Excretory Systems

• Functions of the excretory

systems
• Maintenance of proper internal
levels of inorganic solutes
• Maintenance of proper plasma
water volume
• Maintenance of osmotic balance
• Maintenance of acid-base
balance
• Removal of waste
• Evolution of basic excretory organs
• Simple aquatic animals depend
on diffusion and membrane
transporters.
• More complex aquatic animals
evolved specialized excretory
tissues with transport epithelia.
• Larger aquatic and all terrestrial
animals evolved specialized
tubules lined with transport
epithelia.
• Transport epithelia
• Na+/K+ ATPase pump on basolateral
membrane lowers Na+ concentration inside
the cell by pumping Na+ into the ECF.
• Na+ enters cell from apical side by diffusion
through ENaC channel.
• Cl– is attracted out of the cell through
chloride activated chloride (ClC) or cystic
fibrosis transmembrane-conductance
regulator (CFTR) channels by the charge
gradient produced by Na+ efflux.
• Increased extracellular solute
concentration attracts water across
basolateral membrane.
Salt absorption by a waterproof epithelium
Fluid absorption by a water-permeable epithelium
12.2 Renal Excretory Organs: Overview
• Basic processes taking place in renal
tubules
• Filtration
• Water and small solutes pass through a barrier while
cells and large molecules remain behind.
• Secretion
• Transport epithelia move specific solutes into the
tubule lumen for excretion.
• Reabsorption
• Transport epithelia move specific solutes and water
back into the body from the lumen.
• Osmoconcentration
• Water is removed from the lumen, leaving solutes
behind, producing a more concentrated excretory
fluid.
The principal functions of a tubular renal organ,
such as the vertebrate nephron
12.4 Vertebrate Urinary Systems
and Extrarenal Organs
• Vertebrate urinary
systems
• Kidneys are the urine-
forming organ.
• Paired organs on dorsal
side of abdominal cavity,
one on each side of the
vertebral column
• Blood is supplied by
renal artery and exits via
renal vein.
12.4 Extrarenal Organs Cont’d
• Vertebrate urinary systems cont’d
• Urine drains into two ureters.
• Ureters empty into urinary bladder (in
fish, amphibians, mammals), which
stores the urine, or hindgut (in reptiles
and birds).
• Bladder empties to the outside
through the urethra; hindgut empties
via cloaca.
Urinary system in two vertebrates
• Regions of the kidney
• Renal cortex - outer
• Renal medulla -
inner
• Medulla is divided
into renal pyramids
in larger mammals.
• Renal pelvis is the
drainage area in
center of kidney.
• Regions of the
kidney
• Nephron is the
functional unit of
the kidney.
• 1 million nephrons
in human kidney
• Consists of a
tubule and
associated
vascular
component
• Vasculature of the
nephron
• Afferent arteriole supplies
each nephron.
• Glomerulus is a ball-like knot
of capillaries in renal cortex -
site of filtration of the blood.
• Efferent arteriole exits the
glomerulus.
• Peritubular capillaries
surrounding the tubules
supply the renal tissue with
blood and exchange
materials with the tubular
fluid.
• Functional parts of the
renal tubule
• Bowman’s capsule --
glomerular filtration
• Proximal tubule -- tubular
reabsorption and
secretion
• Loop of Henle --
osmoconcentration
• Descending limb plunges
into medulla
• Ascending limb returns to
cortex
• Functional parts
cont’d
• Distal tubule -
reabsorption/secretion
and osmoconcentration
• Collecting duct -
osmoconcentration
• Empties into renal pelvis
• Juxtaglomerular
apparatus - sensor in
osmoregulation and
blood pressure regulation
12.5 Mammalian Urinary System:
