Gas Exchange

and Circulation
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Introduction High Ratio of Surface Area to Volume

 Oxygen required for, and carbon dioxide produced

by, cellular respiration must be continuously
exchanged with the environment (Respiratory
system)

 In addition, these gases (along with wastes, nutrients,

and other types of molecules) must be transported
throughout the body (Circulatory system)
Figure 42.1

Ventilation
Unidirectional
Surface Area Distance
Buccal Pumping
5. Cellular Ram Ventilation
1. Ventilation 2. Diffusion 3. Circulation 4. Diffusion respiration

Environment

Respiratory surface
O2 O2
Blood
CO2 CO2

Mitochondria Bidirectional or tidal

Respiratory system Circulatory system
Air Ventilation :
Difference in partial • Buccal Pump (Positive
Difference in partial
pressure pressure ventilation)
pressure
• Aspiration Pump (Negative
pressure ventilation
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Dalton's law How Do Oxygen and Carbon Dioxide Behave in

Partial pressure
Air?
Gas Percent of total composition
(mm Hg)
• To understand how gases move by diffusion, it is
Nitrogen (N2) 78.6 597.4
Oxygen (O2) 20.9 158.8
important to express their presence in terms of
Water (H2O) 0.04 3.0 partial pressures instead of percentages
Carbon dioxide (CO2) 0.004 0.3
Others 0.0006 0.5
• The percentage of O2 does not change with elevation
Total composition/total
atmospheric pressure
100% 760.0 • At sea level and at the top of Mt. Everest, O2 is 21%

• The key difference is that far fewer molecules of

oxygen and other atmospheric gases are present per
unit volume of air at higher elevations that sea level
• Because atmospheric pressure is lower at higher
elevations
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Partial Pressures of Oxygen in Air How Do Oxygen and Carbon Dioxide Behave in
Water?
• Aquatic animals live in an environment that contains
much less oxygen than the environment inhabited
by terrestrial animals
• To extract a given amount of oxygen, an aquatic
animal has to process 30 times more water than the
amount of air a terrestrial animal breathes

• Water is about a thousand times denser than air and

much more viscous
• Water breathers have to expend more energy to
ventilate their respiratory surfaces than do air
breathers
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O2 and CO2 Behavior in Water What Affects the Amount of Gas in a Solution?

oxygen
• Oxygen and carbon dioxide diffuse into water from
the atmosphere, but the amount of gas that
dissolves depends on several factors:
1. The solubility of the gas in water
2. The temperature of the water
3. The presence of other solutes
4. The partial pressure of the gas in contact with
the water

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 Figure 42.3

Physical Parameters: The Law of Diffusion

Diffusion constant Difference in partial
• Fick’s law of diffusion states that the rate of (depends on pressure of gas on either
diffusion of a gas depends upon five parameters: solubility of gas side of barrier to diffusion
and temperature)
1. Solubility of the gas
2. Temperature
3. Surface area available for diffusion
4. Differences in partial pressures of the gas across
the gas-exchange surface Area for
gas exchange Distance
5. Thickness of the barrier in diffusion (thickness of
barrier to diffusion)

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The Respiratory Systems Direct diffusion across body surface

 In some animals the gas-exchange surface is the

skin,
 but in most species it is located in a specialized organ
like the lungs of tetrapods, Large Surface
Area to Volume
 the tracheae of insects, Wet enviroment
 or the gills found in mollusks, arthropods, and fish
Respiratory organs provide a greater surface area Respiratory organs provide a greater surface area
for gas exchange for gas exchange

Homeostasis broblem
Osmolarity
Salt balance
Dehydration

How Do Gills Work? How Do Gills Work?

• Gills are outgrowths of the body surface or throat,

used for gas exchange in aquatic animals
(a) External gills are in direct contact with water. (b) Internal gills must have water brought to them.

External
gills • Gills present an extremely large surface area for
oxygen to diffuse across an extremely thin
Internal gills
epithelium
• Among invertebrates, gills can be external or
5 mm 1 cm internal
Polychaeta • Fish gills are located on both sides of the head, and
in teleosts the gills consist of four arches

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Figure 43.21
How Do Gills Work? Gills are outgrowths of the body surface or throat

Coelom

Gills
Gills
Parapodium Tube foot
(functions as gill)

(a) Marine worm (b) Crayfish (c) Sea star

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How Do Fish Ventilate Their Gills? 1. Pumping 2. Ram ventilation

External
gills

1 cm
Figure 42.5

Each gill arch holds

many gill filaments O2-poor blood
Gill
Water IN O2-rich blood
arch
Lamella
Blood
vessels
Gill arch
The operculum Water OUT Water
has been removed flow Operculum
to reveal the gills Oxygen-
rich blood Water flow
Oxygen- Blood flow
to body
Detail of gill poor blood
filament: from heart

Oxygen-rich Countercurrent exchange

water IN PO (mm Hg) in water
2
Water flow 150 120 90 60 30
Gill lamella Gill filaments
Net diffu- 140 110 80 50 20
sion of O2
Blood flow PO (mm Hg)
2
in blood
Oxygen-poor
water OUT
Capillaries
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How Do Insect Tracheae Work?

