光合作用对光和CO2响应模型的研究进展

doi:10.3773/j.issn.1005-264x.2010.06.012

摘要/Abstract

摘要：

光合作用对光和CO₂响应模型是研究植物生理和植物生态学的重要工具, 可为植物光合特性对主要环境因子的响应提供科学依据。该文综述了当前光合作用对光和CO₂响应模型的研究进展和存在的问题, 并在此基础上探讨了这些模型的可能发展趋势。光合作用涉及光能的吸收、能量转换、电子传递、ATP合成、CO₂固定等一系列复杂的物理和化学反应过程。光合作用由原初反应、同化力形成和碳同化3个基本过程构成, 任一个过程均可对光合作用速率产生直接的影响。光合作用对光响应模型只涉及光能的转换, 而光合作用的生化模型包含了同化力形成和碳同化这两个基本过程。把光合作用的原初反应, 即把参与光能吸收、传递和转换的捕光色素分子的物理参数(如捕光色素分子数、捕光色素分子光能吸收截面、捕光色素分子处于激发态的平均寿命等)结合到生化模型中, 可能是今后光合作用对光响应机理模型的发展方向。

关键词: 生化模型, CO₂响应模型, 光响应模型

Abstract:

The light and CO₂ response curve of photosynthesis is an important tool to study plant physiology and plant ecology that can provide a scientific basis for the response of plant photosynthetic properties to environmental factors. This review considered the progress and potential weaknesses of light and CO₂ response models of photosynthesis and discussed research trends. Photosynthesis, which involves energy of light, absorption, energy conversion, electron transfer, ATP synthesis, CO₂ fixation etc., is a complex physical and chemical reaction process. It includes three basic steps: the primary reaction, the assimilatory power forms and the carbon assimilation, and each link may directly influence other processes. Classical models on photosynthetic light response only involve with light energy absorption, and biochemistry models do with the assimilatory power to form as well as carbon assimilation. A future direction of research of the mechanistic model of photosynthetic light response is the primary reaction of photosynthesis, namely participation the energy of light absorption, the transmission and the transformation of the harvesting light pigment member the physical parameter (e.g., the light-harvesting pigment molecules, light energy absorption cross-section of the harvesting pigment, the mean lifetime of the harvesting light pigment) unify in the biochemistry model.

Key words: biochemical model, CO₂ response model of photosynthesis, light-response model of photosynthesis

叶子飘. 光合作用对光和CO₂响应模型的研究进展. 植物生态学报, 2010, 34(6): 727-740. DOI: 10.3773/j.issn.1005-264x.2010.06.012

YE Zi-Piao. A review on modeling of responses of photosynthesis to light and CO₂. Chinese Journal of Plant Ecology, 2010, 34(6): 727-740. DOI: 10.3773/j.issn.1005-264x.2010.06.012

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链接本文: https://www.plant-ecology.com/CN/10.3773/j.issn.1005-264x.2010.06.012

https://www.plant-ecology.com/CN/Y2010/V34/I6/727

图/表 6

图1 冬小麦的光响应曲线(引自文献Yu et al., 2004)。 A, 4个光合作用对光响应模型拟合华北平原冬小麦的光响应曲线。○, 测量点; —, 修正模型的拟合点; ☆, 直角双曲线的拟合点; ┅, 非直角双曲线的拟合点; ▽, 指数方程的拟合点。B, 4个光合作用对光响应模型估算饱和光强的方法。a, 由直线方程结合非直角双曲线得到的饱和光强; b, 由直线方程结合直角双曲线得到的饱和光强; c, 净光合速率为0.9Pnmax所对应光强为饱和光强; d, 由修正模型得到的饱和光强。

Fig. 1 Light-response curve of photosynthesis for Triticum aestivum (Cited from Yu et al., 2004). A, Light-response curve of photosynthesis for T. aestivum fitted by four photosynthetic light response models. ○, measured points; —, points fitted by modified model; ☆, points fitted by rectangular hyperbola; ┅, points fitted by non-rectangular hyperbola; ▽, pointed fitted by exponential equation. B, Saturation irradiance estimated by four photosynthetic light response models. a, saturation irradiance obtained by a method combined non-rectangular hyperbola with line equation; b, saturation irradiance obtained by a method combined rectangular hyperbola with line equation; c, saturation irradiance, i.e. irradiance at 0.9Pnmax; d, saturation irradiance obtained by the modified model.

表1 四个光响应模型拟合温度在20 ℃、CO2浓度在365 μmol·mol-1条件下冬小麦的实测值(引自文献Yu et al., 2004)与拟合结果

Table 1 Results fitted by four models of light-response curve of photosynthesis and measured values for Triticum aestivum at 30 ℃ and 365 μmol·mol-1 CO2 concentration (Cited from Yu et al., 2004)

光合参数 Photosynthetic parameters	初始斜率 Initial slope α	最大净光合速率 P_nmax (μmol·m^-2·s^-1)	饱和光强 I_sat (μmol·m^-2·s^-1)	光补偿点 I_c(μmol·m^-2·s^-1)	暗呼吸速率 R_d (μmol·m^-2·s^-1)	决定系数 Determination coefficient R²
直角双曲线 Rectangular hyperbola	0.076	31.01	671.25*	27.08	-1.93	0.996 2
非直角双曲线 Non-rectangular hyperbola	0.055	26.78	583.18*	23.10	-1.25	0.998 9
指数方程 * Exponential equation*	0.058	24.40	1 288.80**	23.03	-1.30	0.999 1
修正的直角双曲线 Modified rectangular hyperbola	0.062	22.76	1 799.18	23.75	-1.43	0.999 2
测量数据 Measured data	无 None	$\approx $22.87	$\approx $1 800	≈20	-1.25	无 None