Overview and Glomerular Filtration
• Cortical vs. juxtamedullary
nephrons
• Cortical nephrons
• Glomeruli in outer cortex
• Short loops of Henle dip only into outer
medulla.
• Juxtamedullary nephrons
• Glomeruli in inner cortex near the medulla
• Long loops of Henle go into inner medulla.
• Peritubular capillaries form hairpin
vascular loops (vasa recta).
Juxtamedullary and cortical nephrons of a
mammalian kidney
• Filtration apparatus
• Molecular sieve consists of
three layers:
1) Glomerular capillary wall
consists of a single layer of
flattened endothelial cells.
• Perforated with pores
• Pores are too small for
proteins to pass.
•Filtration apparatus
cont’d
2)Basement membrane is a
gelatinous layer
composed of collagen
and glycoproteins.
•Glycoproteins further
limit protein movement.
•Filtration apparatus
cont’d
3) Inner layer of
Bowman’s capsule
consists of podocytes
with filtration slits.
 Layers of the glomerular
membrane of a mammalian
kidney.
Bowman’s
capsule
podocytes
with foot
processes
and
filtration
slits of a
mammalian
kidney
• Glomerular filtration rate (GFR)
• GFR depends on net filtration
pressure, surface area and
permeability of glomerular
membrane
GFR = Kf x net filtration pressure
• Twenty percent of plasma that
enters the glomerulus is filtered.
• GFR in an adult human is 115 -
125 ml/min or ~180 liters/day.
• Controlled adjustments in GFR
• Increased resistance in afferent
arterioles will reduce blood flow to
glomerulus and lower GFR.
• Autoregulation prevents unintentional
shifts in GFR.
• Myogenic mechanism – stretch induced
contraction of arteriolar sm muscle
• Tubuloglomerular feedback via JGA
• increased salt delivery to DCT causes MD to release
ATP and angiotensin, wh/cause vasoconstriction of
afferent art.
Alteration
mechanisms for
glomerular
filtration rate
(GFR) in a
mammalian
kidney
The juxtaglomerular apparatus
in a mammalian kidney
• Controlled adjustments in GFR
cont’d
• Increased sympathetic activity
constricts afferent arterioles and
lowers GFR.
• Contributes to long-term
maintenance of blood pressure
by restoring plasma volume.
Baroreceptor
reflex
influence on
the GFR in
long-term
regulation of
arterial blood
pressure of a
mammal
Change in the
number of
open filtration
slits caused
by podocyte
relaxation and
contraction in
a mammalian
kidney
• Vasopressin controls water
reabsorption.
• Tubular fluid entering the distal tubule is
hypotonic (100 mOsm).
• This fluid empties into the collecting duct
which is bathed by progressively
increasingly concentrated interstitial fluid.
• Permeability of the collecting duct to water
depends on vasopressin (ADH).
• ADH triggers insertion of aquaporins into the
apical membrane of collecting duct cells.
• In dehydration, ADH secretion is increased
and water is reabsorbed from the collecting
duct.
• In overhydration, ADH secretion is low and
water is not reabsorbed.
Mechanism
of action of
vasopressin
in a mammal
Excretion of urine during a water deficit
Excretion of urine during a water excess
• Urea recycling contributes to
medullary hypertonicity.
• Top and middle portions of the
collecting duct are impermeable to
urea.
• Urea concentration in the collecting
duct increases as water is
reabsorbed.
• The lowest section of the collecting
duct is permeable to urea, so urea
diffuses out, increasing the solute
concentration of the medulla.
• Some urea is recycled into the loop of
Henle.
Urea recycling in the mammalian renal medulla
• The ability to osmoconcentrate the
urine varies among mammals.
• Weak osmoconcentrators have
primarily cortical nephrons (e.g.
beaver)

