泛三大洋微微型浮游生物的分布及其与环境因子的相关性
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摘要
海洋微微型浮游生物是地球上种类最丰富，数量最多的有机体，介导着生物地球化学循环过程，在海洋生态系统中起着极其重要的作用。国际上有关微微型浮游植物、异养细菌和浮游病毒在大尺度范围的分布规律报道很少，且在不同海洋环境条件下，微微型浮游植物、异养细菌和浮游病毒之间的相关性及其与环境因子之间的相关性，特别是浮游病毒与宿主细胞间的相关性研究更少。因此，本论文采用流式细胞仪计数法对西太平洋、印度洋、环南大洋及南极水域原绿球藻、聚球藻、微微型真核浮游植物、异养细菌和浮游病毒丰度的分布特点进行了研究，并利用PCA和Pearson相关系数法分析了微微型浮游生物与水温、盐度、营养盐（磷酸盐、硝酸盐+亚硝酸盐和硅酸盐）等环境因子之间的相互关系，同时还分析了不同水域各微微型浮游生物之间的相关性。本文还利用RFLP技术对南极长城湾微微型真核浮游生物的分子多样性进行了分析。本文的研究有助于探讨微微型浮游生物的分布规律、各生物因子之间及环境因子对其分布的影响，并为海洋生态环境的监测、预警及修复提供基础的数据资料。
（1）原绿球藻在西太平洋和印度洋的分布分别延伸至~50°S和~46°S，限制其生长的最低温度分别为9.7°C和11.8°C。在赤道地区丰度最高，随着赤道地区向高纬度水体延伸，丰度逐渐减低直至消失。原绿球藻在西太平洋和印度洋水域的丰度分别为~5.4×10~3–4.20×10~5个mL~(-1)和1.45×10~4–3.73×10~5个mL~(-1)。生物量分别为0.16–15.64mg m-3和0.20–11.6mg m-3。在赤道附近海域，原绿球藻对浮游植物生物量的贡献率达到~52.2%，是浮游植物生物量的主要贡献者。在西太平洋，影响原绿球藻分布的主要因素是温度和营养盐（磷酸盐和硅酸盐），与温度和较低的磷酸盐浓度成显著的正相关性(p<0.01)；与较高的硅酸盐浓度成负相关性(p<0.01)；在印度洋，由于研究海域营养盐水平较高，原绿球藻主要受温度的影响，其生物量与温度呈显著正相关性(p<0.01)。
（2）聚球藻在西太平洋和印度洋的分布分别延伸至57.1°S和54.1°S，限制它们生长的最低温度分别为4.1°C和6.4°C。中纬度水域聚球藻丰度高于低纬度。聚球藻在西太平洋和印度洋水域的丰度分别为1.10×10~3–2.29×10~5个mL~(-1)和1.54×10~3–2.79×10~5个mL~(-1)。生物量分别为0.26–17.74mg m~(-3)和0.3~5–12.93mgm-3。在西太平洋，聚球藻与温度呈显著负相关性(p<0.01)，与硝酸盐＋亚硝酸盐和硅酸盐成显著正相关性(p<0.01)。在印度洋，聚球藻主要受温度的影响，其生物量与温度呈显著正相关性(p<0.01)。
（3）微微型真核浮游植物广泛存在于所研究的海域，在中纬度水域丰度高于低纬度，在高纬度南极水域丰度更低。在南极水域，大陆架水体微微型真核浮游植物的丰度明显低于远洋水体；在垂直分布上，随着水深的增加，它的丰度逐渐减低，直至500m水深完全消失。它在西太平洋、印度洋、环南大洋的丰度分别为5.2×102–4.68×10~4个mL~(-1)，4.9×102–1.73×10~4个mL~(-1)，7.6×102–1.04×10~4个mL~(-1)和1.42×102–4.00×10~3个mL~(-1)。生物量分别为0.26–17.74mg m-3，0.36–29.8mg m-3，0.99–14.1mg m-3。在中纬度水域，微微型真核浮游植物是浮游植物生物量的主要贡献者。在不同大洋，影响微微型真核浮游植物分布的因素不同。在西太平洋，微微型真核浮游植物的分布主要受营养盐影响，与硝酸盐＋亚硝酸盐和硅酸盐成显著正相关性(p<0.01)。在印度洋，它主要受温度的影响，其生物量与温度呈显著正相关性(p<0.01)；在环南大洋，磷酸盐和温度是影响微微型真核浮游植物分布的主要因素。微微型真核浮游植物生物量与磷酸盐成显著的正相关性(p<0.01)，与温度成显著的负相关性(p<0.01)。在南极水域，微微型真核浮游植物丰度与水深、盐度、溶解氧、硝酸盐＋亚硝酸盐、磷酸盐和硅酸盐浓度都有显著的负相关性(p<0.01)。
（4）异养细菌广泛存在于所研究的海域，在中纬度水域丰度高于低纬度，在高纬度南极水域丰度更低。在南极水域，大陆架水体的异养细菌丰度明显低于远洋水体；在水深较浅的断面，异养细菌丰度在垂直分布上无显著差异，在水深较深的断面，在深层水体的丰度明显低于上层水体。在西太平洋、印度洋、环南大洋和南极水域的丰度分别为2.27×10~5–3.80×10~6个mL~(-1)，2.6~5×10~5–1.27×10~6个mL~(-1)，8.4×10~5–1.32×10~6个mL~(-1)和1.0~5×10~5–3.69×10~6个mL~(-1)。生物量分别为4.~54–79.55mg m-3，5.31–25.4mg m~(-3)，1.68–26.4mg m~(-3)和2.10–73.8mg m~(-3)。在西太平洋，异养细菌的分布主要受营养盐影响，与硝酸盐＋亚硝酸盐和硅酸盐成显著正相关性(p<0.01)。在印度洋，异养细菌生物量与硝酸盐＋亚硝酸盐浓度呈显著的负相关性(p<0.01)。在南极水域，异养细菌丰度与深度、盐度、磷酸盐和硅酸盐成显著负相关性(p<0.01)。
（5）浮游病毒广泛存在于所调查的海域，在水平分布上，浮游病毒的丰度与宿主细胞的丰度分布存在密切的联系。在赤道水域存在高值区，与原绿球藻的丰度分布趋势一致；而在中纬度水域，其高值区与聚球藻、微微型真核浮游植物和异养细菌的丰度分布一致。在南极水域浮游病毒在2000m以下水层均有分布，其垂直分布规律与异养细菌类似。在西太平洋、印度洋、环南大洋和南极水域的丰度分别为2.17×10~6–2.49×10~7个mL~(-1)，3.80×10~6–2.41×10~7个mL~(-1)，1.46×10~6–2.98×10~7个mL~(-1)和1.49×10~6–4.39×10~7个mL~(-1)。生物量分别为0.42–4.95mg m~(-3)，0.76–4.82mg m~(-3)，0.11–13.3mg m~(-3)和0.25–8.78mg m~(-3)。在西太平洋，浮游病毒与微微型真核浮游植物、聚球藻和异养细菌生物量之间存在显著的正相关性(p<0.01)，但在赤道水域与原绿球藻生物量存在显著正相关性(p<0.01)；浮游病毒生物量与硝酸盐＋亚硝酸盐和硅酸盐成显著正相关性(p<0.01)，与温度成显著的负相关性(p<0.01)。在印度洋和环南大洋，浮游病毒的分布仅受异养细菌的影响，与异养细菌生物量成显著正相关性(p<0.01)。在南极水域，浮游病毒与叶绿素a、微微型真核浮游植物丰度及异养细菌丰度成显著正相关性(p<0.01)，与水深、盐度、磷酸盐和硅酸盐成显著负相关性(p<0.01)。本文研究表明浮游病毒的分布主要受各海域浮游生物中主要优势类群丰度变化的影响，而营养盐等环境因子通过影响病毒宿主的生长从而间接地影响浮游病毒的空间分布。温度、盐度可能直接或间接地影响病毒的分布。
