Gas Exchange

Basic Principles
Large animals, with high met rates need a circulatory system to
deliver oxygen to the tissues b/c diffusion isnt fast enough to keep up
o They also have a dedicated gas exchange system
This is the point at which O2 enters the animal and CO2
leaves the animal
Regardless of looking at a small animal w/ a small met rate or a larger
complex animal what drives oxygen movement is the same
o He partial pressure difference between the environment and the
tissues
o Cells use O2 meaning there is a partial pressure gradient
between the external environment and the cell
O2 moves down that partial pressure gradient from out of
the cell in
CO2 moves from the cell other external environment
The Oxygen Cascade
o What drives o2 into an animal is diffusion along its partial
pressure gradient
o And this o2 movement can be thought of as a series of steps
called the O2 Cascade
High O2 levels in the external environment
Low O2 in the mitochondria where o2 is being used for
metabolism
o This difference of partial pressure between the environment and
the tissue site of o2 use that allows for o2 movement in other
tissues
o Depending the complexity of the animal there will be a series of
steps
Oxygen is delivered to the exchange surface by some type of
ventilation
Transfer of respiratory gases is Done in 4 steps in vertebrates
o Ventilation
Brings o2 to gas exchange surface
o Diffusion
Across the respiratory epithelium
o Carrying of O2 in the blood to the tissues
o Diffusion from the blood into the tissues
When this is broken down one ca see 2 steps where diffusion is
involved
o At the gas exchange surface

o At the tissues
And there are 2 steps where o2 is being carried by the bulk flow of
fluid
o Ventilation of air and water
o Blood flow
These are convective steps
Diffusion
o This is the movement of molecules by random Brownian motion
o Movement of gas by diffusion is driven by the diffusion gradient
which in the case of gases is the partial pressure gradient
pgas
o Diffusion depends on permeability
Kgas=estimate of permeability
Diffusion of gas is much faster in are than it is in water
and this reflects the permeability
Permeability in part depends on how readily the gas
dissolves into the medium(air, water or blood)
Gases are more mobile in air than in water b/c they
are more mobile in air
o Depends on surface area
Large the surface area the greater diffusion can occur
o And diffusion is inversely proportional to thickness
For fast gas diffusion, a thin barrier is preferred
o Fick equation: Mgas = Pgas Kgas SA/T
o Diffusion depends on P, not C
Convection
o The gas is being moved by the bulk flow of the medium and this
medium can be air water or blood
o The individual o2 molecules are carried by the medium
o This is a much faster transport pathway allows higher rats of
gas transfer
o In terms of o2 delivery; the movement/flow of medium and the
conc of gas in the medium
So how much o2 is present the medium
The flow of the medium times the concentration difference
tells you how much gas is being delivered
o Mgas = Vmedium Cgas
o Convection plays a role in boundary layers
Boundary layers are regions next to the gas exchange
surface that becomes depleted in o2
Area where o2 has diffused from and is now low inn o2
b/c they can become depleted in o2 they slow down
diffusion

o convection eliminates boundary layers by delivering o2 directly

to the gas exchange site/tissues
how are partial pressure and convection are related
o for gases partial pressure determines movement
o in terms of metabolism what really matters is how much gas is
present i.e. the conc of gas
o these two variable are related
see graph in slide 6
o capacitance is a way of relating concentration to partial
pressure (slide 6)
Capacitance = C/ P
if pressure and capacitance is known, one could calculate
conc
Capacitance of Air and liquids need to considered separately
o Air
looking at o2 and co2 in air they have exactly the same
capacitance b/c in air they follow the ideal gas law
when the ideal gas law equation is rearranges on can find
the slope of the relationship= 1/rt
so ALL gases have the same capacitance in air
o Fluid(blood/water)
If air is put overtop of a fluid where initially there is no o2
O2 will move into the fluid according to its partial
pressure gradient until the partial pressures are the same
in the fluid and the air
This will occur by diffusion
Partial pressure of o2 in the fluid is going to
equilibrate with the partial pressure of o2 in the air
over top so there is the same partial pressure in
both locations
The concentration of o2 in the gas vs the liquid will be
different b/c the conc of gas in the liquid will depend on
the solubility of the liquid for oxygen for o2 and
whether there is anything in the liquid to which o2
can bind
Gas will physically dissolve in the liquid and this is
determined by solubility
Knowing solubility you can calculate the conc of gas
in the liquid
There is a chance that o2 will react with the liquid and
this will increase the conc of o2 in the liquid
2 cases
O2 physically dissolves in the liquid

o Amount hat physically dissolves is determined

by the solubility coefficient=
o depends on the particular gas, temperature
and salinity
o as temp goes up solubility goes down
o from we can calculate the physically
dissolved o2
C= P
O2 is involved with additional chemical binding
o Chemical biding of the gas
The gas for o2 in blood and for CO2 in
water and in blood

Lec 8

When water is air equilibrated the CO2 concentration is water and air
is equal but the concentration of O2 in water and air differ
Conc of o2 is water depends on how much o2 can be physically
dissolved in the water and this is given by the following equation:
o [O2]dissolved= PO2 x
When going beyond water you end up with binding of o2 to
respiratory pigments like hemoglobin
If the gas chemically reacts with something in the solution then
physically dissolved gas no longer describes all the gas that is present
o You will have to add in a term to described the amount of
chemically bound gas that is present

