Nitrogenous waste excretion in aquatic species is accomplished
largely at the gills. Because these species have very large
convective water volume requirements in order to extract oxygen
(owing to the low concentration of oxygen in water relative to air),
aquatic species are typically ‘hyperventilated’ with respect to waste
gases such as CO2 and NH3. Thus, in many cases no additional
ventilatory energy needs to be invested by water breathers to
effectively excrete nitrogenous waste to the water, beyond that used
to take up oxygen from the water. This concept was developed early
in the literature. When coupled with another early (but largely
incorrect) notion that molecules like NH3 and urea readily passed
through the lipid portion of the plasma membrane, initial views of
excretion of nitrogenous waste in aquatic organisms held it to be a
largely passive process. [As ammonia can exist as a dissolved
ammonia gas (NH3) and the ammonium ion (NH4
+), this review
will use the convention of the term ‘ammonia’ when the chemical
is not specified, and either the chemical symbols or ‘ammonia gas’
or ‘ammonium ion’ when the chemical form is to be specified.]
Early research also focused on the ionic form of ammonia, i.e. the
ammonium ion, examining the possibility that it could be a
surrogate substrate for transporters viewed as dealing primarily
with hydrogen, potassium or sodium ions, representing an
alternative or parallel pathway for ammonia excretion. Indeed, for
many years much experimentation and debate, regarding the fish
gill in particular, centered on which was more important, NH3 or
NH4
+ excretion. Since most fish were not thought to excrete very
much waste as urea, very little research into urea transport was
undertaken; one notable exception was research to understand how
urea was retained by elasmobranch kidneys and gills in the face of
the massive outwardly directed gradient (e.g. ~400mmoll–1 as a
typical value).