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Chapter 10

Photosynthesis

PowerPoint Lectures for


Biology, Seventh Edition
Neil Campbell and Jane Reece

Lectures by Chris Romero


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Overview: The Process That Feeds the


Biosphere
Photosynthesis
Is the process that converts solar energy into
chemical energy

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Plants and other autotrophs


Are the producers of the biosphere

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Plants are photoautotrophs


They use the energy of sunlight to make
organic molecules from water and carbon
dioxide

Figure 10.1
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Photosynthesis
Occurs in plants, algae, certain other protists,
and some prokaryotes
These organisms use light energy to drive the
synthesis of organic molecules from carbon dioxide
and (in most cases) water. They feed not only
themselves, but the entire living world. (a) On
land, plants are the predominant producers of
food. In aquatic environments, photosynthetic
organisms include (b) multicellular algae, such
as this kelp; (c) some unicellular protists, such
as Euglena; (d) the prokaryotes called
cyanobacteria; and (e) other photosynthetic
prokaryotes, such as these purple sulfur
(a) Plants
bacteria, which produce sulfur (spherical
globules) (c, d, e: LMs).

(c) Unicellular protist 10 m

(e) Pruple sulfur


bacteria

Figure 10.2

(b) Multicellular algae

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(d) Cyanobacteria

40 m

1.5 m

Heterotrophs
Obtain their organic material from other
organisms
Are the consumers of the biosphere

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Concept 10.1: Photosynthesis converts light


energy to the chemical energy of food

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Chloroplasts: The Sites of Photosynthesis in Plants


The leaves of plants
Are the major sites of photosynthesis
Leaf cross section
Vein

Mesophyll

Stomata

Figure 10.3
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CO2

O2

Chloroplasts

Are the organelles in which photosynthesis occurs

Contain thylakoids and grana

Mesophyll

Chloroplast

5 m

Outer
membrane

Stroma Granum

Thylakoid Thylakoid
space

Intermembrane
space

Inner
membrane

1 m

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Tracking Atoms Through Photosynthesis:


Scientific Inquiry
Photosynthesis is summarized as

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

Reactants:

Products:

12 H2O

6 CO2

C6H12O6

Figure 10.4

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6
H2O

6
O2

Photosynthesis as a Redox Process


Photosynthesis is a redox process
Water is oxidized, carbon dioxide is reduced

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The Two Stages of Photosynthesis: A Preview


Photosynthesis consists of two processes
The light reactions

The Calvin cycle

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The light reactions


Occur in the grana

Split water, release oxygen, produce ATP, and


form NADPH

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The Calvin cycle


Occurs in the stroma

Forms sugar from carbon dioxide, using ATP


for energy and NADPH for reducing power

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An overview of photosynthesis

H2O

CO2

Light
NADP
ADP
+ P
LIGHT
REACTIONS

CALVIN
CYCLE
ATP
NADPH

Chloroplast

Figure 10.5

O2

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[CH2O]
(sugar)

Concept 10.2: The light reactions convert solar


energy to the chemical energy of ATP and
NADPH

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The Nature of Sunlight


Light
Is a form of electromagnetic energy, which
travels in waves

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Wavelength
Is the distance between the crests of waves

Determines the type of electromagnetic energy

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The electromagnetic spectrum


Is the entire range of electromagnetic energy, or
radiation
105 nm

103 nm

Gamma
rays

X-rays

UV

1m
106 nm

106 nm

103 nm

1 nm

Infrared

Microwaves

103 m

Radio
waves

Visible light

380

450

500

550

Shorter wavelength

Figure 10.6

Higher energy

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600

650

700
Longer wavelength
Lower energy

750 nm

The visible light spectrum


Includes the colors of light we can see

Includes the wavelengths that drive


photosynthesis

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Photosynthetic Pigments: The Light Receptors


Pigments
Are substances that absorb visible light

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Reflect light, which include the colors we see


Light
Reflected
Light
Chloroplast

Absorbed
light

Granum

Transmitted
light

Figure 10.7
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The spectrophotometer
Is a machine that 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 light absorption versus wavelength
Refracting Chlorophyll
prism
solution

White
light

Photoelectric
tube
Galvanometer
3

100

Slit moves to Green


pass light
light
of selected
wavelength

The high transmittance


(low absorption)
reading indicates that
chlorophyll absorbs
very little green light.

Figure 10.8

Blue
light

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100

The low transmittance


(high absorption) reading
chlorophyll absorbs most blue light.

