Biofilms are complex communities of microorganisms attached to surfaces or associated with interfaces. Despite the focus of modern microbiology research on pure culture, planktonic (free-swimming) bacteria, it is now widely recognized that most bacteria found in natural, clinical, and industrial settings persist in association with surfaces. Furthermore, these microbial communities are often composed of multiple species that interact with each other and their environment. The determination of biofilm architecture, particularly the spatial arrangement of microcolonies (clusters of cells) relative to one another, has profound implications for the function of these complex communities. Numerous new experimental approaches and methodologies have been developed in order to explore metabolic interactions, phylogenetic groupings, and competition among members of the biofilm. To complement this broad view of biofilm ecology, individual organisms have been studied using molecular genetics in order to identify the genes required for biofilm development and to dissect the regulatory pathways that control the plankton-to-biofilm transition. These molecular genetic studies have led to the emergence of the concept of biofilm formation as a novel system for the study of bacterial development. The recent explosion in the field of biofilm research has led to exciting progress in the development of new technologies for studying these communities, advanced our understanding of the ecological significance of surface-attached bacteria, and provided new insights into the molecular genetic basis of biofilm development.

Microbial Biofilms: from Ecology to Molecular Genetics

If stretched end to end, the estimated 1030viruses in the oceans would span farther than the nearest 60 galaxies. This reservoir of genetic and biological diversity continues to yield exciting discoveries and, in this Review, Curtis A. Suttle highlights the areas that are likely to be of greatest interest in the next few years. Viruses are by far the most abundant 'lifeforms' in the oceans and are the reservoir of most of the genetic diversity in the sea. The estimated 1030 viruses in the ocean, if stretched end to end, would span farther than the nearest 60 galaxies. Every second, approximately 1023 viral infections occur in the ocean. These infections are a major source of mortality, and cause disease in a range of organisms, from shrimp to whales. As a result, viruses influence the composition of marine communities and are a major force behind biogeochemical cycles. Each infection has the potential to introduce new genetic information into an organism or progeny virus, thereby driving the evolution of both host and viral assemblages. Probing this vast reservoir of genetic and biological diversity continues to yield exciting discoveries.

Marine viruses — major players in the global ecosystem

Viruses are the most common biological agents in the sea, typically numbering ten billion per litre. They probably infect all organisms, can undergo rapid decay and replenishment, and influence many biogeochemical and ecological processes, including nutrient cycling, system respiration, particle size-distributions and sinking rates, bacterial and algal biodiversity and species distributions, algal bloom control, dimethyl sulphide formation and genetic transfer. Newly developed fluorescence and molecular techniques leave the field poised to make significant advances towards evaluating and quantifying such effects.

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Marine viruses and their biogeochemical and ecological effects

The discovery that viruses may be the most abundant organisms in natural waters, surpassing the number of bacteria by an order of magnitude, has inspired a resurgence of interest in viruses in the aquatic environment. Surprisingly little was known of the interaction of viruses and their hosts in nature. In the decade since the reports of extraordinarily large virus populations were published, enumeration of viruses in aquatic environments has demonstrated that the virioplankton are dynamic components of the plankton, changing dramatically in number with geographical location and season. The evidence to date suggests that virioplankton communities are composed principally of bacteriophages and, to a lesser extent, eukaryotic algal viruses. The influence of viral infection and lysis on bacterial and phytoplankton host communities was measurable after new methods were developed and prior knowledge of bacteriophage biology was incorporated into concepts of parasite and host community interactions. The new methods have yielded data showing that viral infection can have a significant impact on bacteria and unicellular algae populations and supporting the hypothesis that viruses play a significant role in microbial food webs. Besides predation limiting bacteria and phytoplankton populations, the specific nature of virus-host interaction raises the intriguing possibility that viral infection influences the structure and diversity of aquatic microbial communities. Novel applications of molecular genetic techniques have provided good evidence that viral infection can significantly influence the composition and diversity of aquatic microbial communities.

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Virioplankton: viruses in aquatic ecosystems

Viruses exist wherever life is found. They are a major cause of mortality, a driver of global geochemical cycles and a reservoir of the greatest genetic diversity on Earth. In the oceans, viruses probably infect all living things, from bacteria to whales. They affect the form of available nutrients and the termination of algal blooms. Viruses can move between marine and terrestrial reservoirs, raising the spectre of emerging pathogens. Our understanding of the effect of viruses on global systems and processes continues to unfold, overthrowing the idea that viruses and virus-mediated processes are sidebars to global processes.

Viruses in the sea

The concentration of bacteriophages in natural unpolluted waters is in general believed to be low, and they have therefore been considered ecologically unimportant. Using a new method for quantitative enumeration, we have found up to 2.5 x 10(8) virus particles per millilitre in natural waters. These concentrations indicate that virus infection may be an important factor in the ecological control of planktonic micro-organisms, and that viruses might mediate genetic exchange among bacteria in natural aquatic environments.

High abundance of viruses found in aquatic environments.

