1.

Main extracellular cation contributes major osmotic pressure of extracellular

fluidconcrentation gradient set up by sodium pump provedies potential energy for transport of
substances across cell membrane carries inward current for membrane excitation.
2.

Main cytoplasma cation Contributes to osmotic pressure of cytoplasm

establishes the testing potential Activites some enzymes carries outword current for membrane
repolarizatinon.
3.

Low concrentation in cell stabilizes membranes regulates exccytosis and muecle

contraction involved in comenting cells together carries inward current in some meregulates
many enzymes and processes : acts as Second messenger or coupling agen.
4.

intra-and extracellular stabilizes membranes antagonist of calcium Action in many

functions acts as cofactor for many enzymes (e.g myosin ATPase).
5.

inra-and extracellular Buffer

concrentation
6.

7.

Main extracellular anion of tissues counterion for inorganic cations.

12-1 Regulatory systems act as beffers between the external and the internal Environments of the
more advanced groups of animals. The cells and tissues of these animals are protected from large
fluctuations in the external Environment. Such as osmotic extremes. Since the compositon of the
internal extracellular fluids is maintained within Narrow limit.
Table 12-2 elextrolyte composition of the human body fluida.

Note : some of the ions contained within cells are not all freely dissolved in the cytoplsm. But may also
be squestered within cytoplasmic organelles. Thus the true free

concrentation in the cytoplasm is

typically below

meq/kg

O rather than the overall value given in the table. Faiture of anion and
cation total to agree reflects incomplete tabulation.

Have evplved circulatory systems, the blood typically passes through excretory organs, generally termed
kidneys. In terrestial animals , the kidneys not only play an important role in the removal of organic
wastes buta re the primary organs of osmoregulations. In fishes and equatic invertebrates, a major
portion of the osmoregulatory funtions are carried out by other organs, such as the gills, the
integument, and even the intestines.
The ability to maintain a suitable internal Environment in the face of osmotic strees has played a
most important rple in animal evolution. There are two main reasons. First, animals are resticted in their
geographic distribution by environmental factors, one of the most important being the osmotic nature
of the Environment. Second, geographic dispersal followed by genetic isolation is an impoetant
mechanism for the divergence of spesies in the process of evolution. If, for example the arthropods and
the vertebrates had not evolved means of regulating their extracellular compartments, they would have
been far less succesful in their invasion of the osmotically hostile freshwater and terrestial Environment.
As a consequence, the number and diversity of arthrophod and vertebrata spesies would now be
Limited largely to those found in the oceans and fresh Water of the wold, which support far less than
halfthe living spesies of animals. In the absence of terrestrial arthropods and vertebrates, other groups
would have evolved with greater diversity to fill the vacant terrestrial niches.
Thus,the animal groups that have been most successful in colonizing extreme osmotic
Environment have evolved means of regulating their water content and the concentration of solutes
dissolved in their body water.There is always a difference between the optimal intracellular and
extracelluler concentrations of ions.A number of mechanisms are employed to handle osmotic problems
and regulate the differences(1)between intracellular and extracellular compartments and(2)between
the extracellular compartment and the external Environment.these are collectively termed
osmoregulatory mechanisms,a term coined in 1902 by Rudolf hober to refer to the regulation of osmotic
pressure and ionic concentrations in the extracellular compartment of the animal body.
The evolution of efficient osmoregulatory mechanisms had extraordinarily far reaching effects
on other aspects of animal speciation and diversification.The various adaptations and physiologocal
mechanisms evolved by animals to cope with the rigors of the osmotic Environment form aspecialy
fascmating examples of the resourcefulness of evolutionary adaptation.This is the theme of an excellent
book by the late homer Smith (1953) entitled from fish to philosopher.

PROBLEMS OF OSMOREGULATION
Although Three may be hourly and Daily variations in osmotic balance,an animal is generally i nan
osmotic steady state over the long term.that is,on the average,the input-output balance sheet over an
extended periode sums up to zero (figur 12-2).water Enters with food and drink and in a freshwater
Environment it Enders primarily through the respirator epithelia-the gill surfeces of fish and
invertebrates.
12-2 in a strictly osmoregulating animal the amounts of internal and water are held relatively Constant
in the face of Environment changes.This conctancy requires that intake and outflow of water and salts
be equal over an extented period of time.such osmotic homeostasis is maintained at the cost of
metabolic energy.
12-3 two major classes of osmotic exchange between an animal and its Environment.obligatory
exchanges are those that occur in respons to physical factors over which the animals has litlle short-
term physiclogical control.controlled exchanges are those that the animal can vary physiologically to
mainttain internal homeostasis substances entering the animal by either path can leave by the other
path.
And the integument of amphibies and many invertebrates.water leavesthe body in the urine,in the
feces,and by evaporation throught the integument and lungs.
The problem of osmotic regulation Koes not end with the intake and output of water.if,that Wet
so,osmoregulation would be a relatively simple matter a Fort sitting in fresh water far more dilute than
its body fluids would merely have to eliminate the same amount of water as leaked in through its
skin,and a camcl would just stop urine production between oases.Osmorelegulation also includes the
requirement of maintaining favorable solute concentrations,in the extracellulercompartment.Thus,the
Fort immeresed in hypototic pond water is faced not only with the Seed to eliminate excess water but
also with problemof retaining salts,which tend to leak out through the skin,because the skin in
amphibians is much more permeabic than in the other vertebrata clsses.The camel is faced with a rather
different set of problems.in the face of a Limited water supply,it must make a compromise :on the one
hand,it must converse water and eliminate toxic end Products of metabolisme,such as urea on the other
hand,it must regulate the salt concrentation of its extracelluler fluida as it loses water through
evaporation and through the production of urine.
The osmotic exchanges that place between an animal and its Environment can be divided into
two classes(figure12-3):(1)obligatory exchanges-namely,thouse that occur mainly in respons to physical
factors over which the animal has little or no physiological control and(2)regulated exchanges which as
the name indicates are physiologycallycontrolled and serve to aid in maintaining internal
homeostasis.regulation of osmotic exchange forms the major theme of this chapter.The various factors
that contribute to the obligatory exchange have been reviewed by beatley(1971);these are outlined
next.
FACTORS INFLUENCING OBLIGATORY EXCHANGES
1.Gradients between the extracellular compartment and the Environment. The greather the difference
between the concentration of a substance in the external medium and that in the body fluida, the
greather the tendency for net diffusion in the direction of low concentration. Thus, although a Frog
immersed in a pond tends to take up water from its hypotonic Environment, a bony fish in seawater is
faced with the problem of losing water into the hypertonic seawater.