Chapter 10

Photosynthesis

PowerPoint Lectures for

Biology, Seventh Edition
Neil Campbell and Jane Reece

Lectures by Chris Romero

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
 Overview: The Process That Feeds the Biosphere

• Photosynthesis is the process that converts solar

energy into chemical energy
• Directly or indirectly, photosynthesis nourishes
almost the entire living world

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 • Autotrophs 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 from
water and carbon dioxide

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
 • Photosynthesis occurs in plants, algae, certain
other protists, and some prokaryotes
• These organisms feed not only themselves but
also the entire living world

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LE 10-2

Plants

Unicellular protist 10 µm

Purple sulfur 1.5 µm

bacteria

Multicellular algae Cyanobacteria 40 µm

 • Heterotrophs obtain their organic material from
other organisms
• Heterotrophs are the consumers of the biosphere
• Almost all heterotrophs, including humans,
depend on photoautotrophs for food and oxygen

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 Concept 10.1: Photosynthesis converts light energy
to the chemical energy of food
• Chloroplasts are organelles that are responsible
for feeding the vast majority of organisms
• Chloroplasts are present in a variety of
photosynthesizing organisms

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 Chloroplasts: The Sites of Photosynthesis in Plants

• Leaves are the major locations of photosynthesis

• Their green color is from chlorophyll, the green
pigment within chloroplasts
• Light energy absorbed by chlorophyll drives the
synthesis of organic molecules in the chloroplast
• Through microscopic pores called stomata, CO2
enters the leaf and O2 exits

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 • Chloroplasts are found mainly in cells of the
mesophyll, the interior tissue of the leaf
• A typical mesophyll cell has 30-40 chloroplasts
• The chlorophyll is in the membranes of thylakoids
(connected sacs in the chloroplast); thylakoids
may be stacked in columns called grana
• Chloroplasts also contain stroma, a dense fluid

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LE 10-3
Leaf cross section
Vein

Mesophyll

Stomata CO2 O2

Mesophyll cell
Chloroplast

5 µm

Outer
membrane
Thylakoid
Thylakoid Intermembrane
Stroma Granum space
space
Inner
membrane

1 µm
 Tracking Atoms Through Photosynthesis:
Scientific Inquiry

• Photosynthesis can be summarized as the

following equation:

6 CO2 + 12 H2O + Light energy  C6H12O6 + 6 O2 + 6 H2 O

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 The Splitting of Water

• Chloroplasts split water into hydrogen and oxygen,

incorporating the electrons of hydrogen into sugar
molecules

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LE 10-4

Reactants: 6 CO2 12 H2O

Products: C6H12O6 6 H2O 6 O2

 Photosynthesis as a Redox Process

• Photosynthesis is a redox process in which water

is oxidized and carbon dioxide is reduced

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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 water,
release O2, produce ATP, and form NADPH
• 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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LE 10-5_1

H2O

Light

LIGHT
REACTIONS

Chloroplast
LE 10-5_2

H2O

Light

LIGHT
REACTIONS
ATP

NADPH

Chloroplast

O2
LE 10-5_3

H2O CO2

Light

NADP+
ADP
+ Pi
CALVIN
LIGHT CYCLE
REACTIONS
ATP

NADPH

Chloroplast
[CH2O]
O2
(sugar)
 Concept 10.2: 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 a form of electromagnetic energy, also

called electromagnetic radiation
• Like other electromagnetic energy, light travels in
rhythmic waves
• Wavelength = distance between crests of waves
• Wavelength determines the type of
electromagnetic energy
• Light also behaves as though it consists of
discrete particles, called photons

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 • The electromagnetic spectrum is the entire range
of electromagnetic energy, or radiation
• Visible light consists of colors we can see,
including wavelengths that drive photosynthesis

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LE 10-6

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

Gamma Micro- Radio

X-rays UV Infrared
rays waves waves

Visible light

380 450 500 550 600 650 700 750 nm

Shorter wavelength Longer wavelength

Higher energy Lower energy
 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

