The squirmer model of Lighthill and Blake has been widely used to analyse swimming ciliates. However, real ciliates are covered by hair-like organelles, called cilia; the differences between the squirmer model and real ciliates remain unclear. Here, we developed a ciliate model incorporating the distinct ciliary apparatus, and analysed motion using a boundary element-slender-body coupling method. This methodology allows us to accurately calculate hydrodynamic interactions between cilia and the cell body under free-swimming conditions. Results showed that an antiplectic metachronal wave was optimal in the swimming speed with various cell-body aspect ratios, which is consistent with former theoretical studies. Exploiting oblique wave propagation, we reproduced a helical trajectory, like Paramecium, although the cell body was spherical. We confirmed that the swimming velocity of model ciliates was well represented by the squirmer model. However, squirmer modelling outside the envelope failed to estimate the energy costs of swimming; over 90 % of energy was dissipated inside the ciliary envelope. The optimal swimming efficiency was given by the antiplectic wave; the value was 6.7 times larger than in-phase beating. Our findings provide a fundamental basis for modelling swimming micro-organisms.
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