不同海洋生境中超微型浮游生物的分布特点研究
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摘要
本论文研究了不同海洋生境（东海陆架、地中海马赛湾以及黄海胶州湾）中超微型浮游生物群落结构（包括聚球藻、原绿球藻、微微型和微型光合真核生物以及异养细菌）与丰度的时空变化，并分析研究了不同海区中影响超微型生物分布的可能因素。
2006年11月（秋季航次）和2007年2月（冬季航次）东海不同水团中微微型浮游生物群落结构及丰度分布的研究。调查区域主要受到三类水团的影响：沿岸水团，陆架混合水团和黑潮水团。流式细胞仪分析发现东海微微型浮游生物由四个类群组成，即聚球藻、原绿球藻、微微型光合真核生物以及异养细菌。聚球藻、微微型光合真核生物和异养细菌在秋季航次的平均丰度分别为(10.63±10.88)×10~3、(1.61±1.16)×10~3、(3.39±1.27)×10~5cells/mL，冬季航次的平均丰度分别为(6.45±8.60)×10~3、(3.23±2.63)×10~3、(3.76±1.37)×10~5cells/mL。微微型浮游生物群落结构受水团的影响明显，主要表现为原绿球藻很少在表层和沿岸水区域出现，主要存在于陆架混合水及黑潮水区域；秋季聚球藻在陆架混合区域发生明显的亚群分化。微微型浮游生物的丰度分布与水团也有明显的关系，不同类群受水团的影响不完全一致。聚球藻最高丰度值出现在黑潮水区域，最低丰度值出现在沿岸水区域，其在黑潮水区域的丰度明显高于沿岸水区域，这种现象可能是受温度影响的。与聚球藻相比，微微型光合真核生物受水团的影响相对较小，异养细菌受水团的影响最不明显。相关性分析也显示不同水团中微微型浮游生物与环境因子（温度、营养盐和叶绿素a）之间的相关性存在明显差异，可见不同水团对微微型浮游生物分布的影响是有显著差异的。
东海陆架区域远离近岸，受人类活动的影响较小，微微型浮游生物主要是受到水团，即自然环境因素的影响。为了更好的了解人类活动对海洋微生物的影响，我们调查研究了地中海西北部马赛湾自2010年5月至2012年4月超微型浮游生物的组成及季节变化。马赛湾属于开放性的海湾，受人类活动的影响较为明显。马赛湾超微型浮游生物由四个主要类群组成，包括聚球藻、微微型和微型光合真核生物以及异养细菌。已有马赛湾原绿球藻广泛存在的报道，本研究中很少发现原绿球藻。从丰度上看，马赛湾聚球藻是自养型超微型浮游生物的优势类群。聚球藻丰度呈现显著的季节性变化，其高丰度值出现在春季4、5月，高达90×10~3cells/mL，低丰度值出现在秋冬季节，小于15×10~3cells/mL。聚球藻的春季最高值（2010年5月，129.59×10~3cells/mL）和冬季最低值（2012年3月，2.29×10~3cells/mL）之间相差超过40倍。微微型光合真核生物的季节变化与聚球藻相似，高丰度值出现在冬季2、3月，丰度高达10.72×10~3cells/mL。细胞粒径大于2μm的微型光合真核生物丰度比聚球藻和微微型光合真核生物丰度要低的多，其季节变化趋势与聚球藻类似。异养细菌的季节变化波动性非常明显，最大值出现在2012年3月（9.74×10~5cells/mL）。此外，在调查过程中我们发现了一个非常特殊的现象：2010年7月12日的样品中，除了广泛存在的聚球藻、微微型和微型光合真核生物和异养细菌外，原绿球藻也大量存在，此外我们还观察到了一种与聚球藻类似的含有橙色荧光（藻红蛋白）和红色荧光（叶绿素a）、细胞粒径大于2μm的未知类群，该类群丰度高达7104events/mL。通过流式细胞仪分选后在荧光显微镜观察发现，与某些含异形胞的固氮蓝细菌藻丝体类似，它们似乎是一种呈链状、含异形胞的藻丝体，但异形胞并没有位于末端。此外，荧光显微镜照片显示该链状藻丝体是折叠状的。通常情况下，藻丝体都是与硅藻共生的，自由藻丝体很少报道。据我们所知，这种自由藻丝体的大量爆发现象从来没有被报道过。然而，由于样品量少（只有1.5mL）、分析时间晚于采样时间以及荧光显微镜照片分辨率较低等因素限制，我们难以进行物种鉴定。夏季高温引起的水体层化现象、硝酸盐的耗尽等可能与这一现象有关。马赛湾自由藻丝体短暂爆发现象的发生原因有待于进一步研究。马赛湾超微型浮游生物的监测持续进行中，有必要引进高频率原位流式细胞仪，从而避免错过类似的、短暂发生的特殊现象，能够进行进一步的物种鉴定。
