Weak Coupling between Heterotrophic Nanoflagellates and Bacteria in a Eutrophic Freshwater Environment

In a study on the dynamics and trophic role of the heterotrophic nanoflagellate (HNAN) assemblage in the microbial food web of a eutrophic oxbow lake abundances, biomass, and production rates of HNAN and their potential prey organisms, namely heterotrophic bacteria and autotrophic picoplankton, were monitored for a period of 2 years. No coupling between HNAN abundance and biomass and the abundance and biomass of their picoplanktonic prey was observed for the investigation period. The ratio of heterotrophic bacterial to HNAN abundance ranged from 2.2 x 103 to 8.6 x 103 (mean: 4.2 x 103 +/- 1.8 x 103). HNAN carbon consumption could account for only 10% to 40% of bacterial secondary production. The lack of coupling between HNAN and their potential prey and the low HNAN abundance relative to bacterial abundance suggested (a) that HNAN grazing was an insignificant factor in the regulation of bacterial abundance and (b) that HNAN abundance was regulated by predation rather than by prey abundance. This hypothesis was supported by the fact that HNAN growth rates were high (in the range of 0.45 d-1 to 1.00 d-1 during spring and summer, yearly mean: 0.52 d-1), and only weakly correlated with prey abundance and biomass. The results indicated strong top-down control of HNAN and consequently a weak coupling of HNAN and picoplankton in the investigated eutrophic freshwater environment.     

The Role of Nitrogen Fixation in Cyanobacterial Bloom Toxicity in a Temperate,Eutrophic Lake

Toxic cyanobacterial blooms threaten freshwaters worldwide but have proven difficult to predict because the mechanisms of bloom formation and toxin production are unknown, especially on weekly time scales. Water quality management continues to focus on aggregated metrics, such as chlorophyll and total nutrients, which may not be sufficient to explain complex community changes and functions such as toxin production. For example, nitrogen (N) speciation and cycling play an important role, on daily time scales, in shaping cyanobacterial communities because declining N has been shown to select for N fixers. In addition, subsequent N pulses from N₂ fixation may stimulate and sustain toxic cyanobacterial growth. Herein, we describe how rapid early summer declines in N followed by bursts of N fixation have shaped cyanobacterial communities in a eutrophic lake (Lake Mendota, Wisconsin, USA), possibly driving toxic Microcystis blooms throughout the growing season. On weekly time scales in 2010 and 2011, we monitored the cyanobacterial community in a eutrophic lake using the phycocyanin intergenic spacer (PC-IGS) region to determine population dynamics. In parallel, we measured microcystin concentrations, N₂ fixation rates, and potential environmental drivers that contribute to structuring the community. In both years, cyanobacterial community change was strongly correlated with dissolved inorganic nitrogen (DIN) concentrations, and Aphanizomenon and Microcystis alternated dominance throughout the pre-toxic, toxic, and post-toxic phases of the lake. Microcystin concentrations increased a few days after the first significant N₂ fixation rates were observed. Then, following large early summer N₂ fixation events, Microcystis increased and became most abundant. Maximum microcystin concentrations coincided with Microcystis dominance. In both years, DIN concentrations dropped again in late summer, and N₂ fixation rates and Aphanizomenon abundance increased before the lake mixed in the fall. Estimated N inputs from N₂ fixation were large enough to supplement, or even support, the toxic Microcystis blooms.     