The content of carbon, nitrogen, oxygen, phosphorus and sulfur was measured in individual cells from 6 native aquatic samples and 4 samples of cultured bacteria by X-ray microanalysis using a transmission electron microscope (TEM). The molar C N:P ratio for the pooled sample was 50:lO:l From length and width measurements of unfixed air-dried cells we estimated cell volumes over a total range of 0.0026 to 15.8 pm3. and mean C:volume ratios of 30 to 162 fg for the samples included. For the marine samples we found mean N:C ratios of 0.25 to 0.28, while cells from fresh or brackish waters had mean N:C ratios of 0.17 to 0.20, indicating differences in nutrient availability. The P:C ratios for the samples analyzed varied from 0.040 to 0.090, with a pooled mean of 0.052, which is approximately twice that of the Redfield ratio for PC. For 0:C ratios we estimated a pooled mean of 0.37 and a range of 0.22 to 0.77 for all samples. We may conclude that slow-growing or non-growing cells have low 0 C ratios. The mean S:C ratio for all samples was 0.031, with a range of 0 016 to 0.084 for the sample means. A general conclus~on is that single-cell analyses of elemental composition give important information on the physiological conditions of cells and on possible nutrient limitations. The rationale for this is the assumption that changes in macromolecular composition are due to nutrient availability.

https://www.int-res.com/articles/ame/10/a010p015.pdf

Content of carbon, nitrogen, oxygen, sulfur and phosphorus in native aquatic and cultured bacteria

The possible roles of viruses in phytoplankton dynamics were investigated in seawater mesocosms with natural assemblages of phytoplankton growing under various nutrient regimes. Blooms of the marine coccolithophorid Emiliania huxleyi (Lohmann) Hay & Mohler were in some cases succeeded by an increase in the abundance of a morphologically homogeneous population of viruses. The viruses had an hexagonal outline and were about 180 nm in diameter. Similar viruses were found both intracellularly and associated with apparently lysed cells. Viral lysis could account for 25 to 100 % of the net mortality of E huxleyi when the blooms declined under non-limiting nutrient conditions or when the nitrate concentration was low. Production of viruses was limited when the phosphate concentration was low. Dur~ng a bloom of E. huxleyi in Norwegian coastal waters in 1990 we found that the concentration of free algal viruses was increasing at the end of the bloom, indicating that viruses of E. huxleyi may be important under natural conditions as well. These results suggest that viral mortality of phytoplankton may be an important factor regulating community structure, diversity and biomass production in marine environments.

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Viral mortality of the marine alga Emiliania huxleyi (Haptophyceae) and termination of algal blooms

Population sizes of algae, bacteria, heterotrophic flagellates, and viruses were observed through the 1989 spring diatom bloom in Raunefjorden in western Norway. The culmination of the diatom bloom was followed by a peak in the concentration of bacteria and an increase in the concentration of heterotrophic flagellates, a pattern consistent with the concept of a food chain from photosynthetically produced organic material, through bacteria, to bacterivorous flagellates. The concentration of viruses varied through the spring bloom from 5 x 10 in the prebloom situation to a maximum of 1.3 x 10 viruses ml 1 week after the peak of the diatom bloom. Coinciding with the collapse in the diatom bloom, a succession of bacteria and viruses was observed in the mucous layer surrounding dead or senescent diatoms, with an estimated maximum of 23% of the total virus population attached to the diatoms. The dynamic behavior observed for the virus population rules out the possibility that it is dominated by inactive species, and the viruses are suggested to be active members of the microbial food web as agents causing lysis in parts of the bacterial population, diverting part of the bacterial production from the predatory food chain.

Viruses as partners in spring bloom microbial trophodynamics.

Bacteria and virus particles were harvested from water samples by ultracentrifugation directly onto Formvar-coated electron microscopy grids and counted in a transmission electron microscope. With this technique, we have counted and sized bacteria and viruses in marine water samples and during laboratory incubations. By X-ray microanalysis, we could determine the elemental composition and dry-matter content of individual bacteria. The dry weight/volume ratio for the bacteria was 600 fg of dry weight microns-3. The potassium content of the bacteria was normal compared with previous estimates from other bacterial assemblages; thus, this harvesting procedure did not disrupt the bacterial cells. Virus particles were, by an order of magnitude, more abundant than bacteria in marine coastal waters. During the first 5 to 7 days of incubation, the total number of viruses increased exponentially at a rate of 0.4 day-1 and thereafter declined. The high proliferation rate suggests that viral parasitism may affect mortality of bacteria in aquatic environments.

Mikal Heldal

Papers

High abundance of viruses found in aquatic environments.

Content of carbon, nitrogen, oxygen, sulfur and phosphorus in native aquatic and cultured bacteria

Viral mortality of the marine alga Emiliania huxleyi (Haptophyceae) and termination of algal blooms

Viruses as partners in spring bloom microbial trophodynamics.

Enumeration and biomass estimation of planktonic bacteria and viruses by transmission electron microscopy.