与马赛湾开放性海湾不同，胶州湾属于半封闭的浅海海湾，十多条河流入湾，受人类活动的影响更为显著。我们通过2007年1月至2010年12月胶州湾月调查航次，研究了胶州湾微微型浮游生物长期的季节变化。调查航次显示，聚球藻、微微型光合真核生物以及异养细菌普遍存在于胶州湾，而原绿球藻至今尚未被观察到。在季节分布上，无论是在湾内、还是湾口和湾外，微微型浮游生物都呈现出明显的季节变化。聚球藻最高值发生在8月水温最高（>24°C）的时期，自12月到5月（水温<15°C），其丰度维持在低水平。微微型光合真核生物高丰度值出现在4-7月，除8月的其他时期，微微型光合真核生物丰度值都高于聚球藻。异养细菌每年会在8月（最高值）和4月（次高值）出现两个高丰度值，这两个高值恰恰与叶绿素高值出现的时间一致，由此可以推测二者之间存在明显关联。主成分分析结果显示胶州湾微微型浮游生物的季节变化主要受温度和营养盐（硝酸盐、亚硝酸盐、铵盐及硅酸盐等）影响。温度、营养盐与微微型浮游生物丰度之间的正相关关系，也表明了温度和营养盐对胶州湾微微型浮游生物丰度分布的重要性。
马赛湾与胶州湾的自然及人类环境既存在相似点，又有所不同。首先二者都是海湾系统，并受到人类活动的影响。马赛湾属于地中海的一部分，临近法国第二大城市马赛；而胶州湾入海口与黄海相连，受到青岛市人类活动的影响。不同点方面：马赛湾没有淡水河流的影响，胶州湾则受到十多条淡水河流的影响；此外，由于胶州湾夏季降雨量丰富，而马赛湾受到地中海气候的影响，夏季降雨量少，这些特征对于海洋超微型浮游生物的分布都会产生一定的影响。比较马赛湾和胶州湾超微型浮游生物的类群发现，聚球藻、微微型和微型光合真核生物是普遍存在的，而原绿球藻在两个海湾都很少发现。在季节分布上，马赛湾聚球藻和微型光合真核生物高丰度值出现在春季，低丰度值出现在秋季和冬季。微微型光合真核生物高丰度值出现在冬季，比聚球藻稍早；胶州湾聚球藻高丰度值出现在8月，其他月份丰度值较低，微微型和微型光合真核生物高丰度值出现在7月，比聚球藻早一个月。两个海湾超微型浮游生物细胞丰度与细胞荧光含量呈现相反的趋势，即细胞丰度高的季节，细胞较小，细胞荧光含量较低。主成分分析发现马赛湾超微型浮游生物丰度的季节变化主要受到营养盐的影响；胶州湾超微型浮游生物丰度的季节变化则是受到温度和营养盐的双重影响。
通过本论文的实验，我们研究了不同海洋生境中超微型浮游生物的群落结构和丰度变化，发现环境因子和人类活动会在不同程度上对超微型浮游生物产生影响，且不同生境中对海洋超微型浮游生物丰度的影响因子有明显的区别。
In this dissertation, temporal and spatial variations of ultraplankton (Synechococcus,Prochlorococcus, picoeukaryotes, nanoeukaryotes and heterotrophic bacteria) abundanceswere studied in different marine environments (in the East China Sea, Bay of Marseille inthe NW Mediterranean Sea and Jiaozhou Bay in the Yellow Sea) based on flow cytometryanalysis. The relationships between environmental conditions and ultraplankton abundancewere also studied in order to search for the possible controlling factors on ultraplanktondistribution in different marine environments.