Ocean time-series reveals recurring seasonal patterns of virioplankton dynamics in the northwestern Sargasso Sea

There are an estimated 10³⁰ virioplankton in the world oceans, the majority of which are phages (viruses that infect bacteria). Marine phages encompass enormous genetic diversity, affect biogeochemical cycling of elements, and partially control aspects of prokaryotic production and diversity. Despite their importance, there is a paucity of data describing virioplankton distributions over time and depth in oceanic systems. A decade of high-resolution time-series data collected from the upper 300 m in the northwestern Sargasso Sea revealed recurring temporal and vertical patterns of virioplankton abundance in unprecedented detail. An annual virioplankton maximum developed between 60 and 100 m during periods of summer stratification and eroded during winter convective mixing. The timing and vertical positioning of this seasonal pattern was related to variability in water column stability and the dynamics of specific picophytoplankton and heterotrophic bacterioplankton lineages. Between 60 and 100 m, virioplankton abundance was negatively correlated to the dominant heterotrophic bacterioplankton lineage SAR11, as well as the less abundant picophytoplankton, Synechococcus. In contrast, virioplankton abundance was positively correlated to the dominant picophytoplankton lineage Prochlorococcus, and the less abundant alpha-proteobacteria, Rhodobacteraceae. Seasonally, virioplankton abundances were highly synchronous with Prochlorococcus distributions and the virioplankton to Prochlorococcus ratio remained remarkably constant during periods of water column stratification. The data suggest that a significant fraction of viruses in the mid-euphotic zone of the subtropical gyres may be cyanophages and patterns in their abundance are largely determined by Prochlorococcus dynamics in response to water column stability. This high-resolution, decadal survey of virioplankton abundance provides insight into the possible controls of virioplankton dynamics in the open ocean.     

The role of N<Subscript>2</Subscript> fixation in alleviating N limitation in wetland metaphyton: enzymatic,isotopic, and elemental evidence

The role of nitrogen (N₂) fixation in balancing N supply to wetland metaphyton was assessed by comparing primary production with enzymatic, isotopic, and elemental correlates. Primary production, N₂ fixation (acetylene reduction, AR), phosphatase activity, C:N:P ratio, and N isotopic composition of metaphyton were measured along a nutrient gradient in a freshwater marsh during May through September 2004. N₂ fixation and phosphatase activity in metaphyton were negatively correlated with inorganic N and P concentrations, respectively. Although metaphyton N₂ fixation demonstrated a clear spatial pattern along the nutrient gradient, N₂ fixation rates varied monthly and decreased sharply in September. However, the percent contribution of N₂ fixation to N uptake by metaphyton consistently decreased throughout the summer. Furthermore, the decreased contribution of N₂ fixation to N uptake corresponded with an increase in metaphyton N content during the growing season. Nitrogen isotopic data suggested the sustained importance of an atmospheric N₂ source through September at the most downstream (nutrient poor) site even though the percent contribution of N₂ fixition to N uptake was lowest in that month. This suggests that metaphyton were efficiently accumulating and recycling fixed N₂ in support of primary production. Over the course of the summer, metaphyton primary production showed a weak inverse correlation with metaphyton phosphatase activity (r ² = 0.58). The largest residuals in this regression corresponded to the largest vartiation in metaphyton N content. When metaphyton primary production was normalized to metaphyton N content, production rates for the entire growing season were more strongly inversely correlated with metaphyton phosphatase activity (r ² = 0.78). Results of the study suggest that N₂ fixation in N poor areas may adequately supplement community N requirements in metaphyton, thereby causing limitation by other elemental resources such as P.     

Dynamics of prokaryotic picoplankton community in the central and southern Adriatic Sea (Croatia)

This paper addresses the dynamics of the prokaryotic picoplankton community in the coastal and open sea areas of the central Adriatic and in the coastal area of the southern Adriatic. This involved the study, from January to December 2005, of bacteria (total number of non-pigmented bacteria; high nucleic acid content (HNA) bacteria; low nucleic acid content (LNA) bacteria), cyanobacteria (Synechococcus and Prochlorococcus) and heterotrophic nanoflagellates. During the warmer seasons, in the mainly oligotrophic area under investigation into the Adriatic Sea, bacterial densities and bacterial production have shown an increase in values and domination of the LNA group of the bacterial population. In contrast, in those areas influenced by karstic rivers, the domination of HNA bacteria in total abundance of non-pigmented bacteria and high values of bacterial production was estimated throughout the investigated period. Our results show the importance of both HNA and LNA bacterial groups in the total bacterial activity throughout the investigated area. The biomass of bacteria was mostly predominant in the prokaryotic community, while within the autotrophic community Synechococcus biomass mostly predominated. During the warmer seasons, an increase in autotrophic biomass was observed in relation to non-pigmented biomass. The importance of predation in controlling bacteria by heterotrophic nanoflagellates was pronounced during the warmer period and in the coastal areas.     

