不同氮肥水平下结实期灌溉方式对水稻弱势粒灌浆及产量的影响

doi:10.16819/j.1001-7216.2018.7060

摘要/Abstract

摘要： 【目的】

旨在阐明氮肥和灌溉方式对水稻产量、籽粒灌浆及生理特性的影响。

【方法】

以大穗型品种甬优2640和中穗型品种淮稻5号为供试材料进行盆钵试验,大田育秧移栽后设置3种氮肥水平,即0N(不施氮)、MN(2g N/盆)、HN(4g N/盆);抽穗至成熟期设置3种灌溉方式,即CI(保持水层灌溉)、WMD(轻干湿交替灌溉, 土壤水势 -15 kPa时复水)、WSD(重干湿交替灌溉, 土壤水势 -30 kPa时复水)。

【结果】

在CI下,两个品种产量均以MN水平最高;WMD处理下,两个品种产量均以HN水平最高,但与MN下差异不显著,WSD处理下两个品种产量均以HN最高;而在籽粒灌浆上,两个品种强势粒的灌浆速率和最终粒重在各个水氮处理间无显著差异,弱势粒的灌浆速率和最终粒重在良好水势条件CI和轻度水分胁迫WMD下,分别在0 N和MN水平下表现较优;但在重度水分胁迫WSD下,0N水平表现最低,HN最高,但与MN差异不显著。以上都表明产量与弱势粒的灌浆在水氮间存在着明显的交互作用。在品种间,大穗型籼粳杂交稻甬优2640弱势粒灌浆速率及粒重都低于中穗型常规粳稻淮稻5号,其产量优势主要源自较高的每穗粒数。最后,WMD+MN处理下有较高的氮肥利用率,较少的施氮量获得较高的产量,达到节水节氮增产的效果,其次也增加了根系生理活性和叶片光合性能,非结构性碳水化合物(NSC)转运率,促进了地上部的生长发育,同时也加强了物质运转,促进了灌浆中后期弱势粒籽粒的充实,最终达到产量增加的目的,成为本研究最佳水氮运筹方式。

关键词: 水稻, 产量, 强势粒, 弱势粒, 互作, 生理特性

Abstract:

【Objective】 To investigate the effects of water and nitrogen application on yield and grain filling,【Method】 an indica/japonica hybrid Yongyou 2640 with large-panicle and a japonica inbred Huaidao 5 with mid-panicle were grown in pots. After field seedling nursing then transplanting to pots, we designed three N rates, namely 0N (0 g N/pot), medium N level(MN, 2 g N/pot), and high N level(HN, 4 g N/pot), and three irrigation regimes post-anthesis consisting of conventional irrigation (CI, soil water potential was kept at 0 kPa), alternate wetting and moderate drying irrigation (WMD, rewatered when soil water potential reached -15 kPa), and alternate wetting and severe drying irrigation (WSD, rewatered when soil water potential reached -30 kPa). 【Result】 In the CI regime, MN showed the highest grain yield; in the WMD regimes, however, there was no significant difference in grain yield between MN and HN. Furthermore, in the WSD regime, grain yield under HN was the highest. In terms of grain filling, the superior spikelets present no significant difference in grain filling rate and final grain weight compared with all water-nitrogen treatments; Inferior spikeletsperforms better in grain filling rate and final grain weight at 0N and MN under CI and WMD regimes. However, in the WSD regime, 0N led to the lowest while HN showed the highest grain filling and final grain weight, but not significantly differentto MN. The above results showed that there was an obvious interaction between water and nitrogen. Among the varieties, grain filling rate and grain weight of inferior spikeletsof indica/japonica hybrid Yongyou 2640were lower than that ofjaponica inbred Huaidao 5, and the advantage of grain yield ofYongyou 2640 stemed from higher spikelet number per panicle. Finally, in the WMD+MN treatment, there was a higher nitrogen efficiency, creating higher grain yield with less nitrogen, and achieving the purpose of water and nitrogen saving. Secondly, it also eahanced the activity of root and leaves, improving the non-structural carbohydrate(NSC) remobilization, which promoted the upground biomass development and dry matter translocation in order to strengthen the inferior spikelets filling to induce an increase in grain yield. Therefore, it turns out to be the best water-nitrogen management in this research.

Key words: rice, yield, superior spikelets, inferior spikelets, interaction, physiological traits

中图分类号:

朱宽宇, 展明飞, 陈静, 王志琴, 杨建昌, 赵步洪. 不同氮肥水平下结实期灌溉方式对水稻弱势粒灌浆及产量的影响[J]. 中国水稻科学, 2018, 1(1): 155-168.

