Most aquatic vertebrates use suction to capture food, relying on rapid expansion of the mouth cavity to accelerate water and food into the mouth. In ray-finned fishes, mouth expansion is both fast and forceful, and therefore requires considerable power. However, the cranial muscles of these fishes are relatively small and may not be able to produce enough power for suction expansion. The axial swimming muscles of these fishes also attach to the feeding apparatus and have the potential to generate mouth expansion. Because of their large size, these axial muscles could contribute substantial power to suction feeding. To determine whether suction feeding is powered primarily by axial muscles, we measured the power required for suction expansion in largemouth bass and compared it to the power capacities of the axial and cranial muscles. Using X-ray reconstruction of moving morphology (XROMM), we generated 3D animations of the mouth skeleton and created a dynamic digital endocast to measure the rate of mouth volume expansion. This time-resolved expansion rate was combined with intraoral pressure recordings to calculate the instantaneous power required for suction feeding. Peak expansion powers for all but the weakest strikes far exceeded the maximum power capacity of the cranial muscles. The axial muscles did not merely contribute but were the primary source of suction expansion power and generated up to 95% of peak expansion power. The recruitment of axial muscle power may have been crucial for the evolution of high-power suction feeding in ray-finned fishes.epaxial | hypaxial | XROMM | muscle power | volume

Suction-feeding fishes capture food by fast and forceful expansion of the mouth cavity, and axial muscles probably provide substantial power for this feeding behavior. Dorsal expansion of the mouth cavity can only be powered by the epaxial muscles, but both the sternohyoid, shortening against an immobile pectoral girdle to retract the hyoid, and the hypaxial muscles, shortening to retract both the pectoral girdle and hyoid, could contribute ventral expansion power. To determine whether hypaxial muscles generate power for ventral expansion, and the rostrocaudal extent of axial muscle shortening during suction feeding, we measured skeletal kinematics and muscle shortening in largemouth bass (Micropterus salmoides). The threedimensional motions of the cleithrum and hyoid were measured with X-ray reconstruction of moving morphology (XROMM), and muscle shortening was measured with fluoromicrometry, wherein changes in the distance between radio-opaque intramuscular markers are measured using biplanar X-ray video recording. We found that the hypaxials generated power for ventral suction expansion, shortening (mean of 6.2 mm) to rotate the pectoral girdle caudoventrally (mean of 9.3 deg) and retract the hyoid (mean of 8.5 mm). In contrast, the sternohyoid shortened minimally (mean of 0.48 mm), functioning like a ligament to transmit hypaxial shortening to the hyoid. Hypaxial and epaxial shortening were not confined to the rostral muscle regions, but extended more than halfway down the body during suction expansion. We conclude that hypaxial and epaxial muscles are both crucial for powering mouth expansion in largemouth bass, supporting the integration of axial and cranial musculoskeletal systems for suction feeding.

Suction-feeding fish rapidly expand the mouth cavity to generate high-velocity fluid flows that accelerate food into the mouth. Such fast and forceful suction expansion poses a challenge, as muscle power is limited by muscle mass and the muscles in fish heads are relatively small. The largemouth bass powers expansion with its large body muscles, with negligible power produced by the head muscles (including the sternohyoideus). However, bluegill sunfishwith powerful strikes but different morphology and feeding behaviormay use a different balance of cranial and axial musculature to power feeding and different power outputs from these muscles. We estimated the power required for suction expansion in sunfish from measurements of intraoral pressure and rate of volume change, and measured muscle length and velocity. Unlike largemouth bass, the sternohyoideus did shorten to generate power, but it and other head muscles were too small to contribute more than 5-10% of peak expansion power in sunfish. We found no evidence of catapult-style power amplification. Instead, sunfish powered suction feeding by generating high power outputs (up to 438 W kg −1) from their axial muscles. These muscles shortened across the cranial half of the body as in bass, but at faster speeds that may be nearer the optimum for power production. Sunfish were able to generate strikes of the same absolute power as bass, but with 30-40% of the axial muscle mass. Thus, species may use the body and head muscles differently to meet the requirements of suction feeding, depending on their morphology and behavior.