Chapter 8

Urine formation by the

Kidneys
 functions of the kidneys
1. Excretion of metabolic waste products
and foreign chemicals
2. Regulation of water and electrolyte
balances
3. Regulation of acid-base balance
4. Regulation of body fluid osmolarity
and electrolyte concentrations
5. Regulation of arterial pressure
6. Secretion of hormones
 The endocrine function of
kidney
1. Renin
2. Erythropoietin(EPO)
3. 1,25-dihydroxycholecalciferol (1,25-
(OH)2-D3)
4. prostaglandins(PG)
5. kenins
 §1 Physiological anatomy of
the Kidney
1.1 General organization of the urinary system and the kidne

Kidney: paired
organs,about
fist sized, 150 g,
outside peritoneum
against the back.
Kidney Anatomy
 1.2 Nephron
 the basic functional unit of kidney
 1 million nephrons in each kidney
 The kidney cannot regenerate new
nephrons.
 glomerulus
renal corpuscle
Bowman capsule
Nephron

proximal tubule

thick segment of descending limb

renal tubule Loop of Henle thin segment of descending limb
thin segment of ascending limb
thick segment of ascending limb

distal tubule
(afferent arteriole)

(efferent arteriole)
1.3 Cortical nephron and
Juxtamedullary nephron
 Juxtamedullary nephron cortical nephron
Location close to renal medulla cortex
Proportion 10-15% 85-90%
Glomerular volume larger smaller
Loop of Henle long loop deep in medulla short loop
The ratio of the caliber 1:1 2:1
between afferent
arteriole and efferent
arteriole
Vasa recta + -
Juxtaglomerular apparatus few more
Major functions concentrate filtration ，
and dilute urine reabsorption
and secretion
 1.4
Glomerular capillary
membrane
1. Three major layers:
(1) capillary endothelium
(2) basement membrane
(3) epithelium (podocytes) of visceral
layer of Bowman’s capsule
 fenestrae
(fenestration

①
② capillary
endothelium
epithelium
 (1) capillary endothelium
 fenestrae(fenestrat
ion) 70-90nm
 Not act as a major
barrier for plasma
proteins
 (2) basement membrane
 Meshwork of
collagen and
proteoglycan fibrillae
that have spaces( 2-
8nm)
 Filter large amounts
of water and small
solutes, but
effectively prevent
filtration of plasma
proteins
 (3) epithelium (podocytes)
 surrounding the outer
surface of the capillary
basement membrane
 podocytes :long foot-
epithelium
like processes
 pedicels
 slit pores(filtration
slits) ： 25nm
 Provide some
restriction to filtration
 2. The filterability of solutes is
determined by their size and
electrical charge
1. Mechanical barrier: the selective filter of moleculal
weight(MW) :
MW>69 thousand impermeable
MW<69 thousandpermeable
2. Electrochemical barrier: the selective filter of
electric charge
(a layer of negative protein (saliva protein) located at
the surface of filtration membrane)
Molecule with positive charges permeable
Molecule with negative charges impermeable
 1.5 Juxtaglomerular
apparatus
Distribute in cortical nephron mainly.
Consist of Juxtaglomerular cell,
extraglomerular mesangial cell and
Macula densa.
Functions:
Macula densa can perceive the
change of Na+ concentrations in the
distal convoluted tubule .
Juxtaglomerular cell can release renin
when given a suitable stimulus.
Macula densa is a specialized group of epithelial
cells in the initial portion of the distal tubules that
comes in close contact with the afferent and
efferent arteriols.
Juxtaglomerular cell
is in the wall of the
afferent and efferent
arterioles, and can
secrete renin.

