光、氮及其互作对超级粳稻产量和氮、磷、钾吸收的影响

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王亚江, 魏海燕, 颜希亭, 葛梦婕, 孟天瑶, 张洪程, 戴其根, 霍中洋, 许轲, 费新茹. 光、氮及其互作对超级粳稻产量和氮、磷、钾吸收的影响. 作物学报, 2014, 40(7): 1235-1244
WANG Ya-Jiang, WEI Hai-Yan, YAN Xi-Ting, GE Meng-Jie, MENG Tian-Yao, ZHANG Hong-Cheng, DAI Qi-Gen, HUO Zhong-Yang, XU Ke, FEI Xin-Ru. Effects of Light, Nitrogen and Their Interaction on Grain Yield and Nitrogen, Phosphorus and Potassium Absorption inJaponica Super Rice . Acta Agronomica Sinica, 2014, 40(7): 1235-1244 复制到剪切板

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光、氮及其互作对超级粳稻产量和氮、磷、钾吸收的影响

王亚江, 魏海燕^*, 颜希亭, 葛梦婕, 孟天瑶, 张洪程, 戴其根, 霍中洋, 许轲, 费新茹

扬州大学农学院江苏省作物遗传生理国家重点实验室 / 农业部长江流域稻作技术创新中心, 江苏扬州 225009

^* 通讯作者(Corresponding author): 魏海燕, E-mail:wei_haiyan@163.com, Tel: 0514-87979220

第一作者联系方式: E-mail:wyj19900120@163.com

收稿日期:2014-04-16

基金:本研究由国家自然科学基金项目(31101102和30971732), 高等学校博士学科点专项科研基金项目(20103250120003), 国家粮食丰产科技工程项目(2011BAD16B03)和江苏高校优势学科建设工程项目资助;

摘要

大田条件下以超级粳稻南粳44和宁粳3号为材料, 设置2种氮肥水平(N10: 150 kg hm^-2, N20: 300 kg hm^-2)和3种遮光处理(L1: 不遮光, L2: 抽穗前遮光20 d, L3: 抽穗后遮光20 d), 研究光、氮及其互作对超级粳稻产量和氮、磷、钾吸收的影响。结果表明, 同一氮肥水平下产量呈现L1>L3>L2。其中, L2使植株在拔节至抽穗阶段及抽穗期的氮、磷、钾积累量显著下降, 主要由于L2显著降低了抽穗前期的根系α-NA氧化量及根干重, 导致根系吸收养分能力下降, 最终产量显著低于L1, 达30.58%~35.26%。L3使植株在抽穗至成熟阶段及成熟期的氮、磷、钾积累量显著下降, 主要由于L3显著降低了抽穗后期的根系α-NA氧化量及根干重。尽管在抽穗后随着植株根系逐渐衰老及机能下降, L3对根系、养分吸收及最终产量的影响要小于L2, 但最终产量依然显著低于L1, 达10.91%~18.47%。L2和L3条件下, 随着氮肥水平增加, 植株根系α-NA氧化量及根干重显著增加, 导致拔节至成熟期各阶段的氮、磷、钾积累量显著增加, 最终产量及氮肥利用率显著提高。由此可见氮肥施用能部分弥补弱光逆境对超级粳稻氮、磷、钾吸收及产量的影响。

关键词: 超级粳稻; 遮光; 氮肥; 产量; 养分吸收

Effects of Light, Nitrogen and Their Interaction on Grain Yield and Nitrogen, Phosphorus and Potassium Absorption inJaponica Super Rice

WANG Ya-Jiang, WEI Hai-Yan^*, YAN Xi-Ting, GE Meng-Jie, MENG Tian-Yao, ZHANG Hong-Cheng, DAI Qi-Gen, HUO Zhong-Yang, XU Ke, FEI Xin-Ru

Key Laboratory of Crop Genetics and Physiology of Jiangsu Province / Agricultural College of Yangzhou University / Innovation Center of Rice Cultivation Technology in Yangtze Rive Valley, Ministry of Agriculture, Yangzhou 225009, China

Fund:

Abstract

A field experiment was conducted usingjaponica super rice Nanjing 44 and Ningjing 3 with two nitrogen fertilization levels (N10: 150 kg ha^-1; N20: 300 kg ha^-1) and three shading treatments (L1: no shading; L2: shading 20 days before heading; L3: shading 20 days after heading) to study the effects of light, nitrogen and their interaction on grain yield and NPK absorption. Results showed that yield under the same level of nitrogen fertilizer was L1>L3>L2. Compared with L1, L2 significantly reduced NPK accumulation from jointing to heading and at whole heading stage, mainly due to the significant decrease of the root oxidation ability of α-NA and root dry weight before heading, resulting in decreased root nutrient uptake capacity and the final yield decrease of 30.58%-35.26%. L3 significantly reduced NPK accumulation from heading to maturity and at whole maturity stage, mainly due to the significant decrease of the root oxidation ability of α-NA and root dry weight after heading. Although the root was gradual aging with the function declining after heading, resulting in the reduced effect on root system, nutrient absorption and yield in L3 than in L2, the final yield was still 10.91%-18.47% lower in L3 than in L1. Under L2 and L3 conditions, with increasing nitrogen level, the root oxidation ability of α-NA and root dry weight increased significantly, resulting in that NPK accumulation increased significantly in various stages from jointing to maturity, then the final yield and nitrogen use efficiency improved significantly. This indicated that nitrogen fertilizer could partially offset the impact of low light stress on NPK nutrient absorption and grain yield injaponica super rice.

Keyword: Japonica super rice; Shading; Nitrogen; Yield; Nutrient uptake

Show Figures

在超级稻高产形成过程中, 光照时间及光照强度有重大影响^{[ 1, 2]}, 氮肥用量、运筹方式以及氮肥种类也有重要意义^{[ 3, 4, 5]}。根据本课题组已有的研究表明^{[ 9]}, 超级稻生育过程中, 不同时间段的遮光对水稻LAI、光合势、干物质积累量均会造成一定的影响, 导致最终的群体颖花量、结实率、千粒重及产量有显著的差异, 氮肥施用能部分弥补弱光逆境对超级稻物质生产及产量的影响。而在产量的形成过程中, 氮、磷、钾元素对器官建成^{[ 7]}、产量潜力发挥^{[ 8, 9, 10, 11]}具有重要的作用, 但不同生育阶段遮光及氮肥施用对水稻吸收氮、磷、钾元素有何影响尚不清楚, 因此本文在前期研究的基础上, 继续深入研究以探明不同光、氮处理下氮、磷、钾养分吸收特性及其与产量、根系形态生理的关系, 为超级稻超高产栽培及养分管理提供理论依据。

1 材料与方法

1.1 试验地点及供试材料

扬州大学农学院试验农场的土质为沙壤土, 地力较好、营养平衡。前茬为小麦。土壤含全氮0.13%、碱解氮87.45 mg kg^-1、速效磷32.8 mg kg^-1、速效钾88.3 mg kg^-1。

供试品种为超级粳稻品种宁粳3号(生长前期分蘖强, 长势旺, 后期熟相较好, 抗倒性较强, 落粒性中等, 全生育期158 d左右, 株高98 cm左右)和南粳44 (生长前期分蘖较强, 株型紧凑, 后期群体整齐度好, 熟相好, 穗型大, 结实率高, 全生育期158 d左右, 株高100 cm左右)。

1.2 试验设计

2011—2012年采用二因素随机区组设计, 设置2种氮素水平, 即N10 (150 kg hm^-2)和N20 (300 kg hm^-2); 3种遮光处理, 即L1 (不遮光)、L2 (抽穗前遮光20 d)和L3 (抽穗后遮光20 d), 使用黑色遮阳网遮光, 遮光率达50%。于5月17日播种, 6月14日移栽, 栽插密度为27万穴 hm^-2 (14.4 cm × 26.0 cm)。双本栽插。小区面积15 m², 各处理重复3次。为保证单独排灌, 作埂隔离各小区并用塑料薄膜覆盖埂体。为方便根系取样, 移栽前在各小区打入长轴半径为13.0 cm, 短轴半径为7.2 cm, 深30 cm的椭圆形铁管, 将管内泥土挖出, 装入特制的圆形营养袋, 再将袋紧贴管壁, 然后抽出铁管, 保留带土营养袋。营养袋材料为黑色耐氧化聚乙烯膜, 口径21 cm, 高30 cm, 袋底和侧面设8个口径为0.5 cm的滤水透气孔。每小区以S形填埋12个带土营养袋, 然后整地、覆水, 施用基肥, 在每个营养袋中央按照要求栽插水稻秧苗。基肥∶蘖肥∶穗肥=2.5∶2.5∶5.0, 其中穗肥分别于倒四叶和倒二叶叶龄期等量施入。每公顷分别以过磷酸钙和氯化钾的形式基施P₂O₅150 kg和K₂O 150 kg。其他管理措施按常规栽培要求实施。

1.3 测定项目与方法

1.3.1 植株氮、磷、钾的测定分别于拔节、抽穗和成熟期从每小区取有代表性植株4穴, 105℃杀青, 80℃烘至恒重后测定各器官(茎鞘、叶片和穗)及全株的干物质重。植株经粉碎后, 用半微量凯氏定氮法测定氮素含量, 钒钼黄比色法测定磷素含量, 火焰光度计法测定钾素含量。