（6）不同海域，不同类型的微微型浮游生物对海洋生态系统碳循环所贡献的生物量不同。在西太平洋和环南大洋，异养细菌生物量在微微型浮游生物中占据主导地位。在印度洋，微微型真核浮游植物生物量在微微型浮游生物中占主导地位。本文还发现微微型浮游生物存在拮抗和协调生长的作用。一方面，原绿球藻丰度和聚球藻、微微型真核浮游植物丰度成显著的负相关性(p<0.01)，表明原绿球藻与这2者之间存在拮抗作用。在低纬度水域，水温高，营养盐（特别是磷酸盐浓度）较低，体积较小的原绿球藻有更强的生长竞争优势，丰度升高，而聚球藻和微微型真核浮游植物丰度降低；而在中纬度水域，水温降低，营养盐浓度升高，较大个体的藻类生长占优势地位，聚球藻和微微型真核浮游植物丰度升高，而不占竞争优势的原绿球藻丰度急剧降低。另一方面，异养细菌丰度和聚球藻、微微型真核浮游植物丰度之间存在着显著的正相关性(p<0.01)，表明异养细菌与聚球藻、微微型真核浮游植物存在协同生长的作用。在中纬度水域，无机营养盐丰富，聚球藻和微微型真核浮游植物利用无机营养物生长，同时将无机营养物转化为有机营养成分，导致了水体中有机营养物的增加。这些增加的有机营养物被异养细菌利用，异养细菌快速生长，导致在中纬度水域异养细菌丰度随之升高。同时，异养细菌又将制造出大量的无机营养物质，这些物质又可以维持聚球藻和微微型真核浮游植物的生长。（7）本文运用PCR-RFLP技术，在国际上首次揭示了南极长城湾海域微微型真核浮游生物的类群，获得了南极长城湾38个微微型真核浮游生物OTUs，主要类群为横裂甲藻纲(Dinophyceae)、真菌类(Fungi)、丝足虫类(Cercozoa)和纤毛虫亚纲(Ciliates)和未获培养的海洋微微型真核浮游生物。各类群的OTUs数分别为2个、16个、~5个、2个和13个，未获培养的海洋微微型真核浮游生物占总OTUs数的33.4%。
Picoplankton plays an important role in marine ecostystem. Over the past decades,much effort has been devoted to understand picoplankton abundance and their spatialdistribution in Pacific Ocean, India Ocean and the encycling southern Ocean.However，distribution of picophytoplankton, heterotrophic bacteria and viruses onlarge-scale pelagic investigations is practically rare. The correlation amongpicophytoplankton, heterotrophic bacteria and viruses and their relationships withenvironmental factors in different marine environmental conditions, especially thecorrelation between viruses and their host cells are rarer. Therefore, Flow cytometrywas used to study the spatial distribution pattern of Prochlorococcus (Pro),Synechococcus (Syn), Picoeukaryotes (Euk), heterotrophic bacteria and virioplanktonin Pacific Ocean, India Ocean, the encircling southern Ocean and Antarctic waters.PCA (Principal component analysis) and Pearson correlation coefficient analysiswere also used to analyze the relationships between picoplankton and watertemperature, salinity, nutrients (phosphate, nitrate+nitrite and silicate), dissolvedoxygen in different marine environmental conditions and study the relationshipsbetween viruses and their host cells. RFLP (Restriction Fragment LengthPolymorphism) method was also applied to analyze genetic diversity of Eukaryoticpicoplankton in the Great Wall Bay, Antarctica. It hopes to investigate thedistribution characteristics of picoplankton and their relationship with environmentalfactors, in order to provide the basic data for monitoring, early warning and repairingthe marine environment.(1) In this paper, spatial distribution of Pro, Syn, Euk, heterotrophic bacteria and virioplankton were detected in the waters (30°N–69°S,155°E–69°W). Euk, heterotrophic bacteria and virioplankton are distributedwidespreadly in the surveyed area, while pico-photosynthetic prokaryotes are not.The latitudinal distribution of pico-photosynthetic prokaryotes was limited by watertemperature. Pro in the western Pacific and the Indian Ocean was distributed to50°Sand46°S from the north, respectively. The minimum temperature limiting theirgrowth was9.7°C and11.8°C, respectively. Pro abundance was the highest in theequatorial regions, it decreased gradually as the latitude increases and finally