Picoplankton distribution in different water masses of the East China Sea (ECS) wasinvestigated in November2006and February2007. The cruises tracked crossed three majorwater masses: Coastal Water Mass (CWM), Shelf Mixing Water Mass (MWM) andKuroshio Water Mass (KWM). Picoplankton composition was resolved in four main groupsby flow cytometry, namely Synechococcus, Prochlorococcus, picoeukaryotes, andheterotrophic bacteria. The average Synechococcus, picoeukaryotes and heterotrophicbacteria abundances were (10.63±10.88)×10~3,(1.61±1.16)×10~3,(3.39±1.27)×10~5cells/mLin autumn and (6.45±8.60)×10~3,(3.23±2.63)×10~3,(3.76±1.37)×10~5cells/mL in winter,respectively. Prochlorococcus was seldom observed in surface samples and not found inCWM, neither in autumn nor in winter. In contrast, Prochlorococcus was present in MWMand KWM with abundance at around10~3cells/mL in autumn and winter. Synechococcusdistribution showed marked variations with respect to water masses with the maximumabundance in KWM and minimum abundance in CWM. The depth-averaged integratedabundance of Synechococcus was about5-fold higher in KWM than in CWM, which mightbe mainly influenced by temperature. In autumn particularly, at several stations in MWM,Synechococcus was resolved as two subgroups. Picoeukaryotes were less affected bywater-mass differences than Synechococcus. Heterotrophic bacteria depth-averaged integrated abundance exhibited the smallest seasonal variations with respect to watermasses. Correlation analysis showed obvious differences between picoplankton abundancesand environmental parameters (temperature, nutrients and chlorophyll a) in three watermasses, indicated the three water masses are likely to have different influences onpicoplankton (particularly Synechococcus) distribution.
In the Bay of Marseille (NW Mediterranean), the composition of ultraplankton (<10μm) sampled in surface (1m depth) was investigated from May2010to April2012.Ultraplankton (<10μm) composition was resolved in four main groups by flow cytometry,namely Synechococcus, picoeukaryotes, nanoeukaryotes and heterotrophic bacteria.Prochlorococcus was seldom observed in our study, even though in the previous studies,Prochlorococcus has been widely reported. In the Bay of Marseille, Synechococcusdominated the ultraphytoplankton community in terms of cell abundance. Synechococcusshowed a clear seasonal pattern with higher abundance in spring (April and May,>90×10~3cells/mL) and lower abundance in autumn and winter (<15×10~3cells/mL). Synechococcusabundance varied more than40-fold between a spring maximum (May2010,129.59×10~3cells/mL) and winter minimum (March2012,2.29×10~3cells/mL) values. Picoeukaryoteshad a similar distribution pattern as Synechococcus and their high abundance occurred inwinter (January-March,>10.72×10~3cells/mL), slightly earlier than Synechococcus.Nanoeukaryotes exhibited much lower abundance than Synechocccus and picoeukaryotes.The distribution pattern of nanoeukaryotes was similar to that of Synechococcus. FromMay to September in2010, the data of heterotrophic bacteria abundance were missingbecause of technical reasons. Heterotrophic bacteria exhibited pronounced fluctuationsduring the study period and reached their maximum abundance (9.74×10~5cells/mL) inMarch2012. On12July2010, in addition to groups of Synechococcus, Prochlorococcus,picoeukaryotes, and nanoeukaryotes, an unidentified additional cluster was observed.These unknown particles were sorted out by flow cytometry and then observed byepifluorescence microscopy: they appeared to be chain-forming microorganisms, just liketrichomes of some diazotroph cyanobacteria with one heterocyst, but not unambiguouslydistal. The chains observed after cell sorting were not straight, but rather folded. The shortoccurrence of free trichomes in the Bay of Marseille with such a high abundance remains to be explained. High temperature in summer, induced stratification of the water column,and nitrate depletion, may be related to this event. High frequency survey ofultraphytoplankton assemblages at the single cell level appears necessary to observe moreefficiently such phenomenon and document more extensively its full dynamics.
