5.1 Photosynthesis - Part 1 - Hill RXN

TAT1203/SBS1202
Plant Science/Plant Physiology
4.1 Photosynthesis
Part 1
S. L. Ranamukhaarachchi
AgriTech Program
SLTC
October Semester 2023
Reference materials
1. Plant Physiology (third edition) (1982)
F.B. Salisbury & E.W. Ross
Publ. Wadsworth Publishing Co., Inc. California.
2. Biology (sixth edition) (2002)

N.A. Campbell & J.B. Reece
Publ. Pearson Education, Inc.
San Francisco.
*** Special Note – Images were taken from Google images and Biology
Textbook of Campbell and Reece (2002)
4.1 Photosynthesis - S.L. Ranamukhaarachchi - October Semester 2023
1
Photosynthesis
1. Capture of solar energy by plants

2. Photosynthetic carbon reduction in
plants
3. Synthesis of biochemical compounds in plants
4. Factors influencing the rate of
photosynthesis
5. Adaptations of plants to handle unfavourable
variations in the environment.
History of Photosynthesis
• Before 18th Century, scientists believed plants obtain all
requirements from soils.
• In 1771, Joseph Priestly- Green plants could renew air that was
made bad by animals.
• In 1782, Jean Senebier - the noxious gas produced by animals and
plants in darkness (i.e., CO2) stimulates the production of purified air
in the presence of light.
• In 1927, Stephen Hales – A part of the nourishment of plants comes

from the atmosphere that light participates some how in the process.
• In 1930, C. B. Van Neil - There is a similarity between overall

photosynthetic process in green plants and that in certain bacteria.
Robin Hill (1930) supported this. He named the light-driven split of
water in the absence of CO2 fixation as Hill Reaction.
• In 1941 - Evidence on O2 release and that is derived from water was
found.
• In 1954, Daniel Arnon discovered ATP in chloroplast.
2
Main role of light and plants
4/3/2024 4.1 Photosynthesis - S.L. Ranamukhaarachchi - October Semester 2023 5
Fate of light energy in brief………..
❑ The figure shows the fate of

light energy in plants.
❑ Plants capture light energy

through leaves.
❑ Leaf contains Chloroplasts
where light energy is truly
captured.
❑ ATPs are synthesized using

light energy, which could then
be able to release energy with
respiration.
Ref. Campbell & Reece (2002)
3
Light Intensity
• The amount of energy of the light beam per unit area
per unit of time
• Measured in Cal cm-2 s-1, W m-2
• The amount of solar energy impinging (striking) on a
unit surface per unit of time, calories cm-2 s-1, watts m-2
• The energy of an uninterrupted and unattenuated
beam is 2 cal cm-2 min-1
(Cal cm-2 = Langleys, and hence 2 Langleys min-1)
• This is called the “solar constant” but must be
measured outside Earth’s atmosphere with a light
sensor placed perpendicular to the light beam.
• This energy content is found above the Earth’s
atmosphere, otherwise, atmospheric particles absorb
energy in the light beam.
7
TAT1203 Plant Science - S.L. Ranamukhaarachchi - October Semester 2023
Light Quality
• Particularly the wave length
and its influence.
• Photosynthesis – all in the

visible radiation except,
green range, influences.
• Red and far-red have effects

on photomorphogenesis,
i.e., flowering, plant growth
toward sunlight, seed
dormancy, like actions.
4
9
4/3/2024
Light duration
• Light duration influences
photosynthesis and dry matter
production.
• This looks as daily light integral -
DLI.
• Light duration also influences

photomorphogenesis – flowering,
growth patterns of plants, and
indirectly on seed dormancy.
• Temperature of the earth’s
environment is also regulated by
the duration of light – summer
and cold winters.
10
5
Latitude and Radiation Distribution
11
Energy in Solar Radiation
h – Plank’s constant = 1.58 x 10-34 Cal Sec

6.626 x 10-34 J Sec
6.624 x 10-27 Erg Sec
c – Velocity of light = 3.0 x 1010 cm sec-1

3.0 x 10-8 m sec-1
λ - Wavelength
= Red 6.6 x 10-5 cm or 660 nm
= Blue 4.54 x 10-5 cm or 450 nm
TAT401 Controlled Environment Agriculture - S.L. Ranamukhaarachchi - October Semester 2023 12
12
6
3. Light and Light Utilization & Photosynthesis - S.L. Ranamukhaarachchi - August Semester 2021 13
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• Energy of radiation
Blue = 1.58 x 10-34 cal sec x 3.0 x 1010 cm sec-1

