花后高温胁迫对小麦氮素同化利用及产量形成的影响

doi:10.3864/j.issn.0578-1752.2025.19.004

摘要/Abstract

摘要：

【目的】探究花后高温环境下不同耐热型小麦产量的形成过程以及氮素同化能力的变化，为小麦抗逆稳产栽培提供理论支撑。【方法】于2022—2024年，在安徽农业大学国家高新技术农业园开展试验。本试验采用三因素裂区设计，温度作为主区因素，设置高温胁迫（HT）与常温（CK）2个水平；氮肥和品种作为副区因素，其中氮肥设置不施氮（N0）、112.5 kg·hm^-2（N1）和225.0 kg·hm^-2（N2）3个水平；供试材料为6个小麦品种，包括淮麦33（HM33）、龙科1109（LK1109）、安农1589（AN1589）3个耐高温型品种，以及泛麦5号（FM5）、泰农19（TN19）和皖垦1702（WK1702）3个敏感型品种。于小麦花后进行高温胁迫，采用直接称重法测定产量，利用美国LI-6800便携式光合速率测定仪测定光合参数，并通过分光光度法测定氮代谢关键酶活性等指标。【结果】花后高温胁迫对小麦生长发育及生理代谢产生显著抑制效应，导致产量、干物质重量、相对叶绿素含量（SPAD）、叶面积指数（LAI）、净光合速率（P_n）、气孔导度（G_s）、蒸腾速率（T_r）、硝酸还原酶（NR）活性、谷氨酰胺合成酶（GS）活性，以及氮积累量、氮素利用率、氮素吸收效率和氮肥利用率均显著下降。花后高温胁迫下的产量降幅表现出显著的品种耐热性差异与施氮量效应：耐高温品种HM33、LK1109、AN1589在N0、N1和N2处理下的2年平均产量降幅分别为9.71%—6.13%、9.91%—6.24%和11.87%—6.42%，低于敏感型品种FM5、TN19、WK1702的15.26%—10.38%、12.56%—9.84%和12.93%—11.17%。生理机制方面，花后高温胁迫显著降低各施氮处理中小麦品种的P_n，且降幅随施氮量的增加呈现N0>N1>N2的梯度减小趋势，G_s和T_r的变化规律与P_n一致。同时，花后高温胁迫显著降低各施氮处理中小麦旗叶NR和GS活性，而增施氮肥可有效缓解酶活性下降，N2处理下酶活性较高。花后高温胁迫下小麦氮素利用率的降幅也存在品种间差异：耐热型品种HM33的氮素利用率在N2处理下降幅最小，为20.00%；而敏感型品种FM5在N2处理下降幅最大，为31.55%。【结论】花后高温胁迫显著降低小麦产量，增施氮肥（112.5、225.0 kg·hm^-2）可有效缓解此效应。耐热品种通过维持氮代谢酶活性、优化氮同化与转运，减轻小麦产量损失。适量增氮（225.0 kg·hm^-2）显著提升小麦氮素利用效率与籽粒同化物积累。表明协同利用品种耐热性与氮肥管理可增强小麦高温抗性，为小麦稳产栽培提供理论依据。

关键词: 小麦, 高温胁迫, 施氮量, 氮素利用率, 产量

Abstract:

【Objective】This study aimed to investigate the impact of post-anthesis heat stress on yield formation and nitrogen assimilation capacity in wheat genotypes differing in heat tolerance, so as to provide a theoretical foundation for resilient and stable-yielding cultivation strategies.【Method】Field experiments were conducted from 2022 to 2024 at the National High-Tech Agricultural Park of Anhui Agricultural University using a split-plot design. Temperature was set as the main factor with two levels, including high temperature stress (HT) and ambient control (CK), while three nitrogen rates (no nitrogen (N0), 112.5 kg·hm^-2 (N1), and 225.0 kg·hm^-2 (N2)) and six wheat cultivars (three heat-tolerant: HM33, LK1109, AN1589; three heat-sensitive: FM5, TN19, WK1702) were arranged as subplot factors. High temperature stress was applied after anthesis. Grain yield, photosynthetic parameters (LI-6800), and activities of key nitrogen metabolism enzymes (spectrophotometry) were determined.【Result】High-temperature stress after anthesis significantly inhibited wheat growth, development, and physiological metabolism, leading to a marked decrease in yield, dry matter weight, relative chlorophyll content (SPAD), leaf area index (LAI), net photosynthetic rate (P_n), stomatal conductance (G_s), transpiration rate (T_r), nitrate reductase (NR) activity, and glutamine synthetase (GS) activity, as well as nitrogen accumulation, nitrogen use efficiency, nitrogen absorption efficiency, and nitrogen fertilizer use efficiency. The yield reduction due to high-temperature stress exhibited significant varietal heat tolerance differences and nitrogen application effects: the two-year average yield reduction of heat-tolerant varieties of HM33, LK1109, and AN1589 under N0, N1, and N2 treatments were 9.71%-6.13%, 9.91%-6.24%, and 11.87%-6.42%, respectively, lower than the reductions in sensitive varieties of FM5, TN19, and WK1702, which were 15.26%-10.38%, 12.56%-9.84%, and 12.93%-11.17%, respectively. In terms of physiological mechanisms, high-temperature stress after anthesis significantly reduced P_n across all nitrogen treatments, with the reduction showing a decreasing gradient as nitrogen levels increased: N0>N1>N2. The changes in G_s and T_r followed the same pattern as P_n. Furthermore, high-temperature stress significantly reduced NR and GS activity in the flag leaves across all nitrogen treatments, but increased nitrogen fertilization effectively and alleviated the decline in enzyme activity, with the highest enzyme activity observed under the N2 treatment. The reduction in nitrogen use efficiency under high-temperature stress also showed varietal differences: the nitrogen use efficiency of the heat-tolerant variety HM33 decreased the least under the N2 treatment (by 20.00%), while the sensitive variety FM5 exhibited the greatest decrease (31.55%) under the same treatment.【Conclusion】Post-flowering high temperature stress significantly reduced wheat yield, while the application of nitrogen fertilizers (112.5 and 225.0 kg·hm^-2) effectively mitigated this decline. Heat-tolerant varieties minimized yield losses by maintaining the activity of key nitrogen-metabolizing enzymes and optimizing nitrogen assimilation and transport. Increased nitrogen application, particularly at the 225.0 kg·hm^-2, significantly enhanced nitrogen use efficiency and promoted the accumulation of assimilates in grains. These results demonstrated that combining heat-tolerant genotypes with appropriate nitrogen management could improve thermotolerance in wheat, which provided a theoretical foundation for cultivating high and stable yields under heat stress.

