C4植物玉米的光合-光响应曲线模拟研究

doi:10.17521/cjpe.2017.0158

摘要/Abstract

摘要：

光是植物光合作用最基本的一个决定因子, 准确分析光响应曲线及其参数是研究光合生理生态过程对环境变化响应的重要途径; 但相关模型及其模拟的准确性仍待改进。该研究基于C₄作物玉米(Zea mays)不同干旱处理试验资料, 比较研究了现有光响应模型(直角双曲线模型、非直角双曲线模型、直角双曲线修正模型、指数模型、二次函数模型以及新提出的改进模型)的适应性。结果表明, 改进的光响应模型具有较好的精确度, 可较准确地描述光响应曲线, 也能够准确拟合最大净光合速率、光饱和点、光补偿点以及暗呼吸速率4个关键光合参数。该结果为研究植物光合生理生态过程及其环境适应性提供了一个改进的模拟方法。

关键词: 光响应模型, 改进模型, 干旱适用性, 光合参数, 玉米, C₄植物

Abstract: Aims A light response curve can reflect a plant’s ability to utilize light, which is also a key tool in determining the relationship between photosynthetic capacity and environmental factors; however the model accuracies concerning the light response curve remain elusive. The objectives of this study were to compare and assess the model accuracies related to a light response curve and the effects of drought. Methods A field rain shelter was used to control the soil water conditions. To obtain photosynthesis parameters from the light response curve and the drought effects, the relevant models (including the rectangular model, non-rectangular hyperbolic model, modified rectangular hyperbolic model, exponential model, quadratic function model, and a newly modified model) were applied to fit the light response curves. The validity of each model was tested by analyzing the differences between the fitted values obtained by the models and the measured values. Important findings The newly modified model has been proved to performing relatively better in accurately describing the light response curve patterns, and credibly obtaining the crucial photosynthetic parameters such as the maximum net photosynthetic rate, light saturation point, light compensation point, and dark respiration rate, especially under high radiation conditions.

Key words: light response curve, modified model, drought adaptation, photosynthetic parameter, maize, C₄ plants

李义博, 宋贺, 周莉, 许振柱, 周广胜. C₄植物玉米的光合-光响应曲线模拟研究. 植物生态学报, 2017, 41(12): 1289-1300. DOI: 10.17521/cjpe.2017.0158

Li Yi-Bo, SONG He, ZHOU Li, XU Zhen-Zhu, ZHOU Guang-Sheng. Modeling study on photosynthetic-light response curves of a C₄ plant, maize. Chinese Journal of Plant Ecology, 2017, 41(12): 1289-1300. DOI: 10.17521/cjpe.2017.0158

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2017.0158

https://www.plant-ecology.com/CN/Y2017/V41/I12/1289

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处理 Treatment	模型 Model	初始量子效率 α (μmol·μmol^-1)	最大净光合速率 P_nmax (μmol·m^-2·s^-1)	光饱和点 LSP (μmol·m^-2·s^-1)	光补偿点 LCP (μmol·m^-2·s^-1)	暗呼吸速率 R_d (μmol·m^-2·s^-1)
拔节期至抽雄期正常处理 control treatment between jointing stage and tasseling stage	直角双曲线模型 Rectangular hyperbolic model	0.072	52.08	1 287.62	52.73	3.55
	非直角双曲线模型 Non-rectangular hyperbolic model	0.041	36.17	912.31	58.26	2.37
	直角双曲线修正模型 Modified rectangular hyperbolic model	0.049	32.22	1 536.90	57.42	2.76
	指数模型 Exponential model	0.057	34.74	161.57	56.43	2.62
	二次函数模型 Quadratic function model	0.047	32.56	1 512.69	57.23	2.61
	改进模型 Modified model	0.043	31.88	1 596.85	58.80	2.23
	实测值 Measured value	-	31.78	1 775.00	56.05	2.51
拔节期至抽雄期干旱12天处理 continuous 12 days drought treatment between jointing stage and tasseling stage	直角双曲线模型 Rectangular hyperbolic model	0.078	37.05	886.63	59.43	4.14
	非直角双曲线模型 Non-rectangular hyperbolic model	0.040	27.42	674.09	64.88	2.68
	直角双曲线修正模型 Modified rectangular hyperbolic model	0.053	24.41	1 390.46	66.07	3.35
	指数模型 Exponential model	0.055	25.07	117.81	21.66	1.15
	二次函数模型 Quadratic function model	0.042	25.60	1 340.60	59.19	2.42
	改进模型 Modified model	0.047	24.08	1 398.59	67.91	2.88
	实测值 Measured value	-	23.94	1 500.00	68.83	3.12
拔节期至抽雄期干旱26天处理 continuous 26 days drought treatment between jointing stage and tasseling stage	直角双曲线模型 Rectangular hyperbolic model	0.073	17.00	740.68	33.52	2.13
	非直角双曲线模型 Non-rectangular hyperbolic model	0.023	12.84	572.64	36.57	0.85
	直角双曲线修正模型 Modified rectangular hyperbolic model	0.036	14.04	1 033.40	47.55	1.65
	指数模型 Exponential model	0.046	12.43	58.52	41.03	1.16
	二次函数模型 Quadratic function model	0.029	14.61	1 094.81	46.37	1.32
	改进模型 Modified model	0.022	14.18	889.19	56.63	1.33
	实测值 Measured value	-	13.90	800.00	55.27	1.37

