Copepods induce paralytic shellfish toxin production in marine dinoflagellates

Among the thousands of unicellular phytoplankton species described in the sea, some frequently occurring and bloom-forming marine dinoflagellates are known to produce the potent neurotoxins causing paralytic shellfish poisoning. The natural function of these toxins is not clear, although they have been hypothesized to act as a chemical defence towards grazers. Here, we show that waterborne cues from the copepod Acartia tonsa induce paralytic shellfish toxin (PST) production in the harmful algal bloom-forming dinoflagellate Alexandrium minutum. Induced A. minutum contained up to 2.5 times more toxins than controls and was more resistant to further copepod grazing. Ingestion of non-toxic alternative prey was not affected by the presence of induced A. minutum. The ability of A. minutum to sense and respond to the presence of grazers by increased PST production and increased resistance to grazing may facilitate the formation of harmful algal blooms in the sea.     

A multi-phylum study of grazer-induced paralytic shellfish toxin production in the dinoflagellate Alexandrium fundyense: A new perspective on control of algal toxicity

The present study surveyed grazer-induced stimulation of paralytic shellfish toxin (PST) production by the marine dinoflagellate Alexandrium fundyense. The survey included species, known to graze upon A. fundyense, from five phyla: the protists, Polykrikos kofoidii (Dinoflagellata) and Tiarina fusus (Ciliophora), the bivalve molluscs Mytilus edulis and Mya arenaria (Mollusca), the ascidians, Molgula manhattensis and Botrylloides violaceus (Chordata), and the copepod, Eurytemora herdmani (Arthropoda). Direct (grazers in contact with cells of A. fundyense) and indirect (grazers not in contact with cells of A. fundyense) induction assays were carried out with protists and copepods. Only indirect assays were carried out with molluscs and ascidians. Indirect assays also tested whether induction of PST production occurred via kairomones or feeding-related cues. All metazoan grazers induced PST production. By contrast, neither of the two species of protistan grazer induced PST production. Direct and indirect inductions of PST production were evident for the copepod, with direct induction being significantly higher than indirect induction. Effects upon PST production by phylum, species (nested within phylum), and interactions of phylum by diet, and species by diet, were evident. When induction of PST production occurred, a kairomone effect was apparent, except for M. edulis. Similarly, feeding-related cues were evident, except for E. herdmani. An asymptotic relationship between the magnitude of indirect induction of PST production and total cell ingestion by the grazers suggests a saturation response of grazer-induced PST production.     

Grazers and Phytoplankton Growth in the Oceans: an Experimental and Evolutionary Perspective

The taxonomic composition of phytoplankton responsible for primary production on continental shelves has changed episodically through Earth history. Geological correlations suggest that major changes in phytoplankton composition correspond in time to changes in grazing and seawater chemistry. Testing hypotheses that arise from these correlations requires experimentation, and so we carried out a series of experiments in which selected phytoplankton species were grown in treatments that differed with respect to the presence or absence of grazers as well as seawater chemistry. Both protistan (Euplotes sp.) and microarthropod (Acartia tonsa) grazers changed the growth dynamics and biochemical composition of the green alga Tetraselmis suecica, the diatom Thalassiosira weissflogii, and the cyanobacterium Synechococcus sp., increasing the specific growth rate and palatability of the eukaryotic algae, while decreasing or leaving unchanged both parameters in the cyanobacteria. Synechococcus (especially) and Thalassiosira produced toxins effective against the copepod, but ciliate growth was unaffected. Acartia induced a 4-6 fold increase of Si cell quota in the diatom, but Euplotes had no similar effect. The differential growth responses of the eukaryotic algae and cyanobacteria to ciliate grazing may help to explain the apparently coeval radiation of eukaryophagic protists and rise of eukaryotes to ecological prominence as primary producers in Neoproterozoic oceans. The experimental results suggest that phytoplankton responses to the later radiation of microarthropod grazers were clade-specific, and included changes in growth dynamics, toxin synthesis, encystment, and (in diatoms) enhanced Si uptake.     

