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© Springer Science+Business Media Singapore 2016
U. Kulshrestha, P. Saxena (eds.), Plant Responses to Air Pollution,
DOI 10.1007/978-981-10-1201-3_8
Effect of Air Pollutants on Plant
Gaseous Exchange Process: Effect
on Stomata and Respiration
Anshu Gupta
Abstract
Air pollution has become an extremely serious problem. Air pollutants
affect both plants and animals. Under polluted conditions, plants develop
different physiological, morphological and anatomical changes. Pollutants
cause damage to cuticular waxes by which then they enter the leaves
through stomata. This further leads to injury to plants which can be either
acute or chronic. Changes in stomata due to air pollutants which seem to
be small can be of great consequence with respect to survival of the plant
during stress. These effects can further lead to disturbing the water balance
of leaf or whole plant. Respiration also gets affected because of the expo-
sure of plants to air pollutants. The present paper deals with the effect of
air pollutants on stomata as well as on respiration leading to affect gaseous
exchange.
Keywords
Air pollutants • Stomata • Stress • Respiration
8.1 Introduction
Air pollution has become an extremely serious
problem for the modern industrialised world. The
prime concern for today’s world is changes in the
gaseous composition of earth’s atmosphere.
Fossil fuel consumption has accelerated due to
increase in human population, industrial revolu-
tion, technological advancement and urbanisa-
tion (Watson et al. 1990 ). The atmospheric
concentration of CO
2 has increased from about
275 ppm prior to industrial revolution to a present
value of 365 ppm, and it is increasing at the rate
of 1–1.5 ppm/year (Conway et al. 1994 ). Its con-
centration is expected to be doubled by the mid-
dle of the next century (IPCC 1990 ).
Uncontrolled use of fossil fuels in industries
and transport sectors has led to the increase in
concentrations of gaseous pollutants such as SO
2 ,
NO
x , etc. (Rai et al. 2011 ). The general state of
the environment, including air quality, is
A. Gupta (*)
School of Environmental Sciences (SES) , Jawaharlal
Nehru University (JNU) , New Delhi 110067 , India
e-mail: anshu.guptaevs@gmail.com
8
pallavienvironment@gmail.com
86
deteriorating in many cities of the developing
countries. World Bank studies in selected cities
of developing countries have shown that swelling
urban populations and the growth of industrial
activities and automotive traffi c in Asia have
caused serious air pollution (World Bank 2009 ).
The adverse effects of air pollution have been
associated with three major sources: sulphur
dioxide and solid particulates from fossil fuels;
photochemical oxidants and carbon monoxide
from motor vehicles and miscellaneous pollutants
such as hydrogen sulphide, lead and cadmium
emitted by smelters, refi neries, manufacturing
plants and vehicles (Birley and Lock 1999 ).
It is a known fact that 60 % of air pollution in
city is caused by automobiles only. On sensitive
species of both plants and animals, the effect of
these pollutants is observed at acute level. Plants
are considered for investigation of effect of auto
exhaust pollutants. Response of plants towards
air is being assessed by the air pollution tolerance
index (APTI). Some plant species and varieties
are so sensitive that they can be conveniently
employed as biological indicators or monitors of
specifi c pollutants. They can further assist the
planner in managing the urban cities
(Horaginamani and Ravichandran 2010 ).
Agarwal and Bhatnagar ( 1991 ) studied APTI of
some selected plants and described Mangifera
indica as reliable bioaccumulator plant. Air pol-
lution affects plants mainly through the uptake of
pollutants through stomata. Sulphur dioxide and
ozone are the two most important pollutants that
affect the plants (Emberson 2004 ). SO 2 is a wide-
spread phytotoxic air pollutant in the environ-
ment with ambient concentration of about 0.001
ppm in the air (Allen 1990 ).
