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
计量
- 文章访问数: 449
- HTML全文浏览量: 198
- PDF下载量: 79
- 被引次数: 0
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

HTML全文

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

下载: 全尺寸图片幻灯片

图 2 玉米生长季各处理气温变化

Figure 2. Temperature of different treatments in maize growing season

下载: 全尺寸图片幻灯片

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

下载: 全尺寸图片幻灯片

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.

下载: 导出CSV

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.

下载: 导出CSV

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.

下载: 导出CSV

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.

下载: 导出CSV

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.

下载: 导出CSV

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.

下载: 导出CSV

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.

下载: 导出CSV

参考文献(41)

[1]	IPCC. Summary for Policymakers[M]//MASSON-DELMOTTE V, ZHAI P M, PIRANI A, et al. Climate Change 2021: The Physical Science Basis. Contribution of Working GroupⅠ to the Sixth Assessment Report of the Intergovernmental Panel on Climate Change. Cambridge: Cambridge University Press, 2021: 3−32
[2]	宗毓铮, 张函青, 李萍, 等. 大气CO₂与温度升高对北方冬小麦旗叶光合特性、碳氮代谢及产量的影响[J]. 中国农业科学, 2021, 54(23): 4984−4995 doi: 10.3864/j.issn.0578-1752.2021.23.005 ZONG Y Z, ZHANG H Q, LI P, et al. Effects of elevated atmospheric CO₂ concentration and temperature on photosynthetic characteristics, carbon and nitrogen metabolism in flag leaves and yield of winter wheat in North China[J]. Scientia Agricultura Sinica, 2021, 54(23): 4984−4995 doi: 10.3864/j.issn.0578-1752.2021.23.005
[3]	黄辉, 王春乙, 白月明, 等. O₃与CO₂浓度倍增对大豆叶片及其总生物量的影响研究[J]. 中国生态农业学报, 2005, 13(4): 52−55 HUANG H, WANG C Y, BAI Y M, et al. Impact of O₃ and CO₂ concentration doubling on the soybean leaf development and biomass[J]. Chinese Journal of Eco-Agriculture, 2005, 13(4): 52−55
[4]	任宏芳, 李阿立, 郝兴宇, 等. 大气CO₂浓度和气温升高对谷子生长及产量的影响[J]. 生态环境学报, 2020, 29(6): 1123−1129 REN H F, LI A L, HAO X Y, et al. Effects of elevated atmospheric CO₂ concentration and temperature on the growth and yield of millet[J]. Ecology and Environmental Sciences, 2020, 29(6): 1123−1129
[5]	MYERS S S, ZANOBETTI A, KLOOG I, et al. Increasing CO₂ threatens human nutrition[J]. Nature, 2014, 510(7503): 139−142 doi: 10.1038/nature13179
[6]	ZHU C W, KOBAYASHI K, LOLADZE I, et al. Carbon dioxide (CO₂) levels this century will alter the protein, micronutrients, and vitamin content of rice grains with potential health consequences for the poorest rice-dependent countries[J]. Science Advances, 2018, 4(5): 1012−1012 doi: 10.1126/sciadv.aaq1012
[7]	张小琴, 郝兴宇, 宗毓铮, 等. CO₂浓度升高对大豆糖代谢和脂肪代谢影响的研究[J]. 激光生物学报, 2022, 31(1): 70−78 ZHANG X Q, HAO X Y, ZONG Y Z, et al. Effects of elevated CO₂ concentration on sugar metabolism and fat metabolism in soybean[J]. Acta Laser Biology Sinica, 2022, 31(1): 70−78
