Induction Of Enzymes As A Mechanism For The Seasonal Control Of Metabolism In Marine-Invertebrates - Glucose-6-Phosphate Dehydrogenases From The Mantle And Hepatopancreas Of The Common Mussel Mytilus-Edulis-L

1. Glucose-6-phosphate dehydrogenase from the hepatopancreas and mantle tissue of M. edulis was investigated over two years for changes in specific activity (crude enzyme preparations) and the apparent Michaelis constants for G6P and NADP+ (highly purified enzyme preparations). 2. The specific activity of the mantle enzyme was low in summer and autumn and increased in the winter during the time of lipid deposition. In contrast, the specific activity of the hepatopancreas enzyme was high in summer and declined during the autumn and winter. 3. The apparent values for G6P and NADP+ of the mantle enzymechange little during a year. Changes were observed for the hepatopancreas enzyme during the first year but not the second.

Combined Effects Of Body Size Food Concentration And Season On Physiology Of Mytilus-Edulis

Multivariate experiments are used to study the effects of body size, food concentration, and season on the oxygen consumption, ammonia excretion, food assimilation efficiency and filtration rate of Mytilus edulis adults. Food concentrations and season affect both the intercept and the slope of the allometric equation describing oxygen uptake as a function of body size. Multiple regression and response surface techniques are used to describe and illustrate the complex relationship between metabolic rate, ration, season and the body size of M. edulis. Filtration rate has a relatively low weight exponent Q> = 038) and the intercept for the allometric equation is not significantly affected by food concentration, season or acclimation temperatures between 5 and 20 °C. Food assimilation efficiency declines exponentially with increasing food concentration and is dependent on body size at high food levels. The rate of ammonia excretion shows a similar seasonal cycle to that of oxygen consumption. They are both minimal in the autumn/winter and reach a maximum in the spring /summer.

Population-Dynamics Of Mytilicola-Intestinalis In Mytilus-Edulis In South West England

The population dynamics of Mytilicola intestinalis Steuer in mussels (Mytilus edulis L.) from the River Lynher, Cornwall, England, have been studied over 3 years. By transplanting uninfested mussels from the River Erme, South Devon, into the Lynher mussel bed, the study was extended to the growth and development of new infestations under natural conditions. Female Mytilicola intestinalis are shown to breed twice, and two generations of parasites coexist for most of the year, with recruitment taking place in summer and autumn. One generation contributes its first brood to the autumn recruits before overwintering and contributing its second brood to the following summer's recruits. The other generation overwinters as juvenile and immature stages to contribute its two broods successively to the summer and autumn recruits. Environmental temperatures are believed to control the rates of development at all stages rather than acting as triggers in the onset or cessation of breeding at specific times. There is no evidence to support the contention that heavily infested mussels are killed, and parasite mortality is shown to be density-independent.

Seasonal dynamics of the carbonate system in the Western English Channel

We present over 900 carbonate system observations collected over four years (2007–2010) in the Western English Channel (WEC). We determined CO2 partial pressure (pCO2), Total Alkalinity (TA) and Dissolved Inorganic Carbon (DIC) along a series of 40 km transects, including two oceanographic stations (L4 and E1) within a sustained coastal observatory. Our data follow a seasonal pattern of CO2 undersaturation from January to August, followed by supersaturation in September–October and a return to near-equilibrium thereafter. This pattern is explained by the interplay of thermal and biological sinks in winter and spring–summer, respectively, followed by the breakdown of stratification and mixing with deeper, high-CO2 water in autumn. The drawdown of DIC and inorganic N between March and June with a C:N ratio of 8.7–9.5 was consistent with carbon over-consumption during phytoplankton growth. Monthly mean surface pCO2 was strongly correlated with depth integrated chlorophyll a highlighting the importance of subsurface chlorophyll a maxima in controlling C-fluxes in shelf seas. Mixing of seawater with riverine freshwater in near-shore samples caused a reduction in TA and the saturation state of calcite minerals, particularly in winter. Our data show that the L4 and E1 oceanographic stations were small, net sinks for atmospheric CO2 over an annual cycle (−0.52±0.66 mol C m−2 y−1 and −0.62±0.49 mol C m−2 y−1, respectively).

