不同营养环境下海洋微藻生化组成与分子生物学响应特征研究
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摘要
本论文利用实时荧光定量PCR技术，研究了不同营养条件下培养的海洋微藻中四种功能基因（硝酸盐转运蛋白基因、磷酸盐转运蛋白基因、增殖细胞核抗原基因、核酮糖-1，5-二磷酸羧化氧化酶大亚基基因）表达量的变化，分析了这些基因与微藻生长及营养盐状况之间的关系，揭示了浮游植物对外界营养条件响应的分子机制，以期为近海富营养化评价的分子生物学指标体系的建立提供理论基础。研究的主要结果如下：
中肋骨条藻（Skeletonema costatum）硝酸盐转运蛋白基因（Nrt2）表达状况主要与培养液中硝酸盐浓度有关，高浓度硝酸盐抑制Nrt2基因的表达，而低浓度硝酸盐促进Nrt2基因的表达。一次性培养实验表明，当培养液中硝酸盐耗尽时，Nrt2基因表达量迅速升高，并在3天后达到最高值；通过对Nrt2基因表达量与培养液中硝酸盐浓度进行相关性分析发现，二者之间存在极显著负相关关系（P<0.01），并且当硝酸盐浓度低于40μM时，Nrt2基因表达量与硝酸盐浓度之间呈线性相关。在低磷条件下培养的微藻，Nrt2基因表达量显著降低。因此，Nrt2基因表达量是很好的指示微藻氮营养状况的指标。
磷酸盐转运蛋白基因（Pho）表达与伪矮海链藻（Thalassiosira pseudonana）磷营养状况相关。磷缺乏能极大促进Pho基因的表达，而磷添加能抑制Pho基因的表达，且无机磷添加组的表达量最低。相关性分析表明，Pho基因表达量与伪矮海链藻胞内总磷含量有极显著的负相关关系（P<0.01）。因此，Pho基因表达量有可能是指示微藻磷营养状况的良好指标。
不同氮磷浓度下培养的中肋骨条藻光合速率差异显著，光合作用速率随着培养液中的氮磷浓度升高而增大。同时对核酮糖-1，5-二磷酸羧化氧化酶大亚基基因（rbcL）表达量研究发现，rbcL仅与培养液中的硝酸盐浓度显著相关，而与培养液中磷酸盐浓度相关性不显著。这说明N、P营养盐对微藻光合作用的调控机制不同：N从转录水平上即对微藻光合作用产生影响，而P的影响主要体现在转录后水平上。rbcL基因表达量与光合作用速率存在极显著的正相关关系（P <0.01），有潜力成为指示微藻光合能力的分子指标。
中肋骨条藻增殖细胞核抗原基因（PCNA）表达量变化与生长状况相关，与生长率相关性极显著（P <0.01），这种关系为浮游植物现场生长率的测定提供了理论基础，将有可能对赤潮发展动态进行预测。
另外，本文还以我国主要有害藻华原因种东海原甲藻为研究对象，研究了其在不同营养条件下的核酸与蛋白含量变化，结果表明，DNA含量与东海原甲藻所处的生长阶段有关，对营养盐含量的变化不敏感；RNA含量则与N、P浓度均呈极显著相关性（P<0.01），且在不同的生长阶段其含量也会发生变化；RNA/DNA比值与N、P浓度均显著相关（P<0.05），且与生长率呈线性相关关系。培养液中N浓度显著影响到东海原甲藻单位细胞总蛋白含量，P缺乏也会使总蛋白含量显著降低。因此，可将RNA及RNA/DNA作为微藻营养状况的指标。
In this article, transcriptional changes of four functional genes—high affinitynitrate transporter gene (Nrt2), high affinity phosphate transporter gene (Pho),proliferating cell nuclear antigen gene (PCNA) and ribulose-1,5-bisphosphatecarboxylase/oxygenase large subunit gene (rbcL)—were studied in marine eukaryoticphytoplanktons when culture in different nutrient conditions by means of real-timefluorescence quantitative PCR. Relationships between the transcript levels and algaegrowth or nutrient conditions were analyzed. We aimed to elucidate the molecularmechanisms of phytoplankton response to environmental nutrient conditions. Themain results were as follows:
Nrt2transcript levels mainly correlated with the nitrate concentrations in themedium. Higher nitrate concentrations repress the transcript levels of Nrt2and lowernitrate concentrations promote the transcript levels. Results of batch cultures showedthat the Nrt2transcript levels increased rapidly when the nitrate was exhausted andreached the highest values3days after that. Correlation analysis showed that Nrt2transcript levels correlated negatively with the nitrate in the medium. Further more,there was a linear relationship between them when the nitrate was below40μM.Lower phosphate conditions would reduce the Nrt2transcript levels. All these resultsindicated that Nrt2was a good indicator of phytoplankton N conditions.
