Mesozooplankton Grazing Research Articles

Algal Hot Spots in a Changing Arctic Ocean: Sea-Ice Ridges and the Snow-Ice Interface

During the N-ICE2015 drift expedition north-west of Svalbard, we observed the establishment and development of algal communities in first-year ice (FYI) ridges and at the snow-ice interface. Despite some indications of being hot spots for biological activity, ridges are under-studied largely because they are complex structures that are difficult to sample. Snow infiltration communities can grow at the snow-ice interface when flooded. They have been commonly observed in the Antarctic, but rarely in the Arctic, where flooding is less common mainly due to a lower snow-to-ice thickness ratio. Combining biomass measurements and algal community analysis with under-ice irradiance and current measurements as well as light modeling, we comprehensively describe these two algal habitats in an Arctic pack ice environment. High biomass accumulation in ridges was facilitated by complex surfaces for algal deposition and attachment, increased light availability, and protection against strong under-ice currents. Notably, specific locations within the ridges were found to host distinct ice algal communities. The pennate diatoms Nitzschia frigida and Navicula species dominated the underside and inclined walls of submerged ice blocks, while the centric diatom Shionodiscus bioculatus dominated the top surfaces of the submerged ice blocks. Higher light levels than those in and below the sea ice, low mesozooplankton grazing, and physical concentration likely contributed to the high algal biomass at the snow-ice interface. These snow infiltration communities were dominated by Phaeocystis pouchetii and chain-forming pelagic diatoms (Fragilariopsis oceanica and Chaetoceros gelidus). Ridges are likely to form more frequently in a thinner and more dynamic ice pack, while the predicted increase in Arctic precipitation in some regions in combination with the thinning Arctic icescape might lead to larger areas of sea ice with negative freeboard and subsequent flooding during the melt season. Therefore, these two habitats are likely to become increasingly important in the new Arctic with implications for carbon export and transfer in the ice-associated ecosystem.

The fatty acid content of plankton is changing in subtropical coastal waters as a result of OA: Results from a mesocosm study

Ocean Acidification (OA) effects on marine plankton are most often considered in terms of inorganic carbon chemistry, but decreasing pH may influence other aspects of cellular metabolism. Here we present the effects of OA on the fatty acid (FA) content and composition of an artificial phytoplankton community (Phaeodactylum tricornutum, Thalassiosira weissflogii, and Emiliania huxleyi) in a fully replicated, ∼4 m3 mesocosm study in subtropical coastal waters (Wuyuan Bay, China, 24.52°N, 117.18°E) at present day (400 μatm) and elevated (1000 μatm) pCO2 concentrations. Phytoplankton growth occurred in three phases during the 33-day experiment: an initial exponential growth leading to senescence and a subsequent decline phase. Phytoplankton sampled from these mesocosms were fed to mesozooplankton collected by net haul from Wuyuan Bay. Concentrations of saturated fatty acids (SFA) in both phytoplankton and mesozooplankton remained high under acidified and non-acidified conditions. However, polyunsaturated fatty acids (PUFA) and monounsaturated fatty acids (MUFA) increased significantly more under elevated pCO2 during the late exponential phase (Day 13), indicating increased nutritional value for zooplankton and higher trophic levels. Indeed, uptake rates of the essential FA docosahexaenoic acid (C20:5n3, DHA) increased in mesozooplankton under acidified conditions. However, mesozooplankton grazing rates decreased overall with elevated pCO2. Our findings show that these selected phytoplankton species have a relatively high tolerance to acidification in terms of FA production, and local mesozooplankton in these subtropical coastal waters can maintain their FA composition under end of century ocean acidification conditions.

