花叶矢竹叶片色素合成和光合特性

doi:10.17521/cjpe.2023.0258

摘要/Abstract

摘要：

竹子在自然界中存在较多叶色变异, 花叶矢竹(Pseudosasa japonica f. akebonosuji)叶色变异是典型代表, 通过对花叶矢竹不同叶色叶片光合特性的研究, 可解析其叶色变异机理。该研究以花叶矢竹不同叶色叶片为研究对象, 利用紫外分光光度计、高效液相色谱和连续激发式荧光仪等测定不同叶色叶片的光合色素含量、叶绿素合成前体物质相对含量、光系统活性及光合效应差异, 阐明花叶矢竹叶色变异的生理机制。结果表明: (1)花叶矢竹全绿型和花叶型叶片光合色素含量差异显著, 花叶型叶片的叶绿素a/b值显著低于全绿型, 而类胡萝卜素/叶绿素值显著高于绿叶。(2)花叶矢竹白叶和条纹白叶的叶绿素生物合成前体物质粪卟啉III含量显著高于绿叶, 而相邻产物原卟啉IX含量急剧下降, 导致叶绿素a、叶绿素b含量显著降低。(3)条纹叶的净光合速率、表观量子效率等均显著低于绿叶, 全白叶无光合效应。(4)以全绿叶作为对照, 花叶捕获的激子将电子传递到电子传递链中初级醌受体(Q_A)下游的电子受体的概率(Ψ_o)和以吸收光能为基础的性能指数(PI_ABS)均显著降低, 且PI_ABS的降低幅度大于Ψ_o, 白叶复绿后也没有完全恢复, 表明花叶光系统II (PSII)供体侧供应电子的能力和受体侧接收电子的能力都降低, PSII整体性能低于绿叶。白叶和条纹叶叶绿素K相荧光强度(F_k)占F₀ - F_j振幅的比例(W_k)变化值显著高于绿叶(F₀为初始荧光强度, F_j为J相荧光强度), 白叶复绿后其性能基本恢复。绿叶叶绿素F_j占F₀ - F_p振幅的比例(V_j)变化值显著低于白叶和复绿叶, 与条纹绿叶无显著差异, 表明PSII受体侧性能在绿叶和条纹绿叶上表现一致, 但是复绿叶也没有完全恢复至绿叶水平。(5)花叶和复绿叶820 nm处的光吸收量均小于绿叶, 白叶复绿过程中, 光系统I (PSI)最大氧化还原能力(ΔI/I_o)值显著增加并逐渐恢复到稳定绿叶水平, 说明花叶叶绿素I (P700)氧化还原能力都较低, 白叶复绿后氧化能力与绿叶无显著差异, 但P700⁺的还原能力没有恢复到绿叶水平。(6) PSI与PSII之间的协调性变化(Φ_PSI/PSII)在白叶中显著低于绿叶, PSI/PSII的协调性变差, PSII下降幅度大于PSI。复绿叶片Φ_PSI/PSII显著高于绿叶, 说明PSI恢复而PSII没有恢复, 因此, 复绿叶光系统性能减弱主要由PSII引发。总之, 花叶是由叶绿素含量、叶绿素a/b值下降导致的叶绿素a缺乏型突变, 原卟啉IX含量急剧下降导致叶绿素合成障碍; PSII/PSI的协调性变弱、叶片复绿后PSII性能并未恢复到绿叶水平, 因此全株始终保持花叶, 光合能力及利用效率都低于绿叶, 不同叶色之间存在光合生理差异。

关键词: 花叶矢竹, 叶色变异, 光合色素, 光合系统, 光合特性

Abstract:

Aims Variation in leaf colour is natural in bamboo, and the leaf colour variation of Pseudosasa japonicaf. akebonosuji is a typical representative. The mechanism of colour variation can be resolved by studying the photosynthetic properties of different leaf colours.

Methods The photosynthetic pigment content, the relative content of chlorophyll (Chl) synthesis precursors, photosystem activities and photosynthetic effect differences of leaves with different leaf colours were determined by ultraviolet spectrophotometer, high-performance liquid chromatography and continuous excitation fluorescence to elucidate the physiological mechanism of leaf colour variation of P. japonicaf. akebonosuji.

