富营养化水体微囊藻分子生态研究
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摘要
湖泊富营养化和蓝藻水华暴发已经成为世界性的环境难题,蓝藻水华的大规模持续暴发不仅严重破坏了水体水生生态系统的结构,影响和降低其生态功能,同时产生大量蓝藻毒素,给公共安全带来重大威胁。当前,我国湖泊的富营养化问题异常突出,三大淡水湖泊常年暴发以微囊藻属为主的蓝藻水华,弄清水华暴发过程,尤其是产毒微囊藻水华的暴发以及在水体中的动态演化规律,为富营养化水体微囊藻的控制和治理提供理论依据。
采用荧光定量PCR和微囊藻ITS基因克隆文库的构建等手段调查分析了太湖和秦淮河水系中微囊藻、产毒微囊藻的分子多样性,探讨了不同形态型和基因型微囊藻的动态演替过程,揭示了富营养化水体中微囊藻群落结构和数量的动态变化规律及其驱动因子,取得了以下主要成果：
(1)在太湖水体中微囊藻生物量的年变化幅度较大,从1.18×105到2.90×109cells/L,年均值北部湖区最高,南部湖区最低。产毒微囊藻的生物量季节性变化较强,产毒微囊藻占总微囊藻群落生物量的峰值一般出现在水华暴发的前期和水华发生的中后期,最高比例达92%,其他时期的产毒微囊藻含量比例相对较低。统计分析表明影响太湖微囊藻和产毒微囊藻丰度的主要环境因子是温度和正磷酸盐。
(2)太湖共存在8种不同形态型微囊藻,其中水华微囊藻、鱼害微囊藻、铜绿微囊藻和惠氏微囊藻是太湖的优势形态型微囊藻。这4种微囊藻的演替主要受温度的影响,在低水温的水华前期、初期和末期,微囊藻群落优势种为水华和鱼害微囊藻,随着水温的升高,水华中期以后铜绿微囊藻和惠氏微囊藻演替成为优势微囊藻形态型。同时发现,稳定的地理环境有助于微囊藻群落的演替,风浪较大的开阔水域则有利于维持水华微囊藻的优势地位。
(3)共获得太湖微囊藻ITS基因序列4600条,划分为1260种不同的基因型。在全太湖不同点位和水华发生的不同时期,均存在高比例的基因型演替,表明太湖微囊藻群落具有很高的分子多样性和丰富的基因型构成。竺山湾的微囊藻基因型在进化聚类上较好聚为一支,表明相对稳定和封闭的环境有利于微囊藻基因型地域性的形成。太湖的优势微囊藻基因型T1在全太湖周年存在,时间序列上,在水华前期和初期占有较高比例,空间上,在强水动力的开阔水域比例相对较高,有可能是太湖微囊藻水华越冬和复苏的主要种源。
(4)通过单藻株筛选比对,证实基因型T1的形态型属于水华微囊藻。通过对该藻株的研究发现,其最适生长温度为20℃以上,且在20℃的生长速率要高于太湖分离的其他株微囊藻。低温恢复实验表明,与其他微囊藻藻株相比,T1型藻株具有更快的生理恢复速率,可以从低温逆境中快速恢复生理活性,并在高的生长率下快速达到较高的生物量,从而为其在野外春季快速生长并形成优势奠定生理基础。
与单一的强扰动型太湖相比,包含湖泊和众多支流的秦淮河水系的系统取样研究结果表明：
(1)秦淮河的微囊藻群落在不同水环境下(不同支流)显示了较高的遗传多样性,24个主要基因型可以划分为9个不同的基因型组。
(2)低流速和低水力交换条件形成的稳定的地理环境差异可以影响和引导微囊藻群落的分子适应及进化选择,从而在分子进化关系上得到较好地理区分。
(3)pH是影响不同基因型的地理分布及分子适应差异的显著环境因子。
(4)云台山河、俞庄桥、七桥瓮和石臼湖4个样点可能是秦淮河水华大规模增殖和暴发的潜在种子库。
With the increasing development of economy and human activities, environmental problems caused by cyanobacterial blooms in various water bodies especially in freshwater lakes has become a concern all over the world. The characteristic of eutrophication in freshwater ecosystem is the annual cyanobacterial blooms during the high temperature season, which could not only break the structure and the balance of hydrophytic ecology in the water but also pose serious health threats due to the production of toxic compounds. Eutrophication has become a serious environmental problem in most lakes of China. As a prerequisite to the ecosystem management for the cyanobacterial bloom events, it is essential to determine the relationship between the biomass distribution of MC- and non MC-Microcystis and the related environmental factors during the development of the bloom in freshwater ecosystem. However, few systematic investigations in the community structure of Microcystis have been conducted on eutrophic lakes.
