Spatial heterogeneity of cyanobacterial communities and genetic variation of Microcystis populations within large, shallow eutrophic lakes (Lake Taihu and Lake Chaohu, China)

Cyanobacteria, specifically Microcystis, usually form massive blooms in eutrophic freshwater lakes. Cyanobacterial samples were collected from eight sites of both Lake Taihu and Lake Chaohu in late summer to determine the diversity and distribution pattern of cyanobacteria and Microcystis in large, shallow, entropic lakes with significant spatial heterogeneity and long-term Microcystis bloom. Molecular methods based on denaturing gradient gel electrophoresis and clone library analysis were used. A similar heterogeneous distribution pattern of cyanobacteria in both lakes was observed. Most parts of these two lakes with high trophic level were dominated by Microcystis. However, in the regions with low trophic levels as well as low concentrations of chlorophyll a, Synechococcus occupied a considerable percentage. Different morphospecies and genotypes dominated the bloom-forming Microcystis populations in these two lakes. Microcystis viridis and Microcystis novacekii were dominant in Lake Chaohu, whereas Microcystis flos-aquae was dominant in Lake Taihu. Only 2 of the13 Microcystis operational taxonomic units were shared between these two lakes. Analysis of molecular variance based on 16S to 23S internal transcribed spacer sequences indicated the significant genetic differentiation of Microcystis between these two lakes (F_st = 0.19, p < 0.001). However, only 19.46% of the genetic variability was explained by the population variation between lakes, whereas most (80.54%) of the genetic variability occurred within the lakes. Phylogenetic analysis revealed no phylogeographic structure of Microcystis population in these two lakes, as illustrated by their cosmopolitan nature. Our results revealed that spatial heterogeneity within lakes has more impact on the cyanobacterial diversity than geographical isolation in a local scale.     

Effects of elevated CO2 on dynamics of microcystin-producing and non-microcystin-producing strains during Microcystis blooms

In an attempt to elucidate the effects of different CO₂ concentrations (270, 380, and 750 μL/L) on the competition of microcystin-producing (MC-producing) and non-MC-producing Microcystis strains during dense cyanobacteria blooms, an in situ simulation experimentwas conducted in the Meiliang Bay of Lake Taihu in the summer of 2012. The abundance of total Microcystis and MC-producing Microcystis genotypes was quantified based on the 16S rDNA and mcyD gene using real-time PCR. The results showed that atmospheric CO₂ elevation would significantly decrease the pH value and increase the dissolved inorganic carbon (DIC) concentration. Changes in CO₂ concentration did not show significant influence on the abundance of total Microcystis population. However, CO₂ concentrations may be an important factor in determining the subpopulation structure of Microcystis. The enhancement of CO₂ concentrations could largely increase the competitive ability of non-MC-producing over MC-producing Microcystis, resulting in a higher proportion of non-MC-producing subpopulation in treatments using high CO₂ concentrations. Concurrently, MC concentration in water declined when CO₂ concentrations were elevated. Therefore, we concluded that the increase of CO₂ concentrations might decrease potential health risks of MC for human and animals in the future.     

A flow cytometer based protocol for quantitative analysis of bloom-forming cyanobacteria (Microcystis) in lake sediments

A quantitative protocol for the rapid analysis of Microcystis cells and colonies in lake sediment was developed using a modified flow cytometer, the CytoSense. For cell enumeration, diluted sediment samples containing Microcystis were processed with sonication to disintegrate colonies into single cells. An optimized procedure suggested that 5 mg dw (dry weight)/mL dilution combined with 200 W × 2 min sonication yielded the highest counting efficiency. Under the optimized determination conditions, the quantification limit of this protocol was 3.3×10⁴ cells/g dw. For colony analysis, Microcystis were isolated from the sediment by filtration. Colony lengths measured by flow cytometry were similar to those measured by microscopy for the size range of one single cell to almost 400 μupm in length. Moreover, the relationship between colony size and cell number was determined for three Microcystis species, including Microcystis flos-aquae, M. aeruginosa and M. wessenbergii. Regression formulas were used to calculate the cell numbers in different-sized colonies. The developed protocol was applied to field sediment samples from Lake Taihu. The results indicated the potential and applicability of flow cytometry as a tool for the rapid analysis of benthic Microcystis. This study provided a new capability for the high frequency monitoring of benthic overwintering and population dynamics of this bloom-forming cyanobacterium.     