Microbial Community Structure and Dynamics in the Largest Natural French Lake (Lake Bourget)

We investigated the dynamics and diversity of heterotrophic bacteria, autotrophic and heterotrophic flagellates, and ciliates from March to July 2002 in the surface waters (0–50 m) of Lake Bourget. The heterotrophic bacteria consisted mainly of “small” cocci, but filaments (>2 μm), commonly considered to be grazing-resistant forms under increased nanoflagellate grazing, were also detected. These elongated cells mainly belonged to the Cytophaga-Flavobacterium (CF) cluster, and were most abundant during spring and early summer, when mixotrophic or heterotrophic flagellates were the main bacterial predators. The CF group strongly dominated fluorescent in situ hybridization–detected cells from March to June, whereas clear changes were observed in early summer when Beta-proteobacteria and Alpha-proteobacteria increased concomitantly with maximal protist grazing pressures. The analysis of protist community structure revealed that the flagellates consisted mainly of cryptomonad forms. The dynamics of Cryptomonas sp. and Dinobryon sp. suggested the potential importance of mixotrophs as consumers of bacteria. This point was verified by an experimental approach based on fluorescent microbeads to assess the potential grazing impact of all protist taxa in the epilimnion. From the results, three distinct periods in the functioning of the epilimnetic microbial loop were identified. In early spring, mixotrophic and heterotrophic flagellates constituted the main bacterivores, and were regulated by the availability of their resources mainly during April (phase 1). Once the “clear water phase” was established, the predation pressure of metazooplankton represented a strong top-down force on all microbial compartments. During this period only mixotrophic flagellates occasionally exerted a significant bacterivory pressure (phase 2). Finally, the early summer was characterized by the highest protozoan grazing impact and by a rapid shift in the carbon pathway transfer, with a fast change-over of the main predators contribution, i.e., mixotrophic, heterotrophic flagellates and ciliates in bacterial mortality. The high abundance of ciliates during this period was consistent with the high densities of resources (heterotrophic nanoflagellates, algae, bacteria) in deep layers containing the most chlorophyll. Bacteria, as ciliates, responded clearly to increasing phytoplankton abundance, and although bacterial grazing impact could vary largely, bacterial abundance seemed to be primarily bottom-up regulated (phase 3).     

Linkages between bacterioplankton community composition, heterotrophic carbon cycling and environmental conditions in a highly dynamic coastal ecosystem

We used mesocosm experiments to study the bacterioplankton community in a highly dynamic coastal ecosystem during four contrasting periods of the seasonal cycle: winter mixing, spring phytoplankton bloom, summer stratification and autumn upwelling. A correlation approach was used in order to measure the degree of coupling between the dynamics of major bacterial groups, heterotrophic carbon cycling and environmental factors. We used catalysed reporter deposition-fluorescence in situ hybridization to follow changes in the relative abundance of the most abundant groups of bacteria (Alphaproteobacteria, Gammaproteobacteria and Bacteroidetes). Bacterial carbon flux-related variables included bacterial standing stock, bacterial production and microbial respiration. The environmental factors included both, biotic variables such as chlorophyll-a concentration, primary production, phytoplankton extracellular release, and abiotic variables such as the concentration of dissolved inorganic and organic nutrients. Rapid shifts in the dominant bacterial groups occurred associated to environmental changes and bacterial bulk functions. An alternation between Alphaproteobacteria and Bacteroidetes was observed associated to different phytoplankton growth phases. The dominance of the group Bacteroidetes was related to high bacterial biomass and production. We found a significant, non-spurious, linkage between the relative abundances of major bacterial groups and bacterial carbon cycling. Our results suggest that bacteria belonging to these major groups could actually share a function in planktonic ecosystems.     