8.2 Plant Responses to Air
Pollutants
Air pollution may or will have harmful effects on
living things and materials. It may interfere with
biochemical and physiological processes of
plants to an extent, which ultimately leads to
yield losses (Heck et al. 1988 ). Studies have
shown that under polluted conditions, plants
develop different morphological, physiological
and anatomical changes (Inamdar and Chaudhari
1984 ; Iqbal 1985 ; Gravano et al. 2003 ; Dineva
2004 ). Sulphur dioxide, one of the most promi-
nent phytotoxic by-products of fossil fuel burn-
ing, is also rising progressively in large areas
around the world, especially in developing coun-
tries. Both elevated CO
2 and SO
2 are anthropo-
genic stress factors and have potential infl uence
on biological systems including agricultural
crops (Aggarwal and Deepak 2003 ).
Sulphur dioxide is a widespread toxic air pol-
lutant which can cause positive effects on physi-
ological and growth characteristics of plants at
low concentrations, especially in plants growing
in sulphur- defi cient soil (Darrall 1989 ) when the
sulphate might be metabolised to fulfi l the
demand for sulphur as a nutrient (De Kok 1990 ).
Increased uptake of SO
2 can cause toxicity
and reduce growth and productivity of plants due
to accumulation of sulphite or sulphate, by inter-
acting with different physiological processes, and
also it damages tissues and pigments (Darrall
1989 ; Agrawal and Verma 1997 ). In certain cases,
SO
2 -induced reduction in plant growth and alter-
ation of physiological and biochemical processes
are not accompanied with visible foliar symp-
toms (Crittendem and Read 1978 ). Reduction in
yield is also reported without visible symptoms
when plants are treated with low concentration of
SO
2 for long duration (Godzik and Krupa 1982 ).
Sulphur is necessary for the general metabo-
lism of plants because it is a major component of
amino acids, proteins and some vitamins. In
healthy leaves, sulphur content ranges from 500
to 14,000 ppm by dry weight (0.5–14 mg/g dry
weight) depending upon species. Concentrations
below 250 ppm are considered critical, giving
rise to defi ciency symptoms and to the substitu-
tion of selenium (when available) for Sulphur in
amino acids and proteins (Treshow 1970 ). Part,
or all, of the sulphur requirements of plants may
be met by direct uptake of SO
2 from the atmo-
sphere if it is present at very low concentrations.
On the other hand, if the concentration of SO
2
increases beyond a certain critical level that may
vary with species (biochemical threshold level),
it can result in the general disruption of photo-
A. Gupta
pallavienvironment@gmail.com
87
synthesis, respiration and other fundamental
cellular processes. Injury becomes irreversible,
leading to death, as concentration and time of
exposure increase further. Tolerance varies with
many factors of the plant and of its environment
(Malhotra and Hocking 1976 ).
8.3 Entry and Effects
of Pollutants on Plants
The following are the effects and route of the pol-
lutants entering the plant leaf through stomata,
affecting respiration and other gas exchange
processes.
8.3.1 Uptake of Pollutants
The most susceptible part of a plant to injury is
the leaf due to the presence of abundant stomata
which permit the penetration of pollutants into
the tissues of the leaves. Boundary layer resis-
tance is the fi rst barrier of gaseous air pollutants
which varies with a number of factor including
wind speed, size, shape and orientation of leaves
(Heath et al. 2009 ). More pollutants enter the
leaves at higher wind speed as boundary layer
resistance declines. Waxy cuticle is a potential
barrier to most of the pollutants but the cells most
exposed to air pollution action are epidermal
cells. However, cuticular waxes can be dissoci-
ated by acidic gases and these gases can enter the
leaves by penetrating the cuticle (Rai et al. 2011 ).
8.3.2 Effect on Cuticle and Stomata
Cuticle and stomata are the fi rst receptors or tar-
gets where the pollutants encounter. Stomata pro-
vide the direct path through which the gases enter
the leaf, but the direct impact on cuticle must also
be considered. The response of stomata to air pol-
lutants is varying and varies from species to spe-
cies. It also varies with concentration, age of the
plants as well as environmental conditions
(Abeyrante and Illeperuma 2006 ). Plant species
differ in their ability to mitigate traffi c pollution
due to differences in their leaf surface character-
istics which include epicuticular wax, cuticle,
epidermis, stomata and trichomes (Neinhuis and
Barthlatt 1998 ).
