Ocean carbon sequestration: Particle fragmentation by copepods as a significant unrecognised factor?: Explicitly representing the role of copepods in biogeochemical models may fundamentally improve understanding of future ocean carbon storage.

Ocean biology helps regulate global climate by fixing atmospheric CO2 and exporting it to deep waters as sinking detrital particles. New observations demonstrate that particle fragmentation is the principal factor controlling the depth to which these particles penetrate the ocean's interior, and hence how long the constituent carbon is sequestered from the atmosphere. The underlying cause is, however, poorly understood. We speculate that small, particle-associated copepods, which intercept and inadvertently break up sinking particles as they search for attached protistan prey, are the principle agents of fragmentation in the ocean. We explore this idea using a new marine ecosystem model. Results indicate that explicitly representing particle fragmentation by copepods in biogeochemical models offers a step change in our ability to understand the future evolution of biologically-mediated ocean carbon storage. Our findings highlight the need for improved understanding of the distribution, abundance, ecology and physiology of particle-associated copepods.

Remembering John Steele and his models for understanding the structure and function of marine ecosystems.

John Steele (1926-2013) is remembered for his ecosystem modelling studies on the role of biological interactions and environment on the structure and function of marine ecosystems, including consequences for fish production and fisheries management. Here, we provide a scientific tribute to Steele focusing on, by means of example, his modelling of plankton predation [Steele and Henderson (1992) The role of predation in plankton models. J. Plankton Res., 14, 157-172] that showed that differences in ecosystem dynamics between the subarctic Pacific and North Atlantic oceans can be explained solely on the basis of zooplankton mortality. The study highlights Steele's artistry in simplifying the system to a tractable minimal model while paying great attention to the precise functional forms used to parameterize mortality, grazing and other biological processes. The success of this and other works by Steele was in large part due to his effective communication with the rest of the scientific community (especially non-modellers) resulting from his enthusiasm, use of an experiment-like (hypothesis driven) approach to applying his models and by describing simplifications and assumptions in scrupulous detail. We also intend our contribution to remember Steele as the consummate gentleman, notably his humble, behind-the-scenes attitude, his humour and his dedication to enhancing the careers of others.

Intra-medullary nail insertion accuracy: A comparison of the infra-patellar and supra-patellar approach.

AIMS: The anatomical safe zone for intra-medullary nail insertion through the tibial plateau is small, insertion outside of this area risks damage to intra-articular structures and poor fracture reduction. The purpose of this retrospective study was to determine if the new supra-patella (SP) approach confers improved nail insertion accuracy, when compared with the standard infra-patella (IP) technique. PATIENTS AND METHODS: Two hundred cases were included in the study (SP 95, IP 105). Insertion accuracy was assessed on AP and lateral radiographic imaging, and measured as the distances between the central axis of the proximal nail and the ideal entry point. RESULTS: The median distance from the ideal entry point was 4.4 mm (SP) and 5.1 mm (IP) (p = 0.046) in the coronal plane, and 4.0 mm (SP) and 3.7 mm (IP) (p = 0.527) in the sagittal plane. A narrower range in measurements was observed in the SP technique in both sagittal and coronal planes, 17.8 mm vs 28.6 mm, and 19.7 mm vs 30.3 mm respectively. CONCLUSION: We found that the SP technique achieved significantly improved nail insertion accuracy in the coronal plane. Insertion accuracy was equivocal between the two techniques in the sagittal plane. A narrower range in entry points was observed in the SP cohort in both planes suggesting improved control in nail insertion using this technique.

Will Invertebrates Require Increasingly Carbon-Rich Food in a Warming World?