处理 Treatment	模型 Model	初始量子效率 α (μmol·μmol^-1)	最大净光合速率 P_nmax (μmol·m^-2·s^-1)	光饱和点 LSP (μmol·m^-2·s^-1)	光补偿点 LCP (μmol·m^-2·s^-1)	暗呼吸速率 R_d (μmol·m^-2·s^-1)
拔节期至抽雄期正常处理 control treatment between jointing stage and tasseling stage	直角双曲线模型 Rectangular hyperbolic model	0.072	52.08	1 287.62	52.73	3.55
	非直角双曲线模型 Non-rectangular hyperbolic model	0.041	36.17	912.31	58.26	2.37
	直角双曲线修正模型 Modified rectangular hyperbolic model	0.049	32.22	1 536.90	57.42	2.76
	指数模型 Exponential model	0.057	34.74	161.57	56.43	2.62
	二次函数模型 Quadratic function model	0.047	32.56	1 512.69	57.23	2.61
	改进模型 Modified model	0.043	31.88	1 596.85	58.80	2.23
	实测值 Measured value	-	31.78	1 775.00	56.05	2.51
拔节期至抽雄期干旱12天处理 continuous 12 days drought treatment between jointing stage and tasseling stage	直角双曲线模型 Rectangular hyperbolic model	0.078	37.05	886.63	59.43	4.14
	非直角双曲线模型 Non-rectangular hyperbolic model	0.040	27.42	674.09	64.88	2.68
	直角双曲线修正模型 Modified rectangular hyperbolic model	0.053	24.41	1 390.46	66.07	3.35
	指数模型 Exponential model	0.055	25.07	117.81	21.66	1.15
	二次函数模型 Quadratic function model	0.042	25.60	1 340.60	59.19	2.42
	改进模型 Modified model	0.047	24.08	1 398.59	67.91	2.88
	实测值 Measured value	-	23.94	1 500.00	68.83	3.12
拔节期至抽雄期干旱26天处理 continuous 26 days drought treatment between jointing stage and tasseling stage	直角双曲线模型 Rectangular hyperbolic model	0.073	17.00	740.68	33.52	2.13
	非直角双曲线模型 Non-rectangular hyperbolic model	0.023	12.84	572.64	36.57	0.85
	直角双曲线修正模型 Modified rectangular hyperbolic model	0.036	14.04	1 033.40	47.55	1.65
	指数模型 Exponential model	0.046	12.43	58.52	41.03	1.16
	二次函数模型 Quadratic function model	0.029	14.61	1 094.81	46.37	1.32
	改进模型 Modified model	0.022	14.18	889.19	56.63	1.33
	实测值 Measured value	-	13.90	800.00	55.27	1.37

