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Teleost
fish encounter very different osmotic challenges depending on the
salinity of their external environment. In freshwater, fish face osmotic
gain of water and compensate by excreting copious amounts of dilute
urine. However, this strategy involves a loss of vital salts, which
are replaced via ingested food and active uptake at the ‘chloride
cells’ of the gill epithelia. In seawater, fish face osmotic
water loss, coupled with salt gain. Drinking seawater helps to replace
lost water, but this also causes the animal to take in large quantities
of salt. This excess salt is excreted using specialised 'chloride
cells' on the gill epithelium. Seawater adapted fish also produce
small amounts of highly concentrated urine in order to conserve water.
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Euryhaline teleost fish face different osmotic
challenges as they migrate between freshwater and seawater environments.
These fish therefore have to switch strategy, depending on their
external environment. There are two principle types of euryhaline
teleost fish:
Anadromous teleost fish such as Atlantic salmon
(Salmo salar) and sea trout (Salmo trutta) spawn
in freshwater and mature as adults in seawater. Catadromous teleosts
fish such as the European eel, Anguilla anguilla, spawn
in seawater and return to freshwater to mature. Euryhaline teleost
fish must integrate the functions of gill, gut and kidney in order
to adapt successfully to the dramatically different environments
they face as part of their natural life cycle.
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Gill: In freshwater, ‘chloride cells’ remove
divalent ions such as bicarbonate and ammonium, and the Na+K+ATPase
pump on the basolateral membrane works to create an ionic gradient
favoring the movement of Na+ and Cl- into the cell from the external
environment. These chloride cells are primarily found at the base
of the secondary lamellae of the gills and are the principle site
for ionic exchange in euryhaline teleosts. In seawater-adapted fish,
the chloride cells are larger, and function principally to remove
excess Na+ and Cl- ions. Consequently, the Na+K+ATPase pump works
to create a concentration gradient favoring the movement of these
ions into the external environment. Recent research by FERG has demonstrated
increased molecular expression of Na+K+ATPase in long-term seawater
adapted eels. |
Gut: As euryhaline teleost fish adapt to seawater, they must compensate
for the water lost across the gills. This is achieved by drinking
the external medium. One of the principal roles of the peptide hormone,
Angiotensin II, is the control of drinking rate. Angiotensin II is
the biologically active component of the renin angiotensin system
(RAS). In mammals, renin, produced in the juxtaglomerular apparatus
in the kidney, acts on a protein substrate angiotensinogen, produced
in the liver, to form the decapeptide angiotensin I (Ang I). Angiotensin
converting enzyme (ACE) then cleaves Ang I to form the octapeptide
and biologically active component angiotensin II (Ang II). Further
roles for the RAS include, vasopressor effects (constriction of blood
vessels), anti-diuretic effects (reduction in urine production) and
anti-natriuretic effects (reabsorption of sodium).
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The presence of a renin angiotensin
system in teleosts was detected as early as 1942, and confirmed
in the early 1960's. However, it was not until recently that the
biological activity of homologous Ang II was established in teleosts.
Through pharmacological manipulation of the RAS, FERG
has demonstrated some of the key functions of the RAS in teleosts,
including cardiovascular and osmoregulatory roles, using the European
Eel, Anguilla anguilla, as a model. |

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Movement between different environmental
salinities is an energy demanding process which involves significant
physiological stress to the animal. The stress of movement between
freshwater and seawater may result in the animal being more susceptible
to parasitic infection or disease. We have examined these effects
in a range of teleost species using a number of laboratory techniques.
These include the determination of plasma levels of a number of
stress hormones by radioimmunoassay (RIA), including cortisol, ACTH
and MSH. Stress and metabolic effects may also be determined by
examination of the plasma concentrations of glucose and lactate,
and measurement of liver glycogen levels by standard spectrophotometric
and analytical techniques.
Complete adaptation to freshwater or seawater involves
an integration of physiological and molecular processes. Drinking
rate is increased in the seawater environment to compensate for
osmotic water loss and there is a concurrent increase in the expression
of key molecular transporters, such as Na+K+ATPase, to remove excess
sodium and chloride ions.
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