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Focus on
Osmoregulatory organs
contractile vacuole, antennal glands,
protonephridia/flame-bulb system,
metanephridia, malphigian tubules
Basic function of kidney
Types of nitrogenous waste
ammonia, urea and uric acids
Evolution of the vertebrate kidne
Freshwater fish, Marine fish, Amphibians and
Reptiles, Birds and Mammals

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In order for the system to function properly, the
relative concentrations of water and solutes in this
environment must be maintained within somewhat
narrow limits.

Homeostasis is one of the fundamental characteristics
of living things. It is the maintenance of the internal
environment within tolerable limits.

Living organisms achieve homeostasis by monitoring
and regulating a variety of internal parameters and by
behavioral changes.

Excretory systems help in maintaining homeostasis.


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Human homeostasis refers to the body's ability to
regulate its internal physiology to maintain stability in
response to fluctuations in the outside environment.

The liver and kidneys help maintain homeostasis.

The liver is responsible for metabolizing toxic
substances and maintaining carbohydrate
metabolism.

The kidneys are responsible for:
regulating blood water levels
re-absorption of valuable substances into the blood
maintenance of salt and ion levels in the blood
regulation of blood pH
excretion of urea and other wastes.
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Osmoregulation
Osmoregulation is the control of the levels of
water and mineral salts in the blood. It is a
homeostatic mechanism.

There are three important homeostatic
mechanisms:
osmoregulation
thermoregulation
regulation of blood sugar levels.

Homeostasis is important because it results in
our cells being bathed in tissue fluid which has the
correct amount of water, mineral salts, glucose
and temperature
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Osmoregulation is important in regulating
solute concentrations and balances the
gain and loss of water.
Osmoregulation is based largely on
controlled movement of solutes between
internal fluids and the external
environment.
Cells require a balance between osmotic
gain and loss of water. Water uptake and
loss are balanced by various mechanisms
of osmoregulation in different
environments
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Osmotic Challenges
Osmolarity is a measure of the osmotic
pressure exerted by a solution across semi-
permeable membrane (one which allows free
passage of water and completely prevents
movement of solute) compared to pure water.

Osmolarity is also expressed as solute
concentration expressed as molarity.


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Fig. 2.1 Osmolarity
7
Osmoconformers, which are only marine
animals are isoosmotic with their
surroundings and do not regulate their
osmolarity.

Osmoregulators used up their energy to
control water uptake and loss in a
hyperosmotic or hypoosmotic environment.

Most animals are said to be stenohaline
(organism that cannot handle a wide
fluctuation in the salt content of water) and
cannot tolerate substantial changes in
external osmolarity.

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Euryhaline animals such as mollusks and crustaceans
can survive large fluctuations in external osmolarity.

Euryhaline organisms are commonly found in
habitats such as estuaries where the salinity changes
regularly.

However, some organisms are euryhaline because
their life cycle involves migration between
freshwater and marine environments, as is the case
with salmon and eels.

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Freshwater animals
constantly take in water
from their hypoosmotic
environment and lose salts
by diffusion.

They maintain water
balance by excreting large
amounts of dilute urine.
Salts lost by diffusion are
replaced by foods and
uptake across the gills.
Fig. 2.3 Osmoregulation in
freshwater fish
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Freshwater animals show adaptations that reduce water
uptake and conserve solutes. They must rid themselves
of excess water.
Freshwater fish
10
Marine animals consist of marine invertebrates
and marine vertebrates. Most marine
invertebrates are osmoconformers.

Most marine vertebrates and some
invertebrates are osmoregulators.

Marine bony fishes are hypoosmotic to sea
water and lose water by osmosis and gain salt
by both diffusion and from food they eat.
These fishes balance water loss by drinking
seawater.

