Microzooplankton Grazers Research Articles

Killing the predator: impacts of highest-predator mortality on the global-ocean ecosystem structure

Abstract. Recent meta-analyses suggest that microzooplankton biomass density scales linearly with phytoplankton biomass density, suggesting a simple, general rule may underpin trophic structure in the global ocean. Here, we use a set of highly simplified food web models, solved within a global general circulation model, to examine the core drivers of linear predator–prey scaling. We examine a parallel food chain model which assumes microzooplankton grazers feed on distinct size classes of phytoplankton and contrast this with a diamond food web model allowing shared microzooplankton predation on a range of phytoplankton size classes. Within these two contrasting model structures, we also evaluate the impact of fixed vs. density-dependent microzooplankton mortality. We find that the observed relationship between microzooplankton predators and prey can be reproduced with density-dependent mortality on the highest predator, regardless of choices made about plankton food web structure. Our findings point to the importance of parameterizing mortality of the highest predator for simple food web models to recapitulate trophic structure in the global ocean.

Microzooplankton grazers induce chain length plasticity in colonial diatoms

AbstractMany diatoms form long chains and the distribution of chain lengths within a species depends on several environmental factors, among them grazing risk. Larger grazers, such as copepods, efficiently handle and ingest even very long chains but are less efficient with individual cells, whereas most smaller grazers are unable to feed on chains exceeding a few cells in length. Copepod cues make several species shorten their chain length, and theory predicts that cues from small grazers should induce increased, but this remains to be tested, despite the importance of diatoms in marine food webs. Here, we expose three species of chain‐forming diatoms, Skeletonema marinoi, Chaetoceros affinis, and Thalassiosira rotula to cues from various actively feeding micrograzers and record their response. The effect of grazer presence on chain length varies depending on both the type of grazer and the diatom species. For example, S. marinoi increased its chain length when exposed to grazing cues from the heterotrophic dinoflagellate Gyrodinium dominans and the ciliate Euplotes sp. but not to a copepod nauplii (Temora longicornis). C. affinis also responded to cues from grazing G. dominans by increasing chain length, and the response increased with exposure time. Finally, T. rotula did not respond to grazing cues from G. dominans, but rather inhibited the dinoflagellates' ability to feed, presumably through the release of chemical compounds. Our results suggest that some chainforming diatoms can sense and appropriately respond to fast‐growing micrograzers, thus contributing to the success of diatoms in marine environments.

An empirical model of carbon flow through marine viruses and microzooplankton grazers.

Viruses and microzooplankton grazers represent major sources of mortality for marine phytoplankton and bacteria, redirecting the flow of organic material throughout the world's oceans. Here, we investigate the use of nonlinear population models of interactions between phytoplankton, viruses and grazers as a means to quantitatively constrain the flow of carbon through marine microbial ecosystems. We augment population models with a synthesis of laboratory-based estimates of prey, predator and viral life history traits that constrain transfer efficiencies. We then apply the model framework to estimate loss rates in the California Current Ecosystem (CCE). With our empirically parameterized model, we estimate that, of the total losses mediated by viruses and microzooplankton grazing at the focal CCE site, 22 ± 3%, 46 ± 27%, 3 ± 2% and 29 ± 20% were directed to grazers, sloppy feeding (as well as excretion and respiration), viruses and viral lysate respectively. We identify opportunities to leverage ecosystem models and conventional mortality assays to further constrain the quantitative rates of critical ecosystem processes.

Contrasting Controls on Microzooplankton Grazing and Viral Infection of Microbial Prey

The encounter and capture of bacteria and phytoplankton by microbial predators and parasites is fundamental to marine ecosystem organization and activity. Here, we combined biophysical models with published laboratory measurements to infer functional traits, including encounter kernel and capture efficiency, for a wide range of marine viruses and microzooplankton grazers. Despite virus particles being orders of magnitude smaller than microzooplankton grazers, virus encounter kernels and adsorption rates were in many cases comparable in magnitude to grazer encounter kernel and clearance, pointing to Brownian motion as a highly effective method of transport for viruses. Inferred virus adsorption efficiency covered many orders of magnitude, but the median virus adsorption efficiency was between 5 to 25% depending on the assumed host swimming speed. Uncertainty on predator detection area and swimming speed prevented robust inference of grazer capture efficiency, but sensitivity analysis was used to identify bounds on unconstrained processes. These results provide a common functional trait framework for understanding marine host-virus and predator-prey interactions, and highlight the value of theory for interpreting measured life-history traits.