Grazing on a toxic Alexandrium catenella bloom by the lobster krill Munida gregaria (Decapoda: Galatheoidea: Munididae)

Specimens of Munida gregaria were collected within and in the vicinity of a bloom of the toxic dinoflagellate Alexandrium catenella in Queen Charlotte Sound, New Zealand. The crustacean contained paralytic shellfish toxin (PST) with an analogue profile dominated by N-sulfocarbamoyl analogues (C1,2 and GTX5) and carbamate gonyautoxins (GTX1,4), similar to that of the dinoflagellate. A feeding experiment showed that M. gregaria is capable of actively grazing on A. catenella and it may play a role in controlling population growth of the dinoflagellate. This is the first account of the accumulation of PST by M. gregaria. When it is periodically abundant, M. gregaria is an important food item for fish, birds and other marine fauna and they are a vector by which PST may be transferred to higher trophic levels.     

Copepod feeding response to varying Alexandrium spp. cellular toxicity and cell concentration among natural plankton samples

We challenged four species of copepod grazers (Acartia hudsonica, Centropages hamatus, Eurytemora herdmani, Calanus finmarchicus) with natural water samples containing non-toxic algae mixed with one of three clones of Alexandrium spp.—A. tamarense GTCN16 (non-toxic), A. fundyense GTCA28 (moderate toxicity), and A. fundyense BC1 (higher toxicity), each at relatively high (10⁵ cells L⁻¹) and low (10⁴ cells L⁻¹) concentrations. Within any one copepod species, significant differences existed in copepod clearance rates and total food ingested between high and low Alexandrium cell concentrations, and between levels of toxicity, but feeding response did not follow a predictable relationship proportional to toxin levels—rather, the presence or absence of toxin was more important than the level of toxicity. C. finmarchicus behaved differently from the smaller coastal copepods, showing less selectivity and greater concentration dependence. In low Alexandrium concentration treatments, copepod clearance rates on Alexandrium were usually higher, and electivity indices for Alexandrium less negative (indicating less avoidance), compared to high concentration treatments. In high toxicity (BC1) treatments of all copepod species (except C. finmarchicus), total food consumption was consistently less at high Alexandrium concentrations compared to low concentrations, suggesting that high toxicity and concentration suppress overall feeding, while in non-toxic (GTCN 16) treatments total consumption was always higher at high Alexandrium concentrations. Copepod grazers do not follow predictable feeding rules throughout a continuum of conditions, but become more predictable at extremes of concentration and toxicity of prey, consistent with the conclusion that both factors are important. Results support the hypothesis that grazer deterrence imparted by toxicity is only effective at high cell concentrations, but even then will not protect against all grazers.     

Relative importance of nitrogen sources,algal alarm cues and grazer exposure to toxin production of the marine dinoflagellate Alexandrium catenella

Dinoflagellate paralytic shellfish toxin (PST) production is mediated by several abiotic and biotic factors. This study compared the relative importance of nitrogen source and concentration, prey alarm cues and grazer presence on toxin production of the marine dinoflagellate Alexandrium catenella (Group I, strain BF-5). In separate assays run under either nutrient-replete (F/2 medium) or nutrient-depleted (filtered seawater) conditions, PST production of A. catenella was measured as a function of varying concentrations of added nitrogen sources (ammonium and urea), alarm cues from lysed conspecific (A. catenella Group I strains) and interspecific (the diatom, Thalassiosira weissflogii, and the green flagellate, Tetraselmis sp.) algae, and the presence of a grazer (the copepod Acartia hudsonica). Results showed that addition of ammonium or urea did not increase PST production. Unexpectedly, interspecific alarm cues increased toxin production but conspecific ones did not. Grazer presence dramatically induced PST production in A. catenella, irrespective of nutrient conditions, and this effect was an order of magnitude greater than any of the other variables tested. These results corroborate previous studies on grazer-induced PST production, and support the hypothesis that grazer-induced toxin production is not an experimental artifact, but rather a prey defense mechanism.     