Animation: Light and Pigments

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LE 10-7

Light
Reflected
light

Chloroplast

Absorbed Granum
light

Transmitted
light
 • 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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LE 10-8a

White Refracting Chlorophyll Photoelectric

light prism solution tube

Galvanometer

0 100

The high transmittance

Slit moves to Green
(low absorption)
pass light light
reading indicates that
of selected
chlorophyll absorbs
wavelength
very little green light.
LE 10-8b

White Refracting Chlorophyll Photoelectric

light prism solution tube

0 100

Slit moves to The low transmittance

pass light Blue (high absorption)
of selected light reading indicates that
wavelength chlorophyll absorbs
most blue light.
 • An absorption spectrum is a graph plotting a
pigment’s light absorption versus wavelength
• 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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LE 10-9a

Chlorophyll a
Absorption of light by
chloroplast pigments Chlorophyll b

Carotenoids

400 500 600 700

Wavelength of light (nm)
Absorption spectra
 Rate of photo-
LE 10-9b

synthesis (measured
by O2 release)

Action spectrum
 • The action spectrum of photosynthesis was first
demonstrated in 1883 by Thomas 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 aerobic bacteria clustered along the alga
as a measure of O2 production

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LE 10-9c

Aerobic bacteria
Filament
of algae

400 500 600 700

Engelmann’s experiment
 • Chlorophyll a is the main photosynthetic pigment
• Accessory pigments, such as chlorophyll b,
broaden the spectrum used for photosynthesis
• Accessory pigments called carotenoids absorb
excessive light that would damage chlorophyll

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LE 10-10
CH3 in chlorophyll a
CHO in chlorophyll b

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
 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, photons are given off, an afterglow called
fluorescence
• If illuminated, an isolated solution of chlorophyll
will fluoresce, giving off light and heat

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LE 10-11

Excited
e– state

Heat

Photon
(fluorescence)
Photon
Ground
Chlorophyll
molecule state

Excitation of isolated chlorophyll molecule Fluorescence

 A Photosystem: A Reaction Center Associated with
Light-Harvesting Complexes

• A photosystem consists of a reaction center

surrounded by light-harvesting complexes
• The light-harvesting complexes (pigment
molecules bound to proteins) funnel the energy
of photons to the reaction center

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 • A primary electron acceptor in the reaction
center accepts an excited electron from
chlorophyll a
• Solar-powered transfer of an electron from a
chlorophyll a molecule to the primary electron
acceptor is the first step of the light reactions

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

LE 10-12

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

Thylakoid membrane

e–

Transfer Special Pigment

of energy chlorophyll a molecules
molecules
THYLAKOID SPACE
(INTERIOR OF THYLAKOID)
 • There are two types of photosystems in the
thylakoid membrane
• Photosystem II functions first (the numbers reflect
order of discovery) and is best at absorbing a
wavelength of 680 nm
• Photosystem I is best at absorbing a wavelength
of 700 nm
• The two photosystems work together to use light
energy to generate ATP and NADPH

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 Noncyclic Electron Flow

• During the light reactions, there are two possible

routes for electron flow: cyclic and noncyclic
• Noncyclic electron flow, the primary pathway,
involves both photosystems and produces ATP
and NADPH

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

LE 10-13_1
H2O CO2
Light

NADP+
ADP
CALVIN
LIGHT
CYCLE
REACTIONS
ATP

NADPH

O2 [CH2O] (sugar)

Primary
acceptor

e–
Energy of electrons

Light
P680

Photosystem II
(PS II)
LE 10-13_2
H2O CO2
Light

NADP+
ADP
CALVIN
LIGHT
CYCLE
REACTIONS
ATP

NADPH

O2 [CH2O] (sugar)

Primary
acceptor

H2O e–
2 H+
+
Energy of electrons

1/ 2 O2
e–
e–
Light
P680

Photosystem II
(PS II)
LE 10-13_3
H2O CO2
Light

NADP+
ADP
CALVIN
LIGHT
CYCLE
REACTIONS
ATP

NADPH

O2 [CH2O] (sugar)

