How trophic cascades and photic zone nutrient content interact to generate basin-scale differences in the microbial food web

In linear food chains, resource and predator control produce positive and negative correlations, respectively, between biomass at adjacent trophic levels. These simple relationships become more complex in food webs that contain alternative food chains of unequal lengths. We have used a “minimum” model for the microbial part of the pelagic food web that has three such food chains connecting free mineral nutrients to copepods: via diatoms, autotrophic flagellates, and heterotrophic bacteria. Trophic cascades from copepods strongly modulates the balance between the three pathways and, therefore, the functionality of the microbial food web in services such as food production for higher trophic levels, DOM degradation, and ocean carbon sequestration. The result is a theoretical framework able to explain, not only apparent conflicts in Arctic mesocosm experiments, but also biogeochemical features of the Mediterranean. Here, the fundamental difference between Arctic and Mediterranean microbial food webs is the way they are predator driven by seasonal migration of large copepods in the Arctic, but resource driven due to the anti-estuarine circulation in the Mediterranean. In this framework, global change effects on microbial ecosystem functions are more like to come indirectly through changes in these drivers than through direct temperature effects on the microbes.publishedVersio

How trophic cascades and photic zone nutrient content interact to generate basin-scale differences in the microbial food web

In linear food chains, resource and predator control produce positive and negative correlations, respectively, between biomass at adjacent trophic levels. These simple relationships become more complex in food webs that contain alternative food chains of unequal lengths. We have used a “minimum” model for the microbial part of the pelagic food web that has three such food chains connecting free mineral nutrients to copepods: via diatoms, autotrophic flagellates, and heterotrophic bacteria. Trophic cascades from copepods strongly modulates the balance between the three pathways and, therefore, the functionality of the microbial food web in services such as food production for higher trophic levels, DOM degradation, and ocean carbon sequestration. The result is a theoretical framework able to explain, not only apparent conflicts in Arctic mesocosm experiments, but also biogeochemical features of the Mediterranean. Here, the fundamental difference between Arctic and Mediterranean microbial food webs is the way they are predator driven by seasonal migration of large copepods in the Arctic, but resource driven due to the anti-estuarine circulation in the Mediterranean. In this framework, global change effects on microbial ecosystem functions are more like to come indirectly through changes in these drivers than through direct temperature effects on the microbes

Fractal hypothesis of the pelagic microbial ecosystem-can simple ecological principles lead to self-similar complexity in the pelagic microbial food web?

Trophic interactions are highly complex and modern sequencing techniques reveal enormous biodiversity across multiple scales in marine microbial communities. Within the chemically and physically relatively homogeneous pelagic environment, this calls for an explanation beyond spatial and temporal heterogeneity. Based on observations of simple parasite-host and predator-prey interactions occurring at different trophic levels and levels of phylogenetic resolution, we present a theoretical perspective on this enormous biodiversity, discussing in particular self-similar aspects of pelagic microbial food web organization. Fractal methods have been used to describe a variety of natural phenomena, with studies of habitat structures being an application in ecology. In contrast to mathematical fractals where pattern generating rules are readily known, however, identifying mechanisms that lead to natural fractals is not straight-forward. Here we put forward the hypothesis that trophic interactions between pelagic microbes may be organized in a fractal-like manner, with the emergent network resembling the structure of the Sierpinski triangle. We discuss a mechanism that could be underlying the formation of repeated patterns at different trophic levels and discuss how this may help understand characteristic biomass size-spectra that hint at scale-invariant properties of the pelagic environment. If the idea of simple underlying principles leading to a fractal-like organization of the pelagic food web could be formalized, this would extend an ecologists mindset on how biological complexity could be accounted for. It may furthermore benefit ecosystem modeling by facilitating adequate model resolution across multiple scales

Exploring the distance between nitrogen and phosphorus limitation in mesotrophic surface waters using a sensitive bioassay

The balance in microbial net consumption of nitrogen and phosphorus was investigated in samples collected in two mesotrophic coastal environments: the Baltic Sea (Tvarminne field station) and the North Sea (Espegrend field station). For this, we have refined a bioassay based on the response in alkaline phosphatase activity (APA) over a matrix of combinations in nitrogen and phosphorus additions. This assay not only provides information on which element (N or P) is the primary limiting nutrient, but also gives a quantitative estimate for the excess of the secondary limiting element (P+ or N+, respectively), as well as the ratio of balanced net consumption of added N and P over short timescales (days). As expected for a Baltic Sea late springearly summer situation, the Tvarminne assays (n = 5) indicated N limitation with an average P+ = 0.30 +/- 0.10 mu M-P, when incubated for 4 days. For short incubations (1-2 days), the Espegrend assays indicated P limitation, but the shape of the response surface changed with incubation time, resulting in a drift in parameter estimates toward N limitation. Extrapolating back to zero incubation time gave P limitation with N+ approximate to 0.9 mu M-N. The N : P ratio (molar) of nutrient net consumption varied considerably between investigated locations: from 2.3 +/- 0.4 in the Tvarminne samples to 13 +/- 5 and 32 +/- 3 in two samples from Espegrend. Our assays included samples from mesocosm acidification experiments, but statistically significant effects of ocean acidification were not found by this method.Peer reviewe

