广东典型海域微生物群落特征分析
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摘要
微生物在海洋生态系统的物质循环和转化过程中发挥着重要作用。本研究选择广东省两个典型海域：珠海桂山岛海域（受人类活动影响明显）和汕头南澳海域（海水养殖规模大），采用微生物群落水平生理特征（Community-level physiologicalprofiling，CLPP）分析和变性梯度凝胶电泳（Denatured gradient gel electrophoresis,DGGE）分析技术，从微生物群落的生理生化特征和分子生态两方面系统研究了两典型海区微生物群落功能和结构特征。进一步从龙须菜栽培体系分离筛选有益菌，将大型海藻修复与微生物修复相结合，初步探讨了龙须菜-有益菌对水质的协同净化效果，以期为养殖海区环境修复和海水养殖业的可持续发展提供基础数据和科学依据。
主要研究结果如下：
1.于2011年8月，采用CLPP和DGGE方法，研究了珠海桂山岛海域12个采样站点（S1~S12）水体和沉积物微生物群落结构和环境特征。结果表明，养殖区（S6、S7、S8）和受人类活动影响较大站点（S11、S12）表层水体微生物群落代谢活性和多样性指数高于其他站点。表层水体的主要优势菌群为变形细菌，以γ-变形菌和α-变形菌最多。除S10和S12站点外，其他站点的主要优势菌为交替假单胞菌Pseudoalteromonas sp.，属于γ-变形菌。表层沉积物17个主要优势菌分别属于γ-变形菌、-变形菌、拟杆菌、酸杆菌、放线菌等五大类，所占比例分别为27.8%、19.8%、23.1%、22.2%和7.0%。γ-变形菌为表层水体和表层沉积物的共有优势菌类。
基于CLPP和DGGE与环境因子的冗余分析（Redundancy analysis，RDA）发现总磷（Total Phosphorus，TP）和盐度是影响桂山岛海域表层水体微生物群落结构的重要因子。养殖区域和受人类活动影响较大的S6、S8、S11和S12站点的表层水体微生物群落结构受TP、盐度和TN的影响较大。表层沉积物中δ-变形菌所占的相对百分含量与沉积物中重金属Ni、Cr、Fe和Pb含量呈显著正相关（P﹤0.05）。
2.于2012年2月，对汕头南澳岛白沙湾养殖海域鱼类养殖区、龙须菜栽培区和对照区表层水体和表层沉积物微生物群落功能和结构进行了比较研究。结果表明，龙须菜栽培区水体微生物群落具有较高的代谢活性和遗传多样性。该海域表层水体优势细菌主要属于α-变形菌、黄杆菌、γ-变形菌、β-变形菌、厚壁菌、蓝细菌和放线菌等七大类。龙须菜栽培区主要优势菌种类与自然海区一致，为α-变形菌，所占比例分别为39.9%和34.5%，鱼类养殖区的主要优势菌种类为黄杆菌，所占比例为32.2%。龙须菜栽培区水体氮、磷和化学需氧量（Chemical Oxygen Demand，COD）含量均低于鱼类养殖区。
鱼类养殖区表层沉积物微生物群落具有较高的代谢活性，对聚合物、糖类、氨基酸、羧酸、胺类和其他等六大类碳源利用率均为最高，其次为龙须菜栽培区，对照区最低。该海域沉积物主要优势菌属于α-变形菌、γ-变形菌、δ-变形菌、ε-变形菌、酸杆菌、放线菌、厚壁菌、拟杆菌、浮霉菌等九大类群。不同功能区沉积物优势菌均为γ-变形菌和δ-变形菌，龙须菜栽培区这两类细菌所占的比例高达39.6%和19.9%。对照区、龙须菜栽培区和鱼类养殖区表层沉积物变形菌所占比例分别为60.1%，68.8%和66.8%。鱼类养殖区沉积物中总氮（Total Nitrogen，TN）、TP和总有机碳（TotalOrganic Carbon，TOC）含量显著高于龙须菜栽培区和对照区（P<0.05），微生物群落组成受沉积物中TOC、TP和TN的影响显著。
3.从龙须菜栽培环境和龙须菜体表分离、筛选了6株优势芽孢杆菌（3株短小芽孢杆菌GS6、GS11和GS12以及3株地衣芽孢杆菌GS4、EGW1和EGS4），分析了上述六株菌株对碳源的利用能力。据此选择短小芽孢杆菌GS11作为龙须菜-有益菌协同净化研究的实验菌株。
龙须菜-有益菌的协同净化实验表明，龙须菜-菌添加组和龙须菜-菌浸泡组（后称龙须菜-菌协同组）中的芽孢杆菌GS11均能在饲料浸出液生长；经过7d的培养，龙须菜-菌协同组龙须菜增重率达12.2%和11.7%，显著高于龙须菜组（P<0.05）。两种龙须菜-菌协同组均对饲料浸出液中的硝氮、亚硝氮、氨氮、磷酸盐和COD值有吸收或降解作用，尤其对氨氮、磷酸盐和COD的降解效果明显，降解率分别为51.3%和54.4%、63.0%和61.3%、53.6%和52.1%，均高于加菌组和龙须菜组。同时，两种龙须菜-菌协同组饲料浸出液的微生物群落代谢活性也显著高于对照组和龙须菜组（P<0.05）。综合考虑养殖海区的特点，可尝试采用有益菌菌液浸泡的方式，使有益菌与龙须菜协同净化海域环境。
Microorganisms play an important role in the material cycle and transformation process ofmarine ecosystem. Two typical sea areas of Guangdng Province, Guishan Island in Zhuhaiwhich was mainly affected by human activities and Nan’ao Island in Shantao whichoperated large-scale mariculture activities, were chosen in present study. The phylogeneticand metabolic features of microbial communities in these two areas were analyzed usingcommunity-level physiological profiling (CLPP) and denaturing gradient gelelectrophoresis (DGGE). In addition, potential probiotics were isolated and screened fromseaweed Gracilaria lemaneiformis cultivation system. Combining the bioremediationroles of seaweed and probiotic, the purification effects of Gracilaria-probioticcollaborative system on water quality was explored. These results will provide the datasupport and technical reference for environmental remediation and the sustainabledevelopment of mariculture.
