• ISSN 1008-505X
  • CN 11-3996/S

氮磷互作对粳稻不同粒位籽粒灌浆及淀粉合成关键酶活性的调控机制

Regulatory mechanism of nitrogen-phosphorus interaction on grain filling and key enzyme activities involved in starch synthesis in grains at different positions of japonica rice

  • 摘要:
    目的 探究氮磷互作对粳稻产量、不同粒位籽粒灌浆特性以及淀粉合成关键酶活性的影响,为北方粳稻实现稳产增产的氮磷平衡施肥策略提供理论依据。
    方法 于2021和2022年在沈阳农业大学卡力玛水稻实验站进行田间试验,供试品种为粳稻北粳3号。设置9个氮磷组合处理,由3个氮、磷水平组成:N 210 kg/hm2和P2O5 105 kg/hm2 (常规量,N3、P3)、减量15% (N2、P2)和减量30% (N1、P1)。自抽穗开花(即稻穗顶端小穗开花)至成熟期,分别采集稻穗的强势粒、弱势粒和中势粒。一部分烘干、称重,利用Richards方程拟合灌浆过程,并计算灌浆特征相关参数;另一部分用于测定强势粒和弱势粒中腺苷二磷酸葡萄糖焦磷酸化酶(AGPP)、颗粒结合型淀粉合成酶(GBSS)、可溶性淀粉合成酶 (SSS)和淀粉分支酶 (SBE) 活性。在成熟期,调查水稻产量,单位有效穗数,以及强、中、弱势粒的粒数、结实率、千粒重。
    结果 不同氮肥水平下,达到最高产量所需的磷肥水平存在差异,其中N3P3、N3P2、N2P1处理的产量无显著差异,且三者产量均显著高于N3和N2水平下其他处理。N3水平下,适量减磷(P2)对产量构成因素无不利影响,过度减施磷肥(P1)显著降低了单位面积有效穗数、穗粒数、结实率;N2水平下,P3显著降低了单位面积有效穗数和穗粒数,P2显著降低了穗粒数,而P1增加了穗粒数,维持了千粒重;过量减施氮肥显著降低水稻产量构成因素和产量,N1水平下3个磷处理产量均显著低于N3水平。比较产量最高的3个处理,N2P1处理的强、中、弱势粒数均高于N3P3、N3P2处理,强、中势粒结实率与N3P3、N3P2处理差异不显著,但弱势粒结实率显著降低。千粒重受中势粒影响最大,受弱势粒影响次之,增施磷肥有利于提高强、中、弱势粒千粒重。在本研究氮、磷肥用量范围内,氮磷减施呈降低最大灌浆速率、平均灌浆速率、达到最大灌浆速率时的生长量和延长达到最大灌浆速率的时间、活跃生长期的趋势,与N3P3处理相比,N3P2和N2P1处理强、中、弱势粒的活跃生长期分别延长了3.6、2.3、2.3天和3.8、3.3、2.3天。与N3相比,N2抑制了强、弱势粒AGPP活性,而对SSS、SBE和GBSS活性无明显抑制;N1则降低了上述酶活性。磷肥减施抑制了强势粒AGPP和GBSS活性以及弱势粒GBSS活性。相关性分析结果表明,强势粒平均灌浆速率主要由其AGPP、SSS、GBSS活性共同调控,粒重主要受SSS活性的正向影响;弱势粒的粒重与其AGPP和SSS活性均呈极显著正相关。
    结论 磷肥显著影响强势粒灌浆速率、达到最大灌浆速率时的生长量及中势粒达到最大灌浆速率的时间,而氮肥显著影响中势粒达到最大灌浆速率时的生长量和弱势粒达到最大灌浆速率的时间。平衡氮磷用量可协调灌浆期强、弱势粒中淀粉合成关键酶活性,优化籽粒灌浆特性,延长强、中、弱势粒的活跃生长期,有利于发挥产量构成因素的综合优势,实现水稻增产稳产。

     

    Abstract:
    Objectives We analyzed the filling capacities and the starch synthesis activities at different positions of rice grains under different N and P supply levels, so as to provide a theoretical basis for balanced of nitrogen and phosphorus fertilization in japonica rice cultivation in northern China.
    Methods In 2021 and 2022, field trials were conducted at the Kalima Rice Experimental Station of Shenyang Agricultural University using the japonica Beigeng 3. Three N and P application levels were used to compose 9 treatments, conventional level N 210 kg/hm2 and P2O5 105 kg/hm2 (N3, P3), 15% reduction (N2, P2), and 30% reduction (N1, P1). From flowering (flower appeared on the top spikelets of rice) to maturing stage, the superior, medium and inferior grains were separately picked, and part of them were dried to determine the biomass weight, then using Richards equation to fit the filling process and calculated the grain filling parameters. The other part were used for determination of starch synthesis enzyme activities. At harvest, rice yield, effective panicles per unit area, and spikelets per panicle, seed setting rate, 1000-grain weight of superior, medium and inferior grains were investigated.
    Results The P levels required to achieve high yields varied among the N levels. Among N3P3, N3P2, and N2P1 treatments, similar yields were recorded, which were significantly higher than the other treatments. Under N3 level, P2 achieved similar yield components, but P1 achieved significantly lower effective panicles per unit area, spikelets per panicle and seed setting rate. Under the N2 level, P3 significantly decreased the effective panicles per unit area and spikelets per panicle, P2 significantly decreased the spikelets per panicle, while P1 increased the spikelets per panicle, and maintained similar 1000-grain weight. Compared with the N3 level, N1 significantly decreased the yield and its components of rice. Among the three high yield treatments, N2P1 exhibited higher spikelets per panicle of superior, medium, and inferior grains, showed similar seed-setting rate for superior and medium grains, but had a significantly lower seed setting rate for inferior grains compared to N3P3 and N3P2. The 1000-grain weight was most affected by the medium grains, followed by the inferior grains. P levels were beneficial to increase the 1000-grain weight of all the superior, medium and inferior grains. Within the tested nitrogen and phosphorus fertilizer range, reducing N and P reduced the maximum grain filling rate, mean grain filling rate and growth weight reaching maximum grain filling rate, prolonged the time reaching maximum grain filling rate and active growth period. Specifically, active growth period for the N3P2 and N2P1 treatments of superior, medium and inferior grains were extended by 3.6, 2.3, 2.3 days, and 3.8, 3.3, 2.3 days, respectively. Compared with N3, N2 suppressed ADP glucose pyrophosphorylase (AGPP) activity in superior and inferior grains, but the difference in the activities of soluble starch synthase (SSS), starch branching enzyme (SBE) and granule-bound starch synthase (GBSS) were minimal. N1 reduced the activity of these enzymes. Reducing P inhibited the activities of AGPP and GBSS in superior grains and the GBSS activity in inferior grains. Correlation analysis showed that the mean grain filling rate of superior grains was mainly regulated by the activities of AGPP, SSS and GBSS, and the 1000-grain weight was mainly positively affected by the activities of SSS. Notably, the 1000-grain weight of inferior grains exhibited a strong positive relationship with AGPP and SSS activities.
    Conclusions P level significantly influences the grain filling rate and growth weight reaching maximum grain filling rate in superior grains, and the time reaching maximum grain filling rate in medium grains, while N level significantly affects the growth weight reaching maximum grain filling rate in medium grains and the time reaching maximum grain filling rate in inferior grains. Balancing the N and P levels can harmonize the key enzyme activities of starch synthesis in superior and inferior grains during the filling period, affect the comprehensive advantages of grain filling characteristics and yield components of rice, ultimately achieving enhanced and stabilized rice productivity.

     

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