Photo courtesy of Michael S. Quinton

• Desert-dwelling
mammals (e.g.
kangaroo rat)
have all
juxtamedullary
nephrons with
very long loops
of Henle.
• The ability to
osmoconcentrate cont’d
•The “vertical” osmotic
gradient increases with
the length of the loop of
Henle.
•Countercurrent multiplier
is more active in small
mammals with higher
metabolic rates.
The relationship between relative medullary area in the
mammalian kidney (taken at the midline in sagittal section) and
the maximal urine concentration that can be produced.
12.9 Mammalian Urinary System:
Bladder Storage and Micturition
• Urine is temporarily stored in the urinary
bladder.
• Bladder wall is made of smooth muscle lined
by transitional epithelium.
• Umbrella cells are joined by tight junctions
rendering the bladder wall impermeable.
• Bladder can accommodate large fluctuations
in volume.
• Bladder opening is guarded by an internal
urethral sphincter (smooth muscle) and an
external urethral sphincter (skeletal muscle).
12.9 Bladder and Micturition cont’d
• Micturition (urination)
• Micturition spinal reflex
• Stretch receptors in the bladder wall are
stimulated by a distended bladder.
• Stimulate parasympathetic neurons
originating in lower spinal cord.
• Bladder wall contracts due to
parasympathetic stimulation,
mechanically pulling the internal urethral
sphincter open.
• External urethral sphincter opens and
urine is expelled through the urethra.
12.9 Bladder and Micturition cont’d

• Micturition (urination) cont’d

• Cortical control
• Signals reaching the cerebral
cortex stimulate the urge to
urinate.
• Voluntary contraction of the
external urethral sphincter can
delay urination.
Reflex and
higher-
cortical
control of
micturition
(mammal)
12.1 Evolution of Excretory Systems
• Nitrogenous wastes
• Result from the metabolism of proteins and
nucleic acids
• Choice of primary nitrogen waste
correlates with water availability
• Ammonia
• Most aquatic animals that breathe water
• Diluted to nontoxic concentrations
• Urea
• Most terrestrial animals
• More expensive metabolically, less toxic than
ammonia
• Uric acid
• Insects, reptiles and birds
• Most expensive metabolically, highly insoluble
General overview of nitrogen metabolism and excretion in animals.
12.4 Vertebrate Urinary Systems
and Extrarenal Organs
• Fish urinary systems
• Elasmobranches are
isosmotic or hyperosmotic
relative to seawater
• Retain urea and trimethylamine
oxide (TMAO) as osmolytes
• Rectal gland in hindgut
excretes hypertonic fluid high in
NaCl
12.4 Vertebrate Urinary Systems
and Extrarenal Organs cont’d
• Fish urinary systems continued
• Marine bony fishes are hypo-
osmotic
• Drink seawater to reverse water
loss through the gills
• Gills actively transport NaCl
outward and excrete nitrogenous
waste.
• Kidneys remove excess divalent
ions.
Osmoregulation in marine bony fishes
12.4 Vertebrate Urinary Systems
and Extrarenal Organs cont’d

• Fish urinary systems continued

• Freshwater bony fishes are
hyperosmotic
• Take in water through gills and
mouth
• Excrete a large volume of highly
dilute urine
• Gills take in NaCl and excrete NH3
and NH4+
Osmoregulation in freshwater bony fishes.
12.4 Vertebrate Urinary Systems
and Extrarenal Organs cont’d
• Amphibian urinary systems
• Demonstrate the transition to life on
land
• Lungs cannot excrete nitrogenous
wastes nor regulate NaCl
• Kidneys maintain a constant ECF
• Urinary bladder serves as a temporary
water reservoir in case of dehydration
• Arginine vasotocin (AVT) triggers
water uptake through aquaporins in
the bladder wall
12.4 Vertebrate Urinary Systems
and Extrarenal Organs cont’d
• Reptile urinary system
• Nephrons like those of aquatic verts
• Ureters carry urine in liquid or
semisolid form into the cloaca
• Lack a loop of Henle to help conserve
water
• Uric acid is the primary nitrogenous waste
• Cloaca and lower intestine can reabsorb
water
• Nasal salt glands secrete a highly salty
fluid
12.4 Vertebrate Urinary Systems
and Extrarenal Organs cont’d
• Avian urinary system
• Resembles reptiles’ urinary system
• Some mammalian-type nephrons with loops
of Henle further concentrate the urine
• Uric acid crystals are covered with protein
coats to form urate balls
• Marine birds have nasal salt glands located
near the eyes
• Contain blind-end tubules lined with active
salt secreting cells
• Excrete excess salt out of nasal passages
The salt glands of a gull.