[8]	郝兴宇, 李萍, 杨宏斌, 等. 大气CO₂浓度升高对绿豆生长及C、N吸收的影响[J]. 中国生态农业学报, 2011, 19(4): 794−798 doi: 10.3724/SP.J.1011.2011.00794 HAO X Y, LI P, YANG H B, et al. Effects of enriched atmospheric CO₂ on the growth and uptake of N and C in mung bean[J]. Chinese Journal of Eco-Agriculture, 2011, 19(4): 794−798 doi: 10.3724/SP.J.1011.2011.00794
[9]	WANG Y L, ZHANG Y K, SHI Q H, et al. Decrement of sugar consumption in rice young panicle under high temperature aggravates spikelet number reduction[J]. Rice Science, 2020, 27(1): 44−55 doi: 10.1016/j.rsci.2019.12.005
[10]	HU Q, WEISS A, FENG S, et al. Earlier winter wheat heading dates and warmer spring in the US Great Plains[J]. Agricultural and Forest Meteorology, 2005, 135(1/2/3/4): 284−290
[11]	TAO F L, ZHANG S, ZHANG Z, et al. Maize growing duration was prolonged across China in the past three decades under the combined effects of temperature, agronomic management, and cultivar shift[J]. Global Change Biology, 2014, 20(12): 3686−3699 doi: 10.1111/gcb.12684
[12]	刘亮, 郝立华, 李菲, 等. CO₂浓度和温度对玉米光合性能及水分利用效率的影响[J]. 农业工程学报, 2020, 36(5): 122−129 doi: 10.11975/j.issn.1002-6819.2020.05.014 LIU L, HAO L H, LI F, et al. Effects of CO₂ concentration and temperature on leaf photosynthesis and water use efficiency in maize[J]. Transactions of the Chinese Society of Agricultural Engineering, 2020, 36(5): 122−129 doi: 10.11975/j.issn.1002-6819.2020.05.014
[13]	张秀云, 姚玉璧, 雷俊, 等. CO₂浓度升高与增温对马铃薯产量及品质的复合影响[J]. 干旱地区农业研究, 2019, 37(4): 240−246 ZHANG X Y, YAO Y B, LEI J, et al. Collaborative impact of elevated CO₂ concentration and temperature on potato yield and quality[J]. Agricultural Research in the Arid Areas, 2019, 37(4): 240−246
[14]	陈楠楠, 周超, 王浩成, 等. 大气二氧化碳含量升高对稻麦产量影响的整合分析[J]. 南京农业大学学报, 2013, 36(2): 83−90 doi: 10.7685/j.issn.1000-2030.2013.02.014 CHEN N N, ZHOU C, WANG H C, et al. Impacts of elevated atmospheric CO₂ on growth and yield of rice and wheat: a Meta-analysis[J]. Journal of Nanjing Agricultural University, 2013, 36(2): 83−90 doi: 10.7685/j.issn.1000-2030.2013.02.014
[15]	杨小倩, 郅慧, 张辉, 等. 玉米不同部位化学成分、药理作用、利用现状研究进展[J]. 吉林中医药, 2019, 39(6): 837−840 YANG X Q, ZHI H, ZHANG H, et al. Research progress on chemical constituents, pharmacological activity and utilization status of different parts of corn[J]. Jilin Journal of Chinese Medicine, 2019, 39(6): 837−840
[16]	李明, 李迎春, 牛晓光, 等. 大气CO₂浓度升高与氮肥互作对玉米花后碳氮代谢及产量的影响[J]. 中国农业科学, 2021, 54(17): 3647−3665 doi: 10.3864/j.issn.0578-1752.2021.17.008 LI M, LI Y C, NIU X G, et al. Effects of elevated atmospheric CO₂ concentration and nitrogen fertilizer on the yield of summer maize and carbon and nitrogen metabolism after flowering[J]. Scientia Agricultura Sinica, 2021, 54(17): 3647−3665 doi: 10.3864/j.issn.0578-1752.2021.17.008
[17]	HUANG Y L, FANG R, LI Y S, et al. Warming and elevated CO₂ alter the transcriptomic response of maize (Zea mays L.) at the silking stage[J]. Scientific Reports, 2019, 9: 17948 doi: 10.1038/s41598-019-54325-5
[18]	高俊凤. 植物生理学实验指导[M]. 北京: 高等教育出版社, 2006 GAO J F. Experimental Guidance for Plant Physiology[M]. Beijing: Higher Education Press, 2006
