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The Cilium
Now, are any biochemical systems irreducibly complex? Yes, it turns out that many are. A good example is the cilium. Cilia are hairlike structures on the surfaces of many animal and lower plant cells that can move fluid over the cell's surface or "row" single cells through a fluid. Inhumans, for example, cells lining the respiratory tract each have about 200 cilia that beat in synchrony to sweep mucus towards the throat for elimination. What is the structure of a cilium? A cilium consists of bundle of fibers called an axoneme. An axoneme contains a ring of 9 double "microtubules" surrounding two central single microtubules. Each outer doublet consists of a ring of 13 filaments (subfiber A) fused to an assembly of 10 filaments (subfiber B). The filaments of the microtubules are composedof two proteins called alpha and beta tubulin. The 11 microtubules forming an axoneme are held together by three types of connectors: subfibers A are joined to the central microtubules by radial spokes; adjacent outer doublets are joined by linkers of a highly elastic protein called nexin; and the central microtubules are joined by a connecting bridge. Finally, every subfiber A bears two arms, an inner arm and an outer arm, both containing a protein called dynein.

But how does a cilium work? Experiments have shown that ciliary motion results from the chemically-powered "walking" of the dynein arms on one microtubule up a second microtubule so that the two microtubules slide past each other. The protein cross-links between microtubules in a cilium prevent neighboring microtubules from sliding past each other by more than a short distance. These cross-links, therefore, convert the dynein-induced sliding motion to a bending motion of the entire axoneme.

Now, let us consider what this implies. What components are needed for a cilium to work? Ciliary motion certainly requires microtubules; otherwise, there would be no strands to slide. Additionally we require a motor, or else the microtubules of the cilium would lie stiff and motionless. Furthermore, we require linkers to tug on neighboring strands, converting the sliding motion into a bending motion, and preventing the structure from falling apart. All of these parts are required to perform one function: ciliary motion. Just as a mousetrap does not work unless all of its constituent parts are present, ciliary motion simply does not exist in the absence of microtubules, connectors, and motors. Therefore, we can conclude that the cilium is irreducibly complex; an enormous monkey wrench thrown into its presumed gradual, Darwinian evolution.