Countercurrent flow Concurrent flow

(seen in fish gills) (not seen in fish gills)
• Insects have air-filled tubes called tracheae
Water flow over lamellae Water flow over lamellae • Tracheae open to the outside through pores called
(% oxygen) (% oxygen)
spiracles
• They can be closed to minimize water loss by
evaporation

• Air moves from the atmosphere through the

spiracles and into the trachea, which transports air
Blood flow through lamellae Blood flow through lamellae to the tissues and diffuses to the cells from there
(% oxygen) (% oxygen)
• In small insects, this process is sufficient to exchange
gases
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Figure 42.7
Tracheoles Mitochondria Muscle fiber

2.5 m
Spiracles

Body
cell
Tracheae Air
sac Tracheole
Air sacs

Trachea

Air spiracles
External opening
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How Do Insect Tracheae Work? Does physical activity affect air

movement through the tracheal system?
Air moves through the tracheal system faster
during physical activity.
Physical activity does not affect the rate

– In larger species, trachea alternately open and

of air movement.

close
as the wing muscles around them contract and Measure oxygen in
hawkmoth flight

relax Tether muscle during rest

and during flight.

– As a result, the volume of the tracheal system

Flying will increase ventilation of tracheal system,
causing increase in PO2 in wing muscle.
Flying will not increase
changes ventilation of tracheal system; PO2 in flight muscle will decline
steadily during flight.

– As the volume changes, so does the pressure

Flight

– Pressure and volume are inversely related

PO2 (kPa)
– The movement of gases is also aided by larger Recovery of
oxygen levels
as flying
trachea Rest
Initial drop continues
in oxygen
Rest

Time (min)

Muscular contractions may help ventilate the

tracheal system in at least some insects.
 Ground beetle

Tracheae

1 mm

Tracheae dilate when Tracheae compress when

muscles relax muscles contract

200 µm

Figure 42.10
The Biggest insect in the world

Percent O2

Cenozoic
Era
Ratio of Surface Area to Volume? Paleozoic Era
Geologic Time Scale Mesozoic Era
Era: Paleozoic
As evidence, researchers have shown that the diameter Period: Carboniferous*
of the tracheae in large beetles is proportionally much Present
larger than that of small beetles. mya
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 Lung Structure and Ventilation Vary among
How Do Vertebrate Lungs Work?
Species

 Air enters the body through the mouth and nose

 In frogs and other amphibians, the lung is a simple
 Trachea [`trekɪә] sac lined with blood vessels
 Bronchi [`brɑnkaɪ]
 In lizards, the internal lining of the lungs showing
 Bronchioles [`brɑηkɪ,ol] numerous faveoli the collectively give the lining a
honeycomb appearance.
 lung  Mammalian lungs are divided into tiny sacs called
alveoli [ӕl`viә,lɑɪ], which greatly increase the surface
 Animals with lungs include some fish, amphibians, area for gas exchange
reptiles, birds, and mammals – Humans have approximately 150 million alveoli
per lung

Lung Structure and Ventilation Vary among

Species

 Alveoli provide an interface between air and blood (a) Airways into the human lungs (b) Alveoli (c) The alveolar gas-exchange surface

that consists of Smallest

bronchiole Air

– A thin aqueous film Trachea

Lung
Air
Oxygen-rich
blood out
Oxygen
Aqueous film
Oxygen-poor Epithelium

– A layer of epithelial cells blood in

0.2
of alveolus
ECM
µm

– Some extracellular matrix (ECM) material Wall of

capillary

– The wall of a capillary Bronchi

Bronchioles Alveolus Capillaries Blood

extracellular matrix (ECM)

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Lung Structure and Ventilation Vary among Ventilation of the Human Lung
Species
• The mode of ventilation can vary among species • The pressure inside the human chest cavity is about
and animal groups 5 mmHg less than atmospheric pressure
• In the simple lungs of snails and spiders, air • This keeps the lung expanded
movement takes place by diffusion only
• Humans ventilate their lungs by changing pressure
• Vertebrates actively ventilate their lungs by pumping within their chest cavity between about –5 and –8
air via muscular contractions mmHg
• There are two mechanisms for pumping air: • The change in volume is caused by a downward
1. Positive pressure ventilation, used by frogs motion of a thin muscular sheet called the
diaphragm
2. Negative pressure ventilation, used by humans
and other mammals
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(a) Lungs expand and contract in response to changes in

pressure inside the chest cavity. Ventilation of the Human Lung

INHALATION EXHALATION • Two steps are involved—inhalation and exhalation:

Pressure
more
Pressure
less
1. During inhalation, the diaphragm moves down and
negative negative
the pressure in the chest cavity is lowered, causing
Diaphragm
the lungs to expand and air to move in
(b) Ventilatory forces can be modeled by a balloon in a jar.
2. During exhalation, as the diaphragm relaxes, the
肺內腔 chest cavity decreases and air is exhaled
Intrapulmonary cavity
• This is a passive process driven by the elastic recoil
Intrapleural cavity
of the lungs and chest wall as the diaphragm and rib
Pressure
more
Pressure
less
胸膜腔 muscles relax
negative negative

When the When the • It can become energy demanding during exercise
diaphragm diaphragm
is pulled is released,
down, the the balloon
balloon inflates. deflates.
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Pneumothorax

Surfactant
Infant respiratory distress syndrome
The Capacity of the Human Lung Avian Lung Lung, where
gas exchange
occurs

 About 450 mL of air moves into and out of the

Air sacs enable
one-way airflow
through lung

lungs in an average breath Follow one breath (in red) through the avian respiratory system

 Only about two-thirds of this volume actually

Trachea
1. Posterior
Inhalation
air sacs fill

participates in gas exchange, however, because

with outside air.

Posterior air sacs

150 mL of the air occupies dead space—portions Parabronchi

of the air passages, such as the trachea and Exhalation

Lung 2. Lungs fill
with air from

bronchi, that do not have a respiratory surface posterior sacs.

 Another unchanged volume – Residual volume (1.2l) Inhalation 3. Anterior air

sacs fill with
air from lungs.
Anterior
air sacs

Exhalation 4. Anterior air

sacs empty.

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Diagram of parabronchial anatomy

Anterior
air sacs parabronchi

dorsobronchus
posterior
air sacs
mesobronchus

ventrobronchus

© 2014 Pearson Education, Inc. cross-current exchange

 Ventilation of the Bird Lung
Crosscurrent Flow
• Three features make the avian ventilatory system
PO2 is usually higher than what efficient:
would be seen for concurrent, but
1. Dead space in bird lungs is restricted to a short
lower than countercurrent.
stretch of trachea, between the mouth and anterior
air sacs
2. Birds exchange gases during both inhalation and
exhalation
3. The capillaries in bird lungs cross the parabronchi
in a very efficient perpendicular arrangement that
allows for a crosscurrent pattern

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Homeostatic Control of Ventilation

• At rest, the mammalian rate of breathing is

established by the medullary respiratory center, an
area at the base of the brain
• The medullary respiratory center stimulates the rib
and diaphragm muscle to expand and contract

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Homeostatic Control of Ventilation
• During exercise, muscles take up more oxygen from
the blood
• The partial pressure of oxygen in the blood (PO2)
decreases
• At the same time, the muscles release larger
amounts of carbon dioxide, increasing its partial
pressure (PCO2) in the blood
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Homeostatic Control of Ventilation
• Increased CO2 reacts with water in the blood and
cerebrospinal fluid (CSF) to form carbonic acid,
H2CO3, which quickly dissociates into a hydrogen
ion, H+, and a bicarbonate ion, HCO3–
CO2 + H2O ↔ H2CO3 ↔ H+ + HCO3–
• The release of hydrogen ions lowers the blood and
CSF pH, which is sensed by specialized neurons,
leading to the medullary respiratory center and
increasing the breathing rate
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Structure and Function of Hemoglobin
• Red blood cells contain an oxygen-carrying molecule
called hemoglobin
• Hemoglobin consists of four polypeptide chains, each
of which binds to a nonprotein group called a heme
• Each heme contains an iron ion (Fe2+) that can bind to
an oxygen molecule
• Each hemoglobin molecule can thus bind up to four
oxygen molecules
• In blood, 98.5% of the oxygen is bound to hemoglobin
• The other 1.5% is dissolved in plasma
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 Figure 45.13
Hemoglobin
Structure and Function of Hemoglobin Each hemoglobin
molecule can
bind up to four O2
molecules of
oxygen
hemoglobin

O2 from
lung

98.5% of oxygen
binds to hemoglobin
in red blood cells

1.5% of oxygen
dissolves in
blood plasma

The rate of unloading depends on the

partial pressure of oxygen in the tissue
O2 to tissues
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Hemoglobin Unloads Oxygen to Muscle Tissue