o Ex blood contains hemoglobin and oxygen binds to hemoglobin

therefore the conc of o2 in blood is the amount of oxygen that
physically dissolves in the plasma plus the amount of o2
chemically bound to hemoglobin
CO2 chemically reacts with water
o They react to form carbonic acid which then dissociates in
bicarbonate and then this further dissociates to carbonate ions
o So the conc of co2 is not only the physically dissolved CO2 but
on top of that you have to add the amount of co2 that has
chemically reacted with the water
This includes the bicarbonate ions, and the carbonate ions
o Note: the pH of a physiological system is one in which we are
only worrying about the bicarbonate
It dominates b/c the carbonate ions are pretty small and
can be considered negligible most of the time
Co2 in blood
o It is physically dissolved
o Co2 reacts with the water in blood and so the conc of
bicarbonate and carbonate ions are included
o Co2 chemically reacts with respiratory pigments such as
hemoglobin and co2 that is bound to hemoglobin is called
carbamino co2
Carbamino co2 describes the amount of co2 that is bound
to hemoglobin
When there is more than just physically dissolved as, capacitance is
especially useful
o If there is only dissolved gas then one can rely on the solubility
coefficient to give the conc of gas present
o Beyond that(^^) the value of capacitance describes all of the
different ways the gas can be carried in the liquid
It includes things like O2 bound to hemoglobin, or co2
that is chemically reacted with water
is a functional measure of solubility
it will tell you how much gas is in the liquid whereas
the solubility coefficient will only tell you the amount
of gas that physically dissolved in the liquid
movement of o2 into animal can be divided into 4 steps
o oxygen is brought to the gas exchange surface
this involves ventilation so it involves air or water moving
across the gas exchange surface
this is a convective step
o the o2 has to move across the respiratory surface
this is a diffusive step and so it relies on the Fick equation
o o2 is carried in the circulatory system to the tissues sites of use

another convective step b/c the gas is being carried by the

blood
o o2 moves from the blood into the tissue site of use
this is a diffusive step relying on the Ficks equation
o can use this exact same model for co2 however its in reverse
for any convective step, the amount of gas that Is moved Mgas=flow
of medium carry it (air, water or blood) X the concentration
o mgas= amount of gas that is moved
Gas transfer revisited
MO2 = Vm (CiO2-CeO2)
o i=inspired
o E=expired
o How much air youre breathing multiplied by the difference
between your expired and inspired are in the amount of o2 that
is present
o Or it's the amount of water a fish is breathing multiplies by the
difference between inspired water and expired water in the o2
content
Sometimes its difficult to measure concentration and its often easier
to measure pressure
o Because we have a relationship between concentration and
capacitance, concentration can be replaced with capacitance
o Vm mO2 (PiO2-PeO2)
This tells how much O2 is being carried to the gas
exchange surface by the flow of air/water
o V=flow(ml/min)
MO2 = Vb (CaO2-CvO2) = Vb bO2 (PaO2-PvO2)
o Here we can calculate amount of O2 in the blood
o Here we use blood flow so Vb
As well as arterial(a)and venous(v) content Or Pressure
o The flow of blood multiplied by the difference between the
arterial venous blood in terms of how much blood is being
carried
Arterial blood brings O2 to the tissue and the Venous
blood flows away from the tissue
The difference between tells how much o2 the tissue
has acquired
In a well-designed gas exchange system, the amount of O2 moving
through the system will be the same the whole way
o So at each step we can look at a calculation for mO2 but if our
calculate the amount of oxygen that is being carried to the gas
exchange surface we should also note the amount of o2 that is
diffusing into the tissue(in s4)
o The amount of O2 moving should be constant within the system

The end game is to get o2 to the tissues. Everything

preceding that is just moving it into the system
this is a handy property b/c it means we can use things
that are easy to calculate such as the amount of o2 being
delivered to the lungs in the air as a proxy for things can
are really difficult to calculate like the amount of o2 being
delivered to the tissues by diffusion
o its quite difficult to calculate movement by diffusion even when
given the Fick equation b/c parts of the Fick equation are quite
difficult to calculate
MO2 constant across system so can rearrange equations to solve for
unknown variables.
MO2 = DPO2 KO2 SA/T
o Here is a eq of o2 consumption as a function of blood flow and
blood o2 content
o If you want to calculate blood flow; the rearrangement of the eq
will allow it
Blood O2 transport
O2 is carried in the blood both as physically dissolved gas and gas
that is bound to a respiratory pigment(i.e. hemoglobin but there is a
ton)
Hemoglobin increases the amount of o2 the blood can carry
o See slide 14
The amount of just physically dissolved o2 in the blood is about 0.3 vol
%
The amount of o2 present in the blood when you have a respiratory
pigment is 20 vol%
o Vastly more oxygen available b/c f chemical bind fo2 to the resp
pigment
o Hgb is an awesome o2 carrier
Without Hgb in order to deliver the necessary amount of
o2 blood flow must increase
Ex crustacean + fish
o Crusts have to have a higher blood flow b/c
their hemoglobin holds less o2
How high does blood flow need to be in anemic human
beings in order for the human to meet the normal o2
consumption of 1mmO2/g/hr.
See
There is a fish that lacks Hgb
o The ice fish see the blood in slide 16
o Its clear w/out any Hgb
This fish can survive without any Hgb by relying on
dissolved o2 only

This means that the fishes cardiac output will be very high
This means the heart of an ice fish is fairly large
The heart is actually 3x larger than the hearts of
similar red blooded fish to accommodate for the
need of a higher cardiac output
o It is an ectothermic living in very cold temperatures
Has a low met rate
Though the water is old and the solubility of o2 is really
high
Respiratory Pigments
Hemoglobin is not the only respiratory pigment but it is the most
common
It is the respiratory pigment of vertebrates; also found in a lot of
invertebrates
Called a respiratory pigment because it changes colour depending on
whether o2 is bound to it or not.
Hgb where O2 is bound(Hb-O2) is bright red
o Deoxygenated blood deoxy-Hb is a blue-red (so purple...)
o In crustaceans there is hemocyanin
This hemocyanin is copper based meaning when
oxygenated their blood is blue
Clear when deoxygenated
o In annelids they have chlorocruorins which are green when
oxygenated and clear when deoxygenated
o Worms have hemerythrins and this is violet color when
oxygenated and clear when its not.
The Hgb molecule is a tetrameric molecule, so there are 4 subunits
o 2 alpha and 2 beta subunits
o And associated with each globin is an iron based prosthetic
group called the heme group
This is where o2 binds
o 1 molecule of hemoglobin can bind 4 molecules of o2
Due to the subunits each with their own heme group
Lecture 9- Oct 3rd
- Which animals are blue-blooded- crustaceans.
- Can you distinguish between carrying capacity and binding affinity?
o Carrying capacity is the max amount of oxygen the blood can
cary, and the binding affinity is how well the oxygen binds to
hemoglobin.
- Sketch an oxygen equilibrium curve and use it to illustrate the bohr
effect.
o On the x-axis: Partial pressure of Oxygeno y-axis- %saturation of hemoglobim or oxygen saturation in units
and

