The Urinary System

In t his chapter , you wi ll learn t h at

The kidneys maintain the composition of the body's extracellular fluids

starting with and asking and then asking

"' by
acting
-3
How do the
kidneys regulat e
"'
c- ·•· How does t he
,body transport, ·
urine concentrati on store, and
I
next examining
on
and volume? eliminate urine?

25.2 "'
I
I
measured by

"'
Clinical
and asking
evaluation of

"' kidney function

r -and
-- -----,
finally, exploring

Developmental Aspects
of the
L - - - J

Every day the kidneys f ilter nearly 200 liters of fluid

from our bloodstream, allowing toxins, rnetabol ic wastes, and
excess ions to leave the body in uri ne while returning needed
substances to the blood. Much like a water purification plant
CAREER CONNECTION that keeps a city's water drinkable and disposes of its wastes,
the kidneys are usually unappreciated unti l they rnalfu nction
and body fluids become contaminated.
The kidneys perform a chemical balancing act that would be
tricky even for the best chemical engineer. They maintain the
body's internal environment by:
• Regulating the total volume of water in the body and the total
concentration of solutes in that water (osmolality).

Watch a video to learn how • Regulating the concentrations of the various ions in the

O the chapter content is used

in a rea l hea lt h care setting.
extracellular fluids. (Even relatively small changes in some
ion concentrations such as K+ can be fatal.)
• Ensuring long-term acid-base balance.
Go to Mastering A&Pe > Study Area >
An imations and Videos or use quick • Excreting metabol ic wastes and foreign substances such as
access URL https://goo.gl/88srfV drugs or toxins.

974
976 UNIT 4 Mai ntenance of the Body

Figure 25.3 Position of t he kidneys agai nst the posterior

body wall. (a) Cross section viewed from inferior direction. Note
the retroperitoneal position and the supportive tissue layers of the
kidney. (b) Posterior in situ view showing relationship of the kidneys
to the 12th ribs.

Anterior
Inferior
vena cava

Supportive
tissue layers
vein

Renal ---;----,;,t
artery

a....:._;__.,,__ • Fibrous
Bodyof- - -'-s:,- - - - - -- ' capsule
vertebra G.?
Body wal l - - - - -....::,_•.....__ _ _ _ _ _ _-_.,..

Posterior

(a)

12th rib

the superior lumbar region (Figure 25.3) . Extending approxi-

mately frorn T 12 to L3, the kidneys receive sorne protection from
the lower part of the rib cage (Figure 25.3b). The right kidney is (b)
crowded by the liver and lies slightly lower than the left.
An adul t's kidney has a mass of about 150 g (5 ou nces) and
-, HOMEOSTATI C
its average dimensions are 11 cm Jong, 6 cm wide, and 3 cm
thick-about the size of a large bar of soap. The lateral surface
,. . ---&~- IMBALANCE 25.1 CLINICAl!

is convex. The rnedial surface is concave and has a vertical cleft The upper parts of both kidneys are protected by the thoracic
called the r enal hilum (see Figures 25. 1, 2 5.2, and 25.4) that cage, and the perirenal fat provides cushioning. However, the
leads into an internal space called the renal sinus. T he ureter, lower parts of the kidneys, espec ially the right k idney, are
renal blood vessels, lymphatics, and nerves all join each kidney susceptible to blunt trauma such as from falls, motor vehicle
at the hilum and occupy the sinus. Atop each kidney is an adre- accidents, or contact sports injuries. The renal artery is espe-
nal (or suprarenal) gland, an e ndocri ne gland that is function- cially vulnerable to injury from rapid deceleration during car
ally unrelated to the kidney. crashes, and may be lacerated (torn) or develop a thrombosis
Three layers of supportive tissue surround each kidney (Fig- (blood clot). The presence of he1naturia (blood in the urine) is
ure 25.3a). From superficial to deep, these are: an important sign of such renal trauma. Occasionally, surgical
• The renal fascia, an outer layer of dense fibrous connective treatment is required. • •• • •
tissue that anchors the kidney and the adrenal gland to sur-
roundi ng structures Internal Gross Anatomy
• The periren al fat capsu le, a fatty mass that surrou nds the A frontal section through a kidney reveals three distinct regions:
kidney and cushions it against blows cortex, 111.edulla, and pelvis (Figure 25.4) . The rnost superficial
• The fibrous capsule, a transparent capsule that prevents infec- region, the renal cortex, is light-colored and has a granular
tions in surrou nding regions from spreading to the kidney appearance. Deep to the cortex is the darker, reddish-brown
978 UNIT 4 Mai ntenance of the Body

,,,--- Cortical radiate

vein
Aorta Inferior vena cava
_..-- - Cortical radiate
artery
!
Renal artery
f
Renal vein

~---- - - Arcuate artery

Willt-- - -lnterlobar vein !
Segmental artery
t
lnterlobar vein
- - - - lnterlobar a rtery
~ - - Segmental arteries
, - - Renal vein
!
lnterlobar artery
f
~--+ Arcuate vein

- -Renal artery
!
Arcuate artery
t
~ · Cortical radiate vein
~ - - - - -Renal pelvis
!
Cortical radiate artery
:-- ----------}-------,
: Peritubular capillaries
' - - - - or vasa recta

-------{------------J t
Afferent arteriole Efferent arteriole

L J
Glomerulus (capillaries)

---------------------,--------------------
Nephron-associated blood vessels
(see Figure 25.8)

(a) Frontal section illustrating major blood vessels (b) Path of blood flow th rough renal blood vessels

Figure 25.5 Blood vessels of the kidney.

The renal arteries exit at right angles from the abdominal first lumbar splanchnic nerves, which course along with the renal
aorta, and the right renal artery is longer than the left because artery to reach the kidney. These sympathetic vasomotor fibers
the aorta lies to the left of the midl ine. As each renal artery regulate renal blood flow by adjusting the diameter of renal arte-
approaches a kidney, it d ivides into five s egmental arte ries rioles and also influence the formation of urine by the nephron.
(Figure 25.5). Within the renal sinus, each segmental artery
branches further to form several in terlob ar arteries. Check Your Understanding
At the cortex-rnedulla junction, the interlobar arteries branch 1. Zach is hit in the lower back by a n errant baseball. What
into the arcu ate arteries (ar'ku-at) that arch over the bases of protects his kidneys from this mechanical trauma?
the medullary pyramids. Small cortical radia te a rteries (also 2. From inside to outside, list the three layers of supportive tissue
called interlobular arteries) radiate outward from the arcuate that surrou nd each kidney. Where is the parietal peritoneum in
arteries to supply the cortical tissue. More than 90% of the relation to these layers?
blood entering the kidney perfuses the renal cortex. 3. The lumen of the ureter is continuous with a space inside the
Afferent arterioles branching from the cortical radiate arter- kidney. This space has branchi ng extensions. What are the
ies begin a complex arrangement of microscopic blood vessels. names of this space and its extensions?
These vessels are key elements of kidney function, and we will - - - - - - - - - - For answers, see Answers Appendix.
examine them in the next ,nodule when we describe the nephron.
Veins mostly trace the pathway of the arterial supply in
reverse (Figure 2 5.5). Blood leav ing the renal cortex drains Nephrons are the functional
seque ntially into the cortical r adiate, arcn ate, interlobar, and
finally renal veins. (There are no segmental veins.) The renal units of the kidney
veins ex it from the kidneys and empty into the inferior vena Learning Outcome
cava. Because the inferior vena cava lies to the right of the verte- Ill> Describe the anatomy of a nephron.
bral column, the left renal vein is about twice as long as the right.
The ren al plexus, a variable network of autonomic nerve fi- Nephrons (nef' ronz) are the structural and functional units of
bers and ganglia, provides the nerve supply of the kidney and its the kidneys. (A functional un it is the smallest physical structure
ureter. An offshoot of the celiac plexus, the renal plexus is largely capable of carrying out the function of an organ.) If you under-
supplied by sympathetic fibers from the most inferior thoracic and stand how a s ingle nephron works, then you will understand
 Chapt er 25 The Urinary System 979

Renal cortex - - - -

Renal medulla - --

/
/ ,----------
/
/ Glomeru lar capsule: parietal layer
/
/
/
/
/
/
/
- Ureter /
/ Basement
/
/
/ membrane
/
/

Kidney
,, /

/
/

/
/
/
/
/----------------------- ~ ~ ....:.__- Fenestrated endothelium
/ /
Renal corpusc,le / / of the glomerulus
/ /
• Glomerular capsule
/
/
/
/
• Glomerulus - ~ /// // Glomeru lar capsule: visceral layer
/ /
// // \
/
/
/
/
Distal
/ /
\- ,J-- >( / convoluted / [ Apical microvilli Mitochondria
/ tubule

I ~ ""

Proximal
convoluted \[_ Highly infolded
tubule 1----- - basolateral membrane
Proximal convoluted tubule cells
t
Cortex
Apical side

1___________ _
Medulla

Basolatera/ side
Thick segment- - - - - - - - -•
Thin segment - - - t Distal convoluted tubule cells

Nephron loop
• Descending limb -
G--------------------------,- •
•
• Ascending limb- -~ - - - - -I -,
Collecting duct
\
\
Nephron loop (thin-segment) cells
\
\
\
\
\
\
\ [ Pri~ Intercalated cell
\
\
\
\
\
\

Figure 25.6 Locat ion and st ructure of neph rons. Schematic view of a nephron and
collecting duct depicting the structural characteristics of epithelial cells forming various Collecting duct cells
regions.

how the kid neys work. Each kidney contains about I million needs to get rid of. Nearby nephrons empty their processed fil-
nephrons acting in parallel to make urine (Figure 25.6) . Each trate into one of the thousands of collecting ducts, which convey
nephro n creates a cell- and protein-free fil trate from blood. this flu id, now called urine, into the renal pelvis.
From that filtrate, it recovers chemicals the body needs while Each nephron consists of a renal corpuscle and a renal
also secreting into that filtrate selected chemicals that the body tubule. All of the renal corpuscles are located in the renal
980 UNIT 4 Maintenance of the Body

cortex, while the renal tubules begin in the cortex and then pass Proximal Convoluted Tubule (PCT)
into the medulla before returni ng to the cortex. The walls of the proximal convoluted tubule (PCT) are fonned
by cuboidal epithelial cells with large rnitochondria. Their apical
Renal Corpuscle (luminal) surfaces bear dense microvilli (Figure 25.6 and Fig-
Each renal corpuscle consists of a tuft of capillaries called a glo- ure 25.7). Just as in the intestine, this brush border dramatically
merulus (glo-mer'u-lus; glo,n = ball of yam) and a cup-shaped increases the surface area and capacity for reabsorbing water
hollow structure called the glomerular ca psule (or Bo,v1nan's and solutes from the filtrate and secreti ng substances into it.
capsule) (Figures 25.6 and 25.8). The glomentlar capsule is con-
tinuous with its renal tubule and completely surrounds the glo- Neph ron Loop
merulus, much as a well-worn baseball glove encloses a ball. The U-shaped nephron loop (formerly called the loop of
Henle) has descending and ascending li1nbs. The proximal
Glomerulus part of the descending limb is continuous ,vith the proximal
The endothelium of the glomerular capillaries is fenestrated tubule and its cells are similar. The rest of the descending limb,
(penetrated by many pores; .... p. 711 ), which makes these capil- called the descending thin li111b, consists of a simple squamous
laries exceptionally porous. This property allows large amounts ep ithelium. The ep ithelium becomes cuboidal or even low
of solute-rich but virtually protein-free flu id to pass from the columnar in the ascending part of the nephron loop, which is
blood into the glomerular capsule. This plasma-derived flu id therefore called the thick ascending li111b. [n roost nephrons, the
or filtrate is the raw material that the renal tubules process to entire ascending limb is thick, but in some nephrons, the thin
form urine. segment extends around the bend as the ascending thin li111b.
The thick and thin parts of the nephron loop are also referred to
Glomerular Capsule as thick and thin segments.
The glomerular capsule has an external parietal layer and a vis-
ceral layer that clings to the glornerular capillaries. Dist al Convoluted Tubule (DCT)
• The parietal layer is simple squamous epithel ium (Figures The epithelial cells of the d is ta l convoluted tu b ule (DCT),
25.6, 25.7, 25.10, and 25.12a). This layer contributes to the like those of the PCT, are cuboidal and confined to the cortex,
capsule stnrcture but plays no part in forming filtrate.
• The visceral layer, which cli ngs to the glomerular capillaries, Practice Histology questions:
consists of highly modified, branching epithelial cells called MasteringA&P• > Study Area > Lab Tools> PAL
podocytes (pod'o-sits; "foot cells") (see Figure 25.12a-c).
The octopus-like podocytes have foot processes that cling Renal corpuscle Proximal
to the basement membrane of the glomerulus. The clefts or • Squamous epithelium convol uted
of parietal layer of tubule (fuzzy
openings between the foot processes are called filtration lumen due to long
glomerular capsule
slits. Through these slits, filtrate enters the capsular space microvilli)
inside the glomerular capsule. • Glomerular capsular
Distal convoluted
space
We describe the.filtration 1nen1brane, the filter that lies bel\veen tubule (clea r
• Glomerulus lumen)
the blood in the glomerulus a nd the filtrate in the capsular
space, on pp. 984-986.

Renal Tubule and Collecting Duct

The renal tubule is about 3 cm ( 1.2 inches) long and has three
major parts. It leaves the glomerular capsule as the elaborately
coiled proximal convoluted tubule, drops into a hairpin loop
called the nephro11 loop, and then winds and twists again as
the distal convoluted tubule before emptying into a collecting
duct. The terms proxi111al and distal indicate the relationship of
the convoluted tubules to the renal corpuscle-filtrate from the
renal corpuscle passes through the proximal convoluted tubule
first and then the distal convoluted tubule, which is thus "fur-
ther away" from the renal corpuscle. The meandering nature
of the renal tubule increases its length and enhances its filtrate
processing capabilities.
Throughout their length, the renal tubule and collecting duct
consist of a single layer of epithelial cells on a basement mem-
brane. However, each region has a unique histology that reflects
its role in processing filtrate.
Fi gure 25.7 Renal cortical tissue. Photomicrograph (41 SX).
 Chapter 25 The Urinary System 981
Cortical nephron Juxtamedullary nephron
• Short nephron loop • Long nephron loop
• Glomerulus further from the cortex-medulla junction • Glomerulus closer to the cortex-medulla junction
• Efferent arteriole supplies peritubular capillaries • Efferent arteriole supplies vasa recta

RenaJ--1 Glomerulus-, Efferent

corpuscle (capillaries) arteriole
"""".......-- - Cortical radiate vein
__._--+- - Cortical radiate artery

_ __.___ Afferent arteriole

-::;<-- Collecting duct
- -.-- -Distal convoluted tubule
Afferent arteriole
...-r- -----::::::--- Efferent
arteriole

Cortex-
} medulla
Junction
Kidney
Nephron loop
• Ascending limb
!Y
• Descending limb

Fig ure 25.8 Cortica l and juxtamedullary neph rons, and '
t heir blood vessels. Arrows indicate direction of blood flow.
Capillary beds from adjacent nephrons overlap (not shown).

but they are thinner and almost entirely lack microvill i (Fig-
ure 25.6).

Collecting Duct
Each collecting d uct contains two cell types. The more numer-
ous principal cells have sparse, short microvill i and are respon-
sible for mai ntaining the body's water and Na+ balance. The
intercalated cells are cuboidal cells with abu ndant microvill i.
There are two varieties of intercalated cells (types A and B),
and each plays a role in maintaining the acid-base balance of
the blood.
Each collecting duct receives filtrate from many nephrons.
The collecting ducts run side by side through the medu llary pyr-
amids, giving them their striped appearance. As the collecting
ducts approach the renal pelvis, they fuse together and deliver
urine into the minor calyces via papi llae of the pyramids.