Primary
acceptor
Pq
H2O e–
2 H+ Cytochrome
complex
+
Energy of electrons

1/ 2 O 2

e– Pc
e–
Light
P680

ATP

Photosystem II
(PS II)
LE 10-13_4
H2O CO2
Light

NADP+
ADP
CALVIN
LIGHT
CYCLE
REACTIONS
ATP

NADPH

O2 [CH2O] (sugar)

Primary
Primary acceptor
acceptor
Pq e–
H2O e–
2 H+ Cytochrome
complex
+
Energy of electrons

1/ 2 O2
e– Pc
e– P700
Light
P680 Light

ATP

Photosystem I
Photosystem II (PS I)
(PS II)
LE 10-13_5
H2 O CO2
Light

NADP+
ADP
CALVIN
LIGHT
CYCLE
REACTIONS
ATP

NADPH

O2 [CH2O] (sugar)

Primary
Primary acceptor
acceptor Fd
Pq e–
e–
H2O e– e– NADP+
2 H+ Cytochrome
NADP+ + 2 H+
complex
+
Energy of electrons

reductase
1/2 O2 NADPH
e– Pc
+ H+
e– P700
Light
P680 Light

ATP

Photosystem I
(PS I)
Photosystem II
(PS II)
LE 10-14

e–
ATP

e– e–

NADPH
e–
e–
e–

Mill
makes
ATP

e–

Photosystem II Photosystem I
 Cyclic Electron Flow

• Cyclic electron flow uses only photosystem I and

produces only ATP
• Cyclic electron flow generates surplus ATP,
satisfying the higher demand in the Calvin cycle

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LE 10-15

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

Pc

Photosystem I
Photosystem II ATP
 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
• The spatial organization of chemiosmosis
differs in chloroplasts and mitochondria

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LE 10-16
Mitochondrion Chloroplast

MITOCHONDRION CHLOROPLAST
STRUCTURE STRUCTURE

H+ Diffusion
Intermembrane Thylakoid
space space

Electron
Membrane transport
chain

Key ATP
synthase
Higher [H+] Matrix Stroma
Lower [H+] ADP + P i
ATP
H+
 • The current model for the thylakoid membrane
is based on studies in several laboratories
• Water is split by photosystem II on the side of
the membrane facing the thylakoid space
• The diffusion of H+ from the thylakoid space
back to the stroma powers ATP synthase
• ATP and NADPH are produced on the side
facing the stroma, where the Calvin cycle takes
place
Animation: Calvin Cycle

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

LE 10-17
H2 O CO2

Light

NADP+
ADP
CALVIN
LIGHT
CYCLE
REACTIONS
ATP

NADPH

O2 [CH2O] (sugar)
STROMA
(Low H+ concentration) Cytochrome
Photosystem II Photosystem I
complex
Light NADP+
Light
2 H+ reductase
Fd NADP+ + 2H+

NADPH + H+
Pq
Pc

H2O
THYLAKOID SPACE 1/2 O2
(High H+ concentration) +2 H+ 2 H+

To
Calvin
cycle

Thylakoid
membrane ATP
STROMA synthase
ADP
(Low H+ concentration) + ATP
Pi
H+
 Concept 10.3: The Calvin cycle uses ATP and
NADPH to convert 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
• 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

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 • The Calvin cycle has three phases:
– Carbon fixation (catalyzed by rubisco)
– Reduction
– Regeneration of the CO2 acceptor (RuBP)

Play

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LE 10-18_1

H2 O CO2
Input
Light

NADP+
3 (Entering one
ADP CO2 at a time)
LIGHT CALVIN
REACTIONS CYCLE

ATP

NADPH
Phase 1: Carbon fixation

Rubisco
O2 [CH2O] (sugar)
3 P P
Short-lived
intermediate
3 P P 6 P
Ribulose bisphosphate 3-Phosphoglycerate
(RuBP) 6 ATP

