Propulsion by cilia is a fascinating and universal mechanism in biological organisms to generate fluid motion on the cellular level. Cilia are hair-like organelles, which are found in many different tissues and many uni-and multicellular organisms. Assembled in large fields, cilia beat neither randomly nor completely synchronously-instead they display a striking self-organization in the form of metachronal waves (MCWs). It was speculated early on that hydrodynamic interactions provide the physical mechanism for the synchronization of cilia motion. Theory and simulations of physical model systems, ranging from arrays of highly simplified actuated particles to a few cilia or cilia chains, support this hypothesis. The main questions are how the individual cilia interact with the flow field generated by their neighbors and synchronize their beats for the metachronal wave to emerge and how the properties of the metachronal wave are determined by the geometrical arrangement of the cilia, like cilia spacing and beat direction. Here, we address these issues by large-scale computer simulations of a mesoscopic model of 2D cilia arrays in a 3D fluid medium. We show that hydrodynamic interactions are indeed sufficient to explain the self-organization of MCWs and study beat patterns, stability, energy expenditure, and transport properties. We find that the MCW can increase propulsion velocity more than 3-fold and efficiency almost 10-fold-compared with cilia all beating in phase. This can be a vital advantage for ciliated organisms and may be interesting to guide biological experiments as well as the design of efficient microfluidic devices and artificial microswimmers.active matter | mesoscale hydrodynamics | dynamical self-organization F luid transport and locomotion due to motile cilia are ubiquitous phenomena in biological organisms on the cellular level (1, 2). Motile cilia are found in many different tissues-from the brain (3) to the lung and the oviduct-and in many uni-and multicellular organisms-from Clamydomonas (4) and Volvox (5, 6) algae to Paramecium. Motile cilia on the surface of a cell perform an active whip-like motion, which propels the fluid along the surface of cells and tissues. In motile cilia, the beat consists of a fast power stroke in which the cilium has an elongated shape and a slower recovery stroke in which the cilium is curved and closer to the cell surface (Fig. 1A). Due to their typical size in the range of 5-20 μm length and 0.25-1.0 μm thickness, the dynamics of cilia in a fluid are dominated by the balance of force generated by motor proteins (7,8) and fluid viscosity and are thus characterized by small-Reynolds-number hydrodynamics (9). Cilia sometimes act together in pairs, such as in the breast-stroke-like motion of Clamydomonas (4), but much more often in large arrays, such as on the surface of Paramecium and Opalina or the tissue lining the airways of the lung. In all these cases, the beat of different cilia is not random, but strongly synchronized. For many cilia arrays, a wave-like pa...