Application of the European Regional Seas Ecosystem Model (ERSEM) to assessing the eutrophication status in the OSPAR Maritime Area, with particular reference to nutrient discharges from Scottish salmonid aquaculture

Aquaculture production of salmonids in Scotland has grown over the last 15 years, exceeded 150,000 tonnes in 2001. There have been conflicting views as to the likely ecological impact of nutrient discharges from this activity. Whilst quantitative assessments of aquaculture nutrient discharges have been carried out, the debate regarding possible eutrophication impacts of these discharges has so far been largely speculative. In order to provide a quantitative basis for this discussion, a marine ecosystem model was used to simulate the consequences of a 50% reduction in aquaculture nutrient discharges, and the results are presented here

Modelling the behaviour of nutrients in the coastal waters of Scotland

The overall goal of this project was to provide Scotland with a strategic ecosystem simulation tool for identifying maritime areas which could be at risk of eutrophication. The tool should provide spatially resolved output, and be capable of discriminating between different types and locations of nutrient inputs, so as to enable scenario analyses of different reduction options. The specific aims of the project were firstly to simulate the annual cycles of nutrients and ecological properties of Scottish waters and advise on areas which might suffer from eutrophication, and secondly, to determine the contribution of Scottish nutrient discharges to eutrophication in the OSPAR maritime area as a whole

Modelling the behaviour of nutrients in the coastal waters of Scotland - an update on inputs from Scottish aquaculture and their impact on eutrophication status

A previous study estimated that salmon farming contributed approximately 6% of Scotland's nitrogen-nutrient input to coastal waters, and 13% of phosphorus (based on 2001 production figures). However, in some areas of the west of Scotland with small freshwater catchment areas and low levels of human habitation, aquaculture inputs represented greater than 80% of the total. In 2002, FRS published results from an ecosystem modelling study involving a collaboration with the Institute for Marine Research, University of Hamburg, and the Macaulay Land Use Research Institute in Aberdeen, to assess the eutrophication impact of various nutrient inputs to Scottish waters. The results suggested that a 50% reduction in aquaculture salmon production would have only a small impact on water quality which would be undetectable against the background of natural variability due to climate variations. Estimating aquaculture nutrient discharge is a difficult task. The 2002 study was based on data relating to the consented biomass of fish at farm sites in sea lochs. Since then, new data have become available on the actual harvest of fish at all sites in Scotland. In this report, we re-assess the salmon production in Scotland in 2001 and the consequent nutrient discharge, and repeat the ecosystem model runs to estimate the impact of reduction scenarios on eutrophication status. The new data indicate that the previous study had overestimated salmon production and nutrient discharge by approximately 18% Scotland wide. Production and discharge at Shetland and in the Southern Hebrides had been under-estimated, whilst that in the Minches had been over-estimated. New runs of the ecosystem model show that the original conclusions on eutrophication impact were sound. A scenario of 50% reduction in salmon production produced regional changes in water quality which were less than 25% of the natural variability due to climate. New runs simulating a cessation of aquaculture showed that even this extreme reduction scenario produced changes in water quality that were less than half the natural variability

