OSMOREGULATION
I. Introduction
A. Goals of osmoregulatory systems.
1. Volume regulation.
2. Osmotic regulation
3. Ionic regulation.
4. Waste regulation.
B. Definitions and physical-chemical laws
1. Definition: Osmolarity approximately equals the sum of the molarities of the particles in solution. The more precise definition is based on the force exerted to induce water flow. Examples:
a. a 0.1 M glucose has an approximate osmolarity of 0.1 OsM.
b. a 0.3 M KCl has an approximate osmolarity of 0.6 OsM because the KCl will dissociate into 0.3 M K+ and 0.3 M Cl-.
c. a 0.4 M MgCl2 has an approximate osmolarity of 1.2 OsM because the MgCl2 will dissociate into 0.4 M Mg++ and 2*0.4 M Cl -.
d. a solution containing 0.1 M glucose, 0.3 M KCl, and 0.4 M MgCl2 has an approximate osmolarity of 1.9 OsM (0.1 + 0.6 + 1.2).
2. Experimentally determined rule: Water flows from a low osmolarity to a high osmolarity at a rate directly proportional to the difference (gradient) in osmolarity:
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PH2O is the water permeability coefficient across the membrane between compartment A and compartment B;
A is the surface area of the membrane.
Not that the rate becomes zero when the gradient becomes zero (that is, the osmolarities on each side of the membrane are equal). Obviously systems tend toward such an equilibrium.
The phenomenon of water osmotic flow is well determined (by the equation above), but the reasons that osmosis occurs is poorly understood (something to do with the mucking and stability of hydrogen bonds between water molecules).
Examples:
a. If a 10 µL cell is losing water at the rate of 0.2 µL per minute and the cell has an osmolarity of 0.8 OsM and it is suspended in a 90 µL solution of 0.7 M NaCl, then what is the rate of water flow if the cell is 0.4 OsM?
Initially the cell is 0.8 OsM and the surrounding environment is 1.4 OsM (the NaCl disassociates into two particles) thus the initial gradient is 1.4 - 0.8 OsM = 0.7 OsM.
If the cell is 0.4 OsM then the gradient becomes 1.4 - 0.4 = 1.0 OsM. The gradient is now 1.0/0.7 = 1.43 times larger thus the rate should be 1.43 times larger. Thus the new rate is 1.43 * 0.2 µL per minute = 0.286 µL per minute
b. then what will be the equilibrium concentrations when the water flow stops.
The total volume of the system is 90 µL + 10 µL = 100 µL and can't change.
The total moles of solute can't change either and is equal to
0.8 Osmoles/Liter * 10 µL + 1.4 Osmoles/Liter * 90 µL =
8 µOsmoles + 126 µOsmoles = 134 µOsmoles.
Thus the equilibrium concentration is when this number of Osmoles are equally distributed through out the entire volume =
134 µOsmoles / 100 µL = 1.34 µOsM.
3. Experimentally determined rule: Cell membranes are typically very permeable to water. The exceptions are rare but very important (cells impermeable to water can be found in kidneys and salt glands).
4. Experimentally determined rule: Cell membranes are typically mostly impermeable to ions (although that impermeability is not zero and the are some important consequences because it is not zero). Ions primarily get across membranes by facilitated diffusion or active transport.
5. Definitions:
isoosmotic = the osmolarity of two solutions are equal.
hyporosmotic = the solution that is hyporosmotic has a lower osmolarity than the solution to which it is being compared. A hypoosmotic solution has a tendency to gain water.
hyperosmotic = the solution that is hyperosmotic has a higher osmolarity than the solution to which it is being compared. A hyperosmotic solution has a tendency to lose water.
