Appearance of somatostatin and vasoactive intestinal peptide along the developing chicken gut

Abstract
The appearance of somatostatin (SOM)-immunoreactive (IR) and vasoactive intestinal peptide (VIP)-IR neurons in different regions of the embryonic chicken gut was studied by immunostaining wholemounts. The patterns of expression of these peptides in myenteric neurons showed a number of similarities. Both peptides first appeared in the region of the proventriculus-gizzard: SOM at embryonic day (E)4, VIP at E5.5. At later times both peptides were found in positions both rostral and caudal to the gizzard. Both peptides appeared independently in cells at a second site, the cecum of the hindgut: SOM was observed at E6.5 and VIP at E7.5. VIP-IR and SOM-IR cells appear throughout the cecum, then in the rectum, and finally in the ileum. Differences in the patterns of expression were also found. SOM- and VIP-IR neurons appeared at different times along the length of the gut. VIP-IR cells populated the entire gut by E11.5, whereas SOM-IR cells were not present throughout the gut until E13.5. SOM-IR cells appeared in the terminal part of the ganglion of Remak at E4.0. At E6 these SOM-IR cells sent fibers into the wall of the hindgut and later into the midgut. No VIP-IR cells were found in the ganglion of Remak. These findings suggest that neural crest-derived cells first express SOM- and VIP-IR in particular regions of the gut, namely, the proventriculus-gizzard and the cecum. Certain conditions must exist at these sites which favor the expression of these neuropeptides by neural crest-derived cells. The observation of SOM- and VIP-IR cells in the cecum at a stage of development before cells are seen in the ileum supports the concept that sacral neural crest cells contribute precursors for enteric neurons of the avian hindgut.