摘要以江苏省近60年来各阶段具有代表性的12个中粳稻品种(含超级稻)为材料。依据种植推广年代结合株型和基因型将其分为20世纪50年代、60年代、70年代、80年代、90年代和2000年以后6个类型, 进行0N (不施氮, 0N)、240 kg hm-2(中氮, MN)和360 kg hm-2(高氮, HN) 3种氮肥用量处理, 并观察其对水稻产量和群体质量的影响。结果表明, 产量随品种改良逐步提高, 在0N、MN和HN条件下, 由20世纪50年代早期品种到2000年以后的超级稻品种, 产量增幅分别为4.45~4.64、5.89~5.93和8.45~8.62 t hm-2。2000年以前的中粳稻品种产量表现为MN>HN>0N; 2000年以后的超级稻品种则表现为HN>MN>0N。2000年以前的中粳稻品种在MN处理下具有较高群体质量指标(茎蘖成穗率、抽穗至成熟期的干物质积累、有效叶面积和高效叶面积指数、粒叶比、着粒密度、剑叶光合速率和根系氧化力), 2000年以后的超级稻品种则在高氮处理下具有较高的群体质量指标。这些结果表明, 早期品种对氮肥响应较现代超级稻品种敏感, 超级稻品种则在高氮水平下具有更高的产量。群体质量的改善是品种改良增加产量以及超级稻品种在高氮水平下物质生产和产量提高的重要原因。
关键词:粳稻; 品种改良; 氮肥; 产量; 群体质量; 超级稻 Effects of Nitrogen Application Rate on Grain Yield and Population Quality of Mid-seasonJaponicaRice Cultivars at Different Decades in Jiangsu Province CHEN Lu, ZHANG Wei-Yang, WANG Zhi-Qin, ZHANG Hao, LIU Li-Jun, YANG Jian-Chang* Key Laboratory of Crop Genetics and Physiology of Jiangsu Province, Yangzhou University, Yangzhou 225009, China Fund: AbstractGreat progress has been made in the mid-seasonjaponica rice breeding, but little information is available on understanding how nitrogen (N) application rate affects population quality and grain yield during the improvement of cultivars. In this study, 12 typical mid-seasonjaponica rice cultivars (including super rice) applied in the production in Jiangsu Province during the last 60 years were used, and classified into six types of 1950s, 1960s, 1970s, 1980s, 1990s, and after 2000 according to their application times. Three N rates, i.e. zero N (0N), 240 kg ha-1 N (MN), and 360 kg ha-1 N (HN), were applied. Grain yield and some population quality indexes, such as grain-leaf ratio, percentage of productive tillers, leaf photosynthetic rate and root oxidation activity, were determined. The results showed that from 1950s to the present time, the grain yield was increased by 4.45-4.64, 5.89-5.93, and 8.45-8.62 t ha-1 in treatments of 0N, MN, and HN, respectively. The grain yield was progressively increased with the improvement of cultivars, showing a trend of MN > HN > 0N for the cultivars bred in the 20th century, and HN > MN > 0N for the super rice cultivars bred in the 21st century. The percentage of productive tillers, dry matter accumulation from heading to maturity, effective and high effective leaf area index, grain-leaf ratio, leaf photosynthetic rate, and root oxidation activity were the highest at the MN rate for cultivars bred in the 20th century, while at the HN rate for cultivars bred in the 21st century, among the three N rates. The results suggest that the response to N rates is more sensitive for the cultivars bred in the early decades than for the modern super rice cultivars, and the latter could produce a higher grain yield at the HN rate. Enhancement in population quality contributes to the increase in biomass and grain yield in the improvement of rice cultivars and for the super rice at the HN rate.
Keyword:Japonica rice; Improvement of cultivars; Nitrogen fertilizer; Grain yield; Population quality; Super rice Show Figures Show Figures
表2 不同处理下水稻的产量、成熟期干物重、抽穗期总叶面积指数、颖花/叶粒叶比、叶基角、株高和着粒密度的年度与品种间的方差分析 Table 2 Analysis-of-variance of F-values of grain yield, shoot biomass at maturity, total leaf area index at heading, top three leaves angel 10 days after heading, plant height at mature and grains per panicle of rice
变异来源 Source of variation
自由度 df
产量 Yield
颖花/叶 Spikelets/leaf (cm2)
成熟期干物重 Shoot biomass
株高 Plant height
着粒密度 Grains per panicle
品种Variety (V)
11
1416**
313.2**
208.4**
1006**
4215**
氮处理 N treatment (N)
2
6402**
779.2**
3019**
706.3**
777.0*
年度Year (Y)
1
NS
NS
NS
NS
NS
品种×处理(V×N)
22
71.5**
27.6**
12.5**
18.4**
58.3**
年度×品种(Y×V)
11
NS
NS
NS
NS
NS
年度×处理(Y×N)
2
NS
NS
NS
NS
NS
*表示在0.05水平上差异显著;**表示在0.01水平上差异显著; NS在表示0.05水平下不显著。 *,** significant at 0.05 and 0.01 probability levels, respectively. NS, not significant at 0.05 probability level.
