水分与氮素及其互作调控小麦产量和水氮利用效率研究进展
Research Advances in the Effects of Water and Nitrogen and Their Interaction on the Grain Yield, Water and Nitrogen Use Efficiencies of Wheat
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收稿日期: 2023-03-28 修回日期: 2023-05-4
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Received: 2023-03-28 Revised: 2023-05-4
作者简介 About authors
刘颖,主要从事小麦高产高效栽培理论与技术研究,E-mail:
理解土壤水分与氮素及其互作对小麦产量与水氮利用效率的影响,对协同提高小麦产量和水氮利用效率有重要意义。本文综述了小麦节水灌溉技术、氮肥施用技术、水分和氮素对小麦产量与水氮利用效率的互作效应、作物―土壤关系及水氮调控机制等方面的进展。讨论了目前存在的问题,即高产小麦与土壤的水氮互作效应尚不明确,小麦水氮耦合与高效利用的分子机理尚不清楚以及协同提高小麦产量和水氮利用效率的调控途径与关键技术尚未掌握。针对上述问题,今后应重点探究高产小麦与土壤的水氮互作效应与机制、水氮互作调控小麦吸收利用水分和氮素的生理与分子机理,以及协同提高小麦产量和水氮利用效率的调控途径与关键技术。
关键词:
Understanding the effects of water, nitrogen and their interaction on grain yield, water and nitrogen use efficiency is of great importance for synergistic improvement of grain yield, water and nitrogen use efficiency of wheat. This review focused on progress in water-saving irrigation technologies, nitrogen application technologies, interaction effects of water and nitrogen on grain yield, water and nitrogen use efficiency, root-soil relationship and its mechanism regulated by water and nitrogen. Some key problems were discussed: the synergistic interaction between water and nitrogen of soil and yield is still unclear; the molecular mechanism underlying the interaction between water and nitrogen on the efficient uptake and use of water and nitrogen in high-yielding wheat is still unclear; and the regulatory approach of synergistic increases in grain yield and water-nitrogen use efficiency has not been mastered. In view of the above problems, future research should focus on exploring the effect and mechanism of water nitrogen interaction between high-yield wheat and soil, and the physiological and molecular mechanism of water nitrogen interaction regulating water and nitrogen absorption in wheat; and the key regulatory approaches and technologies for synergetic increase of grain yield and water and nitrogen use efficiency in wheat.
Keywords:
本文引用格式
刘颖, 顾昀怿, 张伟杨, 杨建昌.
Liu Ying, Gu Yunyi, Zhang Weiyang, Yang Jianchang.
小麦(Triticum aestivum L.)作为全球最重要的粮食作物之一,提供世界人口消耗热量的20%以上[1]。伴随人口持续增长、城市化和工业化进程不断推进,人们对粮食的需求也在进一步增加。而冬小麦在正常生长情况下生育期耗水量超过400mm,我国北方地区同期降水量远不能满足其生长水分需求。此外,根据联合国粮农组织统计,我国禾谷类作物氮肥用量比欧盟平均高出12.0%,但我国小麦单产(5.2t/hm2)与欧盟平均产量(5.8t/hm2)相比并无显著增加[2]。因此,提高有限资源的投入产出比,实现作物产量、水分与氮素利用效率协同提高是当前面临的一个重大科学问题。在不低估植物遗传作用的情况下,水和氮的有效管理被认为是缩小主要谷类作物产量差距的关键因素[3]。已有研究[4]发现,水分和氮素对作物产量的影响在数量和时间上存在最佳的匹配或耦合,只要水分和氮肥供应合理耦合,就会产生相互促进机制,得到大于水氮2种因子效果叠加的增产作用(即水氮耦合效应),使产量及水氮利用效率协同提高。对于如何同时实现小麦高产和高水氮利用率,国内外学者进行了大量研究[5-6]。本文重点阐述了小麦节水灌溉和氮肥施用技术,探究了水氮互作对小麦产量及其水氮利用效率的影响、根系―土壤关系及其调控机制,对研究过程中发现的问题进行讨论,并对下一步研究和应用进行了展望。
