Chapter 6...

6.
3 Photosynthesis
What is photosynthesis?
• The process of converting solar energy into
chemical energy.
• Can use only water and carbon dioxide to create
sugars = chemical energy.
• Responsible for removal of ~ 200 billion tons of C
from the atmosphere yearly.
Light
6CO2 + 12 H2O C6H12O6 +6O2 +6H2O
p/pigments
• Two stages of photosynthesis

– Light reactions and Dark reactions
• Light reactions convert sunlight into chemical energy
(ATP + NADPH)
• Dark reactions use those products to form sugars (stored
chemical energy)
Overview of Photosynthesis
CO2
Dark Reactions
ATP
NADPH
H2O
+ Light Reactions Sugars
CO2
O2
How can light provide energy for
plants?
• Light is composed of particles: photons
• Light behaves like a wave
– Can be described into wavelength &
frequency
• Only a small portion of the electromagnetic
spectrum.
The electromagnetic spectrum
PAR = photosynthetically available radiation

Pigments
• Pigment = a light absorbing molecule
• Associated with the thylakoid membranes
• Chlorophyll
– Chl a and Chl b (Chl c in some algae)
• Xanthophylls
• Carotenoids
– ß-carotene
Where does photosynthesis take place?
Chloroplast Morphology
Chloroplast Morphology-Terms
• Inner Membrane
• Outer Membrane
• Thylakoid
• Thylakoid lumen
• Stroma
• Granum
Elodea
leaf X
400
The small green structures

within the cells of this plant
are chloroplasts.
Light | Pigments | Overview | Chloroplast | Photosystem II | Electron Transport System | Photosystem I | Calvin Cycle | Photorespiration | C4 plants | Review | Return
Chloroplast Structure
• The chloroplast is surrounded by a double membrane.
• Molecules that absorb light energy (photosynthetic
pigments) are located on disk-shaped structures called
thylakoids.
• The interior portion is the stroma.
Stroma
Double membrane
Thylakoids
The Light Reactions
• Organization of the light reaction system in the

chloroplast.
• Thylakoid membranes and lumen make up both lamellae,
the site of photosynthesis.
• Most reactions take place by the action of enzymes
embedded through the thylakoid membrane.
• Chlorophylls are the most important photosynthetic
pigment, containing a porphyrin ring with a coordinated
Mg2+ ion. Plants use a and b.
Chlorophyll
•Chl a has a methyl
group
•Chl b has a carbonyl

group
Porphyrin ring
Phytol tail
The Light Reactions
• The properties of a molecule determine which

wavelengths of light it can absorb.
• An absorption spectra measures the ability of a
molecule to absorb each wavelength of light.
• This can be determined for any molecule by using a

spectrophotometer.
This graph shows the color of
light absorbed by three
Chlorophyll a different kinds of
photosynthetic pigments.
Absorption Spectrum Notice that they do not absorb

light that is in the green to
Chlorophyll b yellow range.
absorption
Carotenoids
400 500 600 700

Wavelength
Different pigments absorb light
differently
Summary of photosynthesis
Summary of
Light Reactions
Compare:
The Light Reactions
• The electronic state of a molecule is altered when it

absorbs or emits light. The electrons orbitting the
atomic nuclei within the molecule jump to higher
energy levels (shells) and the energy will be lost within a
split second.
Chlorophyll + h = Chlorophyll*
Ground state light excited state

CHLOROPLAST – ENERGY TRANSDUCTION PATHWAYS
Light Reactions 1: Light capture and
redox
• 2 spatially & functionally distinct units =
Photosystems
• Photosystem II (P700) = 1st stage
• Photosystem I (P680)= 2nd stage
• Named after order of discovery and W/L
The Light Reactions
• the light reactions of photosynthesis are energized by light
at two points, called photosystems, in the electron transport
pathway.
Photosystem Light preferred
I Far-red (700) strong enough

to reduce NADP
II Red (680) strong enough

to oxidize water
The Light Reactions
The photosystems are spatially separated.
a. PSII (P680) is located in the grana lamellae.
b. PSI (P700) is located in the edges of the grana and
in the stroma lamellae.
c. They are linked by cytochrome b6 f .
d. This seems to improve the efficiency of each
photosystem
Antenna complexes--harvest light, zap energy to the reaction