o the curve looks like a shaped curve and

o with the Bohr effect we shift either right or left if you shift right its because CO2 has gone up or the other
way if CO2 has gone down.
The capacitance of blood for CO2 is much higher than that of plasma
yet Hb binds relatively little to CO2- explain.
o Its not because hemoglobin can bind CO2, so why does the red
cell matter- because there is a possibility of getting rid of the
products of the hydration reaction so promotes more CO2
loading into the blood (there is also something catalyzing the
reaction).
Respiratory physiology short answer question- 30mL min-1
Haldane effecto related to the Bohr effect- oxygen binding is allosteric inhibitor
of proton binding.
o Ties together CO2 transport and O2 transport in the blood.
o As the blood arrives to the tissues, oxygen is being given up to
the tissues and this mean that hemoglobin is being
deoxygenated and can bind more protons.
o When Hb is deoxygenated it can hold more protons which allows
more CO2 to be entered into the blood.
o The difference between the line of deoxygenated blood and
oxygenated is the Haldane effect.
o Blood moves from tissues to lungs/gills- then oxygen comes in,
binds to Hb, lowering affinity for protons- which allows for
dehydration to give you molecular CO2- benefits for CO2
unloading at tissue and benefits loading at gill or lung.
Diagram- we have air on the left (Va) and blood on the right (Vb)o the red blood cell is loaded up with CO2- most CO2 thats in the
blood is in bicarbonate ions in the plasma so we have to convert
it back to molecular CO2 and then that has to diffuse out of
blood into the air.
o And at the same time, oxygen is moving from the air into the
blood.
o As CO2 diffuses out of the blood, its going to push the reaction
towards CO2 formation (left) because its an equilibrium
reaction.
o To make CO2 you need protons and bicarbonate ions protons will be released by Hb, then Haldane effect will
come into play when oxygen binds to Hb (gives you
protons) nowyou need bicarbonate ions.
Most of them are carried in plasma so they have to get
back into the cell through anion exchanger while chloride
ions are moving out (facilitated diffusion) and gives you

more molecular CO2 which then diffuses out by partial

pressure gradient
. If there is any more HbCO2 (called carbomino CO2)
(small amounts bound to Hb)- it gives up the rest of the
CO2 because there is low levels of CO2 and that also
diffuses out. CA- carbonic hydronaze.
Water breathing teleost fish diagramo difference between lungs- when CO2 is let into the water it
reacts with water and products are protons and HCO3
gooed because it gets rid of CO2 which keeps partial
pressure constant.
o There is also no carbomino CO2 because its almost no
importance in fish.
Dogfisho they have carbonic anhydrase in the blood (floating in plasma),
and its also on the gill membrane where it can catalyze plasma
reactions.
o Up to about half of CO2 excretion happens in the plasma.
o There is also no Haldane effect.
Gas exchange organso gills are outfoldings of body surface and lungs are infoldings of
the body surface.
o Tend to have high surface area, high permeability, thin
membranes and richly visualized surface (lots of blood vessels).
o We can understand this MO2 (amount of O2 thats moving=
deltaPO2 x KO2 x SA/T.
Air vs watero the most important difference is that solubility of oxygen in
water is very low compared to that of air.
o So water holds about 30x less O2 than air does.
o This means that water breathing animals have to move a lot
more water to get the oxygen they need than air breathing
animals do.
MO2= Vm x concentration difference.
o Animal that has same rate of oxygen use, amount of oxygen is
the same but however because capacitance for water for oxygen
is so low, the water breather is going to have a much greater
flow of water.
o Without Hb you can only get dissolved oxygen in water and
thats very low so Hb is important.
o Vw (flow of water past the gill)- Vb (flow of blood past the gill)
there is 10-20x more water.
In air breathers Va:Vb = 1:1 at the gas exchange surface.
This is called the ventilation to perfusion ratio.

o Not only do water breathers have to move more water, but

water is also more dense and viscous than air and this has a
huge impact on how its designed.
Water goes in the mouth, goes through gas exchange
organ and goes out through a separate cavity.
Rather than in and out flow like you find in air breathers,
you have unidirectional flow.
There are a few exceptions- lamprae (jawless fish) water goes in the mouth and comes out through gill
pouches.
When they are attached to prey, they cannot be used
for breathing ther must be an in and out flow of
water in and out of the gill pouches.
o Even with the one way design of water breathers, the cost of
breathing is much higher for them
. For a fish thats just respiring, its 10% of their energy
just to breathe.
o Another difference between air and water is heat capacity1000x in water than in air.
This means that animals that live in water live in a giant
heat sink so most of them cant have body temperatures
that are different than the environment.
o There is a difference between O2 and CO2 in water capacitance of water for CO2 is much higher than it is for
O2 (because it dissolves and CO2 reacts).
Gas exchange through skino very low usually in vertebrates except in amphibians.
o The skin in amphibians can be an important source of O2 and
CO2 movement.
In some amphibians, the skin is the only gas exchange
organ.
o Oxygen moving from pool of air sitting near the skin.
o PO2 in blood starts low and then rises as oxygen moves into the
blood.
o What drives the movement?
Partial pressure of oxygen in the blood and partial
pressure in the water- there is a large difference because
there is no ventilation.
The skin usually isnt permeable, but in amphibians it isbut its not as protective.
There is limited surface area of the skin, its not ventilated,
limits size/metabolic rate.
Gills-

o
o
o

surface area is important for how much O2 uptake can occurand its usually linked to their lifestyle.
Mackerel is a swimmer- so higher surface area.
Internal gills are external gills that have a flap of tissue that
protects them.
It also has advatages for effective ventilation.
Ventilation water comes in the mouth and exits through the flap mouth opens and closes to move the water those two things together pump water across the
gills.
Its not only unidirectional, its non-stop.
Ram ventilation- swimming fast.
Tuna is obligate ram ventilators and they have lost the
capacity to pump so if they stop swimming they cant
breathe.
They have very high metabolic rate, high rates of water
across the gills.
Gill structureo 4 gill arches on side of fishes head that are
like a post supporting structure of the gilo after that there are 2 rows of filaments and
then covering the filament are plate-like
structures called lamellae (site where gas
exchange occurs).
o Maximum contact between water and the gas
exchange surface- lots of surface area.
Why dont gills work in the air?
They need the water to support the gill structure, when in
air surface area is really low.
There are exceptions- lamellae are connected so they
dont collapse in air (mud skipper).
If you look closely at regular gills- they are relatively flat,
thin epithelial layers that are separated by cells that hold
them apart called pillar cells.
As water moves through blood space in one direction,
water moves between lamellae in opposite directioncounter current of water and blood flow.