表2 不同处理下水稻的产量、成熟期干物重、抽穗期总叶面积指数、颖花/叶粒叶比、叶基角、株高和着粒密度的年度与品种间的方差分析 Table 2 Analysis-of-variance of F-values of grain yield, shoot biomass at maturity, total leaf area index at heading, top three leaves angel 10 days after heading, plant height at mature and grains per panicle of rice
2.2 不同氮肥水平下水稻产量及其构成因素在全生育期总施氮量0~360 kg hm-2条件下, 产量均随品种改良大幅度提高(图1-A, B)。当总施氮量为0、240和360 kg hm-2时, 由20世纪50年代早期品种到90年代品种, 产量增幅分别为3.71~ 3.72、4.71~4.96和4.91~4.94 t hm-2左右。在这3种施氮量条件下, 2000年以后的超级稻品种又较20世纪90年代品种分别增产14.33%~15.75%、10.83%~13.49%、42.96%~45.30%。表明超级稻的产量又有了大幅度的提高。在全生育期总施氮量 240 kg hm-2条件下, 2000年以前的中粳稻品种产量较不施氮处理显著增加, 当施氮量增加至 360 kg hm-2时, 产量则呈现下降的趋势, 表现为MN>HN>0N; 对于2000年以后的超级稻品种, 在施氮量0~360 kg hm-2条件下, 产量随着施氮量的增加而增加, 表现为HN>MN>0N。与不施氮肥相比, 施用240 kg hm-2氮肥后, 50年代的早期中粳稻品种的产量增加了86.05%~90.57%, 2000年以后的超级稻品种的产量增加了46.89%~48.21% (图1-A, B)。说明早期品种较超级稻品种对氮肥的响应更敏感。 图1 Fig. 1
图1 江苏省不同氮肥水平下中粳稻品种改良过程中产量及产量构成因素的变化0N: 全生育期不施氮肥; MN: 全生育期施氮240 kg hm-2; HN: 全生育期施氮360 kg hm-2。Fig. 1 Changes of grain yield and its components for japonica rice cultivars in Jiangsu under different nitrogen application rates0N: no nitrogen was applied; MN: 240 kg hm-2 nitrogen was applied; HN: 360 kg hm-2 nitrogen was applied.
图2 江苏省不同氮肥水平下中粳稻品种改良过程中茎蘖动态变化0N: 全生育期不施氮肥; MN: 全生育期施氮240 kg hm-2; HN: 全生育期施氮360 kg hm-2。Fig. 2 Changes of the number of tillers for japonica rice cultivars in Jiangsu under different nitrogen application rates0N: no nitrogen was applied; MN: 240 kg hm-2 nitrogen was applied; HN: 360 kg hm-2 nitrogen was applied.
2.4 不同氮肥水平下水稻干物质积累各年代品种在各生育时期的干物质积累均随着品种改良有显著提高(图3)。在同一施氮量, 各年代品种的干物质积累在分蘖中期差异不显著, 而穗分化始期、抽穗期和成熟期的干物质积累均随着品种改良显著增加。与20世纪50年代的早期中粳稻品种相比, 当施氮量分别为0、240和360 kg hm-2时, 2000年以后的超级稻品种全生育期总干物质增幅分别为7.60、10.15和14.55 t hm-2。2000年以前的中粳稻品种, 总生物产量均随着施氮量的增加呈现先增加后降低的趋势, 表现为MN>HN>0N, 而2000年以后的超级稻品种, 总生物产量随着施氮量的增加而增加, 表现为HN>MN>0N (图3)。 图3 Fig. 3
图3 江苏省不同氮肥水平下中粳稻品种改良过程中干物质积累变化0N: 全生育期不施氮肥; MN: 全生育期施氮240 kg hm-2; HN: 全生育期施氮360 kg hm-2; MT: 分蘖中期; PI: 穗分化始期; HD: 抽穗期; MA: 成熟期。Fig. 3 Changes of the biomass for japonica rice cultivars in Jiangsu under different nitrogen application rates0N: no nitrogen was applied; MN: 240 kg hm-2 nitrogen was applied; HN: 360 kg hm-2 nitrogen was applied; MT: mid tillering; PI: panicle initiation; HD: heading; MA: maturity.
