Chapter 11

Photosynthetic
Processes

Lecture Presentations by
Nicole Tunbridge and
© 2018 Pearson Education Ltd. Kathleen Fitzpatrick
The Process That Feeds the Biosphere

▪ Plants and other photosynthetic organisms contain

organelles called chloroplasts
▪ Photosynthesis is the process that converts solar
energy into chemical energy within chloroplasts
▪ Directly or indirectly, photosynthesis nourishes
almost the entire living world

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▪ Autotrophs are “self-feeders” that sustain
themselves without eating anything derived from
other organisms
▪ Autotrophs are the producers of the biosphere,
producing organic molecules from CO2 and other
inorganic molecules
▪ Almost all plants are photoautotrophs, using the
energy of sunlight to make organic molecules

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Figure 11.2

(d) Cyanobacteria 40
µm
(a) Plants

(b) Multicellular alga

1 µm
(e) Purple sulfur
10 µm

bacteria

(c) Unicellular eukaryotes

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Chloroplasts: The Sites of Photosynthesis
in Plants
▪ Leaves are the major locations of photosynthesis in
plants
▪ Chloroplasts are found mainly in cells of the
mesophyll, the interior tissue of the leaf
▪ Each mesophyll cell contains 30–40 chloroplasts
▪ CO2 enters and O2 exits the leaf through microscopic
pores called stomata

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▪ A chloroplast has an envelope of two membranes
surrounding a dense fluid called the stroma
▪ Thylakoids are connected sacs in the chloroplast
that compose a third membrane system
▪ Thylakoids may be stacked in columns called grana
▪ Chlorophyll, the pigment that gives leaves their
green color, resides in the thylakoid membranes

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Figure 11.4a
Leaf cross section
Chloroplasts Vein

Mesophyll

Stomata
CO2 O2

Chloroplast Mesophyll
cell

20 µm
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Figure 11.4b

Outer
Thylakoid membrane
Thylakoid Intermembrane
Stroma Granum space space
Inner
membrane

Chloroplast 1 µm
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Tracking Atoms Through Photosynthesis:
Scientific Inquiry
▪ Photosynthesis is a complex series of reactions that
can be summarized as the following equation:
6 CO2 + 12 H2O + Light energy → C6H12O6 + 6 O2 +
6 H 2O
▪ The overall chemical change during photosynthesis
is the reverse of the one that occurs during cellular
respiration

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The Splitting of Water

▪ Chloroplasts split H2O into hydrogen and oxygen,

incorporating the electrons of hydrogen into sugar
molecules and releasing oxygen as a by-product

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Figure 11.5

Reactants: 6 CO2 12 H2O

Products: C6H12O6 6 H2O 6 O2

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Photosynthesis as a Redox Process

▪ Photosynthesis reverses the direction of electron

flow compared to respiration
▪ Photosynthesis is a redox process in which H2O is
oxidized and CO2 is reduced
▪ Photosynthesis is an endergonic process; the
energy boost is provided by light

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Figure 11.UN01

becomes reduced

becomes oxidized

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The Two Stages of Photosynthesis: A Preview

▪ Photosynthesis consists of the light reactions

(the photo part) and Calvin cycle (the synthesis
part)
▪ The light reactions (in the thylakoids)
▪ Split H2O
▪ Release O2
▪ Reduce the electron acceptor NADP+ to NADPH
▪ Generate ATP from ADP by photophosphorylation

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▪ The Calvin cycle (in the stroma) forms sugar from
CO2, using ATP and NADPH
▪ The Calvin cycle begins with carbon fixation,
incorporating CO2 into organic molecules