Elevated temperature causes metabolism and respiration to increase in poikilothermic organisms. We hypothesized that invertebrate consumers will therefore require increasingly carbon-rich diets in a warming environment because the increased energetic demands are primarily met using compounds rich in carbon, that is, carbohydrates and lipids. Here, we test this hypothesis using a new stoichiometric model that has carbon (C) and nitrogen (N) as currencies. Model predictions did not support the hypothesis, indicating instead that the nutritional requirements of invertebrates, at least in terms of food quality expressed as C∶N ratio, may change little, if at all, at elevated temperature. Two factors contribute to this conclusion. First, invertebrates facing limitation by nutrient elements such as N have, by default, excess C in their food that can be used to meet the increased demand for energy in a warming environment, without recourse to extra dietary C. Second, increased feeding at elevated temperature compensates for the extra demands of metabolism to the extent that, when metabolism and intake scale equally with temperature (have the same Q10), the relative requirement for dietary C and N remains unaltered. Our analysis demonstrates that future climate-driven increases in the C∶N ratios of autotroph biomass will likely exacerbate the stoichiometric mismatch between nutrient-limited invertebrate grazers and their food, with important consequences for C sequestration and nutrient cycling in ecosystems.

Big in the benthos: Future change of seafloor community biomass in a global, body size-resolved model.

Deep-water benthic communities in the ocean are almost wholly dependent on near-surface pelagic ecosystems for their supply of energy and material resources. Primary production in sunlit surface waters is channelled through complex food webs that extensively recycle organic material, but lose a fraction as particulate organic carbon (POC) that sinks into the ocean interior. This exported production is further rarefied by microbial breakdown in the abyssal ocean, but a residual ultimately drives diverse assemblages of seafloor heterotrophs. Advances have led to an understanding of the importance of size (body mass) in structuring these communities. Here we force a size-resolved benthic biomass model, BORIS, using seafloor POC flux from a coupled ocean-biogeochemistry model, NEMO-MEDUSA, to investigate global patterns in benthic biomass. BORIS resolves 16 size classes of metazoans, successively doubling in mass from approximately 1 µg to 28 mg. Simulations find a wide range of seasonal responses to differing patterns of POC forcing, with both a decline in seasonal variability, and an increase in peak lag times with increasing body size. However, the dominant factor for modelled benthic communities is the integrated magnitude of POC reaching the seafloor rather than its seasonal pattern. Scenarios of POC forcing under climate change and ocean acidification are then applied to investigate how benthic communities may change under different future conditions. Against a backdrop of falling surface primary production (-6.1%), and driven by changes in pelagic remineralization with depth, results show that while benthic communities in shallow seas generally show higher biomass in a warmed world (+3.2%), deep-sea communities experience a substantial decline (-32%) under a high greenhouse gas emissions scenario. Our results underscore the importance for benthic ecology of reducing uncertainty in the magnitude and seasonality of seafloor POC fluxes, as well as the importance of studying a broader range of seafloor environments for future model development.

CO2, the greenhouse effect and global warming: from the pioneering work of Arrhenius and Callendar to today's Earth System Models.

Climate warming during the course of the twenty-first century is projected to be between 1.0 and 3.7°C depending on future greenhouse gas emissions, based on the ensemble-mean results of state-of-the-art Earth System Models (ESMs). Just how reliable are these projections, given the complexity of the climate system? The early history of climate research provides insight into the understanding and science needed to answer this question. We examine the mathematical quantifications of planetary energy budget developed by Svante Arrhenius (1859-1927) and Guy Stewart Callendar (1898-1964) and construct an empirical approximation of the latter, which we show to be successful at retrospectively predicting global warming over the course of the twentieth century. This approximation is then used to calculate warming in response to increasing atmospheric greenhouse gases during the twenty-first century, projecting a temperature increase at the lower bound of results generated by an ensemble of ESMs (as presented in the latest assessment by the Intergovernmental Panel on Climate Change). This result can be interpreted as follows. The climate system is conceptually complex but has at its heart the physical laws of radiative transfer. This basic, or "core" physics is relatively straightforward to compute mathematically, as exemplified by Callendar's calculations, leading to quantitatively robust projections of baseline warming. The ESMs include not only the physical core but also climate feedbacks that introduce uncertainty into the projections in terms of magnitude, but not sign: positive (amplification of warming). As such, the projections of end-of-century global warming by ESMs are fundamentally trustworthy: quantitatively robust baseline warming based on the well-understood physics of radiative transfer, with extra warming due to climate feedbacks. These projections thus provide a compelling case that global climate will continue to undergo significant warming in response to ongoing emissions of CO2 and other greenhouse gases to the atmosphere.

