Principles of Crop Science

LESSON 2. Photosynthesis

Photosynthesis is a reaction occurring in green plants whereby plants utilize

water and the energy of sunlight to fix inorganic carbon dioxide in the form of organic
compounds, releasing oxygen in the process. In other words, the sun’s energy is
transformed by plants through photosynthetic processes into chemical energy usable
by other living organism. The importance of this process is more readily apparent
when we understand that plants are ultimately the source of all food.
The overall photosynthetic process can be written as:

The reaction reads as:

“Six molecules of carbon dioxide plus 6 molecules of water combined in the
presence of light energy and with the aid of chlorophyll to form one molecule of
glucose plus six molecules of oxygen.

1.1. Overview of the process of photosynthesis

The overall process described above is the result of two primary reactions – the
light dependent reaction and light-independent reaction (Figure 1).

Figure 1. Overview of the process of photosynthesis

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The light reaction captures light energy and stores it within two chemicals,
NADPH (nicotinamide adenine dinucleotide phosphate) and ATP (adenosine
triphosphate). These two molecules provide the energy needed to drive the second
stage of photosynthesis, known as the Calvin-Benson cycle, in which carbohydrates
(glucose) are made from CO2 and H2O.
NADPH – reduced form of
nicotinamide adenine
1.2. Light Dependent Reaction dinucleotide phosphate, a
To understand the light reaction, it is small, water-soluble molecule
helpful to focus on the path of three critical that acts as a hydrogen carrier
elements: energy, electrons and protons (hydrogen in biochemical reactions
ions). However, before considering the series of
ATP – adenosine triphosphate,
individual reactions that make up the light
a small, water soluble
reactions, let’s first examine the following: molecu;e that acts as an
energy currency in cells
1.2.a. Chloroplast – the site of photosynthesis
Remembering the internal structure of a leaf, it composed of mesophyll cells.
Within these cells, chloroplast is found which contain the light-absorbing green
pigment called chlorophyll.
The chloroplast is covered by a double membranous structure called the outer
and inner envelope. The proteinaceaous matrix (water-like phase) in the chloroplast
is called stroma. A massive membrane system of lamellae or the thylakoids is found
in the stroma. Chloroplast possess two types of thylakoids-the large ones, known as
stroma thylakoids, extends from one end to the other end of chloroplast, whereas
the small ones are disc shaped thylakoids which are closely packed at places to form
grana and are called grana thylakoids (Figure 2).
The light reactions of photosynthesis occur in the thylakoids while the light
independent reaction occurs in the stroma.

Figure 2. Structure of chloroplast – the site of photosynthesis

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1.2.b. Photosynthetic pigments

The pigments involved in the process of photosynthesis are known as
photosynthetic pigment. Pigments like chlorophylls and carotenoids (carotene and
xanthophylls) may be found in plant cell.
 Chlorophyll is the green pigment in the chloroplast and consists of chlorophyll a
and chlorophyll b.
 Carotenoids are lipids classified into two
Chlorophyll appears green to
groups: carotene is orange in color and our eyes because it absorbs
xanthophylls are yellow in color. Carotenoids light mainly in the red and
complement the absorption of light by the blue parts of the spectrum, so
chlorophyll. The energy absorbed by those only some of the light
pigments is passed to chlorophyll and used in enriched in green wavelengths
photosynthesis and thus act as accessory (about 550 nm) is reflected
pigments. They also protect the chlorophyll into our eyes. Hence, the
molecules from photo-oxidation by picking up green color of the leaves
oxygen radicals and converting them into
harmless molecular stage.

1.2.c. Absorption and Action Spectra

Sunlight is like a rain of photons (light particles) of different frequencies. From
about 400 nm (violet) to about 700 nm (red) in the electromagnetic spectrum (Figure
3) is considered the visible region. Plants capture sunlight by using pigment molecules
that absorb visible light wavelengths (400-700 nm).