To better understand the basic characteristics of Microcystis blooms in Lake Taihu, samples were collected from December 2008 to October 2010 and analyzed morphologically via the distribution and dynamics of different Microcystis morphotypes and genetically via sequencing the clone library of the internal transcribed spacer of the rRNA operon (ITS). We also conducted quantitative PCR to assess total Microcystis abundance and the proportion of microcystin-producing subpopulation. Marked succession of the Microcystis populations in both morphotypes and genotypes occurred during the course of the Microcystis bloom. The T1 genotype, characterized by strains of Microcystis flos-aquae, was the most dominant genotype in Lake Taihu during winter and spring around the whole lake and likely acted as the main inoculum for forming blooms the following year. The 4600 ITS sequences were obtained and were revealed to have 1260 Microcystis genotypes, the highest genetic diversity of Microcystis reported in Lake Taihu. Water temperature temporally affected the succession of both Microcystis genotypes and morphotypes, whereas wind-driven disturbance and the micro-environment influenced spatial distribution of Microcystis genotypes and morphotypes. High ratios of mcyD containing Microcystis subpopulations were detected during the onset and later phase of blooms. Redundancy analysis (RDA) indicated that water temperature and PO4-P were major factors controlling total Microcystis abundance and proportions of microcystin-producing Microcystis population at hyper-eutrophic waters.
To determine the Microcystis population and its spatial distributions along the reaches in Qinhuai River, a convenient approach was applied genetically by sequencing the internal transcribed spacer of the rRNA operon (ITS). The TCS analysis showed that 9 groups were classified from the main 24 genotypes and each group was dominated by one highly represented root sequence. Marked changes in the composition and proportion of the Microcystis ITS genotype were detected from/along the upper to the lower reaches. The possible seed sources of the bloom were located in 4 different locations. Furthermore, it was found that pH was the primary factor affecting the spatial distribution of the main genotype group between samples.
采用荧光定量PCR和微囊藻ITS基因克隆文库的构建等手段调查分析了太湖和秦淮河水系中微囊藻、产毒微囊藻的分子多样性,探讨了不同形态型和基因型微囊藻的动态演替过程,揭示了富营养化水体中微囊藻群落结构和数量的动态变化规律及其驱动因子,取得了以下主要成果：
(1)在太湖水体中微囊藻生物量的年变化幅度较大,从1.18×105到2.90×109cells/L,年均值北部湖区最高,南部湖区最低。产毒微囊藻的生物量季节性变化较强,产毒微囊藻占总微囊藻群落生物量的峰值一般出现在水华暴发的前期和水华发生的中后期,最高比例达92%,其他时期的产毒微囊藻含量比例相对较低。统计分析表明影响太湖微囊藻和产毒微囊藻丰度的主要环境因子是温度和正磷酸盐。
(2)太湖共存在8种不同形态型微囊藻,其中水华微囊藻、鱼害微囊藻、铜绿微囊藻和惠氏微囊藻是太湖的优势形态型微囊藻。这4种微囊藻的演替主要受温度的影响,在低水温的水华前期、初期和末期,微囊藻群落优势种为水华和鱼害微囊藻,随着水温的升高,水华中期以后铜绿微囊藻和惠氏微囊藻演替成为优势微囊藻形态型。同时发现,稳定的地理环境有助于微囊藻群落的演替,风浪较大的开阔水域则有利于维持水华微囊藻的优势地位。
(3)共获得太湖微囊藻ITS基因序列4600条,划分为1260种不同的基因型。在全太湖不同点位和水华发生的不同时期,均存在高比例的基因型演替,表明太湖微囊藻群落具有很高的分子多样性和丰富的基因型构成。竺山湾的微囊藻基因型在进化聚类上较好聚为一支,表明相对稳定和封闭的环境有利于微囊藻基因型地域性的形成。太湖的优势微囊藻基因型T1在全太湖周年存在,时间序列上,在水华前期和初期占有较高比例,空间上,在强水动力的开阔水域比例相对较高,有可能是太湖微囊藻水华越冬和复苏的主要种源。
(4)通过单藻株筛选比对,证实基因型T1的形态型属于水华微囊藻。通过对该藻株的研究发现,其最适生长温度为20℃以上,且在20℃的生长速率要高于太湖分离的其他株微囊藻。低温恢复实验表明,与其他微囊藻藻株相比,T1型藻株具有更快的生理恢复速率,可以从低温逆境中快速恢复生理活性,并在高的生长率下快速达到较高的生物量,从而为其在野外春季快速生长并形成优势奠定生理基础。
与单一的强扰动型太湖相比,包含湖泊和众多支流的秦淮河水系的系统取样研究结果表明：
(1)秦淮河的微囊藻群落在不同水环境下(不同支流)显示了较高的遗传多样性,24个主要基因型可以划分为9个不同的基因型组。
(2)低流速和低水力交换条件形成的稳定的地理环境差异可以影响和引导微囊藻群落的分子适应及进化选择,从而在分子进化关系上得到较好地理区分。
(3)pH是影响不同基因型的地理分布及分子适应差异的显著环境因子。