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Figure 11.6_1

Light H2O

NADP+

ADP
+
LIGHT Pi
REACTIONS

Thylakoid Stroma

Chloroplast

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Figure 11.6_2

Light H2O

NADP+

ADP
+
LIGHT Pi
REACTIONS
ATP
Thylakoid Stroma
NADPH

Chloroplast
O2

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Figure 11.6_3

Light H2O CO2

NADP+

ADP
+
LIGHT Pi CALVIN
REACTIONS CYCLE

ATP
Thylakoid Stroma
NADPH

Chloroplast
O2

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Figure 11.6_4

Light H2O CO2

NADP+

ADP
+
LIGHT Pi CALVIN
REACTIONS CYCLE

ATP
Thylakoid Stroma
NADPH

Chloroplast
O2 [CH2O]
(sugar)
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Concept 11.2: The light reactions convert solar
energy to the chemical energy of ATP and
NADPH
▪ Chloroplasts are solar-powered chemical factories
▪ Their thylakoids transform light energy into the
chemical energy of ATP and NADPH

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The Nature of Sunlight

▪ Light is electromagnetic energy, also called

electromagnetic radiation
▪ Electromagnetic energy travels in rhythmic waves
▪ Wavelength is the distance between crests
of electromagnetic waves
▪ Wavelength determines the type of electromagnetic
energy

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▪ The electromagnetic spectrum is the entire range
of electromagnetic energy, or radiation
▪ Visible light consists of wavelengths (380 nm to
750 nm) that produce colors we can see
▪ Visible light also includes the wavelengths that drive
photosynthesis
▪ Light also behaves as though it consists of discrete
particles, called photons

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Figure 11.7

1m
10–5 nm 10–3 nm 1 nm 103 nm 106 nm (109 nm) 103 m

Gamma Micro- Radio

X-rays UV Infrared waves waves
rays

Visible light

380 450 500 550 600 650 700 750 nm

Shorter wavelength Longer wavelength

Higher energy Lower energy

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Photosynthetic Pigments: The Light Receptors

▪ Pigments are substances that absorb visible light

▪ Different pigments absorb different wavelengths
▪ Wavelengths that are not absorbed are reflected or
transmitted
▪ Leaves appear green because chlorophyll reflects
and transmits green light

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Figure 11.8

Light
Reflected
light

Chloroplast

Absorbed Granum
light

Transmitted
light

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▪ There are three types of pigments in chloroplasts:
▪ Chlorophyll a, the key light-capturing pigment
▪ Chlorophyll b, an accessory pigment
▪ Carotenoids, a separate group of accessory pigments

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Figure 11.10

of light by chloroplast
Chloro-
phyll a Chlorophyll b

Absorption

pigments
Carotenoids

500 400 600 700

Wavelength of light (nm)
(a) Absorption spectra
Rate of photosynthesis
(measured by O2
release)

400 500 600 700

(b) Action spectrum

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Figure 11.11
CH3 in chlorophyll a
CHO in chlorophyll b

CH3

Porphyrin ring:
light-absorbing
“head” of molecule;
note magnesium
atom at center

Hydrocarbon tail:
interacts with hydrophobic
regions of proteins inside
thylakoid membranes of
chloroplasts; H atoms not
shown
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▪ Accessory pigments called carotenoids may
broaden the spectrum of colors that drive
photosynthesis
▪ Some carotenoids function in photoprotection; they
absorb excessive light that would damage
chlorophyll or react with oxygen

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Excitation of Chlorophyll by Light

▪ When a pigment absorbs light, it goes from a ground

state to an excited state, which is unstable
▪ When excited electrons fall back to the ground state,
excess energy is released as heat
▪ In isolation, some pigments also emit light, an
afterglow called fluorescence

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Figure 11.12

Excited
e– state
Energy of electron

Heat

Photon
(fluorescence)
Photon
Ground
Chlorophyll state
molecule

(a) Excitation of isolated chlorophyll molecule (b) Fluorescence

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A Photosystem: A Reaction-Center Complex
Associated with Light-Harvesting Complexes
▪ A photosystem consists of a reaction-center
complex surrounded by light-harvesting complexes
▪ The reaction-center complex is an association of
proteins holding a special pair of chlorophyll a
molecules and a primary electron acceptor

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▪ The light-harvesting complex consists of pigment
molecules bound to proteins
▪ Light-harvesting complexes transfer the energy of
photons to the chlorophyll a molecules in the
reaction-center complex
▪ These chlorophyll a molecules are special because
they can transfer an excited electron to a different
molecule