Figure 3. The electromagnetic spectrum showing the visible region

The absorption spectrum of chlorophyll a and chlorophyll b indicate that these

pigments mainly absorb blue and red light (Figure 4). The action spectrum shows that
maximum photosynthesis takes place in the blue and red regions of the spectrum
(Figure 5). Between blue and red wavelengths, blue light carry more energy. However,

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the most efficient wavelengths of light in photosynthesis are those of red light, green
light is least effective.

Figure 4. The absorption spectrum

Figure 5. The action spectrum

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1.3. Light Reaction (also known as Light Dependent or Photochemical Reaction)

1.3.a. Gathering Sunlight: The Antenna System

Light dependent reaction happens in the thylakoid membrane. Within the
thylakoid membrane, there are two photosystems wherein light energy (photons) are
harvested. Photosystem I is located in stroma (non-stack) lamellae usually absorbs
far-red light of wavelengths (> 680 nm) and Photosystem II in grana (stacked)
lamellae absorbs red light of 680 nm.
Plants capture photons (sunlight energy
particles) by using pigment molecules that absorb
visible light (wavelengths from 400 to 700
nanometers). The main light-absorbing molecule is
chlorophyll and other light-absorbing molecules
(e.g. carotenoids) embedded in the thylakoid
membrane that makes up an antenna system. This
antenna system is designed to absorb light energy
and funnel it to a protein complex called a
reaction center (Figure 6). The reaction center
can use the energy to drive an electron uphill from
one site to another within the reaction center. The Figure 6. The Antenna System
job of the antenna system is to capture photons and change the light energy into
another form of energy known as excitation energy, which is a type of electronic
energy. The excitation energy can be thought of as a packet of energy that jumps
from antenna molecule to antenna molecule until it is trapped by a reaction center.

1.3.b. Electron Transport

The excitation energy trapped by a reaction center provides the energy needed
for electron transfer, which is the next step in the photosynthetic light reactions.
During electron transfer, individual electrons are removed from water and transferred,
by an electron transport chain, to NADP. During the electron transport in
photosynthesis, light energy pushes electrons up an energy hill in the reaction centers.
Subsequent electron flow in the electron transport chain is energetically downhill
and can be used to do work. Figure 7 shows the electron carriers that make up the
photosynthetic electron transport chain in a way that reveals the relative electronic
energy on the vertical scale (known as the Z-scheme).

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Figu
re 7. The Z-scheme diagram of electron flow

The electron transport pathway includes electron transfer from one site to
another within a protein, as well as electron transfer from one molecule to another.
Electron transfer from one molecule to another is possible because certain types of
molecules can easily give up or receive electrons. In biological electron transport
pathways, the electrons are always bound to a molecule (they are too reactive to
hang around free), which means that an oxidation reaction is always coupled to a
reduction reaction. Electrons spontaneously jump from one molecule to another
because some molecules hold onto their electrons more tightly than others.

1.3.c. NADPH Production

Moving an electron from water to NADP requires an input of energy. This job is
done by reaction centers, which use the light energy gathered by the antenna system
to move an electron energetically uphill. As shown in Figure 8, the electron transport
chain in chloroplasts uses two different types of reactions centers: Photosystem II and
Photosystem I. Also, electron transfer from water to NADP requires three membrane-
bound protein complexes: Photosystem II, the cytochrome bf complex (Cyt bf), and
Photosystem I. Electrons are transferred between these large protein complexes by
small mobile molecules (e.g. plastoquinone (PQ) and plastocyanin (PC)).
Photosystem II catalyzes two different chemical reactions:
c.1. Oxidation of water (also known as photolysis of water) - this
process creates O2, which is released, and H ions, which are used in ATP
synthesis. Photosystem II performs this reaction by binding two H2O molecules
and removing one electron at a time. The energy for the removal of a single
electron is provided by a single photon. For Photosystem II to completely

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oxidize two H2O molecules and reduce two molecules of plastoquinone, it

requires four photons.
c.2. Reduction of plastoquinone by adding two electrons taken from
H2O molecules and two protons taken from the outer water phase (stroma)
creates PQH2 (or reduced plastoquinone).

