摘要以新稻20为材料进行土培试验, 设置浅水层灌溉(0 kPa)、轻度水分胁迫(-20 kPa)和重度水分胁迫(-40 kPa) 3种灌溉方式及0氮(0N, 0 kg hm-2)、中氮(MN, 240 kg hm-2)和高氮(HN, 360 kg hm-2) 3种氮水平, 研究不同水氮耦合处理对水稻根长、根冠比、根系伤流、根系有机酸含量、根系玉米素及玉米素核苷与籽粒酶活性的影响。结果表明, 灌溉方式与施氮量存在显著的互作效应, 轻度水分胁迫增加了主要生育期根长、根系伤流量、根系分泌物中有机酸总量、根系玉米素及玉米素核苷含量, 提高籽粒ATP酶、蔗糖合酶及腺苷二磷酸葡萄糖焦磷酸化酶活性, 降低穗分化后水稻根冠比, 且与MN耦合后产量最高, 为本试验最佳的水氮耦合运筹模式; 重度水分胁迫则显著降低主要生育期根长、根系伤流量、根系分泌物中有机酸总量、根系玉米素及玉米素核苷含量, 降低籽粒ATP酶、蔗糖合酶及腺苷二磷酸葡萄糖焦磷酸化酶活性, 增加主要生育期根冠比。水稻籽粒产量与主要生育期水稻根长、根系伤流量、根系分泌物中有机酸总量、根系玉米素及玉米素核苷含量均呈显著或极显著的正相关, 而穗分化至成熟期根冠比与水稻产量呈负相关; 同时水稻根长、根系伤流量、根系分泌物中有机酸总量、根系玉米素及玉米素核苷含量与籽粒ATP酶、蔗糖合酶及腺苷二磷酸葡萄糖焦磷酸化酶活性呈显著或极显著的正相关。表明通过适宜的肥水调控发挥水氮耦合效应, 可以创造良好的根系形态、提高水稻根系代谢能力和籽粒库的生理活性, 促进水稻高产。
关键词:水稻; 水氮耦合; 产量; 根系特性; 籽粒库活性 Effect of Wetting and Drying Alternative Irrigation Coupling with Nitrogen Application on Root Characteristic and Grain-sink Activity XU Guo-Wei1,2, LYU Qiang1, LU Da-Ke1, WANG He-Zheng1, CHEN Ming-Can1 1 Agricultural College, Henan University of Science and Technology, Luoyang 471003, China
2 Key Laboratory of Crop Genetics and Physiology of Jiangsu Province, Yangzhou University, Yangzhou 225009, China
Fund:This study was supported by the National Natural Science Foundation of China (U1304316), the Open Foundation for Key Laboratory of Cultivation and Physiology of Jiangsu Province (027388003K11009), and the Key Project for Science and Technology Research of Henan Provincial Department of Education (13A210266) AbstractSoil moisture and nitrogen nutrient are the two principal factors affecting rice production. Elucidation of their coupling effects on grain yield of rice ( Oryza sativa L.) has great significance for high yielding and high efficiency in production. The purposes of this study were to investigate the coupling effects of water and nitrogen on root traits and grain-sink activity. A field experiment was conducted using a mid-season japonica rice cultivar of Xindao 20 with three treatments of different nitrogen levels, including 0N, MN (240 kg ha-1) and HN (360 kg ha-1), and three irrigation regimes, including submerged irrigation (0 kPa), alternate wetting and moderate drying (-20 kPa) and alternate wetting and severe drying (-40 kPa) in 2013 and 2014. There was a significant interaction between irrigation regimes and nitrogen applications, with a similar result in two years. The grain yield was the highest in the treatment of MN coupling with mild water stress due to improved seed filling rate and grain weight. In the same nitrogen level, root length, root bleeding, organic acid and Z+ZR contents in roots at main growth stages were higher in treatment of alternate wetting and moderate drying than in treatment of submerged irrigation, meanwhile activating of ATPase, sucrose synthase and adenosine phosphate glucose pyrophosphorylase (AGPase) activity in grain were also increased during grain filling period, but root-shoot ratio was lower after panicle initiation stage. The treatment of mild water stress and MN enhanced rice yield and nitrogen use efficiency, being the best water-nitrogen coupling management model in this paper. The opposite result was observed under the condition of alternate wetting and severe drying. Grain yield positively correlated with root length and root metabolism indices at main growth stages and negatively correlated with ratio of root to shoot at the stage of heading. A negative correlation was observed between root-shoot ratio and activating of ATPase, sucrose synthase and AGPase in grain, while a significant or very significant and positive correlation between root length, root metabolism traits and aboveground development. These results suggest that a good root morphology, increasing root metabolism and aboveground development capabilities through the appropriate regulation of water coupling with nitrogen application will be much more beneficial to increasing grain yield in rice.
