Module 3

Section 4

This module will help you gain knowledge on the process of food making in autotrophs and other
photosynthetic organisms aside from the plants. Explain the stages of photosynthesis, chlorophyll, role of
action spectrum of photosynthesis, wavelength of light that drive photosynthesis, light harvesting
complexes and the electron transport chain.

Learning Outcomes:
After examining this module, you are expected to:
1. Discuss the process of photosynthesis.
2. Differentiate the photosynthetic reactions with respiration

Content/Information Sheet:
Photosynthesis is necessary to all living organisms on earth especially the plants and animals
which depends on it. The only biological process that capture energy from the sun and convert it into
chemical compounds(carbohydrates) that every organism uses to power its metabolism

 Joseph Priestley (1772) and later Jan Ingenhousz (1779) showed that plants have the ability to
take up CO2 from the atmosphere and release O2.
 Ingenhousz also discovered that release of O2 by plants was possible only in presence of
sunlight and by the green parts of the plant.
 Robert Hill (1939) demonstrated that isolated chloroplasts evolve O2 when they are
illuminated in the presence of electron acceptor which gets reduced. This reaction called Hill
reaction accounts for the use of water as a source of electrons and protons for CO2 fixation
and release of O2 as by-product
Importance of Photosynthesis
1. Primarily for capturing sunlight’s energy and transform into chemical energy stored in
carbohydrates and other organic molecules as high electrons in the carbon-carbon bonds of the
carbohydrates
2. Carbohydrates synthesis via photosynthesis are the energy source that heterotrophs use to
power ATP synthesis via respiration
3. Photosynthesis powers 99% of Earth’s ecosystem

Heterotroph types:
Heterotrophs are organisms that depend on external source of organic compounds. They are considered
ad other feeders and they must rely on the sugars produced by photosynthetic organisms for thei
energy needs.
Carnivores eat the meat of other animals.
Herbivores eat plants.
Omnivores can eat both meat and plants.

CMB@RJAELE_USM2020 0
 Scavengers eat things left behind by carnivores and herbivores.
Decomposers break down dead plant or animal matter into soil.
Detritivores eat soil and other very small bits of organic matter.

Autotrophs are organisms surviving on carbon dioxide as their principal carbon source.
Photoautotrophs – responsible for capturing the energy that fuels the activities of most
organisms on Earth. They use light to manufacture their own food (self feeders using light). This
include plants, algae and cyanobacteria.

Chemoautotrophs- utilize energy stored in inorganic molecules(such as ammonia, hydrogen

sulfide or nitrites) to convert CO2 into organic compounds. A group of bacteria that synthesize
sugars by not using sunlight energy but by extracting energy from inorganic chemical
compound.

Chloroplast Structure and Function:

 Chloroplast are located predominantly in the mesophyll of leaves,
 Mesophyll -site of photosynthetic activity in the leaves
 Stomata – small regulated (due to flanking of guard cells) openings(either swelling or
shrinking in response to osmotic change) in the leaves where gas exchange of carbon
dioxide and oxygen occurs, located on the leaf underside to minimize water loss
 Chloroplasts of higher plants are generally lens-shaped, their dimensions make chloroplast
as giants among organelles
 Identified as site of photosynthesis in 1881 by T. Engelmann

Source: https://courses.lumenlearning.com/suny-wmopen-biology1/chapter/photosynthesis/
Figure 1. Chloroplast Structure

Parts and Functions of the Chloroplast

1. Double membrane envelope composed of outer membrane and inner membrane
2. Thylakoids – stacked, disc- shaped structures that contain photosynthetic pigments
Thylakoid membrane is where chlorophyll is embedded
 Chlorophyll – pigment that absorbs light and responsible for the initial interaction between
light and plant material and numerous proteins that make up the electron transport chain
Thylakoid lumen – internal space in the thylakoid enclosed by thylakoid membrane
3. Grana- stack of thylakoids
4. Stroma(“bed”) – liquid filled space surrounding the granum

CMB@RJAELE_USM2020 1
Photosynthetic Pigments:
Pigments are compounds that appear colored because they only absorb light of particular
wavelength within visible spectrum. It reflects or transmit the wavelengths they cannot absorb and
making them appear in corresponding color.
Plant pigments are classified as either chlorophylls or carotenoids. found in plants and algae.
1. Chlorophyll
 most important naturally occurring pigment on the planet, found in plants and
cyanobacteria.
 These porphyrin play an important role in the conversion of solar energy to chemical energy
via photosynthesis.
 Chlorophyll and heme groups are referred to as colors of life
 Chlorophyll term(Gk: chloros- green, phyllon-leaf) was coined in 1818 by Joseph Pelletier
and Joseph Bienaime Caventou

