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Lecture 2: LIGHT
REACTION

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COMPETENCIES
Upon mastering the materials of this lecture,
students would be able to explain
1. The characteristics of light as a source of
energy in the process of photosynthesis
2. The process of NADPH formation in the
conversion of radiation energy to be chemical
energy
3. The process of ATP formation in the conversion
of radiation energy to be chemical energy

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62 Slides

LECTURE OUTLINE
What is photosynthesis?
The process of converting solar energy
into chemical energy.
energy.
The process of CO2 reduction into
carbohydrates (sugars) at the expense of
NADPH & ATP
Responsible for removal of ~ 200 billion
tons of C from the atmosphere yearly.
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Primary
electron
acceptor
Energy
to make

Primary
electron
acceptor

NADP

Photosynthesis has two major phases:

1. The absorbance of light and production of chemical forms of
energy (light reactions)
2. The fixation and reduction of carbon and other oxidized
molecules (dark reactions)

2
Light

Light
Primary
electron
acceptor
Reactioncenter
chlorophyll

PS II

PS I

NADPH-producing
photosystem

1
Water-splitting
photosystem
2 H + 1/2

How the Light Reactions Generate NADPH and ATP

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Solar energy is the ultimate source of energy for

life on earth
Solar energy is created at the core of the sun
when hydrogen atoms are fused into helium by
nuclear fusion .

Temperatures of the sun are

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The solar energy is then transmitted in the form of

electromagnetic radiation

about 15,000,0000K at the core,

and
about 5,8000K at the photosphere
(radiative surface of the sun)

Radiation is the transfer of energy through some

material or through space in the form of
electromagnetic waves
Electromagnetic waves are the self-propagating, mutual
oscillation of electric and magnetic fields.
Electromagnetic waves move electromagnetic energy
through space (either empty or filled with transparent
matter)

Most of the electromagnetic radiation emitted

from the sun's surface lies in the visible band
centered at 500 nm

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Longest wavelengths
(Least energetic photons)

Shortest wavelengths
(Most energetic photons)

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The Electromagnetic Spectrum

2.1 Stefan
StefanBoltzmann Law

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This law states that the power emitted per

unit area of the surface of a black body is
directly proportional to the fourth power of
its absolute temperature. That is
R=
R=
T
T4

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Apply StefanStefan-Boltzmann Law To Sun and Earth

R=
R=
T
T4

Sun (6000
( 6000 0K)
RS = (5.67 x 10-8 W/m2 K4) * (5800 0K)4
= 64,164,532 W/m2

Earth (300
(300 0K)
RE = (5.67 x 10-8 W/m2 K4) * (300 0K)4
= 459 W/m2

R = radiation flux (W.m-2 = J.m-2.s-1)

= emissivity (0
(01)
1)
= StefanStefan-Boltzmann constant (5,67032 x 10-8 W.m-2.K-4)
T = absolute temperture (273 + 0C).
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The Sun appears to have been active for 4.6

billion years and has enough fuel to go on for
another five billion years or so.
At the end of its life, the Sun will start to fuse
helium into heavier elements and begin to swell
up, ultimately growing so large that it will
swallow the Earth.
Earth.
After a billion years as a red giant, it will
suddenly collapse into a white dwarf -- the final
end product of a star like ours. It may take a
trillion years to cool off completely.

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Sun emits about 160,000 times more radiation

per unit area than the Earth because Suns
temperature is about 20 times higher than
Earths temperature
temperature 600/300 = 20

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Radiation emitted by a human body

Inverse Square Law

The amount of radiation passing through a specific area is
inversely proportional to the square of the distance of that
area from the energy source.

