植物电子传递速率光响应模型的研究进展

doi:10.17521/cjpe.2022.0409

摘要/Abstract

摘要：

电子传递速率光响应模型是研究植物光合生理和生态学的重要工具, 可为量化原初反应光能的吸收和传递对光的响应提供理论依据。该文综述了目前常用的电子传递速率光响应模型的数学特征, 分析了不同模型的优势及其在实际应用中的潜在问题, 并在此基础上对这些模型可能的发展趋势进行了展望。原初反应包括光能的吸收、光合色素分子的激发和退激发(包括光化学反应、荧光发射和热耗散)、激子共振传递以及光系统II (PSII)反应中心发生电荷分离产生电子传递速率等一系列复杂的物理和生化反应过程。电子传递速率光响应经验模型和半机理模型因不涉及或只涉及部分原初反应过程而难以解释藻类和高等植物的PSII动力学下调、光适应和光保护等现象。电子传递速率光响应机理模型综合考虑了光合色素分子的物理参数(如本征光能吸收截面(σ_ik)、分子处于最低激发态的平均寿命(τ_min)、分子的能级简并度和处于激发态的光合色素分子数(N_k))在整个原初反应过程中的重要作用, 不仅可以获得藻类和高等植物叶片的最大电子传递速率以及对应的饱和光强等光合参数, 还可以获得σ_ik和τ_min等重要的物理参数, 以及有效光能吸收截面($\sigma_{\mathrm{ik}}^{\prime}$)和N_k对光的响应规律等。将环境因子(如温度、CO₂浓度等)耦合到已有的电子传递速率光响应机理模型中, 并明确其与植物的$\sigma_{\mathrm{ik}}^{\prime}$和N_k等参数间的关系, 可能是今后光合电子传递速率光响应机理模型的发展方向。

关键词: 电子传递速率, 原初反应, 经验模型, 半机理模型, 机理模型

Abstract:

The light response curve of electron transport rate is an important tool to investigate plant physiology and ecology. It can provide a theoretical basis for quantifying the absorption and transmission of light energy in primary reaction. In this paper, the mathematical characteristics, the advantages, and the potential weaknesses in practical application and research trends of the light response models of electron transport rate are reviewed and discussed. The primary reaction, which includes absorption of light energy, excitation and de-excitation of photosynthetic pigment molecules (including photochemical reaction, fluorescence emission and heat dissipation), and electron transport rate stemming from charge separation in the photosystem II (PSII) reaction center caused by exciton resonance, is consisted by a series of complex physical and biochemical reactions. The classical and semi-mechanistic models of light response curve of electron transport rate are difficult to explain the dynamic down-regulation of PSII, light adaptation and light protection of algae and higher plant, because they did not involve or only partly involved the primary reaction process. However, the mechanism model takes into account the important role of the physical parameters of the photosynthetic pigment molecule (e.g., the eign-absorption cross-section of light energy (σ_ik), the average life-time of the molecule in the lowest excited state (τ_min), the energy level degeneracy of the molecule and the number of photosynthetic pigment molecules in the excited state (N_k)) in the whole primary reaction process. This model can not only obtain the maximum electron transfer rate and its corresponding saturation light intensity of algae and higher plant, but also get some important physical parameters such as σ_ik and τ_min. Meanwhile, it also can obtain the laws about light-response of the effective light energy absorption cross-section ($\sigma_{\mathrm{ik}}^{\prime}$) and of N_k. It may be the developmental direction of the mechanism model of electron transport rate response to light in the future when the mechanistic model was coupled with some environmental factors (e.g., temperature and CO₂ concentration) and the relationship between light intensity and $\sigma_{\mathrm{ik}}^{\prime}$ and N_k were determined.

Key words: electron transport rate, primary reaction, classical model, semi-mechanistic model, mechanistic model

王复标, 叶子飘. 植物电子传递速率光响应模型的研究进展. 植物生态学报, 2024, 48(3): 287-305. DOI: 10.17521/cjpe.2022.0409

WANG Fu-Biao, YE Zi-Piao. A review on light response models of electron transport rates of plant. Chinese Journal of Plant Ecology, 2024, 48(3): 287-305. DOI: 10.17521/cjpe.2022.0409

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2022.0409

https://www.plant-ecology.com/CN/Y2024/V48/I3/287

图/表 11

图1 不同光系统II (PSII)动力学下调系数(β)值(A)和饱和系数(γ)值(B)对J/I响应曲线的影响。I, 光强; J, 电子传递速率。α, 初始斜率。

Fig. 1 Effect of different values of β (A) and γ (B) on the J/I curves. I, light intensity; J, electron transport rate. α, initial slope; β, coefficient of dynamic down-regulation for photosystem II (PSII); γ, saturation coefficient.