Quantification of microcystin-producing and non-microcystin producing Microcystis populations during the 2009 and 2010 blooms in Lake Taihu using quantitative real-time PCR

Lake Taihu, a large, shallow hypertrophic freshwater lake in eastern China, has experienced lake-wide toxic cyanobacterial blooms annually during summer season in the past decades. Spatial changes in the abundance of hepatotoxin microcystin-producing and non-microcystin producing Microcystis populations were investigated in the lake in August of 2009 and 2010. To monitor the densities of the total Microcystis population and the potential microcystin-producing subpopulation, we used a quantitative real-time PCR assay targeting the phycocyanin intergenic spacer (PC-IGS) and the microcystin synthetase gene (mcyD), respectively. On the basis of quantification by real-time PCR analysis, the abundance of potential toxic Microcystis genotypes and the ratio of the mcyD subpopulation to the total Microcystis varied significantly, from 4.08× 10⁴ to 5.22× 10⁷ copies/mL, from 5.7% to 65.8%, respectively. Correlation analysis showed a strong positive relationship between chlorophyll-a, toxic Microcystis and total Microcystis; the abundance of toxic Microcystis correlated positively with total phosphorus and ortho-phosphate concentrations, but negatively with TN:TP ratio and nitrate concentrations. Meanwhile the proportion of potential toxic genotypes within Microcystis population showed positive correlation with total phosphorus and ortho-phosphate concentrations. Our data suggest that increased phosphorus loading may be a significant factor promoting the occurrence of toxic Microcystis bloom in Lake Taihu.     

洪泽湖有毒和无毒微囊藻丰度及其与环境因子之间的相关分析

为了解洪泽湖有毒和无毒微囊藻丰度及其空间分布,于2014年8月对洪泽湖30个采样点水体的营养盐浓度和Calson富营养化指数(trophic state index,TSI)进行研究,同时采用实时荧光定量PCR技术测定了有毒和无毒微囊藻丰度.结果表明,洪泽湖水体的总氮和总磷浓度平均值分别为1.63 mg·L-1和0.11 mg·L-1,富营养化指数在58.1~73.6之间,洪泽湖水质呈富营养化状态;有毒微囊藻在洪泽湖广泛分布,其丰度在1.13×104~3.51×106copies·m L-1之间,总微囊藻丰度在1.06×105~1.10×107copies·m L-1之间,有毒微囊藻占总微囊藻的比例在8.5%~38.5%之间,平均值为23.6%,有毒微囊藻丰度及其比例均呈现出明显的空间差异性;相关分析结果显示,总微囊藻、有毒微囊藻和有毒微囊藻所占比例三者之间呈极显著正相关性(P0.01),总微囊藻和有毒微囊藻丰度与叶绿素a浓度和TSI有极显著正相关性(P0.01),与透明度有极显著负相关性(P0.01),有毒微囊藻所占比例与叶绿素a、总氮、总磷和TSI有极显著正相关性(P0.01),与TN/TP和透明度有极显著负相关(P0.01).因此,削减洪泽湖总氮、总磷浓度一方面可以降低水体富营养化水平,另一方面有利于抑制有毒微囊藻对无毒微囊藻的竞争优势.     

Spatial and temporal variations of two cyanobacteria in the mesotrophic Miyun reservoir, China

Spatial variations in phytoplankton community within a large mesotrophic reservoir （Miyun reservoir, North China） were investigated in relation to variations in physico-chemical properties, nutrient concentrations, temperature and light conditions over a 5 month period in 2009. The dynamics of phytoplankton community was represented by the dominance of cyanobacteria through summer and fall, following with a short term dominance of chlorophyta in late fall, and a relatively high abundance of diatom in October; on the other hand, maximum phytoplankton biomass was recorded in the north shallow region of Miyun reservoir with a higher nutrients level. Particular attention was paid to the impacts of environmental conditions on the growth of two cyanobacteria genera, the toxin-producing Microcystis and the taste ＆ odor-producing Oscillatoria. Microcystis biomass was in general greatly affected by water temperature and mixing depth/local water depth ratio in this reservoir, while the Oscillatoria biomass in the surface and middle layers was greatly affected by total dissolved phosphorus, and that in the bottom layer was related with the Secchi depth/local water depth ratio. Abundant Oscillatoria biomass was observed only in late September when Microcystis biomass decreased and allowed sufficient light go through.     

