Scientia Agricultura Sinica ›› 2022, Vol. 55 ›› Issue (17): 3303-3320.doi: 10.3864/j.issn.0578-1752.2022.17.004

• TILLAGE & CULTIVATION·PHYSIOLOGY & BIOCHEMISTRY·AGRICULTURE INFORMATION TECHNOLOGY • Previous Articles     Next Articles

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

RU Chen(),HU XiaoTao(),LÜ MengWei,CHEN DianYu,WANG WenE,SONG TianYuan   

  1. Northwest A&F University/Key Laboratory of Agricultural Soil and Water Engineering in Arid and Semiarid Areas, Ministry of Education, Yangling 712100, Shaanxi
  • Received:2021-11-15 Accepted:2022-01-05 Online:2022-09-01 Published:2022-09-07
  • Contact: XiaoTao HU E-mail:chenru1024@nwafu.edu.cn;huxiaotao11@nwsuaf.edu.cn

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 WUEg and WUEb were significantly improved under N2 supply. Adequate water supply under N2 had 19.09% and 19.44% higher NUEg than drought and combined stress under N3, respectively. This indicates adequate water supply under medium N could effectively alleviate the decrease of NUEg induced by drought and heat stress. The increase of NUEg and NUEb 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

Fig. 1

The extreme maximum temperature after anthesis of winter wheat in the main producing areas in Guanzhong Plain of China from 1990 to 2019"

Fig. 2

Changing of daily temperature and soil relative water content during the stress time H represents high temperature; S represents suitable temperature; HF represents heat treatment; SD represents drought treatment; HD represents combined heat and drought stress treatment; SF represents control"

Fig. 3

Effects of N supply on dry weight of aboveground organs in wheat plants under heat and drought stress (a) and (b) represent 10 d after anthesis, (c) and (d) represent 17 d after anthesis, (e) and (f) represent mature; (a), (c) and (e) represent under a suitable temperature, (b), (d) and (f) represent under a high temperature. The values are the means ± SD (n≥3), the different lowercase letters in the same group indicate significant differences (P<0.05). S represents suitable temperature; H represents high temperature; D represents drought; F represents full watering; N represents nitrogen. The right side of the figure shows the variance analysis results of temperature (T), water (W) and nitrogen (N) on the aboveground dry weight. *, ** and *** indicate significant differences at P<0.05, P<0.01 and P<0.001, respectively; NS means no significance. The same as below"

Table 1

Effects of N supply on aboveground N accumulation and transport of wheat plants under heat, drought and combined stress"

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
(%)
花期
Anthesis
花后10 d
10 DAA
花后17 d
17 DAA
成熟期
Mature
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 * ** **

Fig. 4

Effects of N supply on nitrogen distribution rate of wheat organs under heat, drought and combined stress (a) and (b) represent 10 d after anthesis; (c) and (d) represent 17 d after anthesis; (e) and (f) represent mature"

Table 2

Effects of N supply on the contents of protein components and the activity of N metabolism enzymes in wheat grains under heat, drought and combined stress"

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)
清蛋白
Albumin (%)
球蛋白
Globulin (%)
醇溶蛋白
Gliadin (%)
谷蛋白
Glutenin (%)
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

Fig. 5

Effects of N on activities of N metabolism-related enzymes in wheat grains under heat, drought and combined stress NR represents nitrate reductase; GS represents glutamine synthetase; DAA represents days after anthesis. (a) and (c) represent 13 d after anthesis; (b) and (d) represent 19 d after anthesis. The same as below"

Fig. 6

Relationship of activities of N metabolism-related enzymes with grain N accumulation, N use efficiency for grain and protein yield"

Fig. 7

Effects of N supply on wheat yield, yield components and harvest index under heat, drought and combined stress"

Table 3

Variance analysis of temperature, water and nitrogen on yield formation parameters and water and nitrogen utilization efficiency"

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
WUEg
干物重水分利用效率WUEb 籽粒氮
利用效率
NUEg
干物重氮
利用效率
NUEb
氮肥偏
生产力
PFPN
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

Fig. 8

Effects of nitrogen supply on water and nitrogen utilization efficiency and nitrogen harvest index in wheat under heat, drought and combined stress"

Fig. 9

Principal component analysis of yield formation factors in wheat under heat and drought stress The data points of different colors in the ellipse in the figure represent three nitrogen application levels. NUEb: N utilization efficiency for biomass; NUEg: N utilization efficiency for grain; PY: Protein yield; TGW: 1000 grain weight; ADW: Aboveground dry weight; Y: Grain yield; NHI: Nitrogen harvest index; GNA: Grain nitrogen accumulation; SGN: Grain number per spike; ANA: Aboveground N accumulation; SN: Spikes number per pot"

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