Chapter 10 Photosynthesis.

Chapter 10 Photosynthesis

Overview: The Process That Feeds the Biosphere
Photosynthesis is the process that converts solar energy into chemical energy Directly or indirectly, photosynthesis nourishes almost the entire living world © 2011 Pearson Education, Inc.

Autotrophs sustain themselves without eating anything derived from other organisms
Autotrophs are the producers of the biosphere, producing organic molecules from CO2 and other inorganic molecules Almost all plants are photoautotrophs, using the energy of sunlight to make organic molecules © 2011 Pearson Education, Inc.

Figure 10.1 Figure 10.1 How can sunlight, seen here as a spectrum of colors in a rainbow, power the synthesis of organic substances? 4

(a) Plants Photosynthesis occurs in plants, algae, certain other protists, and some prokaryotes These organisms feed not only themselves but also most of the living world (b) Multicellular alga (d) Cyanobacteria (c) Unicellular protists (e) Purple sulfur bacteria © 2011 Pearson Education, Inc.

Heterotrophs obtain their organic material from other organisms
Heterotrophs are the consumers of the biosphere Almost all heterotrophs, including humans, depend on photoautotrophs for food and O2 © 2011 Pearson Education, Inc.

The Earth’s supply of fossil fuels was formed from the remains of organisms that died hundreds of millions of years ago In a sense, fossil fuels represent stores of solar energy from the distant past © 2011 Pearson Education, Inc.

Concept 10.1: Photosynthesis converts light energy to the chemical energy of food
Chloroplasts are structurally similar to and likely evolved from photosynthetic bacteria The structural organization of these cells allows for the chemical reactions of photosynthesis © 2011 Pearson Education, Inc.

Chloroplasts: The Sites of Photosynthesis in Plants
Leaves are the major locations of photosynthesis Their green color is from chlorophyll, the green pigment within chloroplasts Chloroplasts are found mainly in cells of the mesophyll, the interior tissue of the leaf Each mesophyll cell contains 30–40 chloroplasts © 2011 Pearson Education, Inc.

Chloroplasts also contain stroma, a dense interior fluid
CO2 enters and O2 exits the leaf through microscopic pores called stomata The chlorophyll is in the membranes of thylakoids (connected sacs in the chloroplast); thylakoids may be stacked in columns called grana Chloroplasts also contain stroma, a dense interior fluid © 2011 Pearson Education, Inc.

Leaf cross section Chloroplasts Vein Mesophyll Stomata CO2 O2
Figure 10.4 Leaf cross section Chloroplasts Vein Mesophyll Stomata CO2 O2 Chloroplast Mesophyll cell Outer membrane Figure 10.4 Zooming in on the location of photosynthesis in a plant. Thylakoid Intermembrane space Stroma Granum 20 m Thylakoid space Inner membrane 1 m 11

Tracking Atoms Through Photosynthesis: Scientific Inquiry
Photosynthesis is a complex series of reactions that can be summarized as the following equation: 6 CO H2O + Light energy  C6H12O6 + 6 O2 + 6 H2O © 2011 Pearson Education, Inc.

The Splitting of Water Chloroplasts split H2O into hydrogen and oxygen, incorporating the electrons of hydrogen into sugar molecules and releasing oxygen as a by-product Reactants: 6 CO2 12 H2O Products: C6H12O6 6 H2O 6 O2 © 2011 Pearson Education, Inc.

Photosynthesis as a Redox Process
Photosynthesis reverses the direction of electron flow compared to respiration Photosynthesis is a redox process in which H2O is oxidized and CO2 is reduced Photosynthesis is an endergonic process; the energy boost is provided by light becomes reduced Energy  6 CO2  6 H2O C6 H12 O6  6 O2 becomes oxidized © 2011 Pearson Education, Inc.

The Two Stages of Photosynthesis: A Preview
Photosynthesis consists of the light reactions (the photo part) and Calvin cycle (the synthesis part) The light reactions (in the thylakoids) Split H2O Release O2 Reduce NADP+ to NADPH Generate ATP from ADP by photophosphorylation © 2011 Pearson Education, Inc.