itdisappears. The range of Pro abundance in the western Pacific and the Indian Oceanwere5.4×10~3-4.20×10~5cells mL~(-1)and1.45×10~4-3.73×10~5cells mL~(-1),respectively. The Biomasses in the western Pacific and the Indian Ocean were0.16-15.64mg m~(-3)and0.20-11.6mg m~(-3), respectively. Pro made up55.2%of theestimated phytoplankton biomass in the equatorial regions and accounted for themain contributor to phytoplankton biomass in the Western Pacific, In the WesternPacific, the major factors affecting the distribution of Pro were temperature andnutrients. Pro biomass correlated positively with temperature and low phosphate(p<0.01), and negatively with high silicate concentration (p<0.01); Due to highlevels of nutrient waters in the Indian Ocean, Pro was mainly affected by temperatureand Pro biomass were significantly positively correlated with temperature (p <0.01).
(2) Syn in the western Pacific and Indian Ocean distribution was extended to55°S and54.1°S, respectively. The minimum temperatures limiting their growth were4.1°C and6.4°C, respectively. Syn abundance in Mid-latitude waters was higherthan that in the low-latitude waters. Syn in the western Pacific and Indian Oceanwaters abundance were1.10×10~3–2.29×10~5cells mL~(-1)and1.54×10~3–2.79×10~5cells mL~(-1), respectively. The Biomasses were0.26–17.74mg m~(-3)and0.35–12.93mg m~(-3), respectively. In the Western Pacific, Syn was significantly and negativelycorrelated with temperature (p <0.01), and significantly and positively with nitrate+nitrite and silicate (p <0.01). In the Indian Ocean, Syn was mainly affected bytemperature, and syn biomass were significantly and positively correlated withtemperature (p <0.01).
3) Euk abundance in mid-latitude waters was higher than that of low-latitude watersand the lowest in the high-latitude Antarctic waters. Euk abundance in the continentalshelf waters was lower than that of the open waters. Euk abundance in the westernPacific, Indian Ocean, Central Southern Ocean and Antarctic waters were5.2×102–4.68×10~4cells mL~(-1),4.9×102–1.73×10~4cells mL~(-1),7.6×10~2–1.04×10~4cells mL~(-1)and1.42×10~2–4.00×10~3cells mL~(-1), respectively. Their biomasses were0.26–17.74mg m~(-3)，0.36–29.8mg m~(-3)，0.99–14.1mg m~(-3)and0.06–3.21mg m~(-3),respectively. Spatial distribution of Euk was effected by different environmentalfactors in different oceans. In the Western Pacific, the distribution of Euk wasprimarily affected by nutrients and Euk biomass showed a significant and positivecorrelation with nitrate+nitrite and silicate (p <0.01). In the Indian Ocean, Euk ismainly influenced by temperature and Euk biomass was significantly and positivelycorrelated with temperature (p <0.01); in the encircling Southern Ocean, Euk biomassshowed a significant and positive correlation with phosphate (p <0.01), and asignificant and negative correlation with temperature (p <0.01). In Antarctic waters,eukaryotic phytoplankton abundance showed significant and negative correlationswith water depth, salinity, nitrate+nitrite, phosphate and silicate concentrations (p<0.01).
