Highly divergent SSU rRNA genes found in the marine ciliates Myrionecta rubra and Mesodinium pulex

Myrionecta rubra and Mesodinium pulex are among the most commonly encountered planktonic ciliates in coastal marine and estuarine regions throughout the world. Despite their widespread distribution, both ciliates have received little attention by taxonomists. In order to better understand the phylogenetic position of these ciliates, we determined the SSU rRNA gene sequence from cultures of M. rubra and M. pulex. Partial sequence data were also generated from isolated cells of M. rubra from Chesapeake Bay. The M. rubra and M. pulex sequences were very divergent from all other ciliates, but shared a branch with 100% bootstrap support. Both species had numerous deletions and substitutions in their SSU rRNA gene, resulting in a long branch for the clade. This made the sequences prone to spurious phylogenetic affiliations when using simple phylogenetic methods. Maximum likelihood analysis placed M. rubra and M. pulex on the basal ciliate branch, following removal of ambiguously aligned regions. Fluorescent in situ hybridization probes were used with confocal laser scanning microscopy to confirm that these divergent sequences were both expressed in the cytoplasm and nucleolus of M. ruisra and M. pulex. We found that our sequence data matched several recently discovered unidentified eukaryotes in Genbank from diverse marine habitats, all of which had apparently been misattributed to highly divergent amoeboid organisms.     

Natural co-occurrence of Dinophysis acuminata (Dinoflagellata) and Mesodinium rubrum (Ciliophora) in thin layers in a coastal inlet

ABSTRACT. The interdependency of Dinophysis spp., Mesodinium rubrum and Teleaulax spp. has occupied scientists in molecular and ecological domains in recent years. Current knowledge about the predator–prey relationships is based on laboratory investigations. Records on interactions in nature are limited, even though it is known that Dinophysis acuminata and M. rubrum form population maxima in thin layers associated with thermal stratification. We studied the vertical co‐occurrence of these taxa in a stratified coastal inlet in Åland, in the Northern Baltic Sea, SW Finland. Vertical profiles were sampled monthly in the summer of 2008 and observations on diurnal migrational patterns of all species were conducted in September 2008. The population maximum of D. acuminata was almost totally confined to thin layers where the depth maximum of M. rubrum was present. However, this pattern was only observed early in the morning or at noon. The population maxima of M. rubrum and Teleaulax spp. overlapped at noon. Dinophysis acuminata and Teleaulax spp. were restricted to the upper 9 m but M. rubrum was found down to 20 m depth. This study offers circumstantial evidence for the interdependency between the three taxa in nature.     

Temporal stability of marine phytoplankton in a subtropical coastal environment

We investigated the temporal stability of phytoplankton at a subtropical coastal site for 9 months by conducting chlorophyll and flow cytometric measurements at relatively high frequency (roughly at 2–5 day interval). Phytoplankton cells were grouped based on their sizes obtained from flow cytometric signals. We also conducted dilution experiments to estimate the growth and grazing mortality rates of different phytoplankton groups to assess whether the temporal stability of phytoplankton abundances was related with phytoplankton growth/grazing rates. Based on size-fractionated chlorophyll measurements, there was some indication that smaller phytoplankton cells were more stable than larger ones. However, by cytometric counting, there was no evidence for greater stability in small cells. Synechococcus, which had the lowest stability and dominated the <1 μm size class, showed a strong seasonal cycle that was highly dependent on temperature whereas eukaryotes did not have evident seasonal cycles. In general, biomass of a phytoplankton group consisting of several size classes was more stable than that of its sub component, consistent with the hypothesis that higher diversity leads to higher stability, probably related with the effect of statistical averaging (portfolio effect). Stability of heterotrophic bacteria was much higher than that of phytoplankton, leading to the speculation that bacteria were more diverse than phytoplankton. Phytoplankton stability was not related with their growth or grazing mortality rates. Our study suggests that species diversity should be taken into account when considering the temporal stability of phytoplankton.     