DISCOVERY 2010: Spatial and temporal variability in a dynamic polar ecosystem

The Scotia Sea has been a focus of biological- and physical oceanographic study since the Discovery expeditions in the early 1900s. It is a physically energetic region with some of the highest levels of productivity in the Southern Ocean. It is also a region within which there have been greater than average levels of change in upper water column temperature. We describe the results of three cruises transecting the central Scotia Sea from south to north in consecutive years and covering spring, summer and autumn periods. We also report on some community level syntheses using both current-day and historical data from this region. A wide range of parameters were measured during the field campaigns, covering the physical oceanography of the region, air–sea CO2 fluxes, macro- and micronutrient concentrations, the composition and biomass of the nano-, micro- and mesoplankton communities, and the distribution and biomass of Antarctic krill and mesopelagic fish. Process studies examined the effect of iron-stress on the physiology of primary producers, reproduction and egestion in Antarctic krill and the transfer of stable isotopes between trophic layers, from primary consumers up to birds and seals. Community level syntheses included an examination of the biomass-spectra, food-web modelling, spatial analysis of multiple trophic layers and historical species distributions. The spatial analyses in particular identified two distinct community types: a northern warmer water community and a southern cold community, their boundary being broadly consistent with the position of the Southern Antarctic Circumpolar Current Front (SACCF). Temperature and ice cover appeared to be the dominant, over-riding factors in driving this pattern. Extensive phytoplankton blooms were a major feature of the surveys, and were persistent in areas such as South Georgia. In situ and bioassay measurements emphasised the important role of iron inputs as facilitators of these blooms. Based on seasonal DIC deficits, the South Georgia bloom was found to contain the strongest seasonal carbon uptake in the ice-free zone of the Southern Ocean. The surveys also encountered low-production, iron-limited regions, a situation more typical of the wider Southern Ocean. The response of primary and secondary consumers to spatial and temporal heterogeneity in production was complex. Many of the life-cycles of small pelagic organisms showed a close coupling to the seasonal cycle of food availability. For instance, Antarctic krill showed a dependence on early, non-ice-associated blooms to facilitate early reproduction. Strategies to buffer against environmental variability were also examined, such as the prevalence of multiyear life-cycles and variability in energy storage levels. Such traits were seen to influence the way in which Scotia Sea communities were structured, with biomass levels in the larger size classes being higher than in other ocean regions. Seasonal development also altered trophic function, with the trophic level of higher predators increasing through the course of the year as additional predator-prey interactions emerged in the lower trophic levels. Finally, our studies re-emphasised the role that the simple phytoplankton-krill-higher predator food chain plays in this Southern Ocean region, particularly south of the SACCF. To the north, alternative food chains, such as those involving copepods, macrozooplankton and mesopelagic fish, were increasingly important. Continued ocean warming in this region is likely to increase the prevalence of such alternative such food chains with Antarctic krill predicted to move southwards.

The dispersal of phytoplankton populations by enhanced turbulent mixing in a shallow coastal sea

