A theoretical modeling framework for motile and colonial harmful algae.

Climate change is leading to an increase in severity, frequency, and distribution of harmful algal blooms across the globe. For many harmful algae species in eutrophic lakes, the formation of such blooms is controlled by three factors: the lake hydrodynamics, the vertical motility of the algae organisms, and the ability of the organisms to form colonies. Here, using the common cyanobacterium Microcystis aeruginosa as an example, we develop a model that accounts for both vertical transport and colony dynamics. At the core of this treatment is a model for aggregation. For this, we used Smoluchowski dynamics containing parameters related to Brownian motion, turbulent shear, differential setting, and cell-to-cell adhesion. To arrive at a complete description of bloom formation, we place the Smoluchowski treatment as a reaction term in a set of one-dimensional advection-diffusion equations, which account for the vertical motion of the algal cells through molecular and turbulent diffusion and self-regulating buoyant motion. Results indicate that Smoluchowski aggregation qualitatively describes the colony dynamics of M. aeruginosa. Further, the model demonstrates wind-induced mixing is the dominant aggregation process, and the rate of aggregation is inversely proportional to algal concentration. Because blooms of Microcystis typically consist of large colonies, both of these findings have direct consequences to harmful algal bloom formation. While the theoretical framework outlined in this manuscript was derived for M. aeruginosa, both motility and colony formation are common among bloom-forming algae. As such, this coupling of vertical transport and colony dynamics is a useful step for improving forecasts of surface harmful algal blooms.

Microalgal swimming signatures and neutral lipids production across growth phases.

Microalgae have been shown as a potential bioresource for food, biofuel, and pharmaceutical products. During the growth phases with corresponding environmental conditions, microalgae accumulate different amounts of various metabolites. We quantified the neutral lipids accumulation and analyzed the swimming signatures (speed and trajectories) of the motile green alga, Dunaliella primolecta, during the lag-exponential-stationary growth cycle at different nutrient concentrations. We discovered significant changes in the neutral lipid content and swimming signatures of microalgae across growth phases. The timing of the maximum swimming speed coincided with the maximum neutral lipid content and both maxima occurred under nutrient stress at the stationary growth phase. Furthermore, the swimming trajectories suggested statistically significant changes in swimming modes at the stationary growth phase when the maximum intracellular neutral lipid content was observed. Our results provide the potential exploitation of microalgal swimming signatures as possible indicators of the cultivation conditions and the timing of microalgal harvest to maximize the lipid yield for biofuel production. The findings can also be implemented to explore the production of food and antibiotics from other microalgal metabolites with low energy costs.

Increased Denitrification Rates Associated with Shifts in Prokaryotic Community Composition Caused by Varying Hydrologic Connectivity.

While modern developments in agriculture have allowed for increases in crop yields and rapid human population growth, they have also drastically altered biogeochemical cycles, including the biotransformation of nitrogen. Denitrification is a critical process performed by bacteria and fungi that removes nitrate in surface waters, thereby serving as a potential natural remediation strategy. We previously reported that constant inundation resulted in a coupling of denitrification gene abundances with denitrification rates in sediments, but these relationships were not maintained in periodically-inundated or non-inundated environments. In this study, we utilized Illumina next-generation sequencing to further evaluate how the microbial community responds to these hydrologic regimes and how this community is related to denitrification rates at three sites along a creek in an agricultural watershed over 2 years. The hydrologic connectivity of the sampling location had a significantly greater influence on the denitrification rate (P = 0.010), denitrification gene abundances (P < 0.001), and the prokaryotic community (P < 0.001), than did other spatiotemporal factors (e.g., creek sample site or sample month) within the same year. However, annual variability among denitrification rates was also observed (P < 0.001). Furthermore, the denitrification rate was significantly positively correlated with water nitrate concentration (Spearman's ρ = 0.56, P < 0.0001), denitrification gene abundances (ρ = 0.23-0.47, P ≤ 0.006), and the abundances of members of the families Burkholderiaceae, Anaerolinaceae, Microbacteriaceae, Acidimicrobineae incertae sedis, Cytophagaceae, and Hyphomicrobiaceae (ρ = 0.17-0.25, P ≤ 0.041). Prokaryotic community composition accounted for the least amount of variation in denitrification rates (22%), while the collective influence of spatiotemporal factors and gene abundances accounted for 37%, with 40% of the variation related to interactions among all parameters. Results of this study suggest that the hydrologic connectivity at each location had a greater effect on the prokaryotic community than did spatiotemporal differences, where inundation is associated with shifts favoring increased denitrification potential. We further establish that while complex interactions among the prokaryotic community influence denitrification, the link between hydrologic connectivity, microbial community composition, and genetic potential for biogeochemical cycling is a promising avenue to explore hydrologic remediation strategies such as periodic flooding.

