Chapter 11

Photosynthetic
Processes

Lecture Presentations by
Nicole Tunbridge and
Open Rubric
© 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 11.1

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Figure 11.1a

Other organisms also benefit from photosynthesis.

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 Photosynthesis occurs in plants, algae, certain other
unicellular eukaryotes, and some prokaryotes

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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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Figure 11.2a

(a) Plants

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Figure 11.2b

(b) Multicellular alga

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Figure 11.2c

10 µm

(c) Unicellular eukaryotes

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Figure 11.2d

(d) Cyanobacteria
40 µm

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Figure 11.2e

1 µm

(e) Purple sulfur bacteria

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 Heterotrophs obtain organic material from other
organisms
 Heterotrophs are the consumers of the biosphere
 Some eat other living organisms; others, called
decomposers, consume dead organic material or
feces
 Almost all heterotrophs, including humans, depend
on photoautotrophs for food and O2

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 Earth’s supply of fossil fuels was formed from the
remains of organisms that died hundreds of millions
of years ago
 Fossil fuels are being consumed faster then they are
being replenished
 Researchers are exploring methods of using the
photosynthetic process to produce alternative fuels

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Figure 11.3

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Concept 11.1: Photosynthesis converts light
energy to the chemical energy of food
 Chloroplasts are structurally similar to and likely
evolved from photosynthetic bacteria
 The structural organization of these organelles
allows for the chemical reactions of photosynthesis

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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 11.4
Leaf cross section
Chloroplasts Vein

Mesophyll

Stomata
CO2 O2

Chloroplast Mesophyll
cell

Outer
Thylakoid membrane
Thylakoid 20 µm
Stroma Intermembrane
Granum space space
Inner
membrane

Chloroplast 1 µm
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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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Figure 11.4c

Mesophyll
cell

20 µm

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Figure 11.4d

Stroma Granum

Chloroplast 1 µm

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Figure 11.4e

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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 H2 O 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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Animation: Light Energy and Pigments

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 A spectrophotometer measures a pigment’s ability
to absorb various wavelengths
 This machine sends light through pigments and
measures the fraction of light transmitted at each
wavelength

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 An absorption spectrum is a graph plotting a
pigment’s light absorption versus wavelength

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Figure 11.9
White Refracting Chlorophyll Photoelectric
light prism solution tube
Galvanometer
2 3
1 4 0 100

The high transmittance (low

Slit moves to pass light Green absorption) reading indicates
of selected wavelength. light that chlorophyll absorbs
very little green light.

0 100

The low transmittance (high

absorption) reading indicates
Blue
that chlorophyll absorbs
light
most blue 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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 The absorption spectrum of chlorophyll a suggests
that violet-blue and red light work best for
photosynthesis
 An action spectrum profiles the relative
effectiveness of different wavelengths of radiation in
driving a process

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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

Aerobic bacteria
Filament of alga

400 500 600 700

(c) Engelmann’s experiment
Data from T. W. Engelmann, Bacterium photometricum. Ein
Beitrag zur vergleichenden Physiologie des Licht-und
Farbensinnes, Archiv. für Physiologie 30:95–124 (1883).
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Figure 11.10a

of light by chloroplast Chloro-

phyll a Chlorophyll b
Absorption

pigments

Carotenoids

400 500 600 700

Wavelength of light (nm)
(a) Absorption spectra
Data from T. W. Engelmann, Bacterium photometricum. Ein Beitrag
zur vergleichenden Physiologie des Licht-und Farbensinnes, Archiv.
für Physiologie 30:95–124 (1883).

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Figure 11.10b

Rate of photosynthesis
(measured by O2
release)

400 500 600 700

(b) Action spectrum

Data from T. W. Engelmann, Bacterium photometricum. Ein Beitrag
zur vergleichenden Physiologie des Licht-und Farbensinnes, Archiv.
für Physiologie 30:95–124 (1883).