1 小麦节水灌溉技术
1.1 节水灌溉技术简介
1.1.1 喷灌
喷灌是以类似降雨的方式湿润土壤,补充作物所需要的水分,通过影响一些水分参数(水分利用效率、耗水量和耗水特性系数)提高用水效率,减少用水消耗[9-10]。喷灌能够根据作物需水规律进行精确供水,且喷灌下小麦耗水系数(生产0.5kg籽粒所消耗的水量)只相当于畦灌(入畦流量为3~6L/m)耗水系数的25%~30%[9,11]。因此,与畦灌相比,喷灌的效率和精确度更高。在越冬期与拔节期适当喷灌,有助于小麦穗器官的发育,在灌浆前期喷灌处理能增强冬小麦强、弱势粒的起始生长势,缩短小麦达到最大灌浆速率的时间,在灌浆中期和后期,喷灌处理能增加最大灌浆速率、平均灌浆速率及粒重,以提高小麦产量[12⇓-14]。不足在于喷灌需要大量管道设备,且在基建上投资较多,因此需要充分考虑种植作物的经济效益是否能够承担此项花费。
1.1.2 滴灌
1.1.3 渗灌
1.2 亏缺灌溉和部分根区灌溉
1.3 覆膜垄沟复合测墒补灌
覆膜垄沟复合测墒补灌,即覆膜垄沟种植与测墒补灌相结合的灌溉模式[23]。其中覆膜垄沟种植借助起垄、垄上覆膜等措施把地膜覆盖栽培技术与垄沟种植技术有机结合起来,通过起垄覆盖地膜来拦截垄面上的自然降水,汇集起来形成径流,利用水往低处流的特性将垄面部位的降水导流入相邻低凹的沟内种植区,将有限的降水集中在植物的种植沟和生根区,可显著提高土壤含水量,促进植物生长,并提高水分利用效率[23]。测墒补灌是根据目标土壤的相对含水量,计算灌溉量来补充作物关键生育期的可用水量,显著促进中土层土壤持水能力的利用,实现高产和节水[24]。然而,仅应用测墒补灌方式下的冬小麦较易受到倒春寒的危害。因此,一般采用覆膜垄沟复合补灌,以缓解倒春寒的危害,提高小麦的抗逆性和稳产性,保证小麦产量、品质和水肥利用率的协同提高[24-25]。
2 小麦氮肥施用技术
2.1 前氮后移施氮技术
2.2 包膜控释氮肥施用技术
包膜控释尿素指在传统尿素颗粒表面包覆一层疏水材料,减缓肥料颗粒中养分进入土壤的速率[24]。以前有采用硫元素作为尿素包衣,但如果硫元素被过量施入土壤,会严重破坏土壤原有菌群微生物平衡,同时加速土壤酸化[31-32]。近期研究[33⇓-35]表明,天然橡胶、生物炭和树脂等有机材料被广泛用于控释尿素的薄膜制作。采用这些作为肥料涂层对环境污染小,能源浪费少,被称为环境友好型肥料。其中天然橡胶是一种综合性能优越的可再生资源和绿色资源,天然橡胶与生化抑制剂联合包膜控释尿素能够有效抑制土壤中脲酶活性,延缓尿素水解,降低土壤NH3挥发速率,维持适宜的土壤无机氮浓度,减少氮素损失,提高土壤供氮能力,弥补单独天然橡胶包膜控释肥养分释放过快的缺点,并达到增产的效果。生物炭具有孔隙率高、比表面积大、表面电荷、官能团多样等特性,可以有效提高土壤的持水能力、吸附无机氮、增加微生物量、改变土壤细菌群落结构以及影响土壤中氮的硝化和反硝化作用[36]。在相同氮肥水平下,施用生物炭包膜尿素的土壤中总氮和铵态氮的累积淋失量均低于施常规尿素的土壤,氮肥损失比施用常规尿素低约33%,且可以在不影响作物产量的前提下,显著提高氮素利用率[37]。
2.3 有机肥和控释氮肥配施
将化肥与有机肥结合施用,可减少单一化肥使用对环境和经济的负面影响,是可持续农业基本战略之一[38]。相对于化肥,有机肥制备具有所用原料丰富和成本低的优点。应用有机肥可以缓解土壤退化,改善土壤质量,提高土壤养分和有机质水平,减少土壤酸化[39-40],但有机肥养分含量低,释放速率慢,当季养分释放不完全,不利于作物前期对养分的吸收,最终影响产量。而控释尿素能够弥补有机肥肥效不长的缺陷,使肥料养分释放时间、强度与作物养分吸收规律相吻合[33]。并能有效减少氮肥的挥发、淋溶及固定,提高肥料利用效率[41]。但控释肥料价格相对高,且全部施用控释氮肥易因生长前期养分供应不足导致脱肥[42]。因此,生产上一般采用控释氮肥和有机肥配施的方法,其施用比例可根据不同作物、土壤和施肥方式等条件进行合理调整,通过调节养分释放时间使肥料缓慢而稳定地释放到土壤中[43]。
3 小麦根系―土壤关系及其水氮调控机制
土壤中氮素营养与水分状况对土壤微生物群落和土壤酶活性有重要的影响[43⇓-45]。施氮后土壤群落结构发生变化,细菌多样性下降,土壤水分状况通过影响土壤养分的运输、基质有效性和土壤性质来改变土壤微生物群落的组成和活性[46-47]。当土壤含水量为13%~14%时,可能会造成土壤中大量硝酸盐被溶解,导致土壤pH降低,细菌群落多样性指数增加。但还有研究[48]认为,土壤含水量增多会降低土壤的氧含量,使土壤pH升高。酶是土壤中各种生化反应的催化剂,参与养分的循环与转化,其活性高低可以反映土壤肥力的好坏[49-50]。氮肥通过改善土壤理化性质、微生物活性和作物根系环境而影响土壤酶的活性,土壤氮肥增加会引起碳循环有关的水解酶活性的提高[51]。而土壤水分状况主要通过对微生物的调节以及水分造成的土壤厌氧环境来影响土壤酶的活性,若土壤水分降低,土壤蛋白酶的活性也随之下降[52]。
综上所述,根系对于小麦的养分和水分吸收以及生长和最终产量至关重要,水氮互作通过改变土壤环境,引起根系形态和生理变化,进而影响水氮利用效率。
4 水氮耦合对小麦产量与水、肥、氮利用效率的影响及其机制
4.1 农艺原因
