This review summarizes theoretical progress in the field of active matter, placing it in the context of recent experiments. This approach offers a unified framework for the mechanical and statistical properties of living matter: biofilaments and molecular motors in vitro or in vivo, collections of motile microorganisms, animal flocks, and chemical or mechanical imitations. A major goal of this review is to integrate several approaches proposed in the literature, from semimicroscopic to phenomenological. In particular, first considered are ``dry'' systems, defined as those where momentum is not conserved due to friction with a substrate or an embedding porous medium. The differences and similarities between two types of orientationally ordered states, the nematic and the polar, are clarified. Next, the active hydrodynamics of suspensions or ``wet'' systems is discussed and the relation with and difference from the dry case, as well as various large-scale instabilities of these nonequilibrium states of matter, are highlighted. Further highlighted are various large-scale instabilities of these nonequilibrium states of matter. Various semimicroscopic derivations of the continuum theory are discussed and connected, highlighting the unifying and generic nature of the continuum model. Throughout the review, the experimental relevance of these theories for describing bacterial swarms and suspensions, the cytoskeleton of living cells, and vibrated granular material is discussed. Promising extensions toward greater realism in specific contexts from cell biology to animal behavior are suggested, and remarks are given on some exotic active-matter analogs. Last, the outlook for a quantitative understanding of active matter, through the interplay of detailed theory with controlled experiments on simplified systems, with living or artificial constituents, is summarized.

Hydrodynamics of soft active matter

Inspired by the insect-eating Nepenthes pitcher plant, which snares its prey on a surface lubricated by a remarkably slippery aqueous secretion, Joanna Aizenberg and colleagues have synthesized omniphobic surfaces that can self-repair and function at high pressures. Their 'slippery liquid-infused porous surfaces' (or SLIPS) exhibit almost perfect slipperiness towards polar, organic and complex liquids. SLIPS function under extreme conditions, are easily constructed from inexpensive materials and can be endowed with other useful characteristics, such as enhanced optical transparency, through the selection of appropriate substrates and lubricants. Ultra-slippery surfaces of this type might find application in biomedical fluid handling, fuel transport, antifouling, anti-icing, optical imaging and elsewhere. Creating a robust synthetic surface that repels various liquids would have broad technological implications for areas ranging from biomedical devices and fuel transport to architecture but has proved extremely challenging1. Inspirations from natural nonwetting structures2,3,4,5,6, particularly the leaves of the lotus, have led to the development of liquid-repellent microtextured surfaces that rely on the formation of a stable air–liquid interface7,8,9. Despite over a decade of intense research, these surfaces are, however, still plagued with problems that restrict their practical applications: limited oleophobicity with high contact angle hysteresis9, failure under pressure10,11,12 and upon physical damage1,7,11, inability to self-heal and high production cost1,11. To address these challenges, here we report a strategy to create self-healing, slippery liquid-infused porous surface(s) (SLIPS) with exceptional liquid- and ice-repellency, pressure stability and enhanced optical transparency. Our approach—inspired by Nepenthes pitcher plants13—is conceptually different from the lotus effect, because we use nano/microstructured substrates to lock in place the infused lubricating fluid. We define the requirements for which the lubricant forms a stable, defect-free and inert ‘slippery’ interface. This surface outperforms its natural counterparts2,3,4,5,6 and state-of-the-art synthetic liquid-repellent surfaces8,9,14,15,16 in its capability to repel various simple and complex liquids (water, hydrocarbons, crude oil and blood), maintain low contact angle hysteresis (<2.5°), quickly restore liquid-repellency after physical damage (within 0.1–1 s), resist ice adhesion, and function at high pressures (up to about 680 atm). We show that these properties are insensitive to the precise geometry of the underlying substrate, making our approach applicable to various inexpensive, low-surface-energy structured materials (such as porous Teflon membrane). We envision that these slippery surfaces will be useful in fluid handling and transportation, optical sensing, medicine, and as self-cleaning and anti-fouling materials operating in extreme environments.

