65

Effect of Light Intensity on Photosynthesis

Rinukshi Wimalasekera
Department of Botany, Faculty of Applied Sciences, University of Sri Jayewardenepura, Gangodawila, Sri Lanka

4.1 ­Introduction
In plants, the conversion of light energy into chemical energy happens in the process of
photosynthesis. In photosynthesis, higher plants use solar energy to reduce carbon
dioxide, producing carbohydrates, and to oxidize water, releasing oxygen. Light is
essential for three important phases of photosynthesis, namely, light harvested from
sunlight, reduces nicotinamide adenine dinucleotide phosphate (NADP) and produces
adenosine triphosphate (ATP), and converts CO2 into carbohydrates (Mirkovic et al.
2017). The protein complexes photosystem I (PSI) and photosystem II (PSII) in higher
plants are composed of the pigments necessary to harvest photons from specific wave-
lengths of light to catalyze photosynthetic reactions. Light reactions where energy
transduction happens and dark reactions where carbon fixation happens are the two
main types of reactions.
Photosynthesis of higher plants and green algae are regulated by light‐harvesting
functions in response to different light qualities (Ueno et al. 2018). Photosynthetically
active radiation (PAR) ranging from 400 to 700 nm represents the critical energy for
photosynthesis in plants. Although PSI and PSII use the same pigments, their absorp-
tion spectra differ. PS1 preferentially absorbs wavelengths longer than 680 nm in the
far‐red light region and PSII strongly absorbs 475–680 nm with an affinity for red light
of 680 nm (Taiz and Zeiger 2002; Blankenship 2014).
Light intensity or light quantity is the total amount of light that plants receive and it
is an important factor determining the rate of photosynthesis (Chapman and Carter
1976; Taiz and Zeiger 2002). At low light intensities, above the light compensation point
(LCP), photosynthetic rate increases proportionally to the light intensity and reaches a
maximum. However, as light intensity is increased further, chlorophyll can get damaged
and as a result the rate of photosynthesis decreases.
Plant productivity is strongly dependent on the photosynthetic rate. At a given time,
photosynthetic rate is determined by the limiting factors of light or CO2 concentration.
Rubisco and ribulose 1,5‐bisphosphate (RuBp) activities are major metabolic steps that
are important for optimal photosynthetic performance. At high light conditions rubisco

Photosynthesis, Productivity, and Environmental Stress, First Edition.

Edited by Parvaiz Ahmad, Mohammad Abass Ahanger, Mohammed Nasser Alyemeni, and Pravej Alam
© 2020 John Wiley & Sons Ltd. Published 2020 by John Wiley & Sons Ltd.
66 4 Effect of Light Intensity on Photosynthesis

activity is limited and at low light conditions RuBp regeneration is limited. Maintaining
photosynthetic efficiency under changing light conditions requires modification of
light‐harvesting and energy‐transfer processes.
Leaves are capable of adapting to high or low light conditions depending on the
growth environment of plants. Anatomical features of leaves and the arrangement of
chloroplasts regulate the quantity of light absorption thereby preventing damage to the
photosynthetic system by absorption of excessive light. In a similar way, leaf anatomical
and biochemical features help in optimizing light capture when exposed to shade condi-
tions. Plasticity in morphology and biochemical responses are important in maximizing
photosynthetic efficiency (Taiz and Zeiger 2002; Valladares et al. 2005).
Several signaling pathways and underlying regulatory genes responsible for photo-
synthetic light reactions have been identified in many plant species.

4.2 ­Characteristics of Light
Light has both particle and wave characteristics. A light particle is known as a photon
and the energy it contains is called a quantum. Light intensity or light quantity is the
rate at which light spreads over a given surface area. Light intensity is also referred to as
the energy transferred per unit area (Taiz and Zeiger 2002; Blankenship 2014).
Essential parameters in measuring light are spectral quality, quantity, and direction
(Taiz and Zeiger 2002; Both et al. 2015). The quantity of energy that falls on a sensory
plane such as a flat leaf of a known area per unit time is quantified as irradiance in
Wm−2. Photon irradiance in mol m−2 s−1 is the light that is measured as incident quanta.
The direction of light is also an important factor in photosynthesis. Light strikes directly
on a flat surface from above or obliquely (Schirillo 2013). The omnidirectional meas-
urement is called the fluence rate (Rupert and Latarjet 1978). Light interception to
shoots, whole plants, and chloroplasts is mostly not direct. Light can also come from
many directions simultaneously (Schirillo 2013).

