Circulation and Gas Exchange

Problem: How to
transport O2 and nutrients
effectively in multicellular
organisms

 Every organism must exchange

materials with its environment
Exchanges ultimately occur at the
cellular level by crossing the plasma
membrane

rns: 1. The delivery of O2 and nutrients

needed
2. Removal of CO2 and wastes

In unicellular organisms, these

exchanges occur directly with the
environment

 Small molecules can move between

cells and
their surroundings by diffusion
Diffusion is only efficient over small
distances because the time it takes to
diffuse is proportional to the square of
the distance
Tissue thickness,
Rate of Diffusion
Animals have developed cavities and
circulatory systems to transport gasses
and nutrients and to reduces the

The increase in animal

size through evolution
led to many steps in
delivering O2 and
nutrients to where they
are needed

Figure 42.1

Gills are an example of a specialized exchange

system in animals
O2 diffuses from the water into blood vessels
CO2 diffuses from blood into the water

Figure 42.2b

Flatworms have a gastrovascular

cavity and a flat body that minimizes
diffusion distances

Mouth

Gastrovascular 1 mm
Pharynx
cavity
(b) The planarian Dugesia, a flatworm

A gastrovascular cavity functions in both digestion

and distribution of substances throughout the body

The Circulatory System

All circulatory
The Circulatory System
systems, (open or
closed) have:
-Pump heart(s)
-Fluid blood or
hemolymph

-Vessels interconnecting vascular

components

Figure 42.3a

(a) An open circulatory system

Heart

Hemolymph in sinuses

Pores

Tubular heart

Hemolymph baths the organs directly

(b) A closed circulatory system

Heart
Interstitial fluid
Blood
Small
branch vessels
in each organ
Dorsal
vessel
(main
heart)

Auxiliary
hearts

Ventral vessels

blood is confined to vessels and is distinct from

the interstitial fluid

Organization of Vertebrate
Circulatory Systems
Humans and other vertebrates have
a closed circulatory system called
the cardiovascular system
Types of blood vessels: arteries,
arterioles, veins, venules, and
capillaries
Blood flow is one way in these
vessels. Arteries and veins are
distinguished by the direction of
blood flow, not by O2 content

Arteries and Veins

Veins carry blood to the
heart
Arteries carry blood away
from the heart

Copyright 2014 John Wiley & Sons, Inc. All rights reserved.

Single circulation:
fish
- Bony fishes, rays, and
sharks have single
circulation with a twochambered heart
-

In single circulation,
blood leaving the heart
passes through two
capillary beds before
returning
Blood enters through an
atrium and is pumped
out through a ventricle

Gill
capillaries

Artery
Heart:
Atrium (A)
Ventricle (V)
Vein

Key

Oxygen-rich blood
Oxygen-poor blood

Body
capillaries

Double circulation
Amphibians and reptiles,

Frogs and amphibians have

a three-chambered
heart: two atria and one
ventricle

Pulmocutaneous circuit
Lung
and skin
capillaries
Atrium
(A)

Atrium
(A)

Right
Left
Ventricle (V)
Systemic
capillaries
Key

Systemic circuit
Oxygen-rich blood
Oxygen-poor blood

Figure 42.4c

Double circulation:
mammal

Pulmonary circuit

Mammals (and birds) also

have a double circulation

Lung
capillaries

Double circulation maintains

higher blood pressure in the
organs than does single
circulation

A
V

Right

Left

Systemic
capillaries
Systemic circuit
Key

Oxygen-rich blood
Oxygen-poor blood

 Mammals and birds have a four-chambered

heart with two atria and two ventricles
The left side of the heart pumps and receives only
oxygen-rich blood, while the right side receives and
pumps only oxygen-poor blood

Mammals (and birds) have two

circulation paths
Pulmonary circuit blood leaves
the right ventricle, goes to the lungs,
then oxygenated blood enters the
left atrium.
Systemic circuit oxygenated
blood leaves the left ventricle,
pumped through the body, then
deoxygenated blood returns to the
right atrium.