The seasonal variability of picoplankton abundances in the semi-enclosed Jiaozhou Bay(Yellow Sea) were investigated monthly from January2007to December2010by flowcytometic analysis. Picoplankton community in the studied area was comprised ofSynechococcus, picoeukaryotes and heterotrophic bacteria. Prochlorococcus was notdetected at any station during the whole year. A pronounced seasonality was observed inpicoplankton community structure and abundances inside the bay, in the Bay mouth andoutside the bay. Synechococcus exhibited a clear seasonal pattern, with maximumabundance occurring in August when temperature was highest and minimum abundance inwinter and spring. The maximum abundance of picoeukaryotes occurred slightly earlier(April-July) than Synechococcus. Picoeukaryotes abundance outnumbered Synechococcusin most of the year except in August. Heterotrophic bacteria exhibited two peaks ofabundances in August (major peak) and April (minor peak), which were quite similar asChlorophyll a distribution. Principal Component Analysis (PCA) suggested that seasonalvariations of picoplankton were mainly driven by temperature and nutrient concentrations(NO3-, NO2-, NH4-and SiO32-). The significantly positive correlations betweentemperature/nutrients and picoplankton abundances also implied the importance oftemperature and nutrients in picoplankton distribution in Jiaozhou Bay.
The ultraphytoplankton composition, abundance and fluorescence in the Bay ofMarseille and Jiaozhou were compared in2011and2012. In the Bay of Marseille andJiaozhou, Synechococcus, picoeukaryotes and nanoeukaryotes were observed in all thesamples. Prochlorococcus was seldom observed in the Bay of Marseille and neverobserved in the Jiaozhou Bay. Besides, in the Bay of Marseille, a possible bloom of freetrichomes was observed in a short period during summer2010. While in the Jiaozhou Bay,an unknown and abundant cluster was obsvered in all the samples. With flow cytometryand scanning electron microscopy, it is difficult to identify this species. In the next step,molecular technique should be introduced for further identification. In the seasonal scale, ultraphytoplankton in the Bay of Marseille and Jiaohzou exhibited quite differentdistribution patterns. In the Bay of Marseille, Synechococcus and nanoeukaryotes presentedthe maximum abundance in spring and minimum abundances in autumn and winter.Picoeukaryotes showed the maximum abundance in winter, slightly earlier thanSynechococcus and nanoeukaryotes. While in the Jiaozhou Bay, Synechococcus abundancewas highest in August (T>24°C) and lowest in winter and spring (T<15°C). Picoeukaryotesand nanoeukaryotes exhibited highest abundances in July, one month earlier thanSynechococcus, and lower abundances in winter and spring, similar as Synechococcus. Itseemed in the oligotrophic Bay of Marseille, nutrient supply is more important forultraphytoplantkon distribution, while in the Jiaozhou Bay, temperature is the keycontrolling factor for ultraphytoplantkon distribution. In both bays, cell abundanceexhibited a clear opposite trend compared to cellular fluorescence.
