Planktonic communities and chaotic advection in dynamical models of Langmuir circulation

A deterministic mechanism for the production of plankton patches within a typical medium scale oceanic structure is proposed and investigated. By direct numerical simulation of a simple model of Langmuir circulation we quantify the effects of unsteady flows on planktonic communities and demonstrate their importance. Two qualitatively different zones within the flow are identified: chaotic regions that help to spread plankton and locally coherent regions, that do not mix with the chaotic regions and which persist for long periods of time. The relative importance of these regions to both phytoplankton and zooplankton is investigated, taking into account variations in plankton buoyancy. In particular, species-specific retention zone structure is discussed in relation to variations in environmental forcing

A tale of three taxes: photo-gyro-gravitactic bioconvection

The term bioconvection encapsulates the intricate patterns in concentration, due to hydrodynamic instabilities, that may arise in suspensions of non-neutrally buoyant, biased swimming microorganisms. The directional bias may be due to light (phototaxis), gravity (gravitaxis), a combination of viscous and gravitational torques (gyrotaxis) or other taxes. The aim of this study is to quantify experimentally the wavelength of the initial pattern to form from an initially well-mixed suspension of unicellular, swimming green algae as a function of concentration and illumination. As this is the first such study, it is necessary to develop a robust and meticulous methodology to achieve this end. The phototactic, gyrotactic and gravitactic alga Chlamydomonas augustae was employed, with various red or white light intensities from above or below, as the three not altogether separable taxes were probed. Whilst bioconvection was found to be unresponsive to changes in red light, intriguing trends were found for pattern wavelength as a function of white light intensity, depending critically on the orientation of the illumination. These trends are explored to help unravel the mechanisms. Furthermore, comparisons are made with theoretical predictions of initial wavelengths from a recent model of photo-gyrotaxis, encouragingly revealing good qualitative agreement

Analytical approximations for the orientation distribution of small dipolar particles in steady shear flows

Analytic approximations are obtained to solutions of the steady Fokker-Planck equation describing the probability density functions for the orientation of dipolar particles in a steady, low-Reynolds-number shear flow and a uniform external field. Exact computer algebra is used to solve the equation in terms of a truncated spherical harmonic expansion. It is demonstrated that very low orders of approximation are required for spheres but that spheroids introduce resolution problems in certain flow regimes. Moments of the orientation probability density function are derived and applications to swimming cells in bioconvection are discussed. A separate asymptotic expansion is performed for the case in which spherical particles are in a flow with high vorticity, and the results are compared with the truncated spherical harmonic expansion. Agreement between the two methods is excellent

Non-linear bioconvection in a deep suspension of gyrotactic swimming micro-organisms

The non-linear structure of deep, stochastic, gyrotactic bioconvection is explored. A linear analysis is reviewed and a weakly non-linear analysis justifies its application by revealing the supercritical nature of the bifurcation. An asymptotic expansion is used to derive systems of partial differential equations for long plume structures which vary slowly with depth. Steady state and travelling wave solutions are found for the first order system of partial differential equations and the second order system is manipulated to calculate the speed of vertically travelling pulses. Implications of the results and possibilities of experimental validation are discussed

The orientation of swimming bi-flagellates in shear flows

Biflagellated algae swim in mean directions that are governed by their environments. For example, many algae can swim upward on average (gravitaxis) and toward downwelling fluid (gyrotaxis) via a variety of mechanisms. Accumulations of cells within the fluid can induce hydrodynamic instabilities leading to patterns and flow, termed bioconvection, which may be of particular relevance to algal bioreactors and plankton dynamics. Furthermore, knowledge of the behavior of an individual swimming cell subject to imposed flow is prerequisite to a full understanding of the scaled-up bulk behavior and population dynamics of cells in oceans and lakes; swimming behavior and patchiness will impact opportunities for interactions, which are at the heart of population models. Hence, better estimates of population level parameters necessitate a detailed understanding of cell swimming bias. Using the method of regularized Stokeslets, numerical computations are developed to investigate the swimming behavior of and fluid flow around gyrotactic prolate spheroidal biflagellates with five distinct flagellar beats. In particular, we explore cell reorientation mechanisms associated with bottom-heaviness and sedimentation and find that they are commensurate and complementary. Furthermore, using an experimentally measured flagellar beat for Chlamydomonas reinhardtii, we reveal that the effective cell eccentricity of the swimming cell is much smaller than for the inanimate body alone, suggesting that the cells may be modeled satisfactorily as self-propelled spheres. Finally, we propose a method to estimate the effective cell eccentricity of any biflagellate when flagellar beat images are obtained haphazardly

Synthetizing hydrodynamic turbulence from noise: formalism and applications to plankton dynamics

We present an analytical scheme, easily implemented numerically, to generate synthetic Gaussian 2D turbulent flows by using linear stochastic partial differential equations, where the noise term acts as a random force of well-prescribed statistics. This methodology leads to a divergence-free, isotropic, stationary and homogeneous velocity field, whose characteristic parameters are well reproduced, in particular the kinematic viscosity and energy spectrum. This practical approach to tailor a turbulent flow is justified by its versatility when analizing different physical processes occurring in advectely mixed systems. Here, we focuss on an application to study the dynamics of Planktonic populations in the ocean

The interaction of thin-film flow, bacterial swarming and cell differentiation in colonies of Serratia liquefaciens

The rate of expansion of bacterial colonies of S. liquefaciens is investigated in terms of a mathematical model that combines biological as well as hydrodynamic processes. The relative importance of cell differentiation and production of an extracellular wetting agent to bacterial swarming is explored using a continuum representation. The model incorporates aspects of thin film flow with variable suspension viscosity, wetting, and cell differentiation. Experimental evidence suggests that the bacterial colony is highly sensitive to its environment and that a variety of mechanisms are exploited in order to proliferate on a variety of surfaces. It is found that a combination of effects are required to reproduce the variation of bacterial colony motility over a large range of nutrient availability and medium hardness

Bugs on a Slippery Plane : Understanding the Motility of Microbial Pathogens with Mathematical Modelling

Many pathogenic microorganisms live in close association with surfaces, typically in thin films that either arise naturally or that they themselves create. In response to this constrained environment, the cells adjust their behaviour and morphology, invoking communication channels and inducing physical phenomena that allow for rapid colonization of biomedically relevant surfaces or the promotion of virulence factors. Thus, it is very important to measure and theoretically understand the key mechanisms for the apparent advantage obtained from swimming in thin films. We discuss experimental measurements of flows around a peritrichously flagellated bacterium constrained in a thin film, derive a simplified mathematical theory and Green's functions for flows in a thin film with general slip boundary conditions, and establish connections between theoretical and experimental results. This article aims to highlight the importance of mathematics as a tool to unlock qualitative mechanisms associated with experimental observations in the medical and biological sciences

Taylor dispersion of gyrotactic swimming micro-organisms in a linear flow

The theory of generalized Taylor dispersion for suspensions of Brownian particles is developed to study the dispersion of gyrotactic swimming micro-organisms in a linear shear flow. Such creatures are bottom-heavy and experience a gravitational torque which acts to right them when they are tipped away from the vertical. They also suffer a net viscous torque in the presence of a local vorticity field. The orientation of the cells is intrinsically random but the balance of the two torques results in a bias toward a preferred swimming direction. The micro-organisms are sufficiently large that Brownian motion is negligible but their random swimming across streamlines results in a mean velocity together with diffusion. As an example, we consider the case of vertical shear flow and calculate the diffusion coefficients for a suspension of the alga <i>Chlamydomonas nivalis</i>. This rational derivation is compared with earlier approximations for the diffusivity