Bio560 Osmoregulation

Osmoregulation Osmoregulation
and Excretion and Excretion

Water Balance and Waste Disposal Water Balance and Waste Disposal
Excretory Systems Excretory Systems
ntroduction ntroduction
nimal maintain a Iavorable proIile oI solutes nimal maintain a Iavorable proIile oI solutes
and solutions in their intracellular and and solutions in their intracellular and
extracellular body Iluids largely by means oI extracellular body Iluids largely by means oI
the epithelial tissues that Iorm the barrier with the epithelial tissues that Iorm the barrier with
the external environment. the external environment.
Most animal, ion and water balance is Most animal, ion and water balance is
regulated by renal tissues (kidney or kidney regulated by renal tissues (kidney or kidney- -
like tissue)as well as like tissue)as well as extrarenal extrarenal epithelial epithelial
tissues (gills, skins, digestive mucosa) tissues (gills, skins, digestive mucosa)
BIO 560 Osmoregulation BIO 560 Osmoregulation 3 3
Comparative Osmoregulation Comparative Osmoregulation
Water accounts for 60 Water accounts for 60 -- 3S of the weight of 3S of the weight of
animal animal
Water is found in various compartments in Water is found in various compartments in
animals animals -- contains dissolved solutes, including contains dissolved solutes, including
ions and nutrients ions and nutrients
!mportant for animals to maintain appropriate !mportant for animals to maintain appropriate
and correct amounts of water and solutes in and correct amounts of water and solutes in
these various compartments these various compartments
Osmoregulation Osmoregulation -- the process of regulating water the process of regulating water
and solute concentrations and solute concentrations
Comparative Osmoregulation Comparative Osmoregulation
Homeostasis
Osmoregulation
solute balance S gain or loss of water
Excretion
elimination of nitrogenous wastes
%egulating the !nternal Environment %egulating the !nternal Environment --
Water Balance Water Balance
Central to them process of osmoregulation is Central to them process of osmoregulation is
OSNOS!S OSNOS!S
Cells require a balance between water uptake Cells require a balance between water uptake
and loss and loss
Water may enter the body of terrestrial animal Water may enter the body of terrestrial animal
through food, drinking, and oxidative through food, drinking, and oxidative
metabolism metabolism
Water exits the body via evaporation and Water exits the body via evaporation and
excretion excretion
Aquatic animals are not affected by evaporation, Aquatic animals are not affected by evaporation,
but face the problem of osmosis where water but face the problem of osmosis where water
may enter (freshwater) or leave (marine) the may enter (freshwater) or leave (marine) the
body body
Even animals with specialized body coverings that retard Even animals with specialized body coverings that retard
water gain or loss have some unprotected structures water gain or loss have some unprotected structures
exposed to the environment for gas exchange (lungs, gills) exposed to the environment for gas exchange (lungs, gills)
Animals' cells cannot survive a net gain (swell and burst) Animals' cells cannot survive a net gain (swell and burst)
or loss (shrivel and die) of water or loss (shrivel and die) of water
Solutes in the blood help ensure the proper water balance Solutes in the blood help ensure the proper water balance
in the cell in the cell
Osmosis = diffusion of water across a selectively Osmosis = diffusion of water across a selectively
permeable membrane permeable membrane
occurs when two solutions separated by a membrane occurs when two solutions separated by a membrane
differ in osmolarity (total solute concentration) differ in osmolarity (total solute concentration)
if a selectively permeable membrane separates two if a selectively permeable membrane separates two
solutions of differing osmolarities, water flows from the solutions of differing osmolarities, water flows from the
hypotonic solution to the hypertonic solution hypotonic solution to the hypertonic solution
Hypertonic solution = when comparing two Hypertonic solution = when comparing two
solutions, a solution with a greater solute solutions, a solution with a greater solute
concentration, net water movement occurs into concentration, net water movement occurs into
the solution the solution
Hypotonic solution = when comparing two Hypotonic solution = when comparing two
solutions, the solution with lower solute solutions, the solution with lower solute
concentration, net water movement occurs out of concentration, net water movement occurs out of
the solution the solution
!sotonic solution = when comparing two !sotonic solution = when comparing two
solutions, a solution with a solute concentration solutions, a solution with a solute concentration
equal to that of the other solution, no net water equal to that of the other solution, no net water
movement occurs between the solutions movement occurs between the solutions
Concentration usually expressed in terms of mass (seawater = 3S Concentration usually expressed in terms of mass (seawater = 3S
g/L) g/L)
Solutes have physiological effects related to NUNBER OF Solutes have physiological effects related to NUNBER OF
PART!CLES, not mass PART!CLES, not mass -- expressed in terms of NOLES expressed in terms of NOLES
A molar solution has 1 mole solute/L of the final solution = 1N (or A molar solution has 1 mole solute/L of the final solution = 1N (or
mN) mN)
A NOLAL solution has 1 mole solute/kg water = 1m A NOLAL solution has 1 mole solute/kg water = 1m -- only 1 only 1
difference from molarity for seawater difference from molarity for seawater -- thermodynamically more thermodynamically more
correct but harder to measure correct but harder to measure
Osmotic pressure (OP) determines water movement across Osmotic pressure (OP) determines water movement across
membranes membranes
for non for nonelectrolytes = sum of the molarity (or molality) of the electrolytes = sum of the molarity (or molality) of the
different solutes different solutes
described as OSNOLAR!TY (Osmoles/L or mOsm/L) or described as OSNOLAR!TY (Osmoles/L or mOsm/L) or
OSNOLAL!TY (kg OSNOLAL!TY (kg
11
))
Electrolytes Electrolytes -- NaCl has twice the osmotic effect of glucose, as it has NaCl has twice the osmotic effect of glucose, as it has
twice as many particles twice as many particles
Osmotic %esponses of Animals Osmotic %esponses of Animals
Osmotic responses of animals can be classified Osmotic responses of animals can be classified
into two broad categories (depending on how into two broad categories (depending on how
they balance water loss with water gain) they balance water loss with water gain)
1. 1. Osmoconformers Osmoconformers
2. 2. Osmoregulators Osmoregulators
Osmoconformers Osmoconformers = animals that do not = animals that do not
actively adjust their internal actively adjust their internal osmolarity osmolarity
This includes many salt water animals This includes many salt water animals
Body fluids are isotonic with surroundings Body fluids are isotonic with surroundings
824.431472078 824.431472078- -isoosmotic isoosmotic with their with their
environment, most marine invertebrates environment, most marine invertebrates
- - internal internal osmolarity osmolarity nears the external nears the external
environment. (iI external change, internal environment. (iI external change, internal
change in parallel) change in parallel)
Strategies Ior ionic and osmotic
regulation
Osmoregulators Osmoregulators = animals that regulate = animals that regulate
internal internal osmolarity osmolarity by discharging excess water by discharging excess water
or taking in additional water or taking in additional water
!ncludes many saltwater animals, all freshwater !ncludes many saltwater animals, all freshwater
animals, and terrestrial animals animals, and terrestrial animals
Net movement of water in or out, requires a Net movement of water in or out, requires a
concentration gradient concentration gradient -- the maintenance of the maintenance of
which requires energy which requires energy
Osmoregulation Osmoregulation permits animals to live in a permits animals to live in a
variety of habitats, but the tradeoff is that it variety of habitats, but the tradeoff is that it
requires an energy expenditure by the animal requires an energy expenditure by the animal