The Calvin cycle (in the stroma) forms sugar from CO2, using ATP and NADPH
The Calvin cycle begins with carbon fixation, incorporating CO2 into organic molecules © 2011 Pearson Education, Inc.

H2O Light NADP ADP + P i Light Reactions Chloroplast Figure 10.6-1
Figure 10.6 An overview of photosynthesis: cooperation of the light reactions and the Calvin cycle. Chloroplast 17

H2O Light NADP ADP + P i Light Reactions ATP NADPH Chloroplast O2
Figure H2O Light NADP ADP + P i Light Reactions ATP Figure 10.6 An overview of photosynthesis: cooperation of the light reactions and the Calvin cycle. NADPH Chloroplast O2 18

Calvin Cycle Light Reactions
Figure H2O CO2 Light NADP ADP + P i Calvin Cycle Light Reactions ATP Figure 10.6 An overview of photosynthesis: cooperation of the light reactions and the Calvin cycle. NADPH Chloroplast O2 19

Calvin Cycle Light Reactions [CH2O] (sugar)
Figure H2O CO2 Light NADP ADP + P i Calvin Cycle Light Reactions ATP Figure 10.6 An overview of photosynthesis: cooperation of the light reactions and the Calvin cycle. NADPH Chloroplast [CH2O] (sugar) O2 20

Concept 10.2: The light reactions convert solar energy to the chemical energy of ATP and NADPH
Chloroplasts are solar-powered chemical factories Their thylakoids transform light energy into the chemical energy of ATP and NADPH © 2011 Pearson Education, Inc.

The Nature of Sunlight Light is a form of electromagnetic energy, also called electromagnetic radiation Like other electromagnetic energy, light travels in rhythmic waves Wavelength is the distance between crests of waves Wavelength determines the type of electromagnetic energy © 2011 Pearson Education, Inc.

The electromagnetic spectrum is the entire range of electromagnetic energy, or radiation
Visible light consists of wavelengths (including those that drive photosynthesis) that produce colors we can see Light also behaves as though it consists of discrete particles, called photons © 2011 Pearson Education, Inc.

Gamma rays Micro- waves Radio waves
Figure 10.7 1 m 105 nm 103 nm 1 nm 103 nm 106 nm (109 nm) 103 m Gamma rays Micro- waves Radio waves X-rays UV Infrared Visible light Figure 10.7 The electromagnetic spectrum. 380 450 500 550 600 650 700 750 nm Shorter wavelength Longer wavelength Higher energy Lower energy 24

Photosynthetic Pigments: The Light Receptors
Pigments are substances that absorb visible light Different pigments absorb different wavelengths Wavelengths that are not absorbed are reflected or transmitted Leaves appear green because chlorophyll reflects and transmits green light Animation: Light and Pigments © 2011 Pearson Education, Inc.

Light Reflected light Chloroplast Absorbed light Granum
Figure 10.8 Light Reflected light Chloroplast Figure 10.8 Why leaves are green: interaction of light with chloroplasts. Absorbed light Granum Transmitted light 26

A spectrophotometer measures a pigment’s ability to absorb various wavelengths
This machine sends light through pigments and measures the fraction of light transmitted at each wavelength © 2011 Pearson Education, Inc.

Slit moves to pass light of selected wavelength. Green light
Figure 10.9 TECHNIQUE Refracting prism Chlorophyll solution Photoelectric tube White light Galvanometer High transmittance (low absorption): Chlorophyll absorbs very little green light. Slit moves to pass light of selected wavelength. Green light Figure 10.9 Research Method: Determining an Absorption Spectrum Low transmittance (high absorption): Chlorophyll absorbs most blue light. Blue light 28

An absorption spectrum is a graph plotting a pigment’s light absorption versus wavelength
The absorption spectrum of chlorophyll a suggests that violet-blue and red light work best for photosynthesis An action spectrum profiles the relative effectiveness of different wavelengths of radiation in driving a process For the Cell Biology Video Space-Filling Model of Chlorophyll a, go to Animation and Video Files. © 2011 Pearson Education, Inc.