Bioinspired self-repairing slippery surfaces with pressure-stable omniphobicity

Beginning with Emlen (1966) and MacArthur and Pianka (1966) and extending through the last ten years, several authors have sought to predict the foraging behavior of animals by means of mathematical models. These models are very similar,in that they all assume that the fitness of a foraging animal is a function of the efficiency of foraging measured in terms of some "currency" (Schoener, 1971) -usually energy- and that natural selection has resulted in animals that forage so as to maximize this fitness. As a result of these similarities, the models have become known as "optimal foraging models"; and the theory that embodies them, "optimal foraging theory." The situations to which optimal foraging theory has been applied, with the exception of a few recent studies, can be divided into the following four categories: (1) choice by an animal of which food types to eat (i.e., optimal diet); (2) choice of which patch type to feed in (i.e., optimal patch choice); (3) optimal allocation of time to different patches; and (4) optimal patterns and speed of movements. In this review we discuss each of these categories separately, dealing with both the theoretical developments and the data that permit tests of the predictions. The review is selective in the sense that we emphasize studies that either develop testable predictions or that attempt to test predictions in a precise quantitative manner. We also discuss what we see to be some of the future developments in the area of optimal foraging theory and how this theory can be related to other areas of biology. Our general conclusion is that the simple models so far formulated are supported are supported reasonably well by available data and that we are optimistic about the value both now and in the future of optimal foraging theory. We argue, however, that these simple models will requre much modification, espicially to deal with situations that either cannot easily be put into one or another of the above four categories or entail currencies more complicated that just energy.

Citation classic - optimal foraging - a selective review of theory and tests

The boundary layer equations for plane, incompressible, and steady flow are 

$$\matrix{ {u{{\partial u} \over {\partial x}} + v{{\partial u} \over {\partial y}} = - {1 \over \varrho }{{\partial p} \over {\partial x}} + v{{{\partial ^2}u} \over {\partial {y^2}}},} \cr {0 = {{\partial p} \over {\partial y}},} \cr {{{\partial u} \over {\partial x}} + {{\partial v} \over {\partial y}} = 0.} \cr }$$

Boundary Layer Theory

The widespread use of controlled molecular-level motion in key natural processes suggests that great rewards could come from bridging the gap between the present generation of synthetic molecular systems, which by and large rely upon electronic and chemical effects to carry out their functions, and the machines of the macroscopic world, which utilize the synchronized movements of smaller parts to perform specific tasks. This is a scientific area of great contemporary interest and extraordinary recent growth, yet the notion of molecular-level machines dates back to a time when the ideas surrounding the statistical nature of matter and the laws of thermodynamics were first being formulated. Here we outline the exciting successes in taming molecular-level movement thus far, the underlying principles that all experimental designs must follow, and the early progress made towards utilizing synthetic molecular structures to perform tasks using mechanical motion. We also highlight some of the issues and challenges that still need to be overcome.

Synthetic molecular motors and mechanical machines.

Microorganisms such as bacteria and many eukaryotic cells propel themselves with hair-like structures known as flagella, which can exhibit a variety of structures and movement patterns. For example, bacterial flagella are helically shaped and driven at their bases by a reversible rotary engine, which rotates the attached flagellum to give a motion similar to that of a corkscrew. In contrast, eukaryotic cells use flagella that resemble elastic rods and exhibit a beating motion: internally generated stresses give rise to a series of bends that propagate towards the tip. In contrast to this variety of swimming strategies encountered in nature, a controlled swimming motion of artificial micrometre-sized structures has not yet been realized. Here we show that a linear chain of colloidal magnetic particles linked by DNA and attached to a red blood cell can act as a flexible artificial flagellum. The filament aligns with an external uniform magnetic field and is readily actuated by oscillating a transverse field. We find that the actuation induces a beating pattern that propels the structure, and that the external fields can be adjusted to control the velocity and the direction of motion.