Key words: wheat, high temperature stress, nitrogen application rate, nitrogen use efficiency, yield

雷毕欣, 余勇波, 张明通, 崔国际, 洪嘉雯, 胡涛, 犹艾欣, 张文静, 马尚宇, 黄正来, 樊永惠. 花后高温胁迫对小麦氮素同化利用及产量形成的影响[J]. 中国农业科学, 2025, 58(19): 3837-3856.

LEI BiXin, YU YongBo, ZHANG MingTong, CUI GuoJi, HONG JiaWen, HU Tao, YOU AiXin, ZHANG WenJing, MA ShangYu, HUANG ZhengLai, FAN YongHui. Impact of Post-Anthesis Heat Stress on Nitrogen Use Efficiency and Yield Components in Wheat[J]. Scientia Agricultura Sinica, 2025, 58(19): 3837-3856.

图/表 11

表1

图1

表2

花后高温胁迫对小麦产量及产量构成因素的影响"

年份 Year	品种 Cultivar	处理 Treatment	穗数 Spike number (×10⁴·hm^-2)	穗粒数 Grain number	千粒重 1000-grain weight (g)	产量 Yield (kg·hm^-2)
2022—2023	淮麦33 HM33	CKN0	412.67±0.33b	35.00±0.58c	41.57±0.29b	6002.69±61.26e
		CKN1	426.00±2.89a	39.33±0.88b	43.78±0.60a	7332.13±97.57c
		CKN2	433.67±4.41a	44.00±0.58a	44.17±0.61a	8423.64±36.59a
		HTN0	409.33±0.33b	35.00±0.58c	39.00±0.48c	5587.17±115.31e
		HTN1	432.67±2.19a	39.33±0.33b	40.22±0.05bc	6843.79±59.90d
		HTN2	435.67±2.33a	43.33±0.33a	42.26±0.34ab	7977.91±100.50b
	龙科1109 LK1109	CKN0	458.33±4.41b	33.67±0.33c	34.78±0.19c	5365.67±38.42e
		CKN1	520.00±1.15a	37.33±0.33b	37.60±0.25b	7300.37±98.38c
		CKN2	525.67±1.20a	40.00±0.58ab	39.23±0.28a	8247.23±73.81a
		HTN0	463.67±4.10b	33.00±0.58c	32.57±0.25d	4983.97±116.14f
		HTN1	518.33±2.03a	37.67±0.58ab	34.40±0.08c	6715.17±22.63d
		HTN2	521.33±3.18a	41.00±0.58a	36.68±0.14b	7839.32±52.21b
	安农1589 AN1589	CKN0	412.00±2.00c	32.67±0.33c	40.41±0.27b	5438.75±60.75e
		CKN1	425.33±3.18b	35.67±0.33b	42.49±0.24a	6444.22±33.30c
		CKN2	504.33±2.33a	38.67±0.67a	42.89±0.38a	8361.77±81.84a
		HTN0	411.67±1.45c	32.33±0.33c	36.79±0.16d	4896.53±18.61f
		HTN1	433.33±1.67b	35.33±0.33b	38.72±0.23c	5928.12±60.76d
		HTN2	501.00±2.31a	38.33±0.33a	40.75±0.38b	7824.58±37.19b
	泛麦5号 FM5	CKN0	400.00±2.08c	35.00±0.58b	38.53±0.19a	5394.27±94.91e
		CKN1	431.67±3.33b	39.00±0.58a	39.54±0.09a	6654.90±74.84c
		CKN2	482.33±2.67a	40.67±0.33a	39.94±0.42a	7832.87±40.12a
		HTN0	393.33±1.67c	33.67±0.88b	34.47±0.88b	4558.39±32.29f
		HTN1	430.67±0.67b	39.00±0.58a	34.56±0.36b	5804.81±89.32d
		HTN2	482.67±3.93a	40.33±0.67a	35.72±0.28b	6950.52±45.39b
	泰农19 TN19	CKN0	416.67±4.41c	34.67±0.88c	37.46±0.66b	5405.86±15.07d
		CKN1	430.00±1.15bc	41.33±0.33b	38.40±0.32b	6823.33±16.57b
		CKN2	448.33±1.67a	43.67±0.33a	40.31±0.38a	7890.45±76.93a
		HTN0	423.33±4.41c	33.33±0.67c	34.04±0.13c	4802.33±91.17e
		HTN1	428.00±1.53bc	41.00±0.00b	34.60±0.05c	6071.70±17.69c
		HTN2	442.33±5.36ab	43.00±0.58ab	37.17±0.07b	7071.21±133.9b
	皖垦1702 WK1702	CKN0	417.00±1.00b	35.00±0.58b	38.75±0.26ab	5654.77±44.27e
		CKN1	461.33±1.76a	39.67±0.33a	39.65±0.09ab	7255.52±49.71b
		CKN2	465.00±2.65a	40.67±0.88a	40.87±1.04a	7720.82±44.12a