Single-cell vs. bulk activity properties of coastal bacterioplankton over an annual cycle in a temperate ecosystem

The connections between single-cell activity properties of heterotrophic planktonic bacteria and whole community metabolism are still poorly understood. Here, we show flow cytometry single-cell analysis of membrane-intact (live), high nucleic acid (HNA) content and actively respiring (CTC+) bacteria with samples collected monthly during 2006 in northern Spain coastal waters. Bulk activity was assessed by measuring ³H-Leucine incorporation and specific growth rates. Consistently, different single-cell relative abundances were found, with 60–100% for live, 30–84% for HNA and 0.2–12% for CTC+ cells. Leucine incorporation rates (2–153 pmol L⁻¹ h⁻¹), specific growth rates (0.01–0.29 day⁻¹) and the total and relative abundances of the three single-cell groups showed marked seasonal patterns. Distinct depth distributions during summer stratification and different relations with temperature, chlorophyll and bacterial biovolume suggest the existence of different controlling factors on each single-cell property. Pooled leucine incorporation rates were similarly correlated with the abundance of all physiological groups, while specific growth rates were only substantially explained by the percentage of CTC+ cells. However, the ability to reduce CTC proved notably better than the other two single-cell properties at predicting bacterial bulk rates within seasons, suggesting a tight linkage between bacterial individual respiration and biomass production at the community level.     

Primary Production in a Subtropical Stratified Coastal Lagoon—Contribution of Anoxygenic Phototrophic Bacteria

Anaerobic anoxygenic phototrophic bacteria can be found in the suboxic waters of shallow stratified coastal systems, and may play important roles in the total primary production of subtropical stratified coastal lagoons. We investigated the spatiotemporal variability of light CO₂ fixation and net oxygen production in the stratified Conceição Lagoon (Brazil) in summer and fall of 2007, as well as the contribution of bacteriochlorophyll a (BChl a)-containing bacteria to photosynthetically driven electron transfer. Both chlorophyll a (Chl a) and BChl a varied in space, while only BChl a varied in time (three-fold increase from summer to fall). In summer, net oxygen production and light CO₂ fixation were correlated, with both having higher rates with higher Chl a concentrations in the enclosed region of the lagoon. In fall, CO₂ fixation was decoupled from oxygen production. Denaturing gradient gel electrophoresis revealed that bacterial communities of oxic site 12 and suboxic site 33 formed one cluster, different from other oxic samples within the lagoon. In addition, BChl a/Chl a ratios at these sites were high, 40% and 45%, respectively. Light acted as the main factor controlling the BChl a concentration and CO₂ fixation rates. High turbidity within the enclosed area of the lagoon explained high BChl a and decoupling between CO₂ fixation and oxygen production in oxygenated waters. Contribution of purple sulfur bacteria to total bacterial density in suboxic waters was 1.2%, and their biomass contributed to a much higher percentage (12.2%) due to their large biovolume. Our results indicate a significant contribution of anaerobic anoxygenic bacteria to the primary production of the “dead zone” of Conceição Lagoon.     

