The effects of variable nitrogen and phosphorous concentrations on Eriophorum vaginatum tillers grown in nutrient solutions

Eriophorum vaginatum tillers were collected at Eagle Creek, Alaska and cultivated in aerated solutions under controlled environmental conditions. The nutrient solutions contained traces, 1.05 and 21 mg l⁻¹ N (nitrate) and traces, 0.15 and 3.10 mg l⁻¹ P (phosphate), pH was maintained at 5.5. The high N, 21 mg l⁻¹, and P, 3.18 mg l⁻¹, nutrient solution produced significant biomass increases. Functional leaf areas were significantly enhanced by high N and P doses in the solutions. Root surface areas varied considerably between treatments; however, significant differences were not found. The mean root surface area of a tiller reached 126 cm² (range 35–290 cm²), whereas the functional leaf area was 6.8 cm² (range 3.3–20.3 cm²). Tillers growing in the highest N + P solutions produced twice the number of daughter tillers as tillers growing in solutions with trace amounts of N and P.     

Influence of nitrogen supply and sink strength on changes in leaf nitrogen compounds during senescence in two wheat cultivars

Changes in various nitrogen compounds during senescence of the fourth leaf were studied in two cultivars of spring wheat (Triticum aestivum L.). One of the cultivars (Yecora) was supplied with two N levels; the other (Tauro) was grown with the high N level and pruned above the fourth leaf, whereas the control was left intact. In both cultivars grown with high N supply, net nitrogen export from the fourth leaf did not occur until 35 days after sowing (DAS). Loss of leaf soluble proteins started earlier than that of chlorophylis, and coincided initially with an increase in insoluble protein. In N deficient plants the level of total N, soluble protein, and the activity of nitrate reductase (NRA. EC 1.6.6.1) started to decrease about 5 days earlier, and along with chlorophyll, continued to decrease at a faster rate, than in high N plants. Also, with low N supply, the large subunit (LSU, 58 kDa) of ribulose-1.5-bisphosphate carboxylase/oxygenase (Rubisco, EC 4.1.1.39) decreased in greater proportion than other soluble proteins, while with high N supply the decrease in Rubisco LSU was similar to that of other soluble proteins. Nitrogen deficiency caused a greater decrease in soluble proteins than in insoluble proteins, and NRA relative to soluble proteins. The faster senescing Tauro cultivar had lower levels of most parameters, especially NRA, soluble protein and, after 35 DAS. Rubisco LSU as a proportion of soluble protein. The decrease in sink strength due to shoot pruning did generally not affect the level of the various nitrogenous compounds until 35 DAS; thereafter the levels of most parameters, especially soluble protein, Rubisco LSU and, at late stages of senescence, insoluble protein, were higher in pruned than in control shoots. Thus, shoot pruning slows down senescence. The 56- and 78-kDa polypeptides increased, rather than decreased, with leaf age; the level of these two polypeptides showed a negative relationship with Rubisco LSU (r = -0.933 and r = -0.758, respectively).     

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     

Modeling grain nitrogen accumulation and protein composition to understand the sink/source regulations of nitrogen remobilization for wheat

A functional explanation for the regulation of grain nitrogen (N) accumulation in cereal by environmental and genetic factors remains elusive. Here, new mechanistic hypotheses of grain N accumulation are proposed and tested for wheat (Triticum aestivum). First, we tested experimentally the hypothesis that grain N accumulation is mostly source regulated. Four contrasting cultivars, in terms of their grain N concentrations and yield potentials, were grown with non-limiting N supply. Grain number per ear was reduced by removing the top part of the ear at anthesis. Reduction in grain number gave a significant increase in N content per grain for all cultivars, showing that grain N accumulation was source regulated. However, on a per ear basis, cultivars with a high grain number fully compensated their N accumulation for reduced grain number at anthesis. Cultivars with a lower grain number did not compensate completely, and grain N per ear was decreased by 16%. Second, new mechanistic hypotheses of the origins of grain N source regulation and its response to environment were tested by simulation. The hypotheses were: (a). The regulation by N sources of grain N accumulation applies only for the storage proteins (i.e. gliadin and glutenin fractions); (b). accumulation of structural and metabolic proteins (i.e. albumin-globulin and amphiphilic fractions) is sink-regulated; and (c). N partitioning between gliadins and glutenins is constant during grain development and unmodified by growing conditions. Comparison of experimental and simulation results of the accumulation of grain protein fractions under wide ranges of N fertilization, temperatures, and irrigation supported these hypotheses.     

