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The Process That Feeds the Biosphere

 Plants and other photosynthetic organisms contain

organelles called chloroplasts
Chapter 11  Photosynthesis is the process that converts solar
energy into chemical energy within chloroplasts

Photosynthetic  Directly or indirectly, photosynthesis nourishes

almost the entire living world
Processes

Lecture Presentations by
Nicole Tunbridge and
© 2018 Pearson Education Ltd. Kathleen Fitzpatrick © 2018 Pearson Education Ltd.

Figure 11.1

 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 11.1a

 Photosynthesis occurs in plants, algae, certain other

unicellular eukaryotes, and some prokaryotes

Other organisms also benefit from photosynthesis.

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Figure 11.2

 Heterotrophs obtain organic material from other

organisms
(d) Cyanobacteria 40
µm  Heterotrophs are the consumers of the biosphere
(a) Plants
 Some eat other living organisms; others, called
decomposers, consume dead organic material or
feces
 Almost all heterotrophs, including humans, depend
(b) Multicellular alga on photoautotrophs for food and O2
1 µm

(e) Purple sulfur

bacteria
10 µm

(c) Unicellular eukaryotes

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Figure 11.3

 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

© 2018 Pearson Education Ltd. © 2018 Pearson Education Ltd.

Concept 11.1: Photosynthesis converts light Chloroplasts: The Sites of Photosynthesis

energy to the chemical energy of food in Plants
 Chloroplasts are structurally similar to photosynthetic  Leaves are the major locations of photosynthesis in
bacteria plants
 The structural organization of these organelles  Chloroplasts are found mainly in cells of the
allows for the chemical reactions of photosynthesis 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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Figure 11.4a
Leaf cross section
Chloroplasts Vein

 A chloroplast has an envelope of two membranes Mesophyll

surrounding a dense fluid called the stroma
 Thylakoids are connected sacs in the chloroplast
that compose a third membrane system Stomata
CO2 O2
 Thylakoids may be stacked in columns called grana
Chloroplast Mesophyll
 Chlorophyll, the pigment that gives leaves their cell
green color, resides in the thylakoid membranes

20 µm
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Figure 11.4b
Tracking Atoms Through Photosynthesis:
Scientific Inquiry
 Photosynthesis is a complex series of reactions that
can be summarized as the following equation:
Outer
Thylakoid membrane 6 CO2 + 12 H2O + Light energy → C6H12O6 + 6 O2 + 6 H2O
Thylakoid Intermembrane
Stroma Granum space space
 The overall chemical change during
Inner
membrane
photosynthesis is the reverse of the one that
occurs during cellular respiration

Chloroplast 1 µm
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Figure 11.5
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
Reactants: 6 CO2 12 H2O

Products: C6H12O6 6 H2O 6 O2

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Figure 11.UN01
Photosynthesis as a Redox Process

 Photosynthesis reverses the direction of electron

flow compared to respiration

becomes reduced
 Photosynthesis is a redox process in which H2O is
oxidized and CO2 is reduced
becomes oxidized
 Photosynthesis is an endergonic process; the
energy boost is provided by light

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The Two Stages of Photosynthesis: A Preview The Two Stages of Photosynthesis: A Preview

 Photosynthesis consists of two parts:  The light reactions (in the thylakoids)
 light reactions (the photo part)  Split H2O

 Calvin cycle (the synthesis part)  Release O2

 Reduce the electron acceptor NADP+ to NADPH
 Generate ATP from ADP by photophosphorylation

 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 Figure 11.6_2

Light H2O The light reactions Light H2 O The light reactions

(in the thylakoids) (in the thylakoids)

NADP+ NADP+

ADP ADP
+ +
LIGHT Pi LIGHT Pi
REACTIONS REACTIONS
ATP
Thylakoid Stroma Thylakoid Stroma
NADPH