center, where water split and electron transport take place.
Made up of carotenoids, chlorophylls a and b, and their
associated proteins, forming a complex called the light
harvesting complex, such as LHCII which is associated
with PSII.
The Light Reactions
An excited chlorophyll at the reaction center of PSII reduces
(gives its electron away to) an acceptor molecule.
• This electron is passed to

pheophytin w/in PSII.
• This half-filled orbital will be

filled with an electron taken
away from water.
This drawing shows a magnified view of a part of a
thylakoid. The green area is the thylakoid and the blue
area is the stroma of the chloroplast. Photosynthetic
pigments embedded within the membrane form a unit
called an antenna.
Antenna
Stroma
Thylakoid
membrane
Antenna
Photosynthetic pigments such as
chlorophyll A, chlorophyll B and
carotinoids.
A pigment molecule within the antenna absorbs a photon
of light energy. The energy from that pigment molecule is
passed to neighboring pigment molecules and eventually
makes its way to pigment molecule called the reaction
Light energy center. When the reaction center molecule becomes
excited (energized), it loses an electron to an electron
acceptor.
Thylakoid
membrane
Electron acceptor
Reaction Center, Electron Acceptor Reaction center
As a result of gaining an electron (reduction), the electron
acceptor becomes a high-energy molecule. Remember - its
energy came from light.
To understand this transfer of energy, recall that oxidation is
Light energy the loss of an electron and the loss of energy. Reduction is the
gain of an electron and energy. Energy is transferred with the
electron.
Thylakoid
membrane
Reaction Center, Electron Acceptor Electron acceptor
Reaction center
The antenna and electron acceptor are called a photosystem.
There are two kinds of photosystems in plants called photosystem I and photosystem II.
Photosystem I is sometimes called P700 and photosystem II is sometimes P680. The 680
and 700 designations refer to the wavelength of light that they absorb best.
Photosystem
Antenna
Thylakoid
membrane
Antenna, Photosystem Electron acceptor
Reaction center
Phosphorylation
• Chemiosmotic phosphorylation occurred in both

the mitochondria (during cellular respiration)
and in the chloroplast (during photosynthesis).
These two processes are sometimes given
separate names:
– Oxidative phosphorylation (in mitochondria)
– Photophosphorylation (in chloroplast)
Light
Energy Chloroplast
Electron Transport
System
Photosystem II Photosystem I
The three blue circles represent the electron transport system. They are proteins
embedded within the thylakoid membrane.
The first protein receives the electron (and energy) from the electron acceptor.
Stroma
Light
Energy Chloroplast
Electron
H+
Transport System
H+ H+
H+
H+ H+ H+
H+
As a result of gaining an electron (reduction), the first carrier of the

electron transport system gains energy. It uses some of the energy to
pump H+ into the thylakoid.
Thylakoids Stroma
Light
Energy Chloroplast
H+ Electron
H+
Transport
H+
H+
System
H+ H+
H+
The carrier then passes the electron to the next carrier. Because it
used some energy to pump H+, it has less energy (reducing capability)
to pass to the next H+ pump.
Thylakoids Stroma
Light
Energy Chloroplast
H+
H+
H+
H+ H+ H+
H+
H+
Electron
Transport System
This carrier uses some of the remainder of the energy to pump more
H+ into the thylakoid.
Thylakoids Stroma
Light
Energy Chloroplast
H+
H+
H+
H+ H+ H+
H+
H+
Electron
Transport System
The electron is passed to the next carrier which also pumps H+.
Thylakoids Stroma
Light
Energy Chloroplast
H+
H+
H+ H+ H+
H+ H+ H+
Electron Transport
System
The electron transport system functions to create a concentration
gradient of H+inside the thylakoid.
Thylakoids Stroma
Light
Energy Chloroplast
The concentration gradient of H+ is used to synthesize ATP.