Zooplankton grazing and egestion shifts particle size distribution in natural communities

MEPS Marine Ecology Progress Series Contact the journal Facebook Twitter RSS Mailing List Subscribe to our mailing list via Mailchimp HomeLatest VolumeAbout the JournalEditorsTheme Sections MEPS 575:43-56 (2017) - DOI: https://doi.org/10.3354/meps12212 Zooplankton grazing and egestion shifts particle size distribution in natural communities Karen Stamieszkin1,*, Nicole J. Poulton2, Andrew J. Pershing3 1Darling Marine Center, 193 Clark’s Cove Road, Walpole, Maine 04573, USA 2Bigelow Laboratory for Ocean Sciences, 60 Bigelow Drive, East Boothbay, Maine 04544, USA 3Gulf of Maine Research Institute, 350 Commercial Street, Portland, Maine 04101, USA *Corresponding author: karen.stamieszkin@maine.edu ABSTRACT: Marine plankton communities can be viewed in terms of their size structure rather than their taxonomic composition, revealing how allometric relationships affect the functioning of the community. Oceanic particle size spectra can be used to explain and predict variability in carbon export efficiency, because large particles generally sink faster than small particles. Since plankton trophic interactions impact particle size in the surface ocean, this size-structured view is a useful simplification for connecting plankton ecology with biogeochemistry. We conducted a series of grazing experiments to test the hypothesis that mesozooplankton shift particle size spectra toward larger particles, in a predictable manner that reflects their community size structure, through grazing and egestion of fecal pellets. These experiments were carried out over several months, and used natural communities of mesozooplankton and their microplankton prey collected in the coastal Gulf of Maine. After incubation, we analyzed the samples to determine size distribution and taxonomic information. Our results show that mesozooplankton grazing impacts microplankton in proportion to their abundance. Size relationships between plankton predators and prey that have been established at the individual level do not linearly translate to the community level. Further, while grazing itself does not significantly alter the particle size spectrum, repackaging of prey into mesozooplankton fecal pellets shifts particle size spectra toward larger particles. KEY WORDS: Plankton grazing · Egestion · Particle size distribution · Imaging cytometry · Carbon flux Full text in pdf format PreviousNextCite this article as: Stamieszkin K, Poulton NJ, Pershing AJ (2017) Zooplankton grazing and egestion shifts particle size distribution in natural communities. Mar Ecol Prog Ser 575:43-56. https://doi.org/10.3354/meps12212 Export citation RSS - Facebook - Tweet - linkedIn Cited by Published in MEPS Vol. 575. Online publication date: July 20, 2017 Print ISSN: 0171-8630; Online ISSN: 1616-1599 Copyright © 2017 Inter-Research.

Microzooplankton grazing rate in Ensenada of La Paz, B.C.S., Mexico

Phytoplankton growth and microzooplankton grazing rates in situ, were studied in a fixed sampling station from October 2007 to August 2008, at the Ensenada de La Paz, Mexico, using the dilution method. The apparent growth rate of phytoplankton was higher in December (1.33 d-1) and lower in October (0.35 d-1). The grazing rate was highest in April (0.86 d-1) and minimum in February and August (0.38 d-1). Grazing pressure, measured as the percentage of potential production removed (% PPR) varied between 28 and 76%. The relationship between phytoplankton growth and grazing rate indicates that chlorophyll-a production is consumed by approximately 50%, involving other processes that cause mortality of phytoplankton, as sedimentation rate and mesozooplankton grazing. The finding of moderate grazing in autumn and a clear growth and consumption in the winter are features that require monitoring in larger temporal and spatial scales.

Meso‐ and microzooplankton grazing in the Amazon River plume and western tropical North Atlantic

AbstractLargely due to size differences, mesozooplankton are important exporters of carbon and prey for larger organisms, while microzooplankton are important recyclers of nutrients, dominant grazers of phytoplankton, and a key link in the microbial loop. We investigated the relative importance of meso‐ and microzooplankton grazing in the western tropical North Atlantic Ocean (WTNA) and Amazon River plume. Sampling as part of the ANACONDAS project occurred in spring (May–June) 2010 during the peak outflow of the Amazon River and in fall (September–October) 2011 during the plume seasonal retroflection. Mesozooplankton grazing rates decreased with increasing salinity in both seasons, but during the fall both day and nighttime grazing rates were significantly negatively correlated with salinity. Mesozooplankton grazing was highest in plume‐influenced surface waters (0–25 m), and usually dominated by smaller size classes (0.2–0.5 mm and 0.5–1.0 mm). Microzooplankton grazing accounted for approximately 68% of bulk phytoplankton growth across all stations. Comparison of meso‐ and microzooplankton grazing suggests a transition in food web dynamics from a mesozooplankton dominated “export” structure in the plume transitioning to a microzooplankton dominated “retention” structure at mesohaline and oceanic stations above sea surface salinity of 33. Comparison between the seasons suggests a seasonal planktonic succession of low mesozooplankton grazing during the spring peak discharge followed by higher grazing rates and impact by mesozooplankton during the fall retroflection. These results provide important baseline information required for examining effects of climate change on the planktonic food web of the WTNA and for use in biogeochemical models of the region.