Classes of Nephrons
Nephrons are generall y divided into two major groups, cortical
and juxtamedullary (Fig u re 25.8).
982 UNIT 4 Mai ntenance of the Body

,,--- A glomerulus is a special

capil lary bed that is adapted
for filtration.

- - A wide afferent arteriole

supplies each glomerulus.

~ ! -- - A narrower efferent arteriole

drains each glomerulus.

Peritubular capillary beds

Figure 25.9 Blood vessels of the rena l cortex. Scanning electron microg raph of a cast of
blood vessels associated with nephrons (125X). View looking down onto the cortex.

• Cortica l nephrons accou nt for 85% of the nephrons in the Peritubu lar Capill aries
kidneys. Except for small parts of their nephron loops that dip The peritnbnlar capillaries cli ng closely to adjacent re nal
into the outer medulla, they are located entirely in the cortex. tubules and empty into nearby venules. Because they arise from
• J uxta1nednlla ry neph rons (juks"tah-me' du I-ah-re) origi- the efferent arterioles (whic h have high res istance), they only
nate close to (juxta = near to) the cortex-medulla junc- experience low pressure. As a result, these low-pressure, porous
tion, and they play an important role in the kidneys' ability capillaries readily absorb solutes and water from the tubule cells
to produce urine that is co ncentrated (whic h conserves as these substances are reclai med frorn the filtrate. Renal tubules
water). They have long nephron loops that deeply invade the are closely packed together, so the peritubular capillaries of each
medulla, and their ascending limbs have both thin and thick nephron absorb substances from several adjacent nephro ns.
segments.
Vasa Reeta
Nephron Capillary Beds Notice in Figure 25.8 that the efferent arterioles serv ing the
juxtamedullary nephrons tend not to break up into meander-
The renal tu bule of every nephro n is closely associated with
two capillary beds. The firs t capillary bed (the glo1nerulus) ing peritubular capillaries. Instead they form bundles of long
straight vessels called vasa recta (va'sah rek' tah; "straight ves-
produces the fil trate. The second (a combination of peritubular
sels'') that extend deep into the medulla paralleling the lo ng-
capillaries and vasa recta) reclaims most of that filtrate.
est nephron loops. Like all blood vessels, the vasa recta supply
Glomerulus oxygen and nutrients to the tissue through which they pass (the
renal medull a). However, the thin-walled vasa recta also play
The glomerulus, in which the capillaries run in parallel, is spe-
a n important role in forming concentrated urine, as we \viii
c ialized for filtration. It differs from all other capillary beds in
describe shortly.
the body in that it is both fed and drai ned by arterioles-the
afferen t arteriole and efferent a rteriole, respectively (Figure
25.8 and Figure 25.9). Th is arrangement maintains the high
Juxtaglomerular Complex (JGC)
pressure in the glomerulus that is needed for filtration, a process Each nephron has a j nxtaglo,ner nlar complex (JGC) (juks"-
we discuss in Module 25.4. Filtration produces a large amount tah-glo-mer' u-lar), a region where the most distal portion of
of fluid, most (99o/o) of which is reabsorbed by the renal tubule the ascending limb of the nephron loop lies against the affer-
cells and returned to the blood in the peritubular capillary beds. ent arteriole feedi ng the glomerulus (and someti mes the efferent
The afferent arterioles arise from the cortical radiate arter- arteriole) (Figure 25.10). Both the ascending limb and the
ies that run through the renal cortex. The efferent arterioles feed afferent arteriole are modified at the point of contact.
into either the peri tubular capillaries or the vasa recta.
 Cha pter 25 The Urinary System 983
Glomerular capsule Glomerulus
Efferent - -- ~ -4-d,
arteriole - Parietal !ayer Foot processes Podocyte cell body
of glomerular of podocytes (visceral layer of
capsule glomerular capsule)

Capsular
Afferent _ _ / Red blood cell
space
arteriole
Proximal
tubule cell
•
I
Juxtaglo merular
complex

• Macula densa cells - -

of the ascending limb
of nephron loop
• Extraglomerular - -----'--s::-'
;:;-'"'--Lumens of
mesangial cells .__..,,,.-:
glomerular
• Granular cells - - - - ":::.....- -ff capillaries

~ - - Endothelial cell
of glomerular
capillary
•

Glomerular mesangial
cells

Juxtaglomerular complex Renal corpuscle

Figure 25.10 Juxtaglome rular complex (JGC) of a nephron . Mesangial cells that
surrou nd the glomerular capillaries (glomerular mesangial cells) are not part of the JGC.

The JGC includes three populations of cells that help regu - 7. For the juxtamedullary nephron and collecting duct ill ustrated
late the rate of filtrate formation and systemic blood pressure. below, name each of the structu res labeled a- f.
• The 1n a cula d ens a (mak'u-lah den'sah; "dense spot") is a
group of tall, closely packed cells in the ascending limb of
the nephron loop that lies adjacent to the granular cells (Fig-
ure 25. 10). The macula dens a cells are chemoreceptors that
monitor the NaCl content of the filtrate entering the distal
convoluted tubule.
• Gran ular cells [also calledjuxraglo,nerular (JG) cells] are in
the arteriolar walls. They are enlarged smooth muscle cell s
with prominent secretory granules containing the enzyme
renin (seep. 989). Granular cells act as mechanoreceptors
that sense the blood pressure in the afferent arteriole.
• Extraglomerular mesangial cells lie between the arteriole
and tubule cells, and are interconnected by gap junctions.
These cells may pass regulatory signals between macula - - - - - - - - - - For answers, see A nswers Appen dix.
densa and granular cells.
We discuss the physiological role of the JGC on pp. 988-989.
25.3 Overview: Filtration,
Check Your Understanding absorption, and secretion are the key
4. Name the tubular components of a nephron in the order that
filtrate passes th rough them. processes of urine formation
5. What are the structu ral differences between juxtamedullary Learning Outcome
and cortical nephrons, and between their capillary beds? .... List and d efine t he three major renal p rocesses.
6. What type of capillaries are the glomerular capillaries? What is
thei r function' If you had to design a systern to chem ically balance and cleanse
the blood, how would you do it? Conceptually, it's really very
984 UNIT 4 Maintenance of the Body

simple. The body solves this problem in the following way. E) Tubular secretion. Tubular secretion ("selectively adding
First, it "dumps" cell- and protein-free filtered blood into a sep- to the waste container") is the process of selectively mov-
arate "waste container." From this container, it reclaims every- ing substances from the blood into the filtrate. Like tubular
thing the body needs to keep (which is almost everything in the reabsorption, it occurs along the length of the tubule and
container). Finally, the kidney selectively adds specific things collecti ng duct.
to the container, fine-tuning the body's che,n ical balance. Any-
The kidneys process an e normous volu,ne of blood each day.
thing left in the container becomes urine. This is basicall y how
Of the approximately 1200 ml of blood that passes through the
nephrons work. glomeruli each minute, so,ne 650 ml is plasma, and about one-
Urine formation and the adjustment of blood compos ition fifth of this (120-125 ml) is forced into the glo,nerular capsules
involve three processes (Fig ure 25.11): as filtrate. This is equ ivalent to filtering your entire plas1na vol-
O Glomeru la r filtration. Glomerular filtration ("dumping ume more than 60 times each day! Considering the magnitude
filtrate into the waste container") takes place in the renal of their task, it is not surprising that the kidneys (which account
corpuscle and produces a cell- and protein-free filtrate. for o nly I% of body \veight) consume 20-25% of all oxygen
f) Tubular reabsorption. Tubular reabsorption ("reclaiming used by the body at rest.
what the body needs to keep") is the process of selectively Filtrate and urine are quite different. Filtrate contains every-
moving substances from the filtrate back into the blood. It thing found in blood plasma except prote ins. Urine contains
takes place in the renal tubules and collecting ducts. Tubu- unneeded substances such as excess salts and metabolic wastes.
lar reabsorption reclaims almost everything filtered-all of The kidneys process about J80 L (47 gallons!) of blood-derived
the glucose and amino acids, and some 99o/o of the water, fluid daily. Of this amount, less than I% ( 1.5 L) typically leaves
salt, and other components. Anything that is not reabsorbed the body as uri ne; the rest returns to the circulation.
becomes urine.
Check Your Understanding
8. In the kidneys, tubular secretion of a
substance usually results in its excretion as well. Explain the
Afferent arteriole difference between excretion (defined in Chapter 1) and tubular
Glomerular secretion.
capillaries
L...-----==aam- For answers, see Answers Appendix.
Efferent arteriole
Cortical
radiate
Urine formation, step 1: The
artery
glomeruli make filtrate
Glomerular capsule
Learning Outcomes
Ill> Describe the forces (pressures) that promote or counte ract
- - Renal tubule and g lomerular filtration.
collecting duct Ill> Compare t he intrinsic a nd extrinsic controls of the
containing filtrate
g lomerular filtration rate.
Glomerular filtra tion is a passive process in which hydrostatic
pressure forces fluids and solutes through a membrane. The
> - - - - Peritubular
capillary glomeruli can be viewed as s imple mechanical filters because
filtrate formation does not directly consume metabolic energy.
Let's first look at the structure of the filtration membrane and
then see how it works.

To cortical radiate vein The Filtration Membrane

The filtr ation membr ane lies between the blood and the inte-
rior of the glomerular capsule. It is a porous membrane that
Three major allows free passage of water and solutes smaller than plasma
renal processes: Urine proteins. As Figure 25.12d shows, its three layers are:
O II Glomerular filtration
• Fenestrated endothelium of the g lomerular ca pillaries. The
f) II Tubular reabsorption fenestrations (capillary pores) allow a ll blood components
O i. Tubular secretion except blood cells to pass through.

Figure 25.11 The th ree major re nal p rocesses. A single

• Basement membrane. The base,nent membrane lies between
nephron is shown schematically, as if uncoiled. Each kidney actually the other two layers and is composed of their fused basal
has more than a million nephrons acting in parallel. laminae (<Ill p. 118). It forms a physical barrier that blocks all
 Chapter 25 The Urinary System 985

Glomerular
Efferent capsular space
arteriole

,--- Filtration slits

Podocyte - -
Afferent cell body
arteriole

Glomerular capillary-.;
\ _ Parietal Iayer
covered by podocytes
of glomerular Fenestrations - -.,-- -
that form the visceral layer
of glomerular capsule
capsule (pores) ~ •
(a) Renal corpuscle
Glomerular - ~
capillary
endothelium
(podocyte covering and Foot processes
basement membrane removed) of podocyte

(b) Glomerular capillary surrounded by podocytes

Fittration slits

The filtration membrane separates the

blood in the g lomerular capi llaries from
Podocyte the filtrate in t he glomerular capsule .
It has three layers:
cell body
~ - - • Capil lary endothelium
,------ • Basement membrane
• Foot p rocesses of podocyte
of g lomerular capsule

- I
- - - - - - + Filtration
slit
~ - - - - - - - - + - Slit
diaphragm
Blood in Filtrate

Fenestration
-
glomerular

-
capillary
in capsular
space

processes
(pore) of podocyte
(c) Filtration slits between the podocyte foot processes (d} Filtration membrane

Figure 25.12 The filtration membrane. (a) The renal corpuscle consists of a glomerulus
surrounded by a glomerular capsule. (b) Enlargement of a glomerular capillary covered by the
visceral (inner) layer of t he glomerular capsule consisting of podocytes. Some podocytes and the
basement membrane have been removed to show the fenestrations (pores) in t he underlying
capillary wall. (c) Scanning electron micrograph of the visceral layer (6000X). (d) Diagram of a
section through the filtration membrane showing its t hree layers.
986 UNIT 4 Mai ntenance of the Body

but the smallest prote ins whi le still permitti ng most other
solutes to pass. The glycoproteins of the gel -like basement Efferent
membra ne give it a negative charge. As a result, the base- arteriole
ment membrane electrically repels many negatively charged
macromolecular anions such as plasma proteins, rei nforci ng
the blockade based on molecular size.
• Foo t processes o f podocytes of the glome rul a r capsule. •
The visceral layer of the glomerular capsule is made of
podocytes that have fil tration slits between their foot pro- •••
cesses. If any macromolecules manage to make it through
the basement membrane, slit diaphragms- thin membranes
Afferent
that extend across the fil trat ion slits-prevent almost all of arteriole • •••
them from traveli ng farther.
Macromolecules that get "hung up" in the fil tration mem-
brane are engulfed by specialized pericytes (...ii p. 710) called
glo,nerular ,nesangial cells (Figure 25. 10). NFP = Net filtration pressure
Molecules smaller than 3 nm in diameter- such as water, = outward pressures - inward pressures
gl ucose, am ino acids, and nitrogenous wastes- pass freely = (HP90) - (HP05 + OP9 J
= (55) - (1 5 + 30)
from the blood into the glomerular capsule. As a result, these = 10 mmHg
substances usually have similar concentrations in the blood and
the glomerular filtrate. Larger molecules pass with greater dif- Figure 25.13 Forces dete rmining net filtration pressure
ficulty, and those larger than 5 nm are generally barred from (NFP). The pressure values cited in the diagram are approximate.
e nteri ng the tubule. Keeping the plasma proteins in the capillar- (HP9, = hydrostatic pressure in glomerular capillaries, OP9, =
ies maintains the colloid osmotic (oncotic) pressure (...ii p. 731) osmotic pressure in glomerular capillaries, HPcs = hydrostatic
of the glomerular blood, preventing the loss of all its water to pressure in capsular space)
the capsular space. The presence of proteins or blood cells in the
urine usually indicates a problem with the fil tration membrane.
Inward Pressures
Pressures That Affect Filtration Two inward forces inhibit filtrate formation by opposing HPgc·
The principles that govern filtratio n frorn the glomerulus are • The h yd rostatic pressure in the caps ular space (HPcs) is
the same as those that govern fil tration from any capillary bed. the pressure exerted by filtrate in the glomerular capsule. HP,,
Focus on Bulk Flow across Capillary Walls (...ii Focus Figure 19.1,
is much higher than hydrostatic pressure surrou nding most
pp. 732 - 733) s hows fil tration in normal capillary beds.
capillaries because filtrate is confined in a small space with
Figure 25.13 applies these principles to the glomerular capil-
a narrow outlet.
laries of the nephron.
• The colloid os1uotic pressu re in glo1uerular capillaries
Outward Pressures (OP gd is the pressure exerted by the proteins in the blood
Outward pressures promote fil trate formation. that "sucks" water into the capillary.
• The hydrosta tic pr essure in glomerular ca pillaries (HPgJ As shown in Figure 25 . 13, the above pressures determine
is essentially glomerular blood pressure. It is the ch ief force the net filtration pressure (NFP). NFP largely determ ines the
pushing water and solutes out of the blood and across the fil- glomerular filtration rate, which we consider next.
tration membrane. The blood pressure in the glomerulus is
extraordinarily high (approx imately 55 mm Hg compared to Glomerular Filtration Rate (GFR)
an average of 26 mm Hg or so in other capillary beds) and it The glomerular filtration r ate is the volume of fil trate formed
remai ns high across the entire capillary bed. This is because each rn inute by the combined activity of all 2 million glorneruli
the glomerular capill aries are drained by a high-resistance of the kidneys. GFR is directly proportional to each of the fol-
efferent arteriole whose diameter is smaller than the afferent lowi ng factors:
arteriole that feeds them. As a result, fil tration occurs along • Net filt ra tion pressure. NFP is the main controllable fac -
the entire length of each glornerular capillary and reabsorption tor. Of the pressures detenni ning NFP, the most important is
does not occur as it would in other capillary beds. hydrostatic pressure in the glomerulus. This pressure can be
• Theoretically, the colloid os,notic pressure in the capsular controlled by changing the diarneter of the afferent (and sorne-
space of the glomerular capsule would "pull" filtrate into times the efferent) arterioles, as we will see shortly.
the tubule. However, this pressure is essentially zero because
• Total surface area availa ble for filtration. Glomerular cap-
virtually no proteins enter the capsule, so we will not con-
illaries have a huge surface area (collectively equal to the
sider it further.
 Chapter 25 The Urinary System 987
surface area of the skin). Glomerular mesangial cells surrounding these capillar-
ies can fine-tune GFR by contracting to adjust the total surface area available for
filtration.
• Filtration membrane permeability. Glomerular capillaries are thousands of times
more permeable than other capillaries because of their fenestrations.
The huge surface area and high permeability of the filtration membrane explain how Complete an interactive tutorial:
the relatively modest IO mm Hg NFP can produce huge amounts of filtrate. Furthennore, MasteringA&P ' > Study Area>
the NFP in the glomerulus favors filtration over the entire length of the capillary, unlike Interactive Physiology
other capillary beds where filtration occurs only at the arteriolar end and reabsorption
occurs at the venous end. As a result, the adult kidneys produce about 180 L of filtrate
daily, in contrast to the 2 to 4 L formed daily by all other capillary beds combined. This
180 L of filtrate per day translates to the normal GFR of 120-125 ml/min.