Structure of Chlorophyll:

Source: https://students.ga.desire2learn.com/d2l/lor/viewer/viewFile.d2lfile/1798/12577/photosynthesis6.html
Figure 2. Chlorophyll Structure

Parts:
a. tetrapyrrole that contain a chelated metal ion(Magnesium) at the core
b. porphyrin head- composed of four rings of pyrrole with nitrogen arranged in a ring
c. Phytol tail - hydrocarbon chains linked though a carboxylic acid tail

 Chlorophylls reflect green light and absorbs violet, blue and red wavelengths of light used in
photosynthesis.
- Chlorophyll a = absorbs less blue light, blue/green color and is only directly involved in
light reactions of photosynthesis; present in all oxygen-producing photosynthetic
organisms, considered as the reaction center
- chlorophyll b = absorbs more red olive green color and assists chlorophyll a in capturing
light energy and is considered as accessory pigment (harvesting center); present in all
higher plants and green algae, pigment is found in antenna complexes of PSII(light
harvesting complexes)
- chlorophyll c = found in brown algae, diatoms and certain protozoa; co-occurs with
chlorophyll a and carotenoids; has magnesium-phytoporphyrins, contains propenoic
acid at C17 ring that replaces propionic acid side chain found in chlorophylls a and b;
brownish-golden color
- chlorophyll d = found in red algae and some cyanobacteria, minor pigment involved in
capture of the red spectrum of light(far end spectrum of red light), has formyl group at

CMB@RJAELE_USM2020 2
 C3-vinyl group located in chlorophyll a, found in A. marina which makes it them thrive in
environments with less sunlight, absorb light that had been depleted of the red and blue
wavelengths of light
- chlorophyll f = most recent chlorophyll form identified, found in wet lands
cyanobacteria, an accessory pigment in antenna system of cyanobacteria, capable of
absorbing high amounts of near-infrared light, energy is harvested from a different light
wavelength, important discovery given that oxygen producing microorganisms during
photosynthesis were not capable of using infrared light as source of energy.
- bacteriochlorophyll – found only in green and purple bacteria, organisms that do not
produce oxygen during photosynthesis
 While carotenoids, another accessory pigment(another harvesting center) in thylakoid
membrane, reflects red, orange and yellow range.
- Carotenoids are masked by chlorophyll until chlorophyll breaks down during autumn
- Carotenoids are accessory pigments because they cannot transfer sunlight directly to
the photosynthetic pathway but by passing their absorbed energy to chlorophyll and
eventually transfer to the photosynthetic pathway.
- Carotenoids are much larger group of pigments like lycopene(red of tomato), zeaxanthin
(yellow of corn), beta-carotene(orange in orange peel) and these are attracting seed
dispersers.
- Carotenoids in photosynthesis functions as very efficient molecules for disposal of
excess energy and safely dissipate energy as heat.

Source:https://med.libretexts.org/Bookshelves/Nutrition/Book%3A_Intermediate_Nutrition_(Lindshield)/12%3A_Blood_Bones_and_Teeth_Micronutrients/1
2.06%3A_Vitamin_A/12.6A%3A_Carotenoids
Figure 3. Structure and Classification of Carotenoids.

Table 1. Basic classes of photosynthetic pigments .

Pigment Peak Absorbance Reflects
Chlorophyll Chlorophyll a 400-450nm green
Chlorophyll b 450-500nm yellow
Chlorophyll c1
Chlorophyll c2
Chlorophyll d
Chlorophyll f
Bacteripchlorophyll
Carotenoids(alpha and 425-475nm
beta forms

CMB@RJAELE_USM2020 3
 Phycobilins – in red Wavelengths not Red, orange, blue
algae and absorbed by
cyanobacteria chlorophyll a

Uses of Chlorophyll:
 for commercial purposes in agriculture and food industry
 for medicinal purposes
- detoxification
- wound healing
- constipation
- reduce colostomy odor
- reduce bad breaths

Why are leaves green?

Leaves are green because their chloroplast contain large quantities of chlorophyll which absorbs
most strongly in the blue and red wavelength leaving the intermediate green wavelengths to be reflected
to our eyes.

Photosynthetic pigments are large, hydrophobic molecules embedded in photosystems(protein

pigment complexes) that work like antenna to collect sun’s energy. Photosystems are embedded in the
thylakoid membranes of the chloroplasts.
Each pigment type can be identified by the specific pattern of wavelengths it absorbs from
visible light, the absorption spectrum- resulting array of colors ranging from red to violet.(Figure 4).