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Figure 11.10c

Aerobic bacteria
Filament of alga

400 500 600 700

(c) Engelmann’s experiment
Data from T. W. Engelmann, Bacterium photometricum.
Ein Beitrag zur vergleichenden Physiologie des
Licht-und Farbensinnes, Archiv. für Physiologie
30:95–124 (1883).

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 The action spectrum of photosynthesis was first
demonstrated in 1883 by Theodor W. Engelmann
 In his experiment, he exposed different segments of
a filamentous alga to different wavelengths
 Areas receiving wavelengths favorable to
photosynthesis produced excess O2
 He used the growth of aerobic bacteria clustered
along the alga as a measure of O2 production

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 The action spectrum for photosynthesis is broader
than the absorption spectrum of chlorophyll
 Accessory pigments, such as chlorophyll b, broaden
the spectrum used for photosynthesis
 The difference in the absorption spectrum between
chlorophyll a and b is due to a slight structural
difference between the pigment molecules

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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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Figure 11.12a

Excited
e– state
Energy of electron

Heat

Photon
(fluorescence)
Photon
Ground
Chlorophyll state
molecule

(a) Excitation of isolated chlorophyll molecule

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Figure 11.12b

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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Figure 11.13a

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

e–

Transfer Special pair of chloro- Pigment

of energy phyll a molecules molecules

THYLAKOID SPACE
(INTERIOR OF THYLAKOID)
(a) How a photosystem harvests light
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Figure 11.13b

Thylakoid membrane Chlorophyll (green) STROMA

Protein
subunits THYLAKOID
(purple) SPACE
(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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 Some organisms such as purple sulfur bacteria have
PS I but not PS II
 Cyclic electron flow is thought to have evolved
before linear electron flow
 Cyclic electron flow may protect cells from light-
induced damage

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A Comparison of Chemiosmosis in
Chloroplasts and Mitochondria
 Chloroplasts and mitochondria generate ATP by
chemiosmosis, but use different sources of energy
 Mitochondria transfer chemical energy from food to
ATP; chloroplasts transform light energy into the
chemical energy of ATP
 Spatial organization of chemiosmosis differs
between chloroplasts and mitochondria but also
shows similarities

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 In mitochondria, protons are pumped to the
intermembrane space and drive ATP synthesis as
they diffuse back into the mitochondrial matrix
 In chloroplasts, protons are pumped into the
thylakoid space and drive ATP synthesis as they
diffuse back into the stroma

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Figure 11.17

Mitochondrion Chloroplast

Diffusion of
Inter- H+ through
membrane ATP synthase Thylakoid
H+ space
space

Electron
Inner Thylakoid
transport
MITOCHONDRION membrane chain membrane CHLOROPLAST
STRUCTURE STRUCTURE
ATP
Pumping synthase
of H+
Matrix by ETC Stroma
ADP + P i
H+ ATP
Higher [H+]
Lower [H+] H+

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Figure 11.17a

MITOCHONDRION CHLOROPLAST
STRUCTURE STRUCTURE
Diffusion of
H+ through
Inter-
ATP synthase Thylakoid
membrane
space
space H+

Electron
Inner transport Thylakoid
membrane chain membrane

ATP
Pumping synthase
of H+
Matrix by ETC Stroma
ADP + P i
H+ ATP
Higher [H+]
Lower [H+] H+

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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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Figure 11.18a

Cytochrome
Photosystem II complex Photosystem I
+ Light
Light 4 H
Fd

Pq

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

Thylakoid
membrane ATP
STROMA synthase
(low H+ concentration) ADP
+ ATP
Pi H+
© 2018 Pearson Education Ltd.
Figure 11.18b

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

Pq
NADPH
2 Pc

THYLAKOID SPACE
4 H+ (high H+ concentration)

CALVIN
CYCLE

ATP
synthase
ADP STROMA
+ ATP (low H+ concentration)
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.UN04a

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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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Concept 11.4: Alternative mechanisms of
carbon fixation have evolved in hot, arid
climates
 Dehydration is a problem for plants, sometimes
requiring trade-offs with other metabolic processes,
especially photosynthesis
 On hot, dry days, plants close stomata, which
conserves H2O but also limits photosynthesis
 The closing of stomata reduces access to CO2 and
causes O2 to build up
 These conditions favor an apparently wasteful
process called photorespiration

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Photorespiration: An Evolutionary Relic?