82470:,9478 82470:,9478 - - regulate internal regulate internal osmolarity osmolarity
by releasing or taking in water, many marine by releasing or taking in water, many marine
animals, all Ireshwater animals, terrestrials animals, all Ireshwater animals, terrestrials
animals, and humans animals, and humans
- - maintain internal maintain internal osmolarity osmolarity within the within the
narrow range regardless oI the external narrow range regardless oI the external
environment environment
$trategies for ionic and osmotic $trategies for ionic and osmotic
regulation regulation
A large change in external A large change in external osmolarity osmolarity
is fatal to most animals, although is fatal to most animals, although
some can survive radical fluctuations some can survive radical fluctuations
&tenohaline &tenohaline animal animal = animal that = animal that
cannot survive a wide fluctuation in cannot survive a wide fluctuation in
external external osmolarity osmolarity
Euryhaline Euryhaline animal animal = animal that = animal that
can survive wide fluctuations in can survive wide fluctuations in
external external osmolarity osmolarity, may be: , may be:
Osmoconformer Osmoconformer
Osmoregulators Osmoregulators which minimize the which minimize the
osmotic shock by a variety of osmotic shock by a variety of
mechanisms mechanisms
Osmoregulation Osmoregulation in different environments in different environments
Each species has a range oI environmental osmotic Each species has a range oI environmental osmotic
conditions in which it can Iunction: conditions in which it can Iunction:
stenohaline stenohaline - - tolerate a narrow range oI salinities tolerate a narrow range oI salinities
in external environment in external environment - - either marine or either marine or
Ireshwater ranges Ireshwater ranges
euryhaline euryhaline - - tolerate a wide range oI salinities in tolerate a wide range oI salinities in
external environment external environment - - Iresh to saline: Iresh to saline:
short term changes: estuarine short term changes: estuarine - - 10 10 - - 32 32 ppt ppt, ,
intertidal intertidal - - 25 25 - - 40 40
long term changes: long term changes: diadromous diadromous Iishes Iishes
aintaining Water Balance !n aintaining Water Balance !n
Different Environments Different Environments
Narine animals live in a saline environment Narine animals live in a saline environment
consisting of about 36.S water and 3.S consisting of about 36.S water and 3.S
dissolved substances dissolved substances
The dissolved substances are collectively referred The dissolved substances are collectively referred
to as SALTS and the total amount of these to as SALTS and the total amount of these
dissolved substances in the water is SAL!N!TY dissolved substances in the water is SAL!N!TY
Nost marine animals have these problems: losing Nost marine animals have these problems: losing
body water to the ocean and picking up Na body water to the ocean and picking up Na
+ +
from from
the water the water
To solve these problems they drink seawater, To solve these problems they drink seawater,
produce very little urine and in order to get rid of produce very little urine and in order to get rid of
the salt, the salt is pumped using ATP across the the salt, the salt is pumped using ATP across the
gills gills
Nost marine invertebrates are osmoconformers Nost marine invertebrates are osmoconformers
Body fluids are isotonic to the environment Body fluids are isotonic to the environment
Body fluid composition usually differs from the Body fluid composition usually differs from the
external medium due to internal regulation of external medium due to internal regulation of
specific ions specific ions
Some vertebrates of the Class Agnatha (hagfishes) Some vertebrates of the Class Agnatha (hagfishes)
are also osmoconformers are also osmoconformers
Nost cartilaginous fishes, including sharks, Nost cartilaginous fishes, including sharks,
maintain internal salt concentrations lower than maintain internal salt concentrations lower than
seawater by pumping salt out through rectal seawater by pumping salt out through rectal
glands and through the kidneys, yet their glands and through the kidneys, yet their
osmolarity is slightly hypertonic to seawater osmolarity is slightly hypertonic to seawater
Sharks retain high level of urea as a dissolved Sharks retain high level of urea as a dissolved
solute in the body fluids solute in the body fluids
They also produce and retain trimethylamine oxide They also produce and retain trimethylamine oxide
(TNAO), which protects their proteins from (TNAO), which protects their proteins from
denaturation by urea denaturation by urea
Retention of these organic solutes (urea, TNAO) in Retention of these organic solutes (urea, TNAO) in
the body fluids makes them slightly hypertonic to the body fluids makes them slightly hypertonic to
seawater seawater
Sharks do not drink water, but balance osmotic Sharks do not drink water, but balance osmotic
uptake of water by excreting large volume of urine uptake of water by excreting large volume of urine
(also contains Na (also contains Na
+ +
))
Narine bony fishes are Narine bony fishes are
hypotonic to seawater hypotonic to seawater
Compensate for Compensate for
osmotic loss by osmotic loss by
drinking large amounts drinking large amounts
of seawater and of seawater and
pumping excess salt pumping excess salt
out with their gill out with their gill
epithelium epithelium
Excrete only a small Excrete only a small
amount of urine amount of urine
Gain of water and
salt ions from food
and by drinking
seawater
Osmotic water loss
through gills and other parts
of body surface
Excretion of
salt ions
from gills
Excretion of salt ions
and small amounts
of water in scanty
urine from kidneys
Freshwater animals are hypertonic to Freshwater animals are hypertonic to
their environment and constantly take their environment and constantly take
in water by osmosis, at the same in water by osmosis, at the same
time losing Na time losing Na
+ +
to water to water
Freshwater protozoa compensate Freshwater protozoa compensate
with contractile vacuoles that pump with contractile vacuoles that pump
out excess water out excess water
Nany freshwater animals, including Nany freshwater animals, including
fish, compensate by excreting large fish, compensate by excreting large
amounts of very dilute urine amounts of very dilute urine
Since salts are lost in this process, Since salts are lost in this process,
salt is replenished either by eating salt is replenished either by eating
substances with a higher salt content substances with a higher salt content
or, in the case of some fish, by active or, in the case of some fish, by active
uptake of Na uptake of Na
+ +
and Cl and Cl
from the from the

surrounding water by gill epithelium surrounding water by gill epithelium
Uptake of
water and some
ions in food
Osmotic water gain
through gills and other parts
of body surface
Uptake of
salt ions
by gills
Excretion of
large amounts of
water in dilute
urine from kidneys
Anadromous Anadromous fishes such as salmon are fishes such as salmon are
euryhaline euryhaline and migrate between seawater and migrate between seawater
and freshwater and freshwater
While in the ocean, they While in the ocean, they osmoregulate osmoregulate like like
other marine fishes other marine fishes
When in freshwater, they alter their When in freshwater, they alter their
osmoregulation osmoregulation to that of freshwater fishes to that of freshwater fishes
Other marine vertebrates such as reptiles Other marine vertebrates such as reptiles
and and birds have other special salt glands birds have other special salt glands
that secrete excess salt because they that secrete excess salt because they
cannot get rid of it in their urine cannot get rid of it in their urine
Narine mammals, on the other hand, can Narine mammals, on the other hand, can
produce a highly concentrated urine produce a highly concentrated urine
and so get rid of excess salt in their urine and so get rid of excess salt in their urine
Naintaining Water Balance !n Different Naintaining Water Balance !n Different
Environments Environments
Nasal salt gland
Nostril
with salt
secretions
Lumen of
secretory tubule
NaCl
Blood
flow
Secretory cell
of transport
epithelium
Central
duct
Direction
of salt
movement
Transport
epithelium
Secretory
tubule
Capillary
Vein
Artery
How does structure of
epithelial cells govern
osmoregulation?
