B3.1 -Gas exchange as a vital function.

The need for gas exchange

All organisms carry out gas exchange, a vital function that is essential for life. Gas exchange
allows organisms to obtain the gases required for cellular processes such as aerobic
respiration and photosynthesis, and remove waste gases produced in metabolic reactions.
How are organisms, from the tiniest amoeba to a flowering plant to an elephant, adapted to
allow efficient gas exchange?

Gas exchange occurs via a process called diffusion. Diffusion is the random net movement of
particles from an area of higher concentration to an area of lower concentration, leading to
equilibrium – an equal distribution of particles.
Diffusion is the random net movement of particles from an area of higher concentration to
an area of lower concentration, leading to equilibrium – an equal distribution of particles.
Gas exchange occurs via a process called diffusion. Diffusion is the random net movement of
particles from an area of higher concentration to an area of lower concentration, leading to
equilibrium – an equal distribution of particles.
Organisms with a very high surface area-to-volume ratio can rely on diffusion across their
body surface to obtain and release gases, including single-celled organisms such as bacteria
and yeast. As organisms increase in size, surface area-to-volume ratio decreases and there is
a greater need for specialised structures to facilitate efficient gas exchange.

Properties of gas-exchange surfaces

Effective gas-exchange surfaces share some properties that make them efficient at
exchanging gases such as oxygen or carbon dioxide. These properties include: large surface
area – there is more membrane surface available for gases to diffuse across permeability –
the exchange surface must have pores or openings that allow gases to be exchanged across
its surface; for example, plants have stomata – small pores located mostly on the underside
of the leaf through which gases can move into and out of the leaf composed of a thin tissue
layer – so that there is a short distance across which gases need to move having a moist
surface – this helps to dissolve gases before they diffuse across the exchange surface; for
example, alveolar fluid in the alveoli concentration gradient – for diffusion to occur there
has to be a difference in concentration of the gas between two areas. The gas will move
from an area of higher concentration to an area of lower concentration.
Maintenance of concentration gradients
Concentration gradient refers to the difference in concentration of a substance between
two areas. The bigger the difference in concentration, the steeper the concentration
gradient, and the faster the rate of diffusion.
Adaptations in the exchange surfaces of animals to maintain steep concentration gradients
include: a dense network of blood vessels, which means that there is much opportunity for
substances to be exchanged between the surface and the blood.
Ventilation, which ensures the air or water rich in the desired gas is moved across the
exchange surface. Mammals inhale air into the lungs, and exhale to remove air from the
lungs. Fish have various mechanisms to ventilate the gills, including pumping water over the
gills through the mouth
Mammals are also able to maintain steep concentration gradients by separating oxygenated
and deoxygenated blood. This is possible because of a double circulatory system which
helps to ensure that blood transported to respiring cells is highly oxygenated.

Adaptations of the mammalian lungs for

gas exchange
In mammals, the lungs are the organs responsible for the exchange of gases between the air
and the bloodstream.

The structure of the lungs

Alveolar fluid/surfactant Type II pneumocytes secrete alveolar fluid which moistens the
surface of the alveoli, allowing gases to dissolve into the surfactant before diffusing across
the wall of the alveoli and capillary into the blood. Alveolar fluid contains surfactant, a
mixture of lipids and proteins (Figure 3). Surfactant has a key role in reducing the surface
tension of the alveoli and this helps to prevent alveolar collapse during exhalation.

A highly branched network of bronchioles

Each bronchiole branches into many alveoli, the exchange surface of the lungs. This
increases the surface area available for gas exchange, thus increasing the rate of gas
exchange. The high degree of branching also ensures that air is distributed throughout the
lungs. The small diameter of the bronchioles compared with the bronchi and trachea also
helps to slow down rate of the airflow, increasing the efficiency of gas exchange.
Extensive capillary beds surrounding the alveoli
Around the alveoli are many capillaries. This means that there is a very short distance for
oxygen to diffuse from the air in the alveoli into the blood and for carbon dioxide to diffuse
from the blood into the alveoli.

Many alveoli
Many alveoli create a larger surface area, totalling roughly the same area as a tennis court
for the average adult. This means that there is lots of area across which carbon dioxide and
oxygen can be exchanged.

Ventilation of the lungs

For gas exchange to occur in the alveoli, we need to get air into and out of the lungs. How
does this process of ventilation occur?

The following muscles are involved in ventilation;

• The diaphragm, a sheet of muscle found below the ribs.

