CHAPTER 7

PHOTOSYNTHESIS
"Without this flow of
energy from the sun,
channeled largely through
the eukaryotic cells, the
pace of life on this planet
would swiftly diminish and
then, following the
inexorable second law of
thermodynamics would
virtually cease altogether."
Raven, P. H. et al. 1992.

14 pts
PHOTOSYNTHESIS
OVERVIEW
 Photosynthesis - summary
CARBON DIOXIDE + WATER + LIGHT 
CARBOHYDRATE + OXYGEN

6 CO2 + 6 H2O  C6H12O6 + 6 O2 + H2O

• This reaction captures energy from sunlight

• Light energy is used to reduce carbon (removes hydrogen
from water and adds it to carbon)
• Energy is stored in plant tissues as starch, sugars and in
cellulose (wood, coal, gas, oil)
• Stored energy can be released by burning, or by metabolism
IMPORTANCE OF
PHOTOSYNTHESIS
 PHOTOSYNTHETIC ORGANISMS
ARE AT THE BASE OF THE FOOD CHAIN

•AUTOTROPHS (Can make own food, i.e., can

manufacture organic compounds from inorganic
compounds)
•HETEROTROPHS (Eat others to get organic
compounds)
 Photosynthesis is the source of
Oxygen in the atmosphere
 Earth's atmosphere is a layer of gases surrounding the planet
and retained by the Earth's gravity.
 78% nitrogen
 20.95% oxygen
 0.93% argon
 0.038% carbon dioxide (up from 0.028%)
 a variable amount (average around 1%) of water vapor
 Oxygen is reactive with inorganic compounds and does not
stay in the atmosphere. Thus, our current atmosphere is a
balance of oxygen produced by plants and loss to geological
processes
THE NATURE
OF LIGHT
• THE VISIBLE SPECTRUM CAN BE SEEN
WHEN WHITE LIGHT IS PASSED
THROUGH A PRISM.

• THE SPECTRUM IS SEEN AS DIFFERENT

COLORS OF LIGHT
A “SPECTRUM” IS A RANGE OF
WAVELENGTHS OF ELECTROMAGNETIC
RADIATION

One wavelength
Ultraviolet light, X-rays and gamma rays
have too much energy for biological systems.
They break bonds, knock electrons out of
orbit and generally cause cell damage. We
are protected from UV by the ozone layer

Infrared light is heat and has too low an

energy level to be used for photosynthesis
• Pigments are molecules that absorb light. We see
them as being the color of the light that is reflected

• In order to trap light, light must be absorbed. (If

light is reflected, it has no value)
PIGMENTS
 T. W. Engelmann (1883)

• Designed an elegant experiment with spirogyra, an

algae with a single long coiled chloroplast.
• He combined the algae with an oxygen-requiring
bacterium and exposed the algae to light coming
through a prism.
• He showed that the bacteria clustered in the blue
and red areas suggesting that these areas were
producing oxygen.
T. W. Englemann’s experiment 1882

Spirogyra
 PIGMENTS OF PLANTS
•Chlorophyll a
•Chlorophyll b
• Chlorophyll c
(replaces Chlorophyll b in some algae)
•Carotenoids
•Carotene
•Xanthophylls
•Phycobilins (photosynthetic bacteria)
•Phycoerythrin
•Phycocyanin
Carotenoids
•Absorb maximally
between 460 and 550
nm, absorb in the
blue range and
therefore appear red,
orange and yellow
The third major class of
pigments, the phycobilins,
are found in the
cyanobacteria and red algae
only
 REACTIONS OF
PHOTOSYNTHESIS
A. LIGHT vs DARK REACTIONS
 F. F. Blackman (1905)
English Plant Physiologist: Concluded that there
were two reactions, one of which was light dependent
(photochemical reactions) and one of which was
temperature dependent (biochemical reactions or
dark reactions)
 Light reactions take place in the grana thylakoids

Grana Plant cell wall

(stacked
granum Stroma
thylakoids)

Stroma
thylakoids

Chloroplast double membrane

 Closeup of grana and stroma
A stack of folded membranes is referred to as a granum. The stroma
is a watery matrix

