Microorganisms living in gradient environments affect large-scale processes, including the cycling of elements such as carbon, nitrogen or sulfur, the rates and fate of primary production, and the generation of climatically active gases. Aerotaxis is a common adaptation in organisms living in the oxygen gradients of stratified environments. Magnetotactic bacteria are such gradient-inhabiting organisms that have a specific type of aerotaxis that allows them to compete at the oxic-anoxic interface. They biomineralize magnetosomes, intracellular membrane-coated magnetic nanoparticles, that comprise a permanent magnetic dipole that causes the cells to align along magnetic field lines. The magnetic alignment enables them to efficiently migrate toward an optimal oxygen concentration in microaerobic niches. This phenomenon is known as magneto-aerotaxis. Magneto-aerotaxis has only been characterized in a limited number of available cultured strains. In this work, we characterize the magneto-aerotactic behavior of 12 magnetotactic bacteria with various morphologies, phylogenies, physiologies, and flagellar apparatus. We report six different magneto-aerotactic behaviors that can be described as a combination of three distinct mechanisms, including the reported (di-)polar, axial, and a previously undescribed mechanism we named unipolar. We implement a model suggesting that the three magneto-aerotactic mechanisms are related to distinct oxygen sensing mechanisms that regulate the direction of cells' motility in an oxygen gradient.

We show that we can select magnetically steerable nanopropellers from a set of carbon coated aggregates of magnetic nanoparticles using weak homogeneous rotating magnetic fields. The carbon coating can be functionalized, enabling a wide range of applications. Despite their arbitrary shape, all nanostructures propel parallel to the vector of rotation of the magnetic field. We use a simple theoretical model to find experimental conditions to select nanopropellers which are predominantly smaller than previously published ones.

The response of cells to changes in their physico-chemical micro-environment is essential to their survival. For example, bacterial magnetotaxis uses the Earth's magnetic field together with chemical sensing to help microorganisms move towards favoured habitats. The studies of such complex responses are lacking a method that permits the simultaneous mapping of the chemical environment and the response of the organisms, and the ability to generate a controlled physiological magnetic field. We have thus developed a multi-modal microscopy platform that fulfils these requirements. Using simultaneous fluorescence and high-speed imaging in conjunction with diffusion and aerotactic models, we characterized the magneto- aerotaxis of Magnetospirillum gryphiswaldense. We assessed the influence of the magnetic field (orientation; strength) on the formation and the dynamic of a micro-aerotactic band (size, dynamic, position). As previously described by models of magnetotaxis, the application of a magnetic field pointing towards the anoxic zone of an oxygen gradient results in an enhanced aerotaxis even down to Earth's magnetic field strength. We found that neither a ten-fold increase of the field strength nor a tilt of 45° resulted in a significant change of the aerotactic efficiency. However, when the field strength is zeroed or when the field angle is tilted to 90°, the magneto-aerotaxis efficiency is drastically reduced. The classical model of magneto-aerotaxis assumes a response proportional to the cosine of the angle difference between the directions of the oxygen gradient and that of the magnetic field. Our experimental evidence however shows that this behaviour is more complex than assumed in this model, thus opening up new avenues for research.

Studying propulsion mechanisms in low Reynolds number fluid has implications for many fields, ranging from the biology of motile microorganisms and the physics of active matter to micromixing in catalysis and micro- and nanorobotics. The propulsion of magnetic micropropellers can be characterized by a dimensionless speed, which solely depends on the propeller geometry for a given axis of rotation. However, this dependence has so far been only investigated for helical propeller shapes, which were assumed to be optimal. In order to explore a larger variety of shapes, we experimentally studied the propulsion properties of randomly shaped magnetic micropropellers. Surprisingly, we found that their dimensionless speeds are high on average, comparable to previously reported nanofabricated helical micropropellers. The highest dimensionless speed we observed is higher than that of any previously reported propeller moving in a low Reynolds number fluid, proving that physical random shape generation can be a viable optimization strategy.

Magnetic actuation of microscopic devices in a liquid environment has been achieved in various ways, which can be grouped into rolling, propelling and swimming. Previous actuators were designed with a focus on one particular type of magnetic actuation. We have shown earlier that efficient magnetic propellers can be selected from randomly shaped magnetic nanostructures synthesized in solution. Here we show that these synthesized nanostructures can be used for all three types of magnetic actuation. Whereas it might not be surprising that single structures can roll in addition to propelling, swimming is unexpectedly also observed using the same material. In this case, however, the magnetically guided self-assembly of several individual particles into chain-like structures is necessary to obtain swimmers, since individual rigid nanostructures cannot swim. Interestingly, the direction of the swimming motion is not necessarily parallel to the long axis of the chain-like assembly, a finding that had been theoretically expected but experimentally not observed so far. Our findings show that the range of structures that can be effectively actuated by external magnetic fields is much broader than assumed until now. This could open up new opportunities for the design of magnetically actuated devices.