缓控释肥对水稻产量及灌浆期叶片碳氮代谢的影响
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Effects of Slow- and Controlled-Release Fertilizers on Grain Yield and Leaf Carbon and Nitrogen Metabolism during the Grain-Filling Stage in Rice
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收稿日期: 2026-04-15 修回日期: 2026-06-3 网络出版日期: 2026-08-10
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Received: 2026-04-15 Revised: 2026-06-3 Online: 2026-08-10
作者简介 About authors
江峥嵘,主要从事水稻生理生态研究,E-mail:
陈秋丽为共同第一作者,主要从事水稻生理生态研究,E-mail:
水稻灌浆期叶片碳氮代谢是调控产量形成的关键生理过程。为探究不同类型缓控释肥能否通过调控灌浆期叶片碳氮代谢进而影响水稻产量,于2020-2021年水稻生长季,以宁粳8号为供试材料,设置传统分次施肥(CK)及4种缓控释肥处理(速效缓释型肥料UF和SCU,控释型肥料PCU和BBF),开展田间试验。结果表明,UF和SCU处理的产量显著低于CK,2020年分别降低17.65%和5.88%,2021年分别降低13.40%和23.21%;而PCU和BBF处理的产量与CK相当,主要归因于其总颖花量和粒重均维持在CK水平。UF和SCU处理因水稻生育后期氮素供应不足,叶片光合能力下降,蔗糖与淀粉利用效率降低,碳氮代谢关键酶活性较低;PCU处理养分在生育后期大量释放,灌浆期氮素供应充足,促进了叶片碳代谢物的吸收利用,并增强了氮代谢物含量及相关酶活性;双峰释放模式的BBF处理在生育后期维持了适度的氮素供应,叶片光合能力较高,碳氮代谢水平趋于适中,从而使产量达到CK水平。综上,合理选择在水稻生育后期能够适量供应氮素的缓控释肥,有助于维持水稻高产。
关键词:
Carbon and nitrogen metabolism in rice leaves during the grain-filling stage is a key physiological process regulating yield formation. To investigate whether different types of slow- and controlled-release fertilizers influence rice yield by modulating leaf carbon and nitrogen metabolism during the grain-filling stage, a field experiment was conducted using the rice variety Ningjing 8 during the 2020-2021 growing seasons. The treatments comprised conventional split fertilization (CK) and four slow- and controlled-release fertilizers: the fast-acting slow-release fertilizers (UF and SCU), and the controlled-release fertilizers (PCU and BBF). The results showed that yields under UF and SCU were significantly lower than those under CK, with reductions of 17.65% and 5.88% in 2020, and 13.40% and 23.21% in 2021, respectively. By contrast, yields under PCU and BBF were comparable to CK, primarily due to their total spikelet number and grain weight being sustained at CK levels. The UF and SCU treatments resulted in insufficient nitrogen supply during the grain-filling stage, leading to reduced leaf photosynthetic capacity, lower sucrose and starch utilization efficiency, and diminished activities of key carbon and nitrogen metabolism enzymes. The PCU treatment, characterized by concentrated nutrient release during the grain-filling period, ensured adequate nitrogen supply throughout grain filling, thereby promoting the absorption and utilization of carbon metabolites in leaves while enhancing nitrogen metabolite content and associated enzyme activities. The BBF treatment, featuring a bimodal release pattern, provided a moderate nitrogen supply during the grain-filling stage, which supported relatively high leaf photosynthetic capacity and balanced carbon and nitrogen metabolic activity, ultimately achieving yields comparable to CK. In conclusion, selecting slow- and controlled-release fertilizers that ensure an adequate nitrogen supply during grain-filling stage of rice is an effective strategy to sustain high grain yield.
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本文引用格式
江峥嵘, 陈秋丽, 曾娜娜, 刘松, 林最昌, 杨扬, 丁艳锋, 李刚华.
Jiang Zhengrong, Chen Qiuli, Zeng Nana, Liu Song, Lin Zuichang, Yang Yang, Ding Yanfeng, Li Ganghua.
