北大西洋典型海湾浮游异养细菌群落特征研究
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摘要
浮游异养细菌是海洋微食物网的重要组成部分,通过被高营养级生物的摄食,将可溶性有机物质转移到更高的营养级,在海洋生物地化循环中占有重要的地位。本论文以浮游异养细菌生物量和群落结构研究为切入点,利用海上现场调查结合分子生态学技术,研究了北大西洋典型海湾(Logy湾、Goose湾和Melville湖)浮游异养细菌生物量、生长率、摄食死亡率和群落结构的季节变化、空间分布特征以及春季水华期间浮游异养细菌生长和群落结构的时空变化,并进一步深入探讨了浮游异养细菌生长和群落结构变化与环境因子间的关系。该研究对于揭示浮游异养细菌生物量和群落结构的时空变化以及关键影响因素具有重要意义。
一、研究了Logy湾海水表层浮游异养细菌生物量和群落结构季节变化特征和主
要的环境影响因素,结果如下：
(1) Logy湾海水表层浮游异养细菌丰度和生物量均有显著的季节变化。冬季(2月)较低,峰值出现在4月和8月。海水表层浮游异养细菌平均细胞体积在春季最大。利用荧光原位杂交技术,研究了海水表层浮游异养细菌群落结构的季节变化。结果表明,海水表层浮游异养细菌群落结构具有较明显的季节变化。各个季节Alpha-Proteobacteria和Cytophage-Flavobacter (CF) cluster是海洋异养细菌群落中的优势类群,而Beta-Proteobacteria在海洋异养细菌群落中所占比例最低。各主要类群在群落中的相对丰度较高值均出现在春季(4月-6月),而在冬季(2月),相对丰度均较低。
(2)利用稀释法与荧光原位杂交相结合的方法,研究了海水表层浮游异养细菌及群落内主要类群的生长率和受微型浮游动物影响的摄食死亡率的季节变化。结果表明,总异养细菌及其各主要类群的生长率和摄食死亡率具有明显的季节变化,但变化趋势有所不同。总异养细菌的生长率和摄食死亡率夏季较高而冬季较低。Alpha-Proteobacteria在冬季生长率较高,其它类群的生长率和摄食死亡率则是夏季较高而冬季较低。
(3)相关性分析和回归分析结果表明,温度对该海域异养细菌的生长有显著的影响,可以解释40-60%总异养细菌和其主要类群生长率的变化。而叶绿素a浓度和DOC浓度则是影响浮游异养细菌丰度和细菌细胞体积的主要因素。浮游异养细菌与其群落内不同类群的生长率具有显著的相关性,表明异养细菌群落结构的变化与异养细菌群落功能的改变可能有着密切的联系。
二、研究了Goose湾和Mellive湖浮游异养细菌生物量和群落结构的分布特征和主要的环境影响因素,结果如下：
(1)该海域浮游异养细菌丰度在空间上具有显著的分布差异。平面分布上,由河口向远海具有增加的趋势。垂直分布上,浮游异养细菌丰度表层较高,在中部和底层较低。浮游异养细菌细胞体积大小和生物量平面分布上,由河口向远海分布并无显著差异。浮游异养细菌细胞体积大小随深度的增加表现出增大的趋势。浮游异养细菌生物量的垂直分布表现为随着深度的增加,逐渐减小。在远离河口的站位,中部和底部浮游异养细菌生物量的大小并无显著差别,而距河口较近的站位中,中部浮游异养细菌生物量高于表层。
(2)浮游异养细菌群落结构分析结果表明平面分布上Alpha-Proteobacteria和Beta-Proteobacteria表现出显著差异：距河口较近的站位Alpha-Proteobacteria在群落中相对丰度较低,而Beta-Proteobacteria在群落中相对丰度较高。距河口较远站位,Alpha-Proteobacteria在群落中相对丰度较高。垂向分布上,在距河口较近的站位,除CF cluster在中部表现出较高的相对丰度外,Alpha-,Beta-和Gamma-Proteobacteria随深度的增加,相对丰度均未表现出显著的差异。在距河口较远的站位,在垂直分布上,除Alpha-Proteobacteria以外,其他各主要类群在表层的相对丰度较高较高,而Alpha-Proteobacteria的相对丰度表现出较一致的垂直分布。
(3)表层浮游异养细菌及其主要类群的生长率和摄食死亡率空间分布的研究结果显示,不同站位的表层总浮游异养细菌生长率和死亡率均无显著差异。不同优势类群比较,Alpha-Proteobacteria在高盐度站位生长率较高,Beta-Proteobacteria和Gamma-Proteobacteria在低盐度站位表现出较高的生长率,而CF cluster在不同站位的生长率和死亡率均无显著差异。
(4)相关性分析和回归分析结果表明,温度、DOC和DON是影响浮游异养细菌群落丰度和生物量的主要因素,而深度和盐度的变化是影响浮游异养细菌细胞体积的主要因素。生长率与不同类群在异养细菌群落中所占比例变化具有显著的相关性,表明异养细菌对营养物质的利用是影响异养细菌群落结构的关键因素。
三、研究了Logy湾春季水华期间浮游异养细菌生物量和群落结构的变化,并探讨了其与环境因子变化之间的关系,初步明确了该海湾春季水华期间,浮游异养细菌群落结构变化的趋势和控制因素。