Regulation of Glomerular Filtration

GFR is tightly regulated to serve two crucial and sometimes opposing needs. The kid-
neys need a relatively constant GFR to make filtrate and do their job of maintaining
extracellular homeostasis. On the other hand, the body as a whole needs a constant
blood pressure, and this is closely tied to GFR in the following way: Assuming nothing
else changes, an increase in GFR increases urine output, which reduces blood volume
and blood pressure. The opposite holds true for a decrease in GFR.
Two types of controls serve these two different needs. Intrinsic controls (renal
autoregulation) act locally within the kidney to maintain GFR, while extrinsic controls
by the nervous and endocrine systems maintain blood pressure. [n extreme changes of
blood pressure (mean arterial pressure less than 80 or greater than 180 mm Hg), extrin-
sic controls take precedence over intrinsic controls in an effort to prevent damage to
the brain and other crucial organs. These controls are summarized in Table 25.1 and
explained in detail on the next t\vo pages.
GFR can be controlled by changing a single variable-glomerular hydrostatic pres-
sure. All major control mechanisms act primarily to change this one variable. If the
glomerular hydrostatic pressure rises, NFP rises and so does GFR. If the glomerular
hydrostatic pressure falls by as little as 18%, GFR drops to zero. Clearly hydrostatic
pressure in the glomerulus must be tightly controlled. Let's see how the intrinsic and
extrinsic mechanisms accornplish this feat.

Intrinsic Controls: Renal Autoregulation

By adjusting its own resistance to blood flow, a process called renal autoregulatiou,
the kidney can maintain a nearly constant GFR despite fluctuations in systemic arterial
blood pressure. Renal autoregulation uses two different mechanisms: (I) a ,nyogenic

Table 25.1 Summary of Regulation of GFR

INTRINSIC CONTROL
(RENAL AUTOREGULATION) EXTRINSIC CONTROL

Purpose Maintain GFR in spite of changes in Maintain systemic blood pressure

blood pressure (so kidneys can "do
their job")
Mechanisms • Myogenic • Hormonal (renin-angiotensin-
aldosterone)
• Tubuloglomerular feedback
• Neural (baroreceptor reflex via
sympathetic nervous system)
Operating Mean arterial pressure between 80 and Mean arterial pressure far outside
conditions 180mm Hg of the normal range (<80 or >180
mm Hg)
988 UNIT 4 Mai ntenance of the Body

I SYSTEMIC BLOOD PRESSURE

_J
I I I
I
Blood pressure in I GFR Granular cells of r-:;;ibits baroreceptors
afferent a rterioles; I GFR juxtaglomerular I ..i~blood vessels of
complex of kidney systemic circulation
----'--
cp- Release
I Stretch of smooth
muscle in walls of
afferent arterioles
I Filtrate flow and
I NaCl in ascending
limb of nephron loop
[

Catalyzes cascade
resulting in
Renin

I
j Sympathetic
nervous system
J
• formation of
Vasodilation of
afferent arterioles [ Angiotensin II j

Macula densa cells t Aldosterone Vasoconstriction of

of juxtaglomerular secretion by systemic arterioles;
complex of kidney adrenal cortex t peripheral resistance

Release of vasoactiv e
chemicals inhibited
-4 ._
_ _ _ _ __
I I
I Na+ reabsorption
by kidney tubules;
Vasodilation of water follows
afferent arterioles

[ t Blood volume j
l t GFR I . I Syste mic
blood pressure

Tubuloglo merular
Myogenic mechanism Hormonal (renin-angiotensin-aldosterone) Neural controls
of autoregulation mechanism of
mechanism
autoregulation

Intrinsic mechanisms directly regulate GFR despite Extrinsic mechanisms indirectly regulate GFR
moderate changes in blood pressure (between 80 by maintaining systemic blood pressure, which
and 180 mm Hg mean arterial pressure). drives filtration in the kidneys.

Figure 25.14 Regu lat ion of g lomeru lar filtration rate (GFR) in the kidneys. (Note that
while the extrinsic controls are aimed at maintaining blood pressure, they also maintain GFR
since bringing blood pressu re back up allows the kidneys to maintain GFR.)

,nechanis,11 and (2) a tubuloglo,nerular feedback ,nechanism Tubu/og/omerular Feedback Mechanism Autoregulation by
(Figure 25.14, left side). the flow-dependent tubuloglomerular feed back mechanis m
is "directed" by the ,nacula densa cells of the juxtaglo,nerular
Myogenic Mechanism T he m yo genic mechanis1n (m i"o-
co,nplex (see Figure 25. 10). These cells, located in the walls of
jen'ik) reflects a property of vascular smooth muscle-it con-
the ascending limb of the nephron loop, respond to filtrate NaCl
tracts when stretched and relaxes when not stretched (~ p. 727).
concentration (v,hich varies directly with filtrate flov, rate).
Rising system ic blood pressure stretches vascular smooth mus-
When GFR increases, there is not enough time for reabsorption
cle in the arteriolar walls, causing the afferent arterioles to con-
and the concentration of NaCl in the filtrate remains high. The
strict. This constriction restricts blood flow into the glomerulus
macula densa cell s respond to high levels of NaCl in filtrate by
and keeps the GFR at a level ideal for kidney function. Declin-
releasing vasoconstrictor chernicals (ATP and others) that cause
ing systen1ic blood pressure causes dilation of afferent arterioles
intense constriction of the afferent arteriole, reducing blood
and raises glomerular hydrostatic pressure. Both responses help
flow into the glomerulus. This drop in blood flow decreases the
maintain normal NFP and GFR.
 Chapt er 25 The Urinary System 989
NFP and GFR, slowing the flow of filtrate and allowing more through the renal tubules. When macula densa cells sense
time for filtrate processing (NaCl reabsorption). the low NaCl concentration of this sluggishly flowing filtrate,
In contrast, the lov, NaCl concentration of slowly flowing they signal the granular cells to release ren in. T hey may
filtrate inhibits ATP release from macula densa cells, causing signal by releasing less ATP (also thought to be the tubulo-
vasodilation of the afferent arterioles (Figure 25.14). This allows glomerular feedback messenger), by relea~ing nu>re prosta-
more blood to flow into the glomerulus, thus increasing NFP glandin PGEi, or both.
and GFR. • Reduced stretch. Granular cells act as mechanoreceptors. A
Autoregulatory mechanisms maintain a relatively constant drop in mean arterial blood pressure reduces the tension in
GFR over an arterial pressure range from about 80 to 180 mm the granular cells' plasma mernbranes and stimulates thern to
Hg. Consequentl y, normal day-to-day changes in our blood release more renin.
pressure (such as during exercise, sleep, or changes in pos-
ture) do not cause large changes in water and solute excretion. Ot her Fact o rs Affecting GFR
However, the intri nsic controls cannot handle extremely low
Renal cells produce a battery of chemicals, many of which act
systemic blood pressure, such as ,night result from serious hen1-
as paracrines (local signaling molecules) affecting renal arteri-
orrhage (hypovole,nic shock). Once the mean arterial pressure
oles. These include adenosine and prostaglandin E2 (PGE2 ) . In
drops below 80 mm Hg, autoregulation ceases and ex trinsic
addition, the kidney makes its own locally acting angiotensin II
controls take over.
that reinforces the effects of hormonal angiotensin ll described
Ext rinsic Controls: Neural an d Horm onal Mechanisms in Chapter 19 (~ pp. 721 - 723).
The purpose of the extrinsic controls regulating the GFR is to
, HOMEOSTATIC
maintain systemic blood pressure (Figure 25. 14, right side).
~--art""'- IMBALANCE 25.3
CLINICAl!
Sympathetic Nervous System Controls Neural renal controls Abnormally low urinary output (less than 50 ml/day), called
serve the needs of the body as a whole-sometimes to the detri- anuria (ah-nu're-ah), may indicate that glomerular blood pres-
ment of the kidneys. When the volume of the extracellular fluid sure is too low to cause filtration. Renal failure and anuria can
is normal and the sympathetic nervous system is at rest, the also result from situations in which the nephrons stop function-
renal blood vessels are d ilated and renal autoregulation mecha- ing, including acute nephri tis, transfusion reactions, and crush
nisms prevail. However, when the extracellular fluid volume is . . .
lllJUfleS. • • •••
extremely low (as in hypovolemic shock during severe hemor-
rhage), it is necessary to shunt blood to vital organs, and neural
controls may override autoregulatory mechanisms. This could Check Your Understanding
reduce renal blood flow to the point of damaging the kidneys. 9. Extrinsic a nd intrinsic controls of GFR serve two different
When blood pressure falls, norepinephrine released by sym- purposes. What are they?
pathetic nerve fibers (and epinephrine released by the adrenal 10. Which of the pressures that determine NFP is regulated by
medulla) causes vascular smooth muscle to constrict, increas- both intrinsic and extrinsic controls of GFR?
ing peripheral resistance and bringing blood pressure back up 11. t¥!4#i1 Calculate net filtration pressu re given the following
toward normal. [This is the baroreceptor reflex that we dis- values: glomerular hydrostatic pressure = 50 mm Hg, blood
cussed previously ( ~ p. 719).] As part of this reflex, the afferent colloid osmotic pressu re = 25 mm Hg, capsular hydrostatic
arterioles also constrict. Constriction of the afferent arterioles pressu re = 20 Hg.
decreases GFR and so helps restore blood volume and blood - - For answers, see Answers Appendix.
pressure to normal.
.
Renin-Angiotensin -Aldosterone Mechanism As we d is -
cussed in Chapter 19 (~ pp. 722- 723), the r enin-an giotensin-
25.5Urine formation, step 2:
ald osteron e m echanis m is the body's main mechanism for Most of the filtrate is reabsorbed into
increasing blood pressure. Without adequate blood pressure
(as might be due to hemorrhage, dehydration, etc.), glomeru- the blood
lar filtration is not possible, so this mechanism regulates GFR Learning Outcomes
indirectly. II> Describe the mechanisms underlying w ater and sol ute
Low blood pressure causes the granular cells of the juxtaglo- reabsorpt ion from the rena l tubules into the peritubular
merular complex to release r enin. There are three pathways that ca pilla ries.
stimulate granular cells: II> Describe how sod ium a nd w ater reabsorption are
• Syrnpathetic nervous systern. As part of the baroreceptor regulated in the d istal tubu le a nd collecting d uct.
reflex, renal sympathetic nerves activate 13, 1-adrenergic recep- Our total plasma volume fil ters into the renal tubules about
tors that cause the granular cells to release renin. every 22 minutes, so all our plasma would drain away as urine
• Activated ,nacu/a densa cells. Low blood pressure or vasocon- in Jess than 30 minutes were it not for tub ula r reabsorp tion,
striction of the afferent arterioles by the sympathetic ner- which quickly reclaims most of the tubule contents and returns
vous system reduces GFR, slowing down the flow of filtrate them to the blood.
990 UNI T 4 Mai ntenance of the Body

The transcellul ar route

Filtrate Tubule cell Interstitial involves:
in tubule fluid
lumen
Tight Lateral © Transport across the apical
membrane.
junction intercellular
space @ Diffusio n through the
cytosol.
..©
@Transport across the
basolateral membrane. (Often

H 20 and
solutes
(D
... Transcellular route
@ ©
involves the lateral intercellular
spaces because membrane
t ransporters transport ions into
these spaces.)

© Movement through the

interstitial fluid and into the
Capillary - --l capillary.
endothelial The paracellular route involves:
cell
Paracellular route
@. Movement through leaky
t ight junctions, particularly in
the proximal convoluted tubule.
.. ®
~ Basol! eral
membrane
@ Movement through the
interstitial fluid and into the
capil lary.

Figure 25.15 Tu bular reabsorption occurs by transce llular and pa race llula r routes.
Generally, water and solutes move into the peritubular capillaries through intercell ular clefts. For
simpl icity, transporters, ion channels, intercellular clefts, and aquaporins are not depicted.

Tubular reabsorption (shown sche- in wh ich subs tances move down the ir elec troc hem ical
matically at right) is a selective trans- gradients.
epithe/ial process that begins as soon as
( You may wish to review these membrane transport processes
the filtrate enters the prox imal tubules. before you proceed.)
To reach the blood, reabsorbed sub-
stances follow either the transcellular or
paracellular route (Figure 25.15). The Tubular Reabsorption of Sodium
Tubular
transcellular route is j ust like the route reabsorption Sodium ions are the single most abundant cation in the filtrate,
we described for absorptio n in the gut and about 80% of the energy used for active transport is devoted
(~ p. 912). Transported subs tances to reabsorb ing them. Sodium reabsorption is al most always
move through the apical 1ne111bra11e, the active and via the transcellular route. Let's begin with the ATP-
cytosol, and the basolateral 111e1nbra11e driven step.
of the tubule cell and then the endothe-
lium of the peritubular cap illaries. Movement of substances in Sodiu m Transport across the Basolateral Membrane
the paracellular route-between the tubule cells-is limited Na• is ac tively transported out of the tubule cell by primary
by the tight junct ions connecting these cells. In the prox imal active transport-a Na•- K+ ATPase pump in the basolateral
nephron, however, these tight junctions are "leaky" and allow membrane (Figure 25.16 G)). From there, the bul k flow of
water and some important ions (Ca2+, Mg2 • , K•, and sorne Na•) water sweeps Na• into adjacent peritubu lar capillaries. This
to pass through the paracellular route. bulk flow of water and solutes into the peritubular capillaries is
Given healthy kidneys, virtually all organic nutrients such rapid because the blood there has low hydrostatic pressure and
as glucose and amino acids are completely reabsorbed to main- high osmotic pressure (remember, most proteins remain in the
tai n or restore normal plasma concentrations. In contrast, the blood instead of filteri ng out into the tubule).
reabsorption of water and many ions is continuously regulated
and adj usted in response to hormonal s ignals. Depending on the Sodiu m Transport across the Ap ical Membran e
substances transported, the reabsorption process may be active
Active pumping of Na• from the tubule cells results in a strong
or passive.
electrochemical gradient that favors its entry at the apical face
• Active tubular reabsor ption requ ires ATP either directly via secondary active transport (cotransport) carriers (Figure 25.16
(primary active transport; ~ p. 75) or indirectly (secondary @ , @ )or via fac ilitated diffusion through channels (not illus-
active transport; ~ pp. 75-76) for at least one of its steps. trated). This occurs because ( l) the pump mai ntains the intra-
• Passive tu b u lar reabsorption e ncompasses diffusion, cellular Na• concentration at low levels, and (2) the K+ pu mped
facilitated diffusion, and osmosis (~pp. 70- 73)-processes into the tubule cells almost immediately diffuses out into the
 Chapt er 25 The Urinary System 991
interstitial fluid via leakage channels, leaving the interior of the another solute (Figure 25.16@). Cotransported solutes move
tubule cell with a net negative charge. across the basolateral membrane by fac ilitated diffusion via
Because each tubule segment plays a slightly different role other transport proteins (not shown) before moving into the
in reabsorption, the precise mechanism by which Na• is reab- peritubular capillaries.
sorbed at the apical membrane varies.
Passive Tubular Reabsorption of Water
Tubular Reabsorption of Nutrients, Water, The movement of Na+ and other solutes establishes a strong
and Ions osrnotic gradient, and water moves by osmosis into the peritu-
The reabsorption of Na• by primary bular capillaries. Transmembrane proteins called aquaporins
O Complete an interactive (~ p. 71) aid this process by acting as water channels across
active transport provides the energy
and the means for reabsorbing
tutorial: MasteringA&P "
plasma membranes (Figure 25.16 ©).
> Study Area > Interactive
almost every other substance, includ- Physiology In continuously water-permeable regions of the renal tubules,
ing water. such as the PCT, aquaporins are always present in the tubule cell
membranes. Their presence "obliges" the body to absorb water
Secondary Active Transport in the proximal nephron regardless of its state of over- or under-
Substances reabsorbed by secondary active transport (the hydration. This water flow is referred to as obligatory water
"push" comes from the grad ient created by Na•-K+ pumping at reabsorption.
the basolateral membrane) include glucose, amino acids, sorne Aquaporins are virtually absent in the apical membranes
ions, and vitamins. In nearly all these cases, an apical carrier of the collecting duct unless antidiuretic hormone (ADH) is
moves Na+ down its concentration gradient as it cotransports present. Water reabsorption that depends on ADH is called
facultative ,vater reabsorption.