Mesozooplankton production, grazing and respiration in the Bay of Bengal: Implications for net heterotrophy

Mesozooplankton samples were collected from the mixed layer along a central (along 88°E) and a western transect in the Bay of Bengal during four seasons covered between 2001 and 2006 in order to investigate spatio-temporal variability in their biomass. At these stations, grazing and respiration rates were measured from live zooplankton hauled in from the surface during December 2005. Akin to the mesozooplankton “paradox” in the central and eastern Arabian Sea, biomass in the mixed layer was more or less invariant in the central and western Bay of Bengal, even as the chl a showed marginal temporal variation. By empirical equation, the mesozooplankton production rate calculated to be 70–246mgCm−2d−1 is on par with the Arabian Sea. Contrary to the conventional belief, mesozooplankton grazing impact was up to 83% on primary production (PP). Low PP coupled with very high zooplankton production (70% of PP) along with abundant bacterial production (50% of the PP; Ramaiah et al., 2009) is likely to render the Bay of Bengal net heterotrophic, especially during the spring intermonsoon. Greater estimates of fecal pellet-carbon egestion by mesozooplankton compared to the average particulate organic carbon flux in sediment traps, implies that much of the matter is recycled by heterotrophic communities in the mixed layer facilitating nutrient regeneration for phytoplankton growth. We also calculated that over a third of the primary production is channelized for basin-wide zooplankton respiration that accounts for ~52MtC annually. In the current scenario of global warming, if low (primary) productive warm pools like the Bay of Bengal continue to be net heterotrophic, negative implications like enhanced emission of CO2 to the atmosphere, increased particulate flux to the deeper waters and greater utilization of dissolved oxygen resulting in expansion of the existing oxygen minimum zone are imminent.

Mesozooplankton grazing during spring sea-ice conditions in the eastern Bering Sea

Mesozooplankton (copepods and euphausiids) grazing rates and prey preferences were determined during a series of three research cruises to the eastern Bering Sea in spring 2008, 2009, and 2010. Chlorophyll was dominated by large cells (>5µm), especially at bloom locations where they usually comprised greater than 90% of the total chlorophyll biomass. The relative importance of microzooplankton to the prey field biomass decreased with increasing chlorophyll concentration, and was less than 10% of the total prey biomass in ice-edge bloom regions. Overall, microzooplankton was the preferred prey of the mesozooplankton, although phytoplankton/ice algae were the dominant component of the diet because of their much greater biomass, especially during blooms. There were differences between mesozooplankton species in their prey preferences: Metridia pacifica, Pseudocalanus spp. and Calanus spp. had the strongest preference for microzooplankton prey, while euphausiids (Thysanoessa spp.) and Neocalanus flemingeri/plumchrus appeared to feed non-selectively on all prey items. Mesozooplankton exhibited a saturating feeding response to chlorophyll concentration (Holling׳s type II) that could be modeled by Michaelis–Menten equations. Taxa-specific maximum ingestion rates generally followed allometric theory, with smaller zooplankton having higher feeding rates than larger zooplankton, and ranged from about 4–30% body carbon day−1. Trophic cascades during grazing experiments could result in a substantial underestimate of chlorophyll ingestion rates, especially for those taxa that had a strong preference for microzooplankton. Grazing impacts by mesozooplankton on the integrated chlorophyll biomass and primary production were 2.7±4.4 and 26±48% day−1, respectively. Impacts increased significantly with increasing mesozooplankton biomass, which increased from early to late spring. However, grazing impacts were extremely low in ice-edge bloom regions. Our findings suggest that even when grazing by microzooplankton is included in our grazing impact estimates, about 50% of the primary production in phytoplankton blooms during spring on the eastern Bering Sea shelf is not grazed and is available for direct export to the benthic community.

