Candidatus Abditibacter,a novel genus within the Cryomorphaceae,thriving in the North Sea

Coastal phytoplankton blooms are frequently followed by successive blooms of heterotrophic bacterial clades. The class Flavobacteriia within the Bacteroidetes has been shown to play an important role in the degradation of high molecular weight substrates that become available in the later stages of such blooms. One of the flavobacterial clades repeatedly observed over the course of several years during phytoplankton blooms off the coast of Helgoland, North Sea, is Vis6. This genus-level clade belongs to the family Cryomorphaceae and has been resistant to cultivation to date. Based on metagenome assembled genomes, comparative 16S rRNA gene sequence analyses and fluorescence in situ hybridization, we here propose a novel candidate genus Abditibacter, comprising three novel species Candidatus Abditibacter vernus, Candidatus Abditibacter forsetii and Candidatus Abditibacter autumni. While the small genomes of the three novel photoheterotrophic species encode highly similar gene repertoires, including genes for degradation of proteins and algal storage polysaccharides such as laminarin, two of them – Ca. A. vernus and Ca. A. forsetii – seem to have a preference for spring blooms, while Ca. A. autumni almost exclusively occurs in late summer and autumn.     

High-throughput cultivation of heterotrophic bacteria during a spring phytoplankton bloom in the North Sea

On-going studies of phytoplankton-bacterioplankton interactions at the long-term ecological research site Helgoland Roads have indicated that many of the heterotrophic bacterial taxa have not yet been cultivated. A high-throughput approach combining whole cell matrix-assisted laser desorption ionization – time of flight mass spectroscopy with 16S rRNA gene sequencing was applied to the spring bloom of 2016. Aiming at an assessment of cultivability during a spring bloom, cultivation on solid marine media had to be used since dilution to extinction would not have been feasible for a high-throughput approach, as performed in this study. A total of 5023 isolates were obtained from nine weekly samples on eight different solid media between the early-bloom and post-bloom periods. Most of the 4136 strains identified affiliated with Bacteroidetes (13.3%), Gammaproteobacteria (26.9%), Alphaproteobacteria (40.6%) and Actinobacteria (6.7%). Of the 271 operational phylogenetic units (OPUs) identified, 13 are likely to represent novel genera and 143 novel species. A comparison with 16S rRNA gene tag data indicated that most of the isolates were rather rare in surface waters, with the exception of five OPUs affiliating with Rhodobacteraceae, Polaribacter, Psychromonas and Pseudoalteromonas. The effort yielded many novel isolates, yet most of the abundant heterotrophic bacteria still remained elusive. The large strain collection obtained will not only provide insights into the succession of the cultivable fraction of the bacterioplankton, but also enable fine-tuned taxonomic and physiological follow-up studies for improving our knowledge on heterotrophic bacteria in North Sea waters.     

Cultivation of particle-associated heterotrophic bacteria during a spring phytoplankton bloom in the North Sea

Seawater contains free-living and particle-attached bacteria. Only a small fraction is cultivable on plates. As free-living and particle-associated bacteria differ in their physiological traits, their cultivability on plates may coincide with particle association. Using filtration and Imhoff sedimentation cones, particles were collected during a spring phytoplankton bloom off Helgoland (North Sea) in order to obtain particle-associated bacteria as inocula. Direct dilution plating resulted in 526 strains from 3 µm filtration retentates and 597 strains from settled particles. Motile Gammaproteobacteria from the genera Pseudoalteromonas, Shewanella, Psychrobacter, Vibrio and Colwellia, as well as particle-attached Flavobacteriia affiliating with the genera Tenacibaculum and Gramella, were frequently isolated. As a result, a diverse collection comprised of 266 strains was deposited. Two strains were most likely to represent novel genera and 78 strains were probably novel species. Recently, a high-throughput cultivation study from the same site using seawater as an inoculum had retrieved 271 operational phylogenetic units (OPUs) that represented 88% of the 4136 characterized strains at the species level. A comparison of 16S rRNA gene sequences revealed that the collection obtained matched 104 of the 271 seawater OPUs at the species level and an additional 113 at the genus level. This large overlap indicated a significant contribution of particle-associated bacteria to the cultivable microbiome from seawater. The presence of 49 genera not identified in the larger seawater study suggested that sample fractionation was an efficient strategy to cultivate rare members of the planktonic microbiome. The diverse collection of heterotrophic bacteria retrieved in this study will be a rich source for future studies on the biology of particle-associated bacteria.     