Induction of toxin production in dinoflagellates: the grazer makes a difference

The dinoflagellate Alexandrium minutum has previously been shown to produce paralytic shellfish toxins (PST) in response to waterborne cues from the copepod Acartia tonsa. In order to investigate if grazer-induced toxin production is a general or grazer-specific response of A. minutum to calanoid copepods, we exposed two strains of A. minutum to waterborne cues from three other species of calanoid copepods, Acartia clausi, Centropages typicus and Pseudocalanus sp. Both A. minutum strains responded to waterborne cues from Centropages and Acartia with significantly increased cell-specific toxicity. Waterborne cues from Centropages caused the strongest response in the A. minutum cells, with 5 to >20 times higher toxin concentrations compared to controls. In contrast, neither of the A. minutum strains responded with significantly increased toxicity to waterborne cues from Pseudocalanus. The absolute increase in PST content was proportional to the intrinsic toxicity of the different A. minutum strains that were used. The results show that grazer-induced PST production is a grazer-specific response in A. minutum, and its potential ecological importance will thus depend on the composition of the zooplankton community, as well as the intrinsic toxin-producing properties of the A. minutum population.     

Effect of the endoparasite Amoebophrya sp. on toxin content and composition in the paralytic shellfish poisoning dinoflagellate Alexandrium fundyense (Dinophyceae)

Members of the Amoebophrya ceratii complex are endoparasitic dinoflagellates that parasitize a number of their dinoflagellate relatives, including toxic and/or harmful algal bloom-forming species. Despite many studies on the occurrence, prevalence, biology and molecular phylogeny of Amoebophrya spp., little attention has been given to toxin dynamics of host population following parasitism. Using Amoebophrya sp. infecting the paralytic shellfish toxin (PSP)-producing dinoflagellate Alexandrium fundyense, we addressed the following questions: (1) does parasitism by Amoebophrya sp. alter toxin content and toxin profiles of the dinoflagellate A. fundyense over the infection cycle? and (2) do parasite dinospores produced at the end of the infection cycle retain host toxins and thus potentially act as a vector to convey PSP toxin through the marine microbial food-web? Toxin time-course experiments showed that the PSP toxin contents did not vary significantly over the infection cycle, but mean toxin content for infected cultures was significantly higher than that for uninfected cultures. Host toxins were not detected in the free-living, dinospore stage of the parasite. Therefore, our results indicate that Amoebophrya sp. does not function as a vector for transferring PSP toxins to higher trophic levels. Rather, Amoebophrya infections appear to play an important role in maintaining healthy ecosystems by transforming potent toxins-producing dinoflagellates into non-toxic dinospores, representing “edible food” for consumers of the marine microbial food-web during toxic algal bloom event.     

Chemical Response of the Toxic Dinoflagellate Karenia mikimotoi Against Grazing by Three Species of Zooplankton

We investigated the toxicity of Karenia mikimotoi toward three model grazers, the cladoceran Moina mongolica, the copepod Pseudodiaptomus annandalei, and the crustacean Artemia salina, and explored its chemical response upon zooplankton grazing. An induction experiment, where K. mikimotoi was exposed to grazers or waterborne cues from the mixed cultures revealed that K. mikimotoi might be toxic or nutritionally inadequate toward the three grazers. In general, direct exposure to the three grazers induced the production of hemolytic toxins and the synthesis of eicosapentaenoic acid (EPA). Both EPA and the hemolytic toxins from K. mikimotoi decreased the survival rate of the three grazers. In addition, the survival rates of M. mongolica, P. annandalei, and A. salina in the presence of induced K. mikimotoi that had previously been exposed to a certain grazer were lower than their counterparts caused by fresh K. mikimotoi, suggesting that exposure to some grazers might increase the toxicity of K. mikimotoi. The chemical response and associated increased resistance to further grazing suggested that K. mikimotoi could produce deterrents to protect against grazing by zooplankton and that the substances responsible might be hemolytic toxins and EPA.     