Light quality effects on the appearance of tillers of different order in wheat (Triticum aestivum)

Plants of Triticum aestivum cvs Norkin Pan 70, La Paz Inta, Buck Cencerro and Buck Cimarron were grown outdoors in individual pots with two sowing dates and irradiated at the end of the day either with red or far-red light. In red-treated plants the number of tillers was close to the potential calculated from the number of leaves on the main shoot but far-red treated plants produced less tillers. The magnitude of the effect was larger for secondary and tertiary than primary tillers. Thus, the proportion of primary to secondary and tertiary tillers was larger in FR-treated plants. Leaf sheath and lamina were longer under FR in the second sowing date, possibly due to warmer nights. These responses were greatly affected by the genotype. Significant differences in ear number were not found but in one of the sowing dates red-light-treated plants showed a lower grain number and yield than those treated with far-red.     

Effects of source/sink manipulation on net photosynthetic rate and photosynthate partitioning during grain filling in winter wheat

Source-sink relationship, which was influenced by both genotype and environmental factors, contributed to the variation in photosynthesis and photosynthate partitioning of wheat. Source reduction by partial defoliation increased leaf net photosynthetic rate (PN), and sink reduction decreased PN of irrigated wheat. However, the change in PN varied among genotypes. Source reduction enhanced photosynthate translocation into grain in irrigated wheat. However, the enhancement was more evident in cv. Lumai 215953 than incv. Lumai 15. Sink reduction had little effect on the translocation of photosynthate into grain in cv. Lumai 15, but decreased the translocation of photosynthate into grain and increased it into stem in cv. Lumai 215953. In rainfed, non-irrigated wheat, the source or sink manipulation influenced PN only slightly. The source reduction decreased the partitioning of photosynthates into the upper parts (including grains) of plant. However, very little effects of sink reduction on the production of photosynthates occurred in rainfed wheat. This showed that grain sink size was not a factor limiting the production of photosynthates, but controlled the partitioning of photosynthates. Sink reduction decreased photosynthate translocation into grains, and increased it into upper parts of rainfed wheat plant. This revised version was published online in July 2006 with corrections to the Cover Date.     

冬小麦拔节期不同茎蘖对低温胁迫的反应及抗冻性评价

以小麦济南17和山农8355为材料,在低温胁迫条件下,测定了不同茎蘖功能叶和叶鞘超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)活性及丙二醛(MDA)和可溶性蛋白含量,并利用主成分分析、聚类分析对其抗冻性进行综合评价.结果表明： 低温胁迫条件下,小麦拔节期不同茎蘖功能叶和叶鞘中SOD、POD和CAT活性均不同程度地上升,MDA和可溶性蛋白含量则不同程度地上升或下降.利用主成分分析和聚类分析,将济南17不同茎蘖分为3类：主茎和一级分蘖Ⅰ、Ⅱ属强抗冻蘖组,一级分蘖Ⅲ、Ⅳ和二级分蘖Ⅰp属中度抗冻蘖组,二级分蘖Ⅱp属弱抗冻蘖组;将山农8355不同茎蘖分为3类：主茎和一级分蘖Ⅰ、Ⅱ、Ⅲ属强抗冻蘖组,一级分蘖Ⅳ和二级分蘖Ⅰp属中度抗冻蘖组,二级分蘖Ⅱp属弱抗冻蘖组.表明冬小麦拔节期不同茎蘖存在抗冻性差异,且低位蘖较高位蘖抗冻.
     

Carbon dioxide exchange in relation to sink demand in wheat

Summary In this paper, experiments are described which examine the effect of requirement for assimilates by the ear on the rate of net photosynthesis in leaves of wheat (Triticum aestivum L.). Different levels of requirement were achieved by various levels of sterilization of florets just before anthesis, which resulted in a range of grain numbers per ear, and by inhibiting photosynthesis of the intact ear by 3-(3,4-dichlorophenyl)-1,1-dimethylurea (DCMU). Only the ear and two uppermost leaves of the main shoot were considered, all the lower leaves and tiller leaves being excised when the experimental treatments were imposed. In two experiments, tiller regrowth was permitted during the experimental period, while in a third, new tillers were defoliated regularly.The response of leaf photosynthesis to the level of assimilate requirement by the ear was influenced by the treatment of the vegetative tillers. Thus, the net photosynthesis rate of the flag leaf was decreased by a reduction in grain number, or increased by inhibition of photosynthesis in the ear, only when the vegetative tillers were kept defoliated; when these tillers were allowed to re-grow normally, there was no influence of ear treatment on leaf photosynthesis. Temporal changes in leaf photosynthesis were consistent with this response pattern, i.e., when tillers were defoliated, the initial high rates of photosynthesis persisted for much longer.In the experiment where photosynthesis was influenced by the requirement for assimilate in the ear, the variation occurred through change in stomatal conductance on the abaxial surface of the leaf. This surface has a lesser conductance to CO₂ exchange than the adaxial surface. The implication of this finding to rapid methods of plant screening is discussed.     

