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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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Inc.1 Copyright © 2005 Pearson Education. publishing as Benjamin Cummings .• Plants are photoautotrophs – They use the energy of sunlight to make organic molecules from water and carbon dioxide Figure 10.

2 (b) Multicellular algae (d) Cyanobacteria 40 m Copyright © 2005 Pearson Education. (a) On land. such as this kelp. d. but the entire living world. (c) Unicellular protist 10 m (e) Pruple sulfur bacteria 1. (d) the prokaryotes called cyanobacteria. They feed not only themselves. publishing as Benjamin Cummings . e: LMs). certain other protists.• Photosynthesis – Occurs in plants. such as Euglena. photosynthetic organisms include (b) multicellular algae. Inc. plants are the predominant producers of food. such as these purple sulfur (a) Plants bacteria. and (e) other photosynthetic prokaryotes.5 m Figure 10. (c) some unicellular protists. and some prokaryotes These organisms use light energy to drive the synthesis of organic molecules from carbon dioxide and (in most cases) water. algae. which produce sulfur (spherical globules) (c. In aquatic environments.

publishing as Benjamin Cummings .• Heterotrophs – Obtain their organic material from other organisms – Are the consumers of the biosphere Copyright © 2005 Pearson Education. Inc.

• Concept 10. Inc.1: Photosynthesis converts light energy to the chemical energy of food Copyright © 2005 Pearson Education. publishing as Benjamin Cummings .

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

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

Inc.Tracking Atoms Through Photosynthesis: Scientific Inquiry • Photosynthesis is summarized as 6 CO2 + 12 H2O + Light energy  C6H12O6 + 6 O2 + 6 H2 O Copyright © 2005 Pearson Education. publishing as Benjamin Cummings .

4 Copyright © 2005 Pearson Education. incorporating the electrons of hydrogen into sugar molecules Reactants: 6 CO2 12 H2O Products: C6H12O6 6 H2O 6 O2 Figure 10. Inc. publishing as Benjamin Cummings .The Splitting of Water • Chloroplasts split water into – Hydrogen and oxygen.

Inc. carbon dioxide is reduced Copyright © 2005 Pearson Education.Photosynthesis as a Redox Process • Photosynthesis is a redox process – Water is oxidized. publishing as Benjamin Cummings .

The Two Stages of Photosynthesis: A Preview • Photosynthesis consists of two processes – The light reactions – The Calvin cycle Copyright © 2005 Pearson Education. Inc. publishing as Benjamin Cummings .

publishing as Benjamin Cummings . produce ATP.• The light reactions – Occur in the grana – Split water. release oxygen. Inc. and form NADPH Copyright © 2005 Pearson Education.

using ATP for energy and NADPH for reducing power Copyright © 2005 Pearson Education. Inc. publishing as Benjamin Cummings .• The Calvin cycle – Occurs in the stroma – Forms sugar from carbon dioxide.

publishing as Benjamin Cummings .5 O2 [CH2O] (sugar) Copyright © 2005 Pearson Education.• An overview of photosynthesis H2O CO2 Light NADP  ADP + P LIGHT REACTIONS ATP NADPH CALVIN CYCLE Chloroplast Figure 10. Inc.

Inc.• Concept 10. publishing as Benjamin Cummings .2: The light reactions convert solar energy to the chemical energy of ATP and NADPH Copyright © 2005 Pearson Education.

The Nature of Sunlight • Light – Is a form of electromagnetic energy. Inc. which travels in waves Copyright © 2005 Pearson Education. publishing as Benjamin Cummings .

• Wavelength – Is the distance between the crests of waves – Determines the type of electromagnetic energy Copyright © 2005 Pearson Education. Inc. publishing as Benjamin Cummings .

• The electromagnetic spectrum – Is the entire range of electromagnetic energy. Inc. publishing as Benjamin Cummings . 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 Longer wavelength Lower energy 750 nm Shorter wavelength Figure 10.6 Higher energy Copyright © 2005 Pearson Education.

• The visible light spectrum – Includes the colors of light we can see – Includes the wavelengths that drive photosynthesis Copyright © 2005 Pearson Education. Inc. publishing as Benjamin Cummings .

Photosynthetic Pigments: The Light Receptors • Pigments – Are substances that absorb visible light Copyright © 2005 Pearson Education. publishing as Benjamin Cummings . Inc.

publishing as Benjamin Cummings . Inc. which include the colors we see Light Reflected Light Chloroplast Absorbed light Granum Transmitted light Figure 10.– Reflect light.7 Copyright © 2005 Pearson Education.

• The spectrophotometer – Is a machine that sends light through pigments and measures the fraction of light transmitted at each wavelength Copyright © 2005 Pearson Education. Inc. publishing as Benjamin Cummings .

0 100 Figure 10.• 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. Inc. Copyright © 2005 Pearson Education. publishing as Benjamin Cummings .8 Blue light The low transmittance (high absorption) reading chlorophyll absorbs most blue light.

Inc.• The absorption spectra of chloroplast pigments – Provide clues to the relative effectiveness of different wavelengths for driving photosynthesis Copyright © 2005 Pearson Education. publishing as Benjamin Cummings .

• The absorption spectra of three types of pigments in chloroplasts Three different experiments helped reveal which wavelengths of light are photosynthetically important. EXPERIMENT RESULTS Chlorophyll a Absorption of light by chloroplast pigments Chlorophyll b Carotenoids Wavelength of light (nm) (a) Absorption spectra. publishing as Benjamin Cummings .9 Copyright © 2005 Pearson Education. The three curves show the wavelengths of light best absorbed by three types of chloroplast pigments. Figure 10. The results are shown below. Inc.

