Swimbladder Function And Buoyancy Regulation In The Killifish Fundulus Heterochtus
Authors: Goolish Edward M. 
Published: April 2021

ABSTRACT
Killifish, Fundulus heteroclitus, subjected to artificial lift above their center of gravity (10% of body weight) required a minimum of 7–8 days to resorb swimbladder gases completely. The swimbladders of some fish, however, did not fall below 50% of normal volume. The rate of increase in swimbladder volume upon removal of lift varied little among individuals, with approximately 6 days required for complete refilling. Previous deflation of the swimbladder (by syringe) did not result in faster or more complete gas resorption when the fish were subjected to artificial lift. This suggests that the constraint to resorption observed in some fish is not mechanical, e.g. connective tissue, but may reflect individual variability in perception of the stimulus.
Swimbladder dry mass, which scaled as (body mass)0.79, was not affected by exposure to artificial lift. However, fish subjected to 7–11 days of artificial lift displayed slower rates of gas secretion upon removal of lift than control fish whose swimbladders had been evacuated by syringe. The initial rate was 65 % of that of control fish, with two additional days required to achieve normal buoyancy. Also, the rate of swimbladder gas resorption was 24 % faster the second time fish were exposed to artificial lift. These results demonstrate that the capacity for gas secretion and resorption can be altered by previous exposure to hydrostatic challenges.
Killifish buoyancy, expressed as swimbladder volume per weight of the gas-free fish in water, fell from 0.95 to 0.70 ml g−1 after 5 days of exposure to water current. Removal of the pectoral fins eliminated 70% of this decrease, while removal of the pelvic fins had no effect. The rate of gas resorption by fish subjected to artificial lift was also not affected by removal of the pectoral fins. From these results it appears that the decrease in swimbladder volume in fish exposed to water currents is a consequence of lift forces produced by the pectoral fins, but that they are not required for regulation. Fish exposed to water currents or artificial lift swim with a head-down angle of attack. Theoretical estimates show that the vertical force component generated by this swimming behavior is of the appropriate magnitude to compensate for the additional lift.
Fish confined in transparent cages near the surface of the water were less buoyant (0.91 ml g−1) than fish similarly maintained at the bottom of the tank (0.98 ml g−1). However, because this effect was small, 10% of swimbladder volume, visual perception of vertical position is apparently not the primary stimulus for volume regulation. Partial lift (2.65 % of body weight) resulted in the resorption of twice as much swimbladder gas when attachment was anterior to the fish’s center of gravity than when it was an equal distance posterior to the center of gravity. When equal amounts of partial lift and weight were added, lift anterior and weight posterior, no change in swimbladder volume occurred. With the position of these forces reversed, swimbladder volume increased by 31 % to 1.27 ml g−1. These results suggest that fish respond to pitching forces, i.e. longitudinal lift moments, as a stimulus for swimbladder gas secretion and resorption.