• The intercostal muscles, a group of muscles found between and anchored to the ribs.
There are two types of intercostal muscles, the internal and external intercostal
muscles, which are antagonistic due to their opposing actions on the ribs during
breathing.
• The abdominal muscles.

The relationship between volume and pressure

When you inhale, the volume of your thoracic cavity increases. This means that particles
have more space to move, and so collide less frequently with each other and the walls of
the lungs. As a result, pressure in the lungs decreases, allowing air from outside the body to
rush into the lungs. When you exhale, the converse is true; the volume of your thoracic
cavity decreases, so particles have less space to move and collide with each other and the
walls of the lungs more frequently. This results in an increased pressure in the lungs, forcing
air out of the body.

Inspiration
During inspiration:

• The diaphragm contracts and moves downward.

• The external intercostal muscles contract, and the internal intercostal muscles relax,
causing the rib cage to move up and out.
• This increases the volume of the thoracic cavity, and so decreases the pressure in the
lungs.
• As a result, air moves down its pressure gradient into the lungs.

Expiration
During quiet breathing expiration:

• The diaphragm relaxes, moving upwards and inwards.

• The external intercostal muscles relax, and the internal intercostal muscles contract,
causing the rib cage to move down and in.
• This results in a decreased volume of the thoracic cavity, and so increases the
pressure in the thoracic cavity.
• As a result, air moves down its pressure gradient out of the lungs.

During forced breathing expiration:

• The diaphragm relaxes, moving upwards and inwards.

• The external intercostal muscles relax, and the internal intercostal muscles contract,
causing the rib cage to move down and in forcefully.
• This results in a decreased volume of the thoracic cavity, and so increases the
pressure in the thoracic cavity.
• As a result, air moves quickly down its pressure gradient and forcefully out of the
lungs.

The abdominal muscles also play a role in forced expiration, contracting and pushing up,
decreasing the volume of the thoracic cavity and pushing a greater volume of air out of the
lungs.
Measurements of lung volumes
Take a big breath, trying to inhale as much air as possible. The total volume of air in your
lungs is your total lung capacity. The average total lung capacity of a human is 6 litres.

Now exhale as much air as possible. The volume of air that you can forcibly exhale is
your forced vital capacity. The volume of air remaining in your lungs that cannot be exhaled
is the residual volume. Without the residual volume your lungs would collapse.

The volume of air that moves into and out of your lungs with each normal breath is your
tidal volume. After taking a normal breath, the additional volume of air that you can inhale
is the inspiratory reserve. After a normal exhale, the additional volume of air that you can
exhale is the expiratory reserve.

There are several factors that can affect human lung capacities, including:

• age – lung capacities tend to increase with age up to about 25 years of age
• body composition – larger people tend to have larger lungs and so have larger lung
capacities
• biological sex – in general, males tend to have larger lung capacities than females,
although it is important to note that there are more differences within each sex than
there are between the means of the two sexes
• respiratory disease – diseases such as chronic obstructive pulmonary disease (COPD)
and asthma can reduce lung capacities
• Levels of physical activity – regular exercise can increase lung capacities.

Lung volume can be measured using a pulmonary function test called spirometry, a
technique that involves a person breathing into a machine called a spirometer. The volume
and speed of exhaled air is measured and used to evaluate lung function and diagnose
respiratory conditions such as asthma and COPD (Chronic obstructive pulmonary disorder. A
group of diseases that affect the lungs and airways, including emphysema and bronchitis).

Adaptations of the leaves for gas exchange

The leaves are the main organs of gas exchange in a plant.

Dicotyledonous leaves are adapted for the exchange of oxygen, water vapour and carbon
dioxide in the following ways:

• Leaves are thin and flat, meaning they have a large surface area across which gases
can diffuse into and out of the leaf. It also means they have a large surface area to
absorb sunlight.
 • A waxy cuticle covers the cells of the upper epidermis. The waxy cuticle is
hydrophobic, so it is resistant to water and does not allow water to penetrate easily,
providing a waterproof barrier that prevents water loss and dehydration.
• Below the waxy cuticle and on the underside of the leaf is the epidermis. The
epidermis regulates the exchange of gases between the leaf and the external air
through small pores called stomata. Some stomata may also be found on the upper
side of the leaf and on the stem, but most are found on the underside of the leaf to
prevent excess water loss. The stomata are also more protected against physical
factors such as rain and physical contact on the underside of the leaf. The opening
and closing of stomata is regulated by the shape of guard cells. When the stomata
are open the rate of gas exchange increases, and when they are closed the rate of
gas exchange decreases.
• The spongy mesophyll layer provides many air spaces to allow gases to diffuse from
one part of the leaf to another.
• Veins transport water from the roots to the leaves. Veins, called vascular bundles,
are made of xylem and phloem tissue. Xylem tissue transports water and dissolved
minerals, and phloem tissue transports sugars and amino acids. This water will
diffuse out of the leaf in the process of transpiration.

