title: Adaptations of a deep sea scavenger: high ammonia tolerance and active NH₄⁺ excretion by the Pacific hagfish (Eptatretus stoutii).
abstract: The Pacific hagfish (Eptatretus stoutii) has an exceptional ability to both withstand and recover from exposure to high external ammonia (HEA). This tolerance is likely due to the feeding behavior of this scavenger, which feeds on intermittent food falls of carrion (e.g. fish, large marine mammals) during which time it may be exposed to high concentrations of total ammonia (T(Amm)=NH3+NH4(+)) while burrowed inside the decomposing carcass. Here we exposed hagfish to 20 mmol L(-1) T(Amm) for periods of up to 48 h and then let animals recover in ammonia-free seawater. During the 48 h HEA exposure period, plasma T(Amm) increased 100-fold to over 5000 μmol L(-1) while ammonia excretion (J(amm)) was transiently inhibited. This increase in plasma T(Amm) resulted from NH3 influx down massive inwardly directed ΔP(NH3) gradients, which also led to a short-lived metabolic alkalosis. Plasma [T(Amm)] stabilized after 24-48 h, possibly through a reduction in NH3 permeability across the body surface, which lowered NH3 influx. Ammonia balance was subsequently maintained through the re-establishment of J(amm) against an inwardly directed ΔP(NH3). Calculations of the Nernst potential for ammonia strongly indicated that J(amm) was also taking place against a large inwardly directed NH4(+) electrochemical gradient. Recovery from HEA in ammonia-free water was characterized by a large ammonia washout, and the restoration of plasma TAmm concentrations to near control concentrations. Ammonia clearance was also accompanied by a residual metabolic acidosis, which likely offset the ammonia-induced metabolic alkalosis seen in the early stages of HEA exposure. We conclude that restoration of J(amm) by the Pacific hagfish during ammonia exposure likely involves secondary active transport of NH4(+), possibly mediated by Na(+)/NH4(+) (H(+)) exchange.
authors.affiliation: Department of Biological Sciences, University of Alberta, Edmonton, Alberta, T6G 2E9, Canada; Bamfield Marine Sciences Centre, Bamfield, British Columbia, V0R 1B0, Canada. Electronic address: alex.clifford@ualberta.ca.
authors.forename: Alexander M
authors.initials: AM
authors.lastname: Clifford
authors.affiliation: Department of Biological Sciences, University of Alberta, Edmonton, Alberta, T6G 2E9, Canada; Bamfield Marine Sciences Centre, Bamfield, British Columbia, V0R 1B0, Canada.
authors.forename: Greg G
authors.initials: GG
authors.lastname: Goss
authors.affiliation: Bamfield Marine Sciences Centre, Bamfield, British Columbia, V0R 1B0, Canada; Biology Department, Wilfrid Laurier University, Waterloo, Ontario, N2L 3C5, Canada.
authors.forename: Michael P
authors.initials: MP
authors.lastname: Wilkie
mesh_terms: D000136:Acid-Base Equilibrium; D000222:Adaptation, Physiological; D000641:Ammonia / Q000097:blood / Q000378:metabolism* / Q000494:pharmacology; D000818:Animals; D005880:Gills / Q000378:metabolism; D005973:Glutamine / Q000097:blood; D006196:Hagfishes / Q000187:drug effects / Q000502:physiology*; D012623:Seawater
chemical_list: D005973:Glutamine; D000641:Ammonia
keywords: Acid-base balance; Active transport; Agnatha; Ammonia; Gill; Ion regulation; Nernst potential; Nitrogen; Rh glycoprotein; Urea