Absorption of light by chloroplast pigments
Figure 10.10 RESULTS Chloro- phyll a Chlorophyll b Absorption of light by chloroplast pigments Carotenoids (a) Absorption spectra 400 500 600 700 Wavelength of light (nm) Rate of photosynthesis (measured by O2 release) Figure Inquiry: Which wavelengths of light are most effective in driving photosynthesis? (b) Action spectrum 400 500 600 700 Aerobic bacteria Filament of alga Engelmann’s experiment (c) 400 500 600 700 30

The action spectrum of photosynthesis was first demonstrated in 1883 by Theodor W. Engelmann
In his experiment, he exposed different segments of a filamentous alga to different wavelengths Areas receiving wavelengths favorable to photosynthesis produced excess O2 He used the growth of aerobic bacteria clustered along the alga as a measure of O2 production © 2011 Pearson Education, Inc.

Chlorophyll a is the main photosynthetic pigment
CH3 in chlorophyll a CH3 CHO in chlorophyll b Chlorophyll a is the main photosynthetic pigment Accessory pigments, such as chlorophyll b, broaden the spectrum used for photosynthesis Accessory pigments called carotenoids absorb excessive light that would damage chlorophyll Porphyrin ring Hydrocarbon tail (H atoms not shown) © 2011 Pearson Education, Inc.

Excitation of Chlorophyll by Light
When a pigment absorbs light, it goes from a ground state to an excited state, which is unstable When excited electrons fall back to the ground state, photons are given off, an afterglow called fluorescence If illuminated, an isolated solution of chlorophyll will fluoresce, giving off light and heat © 2011 Pearson Education, Inc.

Photon (fluorescence)
Figure 10.12 Excited state e Heat Energy of electron Photon (fluorescence) Photon Figure Excitation of isolated chlorophyll by light. Ground state Chlorophyll molecule (a) Excitation of isolated chlorophyll molecule (b) Fluorescence 34

A Photosystem: A Reaction-Center Complex Associated with Light-Harvesting Complexes
A photosystem consists of a reaction-center complex (a type of protein complex) surrounded by light-harvesting complexes The light-harvesting complexes (pigment molecules bound to proteins) transfer the energy of photons to the reaction center © 2011 Pearson Education, Inc.

THYLAKOID SPACE (INTERIOR OF THYLAKOID)
Figure 10.13 Photosystem STROMA Photon Light- harvesting complexes Reaction- center complex Primary electron acceptor Chlorophyll STROMA e Thylakoid membrane Thylakoid membrane Figure The structure and function of a photosystem. Transfer of energy Special pair of chlorophyll a molecules Pigment molecules Protein subunits THYLAKOID SPACE (INTERIOR OF THYLAKOID) THYLAKOID SPACE (a) How a photosystem harvests light (b) Structure of photosystem II 36

A primary electron acceptor in the reaction center accepts excited electrons and is reduced as a result Solar-powered transfer of an electron from a chlorophyll a molecule to the primary electron acceptor is the first step of the light reactions © 2011 Pearson Education, Inc.

There are two types of photosystems in the thylakoid membrane
Photosystem II (PS II) functions first (the numbers reflect order of discovery) and is best at absorbing a wavelength of 680 nm The reaction-center chlorophyll a of PS II is called P680 © 2011 Pearson Education, Inc.

Photosystem I (PS I) is best at absorbing a wavelength of 700 nm
The reaction-center chlorophyll a of PS I is called P700 © 2011 Pearson Education, Inc.

Linear Electron Flow During the light reactions, there are two possible routes for electron flow: cyclic and linear Linear electron flow, the primary pathway, involves both photosystems and produces ATP and NADPH using light energy © 2011 Pearson Education, Inc.