Pollutants absorbed by guard cells and subsid-
iary cells may initially affect the stomatal aper-
ture. Sulphur dioxide has a notable effect in
stimulating stomatal opening (Mansfi eld and
Majernick 1970 ), interacting with CO
2 and atmo-
spheric moisture.
Different plant species can respond differently
when exposed to same concentrations of SO
2
(Biggs and Davis 1980 ). It can cause opening of
stomata in one species and closing in another
(Mudd 1975 ). It has also been reported that short-
term exposure to SO
2 causes stomatal opening,
whereas long-term exposure can lead to partial
closure (Abeyrante and Illeperuma 2006 ). The
effects of SO
2 and acid deposition are well seen
on cuticular waxes and are well documented
(Fowler et al. 1980 ). Degradation of cuticular
waxes due to air pollution has been seen in spe-
cies such as Scots Pine. Due to air pollution and
acid deposition, the weathering of needle cuticle
is many times faster in unpolluted forest areas.
Similar observations have been described in
lichens and mosses (Huttunen and Lane 1983 ).
Due to this evapotranspiration would be greater
which would be critical in arid environments.
SO
2 had been found to show decrease in photo-
synthesis and respiration in cultured lichen sym-
bionts (Showman and Rudolph 1971 ). Air
pollutants and oxidative stresses can also have a
marked effect on the Ca
2+ homeostasis of guard
cells and the intracellular machinery responsible
for stomatal movement (McAnish et al. 2002 ).
Pollutants like SO
2 enter the leaves mainly
through the stomata, resultant injury is classifi ed
as either acute or chronic. Abeyrante and
Illeperuma ( 2006 ) have given a plot showing the
average values calculated for stomatal pore width
versus SO
2 concentration at the three sampling
sites (Fig. 8.1 ). Sampling site 1 recorded high
SO
2 conc. as compared to other two sites. Site 1
had 50 % of the pore size of the stomatal of leaves
as compared to other two sites.
8 Effect of Air Pollutants on Plant Gaseous Exchange Process: Effect on Stomata and Respiration
pallavienvironment@gmail.com
88
Acute injury results in the appearance of
symptoms like two-sided (bifacial) lesions that
usually occur between veins and along the
margins of the leaves occasionally. Rai and
Kulshrestha ( 2006 ) have suggested that due to air
pollutants the inhibited cell elongation, leaf area
and consequently the increase in cell frequency
resulted in reduction in the size of stomata and
epidermal cells. In order to avoid entry of harm-
ful constituents of exhaust which can otherwise
cause adverse effects, the reduction in the size of
stomata could be considered as an adaptive
response (Satyanarayana et al. 1990 ; Salgare and
Thorat 1990 ).
Distorted shapes of stomata observed in
Pongamia pinnata populations exposed to
exhaust pollution might have resulted due to low-
ering of pH in cytoplasm of guard cells and thus
change in the turgor relations of the stomata com-
plex (Kondo et al. 1980 ) due to physiological
injury within the leaf (Ashenden and Mansfi eld
1978 ). Further, Rai and Mishra ( 2013 ) have illus-
trated that the plants growing along the roadsides
have modifi ed leaf surface characters including
stomata and epidermal cells due to the stress of
automobile exhaust emission with high traffi c
density in urban areas.
Rahul and Jain ( 2014 ) have reported that dust
particles of a range less than 5 mm in diameter
can interfere with the mechanism of stomatal
pores. These small openings are largely respon-
sible for the basic respiration and transpiration
function of plants.
Most of the air pollutants which are known to
show effect on stomata, are natural components of
the atmosphere, but they are present now in higher
concentrations in the atmosphere than their natu-
ral concentration. The changes in the stomata due
to air pollutants which seem to be small can be of
great consequence with respect to survival of a
plant during stress (Robinson et al. 1998 ).
Stomatal resistance should be considered as
the main obstacle to Ozone fl ux (Kollist et al.