籽粒灌浆是水稻产量形成的关键过程。在此过程中,叶片既可作为光合器官合成碳水化合物,又是氮素等营养物质的主要“储存库”和“功能载体”,为水稻灌浆提供物质基础[4]。灌浆期叶片碳氮代谢的协调程度是决定水稻产量形成的关键[5]。碳氮代谢是水稻体内最基本的代谢途径,其相关酶活性与代谢产物之间既相互协调,又相互制约[6]。这种动态平衡关系的变化,直接影响碳、氮化合物在水稻体内的合成、转运与分配,最终决定水稻的产量与品质[7]。在水稻生产中,通过施用氮肥来调控植株体内的碳氮代谢过程,是改善水稻产量与品质的有效手段,且效果十分显著[8-9]。选用氮高效品种并配合氮肥后移,可以在花后促进氮素积累,提高碳氮代谢关键酶活性,促进营养器官中光合同化物的积累及转运[10]。有学者[11]发现,在灌浆过程中氮对叶片中的氮代谢产物氨基酸的积累有显著的正向影响。施氮量过高或者氮肥后移有利于植株氮吸收,消耗水稻植株碳水化合物,为蛋白质合成提供碳源[12];反之,减少氮肥施用有利于提高水稻叶片蔗糖合成酶活性和基因表达,促进植株碳同化物的合成与积累[13]。然而,与传统速效氮肥的多次施用或氮肥后移相比,缓控释肥能否在简化栽培管理的同时,维持灌浆期叶片较高的碳氮代谢活性以提高产量,有待进一步研究。
目前,关于缓控释肥在水稻生产上减少施氮量、减少施肥次数、降低氮肥损失量、提高氮肥利用率、提高水稻产量以及环保等方面的研究较多[14-15],也有研究[16]报道在缓控释肥处理下,土壤中的氮素有效性显著提高,叶片中的氮代谢酶活性提高,水稻的生长条件得到改善,进而提高产量。叶片作为水稻植株最重要的源端营养器官,其在灌浆期的碳氮代谢生理过程中尤为重要。因此,本研究以控氮控肥能力强、氮素利用效率高的宁粳8号为试验材料,选用氮素释放特性差异显著的缓控释肥,系统比较不同类型缓控释肥对水稻灌浆期叶片碳氮代谢过程中代谢产物、酶活性及基因表达量的影响,旨在筛选出能够保障水稻高产且省工节本、增产高效的缓控释肥种类,为水稻绿色高效生产提供理论依据。
1 材料与方法
1.1 试验地概况与试验材料
试验于2020-2021年在江苏省镇江市丹阳市延陵镇宝林村南京农业大学丹阳试验站(31°56′39′′ N,118°59′13′′ E,海拔80 m)进行。试验地位于长江三角洲水稻主产区和高产区,属于湿润的亚热带气候,是长江流域典型气候之一。土壤为黄粘土,水稻田0~20 cm耕层土壤的基本理化性质为pH 6.4、全氮1.3 g/kg、速效氮7.9 mg/kg、速效磷16.9 mg/kg、速效钾139.6 mg/kg、有机质18.3 g/kg。
供试品种为当地高产品种宁粳8号。
1.2 试验设计
试验采用随机区组设计。以传统分次施肥(CK)为对照,分4次施用尿素氮肥,基肥、分蘖肥、促花肥和保花肥的比例为3:2:3:2。试验设置4个缓控释肥施肥处理(图1),分别为硫包膜型缓释肥(SCU)、树脂包膜型缓控释肥(PCU)、脲甲醛(UF)和缓释复混肥(BBF),每个处理3次重复。各缓控释肥均由汉枫农业科技(泰州)有限公司(中国江苏)提供。其中UF和SCU属于速效缓释型肥,呈现出爆发性释放模式,在水稻生育前期释放较快,中、后期释放有限。PCU和BBF则属于控释型肥料。其中,PCU作为SCU的升级产品,具有在水稻生育前期少量释放,后期持续快速释放的特征。BBF是由多种缓释肥和控释肥混合而成的水稻专用缓混肥,由南京农业大学水稻栽培课题组研发,适用于水稻机插。BBF的养分释放特征居于SCU和PCU之间,表现为双峰释放模式,即生育前期释放量较大,后期释放量较小,确保养分供应与水稻生长需求协调匹配[17]。
图1
图1
不同类型缓控释肥的积累氮及阶段氮释放速率
Fig.1
Cumulative N and interval N release rate of slow-controlled release fertilizer
试验施氮量均为210 kg N/hm2,缓控释肥及磷钾肥均为一次性基施,其中P2O5 73.5 kg/hm2、K2O 81.9 kg/hm2,其余栽培管理措施按当地高产要求进行。各小区均于每年5月中下旬播种,6月中下旬机械移栽,机插密度为30.0 cm×12.1 cm,11月上旬收获测产。
1.3 测定项目与方法
1.3.1 产量及其构成因素
于水稻成熟期,每小区普查60穴水稻的有效穗数,并计算平均值,按平均有效穗数取代表穴10穴考种,测定穗粒数、结实率和粒重,并计算产量。各小区水稻全部收获并晒干后测定产量。
