Conservation, development, and function of a cement gland-like structure in the fish Astyanax mexicanus

The larvae of the fish Astyanax mexicanus transiently develop a flat and adhesive structure on the top of their heads that we have called "the casquette" (cas, meaning "hat"). We hypothesized that the cas may be a teleostean homolog of the well-studied Xenopus cement gland, despite their different positions and structures. Here we show that the cas has an ectodermal origin, secretes mucus, expresses bone morphogenic protein 4 (Bmp4) and pituitary homeobox 1/2 (Pitx1/2), is innervated by the trigeminal ganglion and serotonergic raphe neurons, and has a role in the control and the development of the larval swimming behavior. These developmental, connectivity, and behavioral functional data support a level of deep homology between the frog cement gland and the Astyanax cas and suggest that attachment organs can develop in varied positions on the head ectoderm by recruitment of a Bmp4-dependent developmental module. We also show that the attachment organs of the cichlid Tilapia mariae larvae display some of these features. We discuss the possibility that these highly diversified attachment glands may be ancestral to chordates and have been lost repetitively in many vertebrate classes.