HMM/SCM1414-Biology I

CHAPTER 8

GASEOUS EXCHANGE
• Gaseous exchange - uptake of O2 from
environment and discharge of CO2 to
environment.
° Process linked to production of ATP in

cellular respiration.
• Involves respiratory and circulatory

systems - provides O2 for aerobic cellular

respiration and removes CO2.
• Respiratory medium - source of O2:

° Air (atmosphere) - 21% O2 (by volume).

° Water – O2 levels in bodies of water vary,

but are always much less than an

equivalent volume of air.

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8.1 Gaseous Exchange in animal

8.1.1 Respiratory Systems in Animals

(a) Respiratory Surface

• Respiratory surface - the part of an

animal where gases are exchanged with
environment.
° Movements of CO2 and O2 across

respiratory surface occur by diffusion.

• Fick’s Law of Diffusion: "Rate of

transfer of a gas (dV/dt) through a sheet of
tissue is proportional to tissue area (A) and
difference in gas partial pressure between
the 2 sides (P1 – P2) and inversely
proportional to tissue thickness (T)."

Area x Diffusion constant x

(Partial Pressure 1 – Partial
Pressure 2)
Volume of gas =
(per unit time) Thickness

dV = A * D * (P1 – P2)
dt T

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• Fick's Law governs transfer rate of gases

through tissue.
° For respiration, law governs transfer
rate of O2 from alveoli to blood across
thin blood gas barrier, and CO2 in
opposite direction.
° CO2 diffuses ≈ 20x more rapidly than
O2 through tissue sheets.
° Reason: CO2 has higher solubility,
thus increasing diffusion constant (D).
° Diffusion constant is proportional to
solubility divided by square root of
molecular weight.

• Characteristics of respiratory surfaces that

maximize rate of gas exchange:

(1) Large surface area

The larger the animal and the more active
it is, the larger the surface area required
for gaseous exchange. The greater the
area exposed to the environment, the
greater the rate of diffusion.

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(2) Thin
One cell thick – diffusion rapid over very
short distance only. If distance is
doubled, diffusion takes 4x longer.

(3) Moist
Gases must dissolve in water before
diffusing across respiratory surfaces.

(4) A good blood supply

An efficient transport will ensure that
gases will be taken away from
respiratory surface quickly so
maintaining a large concentration
gradient.

(5) A good ventilation gradient

A pumping system, which will
continuously, deliver gas to respiratory
surfaces. This will maintain a large
diffusion gradient across respiratory
surface.

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(b) Types of Respiratory Surfaces

• Structure of respiratory surface depends

on size of organism, whether it lives in
water or on land, and on its metabolic
demands.

(i) Body Surface

• Protists & other unicellular organisms – gas

exchange over entire surface area.
• Simple animals (sponges, cnidarians, &

flatworms) - plasma membrane of every cell

in body is close to outside environment for
gas exchange.
• However, in most animals, bulk of body
lacks direct access to respiratory medium.
° Respiratory surface is a thin, moist

epithelium that separates respiratory

medium from blood or capillaries, which
transport gases to and from rest of body.
(Refer Figure 42.18, Campbell)
• Some animals (earthworms and some

amphibians) use entire outer skin as a

respiratory organ.
° Have dense net of capillaries below moist

skin.

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 HMM/SCM1414-Biology I

° Since respiratory surface must be moist,

their habitats limited to water or damp
places.
° Animals using moist skin as their only
respiratory organ are usually small and
are either long and thin or flat in shape,
with high ratio of surface area to volume.

(ii) Gills

• Respiratory adaptations of most aquatic

animals.
• Gills - outfoldings of body surface that

are suspended in water.

• Water as a respiratory medium:

° Advantage: Cell membranes of respiratory

surface are kept moist since gills are

surrounded by aqueous environment.
° Disadvantage: Low O2 concentrations in

water, especially in warmer and saltier

environments.
• Ventilation - increases flow of

respiratory medium over respiratory

surface, ensuring strong diffusion gradient
exists between gill surface and
environment.