表3 江苏省不同氮肥水平下中粳稻品种改良过程中叶面积指数(LAI)的变化 Table 3 Changes of the leaf area index(LAI) for japonica rice cultivars in Jiangsu under different nitrogen application rates
品种类型 Cultivar type
品种 Cultivar
总LAI Total leaf area index
高效LAI High-effective leaf area index
有效LAI Effective leaf area index
0N
MN
HN
0N
MN
HN
0N
MN
HN
1950s
黄壳早Huangkezao
2.51 de
4.72 fg
5.27 de
1.22 e
2.57 g
2.12 e
1.68 d
3.56 g
2.97 f
桂花球Guihuaqiu
2.25 e
4.38 g
3.80 f
1.12 e
2.39 g
1.90 e
1.53 d
3.30 g
2.66 f
平均Average
2.38
4.55
4.54
1.17
2.48
2.01
1.60
3.43
2.81
1960s
金南风Jinnanfeng
2.46 de
5.45 e
4.96 e
1.79 d
3.22 f
2.84 d
2.53 c
4.37 f
3.84 de
桂花黄Guihuahuang
3.37 bc
5.19 ef
5.22 de
1.74 d
3.11 f
2.73 d
2.47 c
4.16 f
3.68 e
平均Average
2.92
5.32
5.09
1.76
3.19
2.79
2.50
4.26
3.76
1970s
徐稻2号Xudao 2
3.34 b
5.96 d
5.68 d
2.37 c
3.83 e
3.13 d
3.25 b
5.03 e
4.27 d
黎明Liming
2.90 cd
5.33 e
5.55 d
1.75 d
3.12 f
2.88 d
2.40 c
4.10 f
3.92 de
平均Average
3.12
5.65
5.62
2.06
3.48
3.01
2.83
4.56
4.09
1980s
泗稻8号Sidao 8
4.30 a
6.90 bc
6.56 bc
2.52 c
4.27 de
3.86 c
3.40 b
5.62 d
5.11 c
盐粳2号Yanjing 2
4.01 a
6.70 c
6.67 bc
3.02 b
4.60 cd
4.01 c
4.06 a
6.06 cd
5.31 c
平均Average
4.16
6.80
5.62
2.77
4.43
3.93
3.73
5.84
5.21
1990s
镇稻88 Zhendao 88
4.05 a
6.57 c
6.28 c
3.23 b
5.04 bc
4.60 b
3.96 a
6.32 bc
6.00 b
淮稻5号Huaidao 5
4.29 a
6.97 bc
6.98 b
3.38 b
5.26 ab
4.88 b
4.16 a
6.61 b
6.37 b
平均Average
4.17
6.77
6.63
3.30
5.15
4.74
4.06
6.47
6.19
2000-
淮稻9号Huaidao 9
4.06 a
7.72 a
8.81 a
3.92 a
5.77 a
6.34 a
3.97 a
7.82 a
8.25 a
连粳7号Lianjing 7
4.41 a
7.36 ab
8.88 a
4.00 a
5.61 a
6.02 a
4.27 a
7.61 a
7.82 a
平均Average
4.24
8.54
8.85
3.96
5.69
6.18
4.12
7.72
8.03
0N: 全生育期不施氮肥; MN: 全生育期施氮240 kg hm-2; HN: 全生育期施氮360 kg hm-2。同一栏内标以不同字母的值在0.05水平上差异显著。 0N: no nitrogen was applied; MN: 240 kg hm-2 nitrogen was applied; HN: 360 kg hm-2 nitrogen was applied. Values followed by different letters within the same column are significantly different at 0.05 probability level.