Source: https://courses.lumenlearning.com/suny-wmopen-biology1/chapter/photosynthesis/
Figure 4. (a) chlorophyll a, (b) chlorophyll b, (c) beta carotene are hydrophobic organic
pigments found in the thylakoid membrane. (a) and (b) are identical except for the part
indicated in the inbox at upper right corner, are responsible for the green color ofm leaves
(c) is responsible for the orange color in carrots. (d) each pigment has unq=ique absorbance
spectrum.

CMB@RJAELE_USM2020 4
Events of Photosynthesis:
Photosynthesis is a multi-step process that requires sunlight, carbon dioxide (which is low in
energy), and water as substrates. After the process is complete, it releases oxygen and produces
glyceraldehyde-3-phosphate (GA3P), simple carbohydrate molecules (which are high in energy) that can
subsequently be converted into glucose, sucrose, or any of dozens of other sugar molecules. These sugar
molecules contain energy and the energized carbon that all living things need to survive.

Photosynthetic equation: 6CO2 + 12H2O Chlorophyll C6H12O6 + 6H2O + 6O2

Sunlight

2 Phases of Photosynthesis :
A. Light dependent reactions: convert solar energy into chemical energy(ATP and NADPH)
Stage 1
- Occurs in the thylakoids
- Energy is captured from sunlight
- Water is split into hydrogen ions, electrons and oxygen
- Photolysis is splitting of water molecule
- Traps energy from sunlight and does not involve CO2 and does not make sugars
- The oxygen diffuses out of the chloroplasts as by product
Stage 2.
- Occurs in thylakoids
- Light energy is converted to chemical energy which is temporarily stored in ATP and
NADPH
- The electrons that absorbed the energy are passed along the Electron Transport Chain
and store energy in ATP, NADP picks of the H+ to form NADPH and electrons from
photolysis and stores it for later use

Source: https://courses.lumenlearning.com/suny-wmopen-biology1/chapter/photosynthesis/
Figure 6. The reaction centers(Photosystem II and Photosystem I).

What are Reaction Centers?

- Contains a pair of chlorophyll a molecules with special property which undergo
oxidation upon excitation, give up an electron via photoact.
- Site where light energy is converted into an excited electron
 All step involved getting an electron onto the energy carrier NADPH for deliver to the Calvin
cycle(where electron is deposited onto carbon for long term storage in carbohydrate form)

CMB@RJAELE_USM2020 5
Photosystem II and I
- major components of the Electron Transport Chain including cytochrome complex,
- site that converts light energy into chemical energy
- contains a number of antenna proteins to which chlorophyll molecules are bound,
surround the reaction center where photochemistry occurs

Cytochrome complex – enzyme composed of two protein complexes; transfers electrons from the
plastoquinone (Pq a carrier molecule) to the plastocyanin (Pc, a protein)
- Both transfer the protons across the thylakoid membrane and the transfer of electrons
from PSII to PSI

Source: https://www.visionlearning.com/en/library/Biology/2/Photosynthesis-I/192
Figure 7. Electron transfer in Photosystem.

Electron Transport Chain in Photosynthesis:

Photophosphorylation – process where the energy rich electrons pass down the electron transport
system and releases energy which is sufficient to bind inorganic phosphate (Pi) with ADP to form ATP in
the presence of light.
I. Noncyclic Electron transport/ Non cyclic photophosphorylation
Water -> PSII -> ETC -> PSI -> NADPH -> Calvin Cycle
(low E) (energized) (E used for ATP) (re-energized) (high E) (high E)
 The simple noncyclic flow of electrons(z-Scheme, unidirectional) produces ATP and NADPH in equal
amounts.


Source: https://cbseworld.weebly.com/uploads/2/8/1/5/28152469/lesson-11.pdf
Figure 8. Non cyclic photophosphorylation

II. Cyclic Electron Transport/ Cyclic photophosphorylation

Water -> PSII -> ETC -> PSI -> NADPH -> Calvin Cycle

(low E) (energized) (E used for ATP) (re-energized) (high E) (high E)

CMB@RJAELE_USM2020 6
  Cyclic electron flow provides boost of ATP so that Calvin Cycle has the energy requirement
for sugar synthesis

Source: https://cbseworld.weebly.com/uploads/2/8/1/5/28152469/lesson-11.pdf
Figure 9. Cyclic photophosphorylation

Table 1. Difference between types of photophosphorylation.