The Pho gene transcript levels correlated with the phosphorus status of algae. Pdeficiency would promote the Pho transcript levels remarkably, but P addition wouldrepress the transcript levels. The inorganic phosphate added group had the lowesttranscript levels. There was a significant negative relationship between Pho gene transcript levels and cellular phosphorus content of microalgae (P <0.01). Therefore,the Pho gene had the potential to be a good indicator of algal phosphorus status.
Significant differences existed among photosynthetic rates when the algae werecultured in different nutrient conditions. The photosynthetic rate increased with theincrement of nitrate and phosphate concentrations. Simultaneously detected rbcL genetranscript levels showed that, it correlated significantly with nitrate concentrations,but not with phosphate. This indicated the different regulation mechanism between Nand P on the algal photosynthesis: N may influence the photosynthesis from thetranscriptional level, however, the effect of P may from the post transcriptional level.Extreme significant correlations were found between rbcL gene transcript levels andphotosynthetic rates (P <0.01). rbcL gene maybe a good molecular indicator ofphytoplankton primary productions.
PCNA gene transcript levels changed in accordance with the growth of algae,and correlated significantly with the specific growth rates (P <0.01). The relationshipprovided theoretic basis of detecting in situ phytoplankton growth rate and maybeused in the forecast of harmful algae bloom developments.
In addition, responses of nucleic acid and protein content in batch cultures of themarine dinoflagellate Prorocentrum donghaiense were studied in relation to differentnitrate and phosphate concentrations. Results showed that DNA content varied indifferent growth stages, and it was insensitive to nutrient status. RNA content wasstrongly associated with both N and P concentrations (P<0.01). Different growthstages could also change cellular RNA content. The RNA content decreased alongwith the extension of experimental time. RNA/DNA had significant correlation withthe N and P concentrations. There appeared to be a linear relationship between growthrate and RNA/DNA. Cellular total protein content was significantly affected by Nconcentrations in the culture. P deficiency could also reduce cellular protein content.In conclusion, RNA and RNA/DNA could be used as algae nutrient status indicators.
中肋骨条藻（Skeletonema costatum）硝酸盐转运蛋白基因（Nrt2）表达状况主要与培养液中硝酸盐浓度有关，高浓度硝酸盐抑制Nrt2基因的表达，而低浓度硝酸盐促进Nrt2基因的表达。一次性培养实验表明，当培养液中硝酸盐耗尽时，Nrt2基因表达量迅速升高，并在3天后达到最高值；通过对Nrt2基因表达量与培养液中硝酸盐浓度进行相关性分析发现，二者之间存在极显著负相关关系（P<0.01），并且当硝酸盐浓度低于40μM时，Nrt2基因表达量与硝酸盐浓度之间呈线性相关。在低磷条件下培养的微藻，Nrt2基因表达量显著降低。因此，Nrt2基因表达量是很好的指示微藻氮营养状况的指标。
磷酸盐转运蛋白基因（Pho）表达与伪矮海链藻（Thalassiosira pseudonana）磷营养状况相关。磷缺乏能极大促进Pho基因的表达，而磷添加能抑制Pho基因的表达，且无机磷添加组的表达量最低。相关性分析表明，Pho基因表达量与伪矮海链藻胞内总磷含量有极显著的负相关关系（P<0.01）。因此，Pho基因表达量有可能是指示微藻磷营养状况的良好指标。
不同氮磷浓度下培养的中肋骨条藻光合速率差异显著，光合作用速率随着培养液中的氮磷浓度升高而增大。同时对核酮糖-1，5-二磷酸羧化氧化酶大亚基基因（rbcL）表达量研究发现，rbcL仅与培养液中的硝酸盐浓度显著相关，而与培养液中磷酸盐浓度相关性不显著。这说明N、P营养盐对微藻光合作用的调控机制不同：N从转录水平上即对微藻光合作用产生影响，而P的影响主要体现在转录后水平上。rbcL基因表达量与光合作用速率存在极显著的正相关关系（P <0.01），有潜力成为指示微藻光合能力的分子指标。
中肋骨条藻增殖细胞核抗原基因（PCNA）表达量变化与生长状况相关，与生长率相关性极显著（P <0.01），这种关系为浮游植物现场生长率的测定提供了理论基础，将有可能对赤潮发展动态进行预测。
另外，本文还以我国主要有害藻华原因种东海原甲藻为研究对象，研究了其在不同营养条件下的核酸与蛋白含量变化，结果表明，DNA含量与东海原甲藻所处的生长阶段有关，对营养盐含量的变化不敏感；RNA含量则与N、P浓度均呈极显著相关性（P<0.01），且在不同的生长阶段其含量也会发生变化；RNA/DNA比值与N、P浓度均显著相关（P<0.05），且与生长率呈线性相关关系。培养液中N浓度显著影响到东海原甲藻单位细胞总蛋白含量，P缺乏也会使总蛋白含量显著降低。因此，可将RNA及RNA/DNA作为微藻营养状况的指标。
