Stiffness of the human foot and evolution of the transverse arch.

The stiff human foot enables an efficient push-off when walking or running, and was critical for the evolution of bipedalism1-6. The uniquely arched morphology of the human midfoot is thought to stiffen it5-9, whereas other primates have flat feet that bend severely in the midfoot7,10,11. However, the relationship between midfoot geometry and stiffness remains debated in foot biomechanics12,13, podiatry14,15 and palaeontology4-6. These debates centre on the medial longitudinal arch5,6 and have not considered whether stiffness is affected by the second, transverse tarsal arch of the human foot16. Here we show that the transverse tarsal arch, acting through the inter-metatarsal tissues, is responsible for more than 40% of the longitudinal stiffness of the foot. The underlying principle resembles a floppy currency note that stiffens considerably when it curls transversally. We derive a dimensionless curvature parameter that governs the stiffness contribution of the transverse tarsal arch, demonstrate its predictive power using mechanical models of the foot and find its skeletal correlate in hominin feet. In the foot, the material properties of the inter-metatarsal tissues and the mobility of the metatarsals may additionally influence the longitudinal stiffness of the foot and thus the curvature-stiffness relationship of the transverse tarsal arch. By analysing fossils, we track the evolution of the curvature parameter among extinct hominins and show that a human-like transverse arch was a key step in the evolution of human bipedalism that predates the genus Homo by at least 1.5 million years. This renewed understanding of the foot may improve the clinical treatment of flatfoot disorders, the design of robotic feet and the study of foot function in locomotion.

Dynamics and stability of running on rough terrains.

Stability of running on rough terrain depends on the propagation of perturbations due to the ground. We consider stability within the sagittal plane and model the dynamics of running as a two-dimensional body with alternating aerial and stance phases. Stance is modelled as a passive, impulsive collision followed by an active, impulsive push-off that compensates for collisional losses. Such a runner has infinitely many strategies to maintain periodic gaits on flat ground. However, these strategies differ in how perturbations due to terrain unevenness are propagated. Instabilities manifest as tumbling (orientational instability) or failing to maintain a steady speed (translational instability). We find that open-loop strategies that avoid sensory feedback are sufficient to maintain stability on step-like terrains with piecewise flat surfaces that randomly vary in height. However, these open-loop runners lose orientational stability on rough terrains whose slope also varies randomly. The orientational instability is significantly mitigated by minimizing the tangential collision, which typically requires sensory information and anticipatory strategies such as leg retraction. By analysing the propagation of perturbations, we derive a single dimensionless parameter that governs stability. This parameter provides guidelines for the design and control of both biological and robotic runners.

Controllable biomimetic birdsong.

Birdsong is the product of the controlled generation of sound embodied in a neuromotor system. From a biophysical perspective, a natural question is that of the difficulty of producing birdsong. To address this, we built a biomimetic syrinx consisting of a stretched simple rubber tube through which air is blown, subject to localized mechanical squeezing with a linear actuator. A large static tension on the tube and small dynamic variations in the localized squeezing allow us to control transitions between three states: a quiescent state, a periodic state and a solitary wave state. The static load brings the system close to threshold for spontaneous oscillations, while small dynamic loads allow for rapid transitions between the states. We use this to mimic a variety of birdsongs via the slow-fast modulated nonlinear dynamics of the physical substrate, the syrinx, regulated by a simple controller. Finally, a minimal mathematical model of the system inspired by our observations allows us to address the problem of song mimicry in an excitable oscillator for tonal songs.

Curvature-induced stiffening of a fish fin.

How fish modulate their fin stiffness during locomotive manoeuvres remains unknown. We show that changing the fin's curvature modulates its stiffness. Modelling the fin as bendable bony rays held together by a membrane, we deduce that fin curvature is manifested as a misalignment of the principal bending axes between neighbouring rays. An external force causes neighbouring rays to bend and splay apart, and thus stretches the membrane. This coupling between bending the rays and stretching the membrane underlies the increase in stiffness. Using three-dimensional reconstruction of a mackerel (Scomber japonicus) pectoral fin for illustration, we calculate the range of stiffnesses this fin is expected to span by changing curvature. The three-dimensional reconstruction shows that, even in its geometrically flat state, a functional curvature is embedded within the fin microstructure owing to the morphology of individual rays. As the ability of a propulsive surface to transmit force to the surrounding fluid is limited by its stiffness, the fin curvature controls the coupling between the fish and its surrounding fluid. Thereby, our results provide mechanical underpinnings and morphological predictions for the hypothesis that the spanned range of fin stiffnesses correlates with the behaviour and the ecological niche of the fish.