Extraglomerular
mesangial cell:
Phagocytosis ,contraction
 ↓ Arterial pressure Macula densa feedback
mechaniam

(-) ↓glomerular hydrostatic (-)

pressure

↓ GFR

↓ Macula densa
NaCl
↑ renin

↑ Angiotensin II

↑ Efferent arteriolar ↓ Afferent arteriolar

resisance resisance
 Renal blood supply
Renal artery→segmental arteries
→interlobar arteries→arcuate
arteries → interlobular arteries(radial
arteries)→ afferent arterioles
→glomerular capillaries characteristics of
→efferent arterioles renal blood supply:
→peritubular capillaries two capillaries beds

→ interlobular vein →arcuate vein

→interlobar vein →segmental vein
→renal vein.
Renal artery
interlobar arteries

arcuate arteries

interlobular arteries
 two capillaries beds
 glomerular capillaries:
Higher hydrostatic pressure( about 60
mmHg)
--- in favor of rapid fluid filtration ;
 peritubular capillaries:
Lower hydrostatic pressure ( about 13
mmHg)
---in favor of rapid fluid reabsorption;
 The formation of urine by
Kidney
1. glomerular
filtration
2. tubular
reabsorption
3. tubular secretion

Concentration and
dilution of urine Peritubular
capillary blood
Urinary excretion rate=filtration rate-reabsorption rate + secretion rate
 Glomerular filtration
 The first step in urine formation
 when blood flows into the glomerular
capillaries, the water bulk flow of
protein-free plasma filtrate into
Bowman’s capsule through the
glomerular membrane
 ultrafiltrate
 Most substances in
the plasma(except
protein)are freely
filtrated,so that their
concentrations in
Bowman’s capsule
are almost the same
as in the plasma.
 The ultra filtrate contains almost no
protein because the glomerular
membranes restrict the movement of
such high-molecule-weight substance
 GFR (glomerular filtration
rate)
the amount of ultra filtrate formed by two
kidneys per minute.
Normal value:125ml/min,180L/day

Filtration fraction
= GFR / Renal plasma flow
Normal value:about 20% (125/660=19%)
(about 20% of the plasma flowing through the
kidney is filtered by the glomerular capillaries)
The GFR is determined by (1)Effective
filtration pressure (EFP) and
(2)glomerular capillary filtration
coefficient(Kf)

GFR= Kf ☓ EFP
 Effective filtration

pressure,EFP
Represents the sum of the hydrostatic and
colloid osmotic forces that either favor or
oppose filtration.
 Forces favoring filtration:
Glomerular hydrostatic pressure(PG)
Bowman’s capsule colloid osmotic pressure (πB)=0
 Forces opposing filtration ：
Bowman’s capsule hydrostatic pressure (PB)
Glomerular capsule colloid osmotic pressure (πG)
 Glomerular Glomerular
hydrostatic colloid osmotic
Pressure Pressure
(60mmHg) (32mmHg)

Bowman’s
capsule
pressure
(18mmHg)

Glomerular Bowman’s Glomerular

EFP= hydrostatic
-capsule - colloid osmotic
Pressure pressure Pressure
(10mmHg)(60mmHg) (18mmHg) (32mmHg)
 Determinants of GFR
 1. EFP: ↑ GFR ↑
1) Glomerular hydrostatic pressure(PG) :
80~180mmHg, PG ↑ EFP - (fig)
<80 mmHg or ≻ 180mmHg, PG ↑ EFP ↑  GFR ↑
2)Bowman’s
2) capsule hydrostatic pressure (PB) : ↑ EFP↓
Example: oppress and block of ureter
3)Glomerular capsule colloid osmotic pressure (πG): ↑  EFP
↓  GFR ↓
Example:
drinking large quantities of water,decreased albumin;
πG ↓  EFP ↑  GFR ↑
Example :dehydration 脱水
 Determinants of GFR

2. Renal plasma flow (RPF)

RPF ↑
the velocity of πG increase ↓
 filtration areas↑
GFP ↑
 Determinants of GFR
3. glomerular capillary filtration coefficient(Kf)
Kf is the product of the permeability and
filtering surface area of the capillaries.
Kf ↑  GFR ↑
Example:
diabetes mellitus( 糖尿病 ) thickness of glomerular
membrane ↑  Kf ↓  EFP↓  GFR↓
 Renal Blood Flow and it’s
Regulation
Characteristics of RBF:
1. High blood flow:
1200ml/min: ¼ cardiac output
0.4 % of total body weight
A high blood flow is necessary for glomerular
filtration.
2.Distribution:
cortex  94%
outer medulla  5 - 6%
inner medulla  <1% 。
3. Determinants of renal blood flow