In this article, transcriptional changes of four functional genes—high affinitynitrate transporter gene (Nrt2), high affinity phosphate transporter gene (Pho),proliferating cell nuclear antigen gene (PCNA) and ribulose-1,5-bisphosphatecarboxylase/oxygenase large subunit gene (rbcL)—were studied in marine eukaryoticphytoplanktons when culture in different nutrient conditions by means of real-timefluorescence quantitative PCR. Relationships between the transcript levels and algaegrowth or nutrient conditions were analyzed. We aimed to elucidate the molecularmechanisms of phytoplankton response to environmental nutrient conditions. Themain results were as follows:
Nrt2transcript levels mainly correlated with the nitrate concentrations in themedium. Higher nitrate concentrations repress the transcript levels of Nrt2and lowernitrate concentrations promote the transcript levels. Results of batch cultures showedthat the Nrt2transcript levels increased rapidly when the nitrate was exhausted andreached the highest values3days after that. Correlation analysis showed that Nrt2transcript levels correlated negatively with the nitrate in the medium. Further more,there was a linear relationship between them when the nitrate was below40μM.Lower phosphate conditions would reduce the Nrt2transcript levels. All these resultsindicated that Nrt2was a good indicator of phytoplankton N conditions.
The Pho gene transcript levels correlated with the phosphorus status of algae. Pdeficiency would promote the Pho transcript levels remarkably, but P addition wouldrepress the transcript levels. The inorganic phosphate added group had the lowesttranscript levels. There was a significant negative relationship between Pho gene transcript levels and cellular phosphorus content of microalgae (P <0.01). Therefore,the Pho gene had the potential to be a good indicator of algal phosphorus status.
Significant differences existed among photosynthetic rates when the algae werecultured in different nutrient conditions. The photosynthetic rate increased with theincrement of nitrate and phosphate concentrations. Simultaneously detected rbcL genetranscript levels showed that, it correlated significantly with nitrate concentrations,but not with phosphate. This indicated the different regulation mechanism between Nand P on the algal photosynthesis: N may influence the photosynthesis from thetranscriptional level, however, the effect of P may from the post transcriptional level.Extreme significant correlations were found between rbcL gene transcript levels andphotosynthetic rates (P <0.01). rbcL gene maybe a good molecular indicator ofphytoplankton primary productions.
PCNA gene transcript levels changed in accordance with the growth of algae,and correlated significantly with the specific growth rates (P <0.01). The relationshipprovided theoretic basis of detecting in situ phytoplankton growth rate and maybeused in the forecast of harmful algae bloom developments.
In addition, responses of nucleic acid and protein content in batch cultures of themarine dinoflagellate Prorocentrum donghaiense were studied in relation to differentnitrate and phosphate concentrations. Results showed that DNA content varied indifferent growth stages, and it was insensitive to nutrient status. RNA content wasstrongly associated with both N and P concentrations (P<0.01). Different growthstages could also change cellular RNA content. The RNA content decreased alongwith the extension of experimental time. RNA/DNA had significant correlation withthe N and P concentrations. There appeared to be a linear relationship between growthrate and RNA/DNA. Cellular total protein content was significantly affected by Nconcentrations in the culture. P deficiency could also reduce cellular protein content.In conclusion, RNA and RNA/DNA could be used as algae nutrient status indicators.
引文
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