Postanthesis allocation of photosynthates and grain growth in wheat cultivars as affected by source/sink change

Two wheat cultivars, Hesheng 2 with large grain yield potential, and Shannong 505 with small grain yield potential, were used for investigating the responses of postanthesis photosynthesis, dry matter accumulation and allocation, and grain growth to source/sink changes. At the initial grain filling stage, Hesheng 2 was sensitive to source reduction leading to an increase of net photosynthetic rate (P_N) by 10 %; however, little effect of sink reduction was observed. In Shannong 505, P_N was obviously decreased by sink reduction, and changed a little after source reduction. At the rapid grain filling stage, Hesheng 2 was sensitive to both source and sink reduction resulting in the increase or decrease of P_N, respectively. However, the response of P_N in Shannong 505 to source/sink changes was similar to that in previous stage. The dry matter (DM) accumulation after anthesis was affected by source/sink changes. In Hesheng 2, the decrease in DM was higher than that in Shannong 505 after the same source or sink reduction. Source reduction caused a decrease in the allocation of DM to the sheath and stem, and promoted the reserve photosynthates to be reallocated to grain. The effect of sink reduction was contrary. The grain mass of Hesheng 2 was more easily regulated by source/sink changes than that of Shannong 505. The effect source/sink changes on grain mass was in order upper > basal > middle spikelets on spike. As for a spikelet, the effect was found mainly in the grain mass at the positions 3 and 4 from base of the spikelet. This revised version was published online in July 2006 with corrections to the Cover Date.     

Source-sink manipulation effects on postanthesis photosynthesis and grain setting on spike in winter wheat

Source-sink manipulation could regulate the net photosynthetic rate (PN) of winter wheat after anthesis, however, the direction and magnitude of the regulation varied with time after anthesis. The PN was significantly increased by source reduction at the initial time of grain filling, but sink reduction had little influence on the PN, which suggested that the sink (spike) limitation did not occur at this time. Source-sink relation markedly affected PN during rapid grain filling. The PN was increased by source reduction and decreased by sink reduction significantly, which indicated that PN was closely associated with the change of source or sink size. The effect of source-sink manipulation on PN had some relationship with the occurrence of plant senescence at the time of late grain filling. Source reduction accelerated the senescence and dropped the PN, meanwhile, sink reduction delayed the senescence and promoted the PN. A direct relation between the effect of source-sink manipulation on PN and stomatal limitation was not found. Removing one quarter of leaves (RQ) had little influence on spike development after anthesis. In this case there was enough compensation in source production through photosynthesis. Removing one half of leaves (RH) made grain mass per spike and mass of grains lowered, especially the grain mass in the top and base positions of spike declined markedly. The source supply was grain-limiting. Removing one quarter of spikelets (RS) was beneficial to grain-setting in the remaining spikelets, leading to the increase of grain mass. Thus promoting the source supply of photosynthates after anthesis is of major importance for grain to set and to develop.     

灌水量和灌水时期对小麦耗水特性和氮素积累分配的影响

在田间试验条件下,以小麦品种济麦20为材料,研究了不同灌水处理对小麦的耗水特性和氮素积累分配的影响.试验设置7个处理:不浇水(W0);拔节期和开花期浇水,每次灌水量为30mm(W1)、60mm (W2)、90mm(W3);拔节期、开花期和灌浆期浇水,每次灌水量为30mm(W4)、60mm (W5)、90mm(W6).研究结果表明:(1)随灌水量的增加,总耗水量逐渐增加,土壤耗水量和降水量占总耗水量的比例降低.产量和水分利用率最高的W2和W4处理总耗水量分别为413.87,362.15mm;灌溉量、降水量、土壤耗水量分别占总耗水量的比例为29%、36.34%、34.66%,24.85%、41.53%、33.62%;两个处理比较,W4处理提高了对降水的利用比例,但降低了对灌溉水的利用比例.通过对全生育期0～200cm不同土层土壤耗水量的研究得出,W0和W1处理的深层土壤耗水量较低,W3、W5、W6处理的0～200cm 每个土层土壤耗水量均较低,对W2和W4处理,小麦能够利用120～200cm的深层土壤水分,其土壤贮水消耗量显著增加.(2)W2处理的籽粒氮素积累量较高,W1、W4处理籽粒中的氮素分配比例显著高于其它处理,有灌浆水的处理,尤其是灌浆水高于30mm的处理,营养器官氮素转移率和贡献率显著降低;W4处理的籽粒蛋白质含量较高,W2和W4处理的籽粒蛋白质产量显著高于其它处理.(3)籽粒产量随着灌水量的增加先升高后降低,其中W2和W4处理显著高于其它处理;W4处理的产量水分利用效率和蛋白质产量水分利用率显著高于其它处理.结果表明,W4为本试验条件下高产节水的最佳灌水处理.     