Figure A photon hits a pigment and its energy is passed among pigment molecules until it excites P680 An excited electron from P680 is transferred to the primary electron acceptor (we now call it P680+) Primary acceptor 2 e P680 1 Light Figure How linear electron flow during the light reactions generates ATP and NADPH. Pigment molecules Photosystem II (PS II) 41

P680+ is a very strong oxidizing agent
Figure P680+ is a very strong oxidizing agent H2O is split by enzymes, and the electrons are transferred from the hydrogen atoms to P680+, thus reducing it to P680 O2 is released as a by-product of this reaction Primary acceptor 2 e H2O 2 H + 3 1/2 O2 e e P680 1 Light Figure How linear electron flow during the light reactions generates ATP and NADPH. Pigment molecules Photosystem II (PS II) 42

Diffusion of H+ (protons) across the membrane drives ATP synthesis
Figure Each electron “falls” down an electron transport chain from the primary electron acceptor of PS II to PS I Energy released by the fall drives the creation of a proton gradient across the thylakoid membrane Diffusion of H+ (protons) across the membrane drives ATP synthesis Primary acceptor 4 Electron transport chain Pq 2 e H2O 2 H Cytochrome complex + 3 1/2 O2 Pc e e 5 P680 1 Light ATP Figure How linear electron flow during the light reactions generates ATP and NADPH. Pigment molecules Photosystem II (PS II) 43

In PS I (like PS II), transferred light energy excites P700, which loses an electron to an electron acceptor P700+ (P700 that is missing an electron) accepts an electron passed down from PS II via the electron transport chain © 2011 Pearson Education, Inc.

Electron transport chain
Figure Primary acceptor Primary acceptor 4 Electron transport chain Pq e 2 e H2O 2 H Cytochrome complex + 3 1/2 O2 Pc e e P700 5 P680 Light 1 Light 6 ATP Figure How linear electron flow during the light reactions generates ATP and NADPH. Pigment molecules Photosystem I (PS I) Photosystem II (PS II) 45

The electrons are then transferred to NADP+ and reduce it to NADPH
Each electron “falls” down an electron transport chain from the primary electron acceptor of PS I to the protein ferredoxin (Fd) The electrons are then transferred to NADP+ and reduce it to NADPH The electrons of NADPH are available for the reactions of the Calvin cycle This process also removes an H+ from the stroma © 2011 Pearson Education, Inc.

Electron transport chain
Figure Electron transport chain Primary acceptor Primary acceptor 4 7 Electron transport chain Fd Pq e 2 e 8 e e H2O NADP 2 H Cytochrome complex NADP reductase + H + 3 1/2 O2 NADPH Pc e e P700 5 P680 Light 1 Light 6 ATP Figure How linear electron flow during the light reactions generates ATP and NADPH. Pigment molecules Photosystem I (PS I) Photosystem II (PS II) 47

Mill makes ATP NADPH ATP Photosystem II Photosystem I e e e e e
Figure 10.15 e e e Mill makes ATP NADPH e e e Photon Figure A mechanical analogy for linear electron flow during the light reactions. e ATP Photon Photosystem II Photosystem I 48

Cyclic Electron Flow Cyclic electron flow uses only photosystem I and produces ATP, but not NADPH No oxygen is released Cyclic electron flow generates surplus ATP, satisfying the higher demand in the Calvin cycle © 2011 Pearson Education, Inc.

Primary acceptor Primary acceptor Fd Fd NADP + H Pq NADP reductase
Figure 10.16 Primary acceptor Primary acceptor Fd Fd NADP + H Pq NADP reductase Cytochrome complex NADPH Pc Figure Cyclic electron flow. Photosystem I Photosystem II ATP 50

Some organisms such as purple sulfur bacteria have PS I but not PS II
Cyclic electron flow is thought to have evolved before linear electron flow Cyclic electron flow may protect cells from light-induced damage © 2011 Pearson Education, Inc.

Chemiosmosis in Chloroplasts
Chloroplasts generate ATP by chemiosmosis Chloroplasts transform light energy into the chemical energy of ATP In chloroplasts, protons are pumped into the thylakoid space and drive ATP synthesis as they diffuse back into the stroma © 2011 Pearson Education, Inc.