A single tidal cycle survey in a Lagrangian reference frame was conducted in autumn 2010 to evaluate the impact of short-term, episodic and enhanced turbulent mixing on large chain-forming phytoplankton. Observations of turbulence using a free-falling microstructure profiler were undertaken, along with near-simultaneous profiles with an in-line digital holographic camera at station L4 (50° 15′ N 4° 13′ W, depth 50 m) in the Western English Channel. Profiles from each instrument were collected hourly whilst following a drogued drifter. Results from an ADCP attached to the drifter showed pronounced vertical shear, indicating that the water column structure consisted of two layers, restricting interpretation of the Lagrangian experiment to the upper ~ 25 m. Atmospheric conditions deteriorated during the mid-point of the survey, resulting in values of turbulent dissipation reaching a maximum of 10− 4 W kg− 1 toward the surface in the upper 10 m. Chain-forming phytoplankton > 200 μm were counted using the data from the holographic camera for the two periods, before and after the enhanced mixing event. As mixing increased phytoplankton underwent chain breakage, were dispersed by advection through their removal from the upper to lower layer and subjected to aggregation with other suspended material. Depth averaged counts of phytoplankton were reduced from a maximum of around 2050 L− 1 before the increased turbulence, to 1070 L− 1 after, with each of these mechanisms contributing to this reduction. These results demonstrate the sensitivity of phytoplantkon populations to moderate increases in turbulent activity, yielding consequences for accurate forecasting of the role played by phytoplankton in climate studies and also for the ecosystem in general in their role as primary producers.

Assessing the potential of autonomous submarine gliders for ecosystem monitoring across multiple trophic levels (plankton to cetaceans) in shallow shelf seas

A combination of scientific, economic, technological and policy drivers is behind a recent upsurge in the use of marine autonomous systems (and accompanying miniaturized sensors) for environmental mapping and monitoring. Increased spatial–temporal resolution and coverage of data, at reduced cost, is particularly vital for effective spatial management of highly dynamic and heterogeneous shelf environments. This proof-of-concept study involves integration of a novel combination of sensors onto buoyancy-driven submarine gliders, in order to assess their suitability for ecosystem monitoring in shelf waters at a variety of trophic levels. Two shallow-water Slocum gliders were equipped with CTD and fluorometer to measure physical properties and chlorophyll, respectively. One glider was also equipped with a single-frequency echosounder to collect information on zooplankton and fish distribution. The other glider carried a Passive Acoustic Monitoring system to detect and record cetacean vocalizations, and a passive sampler to detect chemical contaminants in the water column. The two gliders were deployed together off southwest UK in autumn 2013, and targeted a known tidal-mixing front west of the Isles of Scilly. The gliders’ mission took about 40 days, with each glider travelling distances of >1000 km and undertaking >2500 dives to depths of up to 100 m. Controlling glider flight and alignment of the two glider trajectories proved to be particularly challenging due to strong tidal flows. However, the gliders continued to collect data in poor weather when an accompanying research vessel was unable to operate. In addition, all glider sensors generated useful data, with particularly interesting initial results relating to subsurface chlorophyll maxima and numerous fish/cetacean detections within the water column. The broader implications of this study for marine ecosystem monitoring with submarine gliders are discussed.

Ocean Net Heat Flux Influences Seasonal to Interannual Patterns of Plankton Abundance

Changes in the net heat flux (NHF) into the ocean have profound impacts on global climate. We analyse a long-term plankton time-series and show that the NHF is a critical indicator of ecosystem dynamics. We show that phytoplankton abundance and diversity patterns are tightly bounded by the switches between negative and positive NHF over an annual cycle. Zooplankton increase before the transition to positive NHF in the spring but are constrained by the negative NHF switch in autumn. By contrast bacterial diversity is decoupled from either NHF switch, but is inversely correlated (r=-0.920) with the magnitude of the NHF. We show that the NHF is a robust mechanistic tool for predicting climate change indicators such as spring phytoplankton bloom timing and length of the growing season.

Assessing the potential of autonomous submarine gliders for ecosystem monitoring across multiple trophic levels (plankton to cetaceans) in shallow shelf seas

