LIGHT-MODULATED PHYSIOLOGICAL RESPONSE TO OCEAN ACIDIFICATION IN PHAEODACTYLUM TRICORNUTUM
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摘要: 光和海洋酸化(CO2浓度升高)分别对海洋硅藻的光合能力具有不同程度的影响, 但两者的耦合响应被较少关注。研究以三角褐指藻作为实验材料, 测定了不同光强下CO2浓度升高对三角褐指藻的生长、净光合速率、生化组分、胞外碳酸酐酶(eCA)活性和核酮糖-1,5-二磷酸羧化/氧化酶(Rubisco)活性的影响。结果显示在低光下, CO2浓度对三角褐指藻的生长和净光合速率(Pn)并没有显著影响, 而在高光下, 具有明显的影响。无论是在高光或是低光下, eCA活性、叶绿素和可溶性蛋白的含量都随着CO2浓度的升高而降低。在低光下, 高浓度CO2 (HC)培养下的Rubisco活性分别是低浓度CO2 (LC)和中浓度CO2 (MC)的2.42和1.39倍, 而在高光下, HC培养下的Rubisco活性分别是LC和MC的6.72和3.45倍。以上结果表明硅藻能够通过调节光合生理特征和CCM运行中能量的分配来适应环境中光强和CO2浓度的变化。
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关键词:
- 光强 /
- 海洋酸化 /
- 三角褐指藻 /
- 碳酸酐酶 /
- 核酮糖-1,5-二磷酸羧化酶/氧化酶
Abstract: There is increasing evidence that different light intensities or ocean acidification (OA) induced by elevated atmospheric CO2 concentration can affect the photosynthetic capacity of marine diatom to different degrees, respectively however, little attention had been paid to their interaction on diatom. In this study, the growth rate, net photosynthetic rate (Pn), biochemical composition, extracellular carbonic anhydrase (eCA) activity, and Ribulose-1,5-bisphosphate carboxylase/oxygenase (RubiscO) activity were investigated when Phaeodactylum tricornutum was grown under different light intensities and CO2 concentrations. The results showed that the specific growth rates and Pn in P. tricornutum were not significantly affected by CO2 concentration under low light intensity (LL), whereas in presence of the high light intensity (HL), elevated CO2 concentration was beneficial to promote the increase of the rate of Pn. The eCA activity, chlorophyll content, and soluble protein content decreased with increase of CO2 concentration, regardless of the high or low light. Under LL, RubiscO activity of HC-grown algae was 2.42 and 1.39 times higher than that of LC- or Medium-CO2 (MC)-grown ones. However, RubiscO activity of HC-grown algae was 6.72 and 3.45 times greater than that of LC- or MC-grown ones under high light. These results indicate that the algae can adapt to changes of light intensity and CO2 concentrations in the environment by adjusting the allocation of energy during the operation of the CO2-concentrating mechanism and photosynthesis. -