Long- and short-term effects of decreased sink demand on carbohydrate levels and photosynthesis in wheat leaves

Wheat (Triticum aestivum L.) ears were removed to investigate long-term regulation of photosynthesis by sink demand at ambient CO₂ and 22 °C. The CO₂ level was also increased to 660 μmol mol^?1 and temperature was lowered to 5 °C to examine short-term responses of photosynthesis to low sink demand. Sink removal inhibited photosynthesis and increased leaf levels of glucose, fructose and ribulose-1, 5-bisphosphate (RuBP), and the glucose-6-phosphate (G6P)/fructose-6-phosphate (F6P) and RuBP/3-phosphoglycerate (PGA) ratios under growth conditions, but had no effect on the activity and activation state of ribulose-1, 5-bisphosphate carboxylase oxygenase (Rubisco) either under growth or short-term conditions, suggesting an inhibition of photosynthesis by decreased in vivo catalysis of Rubisco. Photosynthesis increased similarly in eared and earless shoots after a rise in CO₂ concentration, and the ratio of triose-phosphates (glyceraldehyde 3-phosphate and dihydroxyacetone phosphate, TP) to PGA was similar or higher for removed than intact ears, suggesting that feedback inhibition of photosynthesis was not caused by a limitation of ATP synthesis in chloroplasts. Under short-term conditions (660 μmol mol^?1 CO₂, 5 °C), TP and RuBP levels and the TP/PGA and TP/RuBP ratios were increased by sink removal, indicating an additional limitation of photosynthesis by the rate of RuBP regeneration.     

Experiments with CCC on wheat: effects of spacing, nitrogen and irrigation

In experiments with spring and winter wheat at Rothamsted and Woburn during 4 years CCC increased yield at close spacing (4 in) (10 cm) more than at usual spacing (8 in) (20 cm), but there was no interaction between spacing and yield. Some experiments tested up to 2·4 cwt/acre (300 kg/ha) N to see whether yields continued to increase with more than usual amounts of N, when CCC prevented lodging. There was no evidence of this. When a short dry spell occurred at ear emergence, yield of spring wheat was increased by 6 cwt/acre (750 kg/ha) by CCC and 10 cwt (1250 kg) by irrigation. CCC probably improves yield in these conditions because the larger root system it causes enables more ear-bearing shoots to survive. CCC increases yield in two ways, either by increasing ears or grain per ear. In an unlodged crop CCC usually makes the grains smaller, but by preventing lodging it can also increase size. Usually CCC decreases the leaf area per shoot. The flag leaf may be smaller, unchanged or larger than those of untreated plants. There was no obvious connexion between flag-leaf area and grain yield; when CCC decreased flag-leaf area duration by 25 %, grain yield was unchanged. The results suggest that using CCC gives a more than even chance of a profitable yield increase.     

Growth and photosynthesis in bell-pepper as affected by sink manipulation

Developing fruits on lower nodes were major reproductive sink for photosynthates in bell pepper (Capsicum annuum L.). The removal of these fruits resulted in faster growth of other fruits on upper nodes. Sink manipulated plants have the higher rate of net photosynthetic rate at the later stages of plant development. This revised version was published online in July 2006 with corrections to the Cover Date.     

Concentrative nitrogen allocation to sun-lit branches and the effects on whole-plant growth under heterogeneous light environments

We investigated the nitrogen and carbohydrate allocation patterns of trees under heterogeneous light environments using saplings of the devil maple tree (Acer diabolicum) with Y-shaped branches. Different branch groups were created: all branches of a sapling exposed to full light (L-branches), all branches exposed to full shade (S-branches), and half of the branches of a sapling exposed to light (HL-branches) and the other half exposed to shade (HS-branches). Throughout the growth period, nitrogen was preferentially allocated to HL-branches, whereas nitrogen allocation to HS-branches was suppressed compared to L- and S-branches. HL-branches with the highest leaf nitrogen content (N_area) also had the highest rates of growth, and HS-branches with the lowest N_area had the lowest observed growth rates. In addition, net nitrogen assimilation, estimated using a photosynthesis model, was strongly correlated with branch growth and whole-plant growth. In contrast, patterns of photosynthate allocation to branches and roots were not affected by the light conditions of the other branch. These observations suggest that tree canopies develop as a result of resource allocation patterns, where the growth of sun-lit branches is favoured over shaded branches, which leads to enhanced whole-plant growth in heterogeneous light environments. Our results indicate that whole-plant growth is enhanced by the resource allocation patterns created for saplings in heterogeneous light environments.     

