Lipid-fuelled overwintering by copepods can be a regionally important contribution to carbon sequestration in the deep oceans. Here, we estimate the contribution for Calanus hyperboreus, found in abundance in the northern reaches of the North Atlantic and Arctic Ocean. Estimates for regions with high overwintering populations, Fram Strait, Greenland Sea and Iceland Sea lie between 3.5 gC m 22 yr 21 and 6.0 gC m 22 yr 21 at depths of 1000-3000 m, comparable to the flux of detrital organic carbon at commensurate depths. Apart from the variation in the abundance of overwintering populations, these estimates are most sensitive to mortality rates. We present a general model based on metabolic theory and isomorphism that can be used to constrain estimates for data poor species in other parts of the global ocean.
Diatoms account for 40% of the ocean primary production and play a key role in the oceans’ ability to sequester carbon. The evolutionary success of diatoms and their role in ocean biogeochemistry are related to the siliceous shell that provide partial protection against grazing. The structure and function of phytoplankton communities are governed by environmental constraints and organismal trade‐offs. Defence mechanisms may help explain the high diversity of phytoplankton (incl. diatoms) in the ocean, but only if the defence comes at a cost. Defence costs have been notoriously difficult to demonstrate and quantify in marine phytoplankton. Here, we demonstrate for seven species of planktonic diatoms that their shell thickens and their growth rate declines when cells are exposed to chemical cues from copepods, important predators of diatoms. The responses are proportional to the concentration of grazer cues, but are also highly variable, both between and within species. At our standard experimental condition, the typical decline in growth rate is 10%, and the typical increase in cellular biogenic silica is 16%. The latter value corresponds to a decline in grazing mortality due to small copepods of 11%. Thus, silification in response to grazers is exactly warranted. The similar magnitude of the costs and benefits of silification suggests a flat fitness landscape along the competition‐defence axis. This may help explain the high diversity of coexisting diatoms in the ocean. The significant but variable contribution of diatoms to the downward flux of organic carbon in the ocean depends to a large extent on the silica content of the cells. This is due less to the ballasting effect of silica, but mainly to the different life histories of more or less defended cells that are governed by evolutionary adaptations and—as demonstrated here—plastic responses to grazers. A free Plain Language Summary can be found within the Supporting Information of this article.
The calanoid copepod Pseudodiaptomus annandalei is used as live feed in aquaculture because of its nutritional value and the ability to cope with environmental fluctuations in outdoor ponds. However, little knowledge exists on its ecology. Here we investigated the ecology of P. annandalei in an aquaculture pond in Vietnam. Temperature, salinity, chlorophyll a and biomass of protozoans and copepods were monitored every other day for 1 month. Experiments on protozoan growth and grazing by P. annandalei were also conducted. Copepod fecal pellet production and temperature-dependent egg hatching rates were likewise quantified. Despite very high phytoplankton biomass, biomass of P. annandalei was surprisingly low. Copepod production was estimated from three independent methods: clearance, weight-specific egg production rate (SEP) and specific fecal pellet production rate. SEP proved to be accurate to predict the in situ population growth in the pond. A simple model for production of P. annandalei based on SEP was developed. Our study extends our knowledge of how environmental conditions in the pond may affect the population dynamics and production of copepods. The results have important implications for pond managements ensuring stable copepod production and harvest.
The diatom frustule provides partial protection against copepod grazing. Whether the defense is due to the cells being de‐selected or handled for so long that the grazers lose time for foraging is unknown. The mechanism has implications for the population dynamics of both defended and co‐occurring, undefended nutrient competitors. We use video‐observations to demonstrate that thick‐shelled diatoms captured by the copepod Temora longicornis were rejected more frequently than thin‐shelled diatoms, irrespective of cell size. The thick‐shelled cells of the larger diatoms were handled for much longer, and the time spent handling these limits the consumption of phytoplankton. This may be why many diatoms, even in the presence of dense grazer populations, reach bloom concentrations, and thus, facilitate aggregation and mass sedimentation. This has implications for both carbon sequestration and for securing a large population of cells at depth ready to colonize the pelagic, when growth conditions again become favorable.
Phytoplankton induce defensive traits in response to chemical alarm signals from grazing zooplankton. However, these signals are potentially vulnerable to changes in pH and it is not yet known how predator recognition may be affected by ocean acidification. We exposed four species of diatoms and one toxic dinoflagellate to future pCO2 levels, projected by the turn of the century, in factorial combinations with predatory cues from copepods (copepodamides). We measured the change in growth, chain length, silica content, and toxin content. Effects of increased pCO2 were highly species specific. The induction of defensive traits was accompanied by a significant reduction in growth rate in three out of five species. The reduction averaged 39% and we interpret this as an allocation cost associated with defensive traits. Copepodamides induced significant chain length reduction in three of the four diatom species. Under elevated pCO2Skeletonema marinoi reduced silica content by 30% and in Alexandrium minutum the toxin content was reduced by 30%. Using copepodamides to induce defensive traits in the absence of direct grazing provides a straightforward methodology to assess costs of defense in microplankton. We conclude that copepodamide signalling system is likely robust to ocean acidification. Moreover, the variable responses of different taxa to ocean acidification suggest that there will be winners and losers in a high pCO2 world, and that ocean acidification may have structuring effects on phytoplankton communities.
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