The net power radiated is the difference between

the power emitted and the power absorbed:
P net = P emit P absorb.
Applying the Stefan
StefanBoltzmann law,

I = E(4
E(4R2)/(
)/(4
4r2)
I = Irradiance at the surface of
the outer sphere
E = Irradiance at the surface of
the object (Sun)
R = 6.96 x 105 km (Radius of the Sun)
r = 1.5 x 108 km (Average SunSun-Earth Distance)
I = 64,164,532 W/m2 x(6.96 x 105 )2 /(1.5 x 108 )2
I = 1382 W/m2 (The generally accepted solar constant of
1368 W/m2 is a satellite measured yearly average)
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R = (T4-T04)

A = the total surface area of an adult is about 2 m,

m,
= the
the midmid- and farfar-infrared emissivity of skin and most
clothing is near unity, as it is for most nonmetallic
surfaces.
T = skin temperature is about 33
33C, but clothing reduces
the surface temperature to about 28 C when the
ambient temperature is 20 C.
T0 = the ambient temperature is about 250C in Malang
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WHY ARE
PLANTS
PLA
NTS GRE
GREEN?
EN?

Hence, the net radiative heat loss is about

Pemit =

0.97* 5.67108 W m2 K4 *2 m2(273+28)4

= 902.92 W.m-2 or J.s-1
Pabsorbed =
0.97* 5.67108 W m2 K4 *2 m2(273+27)4
= 890.98 W.m-2 or J.s-1
P net = 902.92-890.98 = 11.94 J.s-1
11.94 x 24 hours x 60 minutes x 60 seconds =
1,031,556.8 J/day

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Why are plants green?

Transmitted light

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2.2 Light

LIGHT AS A WAVE
Wavelength (l
(l ) the distance between crests (or
troughs) of a wave
Frequency (v
(v) the number of crests (or troughs) that
pass by each second.
Speed (c
(c ) the rate at which a crest (or trough) moves
(3.105 km/s).
Crest
l
Maxwell calculated the speed of
propagation of electromagnetic
Trough
waves and found:

Light is an electromagnetic wave

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What is the electromagnetic wave?

It is electricity and magnetism moving through the
space
Light was known to
be a wave
After producing
electromagnetic
waves of other
frequencies, it was
known to be an
electromagnetic wave
as well.

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This is the speed of light in a vacuum.

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2.3 Quantum Theory

Planck's Theory

Light as particles

Light comes in quanta of energy called photons little

bullets of energy.
A photon is a quantum or irreducible quantity of
electromagnetic radiation.

By the 1900's the wave model was accepted by scientist

as how light moved.
Ideas of quantum theory were developed when
classical physics (the wave model) could not explain
several physical phenomena observed in beginning of
the 20th century
light until further heating, then it will glow red, yellow then
"white" hot.
It also did not explain colors given off by various elements as
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they burn

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Energy cannot be absorbed or emitted

unless it is a complete packet.
Planck's theory states that atoms can
only absorb or release energy in fixed
quantum units

The amounts of energy an object emits or absorbs are

called quantum (quanta plural)
Related the Frequency of the radiation to the amount of
energy.
E = h = hc/l
Frequency (v) = c/
h = 6.6262 x 10-34 J-s (joule-seconds)
c = speed of light (3x108 [m/s)
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Visible radiation: visible to our eyes (wavelength

:0.4x10-6 - 0.7x10-6 m)

Dengan memasukkan harga

harga--harga konstanta
konstanta,,
maka

Red = 0.65 mm, Orange = 0.60 mm, Yellow = 0.55 mm,

Green = 0.50 mm, Blue = 0.45 mm & Violet = 0.40 mm

Cahaya dan PAR

Tanaman dalam proses fotosintesis hanya dapat

memanfaatkan pancaran radiasi matahari yang terletak
pada batas panjang gelombang 400 - 700 nm
Radiasi pada batas panjang gelombang 400 - 700 nm
disebut PAR (photosynthetically active radiation) atau
cahaya nampak (visible radiation)
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19,878.1017 J

dimana dalam satuan nano meter (nm)

Sebagai contoh
contoh,, kandungan energi cahaya
merah ( = 680 nm) adalah

1 J (Joule) = 107 erg; 1 c (cal) = 4,2 J; 1 eV

= 1,6.10-12 erg
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Summary of Photons

A.h.c

where
A is Avogadro's number (= 6.02 x 1023 photons/mol)
h is Planck's constant (= 6.62 x 10 -34J s per photon)
c is the speed of light (= 3 x 108 m.s-1).

