溶藻细菌S7溶藻特性、机理及影响因素的研究
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摘要
随着水体富营养化现象的日益严重，有害藻类水华频繁爆发，探索行之有效的抑藻和除藻途径极为迫切，溶藻细菌作为一种新的生物控藻手段显示出了广泛的应用前景。溶藻细菌一般分离自暴发水华的水体，这种土著细菌在工程应用中具有安全、特异、高效等特点。课题组从三峡库区暴发水华的河流中分离出一株溶藻细菌，命名为S7，本文以导致水华暴发的代表性蓝绿藻为研究对象，研究了该菌的溶藻特性、探讨了溶藻机理并分析了溶藻效应的影响因素。论文的主要研究内容和结论如下：
①取生长24h的细菌S7，分别将不同体积的细菌添加到藻液中进行溶藻试验，结果表明：溶藻作用的发生对投加细菌的体积分数有个下限的要求，初始体积分数越大溶藻现象越明显，溶藻时间也越短。当细菌的体积分数大于1:50时，细菌S7在第5d表现出溶藻作用。
②细菌溶藻范围和作用效力试验表明：细菌S7对铜绿微囊藻、水华鱼腥藻、水华微囊藻、栅藻和小球藻均具有溶藻作用，但对水华鱼腥藻去除效果最好，对小球藻去除效果最差，叶绿素a的去除率为42.5%～74.8%。该菌对淡水湖泊中常见蓝绿藻有良好的去除效果。
③利用光学显微镜、扫描电镜观察溶藻过程，结果显示在溶藻过程中藻细胞依次经历了以下过程：细胞膜收缩→细胞膜凹陷→细胞膜形成抑痕→细胞膜破裂→细胞质和色素外泄→细胞死亡。在藻液中分别加入经不同方法处理后的菌液（包括未经处理的原菌液、高温灭菌20min的菌液、0.22μm滤膜过滤液、经10000rpm离心的上清液）进行溶藻试验，结果表明：细菌S7通过释放胞外活性物质进行间接溶藻。
④对细菌S7作用过程中藻细胞的叶绿素a和藻蓝蛋白含量进行监测，结果表明：随着藻细胞的急剧减少，藻细胞叶绿素a和藻蓝蛋白含量下降明显；对藻细胞抗氧化系统的变化进行分析，结果表明：在溶藻初期，超氧化物歧化酶(SOD)和过氧化氢酶（CAT）活性显著升高，表明细菌的溶藻作用激发了藻细胞的抗氧化系统参与反应，但随着细菌的继续作用，SOD和CAT活性下降，表明藻细胞抗氧化系统遭到破坏，丙二醛（MDA）含量也显著提高，说明细菌可以引起藻细胞膜脂过氧化；藻细胞经过PI特异荧光标记后，用流式细胞仪分析藻细胞自发荧光和特异荧光信号，结果表明：藻细胞的核酸物质和细胞膜均遭到破坏，叶绿素a和藻蓝蛋白含量变化与细胞膜的损伤情况具有一致性；藻细胞成分的红外光谱分析也显示藻细胞解体，蛋白质遭到破坏。据此推测细菌S7的溶藻机理为：溶藻物质先损坏藻体的细胞壁和粘质胶被，然后通过改变细胞膜的选择透过性进入藻细胞内部，引起藻细胞膜脂过氧化及活性氧的大量积累，溶藻物质进一步分解叶绿素a，破坏蛋白质，核酸等物质，进而造成藻体正常生理功能的丧失，最终导致细胞破裂死亡。
⑤通过萃取、醇沉、透析、硅胶柱层析对细菌S7分泌的溶藻物质进行分离纯化并利用紫外光谱、红外光谱和荧光光谱对该溶藻活性物质进行研究，结果表明：该活性物质的分子量小于8kDa；甲醇洗脱组分对铜绿微囊藻的溶藻效果最好；对该组分进行200～900nm的紫外光谱扫描，其洗脱组分在330nm处有高强度的吸收峰，表明其中的溶藻活性物质可能含有3～5个共轭单位；红外光谱扫描的结果表明溶藻物质可能含有羟基、饱和烃基和芳香醚；荧光光谱分析结果表明溶藻物质可能具有荧光属性，三维荧光光谱图有两个较强的荧光峰，峰值分别出现在264/436nm和378/369nm,且264/436nm峰荧光强度大于378/369nm峰；该物质可能是一种新的溶藻物质。
⑥单因素试验结果表明：温度、光照和pH值对溶藻效应有较大影响，通过响应曲面Box-Behnken试验对细菌S7溶藻效应的环境因素进行了研究，得出细菌S7对水华鱼腥藻溶藻效应3个环境因子的二次多项式模型，模型显著性高，拟合度好。通过响应面分析得到优化组合，即在培养温度33.06℃，光照强度3312lux和pH为6.26时，溶藻细菌S7对水华鱼腥藻溶藻效果最佳，理论最大去除率为91.40%。
⑦通过响应曲面Central-Composit试验得出了藻细胞去除率与细菌浓度、水体叶绿素a浓度关系的二次多项式模型，为溶藻细菌的工程应用提供了理论参考。从三峡库区临江河回水段试验区采水样进行控藻试验，结果表明：细菌S7可以提高水体透明度、改善水质环境，细菌作用7d对藻细胞叶绿素a去除率为68.27%，说明溶藻细菌可以用于环境水体控制藻类水华，达到改善水质的目的。
In recent years, with eutrophication phenomenon is increasing seriously, theharmful algal blooms break out frequently, so the effective algal inhibition and removalof algae way were explored extremely urgent. Algae-lysing bacteria as a newbiological control species means shows wide application prospect. Algae lysingbacteria isolated from the water bloom general hair, the indigenous bacteria inengineering application is safe, efficient, specific characteristics. This task group fromthe Three Gorges Reservoir blue algae river separates one algae-lysing bacteria,identified Chryseobacterium, named S7. The object of research is discuss the algicidalmechanisms of blue green algae bloom and analyzed the algicidal effect influencefactors. The main research contents and conclusions are as follows:
①Take growth of24h bacteria S7, different volume of the bacteria to the algaeliquid for algicidal experiments respectively. Results show that: The adding bacterialvolume has a lower requirement to algae lysing effect, the greater lytic phenomenon,the more obvious phenomeno and shorter lytic time.When the bacterial volumefraction is greater than1:50, bacterial S7showed a lytic effect in the5d.
