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Motile cilia are hair-like cell extensions that beat periodically to generate fluid flow along various epithelial tissues within the body. In dense multiciliated carpets, cilia were shown to exhibit a remarkable coordination of their beat in the form of traveling metachronal waves, a phenomenon which supposedly enhances fluid transport. Yet, how cilia coordinate their regular beat in multiciliated epithelia to move fluids remains insufficiently understood, particularly due to lack of rigorous quantification. We combine experiments, novel analysis tools, and theory to address this knowledge gap. To investigate collective dynamics of cilia, we studied zebrafish multiciliated epithelia in the nose and the brain. We focused mainly on the zebrafish nose, due to its conserved properties with other ciliated tissues and its superior accessibility for non-invasive imaging. We revealed that cilia are synchronized only locally and that the size of local synchronization domains increases with the viscosity of the surrounding medium. Even though synchronization is local only, we observed global patterns of traveling metachronal waves across the zebrafish multiciliated epithelium. Intriguingly, these global wave direction patterns are conserved across individual fish, but different for left and right noses, unveiling a chiral asymmetry of metachronal coordination. To understand the implications of synchronization for fluid pumping, we used a computational model of a regular array of cilia. We found that local metachronal synchronization prevents steric collisions, i.e., cilia colliding with each other, and improves fluid pumping in dense cilia carpets, but hardly affects the direction of fluid flow. In conclusion, we show that local synchronization together with tissue-scale cilia alignment coincide and generate metachronal wave patterns in multiciliated epithelia, which enhance their physiological function of fluid pumping.

The defects present in solution-processed perovskite light absorbing materials are responsible for deteriorating the performance and stability of perovskite solar cells. Therefore, suppressing the defect formation and passivating the pre-existing defects in perovskite materials are of paramount importance in pushing the efficiency of perovskite solar cells (PSCs) toward the theoretical limit and ensuring the long-term stability of PSCs. In this work, we report an efficient defect passivation route using a bidentate small organic molecule, pyrazine, which is capable of making either a monodentate or a bridging bidentate bond with under-coordinated Pb2+ ions mainly present in the surface and grain boundaries of perovskite films. The various analyses for the defect density and charge carrier lifetime of perovskite light absorbing layers demonstrated that pyrazine effectively passivated defects existing on the surface and grain boundaries of the perovskite films without affecting the morphology and crystallinity via a simple post-treatment procedure. As a result, the best untreated device obtained a power conversion efficiency of 19.14%, while an efficiency of 20.58% was achieved when pyrazine treated.