2000 ), the direct reaction of the pollutant with
cell wall ascorbate is frequently involved (Plochl
et al. 2000 ). The fi rst detoxifying layer which
represents the antioxidant system found in the
cell (apoplasm + symplasm) at the time of Ozone
attack will scavenge ozone and its derivatives
(Fig. 8.2 ). This system is highly linked to the
level of ascorbate and especially apoplastic
ascorbate, which was primarily proposed as a
good indicator for ozone tolerance (Turcsanyi
et al. 2000 ; Tausz et al. 2007 ).
8.3.3 Effect on Plant Water Balance
Many atmospheric pollutants interfere with the
control of stomatal aperture even when present at
low concentrations. Therefore they have potential
to upset the water balance of the leaf or the whole
plant. Pollutants such as SO
2 and CO
2 cause sto-
matal closure at higher concentrations, whereas
at low concentrations the stomatal conductance is
often increased (Robinson et al. 1998 ).
Fig. 8.1 Correlation between
average SO2 concentration and
pore width (Source: Abeyrante
and Illeperuma
2006 )
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89
8.3.4 Effect on Respiration
Exposure to air pollutants may result not only in
damage of leaf, reduction in growth and yield of
crops but it also interferes with physiological
processes (Unsworth and Ormrod 1982 ).
Exposure of plants to air pollutants at high con-
centration for a long period of time results in the
development of symptoms of visible injury and
associated physiological disturbances. These
responses are generally irreversible and may lead
to reduction in plant growth and yield. Many fac-
tors including plants, pollutants and environment
will affect the sensitivity of plants to a range of
pollutants. These include toxicity of the pollut-
ant, concentration, frequency and duration of
exposure to pollutant, stomatal behaviour, pollut-
ant uptake by plants and prevailing environmen-
tal conditions like sunlight, humidity and
temperature.
Although the process of respiration includes
dark respiration and photorespiration which are
important components of carbon budget, the evi-
dences for pollutant-induced modifi cation of
respiration are less well documented than for
photosynthesis. Since these processes of respira-
tion occur in several sites in the cell, including
mitochondria, peroxisomes, cytoplasm, these
processes are very much vulnerable to pollutant
attack (Koziol and Whatley 2013 ).
In a study by Aggarwal and Deepak ( 2003 ),
investigating the long-term infl uence of elevated
concentration of CO
2 and SO
2 , singly and in com-
bination on the physiological and biochemical
characters of two cultivars of wheat ( Triticum
aestivum ), showed that the respiration rate, total
phenolics and total soluble sugars increased in
response to SO
2 . Dark respiration (Rs) increased
in response to SO
2 - and CO
2 + SO
2 -treated plants
as compared to control. In contrast, elevated CO
2
caused decline in Rs insignifi cantly. Rs increased
at individual treatment of SO
2 because the series
of reactions leading to detoxifi cation of SO
2 are
ATP mediated which is provided by respiration.
In contrast to this there was an insignifi cant
decline in Rs due to CO
2 enrichment (Aggarwal
and Deepak 2003 ). Respiration rates were found to
be function of both leaf nitrogen and carbohydrate
Fig. 8.2 Summary of
the relationships
between stomatal
uptake, metabolic
changes and
detoxifi cation system
under chronic ozone
attack in plant cells.
ASC ascorbate, PEPcase
phosphoenolpyruvate
carboxylase, ROS
reactive oxygen species,
Rubisco ribulose-1,5-
bisphosphate
carboxylase (Source:
Dizengremel et al.
2008 )
8 Effect of Air Pollutants on Plant Gaseous Exchange Process: Effect on Stomata and Respiration
pallavienvironment@gmail.com
90
concentration (Tjoelker et al. 1999 ). Aggarwal
and Deepak ( 2003 ) have also reported that rate of
respiration was affected by declining leaf nitrogen
and increasing TNC in response to CO
2 .
A number of studies have been done on the
effect of SO
2 on respiration and oxidative
phosphorylation. In contrast to the above study,
sulphur dioxide has been reported to reduce
respiration in plants (Gilbert 1968 ). Ballantyne
( 1973 ) showed sodium sulphite inhibited ATP
formation in both bean and corn mitochondria.
This inhibition due to sulphite was partially
reversed by the addition of oxidised glutathione
to the reaction mixture following addition of
mitochondria.
