大气CO<sub>2</sub>浓度升高与氮肥互作对玉米光合特性及产量的影响

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牛晓光^1,,
杨荣全¹,
李明¹,
段碧华²,
刁田田¹,
马芬¹,
郭李萍^1,,
1.中国农业科学院农业环境与可持续发展研究所北京 100081
2.北京农学院生物与资源环境学院北京 102206
基金项目: “十三五”国家重点研发计划专项课题2017YFD0300301

详细信息

作者简介:牛晓光, 主要从事农业气象方面的研究。E-mail:nxg18811072358@163.com

通讯作者:郭李萍, 主要研究方向为作物与环境交互作用。E-mail:guoliping@caas.cn

中图分类号:S162.5

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出版历程

收稿日期:2019-09-18
录用日期:2019-11-06
刊出日期:2020-02-01

Effects of interaction between elevated atmospheric CO₂ concentration and nitrogen fertilizer on photosynthetic characteristic and yield of maize

NIU Xiaoguang^1,,
YANG Rongquan¹,
LI Ming¹,
DUAN Bihua²,
DIAO Tiantian¹,
MA Fen¹,
GUO Liping^1,,
1. Institute of Environment and Sustainable Development in Agriculture, Chinese Academy of Agricultural Sciences, Beijing 100081, China
2. College of Bioscience and Resource Environment, Beijing University of Agriculture, Beijing 102206, China
Funds: the National Key Research and Development Program of China2017YFD0300301

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Corresponding author:GUO Liping, E-mail:guoliping@caas.cn

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摘要
摘要:为阐明大气CO₂浓度升高和氮肥交互作用对C₄作物玉米光合生理和产量的影响，本研究利用自由大气CO₂富集（FACE）平台，以玉米品种‘郑单958’为试验材料，在不同施氮量[常氮180 kg（N）·hm^-2、低氮72 kg（N）·hm^-2]下比较大气CO₂浓度[（400±15）μmol·mol^-1]和高CO₂浓度[（550±15）μmol·mol^-1]对玉米生长的影响。结果表明：1）大气CO₂浓度升高使玉米苗期叶片叶绿素浓度显著（P=0.025）增加9.5%，抽雄期净光合速率显著（P=0.009）增加9.0%；低氮和常氮下，高CO₂浓度使玉米各主要生育期胞间CO₂浓度分别显著增加34.8%~48.5%和40.0%~49.4%，气孔导度在大口期和抽雄期分别显著下降21.6%（P=0.015）和22.1%（P=0.010），玉米叶片水分利用效率在大口期、抽雄期和灌浆期分别显著增加12.9%（P=0.002）、9.8%（P=0.019）和18.8%（P=0.001）；高CO₂浓度使玉米非光化学淬灭呈降低趋势、PSII有效光化学量子产量有增加趋势；相同氮水平下，高CO₂浓度对玉米产量没有显著影响。2）高CO₂浓度和合理施氮交互作用对玉米功能叶叶绿素含量、净光合速率、PSⅡ有效光化学量子产量增加有一定的促进作用，如在大口期和抽雄期，常氮+高CO₂浓度处理叶绿素含量比低氮+大气CO₂浓度处理增加17.3%和10.7%，高CO₂浓度和合理施氮量交互作用有增加玉米产量的潜力，合理增加施氮量促进了CO₂肥效的发挥。在未来大气CO₂浓度升高条件下合理施氮对C₄作物玉米生长发育有促进作用。
Abstract:Since the industrial revolution, the concentration of atmospheric CO₂ has increased from 280 μmol·mol^-1 to 400 μmol·mol^-1. Nitrogen is a necessary element for many important enzyme-mediated processes in plant growth and is the primary nutrient needed for plant growth. Among different C₄ crops grown worldwide, including China, maize is the most widely planted crop. Clear answers regarding the effect of elevated atmospheric CO₂ concentration (eCO₂) on corn growth and the interaction between eCO₂ and nitrogen fertilizers (N) are yet not to be attained. Studying the impact of eCO₂ on maize growth under different nitrogen supply conditions is important to assess the role of climate change in the C₄ crop growth. A Free Air CO₂ Enrichment (FACE) facility was used in this experiment. The FACE facility has six octagon loops for eCO₂, (550±15) μmol·mol^-1, and six additional octagon loops for ambient CO₂ concentration of (400±15) μmol·mol^-1 (aCO₂); three of which are eCO₂ experimental loops and the other three are aCO₂ experimental loops applied with conventional nitrogen fertilizer, 180 kg(N)·hm^-2 (CN). The rest are low nitrogen, 72 kg(N)·hm^-2 (LN), application treatments. Twelve experimental loops were arranged randomly in the maize field, with the plants spacing of 25 cm and a rows spacing of 60 cm. Results showed that under eCO₂, the chlorophyll concentration of maize seedling leaves increased significantly by 9.5%, and the net photosynthetic rate increased by 9.0% at the tasseling stage. During the maize growth period, eCO₂ significantly enhanced the intercellular CO₂ concentration by 34.8%-48.5% and 40.0%-49.4% under low nitrogen and conventional nitrogen application conditions, respectively. In addition, the stomatal conductance decreased by 21.6% and 22.1% at the 12-leaf and the tasseling stages, respectively. As a consequence of decreased stomatal conductance, the efficiency of water consumption in maize leaves increased by 12.9%, 9.8%, and 18.8% at the 12-leaf stage, tasseling stage and filling stage, respectively. eCO₂ also decreased Non-Photochemical Quenching (NPQ), and increased PSⅡ effective photochemical quantum yield (F_v'/F_m') value. At the same nitrogen fertilizer (N) level, eCO₂ had no significant effect on the maize yield. Secondly, the interaction of eCO₂ and a reasonable increase of N application rate promoted the chlorophyll content, net photosynthetic rate, and F_v'/F_m' of maize functional leaves. For instance, the chlorophyll content of functional leaves for CN-eCO₂ against LN-aCO₂ increased by 17.3% and 10.7%, respectively, at the 12-leaf and tasseling stages. The combination of eCO₂ and a reasonable increase in the N application achieved the maximized maize yield, indicating the promotional effect of N application under the eCO₂ conditions. Appropriate application of nitrogen fertilizer has the potential to promote the growth and development of maize crop under eCO₂ conditions in future.

