Regional ocean-colour chlorophyll algorithms for the Red Sea

The Red Sea is a semi-enclosed tropical marine ecosystem that stretches from the Gulf of Suez and Gulf of Aqaba in the north, to the Gulf of Aden in the south. Despite its ecological and economic importance, its biological environment is relatively unexplored. Satellite ocean-colour estimates of chlorophyll concentration (an index of phytoplankton biomass) offer an observational platform to monitor the health of the Red Sea. However, little is known about the optical properties of the region. In this paper, we investigate the optical properties of the Red Sea in the context of satellite ocean-colour estimates of chlorophyll concentration. Making use of a new merged ocean-colour product, from the European Space Agency (ESA) Climate Change Initiative, and in situ data in the region, we test the performance of a series of ocean-colour chlorophyll algorithms. We find that standard algorithms systematically overestimate chlorophyll when compared with the in situ data. To investigate this bias we develop an ocean-colour model for the Red Sea, parameterised to data collected during the Tara Oceans expedition, that estimates remote-sensing reflectance as a function of chlorophyll concentration. We used the Red Sea model to tune the standard chlorophyll algorithms and the overestimation in chlorophyll originally observed was corrected. Results suggest that the overestimation was likely due to an excess of CDOM absorption per unit chlorophyll in the Red Sea when compared with average global conditions. However, we recognise that additional information is required to test the influence of other potential sources of the overestimation, such as aeolian dust, and we discuss uncertainties in the datasets used. We present a series of regional chlorophyll algorithms for the Red Sea, designed for a suite of ocean-colour sensors, that may be used for further testing.

Interaction Effects of Light, Temperature and Nutrient Limitations (N, P and Si) on Growth, Stoichiometry and Photosynthetic Parameters of the Cold-Water Diatom Chaetoceros wighamii

Light (20-450 μmol photons m^-2 s^-1), temperature (3-11°C) and inorganic nutrient composition (nutrient replete and N, P and Si limitation) were manipulated to study their combined influence on growth, stoichiometry (C:N:P:Chl a) and primary production of the cold water diatom Chaetoceros wighamii. During exponential growth, the maximum growth rate (~0.8 d^-1) was observed at high temperture and light; at 3°C the growth rate was ~30% lower under similar light conditions. The interaction effect of light and temperature were clearly visible from growth and cellular stoichiometry. The average C:N:P molar ratio was 80:13:1 during exponential growth, but the range, due to different light acclimation, was widest at the lowest temperature, reaching very low C:P (~50) and N:P ratios (~8) at low light and temperature. The C:Chl a ratio had also a wider range at the lowest temperature during exponential growth, ranging 16-48 (weight ratio) at 3°C compared with 17-33 at 11°C. During exponential growth, there was no clear trend in the Chl a normalized, initial slope (α*) of the photosynthesis-irradiance (PE) curve, but the maximum photosynthetic production (P_m) was highest for cultures acclimated to the highest light and temperature. During the stationary growth phase, the stoichiometric relationship depended on the limiting nutrient, but with generally increasing C:N:P ratio. The average photosynthetic quotient (PQ) during exponential growth was 1.26 but decreased to <1 under nutrient and light limitation, probably due to photorespiration. The results clearly demonstrate that there are interaction effects between light, temperature and nutrient limitation, and the data suggests greater variability of key parameters at low temperature. Understanding these dynamics will be important for improving models of aquatic primary production and biogeochemical cycles in a warming climate.

Effects of elevated CO2 and temperature on an intertidal meiobenthic community

In the near future, the marine environment is likely to be subjected to simultaneous increases in temperature and decreased pH. The potential effects of these changes on intertidal, meiofaunal assemblages were investigated using a mesocosm experiment. Artificial Substrate Units containing meiofauna from the extreme low intertidal zone were exposed for 60 days to eight experimental treatments (four replicates for each treatment) comprising four pH levels: 8.0 (ambient control), 7.7 & 7.3 (predicted changes associated with ocean acidification), and 6.7 (CO2 point-source leakage from geological storage), crossed with two temperatures: 12 °C (ambient control) and 16 °C (predicted). Community structure, measured using major meiofauna taxa was significantly affected by pH and temperature. Copepods and copepodites showed the greatest decline in abundance in response to low pH and elevated temperature. Nematodes increased in abundance in response to low pH and temperature rise, possibly caused by decreased predation and competition for food owing to the declining macrofauna density. Nematode species composition changed significantly between the different treatments, and was affected by both seawater acidification and warming. Estimated nematode species diversity, species evenness, and the maturity index, were substantially lower at 16 °C, whereas trophic diversity was slightly higher at 16 °C except at pH 6.7. This study has demonstrated that the combination of elevated levels of CO2 and ocean warming may have substantial effects on structural and functional characteristics of meiofaunal and nematode communities, and that single stressor experiments are unlikely to encompass the complexity of abiotic and biotic interactions. At the same time, ecological interactions may lead to complex community responses to pH and temperature changes in the interstitial environment