Passive Tubular Reabsorption of Solut es

r;-8 As water leaves the tubules, the concentration of solutes in
the filtrate increases and, if able, they too follow their con-
PCT J centration gradients into the peritubular capillaries. This

Filtrate Interstitial G) At the basolateral membrane, Na+

is pumped into the interstitial space by
in tubule fluid
PCT cell the Na+ -K+ AT Pase. Active Na+
lumen
transport creates concentration
gradients that drive:
@ "Downhill" Na+ entry at the
............
Na - ·~, @
3Na+ ----+ 3Na+
apical membrane .

@Reabsorption of organic nutrients

Glucose
Amino acids
....... 2K+ 4
(
<t
J G)
v 2K+
and certain ions by cotransport at the
apical membrane.
Some ions
Vitamins l"_.;;;;;;;~... .c...,._
K+
©
v Reabsorption of water by
osmosis through aquaporins. Water
reabsorption increases the
© concentration of the solutes t hat are
left behind. These solutes can then
be reabsorbed as they move down
their gradients:
Lipid-soluble ® @ Lipid-soluble substances
substances
diffuse by the transcellular route.
Various ions ®various ions (e.g., c1 -, Ca2 +, K +)
and urea Paracellular
\_ Tight junction and urea diffuse by the paracellular
route route.

_.,...~ Primary active transport • Transport protein

• • . . Secondary active transport Ion channel
_.,...~ Passive transport (diffusion)
e Aquaporin

Fig ure 25.16 Tu bular reabsorption of water and nutrients uses active and passive
transport. PCT cells (shown here) reabsorb 800fc, of the filtrate. Microvilli have been omitted
for simplicity.
992 UNIT 4 Mai ntenance of the Body

phenomenon-solutes following solvent-explai ns the passive The rule for water is that it leaves the descending (but not the
reabsorption of a number of solutes present in the filtrate, such ascending) limb of the nephron loop. The opposite is true for
as lipid-soluble substances, certain ions, and some urea (Fig- solutes. Virtually no solute reabsorption occurs in the descend-
ure 25.16 ®· @). It also explains in part why lipid-soluble ing limb, but solutes are reabsorbed both actively and passively
drugs and environmental poll utants are d ifficul t to excrete: in the ascending limb.
Since lipid-soluble compounds can generally pass through In the thin segment of the ascending limb, Na• moves pas-
membranes, they will follow their concentration gradients and sively down the concentration gradient created by water reab-
be reabsorbed, even if this is not "desirable." sorption. In the thick ascending limb, a Na•-K•-2c1- symporter
As Na• ions move through the tubule cells into the peritubu- ( <1111 p. 76) is the main means of Na• entry at the apical surface. A
lar capillary blood, they also establish an electrical gradient that Na•-K• ATPase operates at the basolateral membrane to create
favors passive reabsorption of anions (primarily CI·) to restore
electrical neutrali ty in the filtrate and plasma.
Cortex
Transport Maximum
65°/o of fihrate volume Regulated reabsorption
The transcellular transport systems for the various solutes are reabsorbed • Na+ (by aldosterone;
quite specific and limited. There is a transpor t maximun1 (T ..) •HO c 1- follows)
for nearly every substance that is reabsorbed using a transport
protein in the membrane. The T ,. (reported in mg/min) reflects
the number of transport proteins in the renal tubules available
2+ -
• Na , HC03 , and
many other ions
• Glucose, amino acids,
and other nutrients
• Ca2 + (by parathyroid
hormone)
l
to ferry a particular substance. In general, there are plenty of
transporters for substances such as glucose that need to be
•
retained, and few or no transporters for substances of no use
to the body.
When the tra nsporters are saturated-that is, all bound to
• H+ and NH +
the su bstance they transport- the excess is excreted in urine. • Some drugs • Regulated
secretion
This is what happens in individuals who become hyperglyce- • K+ (by
mic because of uncontrolled diabetes melli tus. As plasma levels
of glucose approach and exceed L80 mg/di, the glucose T"' is -----
aldosterone)

exceeded and large amou nts of glucose may be lost in the urine

.I
even though the renal tubules are still functioning normally. Outer Regulated
medulla reabsorption
• H2 0 (by ADH)
Reabsorptive Capabilities of the Renal • Na+ (by
Tubules and Collecting Ducts aldosterone; c1-
follows)
Table 25.2 compares the reabsorptive ab ilities of various • Urea (increased
regions of the renal tubules and collecting ducts. by ADH)

Proximal Convolut ed Tubule

T he entire renal tubule is involved in reabsorption to some
degree, but the PCT cells are by far the most active reabsorb- Regulated
ers and the events j ust described occur mainly in this tubular secretion
Inner • K+ (by
segment. Normally, the PCT reabsorbs all of the glucose and medulla aldosterone)
am ino acids in the fil trate and 65% of the Na• and water. The
bulk of the electrolytes are reabsorbed by the time the filtrate
reaches the nephron loop. Nearly all of the uric acid and about
• Reabsorption or secretion
half of the urea are reabsorbed in the prox imal tubule, but both to maintain blood pH
are later secreted back into the filtrate. described in Chapter 26;
involves H+, HC03 - ,
Nephron Loop and NH4 +

Beyo nd the PCT, the permeabili ty of the tubule epithel ium

changes dramatically. Here, for the first time, water reabsorp- -+ Reabsorption
__.... Secretion
tion is not coupled to solute reabsorption. Water can leave the
descending limb of the nephron loop but not the ascending Fi gure 25.17 Summary of tubular reabsorption and
limb, where aquaporins are scarce or absent in the tubule cell secretion. The various regions of the renal tubule carry out
membranes. These penneability differences play a vital role in reabsorption and secretion and mai ntain a gradient of osmolality
the kidneys' abili ty to form dilute or concentrated urine. withi n t he medullary interstitial fluid. Color gradients represent
va rying osmolality at different points in the interstitial fluid.
 Chapter 25 The Urinary System 993

Table 25.2 Reabsorption Capabilities of Different Segments of the Renal Tubules and Collecting Ducts
TUBULE SEGMENT SUBSTANCE REA BSORB ED MECHANISM
Proximal Convoluted Tubule (PCT)
Sodium ions (Na•) Primary active transport via basolateral Na•-K• pump; crosses apical
membrane through channels, symporters, or antiporters
Virtually all nutrients (glucose, amino acids, Secondary active transport with Na•
vitamins, some ions)
o-, K+, Mg 2•, Ca 2•, and other ions Passive paracellular d iffusion d riven by electrochemical gradient
HC03- Secondary active transport linked to H• secretion and Na•
reabsorption (see Chapter 26)
Water Osmosis; driven by solute reabsorption (obligatory water
reabsorption)
Lipid-soluble solutes Passive diffusion driven by the concentration gradient created by
reabsorption of water
Urea Primarily passive paracellular diffusion driven by chemical gradient
Nep hron Loop
Descending limb Water Osmosis
Ascending limb Secondary active transport of er-, Na+, and K• via Na•-K•-20-
cotransporter in thick portion; paracellular diffusion; Na•-H• antiport
Ca 2•, Mg 2• Passive paracellular diffusion d riven by electrochemical gradient

Distal Convoluted Tubule (DCT)

Na•, er- Primary active Na• transport at basolateral membrane; secondary
active transport at apical membrane via Na•-Cf- symporter and
channels; aldosterone-regulated at distal portion
ca 2• Passive uptake via PTH-modulated channels in apical membrane;
primary and secondary active t ransport (antiport w ith Na•) in
basolateral membrane

Collecting Duct
Primary active transport of Na• (requires aldosterone); passive
paracellular d iffusion of some er; cotransport of er- and HC03-; K•
is both reabsorbed and secreted (aldosterone dependent), usually
result ing in net K• secretion
Water Osmosis; controlled (facultative) water reabsorption; ADH required
to insert aquaporins
Urea Facilitated d iffusion in response to concent ration gradient in the
deep medulla region; recycles and contributes to medullary osmotic
gradient

the ionic gradient that drives the symporter. The thick ascend- in the OCT and collecting duct (Figure 25.17). Because most
ing limb also has Na+-H+ antiporters. In addition, sorne 50o/o of of the fil tered water and solutes have been reabsorbed by the
Na+ passes via the paracellular route in this region. time the OCT is reached, only a small amount of the filtered load
is subject to this fine tuning (e.g., about 10% of the originally
Distal Convoluted Tubule and Collecting Duct fil tered NaCl and 25% of the water). We introduce hormones
While reabsorption in the PCT and nephron loop does not that act at the OCT and collecting duct here but discuss them in
vary with the body's needs, hormones fine-tune reabsorption more detail in Chapter 26.
994 UNIT 4 Mai ntenance of the Body

• Antid iuretic ho rmone (AD H). As its name reveals, ADH

inhibi ts diuresis (di"u-re' sis), or urine output. ADH makes
the principal cells of the collecti ng ducts more permeable to
water by causing aquaporins (<Ill p. 71 ) to be inserted into
their apical membranes. The amount of ADH determines the
number of aquapori ns, and so regulates the amou nt of water
that is reabsorbed there. When the body is overhydrated,
extracell ular fluid osmolal ity decreases, decreasing ADH
secretion by the posterior pituitary (<Ill p. 613) and mak ing
the collecting ducts relatively impermeable to water. ADH
also increases urea reabsorption by the collecting ducts, as
we will describe later. 15. Primary and secondary active transport
processes are shown in Figure 25.16 (and were introduced in
• Aldosterone. Aldostero ne fine- tunes reabsorption of the Chapter 3). How do they differ?
remaini ng Na•. Decreased blood volume or blood pressure,
r-----------. For answers, see Answers Appendix.
or high extracellular K+ concentration (hyperkalemia), can
cause the adrenal cortex to release aldosterone to the blood
(<Ill pp. 623-624). Except for hyperkalemia (which directly
stimulates the adrenal cortex to secrete aldosterone), these
25.6 Urine formation, step 3: Certain
conditions promote the renin-ang iotensin-aldosterone mech- substances are secreted into the
anism (see Figure 25. 14).
Aldosterone targets the principal cells of the collecti ng
filtrate
ducts and cells of the distal portion of the DCT (prodding Learning Outcome
them to synthesize and retain more apical Na• and K+ chan- .... Describe the importance of tubular secretion and list
nels, and more basolateral Na•-K+ATPases). As a result, little several substances that are secreted.
or no Na+ leaves the body in urine. In the absence of aldos-
The most important way to clear plasma
terone, these segments reabsorb much less Na+ and about 2 %
of unwanted substances is to simply not
of Na+ filtered daily can be lost-an amou nt incompatible
reabsorb them from the fil trate. Another
with life. way is tubular secretion-essentially,
Physiologically, aldosterone's role is to increase blood
reabsorption in reverse (shown schemati-
volume, and therefore blood pressure, by enhancing Na+
cally at right). Tubular secretion moves
reabsorption. In general, water foll ows Na+ if aquaporins are selected subs tances [such as H+, K•,
present. Aldosterone also reduces blood K+ concentrations NH4 + (ammonium ion), creatinine, and Tubu lar
because aldosterone-induced reabsorption of Na+ is coupled secretion
certain organic acids and bases] from
to K+ secretion in the principal cells of the collecti ng duct.
the peritubular capillaries through the
That is, as Na+ enters the cell, K+ moves into the lumen. tubule cells into the filtrate. Also, some
• Atrial natriuretic peptid e (AN P). In contrast to aldosterone, substances (such as HC03·) that are syn-
which acts to conserve Na+, ANP reduces blood Na+, thereby thesized in the tubule cells are secreted.
decreasing blood volume and blood pressure (<Ill p. 624). The urine eventually excreted contains both filtered and
Released by cardiac atrial cells when blood volume or blood secreted substances. With one maj or exception (K•), the PCT
pressure is elevated, AN P exerts several effects that lower is the main site of secretion, but the collecting ducts are also
blood Na• conte nt, including direct inhibition of Na• reab- active (Figure 25.17).
sorption at the collecting ducts. Tubular secretion is important for:
• Parathyroid hormone (PTH). Acting primarily at the DCT, • Disposing of substances, such as certain drugs and ,netab-
PTH increases the reabsorpt ion of Ca2• ( <111 p. 622). olites, that are tightly bound to plas,na proteins. Because
plasma proteins are generally not filtered, the substances they
Check Your Understanding bind are not filtered and so must be secreted.
12. In wh ich part of the nephron does most reabsorption occur? • Eli,ninating undesirable substances or end products that
13. How does the movement of Na• drive the reabsorption of have been reabsorbed by passive processes. Urea and uric
water and solutes? acid, two nitrogenous wastes, are both handled in this way.
14. l•)®WJ In the figure at right above, draw arrows to represent Urea handling in the nephron is complicated and will be dis-
the pathway for transcell ular reabsorption of Na•. If transport cussed o n p. I000, but the net effect is that 40-50% of the
proteins are required for this process, draw and label them urea in the filtrate is excreted.
in the appropriate locations. In each case, state whethe r the
transport of Na• is primary active, secondary active, or passive. • Ridding the body of excess K'. Because virtually all K+ present
in the filtrate is reabsorbed in the PCT and ascending nephron
loop, nearly all K+ in urine comes from aldosterone-driven
 Chapt er 2 5 The Urinary System 995
active tubular secretio n into the late DCT a nd collecting 1\vo types of cou ntercurrent mechanisms determ ine uri ne
ducts. concentration and volume:
• Controlling blood pH. When bl ood p H drops toward the • The countercurrent multiplier is the interaction bet\veen the
acid ic end of its homeostatic range, the renal tubule cells flow of filtrate through the ascending and descending limbs of
actively secrete more H+ into the filtrate and retain and gen- the long nephron loops of juxtamedullary nephrons.
erate more HC03 - (a base). As a result, blood pH rises and • The countercurren t exchanger is the flow of blood through
the urine drains off the excess H•. Conversely, when blood the ascending and descending portions of the vasa recta.
pH approaches the alkaline end of its range, CJ- is reab-
These countercurrent mechanisms establish and mainta in an
sorbed instead of HC0 3-, which is allowed to leave the body
osrn otic gradient extending frorn the cortex through the depths
in urine. We will discuss the kidneys' role in pH homeostasis
of the medulla (Figure 25.18). Th is gradient-the medullary
in more detail in Chapter 26.
osmotic gra dien t-allows the kidneys to vary uri ne concentra-
tion dramatically.
Check Your Understanding How do the kidneys form the osmotic gradient? Focus on the
16. List several substances that are secreted into the kidney tubules. Medullary Osnrotic Gradient (Focus Figure 25.1 on pp. 996-997)
For answers, see Answers Appendix. explores the answer to this question.

