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作物学报

作物学报 ›› 2026, Vol. 52 ›› Issue (4): 1251-1267.doi: 10.3724/SP.J.1006.2026.55057

• 耕作栽培·生理生化 • 上一篇    下一篇

CO2和NaCl对大豆气孔特征、光合性能及抗氧化系统的影响

李坤1(), 籍亚腾1, 田银帅2, 胡民航1, 刘亮1, 李菲1, 李国强1, 郝立华1,*(), 郑云普1,*()   

  1. 1河北工程大学水利水电学院, 河北邯郸056038
    2河北工程大学园林与生态工程学院, 河北邯郸056038
  • 收稿日期:2025-08-21 接受日期:2026-01-22 出版日期:2026-04-12 网络出版日期:2026-02-05
  • 通讯作者: *郑云普, E-mail: zhengyunpu@hebeu.edu.cn; 郝立华, E-mail: haolihua_000@sina.com
  • 作者简介:17836923301@163.com
  • 基金资助:
    国家自然科学基金项目(32071608);河北省自然科学基金项目(E2025402107);河北省自然科学基金项目(E2024402128);河北省硕士在读研究生创新能力培养资助项目(CXZZSS2025081)

Effects of CO2 and NaCl on stomatal traits, photosynthetic performance, and antioxidant systems in soybean

Li Kun1(), Ji Ya-Teng1, Tian Yin-Shuai2, Hu Min-Hang1, Liu Liang1, Li Fei1, Li Guo-Qiang1, Hao Li-Hua1,*(), Zheng Yun-Pu1,*()   

  1. 1School of Water Conservancy and Hydropower, Hebei University of Engineering, Handan 056038, Hebei, China
    2School of Landscape and Ecological Engineering, Hebei University of Engineering, Handan 056038, Hebei, China
  • Received:2025-08-21 Accepted:2026-01-22 Published:2026-04-12 Published online:2026-02-05
  • Contact: *E-mail: zhengyunpu@hebeu.edu.cn; E-mail: haolihua_000@sina.com
  • Supported by:
    National Natural Science Foundation of China(32071608);Hebei Natural Science Foundation of China(E2025402107);Hebei Natural Science Foundation of China(E2024402128);Hebei Province Graduate Innovation Ability Training Project for Master’s Students(CXZZSS2025081)

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摘要：

为深入理解未来大气CO2浓度升高背景下农田生态系统对NaCl胁迫的响应机制, 利用可精准控制CO2浓度的环境生长箱, 探讨了NaCl胁迫条件下CO2浓度升高对大豆气孔形态及其分布、叶片气体交换参数以及抗氧化酶系统的影响。结果表明, 在环境正常CO2浓度条件下, NaCl胁迫降低近轴面的气孔密度、气孔长度、气孔宽度、气孔周长及气孔面积, 而CO2浓度升高增加了气孔密度, 并且导致气孔分布更规则, 从而降低了NaCl对叶片气体交换效率的影响。另外, NaCl胁迫导致净光合速率(Pn)降低, 而CO2浓度升高导致NaCl胁迫(S0、S50、S100和S150)条件下叶片水分利用效率相比环境CO2浓度分别提高188%、243%、97%和85%, 表明CO2浓度升高可以有效缓解NaCl对大豆造成的生理胁迫。另外, CO2浓度升高提高了NaCl胁迫下过氧化物酶活性、超氧化物歧化酶活性以及脯氨酸含量, 且同时降低了重度NaCl胁迫下的丙二醛含量。此外, NaCl胁迫条件下, CO2浓度升高使得GmCLC-d1、GmCLC-d2和GmNHX1的表达量降低, GmNcl1的表达量提高。本研究结果表明, 大气CO2浓度升高可以通过优化气孔形态及分布、增强光化学与生化过程、提升抗氧化防御能力以及调控耐盐基因表达来缓解NaCl胁迫对大豆造成的生理伤害。本研究结果将为深入理解未来气候变化背景下大豆对大气CO2浓度升高和NaCl胁迫的生理及分子响应机制提供理论依据。

关键词: 大气CO2浓度, NaCl胁迫, 气孔特征, 光合性能, 抗氧化系统

Abstract:

To gain a deeper understanding of the response mechanisms of farmland ecosystems under NaCl stress in the context of future elevated atmospheric CO2 concentrations, this study employed environmental growth chambers with precisely controlled CO2 levels to investigate the effects of elevated CO2 on stomatal morphology and distribution, leaf gas exchange parameters, and the antioxidant enzyme system in soybeans under NaCl stress. The results showed that under ambient CO2 conditions, NaCl stress reduced stomatal density, length, width, perimeter, and area on the adaxial surface. In contrast, elevated CO2 increased stomatal density and promoted a more regular stomatal distribution, thereby mitigating the negative effects of NaCl on leaf gas exchange efficiency. Moreover, NaCl stress decreased the net photosynthetic rate (Pn), while elevated CO2 significantly improved leaf water use efficiency under NaCl stress (S0, S50, S100, and S150) by 188%, 243%, 97%, and 85%, respectively, compared to ambient CO2, indicating that elevated CO2 can effectively alleviate NaCl-induced physiological stress in soybeans. Elevated CO2 also enhanced peroxidase and superoxide dismutase activities and increased proline content, while reducing malondialdehyde levels under severe NaCl stress. Furthermore, under NaCl stress, elevated CO2 downregulated the expression of GmCLC-d1, GmCLC-d2, and GmNHX1, and upregulated GmNcl1. These findings demonstrate that elevated atmospheric CO2 can alleviate the physiological damage caused by NaCl stress in soybeans by optimizing stomatal traits, enhancing photochemical and biochemical processes, boosting antioxidant defenses, and regulating salt-tolerance gene expression. This study provides a theoretical basis for understanding the physiological and molecular responses of soybeans to elevated CO2 and NaCl stress under future climate change scenarios.

Key words: atmospheric CO2 concentration, NaCl stress, stomatal traits, photosynthetic performance, antioxidant system

表1

qRT-PCR 引物序列"

基因名称
Gene name		 正向引物
Forward primer (5'-3')		 反向引物
Reverse primer (5'-3')		 产物长度
Product length (bp)
GmCLC-d1 GCTGGTCAATTTGTTCCTGG AGAGATACTGTCATCCGCATTG 150
GmCLC-d2 GCAAAGCCTGTTTCTAGTAAAGG TGCCTAAGACCAAGTTGACG 158
GmNcl1 AGCTGTTCATGGTGCATCTC TTGACCGGACTTTGACTTGG 142
GmNHX1 AGCATCACGATAATCACAGATCC CTTACGCCACAAACGATGAAC 165
GmRCAa AGCTTCAGTTCCCAGTTCAG CCTTTCCATCTGTCCTTGTCTG 149
GmRCAb CTGAATGGTTCTGGAGCTGG CATCTGTCCTTGTCGGTCTG 156
GmRCA11 AGAAGGTTATTGGCTCAAGGG TCCTCTTGTGATGTCCTGTTG 154
GmRCA14 ATTCGGTTCCCAGTTCAGC CCTTTCCATCTGTCCTTGTTTG 151
GmEF-1α CTGACTGTGCTGTTCTTATCATTG ATACCTGGCCTTGGAATACTTG 145

图1

CO2和NaCl对大豆植株生物量的影响 [CO2]: CO2浓度; [NaCl]: NaCl浓度。不同小写字母表示在0.05水平差异显著; *、**和***分别表示在0.05、0.01和0.001水平具有显著差异。"

图2

CO2和NaCl对大豆叶片光合色素含量的影响 处理及缩写同图1。FW: 鲜重。Chl a: 叶绿素 a; Chl b: 叶绿素 b; Tchl: 总叶绿素; Caro: 类胡萝卜素。不同小写字母表示在0.05水平差异显著; *、**和***分别表示在0.05、0.01和0.001水平具有显著差异。"

表2

CO2和NaCl对大豆近轴面气孔形态特征的影响"

气孔特征
Stomatal
trait		 CO2
浓度
[CO2]		 NaCl浓度
[NaCl]		 气孔密度
Stomatal
density
(No. mm-2 )		 气孔长度
Stomatal length
(μm)		 气孔宽度
Stomatal width
(μm)		 气孔周长
Stomatal
perimeter
(μm)		 气孔面积
Stomatal area
(μm2 )		 形状指数
Shape index
(%)
近轴面
Adaxial
surface		 C400 S0 22.50±2.38 c 13.25±0.8 a 4.14±0.6 a 35.12±0.7 ab 53.56±2.0 a 1.26±0.06 c
S50 20.00±1.93 c 12.71±1.2 ab 3.17±0.4 b 33.59±0.5 b 41.85±1.6 b 1.35±0.08 c
S100 12.00±2.16 d 12.10±1.6 ab 1.98±0.3 c 25.95±2.3 c 31.79±0.8 c 1.54±0.07 a
S150 11.75±1.71 d 11.42±1.2 b 1.98±0.4 c 23.43±0.3 d 24.08±0.6 e 1.40±0.05 b
C800 S0 71.75±3.30 a 12.02±1.0 ab 3.62±0.4 ab 36.74±2.3 a 51.71±3.3 a 1.28±0.03 c
S50 69.25±9.54 a 12.36±0.9 ab 3.12±0.7 b 33.70±1.6 b 44.24±2.9 b 1.41±0.07 b
S100 37.75±6.95 b 12.26±0.6 ab 1.94±0.2 c 26.79±0.6 c 32.74±0.8 c 1.59±0.05 a
S150 34.00±1.83 b 11.11±0.7 b 1.95±0.2 c 25.22±0.5 cd 28.06±2.2 d 1.61±0.05 a
CO2浓度[CO2] *** ns ns *** ns ***
NaCl浓度[NaCl] *** ** *** *** *** ***
[CO2]×[NaCl] *** ns ns ns ns **