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Marine animals
11
Fig. 2.2 Osmoregulation in saltwater fish
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Animals That Live in Temporary Waters

Some aquatic invertebrates living in temporary
ponds can lose almost all their body water and
survive in a dormant state. This adaptation is
called anhydrobiosis
a) Hydrated tardigrade (b)Dehydrated tardigrade
Fig. 2.4 Anhydrobiosis. Tardigrade
(water bears) inhabit temporary pond
and droplets of water in soil and on
moist plants

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Land Animals
Land animals are
able to manage
their water
budgets by
drinking, eating
moist foods and
using metabolic
water.
Fig. 2.5 Water balance in two terrestrial mammals
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Desert animals
Get major water savings from simple anatomical features.

EXPERIMENT
Knut and Bodil Schmidt-Nielsen 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 Schmidt-Nielsen 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
Removing the fur of a camel increased
the rate of water loss through
sweating by up to 50%.

CONCLUSION
The fur of camels plays a critical role in
conserving water in the hot desert
environment where they live.

Fig. 2.6 The role of fur in camel in
water conservation.
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Marine birds
Transport epithelia are specialized cells that
regulate solute movement. They are essential
components of osmotic regulation and
metabolic waste disposal. They are arranged
into complex tubular networks.

An example of transport epithelia is found in
the salt glands of marine birds that remove
excess sodium chloride from the blood.

Salt glands is an organ to excrete excess salt.
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(a) An albatrosss 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.

(c) 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.

(b) 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.

Fig. 2.7 Salt-excreting glands in birds
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Types of nitrogenous waste
An animals nitrogenous wastes reflect its
phylogeny and habitat.
The type and quantity of an animals
waste products may have a large impact
on its water balance.
Among the most important wastes are
the nitrogenous breakdown products of
proteins and nucleic acids.
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NITROGENOUS WASTE
PROTEIN
NUCLEIC ACID
Breakdown product from
18
Forms of Nitrogenous Wastes
Different animals
excrete different
forms of nitrogenous
wastes:

Ammonia
Urea
Uric acid
Fig. 2.8 Nitrogenous wastes
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i) Ammonia

Animals that excrete nitrogenous wastes as ammonia need access to
lots of water. This is because ammonia is very soluble but can be
tolerated only at very low concentrations.

Therefore, ammonia excretion is most common in aquatic species.

Ammonia is release across the whole body surface or through the
gills. Many invertebrates release ammonia across the whole body
surface.

In fishes, most of the ammonia is lost as ammonium ions (NH4+) at
the gill epithelium. Freshwater fishes are able to exchange NH4+ for
Na+ from the environment, which helps maintain Na+ concentrations
in body fluids.

Ammonia excretion is much less suitable for land animals. Since
ammonia is so toxic, it can only be transported and excreted in large
volumes of very dilute solutions. Most terrestrial animals and many
marine organisms (which tend to lose water to their environment by
osmosis) do not have access to sufficient water.
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ii) Urea

The liver of mammals and most adult amphibians converts
ammonia to less toxic urea, such as in adult amphibians, sharks,
and some marine bony fishes and turtles.

Urea is synthesized in the liver by combining ammonia with
carbon dioxide and is excreted by the kidneys. The main
advantage of urea is its low toxicity, about 100,000 times less
than that of ammonia. Urea can be transported and stored
safely at high concentrations.

Urea is carried to the kidneys, concentrated and excreted with a
minimal loss of water. The main disadvantage of urea is that
animals must expend energy to produce it from ammonia.

In weighing the relative advantages of urea versus ammonia as
the form of nitrogenous waste, it makes sense that many
amphibians excrete mainly ammonia when they are aquatic
tadpoles. They switch largely to urea when they are land-
dwelling adults.
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iii) Uric Acid

Insects, land snails, and many reptiles, including
birds excrete uric acid as their major nitrogenous
waste.

Uric acid is largely insoluble in water and can
be secreted as a paste with little water loss.
While saving even more water than urea, it is
even more energetically expensive to
produce.

Uric acid and urea represent different adaptations
for excreting nitrogenous wastes with minimal
water loss.
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The Influence of Evolution and
Environment on Nitrogenous Wastes

The kinds of nitrogenous wastes excreted depend on an animals
evolutionary history and habitat. The amount of nitrogenous waste
produced is coupled to the animals energy budget. Mode of
reproduction appears to have been important in choosing among
these alternatives.

