Growth and succession patterns of major phylogenetic groups of marine bacteria during a mesocosm diatom bloom

Our objective was to track microbial processes associated with serial degradation of organic matter derived from algal blooms. To do this, we analyzed population fluctuations and growth responses of major phylogenetic groups of free-living marine bacteria. We used bromodeoxyuridine immunocytochemistry–fluorescence in situ hybridization methodology to examine marine bacterial community development during and after a diatom bloom in a mesocosm. We revealed that the Roseobacter/Rhodobacter, SAR11, Alteromonas, and Bacteroidetes groups were clearly major phylotypes responsible for most free-living bacterial biomass and production throughout the experiment. The clearest bacterial response was a proliferation of the Alteromonas group (cells with large volumes) during development of the bloom (up to 30?% of actively growing cells). Populations of these bacteria declined sharply thereafter, likely due to grazing. Alteromonas group responses suggest that these bacteria strongly influenced the flux of organic matter at an early bloom stage. The growth potential of Bacteroidetes was relatively large as the bloom peaked; this early development probably contributed to the initial stage of bloom decomposition. In contrast, the contribution of Roseobacter/Rhodobacter to total bacterial production increased at a late stage of decomposing of the bloom. The contributions of Betaproteobacteria, SAR11, and SAR86 groups to total bacterial abundance and production were relatively minor throughout the experiment. These results imply that the ability to utilize organic matter derived from diatoms varies among bacterial phylotypes, and, frequently, less abundant but ecological specialist taxa such as Alteromonas may play major roles in the flux of organic matter during diatom blooms.     

Dynamics of fatty acids in newly biosynthesized phytoplankton cells and seston during a spring bloom off the west coast of Hokkaido Island, Japan

¹³C uptake experiments were carried out at a station off the west coast of Hokkaido Island, Japan, during the period between late winter and spring bloom in 1997. The composition of newly biosynthesized particulate fatty acids was determined using a ¹³C tracer and gas chromatography/mass spectrometry (¹³C GC/MS), and compared with that of particulate matter (seston). Two fatty acid compositional ratios revealed that diatoms biomass (16:1(n−7)/16:0, ∑C₁₆/∑C₁₈) were strikingly high in new cells compared with seston on 17 April, coinciding with the dominance of fucoxanthin. In this study, the Polyunsaturation Index (a measure of the percentage of C₁₆ fatty acids that are polyunsaturated) in new cells is generally close to that in seston, while there was a notable discrepancy between new cells and seston in surface (10 m) waters on 17 April. This large difference between new cells and seston is ascribed to: (1) an increase of newly synthesized storage lipids (mainly 16:0 and 16:1) induced by nitrogen limitation in the top 10 m of the water column; (2) an addition of earlier synthesized phytoplankton cells (high Polyunsaturation Index at logarithmic growth stage) and non-phytoplankton components (detritus and bacteria and zooplankton) into seston. The highest Polyunsaturation Index of C₁₆ fatty acids in seston (33%) occurred at 100 m water depth. This may be caused by “settling planktonic aggregates” or “diatom resting spores”. In the present study, it is reasonable to suggest that the Polyunsaturation Index of C₁₆ fatty acids can be a useful indicator for the ecophysiological state of marine diatom populations.     

Variation of dissolved organic matter and fluorescence characteristics before,during and after phytoplankton bloom

The variation of dissolved organic matter (DOM) and fluorescence characteristics during the phytoplankton bloom were investigated in Yashima Bay, at the eastern part of the Seto Inland Sea, Japan. We found significant accumulations of dissolved organic carbon (DOC), dissolved organic nitrogen (DON), chromophoric dissolved organic matter (CDOM) fluorescence, and UV₂₆₀ during the phytoplankton bloom period in 2005, although lower accumulations of DOC and DON and only increases of CDOM fluorescence were observed during the bloom period in 2006. Little or no correlation between DOM and phytoplankton abundance might be due to the composition of DOM, which is a complex mixture of organic materials. The 3D-EEM results revealed that the DOM produced around the phytoplankton bloom period contained tyrosine, tryptophan, and humic-like substances. Our results showed that the occurrence of phytoplankton bloom contributed to the production of DOM in coastal water but the DOM accumulation depended on the type of phytoplankton bloom, the phytoplankton species in particular. From our results, we concluded that phytoplankton have a great role in the dynamics of DOM as a producer in a coastal environment.     

