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Bacteria and phytoplankton are abundant in aquatic environments, forming the base of the food web and mediating elemental cycling at a global scale. Understanding the interactions these microorganisms have with their turbulent fluid environments is an active area of research, largely conducted in laboratory-based flow experiments. In this work, we provide an open-source design and rigorous flow characterization for a 1L, dual oscillating grid turbulence facility, the smallest volume facility to date which produces near-isotropic, homogeneous turbulence. We optimized the tank geometry (grid-to-grid and grid-to-wall spacing), the grid geometry (for both classical and fractal grids: effective mesh size, blockage ratio, and fractal grid parameters), and the grid forcing regimes (for both coupled, antiphase and decoupled, randomized forcing: frequency range, stroke range, and randomized forcing parameters) to minimize mean flows and to produce acceptably homogeneous and isotropic turbulence within the unique constraints of a litre-scale volume. We acquired particle image velocimetry (PIV) measurements for both classical and fractal grids across a wide range of grid forcing regimes. We discuss the resulting length- and timescales relevant to microorganism-flow interactions, from the integral to the Kolmogorov scales. Finally, we discuss how the range of turbulent kinetic energy (TKE) dissipation rates achieved across the operational space of the facility mimics oceanographic turbulence in a range of in situ conditions, from the nearshore to the open ocean. This facility meets a long-standing need in the oceanography community in which feasible experimental working volumes are constrained by labor-intensive culturing requirements for large volumes of aquatic bacteria and phytoplankton.

Turbulence can cause particles to accumulate within specific regions of the flow. One mechanism responsible for this phenomenon, called preferential concentration, consists in particle-fluid interactions yielding inhomogeneous spatial distribution of particles into clusters or depleted regions due to density difference or finite-size effects. In the case of living particles such as plankton, clustering may also originate from their motility or from their behavioral response to turbulent forcing. Preferential concentration of plankton has attracted much attention, because it is a key determinant of encounter rates and therefore relevant for a wide range of ecological processes. However, most studies have focused on microscopic cells, and consequently the case of larger organisms remains poorly studied. Here, we use high-performance particle tracking and three-dimensional Voronoï analysis to test for the emergence of clustering in the spatial distribution of calanoid copepods, the most important metazoans in the oceans in terms of biomass. We found that neither inertia nor motility resulted in significant departure from a random Poisson process over a range of turbulence intensity from very strong to moderate. However, we observed weak clustering in calm water, which may originate from hydrodynamic and olfactory interactions between organisms. Our results improve our understanding of fluid-particle interactions in the zooplankton and have important implications for the modeling of their encounter rates in turbulence.

We present a novel stochastic model for quantifying gross solids (GS) physical disintegration under varying turbulent flow conditions and used a unique experimental setup for model calibration and validation. The stochastic deterioration model predicts faeces size evolution over time. It conceptually entails the two main processes of solid fragmentation, namely breakage and erosion. Model parameters were calibrated on synthetic faeces and validated with real human ones. A cylindrical reactor was used, where turbulent flow was forced by an array of water jets and the physical disintegration of the faeces was monitored using a high speed camera. Image analysis of breakage experiments obtained under backlight illumination allowed determination of the evolution of the solids' size over time. The flow field in the reactor was characterised by particle image velocimetry (PIV) using tracer particles seeded into the water. We found different disintegration behaviours depending on turbulence intensity and water content of the solid. In conditions of low shear stress, dense solids hardly disintegrated. Generally, the model predictions mirrored the broad range in the solids disintegration rate imparted by the high variability in flow conditions and in solids characteristics. It is expected that, similar to our experiments, also in real sewer systems both flow conditions and solid characteristics are highly variable and the stochastic model can be tailored to capture this variability. We thus anticipate that the model can be integrated into existing sewer models predicting sewer flows and solids' movement. From these, shear stress, flow velocities and transport of individual solids can be inferred. The integration of the present solids disintegration model may provide better predictions of hot-spots for solids accumulation and blockages in sewers.

