抗原变异可使得多种致病微生物易于逃避宿主免疫应答。表达在感染红细胞表面的恶性疟原虫红细胞表面蛋白1（PfPMP1）与感染红细胞、内皮细胞、树突状细胞以及胎盘的单个或多个受体作用，在黏附及免疫逃避中起关键的作用。每个单倍体基因组var基因家族编码约60种成员，通过启动转录不同的var基因变异体为抗原变异提供了分子基础。

疟原虫var基因转换速率变化导致抗原变异[英]／Paul H, Robert P, Christodoulou Z, et al//Proc Natl Acad Sci U S A

Differently from passive Brownian particles, active particles, also known as self-propelled Brownian particles or microswimmers and nanoswimmers, are capable of taking up energy from their environment and converting it into directed motion. Because of this constant flow of energy, their behavior can be explained and understood only within the framework of nonequilibrium physics. In the biological realm, many cells perform directed motion, for example, as a way to browse for nutrients or to avoid toxins. Inspired by these motile microorganisms, researchers have been developing artificial particles that feature similar swimming behaviors based on different mechanisms. These man-made micromachines and nanomachines hold a great potential as autonomous agents for health care, sustainability, and security applications. With a focus on the basic physical features of the interactions of self-propelled Brownian particles with a crowded and complex environment, this comprehensive review will provide a guided tour through its basic principles, the development of artificial self-propelling microparticles and nanoparticles, and their application to the study of nonequilibrium phenomena, as well as the open challenges that the field is currently facing.

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Active Particles in Complex and Crowded Environments

Self-propelled particles include both self-phoretic synthetic colloids and various microorganisms. By continually consuming energy, they bypass the laws of equilibrium thermodynamics. These laws enforce the Boltzmann distribution in thermal equilibrium: The steady state is then independent of kinetic parameters. In contrast, self-propelled particles tend to accumulate where they move more slowly. They may also slow down at high density for either biochemical or steric reasons. This creates positive feedback, which can lead to motility-induced phase separation (MIPS) between dense and dilute fluid phases. At leading order in gradients, a mapping relates variable-speed, self-propelled particles to passive particles with attractions. This deep link to equilibrium phase separation is confirmed by simulations but generally breaks down at higher order in gradients: New effects, with no equilibrium counterpart, then emerge. We give a selective overview of the fast-developing field of MIPS, focusing on theory and...

Motility-Induced Phase Separation

Locomotion and transport of microorganisms in fluids is an essential aspect of life. Search for food, orientation toward light, spreading of off-spring, and the formation of colonies are only possible due to locomotion. Swimming at the microscale occurs at low Reynolds numbers, where fluid friction and viscosity dominates over inertia. Here, evolution achieved propulsion mechanisms, which overcome and even exploit drag. Prominent propulsion mechanisms are rotating helical flagella, exploited by many bacteria, and snake-like or whip-like motion of eukaryotic flagella, utilized by sperm and algae. For artificial microswimmers, alternative concepts to convert chemical energy or heat into directed motion can be employed, which are potentially more efficient. The dynamics of microswimmers comprises many facets, which are all required to achieve locomotion. In this article, we review the physics of locomotion of biological and synthetic microswimmers, and the collective behavior of their assemblies. Starting from individual microswimmers, we describe the various propulsion mechanism of biological and synthetic systems and address the hydrodynamic aspects of swimming. This comprises synchronization and the concerted beating of flagella and cilia. In addition, the swimming behavior next to surfaces is examined. Finally, collective and cooperate phenomena of various types of isotropic and anisotropic swimmers with and without hydrodynamic interactions are discussed.

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Physics of microswimmers--single particle motion and collective behavior: a review.

/pdf/physics-of-microswimmers-single-particle-motion-and-2tjtc65h8s.pdf

Physics of Microswimmers - Single Particle Motion and Collective Behavior

We examine a minimal model for an active colloidal fluid in the form of self-propelled Brownian spheres that interact purely through excluded volume with no aligning interaction. Using simulations and analytic modeling, we quantify the phase diagram and separation kinetics. We show that this nonequilibrium active system undergoes an analog of an equilibrium continuous phase transition, with a binodal curve beneath which the system separates into dense and dilute phases whose concentrations depend only on activity. The dense phase is a unique material that we call an active solid, which exhibits the structural signatures of a crystalline solid near the crystal-hexatic transition point, and anomalous dynamics including superdiffusive motion on intermediate time scales.

/pdf/structure-and-dynamics-of-a-phase-separating-active-3ji2j6kwha.pdf

Structure and Dynamics of a Phase-Separating Active Colloidal Fluid

The study of liquid crystals at equilibrium has led to fundamental insights into the nature of ordered materials, as well as to practical applications such as display technologies. Active nematics are a fundamentally different class of liquid crystals, driven away from equilibrium by the autonomous motion of their constituent rod-like particles. This internally generated activity powers the continuous creation and annihilation of topological defects, which leads to complex streaming flows whose chaotic dynamics seem to destroy long-range order. Here, we study these dynamics in experimental and computational realizations of active nematics. By tracking thousands of defects over centimetre-scale distances in microtubule-based active nematics, we identify a non-equilibrium phase characterized by a system-spanning orientational order of defects. This emergent order persists over hours despite defect lifetimes of only seconds. Similar dynamical structures are observed in coarse-grained simulations, suggesting that defect-ordered phases are a generic feature of active nematics.

/pdf/orientational-order-of-motile-defects-in-active-nematics-538k8ga7u2.pdf

Orientational order of motile defects in active nematics

http://absimage.aps.org/image/MAR13/MWS_MAR13-2012-003161.pdf

Motivated by recent experiments, we study a system of self-propelled colloids that experience short-range attractive interactions and are confined to a surface. Using simulations we find that the phase behavior for such a system is reentrant as a function of activity: phase-separated states exist in both the low- and high-activity regimes, with a homogeneous active fluid in between. To understand the physical origins of reentrance, we develop a kinetic model for the system's steady-state dynamics whose solution captures the main features of the phase behavior. We also describe the varied kinetics of phase separation, which range from the familiar nucleation and growth of clusters to the complex coarsening of active particle gels.

Reentrant phase behavior in active colloids with attraction.

The study of liquid crystals at equilibrium has led to fundamental insights into the nature of ordered materials, as well as to practical applications such as display technologies. Active nematics are a fundamentally different class of liquid crystals, driven away from equilibrium by the autonomous motion of their constituent rod-like particles. This internally generated activity powers the continuous creation and annihilation of topological defects, which leads to complex streaming flows whose chaotic dynamics appear to destroy long-range order. Here, we study these dynamics in experimental and computational realizations of active nematics. By tracking thousands of defects over centimetre-scale distances in microtubule-based active nematics, we identify a non-equilibrium phase characterized by system-spanning orientational order of defects. This emergent order persists over hours despite defect lifetimes of only seconds. Similar dynamical structures are observed in coarse-grained simulations, suggesting that defect-ordered phases are a generic feature of active nematics.

Gabriel Redner

Papers

Structure and Dynamics of a Phase-Separating Active Colloidal Fluid

Orientational order of motile defects in active nematics

Structure and Dynamics of a Phase-Separating Active Colloidal Fluid

Reentrant phase behavior in active colloids with attraction.

Orientational order of motile defects in active nematics