Innervation of the monotreme gastrointestinal tract: a study of peptide and catecholamine distribution

The distribution of neurons and endocrine cells containing various peptides or catecholamines was examined in the digestive tracts of the echidna (Tachyglossus aculeatus) and the platypus (Ornithorhynchus anatinus). Comparisons were made with published studies in other species in order to obtain a broader view of the phylogenetic distribution and possible functions of gut peptides and catecholamines. Further comparisons between the echidna and platypus were made in light of their different dietary features and gut histology. The distribution of neurons and axons containing catecholamines or various peptides resembled that in other species (such as the frequent appearance of axons containing substance P and vasoactive intestinal peptide in the intestinal mucosa, and axons containing substance P or enkephalins in the circular muscle). In both species, the stomach histologically resembles the esophagus, being aglandular and lined with stratified squamous epithelium. Innervation of these two organs was similar but not identical, with a greater array of peptides found in the gastric muscle. The intestinal mucosa was densely innervated in both species. The platypus small intestine is unusual in having a thick and deeply folded mucosa (but no villi), in which the superficial epithelium is absent or incomplete at many sites; many axons travel close to these luminal surfaces. Many (putative noradrenergic) axons associated with blood vessels contained neuro-peptide Y, but there was no evidence for intrinsic catecholamine-containing neurons. Somatostatin and cholecystokinin were present in some endocrine cells, but unlike many mammals, absent in neuronal tissue. These studies have shown that there are many strong similarities between monotremes and other mammals in the distribution and array of peptides found within nervous and endocrine tissues of the digestive tract. However, numerous small differences of the echidna and platypus innervation may be correlated with their different digestive structures.