Two solutions that are isoosmotic may still have a subsequent flow of water from one to the other. For example consider a 1 M solution of glucose separated by a membrane permeable only to glucose from a 1 M solution of urea. Even though the two solutions are initially isoosmotic, there will be a subsequent flow of water from the glucose solution into the urea solution because the glucose will move down its concentration gradient via diffusion to the urea side thus increasing the osmolarity on the urea causing water to flow in that direction. Thus, our two original solutions would be regarded as isoosmotic but the glucose solution would be hypertonic to the urea solution. Tonicity looks at the future while osmolarity looks at the present condition.
isotonic = two solutions where there is no subsequent flow of water between them.
Notice that osmolarity only depends upon the osmolarity of two solutions while tonicity additionally depends upon the kind of chemicals that make up the solutions and the nature of the separating membrane.
6. Definitions:
osmoconformer = the osmolarity of the animal changes with the osmolarity of the surrounding environment.
osmoregulator = the osmolarity of the animal is held at a constant level independent of the surrounding osmolarity.
7. Experimentally determined rule: The rate of diffusion of any molecule is directly proportional to the concentration gradient of that molecule:
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where Px is the permeability coefficient of X to the membrane.
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The gradient (subtract the two concentrations) for the situation on the right is three times the gradient on the left, therefore the rate of flow will be three times higher for the situation on the right.
8. Experimentally determined 'general rule of thumb': Measured osmolaritites of plasmas of animals are usually around 250 to 350 mOsM.
Is this the best osmolarity for living reactions?
mOsM Na+ (mM) Cl- (mM) Ocean 1000 450 540 Fresh water 1 0.08 0.05 Goldfish 293 142 107 Flounder 337 180 160 Alligator 278 140 111 Snake 330 160 140 Dove 372 176 136 Mammal 370 269 258 Shark 1075 269 258 The cytoplasm of cells is absolutely equal in osmolarity to that of plasma, but the concentration of Na+ inside the cells is much lower and the concentration of K+ is much higher.
II. Problems of animals living in different environments.
A. Salt water - water "wants" to osmotically flow out and ions in.
B. Fresh water - water "wants" to osmotically flow in while necessary ions can not be obtained by diffusion.
C. Terrestrial - water loss is a problem. Ionic balance depends on the drinking of fresh or salt water.
III. Solutions by considering specific organisms.
A. Lower animals (invertebrates) are sometimes osmoconformers (generally vary with the environment).
B. Flounder
1. Volume regulation
a. Problem: large amounts of water are osmotically pulled out through gills and gut.
b. Solution: salt water is drunk (solves the internal fluid volume problem).
i. Solution creates a new problem: Since water is not actively transported it is necessary for volume control to let the water as well as the salts into the blood stream. (The salts are probably "let" in which induces a water flow.). Thus the problem of too many ions is exacerbated.
c. Solution: excretes little urine (helps control volume problem). The animal does not need to excrete urine to get rid of nitrogen waste because almost all marine organisms get rid of their nitrogen waste by the diffusional outflow of ammonia from the respiratory surfaces (in this case the gills).
2. Osmotic and ionic regulation
a. Problem: Na+ and Cl- uptake through the high surface area of the gills. Also salt uptake in the high surface area of the gut from food intake and the drinking of salt water.
b. Solution: Actively transport out sodium chloride from the gills. There is an energy cost to do this but it is a presumably low cost relative to the whole basal metabolism of the animal (thought to be less than 5%)
i. Special cells called chloride cells do the active transport. It is still not clear if it is the Na+ that is pumped out by Na+-K+ ATPase and Cl- follows the positive charge or if Cl- is pumped out and the Na+ does the following.
C. Brine shrimp (Artemia) - same problems but just more extensive; lives in salt water with concentrations up to 7 M NaCl! Probably solved in the same way.
a. Drinks salt water.
b. Active transport of NaCl out of epithelium and neck organ (larva) via Na+-K+ ATPase.
D. Sea gull
1. Volume regulation
a. drinks salt water
b. little urine excreted (uric acid)
2. Osmotic and ionic regulation
a. Specialized salt glands with a Na+-K+ ATPase. Incredible concentrating abilities (Apparently with membranes with little water permeability). Counter current exchange helps keep the salt concentration high and distal points along the gland.