 △P = Renal artery pressure －

Renal vein pressure
 R total renal vascular resistance
 4. Physiological control of RBF
and GFR
1. Autoregulation of the RBF and GFR
（ 1 ） RBF is relatively constant when BP
fluctuating between 80 ～ 180 mm Hg
even if there are not regulations of nerve
and humoral factors. (Figure)
（ 2 ） Myogenic mechanism:
Tubuloglomerular feedback:
（ 3 ） significance:
to maintain a relatively constant GFR
to allow of renal excretion of water and
solutes under normal conditions.
 ΔP ΔP
Q= = 8ηL
R
ᅲᅲ
R4
 2. Nervous and Hormonal
Regulation of RBF
 Nervous regulation:
 Essentially all the blood vessels of the kidneys, including
both the afferent and efferent arterioles, are richly
innervated of the renal sympathetic nerve fibers.
 defense reaction, brain ischemia, or severe hemorrhage
 renal nerve (sympathetic nerve) activation  NE
releasing α-adrenoceptor  afferent arterioles
contraction  RBF↓ ， GFR ↓ (and the vessels of heart and
brain dilate, blood flow↑)
 Hormonal regulation:
Norepinephrine, Epinephrine ↑ 
afferent arterioles contract 
RBF↓ ， GFR ↓ (the vessels of
heart and brain dilate, blood
flow↑ )
Significance: in emergency 
reallocate of blood,and ensure the
blood supply of brain and heart.
Hormone or autacoid effect on
GFR
NE ↓
E ↓
Endothelin ↓
Angiotensin II — or ↓
NO ↑
Prostaglandins ↑
For many substances, reabsorption plays a much more important role than
does secretion in determining the final urinary excretion rate
 Reabsorption of Renal
tubule and Collecting duct
Conception: the process that some substances
(such as water, solutes) of the tubular fluid
are selectively reabsorbed from the tubules
back into the blood.
Tubular reabsorption is highly selective:
 G,aa,K+ all to be reabsorbed

 H O,Na+Cl-  most to be reabsorbed

2
 Urea  part to be reabsorbed

 Creatinine  not to be reabsorbed

 Filtration,reabsorption,and excretion
rate of different substances by the
kidneys
Amount amount amount % of
filterde
filtered reabsorbed excreted load
reasorbed
Glucose(gm/day) 180 180 0 100
Bicarbonate(mEq/day) 4,320 4,318 2
›99.9
sodium (mEq/day) 25,560 25,410 150
99.4
chloride(mEq/day) 19,440 19,260 180
99.1
urea(gm/day) 46.8 23.4 23.4 50
creatinine(gm/day) 1.8 0 1.8 0
 substance to be reabsorbed
must be transported
 across the tubular
epithelial
membranes into the
renal interstitial fluid
 through the
peritubular capillary
membrane back into
the blood
 The transporting pathways of
substance through the renal
tubular epithelial cells
 Transcellular
pathway:
through the cell
membranes
 Paracellular

pathway:
through the
junctional spaces
 Mechanisms of Reabsorption

1. Passive transport
1). Down electrochemical gradient;
2). not require energy;
3). Mode:Diffusion,Osmosis,facilitated
diffusion
4). Example:H2O
 2. Active transport
1). Against an electrochemical gradient;
2). require energy;
3). Depend on carrier proteins that penetrate
through the membrane
4). divided into two types:
 Primary active transport: coupled directly to an
energy source(hydrolysis of ATP)
 Secondary active transport :coupled indirectly
to an energy source(an ion gradient)
 Primary active transport is
linked to hydrolysis of ATP
 Importance: move solutes against an
electrochemical gradient
 energy source: hydrolysis of ATP
 Example: sodium-potassium ATPase
pump
Na+-K+ ATPase hydrolysis ATP  release energy 
Transport Na+ out of the cell into the interstitium

Transport K+ from the interstitium into the cell
The intracellular concentration of sodium is lower
 (chemical difference)
The cell interior is electrically negative than the
outside (electrical difference)

