First investigations on lamprey responses to elevated total dissolved gas exposure and risk of gas bubble trauma

Abstract

A flexible spill program in the federal Columbia River power system increased the total dissolved gas (TDG) water quality standards (i.e., the gas cap) from 120% to 125%. Spill is used to pass juvenile salmon (Oncorhynchus spp.) over dams, but it can generate elevated TDG, and exposed fish can develop gas bubble trauma (GBT) or experience mortality. Juvenile salmon are monitored for GBT through the Fish Passage Center’s (FPC), and under the flexible spill program, native non-salmonid fishes are also monitored. Pacific Lamprey (Entosphenus tridentatus) are exposed to elevated TDG, but nothing is known about their risk for GBT. This project is the first to evaluate GBT in lamprey, beginning with larval and juvenile lamprey in a controlled laboratory setting. These early life stages were chosen for this initial work because they have been shown to be more sensitive to GBT in other fish species. We modified the FPC protocol for GBT exams to be specific to lamprey and ranked bubbles in the mouth, eyes (juveniles only), gill pores, first and second dorsal fins, caudal fin, anal fin, vent, and body. We followed the FPC ranking criteria and assigned rank based on the proportion of the area occluded with bubbles, as 0=no bubbles, 1=1-5%, 2=6-25%, 3=26-50%, and 4=>50%. Four experiments were completed with larval lamprey from January to September 2022 using small (70 mm total length or less) and large (86 mm total length or greater) larvae in approximately equal proportions. Experiments included: (1) 130% TDG for 31 d, (2) 125% TDG for 91 d, (3) 130% TDG for 20 d with assessments of burrowing performance, and (4) 128-138% TDG for 3-4 d with assessments of predator avoidance ability and the corresponding untreated control groups. The first and second experiments had similar study designs and findings. First, we tested an acute exposure at 130% TDG and then we tested a chronic exposure at 125%, to represent a full spill season. None of the controls (exposed to normally saturated water) experienced mortality or showed GBT signs. Few lamprey in the treatment groups (5% in Experiment 1; 0% in Experiment 2) showed GBT signs, and there were no mortalities (n=200 fish experiment 1; n=100 fish Experiment 2). Lamprey with GBT signs had bubbles on the body, with low severity ranks. During external exams for Experiment 2, we observed bubbles in the gut of several lamprey. The light coloration and transparency of the body made these observations possible, and we confirmed the finding with internal exams. From day 9 to day 91, 70.8% of the lamprey examined had bubbles in the gut. We observed five lamprey that were positively buoyant in the test tanks, and we likely underestimated the prevalence of floating as our procedures were not initially designed to document this sign. In our third experiment, burrowing performance was not significantly different between lamprey exposed to 130% TDG and controls. Mortality was 4.2% in the treatment group, but no GBT signs were observed. The proportion of lamprey with positive buoyancy increased through time, with 87.5% of fish floating on day 20 (end of the test). Bubbles in the gut were observed for some lamprey on each of five sampling dates (day 2 to 20), with prevalence ranging from 50-100%. Median burrow times ranged from 28 to 154 sec for treatment fish and from 40 to 100 sec for controls. We noted some atypical behaviors during burrow performance tests, including lamprey that were positively buoyant and unable to descend through 0.5 m of water to reach the sediment as well as lamprey that were unable to complete burrowing (within 10 min test period). These lamprey were so buoyant that they repeatedly floated to the surface of the water when they stopped or slowed their burrowing movements. Predator avoidance ability was assessed in our fourth experiment by exposing lamprey with GBT signs (floating) and controls to sculpin (Cottus spp.) until about 50% of the fish had been consumed or 2 h had passed. We completed five predation trials, testing the hypothesis that an equal proportion of treatment and control lamprey would be consumed. Treatment groups were generated by exposing 15 lamprey to 128-138% TDG for 3-4 d, until at least 10 lamprey were floating. Overall, 41 treatment and 46 control fish were eaten and there was no evidence that sculpin preferentially preyed upon lamprey with GBT signs. Additional tests with another predator are recommended. Two experiments were completed with juvenile lamprey from March to November 2022: Experiment 5 exposed fish to 125% TDG for 10 d and Experiment 6 exposed fish to 125% for 16 d. Mortality rates for the treatment groups were 21.7% and 20.0% for these experiments, respectively, and few lamprey (4 per experiment) showed GBT signs. With the results from experiments pooled, bubbles were observed in all body areas with low severity ranks (means 0-1). We observed some exophthalmia and bubbles behind the gill pores, in addition to bubbles in the gut and fish floating. The presence of bubbles in or near the gill pores was the likely cause of death as exam findings included enlarged gill pore areas and restricted openings. This project provided the first insights into lamprey responses to elevated TDG, but substantial learning opportunities remain. Our findings highlight that lamprey are vulnerable to GBT, but the effects are generally sublethal and would not be detected using FPC exam procedures. For example, we observed larval and juvenile lamprey that had bubbles in the gut and/or were floating, although these conditions were not consistently linked. More data are needed, but we surmise that it takes some time for bubbles to form in sufficient quantity to create the floatation required to overcome the mass of the lamprey. Positive buoyancy in natural settings could have substantial impacts to the risk of mortality for lamprey. Future studies could test GBT risk for larval lamprey in burrows, investigate the influence of lamprey size, measure performance (e.g., burrowing, swimming, predator avoidance ability) after elevated TDG exposure, and describe the rate that GBT signs dissipate when lamprey are returned to normally saturated water.