• 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. Copyright © 2005 Pearson Education. This graph plots the rate of photosynthesis versus wavelength. Inc. 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. publishing as Benjamin Cummings .

In 1883. Engelmann Aerobic bacteria Filament of alga 500 600 700 400 (c) Engelmann‘s experiment. exposing different segments of the alga to different wavelengths. Theodor W. Bacteria congregated in greatest numbers around the parts of the alga illuminated with violet-blue or red light. Copyright © 2005 Pearson Education. Notice the close match of the bacterial distribution to the action spectrum in part b. Inc. He used aerobic bacteria. which concentrate near an oxygen source. publishing as Benjamin Cummings . to determine which segments of the alga were releasing the most O2 and thus photosynthesizing most. Light in the violet-blue and red portions of the spectrum are most effective in driving CONCLUSION photosynthesis.• The action spectrum for photosynthesis – Was first demonstrated by Theodor W. Engelmann illuminated a filamentous alga with light that had been passed through a prism.

10 Copyright © 2005 Pearson Education. publishing as Benjamin Cummings Hydrocarbon tail: interacts with hydrophobic regions of proteins inside thylakoid membranes of chloroplasts: H atoms not shown .• 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. Inc.

Inc. publishing as Benjamin Cummings .• Other accessory pigments – Absorb different wavelengths of light and pass the energy to chlorophyll a Copyright © 2005 Pearson Education.

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. publishing as Benjamin Cummings .11 A Copyright © 2005 Pearson Education. Inc.

11 B Copyright © 2005 Pearson Education. giving off light and heat Figure 10. Inc.• If an isolated solution of chlorophyll is illuminated – It will fluoresce. publishing as Benjamin Cummings .

A Photosystem: A Reaction Center Associated with Light-Harvesting Complexes Copyright © 2005 Pearson Education. publishing as Benjamin Cummings . Inc.

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

Inc.• When a reaction-center chlorophyll molecule absorbs energy – One of its electrons gets bumped up to a primary electron acceptor Copyright © 2005 Pearson Education. publishing as Benjamin Cummings .

I and II Copyright © 2005 Pearson Education. publishing as Benjamin Cummings . Inc.• The thylakoid membrane – Is populated by two types of photosystems.

Noncyclic Electron Flow • Noncyclic electron flow
– Is the primary pathway of energy transformation in the light reactions

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

• 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

7

e

H2O 2 H+ + O2 3 e– Light 1 e–

e

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)

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

• A mechanical analogy for the light reactions
ATP e–

e–

e–

NADPH e– e– Mill makes ATP e–

e–

Figure 10.14

Photosystem II

Photosystem I

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

publishing as Benjamin Cummings .Cyclic Electron Flow • Under certain conditions – Photoexcited electrons take an alternative path Copyright © 2005 Pearson Education. Inc.

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

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

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

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

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

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

Inc. publishing as Benjamin Cummings .• The Calvin cycle has three phases – Carbon fixation – Reduction – Regeneration of the CO2 acceptor Copyright © 2005 Pearson Education.

Inc. publishing as Benjamin Cummings G3P (a sugar) Output Glucose and other organic compounds .3-Bisphoglycerate 6 NADPH 6 NADPH+ 6 P 5 P (G3P) 6 P Glyceraldehyde-3-phosphate (G3P) Phase 2: Reduction 1 P Figure 10.18 Copyright © 2005 Pearson Education.• 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 P Short-lived intermediate 6 P 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 P 1.

Inc.• Concept 10. arid climates Copyright © 2005 Pearson Education.4: Alternative mechanisms of carbon fixation have evolved in hot. publishing as Benjamin Cummings .

• On hot. Inc. publishing as Benjamin Cummings . plants close their stomata – Conserving water but limiting access to CO2 – Causing oxygen to build up Copyright © 2005 Pearson Education. dry days.

Inc.Photorespiration: An Evolutionary Relic? • In photorespiration – O2 substitutes for CO2 in the active site of the enzyme rubisco – The photosynthetic rate is reduced Copyright © 2005 Pearson Education. publishing as Benjamin Cummings .

Inc. publishing as Benjamin Cummings .C4 Plants • C4 plants minimize the cost of photorespiration – By incorporating CO2 into four carbon compounds in mesophyll cells Copyright © 2005 Pearson Education.

publishing as Benjamin Cummings .• These four carbon compounds – Are exported to bundle sheath cells. Inc. where they release CO2 used in the Calvin cycle Copyright © 2005 Pearson Education.

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

Inc.CAM Plants • CAM plants – Open their stomata at night. incorporating CO2 into organic acids Copyright © 2005 Pearson Education. publishing as Benjamin Cummings .

publishing as Benjamin Cummings . the stomata close – And the CO2 is released from the organic acids for use in the Calvin cycle Copyright © 2005 Pearson Education.• During the day. Inc.

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.20 types of cells. publishing as Benjamin Cummings . In C4 plants. In CAM plants. CALVIN CYCLE CALVIN CYCLE Sugar Sugar Copyright © 2005 Pearson Education.• 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. carbon fixation and the Calvin cycle occur in different Figure 10. carbon fixation and the Calvin cycle occur in the same cells at different times. Inc.

inorganic phosphate. and NADP+ to the light reactions Copyright © 2005 Pearson Education. publishing as Benjamin Cummings .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. Inc.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.

Inc.• Organic compounds produced by photosynthesis – Provide the energy and building material for ecosystems Copyright © 2005 Pearson Education. publishing as Benjamin Cummings .