Favor Na+ to diffuse from the tubular lumen


into the cell through the brush border
 Secondary active transport

 Co – transport:
glucose-sodium transport
amino acids -sodium transport
phosphate -sodium transport
 Counter- transport:
H+-Na+ transport
 Glucose and Amino Acids are
reabsorbed by secondary active
transport
 They are actively transported across
the apical cell membranes of the
epithelial cells
 Their active transport depends on
the sodium gradient across this
membrane
 All other steps are passive
 Co – transport of
Glucose (or amino
Acids) along with
Sodium ions through
The brush border of
The tubular epithelial
cells
 GLUCOSE REABSORPTION HAS A
TUBULAR MAXIMUM
 Threshold for glucose:
the filtered load of
glucose at which
glucose first begins to
appear in the urine
 Transport maximum:
the maximum rate at
which glucose can be
reasorbed from the
tubules
 Passive water reabsorption by
osmosis is coupled mainly to
sodium reabsorption
 Solutes transported out of the tubule
their concentrations inside the tubule
their concentrations in the interstitium

create a concentration difference
 cause water reabsorption by osmosis from the

tubular lumen to the renal interstitium
 Prerequisite: the membrane is permeable to water
 Reabsorption of chloride,urea
and other solutes by passive
diffusion
Na reabsorption
+

H2O reabsorption

Lumen
Negative Luminal Cl- Luminal Urea
potential concentration concentration

Passive Cl- Passive Urea

reabsorption reabsorption
Reabsorption and secretion
along different parts of the
nephron
 Proximal tubule
 reabsorb :
about 65% of the filtered Na+, Cl- ,
HCO3- , K+ , H2O
essentially all the filtered glucose
and amino acids
 secrete:

Organic acids,bases and H+

Solute and water in the loop of
Henle
 The descending thin limb of the loop
of Henle:
o Highly permeable to water(20% filtered water
reabsorbed)
o Moderately permeable to most solutes,including
urea and sodium
 The thick ascending limb of the loop
of Henle:
o Reabsorb about 25% of the filtered loads of Na+, Cl-
and K+ (1Na+-2Cl--1K+ Co – transport )
o Reabsorb large amounts of Ca2+, HCO3- and Mg2+
o impermeable to water(the tubule fluid in the
ascending limb becomes very dilute)
 The thin ascending limb of the loop
of Henle:
much lower reabsorptive capacity than the thick
ascending limb
 Distal tubule

 Early distal tubule:

o Has many of the same
characteristics as the thick ascending
loop of Henle
o Reabsorb Na+, Cl- , Ca2+, and Mg2+
o impermeable to water and urea
 late distal tubule
and cortical collecting
bubule
 Have similar functional characteristics
 Composed of two distinct cell types:
o Principal cells: reabsorb Na+ and
water( controlled by the level of ADH)
secrete K+
o Intercalated cells: reabsorb K+
secrete H+ (role: regulate acid-
base balance)
 impermeable to urea
 Medullary collecting duct
 Reabsorb less than 10% of the
filtered water and sodium( controlled
by the level of ADH)
 permeable to urea
 secrete H+(effects: regulate acid-base
balance)
 Secretion of the Renal
tubules and collecting duct:
Conception: the process that the
epithelia of renal tubules and
collecting duct secrete their metabolic
products or substance of blood into
the tubular lumen.
Include:
1. The secretion of H+
2. The secretion of K+
3. The secretion of NH3
 1. The secretion of H+
Position: mainly in proximal tubule
Achieve by H+-Na+ antiport
Can promote the reabsorption of NaHCO3
Other position:Distal tubule and collecting
duct
H+-K+ ATPase ， hydrogen ATPase
Secreted H+ has three function:
　　　　 H+ + HCO3- 　→ H2CO3
H+ + NaHPO4- → NaH2PO4
H+ + NH3 → NH4+
 2 . The secretion of K+:
 Most of the daily variation in K+
secretion is caused by changes in K+
secretion in the distal tubule and
collecting duct
 Position: principal cells of distal
tubule and collecting duct
 Two steps of K+ secretion
• Uptake from the
interstitium into the
cell by Na+-K+ ATPase
• Passive diffusion from
the interior of the cell
into the tubular fluid
though K+ channels
 3 、 The secretion of NH3
 NH3 in the renal tubule is come from Glutamine
deamination
 NH3 enter tubule by ways of diffusion or NH4 -
Na+ antiport
 The secretion of H+ may promote the secretion
of NH3
 significance ： promote the secretion of H+
and the reabsorption of NaHCO3 , so play an
important role in keep the acid-base balance
 The concentration and
dilution
Hyperosmotic urine:
of urine
lack of water
→ concentrated urine
→ osmotic pressure can up to 1200mmol/L
Hypo-osmotic urine:
excess water
→ dilute urine
→ osmotic pressure can down to 40mmol/L
Isosmotic urine: may be renal failure
 1. Dilution of urine
Excess water in the body
 extracellular fluid osmolarity ↓
 ADH secretion ↓
 reduces the permeability of the distal tubules
and collecting ducts to water
 Solutes of tubular fluid are reabsorbed,
but water is not reabsorbed so much
 lead to dilution of urine.
 2. Concentration of urine
Water deficit in the body
 extracellular fluid osmolarity ↑
 ADH secretion ↑
 increase the permeability of the distal
tubules and collecting ducts to water
 large amounts of water is reabsorbed
 lead to concentration of urine
 Determinants of
concentration and dilution
The osmotic gradient in Medulla
 water absorption force
The level of ADH
ADH increases the permeability of
the distal tubules and collecting
ducts to water
 COUNTERCURRENT MAKES
THE OSMOTIC GRADIENT