Microscopic artificial swimmers

Micropatterned surfaces have been studied extensively as model systems to understand influences of topographic or chemical heterogeneities on wetting phenomena. Such surfaces yield specific wetting or hydrodynamic effects, for example, ultrahydrophobic surfaces, 'fakir' droplets, tunable electrowetting, slip in the presence of surface heterogeneities and so on. In addition, chemical patterns allow control of the locus, size and shape of droplets by pinning the contact lines at predetermined locations. Applications include the design of 'self-cleaning' surfaces and hydrophilic spots to automate the deposition of probes on DNA chips. Here, we discuss wetting on topographically patterned but chemically homogeneous surfaces and demonstrate mechanisms of shape selection during imbibition of the texture. We obtain different deterministic final shapes of the spreading droplets, including octagons, squares, hexagons and circles. The shape selection depends on the topographic features and the liquid through its equilibrium contact angle. Considerations of the dynamics provide a 'shape' diagram that summarizes our observations and suggest rules for a designer's tool box.

/pdf/imbibition-by-polygonal-spreading-on-microdecorated-surfaces-4sy2o5bjwm.pdf

Imbibition by polygonal spreading on microdecorated surfaces

The bacterium Bacillus subtilis produces the molecule surfactin, which is known to enhance the spreading of multicellular colonies on nutrient substrates by lowering the surface tension of the surrounding fluid, and to aid in the formation of aerial structures. Here we present experiments and a mathematical model that demonstrate how the differential accumulation rates induced by the geometry of the bacterial film give rise to surfactant waves. The spreading flux increases with increasing biofilm viscosity. Community associations are known to protect bacterial populations from environmental challenges such as predation, heat, or chemical stresses, and enable digestion of a broader range of nutritive sources. This study provides evidence of enhanced dispersal through cooperative motility, and points to nonintuitive methods for controlling the spread of biofilms.

https://www.pnas.org/content/pnas/106/43/18109.full.pdf

Bacillus subtilis spreads by surfing on waves of surfactant

Following a novel realization of low-Reynolds-number swimming (Dreyfus et al., Nature, vol. 436, 2005, p. 862), in which self-assembled filaments of paramagnetic micron-sized beads are tethered to red blood cells and then induced to swim under crossed uniform and oscillating magnetic fields, the dynamics of magnetoelastic filaments is studied. The filament is modelled as a slender elastica driven by a magnetic body torque. The model is applied to experiments of Goubault et al. (Phys. Rev. Lett., vol. 91, 2003, art. 260802) to predict the lifetimes of metastable static filament conformations that are known to form under uniform fields. A second experimental swimming scenario, complementary to that of Dreyfus et al. (2005), is described: filaments are capable of swimming even if not tethered to red blood cells. Yet, if both ends of the filament are left free and the material and magnetic parameters are uniform along its length then application of an oscillating transverse field can only generate homogeneous torques, and net translation is prohibited by symmetry. It is shown that fore–aft symmetry is broken when variation of the bending stiffness along the filament is accounted for by including elastic defects, which produces results consistent with the swimming phenomenology.

On the dynamics of magnetically driven elastic filaments

Inertial lift forces are exploited within inertial microfluidic devices to position, segregate and sort particles or droplets. However, the forces and their focusing positions can currently only be predicted by numerical simulations, making rational device design very difficult. Here we develop theory for the forces on particles in microchannel geometries. We use numerical experiments to dissect the dominant balances within the Navier–Stokes equations and derive an asymptotic model to predict the lateral force on the particle as a function of particle size. Our asymptotic model is valid for a wide array of particle sizes and Reynolds numbers, and allows us to predict how focusing position depends on particle size.

Marcus Roper

Papers

Microscopic artificial swimmers

Imbibition by polygonal spreading on microdecorated surfaces

Bacillus subtilis spreads by surfing on waves of surfactant

On the dynamics of magnetically driven elastic filaments

Inertial migration of a rigid sphere in three-dimensional Poiseuille flow