Measurement and Modeling of Denitrification in Sand-Bed Streams under Various Land Uses.

Although many studies have measured denitrification in stream sediments, few have utilized these data with local water column and sediment measurements to develop a predictive model for NO uptake. In this study, sediment denitrification was measured from cores in five streams under various land uses in south-central Minnesota using denitrification enzyme activity (DEA) assays and amplification of the gene via real-time, quantitative polymerase chain reaction. Hydraulic and environmental variables were measured in the vicinity of the sediment cores to evaluate the influence of fluid flow and chemical variables on denitrification activity. Potential denitrification rates measured using DEA assays ranged from 0.02 to 10.1 mg N m h, and the abundance of the denitrifier gene was positively correlated with these measurements ( = 0.79, < 0.001) for most of the streams studied. A predictive model to determine NO uptake via denitrification was derived, implementing dimensional analysis of variables that mediate denitrification in sand-bed streams. The proposed model explained 75% of the variability in DEA rates. The results of this study show that denitrification is most dependent on the distribution of sediment organic matter, interstitial pore space, and stream hydraulic characteristics, including shear velocity at the sediment-water interface and stream depth.

Microalga propels along vorticity direction in a shear flow.

Using high-speed digital holographic microscopy and microfluidics, we discover that, when encountering fluid flow shear above a threshold, unicellular green alga Dunaliella primolecta migrates unambiguously in the cross-stream direction that is normal to the plane of shear and coincides with the local fluid flow vorticity. The flow shear drives motile microalgae to collectively migrate in a thin two-dimensional horizontal plane and consequently alters the spatial distribution of microalgal cells within a given suspension. This shear-induced algal migration differs substantially from periodic rotational motion of passive ellipsoids, known as Jeffery orbits, as well as gyrotaxis by bottom-heavy swimming microalgae in a shear flow due to the subtle interplay between torques generated by gravity and viscous shear. Our findings could facilitate mechanistic solutions for modeling planktonic thin layers and sustainable cultivation of microalgae for human nutrition and bioenergy feedstock.

Glossosoma nigrior (Trichoptera: Glossosomatidae) respiration in moving fluid.

Laboratory measurements of dissolved oxygen (DO) uptake by Glossosoma nigrior Banks were conducted in a sealed, recirculating flume under variable fluid flow velocities. Measurements were performed in similar water temperatures, DO concentrations and fluid flow velocities to field conditions in the stream where the larvae were obtained. Total oxygen uptake by both cased larvae and corresponding cases without larvae were quantified. An increased fluid flow velocity corresponded to an increased larval DO uptake rate. Oxygen uptake by the larval cases alone was not as sensitive to changes in the Peclet (Pe) number, the dimensionless ratio of advective to diffusive DO transport, as uptake by larvae themselves. The flux of DO to larvae and their cases was up to seven times larger in a moving fluid in comparison to non-moving fluid conditions in the proximity of larvae for 087, larvae typically remained in their cases. This indicates that oxygen delivery to the larvae at low Pe is insufficient to satisfy the respiratory demands of cased larvae.

Algal swimming velocities signal fatty acid accumulation.

The use of microalgae for biofuel production will be beneficial to society if we can produce biofuels at large scales with minimal mechanical energy input in the production process. Understanding micro-algal physiological responses under variable environmental conditions in bioreactors is essential for the optimization of biofuel production. We demonstrate that measuring micro-algal swimming speed provides information on culture health and total fatty acid accumulation. Three strains of Chlamydomonas reinhardtii were grown heterotrophically on acetate and subjected to various levels of nitrogen starvation. Other nutrient levels were explored to determine their effect on micro-algal kinetics. Swimming velocities were measured with two-dimensional micro-particle tracking velocimetry. The results show an inverse linear relationship between normalized total fatty acid mass versus swimming speed of micro-algal cells. Analysis of RNA sequencing data confirms these results by demonstrating that the biological processes of cell motion and the generation of energy precursors are significantly down-regulated. Experiments demonstrate that changes in nutrient concentration in the surrounding media also affect swimming speed. The findings have the potential for the in situ and indirect assessment of lipid content by measuring micro-algal swimming kinetics.

Three-dimensional lake water quality modeling: sensitivity and uncertainty analyses.