 Outer medulla :
 Active reabsorption of NaCl at thick
ascending limb of loop of Henle;
 Inner medulla :
 Urea diffusing from collecting duct of
inner medulla
 NaCl diffusing from thin ascending limb of
loop of Henle;
THE OSMOTIC GRADIENT
CONCENTRATES THE URINE
WHEN ADH IS PRESENT
 Regulation of urine
formation
Regulative pathway: filtration, reabsorption,
and secretion.
1. Autoregulation
glomerulotubular balance
 The ability of the tubules to increase
reabsorption rate in response to increased
tubular load, even though the percentage of
glomerular filtrate reabsorbed in the proximal
tubule remains relatively constant at about
65% （ constant 　 fraction reabsorption ）
 glomerulotubular balance
 GFR: 125ml/min 150ml/min
The absolute rate of proximal tubular
reabsorption:
81ml/min 97.5ml/min
(65% of GFR) (65% of GFR)
 Importance: help prevent overloading of the tubular
segments when GFR increases.
 2. Humoral regulation:
 ADH
 RAAS
 Antidiuretic hormone, ADH
 Also called vasopressin,VP
 a small peptide with 9 aa;
 Synthetic site: supraoptic nucleus and
paraventricular nucleus of Hypothalamus
 Site of Storage: neurohypophysis (posterior pituitary)
 Site of action: the ADH receptor at distal tubule and
collecting duct
 effects: increase the water permeability of distal
tubule and collecting duct
 Results:More water is reabsorbed
Urine volume is reduced
Fluid is conserved in the body
 Regulation factors of ADH
1). Extracellular fluid osmolarity:
Water dificit  extracellular fluid osmolarity ↑
 excite osmorecepter of Hypothalamus
 excite ADH neurons
 posterior pituitary release ADH ↑
 water permeability of distal tubule and
collecting duct↑
 urine↓
Contrariwise , Extracellular fluid osmolarity ↓ 
urine ↑
 2). Blood volume
Blood volume↑
 Excite cardiopulmonary receptor

 vagi excitation
 Inhibit hypothalamus release ADH
 Urine↓
Contrariwise urine ↑
 3). Else

Baroreceptor excitation  ADH ↓

Pain, nausea, vomit ,ANGⅡ → ADH ↑
Disease of supraoptic nucleus and
paraventricular nucleus
 ADH ↓↓, this phenomenon is called
diabetes insipidus.
Renin-Angiotensin-
Aldosteron System