(1)浮游异养细菌群落的丰度和生物量的峰值都出现在水华的发展阶段。异养细菌细胞体积的峰值则出现在水华的消亡阶段。浮游异养细菌的群落结构的变化表现为,Alpha-Proteobacteria在水华的起始阶段相对丰度较高,而CF cluster在水华的消亡阶段表现出较高的相对丰度。
(2)海水表层浮游异养及群落内部不同类群的生长率表现出一定的差异性：在水华消亡阶段,除Gamma-Proteobacteria外,总浮游异养细菌及群落内主要类群均在水华的消亡阶段表现出较高的生长率。海水表层总浮游异养细菌群落及其各主要类群的摄食死亡率基本一致,较大值出现在水华的消亡阶段。
(3)相关性分析和回归分析结果表明,在水华期间,温度是调控该海域浮游异养细菌生长的关键因素。叶绿素a浓度和DOC浓度分别是控制浮游异养细菌生物量和细胞体积的关键因素。在水华过程中,环境中营养物质浓度的变化可能是控制浮游异养细菌群落结构的主要因素,而微型浮游动物的摄食则在水华的消亡时期调控着表层浮游异养细菌的群落结构。
Heterotrophic bacterioplankton plays an important role in marine biogeochemical cycles. Processes which mediate the transfer of carbon from dissolved pools to higher trophic levels are influenced by the growth and grazing losses of heterotrophic bacteria. The goals of this thesis are:to characterize seasonal and spatial variations in community structure of heterotrophic bacterioplankton in coastal (Logy Bay) and estuarine areas (Goose Bay and Lake Melville) of North Atlantic Ocean, including intensive sampling and experiments during the spring phytoplankton bloom in Logy Bay; to determine relationships among environmental factors, bacterial community structure, growth and grazing loss for bacterial population, as well as individual bacterial phylotypes; to provide insight into mechanism that lead to the observed variations at broad phylogenetic levels.
1.Seasonal changes in heterotrophic bacterial standing stock, community structure, rates of growth and grazing mortality for bacterioplankton, as well as individual phylogenetic groups within the community, are examined in the surface layer of Logy Bay. Factors controlling seasonal changes of these parameters are investigated using Pearson correlation and regression analysis.
(1) Heterotrophic bacterial abundance and biomass show high values in April and minimal in February, while cell volume exhibits maximum in May. Using fluorescence in situ hybridization (FISH), variations in composition of heterotrophic bacterial community were examined. Alpha-Proteobacteria and Cytophaga-Flavobacter cluster dominate the community overall, while Beta-Proteobacteria always comprise the lowest proportion of all detected phylogenetic groups. Relative abundances of all detected individual groups are low during winter (February and March), while higher values are detected during spring (April to June).
