Roll and Yaw of Paramecium swimming in a viscous fluid

Many free-swimming microorganisms like ciliates, flagellates, and invertebrates exhibit helical trajectories. In particular, the Paramecium spirally swims along its anterior direction by the beating of cilia. Due to the oblique beating stroke of cilia, the Paramecium rotates along its long axis as it swims forward. Simultaneously, this long axis turns toward the oral groove side. Combined roll and yaw motions of Paramecium result in swimming along a spiral course. Using Particle Image Velocimetry, we measure and quantify the flow field and fluid stress around Paramecium. We will discuss how the non-uniform stress distribution around the body induces this yaw motion.