表3

CO2和NaCl对大豆远轴面气孔形态特征的影响"

气孔
特征
Stomatal trait		 CO2浓度
[CO2]		 NaCl浓度
[NaCl]		 气孔密度
Stomatal
density
(No. mm-2 )		 气孔长度
Stomatal length
(μm)		 气孔宽度
Stomatal width
(μm)		 气孔周长
Stomatal perimeter
(μm)		 气孔面积
Stomatal area
(μm2 )		 形状指数
Shape index
(%)
远轴面
Abaxial surface		 C400 S0 51.75±3.50 b 13.85±1.2 a 3.33±0.1 a 25.27±0.2 a 44.29±1.3 a 1.33±0.02 a
S50 22.00±4.90 c 14.19±1.4 a 2.91±0.2 ab 24.65±1.8 a 34.11±2.8 b 1.26±0.03 a
S100 23.00±6.68 c 11.82±0.7 ab 2.51±0.2 bc 21.74±1.3 a 32.18±0.7 c 1.24±0.05 a
S150 20.75±2.63 c 12.71±1.9 a 2.31±0.4 c 18.19±3.3 a 35.37±2.0 e 1.30±0.04 a
C800 S0 85.00±3.37 a 10.62±0.3 b 3.19±0.2 a 30.35±1.3 a 35.50±1.4 a 1.27±0.01 a
S50 51.50±10.34 b 13.00±2.0 ab 1.83±0.2 d 29.68±2.3 a 30.20±0.8 b 1.44±0.08 a
S100 49.00±7.07 b 10.23±2.2 b 3.07±0.7 a 29.31±1.3 a 26.97±2.7 c 1.40±0.27 a
S150 44.75±8.85 b 13.20±2.0 ab 2.40±0.2 bc 25.62±1.4 a 34.28±2.3 d 1.35±0.03 a
CO2浓度[CO2] *** ** ns *** *** ns
NaCl浓度[NaCl] *** ** *** *** *** *
[CO2]×[NaCl] ns ** *** ns ** ns

图3

大豆叶片气孔光学显微照片 A: 近轴面环境CO2浓度; B: 远轴面环境CO2浓度; C: 近轴面升高CO2浓度; D: 远轴面升高CO2浓度。标尺: 50 μm。"

图4

CO2和NaCl对大豆气孔空间分布格局的影响 处理同图1。Lhat (d): 最小邻域距离; Upper 95%: 95%置信区间上界; Lower 95%: 95%置信区间下界。"

图5

CO2和NaCl对大豆气体交换参数的影响 处理及缩写同图1。Pn: 净光合速率; Tr: 蒸腾速率; Gs: 气孔导度; WUEI: 瞬时水分利用效率。不同小写字母表示在0.05水平差异显著; **和***分别表示在0.01和0.001水平具有显著差异。"

图6

CO2和NaCl对大豆最大羧化速率(Vcmax)和最大电子传递效率(Jmax)的影响 处理及缩写同图1。Vcmax: 最大羧化速率; Jmax: 最大电子传递速率。不同小写字母表示在0.05水平差异显著; *、**和***分别表示在0.05、0.01和0.001水平具有显著差异。"

图7

CO2和NaCl对抗氧化酶系统的影响 处理及缩写同图1。FW: 鲜重; POD: 过氧化物酶; MDA: 丙二醛; SOD: 超氧化物歧化酶; Pro: 脯氨酸。不同小写字母表示0.05水平差异显著; *、**和***分别表示在0.05、0.01以及0.001水平下具有显著差异。"

图8

CO2和NaCl对大豆Rubisco活化酶编码基因表达量的影响 处理同图1。"

图9

CO2和NaCl对大豆抗盐基因表达量的影响 处理同图1。"

图10

CO2和NaCl处理下大豆叶片Pn与生理生化指标关系拟合 处理同图1。Pn: 净光合速率; ADW: 近轴面气孔宽度; ADP: 近轴面气孔周长; ADA: 近轴面气孔面积; ADD: 近轴面气孔密度; Vcmax: 最大羧化速率; Jmax: 最大电子传递速率。"

图11

CO2和NaCl处理下大豆叶片Pn与植株生物量关系拟合 Pn: 净光合速率。"

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