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O3作为一种强氧化性的空气污染物,会对农作物和树木造成一定的伤害[1-3]。近地层O3浓度升高是由于其前体物(NOx和VOCs)的排放量增加所导致的[4-5]。近年来,虽然O3的峰值浓度已经得到了有效控制,但均值浓度仍然呈逐渐增加的趋势[6-7]。据预测,北半球中纬度的近地层的O3浓度将每年提高0.5%~2.0%[8]。
O3分子主要是通过叶表的气孔进入组织细胞,并发生反应生成活性氧(ROS),包括羟基自由基(·OH)、超氧阴离子(·O2-)和过氧化氢(H2O2)等[9]。O3会引起叶表出现伤害症状,改变植物的生理生化过程,抑制植物的生长[10-12]。研究表明O3浓度升高会降低三叶草的净光合速率(Pn)和气孔导度(Gs),最终导致生物量降低[10]。Calatayud等[13]的研究表明,植物的根生物量和根茎比在O3浓度升高的条件下会降低,这是由于光合作用产物对根系的分配降低所导致的。O3降低Pn的途径有两种:一种是气孔限制,主要通过降低Gs减少CO2的摄入量[14];另一种是非气孔限制,主要通过减少叶绿素(Chl)含量、降低羧化效率以及抑制电子传输,进而减少CO2的同化[15-16]。在非气孔限制为主要因素的情况下,植物通常会启动抗氧化机制,如提高抗坏血酸和总酚的含量[17]。它们是重要的抗氧化物质,可直接或者间接参与清除活性氧的代谢,被证明可以缓解O3对植物造成的伤害[18-19]。作为抗氧化酶系和非酶系(分子)的总水平,植物总抗氧化能力对O3胁迫的响应,往往与植物的敏感性相关[20]。
在北美和欧洲已经有很多树种,例如欧洲山杨(Populus tremula Michx. × Populus alba L. clone INRA 717-1B4)[21]、晚花稠李(Prunus serotina Ehrh.)[22]、美国白蜡(Fraxinus americana L.)[22]、红花槭(Acer rubrum L.)[22]、欧洲山毛榉(Fagus sylvatica)[23]和欧洲山杨(Populus tremula)[24]等,受到O3的危害。在中国,只有少数树种被用来研究对O3浓度升高的响应,且主要是北方树种[25-27]。然而少有关于中国亚热带常绿阔叶树种的研究。本研究的桢楠(Phoebe zhennan S. Lee et F. N. Wei)和闽楠(Phoebe bournei (Hemsl.) Yang),广泛分布于我国亚热带,为我国所特有,也是该区域常用的造林树种,具有较高的经济和生态价值。此外,桢楠和闽楠作为樟科楠属的不同种类植物,种间的生物学特性之间的差异是否会影响其对O3浓度升高的响应,有必要进行研究。本研究的目的是:(1)研究O3浓度升高对光合作用、抗氧化能力和生物量的影响;(2)比较两树种的O3敏感性并分析原因。
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本试验的实际熏气天数为114 d,NF、E1、E2的每日8 h O3浓度均值分别为26.9、99.7、147.0 nmol·mol-1(图 1a)。至试验结束,NF、E1、E2的AOT40(O3浓度超过40 nmol·mol-1的累积)分别为2.4、54.4、96.3 μmol·mol-1·h-1(图 1b)。
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除树种因素对胞间CO2浓度(Ci)无显著影响外,O3、树种、时间分别对所有测定的气体交换参数有显著影响(P < 0.05)(表 1)。O3×树种×时间对所有测定的气体交换参数有显著影响(P < 0.05)(表 1)。在9月份桢楠的测量中,E1对Pn无显著影响(图 2a)。除此之外,E1和E2显著降低了两树种3次测量的Pn(P < 0.05),且E2的Pn低于E1(除8月份闽楠)(图 2a)。Gs与蒸腾速率(Tr)对O3熏蒸的响应与Pn相似(图 2b和d)。E1显著提高了8月份桢楠的Ci,E2显著提高了8月份桢楠的Ci和9月份闽楠的Ci(P < 0.05)(图 2c)。E2显著降低了10月份桢楠的水分利用效率(WUE),显著降低了8月份和10月份闽楠的WUE(P < 0.05)(图 2e)。E1显著降低了8月份桢楠的气孔限制值(Ls),E2显著降低了8月份和10月份桢楠的Ls(P < 0.05)(图 2f)。
表 1 O3、树种、时间及其交互作用对各种测定参数影响的显著性分析
Table 1. Effects of O3, species, time and their interactions on measured indices
项目
ItemsO3 树种
Species时间
TimeO3×树种
O3×SpeciesO3×时间
O3×Time时间×树种
Time×SpeciesO3×时间×树种
O3×Time×Species净光合速率Pn *** ** *** *** *** *** ** 气孔导度Gs *** ** *** ns ns ns * 胞间CO2浓度Ci ** ns *** * ns ** ** 蒸腾速率Tr *** *** *** ns ns ns ** 水分利用效率WUE *** *** *** ** *** *** ** 气孔限制值Ls ** * *** ** ns *** ** 叶绿素Chl *** *** --- *** --- --- --- 类胡萝卜素Car *** *** --- *** --- --- --- 叶绿素/类胡萝卜素Chl/ Car ** ns --- ns --- --- --- 还原性抗坏血酸Reduced ascorbic acid *** *** --- ns --- --- --- 总酚Total phenols *** *** --- ns --- --- --- 总抗氧化能力Total antioxidant activity ** ns --- ns --- --- --- 叶干质量Foliage biomass *** ** --- * --- --- --- 茎干质量Stem biomass * ns --- ns --- --- --- 根干质量Root biomass *** * --- ns --- --- --- 总干质量Total biomass *** ns --- ns --- --- --- 根茎比Below/above ground biomass ratio ns *** --- * --- --- --- 注:***:在0.001水平下差异显著;**:在0.01水平下差异显著;*:在0.05水平下差异显著;ns:无显著性差;---:无时间因素 Note: ***, **, * means significant difference at 0.001, 0.01 and 0.05; ‘ns’ means no significance; ---means no time factor -