小麦产量构成因素包括单位面积穗数、穗粒数和千粒重,这3个因素与水氮供应状况密切相关[62-63]。小麦籽粒灌浆物质主要来源于以下2部分,1/3来源于花前储存在营养器官中物质的再分配,2/3来自开花后的光合产物[64]。拔节期至孕穗期是小麦水分敏感期,这段时间内合理灌溉有利于提高小麦的分蘖成穗率和穗粒数,而此时段不同程度的干旱都会使其分蘖成穗率和结实率降低。随着施氮量的增加,小麦株高、叶面积指数、干物质积累量、产量及其构成因素均呈现先增后减的趋势[65]。过量施氮条件下,小麦分蘖数和成穗数增加,但往往会造成群体过大,影响穗部发育,限制穗粒数增加;施氮过多还易造成小麦灌浆期出现“贪青徒长”,营养器官非正常衰亡,使同化物往籽粒中的转运效率变差,不利于籽粒灌浆充实,导致粒重降低,产量下降[66]。
通常小麦产量在一定范围内随氮肥的增加而增加,但在水分不足的条件下施用过量的肥料可能会导致产量降低。合理施氮会增加土壤储水量,通过提高水分利用率来提高产量[67]。在土壤含水量低的情况下,增施氮肥可以促进小麦根系对土壤深层水分的利用,从而有效提高小麦抗旱性,促进小麦生长和干物质积累,缓解水分亏缺造成的产量降低,达到“以肥调水”的效果[68-69]。可见,水分和氮素供应在对冬小麦产量的影响存在明显交互作用。合理的水肥措施可最大限度地发挥水肥耦合优势在农业生产中的作用,有助于在保证水氮利用效率的情况下提高作物产量[70]。适宜范畴内的水氮配施能有效提高小麦库容量,使源库间的物质运输畅通,但水氮过量会造成源库关系失调。并根据实际生产条件调整不同地区适宜的水分和氮肥阈值。例如,在山东地区,灌溉量为359.8~428.9mm,施氮量为225.4~280.9kg/hm2的水氮耦合模式可保证小麦高产和高经济效益的同时提高水氮利用效率[71]。灌溉定额为390~405mm,施氮量为179~248kg/hm2是干旱地区多砾石砂土的土壤条件下的最佳水肥组合[72]。
4.2 生理机制
光合能力是决定小麦花后产量的主要因素,光合能力的提升与灌溉量和施氮量密切相关[73-74]。合理灌溉能够改善作物的光合性能,提高光合速率,促进氮素在小麦植株中的积累、调配和运转,进而提高小麦产量[75]。分蘖或拔节期合理灌溉能够使小麦叶片早发以及分蘖提前生根,利于小麦植株对水分的吸收利用并进一步提高光合能力;花后充足灌溉能够提升小麦有效绿叶面积,提高光能截获率,向籽粒提供更多的光合产物,从而达到增产的目的[76]。土壤中的氮素主要以有机态氮素和无机态氮素2种形式存在。无机态氮素是植物从土壤中吸收氮素的主要来源,即铵态氮和硝态氮,小麦主要通过硝态氮转运蛋白从土壤中获取硝态氮,在开花后期没有氮素供应限制下,通过硝态氮转运蛋白将硝态氮从根部转运至茎秆或叶片(NRT1.5和NRT1.8是2种重要的硝态氮转运蛋白)。中度干旱胁迫下增施氮肥,能够提高土壤中硝态氮含量,再经过硝酸还原酶和亚硝酸还原酶反应将硝态氮还原成NH4+,被植株进一步吸收利用。而NH4+则通过谷氨酸合成酶(GOGAT)和谷氨酰胺合成酶(GS)循环直接转化为氨基酸,氨基酸通过提高叶绿素含量和叶片光合性能减轻水分亏缺对光系统Ⅱ造成的光损伤以及因干旱胁迫对小麦产量的影响(图1b)。在干旱胁迫下,若氮素营养不良,会改变植株体内激素平衡,尤其产生较多的脱落酸引起气孔关闭,降低光合速率[77-78]。
图1
图1
小麦光合作用与氮素吸收利用模式图
(a) ATP:腺嘌呤核苷三磷酸;ADP:腺苷二磷酸;Pi:磷酸根;[H]:还原氢;NADP+:烟酰胺腺嘌呤二核苷磷酸(氧化态);NADPH:还原型烟酰胺腺嘌呤二核苷酸磷酸;Rubisco:核酮糖-1,5-二磷酸羧化酶/加氧酶;RUBP:1,5-二磷酸核酮糖;PGA:3-磷酸甘油酸;PGP:磷酸甘油磷酸酶;GOX:乙醇酸氧化酶;AGT:丙氨酸乙醛酸转氨酶;GGT:乙醛酸氨基酸转移酶;GDC:甘氨酸脱羧酶;HPR1:过氧体羟基丙酮酸还原酶;NADH:烟酰胺腺嘌呤二核苷酸(还原态);GLYK:甘油酸3-激酶。(b) NR:硝酸还原酶;NiR:亚硝酸还原酶;GOGAT:谷氨酸合成酶;GS:谷氨酰胺合成酶;AS:天冬酰胺合成酶。
Fig.1
Model of photosynthesis and nitrogen uptake and utilization in wheat
(a) ATP: adenosine triphosphate; ADP: adenosine diphosphate; Pi: phosphate; [H]: reduced hydrogen; NADP+: nicotinamide adenine dinucleotide phosphate (oxidized); NADPH: reduced nicotinamide adenine dinucleotide phosphate; Rubisco: ribulose-1,5-bisphosphate carboxylase/oxygenase; RUBP: ribulose- 1,5-diphosphate; PGA: 3-phosphoglycerate; PGP: phosphoglycerate phosphatase; GOX: glycolate oxidase; AGT: alanine glyoxylate transaminase; GGT: glyoxylate amino acid transferase; GDC: glycine decarboxylase; HPR1: peroxisome hydroxypyruvate reductase; NADH: nicotinamide adenine dinucleotide (reduced state); GLYK: glyceric acid 3-kinase. (b) NR: nitrate reductase; NiR: nitrite reductase; GOGAT: glutamate synthase; GS: glutamine synthetase; AS: asparaginase.
