NaCl处理下两种引进红树的光合及抗氧化防御能力

doi:10.3724/SP.J.1258.2013.00046

摘要/Abstract

摘要：

在长期盐胁迫(28天, NaCl浓度从100 mmol·L^-1升至400 mmol·L^-1)下, 比较研究了引进的无瓣海桑(Sonneratia apetala)和拉关木(Laguncularia racemosa)幼苗叶片的气体交换、叶绿素含量、最大光化学效率(F_v/F_m)、O₂^-_·产生速率以及抗氧化酶的活性, 探讨了两种红树幼苗光合、抗氧化防御能力的差异与耐盐性的关系。结果显示: NaCl处理没有明显地影响两种红树幼苗的生长, 表明盐生植物对盐环境的适应性, 但两种红树的生理反应对NaCl处理存在较大的差异。在实验的第28天(苗木的NaCl累计处理浓度递增到400 mmol·L^-1)时, 与对照相比, 无瓣海桑叶片的净光合速率、水分利用效率增加, 气孔导度、蒸腾速率和胞间CO₂浓度/大气CO₂浓度(C_i/C_a)相应降低; 然而, 拉关木叶的净光合速率、蒸腾速率和水分利用效率均回落到对照的水平, 而气孔导度和C_i/C_a均增加, 表明同样的NaCl浓度处理对拉关木叶的净光合速率影响大于无瓣海桑。在NaCl处理期间, 无瓣海桑F_v/F_m一直保持在0.8以上, 而拉关木的F_v/F_m为0.75以下, 说明无瓣海桑具有高于拉关木的潜在最大光合能力。在实验的第7天(NaCl浓度为100 mmol·L^-1)和14天(苗木的NaCl累计处理浓度递增到200 mmol·L^-1)时, 两种红树O₂^-_·产生速率迅速增加, 在实验的第28天(苗木的NaCl累计处理浓度递增到400 mmol·L^-1)时, 无瓣海桑O₂^-_·产生速率是对照的5.3倍, 差异极显著, 此时, 拉关木叶中O₂^-_·产生速率已降低到低于对照的水平。盐处理诱导了两种红树叶中抗氧化酶(超氧化物歧化酶(SOD)、过氧化氢酶(CAT)、抗坏血酸过氧化物酶(APX)、谷胱甘肽还原酶(GR)、过氧化物酶(POD))活性增加, 但拉关木增加的幅度大于无瓣海桑, 表明拉关木能响应盐胁迫并上调抗氧化酶活性, 降低盐诱导的膜脂过氧化, 提高耐盐的能力, 无瓣海桑通过提高水分利用效率来保持体内的水分, 同时, 保持PSII的最大光化学量子产量, 使得无瓣海桑在高盐处理时仍能保持高于对照水平的光合速率。

关键词: 抗氧化酶, 气体交换, 拉关木, 盐胁迫, 无瓣海桑

Abstract:

Aims Sonneratia apetala, a native mangrove species in India, Bengal and Sri-Lanka, was introduced in 1985 to Dongzhaigang Mangrove Nature Reserve in Hainan Island, China from Sundarban, southwest of Bangladesh. Laguncularia racemosa, another mangrove species from Mexico was introduced to the same reserve in 1999. Our objective was to investigate the salinity tolerance mechanism of S. apetala and L. racemosa in order to elucidate adaptive strategies of the halophytes in stressful saline habitat.
Methods We investigated the effects of increasing soil NaCl (100-400 mmol·L^-1) on gas exchange, O₂^-_·production rate, activity of antioxidant enzymes, and the relevance to salt tolerance over four weeks in 1-year-old seedlings of S. apetala and L. racemosa.
Important findings Seedlings of the two mangrove species acclimated to different salinity levels through changing physiological and morphological traits. However, there were significant differences between S. apetala and L. racemosa in photosynthesis and anti-oxidant defense under salt stress. Increasing NaCl stress significantly elevated net photosynthetic rate (P_n), and stomatal conductance (G_s), intercellular CO₂concentration (C_i) and transpiration rate (T_r) decreased. As a result, water use efficiency (WUE) increased in leaves of S. apetala seedlings. But L. racemosa showed a rapid increase of P_n in the initiation of salt stress, and P_n remained lower than control levels at the end of the experiment. As a result, C_iand G_sincrease with the decrease of P_n, T_r and WUE. The reduction occurred after exposure of L. racemosa seedlings to severe salinity, 400 mmol·L^-1 NaCl (28 d). These results indicated that the inhibitory effects of severe salinity were more pronounced in L. racemosa under the same salinity. Moreover, the magnitude of variation of S. apetala maximum photochemical efficiency of PSII (F_v/F_m) was significantly less than that of L. racemosa leaves. We speculated that photosynthetic capacity of S. apetala was higher than L. racemosa. O₂^-_· production rate markedly increased after the two seedlings were subjected to 100 mmol·L^-1 NaCl and 200 mmol·L^-1 NaCl for 7 days and 14 days, respectively. However, O₂^-_· production rate in S. apetala leaves markedly increased upon increasing salinity and reached the highest level after seedlings were subjected to 400 mmol·L^-1 NaCl for 28 days, which was 5.3 fold of that in controls. In contrast to S. apetala, O₂ ^-_· production rate in L. racemosa leaves remained lower than control levels at the end of the experiment. Activity of superoxide dismutase (SOD), ascorbate peroxidase (APX), catalase (CAT), peroxidase (POD) and glutathione reductase (GR) was elevated corresponding to the increased O₂^-_·production in the salt-stressed two mangrove plants; however, the magnitude of increase of L. racemosa antioxidant enzyme activities was significantly greater than that of S. apetala, during the period of salt stress. We suggest that L. racemosa plants were able to sense salt stress and up-regulated the antioxidant enzymes to reduce salt-induced lipid peroxidation and membrane permeability, which contributed to maintenance of membrane integrity and salt tolerance in L. racemosa. Sonneratia apetala seedlings might adapt resistance to severe salinity through improving photosynthesis by higher WUE and maximum photochemical efficiency of PSII (Fv/Fm).

