土壤水分亏缺对水稻茎秆贮藏碳水化合物向籽粒运转的调节

以两个茎秆贮藏物质利用效率不同的水稻(Oryza sativa)杂交组合(‘汕优63’和‘Pc311/早献党’)为材料,进行土壤水分亏缺处理(Water-deficit),以水层灌溉为对照(Well-watered),研究水分亏缺对水稻茎贮藏性碳水化合物运转及其关键酶活性的调节作用。结果表明,水分亏缺促进了水稻茎秆贮藏物质的运转和对籽粒产量的贡献,开花前茎秆贮藏的碳水化合物对产量贡献率分别提高了1.9~3.0倍(与水层灌溉相比)。土壤水分亏缺诱导了水稻茎节间α-淀粉酶、β-淀粉酶、α-葡萄糖苷酶、D-酶活性上升,但淀粉磷酸化酶受到了抑制,说明土壤水分亏缺加强水稻茎秆贮藏淀粉水解途径,而不是磷酸解途径。就蔗糖代谢而言,土壤水分亏缺提高了蔗糖磷酸合成酶的活性和活化状态,抑制蔗糖转化酶活性,促进蔗糖合成,加速贮藏物质快速降解和转移,从而调节稻株贮藏碳水化合物向籽粒的分配。     

Root growth and water uptake in winter wheat under deficit irrigation

Root growth is critical for crops to use soil water under water-limited conditions. A field study was conducted to investigate the effect of available soil water on root and shoot growth, and root water uptake in winter wheat (Triticum aestivum L.) under deficit irrigation in a semi-arid environment. Treatments consisted of rainfed, deficit irrigation at different developmental stages, and adequate irrigation. The rainfed plots had the lowest shoot dry weight because available soil water decreased rapidly from booting to late grain filling. For the deficit-irrigation treatments, crops that received irrigation at jointing and booting had higher shoot dry weight than those that received irrigation at anthesis and middle grain filling. Rapid root growth occurred in both rainfed and irrigated crops from floral initiation to anthesis, and maximum rooting depth occurred by booting. Root length density and dry weight decreased after anthesis. From floral initiation to booting, root length density and growth rate were higher in rainfed than in irrigated crops. However, root length density and growth rate were lower in rainfed than in irrigated crops from booting to anthesis. As a result, the difference in root length density between rainfed and irrigated treatments was small during grain filling. The root growth and water use below 1.4 m were limited by a caliche (45% CaCO₃) layer at about 1.4 m profile. The mean water uptake rate decreased as available soil water decreased. During grain filling, root water uptake was higher from the irrigated crops than from the rainfed. Irrigation from jointing to anthesis increased seasonal evapotranspiration, grain yield, harvest index and water-use efficiency based on yield (WUE), but did not affect water-use efficiency based on aboveground biomass. There was no significant difference in WUE among irrigation treatments except one-irrigation at middle grain filling. Due to a relatively deep root system in rainfed crops, the higher grain yield and WUE in irrigated crops compared to rainfed crops was not a result of rooting depth or root length density, but increased harvest index, and higher water uptake rate during grain filling.     

Physiological mechanisms contributing to the increased water-use efficiency in winter wheat under deficit irrigation