The main results of the present research are as followed.
1. Microbial community structure and environmental characteristic in the water andsediment from12sampling sites (S1~S12) of Guishan Island were analyzed using CLPPand DGGE. The results showed that microbial community metabolic activity and diversityindex of the surface sea waters in the mariculture areas (S6, S7, S8) and the sites affectedby human activities (S11, S12) were higher than that in the other sites. The dominantbacteria in the surface water belonged to Proteobacteria, especially γ-Proteobacteria andα-Proteobacteria. Except for S10and S12, the dominant bacteria specie wasPseudoalteromonas sp., which belonged to γ-Proteobacteria. In the sediment,17dominantbacteria belonged to γ-Proteobacteria,-Proteobacteria, Bacteroidetes, Acidobacteria andActinobacteria, with percentages of27.8%,19.8%,23.1%,22.2%and7%, respectively.γ-Proteobacteria was the only common predominant group occurring both in surface waterand sediment.
The result of redundancy analysis (RDA) based on CLPP and DGGE withenvironmental factors showed that total phosphorus (TP) and salinity were the majorenvironmental factors influencing the microbial community structures in the surface waterin Guishan Island. The bacterial community structures in the mariculture areas and thesites affected by human activities (S6, S8, S11and S12) are greatly influenced by TP,salinity and TN. Furthermore, the relative percentage of-Proteobacteria in the sediment and the concentrations of heavy metals (Ni, Cr, Fe, Pb and so on) had significantlypositive correlation (P <0.05).
2. The phylogenetic and metabolic characteristics of microbial communities insurface water and sediment of Baisha Bay mariculture areas in Nan’ao Island wereexplored in February,2012. G. lemaneiformis cultivation zone (GZ), fish culture zone (FZ)and control zone (CZ) were chosen. The results showed that the bacterial communities inGZ had higher microbial metabolic capabilities and phylogenetic diversities. Forty-fourdominant bacterial species belonged to seven major phylogenetic groups (α-Proteobacteria,Flavobacteria, γ-Proteobacteria, β-Proteobacteria, Cyanobacteria, Firmicutes andActinobacteria). The most dominant bacteria in CZ and GZ were α-Proteobacteria (39.9%and34.5%), while that was Flavobacteria in FZ, with the percentage of32.2%. Moreover,the concentrations of nitrogen, phosphorus and COD in GZ were lower than that in FZ.