Development of lipids during a spring plankton bloom in the northern North Sea: I. Particulate fatty acids

The plankton spring bloom in the northern North Sea was extensively investigated during a period of three months in 1976 at a fixed station occupied by the R.V. “Meteor”. Samples of different depth-profiles, representative of the phytoplankton development, were collected eleven times to analyze the concentration of fatty acids of the particulate matter. The water column was divided into an upper and lower layer according to the thermocline depths, because different processes take place in these layers. During the exponential growth phase the fatty acid concentration rose only slightly due to increases in polyunsaturated fatty acids (18:4, 20:5, 22:6), which are typical for marine plankton. With the exhaustion of nutrients the biochemical composition changed and the fatty acid concentration increased sharply from about 3 to 20 μmol C dm^? finally to about 30% of the particulate carbon. The main proportion consisted of oleic acid (28.3%) and palmitic acid (24.2%). The first phytoplankton bloom, dominated by diatoms (Chaetoceros species), was characterized by the increase in fatty acids with 16 carbon atoms, whereas during the second smaller bloom, with dinoflagellates as the main species, more fatty acids with 18 carbon atoms occurred. After the stationary growth phase the phytoplankton biomass strongly decreased, resulting in an increase of particulate matter below the thermocline. The fatty acid pattern there was similar to that during the stationary phase of the phytoplankton bloom in the upper layer.     

大型水母沙海蜇暴发的温带贫营养海洋生态系统中细菌和浮游植物的关系

Bacterial abundance, phytoplankton community structure and environmental parameters were investigated to study the relationships between bacteria and phytoplankton during giant jellyfish Nemopilema nomurai blooms in the central Yellow Sea during 2013. N. nomurai appeared in June, increased in August, reached a peak and began to degrade in September 2013. Results showed that phosphate was possible a key nutrient for both phytoplankton and bacteria in June, but it changed to nitrate in August and September. Phytoplankton composition significantly changed that pico-phytoplankton relative biomass significantly increased, whereas other size phytoplankton significantly decreased during jellyfish bloom. In June, a significantly positive correlation was observed between chlorophyll a concentration and bacterial abundance(r=0.67, P0.001, n=34).During jellyfish outbreak in August, there was no significant correlation between phytoplankton and bacteria(r=0.11, P0.05, n=25), but the relationship(r=0.71, P0.001, n=31) was rebuilt with jellyfish degradation in September. In August, small size phytoplankton occupied the mixed layer in offshore stations, while bacteria almost distributed evenly in vertical. Chlorophyll a concentration significantly increased from(0.42±0.056) μg/L in June to(0.74±0.174) μg/L in August, while bacterial abundance just slightly increased. Additionally, the negative net community production indicated that community respiration was not entirely determined by the local primary productivity in August. These results indicated that jellyfish blooms potentially affect coupling of phytoplankton and bacteria in marine ecosystems.     

The spatial and temporal development of the spring phytoplankton bloom in the Celtic Sea,April 1979

The results are presented from three hydrogrpahic surveys in April 1979 of a 40 × 50 km region of the Celtic Sea, centred at 7°W and 51°N, using a towed undulating sensor system. In the $\text{5}\text{1}\text{2}$ days between Surveys 1 and 2, the seasonal thermocline was established, with surface to bottom temperature differences reaching 1.5°C, the average surface chlorophyll a level increased from ～ 1 to ～ 5.5 mg m^?3 due mainly to the growth of diatoms, and the surface nitrate concentration decreased from 6 to 1 μM. The third survey was carried out after a further two days and, although surface properties changed little, there was a general deepening of the mixed layer due to stronger winds, and a further increase in the standing stock of phytoplankton.By applying appropriate techniques of horizontal spatial averaging, which took into account possible advective effects, a quantitative comparison was made of the changes at two positions in the survey area which showed significant differences in the rate of development of the phytoplankton population. A simple model of phytoplankton growth was then developed, based on calculations of eddy diffusivity, on measurements of rates of photosynthetic carbon assimilation, and on observations of carbon to chlorophyll ratios, subsurface light attenuation and inorganic nutrient levels. In the absence of any data on zooplankton populations, the loss of phytoplankton by grazing was left as a free parameter.The model was only partly successful in reproducing the observed changes in chlorophyll concentrations during the first $\text{5}\text{1}\text{2}$ days and showed serious limitations for the subsequent 2 days. However, it emphasised several features of the dynamics of spring phytoplankton populations which require further experimental or observational investigation. These include more precise measurements of carbon to chlorophyll ratios and grazing pressure to which the model is sensitive over a rather narrow range (this has important implications in terms both of control by grazing and nutrient limitation), the potential significance of physiological photoadaptation by the plant cells in determining their vertical distribution, and the role of eddy diffusion across the developing thermocline in relation to the sinking of phytoplankton cells and the upward mixing of inorganic nutrients.     