RAAS
 Renin:
Secreted by juxtaglomerular cells
 Angiotensin II:
the body’s most powerful sodium-
retaining hormone
 Aldosterone:
 Secreted by zona glomerulosa cells of adrenal
cortex
 Function:

improve of sodium reabsorption ,water

reabsorption and potassium secretion by
distal tubule and collecting duct
Directly stimulates sodium reabsorption,
especially in the proximal tubules
Increase the quantity of NE released by
sympathetic nerves ending.
Increased the vasoconstrictor tone of
sympathetic vasoconstrictor center.
Aldosterone function: increases the reabsorption of sodium and the
secretion of potassium in principal cells of the distal tubule,
Then increase the reabsorption of water by osmosis
Mechanism of alsosterone effects
 Regulation of secretion of
renin
 Baroreceptors of afferent arterioles
excite
 receptors of macula densa excite
 Renal sympathetic nerve excite
 PGE2,PGI2,NE and Adr  renin ↑
 3. Nervous regulation
 Renal sympathetic nerve: NE
1). Contraction of renal vessel→
RBF↓
→GFR ↓
2). Activate RAAS
3).improve the reabsorption of
Na+,Cl- ,water by proximal tubule
 Renal clearance
Definition:
The renal clearance of a substance is the volume of
plasma that is completely cleared of the
substance by the kidneys per unit time.
the ability of the kidneys to "clear" or remove a
specific substance from the blood.
Clearance equation:
C = U ×V / P （ ml/min ）
 UX×V ＝ GFR × PX － RX ＋ SX
 Urine
Straw yellow ；
Specific gravity: 1.015 - 1.025;
Osmotic pressure : urine > plasma ；
Acidity ： pH: 5.0 - 7.0 ;
Normal quantity of urine of adults:
1000 - 2000ml/ 24h, 1500ml.
>2500ml / 24h , called diuresis ；
100 - 500ml / 24h, called oliguria ；
<100ml /24h, called anuria.
 Urine Transportation,
Storage, and Elimination
 Basic pathway:  urine flows through papillary
ducts into the minor calyces → major calyces →
renal pelvis → ureters → the urinary bladder
→urethra
 Micturition is a two stage process involving the passive
storage and the active voiding of urine.
 The nervous control to micturition includes the
parasympathetic, sympathetic and somatic nervous
systems.
 THE URINARY BLADDER
STORES THE URINE
 Gravity and peristaltic contractions
propel the urine along the ureter
 Parasympathetic stimulation
contracts the bladder and micturition
results if the sphincters (internal and
external urethral sphincters) relax
 The external sphincter is under
voluntary control
 Anatomy
 a. trigone: a triangle-shaped area bounded
by the ureteral openings superiorly and the
opening to the urethra inferiorly.
                b. mucosa: transitional epithelium
+ connective tissue of the lamina propria; 
with rugae
                c. muscularis:  detrusor muscle; 3
layers of smooth muscle
                d. internal urethral sphincter:
formed by circular fibers, around the
opening to the urethra (smooth muscle)
                e. external urethral sphincter :
skeletal muscle
                f. adventitia / serosa
 micturition reflex