A 6.02.10

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photons mol h 6 .62.10 34 J.s photon c 3.10 8 m s

550 .10 9 m

E = 217376.7 J.s
a mol of blue light (l = 400 nm) = 298893.0 J.s
a mol of red light (l = 700 nm) = 170796.0 J.s
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Representation of a Photon

To describe interactions of light with matter, one

generally has to appeal to the particle (quantum)
description of light
A single photon has an energy given by
E = hc/l
where
h = Plancks constant = 6.6x10-34 [J s]
c = speed of light
= 3x108 [m/s]
l = wavelength of the light (in [m])
Photons also carry momentum. The momentum is
related to the energy by:
p = E/c
E/c = h/l
h/l
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For instance,
the energy of "green light" (= 550 nm) is:
E

Photons can be treated as

packets of light
light which behave
as a particle.

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The reduction of CO2 to be carbohydrate

through photosynthesis requires energy
(NADPH & ATP)
Photosynthesis can be divided into two
reactions
Light Reaction

2.
3.

Light Absorption
Pigments
Electron Excitation

Generation of NADPH2
Generation of ATP

Light Reaction
1.

4.
5.

Dark Reaction

Fluorescence
Phosphorescence

Electron transfer & Synthesis NADPH

Proton exchange & Synthesis ATP

Diffusion of CO2
Reduction of CO2

C3, C4 & CAM

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Photosynthesis is the process by which autotrophic

organisms use light energy to make sugar and
oxygen gas from carbon dioxide and water

This is an over simplification approach as H2O never

meets CO2 directly in the photosynthesis

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Chloroplasts

In most plants, photosynthesis occurs primarily in

the leaves, in the chloroplasts

A chloroplast contains:

The leaves have the most chloroplasts

The green color comes from chlorophyll in the
chloroplasts
The pigments absorb light energy
Stroma (a fluid)
Grana (stacks of thylakoids
thylakoids))

The thylakoids contain

chlorophyll

Chlorophyll is the green pigment

that captures light for
photosynthesis
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The location and structure of chloroplasts

Chloroplast
LEAF CROSS SECTION

MESOPHYLL CELL

1.

LEAF

2.

Mesophyll

CHLOROPLAST

Intermembrane space

3.
Outer
membrane

Granum
Grana
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Stroma

4.

Light Absorption
Electron Exitation
Fluorescence
Phosphorescence.
Electron Transfer
NADPH Synthesis

Inner
membrane
Stroma

Thylakoid

Thylakoid
compartment

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LIGHT ABSORBTION
AND TRANSFER
TO THE REACTION
CENTERS

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LIGHT ABSORBTION
AND TRANSFER
TO THE REACTION
CENTERS

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Resonance Energy Transfer - Radiationless

Pigment = a light absorbing molecule
Associated with the thylakoid
membranes
Chlorophyll

Excited state

e-

e-

e-

e-

hv

e-

e-

e-

Xanthophylls
Carotenoids

e-

Ground state

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Chl a and Chl b (Chl

(Chl c in some algae)

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-carotene
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Chl a has a methyl group (CH3)

Chl b has a carbonyl group (CHO)

Porphyrin ring delocalized ePhytol

Phytol tail

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Absorpsi foton mengakibatkan pengaturan

elektron intramolekul pada pusat reaksi yang
diikuti dengan tranfer elektron antar molekul

Pada mulanya, elektron

khlorofil pada pusat
reaksi tereksitasi pada
orbit yang menjauhi inti
atom dan molekul dengan
absorpsi foton langsung
atau lebih mungkin
melalui transfer energi
foton dari antena
pigmen