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° Without ventilation, a region of low O2 &

high CO2 concentrations forms around gill
as it exchanges gas with environment.
° Fish gills ventilated by a current of water
that enters mouth, passes through slits in
pharynx, flows over gills, and exits body.
(Figure 42.20, Campbell)
Because water is dense and contains


little O2 per unit volume, fishes must

expend considerable energy in
ventilating their gills.
• Gas exchange at gill surface enhanced by

opposing flows of water and blood at gills -

countercurrent exchange (Figure 42.21,
Campbell).
° As blood moves through gill capillary, it
becomes more and more loaded with O2,
but it simultaneously encounters water
with even higher O2 concentrations
because it is just beginning its passage
over the gills.
° All along the gill capillary, there is a
diffusion gradient favoring transfer of O2
from water to blood.
° The countercurrent exchange mechanism
is so efficient that gills can remove more
than 80% of O2 from water to blood.

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• Gills unsuited for animal living on land:

° A large surface of wet membrane
exposed to air would lose too much water
by evaporation.
° Gills would collapse as their fine
filaments, no longer supported by water,
cling together, reducing surface area for
exchange.

(iii) Tracheal System

• Tracheal systems & lungs - respiratory

adaptations of terrestrial animals.
• Advantages of air over water as

respiratory medium:
° Higher O2 concentration.

° O2 and CO2 diffuse much faster in air -

respiratory surfaces do not have to be

ventilated as thoroughly as gills.
° If ventilation is required, less energy

needed because air is lighter and easier

to pump and much less volume needs to
be breathed to obtain an equal amount of
O2 .

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• Disadvantages:
° Respiratory surface, which must be large

and moist, continuously loses water to air

by evaporation.
° Problem greatly reduced by having

respiratory surface folded into body.

• Tracheal system of insects composed

of air tubes that branch throughout body.
° Largest tubes (tracheae) open to outside

while finest branches (tracheoles)

extend to surface of nearly every cell
where gas is exchanged by diffusion
across moist epithelium that lines the
terminal ends. (Figure 42.22, Campbell)
° The open circulatory system does not

transport O2 and CO2 since all cells are

located close to respiratory medium.
• For small insect, diffusion through

trachea brings in enough O2 and removes

enough CO2 to support cellular respiration.
° Larger insects with higher energy

demands ventilate their tracheal systems

with rhythmic body movements that
compress and expand the air tubes.

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° An insect in flight has very high metabolic

rate, consuming 10 to 200 times more O2
than it does at rest.
° Alternating contraction and relaxation of
flight muscles compresses and expands
body, rapidly pumping air through
tracheal system.
° Flight muscles are packed with

mitochondria, supplied with O2 by

tracheal tubes.

(iv) Lungs

• Lungs restricted to one location.

° Since respiratory surface of lung is not in

direct contact with all other parts of

body, circulatory system transports gases
between lungs and rest of body.
° Have dense net of capillaries under

epithelium that forms respiratory surface.

° In spiders, terrestrial snails, &

vertebrates.
• Among vertebrates, amphibians have
relatively small lungs that do not provide a
large surface, and many lack lungs
altogether.

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° Rely on diffusion across other body

surfaces, especially moist skin, for gas
exchange.
• In contrast, most reptiles (including all
birds) and all mammals rely entirely on
lungs for gas exchange.
° Turtles may supplement lung breathing
with gas exchange across moist
epithelial surfaces in mouth and anus.
° Lungs and air-breathing have evolved in a

few fish species (lungfishes) as

adaptations to living in O2-poor water or
to spending time exposed to air.
° Besides lungs, birds have eight or nine air

sacs that increase respiratory efficiency.

• In general, size and complexity of lungs
are correlated with an animal’s metabolic
rate (and hence rate of gas exchange).
° For example, lungs of endotherms have a
greater area of exchange surface than
lungs of similar-sized ectotherms.

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8.1.2 Human Respiratory System

• Lungs located in thoracic (chest) cavity.

(Figure 42.23, Campbell)
• Have spongy texture and are lined with a
moist epithelium that functions as
respiratory surface.
• A system of branching ducts conveys air
to the lungs.
• Air enters through nostrils and is then

filtered by hairs, warmed and humidified,

and sampled for odors as it flows through
nasal cavity.
° Nasal cavity leads to pharynx, an

intersection where the paths for air &

food cross.
° Pharynx leads to larynx, a cartilaginous

structure adapted for sound production.

