徐国伟^{1, 2,,},
陆大克¹,
王贺正¹,
贾付俊¹,
陈明灿¹
1.河南科技大学农学院 洛阳 471003
2.扬州大学江苏省作物遗传生理重点实验室 扬州 225009
基金项目: 国家自然科学基金项目U1304316
河南省教育厅科学技术研究重点项目13A210266
江苏省作物栽培生理重点实验室开放基金027388003K11009
河南科技大学学科提升计划A13660002

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作者简介:徐国伟, 主要从事作物栽培生理研究。E-mail:gwxu2007@163.com

中图分类号:S312

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收稿日期:2017-09-30
录用日期:2017-12-29
刊出日期:2018-04-01

Coupling effect of alternate wetting and drying irrigation and nitrogen rate on organic acid in rice root secretion at heading stage

XU Guowei^{1, 2,,},
LU Dake¹,
WANG Hezheng¹,
JIA Fujun¹,
CHEN Mingcan¹
1. College of Agriculture, 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
Funds: the National Natural Science Foundation of ChinaU1304316
the Key Project for Science and Technology Research of Department of Education of Henan Province13A210266
the Open Foundation for Key Laboratory of Cultivation and Physiology of Jiangsu Province027388003K11009
the Henan University of Science and Technology Discipline Improvement and Promotion Plan A13660002

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Corresponding author:XU Guowei, E-mail:gwxu2007@163.com

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摘要
摘要:以水稻品种‘连粳7号’为试验材料进行盆栽试验，设置不施氮（0N，0 kg·hm^-2）、中氮（MN，240 kg·hm^-2）和高氮（HN，360 kg·hm^-2）3种施氮水平及浅水层灌溉（0 kPa）、轻度干湿交替灌溉（-20 kPa）和重度干湿交替灌溉（-40 kPa）3种灌溉方式，研究不同水氮处理对水稻抽穗期根系分泌有机酸总量和组分变化、氨基酸含量及水稻氮肥农学利用率与偏生产力的影响及其耦合效应，探索水氮耦合机理，为水稻氮素高效利用及根际生态提供理论及科学依据。结果表明：轻度干湿交替灌溉增加了水稻根系酒石酸、柠檬酸、草酸、苹果酸、琥珀酸、总有机酸、氨基酸分泌量，分别较浅水层灌溉增加13.2%、8.7%、27.3%、40.0%、6.7%、6.3%及6.4%，水稻氮肥农学利用效率及偏生产力分别增加4.1%及1.7%，显著提高根系分泌有机酸及氨基酸含量；重度干湿交替灌溉减少水稻根系酒石酸、柠檬酸、草酸、苹果酸、琥珀酸的分泌量，显著降低根系分泌有机酸总量、氨基酸含量及水稻的氮肥利用效率。同一水分条件下，施氮显著促进根系酒石酸、乙酸、苹果酸、琥珀酸的分泌，降低了草酸和柠檬酸的分泌量。根系分泌的酒石酸和琥珀酸含量在MN与HN间差异较小。分析表明，根系分泌有机酸总量、氨基酸、苹果酸及琥珀酸的供氮效应为正效应，轻度干湿交替灌溉效应及与供氮的耦合效应为正效应，而重度干湿交替灌溉效应及其与供氮的耦合效应则为负效应。根系分泌的柠檬酸、草酸与氮肥利用率呈显著与极显著正相关，乙酸与氮肥利用间呈显著负相关。结果表明通过轻度干湿交替灌溉与中等施氮调控发挥水肥耦合效应，可以促进水稻根系酒石酸、苹果酸、琥珀酸及氨基酸分泌，提高氮肥利用效率，从而促进水稻高产。
关键词:水稻/
干湿交替灌溉/
水氮耦合/
根系分泌物/
有机酸/
氮肥利用效率
Abstract:Soil water and nitrogen conditions are the principal factors that affect crop growth and formation. Clarifying effects of soil water and nitrogen conditions and their coupling on grain yield of rice (Oryza sativa L.) have significant implications for high production efficiency and yield. There is extensive domestic and foreign research on the interactions of soil water and fertilizer. Specific focus has been put on ground parameters such as crop growth development, dry shoot formation, physiological function, hormone change, nutrient absorption and utilization, water use efficiency, etc. However, there is less work on the characteristics of root secretion, including changes in organic acids in roots. Research on the relationship between organic acid content and nitrogen utilization and the related interactions is also inconsistent and therefore inconclusive. The purpose of this study was to investigate the performance of root secretion under different nitrogen and water regimes and analyze the effect of the interaction of alternate wetting and drying irrigation and nitrogen levels. To that end, a pot-soil experiment was conducted using a mid-season japonica rice cultivar 'Lianjing 7' under different nitrogen rates[0N, MN (240 kg·hm^-2) and HN (360 kg·hm^-2)] and three irrigation regimes[keeping water (0 kPa), alternate wetting and moderate drying (-20 kPa) and alternate wetting and severe drying (-40 kPa)] in 2015 and 2016. The results showed that there was remarkable interaction between nitrogen rate and irrigation regime with the results similar for the two years. Under the same nitrogen rate, tartaric acid, citric acid, oxalic acid, malic acid, succinic acid, total organic acid and amino acid secretion of rice root at heading stage were higher (respectively by 13.2%, 8.7%, 27.3%, 40.0%, 6.7%, 6.3% and 6.4%) in treatment of alternate wetting and moderate drying than in treatment of keeping water flood irrigation. The nitrogen agronomic utilization and partial productivity of rice were increased by 4.1% and 1.7% respectively. The secretion amounts of tartaric acid, citric acid, oxalic acid, malic acid and succinic acid at heading stage decreased (respectively by 16.4%, 4.5%, 12.8%, 41.7% and 5.6%) under the treatment of alternate wetting and severe drying than under the treatment of keeping water. This also reduced total organic acid and amino acid in roots and overall nitrogen use efficiency. Under the same irrigation condition, nitrogen application accelerated the root secretion of tartaric acid, acetic acid, malic acid and succinic acid, but decreased the secretion of oxalic acid and citric acid at heading stage of rice. The differences in tartaric acid, acetic acid, malic acid and succinic acid secretion by roots were small between MN and HN treatments. Nitrogen application has positive effects on total organic acid, amino acid, malic acid, succinic acid secretion. Also its interaction effect with alternate wetting and moderate drying was positive, while the interaction effect with alternate wetting and sever was negative. Correlation analysis indicated that citric and oxalic acids secretion by rice roots was significantly positively correlated with nitrogen utilization. However, acetic acid was negatively correlated with nitrogen use efficiency. The results of the study suggest that increasing root secretion capacity through appropriate regulation of irrigation coupled with nitrogen rate increased nitrogen use efficiency and thereby promoted high yield of rice.
Key words:Rice/
Alternate wetting and drying irrigation/
Water and nitrogen interaction/
Root secretion/
Organic acid/
Nitrogen effi-ciency

