“绿色革命”新进展:赤霉素与氮营养双重调控的表观修饰助力水稻高产高效育种

以半矮秆育种为代表的"绿色革命"极大地提高了作物产量,但也带来氮营养利用效率降低的严重问题。"绿色革命"主要基于调控赤霉素的代谢和信号转导而实现。前期的研究发现,赤霉素信号转导关键因子DELLA蛋白通过调控GRF4而负调控氮素的吸收利用,为半矮秆品系氮利用效率低的问题提供了解决方案。最近的一项研究进一步揭示了GA信号途径与氮响应交叉互作的新机制。该研究发现水稻(Oryza sativa) NGR5是氮素调控分蘖数目的一个关键基因,其表达受氮诱导。通过招募PRC2, NGR5对D14和OsSPL14等分蘖抑制基因所在位点进行H3K27me3甲基化修饰,从而抑制其表达。而在半矮秆背景下超表达NGR5可以提高低氮水平下的水稻产量。NGR5同时也被发现为赤霉素受体GID1的一个新靶标,受到其负调控。该研究发现了调控赤霉素信号通路的新机制,并对高产高效的新一代"绿色革命"育种实践具有重要启示。     

水稻分蘖氮响应调控机理研究进展

氮肥是作物产量增加最主要的驱动因素,然而氮肥滥用会造成生态环境的严重破坏。因此,提高作物氮素利用效率(nitrogen use efficiency,NUE)对未来农业可持续发展至关重要。产量性状对氮素的敏感性是衡量作物氮素利用效率的重要指标。禾本科作物的分蘖数、穗粒数和粒重是产量的直接决定因子,虽然影响三者本身的分子机制已有大量研究,但氮素对这些性状的调控机理仍知之甚少。分蘖数是对氮素响应最为敏感的性状之一,也是氮肥促进作物增产的关键要素。因此,研究氮素如何调控水稻的分蘖发育对于提高作物产量尤为重要。本文总结了水稻氮素利用效率的影响因素和分蘖发育的调控机理,聚焦氮素如何调控水稻分蘖发育的机制,并对该领域未来研究工作进行了展望,以期为作物氮高效精准改良提供参考。     

综述与专论：表观遗传修饰调控阿尔茨海默病的研究进展

阿尔茨海默病（Alzheimer’s disease,AD）是一种临床上常见的以进行性认知功能障碍和记忆减退为主要特征的神经退行性疾病。近些年研究发现，表观遗传修饰如DNA修饰、组蛋白修饰、RNA修饰及非编码RNA在Aβ沉积、Tau蛋白过度磷酸化、神经再生、突触可塑性和认知功能中发挥不同程度的调控作用，进而改善或加剧AD病理进程。临床数据表明表观遗传修饰的改变与AD风险呈显著相关性，运用药物、物理刺激、si RNA等干预手段在AD动物模型中改变表观遗传修饰水平可改善AD病理和认知能力。本文综述了不同的表观遗传修饰在AD中的调控作用，为进一步理解AD的表观遗传学机制及通过干预表观遗传修饰改善或治疗AD的可行性提供理论依据。     

赤霉素和油菜素内酯信号通路双重调控助力小麦新一轮“绿色革命”

自20世纪60年代以来,半矮秆基因Rht-B1b和Rht-D1b的利用显著提高了小麦(Triticum aestivum)抗倒伏能力和收获指数,使得全世界小麦产量翻了一番,引发了农业第1次“绿色革命”。Rht-B1b和Rht-D1b编码植物生长抑制因子DELLA蛋白,是赤霉素(GA)信号转导途径的负调控因子。DELLA蛋白积累抑制细胞分裂和细胞伸长,导致矮化表型;同时也抑制光合作用并降低氮素利用效率,导致半矮化品种需要较高的化肥投入才能获得高产。如何“减肥增效”是实现低碳绿色农业所面临的重大问题。最近,中国农业大学倪中福团队发现了具有育种应用价值的新型“半矮秆”基因模块,证明通过对赤霉素和油菜素内酯(BR)信号通路的双重调控可实现矮秆高产小麦新品种培育。该团队鉴定并克隆了1个控制小麦株高和粒重的数量性状位点(QTL),该QTL在衡597中存在1个约500 kb的r-e-z大片段缺失,其中包括Rht-B1b基因和1个编码RING E3泛素连接酶的ZnF-B基因。研究发现,ZnF-B蛋白与油菜素内酯信号转导途径的抑制因子TaBKI1相互作用,诱导TaBKI1降解,从而促进BR信号转导。Zn...     