ATP and NADPH are produced on the side facing the stroma, where the Calvin cycle takes place
In summary, light reactions generate ATP and increase the potential energy of electrons by moving them from H2O to NADPH © 2011 Pearson Education, Inc.

STROMA (low H concentration) Cytochrome complex NADP reductase
Figure 10.18 STROMA (low H concentration) Cytochrome complex NADP reductase Photosystem II Photosystem I Light 3 Light 4 H+ NADP + H Fd Pq NADPH 2 Pc H2O 1 1/2 O2 THYLAKOID SPACE (high H concentration) +2 H+ 4 H+ To Calvin Cycle Figure The light reactions and chemiosmosis: the organization of the thylakoid membrane. Thylakoid membrane ATP synthase ADP + P i ATP STROMA (low H concentration) H+ 54

Concept 10.3: The Calvin cycle uses the chemical energy of ATP and NADPH to reduce CO2 to sugar
The Calvin cycle regenerates its starting material after molecules enter and leave the cycle The cycle builds sugar from smaller molecules by using ATP and the reducing power of electrons carried by NADPH © 2011 Pearson Education, Inc.

The Calvin cycle has three phases
Carbon enters the cycle as CO2 and leaves as a sugar named glyceraldehyde 3-phospate (G3P) For net synthesis of 1 G3P, the cycle must take place three times, fixing 3 molecules of CO2 The Calvin cycle has three phases Carbon fixation (catalyzed by rubisco) Reduction Regeneration of the CO2 acceptor (RuBP) © 2011 Pearson Education, Inc.

Figure 10.19 The Calvin cycle.
Input 3 (Entering one at a time) CO2 Phase 1: Carbon fixation Rubisco 3 P P Short-lived intermediate 3 P P 6 P Ribulose bisphosphate (RuBP) 3-Phosphoglycerate Figure The Calvin cycle. 57

Input 3 (Entering one at a time) CO2 Phase 1: Carbon fixation Rubisco 3 P P Short-lived intermediate 3 P P 6 P Ribulose bisphosphate (RuBP) 3-Phosphoglycerate 6 ATP 6 ADP Calvin Cycle 6 P P 1,3-Bisphosphoglycerate 6 NADPH 6 NADP 6 P i Figure The Calvin cycle. 6 P Glyceraldehyde 3-phosphate (G3P) Phase 2: Reduction 1 P G3P (a sugar) Glucose and other organic compounds Output 58

Input 3 (Entering one at a time) CO2 Phase 1: Carbon fixation Rubisco 3 P P Short-lived intermediate 3 P P 6 P Ribulose bisphosphate (RuBP) 3-Phosphoglycerate 6 ATP 6 ADP 3 ADP Calvin Cycle 6 P P 3 ATP 1,3-Bisphosphoglycerate 6 NADPH Phase 3: Regeneration of the CO2 acceptor (RuBP) 6 NADP 6 P i Figure The Calvin cycle. 5 P G3P 6 P Glyceraldehyde 3-phosphate (G3P) Phase 2: Reduction 1 P G3P (a sugar) Glucose and other organic compounds Output 59

Concept 10.4: Alternative mechanisms of carbon fixation have evolved in hot, arid climates
Dehydration is a problem for plants, sometimes requiring trade-offs with other metabolic processes, especially photosynthesis On hot, dry days, plants close stomata, which conserves H2O but also limits photosynthesis The closing of stomata reduces access to CO2 and causes O2 to build up These conditions favor an apparently wasteful process called photorespiration © 2011 Pearson Education, Inc.

Photorespiration: An Evolutionary Relic?
In most plants (C3 plants), initial fixation of CO2, via rubisco, forms a three-carbon compound (3-phosphoglycerate) In photorespiration, rubisco adds O2 instead of CO2 in the Calvin cycle, producing a two-carbon compound Photorespiration consumes O2 and organic fuel and releases CO2 without producing ATP or sugar © 2011 Pearson Education, Inc.

