Simple models of the chemical field around swimming plankton

International audienceThe chemical field around swimming plankton depends on the swimming style and speed of the organism and the processes affecting uptake or exudation of chemicals by the organism. Here we present a simple model for the flow field around a neutrally buoyant self-propelled organism at low Reynolds number, and numerically calculate the chemical field around the organism. We show how the concentration field close to the organism and the mass transfer rates vary with swimming speed and style for Dirichlet (diffusion limited transport) boundary conditions. We calculate how the length of the chemical wake, defined as being the distance at which the chemical field drops to 10% of the surface concentration of the organism when stationary, varies with swimming speed and style for both Dirichlet and Neumann (production limited) boundary conditions. For Dirichlet boundary conditions, the length of the chemical wake increases with increasing swimming speed, and the self-propelled organism displays a significantly longer wake than the towed-body model. For the Neumann boundary conditions the converse is true; because swimming enhances the transport of the chemical away from the organism, the surface concentration of chemical is reduced and thus the wake length is reduced

Enhanced sedimentation of elongated plankton in simple flows

Negatively buoyant phytoplankton play an important role in the sequestration of CO_2 from the atmo-sphere and are fundamental to the health of the world’s fisheries. However, there is still much to discoveron transport mechanisms from the upper photosynthetic regions to the deep ocean. In contrast to intuitive expectations that mixing increases plankton residence time in light-rich regions, recent experimental and computational evidence suggests that turbulence can actually enhance sedimentation of negatively buoyant diatoms. Motivated by these studies we dissect the enhanced sedimentation mechanisms using the simplest possible two-dimensional flows, avoiding expensive computations and obfuscation. In particular, we find that in vertical shear, preferential flow alignment and aggregation in down-welling regions both increase sedimentation, whereas horizontal shear reduces the sedimentation due only to alignment. However the magnitude of the shear does not affect the sedimentation rate. In simple vertical Kolmogorov flow elongated particles also have an enhanced sedimentation speed as they spend more time in down-welling regions of the flow with vertically aligned orientation, an effect that increases with the magnitude of shear. An additional feature is identified in horizontal Kolomogorov flow, whereby the impact of shear-dependent sedimentation speed is to cause aggregation in regions of high-shear where the sedimentation speed is minimum. In cellular flow, there is an increase in mean sedimentation speed with aspect ratio and shear strength associated with aggregation in down-welling regions. Furthermore, spatially projected trajectories can intersect and give rise to chaotic dynamics, which is associated with a depletion of particles within so called retention zones

Inherent High Correlation of Individual Motility Enhances Population Dispersal in a Heterotrophic, Planktonic Protist

Quantitative linkages between individual organism movements and the resulting population distributions are fundamental to understanding a wide range of ecological processes, including rates of reproduction, consumption, and mortality, as well as the spread of diseases and invasions. Typically, quantitative data are collected on either movement behaviors or population distributions, rarely both. This study combines empirical observations and model simulations to gain a mechanistic understanding and predictive ability of the linkages between both individual movement behaviors and population distributions of a single-celled planktonic herbivore. In the laboratory, microscopic 3D movements and macroscopic population distributions were simultaneously quantified in a 1L tank, using automated video- and image-analysis routines. The vertical velocity component of cell movements was extracted from the empirical data and used to motivate a series of correlated random walk models that predicted population distributions. Validation of the model predictions with empirical data was essential to distinguish amongst a number of theoretically plausible model formulations. All model predictions captured the essence of the population redistribution (mean upward drift) but only models assuming long correlation times (minute), captured the variance in population distribution. Models assuming correlation times of 8 minutes predicted the least deviation from the empirical observations. Autocorrelation analysis of the empirical data failed to identify a de-correlation time in the up to 30-second-long swimming trajectories. These minute-scale estimates are considerably greater than previous estimates of second-scale correlation times. Considerable cell-to-cell variation and behavioral heterogeneity were critical to these results. Strongly correlated random walkers were predicted to have significantly greater dispersal distances and more rapid encounters with remote targets (e.g. resource patches, predators) than weakly correlated random walkers. The tendency to disperse rapidly in the absence of aggregative stimuli has important ramifications for the ecology and biogeography of planktonic organisms that perform this kind of random walk