Bloom-Forming Cyanobacteria Support Copepod Reproduction and Development in the Baltic Sea

It is commonly accepted that summer cyanobacterial blooms cannot be efficiently utilized by grazers due to low nutritional quality and production of toxins; however the evidence for such effects in situ is often contradictory. Using field and experimental observations on Baltic copepods and bloom-forming diazotrophic filamentous cyanobacteria, we show that cyanobacteria may in fact support zooplankton production during summer. To highlight this side of zooplankton-cyanobacteria interactions, we conducted: (1) a field survey investigating linkages between cyanobacteria, reproduction and growth indices in the copepod Acartia tonsa; (2) an experiment testing relationships between ingestion of the cyanobacterium Nodularia spumigena (measured by molecular diet analysis) and organismal responses (oxidative balance, reproduction and development) in the copepod A. bifilosa; and (3) an analysis of long term (1999–2009) data testing relationships between cyanobacteria and growth indices in nauplii of the copepods, Acartia spp. and Eurytemora affinis, in a coastal area of the northern Baltic proper. In the field survey, N. spumigena had positive effects on copepod egg production and egg viability, effectively increasing their viable egg production. By contrast, Aphanizomenon sp. showed a negative relationship with egg viability yet no significant effect on the viable egg production. In the experiment, ingestion of N. spumigena mixed with green algae Brachiomonas submarina had significant positive effects on copepod oxidative balance, egg viability and development of early nauplial stages, whereas egg production was negatively affected. Finally, the long term data analysis identified cyanobacteria as a significant positive predictor for the nauplial growth in Acartia spp. and E. affinis. Taken together, these results suggest that bloom forming diazotrophic cyanobacteria contribute to feeding and reproduction of zooplankton during summer and create a favorable growth environment for the copepod nauplii.     

The copepod Calanus finmarchicus: A potential vector for trophic transfer of the marine algal biotoxin, domoic acid

The marine algal biotoxin, domoic acid (DA), is produced by certain members of the diatom genus Pseudo-nitzschia. This neurotoxin has been responsible for several mass mortality events involving marine birds and mammals. In all cases, the toxin was transferred from its algal producers through marine food webs by one or more intermediate vectors. The ability of some copepod taxa to serve as vectors for DA has been demonstrated; however, the role played in DA trophic transfer by Calanus finmarchicus, which often dominates N. Atlantic zooplankton assemblages and is a primary dietary component of the highly endangered N. Atlantic right whale (Eubalaena glacialis), has been uncertain. In the present study, we examined the ability of C. finmarchicus to consume DA-producing algae and retain the toxin. Results of grazing and toxin accumulation/depuration experiments showed that C. finmarchicus consumed DA-producing Pseudo-nitzschia multiseries regardless of the presence or absence of morphologically similar, but non-toxic, P. pungens, across initial cell concentrations ranging from 1000-4000 cells mL^− 1. Furthermore, C. finmarchicus did not appear to preferentially consume or avoid either Pseudo-nitzschia species tested. After ingestion of P. multiseries, copepods accumulated DA and retained it for up to 48 h post-removal of the toxin source. These findings provide evidence for the potential of C. finmarchicus to facilitate DA trophic transfer in marine food webs where toxic Pseudo-nitzschia is present.     

Planktonic marine copepods and harmful algae

Marine planktonic copepods are important grazers on harmful algae (HA) species of phytoplankton, and copepods are major entry points for vectorial intoxication of pelagic food webs with HA toxins. Previous reviews (Turner and Tester, 1997, Turner et al., 1998a, Turner, 2006) summarized information on HA interactions with zooplankton grazers, and vectorial intoxication of pelagic food webs, up through approximately 2005. Accordingly, this review will address primarily studies published during the last decade. It will concentrate on generic issues in the developing field of HA:grazer interactions, such as the extent to which HA toxins serve as copepod grazing deterrents, induction of HA grazing deterrents by exposure to copepods, copepod selective feeding to avoid ingesting HA taxa versus non-selective feeding on HA taxa, possible biogeographic aspects of the effects of HA toxins on copepods, impact of copepod grazing on HA bloom development and termination, the role of copepods as entry points for vectorial intoxication of pelagic food webs with HA toxins, and possible reasons and remedies for the highly-variable and conflicting results reported for many studies of copepod grazing on various HA species.     

