不同光强对两种桤木幼苗光合特性和抗氧化系统的影响

doi:10.3724/SP.J.1258.2012.01062

摘要/Abstract

摘要：

通过搭建荫棚设置3种不同的光强, 模拟森林幼苗生长的旷地(砍伐迹地)、林窗和林下光照环境(分别为100%、56.2%和12.5%的全光照), 比较研究了外来种台湾桤木(Alnus formosana)和乡土种桤木(A. cremastogyne)幼苗的叶形态、光合能力、热耗散和抗氧化酶的活性, 探讨了两树种幼苗对光强的适应及光保护策略。结果表明: 在3种光强下, 一定光强范围内随着光强的增加, 两种桤木幼苗的比叶重(LMA)、类胡萝卜素(Cars)、类胡萝卜素/叶绿素(Cars/Chl)和抗氧化酶(超氧化物歧化酶(SOD)、过氧化氢酶(CAT)、抗坏血酸过氧化物酶(APX))活性升高, 最大净光合速率(P_max)、光饱和点(LSP)、光补偿点(LCP)和非光化学猝灭系数(NPQ)具有升高的趋势; Chl含量和瞬时光能利用效率(LUE)降低; 净光合速率(P_n)、气孔导度(G_s)、气孔限制值(L_s)升高, 胞间CO₂浓度(C_i)降低, 推测P_n降低的主要因素是非气孔限制, 表明两种桤木幼苗均能适应不同的生长光强。生长在相同光强下, 桤木幼苗光抑制现象比台湾桤木幼苗严重, 台湾桤木幼苗对光强适应能力较强。随着光照强度的增加, 台湾桤木幼苗NPQ增加不显著, 热耗散较少, 相同光强下P_max和抗氧化酶活性显著高于桤木幼苗, 而桤木幼苗随着光强的增加热耗散显著, 表明在光抑制时, 台湾桤木幼苗主要是通过提高P_max利用光能和抗氧化酶系统进行保护性调节, 桤木幼苗则通过天线系统非辐射耗散将过剩的光能以热能的形式消耗掉。

关键词: 桤木, 台湾桤木, 抗氧化酶, 热耗散, 光抑制, 光适应

Abstract:

Aims Our objective was to investigate the effects of different light intensities on leaf morphology, photosynthetic capacity, heat dissipation and antioxidant enzyme activities in seedlings of Alnus formosana and A. cremastogyne from a hilly region near Lingyan Mountain in northwestern Sichuan Province in China. We also discuss photosynthetic acclimation and photoprotection strategies in seedlings of the two species --Methods-- Three light regimes of 100%, 56.2% and 12.5% of natural light were simulated to match forest openings, forest gaps and forest canopies, respectively. After more than three months, we measured the parameters of gas exchange, including maximum net photosynthetic rate (Pmax), light saturation point (LSP), light compensation point (LCP), net photosynthetic rate (P_n), stomatal conductance (G_s), transpiration rate (T_r), intercellular CO₂concentration (C_i), fluorescent non-photochemical quenching (NPQ), maximum efficiency of PSII photochemistry (F_v/F_m) and light use efficiency (LUE), using a LI-6400 in seedling leaves. We also determined pigment contents per unit leaf area, lamina mass per unit area (LMA) and activity of antioxidant enzymes (superoxide dismutase, catalase and ascorbate peroxidase). --Important findings-- LMA, carotenoid content (Cars), ratio of carotenoid to total chlorophyll (Car/Chl) and activities of superoxide dismutase, catalase and ascorbate peroxidase in leaves increased with the increase of light intensities. P_max, LSP, LCP and NPQ tended to increase, while the chlorophyll content (Chl) and LUE decreased. However, the C_i decreased with the increase of P_n, G_s and stomatal limitation value (L_s). We speculated that non-stomatal limitation was the main factor that inhibited P_n. Seedlings of the two alder species could acclimate to different light regimes in this study through changing of physiological and morphological traits. Under all light regimes, diurnal photoinhibition of photosynthesis, as judged by F_v/F_m, was significantly more severe in A. cremastogyne than in A. formosana. The acclimation capacity to high light regime was stronger in A. formosana than in A. cremastogyne. With the increase of light intensities, P_maxand antioxidant enzyme activities increased significantly, but not NPQ in A. formosana. The opposite trends occurred in A. cremastogyne. At the same light intensity, thermal dissipation was much lower, but P_max was much higher in A. formosana than in A. cremastogyne. These results indicated that A. formosana seedlings might adapt resistance to photoinhibition through improving the use of solar energy by higher P_max and antioxidant enzyme system, while A. cremastogyne seedling avoided photoinhibition mainly through converting excess light energy to heat energy in the form of non-radiative dissipation through the antenna system.