Bifurcation and stability of downflowing gyrotactic micro-organism suspensions in a vertical pipe

In the experiment that first demonstrated gyrotactic behaviour of bottom-heavy swimming microalgae (e.g. Chlamydomonas), Kessler (Nature, vol. 313, 1985, pp. 218-220) showed that a beam-like structure, often referred to as a gyrotactic plume, would spontaneously appear from a suspension of gyrotactic swimmers in a downflowing pipe. Such a plume is prone to an instability to form blips. This work models the gyrotactic plume as a steady parallel basic state and its subsequent breakdown into blips as an instability, employing both the Generalised Taylor Dispersion (GTD) theory and the Fokker-Planck model for comparison. Upon solving for the basic state, it is discovered that the steady plume solution undergoes sophisticated bifurcations. When there is no net flow, there exists a non-trivial solution of the plume structure other than the stationary uniform suspension, stemming from a transcritical bifurcation with the average cell concentration. When a net downflow is prescribed, there exists a cusp bifurcation. Furthermore, there is a critical concentration, at which the cell concentration at the centre would blow up for the GTD model. The subsequent stability analysis using the steady plume solution shows that the Fokker-Planck model is inconsistent with what was experimentally observed, as it predicts stabilisation of axisymmetric blips at high concentration of the plume and destabilisation of the first non-axisymmetric mode at low flow rates

Elongation enhances migration through hydrodynamic shear

Many species of plankton migrate through the water column to avoid predation and acquire resources available only at depth. These vertical migrations, however, can be thwarted by flow which is ubiquitous in the ocean and acts to reorient the motility of swimming plankton in different directions. The torques exerted on an organism by hydrodynamic shear are a strong function of its shape - while spherical organisms only experience the overturning effect of vorticity, more elongated organisms experience additional torques that tend to align them in the direction of principal strain. Recent simulations of directed migration through turbulence indicate that organisms with larger aspect ratios are capable of faster vertical migration, however, the underlying physical processes are not well understood. Here, we use simple models of flow to study how elongation affects the capacity of a swimmer to vertically migrate through shear. We explore how the orientation of a migrating swimmer in simple shear depends both on its aspect ratio and non-dimensional stability, highlighting where a unique stable upwards equilibrium is globally attracting, and where particles can undergo periodic orbits. In addition, we identify that stable up and down equilbria can coexist when there is a non-zero vertical component of vorticity. We go on to investigate how elongation affects the transport and spatial distribution of a migrating swimmer in an inhomogeneous Kolmogorov shear flow. Our results show that a swimmer’s shape can profoundly affect its mean field behaviour in flow and tendency to aggregate in regions of high shear. These findings reveal how flow can select for the elongated morphologies commonly found in marine plankton and may also provide new insights to rationally design microrobots to navigate ambient flows

Sedimentation of inertia-less prolate spheroids in homogenous isotropic turbulence with application to non-motile phytoplankton

Phytoplankton are the foundation of aquatic food webs. Through photosynthesis, phytoplankton draw down CO2 at magnitudes equivalent to forests and other terrestrial plants and convert it to organic material that is then consumed by other planktonic organisms in higher trophic levels. Mechanisms that affect local concentrations and velocities are of primary significance to many encounter-based processes in the plankton, including prey-predator interactions, fertilization and aggregate formation. We report results from simulations of sinking phytoplankton, considered as elongated spheroids, in homogenous isotropic turbulence to answer the question of whether trajectories and velocities of sinking phytoplankton are altered by turbulence. We show in particular that settling spheroids with physical characteristics similar to those of diatoms weakly cluster and preferentially sample regions of downwelling flow, corresponding to an increase of the mean settling speed with respect to the mean settling speed in quiescent fluid. We explain how different parameters can affect the settling speed and what underlying mechanisms might be involved. Interestingly, we observe that the increase in the aspect ratio of the prolate spheroids can affect the clustering and the average settling speed of particles by two mechanisms: first is the effect of aspect ratio on the rotation rate of the particles, which saturates faster than the second mechanism of increasing drag anisotropy