1.3.2 叶色及光合参数
于水稻花后7、21和35 d用叶绿素仪SPAD-502(Minolta,日本)测定挂牌标记剑叶的相对叶绿素含量(SPAD值)。同时,于晴朗上午9:00-11:00,利用LI-6400便携式光合作用测定仪(Li-COR,美国)测定各小区长势一致剑叶的净光合速率(Pn)、气孔导度(Gs)和蒸腾速率(Tr)。各小区测定10片叶片,3次重复。
1.3.3 碳氮代谢产物
于水稻花后0、7、14、21、28、35和42 d,在每个施肥处理下,分别从3个重复小区中各取30片剑叶,经105 ℃杀青、80 ℃烘干后,粉碎过100目筛,用于测定碳氮代谢产物。参考Hendrix[18]的方法并加以修改测定蔗糖含量。采用蒽酮比色法[19]测定叶片中的淀粉含量。用NCH分析仪(Sumika Chemical Analysis Service,Tokyo,日本)的Dumas燃烧法[20]测定氮含量。采用考马斯亮蓝法[21]测定可溶性蛋白含量。用磺基水杨酸提取,上清液用0.22 mm过滤头过滤,采用日立L-8900氨基酸自动分析仪(Hitachi Corp,日本)测定游离氨基酸含量。
1.3.4 碳氮代谢关键酶活性
1.3.5 基因表达
于水稻花后7、21和35 d,在每个施肥处理下,分别从3个重复小区中各取30片新鲜剑叶,进行蔗糖转运蛋白和碳氮同化关键基因表达的测定。叶片中的总RNA由RNeasy Plant Mini Kit(QIAGEN公司,中国)提取纯化,根据试剂盒方法合成cDNA第1链,稀释后用于荧光定量PCR,其中PCR引物由北京擎科生物科技股份有限公司合成,引物信息详见表1。
表1 基因引物信息
Table 1
| 基因Gene | 正向引物Forward primer (5’-3’) | 反向引物Reverse primer (3’-5’) |
|---|---|---|
| OsActin | CAATCGTGAGAAGATGACCC | GTCCATCAGGAAGCTCGTAGC |
| OsSUT2 | TCTTTTATCGGTGGGCTGGT | TTGCAAAGAATGGCCGACAA |
| OsSUT4 | CTCGTGCCCTTTTAGCTGAC | AACGTTTCCAACAGCCATCC |
| OsGS2 | TGGGTTGGAGGAACTGGAATT | ACGTTTGTTAGTAGGGATGGG |
| OsDOF11 | CAATGCCCATCATGCCAGCA | GCTCTGCAGCCTTGGACTAT |
1.4 数据处理
采用Microsoft Excel 2016计算试验数据平均值和标准误差,采用Graphpad prism 7软件作图,采用SPSS 25.0统计软件对数据进行单因素方差分析(ANOVA)和Duncan法多重比较。
2 结果与分析
2.1 缓控释肥对水稻产量及其构成因素的影响
相较于传统分次施肥(CK),不同种类缓控释肥(UF、SCU、PCU和BBF)对水稻产量的影响存在明显差异(表2)。在2020与2021年的水稻生长季,UF和SCU处理的水稻产量显著低于CK处理,其中UF处理产量分别减少17.65%和13.40%,SCU处理产量分别降低5.88%和23.21%;而PCU和BBF处理的产量与CK处理差异不显著。这主要归因于UF和SCU处理的每穗粒数及总颖花量显著低于CK处理,而PCU和BBF处理与CK处理无明显差异。此外,所有缓控释肥处理均能使结实率维持在与CK相当的水平,同时显著提高水稻粒重。年份与处理互作分析结果显示,不同缓控释肥处理在产量、总颖花量和粒重方面均存在显著差异,表明PCU和BBF处理主要通过提升总颖花量和粒重,进而实现与CK相当的产量水平。
表2 2020-2021年不同类型缓控释肥对水稻产量及其构成因素的影响
Table 2
| 年份 Year | 肥料类型 Fertilizer type | 产量 Yield (t/hm2) | 有效穗数 Effective panicle number (/m2) | 每穗粒数 Grain number per panicle | 总颖花量 Total number of spikelets (/hm2) | 结实率 Seed-setting rate (%) | 粒重 Grain weight (mg) |