(2) Using FISH combined with dilution assay, seasonal changes in rates of growth and grazing mortality for bacterial community, as well as individual phylogenetic groups within the community, are determined. Growth rates for bacterial community as well as individual groups are high during summer (July and August), whereas low values are found during winter (February and March), with the exception of Alpha-Proteobacteria. Grazing mortality for bacterial community shows maximum during summer (July and August) and minimum during winter (March). However, no seasonal patterns of grazing mortality are observed among individual phylogenetic groups
(3) Results from Pearson correlation and regression analysis suggest that temperature is the main factor controlling rates of growth for bacteria, explained 40-60% variations in rates of growth for bacterial community as well as individual phylogenetic groups. Chlorophyll a concentration and dissolved organic carbon (DOC) are the controlling factors for bacterial abundance and cell volume, respectively. Correlations between rates of growth for bacterial community and those for individual groups implied bacterial phylogentic groups may share specific ecosystem function.
2.Spatial distributions in heterotrophic bacterial standing stock, community structure, rates of growth and grazing mortality for bacterioplankton, as well as individual phylogenetic groups within bacterial community, are examined in Goose Bay and Lake Melville. Factors related to spatial changes above are investigated using Pearson correlation and regression analysis.
(1) Bacterial abundance show distinct spatial distributions with minimal at low salinity waters and maximal at marine end of estuary. Vertical distribution of bacterial abundance show higher values at surface compared with those in the middle and at bottom. Cell volume and bacterial biomass show similar spatial patterns with each other, with no significant differences among stations. Cell volume increases with depth. Bacterial biomass show no significant differences in the middle and bottom layers at stations where salinity are low, while higher values are observed in the middle layer at stations in high salinity area.
(2) Analyses of heterotrophic bacterial community structure are performed at two selected sampling stations located at the low-(No. 111 station) and high-salinity (No.131 station) stations, respectively. Results indicate that relative abundance of Alpha-Proteobacteria is high at 131 station, whereas Beta-Proteobacteria is the dominant group at 111 station. Relative abundance of Alpha-, Beta-and Gamma-Proteobacteria uniformly distributed throughout the water column.
(3) Rates of growth and grazing mortality are determined using FISH combined with dilution assays at No. 111 station and No.131 station. No significant differences are observed between rates of growth for bacterial community at No. 111 station and No.131 station. Alpha-Proteobacteria growth rates are higher at No.131 station than those of other individual groups, while growth rates for Beta-Proteobacteria, as well as Gamma-Proteobacteria, are high at low-salinity station, No.111 station. No significant differences are found between grazing mortality for bacterial community at No.111 station and No. 131 station.
(4) Results from correlation and regression analysis indicate that dissolved organic carbon (DOC) and dissolved organic nitrogen (DON) are controlling factors for heterotrophic bacterial abundance and biomass. Depth and salinity are primary factors regulating cell volume. Since rates of bacterial growth indicates uptake of DOM, the co-occurrence of high growth rates and high relative abundance of individual phylotypes implies the possible regulations of nutrients uptake on bacterial community structure in this area.
3.Temporal and spatial variations in heterotrophic bacterial community structure are studied during spring phytoplankton bloom in Logy Bay to investigate possible mechanisms that lead to the observed variations in bacterial community composition during the bloom.
(1) Abundance and biomass of heterotrophic bacterioplankton show high values during the developmental stage of the spring bloom, while the maximal of cell volume is found during the post-bloom stage. Alpha-Proteobacteria reach its maximal during the beginning of spring bloom, while Beta-, Gamma-Proteobacteria and CF cluster reach maximum during the post-bloom stage.
(2) Using FISH combined with dilution assays, research indicate that significant differences in patterns of growth rates among bacterial community and detected phylogenetic groups, while patterns of grazing mortality are similar to each other during the study.