氮(磷、钾)素积累量(kg hm^-2) = 该时期地上部干物重×含氮(磷、钾)率

氮(磷、钾)素阶段积累量(kg hm^-2) = 后一时期的氮(磷、钾)素积累量-前一时期的氮(磷、钾)素积累量

氮肥吸收利用率(%) = (施氮区植株吸氮量-氮空白区植株吸氮量)/施氮量

氮(磷、钾)籽粒生产效率(kg kg^-1) = 水稻稻谷产量/水稻吸氮(磷、钾)量

1.3.2 根系相关性状及活性测定于拔节期、抽穗前10 d、抽穗期、抽穗后10 d和成熟期各取埋入土中的4个营养袋, 剥去袋体, 置40目尼龙网袋中用流水冲洗获得完整根系, 其中两穴用于地上部及根干重测定, 将鲜根及地上部105℃杀青30 min, 80℃烘干至恒重, 计算根冠比。用α-NA氧化法测定另外两穴根系对α-NA的氧化量。

1.3.3 数据计算和统计分析两年试验的重复性较好, 处理间各指标值变化趋势一致, 因此, 本文取2012年数据。以Microsoft Excel 2003处理数据, DPS软件进行其他统计分析。

2 结果与分析

2.1 光、氮及其互作对超级粳稻产量的影响

由图1可知, 2个超级粳稻品种年度间产量差异均不大( F= 0.20, F= 3.24)。以2012年为例, 2个超级粳稻品种在2种氮肥水平的产量均呈现L1>L3> L2, 差异显著。其中N10条件下L1比L2高30.58%~35.26%, L1比L3高10.91%~14.43%; N20条件下L1比L2高30.71%~30.75%, L1比L3高15.42%~18.47%。不同光氮处理的产量, 两个品种均呈现N20L1>N20L3>N20L2>N10L1>N10L3>N10L2, 差异显著。

	Figure Option View Download New Window
	图1 光、氮及其互作下超级粳稻的产量Fig. 1 Yield of super rice under effects of light, nitrogen and their interaction

2.2 光、氮及其互作对超级粳稻氮、磷、钾吸收利用的影响

2.2.1 对氮素阶段积累量及比例的影响由表1可知, 较之L1, L2使2个超级粳稻品种在拔节至抽穗阶段及抽穗至成熟阶段的氮素积累量显著下降; L3则使2个超级粳稻品种在抽穗至成熟阶段的氮素积累量显著下降。L2和L3条件下, 随着氮肥水平增加, 各生育阶段的氮素积累量均显著增加。

表1 光、氮及其互作对超级粳稻氮素阶段积累量及比例的影响 Table 1 Effects of light, nitrogen and their interaction on periodical nitrogen accumulation and its ratio in super rice

品种 Cultivar	氮肥水平 Nitrogen level	光处理 Light treatment	拔节前 SO-EG		拔节至抽穗 EG-HD		抽穗至成熟 HD-MA
品种 Cultivar	氮肥水平 Nitrogen level	光处理 Light treatment	N积累量 Nitrogen accumulation (kg hm^-2)	比例 Ratio(%)	N积累量 Nitrogen accumulation (kg hm^-2)	比例 Ratio(%)	N积累量 Nitrogen accumulation (kg hm^-2)	比例 Ratio(%)
南粳44 Nanjing 44	N10	L1	48.73 Bb	34.49 Dd	64.34 Cc	45.53 Cc	28.23 Cc	19.98 Aa
		L2	48.43 Bb	39.55 Aa	50.13 Dd	40.94 Dd	23.89 Dd	19.51 Aa
		L3	48.36 Bb	37.15 Bb	65.15 Cc	50.04 Bb	16.68 Ff	12.81 Cc
	N20	L1	66.32 Aa	32.57 Ee	101.75 Aa	49.97 Bb	35.55 Aa	17.46 Bb
		L2	66.61 Aa	36.84 Bb	82.86 Bb	45.83 Cc	31.35 Bb	17.34 Bb
		L3	66.98 Aa	35.29 Cc	101.51 Aa	53.48 Aa	21.31 Ee	11.23 Dd
	F值 F-value	Nitrogen (N)	503.11^**	440.15^**	6137.75^**	626.29^**	621.69^**	347.96^**
		Light (L)	0.01	687.58^**	603.26^**	808.62^**	854.46^**	1522.38^**
		N × L	0.14	7.05^*	9.79^*	6.35^*	12.59^*	5.99^*
宁粳3号 Ningjing 3	N10	L1	51.41 Bb	33.02 Dd	72.91 Cc	46.82 Cd	31.40 Cc	20.17 Aa
		L2	51.42 Bb	37.76 Aa	57.38 Dd	42.14 Df	27.36 Dd	20.11 Aa
		L3	51.94 Bb	35.64 BCc	73.67 Cc	50.54 Bb	20.15 Ff	13.82 Cc
	N20	L1	71.47 Aa	32.99 Dd	106.62 Aa	49.21 Bc	38.58 Aa	17.80 Bb
		L2	71.41 Aa	36.53 Bb	89.39 Bb	45.73 Ce	34.68 Bb	17.74 Bb
		L3	71.94 Aa	35.13 Cc	107.08 Aa	52.29 Aa	25.77 Ee	12.58 Dd
	F值 F-value	Nitrogen (N)	550.87^**	23.79^**	29263.49^**	179.27^**	974.93^**	181.38^**
		Light (L)	0.16	393.34^**	3317.82^**	504.98^**	1085.91^**	673.69^**
		N × L	0.01	8.31^*	7.41^*	7.85^*	6.43^*	6.44^*

同一品种的相同数据列, 标以不同大小写字母的数值分别在1%和5%水平差异显著。N10: 150 kg hm^-2纯氮; N20: 300 kg hm^-2纯氮; L1: 不遮光; L2: 抽穗前遮光20 d; L3: 抽穗后遮光20 d。^*和^**分别表示在5%和1%水平上差异显著。

Values within the same column for a cultivar followed by different letters are significantly different at 1% (capital) and 5% (lowercase) probability levels, respectively. N10: 150 kg hm^-2 N applied; N20: 300 kg hm^-2 N applied; L1: no shading; L2: shading 20 days before heading; L3: shading 20 days after heading. SO: sowing; EG: elongation; HD: heading; MA: maturing.^* and^** denote significant difference at 5% and 1% probability levels, respectively.

对于氮素阶段积累比例, 较之L1, L2使2个超级粳稻品种在拔节至抽穗阶段显著下降, 而拔节前则显著增加; L3使2个超级粳稻品种在抽穗至成熟阶段显著下降, 而拔节前及拔节至抽穗阶段则显著增加。L2和L3条件下, 随着氮肥水平增加, 拔节至抽穗阶段的氮素积累比例呈增加趋势, 而拔节前及抽穗至成熟阶段呈下降趋势。

2.2.2 对氮素籽粒生产效率及氮肥利用率的影响

图2表明, 2个超级粳稻品种在2种氮肥水平下的氮素籽粒生产效率及氮肥利用率均呈现L1>L3>L2, 差异显著。其中氮素籽粒生产效率方面, N10条件下L1比L2高13.17%~18.27%; N20条件下L1比L2高16.10%~17.93%。2个品种的氮素籽粒生产效率均呈现N10L1>N20L1>N10L3>N20L3>N10L2>N20L2,差异均显著。氮肥吸收利用率方面, N10条件下L1比L2高39.41%~41.96%; N20条件下L1比L2高20.01%~21.49%。2个品种的氮肥吸收利用率均呈现N10L1>N20L1>N20L3>N10L3>N20L2>N10L2, 差异均显著。

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	图2 光、氮及其互作下超级粳稻的氮素籽粒生产效率及氮肥利用率Fig. 2 Nitrogen grain production efficiency and nitrogen use efficiency of super rice under effects of light, nitrogen and their interaction

2.2.3 对磷素阶段积累量及比例的影响由表2可知, 较之L1, L2使2个超级粳稻品种在拔节至抽穗阶段及抽穗至成熟阶段的磷素积累量显著下降; L3则得2个超级粳稻品种在抽穗至成熟阶段的磷素积累量显著下降。L2和L3条件下, 随着氮肥水平增加, 各生育阶段的磷素积累量均显著增加。

表2 光、氮及其互作对超级粳稻磷素阶段积累量及比例的影响 Table 2 Effects of light, nitrogen and their interaction on periodical phosphorus accumulation and its ratio in super rice