Birds pump salt out of
blood
Freshwater fish pump
salts from water into blood
An albatross's salt glands empty via a duct into the nostrils,
and the salty solution either drips off the tip of the beak or
is exhaled in a fine mist
One of several thousand secretory tubules in a salt
excreting gland
Each tubule is lined by a transport epithelium surrounded
by capillaries, and drains into a central duct
The secretory cells actively transport salt from the blood
into the tubules
Blood flows counter to the flow of salt secretion. By
maintaining a concentration gradient of salt in the tubule
(aqua), this countercurrent system enhances salt transfer
from the blood to the lumen of the tubule.
Temporary waters present a special problem to animals that Temporary waters present a special problem to animals that
live in such environments live in such environments
ANHYD%OB!O&!& i ANHYD%OB!O&!& is an adaptation found in a small s an adaptation found in a small
number of aquatic invertebrates which permits them to number of aquatic invertebrates which permits them to
survive when their habitat dries up survive when their habitat dries up
Example Example -- tardigrades tardigrades (any minute invertebrates freshwater (any minute invertebrates freshwater
animal of the phylum animal of the phylum Tardigrada Tardigrada, having a short plump , having a short plump
body and four pairs of short legs) body and four pairs of short legs)
Hydrated animals are about 1 mm long and are about 8S Hydrated animals are about 1 mm long and are about 8S
water water
As the water around the animal disappears, water is lost As the water around the animal disappears, water is lost
from the tissues from the tissues
Tardigrades Tardigrades can survive many years in this state and will can survive many years in this state and will
rehydrate and become active when water returns rehydrate and become active when water returns
(a) Hydrated tardigrade (b) Dehydrated tardigrade
100 m
100 m
Dehydrated and frozen animals face the problem Dehydrated and frozen animals face the problem
of keeping their cell membranes intact of keeping their cell membranes intact
Researchers have found that dehydrated Researchers have found that dehydrated
anhydrobiotic animals contain large amounts of the anhydrobiotic animals contain large amounts of the
disaccharide trehalose along with other sugars disaccharide trehalose along with other sugars
Trehalose appears to replace the water associated Trehalose appears to replace the water associated
with membranes and proteins with membranes and proteins
Trehalose is also found in insects that survive Trehalose is also found in insects that survive
freezing freezing
The threat of desiccation is perhaps the largest regulatory The threat of desiccation is perhaps the largest regulatory
problem confronting terrestrial animals problem confronting terrestrial animals
Humans die if they lose about 12 of their body water Humans die if they lose about 12 of their body water
Adaptations that reduce water loss are key to survival on Adaptations that reduce water loss are key to survival on
land land
Nost terrestrial animals have body coverings that help Nost terrestrial animals have body coverings that help
prevent dehydration prevent dehydration
These include waxy layers in insect exoskeletons, the These include waxy layers in insect exoskeletons, the
shells of land snails, and the multiple layers of dead, shells of land snails, and the multiple layers of dead,
keratinized skin cells keratinized skin cells
Being nocturnal also reduces evaporative water loss Being nocturnal also reduces evaporative water loss
The excretory organs of terrestrial animals are adapted to The excretory organs of terrestrial animals are adapted to
conserve water while eliminating wastes conserve water while eliminating wastes
Land animals Land animals
manage their water manage their water
budgets by drinking budgets by drinking
and eating moist and eating moist
foods and by using foods and by using
metabolic water from metabolic water from
aerobic respiration aerobic respiration
ater
baIance in
a human
00 mL/day
= 100%)
ater
baIance in a
kangaroo rat
mL/day
= 100%)
ngested
in food 0.)
ngested
in food 0)
ngested
in liquid
100)
Derived from
metabolism 0)
Derived from
metabolism 1.8)
ater
gain
Feces 0.9)
Urine
0.4)
Evaporation 1.46)
Feces 100)
Urine
100)
Evaporation 900)
ater
Ioss
Desert animals Desert animals get major water savings from simple get major water savings from simple
anatomical features anatomical features
EXPER!NENT EXPER!NENT
Knut and Bodil SchmidtNielsen and their colleagues from
Duke University observed that the fur of camels exposed to
full sun in the Sahara Desert could reach temperatures of
over 70C, while the animals' skin remained more than
30C cooler
The SchmidtNielsens reasoned that insulation of the skin
by fur may substantially reduce the need for evaporative
cooling by sweating
To test this hypothesis, they compared the water loss rates
of unclipped and clipped camels
RESULTS RESULTS
Removing the fur of a
camel increased the
rate of water loss
through sweating by up
to S0
CONCLUS!ON
The fur of camels plays
a critical role in
their conserving water
in the hot desert
environments where
they live
Control group
Unclipped fur)
Experimental group
Clipped fur)
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Osmoregulation Osmoregulation depends on depends on
transport epithelia transport epithelia
Osmoregulators Osmoregulators utilize transport epithelia to regulate the utilize transport epithelia to regulate the
movement of solutes movement of solutes between their internal fluids and the between their internal fluids and the
external environment external environment
Usually a single sheet of cell, joined by impermeable tight Usually a single sheet of cell, joined by impermeable tight
junctions, facing the external environment junctions, facing the external environment
Nay be a channel that leads to the exterior through an Nay be a channel that leads to the exterior through an
opening on the body surface opening on the body surface
The The molecular composition of the epithelium's plasma molecular composition of the epithelium's plasma
membrane membrane determines the specific determines the specific osmoregulatory osmoregulatory functions functions
(remember that gill epithelium pumps salt out of marine (remember that gill epithelium pumps salt out of marine
fishes and pumps salts into freshwater fishes) fishes and pumps salts into freshwater fishes)
The transport epithelium in the nasal glands of marine bird The transport epithelium in the nasal glands of marine bird
are very efficient at eliminating the excess salts obtained are very efficient at eliminating the excess salts obtained
from drinking water from drinking water
Nay also function in excretion of nitrogenous wastes in some Nay also function in excretion of nitrogenous wastes in some
animals animals
Osmoregulation and Excretion Osmoregulation and Excretion
The The main waste main waste excreted by animals are excreted by animals are
wastes that have nitrogen in them, i.e. wastes that have nitrogen in them, i.e.