Significance Zooplankton possess narrow swimming capabilities, yet are capable of active locomotion amid turbulence. By decoupling the relative velocity of swimming zooplankton from that of the underlying flow, we provide evidence for an active adaptation that allows these small organisms to modulate their swimming effort in response to background flow. This behavioral response results in reduced diffusion at substantial turbulence intensity. Adjusting motility provides fitness advantage because it enables zooplankton to retain the benefits of self-locomotion despite the constraints enforced by turbulence transport. Vigorous swimming and reduced diffusion oppose turbulence advection, can directly affect the dispersal of zooplankton populations, and may help these organisms to actively control their distribution in dynamic environments., Calanoid copepods are among the most abundant metazoans in the ocean and constitute a vital trophic link within marine food webs. They possess relatively narrow swimming capabilities, yet are capable of significant self-locomotion under strong hydrodynamic conditions. Here we provide evidence for an active adaptation that allows these small organisms to adjust their motility in response to background flow. We track simultaneously and in three dimensions the motion of flow tracers and planktonic copepods swimming freely at several intensities of quasi-homogeneous, isotropic turbulence. We show that copepods synchronize the frequency of their relocation jumps with the frequency of small-scale turbulence by performing frequent relocation jumps of low amplitude that seem unrelated to localized hydrodynamic signals. We develop a model of plankton motion in turbulence that shows excellent quantitative agreement with our measurements when turbulence is significant. We find that, compared with passive tracers, active motion enhances the diffusion of organisms at low turbulence intensity whereas it dampens diffusion at higher turbulence levels. The existence of frequent jumps in a motion that is otherwise dominated by turbulent transport allows for the possibility of active locomotion and hence to transition from being passively advected to being capable of controlling diffusion. This behavioral response provides zooplankton with the capability to retain the benefits of self-locomotion despite turbulence advection and may help these organisms to actively control their distribution in dynamic environments. Our study reveals an active adaptation that carries strong fitness advantages and provides a realistic model of plankton motion in turbulence.

Zooplankton live in dynamic environments where turbulence may challenge their limited swimming abilities. How this interferes with fundamental behavioral processes remains elusive. We reconstruct simultaneously the trajectories of flow tracers and calanoid copepods and we quantify their ability to find mates when ambient flow imposes physical constrains on their motion and impairs their olfactory orientation. We show that copepods achieve high encounter rates in turbulence due to the contribution of advection and vigorous swimming. Males further convert encounters within the perception radius to contacts and then to mating via directed motion toward nearby organisms within the short time frame of the encounter. Inertial effects do not result in preferential concentration, reducing the geometric collision kernel to the clearance rate, which we model accurately by superposing turbulent velocity and organism motion. This behavioral and physical coupling mechanism may account for the ability of copepods to reproduce in turbulent environments., eLife, 9, ISSN:2050-084X

The increasing interest for behavioural investigations in aquatic toxicology has heightened the need for developing tools that allow realistic exposure conditions and provide robust quantitative data. Calanoid copepods dominate the zooplankton community in marine and brackish environments. These small organisms have emerged as attractive models because of the sensitivity of their behaviour to important environmental parameters and the significance of self-induced motion in their ecology. Estuarine copepods are particularly relevant in this context because of their incessant exposure to high levels of pollution. We used lipid nanocapsules to deliver sub-lethal concentrations of PAHs (pyrene, phenanthrene and fluoranthene) and PCB 153 into the digestive track of males and females Eurytemora affinis. This novel approach enabled us to achieve both contact and trophic exposure without using phytoplankton, and to expose copepods to small hydrophobic molecules without using organic solvent. We reconstructed the motion of many copepods swimming simultaneously by means of three-dimensional particle tracking velocimetry. We quantified the combined effects of contact and trophic toxicity by comparing the kinematic and diffusive properties of their motion immediately and after 3h and 24h of exposure. Despite the lack of toxicity of their excipients, both empty and loaded capsules increased swimming activity and velocity immediately after exposure. Laser microscopy imaging shows adhesion of nanocapsules on the exoskeleton of the animals, suggesting contact toxicity. The behavioural response resembles an escape reaction allowing copepods to escape stressful conditions. The contact toxicity of empty capsules and pollutants appeared to be additive and nanocapsules loaded with PCB caused the greatest effects. We observed a progressive accumulation of capsules in the digestive track of the animals after 3h and 24h of exposure, which suggests an increasing contribution of systemic toxicity. Nanocapsules filled with PAHs caused a smaller response compared to empty capsules, which we attribute to the narcotic properties of these toxicants. The sharp decrease in velocity after 24h of exposure to capsules loaded with PCB suggests physiological incapacitation following systemic toxicity. Clear differences are visible between genders in their response to empty and loaded capsules, for all exposure durations. Females appear to be less sensitive than males, suggesting different tolerance to stress conditions. Our results confirm the feasibility of using lipid nanocapsules to identify pollutant-induced behavioural alteration in the plankton. They also add new insights into the contact and systemic toxicity of common pollutants. We expect that our results will assist and evoke further research to develop suitable nanocarrier systems for behavioural testing.