Notes on a Mesodinium rubrum red tide in San Francisco Bay (California, USA)

Discrete red patches of water were observed in South San FranciscoBay (USA) on 30 April 1993, and examination of live samplesshowed that this red tide was caused by surface accumulationsof the pigmented ciliate Mesodinium rubrum. Vertical profilesshowed strong salinity and temperature stratification in theupper 5 m, peak chlorophyll fluorescence in the upper meter,and differences in the small-scale density structure and fluorescencedistribution among red patches. Events preceding this Mesodiniumred tide included: (i) heavy precipitation and run-off, allowingfor strong salinity stratification; (ii) a spring diatom bloomwhere the chlorophyll a concentration reached 50 mg m^–3;(ii) depletions of dissolved inorganic N and Si in the photiczone; and (iv) several days of rapid warming and stabilizationof the upper surface layer. These conditions may be generalprerequisites for M.rubrum blooms in temperate estuaries. ¹Present address: Station Marine d'Endoume, Centre d'Oceanologiede Marseille, rue Batterie des Lions, 13007 Marseille, France     

Monitoring phytoplankton, bacterioplankton, and virioplankton in a coastal inlet (Bedford Basin) by flow cytometry.

BACKGROUND: To establish the prevailing state of the ecosystem for the assessment of long-term change, the abundance of microbial plankton in Bedford Basin (Nova Scotia, Canada) is monitored weekly by flow cytometry. METHODS: Phytoplankton are detected by their chlorophyll autofluorescence. Those that contain phycoerythrin are designated as Synechococcus cyanobacteria or cryptophyte algae according to the intensity of light scatter. Bacteria and viruses are stained with DNA-binding fluorochromes and detected by green fluorescence. Distinction is made between bacterial and viral subpopulations exhibiting high and low fluorescence. RESULTS: Time series data are presented for weekly observations from 1991 to 2000. Weekly averages are computed for the complete annual cycle of temperature, salinity, river discharge, nitrate, phosphate, silicate, chlorophyll, total phytoplankton including Synechococcus and cryptophytes, total bacteria including high and low-fluorescence subpopulations, and total viruses including high and low-fluorescence subpopulations. CONCLUSIONS: The microbial biomass in the surface water of Bedford Basin is dominated by phytoplankton. The spring bloom of phytoplankton represents a maximum in algal biovolume, but not in cell number. Phytoplankton, bacteria, and viruses all attain their annual numerical maxima between the summer solstice and the autumn equinox. A vigorous microbial loop and viral shunt is envisioned to occur in the summer.     

Effect of heavy metals on marine phytoplankton

Phytoplankton community responses were used in a heavy metals (HM) marine pollution monitoring biological system. A multifactorial analysis was used in in situ experiments in the White Sea to study HM toxicity to a planktonik algae population in their copresence. It was ascertained that algae with a high rate of increase in population size, producing metabolites, restricting development of phytoplankton community associates, gain advantages in isolation conditions. The algae responded differently to addition of individual metals and their combinations in the environment. Change in the number of dominants resulted in disturbance of the phytoplankton community structure. It has been shown that there are cardinal points in phytoplankton development, determining the growth of individual algae populations and planktonic community on the whole, against a background of competition for biogenic elements and space in conditions of TM environment pollution.     

Seasonal variation in marine phytoplankton and ice algae at a shallow antarctic coastal site

The phytoplankton population near Davis, Vestfold Hills, Antarctica was monitored throughout 1982. Chlorophyll-a determinations and counts of living cells in both the water column and sea ice demonstrated a marked seasonality in phytoplankton abundane and species composition. From April to October nanoplanktonic organisms contributed most of the chlorophyll-a in both the sea ice and water column. Blooms of diatoms occurred in May, November and December in the bottom of the sea-ice and in January and February in the water column. Phaeocystis pouchetii was dominant during December in the water column. Large numbers of dead diatoms were found in winter. The concentrations of nitrate, dissolved inorganic phosphate and dissolved silicate increased throughout the year until December, when the concentrations of nitrate and silicate fell sharply, followed a month later by a reduction in phosphate concentration. The diversity of phytoplankton was greatest during the summer months.     