②The scope and effect of Bacterial lysing experiment shows: the bacterial S7onMicrocystis, Anabaena flos-aquae, bloom Microcystis, Scenedesmus and Chlorella hasalgicidal effect, but best removal effect on the Chlorella vulgaris and worst,chlorophyll a removal with the removal rate is42.5%～74.8%. The result showed thatthe bacteria in freshwater lakes have good removal effect on cyanobacteria and greenalgae.
③The optical microscopy was use to scanning electron microscopy lytic process,results show that algal cells in the lytic process which experiences the followingprocesses: cell membrane contraction, cell membrane, cell membrane formation inmark, rupture of the cell membrane, and cytoplasm and pigment leakage and cell death.In algae solution are added and treated with different methods of bacteria (includinguntreated raw liquid, high temperature sterilization20min bacterial liquid,0.22μmmembrane filtration, the supernatant by10000rpm) of algicidal experiments, theresults show that: the bacterial S7through the extracellular release of activesubstances for indirect algicidal.
④By monitoring on bacterial S7of algae chlorophyll a and phycocyanin content,find that: with the algae decreased dramatically, algae chlorophyll a and algae blueprotein content decreased significantly; on algal cell changes in antioxidative systemwere analyzed, the results show that: in the early lytic, superoxide dismutase enzyme(SOD) and catalase (CAT) activity were significantly increased, suggesting thatbacterial stimulated algal cell antioxidant system involved in the reaction, but with thecontinuing role of bacteria, SOD and CAT activity decreased, indicating cellantioxidant system was destroyed, malondialdehyde (MDA) content increasedsignificantly, bacteria that can cause algal cells membrane lipid peroxidation; algalcells after PI specific fluorescence markers, with flow cytometry analysis of algal cellautofluorescence and specific fluorescence signal, the results show that: the algal cellnucleic acid substances and cell membrane were destroyed, chlorophyll a and algaeblue protein changes associated with cell membrane injury has consistency; algae cellcomponents of infrared spectral analysis also shows that algae cell disintegration,protein damage. It can be inferred that the algae lysing bacteria S7algicidalmechanisms: algicidal substance to damage the algae cell wall and mucilage glue, andthen through changes in cell membrane permselectivity, cause algae cell membranelipid peroxidation and active oxygen accumulation, algicidal substances furtherdecomposition of chlorophyll a, destroy the protein, nucleic acid and other substances,thus causing the algae normal physiological function loss, ultimately leading to cellrupture and death.