nitrogenous wastes nitrogenous wastes -- a byproduct of nucleic a byproduct of nucleic
acids and proteins metabolism acids and proteins metabolism
An animal's nitrogenous wastes reflect its An animal's nitrogenous wastes reflect its
phylogeny and habitat phylogeny and habitat
The type and quantity of an animal's waste The type and quantity of an animal's waste
products may have a large impact on its products may have a large impact on its
water balance water balance
!roteins Nucleic acids
Amino acids Nitrogenous bases
Amino groups
Most aquatic
animals including
most bony fishes
Mammals most
amphibians sharks
some bony fishes
Many reptiles
including
birds) insects
land snails
mmonia Urea Uric acid
O C
C
C
C
O

C O

Ammonia Ammonia
Freshwater Freshwater
Nost toxic Nost toxic
Urea Urea
Terrestrial Terrestrial
Uric acid Uric acid
Egg layers Egg layers
Nost water Nost water
conservative conservative
Node of reproduction Node of reproduction appears to appears to
have been important in choosing have been important in choosing
between the alternatives between the alternatives
(ammonia, urea, or uric acid) (ammonia, urea, or uric acid)
Soluble wastes can diffuse out Soluble wastes can diffuse out of a of a
shell shellless amphibian egg less amphibian egg
(ammonia) (ammonia) or be or be carried away by carried away by
the mother's blood the mother's blood in a mammalian in a mammalian
embryo embryo (urea) (urea)
However, the However, the shelled eggs of birds shelled eggs of birds
and reptiles and reptiles are are not permeable to not permeable to
liquids, liquids, which means that soluble which means that soluble
nitrogenous wastes trapped within nitrogenous wastes trapped within
the egg could accumulate to the egg could accumulate to
dangerous levels (even urea is dangerous levels (even urea is
toxic at very high concentrations) toxic at very high concentrations)
!n these animals, !n these animals, uric acid uric acid
precipitates out of solution precipitates out of solution and can and can
be be stored within the egg stored within the egg as a as a
harmless solid harmless solid left behind when the left behind when the
animal hatches animal hatches
Different animals Different animals excrete nitrogenous wastes in different excrete nitrogenous wastes in different
forms forms
Animals that excrete nitrogenous wastes Animals that excrete nitrogenous wastes as ammonia as ammonia (very (very
small molecule, very soluble, but highly toxic) small molecule, very soluble, but highly toxic)
Need access to lots of water Need access to lots of water
Release it Release it across the whole body surface across the whole body surface or through the or through the
gills gills
The The liver liver of mammals and most adult amphibians of mammals and most adult amphibians
Converts ammonia to less toxic urea Converts ammonia to less toxic urea (takes less energy (takes less energy
to make than uric acid) to make than uric acid)
Urea is Urea is carried to the kidneys carried to the kidneys, concentrated , concentrated
And And excreted with a minimal loss of water excreted with a minimal loss of water
!nsects, land snails, and many reptiles, including !nsects, land snails, and many reptiles, including
birds birds excrete excrete uric acid uric acid as their major nitrogenous as their major nitrogenous
waste waste
Uric acid is largely Uric acid is largely insoluble in water, non insoluble in water, nontoxic toxic (so (so
it it can be accumulated in high concentrations can be accumulated in high concentrations) and ) and
can be can be secreted as a paste with little water loss secreted as a paste with little water loss
The type of nitrogenous waste The type of nitrogenous waste also depends on also depends on
habitat habitat
For e.g., For e.g., terrestrial turtles terrestrial turtles (which often (which often lin lin dry areas) excrete mainly dry areas) excrete mainly
uric acid, uric acid, while while aquatic turtles aquatic turtles excrete both excrete both urea and ammonia urea and ammonia
!n some species, individuals !n some species, individuals can change their nitrogenous wastes can change their nitrogenous wastes
when environmental conditions change when environmental conditions change
For e.g., certain For e.g., certain tortoises tortoises that usually produce that usually produce urea shift to uric acid urea shift to uric acid
when when temperature increases and water becomes less available temperature increases and water becomes less available
Diverse excretory systems Diverse excretory systems are variations are variations
on a tubular theme on a tubular theme
Excretory systems Excretory systems regulate solute regulate solute
movement movement between internal fluids and the between internal fluids and the
external environment external environment
Nost excretory systems Nost excretory systems produce urine by produce urine by
refining a filtrate refining a filtrate derived from body fluids derived from body fluids
Excretory Processes Excretory Processes
Figure 44.9
iltration. The excretory tubule collects a filtrate from the blood.
Water and solutes are forced by blood pressure across the
selectively permeable membranes of a cluster of capillaries and
into the excretory tubule.
%eabsorption. The transport epithelium reclaims valuable
substances from the filtrate and returns them to the body fluids.
&ecretion. Other substances, such as toxins and excess ions,
are extracted from body fluids and added to the contents of the
excretory tubule.
Excretion. The filtrate leaves the system and the body.
Capillary
Excretory
tubule
F
i
l
t
r
a
t
e
U
r
i
n
e
1
2
3
4
Excretory Processes Excretory Processes
Key functions Key functions of most excretory systems are of most excretory systems are
Filtration Filtration pressure pressurefiltering of body filtering of body
fluids producing a filtrate fluids producing a filtrate
Reabsorption Reabsorption reclaiming valuable reclaiming valuable
solutes from the filtrate solutes from the filtrate
Secretion Secretion addition of toxins and other addition of toxins and other
solutes from the body fluids to the solutes from the body fluids to the
filtrate filtrate
Excretion Excretion the filtrate leaves the system the filtrate leaves the system
&urvey of Excretory &ystems &urvey of Excretory &ystems
The systems that perform basic excretory The systems that perform basic excretory
functions functions
vary vary widely among animal groups widely among animal groups
Are generally built on Are generally built on a complex a complex
network of tubules network of tubules
Protonephridia: lame Protonephridia: lame Bulb Bulb
&ystem &ystem
Found in Found in flatworms flatworms
Protonephridium Protonephridium = A network of = A network of closed tubules closed tubules
lacking internal openings lacking internal openings
They They branch branch throughout the body throughout the body
The smallest branches are capped by a cellular The smallest branches are capped by a cellular
unit called unit called a flame bulb a flame bulb
!nterstitial fluid passes !nterstitial fluid passes through a flame bulb and through a flame bulb and
is propelled by a tuft of cilia (in the flame bulb) is propelled by a tuft of cilia (in the flame bulb)
along the branched system of tubules along the branched system of tubules
Nucleus of cap cell
Cilia
nterstitial fluid filters through
membrane where cap cell
and tubule cell interdigitate
interlock)
Tubule cell
Flame bulb
Nephridiopore in body wall
Tubule
!rotonephridia tubules)
Protonephridia: lame Protonephridia: lame Bulb Bulb
&ystem &ystem
!n !n planaria planaria, , this fluid drains into excretory duc this fluid drains into excretory ducts ts
that empty out of the body that empty out of the body through numerous through numerous
nephridiopores nephridiopores
Transport epithelium lining the tubules Transport epithelium lining the tubules function in function in
osmoregulation osmoregulation by by absorbing salts absorbing salts before the fluid before the fluid
exits the body exits the body
Some parasitic flatworms are isotonic to their Some parasitic flatworms are isotonic to their
hosts and this closed system is hosts and this closed system is used mainly to used mainly to
excrete nitrogenous wastes excrete nitrogenous wastes
Protonephridia Protonephridia are also found in other are also found in other
invertebrates and lancets invertebrates and lancets
etanephridia etanephridia
Each segment of most annelids, including Each segment of most annelids, including
earthworms, contains earthworms, contains a pair of a pair of metanephridia metanephridia, ,
excretory excretory tubules that have internal openings to tubules that have internal openings to
collect body fluids collect body fluids
Coelomic Coelomic fluid enters fluid enters the funnel the funnelshaped shaped
nephrostome nephrostome which is surrounded with cilia which is surrounded with cilia
The fluid The fluid passes through the passes through the metanephridium metanephridium and and
empties empties into a storage bladder into a storage bladder that that empties empties
outside the body outside the body through the through the nephridiopore nephridiopore
BIO 560 BIO 560 Osmoregulation Osmoregulation 47 47
Nephrostome
Netanephridia
Nephridiopore
Collecting tubule
Bladder
Capillary network
Coelom
etanephridia etanephridia
The The nephrostome nephrostome collects collects coelomic coelomic fluid fluid from the from the
body segment just anterior body segment just anterior
A network of capillaries envelopes each A network of capillaries envelopes each
metanephridium metanephridium
These These capillaries reabsorb essential salts pumped capillaries reabsorb essential salts pumped
out of the collecting tubules by transport epithelium out of the collecting tubules by transport epithelium
bordering the lumen bordering the lumen
Excretion of hypotonic, dilute urine offsets the Excretion of hypotonic, dilute urine offsets the
continual osmosis of water continual osmosis of water from damp soil across from damp soil across
the skin the skin
alpighian Tubules alpighian Tubules
Nalpighian Nalpighian tubules tubules = = excretory organs of insects and other excretory organs of insects and other
terrestrial arthropods terrestrial arthropods
!nsects produce a relatively !nsects produce a relatively dry waste matter dry waste matter -- an an
important adaptation to terrestrial life important adaptation to terrestrial life
Remove nitrogenous wastes from the Remove nitrogenous wastes from the haemolymph haemolymph and and
function in function in osmoregulation osmoregulation
Are Are outpocketings outpocketings of the gut that open into the digestive of the gut that open into the digestive
tract at the tract at the midgut midguthindgut juncture hindgut juncture
The The tubules dead tubules deadend at the tips end at the tips away from the gut and away from the gut and
are bathed in the are bathed in the haemolymph haemolymph
Transport epithelium Transport epithelium lining each tubule lining each tubule moves solutes moves solutes
(salts and nitrogenous wastes) (salts and nitrogenous wastes) from the from the haemolymph haemolymph
into the tubule's lumen into the tubule's lumen
Digestive tract
Midgut stomach) Malpighian tubules
Rectum
ntestine
indgut
$aIt, water, and
nitrogenous
wastes
Feces and urine
Anus
Malpighian
tubule
Rectum
Reabsorption of H
2
O,
ions, and vaIuabIe
organic moIecuIes
EMOLYM!