小麦对氮和水分的需要量是密切相关的。由于氮吸收依赖于土壤中可溶性硝酸盐的质量与流量,因此,氮的吸收可以说是由水氮施用量决定的。适度水分亏缺与缺氮刺激光呼吸,其中缺氮促进了光呼吸中氮循环,使大量的腺嘌呤核甘三磷酸(ATP)被消耗,降低供碳还原的ATP与还原型烟酰胺腺嘌呤二核苷酸磷酸(NADPH)的比值,导致光合作用削弱;水分亏缺使光呼吸酶对光合电子传递和CO2固定速率的控制作用增强[79-80]。在轻度到中度干旱胁迫下(叶片相对含水量下降到70%),光合作用受到限制主要因为气孔关闭而引起的细胞间CO2浓度降低,当发生严重干旱胁迫时,叶片无机阴离子浓度增加以及Rubisco等光合酶活性降低会导致叶片光合能力下降[57,81]。因此,只有水分和氮素的合理搭配才能有助于提高光能转换过程中的关键酶活性并在花后保持较高的活性水平,使光合能力得到显著提升,从而达到增产的目的[64](图1a)。
4.3 遗传机制
硝态氮和铵态氮是植物吸收和利用的无机氮源,其中硝态氮是小麦吸收利用的主要氮素形态[83]。硝态氮需要硝酸盐转运蛋白硝酸盐转运体1家族(NRT1)和硝酸盐转运体2家族(NRT2)的转运[84]。其中在小麦生长发育过程中,NRT1.5负责将硝态氮运出中柱鞘细胞,装载到木质部,转运至地上部分;NRT1.8主要在根的木质部薄壁细胞中表达,可以把硝态氮从木质部输出[85-86]。然后通过硝酸还原酶和亚硝酸还原酶将硝态氮还原为铵,当植物组织的铵浓度积累到一个较高的水平时,会破坏跨膜质子梯度,抑制呼吸链的电子传递,此时植物会将由硝酸盐同化形成的铵或呼吸产生的铵通过谷氨酰胺―谷氨酸循环同化为氨基酸以减轻对植物造成的伤害[87](图1b)。
水通道蛋白是一种位于细胞膜上的蛋白质,在细胞膜上组成“孔道”,控制水在细胞的进出。氮素可通过调控水通道蛋白基因表达进而影响小麦对于土壤中水分的吸收与利用。目前,关于小麦中水通道蛋白对氮的响应的研究报道很少。水稻根特异性水通道蛋白基因OsPIP1.1、OsPIP2.3-2.5、OsTIP1.1-1.2和OsTIP2.2的表达与氮的可吸收利用性呈正相关。相反,氮缺乏导致水通道蛋白基因表达水平降低,削弱了根系导水率[88]。与此一致,破坏NRT2.1硝酸盐转运蛋白也会对水通道蛋白基因OsPIP1.1、OsPIP1.2、OsPIP2.1、OsPIP2.3和OsPIP2.7的表达水平产生负面影响,导致根系导水率降低[89]。此外,蛋白质组学分析[90]表明,氮的有效性也会影响植物根系中PIP水通道蛋白的水平及其磷酸化状态,进而影响根系的导水能力。这些研究为人们对小麦根系吸收水分和氮素间互作的遗传机理提供了新的参考,小麦的根系氮转运蛋白与水通道蛋白之间的互作关系及相关机制值得深入探究。
5 存在问题与展望
5.1 存在问题
虽然目前对提高小麦产量以及水氮利用效率已经进行了大量研究并取得了重要进展,但仍存在许多问题。
5.1.1 水氮调控作物与土壤的互作效应以及作用机制尚不明确
在不低估植物遗传学作用的情况下,水氮的有效管理已被确定为缩小主要谷类作物产量差距的关键。以往的研究主要集中于水氮耦合效应对小麦生长过程中生理生化特性、如何协同提高小麦产量和水氮利用率以及水氮互作如何影响土壤肥力等方面,但关于水氮互作的作物―土壤效应及其机制缺乏深入研究。
5.1.2 小麦水氮耦合与高效利用的分子机理尚不清楚
目前,主要从农艺和生理水平对小麦水氮耦合与高效利用开展了一些研究和观察,但对于水氮耦合效应的遗传学机制在很大程度上还不清楚。
5.1.3 未完全掌握协同提高小麦产量以及水氮利用效率的调控途径与技术
近年来,我国水分和氮肥消耗量愈渐增长,其中氮肥使用率自2000年以来一直呈现增长状态,消耗量已经达到每年3000万t[91]。水和氮利用效率却未得到显著提高,且目前小麦单产增长率还不能够满足人口增长和社会经济发展的需求。因此,通过何种途径或技术手段实现小麦高产优质高效协同仍亟待解决。
5.2 展望
5.2.1 高产小麦与土壤的水氮互作效应及其机制
通过在田间进行的水氮互作试验,研究在不同灌溉和施氮水平下小麦产量形成与水氮利用效率的动态变化规律;进一步研究水氮互作调控对酶学机制、内源激素、物质生产与运转等方面的影响;构建小麦高产与水氮高效利用的水氮耦合模型;从蛋白质表达、根系形态生理方面阐明小麦对水氮互作的响应与水氮高效利用的机理;揭示高产小麦水氮互作的作物―土壤效应及其机制。
5.2.2 水氮互作对小麦产量和水氮利用效率的分子生物学机制
一些蛋白质和基因可以同时影响水和氮肥利用效率并相互作用,在后续的研究中,应侧重于研究相关蛋白质和基因的具体功能;建立作物遗传图谱,寻找相关数量性状位点;进行水氮利用效率的基因定位和分子标记方面的研究。
5.2.3 协同提高小麦产量和水氮利用效率的调控途径与关键技术
在不同灌溉方式以及水氮互作下研究小麦产量形成及其土壤环境的变化,通过土壤水肥管理等技术研究,建立高产小麦土壤水分管理模式和水肥高效利用模式,开发能够同时满足高产和高水氮利用效率的调控途径和关键技术。
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缓/控释肥料是21世纪肥料发展的重要方向,对维护中国粮食安全、提高肥料利用率,实现农业生产与生态协调发展、节本增效和节能减排等方面均具有十分重要的意义。本文在总结缓/控释肥定义、作用机制、研究现状的基础上,阐述了缓/控释肥在农业生产中的应用效果和在应用及推广中存在的问题,提出了相应的对策和建议,展望了缓/控释肥的发展前景,以期为缓/控释肥的研究提供理论依据。
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Rising world population is expected to increase the demand for nitrogen fertilizers to improve crop yield and ensure food security. With existing challenges on low nutrient use efficiency (NUE) of urea and its environmental concerns, controlled release fertilizers (CRFs) have become a potential solution by formulating them to synchronize nutrient release according to the requirement of plants. However, the most significant challenge that persists is the “tailing” effect, which reduces the economic benefits in terms of maximum fertilizer utilization. High materials cost is also a significant obstacle restraining the widespread application of CRF in agriculture. The first part of this review covers issues related to the application of conventional fertilizer and CRFs in general. In the subsequent sections, different raw materials utilized to form CRFs, focusing on inorganic and organic materials and synthetic and natural polymers alongside their physical and chemical preparation methods, are compared. Important factors affecting rate of release, mechanism of release and mathematical modelling approaches to predict nutrient release are also discussed. This review aims to provide a better overview of the developments regarding CRFs in the past ten years, and trends are identified and analyzed to provide an insight for future works in the field of agriculture.