Changes in the macrozoobenthos of the intertidal zone at Helgoland (German Bight, North Sea): a survey of 1984 repeated in 2002

Changes in the presence and absence of invertebrates as well as in species conspicuousness were documented in a rocky intertidal community based on surveys in 1984 and 2002. In 2002 six vertically and/or morphologically different stations of an intertidal platform were sampled. Five of these six habitats had already been surveyed in 1984. Replicating precisely the method of the first assessment, presence/absence changes as well as changes in species conspicuousness of 83 invertebrate species were documented, indicating that this intertidal community changed considerably during the 18-year interval. Compared with the study in 1984, 27 species newly appeared, whereas 32 species disappeared. Furthermore, 16 species increased in conspicuousness, whereas eight invertebrates decreased. The total number of species in 2002 was 154 versus 158 in 1984. Although algal species were not recorded as thoroughly as invertebrates, a massive decline in cover of Halidrys siliquosa was noted. Conversely, two invasive algal species became established after 1984, Sargassum muticum (since 1988), a cosmopolitan fucoid alga that prefers shallow subtidal areas for colonization, and Mastocarpus stellatus (introduction in the 1980s) that particularly colonized areas in the mid intertidal. In 1984 the mid intertidal zone was dominated by the brown alga Fucus serratus, whereas in 2002 the blue mussel Mytilus edulis and the periwinkle Littorina littorea were the most conspicuous organisms. Annual mean sea surface temperature (BAH measurements) warmed by 1.1°C over the past four decades. Range-related community shifts, introductions of non-indigenous species and the input of pollutants, are considered to explain long-term ecological changes in the invertebrate community at Helgoland.     

Long-term changes in the macrozoobenthos around the rocky island of Helgoland (German Bight,North Sea)

The paper briefly summarizes what is known about long-term changes (facts, causes, consequences) in the macrozoobenthos of intertidal and subtidal hard-bottom communities around the island of Helgoland (German Bight, North Sea). There is increasing observational evidence that these communities (spectrum and abundances of species) are changing on a long-term temporal scale. The reasons are diverse and mainly anthropogenic. A shift in North Sea climate towards more oceanic conditions may be among the most important factors driving the recent changes in species spectrum. Many of the species which have been recorded as new to the Helgoland area during the past decade are southern (oceanic) species which may be considered as indicators of a warming trend.Communicated by K. Wiltshire     

Physiological diversity of Roseobacter clade bacteria co-occurring during a phytoplankton bloom in the North Sea

Organisms of the Roseobacter clade are an important component in marine ecosystems, partially due to their metabolic variety. Not much is known, however, about the physiological diversity of different roseobacters present within one habitat. By using serial dilution cultures with low-nutrient media seven roseobacter strains, co-occurring during a phytoplankton bloom in the southern North Sea, were obtained in this study. Physiological characterization exhibited distinct substrate spectra of the isolates. Although no isolate showed growth on algal osmolyte dimethylsulfoniopropionate (DMSP), feeding experiments revealed that all new strains converted [²H₆]DMSP into a variety of volatile compounds. Six strains mainly decomposed DMSP via the demethylation pathway, but four strains were also capable of cleaving DMSP to DMS and acrylate. It is hypothesized that the great physiological diversity of the roseobacters reflects their ability to inhabit different ecological niches and enables the organisms to cope differently with changing substrate supplies during phytoplankton blooms. Denaturing gradient gel electrophoresis and sequencing of excised bands resulted in detection of five additional roseobacters. Three of these sequences showed affiliation with three of the four major clusters of the Roseobacter clade, consisting predominantly of uncultured organisms (i.e. the Roseobacter clade-affiliated (RCA)), the NAC11-7 and the CHAB-I-5 clusters.     

