大气CO<sub>2</sub>浓度和气温升高对玉米灌浆期碳氮代谢的影响

王娇; 李萍; 宗毓铮; 张东升; 史鑫蕊; 杨净; 郝兴宇

doi:10.12357/cjea.20220395

大气CO₂浓度和气温升高对玉米灌浆期碳氮代谢的影响

doi: 10.12357/cjea.20220395

山西农业大学农学院　太谷　030801

基金项目: 国家自然科学基金面上项目(31871517)、山西省留学人员科技活动择优资助项目(20210041)和山西省基础研究计划青年科学研究项目(20210302124656)资助

详细信息

作者简介:
王娇, 主要研究方向为旱作栽培与作物生理。E-mail: wj0704zsc@126.com

通讯作者:
郝兴宇, 主要研究方向为旱作栽培与作物生理和气候变化对作物的影响。E-mail: haoxingyu1976@126.com

中图分类号: S513
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出版历程
- 收稿日期: 2022-05-23
- 录用日期: 2022-09-15
- 网络出版日期: 2022-11-07
- 刊出日期: 2023-02-10

Effects of increased atmospheric CO₂ concentration and temperature on carbon and nitrogen metabolism in maize at the grain filling stage

College of Agriculture, Shanxi Agricultural University, Taigu 030801, China

Funds: This work was supported by the National Natural Science Foundation of China (31871517), the Program for Selected Young Scientists Studying Abroad of Shanxi Province (20210041) and the Fundamental Research Program of Youth Scientists of Shanxi Province (20210302124656).

More Information

Corresponding author: E-mail: haoxingyu1976@126.com

摘要

摘要: 为探讨C₄作物玉米对CO₂浓度升高、温度升高及其交互作用的响应, 本研究以玉米品种‘先玉335’为材料, 利用人工控制气室设置CK (CO₂浓度为400 μmol∙mol⁻¹, 环境温度)、EC (CO₂浓度为600 μmol∙mol⁻¹, 环境温度)、ET (CO₂浓度为400 μmol∙mol⁻¹, 气温为环境温度+2 ℃)、ECT (CO₂浓度为600 μmol∙mol⁻¹, 气温为环境温度+2 ℃) 4个处理, 测定玉米灌浆期叶片光合生理、糖代谢、氮代谢相关指标, 并在成熟后测定玉米生物量。结果表明: 1) CO₂浓度升高条件下, 玉米叶片叶绿素含量、蔗糖含量、净光合速率及蔗糖合成酶、丙酮酸激酶和α-酮戊二酸脱氢酶活性显著升高(P<0.05), 但谷氨酸合成酶活性显著降低(P<0.05), 地上部生物量和穗重显著升高35.8%和170.2% (P<0.05)。2)气温升高条件下, 叶片净光合速率、蔗糖合成酶和丙酮酸激酶活性显著升高(P<0.05), 但α-酮戊二酸脱氢酶和谷氨酸合成酶活性显著降低(P<0.05), 地上部生物量、叶重、茎重和穗重显著降低37.0%、28.7%、32.3%和62.2% (P<0.05)。3) CO₂浓度和气温均升高条件下, 叶片净光合速率和丙酮酸激酶活性显著升高(P<0.05), 但叶绿素含量、α-酮戊二酸脱氢酶和谷氨酸合成酶活性显著降低(P<0.05), 叶重显著降低23.4% (P<0.05)。总之, CO₂浓度升高可通过促进玉米叶片光合速率, 增加糖代谢相关酶活性和光合代谢产物等缓解温度升高对玉米生物量的负效应; CO₂浓度升高、气温升高以及二者互作下玉米氮代谢受到抑制, 玉米叶片受到氮素胁迫, 或对玉米品质产生不利影响。
- CO₂浓度升高 /
- 气温升高 /
- 玉米 /
- 碳氮代谢 /
- 生物量
Abstract: Future climate change will bring considerable challenges to agricultural production and food security. Presently, research on the effects of elevated CO₂ concentration and increased temperature on crops is mostly focused on C₃ crops, while research on C₄ crops is rare. Maize is the most widely planted C₄ crop in the world, it is of great significance to explore the response of maize to elevated CO₂ concentration, increased temperature, and their combination to assess the impacts of future climate change on C₄ crops. The maize variety ‘Xianyu-335’ was used. Four treatments were set up in controlled chambers: CK (CO₂ concentration 400 μmol·mol⁻¹, ambient temperature), EC (CO₂ concentration 600 μmol·mol⁻¹, ambient temperature), ET (CO₂ concentration 400 μmol·mol⁻¹, 2 ℃ higher than ambient temperature), and ECT (CO₂ concentration 600 μmol·mol⁻¹, 2 ℃ higher than ambient temperature). The related indices of photosynthetic physiology, glucose metabolism, and nitrogen metabolism of maize leaves were measured at the grain-filling stage, and the biomass of maize was measured after ripening. The results showed that: 1) under elevated CO₂ concentrations, the chlorophyll content, sucrose content, net photosynthetic rate, sucrose synthase activity, pyruvate kinase activity, and α-ketoglutarate dehydrogenase activity in leaves were significantly increased (P<0.05), while glutamate synthase activity was significantly decreased (P<0.05). Additionally, aboveground biomass and spike mass were significantly increased by 35.8% and 170.2%, respectively (P<0.05). 2) At increased temperatures, the net photosynthetic rate, and activities of sucrose synthase and pyruvate kinase of leaves were significantly increased (P<0.05), while α-ketoglutarate dehydrogenase and glutamate synthase activities were significantly decreased (P<0.05), and the above-ground biomass and the biomasses of leaf, stem, and spike were significantly decreased by 37.0%, 28.7%, 32.3%, and 62.2%, respectively (P<0.05). 3) Under the combination of elevated CO₂ concentration and increased temperature, the net photosynthetic rate and pyruvate kinase activity of leaves were significantly increased (P<0.05), whereas the chlorophyll content, and activities of α-ketoglutarate dehydrogenase and glutamate synthase were significantly decreased (P<0.05), and the leaf biomass was significantly decreased by 23.4% (P<0.05). In conclusion, elevated CO₂ concentration could alleviate the negative impact of increased temperatures on maize biomass by increasing photosynthesis and the activity of enzymes related to glucose metabolism and photosynthetic metabolites. Under elevated CO₂, increased temperature, or their combination, nitrogen metabolism in maize is inhibited; thus, leaves are subjected to nitrogen stress, which harms maize quality.
- Elevated CO₂ concentration /
- Increased temperature /
- Maize /
- Metabolism of carbon and nitrogen /
- Biomass