(3) Results from correlation and regression analysis indicate that temperature is the controlling factor for the growth of heterotrophic bacteria during the spring bloom, while chlorophyll a and DOC concentration are responsible for the changes of bacterial biomass and cell volume, respectively. Nutrient supplies might be the main factor driving variations in bacterial community structure during the spring bloom, while predation might be a key mechanism for regulation of bacterial community composition during the senescent stage of the bloom.
一、研究了Logy湾海水表层浮游异养细菌生物量和群落结构季节变化特征和主
要的环境影响因素,结果如下：
(1) Logy湾海水表层浮游异养细菌丰度和生物量均有显著的季节变化。冬季(2月)较低,峰值出现在4月和8月。海水表层浮游异养细菌平均细胞体积在春季最大。利用荧光原位杂交技术,研究了海水表层浮游异养细菌群落结构的季节变化。结果表明,海水表层浮游异养细菌群落结构具有较明显的季节变化。各个季节Alpha-Proteobacteria和Cytophage-Flavobacter (CF) cluster是海洋异养细菌群落中的优势类群,而Beta-Proteobacteria在海洋异养细菌群落中所占比例最低。各主要类群在群落中的相对丰度较高值均出现在春季(4月-6月),而在冬季(2月),相对丰度均较低。
(2)利用稀释法与荧光原位杂交相结合的方法,研究了海水表层浮游异养细菌及群落内主要类群的生长率和受微型浮游动物影响的摄食死亡率的季节变化。结果表明,总异养细菌及其各主要类群的生长率和摄食死亡率具有明显的季节变化,但变化趋势有所不同。总异养细菌的生长率和摄食死亡率夏季较高而冬季较低。Alpha-Proteobacteria在冬季生长率较高,其它类群的生长率和摄食死亡率则是夏季较高而冬季较低。
(3)相关性分析和回归分析结果表明,温度对该海域异养细菌的生长有显著的影响,可以解释40-60%总异养细菌和其主要类群生长率的变化。而叶绿素a浓度和DOC浓度则是影响浮游异养细菌丰度和细菌细胞体积的主要因素。浮游异养细菌与其群落内不同类群的生长率具有显著的相关性,表明异养细菌群落结构的变化与异养细菌群落功能的改变可能有着密切的联系。
二、研究了Goose湾和Mellive湖浮游异养细菌生物量和群落结构的分布特征和主要的环境影响因素,结果如下：
(1)该海域浮游异养细菌丰度在空间上具有显著的分布差异。平面分布上,由河口向远海具有增加的趋势。垂直分布上,浮游异养细菌丰度表层较高,在中部和底层较低。浮游异养细菌细胞体积大小和生物量平面分布上,由河口向远海分布并无显著差异。浮游异养细菌细胞体积大小随深度的增加表现出增大的趋势。浮游异养细菌生物量的垂直分布表现为随着深度的增加,逐渐减小。在远离河口的站位,中部和底部浮游异养细菌生物量的大小并无显著差别,而距河口较近的站位中,中部浮游异养细菌生物量高于表层。
(2)浮游异养细菌群落结构分析结果表明平面分布上Alpha-Proteobacteria和Beta-Proteobacteria表现出显著差异：距河口较近的站位Alpha-Proteobacteria在群落中相对丰度较低,而Beta-Proteobacteria在群落中相对丰度较高。距河口较远站位,Alpha-Proteobacteria在群落中相对丰度较高。垂向分布上,在距河口较近的站位,除CF cluster在中部表现出较高的相对丰度外,Alpha-,Beta-和Gamma-Proteobacteria随深度的增加,相对丰度均未表现出显著的差异。在距河口较远的站位,在垂直分布上,除Alpha-Proteobacteria以外,其他各主要类群在表层的相对丰度较高较高,而Alpha-Proteobacteria的相对丰度表现出较一致的垂直分布。
(3)表层浮游异养细菌及其主要类群的生长率和摄食死亡率空间分布的研究结果显示,不同站位的表层总浮游异养细菌生长率和死亡率均无显著差异。不同优势类群比较,Alpha-Proteobacteria在高盐度站位生长率较高,Beta-Proteobacteria和Gamma-Proteobacteria在低盐度站位表现出较高的生长率,而CF cluster在不同站位的生长率和死亡率均无显著差异。
(4)相关性分析和回归分析结果表明,温度、DOC和DON是影响浮游异养细菌群落丰度和生物量的主要因素,而深度和盐度的变化是影响浮游异养细菌细胞体积的主要因素。生长率与不同类群在异养细菌群落中所占比例变化具有显著的相关性,表明异养细菌对营养物质的利用是影响异养细菌群落结构的关键因素。
三、研究了Logy湾春季水华期间浮游异养细菌生物量和群落结构的变化,并探讨了其与环境因子变化之间的关系,初步明确了该海湾春季水华期间,浮游异养细菌群落结构变化的趋势和控制因素。
