花后高温干旱胁迫下氮素对冬小麦氮积累与代谢酶、蛋白质含量及水氮利用效率的影响

doi:10.3864/j.issn.0578-1752.2022.17.004

摘要/Abstract

摘要：

【目的】本研究基于气候室模拟温度日变化特征,旨在探讨氮素对高温、干旱及复合胁迫下冬小麦地上干物质重、氮积累与分配、氮代谢相关酶活性、蛋白质含量、产量及水氮利用效率的影响。【方法】基于人工气候室开展冬小麦盆栽试验,以小偃22号为试验材料,采用裂-裂区随机完全区组设计,以2个温度处理（高温：H;适宜温度：S）为主区,以2个水分水平（干旱：D;充分供水：F）为裂区,3个施氮水平（低氮：N₁;中氮：N₂;高氮：N₃）为裂-裂区,研究冬小麦生长生理特性、产量及水氮利用效率对高温干旱胁迫及各施氮量的响应特征。【结果】高温、干旱及复合胁迫导致地上总干物质重（ADW）和氮积累量（ANA）降低。在成熟期,高温干旱复合胁迫（HD）和干旱胁迫（SD）下N₃处理ANA分别较N₁处理增加7.26%和6.82%。高温、干旱及复合胁迫提高小麦花前氮素对籽粒贡献率（NRR）,HD胁迫各施氮处理NRR均值较对照（SF）增加达38.21%,施氮量的增加扩大这种增加效应。高温、干旱及复合胁迫导致成熟期穗氮分配率降低,特别是复合胁迫。暴露于高温、干旱及复合胁迫下籽粒蛋白质产量（PY）降低,干旱胁迫（7.37%）各施氮处理PY均值较高温胁迫（3.94%）降低更多,无论单一或复合胁迫下籽粒PY均在N₂处理下显著增加。此外,单一的干旱和高温胁迫下降低的谷氨酰胺合成酶（GS）和硝酸还原酶（NR）活性在N₂处理下显著增加,复合胁迫N₁处理NR和GS活性分别较N₃处理提高23.81%和23.07%。与对照相比,干旱胁迫各施氮处理穗粒数、千粒重和产量均值的降幅均高于高温胁迫,N₂处理对高温和干旱胁迫下这些参数存在明显正向调控,产量水分利用效率（WUE_g）和生物量水分利用效率（WUE_b）在N₂处理下得到明显改善。充分供水+N₂处理籽粒（NUE_g）分别较低干旱和复合胁迫N₃处理提高19.09%和19.44%,表明在水分充足条件下中氮能有效地缓解干旱和高温胁迫下籽粒氮利用效率的降低。NUE_g和NUE_b的提高可能归因于合理氮肥调控下增加的GS和NR活性。主成分分析表明胁迫条件下小麦千粒重和ADW与产量的关系更紧密。【结论】高温和干旱胁迫的综合效应比单一胁迫对小麦危害更大。在单一高温和干旱胁迫下,适量增加氮输入能增加氮代谢酶活性并维持更高氮代谢能力,提高籽粒氮积累量及蛋白质产量,将更有利于提高产量及水氮利用效率。然而在花后遭遇高温干旱复合胁迫时,相比低施氮量,增加施氮对小麦产量形成及水氮的吸收利用均产生一定抑制作用,应适当减少氮肥用量。

关键词: 氮肥管理, 高温和干旱胁迫, 氮积累, 蛋白质含量, 产量, 水氮利用效率

Abstract:

【Objective】 The diurnal variation of temperature was simulated based on the growth chamber, this study aimed to investigate the effects of nitrogen (N) on dry matter accumulation, N accumulation and distribution, activities of N metabolism-related enzymes, protein content, yield, and water and N use efficiency of winter wheat plants under heat, drought and combined stress.【Method】 The experiments were carried out based on growth chambers with Xiaoyan 22 as test material. The experiment consisted of three blocks, in which two temperature treatments (high temperature: H; suitable temperature: S) were assigned as the main plot, two watering treatments (drought: D; sufficient water supply: F) were arranged split-plot, and three N supply levels (low N: N₁; medium N: N₂; high N: N₃) were arranged split-split plot to form a completely randomized block design to investigate the response of the growth and physiological characteristics, yield, and water and N use efficiency in wheat plants to heat, drought stress and different N applications.【Result】 Heat, drought and combined stress resulted in the decrease in ADW (aboveground dry weight) and ANA (aboveground nitrogen accumulation). At maturity, the ANA of N₃ supply under HD and SD was higher 7.26% and 6.82% than that under N₁ supply, respectively. Heat, drought and combined stress resulted in the increase in NRR, and the average NRR of three N supplies under HD increased by 38.21% compared with the control, while increasing N supply further expanded this increasing effect. Heat, drought and combined stress led to decrease in N distribution rate of panicle at maturity, especially combined stress. PY decreased significantly when exposed to heat, drought and combined stress. Compared with the control, the decrease of PY under drought stress conditions (7.37%) was more obvious than that under heat stress conditions (3.94%). Under individual and combined stress treatments, PY was significantly increased under N₂ supply. Furthermore, GS and NR activities decreased under individual heat and drought stress, which were significantly increased under regulation of N₂ supply. The NR and GS activities of N₁ supply under HD were 23.81% and 23.07% higher than that of N₃ supply, respectively. Compared with the control, the reduction in grain number per spike, 1000 grain weight and yield under drought stress conditions was greater than that under heat stress. N₂ supply had an obvious positive effect on these parameters of the two stress treatments, and WUE_g and WUE_b were significantly improved under N₂ supply. Adequate water supply under N₂ had 19.09% and 19.44% higher NUE_g than drought and combined stress under N₃, respectively. This indicates adequate water supply under medium N could effectively alleviate the decrease of NUE_g induced by drought and heat stress. The increase of NUE_g and NUE_b might be attributed to increase of GS and NR activities by appropriate N supply. Principal component analysis indicated that TGW and ADW of wheat were more closely related to yield under stresses conditions.【Conclusion】 The results showed that combined effect of drought and heat stress was more detrimental than individual stresses. Under individual heat and drought stress, an appropriate N supply could increase the activities of N metabolism enzymes and maintain higher N metabolism capacity, improve GNA and PY, and would be much more beneficial to increasing grain yield, water and N use efficiency in wheat production. However, when wheat was subjected to the combined stress after anthesis, compared with low N supply, increasing N supply had a restrictive effect on wheat yield formation as well as water and N utilization capacity, while N supply should be appropriately reduced.

Key words: nitrogen management, heat and drought stress, nitrogen accumulation, protein content, grain yield, water and nitrogen use efficiency

汝晨,胡笑涛,吕梦薇,陈滇豫,王文娥,宋天媛. 花后高温干旱胁迫下氮素对冬小麦氮积累与代谢酶、蛋白质含量及水氮利用效率的影响[J]. 中国农业科学, 2022, 55(17): 3303-3320.

RU Chen,HU XiaoTao,LÜ MengWei,CHEN DianYu,WANG WenE,SONG TianYuan. Effects of Nitrogen on Nitrogen Accumulation and Distribution, Nitrogen Metabolizing Enzymes, Protein Content, and Water and Nitrogen Use Efficiency in Winter Wheat Under Heat and Drought Stress After Anthesis[J]. Scientia Agricultura Sinica, 2022, 55(17): 3303-3320.

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处理 Treatment		地上部氮积累量ANA (mg/plant)				成熟期籽粒氮积累 GNA (mg/plant)	花前贮藏氮素转运量 NRA (mg/plant)	花前氮对籽粒贡献率 NRR (%)	花后氮对籽粒贡献率 NCP (%)
处理 Treatment		花期 Anthesis	花后10 d 10 DAA	花后17 d 17 DAA	成熟期 Mature	成熟期籽粒氮积累 GNA (mg/plant)	花前贮藏氮素转运量 NRA (mg/plant)	花前氮对籽粒贡献率 NRR (%)	花后氮对籽粒贡献率 NCP (%)
高温 H	DN₁	—	32.89±1.81f	37.20±1.18e	39.00±2.10f	27.66±0.45f	17.93±1.01d	64.82±2.67d	35.18±1.23c
	DN₂	—	35.03±2.04c	37.24±0.89de	41.70±2.09e	29.30±1.03d	20.18±0.78c	68.87±1.65b	31.13±1.55e
	DN₃	—	34.01±1.79e	36.70±1.60f	41.83±1.32de	28.54±0.69e	21.98±0.90a	77.01±2.80a	22.99±0.62f
	FN₁	—	34.76±1.32d	37.70±0.59c	44.26±2.78c	32.38±0.60c	17.39±0.39e	53.71±1.20f	46.29±1.84a
	FN₂	—	40.14±0.78a	42.27±1.73a	47.61±0.90a	35.32±1.43b	20.29±1.21bc	57.45±3.74e	42.55±2.47b
	FN₃	—	38.63±1.30b	40.40±2.35b	46.63±1.21b	33.82±0.99a	22.46±0.99a	66.41±1.89c	33.59±1.89d
适温 S	DN₁	—	35.42±1.38f	38.96±1.20f	44.29±0.67f	33.10±1.67e	18.08±0.34d	54.62±0.84d	45.38±0.66cd
	DN₂	—	39.94±0.77d	43.37±0.44d	45.28±1.17e	34.12±2.21d	21.25±0.87b	62.59±1.19b	37.41±1.54e
	DN₃	—	39.10±2.20e	42.49±1.14e	47.31±3.01d	34.66±1.11cd	22.62±1.01a	65.26±2.57a	34.74±2.63f
	FN₁	29.27±1.12c	41.08±3.05c	43.06±2.33c	48.88±1.48c	37.16±0.80b	17.55±0.78e	47.23±1.22f	52.77±1.33a
	FN₂	32.58±0.56b	45.00±1.22a	48.64±1.62a	52.95±2.21b	40.83±2.32a	20.26±0.56c	49.86±1.76e	50.14±3.54b
	FN₃	35.27±1.06a	44.09±2.29b	46.09±1.07b	53.23±1.70a	40.23±1.53a	22.27±1.21a	55.36±1.85cd	44.64±1.77d
变异来源 Source of variation	T	—	***	***	***	***	**	***	***
	W	—	***	**	***	***	*	***	***
	N	***	**	**	**	***	***	***	***
	T×W	—	*	*	NS	NS	*	**	NS
	T×N	—	NS	*	**	NS	NS	*	**
	W×N	—	*	NS	*	*	*	NS	**
	T×W×N	—	*	NS	*	NS	*	**	**