Soluble wastes can diffuse out of a shell-less amphibian egg
(ammonia) or be carried away by the mothers blood in a
mammalian embryo (urea). However, the shelled eggs of birds and
reptiles are not permeable to liquids, which mean that soluble
nitrogenous wastes trapped within the egg could accumulate to
dangerous levels. Even urea is toxic at very high concentrations.

Uric acid precipitates out of solution and can be stored within the
egg as a harmless solid left behind when the animal hatches.
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Table 1: Nitrogenous wastes
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Diverse excretory systems are variations on
a tubular theme

The function of excretory systems is to regulate
solute movement between internal fluids and the
external environment.

Excretory Processes

Even though excretory systems are varied, most
of them produce urine by refining a filtrate
derived from body fluids.
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1. Filtration. 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.

2. Reabsorption. The transport epithelium
reclaims valuable substances from the filtrate
and returns them to the body fluids.

3. Secretion. Other substances, such as
toxins and excess ions, are extracted from
body fluids and added to the contents of the
excretory tubule.

4. Excretion. The filtrate leaves the system
and the body.
Fig. 2.9 Overview of key functions of excretory systems
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Key functions of most excretory systems:

1. Filtration - pressure-filtering of body
fluids producing a filtrate
2. Reabsorption - reclaiming valuable
solutes from the filtrate
3. Secretion extraction and addition of
toxins and other solutes from the body
fluids to the content of filtrate in the
excretory tubule.
4. Excretion - the filtrate leaves the system

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Survey of Excretory Systems
The systems that perform basic excretory
functions vary widely among animal
groups.

They are generally built on a complex
network of tubules, for example,
protonephridia, metanephridia,
malpighian tubules, and vertebrate
kidneys
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Protonephridia: Flame-Bulb Systems
Flatworms have an excretory system
called protonephridia.

A protonephridium is a network of dead-
end tubules lacking internal openings. The
tubules branch throughout the body. A
cellular unit called a flame bulb caps the
smallest branches.

These flame bulb with a tuft (bunch) of
cilia draws water and solutes from the
interstitial fluid, through the flame bulb,
and into the tubule system.

These tubules excrete a dilute fluid and
function in osmoregulation.
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The urine in the tubules exits through openings
called nephridiopores. Excreted urine is very
dilute in freshwater flatworms. In fact, the tubules
reabsorb most solutes before the urine exits the
body.

In these freshwater flatworms, the major function
of the flame-bulb system is osmoregulation, while
most metabolic wastes diffuse across the body
surface or are excreted into the gastrovascular
cavity.

However, in some parasitic flatworms,
protonephridia do dispose of nitrogenous wastes.
Protonephridia are also found in rotifers, some
annelids, larval molluscs, and lancelets.
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Fig. 2.10 Protonephridia; the flame-bulb system of a planarian
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Metanephridia
Metanephridia is another tubular excretory system that
consists of internal openings that collect body fluids from
the coelom through a ciliated funnel, the nephrostome, and
release the fluid to the outside through the nephridiopore.

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Fig. 2.11 Metanephridia of an earthworm
32
Each segment of an annelid worm has a pair of open-ended
metanephridia. An earthworms metanephridia have both
excretory and osmoregulatory functions.

As urine moves along the tubule, the transport epithelium
bordering the lumen reabsorbs most solutes and returns them
to the blood in the capillaries.

Nitrogenous wastes remain in the tubule and are excreted
outside. Since earthworms experience a net uptake of water by
osmosis through the skin from damp soil, their metanephridia
balance water influx (entry) by producing dilute urine.
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Malpighian Tubules
In insects and other terrestrial arthropods.

Malpighian tubules remove nitrogenous wastes from
hemolymph and function in osmoregulation. Insects
produce a relatively dry waste matter, which is an
important adaptation to terrestrial life.

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Malphigian tubules open
into the digestive
system and dead-end at
tips that are immersed
in the hemolymph
(circulatory fluid).