The PQH2 molecule unbinds from Photosystem II and diffuses in the

photosynthetic membrane until it encounters a binding site on the cytochrome bf
complex. The cytochrome bf complex removes the electrons from PQH2 and releases
protons into the inner water space of the photosynthetic vesicle (lumen). The
cytochrome bf complex then gives up the electrons to another small molecule,
plastocyanin (PC) then the electrons are transferred to the Photosystem I. The
proton gradient, produced by water oxidation and
oxidation of reduced plastoquinone, is used to
create ATP (1.3.d). Always remember the
The Photosystem I reaction center is like abbreviation: OILRIG
Photosystem II in that it is served by a chlorophyll- (oxidation is loss, reduction is
containing antenna system and uses light energy to gain) meaning when a
move an electron energetically uphill, but compound gains an electron it
is said to be reduced
Photosystem I catalyzes different reactions: it
(reduction), whereas when it
oxidizes plastocyanin and reduces ferredoxin.
gives up an electron it is said
Ferredoxin itself becomes oxidized, losing its to be oxidized (oxidation).
electrons to another acceptor. The last step in the
photosynthetic electron transport chain is reduction
of NADP, producing NADPH.

Figure 8. Overview of the Mechanism of the Light Dependent Reaction

in the thylakoid membrane

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1.3.d. ATP Production

As previously noted, the electron transport chain concentrates protons in the
lumen by the release of protons during the oxidation of water by Photosystem II and
by transporting protons from the stroma to the lumen via plastoquinone (Figure 8). In
addition, electron transport creates a net positive charge on the inner side and a net
negative charge on the outer side of the vesicle, which gives rise to an electric
potential across the membrane. The energy stored in the pH gradient and electric
potential is known as the transmembrane proton electrochemical potential or the
proton motive force.
The conversion of proton electrochemical energy into the chemical-free energy
of ATP is accomplished by a single protein complex known as ATP synthase, which
catalyzes the formation of ATP by the addition of inorganic phosphate (Pi) to ADP.
The reaction is energetically uphill and is driven by the transmembrane proton
electrochemical gradient. Protons move through a channel in the ATP synthase
protein (from the lumen to the stroma) providing the energy for ATP synthesis.
However, the protons are not involved in the chemistry of adding phosphate to ADP at
the catalytic site. Although it has not been proven, it appears that proton flow drives
the rotation part of the ATP. The rotation of ATP synthase can be thought of as
pushing ADP and Pi together to form ATP and water.

Note: From the light reactions to the carbon reduction reaction, the job of the light
reactions is to provide energy in the form NADPH and ATP for the Calvin-Benson cycle.

1.4. Carbon Reduction reaction (also known as Light Independent or Biochemical

Reaction)
Carbon reduction reaction happens in the stroma. The CO2 fixation can occur via
three pathways: 1) C3 or Calvin-Benson Pathway; 2) C4 or Hatch Slack Pathway; 3)
Crassulacean Acid Metabolism or CAM Pathways.

1.4.a. C3 or Calvin Benson Cycle

The C3 cycle is so named because the carbon atom of a molecule of CO2 taken
up by leaf is first detected in the three-carbon compound 3-phosphoglyceric acid
(PGA). PGA is formed when CO2 combines with a 5-carbon sugar, ribulose biphosphate
(RuBP). The reaction is catalyzed by the enzyme RuBP carboxylase/oxygenase
(RuBisCO), an abundant protein in all green tissues.

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During each turn of the

Calvin-Benson cycle, two molecules
of PGA (a total of six carbon atoms)
undergo a complex series of
enzyme-catalyzed transformations
to regenerate RuBP, which then
combines with CO2 to form two
PGAs and complete the cycle. So,
of the six original carbon atoms in
PGA, five give rise to RuBP and the
one remaining appears as one of
the six carbon atoms in the sugar
glucose-6-phosphate (G6P).
Therefore, for every six CO2 Figure 9. Calvin Benson Cycle or
molecules fixed, one G6P leaves C3 pathway of CO2 fixation
the Calvin-Benson cycle for synthesis of starch, sucrose, cellulose, and ultimately all
of the organic constituents of the plant (Figure 9).
The vast majority of higher plants and algae are C3 species.