The absorption spectra of chloroplast pigments


Provide clues to the relative effectiveness of
different wavelengths for driving
photosynthesis

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The absorption spectra of three types of pigments


in chloroplasts
Three different experiments helped reveal which wavelengths of light are photosynthetically
important. The results are shown below.
EXPERIMENT

RESULTS

Absorption of light by
chloroplast pigments

Chlorophyll a
Chlorophyll b

Carotenoids

Wavelength of light (nm)


(a) Absorption spectra. The three curves show the wavelengths of light best absorbed by
three types of chloroplast pigments.
Figure 10.9
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The action spectrum of a pigment

Rate of photosynthesis
(measured by O2 release)

Profiles the relative effectiveness of different


wavelengths of radiation in driving photosynthesis

(b) Action spectrum. This graph plots the rate of photosynthesis versus wavelength.
The resulting action spectrum resembles the absorption spectrum for chlorophyll
a but does not match exactly (see part a). This is partly due to the absorption of light
by accessory pigments such as chlorophyll b and carotenoids.
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The action spectrum for photosynthesis


Was first demonstrated by Theodor W. Engelmann
Aerobic bacteria
Filament
of alga

500
600
700
400
(c) Engelmanns experiment. In 1883, Theodor W. Engelmann illuminated a filamentous alga with light that had
been passed through a prism, exposing different segments of the alga to different wavelengths. He used aerobic
bacteria, which concentrate near an oxygen source, to determine which segments of the alga were releasing the
most O2 and thus photosynthesizing most.
Bacteria congregated in greatest numbers around the parts of the alga illuminated with violet-blue or red light.
Notice the close match of the bacterial distribution to the action spectrum in part b.

CONCLUSION

Light in the violet-blue and red portions of the spectrum are most effective in driving

photosynthesis.
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Chlorophyll a

Is the main photosynthetic pigment

Chlorophyll b

in chlorophyll a

CHO

in chlorophyll b

CH2
CH

CH3

Is an accessory pigment

C
H3C

Mg
N

C
C

H3C

CH3

CH2

H
H

C
O

CH3

CH3

Porphyrin ring:
Light-absorbing
head of molecule
note magnesium
atom at center

C
O

CH2

CH2

O
CH3

CH2

Figure 10.10
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Hydrocarbon tail:
interacts with hydrophobic
regions of proteins inside
thylakoid membranes of
chloroplasts: H atoms not
shown

Other accessory pigments


Absorb different wavelengths of light and pass
the energy to chlorophyll a

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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
e

Excited
state

Heat

Photon
(fluorescence)

Photon

Chlorophyll
molecule

Figure 10.11 A
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Ground
state

If an isolated solution of chlorophyll is


illuminated
It will fluoresce, giving off light and heat

Figure 10.11 B
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A Photosystem: A Reaction Center Associated with


Light-Harvesting Complexes

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A photosystem
Is composed of a reaction center surrounded by a
number of light-harvesting complexes
Thylakoid

Photosystem

Photon

Thylakoid membrane

Light-harvesting
complexes

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STROMA

Primary election
acceptor

Transfer
of energy

Figure 10.12

Reaction
center

Special
chlorophyll a
molecules

Pigment
molecules

THYLAKOID SPACE
(INTERIOR OF THYLAKOID)

The light-harvesting complexes


Consist of pigment molecules bound to
particular proteins
Funnel the energy of photons of light to the
reaction center

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When a reaction-center chlorophyll molecule


absorbs energy
One of its electrons gets bumped up to a
primary electron acceptor

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The thylakoid membrane


Is populated by two types of photosystems, I
and II

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Noncyclic Electron Flow


Noncyclic electron flow
Is the primary pathway of energy
transformation in the light reactions

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Produces NADPH, ATP, and oxygen


H2O

CO2

Light
NADP+
ADP

CALVIN
CYCLE

LIGHT
REACTIONS
ATP
NADPH

O2

[CH2O] (sugar)
Primary
acceptor

Primary
acceptor

Fd
2

2 H+
+
O2

Pq

H2O

NADP+

NADP+
+ 2 H+

reductase

NADPH
PC

+ H+

P700

P680

Light
6
ATP

Figure 10.13

Cytochrome
complex

e
Light
1

Photosystem II
(PS II)

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Photosystem-I
(PS I)

A mechanical analogy for the light reactions


e

ATP

NADPH
e
e

Mill
makes
ATP

Figure 10.14

Photosystem II

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Photosystem I

Cyclic Electron Flow


Under certain conditions
Photoexcited electrons take an alternative path

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In cyclic electron flow


Only photosystem I is used

Only ATP is produced


Primary
acceptor

Primary
acceptor

Fd

Fd

Pq

NADP+
reductase

Cytochrome
complex

NADPH

Pc

Figure 10.15

Photosystem II

ATP

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NADP+

Photosystem I

A Comparison of Chemiosmosis in Chloroplasts


and Mitochondria
Chloroplasts and mitochondria
Generate ATP by the same basic mechanism:
chemiosmosis
But use different sources of energy to
accomplish this

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The spatial organization of chemiosmosis


Differs in chloroplasts and mitochondria
Key
Higher [H+]
Lower [H+]
Chloroplast

Mitochondrion

CHLOROPLAST
STRUCTURE

MITOCHONDRION
STRUCTURE
Intermembrance
space

Membrance

H+ Diffusion
Electron
transport
chain
ATP
Synthase

Matrix

Figure 10.16
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ADP+

Thylakoid
space

Stroma

H+

ATP

In both organelles
Redox reactions of electron transport chains
generate a H+ gradient across a membrane