Seasonal and vertical distribution of planktonic bacteria and heterotrophic nanoflagellates in the middle Adriatic Sea

Temporal and spatial patterns of bacteria and heterotrophic nanoflagellates (HNF) were studied monthly from January 1997 to December 1998 in the middle Adriatic Sea. Bacterial and HNF relationships with phytoplankton biomass and temperature were analyzed to examine how the relative importance of bottom-up and top-down factors may shift over seasons and locations. For the coastal area, an inconsistent relationship between bacterial abundance and chlorophyll a and a stronger relationship between bacterial abundance and bacterial production suggest that other substrates than those of phytoplankton origin are important for bacteria. The analysis of simultaneous effects of temperature and bacterial production on bacterial abundance showed that the effect of temperature obscured the effects of bacterial production, suggesting that bacterial growth itself is highly temperature-dependent. The relationship between HNF abundance and bacterial abundance was slightly improved by the inclusion of in situ temperature, bacterial production or both parameters, as additional independent variables. About 60% of the variability in HNF abundance can be explained by bacterial abundance, bacterial production and temperature. In the open sea, tight coupling of bacterial abundance with chlorophyll a concentrations implied that phytoplankton-derived substrates have a dominant role in controlling bacterial abundance. During the colder months, bacterial abundance was high enough to support higher HNF abundance than observed, suggesting that predation exerted a minor depressing influence on bacterial abundance during that period. During the spring-summer period, HNF controlled bacterial standing stocks by direct cropping of bacterial production.Communicated by: H.-D. Franke     

Microbial and classical food webs: A visit to a hypertrophic lake

Abstract: Plankton community structure and fluxes of carbon for bacteria (production and bacterivory) were investigated in the urban, hypertrophic Lake Rodó (Uruguay) using a short time interval for sampling (5–15 d) during one year. The lake sustains a high phytoplankton biomass (up to 335 μg l⁻¹ chlorophyll a ) always dominated by the filamentous cyanobacteria Planktothrix agardhii . The zooplankton community was numerically dominated by rotifers and ciliates; cladocerans were rare during most of the year. The rotifer abundance was very high (up to 10⁵ individual l⁻¹), the bacterivorous Anuraeopsis fissa being the most abundant species. Predation rates of heterotrophic nanoflagellates (HNF) on bacteria (range: 31–130 bacteria HNF⁻¹ h⁻¹) were higher than those reported in the literature for field studies. A carbon budget showed that HNF can consume on average 91 and 76% of the bacterial carbon production in summer and winter, respectively. Bacterial turnover times are the lowest reported until now from field conditions (5 to 42 h). Consequently, bacterial carbon production was extremely high (72 to 1071 μg C l⁻¹ d⁻¹). Bacterial production was positively correlated to bacterial abundance but the relationship was significantly improved by the inclusion of temperature (82% variability explained). My results support the general trend for increased bacterial production with increasing trophic status, and suggest a lower energy transfer efficiency to higher trophic levels in hypertrophic lakes due to the many trophic interactions involved.     

Effects of modified nutrient concentrations and ratios on the structure and function of the native phytoplankton community in the Neuse River Estuary, North Carolina, USA

A variety of analyses were used to assess the structure (community composition) and function (assimilation number, nitrogen fixation) of phytoplankton in the Neuse River Estuary (NRE), NC under ambient and modified nutrient concentrations. Dilution bioassays were employed to reduce the concentration of nitrogen (N) or both N and phosphorus (P) and thus compare varied DIN:DIP ratios. Experimental manipulations created conditions that may result from mandated N load reductions to the estuary. We hypothesized that unilateral reduction of N loading to the NRE would increase the activity, abundance and diversity of N₂ fixing cyanobacteria. Changes in phytoplankton primary productivity, N₂ fixation (nitrogenase activity), genetic potential for N₂ fixation (presence of nifH), phytoplankton taxonomic composition (diagnostic photopigment concentration) and abundances of N₂ fixing cyanobacteria (microscopy) were determined. Decreasing ambient DIN:DIP ratios in NRE samples resulted in increased rates of N₂ fixation when seed populations were present and environmental conditions were amenable. Decreasing the DIN:DIP ratio did not lead to an increase in the abundance or diversity of N₂ fixing cyanobacteria. Because N₂ fixing cyanobacteria were only actively fixing nitrogen during periods of low riverine N discharge (summer and early autumn), lowering nutrient ratios may not have a major impact on the NRE. However, the maximum potential amount of N from N₂ fixation was calculated using rates from this study and was found to be approximately 3% of total riverine loading of N to the NRE. Because N₂ fixation occurs farther downstream and later in the year than riverine N loading to the NRE, there is potential for N₂ fixation to modify N dynamics. Analyses of the phytoplankton community as a whole in these relatively short term experiments indicated that reduced DIN:DIP may not have a major impact on their structure and function.     