(1)浮游异养细菌群落的丰度和生物量的峰值都出现在水华的发展阶段。异养细菌细胞体积的峰值则出现在水华的消亡阶段。浮游异养细菌的群落结构的变化表现为,Alpha-Proteobacteria在水华的起始阶段相对丰度较高,而CF cluster在水华的消亡阶段表现出较高的相对丰度。
(2)海水表层浮游异养及群落内部不同类群的生长率表现出一定的差异性：在水华消亡阶段,除Gamma-Proteobacteria外,总浮游异养细菌及群落内主要类群均在水华的消亡阶段表现出较高的生长率。海水表层总浮游异养细菌群落及其各主要类群的摄食死亡率基本一致,较大值出现在水华的消亡阶段。
(3)相关性分析和回归分析结果表明,在水华期间,温度是调控该海域浮游异养细菌生长的关键因素。叶绿素a浓度和DOC浓度分别是控制浮游异养细菌生物量和细胞体积的关键因素。在水华过程中,环境中营养物质浓度的变化可能是控制浮游异养细菌群落结构的主要因素,而微型浮游动物的摄食则在水华的消亡时期调控着表层浮游异养细菌的群落结构。
Heterotrophic bacterioplankton plays an important role in marine biogeochemical cycles. Processes which mediate the transfer of carbon from dissolved pools to higher trophic levels are influenced by the growth and grazing losses of heterotrophic bacteria. The goals of this thesis are:to characterize seasonal and spatial variations in community structure of heterotrophic bacterioplankton in coastal (Logy Bay) and estuarine areas (Goose Bay and Lake Melville) of North Atlantic Ocean, including intensive sampling and experiments during the spring phytoplankton bloom in Logy Bay; to determine relationships among environmental factors, bacterial community structure, growth and grazing loss for bacterial population, as well as individual bacterial phylotypes; to provide insight into mechanism that lead to the observed variations at broad phylogenetic levels.
1.Seasonal changes in heterotrophic bacterial standing stock, community structure, rates of growth and grazing mortality for bacterioplankton, as well as individual phylogenetic groups within the community, are examined in the surface layer of Logy Bay. Factors controlling seasonal changes of these parameters are investigated using Pearson correlation and regression analysis.
(1) Heterotrophic bacterial abundance and biomass show high values in April and minimal in February, while cell volume exhibits maximum in May. Using fluorescence in situ hybridization (FISH), variations in composition of heterotrophic bacterial community were examined. Alpha-Proteobacteria and Cytophaga-Flavobacter cluster dominate the community overall, while Beta-Proteobacteria always comprise the lowest proportion of all detected phylogenetic groups. Relative abundances of all detected individual groups are low during winter (February and March), while higher values are detected during spring (April to June).
