Hans Henrik Jakobsen

Grazer cues induce stealth behavior in marine dinoflagellates

Chain formation is common among phytoplankton organisms but the underlying reasons and consequences are poorly understood. Here we show that chain formation is strongly impaired by waterborne cues from copepod grazers in the dinoflagellate Alexandrium tamarense. Chains of Alexandrium cells exposed to copepod cues responded by splitting into single cells or shorter chains. Motion analysis revealed significantly lower swimming velocities for single cells compared with chains, with two- to fivefold higher simulated predator encounter rates for two- and four-cell chains, respectively. In addition, the few remaining two-cell chains in grazed treatments were swimming at approximately half the speed of two-cell chains in treatments without grazers, which reduced encounter rates with grazers to values similar to that of single cells. Chain length plasticity and swimming behavior constitute unique mechanisms to reduce encounters with grazers. We argue that dinoflagellates can regulate the balance between motility and predator avoidance by adjusting chain length. The high predator encounter rate for motile chains may have contributed to the low prevalence of chain formation in motile phytoplankton compared with in nonmotile phytoplankton where chain formation is more common.

High resilience of two coastal plankton communities to twenty-first century seawater acidification: Evidence from microcosm studies

Abstract Increased free CO2 and ocean acidification are among the consequences of anthropogenic carbon emissions. Responses of marine protists to increased levels of CO2 are highly species-specific, and this has been suggested to cause an alteration in plankton species composition, community functions and ultimately biogeochemical cycles. This study aims to test this by performing microcosm incubation experiments at present (pH 8.0) and at three lowered pH levels (pH 7.8, 7.6 and 6.0), corresponding to free CO2 concentrations of 24, 38, 58 & 610 µmol l−1, respectively. Results from two such experiments are reported, and measurements include microscopy counts of ~20 planktonic protist taxa, HPLC pigment analysis, FlowCAM analysis of cell-size spectra, photosynthetic activity and total POC and PON. Initial communities were flagellate (experiment 1) and dinoflagellate and ciliate (experiment 2) dominated, but at pH 8.0, 7.8 and 7.6 a diatom dominance developed during the first days in both experiments. Profound changes in all measured parameters were found as a result of the pH 6.0 treatment, but among the other three pH treatments significant differences were few; Karlodinium spp. was the only taxon to be affected significantly, and plankton group composition, cell sizes and photosynthetic activity all remained unaffected during the 14-day incubation periods. Thus, both of the investigated coastal plankton communities were unaffected by twenty-first century expected changes in pH and free CO2. This may be explained by the large seasonal, and even daily, changes in pH seen in productive marine ecosystems, and the corresponding need for algae to be pH-tolerant.

Unsteady motion: escape jumps in planktonic copepods, their kinematics and energetics.

We describe the kinematics of escape jumps in three species of 0.3–3.0 mm-sized planktonic copepods. We find similar kinematics between species with periodically alternating power strokes and passive coasting and a resulting highly fluctuating escape velocity. By direct numerical simulations, we estimate the force and power output needed to accelerate and overcome drag. Both are very high compared with those of other organisms, as are the escape velocities in comparison to startle velocities of other aquatic animals. Thus, the maximum weight-specific force, which for muscle motors of other animals has been found to be near constant at 57 N (kg muscle)−1, is more than an order of magnitude higher for the escaping copepods. We argue that this is feasible because most copepods have different systems for steady propulsion (feeding appendages) and intensive escapes (swimming legs), with the muscular arrangement of the latter probably adapted for high force production during short-lasting bursts. The resulting escape velocities scale with body length to power 0.65, different from the size-scaling of both similar sized and larger animals moving at constant velocity, but similar to that found for startle velocities in other aquatic organisms. The relative duration of the pauses between power strokes was observed to increase with organism size. We demonstrate that this is an inherent property of swimming by alternating power strokes and pauses. We finally show that the Strouhal number is in the range of peak propulsion efficiency, again suggesting that copepods are optimally designed for rapid escape jumps.