The Role of Microbes in the Nutrition of Detritivorous Invertebrates: A Stoichiometric Analysis.

Detritus represents an important pool in the global carbon cycle, providing a food source for detritivorous invertebrates that are conspicuous components of almost all ecosystems. Our knowledge of how these organisms meet their nutritional demands on a diet that is typically comprised of refractory, carbon-rich compounds nevertheless remains incomplete. "Trophic upgrading" of detritus by the attached microbial community (enhancement of zooplankton diet by the inclusion of heterotrophic protozoans) represents a potential source of nutrition for detritivores as both bacteria and their flagellated protistan predators are capable of biosynthesizing essential micronutrients such as polyunsaturated fatty acids (PUFAs). There is however a trade-off because although microbes enhance the substrate in terms of its micronutrient content, the quantity of organic carbon is diminished though metabolic losses as energy passes through the microbial food web. Here, we develop a simple stoichiometric model to examine this trade-off in the nutrition of detritivorous copepods inhabiting the mesopelagic zone of the ocean, focusing on their requirements for carbon and an essential PUFA, docosahexaenoic acid (DHA). Results indicate that feeding on microbes may be a highly favorable strategy for these invertebrates, although the potential for carbon to become limiting when consuming a microbial diet exists because of the inefficiencies of trophic transfer within the microbial food web. Our study highlights the need for improved knowledge at the detritus-microbe-metazoan interface, including interactions between the physiology and ecology of the associated organisms.

Microbial gardening in the ocean's twilight zone: detritivorous metazoans benefit from fragmenting, rather than ingesting, sinking detritus: fragmentation of refractory detritus by zooplankton beneath the euphotic zone stimulates the harvestable production of labile and nutritious microbial biomass.

Sinking organic particles transfer ∼10 gigatonnes of carbon into the deep ocean each year, keeping the atmospheric CO2 concentration significantly lower than would otherwise be the case. The exact size of this effect is strongly influenced by biological activity in the ocean's twilight zone (∼50-1,000 m beneath the surface). Recent work suggests that the resident zooplankton fragment, rather than ingest, the majority of encountered organic particles, thereby stimulating bacterial proliferation and the deep-ocean microbial food web. Here we speculate that this apparently counterintuitive behaviour is an example of 'microbial gardening', a strategy that exploits the enzymatic and biosynthetic capabilities of microorganisms to facilitate the 'gardener's' access to a suite of otherwise unavailable compounds that are essential for metazoan life. We demonstrate the potential gains that zooplankton stand to make from microbial gardening using a simple steady state model, and we suggest avenues for future research.

Reconciliation of the carbon budget in the ocean's twilight zone.

Photosynthesis in the surface ocean produces approximately 100 gigatonnes of organic carbon per year, of which 5 to 15 per cent is exported to the deep ocean. The rate at which the sinking carbon is converted into carbon dioxide by heterotrophic organisms at depth is important in controlling oceanic carbon storage. It remains uncertain, however, to what extent surface ocean carbon supply meets the demand of water-column biota; the discrepancy between known carbon sources and sinks is as much as two orders of magnitude. Here we present field measurements, respiration rate estimates and a steady-state model that allow us to balance carbon sources and sinks to within observational uncertainties at the Porcupine Abyssal Plain site in the eastern North Atlantic Ocean. We find that prokaryotes are responsible for 70 to 92 per cent of the estimated remineralization in the twilight zone (depths of 50 to 1,000 metres) despite the fact that much of the organic carbon is exported in the form of large, fast-sinking particles accessible to larger zooplankton. We suggest that this occurs because zooplankton fragment and ingest half of the fast-sinking particles, of which more than 30 per cent may be released as suspended and slowly sinking matter, stimulating the deep-ocean microbial loop. The synergy between microbes and zooplankton in the twilight zone is important to our understanding of the processes controlling the oceanic carbon sink.