直播旱作水稻的吸氮特征与土壤氮素表观盈亏

水稻旱作是水稻节水栽培中最有效的方式。通过田间试验研究旱作直播条件下水稻对氮素的吸收利用特征以及土壤矿质氮的动态变化 ,并对土壤氮素的表观盈亏量进行了估算。结果表明 ,直播旱作水稻较水作水稻更注重中后期对氮素养分的吸收 ,尤其是对土壤氮素的吸收 ;幼穗分化后水稻的土壤吸氮量占阶段吸氮总量的 6 9.5 % ,比水作水稻多 17.8%。对 0～ 4 0 cm土层土壤矿质氮含量时空变化的研究表明 ,直播旱作水稻生育前期土壤表层矿质态氮大量累积 ,在灌水和降雨的影响下 ,向下层的迁移增加 ,基肥施用后裸地处理 2 0～ 4 0 cm土层的矿质氮高达 10 4 kg N/hm2 。对水稻各生育期土壤氮素盈亏的计算结果表明 ,自分蘖盛期后旱作各处理都表现出土壤氮素不同程度上的表观亏缺 ,然而就全生育期土壤氮素盈余量而言 ,旱作处理平均高达 12 7kg N/hm2 ,生育前期氮肥的大量投入是氮素盈余的主要原因。本试验结果表明 ,直播旱作水稻生育前期对施用的肥料氮吸收很少 ,提高直播旱作水稻氮肥利用效率的关键在于减少生育前期肥料氮的投入     

植物适应土壤氮素环境的基因选择:以水稻为例

在农业生产过程中,施用无机氮肥是提高作物产量的基础,但氮肥过量施加对生态系统和植物发育进程均会产生不利影响。因此,提高作物氮素利用效率是农业可持续发展的关键。目前,对重要粮食作物水稻(Oryza sativa)的氮高效研究取得了一系列重要进展,并克隆到多个调控氮素吸收、转运和代谢的关键基因。然而,在不断被人工选育的过程...     

氮高效水稻种质资源筛选及相关特性分析

在不施氮和施氮量180 kg/hm22个水平下,对45份水稻种质的氮吸收与利用等相关性状进行分析,并根据参试品种在2个施氮水平下的产量差异将其分为双高效型、高氮高效型、低氮高效型和双低效型等4种类型,重点研究不同氮效率水稻产量、吸氮量以及氮素利用率等性状的相关性及差异分析。结果表明,单株子粒重、单株秸秆重、植株吸氮量和氮生理利用率变幅分别为4.50~29.65 g、6.35~18.70 g、4.80~21.28 g/m2和19.88~62.05 g/g。无论是在低氮还是高氮水平下,水稻产量与有效穗数、穗粒数、结实率、生物量、子粒吸氮量、吸氮量和氮生理利用率均呈显著或极显著正相关,与子粒和秸秆氮浓度呈显著负相关。4种氮效率类型品种间的产量与生物量、吸氮量和氮生理利用率差异均达到极显著水平,双高效型和高氮高效型水稻受氮肥的影响最大,低氮高效型水稻受氮肥的影响最小。研究认为,青马早和陆财早不论是在低氮还是高氮水平下均表现出氮高效利用特性,为典型氮高效型;广陆矮4号在低氮水平下表现出氮高效利用特性,适于低氮条件种植,为典型耐低氮型;早89-01和早籼152在高氮水平下表现出氮高效利用特性,适于高氮条件种植。     

水稻高光效育种研究进展

水稻是世界重要的粮食作物,其较低的光合效率是限制水稻产量的重要因素之一,提高水稻的光能利用率对于进一步提高水稻产量具有关键的作用,本文简要回顾国内外水稻高光效育种的发展历程和研究进展,并对杂交选育、基因工程技术、株型改良等水稻高光效育种研究途径进行了总结与展望。     

Nitrogen Mediates Flowering Time and Nitrogen Use Efficiency via Floral Regulators in Rice

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Effect of Nitrogen Supply on the Nitrogen Use Efficiency of an Annual Herb, Helianthus annuus L.