Chloroplast Chloroplast
O2

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Figure 11.6_3 Figure 11.6_4

Light H2O CO2 Light H2 O CO2

The Calvin cycle The Calvin cycle

(in the stroma) (in the stroma)
NADP+ NADP+

ADP ADP
+ +
LIGHT Pi CALVIN LIGHT Pi CALVIN
REACTIONS CYCLE REACTIONS CYCLE

ATP ATP
Thylakoid Stroma Thylakoid Stroma
NADPH NADPH

Chloroplast Chloroplast
O2 O2 [CH2O]
(sugar)
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Concept 11.2: The light reactions convert solar The Nature of Sunlight
energy to the chemical energy of ATP and
NADPH
 Light is electromagnetic energy, also called
 Chloroplasts are solar-powered chemical factories electromagnetic radiation
 Their thylakoids transform light energy into the  Electromagnetic energy travels in rhythmic waves
chemical energy of ATP and NADPH  Wavelength is the distance between crests
of electromagnetic waves
 Wavelength determines the type of electromagnetic
energy

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Figure 11.7

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

 The electromagnetic spectrum is the entire range Gamma

X-rays UV Infrared
Micro- Radio
rays waves waves
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 Visible light

 Light also behaves as though it consists of discrete

particles, called photons
380 450 500 550 600 650 700 750 nm

Shorter wavelength Longer wavelength

Higher energy Lower energy

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Figure 11.8
Photosynthetic Pigments: The Light Receptors
Light
Reflected
 Pigments are substances that absorb visible light light

 Different pigments absorb different wavelengths Chloroplast

 Wavelengths that are not absorbed are reflected or

transmitted
 Leaves appear green because chlorophyll reflects
and transmits green light

Absorbed Granum
light

Transmitted
light

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Animation: Light Energy and Pigments

 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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Figure 11.9
White Refracting Chlorophyll Photoelectric
light prism solution tube
Galvanometer
2 3
 An absorption spectrum is a graph plotting a 1 4 0 100
pigment’s light absorption versus wavelength

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:  The absorption spectrum of chlorophyll a suggests
 Chlorophyll a, the key light-capturing pigment that violet-blue and red light work best for
photosynthesis
 Chlorophyll b, an accessory pigment
 An action spectrum profiles the relative
 Carotenoids, a separate group of accessory pigments
effectiveness of different wavelengths of radiation in
driving a process

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Figure 11.10 Figure 11.10a

of light by chloroplast

Chloro-
phyll a Chlorophyll b
Absorption

pigments

Chloro-

of light by chloroplast
Carotenoids
phyll a Chlorophyll b
500 400 600 700

Absorption

pigments
Wavelength of light (nm)
(a) Absorption spectra

Carotenoids
Rate of photosynthesis
(measured by O2
release)

400 500 600 700

400 500 600 700
(b) Action spectrum

Aerobic bacteria
Wavelength of light (nm)
Filament of alga
(a) Absorption spectra
Data from T. W. Engelmann, Bacterium photometricum. Ein Beitrag
400 500 600 700 zur vergleichenden Physiologie des Licht-und Farbensinnes, Archiv.
(c) Engelmann’s experiment für Physiologie 30:95–124 (1883).
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 Figure 11.10c

Rate of photosynthesis

Aerobic bacteria
(measured by O2

Filament of alga
release)

400 500 600 700

400 500 600 700 (c) Engelmann’s experiment
Data from T. W. Engelmann, Bacterium photometricum.
(b) Action spectrum Ein Beitrag zur vergleichenden Physiologie des
Licht-und Farbensinnes, Archiv. für Physiologie
Data from T. W. Engelmann, Bacterium photometricum. Ein Beitrag 30:95–124 (1883).
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  The action spectrum for photosynthesis is broader
demonstrated in 1883 by Theodor W. Engelmann than the absorption spectrum of chlorophyll
 In his experiment, he exposed different segments of  Accessory pigments, such as chlorophyll b, broaden
a filamentous alga to different wavelengths the spectrum used for photosynthesis
 Areas receiving wavelengths favorable to  The difference in the absorption spectrum between
photosynthesis produced excess O2 chlorophyll a and b is due to a slight structural
 He used the growth of aerobic bacteria clustered difference between the pigment molecules
along the algae as a measure of O2 production

