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Bio 3302 Lec 2

Mechanisms of oxygen delivery and co2 removal is based upon 2

systems circulatory system and respiratory system
Oxygen moves around in cells by the use of diffusion
o The time for diffusion depends on how far the oxygen has to
o Time required= distance2
As distance gets larger time required to increases fairly
o Larger animals need a system to enhance the rate of delivery of
Metabolic rate also come into play in terms of oxygen delivery
o The higher the metabolic rate the more oxygen needed
Ex large animals with very low metabolic rates may not
really need a circulatory system
Ex 2 small animals with very high met rates may need a
circ system in order for faster O2 delivery
o Once animal that is an exception to this rule of having a high
met rate= having a circ system
These are insects. They have the highest known met rates
but dont use their circ system for oxygen delivery
This is b/c they have a tracheal system which circulates
gas until it is very close to the tissues and then oxygen
diffuses from the end of the tracheal system into the
Circ system can also be used for
o Waste excretion
o Nutrient delivery
o Cell to cell communication
o Hormone transport
o Thermoregulation
Circ syst will transport anything that can move in a fluid
but it can also transport heat
o Generates force
Animals with hydrostatic skeletons rely on fluid force
generated by the circ system
o Animals that need to enlarge or extend an organ also uses the
circ system to do so.
Ex monarch butterfly
Monarch caterpillars and they use the circ system to
Once they are done feeding off milkweed they will
form into a crystal

And they emerge from the crystals with their wings

folded up the force that unfurls their wings is
hydrostatic force provided by a circ system
Circ system has 3 key elements
o A pump
Usually a heart but not always
The pumps work by generating force and pushing the fluid
a head of them(+pressure)
3 basic types of pumps
Peristaltic pump
o Ex type of heart found in insects and
o Its a tube with muscular walls and a wave of
contractions passes along the tube pushing the
blood in front of it
o This heart has force generation and direction
built into one
Direction in which contraction moves
determines the direction in which the
blood moves
Chamber pump with contractile walls
Ex Human heart
o muscle chamber and when muscle contracts it
squeezes the blood pushing it out
o in order for blood to move in the right
direction a series of valves are needed to
direct the flow
chambered pump with non-muscular walls; but is
surrounded by external factor(muscle) that
o this also needs valves to direct flow
o the external muscle will contract squeezing
the chamber and then the valves determines
where the fluid flows
o ex of this type of pump in humans
The large veins in the leg act as
chambers to collect blood and when one
moves their legs muscle contraction
squeeze those veins to return blood to
the heart.
o Network of vessels(vascular system)
3 types of tubes

o Take blood from heart to the periphery of the

o Specialized for exchange of material
o Collect blood from periphery and take it back
to the heart
Not all 3 are present in all animals
Collectively known as vascular system or peripheral
o Circulating fluid
This can be blood or haemolymph
Open vs closed system shows where you see the presence or absence
of the diff types of blood vessels(bvs)
Open Circulatory System
o Animals with open circ system do not have capillaries
o Blood will be pump by the heart into a set of arteries form there
it goes to the tissues directly
The blood bathes the tissues directly; there is only
exchange between the circulating fluid and the tissues
o Circulating fluid is then collected up and returned back to the
heart and this may/may not involve some type of venous system
o It is quite difficult to direct flow with an open system
Tend to be low flow, low pressure, low resistance systems
o If the system of vessels is open one cannot distinguish between
the circulating fluid and the fluid that is bathing the tissues
o Animals with open circ systems have haemolymph instead of
Makes up 30% of animals body mass
Closed Circulatory system
o The blood is in vessels the entire way; from leaving the heart to
returning to the heart
Animals with higher met rates tend to have closed circ
This is a evolutionary trend
o Reason for this trend is the ability to better
control the rate of blood flow and hence the
rate of O2 delivery with a closed separated
As met rate increases you tend to see the trend of
closed circ system
Within these animals a second trend can be seen