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图1干湿交替灌溉与施氮量耦合对水稻抽穗期根系分泌有机酸总量及氨基酸含量的影响
0N:不施氮肥; MN:施氮肥240 kg(N)∙hm^–2; HN:施氮肥360 kg(N)∙hm^–2。不同字母表示不同处理间0.05水平差异显著。
Figure1.Effect of alternate wetting and drying irrigation and nitrogen application on total organic acid and amino acid contents in root secretion at heading stage of rice
0N: no nitrogen application; MN: nitrogen application rate of 240 kg(N)∙hm^–2; HN: nitrogen application rate of 360 kg(N)∙hm^–2. Different letters mean significant differences among treatments at P < 0.05.


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图2干湿交替灌溉与施氮量耦合对水稻抽穗期根系分泌物有机酸含量的影响
0N:不施氮肥; MN:施氮肥240 kg(N)∙hm^–2; HN:施氮肥360 kg(N)∙hm^–2。不同字母表示不同处理间0.05水平差异显著。
Figure2.Effects of alternate wetting and drying irrigation and nitrogen application on organic acids contents in root secretion of rice at heading stage
0N: no nitrogen applied; MN: nitrogen 240 kg(N)∙hm^–2; HN: nitrogen 360 kg(N)∙hm^–2. Different letters mean significant differences among treatments at P < 0.05.


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图3水稻抽穗期根系分泌的有机酸与氮素农学利用率的相关性
Figure3.Correlations between organic acids in root secretion at heading stage with nitrogen agronomic efficiency of rice


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图4水稻抽穗期根系分泌的有机酸与氮素偏生产力的相关性分析
Figure4.Correlations of organic acids in root secretion at heading stage with nitrogen partial factor productivity of rice


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表1干湿交替灌溉与施氮量耦合对水稻根系分泌物有机酸总量及氨基酸效应的影响
Table1.Interaction effect of alternate wetting and drying irrigation and nitrogen application on total organic acid and amino acid contents of rice root secretion at heading stage

项目 Item	效应 Effect	MN			HN
		轻度干湿交替灌溉 Alternate wetting and moderate drying irrigation	重度干湿交替灌溉 Alternate wetting and severe drying irrigation		轻度干湿交替灌溉 Alternate wetting and moderate drying irrigation	重度干湿交替灌溉 Alternate wetting and severe drying irrigation
有机酸总量 Total organic acid content	供氮效应N supplied effect	10.52a	7.03b		9.18a	5.19b
	控水效应Water stress effect	1.72a	-4.14b		1.54a	-4.84b
	耦合效应Coupling effect	1.36a	-2.12b		1.18a	-2.82b
氨基酸 Amino acid content	供氮效应N supplied effect	1.34a	0.90b		1.17a	0.66b
	控水效应Water stress effect	0.22a	-0.53b		0.20a	-0.62b
	耦合效应Coupling effect	0.17a	-0.27b		0.15a	-0.36b
MN:施氮肥240 kg(N)?hm–2; HN:施氮肥360 kg(N)?hm–2。同行不同字母表示0.05水平差异显著。MN: nitrogen 240 kg(N)?hm–2; HN: nitrogen 360 kg(N)?hm–2. Values in the same line followed by different letters are significantly different at P < 0.05.