° During swallowing, epiglottis covers

entrance to larynx, the glottis.

• From larynx, air passes into trachea,

whose shape is maintained by rings of

cartilage.
° Trachea branches into two bronchi, one

leading into each lung.

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° Within lung, each bronchus branches

repeatedly into finer and finer tubes,
bronchioles.
• Epithelium lining major branches of

respiratory tree covered by cilia and thin

film of mucus.
° Mucus traps dust, pollen, and other

particulate contaminants, and beating

cilia move mucus upward to pharynx,
where it is swallowed.
• Bronchioles lead to a cluster of air sacs,

the alveoli.
° Gas exchange occurs across thin
epithelium of alveoli.
° Total surface area of alveoli in human ≈

100 m2 - sufficient to carry out gas

exchange for whole body.
° O2 in air entering alveoli dissolves in

moist film and rapidly diffuses across

epithelium into web of capillaries that
surrounds each alveolus.
° CO2 diffuses in opposite direction.

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8.1.3 Role of Partial Pressure Gradient

• Diffusion of gas depends on differences in

partial pressure, the concentration of a
particular gas to overall total.
° Atmospheric pressure at sea level = 760

mm Hg.
° Since atmosphere is 21% O2 (by volume),

partial pressure of O2 is 0.21 × 760, or

about 160 mm Hg.
° Partial pressure of CO2 is 0.23 mm Hg.

• Gas diffuses from region of higher to

region of lower partial pressure. (Figure 42.27,

Campbell)
• Blood arriving at lungs via pulmonary
arteries has lower partial pressure of O2
and higher partial pressure of CO2 than air
in alveoli.
° As blood enters alveolar capillaries, CO2

diffuses from blood to air within alveoli,

and O2 in alveolar air dissolves in fluid
that coats epithelium and diffuses across
surface into blood.
° By the time blood leaves lungs in

pulmonary veins, its partial pressure of O2

has been raised and its partial pressure
of CO2 has been lowered.

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• In tissue capillaries, gradients of partial

pressure favor diffusion of O2 out of blood
and CO2 into blood.
° Cellular respiration removes O2 from and

adds CO2 to interstitial fluid by diffusion.

° After blood unloads O2 and loads CO2, it is

returned to heart and pumped to lungs

again, where it exchanges gases with air
in alveoli.

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8.1.4 Respiratory pigments

• A diversity of respiratory pigments has

evolved in various animal taxa to support
their normal energy metabolism:
° Hemocyanin: In hemolymph of

arthropods and many molluscs, - has

copper as its O2-binding component,
coloring the blood bluish.
° Hemoglobin: In red blood cells -

respiratory pigment of almost all

vertebrates.
 Consists of four subunits, each with a

heme group (cofactor) that has an iron

atom at its center. (Figure 5.23, Campbell)
 Because iron actually binds the O2,

each hemoglobin molecule can carry

four molecules of O2.

Hb + 4O2 ↔ HbO8 (Oxyhemoglobin)

•Hemoglobin binds O2 reversibly, loading

O2 at lungs or gills and unloading it in other
parts of body.

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8.1.5 Oxygen Transport and Bohr

Effect

• Low solubility of O2 in water is a

fundamental problem for animals that rely
on circulatory systems for O2 delivery.
• Thus, most animals transport most of O2

bound to respiratory pigments instead of

dissolved in solution.

(a) Oxygen Dissociation Curve for

Hemoglobin
(Figure 42.28, Campbell)

 O2 saturation of hemoglobin (%) plotted

against different values of partial pressure
of O2 (PO2) (mmHg)
° Shows how readily hemoglobin
acquires and releases O2 molecules from
its surrounding tissue.
• Where dissociation curve has a steep

slope, even a slight change in PO2 causes

hemoglobin to load or unload a substantial
amount of O2.
° Steep part corresponds to range of partial

pressures found in body tissues.

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° A slight drop in PO2 causes relatively large

increase in amount of O2 unloaded by
blood.