Using Lagrangian‐based process studies to test satellite algorithms of vertical carbon flux in the eastern North Pacific Ocean

AbstractThe biological carbon pump is responsible for the transport of ∼5–20 Pg C yr−1 from the surface into the deep ocean but its variability is poorly understood due to an incomplete mechanistic understanding of the complex underlying planktonic processes. In fact, algorithms designed to estimate carbon export from satellite products incorporate fundamentally different assumptions about the relationships between plankton biomass, productivity, and export efficiency. To test the alternate formulations of export efficiency in remote‐sensing algorithms formulated by Dunne et al. (2005), Laws et al. (2011), Henson et al. (2011), and Siegel et al. (2014), we have compiled in situ measurements (temperature, chlorophyll, primary production, phytoplankton biomass and size structure, grazing rates, net chlorophyll change, and carbon export) made during Lagrangian process studies on seven cruises in the California Current Ecosystem and Costa Rica Dome. A food‐web based approach formulated by Siegel et al. (2014) performs as well or better than other empirical formulations, while simultaneously providing reasonable estimates of protozoan and mesozooplankton grazing rates. By tuning the Siegel et al. (2014) algorithm to match in situ grazing rates more accurately, we also obtain better in situ carbon export measurements. Adequate representations of food‐web relationships and grazing dynamics are therefore crucial to improving the accuracy of export predictions made from satellite–derived products. Nevertheless, considerable unexplained variance in export remains and must be explored before we can reliably use remote sensing products to assess the impact of climate change on biologically mediated carbon sequestration.

Phytoplankton production and grazing balances in the Costa Rica Dome.

We investigated phytoplankton production rates and grazing fates in the Costa Rica Dome (CRD) during summer 2010 based on dilution depth profiles analyzed by flow cytometry and pigments and mesozooplankton grazing assessed by gut fluorescence. Three community production estimates, from 14C uptake (1025 ± 113 mg C m-2 day-1) and from dilution experiments analyzed for total Chla (990 ± 106 mg C m-2 day-1) and flow cytometry populations (862 ± 71 mg C m-2 day-1), exceeded regional ship-based values by 2-3-fold. Picophytoplankton accounted for 56% of community biomass and 39% of production. Production profiles extended deeper for Prochlorococcus (PRO) and picoeukaryotes than for Synechococcus (SYN) and larger eukaryotes, but 93% of total production occurred above 40 m. Microzooplankton consumed all PRO and SYN growth and two-third of total production. Positive net growth of larger eukaryotes in the upper 40 m was balanced by independently measured consumption by mesozooplankton. Among larger eukaryotes, diatoms contributed ∼3% to production. On the basis of this analysis, the CRD region is characterized by high production and grazing turnover, comparable with or higher than estimates for the eastern equatorial Pacific. The region nonetheless displays characteristics atypical of high productivity, such as picophytoplankton dominance and suppressed diatom roles.

Mesozooplankton grazing impact on phytoplankton in the northern regions of the Kara Sea in autumn

The role of mass zooplankton species, copepods Pseudocalanus spp. and Oithona spp. and pteropods Limacina helicina, in grazing impact on phytoplankon in the upper 50 m layer was investigated. Material was collected at 14 stations along two sections crossing the outer shelf, continental slope and adjacent deep-water regions in the western and eastern branches of St. Anna trough (Kara Sea) in September 2011. Feeding rate was measured with fluorescence method. It was shown that total grazing impact on phytoplankton varied in a wide range from 0.2 to 8.7% of biomass and from 2 to 230% of daily primary production. The main contribution to phytoplankton grazing was made by small copepods Oithona spp. and juvenile pteropods. Maximum values of grazing were found in the frontal zones over the slope, minimum–over the outer continental shelf. Significant mesoscale spatial heterogeneity in total zooplankton grazing due to the changes in zooplankton composition and abundance was observed.