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图 1 玉米生长季各处理的CO₂浓度变化

CK: CO₂浓度400 μmol∙mol⁻¹, 环境温度; EC: CO₂浓度600 μmol∙mol⁻¹, 环境温度; ET: CO₂浓度400 μmol∙mol⁻¹, 环境温度+2 ℃; ECT: CO₂浓度600 μmol∙mol⁻¹, 环境温度+2 ℃。CK: CO₂ concentration 400 μmol∙mol⁻¹, ambient temperature; EC: CO₂ concentration 600 μmol∙mol⁻¹, ambient temperature; ET: CO₂ concentration 400 μmol∙mol⁻¹, 2 ℃ higher than ambient temperature; ECT: CO₂ concentration 600 μmol∙mol⁻¹, 2 ℃ higher than ambient temperature.

Figure 1. CO₂ concentrations of different treatments in maize growing season

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图 2 玉米生长季各处理气温变化

Figure 2. Temperature of different treatments in maize growing season

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图 3 玉米地上部生物量与碳氮代谢间皮尔逊相关性

*表示显著相关(P<0.05)。* means significant correlation (P<0.05).

Figure 3. Pearson correlation analysis between above-ground biomass and carbon and nitrogen metabolism in maize

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表 1 CO₂浓度和气温升高对玉米灌浆期叶片光合色素含量的影响

Table 1. Effects of elevated CO₂ concentration and increased temperature on photosynthetic pigment content of maize leaves at grain filling stage