alpighian Tubules alpighian Tubules
Accumulates nitrogenous wastes from the Accumulates nitrogenous wastes from the
haemolymph haemolymph and water and water follows by follows by
osmosis osmosis
The fluid in the tubule The fluid in the tubule then passes then passes
through the hindgut to the rectum through the hindgut to the rectum
Salts and water are reabsorbed across the Salts and water are reabsorbed across the
epithelium of the rectum epithelium of the rectum and and dry dry
nitrogenous wastes nitrogenous wastes are are excreted with excreted with
faeces faeces
ertebrate Kidneys ertebrate Kidneys
The kidneys The kidneys of most vertebrates are of most vertebrates are
compact organs with many excretory compact organs with many excretory
tubules tubules
The structure includes a The structure includes a dense capillary dense capillary
network network intimately associated with the intimately associated with the
tubules tubules
The tubules function in both excretion and The tubules function in both excretion and
osmoregulation osmoregulation
variations of the basic system are found variations of the basic system are found
amongst the vertebrate classes amongst the vertebrate classes
ertebrate Kidneys ertebrate Kidneys
vertebrate kidneys have vertebrate kidneys have six roles six roles in in
homeostasis homeostasis
!on balance !on balance
Osmotic balance Osmotic balance
Blood pressure Blood pressure
pH balance pH balance
Excretion Excretion
Hormone production Hormone production
The ammalian Excretory The ammalian Excretory
&ystem &ystem
The human (mammalian) kidneys are a pair of The human (mammalian) kidneys are a pair of
bean beanshaped organs ~ 10 cm long with two shaped organs ~ 10 cm long with two
layers: cortex and medulla layers: cortex and medulla
(b) Kidney structure
Ureter
Section of kidney from a rat
Renal
medulla
Renal
cortex
Renal
pelvis
&ystem &ystem
Blood enters Blood enters each each
kidney kidney via the renal via the renal
artery artery and and exits via exits via
the renal vei the renal vein n
About 20 of the About 20 of the
blood pumped by each blood pumped by each
heartbeat passes heartbeat passes
through the kidneys through the kidneys
&ystem &ystem
Urine exits Urine exits each kidney each kidney
through a through a ureter ureter and and
both both drain into drain into a a
common common urinary urinary
bladder bladder
Urine leaves the body Urine leaves the body
from the urinary from the urinary
bladder through the bladder through the
urethra urethra
Sphincter muscles Sphincter muscles near near
the junction of the the junction of the
urethra and bladder urethra and bladder
control urination control urination
The Nephron The Nephron
The functional unit The functional unit of of
the kidney the kidney
Composed of multiple Composed of multiple
tubules tubules
Each Each nephron nephron
consists of a single consists of a single
long tubule and a long tubule and a
ball of capillaries ball of capillaries
called the called the
glomerulus glomerulus
Surrounded by a specific vasculature Surrounded by a specific vasculature
BIO 560 BIO 560 Osmoregulation Osmoregulation 59 59
uxta-
medullary
nephron
Cortical
nephron
Collecting
duct
To
renal
pelvis
Renal
cortex
Renal
medulla
0 m
Afferent
arteriole
from renal
artery
Glomerulus
Bowman's capsule
!roximal tubule
!eritubular
capillaries
SEM
Efferent
arteriole from
glomerulus
Branch of
renal vein
Descending
limb
Ascending
limb
Loop
of
enle
Distal tubule
Collecting
duct
(c) ephron
Vasa
recta
(d) FiItrate and
bIood fIow
Each region contains many microscopic Each region contains many microscopic nephrons nephrons
and collecting ducts and collecting ducts
Pathway of the iltrate Pathway of the iltrate
Blood enters Blood enters the the glomerulus glomerulus via the via the afferent afferent
artery, artery, and and leave leave the the glomerulus glomerulus via the via the efferent efferent
artery artery
Filtrate enters the Filtrate enters the lumen of the Bowman's lumen of the Bowman's
capsule capsule
The filtrate then passes through the The filtrate then passes through the proximal proximal
tubule, tubule, the the loop of loop of Henle Henle and the and the distal tubu distal tubule, le,
which empties into a which empties into a collecting duct collecting duct
Filtrate, now called urine, flows from the Filtrate, now called urine, flows from the
collecting ducts into the collecting ducts into the renal pelvis, before renal pelvis, before
draining into the draining into the ureter ureter
Pathway of the iltrate Pathway of the iltrate
!roximal tubule
FiItrate
O
Salts NaCl and others)
CO
3
Urea
Glucose; amino acids
Some drugs
Key
Active transport
!assive transport
CORTEX
OUTER
MEDULLA
NNER
MEDULLA
Descending limb
of loop of
enle
Thick segment
of ascending
limb
Thin segment
of ascending
limb
Collecting
duct
NaCl
NaCl
NaCl
Distal tubule
NaCl Nutrients
Urea
O
NaCl
O CO
3
N
3
CO
3
O
1 4
3 2
3
5
Two types of Two types of nephrons nephrons are found in mammals and are found in mammals and
birds: cortical and birds: cortical and juxtamedullary juxtamedullary nephrons nephrons
Cortical Cortical nephron nephron = = Nephrons Nephrons that that have reduced have reduced
loops of loops of Henle Henle and are and are confined to the renal cortex confined to the renal cortex
80 of the 80 of the nephrons nephrons in humans are cortical in humans are cortical nephrons nephrons
]uxtamedullary ]uxtamedullary nephrons nephrons= = Nephrons Nephrons that have that have
long loops that extend into the renal medulla long loops that extend into the renal medulla
20 of the 20 of the nephrons nephrons are are juxtamedullary juxtamedullary nephrons nephrons
Urine Production Urine Production
The The nephron nephron and collecting duct are and collecting duct are lined lined
with transport epithelium with transport epithelium that processes that processes
the filtrate into urine the filtrate into urine
About 1100 About 11001200 L of blood flow through 1200 L of blood flow through
the human kidneys each day the human kidneys each day
The The nephrons nephrons process 180 L of filtrate process 180 L of filtrate per per
day, and the transport epithelium day, and the transport epithelium
processes this filtrate processes this filtrate to form the ~ 1.S L to form the ~ 1.S L
urine excreted daily urine excreted daily
Urine Production Urine Production
The composition of blood is regulated by The composition of blood is regulated by
transport epithelia of the nephrons and transport epithelia of the nephrons and
collecting ducts through: collecting ducts through:
Filtration Filtration
Reabsorption Reabsorption
Secretion Secretion
iltration of the Blood iltration of the Blood
Filtration occurs as Filtration occurs as blood pressure forces blood pressure forces fluid fluid
from the blood in the from the blood in the glomerulus glomerulus into the lumen into the lumen
of Bowman's capsule of Bowman's capsule
Liquid components of the blood escape into Liquid components of the blood escape into
Bowman's Bowman's capsule capsule blood cells and large blood cells and large
macromolecules are not filtered macromolecules are not filtered
Filtration of small molecules is Filtration of small molecules is nonselective nonselective