• When pO2 is high (alveoli) - hemoglobin

almost fully saturated.
° Significance - hemoglobin picks up O2
as it passes through lungs.
• When the pO2 falls at first (plateau) - little
effect on % saturation.
° Significance - as blood passes
through heart and arteries, pO2 drops
slightly but hemoglobin does not lose
much O2.
• When there is a relatively small change in
pO2 (steep part of curve) - large change in
% saturation of hemoglobin.
° Significance - when blood reaches the
respiring tissues, hemoglobin gives up
most of its O2.
• At low pO2 hemoglobin is unsaturated - it
has given up most of its O2.

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(b) The Bohr Shift

• As in all proteins, hemoglobin’s

conformation is sensitive to a variety of
factors.
° For example, a drop in pH (increase in

CO2/pCO2) lowers affinity of hemoglobin

for O2, an effect called Bohr shift - curve
shifts to the right.
° Increase in temperature also shifts curve

to right.
• Because CO2 reacts with water to form

carbonic acid, an active tissue will lower

the pH of its surroundings and induce
hemoglobin to release more O2.
° Significance - hemoglobin more efficient

at releasing O2 (more oxyhemoglobin

dissociates).
° As tissues become more active, rate of

respiration increases, more CO2 is

released.
° Thus, tissues receive more O2 and can

continue aerobic respiration at same

partial pressure of O2.
• Hemoglobin more efficient at taking up O2
when CO2 levels are low i.e. in lungs.

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• Myoglobin:
° Hemoglobin-like, iron-containing pigmen
tin muscle fibres.
° Consists of single ∝-polypeptide chain
and binds only one O2 molecule.
° O2 dissociation curve is hyperbolic (as
opposed to sigmoidal curve for
hemoglobin) and is to left of that for
hemoglobin.
° Takes up O2 from hemoglobin in blood and
stores O2 within muscle itself.
° Significance - it is an O2 store.
It only releases O2 at very low partial
pressures of O2. Enables muscle to
continue respiring aerobically (to
continue contracting) at low levels of O2.

• Fetal haemoglobin:
° Dissociation curve to left of maternal
("normal") adult hemoglobin.
° Thus, has higher affinity for O2 than
maternal hemoglobin.
° Significance - can take O2 from maternal
haemoglobin at same partial pressure of
O2 in placenta.

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8.1.6 Carbon dioxide transport

• Hemoglobin also helps transport CO2 and

assists in buffering blood pH. (Figure 42.29,
Campbell)
° 7% of CO2 transported in solution.
° 23% binds to amino groups of
hemoglobin.
° 70% transported as bicarbonate ions.
• CO2 from respiring cells diffuses into

blood plasma and then into RBCs.

° CO2 first reacts with water, assisted by

enzyme carbonic anhydrase, to form

H2CO3.
+
° H2CO3 dissociates into hydrogen ion (H )

and bicarbonate ion (HCO3−)

+
° H attaches to hemoglobin (forming

hemoglobinic acid, HHb) and other

proteins, and act as buffer, minimizing
change in blood pH.
−
° HCO3 diffuses into plasma.
−
° As HCO3 diffuses out of RBCs, chloride

ions (Cl−)diffuse into RBCs to maintain

electrical neutrality – process is known
as chloride shift.
• As blood flows through lungs, process is

reversed as diffusion of CO2 out of blood

shifts chemical equilibrium in favor of
conversion of HCO3− to CO2.

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8.1.7 Control of breathing

(Figure 42.26, Campbell)

• Breathing control centers - in medulla

oblongata and pons.
° Medulla’s centers set basic breathing

rhythm – inspiratory center increases

inspiratory rate; expiratory center cuts
off inspiratory activity & promotes
expiration.
° Inspiratory center send impulse via

phrenic nerves to diaphragm and via

thoracic nerves to rib muscles,
stimulating them to contract and making
us inhale.
° Pons helps control transition from
inhalation to exhalation.
° A negative-feedback mechanism via

stretch receptors prevents our lungs

from over-expanding by discharging
inhibitory impulses via vagus nerves to
expiratory center in medulla.

• Medulla’s control center monitors CO2

level of blood and regulates breathing
activity appropriately.