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图12018年玉米生育期间
Figure1.Temperature and precipitation during maize growth period (June-September) in 2018

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图2不同CO₂浓度和氮肥用量处理下玉米主要生育期的功能叶叶绿素含量
LN、CN分别指低氮用量和常规氮用量处理, aCO₂和eCO₂分别指常规浓度CO₂处理和高浓度CO₂处理。V6:六叶期; V12:大口期; VT:抽雄期; R3:灌浆期。不同小写字母表示同一生育期不同处理间差异在P < 0.05水平上显著。ns表示不显著, *和**分别表示同一生育期CO₂、氮肥及其交互作用在P < 0.05和P < 0.01水平显著。
Figure2.Concentrations of chlorophyll in functional leaves of maize under different treatments of CO₂ concentration and nitrogen fertilizer application during different main growth stages
LN and CN mean the treatments of low nitrogen and conventional nitrogen application. aCO₂ and eCO₂ mean the treatments of ambient CO₂ concentration and elevated CO₂ concentration. V6: 6-leaf stage; V12: 12-leaf stage; VT: tasseling stage; R3: filling stage. Different lowercase letters indicate significant differences among treatments during the same growth stage (P < 0.05). "ns" indicates no significant effect, * and ** indicate significant effects of CO₂, nitrogen fertilizer and their interaction during the same growth stage at P < 0.05 and P < 0.01, respectively.