Filtrate at this point Filtrate at this point contains a mixture of glucose, contains a mixture of glucose,
salts, vitamins, nitrogenous wastes, and small salts, vitamins, nitrogenous wastes, and small
molecules molecules in concentrations similar to that in blood in concentrations similar to that in blood
plasma plasma
iltration of the Blood iltration of the Blood
lomerular lomerular capillaries are very leaky capillaries are very leaky
(porous) (porous)
Podocytes Podocytes (specialized cells of the (specialized cells of the
capsule) with capsule) with foot processes foot processes form the form the
filtration structure filtration structure
Nesangial Nesangial cells cells control blood pressure and control blood pressure and
filtration within the filtration within the glomerulus glomerulus
%eabsorption %eabsorption
Proximal tubule Proximal tubule --
reabsorption reabsorption begins begins
Loop of Loop of Henle Henle
(descending limb, (descending limb,
the loop, ascending the loop, ascending
limb) limb) -- most most
reabsorption reabsorption occurs occurs
Distal tubule Distal tubule --
reabsorption reabsorption is is
completed for most completed for most
solutes solutes
Primary urine Primary urine -- initial initial
filtrate filtered in filtrate filtered in
Bowman's capsule Bowman's capsule
that is isosmotic to that is isosmotic to
blood blood
Nost water and salt Nost water and salt
in the primary urine in the primary urine
is reabsorbed using is reabsorbed using
transport proteins transport proteins
and energy and energy
Each zone of the Each zone of the
nephron possess nephron possess
specific transporters specific transporters
Reclaims small molecules essential to the body Reclaims small molecules essential to the body
Nearly all sugar, vitamins, organic nutrients and, Nearly all sugar, vitamins, organic nutrients and,
in mammals and birds, water are reabsorbed in mammals and birds, water are reabsorbed
The concentration of beneficial substances in the The concentration of beneficial substances in the
filtrate is reduced as they are returned to the body filtrate is reduced as they are returned to the body
The concentration of wastes and non The concentration of wastes and nonuseful useful
substances is increased and excreted from the substances is increased and excreted from the
body body
&ecretion &ecretion
Filtrate is joined by substances transported across the Filtrate is joined by substances transported across the
tubule epithelium from the surrounding interstitial fluid as it tubule epithelium from the surrounding interstitial fluid as it
moves through the nephron tubule moves through the nephron tubule
Adds plasma solute to the filtrate Adds plasma solute to the filtrate
The proximal and distal tubules are the most common sites The proximal and distal tubules are the most common sites
of secretion of secretion
Similar to reabsorption, but in reverse Similar to reabsorption, but in reverse
Secretory products: K Secretory products: K
+ +
, NH , NH
44
+ +
, H , H
+ +
, pharmaceuticals, and , pharmaceuticals, and
water watersoluble vitamins soluble vitamins
A very selective process involving both passive and active A very selective process involving both passive and active
transport transport
Countercurrent ultipliers Countercurrent ultipliers
The loop of Henle and The loop of Henle and
collecting duct act as collecting duct act as
countercurrent multipliers to countercurrent multipliers to
create osmotic gradients that create osmotic gradients that
facilitate transport processes facilitate transport processes
The gradients are maintained The gradients are maintained
by by vasa recta vasa recta capillaries capillaries
The osmotic concentration of The osmotic concentration of
the final urine depends on the the final urine depends on the
permeability ( permeability (aquaporins aquaporins) of ) of
the distal tubule and collecting the distal tubule and collecting
duct which can be regulated duct which can be regulated
!mpermeable: dilute urine !mpermeable: dilute urine
Permeable: concentrated urine Permeable: concentrated urine
Figure 11.27
Transport Properties of the Transport Properties of the
Nephron and Collecting Duct Nephron and Collecting Duct
Secretion and reabsorption in the proximal tubule Secretion and reabsorption in the proximal tubule
Substantially alter the volume and composition of Substantially alter the volume and composition of
filtrate filtrate
Reabsorption of water continues Reabsorption of water continues
As the filtrate moves into the descending limb of the As the filtrate moves into the descending limb of the
loop of Henle loop of Henle
As filtrate travels through the ascending limb of the loop of As filtrate travels through the ascending limb of the loop of
Henle Henle
Salt diffuses out of the permeable tubule into the Salt diffuses out of the permeable tubule into the
interstitial fluid interstitial fluid
The distal tubule The distal tubule
Plays a key role in regulating the K Plays a key role in regulating the K
+ +
and NaCl and NaCl
concentration of body fluids concentration of body fluids
The collecting duct The collecting duct
Carries the filtrate through the medulla to the renal Carries the filtrate through the medulla to the renal
pelvis and reabsorbs NaCl pelvis and reabsorbs NaCl
Water Conservation Water Conservation --
concentrated urine production concentrated urine production
The mammalian kidney's ability to conserve water The mammalian kidney's ability to conserve water
is a key terrestrial adaptation is a key terrestrial adaptation
The mammalian kidney can produce urine much The mammalian kidney can produce urine much
more concentrated than body fluids, thus more concentrated than body fluids, thus
conserving water conserving water
The loop of Henle is largely responsible for the The loop of Henle is largely responsible for the
osmotic gradient that concentrates the urine osmotic gradient that concentrates the urine
Two solutes, NaCl and urea, contribute to the Two solutes, NaCl and urea, contribute to the
osmolarity of the interstitial fluid osmolarity of the interstitial fluid causes the causes the
reabsorption of water in the kidney and reabsorption of water in the kidney and
concentrates the urine concentrates the urine
O
NaCl
NaCl
NaCl
NaCl
NaCl
NaCl
NaCl
300
300
100
400
600
900
100
00
400
00
100
Active
transport
!assive
transport
OUTER
MEDULLA
NNER
MEDULLA
CORTEX
O
Urea
O
Urea
O
Urea
O
100
100
900
600
400
300
600
400
300
OsmoIarity of
interstitiaI
fIuid
(mosm/L)
300
Concentrated urine production Concentrated urine production
The countercurrent multiplier system maintains a high salt The countercurrent multiplier system maintains a high salt
concentration in the interior of the kidney, which enables concentration in the interior of the kidney, which enables
the kidney to form concentrated urine the kidney to form concentrated urine
The collecting duct, permeable to water but not salt The collecting duct, permeable to water but not salt
conducts the filtrate through the kidney's osmolarity conducts the filtrate through the kidney's osmolarity
gradient, and more water exits the filtrate by osmosis gradient, and more water exits the filtrate by osmosis