Two sensitivity and uncertainty analysis methods are applied to a three-dimensional coupled hydrodynamic-ecological model (ELCOM-CAEDYM) of a morphologically complex lake. The primary goals of the analyses are to increase confidence in the model predictions, identify influential model parameters, quantify the uncertainty of model prediction, and explore the spatial and temporal variabilities of model predictions. The influence of model parameters on four model-predicted variables (model output) and the contributions of each of the model-predicted variables to the total variations in model output are presented. The contributions of predicted water temperature, dissolved oxygen, total phosphorus, and algal biomass contributed 3, 13, 26, and 58% of total model output variance, respectively. The fraction of variance resulting from model parameter uncertainty was calculated by two methods and used for evaluation and ranking of the most influential model parameters. Nine out of the top 10 parameters identified by each method agreed, but their ranks were different. Spatial and temporal changes of model uncertainty were investigated and visualized. Model uncertainty appeared to be concentrated around specific water depths and dates that corresponded to significant storm events. The results suggest that spatial and temporal variations in the predicted water quality variables are sensitive to the hydrodynamics of physical perturbations such as those caused by stream inflows generated by storm events. The sensitivity and uncertainty analyses identified the mineralization of dissolved organic carbon, sediment phosphorus release rate, algal metabolic loss rate, internal phosphorus concentration, and phosphorus uptake rate as the most influential model parameters.

Kinetic responses of Dunaliella in moving fluids.

The objective of this work was to quantify the kinetic behavior of Dunaliella primolecta (D. primolecta) subjected to controlled fluid flow under laboratory conditions. In situ velocities of D. primolecta were quantified by micron-resolution particle image velocimetry and particle tracking velocimetry. Experiments were performed under a range of velocity gradients and corresponding energy dissipation levels at microscopic scales similar to the energy dissipation levels of natural aquatic ecosystems. An average swimming velocity of D. primolecta in a stagnant fluid was 41 microm/s without a preferential flow direction. In a moving fluid, the sample population velocities of D. primolecta follow a log-normal distribution. The variability of sample population velocities was maximal at the highest fluid flow velocity in the channel. Local fluid velocity gradients inhibited the accrual of D. primolecta by twofold 5 days after the initiation of the experiment in comparison to the non-moving fluid control experiment.

Dissolved oxygen measurements in aquatic environments: the effects of changing temperature and pressure on three sensor technologies.

Dissolved oxygen (DO) is probably the most important parameter related to water quality and biological habitat in aquatic environments. In situ DO sensors are some of the most valuable tools used by scientists and engineers for the evaluation of water quality in aquatic ecosystems. Presently, we cannot accurately measure DO concentrations under variable temperature and pressure conditions. Pressure and temperature influence polarographic and optical type DO sensors compared to the standard Winkler titration method. This study combines laboratory and field experiments to compare and quantify the accuracy and performance of commercially available macro and micro Clark-type oxygen sensors as well as optical sensing technology to the Winkler method under changing pressure and temperature conditions. Field measurements at various lake depths revealed sensor response time up to 11 min due to changes in water temperature, pressure, and DO concentration. Investigators should account for transient response in DO sensors before measurements are collected at a given location. We have developed an effective model to predict the transient response time for Clark-type oxygen sensors. The proposed procedure increases the accuracy of DO data collected in situ for profiling applications.

Do microscopic organisms feel turbulent flows?

Microscopic organisms in aquatic environments are continuously exposed to a variety of physical and chemical conditions. Traditionally, it is accepted that due to their small size the physiology of microscopic organisms is not affected by the moving fluid at their scale. In this study, we demonstrate that the small-scale turbulence significantly modulates algal and bacterial nutrient uptake and growth in comparison to still-water control. The rate of energy dissipation emerges as a physically based scaling parameter integrating turbulence across a range of scales and microscopic organism responses at the cell level. Microbiological laboratory tests and bioassays do not consider fluid motion as an important variable in quantifying the physiological responses of microorganisms. A conceptual model of how to integrate the fluid motion in Monod-type kinetics is proposed. We anticipate our findings will encourage researchers to reconsider the laboratory protocols and modeling procedures in the analysis of microorganism physiological responses to changing physical and chemical environments by integrating the effect of turbulence.

Enhanced uptake of dissolved oxygen and glucose by Escherichia coli in a turbulent flow.