We consider Lagrangian velocity differences of zooplankters swimming in still water and in turbulence. Using cumulants, we quantify the intermittency properties of their motion recorded using three-dimensional particle tracking velocimetry. Copepods swimming in still water display an intermittent behaviour characterized by a high probability of small velocity increments, and by stretched exponential tails. Low values arise from their steady cruising behaviour while heavy tails result from frequent relocation jumps. In turbulence, we show that at short time scales, the intermittency signature of active copepods clearly differs from that of the underlying flow, and reflects the frequent relocation jumps displayed by these small animals. Despite these differences, we show that copepods swimming in still and turbulent flow belong to the same intermittency class that can be modelled by a log-stable model with non-analytical cumulant generating function. Intermittency in swimming behaviour and relocation jumps may enable copepods to display oriented, collective motion under strong hydrodynamic conditions and thus, may contribute to the formation of zooplankton patches in energetic environments., The European Physical Journal E, 38 (10), ISSN:1292-8941, ISSN:1292-895X

Salinity is a major parameter in the ecology of estuarine copepods. However, its effects on swimming behavior are relatively unclear. The study of copepod swimming behavior requires a sufficient number of long and simultaneous trajectories to form a reliable basis for statistical analysis. Using a three-dimensional particle tracking technique, we observed changes in Pseudodiaptomus annandalei swimming behavior in response to a realistic salinity variation. The three adult states i.e. males, females and ovigerous females were actively swimming over the experimental range of salinity. However, an increase in salinity from 5 to 25 caused a decrease in ovigerous female speed and activity. In males and females, swimming speeds were higher at medium salinities and lower at salinities 5 and 25. Slower or faster swimming speeds were explained by variations in frequencies of resting periods (below 1 mm/s), slow swimming (between 1 and 8 mm/s) and fast swimming (above 8 mm/s). Swimming state residence times varied in concordance. Longer durations of resting periods were observed at slow swimming speeds, together with shorter durations of periods of slow and fast swimming. The degree of space occupation of trajectories was estimated through their fractal dimension and was unaffected by salinity. The relative importance of extreme events i.e. large amplitudes of velocity and acceleration indicated intermittency in behavior. Overall, our results show an optimal salinity for swimming activity and highlight differences between adult states. Results add insight into how salinity may influence copepod swimming behavior. Redirection of energy from swimming to osmoregulation when salinity diverges from the optimal range of the species is a possible explanation. Further tests on total metabolism are warranted to confirm this hypothesis.

International audience; Studies on the behavior of copepods require both an appropriate experimental design and the means to perform objectively verifiable numerical analysis. Despite the growing number of publications on copepod behavior, it has been difficult to compare these studies. In this study, we studied two species of copepods, Eurytemora affinis and Pseudodiaptomus annandalei, and employed recently developed scaling and non-scaling methodology to investigate the effects of density and volume on the swimming behavior of individual organisms in still water. We also compared the results of two- and three-dimensional projections of the swimming tracks. A combination of scale-dependent and scale-independent analysis was found to characterize a number of behavioral observations very effectively. We discovered that (i) density has no effect except to increase the time spent in the swimming state of "breaking", (ii) smaller volumes resulted in more complex trajectories, and larger volumes, like density, increased the time spent in the swimming state "breaking", and (iii) three-dimensional projections gave a more accurate estimation of speed and the time spent cruising. When only a vertical 2D projection was used, "cruising" could be confused with "sinking". These results indicate that both experimental conditions and the selection of 2D or 3D projection have important implications regarding the study of copepod behavior. The development of standardized procedures with which to compare the observations made in different studies is an issue of particular urgency.