2006年11月（秋季航次）和2007年2月（冬季航次）东海不同水团中微微型浮游生物群落结构及丰度分布的研究。调查区域主要受到三类水团的影响：沿岸水团，陆架混合水团和黑潮水团。流式细胞仪分析发现东海微微型浮游生物由四个类群组成，即聚球藻、原绿球藻、微微型光合真核生物以及异养细菌。聚球藻、微微型光合真核生物和异养细菌在秋季航次的平均丰度分别为(10.63±10.88)×10~3、(1.61±1.16)×10~3、(3.39±1.27)×10~5cells/mL，冬季航次的平均丰度分别为(6.45±8.60)×10~3、(3.23±2.63)×10~3、(3.76±1.37)×10~5cells/mL。微微型浮游生物群落结构受水团的影响明显，主要表现为原绿球藻很少在表层和沿岸水区域出现，主要存在于陆架混合水及黑潮水区域；秋季聚球藻在陆架混合区域发生明显的亚群分化。微微型浮游生物的丰度分布与水团也有明显的关系，不同类群受水团的影响不完全一致。聚球藻最高丰度值出现在黑潮水区域，最低丰度值出现在沿岸水区域，其在黑潮水区域的丰度明显高于沿岸水区域，这种现象可能是受温度影响的。与聚球藻相比，微微型光合真核生物受水团的影响相对较小，异养细菌受水团的影响最不明显。相关性分析也显示不同水团中微微型浮游生物与环境因子（温度、营养盐和叶绿素a）之间的相关性存在明显差异，可见不同水团对微微型浮游生物分布的影响是有显著差异的。
东海陆架区域远离近岸，受人类活动的影响较小，微微型浮游生物主要是受到水团，即自然环境因素的影响。为了更好的了解人类活动对海洋微生物的影响，我们调查研究了地中海西北部马赛湾自2010年5月至2012年4月超微型浮游生物的组成及季节变化。马赛湾属于开放性的海湾，受人类活动的影响较为明显。马赛湾超微型浮游生物由四个主要类群组成，包括聚球藻、微微型和微型光合真核生物以及异养细菌。已有马赛湾原绿球藻广泛存在的报道，本研究中很少发现原绿球藻。从丰度上看，马赛湾聚球藻是自养型超微型浮游生物的优势类群。聚球藻丰度呈现显著的季节性变化，其高丰度值出现在春季4、5月，高达90×10~3cells/mL，低丰度值出现在秋冬季节，小于15×10~3cells/mL。聚球藻的春季最高值（2010年5月，129.59×10~3cells/mL）和冬季最低值（2012年3月，2.29×10~3cells/mL）之间相差超过40倍。微微型光合真核生物的季节变化与聚球藻相似，高丰度值出现在冬季2、3月，丰度高达10.72×10~3cells/mL。细胞粒径大于2μm的微型光合真核生物丰度比聚球藻和微微型光合真核生物丰度要低的多，其季节变化趋势与聚球藻类似。异养细菌的季节变化波动性非常明显，最大值出现在2012年3月（9.74×10~5cells/mL）。此外，在调查过程中我们发现了一个非常特殊的现象：2010年7月12日的样品中，除了广泛存在的聚球藻、微微型和微型光合真核生物和异养细菌外，原绿球藻也大量存在，此外我们还观察到了一种与聚球藻类似的含有橙色荧光（藻红蛋白）和红色荧光（叶绿素a）、细胞粒径大于2μm的未知类群，该类群丰度高达7104events/mL。通过流式细胞仪分选后在荧光显微镜观察发现，与某些含异形胞的固氮蓝细菌藻丝体类似，它们似乎是一种呈链状、含异形胞的藻丝体，但异形胞并没有位于末端。此外，荧光显微镜照片显示该链状藻丝体是折叠状的。通常情况下，藻丝体都是与硅藻共生的，自由藻丝体很少报道。据我们所知，这种自由藻丝体的大量爆发现象从来没有被报道过。然而，由于样品量少（只有1.5mL）、分析时间晚于采样时间以及荧光显微镜照片分辨率较低等因素限制，我们难以进行物种鉴定。夏季高温引起的水体层化现象、硝酸盐的耗尽等可能与这一现象有关。马赛湾自由藻丝体短暂爆发现象的发生原因有待于进一步研究。马赛湾超微型浮游生物的监测持续进行中，有必要引进高频率原位流式细胞仪，从而避免错过类似的、短暂发生的特殊现象，能够进行进一步的物种鉴定。
与马赛湾开放性海湾不同，胶州湾属于半封闭的浅海海湾，十多条河流入湾，受人类活动的影响更为显著。我们通过2007年1月至2010年12月胶州湾月调查航次，研究了胶州湾微微型浮游生物长期的季节变化。调查航次显示，聚球藻、微微型光合真核生物以及异养细菌普遍存在于胶州湾，而原绿球藻至今尚未被观察到。在季节分布上，无论是在湾内、还是湾口和湾外，微微型浮游生物都呈现出明显的季节变化。聚球藻最高值发生在8月水温最高（>24°C）的时期，自12月到5月（水温<15°C），其丰度维持在低水平。微微型光合真核生物高丰度值出现在4-7月，除8月的其他时期，微微型光合真核生物丰度值都高于聚球藻。异养细菌每年会在8月（最高值）和4月（次高值）出现两个高丰度值，这两个高值恰恰与叶绿素高值出现的时间一致，由此可以推测二者之间存在明显关联。主成分分析结果显示胶州湾微微型浮游生物的季节变化主要受温度和营养盐（硝酸盐、亚硝酸盐、铵盐及硅酸盐等）影响。温度、营养盐与微微型浮游生物丰度之间的正相关关系，也表明了温度和营养盐对胶州湾微微型浮游生物丰度分布的重要性。
