes in the plant’s life cycle, eventually contributing to the final seed yield and quality in all cereal crops. Studies of how the different wheat (Triticum aestivum L.) grain components contribute to the overall development of the seed are very scarce. We performed a proteomics and metabolomics analysis in four different developing components of the wheat grain (seed coat, embryo, endosperm, and cavity fluid) to characterize molecular processes during early and late grain development. In-gel shotgun proteomics analysis at 12, 15, 20, and 26 days after anthesis (DAA) revealed 15 484 identified and quantified proteins, out of which 410 differentially expressed proteins were identified in the seed coat, 815 in the embryo, 372 in the endosperm, and 492 in the cavity fluid. The abundance of selected protein candidates revealed spatially and temporally resolved protein functions associated with development and grain filling. Multiple wheat protein isoforms involved in starch synthesis such as sucrose synthases, starch phosphorylase, granule-bound and soluble starch synthase, pyruvate phosphate dikinase, 14-3-3 proteins as well as sugar precursors undergo a major tissue-dependent change in abundance during wheat grain development suggesting an intimate interplay of starch biosynthesis control. Different isoforms of the protein disulfide isomerase family as well as glutamine levels, both involved in the glutenin macropolymer pattern, showed distinct spatial and temporal abundance, revealing their specific role as indicators of wheat gluten quality. Proteins binned into the functional category of cell growth/division and protein synthesis/degradation were more abundant in the early stages (12 and 15 DAA). At the metabolome level all tissues and especially the cavity fluid showed highly distinct metabolite profiles. The tissue-specific data are integrated with biochemical networks to generate a comprehensive map of molecular processes during grain filling and developmental processes.     

The effects of nitrogen application on growth and nitrogen distribution within the potato canopy

The effect of applying nitrogen (N) fertiliser on the growth and distribution of N within the potato canopy was studied in 1983 and 1984. In both years N was applied either in excess of that required to produce maximum tuber yields, or not at all. The large application of N changed the pattern of canopy growth - stimulating growth of leaves at the top of the stem, particularly lateral branches, for longer during the season, and accelerating the death of (shaded) leaves at the base of the canopy. The pattern of canopy senescence was, therefore, changed from a synchronous to a progressive type. Application of nitrogen fertiliser at supra-optimal rates increased the N contents of leaves, stems and tubers. The extra N in the leaves of these plants was present as reduced N in all leaf positions, and as nitrate (NO^-₃) in the lowermost leaves. In addition, substantial quantities of NO^-₃ were also stored in the stems. Part of this extra N in the canopy was redistributed during subsequent growth, especially to the lateral branches as crop N uptake slowed towards the end of the season. In addition, substantial quantities of N were also potentially available for redistribution to the growing tubers. There was little redistribution of N from the leaves of N-deficient plants. It is suggested that redistribution of N in the canopy of N-replete plants allowed the growth of lateral branches towards the end of the season, thereby maintaining photosynthetically active leaves for longer than N-deficient plants.     

Acclimation of photosynthesis to light and canopy nitrogen distribution: an interpretation

BACKGROUND AND AIMS: Acclimation of photosynthesis to light and its connection with canopy nitrogen (N) distribution are considered. An interpretation of a proportionality between light-saturated photosynthesis and local averaged leaf irradiance is proposed by means of a simple model. MODEL: The model assumes (a) local irradiance drives synthesis of photosynthetic protein from metabolic N; (b) photosynthetic N is slowly degraded over approx. 5-7 d; (c) metabolic N is equally available through the canopy. CONCLUSIONS: The kinetics of acclimation at different light levels may provide a way of parameterizing and testing the model. The model provides a rationale for the proportionality assumption mentioned above, which, while it is consistent with much experimental work, is valuable because it allows canopy photosynthesis to be calculated analytically.     

春小麦灌浆期籽粒沉淀值动态变化及氮磷肥与播期的影响

通过3个品质类型春小麦品种施肥和播期大田试验,建立了灌浆期籽粒沉淀值动态变化的曲线拟合方程,定量揭示籽粒沉淀值动态变化规律及氮磷肥与播期的影响效应.结果表明,自开花15 d始至成熟,籽粒沉淀值随时间变化呈先升高后降低的单峰曲线变化.在灌浆期籽粒沉淀值的动态变化过程中,不同基因型特点不同,且基因型间的关系也随之发生变化.各品种沉淀值积累速度的变化决定了成熟时高蛋白强筋品种沉淀值最高、高蛋白中筋品种次之、低蛋白弱筋品种最低.氮水平增加,高蛋白强筋和低蛋白弱筋品种沉淀值和曲线最高值降低,高蛋白中筋品种则明显升高;高蛋白强筋品种曲线最高值出现时间提前,而高蛋白中筋和低蛋白弱筋品种则推后.磷水平增加,高蛋白品种沉淀值和曲线最高值普遍提高,而低蛋白品种则降低;各基因型曲线最高值出现时间均推迟.氮磷（钾）素平衡配施是形成较高沉淀值的关键.在没有水分胁迫情况下,光温互作是影响籽粒沉淀值动态形成的首要条件,其次为降水;而≥10 ℃积温则为最敏感因子,即在较高光温条件互作前提下,增加灌浆期≥10 ℃积温则提高各基因型品种沉淀值.高蛋白比低蛋白品种更易受氮磷水平和气象条件影响.     