Effects of phytoplankton species composition on absorption spectra and modeled hyperspectral reflectance

Understanding the spectral characteristics of remotely-sensed reflectance by different phytoplankton species can assist in the development of algorithms to identify various algal groups using satellite ocean color remote sensing. One of the main challenges is to separate the effect of species composition on the reflectance spectrum from other factors such as pigment concentration and particle size structure. Measuring the absorption spectra of nine different cultured algae, and estimating the reflectance of the different species, provides a useful approach to study the effects of species composition on the bio-optical properties. The results show that the absorption spectra of different species exhibit different spectral characteristics and that species composition can significantly change the absorption characteristics at four main peaks (438, 536, 600 and 650 nm). A ‘distance angle index’ was used to compare different phytoplankton species. Results indicate that this index can be used to identify species from the absorption spectra, using a database of standard absorption spectra of known species as reference. By taking into account the role of species composition in the phytoplankton absorption model, the performance of the model can be improved by up to 5%. A reflectance-species model is developed to estimate the remotely-sensed reflectance from the absorption spectra, and the reflectance of different phytoplankton species at the same chlorophyll-a concentration is compared, to understand effects of species composition on the reflectance spectra. Different phytoplankton species can cause up to 33% difference in the modeled reflectance at short wavelengths under the condition of the same chlorophyll-a concentration, and variations in the reflectance spectrum correspond to the colors of the algae. The standard deviation of the reflectance among different species shows that the variations from 400 to 450 nm are sensitive to species composition at low chlorophyll-a concentrations, whereas variations in the 510 to 550 nm range are more sensitive under high chlorophyll-a concentrations. For this reason, the green bands may be more suitable for estimating species composition from hyperspectral satellite data during bloom conditions, whereas the blue bands may be more helpful in detection of species under low chlorophyll-a concentrations. In this theoretical approach, variations in reflectance at the same chlorophyll-a concentration can be used to identify phytoplankton species. Another approach to identify phytoplankton species from remotely-sensed hyperspectral reflectance measurements would be to derive the absorption spectra of phytoplankton from the reflectance measurements, and compare these with a standard database of absorption spectra.     

Photoadaptation in marine phytoplankton : changes in spectral absorption and excitation of chlorophyll a fluorescence

The optical properties of marine phytoplankton were examined by measuring the absorption spectra and fluorescence excitation spectra of chlorophyll a for natural marine particles collected on glass fiber filters. Samples were collected at different depths from stations in temperate waters of the Southern California Bight and in polar waters of the Scotia and Ross Seas. At all stations, phytoplankton fluorescence excitation and absorption spectra changed systematically with depth and vertical stability of the water columns. In samples from deeper waters, both absorption and chlorophyll a fluorescence excitation spectra showed enhancement in the blue-to-green portion of the spectrum (470-560 nm) relative to that at 440 nm. Since similar changes in absorption and excitation were induced by incubating sea water samples at different light intensities, the changes in optical properties can be attributed to photoadaptation of the phytoplankton. The data indicate that in the natural populations studied, shade adaptation caused increases in the concentration of photosynthetic accessory pigments relative to chlorophyll a. These changes in cellular pigment composition were detectable within less than 1 day. Comparisons of absorption spectra with fluorescence excitation spectra indicate an apparent increase in the efficiency of sensitization of chlorophyll a fluorescence in the blue and green spectral regions for low light populations.     

Pigments and species composition of natural phytoplankton populations: effect on the absorption spectra

Data from three cruises (Arabesque 1 and 2 cruises in the ArabianSea and the Vancouver Island cruise) were examined to assessthe importance of species composition and accessory pigmentsin modifying specific absorption coefficients. The three cruisesdiffered widely in their phytoplankton assemblages with smallcells dominating the Arabesque 2 cruise and large diatoms theVancouver Island cruise. Absorption spectra from each cruisewere decomposed into 13 Gaussian bands representing absorptionby the major chlorophylls and accessory pigments. The maximumspecific peak height     

Variability in chlorophyll a specific absorption coefficient in marine phytoplankton as a function of cell size and irradiance

Size-dependence of chlorophyll a (Chl a) specific absorptioncoefficient a*(     

Factors Controlling the Abundance and Size Distribution of the Phototrophic Ciliate Myrionecta rubra in Open Waters of the North Atlantic