|---|---|---|---|---|---|---|---|
| 2020 | CK | 10.2ab | 314.4ab | 129.0a | 4.1a | 93.9a | 26.9c |
| UF | 8.4c | 286.2c | 112.3b | 3.2c | 94.9a | 27.6ab | |
| SCU | 9.6b | 305.5abc | 117.5b | 3.6bc | 95.4a | 28.0a | |
| PCU | 10.2ab | 293.3bc | 131.8a | 3.9ab | 94.9a | 27.8a | |
| BBF | 10.8a | 325.3a | 131.2a | 4.3a | 93.6a | 27.2bc | |
| 2021 | CK | 11.2a | 323.0a | 137.9a | 4.4a | 96.8a | 26.1c |
| UF | 9.7b | 335.7a | 110.5c | 3.7b | 93.6a | 28.3a | |
| SCU | 8.6b | 343.3a | 94.2c | 3.2c | 94.9a | 28.1ab | |
| PCU | 11.1a | 368.3a | 116.0abc | 4.3a | 96.3a | 27.2b | |
| BBF | 11.0a | 355.7a | 117.3ab | 4.2a | 95.6a | 27.5ab | |
| F-value | 处理(T) | ** | ns | ** | ** | ns | ** |
| 年份(Y) | ** | ** | * | * | ns | ns | |
| T×Y | ** | ns | ns | ** | ns | * |
不同小写字母表示各处理间差异显著(P < 0.05);“**”和“*”分别表示在P < 0.01和P < 0.05水平上差异显著,ns表示无显著差异。
Different lowercase letters indicate significant differences among different treatments (P < 0.05);“**”and“*”indicate significant differences at
P < 0.01 and P < 0.05 levels, respectively, ns means no significant difference.
2.2 缓控释肥对水稻灌浆期剑叶光合作用的影响
随着灌浆进程的推进(花后0~42 d),剑叶的SPAD值、Pn、Gs及Tr均呈逐渐下降趋势(图2)。在SPAD值方面,PCU和BBF处理在整个灌浆期表现良好,其SPAD值与CK处理无显著差异;而UF和SCU处理的SPAD值在灌浆中、后期显著低于CK处理,叶色褪绿较快。同时,PCU和BBF处理的光合能力整体优于UF和SCU处理。值得注意的是,在2021年,PCU和BBF处理的Pn在灌浆期可达到与CK处理相当的水平。相比之下,UF和SCU处理的Pn在2年中均明显低于CK处理,尤其在灌浆后期下降更为显著。此外,各施肥处理Gs的变化趋势与Pn基本一致,PCU和BBF处理的Gs整体高于UF和SCU处理,但均低于CK处理。因此,PCU和BBF处理在维持灌浆期剑叶光合功能方面表现优于UF和SCU处理,可达到或接近CK处理的水平。
图2
图2
2020-2021年不同类型缓控释肥对水稻灌浆期剑叶SPAD值及光合参数的影响
不同小写字母表示各处理间差异显著(P < 0.05),下同。
Fig.2
Effects of different types of slow-controlled release fertilizer on SPAD value and photosynthetic parameters of flag leaves during grain-filling stage of rice in 2020 and 2021
Different lowercase letters indicate significant differences among different treatments (P < 0.05), the same below.
