Bacterial motility
Bacterial motility is the ability of bacteria to move independently using metabolic energy. Most motility mechanisms that evolved among bacteria also evolved in parallel among the archaea. Most rod-shaped bacteria can move using their own power, which allows colonization of new environments and discovery of new resources for survival. Bacterial movement depends not only on the characteristics of the medium, but also on the use of different appendages to propel. Swarming and [|swimming] movements are both powered by rotating flagella. Whereas swarming is a multicellular 2D movement over a surface and requires the presence of surfactants, swimming is movement of individual cells in liquid environments.
Other types of movement occurring on solid surfaces include twitching, gliding and sliding, which are all independent of flagella. Twitching depends on the extension, attachment to a surface, and retraction of type IV pili which pull the cell forwards in a manner similar to the action of a grappling hook, providing energy to move the cell forward. Bacterial gliding uses different motor complexes, such as the focal adhesion complexes of Myxococcus. Unlike twitching and gliding motilities, which are active movements where the motive force is generated by the individual cell, sliding is a passive movement. It relies on the motive force generated by the cell community due to the expansive forces caused by cell growth within the colony in the presence of surfactants, which reduce the friction between the cells and the surface. The overall movement of a bacterium can be the result of alternating tumble and swim phases. As a result, the trajectory of a bacterium swimming in a uniform environment will form a random walk with relatively straight swims interrupted by random tumbles that reorient the bacterium.
Bacteria can also exhibit taxis, which is the ability to move towards or away from stimuli in their environment. In chemotaxis the overall motion of bacteria responds to the presence of chemical gradients. In phototaxis bacteria can move towards or away from light. This can be particularly useful for cyanobacteria, which use light for photosynthesis. Likewise, magnetotactic bacteria align their movement with the Earth's magnetic field. Some bacteria have escape reactions allowing them to back away from stimuli that might harm or kill. This is fundamentally different from navigation or exploration, since response times must be rapid. Escape reactions are achieved by action potential-like phenomena, and have been observed in biofilms as well as in single cells such as cable bacteria.
Currently there is interest in developing biohybrid microswimmers, microscopic swimmers which are part biological and part engineered by humans, such as swimming bacteria modified to carry cargo.
Background
In 1828, the British biologist Robert Brown discovered the incessant jiggling motion of pollen in water and described his finding in his article "A Brief Account of Microscopical Observations…", leading to extended scientific discussion about the origin of this motion. This enigma was resolved only in 1905, when Albert Einstein published his celebrated essay Über die von der molekularkinetischen Theorie der Wärme geforderte Bewegung von in ruhenden Flüssigkeiten suspendierten Teilchen. Einstein not only deduced the diffusion of suspended particles in quiescent liquids, but also suggested these findings could be used to determine particle size — in a sense, he was the world's first microrheologist.Ever since Newton established his equations of motion, the mystery of motion on the microscale has emerged frequently in scientific history, as famously demonstrated by a couple of articles that should be discussed briefly. First, an essential concept, popularized by Osborne Reynolds, is that the relative importance of inertia and viscosity for the motion of a fluid depends on certain details of the system under consideration. The Reynolds number, named in his honor, quantifies this comparison as a dimensionless ratio of characteristic inertial and viscous forces:
Here, represents the density of the fluid; is a characteristic velocity of the system ; is a characteristic length scale ; and is the viscosity of the fluid. Taking the suspending fluid to be water, and using experimentally observed values for, one can determine that inertia is important for macroscopic swimmers like fish, while viscosity dominates the motion of microscale swimmers like bacteria.
The overwhelming importance of viscosity for swimming at the micrometer scale has profound implications for swimming strategy. This has been discussed memorably by E. M. Purcell, who invited the reader into the world of microorganisms and theoretically studied the conditions of their motion. In the first place, propulsion strategies of large scale swimmers often involve imparting momentum to the surrounding fluid in periodic discrete events, such as vortex shedding, and coasting between these events through inertia. This cannot be effective for microscale swimmers like bacteria: due to the large viscous damping, the inertial coasting time of a micron-sized object is on the order of 1 μs. The coasting distance of a microorganism moving at a typical speed is about 0.1 angstroms. Purcell concluded that only forces that are exerted in the present moment on a microscale body contribute to its propulsion, so a constant energy conversion method is essential.