Development of lipids during a spring plankton bloom in the northern North Sea: II. Dissolved lipids and fatty acids

During the spring plankton bloom (1976) in the northern North Sea the fatty acid and lipid distribution of the chloroform extractable fraction of seawater was analyzed by gas chromatography and thin-layer chromatography. The fatty acids in the seawater made up about 3% of the total dissolved organic matter. Maximum concentrations during the first phytoplankton bloom were in the range of 5 μmol C dm^?3. Palmitic acid (30.4%) and oleic acid (21.4%) as well as myristic, stearic and palmitoleic acids were the main fatty acids. The concentrations of fatty acids were higher below the thermocline in the deeper layers of the water column. The oleic acid showed large fluctuations, especially below the thermocline and seemed to be bound in a polar fraction. Different lipid classes containing fatty acids were determined by thin-layer chromatography. An estimation showed that the free fatty acid fraction made up the main portion (47.9%), followed by the triacylglycerol fraction (22.8%) and a more polar fraction with 25.1%. The triacylglycerol fraction appears to be more stable than the other fraction in the water column.     

Role of dissolved silicate in the occurrence of a phytoplankton bloom

The spring bloom of phytoplankton was studied in March in Funka Bay, Japan, to test the Tsunogai (1979)'s hypothesis regarding the role of silicate in the bloom. The hypothesis comprises two parts. 1) Diatoms are predominant when all the physical and chemical conditions are adequate for plankton growth. 2) Since the Si:P ratio of the diatom body is usually much larger than that of sea water, flagellates (non-siliceous phytoplankton) replace diatoms after dissolved silicate in the sea water has been almost completely consumed by diatoms. At the end of the bloom in late March phosphate still remained in the water but silicate was exhausted and the main species of phytoplankton changed from diatoms to flagellates. Grazing pressure by zooplankton at this time was not so great. A model using the data on assimilation rates of silicate showed a dramatic change of silicate uptake in late March. Poison in scallops caused byProtogonyaulux sp. (dinoflagellates) rapidly increased from mid-April at all stations along the coast of Funka Bay. All of these findings support Tsunogai's hypothesis.     

必需脂肪酸在海洋食物链中的作用

"硅藻-桡足类-鱼类"是海洋生态系统中物质循环和能量流动的主要环节[1].在海洋食物链中,饵料的数量和营养价值是各营养级海洋动物生长和繁殖的主要制约因素[2].影响饵料营养价值的化学组分包括蛋白质、脂类、糖类、矿物质、维生素等,其中脂类物质作为浮游植物营养成分的重要组成部分,在动物的细胞膜、视网膜、脑、神经组织和某些激素的合成中具有不可替代的作用[3].     