		HTN0	413.33±3.28b	34.33±0.33b	35.14±0.06d	4986.85±89.52f
		HTN1	455.33±5.93a	38.67±0.33a	36.14±0.29cd	6363.77±133.80d
		HTN2	455.67±7.54a	39.33±0.67a	38.01±0.17bc	6810.76±137.57c
2023—2024	淮麦33 HM33	CKN0	403.00±0.58c	35.00±0.58c	40.23±0.66c	5671.36±10.82e
		CKN1	421.33±1.86b	38.00±0.58b	43.16±0.83ab	6906.86±54.24c
		CKN2	448.67±1.76a	44.00±0.58a	44.29±0.64a	8747.42±253.63a
		HTN0	400.00±2.31c	34.67±0.88c	35.77±0.59d	4962.18±190.42f
		HTN1	418.33±1.20b	37.33±0.33b	39.81±0.61c	6216.14±67.09d
		HTN2	448.00±1.15a	43.00±0.58a	42.24±0.27b	8138.20±169.86b
	龙科1109 LK1109	CKN0	433.00±0.58c	33.33±0.33b	39.19±0.65c	5655.99±31.03e
		CKN1	496.33±1.33b	35.33±0.33b	41.03±0.74b	7196.21±126.63c
		CKN2	519.00±2.31a	40.00±0.58a	42.97±0.74a	8923.33±210.22a
		HTN0	431.67±1.45c	33.00±0.58b	34.67±1.01d	4937.71±96.83f
		HTN1	495.00±2.31b	34.33±0.67b	38.06±0.89c	6469.13±173.86d
		HTN2	517.67±1.33a	39.00±0.58a	40.88±0.81b	8251.40±58.26b
	安农1589 AN1589	CKN0	410.33±0.88c	33.00±0.58bc	39.85±0.57c	5397.43±139.56d
		CKN1	424.07±2.31b	35.00±0.58b	41.38±0.39b	6142.98±137.91c
		CKN2	445.56±3.53a	39.00±0.58a	43.66±0.48a	7585.08±81.60a
		HTN0	404.80±2.91c	31.33±0.33c	36.70±0.45d	4654.63±85.43e
		HTN1	421.14±1.00b	34.00±1.53b	39.29±0.44c	5629.55±297.32d
		HTN2	443.81±2.03a	38.00±0.58a	42.10±0.40b	7098.49±39.04b
	泛麦5号 FM5	CKN0	402.87±1.33c	34.00±0.58b	38.54±0.39b	5279.13±107.51d
		CKN1	423.33±0.33b	39.67±1.86a	39.62±0.18ab	6652.94±306.09b
		CKN2	446.89±2.67a	41.00±1.15a	40.53±0.26a	7425.04±199.81a
		HTN0	401.53±2.67c	33.00±1.15b	33.90±0.61d	4485.78±79.58e
		HTN1	422.88±1.33b	39.33±0.88a	35.55±0.45c	5909.63±112.43c
		HTN2	445.81±2.19a	40.00±1.53a	37.70±0.11c	6719.96±230.76b
	泰农19 TN19	CKN0	414.87±1.33c	33.00±0.58c	38.39±0.65bc	5255.66±124.53d
		CKN1	421.54±1.33b	43.67±0.88ab	39.53±0.39ab	7276.08±143.18b
		CKN2	434.88±1.33a	44.67±0.88a	40.03±0.20a	7776.84±169.99a
		HTN0	413.54±1.33c	32.00±0.58c	34.19±0.45d	4522.50±59.51e
		HTN1	420.21±2.31b	42.00±1.00b	37.08±0.54c	6544.64±182.77c
		HTN2	432.22±2.31a	43.67±0.67ab	37.38±0.65c	7054.41±156.49b
	皖垦1702 WK1702	CKN0	420.21±2.31c	36.00±0.58b	37.54±0.49c	5676.01±35.50d
		CKN2	432.22±2.31b	40.00±0.58a	40.17±0.38b	6943.89±71.83b
		CKN1	446.89±3.53a	41.67±0.33a	43.95±0.68a	8180.68±2.93a
		HTN0	418.88±1.33c	35.67±0.33b	32.65±0.66e	4878.76±128.01e
		HTN1	429.55±1.33b	39.67±0.88a	35.79±0.64d	6101.30±225.0.84c
		HTN2	445.56±2.67a	41.00±0.58a	40.05±0.37b	7317.06±166.20b
2022—2023	F值 F-value	温度 Temperature	819.10**	45.99**	4729.13**	1076.15**
		氮肥 Nitrogen	232.02**	219.46**	164.83**	4.76**
		温度×氮肥Temperature×Nitrogen	345.65**	3.15**	123.94**	40.19**
2023—2024		温度 Temperature	3353.76**	585.14**	76.84*	60.78*
		氮肥 Nitrogen	1049.74**	5.66*	9.40**	12.17**
		温度×氮肥 Temperature×Nitrogen	1476.43**	28.24**	8.37*	34.09**

