Gyrotactic phytoplankton in laminar and turbulent flows: A dynamical systems approach.

Gyrotactic algae are bottom heavy, motile cells whose swimming direction is determined by a balance between a buoyancy torque directing them upwards and fluid velocity gradients. Gyrotaxis has, in recent years, become a paradigmatic model for phytoplankton motility in flows. The essential attractiveness of this peculiar form of motility is the availability of a mechanistic description which, despite its simplicity, revealed predictive, rich in phenomenology, easily complemented to include the effects of shape, feedback on the fluid and stochasticity (e.g., in cell orientation). In this review we consider recent theoretical, numerical and experimental results to discuss how, depending on flow properties, gyrotaxis can produce inhomogeneous phytoplankton distributions on a wide range of scales, from millimeters to kilometers, in both laminar and turbulent flows. In particular, we focus on the phenomenon of gyrotactic trapping in nonlinear shear flows and in fractal clustering in turbulent flows. We shall demonstrate the usefulness of ideas and tools borrowed from dynamical systems theory in explaining and interpreting these phenomena.

Turbulent fluid acceleration generates clusters of gyrotactic microorganisms.

The motility of microorganisms is often biased by gradients in physical and chemical properties of their environment, with myriad implications on their ecology. Here we show that fluid acceleration reorients gyrotactic plankton, triggering small-scale clustering. We experimentally demonstrate this phenomenon by studying the distribution of the phytoplankton Chlamydomonas augustae within a rotating tank and find it to be in good agreement with a new, generalized model of gyrotaxis. When this model is implemented in a direct numerical simulation of turbulent flow, we find that fluid acceleration generates multifractal plankton clustering, with faster and more stable cells producing stronger clustering. By producing accumulations in high-vorticity regions, this process is fundamentally different from clustering by gravitational acceleration, expanding the range of mechanisms by which turbulent flows can impact the spatial distribution of active suspensions.

Driving potential and noise level determine the synchronization state of hydrodynamically coupled oscillators.

Motile cilia are highly conserved structures in the evolution of organisms, generating the transport of fluid by periodic beating, through remarkably organized behavior in space and time. It is not known how these spatiotemporal patterns emerge and what sets their properties. Individual cilia are nonequilibrium systems with many degrees of freedom. However, their description can be represented by simpler effective force laws that drive oscillations, and paralleled with nonlinear phase oscillators studied in physics. Here a synthetic model of two phase oscillators, where colloidal particles are driven by optical traps, proves the role of the average force profile in establishing the type and strength of synchronization. We find that highly curved potentials are required for synchronization in the presence of noise. The applicability of this approach to biological data is also illustrated by successfully mapping the behavior of cilia in the alga Chlamydomonas onto the coarse-grained model.

Turbulence induces clustering and segregation of non-motile, buoyancy-regulating phytoplankton.

Turbulence plays a major role in shaping marine community structure as it affects organism dispersal and guides fundamental ecological interactions. Below oceanographic mesoscale dynamics, turbulence also impinges on subtle physical-biological coupling at the single cell level, setting a sea of chemical gradients and determining microbial interactions with profound effects on scales much larger than the organisms themselves. It has been only recently that we have started to disentangle details of this coupling for swimming microorganisms. However, for non-motile species, which comprise some of the most abundant phytoplankton groups on Earth, a similar level of mechanistic understanding is still missing. Here, we explore by means of extensive numerical simulations the interplay between buoyancy regulation in non-motile phytoplankton and cellular responses to turbulent mechanical cues. Using a minimal mechano-response model, we show how such a mechanism would contribute to spatial heterogeneity and affect vertical fluxes and trigger community segregation.

Laminar-turbulent boundary in plane Couette flow.

We apply the iterated edge-state tracking algorithm to study the boundary between laminar and turbulent dynamics in plane Couette flow at Re=400. Perturbations that are not strong enough to become fully turbulent or weak enough to relaminarize tend toward a hyperbolic coherent structure in state space, termed the edge state, which seems to be unique up to obvious continuous shift symmetries. The results reported here show that in cases where a fixed point has only one unstable direction, such as for the lower-branch solution in plane Couette flow, the iterated edge tracking algorithm converges to this state. They also show that the choice of initial state is not critical and that essentially arbitrary initial conditions can be used to find the edge state.

Statistical conservation law in two- and three-dimensional turbulent flows.

Particles in turbulence live complicated lives. It is nonetheless sometimes possible to find order in this complexity. It was proposed in Falkovich et al. [Phys. Rev. Lett. 110, 214502 (2013)] that pairs of Lagrangian tracers at small scales, in an incompressible isotropic turbulent flow, have a statistical conservation law. More specifically, in a d-dimensional flow the distance R(t) between two neutrally buoyant particles, raised to the power -d and averaged over velocity realizations, remains at all times equal to the initial, fixed, separation raised to the same power. In this work we present evidence from direct numerical simulations of two- and three-dimensional turbulence for this conservation. In both cases the conservation is lost when particles exit the linear flow regime. In two dimensions we show that, as an extension of the conservation law, an Evans-Cohen-Morriss or Gallavotti-Cohen type fluctuation relation exists. We also analyze data from a 3D laboratory experiment [Liberzon et al., Physica D 241, 208 (2012)], finding that although it probes small scales they are not in the smooth regime. Thus instead of 〈R-3〉, we look for a similar, power-law-in-separation conservation law. We show that the existence of an initially slowly varying function of this form can be predicted but that it does not turn into a conservation law. We suggest that the conservation of 〈R-d〉, demonstrated here, can be used as a check of isotropy, incompressibility, and flow dimensionality in numerical and laboratory experiments that focus on small scales.

Turbulence drives microscale patches of motile phytoplankton.

Patchiness plays a fundamental role in phytoplankton ecology by dictating the rate at which individual cells encounter each other and their predators. The distribution of motile phytoplankton species is often considerably more patchy than that of non-motile species at submetre length scales, yet the mechanism generating this patchiness has remained unknown. Here we show that strong patchiness at small scales occurs when motile phytoplankton are exposed to turbulent flow. We demonstrate experimentally that Heterosigma akashiwo forms striking patches within individual vortices and prove with a mathematical model that this patchiness results from the coupling between motility and shear. When implemented within a direct numerical simulation of turbulence, the model reveals that cell motility can prevail over turbulent dispersion to create strong fractal patchiness, where local phytoplankton concentrations are increased more than 10-fold. This 'unmixing' mechanism likely enhances ecological interactions in the plankton and offers mechanistic insights into how turbulence intensity impacts ecosystem productivity.

Transient turbulence in plane Couette flow.

Plane Couette flow, the flow between two parallel plates moving in opposite directions, belongs to the group of shear flows where turbulence occurs while the laminar profile is stable. Experimental and numerical studies show that at intermediate Reynolds numbers turbulence is transient and that the lifetimes are distributed exponentially. However, these studies have remained inconclusive about a divergence in lifetimes above a critical Reynolds number. The extensive numerical results for flow in a box of width 2pi and length 8pi presented here cover observation times up to 12,000 units and show that while the lifetimes increase rapidly with Reynolds number, they do not indicate a divergence and therefore no transition to persistent turbulence.