表3 江苏省不同氮肥水平下中粳稻品种改良过程中叶面积指数(LAI)的变化 Table 3 Changes of the leaf area index(LAI) for japonica rice cultivars in Jiangsu under different nitrogen application rates
表4 江苏省不同氮肥水平下中粳稻品种改良过程中粒叶比的变化 Table 4 Changes of grain-leaf ratio for japonica rice cultivars in Jiangsu under different nitrogen application rates
品种类型 Cultivar type
品种 Cultivar
颖花/叶 Spikelet/leaf (cm2)
实粒/叶 Filled grains/leaf (cm2)
粒重/叶 Grain yield (mg)/leaf (cm2)
0N
MN
HN
0N
MN
HN
0N
MN
HN
1950s
黄壳早Huangkezao
0.50 d
0.58 b
0.51 d
0.30 g
0.31 g
0.27 f
6.61 i
6.62 h
5.45 h
桂花球Guihuaqiu
0.50 d
0.58 b
0.52 cd
0.39 ef
0.42 e
0.36 e
11.75 f
11.13 e
9.48 e
平均Average
0.50
0.58
0.52
0.34
0.37
0.31
9.18
8.87
7.46
1960s
金南风Jinnanfeng
0.60 bc
0.64 a
0.63 b
0.38 f
0.39 f
0.35 e
10.10 h
9.45 g
8.36 g
桂花黄Guihuahuang
0.50 d
0.54 c
0.49 d
0.43 e
0.43 de
0.37 e
10.85 g
10.36 f
8.92 f
平均Average
0.55
0.59
0.56
0.40
0.41
0.36
10.47
9.91
8.64
1970s
徐稻2号Xudao 2
0.60 bc
0.63 a
0.61 b
0.53 a
0.53 a
0.46 bc
13.08 d
12.31 c
10.62 c
黎明Liming
0.52 d
0.57 c
0.55 c
0.47 cd
0.47 bc
0.42 d
12.27 e
11.68 d
10.00 d
平均Average
0.56
0.60
0.58
0.50
0.50
0.44
12.68
11.99
10.31
1980s
泗稻8号Sidao 8
0.57 c
0.64 a
0.61 b
0.46 d
0.46 bcd
0.43 cd
11.29 fg
11.11 e
10.37 cd
盐粳2号Yanjing 2
0.57 c
0.63 bc
0.61 b
0.49 bcd
0.50 b
0.45 bcd
12.53 e
11.66 d
10.56 c
平均Average
0.57
0.64
0.61
0.48
0.48
0.44
11.91
11.39
10.47
1990s
镇稻88 Zhendao 88
0.57 c
0.65 a
0.63 b
0.52 ab
0.54 a
0.50 a
14.26 c
13.62 a
12.94 a
淮稻5号Huaidao 5
0.58 c
0.63 a
0.61 b
0.50 abc
0.50 b
0.45 bcd
13.87 c
12.86 b
11.77 b
平均Average
0.58
0.64
0.62
0.51
0.52
0.48
14.07
13.24
12.35
2000-
淮稻9号Huaidao 9
0.65 a
0.63 a
0.69 a
0.52 ab
0.45 cde
0.48 ab
16.15 a
13.66 a
13.07 a
连粳7号Lianjing 7
0.63 ab
0.64 a
0.69 a
0.52 ab
0.48 bc
0.51 a
15.55 b
13.70 a
13.35 a
平均Average
0.64
0.64
0.69
0.52
0.47
0.49
15.85
13.68
13.21
0N: 全生育期不施氮肥; MN: 全生育期施氮240 kg hm-2; HN: 全生育期施氮360 kg hm-2。同一栏内标以不同字母的值在0.05水平上差异显著。 0N: no nitrogen was applied; MN: 240 kg hm-2 nitrogen was applied; HN: 360 kg hm-2 nitrogen was applied. Values followed by different letters within the same column are significantly different at 0.05 probability level.
表4 江苏省不同氮肥水平下中粳稻品种改良过程中粒叶比的变化 Table 4 Changes of grain-leaf ratio for japonica rice cultivars in Jiangsu under different nitrogen application rates
图4 江苏省不同氮肥水平下中粳稻品种改良过程中株高(A)与叶基角(B)的变化0N: 全生育期不施氮肥; MN: 全生育期施氮240 kg hm-2; HN: 全生育期施氮360 kg hm-2。Fig. 4 Changes of plant height (A) and top three leaves angle (B) for japonica rice cultivars in Jiangsu under different nitrogen application rates0N: no nitrogen was applied; MN: 240 kg hm-2 nitrogen was applied; HN: 360 kg hm-2 nitrogen was applied.
图5 江苏省不同氮肥水平下中粳稻品种改良过程中根系氧化力(A~C)与光合速率(D~F)的变化0N: 全生育期不施氮肥; MN: 全生育期施氮240 kg hm-2; HN: 全生育期施氮360 kg hm-2; PI: 穗分化始期; HD: 抽穗期; MF: 灌浆中期。Fig. 5 Changes of root oxidation activity (A-C) and photosynthetic rate (D-F) for japonica rice cultivars in Jiangsu under different nitrogen application rates0N: no nitrogen was applied; MN: 240 kg hm-2 nitrogen was applied; HN: 360 kg hm-2 nitrogen was applied; PI: panicle initiation; HD: heading; MF: middle grain filling stage.
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表1 供试中粳稻品种 Table 1 The tested japonica rice cultivars
