Photosynthesis -1

Raj Kumar
Assistant Professor in Botany
Government College
Hisar
Topics to be Studied
 Nature of light
 Historical update
 Pigments of photosynthesis
 Distribution of photosystems
 Light reactions
o Noncyclic photophosphorylation
o Cyclic photophosphorylation
o Pseudocyclic photophosphorylation
•How light energy is first changed to
electrical energy than to chemical
energy (NADPH and ATP)?

•How food is produced in plants?

•Leaf acts as a photosynthetic

machine.
Importance
All life is bottled sunshine’
The use of energy from sunlight by photosynthesis is the
very basis of life on earth.
Food for autotrophs & all other heterotrophs
Biomass
Fossil Fuels
All Oxygen of the earthproduced.
Research for photosynthesis in vitro conditions
Some herbicides function by blocking the light
reaction.
Carotenoids protect chloroplasts from photo-
oxidative damage
 Photosynthetic yield
“If a year's yield of photosynthesis were amassed in
the form of sugar cane, it would form a heap over
two miles high and with a base 43 square miles.”
-G. E. Fogge
•If all of this sugar cane were converted into sugar cubes
(0.5 inch on a side) and stacked end to end, the sugar
cubes would extend 1.6 × 1010 miles, or to the planet
Pluto.
•The photosynthesis by green plant on land is 50 x 109
tons/ year and by ocean plants 180 x 109 tons/ year.
•Plants growing in oceans and the other fresh water
algae account for 90 per cent of total photosynthesis.
Photosynthesis
Anabolic (small molecules combined)
Endergonic (stores energy)
Carbon dioxide (CO2) requiring process that
uses light energy (photons) and water (H2O) to
produce organic macromolecules (glucose).

6CO2 + 6H2O  C6H12O6 + 6O2

SUN
photons

glucose
6
Structure of leaf
Photosynthetic pigments
 Located in chloroplast’s
thylakoidal membranes.
 Three types

1. Chlorophylls
2. Carotenoids
3. Phycobillins

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publishing as Benjamin Cummings
Chlorophylls
Chlorophylls (Chl a, b, c, d, e) and
bacteriochl. (Bchl a, b , c, d,
bacterioviridin) are found in
photosynthetic organisms.
But all organisms contain a mixture of more
than one kind of pigments.
Chlorophyll a and b are abundant in green
algae and higher plants.
Several spectrally distinct forms of Chl a
shown by absorption peaks exist e.g. Chl
a670–673, Chl a680–683, P700, P680
Chlorophyll Molecules
Located in the thylakoid membranes
Chlorophyll have Mg+ in the center
Chlorophyll pigments harvest energy (photons)
by absorbing certain wavelengths (blue-420 nm
and red-660 nm are most important)
Plants are green because the green wavelength
is reflected, not absorbed.
absorbed

12
All chl. consist of a
tetrapyrolle ring having
conjugated double bonds
and a phytol tail (long chain
alcohol).
The centre of the
tetrapyrolle ring has a Mg.
co-ordinatively held to N
atoms of pyrolle rings.
Side chains attached to
pyrolle ring vary in different
chl.
Phytol chain imparts the
molecule lipid solubility &
water insolubility.
Chl b is minor chl in higher
plants and green algae
absorbtion peaks 455, 640
 •Chl a has a methyl
group

•Chl b has a carbonyl

group

Porphyrin ring
delocalized e-

Phytol tail
Carotenoids:
Supplementary light receptors
Yellow, red or purple pigments.
β-carotene (m.imp) is a red
orange isoprenoid compound;
precursor of vit A in animals
Xanthophyll is yellow.
Abs 400-500 nm.
Protect cht against photo-
oxidative damage.
Xanthophylls like lutein &
zeaxanthin are oxygen
derivatives of carotenes containg
1-8 oxygen atoms.
http://www.plantstress.com/Articles/O
xidative%20Stress_files/IMG00029.gif
Phycobilins:
Found in phycobilisomes.
Linear tetrapyroles that have extended polyene
system found in chlorophylls but not their
cyclic structure or central magnesium.
e.g. phycoerythrin (490, 546, 570 nm) and
phycocyanin (608 nm).
Why are plants green?
h t
lig
c ted Transmitted light
efl e
R
 THE COLOR OF LIGHT SEEN IS THE
COLOR NOT ABSORBED

Chloroplasts absorb
light energy and
Reflected
convert it to Light light
chemical energy

Absorbed
light

Transmitted Chloroplast
light
Wavelength of Light (nm)

400 500 600 700

Short wave Long wave

(more energy) (less energy)

23
Absorption of Light by
Chlorophyll
Chlorophyll absorbs blue-violet & red light best

Absorption

violet blue green yellow orange red

wavelength
24
Question:

During the fall,

what causes the
leaves to change
colors?
25
Fall Colors
In addition to the chlorophyll pigments, there
are other pigments present
During the fall, the green chlorophyll
pigments are greatly reduced revealing the
other pigments
Carotenoids are pigments that are either
red, orange, or yellow

26
What is Photosynthesis
The overall process by which plants, algae
and prokaryotes use light energy to
synthesize organic compounds is called
photosynthesis
It uses carbon dioxide (CO2), water (H2O),
minerals and light to produce carbohydrates
(food) and oxygen (for breathing).
Summary of photosynthesis…
sunlight
CO2 + H2O C6H12O6 + O2
Photosynthesis Overview

Light reactions (occur in thyakoids) convert

light energy to NADPH and ATP.

