URINARY SYSTEM

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 Urinary system
- Whether an animal lives on land, in salt water, or in fresh water, water balance
depends on the regulation of solute movement between internal fluids and the
external environment. Much of this movement is handled by excretory systems.
These systems are central to homeostasis because they dispose of metabolic wastes
and control body fluid composition.

Excretory Processes

- Animals across a wide range of species produce a fluid waste called urine through
the basic steps shown in the Figure.

• In the first step, body fluid (blood,

coelomic fluid, or hemolymph) is
brought in contact with the selectively
permeable membrane of a transport
epithelium. In most cases, hydrostatic
pressure (blood pressure in many
animals) drives a process of filtration.

Cells, as well as proteins and other large

molecules, cannot cross the epithelial
membrane and remain in the body fluid. In
contrast, water and small solutes, such as
salts, sugars, amino acids, and nitrogenous
wastes, cross the membrane, forming a
solution called the filtrate.

- The filtrate is converted to a waste fluid by

the specific transport of materials into or out of
the filtrate.

• The process of selective reabsorption recovers useful molecules and water

from the filtrate and returns them to the body fluids.

- Valuable solutes—including glucose, certain salts, vitamins, hormones, and

amino acids—are reabsorbed by active transport.

• Nonessential solutes and wastes are left in the filtrate or are added to it by
selective secretion, which also occurs by active transport.

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The pumping of various solutes adjusts the osmotic movement of water into or out of
the filtrate.

• In the last step—excretion—the processed filtrate containing nitrogenous

wastes is released from the body as urine.

The major function of the urinary system

Its major function is to control the composition and volume of body fluids. The
kidneys perform this function through multiple processes:

1. Excretion. The kidneys are the major excretory organs of the body. They
remove waste products from the blood. Many waste products are toxic, but most
are metabolic by-products of cells and substances absorbed from the intestine.
The skin, liver, lungs, and intestines eliminate some of these waste products, but
they cannot compensate if the kidneys fail to function.
2. Regulation of blood volume and pressure. The kidneys play a major role in
controlling the extracellular fluid volume in the body. They can produce either a
large volume of dilute urine or a small volume of concentrated urine. Thereby,
the kidneys regulate blood volume and blood pressure.
3. Regulation of the concentration of solutes in the blood. The kidneys help
regulate the concentration of the major molecules and ions, such as glucose,
Na+, Cl−, K+, Ca2+, HCO3-, and HPO42-.
4. Regulation of extracellular fluid pH. The kidneys excrete variable amounts of
H+ to help regulate extracellular fluid pH.
5. Regulation of red blood cell synthesis. The kidneys secrete a hormone,
erythropoietin, which regulates the synthesis of red blood cells in bone
marrow.
6. Regulation of vitamin D synthesis. The kidneys play an important role in
controlling blood levels of Ca2+ regulating the synthesis of vitamin D.

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- In humans, the excretory system consists of a pair of kidneys, bean-shaped organs
about 10 cm in length, as well as organs
for transporting and storing urine.

- Urine produced by each kidney exits

through a duct called the ureter; the two
ureters drain into a common sac called
the urinary bladder.

- During urination, urine is expelled

from the bladder through a tube called
the urethra, which empties to the
outside near the vagina in females and
through the penis in males. Sphincter
muscles near the junction of the urethra
and bladder regulate urination.

- Each kidney has an outer renal cortex

and an inner renal medulla. Both regions are
supplied with blood by a renal artery and
drained by a renal vein. Within the cortex and
medulla lie tightly packed excretory tubules and
associated blood vessels. The inner renal
pelvis collects urine from the excretory tubules
and passes it to the urinary bladder.

- Weaving back and forth across the renal

cortex and medulla are the nephrons, the
functional units of the vertebrate kidney. Of
the roughly 1 million nephrons in a human kidney, 85% are cortical nephrons, which
reach only a short distance into the medulla. The remainder, the juxtamedullary
nephrons, extend deep into the medulla. Juxtamedullary nephrons are essential for
production of urine that is hyperosmotic to body fluids, a key adaptation for water
conservation in mammals.

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- Each nephron consists of a single long tubule as well as a ball of capillaries called
the glomerulus.

- The blind end of the tubule forms a cup-shaped swelling, called Bowman’s capsule,
which surrounds the glomerulus. Filtrate is formed when blood pressure forces fluid
from the blood in the glomerulus into the lumen of Bowman’s capsule.

- Processing occurs as the filtrate passes through three major regions of the nephron:
the proximal tubule, the loop of Henle (a hairpin turn with a descending limb and
an ascending limb), and the distal tubule.

- A collecting duct receives processed filtrate

from many nephrons and transports it to the
renal pelvis.

- Each nephron is supplied with blood by an

afferent arteriole, an offshoot of the renal
artery that branches and forms the
capillaries of the glomerulus. The capillaries
converge as they leave the glomerulus, forming
an efferent arteriole.

- Branches of this vessel form the peritubular

capillaries, which surround the proximal and
distal tubules. Other branches extend downward
and form the vasa recta, hairpin-shaped
capillaries that serve the renal medulla, including
the long loop of Henle of juxtamedullary nephrons.

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The nephron is organized for stepwise processing of blood filtrate

- The porous capillaries and specialized cells of Bowman’s capsule are permeable to
water and small solutes, but not blood cells or large molecules, such as plasma
proteins. Thus, the filtrate produced in the capsule contains salts, glucose, amino
acids, vitamins, nitrogenous wastes, and other small molecules.

- Because such molecules pass freely between glomerular capillaries and Bowman’s
capsule, the concentrations of these substances in the initial filtrate are the same as
those in blood plasma.

