Endocrine Control of Osmoregulation in Teleost

Fish1
As the primary link between environmental change and physiological response,
the neuroendocrine system is a critical part of osmoregulatory adaptations.
Cortisol has been viewed as the seawater-adapting hormone in fish and
prolactin as the fresh water adapting hormone. Recent evidence indicates that
the growth hormone/insulin-like growth factor I axis is also important in
seawater adaptation in several teleosts of widely differing evolutionary lineages.
In salmonids, growth hormone acts in synergy with cortisol to increase seawater
tolerance, at least partly through the upregulation of gill cortisol receptors.
Cortisol under some conditions may promote ion uptake and interacts with
prolactin during acclimation to fresh water. The osmoregulatory actions of
growth hormone and prolactin are antagonistic. In some species, thyroid
hormones support the action of growth hormone and cortisol in promoting
seawater acclimation. Although a broad generalization that holds for all teleosts
is unlikely, our current understanding indicates that growth hormone promotes
acclimation to seawater, prolactin promotes acclimation to fresh water, and
cortisol interacts with both of these hormones thus having a dual
osmoregulatory function.
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The capacity to regulate plasma ions in the face of changing external salinity is
an obvious necessity for fish that live in estuaries or that move between fresh
water and seawater as part of their normal life cycle. The need to respond to
salinity change may be rapid, such as during tidal cycles or rapid movements
through estuaries, or slow, such as in the seasonal or ontogenetic acquisition of
salinity tolerance in anadromous fish. The former requires the rapid activation
of existing mechanisms (transport proteins and epithelia), whereas the second
requires the differentiation of transport epithelia and synthesis of new transport
proteins. As the primary link between environmental change and physiological
response, the neuroendocrine system is a critical part of these osmoregulatory
adaptations. In this paper I will review recent evidence for the hormones
involved in development and differentiation of transport epithelia that control
the ability of teleost fish to move between fresh water and seawater. Our
previous textbook view of the endocrinology of osmoregulation has been that
cortisol is the seawater-adapting hormone and prolactin is the fresh wateradapting hormone, clearly defined for the first time by Utida et al. (1972).

Evidence collected in the last 15 yr indicates that the growth hormone/insulinlike growth factor I axis is also important in the seawater acclimation process
of teleosts. Recent findings on the importance of cortisol in ion uptake will also
be presented, and these indicate that cortisol has a dual osmoregulatory function
in many teleosts.
Osmoregulation in fish
Because of their environment, osmoregulation in fish presents specific
problems. Fish have adaptations that enable them to deal with these problems.
Those which live in fresh water or sea water have different problems.
Freshwater fish
Problems:

These fish are hypertonic to their surroundings. This means their blood
has a lower water concentration than the surrounding fresh water.

As fresh water passes through the mouth and over the gill membranes,
watermolecules diffuse from the fresh water into the blood by osmosis.

These fish must produce a very large volume of urine to balance this large
intake of water.

This large volume of urine carries salt with it, and the salt has to be
replaced.
Solutions:

To produce a large volume of urine the fish must remove a large volume
of water from the blood by having a high rate of filtration into
the kidneytubules.

This is done by having a kidney with many large glomeruli - capillary

networks from which fluid is filtered at the start of the kidney tubules.

Salt replacement is solved by chloride secretory cells in the gills, which

actively transport salts from the surrounding water into the blood.
Saltwater fish

These fish are hypotonic to their surroundings. This means their blood has
a higher water concentration than the surrounding sea water.

As sea water passes through the mouth and over the gill membranes,
water molecules diffuse out of the blood into the sea water by osmosis.

These fish must replace the water which they constantly lose by osmosis

They can also only afford to produce a very small volume of urine.
Drinking sea water brings a large quantity of salt into the blood and this
has to be removed.

Solutions:

To replace the water they lose, saltwater fish drink sea water.
To produce a small a volume of urine they must have a low rate of
filtration of water into the kidney tubules.

This is done by having a kidney with relatively few small glomeruli.

Salt is removed by chloride secretory cells in the gills, which actively

transport salts from the blood into the surrounding water.

Osmoregulation in salmon

Salmon begin their lives in rivers and migrate to the sea, returning to the same
rivers later in their lives. They are able to cope with these changes by altering
the way they osmoregulate.

Water balance in plants

Transpiration stream: First Stage

Water taken up by the roots of a plant is transported through a plant to the

leaves and lost into the air. The stages of the process are:

Water enters root hair cells by osmosis.

The root hair cell is hypertonic to the surrounding soil water. This means
that it has a lower water molecule concentration.

Water then moves from cell to cell through the root cortex by osmosis
along a concentration gradient; this means that each cell is hypertonic to the one
before it.

In the centre of the root the water enters the xylem vessels.

Water may move by diffusion through the cell walls and intercellular
spaces.
Second Stage

In the leaves, water molecules leave the xylem vessels and move from cell to
cell. They move through the spongy mesophyll layer by osmosis along a
concentration gradient. Water then evaporates into spaces behind
the stomataand diffuses through the stomata into the surrounding air.
Water rises from the roots to the leaves through the xylem vessels because of
two properties of water molecules:

Adhesion
Water rises in the narrow vessels partly because water molecules are attracted to
the walls of the vessels.

Cohesion
Water molecules are attracted to each other, and as water evaporates from the
leaves columns of water are drawn up through the xylem vessels.
The loss of water from the leaves of a plant is called transpiration, and the
resulting flow of water through the plant is called the transpiration stream.
The transpiration stream is important because:

it carries water for photosynthesis to the palisade cells in the leaves

the water carries essential mineral salts in solution
evaporation from the leaves has a cooling effect