The Dark reactions or Calvin Cycle (occur

in stroma) uses NADPH & ATP to fix CO2
to make C6H12O6 (sugars).
 Light show both particulate
Nature of Light and wave nature.
A simple equation relates the
wavelength (λ), frequency (ν is
no. of wave crests at a given
time) and speed or velocity (c
= 3.0 x 108 ms-1) of the wave.
c = λν
Light is a transverse (side to
side) electromagnetic wave in
which both electric and
magnetic fields oscillate
perpendicularly to the
direction of the propagation
http://www.pas.rochester.edu/~afrank/ of the wave and at 90o with
A105/LectureV/FG03_006_PCT respect to each other.
Light is also a particle which is called photon having
an amount of energy called quantum (pl: quanta).
The energy (E) of a photon is shown by Planck’s law E
= hν
h = Planck’s constant (6.626 x 10-34 Js); ν = frequency of
light.
Sunlight is like a rain of photons of different
frequencies.
Our eyes are sensitive to small range i.e. the visible
light.
Wavelength (λ) and frequency are inversely
proportional.
Different wavelengths of visible light are seen by
the human eye as different colors.

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publishing as Benjamin Cummings
Gamma Infrared &
rays X-rays UV Microwaves Radio waves

Visible light

Wavelength (nm)
Absorption spectrum
relates the amount of light
energy taken up or absorbed
by a molecule or a substance
w.r.t. the wavelength of light.
The absorption spectra of
Chl a and Chl b show that
these pigments chiefly
absorb blue (480nm) and red
regions (662 nm) of
spectrum.
Chlorophyll absorbs blue-
violet & red light best
The exact position of peaks
depend upon the solvent
used for extraction.
The transmitted and
reflected light are vastly rich
in green (500 -600 nm) gives
leaves their green colour.
Action spectrum shows
the magnitude of
photosyn (or biological
process) to light (i.e.
different wavelengths).
The magnitude of
photosyn is measured
and plotted by amount
of action (e.g. CO2
fixation, O2 production,
NADP+ reduction,
change in biomass etc.)
2004 Pearson Education, Inc.
publishing as Benjamin Cummings
Historical update
Aristotle & Theophrastus 320 B.C. plants derive
nutrition directly from soil.
Van Helmont (1648): all food for green plants
comes from water.
Stephen Hales (1727): green plants require
sunlight for their nutrition.
Joseph Pristley (1772): green plants purify foul
air (de phlogiston) produced by burning of candle
and covert it into pure air (by producing phlogiston
i.e. oxygen)
Ingen Housz (1779): plants purify the noxious
air only in the presence of light.
Lavosier (1783): identified pure air produced by
plants as oxygen and foul air as CO2.
Historical update
Nicolas de Saussare (1804): water was used in
photosynthesis
Dutrochet (1837): Chlorophyll is important for
photosynthesis.
Liebig (1845): main source of carbon in plants is
CO2.
Van Meyer (1845): green plants transform light
energy into chemical energy of organic matter.
J.V. Sachs (1862): First visible product is the
starch and chl. is present in chloroplast.
T.W. Englemann (1888): chloroplast is seat of
photosynthesis & plotted the 1st action spectrum
using spirogyra.
 Cont… Historical update
Blackmann (1905): proposed two steps i.e. light & dark
reactions. Proposed law of limiting factors.
Van Neil (1931): some bacteria use H2S instead of H2O in
photosyn. Photosynthetic rxn are redox reactions.
 CO2 + 2 H2S + light energy → CH2O + H2O + 2S
Emerson and Arnold (1932): by flashing light (10-5 s)
experiments at different temp. showed the existence of
light and dark reactions.
Robert Hill (1937): Cht cause photo-oxidation of water
to produce a hydrogen donor.
Ruben, Hassid and Kamen (1941): using labelled
oxygen showed oxygen evolved comes from water and
not from CO2.
Arnon, Allen and Whatley (1954) used 14CO2 to show
that CO2 is fixed by isolated chloroplast.
Emerson et al (1957) Emerson enhancement effect
and red drop
 Historical update
Melvin Calvin (1954) Calvin cycle or dark reaction.
Nobel Prize for Chemistry in 1961
Hill & Bendell (1960): proposed Z scheme & suggested
that two photosystems operate in series.
Arnon (1961): photophosphorylation.
Peter Mitchell (1961) chemiosmotic coupling
hypothesis. Nobel Prize in Chemistry in 1978
Hatch and Slack (1966) C4 cycle in sugarcane
Johann Diesenhofer, Robert Huber and Hartmut
Michel (1985) West Germany crystallized
photosynthetic reaction centre of bacterium
Rhizobacter virdis. Nobel prize in 1988.
Rudolph A. Marcus (1985): theory of electron
transfer reaction in chemical systems. Nobel Prize in
Chemistry in 1992 for his contributions to the. The
electron transfer reactions are rapid (as fast as a few
picoseconds) and highly specific.
•Carotenoids, chl b & chla