Cyclic photophosphorylation
 Only PSI is functional
 Electron comes from the chlorophyll
P700 molecule and returns to the same
chlorophyll P700
 Mainly found in photosynthetic
eubacteria e.g. purple Sulphur bacteria
Non cyclic photophosphorylation
 Both PSI and PSII are functional
 Water is the primary source of the
electrons and H+, undergo photolysis and
NADP is the final acceptor of the
electrons and H+ ions
 Oxygen is evolved as by-product
 Takes place in all green plants and
cyanobacteria except photosynthetic
eubacteria

PSII (P680)
- Captures energy to create proton gradients to produce ATP
- Delivers high energy electrons to primary electron acceptor and through the Electron
Transport Chain (Pq to cytochrome complex to Pc) to PSI
- Missing electron is replaced by extracting a low-energy electron from water(photolysis-
water splitting)
- Re-reduced after every photoact
Photolysis - water splitting, process to release of electrons ( 2H+ and O2 atoms as waste product)
- Required to form one molecule of diatomic oxygen gas
- 10% of oxygen is used by mitochondria in the leaf to support oxidative phosphorylation
and the remaining O2 escapes to the atmosphere used by aerobic organisms to support
respiration
 Electron moving through the proteins between PSII and PSI lose energy
 The lost energy is used to move H+ atoms from stromal side of the membrane to the
thylakoid lumen
 In the thylakoid lumen, H+ atoms and ones produced from photolysis are accumulated and
later used to synthesize ATP

CMB@RJAELE_USM2020 7
PSI(P700)
- Captures energy to reduce NADP+ into NADPH
- Site where lost energy prior to arrival at PSI is re-energized by PSI, by absorbing another
photon in the PSI antenna
- Oxidized and sends a high energy electron to NADP+ to form NADPH

***The two photosystems work together to guarantee that the production of NADPH will roughly equal
the production of ATP. Other mechanisms exist to fine tune that ratio to exactly match the chloroplast’s
constantly changing energy needs.

Table 2. Differences between Photosystems I and II

Photosystem I
 Has a reaction centre of chlorophyll a
molecule with maximum light
absorption at 700nm wavelength and
is referred to as P700
 Primary electron acceptor is an iron
protein (Fe-S protein)
 A set of electron carriers are
plastocyanin, ferredoxin and
cytochrome
Photosystem II
 Has a reaction center of chlorophyll a
molecule with maximum light
absorption at 680nm and is referred
to as P680
 Primary electron acceptor,
pheophytin is a modified chlorophyll
a molecule with 2 H atoms in place
of magnesium ion
 A set of electron carriers are
pheophytin, plastoquinone,
cytochromes

ATP Production:
- Happens in the thylakoid lumen creating a concentration gradient (like in respiration H
build up happens in the intermembrane space of the mitochondria)
- Passive diffusion of H+ ions from high concentration (in the thylakoid lumen) to low
concentration in stroma is harnessed for ATP generation just like in the Electron
Transport Chain of cellular respiration
- Ions build up energy because of diffusion and all have same electrical charge, repelling
each other
- To release the energy, H+ ions will rush through the opening in the specialized protein
channel of the ATP synthase by attaching to a third phosphate group of ADP, forming
the ATP molecule see Figure 8)
Chemiosmosis
- Process in which energy is stored as H+ gradient across a membrane
- Used to synthesize ATP from ADP and Pi with ATP synthase and its power source is
difference in the H+ ion concentration on opposite sides of the membrane
- flow of hydrogen ions through ATP synthase; ions move from an area of high
concentration to an area of low concentration through a semi-permeable structure.
 Chloroplast use chemiosmosis to generate ATP during photosynthesis
 Prokaryotes lack organelles like mitochondria and chloroplasts to generate H+ gradients
across plasma membranes and cannot use it for ATP synthesis
 Proposed by Peter Mitchell and won a Nobel Prize in 1978

CMB@RJAELE_USM2020 8
 B. Light Independent Reactions/Biosynthetic Pathways
- Occurs in the stroma of the chloroplasts
- Chemical energy stored in ATP and NADPH powers the formation of organic compounds(sugars) using
CO2 and H+ from NADPH
- Series of reactions that catalyze the CO2 reduction to carbohydrates (CO2 fixation)
- Reactions are independent of light i.e. light is not necessary but can continue in light as
well if products of the light reaction are available
- Carbon fixation reactions produce sugar in the leaves of the plant from where it is
exported to other plant tissues as the source of both organic molecules and energy for
growth and metabolism

Source: https://courses.lumenlearning.com/suny-wmopen-biology1/chapter/photosynthesis/
Figure 10. Light Dependent Reactions

- In plants, carbon dioxide (CO2) enters the leaves through stomata, where it diffuses over
short distances through intercellular spaces until it reaches the mesophyll cells.
- Once in the mesophyll cells, CO2 diffuses into the stroma of the chloroplast—the site of
light-independent reactions of photosynthesis.
- Calvin cycle, is named for the man who discovered it Melvin Calvin, and because these
reactions function as a cycle. Others call it the Calvin-Benson cycle to include the name of
another scientist involved in its discovery and C3 cycle is another term used for Calvin
Cycle as it initially generates a PGA (3-phospholglyceric acid) a first 3C stable product
through carboxylation (Figure 10).