ATP is produced from ADP and Pi when hydrogen ions pass
out of the thylakoid through ATP synthase.
H+
H+ H+ H+
H+ H+ ATP
H+
Photophosphorylatio ADP + Pi
n H+
Thylakoids Stroma
Light
Energy Chloroplast
This method of synthesizing ATP by using a H+ gradient

in the thylakoid is called photophosphorylation.
H+
H+ H+ H+
H+ H+ ATP
Photophosphorylati H+
ADP + Pi
on H+
Thylakoids Stroma
A Thylakoid H+
H+
H+ H+ H+ + H + H+ +
H+ H H H +
H+ H+ H+ H+
In order to synthesize ATP, hydrogen ions H+

must first be pumped into the thylakoid. This
process requires energy.
A Thylakoid H+
H+
H+ H+ H+ + H + H+ +
H+ H H H +
H+ H+ H+ H+
A concentration gradient of hydrogen ions is

established. The osmotic pressure from this
H+
gradient can be used as an energy source for
producing ATP.
Chemiosmotic Phosphorylation
Osmotic pressure forces hydrogen ions through this H+
protein (ATP synthase) as they return
H+ to the stroma.
H+ H+ H+ + H + H+ +
H+ H H H +
ADP + Pi H+ H+ H+ H+
ATP synthase produces ATP by

ATP phosphorylating ADP. The energy
comes from hydrogen
+ ions forcing
H
H+ their way into the stroma as they pass
through the ATP synthase under
pressure (osmotic pressure).
PSII electron transport chain
H+ H+
1. Q accepts 2 e- from
Q
e- e- P680 and removes 2 H+
from the stroma
2. Q passes the e- to
e- e- cytochrome b/f complex &
pumps the 2H+ into thylakoid
cytochrome lumen
b/f 3. As the e- moves through
complex
b/f more H+ are pumped into
lumen
The PS II Electron transport chain
H+ H+
Q
e- e-
cytochrome
b/f
complex
2H+ e-
H+ plastocyanin
PSI electron transport chain
Ferredoxin
e-
1. Ferredoxin receives e- from P700*
Fe/S
2. e- moves through FAD (flavin
FAD adenine dinucleotide)
H+
FAD-NAPD e-
Reductase 3. e- plus stromal H+ are used to
NADP+ reduce NADP+ to NADPH
NADPH = FINAL PRODUCT!

PS II to PS I
e- 4. e- end up on
plastocyanin
plastocyanin: a soluble
electron carrier in the lumen
5. Plastocyanin serves as
e- donor for PSI reaction
center Chl a
e-
plastocyanin
H2O O2
light Summary of
light reactions
Photosynthesis
ATP NADPH
ADP
C02 light-independent reactions NADP+
(Calvin cycle)
C6H12O6
Light
Energy Chloroplast
H+
H+ H+ H+
H+ H+ ATP
H+
Photosystem ADP + Pi
H+
I
At this point, the electron has little reducing capability
(little energy is left). It is passed to the P700 antenna.
Thylakoids Stroma
Light
Energy Chloroplast
H+
H+ H+ H+
H+ H+ ATP
H+
P700 ADP + Pi
Antenna H+
A pigment molecule in the P700 antenna absorbs a photon of

solar energy. The energy from that molecule is passed to
neighboring molecules within the antenna. The energy is
eventually passed to the reaction center of this antenna.
Thylakoids Stroma
Light
Energy Chloroplast
H+
H+ H+ H+
H+ H+ ATP
H+
Electron ADP + Pi
Acceptor H+
As a result of being energized, the P700 reaction

center loses the electron to an electron acceptor.
Thylakoids Stroma
Light
Energy Chloroplast
NADP+ + H+
NADPH
H+
H+ H+ H+
H+ H+ ATP
H+
ADP + Pi
NADP+ H+
The acceptor passes it to NADP+, which becomes

reduced to NADPH.
NADP+ + 2e- + H+ → NADPH
Thylakoids Stroma
Light
Energy Chloroplast
NADP+ + H+
NADPH
H+
H+ H+ H+
H+ H+ ATP
H+
ADP + Pi
Splitting H2O H+
The electron that was initially lost by

photosystem II (P680) must be replaced.
Thylakoids Stroma
Light
Energy Chloroplast
NADP+ + H+
NADPH
H+
H+ H+ H+
H+ H+ ATP
H+
H2O→2e- + 2H+ + ½ O2
ADP + Pi
Splitting H2O H+
A hydrogen atom contains one electron (e-) and one

proton (H+). The two hydrogen atoms in a water
molecule can therefore be used to produce 2e- and 2H+.
Thylakoids Stroma
Photosystem I
e-* 1. Charge
separation and
photo-oxidation
are similar to PSII
Optimal
wavelength =
+
700nm
P700
e- 2. Plastocyanin acts as
plastocyanin reducing agent on P700
Chl a
PSI electron transport chain
e-
Ferredoxin
e- Fe/S
e-
FAD H+
FAD-NAPD e-
Reductase
NADP+ NADPH
ATP synthesis
H+ 1. O2 evolving complex
H+ liberates H+ into lumen from
stroma
water
2. Q and Cyt b/f pump H+ from

stroma into lumen.
lumen
H+
H+ 3. NADP+ scavenges protons
H+ H+ H+ from the stroma
H+
H+ H+
pH separation across
H+ H+
H+ membrane = Proton Motive
Force!
ATP synthesis
ADP + Pi
F0 F1 H+
ATP complex
stroma
H+
lumen H+ The energy released as