ABSTRACT. Myrionecta rubra, a ubiquitous planktonic ciliate, has received much attention due to its wide distribution, occurrence as a red tide organism, and unusual cryptophyte endosymbiont. Although well studied in coastal waters, M. rubra is poorly examined in the open ocean. In the Irminger Basin, North Atlantic, the abundance of M. rubra was 0–5 cells/ml, which is low compared with that found in coastal areas. Distinct patchiness (100 km) was revealed by geostatistical analysis. Multiple regression indicated there was little relationship between M. rubra abundance and a number of environmental factors, with the exception of temperature and phytoplankton biomass, which influenced abundance in the spring. We also improve on studies that indicate distinct size classes of M. rubra; we statistically recognise four significantly distinct width classes (5–16, 12–23, 18–27, 21–33 μm), which decrease in abundance with increasing size. A multinomial logistic regression revealed the main variable correlated with this size distribution was ambient nitrate concentration. Finally, we propose a hypothesis for the distribution of sizes, involving nutrients, feeding, and dividing of the endosymbiont.     

Effect of (a)synchronous light fluctuation on diversity,functional and structural stability of a marine phytoplankton metacommunity

Disentangling the mechanisms that maintain the stability of communities and ecosystem properties has become a major research focus in ecology in the face of anthropogenic environmental change. Dispersal plays a pivotal role in maintaining diversity in spatially subdivided communities, but only a few experiments have simultaneously investigated how dispersal and environmental fluctuation affect community dynamics and ecosystem stability. We performed an experimental study using marine phytoplankton species as model organisms to test these mechanisms in a metacommunity context. We established three levels of dispersal and exposed the phytoplankton to fluctuating light levels, where fluctuations were either spatially asynchronous or synchronous across patches of the metacommunity. Dispersal had no effect on diversity and ecosystem function (biomass), while light fluctuations affected both evenness and community biomass. The temporal variability of community biomass was reduced by fluctuating light and temporal beta diversity was influenced interactively by dispersal and fluctuation, whereas spatial variability in community biomass and beta diversity were barely affected by treatments. Along the establishing gradient of species richness and dominance, community biomass increased but temporal variability of biomass decreased, thus highest stability was associated with species-rich but highly uneven communities and less influenced by compensatory dynamics. In conclusion, both specific traits (dominance) and diversity (richness) affected the stability of metacommunities under fluctuating conditions.     

An automatic device for in vivo absorption spectra acquisition and chlorophyll estimation in phytoplankton cultures

In order to aid the study of photoacclimation, a new programmable deviceis described which provides automatic on-line acquisition of in vivo cellabsorption in phytoplankton cultures. The system was used for a long-termstudy of Rhodomonas salina grown at constant photon flux density ina nitrate-limited continuous culture with different dilution rates. Particulate absorption measured at the red chlorophyll a (Chl a)maximum was not a good proxy of biomass, because of the large variabilityof cellular chlorophyll induced by nitrogen limitation. However, thedevice is well suited to automatic assessment of Chl a andphycoerythrin (PE) concentrations in phytoplankton cultures, if algal cellsize and concentration are measured in parallel to correct the packagingeffect. The effects of nitrogen limitation on Chl a and PE contentsand particle absorbance are discussed.     

Influence of UV-B radiation on photosynthetic light-response curves, absorption spectra and motility of four phytoplankton species

Several unicellular algae were exposed to artificial UV-B (280–320 nm) radiation after adaptation to high (43 W m⁻²) and low (19 W m⁻²) visible light. UV-B radiation had different effects on rates of photosynthesis, motility and absorption spectra for these species. Photosynthesis of Euglena gracilis and the diatom Phaeodactylum tricomution was more sensitive to UV-B inhibition than that of the dinoflagellates Heterocapsa triquetra and Prorocentrum minimum . Not only UV-B radiation but also high visible light had a photoinhibitory effect on photosynthesis in all four organisms. The effect on photosynthesis was observed both on the quantum yield and on the light saturation rate of photosynthesis. The dinoflagellates, in contrast to E. gracilis and P. tricorntum , absorbed strongly in the UV region (334 nm) and their absorption peaks increased after growth under high visible light or with or without UV-B radiation for one week. The swimming speed of H. triquetra decreased more after low visible light and UV-B radiation compared to high visible light and UV-B radiation. The negative effects of UV-B radiation on P. minimum and E. gracilis were most pronounced after high visible light.     

