Periodic fluid extrusion and models of digesta mixing in the intestine of a herbivore, the common brushtail possum (Trichosurus vulpecula)

Abstract

The digesta in four gut compartments (proximal and distal halves of small intestine, caecum, and proximal colon) of a wild hindgut fermenting herbivore, the common brushtail possum (Trichosurus vulpecula), were investigated by rheometry and permeametry. Digesta from all compartments were highly viscous and exhibited shear-thinning. Apparent viscosity was positively related to dry matter content, and increased from proximal small intestine to colon. Dynamic rheological measurements showed that in small intestinal digesta the elastic modulus was greater than the viscous modulus and their ratios were characteristic of weak gels, indicating that digesta could sustain compression. The apparent viscosity of distal small intestinal digesta was markedly lower when measured by capillary viscometry than by rotatory viscometry, indicating that plug flow was likely to be facilitated by lubrication from a peripheral layer of less viscous fluid; i.e., there was an augmented plug flow. Permeametry showed that fluid was extruded from all digesta on compression at physiological pressures, that there was significant permeability of proximal and distal small intestinal digesta, but that digesta became progressively compacted during permeation, with a concomitant reduction in permeability as dry matter content increased. It is proposed that conditions within the small intestine differ from those of an ideal plug flow reactor as radial mixing and turbulence cannot occur. Instead, we suggest that segmentation and peristalsis aid radial mixing of the fluid phase by compressing the solid phase, with extrusion of fluid through the digesta plug. This extrusion may be followed by resorption of fluid back into the plug when the elasticity of the solid phase of digesta is Hookean, thus aiding the mixing of secreted enzymes with insoluble substrates within the plug.