氮素缓解春小麦花后高温早衰的荧光特性研究

doi:10.3864/j.issn.0578-1752.2021.15.018

摘要/Abstract

摘要：

【目的】为了研究高温条件下氮素对春小麦荧光特性影响机制,阐明氮素调控光反应中心的内在机理,从而制定缓解高温危害的氮肥运筹措施。【方法】于2019年和2020年开展田间试验,采用裂区试验设计,主区为5个施氮量,依次为0（N0）、75 kg·hm^-2（N1）、150 kg·hm^-2(N2）、225 kg·hm^-2（N3）、300 kg·hm^-2（N4）,副区为温度,分别为25℃±2℃（CK）和35℃±2℃（HT）。分析花后不同温度处理下施氮量与叶片含氮量、叶绿素、光系统Ⅱ(PSⅡ）反应中心活性参数、PSⅡJ相可变荧光（V_J）、PSⅡ能量分配率、PI、F_v/F₀和F_v/F_m之间的相互关系。【结果】施氮量与温度对叶片含氮量、叶绿素a、叶绿素b、ABS/RC、DI₀/RC、V_J、φE₀、φD₀和产量的影响显著。2年结果表明,随着施氮量的增加,产量均呈先增后降的趋势,且在施氮量为N3（225 kg·hm^-2）时产量最高,2年平均产量分别为9.51t·hm^-2（CK）和8.73 t·hm^-2（HT）。同一氮肥处理下,高温与常温间各指标差异明显,高温处理后叶绿素a、ABS/RC、ET₀/RC、TR₀/RC、PI、F_v/F₀、F_v/F_m和产量均有所降低,说明高温对荧光参数和PSⅡ活性的影响具有负效应。同一温度处理下,随着施氮量增加,春小麦叶绿素含量、ABS/RC、ET₀/RC、PI、F_v/F₀和F_v/F_m等均呈先增后降趋势,DI₀/RC和V_J呈先降后增趋势,且在N3时达到峰值,说明施氮量对叶绿素荧光参数和PSⅡ活性的影响具有补偿效应,适宜的施氮量可以有效增强其活性。温度对ABS/RC、TR₀/RC、ET₀/RC、F_v/F₀和F_v/F_m的影响不显著,而施氮量与温度对ABS/RC、TR₀/RC、ET₀/RC、F_v/F₀和F_v/F_m的交互影响达到显著（P<0.05）或极显著（P<0.01）水平。【结论】施氮量和温度对春小麦荧光特性和产量的影响存在交互作用,其中主导因素为温度,而施氮量对其存在补偿效应。合理的施氮量能有效增加小麦旗叶氮含量、叶绿素含量和PSⅡ反应中心活性,提高植物对光能的捕获、吸收、转化及最大光化学效率,并降低能量耗散率,从而抵御高温胁迫对春小麦光合系统造成的损伤。本试验条件下,选用宁春50号,采用N3（225 kg·hm^-2）的施氮量能有效抵御高温胁迫,并提高春小麦的产量,可为当地春小麦高产稳产提供理论依据和技术支撑。

关键词: 春小麦, 施氮量, 花后高温, 荧光特性, 产量

Abstract:

【Objective】 The aim of this study was to investigate the influence mechanism of nitrogen on the fluorescence characteristics of spring wheat under high temperature, and to clarify the internal mechanism of nitrogen regulation of light reaction center, so as to formulate the nitrogen fertilizer operation measures to alleviate the harm of high temperature.【Method】 The experiment was carried out in 2019 and 2020, using the split-plot trial design. The main zones were 5 nitrogen application, including 0 (N0), 75 kg·hm^-2(N1), 150 kg·hm^-2(N2), 225 kg·hm^-2 (N3), and 300 kg·hm^-2(N4), and the sub-zones were temperatures of 25℃±2℃ (CK) and 35℃±2℃(HT). The interrelationships between nitrogen application and leaf nitrogen content, chlorophyll, PSⅡreaction center activity parameters, PSⅡJ phase variable fluorescence (V_J), PSⅡenergy allocation rate, PI, F_v/F₀, and F_v/F_m at high post-flower temperatures were analyzed. 【Result】 The results showed that the effects of nitrogen application and temperature on leaf nitrogen, chlorophyll a, chlorophyll b, ABS/RC, DI₀/RC, V_J, φE₀ and φD₀and yield were significant. With the increase of nitrogen application, the yield increased first and then decreased, and reached at the highest yield under N3 with normal and high temperature treatments, which was 9.03 t·hm^-2(CK) and 8.37 t·hm^-2(HT). The difference between high temperature and normal temperature was obvious under different temperature treatment with the same nitrogen application, and the chlorophyll a, ABS/RC, ET₀/RC, TR₀/RC, PI, F_v/F₀, F_v/F_m and yield decreased after high temperature treatment, which indicated that the effects of high temperature on fluorescence parameters and PSⅡactivity had negative effect. Under the treatments of different nitrogen application at the same temperature, the chlorophyll content and ABS/RC, ET₀/RC, PI, F_v/F₀, F_v/F_m of spring wheat increased first and then decreased, while the DI₀/RC and V_J decreased first and then increased, and reached the peak at N3(225 kg·hm^-2), indicating that the effects of nitrogen application on chlorophyll fluorescence parameters and PSⅡactivity had compensatory effect, and the appropriate nitrogen application amount could effectively enhance its activity. The effects of temperature on ABS/RC, TR₀/RC, ET₀/RC, F_v/F₀ and F_v/F_m was not significant, but the interaction between nitrogen application and temperature reached significant level (P<0.05) and extremely significant level (P<0.01). 【Conclusion】 To sum up, the effects of nitrogen application and temperature on the fluorescence characteristics and yield of spring wheat was interactive, in which the dominant factor was temperature, while the amount of nitrogen application had a compensatory effect on it. A reasonable amount of nitrogen application could effectively increase the nitrogen content, chlorophyll content and PSⅡreaction center activity of wheat flag leaves, increase the capture, absorption, transformation and maximum photochemical efficiency of light energy by plants, and reduce the energy dissipation rate, so as to resist the damage caused by high temperature stress to the photosynthetic system of spring wheat. According to this test condition, the nitrogen application amount of N3 could effectively resist high temperature stress and increase the yield of spring wheat. The yield of high spring wheat could provide theoretical basis and technical support for high and stable yield of local spring wheat.

Key words: spring wheat, nitrogen application, post-flower high temperature, fluorescence characteristics, yield

坚天才,吴宏亮,康建宏,李鑫,刘根红,陈倬,高娣. 氮素缓解春小麦花后高温早衰的荧光特性研究[J]. 中国农业科学, 2021, 54(15): 3355-3368.

JIAN TianCai,WU HongLiang,KANG JianHong,LI Xin,LIU GenHong,CHEN Zhuo,GAO Di. Fluorescence Characteristics Study of Nitrogen in Alleviating Premature Senescence of Spring Wheat at High Temperature After Anthesis[J]. Scientia Agricultura Sinica, 2021, 54(15): 3355-3368.

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https://www.chinaagrisci.com/CN/Y2021/V54/I15/3355

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年份 Year	测定指标 Indicator for determination	处理 Treatment	处理时间段内各指标的平均值 Average of indicators in the processing period
年份 Year	测定指标 Indicator for determination	处理 Treatment	1 d	2 d	3 d
2019	气温 Temperature (℃)	CK	26.63	27.17	25.29
	气温 Temperature (℃)	HT	35.13	35.28	35.03
	光照强度 Light intensity (LX)	CK	73487.5	73970.8	72680.2
	光照强度 Light intensity (LX)	HT	69309.5	69154.2	68734.4
2020	气温 Temperature (℃)	CK	25.32	25.59	26.33
	气温 Temperature (℃)	HT	36.15	35.92	35.83
	光照强度 Light intensity (LX)	CK	75164.3	79542.4	70158.0
	光照强度 Light intensity (LX)	HT	70535.9	70135.2	67384.3