6株蓝藻伪空胞的临界破裂压力研究

通过改进的压力毛细管法,研究了6株蓝藻伪空胞的临界破裂压力,并研究了过滤（真空度为0.02 MPa）和离心（离心力低于500 r/min）2种细胞浓缩方法对细胞内伪空胞含量测定的影响.结果表明,对于分散的单细胞的微囊藻,无论是采用过滤还是离心的方法,都难以达到理想的浓缩效果;对于群体形态的微囊藻,可以采用离心的方法进行浓缩,对于丝状的浮游蓝丝藻,宜采用过滤的方法进行浓缩. 此2种浓缩方法对细胞伪空胞含量的测定影响很小,伪空胞破裂率<7％. 6株蓝藻均为浅水湖泊藻类,由于长期自然选择的结果,6株蓝藻伪空胞的临界破裂压力比较接近. 5株微囊藻伪空胞的临界破裂压力为0.64～0.67 MPa之间;孟氏浮游蓝丝藻伪空胞的临界破裂压力为0.715 MPa,与深水湖泊或水库的蓝藻相比,6株蓝藻的伪空胞平均临界破裂压力均较小;在相同光照和温度条件下,单细胞微囊藻的膨压比群体微囊藻的膨压略大.     

鄱阳湖微囊藻毒素时空分布格局及其与理化和生物因子的关系

本文分别于2012年4月(春季)、8月(夏季)、10月(秋季)和2013年1月(冬季)在鄱阳湖进行了4次野外调查和样品采集,采用超高效液相色谱-串联质谱仪(UPLC-MS/MS)对鄱阳湖水柱中常见的3种微囊藻毒素(MC-RR、MC-YR和MCLR)的质量浓度进行定量分析,探寻了微囊藻毒素(microcystins,MCs)质量浓度的时空变化规律及其与湖泊环境理化因子和生物因子之间的关系.结果表明,MC-RR是鄱阳湖MCs的主要毒素种类.胞内微囊藻毒素(intracellular MCs,IMCs)与微囊藻生物量(r=0.47,P0.01)、颤藻生物量(r=0.68,P0.01)、蓝藻生物量(r=0.56,P0.01)、Chl-a(r=0.28,P0.01)之间均具有显著的正相关关系,但与鱼腥藻生物量之间没有明显的相关关系(P0.05),这说明微囊藻和颤藻是鄱阳湖微囊藻毒素的主要产毒藻类.IMCs与水温(r=0.51,P0.01)、透明度(r=0.69,P0.01)、Fe(r=0.43,P0.01)和Zn(r=0.43,P0.01)之间均呈现显著的正相关关系,与TN(r=-0.44,P0.01)、TP(r=-0.29,P0.01)、NO-2-N(r=-0.28,P0.05)、NH+4-N(r=-0.33,P0.05)、Ca(r=-0.34,P0.01)和Mg(r=-0.35,P0.05)均呈现显著的负相关关系,但与pH、PO3-4-P、NO-3-N、电导率、高锰酸盐指数、Cu离子之间的相关关系不显著(P0.05),这说明光照强度(透明度反映)、氮、磷、水温是控制鄱阳湖微囊藻毒素产生的重要因子,金属离子Ca、Mg、Fe、Zn在一定程度上能影响微囊藻毒素的产生.鄱阳湖IMCs和胞外微囊藻毒素(extracellular MCs,EMCs)呈现相似的季节变化趋势,即夏季最高,其次为秋季,冬季和春季较低.鄱阳湖IMCs的空间分布整体上呈现东部湖汊群、松门山周围以及蚌湖和蚌湖进入鄱阳湖的入湖口较高,其它区域较低的趋势,而EMCs的高值区位于松门山周围和蚌湖及其入湖口.     

