THE GUT
Osmotic, Ionic and Nitrogenous–Waste Balance
Contents
Osmoregulation in Fishes: An Introduction
Mechanisms of Gill Salt Secretion in Marine Teleosts
Mechanisms of Ion Transport in Freshwater Fishes
Water Balance and Aquaporin
Osmosensing
Osmoregulation in Fishes: An Introduction
DH Evans, University of Florida, Gainesville, FL, USA
ª 2011 Elsevier Inc. All rights reserved.
Introduction
Paleoecology of Fishes
Osmoregulation in Marine Fishes
Osmoregulation in Freshwater Fishes
Glossary
Diffusion Net movement of a solute from an area of
higher concentration to an area of lower concentration.
Glomerulus Tuft of capillaries surrounded by the
Bowman’s capsule of the vertebrate kidney where blood
is filtered to produce urine in the proximal tubule.
Hyperosmotic Containing a greater concentration of
solutes than another solution. If the solutions are
separated by a permeable membrane (such as gill
epithelium), solute will diffuse from a hyperosmotic to a
hyposmotic solution and water will move from the
hyposmotic to hyperosmotic solution.
Hyposmotic Containing a lesser concentration of
solutes than another solution.
Isosmotic Containing the same total solute
concentration as another solution. If the solutions are
separated by a permeable membrane, there will be no
Introduction
Osmoregulation is a fundamental process of living sys-
tems, equivalent in importance to respiration, digestion,
or reproduction. Osmoregulatory processes are those that
1348
The Origin of Ionic Uptake Systems in the Early
Vertebrates
Why Aren’t All Fishes Euryhaline?
Further Reading
net movement of either solute or water from one solution
to the other.
Osmoregulation The maintenance of
consistent cellular or organismal fluid composition and
volume.
Osmosis Essentially diffusion for water molecules. As
with diffusion for dissolved solutes, water moves due to
random thermal oscillations, in a net sense, from where
there is a higher water concentration (amount per unit
volume) to where there is a lower water concentration.
This is generally inversely related to the situation for
dissolved solutes, as higher levels of these occupy
space leaving less space for water and reducing its
concentration. Water therefore (again in a net sense)
moves from where the osmolality is lower to where it is
higher (the opposite of what occurs for the diffusion of
solutes).
enable a fish to maintain its cellular fluid composition and
volume. This is of critical importance because the pro­
tein-based processes of cells are sensitive to cytoplasmic
ionic concentrations, and cell membranes tolerate
 Osmotic, Ionic and Nitrogenous-Waste Balance | Osmoregulation in Fishes: An Introduction 1349

relatively small deviations in cell volume (they will either
collapse or explode). While terrestrial organisms are con­
tinually faced with the problem of dehydration, fishes are
surrounded by water, and the water can tend to either
dehydrate (in seawater, SW) or hydrate them (in fresh­
water, FW).
The differences in the ionic and total osmotic concen­
trations of the water and those of the fish body fluids
determine the magnitude and the direction of ion (diffu­
sion) and water (osmosis) movements across the fish gill
epithelium. Fishes can limit ion and water exchanges by
creating barriers at the skin surface (e.g., scales and mucus
layers) that limit permeability. However, they still possess
gills that are specialized for gas exchange (large surface
area, thin, highly vascularized), which remains an excel­
lent site for osmotic movement of ions and water. In fact,
when oxygen and carbon dioxide diffusion across the gills
increases during activity, ion and water diffusion increases
too. This creates a problem for a fish called the osmo­
respiratory compromise (see also Role of the Gills: The
Osmorespiratory Compromise). Therefore, post-exercise
fish not only have to rectify the depletion of oxygen stores,
but they must also deal with ions and water they may have
lost or gained as a result of breathing more! When terres­
trial animals exercise, they only lose water via their
respiratory surface.
With the exception of the hagfishes, the ionic concen­
trations of all fish plasma are intermediate between SW
and FW levels (Table 1). Consequently, excluding
hagfishes, all fishes must osmoregulate because they
usually live in either FW (hyposmotic to fish plasma) or
SW (hyperosmotic to fish plasma). Hagfishes are excep­
tional because their plasma has similar ionic and osmotic
concentrations as the SW environment (termed isosmotic),
and osmoregulation is not necessary. Animals that maintain
their blood plasma isosmotic with the environment are able
to save metabolic energy, because life in nonisosmotic
solutions requires compensatory transport of solutes and
water by cell membranes, epithelial tissues, or specific
organs and these processes are energy demanding
(variously estimated at 2–8% of resting metabolic rate).