Important findings (1) There were significant differences in photosynthetic pigment content between the all-green and mosaic leaves of P. japonica f. akebonosuji. The mosaic leaves’s chl a/b value was significantly lower than the all-green leaves, while the carotenoid/Chl a+b value was substantially higher than all-green leaves. (2) The chlorophyll biosynthetic precursor substance, Coprogen III, was significantly higher in albino and striped albino leaves than in green leaves. At the same time, the content of protoporphyrin IX decreased sharply, resulting in significant reductions in Chl a and Chl b content. (3) The net photosynthetic rate and apparent quantum yield of striped leaves were significantly lower than in green leaves, and there was no photosynthetic effect in all-albino leaves. (4) Using all-green leaves as a control, the overall performance of photosystem II in mosaic leaves was lower than that of green leaves. The proportion of fluorescence intensity at J-step (F_j)in F₀ - F_p (F₀, minimal fluorescence intensity; F_p, fluorescence intencity at P-step) amplitude of chlorophyll value of green leaves was significantly lower than that of albino and re-greened leaves. There was no significant difference with striped green leaves, indicating that PSII receptor side performance was consistent on green and striped green leaves, but re-greened leaves did not fully recover to the level of green leaves. (5) The light absorption at 820 nm of mosaic leaves and re-greened leaves was lesser than that of green leaves. During the re-greening of albino leaves, the maximum redox capacity of photosystem I (PSI) (ΔI/I_o) values increased significantly. They gradually returned to the level of stable green leaves, indicating that the redox capacity of chlorophyll I (P700) was low. The oxidative capacity of albino leaves after re-greening was not significantly different from that of green leaves. (6) The change in coordination between PSI and PSII (Φ_PSI/PSII) in albino leaves was significantly lower than in green leaves, and the coordination of PSI/PSII deteriorated, with PSII decreasing more than PSI. PSII mainly triggered the weakening of the photosystem performance of re-greened leaves. The mosaic leaves a Chl a-deficient mutation caused by reduced chlorophyll content and Chl a/b value, and a sharp reduction in protoporphyrin IX content impairs chlorophyll synthesis. Therefore, the whole plant always retains the mosaic leaves, and the photosynthetic capacity and utilization efficiency are lower than green leaves. There are physiological differences in photosynthesis between the different leaf colours.

Key words: Pseudosasa japonica f. akebonosuji, leaf color variation, photosynthetic pigment, photosynthetic system, photosynthetic characteristics

王妮, 李朝娜, 郑旭理, 姜思成, 杨海芸. 花叶矢竹叶片色素合成和光合特性. 植物生态学报, 2024, 48(11): 1536-1546. DOI: 10.17521/cjpe.2023.0258

WANG Ni, LI Zhao-Na, ZHENG Xu-Li, JIANG Si-Cheng, YANG Hai-Yun. Pigment synthesis and photosynthetic characteristics of leaves in Pseudosasa japonica f. akebonosuji. Chinese Journal of Plant Ecology, 2024, 48(11): 1536-1546. DOI: 10.17521/cjpe.2023.0258

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2023.0258

https://www.plant-ecology.com/CN/Y2024/V48/I11/1536

图/表 12

图1 花叶矢竹不同叶片类型和不同复绿阶段叶片。

Fig. 1 Different types of leaves and different degrees of return-green leaves of Pseudosasa japonica f. akebonosuji. AL0 (AL), albino leaf; AL2, albino leaf unfold for 2 months; AL6, albino leaf unfold for 6 months; AL12, albino leaf unfold for 12 months; GL, green leaf; SA, striped albino leaf; SG, striped green leaf.

表1 花叶矢竹不同叶片光合色素含量的变化(平均值±标准差)

Table 1 Variation of photosynthetic pigment content of Pseudosasa japonica f. akebonosuji (mean ± SD)

材料 Material	叶绿素a Chl a (µg·g^-1)	叶绿素b Chl b (µg·g^-1)	总叶绿素 Chl (a+b) (µg·g^-1)	类胡萝卜素 Car (µg·g^-1)	叶绿素a/b Chl a/b	类胡萝卜素/叶绿素 Car/Chl
绿叶 GL	408.9 ± 49.0^a	180.8 ± 24.3^a	589.7 ± 73.3^a	61.1 ± 12.1^a	2.3 ± 0.1^a	0.097 ± 0.021^c
条纹绿叶 SG	355.7 ± 17.5^a	195.3 ± 12.6^a	551.1 ± 21.1^a	50.8 ± 9.5^a	1.9 ± 0.4^b	0.091 ± 0.044^c
条纹白叶 SA	17.8 ± 3.1^b	10.7 ± 1.5^b	28.6 ± 4.6^b	8.8 ± 0.8^b	1.6 ± 0.1^b	0.338 ± 0.076^b
白叶 AL0	10.0 ± 1.0^b	5.7 ± 0.8^b	15.7 ± 1.7^b	7.8 ± 0.4^b	1.9 ± 0.5^b	0.534 ± 0.055^a