2.3 缓控释肥对水稻灌浆期叶片中碳代谢物含量及其关键酶活性的影响
灌浆期剑叶中蔗糖含量呈先降低后升高的趋势(图3)。与CK处理相比,BBF处理的剑叶蔗糖含量在整个灌浆期均显著升高,而SCU和PCU处理大多未表现出显著差异。此外,UF处理剑叶蔗糖含量仅在花后0、35和42 d显著高于CK处理。就淀粉含量而言,各处理间差异显著。值得注意的是,PCU处理剑叶淀粉含量在整个灌浆期间大多显著低于CK处理,或与CK处理无显著差异;而UF与SCU处理在花后28~42 d均显著高于CK处理;BBF处理则处于各处理组的中等水平。剑叶中碳代谢关键酶SPS和AGP的活性随灌浆进程呈逐渐下降趋势。与CK相比,UF和SCU处理的碳代谢酶活性多显著降低,而PCU和BBF处理则维持在接近CK的水平。然而,PCU处理的剑叶淀粉含量显著低于CK,表明其叶片光合固定的碳并未滞留积累于叶片,而是被高效转运至籽粒用于灌浆。相比之下,BBF处理酶活性稳定,淀粉含量中等,呈现相对平衡的碳分配。
图3
图3
不同类型缓控释肥对水稻灌浆期剑叶中碳代谢物含量及其关键酶活性的影响
Fig.3
Effects of different types of slow-controlled release fertilizer on the change of carbon metabolite contents and key enzyme activities in flag leaves of rice
2.4 缓控释肥对水稻灌浆期叶片氮代谢物含量及其关键酶活性的影响
灌浆期,剑叶中氮代谢物含量及其关键酶活性多呈逐渐下降的趋势(图4)。就氮代谢物而言,PCU处理下剑叶氮含量、总游离氨基酸含量及可溶性蛋白含量均维持在接近CK处理的水平;而UF和SCU处理下,这些氮代谢物含量多显著低于CK处理。氮代谢关键酶(NR、GS和GOGAT)活性的变化趋势与氮代谢物含量基本一致。其中,BBF处理下剑叶氮代谢物含量及关键酶活性均处于各处理组中的适中水平,表明其对氮代谢的调控作用较为平衡。
图4
图4
不同类型缓控释肥对水稻灌浆期剑叶中氮代谢物含量及其关键酶活性的影响
Fig.4
Effects of different types of slow-controlled release fertilizer on the change of nitrogen metabolite contents and key enzyme activities in flag leaves of rice
2.5 水稻产量构成因素与剑叶碳氮代谢关键酶活性的相关性
进一步分析产量构成因素与灌浆期剑叶碳氮代谢关键酶活性的相关性分析(图5)可知,不同缓控释肥处理下,水稻产量及总颖花数多与灌浆期剑叶碳氮代谢关键酶活性呈显著正相关,其中与SPS、GS和GOGAT的活性关系最为密切。值得注意的是,灌浆期各处理下粒重与上述剑叶酶活性多呈负相关。这可能是由于灌浆期叶片逐渐衰老导致碳氮代谢酶活性下降,而此时籽粒灌浆仍在进行,并且粒重的最终积累多依赖于灌浆前期叶片光合产物的储备与转运效率。其中,AGP主要在灌浆前期和中期与产量构成参数呈显著正相关,表明其在灌浆前期和中期淀粉转化过程中发挥关键作用;NR在灌浆前期和后期与产量呈显著正相关,在灌浆前期和中期与总颖花数和粒重存在相关性,反映出氮同化过程在水稻奠定颖花数和维持叶片功能、促进氮素再转运的双重贡献。
图5
图5
缓控释肥处理下水稻产量构成因素与剑叶碳氮代谢关键酶活的相关性分析
Y:产量,TNS:总颖花量,GW:粒重。7DAA、21DAA、35DAA分别表示花后7 d(灌浆前期)、21 d(灌浆中期)和35 d(灌浆后期)。
Fig.5
Correlation analysis between yield components and key enzyme activities of carbon and nitrogen metabolism in flag leaves of rice
Y; yield, TNS: total number of spikelets, GW: grain weight. 7DAA, 21DAA, and 35DAA represent 7 (early grain filling), 21 (mid-grain filling), and 35 (late grain filling) days after anthesis, respectively. *: P < 0.05, **: P < 0.01.
