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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

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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
bacteria, which produce sulfur (spherical (a) Plants
globules) (c, d, e: LMs).

(c) Unicellular protist 10 µm

(e) Pruple sulfur 1.5 µm


bacteria

Figure 10.2 (b) Multicellular algae (d) Cyanobacteria 40 µm

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• 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

CO2 O2
Stomata

Figure 10.3
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• Chloroplasts
– Are the organelles in which photosynthesis occurs
– Contain thylakoids and grana Mesophyll

Chloroplast

5 µm

Outer
membrane

Thylakoid Thylakoid Intermembrane


Stroma Granum space space
Inner
membrane

1 µm
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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: 6 CO2 12 H2O

Products: C6H12O6 6 H2O 6 O2

Figure 10.4

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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 CALVIN
REACTIONS CYCLE

ATP

NADPH

Chloroplast
[CH2O]
O2
Figure 10.5 (sugar)

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• 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
1m
10–5 nm 10–3 nm 1 nm 103 nm 106 nm 106 nm 103 m

Gamma Micro- Radio


rays X-rays UV Infrared waves waves

Visible light

380 450 500 550 600 650 700 750 nm

Shorter wavelength Longer wavelength


Higher energy Lower energy
Figure 10.6
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• 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 Granum
light

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
White Refracting Chlorophyll Photoelectric
light prism solution tube
Galvanometer
2 3

1 0 100
4

Slit moves to Green The high transmittance


pass light light (low absorption)
of selected reading indicates that
wavelength chlorophyll absorbs
very little green light.

0 100

The low transmittance


Blue
(high absorption) reading
light
Figure 10.8 chlorophyll absorbs most blue light.
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• 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
EXPERIMENT Three different experiments helped reveal which wavelengths of light are photosynthetically
important. The results are shown below.

RESULTS

Chlorophyll a

Chlorophyll b
Absorption of light by
chloroplast pigments

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
– Profiles the relative effectiveness of different
wavelengths of radiation in driving photosynthesis
(measured by O2 release)
Rate of 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

400 500 600 700


(c) Engelmann‘s 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
CH3 in chlorophyll a

• Chlorophyll b CH2
CHO in chlorophyll b

CH H CH3

– Is an accessory pigment H3 C C
C
C
C
C
C
C CH2 CH3 Porphyrin ring:
Light-absorbing
C N N C
“head” of molecule
H C Mg C H
note magnesium
H3 C C N N C
atom at center
C C C C CH3
H C C C
CH2 H
H C C
CH2 O O

C O O

O CH3
CH2

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

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• 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
Excited
e– state
Energy of election

Heat

Photon
(fluorescence)

Ground
Photon Chlorophyll state
molecule

Figure 10.11 A
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• 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

Photon Photosystem

STROMA
Light-harvesting Reaction Primary election
complexes center acceptor

Thylakoid membrane

e–

Transfer Special Pigment


of energy chlorophyll a molecules
molecules
THYLAKOID SPACE
Figure 10.12 (INTERIOR OF THYLAKOID)

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• 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 H2 O CO2
Light
NADP+
ADP
LIGHT CALVIN
REACTIONS CYCLE
ATP

NADPH

El
Tra ectro
O2 [CH2O] (sugar) ns n
ch por
ain t
Primary
acceptor 7
Primary
Elec 4
acceptor tron t Fd
rans
Pq por t
2 chai e 8
n e–
H2 O e NADP+
Cytochrome
2 H+ NADP + + 2 H+
complex
+ reductase
3
O2 NADPH
PC
e–
+ H+
e– 5 P700
Light
1 P680 Light

ATP

Photosystem-I
Photosystem II
Figure 10.13 (PS II) (PS I)

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• A mechanical analogy for the light reactions

e–
ATP

e– e–

NADPH
e–
e–
e –

Mill

on
makes

Phot
ATP

e–
n
Photo

Photosystem II Photosystem I
Figure 10.14

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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
Primary acceptor
Fd
acceptor Fd

Pq NADP+
NADP+
reductase
Cytochrome NADPH
complex

Pc

ATP Photosystem I
Photosystem II
Figure 10.15

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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+]
Mitochondrion Chloroplast

MITOCHONDRION CHLOROPLAST
STRUCTURE STRUCTURE
H+ Diffusion
Intermembrance Thylakoid
space Electron space
Membrance transport
chain
ATP
Synthase Stroma
Matrix
ADP+ P
ATP
H+
Figure 10.16

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• 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
CALVIN
LIGHT
CYCLE
REACTOR
ATP
NADPH

STROMA O2 [CH2O] (sugar)


(Low H+ concentration) Cytochrome
Photosystem II complex Photosystem I
Light NADP+
2 H+ reductase
3
Fd NADP+ + 2H+

NADPH + H+
Pq
Pc
2
H2 O 1
⁄2 O2
THYLAKOID SPACE 1 2 H+
+2 H+
(High H+ concentration)
To
Calvin
cycle

ATP
Thylakoid synthase
STROMA membrane ADP
(Low H+ concentration) ATP
P
H +

Figure 10.17
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• 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
H2O CO2
Light Input
NADP+ 3 (Entering one
ADP
CALVIN
CO2 at a time)
LIGHT
REACTION CYCLE
ATP
NADPH Phase 1: Carbon fixation

Rubisco
O2 [CH2O] (sugar)
3 P P

Short-lived
intermediate 6 P
3 P P
Ribulose bisphosphate 3-Phosphoglycerate
(RuBP) 6 ATP
6 ADP

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

1 P
G3P Glucose and
Figure 10.18 (a sugar) other organic
Output compounds
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• 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
Photosynthetic cell CO
CO
PEP carboxylase 2 2
cells of C4 plant Bundle-
leaf sheath
cell

Oxaloacetate (4 C) PEP (3 C)
Vein ADP
(vascular tissue)
Malate (4 C) ATP

C4 leaf anatomy
Bundle- Pyruate (3 C)
Sheath
cell 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

Sugarcane Pineapple

C4 CAM
CO2 CO2
Mesophyll Cell
Night
Organic acid 1 CO2 incorporated Organic acid
Bundle- into four-carbon
sheath organic acids
(carbon fixation) Day
cell
(a) Spatial separation (b) Temporal separation
of steps. In C4 CALVIN 2 Organic acids CALVIN of steps. In CAM
plants, carbon fixation CYCLE release CO2 to CYCLE plants, carbon fixation
and the Calvin cycle Calvin cycle and the Calvin cycle
occur in different occur in the same cells
Sugar Sugar
Figure 10.20 types of cells. at different times.

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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 G3P
NADPH Starch
(storage)
Amino acids
Fatty acids
Chloroplast
O2 Sucrose (export)
Calvin cycle reactions:
Light reactions:
• Take place in the stroma
• Are carried out by molecules in the
• Use ATP and NADPH to convert
thylakoid membranes
CO2 to the sugar G3P
• Convert light energy to the chemical
energy of ATP and NADPH • Return ADP, inorganic phosphate,
• Split H2O and release O2 to the and
Figure 10.21 NADP+ to the light reactions
atmosphere

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• Organic compounds produced by
photosynthesis
– Provide the energy and building material for
ecosystems

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