The Pulmonary Circuit

Blood begins its flow with the right
ventricle pumping blood to the lungs
via the pulmonary arteries
In the lungs, the blood loads O2 and
unloads CO2
Oxygen-rich blood from the lungs
enters the heart at the left atrium
via the pulmonary veins

Figure 42.5

Superior
vena cava

Capillaries of
head and
forelimbs

Pulmonary
artery
Capillaries
of right lung

Pulmonary
artery
Aorta

6
2

11

Pulmonary
vein
Right atrium
Right ventricle

Capillaries
of left lung

Pulmonary
vein
Left atrium
Left ventricle

10

Aorta

Inferior
vena cava

Capillaries of
abdominal organs
and hind limbs

The Systemic Circuit

Blood is pumped through the aorta to the
body tissues by the left ventricle
(strongest muscle of the heart)
Blood returns to the heart through the
superior vena cava (blood from head,
neck, and forelimbs) and inferior vena
cava (blood from trunk and hind limbs)
The superior vena cava and inferior vena
cava flow into the right atrium

ings to Remember about a Mammalian H

The mammalian heart has a double circulation:
heart
lungs
heart
systemic
The two atria have relatively thin walls and serve as
collection chambers for blood returning to
the heart
The ventricles have thicker walls ( left is the thickest)
and contract much more forcefully

The aorta provides blood to the

heart through the coronary arteries

Pericardium

The heart is enclosed and held in place

by the pericardium

Copyright 2014 John Wiley & Sons, Inc. All rights reserved.

Valves open and close to allow

blood flow between chambers
Aorta

Pulmonary artery

Pulmonary
artery

Right
atrium

Left
atrium

Semilunar
(pulmonary) valve

Semilunar (aortic)
Valve

Atrioventricular
(AV) valve
(tricuspid)

Atrioventricular
(AV) valve
(bicuspid or mitral)
Right
ventricle

Left
ventricle

rt Valves Open and Close in Response to Press

Differences

Fig. 9-19, p.377

Valves are One-way

Copyright 2014 John Wiley & Sons, Inc. All rights reserved.

CUSP OF
TRICUSPID
VALVE
CHORDAE
TENDINEAE

PAPILLARY
MUSCLE

(c) Tricuspid valve open

Copyright 2014 John Wiley & Sons, Inc. All rights reserved.

The heart contracts and relaxes in a rhythmic cycle

called the cardiac cycle

Systole - The contraction, or

pumping, phase
Diastole - The relaxation, or filling,
phase
*One
complete cycle of pumping and filling
is called the Cardiac Cycle
(In reference to ventricles)

2 Atrial systole and

ventricular diastole

The Cardiac Cycle

one cardiac cycle = 0.8 sec.

1 Atrial and
ventricular diastole

Lub Dub
0.1
sec

Lub = onset of systole

(AV valve closes)
Dub = onset of diastole
(Aortic valve closes)

0.4
sec

0.3 sec

3 Ventricular systole
and atrial diastole

How much blood moves through the hear

heart rate, -the number of beats per minute:
72 beats/min
stroke volume- the amount of blood
pumped in a single contraction:
70 mL
cardiac output - the volume of blood
pumped into the systemic circulation per
minute
5 L/min = 72 beats/min x 70 mL/beat

diac output = heart rate x stroke vo

What determines Cardiac Output?

CO(mL/min)

Stroke Volume and Heart

Rate
- When the heart beats faster,
more blood is pumped into the
circulation per minute
- When the heart muscles
contract harder, more blood is
emptied into system

The Conduction System

Cardiac muscle cells are self-excitable,
and therefore, autorhythmic
Cardiac muscle cells repeatedly
generate spontaneous action potentials
that then trigger heart contractions

Copyright 2014 John Wiley & Sons, Inc. All rights reserved.

Muscle cells are separated by intercalated

discs which contain gap-junctions
Intercalated discs
Electrical
impulses
travel
through gapjunctions

Action potentials
travels through
gap junctions

Atria and ventricles

have separate
conductive muscles
- no gap junctions
between them

Maintaining the Hearts Rhythmic

Beat

Autorhythmatic contract and relax without signal

from nervous system
Signals (yellow)
from SA node
spread through
atria.

SA node
(pacemaker)
ECG

The sinoatrial (SA)

node, or pacemaker,
sets the rate and timing
at which cardiac muscle
cells contract

Impulses from the SA node travel to the

atrioventricular (AV) node, where
they are delayed
1 Signals (yellow)
from SA node
spread through
atria.

SA node
(pacemaker)

ECG

Signals are
delayed at AV
node.

