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作物学报 ›› 2023, Vol. 49 ›› Issue (7): 1968-1978.doi: 10.3724/SP.J.1006.2023.24152

• 耕作栽培·生理生化 • 上一篇    下一篇

施氮量对高粱籽粒灌浆及淀粉累积的影响

王媛1,2(), 王劲松1, 董二伟1, 刘秋霞1, 武爱莲1, 焦晓燕1,*()   

  1. 1山西农业大学资源环境学院, 山西太原 030031
    2高粱遗传与种质创新山西省重点实验室, 山西晋中 030600
  • 收稿日期:2022-06-30 接受日期:2022-10-10 出版日期:2023-07-12 网络出版日期:2022-10-19
  • 通讯作者: *焦晓燕, E-mail: jiaoxiaoyan@sxagri.ac.cn
  • 作者简介:E-mail: wangyuan1520@126.com
  • 基金资助:
    本研究由高粱遗传与种质创新山西省重点实验室开放基金项目(2019K-2);财政部和农业农村部: 国家现代农业产业技术体系建设专项(CARS-06-13.5-A20);山西省应用基础青年科技研究基金项目(201901D211559)

Effect of nitrogen application level on grain starch accumulation at grain filling stage in sorghum spikelets

WANG Yuan1,2(), WANG Jin-Song1, DONG Er-Wei1, LIU Qiu-Xia1, WU Ai-Lian1, JIAO Xiao-Yan1,*()   

  1. 1College of Resources and Environment, Shanxi Agricultural University, Taiyuan 030031, Shanxi, China
    2Key Laboratory of Sorghum Genetics & Germplasm Innovation of Shanxi Province, Jinzhong 030600, Shanxi, China
  • Received:2022-06-30 Accepted:2022-10-10 Published:2023-07-12 Published online:2022-10-19
  • Contact: *E-mail: jiaoxiaoyan@sxagri.ac.cn
  • Supported by:
    The Shanxi Province Key Laboratory of Sorghum Genetics & Germplasm Innovation(2019K-2);The China Agriculture Research System of MOF and MARA(CARS-06-13.5-A20);The Science and Technology Research Project for Youth in Shanxi Province(201901D211559)

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摘要：

本试验于2019—2020年以汾酒粱1号为材料, 在0、75、150、225、300 和450 kg N hm-2 6个施氮水平下, 于花后每隔7 d采集不同穗位籽粒分析其灌浆特性及淀粉形成过程, 探究不同氮素用量对高粱籽粒灌浆及单粒淀粉累积的影响。结果表明, 与不施氮相比施氮75 kg hm-2显著提高了穗粒数和产量, 但随施氮量的增加产量没有显著变化; 氮素对优势粒(始花日开始2 d内开花的籽粒)和劣势粒(始花日开始5~6 d间开花的籽粒)的单粒重及单粒体积、灌浆特性和单粒淀粉累积的影响趋势基本一致, 施氮条件下单粒重、单粒体积和灌浆速率随施氮量增加而增加, 但不施氮处理的单粒重和单粒体积仍高于各施氮处理, 且缺氮显著延长了灌浆活跃期。籽粒淀粉累积速率与参与籽粒淀粉合成的关键酶ADP-葡萄糖焦磷酸化酶(AGPase)和可溶性淀粉合酶(SSS)活性显著相关; 过量施氮(450 kg N hm-2)灌浆前期籽粒中AGPase和SSS的活性最高, 促进了灌浆前期籽粒淀粉累积; 施氮75 kg hm-2灌浆前期籽粒中AGPase和SSS的活性和淀粉累积速率次之; 虽然缺氮降低了灌浆前期籽粒中AGPase和SSS的活性, 但在灌浆后期维持较高活性而延长了灌浆活跃期, 因而后期具有较高的单粒淀粉累积速率, 提升了单粒淀粉累积量和单粒重。

关键词: 高粱, 氮, 优势粒, 劣势粒, 籽粒灌浆, 籽粒形态, 淀粉积累, 籽粒淀粉合成关键酶

Abstract:

To investigate the effect of N application level on grain-filling and starch accumulation in individual sorghum grains, sorghum variety Fenjiuliang 1 was used as the experimental material in 2019 and 2020. Six N rates of 0, 75, 150, 225, 300, and 450 kg N hm-2 were applied before sowing to experimental plots in Shanxi, China. To analyze sorghum grain-filling and starch accumulation by the Richards’ growth equation, the superior and inferior spikelets were sampled at seven days intervals at each sampling from anthesis to maturity. The rational N application level (75 kg N hm-2) showed the maximum grain number per panicle resulting in the maximum yield per hectare. For both superior and inferior spikelets, N had similar effects on grain weight, grain morphology structure, the characteristics of grain-filling, and grain starch accumulation. The grain weight, grain volume, and grain-filling rate increased with the increase of N application rate, whereas the maximum grain weight and grain volume was obtained with the zero N treatment. The grain starch accumulation rate was highly correlated with the activity of ADP-glucose pyrophosphorylase (AGPase) and soluble starch synthase (SSS). Compared to the zero N treatment, N application enhanced grain starch accumulation rate at early grain-filling stage and grain-filling rate, which may be due to the increased AGPase and SSS activity at the early grain-filling stage. Compared with rational N (75 kg N hm-2), excessive N (450 kg N hm-2) promoted grain starch accumulation by enhancing the activity of AGPase and SSS at the early grain filling stage, whereas zero N application enhanced sorghum grain weight and grain starch accumulation by extending the grain-filling duration and enhancing the activity of key enzymes in grain involved in sucrose-to-starch conversion at the late grain-filling stage.

Key words: sorghum, nitrogen, superior grains, inferior grains, grain filling, grain morphology, starch accumulation, starch biosynthesis enzymes

图1

2019年和2020年生长季日平均气温和降雨量"

图2

不同施氮水平下的高粱产量(a)和每穗粒数(b) N0: 施氮量为0 kg hm-2; N75: 施氮量为75 kg hm-2; N150: 施氮量为150 kg hm-2; N225: 施氮量为225 kg hm-2; N300: 施氮量为300 kg hm-2; N450: 施氮量为450 kg hm-2。不同字母表示同一年度不同处理间在0.05概率水平差异显著。"

表1

不同施氮水平下高粱籽粒特点及籽粒灌浆特性"

年份
Year		 穗位
Spikelet
position		 处理
N treatment
(kg hm-2)		 单粒重
Weight per
grain
(mg)		 籽粒体积
Volume per grain
(mm3)		 籽粒密度
Grain
density
(g L-1)		 活跃灌浆期
Active grain-filling period (d)		 灌浆速率
Grain-filling
rate
(mg grain-1 d-1)		 R2
2019 优势粒
Superior spikelets		 0 20.3±0.17 d 16.83±0.17 d 1192.9±33.2 a 36.4±0.8 c 0.601±0.020 a 0.995**
75 18.6±0.11 ab 15.00±0.50 b 1240.0±24.7 ab 33.2±1.2 b 0.604±0.016 a 0.998**
150 18.1±0.30 a 13.40±0.23 a 1327.1±11.2 c 29.2±0.4 a 0.638±0.012 b 0.996**
225 18.0±0.20 a 14.83±0.09 b 1245.5±11.2 ab 29.9±1.1 a 0.623±0.018 b 0.997**
300 18.9±0.20 bc 14.87±0.13 b 1238.7±7.9 ab 31.8±1.1 ab 0.627±0.021 b 0.996**
450 19.3±0.22 c 15.82±0.22 c 1261.0±4.1 b 31.5±0.6 ab 0.637±0.025 b 0.997**
劣势粒
Inferior spikelets		 0 18.2±0.18 c 15.50±0.29 c 1215.3±18.9 a 44.0±0.5 c 0.416±0.012 a 0.997**
75 16.7±0.07 a 14.67±0.73 bc 1192.6±43.1 a 36.9±0.9 a 0.472±0.014 cd 0.998**
150 16.7±0.10 a 12.92±0.36 a 1333.5±43.7 b 38.1±0.9 ab 0.452±0.013 b 0.996**
225 16.7±0.03 a 14.33±0.17 bc 1192.8±17.0 a 39.3±0.2 b 0.443±0.012 b 0.994**
300 16.9±0.03 a 14.00±0.12 ab 1239.4±12.4 ab 39.0±0.5 b 0.461±0.017 bc 0.995**
450 17.6±0.26 b 14.67±0.44 bc 1239.1±24.9 ab 38.6±0.2 ab 0.481±0.022 c 0.998**
2020 优势粒
Superior spikelets		 0 20.4±0.26 c 18.50±0.58 c 1103.0±41.2 a 40.6±0.3 c 0.578±0.013 a 0.996**
75 18.8±0.11 a 15.67±0.33 b 1200.8±24.9 ab 36.8±0.4 ab 0.623±0.014 b 0.997**
150 18.9±0.12 a 14.00±0.50 a 1352.0±45.4 c 36.6±0.5 ab 0.642±0.018 b 0.996**
225 18.7±0.17 a 14.83±0.60 ab 1264.2±60.8 bc 37.7±0.6 b 0.616±0.011 b 0.996**
300 19.3±0.19 ab 15.00±0.50 ab 1288.5±34.3 bc 35.5±0.4 a 0.591±0.018 b 0.997**
450 19.8±0.24 b 15.33±0.33 ab 1289.9±38.9 bc 35.7±0.4 a 0.608±0.014 b 0.995**
劣势粒
Inferior spikelets		 0 18.8±0.06 c 16.83±0.33 b 1119.4±18.1 a 46.3±0.4 d 0.400±0.009 a 0.994**
75 17.6±0.24 a 14.33±0.33 a 1232.0±30.2 b 42.4±0.3 bc 0.408±0.008 a 0.996**
150 18.3±0.28 b 14.83±0.44 a 1234.1±30.8 b 40.3±0.9 a 0.451±0.011 a 0.998**
225 18.1±0.05 ab 14.67±0.44 a 1233.3±34.0 b 40.8±0.7 ab 0.437±0.010 ab 0.996**
300 18.0±0.08 ab 14.83±0.17 a 1215.7±8.9 b 40.6±0.1 a 0.438±0.013 b 0.995**
450 19.1±0.11 c 15.17±0.17 a 1261.3±7.2 b 43.1±0.4 c 0.428±0.014 b 0.995**
方差分析ANOVA
Panicle position (P) ** ** * ** **
N treatment (N) ** ** ** ** **
Year (Y) ** ** NS ** **
N×P * NS NS NS NS
N×Y * * * NS NS
P×Y ** NS NS ** NS
N×P×Y NS NS NS ** NS