Microbial communities in surface sediment in FZ had the highest metabolic activitiesand utilization for six kinds of carbon source. The dominant bacterial species belonged tonine major phylogenetic groups (α-Proteobacteria, γ-Proteobacteria, δ-Proteobacteria,ε-Proteobacteria, Acidobacteria, Actinobacteria, Firmicutes, Bacteroidetes andPlanctomycetes). γ-Proteobacteria and δ-Proteobacteria were the predominant group, andthe percentage were39.6%and19.9%in GZ. The phylum of Proteobacteria in CZ, GZand FZ were account for60.1%,68.8%and66.8%, respectively. The concentrations of TN,TP and TOC in surface sediment of FZ were significantly higher than that of GZ and CZ(P<0.05). Microbial community composition was influenced by the TOC, TP and TN inthe sediments.
3. Six strains of Bacillus (three Bacillus pumilus GS6, GS11, GS12and three Bacilluslicheniformis GS4, EGW1, EGS4) were isolated and screened from Gracilaria cultivationsystem. According to the utilization of carbon source, B. pumilus GS11was selected forthe research on purifying effect of Gracilaria-probiotic collaborative system.
The purifying effect of Gracilaria-probiotic collaborative system showed that B.pumilus GS11could grow well in the feed leaching liquor in the group ofGracilaria-bacteria added directly and the group of bacteria immersed. The growth ratesof Gracilaria-bacteria collaborative group were significantly higher than the group ofGracilaria (P<0.05) after7d cultivation, with12.2%and11.7%, respectively. In thesetwo collaborative groups, nitrate, nitrite, ammonia, phosphate and COD in the feedleaching liquor were absorbed or degraded. The degradation effect on ammonia, phosphateand COD were significant in these two groups, with the percentage of51.3%and54.4%, 63.0%and61.3%,53.6%and52.1%, respectively. Furthermore, the microbial metaboliccapabilities in the feed leaching liquor in two collaborative groups were significantlyhigher than that of the control group and Gracilaria group (P<0.05). Given to thecharacteristics of mariculture ecosystem, Gracilaria cultivation system with probioticimmersed bacteria would be a possible way to improve the mariculture environment.
主要研究结果如下：
1.于2011年8月，采用CLPP和DGGE方法，研究了珠海桂山岛海域12个采样站点（S1~S12）水体和沉积物微生物群落结构和环境特征。结果表明，养殖区（S6、S7、S8）和受人类活动影响较大站点（S11、S12）表层水体微生物群落代谢活性和多样性指数高于其他站点。表层水体的主要优势菌群为变形细菌，以γ-变形菌和α-变形菌最多。除S10和S12站点外，其他站点的主要优势菌为交替假单胞菌Pseudoalteromonas sp.，属于γ-变形菌。表层沉积物17个主要优势菌分别属于γ-变形菌、-变形菌、拟杆菌、酸杆菌、放线菌等五大类，所占比例分别为27.8%、19.8%、23.1%、22.2%和7.0%。γ-变形菌为表层水体和表层沉积物的共有优势菌类。