Two types of
photosystems
cooperate in the
light reactions
ATP
mill

Water-splitting
photosystem
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NADPH-producing
photosystem

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Dua orbit dipertimbangkan sebagai

tempat eksitasi elektron yaitu orbit I dan
II (eksitasi
(eksitasi I & II)
Elektron yang tereksitasi tidak terikat
kuat pada molekul khlorofil dan mudah
ditransfer ke molekul lain disekitarnya
disekitarnya..
Pusat reaksi yang tereksitasi adalah
reduktor yang kuat
kuat,, dan bahkan cukup
kuat untuk mereduksi molekul lain yang
tidak siap menerima elektron.

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Partial energy diagram for a photoluminescent system.

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Fluorescence: Emisi cahaya dari molekul yang

Fluorescence:
sedang diiradiasi sebagai akibat dari penurunan
elektron dari orbait 1 ke orbit dasar
dasar.. Proses ini
tidak tergantung suhu dan berlangsung cepat
(lifetime 10-8 detik
detik).
). Panjang gelombang lebih
besar dari panjang gelombang yang diabsorpsi
(chlorophyll a mengabsorpsi cahaya pada 430 &
630 nm, dan mengemisi cahaya pada 668 nm).
Phosphorescence:: Emisi cahaya dari molekul
Phosphorescence
sebagai akibat penurunan elektron dari triplet
state ke orbit dasar
dasar.. Cahaya yang dihasilkan
berlangsung relatif perlahan (10-4 2 detik
detik),
),
dan panjang gelombang relatif sangat panjang

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Excitation of chlorophyll
in a chloroplast

Red light absorbed by photosystem II

(PSII) produces a strong oxidant and a
weak reductant
reductant..
Far
Far--red light absorbed by photosystem I
(PSI) produces a weak oxidant and a
strong reductant
reductant..
The strong oxidant generated by PSII
oxidizes water,
water, while the strong
reductant produced by PSI reduces
NADP..
NADP

Loss of energy due to heat

causes the photons of light to be
less energetic.
Less energy translates into
longer wavelength.

Excited
state

Light

Heat

Energy = (Plancks constant) x

(velocity of light)/(wavelength of
light)
Transition toward the red end of
the visible spectrum.

Light
(fluorescence)
Photon
Ground
state
Chlorophyll
molecule

(a) Absorption of a photon

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(b) fluorescence of isolated chlorophyll in solution

1.
2.
3.
4.
5.

6.
7.

H 2O
Z (PSII)
P 680
680** (PSII reaction
center chlorophyll)
Pheo ( pheophytin
pheophytin))
Q A and Q B
( plastoquinone
acceptors)
Cytochrome b.
b.--f
complex
PC ( plastocyanin
plastocyanin))

8.
9.
10.
11.

12.
13.

14.
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P700+ (PSI reaction

center chlorophyll)
A 0 (chlorophyll ?)
A 1 ( quinone
quinone?)
?)
FeSx,, FeSB, & FeSA
FeSx
( membrane
membrane--bound
iron--sulfur proteins
iron
proteins))
Fd ( soluble ferredoxin
ferredoxin))
Fp ( flavoprotein
ferredoxin--NADP
ferredoxin
reductase))
reductase
NADP
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Noncyclic Photophosphorylation

Plants produce O2 gas by splitting H2O

Photosystem II regains electrons by splitting water, leaving

O2 gas as a byby-product

Primary
electron acceptor

The O2 liberated by photosynthesis is made

from the oxygen in water (H+ and e-)

Primary
electron acceptor

Photons

Energy for
synthesis of
PHOTOSYSTEM I
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PHOTOSYSTEM II

53

by chemiosmosis

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1.
The red X indicates that protons
do not directly pass through the
cytochrome complex.

2.
X

3.

