Mechanisms for bacterial gliding motility on soft substrates

The motility mechanism of certain rod-shaped bacteria has long been a mystery, since no external appendages are involved in their motion which is known as gliding. However, the physical principles behind gliding motility still remain poorly understood. Using myxobacteria as a canonical example of such organisms, we identify here the physical principles behind gliding motility, and develop a theoretical model that predicts a two-regime behavior of the gliding speed as a function of the substrate stiffness. Our theory describes the elastic, viscous, and capillary interactions between the bacterial membrane carrying a traveling wave, the secreted slime acting as a lubricating film, and the substrate which we model as a soft solid. Defining the myxobacterial gliding as the horizontal motion on the substrate under zero net force, we find the two-regime behavior is due to two different mechanisms of motility thrust. On stiff substrates, the thrust arises from the bacterial shape deformations creating a flow of slime that exerts a pressure along the bacterial length. This pressure in conjunction with the bacterial shape provides the necessary thrust for propulsion. However, we show that such a mechanism cannot lead to gliding on very soft substrates. Instead, we show that capillary effects lead to the formation of a ridge at the slime-substrate-air interface, which creates a thrust in the form of a localized pressure gradient at the tip of the bacteria. To test our theory, we perform experiments with isolated cells on agar substrates of varying stiffness and find the measured gliding speeds to be in good agreement with the predictions from our elasto-capillary-hydrodynamic model. The physical mechanisms reported here serve as an important step towards an accurate theory of friction and substrate-mediated interaction between bacteria in a swarm of cells proliferating in soft media.Comment: Main article (8 pages and 8 figures) and Supplementary Informatio

Roadmap on emerging concepts in the physical biology of bacterial biofilms: from surface sensing to community formation

Bacterial biofilms are communities of bacteria that exist as aggregates that can adhere to surfaces or be free-standing. This complex, social mode of cellular organization is fundamental to the physiology of microbes and often exhibits surprising behavior. Bacterial biofilms are more than the sum of their parts: single-cell behavior has a complex relation to collective community behavior, in a manner perhaps cognate to the complex relation between atomic physics and condensed matter physics. Biofilm microbiology is a relatively young field by biology standards, but it has already attracted intense attention from physicists. Sometimes, this attention takes the form of seeing biofilms as inspiration for new physics. In this roadmap, we highlight the work of those who have taken the opposite strategy: we highlight the work of physicists and physical scientists who use physics to engage fundamental concepts in bacterial biofilm microbiology, including adhesion, sensing, motility, signaling, memory, energy flow, community formation and cooperativity. These contributions are juxtaposed with microbiologists who have made recent important discoveries on bacterial biofilms using state-of-the-art physical methods. The contributions to this roadmap exemplify how well physics and biology can be combined to achieve a new synthesis, rather than just a division of labor

Novel mechanisms power bacterial gliding motility.

For many bacteria, motility is essential for survival, growth, virulence, biofilm formation and intra/interspecies interactions. Since natural environments differ, bacteria have evolved remarkable motility systems to adapt, including swimming in aqueous media, and swarming, twitching and gliding on solid and semi-solid surfaces. Although tremendous advances have been achieved in understanding swimming and swarming motilities powered by flagella, and twitching motility powered by Type IV pili, little is known about gliding motility. Bacterial gliders are a heterogeneous group containing diverse bacteria that utilize surface motilities that do not depend on traditional flagella or pili, but are powered by mechanisms that are less well understood. Recently, advances in our understanding of the molecular machineries for several gliding bacteria revealed the roles of modified ion channels, secretion systems and unique machinery for surface movements. These novel mechanisms provide rich source materials for studying the function and evolution of complex microbial nanomachines. In this review, we summarize recent findings made on the gliding mechanisms of the myxobacteria, flavobacteria and mycoplasmas

Novel mechanisms power bacterial gliding motility.

For many bacteria, motility is essential for survival, growth, virulence, biofilm formation and intra/interspecies interactions. Since natural environments differ, bacteria have evolved remarkable motility systems to adapt, including swimming in aqueous media, and swarming, twitching and gliding on solid and semi-solid surfaces. Although tremendous advances have been achieved in understanding swimming and swarming motilities powered by flagella, and twitching motility powered by Type IV pili, little is known about gliding motility. Bacterial gliders are a heterogeneous group containing diverse bacteria that utilize surface motilities that do not depend on traditional flagella or pili, but are powered by mechanisms that are less well understood. Recently, advances in our understanding of the molecular machineries for several gliding bacteria revealed the roles of modified ion channels, secretion systems and unique machinery for surface movements. These novel mechanisms provide rich source materials for studying the function and evolution of complex microbial nanomachines. In this review, we summarize recent findings made on the gliding mechanisms of the myxobacteria, flavobacteria and mycoplasmas

Mechanisms for bacterial gliding motility on soft substrates.

The motility mechanism of certain prokaryotes has long been a mystery, since their motion, known as gliding, involves no external appendages. The physical principles behind gliding still remain poorly understood. Using myxobacteria as an example of such organisms, we identify here the physical principles behind gliding motility and develop a theoretical model that predicts a 2-regime behavior of the gliding speed as a function of the substrate stiffness. Our theory describes the elasto-capillary-hydrodynamic interactions between the membrane of the bacteria, the slime it secretes, and the soft substrate underneath. Defining gliding as the horizontal translation under zero net force, we find the 2-regime behavior is due to 2 distinct mechanisms of motility thrust. On mildly soft substrates, the thrust arises from bacterial shape deformations creating a flow of slime that exerts a pressure along the bacterial length. This pressure in conjunction with the bacterial shape provides the necessary thrust for propulsion. On very soft substrates, however, we show that capillary effects must be considered that lead to the formation of a ridge at the slime-substrate-air interface, thereby creating a thrust in the form of a localized pressure gradient at the bacterial leading edge. To test our theory, we perform experiments with isolated cells on agar substrates of varying stiffness and find the measured gliding speeds in good agreement with the predictions from our elasto-capillary-hydrodynamic model. The mechanisms reported here serve as an important step toward an accurate theory of friction and substrate-mediated interactions between bacteria proliferating in soft media