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° Chemoreceptors in medulla detect

changes in pH (increase in CO2) of blood
and cerebrospinal fluid bathing brain.
 CO2 reacts with water to form carbonic

acid, which dissociates into HCO3− and

H+, which lowers pH.
° Chemoreceptors in walls of aorta &
carotid arteries also detect changes in
pH.
 Nerves impulses relay changes to

medulla.
° Medulla’s control center increases depth
and rate of breathing, and excess CO2 is
eliminated in exhaled air.

• O2 concentrations in blood usually have

little effect on breathing control centers.
° But, if O2 level falls markedly, example, at

high altitudes, chemoreceptors in aorta

and carotid arteries in neck send signals
to medulla to increase breathing rate.

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Lungs deflated

6. Expiration

Diaphragm &
Inhibitory impulse
intercostals
no longer fired
muscles relax

Step 1 begins
5. No impulse
again

CO2 from
Voluntary BREATHING 1. Stimulation
FOREBRAIN control respiratory
CENTER
activity

4. Inhibition of 2. Nerve
inspiration impulse fired

Inhibitory impulse Diaphragm &

fired from stretch intercostals
receptors in lungs muscles contract

3. Inspiration

Lungs inflated

Figure: Control of Ventilation

From: Biological Science: Green, N.P.O., Stout, G.W., &

Taylor, D.J., Cambridge University Press.

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8.1.8 Lung Volumes

• Volume of air inhaled and exhaled can be

measured using a spirometer.
Lung volume/dm3

Maximum inspiratory
level
5.0
Inspiratory
Inspiratory reserve volume Respiratory
capacity inspiratory level
3.45
Vital Tidal
capacity volume
3.0 Resting
Expiratory expiratory level
reserve volume
Functional
1.5 residual Maximum expiratory
capacity level
Residual
volume
• Terms to describe volume changes of lungs
0 during breathing:
Time

a) Tidal Volume (TV) - The average

volume of gas inspired and exhaled during
normal breathing.
b) Inspiratory Reserve Volume (IRV) -
The maximum amount of gas that can be
inspired from the inspiratory point of a
normal tidal volume.

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c) Expiratory Reserve Volume (ERV) -

The maximum volume of gas that can be
exhaled from the resting end-expiratory level.
d) Residual Volume (RV) - The volume of
gas that remains in the lungs after a
maximum exhalation.
e) Inspiratory Capacity (IC) - The
maximum volume of gas that can be inspired
from the resting end-expiratory level.
f) Vital Capacity (VC) - The maximum
amount of gas that can be exhaled after a
maximum inspiration.
g) Functional Residual Capacity (FRC) -
The volume of gas that remains in the lungs
at the end of a normal exhalation.
h) Tidal Lung Capacity (TLC) - The total
volume of gas contained in the lungs after a
maximum inspiration.

• Summary of Lung Capacities

° IC = VT + IRV
° FRC = ERV + RV
° VC = VT + IRV + ERV
° TLC = VT + IRV + ERV + RV

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• Normal values for lung volumes &

capacities (For normal adult male)

Volumes Capacities (ml)

1. IRV 3100
2. TV 500
3. ERV 1200
4. RV 1200
5. IC 3600
6. VC 4800
7. FRC 2400
8. TLC 6000

http://www.mededsys.com/courses_online/208/208.html#lungvol

• Lungs hold more air than the vital capacity

- some air, the residual volume, remains
in lungs because alveoli do not completely
collapse.
• Since lungs do not completely empty and

refill with each breath cycle, newly inhaled

air is mixed with O2-depleted residual air.
° Thus, maximum O2 concentration in

alveoli is considerably less than in

atmosphere.
° Although this limits effectiveness of gas

exchange, CO2 in residual air is critical

for regulating pH of blood and breathing
rate in mammals.

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Respiratory diseases

Disease Respiratory failure

Asthma Constriction of smooth muscles in
the bronchiolar and bronchial wall,
excess mucus secretion and
insufficient recoil of the alveoli.
Caused by allergy and emotional
upset. Results in difficulty in
breathing.
Pneumonia Alveoli filled of fluid, caused by
chemical, bacteria (Streptococcus),
viruses, protozoa or fungi.
Tuberculosi Mycobacterium tuberculosis,
s water-borne bacteria causes lung
damage in variety of ways. The
infectious bacteria are normally
spread through air by coughing and
sneezing.

