Michael Shelley

Flexible filaments in a flowing soap film as a model for one-dimensional flags in a two-dimensional wind.

The dynamics of swimming fish and flapping flags involves a complicated interaction of their deformable shapes with the surrounding fluid flow. Even in the passive case of a flag, the flag exerts forces on the fluid through its own inertia and elastic responses, and is likewise acted on by hydrodynamic pressure and drag. But such couplings are not well understood. Here we study these interactions experimentally, using an analogous system of flexible filaments in flowing soap films. We find that, for a single filament (or ‘flag’) held at its upstream end and otherwise unconstrained, there are two distinct, stable dynamical states. The first is a stretched-straight state: the filament is immobile and aligned in the flow direction. The existence of this state seems to refute the common belief that a flag is always unstable and will flap. The second is a flapping state: the filament executes a sinuous motion in a manner akin to the flapping of a flag in the wind. We study further the hydrodynamically coupled interaction between two such filaments, and demonstrate the existence of four different dynamical states.

Drag reduction through self-similar bending of a flexible body

The classical theory of high-speed flow predicts that a moving rigid object experiences a drag proportional to the square of its speed. However, this reasoning does not apply if the object in the flow is flexible, because its shape then becomes a function of its speed—for example, the rolling up of broad tree leaves in a stiff wind. The reconfiguration of bodies by fluid forces is common in nature, and can result in a substantial drag reduction that is beneficial for many organisms. Experimental studies of such flow–structure interactions generally lack a theoretical interpretation that unifies the body and flow mechanics. Here we use a flexible fibre immersed in a flowing soap film to measure the drag reduction that arises from bending of the fibre by the flow. Using a model that couples hydrodynamics to bending, we predict a reduced drag growth compared to the classical theory. The fibre undergoes a bending transition, producing shapes that are self-similar; for such configurations, the drag scales with the length of self-similarity, rather than the fibre profile width. These predictions are supported by our experimental data.

The mechanics of slithering locomotion

In this experimental and theoretical study, we investigate the slithering of snakes on flat surfaces. Previous studies of slithering have rested on the assumption that snakes slither by pushing laterally against rocks and branches. In this study, we develop a theoretical model for slithering locomotion by observing snake motion kinematics and experimentally measuring the friction coefficients of snakeskin. Our predictions of body speed show good agreement with observations, demonstrating that snake propulsion on flat ground, and possibly in general, relies critically on the frictional anisotropy of their scales. We have also highlighted the importance of weight distribution in lateral undulation, previously difficult to visualize and hence assumed uniform. The ability to redistribute weight, clearly of importance when appendages are airborne in limbed locomotion, has a much broader generality, as shown by its role in improving limbless locomotion.

Instabilities, pattern formation, and mixing in active suspensions

Suspensions of self-propelled particles, such as swimming micro-organisms, are known to undergo complex dynamics as a result of hydrodynamic interactions. To elucidate these dynamics, a kinetic theory is developed and applied to study the linear stability and the nonlinear pattern formation in these systems. The evolution of a suspension of self-propelled particles is modeled using a conservation equation for the particle configurations, coupled to a mean-field description of the flow arising from the stress exerted by the particles on the fluid. Based on this model, we first investigate the stability of both aligned and isotropic suspensions. In aligned suspensions, an instability is shown to always occur at finite wavelengths, a result that extends previous predictions by Simha and Ramaswamy [“Hydrodynamic fluctuations and instabilities in ordered suspensions of self-propelled particles,” Phys. Rev. Lett. 89, 058101 (2002)]. In isotropic suspensions, we demonstrate the existence of an instability for th...

An egalitarian network model for the emergence of simple and complex cells in visual cortex.

We explain how simple and complex cells arise in a large-scale neuronal network model of the primary visual cortex of the macaque. Our model consists of ≈4,000 integrate-and-fire, conductance-based point neurons, representing the cells in a small, 1-mm2 patch of an input layer of the primary visual cortex. In the model the local connections are isotropic and nonspecific, and convergent input from the lateral geniculate nucleus confers cortical cells with orientation and spatial phase preference. The balance between lateral connections and lateral geniculate nucleus drive determines whether individual neurons in this recurrent circuit are simple or complex. The model reproduces qualitatively the experimentally observed distributions of both extracellular and intracellular measures of simple and complex response.

The long-time motion of vortex sheets with surface tension

We study numerically the simplest model of two incompressible, immiscible fluids shearing past one another. The fluids are two-dimensional, inviscid, irrotational, density matched, and separated by a sharp interface under a surface tension. The nonlinear growth and evolution of this interface is governed by only the competing effects of the Kelvin–Helmholtz instability and the dispersion due to surface tension. We have developed new and highly accurate numerical methods designed to treat the difficulties associated with the presence of surface tension. This allows us to accurately simulate the evolution of the interface over much longer times than has been done previously. A surprisingly rich variety of behavior is found. For small Weber numbers, where there are no unstable length-scales, the flow is dispersively dominated and oscillatory behavior is observed. For intermediate Weber numbers, where there are only a few unstable length-scales, the interface forms elongating and interpenetrating fingers of fluid. At larger Weber numbers, where there are many unstable scales, the interface rolls-up into a “Kelvin-Helmholtz” spiral with its late evolution terminated by the collision of the interface with itself, forming at that instant bubbles of fluid at the core of the spiral. Using locally refined grids, this singular event (a “topological” or “pinching” singularity) is studied carefully. Our computations suggest at least a partial conformance to a local self-similar scaling. For fixed initial data, the pinching singularity times decrease as the surface tension is reduced, apparently towards the singularity time associated with the zero surface tension problem, as studied by Moore and others. Simulations from more complicated, multi-modal initial data show the evolution as a combination of these fingers, spirals, and pinches.