Transpiration as a consequence of gas

exchange in the leaves
Transpiration is the loss of water vapour through the leaves, stems, and other above-ground
parts of the plant.

Capillary action occurs due to the cohesive and adhesive properties of water.

The diffusion of water out into the atmosphere through the stomata results in a negative
pressure, which pulls on water molecules in the xylem in a process called capillary action.
Capillary action occurs due to the cohesive and adhesive properties of water.

As well as allowing a plant to absorb water and nutrients from the soil, and transport these
throughout the plant, transpiration plays a role in regulating the temperature of the plant.
Water that is transpired can carry away excess heat, helping to keep the plant cool. On a
larger scale, the water that is transpired from many plants, such as the Amazon rainforest,
generates atmospheric water vapour which leads to cooling, airflow and rainfall.
Measuring the rate of transpiration
The rate of transpiration can be estimated using a photometer. A potometer measures the
volume of water taken up by a plant, which will be approximately equal to the volume of
water lost from the plant in the process of transpiration during that time.

Factors affecting the rate of transpiration:

- Temperature – The higher the temperature, the more kinetic energy water
molecules have, and so the faster they evaporate and diffuse out of the stomata.
High temperatures also increase the saturation point of the air outside the leaf,
meaning the air is able to hold more water vapour molecules.
- Humidity – Humidity refers to the amount of moisture in the air. The higher the
humidity of the surrounding air, the more water vapour molecules in the air outside
the leaf. As a result, there is a lower concentration gradient between the leaf and
the surrounding air, slowing the rate of diffusion of water vapour.
- Wind intensity – The higher the wind speed, the faster water molecules are moved
away from the leaf once transpiration has occurred. This results in an increased
concentration gradient between the stomata and the air outside the leaf, increasing
the rate of diffusion of water molecules.
- Light intensity – When light intensity is higher, guard cells cause the stomata to open
wider to allow more carbon dioxide to diffuse into the leaf for photosynthesis. This
means that more water evaporates from the plant.

Stomatal density
Stomatal density is the number of stomata in a particular unit of area of a leaf or other plant
organ.

Knowing the stomatal density is useful for a number of reasons, including:

❖ Plants with more stomata tend to have a higher rate of transpiration, and therefore
the stomatal density can indicate how efficiently a plant uses water.
❖ Changes in stomatal density in different environments can help us to understand the
effect of climates and climate change on plant physiology. Similarities and
differences in stomatal density can indicate phylogenetic relationships between
plants physiology.
❖ Similarities and differences in stomatal density can indicate phylogenetic
relationships between plants.
❖ Stomatal density of plant fossils can be used as an indicator of past environmental
conditions.
Determining stomatal density
To determine stomatal density, you need to know:

The area of your field of view. You can use a stage micrometre and graticule to determine
the measurements of your field of view. See section A2.2.1–2 for a reminder on using stage
micrometres and graticules. Once you have measured these values you can calculate the
area of your field of view using the equation area = πr2.

The number of stomata in the field of view.

Transport of oxygen (HL)

Vertebrates tend to have a high metabolic rate. Because of this they require a high rate of
oxygen delivery to, and carbon dioxide removal from, cells.

Adaptations of foetal and adult haemoglobin

Haemoglobin is a protein that is found in the red blood cells of vertebrates. Haemoglobin is
responsible for:

▪ Transporting oxygen from the lungs to respiring tissues

▪ Transporting carbon dioxide from respiring tissues back to the lungs where it can be
exhaled.

When haemoglobin is bound to oxygen, the molecule is called an oxyhaemoglobin complex.

Haemoglobin that is not bound to oxygen is called deoxyhaemoglobin.

Foetal haemoglobin
Foetal haemoglobin (HbF) is a type of haemoglobin produced by the developing foetus
during pregnancy.

It is the dominant form of haemoglobin during foetal development and will remain in the
infant until 6 months old, being gradually replaced by adult haemoglobin.

HbF has a quaternary structure, being composed of two alpha and two gamma polypeptide
chains, each of which contains a haem-group that can bind reversibly to oxygen.
The structure of foetal haemoglobin. The two alpha polypeptide chains are shown in green
and blue, and the two gamma polypeptide chains are shown in orange and pink. Each
polypeptide chain has a haem group, which contains an iron atom to which oxygen binds
reversibly.