Key words: Alnus cremastogyne, A. formosana, antioxidant enzyme, heat dissipation, photoinhibition, photosynthetic acclimation

刘柿良, 马明东, 潘远智, 魏刘利, 何成相, 杨开茂. 不同光强对两种桤木幼苗光合特性和抗氧化系统的影响. 植物生态学报, 2012, 36(10): 1062-1074. DOI: 10.3724/SP.J.1258.2012.01062

LIU Shi-Liang, MA Ming-Dong, PAN Yuan-Zhi, WEI Liu-Li, HE Cheng-Xiang, YANG Kai-Mao. Effects of light regimes on photosynthetic characteristics and antioxidant system in seedlings of two alder species. Chinese Journal of Plant Ecology, 2012, 36(10): 1062-1074. DOI: 10.3724/SP.J.1258.2012.01062

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链接本文: https://www.plant-ecology.com/CN/10.3724/SP.J.1258.2012.01062

https://www.plant-ecology.com/CN/Y2012/V36/I10/1062

图/表 10

图1 光强对两种桤木幼苗叶片的比叶重和色素含量的影响(平均值±标准误差)。LT1, 全光照; LT2, (56.2 ± 1.03)%全光照; LT3, (12.5 ± 0.21)%全光照。L, 光照; S, 树种; S × L, 树种和光照的交互作用。**, p < 0.01; *, p < 0.05。不同大写字母表示相同光照处理下不同树种之间差异显著(p < 0.05); 不同小写字母表示相同树种在不同光照处理之间差异显著(p < 0.05)。

Fig. 1 Effect of light regimes on the lamina mass per unit area (LMA) and pigment content in seedling leaves of two Alnus species (mean ± SE). LT1, full light; LT2, (56.2 ± 1.03)% full light; LT3, (12.5 ± 0.21)% full light. L, light; S, tree species; S × L, interaction of tree species and light. **, p < 0.01; *, p < 0.05. Different capital letters indicate significant differences among two tree species under the same light regimes (p < 0.05); Different small letters indicate significant differences among three light regimes of the same species (p < 0.05).

图2 不同光强下两种桤木幼苗光合作用的光响应曲线。LT1, 全光照; LT2, (56.2 ± 1.03)%全光照; LT3, (12.5 ± 0.21)%全光照。

Fig. 2 Light-photosynthetic response curves of two Alnus seedlings under different light regimes. LT1, full light; LT2, (56.2 ± 1.03)% full light; LT3, (12.5 ± 0.21)% full light.

表1 不同光强下两种桤木幼苗的最大净光合速率(Pmax)、光饱和点(LSP)和光补偿点(LCP) (平均值±标准误差)

Table 1 Maximum net photosynthetic rate (Pmax), light saturation point (LSP) and light compensation point (LCP) for the seedlings of two Alnus species under different light regimes (mean ± SE)

