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作物学报 ›› 2025, Vol. 51 ›› Issue (4): 1077-1090.doi: 10.3724/SP.J.1006.2025.41035

• 耕作栽培·生理生化 • 上一篇    下一篇

花后高温干旱逆境对冬小麦光合特性和产量的影响

李乔1,2(), 叶杨春2, 常旭虹2, 王德梅2, 王艳杰2, 杨玉双2, 马瑞琦2, 赵广才2, 蔡瑞国1, 张敏1,*(), 刘希伟2,*()   

  1. 1河北科技师范学院农学与生物科技学院 / 河北省作物逆境生物学重点实验室, 河北昌黎 066004
    2中国农业科学院作物科学研究所 / 农业农村部作物生理生态重点实验室, 北京 100081
  • 收稿日期:2024-05-20 接受日期:2024-12-12 出版日期:2025-04-12 网络出版日期:2024-12-18
  • 通讯作者: 刘希伟, E-mail: liuxiwei@caas.cn; 张敏, E-mail: zhangmin625@126.com
  • 作者简介:E-mail: liqiao3820@163.com
  • 基金资助:
    国家重点研发计划项目(2023YFD2300202);财政部和农业农村部国家现代农业产业技术体系建设专项(CARS-03-16);中国农业科学院作物科学研究所“所级统筹”基本科研业务费(S2023QH06)

Effects of high temperature and drought stresses on photosynthetic characteristics and yield of winter wheat after anthesis

LI Qiao1,2(), YE Yang-Chun2, CHANG Xu-Hong2, WANG De-Mei2, WANG Yan-Jie2, YANG Yu-Shuang2, MA Rui-Qi2, ZHAO Guang-Cai2, CAI Rui-Guo1, ZHANG Min1,*(), LIU Xi-Wei2,*()   

  1. 1College of Agronomy and Biotechnology, Hebei Normal University of Science and Technology / Hebei Key Laboratory of Crop Stress Biology, Changli 066004, Hebei, China
    2Institute of Crop Sciences, Chinese Academy of Agricultural Sciences / Key Laboratory of Crop Physiology and Ecology, Ministry of Agriculture and Rural Affairs, Beijing 100081, China
  • Received:2024-05-20 Accepted:2024-12-12 Published:2025-04-12 Published online:2024-12-18
  • Contact: E-mail: liuxiwei@caas.cn; E-mail: zhangmin625@126.com
  • Supported by:
    National Key Research and Development Program of China(2023YFD2300202);China Agriculture Research System of MOF and MARA(CARS-03-16);Institute of Crop Sciences, Chinese Academy of Agricultural Sciences “Institute-level Coordination” Basic Research Business Fees(S2023QH06)

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摘要：

为了明确花后高温干旱逆境对小麦产量的影响及其生理机制, 本研究于2022—2023年冬小麦生长季, 选用中麦36 (ZM36)和济麦22 (JM22) 2个小麦品种, 在北京和河北赵县大田试验条件下设置花后高温(HT)、干旱(DS)、高温干旱复合胁迫(DHS) 3种逆境处理, 以自然环境为对照(CK), 比较了不同逆境处理对小麦光合特性、叶片衰老和籽粒产量的影响。 结果表明,与CK相比, 在逆境下北京试验点ZM36的产量、穗粒数、千粒重分别降低了18.0%~40.2%、10.4%~16.3%和6.9%~22.7%; JM22分别降低了18.2%~32.8%、3.1%~8.7%和4.0%~14.6%。赵县试验点ZM36的产量、穗粒数、千粒重分别降低了6.4%~27.8%、8.2%~23.1%和2.9%~11.0%; JM22分别降低了6.8%~35.3%、8.0%~19.0%和0.6%~7.7%。2个试验点不同逆境对2个小麦品种产量的影响程度均为DHS>DS>HT。花后逆境下2个小麦品种叶面积指数(LAI)降低14.4%~36.9%, 旗叶叶绿素相对含量(SPAD)减少11.2%~24.6%、叶片持绿时间(Chltotal)缩短1.8~5.0 d, PSII最大光化学效率(Fv/Fm)降低3.5%~10.5%, 非光化学淬灭系数(NPQ)增加9.3%~27.8%, 致使旗叶净光合速率(Pn)降低14.3%~39.6%, 且花后高温干旱复合胁迫对小麦旗叶光合特性的影响远大于单一干旱、高温胁迫。由结构方程可知, 叶片温度(Tleaf)与土壤体积含水量(SVC)、SPAD、Pn呈负相关, SVC与LAI、SPAD、Pn、Fv/Fm呈正相关, 表明较高的土壤体积含水量(0~20 cm土层30%~32%)可降低冠层和叶片温度, 延缓小麦叶片衰老, 提高光合效率。研究结果为小麦高产稳产抗逆栽培提供理论依据。