Exploring nonlinear functional responses of zooplankton grazers in dilution experiments via optimization techniques

AbstractUnderstanding complex dynamics of planktonic ecosystems requires good knowledge of predator functional responses, especially for microzooplankton grazers. Although natural microzooplankton communities are highly diverse, we often need to quantify microzooplankton grazing at the macrolevel, considering them as a single “box.” Here, we consider community microzooplankton predation rates as a function of their prey abundance. Estimating the functional response of microzooplankton as a function of bulk prey often exploits the dilution approach: joint incubation experiments of grazers and their prey where samples are subjected to a range of dilution factors. However, the classic methodology appears reliable only for a simple linear functional response of grazers and it requires appropriate adjustments for nonlinearity and saturation of food consumption at high prey abundance. We explore how recently suggested methods applying nonlinear functional responses of bulk microzooplankton grazing can be enhanced to provide more robust results. We argue that different formulations of the functional response should be fitted to the data and that an appropriate choice of dilution factors in experiments can substantially improve the accuracy of feeding rate estimation. Pragmatically, our work shows that for oligotrophic ecosystems, it is best to run incubations where the majority of dilution factors are close to D = 1 (undiluted), whereas, for eutrophic waters, dilution factors should be run close to D = 0 as well as D = 1. Furthermore, running equidistant dilution factors represents a waste of effort; it is more important to run at fewer values of D with greater replication.

Altering phytoplankton growth rates changes their value as food for microzooplankton grazers

Altering phytoplankton growth rates changes their value as food for microzooplankton grazers

Growth and mortality of coccolithophores during spring in a temperate Shelf Sea (Celtic Sea, April 2015)

Coccolithophores are key components of phytoplankton communities, exerting a critical impact on the global carbon cycle and the Earth’s climate through the production of coccoliths made of calcium carbonate (calcite) and bioactive gases. Microzooplankton grazing is an important mortality factor in coccolithophore blooms, however little is currently known regarding the mortality (or growth) rates within non-bloom populations. Measurements of coccolithophore calcite production (CP) and dilution experiments to determine microzooplankton (≤63 µm) grazing rates were made during a spring cruise (April 2015) at the Central Celtic Sea (CCS), shelf edge (CS2), and within an adjacent April bloom of the coccolithophore Emiliania huxleyi at station J2.CP at CCS ranged from 10.4 to 40.4 µmol C m−3 d−1 and peaked at the height of the spring phytoplankton bloom (peak chlorophyll-a concentrations ∼6 mg m−3). Cell normalised calcification rates declined from ∼1.7 to ∼0.2 pmol C cell−1 d−1, accompanied by a shift from a mixed coccolithophore species community to one dominated by the more lightly calcified species E. huxleyi and Calciopappus caudatus. At the CCS, coccolithophore abundance increased from 6 to 94 cells mL−1, with net growth rates ranging from 0.06 to 0.21 d−1 from the 4th to the 28th April. Estimates of intrinsic growth and grazing rates from dilution experiments, at the CCS ranged from 0.01 to 0.86 d−1 and from 0.01 to 1.32 d−1, respectively, which resulted in variable net growth rates during April. Microzooplankton grazers consumed 59 to >100% of daily calcite production at the CCS. Within the E. huxleyi bloom a maximum density of 1986 cells mL−1 was recorded, along with CP rates of 6000 µmol C m−3 d−1 and an intrinsic growth rate of 0.29 d−1, with ∼80% of daily calcite production being consumed.Our results show that microzooplankton can exert strong top-down control on both bloom and non-bloom coccolithophore populations, grazing over 60% of daily growth (and calcite production). The fate of consumed calcite is unclear, but may be lost either through dissolution in acidic food vacuoles, and subsequent release as CO2, or export to the seabed after incorporation into small faecal pellets. With such high microzooplankton-mediated mortality losses, the fate of grazed calcite is clearly a high priority research direction.