品种 Cultivar	氮肥水平 Nitrogen level	光处理 Light treatment	拔节前 SO-EG		拔节至抽穗 EG-HD		抽穗至成熟 HD-MA
品种 Cultivar	氮肥水平 Nitrogen level	光处理 Light treatment	P积累量 Phosphorus accumulation (kg hm^-2)	比例 Ratio(%)	P积累量 Phosphorusaccumulation (kg hm^-2)	比例 Ratio(%)	P积累量 Phosphorus accumulation (kg hm^-2)	比例 Ratio(%)
南粳44 Nanjing 44	N10	L1	12.75 Bb	23.93 CDcd	25.18 Cc	47.27 Dd	15.34 Cc	28.80 Aa
		L2	12.75 Bb	27.07 Aa	20.93 Dd	44.44 Ee	13.42 De	28.49 Aa
		L3	13.05 Bb	25.89 Bb	25.97 Cc	51.51 Bb	11.40 Ef	22.60 Cc
	N20	L1	18.24 Aa	23.40 Dd	40.09 Aa	51.43 Bb	19.62 Aa	25.17 Bb
		L2	18.13 Aa	26.35 ABb	33.51 Bb	48.68 Cc	17.19 Bb	24.97 Bb
		L3	17.89 Aa	24.35 Cc	40.62 Aa	55.29 Aa	14.96 Cd	20.37 Dd
	F值 F-value	Nitrogen (N)	133.55^**	60.21^**	3507.50^**	5619.88^**	2292.27^**	356.35^**
		Light (L)	0.01	215.30^**	263.93^**	5377.13^**	943.36^**	468.41^**
		N × L	0.20	6.69^*	9.75^*	6.86^*	6.99^*	7.36^*
宁粳3号 Ningjing 3	N10	L1	13.03 Bb	22.28 De	28.61 Cc	48.91 Cd	16.85 Cc	28.81 Aa
		L2	13.20 Bb	26.01 Ab	22.97 Dd	45.29 De	14.56 Ee	28.70 Aa
		L3	12.83 Bb	23.33 Cd	29.63 Cc	53.85 Bb	12.55 Ff	22.82 Cc
	N20	L1	19.14 Aa	23.09 Cd	43.42 Aa	52.39 Bc	20.32 Aa	24.52 Bb
		L2	19.47 Aa	26.44 Aa	36.39 Bb	49.40 Cd	17.79 Bb	24.16 Bb
		L3	19.42 Aa	24.50 Bc	44.07 Aa	55.60 Aa	15.78 Dd	19.90 Dd
	F值 F-value	Nitrogen (N)	194.73^**	88.73^**	6995.17^**	179.05^**	10611.45^**	440.67^**
		Light (L)	0.12	592.17^**	708.01^**	336.58^**	6324.49^**	344.10^**
		N × L	0.10	6.28^*	6.04^*	9.19^*	6.20^*	7.28^*

对于磷素阶段积累比例, 较之L1, L2使2个超级粳稻品种在拔节至抽穗阶段显著下降, 而拔节前显著增加; L3使2个超级粳稻品种在抽穗至成熟阶段显著下降, 而拔节前及拔节至抽穗阶段显著增加。L2和L3条件下, 随着氮肥水平增加, 拔节至成熟阶段的磷素积累比例呈增加趋势, 而抽穗至成熟阶段呈下降趋势。

2.2.4 对磷素籽粒生产效率的影响图3表明, 2个超级粳稻品种在2种氮肥水平下的磷素籽粒生产效率均呈现L1>L3>L2, 差异显著。其中N10条件下L1比L2高15.46%~17.32%; N20条件下L1比L2高15.45%~16.15%。对于磷素籽粒生产效率, 南粳44呈现N10L1 > N20L1 > N10L3 > N10L2 > N20L3 > N20L2; 宁粳3号呈现N10L1 > N10L3 > N20L1 > N20L3 > N10L2 > N20L2, 差异均显著。

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	图3 光、氮及其互作下超级粳稻的磷素籽粒生产效率Fig. 3 Phosphorus grain production efficiency of super rice under effects of light, nitrogen and their interaction

2.2.5 对钾素阶段积累量及比例的影响由表3可知, 较之L1, L2使2个超级粳稻品种在拔节至抽穗阶段及抽穗至成熟阶段的钾素积累量显著下降; L3则使2个超级粳稻品种在抽穗至成熟阶段的钾素积累量显著下降。L2和L3条件下, 随着氮肥水平增加, 各生育阶段的钾素积累量均显著增加。

表3 光、氮及其互作对超级粳稻钾素阶段积累量及比例的影响 Table 3 Effects of light, nitrogen and their interaction on periodical potassium accumulation and its ratio in super rice

品种 Cultivar	氮肥水平 Nitrogen level	光处理 Light treatment	拔节前 SO-EG		拔节至抽穗 EG-HD		抽穗至成熟 HD-MA
品种 Cultivar	氮肥水平 Nitrogen level	光处理 Light treatment	K积累量 Potassium accumulation (kg hm^-2)	比例 Ratio(%)	K积累量 Potassiumaccumulation (kg hm^-2)	比例 Ratio(%)	K积累量 Potassium accumulation (kg hm^-2)	比例 Ratio(%)
南粳44 Nanjing 44	N10	L1	100.13 Bb	47.05 Cc	103.32 Cc	48.55 De	9.35 Cc	4.40 Aa
		L2	100.82 Bb	51.07 Aa	88.29 Dd	44.73 Ef	8.30 Dd	4.20 Aa
		L3	101.01 Bb	48.53 Bb	104.22 Cc	50.08 Cc	2.88 Ff	1.39 Cc
	N20	L1	145.50 Aa	43.82 Ee	174.77 Aa	52.63 Bb	11.79 Aa	3.55 Bb
		L2	145.48 Aa	47.17 Cc	152.74 Bb	49.52 Cd	10.23 Bb	3.32 Bb
		L3	144.40 Aa	44.84 Dd	174.33 Aa	54.02 Aa	4.01 Ee	1.14 Cc
	F值 F-value	Nitrogen (N)	2472.61^**	2514.77^**	7696.90^**	1873.39^**	415.84^**	66.67^**
		Light (L)	0.09	905.23^**	252.22^**	876.63^**	2386.05^**	463.10^**
		N × L	0.41	7.57^*	7.51^*	7.11^*	18.34^**	6.44^*
宁粳3号 Ningjing 3	N10	L1	112.71 Bb	45.28 BCc	126.05 Cc	50.64 Cc	10.18 Cc	4.09 Aa
		L2	112.83 Bb	50.63 Aa	101.30 Dd	45.45 Dd	8.74 Dd	3.92 Aa
		L3	112.83 Bb	46.43 Bb	127.06 Cc	52.28 Bb	3.14 Ff	1.29 Cc
	N20	L1	158.68 Aa	43.51 Dd	192.85 Aa	52.88 ABb	13.14 Aa	3.60 Bb
		L2	158.85 Aa	46.83 Bb	168.78 Bb	49.76 Cc	11.54 Bb	3.40 Bb
		L3	158.11 Aa	44.58 CDc	192.80 Aa	54.31 Aa	4.12 Ee	1.11 Cc
	F值 F-value	Nitrogen (N)	2899.39^**	132.60^**	100580.90^**	151.10^**	344.89^**	80.36^**
		Light (L)	0.06	146.56^**	6112.04^**	213.45^**	1657.67^**	1484.78^**
		N × L	0.08	9.55^*	5.80^*	9.78^*	27.55^**	6.03^*

对于钾素阶段积累比例, 较之L1, L2使2个超级粳稻品种在拔节至抽穗阶段显著下降, 而拔节前显著增加; L3使2个超级粳稻品种在抽穗至成熟阶段显著下降, 而拔节前及拔节至抽穗阶段显著增加。L2和L3条件下, 随着氮肥水平增加, 拔节至成熟阶段的钾素积累比例呈增加趋势, 而拔节前及抽穗至成熟阶段呈下降趋势。

2.2.6 对钾素籽粒生产效率的影响图4表明, 2个超级粳稻品种在2种氮肥水平下的钾素籽粒生产效率均呈现L1>L3>L2, 差异显著。其中N10条件下L1比L2高21.09%~21.14%; N20条件下L1比L2高21.46%~ 21.57%。2个品种的钾素籽粒生产效率均呈现N10L1> N20L1>N10L3>N10L2>N20L3>N20L2, 差异均显著。

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	图4 光、氮及其互作下超级粳稻的钾素籽粒生产效率Fig. 4 Potassium grain production efficiency of super rice under effects of light, nitrogen and their interaction

2.3 光、氮及其互作对超级粳稻根系形态生理的影响

2.3.1 对根系α-NA氧化量的影响由表4可知,

较之L1, L2使2个超级粳稻品种在抽穗前期的根系α-NA氧化量显著下降, 但遮光解除后水稻根系α-NA氧化量则显著增加; L3使2个超级粳稻品种在抽穗后期根系α-NA氧化量显著下降, 但遮光解除后水稻根系α-NA氧化量也显著增加。L2和L3条件下, 随着氮肥水平增加, 各生育期的根系α-NA氧化量均显著增加。

表4 光、氮及其互作对超级粳稻根系α-NA氧化量的影响 Table 4 Effects of light, nitrogen and their interaction on root oxidation ability of α-NA of super rice (µg h^-1 g^-1)

2.3.2 对根干重的影响由表5可知, 较之L1, L2使2个超级粳稻品种在抽穗期及成熟期的根干重显著下降; L3使2个超级粳稻品种在成熟期的根干重显著下降。L2和L3条件下, 随着氮肥水平增加, 各生育期的根干重均显著增加。

表5 光、氮及其互作对超级粳稻根干重的影响 Table 5 Effects of light, nitrogen and their interaction on root dry weight of super rice (t hm^-2)

2.3.3 对根冠比的影响由表6可知, 较之L1, L2使2个超级粳稻品种在抽穗期及成熟期的根冠比显著增加; L3使2个超级粳稻品种在成熟期的根冠比显著增加, 而根冠比显著增加的主要原因是遮光

条件下植株地上部干物重降低的幅度远远大于根部干物重降低的幅度。L2和L3条件下, 随着氮肥水平增加, 各生育期的根冠比均显著下降。

表6 光、氮及其互作对超级粳稻根冠比的影响 Table 6 Effects of light, nitrogen and their interaction on ratio of root to shoot of super rice

2.4 超级粳稻氮、磷、钾积累量、根系形态生理指标与产量的相关

相关分析表明(表7), 不同光、氮条件下, 产量与主要生育期的氮、磷、钾元素积累量、根系干重和根系α-NA氧化量均呈正相关, 除与抽穗后10 d及成熟期的根系α-NA氧化量相关性不显著外, 其余都达到显著或极显著水平。而与根冠比呈极显著负相关。