Deficit irrigation in winter wheat has been practiced in the areas with limited irrigation water resources. The objectives of this study were to (i) understand the physiological basis for determinations of grain yield and water-use efficiency in grain yield (WUE) under deficit irrigation; and (ii) investigate the effect of deficit irrigation on dry matter accumulation and remobilization of pre-anthesis carbon reserves during grain filling. A field experiment was conducted in the Southern High Plains of the USA and winter wheat (cv. TAM 202) was grown on Pullman clay loam soil (fine mixed thermic Torretic Paleustoll). Treatments consisted of rain-fed, deficit irrigation from jointing to the middle of grain filling, and full irrigation. The physiological measurements included leaf water potential, net photosynthetic rate (Pn), stomatal conductance (Gs), and leaf area index. The rain-fed treatment had the lowest seasonal evapotranspiration (ET), biomass, grain yield, harvest index (HI) and WUE as a result of moderate to severe water stress from jointing to grain filling. Irrigation application increased seasonal ET, and ET increased as irrigation frequency increased. The seasonal ET increased 20% in one-irrigation treatments between jointing and anthesis, 32-46% in two-irrigation treatments, and 67% in three- and full irrigation treatments. Plant biomass, grain yield, HI and WUE increased as the result of increased ET. The increased yield under irrigation was mainly contributed by the increased number of spikes, and seeds per square meter and per spike. Among the irrigation treatments, grain yield increased significantly but the WUE increased slightly as irrigation frequency increased. The increased WUE under deficit irrigation was contributed by increased HI. Water stress during grain filling reduced Pn and Gs, and accelerated leaf senescence. However, the water stress during grain filling induced remobilization of pre-anthesis carbon reserves to grains, and the remobilization of pre-anthesis carbon reserves significantly contributed to the increased grain yield and HI. The results of this study showed that deficit irrigation between jointing and anthesis significantly increased wheat yield and WUE through increasing both current photosynthesis and the remobilization of pre-anthesis carbon reserves.     

A proteomics view on the role of drought-induced senescence and oxidative stress defense in enhanced stem reserves remobilization in wheat

Drought is one of the major factors limiting the yield of wheat (Triticum aestivum L.) particularly during grain filling. Under terminal drought condition, remobilization of pre-stored carbohydrates in wheat stem to grain has a major contribution in yield. To determine the molecular mechanism of stem reserve utilization under drought condition, we compared stem proteome patterns of two contrasting wheat landraces (N49 and N14) under a progressive post-anthesis drought stress, during which period N49 peduncle showed remarkably higher stem reserves remobilization efficiency compared to N14. Out of 830 protein spots reproducibly detected and analyzed on two-dimensional electrophoresis gels, 135 spots showed significant changes in at least one landrace. The highest number of differentially expressed proteins was observed in landrace N49 at 20days after anthesis when active remobilization of dry matter was observed, suggesting a possible involvement of these proteins in effective stem reserve remobilization of N49. The identification of 82 of differentially expressed proteins using mass spectrometry revealed a coordinated expression of proteins involved in leaf senescence, oxidative stress defense, signal transduction, metabolisms and photosynthesis which might enable N49 to efficiently remobilized its stem reserves compared to N14. The up-regulation of several senescence-associated proteins and breakdown of photosynthetic proteins in N49 might reflect the fact that N49 increased carbon remobilization from the stem to the grains by enhancing senescence. Furthermore, the up-regulation of several oxidative stress defense proteins in N49 might suggest a more effective protection against oxidative stress during senescence in order to protect stem cells from premature cell death. Our results suggest that wheat plant might response to soil drying by efficiently remobilize assimilates from stem to grain through coordinated gene expression.     

Contribution of Pre-Anthesis Assimilates and Current Photosynthesis to Grain Yield,and their Relationships to Drought Resistance in Wheat Cultivars Grown Under Different Soil Moisture

We investigated the relative importance of pre-anthesis assimilates stored in plant parts, mainly in the stem, and post-anthesis photosynthesis to drought resistance in wheat (Triticum aestivum L.) cultivars Hongwangmai (drought resistant) and Haruhikari (drought sensitive) subjected to two soil moisture regimes: irrigated and non-irrigated. In the irrigated treatment, soil moisture was maintained near field capacity throughout the growing season, while in the non-irrigated treatment water was withheld from 81 d after sowing until maturity. Drought stress reduced grain yield of Hongwangmai and Haruhikari by 41 and 60 %, respectively. Remobilization of pre-anthesis assimilates to the grain (remobilization) was reduced by drought in Hongwangmai but increased in Haruhikari. The contribution of pre-anthesis assimilates to the grain decreased under non-irrigated treatment in Hongwangmai. However, under water stress, Hongwangmai maintained a higher net photosynthetic rate in the flag leaf than Haruhikari. These results indicated that maintenance of post-anthesis photosynthetic rate was related to drought resistance in Hongwangmai rather than to remobilization under drought stress.     