Climate change and the timing, magnitude, and composition of the phytoplankton spring bloom

In this article, we show by mesocosm experiments that winter and spring warming will lead to substantial changes in the spring bloom of phytoplankton. The timing of the spring bloom shows only little response to warming as such, while light appears to play a more important role in its initiation. The daily light dose needed for the start of the phytoplankton spring bloom in our experiments agrees well with a recently published critical light intensity found in a field survey of the North Atlantic (around 1.3 mol photons m^?2 day^?1). Experimental temperature elevation had a strong effect on phytoplankton peak biomass (decreasing with temperature), mean cell size (decreasing with temperature) and on the share of microplankton diatoms (decreasing with temperature). All these changes will lead to poorer feeding conditions for copepod zooplankton and, thus, to a less efficient energy transfer from primary to fish production under a warmer climate.     

Further studies of a spring phytoplankton bloom in an enclosed experimental ecosystem

A detailed characterization is presented of the spring diatom bloom which occurred in the enclosed experimental ecosystem bags at Loch Ewe, Scotland, during March–April 1983. The nutrient condition and bacterial biomass of the water column, phytoplankton species distribution, gross biochemical composition and detailed lipid composition (lipid class, fatty acid and free sterol) of the phytoplankton are reported throughout the bloom period. The results are compared with results from previous years. The conclusions are that major changes take place in the biochemical composition of a rapidly growing diatom population which affect both the gross composition and also the more detailed lipid composition. Such changes can take place over a matter of days and appear to be very dependent upon available growth conditions. Both carbohydrate and lipids levels increase towards the end of the bloom as nitrate and silicate levels are depleted in the water. Neutral lipids are shown to be important lipid components of the phytoplankton populations and long-chain ω3 polyunsaturated fatty acids are found to be only minor components of the structural polar lipids. The fatty acid and sterol data are discussed in relation to present knowledge concerning phytoplankton lipid composition.     

Physical processes and their role on the spatial and temporal variability of the spring phytoplankton bloom in the central Yellow Sea

The spatial and temporal variability of the spring phytoplankton bloom (SPB) in the central Yellow Sea is studied, using SeaWiFS surface chlorophyll remote-sensing data, AVHRR sea surface temperatures (SST), QuikSCAT sea surface wind speed (SSW) from 1998 to 2009 and the cruise survey data in 2007 and 2009. The influences of the hydrological conditions on the SPB are significant. (1) The rising SST and low SSW in spring play an important role in the development of the SPB. The SPB in the central Yellow Sea occurs primarily in April (from April 3 to April 24) and at this period the mean SST is generally greater than 10 °C, and 24 h averaged SSW is less than 5.4 m/s. The 99% of the SPB occurs when the SST is 9–14 °C and SSW is 0–7.9 m/s. (2) Specifically, the development of the SPB is from April 4 to April 7 and from April 4 to April 22 in 2007 and 2009 respectively. The longer duration of the SPB in 2009 than that in 2007 is related to the high SST in 2009, which is approximately 2 °C greater than that in 2007, and the weak SSW in 2009, which is much lower than that in 2007, which further indicate that these two factors are critical to the duration of the SPB. (3) The horizontal distribution of surface chlorophyll a in 2007 is found to be greater and located more northward than that in 2009. Comparing the spatial coverage of the SPB, the path of the Yellow Sea warm current, which is warmer and salty, and the location of maximum SST reveals that the spatial coverage of the SPB locates in the warmer temperature (>9 °C) and higher salinity (>33) waters in March and this warm and salty water was much more evident in 2007 than that in 2009 because a stronger warm current of the Yellow Sea in winter. (4) The vertical depths of the maximum chlorophyll layer (MCL) appear at three levels: surface, 10 m and 30 m in 2007, whereas the MCL occurs only at a sub-surface level in 2009 which is related to vertical stability of the water column, i.e., the water is vertically mixed in 2007 while vertically stratified in 2009.     