The differences in the ionic (and total osmotic) con­
centrations between SW and FW are extreme (1000 times
in most cases), by comparison with the difference in
plasma ionic concentrations among fish (only �3 times).
Most fishes are adapted to either FW or SW, not both.
Fishes that are only able to osmoregulate over a very
narrow range of salinities are termed ‘stenohaline’, those
that can tolerate a wide range of salinities (e.g., both FW
and SW) are termed ‘euryhaline’. Freshwater fishes can
often tolerate salinities up to isosmolarity and marine
species can often tolerate salinities down to isosmolarity.
Euryhaline species can osmoregulate on both sides of the
isosmolarity point. Estuarine environments provide a full
range of salinities in which many fish live; isosmolar
habitats may exist here too. Very few fish species are
adapted to hypersaline conditions. (see also Intertidal
Fishes: Intertidal Habitats).

Table 1 Ionic compositions of plasma in fishes in SW and FW

Salinity or species Total osmolaritya Na+b Cl� K+ Ca2+ Mg2+ SO2�

4 Urea TMAO

Seawater 1050 439 513 9.3 9.6 50 26

Hagfish 1035 486 508 8.2 5.1 12 3
Anadromous lamprey 333 156 159 5.6 3.5 7.0
Euryhaline bull shark 1067 289 296 5.8 4.4 1.8 370 47
Dogfish shark 1118 255 241 6.0 5.0 3.0 0.5 441 72
Euryhaline Atlantic stingray 953 319 295 330
Euryhaline steelhead trout 325 153 135 4.0 1.4 0.65
Euryhaline puffer 363 179 144 2.8 3.1 1.8
Freshwater (soft) 1.0 0.25 0.23 0.01 0.07 0.04 0.05
Anadromous lamprey 112 99.6 2.3 1.8 1.5
Landlocked lamprey 272 120 104 3.9 2.5 2.0
Euryhaline bull shark 595 221 220 4.2 3.0 1.3 151 19
Euryhaline Atlantic stingray 621 212 209 196
Freshwater stingray 319 178 146 1.22
Gar 159 133 4.2 6.1 0.3
Bowfin 279 133 110 1.5
Carp 274 130 125 2.9 2.1 1.2
Euryhaline steelhead trout 260 153 133 3.8 1.4 0.5
Euryhaline puffer 346 166 128 3.0 3.5 1.3
a
mOsm.l�1.

b
mM.l�1 or mM.kg�1.

Modified from Evans DH and Claiborne JB (2009) Osmotic and ionic regulation in fishes. In: Evans DH (ed.) Osmotic and Ionic Regulation: Cells and

Animals, pp. 295–366. Boca Raton, FL: CRC Press.

1350 Osmotic, Ionic and Nitrogenous-Waste Balance | Osmoregulation in Fishes: An Introduction
Over 95% of the total solute concentration (32–35
parts per thousand, ‰) of SW is due to the salts listed
in Table 1, but over 80 other elements and ions are
present in measurable quantities. The same elements
predominate in FW but at much lower concentrations
and more variable compositions because of differences
in input from the terrestrial substrates (Table 1). One of
the more important variables in FW is the calcium con­
centration, which, although lower than that in SW, may
affect the permeabilities (salt and water) of biological
membranes, thereby indirectly influencing osmoregula­
tion (see below).
Paleoecology of Fishes
Much can be learned about the possible evolutionary
course of osmoregulatory strategies and mechanisms by
comparing the plasma ionic compositions and habitats
among the major phylogenetic fish groups. The finding
that the plasma ionic concentration of all fishes (except
for the hagfishes) is 30–40% that of SW has intrigued
physiologists for over a century. Based upon the fossil
record and the fact that the most primitive bony fishes
(sturgeons, gar and bowfin, and bichirs) are FW fishes,
and the fact that euryhaline marine species reduce their
plasma salt concentrations as they enter lower salinities, it
is generally agreed that ancestral fish species entered FW
over 500 million year ago from a marine origin.