(2) Using FISH combined with dilution assay, seasonal changes in rates of growth and grazing mortality for bacterial community, as well as individual phylogenetic groups within the community, are determined. Growth rates for bacterial community as well as individual groups are high during summer (July and August), whereas low values are found during winter (February and March), with the exception of Alpha-Proteobacteria. Grazing mortality for bacterial community shows maximum during summer (July and August) and minimum during winter (March). However, no seasonal patterns of grazing mortality are observed among individual phylogenetic groups
(3) Results from Pearson correlation and regression analysis suggest that temperature is the main factor controlling rates of growth for bacteria, explained 40-60% variations in rates of growth for bacterial community as well as individual phylogenetic groups. Chlorophyll a concentration and dissolved organic carbon (DOC) are the controlling factors for bacterial abundance and cell volume, respectively. Correlations between rates of growth for bacterial community and those for individual groups implied bacterial phylogentic groups may share specific ecosystem function.
2.Spatial distributions in heterotrophic bacterial standing stock, community structure, rates of growth and grazing mortality for bacterioplankton, as well as individual phylogenetic groups within bacterial community, are examined in Goose Bay and Lake Melville. Factors related to spatial changes above are investigated using Pearson correlation and regression analysis.
(1) Bacterial abundance show distinct spatial distributions with minimal at low salinity waters and maximal at marine end of estuary. Vertical distribution of bacterial abundance show higher values at surface compared with those in the middle and at bottom. Cell volume and bacterial biomass show similar spatial patterns with each other, with no significant differences among stations. Cell volume increases with depth. Bacterial biomass show no significant differences in the middle and bottom layers at stations where salinity are low, while higher values are observed in the middle layer at stations in high salinity area.
(2) Analyses of heterotrophic bacterial community structure are performed at two selected sampling stations located at the low-(No. 111 station) and high-salinity (No.131 station) stations, respectively. Results indicate that relative abundance of Alpha-Proteobacteria is high at 131 station, whereas Beta-Proteobacteria is the dominant group at 111 station. Relative abundance of Alpha-, Beta-and Gamma-Proteobacteria uniformly distributed throughout the water column.
(3) Rates of growth and grazing mortality are determined using FISH combined with dilution assays at No. 111 station and No.131 station. No significant differences are observed between rates of growth for bacterial community at No. 111 station and No.131 station. Alpha-Proteobacteria growth rates are higher at No.131 station than those of other individual groups, while growth rates for Beta-Proteobacteria, as well as Gamma-Proteobacteria, are high at low-salinity station, No.111 station. No significant differences are found between grazing mortality for bacterial community at No.111 station and No. 131 station.
(4) Results from correlation and regression analysis indicate that dissolved organic carbon (DOC) and dissolved organic nitrogen (DON) are controlling factors for heterotrophic bacterial abundance and biomass. Depth and salinity are primary factors regulating cell volume. Since rates of bacterial growth indicates uptake of DOM, the co-occurrence of high growth rates and high relative abundance of individual phylotypes implies the possible regulations of nutrients uptake on bacterial community structure in this area.
3.Temporal and spatial variations in heterotrophic bacterial community structure are studied during spring phytoplankton bloom in Logy Bay to investigate possible mechanisms that lead to the observed variations in bacterial community composition during the bloom.
(1) Abundance and biomass of heterotrophic bacterioplankton show high values during the developmental stage of the spring bloom, while the maximal of cell volume is found during the post-bloom stage. Alpha-Proteobacteria reach its maximal during the beginning of spring bloom, while Beta-, Gamma-Proteobacteria and CF cluster reach maximum during the post-bloom stage.
(2) Using FISH combined with dilution assays, research indicate that significant differences in patterns of growth rates among bacterial community and detected phylogenetic groups, while patterns of grazing mortality are similar to each other during the study.
(3) Results from correlation and regression analysis indicate that temperature is the controlling factor for the growth of heterotrophic bacteria during the spring bloom, while chlorophyll a and DOC concentration are responsible for the changes of bacterial biomass and cell volume, respectively. Nutrient supplies might be the main factor driving variations in bacterial community structure during the spring bloom, while predation might be a key mechanism for regulation of bacterial community composition during the senescent stage of the bloom.
引文
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