Microorganisms have optimized their metabolism for continuous energy production, while purely artificial microswimmers must obtain energy from the environment, since their on-board-storage-capacity is very limited. As a further consequence of the continuous dissipation of energy, biological and artificial microswimmers do not obey the laws of equilibrium statistical physics, and need to be described by non-equilibrium dynamics. Mathematically, Purcell explored the implications of low Reynolds number by taking the Navier-Stokes equation and eliminating the inertial terms:
where is the velocity of the fluid and is the gradient of the pressure. As Purcell noted, the resulting equation — the Stokes equation — contains no explicit time dependence. This has some important consequences for how a suspended body can swim through periodic mechanical motions or deformations. First, the rate of motion is practically irrelevant for the motion of the microswimmer and of the surrounding fluid: changing the rate of motion will change the scale of the velocities of the fluid and of the microswimmer, but it will not change the pattern of fluid flow. Secondly, reversing the direction of mechanical motion will simply reverse all velocities in the system. These properties of the Stokes equation severely restrict the range of feasible swimming strategies.
As a concrete illustration, consider a mathematical scallop that consists of two rigid pieces connected by a hinge. Can the "scallop" swim by periodically opening and closing the hinge? No: regardless of how the cycle of opening and closing depends on time, the scallop will always return to its starting point at the end of the cycle. Here originated the striking quote: "Fast or slow, it exactly retraces its trajectory and it's back where it started". In light of this scallop theorem, Purcell developed approaches concerning how artificial motion at the micro scale can be generated. This paper continues to inspire ongoing scientific discussion; for example, recent work by the Fischer group from the Max Planck Institute for Intelligent Systems experimentally confirmed that the scallop principle is only valid for Newtonian fluids.
Motile systems have developed in the natural world over time and length scales spanning several orders of magnitude, and have evolved anatomically and physiologically to attain optimal strategies for self-propulsion and overcome the implications of high viscosity forces and Brownian motion, as shown in the diagram on the right.
Some of the smallest known motile systems are motor proteins, i.e., proteins and protein complexes present in cells that carry out a variety of physiological functions by transducing chemical energy into mechanical energy. These motor proteins are classified as myosins, kinesins, or dyneins. Myosin motors are responsible for muscle contractions and the transport of cargousing actin filaments as tracks. Dynein motors and kinesin motors, on the other hand, use microtubules to transport vesicles across the cell. The mechanism these protein motors use to convert chemical energy into movement depends on ATP hydrolysis, which leads to a conformation modification in the globular motor domain, leading to directed motion.
Bacteria can be roughly divided into two fundamentally different groups, gram-positive and gram-negative bacteria, distinguished by the architecture of their cell envelope. In each case the cell envelope is a complex multi-layered structure that protects the cell from its environment. In gram-positive bacteria, the cytoplasmic membrane is only surrounded by a thick cell wall of peptidoglycan. By contrast, the envelope of gram-negative bacteria is more complex and consists of the cytoplasmic membrane, a thin layer of peptidoglycan, and an additional outer membrane, also called the lipopolysaccharide layer. Other bacterial cell surface structures range from disorganised slime layers to highly structured capsules. These are made from secreted slimy or sticky polysaccharides or proteins that provide protection for the cells and are in direct contact with the environment. They have other functions, including attachment to solid surfaces. Additionally, protein appendages can be present on the surface: fimbriae and pili can have different lengths and diameters and their functions include adhesion and twitching motility.
Specifically, for microorganisms that live in aqueous environments, locomotion refers to swimming, and hence the world is full of different classes of swimming microorganisms, such as bacteria, protozoa, and algae. Bacteria move due to rotation of hair-like filaments called flagella, which are anchored to a protein motor complex on the bacteria cell wall.