图2 不同光系统II (PSII)动力学下调系数(β)值(A)和饱和系数(γ)值(B)对$\sigma_{\mathrm{ik}}^{\prime}$/I响应曲线的影响。$\sigma_{\mathrm{ik}}^{\prime}$, 有效光能吸收截面; I, 光强。α, 初始斜率。

Fig. 2 Effect of different values of β (A) and γ (B) on the$\sigma_{\mathrm{ik}}^{\prime}$/I curves. $\sigma_{\mathrm{ik}}^{\prime}$, effective light energy absorption cross-section; I, light intensity. α, initial slope; β, coefficient of dynamic down-regulation for photosystem II (PSII); γ, saturation coefficient.

图3 3个模型拟合小麦(A)和大豆(B)的J/I响应曲线(平均值±标准误, n = 3)。I, 光强; J, 电子传递速率; 公式(16),$ J=\alpha \frac{1-\beta I}{1+\gamma I} I$。

Fig. 3 J/I curves fitted by three models for Triticum aestivum (A) and Glycine max (B) (mean ± SE, n = 3). I, light intensity; J, electron transport rate; Eq. (16), $ J=\alpha \frac{1-\beta I}{1+\gamma I} I$.

表1 3个模型拟合小麦和大豆的J/I响应曲线得到的光合参数(平均值±标准误)

Table 1 Photosynthetic parameters of J/I curves fitted by three models for Triticum aestivum and Glycine max (mean ± SE)

参数 Parameter	小麦 Triticum aestivum				大豆 Glycine max
参数 Parameter	公式(16) Eq. (16)	非直角双曲线模型 NRH model	双指数函数模型 Double exponent model	观测 Observation	公式(16) Eq. (16)	非直角双曲线模型 NRH	双指数函数模型 Double exponent model	观测 Observation
曲度 Curvature (θ)	-	0.816 ± 0.009		-	-	0.924 ± 0.005		-
初始斜率 Initial slope (α)	0.295 ± 0.012^a	0.282 ± 0.012^a	0.282 ± 0.012^a	-	0.299 ± 0.006^a	0.282 ± 0.005^a	0.282 ± 0.012^a	-
光系统II动力学下调系数 Coefficient of dynamic down-regulation for photosystem II (β)	(2.42 ± 0.28) × 10^-3	-	37.98 ± 1.27	-	(3.07 ± 0.08) × 10^-4	-	(6.13 ± 0.27) × 10³	-
饱和系数 Saturation coefficient (γ)	(1.26 ± 0.66) × 10^-4	-	-	-	(-1.50 ± 0.24) × 10^-4	-	-	-
β = 0时最大电子传递速率 Maximum photosynthetic electron flow when β = 0 (J_s)	-	-	(8.47± 0.35) × 10⁵	-	-	-	(1.81 ± 0.23) × 10⁸	-
最大电子传递速率 Maximum photosynthetic electron flow (J_max)	257.23 ± 7.36^b	304.91 ± 7.11^a	264.74 ± 8.45^b	261.56± 7.32^b	332.79 ± 5.16^b	373.87 ± 5.47^a	367.61 ± 8.13^a	332.86 ± 5.01^b
饱和光强 Saturation irradiance (PAR_sat)	1 873.37 ± 109.46^b	-	2 221.35 ± 125.89^a	1 734.16 ± 66.15^b	1 906.01 ± 19.97^b	-	2 948.74 ± 105.78^a	1 933.23 ± 66.27^b
确定系数 Determined coefficient (R²)	0.998 4 ± 0.000 9	0.998 2 ± 0.000 7	0.999 4 ± 0.000 6	-	0.998 9 ± 0.000 1	0.999 2 ± 0.000 1	0.998 0 ± 0.000 2	-

图4 3个水稻品种的电子传递速率光响应曲线(平均值±标准误, n = 3)。I, 光强; J, 电子传递速率; 公式(16), $ J=\alpha \frac{1-\beta I}{1+\gamma I} I$。

Fig. 4 Light response of electron transport rate for three Oryza sativa cultivars (mean ± SE, n = 3). I, light intensity; J, electron transport rate; Eq. (16), $ J=\alpha \frac{1-\beta I}{1+\gamma I} I$.

表2 3个水稻品种电子传递速率光响应的特征参数(平均值±标准误)

Table 2 Characteristic parameters of light response curves of electron transport rate for three Oryza sativa cultivars (mean ± SE)

参数 Parameter	品种 Cultivars
参数 Parameter	‘深95优1326’ ‘Shen95you1326’	‘五丰优1326’ ‘Wufengyou1326’	‘赣香优1326’ ‘Ganxiangyou1326’
初始斜率 Initial slope (α) (μmol·μmol^-1)	0.308 ± 0.017^a	0.321 ± 0.003^a	0.281 ± 0.005^b
最大电子传递速率 Maximum electron transport rate (J_max) (μmol·m^-2·s^-1)	116.99 ± 6.97^a	115.63 ± 2.16^a	102.48 ± 0.58^b
饱和光强 Saturation irradiance (PAR_sat) (μmol·m^-2·s^-1)	1 155.84 ± 16.41^a	1 181.05 ± 11.32^a	1 076.22 ± 13.55^a
叶绿素含量 Chlorophyll content (mg·dm^-2)	4.58 ± 0.34^b	6.11 ± 0.31^a	5.23 ± 0.25^ab
叶绿素a/叶绿素b Chlorophyll a/ chlorophyll b	2.80 ± 0.54^a	2.23 ± 0.07^a	2.37 ± 0.02^a
本征截面 Eign cross-section (σ_ik) (×10^-21m²)	2.69 ± 0.27^a	2.09 ± 0.10^b	2.13 ± 0.09^b
最小平均寿命 Minimum average life-time (τ_min) (×10^-3s)	1.41 ± 0.21^b	1.62 ± 0.03^a	1.43 ± 0.04^b
确定系数 Determined coefficient (R²)	0.998 3	0.999 3	0.989 3