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DOI:10.3864/j.issn.0578-1752.2018.02.009
[本文引用: 2]
【目的】通过对华北地区一年两熟种植模式下冬小麦生长季不同施肥和灌溉处理下土壤细菌群落的研究,揭示长期不同施肥和灌溉制度下土壤细菌数量、多样性和群落结构的变化规律。为科学施肥和灌溉,提高农田地力和维持土壤微生物多样性等提供依据。【方法】依托中国农业大学吴桥实验站,选取长期施肥和灌溉定位试验的6个处理冬小麦收获后耕层土壤为研究对象,分别为化肥+不灌溉(CI0)、化肥+拔节期灌溉(CI1)、化肥+拔节期灌溉+灌浆期灌溉(CI2)、有机肥+不灌溉(MI0)、有机肥+拔节期灌溉(MI1)和有机肥+拔节期灌溉+灌浆期灌溉(MI2)。借助荧光定量PCR技术和Illumina Miseq高通量测序平台,以16S rRNA基因为标靶,研究长期不同施肥和灌溉制度对土壤细菌数量、多样性和群落结构的影响,并分析细菌数量、多样性和群落结构变化与土壤理化性质的相关性。【结果】灌溉显著提高了土壤含水量和土壤pH,施有机肥比施化肥显著提高了土壤有机碳含量。不同处理细菌16S rRNA基因拷贝数为每克干土4.34×109—1.39×1010。灌溉显著提高了细菌数量,化肥和有机肥处理分别提高了1.17—1.60和0.76—1.93倍。多样性指数结果表明灌溉显著影响细菌群落α多样性指数,施肥对细菌群落α多样性指数的影响均不显著。门水平上,18个样品共获得39个类群,其中变形菌门(Proteobacteria)、放线菌门(Actinobacteria)、绿弯菌门(Chloroflexi)、酸杆菌门(Acidobacteria)和拟杆菌门(Bacteroidetes)为优势类群,相对丰度共占77.22%—86.28%。不同处理间放线菌门(11.09%—27.01%)、拟杆菌门(5.45%—12.13%)和Saccharibacteria(2.41%—3.77%)的相对丰度差异显著。灌溉显著降低了放线菌门和Saccharibacteria的相对丰度,化肥和有机肥处理分别降低了36.48%—48.03%、22.17%—33.67%和15.21%—45.54%、13.40%—23.97%。层次聚类和主成分分析结果显示施肥和灌溉对细菌群落结构都产生影响,相同灌溉次数处理的细菌群落结构相似,而相同施肥处理间细菌群落结构差异较大,表明灌溉对细菌群落结构的影响强于施肥。此外,土壤含水量、土壤pH、全氮含量和有机碳含量与细菌数量、α多样性指数和群落结构存在一定的显著相关关系。【结论】灌溉显著改变了细菌数量、多样性和群落结构,施肥对细菌数量和群落结构的影响较小。土壤含水量和土壤pH是造成土壤细菌数量、多样性和群落结构差异的主要原因。
Plant community and soil chemistry responses to long‐term nitrogen inputs drive changes in alpine bacterial communities
Bacterial community composition and diversity was studied in alpine tundra soils across a plant species and moisture gradient in 20 y-old experimental plots with four nutrient addition regimes (control, nitrogen (N), phosphorus (P) or both nutrients). Different bacterial communities inhabited different alpine meadows, reflecting differences in moisture, nutrients and plant species. Bacterial community alpha-diversity metrics were strongly correlated with plant richness and the production of forbs. After meadow type, N addition proved the strongest determinant of bacterial community structure. Structural Equation Modeling demonstrated that tundra bacterial community responses to N addition occur via changes in plant community composition and soil pH resulting from N inputs, thus disentangling the influence of direct (resource availability) vs. indirect (changes in plant community structure and soil pH) N effects that have remained unexplored in past work examining bacterial responses to long-term N inputs in these vulnerable environments. Across meadow types, the relative influence of these indirect N effects on bacterial community structure varied. In explicitly evaluating the relative importance of direct and indirect effects of long-term N addition on bacterial communities, this study provides new mechanistic understandings of the interaction between plant and microbial community responses to N inputs amidst environmental change.