AV
node

Next, the impulses travel through the

bundle branches (Bundle of His) to the
heart apex, where
1 Signals (yellow)
from SA node
spread through
atria.

SA node
(pacemaker)

ECG

2 Signals are
delayed at AV
node.

AV
node

3 Bundle branches
pass signals to
heart apex.

Bundle
branches

Heart
apex

 they travel to the Purkinje fibers which

make the ventricles contract.
1 Signals (yellow)
from SA node
spread through
atria.

SA node
(pacemaker)

ECG

2 Signals are
delayed at AV
node.

AV
node

3 Bundle branches
pass signals to
heart apex.

Bundle
branches

Heart
apex

4 Signals
spread
throughout
ventricles.

Purkinje
fibers

The Electrocardiogram, (ECG or

EKG)
Electrical activity in body fluids: the sum of
cardiac muscle activity
P wave atrial contraction
QRS complex ventricular
contraction
T wave ventricular
relaxation

Nervous system regulation

of the pacemaker
The sympathetic division speeds
up the pacemaker norepinephrine
and epinephrine
The parasympathetic division
slows down the pacemaker acetylcholine

Vessels in the circulatory

system

Arteries

Arterioles
Capillaries

Back to the heart

venules
Veins

Arteries have thicker walls than veins to

accommodate the high pressure of blood
pumped from the heart

Vein

Artery

Red blood cells

Figure 42.9c

Capillaries are only slightly wider than a

red
blood cell

Capillaries have thin walls, the endothelium plus its

basal lamina, to facilitate the exchange of materials
Red blood cell

Capillary

The epithelial layer that lines blood vessels is

called the endothelium. It is smooth and
minimizes resistance.

Valve

Basal lamina
Endothelium
Smooth
muscle
Connective
Capillary
tissue

Endothelium
Smooth
muscle
Connective
tissue

Artery

Vein

Arteriole

Venule

Velocity of blood is lower in veins than in

arteries. So, why does blood continue to
flow towards heart?
Direction of blood flow
in vein (toward heart)
Valve (open)

One-way valves
prevent backflow
Skeletal muscle
contraction also
moves blood along

Skeletal muscle

Valve (closed)

Blood Flow Velocity

Speed in which blood flows is
inversely proportional the total
cross-sectional area of the vessels
Capillaries have the highest
cumulative cross-sectional area,
therefore blood moves slowest
through the capillaries.

The the
crosssectional
area, the
the velocity of
blood

50
40
30
20
10
0
Systolic
pressure

Pressure is
greatest in
the Aorta
Venae
cavae

Veins

Venules

Capillaries

Arterioles

Diastolic
pressure
Arteries

120
100
80
60
40
20
0

Aorta

Pressure
(mm Hg)

Velocity
(cm/sec)

Area (cm2)

Collectively,
capillaries
have the
greatest area

5,000
4,000
3,000
2,000
1,000
0

 Velocity of blood flow is slowest in the capillary

beds, as a result of the high resistance and large
total cross-sectional area
Blood flow in capillaries is necessarily slow for
exchange of materials

Capillary Function
Two mechanisms regulate
distribution of blood in capillary beds
Constriction or dilation of arterioles
that supply
capillary beds (regulated by nerve
impulses , hormones, or other
chemicals)
Precapillary sphincters that control
flow of blood
between arterioles and venules

Figure 42.13

Precapillary sphincters

Arteriole

Thoroughfare
channel

Capillaries
Venule

(a) Sphincters relaxed

Arteriole
(b) Sphincters contracted

Venule

Endothelial
cell

Endothelial cells of capillaries fit together like a

jigsaw puzzle

Lipid soluble substances

do not need the pores,
but pass through the cells
directly

Pores
Capillary

Allows passage
of water soluble

(a)

substances

The exchange of substances across

capillary walls.
-Regulated by blood pressure and osmotic
pressure
-Blood proteins rarely pass through endothelium
and help maintain osmotic pressure
INTERSTITIAL
FLUID

Net fluid movement out

Body cell

Blood
pressure
Osmotic
pressure

Arterial end
of capillary

Direction of blood flow

Venous end
of capillary

Blood
Composition and
Function
Plasma = 55% of whole blood
Platelets
Buffy coat
<1%
White blood cells

Packed cell
volume, or
hematocrit

Red blood cells =

45% of whole blood
% hematocrit varies with species and with
stress levels.