表2

图2

图3

图4

图5

图6

图7

表3

花后高温胁迫对小麦氮素利用率的影响"

品种 Cultivar	处理 Treatment	氮积累量 Nitrogen accumulation (kg·hm^-2)	氮素利用率 Nitrogen use efficiency (kg·kg^-1)	氮素吸收效率 Nitrogen uptake efficiency (%)	氮肥利用率 Nitrogen recovery efficiency (%)
淮麦33 HM33	CKN0	103.67±2.17e	—	—	—
	CKN1	153.79±1.63c	15.80±0.76b	27.20±0.48c	35.71±1.16a
	CKN2	188.57±2.06a	19.31±1.00a	37.15±0.54a	33.58±0.18b
	HTN0	92.59±2.72f	—	—	—
	HTN1	127.61±3.96d	12.11±0.76c	22.57±0.74d	24.95±1.02c
	HTN2	165.98±5.81b	15.45±0.30b	32.69±1.14b	29.03±1.75d
龙科 1109 LK1109	CKN0	104.81±4.73e	—	—	—
	CKN1	142.88±7.30c	17.94±0.33b	25.27±1.20c	27.13±2.75a
	CKN2	188.93±4.30a	21.26±1.02a	37.21±0.75a	33.27±0.52b
	HTN0	95.65±2.58f	—	—	—
	HTN1	132.35±4.82d	12.74±1.10d	20.24±3.67d	26.15±2.30b
	HTN2	166.36±2.42b	15.98±0.76c	32.77±0.41b	27.97±0.28b
安农1589 AN1589	CKN0	169.34±15.72cd	—	—	—
	CKN1	214.41±15.75b	14.01±0.53b	37.91±2.53b	32.12±0.98a
	CKN2	253.52±12.26a	20.76±0.75a	49.93±2.24a	33.29±1.38a
	HTN0	118.14±8.84e	—	—	—
	HTN1	154.87±8.46d	10.14±0.15c	31.05±1.61c	26.17±0.31b
	HTN2	186.94±9.36c	15.42±0.73b	42.71±3.68b	27.21±0.37b
泛麦5号FM5	CKN0	156.26±7.32d	—	—	—
	CKN1	204.04±5.08b	16.75±1.21b	36.09±0.64b	34.04±1.81a
	CKN2	240.70±11.14a	21.57±1.17a	47.41±2.01a	33.39±1.53a
	HTN0	104.50±7.99e	—	—	—
	HTN1	141.99±8.45d	10.41±0.28d	25.11±1.38c	26.72±0.43b
	HTN2	178.10±9.86c	13.90±0.51c	35.08±1.84b	29.11±0.94b
泰农19 TN19	CKN0	126.34±7.60c	—	—	—
	CKN1	172.85±5.12b	15.03±0.35b	30.57±0.82c	33.14±2.24a
	CKN2	209.33±8.44a	17.75±0.57a	41.23±1.67a	32.82±0.65a
	HTN0	94.06±4.95d	—	—	—
	HTN1	130.23±5.09c	10.12±0.17d	23.03±0.76d	25.77±1.54c
	HTN2	167.72±6.25b	12.14±0.53c	33.04±1.17b	29.13±0.60b
皖垦1702 WK1702	CKN0	137.08±6.85d	—	—	—
	CKN1	183.25±5.32b	17.22±0.75b	32.41±0.96b	32.90±1.21a
	CKN2	219.42±5.32a	21.83±0.90a	43.22±1.06a	32.57±0.72a
	HTN0	89.40±3.80f	—	—	—
	HTN1	124.69±6.70e	11.17±1.76d	22.88±0.47c	25.15±2.56c
	HTN2	162.74±0.53c	14.94±1.17c	32.06±0.20b	29.01±1.66b
F-value	温度 Temperature	1.64	127.18**	0.78	7.86
	氮肥 Nitrogen	30.80**	116.16**	0.58	203.04**
	温度×氮肥Temperature×Nitrogen	55.93**	6.18	162.60**	4.90