The transport
epithelium lining the
tubules secretes certain
solutes, including
nitrogenous wastes,
from the hemolymph
into the lumen of the
tubule.
35
Water follows the solutes into
the tubule by osmosis, and the
fluid then passes back to the
rectum, where most of the
solutes are pumped back into
the hemolymph.

Water again follows the
solutes, and the nitrogenous
wastes, primarily insoluble uric
acid, are eliminated as almost
dry matter along with the
feces.

This system is highly effective
in conserving water and is one
of several key adaptations
contributing to the great
success of insects on land.
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Fig. 2.12 Malphigian tubules of insects
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Vertebrate Kidneys
Kidneys as the excretory organs of vertebrates,
function in both excretion and osmoregulation.

Nephrons and associated blood vessels are the
functional units of the mammalian kidney.

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The mammalian excretory system
centers on paired kidneys, which
are also the principal site of
water balance and salt
regulation.

Mammals have a pair of bean-
shaped kidneys. Each kidney is
supplied with blood by a renal
artery and drained by a renal
vein.
38
In humans, the kidneys account for
less than 1% of body weight, but they
receive about 20% of resting cardiac
output.

Urine exits each kidney through a duct
called the ureter, and both ureters
drain through a common urinary
bladder.

During urination, urine is expelled
from the urinary bladder through a
tube called the urethra, which
empties to the outside near the vagina
in females or through the penis in
males.

Sphincter muscles near the junction of
the urethra and the bladder control
urination.

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Structure and Function of the
Nephron and Associated Structures

Each nephron consists of a single long
tubule and a ball of capillaries, called the
glomerulus.

The blind end of the tubule forms a cup-
shaped swelling, called Bowmans
capsule, that surrounds the glomerulus.
Each human kidney contains about a
million nephrons, with a total tubule
length of 80 km.

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The mammalian kidney has two distinct regions, an outer
renal cortex and an inner renal medulla.

Both regions are packed with microscopic excretory
tubules, nephrons, and their associated blood vessels.

40
Nephron: functional unit of kidney
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Fig2.13 The mammalian excretory system
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Filtration of the blood
Filtration occurs as blood pressure forces fluid
from the blood in the glomerulus into the
lumen of Bowmans capsule.

The porous capillaries, along with specialized
capsule cells called podocytes, are permeable
to water and small solutes but not to
blood cells or large molecules such as plasma
proteins.

The filtrate in Bowmans capsule contains salt,
glucose, amino acids, vitamins, nitrogenous
wastes such as urea, and other small
molecules.

Filtration of small molecules is nonselective
and the filtrate in Bowmans capsule is a
mixture that reflects the concentration of
various solutes in the blood plasma.

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Pathway of the Filtrate
From Bowmans capsule, the filtrate passes
through three regions of the nephron: the
proximal tubule; the loop of Henle, a
hairpin turn with a descending limb and an
ascending limb; and the distal tubule.
The distal tubule empties into a collecting
duct, which receives processed filtrate
from many nephrons. The many collecting
ducts empty into the renal pelvis, which
is drained by the ureter.
The nephron and the collecting duct are
lined by a transport epithelium that
processes the filtrate to form the urine.
Their most important task is to reabsorb
solutes and water. The nephrons and
collecting ducts reabsorb nearly all of the
sugar, vitamins, and other organic nutrients
from the initial filtrate and about 99% of
the water. This reduces 180 L of initial
filtrate to about 1.5 L of urine to be
voided.

Mimi Sophia 44
Blood Vessels Associated with the Nephrons

Each nephron is supplied with blood by
an afferent arteriole, a branch of the
renal artery that subdivides into the
capillaries of the glomerulus.

The capillaries converge as they leave the
glomerulus forming an efferent arteriole.

The vessels subdivide again forming the
peritubular capillaries, which surround
the proximal and distal tubules.

Additional capillaries extend downward
to form the vasa recta, a loop of
capillaries that serves the loop of Henle.
The tubules and capillaries are immersed
in interstitial fluid, through which
substances diffuse.
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Although the excretory tubules and their
surrounding capillaries are closely
associated, they do not exchange materials
directly.