处理 Treatment	叶绿素a Chlorophyll a	叶绿素b Chlorophyll b	类胡萝卜素 Carotenoids	总叶绿素(a+b) Chlorophyll
mg∙g⁻¹(FW)
CK	2.10±0.06b	0.15±0.01b	0.46±0.01a	2.24±0.10b
EC	2.38±0.02a	0.21±0.01a	0.48±0.01a	2.59±0.02a
ET	2.24±0.08ab	0.19±0.01a	0.45±0.02a	2.43±0.09ab
ECT	1.64±0.11c	0.12±0.01b	0.35±0.02b	1.77±0.12c
P_CO²	0.07	0.65	0.04	0.09
P_T	0.00	0.03	0.00	0.01
P_CO²_×T	0.00	0.00	0.01	0.00
CK: CO₂浓度400 μmol∙mol⁻¹, 环境温度; EC: CO₂浓度600 μmol∙mol⁻¹, 环境温度; ET: CO₂浓度400 μmol∙mol⁻¹, 环境温度+2 ℃; ECT: CO₂浓度600 μmol∙mol⁻¹, 环境温度+2 ℃。CO₂: CO₂浓度; T: 温度。同列不同小写字母表示处理间差异显著(P<0.05)。CK: CO₂ concentration 400 μmol∙mol⁻¹, ambient temperature; EC: CO₂ concentration 600 μmol∙mol⁻¹, ambient temperature; ET: CO₂ concentration 400 μmol∙mol⁻¹, 2 ℃ higher than ambient temperature; ECT: CO₂ concentration 600 μmol∙mol⁻¹, 2 ℃ higher than ambient temperature. Different lowercase letters in the same column mean significant differences among treatments at P<0.05 level.

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表 2 CO₂浓度和气温升高对玉米灌浆期叶片光合参数的影响

Table 2. Effects of elevated CO₂ concentration and increased temperature on photosynthetic parameters of maize leaves at grain filling stage

处理 Treatment	净光合速率 Net photosynthetic rate (μmol∙m⁻²∙s⁻¹)	气孔导度 Stomatal conductance [mol(H₂O)∙m⁻²∙s⁻¹]	蒸腾速率 Transpiration rate [mmol(H₂O)∙m⁻²∙s⁻¹]	水分利用效率 Water use efficiency [μmol(CO₂)∙mmol⁻¹(H₂O)]
CK	23.29±0.61c	0.12±0.01c	3.14±0.06d	7.42±0.06b
EC	33.32±0.45a	0.13±0.01b	3.35±0.08c	9.94±0.20a
ET	34.55±0.48a	0.14±0.01b	3.56±0.04b	9.70±0.11a
ECT	26.65±0.46b	0.15±0.01a	3.80±0.04a	7.01±0.06c
P_CO²	0.04	0.00	0.00	0.55
P_T	0.00	0.00	0.00	0.01
P_CO²×T	0.00	0.56	0.78	0.00
CK: CO₂浓度400 μmol∙mol⁻¹, 环境温度; EC: CO₂浓度600 μmol∙mol⁻¹, 环境温度; ET: CO₂浓度400 μmol∙mol⁻¹, 环境温度+2 ℃; ECT: CO₂浓度600 μmol∙mol⁻¹, 环境温度+2 ℃。CO₂: CO₂浓度; T: 温度。同列不同小写字母表示处理间差异显著(P<0.05)。CK: CO₂ concentration 400 μmol∙mol⁻¹, ambient temperature; EC: CO₂ concentration 600 μmol∙mol⁻¹, ambient temperature; ET: CO₂ concentration 400 μmol∙mol⁻¹, 2 ℃ higher than ambient temperature; ECT: CO₂ concentration 600 μmol∙mol⁻¹, 2 ℃ higher than ambient temperature. Different lowercase letters in the same column mean significant differences among treatments at P<0.05 level.

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表 3 CO₂浓度和气温升高对玉米灌浆期叶片蔗糖和淀粉含量、蔗糖合成酶和蔗糖磷酸合成酶活性的影响

Table 3. Effects of elevated CO₂ concentration and increased temperature on the contents of sucrose and starch, the activities of sucrose synthase and sucrose phosphate synthase in maize leaves at grain filling stage