4.2.1 Photosynthetically Active Radiation (PAR)

The total radiant energy from the sun is called the total solar irradiance. Only about 5%
of energy reaching the earth can be converted into carbohydrates by photosynthesis
because the wavelengths of the major fractions of the incident light cannot be absorbed
by the photosynthetic pigments. The radiant energy of the sun is comprised of a range
of wavelengths. PAR represents the fraction of sunlight from 400 to 700 nm and is the
crucial source of energy for plants (Taiz and Zeiger 2002; Blankenship 2014). Depending
on the time of the day and latitude PAR fluctuates and also seasonal variations of PAR
can be observed. The availability of PAR for photosynthesis is affected by cloud cover,
shading by trees, buildings, and air pollution (Rahman et al. 2017). PAR values span
from 0 to 3000 μmol m−2 s−1 and usually PAR irradiance and PAR fluence rates are about
2000 μmol m−2 s−1.
Net primary productivity or biomass production shows a linear relationship with
intercepted PAR. Radiation use efficiency (RUE) is dependent on the low availability
of water, nutrient deficiency, or any adverse stress conditions that negatively affect
metabolic activities (Hammer and Wright 1994; Ruimy et al. 1995). PAR intensity
 4.3 Light Absorption and Pigments 67

requirements for different species at given growth condition differ. Ubiernai et al.
(2013) reported that Zea mays, Miscanthus x giganteus and Flaveria bidentis show
varied C4 photosynthesis under a series of different PARs. Too high or too low PAR
intensity causes photoinhibition and disrupts the photosynthetic machinery in
Thalassia testudinum seedlings in response to changes in light (Howarth and Durako
2013). PAR changes constantly within a day. Plants have developed mechanisms to
maintain a balance between converting radiation energy thereby protecting the pho-
tosynthetic apparatus from photoinhibition and repairing possible damage (Bertamini
and Nedunchezhian 2003).

4.3 ­Light Absorption and Pigments

Chlorophylls are the main light absorbing pigments present in the chloroplasts of pho-
tosynthesizing plants. The thylakoid reactions of photosynthesis begin with the absorp-
tion of light, electron transport reactions then occur that lead to the reduction of
NADP+ to NADPH and production of ATP takes place in the internal membranes of
thylakoids in chloroplasts and carbon fixation takes place in the stroma (Lodish et al.
2000; Taiz and Zeiger 2002). Chlorophylls a and b are the most abundant photosyn-
thetic pigments that absorb light and drive the power for photosynthesis. Carotenoids
which are integral constituents of the thylakoid membranes act as accessory pigments
to absorb light and transfer to chlorophylls for photosynthesis. Pigments act as com-
plexes of antenna that absorb light and transfer energy to the reaction center complex.
Light harvesting complexes (LHCs) contain a high concentration of chlorophyll a and
depending on the species, chlorophyll b and other pigments (Nicholls and Ferguson
2013). In higher plants PSI and PSII consist of pigments necessary to harvest photons
from specific wavelengths of light. A decrease in PSII results in a substantial decrease in
photosynthesis (Demmig‐Adams and Adams 1992). If excess light is not dissipated,
over reduction of reaction centers takes place by damaging the PSII. Studies with
mutants disrupted in regulatory functions of thylakoid reactions of photosynthesis pro-
vide important information on the regulatory genes and signaling pathways.

4.3.1 Dissipation of Excess Light Energy

Plants have developed some photoprotective mechanisms to dissipate excess light and
thereby to avoid photodamage to PSII (Demmig‐Adams and Adams 1992; Qiu et al.
2003). Quenching of chlorophyll fluorescence, which transfers excessively absorbed
light energy away from electron transport toward heat production, is one such mecha-
nism of dissipating surplus energy. When exposed to excess light, leaves dissipate the
surplus absorbed light energy in order to protect the photosynthetic apparatus from
damage. Dissipated light energy is based on irradiance, species, growth conditions,
nutrition conditions, and temperature (Qiu et al. 2003; Zhou et al. 2007).
It has been shown that in rice, high light stress limits the metabolic processes of photo-
synthesis (Zhou et al. 2007). Photoinactivation of PSII complexes and photoprotection
was observed in Capsicum annuum L. leaves when exposed to 500 μmol photons m−2 s−1
(high light) (Lee et al. 2001). The xanthophyll cycle plays an important role in protecting
plants from excess light energy (Lodish et al. 2000; Taiz and Zeiger 2002). High light
68 4 Effect of Light Intensity on Photosynthesis

intensity receiving leaves contain higher levels of xanthophyll than shade leaves. The xan-
thophyll cycle is also active in low light receiving plants in the forest understory which
usually experience sun flecks. Xanthophylls are useful in photoprotection of PSII (Jahns
and Holzwarth 2012). Regulation of the xanthophyll cycle is also helpful in protecting
against damage caused by high leaf temperatures that often results from high light levels.