The tubules and capillaries are immersed in
interstitial fluid, through which various
materials diffuse between the plasma in the
capillaries and the filtrate within the
nephron tubule.
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From Blood Filtrate to Urine:
A Closer Look

Filtrate from Bowmans capsule becomes urine as it flows through the
mammalian nephron and collecting ducts.

Secretion and reabsorption in the proximal tubule significantly alter
the volume and composition of filtrate. For example, the cells of the
transport epithelium help maintain a constant pH in body fluids by
controlled secretions of hydrogen ions or ammonia. The cells also
synthesize and secrete ammonia, which neutralizes the acid.

The proximal tubules reabsorb about 90% of the important buffer
bicarbonate (HCO3).

Drugs and other poisons that have been processed in the liver pass
from the peritubular capillaries into the interstitial fluid and then
across the epithelium to the nephrons lumen. Valuable nutrients,
including glucose, amino acids, and K+, are actively or passively
absorbed from filtrate.
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One of the most important functions of the proximal tubule
is reabsorption of most of the NaCl and water from the
initial filtrate volume.

Salt in the filtrate diffuses into the cells of the transport
epithelium. The epithelial cells actively transport Na+ into the
interstitial fluid. This transfer of positive charge is balanced by
the passive transport of Cl out of the tubule. As salt moves
from the filtrate to the interstitial fluid, water follows by
osmosis.

Reabsorption of water continues as the filtrate moves into the
descending limb of the loop of Henle. This transport
epithelium is freely permeable to water but not very
permeable to salt and other small solutes.

For water to move out of the tubule by osmosis, the
interstitial fluid bathing the tubule must be hyperosmotic to
the filtrate.
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Since the osmolarity of the interstitial fluid becomes increasingly
greater from the outer cortex to the inner medulla, the filtrate
moving within the descending loop of Henle continues to lose
water.

In contrast to the descending limb, the transport epithelium of
the ascending limb of the loop of Henle is permeable to salt,
not water.

As filtrate ascends the thin segment of the ascending limb of the
loop of Henle, NaCl diffuses out of the permeable tubule into
the interstitial fluid, increasing the osmolarity of the medulla.

The active transport of salt from the filtrate into the interstitial
fluid continues in the thick segment of the ascending limb.

By losing salt without giving up water, the filtrate becomes more
and more dilute as it moves up to the cortex in the ascending
limb of the loop.

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The distal tubule plays a key role in regulating the K+
and NaCl concentrations in body fluids by varying
the amount of K+ that is secreted into the filtrate and
the amount of NaCl reabsorbed from the filtrate.

Like the proximal tubule, the distal tubule also
contributes to pH regulation by controlled secretion of
H+ and the reabsorption of bicarbonate (HCO
3
).

The collecting duct carries the filtrate through the
medulla to the renal pelvis and reabsorbs NaCl.

By actively reabsorbing NaCl, the transport epithelium
of the collecting duct plays a large role in determining
how much salt is actually excreted in the urine. Though
the degree of its permeability is under hormonal
control, the epithelium is permeable to water but not
to salt or (in the renal cortex) to urea.

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As the collecting duct go across the gradient of
osmolarity in the kidney, the filtrate becomes
increasingly concentrated as it loses more and more
water by osmosis to the hyperosmotic interstitial
fluid.

In the inner medulla, the duct becomes permeable
to urea. Because of the high urea concentration in
the filtrate at this point, some urea diffuses out of
the duct and into the interstitial fluid. Along with
NaCl, this urea contributes to the high osmolarity
of the interstitial fluid in the medulla.

This high osmolarity enables the mammalian kidney
to conserve water by excreting urine that is
hyperosmotic to general body fluids.

Mimi Sophia 51
Fig. 2.14 The nephron and collecting duct: regional functions of the transport epithelium.
Mimi Sophia 52
The mammalian kidneys ability to conserve
water is a key terrestrial adaptation
The mammalian kidney can produce urine much more
concentrated than body fluids, thus conserving water.

