Lagrangian Coherent Structures: applications to living systems. Bradly Alicea

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Lagrangian Coherent Structures: applications to living systems Bradly Alicea http://www.msu.edu/~aliceabr

Introduction: basic scientific problem Q: How can we quantify emergent structures? Use Lagrangian Coherent Structures (LCS) approach. * hard to capture higher-level relationships using simple pairwise comparisons or aggregate measurements (curse of dimensionality). * swarms, herds, colonies, and flocks: all share a common set of principles selforganized behaviors (weak, strong emergence). * behaviors, formation of structure occurs in a medium (air, liquid, etc) and so are analogous to or explicitly involve flows (mechanical representation). * interactions are also dynamic need a dynamical systems approach.

Introduction: key people and papers Key people/hot papers in this area: George Haller (MIT): http://georgehaller.com/, see Uncovering the Lagrangian Skeleton of Turbulence. PRL, 98, 144502 (2007). John Dabiri (Caltech): http://dabiri.caltech.edu/, see A Lagrangian approach to identifying vortex pinch-off. Chaos, 20, 017513 (2010). Emilie Tew Kai (IRD, France): see Top marine predators track Lagrangian coherent structures. PNAS, 106(20), 8245 8250 (2009).

Lagrangian introduction Lagrangian: function that summarizes system dynamics (special case of classical mechanics). L = T - V If Lagrangian known, equations of motion used instead, becomes PDE. Lagrange points around common COG. Courtesy: NASA www.scholarpedia.org/article/lagrangian-mechanics en.wikipedia.org/wiki/lagrangian T = kinetic energy, V = potential energy. * combines conservation of momentum with conservation of energy. First-order Lagrangian: constraints (motion energy) explicit in form of equations (Lagrangian multipliers). Second-order Lagrangian: constraints (motion energy) explicit in form of a generalized coordinate system (LCS method). Stationary action: assume well-defined trajectory bounded in space and time. S = S (Joules sec)

Finite Lyapunov exponents Key tool for LCS: finite-time and finite-size Lyapunov exponents (FTLE or FSLE): * FTLE = evolution of collective particle trajectories over time (find distance for given time interval). * FSLE = evolution of collective particle trajectories across space (find time for given set of distances). LCS is a flexible dynamical system: * Lagrangian representation of mechanics (special case of classical mechanics). Use where Eulerian representations are incomplete. Points in a flow have an initial position. The relationship between these points deformed (flow travels apart) over time: * distance between points quantified using Lyapunov exponent. * ridges form in flow map over time/space = coherent structures.

Finite Lyapunov exponents (con t) Finite Lyapunov exponent (related to γ, key parameter in dynamical systems) defined by: Lyapunov exponent (scalar value) Movement of particles (advection) Coordinate system (interpolation) http://amath.colorado.edu/index.php?page=finite-time-lyapunov-exponents-ftle A matter of diffusion.. * particles diffuse across a coordinate system at rate r. * if there is underlying order, particles will aggregate into larger-scale structures. * stochastic component (diffusion speed), deterministic component (interface between regimes).

Flow maps and transport dynamics Flow maps: visualizations of how particles diffuse, travel, and aggregate over time. Major structures visible in flow map: vortex rings, streaks, tracks, streams, etc. Characterized indirectly using LCS method (indicator of, influential in natural processes). Flow map examples: Streaks Vortex rings Tracks Streams http://www.vacet.org/ga llery/images_video/jet4 -ftle-0.012_sm.png http://www.rsmas.miami.edu/i nfo/soundings/2008/03/image s/olascoaga-graph.jpg http://www.pa.msu.edu/~stein r/images/vorticity2.jpg

Flow maps and transport dynamics (con t) Coherent structures = distinguished set of fluid particles. Stability properties = impact on fluid mixing. Courtesy: Getty Images In time-periodic laminar flow ~ complex tangles predicted for chaotic advection. * velocity measurements for small # of tracer particles (e.g. dye) can provide a highly resolved velocity field. * particles yield more information about velocity field as time progresses. * high degree of noise, spatial complexity work against this. Re (Reynolds #): Inertial Ro (Rossby #): Length scale Velocity scale Coriolis frequency Re >> 0, turbulent (inertial). Re ~ 0, laminar (viscous). Viscous Ro >> 0, centrifugal, inertial. Ro ~ 0, coriolis.

Flow maps and transport dynamics (con t) Ro = ratio of inertial to Coriolis forces. Re = ratio of inertial to viscous forces. : : FTLE (time): FSLE (size): Actual trajectory unknown, inferred by repeated measurements of λ over interval Courtesy: http://www.cds.caltech.edu/~marsden/ \lcs/fluid-transport

Flow maps and transport dynamics (con t) Experiment: 50x40 cm tank, 0.4 Hz rotation. 2D flow laminar flow regime at top of tank. * Re = 1000, Ro = 0.3 * velocity field = 1500x1500 grid. * Δt =.004 sec. * L t (x o ) along particle paths is large only in small regions of flow. Integrated Lagrangian divergence: Dabiri s Pinch-off (Chaos, 2010) approach: Direct Lyapunov exponent (DLE): Flow map (maps fluid particles from initial position t o over interval T): Linearized divergence of trajectories over interval T FTLE: Deformation tensor

Flow maps and transport dynamics (con t) Features of flow: * long-lived vortices and jets in laminar regime with diversity of sizes. * coherent structures are columnar (vertical across 2D, quasi- 2D and 3D turbulence regimes). Visualizations of LCS: * t >> t 0 = repelling LCS at t 0 are locally maximizing curves (ridges). * t << t 0 = attracting LCS at t 0 are locally maximizing curves (ridges). Results invariant for T values (integration) for 1-16. Integration = ratio of deformed area at t + T: original area around x 0 at t.