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硅藻CCMs的运行是个耗能的过程, 其活性不仅取决于外部无机碳(Inorganic carbon, Ci)浓度, 还与光能的获取有着密切的联系[5, 18]。藻类的光合作用为Ci的转运提供能量, 对CCMs的活性具有直接的影响[19—21]。光和CO2浓度是2个重要的环境因子, 其对藻类的相互作用会影响藻类的光合生理和海洋生态环境[22]。在酸化和200 μmol photons/(m2·s)的光强下, 三角褐指藻能够减少CCMs运行过程中的能量支出, 并将这部分能量用于细胞的增长[16]。此外, CO2和光强的交互作用还会对浮游植物群落造成影响。当CO2浓度升高时, 低光强能够促进浮游植物群落的生长和碳的固定, 而在高光强下浮游植物群落初级生产力下降[23]。光和CO2对藻的生长和光合特性的影响是存在差异的, 涉及藻类对光和海洋酸化的响应机制还需要进一步探索[15, 22]。
本文利用海洋浮游硅藻的模式藻种三角褐指藻作为实验材料, 通过测定不同光强和CO2浓度下三角褐指藻光合生理、碳酸酐酶(Carbonic anhydrase, CA)和Rubisco活性的变化, 探讨光强和CO2浓度对于三角褐指藻CCM的调节作用。
1. 材料与方法
三角褐指藻bac-2 (Phaeodactylum tricornutum
Bohlin)取自中国科学院海洋研究所。藻细胞被培养在f/2加富的天然海水中, 温度为(20±1)℃, 设置高、低光的光强分别为160和50 μmol photos/(m2·s), 培养液的pH分别控制在7.80±0.05、8.00±0.05和8.20±0.05, 相当于CO2浓度分别为25 μmol/L (High carbon dioxide concentration, HC)、16 μmol/L (Medium carbon dioxide concentration, MC)和11 μmol/L (Low carbon dioxide concentration, LC)。三角褐指藻的接种起始密度为500 cell/mL, 通过添加饱和CO2海水以维持培养液中pH。培养至18—20代后用0.45 μm聚酰胺滤纸过滤收集用于实验。 1.1 海水碳酸盐化学
通过滴定法测定海水[温度: (20±1)℃; 盐度:30‰]的总碱度(Total alkalinity, TA)[24, 25]。通过TA、pH和硼酸浓度计算培养液中的碳酸碱度[26], CO2的浓度通过碳酸碱度计算获得[27]。
1.2 生长测定
用血球计数板在显微镜下对藻细胞进行计数。比生长速率(Specific growth rate, SGR)通过以下公式计算: μ=(lnX2–lnX1)/(T2–T1), 其中, X1和X2分别代表三角褐指藻在T1和T2时的细胞密度。
1.3 光合作用测定
采用Clark型氧电极(YSI 5300A, YSI, USA)在400 μmol photons/(m2·s)的光强下测定藻细胞的净光合放氧速率。用恒温循环水浴连接反应槽, 使反应槽中的温度维持在(20±0.1)℃, 测定过程中三角褐指藻的细胞密度约为6.0×106 cell/mL。
1.4 可溶性蛋白含量测定
用考马斯亮蓝G-250染料结合法测定可溶性蛋白(Soluble protein, SP)含量 [28]。将培养样品在4℃, 12000×g的条件下离心3min收集藻细胞。在收集的藻细胞中加入5 mL的蒸馏水, 然后用超声波细胞破碎仪破碎藻细胞, 并在5000 r/min下离心10min。取1 mL上清液, 再加入5 mL考马斯亮蓝G-250溶液, 用紫外可见分光光度计(UV-1800, 岛津, 日本)测定其在595 nm处的吸光度, 再用标准曲线计算SP含量。用牛血清蛋白作为标准。
1.5 叶绿素含量测定
通过离心(12000×g, 5min)收集藻细胞, 弃上清液, 加入5 mL 90%的丙酮, 然后在4℃的暗环境中静置12h。提取液在20℃, 5000 r/min的条件下离心10min后, 取上清液, 于紫外可见分光光度计(UV-1800, 岛津, 日本)上测定各波长下的吸光度。根据以下公式计算藻细胞的Chl. a, c含量[29]:
Chl. a (μg/mL)=11.47×A664–0.40×A630
Chl. c (μg/mL)=24.36×A630–3.73×A664
式中, A630和A664分别代表在波长630和664 nm下的吸光度。
1.6 碳酸酐酶活性测定
CA活性用调整后的电量法进行测定[30]。将纯水置于4℃恒温循环水浴中, 通入高纯度CO2气体约60min, 制取CO2饱和水。将反应槽连接在恒温水浴循环器上, 使反应槽中的温度维持在4℃。取5 mL含藻细胞(6.0×106 cell/mL)的巴比妥溶液(20 mmol/L, pHnbs 8.3)置于反应槽中, 然后加入4 mL的CO2饱和水(4℃)测定pHnbs 8.3降至pHnbs 7.3所需的时间。CA活性通过以下公式测定: Units CA=10×(T0/T–1), 其中, T0和T分别表示不存在和存在酶的情况下pH从8.3下降到7.3所需的时间。
1.7 Rubisco活性测定