The influence of viruses on phytoplankton and bacterial productivity in Hamilton Harbour, an impaired embayment of Lake Ontario

Abstract This study investigated the ecological effects of viruses on the microbial community in Hamilton Harbour, a Lake Ontario embayment. A modified Landry-Hassett style dilution assay was used to assess the relative impact of microzooplankton grazers and viruses on phytoplankton community and population growth and mortality in August 2013. Although viral effects were not observed in this experiment, significant grazing mortality was detected in all instances. Similar dilution methodology was then applied in conjunction with standard bacterial and size-fractionated primary productivity assays (using 3 H–leucine and Na 14 CO 3 , respectively). The first iteration of this experiment (August 2013) involved the use of a single dilution treatment with grazer-and-virus-free filtrate, and led to the observation that the combined activity of grazers and viruses appeared to stimulate productivity for phytoplankton 20 μm may have first been limited by viral infection/lysis and then stimulated by grazer dynamics. Our results provide the first evidence that viruses can actively affect microbial productivity in the Harbour.

Allochthonous carbon is a major driver of the microbial food web – A mesocosm study simulating elevated terrestrial matter runoff

Climate change predictions indicate that coastal and estuarine environments will receive increased terrestrial runoff via increased river discharge. This discharge transports allochthonous material, containing bioavailable nutrients and light attenuating matter. Since light and nutrients are important drivers of basal production, their relative and absolute availability have important consequences for the base of the aquatic food web, with potential ramifications for higher trophic levels. Here, we investigated the effects of shifts in terrestrial organic matter and light availability on basal producers and their grazers. In twelve Baltic Sea mesocosms, we simulated the effects of increased river runoff alone and in combination. We manipulated light (clear/shade) and carbon (added/not added) in a fully factorial design, with three replicates. We assessed microzooplankton grazing preferences in each treatment to assess whether increased terrestrial organic matter input would: (1) decrease the phytoplankton to bacterial biomass ratio, (2) shift microzooplankton diet from phytoplankton to bacteria, and (3) affect microzooplankton biomass. We found that carbon addition, but not reduced light levels per se resulted in lower phytoplankton to bacteria biomass ratios. Microzooplankton generally showed a strong feeding preference for phytoplankton over bacteria, but, in carbon-amended mesocosms which favored bacteria, microzooplankton shifted their diet towards bacteria. Furthermore, low total prey availability corresponded with low microzooplankton biomass and the highest bacteria/phytoplankton ratio. Overall our results suggest that in shallow coastal waters, modified with allochthonous matter from river discharge, light attenuation may be inconsequential for the basal producer balance, whereas increased allochthonous carbon, especially if readily bioavailable, favors bacteria over phytoplankton. We conclude that climate change induced shifts at the base of the food web may alter energy mobilization to and the biomass of microzooplankton grazers.

Mixotrophy in the Marine Plankton.

Mixotrophs are important components of the bacterioplankton, phytoplankton, microzooplankton, and (sometimes) zooplankton in coastal and oceanic waters. Bacterivory among the phytoplankton may be important for alleviating inorganic nutrient stress and may increase primary production in oligotrophic waters. Mixotrophic phytoflagellates and dinoflagellates are often dominant components of the plankton during seasonal stratification. Many of the microzooplankton grazers, including ciliates and Rhizaria, are mixotrophic owing to their retention of functional algal organelles or maintenance of algal endosymbionts. Phototrophy among the microzooplankton may increase gross growth efficiency and carbon transfer through the microzooplankton to higher trophic levels. Characteristic assemblages of mixotrophs are associated with warm, temperate, and cold seas and with stratification, fronts, and upwelling zones. Modeling has indicated that mixotrophy has a profound impact on marine planktonic ecosystems and may enhance primary production, biomass transfer to higher trophic levels, and the functioning of the biological carbon pump.

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.