淀山湖水华高发期浮游植物群落变化特征研究

根据2009年5～10月淀山湖浮游植物数量和种类的观测结果,对水华高发季节浮游植物群落组成特点及优势种交替过程进行分析,初步探讨这一时期浮游植物群落特征与环境因子的关系.结果表明,这一时期浮游植物主要由蓝藻和绿藻组成,蓝藻在密度上、绿藻在种类上占优势.水平分布上,西部和西南部的蓝藻密度值和种类数较高.总密度最高峰出现在9月,其值为23.40×107cells.L-1,其中蓝藻数量占90.3%,各样点间总密度差异显著(ANOVA,P<0.05).优势种在调查期间存在明显的交替现象,蓝藻门微囊藻属(Microcystis)藻类逐步成为优势种并形成水华.水温与pH是影响水华发生的重要环境因子,微囊藻受风向影响容易向下风向水域聚集形成水华.淀山湖中部和西部的浮游植物多样性较差,水华暴发时,全湖多样性指数降低,群落结构趋于简单.     

营养盐恢复对氮磷饥饿铜绿微囊藻生长的影响

在自然水体中微囊藻经常经历营养盐的限制和再补充,营养盐浓度的波动影响着微囊藻的生长和暴发.本文利用14C标记和荧光定量PCR研究了营养盐恢复对氮磷饥饿的铜绿微囊藻细胞生理和光合作用相关基因表达的影响.结果表明经过氮磷饥饿的铜绿微囊藻在补充营养盐后,都能快速生长,氮饥饿的藻细胞生长更快,但其经过快速增长期后很快开始衰亡.补充营养盐后,经过磷饥饿的藻细胞固碳能力和光合作用相关基因表达很快恢复至未经饥饿藻细胞的水平,而经过氮饥饿的藻细胞其固碳能力和相关基因虽然有很大提升,却难以恢复至未经饥饿的藻细胞的水平.这可能也是氮饥饿的藻细胞在饥饿解除后虽然能暂时地快速生长,但很快又进入衰亡的原因之一.     

湖北枝江市石子岭水库蓝藻水华研究

2009年8月21日对枝江市作为饮用水源的石子岭水库浮游植物和水体营养盐状况进行了调查.结果表明,该水库已经发生了微囊藻水华;浮游植物群落结构定量分析和多样性指数计算表明石子岭水库明显富营养化,惠氏微囊藻是是群落中绝对优势种.同时浮游植物生态位测度值表明微囊藻等蓝藻在时下的生态因子作用下将呈现逐渐衰退,而绿藻种群扩张,水库中浮游植物明显呈现由蓝藻优势向绿藻优势群落演替趋势.测定叶绿素荧光参数,以了解微囊藻水华的生长状态,比较研究中午强光下和经过24 h暗恢复及与室内培养的惠氏微囊藻三者的差异,表明夏末秋初阶段水库中微囊藻受到了一定的光照胁迫.水库中营养盐分析表明过高的总氮、总磷浓度和适宜的N/P比可能是水华暴发的主要原因,同时也是导致石子岭水库浮游植物呈现特定的群落结构特征的关键因素.     

电子束辐射对铜绿微囊藻藻毒素产生和释放的抑制作用研究

铜绿微囊藻及其所产生的藻毒素严重威胁人类健康和生命安全,给水处理带来很大困难.利用直线加速器产生的电子束对铜绿微囊藻分别进行1、2、3、4和5 kGy的辐射,通过胞内毒素和胞外毒素的变化,分析了电子束辐射对藻细胞毒素产生和释放过程的影响.结果表明,2～5 kGy的辐射使藻细胞叶绿素a含量迅速大幅下降,对胞外毒素的去除率分别为76.9%、76.2%、76.2%和97.7%,表明电子束辐射对微囊藻毒素的降解作用.胞内毒素、胞外毒素与叶绿素a浓度之间的相关性拟合进一步证实电子束辐射抑制了胞内毒素的产生.     

Nutrients regeneration pathway, release potential, transformation pattern and algal utilization strategies jointly drove cyanobacterial growth and their succession

In order to better understand the contribution of nutrients regeneration pathway, release potential and transformation pattern to cyanobacterial growth and succession, 7 sampling sites in Lake Chaohu with different bloom degree were studied every two months from February to November 2018. The carbon, nitrogen (N) and phosphorus (P) forms or fractions in surface, interstitial water and sediments as well as extracellular enzymatic activities, P sorption, specific microbial abundance and community composition in sediments were analyzed. P regeneration pathway was dominated by iron-bound P desorption and phosphorus-solubilizing bacteria solubilization in severe-bloom and slight-bloom area respectively, which both resulted in high soluble reactive phosphorus (SRP) accumulation in interstitial water. However, in severe-bloom area, higher P release potential caused the strong P release and algal growth, compared to slight-bloom area. In spring, P limitation and N selective assimilation of Dolichospermum facilitated nitrate accumulation in surface water, which provided enough N source for the initiation of Microcystis bloom. In summer, the accumulated organic N in Dolichospermum cells during its bloom was re-mineralized as ammonium to replenish N source for the sustainable development of Microcystis bloom. Furthermore, SRP continuous release led to the replacement of Dolichospermum by Microcystis with the advantage of P quick utilization, transport and storage. Taken together, the succession from Dolichospermum to Microcystis was due to both the different forms of N and P in water column mediated by different regeneration and transformation pathways as well as release potential, and algal N and P utilization strategies.     