(4) The abundance of heterotrophic bacteria was the lowest in Antarctic waters, Theanalysis on vertical profile of bacterial abundance indicated that there was nosignificant decrease (p>0.05) in the IS Transect, which depth is shallow. Theirabundance in the bottom layer was lower than in the upper layers in those deeptransects, which the viral vertical distribution was similar to in Antarctic waters. Theabundance in the Western Pacific, the Indian Ocean, Southern Ocean and Antarcticwaters were2.27×10~5–3.80×10~6cells mL~(-1),2.65×10~5–1.27×10~6cells mL~(-1),8.4×10~5–1.32×10~6个mL~(-1)and1.05×10~5–3.69×10~6cells mL~(-1), respectively. The biomasseswere4.54–79.55mg m~(-3),5.31–25.4mg m~(-3),1.68–26.4mg m~(-3)and2.10–73.8mgm~(-3), respectively. In the Western Pacific, spatial distribution of heterotrophic bacteriais mainly affected by nutrients and bacteria biomass was significantly and positivelycorrelated with nitrate+nitrite and silicate (p <0.01). In the Indian Ocean, heterotrophic bacteria biomass was significantly and negetively correlated withnitrate+nitrite concentration (p <0.01). In Antarctic waters, heterotrophic bacteriaabundance showed a significant and negative correlation with water depth, salinity,phosphate and silicate (p <0.01).
(5) Viral abundance in the western Pacific, Indian Ocean, encircling Southern Oceanand Antarctic waters were2.17×10~6–2.49×10~7particles mL~(-1),3.80×10~6–2.41×10~7particles mL~(-1),1.46×10~6–2.98×10~7particles mL~(-1)and1.49×10~6–4.39×10~7particlesmL~(-1), respectively. The biomasses were0.42–4.95mg m~(-3),0.76–4.82mg m~(-3),0.11–13.3mg m~(-3)and0.25–8.78mg m~(-3), respectively. The spatial distribution ofvirus was mainly affected by the distribution of the host cells. In the western Pacific,viral distribution is affected by host cells and environmental factors. Viral biomasscorrelated positively with the Syn, Euk, heterotrophic bacteria biomass, nitrate+nitrite and silicate (p<0.01), and negatively with temperature (p<0.01). In the IndianOcean and Southern Ocean, the distribution of viruses were only affected byheterotrophic bacterial and viral biomass showed a significant and positivecorrelation with heterotrophic bacteria biomass (p <0.01). In Antarctic waters, viralabundance was significantly and positively correlated with chlorophyll a, eukabundance and bacteria abundance (p <0.01), and significantly and negativelycorrelated with water depth, salinity, phosphate and silicate were (p <0.01).
(6) According to analyze the biomass of each group of picoplankton in the westernPacific, Indian Ocean and encircling Southern Ocean (including the three partitions ofthe Pacific, Indian Ocean and the Atlantic), it was found that heterotrophic bacteriaaccounted for the vast majority of the estimated picoplankton biomass in the Pacificand Atlantic, while the contribution of Euk to total picoplankton biomass dominatedin the Indian Ocean. Our results also suggested that antagonistic and synergisticeffects existed among the groups of picoplankton. In one hand, Pro showed asignificant and negative correlation with Syn, Euk and heterotrophic bacteria (p<0.01), which suggested that pro showed the antagonistic effects with Syn and Euk.In the low-latitude waters, Pro abundance was higher than that of Syn and Eukbecause of high temperature and low nutrients. In the mid-latitude waters, it benifited for the growth of the bigger algae cells with the nutrients increased, then Euk and Synabundance increased while pro growth was limited and its abundance decreased. Onthe other hand, bacteria showed a significant and positive correlation with Syn andEuk (p <0.01), which which suggested that bacteria showed the synergistic effectswith Syn and Euk. In the mid-latitude waters, Syn and Euk utilized inorganicnutrients for the purpose of growth and thereby changed them into organic nutrients,this then leaded to an increase in the level of organic material. This increased level oforganic nutrients was then utilized to the benefit of heterotrophic bacteria, wherebacterial growth turned organic into inorganic matter. Conversely, the producedinorganic nutrients of bacterial origin could further sustain the growth of Syn andEuk.