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Figure 11.11
CH3 in chlorophyll a
CHO in chlorophyll b

CH3

Porphyrin ring:
light-absorbing  Accessory pigments called carotenoids may
“head” of molecule;
note magnesium
broaden the spectrum of colors that drive
atom at center photosynthesis
 Some carotenoids function in photoprotection; they
absorb excessive light that would damage
chlorophyll or react with oxygen

Hydrocarbon tail:
interacts with hydrophobic
regions of proteins inside
thylakoid membranes of
chloroplasts; H atoms not
shown
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Figure 11.12
Excitation of Chlorophyll by Light

Excited
 When a pigment absorbs light, it goes from a ground e– state
state to an excited state, which is unstable

Energy of electron
Heat
 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 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  The light-harvesting complex consists of pigment
complex surrounded by light-harvesting complexes molecules bound to proteins
 The reaction-center complex is an association of  Light-harvesting complexes transfer the energy of
proteins holding a special pair of chlorophyll a photons to the chlorophyll a molecules in the
molecules and a primary electron acceptor reaction-center complex
 These chlorophyll a molecules are special because
they can transfer an excited electron to a different
molecule

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Figure 11.13

Photosystem STROMA
 A primary electron acceptor in the reaction center Photon Light- Reaction-
harvesting center Primary
accepts excited electrons and is reduced as a result complexes complex electron
acceptor
 Solar-powered transfer of an electron from a

Thylakoid membrane
chlorophyll a molecule to the primary electron
Chlorophyll (green)

Thylakoid membrane
acceptor is the first step of the light reactions e–
STROMA

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 Figure 11.13b

Photosystem STROMA
Photon Light- Reaction-
harvesting center Primary
complexes complex electron
acceptor
Chlorophyll (green) STROMA

Thylakoid membrane
Thylakoid membrane

e–

Protein
subunits THYLAKOID
(purple) SPACE
Transfer Special pair of chloro- Pigment (b) Structure of a photosystem
of energy phyll a molecules molecules

THYLAKOID SPACE
(INTERIOR OF THYLAKOID)
(a) How a photosystem harvests light
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 There are two types of photosystems in the thylakoid  Photosystem I (PS I) is best at absorbing a
membrane wavelength of 700 nm
 Photosystem II (PS II) functions first (the numbers  The reaction-center chlorophyll a of PS I is called
reflect order of discovery) P700
 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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Linear Electron Flow

 During the light reactions, there are two possible  There are eight steps in linear electron flow:
routes for electron flow: cyclic and linear 1. A photon hits a pigment in a light-harvesting
 Linear electron flow, the primary pathway, involves complex of PS II, and its energy is passed among
both photosystems and produces ATP and NADPH pigment molecules until it excites P680
using light energy 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 Figure 11.14_1

H2O Primary
CO2 electron
acceptor
Light
e–
2
NADP+
ADP

LIGHT CALVIN
REACTIONS CYCLE

ATP 1 P680
NADPH Light

O2 [CH2O] (sugar)
Pigment
molecules
Photosystem II
(PS II)

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Figure 11.14_2 Figure 11.14_3

4 Electron
transport chain
Primary Primary
electron electron
acceptor acceptor Pq

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

1
e–
P680 1
e–
P680 5
Light Light
ATP

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

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Figure 11.14_4

3. H2O is split by enzymes, and the electrons are 4 Electron Primary

transport chain electron
transferred from the hydrogen atoms to P680+, thus Primary
electron
acceptor

reducing it to P680 acceptor Pq e–

2 H+ e–
2 Cytochrome
H2O
 P680+ is the strongest known biological oxidizing agent +
complex
1/2 O2 3 Pc
 The H+ are released into the thylakoid space e– P700
1
e–
P680 5 Light
 O2 is released as a by-product of this reaction Light 6
ATP

