Recently developed techniques for estimating bacterial biomass and productivity indicate that bacterial biomass in the sea is related to phytoplankton concentration and that bacteria utilise 10 to 50 % of carbon fixed by photosynthesis. Evidence is presented to suggest that numbers of free bacteria are controlled by nanoplankton~c heterotrophic flagellates which are ubiquitous in the marine water column. The flagellates in turn are preyed upon by microzooplankton. Heterotrophic flagellates and microzooplankton cover the same size range as the phytoplankton, thus providing the means for returning some energy from the 'microbial loop' to the conventional planktonic food chain.

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The Ecological Role of Water-Column Microbes in the Sea*

Heterotrophic bacteria are thought to be important components of aquatic ecosystems in several ways. These bacteria remineralize organic materials and convert some organic material into bacterial biomass. We examined data from 70 studies in which estimates of production of heterotrophic bacterial biomass (bacterial production) were reported for freshand saltwater ecosystems. In sediments, bacterial production was sigdicantly (p <0.001), positively correlated to sediment organic C content. Systems which had hlgh rates of benthic primary production (such as coral reefs) had rates of bacterial production greater than those predicted by sediment organic C content alone. In the photic zone of lakes and the ocean, bacterial production was significantly correlated with planktonic primary production, chlorophyll a, or numbers of planktonic bacteria. For all planktonic systems analysed, bacterial production ranged from 0.4 to 150 pg C 1-' d-' and averaged 20 % (median 16.5 %) of planktonic primary production. On an area1 basis for the entire water column, bacterial production ranged from 118 to 2439 mg m-2 d-' and averaged 30 % (median 27 %) of water column primary production. Heterotrophic bacterial production is, thus, a large component of total secondary production and is roughly twice as large as the production of macrozooplankton for a given level of primary production.

/pdf/bacterial-production-in-fresh-and-saltwater-ecosystems-a-3058k94tmt.pdf

Bacterial production in fresh and saltwater ecosystems: a cross-system overview

Despite the impressive advances that have been made in assessing the diversity of marine microorganisms, the mechanisms that underlie the participation of microorganisms in marine food webs and biogeochemical cycles are poorly understood. Here, we stress the need to examine the biochemical interactions of microorganisms with ocean systems at the nanometre to millimetre scale--a scale that is relevant to microbial activities. The local impact of microorganisms on biogeochemical cycles must then be scaled up to make useful predictions of how marine ecosystems in the whole ocean might respond to global change. This approach to microbial oceanography is not only helpful, but is in fact indispensable.

Microbial structuring of marine ecosystems

Microscopic estimation of bacterial biomass requires determination of both biovolume and biovolume-to-biomass conversion. Both steps have uncertainty when applied to the very small bacteria typically found in natural seawater. In the present study, natural bacterioplankton assemblages were freshly collected, passed through 0.6-μm-pore-size Nuclepore filters to remove larger particulate materials, and diluted for growth in 0.22-μm-pore-size Millipore filter-sterilized unenriched seawater. This provided cells comparable in size and morphology to those in natural seawater, but the cultures were free of the interfering particulate detritus naturally present. Cells were collected on glass-fiber GF/F filters, and biovolumes were corrected for cells passing these filters; C and N were measured with a CHN analyzer. Our criteria for size measurement by epifluorescence photomicrography were confirmed with fluorescent microspheres of known diameters. Surprisingly, in six cultures with average per-cell biovolumes ranging from 0.036 to 0.073 μm3, the average per-cell carbon biomass was relatively constant at 20 ± 0.08 fg of C (mean ± standard error of the mean). The biovolume-to-biomass conversion factor averaged 0.38 ± 0.05 g of C cm−3, which is about three times higher than the value previously estimated from Escherichia coli, and decreased with increasing cell volume. The C:N ratio was 3.7 ± 0.2. We conclude that natural marine bacterial biomass and production may be higher than was previously thought and that variations in bacterial size may not reflect variations in biomass per cell. Images

Relationships between Biovolume and Biomass of Naturally Derived Marine Bacterioplankton.

In his commentary Azam summarizes new work indicating that microbes in the ocean lead much different lives than we had supposed. They ingest both dissolved and particulate carbon and colonize particles of marine snow and other detritus. They can have enormous influence on the overall carbon flux in the ocean9s ecosystem.

Microbial Control of Oceanic Carbon Flux: The Plot Thickens

Frequency of dividing cells is suggested to be an indirect measure of the mean growth rate of an aquatic bacterial community. Seasonal changes in frequency of dividing cells were found which covariated with the bacterial uptake of 14C-labeled phytoplankton exudates. Batch and continuous culture growth experiments, using brackish water bacteria in pure and mixed enrichment cultures, were performed to establish a relationship between frequency of dividing cells and growth rate. An improved technique for bacterial direct counts, using fluorescent staining and epifluorescence microscopy, is presented. Based on a 6-month survey in a coastal area of the Baltic Sea, the bacterial production in the photic zone is estimated. Compared to the total primary production in the area, the bacterial population during this period utilized approximately 25% of the amount of carbon originally fixed by the primary producers.

P. Hörstedt

Papers

Frequency of Dividing Cells, a New Approach to the Determination of Bacterial Growth Rates in Aquatic Environments