强筋与弱筋小麦籽粒蛋白质组分与加工品质对灌浆期弱光的响应

选用强筋小麦品种济麦20和弱筋小麦品种山农1391,在大田试验条件下,分别于籽粒灌浆前期(花后6—9 d)、中期(花后16—19 d)和后期(花后26—29 d)对小麦进行弱光照处理,研究了籽粒产量、蛋白质组分及加工品质的变化。灌浆期弱光显著降低小麦籽粒产量,灌浆中期对济麦20和灌浆后期对山农1391的产量降幅最大。弱光处理后,籽粒氮素积累量及氮素收获指数减少。但弱光使籽粒蛋白质含量显著升高,其中灌浆中期弱光升幅最大,原因可能是由于其粒重降低造成的。弱光对可溶性谷蛋白无显著影响,但增加不溶性谷蛋白含量,使谷蛋白聚合指数显著升高,面团形成时间和稳定时间亦升高,籽粒灌浆中、后期弱光对上述指标的影响较前期大。灌浆期短暂的弱光照对改善强筋小麦粉质仪参数有利,但使弱筋小麦变劣;并均伴随籽粒产量的显著降低这一不利影响。     

Hormonal changes with uniconazole trigger canopy apparent photosynthesis and grain filling in wheat crop in a semi-arid climate

Protoplasma - Phytohormones are important for the growth and development of plants. The objective of the experiment was to investigate the effect of foliar application of uniconazole (UCZ) at the...     

枣麦间作系统中光能在作物群体内分布的数值模拟

应用农业气象学原理和方法,依据实测资料,分析了枣麦间作系统中枣树带遮荫宽度及阴影内的相对光照度的时空变化规律．以及遮荫区、非遮荫区内小麦群体的光能垂直分布特点;应用数理方法,模拟了遮荫区及非遮荫区内小麦群体光能垂直分布曲线．结果表明,南北走向的枣树带每天早、晚其遮荫区域霉宽,相对光照度较大;中午前后,遮荫区域宽度较窄,相对光照度较小．尽管遮荫区与非遮荫区内小麦群体的光能垂直分布有明显的差异。但二者间都符合高斯曲线的递减规律．     

Subtilisin-like serine proteases involved in N remobilization during grain filling in wheat

The induction of two subtilisin-like proteases (P1 and P2) associated with stress-induced senescence in young plants was investigated in adult wheat plants during the grain-filling period. Western blot analysis of flag leaf extracts showed that P1 was induced very late in the life cycle of the plants (9 days post-anthesis) and that 7 days later it reached a 2.5-fold increase with respect to the initial value at anthesis. On the other hand, the P2 signal was already detected previous to anthesis and increased soon after anthesis, reaching a fourfold increase by the end of the grain-filling period. The induction of P1 and P2 temporally correlates with the degradation of the Rubisco small and large subunits in the flag leaf, as well as with nitrogen (N) accumulation in the ears. At the same time, a decrease in the endogenous concentration of the cytokinins isopentenyladenine and isopentenyladenosine (iP + iPA) in the leaves was observed. In detached leaves senescing in the dark, the levels of both proteases were affected by 6-benzylaminopurine application: the induction of P1 was completely prevented, whereas the induction of P2 was reduced. Our findings demonstrate that both P1 and P2 are expressed in leaves of adult plants and are induced during natural senescence. These results enable us to postulate their participation in N remobilization to developing grains during monocarpic senescence and their regulation by a cytokinin-mediated mechanism.     