表7 超级粳稻氮、磷、钾积累量、根系形态生理指标与产量的相关系数 Table 7 Correlation coefficients of nitrogen, phosphorus and potassium accumulation, root characteristics with yield of super rice

3 讨论

3.1 光、氮及其互作对超级粳稻产量及氮、磷、钾养分吸收特性的影响

前人研究表明, 不同光照条件对水稻产量有显

著影响, 如各生育阶段遮光均导致水稻产量显著下降^{[ 9, 15]}。氮、磷、钾是水稻正常生长发育必不可少的三大营养元素, 对水稻产量潜力的发挥起着重要作用, 然而不同时期遮光对超级粳稻氮、磷、钾养分吸收又有怎样的影响呢？本研究表明, L2使植株拔节至抽穗阶段及抽穗期的氮、磷、钾元素积累量显著下降, 最终产量显著下降, 同时氮肥利用率及氮、磷、钾元素籽粒生产效率也显著下降; L3使植株抽穗至成熟阶段及成熟期的氮、磷、钾元素积累量显著下降, 最终产量显著下降, 同时氮肥利用率及氮、磷、钾元素籽粒生产效率也显著下降, 但下降幅度均低于L2, 主要因为拔节至抽穗阶段是水稻一生中氮、磷、钾元素积累的高峰期, 该阶段的养分积累与群体叶面积发展、颖花分化及最终产量密切相关^{[ 13]}。L2和L3条件下, 随着氮肥水平增加, 拔节至抽穗阶段、抽穗期及抽穗至成熟阶段、成熟期的氮、磷、钾元素积累量均显著增加, 最终产量及氮肥利用率显著提高, 这说明氮肥施用能部分弥补因弱光逆境对超级粳稻氮、磷、钾元素吸收特性及产量的影响。同时, 不同光、氮条件下的氮、磷、钾养分吸收还具有协同性。

3.2 光、氮及其互作对超级粳稻根系形态生理及氮、磷、钾养分吸收特性的影响

根系不仅是水稻的支持器官, 水分、养分的吸收器官, 同时也是植株地下部与地上部物质交换和代谢的重要器官。水稻根系与植株氮^{[ 14, 15, 16]}、磷^{[ 17, 18]}、钾^{[ 19, 20]}元素的吸收效率有密切的关系。本研究表明, L2显著降低了植株抽穗前期的根系活力及根干重, 导致抽穗前期的氮、磷、钾元素积累量显著下降, 最终氮肥利用率及氮、磷、钾元素籽粒生产效率显著下降。L3显著降低了植株抽穗后期的根系活力及根干重, 导致抽穗后期的氮、磷、钾元素积累量显著下降, 最终氮肥利用率及氮、磷、钾元素籽粒生产效率显著下降, 但下降幅度均低于L2, 主要由于抽穗后植株根系逐渐衰老及机能下降^{[ 21]}。此外, 弱光逆境解除后, 水稻根系活力得以适当恢复, 表现为抽穗期及成熟期根系α-NA氧化量高于不遮光处理。L2和L3条件下, 通过增施氮肥可促进根系生长, 提高根系活力, 延缓后期根系衰老, 进而促进生育中后期水稻根系对氮、磷、钾养分的吸收以提高最终产量。这说明氮肥施用能部分弥补因弱光逆境对超级粳稻根系形态生理及氮、磷、钾养分吸收特性的影响。

4 结论

光、氮及其互作对超级粳稻产量及氮、磷、钾养分吸收特性有显著的影响。L2显著降低了植株抽穗前期的根干重及根系活力, 导致该阶段的氮、磷、钾元素积累量显著下降, 最终产量显著低于L1。L3显著降低了植株抽穗后期的根干重及根系活力, 导致该阶段的氮、磷、钾元素积累量显著下降, 尽管抽穗后植株根系逐渐衰老及机能下降, L3对根系、养分吸收及最终产量的影响小于L2, 但最终产量依然显著低于L1。L2和L3条件下, 随着氮肥水平增加, 植株根干重及根系活力显著增加, 导致植株生育中后期的养分吸收能力提高, 最终产量及氮肥利用率显著提高。由此可见氮肥施用能部分弥补弱光逆境下超级粳稻对氮、磷、钾养分吸收能力, 使产量有所提高。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。

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[15]	魏海燕, 张洪程, 张胜飞, 杭杰, 戴其根, 霍中洋, 许轲, 马群, 张庆, 刘艳阳. 不同氮利用效率水稻基因型的根系形态与生理指标的研究. 作物学报, 2008, 34: 429-436 Wei H Y, Zhang H C, Zhang S F, Hang J, Dai Q G, Huo Z Y, Xu K, Ma Q, Zhang Q, Liu Y Y. Root morphological and physiological characteristics in rice genotypes with different N use efficiencies. Acta Agron Sin, 2008, 34: 429-436 (in Chinese with English abstract) [本文引用:2] [CJCR: 1.667]
[16]	戢林, 李廷轩, 张锡洲, 余海英. 氮高效利用基因型水稻根系形态和活力特征. 中国农业科学, 2012, 45: 4770-4781 Ji L, Li T X, Zhang X Z, Yu H Y. Root morphological and activity characteristics of rice genotype with high nitrogen utilization efficiency. Sci Agric Sin, 2012, 45: 4770-4781 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.889]
[17]	李锋, 曲雪艳, 潘晓华, 姚锋先, 杨福孙. 不同水稻品种对难溶性磷利用能力的初步研究. 植物营养与肥料学报, 2003, 9: 420-424 Li F, Qu X Y, Pan X H, Yao F X, Yang F S. Preliminary study on the utilization ability of insoluble phosphate in various rice cultivars. Plant Nutr Fert Sci, 2003, 9: 420-424 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.883]
[18]	郭玉春, 林文雄, 石秋梅, 梁义元, 何华勤, 陈芳育. 低磷胁迫下不同磷效率水稻苗期根系的生理适应性研究. 应用生态学报, 2003, 14: 61-65 Guo Y C, Lin W X, Shi Q M, Liang Y Y, He H Q, Chen F Y. Physiological adaptability of seeding rice genotypes with different P uptake effciency under low P-deficient stress. Chin J Appl Ecol, 2003, 14: 61-65 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.742]
[19]	刘建祥, 杨肖娥, 杨玉爱, 吴良欢. 低钾胁迫下水稻钾高效基因型若干生长特性和营养特性的研究. 植物营养与肥料学报, 2003, 9: 190-195 Liu J X, Yang X E, Yang Y A, Wu L H. Some agronomic and nutritional characteristics for potassium efficient rice genotypes under low potassium stress. Plant Nutr Fert Sci, 2003, 9: 190-195 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.883]
[20]	台德卫, 张效忠, 苏泽胜, 王元垒, 罗彦长, 夏家发. 全球水稻分子育种核心种质资源耐低钾品种的苗期筛选. 植物遗传资源学报, 2004, 5: 356-359 Tai D W, Zhang X Z, Su Z S, Wang Y L, Luo Y C, Xia J F. Screening for low-kalium tolerance varieties at seedling stage from the core germplasm of integrated international rice. J Plant Genet Resour, 2004, 5: 356-359 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.1628]
[21]	褚光, 杨凯鹏, 王静超, 张耗. 水稻根系形态与生理研究进展. 安徽农业科学, 2012, 40: 5097-5101 Zhu G, Yang K P, Wang J C, Zhang H. Research progress on root morphology and physiology of rice. J Anhui Agric Sci, 2012, 40: 5097-5101 (in Chinese with English abstract) [本文引用:1] [CJCR: 0.687]

2009

0.0

1.474

. 2009, 30:219-222

Effects of ecological and climatic conditions on super rice yields of Zhunliangyou 527 in Southeast Guizhou

以贵州省黔东南州16个气象观测站2005-2007年的气温、降水量、日照时数以及海拔高度、播种期作为主要生态气候因子,分析其对准两优527超级交杂水稻产量的影响.结果表明:准两优527最适宜在海拔480-750m的地区栽培,尤其是海拔620m上下地带容易获得高产;最适宜播种期为3月28日-4月7日,播种期每推迟10d,产量减少48.2～376.3kg/667m2;最适宜积温条件为3200～3800℃·d,积温每增加100℃·d,产量减少8.32～140.43kg/667m2;最适宜降水量条件为460～800mm,降水量每增加或减少100mm,产量减少7.41～338.36kg/667m2;最适宜光照条件为620～730h,光照少于500h或多于900h,产量降低明显.

... 在超级稻高产形成过程中, 光照时间及光照强度有重大影响^[1,2], 氮肥用量、运筹方式以及氮肥种类也有重要意义^[3,4,5] ...

2008

0.0

1.667

. 2008, 34:2003-2009 DOI:10.3724/SP.J.1006.2008.02003

Effect of shading on the photosynthetic characteristics and yield at later growth stage of hybrid rice combination

1 Rice Research Institute of Sichuan Agricultural University, Wenjiang 611130, Sichuan; 2 Agricultural College of Sichuan Agricultural University, Ya’an 625014, Sichuan, China

Photosynthetic characteristics in shading treatment and adaptation mechanism under adverse conditions were studied using in six hybrid rice combinations with different photosensitivity. The results showed that the grain yield was significantly reduced under shading treatment (80% shading rate) at heading stage, then recovered to natural illumination; the chlorophyll (a+b) and chlorophyll b contents were increased, but failed to reach the normal level after the recovery period; low light reduced the photosynthetic rate of the functional leaves and the quantity of photosynthetic products transported to grain, meanwhile, the content of malondialdehyde (MDA) and leaf area index were increased, the activity of SOD varied differently. There was a great difference in adaptability to low light environment among combinations. It is the physiological basis of rice tolerance to shading that the flag leaf under low light stress can maintain a higher net photosynthetic rate (P_n), quantum efficiency, chlorophyll content, and lower content of malondialdehyde (MDA) and lower SOD activity.