Lec 10
Water breathing teleost fish diagram-(Slide 31)

Blood flow on one side of the gas exchange organ(gill) and water
instead of air on the other side of the gill
o So when CO2 leaves the gills it is let into the water and reacts
with water producing protons and HCO3
This is good because CO2 that reacts with water does not
affect the partial pressure
Co2 diffuses out of the gill down its partial pressure
gradient and reacts with water
Meaning it keeps partial pressure constant.
This means water breather are able to get rid of CO2 more
readily than air breathers
o There is also no carbamino CO2 because its almost no
importance in fish.
Elasmobranch Fish (dog fish; slide 32)
Carbonic anhydrase is present, membrane bound in the gill and
floating around in the plasma
o This means that the plasma reactions are catalyzed
o This plasma reaction is catalyzed, meaning it occurs quickly
playing an important role in CO2 excretion
The red cells become less important b/c these fish lack the Haldane
effect
o Hb is a good buffer but in the absence of the Haldane effect the
contribution of hemoglobin to proton binding is lessened
Conclusion
o There less room in the blood cell to mop up protons and there
are catalyzed CO2 reactions in the plasma
Due to these two factors both plasma bicarbonate
dehydration and red cell bicarbonate dehydration
contribute to the excretion of CO2
These fish are very different than other fish in how this fundamental
process occurs
Gas exchange Organs
In vertebrates there are 3 possible gas exchange organs
o Integument
Body covering/skin
o Gills
Out folding of the skin
Evaginated highly folded extension of the skin
o Lungs
Infolding of the body surfaces
Internalized invaginated, highly folded infolding of body
surface

All three of these gas exchange organs have a series of design

features
o high surface area
o high permeability
o thin membranes and richly vascularized surface (lots of blood
vessels)
o Fick equation=MO2=PO2 x KO2 x SA/T
The respiratory medium has a big impact on the design of the gas
exchange organ itself
Eff ects of respiratory mediums (Air vs Water)
Water breathers
o Solubility of oxygen in water is very low compared to that of air.
So water holds about 30x less O2 than air does.
This means that water breathing animals have to move a
lot more water to get the oxygen they need than air
breathing animals do.
MO2= Vm x concentration difference(can be replaced
with KO2 and PO2)
o If we have 2 animal that have same rate of oxygen use and the
same partial pressure difference however on is an air breather
and the other is a water breather
Because capacitance for water for oxygen is so low, the
water breather is going to have a much greater flow of
water.
(if the carry capacity for oxygen is low then to deliver an
appropriate amount of oxygen you will need a higher flow
of respiratory medium)
Water breathers have to breathe 10-20X more water to
gain the same amount of oxygen as an air breather would
have to breathe air
o The flow of medium relative to blood flow
In water breathers the capacitance of water for oxygen is
much lower than the capacitance of blood for oxygen
Blood contain respiratory pigment and water doesnt
When O2 moves from water to blood you need a high flow
of water to deliver the oxygen
And you need a low flow of blood to take the O2
away from the gas exchange organ.
Water flow has to be 10-20X higher than blood flow to
move the same amount of oxygen
o The Viscosity of water
Water is both denser and more viscous than air

When the water breather move the water against their gas
exchange organs the water that they are moving is 800X
more dense and 50X more viscous than air
In order to overcome this water breathers use
unidirectional flow
o Water goes in the mouth, passes of the gills
and exits out through the apicular cavity
These are the flaps on either side of the
fishes heads
There are a few exceptions- lamprey (jawless fish)
o This fish has a plat like mouth which it uses to
suck on its prey
o When unattached water goes in the mouth and
comes out through gill pouches.
o When they are attached to prey, theyre mouth
cannot be used for ventilation
So instead of a unidirectional flow of
water it has to use tidal flow of water in
and out of the gill pouches(less effective)
o Heat Capacity of Water
Heat capacity is 1000x higher in water than in air.
Water acts as giant heat sink, any heat that fish produces
as the water passes across the gills the heat is transferred
from the fish to the environment
Consequence of this high heat capacity is that most water
breathers are not endothermic animals
In water the capacitance of water for CO2 is much higher
than it is for O2 (because it dissolves and CO2 reacts).
This makes it easy for water breathers to get rid of
carbon dioxide even though its difficult for them to
gain O2
o When looking at what regulates ventilation in
water breathers it is usually O2
o This is b/c they have difficulty getting O2 from
the environment
For air breathers
o blood and air hold similar amounts of oxygen (the same
capacitance)
o The flow of air and the flow of blood is the same
o The amount of air needed to deliver O2 to the lungs is equal to
the amount in blood needed to carry that oxygen away from the
lung
o Air breathers are tidal breathers
Air is sucked into the lung and then back out

In and out through the same set of tubes

o Ventilation in this case is regulated by CO2
This is because it is more difficult for them to get rid of
CO2
This comparison of ventilation flow and blood flow is called the
ventilation to profusion ration
Profusion means the flow of blood through the gas exchange organ
Able to distinguish a water breather from an air breather b/c the
ventilation to profusion ration in a water breather is usually around
10-20:1
Ventilation: profusion is air breathers = 1:1

Integument
Gas exchange through the skin is not very important in
mammals/bird/reptile and most fish
However this is important in amphibians
o Amphibians have thin permeable skin that is suitable for gas
exchange
o Frog n slide 37- some O2 across the skin; large amount of CO2
This large amount of CO2 excretion could be accounted
for by the fact that CO2 loss is easier in water
o The lungless salamanders
They do not have lungs or gills and rely entirely on their
sin for gas exchange
They have a high SA: V ration, low met rate,
For most vertebrates the skin is not a great gas exchange organ
o Oxygen moving from pool of air sitting near the skin.
o PO2 in blood starts low and then rises as oxygen moves into the
blood.
This movement is driven by the partial pressure of oxygen
in the blood and partial pressure in the water/air
Note: Slide 40; environmental partial pressure is still
much higher than the PO2 of blood even when oxygenated
This reflects the limitations of skin as a gas
exchange organ
This occurs because the skin is ventilated meaning there
s=is no mechanism to move air/water across the skin
other than the animal swimming or moving around
This is a disadvantage b/c the pool of O2 sitting
above the skin can become depleted leading to the
formation of boundary layer
o Meaning less O2 moving into the animal

