Microbial spatial variability: An example from the Celtic Sea

In July 2004, dominant populations of microbial ultraplankton (<5 μm), in the surface of the Celtic Sea (between UK and Eire), were repeatedly mapped using flow cytometry, at 1.5 km resolution over a region of diameter 100 km. The numerically dominant representatives of all basic functional types were enumerated including one group of phototrophic bacteria (Syn), two groups of phytoplankton (PP, NP), three groups of heterotrophic bacterioplankton (HB) and the regionally dominant group of heterotrophic protists (HP). The distributions of all organisms showed strong spatial variability with little relation to variability in physical fields such as salinity and temperature. Furthermore, there was little agreement between distributions of different organisms. The only linear correlation consistently explaining more than 50% of the variance between any pairing of the organism groups enumerated is between two different groups of HB. Specifically, no linear, or non-linear, relationship is found between any pairings of SYB, PP or HB groups with their protist predators HP. Looking for multiple dependencies, factor analysis reveals three groupings: Syn, PP and low nucleic acid content HB (LNA); high nucleic acid content HB (HNA); HP and NP. Even the manner in which the spatial variability of Syn, PP and HB abundance varies as a function of lengthscale (represented by a semivariogram) differs significantly from that for HP. In summary, although all microbial planktonic groups enumerated are present and numerically dominant throughout the region studied, at face value the relationships between them seem weak. Nevertheless, the behaviour of a simple, illustrative ecological model, with strongly interacting phototrophs and heterotrophs, with stochastic forcing, is shown to be consistent with the observed poor correlations and differences in how spatial variability varies with lengthscale. Thus, our study suggests that a comparison of microbial abundances alone may not discern strong underlying trophic interactions. Specific knowledge of these processes, in particular grazing, will be required to explain the causes of the observed microbial spatial variability and its resulting consequences for the functioning of the ecosystem.

Tropical Marginal Seas: Priority Regions for Managing Marine Biodiversity and Ecosystem Function

Tropical marginal seas (TMSs) are natural subregions of tropical oceans containing biodiverse ecosystems with conspicuous, valued, and vulnerable biodiversity assets. They are focal points for global marine conservation because they occur in regions where human populations are rapidly expanding. Our review of 11 TMSs focuses on three key ecosystems—coral reefs and emergent atolls, deep benthic systems, and pelagic biomes—and synthesizes, illustrates, and contrasts knowledge of biodiversity, ecosystem function, interaction between adjacent habitats, and anthropogenic pressures. TMSs vary in the extent that they have been subject to human influence—from the nearly pristine Coral Sea to the heavily exploited South China and Caribbean Seas—but we predict that they will all be similarly complex to manage because most span multiple national jurisdictions. We conclude that developing a structured process to identify ecologically and biologically significant areas that uses a set of globally agreed criteria is a tractable first step toward effective multinational and transboundary ecosystem management of TMSs.

Ecosystem function and services provided by the deep sea

The deep sea is often viewed as a vast, dark, remote, and inhospitable environment, yet the deep ocean and seafloor are crucial to our lives through the services that they provide. Our understanding of how the deep sea functions remains limited, but when treated synoptically, a diversity of supporting, provisioning, regulating and cultural services becomes apparent. The biological pump transports carbon from the atmosphere into deep-ocean water masses that are separated over prolonged periods, reducing the impact of anthropogenic carbon release. Microbial oxidation of methane keeps another potent greenhouse gas out of the atmosphere while trapping carbon in authigenic carbonates. Nutrient regeneration by all faunal size classes provides the elements necessary for fueling surface productivity and fisheries, and microbial processes detoxify a diversity of compounds. Each of these processes occur on a very small scale, yet considering the vast area over which they occur they become important for the global functioning of the ocean. The deep sea also provides a wealth of resources, including fish stocks, enormous bioprospecting potential, and elements and energy reserves that are currently being extracted and will be increasingly important in the near future. Society benefits from the intrigue and mystery, the strange life forms, and the great unknown that has acted as a muse for inspiration and imagination since near the beginning of civilization. While many functions occur on the scale of microns to meters and timescales up to years, the derived services that result are only useful after centuries of integrated activity. This vast dark habitat, which covers the majority of the globe, harbour processes that directly impact humans in a variety of ways; however, the same traits that differentiate it from terrestrial or shallow marine systems also result in a greater need for integrated spatial and temporal understanding as it experiences increased use by society. In this manuscript we aim to provide a foundation for informed conservation and management of the deep sea by summarizing the important role of the deep sea in society.

