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Brain cilia are non-motile at early larval stage; some are motile at late larval stage. a, c Arl13b: GFP is expressed in cilia throughout the developing fish including the neuroepithelium and spinal canal. b, d Quantification of ciliary beat frequency (Hz) for motile cilia in each region at early (27 hpf) and late (51 hpf) larval stages, if any. e, f Visualization of ciliary movement for cilia in each region at 27 and 51 hpf, if any. Each of the three panels is an overlay of two sequential image acquisition frames (images were taken in the same region at distinct times at 57.22 fps, so each overlay represents a time difference of ~0.017 s). See Additional file 2: Movie 1 and Additional file 3: Movie 2 for full dataset. Colors were selected such that unmoved pixels appear black (Frame 1: green + magenta; Frame 2: red + cyan). Horizontal lines represent average vmax and error bars denote SEM. T telencephalic ventricle, D/M diencephalic/mesencephalic ventricle, R rhombencephalic ventricle, SC spinal canal ventricle analysis region, hpf hours post-fertilization, N.A. quantification of cilia frequency is not applicable because no cilia are motile. Scale bars: a, c: 100 μm; e, f: 5 μm
At early larval stages no motile cilia are present in the neuroepithelium and are therefore unlikely to contribute to the CSF movement observed. The non-motile cilia at this time may be able to detect CSF movement, potentially to activate planar cell polarity signaling described by Ohata and colleagues [21], or to guide migrating neurons or extending axons [54]. At late larval stages, cilia along the D ventricle are highly motile and may indicate differentiating CP, as CP cells have motile cilia [52]. As discussed above, these ciliary movements or CSF production may drive CSF movement from D to T ventricles. Mutants that disrupt the primary cilia present with hydrocephalus [53], which would preclude clear comparison of CSF movement. However, future studies acutely disrupting cilia are needed to fully address potential roles of cilia in CSF movement.
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