Phycotoxin accumulation in zooplankton feeding on Alexandrium fundyense--vector or sink?

Zooplankton can consume toxic Alexandrium spp. dinoflagellatesin the Gulf of Maine and retain paralytic shellfish poisoning(PSP) toxins, potentially acting as toxin vectors. We performedexperiments to determine toxin budgets for common species ofcopepods (Acartia hudsonica, Eurytemora herdmani, Centropageshamatus) feeding on toxic Alexandrium fundyense, offered asmonocultures or in mixtures of algal prey, by comparing calculatedtoxin ingestion rates and toxin content of copepod body tissueand fecal pellets. When fed monocultures, both copepod tissueand fecal pellet fractions accounted for 5% each of the calculatedingested toxin, and thus by difference 90% was lost as a dissolvedfraction into the seawater medium. The presence of alternativefood did not significantly alter the efficiency of toxin retention.Sloppy feeding or regurgitation are probable mechanisms forrelease of toxin to sea water. Experiments using varying concentrationsof A. fundyense and alternativenon-toxic species did not showsignificant effects of cell concentration on toxin retentionefficiency. Total toxin retained and efficiency of retentionvaried among copepod species. Toxin profiles (% molar composition)of dinoflagellates, copepod tissues and fecal pellets differedslightly, suggesting some metabolic transformation. Becauseof their low retention efficiency, copepod grazers can effectivelydisperse PSP toxins produced by Alexandrium spp. into the environment,where they are much less likely to be harmful—zooplanktonact as a sink for PSP toxins. Nevertheless, sufficient toxinbody burdens are attained to contribute to propagation of PSPtoxins to other trophic levels.     

Detection of cyanobacterial sxt genes and paralytic shellfish toxins in freshwater lakes and brackish waters on Åland Islands,Finland

Harmful cyanobacteria are a globally growing concern. They produce a large variety of toxic compounds, including saxitoxin and its many structural variants, a group of potent neurotoxins collectively called paralytic shellfish toxins or PST. Nucleic acid based detection methods, such as qPCR, have been proposed as potential screening and monitoring tools for toxic cyanobacteria, but it is not clear how well the presence and quantity of saxitoxin biosynthesis (sxt) genes can be used to predict the production of PST in the environment. In this study, the prevalence of three sxt genes and their co-occurrence with paralytic shellfish toxins in the environment was investigated. The sxtA, sxtG and sxtB genes were present on average in 31% of the samples collected from lakes and brackish coastal waters on Åland Islands, Finland, during the three-year monitoring period. PST detection frequency varied from 13% to 59% from year to year, and concentrations were generally low. On average higher sxtB copy numbers were associated with PST detection, and although a positive correlation between gene copy numbers and toxin concentrations was observed (Spearman rank correlation, ρ = 0.53, P = 0.012), sxt gene presence or quantity didn’t reliably predict PST production. Sequencing of sxtA fragments and identification of main cyanobacteria indicated that the likely candidate responsible for PST production in the samples belonged to the genus Anabaena.     

No evidence for induction or selection of mutant sodium channel expression in the copepod Acartia husdsonica challenged with the toxic dinoflagellate Alexandrium fundyense

Some species in the dinoflagellate genus Alexandrium spp. produce a suite of neurotoxins that block sodium channels, known as paralytic shellfish toxins (PST), which have deleterious effects on grazers. Populations of the ubiquitous copepod grazer Acartia hudsonica that have co‐occurred with toxic Alexandrium spp. are better adapted than naïve populations. The mechanism of adaptation is currently unknown. We hypothesized that a mutation in the sodium channel could account for the grazer adaptation. We tested two hypotheses: (1) Expression of the mutant sodium channel could be induced by exposure to toxic Alexandrium fundyense; (2) in the absence of induction, selection exerted by toxic A. fundyense would favor copepods that predominantly express the mutant isoform. In the copepod A. hudsonica, both isoforms are expressed in all individuals in varying proportions. Thus, in addition to comparing expression ratios of wild‐type to mutant isoforms for individual copepods, we also partitioned copepods into three groups: those that predominantly express the mutant (PMI) isoform, the wild‐type (PWI) isoform, or both isoforms approximately equally (EI). There were no differences in isoform expression between individuals that were fed toxic and nontoxic food after three and 6 days; induction of mutant isoform expression did not occur. Furthermore, the hypothesis that mutant isoform expression responds to toxic food was also rejected. That is, no consistent evidence showed that the wild‐type to mutant isoform ratios decreased, or that the relative proportion of PMI individuals increased, due to the consumption of toxic food over four generations. However, in the selected line that was continuously exposed to toxic food sources, egg production rate increased, which suggested that adaptation occurred but was unrelated to sodium channel isoform expression.     