对6个光敏感性不同的杂交稻组合在遮光处理条件下的光合特性及其对逆境的适应机理进行了研究。结果表明，在田间通过抽穗期遮光处理(80%遮光率) 15 d，及恢复自然光照15 d后，水稻的产量显著降低；水稻剑叶叶绿素含量增加，恢复生长15 d后，并未恢复到正常水平；弱光降低功能叶片的净光合速率，减少光合产物向籽粒的运输，增加MDA含量和叶面积指数，而SOD活性变化不一。杂交水稻适应弱光环境的能力是由自身因素决定的，组合之间存在很大差异。弱光胁迫下剑叶能保持较高的净光合速率、量子效率、叶绿素含量以及较低的MDA含量、SOD活性，是耐弱光水稻品种的生理基础。

... 在超级稻高产形成过程中, 光照时间及光照强度有重大影响^[1,2], 氮肥用量、运筹方式以及氮肥种类也有重要意义^[3,4,5] ...

2003

0.0

1.889

. 2003, 36:800-806

Effects of N-application rate on yield, quality and characters of nitrogen uptake of hybrid rice variety Liangyoupeijiu

扬州大学农学院

在大田条件下 ,以汕优 6 3为对照 ,研究不同施氮量对杂交稻两优培九的影响。结果表明 ,随施氮量的增加 ,两优培九的吸氮量、干物质累积量、百公斤籽粒需氮量均增加 ;产量以及氮肥利用率均随施氮量的增加而增加 ,至中肥处理达最大值 ,而高肥处理则显著降低 ;相同施氮水平 ,两优培九的产量、吸氮量、干物质累积量以及肥料利用率均高于汕优 6 3;在 90 0 0～ 10 5 0 0kg·ha-1的产量水平下 ,百公斤籽粒需氮量为 2 .0～ 2 .2kg ,氮肥利用率在 40 %左右 ;适当增施氮肥有利于营养品质、加工品质、蒸煮食味品质的提高 ,而高肥不利于外观品质的提高。在品种间产量和品质均存在显著差异 ,4个氮肥处理水平下 ,两优培九的产量均高于汕优 6 3、品质优于汕优 6 3。

... 在超级稻高产形成过程中, 光照时间及光照强度有重大影响^[1,2], 氮肥用量、运筹方式以及氮肥种类也有重要意义^[3,4,5] ...

2009

0.0

0.687

. 2009, 37(2):705-707

Effects of nitrogen application on growth and yield of super rice

[目的]分析在同等施氮条件下前后期施氮比例对超级稻产量、叶面积指数及干物质积累量的影响.[方法]在大田栽培条件下,以超级稻品种宁粳3号为材料,设置氮肥运筹试验.[结果]按基蘖肥:穗肥为6∶ 4、5∶ 5施用,可提高单位面积穗数、颖花量、粒叶比,优化株型和群体,增加拔节-抽穗、抽穗-成熟的物质积累.[结论]按基蘖肥:穗肥为6∶ 4、5∶ 5施用可提高产量.

... 在超级稻高产形成过程中, 光照时间及光照强度有重大影响^[1,2], 氮肥用量、运筹方式以及氮肥种类也有重要意义^[3,4,5] ...

2011

0.0

0.605

... 在超级稻高产形成过程中, 光照时间及光照强度有重大影响^[1,2], 氮肥用量、运筹方式以及氮肥种类也有重要意义^[3,4,5] ...

2014

0.0

1.667

. 2014, 40:154-165 DOI:10.3724/SP.J.1006.2014.00154

Effects of light, nitrogen and their interaction on grain yield and matter production characteristics of japonica super rice

Key Laboratory of Crop Genetics and Physiology of Jiangsu Province, Agricultural College of Yangzhou University / Innovation Center of Rice Cultivation Technology in Yangtze Rive Valley, Ministry of Agriculture, Yangzhou 225009, China

Field experiment was conducted usingjaponica super rice Nanjing 44 and Ningjing 3 with two nitrogen fertilization levels (N10: 150 kg ha^-1, N20: 300 kg ha^-1) and three shading treatments (L1: no shading, L2: shading 20 days before heading, L3: shading 20 days after heading) to study theeffects of light, nitrogen and their interaction on grain yield and matter production. Results showed that, compared with L1 treatment, L2 not only reduced the effective panicles and grains per panicle, resulting in the decline of total spikelets by 24.81% to 35.63%, but also significantly reduced the tiller number and leaf area index at heading by 2.90%–6.44% and 19.02%–27.17% respectively, causing photosynthetic potential, dry matter accumulation decreased significantly from heading to maturity, and at last the final yield decreased by 27.23%–35.26%. Compared with L1 treatment, L3 mainly affected photosynthate accumulation from heading to maturity, leading to seed setting rate and grain weight significantly decreased by 1.49%–4.48% and 5.54%–9.17% respectively, and the final yield decreased by 10.91%–18.47%. UnderL2 condition, with increasing nitrogen level, tiller number and leaf area index at heading increased significantly, resulting in photosynthetic potential, dry matter accumulation increased significantly from heading to maturity, and the ultimately effective panicles, number of grains per panicle, total number of spikelets and the yield improved significantly. Under L3 condition, with increasing nitrogen level, photosynthate accumulation from heading to maturity increased significantly, dry matter translocation from stems and leavesto the panicle increased significantly, meanwhile, translocation rate and contribution rate were further improved, and the ultimate yield improved significantly. This indicated that nitrogen fertilizer could partially offset the impact of low light stress on the matter production and yield of japonica super rice.

大田条件下以超级粳稻南粳44和宁粳3号为材料，设置2种氮肥水平(N10：150 kg hm^-2，N20：300 kg hm^-2)和3种遮光处理(L1：不遮光，L2：抽穗前遮光20 d，L3：抽穗后遮光20 d)，研究光、氮及其互作对超级粳稻产量和物质生产特征的影响。结果表明，较之L1，L2不仅减少了有效穗数和每穗粒数，导致总颖花量下降，降幅达24.81%~35.63%，而且还显著降低了抽穗期茎蘖数和叶面积指数，降幅达2.90%~6.44%和19.02%~27.17%，导致抽穗至成熟阶段的光合势、干物质积累量显著下降，最终产量显著下降，降幅达27.23%~35.26%。较之L1，L3主要影响了抽穗至成熟阶段的光合物质积累，导致结实率和千粒重显著下降，降幅达1.49%~4.48%和5.54%~9.17%，最终产量显著下降，降幅达10.91%~18.47%。L2条件下，随着氮肥水平增加，抽穗期茎蘖数与叶面积指数均显著增加，导致抽穗至成熟阶段光合势、干物质积累显著增加，最终有效穗数、每穗粒数、总颖花量以及产量显著提高。L3条件下，随着氮肥水平增加，抽穗至成熟阶段的光合物质积累显著提高，其中茎叶干物质向穗部转运量显著增加，转运率和贡献率也进一步提高，最终产量显著提高。由此可见氮肥施用能部分弥补因弱光逆境对超级粳稻物质生产及其产量的影响。

2000

0.0

0.849

... 而在产量的形成过程中, 氮、磷、钾元素对器官建成^[7]、产量潜力发挥^[8,9,10,11]具有重要的作用, 但不同生育阶段遮光及氮肥施用对水稻吸收氮、磷、钾元素有何影响尚不清楚, 因此本文在前期研究的基础上, 继续深入研究以探明不同光、氮处理下氮、磷、钾养分吸收特性及其与产量、根系形态生理的关系, 为超级稻超高产栽培及养分管理提供理论依据 ...