(7) In the present study, molecular tools were used to investigate the marineeukaryotic communities of Great Wall Bay, Antarctica.18s rDNA genetic library wasconstructed, and then the library of eukaryotic ribosomal DNA were screened byrestriction fragment length polymorphism analysis. The phylogenetic diversity in thelibrary was rather great,430positive clones and38OTUs have been isolated, whichindicates that hat the microeukaryotes in this area are diverse. The38OTUs arefound to distribute in Fungi, Dinophyceae, Cercozoa Ciliates, and Uncultured marineeukaryote clones. The microeukaryotic groups are arranged from the most diverse tothe least diverse as: uncultured Dinophyceae, Fungi, Cercozoa, Ciliates and marinemicroeukaryotes. And the number of OTU of these groups is:2,16,5,2and13,respectively. And the uncultured marine microeukaryotes amounts33.4%of the totalnumbers.
（1）原绿球藻在西太平洋和印度洋的分布分别延伸至~50°S和~46°S，限制其生长的最低温度分别为9.7°C和11.8°C。在赤道地区丰度最高，随着赤道地区向高纬度水体延伸，丰度逐渐减低直至消失。原绿球藻在西太平洋和印度洋水域的丰度分别为~5.4×10~3–4.20×10~5个mL~(-1)和1.45×10~4–3.73×10~5个mL~(-1)。生物量分别为0.16–15.64mg m-3和0.20–11.6mg m-3。在赤道附近海域，原绿球藻对浮游植物生物量的贡献率达到~52.2%，是浮游植物生物量的主要贡献者。在西太平洋，影响原绿球藻分布的主要因素是温度和营养盐（磷酸盐和硅酸盐），与温度和较低的磷酸盐浓度成显著的正相关性(p<0.01)；与较高的硅酸盐浓度成负相关性(p<0.01)；在印度洋，由于研究海域营养盐水平较高，原绿球藻主要受温度的影响，其生物量与温度呈显著正相关性(p<0.01)。
（2）聚球藻在西太平洋和印度洋的分布分别延伸至57.1°S和54.1°S，限制它们生长的最低温度分别为4.1°C和6.4°C。中纬度水域聚球藻丰度高于低纬度。聚球藻在西太平洋和印度洋水域的丰度分别为1.10×10~3–2.29×10~5个mL~(-1)和1.54×10~3–2.79×10~5个mL~(-1)。生物量分别为0.26–17.74mg m~(-3)和0.3~5–12.93mgm-3。在西太平洋，聚球藻与温度呈显著负相关性(p<0.01)，与硝酸盐＋亚硝酸盐和硅酸盐成显著正相关性(p<0.01)。在印度洋，聚球藻主要受温度的影响，其生物量与温度呈显著正相关性(p<0.01)。
（3）微微型真核浮游植物广泛存在于所研究的海域，在中纬度水域丰度高于低纬度，在高纬度南极水域丰度更低。在南极水域，大陆架水体微微型真核浮游植物的丰度明显低于远洋水体；在垂直分布上，随着水深的增加，它的丰度逐渐减低，直至500m水深完全消失。它在西太平洋、印度洋、环南大洋的丰度分别为5.2×102–4.68×10~4个mL~(-1)，4.9×102–1.73×10~4个mL~(-1)，7.6×102–1.04×10~4个mL~(-1)和1.42×102–4.00×10~3个mL~(-1)。生物量分别为0.26–17.74mg m-3，0.36–29.8mg m-3，0.99–14.1mg m-3。在中纬度水域，微微型真核浮游植物是浮游植物生物量的主要贡献者。在不同大洋，影响微微型真核浮游植物分布的因素不同。在西太平洋，微微型真核浮游植物的分布主要受营养盐影响，与硝酸盐＋亚硝酸盐和硅酸盐成显著正相关性(p<0.01)。在印度洋，它主要受温度的影响，其生物量与温度呈显著正相关性(p<0.01)；在环南大洋，磷酸盐和温度是影响微微型真核浮游植物分布的主要因素。微微型真核浮游植物生物量与磷酸盐成显著的正相关性(p<0.01)，与温度成显著的负相关性(p<0.01)。在南极水域，微微型真核浮游植物丰度与水深、盐度、溶解氧、硝酸盐＋亚硝酸盐、磷酸盐和硅酸盐浓度都有显著的负相关性(p<0.01)。
（4）异养细菌广泛存在于所研究的海域，在中纬度水域丰度高于低纬度，在高纬度南极水域丰度更低。在南极水域，大陆架水体的异养细菌丰度明显低于远洋水体；在水深较浅的断面，异养细菌丰度在垂直分布上无显著差异，在水深较深的断面，在深层水体的丰度明显低于上层水体。在西太平洋、印度洋、环南大洋和南极水域的丰度分别为2.27×10~5–3.80×10~6个mL~(-1)，2.6~5×10~5–1.27×10~6个mL~(-1)，8.4×10~5–1.32×10~6个mL~(-1)和1.0~5×10~5–3.69×10~6个mL~(-1)。生物量分别为4.~54–79.55mg m-3，5.31–25.4mg m~(-3)，1.68–26.4mg m~(-3)和2.10–73.8mg m~(-3)。在西太平洋，异养细菌的分布主要受营养盐影响，与硝酸盐＋亚硝酸盐和硅酸盐成显著正相关性(p<0.01)。在印度洋，异养细菌生物量与硝酸盐＋亚硝酸盐浓度呈显著的负相关性(p<0.01)。在南极水域，异养细菌丰度与深度、盐度、磷酸盐和硅酸盐成显著负相关性(p<0.01)。
（5）浮游病毒广泛存在于所调查的海域，在水平分布上，浮游病毒的丰度与宿主细胞的丰度分布存在密切的联系。在赤道水域存在高值区，与原绿球藻的丰度分布趋势一致；而在中纬度水域，其高值区与聚球藻、微微型真核浮游植物和异养细菌的丰度分布一致。在南极水域浮游病毒在2000m以下水层均有分布，其垂直分布规律与异养细菌类似。在西太平洋、印度洋、环南大洋和南极水域的丰度分别为2.17×10~6–2.49×10~7个mL~(-1)，3.80×10~6–2.41×10~7个mL~(-1)，1.46×10~6–2.98×10~7个mL~(-1)和1.49×10~6–4.39×10~7个mL~(-1)。生物量分别为0.42–4.95mg m~(-3)，0.76–4.82mg m~(-3)，0.11–13.3mg m~(-3)和0.25–8.78mg m~(-3)。在西太平洋，浮游病毒与微微型真核浮游植物、聚球藻和异养细菌生物量之间存在显著的正相关性(p<0.01)，但在赤道水域与原绿球藻生物量存在显著正相关性(p<0.01)；浮游病毒生物量与硝酸盐＋亚硝酸盐和硅酸盐成显著正相关性(p<0.01)，与温度成显著的负相关性(p<0.01)。在印度洋和环南大洋，浮游病毒的分布仅受异养细菌的影响，与异养细菌生物量成显著正相关性(p<0.01)。在南极水域，浮游病毒与叶绿素a、微微型真核浮游植物丰度及异养细菌丰度成显著正相关性(p<0.01)，与水深、盐度、磷酸盐和硅酸盐成显著负相关性(p<0.01)。本文研究表明浮游病毒的分布主要受各海域浮游生物中主要优势类群丰度变化的影响，而营养盐等环境因子通过影响病毒宿主的生长从而间接地影响浮游病毒的空间分布。温度、盐度可能直接或间接地影响病毒的分布。