Subsequent evolution of modern marine fishes (both
bony and cartilaginous), as well as tetrapods (amphibians,
reptiles, birds, and mammals), took place after this FW
invasion. Extant lampreys, which are either FW or eur­
yhaline migrants (FW to SW to FW) as they mature and
reproduce (see also Hagfishes and Lamprey: Lampreys:
Energetics and Development and Toxicology: The
Effects of Toxicants on Olfaction in Fishes), are probably
modern descendents of these ancestral fish species. The
stenohaline, marine hagfishes are thought to be remnants
of the ancestral, marine pre-vertebrates, largely because,
Seawater
Na+, Cl−
H2O
Ingest
Filtration
SW
+ −
Na .Cl .H2O
Na+, Cl−
Figure 1 General pattern for SW teleost osmoregulation. From Evans DH (2008) Teleost fish osmoregulation: What have we learned
since August Krogh, Homer Smith, and Ancel Keys? American Journal of Physiology – Regulatory, Integrative and Comparative
Physiology 295: R704–R713.
like marine invertebrates and unlike any other chordates,
they are ionically isosmotic to SW.
Osmoregulation in Marine Fishes
Hagfishes
Although hagfish plasma is isosmotic to SW, there are
ionic differences, most notably greater Na+ but lesser
Cl�, Mg2+, and SO42– concentrations (Table 1). The
mechanisms maintaining these ionic gradients are largely
unknown, although the copious slime production by
hagfishes may be an excretory pathway for at least the
divalent ions (see also Hagfishes and Lamprey: Hagfish
and The Skin: Hagfish Slime). The slightly greater
plasma Na+, and lesser Cl� concentration may be sec­
ondary to the presence of gill Na+/H+ and Cl�/HCO3�
exchangers used in net acid secretion for acid–base
regulation or because of the charge distribution second­
ary to plasma organics or excretion of unwanted divalent
ions (e.g., Ca2+ or Mg2+), but these propositions are
largely untested.
Teleosts
Marine teleosts face dehydration and net ionic loading
because they are hyposmotic to SW (Table 1). In the
1930s, two pioneers in fish osmoregulation (Homer Smith
and Ancel Keys) demonstrated that teleosts ingest SW to
offset the osmotic loss of water and excrete the excess
NaCl, using the gills, rather than the kidneys (as mam­
mals and birds do). Specialized ‘chloride’ cells (now
termed mitochondrion-rich cells, MRCs) were described
in the marine fish gill by Keys, and substantial evidence
now exists that MRCs are the site of NaCl extrusion by
the marine teleost gill epithelium (see also Osmotic,
Ionic and Nitrogenous-Waste Balance: Mechanisms of
Gill Salt Secretion in Marine Teleosts). The ingested
divalent ions (Ca2+, Mg2+, and SO42–) are excreted by
the intestine and kidneys (Figure 1). In fact, the ingested
1000 mOsm
Na+.Cl−.H2O
400 mOsm
Low Vol
Isotonic Na+,Cl−
 Osmotic, Ionic and Nitrogenous-Waste Balance | Osmoregulation in Fishes: An Introduction
Ca2+ is precipitated out as a bicarbonate salt, which is
excreted and actually appears to play a significant role in
marine carbon cycling (see also Role of the Gut: Gut
Ion, Osmotic and Acid-Base Regulation). The urine flows
of marine teleosts are quite low, and there are some
marine teleosts that have secondarily ‘lost’ the blood-
filtering renal glomerulus and produce urine by ionic
secretion in the proximal tubule (see also Role of the
Kidneys: The Kidney). Indeed, it was investigations of
these aglomerular species that provided direct evidence
for proximal tubular ionic secretion in the vertebrate
kidney (including mammals), which was controversial
until the aglomerular teleost species were discovered.
Elasmobranchs
The marine sharks, skates, and rays have evolved very
different mechanisms of osmoregulation, and it is not
clear why there is such disparity between the teleost and
elasmobranch mode of marine osmoregulation. This is a
good case of ‘divergent evolution’. Elasmobranch plasma
contains exceedingly high concentrations of the organic
solutes urea and trimethylamine oxide (TMAO;
Table 1). Indeed, the urea concentration in these fishes
is 5–10 times higher than coma-producing levels in mam­
mals. Importantly, it is the relatively high TMAO
concentrations that counteract the toxic effects of the
extraordinarily high urea concentrations in elasmo­
branchs (see also Responses and Adaptations to the
Environment: General Principles of Biochemical
Adaptations).