From Blood Filtrate to Urine: A Closer Look

- In this section, we will follow filtrate along its path in the nephron and collecting
duct, examining how each region contributes to the stepwise processing of filtrate into
urine.

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1. Proximal tubule.

Reabsorption in the proximal tubule is critical for the recapture of ions, water, and
valuable nutrients from the huge volume of initial filtrate.

• NaCl (salt) in the filtrate diffuses into the cells of the transport epithelium,
where Na+ is actively transported into the interstitial fluid. This transfer of
positive charge out of the tubule drives the passive transport of Cl-, as well as
the movement of more Na+ from the lumen into the cells of the tubule wall by
facilitated diffusion and cotransport mechanisms.
• As salt moves from the filtrate to the interstitial fluid, water follows by
osmosis. The salt and water then diffuse from the interstitial fluid into the
peritubular capillaries.
• Glucose, amino acids, potassium ions (K+), and other essential substances
are also actively or passively transported from the filtrate to the interstitial
fluid and then into the peritubular capillaries.

- Processing of filtrate in the proximal tubule helps maintain a relatively constant pH

in body fluids.

- Cells of the transport epithelium

secrete H+ into the lumen of the
tubule but also synthesize and
secrete ammonia, which acts as a
buffer to trap H+ in the form of
ammonium ions (NH4+).

• The more acidic the filtrate,

the more ammonia the cells
produce and secrete, and a
mammal’s urine usually
contains some ammonia
from this source (even Reabsorption in the Proximal Convoluted Tubule
though most nitrogenous waste is excreted as
urea).

- The proximal tubules also reabsorb about 90% of the buffer bicarbonate (HCO3-
) from the filtrate, contributing further to pH balance in body fluids.

- As the filtrate passes through the proximal tubule, materials to be excreted become
concentrated. Many wastes leave the body fluids during the nonselective filtration
process and remain in the filtrate while water and salts are reabsorbed.

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 o Urea, for example, is reabsorbed at a much lower rate than are salt and
water.
• Some other toxic materials are actively secreted into filtrate from surrounding
tissues.
o For example, drugs and toxins that have been processed in the liver pass
from the peritubular capillaries into the interstitial fluid. These molecules
then enter the proximal tubule, where they are actively secreted from the
transport epithelium into the lumen.

2. Descending limb of the loop of Henle.

- Reabsorption of water continues as the filtrate moves into the descending limb of
the loop of Henle. Here numerous water channels formed by aquaporin proteins
make the transport epithelium freely permeable to water. In contrast, there are almost
no channels for salt and other small solutes, resulting in very low permeability for
these substances.

- For water to move out of the tubule by osmosis, the interstitial fluid bathing the
tubule must be hyperosmotic to the filtrate. This condition is met along the entire
length of the descending limb, because the osmolarity of the interstitial fluid increases
progressively from the outer cortex to the inner medulla of the kidney. As a result, the
filtrate loses water—and therefore its solute concentration increases—all along its
journey down the descending limb.

3. Ascending limb of the loop of Henle. The filtrate reaches the tip of the loop and then travels
within the ascending limb as it returns to the cortex.

- Unlike the descending limb, the ascending limb has a transport epithelium
studded with ion channels, but not water channels. Indeed, this membrane is
impermeable to water. Impermeability to water is very rare among biological
membranes and is critical to the function of the ascending limb.

- The ascending limb has two specialized regions: a thin segment near the loop tip
and a thick segment adjacent to the distal tubule.

• As filtrate ascends in the thin segment, NaCl, which became concentrated in

the descending limb, diffuses out of the permeable tubule into the interstitial
fluid. This movement of NaCl out of the tubule helps maintain the osmolarity of
the interstitial fluid in the medulla.
• In the thick segment of the ascending limb, the movement of NaCl out of the
filtrate continues. Here, however, the epithelium actively transports NaCl into
the interstitial fluid. As a result of losing salt but not water, the filtrate becomes

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 progressively more dilute as it moves up to the cortex in the ascending limb of
the loop.

4. Distal tubule. The distal tubule plays a key role in regulating the K+ and NaCl
concentration of body fluids.

- This regulation involves variation in the amount of K+ secreted into the filtrate as
well as the amount of NaCl reabsorbed from the filtrate. Like the proximal tubule, the
distal tubule contributes to pH regulation by the controlled secretion of H+ and
reabsorption of HCO3-.

5. Collecting duct. The collecting duct carries the filtrate through the medulla to the renal
pelvis.

- The transport epithelium of the nephron and collecting duct processes the filtrate,
forming the urine. One of this epithelium’s most important tasks is reabsorption of
solutes and water.

✓ Under normal conditions, approximately 1,600 L of blood flows through a pair

of human kidneys each day, about 300 times the total volume of blood in the
body.

From this enormous traffic of blood, the nephrons and collecting ducts process about
180 L of initial filtrate.

✓ Of this, about 99% of the water and nearly all of the sugars, amino acids,
vitamins, and other organic nutrients are reabsorbed into the blood, leaving only
about 1.5 L of urine to be transported to the bladder.

- As filtrate passes along the transport epithelium of the collecting duct, hormonal
control of permeability and transport determines the extent to which the urine becomes
concentrated.

- When the kidneys are conserving water, aquaporin channels in the collecting duct
allow water molecules to cross the epithelium. At the same time, the epithelium
remains impermeable to salt and, in the renal cortex, to urea.

- As the collecting duct traverses the gradient of osmolarity in the kidney, the filtrate
becomes increasingly concentrated, losing more and more water by osmosis to the
hyperosmotic interstitial fluid.

- In the inner medulla, the duct becomes permeable to urea. Because of the high
urea concentration in the filtrate at this point, some urea diffuses out of the duct and

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