Physical processes and their role on the spatial and temporal variability of the spring phytoplankton bloom in the central Yellow Sea

The spatial and temporal variability of the spring phytoplankton bloom (SPB) in the central Yellow Sea is studied, using SeaWiFS surface chlorophyll remote-sensing data, AVHRR sea surface temperatures (SST), QuikSCAT sea surface wind speed (SSW) from 1998 to 2009 and the cruise survey data in 2007 and 2009. The influences of the hydrological conditions on the SPB are significant. (1) The rising SST and low SSW in spring play an important role in the development of the SPB. The SPB in the central Yellow Sea occurs primarily in April (from April 3 to April 24) and at this period the mean SST is generally greater than 10 °C, and 24 h averaged SSW is less than 5.4 m/s. The 99% of the SPB occurs when the SST is 9–14 °C and SSW is 0–7.9 m/s. (2) Specifically, the development of the SPB is from April 4 to April 7 and from April 4 to April 22 in 2007 and 2009 respectively. The longer duration of the SPB in 2009 than that in 2007 is related to the high SST in 2009, which is approximately 2 °C greater than that in 2007, and the weak SSW in 2009, which is much lower than that in 2007, which further indicate that these two factors are critical to the duration of the SPB. (3) The horizontal distribution of surface chlorophyll a in 2007 is found to be greater and located more northward than that in 2009. Comparing the spatial coverage of the SPB, the path of the Yellow Sea warm current, which is warmer and salty, and the location of maximum SST reveals that the spatial coverage of the SPB locates in the warmer temperature (>9 °C) and higher salinity (>33) waters in March and this warm and salty water was much more evident in 2007 than that in 2009 because a stronger warm current of the Yellow Sea in winter. (4) The vertical depths of the maximum chlorophyll layer (MCL) appear at three levels: surface, 10 m and 30 m in 2007, whereas the MCL occurs only at a sub-surface level in 2009 which is related to vertical stability of the water column, i.e., the water is vertically mixed in 2007 while vertically stratified in 2009.     

Copepod feeding behaviour and egg production during a dinoflagellate bloom in the North Sea

Feeding strategies of copepods were studied during a dinoflagellate-dominated bloom in the North Sea in August 2001. The aim of this study was to evaluate the importance of mesozooplankton grazing as a biological loss factor of harmful algal blooms under natural conditions. Therefore, ingestion, egestion and egg production experiments were performed with the most abundant copepod species Calanus helgolandicus, Temora longicornis and Acartia sp. feeding on the natural phytoplankton community. Dinophysis norvegica and Ceratium furca were the most abundant dinoflagellate species at the time of the experiments. Grazing experiments as well as examination of fecal pellet content revealed C. helgolandicus fed efficiently on D. norvegica. Ingestion rates up to 47 cells female⁻¹ h⁻¹ were measured and a large proportion of the C. helgolandicus fecal pellets contained intact D. norvegica cells. Dinophysis cells were rarely seen in fecal pellets produced by T. longicornis, and never observed in pellets produced by Acartia sp. The ingestion rate of C. furca, which was the dominating Ceratium species, mimicked that of D. norvegica. C. helgolandicus grazed significantly on C. furca (16 cells female⁻¹ h⁻¹), while the ingestion rate of T. longicornis was low and Acartia sp. was not able to graze on C. furca. Egg production experiments revealed that 92% of the C. helgolandicus females produced eggs. The specific egg production rate and the proportion of females producing eggs among T. longicornis were low. This field experiment clearly shows that some copepod species feed efficiently on D. norvegica and C. furca under natural conditions, which may affect the bloom development of these dinoflagellates.     

Earlier onset of the spring phytoplankton bloom in lakes of the temperate zone in a warmer climate

The decoupling of trophic interactions is potentially one of the most severe consequences of climate warming. In lakes and oceans the timing of phytoplankton blooms affects competition within the plankton community as well as food–web interactions with zooplankton and fish. Using Upper Lake Constance as an example, we present a model‐based analysis that predicts that in a future warmer climate, the onset of the spring phytoplankton bloom will occur earlier in the year than it does at present. This is a result of the earlier occurrence of the transition from strong to weak vertical mixing in spring, and of the associated earlier onset of stratification. According to our simulations a shift in the timing of phytoplankton growth resulting from a consistently warmer climate will exceed that resulting from a single unusually warm year. The numerical simulations are complemented by a statistical analysis of long‐term data from Upper Lake Constance which demonstrates that oligotrophication has a negligible effect on the timing of phytoplankton growth in spring and that an early onset of the spring phytoplankton bloom is associated with high air temperatures and low wind speeds.     