Elasmobranchs do not ingest appreciable volumes of
SW because they are slightly hyperosmotic to SW, so
there is no osmotic loss of water across the gills. In fact,
the osmotic uptake of water is balanced by proportional
urine excretion in elasmobranchs, produced by glomerular
filtration and, relatively unstudied, tubular secretory and
reabsorptive processes in the exceedingly complex elasmo­
branch renal nephrons. There are also ionic gradients
producing a net, diffusional gain of NaCl, which is offset
by extra-renal and extra-branchial salt secretion by a
specialized rectal gland that secretes a hypersaline
solution into the cloaca (see also Role of the Gut:
Dogfish Rectal Gland). The rectal gland has extremely
high concentrations of the ubiquitous ion transport
enzyme, Na+/K+-activated ATPase, which plays a cen­
tral role in the mechanisms of NaCl secretion by this
gland, as well as by the teleost gill epithelium (see also
Osmotic, Ionic and Nitrogenous-Waste Balance:
Mechanisms of Gill Salt Secretion in Marine Teleosts).
In fact, the transporting cell in the shark rectal gland
looks very much like the teleost gill MRC. MRCs are
also found in the elasmobranch gill epithelium, but there
is no evidence that they function in net salt secretion. It
is apparent that the elasmobranch gill MRC function in
1351
acid–base regulation by secreting variable amounts of
H+ versus HCO� 3.
Lampreys
Lampreys are anadromous species that breed in FW; the
resulting larval forms (ammocoetes) metamorphose after a
few years into juveniles that migrate downstream and live
most of their adult life in the marine environment (see
also Hagfishes and Lamprey: Lampreys: Energetics and
Development). Because the marine adults are so rarely
captured, and the upstream migrating, premating adults
have lost the ability to osmoregulate in SW, little is known
about the mechanisms for osmoregulation in SW lampreys
(see also Hagfishes and Lamprey: Lampreys:
Environmental Physiology). Their plasma is hyposmotic
to SW (Table 1), so studies have found that, like teleosts,
they ingest SW, and presumably excrete the excess NaCl
across the gill, because the kidney cannot produce urine
that has greater NaCl concentrations than the plasma. The
branchial epithelium has MRCs, but no functional or
molecular studies have been published to support the
logical suggestion that the pathways and mechanisms for
gill salt extrusion are the same as have been well docu­
mented in teleosts.
Osmoregulation in Freshwater Fishes
Teleosts
Freshwater teleosts are hyperosmotic to their environ­
ment (Table 1) and therefore face a net influx of water
and loss of NaCl across their branchial epithelium. They
balance the osmotic influx by producing a much larger
volume of urine than their marine relatives, with a much
lesser ionic concentrations to conserve salt. Nevertheless,
renal loss of salt adds to the diffusional loss, and must be
offset by some uptake mechanism (Figure 2). Some salt
may be gained by ingestion of food, but this has rarely
been quantified and would limit osmoregulation during
periods of low food availability. In the 1930s, August
Krogh provided direct evidence for the independent
uptake of Na+ and Cl� by a variety of freshwater animals,
including fishes. He also proposed that, in order to avoid
bioelectrical problems, it was likely that the uptake of
each ion was in exchange for a counter-ion that was
relatively abundant in the plasma: H+ (or NH+4 ) and
HCO� 3 , respectively. Subsequent studies have confirmed
this proposition, although the actual cell and pathways are
still under debate, and may be species specific. It is prob­
able that MRCs are involved, and most species that have
been studied possess MRC subtypes that may have sepa­
rate functions (Osmotic, Ionic and Nitrogenous-Waste
Balance: Mechanisms of Ion Transport in Freshwater
Fishes). The current debate centers on whether the
1352 Osmotic, Ionic and Nitrogenous-Waste Balance | Osmoregulation in Fishes: An Introduction
Freshwater
Na+, Cl−
H2O
Filtration
Na+, Cl−
Figure 2 FW teleost osmoregulation.. From Evans DH (2008) Teleost fish osmoregulation: What have we learned since August Krogh,
Homer Smith, and Ancel Keys? American Journal of Physiology – Regulatory, Integrative and Comparative Physiology 295: R704–R713.
Na+/H+ exchange is via an electroneutral chemical link­
age (e.g., via a class of proteins termed Na-H exchangers
(NHEs), which mediate this exchange in other tissues) or
via an electrogenic proton pump (H-ATPase) which pro­
duces a cytoplasmic negativity that draws in Na+ from the
freshwater via a Na+ channel. There is also emerging
evidence that a coupled transport of Na and Cl may
play some role in this uptake process.