表3

图8

参考文献 48

[1]	张玉. 浅析小麦种植技术推广意义及有效途径. 种子科技, 2018, 36(5): 25.
	ZHANG Y. Analysis on the significance and effective ways of wheat planting technology popularization. Seed Science & Technology, 2018, 36(5): 25. (in Chinese)
[2]	CHEN Z L, GALLI M, GALLAVOTTI A. Mechanisms of temperature-regulated growth and thermotolerance in crop species. Current Opinion in Plant Biology, 2022, 65: 102134.
[3]	曹彩云, 党红凯, 郑春莲, 李科江, 马俊永. 阶段高温对不同小麦品种产量的影响及其耐热性差异研究. 麦类作物学报, 2020, 40(3): 351-364.
	CAO C Y, DANG H K, ZHENG C L, LI K J, MA J Y. Impacts of high temperature in different periods on yield and evaluation of heat tolerance in wheat varieties. Journal of Triticeae Crops, 2020, 40(3): 351-364. (in Chinese)
[4]	石晓丽, 史文娇. 极端高温对黄淮海平原冬小麦产量的影响. 生态与农村环境学报, 2016, 32(2): 259-269.
	SHI X L, SHI W J. Impacts of extreme high temperature on winter wheat yield in the Huang-Huai-Hai Plain. Journal of Ecology and Rural Environment, 2016, 32(2): 259-269. (in Chinese)
[5]	李世平. 冬小麦耐热性及相关性状数量位点遗传剖析[D]. 杨凌: 西北农林科技大学, 2013.
	LI S P. Genetic dissection of quantitative traits loci for heat tolerance index and traits associated with heat tolerance in winter wheat (Triticum aestivum L.)[D]. Yangling: Northwest A & F University, 2013. (in Chinese)
[6]	FAROOQ M, BRAMLEY H, PALTA J A, SIDDIQUE K H M. Heat stress in wheat during reproductive and grain-filling phases. Critical Reviews in Plant Sciences, 2011, 30(6): 491-507.
[7]	滕中华, 智丽, 宗学凤, 王三根, 何光华. 高温胁迫对水稻灌浆结实期叶绿素荧光、抗活性氧活力和稻米品质的影响. 作物学报, 2008, 34(9): 1662-1666.
	TENG Z H, ZHI L, ZONG X F, WANG S G, HE G H. Effects of high temperature on chlorophyll fluorescence, active oxygen resistance activity, and grain quality in grain-filling periods in rice plants. Acta Agronomica Sinica, 2008, 34(9): 1662-1666. (in Chinese)
[8]	王娇, 白海霞, 韩语燕, 梁惠, 冯雅楠, 张东升, 李萍, 宗毓铮, 史鑫蕊, 郝兴宇. CO₂浓度升高、升温及其交互作用对良星99冬小麦叶片碳氮代谢的影响. 作物学报, 2025, 51(4): 1061-1076. doi: 10.3724/SP.J.1006.2025.41054
	WANG J, BAI H X, HAN Y Y, LIANG H, FENG Y N, ZHANG D S, LI P, ZONG Y Z, SHI X R, HAO X Y. Effects of elevated CO₂ concentration, increased temperature and their interaction on the carbon and nitrogen metabolism in Liangxing 99 winter wheat leaves. Acta Agronomica Sinica, 2025, 51(4): 1061-1076. (in Chinese)
[9]	TISCHNER R. Nitrate uptake and reduction in higher and lower plants. Plant, Cell & Environment, 2000, 23(10): 1005-1024.
[10]	LIANG C G, CHEN L P, WANG Y, LIU J, XU G L, LI T. High temperature at grain-filling stage affects nitrogen metabolism enzyme activities in grains and grain nutritional quality in rice. Rice Science, 2011, 18(3): 210-216.
[11]	冯万军, 邢国芳, 牛旭龙, 窦晨, 韩渊怀. 植物谷氨酰胺合成酶研究进展及其应用前景. 生物工程学报, 2015, 31(9): 1301-1312.
	FENG W J, XING G F, NIU X L, DOU C, HAN Y H. Progress and application prospects of glutamine synthase in plants. Chinese Journal of Biotechnology, 2015, 31(9): 1301-1312. (in Chinese)
[12]	朱保磊, 陈金平, 李刚, 周国勤, 赵万兵, 赵永涛, 陈杰, 陈建辉, 石守设, 詹克慧. 小麦不溶性谷蛋白含量与HMW-GS及品质的关系. 麦类作物学报, 2021, 41(9): 1099-1104.
	ZHU B L, CHEN J P, LI G, ZHOU G Q, ZHAO W B, ZHAO Y T, CHEN J, CHEN J H, SHI S S, ZHAN K H. Relationship between insoluble glutenin content and HMW-GS and quality in wheat. Journal of Triticeae Crops, 2021, 41(9): 1099-1104. (in Chinese)
[13]	LENKA N K, LENKA S, YASHONA D S, SHUKLA A K, ELANCHEZHIAN R, DEY P, PATRA A K. Carbon dioxide and/or temperature elevation effect on yield response, nutrient partitioning and use efficiency of applied nitrogen in wheat crop in central India. Field Crops Research, 2021, 264: 108084.