4.545 x 10-5 cm
= 10.5 x 10-16 Cal/photon
= 6.02 x 1023 photons/Einstein

= 10.5 x 10-16 Cal/photon x 6.02 x 1023 photons/Einstein
= 63,400 Cal/Einstein
Red = 1.58 x 10-34 cal sec x 3.0 x 1010 cm sec-1

6.6 x 10-5 cm
= 7.18 x 10-16 Cal/photon
= 43,234 Cal/Einstein
Einstein is a mole of photon.

Einstein = Avogadro number of photons, i.e., 6.02 x 1023 photons/mole.
TAT401 Controlled Environment Agriculture - S.L. Ranamukhaarachchi - October Semester 2023 14
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4.1 Photosynthesis - S.L. Ranamukhaarachchi - October Semester 2023 15
15
Measurement of Light Energy

• Einstein → Energy of one mole of photon
• PPF → Microeinsteins (= micromoles of photon)
• PPFD → (µE/m2/S) = (µMol/m2/S)
• When light energy reacts with other substances, it work

on mole basis.
• One mole of photon = Avogadro number of
photons/mole, i.e., 6.02 x 1023).
• PPFD works on per unit area basis, an intensity or
intensity of energy received.
TAT401 Controlled Environment Agriculture - S.L. Ranamukhaarachchi - October Semester 2023
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8
PAR, PPF and PPFD
• PAR - photosynthetic active radiation.
PAR describes the wavelengths of light in the visible range of 400-700nm.
• PPF – photosynthetic photon flux.

PPF applies to define the measurement of PAR. It stands for how much PAR
is being produced by any one lighting system over the period of a second.
• PPFD - photosynthetic photon flux density. PPFD is more relevant than the PPF, and
measures the PPF for a given surface area.
PPFD is measured in micromoles per square metre per second (µMol/m2/S)
It says exactly how many PAR photons are landing on a specific surface area.
Unit → (µE/m2/S) = (µMol/m2/S)
PPFD shows how many photosynthetic photons are actually impacting the
grow area and working when it comes to crop growth & yield.
TAT401 Controlled Environment Agriculture - S.L. Ranamukhaarachchi - October Semester 2023
17
Pigments and Energy
18
9
Estimation of Energy in different Wavelengths of
Solar Beam
One can estimate the energy
content of the light beam
practically by sending through
a prism and measuring the
energy content of each wave
length with a galvanometer.
Remember, chlorophyll
absorbs the energy content of
both blue and red wave
lengths, and hence
galvanometer records only a
slight energy level.
Remember, green wave

lengths are not utilized by
chlorophyll in plants.
19
Absorption of light energy
20
20
10
Pigments and absorption Spectra
https://bio.libretexts.org/Bookshelves/Introductory_and_General_Biology/Book%3A_General_Biology_(Boundless)/8%3A
_Photosynthesis/8.2%3A_The_Light-Dependent_Reactions_of_Photosynthesis/8.2B%3A_Absorption_of_Light
21
Chlorophyll a & b
• Chlorophyll a and b are

common in plants, and
contain four porphyrin rings
connected in the center by
Mg.
• The difference between

Chlorophyll a and b are
shown in the figure.
22
11
Ref: https://www.horti-growlight.com/en-gb/par-ppf-ypf-ppfd-dli
23
Wavelengths of solar beam and effectiveness in

photosynthesis
• The upper diagram shows the absorption

spectra of chlorophyll a and b.
• Chlorophyll a and b absorb usually blue wave

lengths more than red.
• Meanwhile, orange wave lengths are absorbed

by carotenoids.
• In the overall absorption pattern, blue and red

are absorbed very high and green in the middle
is absorbed minimally.
24
12
Pigments and light energy
absorbance
• Chlorophyll a - Blue-violet range – reflects green

• Chlorophyll b - Red & blue – reflects green
• Beta Carotene - Blue and orange
• Xanthophyll - Yellow
• Anthocyanin - Red
25
Photosynthetic tissues
❑ Leaf – the main part that

captures solar radiation.
❑ However, green materials

including stem, stalk, fruit
skin, etc. also can capture
solar radiation and carbon
fixation.
❑ Broadleaf and grass types

are having different leaf
structures.
26
13
Anatomy of leaves
Broadleaf (C3) Grass leaf (C4)
27
How radiation behaves in nature