Lung cancer Any inhaled irritant stimulates cell

( pulmonary to grow abnormally. Individual has
carcinoma) difficulty to breathe; chest pains
and spitting blood. Cigarette
smokers have 20 times more risk
than non smokers of having this
disease.

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8.2 Gaseous Exchange in Plants

8.2.1 Structure of stoma

(Figure 36.13, Campbell)

• Stoma – physical gap between two

specialized epidermal cells, guard cells.
• Guard cells are suspended by other

epidermal cells over an air chamber,

leading to internal air space.
• Guard cells control diameter of stoma by
changing shape, thereby widening or
narrowing gap between the two cells.
° When guard cells take in water by

osmosis, they become more turgid, and

guard cells become bowed due to
orientation of cellulose microfibrils.
 This increases gap between cells.
° When guard cells lose water and become
flaccid, they become less bowed, and
space between them closes.

• Why guard cells bow apart when turgid:

(i) They are fused at their ends.
(ii) Inner cell walls which form stoma are
thicker than outer walls.
(iii) Cellulose microfibrils are oriented
radially rather than longitudinally.

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8.2.2 Mechanism of stomatal opening

and closing
(Figure 36.13, Campbell)

• Changes in water potential (Ψ w) that open

and close stomata result from reversible
uptake and loss of K+ by guard cells.
° Stomata open when guard cells actively

accumulate K+ into vacuole.

° Water potential in guard cells decreases,

leading to inflow of water by osmosis.

+
° Stomata close due to exodus of K from

guard cells, leading to osmotic loss of

water.
° Regulation of aquaporins may also be
involved in swelling and shrinking of
guard cells by varying permeability of
membranes to water.
+
• K fluxes across guard cell membranes

are coupled to generation of membrane

potentials by proton pumps.
° Stomatal opening correlates with active

transport of H+ out of guard cells.

° The resulting voltage (membrane

potential) drives K+ into cell through

specific membrane channels.

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(a) Stomatal opening

• Membrane proton pump is activated -

pumps H+ out of cell:
° Generates a stronger membrane potential

(gets more negative, originally -100 mV, it

goes to -150 or -180 mV), i.e., membrane
is hyperpolarized.
• Triggers inward-specific K+ channels to
open.
° K diffuses in down its electrochemical
+

gradient.
° K concentration can increase from ≈ 100
+

mM to 400 - 800 mM
• Cl- also diffuses in to balance positive
charge of K+.
° Guard cells also make malate to
2-

balance the K+ and lower the pH.

• Accumulation of ions makes water
potential (Ψ w) of guard cells more negative.
° Water enters cells, moving down water

potential gradient, causing guard cells to

swell.
• Stomata open due to changes in volume of
guard cells.
° Radial arrangement of cellulose

microfibrils in guard cell walls forces

pore to open when guard cells swell.

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(b) Stomatal closure

• Closure initiated by:

° Shutting down proton pump.

° Opening anion (like Cl ) channels,

-

allowing anions to flow out.

° This dissipates much of the membrane

potential (charge difference becomes

less negative, going back to original
membrane potential of ≈ -100 mV)
• Inward-specific K+ channels close,
outward-specific K+ channels open.
° K diffuses out of cell, again down its
+

electrochemical potential.
° Water potential in guard cells becomes

less negative.
° Water flows out of cells & they shrink.

• Reduced volume of guard cells causes

stomatal pore to collapse shut.

http://www.esf.edu/efb/course/EFB530/lectures/stomata.htm

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•Factors (cues) that trigger stomatal

opening:

(i) Blue-light receptors in guard cells

stimulate activity of ATP-powered proton
pumps in plasma membrane, promoting
uptake of K+.
(ii) Depletion of CO2 within air spaces of leaf
as photosynthesis begins.
(iii) Circadian rhythm - internal “clock”
located in guard cells that regulate
cyclic processes.

• Factors that trigger stomatal closing:

(i) Darkness.
(ii) High internal CO2 concentration.
(iii) Abscisic acid - produced by mesophyll
cells in response to water deficiency/
stress.
(iv) Circadian rhythm.

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