[14]	MANDIC V, KRNJAJA V, TOMIC Z, BIJELIC Z, SIMIC A, RUZIC MUSLIC D, GOGIC M. Nitrogen fertilizer influence on wheat yield and use efficiency under different environmental conditions. Chilean Journal of Agricultural Research, 2015, 75(1): 92-97.
[15]	MA T, CHEN K W, HE P R, DAI Y, YIN Y Q, PENG S H, DING J H, YU S E, HUANG J S. Sunflower photosynthetic characteristics, nitrogen uptake, and nitrogen use efficiency under different soil salinity and nitrogen applications. Water, 2022, 14(6): 982.
[16]	胡晨曦. 夜间增温对土壤-小麦系统氮素平衡及利用的影响[D]. 南京: 南京农业大学, 2019.
	HU C X. Effects of nighttime warming on nitrogen balance and utilization in the soil-wheat system[D]. Nanjing: Nanjing Agricultural University, 2019. (in Chinese)
[17]	王潭刚, 孙婷, 王冀川, 李慧琴, 高振, 石元强. 播期和密度对滴灌冬小麦群体结构与抗倒特性的影响. 浙江农业学报, 2021, 33(2): 193-202. doi: 10.3969/j.issn.1004-1524.2021.02.02
	WANG T G, SUN T, WANG J C, LI H Q, GAO Z, SHI Y Q. Effects of sowing date and density on population structure and lodging resistance of winter wheat under drip irrigation. Acta Agriculturae Zhejiangensis, 2021, 33(2): 193-202. (in Chinese) doi: 10.3969/j.issn.1004-1524.2021.02.02
[18]	初金鹏, 朱文美, 尹立俊, 石玉华, 邓淑珍, 张良, 贺明荣, 代兴龙. 宽幅播种对冬小麦‘泰农18’产量和氮素利用率的影响. 应用生态学报, 2018, 29(8): 2517-2524. doi: 10.13287/j.1001-9332.201808.027
	CHU J P, ZHU W M, YIN L J, SHI Y H, DENG S Z, ZHANG L, HE M R, DAI X L. Effects of wide-range planting on the yield and nitrogen use efficiency of winter wheat cultivar Tainong 18. Chinese Journal of Applied Ecology, 2018, 29(8): 2517-2524. (in Chinese)
[19]	邹琦. 植物生理学实验指导. 北京: 中国农业出版社, 2000.
	ZOU Q. Experimental Instruction of Plant Physiology. Beijing: China Agriculture Press, 2000. (in Chinese)
[20]	李泽松, 林清华, 张楚富, 彭进, 魏国威. 不同氮源对水稻幼苗根氨同化酶的影响. 武汉大学学报(自然科学版), 2000, 46(6): 729-732.
	LI Z S, LIN Q H, ZHANG C F, PENG J, WEI G W. Effect of different nitrogen sources on ammonia-assimilating enzymes of the roots in rice seedlings. Journal of Wuhan University (Natural Science Edition), 2000, 46(6): 729-732. (in Chinese)
[21]	樊永惠, 李宇星, 马亮亮, 吕钊彦, 武倩倩, 张文静, 马尚宇, 黄正来. 灌浆期高温胁迫下外源水杨酸对小麦旗叶抗氧化生理特性的影响. 核农学报, 2022, 36(9): 1878-1886. doi: 10.11869/j.issn.100-8551.2022.09.1878
	FAN Y H, LI Y X, MA L L, LÜ Z Y, WU Q Q, ZHANG W J, MA S Y, HUANG Z L. Effects of exogenous salicylic acid on antioxidant physiological characteristics of wheat flag leaves under high temperature stress at grain filling stage. Journal of Nuclear Agricultural Sciences, 2022, 36(9): 1878-1886. (in Chinese) doi: 10.11869/j.issn.100-8551.2022.09.1878
[22]	封超年, 郭文善, 施劲松, 彭永欣, 朱新开. 小麦花后高温对籽粒胚乳细胞发育及粒重的影响. 作物学报, 2000, 26(4): 399-405.
	FENG C N, GUO W S, SHI J S, PENG Y X, ZHU X K. Effect of high temperature after anthesis on endosperm cell development and grain weight in wheat. Acta Agronomica Sinica, 2000, 26(4): 399-405. (in Chinese)
[23]	VERMA L, SINGH A K, SINGH S, TIWARI D, ZAIDI S T, YADAV R K, MISHRA S R, SINGH A K. Temperature stress its impact on yield of various wheat varieties at different growth stages. National Academy Science Letters, 2024, 47(3): 219-225.
[24]	岳鹏莉, 王晨阳, 卢红芳, 刘卫星, 马耕, 王强, 胡阳阳. 灌浆期高温胁迫对小麦灌浆的影响及叶面喷剂的缓解作用. 中国生态农业学报, 2016, 24(8): 1103-1113.
	YUE P L, WANG C Y, LU H F, LIU W X, MA G, WANG Q, HU Y Y. Impact of high temperature stress on grain-filling and the relief effect of foliage sprays during grain-filling stage of wheat. Chinese Journal of Eco-Agriculture, 2016, 24(8): 1103-1113. (in Chinese)
[25]	罗宏海, 李俊华, 张宏芝, 何在菊, 勾玲, 张旺锋. 源库调节对新疆高产棉花产量形成期光合产物生产与分配的影响. 棉花学报, 2009, 21(5): 371-377. doi: 10.11963/cs090507