⑤Through extraction, alcohol precipitation, dialysis, silica gel columnchromatography separation and purification method on bacterial S7secretion ofalgicidal substances were isolated and purified by ultraviolet spectroscopy, infraredspectroscopy and fluorescence spectroscopy of the lytic activity substances werestudied. The results showed that the active substances had a molecular weight less than8kDa; methanol elution fractions on Microcystis algae lysing effect is best; thefractions were200～900nm in the UV spectrum, the elution fractions in at330nm highintensity of absorption peak, indicating that the algicidal substances may contain3to5conjugated unit; infrared spectrum scanning results showed active material maycontain hydroxyl, unsaturated alkyl and aryl ether; fluorescence spectra showed thatalgicidal substances may have fluorescent properties, three-dimensional fluorescencespectrum has two strong fluorescence peak, the peaks appeared at264/436nm and 378/369nm, and264/436nm fluorescence intensity greater than378/369nm peak; thesubstance may be a new kind of algicidal substances.
⑥The single factor experiments showed that the temperature、illumination andpH has great effect on lytic influence; Box-Behnken test on bacterial S7lytic effect ofenvironmental factors were studied, that bacterial S7on Anabaena flos-aquae lyticeffect of3environmental factors of two many model, model obviously high, fitwell.through response surface analysis optimized combination, namely in the culturetemperature33.06℃, light intensity of3312lux and pH6.26, algicidal bacteria S7onAnabaena flos-aquae lytic effect best, theoretical maximum removal rate of91.40%.
⑦Through Central-Composit test in the response surface find that two degreepolynomial model of algae lysing bacteria between he algae removal rate and bacterialconcentration, to provide theoretical reference for engineering applications. From theThree Gorges Reservoir in Linjiang River backwater period of Experimental Zonewater samples for algae control test, the results show that: the bacterial S7canimprove the transparency of the water body, improving the water quality of theenvironment, bacteria algae chlorophyll7d removal rate of a was68.27%, descriptionof algae lysing bacteria can be used for water environment control of algal bloom forthe purposep of improving water quality.
①取生长24h的细菌S7，分别将不同体积的细菌添加到藻液中进行溶藻试验，结果表明：溶藻作用的发生对投加细菌的体积分数有个下限的要求，初始体积分数越大溶藻现象越明显，溶藻时间也越短。当细菌的体积分数大于1:50时，细菌S7在第5d表现出溶藻作用。
②细菌溶藻范围和作用效力试验表明：细菌S7对铜绿微囊藻、水华鱼腥藻、水华微囊藻、栅藻和小球藻均具有溶藻作用，但对水华鱼腥藻去除效果最好，对小球藻去除效果最差，叶绿素a的去除率为42.5%～74.8%。该菌对淡水湖泊中常见蓝绿藻有良好的去除效果。
③利用光学显微镜、扫描电镜观察溶藻过程，结果显示在溶藻过程中藻细胞依次经历了以下过程：细胞膜收缩→细胞膜凹陷→细胞膜形成抑痕→细胞膜破裂→细胞质和色素外泄→细胞死亡。在藻液中分别加入经不同方法处理后的菌液（包括未经处理的原菌液、高温灭菌20min的菌液、0.22μm滤膜过滤液、经10000rpm离心的上清液）进行溶藻试验，结果表明：细菌S7通过释放胞外活性物质进行间接溶藻。