Zooplankton community grazing impact on a toxic bloom of Alexandrium fundyense in the Nauset Marsh System, Cape Cod, Massachusetts, USA

Embayments and salt ponds along the coast of Massachusetts can host localized blooms of the toxic dinoflagellate Alexandrium fundyense. One such system, exhibiting a long history of toxicity and annual closures of shellfish beds, is the Nauset Marsh System (NMS) on Cape Cod. In order to measure net growth rates of natural A. fundyense populations in the NMS during spring 2012, incubation experiments were conducted on seawater samples from two salt ponds within the NMS (Salt Pond and Mill Pond). Seawater samples containing natural populations of grazers and A. fundyense were incubated at ambient temperatures. Concentrations of A. fundyense after incubations were compared to initial abundances to determine net increases from population growth, or decreases presumed to be primarily due to grazing losses. Abundances of both microzooplankton (ciliates, rotifers, copepod nauplii and heterotrophic dinoflagellates) and mesozooplankton (copepodites and adult copepods, marine cladocerans, and meroplankton) grazers were also determined. This study documented net growth rates that were highly variable throughout the bloom, calculated from weekly bloom cell counts from the start of sampling to bloom peak in both ponds (Mill Pond range=0.12–0.46d−1; Salt Pond range=−0.02 to 0.44d−1). Microzooplankton grazers that were observed with ingested A. fundyense cells included polychaete larvae, rotifers, tintinnids, and heterotrophic dinoflagellates of the genera Polykrikos and Gymnodinium. Significant A. fundyense net growth was observed in two incubation experiments, and only a single experiment exhibited significant population losses. For the majority of experiments, due to high variability in data, net changes in A. fundyense abundance were not significant after the 24-h incubations. However, experimental net growth rates through bloom peak were not statistically distinguishable from estimated long-term average net growth rates of natural populations in each pond (Mill Pond=0.27d−1 and Salt Pond=0.20d−1), which led to peak bloom concentrations on the order of 106cellsl−1 in both ponds. Experimental net growth rates from the incubations underestimated the observed natural net growth rates at several time intervals prior to bloom peak, which may indicate that natural populations experienced additional sources of vegetative cells or periods of reduced losses that the 24-h incubation experiments did not capture, or that the experimental procedure introduced containment artifacts.

Mesozooplankton abundance in relation to the chlorophyll maximum in the Neuse River Estuary, North Carolina, USA: Implications for trophic dynamics

Estuaries often have distinct zones of high chlorophyll a concentrations, known as chlorophyll maximum (CMAX). The persistence of these features is often attributed to physical (mixing and light availability) and chemical (nutrient availability) features, but the role of mesozooplankton grazing is rarely explored. We measured the spatial and temporal variability of the CMAX and mesozooplankton community in the eutrophic Neuse River Estuary, North Carolina. We also conducted grazing experiments to determine the relative impact of mesozooplankton grazing on the CMAX during the phytoplankton growing season (spring through late summer). The CMAX was consistently located upriver of the zone of maximum zooplankton abundance, with an average spatial separation of 18 km. Grazing experiments in the CMAX region revealed negligible effect of mesozooplankton on chlorophyll a during March, and no effect during June or August. These results suggest that the spatial separation of the peak in chlorophyll a concentration and mesozooplankton abundance results in minimal impact of mesozooplankton grazing, contributing to persistence of the CMAX for prolonged time periods. In the Neuse River Estuary, the low mesozooplankton abundance in the CMAX region is attributed to lack of a low salinity tolerant species, predation by the ctenophore Mnemiopsis leidyi, and/or physiologic impacts on mesozooplankton growth rates due to temperature (in the case of low wintertime abundances). The consequences of this lack of overlap result in exacerbation of the effects of eutrophication; namely a lack of trophic transfer to mesozooplankton in this region and the sinking of phytodetritus to the benthos that fuels hypoxia.