Mechanisms and feasibility of prey capture in ambush-feeding zooplankton

Many marine zooplankters, particularly among copepods, are “ambush feeders” that passively wait for their prey and capture them by fast surprise attacks. This strategy must be very demanding in terms of muscle power and sensing capabilities, but the detailed mechanisms of the attacks are unknown. Using high-speed video we describe how copepods perform spectacular attacks by precision maneuvering during a rapid jump. We show that the flow created by the attacking copepod is so small that the prey is not pushed away, and that the attacks are feasible because of their high velocity (≈100 mm·s−1) and short duration (few ms), which leaves the prey no time for escape. Simulations and analytical estimates show that the viscous boundary layer that develops around the attacking copepod is thin at the time of prey capture and that the flow around the prey is small and remains potential flow. Although ambush feeding is highly successful as a feeding strategy in the plankton, we argue that power requirements for acceleration and the hydrodynamic constraints restrict the strategy to larger (> 0.25 mm), muscular forms with well-developed prey perception capabilities. The smallest of the examined species is close to this size limit and, in contrast to the larger species, uses its largest possible jump velocity for such attacks. The special requirements to ambush feeders with such attacks may explain why this strategy has evolved to perfection only a few times among planktonic suspension feeders (few copepod families and chaetognaths).

Dynamics of Dissolved and Particulate Polyunsaturated Aldehydes in Mesocosms Inoculated with Different Densities of the Diatom Skeletonema marinoi

A survey of the production of polyunsaturated aldehydes (PUA) of manipulated plankton communities is presented here. PUA are phytoplankton-derived metabolites that are proposed to play an important role in chemically mediated plankton interactions. Blooms of different intensities of the diatom Skeletonema marinoi were generated in eight mesocosms filled with water from the surrounding fjord by adding different amounts of a starting culture and nutrients. This set-up allowed us to follow PUA production of the plankton community over the entire induced bloom development, and to compare it with the natural levels of PUA. We found that S. marinoi is a major source for the particulate PUA 2,4-heptadienal and 2,4-octadienal (defined as PUA released upon wounding of the diatom cells) during the entire bloom development. Just before, and during, the decline of the induced diatom blooms, these PUA were also detected in up to 1 nM concentrations dissolved in the water. In addition, we detected high levels of the PUA 2,4-decadienal that was not produced by the diatom S. marinoi. Particulate decadienal correlated well with the cell counts of the prymnesiophyte Phaeocystis sp. that also developed in the fertilized mesocosms. Particulate decadienal levels were often even higher than those of diatom-derived PUA, indicating that PUA sources other than diatoms should be considered when it comes to the evaluation of the impact of these metabolites.

Future Climate Scenarios for a Coastal Productive Planktonic Food Web Resulting in Microplankton Phenology Changes and Decreased Trophic Transfer Efficiency

We studied the effects of future climate change scenarios on plankton communities of a Norwegian fjord using a mesocosm approach. After the spring bloom, natural plankton were enclosed and treated in duplicates with inorganic nutrients elevated to pre-bloom conditions (N, P, Si; eutrophication), lowering of 0.4 pH units (acidification), and rising 3°C temperature (warming). All nutrient-amended treatments resulted in phytoplankton blooms dominated by chain-forming diatoms, and reached 13–16 μg chlorophyll (chl) a l−1. In the control mesocosms, chl a remained below 1 μg l−1. Acidification and warming had contrasting effects on the phenology and bloom-dynamics of autotrophic and heterotrophic microplankton. Bacillariophyceae, prymnesiophyceae, cryptophyta, and Protoperidinium spp. peaked earlier at higher temperature and lower pH. Chlorophyta showed lower peak abundances with acidification, but higher peak abundances with increased temperature. The peak magnitude of autotrophic dinophyceae and ciliates was, on the other hand, lowered with combined warming and acidification. Over time, the plankton communities shifted from autotrophic phytoplankton blooms to a more heterotrophic system in all mesocosms, especially in the control unaltered mesocosms. The development of mass balance and proportion of heterotrophic/autotrophic biomass predict a shift towards a more autotrophic community and less-efficient food web transfer when temperature, nutrients and acidification are combined in a future climate-change scenario. We suggest that this result may be related to a lower food quality for microzooplankton under acidification and warming scenarios and to an increase of catabolic processes compared to anabolic ones at higher temperatures.

Co-transport of polycyclic aromatic hydrocarbons by motile microorganisms leads to enhanced mass transfer under diffusive conditions.

The environmental chemodynamics of hydrophobic organic chemicals (HOCs) are often rate-limited by diffusion in stagnant boundary layers. This study investigated whether motile microorganisms can act as microbial carriers that enhance mass transfer of HOCs through diffusive boundary layers. A new experimental system was developed that allows (1) generation of concentration gradients of HOCs under the microscope, (2) exposure and direct observation of microorganisms in such gradients, and (3) quantification of HOC mass transfer. Silicone O-rings were integrated into a Dunn chemotaxis chamber to serve as sink and source for polycyclic aromatic hydrocarbons (PAHs). This resulted in stable concentration gradients in water (>24 h). Adding the model organism Tetrahymena pyriformis to the experimental system enhanced PAH mass transfer up to hundred-fold (benzo[a]pyrene). Increasing mass transfer enhancement with hydrophobicity indicated PAH co-transport with the motile organisms. Fluorescence microscopy confirmed such transport. The effective diffusivity of T. pyriformis, determined by video imaging microscopy, was found to exceed molecular diffusivities of the PAHs up to four-fold. Cell-bound PAH fractions were determined to range from 28% (naphthalene) to 92% (pyrene). Motile microorganisms can therefore function as effective carriers for HOCs under diffusive conditions and might significantly enhance mobility and availability of HOCs.