Urea diffuses out of the collecting duct as it traverses the Urea diffuses out of the collecting duct as it traverses the
inner medulla inner medulla
Urea and NaCl form the osmotic gradient that enables the Urea and NaCl form the osmotic gradient that enables the
kidney to produce urine that is hyperosmotic to the blood kidney to produce urine that is hyperosmotic to the blood
Fig. 12-23, p.552
%abIe 12-3, p.552
Excretion Excretion
After the urine is produced, it leaves the After the urine is produced, it leaves the
kidney and enters the urinary bladder kidney and enters the urinary bladder
Urine leaves the bladder via the urethra Urine leaves the bladder via the urethra
Sphincters of smooth muscle control the Sphincters of smooth muscle control the
flow of urine out of the bladder flow of urine out of the bladder
The opening and closing of the sphincters is The opening and closing of the sphincters is
controlled by a spinal cord reflex arc controlled by a spinal cord reflex arc
(micturition reflex) and can be influenced (micturition reflex) and can be influenced
by voluntary controls by voluntary controls
%egulation of Kidney unction %egulation of Kidney unction
The osmolarity of the urine The osmolarity of the urine is regulated by is regulated by
nervous and hormonal control of water nervous and hormonal control of water
and salt reabsorption in the kidneys and salt reabsorption in the kidneys
Endocrine hormones that affect kidney Endocrine hormones that affect kidney
function: function:
Diuretics Diuretics -- stimulate excretion of water stimulate excretion of water
Antidiuretics Antidiuretics -- reduce excretion of water reduce excretion of water
Steroid hormones Steroid hormones -- slow response slow response
Peptide hormones Peptide hormones -- rapid response rapid response
lomerular iltration %ate (% lomerular iltration %ate (%
Changes in the FR occur Changes in the FR occur
primarily as a result of primarily as a result of
changes in glomerular changes in glomerular
capillary blood pressure capillary blood pressure
Determined by three Determined by three
main forces: main forces:
lomerular capillary blood lomerular capillary blood
pressure pressure
Bowman's capsule Bowman's capsule
hydrostatic pressure hydrostatic pressure
Plasma Plasmacolloid osmotic colloid osmotic
pressure pressure -- osmotic osmotic
pressure due to protein pressure due to protein
concentration concentration
%abIe 12-1, p.535
FR depends not only on the net filtration FR depends not only on the net filtration
pressure but also on the glomerular surface area pressure but also on the glomerular surface area
available for filtration, and the permeability of the available for filtration, and the permeability of the
glomerular membrane glomerular membrane
Changes in any of the physical forces can affect Changes in any of the physical forces can affect
FR FR
Plasma Plasmacolloid osmotic pressure and Bowman's colloid osmotic pressure and Bowman's
capsule hydrostatic pressure are not subject to capsule hydrostatic pressure are not subject to
regulation regulation -- do not vary substantially under do not vary substantially under
normal conditions normal conditions
Can change pathologically and thus affect FR Can change pathologically and thus affect FR
A decrease in plasma protein concentration A decrease in plasma protein concentration
FR (and vice versa) FR (and vice versa)
Bowman's capsule hydrostatic pressure can Bowman's capsule hydrostatic pressure can
become elevated (obstruction to the urinary become elevated (obstruction to the urinary
tract, due to kidney stone or prostatic tract, due to kidney stone or prostatic
enlargement enlargement FR FR
Damming up of fluid behind the obstruction Damming up of fluid behind the obstruction
elevates capsular hydrostatic pressure elevates capsular hydrostatic pressure
lomerular capillary BP can be controlled to adjust the FR lomerular capillary BP can be controlled to adjust the FR
As the glomerular capillary BP As the glomerular capillary BP (assuming other factors (assuming other factors
remain constant), the net filtration pressure remain constant), the net filtration pressure and FR and FR
The capillary BP depends on the rate of blood flow within The capillary BP depends on the rate of blood flow within
each glomerulus each glomerulus -- is determined largely by the mean is determined largely by the mean
systemic arterial BP and the resistance of the arterioles systemic arterial BP and the resistance of the arterioles
!f resistance in the afferent arteriole !f resistance in the afferent arteriole less blood flows into less blood flows into
the glomerulus the glomerulus -- leads to leads to FR FR
!f resistance in the afferent arteriole !f resistance in the afferent arteriole leads to leads to FR FR
!f resistance in the efferent arteriole !f resistance in the efferent arteriole - - leads to leads to FR FR
Fig. 12-11a, p.536
Fig. 12-11b, p.536
Detection by aortic arch and
carotid sinus baroreceptors
$ympathetic activity
GeneraIized
arterioIar vasoconstriction
fferent arterioIar
vasoconstriction
GFR
GIomeruIar capiIIary
bIood pressure
Cardiac
output
%otaI
peripheraI
resistance
$hort-term
adjustment for
rteriaI
bIood pressure
Long-term
adjustment for
Urine voIume
Conservation of
fIuid and saIt
rteriaI bIood pressure
$timuIation of macuIa densa ceIIs
to reIease vasoactive chemicaIs
ChemicaIs reIeased that induce
afferent arterioIar vasoconstriction
Driving pressure into gIomeruIus
GIomeruIar capiIIary pressure
GFR
Rate of fIuid fIow
through tubuIes
BIood fIow into gIomeruIus
GIomeruIar capiIIary pressure
to normaI
GFR to normaI
Fig. 12-18, p.545
Fig. 12-19, p.546
a
+
/ ECF voIume/ arteriaI pressure
Renin
ngiotensin I
%ubuIar a
+
reabsorption %ubuIar K
+
secretion
Urinary a
+
excretion Urinary K
+
excretion
ngiotensin II PIasma K+
Idosterone

%egulation of % %egulation of %
Two major control mechanisms: Two major control mechanisms:
Autoregulation Autoregulation -- aimed at preventing aimed at preventing
spontaneous changes in FR spontaneous changes in FR
Extrinsic sympathetic control Extrinsic sympathetic control -- aimed at long aimed at long
term regulation term regulation
Such spontaneous changes in FR are Such spontaneous changes in FR are
largely prevented by intrinsic regulatory largely prevented by intrinsic regulatory
mechanisms initiated by the kidneys mechanisms initiated by the kidneys
AUTOREULAT!ON AUTOREULAT!ON
%egulation of % %egulation of %
The extrinsic control mechanisms can override the The extrinsic control mechanisms can override the
autoregulatory responses autoregulatory responses
The extrinsic control is mediated by sympathetic NS The extrinsic control is mediated by sympathetic NS
input to the afferent arterioles input to the afferent arterioles -- aimed at regulating aimed at regulating
arterial blood pressure arterial blood pressure
arterial BP is detected by the arterial carotid sinus and arterial BP is detected by the arterial carotid sinus and
aortic arch baroreceptors, which initiate neural reflexes aortic arch baroreceptors, which initiate neural reflexes
to increase BP to increase BP -- coordinated by the cardiovascular coordinated by the cardiovascular