❶urine volume > 200-400 ml → bladder pressure↑

❷ → excites sensory stretch receptors in bladder
wall(the bladder feels "full" )
→conduct sensory signals by sensory nerve fiber of
pelvic nerves
❸ → sacral micrurition centers(S2-S4) and brain
❹ → parasympathetic nerve fiber of pelvic nerves to
the urinary bladder wall
→the detrusor muscle contract and the internal
urethral sphincter relax
→ urine enters posterior urethra
→ further excites sensory stretch receptors
in posterior urethra and bladder wall
→ further increase in reflex contraction of
bladder(self-regeneration,positive
feedback)
❺ (meanwhile of ❹) → inhibit pudendal
nerve under voluntary control → the
external urethral sphincter  diastole
❻ →urinate
 cystometrogram
 Cystometric study uses a device to
pump water into the bladder. The
device then measures the amount of
fluid present in the bladder when you
first feel the need to urinate, when
you are able to sense fullness, and
when your bladder is completely full.
 Facilitation or inhibition of
micturation by the brain
 Centers in the brain:
 pons: strong facilitory and
inhibitory centers in the brain stem
 several centers located in the
cebebral cortex: mainly inhibitory
 Higher center exert final
control of micturition
 The higher centers keep the micturition reflex partially
inhibited except when micturition is desired
 The higher centers can prevent micturition, even if the
micturition reflex does occur,by continual tonic contraction
of the external bladder sphincter until a convenient time
presents itself.
 When it is time to urinate,the cortical centers can facilitate
the sacral micturition centers to help initiate a micturition
reflex and at the same time inhibit the external urinary
sphincter so that urination can occur.
 Reabsorption of Na+
(1). About 65 - 70% proximal tubule;
20%  loop of Henle; 10% distal
tubule; 3%  collecting duct.
(2). Filtration of Na+ > 500g/D, excretion of
Na+ is 3 ~ 5 g/d, 99% of filtration is
reabsorbed.
(3). Pathway of reabsorption:
Transcellular pathway
Paracellular pathway
 The net reabsorption of sodium
ions involves at least three steps
(1). Sodium diffuses across the luminal membrane into the
cell down an electrochemical gradient
established by the sodium pump on the basolateral side of
the membrane.
(2). Sodium is transported across the memebrane into
interstitial fluid against an electrochemical gradient by
the sodium pump.
(3). Sodium is reabsorbed from the interstitial fluid into the
peritubular capillaries by ultrafiltration.
 The mechanism of Na+
reabsorption
 The first step of Na+ reabsorption in
different tubular segments has it’s own
characteristics.
 The second step and the third step of
Na+ reabsorption in the renal tubules
are similar.
 The mechanism of Na+
reabsorption
1. Early Proximal tubule :
Sodium enter into cells by way of:
(1) Na+-H+ antiport;
(2) Na+- G symport;
Na+- aa symport;
Na+- latic acid symport;
 2. Later Proximal tubule:

Sodium enter into cells by way of:

(1). Na+-H+ antiport;
(2). Paracellular pathway: 顺电势梯度被
动吸收
 3.Thick segment of loop
ascending limb
Sodium enter into cells by way of:
1- sodium, 2- chloride, 1- potasium symport
End result of three kinds of ions:
Na+ is pumped into interstitial fluid by sodium
pump and reabsorbed ;
Cl- diffuse into cell gap down electrochemical
gradient through Cl- channels :
K+ diffuse back to tubular lumen down
electrochemical gradient by K+ channels;
Furosemide can inhibit the symport of 1Na+-
2Cl-1K+ and play role in diuresis.
 4. Early Distal tubule:
Sodium enter into cells by way of:

 Na+- Cl- symport

 Thiazide （噻嗪类）
 5. Later Distal tubule and
Collecting Duct:
 Sodium diffuses across the apical
membrane into the Principal cell （
Sodium channel ）
 Reaborsoption of Na+ at distal
tubule and collecting duct
regulated by Aldosterone;
 Reabsorption of water:
 1. Quantity of reasorption:99%
 2. Passive reabsorption: osmotic pressure
 3. Position and siginificance:
 Proximal tubule:
 65-70%;
 Accompanied by the reabsorption of NaCl
 Has nothing to do with whether the body lack
water or not.
 Not regulated by hormones;
 Distal tubule: 10%;
 Collecting duct:10% ；
 Relation to whether body lack water
or not;
 Regulated by ADH and Aldosterone;
 Descending limb:10%;
 Reabsorption of HCO3- :
 About 80 ~ 85 per cent in proximal
tubule.
 Reabsorbed in a form of CO2 .
 Relate to the H+ secretion of tubular
epithelium.
 Significance:maintain the acid- base
balance of extracellular fluid.
Acetazolamide can inhibit the action
of Carbonic Anhydrase, diuretic
 Reabsorption of K+ :
All the filtrated K+ is reabsorbed,
and the K+ in the urine come from
excretion of distal tubule and
collecting duct .
Position: most at proximal tubule
Active transport.
 Reabsorption of glucose:
 Position: proximal tubule.
 All the filtrated glucose is
reabsorbed under normal condition.
 Secondary active transport,
accompanied by the primary active
transport of sodium .
Renal threshold for glucose:
 the maximal blood sugar
concentration which can not result in
glucosuria.
 Reasons: there is a limit to the
amount of transporter proteins and
binding site.
维持稳态的控制系统的组成要素
体液的组成
肾功能机制