马赛湾与胶州湾的自然及人类环境既存在相似点，又有所不同。首先二者都是海湾系统，并受到人类活动的影响。马赛湾属于地中海的一部分，临近法国第二大城市马赛；而胶州湾入海口与黄海相连，受到青岛市人类活动的影响。不同点方面：马赛湾没有淡水河流的影响，胶州湾则受到十多条淡水河流的影响；此外，由于胶州湾夏季降雨量丰富，而马赛湾受到地中海气候的影响，夏季降雨量少，这些特征对于海洋超微型浮游生物的分布都会产生一定的影响。比较马赛湾和胶州湾超微型浮游生物的类群发现，聚球藻、微微型和微型光合真核生物是普遍存在的，而原绿球藻在两个海湾都很少发现。在季节分布上，马赛湾聚球藻和微型光合真核生物高丰度值出现在春季，低丰度值出现在秋季和冬季。微微型光合真核生物高丰度值出现在冬季，比聚球藻稍早；胶州湾聚球藻高丰度值出现在8月，其他月份丰度值较低，微微型和微型光合真核生物高丰度值出现在7月，比聚球藻早一个月。两个海湾超微型浮游生物细胞丰度与细胞荧光含量呈现相反的趋势，即细胞丰度高的季节，细胞较小，细胞荧光含量较低。主成分分析发现马赛湾超微型浮游生物丰度的季节变化主要受到营养盐的影响；胶州湾超微型浮游生物丰度的季节变化则是受到温度和营养盐的双重影响。
通过本论文的实验，我们研究了不同海洋生境中超微型浮游生物的群落结构和丰度变化，发现环境因子和人类活动会在不同程度上对超微型浮游生物产生影响，且不同生境中对海洋超微型浮游生物丰度的影响因子有明显的区别。
In this dissertation, temporal and spatial variations of ultraplankton (Synechococcus,Prochlorococcus, picoeukaryotes, nanoeukaryotes and heterotrophic bacteria) abundanceswere studied in different marine environments (in the East China Sea, Bay of Marseille inthe NW Mediterranean Sea and Jiaozhou Bay in the Yellow Sea) based on flow cytometryanalysis. The relationships between environmental conditions and ultraplankton abundancewere also studied in order to search for the possible controlling factors on ultraplanktondistribution in different marine environments.
Picoplankton distribution in different water masses of the East China Sea (ECS) wasinvestigated in November2006and February2007. The cruises tracked crossed three majorwater masses: Coastal Water Mass (CWM), Shelf Mixing Water Mass (MWM) andKuroshio Water Mass (KWM). Picoplankton composition was resolved in four main groupsby flow cytometry, namely Synechococcus, Prochlorococcus, picoeukaryotes, andheterotrophic bacteria. The average Synechococcus, picoeukaryotes and heterotrophicbacteria abundances were (10.63±10.88)×10~3,(1.61±1.16)×10~3,(3.39±1.27)×10~5cells/mLin autumn and (6.45±8.60)×10~3,(3.23±2.63)×10~3,(3.76±1.37)×10~5cells/mL in winter,respectively. Prochlorococcus was seldom observed in surface samples and not found inCWM, neither in autumn nor in winter. In contrast, Prochlorococcus was present in MWMand KWM with abundance at around10~3cells/mL in autumn and winter. Synechococcusdistribution showed marked variations with respect to water masses with the maximumabundance in KWM and minimum abundance in CWM. The depth-averaged integratedabundance of Synechococcus was about5-fold higher in KWM than in CWM, which mightbe mainly influenced by temperature. In autumn particularly, at several stations in MWM,Synechococcus was resolved as two subgroups. Picoeukaryotes were less affected bywater-mass differences than Synechococcus. Heterotrophic bacteria depth-averaged integrated abundance exhibited the smallest seasonal variations with respect to watermasses. Correlation analysis showed obvious differences between picoplankton abundancesand environmental parameters (temperature, nutrients and chlorophyll a) in three watermasses, indicated the three water masses are likely to have different influences onpicoplankton (particularly Synechococcus) distribution.