Flow maps and transport dynamics (con t) Perform time-integration in two ways, derive two distinct structural modes: Attracting LCS, backwards in time: Maximum values of FSLE = lines that approximate unstable manifolds. Neighboring fluid trajectory attracted, escapes hyperbolic points. Example: fluid funneling down a drain (towards attractor point, away from slopes). Repelling LCS, forwards in time: Maximum values of FSLE form "ridges" that delineate fluids with distinct origins. Neighboring trajectories repelled and form lines that strongly modulate fluid flow, act as transport barriers. Example: advancing front of soap scum or ripple of water. Again, a matter of diffusion: For example: uniform diffusion from a glob of particles (initial condition) can create a repelling LCS that forms a continuous ridge (ring).

Dabiri Paper (Chaos, 2010) Vortex Rings: Wide variety of biological flows, fluid column of length (L) pushed through aperture of distance (D). L/D = formation #. * boundary layers form with movement (piston up and down in column), rolls up into a vortex ring. * vorticity flux fuels growth. L/D > 4, wake formation begins to trail ring. Left: Nature, 424, 621-622 (2003). * disconnection of velocity and vorticity fields = pinch-off. Challenges: previous approaches to understanding pinch off. * pinch-off was not directly observed during vortex ring evolution. * trouble controlling, disentangling interactions in the dynamics. * in low Re flows, vortex ring evolution confused with viscous diffusion.

Dabiri Paper (Chaos, 2010 - con t) Proposed criterion to ID vortex ring pinch-off in axisymmetric jet flow: * starting jet of L/D ~ 12. * termination of previous LCS precedes appearance of new LCS (independent). Marks initiation of pinch-off. * LCS can either form a spiral (persistent vortices) or merge with existing LCS (merger of vortices in shear layer). LCS = finite-time invariant manifolds in time-dependent, aperiodic system: Repelling: diverge forward in time. Attracting: diverge backward in time.

Dabiri Paper (Chaos, 2010 - con t) Streamlines (left) and pathlines (right) for three different times. Streaklines (left) and timelines (right) for three different times.

Dabiri Paper (Chaos, 2010 - con t) LCS are most effective as a quantitative indicator of relative Lagrangian trajectories over time, not pinch-off per se. * regions of high particle seperation between regions of qualitatively different flows. * suited to ID seperation different regions of vorticity-carrying flows that occurs during pinch-off. * robust ID of vortex structure in unsteady flows. * frame-independent FTLE allows for time-history of motion. LCS may be useful to understand optimal vortex formation, implications for a number of biological systems: Courtesy: Sinauer & Associates. * flow of blood/particulates through circulatory system. * swimming and flying motions.

Tew Kai Paper (PNAS, 2009) Problem: ocean eddies (turbulence) influence primary production, community structure, and predation. * predators use boundary between 2 eddies, generates strong dynamic interface. * occurs at multiple temporal and spatial scales, but especially at spatial scales (<10km), not directly observable using satellite altimetry. How do predators locate these structures? * map observations to horizontal velocity field. Apply to FSLE. * compute from marine surface velocity data, mixing activity, and coherent structures. 1) FSLE: measures how fast fluid particles separate to a specified distance. 2) LCS: ID as ridges (location with maximum value of Lyapunov exponent fields). * dispersion rates of tracer particles - integrate trajectories towards the future (forward) or towards past (backward).

Tew Kai Paper (PNAS, 2009- con t) Quantifiers: 1) FSLE f : repel neighboring trajectories (positive value). 2) FSLE b : attract neighboring trajectories (negative value). * dispersion rates of tracer particles allow us to integrate trajectories towards the future (FSLE f ) or towards past (FSLE b ). LCS by FSLE: stretching and contractile properties of transport. * compute time τ it takes 2 tracer particles initially separated by distance δ o to reach final seperation distance δ f in velocity field. FSLE at position x and time t: * depend on choice of 2 length scales = δ o and δ f. Here, δ o =.025 and δ f = 1. * other studies: FSLE adequate to characterize horizontal mixing and transport structures, correlate with tracer fields like temperature and chlorophyll.

Tew Kai Paper (PNAS, 2009- con t) Top: flow maps for geographical region of interest. Show structure of FSLEs. Lower left: histograms for relative frequency of FSLEs. Lower right: migration of birds during observed trips (follow FSLEs). * as predicted by dissipative structures (Prigogine) and constructal theory (Bejan). * higher energy density structures in active flows.

Applications of LCS methodology Drug delivery and diagnosis: * characterize flow patterns in bloodstream as indicative of arterial clogging. * where in circulatory system are laminar, other flow conditions maximized? * jet flows for transport (see Proc. Royal Society B, 272, 1557 1560, 2005). * fastest route to delivery (circulatory system as network physiological TSP)? Morphogenesis: * so-called morphogen gradients may be characterized as ridges from different sources. * especially interesting in tissue engineering contexts (using complex forces to differentiate stem cells). Courtesy: Nature, 413, 797-803 (2001).

Applications of LCS methodology (con t) Analysis of Biological Complexity: LCS ridges as basis for rugged energy, fitness landscapes. Soft Active Material Dynamics: How do we describe higher-level order in complex monolayers, bilayers? * nematic (left) to smectic (right) transitions in self-assembly. * complex patterns in materials, surfaces. Reaction-Diffusion Computing: * Stu Kauffman s NK-boolean networks, ruggedness = complexity. * characterizing patterns of local (and even global) minima/maxima may allow for higher-order interpretation. * computers that use B-Z reactions, other physical models. * how do we characterize outputs (binary and nonbinary)?