图2 低光强条件下生长的三叶鬼针草的光响应曲线(引自文献 Ye & Zhao, 2008)。 A, 4个光合作用对光响应模型拟合低光强条件下生长的三叶鬼针草的光响应曲线。○, 测量点; —, 修正模型的拟合点; ☆, 直角双曲线的拟合点; ┅ , 非直角双曲线的拟合点; ▽, 指数方程的拟合点。B, 4个光合作用对光响应模型估算饱和光强的方法。a, 由直线方程结合非直角双曲线得到的饱和光强; b, 由直线方程结合直角双曲线得到的饱和光强; c, 净光合速率为0.9Pnmax所对应光强为饱和光强; d, 由修正模型得到的饱和光强。

Fig. 2 Light-response curve of photosynthesis for Bidens pilosa grown under low light condition (Cited from Ye & Zhao, 2008). A, Light-response curve of photosynthesis for Bidens pilosa fitted by four photosynthetic light response models. ○, measured points; —, points fitted by modified model; ☆, points fitted by rectangular hyperbola; ┅, points fitted by non-rectangular hyperbola; ▽, pointed fitted by exponential equation. B, Saturation irradiance estimated by four photosynthetic light response models. a, saturation irradiance obtained by a method combined non-rectangular hyperbola with line equation; b, saturation irradiance obtained by a method combined rectangular hyperbola with line equation; c, saturation irradiance, i.e. irradiance at 0.9Pnmax; d, saturation irradiance obtained by the modified model.

表2 四个光响应模型拟合温度在27.3 ℃、CO2浓度在390 μmol·mol-1条件下三叶鬼针草的实测值(引自文献 Ye & Zhao, 2008)与拟合结果

Table 2 Results fitted by four models of light-response curve of photosynthesis and measured values for Bidens pilosa grown under low light condition (Cited from Ye & Zhao, 2008)

光合参数 Photosynthetic parameters	初始斜率 Initial slope α	最大净光合速率 P_nmax (μmol·m^-2·s^-1)	饱和光强 I_sat(μmol·m^-2·s^-1)	光补偿点 I_c(μmol·m^-2·s^-1)	暗呼吸速率 R_d (μmol·m^-2·s^-1)	决定系数 Determination coefficient R²
直角双曲线模型 Rectangular hyperbola	0.118	6.69	199.58*	4.39	-0.48	0.939 6
非直角双曲线模型 Non-rectangular hyperbola	0.042	5.79	171.33*	无 None	0.02	0.964 6
指数方程*** Exponential equation	0.068	6.02	247.92**	3.45	-0.23	0.966 6
直角双曲线的修正模型 Modified rectangular hyperbola	0.071	6.49	646.70	3.17	-0.22	0.996 8
测量数据 Measured data	无 None	$\approx $6.5	$\approx $650	≈3	-0.15	无 None

图3 冬小麦的光合作用对胞间CO2浓度的响应曲线(引自文献Yu et al., 2004)。 A, 3个光合作用对CO2响应模型拟合华北平原冬小麦的光响应曲线。○, 测量点; —, 修正模型的拟合点; ┅, 直角双曲线或Michaelis-Menten模型的拟合点。B, 3个光合作用对CO2响应模型估算饱和CO2浓度的方法。a,净光合速率为0.6Pnmax的CO2浓度为饱和胞间CO2浓度; b, 由修正模型得到的饱和胞间CO2浓度。

Fig. 3 Intercellular CO2 response of photosynthesis for Triticum aestivum (Cited from Yu et al., 2004). A, CO2 response curve of photosynthesis for T. aestivum fitted by three CO2 response models of phtosynthesis. ○, measured points; —, points fitted by a modified model; ┅, points fitted by rectangular hyperbola or Michaelis-Menten model. B, Saturation CO2 estimated by three CO2 response models of photosynthesis. a, saturation intercellular CO2 concentration, i.e. Ci at 0.6Pnmax; b, saturation intercellular CO2 concentration obtained by the modified model.

表3 三种模型拟合冬小麦光合作用对胞间CO2浓度响应的结果

Table 3 Simulated results of CO2 response of photosynthesis with three models for Triticum aestivum

光合参数 Photosynthesis parameter	MichaelisMenten 模型 MichaelisMenten model	直角双曲线模型 Rectangular hyperbola model	修正模型 Modified model	测量值 Measured data
初始羧化效率 (mol·m^-2·s^-1) Initial slope α	无None	0.331	0.228	无None
光合能力 (μmol·m^-2·s^-1) Photosynthetic capacity	68.54	68.54	42.26	$\approx $42.3
饱和胞间CO₂浓度(μmol·mol^-1) saturation intercellular CO₂concentration C_isat	687.02*	687.02*	886.74	$\approx $900
CO₂补偿点 (μmol·mol^-1) CO₂ compensation point Γ	45.58	45.58	47.40	$\approx $48
光呼吸速率 (μmol·m^-2·s^-1) Rate of photorespiration R_p	-12.36	-12.36	-9.51	无 None
决定系数 Determination coefficient (R²)	0.991 3	0.991 3	0.998 7	无 None

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光合作用对光和CO₂响应模型的研究进展

A review on modeling of responses of photosynthesis to light and CO₂

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