(4)云台山河、俞庄桥、七桥瓮和石臼湖4个样点可能是秦淮河水华大规模增殖和暴发的潜在种子库。
With the increasing development of economy and human activities, environmental problems caused by cyanobacterial blooms in various water bodies especially in freshwater lakes has become a concern all over the world. The characteristic of eutrophication in freshwater ecosystem is the annual cyanobacterial blooms during the high temperature season, which could not only break the structure and the balance of hydrophytic ecology in the water but also pose serious health threats due to the production of toxic compounds. Eutrophication has become a serious environmental problem in most lakes of China. As a prerequisite to the ecosystem management for the cyanobacterial bloom events, it is essential to determine the relationship between the biomass distribution of MC- and non MC-Microcystis and the related environmental factors during the development of the bloom in freshwater ecosystem. However, few systematic investigations in the community structure of Microcystis have been conducted on eutrophic lakes.
To better understand the basic characteristics of Microcystis blooms in Lake Taihu, samples were collected from December 2008 to October 2010 and analyzed morphologically via the distribution and dynamics of different Microcystis morphotypes and genetically via sequencing the clone library of the internal transcribed spacer of the rRNA operon (ITS). We also conducted quantitative PCR to assess total Microcystis abundance and the proportion of microcystin-producing subpopulation. Marked succession of the Microcystis populations in both morphotypes and genotypes occurred during the course of the Microcystis bloom. The T1 genotype, characterized by strains of Microcystis flos-aquae, was the most dominant genotype in Lake Taihu during winter and spring around the whole lake and likely acted as the main inoculum for forming blooms the following year. The 4600 ITS sequences were obtained and were revealed to have 1260 Microcystis genotypes, the highest genetic diversity of Microcystis reported in Lake Taihu. Water temperature temporally affected the succession of both Microcystis genotypes and morphotypes, whereas wind-driven disturbance and the micro-environment influenced spatial distribution of Microcystis genotypes and morphotypes. High ratios of mcyD containing Microcystis subpopulations were detected during the onset and later phase of blooms. Redundancy analysis (RDA) indicated that water temperature and PO4-P were major factors controlling total Microcystis abundance and proportions of microcystin-producing Microcystis population at hyper-eutrophic waters.
To determine the Microcystis population and its spatial distributions along the reaches in Qinhuai River, a convenient approach was applied genetically by sequencing the internal transcribed spacer of the rRNA operon (ITS). The TCS analysis showed that 9 groups were classified from the main 24 genotypes and each group was dominated by one highly represented root sequence. Marked changes in the composition and proportion of the Microcystis ITS genotype were detected from/along the upper to the lower reaches. The possible seed sources of the bloom were located in 4 different locations. Furthermore, it was found that pH was the primary factor affecting the spatial distribution of the main genotype group between samples.
引文
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