Matilda Backholm

Viscoelastic properties of the nematode Caenorhabditis elegans, a self-similar, shear-thinning worm

Undulatory motion is common to many creatures across many scales, from sperm to snakes. These organisms must push off against their external environment, such as a viscous medium, grains of sand, or a high-friction surface; additionally they must work to bend their own body. A full understanding of undulatory motion, and locomotion in general, requires the characterization of the material properties of the animal itself. The material properties of the model organism Caenorhabditis elegans were studied with a micromechanical experiment used to carry out a three-point bending measurement of the worm. Worms at various developmental stages (including dauer) were measured and different positions along the worm were probed. From these experiments we calculated the viscoelastic properties of the worm, including the effective spring constant and damping coefficient of bending. C. elegans moves by propagating sinusoidal waves along its body. Whereas previous viscoelastic approaches to describe the undulatory motion have used a Kelvin–Voigt model, where the elastic and viscous components are connected in parallel, our measurements show that the Maxwell model, where the elastic and viscous components are in series, is more appropriate. The viscous component of the worm was shown to be consistent with a non-Newtonian, shear-thinning fluid. We find that as the worm matures it is well described as a self-similar elastic object with a shear-thinning damping term and a stiffness that becomes smaller as one approaches the tail.

Direct Measurements of Drag Forces in C. elegans Crawling Locomotion

With a simple and versatile microcantilever-based force measurement technique, we have probed the drag forces involved in Caenorhabditis elegans locomotion. As a worm crawls on an agar surface, we found that substrate viscoelasticity introduces nonlinearities in the force-velocity relationships, yielding nonconstant drag coefficients that are not captured by original resistive force theory. A major contributing factor to these nonlinearities is the formation of a shallow groove on the agar surface. We measured both the adhesion forces that cause the worms body to settle into the agar and the resulting dynamics of groove formation. Furthermore, we quantified the locomotive forces produced by C. elegans undulatory motions on a wet viscoelastic agar surface. We show that an extension of resistive force theory is able to use the dynamics of a nematodes body shape along with the measured drag coefficients to predict the forces generated by a crawling nematode.

Undulatory microswimming near solid boundaries

The hydrodynamic forces involved in the undulatory microswimming of the model organism C. elegans are studied in proximity to solid boundaries. Using a micropipette deflection technique, we attain direct and time-resolved force measurements of the viscous forces acting on the worm near a single planar boundary as well as confined between two planar boundaries. We observe a monotonic increase in the lateral and propulsive forces with increasing proximity to the solid interface. We determine normal and tangential drag coefficients for the worm, and find these to increase with confinement. The measured drag coefficients are compared to existing theoretical models. The ratio of normal to tangential drag coefficients is found to assume a constant value of 1.5 ± 0.1(5) at all distances from a single boundary, but increases significantly as the worm is confined between two boundaries. In response to the increased drag due to confinement, we observe a gait modulation of the nematode, which is primarily characteri...

The effects of viscosity on the undulatory swimming dynamics of C. elegans

The undulatory swimming dynamics of the millimetric nematode Caenorhabditis elegans was investigated in fluids with different viscosities. The technique of micropipette deflection was used to directly measure the drag forces experienced by the swimming worm in both the lateral and propulsive directions. Gait modulation due to increasing viscosity in our tethered system was found to be qualitatively similar to that of freely swimming worms. Resistive force theory was used to determine the drag coefficients of the slender swimmer, and the experimental values were compared to the classical theories of Lighthill as well as Gray and Hancock. The gait modulation was shown to be independent of how the environmental resistance is changed, indicating the relevance of only the fluid resistance on the swimming kinematics and dynamics of the nematode.

Relaxation and Intermediate Asymptotics of a Rectangular Trench in a Viscous Film

The surface of a thin liquid film with nonconstant curvature flattens as a result of capillary forces. While this leveling is driven by local curvature gradients, the global boundary conditions greatly influence the dynamics. Here, we study the evolution of rectangular trenches in a polystyrene nanofilm. Initially, when the two sides of a trench are well separated, the asymmetric boundary condition given by the step height controls the dynamics. In this case, the evolution results from the leveling of two noninteracting steps. As the steps broaden further and start to interact, the global symmetric boundary condition alters the leveling dynamics. We report on full agreement between theory and experiments for the capillary-driven flow and resulting time dependent height profiles, a crossover in the power-law dependence of the viscous energy dissipation as a function of time as the trench evolution transitions from two noninteracting to interacting steps, and the convergence of the profiles to a universal self-similar attractor that is given by the Greens function of the linear operator describing the dimensionless linearized thin film equation.

The nematode C. elegans as a complex viscoelastic fluid

The viscoelastic material properties of the model organism C. elegans were probed with a micropipette deflection technique and modelled with the standard linear solid model. Dynamic relaxation measurements were performed on the millimetric nematode to investigate its viscous characteristics in detail. We show that the internal properties of C. elegans can not be fully described by a simple Newtonian fluid. Instead, a power-law fluid model was implemented and shown to be in excellent agreement with experimental results. The nematode exhibits shear thinning properties and its complex fluid characteristics were quantified. The bending-rate dependence of the internal damping coefficient of C. elegans could affect its gait modulation in different external environments.Graphical abstract