Relationship between Carbon Isotope Discrimination and Grain Yield in Spring Wheat Cultivated under Different Water Regimes

In C₃ plants, carbon isotope discrimination (Δ) has been proposed as an indirect selection criterion for grain yield. Reported correlations between Δ and grain yield however, differ highly according to the analyzed organ or tissue, the stage of sampling, and the environment and water regime. In a first experiment carried out in spring wheat during two consecutive seasons in the dry conditions of northwest Mexico (Ciudad Obregon, Sonora), different water treatments were applied, corresponding to the main water regimes available to spring wheat worldwide, and the relationships between Δ values of different organs and grain yield were examined. Under terminal (post‐anthesis) water stress, grain yield was positively associated with Δ in grain at maturity and in leaf at anthesis, confirming results previously obtained under Mediterranean environments. Under early (pre‐anthesis) water stress and residual moisture stress, the association between grain Δ and yield was weaker and highly depended on the quantity of water stored in the soil at sowing. No correlation was found between Δ and grain yield under optimal irrigation. The relationship between Δ and grain yield was also studied during two consecutive seasons in 20 bread wheat cultivars in the Ningxia region (Northern China), characterized by winter drought (pre‐anthesis water stress). Wheat was grown under rainfed conditions in two locations (Guyuan and Pengyang) and under irrigated conditions in another two (Yinchuan and Huinong). In Huinong, the crop was also exposed to salt stress. Highly significant positive associations were found between leaf and grain Δ and grain yields across the environments. The relationship between Δ and yield within environments highly depended on the quantity of water stored in the soil at sowing, the quantity and distribution of rainfall during the growth cycle, the presence of salt in the soil, and the occurrence of irrigation before anthesis. These two experiments confirmed the value of Δ as an indirect selection criterion for yield and a phenotyping tool under post‐anthesis water stress (including limited irrigation).     

Contributions of root uptake and remobilization to grain zinc accumulation in wheat depending on post-anthesis zinc availability and nitrogen nutrition

Background and aims

Whether root Zn uptake during grain filling or remobilization from pre-anthesis Zn stores contributes more to grain Zn in wheat is subject to an on-going debate. This study investigated the effects of N nutrition and post-anthesis Zn availability on the relative importance of these sources.

Methods

Durum wheat plants were grown in nutrient solution containing adequate Zn (0.5?μM) and three different N levels (0.5; 1.5; 4.5?mM). One third of the plants were harvested when they reached anthesis. One half of the remaining plants were grown to maturity with adequate Zn, whereas the Zn supply to the other half was discontinued at anthesis. Roots, straw and grains were harvested separately and analyzed for Zn and N.

Results

Depending on the N supply, Zn remobilization from pre-anthesis sources provided almost all of grain Zn when the Zn supply was withheld at anthesis; otherwise up to 100?% of grain Zn could be accounted for by Zn taken up post-anthesis. By promoting tillering and grain yield and extending the grain filling, higher N supply favored the contribution of Zn uptake to grain Zn accumulation.

Conclusion

Remobilization is critical for grain Zn accumulation when Zn availability is restricted during grain filling. However, where root uptake can continue, concurrent Zn uptake during grain development, favored by higher N supply, overshadows net remobilization.     