ATP synthase
Uses this proton-motive force to make ATP

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The light reactions and chemiosmosis: the


organization of the thylakoid membrane
H2O

CO2

LIGHT
NADP+
ADP

LIGHT
REACTOR

CALVIN
CYCLE

ATP

NADPH

STROMA
(Low H+ concentration)

O2

[CH2O] (sugar)

Cytochrome
Photosystem II
complex

Photosystem I
NADP+
reductase

Light

2 H+
Fd

NADP+ + 2H+

NADPH + H+
Pq

Pc
2

H2O

THYLAKOID SPACE
1
(High H+ concentration)

1
2

O2
+2 H+

2 H+

To
Calvin
cycle

STROMA
(Low H+ concentration)

Thylakoid
membrane

ATP
synthase
ADP
ATP
P

Figure 10.17
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H+

Concept 10.3: The Calvin cycle uses ATP and


NADPH to convert CO2 to sugar
The Calvin cycle
Is similar to the citric acid cycle
Occurs in the stroma

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The Calvin cycle has three phases


Carbon fixation

Reduction
Regeneration of the CO2 acceptor

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The Calvin cycle


Light

H2 O

CO2

Input
3 (Entering one
CO2 at a time)

NADP+
ADP

CALVIN
CYCLE

LIGHT
REACTION
ATP

Phase 1: Carbon fixation

NADPH

O2

Rubisco

[CH2O] (sugar)

3 P

3 P

Short-lived
intermediate

Ribulose bisphosphate
(RuBP)

3-Phosphoglycerate

ATP

6 ADP

CALVIN
CYCLE

3 ADP
3

ATP
Phase 3:
Regeneration of
the CO2 acceptor
(RuBP)

6 P

1,3-Bisphoglycerate
6 NADPH
6 NADPH+
6 P
P

(G3P)

Glyceraldehyde-3-phosphate
(G3P)

Figure 10.18
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G3P
(a sugar)
Output

Glucose and
other organic
compounds

Phase 2:
Reduction

Concept 10.4: Alternative mechanisms of


carbon fixation have evolved in hot, arid
climates

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On hot, dry days, plants close their stomata


Conserving water but limiting access to CO2

Causing oxygen to build up

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Photorespiration: An Evolutionary Relic?


In photorespiration
O2 substitutes for CO2 in the active site of the
enzyme rubisco
The photosynthetic rate is reduced

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C4 Plants
C4 plants minimize the cost of photorespiration
By incorporating CO2 into four carbon
compounds in mesophyll cells

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These four carbon compounds


Are exported to bundle sheath cells, where
they release CO2 used in the Calvin cycle

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C4 leaf anatomy and the C4 pathway


Mesophyll
cell

Mesophyll cell
Photosynthetic
cells of C4 plant
leaf

CO
CO
2 2

PEP carboxylase

Bundlesheath
cell
PEP (3 C)
ADP

Oxaloacetate (4 C)
Vein
(vascular tissue)
Malate (4 C)

ATP

C4 leaf anatomy
BundleSheath
cell

Pyruate (3 C)
CO2

Stoma
CALVIN
CYCLE

Sugar

Vascular
tissue

Figure 10.19
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CAM Plants
CAM plants
Open their stomata at night, incorporating CO2
into organic acids

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During the day, the stomata close


And the CO2 is released from the organic acids
for use in the Calvin cycle

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The CAM pathway is similar to the C4 pathway

Pineapple

Sugarcane
C4
Mesophyll Cell

Organic acid
Bundlesheath
cell
(a) Spatial separation
of steps. In C4
plants, carbon fixation
and the Calvin cycle
occur in different
Figure 10.20 types of cells.

CAM

CO2

CALVIN
CYCLE

CO2

1 CO2 incorporated Organic acid


into four-carbon
organic acids
(carbon fixation)
2 Organic acids
release CO2 to
Calvin cycle

Sugar

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CALVIN
CYCLE

Sugar

Night

Day
(b) Temporal separation
of steps. In CAM
plants, carbon fixation
and the Calvin cycle
occur in the same cells
at different times.

The Importance of Photosynthesis: A Review


A review of photosynthesis
Light reaction

Calvin cycle

H2O

CO2

Light
NADP+
ADP
+P1

RuBP

3-Phosphoglycerate

Photosystem II
Electron transport chain
Photosystem I
ATP

NADPH

G3P
Starch
(storage)
Amino acids
Fatty acids

Chloroplast

Figure 10.21

O2

Light reactions:
Are carried out by molecules in the
thylakoid membranes
Convert light energy to the chemical
energy of ATP and NADPH
Split H2O and release O2 to the
atmosphere

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Sucrose (export)

Calvin cycle reactions:


Take place in the stroma
Use ATP and NADPH to convert
CO2 to the sugar G3P
Return ADP, inorganic phosphate,
and
NADP+ to the light reactions

Organic compounds produced by


photosynthesis
Provide the energy and building material for
ecosystems

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