物种 Species	处理 Treatment	P_max (μmol CO₂ ?m^-2?s^-1)	LSP (μmol CO₂?m^-2?s^-1)	LCP (μmol CO₂ ?m^-2?s^-1)
台湾桤木 A. formosana	LT1	8.52 ± 0.45^{A, a}	1205.7 ± 104.0^{A, a}	98.05 ± 1.49^{B, a}
	LT2	7.48 ± 0.43^{A, a}	995.8 ± 55.4^{A, ab}	83.22 ± 1.76^{A, b}
	LT3	5.46 ± 0.27^{A, b}	755.9 ± 79.5^{A, b}	38.42 ± 3.12^{B, c}
桤木 A. cremastogyne	LT1	7.13 ± 0.05^{B, a}	877.3 ± 60.9^{A, a}	104.57 ± 3.02^{A, a}
	LT2	6.62 ± 0.30^{B, a}	749.4 ± 56.1^{B, a}	86.93 ± 5.04^{A, b}
	LT3	4.75 ± 0.18^{A, b}	489.6 ±16.0^{B, b}	84.69 ± 1.06^{A, b}

表2 不同光强下两种桤木幼苗叶片的气体交换参数和瞬时光能利用效率(平均值±标准误差)

Table 2 Gas exchange parameters and instantaneous light utilization efficiency in seedling leaves of two Alnus species under different light regimes (mean ± SE)

物种 Species	处理 Treatment	P_n (μmol CO₂·m^-2·s^-1)	T_{r (}mmol H₂O·m^-2·s^-1₎	G_s(mmol H₂O·m^-2 ·s^-1)	C_i (μmol CO₂?m^-2?s^-1)	LUE (mmol?mol^-1)	L_s
台湾桤木 A. formosana	LT1	7.32 ± 0.42^{A, a}	5.61 ± 0.21^{A, a}	236.91 ± 13.48^{A, a}	249.15 ± 15.91^{A, b}	8.81 ± 0.77^{A, b}	0.24 ± 0.01^{B, a}
台湾桤木 A. formosana	LT2	7.45 ± 0.32^{A, a}	4.12 ± 0.14^{A, b}	192.41 ± 4.20^{A, b}	281.85 ± 12.71^{A, ab}	13.11 ± 2.12^{B, b}	0.27 ± 0.01^{A, a}
	LT3	2.96 ± 0.36^{A, b}	2.93 ± 0.09^{A, c}	161.09 ± 13.27^{A, b}	337.01 ± 21.78^{B, a}	35.98 ± 1.11^{B, a}	0.14 ± 0.01^{A, b}
桤木 A.cremastogyne	LT1	6.66 ± 084^{B, a}	3.73 ± 0.49^{B, a}	196.32 ± 15.35^{A, a}	262.24 ± 10.86^{A, b}	9.63 ± 0.33^{A, b}	0.29 ± 0.01^{A, a}
桤木 A.cremastogyne	LT2	5.47 ± 0.78^{B, a}	2.89 ± 0.09^{B, a}	151.46 ± 11.87^{B, ab}	291.87 ± 7.97^{A, b}	15.25 ± 3.09^{A, b}	0.26 ± 0.02^{A, a}
	LT3	2.30 ± 0.45^{A, b}	1.71 ± 0.22^{B, b}	108.16 ± 25.93^{A, b}	340.35 ± 8.39^{A, a}	30.81 ± 3.70^{A, a}	0.15 ± 0.01^{A, b}

表3 两种桤木幼苗叶片气体交换参数和瞬时光能利用效率之间的相关性(n = 18)

Table 3 Correlation coefficients among gas exchange parameters and LUE in seedling leaves of two Alnus species (n = 18)

	P_n	T_r	G_s	C_i	L_s	LUE
P_n	1.00
T_r	0.77^**	1.00
G_s	0.72^**	0.87^**	1.00
C_i	-0.73^**	-0.80^**	-0.65^**	1.00
L_s	0.81^**	0.50^*	0.49^*	-0.73^**	1.00
LUE	-0.89^**	-0.66	-0.68^**	0.79^**	-0.89^**	1.00

图3 不同光强下两种桤木幼苗叶片光系统II最大光能转化效率(Fv/Fm)日变化(平均值±标准误差)。LT1, 全光照; LT2, (56.2 ± 1.03)%全光照; LT3, (12.5 ± 0.21)%全光照。

Fig. 3 Diurnal changes of maximum efficiency of PSII photochemistry (Fv/Fm) for the seedling leaves of two Alnus species under different light regimes (mean ± SE). LT1, full light; LT2, (56.2 ± 1.03)% full light; LT3, (12.5 ± 0.21)% full light.