Relationship between Carbon Isotope Discrimination and Grain Yield in Spring Wheat Cultivated under Different Water Regimes

In C3 plants, carbon isotope discrimination (△) has been proposed as an indirect selection criterion for grain yield. Reported correlations between △ and grain yield however, differ highly according to the analyzed organ or tissue, the stage of sampling, and the environment and water regime. In a first experiment carried out in spring wheat during two consecutive seasons in the dry conditions of northwest Mexico (Ciudad Obregon, Sonora), different water treatments were applied,corresponding to the main water regimes available to spring wheat worldwide, and the relationships between △ values of different organs and grain yield were examined. Under terminal (post-anthesis) water stress, grain yield was positively associated with △ in grain at maturity and in leaf at anthesis, confirming results previously obtained under Mediterranean environments. Under early (pre-anthesis) water stress and residual moisture stress, the association between grain △ and yield was weaker and highly depended on the quantity of water stored in the soil at sowing. No correlation was found between △ and grain yield under optimal irrigation. The relationship between △ and grain yield was also studied during two consecutive seasons in 20 bread wheat cultivars in the Ningxia region (Northern China), characterized by winter drought(pre-anthesis water stress). Wheat was grown under rainfed conditions in two locations (Guyuan and Pengyang) and under irrigated conditions in another two (Yinchuan and Huinong). In Huinong, the crop was also exposed to salt stress.Highly significant positive associations were found between leaf and grain △ and grain yields across the environments.The relationship between △ and yield within environments highly depended on the quantity of water stored in the soil at sowing, the quantity and distribution of rainfall during the growth cycle, the presence of salt in the soil, and the occurrence of irrigation before anthesis. These two experiments confirmed the value of △ as an indirect selection criterion for yield and a phenotyping tool under post-anthesis water stress (including limited irrigation).     

Response of Spring Wheat (Triticum aestivum L.) Quality Traits and Yield to Sowing Date

The unpredictability and large fluctuation of the climatic conditions in rainfed regions do affect spring wheat yield and grain quality. These variations offer the opportunity for the production of better quality wheat. The effect of variable years, locations and sowing managements on wheat grain yield and quality was studied through field experiments using three genotypes, three locations for two years under rainfed conditions. The two studied years as contrasting years at three locations and sowing dates depicted variability in temperature and water stress during grain filling which resulted considerable change in grain yield and quality. Delayed sowing, years (2009–10) and location (Talagang) with high temperature and water stress resulted increased proline, and grain quality traits i.e. grain protein (GP) and grain ash (GA) than optimum conditions (during 2008–09, at Islamabad and early sowing). However, opposite trend was observed for dry gluten (DG), sodium dodecyl sulphate (SDS), SPAD content and grain yield irrespective of genotypes. The influence of variable climatic conditions was dominant in determining the quality traits and inverse relationship was observed among some quality traits and grain yield. It may be concluded that by selecting suitable locations and different sowing managements for subjecting the crop to desirable environmental conditions (temperature and water) quality traits of wheat crop could be modified.     

高温下干旱和渍水对冬小麦花后旗叶光合特性和物质转运的影响

以蛋白质含量不同的两个冬小麦品种扬麦9号和豫麦34为材料,研究了不同温度和水分条件下小麦花后旗叶光合特性的变化、营养器官花前贮藏干物质和氮素转运特征及其与籽粒产量和品质形成的关系.结果表明,高温及干旱和渍水均明显降低了旗叶光合速率和叶绿素含量（SPAD值）,但高温下干旱和渍水对光合作用的影响加重.小麦营养器官花前贮藏干物质、氮素转运量和转运率在适温下表现为干旱>对照>渍水,高温下则表现为对照>干旱>渍水.适温下花后同化物积累量表现为对照>渍水>干旱,高温下则表现为对照>干旱>渍水.花后氮素积累量在适温和高温下均表现为对照>渍水>干旱.籽粒淀粉含量以适温适宜水分处理最高,高温渍水下最低;蛋白质含量以高温干旱下最高,适温渍水下最低.温度和水分逆境下小麦粒质量和淀粉含量的降低与花后较低的光合能力及干物质积累有关,而蛋白质含量则与花前贮藏氮素的转运量和转运率有关.     

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.     

