The effect of viral plasticity on the persistence of host-virus systems.

Phenotypic plasticity plays an important role in the survival of individuals. In microbial host-virus systems, previous studies have shown the stabilizing effect that host plasticity has on the coexistence of the system. By contrast, it remains uncertain how the dependence of the virus on the metabolism of the host (i.e. "viral plasticity") shapes bacteria-phage population dynamics in general, or the stability of the system in particular. Moreover, bacteria-phage models that do not consider viral plasticity are now recognised as overly simplistic. For these reasons, here we focus on the effect of viral plasticity on the stability of the system under different environmental conditions. We compared the predictions from a standard bacteria-phage model, which neglects plasticity, with those of a modification that includes viral plasticity. We investigated under which conditions viral plasticity promotes coexistence, with or without oscillatory dynamics. Our analysis shows that including viral plasticity reveals coexistence in regions of the parameter space where models without plasticity predict a collapse of the system. We also show that viral plasticity tends to reduce population oscillations, although this stabilizing effect is not consistently observed across environmental conditions: plasticity may instead reinforce dynamic feedbacks between the host, the virus, and the environment, which leads to wider oscillations. Our results contribute to a deeper understanding of the dynamic control of bacteriophage on host populations observed in nature.

Seasonal copepod lipid pump promotes carbon sequestration in the deep North Atlantic.

Estimates of carbon flux to the deep oceans are essential for our understanding of global carbon budgets. Sinking of detrital material ("biological pump") is usually thought to be the main biological component of this flux. Here, we identify an additional biological mechanism, the seasonal "lipid pump," which is highly efficient at sequestering carbon into the deep ocean. It involves the vertical transport and metabolism of carbon rich lipids by overwintering zooplankton. We show that one species, the copepod Calanus finmarchicus overwintering in the North Atlantic, sequesters an amount of carbon equivalent to the sinking flux of detrital material. The efficiency of the lipid pump derives from a near-complete decoupling between nutrient and carbon cycling­a "lipid shunt," and its direct transport of carbon through the mesopelagic zone to below the permanent thermocline with very little attenuation. Inclusion of the lipid pump almost doubles the previous estimates of deep-ocean carbon sequestration by biological processes in the North Atlantic.

Projected impacts of 21st century climate change on diapause in Calanus finmarchicus.

Diapause plays a key role in the life cycle of high latitude zooplankton. During diapause, animals avoid starving in winter by living in deep waters where metabolism is lower and met by lipid reserves. Global warming is therefore expected to shorten the maximum potential diapause duration by increasing metabolic rates and by reducing body size and lipid reserves. This will alter the phenology of zooplankton, impact higher trophic levels and disrupt biological carbon pumps. Here, we project the impacts of climate change on the key North Atlantic copepod Calanus finmarchicus under IPCC RCP 8.5. Potential diapause duration is modelled in relation to body size and overwintering temperature. The projections show pronounced geographic variations. Potential diapause duration reduces by more than 30% in the Western Atlantic, whereas in the key overwintering centre of the Norwegian Sea it changes only marginally. Surface temperature rises, which reduce body size and lipid reserves, will have a similar impact to deep-water changes on diapause in many regions. Because deep-water warming lags that at the surface, animals in the Labrador Sea could offset warming impacts by diapausing in deeper waters. However, the ability to control diapause depth may be limited.

Understanding patterns and processes in models of trophic cascades.

Climate fluctuations and human exploitation are causing global changes in nutrient enrichment of terrestrial and aquatic ecosystems and declining abundances of apex predators. The resulting trophic cascades have had profound effects on food webs, leading to significant economic and societal consequences. However, the strength of cascades-that is the extent to which a disturbance is diminished as it propagates through a food web-varies widely between ecosystems, and there is no formal theory as to why this should be so. Some food chain models reproduce cascade effects seen in nature, but to what extent is this dependent on their formulation? We show that inclusion of processes represented mathematically as density-dependent regulation of either consumer uptake or mortality rates is necessary for the generation of realistic 'top-down' cascades in simple food chain models. Realistically modelled 'bottom-up' cascades, caused by changing nutrient input, are also dependent on the inclusion of density dependence, but especially on mortality regulation as a caricature of, e.g. disease and parasite dynamics or intraguild predation. We show that our conclusions, based on simple food chains, transfer to a more complex marine food web model in which cascades are induced by varying river nutrient inputs or fish harvesting rates.

Elevated fish densities extend kilometres from oil and gas platforms.