Rubisco活性测定参照Helbling等[31]的方法。离心收集藻细胞后, 将酶粗提取液加入到离心管中, 摇匀后将细胞置于4℃的冷水浴中用超声波进行破碎, 然后在12000×g, 4℃下离心15min, 取上清液。酶粗提的缓冲溶液(pHnbs8.00)包含50.00 mmol/L Tril-HCl、20.00 mmol/L MgCl2、0.20 mmol/L EDTA (Ethy-lenediaminetetraacetic acid, EDTA)和5.00 mmol/L Glutathione。Rubisco活性测定混合溶液(pHnbs8.00)中包含0.20 mmol/L还原性辅酶Ⅰ(Nicotinamide adenine dinucleotide, NADH)、3.00 mmol/L ATP (Adenosine triphosphate, ATP)、5.00 mmol/L磷酸肌酸(Phosphatecreatine, CP)、25.00 mmol/L碳酸氢钠(Sodium Bicarbonate, NaHCO3)、22.00 units磷酸肌酸激酶(Creatine phosphokinase, CPK)、18.00 units 3-磷酸甘油磷酸激酶(3-Phosphoglyceric phosphokinase)和9.00 units甘油醛-3-磷酸脱氢酶(Glyceraldehyde-3-phosphate dehydrogenase, GAPDH)。将酶粗提取液加入到Rubisco测定的混合溶液中, 然后用紫外可见分光光度计(UV-1800, 岛津, 日本)在20℃下测定NADH在340 nm下吸光度的变化。藻细胞Rubisco活性以mAbs 340/mg protein·s表示。
1.8 数据分析
应用软件Excel和Origin 8.51进行数据处理与分析, 用单因素方差分析(One-way ANOVA)和Student’s t-test分析不同实验处理间的差异性。直方图上不同字母表示在同一光强下不同浓度CO2处理之间的显著差异, 以P<0.05为差异显著水平。数值表示为平均值± SD,n=3。
2. 结果
2.1 生长
如图 1所示, 在低光强下, 不同的CO2浓度下培养的三角褐指藻比生长速率并没有存在显著的差异(P>0.05); 而在高光强下, HC条件下培养的三角褐指藻比生长速率比MC和LC条件下培养的分别增加了2.20% (P<0.05)和3.2% (P>0.05)。
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图 1 不同光强和CO2浓度对三角褐指藻比生长速率的影响Figure 1. Effect of light intensity and CO2 concentration on specific growth rate of Phaeodactylum tricornutum2.2 净光合速率
在低光强下, 不同浓度CO2下培养的三角褐指藻的净光合速率并无显著差异(P>0.05) (图 2)。而在高光强下培养的三角褐指藻的净光合速率随着CO2浓度的增加而显著升高(P<0.05), 在HC下培养的三角褐指藻的净光合速率分别是在MC和LC下培养的1.37 (P>0.05)和1.78倍(P<0.05) (图 2)。
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图 2 不同光强和CO2浓度对三角褐指藻净光合速率的影响Figure 2. Effect of light intensity and CO2 concentration on net photosynthetic rate of Phaeodactylum tricornutum2.3 叶绿素含量
在低光强下, 不同浓度CO2下培养的三角褐指藻Chl. a和Chl. c含量并无显著差异(P>0.05) (图 3)。在高光强下, LC下培养的Chl. a含量分别是MC和HC下培养的1.38 (P>0.05)和2.17倍(P<0.05); LC下培养的Chl.c含量分别是在MC和HC下培养的1.20 (P>0.05)和2.24倍(P<0.05) (图 3)。
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图 3 不同光强和CO2浓度对三角褐指藻Chl. a (a) 和Chl. c (b)含量的影响Figure 3. Effect of light intensity and CO2 concentration on the contents of Chl. a (a) and Chl. c (b) of Phaeodactylum tricornutum2.4 可溶性蛋白含量
在低光强下, 不同浓度CO2下培养的三角褐指藻可溶性蛋白含量并无显著差异(P>0.05) (图 4)。而在高光强下, MC和HC下培养的可溶性蛋白含量比在LC下培养的分别下降了20.0% (P<0.05)和28.8% (P<0.05) (图 4)。
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图 4 不同光强和CO2浓度对三角褐指藻可溶性蛋白含量的影响Figure 4. Effect of light intensity and CO2 concentration on soluble protein content of Phaeodactylum tricornutum2.5 碳酸酐酶活性