[19]	郭茜茜. 大豆子粒蛋白质积累与碳代谢关系的研究[D]. 哈尔滨: 东北农业大学, 2010 GUO X X. Research on the relationship between protein accumulation and carbon metabolism in soybean grain[D]. Harbin: Northeast Agricultural University, 2010
[20]	BHATT R, BAIG M J, TIWARI H S, et al. Growth, yield and photosynthesis of Panicum maximum and Stylosanthes hamata under elevated CO₂[J]. Journal of Environmental Biology, 2010, 31: 549−52
[21]	金奖铁, 李扬, 李荣俊, 等. 大气二氧化碳浓度升高影响植物生长发育的研究进展[J]. 植物生理学报, 2019, 55(5): 558−568 doi: 10.13592/j.cnki.ppj.2018.0533 JIN J T, LI Y, LI R J, et al. Advances in studies on effects of elevated atmospheric carbon dioxide concentration on plant growth and development[J]. Plant Physiology Journal, 2019, 55(5): 558−568 doi: 10.13592/j.cnki.ppj.2018.0533
[22]	吴雪霞, 张圣美, 张爱冬, 等. 外源褪黑素对高温胁迫下茄子幼苗光合和生理特性的影响[J]. 植物生理学报, 2019, 55(1): 49−60 WU X X, ZHANG S M, ZHANG A D, et al. Effect of exogenous melatonin on photosynthetic and physiological characteristics of eggplant seedlings under high temperature stress[J]. Plant Physiology Journal, 2019, 55(1): 49−60
[23]	刘紫娟, 李萍, 宗毓铮, 等. 大气CO₂浓度升高对谷子生长发育及玉米螟发生的影响[J]. 中国生态农业学报, 2017, 25(1): 55−60 LIU Z J, LI P, ZONG Y Z, et al. Effect of elevated [CO₂] on growth and attack of Asian corn borers (Ostrinia furnacalis) in foxtail millet (Setaria italica)[J]. Chinese Journal of Eco-Agriculture, 2017, 25(1): 55−60
[24]	赵天宏, 黄国宏. 大气二氧化碳浓度升高对植物影响的研究进展[J]. 作物杂志, 2003(3): 3−6 ZHAO T H, HUANG G H. Research Progress on the effects of elevated atmospheric carbon dioxide concentration on plants[J]. Crops, 2003(3): 3−6
[25]	AINSWORTH E, ROGERS A. The response of photosynthesis and stomatal conductance to rising [CO₂]: mechanisms and environmental interactions[J]. Plant, Cell & Environment, 2007, 30(3): 258−270
[26]	CURTIS P S, WANG X Z. A meta-analysis of elevated CO₂ effects on woody plant mass, form, and physiology[J]. Oecologia, 1998, 113(3): 299−313 doi: 10.1007/s004420050381
[27]	廖轶, 陈根云, 张海波, 等. 水稻叶片光合作用对开放式空气CO₂浓度增高(FACE)的响应与适应[J]. 应用生态学报, 2002, 13(10): 1205−1209 doi: 10.3321/j.issn:1001-9332.2002.10.002 LIAO Y, CHEN G Y, ZHANG H B, et al. Response and acclimation of photosynthesis in rice leaves to free-air CO₂ enrichment (FACE)[J]. Chinese Journal of Applied Ecology, 2002, 13(10): 1205−1209 doi: 10.3321/j.issn:1001-9332.2002.10.002
[28]	谢辉, 范桂枝, 荆彦辉, 等. 植物对大气CO₂浓度升高的光合适应机理研究进展[J]. 中国农业科技导报, 2006, 8(3): 29−34 XIE H, FAN G Z, JING Y H, et al. Progress of research on photosynthetic acclimation of plant to elevated atmospheric CO₂[J]. Review of China Agricultural Science and Technology, 2006, 8(3): 29−34
[29]	胡晓雪, 杜维俊, 杨珍平, 等. 大气CO₂浓度和气温升高对野生大豆光合作用的影响[J]. 山西农业科学, 2015, 43(7): 798−801, 853 HU X X, DU W J, YANG Z P, et al. Effect of elevated CO₂ concentration and increased temperature on the photosynthesis of wild soybean[J]. Journal of Shanxi Agricultural Sciences, 2015, 43(7): 798−801, 853
[30]	赵娜, 孙艳, 王德玉, 等. 外源褪黑素对高温胁迫条件下黄瓜幼苗氮代谢的影响[J]. 植物生理学报, 2012, 48(6): 557−564 doi: 10.13592/j.cnki.ppj.2012.06.007 ZHAO N, SUN Y, WANG D Y, et al. Effects of exogenous melatonin on nitrogen metabolism in cucumber seedlings under high temperature stress[J]. Plant Physiology Journal, 2012, 48(6): 557−564 doi: 10.13592/j.cnki.ppj.2012.06.007