2.6 缓控释肥对水稻灌浆期叶片碳氮同化关键基因表达的影响
不同类型缓控释肥处理对叶片碳氮同化关键基因的表达具有差异化调控作用(图6)。其中,蔗糖装载关键基因(OsSUT2,OsSUT4)、氮同化关键基因(OsGS2)以及协同调控碳氮同化的关键转录因子(OsDOF11)在不同处理下的表达水平存在明显差异。在灌浆期,PCU处理下这些基因的表达量均显著上调,尤其在灌浆前期和中期最为明显。相比之下,UF和SCU处理下各基因的表达水平多与CK无显著差异,部分基因表达量甚至低于CK处理水平。BBF处理对剑叶碳氮同化关键基因表达的调控作用较为适中,多数基因呈显著上调或维持在与CK相当的水平,但整体上调幅度未超过PCU处理。因此,PCU处理可增强灌浆前、中期碳氮协同代谢关键基因的表达,而BBF处理则表现出较为适中的调控模式。
图6
图6
不同类型缓控释肥对水稻灌浆期剑叶中碳氮同化关键基因表达的影响
Fig.6
Effects of different types of slow-controlled release fertilizer on the key genes expression of carbon and nitrogen assimilate in flag leaves of rice
3 讨论
缓控释肥是一种利用效率高、省时省工的新型肥料,可满足作物养分的供需平衡[26]。其中,速效缓释型肥料UF和SCU在水稻生育前期氮素释放较快,生育中、后期释放有限;而控释型肥料PCU和BBF可在全生育期释放氮素,并且BBF肥料呈现双峰释放模式。本研究中,UF和SCU处理的水稻产量明显低于CK处理,而PCU和BBF处理使产量维持在与CK相当的水平。缓控释肥的多级释放机制可满足作物全生育期的生长需求,对水稻产量和品质具有一定的提升作用[27]。这说明,在水稻生长阶段持续提高营养物质的PCU和BBF肥料可充分提供水稻生长发育所需养分,在保证产量的基础上实现省工省力。其中,PCU和BBF处理的总颖花量和粒重显著高于其他缓控释肥处理,并且与CK处理无明显差异,这可归因于PCU和BBF在孕穗期及灌浆期均可持续性释放,为水稻的孕穗和灌浆过程充分提供养分。
水稻籽粒灌浆期间,叶片提供了所需碳同化物的60%~80%,其中剑叶扮演着至关重要的角色[28]。本研究发现,PCU和BBF处理下剑叶的SPAD值及光合参数均可达到CK的水平,而UF和SCU处理则明显低于CK。已有研究[29]表明,氮肥后移对禾本科作物剑叶的光合能力具有显著的促进作用。这说明在灌浆期能够持续释放养分的PCU和BBF肥料,可使水稻的光合能力维持在与CK处理相当的水平。灌浆期,水稻叶片在光照期间通过光合作用合成蔗糖,既可通过碳代谢关键酶SPS和AGP等将蔗糖转化成淀粉临时储存在叶片中,也可通过叶片蔗糖装载的模式立即转运至籽粒[30]。本研究中,UF和BBF处理下蔗糖含量略高于CK处理,其余处理与CK无差异。同时,UF和SCU处理灌浆中、后期淀粉含量显著高于CK,PCU处理则在整个灌浆期显著低于CK,呈低淀粉滞留特征。从碳代谢关键酶活性来看,UF和SCU处理的SPS和AGP活性多显著低于CK处理,而PCU和BBF处理的酶活性则接近CK处理。在碳代谢物转运关键基因表达方面,蔗糖转运子OsSUT2和OsSUT4可以以质外体方式,将叶片中的蔗糖装载入筛管伴胞细胞并运输至未成熟的籽粒中,供其生长发育[31]。PCU处理下剑叶OsSUT2和OsSUT4的表达量均明显高于CK处理,表明PCU肥料可显著增强叶片的碳同化物转运能力。相比之下,BBF处理下剑叶碳代谢物含量及碳同化关键基因表达水平均处于各缓控释肥处理的适中水平。因此,PCU和BBF均可通过促进叶片碳同化物积累和转运能力以改善水稻籽粒灌浆。其中,PCU表现为高效转运和低淀粉滞留的特征,而双峰释放模式的BBF则在碳分配与积累之间表现出更好的平衡性。
氮素营养对叶片中的游离氨基酸和蛋白质有正向影响[9]。可溶性蛋白作为植物体中最活跃的蛋白质之一,参与植物生理代谢中多种酶类的调节[32]。在缓控释肥施用下,PCU处理在灌浆期剑叶中的氮含量、游离氨基酸及可溶性蛋白含量均可达到CK的水平,而UF和SCU处理则显著低于CK,BBF处理居中。随着施氮量的提高,氮代谢酶活性增强,可促进氨基酸的合成与积累。叶片中NR、GS、GOGAT是3种最主要的氮代谢酶[33]。本研究中,各缓控释肥处理下这些酶活性的变化趋势与氮代谢物含量基本一致:UF和SCU处理较低,PCU和CK处理较高,BBF处理居中。已有研究[34]表明,氮肥后移或增加施氮量,可提高氮素营养,进而提高叶片中的可溶性蛋白含量,延缓叶片衰老。因此,全生育期持续释放养分的PCU和BBF肥料,在灌浆期仍能释放氮素,通过提高氮代谢关键酶(NR、GS、GOGAT)活性促进叶片中氮代谢物的合成与积累;而生育前期爆发性释放的UF和SCU肥料,因灌浆期供氮不足,导致氮代谢酶活性下降、氮代谢物含量降低,无法维持叶片正常的氮素营养水平。
不同缓控释肥处理下碳氮代谢关键酶活性与水稻的总颖花数、粒重和产量显著相关。生育后期大量释放营养的PCU肥料明显可加强叶片中的碳转运能力和氮代谢能力,使得叶片中淀粉积累较少,且氮代谢物含量较高。碳氮同化关键基因OsGS2与OsDOF11均为调控水稻产量形成与籽粒灌浆过程的重要转录因子。其中,OsGS2是氮同化过程中GS/GOGAT循环的关键调控基因,核心功能在于调控籽粒大小与产量[35];OsDOF11属于DOF转录因子家族,主要通过调控糖分运输,并间接影响氮素吸收与同化能力,从而维持水稻的碳氮平衡[36]。生育期双峰释放模式的BBF肥料在灌浆期适量释放营养,导致叶片中碳氮代谢关键酶活及OsSUT2/4、OsGS2和OsDOF11的表达模式趋于适中,使得叶片具有较好的碳氮代谢平衡能力。因此,全生育期持续释放养分的缓控释肥(如PCU和BBF肥料),有助于维持叶片的碳氮代谢能力,从而提高水稻产量。然而,在灌浆期适量释放养分的BBF肥料,则更有利于维持叶片的碳氮代谢平衡。