The kidneys create and use an This greatly enlarged nephron and collecting duct
show you the arrangement of structures that pass
osmotic gradient to regulate urine through the osmotic gradient in the renal medulla.
If you keep t his anatomy in mind, it wil l help you
concentration and volume understand the process of ma king either very
d ilute or very concentrated urine.
Learning Outcomes
II- Describe t he mecha nisms responsible for the medu lla ry
The osmolality of the
osmotic grad ient. interstitial fluid in the
II- Explain how d ilute and concentrated u rine a re formed. renal cortex is isotonic
at 300 mOsm .
From day to day, and even hour to hour, our intake and loss of
fluids can vary dramatically. For example, when you run on
a hot summer day, you dehydrate as you rapidly lose fl uid as
sweat. On the other hand, if you drink a pitcher of lemonade
while sitting on the porch, you may overhydrate. In response, The osmolality of the
interstitial fluid of the
the kidneys make adjustments to keep the solute concentration renal medulla increases
of body fluids constant at about 300 mOsm, the normal osmotic prog ressively f ro m 300
concentration of blood plasma. Maintaining constant osmolality mOsm at the
cortex-medulla
of extracellular fluids is crucial for preventing cells, particularly junction to 1200 mOsm
in the brain, frorn shrinking or swelling from the osmotic move- at the medulla-pelvis
ment of water. junction.
Recall from Chapter 3 (pp. 71-73) that a solution's osmo-
lal ity is the co ncentration of solute particles per kilogram of
water. Because I osmol (equivalent to I mole of part icles) is a
fairly large un it, the milliosmol (mOsm) (mil"e-oz' mol), equal
to 0 .001 osmol, is generally used. In the discussion that follows,
we use mOsm to indicate mOsm/kg.
The kidneys keep the solute load of body fluids constant by
regulating urine concentration and volume.
• When you dehydrate, your kidneys produce a small volume of
concentrated urine. Figure 25.18 Osmotic gradient in t he re nal medulla.
• When you overhydrate, your kidneys produce a large voltune
of dilute urine.
*The term "countercurrent" is commonJy misunderstood to mean that the
The kidneys accomplish this feat using cou ntercurrent mech-
direction of fluid flow in the nephron loops is opposite that of the blood in
anisms. In the kidneys, the term countercurrent means that fluid the vasa recta. In fact, there is no one•to·one relationship between individual
flows in opposite directions through adjacent segments of the nephron loops and capillaries of the vasa recta as might be suggested by two-
same tube connected by a hairpin turn* (see Focus on the Med- dimensional diagrams such as Focus Figure 25. I. Instead, there are many
tubules and capillaries packed together like a bundle of straws. Each tubule is
ullary Osn,otic Gradient, Focus Figure 25.1, pp. 996-997). This surrounded by many blood vessels, whose flow is not ne.cessariJy counter to
arrangement makes it possible to exchange materials between flow in that tubule (see Figure 25.8}.
the t\vo segments. (Texr co111i1111es on p. 998.)
Juxtamedullary nephrons create an osmotic gradient within the renal medulla
that allows the kidney to produce urine of varying concentration.

The big picture:

Three key players
interact with the medullary
osmotic gradient.
300

300

The osmotic gradient

The osmolality (solute
concentration) of the
400 l
medullary interstitial fluid 600 The long nephron loops of
progressively increases juxtamedullary nephrons create
from the 300 mOsm of the gradient. They act as
normal body fluid to 1200 800 countercurrent multipliers.
mOsm at the deepest part
of the medulla.

300

Lower concentration - - - 300

of solutes

400

600 I The vasa recta preserve the

gradient. They act as
counterc urr ent exchangers
800 (see Figure 25. 19).

Higher concentration - - -
of solutes

300

300

400
The collecting ducts of
all nephrons use the gradient
600
to adjust urine osmolality
(see Figure 25.20). I

800

996
 Long nephron loops of juxtamedullary nephrons create the gradient.
The countercurrent multiplier depends on three properties These properties establish a positive feedback cycle that
of the nephron loop to establish the osmotic gradient. uses the flow of fluid to multiply the power of the salt pumps.

Filtrate f lows in t he
opposite direction
(cou ntercu rrent)
t hro ug h t wo
adjacent parallel
sections of a
t Water leaves the
descending limb
t Interstitial fluid
osmolality
nephron loop.

H20 4 NaCl H20 4 NaCl

The descending
limb is permeable
to water, but not
t The ascending limb
is impermeable to
w ater, and p umps
t Osmolality of filtrate
in descending limb
t Salt is pumped out
of the ascending limb
to salt. out salt.

~ Active transport

~ Passive transport

- Water impermeable
t Osmolality of filtrate
entering the ascending
limb

As water and solutes are reabsorbed, the loop first concentrates the filtrate, then dilutes it.

300 r 100
\
Cortex (__,.. 100
300 300
G) Filt rate entering the
nephron loop is isosmotic to
both blood plasma and cortical
H20 + - l NaCl
@ Filtrate is at its most dilute
as it leaves the nephron loop . At
100 mOsm, it is hypo-osmotic to

- E
interstitial fluid .
H20 + -
400
NaCl
200
the interstitial fluid.

-~
400

~
;;:::
H20 + - l NaCl
t @ Na • and c1 - are pumped out
of the filtrate. This increases the
Outer
interstitial fluid osmolality.
"'=600
I!
medulla H20 + -
6
NaCl
400
JI
,s; @water moves out of the ~ +-
0 filtrate in the descending limb
~ down its osmotic gradient. NaCl
This concentrates the filtrate. ~~

J 900
H:z()~ @ Filtrate reaches its highest
concentration at the bend of the
loop.

997
998 UNI T 4 Mai ntenance of the Body

The Countercurrent Multiplier difference reflects the power of the ascending limb 's NaCl
pumps, which are just powerful enough to create a 200 mOsm
Take some ti me to study the mechanism of the cou ntercurrent
difference between the inside and outside of the ascending limb.
multiplier in Focus Figure 25. l (pp. 996-997). The countercur-
A 200 mOsm gradient by itself would not be e nough to allow
rent multiplier depends o n actively transporting solutes out of
excretion of very concentrated urine. The beauty of this system
the ascending limb ("S tart" of the positive feedback cycle).
lies in the fact that, because of countercurrent flow, the nephron
Although the two limbs of the nephron loop are not in direct
loop is able to "rnultiply" these small changes in solute concen-
contact with each other, they are close e nough to influence
tration into a gradient change alo ng the vertical length of the
each other's excha nges with the interstitial fluid they share.
loop (both inside and outside) that is closer to 900 mOsm ( 1200
The more NaCl the ascending limb extrudes, the more water
mOsm - 300 mOsm).
diffuses out of the descending li mb and the saltier the filtrate in
Notice also that while much of the Na+ and c 1- reabsorption
the descending li mb becornes. The ascending limb then uses the
in the ascending limb is active (via Na+-K+-2c1- cotransporters
increasingly "salty" filtrate left behind in the descending limb
in the thick ascending limb), some is passive (mostly in the thin
to raise the osmolal ity of the medullary interst itial fluid even
portion of the ascending limb).
further. This establishes a positive feedback cycle that produces
the high osmolality of the flu ids in the descending limb and
interstitial fluid.
The Countercurrent Exchanger
Notice at the bottom of the rig ht page of Focus Figure The vasa recta act as countercurrent exchangers (Figure 25.19).
25. l that there is a constant difference in filtrate concentra- Countercurrent exchange does not create the medullary gradi-
tion (200 mOsm) between the two limbs of the nephron loop, ent, but preserves it by ( l) preventing rapid removal of salt from
and between the ascend ing limb and the interst itial fluid. This the medullary interstitial space, a nd (2) removing reabsorbed
water. As a result, blood leaving and reentering the cortex via
the vasa recta has nearly the same solute concentration.
The water picked up by the ascend ing vasa recta includes
( Vasa recta preserve the gradient. ) not o nly water lost from the descending vasa recta, but also
water reabsorbed from the nephron loop and collecting duct.
• The vasa recta are highly permeable to water and solutes.
• Countercurrent exchanges occur between each section of
As a result, the volume of blood at the end of the vasa recta is
th e vasa recta and its surround ing fluid. As a result: greater than at the beginning.
• The blood within the vasa recta remains nearly isosmotic
to the surrounding fluid.
• The vasa recta are able to reabsorb water and solutes into
Formation of Dilute or
the general circulation without undoing the osmotic gradient Concentrated Urine
created by the countercurrent multiplier.
As we have j ust seen, the kidneys go to a great deal of trou-
ble to create the medullary osmotic gradient. But for what pur-
Blood from ' \ pose? Without this gradie nt, you would not be able to raise the
efferent
300 arteriole concentration of urine above 300 mOsm-the osmolality of
interstitial fluid. As a result, you would not be able to conserve
3
water when you are dehydrated.
NaCl d
H20 Figure 25.20 shows the body's response to either over-
3 hydration or dehydration and ADH's role in co ntrolling the
400
The counter- production of d ilute or concentrated urine. When we are over-
current flow
of flu id moves
hydrated, ADH production decreases (-<Ill p. 613) and the osmo-
through two lality of urine falls as low as I00 mOsm. If aldosterone (not
adjacent shown) is present, the DCT and collecting duct cells can remove

600
t
600........
parallel
sections of
Na+ and selected other ions from the filtrate, maki ng the urine
that enters the renal pelvis even more dilute. The osmolality of
vasa recta.
uri ne can plunge as low as 50 mOsm, about one-sixth the con-
centration of glomerular filtrate or blood plas ma.
When we are dehydrated, the posterior pitu itary releases
900 9001=== large amounts of ADH a nd the solute concentratio n of urine
may rise as high as l 200 ,nOsm, the concentration of interstitial
NaCl - + - + NaCl
900 fluid in the deepest part of the medulla. With max imal ADH
H2 0 +- +- H20
secretion, up to 99% of the water in the filtrate is reabsorbed
1200 L= = = = \
Vasa recta---
1200 j =========~ and returned to the blood, and only half a liter per day of highly
concentrated urine is excreted. The ability of our kid neys to
produce such concentrated urine is critically tied to our abili ty
Figure 25.19 Countercurrent exchange. to survive for a limited time without water.
 Chapter 25 The Urinary System 999

( Collecting ducts use the gradient. )

(a) If we were so overhydrated we had no ADH ... (b) If we were so dehydrated we had maximal ADH ...

I Osmolality of extracellular fluids t Osmolality of extracellular fluids

I
•
I ADH release from posterior pituitary t ADH release from posterior pituitary
I
I Number of aquaporins (H2 0 •
channels) in collecting duct t Number of aquaporins (H2 0 channels) in collecting duct

•
I H2 0 reabsorption from collecting duct

•
Large volume of d ilute urine
t H2 0
'
reabsorption from collecting duct
I
Small volume of concentrated urine

- - Collecting Collecting duct - -

duct
Descending limb 300 Descending limb 300
of nephron loop of nephron loop
100 100 150

i OCT 100
i (
obT 300

Cortex Cortex
300 100 300
-
E
300 100 300 -
E
NaCl ~ 0E NaCl ~ 0E
~

- NaCl ~
-
600 400 600
100
Outer
medulla -
0
.~
Outer
medulla
H20 ~
NaCl ~ "'
0 NaCl ~ , - .urea ~
900 ~ 7 900
0

Urea ~

Inner Inner
medulla medulla
100
1200 1200
t
-'4~~ Active transport
-ol~~ Passive transport
Large volume
of d ilute urine
Urea contributes to the
osmotic gradient. ADH
increases its recycling .
'
Small volume of
concentrated urine

Figure 25.20 Mechanism for forming dilute or concentrated urine. Under the control
of antidiuretic hormone (ADH), the collecting ducts fi ne-tune urine concentration using the
medullary osmotic gradient.
1000 UNIT 4 Maintenance of the Body