Copepod grazing during an iron-induced diatom bloom in the Antarctic Circumpolar Current (EisenEx): I. Feeding patterns and grazing impact on prey populations

Feeding activity, selective grazing and the potential grazing impact of two dominant grazers of the Polar Frontal Zone, Calanus simillimus and Rhincalanus gigas, and of copepods < 2 mm were investigated with incubation experiments in the course of an iron fertilized diatom bloom in November 2000. All grazers were already actively feeding in the low chlorophyll waters prior to the onset of the bloom. C. simillimus maintained constant clearance rates and fed predominantly on diatoms. R. gigas and the small copepods strongly increased clearance and ingestion of diatoms in response to their enhanced availability. All grazers preyed on microzooplankton, most steadily on ciliates, confirming the view that pure herbivory appears to be the exception rather than the rule in copepod feeding. The grazers exhibited differences in feeding behavior based on selectivity indices. C. simillimus and R. gigas showed prey switching from dinoflagellates to diatoms in response to the phytoplankton bloom. All grazers most efficiently grazed on large diatoms leading to differences in daily losses for large and small species, e.g. Corethron sp. or Thalassionema nitzschioides. Species-specific diatom mortality rates due to grazing suggest that the high feeding activity of C. simillimus prior to and during the bloom played a role in shaping diatom population dynamics.     

Paralytic shellfish toxins or spirolides? The role of environmental and genetic factors in toxin production of the Alexandrium ostenfeldii complex

Dinoflagellates of the Alexandrium ostenfeldii complex (A. ostenfeldii, A. peruvianum) are capable of producing different types of neurotoxins: paralytic shellfish toxins (PSTs), spirolides and gymnodimines, depending on the strain and its geographic origin. While Atlantic and Mediterranean strains have been reported to produce spirolides, strains originating from the brackish Baltic Sea produce PSTs. Some North Sea, USA and New Zealand strains contain both toxins. Causes for such intraspecific variability in toxin production are unknown. We investigated whether salinity affects toxin production and growth rate of 5 A. ostenfeldii/peruvianum strains with brackish water (Baltic Sea) or oceanic (NE Atlantic) origin. The strains were grown until stationary phase at 7 salinities (6–35), and their growth and toxin production was monitored. Presence of saxitoxin (STX) genes (sxtA1 and sxtA4 motifs) in each strain was also analyzed. Salinity significantly affected both growth rate and toxicity of the individual strains but did not change their major toxin profile. The two Baltic Sea strains exhibited growth at salinities 6–25 and consistently produced gonyautoxin (GTX) 2, GTX3 and STX. The two North Sea strains grew at salinities 20–35 and produced mainly 20-methyl spirolide G (20mG), whereas the strain originating from the northern coast of Ireland was able to grow at salinities 15–35, only producing 13-desmethyl spirolide C (13dmC). The effects of salinity on total cellular toxin concentration and distribution of toxin analogs were strain-specific. Both saxitoxin gene motifs were present in the Baltic Sea strains, whereas the 2 North Sea strains lacked sxtA4, and the Irish strain lacked both motifs. Thus sxtA4 only seems to be specific for PST producing strains. The results show that toxin profiles of A. ostenfeldii/peruvianum strains are predetermined and the production of either spirolides or PSTs cannot be induced by salinity changes. However, changes in salinity may lead to changed growth rates, total cellular toxin concentrations as well as relative distribution of the different PST and spirolide analogs, thus affecting the actual toxicity of A. ostenfeldii/peruvianum populations.     