野外模拟扰动方式对太湖浮游植物群落结构的影响

浮游植物是水生生态系统的重要生产者,扰动是水体的固有特征.为探究扰动方式对浮游植物的影响,通过野外模拟试验探讨不同扰动方式（持续扰动和间歇扰动）对太湖浮游植物的影响,并在试验期间测定各扰动条件下浮游植物生长和群落结构变化情况.结果表明:①对照组、间歇扰动组和持续扰动组中浮游植物平均生物量分别为5.92、19.42、58.27 mg/L,平均细胞密度为1.06×108、1.23×108、3.48×108 L-1,持续扰动组浮游植物生物量和细胞密度显著高于对照组和间歇扰动组.②在间歇扰动组中,浮游植物优势种始终是微囊藻,其占比平均值为84.61%;对照组和持续扰动组中,浮游植物主要优势种由微囊藻逐渐转变为湖生伪鱼腥藻,其中微囊藻占比从试验开始时的77.84%到试验结束时分别降至4.72%和5.62%,而湖生伪鱼腥藻从试验开始时的0.55%到试验结束时分别升至94.40%和90.52%.③对照组、间歇扰动组和持续扰动组中Shannon-Wiener指数分别为1.02、1.03和1.63,持续扰动组显著高于对照组和间歇扰动组.研究显示,在该试验条件下,间歇扰动有利于维持微囊藻在太湖浮游植物中的优势地位,同时表明扰动方式显著影响太湖浮游植物生长和群落结构.      

Physiological cellular responses and adaptations of Rhodococcus erythropolis IBBPo1 to toxic organic solvents

A new Gram-positive bacterium, Rhodococcus erythropolis IBBPo1（KF059972.1） was isolated from a crude oil-contaminated soil sample by enrichment culture method. R. erythropolis IBBPo1 was able to tolerate a wide range of toxic compounds, such as antibiotics（800–1000 μg/mL）,synthetic surfactants（50–200 μg/mL）, and organic solvents（40%–100%）. R. erythropolis IBBPo1 showed good tolerance to both alkanes（cyclohexane, n-hexane, n-decane） and aromatics（toluene, styrene, ethylbenzene） with logPOW（logarithm of the partition coefficient of the solvent in octanol–water mixture） values between 2.64 and 5.98. However, alkanes were less toxic for R. erythropolis IBBPo1 cells, compared with aromatics. The high organic solvent tolerance of R. erythropolis IBBPo1 could be due to the presence in their large genome of some catabolic（alkB, alkB1, todC1, todM, xylM）, transporter（HAE1） and trehalose-6-phosphate synthase（otsA1, KF059973.1） genes. Numerous and complex physiological cellular responses and adaptations involved in organic solvent tolerance were revealed in R. erythropolis IBBPo1 cells exposed 1 and 24 hr to 1% organic solvents. R. erythropolis IBBPo1 cells adapt to 1% organic solvents by changing surface hydrophobicity, morphology and their metabolic fingerprinting.Considerable modifications in otsA1 gene sequence were also observed in cells exposed to organic solvents（except ethylbenzene）.     