Marine biodiversity and ecosystem function relationships:the potential for practical monitoring applications

There is an increasing demand for environmental assessments of the marine environment to include ecosystem function. However, existing schemes are predominantly based on taxonomic (i.e. structural) measures of biodiversity. Biodiversity and Ecosystem Function (BEF) relationships are suggested to provide a mechanism for converting taxonomic information into surrogates of ecosystem function. This review assesses the evidence for marine BEF relationships and their potential to be used in practical monitoring applications (i.e. operationalized). Five key requirements were identified for the practical application of BEF relationships: (1) a complete understanding of strength, direction and prevalence of marine BEF relationships, (2) an understanding of which biological components are influential within specific BEF relationships, (3) the biodiversity of the selected biological components can be measured easily, (4) the ecological mechanisms that are the most important for generating marine BEF relationships, i.e. identity effects or complementarity, are known and (5) the proportion of the overall functional variance is explained by biodiversity, and hence BEF relationships, has been established. Numerous positive and some negative BEF relationships were found within the literature, although many reproduced poorly the natural species richness, trophic structures or multiple functions of real ecosystems (requirement 1). Null relationships were also reported. The consistency of the positive and negative relationships was often low that compromised the ability to generalize BEF relationships and confident application of BEF within marine monitoring. Equally, some biological components and functions have received little or no investigation. Expert judgement was used to attribute biological components using spatial extent, presence and functional rate criteria (requirement 2). This approach highlighted the main biological components contributing the most to specific ecosystem functions, and that many of the particularly influential components were found to have received the least amount of research attention. The need for biodiversity to be measureable (requirement 3) is possible for most biological components although difficult within the functionally important microbes. Identity effects underpinned most marine BEF relationships (requirement 4). As such, processes that translated structural biodiversity measures into functional diversity were found to generate better BEF relationships. The analysis of the contribution made by biodiversity, over abiotic influences, to the total expression of a particular ecosystem function was rarely measured or considered (requirement 5). Hence it is not possible to determine the overall importance of BEF relationships within the total ecosystem functioning observed. In the few studies where abiotic factors had been considered, it was clear that these modified BEF relationships and have their own direct influence on functional rate. Based on the five requirements, the information required for immediate ‘operationalization’ of BEF relationships within marine functional monitoring is lacking. However, the concept of BEF inclusion within practical monitoring applications, supported by ecological modelling, shows promise for providing surrogate indicators of functioning.

The assessment of a global marine ecosystem model on the basis of emergent properties and ecosystem function: a case study with ERSEM

Ecosystem models are often assessed using quantitative metrics of absolute ecosystem state, but these model-data comparisons are disproportionately vulnerable to discrepancies in the location of important circulation features. An alternative method is to demonstrate the models capacity to represent ecosystem function; the emergence of a coherent natural relationship in a simulation indicates that the model may have an appropriate representation of the ecosystem functions that lead to the emergent relationship. Furthermore, as emergent properties are large-scale properties of the system, model validation with emergent properties is possible even when there is very little or no appropriate data for the region under study, or when the hydrodynamic component of the model differs significantly from that observed in nature at the same location and time. A selection of published meta-analyses are used to establish the validity of a complex marine ecosystem model and to demonstrate the power of validation with emergent properties. These relationships include the phytoplankton community structure, the ratio of carbon to chlorophyll in phytoplankton and particulate organic matter, the ratio of particulate organic carbon to particulate organic nitrogen and the stoichiometric balance of the ecosystem. These metrics can also inform aspects of the marine ecosystem model not available from traditional quantitative and qualitative methods. For instance, these emergent properties can be used to validate the design decisions of the model, such as the range of phytoplankton functional types and their behaviour, the stoichiometric flexibility with regards to each nutrient, and the choice of fixed or variable carbon to nitrogen ratios.

The Impact of Polystyrene Microplastics on Feeding, Function and Fecundity in the Marine CopepodCalanus helgolandicus

Microscopic plastic debris, termed “microplastics”, are of increasing environmental concern. Recent studies have demonstrated that a range of zooplankton, including copepods, can ingest microplastics. Copepods are a globally abundant class of zooplankton that form a key trophic link between primary producers and higher trophic marine organisms. Here we demonstrate that ingestion of microplastics can significantly alter the feeding capacity of the pelagic copepod Calanus helgolandicus. Exposed to 20 μm polystyrene beads (75 microplastics mL–1) and cultured algae ([250 μg C L–1) for 24 h, C. helgolandicus ingested 11% fewer algal cells (P = 0.33) and 40% less carbon biomass (P < 0.01). There was a net downward shift in the mean size of algal prey consumed (P < 0.001), with a 3.6 fold increase in ingestion rate for the smallest size class of algal prey (11.6–12.6 μm), suggestive of postcapture or postingestion rejection. Prolonged exposure to polystyrene microplastics significantly decreased reproductive output, but there were no significant differences in egg production rates, respiration or survival. We constructed a conceptual energetic (carbon) budget showing that microplastic-exposed copepods suffer energetic depletion over time. We conclude that microplastics impede feeding in copepods, which over time could lead to sustained reductions in ingested carbon biomass.