H+ protons travel down their
H+ H+ H+ concentration gradient is
H+ used to fuel an ATP
H+ H+
synthase
H+
H+
NADPH
light e- acceptor
NADP+
e- acceptor
ATP
electron
transport
system
P 700 antenna
Summary of Light
Reactions
complex
P680 antenna
complex H2O → 2e- + 2H+ + O
Light
Energy Chloroplast
NADP+ + H+ CO2
NADPH
H+ Calvin
H+ H+ H+
H+ H+ ATP Cycle
H+
H2O→2e- + 2H+ + ½ O2
Calvin ADP + Pi
Cycle H+ glucose
The next several slides show how the products of

the light reactions (ATP and NADPH) are used to
reduce CO2 to carbohydrate in the Calvin cycle.
Thylakoids Stroma
Light
Energy Chloroplast
NADP+ + H+ CO2
NADPH
H+ Calvin
H+ H+ H+
H+ H+ ATP Cycle
H+
H2O→2e- + 2H+ + ½ O2
ADP + Pi
Calvin Cycle H+ glucose
The reactions of the Calvin cycle occur in the stroma of the chloroplast.
Thylakoids Stroma
Light Reactions Products
In: CO2, H2O, sunlight
Out: O2, ATP, NADPH

This slide summarizes photosynthesis.
H2O O2 6CO2 + 6H2O + E → C6H12O6 + 6O2
light Summary of
light reactions
Photosynthesis
ATP NADPH
ADP
C02 light-independent reactions NADP+
(Calvin cycle)
C6H12O6
H2O O2
Light
Energy Chloroplast
NADP+ + H+
light
NADPH
H+ + H+
H H+ H+ H+
lightofreactions
H+ ATP
H2O→2e- + 2H+ + ½ O2
H+
ADP + Pi
Summary Photosynthesis
Thylakoids Stroma
ADP
ATP NADPH
NADP+
C02 light-independent reactions

(Calvin cycle)
C6H12O6
The Calvin cycle - uses the chemical
energy of ATP and NADPH to reduce
CO2 to sugar
• The Calvin cycle, like the citric acid cycle,
regenerates its starting material after molecules enter
and leave the cycle
• The cycle builds sugar from smaller molecules by
using ATP and the reducing power of electrons
carried by NADPH
© 2011 Pearson Education, Inc.

• Carbon enters the cycle as CO2 and leaves as a
sugar named glyceraldehyde 3-phospate (G3P)
• For net synthesis of 1 G3P, the cycle must take
place three times, fixing 3 molecules of CO2
• The Calvin cycle has three phases:
– Carbon fixation (catalyzed by rubisco)

– Reduction
– Regeneration of the CO2 acceptor (RuBP)
© 2011 Pearson Education, Inc.

CO2 Fixation
• CO2 fixation refers to bonding CO2 to an

organic molecule to make a larger
molecule.
• C5 + CO2 → C6
“C5” is an abbreviation that means that this

molecule has 5 carbon atoms.
6 CO2
6 C-C-C-C-C-C
CO 2 Fixation
RuBP The enzyme that catalyzes this
Carboxylase reaction is ribulose biphosphate
(rubisco) carboxylase (rubisco).
6 C-C-C-C-C
CO2 fixation refers to bonding CO2 to an organic

molecule to make a larger molecule.
Each CO2 is bonded to ribulose biphosphate
(RuBP).
C5 + CO2 → C6
C3 Photosynthesis
6 CO
– Calvin Cycle
2
6 C-C-C-C-C-C
RuBP
Carboxylase
(rubisco)
RuBP 6 C-C-C-C-C PGA 12 C-C-C
Each of these 6-carbon