Kuanyu ZHU, Mingfei ZHAN, CHENJing, Zhiqin WANG, Jianchang YANG, Buhong ZHAO. Effects of Irrigation Regimes DuringGrain Filling Under Different Nitrogen Rates on Inferior SpikeletsGrain-Filling and Grain Yield of Rice[J]. Chinese Journal OF Rice Science, 2018, 1(1): 155-168.

图/表 11

表1 产量、籽粒灌浆及其生理特性在年度间、品种间及处理间的方差分析

Table 1 Analysis-of-variance (F-values) for grain yield, grain filling characteristics and physiologicaltraits of riceamong years, cultivars and treatments.

变异来源 Sourceofvariance	自由度df	产量 Yield	平均灌浆速率 Mean of grain filling rate	最终粒重 Final weight	NSC转运率 NSC remobiliza -tion rate	根系氧化力 Root oxidation	光合速率 Photosynthe -tic rate	蒸腾速率 Transpiration rate		根系IAA含量 IAA content of Root
年度Year(Y)	1	7.95^ns	8.58^ns	2.45^ns	3.32^ns	4.67^ns	4.77^ns	19.81^ns	16.35^ns		12.89^ns
品种Cultivar(C)	1	63.65^**	141.36^**	71.91^**	20.99^**	25.68^**	46.93^**	98.32^**	33.66^**		34.78^**
灌溉Irrigation(I)	2	18.98^**	148.92^**	89.15^**	28.22^**	86.82^**	48.90^**	484.03^**	14.15^**		16.52^**
氮肥Nitrogen(N)	2	15.09^**	24.66^*	54.88^**	11.62^**	53.53^*	54.91^*	5.66^*	40.68^**		42.26^**
Y×C	1	138.98^ns	148.92^ns	89.15^ns	202.22^ns	486.82^ns	486.90^ns	484.03^ns	12.15^ns		616.52^ns
Y×I	2	8.99^ns	6.08^ns	11.43^ns	102.62^ns	68.52^ns	57.14^ns	2.87^ns	59.37^ns		51.42^ns
Y×N	2	30.90^ns	11.82^ns	96.13^ns	22.54^ns	78.65^ns	33.64^ns	49.25^ns	25.45^ns		22.06^ns
C×I	2	99.57^**	0.35^ns	0.02^ns	0.95^ns	1.23^ns	1.22^ns	1.22^ns	64.53^**		64.56^**
C×N	2	152.52^**	0.21^ns	3.32^*	17.55^**	140.56^*	140.58^*	139.75^**	27.71^**		27.74^**
I×N	4	97.29^**	18.00^**	66.65^**	1.29^ns	0.51^ns	0.50^ns	0.50^ns	4.21^**		4.25^**
Y×C×I	2	2.68^ns	0.09^ns	0.93^ns	0.82^ns	0.05^ns	0.72^ns	0.01^ns	0.23^ns		0.21^ns
Y×C×N	2	3.62^ns	0.08^ns	0.97^ns	6.75^ns	5.23^ns	0.43^ns	0.83^ns	0.10^ns		0.09^ns
C×I×N	4	11.38^**	0.18^ns	2.05^ns	5.54^*	10.15^**	10.16^**	10.10^**	0.02^ns		0.02^ns
Y×I×N	4	2.11^ns	0.03^ns	0.90^ns	5.32^ns	44.24^ns	0.81^ns	0.11^ns	0.02^ns		0.01^ns
Y×C×I×N	4	3.92^ns	0.04^ns	0.92^ns	112.52^ns	68.93^ns	42.34^ns	60.52^ns		33.21^ns	8.97^ns

表2 不同氮肥水平下结实期灌溉方式对水稻产量及其构成因素的影响

Table 2 Effects of irrigation regimes during grain filling under different nitrogen rates on the yield components of rice.