In the human kidney, about 80% of the nephrons, the
cortical nephrons, have reduced loops of Henle and
are almost entirely confined to the renal cortex.The
other 20%, the juxtamedullary nephrons, have well-
developed loops that extend deeply into the renal
medulla.

Only mammals and birds have juxtamedullary nephrons;
the nephrons of other vertebrates lack loops of Henle.
The juxtamedullary nephrons enable mammals to
produce urine that is hyperosmotic (hypertonic) to
body fluids and thus conserving water.
Mimi Sophia 53
Solute Gradients and
Water Conservation
In a mammalian kidney, the joint action and precise
arrangement of the loops of Henle and the collecting ducts are
largely responsible for the osmotic gradient that concentrates
the urine
Fig. 2.15 How the human kidney concentrates urine: the two-solute model
Mimi Sophia 54
Two solutes, NaCl and urea, contribute to the
osmolarity of the interstitial fluid which causes the
reabsorption of water in the kidney and
concentrates the urine.

Before leaving the kidney, the urine may attain
osmolarity of the interstitial fluid in the inner
medula, which can be as high as 1200 mosm/L.

Although urine is isoosmotic to the inner medula
interstitial fluid, it is hyperosmotic to blood and
interstitial fluid elsewhere in the body.

The high osmolarity allows the solutes remaining in
the urine to be excreted from the body with
minimal water loss.
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The countercurrent multiplier system involving
the loop of Henle maintains a high salt
concentration in the interior of the kidney,
which enables the kidney to form
concentrated urine.

The collecting duct, permeable to water but
not salt conducts the filtrate through the
kidneys osmolarity gradient, and more water
exits the filtrate by osmosis.

Urea diffuses out of the collecting duct as it
traverse (go across) the inner medulla. Urea
and NaCl, form the osmotic gradient that
enables the kidney to produce urine that is
hyperosmotic to the blood.

Mimi Sophia 56
Regulation of Kidney Function
Fig 2.16 Hormonal Control of Kidney
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1) Antidiuretic hormone (ADH)
2) Renin-Angiotensin-Aldosterone System
(RAAS)
57
Regulation of blood osmolarity is maintained by
hormonal control of the kidney by negative feedback
circuits.

One hormone important in regulating water balance is
antidiuretic hormone (ADH).
[Diuretic: substance tend to increase flow of urine]

ADH
* produced in the hypothalamus of the brain
* stored in and released from the pituitary gland, which
lies just below the hypothalamus
* osmoreceptor cells in the hypothalamus monitor the
osmolarity of the blood.

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When blood osmolarity rises
above a set point, more ADH is
released into the bloodstream
and reaches the kidney.

ADH induces the epithelium
of the distal tubules and
collecting ducts to become
more permeable to water.

This will increase water
reabsorption. This reduces urine
volume and helps prevent further
increase of blood osmolarity
above the set point.

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HIGH:
Osmolarity
59
When blood osmolarity rises
above a set point, more ADH is
released into the bloodstream
and reaches the kidney.

ADH induces the epithelium
of the distal tubules and
collecting ducts to become
more permeable to water.

This will increase water
reabsorption. This reduces urine
volume and helps prevent further
increase of blood osmolarity
above the set point.

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ADH Level
Water
reabsorption
Osmolarity
(High Na+)
HIGH:
Osmolarity
60
Mimi Sophia 61
By negative feedback, the drop in
osmolarity of the blood reduces the
activity of osmoreceptor cells in the
hypothalamus, and less ADH is secreted.

Only a gain of additional water in food and
drink can bring osmolarity all the way back
down to the set point. ADH alone only
prevents further movements away from the
set point.

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Renin-angiotensin-aldosterone system
(RAAS)
A second regulatory mechanism
involves a special tissue called the
juxtaglomerular apparatus
(JGA), located near the afferent
arteriole that supplies blood to the
glomerulus.

When blood pressure or blood
volume in the afferent arteriole drops,
the enzyme renin initiates chemical
reactions that convert a plasma
protein angiotensinogen to a peptide
called angiotensin II.