6 ADP

CALVIN
CYCLE
LE 10-18_2
H2O CO2

Light
Input

NADP+
3 (Entering one
ADP CO2 at a time)
LIGHT CALVIN
REACTIONS CYCLE

ATP

NADPH
Phase 1: Carbon fixation

Rubisco
O2 [CH2O] (sugar)
3 P P
Short-lived
intermediate
3 P P 6 P
Ribulose bisphosphate 3-Phosphoglycerate
(RuBP) 6 ATP

6 ADP

CALVIN
CYCLE
6 P P
1,3-Bisphosphoglycerate
6 NADPH

6 NADP+
6 Pi

6 P
Glyceraldehyde-3-phosphate Phase 2:
(G3P) Reduction

1 P
G3P Glucose and
(a sugar) other organic
Output compounds
LE 10-18_3
H2O CO2

Light
Input

NADP+
3 (Entering one
ADP CO2 at a time)
LIGHT CALVIN
REACTIONS CYCLE

ATP

NADPH
Phase 1: Carbon fixation

Rubisco
O2 [CH2O] (sugar)
3 P P
Short-lived
intermediate
3 P P 6 P
Ribulose bisphosphate 3-Phosphoglycerate
(RuBP) 6 ATP

6 ADP

3 ADP CALVIN
CYCLE
6 P P
3 ATP
1,3-Bisphosphoglycerate
6 NADPH
Phase 3:
Regeneration of 6 NADP+
the CO2 acceptor 6 Pi
(RuBP)
5 P
G3P P
6
Glyceraldehyde-3-phosphate Phase 2:
(G3P) Reduction

1 P
G3P Glucose and
(a sugar) other organic
Output compounds
 Concept 10.4: Alternative mechanisms of carbon
fixation have evolved in hot, arid climates

• Dehydration is a problem for plants, sometimes

requiring tradeoffs with other metabolic processes,
especially photosynthesis
• On hot, dry days, plants close stomata, which
conserves water but also limits photosynthesis
• The closing of stomata reduces access to CO2 and
causes O2 to build up
• These conditions favor a seemingly 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
• In photorespiration, rubisco adds O2 to the Calvin
cycle instead of CO2
• 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
• 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 in
mesophyll cells
• These four-carbon compounds are exported to
bundle-sheath cells, where they release CO2 that
is then used in the Calvin cycle

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LE 10-19

Mesophyll
Mesophyll cell cell CO2
Photosynthetic PEP carboxylase
cells of C4 plant Bundle-
The C4 pathway
leaf sheath
cell
Oxaloacetate (4 C) PEP (3 C)
Vein ADP
(vascular tissue)
Malate (4 C) ATP
C4 leaf anatomy
Pyruvate (3 C)
Bundle-
Stoma sheath CO2
cell
CALVIN
CYCLE

Sugar

Vascular
tissue
 CAM Plants

• CAM plants open their stomata at night,

incorporating CO2 into organic acids
• Stomata close during the day, and CO2 is released
from organic acids and used in the Calvin cycle

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LE 10-20

Sugarcane Pineapple

C4 CAM
CO2 CO2
Mesophyll CO2 incorporated Night
cell Organic acid into four-carbon Organic acid
organic acids
(carbon fixation)
Bundle- CO2 CO2 Day
sheath
cell Organic acids
CALVIN CALVIN
release CO2 to
CYCLE CYCLE
Calvin cycle

Sugar Sugar

Spatial separation of steps Temporal separation of steps

 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
• In addition to food production, photosynthesis
produces the oxygen in our atmosphere

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LE 10-21

Light reactions Calvin cycle

H2O CO2

Light
NADP+
ADP
+ Pi

RuBP 3-Phosphoglycerate
Photosystem II
Electron transport
chain
Photosystem I
ATP G3P
Starch
NADPH (storage)

Chloroplast Amino acids

Fatty acids

O2 Sucrose (export)