关键词: 小麦, 高温, 干旱, 复合胁迫, 光合特性, 产量

Abstract:

This study aimed to investigate the effects of high temperature and drought stresses on wheat yield and their underlying physiological mechanisms. Two wheat varieties, Zhongmai 36 (ZM36) and Jimai 22 (JM22), were selected during the winter wheat growing season from 2022 to 2023. Three stress treatments—high temperature (HT), drought (DS), and combined high temperature and drought stress (DHS)—were applied after anthesis under field conditions in Beijing and Zhaoxian, Hebei province, along with a natural environment control (CK). The effects of the stress treatments on photosynthetic characteristics, leaf senescence, and grain yield were compared. Over the two-year period, the yield, grain number per spike, and 1000-grain weight of ZM36 in Beijing decreased by 18.0%-40.2%, 10.4%-16.3%, and 6.9%-22.7%, respectively, while for JM22, the reductions were 18.2%-32.8%, 3.1%-8.7%, and 4.0%-14.6%, respectively. In Zhaoxian, the yield, grain number per spike, and 1000-grain weight of ZM36 declined by 6.4%-27.8%, 8.2%-23.1%, and 2.9%-11.0%, respectively, while JM22 experienced decreases of 6.8%-35.3%, 8.0%-19.0%, and 0.6%-7.7%, respectively. The yield reductions followed the order: DHS > DS > HT. Additionally, the leaf area index (LAI) of both wheat varieties decreased by 14.4%-36.9%, the relative chlorophyll content (SPAD) of the flag leaf declined by 11.2%-24.6%, and the leaf stay-green duration (Chltotal) was shortened by 1.8-5.0 days. As a result, the net photosynthetic rate (Pn) of the flag leaf decreased by 14.3%-39.6%, the maximum photochemical efficiency of PSII (Fv/Fm) was reduced by 3.5%-10.5%, and the non-photochemical quenching coefficient (NPQ) increased by 9.3%-27.8%. The combined high temperature and drought stress after anthesis had a much greater impact on the photosynthetic characteristics of flag leaves than individual drought or high temperature stress. Structural equation modeling revealed that leaf temperature (Tleaf) was negatively correlated with soil volumetric water content (SVC), SPAD, and Pn, while SVC was positively correlated with LAI, SPAD, Pn, and Fv/Fm. Furthermore, higher soil volumetric water content (30%-32% in the 0-20 cm soil layer) reduced canopy and leaf temperatures, delayed wheat leaf senescence, and improved photosynthetic efficiency. These findings provide a theoretical basis for strategies to achieve high and stable wheat yields under stress conditions.

Key words: wheat, high temperature, drought, combined stresses, photosynthetic characteristics, yield

图1

2022-2023年度北京试验点和赵县试验点逆境处理期间试验小区的小气候变化 CK: 对照; HT: 高温; DS: 干旱; DHS: 高温干旱复合逆境。"

表1

花后高温、干旱及高温干旱复合逆境对小麦产量及产量构成因素的影响"