图2 花叶矢竹不同类型叶片间叶绿素合成前体物质相对含量(平均值±标准差)。设全绿叶(GL)中各产物含量为1, A、B、C分别为条纹绿叶(SG)、条纹白叶(SA)和全白叶(AL0)相对于绿叶(GL)的含量。ALA, δ-氨基乙酰丙酸; Chl, 叶绿素; Coprogen III, 粪卟啉原III; Mg-ProtoIX, Mg-原卟啉IX; PBG, 单卟啉胆色素; Pchlide, 原叶绿素酸酯; Proto IX, 原卟啉IX; Urogen III, 尿卟啉原III。不同小写字母表示不同类型叶片间差异显著(p < 0.05)。

Fig. 2 Relative content of precursor substances for chlorophyll synthesis of Pseudosasa japonica f. akebonosuji among different leaf types (mean ± SD). Supposing content of green leaf is 1, A, B, C are contents of striped green leaf (SG), striped albino leaf (SA) and albino leaf (AL0) relative to green leaf (GL). ALA, δ-aminolevulinic acid; Chl, chlorophyll; Coprogen III, coproporphyrinogen III; Mg-Proto, Mg-protoporphyrin IX; PBG, porphobilinogen; Pchlide, protochlorophyllide; Proto IX, protoporphyrin IX; Urogen III, uroporphyrino-gen III. Different lowercase letters indicate significant differences among different leaf types (p < 0.05).

图3 花叶矢竹不同类型叶片间的光响应曲线(平均值±标准差)。GL, 绿叶; SA, 条纹白叶; SG, 条纹绿叶。

Fig. 3 Light response curves of net photosynthetic rate (Pn) in Pseudosasa japonica f. akebonosuji among different leaf types (mean ± SD). GL, green leaf; SA, albino sector in leaf with strips; SG, green sector in leaf with strips. PAR, photosynthetically active radiation.

表2 花叶矢竹不同类型叶片间对光响应特性的影响(平均值±标准差)

Table 2 Effects of different leaf types on photosynthesis-light response parameters of Pseudosasa japonica f. akebonosuji (mean ± SD)

叶型 Leaf type	最大净光合速率 P_max(μmol·m^-2·s^-1)	表观量子效率 AQY (μmol·mol^-1)	暗呼吸速率 R_d(μmol·m^-2·s^-1)	光补偿点 LCP (μmol·m^-2·s^-1)	光饱和点 LSP (μmol·m^-2·s^-1)
绿叶 GL	13.70 ± 2.44^a	0.057 ± 0.005^a	1.18 ± 0.18^a	21.30 ± 2.02^b	424.50 ± 39.01^b
条纹绿叶 SG	8.65 ± 2.55^b	0.004 ± 0.006^b	0.97 ± 0.02^a	26.16 ± 3.75^b	489.30 ± 86.63^b
条纹白叶 SA	0.93 ± 0.01^c	0.003 ± 0.000^c	0.60 ± 0.12^b	260.02 ± 15.45^a	628.80 ± 42.80^a
白叶 AL0	-	-	-	-	-

表3 花叶矢竹不同类型叶片间CO2响应曲线模拟的光合参数(平均值±标准差)

Table 3 Photosynthesis parameters for CO2 response curve for different leaf types in Pseudosasa japonica f. akebonosuji (mean ± SD)

叶型 Leaf type	V_cmax(μmol·m^-2·s^-1 )	J_max(μmol·m^-2·s^-1)	J_max/V_cmax
绿叶 GL	86.22 ± 3.78^a	117.67 ± 4.36^a	1.37 ± 0.01^c
条纹绿叶 SG	85.60 ± 8.29^a	120.77 ± 13.87^a	1.41 ± 0.03^b
条纹白叶 SA	15.34 ± 1.58^b	30.61 ± 3.05^b	2.00 ± 0.01^a
白叶 AL0	-	-	-

图4 花叶矢竹不同类型叶片间叶绿素荧光诱导动力学曲线和相对可变荧光强度差值。AL0 (AL), 白叶; AL2, 白叶展开2个月; AL6, 白叶展开6个月; AL12, 白叶展开12个月; GL, 绿叶; SA, 条纹白叶; SG, 条纹绿叶。

Fig. 4 Dynamic curves for chlorophyll a fluorescein and the relative variable fluorescence intensity with different leaf types of Pseudosasa japonica f. akebonosuji. AL0 (AL), albino leaf; AL2, albino leaf unfold for 2 months; AL6, albino leaf unfold for 6 months; AL12, albino leaf unfold for 12 months; GL, green leaf; SA, striped albino leaf; SG, striped green leaf.