Combined effects of reduced irrigation and water quality on the soil microbial community of a citrus orchard under semi-arid conditions
DOI:10.1016/j.soilbio.2016.10.024 URL [本文引用: 1]
Nitrogen fertilization directly affects soil bacterial diversity and indirectly affects bacterial community composition
DOI:10.1016/j.soilbio.2015.09.018 URL [本文引用: 1]
Influences of irrigation amount on the rhizospheric microorganism composition and carbon dioxide flux of maize crops
DOI:10.1016/j.geoderma.2019.02.022 URL [本文引用: 1]
Response of the abundance of key soil microbial nitrogen-cycling genes to multi-factorial global changes
DOI:10.1371/journal.pone.0076500 URL [本文引用: 1]
The bacterial biogeography of British soils
DOI:10.1111/j.1462-2920.2011.02480.x
PMID:21507180
[本文引用: 1]
Despite recognition of the importance of soil bacteria to terrestrial ecosystem functioning there is little consensus on the factors regulating belowground biodiversity. Here we present a multi-scale spatial assessment of soil bacterial community profiles across Great Britain (> 1000 soil cores), and show the first landscape scale map of bacterial distributions across a nation. Bacterial diversity and community dissimilarities, assessed using terminal restriction fragment length polymorphism, were most strongly related to soil pH providing a large-scale confirmation of the role of pH in structuring bacterial taxa. However, while α diversity was positively related to pH, the converse was true for β diversity (between sample variance in α diversity). β diversity was found to be greatest in acidic soils, corresponding with greater environmental heterogeneity. Analyses of clone libraries revealed the pH effects were predominantly manifest at the level of broad bacterial taxonomic groups, with acidic soils being dominated by few taxa (notably the group 1 Acidobacteria and Alphaproteobacteria). We also noted significant correlations between bacterial communities and most other measured environmental variables (soil chemistry, aboveground features and climatic variables), together with significant spatial correlations at close distances. In particular, bacterial and plant communities were closely related signifying no strong evidence that soil bacteria are driven by different ecological processes to those governing higher organisms. We conclude that broad scale surveys are useful in identifying distinct soil biomes comprising reproducible communities of dominant taxa. Together these results provide a baseline ecological framework with which to pursue future research on both soil microbial function, and more explicit biome based assessments of the local ecological drivers of bacterial biodiversity.© 2011 Society for Applied Microbiology and Blackwell Publishing Ltd.
Soil enzyme activity response under the amendment of different types of biochar
DOI:10.3390/agronomy12030569
URL
[本文引用: 1]
Biochar (BC) is a material that finds many applications in agriculture and environmental activities. The aim of the study was to define the influence of biochar produced from various organic materials: mellow compost (MC), stabilized municipal sewage sludge (MSS), pine sawdust (PS), sycamore sawdust (SS) and oak leaves (OL) on soil enzyme activity, as well as its relations with carbon and nitrogen content. After a 60-day incubation of soil and BC, the activity of dehydrogenases (DEH), catalase (CAT), alkaline (AlP) and acid (AcP) phosphatases was investigated. The basic parameters of soil were also determined: TOC, TN, DOM, pH in H2O, available phosphorus (AP). The highest AP content was obtained in the S + MSS, S + OL and S + MC variants. Enzyme activity was highest in soil with MSS BC, regardless of incubation time. After 60 days, the activity of soil enzymes was inhibited. The obtained results indicate that the response of enzymatic activity to biochar depends on the feedstock material and the incubation time. When using BC as an exogenous matter, it is necessary to determine the TOC/TN ratio. For the very wide range of this parameter, supplemental nitrogen fertilization or mixtures of different biochars should be applied.
Variations in soil microbial community composition and enzymatic activities in response to increased N deposition and precipitation in Inner Mongolian grassland
DOI:10.1016/j.apsoil.2017.06.041 URL [本文引用: 1]
Changes in soil enzymes related to C and N cycle and in soil C and N content under prolonged warming and drought in a Mediterranean shrubland
DOI:10.1016/j.apsoil.2007.12.011 URL [本文引用: 1]
Perennial grain crop roots and nitrogen management shape soil food webs and soil carbon dynamics
DOI:10.1016/j.soilbio.2019.107573 URL [本文引用: 1]
Effects of drip irrigation on root activity pattern, root-sourced signal characteristics and yield stability of winter wheat
DOI:10.1016/j.agwat.2022.107783 URL [本文引用: 1]
Interactions between water and nitrogen in Australian cropping systems: physiological, agronomic, economic, breeding and modelling perspectives
DOI:10.1071/CP16027
URL
[本文引用: 2]