Urea Recycling and the Medullary ll> list several abnormal urine com ponents, and name the
condition characterized by the presence of detectable
Osmotic Gradient amounts of each.
We're not quite done yet. There's one last piece of the puzzle Jeft-
urea. We usually think of urea a~ simply a metabolic waste prod- Since the tirne of the ancient Greek Hippocrates, physicians
have exam ined their patients' urine for s igns of disease. Uri-
uct, but conserving water is so important that the kidneys actually
n alysis, the analysis of urine, can aid in the diag nosis of dis-
use urea to help form the medullary gradient (Figure 25.20).
eases or de tect illegal substances. However, to measure how
1. Urea enters the filtrate by facilitated diffusion in the ascend- well the kidneys are functioning, we need to analyze both blood
ing thin limb of the nephron loop. and urine. Renal function can quickly be assessed by measuring
2. As the fil trate moves on, the cortical collect ing duct usually levels of nitrogenous wastes in blood. But to precisely measure
reabsorbs water, leaving urea behind. kidney function, we need to determine the renal clearance, which
3. When filtrate reaches the portion of the collecting duct in requires that both blood and urine be tested. Normal blood and
the deep medullary region, the now highly concentrated urea urine values for various substances are found in Appendix F.
moves by fac ilitated diffusion out of the collecting duct into
the interstitial fluid of the medulla. These movements form Renal Clearance
a pool of urea that recycles back into the ascending thin limb
of the nephron loop. In this way, urea co ntributes substan- R enal clear ance refers to the volume of plasma frorn wh ich the
kidneys clear (completely remove) a particular substance in a
tially to the high osmolality in the medulla.
given time, usually I minute. Renal clearance tests are done to
Antidiuretic hormone e nha nces urea transport out of the determine the GFR, which allows us to detect glornerular dam-
medullary collecting duct. Whe n ADH is present, it increases age and follow the progress of renal disease.
urea recycli ng and strengthens the medullary osmotic gradient, The renal clearance rate ( C) of any substance, in ml/min, is
allowing more concentrated uri ne to be formed. calculated from the equation
C = UV/P
Diuretics
There are several types of diuretics, chemicals that enhance where
urinary output. Alcoho l e ncourages diuresis by inhibiting U = concentration of the substance in urine (mg/ml)
release of ADH. Other diuretics increase urine flow by inhibit-
V = flow rate of uri ne formation (ml/min)
ing Na+ reabsorption and the obligatory water reabsorption that
normally follows. Examples include many drugs prescribed for P = concentration of the substance in plasma (mg/ml)
hypertension ( <1111 p. 724) or the edema of congestive heart failure Because it is freely fi ltered a nd neither reabsorbed nor
( <1111 p. 700). Most diuretics inhibit Na+-associated symporters. secreted by the kidneys, inulin (i n'u-lin) is the substance used
"Loop diuretics" (l ike furosem ide) are powerful because th ey to deterrnine the GFR. lnuli n is a plant polysaccharide that has a
inhibit formation of the medullary gradient by acting at the renal clearance value equal to the GFR. When inulin is infused
ascending limb of the nephron loop. Thiazides are Jess potent suc h that its plasma concentrat ion is I mg/ml (P = I mg/ml),
and act at the DCT. An os,notic diuretic is a substance that is then generally U = 125 mg/ml, and V = I rn l/min. Therefore,
not reabsorbed and that carries water out with it (for example, its renal clearance is C = ( 125 x I)/1 = 125 ml/min, meaning
the high blood glucose of a diabetes mellitus patient). that in I minute the kidneys have cleared all the inulin present
in 125 rn l of plasma.
Check Your Understanding The clearance value tells us about the net handling of a sub-
17. Describe the special characteristics of the descending and stance by the kidneys. There are three possible cases:
ascending limbs of the nephron loop that cause the formation
• A cleara nce value Jess than that of inul in means that the
of the medullary osmotic gradient.
substance is reabsorbed. For example, C is normally 70 ml/
18. Under what conditions is ADH released from the posterior
pituitary? What effect does ADH have on the collecting ducts? min for urea, meaning that of the 125 ml of glomerular fil-
trate formed each mi nute, approximately 70 ml is completely
- - - - - - - - - - For answers, see Answers Appendix.
cleared of urea, while the urea in the remaining 55 ml is
---
recovered and returned to the plasma. If C is zero (such as for
glucose in healthy individuals), reabsorption is complete or
25 .8 Renal function is evaluated by the substance is not filtered.
analyzing blood and urine • If C is equal to that of inulin, there is no net reabsorption or
Learning Outcomes secretion.
ll> Define renal clearance and explain how this value • If C is greater than that of inulin, the tubule cells are secreti ng
summarizes the way a substance is hand led by t he the substance into the filtrate. This is the case with most drug
kid ney. metabolites. Knowing a drug's renal clearance value is essen-
ll> Describe the normal physical and chemical properties o f tial because if it is high, the drug dosage must also be high
urine. and administered frequently to maintain a therapeutic level.
 Chapter 25 The Urinary System 1001
Creati nine, which has a C value of 140 ml/m in, is freely I retains _or adds needed substances, while removing wastes and
filtered but also secreted in small amounts. Nevertheless, its E ess ions. • • •••
measured blood value is often used to g ive an estimate of GFR.
This is convenie nt because creatinine does not need to be intra-
venously infused into the patie nt as does inulin. Urine
Che mica l Composition
.- HOMEOSTATIC
Water accounts for about 95% of urine volume; the remain-
~--A ~- IMBALANCE 25.4 ing 5% consists of solutes. T he largest compone nt of urine by
weight, apart from water, is urea, which is derived from the
Chronic renal disease, defined as a GFR of less than 60 ml/
normal breakdown of amino acids. Other nitrogenous ,vastes
min for at least three months, often develops silently over many
in uri ne include uric acid (an end product of nucleic acid
years. Filtrate formation decreases gradually, nitrogenous wastes
meta bolisrn) and cr eatinine (a metabolite of creatine phos-
accumulate in the blood, and blood pH drifts toward the acidic
phate, which is found in large amounts in skeletal muscle tissue
range. The leading cause of c hronic renal disease is diabetes
where it stores energy to regenerate ATP; <4 p. 304).
melli tus (44o/o of new cases), with hypertension a close second
Normal solute constituents of urine, in order of decreasi ng
(28% of new cases). Other causes include repeated kidney infec-
concentration, are urea, Na+, K', P0 43 -, S04 2-, creatinine, and
tions, physical trauma, and heavy metal poisoning.
uric acid. Much smaller but highly variable amounts of Ca2 +,
In r ena l failu r e (GF R < 15 ml/m in), fil trate formation
Mg2 +, and HC03- are also present.
decreases or stops completely. The clinical syndrome associ-
Unusually high concentrations of any solute, or the presence
ated with renal failure is called uremia (literally "uri ne in the
of abnormal substances such as plasma proteins, WBCs (pus),
blood") and includes fatigue, anorexia, nausea, mental changes,
or bile pigments, may indicate pathology (Table 25.3). (Nor-
and muscle cramps.
mal urine values are listed in Appendix F.)
Whi le uremia was once attributed to accumulation of nitro-
genous wastes (particularly urea), we now know that urea is not Ph ysical Characteristics
especially toxic. Rather, this multiorgan failure is caused by the
Color and Transparency Freshly voided urine is clear and
interplay of multiple factors. These include ionic and hormo-
pale to deep yellow. Its yellow color is due to u rochron1e (u'ro-
nal imbalances (i ncluding anemia due to lack of erythropoietin;
krom), a pigment that results when the body destroys hemo-
<4 p. 648), as well as metabolic abnormalities and accumulation
globin. The more concentrated the urine, the deeper the color.
of various toxic molecules that interfere with normal metabolis m.
An abnormal color (such as pink, brown, or a smoky tinge)
Current treatment opt ions are hemodialysis or a kidney may result from eat ing certain foods (beets, rhubarb) or from
transplant. Hemodialysis uses an "artificial kidney" apparatus,
the presence of bile pigments or blood in the urine. Additio n-
passi ng the patient's blood through a membrane tubing that is
ally, some commo nly prescribed drugs and vitam in supple-
permeable only to selected substances. The tubing is irn mersed
ments alter the color of urine. Cloudy urine may indicate a
in a solution that differs slightly from normal cleansed plasma.
uri nary tract infection.
As blood circulates through the tubi ng, substances such as
nitrogenous wastes and K+ present in the blood (but not in the Odor Fresh uri ne is slightly aromatic, but if allowed to stand,
bath) diffuse out of the blood into the surrounding solution. it develops an ammonia odor as bacteria metabolize its urea sol-
Meanwhile, substances to be added to the blood, mainly buff- utes. Some drugs and vegetables al ter the odor of urine, as do
ers for H+ (and glucose for malnourished patients), move from some diseases. For example, in uncontrolled diabetes mellitus
the bathing solut ion into the blood. In this way, hemodialysis the uri ne smells fruity because of its acetone content.

Table 25.3 Abnormal Urinary Constituents

SUBSTANCE NAME O F CONDITIO N POSSIBLE CA USES

Glucose Glycosuria Diabetes mellitus

Proteins Proteinuria, albuminuria Nonpatho logical: excessive physical exertion, preg nancy
Patholog ical (over 150 mg/day): glomerulo nephritis (see Related Clinical Terms on p. 1008),
severe hypertensio n, heart failure, often an initial sign of renal disease
Ketone bod ies Ketonuria Excessive formation and accu mulation of ketone bodies, as in starvation and unt reated
diabet es mellit us
Hemog lobi n Hemoglobinuria Vario us: t ransfusion reactio n, hemolytic anemia, severe burns, etc.
Bi le pig ments Bilirubinuria Liver disease (hepatitis, cirrhosis) or obstruction of bi le d ucts fro m liver or g allbladder
Erythrocytes Hematuria Bleeding urinary t ract (due to trauma, kidney stones, infection, or cancer)
l eu kocytes (pus) Pyuria Urinary tract infection
1002 UNIT 4 Mainten ance of the Body

pH Urine is usually slightly acidic (around pH 6), but changes

in body metabolism or diet may cause the p H to vary from
about 4.5 to 8.0. A predorninantly acidic diet that contains large
amounts of protein and whole wheat products produces acidic , , , - - Lumen
uri ne . A vegetarian (alkaline) diet, bacterial infection of the uri-
Mucosa
nary tract, or prolonged drai nage of the stomach via a nasogastric , , - - - • Transitional
tube all cause the urine to become alkaline . epithelium
k =-- · Lamina
Specific Gravity The ratio of the mass of a substance to the propria
mass of an equal volume of disti lled water is its specific gravity.
Because urine is water plus solutes, a given volurne has a greater Muscularis
mass than the same volume of distilled water. The specific grav- ' ~ - -· Longitudinal
layer
ity of distilled water is 1.0 and that of urine ranges frorn 1.00 I to
~ - - · Circular
1.035, depending on its solute concentration. layer

Check Your Understanding

19. What are the three major nitrogenous wastes excreted in the
urine?
20. j@;l4•)Gil What would you expect the normal clearance value
... - '

for amino acids to be? Explai n.

Figure 25.21 Cross section of the uret er wa ll (10x). The
==--------- For answers, see Answers Appendix. prominent mucosal folds seen in an empty ureter stretch and flatten
to accommodate large pulses of urine.

The ureters, bladder, and Practice Histology questions:

urethra transport, store, and MasteringA&P" > Study Area > Lab Tools> PAL

eliminate urine
Learning Outcomes The ureter plays an active role in transporti ng urine. Incom-
.... Describe the general location, structure, a nd funct ion of ing urine distends the ureter and sti mulates its muscularis to
the ureters, urinary bladder, and u rethra. co ntract, propelling urine into the bladder. ( Uri ne does not
.... Com pare the course, length, and functions o f t he male reach the bladder through grav ity alone.) The strength a nd
urethra w ith t hose of the fema le. frequency of the peristaltic waves are adjusted to the rate of
.... Define micturition a nd describe its neural control. urine formation. Both sympathetic and parasympathetic fibers
innervate each ureter, but neural control of peristalsis appears
The kidneys forrn urine continuously and the ureters transport it to be insignificant compared to the way ureteral smooth muscle
to the bladder. It is usually stored in the bladder until its release responds to stretch.
through the urethra in a process called micturition.

Ureters , HOMEOSTATIC
~-""J.r"- IMBALANCE 25.5
CLINICAi.!
The ureters are slender tubes that convey urine from the kidneys
to the bladder (Figures 25.1, 25.2, and 25.23). Each ureter begi ns Whe n calc ium or magnesium salts or uric acid are concentrated
at the level of L 2 as a co ntinuatio n of the renal pelvis. From in urine, they ,nay crystallize and precipitate as " kidney stones,"
there, it descends behind the peritoneum a nd runs obliquely or r en al calculi (kal 'ku-li; calculus = little stone). If stones
through the posterior bladder wal l. This arrangement prevents obstruct urine flow, they cause excruciati ng pain radiating from
backflow of urine because any increase in bladder pressure corn- the flank to the abdomen. Nephrolithiasis (litho = stone; iasis =
presses and closes the distal ends of the ureters. condition) refers to calculi in the kidney, whereas ureterolithi-
Histologically, the ureter wall has three layers (Figure 25.21). asis refers to calculi lodged in the ureters.
From the inside out: Whi le most small calculi pass without intervention, larger
• The 111.ucosa contains a transitional epithelium ( ~ p. 123) that calculi or those that lodge in the ureter can be removed endo-
is continuous with the mucosae of the kidney pelvis superi- scopically or surgically. Many calculi, especially those in the
orly and the bladder medially. kidney, can be treated by lithotripsy, a noninvas ive procedure
• The mu.scularis is composed chiefly of !\Yo smooth rnuscle that uses acoustic wave energy to break stones into smaller
sheets: the internal lo ngitudinal layer and the ex ternal c ir- pieces that may be passed more eas ily.
cular layer. An add itional smooth muscle layer, the external Obesity and elevated blood calcium are risk factors for the
longitudinal layer, appears in the lower third of the ureter. formation of renal calculi. The best way to prevent recurrence
is to maintain d ilute urine by adequate hydration. • •• • •
• The adventitia covering the ureter's external surface is typical
fibrous connective tissue.
 Chapter 25 The Urin ary System 1003
- - - - - - - - Peritoneum

- - - - - Rugae- --..,.
·- ~ - - - Detrusor - ,

- - - Adventitia

.------- - - - Trigone of bladder

~ - - - - , f-- - - Bladder neck----._,_

' - ~ 2 ~~ - -- - lnternal urethral - ~,

- sphincter
.....-- - - - - - Prostate
- -- ,' - - - - - - - Prostatic urethra
r-+-- - - - - - l n t e rmediate part
of the urethra
--r---::--- - - - Extemal urethral - - - - - -- - ---:c-
~ sphincter ~
r " " - - - Urogenital diaphragm ~ r e t h r a - -~ -~ JT-,
External urethral_;
orifice
(b) Female.

Figure 25.22 Structure of the urinary bladder a nd urethra.

The anterior wall of the bladder has been cut away to reveal t he
position of the t rigone.

Erectile tissue
of penis
(tri'gon; trigon = triangle), important clinically because infec-
tions tend to persist in this region.
The bladder wall has three layers: a mucosa containing
transitional epithelium, a thick muscular layer, and a fibrous
adventitia (except on its superior surface, where it is covered
by the peritoneum). The muscular layer, called the detrusor
(de-tru'sor; "to thrust out"), consists of intermingled smooth
(a) Male. The long male urethra has three regions: prostatic, muscle fibers arranged in inner and outer longitudinal layers
intermediate, and spongy.
and a middle circular layer.

Urine Storage Ca pacity

The bladder is very distensible. When empty, the bladder col-
Urinary Bladder lapses into its basic pyramidal shape. Its walls are thick and the
The urinary bladder is a smooth, collapsible, muscular sac mucosa is thrown into folds (rugae). As urine accumulates, the
that stores urine temporarily. bladder expands, becomes pear shaped, and rises superiorly in
the abdominal cavity. The transitional epithelium and muscu-
Urinary Bladder Anatomy lar wall stretch and thin, a nd rugae disappear. These changes
The bladder is located retroperitoneally on the pelvic floor just allow the bladder to store more urine without a significant rise
posterior to the pubic symphysis. The prostate (part of the male in internal pressure.
reproductive system) lies inferior to the bladder neck, which Usually, a moderately full bladder holds approximately 500
empties into the urethra. In females, the bladder is anterior to ml ( I pint) of urine. However, if urine flow has been partially
the vagina and uterus (see Figure 27 .15 on p. I061 ). obstructed for a long time (as by an enlarged prostate), the n
The interior of the bladder has openings for both ureters and the bladder can become e nlarged to hold more than t\vice that
the urethra (Figure 25.22). The smooth, triangular region of amount. When te nse with urine, it can be palpated well above
the bladder base outlined by these three openings is the trigone the pubic symphysis.
1004 UNIT 4 Maintenance of the Body

• The pros tatic ur ethra , about 2.5 cm ( I inch) long, ru ns

within the prostate.
Kidney • The intermediate part of the urethra (or me111branous u re-
thra), which runs through the urogenital cliaphragm, extends
about 2 cm from the prostate to the beginning of the penis.
Renal
• The spongy urethra, about 15 cm long, passes through the
pelvis penis and opens at its tip via the external urethral orifice.
The male urethra has a double function: It carries semen as well
as urine out of the body. We discuss its reproductive function in
Ureter Chapter 27.