Nitrogen use efficiency (NUE) is the product of nitrogen productivity (NP) and the mean residence time of nitrogen (MRT). Theory suggests that there should be a trade-off between both components,but direct experimental evidence is still scarce. To test this hypothesis, we analyzed the effect of varying nitrogen supply levels on NUEand its two components (NP, MRT) in Helianthus annuus L., an annual herb.The plants investigated were subjected to six nitrogen levels (0, 2, 4, 8, 16, and 32 g N/m2). Total plant production increased substantially with increasing nitrogen supply. Nitrogen uptake and loss also in creased with nitrogen supply. Nitrogen influx (rin) and outflux (rout) were defined as the rates of nitrogen uptake and loss per unit aboveground nitrogen, respectively. Both rin and rout increased with increasing nitrogen supply. In addition, rin was far higher than rout. Consequently, the relative rate of nitrogen incre ment (rin- rout) also increased with nitrogen supply. There were marked differences between treatments with respect to parameters related to the stress resistance syndrome: nitrogen pool size, leaf nitrogen concentration,and net aboveground productivity increased with nitrogen supply. Plants at high nitrogen levels showed a higher NP (the growth rate per unit aboveground nitrogen) and a shorter MRT (the inverse of rout), whereas plants at low nitrogen levels displayed the reverse pattern. Shorter MRT for plants at high nitrogen levels was caused by the abscission of leaves that contained relatively large fractions of total plant nitrogen. We found a negative relationship between NP and MRT, the components of NUE, along the gradient of nitrogen availability, suggesting that there was a trade-off between NP and MRT. The NUE increased with increasing nitrogen availability, up to a certain level, and then decreased. These results offer support for the hypoth esis that adaptation to infertile habitats involves a low nitrogen loss (long MRT in the plant) rather than a high NUE per se. The higher NUE at the plant level was a result, in part, of greater nitrogen resorption during senescence. We suggest that a long MRT (an index of nitrogen conservation) is a potentially successful strategy in nitrogen-poor environments.     

Effects of Irrigation and Nitrogen on the Performance of Aerobic Rice in Northern China

Aerobic rice is a new production system in which specially-developed varieties are grown under non-flooded, non-puddled, and non-saturated soil conditions. In 2003-2004, irrigation x Nitrogen experiments were carried out near Beijing using variety HD297. Water treatments included four irrigation levels, and Nitrogen treatments included different fertilizer N application rates and different numbers of N splits. The highest yields were 4460 kg/ha with 688 mm of total (rain plus irrigation) water input in 2003 and 6 026 kg/ha with 705 mm of water input in 2004. Because of the quite even distribution of rainfall in both years, the four irrigation treatments did not result in large differences of soil water conditions. There were few significant effects of irrigation on biomass accumulation, but yield increased with the total amount of water applied. High yields coincided with high harvest index and high percentages of grain filling. The application of fertilizer N either reduced biomass and yield or kept it at the same level as 0 N and consistently reduced the percentage grain filling and 1 000-grain weight. With the highest water application, five splits of N gave higher yield than three splits, whereas three splits gave higher yield than five splits with lower water applications.     

Nitrogen Use Efficiency of Crop Plants: Physiological Constraints upon Nitrogen Absorption

Current global nitrogen fertilizer use has reached approximately one hundred billion kg per annum. In many agricultural systems, a very substantial portion of this applied nitrogen fertilizer is lost from soil to groundwaters, rivers and oceans. While soil physicochemical properties play a significant part in these losses, there are several characteristic features of plant nitrogen transporter function that facilitate N losses. Nitrate and ammonium efflux from roots result in a reduction of net nitrogen uptake. As external nitrate and ammonium concentrations, respectively, are increased, particularly into the range of concentrations that are typical of agricultural soils, elevated rates of nitrate and ammonium efflux result. The rapid down-regulation of high-affinity influx as plants become nitrogen replete further reduces the root's capacity to acquire external nitrogen; only nitrogen-starved roots absorb with both high capacity and high affinity. The results of studies using molecular biology methods demonstrate that genes encoding nitrate and ammonium transporters are rapidly down-regulated when nitrogen is resupplied to nitrogen-starved plants. Provision of ammonium to roots of plants actively absorbing nitrate imposes a block on nitrate uptake, the extent of which depends on the ammonium concentration, thus further reducing the efficient utilization of soil nitrate. During the daily variation of incoming light and during periods of low incident irradiation (i.e. heavy cloud cover) the expression levels of genes encoding nitrate and ammonium transporters, and rates of nitrate and ammonium uptake, are substantially reduced. Low temperatures reduce growth and nitrogen demand, and appear to discriminate against high-affinity nitrogen influx. In sum, these several factors conspire to limit rates of plant nitrogen uptake to values that are well below capacity. These characteristics of the plant's nitrogen uptake systems facilitate nitrogen losses from soils.     