As met rate goes up there tends to be more

separating of blood flow to the gas exchange
organs(resp circ) and blood flow to the rest of the
tissues(systemic circ)
o As met rate goes up there is an increased
separation between the respiratory and
systemic circulatory systems
o The animals with higher metabolic rates are
air breathers and along with this there is a
shift to separating out the respiratory and
systemic circulation
Basic fish plan;
Closed implies that there is a complete set of vessels
Blood leaves the heart and arteries exchange between the
tissue and blood occurs in the capillaries and blood
returns to the heart in veins
It is a continuous circuit of vessels with an exchange
between the circulating fluid and the tissues happening at
the capillaries
These circulatory systems are found in all vertebrates(fish,
amphibians, reptiles, mammals, birds) and in a number of
invertebrates particularly ones that have a high metabolic
rate(cephalopods-squid and octopus; or in animals that use the
circ system for a hydrostatic skeleton-earth worm)
b/c blood is circulating in vessels all the time the circ system
provides quite a bit of resistance to flow and so higher pressures
must be generated in order to move the blood through the
closed systems of vessels
these are often considered to be high pressure/high
resistance systems
more force must be generated to move the fluid around
but they are able to direct and control the speed of the
In this case one can distinguished between the circulating
fluid(blood) and the fluid that bathes the tissues (interstitial
fluid ISF)
Blood contains red blood cells and proteins in the palms
ISF lacks both rbcs and proteins
Keeping blood and ISF separate is quite important
Plasma proteins that escape from the blood and into
the ISF have to be collected up and returned to the
o The is a function of the lymphatic system
Together the blood and ISF make up the ECF(in both closed and
open systems)

ECF accounts for about 30% of body mass

But in this case blood is only about 5% of that total with
the remaining amount being the ISF

o The circulatory plan in the fish is fairly simple
Blood is pumped by the heart, goes to the gills and is
oxygenated, the oxygenated blood is carried to the
systemic tissues where it gives up its oxygen and the
blood returns to the heart
o The problem with this circ sys is that the heart has to pump
blood to the gills and the systemic circulation both of which are
sites of resistance
Heart must generate relatively high pressures
The gills as the first point of the system are going to see
relatively high points of pressure
They are able to withstand this because there is
water on the other side of the gill to counter balance
this pressure
Pressure of the system as a whole is limited to the
pressure that the gills can withstand
o Fish heart has 4 chambers in a series
Blood enters the sinus venoses the atrium
ventriclebulbous or conus arteriosis
The ventricle is where the main force generation
Bulbous and conus arteriosis are structurally
different but share the same purpose functionally
which is to maintain blood flow when the heart
o Same function as the aorta in a mammal
o Sinus venosus is only found in fish; it disappears it land living
Mammals and birth
o Essential there are 2 two chambered hearts
They are physically together in the same organ however
when broken down you have 2, 2 chambered hearts
o The left side of the heart and a right side.
Left side Always on the right when looking at a diagram
and right side is the left in the diagram
o The left side of the hear takes blood returning from the lungs
and pumps it out to the systemic circulation
o The right side of the heart takes blood returning from systemic
circulation(deoxygenated) and it pumps it to the lungs to be