Another problem is that there is a conflict between the

skin as a protection covering and the skin as a gas
exchange organ
For the skin to be an effective protective organ it
needs to be thick/tough/impermeable
o However for the skin to be a gas exchange
organ it should be the opposite(thin,
permeable)
The skin is also relatively limited in surface area
In slide 39 this frog lives in a high altitude area with
fairly hypoxic water causing it to remain immersed
in the water
Because its gas exchange organ on the sin it forms
folds in the skin to increase surface area increasing
gas exchange
o These fold of skin are not really ideal for
protection or locomotion

Gills
Specialized gas exchange organs
Out folding/evagination of the body surface
High folded to increase SA
o Surface area is important for how much O2 uptake can occur
and its usually linked to their lifestyle.
Ex. Mackerel is a swimmer- so higher surface area.
Internal gills are external gills that have a flap of tissue that protects
them.
o This provided protection and effective ventilation.
The presence of the apicular flow allows for ventilation of
the gills
What happens is that water goes in the mouth
which opens and closes to pump water
the gill flaps also open and close to help move water
across the gills(apicular pump)
this gives 2 pump actions in order to in the mouth
and exits through the flap
o this give a unidirectional non-stop motion of
water
this is essential b/c of the low solubility of O2 in water and
the fact that water is dense and viscous
even in this one way mechanism the cost of
ventilation in water breathing fish is much higher
than the cost of ventilation in an air breather
at rest, breathing would cost water breathing fish
about 20-30% or its resting met rate

Ram

o this is because they need to move more water

and it is dense and viscous
fish generally do whatever it takes to lower cost of
ventilation
o ex. ram ventilation
ventilation
When ram ventilation the fish opens its mouth and
water is able to flow in the mouth and across the
gills without having to pump water
This involves swimming fast.
This means they actively stop breathing and regain
the energy that would be spent on pumping water
across the gills
o Most fish will do this if they are swimming in a
current, or do it when they are swimming fast
enough; they can also get a ride in order to
ram ventilate
Ex the remora links/attaches to a shark
and stops pumping instead using the
flow of water generated by the sharks
swimming for ventilation(facultative ram
ventilation)
o Tuna is obligate ram ventilators as they have
lost the capacity to pump so if they stop
swimming they suffocate
They have very high metabolic rate, high
rates of water moving across the gills.

Gill structure
o 4 gill arches on either side of fishes head that are like a post
supporting structure of the gill
These have cartilage to help support its shape
o On each gill arch there are 2 rows of filaments
o covering the filament are plate-like structures called lamellae
this is the site where gas exchange occurs
they are equivalent to alveoli in mammal lung and par
bronchi in fish lung
they form a type of sieve that that water passes through
Maximum contact between water and the gas exchange
surface- lots of surface area.
o Filaments and lamellae shape is supported by the flow of water
Why dont gills work in the air?
o They need the water to support the gill structure, when in air
the surface area is reduced.

There are exceptions- lamellae are connected so they

dont collapse in air (mud skipper).
o If you look closely at regular gills- they are relatively flat, thin
epithelial layers that are separated by cells that hold them apart
called pillar cells.
As water moves through blood space in one direction,
water moves between lamellae in opposite directioncounter current of water and blood flow.
o The lamellae structure
Consists of
Two respiratory epithelial sheets which are separated by
blood space.
Pillar cells
These are cells that hold two epithelial cells at a
good distance.
oxygen has to travel from the water through the
diffusion barrier to get to the blood cell
the diffusion barrier is about 5um thick in most fish
and consist of
o a mucus layer
o respiratory epithelium
o blood
The epithelium is relatively thick because there is water
on one side and blood on the other and it needs to be
tough enough to withstand the movement of both passed
it.
the thickness of the diffusion barrier correlates with
lifestyle
The tuna has a thin epithelial layer because it has a
high performance lifestyle- has to take up a lot of
oxygen so has a very high surface area also.
Model of gas exchange
o Blood flow and water flow along epithelium are in opposite
directions(counter current)
o The counter current flow of blood and water is the basis of the
fish gills efficiency
Has the higher efficient
o In this counter current system the water that is entering the gillthe best oxygenated water- comes into contact with the blood
that is leaving the gill (the most oxygenated blood)
At the other end the water that is leaving the gill-which
has been depleted of O2-meets the blood that is depleted
of O2
o In countercurrent exchange

As blood leaves the gill it is trying to come to equilibrium

with inspired PO2(PiO2)
So the gradient may not be large at any point but
the final PO2 achieved in the blood is high b/c the
blood is trying to equilibrium with PiO2 water
This makes the countercurrent system more efficient.
o In concurrent flow water and blood go in the same direction
It starts off with a much higher gradient b/c the inspired
water(highly oxygenated) meets the venous blood which
has the lowest O2 giving a large gradient
Here the blood that is entering the gills is coming into
equilibrium with water that is leaving the grill
Highest PO2 achieved is whatever the expired
PO2(PeO2) is

Lungs
Lungs are highly folded with the surface area correlating with the
activity of the animal.
Total surface area in a human lung is about 80m2- folded in.
They are typically ventilated that moves air in and out of the
lung.
o In amphibians that is a positive pressure mechanism
where they gulp air and push down int the lung
o Lung in reptiles, mammals and birds/lungfish are suction
lungs where the volume of the lung is increased and this
pulls/sucks air into the lung
Diaphragm in mammals helps with mechanism for breathing.
Most reptiles cant move and breathe at the same time
o This is because muscles used for movement are also needed to
help pull the lungs outward to allow for the drawing of air into
the lungs.
o To be able to run and gain oxygen at the same time, they lift
their front legs and run on the back so they can use the muscles
in the front to breathe.
Other lizards stop breathing when they move
o This is why you see relatively low metabolic rates in lizards.
Mammalian lungo The functional unit as alveoli
This is a small, balloon like structure- very large surface
area required for maximizing oxygen uptake.
The alveoli also provides High surface area, it is relatively
permeable, thin, and it is actively ventilated (suction
mechanism),
It is also surrounded by a capillary network.