A combination of scientific, economic, technological and policy drivers is behind a recent upsurge in the use of marine autonomous systems (and accompanying miniaturized sensors) for environmental mapping and monitoring. Increased spatial–temporal resolution and coverage of data, at reduced cost, is particularly vital for effective spatial management of highly dynamic and heterogeneous shelf environments. This proof-of-concept study involves integration of a novel combination of sensors onto buoyancy-driven submarine gliders, in order to assess their suitability for ecosystem monitoring in shelf waters at a variety of trophic levels. Two shallow-water Slocum gliders were equipped with CTD and fluorometer to measure physical properties and chlorophyll, respectively. One glider was also equipped with a single-frequency echosounder to collect information on zooplankton and fish distribution. The other glider carried a Passive Acoustic Monitoring system to detect and record cetacean vocalizations, and a passive sampler to detect chemical contaminants in the water column. The two gliders were deployed together off southwest UK in autumn 2013, and targeted a known tidal-mixing front west of the Isles of Scilly. The gliders’ mission took about 40 days, with each glider travelling distances of >1000 km and undertaking >2500 dives to depths of up to 100 m. Controlling glider flight and alignment of the two glider trajectories proved to be particularly challenging due to strong tidal flows. However, the gliders continued to collect data in poor weather when an accompanying research vessel was unable to operate. In addition, all glider sensors generated useful data, with particularly interesting initial results relating to subsurface chlorophyll maxima and numerous fish/cetacean detections within the water column. The broader implications of this study for marine ecosystem monitoring with submarine gliders are discussed.

Spatio-temporal variability in ammonia oxidation and ammonia oxidising bacteria and archaea in coastal sediments of the Western English Channel

The abundance of ammonia-oxidising bacterial (AOB) and ammonia-oxidising archaeal (AOA) (amoA) genes and ammonia oxidation rates were compared bimonthly from July 2008 to May 2011 in 4 contrasting coastal sediments in the western English Channel. Despite a higher abundance of AOA amoA genes within all sediments and at all time-points, rates of ammonia oxidation correlated with AOB and not AOA amoA gene abundance. Sediment type was a major factor in determining both AOB amoA gene abundance and AOB community structure, possibly due to deeper oxygen penetration into the sandier sediments, increasing the area available for ammonia oxidation. Decreases in AOB amoA gene abundance were evident during summer and autumn, with maximum abundance and ammonia oxidation rates occurring in winter and early spring. PCR-DGGE of AOB amoA genes indicated that no seasonal changes to community composition occurred; however, a gradual movement in community composition occurred at 3 of the sites studied. The lack of correlation between AOA amoA gene abundance and ammonium oxidation rates, or any other environmental variable measured, may be related to the higher spatial variation amongst measurements, obscuring temporal trends, or the bimonthly sampling, which may have been too infrequent to capture temporal variability in the deposition of fresh organic matter. Alternatively, AOA may respond to changing substrate concentrations by an increase or decrease in transcript rather than gene abundance.

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.

Annual study of oxygenated volatile organic compounds in UK shelf waters

We performed an annual study of oxygenated volatile organic compound (OVOC) seawater concentrations at a site off Plymouth, UK in the Western English Channel over the period of February 2011–March 2012. Acetone concentrations ranged from 2–10 nM (nanomole/L) in surface waters with a maximum observed in summer. Concentrations correlated positively with net shortwave radiation and UV light, suggestive of photochemically linked acetone production. We observed a clear decline in acetone concentrations below the mixed layer. Acetaldehyde varied between 4–37 nM in surface waters with higher values observed in autumn and winter. Surface concentrations of methanol ranged from 16–78 nM, but no clear annual cycle was observed. Methanol concentrations exhibited considerable inter-annual variability. We estimate consistent deposition to the sea surface for acetone and methanol but that the direction of the acetaldehyde flux varies during the year.

Nanoplankton and picoplankton in the Western English Channel: abundance and seasonality from 2007–2013