Arsenic,antimony and selenium speciation during a spring phytoplankton bloom in a closed experimental ecosystem

A study has been made of the effects of a spring diatom bloom on the levels and speciation of dissolved arsenic, antimony and selenium in the water enclosed in an experimental ecosystem moored in Loch Ewe (NW Scotland). Primary productivity resulted in severe depletion of phosphate and silicate in the bag, but had little effect on the levels and speciation of arsenic and antimony. Calculations based on phosphate depletion data strongly suggest that the field diatom population present during the experiment was capable of some degree of discrimination between phosphate and arsenate ions. Whilst biomethylation of arsenic was not observed in the upper region of the bag, where the phytoplankton population was at its greatest, the methylated form accounted for 64% of the dissolved arsenic at the base of the bag. In this region, however, the total dissolved arsenic levels were not higher than in the rest of the bag, suggesting microbial methylation of dissolved arsenic rather than the release of methylated arsenic from decaying phytoplankton.Total dissolved selenium and selenium(IV) showed some evidence of depletion during the development of the phytoplankton bloom, in support of previous observations of preferential selenite assimilation.     

Factors controlling the temporal and spatial variations in Synechococcus abundance in a monsoonal estuary

Temporal and spatial variations in Synechococcus abundance were investigated over an annual cycle (February'10–January'11) along a salinity gradient (0–35) in the tropical Zuari estuary, influenced by south-west monsoons. Synechococcus exhibited salinity preferences with phycoerythrin-rich cells at salinities >2 (Synechococcus-PEI), >20 (Synechococcus-PEII) and <1 (Synechococcus-PEIII) whereas phycocyanin-rich (Synechococcus-PC) dominant at lower salinities. Downstream stratification during monsoon caused Synechococcus group segregation in the surface and near-bottom waters. During monsoon-break and non-monsoon period stabilized waters, increased salinity, temperature, solar radiation and low rainfall favored high Synechococcus abundance whereas unstable waters, increased turbidity and low solar radiation during active monsoon lowered abundance. SYN-PC positively co-related with nitrate and phosphate and SYN-PEI with phosphate. Synechococcus contribution to phytoplankton carbon biomass ranged from 9 to 29%. In monsoonal estuaries, rainfall intensity regulates freshwater runoff which modulates the estuarine environment, creating temporal–spatial niche segregation of Synechococcus groups thereby serving as indicator organisms of the estuarine hydrodynamics.     

Enrichment of alkanes within a phytoplankton bloom during an in situ iron enrichment experiment in the western subarctic Pacific

During the Subarctic Pacific Iron Experiment for Ecosystem Dynamics Study ΙΙ (SEEDS-II), we monitored variations in the concentrations of non-methane hydrocarbons (NMHCs), CH₃Cl, N₂O, and CH₄ within a phytoplankton bloom. Stable isotopic compositions were also determined to evaluate the sources of the variations. Although there was little variation in either the concentrations or the stable isotopic compositions of alkenes, CH₃Cl, N₂O, and CH₄ during the 23-day observation period, alkane concentrations increased substantially as the phytoplankton bloomed. The column-integrated quantities of alkanes increased to 3 times pre-bloom levels for C₂H₆, 5 times for C₃H₈, and 20 times for n-C₄H₁₀. The δ¹³C values of both C₂H₆ and C₃H₈ remained almost constant while concentrations increased, whereas that of n-C₄H₁₀ increased by about 12‰. To evaluate the sources of the alkanes produced during the bloom, we compared their δ¹³C values with those of alkanes produced in axenic phytoplankton cultures in our laboratory. We concluded that during the SEEDS-ΙΙ experiment the major portions of C₂H₆ and C₃H₈ were produced during the autolysis of diatoms cells, whereas n-C₄H₁₀ was produced during autolysis of other phytoplankton cells such as cryptophytes and dinoflagellates.     

Changes in structural and functional diversity of nematode communities during a spring phytoplankton bloom in the southern North Sea

The response of nematode communities to the sedimentation of a spring phytoplankton bloom in a sandy, well-oxygenated sediment at a single station (station 330) in the Southern North Sea was investigated monthly from early March to July 1999. Both structural (nematode density, diversity, vertical distribution and community composition) and functional (feeding type distributions, number of species within feeding groups) characteristics showed considerable changes shortly after the arrival of fresh organic material at the sediment surface. The general increase in numerical densities and diversity was related to changes within the groups of selective deposit-feeding and epistrate-feeding nematodes. It is hypothesised that sedimentation and subsequent remineralisation of fresh organic matter during the spring phytoplankton bloom result in an increase of suitable food items (both living and dead). This, combined with the availability of oxygen, creates conditions in which many nematode species can co-exist.     