Due to the presence of the gamma polypeptides, HbF has a higher affinity for oxygen than
does adult haemoglobin, which contains beta polypeptides instead.

This increases the efficiency with which the foetus obtains oxygen from the mother’s blood
across the placenta. The high affinity of HbF to oxygen is important for foetal development
and survival as the oxygen concentration in foetal blood circulation is much lower than the
oxygen concentration in the mother’s circulation.

Adult haemoglobin
Like HbF, adult haemoglobin (HbA) has a quaternary structure, being composed of four
subunits – two alpha and two beta polypeptide chains.
Each polypeptide chain contains a prosthetic group called haem group, which contains an
iron atom that can bind reversibly to a molecule of oxygen.
HbA has a lower affinity for oxygen than foetal haemoglobin. This is because it has a higher
affinity for a molecule called 2,3-bisphosphoglyceric acid (2,3 BPG). An organic phosphate
found in red blood cells. 2,3 BPG normally competes with oxygen for the binding to
haemoglobin.
The structure of adult haemoglobin. The two alpha polypeptide chains are shown in green
and blue, and the two beta polypeptide chains are shown in orange and pink. Each
polypeptide chain has a haem group, which contains an iron atom to which oxygen binds
reversibly.
You may also come across a third type of haemoglobin in your reading, myoglobin, the
transport molecule found in muscle tissue. Myoglobin consists of only one polypeptide
chain and has a much higher affinity for oxygen than either HbF or HbA.

Cooperative binding of oxygen to haem groups

When one oxygen binds to one haemoglobin subunit, this changes the conformation of the
molecule, increasing the affinity of haemoglobin to oxygen and therefore making it easier
for other oxygen molecules to bind to the remaining haem groups. This process is called
cooperative binding.
Conversely, when an oxygen molecule is released from a haem group, the conformation of
the haemoglobin changes, reducing the affinity of haemoglobin to oxygen.
The oxygen dissociation curve plots the partial pressure (concentration) of oxygen on the x-
axis, and the percentage saturation of haemoglobin with oxygen on the y-axis. Cooperative
binding of haemoglobin to oxygen results in a sigmoidal (S-shaped) oxygen dissociation
curve. The rate of oxygen uptake by haemoglobin increases rapidly as the partial pressure of
oxygen increases. But it eventually levels off as the haemoglobin becomes fully saturated
with oxygen in areas of higher partial pressures of oxygen, such as in the lungs.

Allosteric binding of carbon dioxide

As well as transporting oxygen, haemoglobin can transport a small amount of carbon
dioxide. Unlike oxygen, carbon dioxide binds to an allosteric region of haemoglobin, rather
than the haem group. This is known as allosteric binding and causes haemoglobin to form
carbaminohaemoglobin, undergoing a conformational change which results in a decreased
affinity for oxygen. This is important for ensuring that haemoglobin unloads oxygen in areas
of low partial pressure of oxygen, such as respiring tissues.

The Bohr shift

Cooperative binding of oxygen and the allosteric binding of carbon dioxide result in the
sigmoidal or ‘S’-shaped oxygen dissociation curve.

The oxygen dissociation curve can be shifted to the right with higher partial pressures of
carbon dioxide in the blood in a process called the Bohr shift.

Although some carbon dioxide will be transported while bound to haemoglobin, most will
be dissolved in the plasma, where, in its dissolved form, it will react with water to form
carbonic acid, lowering the pH of the blood. This, in turn, changes the tertiary structure of
haemoglobin, decreasing its affinity for oxygen and shifting the oxygen dissociation curve to
the right. The same effect can occur due to the presence of lactic acid in the blood and
increased blood temperature.

As a result of the decreased affinity for oxygen, haemoglobin releases more oxygen into the
tissues with higher partial pressures of carbon dioxide, such as those found in respiring
muscles. Conversely, it loads up more oxygen in areas with higher partial pressures of
oxygen, such as in the lungs. This shift to the right in the oxygen dissociation curve is called
the Bohr shift.

The Bohr shift occurs in both HbF and HbA, although to a lesser effect in HbF. It is believed
to play a role in facilitating oxygen transfer from the mother to the foetus to help to ensure
proper foetal development.
~In areas of high partial pressures of oxygen the affinity of haemoglobin to oxygen
decreases, shifting the oxygen dissociation curve to the right.