Locomotion in sturgeon: function of the pectoral fins

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Abstract
Pectoral fins are one of the major features of locomotor design in ray-finned fishes and exhibit a well-documented phylogenetic transition from basal to derived clades. In percomorph fishes, the pectoral fins are often used to generate propulsive force via oscillatory movements, and pectoral fin propulsion in this relatively derived clade has been analyzed extensively. However, in the plesiomorphic pectoral fin condition, exemplified by sturgeon, pectoral fins extend laterally from the body in a generally horizontal orientation, have been assumed to generate lift to balance lift forces and moments produced by the heterocercal tail, and are not oscillated to generate propulsive force. The proposal that pectoral fins in fishes such as sturgeon generate lift during horizontal locomotion has never been tested experimentally in freely swimming fishes. In this paper, we examine the function of pectoral fins in sturgeon swimming at speeds from 0.5–3.0 L s−1, where L is total body length. Sturgeon were studied during steady horizontal locomotion as well as while sinking and rising in the water column. Pectoral fin function was quantified using three-dimensional kinematics to measure the orientation of the fin surface, digital particle image velocimetry (DPIV) was used to describe flow in the wake of the fin and to estimate force exerted on the water, and electromyography was used to assess pectoral fin muscle function. Sturgeon (size range 25–32 cm total length) swam horizontally using continuous undulations of the body with a positive body angle that decreased from a mean of 20 ° at 0.5 L s−1 to 0 ° at 3.0 L s−1.Both the angle of the body and the pectoral fin surface angle changed significantly when sturgeon moved vertically in the water column. Three-dimensional kinematic analysis showed that during steady horizontal swimming the pectoral fins are oriented with a negative angle of attack predicted to generate no significant lift. This result was confirmed by DPIV analysis of the pectoral fin wake, which only revealed fin vortices, and hence force generation, during maneuvering. The orientation of the pectoral fins estimated by a two-dimensional analysis alone is greatly in error and may have contributed to previous suggestions that the pectoral fins are oriented to generate lift. Combined electromyographic and kinematic data showed that the posterior half of the pectoral fin is actively moved as a flap to reorient the head and body to initiate rising and sinking movements. A new force balance for swimming sturgeon is proposed for steady swimming and vertical maneuvering. During steady locomotion, the pectoral fins generate no lift and the positive body angle to the flow is used both to generate lift and to balance moments around the center of mass. To initiate rising or sinking, the posterior portion of the pectoral fins is actively moved ventrally or dorsally, respectively, initiating a starting vortex that, in turn, induces a pitching moment reorienting the body in the flow. Adjustments to body angle initiated by the pectoral fins serve as the primary means by which moments are balanced.