Swimming behavior and prey retention of the polychaete larvae Polydora ciliata (Johnston)

SUMMARY The behavior of the ubiquitous estuarine planktotrophic spionid polychaete larvae Polydora ciliata was studied. We describe ontogenetic changes in morphology, swimming speed and feeding rates and have developed a simple swimming model using low Reynolds number hydrodynamics. In the model we assumed that the ciliary swimming apparatus is primarily composed of the prototroch and secondarily by the telotroch. The model predicted swimming speeds and feeding rates that corresponded well with the measured speeds and rates. Applying empirical data to the model, we were able to explain the profound decrease in specific feeding rates and the observed increase in the difference between upward and downward swimming speeds with larval size. We estimated a critical larval length above which the buoyancy-corrected weight of the larva exceeds the propulsion force generated by the ciliary swimming apparatus and thus forces the larva to the bottom. This modeled critical larval length corresponded to approximately 1 mm, at which, according to the literature, competence for metamorphosis and no more length increase is observed. These findings may have general implications for all planktivorous polychaete larvae that feed without trailing threads. We observed bell shaped particle retention spectra with a minimum prey size of approximately 4 μm equivalent spherical diameter, and we found that an ontogenetic increase in maximum prey size add to a reduction in intra-specific food competition in the various larval stages. In a grazing experiment using natural seawater, ciliates were cleared approximately 50% more efficiently than similar sized dinoflagellates. The prey sizes retainable for P. ciliata larvae covers the microplankton fraction and includes non-motile as well as motile prey items, which is why the larvae are trophically positioned among the copepods and dinoflagellates. Not only do larval morphology and behavior govern larval feeding, prey behavior also influences the feeding efficiency of Polydora ciliata.

LOW THERMAL LIMIT OF GROWTH RATE OF SYMBIODINIUM CALIFORNIUM (DINOPHYTA) IN CULTURE MAY RESTRICT THE SYMBIONT TO SOUTHERN POPULATIONS OF ITS HOST ANEMONES (ANTHOPLEURA SPP.; ANTHOZOA, CNIDARIA)1

Symbiodinium californium (#383, Banaszak et al. 1993 ) is one of two known dinoflagellate symbionts of the intertidal sea anemones Anthopleura elegantissima, A. xanthogrammica, and A. sola and occurs only in hosts at southern latitudes of the North Pacific. To investigate if temperature restricts the latitudinal distribution of S. californium, growth and photosynthesis at a range of temperatures (5°C–30°C) were determined for cultured symbionts. Mean specific growth rates were the highest between 15°C and 28°C (μ 0.21–0.26 · d−1) and extremely low at 5, 10, and 30°C (0.02–0.03 · d−1). Average doubling times ranged from 2.7 d (20°C) to 33 d (5, 10, and 30°C). Cells cultured at 10°C had the greatest cell volume (821 μm3) and the highest percentage of motile cells (64.5%). Growth and photosynthesis were uncoupled; light‐saturated maximum photosynthesis (Pmax) increased from 2.9 pg C · cell−1 · h−1 at 20°C to 13.2 pg C · cell−1 · h−1 at 30°C, a 4.5‐fold increase. Less than 11% of daily photosynthetically fixed carbon was utilized for growth at 5, 10, and 30°C, indicating the potential for high carbon translocation at these temperatures. Low temperature effects on growth rate, and not on photosynthesis and cell morphology, may restrict the distribution of S. californium to southern populations of its host anemones.

A Redescription of Balanion comatum Wulff, 1919 (Prorodontida, Ciliophora), with Notes on its Cultivation and Behaviour

ABSTRACT Morphological and behavioural information (from live, protargol‐stained, SEM, and TEM materials) were used to re‐describe the type species of Balanion, isolated from Danish waters. The ciliate was compared to the original description and to the only other described species, Balanion planctonicum, Balanion comatum was neotypified and distinguished from B. planctonicum by having 12–26 vs. 7–12 kinetosomes in the somatic kinetics and having 9 vs. 4 microtubules per oral flap.