Functional cytology of the hepatopancreas of decapod crustaceans

This article reviews the morphogenesis, morphology, histology, ultrastructure, and structural-functional relationships of the hepatopancreas, the main metabolic organ of the Decapoda. The hepatopancreas develops in early larval stages from a pair of lateral lobes of the midgut anlage. In adults, it consists of hundreds of blindly ending tubules that are enveloped by a muscle net consisting of longitudinal and circular fibers. Stem cells at the distal ends of the tubules give rise to three ultrastructurally different epithelial cell types, the R-, F-, and B-cells. Histochemistry, immunohistochemistry, in situ hybridization, and monitoring of ultrastructural changes under different experimental conditions allowed the attribution of functions to these cell types. R-cells serve for the absorption and metabolization of nutrients, storage of energy reserves and minerals, synthesis of lipoproteins for export to other organs, detoxification of heavy metals, and excretion of uric acid. F-cells synthesize digestive enzymes and blood proteins involved in oxygen transport and immune defense. They also detoxify some heavy metals and probably organic xenobiotics. B-cells are assumed to produce and recycle fat emulsifiers. The hepatopancreas tubules lack nerves. The presence of scattered M-cells with putative endocrine function in the epithelium suggests that the hepatopancreas is mainly hormonally controlled. M-cells probably represent a self-perpetuating cell lineage independent from E-cells. The interstitium between the tubules contains connective tissue, arterioles, hemolymph with circulating hemocytes, and fixed phagocytes that eliminate pathogens. The hepatopancreas is histologically and ultrastructurally uniform throughout the Decapoda, despite their broad variety in body size, morphology, life style, and ecology. However, in a few cavernicolous and deep-sea shrimps parts of the hepatopancreas are transformed into large oil storing and bioluminescent compartments. Within the malacostracan crustaceans, the hepatopancreas of the Decapoda is most similar to the digestive gland of the Euphausiacea, supporting close taxonomic relationship of these two taxa.