Capillaries are pressed tight against respiratory

epithelium
o The distance of the respiratory epithelium is
about 0.5um (similar value of tuna)
o Low thickness=better/easier O2 transfer
o This is 10x thinner than a fish gill
o Birds around 0.1 (thickness of barrier).
o Air is moved through a series of conducting airways
Air comes in through the trachea, which is a single tube,
that splits into two bronchi
Air continues down, until terminal bronchioles lead to
alveoli.
This structure is not involved in gas exchange at all
instead it just gets air to where it needs to be.
The only place where you can find gas exchange
besides in the alveoli is in the smallest bronchioles
leading to the alveoli
o These two are the respiratory airways.
Small changes in radius in the airways causes a big
difference in resistance
Beta 2 receptors on muscle that allows them to
dilate are the same receptors in lungs.
People with asthma take 2 agonist to promote
dilation.
o Ventilation
lungs are bounded by thoracic cage which consists of ribs
and diaphragm
for them to expand properly, the lung cannot be
physically attached to either structure
o So the lung is kept in a fluid filled space called
pleural space which is filled with fluid called
pleural fluid.
o This acts as a suction mechanism to hold lung
on to the wall of the thoracic cage.
o If the seal is broken the lungs own tension will
cause it to collapse
The lungs are expanded by the outward movement of the
ribcage and the downward movement of the diaphragm.
Model of gas exchange Mammalian lungs are not a very effective gas exchange surface
Here there is a pool of air on top of the blood vessel in alveolus
o This pool of air is tuned over/refreshed meaning there is
ventilation which improves the gas exchange model
o This is very similar to the mode of gas transfer across the skin

PiO2 and PeO2 and there is diffusion of oxygen in the air from the
alveolus into the blood
o PO2 starts high in the air and then decreases from inspired to
expired
o O2 moves into the blood starting from a low pressure to a
higher partial pressure
The PO2 of air in the lung is lower than that in the environment
making this model less efficient
o Meaning lungs contain stale air
o Po2 of air in lungs < PO2 of air in environment
The reason for this poorer air quality is the fact that the lungs contain
2L of air but with each breath taken one only inhales 500mL
o So the bets one can do is to turn over about of the air in the
lungs
o The conducting airways occupy about 150ml of volume and so of
the 500ml that one breathe only about 350ml actually reach the
alveoli
Not much air is turn over in the lugs at any given breath
o Because of this alveolar PO2 sis lower than inspired PO2 and
this limits the arterial PO2 that can be achieved
arterial PO2 is one that matches expired PO2
o If youre snorkeling, youre breathing through a tube. What
happens to arterial PO2? And alveolar PO2?
Arterial PO2+Aveolar
Will decrease because you are increasing the
amount of dead space(the tube that is used acts like
an extended conducting airway)
This make it more difficult for O2 uptake
o Less air in the lungs leading to arterial PO2 to
fall
This however doesnt happen because
o When a trend for arterial PO2 to fall the
monitoring systems(chemocenters) that detect
the blood kick in and increase tidal volume
This compensates for the anatomical
dead space
Bird lungs
o They have relatively compact lungs which are about half the size
of mammalian lungs
Instead of alveoli their lungs are made up of tiny tubes
made up of parabronchi (site of gas exchange).
o In addition there are respiratory sacs which are involved in
ventilating the lung and they hold about 80% of the total volume
of the system

They are not involved in gas exchange, instead they are a

part of the ventilatory mechanism
This allows the bird lungs to be more efficient in
comparison to mammalian lungs
They also have a very low diffusion distance relative to
mammals (0.1 micron)
Correlation between lifestyle and thickness of respiratory
epithelium
Birds with thick respiratory epithelium are usually non
flying birds(ostrich, emu, penguin)-similar to mammals
This low diffusion distance however makes birds susceptible to
respiratory problems
Bird Lung Structure
In terms of lungs as a whole, flow is tidal
However when looking at gas exchange surface
itself(parabronchus) flow is one-way and continuous
This is the advantage of using tubes instead of
balloons as the gas exchange site
Air comes in through the bronchus and down to the
mesobronchus from which it enters the dorsobronchus
and goes through the parabronchi be collected by the
ventobronchus and be collected again
A loop through the lung itself with a tidal component
in and out
Capillaries surround the parabronchi
O2 moves form air in parabronchi and into the
capillaries
In this model the PO2 of air depends on where it is in the
air tube
If blood meets air at the inspired end of the tub the
air has a High PO2
If the capillary is at the expired end of the tube the
blood is going to be meeting air with a low PO2
o Air PO2 is low so PbO2 is low
What comes out of the lung is a mix of all these diff
situations
o On average experiment show that the arterial
PO2 is just above expired PO2
o So this model(cross current arrangement) is
much better than the pool type arrangement in
the mammalian lung but not as good as the
counter current arrangement in fish lungs
Air Movement through the bird lung
Ventilatory innovation of bird lings are air sacs

o
o

Two set of air sacs

Posterior and anterior
It takes 2 complete respiratory cycles to move a bolus of
air through the lungs
Inhalation 1
The bolus of air(green in slide59) is pulled into the
air sacs and the dorsobronchus by the expansion of
the air sac
The bird exhales and this compresses the air sac
pushing air out of them and forcing air through the
parabronchi
This completes one respiratory cycle and the bolus
of air is halfway through the line
Inhalation 2
The bird inhales again, expanding the air sacs
pulling air in this continues movement through the
parabronchi
The bird exhales for the second time and this
compresses the air out the ventobronchus and back
out in a tidal fashion
Because of these air sacs and the fact that the bird lung is
made up of tubes other than balloons you end up with a
functional gas exchange unit that is a continues one way
flow of air
This allows for a more efficient model of gas transfer
Some recent work suggest that it may be found in
realties with high metabolic rate
o Bird dont have counter current blood flow because there is a
sea of capillaries winding around the air tube and so air is
moving through the tube and blood Is going across at the same
time
This is a cross current way of blood flow

Extraction eff iciency

This is the difference in inspired to expired PO2 divided by inspire
Po2
o Essentially saying how much O2 is removed from the air that is
breathed as a percentage of the inspired Po2
For mammals efficiency is usually around 20-25%
o Pool type arrangement which isnt efficient at all
Bird: can be up to 40%
o Cross current blood/air flow allow for higher efficiencies to be
achieved
Fish=20- 60%

o Design of fish gills allow for more efficient transmission of gas

o This is because they have the counter current arrangement of
blood and water flow
Mimic effects of high altitude on birds contrast and compare birds
and mammals. Bird has been found at 9000 meters.
Extraction efficiency is how much PO2 falls when gas exchange
happens. In mammalian- 20-25%, bird is 40% and fish can be as high
as 60%. So design of fish gill allows for more efficient transmission of
gas. Why is the fish gill the most efficient design?