4.3.2 Photoinhibition
Photoinhibition is the light‐induced decrease of photosynthetic activity that occurs
when leaves receive more light than they can utilize. When exposed to high light levels,
the reaction center of PSII is inactivated and damaged (Keren and Krieger‐Liszkay
2011). Dynamic photoinhibition can be observed under moderately excess light espe-
cially during midday when leaves are exposed to maximum light. Photoinhibition is
triggered by the diversion of absorbed energy toward heat dissipation. Photoinhibition
is reversed when photon flux levels reduce below the saturation levels. Prolonged pho-
toinhibition results from exposure to excess light that damages the photosynthetic sys-
tem and lowers both quantum efficiency and photosynthetic rate (Aro et al. 1993). The
reaction center of PSII gets severely damaged and the damage persists for longer peri-
ods of time, severely affecting photosynthesis. Photoinhibition could be observed in
peas acclimated to a series of high 11 irradiance levels (Aro et al. 1993).
A short‐term decrease in quantum efficiency can be considered as a protective meas-
ure for the photosynthetic apparatus. Several studies show protective mechanisms in
response to high light intensities. It has been found that excessive light energy absorbed
by lichen soil crusts is dissipated by increasing non‐photochemical quenching, provid-
ing some protection for lichen soil crusts (Wu et al. 2017).

4.4 ­Light Absorption by Leaves

Leaves are specialized organs for light utilization and carbon fixation. Photosynthesis
greatly depends upon the absorption of light by pigments, especially chlorophyll a in the
plant leaves. Chlorophyll absorbs blue and red regions of the spectrum and about 85–90%
of PAR is absorbed by the leaf surface and the remainder is reflected or transmitted (Taiz
and Zeiger 2002; Blankenship 2014). Because of high light absorption by chlorophyll in
the upper region of the leaf, a light gradient is created from the leaf surface into the tissue
inducing different light regimes. Under strong light intensities, the chloroplasts at the
surface of the leaf are saturated and most of the light cannot be used for photosynthesis,
but the chloroplasts in the lower surface of the leaf might be light deprived. Quantification
of light absorption by isolated chloroplasts and leaves show considerable differences due
to light scattering and “packing” effects (Merzlyak et al. 2009).

4.4.1 Light Absorption and the Anatomy, Morphology, and Biochemical

Characteristics of Leaves
Leaves show highly specialized anatomical features for light absorption. The impor-
tance of leaf anatomy for photosynthesis is reflected by the differential anatomical
­features exhibited by leaves that develop under different light conditions (Vogelmann
et al. 1996; Xiao et al. 2016).
 4.4 Light Absorption by Leaves 69

The epidermal cells are transparent to light and can focus the light to chloroplasts
because of the convex nature of epidermal cells. The amount of light that is finally
received by the chloroplasts can be several times higher than the amount of ambient
light (Vogelmann et al. 1996). The distribution pattern of chlorophylls creating gaps in
the chloroplasts leads to differential absorption of light and this phenomenon is known
as the sieve effect. Light penetrates the palisade cells because of the sieve effect and light
channeling. A fraction of incident light is propagated through the central vacuole of the
palisade and air spaces between the cells. Transmission of light into the leaf interior
occurs by facilitating light channeling (Atrashevskii et al. 1999). The multiple reflec-
tions of light that occur between cell–air interfaces greatly enhance the length of the
path of photon travel. This phenomenon of light scattering is important in increasing
the probability of light absorption and thereby enhancing photosynthesis. Light scatter-
ing occurs to a greater extent when light penetrates through the air space surrounding
spongy mesophyll cells (Vogelmann et al. 1996; Pierantoni et al. 2017). In a study with
24 plant species, chloroplast movement is reported to be influenced by anatomy and
optical properties of leaves (Davis et al. 2011). There are some anatomical differences in
shade exposed and sun exposed plants. Sun grown leaves show characteristic features
like specialization of upper surface of leaves e.g. longer palisade cells (Davis et al. 2011).