o Each side of the heart both have an atrium and a ventricle

Left for systemic; right for pulmonary
o Low pressure is needed in the pulmonary circuit and high
pressure in the systemic circulation
This is possible because in effect they have 2 hearts
The left/right sides are operating independently
So the left ventricle can generate high pressure to send
blood to the systemic circulation
The right ventricle generates lower pressure to send blood
to the pulmonary circuit
o The only problem with this circulatory design is that blood flow
always has to go to both parts
As it comes back from the systemic circulation it has to go
back to the pulmonary circuit
There is no way to stop blood flow to the lungs (like when
you hold your breath...doesnt happen bud)
Intermittent air breathers- amphibians; reptiles+ air breathing fish
o Typically have lower metabolic rate
o Heart has 2 atria; one for blood collection from the lungs and
one that is collecting blood from the systemic circulation
o Typically has one ventricle- that directs the flow of blood to the
In reptiles its partially separated into a left and right
In amphibians there is only one ventricle
So deoxygenated blood from the tissues and oxygenated
blood from the lungs come into the same ventricle and are
pumped out to the lungs and tissue
o Even though there is a single ventricle the oxygenated and
deoxygenated blood is relatively separated
The mechs for this are not well understood
o Blood pressure in this animal as a whole is fairly low
Having a high bp will destroy its lung
Bp is limited by what the lungs can withstand
Low bp and low met rate animals
o The advantage of this is that when the animal is under water/
holding its breath the lungs become useless
So the system reduces blood flow to the lungs and
redistributes it to the skin where it can still take up
o When it resurfaces it redistributes the blood back to the blood
It has the capacity to shunt blood to diff organs depending
on its require
The presence of a single ventricle makes this possible

o Same idea is seen in reptiles though there is as light separation

in ventricles
o Same plan in air breathing fish
lung fish
o These fish can drown
o Obligate air breathing fish without any access to air they will
o They have a set out gills, a lung and a heart
o They have low met rates
o Tend to breath once every five minutes
During the rime spent underwater the animal is depleting
the oxygen in its lung
o There is a single 4 chambered heart (like a standard fish heart)
As the blood leaves the heart it is directed to the gills
The gills are separated into anterior gills which are not
used for O2 uptake and posterior gills which are used for
O2 uptake
From the posterior gills the blood moves up either to the
lung-if there is usable oxygen here- or if there is no
useable O2 in the lung the ductus opens up and directs
the blood from the gills to the systemic tissues
Oxygenated blood is directed by anterior gills to the
Deoxygenate blood is directed to posterior gills which
direct to lungs if there is air there or it its directed to the
The state of the ductus determines whether the blood goes
to the tissues or to the lungs
o The ventricle generates pressure for the lungs, gills, systemic
tissues. Systemic circuit as a whole
Must be a low pressure pump so the lungs are blown
These are low met rate animals
Lecture 3
Mammalian heart
Systemic half on the left has much thicker walls which allows for
higher levels of pressure
Blood returning from lung enters the left atrium and from there goes
into the left ventricle and is pumped out to the systemic tissue where
it is deoxygenated
o The deoxygenated blood is pumped into the right atrium and
then into the right ventricle and is pumped out to the lung
This heart is a chambered heart with contractile walls and based on
this there are valves that direct blood flow

In the mammalian heart there are vales between the atria and the
ventricles to control blood flow
Heart is made of cardiac muscle
o It is similar to skeletal muscle
o These are considered to be striated because they contain
o They are quite short and are connected end to end by
intercalated discs
In these discs one will find gap junctions
Which are communicating junctions that allow
electrical communication between the different cells
of the heart so they can contract as a unit
o The heart is innervated by the ANS
Within the ANS; there are 2 divisions
o Responsible for fight/flight alarm type
o Neurotransmitter of choice is noradrenaline
Interact with receptor in the tissues that
are adrenergic receptors
o B-1 receptors are more important for the heart
o Responsible for housekeeping or vegetative
o the rest and digest system
o Responsible for functions outside of alarm
o Neurotransmitter of choice is acetylcholine
Interacts with muscular genic receptors
in the effector organ
Main one for the heart is the m2
Both sympathetic and parasympathetic divisions are active
and in most tissues you can find innervation from both
Heart is innervated by both but they have opposite
o Sympathetic will increase heart rate to allow
for response in emergency situation
o Parasympathetic will slow it down
They have antagonistic effects
ANS allows good control of internal functions without any
conscious control
Most of the heart is made of cardiac muscle cells and these come in
two types