Elasmobranchs
Freshwater elasmobranchs are rare; the most notable
example is the euryhaline bull shark (Carcharhinus leucus)
that can migrate into freshwater in Central America,
Australia, and India (and even Illinois, where one was
captured in the Mississippi River 3800 km from the Gulf
of Mexico). Another example is the Atlantic stingray
(Dasyatis sabina) that has a resident population in lakes
in Central Florida. The only truly freshwater stingray,
Potamotrygon sp., is found in tributaries of the Amazon
and Orinoco Rivers in South America. The euryhaline
species all display reduced plasma NaCl and urea levels
when captured in lowered salinities, and Potamotrygon
plasma contains nearly no urea (Table 1). Indeed,
Potamotrygon can neither increase the plasma urea con­
centrations nor tolerate substantial increases in salinity.
The rates of urea synthesis are much lower than in
marine elasmobranchs, and the inability to increase
plasma urea concentrations may also be associated with
the loss of specific tubules in the kidney. These tubules
have been implicated in renal reabsorption of urea
filtered from the plasma. As might be expected, the rectal
gland is much reduced in freshwater elasmobranchs, and
the osmotic uptake of water is balanced by excretion of
extremely high volumes of dilute urine. The diffusional,
renal, and small rectal gland loss of salts must be
balanced by branchial uptake. At least in D. sabina, the
extraction of NaCl from the hyposmotic environment is
1 mOsm
Na+, Cl−
300 mOsm
High vol
Low Na+, Cl−
mediated by two cells that have the apical proteins
necessary for independent Na+ and Cl� uptake.
Lampreys
Larval, juvenile, and adult lampreys in freshwater show
much reduced plasma ionic concentrations compared to
the marine adults (Table 1). The osmotic uptake of water
is balanced by production of large volumes of very dilute
urine. The renal and diffusional loss of ions is offset by
active branchial epithelial uptake via what appears to be
two types of freshwater MRCs that disappear as the
lamprey migrates into SW and reappear when the adults
return to FW. These cells are thought to extract needed
NaCl via a combination of the ionic transporters pro­
posed for freshwater teleosts (see also Osmotic, Ionic
and Nitrogenous-Waste Balance: Mechanisms of Ion
Transport in Freshwater Fishes, Cellular, Molecular,
Genomics, and Biomedical Approaches: Physiology of
Triploid Fish; Mechanisms of Ion Transport in
Freshwater Fishes; Hagfishes and Lamprey: Lampreys:
Environmental Physiology).
The Origin of Ionic Uptake Systems in the
Early Vertebrates
Despite the uncertainty of the specific transport pathways
that mediate NaCl uptake in freshwater fishes, it is clear that
the evolution of this system was critical for the entry of
primitive marine fishes into the freshwater environment
nearly 500 million years ago. Where did it come from?
Unlike mammals, fishes cannot substantially alter their
blood pH by changes in renal secretion of H+ or HCO� 3
or changes in respiratory excretion of CO2. Thus, it has
been found that acid–base regulation in fishes is mediated
by differential excretion of H+ or HCO�3 by the gill epithe­
lium (see also Role of the Gut: Gut Ion, Osmotic and Acid-
Base Regulation). The demonstration that marine teleosts,
 Osmotic, Ionic and Nitrogenous-Waste Balance | Osmoregulation in Fishes: An Introduction
elasmobranchs, and even hagfishes use these gill transport
mechanisms for acid–base regulation led to the proposition
that the evolution of ionic exchange systems that could
excrete blood H+ and HCO� 3 in exchange for external
Na+ and Cl� predated the evolutionary entrance into fresh­
water. Thus, the origin of ionic exchange mechanisms for
acid–base regulation presumably afforded a pre-adaptation
for ionic regulation in reduced salinities.
Why Aren’t All Fishes Euryhaline?
If the gill epithelium of all fishes, including hagfishes,
possess the ionic exchange systems (for acid–base regula­
tion) that potentially can be used for ionic uptake in
freshwater, why aren’t all fishes euryhaline? One might
propose that intestinal and renal functions are limiting,
but these are usually quantitative rather than qualitative
differences. Presence of a putative ionic uptake system is
qualitative; without it, freshwater osmoregulation is not
possible. It appears that the limiting factor for euryhali­
nity is the efficiency of the ionic uptake systems versus
the ionic permeability of the branchial epithelium itself.