④对细菌S7作用过程中藻细胞的叶绿素a和藻蓝蛋白含量进行监测，结果表明：随着藻细胞的急剧减少，藻细胞叶绿素a和藻蓝蛋白含量下降明显；对藻细胞抗氧化系统的变化进行分析，结果表明：在溶藻初期，超氧化物歧化酶(SOD)和过氧化氢酶（CAT）活性显著升高，表明细菌的溶藻作用激发了藻细胞的抗氧化系统参与反应，但随着细菌的继续作用，SOD和CAT活性下降，表明藻细胞抗氧化系统遭到破坏，丙二醛（MDA）含量也显著提高，说明细菌可以引起藻细胞膜脂过氧化；藻细胞经过PI特异荧光标记后，用流式细胞仪分析藻细胞自发荧光和特异荧光信号，结果表明：藻细胞的核酸物质和细胞膜均遭到破坏，叶绿素a和藻蓝蛋白含量变化与细胞膜的损伤情况具有一致性；藻细胞成分的红外光谱分析也显示藻细胞解体，蛋白质遭到破坏。据此推测细菌S7的溶藻机理为：溶藻物质先损坏藻体的细胞壁和粘质胶被，然后通过改变细胞膜的选择透过性进入藻细胞内部，引起藻细胞膜脂过氧化及活性氧的大量积累，溶藻物质进一步分解叶绿素a，破坏蛋白质，核酸等物质，进而造成藻体正常生理功能的丧失，最终导致细胞破裂死亡。
⑤通过萃取、醇沉、透析、硅胶柱层析对细菌S7分泌的溶藻物质进行分离纯化并利用紫外光谱、红外光谱和荧光光谱对该溶藻活性物质进行研究，结果表明：该活性物质的分子量小于8kDa；甲醇洗脱组分对铜绿微囊藻的溶藻效果最好；对该组分进行200～900nm的紫外光谱扫描，其洗脱组分在330nm处有高强度的吸收峰，表明其中的溶藻活性物质可能含有3～5个共轭单位；红外光谱扫描的结果表明溶藻物质可能含有羟基、饱和烃基和芳香醚；荧光光谱分析结果表明溶藻物质可能具有荧光属性，三维荧光光谱图有两个较强的荧光峰，峰值分别出现在264/436nm和378/369nm,且264/436nm峰荧光强度大于378/369nm峰；该物质可能是一种新的溶藻物质。
⑥单因素试验结果表明：温度、光照和pH值对溶藻效应有较大影响，通过响应曲面Box-Behnken试验对细菌S7溶藻效应的环境因素进行了研究，得出细菌S7对水华鱼腥藻溶藻效应3个环境因子的二次多项式模型，模型显著性高，拟合度好。通过响应面分析得到优化组合，即在培养温度33.06℃，光照强度3312lux和pH为6.26时，溶藻细菌S7对水华鱼腥藻溶藻效果最佳，理论最大去除率为91.40%。
⑦通过响应曲面Central-Composit试验得出了藻细胞去除率与细菌浓度、水体叶绿素a浓度关系的二次多项式模型，为溶藻细菌的工程应用提供了理论参考。从三峡库区临江河回水段试验区采水样进行控藻试验，结果表明：细菌S7可以提高水体透明度、改善水质环境，细菌作用7d对藻细胞叶绿素a去除率为68.27%，说明溶藻细菌可以用于环境水体控制藻类水华，达到改善水质的目的。
In recent years, with eutrophication phenomenon is increasing seriously, theharmful algal blooms break out frequently, so the effective algal inhibition and removalof algae way were explored extremely urgent. Algae-lysing bacteria as a newbiological control species means shows wide application prospect. Algae lysingbacteria isolated from the water bloom general hair, the indigenous bacteria inengineering application is safe, efficient, specific characteristics. This task group fromthe Three Gorges Reservoir blue algae river separates one algae-lysing bacteria,identified Chryseobacterium, named S7. The object of research is discuss the algicidalmechanisms of blue green algae bloom and analyzed the algicidal effect influencefactors. The main research contents and conclusions are as follows:
①Take growth of24h bacteria S7, different volume of the bacteria to the algaeliquid for algicidal experiments respectively. Results show that: The adding bacterialvolume has a lower requirement to algae lysing effect, the greater lytic phenomenon,the more obvious phenomeno and shorter lytic time.When the bacterial volumefraction is greater than1:50, bacterial S7showed a lytic effect in the5d.
②The scope and effect of Bacterial lysing experiment shows: the bacterial S7onMicrocystis, Anabaena flos-aquae, bloom Microcystis, Scenedesmus and Chlorella hasalgicidal effect, but best removal effect on the Chlorella vulgaris and worst,chlorophyll a removal with the removal rate is42.5%～74.8%. The result showed thatthe bacteria in freshwater lakes have good removal effect on cyanobacteria and greenalgae.