;Bio3302Lec 12-13
Control of Ventilation
There are two parts to ventilation
o generating basic ventilatory movements (breathing movements)
o how you match ventilation to the requirements of the animal
flying bird is at a high need
There is a fairly complex set of brain neurons that generate motor
output to breathing muscles (located in brain stem, medulla).
o On top of that, there are levels of control that allow breathing
methods to be controlled
Brain centers that allow you to hold your breath, or
voluntary control.
In addition to conscious control, there are mechanisms to match size
and frequency of breaths to match needs of animal.
o Most important of these are chemoreceptors that are involved to
detecting o2 and co2 levels in body, and matching requirements.
Mammalian chemoreceptors
2 sets of chemoreceptors that regulate ventilation
o central (in brainstem)
The most important component in the frequency and size
of breaths in mammals.
The chemoreceptors are proton sensors (detect hydrogen
ions)
however, they typically respond to Co2, because of
the blood brain barrier
o Protons dont readily pass blood brain barrier
but CO2can and so it reacts with water to form
protons.
As CO2 levels rise, more of it enters, reacts with water,
and proton detecting cells respond to increase in proton
levels by increase in breathing.
A fall in pH (increase in proton levels) stimulates
breathing (brings CO2 levels back to where they should
be).

Fish

o peripheral chemoreceptors
These are capable of detecting O2, CO2 and protons, but
more sensitive to CO2 and protons than they are to O2.
Found in two locations (called glomus cells) one in the
carotid body, and the aortic bodies,
Detect blood going to the brain to make sure that
the brain gets sufficient oxygen delivery.
chemoreceptors
There are no central chemoreceptors (nothing in the brain).
They have peripheral chemoreceptors and are located on the gills and
called neuroepithelial cells- similar to glomus.
Through evolution, the structure in the gill arch in the fish became
incorporated in the homologous structure in mammal, leading people
to believe that neuroepithelial cells and glomus are homologous.
o However this is not the case
o Neuroepithelial cells detect CO2+ O2levels of blood and water
They are not sensitive to protons
o Glomus are found only blood levels(not found in the fish doe)
Why is ventilation keyed to O2 in water breathers, but CO2/pH in air
breathers?
o In Fish ventilation is keyed to O2
In fish ventilation primarily response to oxygen levels
In water O2 is limiting and therefore animals that breathe
water need to monitor O2 very carefully in order to get
sufficient O2
Remember CO2 is able to react with water but not with air
making to easier to get of CO2
o In mammals
Ventilation changes primarily to changes in CO2 or pH
It is more difficult to get rid of CO2 in air
For air breathers it is much more easier to get O2 in air
and so they breathe less to get the same amount of O2 as
water breather
By breathing less CO2 levels become higher
o So here there are 2 factors at play
O2 is difficult to obtain in O2 so water breathers maintain
O2 but in air breathers they are able to breathe less to
get the required amount of O2 and therefore CO2 levels
tend to accumulate
So the convection requirement(how much one has to
breather in ait much lower than in water in air and b/c of
the low convection requirement in air CO2 levels tend to
higher
Humans PCO2=40Torr

 Fish PCO2= 4Torr

Diving and Divers-respiratory and cardiovascular adjustments
Divers are air breathers that go under water, where they no longer
have access to respiratory medium, and have to cope.
o Often they are exercising at this time (swimming) without
oxygen.
o Those that dive to great depths can run into problems because
of pressure and temperature.
o Animals that are adapted divers use the same responses as
animals that arent; they just take it to a higher level.
Opportunistic nature of natural selection is the result.
First problem is holding their breath.
o When you hold your breath, CO2 levels rise, which are the
strongest ventilatory stimulus for air breathers.
Slide 70
o This line on the left is a just ordinary humans that are subjected
to the test, so as CO2 levels rise, breathing rises.
The three lines on the right are for 3 people who are
trained as divers, you can see that they tolerate much
higher levels of CO2 before ventilation starts to go up.
Adapted divers exhibit blunted ventilatory sensitivity
to CO2
By hyperventilating before diving they lower Co2 levels
so that the levels dont go up as fast when holding breath.
What about exercising?
o For divers, the only oxygen available is what they take with
them (blood, lungs and muscles).
o Most adapted divers dont take oxygen into their lungs.
So whatever is left is in the blood and muscle.
The bottom figure in slide 71 shows total oxygen thats
available.
In general, most is available in the blood.
Adapted divers increase blood volume and
hematocrit (Hb levels).
o In some cases, they also have high levels of
muscle oxygen levels, which they achieve by
having higher levels or myoglobin.
A problem with high hematocrit is the strain on the
heart but they deal with it by only releasing red
blood cells when they dive (hematocrit increases
during the dime, and decreases when it emerges).
o They use the spleen as RBC reservoir, release
it then re-sequester into the spleen after the
dive