\n\nThis paper reviews the interactions between water and nitrogen from physiological, agronomic, economic, breeding and modelling perspectives. Our primary focus is wheat; we consider forage crops, sorghum and legumes where relevant aspects of water–nitrogen interactions have been advanced.\nFrom a physiological perspective, we ask: How does nitrogen deficit influence the water economy of the crop? How does water deficit influence the nitrogen economy of the crop? How do combined water and nitrogen deficit affect crop growth and yield? We emphasise synergies, and the nitrogen-driven trade-off between the efficiency in the use of water and nitrogen. The concept of nitrogen–water co-limitation is discussed briefly.\nFrom agronomic and economic perspectives, the need to match supply of nitrogen and water is recognised, but this remains a challenge in dryland systems with uncertain rainfall. Under-fertilisation commonly causes gaps between actual and water-limited potential yield. We discuss risk aversion and the role of seasonal rainfall forecasts to manage risk.\nFrom a breeding perspective, we ask how selection for yield has changed crop traits relating to water and nitrogen. Changes in nitrogen traits are more common and profound than changes in water-related traits. Comparison of shifts in the wheat phenotype in Australia, UK, Argentina and Italy suggests that improving yield per unit nitrogen uptake is straightforward; it requires selection for yield and allowing grain protein concentration to drift unchecked. A more interesting proposition is to increase nitrogen uptake to match yield gains and conserve protein in grain. Increased stomatal conductance is a conspicuous response to selection for yield which partially conflicts with the perception that reduced conductance at high vapour pressure deficit is required to increase water- use efficiency; but high stomatal conductance at high vapour pressure deficit may be adaptive for thermal stress.\nFrom a modelling perspective, water and nitrogen are linked in multiple ways. In crops where water limits growth, reduced biomass reduces nitrogen demand. Reciprocally, nitrogen limitation during crop expansion reduces leaf area index and increases the soil evaporation : transpiration ratio. Water–nitrogen interactions are also captured in the water-driven uptake of nitrogen by mass flow and diffusion and in the water-driven processes of nitrogen in soil (e.g. mineralisation).\nThe paper concludes with suggestions for future research on water-nitrogen interactions.\n
Dynamic responses of wheat to drought and nitrogen stresses during rewatering cycles
DOI:10.1016/j.agwat.2014.08.006 URL [本文引用: 3]
不同测墒补灌水平对小麦水氮利用及土壤硝态氮淋溶的影响
于2012—2014年两个小麦生长季,以全生育期不灌水(W<sub>0</sub>)为对照,设置3个测墒补灌处理,即拔节和开花期使0~140 cm土层土壤平均相对含水量分别为65%(W<sub>1</sub>)、70%(W<sub>2</sub>)和75%(W<sub>3</sub>),研究其对土壤水利用、小麦氮素积累转运和土壤硝态氮分布及籽粒产量的影响.结果表明: W<sub>2</sub>处理土壤贮水消耗量及占总耗水量的比例和灌溉水占总耗水量的比例较高,且吸收利用100~140 cm土层土壤贮水量较高.开花期营养器官氮素积累量及开花后氮素积累量均为W<sub>2</sub>、W<sub>3</sub>>W<sub>1</sub>>W<sub>0</sub>,成熟期营养器官氮素积累量为W<sub>3</sub>>W<sub>2</sub>>W<sub>1</sub>>W<sub>0</sub>,营养器官氮素向籽粒中的转移量和成熟期籽粒氮素积累量均为W<sub>2</sub>>W<sub>3</sub>>W<sub>1</sub>>W<sub>0</sub>.成熟期0~60 cm土层硝态氮含量表现为W<sub>0</sub>>W<sub>1</sub>>W<sub>2</sub>>W<sub>3</sub>,80~140 cm土层为W<sub>3</sub>显著高于其他处理,140~200 cm土层各处理间无显著差异.W<sub>2</sub>处理的籽粒产量、水分利用效率、氮素吸收效率及氮肥偏生产力均最高.在本试验条件下,综合考虑籽粒产量、水分利用效率、氮素吸收效率及土壤硝态氮的淋溶,W<sub>2</sub>处理是高产节水生态安全的最佳灌溉处理.
Nitrogen use by winter wheat and changes in soil nitrate nitrogen levels with supplemental irrigation based on measurement of moisture content in various soil layers
DOI:10.1016/j.fcr.2014.05.016 URL [本文引用: 1]
Strategies to increase barley production and water use efficiency by combining deficit irrigation and nitrogen fertilizer
DOI:10.1007/s00271-022-00811-0 [本文引用: 2]
Effects of reducing nitrogen application rate under different irrigation methods on grain yield, water and nitrogen utilization in winter wheat
DOI:10.3390/agronomy12081835
URL
[本文引用: 1]
We conducted a two-year field experiment on winter wheat (Triticum aestivum L.) from 2016–2018 to compare the effects of reducing nitrogen application rate in spring under three irrigation methods on grain yield (GY), water and nitrogen use efficiency in the North China Plain (NCP). Across the two years, GY of conventional irrigation (CI), micro-sprinkling irrigation (SI) and drip irrigation (DI) decreased by 6.35%, 9.84% and 6.83%, respectively, in the reduced nitrogen application rate (N45) than the recommended nitrogen application rate (N90). However, micro-irrigation (SI and DI) significantly increased GY relative to CI under the same nitrogen application rate, and no significant difference was observed in GY between SI and DI under N45, while SI obtained the highest GY under N90. The difference among different treatments in GY was mainly due to the variation in grain weight. The seasonal evapotranspiration (ET) in N45 was decreased more significantly than N90, and there was no significantly difference in ET among different irrigation methods under N45, but micro-irrigation significantly decreased the ET relative to CI under N90. Micro-irrigation significantly improved water use efficiency (WUE) compared to CI at the same nitrogen application rate. Under N45, compared with CI, WUE in SI and DI increased by 9.09% and 4.70%, respectively; however, the WUE increased by 15.9% and 7.23%, respectively, under N90. Reducing nitrogen application rate did not have a significant impact on WUE under CI, but it did have a substantial negative impact on SI and DI. Nitrogen accumulation in wheat plants at maturity (NAM) in N45 deceased significantly compared with N90 under the same irrigation method. Compared with CI under the same nitrogen application rate, micro-irrigation treatments significantly increased NAM, while SI was the largest. In comparison to N90, under three irrigation methods, N45 significantly increased nitrogen fertilizer use efficiency (NfUE). The highest NfUE was attained in SI, followed by DI, while CI was the lowest. Moreover, N45 significantly decreased soil NO3−-N accumulation (SNC) in three irrigation methods, and micro-irrigation significantly decreased the SNC in deep soil layers compared with CI when nitrogen is applied at the same level. Overall, micro-irrigation with a reduced nitrogen application rate in spring can achieve a relatively higher production of winter wheat while increasing the use efficiency of water and nitrogen and reducing soil NO3−-N leaching into deep soil layers in the NCP.