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▪ A primary electron acceptor in the reaction center
accepts excited electrons and is reduced as a result
▪ Solar-powered transfer of an electron from a
chlorophyll a molecule to the primary electron
acceptor is the first step of the light reactions

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Figure 11.13

Photosystem STROMA
Photon Light- Reaction-
harvesting center Primary
complexes complex electron
acceptor
Thylakoid membrane

Thylakoid membrane
Chlorophyll (green) STROMA
e–

Transfer Special pair of chloro-

of energy phyll a molecules Protein
Pigment
THYLAKOID SPACE molecules subunits THYLAKOID
(INTERIOR OF THYLAKOID) (purple) SPACE
(a) How a photosystem harvests light (b) Structure of a photosystem

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▪ There are two types of photosystems in the thylakoid
membrane
▪ Photosystem II (PS II) functions first (the numbers
reflect order of discovery)
▪ The reaction-center chlorophyll a of PS II is called
P680 because it is best at absorbing a wavelength of
680 nm

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▪ Photosystem I (PS I) is best at absorbing a
wavelength of 700 nm
▪ The reaction-center chlorophyll a of PS I is called
P700

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Linear Electron Flow

▪ During the light reactions, there are two possible

routes for electron flow: cyclic and linear
▪ Linear electron flow, the primary pathway, involves
both photosystems and produces ATP and NADPH
using light energy

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▪ There are eight steps in linear electron flow:
1. A photon hits a pigment in a light-harvesting
complex of PS II, and its energy is passed among
pigment molecules until it excites P680
2. An excited electron from P680 is transferred to the
primary electron acceptor (we now call it P680+)

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Figure 11.UN02

H2O CO2

Light

NADP+
ADP

LIGHT CALVIN
REACTIONS CYCLE

ATP
NADPH

O2 [CH2O] (sugar)

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Figure 11.14_1

Primary
electron
acceptor

e–
2

1 P680
Light

Pigment
molecules
Photosystem II
(PS II)

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Figure 11.14_2

Primary
electron
acceptor

2 H+ e–
2
H2O
+
1/2 O2 3
e–
e–
1 P680
Light

Pigment
molecules
Photosystem II
(PS II)

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Figure 11.14_3

4 Electron
transport chain
Primary
electron
acceptor Pq

2 H+ e–
2 Cytochrome
H2O
complex
+
1/2 O2 3 Pc
e–

1
e–
P680 5
Light
ATP

Pigment
molecules
Photosystem II
(PS II)

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Figure 11.14_4

4 Electron Primary
transport chain electron
Primary acceptor
electron
acceptor Pq e–

2 H+ e–
2 Cytochrome
H2O
complex
+
1/2 O2 3 Pc
e– P700
1
e–
P680 5 Light
Light 6
ATP

Pigment
molecules Photosystem I
Photosystem II (PS I)
(PS II)

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Figure 11.14_5

7 Electron
transport chain
4 Electron Primary
transport chain electron Fd 8
Primary acceptor e– e– NADP+
electron NADP+ + H+
acceptor Pq e–
reductase NADPH
2 H+ e–
2 Cytochrome
H2O
complex
+
1/2 O2 3 Pc
e– P700
1
e–
P680 5 Light
Light 6
ATP

Pigment
molecules Photosystem I
Photosystem II (PS I)
(PS II)

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 3. H2O is split by enzymes, and the electrons are
transferred from the hydrogen atoms to P680+, thus
reducing it to P680
▪ P680+ is the strongest known biological oxidizing agent
▪ The H+ are released into the thylakoid space
▪ O2 is released as a by-product of this reaction

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 4. Each electron “falls” down an electron transport
chain from the primary electron acceptor of PS II to
PS I. Energy released by the fall drives the creation
of a proton gradient across the thylakoid membrane
5. Potential energy stored in the proton gradient drives
production of ATP by chemiosmosis

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 6. In PS I (like PS II), transferred light energy excites
P700, which loses an electron to the primary electron
acceptor
▪ P700+ (P700 that is missing an electron) accepts an
electron passed down from PS II via the electron
transport chain