E. Shark
1. Volume regulation - isoosmotic because of high urea and TMAO content in blood. Note that she still has close to normal Na+ and Cl- concentrations but tolerates high urea. The TMAO, somehow, seems to overcome some of the toxic effects of the urea.
2. Ionic balance is not as serious, but those ions that do leak in can be removed by a Na+-K+ ATPase in the rectal glands and gills.
F. Salt frog - a good example of convergent evolution: solves problems in same way as shark with high urea in blood.
G. Whale - ionic balance achieved by ion concentrating kidneys. Only birds and mammals have loops of Henle.
H. Crappie - a fresh water animal.
1. Volume regulation
a. Problem: large amounts of water are osmotically pulled in through gills.
b. Solution: large amounts of dilute urine are released.
i. Solution creates a new problem: Since the water in urine is coming from the blood and perfectly pure water can not be made, then some ions are going to be lost this way (in addition to the ions lost at the gills)..
c. Solution: does not drink water.
2. Osmotic and ionic regulation
a. Problem: Na+ and Cl- loss through the high surface area of the gills. Also salt loss in the urine.
b. Solution: Actively transport in sodium chloride using the gills. There is an energy cost to do this but it is a presumably low cost relative to the whole basal metabolism of the animal (thought to be less than 5%)
i. it is probably not a simple reversal of the Cl- cells discussed above.
I. Frog - Water uptake through the skin is high therefore it releases copious amounts of urine. Ions transported into the body through the skin.
J. Kangaroo rat - desert, terrestrial (i.e. water is at a premium)
1. Volume regulation
a. Problem: Water is lost through respiration, urine, and evaporation. Little free water is available to drink and there is little water content in the food. In stressful times the only available water is metabolic (oxidative) water:
C6H12O6 + 6 O2 ----> 6 CO2 + 6 H2O + energy
If it is going to survive, it must survive off of this water. Getting enough food to burn to get the water could be a problem.
b. Solution: minimize water loss.
i. Behavior- be active only during cooler times (night) and minimize evaporative water loss.
ii. It has not evolved around the problem of getting rid of its nitrogen waste via urea - and urea is a problem because it must be dissolved in water. But it does excrete a very concentrated urine and the animal has excellent kidneys to carry out that process.
iii. Reduces the amount of heat lost through respiration by a nose heat exchanger. The evaporation of water in the early part of the nasal passages causes a cooling in the nasal passages so that the temperature there is less than the body temperature. When the animal exhales , the water saturated air at 38° C from the lungs comes to this cooler area and some water condenses out and is saved.
No heat exchange: water loss per liter:
Air in at 10% at 38 °C minus air out at 100% at 38 °C
5 mg/L - 50 mg/L = -45 mg/L
Heat exchange
Air in at 10% at 38 °C minus air out at 100% at 20 °C
5 mg/L - 17 mg/L = -12 mg/L
A loss of 12 mg versus 45 is a big savings.
Since he is essentially cooling the environment, this must cost the animal energy. Since incoming air is only 10% humidity, evaporation automatically causes cooling. Upon exhaling the warm saturated will become cool saturated air thus inducing condensation (rain). The longer and more convoluted and narrower the passage ways, the more easily the latter process occurs.
All animals carry out this phenomenon, but kangaroo rats have made an art out of it.
2. Osmotic and ionic regulation
a. Problem: Not a big problem. The kidney handles the problems that do occur.
E. Rattlesnake - low metabolism, impermeable skin, uric acid excretion, free water in food.
F. Flour beetle
1. Water regulation - impermeable, wax covered, stores uric acid rather than excretes it.
2. Ion regulation - Malpighian tubules (gather wastes - uric acid, salts - by active secretion since there is no "blood" pressure). Most material passes out of gut into hemolymph and the tubules pull out of that wastes which are then deposited into the hindgut.
H. Us and all mammals - the kidney