4 结论
不同类型缓控释肥对水稻灌浆期剑叶光合特性、碳氮代谢及产量具有显著的差异化调控作用。其中,生育后期快速释放型肥料(PCU)处理下,剑叶碳氮代谢物含量、关键酶活性(SPS、AGP、NR、GS和GOGAT)及相关基因表达水平均可达到甚至超过CK水平,同时剑叶SPAD值、Pn和Gs维持在较高水平,为籽粒灌浆提供了充足的光合同化物,进而提升了总颖花量和粒重。生育期双峰释放型肥料(BBF)则表现出适中的碳氮代谢调控模式,有利于维持碳氮平衡,同样实现了与CK相当的产量。相比之下,生育期速效释放型肥料(UF、SCU)处理下,多数碳氮代谢指标及光合性能均低于CK。因此,合理选择以BBF为代表的缓控释肥,可在简化施肥方式的同时维持水稻叶片碳氮代谢平衡,实现与CK处理相当的产量水平。
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PMID:35579477
[本文引用: 1]
The grain filling of rice depends on photoassimilates from leaves and stems. Phloem loading is the first crucial step for the transportation of sucrose to grains. However, phloem loading mechanisms in rice leaves and stems and their response to nitrogen (N) remain unclear. Here, using a combination of electron microscopy, transportation of a phloem tracer and 13C labeling, phloem loading was studied in rice leaves and stems. The results showed that the sieve element-companion cell complex lacked a symplastic connection with surrounding parenchyma cells in leaves and stems. The genes expression and protein levels of sucrose transporter (SUTs) and sugars will eventually be exported transporters (SWEETs) were detected in the vascular bundle of leaves and stems. A decrease in the 13C isotope remobilization from leaves to stems and panicles following treatment with p-chloromercuribenzenesulfonic acid indicated that rice leaves and stems actively transport sucrose into the phloem. Under low-N (LN) treatment, the activities of α-amylase, β-amylase and sucrose phosphate synthase (SPS) in stems and activity of SPS in leaves increased; genes expression and protein levels of SUTs and SWEETs in leaves and stems increased; the 13C isotope reallocation in panicles increased. These indicated that LN enhanced apoplastic phloem loading in stems and leaves. This improved the translocation of photoassimilates and consequently increased grain filling percentage, grain weight and harvest index. This study provides evidence that rice leaves and stems utilize an apoplastic loading strategy and respond to N stimuli by regulating the genes expression and protein levels of SUTs and SWEETs.© The Author(s) 2022. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.