Keyword:Rice; Water and nitrogen coupling; Yield; Root characteristic; Sink activity in grain Show Figures Show Figures
表1 水氮耦合下产量及抽穗期主要根系性状的方差分析 Table 1 Analysis of variance of rice yield and main root traits under the condition of water and nitrogen coupling
变异来源 Source of variation
自由度 df
产量 Grain yield (g m-2)
根长 Root length (m hill-1)
根冠比 Root-shoot ratio
根系伤流量 Root bleeding (mg h-1plant-1)
根系氧化力 Root oxidation activity (μ g α -NA h-1 g-1)
根系有机酸总量 Root total organic acid content (mg g-1)
根系细胞分裂素含量 Z+ZR content (pmol g-1)
Y
1
NS
NS
NS
NS
NS
NS
NS
W
2
387.5* *
362.6* *
18.7* *
274.1* *
421.5* *
38.7* *
564.7* *
N
2
562.3* *
1002.7* *
42.4* *
536.3* *
358.4* *
152.5* *
379.2* *
Y× W
2
NS
NS
NS
NS
NS
NS
NS
Y× N
2
NS
NS
NS
NS
NS
NS
NS
W× N
4
19.5* *
34.7* *
3.0*
31.4* *
19.9* *
4.6*
9.5* *
Y× W× N
4
NS
NS
NS
NS
NS
NS
NS
NS: not significant (P > 0.05). * and * * represent P< 0.05 and P < 0.01, respectively. The indicator data were determined at heading stage, expect for grain yield. Y, W, and N represent year, water and nitrogen, respectively. NS表示在0.05水平上差异不显著。* 与* * 表示在0.05和0.01水平上差异显著与极显著。除产量外其余指标均为抽穗期测定数据。Y表示年度间, W表示水分处理, N表示施氮量。
表1 水氮耦合下产量及抽穗期主要根系性状的方差分析 Table 1 Analysis of variance of rice yield and main root traits under the condition of water and nitrogen coupling
表2 水氮耦合对水稻产量及产量构成因素的影响 Table 2 Effect of water and nitrogen coupling on yield and its components in whole growing period
年代 Year
处理 Treatment
单位面积穗数 Panicles m-2
每穗粒数 Spikelet per panicle
结实率 Filled grain rate (%)
千粒重 1000-grain weight (g)
产量 Yield (g m-2)
2014
0N (0 kPa)
256.7 d
123.3 bc
84.2 b
25.0 c
666.2 d
0N (-20 kPa)
252.7 d
118.0 c
87.6 a
25.7 b
671.2 d
0N (-40 kPa)
226.0 e
112.5 d
81.7 bc
24.8 c
515.2 f
MN (0 kPa)
390.7 ab
131.0 a
77.9 d
25.6 b
1020.6 ab
MN (-20 kPa)
386.7 ab
126.1 ab
81.4 bc
26.3 a
1043.8 a
MN (-40 kPa)
356.0 c
116.3 c
76.2 d
25.1 c
791.9 c
HN (0 kPa)
406.7 a
134.3 a
72.0 e
25.1 c
987.0 b
HN (-20 kPa)
393.3 a
130.5 a
76.4 d
25.8 b
1011.8 ab
HN (-40 kPa)
364.0 c
102.9 e
65.1 f
24.7 c
602.3 e
2013
0N (0 kPa)
260.8 d
120.7 cd
82.9 b
25.3 c
660.2 d
0N (-20 kPa)
254.1 e
118.9 d
86.7 a
25.8 b
675.8 d
0N (-40 kPa)
230.8 f
110.3 e
81.6 bc
24.9 d
517.3 f
MN (0 kPa)
415.7 a
134.0 ab
79.1 cd
25.4 bc
1119.2 ab
MN (-20 kPa)
411.3 ab
128.0 bc
81.0 bc
26.5 a
1130.1 a
MN (-40 kPa)
381.9 c
114.1 de
73.7 ef
24.9 d
799.7 c
HN (0 kPa)
430.1 a
142.6 a
70.1 f
25.0 cd
1074.8 b
HN (-20 kPa)
420.9 a
137.2 a
74.1 e
25.9 b
1108.3 ab
HN (-40 kPa)
384.1 c
101.1 f
63.4 g
24.6 de
605.6 e
0N: no nitrogen applied; MN: nitrogen 240 kg hm-2; HN: nitrogen 360 kg hm-2. Values within the same column followed by different letters are significantly different at P< 0.05. 0N: 不施氮肥; MN: 240 kg N hm-2; HN: 360 kg N hm-2; 同一列标以不同字母的值在0.05水平上差异显著。
表2 水氮耦合对水稻产量及产量构成因素的影响 Table 2 Effect of water and nitrogen coupling on yield and its components in whole growing period