如图 5所示, 无论在高光强或低光强下, 三角褐指藻的胞外碳酸酐酶(eCA)活性均随CO2浓度升高而降低。在低光强下, MC和HC下培养的细胞其eCA活性分别是LC下培养的59.09% (P<0.05)和22.73% (P<0.05)。在高光强下, MC和HC下培养的eCA活性分别是LC下培养的62.71% (P<0.05)和39.98% (P<0.05) (图 5)。
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图 5 不同光强和CO2浓度对三角褐指藻eCA活性的影响Figure 5. Effect of light intensity and CO2 concentration on eCA activity of Phaeodactylum tricornutum2.6 Rubisco活性
如图 6所示, 在低光强下, Rubisco活性随CO2浓度增加而升高, 其均值分别为0.52、0.91和1.27×10–3 mAbs/μg(SP)·s, 其中在HC下培养的Rubisco活性分别是LC和MC下培养的2.42 (P<0.05)和1.39倍(P<0.05)。在高光强下Rubisco活性随CO2浓度增加而升高, 其均值分别为0.89、1.74和6.0×10–3 mAbs/μg(SP)·s, 其中HC下培养的Rubisco活性分别是LC和MC下培养的6.72 (P<0.05)和3.45倍(P<0.05)。
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图 6 不同光强和CO2浓度对三角褐指藻Rubisco活性的影响Figure 6. Effect of light intensity and CO2 concentration on Rubisco activity of Phaeodactylum tricornutum3. 讨论
3.1 生长与光合作用
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Sinutok等[36]的研究结果显示, 在光强250 μmol/(m2·s)和海洋酸化条件下Halimeda macroloba和Hali-meda cylindracea的叶绿素含量明显下降, 这与我们的结果是相似的。在低光强下, 藻细胞通过增加叶绿素含量来提高光的捕获能力, 为CCM的运行提供更充足的能量[16]; 而在高光下, 藻细胞通过减少色素含量以达到光保护的目的[37]。此外, 在酸化条件下CCMs活性被削弱, 减少了能量的消耗, 促使藻细胞通过减少叶绿素含量来避免过度的光能吸收以达到光保护的目的[34]。叶绿素和SP的合成需要大量的N, 在低光下, ATP的合成减少, 导致藻细胞对N的同化能力受到限制并影响叶绿素和SP的合成, 从而削弱CO2浓度对叶绿素和SP含量的影响[38]。而在高光强下, CO2浓度的升高导致三角褐指藻SP的含量减少, 这是由于在酸化条件下, 更多的N被用于藻细胞生长过程中的氮代谢和结构同化[39, 40]。
3.2 酶活性
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图 1 不同光强和CO2浓度对三角褐指藻比生长速率的影响
Figure 1. Effect of light intensity and CO2 concentration on specific growth rate of Phaeodactylum tricornutum
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图 2 不同光强和CO2浓度对三角褐指藻净光合速率的影响
Figure 2. Effect of light intensity and CO2 concentration on net photosynthetic rate of Phaeodactylum tricornutum
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图 3 不同光强和CO2浓度对三角褐指藻Chl. a (a) 和Chl. c (b)含量的影响
Figure 3. Effect of light intensity and CO2 concentration on the contents of Chl. a (a) and Chl. c (b) of Phaeodactylum tricornutum
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图 4 不同光强和CO2浓度对三角褐指藻可溶性蛋白含量的影响
Figure 4. Effect of light intensity and CO2 concentration on soluble protein content of Phaeodactylum tricornutum
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图 5 不同光强和CO2浓度对三角褐指藻eCA活性的影响
Figure 5. Effect of light intensity and CO2 concentration on eCA activity of Phaeodactylum tricornutum
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