处理 Treatment		蛋白质组分含量 Protein component content				谷/醇比 Glu/Gli	总蛋白质含量 Total protein content (%)	蛋白质产量 Protein yield (mg/grain)
处理 Treatment		清蛋白 Albumin (%)	球蛋白 Globulin (%)	醇溶蛋白 Gliadin (%)	谷蛋白 Glutenin (%)	谷/醇比 Glu/Gli	总蛋白质含量 Total protein content (%)	蛋白质产量 Protein yield (mg/grain)
高温 H	DN₁	1.74±0.02a	1.52±0.02b	3.16±0.12d	4.11±0.17e	1.30±0.02ab	12.41±0.56f	4.27±0.10f
	DN₂	1.75±0.05a	1.60±0.05ab	3.22±0.07cd	4.40±0.21cd	1.37±0.05ab	13.01±0.29a	4.71±0.23d
	DN₃	1.71±0.03b	1.73±0.04a	3.31±0.08bcd	4.27±0.77de	1.29±0.01b	12.87±0.19c	4.50±0.15e
	FN₁	1.58±0.06d	1.20±0.09e	3.42±0.12ab	4.70±0.19b	1.37±0.07ab	12.56±0.44e	4.84±0.25c
	FN₂	1.60±0.02c	1.22±0.01de	3.52±0.09a	4.94±0.09a	1.40±0.05a	12.89±0.72bc	5.09±0.31a
	FN₃	1.71±0.05b	1.31±0.01cde	3.46±0.10ab	4.78±0.23ab	1.38±0.08ab	12.65±0.30de	4.93±0.18bc
适温 S	DN₁	1.53±0.07c	1.19±0.04de	3.52±0.08a	4.87±0.30e	1.38±0.01e	12.21±0.14bc	4.68±0.24de
	DN₂	1.62±0.03a	1.22±0.07c	3.48±0.18a	5.09±0.19cd	1.46±0.04d	12.45±0.88a	4.99±0.30bc
	DN₃	1.57±0.01b	1.18±0.03e	3.60±0.13a	4.95±0.09de	1.38±0.05e	12.55±0.26a	4.66±0.15e
	FN₁	1.44±0.03f	1.22±0.02c	3.01±0.21d	5.42±0.25b	1.80±0.06bc	11.60±0.50e	4.91±0.29c
	FN₂	1.51±0.06d	1.26±0.08a	3.10±0.04dcd	5.74±0.16a	1.85±0.04a	11.88±0.63d	5.25±0.41a
	FN₃	1.47±0.03e	1.24±0.03b	3.20±0.17bc	5.64±0.37a	1.79±0.02bc	12.10±0.66c	5.31±0.27a
变来源异 Source of variation	T	*	**	**	**	***	**	***
	W	*	**	**	**	**	***	**
	N	NS	*	*	***	**	NS	*
	T×W	NS	**	**	**	*	***	***
	T×N	NS	**	NS	NS	**	**	**
	W×N	NS	NS	NS	NS	*	NS	NS
	T×W×N	**	NS	NS	NS	*	NS	NS

变异来源 Source of variation	穗数 Spike number	穗粒数 Grain number per spike	千粒重 1000 grain weight	产量 Yield	收获指数 Harvest index	水分利用效率 WUE_g	干物重水分利用效率WUE_b	籽粒氮利用效率 NUE_g	干物重氮利用效率 NUE_b	氮肥偏生产力 PFP_N	氮收获指数 NHI
T	NS	*	**	***	***	NS	NS	**	**	***	*
W	NS	*	**	***	**	*	**	**	*	***	***
N	***	**	*	**	*	NS	*	*	**	***	**
T×W	NS	NS	*	**	**	**	**	*	*	**	NS
T×N	**	**	*	NS	**	**	*	*	**	**	*
W×N	*	**	NS	NS	NS	**	*	*	NS	***	**
T×W×N	NS	*	NS	NS	**	NS	NS	NS	NS	**	NS