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 characterized by a decrease in the swimming amplitude.
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See supplementary material at http://dx.doi.org/10.1063/1.4897651 for examples of RFT fits for swimming near one and two boundaries, and for movies of the worm swimming before and after confinement.
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