In the Bay of Marseille (NW Mediterranean), the composition of ultraplankton (<10μm) sampled in surface (1m depth) was investigated from May2010to April2012.Ultraplankton (<10μm) composition was resolved in four main groups by flow cytometry,namely Synechococcus, picoeukaryotes, nanoeukaryotes and heterotrophic bacteria.Prochlorococcus was seldom observed in our study, even though in the previous studies,Prochlorococcus has been widely reported. In the Bay of Marseille, Synechococcusdominated the ultraphytoplankton community in terms of cell abundance. Synechococcusshowed a clear seasonal pattern with higher abundance in spring (April and May,>90×10~3cells/mL) and lower abundance in autumn and winter (<15×10~3cells/mL). Synechococcusabundance varied more than40-fold between a spring maximum (May2010,129.59×10~3cells/mL) and winter minimum (March2012,2.29×10~3cells/mL) values. Picoeukaryoteshad a similar distribution pattern as Synechococcus and their high abundance occurred inwinter (January-March,>10.72×10~3cells/mL), slightly earlier than Synechococcus.Nanoeukaryotes exhibited much lower abundance than Synechocccus and picoeukaryotes.The distribution pattern of nanoeukaryotes was similar to that of Synechococcus. FromMay to September in2010, the data of heterotrophic bacteria abundance were missingbecause of technical reasons. Heterotrophic bacteria exhibited pronounced fluctuationsduring the study period and reached their maximum abundance (9.74×10~5cells/mL) inMarch2012. On12July2010, in addition to groups of Synechococcus, Prochlorococcus,picoeukaryotes, and nanoeukaryotes, an unidentified additional cluster was observed.These unknown particles were sorted out by flow cytometry and then observed byepifluorescence microscopy: they appeared to be chain-forming microorganisms, just liketrichomes of some diazotroph cyanobacteria with one heterocyst, but not unambiguouslydistal. The chains observed after cell sorting were not straight, but rather folded. The shortoccurrence of free trichomes in the Bay of Marseille with such a high abundance remains to be explained. High temperature in summer, induced stratification of the water column,and nitrate depletion, may be related to this event. High frequency survey ofultraphytoplankton assemblages at the single cell level appears necessary to observe moreefficiently such phenomenon and document more extensively its full dynamics.
The seasonal variability of picoplankton abundances in the semi-enclosed Jiaozhou Bay(Yellow Sea) were investigated monthly from January2007to December2010by flowcytometic analysis. Picoplankton community in the studied area was comprised ofSynechococcus, picoeukaryotes and heterotrophic bacteria. Prochlorococcus was notdetected at any station during the whole year. A pronounced seasonality was observed inpicoplankton community structure and abundances inside the bay, in the Bay mouth andoutside the bay. Synechococcus exhibited a clear seasonal pattern, with maximumabundance occurring in August when temperature was highest and minimum abundance inwinter and spring. The maximum abundance of picoeukaryotes occurred slightly earlier(April-July) than Synechococcus. Picoeukaryotes abundance outnumbered Synechococcusin most of the year except in August. Heterotrophic bacteria exhibited two peaks ofabundances in August (major peak) and April (minor peak), which were quite similar asChlorophyll a distribution. Principal Component Analysis (PCA) suggested that seasonalvariations of picoplankton were mainly driven by temperature and nutrient concentrations(NO3-, NO2-, NH4-and SiO32-). The significantly positive correlations betweentemperature/nutrients and picoplankton abundances also implied the importance oftemperature and nutrients in picoplankton distribution in Jiaozhou Bay.
The ultraphytoplankton composition, abundance and fluorescence in the Bay ofMarseille and Jiaozhou were compared in2011and2012. In the Bay of Marseille andJiaozhou, Synechococcus, picoeukaryotes and nanoeukaryotes were observed in all thesamples. Prochlorococcus was seldom observed in the Bay of Marseille and neverobserved in the Jiaozhou Bay. Besides, in the Bay of Marseille, a possible bloom of freetrichomes was observed in a short period during summer2010. While in the Jiaozhou Bay,an unknown and abundant cluster was obsvered in all the samples. With flow cytometryand scanning electron microscopy, it is difficult to identify this species. In the next step,molecular technique should be introduced for further identification. In the seasonal scale, ultraphytoplankton in the Bay of Marseille and Jiaohzou exhibited quite differentdistribution patterns. In the Bay of Marseille, Synechococcus and nanoeukaryotes presentedthe maximum abundance in spring and minimum abundances in autumn and winter.Picoeukaryotes showed the maximum abundance in winter, slightly earlier thanSynechococcus and nanoeukaryotes. While in the Jiaozhou Bay, Synechococcus abundancewas highest in August (T>24°C) and lowest in winter and spring (T<15°C). Picoeukaryotesand nanoeukaryotes exhibited highest abundances in July, one month earlier thanSynechococcus, and lower abundances in winter and spring, similar as Synechococcus. Itseemed in the oligotrophic Bay of Marseille, nutrient supply is more important forultraphytoplantkon distribution, while in the Jiaozhou Bay, temperature is the keycontrolling factor for ultraphytoplantkon distribution. In both bays, cell abundanceexhibited a clear opposite trend compared to cellular fluorescence.
引文
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