氮肥运筹对小麦产量、氮素利用效率和光能利用率的影响

连续2年在西南冬麦区的重庆、仁寿、广汉、西昌4个地点,开展3种施氮水平(每公顷纯氮0、120、180 kg,简写为N₀、N₁₂₀、N₁₈₀)和3种氮肥分配模式(N_A:底肥100%;N_B:底肥70%+苗肥30%;N_C:底肥60%+拔节肥40%)的田间试验,监测小麦花后冠层叶片SPAD值、群体光合速率(CAP)、光能利用等生理参数和籽粒产量,计算氮素利用效率、光能利用率等.结果表明: 随施氮水平增加,小麦上三叶SPAD值、CAP、光合有效辐射(PAR)截获率和产量均呈增加趋势,而氮肥农学利用效率、生产效率、吸收效率和利用效率呈降低趋势.氮肥后移的增效作用因施氮水平而异,SPAD于N₁₈₀增效明显,而CAP于N₁₂₀增效明显,不同氮肥管理模式的光能利用率因地点而异.氮肥后移能明显提高小麦氮肥农学效率、生产效率、吸收效率和氮素表观回收率,但氮肥利用效率则略有减少.氮肥后移效果N_C总体优于N_B处理.不同地点比较,广汉的SPAD值、CAP、PAR截获率、氮肥利用参数较高,其产量也相应最高;西昌的产量、SPAD值及氮素利用效率较高,但其光能利用率和CAP较低;重庆和仁寿的SPAD值、光能利用率及氮素利用效率均较低,其产量也最低.小麦生物产量与各地点的籽粒产量、CAP、SPAD值和PAR累积截获量均呈显著或极显著的正相关关系.表明不同生态区域增施氮肥都能促进小麦增产,氮肥后移可进一步优化产量结构、改善氮肥和光能利用效率,但存在年份和地点差异,各地需要制定有针对性的氮肥管理模式.     

灌浆期高温和水分逆境对冬小麦籽粒蛋白质和淀粉含量的影响

灌浆期高温和水分逆境是影响小麦籽粒产量和品质的关键气候因子。以扬麦9号、徐州26和豫麦34三个小麦品种为材料,利用人工气候室模拟灌浆期高温和水分胁迫环境,研究了花后高温及温度和水分互作对小麦籽粒蛋白质和淀粉形成的影响。结果表明,高温显著提高了小麦籽粒蛋白质含量及清蛋白、球蛋白和醇溶蛋白含量,但降低了谷蛋白含量,导致麦谷蛋白／醇溶蛋白比值降低。高温显著降低了籽粒总淀粉和支链淀粉含量及支／直比。籽粒蛋白质和淀粉及其组分形成所需的适宜昼夜温差随小麦品质类型而异,但温度水平对籽粒蛋白质和淀粉的影响较温差大。在高温和水分逆境下,温度对籽粒蛋白质和淀粉含量的影响较水分逆境大,且存在显著的互作效应。小麦籽粒蛋白质含量均表现为干旱〉对照〉渍水,以高温干旱最高,适温渍水最低;淀粉含量为对照〉干旱〉渍水,以适温对照最高,而高温渍水最低。高温和水分逆境显著提高了籽粒醇溶蛋白含量而降低了谷蛋白含量及支链淀粉含量,使蛋白质谷／醇比和淀粉支／直比降低,以高温渍水对籽粒蛋白质和淀粉组分的影响最为显著。不同品种之间,高蛋白小麦籽粒蛋白质和组分的形成受高温和水分逆境的影响更大,而低蛋白品种籽粒淀粉形成显著受温度和水分逆境的调节。分析表明,在高温和水分逆境下籽粒蛋白质含量与清蛋白和醇溶蛋白显著正相关,籽粒淀粉含量与谷蛋白、支链淀粉含量及支／直比显著正相关。     

Partitioning of leaf nitrogen with respect to within canopy light exposure and nitrogen availability in peach (Prunus persica)

Summary Relationships between leaf nitrogen content and within canopy light exposure were studied in mature nectarine peach trees (Prunus persica cv. Fantasia) that had received 0, 112, 196, 280 or 364 kg of fertilizer nitrogen per hectare per year for the previous 3 years. The relationships between light saturated leaf CO₂ assimilation rates and leaf nitrogen concentration were also determined on trees in the highest and lowest nitrogen fertilization treatments. The slope of the linear relationship between leaf N content per unit leaf area and light exposure was similar for all nitrogen treatments but the y-intercept of the relationship increased with increasing N status. The slope of the relationship between leaf N content per unit leaf area and light saturated CO₂ assimilation rates was greater for the high N trees than the low N trees, but maximum measured leaf CO₂ assimilation rates were similar for both the high and low N treatments. A diagrammatic model of the partitioning of leaf photosynthetic capacity with respect to leaf light exposure for high and low nitrogen trees suggests that the major influence of increased N availability is an increase in the photosynthetic capacity of partially shaded leaves but not of the maximum capacity of highly exposed leaves.     

Leaf nitrogen remobilisation for plant development and grain filling

A major challenge of modern agriculture is to reduce the excessive input of fertilisers and, at the same time, to improve grain quality without affecting yield. One way to achieve this goal is to improve plant nitrogen economy through manipulating nitrogen recycling, and especially nitrogen remobilisation, from senescing plant organs. In this review, the contribution of nitrogen remobilisation efficiency (NRE) to global nitrogen use efficiency (NUE), and tools dedicated to the determination of NRE are described. An overall examination of the physiological, metabolic and genetic aspects of nitrogen remobilisation is presented.     