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表2抽穗期水氮耦合对水稻根系分泌物有机酸效应的影响
Table2.Interaction effects of alternate wetting and drying irrigation and nitrogen application on organic acids of root secretion of rice at heading stage

项目 Item	效应 Effect	MN			HN
		轻度干湿交替灌溉 Alternate wetting and moderate drying irrigation	重度干湿交替灌溉 Alternate wetting and severe drying irrigation		轻度干湿交替灌溉 Alternate wetting and moderate drying irrigation	重度干湿交替灌溉 Alternate wetting and severe drying irrigation
酒石酸 Tartaric acid	供氮效应N supplied effect	0.09a	0.09a		0.10a	0.08a
	控水效应Water stress effect	0.04a	-0.13b		0.04a	-0.15b
	耦合效应Coupling effect	0.05a	-0.03c		0.01b	-0.02bc
柠檬酸 Citric acid	供氮效应N supplied effect	-0.02a	-0.02a		-0.01a	-0.06b
	控水效应Water stress effect	0.05a	-0.05b		0.06a	-0.04b
	耦合效应Coupling effect	0.01b	0.01b		0.05a	0.01b
乙酸 Acetic acid	供氮效应N supplied effect	0.01a	0.01a		0.01a	0.04a
	控水效应Water stress effect	-0.01a	-0.01a		-0.02a	-0.04a
	耦合效应Coupling effect	0.01a	0.02a		-0.01a	0.02a
草酸 Oxalic acid	供氮效应N supplied effect	-0.04b	-0.01a		-0.06b	-0.06b
	控水效应Water stress effect	0.09b	-0.12c		1.00a	-0.15c
	耦合效应Coupling effect	0.05a	0.03a		0.01a	0.01a
苹果酸 Malic acid	供氮效应N supplied effect	0.16a	0.09b		0.13a	0.06b
	控水效应Water stress effect	0.10b	-0.09c		1.00a	-0.09c
	耦合效应Coupling effect	0.04a	-0.04b		0.04a	-0.04b
琥珀酸 Succinic acid	供氮效应N supplied effect	0.12b	0.06c		1.00a	0.04c
	控水效应Water stress effect	0.09a	-0.04b		0.09a	-0.03b
	耦合效应Coupling effect	0.06a	-0.01b		0.06a	-0.03b
MN:施氮肥240 kg(N)?hm–2; HN:施氮肥360 kg(N)?hm–2。同行不同字母表示0.05水平差异显著。MN: nitrogen application rate of 240 kg(N)?hm–2; HN: nitrogen application rate of 360 kg(N)?hm–2. Values in the same line followed by different letters are significantly different at P < 0.05.

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表3干湿交替灌溉与施氮量耦合对水稻产量及氮素利用的影响
Table3.Effect of alternate wetting and drying irrigation and nitrogen application on grain yield and nitrogen use efficiency of rice

处理Treatment		每盆施氮量 Total N fertilizer per pot (g)	每盆水稻产量 Rice yield per pot (g)	氮肥农学利用率 Agronomic efficiency of N fertilizer [kg(grain)·kg-1]	氮肥偏生产力 Partial factor productivity of N fertilizer [kg(grain)·kg-1]
施氮量 N application (g?kg-1)	灌溉 Irrigation
0	保持水层 Keeping water	0	31.7±1.5d	—	—
	轻度干湿交替灌溉 Alternate wetting and moderate drying irrigation	0	32.3±1.7d	—	—
	重度干湿交替灌溉 Alternate wetting and severe drying irrigation	0	24.1±1.0f	—	—
240	保持水层 Keeping water	1.77	51.9±2.4a	24.2±0.8a	44.1±1.8a
	轻度干湿交替灌溉 Alternate wetting and moderate drying irrigation	1.77	52.7±2.6a	25.2±1.2a	44.8±2.3a
	重度干湿交替灌溉 Alternate wetting and severe drying irrigation	1.77	37.6±1.5c	16.3±0.6b	32.0±1.4b
360	保持水层 Keeping water	2.65	50.6±2.3ab	15.1±0.7b	28.6±1.2c
	轻度干湿交替灌溉 Alternate wetting and moderate drying irrigation	2.65	51.3±2.5ab	15.7±1.0b	29.1±0.6c
	重度干湿交替灌溉 Alternate wetting and severe drying irrigation	2.65	29.1±1.1e	4.0±0.4c	16.5±0.9d
同列不同字母表示0.05水平差异显著。Values in the same column followed by different letters are significantly different at 0.05 level.

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