品种 Cultivar	处理 Treatment	每盆穗数 No.of panicle per pot	每穗粒数 Spikelet number per panicle	结实率 Seed setting rate /%	千粒重 1000-grain weight /g	产量 Grain yield /g·pot^-1
甬优2640 Yongyou 2640	CI+0N	11.2c	229.5c	90.5c	24.5b	57.0g
甬优2640 Yongyou 2640	CI+MN	18.7b	276.3b	89.5d	24.6b	114.3b
	CI+HN	21.2a	289.4a	75.9f	23.6c	110.2c
	WMD+0N	11.2c	230.1c	93.8a	25.5a	61.7f
	WMD+MN	18.6b	276.8b	93.2b	25.6a	122.7a
	WMD+HN	21.2a	289.2a	80.1e	25.4a	124.8a
	WSD+0N	11.4c	231.9c	74.7h	22.9e	44.7h
	WSD+MN	18.5b	274.9b	75.3g	23.4d	91.4e
	WSD+HN	21.1a	288.5a	75.5g	23.0d	105.7d
淮稻5号 Huaidao 5	CI+0N	13.8c	141.5c	91.9b	27.1b	48.6g
淮稻5号 Huaidao 5	CI+MN	23.0b	171.6b	91.7b	27.2b	97.0b
	CI+HN	24.2a	177.9a	80.7d	26.4c	91.7c
	WMD+0N	13.9c	141.9c	92.9a	27.7a	50.7f
	WMD+MN	22.8b	171.3b	92.5a	27.5a	99.9a
	WMD+HN	24.2a	177.0a	85.4c	27.6a	100.9a
	WSD+0N	13.7c	141.6c	80.1e	25.1e	39.0h
	WSD+MN	22.6b	169.8b	81.9d	25.8d	81.1e
	WSD+HN	24.0a	175.6a	82.2d	25.9d	89.7d

图1 不同氮肥水平下结实期灌溉方式对水稻氮肥利用率和收获指数的影响 CI–常规灌溉;WMD–轻干湿交替灌溉;WSD–重干湿交替灌溉; MN–中氮;HN–高氮。

Fig.1. Effects of irrigation regimes during grain filling under different nitrogen rates on the nitrogen use efficiency and harvest index of rice. CI,Conventional irrigation; WMD,Alternate wetting and moderate soil drying; WSD,Alternate wetting and severe soil drying.MN,Medium nitrogen level; HN,High nitrogrn level.

图2 不同氮肥水平下结实期灌溉方式对水稻强弱势粒增重状态的影响 CI+0N–常规灌溉+不施氮;CI+MN–常规灌溉+中氮;CI+HN–常规灌溉+高氮;WMD+0N–轻干湿交替灌溉+不施氮; WMD+MN–轻干湿交替灌溉+中氮;WMD+HN–轻干湿交替灌溉+高氮;WSD+0N–重干湿交替灌溉+不施氮;WSD+MN–重干湿交替灌溉+中氮;WSD+HN–重干湿交替灌溉+高氮。S–强势粒;I–弱势粒。

Fig.2. Effects of irrigation regimes during grain filling under different nitrogen rates on grain weight of rice. CI+0N, Conventional irrigation+0 nitrogen fertilizer;CI+MN,Conventional irrigation+mediumnitrogen level;CI+HN,Conventional irrigation+high nitrogen level;WMD+0N,Alternate wetting and moderate soil drying+0 nitrogen fertilizer;WMD+MN,Alternate wetting and moderate soil drying+medium nitrogen level;WMD+HN,Alternate wetting and moderate soil drying+medium nitrogen level;WSD+0N,Alternate wetting and severe soil drying+0 nitrogen fertilizer;WSD+MN,Alternate wetting and severe soil drying+medium nitrogen level;WSD+HN,Alternate wetting and severe soil drying+high nitrogen level. S, Superior spikelets; I, Inferior spikelets.

表3 不同氮肥水平下结实期灌溉方式对水稻籽粒灌浆参数的影响

Table 3 Effects of irrigation regimes during grain filling under different nitrogen rates on the grain filling characters of rice.