J HOMEOSTATIC
~-""'Ji~- IMBALANCE 25. 6
Urinary tract infection (UTI) is a common problem, with up
to 50% of women experiencing a UTI during their lives. The
location of the female urethra near the anus predisposes it to
colonization by fecal bacteria. Sexual activity and the use of
spermicides increase the risk. In both men and women, cath·
eterization or any condition that obstructs the normal flow of
Urinary uri ne also predisposes an individual to acquiri ng a UTT.
bladder Infectio n of the urinary bladder, or cystitis, causes urinary
frequency (increased void ing) and dysuria (painful urination).
The urine may be blood tinged or foul smelling. If the infec-
Figure 25.23 Pyelogra m. This X-ray image was obtained using a tion ascends to the kidney, the more serious pyelonephritis (see
contrast medium to show the ureters, kidneys, and urinary bladder. p. 977) results, accompanied by fever, bac k pain, a nd other
(Artificially colored.) symptoms of systemic ill ness. After the offending bacteria are
identified by culturing the urine, antibiotics are used to treat
the problem. • •• • •
T he urinary bladder and ure ters can be seen in a special
X ray called a pyelogram (Figure 25.23). Micturition
Micturition (mik"tu-rish' un; mictur = urinate), also cal led uri-
Urethra nation or voiding, is the act of emptyi ng the uri nary bladder.
The urethra is a thin-walled muscular tube that drains urine For micturition to occur, three things must happen simultane-
from the bladder and conveys it out of the body. The epithe- ously: ( I) the detrusor must contract, (2) the internal urethral
lium of its mucosa) lining is mostly pseudostratified columnar sphincter must ope n, and (3) the external urethral sphincter
epithel ium. However, near the bladder it becomes transitional must open.
epithelium, and near the ex ternal opening it cha nges to a pro- The detrusor and its internal urethral sphincter are composed
tective stratified squamous epitheliurn. of smooth muscle and are innervated by both the parasympa-
At the bladder-urethra junction, the detrusor smooth muscle thetic and sympathetic nervous systems, which have opposing
thickens to fonn the internal urethral sphincter (Figure 25.22). actions. The external urethral sphincter, in contrast, is skeletal
This involuntary sphincter, controlled by the autonomic nervous muscle, and therefore is innervated by the somatic nervous
system, keeps the urethra closed when uri ne is not bei ng passed system.
and prevents leaking between voiding. How are the three events required for micturition coordi-
The external urethral sphincter surrounds the urethra as nated? Mic turition is most easily understood in infants where
it passes through the urogenital diaphragm. This sphincter is a spinal reflex coordinates the process. As urine accumulates,
formed of skeletal muscle and is volu ntarily controlled. The distension of the bladder activates stretch receptors in its walls.
Leva tor ani muscle of the pelvic floor also serves as a voluntary Irnpulses from the activated receptors travel via visceral afferent
constrictor of the urethra(~ Table 10.7, pp. 348-349) . fibers to the sacral region of the spinal cord. Visceral afferent
The length a nd functions of the urethra differ in the two impulses, relayed by sets of interneurons, excite parasympa-
sexes. In females the urethra is o nly 3-4 cm ( 1.5 inches) long thetic neurons and inhibi t sympathetic neurons (Figure 25.24).
and fibrous connective tissue bi nds it tightly to the anterior vag- As a result , the detrusor contracts and the internal sphincter
inal wall. Its exten1al opening, the external urethral orifice, opens. Visceral afferent impulses also decrease the firing rate
lies anterior to the vaginal opening and posterior to the cli toris. of somatic effere nts that normally keep the external urethral
In males the urethra is approximately 20 cm (8 inches) long sphincter closed. This allows the sphincter to relax so urine can
and has three regions. flow.
 Chapter 25 The Urinary System 1005

Brain
Higher brain
Urinary bladder centers
fills, stretching
bladder wall
Allow or inhibit micturition
as appropriate

Afferent impulses Pontine micturition Pontine storage

from stretch center oenter
receptors

Simple Promotes micturition Inhibits micturition by

spinal by acting on all three acting on all three spinal
reflex spinal efferents efferents

Spinal
cord I Spinal
cord
.
ifParasympathetic J !Tsympathe~ 1'5omatic mo; ; - i I Parasympathetic activity
L act,v,ty L _J
activity ~ erve actlVI~ t Sympathetic activity
t Somatic motor nerve activity
I
..
•
Detrusor contracts; External urethral
internal urethral sphincter opens
sphincter opens

I
l
Micturition ~ ................. ••••••••••••••••••
lnhib,'t s

Figure 25.24 Cont rol of micturit ion .

Between ages 2 and 3, descending circuits from the brain , HOMEOSTATI C

~-"'Ji~- IMBALANCE 25.7 CLINICAi!
have matured enough to begin to override reflexive urination.
The pons ( <1111 p. 452) has two centers that participate in control In adu lts, urinary incontinence (the inability to control uri-
of micturition. The pontine storage center inhibits micturition, nation) is usually a result of weakened pelvic muscles follow-
\vhereas the ponline ,nicturition center promotes this reflex. ing childbirth or surgery, physical pressure during pregnancy,
Afferent impulses from bladder stretch receptors are relayed or nervous system problems. In stress incontinence, a sudden
to the pons, as \veil as to higher brain centers that provide the increase in intra-abdom inal pressure (during laughing and
conscious awareness of bladder fullness. coughing) forces urine through the ex ternal sphincter. This
Lower bladder volumes primarily activate the pontine stor- condition is common during pregnancy when the heavy uterus
age center, which inhibits urination by suppressing parasympa- stretches the muscles of the pelvic floor and the urogenital dia-
thetic and enhanci ng sympathe tic output to the bladder. When phragm ( <1111 p. 348) that support the external sphincter. In over-
a person chooses not to void, reflex bladder contractions sub- flow incontinence, urine dribbles from the urethra whenever the
side within a minute or so and urine continues to accumulate. bladder overfills.
Because the external sphincter is voluntarily controlled, we In urinary retention, the bladder is unable to expel its
can choose to keep it closed and postpone bladder emptying contained urine. Urinary retentio n is common after general
temporarily. After additional urine has collected, the micturi - anesthesia (i t takes a little time for the detrusor to regain its
tion reflex occurs again and, if urinati on is delayed again, is activity). Urinary retention in men often reflects hypertrophy
damped once rnore. of the prostate, which narrows the urethra, making it difficult
The urge to void gradually becomes greater and greater, and to void. When urinary retention is prolonged, a slender drain-
micturition usually occurs before urine volume exceeds 400 ml. age tube called a catheter (kath'e-ter) must be inserted through
After normal micturition, o nly about 10 ml of urine remains in the urethra to drain the urine and prevent bladder trauma from
the bladder. excessive stretching. • •• • •
 Chapter 25 The Urinary System 1007
successively higher sources. Although the lower blood vessels • Autoso,nal recessive PKD is more severe and rnuch less corn-
usually degenerate, they sometimes persist, and for this reason mon, affect ing I in 20,000 people. Almost half of newborns
multiple renal arteries are common. The metanephric kidneys with recessive PKD die j ust after birth, and survivors gener-
excrete urine by the third month of fetal life, and most of the ally develop renal failure in early childhood. •• • •
amniotic fluid that surrounds a developing fetus is fetal uri ne.
Nonetheless, the fetal kid neys do not work nearly as hard as Because its bladder is very small and its kidneys are less
they will after birth because exchange through the placenta able to concentrate urine for the first two months, a newborn
allows the mother's urinary system to clear most of the undesir- baby voids 5 to 40 tirnes dai ly, depending on fluid intake. By 2
able substances from the fetal blood. months, infants void approximately 400 ml/day, and the a,nount
As the metanephros is developing, the cloaca subdivides to steadily increases u ntil adolescence, when adult urine output
form the future rectum and anal canal and the ur ogenital sin us, (about 1500 ml/day) is achieved.
into which the uri nary and genital ducts empty. The urinary Incontinence is normal in infants because their nervous sys-
bladder and the urethra then develop from the urogenital sinus tems have not matured enough to control the external urethral
(Figure 25.2Sb-d). sphincter. Reflex voidi ng occurs each tirne a baby's bladder fills
enough to activate the stretch receptors. Control of the volun-
, HOMEOSTATIC tary urethral sphincter goes hand in hand with nervous system
~--•.---~ IMBALANCE 25. 8 development. By 15 months, most toddlers know when they
have voided. By 24 months, some children are ready to begin
Three of the most common congenital abnormalities of the uri-
toilet trai ning. Daytime control usually is achieved first. It is
nary systern are horseshoe kidney, hypospadias, and polycystic
unrealistic to expect corn plete nighttime control before age 4.
kidney.
From childhood through late middle age, most urinary sys-
Whe n ascending from the pelvis the kidneys are very close
tem problems are infectious conditions. Escherichia coli bac-
together, and in 1 out of 600 people they fuse across the mid-
teria are normal residents of the digestive tract and generally
line, forming a sing le, U-shaped horseshoe kidney . T his con-
cause no proble,ns there, but these bacteria accou nt for 80o/o of
d ition is usually asymptomatic, but it may be associated with
all uri nary tract infections. Sexually trans,nitted infections can
other kid ney abnormalities, such as obstructed drainage, that
also inflame the urinary tract and clog some of its ducts. Child-
increase the risk of frequent kidney infections.
hood streptococcal infections such as strep throat and scarlet
Hypospadias (hi"po-spa'de-as), found in male infants only,
fever, if not treated promptly, may cause long-term inflamma-
is the most common congenital abnormal ity of the urethra. It
tory renal damage.
occurs when the urethral orifice is located on the ventral surface
Only about 3o/o of elderly people have histologically normal
of the penis. This problem may be corrected surgically when
kidneys, and kidney function declines with advancing age. The
the child is around 12 months old.
kidneys shrink as the nephrons decrease in size and number,
Polycystic kidney disease (PKD) is a group of disorders char-
and the tubule cells become less efficient. By age 80, the GFR
acterized by the presence of many fluid-filled cysts in the kid-
of healthy seniors is only about 70% that of young adults. Dia-
neys. These interfere with renal fu nction, ultimately leading to
betics are particularly at risk for renal disease, accounting for
renal failure. These disorders can be grouped into two general
almost half of new cases.
forms based on their pattern of inheritance:
Loss of bladder to ne w ith age can cause an an noyi ng
• Autoso,nal do1nina11t PKD, the less severe form, is much increase in frequency of micturition. Nocturia (nok-tu're-ah),
more com,non, affecting 1 in 500 people. The cysts develop the need to get up during the night to urinate, plagues almost
so gradually that they produce no symptoms until about 40 two-thirds of this population. Many people eventually experi-
years of age. ence incontinence, which can usually be treated with exercise,
medications, or surgery.

Bladder ca ncer Bladder cancer, three times more common in men common result of tearing of the pelvic floor muscles during
than in women, accounts for about 2% of al l cancer deaths. It childbirth.
usually involves neoplasms of the bladder's lining epithelium Cystoscopy (sis-tos' ko-pe; cyst = bladder; scopy = observation)
and may be induced by carcinogens from the environment Procedure in which a thin viewing tube is threaded into the
or the workplace that end up in urine. Smoking, exposure to bladder through the urethra to examine the bladder's mucosal
industrial chemicals, and arsenic in drinking water have also surface.
been linked to bladder cancer. B lood in the urine is a common
. . Diabetes insipidus (in-si'pT-dus; insipid= ta~teless, bland) See
warmng sign.
Chapter 16, p. 614. Nephrogenic diabetes insipidus is due to
Cystoc.ele (sis'to-sel; cyst = a sac, the bladder; ce/e = hernia, lack of ADH receptors in the collecting duct.
rupture) Herniation of the urinary bladder into the vagina; a
1008 UNIT 4 Maintenance of the Body

Glom erulonephritis (G N) (glo-mer"u-lo-nef-ri' tis) In flammation Nephrotoxi n A substance (heavy me tal, organic solvent, o r
of the g lomeruli, leading to increased permeabili ty of the bacterial toxin) that is toxic to the lcidneys.
filtratio n membrane. In some cases, circulating im1nune Noct ur nal e nuresis (NE) (en"u-re'sis) An inabili ty to control
complexes (antibodies bo und to foreign substances, such as urination at night d uring sleep; bed-wetting. In chi ldren
streptococcal bacteria) become trapped in the glome rular over 6, called primary NE if control has never been achieved
basement membranes. In other cases, immune responses and secondary NE if control was achieved an d then lost.
are mounted against o ne's own kidney tissues, leading to Secondary NE often has psychological causes. Primary NE is
glomerular dainage. In either case, the in flam1natory response more common and resulL~ from a combination of inadequate
that follows damages the filtration membrane, allowing blood nocturnal ADH productio n, unusually sound sleep, or a small
proteins and even blood cells to pass into the re nal tubules and b ladder capacity. Synthetic ADH often corrects the problem.
into the urine. As the osmotic pressure of blood drops, fluid
R enal infa r ct Area of dead, or necrotic, renal tissue d ue to
seeps from the b loodstream into the tissue spaces, causing
b lockage of the va~cular supply to the lcidney or hemorrhage.
bodywide edema Renal s hutdown requiring dialysis may occur
temporarily, but normal re nal function usually returns within A com1non cause of localized renal infarct is an obstructed
interlobar artery. Because interlobar arteries do not
a few months. If permanent g lomerular damage occurs, acute
anastomose, their obstruction leads to ischemic necrosis of
GN becomes chronic GN and ultimately renal failure resulL~.
the portions of the kidney they supply.
Intra,•enous pyelogr am (IVP) (pi'e-lo-gra1n; pyelo = lcidney pelvis;
Urologist (u-rol'o-jist) Physician who specializes in diseases
gmm = written) An X ray of the lcidneys and ureters obtained after
of urinary structures in bo th sexes an d in diseases of the
intravenous injection of a contrast mediu1n (as in Figure 25.23).
reproductive tract of males.