The response of alpha-amylase activity during wheat grain development to nitrogen fertiliser

The aim of this study was to test whether the effect of nitrogen fertiliser on Hagberg falling number of winter wheat (Triticum aestivum) grain in the absence of sprouting is mediated by pre-maturity alpha-amylase activity and is related to grain drying rate. A field experiment with two cultivars (Avalon and Mercia) in 1990 and 1991 compared four application rates of nitrogen. Samples of grain were taken at intervals during development for moisture determination and alpha-amylase assay. Grains from plots given nitrogen dried faster in both years, but alpha-amylase activity and Hagberg falling number responded differently to nitrogen in the two years. In the warmer and drier year of 1990, alpha-amylase activity declined throughout development leading to very high Hagberg falling number at harvest, with little effect of nitrogen. In the cooler and wetter year of 1991, alpha-amylase activity declined until about 30% moisture. After this stage, alpha-amylase activity increased in the absence of sprouting in grain from plots receiving little or no nitrogen. This resulted in a linear increase in Hagberg falling number in response to nitrogen fertiliser. Electrophoresis of alpha-amylase isozymes indicated that the increase in Hagberg falling number in response to nitrogen was not mediated by a decrease in retained pericarp alpha-amylase activity, but by a reduction in pre-maturity alpha-amylase activity. These results support the hypothesis that slow grain drying enhances pre-maturity alpha-amylase formation, and also support the hypothesis that an additional environmental factor varying between seasons is involved in pre-maturity alpha-amylase formation.     

Wheat nitrogen metabolism during grain filling: comparative role of glumes and the flag leaf

The mobilization of nitrogen (N) compounds and the roles played by glumes and the flag leaf during grain filling were studied in bread wheat (Triticum aestivum L. cv. Florida) grown under field conditions. Glumes lost twice as much of their total N content as that lost by the flag leaf between the milk and early dough stages. In the flag leaf, glumes and grains, Glu, Asp, Ser and Ala accounted for 85% of all the reductions in the free amino acid pool. Principal component analysis of free amino acid pools separated grains from the glumes and the flag leaf, suggesting grain specific regulations in the use of free amino acids in protein synthesis. In all three organs, no decrease in Gln was detected, probably due to steady glutamine synthetase (GS; EC 6.3.1.2) activities per soluble protein in both the flag leaf and glumes. Compared with the flag leaf, glumes presented relatively smaller amounts of the chloroplast GS associated isoform. This we show is due to a lower relative number of mesophyll cells in glumes as supported by the different anatomy and the cellular pattern of the GS immunolocalization. We argue that cellular distribution plays a key role in supporting metabolism to enable the various functions undertaken by glume tissue.     

Interactions between polyamines and ethylene during grain filling in wheat grown under water deficit conditions

This study was to test the hypothesis that polyamines (PAs) and ethylene may be involved in mediating the effect of water deficit on grain filling. Two wheat cultivars, drought-tolerant Shannong16 (SN16) and drought-sensitive Jimai22 (JM22), were used and subjected to well-watered and severe water deficit (SD) during grain filling. SD reduced the weight of superior and inferior grains, by 7.38 and 23.54 % in JM22, 13.8 and 2.2 % in SN16, respectively. Higher free-spermidine (Spd) and free-spermine (Spm) concentration and lower free-putrescine (Put) concentration, ethylene evolution rate (EER) and 1-aminocylopropane-1-carboxylic acid (ACC) concentration were found in superior grains than those in inferior ones. Opposite to the variations of Spd and Spm concentration, ACC, Put concentration and EER were significantly increased under SD. The percentage variation of PAs and ACC differed with cultivars and grain types. ACC concentration of superior and inferior grains under SD increased significantly at 21 days post-anthesis, by 90 and 164 % in JM22, 65 and 13.2 % in SN16, respectively. The equivalent value of Put concentration was 1.04 and 7.9 % in JM22, 34.4 and 10.3 % in SN16. Spd concentration of superior grains showed a higher decrease than that of inferior ones in both cultivars, while Spm exhibited an opposite trend between both grain types. These percentage variations were highly consistent with the differed responses of weight of both grain types to SD in JM22 and SN16. Grain filling rate was negatively correlated with EER and ACC concentration, while positively correlated with Spd and Spm concentration as well as the ratio of Spd or Spm to ACC. Exogenous Spd or aminoethoxyvinylglycine (an inhibitor of ethylene synthesis by inhibiting ACC synthesis) obviously reduced ACC concentration and EER and increased Spd and Spm concentration, while exogenous ethephon (an ethylene-releasing agent) or methylglyoxal-bis (an inhibitor of Spd and Spm synthesis) showed the opposite effects. The results suggested that it would be good for wheat to have the physiological traits of higher Spd and Spm, as well as a higher Spd/ACC or Spm/ACC, under SD.