Countercurrent multiplication

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A countercurrent multiplier system is a mechanism that expends energy to create

a concentration gradient.
It is found widely in nature and especially in mammalian organs. For example, it can refer to the
process that is underlying the process of urine concentration, that is, the production
ofhyperosmotic urine by the mammalian kidney.[1] The ability to concentrate urine is also present
in birds.[1]
Countercurrent multiplication is frequently mistaken for countercurrent exchange, a similar but
different mechanism where gradients are maintained, but not established.

Physiological principles[edit]
The term derives from the form and function of the loop of Henle, which consists of two parallel
limbs of renal tubules running in opposite directions, separated by the interstital space of therenal
medulla.[citation needed]

The descending limb of the loop of Henle is permeable to water but impermeable to solutes,
due to the presence of aquaporin 1 in its tubular wall. Thus water moves across the tubular
wall into the medullary space, making the filtrate hypertonic.[citation needed]
The ascending limb is impermeable to water (because of a lack of aquaporin, a common
transporter protein for water channels in all cells except the walls of the ascending limb of
theloop of Henle) but permeable to solutes, but here Na+, Cl, and K+ are actively transported
into the medullary space, making the filtrate hypotonic (with a higher water potential). This
constitutes the single effect of the countercurrent multiplication process.[citation needed]
Active transport of these ions from the thick ascending limb creates an osmotic pressure
drawing water from the descending limb into the hyperosmolar medullary space, making the
filtrate hypertonic (with a lower water potential).[citation needed]
The countercurrent flow within the descending and ascending limb thus increases,
or multiplies the osmotic gradient between tubular fluid and interstitial space.[citation needed]

Details[edit]
Countercurrent multiplication was originally studied as a mechanism whereby urine is
concentrated in the nephron. Initially studied in the 1950s by Gottschalk and Mylle
following Werner Kuhn's postulations,[2] this mechanism gained popularity only after a series of
complicated micropuncture experiments.[3]
The proposed mechanism consists of pump, equilibration, and shift steps. In the proximal tubule,
the osmolarity is isomolar to plasma (300 mOsm/L). In a hypothetical model where there was no
equilibration or pump steps, the tubular fluid and interstitial osmolarity would be 300 mOsm/L as
well.{Respicius Rwehumbiza, 2010}
Pump: The Na+/K+/2Cl transporter in the ascending limb of the loop of Henle helps to create a
gradient by shifting Na+ into the medullary interstitium. The thick ascending limb of the loop of
Henle is the only part of the nephron lacking in aquaporina common transporter protein for
water channels. This makes the thick ascending limb impermeable to water. The action of the
Na+/K+/2Cl transporter therefore creates a hypoosmolar solution in the tubular fluid and a
hyperosmolar fluid in the interstitium, since water cannot follow the solutes to produce osmotic
equilibrium.[citation needed]
Equilibration: Since the descending limb of the loop of henle consists of very leaky epithelium,
the fluid inside the descending limb becomes hyperosmolar.[citation needed]
Shift: The movement of fluid through the tubules causes the hyperosmotic fluid to move further
down the loop. Repeating many cycles causes fluid to be near isosmolar at the top of Henle's

loop and very concentrated at the bottom of the loop. Interestingly, animals with a need for very
concentrated urine (such as desert animals) have very long loops of Henle to create a very large
osmotic gradient. Animals that have abundant water on the other hand (such as beavers) have
very short loops. The vasa recta have a similar loop shape so that the gradient does not
dissipate into the plasma.[citation needed]
The mechanism of counter current multiplication works together with the vasa recta's counter
current exchange to prevent the wash out of salts and maintain a high osmolarity at the inner
medulla.[citation needed]