Key words: antioxidant enzyme, gas exchange, Laguncularia racemosa, salt stress, Sonneratia apetala

陈坚,李妮亚,刘强,钟才荣,黄敏,曾佳. NaCl处理下两种引进红树的光合及抗氧化防御能力. 植物生态学报, 2013, 37(5): 443-453. DOI: 10.3724/SP.J.1258.2013.00046

CHEN Jian,LI Ni-Ya,LIU Qiang,ZHONG Cai-Rong,HUANG Min,ZENG Jia. Antioxidant defense and photosynthesis for non-indigenous mangrove species Sonneratia apetala and Laguncularia racemosa under NaCl stress. Chinese Journal of Plant Ecology, 2013, 37(5): 443-453. DOI: 10.3724/SP.J.1258.2013.00046

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链接本文: https://www.plant-ecology.com/CN/10.3724/SP.J.1258.2013.00046

https://www.plant-ecology.com/CN/Y2013/V37/I5/443

图/表 6

表1 NaCl处理对两种红树气体交换的影响(平均值±标准误差, n = 20)

Table 1 Effect of NaCl treatment on gas exchange of two mangrove plants (mean ± SE, n = 20)

树种 Species	盐处理 NaCl treatment (d)	净光合速率 P_n (μmol CO₂·m^{- 2}·s^{- 1} )	气孔导度 G_s (mmol H₂O·m^{- 2}·s^{- 1})	蒸腾速率 T_r (mmol H₂O·m^{- 2}·s^{- 1})	水分利用效率 WUE (μmol CO₂·mmol^-1H₂O)	胞间CO₂浓度/ 大气CO₂浓度 C_i/C_a
无瓣海桑 Sonneratia apetala	对照 CK	7.813^a	247.083^a	5.240^a	1.491^a	0.820^a
无瓣海桑 Sonneratia apetala	7	6.135^a	309.917^b	3.282^B	1.870^B	0.897^B
	14	7.183^a	94.058^C	1.176^C	6.111^C	0.672^C
	21	8.757^a	146.183^D	2.192^D	3.995^D	0.723^D
	28	10.074^b	206.000^a	3.148^B	3.200^E	0.763^E
拉关木 Laguncularia racemosa	对照 CK	5.472^a	144.092^a	3.417^a	1.601^a	0.793^a
	7	7.914^B	202.200^b	2.120^b	3.730^B	0.802^a
	14	6.263^a	224.167^C	3.114^a	2.011^C	0.855^B
	21	6.025^a	170.167^b	2.939^a	2.050^c	0.828^C
	28	5.124^a	197.167^C	3.337^a	1.750^a	0.862^B

图1 NaCl处理下2种红树植物幼苗叶片叶绿素SPAD值和最大光化学效率(Fv/Fm) (平均值±标准误差, n = 20)。NaCl累计处理浓度: 1/4 Hoagland营养液(对照)、100 mmol·L-1 NaCl (7天)、200 mmol·L-1 NaCl (14天)、300 mmol·L-1 NaCl (21天)、400 mmol·L-1 NaCl (28天)。*, 对照和NaCl处理之间差异显著(p < 0.05); **, 对照和NaCl处理之间差异极显著(p < 0.01)。

Fig. 1 Maximum quantum efficiency of PSII photochemistry (Fv/Fm) and SPAD value of seedlings leaves of two mangrove plants under salt treatment (mean ± SE, n = 20). NaCl treatments: 1/4 strength Hoagland’s nutrient solution (CK), 100 mmol·L-1 NaCl (7 d), 200 mmol·L-1 NaCl (14 d), 300 mmol·L-1 NaCl (21 d)、400 mmol·L-1 NaCl (28 d). *, significant difference between control and NaCl treatments (p < 0.05); **, highly significant difference between control and NaCl treatments (p < 0.01).