control centre in the brain stem control centre in the brain stem -- mediated primarily mediated primarily
thorugh increased sympathetic activity to the heart and thorugh increased sympathetic activity to the heart and
blood vessels blood vessels
Compensation for a depleted plasma volume is reduced Compensation for a depleted plasma volume is reduced
urine output urine output -- accomplished in part by reducing FR accomplished in part by reducing FR
%ole of Baroreceptor %eflex in %ole of Baroreceptor %eflex in
Extrinsic Control Extrinsic Control
FR is reduced by the baroreceptor reflex FR is reduced by the baroreceptor reflex
response to a fall in BP response to a fall in BP
Sympathetically vasoconstriction occurs in most Sympathetically vasoconstriction occurs in most
arterioles (including afferent arterioles) arterioles (including afferent arterioles)
throughout the body throughout the body -- to increase total to increase total
peripheral resistance peripheral resistance
vasoconstriction vasoconstriction less blood flow into the less blood flow into the
glomeruli glomeruli glomerular capillary BP glomerular capillary BP
The resultant decrease in FR in turn reduces The resultant decrease in FR in turn reduces
urine volume urine volume -- restore the plasma volume restore the plasma volume
!ncreased thirst also contribute to long !ncreased thirst also contribute to longterm term
maintenance of BP maintenance of BP
Fig. 12-11a, p.536
!ntrinsic and Extrinsic !ntrinsic and Extrinsic
%egulators of % %egulators of %
Three intrinsic pathways Three intrinsic pathways
Nyogenic regulation Nyogenic regulation
Tubuloglomerular feedback Tubuloglomerular feedback
Nesangial control Nesangial control
Extrinsic endocrine hormones Extrinsic endocrine hormones
Angiotensin !! is a vasoconstrictor Angiotensin !! is a vasoconstrictor
Prostaglandins are vasodilators Prostaglandins are vasodilators
Figure 11.30
Antidiuretic Hormone Antidiuretic Hormone
(asopressin (asopressin
A peptide hormone (antidiuretic hormone A peptide hormone (antidiuretic hormone
ADH) produced in the hypothalamus and ADH) produced in the hypothalamus and
released by the pituitary gland released by the pituitary gland
!ncreases water reabsorption by the collecting !ncreases water reabsorption by the collecting
duct by increasing the number of aquaporins duct by increasing the number of aquaporins
Release: increasing plasma osmolarity detected Release: increasing plasma osmolarity detected
by osmoreceptors in the hypothalamus by osmoreceptors in the hypothalamus
!nhibition: increasing blood pressure detected by !nhibition: increasing blood pressure detected by
stretch receptors in the atria and baroreceptors stretch receptors in the atria and baroreceptors
in the carotid and aortic bodies in the carotid and aortic bodies
Osmoreceptors
in hypothalamus
Drinking reduces
blood osmolarity
to set point
H
2
O reab
sorption helps
prevent further
osmolarity
increase
ST!NULUS:
The release of ADH is
triggered when osmo
receptor cells in the
hypothalamus detect an
increase in the osmolarity
of the blood
Homeostasis:
Blood osmolarity
Hypothalamus
ADH
Pituitary
gland
!ncreased
permeability
Thirst
Collecting duct
Distal
tubule
(a Antidiuretic hormone (ADH) enhances fluid retention by making
the kidneys reclaim more water.
Aldosterone Aldosterone
A steroid hormone produced by the A steroid hormone produced by the
adrenal cortex adrenal cortex
Targets cells in the distal tubule and Targets cells in the distal tubule and
collecting ducts collecting ducts
Stimulates Na Stimulates Na
+ +
reabsorption (and reabsorption (and
secondarily water recovery from the urine) secondarily water recovery from the urine)
and enhances K and enhances K
+ +
excretion excretion
Stimulated by increases in circulating K Stimulated by increases in circulating K
+ +
%enin %enin Angiotensin Angiotensin Aldosterone Aldosterone
Pathway Pathway
Steps Steps
]uxtaglomerular cells secrete the enzyme ]uxtaglomerular cells secrete the enzyme
renin renin
Converts angiotensinogen to angiotensin ! Converts angiotensinogen to angiotensin !
Angiotensin converting enzyme (ACE) from Angiotensin converting enzyme (ACE) from
epithelia of blood vessels epithelia of blood vessels converts converts angiotensin angiotensin
! ! to angiotensin !! to angiotensin !!
Angiotensin !! causes the synthesis and Angiotensin !! causes the synthesis and
release of aldosterone from the adrenal cortex release of aldosterone from the adrenal cortex
%enin %enin Angiotensin Angiotensin Aldosterone Aldosterone
Pathway Pathway
Secretion of renin is controlled in three Secretion of renin is controlled in three
ways ways
Baroreceptors in juxtaglomerular cells release Baroreceptors in juxtaglomerular cells release
renin in response to low BP renin in response to low BP
Sympathetic neurons in the cardiovascular Sympathetic neurons in the cardiovascular
control center of the medulla oblongata control center of the medulla oblongata
trigger renin secretion in response to low BP trigger renin secretion in response to low BP
Nacula densa cells in the Nacula densa cells in the distal tubule distal tubule
respond to decreases in flow by releasing a respond to decreases in flow by releasing a
paracrine signal that induces juxtaglomerular paracrine signal that induces juxtaglomerular
cells to release renin cells to release renin
!ncreased Na
+
and H
2
O reab
sorption in
distal tubules
Homeostasis:
Blood pressure,
volume
ST!NULUS:
The juxtaglomerular
apparatus (]A) responds
to low blood volume or
blood pressure (such as due
to dehydration or loss of
blood)
Aldosterone
Adrenal gland
Angiotensin !!
Angiotensinogen
Renin
production
Renin
Arteriole
constriction
Distal
tubule
]A
(b The reninangiotensinaldosterone system (RAAS) leads to an increase
in blood volume and pressure.
Atrial Natriuretic Peptide Atrial Natriuretic Peptide
(ANP (ANP
!ncreases the release of Na !ncreases the release of Na
+ +
in the urine in the urine
Produced in specialized cells within the atria Produced in specialized cells within the atria
Excreted in response to stretch associated with Excreted in response to stretch associated with
an increase in blood volume an increase in blood volume
!ncreases FR by causing relaxation of the !ncreases FR by causing relaxation of the
contractile cells that control the size of the contractile cells that control the size of the
filtration slits of the glomerulus filtration slits of the glomerulus
Also decreases aldosterone, renin, and ADH Also decreases aldosterone, renin, and ADH
release and acts as an antagonist with the renin release and acts as an antagonist with the renin
angiotensin !! and aldosterone pathway angiotensin !! and aldosterone pathway
Thirst Thirst
Detected and controlled by the Detected and controlled by the
hypothalamus hypothalamus

Bio560 Osmoregulation

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Osmoregulation Osmoregulation

and Excretion and Excretion

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Osmoregulation and Excretion Osmoregulation and Excretion

BIO 560 Osmoregulation BIO 560 Osmoregulation 94 94

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