Wheat growth, yield, and quality under water deficit and reduced nitrogen supply. A review
DOI:10.1007/s10343-021-00615-w [本文引用: 1]
A nitrogen fertilizer strategy for simultaneously increasing wheat grain yield and protein content: Mixed application of controlled-release urea and normal urea
DOI:10.1016/j.fcr.2021.108405 URL [本文引用: 1]
Impacts of straw, biogas slurry, manure and mineral fertilizer applications on several biochemical properties and crop yield in a wheat-maize cropping system
DOI:10.17221/467/2018-PSE URL [本文引用: 1]
Nitrogen fertilization improved water-use efficiency of winter wheat through increasing water use during vegetative rather than grain filling
DOI:10.1016/j.agwat.2017.11.010 URL [本文引用: 1]
水氮互作对滴灌小麦土壤硝态氮运移、氮平衡及水氮利用效率的影响
经过2年滴灌小麦大田试验,研究不同灌水定额和不同施氮水平,这2个试验因素下小麦土壤硝态氮运移、氮平衡及水氮利用效率的变化情况。结果表明:2年内小麦各生育阶段耗水量和耗水模数均表现为抽穗扬花期=灌浆期>拔节孕穗期>分蘖期=成熟期>出苗期。在0~100 cm各处理在各生育期的硝态氮含量随土壤深度呈现减小趋势,表现出“上高下低”的趋势;其中土壤硝态氮含量均在0~20 cm出现最大值。在0~60 cm硝态氮含量与施氮量成正比,各处理下各生育期在80~100 cm土层的硝态氮变幅不大。当施氮量超过248 kg/hm<sup>2</sup>再增加施氮量,籽粒产量增加不明显,甚至有降低趋势,且影响籽粒吸收氮素。当灌水量大于390 mm,有很大部分水量因深层渗漏而损失,大量硝态氮也随着水分的渗漏而淋溶。此试验氮素损失中,淋溶的损失是主要途径,大部分硝态氮随水分渗漏被淋洗至60 cm以下的深层土壤中,且随施氮量的增加硝态氮淋溶更为严重。经过2年试验表明,施肥量在179~248 kg/hm<sup>2</sup>,灌水定额为45 mm,灌溉定额为390~405 mm的水氮组合,其表观损失较低,水氮利用效率较高,是适宜干旱地区多砾石砂土条件下的最佳水肥组合。
Optimized split nitrogen fertilizer increase photosynthesis, grain yield, nitrogen use efficiency and water use efficiency under water-saving irrigation
DOI:10.1038/s41598-019-56847-4
[本文引用: 1]
A large body of literature is available on wound healing in humans. Nonetheless, a standardized ex vivo wound model without disruption of the dermal compartment has not been put forward with compelling justification. Here, we present a novel wound model based on application of negative pressure and its effects for epidermal regeneration and immune cell behaviour. Importantly, the basement membrane remained intact after blister roof removal and keratinocytes were absent in the wounded area. Upon six days of culture, the wound was covered with one to three-cell thick K14+Ki67+ keratinocyte layers, indicating that proliferation and migration were involved in wound closure. After eight to twelve days, a multi-layered epidermis was formed expressing epidermal differentiation markers (K10, filaggrin, DSG-1, CDSN). Investigations about immune cell-specific manners revealed more T cells in the blister roof epidermis compared to normal epidermis. We identified several cell populations in blister roof epidermis and suction blister fluid that are absent in normal epidermis which correlated with their decrease in the dermis, indicating a dermal efflux upon negative pressure. Together, our model recapitulates the main features of epithelial wound regeneration, and can be applied for testing wound healing therapies and investigating underlying mechanisms.
Effects of nitrogen fertilizer and planting density on the leaf photosynthetic characteristics, agronomic traits and grain yield in common buckwheat (Fagopyrum esculentum M.)
DOI:10.1016/j.fcr.2018.02.001 URL [本文引用: 1]
Variability of leaf photosynthetic characteristics in rice and its relationship with resistance to water stress under different nitrogen nutrition regimes
DOI:10.1111/ppl.v167.4 URL [本文引用: 1]
Involvement of abscisic acid and cytokinins in the senescence and remobilization of carbon reserves in wheat subjected to water stress during grain filling
DOI:10.1046/j.1365-3040.2003.01081.x URL [本文引用: 1]
The role of photorespiration during drought stress: an analysis utilizing barley mutants with reduced activities of photorespiratory enzymes
DOI:10.1046/j.1365-3040.1999.00410.x URL [本文引用: 1]
Ammonia production and assimilation: its importance as a tolerance mechanism during moderate water deficit in tomato plants
DOI:10.1016/j.jplph.2010.11.018 URL [本文引用: 1]
Effect of IW: CPE-based irrigation scheduling and N-fertilization rate on yield, water and n-use efficiency of wheat (Triticum aestivum)
DOI:10.1007/s40003-020-00489-w [本文引用: 1]
Physiological mechanisms underlying reduced photosynthesis in wheat leaves grown in the field under conditions of nitrogen and water
DOI:10.1016/j.cj.2022.06.010 URL [本文引用: 2]
The Arabidopsis nitrate transporter NPF7.3/NRT1.5 is involved in lateral root development under potassium deprivation
Responses of barley root and shoot proteomes to long‐term nitrogen deficiency, short‐term nitrogen starvation and ammonium
DOI:10.1111/pce.2011.34.issue-12 URL [本文引用: 1]
Aquaporins and water permeability of plant membranes
Nitrogen availability affects hydraulic conductivity of rice roots, possibly through changes in aquaporin gene expression
DOI:10.1007/s11104-014-2070-4 URL [本文引用: 1]
Exposure to high nitrogen triggered a genotype‐dependent modulation of cell and root hydraulics, which can involve aquaporin regulation
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