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

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4. Each electron “falls” down an electron transport 6. In PS I (like PS II), transferred light energy excites
chain from the primary electron acceptor of PS II to P700, which loses an electron to the primary electron
PS I. Energy released by the fall drives the creation acceptor
of a proton gradient across the thylakoid membrane  P700+ (P700 that is missing an electron) accepts an
5. Potential energy stored in the proton gradient drives electron passed down from PS II via the electron
production of ATP by chemiosmosis transport chain

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Figure 11.14_5

7 Electron
transport chain
4 Electron Primary 7. Each electron “falls” down an electron transport
transport chain electron Fd 8
Primary acceptor e– e– NADP+ chain from the primary electron acceptor of PS I to
electron NADP+ + H+
acceptor Pq e– reductase NADPH
the protein ferredoxin (Fd)
2 H+ e–
2 Cytochrome
H2O
+
complex
8. NADP+ reductase catalyzes the transfer of electrons
1/2 O2 3
e–
Pc
P700 to NADP+, reducing it to NADPH
1
e–
P680 5 Light
 The electrons of NADPH are available for the reactions
Light 6
of the Calvin cycle
ATP
 This process also removes an H+ from the stroma
Pigment
molecules Photosystem I
Photosystem II (PS I)
(PS II)

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Figure 11.15

e–

 The energy changes of electrons during linear flow

through the light reactions can be shown in a e– e–
Mill
mechanical analogy makes
NADPH
e– ATP
e–
e–

e–
ATP

Photosystem II Photosystem I
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Figure 11.16
Cyclic Electron Flow

 In cyclic electron flow, electrons cycle back from Primary

acceptor
Fd to the PS I reaction center via a plastocyanin Primary
acceptor
Fd
Fd
molecule (Pc) Pq
NADP+
NADP+ + H+
 Cyclic electron flow uses only photosystem I and Cytochrome
reductase
NADPH
produces ATP, but not NADPH complex

 No oxygen is released Pc

Photosystem I
Photosystem II ATP

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A Comparison of Chemiosmosis in
Chloroplasts and Mitochondria
 Some organisms such as purple sulfur bacteria have  Chloroplasts and mitochondria generate ATP by
PS I but not PS II chemiosmosis, but use different sources of energy
 Cyclic electron flow is thought to have evolved  Mitochondria transfer chemical energy from food to
before linear electron flow ATP; chloroplasts transform light energy into the
 Cyclic electron flow may protect cells from light- chemical energy of ATP
induced damage  Spatial organization of chemiosmosis differs
between chloroplasts and mitochondria but also
shows similarities

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Figure 11.17

Mitochondrion Chloroplast

 In mitochondria, protons are pumped to the

intermembrane space and drive ATP synthesis as
they diffuse back into the mitochondrial matrix
Diffusion of
 In chloroplasts, protons are pumped into the Inter- H+ through
ATP synthase Thylakoid
membrane
thylakoid space and drive ATP synthesis as they space H+ space

diffuse back into the stroma Inner

Electron
transport Thylakoid
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  ATP and NADPH are produced on the side facing
Inter-
H+ through the stroma, where the Calvin cycle takes place
ATP synthase Thylakoid
membrane
space
space H+  In summary, light reactions generate ATP and
increase the potential energy of electrons by moving
Electron
Inner transport Thylakoid them from H2O to NADPH
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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Figure 11.UN03 Figure 11.18

H2O CO2
Cytochrome NADP+
Light Photosystem II complex Photosystem I reductase
4 H+ Light 3
Light NADP+ + H+
Fd
NADP+
Pq
ADP NADPH
e–
2 Pc
e–
CALVIN H2O