Thousands of offshore oil and gas platforms have been installed throughout the world's oceans and more structures are being installed as part of the transition to renewable energy. These structures increase the availability of ecological niches by providing hard substrate in midwater and complex 3D habitat on the seafloor. This can lead to 'hotspots' of biodiversity, or increased densities of flora and fauna, which potentially spill over into the local area. However, the distances over which these higher densities extend (the 'range of influence') can be highly variable. Fish aggregate at such structures, but the range of influence and any implications for wider fish populations, are unclear. We investigated the relationship between fish and platform areal densities using high resolution fisheries acoustic data. Data were collected in the waters surrounding the vessel exclusions zones around 16 oil and gas platforms in the North Sea, and throughout the wider area. We estimated densities of schooling fish using echo-integration, and densities of non-schooling fish using echo-counting. At 10 platforms, non-schooling fish densities were elevated near the platform relative to background levels in the equivalent wider area. The range of influence, defined here as the range to which fish densities were elevated above background, varied from 0.8 to 23 km. In areas of high platform density, fish schools were encountered more often, and non-schooling fish densities were higher, when controlling for other sources of environmental variation. This is the first time such long-range effects have been identified; previously, ranges of influence have been reported in the order of just 10s-100s of metres. These findings suggest that the environmental impact of these structures may extend further than previously thought, which may be relevant in the context of upcoming management decisions around the decommissioning of these structures.

Unconstrained coevolution of bacterial size and the latent period of plastic phage.

Viruses play critical roles in the dynamics of microbial communities. Lytic viruses, for example, kill significant fractions of autotrophic and heterotrophic microbes daily. The dynamic interplay between viruses and microbes results from an overlap of physiological, ecological, and evolutionary responses: environmental changes trigger host physiological changes, affecting the ecological interactions of host and virus and, ultimately, the evolutionary pressures influencing the two populations. Recent theoretical work studied how the dependence of viral traits on host physiology (viral plasticity) affects the evolutionarily stable host cell size and viral infection time emerging from coevolution. Here, we broaden the scope of the framework to consider any coevolutionary outcome, including potential evolutionary collapses of the system. We used the case study of Escherichia coli and T-like viruses under chemostat conditions, but the framework can be adapted to any microbe-virus system. Oligotrophic conditions led to smaller, lower-quality but more abundant hosts, and infections that were longer but produced a reduced viral offspring. Conversely, eutrophic conditions resulted in fewer but larger higher-quality hosts, and shorter but more productive infections. The virus influenced host evolution decreasing host size more noticeably for low than for high dilution rates, and for high than for low nutrient input concentration. For low dilution rates, the emergent infection time minimized host need/use, but higher dilution led to an opportunistic strategy that shortened the duration of infections. System collapses driven by evolution resulted from host failure to adapt quickly enough to the evolving virus. Our results contribute to understanding the eco-evolutionary dynamics of microbes and virus, and to improving the predictability of current models for host-virus interactions. The large quantitative and qualitative differences observed with respect to a classic description (in which viral traits are assumed to be constant) highlights the importance of including viral plasticity in theories describing short- and long-term host-virus dynamics.

Ecosystem approach to harvesting in the Arctic: Walking the tightrope between exploitation and conservation in the Barents Sea.

Projecting the consequences of warming and sea-ice loss for Arctic marine food web and fisheries is challenging due to the intricate relationships between biology and ice. We used StrathE2EPolar, an end-to-end (microbes-to-megafauna) food web model incorporating ice-dependencies to simulate climate-fisheries interactions in the Barents Sea. The model was driven by output from the NEMO-MEDUSA earth system model, assuming RCP 8.5 atmospheric forcing. The Barents Sea was projected to be > 95% ice-free all year-round by the 2040s compared to > 50% in the 2010s, and approximately 2 °C warmer. Fisheries management reference points (FMSY and BMSY) for demersal fish (cod, haddock) were projected to increase by around 6%, indicating higher productivity. However, planktivorous fish (capelin, herring) reference points were projected to decrease by 15%, and upper trophic levels (birds, mammals) were strongly sensitive to planktivorous fish harvesting. The results indicate difficult trade-offs ahead, between harvesting and conservation of ecosystem structure and function.

Evolutionarily Stable Coevolution Between a Plastic Lytic Virus and Its Microbial Host.

Hosts influence and are influenced by viral replication. Cell size, for example, is a fundamental trait for microbial hosts that can not only alter the probability of viral adsorption, but also constrain the host physiological processes that the virus relies on to replicate. This intrinsic connection can affect the fitness of both host and virus, and therefore their mutual evolution. Here, we study the coevolution of bacterial hosts and their viruses by considering the dependence of viral performance on the host physiological state (viral plasticity). To this end, we modified a standard host-lytic phage model to include viral plasticity, and compared the coevolutionary strategies emerging under different scenarios, including cases in which only the virus or the host evolve. For all cases, we also obtained the evolutionary prediction of the traditional version of the model, which assumes a non-plastic virus. Our results reveal that the presence of the virus leads to an increase in host size and growth rate in the long term, which benefits both interacting populations. Our results also show that viral plasticity and evolution influence the classic host quality-quantity trade-off. Poor nutrient environments lead to abundant low-quality hosts, which tends to increase viral infection time. Conversely, richer nutrient environments lead to fewer but high-quality hosts, which decrease viral infection time. Our results can contribute to advancing our understanding of the microbial response to changing environments. For instance, both cell size and viral-induced mortality are essential factors that determine the structure and dynamics of the marine microbial community, and therefore our study can improve predictions of how marine ecosystems respond to environmental change. Our study can also help devise more reliable strategies to use phage to, for example, fight bacterial infections.