Skating on a film of air: drops impacting on a surface.

The commonly accepted description of drops impacting on a surface typically ignores the essential role of the air that is trapped between the impacting drop and the surface. Here we describe a new imaging modality that is sensitive to the behavior right at the surface. We show that a very thin film of air, only a few tens of nanometers thick, remains trapped between the falling drop and the surface as the drop spreads. The thin film of air serves to lubricate the drop enabling the fluid to skate on the air film laterally outward at surprisingly high velocities, consistent with theoretical predictions. Eventually this thin film of air breaks down as the fluid wets the surface via a spinodal-like mechanism. Our results show that the dynamics of impacting drops are much more complex than previously thought, with a rich array of unexpected phenomena that require rethinking classic paradigms.

Algorithm for a microfluidic assembly line.

Microfluidic technology has revolutionized the control of flows at small scales giving rise to new possibilities for assembling complex structures on the microscale. We analyze different possible algorithms for assembling arbitrary structures, and demonstrate that a sequential assembly algorithm can manufacture arbitrary 3D structures from identical constituents. We illustrate the algorithm by showing that a modified Hele-Shaw cell with 7 controlled flow rates can be designed to construct the entire English alphabet from particles that irreversibly stick to each other.

Precursors to splashing of liquid droplets on a solid surface.

A high velocity impact between a liquid droplet and a solid surface produces a splash. Classical work traced the origin of the splash to a thin sheet of fluid ejected near the impact point. Mechanisms of sheet formation have heretofore relied on initial contact of the droplet and the surface. We demonstrate that, neglecting intermolecular forces between the liquid and the solid, the liquid does not contact the solid, and instead spreads on a very thin air film. The interface of the droplet develops a high curvature and emits capillary waves.

Short-time dynamics of partial wetting.

When a liquid drop contacts a wettable surface, the liquid spreads over the solid to minimize the total surface energy. The first moments of spreading tend to be rapid. For example, a millimeter-sized water droplet will wet an area having the same diameter as the drop within a millisecond. For perfectly wetting systems, this spreading is inertially dominated. Here we identify that even in the presence of a contact line, the initial wetting is dominated by inertia rather than viscosity. We find that the spreading radius follows a power-law scaling in time where the exponent depends on the equilibrium contact angle. We propose a model, consistent with the experimental results, in which the surface spreading is regulated by the generation of capillary waves.

Mechanisms for liquid slip at solid surfaces.

One of the oldest unresolved problems in fluid mechanics is the nature of liquid flow along solid surfaces. It is traditionally assumed that across the liquid-solid interface, liquid and solid speeds exactly match. However, recent observations document that on the molecular scale, liquids can slip relative to solids. We formulate a model in which the liquid dynamics are described by a stochastic differential-difference equation, related to the Frenkel-Kontorova equation. The model, in agreement with molecular dynamics simulations, reveals that slip occurs via two mechanisms: localized defect propagation and concurrent slip of large domains. Well-defined transitions occur between the two mechanisms.

A simple model illustrating the role of turbulence on phytoplankton blooms.

The problem of the vertical distribution of phytoplankton is considered in the presence of gravitational settling, turbulent mixing, population growth due to cell division and a constant rate of loss due to predation and natural death. Nutrients are assumed to be plentiful so that the production rate depends only on the light available for photosynthesis. The non-linear saturation of plankton growth is modeled by allowing the attenuation rate of light to be a linear function of the plankton density. The turbulent diffusivity is assumed constant which corresponds to a mixed layer depth very much greater than the depth of light penetration (euphotic depth). It is shown that an exact analytical solution of this non-linear problem is possible for an idealized model in which the functional dependence of production on light intensity is assumed to be a step function. Non-zero solutions are shown to exist only if the parameters characterizing the system are above a certain critical curve in a two dimensional parameter space. Numerical simulations using functional forms of the production curve that resemble the measured photosynthetic response of plankton, show, that the qualitative behavior of the system is similar to that of the idealized model presented. Comparisons are made with other analytical approaches to the problem.