Fig. 9-3, p.361

What is Plasma?
Plasma contains inorganic salts, sometimes
called electrolytes
Plasma proteins influence blood pH and
help maintain osmotic balance between
blood and interstitial fluid
Particular plasma proteins function in lipid
transport, immunity, and blood clotting
Plasma is similar in composition to
interstitial
fluid, but plasma has a much higher protein
concentration

Cellular Elements of Blood

Suspended in blood plasma are two types
of cells
Red blood cells (erythrocytes) transport O2.
The hormone erythropoietin (EPO)
stimulates erythrocyte production when O2
delivery is low
White blood cells (leukocytes) function in
defense

Platelets are fragments of cells that are

involved in clotting

Pluripotent Stem Cells give rise to

Lymphoid and Myeloid cell types

Erythrocytes
Red blood cells, or erythrocytes, are the
most numerous blood cells
They contain hemoglobin, the ironcontaining protein that transports O2
Each molecule of hemoglobin binds up to
four molecules of O2
In mammals, mature erythrocytes
lack nuclei and mitochondria
*One erythrocyte contains ~
250 billion hemoglobin
molecules

Iron
Heme
Hemoglobin

Red Blood Cells

8m

Have no nucleus or
membrane bound
organelles
Surface view

How would the

biconcave shape of the
cells benefit diffusion of
O2?

Sectioned view
(a) RBC shape

RBCS (Erythrocytes)

Lack intracellular organelles necessary for cellular repair, growth, division

Short Life Span (~120 days)

Ruptured RBCs are destroyed in spleen, liver or bone marrow

Phagocytic WBCs clear the debris

Respiration
Gas exchange supplies O2 for cellular
respiration and disposes of CO2
Gas exchange occurs across specialized
respiratory surfaces

Ventilation moves the respiratory

medium over the respiratory surface

Coelom
Gills
Parapodium
(functions as gill)
(a) Marine worm

Gills

(b) Crayfish

Tube foot
(c) Sea star

Figure 42.22a

Gill
arch
Blood
vessels
Gill arch
Water
flow

Operculum

In fish gills, more than

80% of the O2
dissolved in the water
is removed as water
passes over the

Gill filaments

Tracheoles Mitochondria

Muscle fiber

2.5 m

Figure 42.23

Tracheae
Air sacs

Body
cell

Air
sac

Tracheole

Trachea
External opening (spiracles)

Air

Book lungs of
the Spider

Mammalian Respiratory System

Nasal
cavity

Pharynx
Larynx

Left lung

(Esophagus)
Trachea
Right lung
Bronchus
Bronchiole
Diaphragm
(Heart)

Lungs are encased in a membraneous

pleural sac
that is filled
with fluid
between its
cavity
Lung

Lollipop

Water-filled
balloon

Pleural sac
(a)

Left pleural sac

Right pleural sac

Thoracic wall

Left
lung

Right
lung

Parietal pleura
Visceral pleura
Pleural cavity filled
with intrapleural fluid

Diaphragm
(b)

Fig. 11-14, p.477

The right lung has three lobes, the

left lung has two lobes

Apex
Superior lobe

ANTERIOR

Oblique fissure
Inferior lobe
POSTERIOR
(b) Lateral view of right
lung

Inferior lobe

Middle lobe
Base

POSTERIOR
(c) Lateral view of left
lung

Trachea

Bronchus

Bronchiole

(Heart)

Diaphragm

Anatomy of Alveoli
Branch of
pulmonary vein
(oxygen-rich
blood)
Terminal
bronchiole

Branch of
pulmonary artery
(oxygen-poor
blood)

Alveoli

exchange occurs in the alveoli

Capillaries

200 300 million alveoli in a human lung

 Alveoli lack cilia and are susceptible

to contamination
Secretions called surfactants coat
the surface of the alveoli
Preterm babies lack surfactant and
are vulnerable to respiratory distress
syndrome; treatment is provided by
artificial surfactants

How much air can the lungs hold?

56L
The the volume of air inhaled with
each breath - ~ 500ml (Tidal Volume)
Forced inhalation/exhalation ~ 4 L
(Vital capacity)
There is always air left behind in the
lungs ~1.2L (residual volume)

Mammals ventilate their lungs by negative

pressure breathing, which pulls air into the
lungs
Rib cage
Rib cage
gets smaller
as rib muscles
relax.

expands as
rib muscles
contract.