（6）不同海域，不同类型的微微型浮游生物对海洋生态系统碳循环所贡献的生物量不同。在西太平洋和环南大洋，异养细菌生物量在微微型浮游生物中占据主导地位。在印度洋，微微型真核浮游植物生物量在微微型浮游生物中占主导地位。本文还发现微微型浮游生物存在拮抗和协调生长的作用。一方面，原绿球藻丰度和聚球藻、微微型真核浮游植物丰度成显著的负相关性(p<0.01)，表明原绿球藻与这2者之间存在拮抗作用。在低纬度水域，水温高，营养盐（特别是磷酸盐浓度）较低，体积较小的原绿球藻有更强的生长竞争优势，丰度升高，而聚球藻和微微型真核浮游植物丰度降低；而在中纬度水域，水温降低，营养盐浓度升高，较大个体的藻类生长占优势地位，聚球藻和微微型真核浮游植物丰度升高，而不占竞争优势的原绿球藻丰度急剧降低。另一方面，异养细菌丰度和聚球藻、微微型真核浮游植物丰度之间存在着显著的正相关性(p<0.01)，表明异养细菌与聚球藻、微微型真核浮游植物存在协同生长的作用。在中纬度水域，无机营养盐丰富，聚球藻和微微型真核浮游植物利用无机营养物生长，同时将无机营养物转化为有机营养成分，导致了水体中有机营养物的增加。这些增加的有机营养物被异养细菌利用，异养细菌快速生长，导致在中纬度水域异养细菌丰度随之升高。同时，异养细菌又将制造出大量的无机营养物质，这些物质又可以维持聚球藻和微微型真核浮游植物的生长。（7）本文运用PCR-RFLP技术，在国际上首次揭示了南极长城湾海域微微型真核浮游生物的类群，获得了南极长城湾38个微微型真核浮游生物OTUs，主要类群为横裂甲藻纲(Dinophyceae)、真菌类(Fungi)、丝足虫类(Cercozoa)和纤毛虫亚纲(Ciliates)和未获培养的海洋微微型真核浮游生物。各类群的OTUs数分别为2个、16个、~5个、2个和13个，未获培养的海洋微微型真核浮游生物占总OTUs数的33.4%。
Picoplankton plays an important role in marine ecostystem. Over the past decades,much effort has been devoted to understand picoplankton abundance and their spatialdistribution in Pacific Ocean, India Ocean and the encycling southern Ocean.However，distribution of picophytoplankton, heterotrophic bacteria and viruses onlarge-scale pelagic investigations is practically rare. The correlation amongpicophytoplankton, heterotrophic bacteria and viruses and their relationships withenvironmental factors in different marine environmental conditions, especially thecorrelation between viruses and their host cells are rarer. Therefore, Flow cytometrywas used to study the spatial distribution pattern of Prochlorococcus (Pro),Synechococcus (Syn), Picoeukaryotes (Euk), heterotrophic bacteria and virioplanktonin Pacific Ocean, India Ocean, the encircling southern Ocean and Antarctic waters.PCA (Principal component analysis) and Pearson correlation coefficient analysiswere also used to analyze the relationships between picoplankton and watertemperature, salinity, nutrients (phosphate, nitrate+nitrite and silicate), dissolvedoxygen in different marine environmental conditions and study the relationshipsbetween viruses and their host cells. RFLP (Restriction Fragment LengthPolymorphism) method was also applied to analyze genetic diversity of Eukaryoticpicoplankton in the Great Wall Bay, Antarctica. It hopes to investigate thedistribution characteristics of picoplankton and their relationship with environmentalfactors, in order to provide the basic data for monitoring, early warning and repairingthe marine environment.(1) In this paper, spatial distribution of Pro, Syn, Euk, heterotrophic bacteria and virioplankton were detected in the waters (30°N–69°S,155°E–69°W). Euk, heterotrophic bacteria and virioplankton are distributedwidespreadly in the surveyed area, while pico-photosynthetic prokaryotes are not.The latitudinal distribution of pico-photosynthetic prokaryotes was limited by watertemperature. Pro in the western Pacific and the Indian Ocean was distributed to50°Sand46°S from the north, respectively. The minimum temperature limiting theirgrowth was9.7°C and11.8°C, respectively. Pro