% O2 changes

O2 saturation (%) of hemoglobin

slowly

 Blood leaving human lungs has a PO2 greater than

that of muscles and other tissues
– This difference creates a diffusion gradient that
unloads O2 from hemoglobin to the tissues
In this part of the graph, a
small change in PO2 leads to
% O2 a large change in how much
changes O2 unloads from hemoglobin
slowly cooperative binding

PO2 (mm Hg) in blood within tissue

Oxygen–Hemoglobin Equilibrium Curve, or
© 2014 Pearson Education, Inc. oxygen dissociation curve
Figure 42.16

How Do pH and Temperature Affect Oxygen

(a) With cooperative binding, there is a large
difference the amount of O2 delivered to resting
and exercising tissues.
Unloading from Hemoglobin?
• Hemoglobin is sensitive to changes in pH and
(b) Without cooperative binding, there is a
smaller difference in the amount of O2 delivered temperature
to resting and exercising tissues.

• Decreases in pH alter hemoglobin’s conformation

O2 saturation (%) of hemoglobin

O2 saturation (%) of hemoglobin

Resting
Exercising
Resting such that it is more likely to release O2 at all values of
Exercising PO2, causing a right shift in the hemoglobin curve
• This phenomenon is known as the Bohr shift
• The Bohr shift makes hemoglobin more likely to
release oxygen during exercise in which PCO2 is high,
PO2 (mm Hg) in blood within tissue
PO2 (mm Hg) in blood within tissue
pH is low, and tissues are under oxygen stress
• As temperature rises during exercise, the same
effect is observed, causing a right shift
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Figure 42.17 Figure 45.17

O2 saturation (%) of hemoglobin

O2 saturation (%) of hemoglobin

pH 7.4 pH 7.2
Fetal
Bohr shift hemoglobin

Adult (maternal)
hemoglobin

PO2 (mm Hg) in blood within tissue

PO2 (mm Hg) in blood within tissue
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Oxygen-storage function
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• enhanced gas-exchange efficiency
in the lungs,
• maintenance of O2 delivery and
oxygenation in the brain during
hypoxia,
• augmented O2 diffusion capacity in
peripheral tissues and a high
aerobic capacity.
high-altitude adaptation, at every step
in the O2 pathway of highland species.
• an enhanced hypoxic ventilatory
response,
• an effective breathing pattern,
• larger lungs,
• haemoglobin with a higher O2
affinity,
• alterations in the metabolic
properties of cardiac and skeletal
muscle.
Journal of Experimental Biology 2011 214: 2455-2462; doi: 10.1242/jeb.052548
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Respiratory Adaptations of Diving Mammals
• These animals have a high blood to body
volume ratio
• Deep-diving air breathers stockpile O2 and deplete it
slowly
• Diving mammals can store oxygen in their muscles in
myoglobin proteins
• Diving mammals also conserve oxygen by
– Changing their buoyancy to glide passively
– Decreasing blood supply to muscles
– Deriving ATP in muscles from fermentation once
oxygen is depleted
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 (A) An 8.5-h record of heart rate and depth profiles of a blue whale.

J. A. Goldbogen et al. PNAS 2019;116:50:25329-25332

©2019 by National Academy of Sciences

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Adaptions living in high altitude CO2 Transport and the Buffering of Blood pH

Transported
CO2 from Transport
in plasma
tissue protein in
cell membrane

Carbonic
anhydrase

Binds to
hemoglobin

Red blood cell

Plasma
2,3-diphosphoglyceric acid (2,3-DBG)
Carbonic Anhydrase and Hemoglobin When Blood Returns to the Lungs

 Carbonic anhydrase activity in red blood cells is
important for two reasons:
1. The protons produced by the carbonic anhydrase
reaction induce the Bohr shift, which makes
hemoglobin more likely to release oxygen
2. The PCO2 in blood drops when CO2 is converted to
bicarbonate, maintaining a strong partial-pressure
gradient favoring the entry of CO2 into red blood cells
 Much of the H+ produced by the dissociation of
carbonic acid is taken up by hemoglobin
– Therefore, hemoglobin also acts as a buffer,
minimizing changes in pH
 In the alveoli, a partial-pressure gradient favors the
diffusion of CO2 from plasma and RBCs to the
atmosphere

When Blood Returns to the Lungs

 Hemoglobin releases protons, which combine with

bicarbonate to form CO2
– Which then diffuses into the alveoli and is exhaled
from the lungs
 Hemoglobin picks up O2 during inhalation, and the
cycle begins again