LIGHT CYCLE THYLAKOID SPACE

1 1/2 O2
+2 H+ 4 H+
(high H+ concentration)
REACTIONS
CALVIN
ATP CYCLE

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

O2 [CH2O] (sugar)
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Figure 11.18a Figure 11.18b

Cytochrome Cytochrome NADP+

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

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

CALVIN
CYCLE

Thylakoid
membrane ATP ATP
STROMA synthase synthase
(low H+ concentration) ADP ADP STROMA
+ ATP + ATP (low H+ concentration)
Pi H+ 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
 Carbon enters the cycle as CO2 and leaves as a
 The Calvin cycle, like the citric acid cycle, sugar named glyceraldehyde 3-phospate (G3P)
regenerates its starting material after molecules  For net synthesis of one G3P, the cycle must take
enter and leave the cycle place three times, fixing three molecules of CO2
 The Calvin cycle is anabolic; it builds sugar from  The Calvin cycle has three phases:
smaller molecules by using ATP and the reducing
1. Carbon fixation (catalyzed by rubisco)
power of electrons carried by NADPH
2. Reduction
3. Regeneration of the CO2 acceptor (RuBP)

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Figure 11.19_1 Figure 11.19_2

Input 3 CO2, entering one per cycle Input 3 CO2, entering one per cycle

Phase 1: Carbon fixation Phase 1: Carbon fixation

Rubisco Rubisco
3 P P 3 P P

3 P P 6 P 3 P P 6 P
RuBP 3-Phosphoglycerate RuBP 3-Phosphoglycerate 6 ATP

6 ADP

Calvin Calvin
Cycle 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 Concept 11.5: Life depends on photosynthesis
Phase 1: Carbon fixation The Importance of Photosynthesis: A Review
Rubisco
3 P P  The energy entering chloroplasts as sunlight gets
3 P P 6 P stored as chemical energy in organic compounds
RuBP 3-Phosphoglycerate 6 ATP

6 ADP  Sugar made in the chloroplasts supplies chemical

3 ADP Calvin energy and carbon skeletons to synthesize the
Cycle 6 P P
3 ATP
1,3-Bisphosphoglycerate organic molecules of cells
6 NADPH
Phase 3:
Regeneration
6 NADP+
6 Pi
 Plants store excess sugar as starch in chloroplasts
of RuBP 5
G3P
P
6 P
and other structures such as roots, tubers, seeds,
G3P Phase 2:
Reduction
and fruits

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

O2 CO2 Chloroplast
H2O CO2

Light
H2O

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

O2 Sucrose (export)
H2O
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Figure 11.23 Figure 11.23a

MAKE CONNECTIONS: The Working Cell

Movement Across Cell Membranes
DNA (Chapter 8)
1
Nucleus
Nuclear
mRNA Energy Transformations in the Cell:
1 DNA
pore Photosynthesis and Cellular
2 Rough endoplasmic
Respiration (Chapters 6, 10, and 11) Nucleus
reticulum (ER)
Protein
Protein
3 in vesicle mRNA
Ribosome mRNA Vacuole Nuclear
pore
4
Vesicle 7 Photosynthesis CO2
2
Golgi in chloroplast
apparatus
forming
Protein
H2O Rough endoplasmic
Plasma 6 Organic
ATP
Transport Protein Protein reticulum (ER)
molecules 8
membrane 5
O2 Cellular respiration ATP
ATP pump
3 in vesicle
11
Flow of Genetic in mitochondrion ATP

Information in the Cell:

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

Cell wall O2
Flow of Genetic Information in the Cell:
H2O
CO2 DNA → RNA → Protein (Chapters 5, 7, 8)

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Figure 11.23b Figure 11.23c

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

10
9 Movement Across Cell
Membranes
(Chapter 8)
Cell wall O2
Energy Transformations
CO2
in the Cell: Photosynthesis
H2O
Flow of Genetic Information in the Cell: and Cellular Respiration
DNA → RNA → Protein (Chapters 5, 7, 8) (Chapters 6, 10, 11)
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