In food webs, interactions between competition and defence control the partitioning of limiting resources. As a result, simple models of these interactions contain links between biogeochemistry, diversity, food web structure and ecosystem function. Working at hierarchical levels, these mechanisms also produce self-similarity and therefore suggest how complexity can be generated from repeated application of simple underlying principles. Reviewing theoretical and experimental literature relevant to the marine photic zone, we argue that there is a wide spectrum of phenomena, including single cell activity of prokaryotes, microbial biodiversity at different levels of resolution, ecosystem functioning, regional biogeochemical features and evolution at different timescales; that all can be understood as variations over a common principle, summarised in what has been termed the 'Killing-the-Winner' (KtW) motif. Considering food webs as assemblages of such motifs may thus allow for a more integrated approach to aquatic microbial ecology.

Microalgae could provide a sustainable alternative to fish oil as a source for the omega-3 polyunsaturated fatty acids eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA). However, growing microalgae on a large-scale is still more cost-intensive than fish oil production, and outdoor productivities vary greatly with reactor type, geographic location, climate conditions and microalgae species or even strains. The diatom Phaeodactylum tricornutum has been intensively investigated for its potential in large-scale production, due to its robustness and comparatively high growth rates and EPA content. Yet, most research have been performed in southern countries and with a single commercial P. tricornutum strain, while information about productivities at higher latitudes and of local strains is scarce. We examined the potential of the climate conditions in Bergen, western Norway for outdoor cultivation of P. tricornutum in flat panel photobioreactors and cultivated three different strains simultaneously, one commercial strain from Spain (Fito) and two local isolates (M28 and B58), to assess and compare their biomass and EPA productivities, and fatty acid (FA) profiles. The three strains possessed similar biomass productivities (average volumetric productivities of 0.20, 0.18, and 0.21 g L− 1 d− 1), that were lower compared to productivities reported from southern latitudes. However, EPA productivities differed between the strains (average volumetric productivities of 9.8, 5.7 and 6.9 mg L− 1 d− 1), due to differing EPA contents (average of 4.4, 3.2 and 3.1% of dry weight), and were comparable to results from Italy. The EPA content of strain Fito of 4.4% is higher than earlier reported for P. tricornutum (2.6–3.1%) and was only apparent under outdoor conditions. A principal component analysis (PCA) of the relative FA composition revealed strain-specific profiles. However, including data from laboratory experiments, revealed more significant differences between outdoor and laboratory-grown cultures than between the strains, and higher EPA contents in outdoor grown cultures.

Factors controlling the community composition of marine heterotrophic prokaryotes include organic-C, mineral nutrients, predation, and viral lysis. Two mesocosm experiments, performed at an Arctic location and bottom-up manipulated with organic-C, had very different results in community composition for both prokaryotes and viruses. Previously, we showed how a simple mathematical model could reproduce food web level dynamics observed in these mesocosms, demonstrating strong top-down control through the predator chain from copepods via ciliates and heterotrophic nanoflagellates. Here, we use a steady-state analysis to connect ciliate biomass to bacterial carbon demand. This gives a coupling of top-down and bottom-up factors whereby low initial densities of ciliates are associated with mineral nutrient-limited heterotrophic prokaryotes that do not respond to external supply of labile organic-C. In contrast, high initial densities of ciliates give carbon-limited growth and high responsiveness to organic-C. The differences observed in ciliate abundance, and in prokaryote abundance and community composition in the two experiments were in accordance with these predictions. Responsiveness in the viral community followed a pattern similar to that of prokaryotes. Our study provides a unique link between the structure of the predator chain in the microbial food web and viral abundance and diversity.

Theoretical work has suggested an important role of lytic viruses in controlling the diversity of their prokaryotic hosts. Yet, providing strong experimental or observational support (or refutation) for this has proven evasive. Such models have usually assumed “host groups” to correspond to the “species” level, typically delimited by 16S rRNA gene sequence data. Recent model developments take into account the resolution of species into strains with differences in their susceptibility to viral attack. With strains as the host groups, the models will have explicit viral control of abundance at strain level, combined with explicit predator or resource control at community level, but the direct viral control at species level then disappears. Abundance of a species therefore emerges as the combination of how many strains, and at what abundance, this species can establish in competition with other species from a seeding community. We here discuss how species diversification and strain diversification may introduce competitors and defenders, respectively, and that the balance between the two may be a factor in the control of species diversity in mature natural communities. These models can also give a dominance of individuals from strains with high cost of resistance; suggesting that the high proportion of “dormant“ cells among pelagic heterotrophic prokaryotes may reflect their need for expensive defense rather than the lack of suitable growth substrates in their environment.