Laboratory experiments were conducted to study the effect of turbulence on Escherichia coli cells in an oscillating grid reactor under conditions of no oxygen transfer to the liquid phase. Fluid flow was quantified at a submillimeter resolution using a particle image velocimetry measuring technique. The root-mean-square estimates of the velocity gradient tensor components indicated the dominance of shear rate deformation in the fluid surrounding E. coli. The E. coli growth rate, dissolved oxygen (DO), and glucose uptake rates were facilitated by fluid-flow energy dissipation in the turbulent fluid. The Kolmogorov length scale (eta(K)) and velocity (u( K )) underlined characteristic scales at which enhanced DO and glucose uptake by E. coli were determined in a turbulent flow in comparison to still-water controls. A first-order power-law relation between the mass transport to the cells and the moving fluid is developed. The combined effects of the enhanced rate of strain at eta(K) scale and uniform velocity at u(K) determined the facilitated DO and glucose fluxes to E. coli. The mass transport to the E. coli was modeled by the Sherwood (Sh)-Péclet (Pe) number relationship by Sh = 1 + 1.08Pe(uK)(0.62) where Pe(uK) is the Péclet number defined by the u(K) velocity scale. The proposed first-order model described experimental data fairly well.

Enhancement and inhibition of denitrification by fluid-flow and dissolved oxygen flux to stream sediments.

Microscale measurements of nitrate (NO3-) and dissolved oxygen (DO) concentrations in sediments were made in a laboratory channel under turbulent fluid-flow conditions to examine the effects of DO flux on denitrification rates. DO concentrations and flux within sediments increased with increasing velocity in the surface water. Under low fluid-flow conditions (shear stress velocity, u* < 0.23 cm s(-1)), increasing velocity increased NO3- loss from the bulk flow. For high fluid-flow conditions (u* > 0.39 cm s(-1)), increasing velocity inhibited NO3-loss. Sediment cores were collected and sliced to measure the depth distribution of denitrifying biomass in sediments. Quantities of nirK and nirS genes were higher within the surface layer and decreased with depth in the sediments. Microscale concentration profiles of DO and NO3- revealed that denitrification occurs within a thin region just below the oxic-anoxic interface in sediments. The interplay of mass transfer and DO flux generated threshold conditions for NO3- loss by denitrification. These results suggest that for a given sediment and environmental conditions (chemical, physical, microbiological), there exists an optimal range in velocities for enhancing denitrification in aquatic systems.

Laboratory study of heavy metal phytoremediation by three wetland macrophytes.

Detention ponds and constructed wetlands have proven to be effective in reducing peak stormwater runoff volume and flow, and recent interest has extended to utilizing them to improve stormwater runoff quality. A review of stormwater runoff studies indicated that lead, zinc, copper, cadmium, phosphorus, and chloride are contaminants of primary concern. In laboratory settings, the uptake of contaminants by three wetland plant species, Glyceria grandis, Scirpus validus, and Spartina pectinata, was examined and removal rates from nutrient solutions inflow and nonflow reactors were measured. The removal rates varied by plant species and target contaminant, and no one species was the best accumulator of all six contaminants. Belowground tissues of all three species accumulated higher concentrations of the four heavy metals and aboveground tissues accumulated higher concentrations of phosphorus and chloride. Plants grown in flow reactors showed significantly higher accumulation rates than those grown in nonflow reactors. Also, plants grown hydroponically accumulated higher concentrations of the six target contaminants than those grown in sand reactors. However, those grown in sand had a much greater increase of biomass and removed a greater mass of the six target contaminants. Removal rates measured in these experiments can be used to design detention ponds to maximize stormwater remediation.

Advective velocity and energy dissipation rate in an oscillatory flow.

Characterizing the transport processes at the sediment-water interface along sloping boundaries in lakes and reservoirs is of fundamental interest in lake and reservoir water quality management. The turbulent bottom boundary layer (TBBL) along a slope, induced by the breaking of internal waves in a linearly stratified fluid, was investigated through laboratory measurements. Fast response micro-scale conductivity and temperature probes in conjunction with laser-Doppler velocimetry were used to measure the time series of salinity, temperature, and velocity along a sloping boundary. Turbulent energy spectra were computed from the velocity data using a time-dependent advective velocity and Taylor's hypothesis. The energy spectra were used to estimate the energy dissipation rate at different positions in the TBBL. The advective velocity in this near-zero mean shear flow is based on an integral time scale (T(int)). The integral time scale is related to the average frequency of the spectral energy density of the flow velocity. The energy dissipation rate estimated from the variable advective velocity with an averaging time window equal to the integral time scale (T=T(int)) was 43% higher than the energy dissipation rate estimated from a constant advective velocity. The estimated dissipation rates with T=T(int) were comparable to values obtained by curve-fitting a theoretical Batchelor spectrum for the temperature gradient spectra. This study proposes the integral time scale to be used for the oscillatory flows as (a) a time-averaging window to estimate the advective velocity and associated energy dissipation level, and (b) a normalizing parameter in the energy spectrum.