Mid-winter limnological surveys of Lake Erie captured extremes in ice extent ranging from expansive ice cover in 2010 and 2011 to nearly ice-free waters in 2012. Consistent with a warming climate, ice cover on the Great Lakes is in decline, thus the ice-free condition encountered may foreshadow the lakes future winter state. Here, we show that pronounced changes in annual ice cover are accompanied by equally important shifts in phytoplankton and bacterial community structure. Expansive ice cover supported phytoplankton blooms of filamentous diatoms. By comparison, ice free conditions promoted the growth of smaller sized cells that attained lower total biomass. We propose that isothermal mixing and elevated turbidity in the absence of ice cover resulted in light limitation of the phytoplankton during winter. Additional insights into microbial community dynamics were gleaned from short 16S rRNA tag (Itag) Illumina sequencing. UniFrac analysis of Itag sequences showed clear separation of microbial communities related to presence or absence of ice cover. Whereas the ecological implications of the changing bacterial community are unclear at this time, it is likely that the observed shift from a phytoplankton community dominated by filamentous diatoms to smaller cells will have far reaching ecosystem effects including food web disruptions.
Recent discoveries have altered prevailing paradigms concerning the conditions under which nitrification takes place and the organisms responsible for nitrification in aquatic ecosystems. In Lake Superior, nitrate (NO is generated by nitrification within the lake, important questions remain concerning the magnitude and controls of nitrification, and which microbial groups are primarily responsible for this process. We measured water-column nitrification rates in the western basin of Lake Superior during five research cruises from November 2009 to March 2011. Using in situ bottle incubations at 10 depths, we quantified nitrification rates using both the oxidation of 15 N-labeled ammonium (NH z 4 ) and the uptake of 14 C associated with nitrification. Average rates of NH z 4 oxidation ranged from 18-34 nmol N L 21 d 21 across the five cruises, similar to values reported for the coastal ocean, and two orders of magnitude lower than values reported from other lakes. Low nitrification rates observed in the epilimnion corresponded to the absence of ammonium-oxidizing archaea and nitrite-oxidizing bacteria. The measured rates of nitrification are . 50-fold greater than the long-term NO { 3 rise in the lake, indicating that N is actively cycling and that long-term change in this ecosystem is mediated by internal dynamics.
In September 2004 a large, nearly monospecific diatom bloom of Pseudo-nitzschia cuspidata off the coast of the state of Washington reached cell concentrations of 6.1 3 10 6 cells L 21 and produced maximum particulate domoic acid (pDA), dissolved domoic acid (dDA), and cellular domoic acid concentrations of 43 nmol L 21 , 4 nmol L 21 , and 63 pg cell 21 , respectively. This bloom co-dominated the phytoplankton assemblage with the euglenoid Eutreptiella sp. in the Juan de Fuca eddy region, a known initiation site for toxigenic Pseudo-nitzschia blooms. Two isolates of P. cuspidata collected during separate cruises produced domoic acid (DA) in culture. During the September 2004 survey, 84% of the stations (n 5 98) had detectable Pseudo-nitzschia and 78% had detectable pDA. There were no significant correlations between either pDA or cellular DA and ambient concentrations of macronutrients; however, when considering only those stations where Pseudo-nitzschia was present, pDA was positively correlated with chlorophyll a and negatively correlated with temperature (p # 0.01) at both 1-and 5-m depths. Correlations between cellular DA concentrations and total bacteria or cyanobacteria abundances were not significant. Variable ratios of pDA : dDA in the eddy region suggest that DA release was under cellular regulation by Pseudo-nitzschia. Stations where dissolved Fe concentrations were limiting (,0.5 nmol L 21 ) had the highest Pseudo-nitzschia abundances and pDA and cellular DA values. These results provide enticing field evidence of the role of Fe limitation in controlling cellular DA levels.