植物叶片氮分配及其影响因子研究进展

氮是植物生长的基本限制性因子,它的有效利用可以增加植物的适应性。叶片氮分配是指氮在植物叶片细胞各细胞结构以及游离化合物中所分配的比例。叶片氮的分配方式决定了叶片光合作用的强弱,影响叶片的坚韧程度以及化学防御强度,因此研究氮在植物叶片内的分配方式具有重要意义。阐述了叶片氮分配的方式,分析了影响叶片氮分配的生物和非生物因子(CO2,光,土壤养分),介绍了常用的叶片氮分配的研究方法,并对未来的研究进行了展望。     

小麦与蚕豆间作系统氮肥调控对小麦白粉病发生及氮素累积分配的影响

通过田间小区试验,设N₀(0 kg·hm^-2)、N₁(112.5 kg·hm^-2)、N₂( 225 kg·hm^-2)、N₃( 337.5 kg·hm^-2) 4个施氮水平,研究不同施氮水平下小麦与蚕豆间作对小麦白粉病发生、植株氮含量和氮素累积分配的影响,探讨间作系统氮肥调控下小麦植株氮素含量、氮素累积分配与白粉病发生的关系.结果表明: 无论单作还是间作,施氮(N₁、N₂和N₃)均增加了小麦籽粒产量,以N₂水平下产量最高,单、间作分别为4146和4679 kg·hm^-2;施氮加重了小麦白粉病的发生与危害,N₁、N₂和N₃水平下病害进展曲线下的面积(AUDPC)分别平均增加39.6%～55.6%(基于发病率DI)和92.5%～217.0%(基于病情指数DSI),病情指数受氮素调控的影响较发病率大;施氮显著提高小麦植株氮含量(8.4%～51.6%)和氮素累积量(19.7%～133.7%),对氮素分配比例无显著影响.与单作相比,间作小麦产量平均增加12.0%;AUDPC(DI)和AUDPC(DSI)分别平均降低11.5%和30.7%,间作对病情指数的控制效果优于发病率.间作显著降低发病盛期小麦氮含量、阶段累积量和叶片中氮素分配比例(降幅6.6%～12.5%、1.4%～6.9%和9.0%～15.5%).在本研究条件下,兼顾控病效果和产量效应,小麦施氮量不应超过225 kg·hm^-2.     

氮肥运筹对晚播冬小麦氮素和干物质积累与转运的影响

氮素平衡对干物质积累与分配的影响是农业生态系统研究的重要内容,在保障产量前提下减少氮肥施用量可减少环境污染与温室气体排放。以晚播冬小麦为研究对象,设置4个施氮量水平:0 kg/hm2(N0)、168.75 kg/hm2(N1)、225 kg/hm2(N2)、281.25 kg/hm2(N3),每个施氮量水平下设置2个追氮时期处理:拔节期(S1)、拔节期+开花期(S2),研究了氮肥运筹对晚播冬小麦氮素和干物质积累与转运及氮肥利用率的影响。结果表明:拔节期追施氮肥(S1)条件下,在225 kg/hm2(N2)基础上增施25%氮肥(N3)对开花期氮素积累总量和营养器官氮素转运量无显著影响;拔节期+开花期追施氮肥(S2)条件下,随施氮量增加,开花期氮素积累总量和花后营养器官氮素转运量升高;S2较S1显著提高成熟期籽粒及营养器官氮素积累量、花后籽粒氮素积累量及其对籽粒氮素积累的贡献率。同一施氮量条件下,S2较S1提高了成熟期的干物质积累量、开花至成熟阶段干物质积累强度和花后籽粒干物质积累量。同一追氮时期条件下,籽粒产量N2与N3无显著差异,氮肥偏生产力随施氮量增加而降低;同一施氮量条件下,S2较S1提高了晚播冬小麦的籽粒产量和氮肥吸收利用率。拔节期+开花期追施氮肥,总施氮量225kg/hm2为有利于实现晚播冬小麦高产和高效的最优氮肥运筹模式。