compounds splits to form two
3-carbon compounds called
phosphoglycerate.
6 CO2 Calvin Cycle
6 C-C-C-C-C-C
RuBP
Carboxylase
(rubisco)
PGAL
12 C-C-C
The two molecules of PGA are

reduced to form PGAL
(phosphoglyceraldehyde).
6 CO2 Calvin Cycl
6 C-C-C-C-C-C
RuBP
Carboxylase
(rubisco)
12 ATP
PGAL
12 C-C-C 12 ADP + P
12 NADPH
12 NADP+
6 CO2 Calvin Cycle
6 C-C-C-C-C-C
RuBP
Carboxylase
(rubisco)
RuBP 6 C-C-C-C-C 6 ADP + P PGA 12 C-C-C
6 ATP
Two of the PGAL are used to form glucose phosphate, then
glucose.
10 C-C-C 12 ATP
PGAL
12 C-C-C 12 ADP + P
12 NADPH
C-C-C-C-C-C Glucose 12 NADP+
6 CO2 Calvin Cycle
6 C-C-C-C-C-C
RuBP
Carboxylase
(rubisco)

6 ADP + P The remaining 10 PGAL are
rearranged to form 6 RuBP.
6 ATP
10 C-C-C 12 ATP
PGAL
12 C-C-C 12 ADP + P
12 NADPH
6 CO2 Calvin Cycle
6 C-C-C-C-C-C
RuBP
Carboxylase
(rubisco)

6 ADP + P This process requires
energy in the form of ATP.
6 ATP
10 C-C-C 12 ATP
PGAL
12 C-C-C 12 ADP + P
12 NADPH
Summary:
Light
energy
ECOSYSTEM
Photosynthesis
in chloroplasts
CO2 + H2O Organic
THE + O2
molecules
Cellular respiration
ENERGY in mitochondria
CYCLE
ATP powers
ATP
most cellular work
Heat
energy

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6.

• Two stages of photosynthesis

PAR = photosynthetically available radiation

The small green structures

• Organization of the light reaction system in the

•Chl b has a carbonyl

• The properties of a molecule determine which

• This can be determined for any molecule by using a

Absorption Spectrum Notice that they do not absorb

400 500 600 700

• The electronic state of a molecule is altered when it

Ground state light excited state

Photosystem Light preferred

I Far-red (700) strong enough

II Red (680) strong enough

Antenna complexes--harvest light, zap energy to the reaction

• This electron is passed to

• This half-filled orbital will be

Reaction Center, Electron Acceptor Electron acceptor

Antenna, Photosystem Electron acceptor

• Chemiosmotic phosphorylation occurred in both

As a result of gaining an electron (reduction), the first carrier of the

The concentration gradient of H+ is used to synthesize ATP.

This method of synthesizing ATP by using a H+ gradient

In order to synthesize ATP, hydrogen ions H+

A concentration gradient of hydrogen ions is

ATP synthase produces ATP by

NADPH = FINAL PRODUCT!

A pigment molecule in the P700 antenna absorbs a photon of

As a result of being energized, the P700 reaction

The acceptor passes it to NADP+, which becomes

The electron that was initially lost by

A hydrogen atom contains one electron (e-) and one

2. Q and Cyt b/f pump H+ from

lumen H+ The energy released as

The next several slides show how the products of

Calvin Cycle H+ glucose

In: CO2, H2O, sunlight

Out: O2, ATP, NADPH

C02 light-independent reactions

© 2011 Pearson Education, Inc.

– Carbon fixation (catalyzed by rubisco)

© 2011 Pearson Education, Inc.

• CO2 fixation refers to bonding CO2 to an

“C5” is an abbreviation that means that this

CO2 fixation refers to bonding CO2 to an organic

RuBP 6 C-C-C-C-C PGA 12 C-C-C

Each of these 6-carbon

RuBP 6 C-C-C-C-C PGA 12 C-C-C

The two molecules of PGA are

RuBP 6 C-C-C-C-C PGA 12 C-C-C

RuBP 6 C-C-C-C-C 6 ADP + P PGA 12 C-C-C

C-C-C-C-C-C Glucose 12 NADP+

RuBP 6 C-C-C-C-C PGA 12 C-C-C

C-C-C-C-C-C Glucose 12 NADP+

RuBP 6 C-C-C-C-C PGA 12 C-C-C

C-C-C-C-C-C Glucose 12 NADP+

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