The role of mixotrophic protists in the biological carbon pump

Abstract. The traditional view of the planktonic food web describes consumption of inorganic nutrients by photoautotrophic phytoplankton, which in turn supports zooplankton and ultimately higher trophic levels. Pathways centred on bacteria provide mechanisms for nutrient recycling. This structure lies at the foundation of most models used to explore biogeochemical cycling, functioning of the biological pump, and the impact of climate change on these processes. We suggest an alternative new paradigm, which sees the bulk of the base of this food web supported by protist plankton communities that are mixotrophic – combining phototrophy and phagotrophy within a single cell. The photoautotrophic eukaryotic plankton and their heterotrophic microzooplankton grazers dominate only during the developmental phases of ecosystems (e.g. spring bloom in temperate systems). With their flexible nutrition, mixotrophic protists dominate in more-mature systems (e.g. temperate summer, established eutrophic systems and oligotrophic systems); the more-stable water columns suggested under climate change may also be expected to favour these mixotrophs. We explore how such a predominantly mixotrophic structure affects microbial trophic dynamics and the biological pump. The mixotroph-dominated structure differs fundamentally in its flow of energy and nutrients, with a shortened and potentially more efficient chain from nutrient regeneration to primary production. Furthermore, mixotrophy enables a direct conduit for the support of primary production from bacterial production. We show how the exclusion of an explicit mixotrophic component in studies of the pelagic microbial communities leads to a failure to capture the true dynamics of the carbon flow. In order to prevent a misinterpretation of the full implications of climate change upon biogeochemical cycling and the functioning of the biological pump, we recommend inclusion of multi-nutrient mixotroph models within ecosystem studies.

Virally-Mediated Versus Grazer-Induced Mortality Rates in a Warm-Temperate Inverse Estuary (Spencer Gulf, South Australia)

We investigated the seasonal dynamics of flow cytometrically-defined populations of viruses, heterotrophic bacteria, and the picoeukaryotic and prokaryotic phytoplankton at three sites in the temperate oligotrophic inverse estuary of Spencer Gulf (South Australia). We consistently identified two sub-populations of viruses, three sub-populations of heterotrophic bacteria, one population of picoeukaryotic phytoplankton and two populations of prokaryotic phytoplankton (cyanobacteria Prochlorococcus and Synechococcus). Both the cytometric community composition and the abundance of viruses, heterotrophic bacteria and both prokaryotic (Synechococcus and Prochlorococcus) and eukaryotic picophytoplankton were consistent with previous observations conducted in South Australian continental shelf waters. Noticeably LDNA bacteria (i.e. inactive or dormant cells) were consistently significantly the most abundant group of heterotrophic bacteria (totaling from 29% to 68% of total bacterial abundance) and were up to 10-fold more abundant than that previously reported in South Australian continental shelf waters, including the nearby Saint Vincent Gulf. These results suggest an overall low activity of the microbial community, and are consistent with previous evidence that LDNA cells may play a greater role in heterotrophic processes than HDNA cells in oligotrophic waters. In an attempt to further assess the qualitative and quantitative nature of the mortality of these organisms, we used a specific dilution assay to assess the relative contribution of viruses and microzooplankton grazers to the mortality rates of heterotrophic bacteria, and picoeukaryotic and prokaryotic phytoplankton. We consistently reported site-specific, population specific and sea-son-specific viral lysis and grazing rates of heterotrophic bacteria and the picoeukaryotic and prokaryotic (cyanobacteria Prochlorococcus and Synechococcus) phytoplankton across sites and seasons. Specifically, both viral lysis and micro-zooplankton grazing rates of heterotrophic bacteria were consistently relatively low across sites and seasons, even though their seasonality suggested an overall dominance of grazing over viral lysis in both summer and winter. In contrast, no seasonality is found in either lysis or grazing rates of prokaryotic and eukaryotic picophytoplankton, which are comparable to previous observations conducted in oligotrophic waters, suggesting the mortality dynamics of these populations is similar to those encountered in other oligotrophic waters. The observed patterns of mortality rates of heterotrophic bacteria and both prokaryotic and eukaryotic picophytoplankton are consistent with the low chlorophyll concentration and production previously observed in the waters of the Spencer Gulf.

Role of infochemical mediated zooplankton grazing in a phytoplankton competition model

Infochemicals released by marine phytoplankton play important roles in food web interactions by influencing the feeding behavior and selectivity of zooplanktonic predators. Recent modeling efforts have focused on the role of such chemicals as toxic grazing deterrents in phytoplankton competition. However, infochemicals may also be utilized as grazing cues, leading predators to profitable foraging patches. Here we investigate the role of infochemical mediated zooplankton grazing in a standard 3-species phytoplankton competition model, with the aim of further elucidating the ecological role of phytoplankton derived infochemicals. We then extend this to consider a more realistic 4-species model. The models produce a range of solutions depending on the strength of competition and microzooplankton grazing selectivity. Our key result is that infochemical chemoattractants, which increase the susceptibility of the producer to grazing, can provide a refuge for both competing phytoplankton species by attracting carnivorous copepods to consume microzooplankton grazers in a multi-trophic interaction. Our results indicate that infochemicals potentially have important consequences for the dynamics of marine food webs.