CMB@RJAELE_USM2020 9
 Source: https://students.ga.desire2learn.com/d2l/lor/viewer/viewFile.d2lfile/1798/12577/photosynthesis6.html
Figure 11. Calvin Cycle

B.1 Calvin Cycle Stages

1. Fixation: enzyme RuBisCO (ribulose bisphosphate carboxylase) catalyze reaction between CO2
and 3 molecules of RuBP ( ribulose bisphosphate) to produce 2 molecules of 3-phosphoglyceric
acid
 Reaction between 1 molecule of CO2 and RuBP gives 2 molecules of 3-PGA
 In the stroma, in addition to CO2, two other components are present to initiate the light-
independent reactions: an enzyme called ribulose bisphosphate carboxylase (RuBisCO), and
three molecules of ribulose bisphosphate (RuBP), as shown in Figure 8. RuBP has five atoms
of carbon, flanked by two phosphates.
 RuBisCO catalyzes a reaction between CO2 and RuBP. For each CO2 molecule that reacts with
one RuBP, two molecules of another compound (3-PGA) form.
 PGA has three carbons and one phosphate. Each turn of the cycle involves only one RuBP and
one carbon dioxide and forms two molecules of 3-PGA. The number of carbon atoms remains
the same, as the atoms move to form new bonds during the reactions (3 atoms from 3CO 2 +
15 atoms from 3RuBP = 18 atoms in 3 atoms of 3-PGA). This process is called carbon fixation,
because CO2 is “fixed” from an inorganic form into organic molecules.
2. Reduction: products of light dependent reaction (ATP and NADPH) provide electrons that are used
to convert 3-PGA molecules into glyceraldehyde3-phosphate (See Figure 11)
 6 molecules of ATP and NADPH produce six 3C glyceraldehyde 3-phosphate
 ATP and NADPH are used to convert the six molecules of 3-PGA into six molecules of a
chemical called glyceraldehyde 3-phosphate (G3P).
 That is a reduction reaction because it involves the gain of electrons by 3-PGA.
 Six molecules of both ATP and NADPH are used. For ATP, energy is released with the loss of
the terminal phosphate atom, converting it into ADP; for NADPH, both energy and a hydrogen
atom are lost, converting it into NADP+. Both of these molecules return to the nearby light-
dependent reactions to be reused and reenergized.
3. Regeneration: single molecule of glyceraldehyde 3-phosphate is transported to the cytoplasm to
form various components needed by the cell, remaining molecules are used to regenerate RuBP
(see Figure 12).
 Interestingly, at this point, only one of the G3P molecules leaves the Calvin cycle and is
sent to the cytoplasm to contribute to the formation of other compounds needed by the
plant.

CMB@RJAELE_USM2020 10
  Because the G3P exported from the chloroplast has three carbon atoms, it takes three
“turns” of the Calvin cycle to fix enough net carbon to export one G3P. But each turn
makes two G3Ps, thus three turns make six G3Ps.
 One is exported while the remaining five G3P molecules remain in the cycle and are used
to regenerate RuBP, which enables the system to prepare for more CO2 to be fixed. Three
more molecules of ATP are used in these regeneration reactions.

Source: https://courses.lumenlearning.com/suny-wmopen-biology1/chapter/photosynthesis/
Figure 12. Calvin Cycle

B.2 C4 Cycle or Hatch –Slack Cycle

 This cycle is an adaptation for plants growing under dry hot environment( can photosynthesize
even in the conditions of very low CO2 concentration and under partial stomatal closure
 Sugarcane and maize can grow at low water content, high temperature and high light intensity
 Undergo photorespiration (RuBP oxidation with O2 present) is absent in C4 plants so
photosynthetic rate is high
 Leaf anatomy of C4 plants show Kranz anatomy (presence of dimorphic chloroplasts- Figure 13)

Source: https://cbseworld.weebly.com/uploads/2/8/1/5/28152469/lesson-11.pdf
Figure 13. Kranz anatomy in maize leaf (c.s.)

o Vascular bundles have a sheath of large parenchyma cells around then in wreath
form, thus Kranz anatomy (Kranz = wreath)
o Leaves possess dimorphic chloroplasts(2 types)
o Chloroplasts in mesophyll cells are smaller and have well developed grana(granal
chloroplasts) but do not accumulate starch
o Chloroplasts in the bundle sheath cells are larger and lack grana(agranal
chloroplasts) but contain numerous starch grains
 The initial acceptor of CO2 in C4 plants is phosphoenol pyruvic acid or PEP(3C compound). It
combines with CO2 in presence of phosphoenol pyruvate carboxylase (PEP carboxylase) and form