Combining metagenomics with metaproteomics and stable isotope probing reveals metabolic pathways used by a naturally occurring marine methylotroph

A variety of culture-independent techniques have been developed that can be used in conjunction with culture-dependent physiological and metabolic studies of key microbial organisms, in order to better understand how the activity of natural populations influences and regulates all major biogeochemical cycles. In this study, we combined DNA-stable isotope probing with metagenomics and metaproteomics to characterize an as yet uncultivated marine methylotroph that actively incorporated carbon from 13C-labeled methanol into biomass. By metagenomic sequencing of the heavy DNA, we retrieved virtually the whole genome of this bacterium and determined its metabolic potential. Through protein-stable isotope probing, the RuMP cycle was established as the main carbon assimilation pathway, and the classical methanol dehydrogenase-encoding gene mxaF, as well as three out of four identified xoxF homologues were found to be expressed. This proof-of-concept study is the first in which theculture-independent techniques of DNA- and protein-stable isotope probing have been used to characterize the metabolism of a naturally-ocurring Methylophaga-like bacterium in the marine environment (i.e. M. thiooxydans L4) and thus provides a powerful approach to access the genome and proteome of uncultivated microbes involved in key processes in the environment

Satellite estimates of net community production indicate predominance of net autotrophy in the Atlantic Ocean

There is ongoing debate as to whether the oligotrophic ocean is predominantly net autotrophic and acts as a CO2 sink, or net heterotrophic and therefore acts as a CO2 source to the atmosphere. This quantification is challenging, both spatially and temporally, due to the sparseness of measurements. There has been a concerted effort to derive accurate estimates of phytoplankton photosynthesis and primary production from satellite data to fill these gaps; however there have been few satellite estimates of net community production (NCP). In this paper, we compare a number of empirical approaches to estimate NCP from satellite data with in vitro measurements of changes in dissolved O2 concentration at 295 stations in the N and S Atlantic Ocean (including the Antarctic), Greenland and Mediterranean Seas. Algorithms based on power laws between NCP and particulate organic carbon production (POC) derived from 14C uptake tend to overestimate NCP at negative values and underestimate at positive values. An algorithm that includes sea surface temperature (SST) in the power function of NCP and 14C POC has the lowest bias and root-mean square error compared with in vitro measured NCP and is the most accurate algorithm for the Atlantic Ocean. Nearly a 13 year time series of NCP was generated using this algorithm with SeaWiFS data to assess changes over time in different regions and in relation to climate variability. The North Atlantic subtropical and tropical Gyres (NATL) were predominantly net autotrophic from 1998 to 2010 except for boreal autumn/winter, suggesting that the northern hemisphere has remained a net sink for CO2 during this period. The South Atlantic subtropical Gyre (SATL) fluctuated from being net autotrophic in austral spring-summer, to net heterotrophic in austral autumn–winter. Recent decadal trends suggest that the SATL is becoming more of a CO2 source. Over the Atlantic basin, the percentage of satellite pixels with negative NCP was 27%, with the largest contributions from the NATL and SATL during boreal and austral autumn–winter, respectively. Variations in NCP in the northern and southern hemispheres were correlated with climate indices. Negative correlations between NCP and the multivariate ENSO index (MEI) occurred in the SATL, which explained up to 60% of the variability in NCP. Similarly there was a negative correlation between NCP and the North Atlantic Oscillation (NAO) in the Southern Sub-Tropical Convergence Zone (SSTC),which explained 90% of the variability. There were also positive correlations with NAO in the Canary Current Coastal Upwelling (CNRY) and Western Tropical Atlantic (WTRA)which explained 80% and 60% of the variability in each province, respectively. MEI and NAO seem to play a role in modifying phases of net autotrophy and heterotrophy in the Atlantic Ocean.

Mechanisms shaping size structure and functional diversity of phytoplankton communities in the ocean

The factors regulating phytoplankton community composition play a crucial role in structuring aquatic food webs. However, consensus is still lacking about the mechanisms underlying the observed biogeographical differences in cell size composition of phytoplankton communities. Here we use a trait-based model to disentangle these mechanisms in two contrasting regions of the Atlantic Ocean. In our model, the phytoplankton community can self-assemble based on a trade-off emerging from relationships between cell size and (1) nutrient uptake, (2) zooplankton grazing, and (3) phytoplankton sinking. Grazing 'pushes' the community towards larger cell sizes, whereas nutrient uptake and sinking 'pull' the community towards smaller cell sizes. We find that the stable environmental conditions of the tropics strongly balance these forces leading to persistently small cell sizes and reduced size diversity. In contrast, the seasonality of the temperate region causes the community to regularly reorganize via shifts in species composition and to exhibit, on average, bigger cell sizes and higher size diversity than in the tropics. Our results raise the importance of environmental variability as a key structuring mechanism of plankton communities in the ocean and call for a reassessment of the current understanding of phytoplankton diversity patterns across latitudinal gradients.