参数Indicator	说明Illustration
F₀	初始荧光产量Initial fluorescence yield
F_m	最大荧光产量Maximum fluorescence yield
PI	所有荧光参数的综合指标 Synthesis of all fluorescence parameters
F_v/F_m	最大光化学效率Maximum photochemical efficiency
F_v/F₀	捕获光能与热耗散能量的比值 Ratio of captured light energy to heat dissipated energy
V_J	电子经过质体醌A(Q)时的能量耗散比率Energy dissipation ratio of electrons passing through plastid quinone A (Q)
ψ₀=ET₀ /TR₀= 1-V_J	反应中心捕获的激子中用来推动电子传递到电子传递链中超过QA的其他电子受体的激子占用来推动QA还原激子的比率 Rate of excitonic occupancy in excitons captured by reaction centers that push electrons to transfer to electron transport chains that exceed QA of other electron acceptors
φE₀=[1-(F₀/F_m)]×ψ₀	吸收的能量用于电子传递的量子产额Quantum yield of absorbed energy for electron transfer
φD₀=1-φP₀=F₀/F_m	用于热耗散的量子比率 Quantum ratios for heat dissipation
ABS/RC=M₀×(1/V_J) ×(1 /φP₀)	单位反应中心吸收的光能 Light energy absorbed by the unit reaction center
TR₀/RC=M₀×(1/V_J)	单位反应中心捕获的用于还原Q_A的能量 Energy captured by the unit reaction center for reducing QA
ET₀ /RC=M₀×(1/Vj) ×ψ₀	单位反应中心捕获的用于电子传递的能量 Energy captured by a unit reaction center for electron transfer
DI₀ /RC=(ABS/RC)–( TR₀ / RC)	单位反应中心耗散掉的能量 Energy dissipated in the unit reaction center

年份	温度 Temperature	氮肥 Nitrogenous	花后不同时间的荧光参数 Fluorescence parameters at different time after flowering
			V_J		ψ₀		φE₀		φD₀
			25 d	30 d	25 d	30 d	25 d	30 d	25 d	30 d
2019	CK	N0	0.69ab	0.79ab	0.22b	0.10c	0.20b	0.11d	0.33b	0.55a
		N1	0.74ab	0.78ab	0.26b	0.22b	0.24b	0.12d	0.32b	0.43b
		N2	0.79a	0.72bc	0.31ab	0.28ab	0.27ab	0.20b	0.34b	0.30c
		N3	0.63b	0.67c	0.37a	0.33a	0.30a	0.25a	0.20d	0.24d
		N4	0.67b	0.72bc	0.33ab	0.28ab	0.24b	0.19b	0.26c	0.30c
	HT	N0	0.82a	0.87a	0.18b	0.23b	0.09d	0.15c	0.39a	0.37bc
		N1	0.79a	0.85a	0.20b	0.15bc	0.13c	0.17bc	0.37a	0.57a
		N2	0.69ab	0.81a	0.30ab	0.19bc	0.22b	0.13cd	0.26c	0.35bc
		N3	0.67b	0.69bc	0.33ab	0.30a	0.26ab	0.22b	0.22d	0.28cd
		N4	0.68b	0.82a	0.32ab	0.18bc	0.26ab	0.11d	0.26c	0.39bc
2020	CK	N0	0.58bc	0.60d	0.22c	0.20d	0.24c	0.20b	0.20e	0.25d
		N1	0.70a	0.68bc	0.30c	0.32b	0.16e	0.23ab	0.28c	0.29b
		N2	0.58bc	0.64c	0.40ab	0.36a	0.30ab	0.27a	0.32b	0.30c
		N3	0.48c	0.54d	0.42a	0.30b	0.32a	0.25a	0.25d	0.20e
		N4	0.57bc	0.65c	0.43a	0.31b	0.27b	0.26a	0.16f	0.24d
	HT	N0	0.61b	0.84a	0.29c	0.18d	0.28b	0.12c	0.27d	0.28b
		N1	0.68b	0.73b	0.38b	0.21cd	0.19d	0.19b	0.29c	0.30c
		N2	0.62b	0.70b	0.38b	0.30b	0.27b	0.21ab	0.21e	0.35a
		N3	0.48c	0.64c	0.42a	0.36a	0.34a	0.27a	0.39a	0.25d
		N4	0.62b	0.68bc	0.38b	0.26c	0.21c	0.20b	0.26d	0.32a
2019	方差分析 Variance analysis	温度 Temperature (T)	**		**		**		**
		施氮量 Nitrogenous (N)	NS		NS		NS		**
		温度×施氮量 T×N	*		**		**		**
2020	方差分析 Variance analysis	温度 Temperature (T)	**		**		**		**
		施氮量 Nitrogenous (N)	NS		*		NS		**
		温度×施氮量 T×N	*		**		**		**