The figure shows how light
behaves when strikes on a
chloroplast.
Three types of actions take

place for its energy when a
light bean impinges on a
chloroplast.
1. Reflection
2. Absorption
3. Transmission
The diagram also shows

the separation of light
beam into its original
colors that make visible
radiation when passing Ref. Campbell & Reece (2002)
through thylakoids.
28
14
Reflectivity, Absorptivity and Transmissivity
• Reflectivity (Ir)– Percentage light
reflected from total incidence
radiation. This refers to total
incidence radiation. Albedo
refers to reflectivity within visible
radiation (max 20%, generally
8%).
• Absorptivity (Ia) – Percentage
absorbed from incidence
radiation.
• Transmissivity (It) – Percentage
transmitted from incidence
radiation.
• Total radiation, I= Ir + Ia + It
29
30
15
31
32
16
33
Photosynthesis….
34
17
Reaction Center
• Chlorophyll a in PS II is called the reaction centre.
• This is the pigment that first become excited and
received an electron from water (convert physical energy
to chemical energy).
• This is where the electron harvest begins. In photosystem
II (PSII), chlorophyll a with the peak absorption at 680 nm
(P680) act as reaction centres.
• All other pigment such as chlorophyll b and carotenoids
are called accessory pigments. This is because other
pigments pass absorbed energy to the chlorophyll a
molecule.
• Chlorophyll a (reaction centre) and the accessory
pigment (harvesting centre) are packed into functional
clusters called photosystems.
• Photosystems are of two types PSI and PSII.
➢ In photosystem I (PSI), chlorophyll a with the maximum
absorption at 700 nm (P700) and
35
• Leaf arrangement – Upper side

outermost layer is epidermis and under
Leaf structure & Functions side is lower epidermis, and the role is
mainly to protect cells.
• Stomata provide the opportunity for

CO2, O2 and water vaporto enter
(influx) and exit (efflux).
• Stomata protects the leaf interior from

drying, CO2 reaching toxic levels and
releasing O2 during photosynthesis.
• Chloroplasts in mesophyll cells and

other tissues engage in capturing light
energy and photosynthesis.
• Vascular tissues provide inputs like

water and nutrients for PS and remove
the end products from leaf and
facilitate chloroplasts to continue with
its PS rates.
36
18
Chloroplast
• CO2 is fixed in the chloroplasts.
• Chlorophylls pick up energy in solar
radiation to do that.
• Chloroplast has a double membrane, like
the mitochondria.
• There are coin-like thylakoids stacked
inside. Each thylakoid has a wall, is made
of the normal cell membrane materials,
i.e., fatty acids and phospholipids as a
double layer, in which protein bodies are
located for specific functions.
• The protein bodies have different
characteristics and functions.
• ATPase enzyme capable of forming ATPs
in protein bodies.
37
Harvesting Light Energy

❑ The light is harvested by pigments chlorophyll
a & b, and Carotenoids in plants.
❑ In addition, Xanthophylls harvest energy in the

yellow wave lengths.
❑ Light energy harvesting step is known as Hill

reaction.
❑ Energy in the green wavelength of solar beam

is not harvested by plants. Only human eye is
sensitive to green wave lengths.
❑ Estimation of energy contents of wave lengths

could be done on
a. Total energy – by pyranometric sensor
b. PAR energy - by Quantum sensor
c. Green range – by photometric sensor
❑ A sensor looks like the one in the diagram.

38
19
Arrangement of chloroplast ❑ Stack of thylakoids are known as
granum.
❑ Each thylakoids has a central cavity,

known as thylakoid space.
❑ The cavity of one thylakoid connects

with that of another through a
channel, (“thylakoid channel”)
which then connect all thylakoids in
a chloroplast.
❑ Outside the granum, cavity is known

as “Stroma” where Calvin cycle
operates.
❑ Thylakoid membrane contains

pigment molecules embedded as
clusters that convert the physical
form of energy of solar radiation to
chemical form.
39
Energy capture….
• Two energy harvesting points known as

photosystem I & II which are specialized protein
complexes.
• Photosystem I (P700) works on NADPH
synthesis, and it receives the electrons from PS II.
• PS II contains both chlorophyll a and b up to 50%