Estuarine copepods experience strong and frequent salinity variations caused by alternating tides. Salinity is known to be a major parameter in the ecology of copepods from brackish waters but its effects on small scale behavior are relatively unclear. In this study, we used two-dimensional laboratory filming techniques to record swimming paths of the three reproductive stages (i.e. male, non-ovigerous female and ovigerous female) of Eurytemora affinis from the Seine estuary, under different salinity conditions. We show that the probability density of both the instantaneous swimming speed and the duration of the two most frequent swimming states (i.e. break and slow swimming) followed a power-law trend regardless of the salinity. Increase in salinity (i.e. 5, 15, 25 and 30) steadily decreased the instantaneous speed of E. affinis reproductive stages. At the extreme salinity of 0.5, the mean speed decreased for non-ovigerous females and ovigerous females but increased for males, when compared with higher salinities. Maximum speed was observed at salinity 0.5 for males and at salinity 5 for non-ovigerous females and ovigerous females. Low swimming speeds were associated with break events of high frequency and long duration. High swimming speeds were exhibited with cruising states of high frequency and break events of short duration. This supports laboratory and field studies indicating a preference of E. affinis for low to medium salinities.

Calanoid copepods represent a major component of the plankton community. These small animals reside in constantly flowing environments. Given the fundamental role of behaviour in their ecology, it is especially relevant to know how copepods perform in turbulent flows. By means of three-dimensional particle tracking velocimetry, we reconstructed the trajectories of hundreds of adult Eurytemora affinis swimming freely under realistic intensities of homogeneous turbulence. We demonstrate that swimming contributes substantially to the dynamics of copepods even when turbulence is significant. We show that the contribution of behaviour to the overall dynamics gradually reduces with turbulence intensity but regains significance at moderate intensity, allowing copepods to maintain a certain velocity relative to the flow. These results suggest that E. affinis has evolved an adaptive behavioural mechanism to retain swimming efficiency in turbulent flows. They suggest the ability of some copepods to respond to the hydrodynamic features of the surrounding flow. Such ability may improve survival and mating performance in complex and dynamic environments. However, moderate levels of turbulence cancelled gender-specific differences in the degree of space occupation and innate movement strategies. Our results suggest that the broadly accepted mate-searching strategies based on trajectory complexity and movement patterns are inefficient in energetic environments.

Estuarine waters contain a variety of chemicals which affect to various extents the behavior of aquatic organisms. Little is known, however, on the behavioral response of copepods. The present study shows the results of laboratory experiments investigating the immediate effects of sub-lethal concentrations of three commonly found contaminants on the three-dimensional swimming behavior of the estuarine calanoid copepod Eurytemora affinis. Nonylphenol at 2 μg L⁻¹, cadmium at 45 n gL⁻¹ and a mixture of low to medium molecular weight polycyclic aromatic hydrocarbons at 40 ng L⁻¹ all affected the swimming behavior of E. affinis adults, increasing both swimming speed and activity. In most cases, effects were observable within 30 min of exposure and persisted or faded during a period of depuration in uncontaminated water of similar duration. In ovigerous females exposed to Cd and PAHs, effects appeared to be more pronounced during the depuration period, suggesting that carrying ovisacs may impair recovery. We quantified differences in the distribution of swimming speed values by considering the relative frequencies of periods of break, slow and fast swimming and we observed a trend toward faster movements in the presence of pollutants. The degree of trajectory complexity, estimated through their fractal dimension, was unaffected by pollutants. Since both narcotic and non-narcotic pollutants induced hyperactivity, our results suggest that changes in behavior after a short-term exposure may be independent of the general mode of action of the chemicals. The increase in speed and activity resembles an escape reaction permitting copepods to evade stressful conditions. Overall, these results indicate that environment-relevant concentrations of pollutants can induce rapid changes in copepod behavior. Since behavioral processes represent a fundamental element in the ecology of copepods, our results raise concern about the effects of background levels of pollution on a major component of the plankton community. The long-term response of copepods to waterborne pollutants, their synergistic effects and their interactions with other environmental factors need further investigation.