1.4.b. C4 or Hatch-Slack Pathway

The defining feature of CO2 fixation in C4 pathway is involvement of two
distinct cell types. Microscopic examination of leaf sections of C4 plants reveals two
chloroplast-containing cell types in an arrangement termed Kranz anatomy. Bundle
sheath cells form a cylindrical layer one cell deep around each leaf vein. These cells
are typically enlarged, thick walled, and densely packed with chloroplasts. At least
two layers of loosely packed mesophyll cells separate adjacent bundle sheath strands
(Figure 10).

Kranz Anatomy

Figure 10. Comparative anatomy of C3 and C4 leaf

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In C4 pathway, when CO2 enters the leaf it is first fixed in the mesophyll cells
by the enzyme phosphoenolpyruvate (PEP) carboxylase. The carbon atom from CO2
is first detected in the four-carbon organic acid oxaloacetic acid (OAA), hence the
name C4. The OAA is then reduced to malic acid or converted to aspartic acid
depending on species. Malate and aspartate are transported to bundle sheath cells
where they are decarboxylated, thereby
releasing CO2. This newly formed CO2 is
refixed by RuBisCO and metabolized by
the Calvin-Benson cycle present in the
bundle sheath. The remaining three
carbon atoms derived from the malate
and aspartate are transported back to
the mesophyll cells as pyruvic acid to
regenerate the three-carbon PEP (Figure
11).
Familiar species possessing the C4
photosynthesis mechanism are maize,
sorghum, sugarcane, and several
Figure 11. C4 pathway of CO2 fixation
common weeds.

1.4.c. Crassulacean acid metabolism (CAM) pathway

The crassulacean acid metabolism (CAM) pathway was discovered first in plants
of the family Crassulaceae but familiar species include pineapple and cacti. It is
considered an adaptation to life in arid environments. CAM photosynthesis resembles
C4 photosynthesis in terms of the pathway of fixation of carbon. The prominent
difference, however, is that CAM plants take up CO2 from the atmosphere at night
and synthesize malic acid via PEP carboxylase. During daytime, the stomata that
admit CO2 close to conserve water. Malic acid is decarboxylated and the CO2 is refixed
by the Calvin-Benson Cycle. Some of the starch accumulated during daytime is
converted to PEP at night to support CO2 fixation (Figure 12).

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Figure 12. CAM Pathway of CO2 fixation

1.5. Distinct Features of C3, C4, and CAM Plants

C4 plants are less efficient than C3 plants in terms of the energy requirements
in fixing CO2. To fix one molecule of CO2, C4 plants need five ATPs whereas C3 plants
need only three. However, C4 plants have higher photosynthetic rates than C3 plants
and also are able to continue photosynthesizing under conditions such as high
temperatures and high light intensity when C3 plants cannot. Generally, C4 plants are
adapted to tropical conditions. Under hot, sunny skies, C3 plants undergo a process
called photorespiration (light-dependent production of glycolic acid in chloroplast and
its subsequent oxidation in peroxisomes). C4 plants use CO2 more efficiently and
hence are able to function at only partially closed stomata, as occurs on hot, sunny
days.
CAM plants are relatively slower growing that C3 or C4 plants under favourable
conditions. They grow more slowly because plants, by nature, tend to conserve
moisture and in so doing close the stomata most of the day, thus limiting the CO2
intake needed for fixation.