③The optical microscopy was use to scanning electron microscopy lytic process,results show that algal cells in the lytic process which experiences the followingprocesses: cell membrane contraction, cell membrane, cell membrane formation inmark, rupture of the cell membrane, and cytoplasm and pigment leakage and cell death.In algae solution are added and treated with different methods of bacteria (includinguntreated raw liquid, high temperature sterilization20min bacterial liquid,0.22μmmembrane filtration, the supernatant by10000rpm) of algicidal experiments, theresults show that: the bacterial S7through the extracellular release of activesubstances for indirect algicidal.
④By monitoring on bacterial S7of algae chlorophyll a and phycocyanin content,find that: with the algae decreased dramatically, algae chlorophyll a and algae blueprotein content decreased significantly; on algal cell changes in antioxidative systemwere analyzed, the results show that: in the early lytic, superoxide dismutase enzyme(SOD) and catalase (CAT) activity were significantly increased, suggesting thatbacterial stimulated algal cell antioxidant system involved in the reaction, but with thecontinuing role of bacteria, SOD and CAT activity decreased, indicating cellantioxidant system was destroyed, malondialdehyde (MDA) content increasedsignificantly, bacteria that can cause algal cells membrane lipid peroxidation; algalcells after PI specific fluorescence markers, with flow cytometry analysis of algal cellautofluorescence and specific fluorescence signal, the results show that: the algal cellnucleic acid substances and cell membrane were destroyed, chlorophyll a and algaeblue protein changes associated with cell membrane injury has consistency; algae cellcomponents of infrared spectral analysis also shows that algae cell disintegration,protein damage. It can be inferred that the algae lysing bacteria S7algicidalmechanisms: algicidal substance to damage the algae cell wall and mucilage glue, andthen through changes in cell membrane permselectivity, cause algae cell membranelipid peroxidation and active oxygen accumulation, algicidal substances furtherdecomposition of chlorophyll a, destroy the protein, nucleic acid and other substances,thus causing the algae normal physiological function loss, ultimately leading to cellrupture and death.
⑤Through extraction, alcohol precipitation, dialysis, silica gel columnchromatography separation and purification method on bacterial S7secretion ofalgicidal substances were isolated and purified by ultraviolet spectroscopy, infraredspectroscopy and fluorescence spectroscopy of the lytic activity substances werestudied. The results showed that the active substances had a molecular weight less than8kDa; methanol elution fractions on Microcystis algae lysing effect is best; thefractions were200～900nm in the UV spectrum, the elution fractions in at330nm highintensity of absorption peak, indicating that the algicidal substances may contain3to5conjugated unit; infrared spectrum scanning results showed active material maycontain hydroxyl, unsaturated alkyl and aryl ether; fluorescence spectra showed thatalgicidal substances may have fluorescent properties, three-dimensional fluorescencespectrum has two strong fluorescence peak, the peaks appeared at264/436nm and 378/369nm, and264/436nm fluorescence intensity greater than378/369nm peak; thesubstance may be a new kind of algicidal substances.
⑥The single factor experiments showed that the temperature、illumination andpH has great effect on lytic influence; Box-Behnken test on bacterial S7lytic effect ofenvironmental factors were studied, that bacterial S7on Anabaena flos-aquae lyticeffect of3environmental factors of two many model, model obviously high, fitwell.through response surface analysis optimized combination, namely in the culturetemperature33.06℃, light intensity of3312lux and pH6.26, algicidal bacteria S7onAnabaena flos-aquae lytic effect best, theoretical maximum removal rate of91.40%.
⑦Through Central-Composit test in the response surface find that two degreepolynomial model of algae lysing bacteria between he algae removal rate and bacterialconcentration, to provide theoretical reference for engineering applications. From theThree Gorges Reservoir in Linjiang River backwater period of Experimental Zonewater samples for algae control test, the results show that: the bacterial S7canimprove the transparency of the water body, improving the water quality of theenvironment, bacteria algae chlorophyll7d removal rate of a was68.27%, descriptionof algae lysing bacteria can be used for water environment control of algal bloom forthe purposep of improving water quality.
引文
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