 In most plants (C3 plants), initial fixation of CO2, via

rubisco, forms a three-carbon compound
(3-phosphoglycerate)
 In photorespiration, rubisco adds O2 instead of
CO2 in the Calvin cycle, producing a two-carbon
compound
 Photorespiration consumes O2 and organic fuel and
releases CO2 without producing ATP or sugar

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 Photorespiration may be an evolutionary relic
because rubisco first evolved at a time when the
atmosphere had far less O2 and more CO2
 Photorespiration limits damaging products of light
reactions that build up in the absence of the Calvin
cycle
 In many plants, photorespiration is a problem
because on a hot, dry day it can drain as much as
50% of the carbon fixed by the Calvin cycle

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C4 Plants

 C4 plants minimize the cost of photorespiration by

incorporating CO2 into four-carbon compounds
 There are two distinct types of cells in the leaves of
C4 plants:
 Bundle-sheath cells are arranged in tightly packed
sheaths around the veins of the leaf
 Mesophyll cells are loosely packed between the
bundle sheath and the leaf surface

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 Sugar production in C4 plants occurs in a three-step
process:
1. The production of the four-carbon precursors is
catalyzed by the enzyme PEP carboxylase in the
mesophyll cells
 PEP carboxylase has a higher affinity for CO2 than
rubisco does; it can fix CO2 even when CO2
concentrations are low

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 2. These four-carbon compounds are exported to
bundle-sheath cells
3. Within the bundle-sheath cells, they release CO2
that is then used in the Calvin cycle

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Figure 11.20

C4 leaf anatomy The C4 pathway

Mesophyll
Photo- Mesophyll cell CO2
synthetic cell PEP carboxylase
cells of Bundle-
C4 plant sheath
leaf cell
Oxaloacetate (4C) PEP (3C)
Vein ADP
(vascular Malate (4C) ATP
tissue)

Pyruvate
Plasmodesma (3C)
CO2
Stoma Bundle-
sheath Calvin
cell Cycle

Sugar

Vascular
tissue

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Figure 11.20a

C4 leaf anatomy

Photo- Mesophyll
synthetic cell
cells of
C4 plant Bundle-
leaf sheath
cell

Vein
(vascular
tissue)

Stoma
© 2018 Pearson Education Ltd.
Figure 11.20b
The C4 pathway
Mesophyll
cell PEP carboxylase CO2

Oxaloacetate (4C) PEP (3C)

ADP
Malate (4C) ATP

Pyruvate
Plasmodesma CO (3C)
2
Bundle-
sheath Calvin
cell Cycle

Sugar

Vascular
tissue
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 Since the Industrial Revolution in the 1800s, CO2
levels have risen greatly
 Increasing levels of CO2 may affect C3 and C4 plants
differently, perhaps changing the relative abundance
of these species
 The effects of such changes are unpredictable and a
cause for concern

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 Suitable agricultural land is decreasing due to the
effects of climate change, while the world demand
for food continues to increase
 C4 photosynthesis uses less water and resources
than C3 photosynthesis
 Scientists have genetically modified rice, a C3 plant,
to carry out C4 photosynthesis
 They estimate 30–50% increase in yield compared
to C3 rice

© 2018 Pearson Education Ltd.