The nano- and picoplankton community at Station L4 in the Western English Channel was studied between 2007 and 2013 by flow cytometry to quantify abundance and investigate seasonal cycles within these communities. Nanoplankton included both photosynthetic and heterotrophic eukaryotic single-celled organisms while the picoplankton included picoeukaryote phytoplankton, Synechococcus sp. cyanobacteria and heterotrophic bacteria. A Box–Jenkins Transfer Function climatology analysis of surface data revealed that Synechococcus sp., cryptophytes, and heterotrophic flagellates had bimodal annual cycles. Nanoeukaryotes and both high and low nucleic acid-containing bacteria (HNA and LNA, respectively) groups exhibited unimodal annual cycles. Phaeocystis sp., whilst having clearly defined abundance maxima in spring was not detectable the rest of the year. Coccolithophores exhibited a weak seasonal cycle, with abundance peaks in spring and autumn. Picoeukaryotes did not exhibit a discernable seasonal cycle at the surface. Timings of maximum group abundance varied through the year. Phaeocystis sp. and heterotrophic flagellates peaked in April/May. Nanoeukaryotes and HNA bacteria peaked in June/July and had relatively high abundance throughout the summer. Synechococcus sp., cryptophytes and LNA bacteria all peaked from mid to late September. The transfer function model techniques used represent a useful means of identifying repeating annual cycles in time series data with the added ability to detect trends and harmonic terms at different time scales from months to decades.

Feeding selectivity of bivalve larvae on natural plankton assemblages in the Western English Channel

Meroplankton, including bivalve larvae, are an important and yet understudied component of coastal marine food webs. Understanding the baseline of meroplankton ecology is imperative to establish and predict their sensitivity to local and global marine stressors. Over an annual cycle (October 2009–September 2010), bivalve larvae were collected from the Western Channel Observatory time series station L4 (50°15.00′N, 4°13.02′W). The morphologically similar larvae were identified by analysis of the 18S nuclear small subunit ribosomal RNA gene, and a series of incubation experiments were conducted to determine larval ingestion rates on natural plankton assemblages. Complementary gut content analysis was performed using a PCR-based method for detecting prey DNA both from field-collected larvae and those from the feeding experiments. Molecular identification of bivalve larvae showed the community composition to change over the course of the sampling period with domination by Phaxas in winter and higher diversity in autumn. The larvae selected for nanoeukaryotes (2–20 µm) including coccolithophores (<20 µm) which together comprised >75 % of the bivalve larvae diet. Additionally, a small percentage of carbon ingested originated from heterotrophic ciliates (<30 µm). The molecular analysis of bivalve larvae gut content provided increased resolution of identification of prey consumed and demonstrated that the composition of prey consumed established through bottle incubations conferred with that established from in situ larvae. Despite changes in bivalve larvae community structure, clearance rates of each prey type did not change significantly over the course of the experiment, suggesting different bivalve larvae species may consume similar prey.

How does Calanus helgolandicus maintain its population in a variable environment? Analysis of a 25-year time series from the English Channel

Calanus helgolandicus is a key copepod of the NE Atlantic and fringing shelves, with a distribution that is expanding northwards with oceanic warming. The Plymouth L4 site has warmed over the past 25-years, and experiences large variations in the timing and availability of food for C. helgolandicus. Here we examine the degree to which these changes translate into variation in reproductive output and subsequently C. helgolandicus population size. Egg production rates (eggs female−1 day−1) were maximal in the spring to early-summer period of diatom blooms and high ciliate abundance, rather than during the equally large autumn blooms of autotrophic dinoflagellates. Egg hatch success was lower in spring however, with a greater proportion of naupliar deformities then also. Both the timing and the mean summer abundance of C. helgolandicus (CI–CVI) reflected those of spring total reproductive output. However this relationship was driven by inter-annual variability in female abundance and not that of egg production per female, which ranged only two-fold. Winter abundance of C. helgolandicus at L4 was much more variable than abundance in other seasons, and reflected conditions from the previous growing season. However, these low winter abundances had no clear carry-over signal to the following season’s population size. Overall, the C. helgolandicus population appears to be surprisingly resilient at this dynamic, inshore site, showing no long-term phenology shift and only a four-fold variation in mean abundance between years. This dampening effect may reflect a series of mortality sources, associated with the timing of stratification in the early part of the season, likely affecting egg sinking and loss, plus intense, density-dependent mortality of early stages in mid-summer likely through predation.