together serve as an light
harvesting antenna, collecting
light and transferring its
energy to the rxn centre (P700/
P680),
where chemical reactions
store some of the energy by
transferring electrons from a
chl pigment to an e- acceptor.
•An electron donor then
reduces the chl again.
•The transfer of energy in the
antenna is a purely physical
phenomenon and involves no
chemical changes
FIGURE 19-52 Organization of photosystems in the thylakoid membrane
Mechanism of light reactions
• The chemical rxns involved in electron
transfer during photosynthesis are three
types
 Non cyclic photophosphorylation or Z
scheme
 Cyclic photophosphorylation
 Pseudocyclic photophosphorylation
• There are two photosystems or pigment
systems I & II and Cytchrome b6f complex
embedded in thylakoidal membranes are
involved in these reactions.
Two photosystems
Oxygen evolving organisms have two photosystems
that operate in series
PS I absorbs far red light of wavelengths greater
than 680 nm; PS II absorb red light of 680 nm.
The two PS each have antenna pigments and
reaction centres.
The two PS are linked by an electron transport
chain.
Located along with their respective antenna
pigments in thylakoidal membranes.
The reaction centers, the antenna pigment-protein
complexes and most of the electron transport
enzymes are all integral membrane proteins
Arrangement of the thykaloid membrane
•Folded regions & appressed regions called grana are rich in PS II.
LHCII and Cyt b6f complex also present.
•Stroma lamellae (non appressed regions) are rich in PS I. Cyt b6f-
complex & ATPsynthase also present.
Photosystem I (PS I)
PS I is a large multi subunit
complex.
Light harvesting complex
or antenna of carotenoids
and chl’s and rxn centre
with special chl a molecule
P700 bound to two
proteins Psa A & Psa B with
m. mass 66 to 70 Kda.
Total no. of chl. 200.
The early e- acceptors are
A0 ( a chl. A molecule), A1
(a quinone called
phylloquinone or vit K1).
Other are iron –sulfur
proteins or Fe-S centers :
FeSx, FeSA and FeSB. http://upload.wikimedia.org/wikipedia/comm
ons/8/81/P_700.jpg
FeSX is the part of the
P700 binding proteins.
FeSA and FeSB present on
8 Kda protein that is part
of PS I reaction center
complex.
Electrons are transferred
through centers A & B to
Fd (a small water solube
Fe-S protein). The
membrane associated
flavoprotein Fd-NADP
reductase (FNR) reduces
NADP+ to NADPH.
PS I also supply the
reductants to reduce
nitrate and regulation of
some of carbon fixation
enzymes.
http://upload.wikimedia.org/wikipedia/comm
ons/8/81/P_700.jpg
Photosystem II
It is an integral memb.
complex containing P680
rxn center.
It contains 20 proteins.
Two proteins D1 and D2
binds e- carriers P680,
pheophytin and
plastoquinone.
Other proteins such as CP43
and CP47 binds Chl a
antenna pigments
Still other PSII proteins
(33,23 & 17 Kda) are involved
in water oxidation.
Function of remaining low http://www.biochem.arizona.edu/classes/bioc462/462b/Miesf
eld/photo_phos/text_figures/476_figure-19-54c.jpg
m. mass subunits including
Cyt b559 remains unknown. •PS II function as light
dependent water
plastoquinone oxidoreductase
FIGURE Photosystem II of the cyanobacterium Synechococcus elongates.
Cytochrome b6f complex
Large multisubunit protein.
It contains two cytochromes (Cyt b
and Cyt f)
It has a Rieske iron-sulfur protein
(FeSR) in which two iron atoms are
attached to two sulfur atoms.
It also contains an additional heme,
a chl and a carotenoid whose
functions are yet to be resolved.
The e- and protons flow through
complex by a Q cycle.
For a pair of e- four H+ are pumped
from the stroma in the thylakoid
lumen.
It mediates transfer of e- from
Plastoquinone to plastocyanin. http://www.biochem.arizona.edu/classes/bioc46
2/462b/Miesfeld/photo_phos/text_figures/476_
figure-19-54c.jpg
 Non cyclic Photophosphorylation or Z scheme
In this case the electron expelled by the excited
chlorophyll molecule (P680) never comes back.
In the picture the redox carriers are placed at their mid
point redox potentials (at pH 7).
The excited P680* transfer an electron to pheophytin.
P680 oxidized by light is re-reduced by Yz (an tyrosine
residue in D1 protein of PS II) which in turn receives e-
from oxidation of water by OEC.
Pheo transfer e- to QA and QB (plastoquinones).
Cyt b6f complex trasfer e- to PC soluble protein.
Plstocyanin (PC) reduces oxidized P700 (P700+)of PS I.
A0 (chl) accepts e- from P700+. Next acceptor is A1 (quinone)
A series of memb bound iron sulfur proteins (FeSX, FeSA
and FeSB transfer e- to soluble ferredoxin (fd).
The soluble flavoprotein ferredoxin-NADP reductase
(FNR) reduces NADP+ to NADPH (used in Calvin cycle).
Non cyclic Photophosphorylation or Z scheme