CHAPTER SUMMARY
nephrons are located near the cortex-medulla j unction, and their
fJ.11 The kidneys have three distinct regions and a rich nephron loops dip deeply into the medulla. Instead of directly
blood supply (pp. 975-978) forming peritubular capillaries, the efferent arterioles of many
Location and External Anat omy (p p. 975-976) j uxtamedullary nephrons fonn uniq ue bundles of straight
1. The paired lcidneys are retroperitoneal in the superior lumbar vessels, cal led vasa recta, that serve tubule segments in the
region . meduJla. Juxtainedullary nephrons and the vasa recta play an
2. A fibrous capsule, a perirenal fat capsule, and renal fascia important role in establishing the medullary osmotic gradie nt.
surround each lcidney. The perirenal fat capsule helps hold the 4. Collecting ducts receive urine from many nephrons and help
lcidneys in position . concentrate urine. They form the medullary pyramids.
5. The j uxtaglomerular complex is at the point of contact between
Internal Gross Anatomy (pp. 976-977) the afferent arteriole and the most distal part of the ascending
3. A kidney ha~ a superficial cortex, a deeper medulla consisting limb of the nephron loop. It consist~ of the granu lar cells, the
mainly of medullary pyramids, and a medial pelvis. Extensions macula densa, and extraglo me rular mesangial cells.
of the pelvis (calyces) surround and collect urine draining from
the apices of the medullary pyramids. fJ.il Overview: Filtration, absorption, and secretion are
the key processes of urine formation (p p. 983-984)
Blood and Nerve Supply (pp. 977-978)
1. Functions of the nephrons include g lomerular filtration, tubular
4. The kidneys receive 25% of the total cardiac output per minute. reabsorption, and tubular secretion. Via these functional
5. The vascular pathway through a lcidney is as follows: renal artery processes, the kidneys regulate the volume, composi tion, and
--> segmental arteries--> interlobar arteries--> arcuate arteries pH of the blood, and eli minate nitrogenous metabo lic wastes.
--> cortical radiate arteries--> afferent arterioles--> g lome ruti -->
efferent arterioles--> peritubular capillary beds--> cortical f).j1 Urine formation, step 1: The glomeruli make
radiate veins --> arcuate veins --> interlobar veins --> renal vein. filtrate (pp. 984-989)
6. The nerve supply of the kidneys is derived from the renal plexus.
1. The fi ltration me mbrane consists of the fenestrated g lomerular
f).fj Nephrons are the functional units of t he endothelium, the intervening basement membrane, and the podocyte-
kidney (pp. 978-983) containing visceral layer of the glomerular capsule. It permits free
pa~sage of substances smaller than (most) pla~ma proteins.
1. Nephrons are the structural and functional units of the kidneys.
2. The glomeruli function a~ filters. High glomerular blood
2. Each nephron consists of a glo me rulus (a high-pressure
pressure (55 mm Hg) occurs because the glomeruli are fed and
capillary bed), a glomerular capsule, and a re nal tubule that is
drained by arterioles, and the afferent arterioles are larger in
continuous with the capsule. S ubdivisions of the renal tubule
diameter than the efferent arterioles.
(from the glo me rular capsule) are the proximal convoluted
3. Abo ut o ne -fifth of the plasma flowing through the lcidneys is
tubule, nephron loop, and distal convoluted tubule. A second
filtered from the glomeruli into the g lomerular capsule.
capillary bed, the low-pressure peritubular capillary bed, is
4. Usually about IO mm Hg, the net filtration pressure (NFP) is
closely associated with the renal tubule of each nephron.
determined by the relationship between forces favoring fi ltration
3. The more numerous cortical nephrons are located almost
(glomerular hydrostatic pressure) and forces that oppose it
e ntirely in the cortex; only a s1nall part of their nephron loop
(capsular hydrostatic pressure and blood colloid osmotic pressure).
pene trates into the medulla. Glomeruli of j uxtamedullary
 Chapter 25 The Urinary System 1009
5. The glome rular filtration rate (GFR) is directly proportional to
the net filtration pressure and is about 125 ml/min (180 Uday).
lift The kidneys create and use an osmotic gradient
to regulate urine concentration and volume
6. Intrinsic renal contro l, o r renal autoregulation, e nables the
(pp. 995-1000)
kidneys to ma intain a re latively constant renal b lood flow and
glomerular fi ltration rate. Intrinsic control involves a myogenic 1. The graduated hyperosmo lality of the medullary fl uid~ (largeIy
mechanism and a tubuloglomerular feedback mechanism due to the cycling of NaCl and urea) ensures that the filtrate
mediated by the macula densa. reaching the distal convoluted tubule is dilute (hypo-osmolar).
7. Extrinsic control of GFR, via nerves and honno nes, maintains This allows urine wi th osmolalities ranging from 50 to 1200
blood pressure. Strong sympathetic nervous syste m activation mOsm to be formed .
causes constriction of the afferent arterioles, which decreases • The descending li mb of the nephron loop is permeable to
filtrate formation . water, which leaves the fi ltrate and enters the medullary
8. The renin-angiotensin-aldosterone mechanism raises systemic interstitial space. T he fi ltrate and medullary flu id at the bend
blood pressure by generating angiotensin II. Re nin is re leased of the ne phro n loop are hyperosmolar.
from granular cells in response to ( 1) direct sympathetic ne rvous • The ascending li mb is impermeable to water. Na• and c1-
system stimulation, (2) paracrines released by the macula densa, move out of the fi ltrate into the interstitial space, passively in
and (3) reduced stretch of granular cell membranes. the thin portion and actively in the thick portion. The filtrate
becomes more di Iu te.
• As filtrate flows through the collecting d uc ts in the inner
Complete an interactive tutorial: MasteringA&P" > Study Area>
medulla, urea diffuses into the interstitial space. F rom here,
Interactive Physiology> Urinary System> Glomerular Filtration
urea reenters the ascending th in limb and is recycled.
• The blood flow in the vasa recta is sluggish, and the
f).j.j Urine f ormation, step 2: Most of the filtrat e is contained blood equilibrates with the medullary interstitial
fluid . He nce, blood ex iting the medulla in the vasa recta
reabsorbed into t he blood (pp. 989- 994)
is nearly isotonic to blood plasma and the hig h solute
1. During tubular reabsorption, needed substances are removed concentration of the medulla is maintai ned .
from the fi ltrate by the tubule cells and returned to the peritubular 2. In the a bsence of antidiuretic hormone, di lute urine is formed
capillary b lood. T he primary active transport of Na• by a Na•-K• because the dilute filtrate reaching the collecting duct is simply
ATPase pump at the basolateral membrane accounts for Na• allowed to pass from the kidneys.
reabsorption and establishes the electrochemical gradient that 3. When extracellular flu id osmolality rises, blood levels of
drives the reabsorption of most other solutes and H20. Na• enters antidiure tic honno ne rise, and the collecting d uc ts become
a t the apical s urface of the tubule cell via faci litated diffusion mo re pe rmeable to water. \Yater moves out of the fi ltrate as it
through c hanne ls o r as part of a cotransport mechanism. flows thro ugh the hyperosmotic medullary areas. Consequently,
2. Passive tubular reabsorption is driven by electroche mical mo re concentrated urine is produced , and in smalle r amounts.
gradients established by active reabsorption of sodium ions.
Water, many ions, and various othe r substances (for example, f).j:j Renal funct ion is evaluated by analyzing blood
urea) are reabsorbed passively by diffusion via transcellular or and urine (pp. 1000-1002)
paracellular path ways.
3. Secondary active tubular reabsorption occurs by cotransport Renal Clearance (pp. 1000-1001)
wi th Na• via transport proteins. Transpo rt of s uch s ubstances is 1. Re nal clearance is the volume of plasma that is completely
limited by the number of carrie rs available. Actively reabsorbed cleared of a particular s ubstance per minute. Studies of renal
s ubstances include glucose, amino acids, and some ions. clearance provide in formation about renal function or the
4. The prox imal tubule cells are most active in reabsorption . M ost course o f renal disease.
of the nutrients, 65o/o of the water and sodium ions, and the 2. Re nal failure has serious consequences: T he kidneys are unable
bulk of actively transpo rted ions are reabsorbed in the proximal to concentrate urine, acid-base and e lectro lyte imbalances
convoluted tubules. occur, and nitrogenous wastes accumulate in the blood .
5. Reabsorption of additional sodium ions and water occurs in the
distal tubules and collecting duel~ and is hormonally contro lled. Urine (pp. 1001-1002)
Aldosterone increases the reabsorption of sodium; antidiure tic 3. Urine is 95% water; solutes include nitrogenous wastes (urea,
hormone (ADH) e nhances water reabsorption by the collecting uric acid , and creatinine) and various ions (always sodium,
d uc ts. pota~sium, s ulfate, and phosphate).
4. Substances not normally found in urine include g lucose, proteins,
Complete an interactive tutorial: MasteringA&P" > Study Area> erythrocytes, leukocytes, he moglobin, and bi le pigment~.
Interactive Physiology> Urinary System> Tubular Reabsorption 5. Urine is typically clear, yellow, aromatic, and slightly acid ic. Its
and Secretion specific gravity ranges from 1.001 to 1.035.

fftJ The ureters, bladder, and urethra transport, store,

fiff Urine f ormation, step 3: Certain substances are and eliminate urine (p p. 1002-1006)
secreted into the filtrat e (p p. 994-995) Ureters (p. 1002)
1. Tubular secretio n adds s ubstances to the filtrate (from the 1. The ureters are sle nder tubes running re troperitoneally from
blood o r tubule cells). It is an active process that is important each kidney to the bladder. They cond uct urine by peristalsis
in eliminating drugs, certain wastes, and excess ions and in from the re nal pelvis to the urinary bladder.
maintaining the acid-base balance of the blood.
1010 UNIT 4 Maintenance of the Body

Uri nary Bladder (pp. 1003-1004) inhibited), causing the detrusor to contract and the internal
2. The urinary bladder, which functions to store urine, is a urethral sphincter to open. In adults, pontine storage and
distensible muscular sac that lies posterior to the pubic symphysis. micturition centers can override this simple reflex.
It has two inlet~ (ureters) and one outlet (urethra) that outline the 9. Because the external sphincter is voluntari ly controlled,
trigone. In males, the prostate s urround~ the bladder outlet. micturition can usuaJly be delayed temporarily.
3. The b ladder wall consists of a mucosa containing transitional Developmental Aspects of the Urinary
epithelium; a three-layered detrusor; and an adventitia.
System (pp. 1006--1007)
Urethra (p. 1004) 1. Three set~ of kidneys (pronephric, mesonephric, and
4. The urethra is a muscular tube that conveys urine from the me tanephric) develop from the intermediate mesoderrn . The
me tanephros excretes urine by the third month of development.
bladder to the body exterior.
5. W here the urethra leaves the bladder, it is surroun ded by an 2. Common congenital abnormalities are horseshoe kidney,
internal urethral sphincter, an involuntary s1nooth muscle hypospadias, and polycystic kidney disease (PKO).
3. The kidneys of newborns are less able to concentrate urine;
sphincte r. Where it passes through the urogenital diaphragm, the
their b ladder is small and voiding is frequent. Neuromuscular
voluntary external urethral sphincter is formed by skeletal muscle.
6. In females the urethra is ~ c m long and conducts o n! y urine. maturation gene rally allows toi le t training for micturition to
begin by 24 months of age.
In ma les it is 20 cm long and conduc ts both urine and semen .
4. The most common urinary syste m pro blems in c hildren and
M icturit ion (pp. 1004-1005) young to middle-aged adult~ are bacterial infections.
5. With age, nephrons are lost, the filtration rate decreases, and
7. Micturition is e mptying of the b ladder.
tubule cells become less efficient at concentrating urine.
8. Accumulating urine stretches the bladder wal l, which initiates
6. Bladder capacity and tone decrease with age, leading to
the mic turition reflex . In in fants, this is a simple spinal reflex :
freque nt mic turition and (often) incontinence. Urinary retention
Parasympathetic fibers are excited (and sympathetic fibers
is a common problem of elderly men .

REVIEW QUESTIONS To access additional practice questions using your smartphone, tablet,
or computer: MasteringA&P• > Study Area > Practice Tests & Quizzes

Level 1 Remember/Understand 9. Which of the following is/are true about ADH? (a) It promotes
obligatory water reabsorption, (b ) it is secre ted in response to an
(Some questions have more than o ne correct answer. Select the best
inc rease in extracellular Auid osmolality, (c) it causes insertion
answer or answers from the c ho ices given.)
of aquaporins in the PCT, (d) it promotes Na• reabsorption .
1. The lowest blood concentration of nitrogenous waste occurs in 10. What is the importance of the perire nal fat capsule that
the (a ) hepatic vein, (b) inferior vena cava, (c) renal artery, surrounds the kidney?
(d) renal vein. 11. Defi ne micturition and describe the micturition reAex.
2. The g lome rular capillaries differ from o ther capillary networks
in the body because they (a) have a larger area of anastomosis, Level 2 Apply/Analyze
(b) are derived from and drain into arterioles, (c) are not made 12. Describe the mechanisms that contribute to renal autoregulation.
of endothelium, (d) are sites of filtrate formation . 13. Describe the mechanisms of extrinsic regulation of GFR, and
3. Damage to the renal medulla would interfere first with the their physiological role.
functioning of the (a) glo me rular capsules, (b) distal convoluted 14. Describe what is involved in active and passive tubular
tubules, (c) collecting duc ts, (d ) proximal convoluted tubules. reabsorption.
4. W hich is reabsorbed by the proximal convoluted tubule cells? 15. Explain how the peritubular capi llaries are adapted for receiving
(a) Na+, (b) K+, (c) amino acids, (d) all of the a bove. reabsorbed s ubstances.
5. Glucose is not normally found in the urine because it (a) does 16. Explain the process and purpose of tubular secretion.
not pass thro ugh the walls of the glo me rulus, (b ) is kept in the 17 . How does urinary b ladder anatomy s upport its storage function?
blood by colloid osmo tic pressure, (c) is reabsorbed by the 18. Describe the c hanges that occur in kidney and b ladder anatomy
tubule cells, (d) is removed by the body cells before the blood and physiology in o ld age.
reaches the kidney.
6. Filtration at the glo me rulus is inversely related to (a ) water Level 3 Evaluate/Synthesize
reabsorption, (b) capsular hydrostatic pressure, (c) arterial 19. Trace the pathway a creatinine molecule takes from a
blood pressure, (d) acidity of the urine. g lomerulus to the urethra. Name every mic roscopic or gross
7. Tubular reabsorption (a) of glucose and many other substances structure it passes through on its journey.
is a Tm-li mited active transport process, (b) of chloride is always 20. Explain the important diffe rences between blood plasma and
linked to the passive transport of Na+, (c) is the moveme nt of g lomerular fi ltrate, and relate the differences to the structure of
s ubstances from the b lood into the nephron, (d) of sodium the fi ltration me mbrane.
occurs o nly in the proximal tubule. 21. How does aldosterone modify the c hemical composition of urine?
8. If a freshly voided urine sam ple contains excessive ainounts 22. Explain why the filtrate becomes hypotonic as it Aows thro ug h
of urochrome, it has (a ) an ainmonia-like odor, (b) a pH below the ascending li mb of the nephron loop. Also explain why the
normal, (c) a dark yellow color, (d) a pH above nonnal . filtrate at the bend of the nephron loop (and the interstitial Auid
of the deep portions of the medulla) is hypertonic.
 Chapter 25 The Urinary System 1011
23. Mrs. Bigda, a 60-year-old woman, was brought to the hospital
by the police after falling to the pavement. She is found to c. The kidneys normally increase both their rate of fi ltra-
have alcoholic hepatitis. She is put on a salt- and protein- tion and reabsorption when blood pressure increases.
restricted diet and diuretics are prescribed to manage her ascites d. The kidneys release more renin when blood pressure
(accumulated Auid in the peritoneal cavity). How will diuretics increases, activating the renin-angiotensin-aldosterone
reduce this excess Auid? Name and describe the mechanisms pathway.
of action of three types of diuretics. Why is her diet salt- 3. Where and how do thiazide diuretics act in the kidneys
restricted? and how does this reduce blood pressure?
24. While repairing a frayed uti lity wire, Kevin, an experienced
At his two-week appointment at the outpatient clinic, Mr.
lineman, slips and fal ls to the ground. Medical examination
reveals a fracture of his lower spine and transection of the Boulard complains of fatigue, weakness, muscle cramps, and
lumbar region of the spinal cord. How will Kevin's micturition irregular heartbeats. A physical examination and Jab tests
produce the following observations:
be controlled from this point on? Will he ever again feel
the need to void? Will there be dribbling of urine between • BP 133/90, HR 75, temperature 37.3•c
voidings? Explain the reasoning behind all your responses. • Blood K+ 2.9 mEq/L (norrnal 3.5-5.5 mEq/L); blood Na•
25. Patty, aged 55, is awakened by excruciating pain that radiates 135 mEq/L (normal 135-145 mEq/L)
from her right abdomen to the Join and groin regions on the • Urine K• 55 mEq/L (normal <40 mEq/L); urine Na• 21
same side. The pain is not continuous but recurs at intervals mEq/L (norrnal >20 mEq/L)
of3 to 4 minutes. D iagnose her problem, and cite factors that
4. What is Mr. Boulard's main problem at this point?
might favor its occurrence.
26. Why are renal failure patients undergoing dialysis at risk for Explain how the thiazide d iuretic might have caused this
anemia and osteoporosis? What medications or supplements problem.
could you give them to prevent these problems? When asked about his medications, Mr. Boulard admits
that he did not fill his ACE inhibitor prescription because it
was too expensive. He could only afford the thiazide medica-
tions along with his insulin.
5. How do ACE inhibitors reduce blood pressure?
32-Year-Old Diabetic Male on a Diuretic 6. You've counseled Mr. Boulard about
how to prevent recurrence of his hypokalemia. Which
Let's return to Kyle
of the statements he makes indicates a need for further
Boulard, whom we
instruction?
met in the previous a. "There's lots of potassi um in fruits and vegetables, so
chapter. Mr. Boulard I' II eat more bananas."
has recovered from his
b. "There's lots of potassi um in regular table salt, so I'll
acute diabetic crisis. just use more."
The last update on c. ''I guess I need to come in regularly and have potas-
his chart before he is sium levels checked."
discharged includes d. "Maybe I should take a potassium supplement."
the following:
• BP I 50/95, HR 75,
temperature 37.2°C
• Urine: pH 6.9, negative for glucose and ketones; 24-hour
urine collection reveals 170 mg albumin in urine per day
Mr. Boulard is prescribed a thiazide diuretic and an angio-
tensin converting enzyme (ACE) inhibitor. He is counseled
o n the importance of taking his medications regularly and
keeping his outpatient follow-up appointments.
1. What is albumin? Is it normaJJy found in the urine? What
does its presence suggest?
2. Mr. Boulard's blood pressure is quite high.
How would this affect his nephron function? Choose the
statement below that is most accurate.
a. The kidneys normally increase their rate of filtration
when blood pressure increases.
b. The kidneys normally increase their rate of reabsorp-
tion when blood pressure increases.