图5 3个水稻品种光合色素分子的有效光能吸收截面($\sigma_{\mathrm{ik}}^{\prime}$)对光的响应(平均值±标准误, n = 3)。I, 光强。

Fig. 5 Light response of the effective light energy absorption cross-section ($\sigma_{\mathrm{ik}}^{\prime}$) for three Oryza sativa cultivars (mean ± SE, n = 3). I, light intensity.

图6 3个水稻品种处于激发态的光合色素分子数(Nk)对光的响应(平均值±标准误, n = 3)。I, 光强。

Fig. 6 Light response of molecules in the excited state (Nk) for three Oryza sativa cultivars (mean ± SE, n = 3). I, light intensity.

图7 杂种杜鹃在遮阴和全日照条件下的J/I响应曲线和NPQ/I响应曲线(平均值±标准误, n = 5)。I, 光强; J, 电子传递速率; NPQ, 非光化学淬灭; 公式(16),$ J=\alpha \frac{1-\beta I}{1+\gamma I} I$。

Fig. 7 J/I and NPQ/I curves for Rhododendron ‘Hybrida’ under shading and full sunlight conditions (mean ± SE, n = 5). I, light intensity; J, electron transport rate; NPQ, non-photochemical quenching; Eq. (16), $J=\alpha \frac{1-\beta I}{1+\gamma I} I$.

图8 杂种杜鹃在遮阴和全日照条件下的Nk/I和NPQ/I响应曲线(平均值±标准误, n = 5)。I, 光强; Nk, 处于激发态的光合色素分子数; NPQ, 非光化学淬灭; 公式(21),$ N_{\mathrm{k}}=\frac{1}{1+g_{\mathrm{i}} / g_{\mathrm{k}}} \frac{\gamma I}{1+\gamma I} N_{0}$。

Fig. 8 Nk/I and NPQ/I curves for Rhododendron ‘Hybrida’ under shading and full sunlight conditions(mean ± SE, n = 5). I, light intensity; Nk, molecules in the excited state; NPQ, non-photochemical quenching; Eq. (21),$ N_{\mathrm{k}}=\frac{1}{1+g_{\mathrm{i}} / g_{\mathrm{k}}} \frac{\gamma I}{1+\gamma I} N_{0}$.

图9 光合作用电子传递的Z链。A0和A1, 光系统I (PSI)原初电子受体; Cytb363HP, 363HP细胞色素b; Cytb363LP, 363LP细胞色素b; Cytf, 细胞色素f; FA/FB, PSI次级电子受体; FeS, 铁硫中心; Fd, 铁氧还蛋白; FNR, Fd-NADP还原酶; FX, 非血红素铁硫蛋白; hv, 光照; Mn, 锰簇; NADP+, 脱氢烟酰胺腺嘌呤二核苷磷酸; NADPH, 烟酰胺腺嘌呤二核苷磷酸; PC, 质蓝素; Phe, 脱镁叶绿素; PQ, 质体醌; P700, PSI反应中心色素; P700*, 激发态的PSI反应中心色素; P680, 光系统II (PSII)反应中心色素; P680*, 激发态的PSII反应中心色素; QA, PSII初级醌受体; QB, PSII次级醌受体; Tyr, 酪氨酸。

Fig. 9 Z chain of electron transport in photosynthesis. A0 and A1, the primary electron acceptor of photosystem I (PSI); Cytb363HP, 363HP cytochrome b; Cytb363LP, 363LP cytochrome b; Cytf, cytochrome f; FA/FB, the secondary electron acceptor of PSI; FeS, iron-sulifide protein; Fd, ferredoxin; FNR, Fd-NADP reductase; Fx, non-heme iron sulfide protein; hv, irradiation with light; Mn, manganese cluster; NADP+, dehydronicotinamide adenine dinucleotide phosphate; NADPH, nicotinamide adenine dinucleotide phosphate; PC, plasto cynin; Phe, phephytin; PQ, plastoquinone; P700, the pigment of PSI reaction center; P700*, the excited pigment of PSI reaction center; P680, the pigment of photosystem II (PSII) reaction center; P680*, the excited pigment of PSII reaction center; QA, the primary quinone electron acceptor of PSII; QB, the secondary quinone electron acceptor of PSII; Tyr, tyrosine.

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