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Table 1. Summary of the Characteristics of C3, C4 and CAM Plants

Characteristics C3 plants C4 plants CAM plants
Plant species Rice, taro, sweet Corn, sorghum, Agave, bromeliads
following the potato, cassava, sugarcane, amaranth such as pineapple,
pathway mungbean, spinach, orchids, succulents
wheat, potato,
tobacco, sugarbeet,
soybean, sunflower

Biomass Moderately productive, Highly productive, 80 Very poor

Production 30 tons per ha. is tons per ha. for productivity
possible for sunflower sugarcane

Leaf anatomy Mesophyll with no Mesophyll with Mesophyll with large

distinct bundle sheath distinct bundle sheath vacuoles
(Kranz anstomy)

Initial CO2 Ribulose 1,5 Phosphoenol pyruvate PEP in the dark;

acceptor biphosphate (RuBP) (PEP) RuBP in the light

First stable 3-phosphoglycerate (3- Oxaloacetate (OAA), a OAA in the dark and
product PGA), a 3-carbon 4-carbon compound 3-PGA in the light
compound

CO2 fixation Only one CO2 fixation Two CO2 fixation Two CO2 fixation
pathway pathway are pathways are
separated in space separated in time

The triphosphate produced in the Calvin-Benson Cycle (in C3, C4, and CAM
pathways) is the precursor to produce sucrose and starch, the two major end products
of photosynthesis, is starting point for a wide range of biosynthetic pathways. Sucrose
is the main form of carbohydrates translocated in the plant. It is therefore and
important raw material for the synthesis in different parts of the plant of many other
organic molecules that are necessary for growth, maintenance, and yield formation.

1.6. Factors Affecting Photosynthesis

Photosynthesis and consequently, crop productivity are influenced by a host of
variable internal and external factors.

1.6.a. Internal Factors

a.1. Nutritional status of plant. A deficiency of any of the essential
element will cause a decrease in the photosynthetic rate of the leaves. For

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example, deficiency of N and Mg in the chlorophyll may affect its formation,

Fe deficiency my affect chlorophyll synthesis, N deficiency may affect
photosynthetic rate in rice

a.2. Stomatal aperture. The stomata openings must be great enough for
passage of adequate levels of CO2. Stomatal aperture is reduced by water
stress, insufficient light and high CO2 concentration

a.3. Leaf age. The photosynthetic rate of young leaves is usually low but
increases as the leaf approaches full expansion. Old, senescent leaves
eventually become yellow and are unable to photosynthesize because of
chlorophyll breakdown.

1.6.b. External Factors

b.1. Light (irradiance). Photosynthetic
rate is partially a function of the level
of irradiance. In total darkness, no
photosynthesis occurs and as
irradiance increases, photosynthesis
increases until the light compensation
point is reached. The doubling of
available light during summer explains
why light is often a limiting factor to
high yields during wet season. Figure
13 shows the marked difference in
light response curves of C3 and C4
Figure 13. Light response curves of
plants. typical C3 and C4 plants

b.2. Temperature. The light dependent reaction of photosynthesis is little

affected by temperature but the carbon reduction reaction is very
dependent. Enzymatically controlled reactions such as photosynthesis and
respiration occur in a temperature range greater than 0ºC but less than 50ºC.
b.3. CO2 concentration. Photosynthetic rates are enhanced by higher CO2
concentration unless stomata are closed by drought or water stress. The
response of photosynthesis to CO2 is similar to its response to irradiance.

b.4. Moisture. Water is needed for photosynthesis but only about 0.1% of
the total water absorbed is used by the plant for this purpose, mostly is
used in transpiration process. Only that supply of water affects stomatal

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opening (when water is deficient, stomata is closed hence the rate of

photosynthesis is reduced).

Suggested instructional videos to watch:

 Photosynthesis Overview:
https://www.youtube.com/watch?v=PiAUPg4UrrE
 Light Dependent Stage:
https://www.youtube.com/watch?v=sYDzACaYLhc&feature=share&fbclid=Iw
AR1SAZEsFKX-pjwccqMkNaSAM_QQEJwAL9qK8l_FPENe3ixIQSyNjBsHRPk
 Light Independent Stage:
https://www.youtube.com/watch?v=dO987q1v0c0&feature=share&fbclid=Iw
AR0tGecQLtTxpz7JvVBVzpRCD_RIwVB2dqHDxMl0aQpcrPyTYn5rptEh4ss

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