Temporal dynamics of carbon flow through the microbial plankton community in a coastal upwelling system off northern Baja California, Mexico

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 461:31-46 (2012) - DOI: https://doi.org/10.3354/meps09782 Temporal dynamics of carbon flow through the microbial plankton community in a coastal upwelling system off northern Baja California, Mexico Lorena Linacre1, 2,*, Michael R. Landry3, Ramón Cajal-Medrano4, J. Rubén Lara-Lara5, J. Martín Hernández-Ayón6, Rosa R. Mouriño-Pérez2, Ernesto García-Mendoza5, Carmen Bazán-Guzmán5 1Programa de Doctorado en Oceanografía Costera, Facultad de Ciencias Marinas/Instituto de Investigaciones Oceanológicas, 4Facultad de Ciencias Marinas, and 6Instituto de Investigaciones Oceanológicas; Universidad Autónoma de Baja California (UABC), Ensenada, Baja California 22860, Mexico 2Departamento de Microbiología, División de Biología Experimental y Aplicada, and 5Departamento de Oceanografía Biológica, División de Oceanología; Centro de Investigación Científica y de Educación Superior de Ensenada (CICESE), Ensenada, Baja California 22860, Mexico 3Scripps Institution of Oceanography, University of California at San Diego, La Jolla, California 92093-0227, USA *Email: lorenalinacre@gmail.com ABSTRACT: We investigated the temporal dynamics of carbon flow through the microbial food web of a coastal upwelling system (ENSENADA station) off northern Baja California during 6 cruises (September 2007 to November 2008). Carbon biomass assessments for major autotrophic size groups (pico- to micro-sized cells) and their microzooplankton grazers were based on analyses using flow cytometry, HPLC pigments and epifluorescence microscopy. Taxon-specific phytoplankton growth and microzooplankton grazing rates were determined from 24 h in situ incubations in the euphotic zone using an abbreviated 3-treatment dilution technique. Carbon biomass and instantaneous growth and grazing rate determinations were used to estimate daily rates of taxon-specific production and losses due to microzooplankton grazing. Overall, microbial biomass showed a close balance between autotrophic and heterotrophic components, except during a period of very strong upwelling (April 2008), which favored large phytoplankters and high primary production. Throughout a wide range of environmental conditions, the community primary production (PP) attributed both to small (mostly picophytoplankton and prasinophytes) and large (mostly diatoms and autotrophic dinoflagellates) autotrophs was significantly grazed (78 ± 9% of PP) by small (<20 µm) and large (>20 µm) ciliates and flagellates (including mixotrophic dinoflagellates), respectively, showing complementary temporal shifts in protistan grazer types that matched the dominant phytoplankton. While large diatoms were strongly consumed by large ciliates during the 2 most productive periods (September 2007 and April 2008), pico- and nano-sized phytoplankton were grazed most by nanoflagellates and small ciliates from November 2007 to January 2008. Consequently, biogenic carbon production in this ecosystem is transferred through a multivorous food web. KEY WORDS: Biogenic carbon flow · Multivorous food web · Phytoplankton growth rate · Phytoplankton grazing rate · Primary production Full text in pdf format PreviousNextCite this article as: Linacre L, Landry MR, Cajal-Medrano R, Lara-Lara JR and others (2012) Temporal dynamics of carbon flow through the microbial plankton community in a coastal upwelling system off northern Baja California, Mexico. Mar Ecol Prog Ser 461:31-46. https://doi.org/10.3354/meps09782 Export citation RSS - Facebook - Tweet - linkedIn Cited by Published in MEPS Vol. 461. Online publication date: August 08, 2012 Print ISSN: 0171-8630; Online ISSN: 1616-1599 Copyright © 2012 Inter-Research.