Population density and temperature correlate with long-term trends in somatic growth rates and maturation schedules of herring and sprat.

We examine long-term trends in the average growth rates and maturation schedules of herring and sprat populations using survey data collected from the North Sea and west of Scotland since the 1960s and 1980s respectively. Otolith age data and maturity data are used to calculate time series of mean lengths at age, von Bertalanffy growth parameters, and probabilistic maturation reaction norms. As the growth and maturation of fish is known to be influenced by temperature and stock abundances, we account for these variables using Generalised Additive Models. Each of the herring populations displayed either steady declines in mean length across multiple age groups, or declines in length followed years later by some recovery. Depending on region, lengths at age of sprat increased or decreased over time. Varying temporal trends in maturation propensity at age and length were observed across herring populations. Many of the trends in growth rate and maturation were correlated to population abundance and/or temperature. In general, abundance is shown to be negatively correlated to growth rates in herring and sprat, and positively correlated with maturation propensity in herring. Temperature is also shown to be correlated to growth and maturation, and although the effect is consistent within species, the temperature effects differ between herring and sprat. This study provides detailed information about long-term trends in growth and maturation, which is lacking for some of these pelagic stocks, especially in the west of Scotland.

Landscape effects on demersal fish revealed by field observations and predictive seabed modelling.

Nature conservation and fisheries management often focus on particular seabed features that are considered vulnerable or important to commercial species. As a result, individual seabed types are protected in isolation, without any understanding of what effect the mixture of seabed types within the landscape has on ecosystem functions. Here we undertook predictive seabed modelling within a coastal marine protected area using observations from underwater stereo-video camera deployments and environmental information (depth, wave fetch, maximum tidal speeds, distance from coast and underlying geology). The effect of the predicted substratum type, extent and heterogeneity or the diversity of substrata, within a radius of 1500 m around each camera deployment of juvenile gadoid relative abundance was analysed. The predicted substratum model performed well with wave fetch and depth being the most influential predictor variables. Gadus morhua (Atlantic cod) were associated with relatively more rugose substrata (Algal-gravel-pebble and seagrass) and heterogeneous landscapes, than Melanogrammus aeglefinus (haddock) or Merlangius merlangus (whiting) (sand and mud). An increase in M. merlangus relative abundance was observed with increasing substratum extent. These results reveal that landscape effects should be considered when protecting the seabed for fish and not just individual seabed types. The landscape approach used in this study therefore has important implications for marine protected area, fisheries management and monitoring advice concerning demersal fish populations.

Hind-casting the quantity and composition of discards by mixed demersal fisheries in the North Sea.

Many commercial fisheries seek to maximise the economic value of the catch that they bring ashore and market for human consumption by discarding undersize or low value fish. Information on the quantity, size and species composition of discarded fish is vital for stock assessments and for devising legislation to minimise the practice. However, except for a few major species, data are usually extremely sparse and reliant on observers aboard a small sample of fishing vessels. Expanding these data to estimate total regional discards is highly problematic. Here, we develop a method for utilising additional information from scientific trawl surveys to model the quantities of fish discarded by the commercial fisheries. As a case-study, we apply the model to the North Sea over the period 1978-2011, and show a long-term decline in the overall quantity of fish discarded, but an increase in the proportion of catch which is thrown away. The composition of discarded catch has shifted from predominantly (∼80%) roundfish, to >50% flatfish. Undersized plaice constitute the largest single fraction of discards, unchanged from the beginning of the 20th century. Overall, around 60% of discarded fish are rejected on the basis of size rather than for reasons of species value or quota restrictions. The analysis shows that much more information can be gained on discarding by utilising additional sources of data rather than relying solely on information gathered by observers. In addition, it is clear that reducing fishing intensity and rebuilding stocks is likely to be more effective at reducing discards in the long term, than any technical legislation to outlaw the practice in the short term.

Cascading ecological effects of eliminating fishery discards.

Discarding by fisheries is perceived as contrary to responsible harvesting. Legislation seeking to end the practice is being introduced in many jurisdictions. However, discarded fish are food for a range of scavenging species; so, ending discarding may have ecological consequences. Here we investigate the sensitivity of ecological effects to discarding policies using an ecosystem model of the North Sea--a region where 30-40% of trawled fish catch is currently discarded. We show that landing the entire catch while fishing as usual has conservation penalties for seabirds, marine mammals and seabed fauna, and no benefit to fish stocks. However, combining landing obligations with changes in fishing practices to limit the capture of unwanted fish results in trophic cascades that can benefit birds, mammals and most fish stocks. Our results highlight the importance of considering the broader ecosystem consequences of fishery management policy, since species interactions may dissipate or negate intended benefits.

Comment on "A global map of human impact on marine ecosystems".

Halpern et al. (Reports, 15 February 2008, p. 948) integrated spatial data on 17 drivers of change in the oceans to map the global distribution of human impact. Although fishery catches are a dominant driver, the data reflect activity while impacts occur at different space and time scales. Failure to account for this spatial disconnection could lead to potentially misleading conclusions.