品种 Variety	处理 Treament	最大灌浆速率 G_max/(mg·grain^-1d^-1)		达最大灌浆速率时间 T_max/d		平均灌浆速率 G_mean/(mg·grain^-1d^-1)		活跃灌浆期 Active filling period/d		最终粒重A /(mg·grain^-1)
品种 Variety	处理 Treament	S	I	S	I	S	I	S	I	S	I
甬优2640	CI+0N	2.08a	1.16ab	14.37a	33.42f	1.49a	0.54b	18.43a	30.13d	27.46a	16.27c
Yongyou 2640	CI+MN	2.10a	1.13ab	14.42a	35.88e	1.48a	0.53b	18.52a	31.26c	27.41a	16.56c
	CI+HN	2.04a	1.02ab	14.83b	41.86b	1.46a	0.46c	18.55a	33.63a	27.08a	15.46d
	WMD+0N	2.08a	1.23a	14.38a	31.32g	1.50a	0.61a	18.30a	28.28e	27.45a	17.25b
	WMD+MN	2.10a	1.18ab	14.40a	32.12g	1.51a	0.62a	18.33a	30.12d	27.68a	18.67a
	WMD+HN	2.08a	1.15ab	14.50a	38.37d	1.47a	0.59a	18.46a	32.58b	27.14a	18.63a
	WSD+0N	2.05a	0.88c	14.95b	37.51d	1.47a	0.33e	18.35b	22.65g	26.97a	7.47f
	WSD+MN	2.11a	1.02b	14.45a	40.32c	1.48a	0.40d	18.39a	27.95f	27.22a	11.18e
	WSD+HN	2.09a	1.04b	14.52a	44.68a	1.46a	0.41cd	18.42a	28.12f	26.89a	11.52e
淮稻5号	CI+0N	2.26a	1.32ab	15.35a	26.56h	1.68a	0.80b	17.54a	24.26c	29.47a	19.40b
Huaidao 5	CI+MN	2.23a	1.27ab	15.38a	28.48e	1.70a	0.76b	17.56a	25.71b	29.85a	19.53b
	CI+HN	2.20b	1.2ab	15.37a	30.15b	1.69a	0.70c	17.65a	26.33a	29.83a	18.43c
	WMD+0N	2.28a	1.43a	15.38a	26.09i	1.70a	0.89a	17.52a	23.16d	29.78a	21.32a
	WMD+MN	2.25a	1.37ab	15.40a	27.31g	1.69a	0.91a	17.58a	24.21c	29.71a	22.03a
	WMD+HN	2.23a	1.24ab	15.42a	29.12d	1.67a	0.87a	17.68a	25.32b	29.53a	22.02a
	WSD+0N	2.22a	0.95c	15.33a	27.68f	1.71a	0.54e	17.42a	19.86f	29.79a	10.72e
	WSD+MN	2.24a	1.13b	15.41a	29.66c	1.70a	0.63d	17.54a	22.56e	29.82a	14.21d
	WSD+HN	2.19a	1.15b	15.42a	31.85a	1.69a	0.61d	17.52a	23.18d	29.61a	14.14d

表3 不同氮肥水平下结实期灌溉方式对水稻籽粒灌浆参数的影响

Table 3 Effects of irrigation regimes during grain filling under different nitrogen rates on the grain filling characters of rice.

品种 Variety	处理 Treament	最大灌浆速率 G_max/(mg·grain^-1d^-1)		达最大灌浆速率时间 T_max/d		平均灌浆速率 G_mean/(mg·grain^-1d^-1)		活跃灌浆期 Active filling period/d		最终粒重A /(mg·grain^-1)
品种 Variety	处理 Treament	S	I	S	I	S	I	S	I	S	I
甬优2640	CI+0N	2.08a	1.16ab	14.37a	33.42f	1.49a	0.54b	18.43a	30.13d	27.46a	16.27c
Yongyou 2640	CI+MN	2.10a	1.13ab	14.42a	35.88e	1.48a	0.53b	18.52a	31.26c	27.41a	16.56c
	CI+HN	2.04a	1.02ab	14.83b	41.86b	1.46a	0.46c	18.55a	33.63a	27.08a	15.46d
	WMD+0N	2.08a	1.23a	14.38a	31.32g	1.50a	0.61a	18.30a	28.28e	27.45a	17.25b
	WMD+MN	2.10a	1.18ab	14.40a	32.12g	1.51a	0.62a	18.33a	30.12d	27.68a	18.67a
	WMD+HN	2.08a	1.15ab	14.50a	38.37d	1.47a	0.59a	18.46a	32.58b	27.14a	18.63a
	WSD+0N	2.05a	0.88c	14.95b	37.51d	1.47a	0.33e	18.35b	22.65g	26.97a	7.47f
	WSD+MN	2.11a	1.02b	14.45a	40.32c	1.48a	0.40d	18.39a	27.95f	27.22a	11.18e
	WSD+HN	2.09a	1.04b	14.52a	44.68a	1.46a	0.41cd	18.42a	28.12f	26.89a	11.52e
淮稻5号	CI+0N	2.26a	1.32ab	15.35a	26.56h	1.68a	0.80b	17.54a	24.26c	29.47a	19.40b
Huaidao 5	CI+MN	2.23a	1.27ab	15.38a	28.48e	1.70a	0.76b	17.56a	25.71b	29.85a	19.53b
	CI+HN	2.20b	1.2ab	15.37a	30.15b	1.69a	0.70c	17.65a	26.33a	29.83a	18.43c
	WMD+0N	2.28a	1.43a	15.38a	26.09i	1.70a	0.89a	17.52a	23.16d	29.78a	21.32a
	WMD+MN	2.25a	1.37ab	15.40a	27.31g	1.69a	0.91a	17.58a	24.21c	29.71a	22.03a
	WMD+HN	2.23a	1.24ab	15.42a	29.12d	1.67a	0.87a	17.68a	25.32b	29.53a	22.02a
	WSD+0N	2.22a	0.95c	15.33a	27.68f	1.71a	0.54e	17.42a	19.86f	29.79a	10.72e
	WSD+MN	2.24a	1.13b	15.41a	29.66c	1.70a	0.63d	17.54a	22.56e	29.82a	14.21d
	WSD+HN	2.19a	1.15b	15.42a	31.85a	1.69a	0.61d	17.52a	23.18d	29.61a	14.14d