[31]	LI S H, LI Y M, GAO Y, et al. Effects of CO₂ enrichment on non-structural carbohydrate metabolism in leaves of cucumber seedlings under salt stress[J]. Scientia Horticulturae, 2020, 265: 109275 doi: 10.1016/j.scienta.2020.109275
[32]	曹新超. 铁皮石斛蔗糖代谢及抗氧化酶对逆境胁迫的响应[D]. 杭州: 浙江农林大学, 2018 CAO X C. Responses of sucrose metabolism and antioxidant enzymes of Dendrobium catenatum to stress[D]. Hangzhou: Zhejiang A & F University, 2018
[33]	王军可, 王亚梁, 陈惠哲, 等. 灌浆初期高温影响水稻籽粒碳氮代谢的机理[J]. 中国农业气象, 2020, 41(12): 774−784 doi: 10.3969/j.issn.1000-6362.2020.12.003 WANG J K, WANG Y L, CHEN H Z, et al. Mechanism of high temperature affecting carbon and nitrogen metabolism of rice grain at the early stage of grain filling[J]. Chinese Journal of Agrometeorology, 2020, 41(12): 774−784 doi: 10.3969/j.issn.1000-6362.2020.12.003
[34]	刘昭霖, 宗毓铮, 张东升, 等. 大气CO₂浓度和气温升高对大豆叶片光合特性及氮代谢的影响[J]. 中国农业气象, 2021, 42(5): 426−437 doi: 10.3969/j.issn.1000-6362.2021.05.007 LIU Z L, ZONG Y Z, ZHANG D S, et al. Effects of elevated atmospheric CO₂ concentration and increased air temperature on photosynthetic characteristics and nitrogen metabolism of soybean leaves[J]. Chinese Journal of Agrometeorology, 2021, 42(5): 426−437 doi: 10.3969/j.issn.1000-6362.2021.05.007
[35]	LI Y S, YU Z H, LIU X B, et al. Elevated CO₂ increases nitrogen fixation at the reproductive phase contributing to various yield responses of soybean cultivars[J]. Frontiers in Plant Science, 2017, 8: 1546 doi: 10.3389/fpls.2017.01546
[36]	李春华, 曾青, 沙霖楠, 等. 大气CO₂浓度和温度升高对水稻地上部干物质积累和分配的影响[J]. 生态环境学报, 2016, 25(8): 1336−1342 LI C H, ZENG Q, SHA L N, et al. Impacts of elevated atmospheric CO₂ and temperature on above-ground dry matter accumulation and distribution of rice (Oryza sativa L.)[J]. Ecology and Environmental Sciences, 2016, 25(8): 1336−1342
[37]	SCHIERMEIER Q. Climate change offers bleak future[J]. Nature, 2001, 409(6823): 971
[38]	WANG J Q, LIU X Y, ZHANG X H, et al. Size and variability of crop productivity both impacted by CO₂ enrichment and warming — A case study of 4 years field experiment in a Chinese paddy[J]. Agriculture, Ecosystems & Environment, 2016, 221: 40−49
[39]	MOHAMMED A R, TARPLEY L. High nighttime temperatures affect rice productivity through altered pollen germination and spikelet fertility[J]. Agricultural and Forest Meteorology, 2009, 149(6/7): 999−1008
[40]	宋磊, 次仁央金, 王小强, 等. 小麦对高温胁迫的响应机制研究进展[J]. 中国农学通报, 2021, 37(36): 6−12 doi: 10.11924/j.issn.1000-6850.casb2021-0681 SONG L, CIRENYANGJIN, WANG X Q, et al. Response mechanism of wheat to high temperature stress: a review[J]. Chinese Agricultural Science Bulletin, 2021, 37(36): 6−12 doi: 10.11924/j.issn.1000-6850.casb2021-0681
[41]	伍卫, 李永明, 普丽花, 等. 高温多雨寡照对西双版纳州玉米的影响[J]. 现代化农业, 2022(2): 20−22 doi: 10.3969/j.issn.1001-0254.2022.02.007 WU W, LI Y M, PU L H, et al. Effects of high temperature and rainfall on maize in Xishuangbanna Prefecture[J]. Modernizing Agriculture, 2022(2): 20−22 doi: 10.3969/j.issn.1001-0254.2022.02.007