each.
• The key function of the photosystem II is in the
hydrolysis/split of water and ATP synthesis.
• The key part of PS II (p680) is the reaction center,
which is a chlorophyll-a molecule called P680 which
absorbs red light (at 680 nm).
• Each reaction center has multiple light-harvesting
complexes (LHCs), which contains predominantly
chlorophylls a and b, carotenoids, and other
pigments that absorb light at multiple wave lengths.
40
20
Reaction Center…
❑ Each antenna complex has 250 to
400 pigment molecules.
❑ Antenna pigments play a vital role in the

harnessing of photon energy. They function
solely to harvest light energy and transfer
reaction center.
❑ The energy absorb is shuttled by

resonance (a method of energy transfer by
vibration) to the reaction center.
❑ When solar energy strikes on a chlorophyll

molecule, an electron becomes excited and
then shift to a higher orbit/sphere. Ref. Campbell & Reece (2002)
❑ The excited electron is picked up by the

primary electron acceptor, Quinone in the • Missing electron in the chlorophyll
Photosystem II and transfer to other is replaced from water in the
members using oxidation-reduction thylakoid channel.
process.
41
Excitation of electrons in chlorophyll in the presence of solar energy
❑ The solar energy leads to

excitation of an electron
from chlorophyll molecule.
❑ Excited electron will
automatically shift to a
higher orbit from the
nucleus.
❑ This energy is then
released as heat and also
emits as light, known as
fluorescence. The
process is called
fluorescence.
Once the absorbed energy is released, the electron then

returns to the original orbit. Or sometimes, a photon is
released, and which will lead to fluorescence and that can Ref. Campbell & Reece, (2002)
be captured by a digital camera when the event is isolated.
42
21
Arrangement of chloroplast
❑ Electron Transport
System (ETS) is in the
thylakoid membrane.
❑ ETS picks up electrons

from water and provide
to NADP+ forming
NADPH and ATP as
end products.
❑ Hill reaction occurs in

the Thylakoid
membrane of the
chloroplast
43
Energy capture process by plants through chlorophylls ….
❑ Capture of energy by plants occurs

by picking up excited electron and
taking it away from chlorophyll
molecule.
❑ As a photon of solar beam strikes

on chlorophyll molecules, then an
electron will get excited and jump
out of its orbit.
❑ This electron usually returns to

chlorophyll when the energy is
released. But the system quickly
picks up that electron and will not
allow to return.
❑ This process goes on in
chloroplasts during the daytime. Ref. Campbell & Reece (2002)
44
22
Energy capture process by plants through chlorophylls ….
❑ It is vital that the member in the
right place of the team picks up
the excited e-. The compound
that capture the excited electron is
known as the primary electron
acceptor.
❑ After that the electron will continue
to take away through several
compounds.
❑ Then it comes to another site
(PS I or P700), where it has
Chlorophyll a and makes similar
excitation to transfer the e- to
ferredoxin, another primary
electron acceptor.
❑ Finally, the electron will be picked
up by NADP+ by forming NADPH.
45
Photosynthesis….
46
23
Energy Capture Process – Hill Reaction …
Electron Transport System (ETS) located in the thylakoid membrane one can
visualize. ETS picks up electrons from water and provide well organized access to
reduce NADP forming NADPH and ATP as end products.
47
48
24
Hill Reaction.
In the Hill Reaction, excited electron are

first picked up by quinone (Q), which
then transfers electrons Plastoquinone
(PQ). This process occurs in a Redox
manner, which means PQ reduces with
the acceptance of electron and Q
oxidizes.
Interesting part is that when an e-
passes from Q to PQ, there is a proton
(H+) generated on the Thylakoid
channel. This H+ will pass from
Thylakoid space through protein body
having ATPase enzyme, thus Ref. Campbell & Reece (2002)
generating an ATP molecule using ADP.
The e- received by PQ will, in the same way, is transferred to cytochrome

oxidase complex, which contains Fe. Fe+++ then becomes → Fe++ when the
e- is received, and PQ is oxidized. Cytochrome oxidase then will transfer the
electron to plastocyanin (PC) in the same Redox Rxns.
49
Photosynthesis….
• At the PC step, the electron will be

received by PS I. This happens when
light excites an electron in that
chlorophyll molecule and is picked up
by Ferredoxin (Fd – a 1o electron
acceptor).
• That deficit electron of PS I is now
recovered from PC.
• The electron from Fd is transferred
through Fd-NADP+ reductase complex.
• In here Redox reactions are mediated
through FAD → FADH2 (Flavin adenine
di-hydrogenase enzyme).
• The electron reduces NADP+ forming Ref. Campbell & Reece (2002)
NADPH (reduced form of Nicotine
Adenine Di-Phosphate) which is a
product of ETS.
The three end products of ETS are ATP, NADPH and gas O2.
50
25
• Hill reaction completes its role after
forming NADPH+H and ATPs.
• After Hill Reaction, electrons and energy

will support Calvin cycle that is in the
Stroma side of the chloroplast.
51
Continued…..
52
26