年份	温度 Temperature	氮肥 Nitrogenous	花后不同时间的荧光参数 Fluorescence parameters at different time after flowering
			PI		F_v/F₀		F_v/F_m		Rubisco (mmol CO₂·min^-1·mL^-1)
			25 d	30 d	25 d	30 d	25 d	30 d	25 d	30 d
2019	CK	N0	1.54d	0.44cd	1.98de	0.92d	0.53b	0.44bc	0.33e	0.23d
		N1	1.92cd	0.62bc	2.18d	1.30cd	0.58b	0.54b	0.51d	0.38c
		N2	2.03cd	0.78bc	2.46cd	2.37b	0.63ab	0.66ab	0.62c	0.49b
		N3	4.91a	1.53a	4.02a	3.10a	0.75a	0.76a	0.87a	0.62a
		N4	3.06b	1.03b	2.78c	2.44b	0.73a	0.68ab	0.79b	0.42c
	HT	N0	0.26f	0.23d	0.98f	0.66e	0.44c	0.31c	0.11f	0.06e
		N1	0.32f	0.18e	1.24e	0.74e	0.74a	0.44bc	0.35e	0.22d
		N2	1.01e	0.21d	2.86c	1.95c	0.78a	0.65ab	0.52d	0.41c
		N3	2.96b	0.96b	3.52b	2.58b	0.79a	0.71a	0.58c	0.50b
		N4	1.86cd	0.28d	3.60b	1.45cd	0.67ab	0.48bc	0.51d	0.33cd
2020	CK	N0	2.15c	0.84d	2.51d	1.51c	0.48c	0.40c	0.39e	0.28e
		N1	2.31b	0.97d	2.80c	1.94b	0.50c	0.46b	0.66c	0.44c
		N2	3.44b	1.21cd	3.03b	1.85b	0.61a	0.54ab	0.68c	0.42c
		N3	5.83a	2.39a	3.50a	2.36a	0.62a	0.58a	1.02a	0.69a
		N4	3.41b	1.50c	3.40a	1.98b	0.56bc	0.59a	0.82b	0.40cd
	HT	N0	0.83e	0.45f	1.59f	0.54f	0.32d	0.15d	0.19f	0.10f
		N1	1.21d	0.88d	2.04e	1.06e	0.46c	0.39c	0.14f	0.09f
		N2	2.06cd	1.33cd	2.53d	1.31d	0.55bc	0.40c	0.43e	0.31e
		N3	3.20bc	1.92b	3.05b	1.83b	0.58a	0.49b	0.50d	0.38d
		N4	2.16c	0.67e	2.63cd	1.59c	0.50b	0.47b	0.62c	0.51b
2019	方差分析 Variance analysis	温度 Temperature (T)	*		NS		NS		**
		施氮量 Nitrogenous (N)	**		**		**		*
		温度×施氮量 T×N	*		*		*		*
2020	方差分析 Variance analysis	温度 Temperature (T)	NS		NS		NS		**
		施氮量 Nitrogenous (N)	**		*		**		*
		温度×施氮量 T×N	*		*		*		*

年份 Year	指标 Indicator	叶片含氮量 Leaf nitrogen	叶绿素a Chlorophyll a	叶绿素b Chlorophyll b	ABS/RC	TR₀/RC	ET₀/RC	DI₀/RC	PI	F_v/F_m	F_v/F₀
2019	产量 Yield	0.683*	0.453*	0.529*	0.836**	0.857**	0.728**	-0.883**	0.608**	0.825**	0.749**
2020	产量 Yield	0.468**	0.552**	0.324*	0.633*	0.415**	0.941*	-0.682**	0.711**	0.693**	0.675**