Contrasting complexing capacity of dissolved organic matter produced during the onset,development and decay of a simulated bloom of the marine diatom Skeletonema costatum

The capacity of natural dissolved organic matter (DOM) produced during the onset, development and decay of a simulated bloom of the marine diatom Skeletonema costatum to complex free copper has been followed for a 2 week period. Copper binding capacity of the culture was measured by anodic stripping voltammetry (ASV) with a hanging mercury drop electrode (HMDE). The concentration of dissolved organic carbon (DOC) and two fluorophores, M (humic-like, Ex/Em: 320 nm/410 nm) and T (protein-like, Ex/Em: 280 nm/350 nm), were followed during the course of the incubation. Models using DOC concentrations alone could not accurately predict the complexing capacity of the culture, especially at the end of the bloom, and better predictions were obtained when fluorescence measurements were considered. They were helpful in characterising two types of copper ligands produced in the culture. The first type, traced by the fluorescence of peak T, was related to labile DOC directly exuded by phytoplankton. The second type, traced by the fluorescence of peak M, was the refractory humic-like material presumably produced in situ as a by-product of the bacterial degradation of phytogenic materials. During the onset and development of the bloom (days 0 to 7), the fluorescence of peak T explains 60–80% of the total complexing capacity of the culture, suggesting that exuded “protein-like” compounds among other exuded complexing agents efficiently complexed free copper. On the contrary, during the decay (days 8 to 13), these ligands were replaced by humic substances as the complexing agent for copper.     

Factors controlling the timing of major spring bloom events in an UK south coast estuary

Factors controlling the timing of major (>10 mg chlorophyll a m⁻³) spring bloom events in the estuarine waters of the Solent, on the south coast of the UK, have been investigated. Winter to summer variations in chlorophyll a concentration together with relevant meteorological and hydrographical data have been analysed for 5 years (1988, 1992, 2001, 2002 and 2003). Mean water column irradiance is demonstrated to be the main factor controlling the timing of the first major spring bloom event, usually dominated by large chain-forming diatoms. When chlorophyll a concentration first exceeds 10 mg m⁻³ in spring (usually in May) the mean water column photosynthetic active radiation (PAR) averaged for one week prior to the sampling date was always >380 W h m⁻² d⁻¹. Prior to the main spring bloom event surface incident radiation and water turbidity combine to limit chlorophyll a concentration to levels <10 mg m⁻³. Chlorophyll a concentrations >10 mg m⁻³ do not occur in the Solent until almost the entire 10 m water column is within the euphotic zone (i.e. above 1% light level) and light extinction coefficient (k) is ca. ≤0.5 m⁻¹. Statistically, river flow explains the largest percentage of the variations in k and the delayed bloom in June 2002 is due to increased cloud cover and high levels of rainfall in May, which caused a reduction in surface incident irradiance and increased turbidity. Chlorophyll a peaks during these major bloom events generally occur on spring tides when increased mixing rates favour net growth of diatoms.     

Mechanisms controlling the temporal variation of the sea ice edge in the Southern Ocean

This paper examines the mechanism controlling the short time-scale variation of sea ice cover over the Southern Ocean. Sea ice concentration and ice velocity datasets derived from images of the Defense Meteorological Satellite Program (DMSP) Special Sensor Microwave Imager (SSM/I) are employed to reveal this mechanism. The contribution of both dynamic and thermodynamic processes to the change in ice edge location is examined by comparing the meridional velocity of ice edge displacement and sea ice drift. In the winter expansion phase, the thermodynamic process of new ice production off the ice edge plays an important role in daily advances of ice cover, whereas daily retreats are mostly due to southward ice drift. On the other hand, both advance and retreat of ice edges in the spring contraction phase are mostly caused by the dynamic process of the ice drift. Based on the above mechanism and the linear relation between the degree of ice production at the ice edge and northward wind speed, the seasonal advance of ice cover can be roughly reproduced using the meridional velocity of ice drift at the ice edge.     