图2 NaCl处理下无瓣海桑和拉关木叶片O2-· 产生速率和超氧化物歧化酶(SOD)活性的变化(平均值±标准误差, n = 3)。NaCl累计处理浓度: 1/4 Hoagland营养液(对照)、100 mmol·L-1 NaCl (7天)、200 mmol·L-1 NaCl (14天)、300 mmol·L-1 NaCl (21天)、400 mmol·L-1NaCl (28天)。*, 对照和NaCl处理之间差异显著(p < 0.05); **, 对照和NaCl处理之间差异极显著(p < 0.01)。

Fig. 2 Changes of O2-· production rate and superoxide dismutase (SOD) activity in leaves of Sonneratia apetala and Laguncularia racemosa under NaCl treatments (mean ± SE, n = 3). NaCl treatments: 1/4 strength Hoagland’s nutrient solution (CK), 100 mmol·L-1 NaCl (7 d), 200 mmol·L-1 NaCl (14 d), 300 mmol·L-1 NaCl (21 d)、400 mmol·L-1 NaCl (28 d). *, significant difference between control and NaCl treatments (p < 0.05); **, highly significant difference between control and NaCl treatments (p < 0.01).

图3 NaCl处理下无瓣海桑和拉关木叶片抗坏血酸过氧化物酶(APX)和过氧化氢酶(CAT)活性的变化(平均值±标准误差, n = 3)。NaCl累计处理浓度: 1/4 Hoagland营养液(对照)、100 mmol·L-1 NaCl (7天)、200 mmol·L-1 NaCl (14天)、300 mmol·L-1 NaCl (21天)、400 mmol·L-1 NaCl (28天)。*, 对照和NaCl处理之间差异显著(p < 0.05); **, 对照和NaCl处理之间差异极显著(p < 0.01)。

Fig. 3 Changes of ascorbate peroxidase (APX) and catalase (CAT) activity in leaves of Sonneratia apetala and Laguncularia racemosa under NaCl treatment (mean ± SE, n = 3). NaCl treatment: 1/4 strength Hongland’s nutrient solution (CK), 100 mmol·L-1 NaCl (7 d), 200 mmol·L-1 NaCl (14 d), 300 mmol·L-1 NaCl (21 d)、400 mmol·L-1 NaCl (28 d). *, significant difference between control and NaCl treatments (p < 0.05); **, highly significant difference between control and NaCl treatments (p < 0.01).

图4 NaCl处理对无瓣海桑和拉关木叶片谷胱甘肽还原酶(GR)和过氧化物酶(POD)活性的变化(平均值±标准误差, n = 3)。NaCl累计处理浓度: 1/4 Hoagland营养液(对照)、100 mmol·L-1 NaCl (7天)、200 mmol·L-1 NaCl (14天)、300 mmol·L-1 NaCl (21天)、400 mmol·L-1 NaCl (28天)。*, 对照和NaCl处理之间差异显著(p < 0.05); **, 对照和NaCl处理之间差异极显著(p < 0.01)。

Fig. 4 Changes of glutathione reductase (GR) and peroxidase (POD) activity in leaves of Sonneratia apetala and Laguncularia racemosa under NaCl treatment (mean ± SE, n = 3). NaCl treatment: 1/4 strength Hongland’s nutrient solution (CK), 100 mmol·L-1 NaCl (7 d), 200 mmol·L-1 NaCl (14 d), 300 mmol·L-1 NaCl (21 d)、400 mmol·L-1 NaCl (28 d). *, significant difference between control and NaCl treatments (p < 0.05); **, highly significant difference between control and NaCl treatments (p < 0.01).

表2 海南东寨港拉关木与海桑属植物生长表现比较

Table 2 Comparison of growth state for Laguncularia racemosa and Sonneratia in Dongzhai Harbor of Hainan Island, China

树种 Species	树龄 Age (a)	树高 Height (m)	地径 Ground diameter (cm)	胸径 Diameter at breast height (cm)	冠幅 Canopy (m × m)
拉关木 Laguncularia racemoca	5	6.3	14.4	8.5	2.1 × 2.6
无瓣海桑 Sonneratia apetala	5	9.2	20.8	12.9	4.5 × 4.2
海桑 S. caseolaris	5	6.6	16.5	10.2	2.8 × 2.4
杯萼海桑 S. alba	5	5.2	14.7	8.5	3.7 × 2.9
卵叶海桑 S. ovata	5	2.4	9.8	3.7	1.8 × 1.7

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