abundance was the highest in theequatorial regions, it decreased gradually as the latitude increases and finally itdisappears. The range of Pro abundance in the western Pacific and the Indian Oceanwere5.4×10~3-4.20×10~5cells mL~(-1)and1.45×10~4-3.73×10~5cells mL~(-1),respectively. The Biomasses in the western Pacific and the Indian Ocean were0.16-15.64mg m~(-3)and0.20-11.6mg m~(-3), respectively. Pro made up55.2%of theestimated phytoplankton biomass in the equatorial regions and accounted for themain contributor to phytoplankton biomass in the Western Pacific, In the WesternPacific, the major factors affecting the distribution of Pro were temperature andnutrients. Pro biomass correlated positively with temperature and low phosphate(p<0.01), and negatively with high silicate concentration (p<0.01); Due to highlevels of nutrient waters in the Indian Ocean, Pro was mainly affected by temperatureand Pro biomass were significantly positively correlated with temperature (p <0.01).
(2) Syn in the western Pacific and Indian Ocean distribution was extended to55°S and54.1°S, respectively. The minimum temperatures limiting their growth were4.1°C and6.4°C, respectively. Syn abundance in Mid-latitude waters was higherthan that in the low-latitude waters. Syn in the western Pacific and Indian Oceanwaters abundance were1.10×10~3–2.29×10~5cells mL~(-1)and1.54×10~3–2.79×10~5cells mL~(-1), respectively. The Biomasses were0.26–17.74mg m~(-3)and0.35–12.93mg m~(-3), respectively. In the Western Pacific, Syn was significantly and negativelycorrelated with temperature (p <0.01), and significantly and positively with nitrate+nitrite and silicate (p <0.01). In the Indian Ocean, Syn was mainly affected bytemperature, and syn biomass were significantly and positively correlated withtemperature (p <0.01).
3) Euk abundance in mid-latitude waters was higher than that of low-latitude watersand the lowest in the high-latitude Antarctic waters. Euk abundance in the continentalshelf waters was lower than that of the open waters. Euk abundance in the westernPacific, Indian Ocean, Central Southern Ocean and Antarctic waters were5.2×102–4.68×10~4cells mL~(-1),4.9×102–1.73×10~4cells mL~(-1),7.6×10~2–1.04×10~4cells mL~(-1)and1.42×10~2–4.00×10~3cells mL~(-1), respectively. Their biomasses were0.26–17.74mg m~(-3)，0.36–29.8mg m~(-3)，0.99–14.1mg m~(-3)and0.06–3.21mg m~(-3),respectively. Spatial distribution of Euk was effected by different environmentalfactors in different oceans. In the Western Pacific, the distribution of Euk wasprimarily affected by nutrients and Euk biomass showed a significant and positivecorrelation with nitrate+nitrite and silicate (p <0.01). In the Indian Ocean, Euk ismainly influenced by temperature and Euk biomass was significantly and positivelycorrelated with temperature (p <0.01); in the encircling Southern Ocean, Euk biomassshowed a significant and positive correlation with phosphate (p <0.01), and asignificant and negative correlation with temperature (p <0.01). In Antarctic waters,eukaryotic phytoplankton abundance showed significant and negative correlationswith water depth, salinity, nitrate+nitrite, phosphate and silicate concentrations (p<0.01).
