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Elasmobranch sh typically maintain body uids slightly hyperosmotic
to the surrounding environment. This is achieved by regulating the concentra-
tions of organic and inorganic osmolytes, principally sodium, chloride, urea,
and trimethylamine oxide. The concentrations of these osmolytes are altered,
independently of each other, in response to changes in environmental salin -
ity. These alterations are necessarily coupled with changes in the direction
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