Photorespiration may be an evolutionary relic because rubisco first evolved at a time when the atmosphere had far less O2 and more CO2 Photorespiration limits damaging products of light reactions that build up in the absence of the Calvin cycle In many plants, photorespiration is a problem because on a hot, dry day it can drain as much as 50% of the carbon fixed by the Calvin cycle © 2011 Pearson Education, Inc.

C4 Plants C4 plants minimize the cost of photorespiration by incorporating CO2 into four-carbon compounds in mesophyll cells This step requires the enzyme PEP carboxylase PEP carboxylase has a higher affinity for CO2 than rubisco does; it can fix CO2 even when CO2 concentrations are low These four-carbon compounds are exported to bundle-sheath cells, where they release CO2 that is then used in the Calvin cycle © 2011 Pearson Education, Inc.

Photosynthetic cells of C4 plant leaf PEP carboxylase
Figure 10.20 C4 leaf anatomy The C4 pathway Mesophyll cell Mesophyll cell CO2 Photosynthetic cells of C4 plant leaf PEP carboxylase Bundle- sheath cell Oxaloacetate (4C) PEP (3C) Vein (vascular tissue) ADP Malate (4C) ATP Pyruvate (3C) Bundle- sheath cell Stoma CO2 Calvin Cycle Figure C4 leaf anatomy and the C4 pathway. Sugar Vascular tissue 64

CAM Plants Some plants, including succulents, use crassulacean acid metabolism (CAM) to fix carbon CAM plants open their stomata at night, incorporating CO2 into organic acids Stomata close during the day, and CO2 is released from organic acids and used in the Calvin cycle © 2011 Pearson Education, Inc.

Calvin Cycle Calvin Cycle
Figure 10.21 Sugarcane Pineapple C4 CAM CO2 CO2 1 CO2 incorporated (carbon fixation) Mesophyll cell Organic acid Organic acid Night Figure C4 and CAM photosynthesis compared. CO2 CO2 Bundle- sheath cell 2 CO2 released to the Calvin cycle Day Calvin Cycle Calvin Cycle Sugar Sugar (a) Spatial separation of steps (b) Temporal separation of steps 66

The Importance of Photosynthesis: A Review
The energy entering chloroplasts as sunlight gets stored as chemical energy in organic compounds Sugar made in the chloroplasts supplies chemical energy and carbon skeletons to synthesize the organic molecules of cells Plants store excess sugar as starch in structures such as roots, tubers, seeds, and fruits In addition to food production, photosynthesis produces the O2 in our atmosphere © 2011 Pearson Education, Inc.

H2O CO2 Light NADP ADP + P i Light Reactions: RuBP 3-Phosphoglycerate
Figure 10.22 H2O CO2 Light NADP ADP + P i Light Reactions: Photosystem II Electron transport chain Photosystem I Electron transport chain RuBP 3-Phosphoglycerate Calvin Cycle ATP G3P Figure A review of photosynthesis. Starch (storage) NADPH Chloroplast O2 Sucrose (export) 68

Electron transport chain Electron transport chain
Figure 10.UN02 Primary acceptor Electron transport chain Primary acceptor Electron transport chain Fd NADP + H H2O Pq NADP reductase O2 Cytochrome complex NADPH Pc Figure 10.UN02 Summary figure, Concept 10.2 Photosystem I ATP Photosystem II 69

Regeneration of CO2 acceptor
Figure 10.UN03 3 CO2 Carbon fixation 3  5C 6  3C Calvin Cycle Regeneration of CO2 acceptor 5  3C Figure 10.UN03 Summary figure, Concept 10.3 Reduction 1 G3P (3C) 70

Figure 10.UN04 pH 4 pH 7 pH 4 pH 8 Figure 10.UN04 Test Your Understanding, question 9 ATP 71

Figure 10.UN05 Figure 10.UN05 Appendix A: answer to Figure legend question 72

Figure 10.UN06 Figure 10.UN06 Appendix A: answer to Figure legend question 73

Figure 10.UN07 Figure 10.UN07 Appendix A: answer to Summary of Concept 10.3 Draw It question 74

Figure 10.UN08 Figure 10.UN08 Appendix A: answer to Test Your Understanding, question 9 75

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