CAM Plants

 Some plants, including succulents, use

crassulacean acid metabolism (CAM) to
fix carbon
 CAM plants open their stomata at night,
incorporating CO2 into organic acids that are stored
in the vacuoles
 Stomata close during the day, and CO2 is released
from organic acids and used in the Calvin cycle

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 The CAM pathway is similar to the C4 pathway in
that they both incorporate CO2 into organic
intermediates before it enters the Calvin cycle
 The C4 pathway structurally separates the initial
steps of carbon fixation from the Calvin cycle
 In the CAM pathway, these steps occur in the same
cell, but are separated in time

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Figure 11.21

Sugarcane Pineapple
1 1
CO2 CO2
C4 CAM
Mesophyll Organic Organic Night
cell acid acid

CO2 2 CO2 2
Bundle-
Day
sheath Calvin Calvin
cell Cycle Cycle

Sugar Sugar

(a) Spatial separation (b) Temporal separation

of steps of steps
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Figure 11.21a

Sugarcane

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Figure 11.21b

Pineapple

© 2018 Pearson Education Ltd.

Concept 11.5: Life depends on photosynthesis
The Importance of Photosynthesis: A Review
 The energy entering chloroplasts as sunlight gets
stored as chemical energy in organic compounds
 Sugar made in the chloroplasts supplies chemical
energy and carbon skeletons to synthesize the
organic molecules of cells
 Plants store excess sugar as starch in chloroplasts
and other structures such as roots, tubers, seeds,
and fruits

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Figure 11.22a

O2 CO2

H2O

Sucrose
(export)

H2O
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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 11.23

MAKE CONNECTIONS: The Working Cell

Movement Across Cell Membranes
DNA (Chapter 8)
1
Nucleus
Nuclear
mRNA Energy Transformations in the Cell:
pore Photosynthesis and Cellular
2 Rough endoplasmic
reticulum (ER)
Respiration (Chapters 6, 10, and 11)
Protein Protein
3 in vesicle
Ribosome mRNA Vacuole

4 7 Photosynthesis
Vesicle CO2
Golgi forming in chloroplast
apparatus H2O
Protein ATP
6 Organic Transport
Plasma
molecules 8 ATP
pump
membrane 5
O2 Cellular respiration ATP
11
Flow of Genetic in mitochondrion ATP

Information in the Cell:

DNA → RNA → Protein
10
(Chapters 5, 7, and 8) 9

Cell wall O2
CO2
H2O

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Figure 11.23a

1 DNA
Nucleus
mRNA
Nuclear
pore
2
Rough endoplasmic
Protein Protein reticulum (ER)
3 in vesicle

Ribosome mRNA
Flow of Genetic Information in the Cell:
DNA → RNA → Protein (Chapters 5, 7, 8)

© 2018 Pearson Education Ltd.

Figure 11.23b

4
Vesicle
Golgi forming
apparatus
Protein
6
Plasma
membrane 5

Cell wall

Flow of Genetic Information in the Cell:

DNA → RNA → Protein (Chapters 5, 7, 8)
© 2018 Pearson Education Ltd.
Figure 11.23c

7 Photosynthesis CO2
in chloroplast
H2O

ATP
Organic Transport
molecules 8 ATP pump

O2 Cellular respiration ATP

11
in mitochondrion ATP

10
9 Movement Across Cell
Membranes
(Chapter 8)
O2
Energy Transformations
CO2
in the Cell: Photosynthesis
H2O
and Cellular Respiration
(Chapters 6, 10, 11)
© 2018 Pearson Education Ltd.
Figure 11.UN04b

Corn plant

Velvetleaf
plant

Corn plant surrounded by

invasive velvetleaf plants

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Figure 11.UN05

Primary
electron Fd
Primary
electron acceptor NADP+
acceptor NADP+ + H+
reductase NADPH
Pq
H2O
O2 Cytochrome
complex

Pc

Photosystem I
ATP
Photosystem II

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Figure 11.UN06
3 CO2

Carbon fixation

3 x 5C 6 x 3C

Calvin
Cycle
Regeneration of
CO2 acceptor
5 x 3C

Reduction

1 G3P (3C)
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Figure 11.UN07

pH 4
pH 7

pH 4 pH 8
ATP

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Figure 11.UN08

© 2018 Pearson Education Ltd.

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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