Dynamics of periphyton nitrogen fixation in short-hydroperiod wetlands revealed by high-resolution seasonal sampling

Periphyton N₂ fixation plays a key role in N cycling of aquatic systems, but temporal studies of this process are often lacking, especially in systems with only seasonal flooding. We used seven samplings to characterize nitrogenase activity (acetylene reduction method) of periphyton in short-hydroperiod marl prairies and two wetlands restored from agricultural disturbance in Everglades National Park, USA. We hypothesized that the seasonal drying and rewetting would increase the temporal dynamics of the process. All sites showed significant periphyton N₂ fixation, but in restored areas highest rates were observed only in the early wet season (July), while in reference sites fixation was spread throughout the summer. Most N₂ fixation in the restored areas was confined to a 3-month-period resulting in large underestimates of annual fixation in previous studies with few seasonal measurements. N₂ fixation rates correlated with total P, N and TN:TP ratio, and periphyton moisture content in the dry season. N stable isotopic signature was a good indicator of N₂ fixation rates between sites, but did not correctly indicate seasonal patterns. These findings improve our understanding of N cycling in wetlands like the Everglades and indicate a need for more detailed measurements of processes in seasonally flooded systems.     

Distribution Pattern of Photosynthetic Picoplankton and Heterotrophic Bacteria in the Northern South China Sea

The environmental regulation of plcoplankton distribution in the northern South China Sea was examined In winter and summer of 2004. The average abundance of Synechococcus, Prochlorococcus, and heterotrophlc bacteria was lower In winter （30, 21, and 780×10^3 cells/cm^3, respectively） than In summer （53, 85, and 1 090×10^3 cells/cm^3, respectively）, but the seasonal pattern was opposite for plcoeukaryotlc phytoplankton （4 500 and 3 200 cells/cm^3 In winter and summer, respectively）. Synechococcus, picoeukaryotes, and bacteria were most abundant in the nutrient-rich coastal zone and continental shelf, but Prochlorococcus was most abundant In the continental slope and open ocean. The vertical distribution of each photosynthetic group and heterotrophlc bacteria changed between the two seasons. Synechococcus populations with apparently different phycoerythrobilin content occurred at many stations In the summer. In addition, two different populations of Prochlorococcus were found： （i） small, weakly fluorescing cells in the surface layer; and （ii） larger, strongly fluorescent cells In the deep layer. The distribution pattern of photosynthetic plcoplankton and heterotrophlc bacteria depends on environmental effects and their ecophyslologlcal differences. The distribution of Synechococcus appeared to be related to nutrient availability, whereas the distribution of Prochlorococcus appeared to be limited by temperature. Synechococcus was the only plcophytoplankton with a consistent strong relationship with bacteria.     

Nitrogenase Activity and nifH Expression in a Marine Intertidal Microbial Mat

N₂ fixation, diazotrophic community composition, and organisms actively expressing genes for N₂ fixation were examined over at 3−year period (1997–1999) for intertidal microbial mats on a sand flat located in the Rachel Carson National Estuarine Research Reserve (RCNERR) (Beaufort, NC, USA). Specifically, diel variations of N₂ fixation in the mats from the RCNERR were examined. Three distinct diel patterns of nitrogenase activity (NA) were observed. NA responses to short-term inhibitions of photosynthesis corresponded to one of the three patterns. High rates of NA were observed during peak O₂ production periods for diel experiments during summer months. Different types of NA diel variations correspond to different stages of mat development. Chloramphenicol treatments indicated that the mechanism of protein synthesis supporting NA changed throughout the day. Analysis of mat DNA and RNA gave further evidence suggesting that in addition to cyanobacteria, other functional groups were responsible for the NA observed in the RCNERR mats. The role of microbial diversity in the N₂ fixation dynamics of these mats is discussed.     