Combining a minimum food web model with Arctic microbial community dynamics, we have suggested that top-down control by copepods can affect the food web down to bacterial consumption of organic carbon. Pursuing this hypothesis further, we used the minimum model to design and analyse a mesocosm experiment, studying the effect of high (+Z) and low (-Z) copepod density on resource allocation, along an organic-C addition gradient. In the Arctic, both effects are plausible due to changes in advection patterns (affecting copepods) and meltwater inputs (affecting carbon). The model predicts a trophic cascade from copepods via ciliates to flagellates, which was confirmed experimentally. Auto- and heterotrophic flagellates affect bacterial growth rate and abundance via competition for mineral nutrients and predation, respectively. In +Z, the model predicts low bacterial abundance and activity, and little response to glucose; as opposed to clear glucose consumption effects in –Z. We observed a more resilient bacterial response to high copepods and demonstrate this was due to changes in bacterial community equitability. Species able to use glucose to improve their competitive and/or defensive properties, became predominant. The observed shift from a SAR11-to a Psychromonodaceae – dominated community suggests the latter was pivotal in this modification of ecosystem function. We argue that this group used glucose to improve its defensive or its competitive abilities (or both). Adding such flexibility in bacterial traits to the model, we show how it creates the observed resilience to top-down manipulations observed in our experiment.

SummaryFor more than 25 years, virus‐to‐bacteria ratios (VBR) have been measured and interpreted as indicators of the importance of viruses in aquatic ecosystems, yet a generally accepted theory for understanding mechanisms controlling VBR is still lacking. Assuming that the denominator (total bacterial abundance) is primarily predator controlled, while viral lysis compensates for host growth rates exceeding this grazing loss, the numerator (viral abundance) reflects activity differences between prokaryotic hosts. VBR is then a ratio between mechanisms generating structure within the bacterial community and interactions between different plankton functional types controlling bacterial community size. We here show how these arguments can be formalized by combining a recently published model for co‐evolutionary host‐virus interactions, with a previously published “minimum” model for the microbial food web. The result is a framework where viral lysis links bacterial diversity to microbial food web structure and function, creating relationships between different levels of organization that are strongly modified by organism‐level properties such as cost of resistance.

Interactions of bottom-up factors such as the availability of mineral and organic nutrients, and top-down factors like predation and viral infection affect microbial communities of the pelagic food web. Hypothesis derived from previous experimental and modeling work suggest some general mechanisms on how these factors may be linked and call for an experiment to test interactions between (1) nitrogen source and diatom cell size, (2) bacterial growth rate limitation by mineral nutrients versus organic carbon, and (3) enhanced versus decreased predation pressure by copepods (Calanus finmarchicus). We performed a mesocosm experiment using a replicated three factor--two level full factorial design (NH 1 4 /NO 2 3 , 1/2 glucose, and 1/2 copepods). Diatoms, pico-and nano-sized autotrophs, prokaryotes, heterotrophic nanoflagellates, micro-and mesozooplankton abundances were monitored for 19 days. The main finding was a system that responded remarkably little to these manipulations with no effects of nitrogen source and only moderate effects of carbon treatments. Zooplankton manipulations had the strongest impact on microbial communities, but, opposite to our hypothesis based on previous studies, microzooplankton and diatoms increased during the first 7-10 days when mesozooplankton abundances were experimentally enhanced. We suggest that high abundances of dinoflagellates and ciliates and occurrence of rotifers, which dominated microzooplankton community in the first half of the experiment complicated trophic linkages between copepods, microzooplankton and diatoms. Apart from this dampened copepod mediated trophic cascade, we found tight couplings between communities of the microbial food web, with maxima of predator communities coinciding with low abundances of prey communities.

Quantification of grazing losses of marine heterotrophic bacteria is critical for understanding nutrient and carbon pathways in aquatic systems. The dilution method is a commonly used experimental approach for quantifying bacterivory. However, valid estimates of grazing rates obtained using this method depend on several methodological assumptions including that the method does not influence specific growth rates of bacteria. Here, we hypothesize that filtration during the set-up of a dilution experiment has the potential to release allelochemicals from phytoplankton cells and thereby stimulate or inhibit bacterial growth with the consequence of biased grazing estimates. We tested this hypothesis during a natural Phaeocystis pouchetii bloom at two different locations within an Arctic fjord. Results from the dilution experiments suggest higher gross growth rate and grazing impact for bacteria in the outer fjord compared with the inner fjord. However, specific growth rates estimated by bacterial production cell 21 were significantly elevated in dilutions of water from the outer fjord but not the inner fjord. The analysis of dissolved metabolites in the seawater from both experiments prior and after filtration revealed altered metabolic profiles after filtration at both stations. As unaffected specific growth of prey on dilution is one of three fundamental assumptions of the dilution method, we conclude that it is important that empirically estimated bacterial specific growth rates be routinely included when using the dilution method to quantify bacterivory.