The best example of this relationship is the finding, nearly
80 years ago, that normally stenohaline marine reef fishes
(e.g., angel fish, puffer, gray snapper, and horse-jack) are
found in freshwater lakes in the Bahamas and springs in
western Florida (e.g., Homosassa Springs). Because these
water bodies have relatively high concentrations of Ca2+
(studies have shown that it can decrease the passive loss of
ions from fishes), it has been suggested that one of the
primary limiting factors for invasion into freshwater is the
water calcium concentrations and the overall permeabil­
ity of the gill.
See also: Hagfishes and Lamprey: Hagfish; Lampreys:
Energetics and Development; Lampreys: Environmental
Physiology. Osmotic, Ionic and Nitrogenous-Waste
Balance: Mechanisms of Ion Transport in Freshwater
Fishes; Mechanisms of Gill Salt Secretion in Marine
Teleosts. Responses and Adaptations to the
Environment: General Principles of Biochemical
Adaptations. Role of the Gills: The Osmorespiratory
1353
Compromise. Role of the Gut: Dogfish Rectal Gland; Gut
Ion, Osmotic and Acid-Base Regulation. Role of the
Kidneys: The Kidney. The Skin: Hagfish Slime.
Toxicology: The Effects of Toxicants on Olfaction in
Fishes.
Further Reading
Bartels H and Potter IC (2004) Cellular composition and ultrastructure of
the gill epithelium of larval and adult lampreys: Implications for
osmoregulation in fresh and seawater. Journal of Experimental
Biology 207: 3447–3462.
Beyenbach KW (2004) Kidneys sans glomeruli. American Journal of
Physiology, Renal Physiology 286: F811–F827.
Carrier JC and Evans DH (1976) The role of environmental calcium in
freshwater survival of the marine teleost, Lagodon rhomboides.
Journal of Experimental Biology 65: 529–538.
Claiborne JB, Edwards SL, and Morrison-Shetlar AI (2002) Acid–base
regulation in fishes: Cellular and molecular mechanisms. Journal of
Experimental Zoology 293: 302–319.
Evans DH (1984) Gill Na/H and Cl/HCO3 exchange systems evolved
before the vertebrates entered fresh water. Journal of Experimental
Biology 113: 464–470.
Evans DH (2008) Teleost fish osmoregulation: What have we learned
since August Krogh, Homer Smith, and Ancel Keys? American
Journal of Physiology – Regulatory, Integrative and Comparative
Physiology 295: R704–R713.
Evans DH and Claiborne JB (2009) Osmotic and ionic regulation in
fishes. In: Evans DH (ed.) Osmotic and Ionic Regulation: Cells and
Animals, pp. 295–366. Boca Raton, FL: CRC Press.
Evans DH, Piermarini PM, and Choe KP (2004)
Homeostasis: Osmoregulation, pH regulation, and nitrogen
excretion. In: Carrier J, Musick J, and Heithaus J (eds.) Biology of
Sharks and Their Relatives, pp. 447–475. Boca Raton, FL: CRC
Press.
Evans DH, Piermarini PM, and Choe KP (2005) The multifunctional fish
gill: Dominant site of gas exchange, osmoregulation, acid–base
regulation, and excretion of nitrogenous waste. Physiological
Reviews 85: 97–177.
Marshall WS and Grosell M (2006) Ion transport, osmoregulation and
acid–base balance. In: Evans DH and Claiborne JB (eds.) The
Physiology of Fishes, pp. 177–230. Boca Raton, FL: CRC Press.
Olson KR (1999) Rectal gland and volume homeostasis. In: Hamlett WC
(ed.) Sharks, Skates, and Rays, pp. 329–352. Baltimore, MD: Johns
Hopkins University Press.
Parks SK, Tresguerres M, and Goss GG (2007) Blood and gill responses
to HCl infusions in the Pacific hagfish (Eptatretus stoutii). Canadian
Journal of Zoology 85: 855–862.
Silva P, Solomon RJ, and Epstein FH (1996) The rectal gland of Squalus
acanthias: A model for the transport of chloride. Kidney International
49: 1552–1556.
Wilson RW, Millero FJ, Taylor JR, et al. (2009) Contribution of fish to
marine inorganic carbon cycle. Science 323: 359–362.