图3 不同氮肥水平下结实期灌溉方式对水稻氮根系氧化力的影响 CI–常规灌溉;WMD–轻干湿交替灌溉;WSD–重干湿交替灌溉;0N–不施氮;MN–中氮;HN–高氮。

Fig.3. Effects of irrigation regimes during grain filling under different nitrogen rates on the root oxidation activity of rice. CI,Conventional irrigation; WMD,Alternate irrigation with wetting and moderate drying; WSD,Alternate irrigation with wetting and severedrying; 0N,Zero nitrogenfertilizer; MN,Medium nitrogen level; HN,High nitrogen level.

图4 不同氮肥水平下结实期灌溉方式对水稻氮根系激素的影响 CI–常规灌溉;WMD–轻干湿交替灌溉;WSD–重干湿交替灌溉;0N–不施氮;MN–中氮;HN–高氮。

Fig.4. Effects of irrigation regimes during grain filling under different nitrogen rates on the root hormone of rice. CI,Conventional irrigation; WMD,Alternate irrigation with wetting and moderate drying; WSD,Alternate irrigation with wetting and severe drying; 0N, 0 nitrogenfertilizer; MN,Medium nitrogen level; HN,High nitrogen level.

图5 不同氮肥水平下结实期灌溉方式对水稻叶片光合速率(A-F)和蒸腾速率(G-L)的影响 CI–常规灌溉;WMD–轻干湿交替灌溉;WSD–重干湿交替灌溉;0N–不施氮;MN–中氮;HN–高氮。

Fig.5. Effects of irrigation regimes during grain filling under different nitrogen rates on leaf photosynthesis(A-F) and transpiration(G-L) of rice. CI,Conventional irrigation; WMD,Alternate irrigation with wetting and moderate drying; WSD,Alternate irrigation with wetting and severe drying; 0N, 0 nitrogenfertilizer; MN,Medium nitrogen level; HN,High nitrogen level.

图6 不同氮肥水平下结实期灌溉方式对水稻茎秆中非结构性碳水化合物NSC含量(A~F)、转运率(C, G)、贡献率(D, H)的影响 CI–常规灌溉;WMD–轻干湿交替灌溉;WSD–重干湿交替灌溉;0N–不施氮;MN–中氮;HN–高氮。

Fig.6. Effects of irrigation regimes during grain filling under different nitrogen rates on non-structural carbohydrate(NSC) content(A-F), remobilization(C, G), accumulation(D, H) of stem in rice. CI,Conventional irrigation; WMD,Alternate irrigation with wetting and moderate drying; WSD,Alternate irrigation with wetting and severe drying; 0N, 0 nitrogenfertilizer; MN,Medium nitrogen level; HN,High nitrogen level.

表4 产量构成因素及生理特性对产量的贡献

Table 4 Contributions of yield components and physiological traits to grain yield.

产量构成及生理指标 Yield components and physiological traits	直接作用 Direct contribution	间接作用 Indirect contribution
产量构成及生理指标 Yield components and physiological traits	直接作用 Direct contribution	x₁	x₂	x₃	x₄	x₅	x₆	x₇	x₈
x₁	0.079		0.645	-0.083	-0.002	-0.045	-0.005	-0.027	-0.049
x₂	0.647	0.078		-0.081	-0.001	-0.036	-0.003	-0.019	-0.045
x₃	0.214	-0.031	-0.243		0.029	-0.040	0.030	0.017	0.084
x₄	-0.036	-0.004	-0.031	0.174		-0.002	-0.002	-0.013	0.021
x₅	0.543	-0.007	-0.043	-0.072	-0.001		0.026	0.156	-0.018
x₆	0.038	-0.009	-0.056	0.016	-0.002	0.376		0.225	0.032
x₇	0.225	-0.009	-0.056	0.016	-0.002	0.376	0.226		0.032
x₈	0.172	-0.022	-0.171	0.106	0.005	-0.288	0.007	0.042

图7 弱势粒灌浆速率和粒重与结实率和千粒重的相关性**表示在0.01水平上显著。

Fig.7. Correlations between grain filling rate, final weight of inferior spikelets and seed setting rate, 1000-grain weight. ** Significant at 0.01 level.

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