2008

0.0

1.889

. 2008, 41:3123-3132

Characteristics of nutrient uptake and utilization of super hybrid rice under different fertilizer application rates

<FONT face=Verdana>湖南农业大学农学院</FONT><BR>

【目的】探索超级杂交稻对氮、磷、钾养分的吸收利用规律。【方法】于2004年和2005年，选用超级杂交稻品种准两优527和两优293为供试材料，在湖南省5个不同水稻种植生态区进行田间小区试验，研究在农民实际平均施肥量及分别减少和增加25%施肥量条件下，超级杂交稻抽穗期和成熟期植株体内的氮、磷、钾养分含量和吸收积累规律。【结果】无论在抽穗期，还是成熟期，不同施肥水平条件下水稻植株体内氮、磷、钾养分的含量均无显著差异，其在不同地点间的变化幅度低于相应的水稻产量和养分吸收量；养分吸收量差异主要由单位面积干物质生产量不同所引起。在不同施肥水平下，随着产量升高，氮、磷、钾收获指数呈上升趋势，但生产单位稻谷所需养分量呈下降趋势。在产量最高的桂东点，其植株体内氮、磷、钾养分含量和积累量均处于中等水平。【结论】采用多次施肥，不同施肥水平（135～225 kg N•ha-1、29.7～49.5kg P•ha-1、112.1～186.8 kg K•ha-1）对超级杂交稻株体内的氮、磷、钾养分吸收积累影响不明显；随着产量的提高，超级杂交稻对氮、磷、钾养分的吸收利用率也可提高，从而实现高产与养分高效利用的协调统一。

2008

0.0

0.951

. 2008, 23:267-273

Effects of different nutrient levels in different fertility soils on super rice yields and nutrient use efficiency

在不同土壤养分条件下,对氮、磷、钾三大营养元素设置不同水平施用量,研究地力条件和养分水平对超级稻产量和养分利用率的影响.结果表明:随着施氮量的增加,产量呈现先升后降的趋势,平均以施180 kg/hm2产量最高.土壤氮素背景值与空白区产量呈正相关关系,R=0.893 0,也就是说土壤氮含量高,不施氮肥水稻也能高产.氮素利用率与施氮水平呈极显著的负相关关系,与土壤背景氮呈单峰曲线关系.氮素收获指数随施氮水平的提高而下降,土壤氮含量对其并无影响.在土壤严重缺磷和钾素情况下,施磷钾有显著的增产效果,产量和施肥量有正相关关系,R分别为0.984 1和0.940 8,随着土壤磷素和钾素量的上升,施肥增产效果逐渐下降,在土壤含磷＞10 mg/kg状态下甚至出现施磷减产的现象.磷钾肥农学利用率和表观利用率与其施用量呈极显著负相关,磷肥R分别为-0.961 1和-0.904 9,钾肥R分别为-0.953 4和-0.947 2,生理利用率受土壤背景值影响,与施用量的关系表观较复杂.磷肥农学利用率、表观利用率均与土壤背景磷含量呈单峰曲线关系,以土壤磷含量在少量缺乏的情况下,磷肥利用率最高.钾肥农学利用率与土壤含钾量的关系和磷肥相似,但其表观利用率却与土壤含钾量呈负相关关系.磷钾肥的收获指数与氮相似,随施肥水平的提高而下降,土壤磷钾含量对其并无影响.以上表明,超级稻生产在施肥措施中要考虑当地的土壤条件,尽量减少施用量,提高其利用效率,达到增产增效的目的.

... 根据本课题组已有的研究表明^[9], 超级稻生育过程中, 不同时间段的遮光对水稻LAI、光合势、干物质积累量均会造成一定的影响, 导致最终的群体颖花量、结实率、千粒重及产量有显著的差异, 氮肥施用能部分弥补弱光逆境对超级稻物质生产及产量的影响 ...

... 著影响, 如各生育阶段遮光均导致水稻产量显著下降^[9,15] ...

2002

0.0

0.916

. 2002, 25(4):6-10

Nitrogen, phosphorous and potassium uptake characteristics of rice and its relationship with grain yield

在田间条件下,对包括两系杂交稻在内的8个水稻品种的氮磷钾吸收特性及其与产量的关系进行了分析研究.结果表明:早稻仅在分蘖-拔节期出现一个吸氮高峰,约占总吸氮量的60%;晚稻在分蘖期和孕穗-抽穗期出现两个吸氮高峰,分别占总吸氮量的40%和24%左右.早稻吸收磷钾的高峰期可从分蘖期一直持续到抽穗期以后;而晚稻吸收磷钾的高峰期仅在分蘖期,其吸收量占全生育期吸收量的55%以上.在稻谷产量、稻草产量和干物质总积累量方面,两系杂交稻比其它品种具有明显的优势,分别高出21%,46%和32%;在这些方面晚稻比早稻具有更明显的优势,晚稻的磷钾养分(尤其是磷素)利用效率比早稻高,晚稻吸收较少的养分而生产较多的稻谷.相关分析表明,水稻各生育阶段的吸氮量与稻谷产量均呈显著正相关;而中后期的磷钾吸收多为奢侈吸收,它与稻谷产量无相关性.

2011

0.0

1.667

. 2011, 37:882-894 DOI:10.3724/SP.J.1006.2011.00882

Characteristics of nutrient uptake and utilization of mid-season hybrid rice under different nitrogen application rates in different locations of Southwest China

Rice and Sorghum Institute, Sichuan Academy of Agricultural Sciences, Luzhou 646000, China

The characteristics of nitrogen (N), phosphorus (P) and potassium (K) uptake and their utilization of mid-season hybrid rice were studied. Field experiment were conducted with mid-season rice combinations II-you 7 and Yuxiangyou 203 grown under the condition of different ecological sites, chemical qualities of soil and the levels of N application in 2009. By using variance, correlation, regression and partial correlation analysis, the results indicated that significant or highly significant differences of grain yield, dry matter production, uptake amount and harvest indexes of N, P, K and N, P, K requirements of above ground plants for producing of 1 000 kg grains (RAGPPG) were found among seven locations. There were significant or highly significant effects of fertilizer application treatments on grain yield, dry matter production, uptake amount, harvest index and N among RAGPPG, but no effect on P and K among RAGPPG. Highly significantly positive correlations were observed among harvest indexes of N, P, K and among RAGPPG, but not between harvest indexes of N, P, K; RAGPPG and grain yield. Multiple stepwise regression analysis showed that harvest indexes of N, P, K and RAGPPG were significantly influenced by different ecological sites, chemical qualities of soil and the levels of N application, and the determination coefficients ranged from 0.5972–0.8404 and 0.7637–0.8804。The results provide a scientific basis for the determination of the efficient amount of N, P, and K applied for full scale rice cultivation.

以杂交中稻II优7号和渝香优203为材料，在西南稻区4省(市)的7个生态点采用相同的试验方案，研究了地理位置、土化特性、施氮量对植株氮、磷、钾积累和分配的影响。结果表明，不同试验地点间稻谷产量、干物质产量、氮磷钾的吸收量、收获指数和每生产1 000 kg稻谷的氮、磷、钾需要量(RAGPPG)差异显著或极显著。施肥处理对稻谷产量、干物质产量、氮的吸收量、收获指数和RAGPPG中的氮有显著或极显著影响，对RAGPPG中的磷、钾影响不显著。氮、磷、钾收获指数间和RAGPPG间均呈极显著正相关，RAGPPG和收获指数均与稻谷产量水平没有相关性。经逐步回归分析，RAGPPG和氮、磷、钾收获指数均分别与试验点所处地理位置、施肥水平及土化特性呈极显著线性关系，决定系数分别为0.5972~0.8404和0.7637~0.8804。可作为制定各地水稻高产高效相应的氮、磷、钾施肥量的科学依据。

1999

0.0

1.742

. 1999, 10:193-196

Effect of shading on growth, development and yield formation of rice

Institute of Tropical and Subtropical Ecology, South China Agricultural University, Guangzhou 510642

Field experiments under 45% shading at different stages(early, middle 1, middle 2 and later stages) of rice growth show that regardless of different stages, shading decreased the accumulation rate of dry matter and the uptake of nutrients, but increased the nutrient content in rice plant, and affected its yield in varying degrees. Shading at early growth stage mainly decreased tiller number, effective panicle number and leaf area index, and reduced yield by 11.56%, but increased plant height. At middle 1 stage, shading showed no significant influence on rice growth, and the yield was reduced by only 5.46%. At middle 2 stage, shading mainly decreased grains per panicle and 1000 grain weight, and the yield was reduced markedly by 30.80%. At later stage, shading decreased the percentage of filled grains and 1000 grain weight, and the yield was the lowest, reduced by 55.40%. Suggestions on the way of regulating sowing and transplanting dates and making fertilization decision under low light intensity also put forward in this paper.

研究了不同生育期遮光45%(水稻生长的前期、中1期、中2期、后期)对水稻形态发育、光合产物积累、稻株养分含量、产量构成等的影响,并重点研究了植株对N、P、K养分的吸收规律。结果表明,各期遮光后均使干物质积累速率降低,植株N、P、K养分吸收量减少,对产量有不同程度的影响,但植株体内N、P、K养分含量上升。前期遮光主要使分蘖数急剧下降,有效穗数减少,叶面积系数下降,产量下降11.56%,但株高增加;中1期遮光后对水稻生长影响不大,产量仅降低5.46%;中2期遮光主要使每穗粒数和千粒重下降,对产量影响较大,降低幅度达30.80%;后期遮光主要影响结实粒和千粒重,其产量最低,降低55.40%.文中还对弱光照条件下的生育期安排和施肥决策提出了建议.

2000

0.0

... L3使植株抽穗至成熟阶段及成熟期的氮、磷、钾元素积累量显著下降, 最终产量显著下降, 同时氮肥利用率及氮、磷、钾元素籽粒生产效率也显著下降, 但下降幅度均低于L2, 主要因为拔节至抽穗阶段是水稻一生中氮、磷、钾元素积累的高峰期, 该阶段的养分积累与群体叶面积发展、颖花分化及最终产量密切相关^[13] ...