Deserts on the sea floor: Edward Forbes and his azoic hypothesis for a lifeless deep ocean.

While dredging in the Aegean Sea during the mid-19th century, Manxman Edward Forbes noticed that plants and animals became progressively more impoverished the greater the depth they were from the surface of the water. By extrapolation Forbes proposed his now infamous azoic hypothesis, namely that life would be extinguished altogether in the murky depths of the deep ocean. The whole idea seemed so entirely logical given the enormous pressure, cold and eternal darkness of this apparently uninhabitable environment. Yet we now know that the sea floor is teeming with life. Curiously, it took 25 years for the azoic hypothesis to fall from grace. This was despite the presence of ample contrary evidence, including starfishes, worms and other organisms that seemingly originated from the deep seabed. This is a tale of scientists ignoring observations that ran counter to their deep-seated, yet entirely erroneous, beliefs.

Statin-induced rhabdomyolysis.

Simvastatin and other HMG-CoA reductase inhibitors (statins) are one of the most frequently prescribed class of medications in the United States, with over 15 million Americans taking these drugs. Relatively rare adverse effects related to the known toxic effects of these drugs are more common than generally realized. Clinically significant statin-induced rhabdomyolysis is an uncommon but life-threatening adverse effect. We describe a case of simvastatin-induced rhabdomyolysis. Current knowledge of the pharmacology of the HMG-CoA reductase inhibitors and the drug interactions that potentiate these adverse effects are discussed. The clinical features of rhabdomyolysis and current treatment recommendations are presented.

Metabolic stoichiometry and the fate of excess carbon and nutrients in consumers.

Animals encountering nutritionally imbalanced foods should release elements in excess of requirements in order to maintain overall homeostasis. Quantifying these excesses and predicting their fate is, however, problematic. A new model of the stoichiometry of consumers is formulated that incorporates the separate terms in the metabolic budget, namely, assimilation of ingested substrates and associated costs, protein turnover, other basal costs, such as osmoregulation, and the use of remaining substrates for production. The model indicates that release of excess C and nonlimiting nutrients may often be a significant fraction of the total metabolic budget of animals consuming the nutrient-deficient forages that are common in terrestrial and aquatic systems. The cost of maintenance, in terms of not just C but also N and P, is considerable, such that food quality is important even when intake is low. Many generalist consumers experience short-term and unpredictable fluctuations in their diets. Comparison of model output with data for one such consumer, Daphnia, indicates that mechanisms operating postabsorption in the gut are likely the primary means of regulating excess C, N, and P in these organisms, notably respiration decoupled from biochemical or mechanical work and excretion of carbon and nutrients. This stoichiometrically regulated release may often be in organic rather than inorganic form, with important consequences for the balance of autotrophic and heterotrophic processes in ecosystems.

Stoichiometry and food-chain dynamics.

Traditional models of chemostat systems looking at interactions between predator, prey and nutrients have used only a single currency, such as energy or nitrogen. In reality, growth of autotrophs and heterotrophs may be limited by various elements, e.g. carbon, nitrogen, phosphorous or iron. In this study we develop a dynamic energy budget model chemostat which has both carbon and nitrogen as currencies, and examine how the dual availability of these elements affects the growth of phytoplankton, trophic transfer to zooplankton, and the resulting stability of the chemostat ecosystem. Both species have two reserve pools to obtain a larger metabolic flexibility with respect to changing external environments. Mineral nitrogen and carbon form the base of the food chain, and they are supplied at a constant rate. In addition, the biota in the chemostat recycle nutrients by means of respiration and excretion, and organic detritus is recycled at a fixed rate. We use numerical bifurcation analysis to assess the model's dynamic behavior. In the model, phytoplankton is nitrogen limited, and nitrogen enrichment can lead to oscillations and multiple stable states. Moreover, we found that recycling has a destabilizing effect on the food chain due to the increased repletion of mineral nutrients. We found that both carbon and nitrogen enrichment stimulate zooplankton growth. Therefore, we conclude that the concept of single-element limitation may not be applicable in an ecosystem context.