 Another way of maximizing oxygen stores is stripping as much

oxygen as possible from blood to bring to muscle.
o For animals that have a lot of blood oxygen have a right
shifted oxygen curve (high p50 value, lower affinity for Hb
to oxygen makes it easier for oxygen to unload from
blood).
o As animal mass increases, p50 values fall because small
mammals have higher metabolic rates and need to
enhance oxygen delivery.
Both seals have higher p50 than expected and large
caticaious.
This is thought to be adaptation for oxygen delivery
during a dive. Small cateceans (dolphins) have
higher affinity for oxygen (lower p50) than you
would expect- easier to take up oxygen from lowoxygen environments. The reason is because they
dive on a breath of air- shallow divers so their high
affinity Hb is useful for taking oxygen from that lung
full of air as oxygen gets depleted. Some have strong
Bohr effect- shifts curve to right to maximize oxygen
delivery.
In addition, these animals use oxygen as slowly as possible.
o One strategy is to not exercise when they dont have to.
o Rather than actively swimming, these animals glide down and
then swim up and by gliding; they use less oxygen and can stay
down longer.
The other strategy is to become slightly hypothermic.
o Its been difficult to document this in free diving animals.
o It looks like body temp falls when animal dives.
These animals allow themselves to become a bit
hypothermic because it lowers metabolic rate,
they suppress the shivering only when theyre
diving;
make oxygen stores last much longer

Exercising, swimming around without oxygen

o they can increase blood volume, try and minimize way that they
use the oxygen by
Lowering body temperature and directing the blood flow
to the tissues that need it most.
Brain, heart, lungs

o This happens across a variety of tissues.

It is important to note that blood flow to the heart is
greatly reduced during diving, even though its oxygen
sensitive- when it dives, the heart doesnt beat as fast and
therefore doesnt need as much oxygen/blood flow.
Slide 77
o A- Animal at surface, dives then comes back up (HR falls when
diving).
Dive response
This is a fall in heat rate and a selective O2 delivery
system
It is present in most air-breathing vertebrates,
Particularly strong to animals who have diving lifestyle
(natural selection).
o B)Adapted divers show unusual tolerance to low blood
oxygen levels(tolerance of hypoxia)
Divers have a pronounced tolerance for hypoxemia
Data for seals as they did natural dives, as it dives
blood oxygen levels fall and lowest levels are very
low.
The critical PO2 for a seal is about 10mmHg
o They deplete 90% of blood O2 reserves
during a routine dive
o This is the point where there is
irreversible brain damage,
o when they dive they are barely above that
value
For us, lowest PO2 are 20-30mmHg
This measurement was taken from human
blood that was subjected to climbing Mount
Everest.
Anaerobic metabolism
during the dive vasoconstriction of the arterioles going to skeletal
muscles occurs
however muscles are able to work anaerobically and when they do
they produce lactic acid
o This acid accumulates in the muscle not in the blood.
o This is a benefit of peripheral vasoconstriction because it traps
the acid.
When animal gets back to the surface, the lactic acid is released into
the blood and has to metabolize the acid, get rid of CO2 (recovery)
and its anaerobic.
o Adapted divers have high blood buffer values so they can
tolerate high levels of lactic acid.

o Because of the costs of recovery of diving, most divers dont use

this type of diving unless they have to.
Only for dives that are longer than 20 mins in length lead to the
accumulation lactic acid.
o Dives that are over this length are a lot less frequent because
the animal avoids this recovery period.
In addition to exercise without oxygen, the animals have another
problem to face and its the high pressure at depth.
o Every 10m, pressure increases by 1atm so at 30m, its 4atm.
At the depth gas exchange between lung and tissues
means that the tissues experience these high partial
pressures that are in the lungs
o This isnt really a problem for the tissues, but high pressure
compresses the lung which results in high partial pressure
thats in the lung.
When the animal surfaces, the gases come out of solution
which causes gas bubbles and this could be a fatal
problem.
This problem can be avoided if the animal comes up slowly
enough, to allow the blood to equilibrate with the new
partial pressures.
Animals that dive however dont have this option
Animals avoid the bends by doing the following
1. dive on empty lungs as much as possible (exhale rather than
inhale before they dive),
They have larger tidal volumes than usual so they are
more effective at emptying the lungs than humans would
be.
2. When they dive their alveoli collapse and when this occurs air is
pushed into the conducting airways
This is an advantage because the gas thats trapped here
cant equilibrate with the blood.
Humans have opposite response, its not alveoli that
collapse but lungs.
o High pressure depth causes the rib to
collapse inwards and helps to empty the lungs
to minimize equilibration between air in lungs
and blood.
3. There is also strong peripheral vasoconstriction
Blood isnt circulating to the tissues so minimizes
chance of equilibration of tissues.
Summarize specialization of vertebrate diverso vasoconstriction of peripheral vasculature,
o lower heart rate

o
o
o
o
o
o
o

Birds
o

(both are for dive response)

blunted ventilation (CO2),
high levels of Mb and Hb (expand oxygen stores),
ability to cope with high levels of lactic acid (blood buffering),
strong facility for anaerobic metabolism,
drop in body temperature
Gliding rather than active swimming.
Pulmonary
alveoli can collapse
dive on an exhalation rather than inhalation.
are ableto perform better at higher altitudes thanmammasl
This can be attributed to
Higher SA of gas exchange surface
Blood to air diffusion barrier is thinner
They are able to withdtasn ntracranial pressure better
than mammals do
When you put birds and mammals at high altitudes
they hyperventilate and as s result of this CO2 levels
fall
When co2 levlesfallthis affects brian blood flow in
mmamals but not in birds
o Birds are able to cope with hypercapnia(low
co2 levels)
Better neurons are more tolerant of low O2than
mammalian neurons there fore they can withstand hypoxia
better
Because birds use tubes ratherthen balloons they can
have unidirectional constsnt air flow
They can also use coress current blood/air
flowgivinfgimproved efficiency of O2 extraction
Theu do not show hypoxic pulmonary vascontrciiton
This helps at high atlitudes as you are breathing air
of low O2 levels
In this situation there would be vasoconstriction in
mammalian lungs
o This cnan lead to oedoma
Capillary to muscle cells ditacnce is lower in bird than in
the muscles making it easy to deliver O2 under hypoxic
condiions
Birds have larger hearts and lungs for better o2 delivery
This suggest that birds seem to do better
These mechanism occurred more in birds than in
mammals because of the birds ability for flight

O2 delivery requirement for flight are much higher

than they ar for swimming or running and therefore
all of these adaptaions came into play for O2
delivery during flight
o One mammal where one cn see these suit of traits are in bats
They have larger than normal hearts, larger than normal
lung SA, thinner than typical blood to air diffusion ditnces.