Geographical distribution of red and green Noctiluca scintillans

The dinoflagellate Noctiluca scintillans is one of the most important and abundant red tide organisms and it is distributed world-wide. It occurs in two forms. Red Noctiluca is heterotrophic and fills the role of one of the microzooplankton grazers in the foodweb. In contrast, green Noctiluca contains a photosynthetic symbiont Pedinomonas noctilucae (a prasinophyte), but it also feeds on other plankton when the food supply is abundant. In this review, we document the global distribution of these two forms and include the first maps of their global distribution. Red Noctiluca occurs widely in the temperate to sub-tropical coastal regions of the world. It occurs over a wide temperature range of about 10°C to 25°C and at higher salinities (generally not in estuaries). It is particularly abundant in high productivity areas such as upwelling or eutrophic areas where diatoms dominate since they are its preferred food source. Green Noctiluca is much more restricted to a temperature range of 25°C–30°C and mainly occurs in tropical waters of Southeast Asia, Bay of Bengal (east coast of India), in the eastern, western and northern Arabian Sea, the Red Sea, and recently it has become very abundant in the Gulf of Oman. Red and green Noctiluca do overlap in their distribution in the eastern, northern and western Arabian Sea with a seasonal shift from green Noctiluca in the cooler winter convective mixing, higher productivity season, to red Noctiluca in the more oligotrophic warmer summer season.

The role of ciliates, heterotrophic dinoflagellates and copepods in structuring spring plankton communities at Helgoland Roads, North Sea

Mesocosm experiments coupled with dilution grazing experiments were carried out during the phytoplankton spring bloom 2009. The interactions between phytoplankton, microzooplankton and copepods were investigated using natural plankton communities obtained from Helgoland Roads (54°11.3′N; 7°54.0′E), North Sea. In the absence of mesozooplankton grazers, the microzooplankton rapidly responded to different prey availabilities; this was most pronounced for ciliates such as strombidiids and strobilids. The occurrence of ciliates was strongly dependent on specific prey and abrupt losses in their relative importance with the disappearance of their prey were observed. Thecate and athecate dinoflagellates had a broader food spectrum and slower reaction times compared with ciliates. In general, high microzooplankton potential grazing impacts with an average consumption of 120% of the phytoplankton production (Pp) were measured. Thus, the decline in phytoplankton biomass could be mainly attributed to an intense grazing by microzooplankton. Copepods were less important phytoplankton grazers consuming on average only 47% of Pp. Microzooplankton in turn contributed a substantial part to the copepods’ diets especially with decreasing quality of phytoplankton food due to nutrient limitation over the course of the bloom. Copepod grazing rates exceeded microzooplankton growth, suggesting their strong top-down control potential on microzooplankton in the field. Selective grazing by microzooplankton was an important factor for stabilising a bloom of less-preferred diatom species in our mesocosms with specific species (Thalassiosira spp., Rhizosolenia spp. and Chaetoceros spp.) dominating the bloom. This study demonstrates the importance of microzooplankton grazers for structuring and controlling phytoplankton spring blooms in temperate waters and the important role of copepods as top-down regulators of microzooplankton.

Dinoflagellates and ciliates at Helgoland Roads, North Sea

A monitoring programme for microzooplankton was started at the long-term sampling station “Kabeltonne” at Helgoland Roads (54°11.3′N; 7°54.0′E) in January 2007 in order to provide more detailed knowledge on microzooplankton occurrence, composition and seasonality patterns at this site and to complement the existing plankton data series. Ciliate and dinoflagellate cell concentration and carbon biomass were recorded on a weekly basis. Heterotrophic dinoflagellates were considerably more important in terms of biomass than ciliates, especially during the summer months. However, in early spring, ciliates were the major group of microzooplankton grazers as they responded more quickly to phytoplankton food availability. Mixotrophic dinoflagellates played a secondary role in terms of biomass when compared to heterotrophic species; nevertheless, they made up an intense late summer bloom in 2007. The photosynthetic ciliate Myrionecta rubra bloomed at the end of the sampling period. Due to its high biomass when compared to crustacean plankton especially during the spring bloom, microzooplankton should be regarded as the more important phytoplankton grazer group at Helgoland Roads. Based on these results, analyses of biotic and abiotic factors driving microzooplankton composition and abundance are necessary for a full understanding of this important component of the plankton.