Induced defences in marine and freshwater phytoplankton: a review

Many organisms have developed defences to avoid predation by species at higher trophic levels. The capability of primary producers to defend themselves against herbivores affects their own survival, can modulate the strength of trophic cascades and changes rates of competitive exclusion in aquatic communities. Algal species are highly flexible in their morphology, growth form, biochemical composition and production of toxic and deterrent compounds. Several of these variable traits in phytoplankton have been interpreted as defence mechanisms against grazing. Zooplankton feed with differing success on various phytoplankton species, depending primarily on size, shape, cell wall structure and the production of toxins and deterrents. Chemical cues associated with (i) mechanical damage, (ii) herbivore presence and (iii) grazing are the main factors triggering induced defences in both marine and freshwater phytoplankton, but most studies have failed to disentangle the exact mechanism(s) governing defence induction in any particular species. Induced defences in phytoplankton include changes in morphology (e.g. the formation of spines, colonies and thicker cell walls), biochemistry (such as production of toxins, repellents) and in life history characteristics (formation of cysts, reduced recruitment rate). Our categorization of inducible defences in terms of the responsible induction mechanism provides guidance for future work, as hardly any of the available studies on marine or freshwater plankton have performed all the treatments that are required to pinpoint the actual cue(s) for induction. We discuss the ecology of inducible defences in marine and freshwater phytoplankton with a special focus on the mechanisms of induction, the types of defences, their costs and benefits, and their consequences at the community level.     

Effect of short-term exposure to UVA and UVB on potential phytoplankton production in UK coastal waters

The influence of vertical mixing on phytoplankton sensitivityto UV light has been assessed over an annual cycle. Photosynthesisrates of natural assemblages were compared in samples that wereincubated at fixed position in a light gradient and with duplicatesamples that simulated vertical mixing by movement in the samegradient with a periodicity of 4 h. This is the typical time-scaleof vertical mixing in coastal waters in the English Channel.There were clear seasonal differences in the short-term responseof phytoplankton to enhanced UVA+UVB. For most of the year,there was no detectable effect of UV on photosynthetic carbonfixation. But natural assemblages in late winter/early spring,when high UV light may sporadically occur at this latitude,were sensitive to UVA+UVB. In some samples, primary productionwas 40% of that measured in the absence of UV light. At thetime of maximum sensitivity to UV, the phytoplankton assemblagewas dominated by diatoms. Simulated vertical mixing resultedin more inhibition of photosynthesis by UVA+UVB light than whensamples were at constant light with the same time-integratedirradiance. Transient increases in UVA+UVB due to ozone depletion,such as have been observed over Northern Europe, could havea serious impact on coastal phytoplankton production in latewinter/early spring.     

Seasonal changes in bacterial and phytoplankton biomass in a subantarctic coastal area (Kerguelen Islands)

The seasonal variations of bacterial and phytoplanktonic biomass were studied during several pluri-annual surveys in the subantarctic Morbihan Bay (Kerguelen Islands, 49 ° 20 S; 70 ° 10 E). Large interannual variation was observed. Phytoplanktonic biomass showed moderate values during winter and autumn. They increased sharply in spring, reaching a maximum value of about 1 mg C l^–1 corresponding to an important depletion of nutrients. A second phytoplanktonic bloom of similar amplitude occurred in late summer. During algal blooms which were roughly associated with optimal values of solar irradiation for the first one and with the highest temperatures for the second one, phytoplanktonic material represented near 100% of particulate and living carbon. Bacteria showed maximal abundance (0.2 to 0.7 mg C l^–1) during summer or autumn. Their relative abundance, which represented less than 1% of the living biomass in spring and summer, can reach more than 95% in autumn and winter.