The effect of inorganic particle concentration on bacteria-virus-nanoflagellate dynamics

The effect of inorganic particle concentrations on bacteria-virus-nanoflagellate dynamics in an oligotrophic coastal system was investigated using a model aluminosilicate, kaolinite, with a modal size of 2.1 μm. Virus-only, bacteria-only and bacteria-virus-nanoflagellate incubations were carried out at increasing kaolinite concentrations to elucidate the microbial response. The sorption of bacteria and viruses to kaolinite particles was negligible over a concentration range of 1-50 mg l(-1). In contrast, the abundance of heterotrophic nanoflagellates was negatively correlated with kaolinite concentrations following both 48 and 96 h incubations. Calculated nanoflagellate bacterial ingestion rates were reduced by 5-35% depending on kaolinite particle concentration. In the bacteria-virus-nanoflagellate incubations viral production increased by 56 × 10(3) to 104 × 10(3) VLPs ml(-1) h(-1) as a function of kaolinite particle concentration. Our results demonstrate for the first time that the interaction of microbial populations with inorganic particles can shift the balance between protist and virally mediated mortality of marine heterotrophic prokaryotes.     

Bacterial Activity in a Reservoir Determined by Autoradiography and its Relationships to Phyto- and Zooplankton

In the drinking water reservoir Římov (Southern Bohemia) bacterioplankton was studied during 1983. Special attention was given to the relationships between parameters of bacterial abundance, total and individual activity. Bacterial counts and biomass was assessed and autoradiographic determinations of the proportion of active bacteria incorporating thymidine (Th) and mixture of amino acids (AA) and total uptake rate of AA were made over a year in the surface layer and during summer stratification from the thermocline and 15 m depth. Specific activity of metabolically active bacteria (SAMAB) and specific activity per unit of biomass (SAUB) were negatively correlated with counts of metabolizing cells and with bacterial biomass, respectively. Total and individual heterotrophic activity and counts of bacteria coincided with the changes of phytoplankton biomass, whereas bacteria incorporating Th were more tightly correlated with primary production. The most significant relation of metabolically active bacteria was found to cladoceran biomass. Thus, this part of heterotrophic bacterial activity seems to be stimulated by leakage of dissolved organic matter from phytoplankton being disrupted and incompletely digested by cladocerans rather than from healthy photosynthetizing cells.     

Abundance and Biomass of Heterotrophic Flagellates, and Factors Controlling Their Abundance and Distribution in Sediments of Botany Bay

The abundance and biomass of heterotrophic flagellates were estimated monthly in sediments of Botany Bay during March 1999-February 2000. The annual abundance and biomass were in the ranges of 0.46-4.70 x 10(5) cells/cm(3) and of 0.30-8.61 micro g C/cm(3), respectively. The majority of heterotrophic flagellates (93-100%) were less than 10 mm in length and few flagellates were larger than 10 mm. Of the total microbial carbon biomass, heterotrophic flagellates made up about 5% (but at times up to 35%). The contribution of heterotrophic flagellates varied from month to month, and among the sites. The abundance of heterotrophic flagellates was negatively correlated with sediment grain size and positively correlated with the abundance of bacteria, algae (autotrophic flagellates and diatoms), and their probable grazers. A best subsets regression analysis showed that bacterial and algal abundance are the most important factors controlling the abundance of heterotrophic flagellates. When the previously reported grazing rates on bacteria were applied, heterotrophic flagellates would consume a maximum of 64% of bacterial standing stock daily in Botany Bay, suggesting that heterotrophic flagellates are important as bacterivores. However, the importance of heterotrophic flagellate grazing probably varies significantly among the sites and from month to month.