Mexican hats and pinwheels in visual cortex

Many models of cortical function assume that local lateral connections are specific with respect to the preferred features of the interacting cells and that they are organized in a Mexican-hat pattern with strong “center” excitation flanked by strong “surround” inhibition. However, anatomical data on primary visual cortex indicate that the local connections are isotropic and that inhibition has a shorter range than excitation. We address this issue in an analytical study of a neuronal network model of the local cortical circuit in primary visual cortex. In the model, the orientation columns specified by the convergent lateral geniculate nucleus inputs are arranged in a pinwheel architecture, whereas cortical connections are isotropic. We obtain a trade-off between the spatial range of inhibition and its time constant. If inhibition is fast, the network can operate in a Mexican-hat pattern with isotropic connections even with a spatially narrow inhibition. If inhibition is not fast, Mexican-hat operation requires a spatially broad inhibition. The Mexican-hat operation can generate a sharp orientation tuning, which is largely independent of the distance of the cell from the pinwheel center.

How flexibility induces streamlining in a two-dimensional flow

Recent work in bio-fluid dynamics has studied the relation of fluid drag to flow speed for flexible organic structures, such as tree leaves, seaweed, and coral beds, and found a reduction in drag growth due to body reconfiguration with increasing flow speed. Our theoretical and experimental work isolates the role of elastic bending in this process. Using a flexible glass fiber wetted into a vertical soap-film flow, we identify a transition in flow speed beyond which fluid forces dominate the elastic response, and yield large deformations of the fiber that greatly reduce drag. We construct free-streamline models that couple fluid and elastic forces and solve them in an efficient numerical scheme. Shape self-similarity emerges, with a scaling set by the balance of forces in a small “tip region” about the flow’s stagnation point. The result is a transition from the classical U2 drag scaling of rigid bodies to a new U4/3 drag law. We derive an asymptotic expansion for the fiber shape and flow, based on the le...

States of high conductance in a large-scale model of the visual cortex

This paper reports on the consequences of large, activity dependent, synaptic conductances for neurons in a large-scale neuronal network model of the input layer 4Cα of the Macaque primary visual cortex (Area V1). This high conductance state accounts for experimental observations about orientation selectivity, dynamics, and response magnitude (D. McLaughlin et al. (2000) Proc. Natl. Acad. Sci. USA 97: 8087–8092), and the linear dependence of Simple cells on visual stimuli (J. Wielaard et al. (2001) J. Neuroscience 21: 5203–5211). The source of large conductances in the model can be traced to inhibitory corticocortical synapses, and the models predictions of large conductance changes are consistent with recent intracellular measurements (L. Borg-Graham et al. (1998) Nature 393: 369–373; J. Hirsch et al. (1998) J. Neuroscience 15: 9517–9528; J.S. Anderson et al. (2000) J. Neurophysiol. 84: 909–926). During visual stimulation, these conductances are large enough that their associated time-scales become the shortest in the model cortex, even below that of synaptic interactions. One consequence of this activity driven separation of time-scales is that a neuron responds very quickly to temporal changes in its synaptic drive, with its intracellular membrane potential tracking closely an effective reversal potential composed of the instantaneous synaptic inputs. From the effective potential and large synaptic conductance, the spiking activity of a cell can be expressed in an interesting and simplified manner, with the result suggesting how accurate and smoothly graded responses are achieved in the model network. Further, since neurons in this high-conductance state respond quickly, they are also good candidates as coincidence detectors and burst transmitters.

Surprising behaviors in flapping locomotion with passive pitching

To better understand the role of wing and fin flexibility in flapping locomotion, we study through experiment and numerical simulation a freely moving wing that can “pitch” passively as it is actively heaved in a fluid. We observe a range of flapping frequencies corresponding to large horizontal velocities, a regime of underperformance relative to a clamped (nonpitching) flapping wing, and a surprising, hysteretic regime in which the flapping wing can move horizontally in either direction (despite left/right symmetry being broken by the specific mode of pitching). The horizontal velocity is shown to peak when the flapping frequency is near the immersed system’s resonant frequency. Unlike for the clamped wing, we find that locomotion is achieved by vertically flapped symmetric wings with even the slightest pitching flexibility, and the system exhibits a continuous departure from the Stokesian regime. The phase difference between the vertical heaving motion and consequent pitching changes continuously with ...