Comparison of acid mucin goblet cell distribution and Hox13 expression patterns in the developing vertebrate digestive tract.

The digestive tract of vertebrates is a complex organ system required for the digestion of food and the absorption of nutrients. The colon evolved as a water absorption organ essential for vertebrates to survive on land. In contrast to land vertebrates, the Chondrichthyes (sharks, skates and rays) are nearly iso-osmotic with their ocean environment and do not reabsorb water from food waste. To understand the origin of the vertebrate colon, we examined the distribution of sulfated and sialyated mucus-producing cells in the little skate, Raja erinacea, as an indication of water absorption function in the chondrichthian digestive tract. The percentage of acid mucin producing goblet cells was analyzed in the spiral valve and hindgut of little skate and the small intestine and colon of mouse embryos. Levels of acid mucins in the hindgut of the little skate was comparable to that of the small intestines of terrestrial vertebrates, whereas the distal region of the spiral valve contained high levels of acid mucin producing cells similar to the colon of mouse and chick. The low numbers of acid mucins in the little skate hindgut confirms that a functional colon for water absorption is absent in the Chondrichthyes. Interestingly, the presence of high levels of acid mucins in the posterior spiral valve provides evidence for a possible primordial water-absorbing organ in the elasmobranchs. Hoxd13 patterns acid mucins in the colons of terrestrial vertebrates. Expression of Hoxd13 and Hoxa13 in R. erinacea suggests conserved roles for Hox genes in patterning the early hindgut.