Production of viruses during a spring phytoplankton bloom in the South Pacific Ocean near of New Zealand

Lagrangian studies of virus activity in pelagic environments over extended temporal scales are rare. To address this, viruses and bacteria were examined during the course of a natural phytoplankton bloom in the pelagic South Pacific Ocean east of New Zealand. Daily samples were collected in a mesoscale eddy from year days 263-278 (September 19th-October 4th, 2008). The productive bloom transitioned from a diatom to a pico- and nanoplankton-dominated system, resulting in chlorophyll a concentrations up to 2.43?μg?L(-1) . Virus abundances fluctuated c.?10-fold (1.8?×?10(10) -1.3?×?10(11) L(-1) ) over 16?days. The production rates of virus particles were high compared with those reported in other marine systems, ranging from 1.4?×?10(10) to 2.1?×?10(11) L(-1) day(-1) . Our observations suggest viruses contributed significantly to the mortality of bacteria throughout the bloom, with 19-216% of the bacterial standing stock being lysed daily. This mortality released nutrient elements (N, Fe) that likely helped sustain the bloom through the sampling period. Parametric analyses found significant correlations with both biotic (e.g. potential host abundances) and abiotic parameters (e.g. nutrient concentrations, temperature). These observations demonstrate that viruses may be critical in the extended maintenance of regeneration-driven biological production.     

Complete genome sequence of strain s85, a novel member of the family Flavobacteriaceae

An agar-degrading marine bacterium identified as a novel member of the family Flavobacteriaceae (strain S85) was isolated from seawater in Micronesia. The sequenced strain S85 genome is composed of 3,384,629 bp in a circular chromosome, which includes 2,883 complete open reading frames.     

Preliminary investigation of the contribution of fast-ice algae to the spring phytoplankton bloom in Ellis Fjord,eastern Antarctica

 Algae released from fast-ice in Ellis Fjord, eastern Antarctica, made little contribution to subsequent phytoplankton growth. Dominant taxa in the interior ice community included Nitzschia cylindrus (Grun) Hasle, Navicula glaciei V.H. and a dinoflagellate cyst. Diatom mortality within the ice was high. The algal contribution to the phytoplankton from the fast ice was estimated by calculating the difference between algal biomass in ice cores taken on 14 November with those taken on 18 December 1992. The biomass of sedimenting phytoplankton was estimated using sediment traps; weekly cell counts of water were used to monitor net phytoplankton growth. The low contribution from the fast-ice of Ellis Fjord to the phytoplankton is similar to results from other Antarctic fast-ice communities but is not necessarily reflective of processes occurring within either Antarctic or Arctic pack ice communities. An algal mat growing on the base of the fast-ice had a carbon standing crop of between 0.231 gC m^-2 and 0.022 gC m^-2. Much of this was delivered to the water column as the ice melted while the remainder was exported. Received: 15 March 1995/Accepted: 4 September 1995     

Dynamics in bacterial surface properties of a natural bacterial community in the coastal North Sea during a spring phytoplankton bloom

The hydrophilic and hydrophobic properties of single cells of natural bacterioplankton communities were determined using a recently developed staining method combined with confocal laser scanning microscopy and advanced image analysis. On an average, about 50% of the bacterial cell area was covered by hydrophobic and only 16% by hydrophilic properties, while about 72% was covered by the genome. However, the size of these properties was independent of the bacterial cell size. Bacterial hydrophobicity was positively correlated with ambient NH(4)(+) concentrations and negatively correlated with overall bacterial abundance. The expression of hydrophilicity was more dynamic. Over the spring phytoplankton bloom, the bacterioplankton ratio(phil/phob) repeatedly reached highest values shortly before peaks in bacterioplankton abundance were observed, indicating a direct and fast response of bacterial surface properties, especially hydrophilicity, to changing environmental conditions. Compared to bacterial strains, recently studied with the same method, cells of marine bacterioplankton communities are much smaller and less frequently covered by hydrophobic or hydrophilic properties. While the percentage area covered by the genome is essentially the same, the percentage area covered by hydrophobic or hydrophilic properties is much smaller.     

