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作物学报 ›› 2025, Vol. 51 ›› Issue (4): 1061-1076.doi: 10.3724/SP.J.1006.2025.41054

• 耕作栽培·生理生化 • 上一篇    下一篇

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

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()   

  1. 山西农业大学农学院, 山西太谷 030801
  • 收稿日期:2024-08-08 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: 郝兴宇, E-mail: haoxingyu1976@126.com
  • 作者简介:E-mail: wj0704zsc@126.com
  • 基金资助:
    山西农业大学青年拔尖创新人才支持计划项目(BJRC201602);山西农业大学农学院研究生教育改革与质量提升工程顶目(2023YDT03);山西农业大学农学院研究生教育改革与质量提升工程顶目(2023YCX05);山西农业大学农学院研究生教育改革与质量提升工程顶目(2023YCX42)

Effects of elevated CO2 concentration, increased temperature and their interaction on the carbon and nitrogen metabolism in Liangxing 99 winter wheat leaves

WANG Jiao(), BAI Hai-Xia, HAN Yu-Yan, LIANG Hui, FENG Ya-Nan, ZHANG Dong-Sheng, LI Ping, ZONG Yu-Zheng, SHI Xin-Rui, HAO Xing-Yu()   

  1. College of Agriculture, Shanxi Agricultural University, Taigu 030801, Shanxi, China
  • Received:2024-08-08 Accepted:2024-12-12 Published:2025-04-12 Published online:2024-12-18
  • Contact: E-mail: haoxingyu1976@126.com
  • Supported by:
    Youth Talents Support Program of Shanxi Agricultural University(BJRC201602);Graduate Education Reform and Quality Improvement Program of College of Agriculture, Shanxi Agricultural University(2023YDT03);Graduate Education Reform and Quality Improvement Program of College of Agriculture, Shanxi Agricultural University(2023YCX05);Graduate Education Reform and Quality Improvement Program of College of Agriculture, Shanxi Agricultural University(2023YCX42)

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摘要：

单一的大气CO2浓度升高或温度升高均已被证实对小麦的生长和产量产生显著影响, 然而, 关于CO2浓度和温度共同升高对小麦整个生育期的影响研究很少。本研究以冬小麦品种“良星99”为材料, 利用人工控制气候室, 设置2种CO2浓度水平(环境大气CO2浓度和环境大气CO2浓度+200 μmol mol-1)和2种温度水平(环境温度和环境温度+2℃), 测定其对冬小麦物候期、光合作用、碳代谢、氮同化和产量的影响。 结果表明:大气CO2浓度升高使拔节期、开花期和灌浆期净光合速率和水分利用效率增加, 使灌浆期可溶性糖含量升高, 虽灌浆期谷氨酰胺合成酶和谷丙转氨酶活性降低, 但仍可通过增加穗数, 进而使生物量和产量增加32.8%和30.0%。升温会缩短冬小麦全生育期, 使拔节期谷氨酸合成酶活性下降, 开花期水分利用效率、可溶性糖、淀粉和蔗糖含量、谷氨酸合成酶活性降低, 灌浆期蔗糖合成酶和谷氨酰胺合成酶活性下降, 生物量降低12.2%, 产量无显著影响。大气CO2浓度升高促进升温下冬小麦提前开花, 并延长花后籽粒灌浆时间。升温条件下, 大气CO2浓度升高通过上调拔节期TaRUBP1的表达进而提高净光合速率, 提高拔节期可溶性糖含量和开花期蔗糖含量, 从而缓解升温对光合同化的负面影响。升温条件下, CO2浓度升高上调开花期TaGS2和灌浆期TaNR的表达进而增加开花期谷氨酰胺合成酶活性和灌浆期硝酸还原酶活性, 缓解升温对氮同化的抑制作用。此外, CO2浓度升高减轻升温对冬小麦生物量的负面影响, 并通过增加穗数提高升温条件下冬小麦产量23.9%。总之, CO2浓度通过提高光合能力、增加光合同化物积累、促进氮同化、延长灌浆时间等缓解升温对冬小麦生物量的负效应, 并通过增加穗数提高冬小麦产量。

关键词: 冬小麦, 碳氮代谢, 生长, 产量, CO2浓度升高, 温度升高

Abstract:

It is well established that either elevated atmospheric CO2 concentration or increased temperature alone can significantly influence wheat growth and yield. However, limited research has explored the combined effects of elevated CO2 and increased temperature on wheat throughout its entire growth period. In this study, winter wheat (Triticum aestivum “Liangxing99”) was cultivated in environment-controlled chambers under two CO2 concentrations (ambient and ambient + 200 μmol mol-1 and two temperature regimes (ambient and ambient + 2℃). Phenological development, photosynthesis, carbohydrate metabolism, nitrogen assimilation, and yield were systematically investigated. Elevated CO2 enhanced the net photosynthetic rate and water use efficiency during the elongation, anthesis, and grain filling stages, while also increasing soluble sugar content during the grain filling stage. Although the activities of glutamine synthetase (GS) and glutamic-pyruvic transaminase (GPT) decreased during grain filling, elevated CO2 still increased biomass and yield by 32.8% and 30.0%, respectively, primarily by increasing the number of grains. In contrast, increased temperature shortened the overall growth period of winter wheat, reduced glutamate synthetase (GOGAT) activity during the elongation stage, and decreased soluble sugar, starch, and sucrose contents. Additionally, increased temperature lowered water use efficiency and GOGAT activity at anthesis, as well as sucrose synthase and GS activities during grain filling, resulting in a 12.2% reduction in biomass, though yield was not significantly affected. Elevated CO2 mitigated the adverse effects of increased temperature by advancing flowering, extending the grain filling period, and alleviating the inhibition of photosynthesis and nitrogen assimilation. Specifically, elevated CO2 upregulated TaRUBP1 expression during elongation, which enhanced the net photosynthetic rate and increased soluble sugar content at elongation and sucrose content at anthesis. Moreover, elevated CO2 upregulated TaGS2 expression at anthesis and TaNR expression during grain filling, which increased GS activity at anthesis and nitrate reductase (NR) activity during grain filling. These responses alleviated the inhibition of nitrogen assimilation caused by high temperature. Consequently, elevated CO2 mitigated the negative effects of increased temperature on biomass production, while enhancing yield by 23.9% under elevated temperature, primarily through an increased number of grains. In summary, elevated CO2 alleviated the negative impacts of increased temperature on winter wheat biomass by enhancing photosynthetic capacity, promoting the accumulation of photosynthetic assimilates, improving nitrogen assimilation, and extending the grain filling period. Additionally, it increased yield by boosting the number of grains under elevated temperature conditions.

Key words: winter wheat, carbon and nitrogen metabolism, growth, yield, elevated CO2 concentration, increased temperature

图1

小麦生长季各处理CO2浓度变化 CK: 环境CO2浓度, 环境温度; EC: 环境CO2浓度+200 μmol mol-1, 环境温度; ET: 环境CO2浓度, 环境温度+2℃; ECT: 环境CO2浓度+200 μmol mol-1, 环境温度+2℃。"

图2

小麦生长季各处理气温变化 处理同图1。"

表1

基因的引物信息"

基因
Gene		 基因登录号
Gene ID		 基因功能
Gene function		 引物序列
Primer sequence (5′-3′)
TaActin AB181991 内参基因
Reference gene		 F: GGAGAAGCTCGCTTACGTG
R: GGGCACCTGAACCTTTCTGA
TaRUBP KF801504.1 1,5-二磷酸核酮糖羧化/转移酶Ribulose-1,5-bisphosphate carboxylase/oxygenase F: ATTACTTGAATGCGACTGCG
R: CGGCAATAATGAGCCAAAGT
TaPATPaseβ-sub M16843.1 叶绿体ATP合成酶β亚基
ATP synthase β-subunit, chloroplastic		 F: AATGATGCGGAACTTGGT
R: CATACGGCGGGAGTCAT
TaFd1 X75089.1 铁氧还蛋白
Ferredoxin		 F: TCACCTGCCACGCCTAC
R: CATCGAGCAATTCATTCG
TaD1 M21352.1 PSII复合体D1蛋白
PSII protein D1		 F: TGGGCTGACTTGGTTGAC
R: GAAGTTGCGGTCAATAAGGT
TaD2 EU492899.1 PSII复合体D2蛋白
PSII protein D2		 F: AACTTGCTCGGTCTGTTC
R: GCCGCACCTAATACTCC
TaNR TC234027 硝酸还原酶
Nitrate reductase		 F: GGCCAATTCYTTCATCTCCTTCTG
R: TACRTSCACAGATTGATGCGTCSA
TaNIR FJ527909.1 亚硝酸还原酶
Nitrite reductase		 F: CAGGAGAAGGTGAAGCTGG
R: TCATGAACCGCCCATACTG
TaGS1 DQ124209.1 胞质谷氨酰胺合成酶
Glutamine synthetase, cytosolic		 F: CGAGTCGATGAGGAAGGAC
R: CCCCAGCTGAAGGTGTTGA
TaGS2 DQ124212.1 质体谷氨酰胺合成酶
Glutamine synthetase, plastidial		 F: AGAGAATCACGGAGCAAGC
R: ATCACGTCGAAACCTCCATC
TaFd-GOGAT TC394038 质体谷氨酸合酶
Glutamate synthetase, plastidial		 F: TGGTGCCACCCAGCGAAG
R: AGCTCGTTTCCAGAAGATGCCTT