Energy production and ecosystem services

Ecosystems provide a range of goods and services that contribute toward human well-being. It is increasingly recognized that factors such as a growing and increasingly affluent world population, coupled with increased globalization of trade, are significantly influencing the delivery of ecosystem goods and services. This chapter argues that future energy policy must be designed based on a broad set of environmental and social considerations that examine the national and international implications of each energy technology. This approach ensures a more holistic overview of the costs and benefits associated with energy production, allowing society to make more informed choices about their futures, including how their energy is sourced, generated, and delivered.

Energy and ecosystem service impacts

Ecosystems provide a range of goods and services that contribute toward human well-being through the environmental, economic, and cultural benefits they provide. Although the importance of these services is increasingly being recognized by governments, our understanding of the implications of different energy technologies on the provision of these services is limited. The chapter presents an assessment of four key energy technologies that considers the ecosystem services impacts across the entire lifecycle. In demonstrating the global implications of these energy technologies, the chapter makes the case that assessment of UK energy policy must consider a broad range of environmental and societal indicators both within the UK and overseas.

Phytoplankton phenology indices in coral reef ecosystems: Application to ocean-color observations in the Red Sea

Phytoplankton, at the base of the marine food web, represent a fundamental food source in coral reef ecosystems. The timing (phenology) and magnitude of the phytoplankton biomass are major determinants of trophic interactions. The Red Sea is one of the warmest and most saline basins in the world, characterized by an arid tropical climate regulated by the monsoon. These extreme conditions are particularly challenging for marine life. Phytoplankton phenological indices provide objective and quantitative metrics to characterize phytoplank- ton seasonality. The indices i.e. timings of initiation, peak, termination and duration are estimated here using 15 years (1997–2012) of remote sensing ocean-color data from the European Space Agency (ESA) Climate Change Initiative project (OC-CCI) in the entire Red Sea basin. The OC-CCI product, comprising merged and bias-corrected observations from three independent ocean-color sensors (SeaWiFS, MODIS and MERIS), and processed using the POLYMER algorithm (MERIS period), shows a significant increase in chlorophyll data cover- age, especially in the southern Red Sea during the months of summer NW monsoon. In open and reef-bound coastal waters, the performance of OC-CCI chlorophyll data is shown to be comparable with the performance of other standard chlorophyll products for the global oceans. These features have permitted us to investigate phytoplankton phenology in the entire Red Sea basin, and during both winter SE monsoon and summer NW monsoon periods. The phenological indices are estimated in the four open water provinces of the basin, and further examined at six coral reef complexes of particular socio-economic importance in the Red Sea, including Siyal Islands, Sharm El Sheikh, Al Wajh bank, Thuwal reefs, Al Lith reefs and Farasan Islands. Most of the open and deeper waters of the basin show an apparent higher chlorophyll concentration and longer duration of phyto- plankton growth during the winter period (relative to the summer phytoplankton growth period). In contrast, most of the reef-bound coastal waters display equal or higher peak chlorophyll concentrations and equal or lon- ger duration of phytoplankton growth during the summer period (relative to the winter phytoplankton growth period). The ecological and biological significance of the phytoplankton seasonal characteristics are discussed in context of ecosystem state assessment, and particularly to support further understanding of the structure and functioning of coral reef ecosystems in the Red Sea.

Altered Lipid Accumulation in Nannochlropsis Salina CCAP849/3 Following EMS and UV Induced Mutagenesis

Microalgae have potential as a chemical feed stock in a range of industrial applications. Nannochloropsis salina was subject to EMS mutagenesis and the highest lipid containing cells selected using fluorescence-activated cell sorting. Assessment of growth, lipid content and fatty acid composition identified mutant strains displaying a range of altered traits including changes in the PUFA content and a total FAME increase of up to 156% that of the wild type strain. Combined with a reduction in growth this demonstrated a productivity increase of up to 76%. Following UV mutagenesis, lipid accumulation of the mutant cultures was elevated to more than 3 fold that of the wild type strain, however reduced growth rates resulted in a reduction in overall productivity. Changes observed are indicative of alterations to the regulation of the omega 6 Kennedy pathway. The importance of these variations in physiology for industrial applications such as biofuel production is discussed.