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图3不同CO₂浓度和氮肥用量处理下玉米主要生育期的功能叶净光合速率
LN、CN分别指低氮用量和常规氮用量处理, aCO₂和eCO₂分别指常规浓度CO₂处理和高浓度CO₂处理。V6:六叶期; V12:大口期; VT:抽雄期; R3:灌浆期。不同小写字母表示同一生育期不同处理间差异在P < 0.05水平上显著。ns表示不显著, *和**分别表示同一生育期CO₂、氮肥及其交互作用在P < 0.05和P < 0.01水平显著。
Figure3.Net photosynthetic rate of functional leaves of maize under different treatments of CO₂ concentration and nitrogen fertilizer application during different main growth stages
LN and CN mean the treatments of low nitrogen and conventional nitrogen application. aCO₂ and eCO₂ mean the treatments of ambient CO₂ concentration and elevated CO₂ concentration. V6: 6-leaf stage; V12: 12-leaf stage; VT: tasseling stage; R3: filling stage. Different lowercase letters indicate significant differences among treatments during the same growth stage (P < 0.05). "ns" indicates no significant effect, * and ** indicate significant effects of CO₂, nitrogen fertilizer and their interaction during the same growth stage at P < 0.05 and P < 0.01, respectively.

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图4玉米主要生育期不同CO₂浓度和氮肥用量处理功能叶胞间CO₂浓度
LN、CN分别指低氮用量和常规氮用量处理, aCO₂和eCO₂分别指常规浓度CO₂处理和高浓度CO₂处理。V6:六叶期; V12:大口期; VT:抽雄期; R3:灌浆期。不同小写字母表示同一生育期不同处理间差异在P < 0.05水平上显著。ns表示不显著, *和**分别表示同一生育期CO₂、氮肥及其交互作用在P < 0.05和P < 0.01水平显著。
Figure4.Intercellular CO₂ concentrations in functional leaves of maize under different treatments of CO₂ concentration and nitrogen fertilizer application during different main growth stages
LN and CN mean the treatments of low nitrogen and conventional nitrogen application. aCO₂ and eCO₂ mean the treatments of ambient CO₂ concentration and elevated CO₂ concentration. V6: 6-leaf stage; V12: 12-leaf stage; VT: tasseling stage; R3: filling stage. Different lowercase letters indicate significant differences among treatments during the same growth stage (P < 0.05). "ns" indicates no significant effect, * and ** indicate significant effects of CO₂, nitrogen fertilizer and their interaction during the same growth stage at P < 0.05 and P < 0.01, respectively.

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图5不同CO₂浓度和氮肥用量处理下玉米主要生育期功能叶叶片的气孔导度、蒸腾速率和水分利用效率
LN、CN分别指低氮用量和常规氮用量处理, aCO₂和eCO₂分别指常规浓度CO₂处理和高浓度CO₂处理。V6:六叶期; V12:大口期; VT:抽雄期; R3:灌浆期。不同小写字母表示同一生育期不同处理间差异在P < 0.05水平上显著。ns表示不显著, *和**分别表示同一生育期CO₂、氮肥及其交互作用在P < 0.05和P < 0.01水平显著。
Figure5.Stomatal conductance, transpiration rate and water use efficiency of functional leaves of maize under different treatments of CO₂ concentration and nitrogen fertilizer application during different main growth stages
LN and CN mean the treatments of low nitrogen and conventional nitrogen application. aCO₂ and eCO₂ mean the treatments of ambient CO₂ concentration and elevated CO₂ concentration. V6: 6-leaf stage; V12: 12-leaf stage; VT: tasseling stage; R3: filling stage. Different lowercase letters indicate significant differences among treatments during the same growth stage (P < 0.05). "ns" indicates no significant effect, * and ** indicate significant effects of CO₂, nitrogen fertilizer and their interaction during the same growth stage at P < 0.05 and P < 0.01, respectively.