We present evidence for the directed formation of ice by planktonic communities dominated by filamentous diatoms sampled from the ice-covered Laurentian Great Lakes. We hypothesize that ice formation promotes attachment of these non-motile phytoplankton to overlying ice, thereby maintaining a favorable position for the diatoms in the photic zone. However, it is unclear whether the diatoms themselves are responsible for ice nucleation. Scanning electron microscopy revealed associations of bacterial epiphytes with the dominant diatoms of the phytoplankton assemblage, and bacteria isolated from the phytoplankton showed elevated temperatures of crystallization (T c ) as high as À 3 1C. Ice nucleation-active bacteria were identified as belonging to the genus Pseudomonas, but we could not demonstrate that they were sufficiently abundant to incite the observed freezing. Regardless of the source of ice nucleation activity, the resulting production of frazil ice may provide a means for the diatoms to be recruited to the overlying lake ice, thereby increasing their fitness. Bacterial epiphytes are likewise expected to benefit from their association with the diatoms as recipients of organic carbon excreted by their hosts. This novel mechanism illuminates a previously undescribed stage of the life cycle of the meroplanktonic diatoms that bloom in Lake Erie and other Great Lakes during winter and offers a model relevant to aquatic ecosystems having seasonal ice cover around the world.
Ice-nucleating particles (INPs) associated with fresh waters are a neglected, but integral component of the water cycle. Abundant INPs were identified from surface waters of both the Maumee River and Lake Erie with ice nucleus spectra spanning a temperature range from −3 to −15 °C. The majority of river INPs were submicron in size and attributed to biogenic macromolecules, inferred from the denaturation of ice-nucleation activity by heat. In a watershed dominated by row-crop agriculture, higher concentrations of INPs were found in river samples compared to lake samples. Further, ice-nucleating temperatures differed between river and lake samples, which indicated different populations of INPs. Seasonal analysis of INPs that were active at warmer temperatures (≥−10 °C; INP −10 ) showed their concentration to correlate with river discharge, suggesting a watershed origin of these INPs. A terrestrial origin for INPs in the Maumee River was further supported by a correspondence between the ice-nucleation signatures of river INPs and INPs derived from the soil fungus Mortierella alpina. Aerosols derived from turbulence features in the river carry INP −10 , although their potential influence on regional weather is unclear. INP −10 contained within aerosols generated from a weir spanning the river, ranged in concentration from 1 to 11 INP m −3 , which represented a fold-change of 3.2 over average INP −10 concentrations sampled from aerosols at control locations.
Chlamydomonas raudensis H. Ettl (UWO 241) is a psychrophilic green alga endemic to Lake Bonney, Antarctica. The objective of this study was to investigate the response of UWO 241 to incubation at 24°C, a temperature close to optimum for related mesophilic species. Using chl a fluorescence analysis, shifting cells from a growth temperature of 10°C-24°C resulted in a decline in PSII photochemical efficiency with light energy being directed away from photochemistry and toward dissipative pathways. Using the SYTOX Green assay, it was determined that UWO 241 cells die when incubated at 24°C under growth irradiance with a half-time of 34.9 h. The role of light in cell death was minor as cell death occurred in darkness at 24°C with a half-time of 43.7 h. To examine the plasticity of UWO 241 to temperature stress, 10°C-grown cells were shifted to 24°C for 12 h and then returned to 10°C to recover. The 12 h incubation at 24°C, which resulted in <10% cell death, led to declines in both light-saturated rates of photosynthesis and respiration, PSII photochemistry and energy partitioning, and changes to transcript abundances-those associated with the light-harvesting protein of PSII and ferredoxin declining rapidly, whereas transcripts of specific heat-shock proteins (HSPs) increased. Within 24-48 h of being transferred back to 10°C, all parameters returned to levels occurring in 10°C-grown cells. This research shows, for the first time, that 24°C is a temperature that is lethal to UWO 241, and yet this organism displays considerable physiological and molecular plasticity.
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