Effects of nitrogen fertilization on nitrogen metabolism and grain protein in premium-tasty japonica rice
DOI:10.1007/s42976-023-00480-5 [本文引用: 1]
Nitrogen modulates grain yield,nitrogen metabolism, and antioxidant response in different rice genotypes
DOI:10.1007/s00344-022-10684-4 [本文引用: 1]
Comparative transcriptome analysis reveals major genes, transcription factors and biosynthetic pathways associated with leaf senescence in rice under different nitrogen application
DOI:10.1186/s12870-024-05129-x
PMID:38760728
[本文引用: 1]
Rice (Oryza sativa L.) is one of the most important food crops in the world and the application of nitrogen fertilizer is an effective means of ensuring stable and high rice yields. However, excessive application of nitrogen fertilizer not only causes a decline in the quality of rice, but also leads to a series of environmental costs. Nitrogen reutilization is closely related to leaf senescence, and nitrogen deficiency will lead to early functional leaf senescence, whereas moderate nitrogen application will help to delay leaf senescence and promote the production of photosynthetic assimilation products in leaves to achieve yield increase. Therefore, it is important to explore the mechanism by which nitrogen affects rice senescence, to search for genes that are tolerant to low nitrogen, and to delay the premature senescence of rice functional leaves.The present study was investigated the transcriptional changes in flag leaves between full heading and mature grain stages of rice (O. sativa) sp. japonica 'NanGeng 5718' under varying nitrogen (N) application: 0 kg/ha (no nitrogen; 0N), 240 kg/ha (moderate nitrogen; MN), and 300 kg/ha (high nitrogen; HN). Compared to MN condition, a total of 10427 and 8177 differentially expressed genes (DEGs) were detected in 0N and HN, respectively. We selected DEGs with opposite expression trends under 0N and HN conditions for GO and KEGG analyses to reveal the molecular mechanisms of nitrogen response involving DEGs. We confirmed that different N applications caused reprogramming of plant hormone signal transduction, glycolysis/gluconeogenesis, ascorbate and aldarate metabolism and photosynthesis pathways in regulating leaf senescence. Most DEGs of the jasmonic acid, ethylene, abscisic acid and salicylic acid metabolic pathways were up-regulated under 0N condition, whereas DEGs related to cytokinin and ascorbate metabolic pathways were induced in HN. Major transcription factors include ERF, WRKY, NAC and bZIP TF families have similar expression patterns which were induced under N starvation condition.Our results revealed that different nitrogen levels regulate rice leaf senescence mainly by affecting hormone levels and ascorbic acid biosynthesis. Jasmonic acid, ethylene, abscisic acid and salicylic acid promote early leaf senescence under low nitrogen condition, ethylene and ascorbate delay senescence under high nitrogen condition. In addition, ERF, WRKY, NAC and bZIP TF families promote early leaf senescence. The relevant genes can be used as candidate genes for the regulation of senescence. The results will provide gene reference for further genomic studies and new insights into the gene functions, pathways and transcription factors of N level regulates leaf senescence in rice, thereby improving NUE and reducing the adverse effects of over-application of N.© 2024. The Author(s).
A new gain-of-function OsGS2/GRF4 allele generated by CRISPR/Cas 9 genome editing increases rice grain size and yield
DOI:10.1016/j.cj.2022.01.004 URL [本文引用: 1]
OsDOF11 affects nitrogen metabolism by sucrose transport signaling in rice (Oryza sativa L.)
DOI:10.3389/fpls.2021.703034
URL
[本文引用: 1]
Carbon and nitrogen antagonistically regulate multiple developmental processes. However, the molecular mechanism affecting nitrogen metabolism by sucrose transport remains poorly defined. Previously, we noted that Oryza sativa DNA BINDING WITH ONE FINGER 11 (OsDOF11) mediated sucrose transport by binding to the promoter regions of Sucrose Transporter 1 (SUT1), Oryza sativa Sugars Will Eventually be Exported Transporters 11 (OsSWEET11), and OsSWEET14. Here, we note that OsDOF11 promotes nitrogen uptake and then maintains the ratio of fresh weight to dry weight in seedling plants and the effective leaf blade at flowering stages. Mutants of the sucrose transporter gene OsSWEET14 displayed a phenotype similar to that of OsDOF11. By microarray analysis and qRT-PCR in OsDOF11 mutant plants, OsDOF11 affected the transcription level of amino acid metabolism-related genes. We further found that mainly amino acid contents were reduced in flag leaves but increased in seeds. Both sugar and organic nitrogen changes caused the ratio of fresh weight to dry weight to decrease in OsDOF11 mutant seedling plants and mature leaves, which might result in vigorous reduced metabolic activity and become less susceptible to stress. These results demonstrated that OsDOF11 affected nitrogen metabolism by sugar distribution in rice, which provided new insight that OsDOF11 coordinated with C and N balance to maintain plant growth activity.
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