表3 Table 3 表3(Table 3)
表3 水氮耦合对水稻根长及根冠比的影响 Table 3 Effect of water and nitrogen coupling on root length and root-shoot ratio in whole growing period
处理 Treatment
根长 Root length ( m hill-1)
根冠比 Root-shoot ratio
分蘖盛期Mid-tillering
穗分化始期 PI
抽穗期 Heading
成熟期 Maturity
分蘖盛期Mid-tillering
穗分化始期 PI
抽穗期 Heading
成熟期 Maturity
0N (0 kPa)
36.4 g
75.5 f
69.4 ef
41.0 e
0.30 e
0.21 de
0.19 b
0.03 a
0N (-20 kPa)
50.2 f
89.0 e
77.7 e
48.8 d
0.29 e
0.22 cd
0.17 cd
0.03 a
0N (-40 kPa)
30.7 h
63.4 g
54.3 g
32.8 f
0.22 f
0.24 b
0.21 a
0.03 a
MN (0 kPa)
84.2 c
198.5 b
169.8 b
89.8 b
0.35 d
0.21 de
0.15 ef
0.03 a
MN (-20 kPa)
91.3 ab
209.1 a
181.4 a
102.9 a
0.36 d
0.21 de
0.14 f
0.03 a
MN (-40 kPa)
68.8 d
144.3 c
131.8 c
71.4 c
0.41 ab
0.24 b
0.17 cd
0.03 a
HN (0 kPa)
89.2 bc
203.3 b
174.9 ab
96.5 b
0.38 c
0.23 bc
0.17 cd
0.03 a
HN (-20 kPa)
96.6 a
212.5 a
184.3 a
105.5 a
0.39 bc
0.23 bc
0.15 ef
0.04 a
HN (-40 kPa)
59.3 e
127.1 d
106.9 d
65.9 c
0.43 a
0.26 a
0.19 b
0.04 a
0N: no nitrogen applied; MN: nitrogen 240 kg hm-2; HN: nitrogen 360 kg hm-2; PI: panicle initiation. Values within the same column followed by different letters are significantly different at P< 0.05. 0N: 不施氮肥; MN: 240 kg N hm-2; HN: 360 kg N hm-2; 同一列不同字母表示在0.05水平上差异显著。
表3 水氮耦合对水稻根长及根冠比的影响 Table 3 Effect of water and nitrogen coupling on root length and root-shoot ratio in whole growing period
图1 水氮耦合对水稻生育过程中根系伤流强度的影响0N: 不施氮肥; MN: 施氮240 kg· hm-2; HN: 施氮360 kg hm-2; PI: 幼穗分化始期; 20DAA: 抽穗后20 d; 同一生育期柱上不同字母表示各处理在0.05水平上差异显著。Fig. 1 Effect of water and nitrogen coupling on root bleeding intensity during the growth and development of rice cultivars0N: no nitrogen applied; MN: nitrogen 240 kg hm-2; HN: nitrogen 360 kg hm-2; PI: initiation of panicle differentiation; 20DAA: 20 days after anthesis; Bars within the same growth period represented by different letters are significantly different at P< 0.05.
图2 水氮耦合对水稻生育过程中根系有机酸总量的影响0N: 不施氮肥; MN: 施氮240 kg· hm-2; HN: 施氮360 kg hm-2; PI: 幼穗分化始期; 20DAA: 抽穗后20 d。同一生育期不同字母表示各处理在0.05水平上差异显著。Fig. 2 Effect of water and nitrogen coupling on root organic acid contents during growth and development of rice0N: no nitrogen applied; MN: nitrogen 240 kg hm-2; HN: nitrogen 360 kg hm-2; PI: initiation of panicle differentiation; 20DAA: 20 days after anthesis. Bars within the same growth period represented by different letters are significantly different at P < 0.05.
图3 水氮耦合对水稻生育过程中根系玉米素及玉米素核苷含量的影响0N: 不施氮肥; MN: 施氮240 kg hm-2; HN: 施氮360 kg hm-2; PI: 幼穗分化始期; 20DAA: 抽穗后20 d。同一生育期不同字母表示各处理在0.05水平上差异显著。Fig. 3 Effect of water and nitrogen coupling on root Z+ZR content during growth and development of rice cultivars0N: no nitrogen applied; MN: nitrogen 240 kg hm-2; HN: nitrogen 360 kg hm-2; PI: initiation of panicle differentiation; 20DAA: 20 days after anthesis. Bars within the same growth period represented by different letters are significantly different at P < 0.05.