Grazing suppression of dimethylsulphoniopropionate (DMSP) accumulation in iron-fertilised, sub-Antarctic waters

The impact of in situ iron fertilisation on the production of particulate dimethylsulphoniopropionate (DMSPp) and its breakdown product dimethyl sulphide (DMS) was monitored during the SOLAS air–sea gas exchange experiment (SAGE). The experiment was conducted in the high nitrate, low chlorophyll (HNLC) waters of the sub-Antarctic Southern Ocean (46.7°S 172.5°E) to the south-east of New Zealand, during March–April, 2004. In addition to monitoring net changes in the standing stocks of DMSPp and DMS, a series of dilution experiments were used to determine the DMSPp production and consumption rates in relation to increased iron availability. In contrast to previous experiments in the Southern Ocean, DMS concentrations decreased over the course of the 15-d iron-fertilisation experiment, from an integrated volume-specific concentration in the mixed layer on day 0 of 0.78 nM (measured values 0.65–0.91 nM) to 0.46 nM (measured values 0.42–0.47 nM) by day 15, in parallel with the surrounding waters. DMSPp, chlorophyll a and the abundance of photosynthetic picoeukaryotes exhibited indiscernible or only moderate increases in response to the raised iron availability, despite an obvious physiological response by the phytoplankton. High specific growth rates of DMSPp, equivalent to 0.8–1.2 doublings d −1, occurred at the simulated 60% light level of the dilution experiments. Despite the high production rates, DMSPp accumulation was suppressed in part by microzooplankton grazers who consumed between 61% d −1 and 126% d −1 of the DMSPp production. Temporal trends in the rates of production and consumption illustrated a close coupling between the DMSP-producing phytoplankton and their microzooplankton grazers. Similar grazing and production rates were observed for the eukaryotic picophytoplankton that dominated the phytoplankton biomass, partial evidence that picoeukaryotes contributed a substantial proportion of the DMSP synthesis. These rates for DMSPp and picoeukaryotes were considerably higher than for chlorophyll a, indicating higher cycling rates of the DMSP-producing taxa than for the bulk phytoplankton community. When compared to the total phytoplankton community, there was no evidence of selection against the DMSP-containing phytoplankton by the microzooplankton grazers; the opposite appeared to be the case. SAGE demonstrated that increased iron availability in the HNLC waters of the Southern Ocean does not invariably lead to enhanced DMS sea–air flux. The potential suppression of DMSPp accumulation by grazers needs to be taken into account in future attempts to elevate DMS emission through in situ iron fertilisation and in understanding the hypothesised link between levels of Aeolian iron deposition in the Southern Ocean, DMS emission and global albedo.

Growth and grazing of microzooplankton in response to the harmful alga Heterosigma akashiwo in prey mixtures

Microzooplankton grazers may play a role in regulating blooms of the ichthyotoxic alga Heterosigma akashiwo. This study tested the effects of H. akashiwo, when part of a mixed-prey assemblage, on the growth and feeding of microzooplankton. Laboratory cultures of 3 ciliates, Favella sp., Metacylis sp., and Strombidinopsis acuminatum, were exposed to reciprocal concentra- tions of H. akashiwo and a non-toxic prey, at saturating prey concentrations. Heterosigma akashiwo was toxic to Favella sp. and Metacylis sp. when H. akashiwo was the sole prey species; however, this toxicity was eliminated in the mixed-prey treatments, likely because of avoidance of H. akashiwo and selective feeding on non-toxic prey. In contrast, the growth rate of S. acuminatum was unaffected by H. akashiwo. Both Favella sp. and S. acuminatum ingested H. akashiwo but selected against the alga when other prey was available. In addition, natural planktonic communities, collected from East Sound, Orcas Island, northern Puget Sound, in September and October 2007, were exposed to bloom- level concentrations of H. akashiwo. Abundances of the major microzooplankton types, primarily cil- iates and Gyrodinium/Gymnodinium dinoflagellates, were unaffected by H. akashiwo, although slight changes in grazer size structure did occur. Heterosigma akashiwo was harmful to the smallest grazers, mainly aloricate ciliates and small Gyrodinium/Gymnodinium dinoflagellates, and beneficial to larger Gyrodinium/Gymnodinium dinoflagellates that were able to ingest and grow on the alga. The alga was not consumed by the majority of grazers. Preferential feeding on alternate prey reduces toxic effects of H. akashiwo on microzooplankton. Avoidance of H. akashiwo by a major group of grazers would promote bloom formation by reducing H. akashiwo mortality and focusing community grazing pressure on potential competitor species.