地点
Location		 品种
Cultivar		 处理
Treatment		 产量
Yield
(kg hm-2)		 穗数
Spikes
(×104 hm-2)		 穗粒数
Kernels per spike		 千粒重
1000-grain weight (g)		 不孕小穗数
Infertile spikelet
per spike
北京
Beijing		 中麦36
ZM36		 CK 8933.0 a 520.1 a 44.2 a 46.2 a 1.6 b
HT 7326.9 b 498.3 a 39.6 b 43.0 b 2.3 ab
DS 5464.8 c 506.7 a 37.8 c 37.7 c 2.1 b
DHS 5345.3 c 496.7 a 37.0 c 35.7 d 2.4 a
济麦22
JM22		 CK 9229.8 a 538.8 a 42.4 a 47.1 a 1.0 a
HT 7548.9 b 522.5 a 41.1 ab 45.2 b 1.0 a
DS 6786.2 c 522.5 a 40.5 b 42.6 c 1.1 a
DHS 6199.7 d 515.0 a 38.7 c 40.2 d 1.4 a
河北赵县Zhaoxian, Hebei 中麦36
ZM36		 CK 9762.9 a 689.1 a 36.4 a 48.4 a 2.7 b
HT 9133.8 b 685.1 a 30.6 bc 47.0 b 3.1 ab
DS 8996.7 b 681.0 a 33.4 ab 45.7 c 2.6 b
DHS 7052.1 c 672.0 a 28.0 c 43.1 d 4.5 a
济麦22
JM22		 CK 9575.6 a 716.1 a 31.1 a 49.3 a 2.9 b
HT 8924.9 b 709.1 a 26.2 bc 49.0 a 3.9 b
DS 8088.8 c 695.1 a 28.6 ab 47.3 b 2.8 a
DHS 6199.8 d 708.0 a 25.2 c 45.5 c 4.1 a
地点 Location (L) ** ** ** ** **
品种 Cultivar (C) ns ** ** ** **
处理 Treatment (T) ** ns ** ** **
L×C ** ns ** * **
L×T ** ns ** ** **
C×T ns ns ** ** ns
L×C×T ** ns ns ** ns

表2

不同处理产量与产量因素的相关系数和通径系数"

因素
Factor		 与产量的简单相关系数
Correlation coefficient with yield		 直接通径系数
Direct path coefficient		 间接通径系数
Indirect path coefficient
X1 X2 X3
X1 0.50** -0.01 — 0.21 -0.69
X2 -0.20 0.27* -0.01 — -0.37
X3 0.84** 0.97** -0.01 -0.10 —

图2

花后不同逆境对小麦叶面积指数(LAI)和旗叶叶绿素相对含量(SPAD)的影响 处理同图1。不同小写字母表示不同处理间存在显著差异(P < 0.05); ns: 不显著; n.d.: 未测量。"

表3

不同逆境处理下小麦旗叶衰老模型及衰老相关参数"

地点
Location		 品种
Cultivar		 处理
Treatment		 旗叶衰老模型
Flag leaf senescence model		 最大衰老速率
Rmax		 平均衰老速率
Rave		 衰老速率达
最大的时期
M		 叶绿素
总持续期
Chltotal		 叶绿素稳定持续期
Chlper		 快速
衰老期
Chlloss
北京
Beijing		 中麦36
ZM36		 CK G=55.68 × e(-0.0005 × e(0.24t)) R2=0.998 5.02 b 2.51 b 30.88 a 34.31 a 21.47 a 12.84 a
HT G=55.65 × e(-0.0017 × e(0.24t)) R2=0.996 4.83 b 2.41 b 28.72 b 32.22 b 18.16 b 14.07 a
DS G=54.18 × e(-0.0004 × e(0.29t)) R2=0.991 5.78 a 2.89 a 26.88 c 29.69 c 17.09 bc 12.60 a
DHS G=53.82 × e(-0.0005 × e(0.29t)) R2=0.993 5.48 ab 2.74 ab 26.32 d 29.30 c 16.61 c 12.68 a
济麦22
JM22		 CK G=61.40 × e(-0.0003 × e(0.26t)) R2=0.990 4.86 bc 2.43 bc 31.24 a 34.39 a 22.23 a 12.16 b
HT G=62.13 × e(-0.0050 × e(0.18t)) R2=0.998 4.20 c 2.10 c 28.76 b 33.33 b 17.09 b 16.24 a
DS G=61.02 × e(-0.0021 × e(0.23t)) R2=0.999 5.22 ab 2.61 ab 26.58 c 30.19 c 16.86 b 13.32 b
DHS G=61.22 × e(-0.0034 × e(0.22t)) R2=0.993 6.01 a 3.00 a 26.31 c 30.19 c 16.07 b 14.12 b
河北
赵县
Zhaoxian, Hebei		 中麦36
ZM36		 CK G=54.76 × e(-0.0002 × e(0.27t)) R2=0.996 5.43 a 2.72 a 30.64 a 33.71 a 21.41 a 12.30 a
HT G=53.72 × e(-0.0003 × e(0.29t)) R2=0.993 5.82 a 2.91 a 28.53 b 31.37 b 19.95 ab 11.42 a
DS G=51.82 × e(-0.0004 × e(0.29t)) R2=0.996 5.46 a 2.73 a 27.26 c 30.12 c 18.06 bc 12.06 a
DHS G=50.62 × e(-0.0003 × e(0.30t)) R2=0.999 5.55 a 2.78 a 26.50 d 29.24 d 17.24 c 12.00 a
济麦22
JM22		 CK G=60.28 × e(-0.0009 × e(0.22t)) R2=0.998 4.96 ab 2.48 ab 31.15 a 34.88 a 20.90 a 13.98 ab
HT G=60.06 × e(-0.0035 × e(0.20t)) R2=0.998 4.41 b 2.21 b 28.80 b 33.03 b 17.65 b 15.39 a
DS G=58.45 × e(-0.0013 × e(0.25t)) R2=0.999 5.37 ab 2.69 ab 26.84 c 30.19 c 17.52 b 12.68 b
DHS G=58.28 × e(-0.0009 × e(0.27t)) R2=0.999 5.90 a 2.95 a 25.55 d 28.60 d 17.06 b 11.54 b