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TAT1203/SBS1202

Plant Science/Plant Physiology

2. Biology (sixth edition) (2002)

1. Capture of solar energy by plants

4.1 Photosynthesis - S.L. Ranamukhaarachchi - October Semester 2023

• In 1927, Stephen Hales – A part of the nourishment of plants comes

• In 1930, C. B. Van Neil - There is a similarity between overall

4.1 Photosynthesis - S.L. Ranamukhaarachchi - October Semester 2023

4/3/2024 4.1 Photosynthesis - S.L. Ranamukhaarachchi - October Semester 2023 5

Fate of light energy in brief………..

❑ The figure shows the fate of

❑ Plants capture light energy

❑ ATPs are synthesized using

Ref. Campbell & Reece (2002)

4.1 Photosynthesis - S.L. Ranamukhaarachchi - October Semester 2023

• Photosynthesis – all in the

• Red and far-red have effects

• Light duration also influences

TAT1203 Plant Science - S.L. Ranamukhaarachchi - October Semester 2023

TAT1203 Plant Science - S.L. Ranamukhaarachchi - October Semester 2023

Energy in Solar Radiation

h – Plank’s constant = 1.58 x 10-34 Cal Sec

c – Velocity of light = 3.0 x 1010 cm sec-1

TAT401 Controlled Environment Agriculture - S.L. Ranamukhaarachchi - October Semester 2023 12

Blue = 1.58 x 10-34 cal sec x 3.0 x 1010 cm sec-1

= 6.02 x 1023 photons/Einstein

Red = 1.58 x 10-34 cal sec x 3.0 x 1010 cm sec-1

Einstein is a mole of photon.

TAT401 Controlled Environment Agriculture - S.L. Ranamukhaarachchi - October Semester 2023 14

Measurement of Light Energy

• When light energy reacts with other substances, it work

TAT401 Controlled Environment Agriculture - S.L. Ranamukhaarachchi - October Semester 2023

• PPF – photosynthetic photon flux.

TAT401 Controlled Environment Agriculture - S.L. Ranamukhaarachchi - October Semester 2023

Pigments and Energy

4.1 Photosynthesis - S.L. Ranamukhaarachchi - October Semester 2023

Remember, green wave

4.1 Photosynthesis - S.L. Ranamukhaarachchi - October Semester 2023

Absorption of light energy

4.1 Photosynthesis - S.L. Ranamukhaarachchi - October Semester 2023

• Chlorophyll a and b are

• The difference between

4.1 Photosynthesis - S.L. Ranamukhaarachchi - October Semester 2023

4.1 Photosynthesis - S.L. Ranamukhaarachchi - October Semester 2023

Wavelengths of solar beam and effectiveness in

• The upper diagram shows the absorption

• Chlorophyll a and b absorb usually blue wave

• Meanwhile, orange wave lengths are absorbed

• In the overall absorption pattern, blue and red

4.1 Photosynthesis - S.L. Ranamukhaarachchi - October Semester 2023

• Chlorophyll a - Blue-violet range – reflects green

4.1 Photosynthesis - S.L. Ranamukhaarachchi - October Semester 2023

❑ Leaf – the main part that

❑ However, green materials

❑ Broadleaf and grass types

4.1 Photosynthesis - S.L. Ranamukhaarachchi - October Semester 2023

4.1 Photosynthesis - S.L. Ranamukhaarachchi - October Semester 2023

How radiation behaves in nature

Three types of actions take

The diagram also shows

4.1 Photosynthesis - S.L. Ranamukhaarachchi - October Semester 2023

4.1 Photosynthesis - S.L. Ranamukhaarachchi - October Semester 2023

4.1 Photosynthesis - S.L. Ranamukhaarachchi - October Semester 2023