An unusually large phytoplankton spring bloom drives rapid changes in benthic diversity and ecosystem function

In 2012, the Western English Channel experienced an unusually large and long-lived phytoplankton spring bloom. When compared with data from the past 20 years, average phytoplankton biomass at Station L4 (part of the Western Channel Observatory) was approximately 3× greater and lasted 50% longer than any previous year. Regular (mostly weekly) box core samples were collected from this site before, during and after the bloom to determine its impact on macrofaunal abundance, diversity, biomass, community structure and function. The spring bloom of 2012 was shown to support a large and rapid response in the majority of benthic taxa and functional groups. However, key differences in the precise nature of this response, as well as in its timing, was observed between different macrofauna feeding groups. Deposit feeders responded almost instantly at the start of the bloom, primarily thorough an increase in abundance. Suspension feeders and opportunistic/predatory/carnivorous taxa responded slightly more slowly and primarily with an increase in biomass. At the end of the bloom a rapid decline in macrobenthic abundance, diversity and biomass closely followed the decline in phytoplankton biomass. With suspension feeders showing evidence of this decline a few weeks before deposit feeders, it was concluded that this collapse in benthic communities was driven primarily by food availability and competition. However, it is possible that environmental hypoxia and the presence of toxic benthic cyanobacteria could also have contributed to this decline. This study shows evidence for strong benthic–pelagic coupling at L4; a shallow (50 m), coastal, fine-sand habitat. It also demonstrates that in such habitats, it is not just planktonic organisms that demonstrate clear community phenology. Different functional groups within the benthic assemblage will respond to the spring bloom in specific manner, with implications for key ecosystem functions and processes, such as secondary production and bioturbation. Only by taking integrated benthic and pelagic observations over such fine temporal scales (weekly) was the current study able to identify the intimate structure of the benthic response. Similar studies from other habitats and under different bloom conditions are urgently needed to fully appreciate the strength of benthic–pelagic coupling in shallow coastal environments.

Coastal upwelling and downwelling forcing of circulation in a semi-enclosed bay: Ria de Vigo

Semi-enclosed bays in upwelling regions are exposed to forcing related to winds, currents and buoyancy over the shelf. The influence of this external forcing is moderated by factors such as connectivity to the open ocean, shelter by surrounding topography, dimensions of the bay, and freshwater outflows. Such bays, preferred locations for ports, mariculture, marine industry, recreational activities and coastal settlement, present a range of characteristics, understanding of which is necessary to their rational management. Observations in such a semi-enclosed bay, the Ria de Vigo in Spain, are used to characterize the influence of upwelling and downwelling pulses on its circulation. In this location, near the northern limit of the Iberian upwelling system, upwelling events dominate during a short summer season and downwelling events the rest of the year. The ria response to the external forcing is central to nutrient supply and resultant plankton productivity that supports its high level of cultured mussel production. Intensive field studies in September 2006 and June 2007 captured a downwelling event and an upwelling event, respectively. Data from eight current profiler moorings and boat-based MiniBat/ADCP surveys provided an unprecedented quasi-synoptic view of the distribution of water masses and circulation patterns in any ria. In the outer ria, circulation was dominated by the introduction of wind-driven alongshore flow from the external continental shelf through the ria entrances and its interaction with the topography. In the middle ria, circulation was primarily related to the upwelling/downwelling cycle, with a cool, salty and dense lower layer penetrating to the inner ria during upwelling over the shelf. A warmer, lower salinity and less dense surface layer of coastal waters flowed inward during downwelling. Without external forcing, the inner ria responded primarily to tides and buoyancy changes related to land runoff. Under both upwelling and downwelling conditions, the flushing of the ria involved shelf responses to wind pulses. Their persistence for a few days was sufficient to allow waters from the continental shelf to penetrate the innermost ria. Longer term observations supported by numerical modeling are required to confirm the generality of such flushing events in the ria and determine their typical frequency, while comparative studies should explore how these scenarios fit into the range of conditions experienced in other semi-enclosed bays.