Stack of
granum
thylakoids

Stroma
Dark or biochemical rxns
 REACTIONS OF
PHOTOSYNTHESIS
B. LIGHT REACTION
Light-trapping pigments float in the thylakoid membranes
 The pigment aggregates have been given the name
antenna complex because it acts as an antenna to trap
light and transfer the absorbed energy

Antenna complex =
Chlorophyll a,
chlorophyll b and
carotenoids
When light has the correct energy level, it is absorbed
by an electron. The electron must move to a new
orbital whose energy level corresponds to the
electron’s new energy load

Photon

e-
The excited state is unstable. The atom can become
stable if the electron drops back into lower orbit.
When this happens the excess energy may be released
as visible light (fluorescence) or heat (infared light)

e-

Heat or light
Alternately, the electron can move to a more stable
orbital on an entirely different atom. This electron
transfer is the critical step in photosynthesis
Reaction center: Within the antenna complex, there is
a special pair of chlorophyll A molecules and
associated proteins called a reaction center
Function of the reaction center: accepts electrons
transferred from the antenna complex and boosts
them into higher orbit where they can be transferred
to another electron acceptor molecule
There are two different types of reaction
center/antenna complexes in the
thylakoid membranes (light reaction).
• The reaction center of Photosystem I absorbs
maximally at 700 nm. The reaction center is called
P700

• The reaction center of Photosystem II absorbs

maximally at 680 nm. The reaction center is called
P680
Energy depiction of Photosynthesis, the “Z” scheme
 Photophosphorylation
Photophosphorylation = LIGHT driven production of ATP
1. Electrons are passed along a series of electron carriers

When electrons reach transmembrane pumps, protons are

pumped into the lumen of the thylakoid membrane

This proton gradient drives

synthesis of ATP into the
stroma (cytoplasm) of the
chloroplast

2. Similar in principle to
oxidative phosphorylation
in mitochondria
 Light reactions - summary
• Requires light
• Independent of temperature (an indication that
enzymes are not involved)
• Light is used to make ATP and to reduce electron
carrier molecules
 Summary of light reaction
Eight photons used in the light (photochemical) reaction

Produce 3 ATPs
Reduce 2 molecules of
NADP+ to NADPH
Oxygen is produced
from splitting water
 CYCLIC ELECTRON FLOW
In photosynthetic prokaryotes, only part of the
reaction is present (photosystem 1). This is the
older system. Green plants have added a
second system to increase energy capture.
•No water is split
•No NADPH is made
a. Cyclic phosphorylation alone may have existed for a billion years in bacteria who
used sulfur as an electron donor but modern plants use water as a source of
electrons and that required a major change because the energy in P700 isn’t high
enough to remove an electron from water.
b. Plants overcame this problem by grafting on a second, more powerful
photosystem which could harvest shorter, higher energy wavelengths of light (Z
scheme)
 REACTIONS OF
PHOTOSYNTHESIS
B. DARK REACTIONS
PHOTOSYNTHESIS IS COMPOSED OF
TWO SETS OF REACTIONS

LIGHT DEPENDENT = PHOTOCHEMICAL

TEMPERATURE INDEPENDENT
LIGHT DEPENDENT

DARK REACTIONS = CARBON FIXATION

RXNS = BIOCHEMICAL RXNS
TEMPERATURE DEPENDENT
LIGHT INDEPENDENT
Berkeley scientist, Melvin Calvin (1950) was
the first to elucidate the steps in the dark
reaction of photosynthesis
 Dark dependent stage
• Occurs in the light but does not require light
• Temperature dependent or sensitive reactions. Reactions
increase up to 30C then decrease. This temperature profile is
an indication that enzymes are involved
• Energy products of the light reaction are used to reduce
carbon from CO2 to a sugar (carbon fixation)
 RUBISCO

Ribulose-1,5-bisphosphate or RuBP
8 large subunits (gene in
chloroplast)
8 small subunits (gene in
nucleus)
Most abundant protein on earth
 Sugar