Adjusting Nitrogen Application in Accordance with Soil Water Availability Enhances Yield and Water Use by Regulating Physiological Traits of Maize under Drip Fertigation

Knowledge of the interactive effects of water and nitrogen (N) on physio-chemical traits of maize (Zea mays L.) helps to optimize water and N management and improve productivity. A split-plot experiment was conducted with three soil water conditions (severe drought, moderate drought, and fully water supply referring to 45%–55%, 65%–75%, and 85%–95% field capacity, respectively) and four N application rates (N₀, N₁₅₀, N₂₄₀, and N₃₃₀ referring to 0, 150, 240, 330 kg N ha^–1 respectively) under drip fertigation in 2014 and 2015 in the Huang-Huai-Hai Plain of China. The results indicated that drought stress inhibited physiological activity of plants (leaf relative water content, root bleeding sap, and net photosynthetic rate), resulting in low dry matter accumulation after silking, yield, and N uptake, whereas increased WUE and NUE. N application rates over than 150 kg ha^–1 aggravated the inhibition of physiological activity under severe drought condition, while it was offset under moderate drought condition. High N application rates (N₃₃₀) still revealed negative effects under moderate drought condition, as it did not consistently enhance plant physiological activity and significantly reduced N uptake as compared to the N₂₄₀ treatment. With fully water supply, increasing N application rates synergistically enhanced physiological activity, promoted dry matter accumulation after silking, and increased yield, WUE, and N uptake. Although the N₂₄₀ treatment reduced yield by 5.4% in average, it saved 27.3% N under full water supply condition as compared with N₃₃₀ treatment. The results indicated that N regulated growth of maize in aspects of physiological traits, dry matter accumulation, and yield as well as water and N use was depended on soil water status. The appropriate N application rates for maize production was 150 kg ha^–1 under moderate drought or 240 kg ha^–1 under fully water supply under drip fertigation, and high N supply (>150 kg ha^–1) should be avoided under severe drought condition.     

Effect of Reaction-Finished Solution of Hydrochar (HRFS) Application on Rice Grain Yield and Nitrogen Use Efficiency in Saline Soil

We conducted a pot experiment to examine the feasibility of applying a reaction-finished solution of hydrochar (HRFS) to enhance rice production in a saline soil. With this purpose, HRFS was applied (0, 10, 20, 40, 60, 80 and 100 mL/pot) and rice yield and nitrogen (N) use efficiency (NUE) were determined. HRFS application significantly (P < 0.05) increased rice grain yield by 19.6%–30.0% compared to the control treatment (CKU, with N but without HRFS addition). Moreover, HRFS application promoted plant height and straw biomass of rice. Increases of rice yield were mainly achieved by increases in the number of panicles and grains per panicle. Compared with the CKU treatment, the NUE of HRFS amendments significantly (P < 0.05) increased by 56.3%–71.7%. This indicated that the improvement of NUE was one of the mechanisms to improve rice grain yield with HRFS amendment. The results of regression analysis showed that there was a positive relationship (R²=0.8332) between rice yield and HRFS application rate within an appropriate range. The highest rice yield was recorded with the HRFS application of 40 mL/pot, but a further increase in HRFS application rate appeared to reduce rice yield. Based on the results of this pot study, HRFS application can increase rice yield in a saline soil by regulating its yield components and enhancing NUE. However, impact of HRFS on these variables showed a “dose effect”.     

翻压山黧豆绿肥与氮肥减施对水稻生长及其养分吸收与产量的影响

为探明种植翻压山黧豆绿肥与减施氮肥下的水稻生产潜力,通过3年田间定位试验,设置冬闲+不施肥(NF)、山黧豆绿肥(GM)、冬闲+常规氮肥(100%N,CK)、山黧豆绿肥+80%常规氮肥(GM+80%N)、山黧豆绿肥+70%常规氮肥(GM+70%N)、山黧豆绿肥+60%常规氮肥(GM+60%N)6个处理,研究不同处理对水稻生长、养分吸收及产量的影响。结果表明：(1)与CK相比,翻压山黧豆绿肥并减施氮肥处理均能够显著提升水稻株高、增加水稻分蘖数、提高水稻干物质积累量,其中以GM+70%N施肥处理提升效果最为明显。(2)GM+70%N施肥处理下,不同生育时期水稻株高、有效分蘖数分别较对照常规施肥(100%N)提升了13.32%~ 15.73%和33.98%~59.47%,水稻干物质积累量提高了23.19%~144.18%,且随着生育时期的推进增加速率依次降低。(3)种植翻压山黧豆绿肥并减施氮肥处理下水稻产量均有所提高,其中GM+70%N和GM+80%N处理显著提高,增产分别达13.84%,7.25%,且GM+70%N处理下水稻植株和籽粒养分吸收更为全面。研究发现,种植翻压山黧豆并适量减施氮肥能有效促进水稻生长和养分的吸收积累,显著提高水稻产量,说明翻压山黧豆绿肥可替代稻田30%~40%的氮肥施入量,并可在避免水稻旺长的同时实现水稻高产,是四川水稻种植较好的耕作措施。