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 µm

(e) Pruple sulfur bacteria

1.5 µm

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

Stomata

CO2

O2

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

5 µm

Outer membrane Thylakoid Thylakoid space Intermembrane space

Stroma Granum

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 REACTIONS ATP NADPH CALVIN CYCLE

Chloroplast

Figure 10.5

O2

[CH2O] (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
10–5 nm 10–3 nm 1 nm 103 nm 106 nm 1m 106 nm 103 m

Gamma rays

X-rays

UV

Infrared

Microwaves

Radio waves

Visible light

380

450

500

550

600

650

700

750 nm

Shorter wavelength

Longer wavelength Lower energy

Figure 10.6

Higher energy

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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 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
White light Refracting Chlorophyll prism solution
2 1 4 3 0 100

Photoelectric tube Galvanometer

Slit moves to Green pass light light of selected wavelength

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

0

100

Figure 10.8

Blue light

The low transmittance (high absorption) reading 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
Three different experiments helped reveal which wavelengths of light are photosynthetically important. The results are shown below.
EXPERIMENT

RESULTS
Chlorophyll a Absorption of light by chloroplast pigments 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
– Profiles the relative effectiveness of different wavelengths of radiation in driving photosynthesis
Rate of photosynthesis (measured by O2 release) (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) 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
photosynthesis.

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

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• Chlorophyll a
– Is the main photosynthetic pigment
CH3 CHO CH2 CH H C N Mg C C H C CH2 CH2 C O CH2 O O CH3 N C H H C C O N C C C C C O CH3 C C N CH3 C C C C H CH2 CH3 in chlorophyll a in chlorophyll b

• Chlorophyll b
– Is an accessory pigment
H3 C H H3 C C C

C C

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

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

Energy of election

Heat

Photon (fluorescence) Photon Chlorophyll molecule Ground state

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 Light-harvesting complexes Reaction center Primary election acceptor

STROMA

Thylakoid membrane

e–

Transfer of energy

Special chlorophyll a molecules

Pigment molecules

Figure 10.12
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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
H2 O CO2
NADP+ ADP

Light

LIGHT REACTIONS
ATP NADPH

CALVIN CYCLE

O2

[CH2O] (sugar) Primary acceptor
Elec tr

El Tra ectro ns n ch por ain t

Primary acceptor 2 2 H+ + O2 H2 O 3 e– Light 1 e– P680 e Pq

7 Fd

4
on t rans por t chai n

e

e–

8 NADP reductase
+

Cytochrome complex
PC

NADP+ + 2 H+ NADPH

5

P700 Light 6

+ H+

ATP

Figure 10.13

Photosystem II (PS II)

Photosystem-I (PS I)

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• A mechanical analogy for the light reactions
ATP e–

e–

e– NADPH e– e Mill makes ATP

e–

e–
Photo n

Figure 10.14

Photosystem II

Photosystem I

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on Phot

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

Figure 10.15

Photosystem II

ATP

Photosystem I

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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 STRUCTURE Intermembrance space Membrance H+ Diffusion Electron transport chain ATP Synthase ADP+
P

CHLOROPLAST STRUCTURE Thylakoid space

Stroma H+ ATP

Matrix

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

LIGHT REACTOR

CALVIN CYCLE

ATP

NADPH

STROMA (Low H+ concentration)

O2

[CH2O] (sugar)

Cytochrome Photosystem II complex
2 H+

Photosystem I

Light Fd

NADP+ reductase

3

NADP+ + 2H+

NADPH + H+ Pq H2 O 2
1

Pc

THYLAKOID SPACE (High H+ concentration)

1

⁄2 O2 +2 H+ 2 H+

To Calvin cycle ATP synthase
ADP ATP P H
+

STROMA (Low H+ concentration)

Thylakoid membrane

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
Light
H2O
NADP+ ADP CO2

LIGHT REACTION

CALVIN CYCLE
ATP

Input 3 (Entering one CO2 at a time)
Phase 1: Carbon fixation

NADPH

O2

[CH2O] (sugar)

Rubisco
3 P P

Ribulose bisphosphate (RuBP)

3 P

P

Short-lived intermediate

6

P

3-Phosphoglycerate

6 6 ADP

ATP

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

CALVIN CYCLE

6 P

P

1,3-Bisphoglycerate
6 NADPH
6 NADPH+ 6 P 5 P

(G3P)

6

P

Glyceraldehyde-3-phosphate (G3P)

Phase 2: Reduction

1

Figure 10.18
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G3P (a sugar) Output

P

Glucose and other organic compounds

• 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
Photosynthetic cells of C4 plant leaf Mesophyll cell Bundlesheath cell Mesophyll cell PEP carboxylase CO CO2 2

Oxaloacetate (4 C) Vein (vascular tissue) Malate (4 C) C4 leaf anatomy BundleSheath cell Stoma

PEP (3 C) ADP ATP

Pyruate (3 C) CO2

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
C4 Mesophyll Cell 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.

Pineapple
CAM

CO2

CO2 Night

Organic acid

1 CO2 incorporated Organic acid into four-carbon organic acids (carbon fixation) 2 Organic acids release CO2 to Calvin cycle
CALVIN CYCLE

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

CALVIN CYCLE

Sugar

Sugar

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The Importance of Photosynthesis: A Review • A review of photosynthesis
Light reaction H2O Light
NADP+ ADP +P1 RuBP Photosystem II Electron transport chain Photosystem I ATP NADPH 3-Phosphoglycerate

Calvin cycle CO2

G3P
Starch (storage) Amino acids Fatty acids

Chloroplast

O2

Sucrose (export)

Figure 10.21

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

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

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

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