(4) The abundance of heterotrophic bacteria was the lowest in Antarctic waters, Theanalysis on vertical profile of bacterial abundance indicated that there was nosignificant decrease (p>0.05) in the IS Transect, which depth is shallow. Theirabundance in the bottom layer was lower than in the upper layers in those deeptransects, which the viral vertical distribution was similar to in Antarctic waters. Theabundance in the Western Pacific, the Indian Ocean, Southern Ocean and Antarcticwaters were2.27×10~5–3.80×10~6cells mL~(-1),2.65×10~5–1.27×10~6cells mL~(-1),8.4×10~5–1.32×10~6个mL~(-1)and1.05×10~5–3.69×10~6cells mL~(-1), respectively. The biomasseswere4.54–79.55mg m~(-3),5.31–25.4mg m~(-3),1.68–26.4mg m~(-3)and2.10–73.8mgm~(-3), respectively. In the Western Pacific, spatial distribution of heterotrophic bacteriais mainly affected by nutrients and bacteria biomass was significantly and positivelycorrelated with nitrate+nitrite and silicate (p <0.01). In the Indian Ocean, heterotrophic bacteria biomass was significantly and negetively correlated withnitrate+nitrite concentration (p <0.01). In Antarctic waters, heterotrophic bacteriaabundance showed a significant and negative correlation with water depth, salinity,phosphate and silicate (p <0.01).
(5) Viral abundance in the western Pacific, Indian Ocean, encircling Southern Oceanand Antarctic waters were2.17×10~6–2.49×10~7particles mL~(-1),3.80×10~6–2.41×10~7particles mL~(-1),1.46×10~6–2.98×10~7particles mL~(-1)and1.49×10~6–4.39×10~7particlesmL~(-1), respectively. The biomasses were0.42–4.95mg m~(-3),0.76–4.82mg m~(-3),0.11–13.3mg m~(-3)and0.25–8.78mg m~(-3), respectively. The spatial distribution ofvirus was mainly affected by the distribution of the host cells. In the western Pacific,viral distribution is affected by host cells and environmental factors. Viral biomasscorrelated positively with the Syn, Euk, heterotrophic bacteria biomass, nitrate+nitrite and silicate (p<0.01), and negatively with temperature (p<0.01). In the IndianOcean and Southern Ocean, the distribution of viruses were only affected byheterotrophic bacterial and viral biomass showed a significant and positivecorrelation with heterotrophic bacteria biomass (p <0.01). In Antarctic waters, viralabundance was significantly and positively correlated with chlorophyll a, eukabundance and bacteria abundance (p <0.01), and significantly and negativelycorrelated with water depth, salinity, phosphate and silicate were (p <0.01).
(6) According to analyze the biomass of each group of picoplankton in the westernPacific, Indian Ocean and encircling Southern Ocean (including the three partitions ofthe Pacific, Indian Ocean and the Atlantic), it was found that heterotrophic bacteriaaccounted for the vast majority of the estimated picoplankton biomass in the Pacificand Atlantic, while the contribution of Euk to total picoplankton biomass dominatedin the Indian Ocean. Our results also suggested that antagonistic and synergisticeffects existed among the groups of picoplankton. In one hand, Pro showed asignificant and negative correlation with Syn, Euk and heterotrophic bacteria (p<0.01), which suggested that pro showed the antagonistic effects with Syn and Euk.In the low-latitude waters, Pro abundance was higher than that of Syn and Eukbecause of high temperature and low nutrients. In the mid-latitude waters, it benifited for the growth of the bigger algae cells with the nutrients increased, then Euk and Synabundance increased while pro growth was limited and its abundance decreased. Onthe other hand, bacteria showed a significant and positive correlation with Syn andEuk (p <0.01), which which suggested that bacteria showed the synergistic effectswith Syn and Euk. In the mid-latitude waters, Syn and Euk utilized inorganicnutrients for the purpose of growth and thereby changed them into organic nutrients,this then leaded to an increase in the level of organic material. This increased level oforganic nutrients was then utilized to the benefit of heterotrophic bacteria, wherebacterial growth turned organic into inorganic matter. Conversely, the producedinorganic nutrients of bacterial origin could further sustain the growth of Syn andEuk.
(7) In the present study, molecular tools were used to investigate the marineeukaryotic communities of Great Wall Bay, Antarctica.18s rDNA genetic library wasconstructed, and then the library of eukaryotic ribosomal DNA were screened byrestriction fragment length polymorphism analysis. The phylogenetic diversity in thelibrary was rather great,430positive clones and38OTUs have been isolated, whichindicates that hat the microeukaryotes in this area are diverse. The38OTUs arefound to distribute in Fungi, Dinophyceae, Cercozoa Ciliates, and Uncultured marineeukaryote clones. The microeukaryotic groups are arranged from the most diverse tothe least diverse as: uncultured Dinophyceae, Fungi, Cercozoa, Ciliates and marinemicroeukaryotes. And the number of OTU of these groups is:2,16,5,2and13,respectively. And the uncultured marine microeukaryotes amounts33.4%of the totalnumbers.
引文
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