图3

花后不同逆境对小麦净光合速率(Pn)、气孔导度(Gs)和蒸腾速率(Tr)的影响 处理同图1。不同小写字母表示不同处理间存在显著差异(P < 0.05); ns: 不显著。"

图4

不同逆境下叶片温度与叶面积指数(LAI)、叶绿素相对含量(SPAD)、叶表相对湿度(RHsfc)、净光合速率(Pn)的相关性分析 处理同图1。*: P < 0.05; **: P < 0.01。"

图5

花后不同逆境对小麦旗叶荧光参数的影响 处理同图1。F0: 基础荧光; Fm: 最大荧光; Fv: 可变荧光; Fm-less: 稳态荧光; Fv/Fm: PSII最大光化学效率; Fv/F0: PSII潜在光化学效率; NPQ: 非光化学淬灭系数。不同小写字母表示不同处理间存在显著差异(P < 0.05); ns: 不显著。"

表4

不同处理下粒重与相关性状的灰色关联度"

指标
Indicator		 CK HT DS DHS
关联度
Correlation
degree		 排名
Ranking		 关联度
Correlation
degree		 排名
Ranking		 关联度
Correlation
degree		 排名
Ranking		 关联度
Correlation
degree		 排名
Ranking
穗粒数 Kernels per spike 0.8135 1 0.5481 8 0.5256 6 0.7887 1
平均衰老速率 Rave 0.7079 2 0.5096 9 0.7481 1 0.5720 7
叶绿素总持续时间 Chltotal 0.5170 9 0.7506 2 0.5525 5 0.6818 4
净光合速率 Pn 0.6661 4 0.6108 7 0.6888 2 0.7039 3
气孔导度 Gs 0.6170 6 0.6695 6 0.4802 9 0.6718 6
蒸腾速率 Tr 0.6300 5 0.7016 5 0.5086 7 0.5447 8
PSII最大光化学效率 Fv/Fm 0.5352 8 0.7375 3 0.6027 3 0.6753 5
PSII潜在光化学效率 Fv/F0 0.5866 7 0.7328 4 0.5086 7 0.7306 2
非光化学淬灭系数 NPQ 0.6681 3 0.7595 1 0.5554 4 0.5231 9

图6

旗叶光合特性与形态性状的关系 (a): 左下济麦22, 右上中麦36; (b): 主成分分析图; (c): 结构方程模型(SEM)模型中χ2=9.80, df=8, P=0.28, GFI=0.97, CFI=0.99, NFI=0.99, TLI=0.99, RMR=0.002, RMSEA=0.049, 黑色和红色实线箭头分别表示显著性正、负相关, 虚线箭头表示无显著性相关; *: P < 0.05, **: P < 0.01。SVC: 土壤体积含水量; LAI: 叶面积指数; SPAD: 叶绿素相对含量; Tleaf: 叶片温度; Pn: 净光合速率; Gs: 气孔导度; Tr: 蒸腾速率; RHsfc: 叶表相对湿度; F0: 基础荧光; Fm: 最大荧光; Fv: 可变荧光; Fm-less: 稳态荧光; Fv/Fm: PSII最大光化学效率; Fv/F0: PSII潜在光化学效率; NPQ: 非光化学淬灭系数。"

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