Subtropical zooplankton assemblage promotes the harmful cyanobacterium Cylindrospermopsis raciborskii in a mesocosm experiment

Harmful algal blooms (HABs) with public health impacts threaten freshwater ecosystems, including drinking water reservoirs, globally. Subtropical systems are often dominated by filamentous and colonial cyanobacteria, algae that are potentially less accessible for consumption by resident meso-zooplankton grazers. Less understood than selective grazing is the role of zooplankton in regenerating nutrients and facilitating growth of algae with efficient uptake strategies, such as the toxin-producing cyanobacterium, Cylindrospermopsis raciborskii. Using 800-L bags suspended in the upper 3 m of the water column, we examined the growth of C. raciborskii under four treatments: 3 ambient zooplankton biomass, 10 zooplankton, 10 zooplankton plus inorganic P addition and a no amendment control (3Z, 10Z, 10ZP, control, respectively). After 4 days, C. raciborskii relative abundance doubled in the 10Z and 10ZP treatments compared with the control and 3Z treatments, and after 7 days P addition resulted in 20% higher relative C. raciborskii biomass compared with other treatments, and an order of magnitude increase in N-fixing phytoplankton. The particulate C : P ratio declined in the 10Z and 10ZP mesocosms, indicating that meso-zooplankton facilitated P transfer to algae. Overall, the copepod dominated subtropical meso-zooplankton assemblage promoted C. raciborskii abundance and biomass over the short-term, demonstrating their facilitation of subtropical freshwater HAB formation.

Particle flux characterisation and sedimentation patterns of protistan plankton during the iron fertilisation experiment LOHAFEX in the Southern Ocean

The taxonomic composition and types of particles comprising the downward particle flux were examined during the mesoscale artificial iron fertilisation experiment LOHAFEX. The experiment was conducted in low-silicate waters of the Atlantic Sector of the Southern Ocean during austral summer (January–March 2009), and induced a bloom dominated by small flagellates. Downward particle flux was low throughout the experiment, and not enhanced by addition of iron; neutrally buoyant sediment traps contained mostly faecal pellets and faecal material apparently reprocessed by mesozooplankton. TEP fluxes were low, ≤5mgGXeq.m−2d−1, and a few phytodetrital aggregates were found in the sediment traps. Only a few per cent of the POC flux was found in the traps consisting of intact protist plankton, although remains of taxa with hard body parts (diatoms, tintinnids, thecate dinoflagellates and foraminifera) were numerous, far more so than intact specimens of these taxa. Nevertheless, many small flagellates and coccoid cells, belonging to the pico- and nanoplankton, were found in the traps, and these small, soft-bodied cells probably contributed the majority of downward POC flux via mesozooplankton grazing and faecal pellet export. TEP likely played an important role by aggregating these small cells, and making them more readily available to mesozooplankton grazers.

Mesozooplankton Grazing on Picocyanobacteria in the Baltic Sea as Inferred from Molecular Diet Analysis

Our current knowledge on the microbial component of zooplankton diet is limited, and it is generally assumed that bacteria-sized prey is not directly consumed by most mesozooplankton grazers in the marine food webs. We questioned this assumption and conducted field and laboratory studies to examine picocyanobacteria contribution to the diets of Baltic Sea zooplankton, including copepods. First, qPCR targeting ITS-1 rDNA sequence of the picocyanobacteria Synechococcus spp. was used to examine picocyanobacterial DNA occurrence in the guts of Baltic zooplankton (copepods, cladocerans and rotifers). All field-collected zooplankton were found to consume picocyanobacteria in substantial quantities. In terms of Synechococcus quantity, the individual gut content was highest in cladocerans, whereas biomass-specific gut content was highest in rotifers and copepod nauplii. Moreover, the gut content in copepods was positively related to the picocyanobacteria abundance and negatively to the total phytoplankton abundance in the water column at the time of sampling. This indicates that increased availability of picocyanobacteria resulted in the increased intake of this prey and that copepods may rely more on picoplankton when food in the preferred size range declines. Second, a feeding experiments with a laboratory reared copepod Acartia tonsa fed a mixture of the picocyanobacterium Synechococcus bacillaris and microalga Rhodomonas salina confirmed that copepods ingested Synechococcus, even when the alternative food was plentiful. Finally, palatability of the picocyanobacteria for A. tonsa was demonstrated using uptake of 13C by the copepods as a proxy for carbon uptake in feeding experiment with 13C-labeled S. bacillaris. These findings suggest that, if abundant, picoplankton may become an important component of mesozooplankton diet, which needs to be accounted for in food web models and productivity assessments.

