Oropharyngeal morphology related to filtration mechanisms in suspension-feeding American shad (Clupeidae)

To assess potential filtration mechanisms, scanning electron microscopy was used in a comprehensive quantification and analysis of the morphology and surface ultrastructure for all five branchial arches in the ram suspension-feeding fish, American shad (Alosa sapidissima, Clupeidae). The orientation of the branchial arches and the location of mucus cells on the gill rakers were more consistent with mechanisms of crossflow filtration and cross-step filtration rather than conventional dead-end sieving. The long, thin gill rakers could lead to a large area for the exit of water from the oropharyngeal cavity during suspension feeding (high fluid exit ratio). The substantial elongation of gill rakers along the dorsal-ventral axis formed d-type ribs with a groove aspect ratio of 0.5 and a Reynolds number of approximately 500, consistent with the potential operation of cross-step filtration. Mucus cell abundance differed significantly along the length of the raker and the height of the raker. The mucus cell abundance data and the observed sloughing of denticles along the gill raker margins closest to the interior of the oropharyngeal cavity suggest that gill raker growth may occur primarily at the raker tips, the denticle bases, and the internal raker margins along the length of the raker. These findings will be applied in ongoing experiments with 3D-printed physical models of fish oral cavities in flow tanks, and in future ecological studies on the diet and nutrition of suspension-feeding fishes.