4.4.2 Light‐Mediated Leaf Movement

Leaves are capable of regulating the incident light they absorb by chloroplast and leaf
movement. Under low light conditions, photosynthetic rate is increased by gathering
chloroplasts at the cell surface parallel to the plane of the leaf to maximize the absorp-
tion of light. Under high light, excess absorption of light is avoided by moving chloro-
plasts to the cell surface parallel to incident light (Taiz and Zeiger 2002; Davis et al. 2011).
Leaves are able to move diurnally and absorb light, orienting leaf blades (lamina) per-
pendicular or parallel to the sun rays (Ehleringer and Forseth 1980; Koller 1990, 2000;
Vandenbrink et al. 2014). This phenomenon of solar tracking occurs widely in the pho-
tosynthetic pathway of C3 and C4 plants in many families. Sun‐induced leaf movement
is often described as heliotropism and sun‐avoidant leaf movement as paraheliotropism.
Leaves that maximize light interception by solar tracking are called diaheliotropic
leaves. Leaves orientate perpendicular to direct sun rays for tracking, absorbing maxi-
mum light throughout the day thereby increasing photosynthetic rates. Some solar‐
tracking plants avoid direct exposure to sunlight. Leaves that are oriented parallel to the
direct sun rays show reduced leaf temperatures and decreased water loss through tran-
spiration (Ehleringer and Forseth 1980; Koller 1990, 2000). Solar tracking of leaves sig-
nificantly contributes to improving photosynthesis, which especially beneficial for
plants that grow in arid regions. Recent studies provide evidence for the function of the
circadian clock in modulating heliotropic movement during the day and reorientation
of movement during the night (Harmer 2009; Kutschera and Briggs 2016).

4.4.3 Light Absorption by Sun and Shade Adapted Leaves

Some of the plants have a great ability to adapt to light regimes of their habitats. Leaves
that are adapted to low light intensity environments are unable to grow in high light. In
shade species, light absorption is greatly dependent on chloroplast movement and more
70 4 Effect of Light Intensity on Photosynthesis

than 10% of absorptance changes are observed between high‐ and low‐light treatments
(Davis et al. 2011). Besides the differences of anatomical characteristics of shade and
sun adapted leaves, shade leaves have more total chlorophyll and the ratio of chloro-
phyll b to chlorophyll a is high. Sun adapted leaves have more rubisco and xanthophyll
cycle components. Light harvesting mechanisms are different in shade and sun plants.
Some of the shade plants exhibit a 3 : 1 ratio of PSII to PSI reaction centers while the
ratio is 2 : 1 in sun plants (Anderson 1986). Some shade plants add more chlorophyll to
PSII. All these adaptations are for the enhancement of light absorption.

4.5 ­Light and Photosynthetic Responses

The relationship between photosynthetic CO2 assimilation and photon fluxes provide
useful information on photosynthetic properties. CO2 uptake exactly balances the CO2
released at a certain point and it is called the LCP. The LCP varies with species, growth,
and developmental conditions of plants. Plants grown in full sunlight and those grown
in the shade show a significant difference in LCP, determining a plant’s tolerance to low
light levels (Walters and Reich 1999). Shade plants, with lower LCPs, survive in light‐
limited environments (Taiz and Zeiger 2002; Craine and Reich 2005).
Above the LCP, photosynthetic rate increases proportionally to photon flux and
reaches the maximum quantum yield. In intact leaves, measured quantum yields for
CO2 fixation is estimated to be in the range of 0.04 and 0.06 (Taiz and Zeiger 2002).
Quantum yield also varies with parameters such as temperature and CO2 concentra-
tion. Below 30 °C, quantum yields are generally higher in C3 plants than those of C4
plants and above 30 °C the quantum yield of C4 plants are higher. The level of light satu-
ration is lower in shade plants than those of sun plants. At higher photon fluxes the
photosynthetic rate saturates, indicating that other factors such as electron transport
and rubisco activity can limit photosynthesis.
Leaves are unable to utilize full sunlight. At a given time, only a small fraction of
leaves are exposed to full sunlight. Crop yield is highly dependent on the amount of light
received during the growth season when other factors are not limiting.

4.6 ­Conclusion and Future Prospects

In higher plants photosynthetic reactions are regulated by the photons harvested
from specific wavelengths of light by the photosystems that are composed of chloro-
phylls. The PAR intensity requirement for different species at a given growth condi-
tion differs and leaves are capable of regulating the incident light they absorb. Too
high or too low PAR intensity causes photoinhibition and disrupts the photosynthetic
machinery. Photoprotective reactions dominate the regulation of highly dynamic
photosynthetic processes in response to a fluctuating light environment. Recent
genetics, molecular, physiological, biochemical, and functional genomics studies have
revealed a variety of genes and signaling molecules that are implicated in light‐medi-
ated regulation of photosynthesis. The majority of plants including important crop
species growing in a wide geographic range deal successfully with the excessive light
they receive by avoiding photodamage, often with a trade‐off between photosynthetic
 ­  References 71

efficiency and photoprotection. It is not well understood how agricultural crops

maximize crop productivity under high light intensities sensitizing the environment.
Understanding the genetic variations present in certain crop varieties to maintain
high photosynthetic efficiency at high light intensities can be manipulated to improve
photosynthetic productivity. Some plant varieties grown under shade conditions are
well adapted to maximize their photosynthetic efficiency under low light intercep-
tion. The molecular mechanisms that operate in these plants can be further studied
and applied to improve the photosynthetic efficiency of plants growing especially
under canopy cover and at high latitudes, ultimately increasing biomass production.

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