Growth and decline of a diatom spring bloom phytoplankton species composition, formation of marine snow and the role of heterotrophic dinoflagellates

During the spring of 1994, we determined the factors responsiblefor the decline of the seasonal diatom bloom in the Gullmarfjord, on the west coast of Sweden. Four species constituted>75% of the biomass—Detonula confervacea, Chaetocerosdiadema, Skeletonema costatum and Thalassiosira nordenskioeldii—reachingconcentrations of 4900, 350, 8200 and 270 cells ml^–1,respectively. Growth of phytoplankton was exponential (growthrate = 0.12 day^–1) from 3 to 21 March, after which a galewith winds >15 m s^–1 caused massive aggregation. Amaximum of 130 p.p.m. (v/v) of marine snow aggregates was observedby in situ video at the peak of the bloom. Critical concentrations(Jackson, Deep-Sea Res., 37, 1197–1211, 1990) were similarto observed showing that coagulation theory could explain thesudden decline of the bloom. The heterotrophic dinoflagellateGyrodinium cf. spirale increased exponentially after the peakof the bloom with maximum (temperature-adjusted) growth rates.After the rapid aggregation and sedimentation of the bloom,they were able to control any further growth of diatoms. Nitrateand silicate were never depleted, but phosphate may have beenlimiting by the end of the study period. We conclude that massaggregation during a gale marked the end of the bloom, and thatintense grazing by heterotrophic dinoflagellates prevented anysubsequent increase of diatoms.     

Changing expression patterns of TonB-dependent transporters suggest shifts in polysaccharide consumption over the course of a spring phytoplankton bloom

Algal blooms produce large quantities of organic matter that is subsequently remineralised by bacterial heterotrophs. Polysaccharide is a primary component of algal biomass. It has been hypothesised that individual bacterial heterotrophic niches during algal blooms are in part determined by the available polysaccharide substrates present. Measurement of the expression of TonB-dependent transporters, often specific for polysaccharide uptake, might serve as a proxy for assessing bacterial polysaccharide consumption over time. To investigate this, we present here high-resolution metaproteomic and metagenomic datasets from bacterioplankton of the 2016 spring phytoplankton bloom at Helgoland island in the southern North Sea, and expression profiles of TonB-dependent transporters during the bloom, which demonstrate the importance of both the Gammaproteobacteria and the Bacteroidetes as degraders of algal polysaccharide. TonB-dependent transporters were the most highly expressed protein class, split approximately evenly between the Gammaproteobacteria and Bacteroidetes, and totalling on average 16.7% of all detected proteins during the bloom. About 93% of these were predicted to take up organic matter, and for about 12% of the TonB-dependent transporters, we predicted a specific target polysaccharide class. Most significantly, we observed a change in substrate specificities of the expressed transporters over time, which was not reflected in the corresponding metagenomic data. From this, we conclude that algal cell wall-related compounds containing fucose, mannose, and xylose were mostly utilised in later bloom stages, whereas glucose-based algal and bacterial storage molecules including laminarin, glycogen, and starch were used throughout. Quantification of transporters could therefore be key for understanding marine carbon cycling.Subject terms: Water microbiology, Molecular ecology, Proteomics     

Abundance,population structure and vertical distribution of dominant calanoid copepods on the eastern Weddell Sea shelf during a spring phytoplankton bloom

The main emphasis of this study was to analyse the short-term development of abundance, population structure and vertical distribution of the dominant calanoid copepods during a phytoplankton bloom in the coastal area of the eastern Weddell Sea in December 2003. Microcalanus pygmaeus was by far the most abundant calanoid species. Metridia gerlachei, Ctenocalanus citer, Calanoides acutus, Calanus propinquus and the ice-associated Stephos longipes were also present in considerable proportions. The observed changes in the population characteristics and parameters of these species are described in detail and discussed in the context of the spring phytoplankton bloom. A conspicuous event occurring during the final stage of the study was the development of a strong storm. While the results suggest that this storm did not have any considerable influence on the populations of all other investigated copepod species, it very likely caused pronounced changes in the S. longipes population present in the water column. Before the storm, S. longipes was found primarily in the upper 100 m of the water column, and its population was dominated by adults (mean proportion = 41%) and the copepodite stage I (mean proportion = 30%). After the storm, the abundance increased considerably, and the copepodite stage I contributed by far the largest proportion (53%) of the total population indicating that the early copepodite stages probably had been released from the sea ice into the under ice water layer due to ice break-up and ice melt processes caused by the storm.