Changes in the swimming behavior of Pseudodiaptomus annandalei (Copepoda, Calanoida) adults exposed to the diatom toxin 2-trans, 4-trans decadienal

Diatoms are broadly present in marine habitats and often dominate seasonal phytoplankton blooms in polar and temperate latitudes. Certain species produce polyunsaturated aldehydes upon mechanical wounding caused by mesozooplankton grazing. Ample evidence is available on toxin-induced reproductive failure in copepods, yet their behavioral effects remain unclear. Here we present results of laboratory experiments in which we investigated the immediate effects of the diatom-derived aldehyde 2-trans, 4-trans decadienal on the three-dimensional swimming behavior of the calanoid copepod Pseudodiaptomus annandalei. Short-term direct exposure to the toxin at 3μM, 6μM and 12μM induced hyperactivity in the three adult states, as evidenced by a marked and dose-dependent increase in the number of trajectories. It also caused a higher proportion of vertical movements. In males and ovigerous females exposed to decadienal at 3μM and 6μM, hyperactivity came with an equally specific dose-dependent decrease in swimming speed. Males and ovigerous females swam faster at 12μM than at 6μM, suggesting a complex mode of action of the toxin. In non-ovigerous females, decadienal had little effects on swimming speed, supporting the assumption that female copepods are less affected by certain environmental stressors. Multifractal analysis revealed differences in the statistical properties of the swimming behavior between experimental conditions. The moment structure function of the displacement appeared to be moderately multifractal in the three adult states swimming in control water. Ethanol as carrier solvent at 200ppm caused an increase in swimming speed and a switch toward a more ballistic motion in males and ovigerous females. On the opposite, exposure to the toxin reduced or cancelled the effects of ethanol and resulted in a more Brownian motion for high moment values. Decadienal had little effects on the behavior of non-ovigerous females except at the highest concentration. Our results demonstrate that decadienal, a model diatom aldehyde, impairs the behavior of adult copepods. They provide further information on the interaction between diatoms and their main predator.

Planktonic food webs revisited: Reanalysis of results from the linear inverse approach

Identification of the trophic pathway that dominates a given planktonic assemblage is generally based on the distribution of biomasses among food-web compartments, or better, the flows of materials or energy among compartments. These flows are obtained by field observations and a posteriori analyses, including the linear inverse approach. In the present study, we re-analysed carbon flows obtained by inverse analysis at 32 stations in the global ocean and one large lake. Our results do not support two “classical” views of plankton ecology, i.e. that the herbivorous food web is dominated by mesozooplankton grazing on large phytoplankton, and the microbial food web is based on microzooplankton significantly consuming bacteria; our results suggest instead that phytoplankton are generally grazed by microzooplankton, of which they are the main food source. Furthermore, we identified the “phyto-microbial food web”, where microzooplankton largely feed on phytoplankton, in addition to the already known “poly-microbial food web”, where microzooplankton consume more or less equally various types of food. These unexpected results led to a (re)definition of the conceptual models corresponding to the four trophic pathways we found to exist in plankton, i.e. the herbivorous, multivorous, and two types of microbial food web. We illustrated the conceptual trophic pathways using carbon flows that were actually observed at representative stations. The latter can be calibrated to correspond to any field situation. Our study also provides researchers and managers with operational criteria for identifying the dominant trophic pathway in a planktonic assemblage, these criteria being based on the values of two carbon ratios that could be calculated from flow values that are relatively easy to estimate in the field.

Mechanistic origins of variability in phytoplankton dynamics. Part II: analysis of mesocosm blooms under climate change scenarios

Driving factors of phytoplankton spring blooms have been discussed since long, but rarely analyzed quantitatively. Here, we use a mechanistic size-based ecosystem model to reconstruct observations made during the Kiel mesocosm experiments (2005–2006). The model accurately hindcasts highly variable bloom developments including community shifts in cell size. Under low light, phytoplankton dynamics was mostly controlled by selective mesozooplankton grazing. Selective grazing also explains initial dominance of large diatoms under high light conditions. All blooms were mainly terminated by aggregation and sedimentation. Allometries in nutrient uptake capabilities led to a delayed, post-bloom dominance of small species. In general, biomass and trait dynamics revealed many mutual dependencies, while growth factors decoupled from the respective selective forces. A size shift induced by one factor often changed the growth dependency on other factors. Within climate change scenarios, these indirect effects produced large sensitivities of ecosystem fluxes to the size distribution of winter phytoplankton. These sensitivities exceeded those found for changes in vertical mixing, whereas temperature changes only had minimal impacts.