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表12018年夏玉米播前土壤理化性状(0~20 cm)
Table1.Basic soil properties at 0-20 cm layer before sowing the maize in 2018

处理 Treatment	有机质 Organic matter (g·kg^-1)	全氮 Total N (g·kg^-1)	碱解氮 Alkali-hydrolyzable N (mg·kg^-1)	速效磷 Olsen-P (mg·kg^-1)	速效钾 Available K (mg·kg^-1)	pH
常规CO₂浓度 Ambient CO₂ concentration	28.1±1.2a	1.55±0.07a	105.5±2.9a	38.2±1.9a	158.6±7.8a	8.39±0.02b
高CO₂浓度 Elevated CO₂ concentration	30.7±1.9a	1.65±0.09a	118.0±5.1a	41.1±1.8a	155.6±5.4a	8.50±0.03a
同列不同小写字母表示两个处理间差异在P < 0.05水平显著。Different lowercase letters in a column indicate significant differences between two treatments at 0.05 level.

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表2不同CO₂浓度和氮肥用量处理对玉米主要生育期功能叶叶绿素荧光参数的影响
Table2.Major chlorophyll fluorescence parameters of functional leaves of maize under different treatments of CO₂ concentration and nitrogen fertilizer application during main growth stages

生育期 Growth stage	处理 Treatment		F_v'/F_m'	Qp	NPQ
六叶期 Six-leaf stage (V6)	低氮用量 Low nitrogen	常规CO₂浓度Ambient CO₂ concentration	0.46±0.04a	0.75±0.03a	0.63±0.09a
	低氮用量 Low nitrogen	高CO₂浓度Elevated CO₂ concentration	0.50±0.03a	0.81±0.02a	0.53±0.06a
	常规氮用量 Control nitrogen	常规CO₂浓度Ambient CO₂ concentration	0.49±0.02a	0.76±0.02a	0.55±0.02a
	常规氮用量 Control nitrogen	高CO₂浓度Elevated CO₂ concentration	0.51±0.01a	0.77±0.02a	0.55±0.04a
大口期 Twelve-leaf stage (V12)	低氮用量Low nitrogen	常规CO₂浓度Ambient CO₂ concentration	0.58±0.03a	0.80±0.03a	0.43±0.00a
	低氮用量Low nitrogen	高CO₂浓度Elevated CO₂ concentration	0.61±0.03a	0.84±0.03a	0.40±0.04a
	常规氮用量 Control nitrogen	常规CO₂浓度Ambient CO₂ concentration	0.59±0.02a	0.82±0.01a	0.41±0.03a
	常规氮用量 Control nitrogen	高CO₂浓度Elevated CO₂ concentration	0.62±0.02a	0.82±0.03a	0.39±0.06a
抽雄期 Tasseling stage (VT)	低氮用量Low nitrogen	常规CO₂浓度Ambient CO₂ concentration	0.43±0.02b	0.68±0.03bc	0.53±0.04a
	低氮用量Low nitrogen	高CO₂浓度Elevated CO₂ concentration	0.51±0.03ab	0.75±0.02ac	0.47±0.01ab
	常规氮用量 Control nitrogen	常规CO₂浓度Ambient CO₂ concentration	0.48±0.01ab	0.66±0.03b	0.45±0.04ab
	常规氮用量 Control nitrogen	高CO₂浓度Elevated CO₂ concentration	0.55±0.03a	0.77±0.02a	0.43±0.03b
灌浆期 Filling stage (R3)	低氮用量Low nitrogen	常规CO₂浓度Ambient CO₂ concentration	0.47±0.05a	0.76±0.03a	0.62±0.03a
	低氮用量Low nitrogen	高CO₂浓度Elevated CO₂ concentration	0.48±0.03a	0.73±0.01a	0.61±0.08a
	常规氮用量 Control nitrogen	常规CO₂浓度Ambient CO₂ concentration	0.56±0.04a	0.75±0.04a	0.61±0.10a
	常规氮用量 Control nitrogen	高CO₂浓度Elevated CO₂ concentration	0.58±0.02a	0.75±0.04a	0.58±0.05a
同列不同小写字母表示在同一时期不同处理间差异达5%显著水平。Different lowercase letters in the same column indicate significant differences among different treatments at the same stage (P < 0.05).