CMB@RJAELE_USM2020 11
 oxaloacetic acid(OAA)- first 4C stable product of the cycle, thus its name. This fixation of CO2
occurs in the cytosol of the mesophyll cells of the leaf.
 OAA travels to chloroplasts of bundle sheath cells where it releases the fixed CO2 and is also C3
cycle operates, thus CO2 immediately combines with RuBP in C3 cycle to produce
sugars(Figure14)
 C4 cycle involves PEP carboxylase(PEPCo) in the mesophyll and RuBP carboxylase (Rubisco) in the
bundle sheath cells

Source: https://cbseworld.weebly.com/uploads/2/8/1/5/28152469/lesson-11.pdf
Figure 14. C4 Pathway

Table 3. Differences between Independent Pathways.

C3 Plants C4 Plants CAM plants
1. Definition Plants whose first Conversion of sunlight Plants which store
product after carbon energy into 4C energy from the
assimilation from oxaloacetic acid(OAA) by sun and convert it
sunlight is 3C tropical plants, takes lace into energy during
phosphoglyceric acid( before C3 cycle and night follows the
3-PGA) for the further convert into CAM
production of energy energy
called C3 plants and More efficient that C3
pathway pathway
Used commonly by
plants
2. Cells involved Mesophyll cells Mesophyll cells Both C3 and C4
Bundle sheath cells same mesophyll
3. examples Sunflower, rice, beans Sugarcane, maize, Cacti, orchids
sorghum
4. seen in All photosynthetic Tropical plants Semi-arid
plants conditions
CMB@RJAELE_USM2020 12
 5. Kranz Leaf Only one type of Dimorphic chloroplasts or Absent ,
anatomy chloroplasts Kranz anatomy present xeromorphic
Kranz anatomy is
absent
6. Photorespiration Present in high rate, Absent, photosynthetic Detectable in the
excess of oxygen is an efficiency is further afternoon
inhibitor of increased
photosynthesis
7. Glucose 12NADPH and 18 ATPs 12 NADPH and 30 ATPs 12NADPH and 39
production are required are required ATPs are required
8. Plant types using Mesophytic, mesophytic xerophytic
the cycle hydrophytic
xerophytic
9. First stable 3-phosphoglyceric Oxaloacetate(OAA) Oxaloacetate(OAA)
product acid(3-PGA) at night and 3PGA
at daytime
10. Calvin cycle alone Along with Hatch and C3 and Hatch and
operative Slack cycle Slack cycle
11. Optimum 15-250C 30-400C >400C
temperature for
PS
12. Carboxylating RuBP carboxylase In mesophyll: PEP In the dark: PEP
enzyme carboxylase carboxylase
In bundle sheath: RuBP In light: RuBP
carboxylase carboxylase
13. CO2:ATP:NADPH2 1:3:2 1:5:2 1:6.5:2
ratio
14. Initial CO2 RuBP(ribulose-1,5- PEP(phosphoenol PEP(phosphoenol
acceptor phosphate pyruvate) pyruvate)
15. CO2 30-70 6-10 0-5 in dark
compensation
point
16. Typical all Tropical, elevated Dry,arid
environments daytime temperatures,
drought
17. Stoma open yes yes no
during the day?
18. Number of steps 1 2 2
in pathway

Crassulacean Acid Metabolism(CAM):

- A carbon fixation pathway that evolved in some plants as an adaptation to arid conditions, where
the stomata in leaves remain shut/close during the day to reduce evapotranspiration and open at
night to collect CO2
- Process of photosynthesis in desert plants (xerophytes – cacti and most succulents) to conserve
water and fix CO2 at night when stomates are open

CMB@RJAELE_USM2020 13
- Efficient use of CO2 allows plant to adapt to living with less water
- Temporary carbon fixation and storage process, conserve water due to cooler temperature at night
and carry out low levels of photosynthesis without opening the stomata at all ( mechanism for
surviving extremely dry periods
- Use phosphoenol pyruvate(PEP) carboxylase to capture CO2, which physically separate the CO2
fixation temporally(separated by time).
- CAM plants store CO2 mostly in malic acid via carboxylation of PEP to OAA which is the reduced to
malate
- Decarboxylation of malate during the day releases CO2 inside the leaves which allows carbon fixation
to 3-phosphoglycerate by RuBisCo
-

Source: https://biologydictionary.net/c3-c4-cam-plants/
Figure 15. CAM carbon fixation pathway

Factors Affecting Rate of Photosynthesis:

Internal Factors:
1. Chlorophyll- The amount of chlorophyll present has a direct relationship with the rate of
photosynthesis because this pigment is directly involved in trapping light energy responsible for
the light reactions
2. Leaf age and anatomy : Newly expanding leaves show gradual increase in rate of photosynthesis
and the maximum is reached when the leaves achieve full size. Chloroplast functions decline as
the leaves age. Rate of photosynthesis is influenced by variation in
a. number, structure and distribution of stomata,
b. size and distribution of intercellular spaces
c. relative proportion of palisade and spongy tissues and (iv) thickness of cuticle.
3. Demand for photosynthate : Rapidly growing plants show increased rate of photosynthesis in
comparison to mature plants. When demand for photosynthesis is lowered due to poor
meristematic activity, the photosynthetic rate declines.