4.
Protons cross the membrane via
oxidation and reduction of quinones
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Z is a tyrosine side chain on the reaction

center protein D1. Electrons are extracted
from water (H2O) by the oxygen
oxygen--evolving
complex and rereduce Z+.
On the oxidizing side of PSII (to the left of
the arrow joining P680 with P680*), P680+
is rereduced by Z, the immediate donor to
PSII.
The excited PSII reaction center chlorophyll,
(P680*) transfers an electron to pheophytin
(Pheo
Pheo).
).
On the reducing side of PSII (to the right of
the arrow joining P680 with P680*), the
pheophytin transfers electrons to the
plastoquinone acceptors QA and QB.

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5.

6.

7.

8.

9.

The cytochrome b.
b.--f complex transfers
electrons to plastocyanin (PC), which in turn
+
reduces P700 .
The b,
b,--f complex contains a Rieske iron
iron--sulfur
protein (FeSR
(FeSR),
), two b-type cytochromes (cyt
b), and cytochrome f (cyt f).
The acceptor of electrons from P700* (A0) is
thought to be a chlorophyll, and the next
acceptor (A1) may be a quinone
quinone..
A series of membrane
membrane--bound iron
iron--sulfur
proteins (FeSx
(FeSx,, FeSB, and FeSA) transfer
electrons to soluble ferredoxin (Fd
Fd).
).
The flavoprotein ferredoxin
ferredoxin--NADP reductase
(Fp
Fp)) serves to reduce NADP, which is used in
the Calvin cycle to reduce CO2.

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10.
11.

12.

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13. The transfer of electron occurs

from A0
to A1 (quinone
(quinone),
), FeSx
FeSx,, FeSB, & FeSA
(membrane
membrane--bound iron
iron--sulfur proteins
proteins),
),
Fd (soluble ferredoxin ), Fp
(flavoprotein ferredoxin
ferredoxin--NADP
reductase)) and finally to NADP+
reductase
14. It was found antagonistic effects of light
on cytochrome oxidation.

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The dashed line indicates cyclic electron flow around

PSI.
PSII produces electrons that reduce the cytochrome
b.--f complex, while PSI produces an oxidant that
b.
oxidizes the cytochrome b.
b.--f complex.
P680 and P700 refer to the wavelengths of maximum
absorption of the reaction center chlorophylls in PSII
and PSI
The electron of PSI is then excited upon absorption of
radiation energy and transfer to A0 (chlorophyll)

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H2O Z (PSII)
(PSII)P680
680** (PSII reaction center
chlorophyll) Pheo (pheophytin
pheophytin))QA and
QB (plastoquinone acceptors)
acceptors)
Cytochrome
b.--f complex PC (plastocyanin
b.
plastocyanin))P700+
(PSI reaction center chlorophyll) A0
(chlorophyll ?)
?)A1 (quinone
quinone?)
?) FeSx
FeSx,,
FeSB, & FeSA (membrane
membrane--bound iron
iron--sulfur
proteins)) Fd (soluble ferredoxin )Fp
proteins
(flavoprotein ferredoxin
ferredoxin--NADP
reductase))NADP
reductase

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1.

2.
3.
4.

Eksitasi 1 mol e pada setiap pusat reaksi (PSII &

PSI) membutuhkan
membutuhkan1
1 kuanta cahaya
cahaya.. Reduksi 1
mol NADP 1 mol NADPH membutuhkan 2 mol
e
Berapa kuanta cahaya dibutuhkan untuk
pembentukan 1 mol NADPH ?
Berapa NADPH dihasil dari hasil fotolisis air ?
Tingkat Cahaya di Malang sekitar 1 mmol.s-1,
berapa NADPH yang dihasilkan dengan tingkat
cahaya demikian ?

1.
2.
3.
4.

5.

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Where does the light reaction happen ?

What is the function of water in
photosynthesis ?
What is the event to happen after the light
interception by pigments
What is the first molecule receiving
electrons from the pigments (chlorophyll)
excited at PS I
What is the first molecule receiving
electrons from the pigments (chlorophyll)
excited at PS II

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