Cross-shelf variability in the Iberian Peninsula Upwelling System: Impact of a mesoscale filament

Based on a multidisciplinary survey in the Iberian upwelling during late summer 2007, this paper analysed comparatively the cross-shore variability and offshore transport across the upwelling front and within a mesoscale filament.Along the East–West (EW) sections, transient upwelling pulses bring regularly cold, fresh and nutrient-enriched waters to the surface, triggering intense biological responses. Offshore advection by wind-forced Ekman drift of the successive fronts, interrupted by relaxation periods, drive the variability of the planktonic communities. While the near-shore areas are dominated by relatively small phytoplankton controlled by mesozooplankton grazing, large cells of diatoms appear after a short decay. Although microphytoplankton dominates largely the shelf communities, the species composition varies during the offshore drift with the apparition of dinoflagellates and the gradual development of large zooplankton individuals. The oligotrophic ecosystem characterised by small organisms and low biomass (∼80km offshore) contrasts strongly with the transitional area and the coastal upwelling.The low density waters within the filament and the existence of a pair of opposite rotating eddies at its base and tip promote its generation and rapid seaward extension. The intensified offshore advection of coastal enriched waters considerably increases the area favouring a productive ecosystem (until ∼160km off the coast). Cross-shelf variability of bio-physical variables is observed in the filament as along EW sections, although a subsequent homogenisation within the mesoscale structure erases the sharp fronts. Off the shelf within the filament, the chlorophyll a is distinctly organised as a shallow subsurface maximum dominated by nano-phytoplankton. The relative physical isolation of a dynamical food-web in the filament is also promoting nutrient remineralisation under the structure.Finally, we estimate that mesoscale filaments, although being less extended meridionally than the upwelling front itself (∼40% of the length of the front) are responsible for a greater offshore transport of chlorophyll (∼60% of total cross-shelf exchanges) over the Iberian system. Despite the favourable wind pulses advecting westward the successive upwelling fronts, self-propelled filaments provide permanent offshore transport, even under wind relaxation period, thus playing a major role in cross-shelf exchanges.

A <sup>13</sup>C labelling study on carbon fluxes in Arctic plankton communities under elevated CO<sub>2</sub> levels

Abstract. The effect of CO2 on carbon fluxes (production, consumption, and export) in Arctic plankton communities was investigated during the 2010 EPOCA (European project on Ocean Acidification) mesocosm study off Ny Ålesund, Svalbard. 13C labelled bicarbonate was added to nine mesocosms with a range in pCO2 (185 to 1420 μatm) to follow the transfer of carbon from dissolved inorganic carbon (DIC) into phytoplankton, bacterial and zooplankton consumers, and export. A nutrient–phytoplankton–zooplankton–detritus model amended with 13C dynamics was constructed and fitted to the data to quantify uptake rates and carbon fluxes in the plankton community. The plankton community structure was characteristic for a post-bloom situation and retention food web and showed high bacterial production (∼31% of primary production), high abundance of mixotrophic phytoplankton, low mesozooplankton grazing (∼6% of primary production) and low export (∼7% of primary production). Zooplankton grazing and export of detritus were sensitive to CO2: grazing decreased and export increased with increasing pCO2. Nutrient addition halfway through the experiment increased the export, but not the production rates. Although mixotrophs showed initially higher production rates with increasing CO2, the overall production of POC (particulate organic carbon) after nutrient addition decreased with increasing CO2. Interestingly, and contrary to the low nutrient situation, much more material settled down in the sediment traps at low CO2. The observed CO2 related effects potentially alter future organic carbon flows and export, with possible consequences for the efficiency of the biological pump.