Use of ocean colour remote sensing to monitor sea surface suspended sediments

Ocean colour remote sensing (OCRS) from satellite platforms has revolutionised our ability to monitor the interplay of physical and biogeochemical processes in surface waters of the ocean. Since the launch of SeaWiFS in 1996, a continuous time series of OCRS data has been accumulated from a series of satellite sensors giving near daily global coverage. These sensors measure top of atmosphere (TOA) spectral radiance which is corrected for atmospheric effects (~80% of the measured signal in the blue - Gordon 1978) to give water leaving radiances. From these putrely optical signals, it is possible to derive a wide range of higher level products such as chlorophyll concentration, diffuse attenuation coefficients, photosynthetically available radiation (PAR) and a wide range of inherent optical properties (IOPs) to name but a few. In terms of surface area and primary productivity, the global ocean is heavily dominated by deep, oceanic waters, where the optical properties are driven by phytoplankton, associated dissolved organics and water itself. It is little surprise then that early standard OCRS products were developed for optimal performance over these globally significant regions. Standard chlorophyll algorithms were developed using changes in blue-green reflectance ratios (e.g. O’Reilley et al., 1998) that can be related to the effect of changing concentrations of microscopic scale (1µm-200µm) phytoplankton (Kirk,1983) forming blooms that can stretch for thousands of km. More recently, attention has shifted to economically important coastal regions where, for example, harmful algal blooms have potential to cause significant societal and economic impact. OCRS algorithms have been developed to specifically aid in the monitoring of both toxic species e.g. Karenia brevis in the Gulf of Mexico (Stumpf et al., 2003), and also to monitor for extreme eutrophication events where excessive levels of phytoplankton cause the reduction of oxygen dissolved in the water column (hypoxia) leading to animal mortality (e.g. Mallin et al., 2006). The optically complex nature of coastal waters, more generally, presents a particular problem for OCRS applications in these regions. Shallow shelf seas and other inshore waters are subject to the influence of sediment resuspension and freshwater discharge bringing additional loads of coloured dissolved organic materials (CDOM). This results in multiple, independently varying, optically significant components, each of which influences the water leaving radiance spectrum making interpretation of spectral changes significantly more difficult. Many studies have demonstrated the breakdown in performance of standard algorithms (e.g. Chl, McKee et al. 2007) in optically complex coastal waters. In this paper we will focus on the effect of suspended sediment on optical properties of the water column. Suspended sediment has long been known to influence light penetration (Gordon and McCluney, 1975) which can limit primary production and also contribute to hypoxia (Greig et al., 2005). There is interest in monitoring sediment concentration for coastal erosion applications and various OCRS algorithms have been developed that exploit the relatively strong backscattering properties of sediment. For example, Doxaran et al. (2002) successfully presented a sediment algorithm for the highly turbid Gironde estuary. More recently a radiative transfer approach was used to refine this type of approach to incorporate the potential impact of other materials on the red reflectance values that support sediment algorithms (Neil et al., 2011). This algorithm provides estimates of maximum and minimum sediment load concentrations assuming reasonable potential ranges of Chl and CDOM for coastal waters. The aim of this paper is to determine the extent to which the Neil et al. algorithm, which was developed for Irish Sea waters, can be applied to data collected in the North Sea. The ultimate goal is to assess the potential for using OCRS data to monitor suspended sediment concentrations in coastal waters, with monitoring marine turbine arrays an obvious and potentially important application

Scoping the impact of tidal and wave energy extraction on suspended sediment concentrations and underwater light climate

The depth to which sunlight penetrates below the sea surface is one of the key factors determining the species composition and productivity of marine ecosystems. The effects range from the rate and fate of primary production, through the performance of visual predators such as fish, the potential for refuge from predators by migrating to depth, to the scope for seabed stabilisation by algal mats. Light penetration depends partly on spectral absorption by seawater and dissolved substances, but mainly on the scattering caused by suspended particulate material (SPM). Some of this SPM may be of biological origin, but in coastal waters the majority is mineral material originating ultimately from seabed disturbance and land erosion, the latter being deposited in the sea by rivers and aerial processes. SPM is maintained in the water column or deposited on the seabed depending on combinations of hydrodynamic processes including baroclinic (density-driven) or barotropic (mainly tidal and wind driven) currents, and wave action (Ward et al. 1984; Huettel et al. 1996). Since tidal and wave energy extraction must alter these hydrodynamic properties at some scales depending on the nature of the extraction process, we can expect some kind of impact on the concentration of the SPM. If these are large enough, we may have to consider the extent to which these may impact the underwater light environment and the local or regional ecology. Whilst several coupled hydrodynamic-sediment models exist to predict SPM distributions in aquatic systems, their skill level in open coastal and offshore marine waters is acknowledged to be relatively low. This is largely because the processes are not well understood and the formulations are largely based on empirical relationships rather than fundamental physical principles. The models are also highly demanding in terms of calibration data and computational resources. Hence their utility for predicting relatively subtle effects arising from changes in flow or wave environments due to energy extraction devices seems rather low. Here, we summarise the key mathematical functions describing the processes involved in sediment suspension, and propose a lightweight one-dimensional (vertical) model which can be used to scope the effects of changes in flow and wave energy on SPM

New primary production in northwest European shelf seas, 1960–2003

Spatial and temporal patterns from 1960 to 2003 in annual potential new primary production (PNP) of the NW European shelf seas were derived from general additive models of nitrate concentrations and from data on riverine and atmospheric fluxes of oxidized nitrogen. Average PNP was highest in the seasonally stratified outer shelf regions (>70 gC m-2 yr-1), where the proportion of PNP accounted for by vertical fluxes from deep water (>65%) was correlated with the North Atlantic Oscillation (NAO) index. PNP was lowest in the central North Sea (~30 gC m-2 yr-1) and in the southern North Sea was correlated with river inputs that accounted for 24% of the annual total (average ~50 gC m-2 yr-1). Atmospheric deposition accounted for ~3% of annual PNP region-wide, but in the northern North Sea this was higher than the contribution from rivers. Tidal fronts are traditionally considered to be highly productive zones, but we find them to have characteristically low PNP and conclude that they must be loci of high recycled production. The results indicate an exceptional flux of nitrate-rich ocean water onto the shelf in the early 1990s, which resulted in a pulse of PNP coincident with a well-documented 'regime shift' in the pelagic food web. North Sea-wide, long-term average PNP was approximately equal to production by all higher trophic levels combined, though trophic propagation of inter-annual variations was weakly defined. Nevertheless, there is a case for proposing that harvesting in areas and periods of low PNP should be managed more conservatively to minimize the risk of detrimental effects on the food web