Effects of temperature and salinity on the growth of Alexandrium (Dinophyceae) isolates from the Salish Sea

Toxin‐producing blooms of dinoflagellates in the genus Alexandrium have plagued the inhabitants of the Salish Sea for centuries. Yet the environmental conditions that promote accelerated growth of this organism, a producer of paralytic shellfish toxins, is lacking. This study quantitatively determined the growth response of two Alexandrium isolates to a range of temperatures and salinities, factors that will strongly respond to future climate change scenarios. An empirical equation, derived from observed growth rates describing the temperature and salinity dependence of growth, was used to hindcast bloom risk. Hindcasting was achieved by comparing predicted growth rates, calculated from in situ temperature and salinity data from Quartermaster Harbor, with corresponding Alexandrium cell counts and shellfish toxin data. The greatest bloom risk, defined at μ >0.25 d^?1, generally occurred from April through November annually; however, growth rates rarely fell below 0.10 d^?1. Except for a few occasions, Alexandrium cells were only observed during the periods of highest bloom risk and paralytic shellfish toxins above the regulatory limit always fell within the periods of predicted bloom occurrence. While acknowledging that Alexandrium growth rates are affected by other abiotic and biotic factors, such as grazing pressure and nutrient availability, the use of this empirical growth function to predict higher risk time frames for blooms and toxic shellfish within the Salish Sea provides the groundwork for a more comprehensive biological model of Alexandrium bloom dynamics in the region and will enhance our ability to forecast blooms in the Salish Sea under future climate change scenarios.     

Zooplankton community grazing impact on a bloom of Alexandrium fundyense in the Gulf of Maine

Shipboard grazing experiments were conducted in the Gulf of Maine and on Georges Bank during of June 2006 to estimate zooplankton community grazing impact on a natural bloom of the toxic dinoflagellate Alexandrium fundyense. Surface seawater samples containing natural populations of grazers and A. fundyense from 23 stations were incubated at ambient temperatures. Concentrations of A. fundyense after incubations were compared to those at the start of each experiment to determine net increases due to population growth, or decreases presumed to be primarily due to grazing losses. Abundances of both microzooplankton (tintinnids, oligotrich ciliates, rotifers, copepod nauplii and heterotrophic dinoflagellates) and mesozooplankton (copepod nauplii, copepodites and adult copepods, rotifers, marine cladocerans, and meroplankton) grazers in experimental aliquots were also determined. The total zooplankton community had minimal grazing impact on natural populations of A. fundyense at most stations. At 70% of the stations where grazing experiments were performed, there were no significant differences in initial and final concentrations of A. fundyense. This indicated that growth of, and grazing on A. fundyense were in approximate balance. At 2 stations, which had the highest A. fundyense abundances of the cruise (>10⁴ cells l⁻¹), % of the A. fundyense population grazed per day was significantly negative, indicating that net population growth of A. fundyense exceeded grazing losses. At 5 stations, which had low concentrations of A. fundyense (10²–10³ cells l⁻¹), % of the A. fundyense population grazed per day was significantly positive, indicating that losses of A. fundyense due to grazing exceeded net population growth. For stations with significant differences between Initial and Grazed concentrations of A. fundyense, grazing had the greatest impact at lower concentrations of A. fundyense, and grazing impact by the larger mesozooplankton was inversely related to zooplankton abundance. There was no relationship between microzooplankton abundance and grazing impact on A. fundyense. Grazing exceeded growth only where A. fundyense abundance was low, and growth exceeded grazing only where A. fundyense abundance was high. The inverse relationship between grazing impact and A. fundyense abundance implies that grazing may be capable of retarding bloom development at low concentrations typical of the early stages of a bloom, but at higher concentrations once a bloom becomes established, either grazing maintains a balance with A. fundyense growth, or growth exceeds grazing losses at highest concentrations.