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表3不同CO₂浓度和氮肥用量对玉米地上部干物质量的影响
Table3.Effects of different CO₂ concentration and nitrogen fertilizer application treatments on dry matter weight of aboveground parts of maize

处理 Treatment		干物质Dry matter (g·plant^-1)
处理 Treatment		六叶期 Six-leaf stage (V6)	大口期 Twelve-leaf stage (V12)	抽雄期 Tasseling stage (VT)	灌浆期 Filling stage (R3)	成熟期 Maturity (R6)
低氮用量 Low nitrogen	常规CO₂浓度Ambient CO₂ concentration	6.8±0.8a	87.5±1.2b	120.4±2.0a	178.8±7.2b	367.5±17.1a
低氮用量 Low nitrogen	高CO₂浓度Elevated CO₂ concentration	7.5±0.5a	97.6±3.7ab	114.5±3.2a	203.6±4.1a	368.3±18.6a
常规氮用量 Control nitrogen	常规CO₂浓度Ambient CO₂ concentration	7.6±0.4a	96.0±5.8ab	116.1±2.9a	214.4±8.1a	391.9±12.5a
常规氮用量 Control nitrogen	高CO₂浓度Elevated CO₂ concentration	8.3±0.4a	106.2±4.7a	126.5±5.6a	217.7±6.0a	392.5±11.2a
显著性 Significant	CO₂	ns	ns	ns	ns	ns
	N	ns	ns	ns	**	ns
	CO₂×N	ns	ns	ns	ns	ns
同列不同小写字母表示在同一时期不同处理间差异达5%显著水平。ns表示影响不显著, 表示同一生育期CO₂、氮肥及其交互作用的影响在P < 0.01水平显著。Different lowercase letters in the same column indicate significant differences among different treatments during the same growth stage (P < 0.05). “ns” denotes no significant effect, indicate significant effects of CO₂, nitrogen fertilizer and their interaction during the same growth stage at P < 0.01.

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表4不同CO₂浓度和氮肥用量对玉米产量及其构成因素和部分穗性状的影响
Table4.Effects of different CO₂ concentration and nitrogen fertilizer application treatments on maize yield and its components and some ear characters

处理 Treatment		穗长 Ear length (cm)	穗粗 Ear diameter (cm)	穗粒数 Kernels per plant	穗粒重 Kernel weight (g)	千粒重 1000-kernel weight (g)	产量 Grain yield (kg·hm^-2)
处理 Treatment		低氮用量(LN) Low nitrogen	常规CO₂浓度Ambient CO₂ concentration	21.1±0.3a	16.1±0.1bc	630.2±11.2a	192.6±5.1b	317.7±4.8b	8 379±226b
高CO₂浓度Elevated CO₂ concentration	21.2±0.4a	低氮用量(LN) Low nitrogen	15.8±0.2b	626.6±11.2a	193.4±4.5b	328.4±7.8ab	8 399±112b
常规氮用量(CN) Control nitrogen	常规CO₂浓度Ambient CO₂ concentration	21.3±0.2a	16.5±0.1ac	638.0±12.3a	213.0±4.5a	344.9±4.3a	9 219±65a
常规氮用量(CN) Control nitrogen	高CO₂浓度Elevated CO₂ concentration	21.1±0.1a	16.7±0.2a	629.1±9.1a	210.3±2.3a	337.7±5.2a	9 311±187a
差异显著性 Significant	CO₂	ns	ns	ns	ns	ns	ns
	N	ns	**	ns	**	*	**
	CO₂ × N	ns	ns	ns	ns	ns	ns
同列不同小写字母表示不同处理间差异达5%显著水平。ns表示影响不显著, 和表示同一生育期CO₂、氮肥及其交互作用的影响在P < 0.05和P < 0.01水平显著。Different lowercase letters in the same column indicate significant differences among different treatments during the same growth stage (P < 0.05). “ns” denotes no significant effect, and ** indicate significant effects of CO₂, nitrogen fertilizer and their interaction during the same growth stage at P < 0.05 and P < 0.01, respectively.

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