巴夫藻脂肪酸和奋醇的同步测定研究

建立了一种脂肪酸甾醇分步衍生气相色谱／质谱(GC／MS)同步分析新方法，使用一份干质量约20mg的巴夫藻(Pavlova virdis)样品，采用SPB-5色谱柱，在岛津QP2010气相色谱／质谱联用分析仪上，实现可靠的定量定性分析。方法回收率为93％～107％；方法的重现性较好，各组分3次测定结果的标准相对偏差都小于10％；方法的检测限为3．5ng。选取巴夫藻(Pavlova sp．)进行分析，可初步确定出26种脂肪酸和10种甾类化合物以及它们的含量。     

Spring phytoplankton bloom in the fronts of the East China Sea

Frontal areas between warm and saline waters of the Kuroshio currents and colder and diluted waters of the East China Sea (ECS) influenced by the Changjiang River were identified from the satellite thermal imagery and hydrological data obtained from the Coastal Ocean Process Experiment (COPEX) cruise during the period between March 1^st and 10^th, 1997. High chlorophyll concentrations appeared in the fronts of the East China Seas with the highest chlorophyll-a concentration in the southwestern area of Jeju Island (~2.9 mg/m³) and the eastern area of the Changjiang River Mouth (~2.8 mg/m³). Vertical structures of temperature, salinity and density were similar, showing the fronts between ECS and Kuroshio waters. The water column was well mixed in the shelf waters and was stratified around the fronts. It is inferred that the optimal condition for light utilization and nutrients induced both from the coastal and deep waters enhances the high phytoplankton productivity in the fronts of the ECS. In addition, the high chlorophyll-a in the fronts seems to have been associated with the water column stability as well.     

筼筜湖绿潮期间颗粒有机物及沉积有机物的来源研究

以原子碳氮比C_at/N_at、TOC/Chl a、δ13C和δ15N等为指标,分析了筼筜湖绿潮爆发期间悬浮颗粒有机物(POM)和沉积有机物(SOM)的来源。结果显示,筼筜湖的POM主要以外源输入为主。在靠近海水入口的引水渠,POM主要来自厦门西海域的陆源有机碎屑;位于筼筜湖上游的干渠,生活污水及餐饮业废水的有机质是其POM的主要贡献者;内、外湖POM的C_at/N_at(6.94~7.08)与浮游植物接近,但它们并不以浮游植物为主,而主要来自引水渠和干渠有机质的输入。以内湖为例,它们对内湖POM的联合贡献高达54%~97%。筼筜湖SOM的潜在来源多样,但不同湖区差异显著:在大型海藻覆盖区,主要以大型海藻和POM为主,而在无海藻覆盖的区域,则主要来自POM的自然沉降或与底栖微藻的联合贡献。结果表明,在来源复杂的潟湖系统,有机物的化学组成并不能很好的指示有机物的来源和成因,它在有机质的示踪方面并不如稳定同位素来得有效可靠。不过,基于多种指标的分析结果可能更准确。     

河北昌黎生态监控区海洋浮游植物多样性变化趋势研究

根据2006-2010年,每年8月河北昌黎生态监控区采集到的浮游植物种类和细胞数量等调查数据,以Simpson优势度为指标分析其物种组成及群落结构的年际变化情况。运用3种多样性指标,即Pielou均匀度指数(J)、Margalef丰富度指数(dM)a和Shannon-Wiener物种多样性指数(H)′分别进行站位分析及均值分析,深入探究浮游植物多样性的年际变化趋势及规律。利用水质综合污染指数(P)对海水环境污染程度进行指标评价。结果显示:5年中浮游植物大都隶属于硅藻和甲藻类,且前者占绝对优势,群落结构较单一,物种数目逐年减少;站位分析中,除个别站位外3种指标的变化趋势均呈逐年波动性减少;均值分析中,J值均处于中等水平,dMa值为波动性下降,H′值则呈逐年降低之趋势,表明浮游植物多样性总体水平逐年下降。同时,P值呈逐年波动性上升趋势,前4年为中度污染,第5年为严重污染。因此,监控区海域生态环境状况每况愈下。针对上述结果,探讨了浮游植物多样性变化的主要原因。对全面掌握海洋浮游植物物种演变,改善海域生态环境现状,实现海洋资源可持续利用和海洋生态系统可持续发展,具有重要的现实意义。