处理 Treatment	模型 Model	R²	拟合MSE Fitted MSE	拟合MAE Fitted MAE	实测MSE Test MSE	实测MAE Test MAE
拔节期至抽雄期正常处理 control treatment between jointing stage and tasseling stage	直角双曲线模型 Rectangular hyperbolic model	0.991β8	1.68	1.08	3.74	1.50
	非直角双曲线模型 Non-rectangular hyperbolic model	0.999β6	0.08	0.24	0.86	0.81
	直角双曲线修正模型 Modified rectangular hyperbolic model	0.999β4	0.12	0.29	1.01	0.82
	指数模型 Exponential model	0.997β0	0.74	0.69	4.05	1.87
	二次函数模型 quadratic function model	0.999β3	0.15	0.30	0.75	0.72
	改进模型 Modified model	0.999β7	0.05	0.19	0.91	0.80
拔节期至抽雄期干旱12天处理 continuous 12 days drought treatment between jointing stage and tasseling stage	直角双曲线模型 Rectangular hyperbolic model	0.984β7	1.52	1.03	3.19	1.22
	非直角双曲线模型 Non-rectangular hyperbolic model	0.995β8	0.40	0.52	0.58	0.49
	直角双曲线修正模型 Modified rectangular hyperbolic model	0.997β3	0.25	0.45	0.49	0.61
	指数模型 Exponential model	0.993β0	0.71	0.73	5.42	2.05
	二次函数模型 Quadratic function model	0.992β7	0.74	0.76	0.71	0.57
	改进模型 Modified model	0.997β2	0.26	0.39	0.39	0.36
拔节期至抽雄期干旱26天处理 continuous 26 days drought treatment between jointing stage and tasseling stage	直角双曲线模型 Rectangular hyperbolic model	0.920β5	2.71	1.34	4.31	1.83
	非直角双曲线模型 Non-rectangular hyperbolic model	0.961β0	1.24	0.90	3.31	1.45
	直角双曲线修正模型 Modified rectangular hyperbolic model	0.997β1	0.10	0.26	0.37	0.48
	指数模型 Exponential model	0.950β8	1.73	1.06	5.20	1.60
	二次函数模型 quadratic function model	0.992β0	0.31	0.48	0.61	0.71
	改进模型 Modified model	0.999β2	0.03	0.14	0.14	0.21

处理 Treatment	模型 Model	R²	拟合MSE Fitted MSE	拟合MAE Fitted MAE	实测MSE Test MSE	实测MAE Test MAE
拔节期至抽雄期正常处理 control treatment between jointing stage and tasseling stage	直角双曲线模型 Rectangular hyperbolic model	0.991β8	1.68	1.08	3.74	1.50
	非直角双曲线模型 Non-rectangular hyperbolic model	0.999β6	0.08	0.24	0.86	0.81
	直角双曲线修正模型 Modified rectangular hyperbolic model	0.999β4	0.12	0.29	1.01	0.82
	指数模型 Exponential model	0.997β0	0.74	0.69	4.05	1.87
	二次函数模型 quadratic function model	0.999β3	0.15	0.30	0.75	0.72
	改进模型 Modified model	0.999β7	0.05	0.19	0.91	0.80
拔节期至抽雄期干旱12天处理 continuous 12 days drought treatment between jointing stage and tasseling stage	直角双曲线模型 Rectangular hyperbolic model	0.984β7	1.52	1.03	3.19	1.22
	非直角双曲线模型 Non-rectangular hyperbolic model	0.995β8	0.40	0.52	0.58	0.49
	直角双曲线修正模型 Modified rectangular hyperbolic model	0.997β3	0.25	0.45	0.49	0.61
	指数模型 Exponential model	0.993β0	0.71	0.73	5.42	2.05
	二次函数模型 Quadratic function model	0.992β7	0.74	0.76	0.71	0.57
	改进模型 Modified model	0.997β2	0.26	0.39	0.39	0.36
拔节期至抽雄期干旱26天处理 continuous 26 days drought treatment between jointing stage and tasseling stage	直角双曲线模型 Rectangular hyperbolic model	0.920β5	2.71	1.34	4.31	1.83
	非直角双曲线模型 Non-rectangular hyperbolic model	0.961β0	1.24	0.90	3.31	1.45
	直角双曲线修正模型 Modified rectangular hyperbolic model	0.997β1	0.10	0.26	0.37	0.48
	指数模型 Exponential model	0.950β8	1.73	1.06	5.20	1.60
	二次函数模型 quadratic function model	0.992β0	0.31	0.48	0.61	0.71
	改进模型 Modified model	0.999β2	0.03	0.14	0.14	0.21

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C₄植物玉米的光合-光响应曲线模拟研究

Modeling study on photosynthetic-light response curves of a C₄ plant, maize

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