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 7. Each electron “falls” down an electron transport
chain from the primary electron acceptor of PS I to
the protein ferredoxin (Fd)
8. NADP+ reductase catalyzes the transfer of electrons
to NADP+, reducing it to NADPH
▪ The electrons of NADPH are available for the reactions
of the Calvin cycle
▪ This process also removes an H+ from the stroma

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▪ The energy changes of electrons during linear flow
through the light reactions can be shown in a
mechanical analogy

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Figure 11.15

e–

e– e–
Mill
makes
NADPH
e– ATP
e–
e–

e–

ATP

Photosystem II Photosystem I
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Cyclic Electron Flow

▪ In cyclic electron flow, electrons cycle back from

Fd to the PS I reaction center via a plastocyanin
molecule (Pc)
▪ Cyclic electron flow uses only photosystem I and
produces ATP, but not NADPH
▪ No oxygen is released

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Figure 11.16

Primary
acceptor
Primary Fd
acceptor Fd
NADP+
Pq
NADP+ + H+
reductase
Cytochrome NADPH
complex

Pc

Photosystem I
Photosystem II ATP

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▪ ATP and NADPH are produced on the side facing
the stroma, where the Calvin cycle takes place
▪ In summary, light reactions generate ATP and
increase the potential energy of electrons by moving
them from H2O to NADPH

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Figure 11.UN03

H2O CO2

Light

NADP+
ADP
CALVIN
LIGHT CYCLE
REACTIONS
ATP
NADPH

O2 [CH2O] (sugar)
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Figure 11.18

Cytochrome NADP+
Photosystem II complex Photosystem I reductase
4 H+ Light 3
Light NADP+ + H+
Fd

Pq
NADPH
e– 2 Pc
e–
H2O
1 1/2 O2
THYLAKOID SPACE +2 H+ 4 H+
(high H+ concentration)

CALVIN
CYCLE

Thylakoid
membrane ATP
STROMA synthase
(low H+ concentration) ADP
+ ATP
Pi H+

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Concept 11.3: 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 Calvin cycle is anabolic; it builds sugar from
smaller molecules by using ATP and the reducing
power of electrons carried by NADPH

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

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Figure 11.19_1
Input 3 CO2, entering one per cycle

Phase 1: Carbon fixation

Rubisco
3 P P

3 P P 6 P
RuBP 3-Phosphoglycerate

Calvin
Cycle

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Figure 11.19_2
Input 3 CO2, entering one per cycle

Phase 1: Carbon fixation

Rubisco
3 P P

3 P P 6 P
RuBP 3-Phosphoglycerate 6 ATP

6 ADP

Calvin
Cycle 6 P P
1,3-Bisphosphoglycerate
6 NADPH

6 NADP+
6 Pi

6 P
G3P Phase 2:
Reduction

1 P
Glucose and
G3P other organic
Output compounds
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Figure 11.19_3
Input 3 CO2, entering one per cycle

Phase 1: Carbon fixation

Rubisco
3 P P

3 P P 6 P
RuBP 3-Phosphoglycerate 6 ATP

6 ADP

3 ADP Calvin
3 ATP
Cycle 6 P P
1,3-Bisphosphoglycerate
6 NADPH

Phase 3: 6 NADP+
Regeneration 6 Pi
5 P
of RuBP
G3P 6 P
G3P Phase 2:
Reduction

1 P
Glucose and
G3P other organic
Output compounds
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Figure 11.22b

Chloroplast
H2O CO2

Light

NADP+
ADP 3-Phosphoglycerate
LIGHT
+
REACTIONS:
Photosystem II Pi RuBP CALVIN
CYCLE
Electron transport chain
Photosystem I
ATP
Electron transport chain G3P
NADPH Starch
(storage)

O2 Sucrose (export)
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Figure 11.22c

LIGHT REACTIONS CALVIN CYCLE REACTIONS

• Are carried out by molecules • Take place in the stroma

in the thylakoid membranes • Use ATP and NADPH to convert
• Convert light energy to the chemical CO2 to the sugar G3P
energy of ATP and NADPH • Return ADP, inorganic phosphate,
• Split H2O and release O2 and NADP+ to the light reactions
to the atmosphere

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