Climate fluctuations and the spring invasion of the North Sea by Calanus finmarchicus

The population of Calanus finmarchicus in the North Sea is replenished each spring by invasion from an overwintering stock located beyond the shelf edge. A combincation of field observations, statistical analysis of Continuous Plankton Recorder (CPR) data, and particle tracking model simulations, was used to investigate the processes involved in the cross-shelf invasion. The results showed that the main source of overwintering animals entering the North Sea in the spring is at depths of greater than 600m in the Faroe Shetland Channel, where concentrations of up to 620m -3 are found in association with the overflow of Norwegian Sea Deep Water (NSDW) across the Iceland Scotland Ridge. The input of this water mass to the Faroe Shetland Channel, and hence the supply of overwintering C. finmarchicus, has declined since the late 1960s due to changes in convective processes in the Greenland Sea. Beginning in February, animals start to emerge from the overwintering state and migrate to the surface waters, where their transport into the North Sea is mainly determined by the incidence of north-westerly winds that have declined since the 1960s. Together, these two factors explain a high proportion of the 30-year trends in spring abundance in the North Sea as measured by the CPR survey. Both the regional winds and the NSDW overflow are connected to the North Atlantic Oscillation Index (NAO), which is an atmospheric climate index, but with different time scales of response. Thus, interannual fluctuations in the NAO can cause immediate changes in the incidence of north-westerly winds without leading to corresponding changes in C. finmarchicus abundance in the North Sea, because the NSDW overflow responds over longer (decadal) time scales

Winter distribution of Calanus finmarchicus in the Northeast Atlantic

Data from plankton sampling and Optical Plankton Counter deployments during six cruises between December of 1994 and 1999 have been used to derive a composite three-dimensional distribution of the abundance of Calanus finmarchicus during winter (December-January) in the Norwegian Sea and Northeast Atlantic. There are two centres of abundance, one in the eastern Norwegian Sea and Faroe-Shetland Channel, associated with the interface between Norwegian Sea Deep Water and Intermediate Water layers, and another in the Irminger Sea southwest of Iceland in association with Labrador Sea Water. In the open Northeast Atlantic, the concentration of wintering animals is around 30% of that in the Norwegian Sea and the vertical distribution ismore diffuse and on average deeper. Modelling studies have shown that the overwinter distribution and transport are key factors determining the spatial persistence of C. finmarchicus but, apart from the data presented here, there is little knowledge of these large-scale properties

Comparative analysis of Calanus finmarchicus demography at locations around the Northeast Atlantic

Standardized time-series sampling was carried out throughout 1997 at seven locations around the Northeast Atlantic to investigate regional variations in the seasonal demography of Calanus finmarchicus. Sites ranged from an inshore location in the North Sea, where C. finmarchicus formed only a small component of the zooplankton (2000 mgC m-2 during spring and summer). The internal consistency of the demographic time-series from each site was investigated by three partial models of life-cycle processes. In general, the demography of late copepodites could be accounted for by a relatively simple forecast model of stage development and diapause. However, there was a large discrepancy between nowcast estimates of egg production based on female abundance, temperature, and chlorophyll, and hindcast simulations of the egg production required to account for the observed abundance of early copepodite stages. The results point to a gap in our understanding of seasonal variations in rates of egg production and/or survival of nauplii. Overall, the population sampled at Weathership M appeared to be reasonably self-contained, but all other sites were reliant on invasion of overwintered stock in spring. At least two generations were observed at all but one site, but the extent to which these were generated by discrete bursts of egg production varied between sites and seemed to be partly dependent on the proximity to an overwintering location

Human axial progenitors generate trunk neural crest cells

The neural crest (NC) is a multipotent embryonic cell population generating distinct cell types in an axial position-dependent manner. The production of NC cells from human pluripotent stem cells (hPSCs) is a valuable approach to study human NC biology. However, the origin of human trunk NC remains undefined and therefore current in vitro differentiation strategies induce only a modest yield of trunk NC cells. Here we show that hPSC-derived axial progenitors, the posteriorly-located drivers of embryonic axis elongation, give rise to trunk NC cells and their derivatives. Moreover, we define the molecular signatures associated with the emergence of human NC cells of distinct axial identities in vitro. Collectively, our findings indicate that there are two routes toward a human post-cranial NC state: the birth of cardiac and vagal NC is facilitated by retinoic acid-induced posteriorisation of an anterior precursor whereas trunk NC arises within a pool of posterior axial progenitor