Lung
Diaphragm
1 INHALATION: Diaphragm

contracts (moves down).

2 EXHALATION: Diaphragm

relaxes (moves up).

Lung volume increases as the rib muscles

and diaphragm contract

Partial Pressure Gradients in Gas

Exchange
Partial pressure is the pressure
exerted by a particular gas in a
mixture of gases
Gases undergo net diffusion from a
region of higher partial pressure to a
region of lower
partial pressure

When the chest expands, the volume

of the lungs increases. The pressure
exerted by oxygen within the lungs
decreases, therefore oxygen diffuses
into the
O2P lungs
= 160 mmHg
02

Right pleural sac

P02 < 160 mmHg

Diaphragm

Figure 42.29
160
120
27

6 Exhaled air

Inhaled air

PO 2 PCO 2

PO 2 PCO 2
Alveolar
epithelial
cells

40

CO2

O2

Alveolar
spaces 2
Alveolar
capillaries

Blood
entering
alveolar
capillaries

45

PO 2 PCO 2

0.2

104
40

PO 2 PCO 2

5
Pulmonary
veins and
systemic 3
arteries

Pulmonary
arteries
and systemic
veins

104
40

PO 2 PCO 2

<40

>45

PO 2 PCO 2

4 Body
tissue

CO2

O2

Systemic
capillaries

Respiratory Pigments
Respiratory pigments, proteins that
transport oxygen, greatly increase the
amount of oxygen that blood can
carry
Arthropods and many molluscs have
hemocyanin with copper as the
oxygen-binding component
In vertebrates, hemoglobin is
contained within erythrocytes

Hemoglobin (Hb)

Polypeptide chain

Each hemoglobin
molecule binds to
four O2 molecules

Structure:
Four polypeptide
chains
Four iron-containing
heme groups

Iron
atom
within
heme
group

Polypeptide chain
Heme groups

Polypeptide chain

PO2 =
PO2
O

O
O
O

PO2 > PO2

= O2

O
O

O
O
O
O

O
O

O
O

PO2 = PO2

O O
O O

= Hb

Only unbound O2 contributes to PO2

O2 saturation of hemoglobin (%)

Figure 42.30a

100

O2 unloaded
to tissues
at rest

80

O2 unloaded
to tissues
during
exercise

60
40
20
0

20

40

60

Tissues during Tissues

exercise
at rest
PO 2 (mm Hg)

80

100
Lungs

(a) PO 2 and hemoglobin dissociation at pH 7.4

O2 saturation of hemoglobin (%)

Bohr Shift
100
pH 7.4
80

pH 7.2
Hemoglobin
retains less
O2 at lower pH
(higher CO2
concentration)

60
40
20
0

20

40

60

80

PO 2 (mm Hg)
(b) pH and hemoglobin dissociation

100

Carbon Dioxide Transport

Some CO2 from respiring cells diffuses into
the blood and is transported in blood
plasma, bound to hemoglobin
The remainder diffuses into erythrocytes
and reacts with water to form H2CO3, which
dissociates into H+ and bicarbonate ions
(HCO3)
In the lungs the relative partial pressures of
CO2 favor the net diffusion of CO2 out of the
blood

Control of Breathing in
Humans

In humans, breathing is usually

regulated by involuntary mechanisms
The breathing control centers are found
in the medulla oblongata of the brain
The medulla regulates the rate and
depth of breathing in response to pH
changes in the cerebrospinal fluid

Figure 42.28

NORMAL BLOOD pH
(about 7.4)
Blood CO2 level falls
and pH rises.

Medulla detects
decrease in pH of
cerebrospinal fluid.
Cerebrospinal
fluid
Signals from
medulla to rib
muscles and
diaphragm
increase rate
and depth of
ventilation.

Carotid
arteries
Aorta

Medulla
oblongata

Blood pH falls
due to rising levels of
CO2 in tissues (such as
when exercising).

Medulla receives
signals from major
blood vessels.

Sensors in major
blood vessels
detect decrease
in blood pH.

 Sensors in the aorta and carotid arteries

monitor O2 and CO2 concentrations in the
blood
These signal the breathing control centers,
which respond as needed
Additional modulation of breathing takes
place in the pons, next to the medulla

CO2 is the main determinant of pH in

the blood
How is pH regulated?