Characterization of typical taste and odor compounds formed by Microcystis aeruginosa

Production and characteristics of typical taste and odor (T&O) compounds by Microcystis aeruginosa were investigated. A few terpenoid chemicals, including 2-MIB, β-cyclocitral, and β-ionone, and a few sulfur compounds, such as dimethyl sulfide and dimethyl trisulfide, were detected. β-Cyclocitral and β-carotene concentrations were observed to be relevant to the growth phases of Microcystis. During the stable growth phase, 41-865 fg/cell of β-cyclocitral were found in the laboratory culture. β-Cyclocitral concentrations correlated closely with β-carotene concentrations, with the correlation coefficient R2 = 0.96, as it is formed from the cleavage reaction of β-carotene. For dead cell cases, a high concentration of dimethyl trisulfide was detected at 3.48-6.37 fg/cell. Four T&O compounds, including β-cyclocitral, β-ionone, heptanal and dimethyl trisulfide, were tested and found to be able to inhibit and damage Microcystis cells to varying degrees. Among these chemicals, β-cyclocitral has the strongest ability to quickly rupture cells.     

新安江水库河口区水质及藻类群落结构高频变化

水库库尾区的水环境多变,是水库生态系统突变的重要策源地.为探究大型水库水源地水环境演变特征及其突变的促发机制,以新安江水库为例,通过库尾河口断面18个月水质浮标的高频记录及3 d一次的藻类群落结构人工鉴定数据等,分析了气象水文过程影响下的水库库尾区的水温、溶解氧、浊度及营养盐等环境指标及藻类群落结构的高频变化特征,揭示了降雨、入流及季节温度变化等关键气象水文过程对水库水质及藻类群落结构的影响机制.结果表明:①在27 m深的河流入库区的水体温度和溶解氧存在明显的季节分层,相应水体藻类叶绿素a和营养盐等指标也同步发生分层,水温分层从气温达到14℃以上的3月中旬开始,至气温降至24℃后的10月中旬结束,期间较大降雨和入流多次破坏水温分层;②河道入库区水体氮、磷等营养盐变幅大,总磷浓度变幅为0. 011～0. 188 mg·L^-1之间,总氮浓度变幅为0. 75～2. 76 mg·L^-1之间,总磷和总氮中的溶解态占比分别为56%及88%,降雨入流对水体营养盐浓度影响巨大,3 d的累积降雨与水体氮、磷浓度显著正相关,3～6月(雨季)的营养盐含量明显高于其...     

水质改善的浮游植物群落演替机制模拟研究

模拟了广州市某河流的不同水质改善阶段,并研究其对铜绿微囊藻（Microcystis aeruginosa）、斜生栅藻（Scenedesmus obliquus）、短线脆杆藻（Fragilaria brevistriata）和易变裸藻（Euglena mutabilis）生长及超氧化物歧化酶（SOD）活性的影响.同时,考察了由4种藻类构成的人工群落的生长特征和群落演替率,以探讨河流水质改善过程中浮游植物群落演替机制.结果表明：4种藻类对氮、磷营养盐和COD_Cr等胁迫的响应程度差异及种间相互竞争作用等是水质改善过程中导致浮游植物群落演替的影响因素;水质改善初期（1~3 d）,群落演替率与改善程度呈显著的正相关关系（p<0.05）;但较长时间（约13 d）处于相对稳定的环境条件下时,群落演替速率将趋于零,从而达到一个新的稳定状态.水质的不断改善将导致浮游植物的种类和数量呈现出蓝藻和裸藻逐渐减少,而绿藻和硅藻逐渐增多的变化趋势.     

Submerged vegetation removal promotes shift of dominant phytoplankton functional groups in a eutrophic lake

Historical data indicate that the dominance of submerged plants in Dianchi Lake in the 1960 s was characterized by low algal density with dominance of non-toxic group J（Scenedesmus,Pediastrum,etc.）. The removal of submerged plants,which began in the 1970 s,resulted in the expansion of bloom-forming Microcystis（group M）. Laboratory experiments suggested that Microcystis aeruginosa was inclined to grow and develop at elevated temperatures. The growth of Scenedesmus obliquus was slower than that of co-cultivated M. aeruginosa in the absence of Ceratophyllum demersum,especially at higher temperatures. The existence of submerged plant C. demersum could inhibit the growth of the harmful algae M. aeruginosa and this inhibitory effect by C. demersum was enhanced with an increase in temperature. Instead,with C. demersum,the growth of S. obliquus was not inhibited,but the co-cultivated M. aeruginosa was eliminated in a short time. Combined with the historical data and laboratory experiments,it was indicated that the submerged plants might play important roles in the dominance of the non-toxic group J in the historical succession. Consequently,the introduction of the submerged plant such as C. demersum might alter the dominant phytoplankton functional groups from M to J and benefit the restoration of the eutrophic lake.