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.

/pdf/active-particles-in-complex-and-crowded-environments-4pyirdi1aj.pdf

Active Particles in Complex and Crowded Environments

Active Brownian particles, also referred to as microswimmers and nanoswimmers, are biological or manmade microscopic and nanoscopic particles that can self-propel. Because of their activity, their behavior can only be explained and understood within the framework of nonequilibrium physics. In the biological realm, many cells perform active Brownian motion, for example, when moving away from toxins or towards nutrients. Inspired by these motile microorganisms, researchers have been developing artificial active particles that feature similar swimming behaviors based on different mechanisms; these manmade micro- and nanomachines hold a great potential as autonomous agents for healthcare, sustainability, and security applications. With a focus on the basic physical features of the interactions of active Brownian particles with a crowded and complex environment, this comprehensive review will put the reader at the very forefront of the field of active Brownian motion, providing a guided tour through its basic principles, the development of artificial self-propelling micro- and nanoparticles, and their application to the study of nonequilibrium phenomena, as well as the open challenges that the field is currently facing.

/pdf/active-brownian-particles-in-complex-and-crowded-3ym4we2m8l.pdf

Active Brownian 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.

/pdf/physics-of-microswimmers-single-particle-motion-and-4vxoi085wf.pdf

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

We study experimentally and numerically a (quasi-)two-dimensional colloidal suspension of self-propelled spherical particles. The particles are carbon-coated Janus particles, which are propelled due to diffusiophoresis in a near-critical water-lutidine mixture. At low densities, we find that the driving stabilizes small clusters. At higher densities, the suspension undergoes a phase separation into large clusters and a dilute gas phase. The same qualitative behavior is observed in simulations of a minimal model for repulsive self-propelled particles lacking any alignment interactions. The observed behavior is rationalized in terms of a dynamical instability due to the self-trapping of self-propelled particles.

/pdf/dynamical-clustering-and-phase-separation-in-suspensions-of-5st4dsr3sy.pdf

Dynamical clustering and phase separation in suspensions of self-propelled colloidal particles.

Tiny self-propelled swimmers capable of autonomous navigation through complex environments provide appealing opportunities for localization, pick-up and delivery of microscopic and nanoscopic objects. Inspired by motile cells and bacteria, man-made microswimmers have been created and their motion in homogeneous environments has been studied. As a first step towards more realistic conditions under which such microswimmers will be employed, here we study, experimentally and with numerical simulations, their behavior in patterned surroundings that present complex spatial features where frequent encounters with obstacles become important. To study the microswimmers as a function of their swimming behavior, we develop a novel species of microswimmers whose active motion is due to the local demixing of a critical binary liquid mixture and can be easily tuned by illumination. We show that, when microswimmers are confined to a single pore whose diameter is comparable with their swimming length, the probability of finding them at the confinement walls significantly increases compared to Brownian particles. Furthermore, in the presence of an array of periodically arranged obstacles, microswimmers can steer even perpendicularly to an applied force. Since such behavior is very sensitive to the details of their specific swimming style, it can be employed to develop advanced sorting, classification and dialysis techniques.

Microswimmers in patterned environments

Micron-sized self-propelled (active) particles can be considered as model systems for characterizing more complex biological organisms like swimming bacteria or motile cells. We produce asymmetric microswimmers by soft lithography and study their circular motion on a substrate and near channel boundaries. Our experimental observations are in full agreement with a theory of Brownian dynamics for asymmetric self-propelled particles, which couples their translational and orientational motion.

/pdf/circular-motion-of-asymmetric-self-propelling-particles-47mj1fyrru.pdf

Circular motion of asymmetric self-propelling particles.

We demonstrate with experiments and simulations how microscopic self-propelled particles navigate through environments presenting complex spatial features, which mimic the conditions inside cells, living organisms and future lab-on-a-chip devices. In particular, we show that, in the presence of periodic obstacles, microswimmers can steer even perpendicularly to an applied force. Since such behaviour is very sensitive to the details of their specific swimming style, it can be employed to develop advanced sorting, classification and dialysis techniques.

Microswimmers in Patterned Environments

The assembly of artificial nanostructured and microstructured materials which display structures and functionalities that mimic nature’s complexity requires building blocks with specific and directional interactions, analogous to those displayed at the molecular level. Despite remarkable progress in synthesizing “patchy” particles encoding anisotropic interactions, most current methods are restricted to integrating up to two compositional patches on a single “molecule” and to objects with simple shapes. Currently, decoupling functionality and shape to achieve full compositional and geometrical programmability remains an elusive task. We use sequential capillarity-assisted particle assembly which uniquely fulfills the demands described above. This is a new method based on simple, yet essential, adaptations to the well-known capillary assembly of particles over topographical templates. Tuning the depth of the assembly sites (traps) and the surface tension of moving droplets of colloidal suspensions enables controlled stepwise filling of traps to “synthesize” colloidal molecules. After deposition and mechanical linkage, the colloidal molecules can be dispersed in a solvent. The template’s shape solely controls the molecule’s geometry, whereas the filling sequence independently determines its composition. No specific surface chemistry is required, and multifunctional molecules with organic and inorganic moieties can be fabricated. We demonstrate the “synthesis” of a library of structures, ranging from dumbbells and triangles to units resembling bar codes, block copolymers, surfactants, and three-dimensional chiral objects. The full programmability of our approach opens up new directions not only for assembling and studying complex materials with single-particle-level control but also for fabricating new microscale devices for sensing, patterning, and delivery applications.

https://advances.sciencemag.org/content/advances/2/4/e1501779.full.pdf

Ivo Buttinoni

Papers

Dynamical clustering and phase separation in suspensions of self-propelled colloidal particles.

Microswimmers in patterned environments

Circular motion of asymmetric self-propelling particles.

Microswimmers in Patterned Environments

Programmable colloidal molecules from sequential capillarity-assisted particle assembly.