基于CLPP和DGGE与环境因子的冗余分析（Redundancy analysis，RDA）发现总磷（Total Phosphorus，TP）和盐度是影响桂山岛海域表层水体微生物群落结构的重要因子。养殖区域和受人类活动影响较大的S6、S8、S11和S12站点的表层水体微生物群落结构受TP、盐度和TN的影响较大。表层沉积物中δ-变形菌所占的相对百分含量与沉积物中重金属Ni、Cr、Fe和Pb含量呈显著正相关（P﹤0.05）。
2.于2012年2月，对汕头南澳岛白沙湾养殖海域鱼类养殖区、龙须菜栽培区和对照区表层水体和表层沉积物微生物群落功能和结构进行了比较研究。结果表明，龙须菜栽培区水体微生物群落具有较高的代谢活性和遗传多样性。该海域表层水体优势细菌主要属于α-变形菌、黄杆菌、γ-变形菌、β-变形菌、厚壁菌、蓝细菌和放线菌等七大类。龙须菜栽培区主要优势菌种类与自然海区一致，为α-变形菌，所占比例分别为39.9%和34.5%，鱼类养殖区的主要优势菌种类为黄杆菌，所占比例为32.2%。龙须菜栽培区水体氮、磷和化学需氧量（Chemical Oxygen Demand，COD）含量均低于鱼类养殖区。
鱼类养殖区表层沉积物微生物群落具有较高的代谢活性，对聚合物、糖类、氨基酸、羧酸、胺类和其他等六大类碳源利用率均为最高，其次为龙须菜栽培区，对照区最低。该海域沉积物主要优势菌属于α-变形菌、γ-变形菌、δ-变形菌、ε-变形菌、酸杆菌、放线菌、厚壁菌、拟杆菌、浮霉菌等九大类群。不同功能区沉积物优势菌均为γ-变形菌和δ-变形菌，龙须菜栽培区这两类细菌所占的比例高达39.6%和19.9%。对照区、龙须菜栽培区和鱼类养殖区表层沉积物变形菌所占比例分别为60.1%，68.8%和66.8%。鱼类养殖区沉积物中总氮（Total Nitrogen，TN）、TP和总有机碳（TotalOrganic Carbon，TOC）含量显著高于龙须菜栽培区和对照区（P<0.05），微生物群落组成受沉积物中TOC、TP和TN的影响显著。
3.从龙须菜栽培环境和龙须菜体表分离、筛选了6株优势芽孢杆菌（3株短小芽孢杆菌GS6、GS11和GS12以及3株地衣芽孢杆菌GS4、EGW1和EGS4），分析了上述六株菌株对碳源的利用能力。据此选择短小芽孢杆菌GS11作为龙须菜-有益菌协同净化研究的实验菌株。
龙须菜-有益菌的协同净化实验表明，龙须菜-菌添加组和龙须菜-菌浸泡组（后称龙须菜-菌协同组）中的芽孢杆菌GS11均能在饲料浸出液生长；经过7d的培养，龙须菜-菌协同组龙须菜增重率达12.2%和11.7%，显著高于龙须菜组（P<0.05）。两种龙须菜-菌协同组均对饲料浸出液中的硝氮、亚硝氮、氨氮、磷酸盐和COD值有吸收或降解作用，尤其对氨氮、磷酸盐和COD的降解效果明显，降解率分别为51.3%和54.4%、63.0%和61.3%、53.6%和52.1%，均高于加菌组和龙须菜组。同时，两种龙须菜-菌协同组饲料浸出液的微生物群落代谢活性也显著高于对照组和龙须菜组（P<0.05）。综合考虑养殖海区的特点，可尝试采用有益菌菌液浸泡的方式，使有益菌与龙须菜协同净化海域环境。
Microorganisms play an important role in the material cycle and transformation process ofmarine ecosystem. Two typical sea areas of Guangdng Province, Guishan Island in Zhuhaiwhich was mainly affected by human activities and Nan’ao Island in Shantao whichoperated large-scale mariculture activities, were chosen in present study. The phylogeneticand metabolic features of microbial communities in these two areas were analyzed usingcommunity-level physiological profiling (CLPP) and denaturing gradient gelelectrophoresis (DGGE). In addition, potential probiotics were isolated and screened fromseaweed Gracilaria lemaneiformis cultivation system. Combining the bioremediationroles of seaweed and probiotic, the purification effects of Gracilaria-probioticcollaborative system on water quality was explored. These results will provide the datasupport and technical reference for environmental remediation and the sustainabledevelopment of mariculture.
The main results of the present research are as followed.