	LUO H H, LI J H, ZHANG H Z, HE Z J, GOU L, ZHANG W F. Effects of source and sink manipulation on transportation and allocation of leaf photosynthetic products during flowering and boll-setting stage in high-yield cotton of Xinjiang. Cotton Science, 2009, 21(5): 371-377. (in Chinese)
[26]	COSSANI C M, REYNOLDS M P. Physiological traits for improving heat tolerance in wheat. Plant Physiology, 2012, 160(4): 1710-1718. doi: 10.1104/pp.112.207753 pmid: 23054564
[27]	BALA S, ASTHIR B, TAGGAR M S. Grain carbon metabolism and stem reserve mobilization compensate high temperature stress in wheat. Agrochimica: International Journal of Plant Chemistry, Soil Science and Plant Nutrition of the University of Pisa, 2021, 65(1): 39-52.
[28]	SOLTANI A, WERADUWAGE S M, SHARKEY T D, LOWRY D B. Elevated temperatures cause loss of seed set in common bean (Phaseolus vulgaris L.) potentially through the disruption of source- sink relationships. BMC Genomics, 2019, 20(1): 312.
[29]	王磊. 硫化氢信号对干旱胁迫下十字花科植物光合作用的调节[D]. 太原: 山西大学, 2020.
	WANG L. Regulation of hydrogen sulfide signal on photosynthesis of cruciferous plants under drought stress[D]. Taiyuan: Shanxi University, 2020. (in Chinese)
[30]	江珊, 吴龙英, 赵宝生, 黄佳惠, 蒋宇喆, 焦元, 黄进. 植物耐受高温胁迫的分子机制研究进展. 中国农学通报. 2024, 40(9): 132-138. doi: 10.11924/j.issn.1000-6850.casb2023-0544
	JIANG S, WU L Y, ZHAO B S, HUANG J H, JIANG Y Z, JIAO Y, HUANG J. Molecular mechanism of heat stress tolerance in plants: A review. Chinese Agricultural Science Bulletin, 2024, 40(9): 132-138. (in Chinese) doi: 10.11924/j.issn.1000-6850.casb2023-0544
[31]	LI R Q, HE Y, CHEN J Y, ZHENG S Y, ZHUANG C X. Research progress in improving photosynthetic efficiency. International Journal of Molecular Sciences, 2023, 24(11): 9286.
[32]	卞晓波, 陈丹丹, 王强盛, 王绍华. 花后开放式增温对小麦产量及品质的影响. 中国农业科学, 2012, 45(8): 1489-1498. doi:10.3864/j.issn.0578-1752.2012.08.004.
	BIAN X B, CHEN D D, WANG Q S, WANG S H. Effects of different day and night temperature enhancements on wheat grain yield and quality after anthesis under free air controlled condition. Scientia Agricultura Sinica, 2012, 45(8): 1489-1498. doi:10.3864/j.issn.0578-1752.2012.08.004. (in Chinese)
[33]	余梦奇, 路梦莉, 张雅婷, 陈志英, 李文阳. 灌浆期高温对玉米叶片光合特性及抗氧化酶活性的影响. 中国农业气象, 2023, 44 (7): 599-610.
	YU M Q, LU M L, ZHANG Y T, CHEN Z Y, LI W Y. Effects of high temperature on photosynthetic characteristics and antioxidant enzyme activities of maize leaves during filling stage. Chinese Journal of Agrometeorology, 2023, 44(7): 599-610. (in Chinese)
[34]	姜磊, 李焕勇, 张芹, 张会龙, 乔艳辉, 张华新, 杨秀艳. AM真菌对盐碱胁迫下杜梨幼苗生长与生理代谢的影响. 南京林业大学学报:自然科学版, 2020, 44(6): 152-160.
	JIANG L, LI H Y, ZHANG Q, ZHANG H L, QIAO Y H, ZHANG H X, YANG X Y. Effects of arbuscular mycorrhiza fungi on the growth and physiological metabolism of Pyrus betulaefolia Bunge seedlings under saline-alkaline stress. Journal of Nanjing Forestry University: Natural Sciences Edition, 2020, 44(6): 152-160. (in Chinese)
[35]	付景, 孙宁宁, 刘天学, 马俊峰, 杨豫龙, 赵霞, 穆心愿, 李潮海. 穗期高温对玉米子粒灌浆生理及产量的影响. 作物杂志, 2019(3): 118-125.
	FU J, SUN N N, LIU T X, MA J F, YANG Y L, ZHAO X, MU X Y, LI C H. The effects of high temperature at spike stage on grain- filling physiology and yield of maize. Crops, 2019(3): 118-125. (in Chinese)
[36]	赵晶晶. 花后高温胁迫下不同施氮量对春小麦产量形成的影响机理[D]. 银川: 宁夏大学, 2015.
	ZHAO J J. The mechanism of the effect of different nitrogen application rates on spring wheat yield formation under post-anthesis high temperature stress[D]. Yinchuan: Ningxia University, 2015. (in Chinese)