除树种因素对Chl/Car无显著影响外,O3和树种因素分别对Chl、Car含量和Chl/Car具有显著影响(P < 0.01)(表 1)。O3和树种的交互作用对Chl和Car含量具有显著影响(P < 0.001),这说明O3对Car和Chl含量的影响在两树种间具有显著性差异。E1和E2分别显著降低了桢楠的Chl含量23.7%和30.0%(P < 0.05)(图 3a)。E1显著降低了闽楠的Chl含量(P < 0.05)(图 3a)。两树种的Car与Chl含量表现出相似的变化趋势(图 3b)。O3熏蒸略微降低了两树种的Chl/Car,但不显著(图 3c)。
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除树种因素对总抗氧化能力无显著影响外,O3和树种因素分别对还原型抗坏血酸、总酚和总抗氧化能力水平具有显著影响(P < 0.01)(表 1)。O3和树种因素的交互作用对3种测定的抗氧化物质水平无显著影响(表 1)。O3熏蒸的两树种还原型抗坏血酸、总酚和总抗氧化能力水平高于NF处理(图 4a、b和c)。与NF相比,E1显著提高了桢楠的还原型抗坏血酸含量,显著提高了闽楠的还原型抗坏血酸和总酚含量(P < 0.05)(图 4a、b和c)。E2显著提高了桢楠的3种抗氧化物质水平,显著提高了闽楠的总酚含量(P < 0.05)(图 4a、b和c)。
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O3因素对所有测定的生物量指标具有显著影响(根茎比除外)(P < 0.05)(表 1)。树种因素对叶干质量、根干质量和根茎比具有显著影响(P < 0.05),但对总干质量和茎干质量的影响不显著(表 1)。O3和树种间的交互作用对叶干质量和根茎比有显著的影响(P < 0.05),说明O3对叶干质量和根茎比的影响在树种间有显著差异(表 1)。除E1对闽楠的茎干质量无显著影响外,E1和E2显著降低了两树种的根、茎、叶和总干质量(P < 0.05),但E1和E2之间无显著差异(图 5a和b)。随着O3浓度升高,闽楠的根茎比显著降低,而桢楠的根茎比变化不显著(图 5c)。
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AOT40与两树种的Pn、总干质量、叶、根干质量呈现显著的负相关关系(P < 0.01)(图 6a~d)。闽楠的Pn、总干质量、叶、根干质量减少速率均大于桢楠。闽楠的Pn-AOT40回归方程的斜率较桢楠减少了45.2%(图 6a)。在所有测定的生物量指标-AOT40的回归方程中,闽楠的叶生物量表现出与桢楠的最大差距,较桢楠减少了61.6%(图 6b)。
O3浓度升高对桢楠和闽楠幼苗光合作用、抗氧化能力及生物量的影响
Effects of Elevated O3 Level on Photosynthesis, Antioxidant Capacity and Biomass of Phoebe zhennan and Phoebe bournei Seedlings in Subtropical China
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摘要:
目的 探讨O3浓度升高对两种亚热带树木幼苗的影响,并分析其敏感性差异及原因。 方法 本试验以1年生桢楠和闽楠幼苗为材料,采用开顶式气室(OTCs),研究未过滤大气、100 nmol·mol-1(E1)、150 nmol·mol-1(E2)O3处理对光合作用、抗氧化能力和生物量的影响。 结果 研究表明:(1)O3熏蒸主要通过非气孔限制因素降低了两树种的净光合速率(Pn)。其中对于桢楠,E2对Pn的不利影响大于E1。而对于闽楠,8月份E2的Pn大于E1,而9月份和10月份小于E1。(2)O3熏蒸提高了两树种的抗氧化能力,表现为总酚含量和总抗氧化能力随着O3浓度的升高逐渐升高,而E2的还原型抗坏血酸含量低于E1。(3)O3熏蒸降低了两树种根、茎、叶及总干质量,及降低了闽楠的根茎比。 结论 O3熏蒸降低了两树种的光合作用,提高了抗氧化能力,最终减少了生物量。根据Pn和生物量,确定闽楠较桢楠对O3浓度升高更加敏感。两树种间的O3敏感性差异与气孔导度和抗氧化物质的背景水平及其对O3浓度升高的响应有关。 Abstract:Objective Considerable researches have documented the negative effects of O3 on woody species in North America and Europe, however, little is known about how woody tree species respond to elevated O3 in subtropical China. The objectives of this study are to investigate the effects of elevated O3 on two evergreen tree species in subtropical China, and compare the O3 sensitivity between them and explore potential mechanisms on differential responses to O3. Method One-year-old seedlings of Phoebe zhennan S. Lee et F. N. Wei and Phoebe bournei (Hemsl.) Yang were exposed to non-filtered air, 100 nmol mol-1 O3 air (E1) and 150 nmol mol-1 O3 air (E2) to investigate the O3 effects on photosynthesis, antioxidant capacity and biomass of these two species. Result The results are as follows:(1) O3 fumigation reduced net photosynthesis rate (Pn) mainly through non-stomatal factors in both species. Pn was lower under E2 than that under E1 in P. zhennan. Whereas for P. bournei, Pn was greater under E2 than that under E1 in August measurement, while the reverse patterns were observed in the measurements in September and October. (2) O3 fumigation increased measured antioxidant substances levels in both species. The levels of total phenols and total antioxidant capacity increased with O3 concentrations increasing. However, reduced ascorbic acid contents were lower under E2 than that under E1. (3) O3 fumigation decreased root, stem, foliage and total biomass in both species. The reduced below/above ground biomass ratio was only observed in P. bournei. Conclusion O3 fumigation decreases photosynthesis, increases the antioxidant capacity, and ultimately inhibits the growth of the two species. The O3 sensibility of P. bournei is stronger than that of P. zhennan based on Pn and biomass. The differential sensitivity to O3 is related to (i) stomatal conductance, and (ii) antioxidant substances responses to elevated O3 and their background level. -
Key words:
- O3
- / photosynthesis
- / antioxidant capacity
- / biomass
- / Phoebe zhennan
- / Phoebe bournei
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表 1 O3、树种、时间及其交互作用对各种测定参数影响的显著性分析
Table 1. Effects of O3, species, time and their interactions on measured indices
项目
ItemsO3 树种
Species时间
TimeO3×树种
O3×SpeciesO3×时间
O3×Time时间×树种
Time×SpeciesO3×时间×树种
O3×Time×Species净光合速率Pn *** ** *** *** *** *** ** 气孔导度Gs *** ** *** ns ns ns * 胞间CO2浓度Ci ** ns *** * ns ** ** 蒸腾速率Tr *** *** *** ns ns ns ** 水分利用效率WUE *** *** *** ** *** *** ** 气孔限制值Ls ** * *** ** ns *** ** 叶绿素Chl *** *** --- *** --- --- --- 类胡萝卜素Car *** *** --- *** --- --- --- 叶绿素/类胡萝卜素Chl/ Car ** ns --- ns --- --- --- 还原性抗坏血酸Reduced ascorbic acid *** *** --- ns --- --- --- 总酚Total phenols *** *** --- ns --- --- --- 总抗氧化能力Total antioxidant activity ** ns --- ns --- --- --- 叶干质量Foliage biomass *** ** --- * --- --- --- 茎干质量Stem biomass * ns --- ns --- --- --- 根干质量Root biomass *** * --- ns --- --- --- 总干质量Total biomass *** ns --- ns --- --- --- 根茎比Below/above ground biomass ratio ns *** --- * --- --- --- 注:***:在0.001水平下差异显著;**:在0.01水平下差异显著;*:在0.05水平下差异显著;ns:无显著性差;---:无时间因素 Note: ***, **, * means significant difference at 0.001, 0.01 and 0.05; ‘ns’ means no significance; ---means no time factor -
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