图3

CO2浓度和温度升高对冬小麦物候期的影响 处理同图1。"

图4

CO2浓度和温度升高对冬小麦叶绿素的影响 处理同图1。不同字母表示处理间差异显著(P ≤ 0.05)。"

图5

CO2浓度和温度升高对冬小麦光合参数的影响 处理同图1。不同字母表示处理间差异显著(P ≤ 0.05)。"

图6

CO2浓度和温度升高对冬小麦光合相关基因的影响 处理同图1。不同字母表示处理间差异显著(P ≤ 0.05)。"

图7

CO2浓度和温度升高对冬小麦碳水化合物代谢的影响 处理同图1。不同字母表示处理间差异显著(P ≤ 0.05)。"

图8

CO2浓度和温度升高对冬小麦氮同化的影响 处理同图1。不同字母表示处理间差异显著(P ≤ 0.05)。"

图9

CO2浓度和温度升高对冬小麦氮同化相关基因的影响 处理同图1。不同字母表示处理间差异显著(P ≤ 0.05)。"

图10

CO2浓度和温度升高对冬小麦生物量和产量的影响 处理同图1。不同字母表示处理间差异显著(P ≤ 0.05)。"

表2

CO2浓度和温度升高对冬小麦产量构成的影响"

年份
Year		 处理
Treatment		 单穗粒数
Grain number per spike		 单位面积穗数
Spike number (m-2)		 千粒重
1000-grain weight (g)
2019-2020 CK 35.13±0.79 ab 259.99±37.41 b 40.85±5.49 a
EC 36.28±1.62 a 331.49±23.88 b 44.84±2.27 a
ET 31.02±3.03 ab 285.99±13.20 b 40.55±1.92 a
ECT 28.22±3.28 b 412.74±19.63 a 38.73±0.99 a
2020-2021 CK 29.46±1.53 a 235.62±6.29 b 43.18±1.02 b
EC 29.55±1.64 a 328.24±25.38 a 46.47±1.22 a
ET 32.01±1.92 a 247.81±10.75 b 43.08±0.07 b
ECT 27.87±1.40 a 341.24±5.14 a 42.15±0.82 b
P-value 年份Year ns * ns
CO2 ns ** ns
T ns * ns
CO2´T ns ns ns

图11

CO2浓度和温度升高对冬小麦生物量和产量影响的PLS-PM分析 ES: 拔节期; AS: 开花期; GFS: 灌浆期。光合能力(潜变量)由指标净光合速率(观测变量)反映。碳代谢(潜变量)由指标可溶性糖、淀粉、蔗糖、SUS和SPS (观测变量)反映。氮同化(潜变量)由指标NR、GS、GOGAT和GPT (观测变量)反映。红色实线箭头和蓝色实线箭头分别指示正向和负向的因果关系; 灰色虚线箭头指示路径关系中的非显著性。箭头旁边的数字及其粗细程度共同体现路径系数的大小。潜变量的决定系数通过R2值具体呈现。***表示在P ≤ 0.001水平上显著; **表示在P ≤ 0.01水平上显著; *表示在P ≤ 0.05水平上显著。该模型的拟合优度值为0.6。"

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