Chapter 10

Photosynthesis

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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

1.5 m

Figure 10.2

(b) Multicellular algae

(d) Cyanobacteria

40 m

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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

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

5 m

Outer membrane
Thylakoid Thylakoid space Intermembrane space

Stroma Granum

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:

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
105 nm 103 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 Longer wavelength Lower energy

750 nm

Shorter wavelength

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.

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

CONCLUSION
photosynthesis.

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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 C C O N C C C O C C CH3 C C N CH3 C C C C H CH2 CH3 in chlorophyll a in chlorophyll b

Chlorophyll b
Is an accessory pigment
H3C H H3C C C

C C

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

H H

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) Chlorophyll molecule Ground state

Photon

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

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

H2O Light
NADP+ ADP

CO2

LIGHT REACTIONS
ATP NADPH

CALVIN CYCLE

O2

[CH2O] (sugar) Primary acceptor 4 Fd 2 Pq Cytochrome complex

PC

Primary acceptor

H2O 2 H+ + O2 3 e Light 1 e

8 NADP+ reductase NADPH NADP+ + 2 H+

5 P680

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

NADPH e e Mill makes ATP e

Figure 10.14

Photosystem II

Photosystem I

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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 acceptor Fd Primary acceptor Fd NADP+ NADPH

Pq
Cytochrome complex Pc

NADP+ reductase

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 H+ Diffusion Electron transport chain ATP Synthase ADP+
P

CHLOROPLAST STRUCTURE Thylakoid space

Membrance

Stroma H+

Matrix

ATP

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 NADP+ reductase

Fd 3

Light

NADP+ + 2H+

NADPH + H+ Pq 2 H2O Pc

THYLAKOID SPACE 1 (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
H2 O
NADP+ ADP CO2

LIGHT REACTION
ATP NADPH

CALVIN CYCLE

Input 3 (Entering one CO2 at a time)

Phase 1: Carbon fixation

O2

[CH2O] (sugar)

Rubisco
3 P P

3 P

Short-lived intermediate

Ribulose bisphosphate (RuBP)

3-Phosphoglycerate

6 6 ADP

ATP

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

CALVIN CYCLE

6 P

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

(G3P)

Glyceraldehyde-3-phosphate (G3P)

Phase 2: Reduction

Figure 10.18
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G3P (a sugar) Output

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

Mesophyll cell Photosynthetic cells of C4 plant leaf Bundlesheath cell Oxaloacetate (4 C) Vein (vascular tissue) Malate (4 C) C4 leaf anatomy BundleSheath cell Stoma CALVIN CYCLE Pyruate (3 C) CO2 PEP (3 C) ADP ATP Mesophyll cell PEP carboxylase CO CO 2 2

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

Pineapple
CAM

CO2

CO2 Night

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.

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

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

Calvin cycle CO2

RuBP
Photosystem II Electron transport chain Photosystem I ATP

3-Phosphoglycerate

G3P
Starch (storage) Amino acids Fatty acids

NADPH

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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