External Factors:
a. Light Intensity – The rate of photosynthesis increases with increase of intensity of light
within physiological limits or rate of photosynthesis is directly proportional to light
intensity. Except on a cloudy day and at nights, light is never a limiting factor in
photosynthesis in nature.
- At a certain light intensity the amount of CO2 used in photosynthesis and the amount of
CO2 produced in respiration are the same. This point of light intensity is known as
compensation point.

CMB@RJAELE_USM2020 14
 Wavelength of light absorbed by photosynthetic pigments affects rate of photosynthesis.
Red light and to some extent blue light has an enhancing influence on photosynthesis. The
proportion of the total incident sunlight on earth, absorbed by green plants is generally a
limiting factor. As per the estimates of the total incident light reaching the green plants,
only about 1-2% is actually absorbed, because 70% is transmitted, and 28-29% is reflected
back into the atmosphere.
b. CO2 levels –carbon dioxide , one of the raw materials for photosynthesis, its concentration
affects the rate of photosynthesis. Because of its very low concentration (0.03%) in the
atmosphere, it acts as limiting factor in natural photosynthesis. At optimum temperature
and light intensity, if carbon dioxide supply is increased the rate of photosynthesis
increases markedly until CO2 conc. is as high as 3.0%. Thus, CO2 conc. in the atmosphere
is always a limiting factor for photosynthesis.
-increasing CO2 levels stimulate photosynthesis until it reaches plateau
c. Temperature- Raising temperature speed up the chemical reaction in photosynthesis,
directly proportional where rate of photosynthesis increases with temperature increase.
- It peaks at certain temperature and decreases when temperature is further increased
- Very high and very low temperature affect the rate of photosynthesis adversely.
- Rate of photosynthesis will rise with temperature from 5°-37°C beyond which there is a
rapid fall, as the enzymes involved in the process of the dark reaction are denatured at
high temperature.
- Between 5°-35°C, with every 10°C rise in temperature rate of photosynthesis doubles or
Q10 is 2 (Q = quotient), or slightly less than two.
d. Water - has an indirect effect on the rate of photosynthesis. Loss of water in the soil is
immediately felt by the leaves, which get wilted and their stomata close down thus
hampering the absorption of CO2 from the atmosphere. This causes decline in
photosynthesis
e. Oxygen - Concentration of oxygen as an external factor, is never a limiting factor for
photosynthesis because it is a by-product of photosynthesis, and it easily diffuses into the
atmosphere from the photosynthesizing organ, the leaf. However, excess of O2
surrounding a green plant, reduces photosynthetic rate by promoting the rate of aerobic
respiration.
f. Mineral elements - Some mineral elements like magnesium, copper, manganese and
chloride ions, which are components of photosynthetic enzymes, and magnesium as a
component of chlorophylls are important, and their deficiency would affect the rate of
photosynthesis indirectly by affecting the synthesis of photosynthetic enzymes and
chlorophyll, respectively

What is Concept of Limiting Factors?

When a process is affected by various factors, the rate of the process depends upon the pace of
the slowest factor. Let us consider three factors like light, carbon dioxide and temperature. It is seen that
when all three factors are optimum, the rate of photosynthesis is maximum. However, of the three factors
even if one of the factors becomes suboptimal and the other factors remain optimal, the rate of the
photosynthetic process declines substantially. This is known as law of limiting factors shown by Blackman
in 1905. It is defined as when a process is conditioned as to its rapidity by a number of separate factors,
the rate of the process is limited by the pace of the slowest factor which is known as the limiting factor.