处理 Treatment	蔗糖含量 Sucrose content [mg∙g⁻¹(FW)]	蔗糖合成酶活性 Sucrose synthase activity [mg∙g⁻¹(FW)∙h⁻¹]	蔗糖磷酸合成酶活性 Sucrose phosphate synthase activity [mg∙g⁻¹(FW)∙h⁻¹]	淀粉含量 Starch content [mg∙g⁻¹(FW)]
CK	0.05±0.01b	227.84±7.90b	341.00±61.16a	0.42±0.06ab
EC	0.27±0.03a	328.01±15.25a	342.69±6.25a	0.32±0.05b
ET	0.01±0.01b	322.64±13.04a	347.45±1.79a	0.53±0.07a
ECT	0.04±0.02b	242.61±27.74b	342.24±4.67a	0.33±0.02b
P_CO²	0.00	0.58	0.91	0.04
P_T	0.00	0.80	0.93	0.27
P_CO²×T	0.00	0.00	0.96	0.50
CK: CO₂浓度400 μmol∙mol⁻¹, 环境温度; EC: CO₂浓度600 μmol∙mol⁻¹, 环境温度; ET: CO₂浓度400 μmol∙mol⁻¹, 环境温度+2 ℃; ECT: CO₂浓度600 μmol∙mol⁻¹, 环境温度+2 ℃。CO₂: CO₂浓度; T: 温度。同列不同小写字母表示处理间差异显著(P<0.05)。CK: CO₂ concentration 400 μmol∙mol⁻¹, ambient temperature; EC: CO₂ concentration 600 μmol∙mol⁻¹, ambient temperature; ET: CO₂ concentration 400 μmol∙mol⁻¹, 2 ℃ higher than ambient temperature; ECT: CO₂ concentration 600 μmol∙mol⁻¹, 2 ℃ higher than ambient temperature. Different lowercase letters in the same column mean significant differences among treatments at P<0.05 level.

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表 4 CO₂浓度和气温升高对玉米灌浆期叶片丙酮酸激酶、α-酮戊二酸脱氢酶和琥珀酸脱氢酶活性的影响

Table 4. Effects of elevated CO₂ concentration and increased temperature on the activities of pyruvate kinase, α-ketoglutarate dehydrogenase and succinate dehydrogenase in maize leaves at grain filling stage

处理 Treatment	丙酮酸激酶活性 Pyruvate kinase activity	α-酮戊二酸脱氢酶活性 α-Ketoglutarate dehydrogenase activity	琥珀酸脱氢酶活性 Succinate dehydrogenase activity
U·g⁻¹(FW)
CK	260.10±20.91d	230.97±1.68b	123.41±36.08a
EC	694.74±1.47a	305.32±1.81a	51.12±17.47a
ET	504.39±10.92b	55.18±23.53c	119.01±72.04a
ECT	333.38±2.44c	47.76±3.98c	66.29±13.08a
P_CO²	0.00	0.02	0.12
P_T	0.00	0.00	0.88
P_CO²×T	0.00	0.01	0.79
CK: CO₂浓度400 μmol∙mol⁻¹, 环境温度; EC: CO₂浓度600 μmol∙mol⁻¹, 环境温度; ET: CO₂浓度400 μmol∙mol⁻¹, 环境温度+2 ℃; ECT: CO₂浓度600 μmol∙mol⁻¹, 环境温度+2 ℃。CO₂: CO₂浓度; T: 温度。同列不同小写字母表示处理间差异显著(P<0.05)。CK: CO₂ concentration 400 μmol∙mol⁻¹, ambient temperature; EC: CO₂ concentration 600 μmol∙mol⁻¹, ambient temperature; ET: CO₂ concentration 400 μmol∙mol⁻¹, 2 ℃ higher than ambient temperature; ECT: CO₂ concentration 600 μmol∙mol⁻¹, 2 ℃ higher than ambient temperature. Different lowercase letters mean significant differences among treatments at P<0.05 level.

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表 5 CO₂浓度和气温升高对玉米灌浆期叶片硝酸还原酶、谷氨酰胺合成酶和谷氨酸合成酶活性的影响

Table 5. Effects of elevated CO₂ concentration and increased temperature on the activities of nitrate reductase, glutamine synthase and glutamate synthase in maize leaves at grain filling stage