不同生育时期干旱对冬小麦氮素吸收与利用的影响

以抗旱性强的‘石家庄8号’和抗旱性弱的‘偃麦20’冬小麦(Triticum aestivum)为材料, 在田间遮雨棚条件下, 研究返青-拔节期、拔节-开花期和灌浆后期3个生育期不同干旱程度对冬小麦产量、氮素吸收、分配和利用的影响。结果表明, 在干旱条件下, 抗旱性强的‘石家庄8号’产量高于抗旱性弱的‘偃麦20’, 并且其3个生育时期轻度干旱均可提高产量。拔节-开花期干旱对两个冬小麦品种氮素的吸收和运转影响均最大, 其次为返青-拔节期, 而灌浆后期影响较小。不同生育期中度和重度干旱均降低了花前贮藏氮素向籽粒中的转移, 并且氮肥利用效率和生产率也较低, 而在返青-拔节和灌浆后期轻度干旱有利于营养器官的氮素向籽粒中转移, 提高了氮肥利用效率和生产率。在干旱条件下, 抗旱性强的‘石家庄8号’籽粒氮素积累对花前贮藏氮素再运转的依赖程度高, 而‘偃麦20’对花后氮素的积累和转移依赖较高。综合产量和氮素的转移特点, 在生产实践中, 返青-拔节期和灌浆后期要注意对小麦进行适度的干旱处理, 在拔节-开花期要保证冬小麦的充分灌溉, 从而有利于氮素的积累和分配。     

不同水分管理下优质稻花后植株碳氮流转与籽粒生长及品质的相关性

以桂华占、八桂香为材料,在干湿交替灌溉、亏缺灌溉、淹水灌溉3种水分条件下,研究优质稻花后植株碳氮流转与籽粒生长及品质的相关性。结果表明:不同水分管理下,桂华占和八桂香花后碳氮流转与籽粒的生长间存在密切相关。主要表现在:(1)茎鞘和叶片干物质转运对籽粒干物质积累的贡献率为16.86%～25.68%,花后茎叶干物质运转速度和运转率与籽粒起始灌浆势呈显著甚至极显著正相关;籽粒最大灌浆速率、活跃灌浆期、持续灌浆时间与叶片干物质运转速度和运转率呈极显著正相关,与茎鞘干物质运转速度和运转率呈极显著负相关;(2)茎鞘碳同化物转运对籽粒的产量和淀粉产量的贡献率则为干湿交替灌溉>亏缺灌溉>淹水灌溉;但叶片碳同化物转运对籽粒的产量和淀粉产量的贡献率则为淹水灌溉>亏缺灌溉>干湿交替灌溉;茎叶可溶性糖积累量的减少和籽粒直链淀粉含量和积累量增加是同步的,且茎叶可溶性糖积累量快速递减期(花后3～12d)与直链淀粉含量和积累量快速递增期(花后6～12d)同步;(3)茎鞘和叶片氮素转运对籽粒氮素积累的贡献率为44.05%～117.66%,叶片总氮转运对籽粒氮素积累的贡献率大于茎鞘,茎鞘和叶片氮同化物对籽粒氮素的贡献率以淹水灌溉处理的最大,亏缺灌溉处理的次之,干湿交替灌溉处理的最小。     