图4 不同光强下两种桤木幼苗叶片荧光非光化学猝灭系数(NPQ)日变化(平均值±标准误差)。LT1, 全光照; LT2, (56.2 ± 1.03)%全光照; LT3, (12.5 ± 0.21)%全光照。

Fig. 4 Diurnal changes of non-photochemical quenching (NPQ) for the seedling leaves of two Alnus species grown under different light regimes (mean ± SE). LT1, full light; LT2, (56.2 ± 1.03)% full light; LT3, (12.5 ± 0.21)% full light.

图5 不同光强下两种桤木幼苗叶片的丙二醛(MDA)和H2O2含量(平均值±标准误差)。LT1, 全光照; LT2, (56.2 ± 1.03)%全光照; LT3, (12.5 ± 0.21)%全光照。L, 光照; S, 树种; S × L, 树种和光照的交互作用。**, p < 0.01; *, p < 0.05; ns, p > 0.05。不同大写字母表示相同光照处理下不同树种之间差异显著(p < 0.05); 不同小写字母表示相同树种在不同光照处理之间差异显著(p < 0.05)。

Fig. 5 Malondialdehyde (MDA) and H2O2 contents in seedling leaves of two Alnus species under different light regimes (mean ± SE). LT1, full light; LT2, (56.2 ± 1.03)% full light; LT3, (12.5 ± 0.21)% full light. L, light; S, tree species; S × L, interaction of tree species and light. **, p < 0.01; *, p < 0.05; ns, p > 0.05. Different capital letters indicate significant differences among two tree species under the same light regimes (p < 0.05); Different small letters indicate significant differences among three light regimes of the same species (p < 0.05).

表4 两种桤木幼苗叶片中丙二醛(MDA)含量、过氧化氢(H2O2)含量、超氧化物歧化酶(SOD)活性、过氧化氢酶(CAT)活性和抗坏血酸过氧化物酶(APX)活性之间的相关性(n = 18)

Table 4 Correlation coefficients among malondialdehyde (MDA) content, H2O2 content, superoxide dismutase (SOD) activity, catalase (CAT) activity and ascorbate peroxidase (APX) activity in seedling leaves of two Alnus species (n = 18)

	MDA含量 MDA content	H₂O₂含量 H₂O₂ content	CAT活性 CAT activity	SOD活性 SOD activity	APX活性 APX activity
MDA含量 MDA content	1.00
H₂O₂含量 H₂O₂ content	0.20	1.00
CAT活性 CAT activity	0.15	0.82^**	1.00
SOD活性 SOD activity	-0.21	0.81^**	0.84^**	1.00
APX活性 APX activity	-0.04	0.80^**	0.91^**	0.89^**	1.00

图6 不同光强下两种桤木幼苗叶片抗氧化酶活性(平均值±标准误差)。APX, 抗坏血酸过氧化物酶; CAT, 过氧化氢酶; SOD, 超氧化物歧化酶。LT1, 全光照; LT2, (56.2 ± 1.03)%全光照; LT3, (12.5 ± 0.21)%全光照。L, 光照; S, 树种; S × L, 树种和光照的交互作用。**, p < 0.01; *, p < 0.05; ns, p > 0.05。不同大写字母表示相同光照处理下不同树种之间差异显著(p < 0.05); 不同小写字母表示相同树种在不同光照处理之间差异显著(p < 0.05)。

Fig. 6 Activities of antioxidant enzyme in seedling leaves of two Alnus species under different light regimes (mean ± SE). APX, ascorbate peroxidase; CAT, catalase; SOD, superoxide dismutase. LT1, full light; LT2, (56.2 ± 1.03)% full light; LT3, (12.5 ± 0.21)% full light. L, light; S, tree species; S × L, interaction of tree species and light. **, p < 0.01; *, p < 0.05; ns, p > 0.05. Different capital letters indicate significant differences among two tree species under the same light regimes (p < 0.05); Different small letters indicate significant differences among three light regimes of the same species (p < 0.05).

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