CMB@RJAELE_USM2020 15
Chemosynthesis:
- Process of utilizing chemical energy released during biological oxidation of certain
inorganic substances (H2S and NH3) to reduce CO2 to carbohydrates by chemosynthetic
autotrophs
- Method of carbon assimilation when CO2 reduction is carried out in darkness
Chemosynthetic autotrophs – colorless bacteria that uses chemical energy to reduce CO2
a. Nitrifying bacteria – Nitrosomonas and Nitrobacter – oxidize NH3 to NO2
b. Sulphur bacteria
c. Iron bacteria
d. Hydrogen and methane producing bacteria

Table 4. Difference between photosynthesis and chemosynthesis

Chemosynthesis Photosynthesis
It occurs only in colourless anaerobic 1. This process occurs in all green plants bacteria
including green bacteria.
During this process CO2 is reduced light 2. CO2 and H2O are converted into to
carbohydrates in the presence of and chlorophyll. carbohydrates without light and chlorophyll
Here chemical energy released during oxidation of 3. Light energy is converted into chemical and
inorganic energy substances is used up to stored in the form of carbohydrates
synthesize carbohydrates.
No pigment molecule is involved and oxygen is not 4. Several pigments are involved and oxygen is
evolved. evolved as a by-product
No photophosphorylation takes place Photophosphorylation takes place i.e. ATP is
produced.

Summary:
 Photosynthesis transform life on earth
 Photosynthesis evolved to allow living things access to enormous amounts of energy to
build new structures and achieve biodiversity.
 Only photoautotrophs can perform photosynthesis through presence of chlorophyll
 Eukaryotic autotrophs(plants and algae) have chloroplasts where photosynthesis takes
place and starch accumulates while in prokaryotes(cyanobacteria) the process happens
within the folded membranes of the plasma membrane and the cytoplasm
 Chlorophyll a specialized pigment that absorbs certain portions of the visible spectrum
and capture energy from sunlight
 Photosynthesis needs carbon dioxide and water to assemble carbohydrate molecules and
release oxygen as waste product into the atmosphere
 Green plants are capable of synthesizing carbohydrates from CO2 and H2O in the
presence of light, by the process of photosynthesis
 During photosynthesis ‘light energy’, which is captured by the photosynthetic pigments
(chlorophyll, carotenoids and xanthophyll) present in the chloroplasts, is converted into
chemical energy.
 Photosynthesis in general is expressed by the following equation:
6CO2 + 12H2O -Chlorophyll/ Light → C6H12O6 + 6H2O + 6O2

CMB@RJAELE_USM2020 16
  Light reactions: take place in grana or thylakoids of chloroplasts only in the presence of
light.
 Dark reactions: occur in the stroma of chloroplast and are independent of light, if products
of light reaction are provided.
 Light energy is used for splitting of water, and production of ATP and NADPH2 and actual
reduction of CO2 takes place in the dark reaction
 Light reaction occurs with the help of two functional units, photosystem-I and
photosystem-II.
 During light reaction phosphorylation of ADP to ATP may occur in two ways, via cyclic
and non-cyclic.
 During dark reactions CO2 is accepted by Ribulose bi-phosphate (RuBP) and the first
stable product 3-PGA (3 phosphoglyceric acid) is formed, which by further cyclic reactions
(Calvin Cycle) leads to the formation of carbohydrates as well as in regeneration of RuBP.
 In C4 plants like maize and sugarcane the primary acceptor of CO2 is in mesophyll cells
and the first detectable product of dark reaction is oxaloacetic acid (OAA), whereas in the
bundle sheath cells CO2 fixation occurs through. Calvin cycle. CAM plants on the contrary
occurs temporally - differences in time.
 Occurrence of dimorphic chloroplasts in C4 plants is known as “Kranz anatomy” and is
characterized by the presence of a sheath of parenchyma cells around a vascular bundle
(bundle sheath). Cells of this sheath have larger chloroplasts which lack grana and are
filled with starch grains. In contrast mesophyll cells contain chloroplasts which are smaller
but have well developed grana.
 Rate of photosynthesis is influenced by (i) environmental factors such as light,
temperature, carbon dioxide concentration and water, and (ii) internal factors which
include age of leaf, chlorophyll content and leaf anatomy

References:
Chlorophyll: Definition, Structure, Function and Photosynthesis. 2020. Retrieved from:
https://www.microscopemaster.com/chlorophyll.html
https://byjus.com/biology/photosynthesis/
https://www.etsu.edu/uschool/faculty/tadlockd/documents/apbio_chp_10_lectout.pdf
https://www.researchgate.net/publication/221924762_Photosynthesis_How_and_Why
Lents, N.H. and J. Nishan. 2014. Photosynthesis I: Harnessing the Energy of the Sun. Retrieved from:
https://www.visionlearning.com/en/library/Biology/2/Photosynthesis-I/192
Lesson -11. Photosynthesis. Retrieved from
https://cbseworld.weebly.com/uploads/2/8/1/5/28152469/lesson-11.pdf
Overview of Photosynthesis. Retrieved from:
https://www.oercommons.org/courseware/lesson/14976/overview
Photosynthesis. Retrieved from: https://courses.lumenlearning.com/suny-wmopen-
biology1/chapter/photosynthesis/

CMB@RJAELE_USM2020 17