1. Microbial community structure and environmental characteristic in the water andsediment from12sampling sites (S1~S12) of Guishan Island were analyzed using CLPPand DGGE. The results showed that microbial community metabolic activity and diversityindex of the surface sea waters in the mariculture areas (S6, S7, S8) and the sites affectedby human activities (S11, S12) were higher than that in the other sites. The dominantbacteria in the surface water belonged to Proteobacteria, especially γ-Proteobacteria andα-Proteobacteria. Except for S10and S12, the dominant bacteria specie wasPseudoalteromonas sp., which belonged to γ-Proteobacteria. In the sediment,17dominantbacteria belonged to γ-Proteobacteria,-Proteobacteria, Bacteroidetes, Acidobacteria andActinobacteria, with percentages of27.8%,19.8%,23.1%,22.2%and7%, respectively.γ-Proteobacteria was the only common predominant group occurring both in surface waterand sediment.
The result of redundancy analysis (RDA) based on CLPP and DGGE withenvironmental factors showed that total phosphorus (TP) and salinity were the majorenvironmental factors influencing the microbial community structures in the surface waterin Guishan Island. The bacterial community structures in the mariculture areas and thesites affected by human activities (S6, S8, S11and S12) are greatly influenced by TP,salinity and TN. Furthermore, the relative percentage of-Proteobacteria in the sediment and the concentrations of heavy metals (Ni, Cr, Fe, Pb and so on) had significantlypositive correlation (P <0.05).
2. The phylogenetic and metabolic characteristics of microbial communities insurface water and sediment of Baisha Bay mariculture areas in Nan’ao Island wereexplored in February,2012. G. lemaneiformis cultivation zone (GZ), fish culture zone (FZ)and control zone (CZ) were chosen. The results showed that the bacterial communities inGZ had higher microbial metabolic capabilities and phylogenetic diversities. Forty-fourdominant bacterial species belonged to seven major phylogenetic groups (α-Proteobacteria,Flavobacteria, γ-Proteobacteria, β-Proteobacteria, Cyanobacteria, Firmicutes andActinobacteria). The most dominant bacteria in CZ and GZ were α-Proteobacteria (39.9%and34.5%), while that was Flavobacteria in FZ, with the percentage of32.2%. Moreover,the concentrations of nitrogen, phosphorus and COD in GZ were lower than that in FZ.
Microbial communities in surface sediment in FZ had the highest metabolic activitiesand utilization for six kinds of carbon source. The dominant bacterial species belonged tonine major phylogenetic groups (α-Proteobacteria, γ-Proteobacteria, δ-Proteobacteria,ε-Proteobacteria, Acidobacteria, Actinobacteria, Firmicutes, Bacteroidetes andPlanctomycetes). γ-Proteobacteria and δ-Proteobacteria were the predominant group, andthe percentage were39.6%and19.9%in GZ. The phylum of Proteobacteria in CZ, GZand FZ were account for60.1%,68.8%and66.8%, respectively. The concentrations of TN,TP and TOC in surface sediment of FZ were significantly higher than that of GZ and CZ(P<0.05). Microbial community composition was influenced by the TOC, TP and TN inthe sediments.
3. Six strains of Bacillus (three Bacillus pumilus GS6, GS11, GS12and three Bacilluslicheniformis GS4, EGW1, EGS4) were isolated and screened from Gracilaria cultivationsystem. According to the utilization of carbon source, B. pumilus GS11was selected forthe research on purifying effect of Gracilaria-probiotic collaborative system.
The purifying effect of Gracilaria-probiotic collaborative system showed that B.pumilus GS11could grow well in the feed leaching liquor in the group ofGracilaria-bacteria added directly and the group of bacteria immersed. The growth ratesof Gracilaria-bacteria collaborative group were significantly higher than the group ofGracilaria (P<0.05) after7d cultivation, with12.2%and11.7%, respectively. In thesetwo collaborative groups, nitrate, nitrite, ammonia, phosphate and COD in the feedleaching liquor were absorbed or degraded. The degradation effect on ammonia, phosphateand COD were significant in these two groups, with the percentage of51.3%and54.4%, 63.0%and61.3%,53.6%and52.1%, respectively. Furthermore, the microbial metaboliccapabilities in the feed leaching liquor in two collaborative groups were significantlyhigher than that of the control group and Gracilaria group (P<0.05). Given to thecharacteristics of mariculture ecosystem, Gracilaria cultivation system with probioticimmersed bacteria would be a possible way to improve the mariculture environment.
引文
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