o 99% is made up of strongly contractile muscle cells

Packed full of sarcomeres; responsible for generating
o Other 1% are the cells of the conducting system
Pacemaker cells
They do not have a lot of contractile apparatus
They are specialized for controlling and coordinating
heart beat
Include the cells of the sinoatrial node, the atrial
ventricular node and the other cells forming the
conducting system that direct electrical activity over the
heart as a whole
The pacemaker cells have a resting potential that is quite unstable
o They have arresting potential that changes over time-pacemaker
potential and the presence of this changing resting potential is
the key characteristic of a pacemaker cell
o The pacemaker cells slowly depolarize because of the changing
membrane potential and when they reach a certain threshold
they trigger an action potential which the trigger a heartbeat/
contraction of heart
Initiate heart beat+ control it
o The pacemaker potential reflects the presence of the funny
It is considered to be funny because it is open and then
slowly closes and it functions as a Na+ leak channel
Allows Na+ to leak into the cell causing a gradual
o Also present in the cell are voltage gated Ca2+ channels(ttypes)
when the membrane potential depolarizes to the
threshold these channels open and it creates an action
o this is what initiates the heart beat in a vertebrate heart and
because it is a muscle cell for the initiation of this heart beat;
vertebrates hearts are considered to be myogenic
neurogenic: nervous activity activate heart beat
gap junctions allow electrical communication between cells
o once an action potential is fired within a cell that is then spread
to the other cells in their heart
o spreads from one heart cell to another by the gap junctions
o the heart rate is then set by the cells that depolarize most
quickly and these happen to be the cell of the sinuses venoses in
fish or sinoatrial node in mammals
pacemaker cells set the heart rate

the pacemaker cells in human sinoatrial node fire at rate of 100 time
per minute
o 100 depolarizations a minute
o This means the heart rate should be 100beats/min
o Heart beat is controlled by other mechanisms
There is input from the parasympathetic nervous system
that regulates heart rate
Sinoatrial node is in the atrium of the mammalian heart and it fires its
action potential first and this spreads across the atria real quick
o It slows down when it gets to the atrial ventricular node
o There is a layer of connective tissue between the two atria and
the two ventricle
o The cells of the atria re connected by gap junctions and the cells
of the ventricle are connected by gap junctions
But the atria and the ventricles do not communicate
This layer of connective tissue means there is no electrical
communication between the atria and ventricles except
through the atrial ventricular node
Conduction though this part of the system is quite slow
Once it gets through this system it speeds up
tremendously and shoots up through the ventricles at a
rate of 4or 5 m/s
o Measures the consequence of all the action potentials in the bod
fluid as a whole
It is the sum effect of all the action potentials happening
o Measured by placing electrodes upon the bodys surface not by
trying to impale a single heart cell to measure membrane
potential of that cell
o In the standard mammalian EKG there is a small wave called the
p wave
This shows the atria depolarizing
o There is a larger wave(QRS): this is the ventricles depolarizing
o Small t wave- this is the ventricular repolarization
o The repolarizing of the atrium is not seen because it occurs at
the same time that the ventricles depolarize and so it is masked
by this
o Value of the EKG tries to figure out what is going on in the heart
The heart as a pump
o Electrical activity in the conducting system has to be converted
into muscle contractions in 99% contractile part of the heart
To get these cells to contract the heart first has to be

A strong action potential is needed in the muscle

contractile cells (an ex can be found in slide 22/23)
There is stable resting potential; there is a strong
depolarization, then there is a plateau phase

Slide 22
Action potential f contractile cells
o Stable resting potential and when it is triggered it undergoes
depolarization and a voltage gated Na+ channel opens allowing
Na+ to pour into the cell and this gives you a strong
There is also a voltage gated Ca2+ channel that opens a
little bit later and this calcium channel stays open for a
while and that is how one gets the plateau region
This is an L-type gates calcium channel; in
pacemaker there is a t-type calcium channel
The flow of calcium into the cell triggers calcium
release from stores within the cell
o Not only have calcium moving into the cell
through the voltage gated channels you also
have calcium being released from stores inside
the cell
o The release of these stored calcium the
triggers the contraction of the cell
Calcium inflow + calcium
released=occurrence of muscle
o The cell then repolarizes and this represents a K+ channel that
is opened which allows K+ ions to flow out.
o Note in the diagram there is a prolonged plateau phase and a
refractory period
This is long refractory period is important because the
heart has to relax and refill with blood before it can
contract again
So the refractory period allows for the cells to rest
and reset as well as gives time for the heart to relax
and be refilled with blood
The long plateau in the action potential is necessary
because it allows all the muscle cells to contract at the
same time
This also makes sure that the heart cannot go into a
tetanic contraction-no muscle cramp in the heart
Refractory period: period where the cell is resting before
another AP can occur
Slide 23