[37]	FENG B, LIU P, LI G, DONG S T, WANG F H, KONG L A, ZHANG J W. Effect of heat stress on the photosynthetic characteristics in flag leaves at the grain-filling stage of different heat‐resistant winter wheat varieties. Journal of Agronomy and Crop Science, 2014, 200(2): 143-155.
[38]	GAO C H, SUN M, ANWAR S, FENG B, REN A X, LIN W, GAO Z Q. Response of physiological characteristics and grain yield of winter wheat varieties to long-term heat stress at anthesis. Photosynthetica, 2021, 59(4): 640-651.
[39]	蒋志敏, 王威, 储成才. 植物氮高效利用研究进展和展望. 生命科学, 2018, 30(10): 1060-1071.
	JIANG Z M, WANG W, CHU C C. Towards understanding of nitrogen use efficiency in plants. Chinese Bulletin of Life Sciences, 2018, 30(10): 1060-1071. (in Chinese)
[40]	刘颖, 顾昀怿, 张伟杨, 杨建昌. 水分与氮素及其互作调控小麦产量和水氮利用效率研究进展. 作物杂志, 2023(4): 7-15.
	LIU Y, GU Y Y, ZHANG W Y, YANG J C. Research advances in the effects of water and nitrogen and their interaction on the grain yield, water and nitrogen use efficiencies of wheat. Crops, 2023(4): 7-15. (in Chinese)
[41]	曹珍珍, 张其芳, 韦克苏, 杨卫丽, 刘光快, 程方民. 水稻籽粒氮代谢几个关键酶对花后高温胁迫的响应及其与贮藏蛋白积累关系. 作物学报, 2012, 38(1): 99-106.
	CAO Z Z, ZHANG Q F, WEI K S, YANG W L, LIU G K, CHENG F M. Response of some key enzyme activities involved in nitrogen metabolism to high temperature at filling stage and its relation to storage protein accu-mulation in rice grain. Acta Agronomica Sinica, 2012, 38(1): 99-106. (in Chinese)
[42]	王利群, 王勇, 董英, 王文兵. 硝酸盐对硝酸还原酶活性的诱导及硝酸还原酶基因的克隆. 生物工程学报, 2003, 19(5): 632-635.
	WANG L Q, WANG Y, DONG Y, WANG W B. Induced activity of nitrate reductase by nitrate and cloning of nitrate reductase gene. Chinese Journal of Biotechnology, 2003, 19(5): 632-635. (in Chinese)
[43]	叶利庭, 吕华军, 宋文静, 图尔迪, 沈其荣, 张亚丽. 不同氮效率水稻生育后期氮代谢酶活性的变化特征. 土壤学报, 2011, 48(1): 132-140.
	YE L T, LÜ H J, SONG W J, TU E D, SHEN Q R, ZHANG Y L. Variation of activity of n metabolizing enzymes in rice plants different in n use efficiency at their late growth stages. Acta Pedologica Sinica, 2011, 48(1): 132-140. (in Chinese)
[44]	DANIEL R M, DANSON M J. Temperature and the catalytic activity of enzymes: A fresh understanding. FEBS Letters, 2013, 587(17): 2738-2743. doi: 10.1016/j.febslet.2013.06.027 pmid: 23810865
[45]	刘磊, 宋娜娜, 齐晓丽, 崔克辉. 水稻根系特征与氮吸收利用效率关系的研究进展. 作物杂志, 2022(1): 11-19.
	LIU L, SONG N N, QI X L, CUI K H. Research advances on the relationship between root characteristics and nitrogen uptake and utilization efficiency in rice. Crops, 2022(1): 11-19. (in Chinese)
[46]	彭超军, 董海滨, 齐学礼, 华夏, 高崇, 赵明忠, 胡琳. 不同籽粒氮含量小麦的氮素转运、分配和相关生理特性研究. 麦类作物学报, 2025, 1-8.
	PENG C J, DONG H B, QI X L, HUA X, GAO C, ZHAO M Z, HU L. Research on N translocation, distribution, and related physiological characteristics of wheat with different grain N contents. Journal of Cereal Crops, 2025, 1-8. (in Chinese)
[47]	ANAS M, LIAO F, VERMA K K, SARWAR M A, MAHMOOD A, CHEN Z L, LI Q, ZENG X P, LIU Y, LI Y R. Fate of nitrogen in agriculture and environment: Agronomic, eco-physiological and molecular approaches to improve nitrogen use efficiency. Biological Research, 2020, 53(1): 47. doi: 10.1186/s40659-020-00312-4 pmid: 33066819
[48]	BAI E, LI S L, XU W H, LI W, DAI W W, JIANG P. A meta-analysis of experimental warming effects on terrestrial nitrogen pools and dynamics. New Phytologist, 2013, 199(2): 441-451.

年份 Year	全氮 Total N (g·kg^-1)	有机质 Organic matter (g·kg^-1)	有效磷 Available phosphorous (mg·kg^-1)	碱解氮 Alkali hydrolyzed nitrogen (mg·kg^-1)	速效钾 Rapidly-available potassium (mg·kg^-1)
2022—2023	1.28	13.64	31.43	73.63	210.59
2023—2024	0.98	10.65	33.67	61.97	212.79