表4 全生育期水氮耦合对水稻灌浆期籽粒酶活性的影响 Table 4 Effect of water and nitrogen coupling on enzyme activities in grain at filling stage
处理 Treatment
抽穗期 Heading stage
花后20 d 20 DAA
ATP酶 ATPase (μ mol Pi mg-1 Pro h-1)
蔗糖合酶 Sucrose synthase (μ mol g-1 FW min-1)
腺苷二磷酸葡萄糖焦磷酸化酶 AGPase (μ mol g-1 FW min-1)
ATP酶 ATPase (μ mol Pi mg-1 Pro h-1)
蔗糖合酶 Sucrose synthase (μ mol g-1 FW min-1)
腺苷二磷酸葡萄糖焦磷酸化酶 AGPase (μ mol g-1 FW min-1)
0N (0 kPa)
3.8 ef
8.3 cd
6.1 e
3.0 d
3.8 e
2.1 g
0N (-20 kPa)
4.4 cd
9.0 c
6.5 d
3.5 c
4.7 d
2.6 f
0N (-40 kPa)
3.3 g
7.6 de
5.6 f
2.6 e
3.1 f
1.8 h
MN (0 kPa)
5.2 b
11.2 b
7.5 bc
4.3 b
5.8 b
3.8 c
MN (-20 kPa)
6.1 a
13.1 a
7.9 a
5.0 a
6.4 a
4.6 a
MN (-40 kPa)
4.6 c
8.8 cd
6.1 e
3.8 c
4.6 d
3.1 e
HN (0 kPa)
4.8 bc
10.4 b
7.2 c
3.7 c
5.2 c
3.4 d
HN (-20 kPa)
5.7 a
12.6 a
7.7 ab
4.4 b
5.9 b
4.3 b
HN (-40 kPa)
4.1 de
8.0 d
5.8 ef
3.1 d
3.8 e
2.5 f
0N: no nitrogen applied; MN: nitrogen 240 kg hm-2; HN: nitrogen 360 kg hm-2; PI: initiation of panicle differentiation; 20DAA: 20 days after anthesis. Values within a column followed by different letters are significantly different at P< 0.05. 0N: 不施氮肥; MN: 施氮240 kg hm-2; HN: 施氮360 kg· hm-2; PI: 幼穗分化始期; 20DAA: 抽穗后20 d。同一列中标以不同字母的值在0.05水平上差异显著。
表4 全生育期水氮耦合对水稻灌浆期籽粒酶活性的影响 Table 4 Effect of water and nitrogen coupling on enzyme activities in grain at filling stage
表5 Table 5 表5(Table 5)
表5 根系指标与产量及籽粒库活性的相关关系分析 Table 5 Correlation coefficients of yield and enzyme activities in grain with root traits at different growth stages
根系性状 Root trait
与产量相关系数 Correlation coefficients of yield with
与籽粒库活性部相关 Correlation coefficients between sink activity
分蘖盛期 Mid-tillering
穗分化始期 PI
抽穗期 Heading
成熟期 Maturity
ATP酶 ATPase (μ mol Pi mg-1 Pro h-1)
蔗糖合酶 Sucrose synthase (μ mol g-1 FW min-1)
腺苷二磷酸葡萄糖焦磷酸化酶 AGPase (μ mol g-1 FW min-1)
根长 Root length
0.821*
0.837*
0.852*
0.811*
0.822*
0.722
0.672
根冠比 Root-shoot ratio
0.156
-0.312
-0.827*
-0.018
-0.944*
-0.847*
-0.854*
根系伤流量 Root bleeding
0.669
0.933* *
0.931* *
—
0.908* *
0.979* *
0.972* *
根系有机酸 Root organic acid
0.805*
0.932* *
0.931* *
—
0.908* *
0.980* *
0.971* *
根系生长素 Z+ZR in root
0.684
0.920* *
0.911* *
—
0.903* *
0.906* *
0.899* *
“ — ” no data available. 0N: no nitrogen applied; MN: nitrogen 240 kg hm-2; HN: nitrogen 360 kg hm-2; PI: initiation of panicle differentiation; 20DAA: 20 days after anthesis. * Significance of correlation at P < 0.05.* * Significance of correlation at P < 0.01. “ — ” : 未测定; 0N: 不施氮肥; MN: 施氮240 kg hm-2; HN: 施氮360 kg hm-2; PI: 幼穗分化始期; 20DAA: 抽穗后20 d。* 相关在0.05水平显著; * * 相关在0.01水平显著。
表5 根系指标与产量及籽粒库活性的相关关系分析 Table 5 Correlation coefficients of yield and enzyme activities in grain with root traits at different growth stages
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表1 水氮耦合下产量及抽穗期主要根系性状的方差分析 Table 1 Analysis of variance of rice yield and main root traits under the condition of water and nitrogen coupling