o When heart is contracting
o When the heart is relaxing
Stroke volume
o Volume of blood pumped b the heart
o When heart contracts it ejects a volume of blood called the SV
and this is the difference in volume of the heart when it is
full(end diastolic volume) and after it contracts(end systolic
Most of the filling of the ventricles is due to venous pressure
o The valves between the atria and ventricle sis open and so the
blood flow through the valves to the ventricle from the atria
o This is driven by venous pressure; the atrial contraction helps
but it is only responsible for 1/3 of the volume in the ventricle
Venous pressure does all the work
For the heart to function properly output from the heart must match
the need for oxygen delivery
o This can be achieved by increasing heart rate or stroke volume
o Regulation of stroke volume is referred to as inotropic control
and its caused by both a mechanical relationship and neural and
hormonal control
The mechanical relationship is the frank-starling
See in slide 23
It shows that as end diastolic volume increases(as
heart if filled) SV also increases
o The fuller the heart the more forceful the
contraction and this ejects more blood
It is ;mechanical b/c as you fill the heart fuller it
stretches out muscle and this result in a more
forceful contraction
Very important because you always want to empty
the heart to the same point so by filling the heart
fuller you need more blood out of it to get back to
the same empty point
As heart rate goes higher there is less time needed
to fill it with blood and so the SV gets smaller there
hormonal and neural; mechanisms to accompany the
mechanical relationship and these mechs work to
stoop stroke volume from falling
o Neural/hormonal control of the heart
The sympathetic nerves innervate the ventricles and
strongly contractile muscle cells they release

noradrenaline which acts on 1 androgenic receptors and

this increases the force of contraction
As you increase sympathetic activity at any given
volume more blood is released from the heart; vice
versa for decreased sympathetic activity
Frank Starling relationship describes the change in
SV for a change in filling
o For any given level of sympathetic stimulation
there is a spec frank starling curve
o (only sympathetic control over SV)
o Circulating catecholamines
Sympathetic cells release noradrenaline
The cell that you find in the centre of the adrenal
gland are sympathetic neurons but instead of
releasing neurotransmitter onto the nerve it
releases the NTM into the blood and it circulates as
a hormone
Which is responsible for the fight or flight response
This too can regulate strong volume by acting on the
1 adrenergic receptors
heart rate
o controlling heart rate is the chonotropic effect
o the pacemaker is innervated by both the sympathetic and
parasympathetic systems
contains 1andrenrgic receptors activated by sympathetic
system or by circulating catecholamines
also contains m2 muscarinic receptors activate by ACh
which are activated by the parasympathetic system
sympathetic leads to an increase; parasympathetic causes
it to decrease
o Sympathetic nervous system
When the 1 receptor is activated by adrenaline or
noradrenaline it acts to open the sodium channels more to
get a faster depolarization(faster pacemaker potential)
o parasympathetic
When ACh acts on the m2 receptors it has the effect of
opening up potassium channels which allows positively
charged ions escape from the cell and there for it
depolarizes more slowly
The consequence of this is as low heart rate
o At rest both the sympathetic and parasympathetic systems are
o Sympathetic tone which helps set the resting heart rate
o Parasympathetic tone

Parasympathetic nerve that goes to heart is the vagus

nerve(can also be called vagal tone)
This also helps set the resting heart rate.
o Chonotropic effects only affects the pacemaker potential