处理 Treatment	硝酸还原酶活性 Nitrate reductase activity [μg(NO₂)·g⁻¹(FW)·h⁻¹]	谷氨酰胺合成酶活性 Glutamine synthetase activity [U·g⁻¹(FW)]	谷氨酸合成酶活性 Glutamate synthetase activity [U·g⁻¹(FW)]
CK	3.36±0.85a	3.85±0.16a	248.05±67.93a
EC	1.45±0.30a	7.30±0.85a	65.17±32.23b
ET	9.42±5.09a	4.12±1.43a	83.32±23.39b
ECT	9.35±3.71a	6.67±2.15a	71.91±12.09b
P_CO²	0.76	0.06	0.04
P_T	0.06	0.90	0.08
P_CO²×T	0.78	0.75	0.06
CK: CO₂浓度400 μmol∙mol⁻¹, 环境温度; EC: CO₂浓度600 μmol∙mol⁻¹, 环境温度; ET: CO₂浓度400 μmol∙mol⁻¹, 环境温度+2 ℃; ECT: CO₂浓度600 μmol∙mol⁻¹, 环境温度+2 ℃。CO₂: CO₂浓度; T: 温度。同列不同小写字母表示处理间差异显著(P<0.05)。CK: CO₂ concentration 400 μmol∙mol⁻¹, ambient temperature; EC: CO₂ concentration 600 μmol∙mol⁻¹, ambient temperature; ET: CO₂ concentration 400 μmol∙mol⁻¹, 2 ℃ higher than ambient temperature; ECT: CO₂ concentration 600 μmol∙mol⁻¹, 2 ℃ higher than ambient temperature. Different lowercase letters mean significant differences among treatments at P<0.05 level.

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表 6 CO₂浓度和气温升高对玉米成熟期形态指标的影响

Table 6. Effects of elevated CO₂ concentration and increased temperature on morphological index of maize at maturity stage

处理 Treatment	株高 Plant height (cm)	茎粗 Stem-diameter (mm)	节数 Internodes number	穗位节 Ear node
CK	206.97±11.46a	7.87±0.79b	12.00±0.00b	7.00±0.00a
EC	222.50±5.77a	11.17±0.36a	13.33±0.33a	7.33±0.33a
ET	161.17±15.31b	7.71±0.57b	11.33±0.67b	4.33±0.33b
ECT	225.67±0.83a	7.94±0.72b	13.67±0.33a	7.33±0.33a
P_CO²	0.00	0.02	0.00	0.00
P_T	0.07	0.03	0.69	0.00
P_CO²×T	0.04	0.04	0.26	0.00
CK: CO₂浓度400 μmol∙mol⁻¹, 环境温度; EC: CO₂浓度600 μmol∙mol⁻¹, 环境温度; ET: CO₂浓度400 μmol∙mol⁻¹, 环境温度+2 ℃; ECT: CO₂浓度600 μmol∙mol⁻¹, 环境温度+2 ℃。CO₂: CO₂浓度; T: 温度。同列不同小写字母表示处理间差异显著(P<0.05)。CK: CO₂ concentration 400 μmol∙mol⁻¹, ambient temperature; EC: CO₂ concentration 600 μmol∙mol⁻¹, ambient temperature; ET: CO₂ concentration 400 μmol∙mol⁻¹, 2 ℃ higher than ambient temperature; ECT: CO₂ concentration 600 μmol∙mol⁻¹, 2 ℃ higher than ambient temperature. Different lowercase letters in the same column mean significant differences among treatments at P<0.05 level.

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表 7 CO₂浓度和气温升高对玉米成熟期生物量的影响

Table 7. Effects of elevated CO₂ concentration and increased temperature on biomass of maize at maturity stage

处理 Treatment	地上部生物量 Above-ground biomass	叶干重 Leaf dry mass	茎干重 Stem dry mass	穗干重 Spike dry mass
g∙plant⁻¹
CK: CO₂浓度400 μmol∙mol⁻¹, 环境温度; EC: CO₂浓度600 μmol∙mol⁻¹, 环境温度; ET: CO₂浓度400 μmol∙mol⁻¹, 环境温度+2 ℃; ECT: CO₂浓度600 μmol∙mol⁻¹, 环境温度+2 ℃。CO₂: CO₂浓度; T: 温度。同列不同小写字母表示处理间差异显著(P<0.05)。CK: CO₂ concentration 400 μmol∙mol⁻¹, ambient temperature; EC: CO₂ concentration 600 μmol∙mol⁻¹, ambient temperature; ET: CO₂ concentration 400 μmol∙mol⁻¹, 2 ℃ higher than ambient temperature; ECT: CO₂ concentration 600 μmol∙mol⁻¹, 2 ℃ higher than ambient temperature. Different lowercase letters in the same column mean significant differences among treatments at P<0.05 level.

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