http://upload.wikimedia.org/wikipedia/commons/thumb/1/18/T
hylakoid_membrane.png/300px-Thylakoid_membrane.png
Oxidation of water
Water is oxidized by oxygen
evolving complex.
Water is very stable.
OEC is source of all atmospheric
O2.
OEC exist in five different
oxidation states (S0, S1, S2, S3, S4)
Mn complex0 to Mn complex4+
S4 state is strong oxidant that
evolves O2.
Yz (a tyrosin residue forms the
link b/n P680+ and OEC. http://upload.wikimedia.org/wikipedia/en/
e/e7/Oxygen_evolving_complex.png
Cyclic photophosphorylation
 Involves only PS I.
 e- passed from P700 to ferredoxin do
not continue to NADP+ but move
back through the cytochrome b6f
complex to PC.
 PC donates e- to P700 the
illumination of which promotes e-
transfer to ferredoxin.
 Cyclic e- flow is not accompanied
by net formation of NADPH or the
evolution of O2.
 It is accompanied by proton
pumping from stroma to thylakoid
lumen creating proton gradient
leading to phosphorylation of ADP
to ATP referred as cyclic
photophosphorylation.
 Conditions when it occur: When
the plant cell has sufficient NADPH
but requires additional ATP for
other metabolic processes.
 Pseudocyclic
Mehler reaction or
 Pseudocyclic photophosphorylation
photophosphorylation
reaction
 From Fd e- are passed to O2
forming H2O2
 H2O2 disrupts memb of cht.
 Catalase detoxify H2O2 to
H2O + O2
 Electrons come from water to
oxygen and back to water but
the same e- are not recycled
like the cyclic flow do and for
this reason that is why is also
not referred to as cyclic flow.
 Occur - O2 conc. very high or
when CO2 fixation is very low
and it saturates at very high
light intensities.
 Measured by the rates of http://www.und.ac.za/und/icd/citte/paper/
oxygen uptake caused by light. net2/fig6.gif
Mode of Action of Some
Herbicides
Interupt photosynthetic e- flow
DCMU (diuron), block e- flow at the
quinone acceptors of PS II by
competing for the binding site of
plastoquinone with QB.
Paraquat intercepts e- between the
bound ferredoxin acceptors and
NADP and reduces O2 to
superoxide (O2–) that reacts
nonspecifically with a number of
molecules (especially lipids) in the
cht & damages them.
In herbicide-resistant biotypes of
common weeds the resistance factor
has been traced to a change in the
D1 protein, in the quinone (and Taiz & Zeiger 2006 Plant physiology
herbicide) binding region of the
peptide, that lowers the binding
affinity of the herbicide.
Conclusion
The light reactions prepares the energy and reductant
pool which are required in the next step of
photosynthesis i.e. the Dark Reactions or Calvin
Cycle.
O2 a by product of the reaction is important for life on
earth.
For queries
e mail:
rajkumarbot@yahoo.com
References
Plant Physiology Taiz and Zeiger (2006)

Biochemistry and Molecular Biology Buchanan et. al

of Plants 2000
Modern’s Botany Vol. III B.B. Arora (2008)

Plant Physiology, Biochemisty and HS Srivastava (2006)

Biotechnology

Molecular Biology of a Cell Anderson

Plant Physiology Salisbury and Ross (2004)

Plant Biochemistry Day and Harbour

http://www.biochem.arizona.edu/classes/bi http://jpkc.yzu.edu.cn/course/zhwshl/ppeboo
k/04z/imgch4/ch40406.gif
oc462/462b/Miesfeld/photo_phos/text_figu http://en.wikipedia.org
res/476_figure-19-54c.jpg
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