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LOW:
- Blood volume
- Blood pressure
63
Acting as a hormone, angiotensin II increases
blood pressure and blood volume in several ways:

It raises blood pressure by constricting
arterioles, decreasing blood flow to many
capillaries, including those of the kidney.

Stimulates the proximal tubules to reabsorb
more NaCl and water. This reduces the
amount of salt and water excreted and,
consequently, raises blood pressure and volume.

It also stimulates the adrenal glands, located atop
the kidneys, to release a hormone called
aldosterone. This acts on the distal tubules,
which reabsorb Na+ and water, increasing
blood volume and pressure.
Mimi Sophia 64
In summary, the renin-angiotensin-aldosterone
system (RAAS) is part of a complex feedback
circuit that functions in homeostasis.

A drop in blood pressure triggers a release of renin
from the JGA. In turn, the rise in blood pressure
and volume resulting from the various actions of
angiotensin II and aldosterone reduce the release
of renin.

Normally, ADH and the RAAS are partners in
homeostasis. ADH alone would lower blood Na+
concentration by stimulating water reabsorption in
the kidney. But the RAAS helps maintain balance by
stimulating Na+ reabsorption.
Mimi Sophia 65
EVOLUTION OF VERTEBRATE KIDNEY
Diverse adaptations of the vertebrate kidney have evolved in
different environments

The form and function of nephrons in various vertebrate classes
are related primarily to the requirements for osmoregulation in the
animals habitat

Exploring environmental adaptations of the vertebrate
kidney

Variations in nephron structure and function equip the kidneys of
different vertebrates for osmoregulation in their various habitats.

Mammals that excrete the most hyperosmotic urine, such as
hopping mice and other desert mammals, have exceptionally long
loops of Henle. This maintains steep osmotic gradients, resulting in
very concentrated urine.
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In contrast, beavers, which rarely face problems of dehydration, have
nephrons with short loops, resulting in a much lower ability to
concentrate urine.

Birds, like mammals, have kidneys with juxtamedullary nephrons that
specialize in conserving water. However, the nephrons of birds have much
shorter loops of Henle than do mammalian nephrons.

Bird kidneys cannot concentrate urine to the osmolarities achieved by
mammalian kidneys. The main water conservation adaptation of birds is
the use of uric acid as the nitrogen excretion molecule.

The kidneys of other reptiles, having only cortical nephrons, produce
urine that is, at most, isoosmotic to body fluids. However, the epithelium
of the cloaca helps conserve fluid by reabsorbing some of the water
present in urine and feces. Also, like birds, most other terrestrial reptiles
excrete nitrogenous wastes as uric acid.
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Cloaca is the posterior opening that serves as the
only such opening for the intestinal and urinary
tracts of certain animal species.

Birds, reptiles, amphibian excrete both urine and
feces through cloaca. For placental mammals have
different opening for evacuation.

In contrast to mammals and birds, a freshwater fish
must excrete excess water because the animal is
hyperosmotic to its surroundings. Instead of
conserving water, the nephrons produce a large
volume of very dilute urine.

Freshwater fishes conserve salts by reabsorption of
ions from the filtrate in the nephrons.

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Amphibian kidneys function much like those of
freshwater fishes. When in fresh water, the skin of
the frog accumulates certain salts from the water
by active transport, and the kidneys excrete dilute
urine.

On land, where dehydration is the most critical
problem, frogs conserve body fluid by reabsorbing
water across the epithelium of the urinary bladder.

Marine bony fishes, being hypoosmotic (hypotonic)
to their surroundings, have the opposite problem
of their freshwater relatives. In many species,
nephrons have small glomeruli or lack glomeruli
altogether. Concentrated urine is produced by
secreting ions into excretory tubules.
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The kidneys of marine fishes excrete very
little urine and function mainly to get rid of
divalent ions such as Ca
2+
, Mg
2+
, and SO
4
2
,
which the fish takes in by its nonstop
drinking of seawater.

Its gills excrete mainly monovalent ions
such as Na+ and Cl and the bulk of its
nitrogenous wastes in the form of NH
4
+.

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