2012

0.0

1.667

. 2012, 38:648-656 DOI:10.3724/SP.J.1006.2012.00648

Root morphological and physiological characteristics of rice cultivars with high yield and high nitrogen use efficiency

1 Innovation Center of Rice Technology in Yangtze Rice Valley, Ministry of Agriculture / Key Laboratory of Crop Genetics and Physiology of Jiangsu Province, Yangzhou University, Yangzhou 225009, China; 2 Rice Research Institute of Guizhou Province, Guiyang 550006, China

The difference of root morphological and physiological characteristics of low-yielding and low N-efficiency, high-yielding and medium N-efficiency, high-yielding and high N-efficiency rice cultivars was investigated using six representative japonica varieties under their optimum N levels, respectively. The results showed that the high-yielding genotypes showed distinct advantages over the low-yielding ones in root dry weight, root volume, total and active absorbing surface areas of root system, root oxidation ability of α-NA and root bleeding intensity at each growth stage, indicating that the increase of productivity was accompanied by the improvement of root morphological traits and the enhancement of root physiological activities. Cultivars with different N-efficiency displayed diverse root morphological and physiological characteristics, even though they were all on a high-yielding level. Comparing with medium N-efficiency genotypes, the high N-efficiency ones showed lower root dry weight, root volume, root bleeding intensity and root total absorbing surface area for populations, but for single plants, the root dry weight, root volume, root bleeding intensity, active absorbing surface area and α-NA oxidation amount were superior in a significant or extremely significant degree. The results above suggest that the coordination of high-yielding with high N-efficiency could be achieved through controlling population growth properly, facilitating the concordant development of plant population and individuals, and endeavoring to improve the single stem root quality after heading.

选用低产氮低效型、高产氮中效型和高产氮高效型具有代表性的6个粳稻品种，在各自最适氮素水平下，研究了根系形态生理特征的差异。结果表明，较之低产类型品种，高产类型品种在根干重、根体积、根系总吸收表面积、活跃吸收表面积、根系α-NA氧化量及根系伤流强度等方面在各个生育时期均存在着明显的优势，说明生产力的提高伴随着根系形态特征的改善和生理活性的加强。同为高生产力类型品种，因氮利用率的差异根系形态生理特征表现不同。较之高产氮中效类型，高产氮高效型水稻的群体根干重、群体根体积、群体根系伤流强度和根系总吸收表面积均有所降低，而单茎根干重、单茎根体积、单茎根系伤流强度、活跃吸收表面积比及根系α-NA氧化量却有显著或极显著提高。表明适当控制高生产力水稻的群体生长量，促进群体和个体协调发展，着力提高抽穗后单茎根系质量，将是水稻高产和氮高效协调统一的可靠途径。

... 水稻根系与植株氮^[14,15,16]、磷^[17,18]、钾^[19,20]元素的吸收效率有密切的关系 ...

2008

0.0

1.667

. 2008, 34:429-436 DOI:10.3724/SP.J.1006.2008.00429

Root morphological and physiological characteristics in rice genotypes with different N use efficiencies

1 Jiangsu Province Key Laboratory of Crop Genetics and Physiology; 2 Innovation Center of Rice Cultivation Technology in Yangtze Valley, Ministry of Agriculture, Yangzhou University, Yangzhou 225009, Jiangsu, China <BR>

Nitrogen is the most important input required in rice production although over use of N causes so many environment problems. It is well known that N use efficiency is varied in different rice genotypes. Therefore, it is important to explore and exploit the potential contributing to the N use efficiency. As an integral part of plant organs, rice root plays an important role in absorption and utilization of N from soil and fertilizer. In this research, a field experiment with 225 kg ha-1 N fertilizer application was carried out in 2006 on the farm of Yangzhou University, Jiangsu province, China. A total of 12 rice genotypes ( 6 N-efficient and 6 N-inefficient ) selected from 120 rice cultivars grown in Yangzhou during 2004 and 2005 were adopted to investigate the charac-teristics of root morphology and physiology in rice types with different N use efficiencies. Relationship between N use efficiency and indexes of root morphology and physiology was also analyzed. The results showed that at different growth stages, the indexes of root morphology and physiology including the root dry weight, root volume, total absorbing surface area of root, active absorbing surface area of root, ratio of active absorbing surface area to total absorbing surface area in N efficient rice type were obviously higher than those in N inefficient rice type. At the critical stage of productive tillering and the stage of elongating, the ratio of root to shoot in N efficient rice type was significantly higher than that in N inefficient rice type while the trend was con-trary at the stages of heading and maturing. Before the stage of maturing, the root oxidation ability of α-NA in N efficient rice type was superior to that in N inefficiency rice type while at the stage of maturing the root oxidation ability of α-NA in N efficient hybrid rice was appreciably lower than that in some N efficient rice cultivars. N use efficiency positively correlated to all the above morphological and physiological indices at all growth stages, with the exception of negatively correlation to the ratio of root to shoot at the stage of heading and maturing. It could be concluded that the root morphology is good and the root is vigorous in N efficient rice type, its morphological and physiological characteristics ensures the efficient absorption and utilization of N in all its life, and proper ratio of root to shoot and the harmonious growth of root and shoot can also improve the efficiency of N absorption and utilization.

选用氮素利用高效型和低效型具有代表性的12个粳稻品种, 研究了225 kg hm-2施氮条件下根系形态和生理指标的差异及其与氮素利用效率的相互关系。结果表明, 有效分蘖临界叶龄期、拔节期、抽穗期和成熟期, 氮高效型水稻的根干重、根系体积、总吸收表面积、活跃吸收表面积和活跃吸收表面积比均显著或极显著大于氮低效型水稻。有效分蘖临界叶龄期和拔节期, 氮高效类型水稻的根冠比显著大于氮低效型, 而抽穗和成熟期则表现相反趋势。成熟期前, 氮高效型水稻的根系α-NA氧化量极显著大于氮低效型, 而成熟期, 氮高效型中杂交水稻的根系α-NA氧化量略低于个别氮低效型水稻品种。相关性分析表明, 有效分蘖临界叶龄期、拔节期、抽穗期和成熟期, 水稻的根干重、根系体积、根系的α-NA氧化量、总吸收表面积、活跃吸收表面积和活跃吸收表面积比与氮素利用效率呈显著或极显著正相关。有效分蘖临界叶龄期和拔节期, 植株的根冠比与氮素利用效率呈显著正相关, 而抽穗和成熟期则呈极显著负相关。氮高效型水稻在其一生中具有良好的根系形态和保持较强的根系活力, 为植株大量吸收和高效利用氮素奠定了良好基础。同时, 在水稻的生长过程中, 地下部与地上部的合理比例及协调生长也是促进植株高效吸收利用氮素的重要因素。

... 著影响, 如各生育阶段遮光均导致水稻产量显著下降^[9,15] ...

... 水稻根系与植株氮^[14,15,16]、磷^[17,18]、钾^[19,20]元素的吸收效率有密切的关系 ...

2012

0.0

1.889

. 2012, 45:4770-4781 DOI:10.3864/j.issn.0578-1752.2012.23.003

Root morphological and activity characteristics of rice genotype with high nitrogen utilization efficiency

College of Resource and Environmental Science, Sichuan Agricultural University, Chengdu 611130

【Objective】 The substantial reduction in fertilizer inputs and the selection of genotypes with high nutrient utilization efficiency or aggressive nutrient acquisition are beneficial to soil fertility in crop production by low-input approach. In order to provide a theoretical basis for cultivation regulation and breeding improvement of roots, roots morphological and activity characteristics of rice genotype (Oryza sativa) with different nitrogen utilization efficiencies (NUE) were investigated under different nitrogen treatments. 【Method】 Three pot culture experiments were carried out in a net room on the farm of Sichuan Agricultural University, Sichuan province, China in 2008 and 2009, including soil culture experiment with nitrogen treatments of 200 mg?kg-1, hydroponics of root morphological experiment with nitrogen treatments of 20 mg?L-1, 40 mg?L-1, and hydroponics of root physiological experiment with nitrogen treatments of 10 mg?L-1, 20 mg?L-1, 40 mg?L-1, 60 mg?L-1. In this research, root morphological characteristics of fine lateral root, coarse lateral root, and adventitious root were analyzed by root analysis system of WinRHIZO. The relationship between the root characteristics and nitrogen accumulation was also analyzed. 【Result】Rice grain yield and nitrogen utilization efficiency showed extremely significant genotype differences. Grain yield, nitrogen accumulation of grain, nitrogen dry matter production efficiency, nitrogen utilization efficiency and nitrogen harvest index of high NUE were 50.20%, 34.20%, 11.48%, 26.01% and 12.50% higher than those of low NUE. Dry matter weight, nitrogen accumulation and grain yield, nitrogen utilization of rice showed a significant or very significant positive correlation at jointing stage and heading stage. Under the low nitrogen treatment, a significant difference in root morphological characteristics of different types between high NUE and low NUE. Fine lateral root length of high NUE was 32.09%, 14.66%, 14.40% and 12.69% higher than low NUE at seeding, tillering, jointing and heading stages, respectively. Coarse lateral root surface area of high NUE was 94.70% and 64.38% higher than low NUE at jointing and heading stages, while root volume was 90.24% and 58.18% higher. Adventitious root length, surface area and volume of high NUE were 40.84%, 44.90% and 51.02% higher than those of low NUE at jointing stage. Root total absorbing surface area, active absorbing surface area and reducing capacity of high NUE decreased significantly with the increase of nitrogen at different growing stages. Under the same nitrogen treatment, root total absorbing surface area, active absorbing surface area, oxidation ability and reducing capacity of high NUE were 1.3-2.1 times, 1.1-3.2 times, 1.0-3.0 times and 1.4-2.2 times higher than those of low NUE. Under the low nitrogen treatment, length, surface area and volume of coarse lateral root had the greatest influence on nitrogen accumulation, the influence degree was 47.1-78.4%. 【Conclusion】 Good root morphology and physiological activity are the important characteristics of high NUE under the low nitrogen treatment. Rice genotype with high nitrogen utilization efficiency can be cultivated through improving the quality and quantity of coarse lateral root in order to obtain good root configuration.