图5 花叶矢竹不同类型叶片间捕获的激子将电子传递到电子传递链中初级醌受体(QA)下游的电子受体的概率(Ψo)和以吸收光能为基础的性能指数(PIABS) (平均值±标准差)。AL0 (AL), 白叶; AL2, 白叶展开2个月; AL6, 白叶展开6个月; AL12, 白叶展开12个月; GL, 绿叶; SA, 条纹白叶; SG, 条纹绿叶。不同小写字母表示不同类型叶片间差异显著(p < 0.05)。

Fig. 5 Probability of that a trapped exciton the moves an electron further than QA by trapped exciton (Ψo) and performance index (PIABS) of Pseudosasa japonica f. akebonosuji under different leaf types (mean ± SD). AL0 (AL), albino leaf; AL2, albino leaf unfold for 2 months; AL6, albino leaf unfold for 6 months; AL12, albino leaf unfold for 12 months; GL, green leaf; SA, striped albino leaf; SG, striped green leaf. Different lowercase letters indicate significant differences among different leaf types (p < 0.05).

图6 花叶矢竹不同类型叶片间叶绿素Fk占F0 - Fj振幅的比例(Wk)和Fj占F0 - Fp振幅的比例(Vj) (平均值±标准差)。AL0 (AL), 白叶; AL2, 白叶展开2个月; AL6, 白叶展开6个月; AL12, 白叶展开12个月; GL, 绿叶; SA, 条纹白叶; SG, 条纹绿叶; F0, 初始荧光; Fj, J相荧光强度; Fk, K相荧光强度; Fp, P相荧光强度。不同小写字母表示不同类型叶片间差异显著(p < 0.05)。

Fig. 6 Proportion of Fk in F0 - Fj amplitude of chlorophyll (Wk) and proportion of Fj in F0 - Fp amplitude of chlorophyll (Vj) of Pseudosasa japonica f. akebonosuji under different leaf types (mean ± SD). AL0 (AL), albino leaf; AL2, albino leaf unfold for 2 months; AL6, albino leaf unfold for 6 months; AL12, albino leaf unfold for 12 months; GL, green leaf; SA, striped albino leaf; SG, striped green leaf; F0, minimal fluorescence; Fj, fluorescence intensity at J-step; Fk, fluorescence intensity at K-step; Fp, fluorescence intensity at P-step. Different lowercase letters indicate significant differences among different leaf types (p < 0.05).

图7 花叶矢竹不同类型叶片间光系统I (PSI) 820 nm相对吸收值变化(平均值±标准差)。AL0 (AL), 白叶; AL2, 白叶展开2个月; AL6, 白叶展开6个月; AL12, 白叶展开12个月; GL, 绿叶; SA, 条纹白叶; SG, 条纹绿叶。

Fig. 7 Changes of relative absorbtion of photosystem I (PSI) (PSI) 820 nm of different leaf types in Pseudosasa japonica f. akebonosuji (mean ± SD). AL0 (AL), albino leaf; AL2, albino leaf unfold for 2 months; AL6, albino leaf unfold for 6 months; AL12, albino leaf unfold for 12 months; GL, green leaf; SA, striped albino leaf; SG, striped green leaf.

图8 花叶矢竹不同类型叶片间光系统I (PSI)最大氧化还原能力(ΔI/Io)的差异(平均值±标准差)。AL0 (AL), 白叶; AL2, 白叶展开2个月; AL6, 白叶展开6个月; AL12, 白叶展开12个月; GL, 绿叶; SA, 条纹白叶; SG, 条纹绿叶。不同小写字母表示不同类型叶片间差异显著(p < 0.05)。

Fig. 8 Difference of photosystem I (PSI) maximum ability of oxido-reduction ΔI/Io of different leaf types in Pseudosasa japonica f. akebonosuji (mean ± SD). AL0 (AL), albino leaf; AL2, albino leaf unfold for 2 months; AL6, albino leaf unfold for 6 months; AL12, albino leaf unfold for 12 months; GL, green leaf; SA, striped albino leaf; SG, striped green leaf. Different lowercase letters indicate significant differences among different leaf types (p < 0.05).

图9 花叶矢竹不同类型叶片间光系统I (PSI)与光系统II (PSII)之间的协调性变化(ΦPSI/PSII) (平均值±标准差)。AL0 (AL), 白叶; AL2, 白叶展开2个月; AL6, 白叶展开6个月; AL12, 白叶展开12个月; GL, 绿叶; SA, 条纹白叶; SG, 条纹绿叶。不同小写字母表示不同类型叶片间差异显著(p < 0.05)。

Fig. 9 Changes of coordination between photosystem I (PSI) and photosystem II (PSII) of Pseudosasa japonica f. akebonosuji among different leaf types (ΦPSI/PSII) (mean ± SD). AL0 (AL), albino leaf; AL2, albino leaf unfold for 2 months; AL6, albino leaf unfold for 6 months; AL12, albino leaf unfold for 12 months; GL, green leaf; SA, striped albino leaf; SG, striped green leaf. Different lowercase letters indicate significant differences among different leaf types (p < 0.05).

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