Nutrient pathways through the microbial food web: principles and predictability discussed, based on five different experiments

Although explanatory and predictive powers are 2 closely interconnected aspects of conceptual and mathematical models of complex systems, the two are not equivalent. The 2 aspects are discussed here for the microbial part of photic zone food webs of the marine pelagic. We focus on the specific question of how limiting nutrients are transferred from the dissolved form, through the microbial food web, to mesozooplankton. For this purpose, 5 different nutrient addition experiments are reviewed and compared to a ‘simplest possible’ conceptual food web model. The experiments range in scale from artificial food webs constructed in laboratory chemostats, via mesocosm experiments, to a Lagrangian open-ocean addition experiment and cover time scales from days to weeks. We conclude that main system responses in all cases can be explained within the framework of the simple model, and that each experiment therefore also adds credibility to the basic concepts of this model. However, different system attributes profoundly affect the pathway and speed of nutrient transfer in each experiment. A re-occurring theme seems to be how the interactions between flexible stoichiometry and predatory processes modify experimental outcomes. Understanding the flexibility in the behavior of the system has thus increased with each experiment, but the requirement for new ad hoc assumptions to be added to the basic model structure in each case makes reliable predictions of the experimental outcome appear only possible with further model elaboration

What difference does it make if viruses are strain-, rather than species-specific?

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 represented by 16S rDNA 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 suggest 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

Optimal defense strategies in an idealized microbial food web under trade-off between competition and defense

Trophic mechanisms that can generate biodiversity in food webs include bottom-up (growth rate regulating) and top-down (biomass regulating) factors. The top-down control has traditionally been analyzed using the concepts of “Keystone Predation” (KP) and “Killing-the-Winner” (KtW), predominately occuring in discussions of macro- and micro-biological ecology, respectively. Here we combine the classical diamond-shaped food web structure frequently discussed in KP analyses and the KtW concept by introducing a defense strategist capable of partial defense. A formalized description of a trade-off between the defense-strategist's competitive and defensive ability is included. The analysis reveals a complex topology of the steady state solution with strong relationships between food web structure and the combination of trade-off, defense strategy and the system's nutrient content. Among the results is a difference in defense strategies corresponding to maximum biomass, production, or net growth rate of invading individuals. The analysis thus summons awareness that biomass or production, parameters typically measured in field studies to infer success of particular biota, are not directly acted upon by natural selection. Under coexistence with a competition specialist, a balance of competitive and defensive ability of the defense strategist was found to be evolutionarily stable, whereas stronger defense was optimal under increased nutrient levels in the absence of the pure competition specialist. The findings of success of different defense strategies are discussed with respect to SAR11, a highly successful bacterial clade in the pelagic ocean

Linking internal and external bacterial community control gives mechanistic framework for pelagic virus-to-bacteria ratios

For 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

Organic carbon and mineral nutrient limitation of oxygen consumption, bacterial growth and efficiency in the Norwegian Sea

To evaluate the role of bacteria in the transformation of organic matter in subarctic waters, we investigated the effect of mineral nutrients (ammonia and phosphate) and organic carbon (glucose) enrichment on heterotrophic bacterial processes and community structure. Eight experiments were done in the Norwegian Sea during May and June 2008. The growth-limiting factor (carbon or mineral nutrient) for heterotrophic bacteria was inferred from the combination of nutrient additions that stimulated highest bacterial oxygen consumption, biomass, production, growth rate and bacterial efficiency. We conclude that heterotrophic bacteria were limited by organic carbon and co-limited by mineral nutrients during the prevailing early nano-phytoplankton (1–10 lm) bloom conditions. High nucleic acid (HNA) bacteria became dominant ([80%) only when labile carbon and mineral nutrient sources were available. Changes in bacterial community structure were investigated using denaturing gradient gel electrophoresis (DGGE) of polymerase chain reaction (PCR)-amplified 16S ribosomal RNA genes. The bacterial community structure changed during incubation time, but neither carbon nor mineral nutrient amendment induced changes at the end of the experiments. The lack of labile organic carbon and the availability of mineral nutrients are key factors controlling bacterial activity and the role of the microbial food web in carbon sequestration

Competition for inorganic and organic forms of nitrogen and phosphorous between phytoplankton and bacteria during an Emiliania huxleyi spring bloom (PeECE II)

SCOPUS: ar.jinfo:eu-repo/semantics/publishe