干旱调控下小麦RIL群体灌浆期茎秆可溶性碳水化合物积累与转运的遗传分析

以小麦RIL群体(陇鉴19×Q9086,F₈)120个株系及其亲本为供试材料,研究雨养(DS)和正常灌溉(WW)条件下,小麦灌浆期不同发育阶段主茎不同节位可溶性碳水化合物(WSC)含量、转运率及其对籽粒的贡献率,以及穗粒重的遗传特点及各目标性状间的相关性.结果表明: 在两种水分条件下,小麦RIL群体各目标性状变异广泛,变异系数在2.7%～62.1%(DS)和1.9%～52.1%(WW),多样性指数在0.61～0.90(DS)和0.64～0.89(WW),且存在超亲分离.各目标性状表型受基因型、水分环境、节位和发育时期的显著影响.其中,WSC含量受发育时期的影响较大,WSC转运率具有显著的水分和节位主导效应,而WSC转运对籽粒的贡献率受基因型、节位和水分的共同作用.开花初期和灌浆中期WSC含量、花前WSC转运率对籽粒的贡献率之间普遍表现为显著或极显著正相关,且干旱条件下其相关系数更高.各目标性状的遗传力较低(hB₂在干旱条件下为0.31～0.56,灌溉条件下为0.44～0.67),控制各目标性状的遗传基因对数在6～29对(DS)和3～19对(WW).表明该群体对所考察性状有贡献的等位基因在其后代群体中得到广泛分离,其表达易受水分环境的影响,符合典型数量性状特点.     

氮肥后移对抽穗后水分胁迫下冬小麦光合特性及产量的影响

采用子母桶栽土培法模拟冬小麦抽穗后不同的水分胁迫状态,研究了氮肥后移对冬小麦光合特性及产量的影响.设置3个氮肥处理,分别为N₁(基肥∶拔节肥∶开花肥=10∶0∶0)、N₂(6∶4∶0)和N₃(4∶3∶3),模拟冬小麦抽穗后2种水分胁迫(渍水胁迫、干旱胁迫),设正常供水为对照.结果表明：相同供水条件下,N₂和N₃处理较N₁处理显著提高冬小麦灌浆期旗叶的SPAD和光合速率,确保了收获时较高的穗数、穗粒数和地上部分生物量;氮肥后移处理显著提高了冬小麦的耗水量,但其籽粒产量和水分利用效率也显著提高.相同氮肥条件下,干旱胁迫和渍水胁迫处理较正常供水显著降低了冬小麦开花期和灌浆期旗叶的光合速率、千粒重、穗粒数和产量.与正常供水相比,各氮肥条件下干旱胁迫和渍水胁迫处理花后旗叶光合速率及籽粒产量的减小幅度均表现为N₁＞N₂＞N₃.表明氮肥后移通过提高旗叶SPAD、减缓花后旗叶光合速率的下降幅度、增加地上部分干物质积累量,调控产量及其构成要素,以减轻逆境灾害(干旱和渍水胁迫)对产量的影响.     

Traits and selection strategies to improve root systems and water uptake in water-limited wheat crops

Wheat yields globally will depend increasingly on good management to conserve rainfall and new varieties that use water efficiently for grain production. Here we propose an approach for developing new varieties to make better use of deep stored water. We focus on water-limited wheat production in the summer-dominant rainfall regions of India and Australia, but the approach is generally applicable to other environments and root-based constraints. Use of stored deep water is valuable because it is more predictable than variable in-season rainfall and can be measured prior to sowing. Further, this moisture is converted into grain with twice the efficiently of in-season rainfall since it is taken up later in crop growth during the grain-filling period when the roots reach deeper layers. We propose that wheat varieties with a deeper root system, a redistribution of branch root density from the surface to depth, and with greater radial hydraulic conductivity at depth would have higher yields in rainfed systems where crops rely on deep water for grain fill. Developing selection systems for mature root system traits is challenging as there are limited high-throughput phenotyping methods for roots in the field, and there is a risk that traits selected in the lab on young plants will not translate into mature root system traits in the field. We give an example of a breeding programme that combines laboratory and field phenotyping with proof of concept evaluation of the trait at the beginning of the selection programme. This would greatly enhance confidence in a high-throughput laboratory or field screen, and avoid investment in screens without yield value. This approach requires careful selection of field sites and years that allow expression of deep roots and increased yield. It also requires careful selection and crossing of germplasm to allow comparison of root expression among genotypes that are similar for other traits, especially flowering time and disease and toxicity resistances. Such a programme with field and laboratory evaluation at the outset will speed up delivery of varieties with improved root systems for higher yield.     

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.     

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。