【目的】研究水稻（Oryza sativa）氮高效利用基因型根系形态和活力变化，为根系的栽培调控和育种改良提供理论依据和技术参考。【方法】选择前期筛选出的水稻氮利用效率高低不一的基因型为试验材料，在比较氮利用效率基因型差异的基础上，采用水培试验，利用根系分析系统提取苗期至抽穗期不定根、粗分枝根和细分枝根的长度、表面积和体积等形态指标数据，探讨各类根形态与氮吸收的关系，同时分析氮高效利用基因型中典型材料不同供氮水平下根系活力变化。【结果】（1）水稻产量和氮利用效率呈现极显著的基因型差异，氮高效利用基因型籽粒产量、籽粒氮积累量、氮素干物质生产效率、氮素籽粒生产效率和氮素收获指数比低效基因型高50.20%、34.20%、11.48%、26.01%和12.50%。拔节期和抽穗期水稻干物质量、氮积累量与籽粒产量、氮素籽粒生产效率、氮素收获指数均呈现显著或极显著正相关，抽穗前（特别是拔节期和抽穗期）的物质积累和氮的吸收显著影响水稻产量和氮利用效率的提高。（2）低氮水平下，氮高、低利用效率基因型间的根系形态指标差异显著。细分枝根根长占水稻总根长的比重最大，为73.40%，且高效基因型在苗期、分蘖期、拔节期和抽穗期比低效基因型分别高32.09%、14.66%、14.40%和12.69%；粗分枝根表面积和体积分别占水稻总表面积和总体积29.81%和43.50%，其中高效基因型粗分枝根表面积在拔节期和抽穗期比低效基因型分别高94.70%和64.38%，体积分别高90.24%和58.18%；不定根根长、表面积和体积分别占水稻总根长、总表面积和总体积19.68%、36.66%和41.19%，且高效基因型不定根根长、表面积和体积在拔节期比低效基因型高40.84%、44.90%和51.02%，差异最大。（3）氮高效利用基因型根系吸收面积和还原力随着氮水平的提高显著降低，而氧化力变化不大。相同氮水平下，氮高效利用基因型拔节后总吸收面积、活跃吸收面积、氧化力、还原力分别为低效基因型的1.3—2.1倍、1.1—3.2倍、1.0—3.0倍、1.4—2.2倍。（4）低氮水平下，粗分枝根的根长、表面积和体积对氮积累量影响程度最大，为47.1%—78.4%。粗分枝根的发育情况直接影响氮的吸收，从而影响水稻产量和氮利用效率。【结论】低氮条件下良好的根系形态和生理活性是水稻氮高效利用的重要特征。培育氮高效利用基因型，可对水稻营养生长期根系形态和活性加以遗传改良，尤其是提高粗分枝根的比例，以期塑造良好的根系构型。

... 水稻根系与植株氮^[14,15,16]、磷^[17,18]、钾^[19,20]元素的吸收效率有密切的关系 ...

2003

0.0

1.883

. 2003, 9:420-424 DOI:10.11674/zwyf.2003.0408

Preliminary study on the utilization ability of insoluble phosphate in various rice cultivars

College of Agronomy;Jiangxi Agricultural University;Nanchang 330045; China

A horizontal division of root system was used to study the utilization ability of insoluble phosphate in low-phosphorus-tolerant cultivars(Ltz3) and l ow-phosphorus-sensitive cultivars(Xsbl). The results were as follows: Compare d to treatment with normal level phosphorus of solution in the lower pot, the dry matter weight, root oxidation decreased, MDA content increased in the treatment without phosphorus in whole system, but in the treatment with insoluble phosphat e in quartz in upper pot, the indexes changed more slightly and had higher phosp horus content, which suggested the active mobilization of phosphate of root had functioned; Low-phosphorus-tolerant cultivar had higher dry matter weight, roo t oxidation, phosphorus content and lower MDA content than low-phosphorus-sensit iv e cultivar, indicating its utilization ability of insoluble phosphate was strong er. The differences of each index among treatments with different insoluble phos phate might be related to the different availability of different insoluble phos phate on rice. The ratio of root to shoot increased under low phosphorus or phos phorus deficiency condition, and the lower phosphate availability, the ratio of root to shoot was higher.

采用子母盆栽水平分根试验,研究了水稻耐低磷品种莲塘早3号和低磷敏感品种新三百粒对Fe-P、Al-P、Ca-P3种难溶性磷的利用能力。结果表明,与母盆营养液中含正常磷的处理相比,子盆和母盆均不含磷的处理两品种的生物量、根系氧化力下降、MDA含量上升,以难溶性磷为唯一磷源加入子盆石英砂的处理其变化幅度更小,磷含量也比完全缺磷的明显要高,说明根系对难溶性磷的主动活化在起作用;耐低磷品种较高的生物量、磷含量和根系氧化力以及较低的MDA含量表明其活化利用难溶性磷的能力比低磷敏感品种强;不同难溶性磷处理间,各指标的差异与不同形态难溶性磷对水稻有效性的差异有关。另外,缺磷或低磷条件下根冠比增加;磷的有效性越低,其根冠比越高。

... 水稻根系与植株氮^[14,15,16]、磷^[17,18]、钾^[19,20]元素的吸收效率有密切的关系 ...

2003

0.0

1.742

. 2003, 14:61-65

Physiological adaptability of seeding rice genotypes with different P uptake effciency under low P-deficient stress

College of Crop Science, Fujian Agriculture and Forestry University, Fuzhou 350002, China

The genotypic differences in Puptake efficiency of three rice varieties (IR74, IR71331 and IR71379) were studied under the P-deficiency condition with hydroponics, and their adaptability to low phosphorus stress about physio-biochemical mechanisms was further studied. The results showed that rice genotypes tolerated low Pstress resulted from the co-ordination of Puptake efficiency, internal utilization efficiency and its translocation efficiency. The higher P-efficiency type IR74 and the middle type IR71331 had a higher Puptake efficiency. The rice genotype with higher P-efficiency was characterized by higher activity and desirable kinetic parameters of H₂PO₄^- uptake, showing lower K_m and C_min, but higher I_max values and relative APase activity,small amount of K_m and C_min. Moreover,under low Pstress, the activity of RN ase was about ten to fifteen times as high as that of the control (CK), but it had little genotypic differences.

以P效率差异显著的IR74(P高效型)、IR71331(中间型)及IR71379(P低效型)等3个品种为供试材料,采用水培法研究了这3类水稻在低P胁迫下P的吸收、转运及利用效率等P效率差异的原因以及P吸收动力学特征参数、R/S、酸性磷酸酯酶(acidphosphoesterase,简称APase)与核糖核酸酶(ribonuclease,简称RNase)等表现.结果表明,在低P胁迫下,P效率的高低是由水稻对P的吸收率、转运率以及利用效率综合作用的结果,存在基因型差异,P高效基因型IR74和P效率中间型的IR71331具有高的P吸收率,而IR71379P的吸收率低.不同P效率水稻在低P胁迫下,其K_m、C_min、I_max、R/S与Apase相对活性等参数表现出基因型的差异,小的K_m和C_min,大的I_max和R/S及高的APase升幅是水稻对P胁迫的生理适应性特征,也是植株在低P胁迫下较为敏感生理指标.其中各类型水稻叶片中RNase活性在低P胁迫下均表现大幅度上升,在品种间无显著差异,说明该酶是逆境胁迫并非低P胁迫的特征反应.

... 水稻根系与植株氮^[14,15,16]、磷^[17,18]、钾^[19,20]元素的吸收效率有密切的关系 ...

2003

0.0

1.883

. 2003, 9:190-195

Some agronomic and nutritional characteristics for potassium efficient rice genotypes under low potassium stress

采用溶液培养筛选,结合田间试验,提出了采用低钾胁迫下水稻体内钾利用效率作为衡量水稻钾利用效率的指标;探讨了钾高效基因型水稻的若干生长特性和营养特性;指出低钾胁迫导致水稻生物量减少, 植株生长缓慢,分蘖能力差,根系生长受到抑制,根系吸收的钠增加.水稻钾高效基因型低钾胁迫下仍具有较强的生长势(相对干重、相对株高、相对根长较大),其地上部钾/钠比值高而根部钾/钠比值较低,地上部和根部钾/氮吸收量比值较低.

... 水稻根系与植株氮^[14,15,16]、磷^[17,18]、钾^[19,20]元素的吸收效率有密切的关系 ...

2004

0.0

1.1628

. 2004, 5:356-359

Screening for low-kalium tolerance varieties at seedling stage from the core germplasm of integrated international rice

以全球水稻分子育种计划提供的117份核心种质资源为供试材料,进行苗期水培试验,比较两种不同K+浓度处理下的苗期根茎性状表现.结果表明:两种处理条件下,品种间存在显著差异;以发根数、总根长、苗高、地下部干重、地上部干重等5个性状相对指数的平均值构成品种的综合指数,作为筛选耐低钾品种的评判标准,结合稻苗生长情况,从117份核心种质资源中筛选出9个耐低钾品种和32个较耐低钾品种.

... 水稻根系与植株氮^[14,15,16]、磷^[17,18]、钾^[19,20]元素的吸收效率有密切的关系 ...

2012

0.0

0.687

... L3显著降低了植株抽穗后期的根系活力及根干重, 导致抽穗后期的氮、磷、钾元素积累量显著下降, 最终氮肥利用率及氮、磷、钾元素籽粒生产效率显著下降, 但下降幅度均低于L2, 主要由于抽穗后植株根系逐渐衰老及机能下降^[21] ...