Although most of the workers have investi-
gated photosynthetic response, effects on respira-
tory processes have also been observed.
8.3.4.1 Respiratory Response to High
Concentration of Pollutants
When plants get exposed to high concentration of
pollutants, plants develop visible injury to the tis-
sues. Depending upon the degree of injury, the
respiration is either inhibited or stimulated. As
the repair processes utilise energy because of
this, the rate of respiration in the non-damaged
tissues adjacent to these necrotic areas is
increased. A wasteful loss of carbohydrate and
energy which is normally used in growth occurs
due to the enhancement of respiration in response
to high concentration of pollutants. If exposures
are not extreme, physiological processes are
altered but no visible damage occurs. If stress
periods are prolonged, these effects may lead to
reductions in growth in the long run (Koziol and
Whatley 2013 ). Reduced respiration in plants
grown at elevated CO
2 has common response but
not universal (Ziska and Bunce 1993 ). Carbon
dioxide serves as the substrate for photosynthe-
sis. Results of several experiments at elevated
CO
2 have indicated stimulation of photosynthesis
and reduction in photorespiration, thereby
increasing the growth and productivity of plants
(Allen 1990 ). Under CO
2 enrichment, the amount
of carbon fi xed is greater than the amount of car-
bon lost and therefore growth and productivity
are enhanced (Ryan 1991 ).
8.3.4.2 Respiratory Response to Low
Concentration of Pollutants
On exposure to a low concentration of pollutants,
stimulation of respiration is usually exhibited by
plants, which may be due to the operation of
detoxifi cation and repair mechanism. In response
to pollutants, shift from the glycolytic pathway to
the pentose phosphate pathway is often observed.
If the pollutant exposure periods are short, this
enhanced use of energy by the plant is likely to be
of benefi t. So it prevents the pollutants to reach
the sensitive metabolic sites like photosynthetic
pathways within the cell. Prior to any observed
depression of photosynthesis, indeed respiration
can be affected (Koziol and Whatley 2013 ).
8.3.4.3 Effect of Pollutants
on Photorespiration
Evidences generally are not available to allow an
assessment of effect of pollutants on photorespi-
ration. This is due to some reasons like diffi cul-
ties involved in measuring rates of respiration in
the light and also due to the fact that early inves-
tigations were unaware of the existence of this
process. Indeed, photorespiration is a wasteful
process; pollutant-induced effects may be benefi -
cial to the growth of the plants because in the
short term, the rates of net photosynthesis will
increase (Koziol and Whatley 2013 ).
8.3.4.4 Changes in Respiration
in Association
with Photosynthesis
If the rate of photosynthesis in plants is very
high, a small change in the rate of respiration will
not effect signifi cantly on the carbon balance of
the plant. On the other hand, if the rate of photo-
synthesis is very low, change in respiration can
lead to change in growth and yield of the plant
(Koziol and Whatley 2013 ).
If the environmental conditions like light and
temperature are limiting for the plant photosyn-
thesis and plant is exposed to very high
concentration of pollutants, under these condi-
tions, photosynthesis will be severely reduced.
Under such conditions change in respiration rate
could alter signifi cantly the carbon balance of the
plant. This may lead to premature leaf drop,
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91
senescence. Jones and Mansfi eld ( 1982 ) have
reported that greater reduction in photosynthetic
rates has been seen in plants exposed to higher
level of pollutants under light-limited conditions
than the plants under light higher irradiance.
Evidences are there to show the response to
pollutants from the non-photosynthetic portion of
the plant such as roots. A reduction in the activity
of root will have consequences not only upon
root growth but also for the whole plant, if the
plant is growing in a stressful environment.
8.4 Conclusion
The study shows that leaf characters including
cuticle, stomata, epidermal cells, and guard cells
get affected due to stress induced by the air pol-
lutants. This further affects the gaseous exchange
as well as respiration in plants. This is an indica-
tor of environmental stress. The effects of indi-
vidual pollutants are quite variable because they
vary from species to species. Changes in leaf in
characters induced due to the effect of air pollut-
ants seem to be small, but during the survival of
the plant in stress, they can be of great
consequence.
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