图3

不同施氮量下高粱优势粒(a, c, e, g)和劣势粒(b, d, f, h)的淀粉累积(a, b, c, d)和淀粉累积速率(e, f, g, h) 根据Richards方程拟合淀粉累积速率。处理同图2。"

表2

花后14~28 d不同施氮量对高粱优势粒淀粉相关酶活性的影响(2019年)"

花后天数 Days post-anthesis 处理
N treatment
(kg hm-2)		 酶活性Enzyme activity
ADP-葡萄糖焦磷酸化酶
ADP-glucose
pyrophosphorylase
(μmol grain-1 h-1)		 可溶性淀粉合酶
Soluble starch
synthase
(μmol grain-1 h-1)		 颗粒结合淀粉合酶
Granule-bound
starch synthase
(μmol grain-1 h-1)		 淀粉分支酶
Starch branching
enzyme
(U grain-1 min-1)
14 d N0 0.258±0.007 a 0.21±0.013 a 0.13±0.0061 b 185.8±14.2 ab
N75 0.346±0.026 a 0.27±0.011 a 0.1±0.005 a 233.6±12.4 b
N450 0.549±0.025 b 0.37±0.035 b 0.16±0.0038 c 151.3±23.1 a
21 d N0 0.211±0.007 a 0.23±0.02 ab 0.18±0.006 b 406.5±15.2 b
N75 0.227±0.009 a 0.20±0.022 a 0.14±0.003 a 318.1±21.1 a
N450 0.258±0.013 b 0.31±0.033 b 0.20±0.004 c 327.8±5.2 a
28 d N0 0.200±0.011 b 0.18±0.013 a 0.14±0.002 a 452.1±25.7 b
N75 0.105±0.041 a 0.15±0.013 a 0.13±0.005 a 441.7±8.3 b
N450 0.128±0.036 a 0.19±0.012 a 0.13±0.011 a 367.2±9.5 a

图4

不同施氮量下高粱优势粒AGPase (a)、SSS (b)、GBSS (c)和SBE (d)的酶活性(2019年) 处理同图2。"

表3

高粱优势粒淀粉快速累积时期(花后7~35 d) AGPase、SSS、GBSS和SBE的酶活性与淀粉累积速率的相关性"

酶活性
Activity of enzymes		 淀粉累积速率 Starch accumulation rate
N0 N75 N450 所有处理All treatment
ADP-葡萄糖焦磷酸化酶ADP-glucose pyrophosphorylase 0.935* 0.979** 0.950* 0.989**
可溶性淀粉合酶Soluble starch synthase 0.868 0.977** 0.976** 0.960**
颗粒结合淀粉合酶Granulebound starch synthase 0.631 0.412 0.785 0.652
淀粉分支酶Starch branching enzyme 0.220 0.240 0.065 0.166
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