PGAL is an intermediate in respiration during the oxidation of

glucose to pyruvate
 C3 PLANTS

PLANTS WHICH USE ONLY THE CALVIN CYCLE TO

FIX CO2 INTO SUGARS ARE CALLED C3 PLANTS
(BECAUSE A THREE-CARBON MOLECULE IS THE
FIRST STABLE MOLECULE IN THE CYCLE)

MOST PLANT SPECIES ARE C3 PLANTS

CEREALS
PEANUTS
COTTON
SOYBEANS
MOST TREES
MOST LAWN GRASSES
 REACTIONS OF
PHOTOSYNTHESIS
Photorespiration
 PHOTOSYNTHESIS IS NOT PERFECT:
PHOTORESPIRATION
•When CO2 levels in the leaf get too low, RUBISCO starts
binding oxygen instead of CO2
•Happens when stomates close to prevent water loss. (This
prevents CO2 diffusion into the leaf)
•Forces another pathway
O2 + RuBP  PGA + phosphoglycolic acid
•Reactions called photorespiration because they occur in the light,
use oxygen and release CO2
•Unlike normal respiration, it produces no ATP
•It is a wasteful salvage pathway
 RUBISCO CO2

•RUBISCO binds
CO2and O2 in the same
site
•If oxygen is bound, the
Calvin cycle cannot take O2

place

•Instead, photorespiration
takes place
 REACTIONS OF
PHOTOSYNTHESIS
The C4 pathway
 C4 PLANTS
• Some plants have developed mechanisms to bypass
the RUBISCO carboxylase/oxygenase problem
• These plants are called C4 plants because they fix
carbon into a 4-carbon sugar
• Plants out-compete C3 plants in hot dry weather only.
C4 carbon fixation has
evolved on up to 40
independent occasions in
different groups of plants,
making it an example of
convergent evolution. Plants
which use C4 metabolism
include sugarcane, maize,
sorghum, finger millet,
amaranth, and switchgrass.
C4 plants arose around
25 to 32 million years ago
C4 plants have a specific kind of leaf anatomy. Carbon is fixed into
malate in mesophyll cells. The Calvin cycle occurs in bundle sheath
cells that surround the vascular system (few thylakoid grana and
protected from oxygen). Thus, the two processes are separated in
space. Calvin cycle Malate
Mesophyll cell with granna

Bundle sheath cell

Chloroplasts lack grana
Malate
Plasmodesmata

Calvin cycle-note few grana

 REACTIONS OF
PHOTOSYNTHESIS
CAM Plants
Crassulacean acid metabolism
Separates CO2 intake and the Calvin
cycle by time rather than by space

Crassula ovata – Jade Plant

Crassulacean acid metabolism
The majority of plants
possessing
Crassulacean Acid
Metabolism are either
epiphytes (e.g. orchids,
bromeliads) or
succulent xerophytes
(e.g. cacti, cactoid
Euphorbias).

16,000 species (7% of

plants). Most are
angiosperms but
includes some
gymnosperms, ferns
and quillworts
 REACTIONS OF
PHOTOSYNTHESIS
Carbon Cycle
Global Warming
Another important carbon “sink” is the arctic tundra which is
now decomposing and releasing CO2 as the arctic thaws.
 Carbon Dioxide in the atmosphere
GREENHOUSE
EFFECT – HUMANS
ARE RELEASING
STORED CARBON
WHICH INCREASES
ATMOSPHERIC CO2

The present level is higher than at any time during the last 800 thousand
years,[6] and likely higher than in the past 20 million years.[7]
6.Amos, Jonathan (2006-09-04). "Deep ice tells long climate story". BBC News.
http://news.bbc.co.uk/2/hi/science/nature/5314592.stm. Retrieved 2010-04-28.
7. Climate Change 2001: The Scientific Basis
 TRENDS
•TOTAL GHG EMMISIONS INCREASED 14.2% SINCE 1990
•DOMINANT GAS EMITTED IS CO2, MOSTLY FROM FOSSIL FUEL
COMBUSTION (oxidation of sugars produced by fossil plants)
 Summer
NOAA CO2 tracker

Red=high CO2 emissions

Blue = low CO2 emissions

Winter
NOAA CO2 tracker
END