The effects of short-term acidification on the intertidal macroalgae Palmaria palmate and Saccharina latissima

Ocean acidification, the result of increased dissolution of carbon dioxide (CO2) in seawater, is a leading subject of current research. The effects of acidification on non-calcifying macroalgae are, however, still unclear. The current study reports two 1-month studies using two different macroalgae, the red alga Palmaria palmata (Rhodophyta) and the kelp Saccharina latissima (Phaeophyta), exposed to control (pHNBS = ∼8.04) and increased (pHNBS = ∼7.82) levels of CO2-induced seawater acidification. The impacts of both increased acidification and time of exposure on net primary production (NPP), respiration (R), dimethylsulphoniopropionate (DMSP) concentrations, and algal growth have been assessed. In P. palmata, although NPP significantly increased during the testing period, it significantly decreased with acidification, whereas R showed a significant decrease with acidification only. S. latissima significantly increased NPP with acidification but not with time, and significantly increased R with both acidification and time, suggesting a concomitant increase in gross primary production. The DMSP concentrations of both species remained unchanged by either acidification or through time during the experimental period. In contrast, algal growth differed markedly between the two experiments, in that P. palmata showed very little growth throughout the experiment, while S. latissima showed substantial growth during the course of the study, with the latter showing a significant difference between the acidified and control treatments. These two experiments suggest that the study species used here were resistant to a short-term exposure to ocean acidification, with some of the differences seen between species possibly linked to different nutrient concentrations between the experiments.

Arctic Oceanography - Oceanography: Atmosphere-Ocean Exchange, Biogeochemistry & Physics

The Arctic Ocean is, on average, the shallowest of Earth’s oceans. Its vast continental shelf areas, which account for approximately half of the Arctic Ocean’s total area, are heavily influenced by the surrounding land masses through river run-off and coastal erosion. As a main area of deep water formation, the Arctic is one of the main «engines» of global ocean circulation, due to large freshwater inputs, it is also strongly stratified. The Arctic Ocean’s complex oceanographic configuration is tightly linked to the atmosphere, the land, and the cryosphere. The physical dynamics not only drive important climate and global circulation patterns, but also control biogeochemical cycles and ecosystem dynamics. Current changes in Arctic sea-ice thickness and distribution, air and water temperatures, and water column stability are resulting in measurable shifts in the properties and functioning of the ocean and its ecosystems. The Arctic Ocean is forecast to shift to a seasonally ice-free ocean resulting in changes to physical, chemical, and biological processes. These include the exchange of gases across the atmosphere-ocean interface, the wind-driven ciruclation and mixing regimes, light and nutrient availability for primary production, food web dynamics, and export of material to the deep ocean. In anticipation of these changes, extending our knowledge of the present Arctic oceanography and these complex changes has never been more urgent.

Marine and coastal policy in the UK: Challenges and opportunities in a new era

Marine and coastal policy in the UK has faced a number of significant changes in recent years, most notably the passing of the Marine and Coastal Access Act in 2009. These changes have brought significant challenges and opportunities for all those involved in the management and use of the UK's marine and coastal environment. This new era of marine policy inspired the UK's first Marine and Coastal Policy forum held in June 2011. In this introductory paper the global context of marine policy changes and the themes which emerged from the forum, forming the basis of the articles in this special issue, are outlined. It is concluded that there is a high level of engagement, capacity and willingness of key stakeholders to work collaboratively to address the environmental, social and economic complexities of managing the marine and coastal environment. It is both evident and encouraging that progress is being made and the many challenges faced in this new era give rise to a number of opportunities to develop new ideas and effective mechanisms for finding solutions

Hierarchical random walks in trace fossils and the origin of optimal search behavior

Efficient searching is crucial for timely location of food and other resources. Recent studies show diverse living animals employ a theoretically optimal scale-free random search for sparse resources known as a Lévy walk, but little is known of the origins and evolution of foraging behaviour and the search strategies of extinct organisms. Here we show using simulations of self-avoiding trace fossil trails that randomly introduced strophotaxis (U-turns) – initiated by obstructions such as ¬¬¬self-trail avoidance or innate cueing – leads to random looping patterns with clustering across increasing scales that is consistent with the presence of Lévy walks. This predicts optimal Lévy searches can emerge from simple behaviours observed in fossil trails. We then analysed fossilized trails of benthic marine organisms using a novel path analysis technique and find the first evidence of Lévy-like search strategies in extinct animals. Our results show that simple search behaviours of extinct animals in heterogeneous environments give rise to hierarchically nested Brownian walk clusters that converge to optimal Lévy patterns. Primary productivity collapse and large-scale food scarcity characterising mass extinctions evident in the fossil record may have triggered adaptation of optimal Lévy-like searches. The findings suggest Lévy-like behaviour has been employed by foragers since at least the Eocene but may have a more ancient origin, which could explain recent widespread observations of such patterns among modern taxa.