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Chapter 10 Photosynthesis PowerPoint Lecture Presentations for Biology Eighth Edition Neil Campbell and Jane Reece Lectures by Chris Romero, updated by Erin Barley with contributions from Joan Sharp Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 1

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 Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 2

Autotrophs sustain themselves without eating anything derived from other organisms Autotrophs are the producers of the biosphere, producing organic molecules from CO 2 and other inorganic molecules Almost all plants are photoautotrophs, using the energy of sunlight to make organic molecules from H 2 O and CO 2 Photosynthesis occurs in plants, algae, certain other protists, and some prokaryotes These organisms feed not only themselves but also most of the living world 3 Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

BioFlix: Photosynthesis (a) Plants (c) Unicellular protist 10 µm (e) Purple sulfur bacteria 1.5 µm (b) Multicellular alga (d) Cyanobacteria 40 µm 4

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 Light energy absorbed by chlorophyll drives the synthesis of organic molecules in the chloroplast CO 2 enters and O 2 exits the leaf through microscopic pores called stomata Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 5

Chloroplasts are found mainly in cells of the mesophyll, the interior tissue of the leaf A typical mesophyll cell has 30 40 chloroplasts There are about 500, 000 chloroplasts per mm 2 of leaf surface. 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 fluid Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 6

Leaf cross section Vein Mesophyll Stomata CO 2 O 2 Chloroplast Mesophyll cell 5 µm 7

Chloroplast Stroma Granum Thylakoid Thylakoid space Inner membrane Outer membrane Intermembrane space 1 µm 8

AN OVERVIEW OF PHOTOSYNTHESIS Light energy 6 CO 2 + 6 H 2 O C 6 H 12 O 6 + 6 O 2 Carbon dioxide Water Glucose Photosynthesis Oxygen gas Photosynthesis can be summarized as the following equation: 6 CO 2 + 12 H 2 O + Light energy C 6 H 12 O 6 + 6 O 2 + 6 H 2 O 9

Chloroplasts split H 2 O into hydrogen and oxygen, incorporating the electrons of hydrogen into sugar molecules Reactants: 6 CO 2 12 H 2 O Products: C 6 H 12 O 6 6 H 2 O 6 O 2 Photosynthesis is a redox process in which H 2 O is oxidized and CO 2 is reduced 10

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 H 2 O Release O 2 Reduce NADP + to NADPH Generate ATP from ADP by photophosphorylation Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 11

The Calvin cycle (in the stroma) forms sugar from CO 2, using ATP and NADPH The Calvin cycle begins with carbon fixation, incorporating CO 2 into organic molecules Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 12

H 2 O CO 2 Light An overview of photosynthesis: cooperation of the light reactions and the Calvin cycle Light Reactions NADP + ADP + P ATP i Calvin Cycle NADPH Chloroplast O 2 [CH 2 O] (sugar) 13

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 Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 14

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 Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 15

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 Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 16

10 5 nm 10 3 nm 1 nm 10 3 nm 10 6 nm 1 m (10 9 nm) 10 3 m Gamma rays X-rays UV Infrared Microwaves Radio waves Visible light 380 450 500 550 600 650 700 750 nm Shorter wavelength Higher energy Longer wavelength Lower energy 17

Photosynthetic Pigments: The Light Receptors Pigments are molecules that absorb visible light Different pigments absorb different wavelengths Wavelengths that are not absorbed are reflected or transmitted Animation: Light and Pigments 18 Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 10-7 Leaves appear green because chlorophyll reflects and transmits green light Chloroplast Light Reflected light Absorbed light Granum Transmitted light 19

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 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 Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 20

RESULTS Chlorophyll a Chlorophyll b Carotenoids (a) Absorption spectra 400 500 600 700 Wavelength of light (nm) (b) Action spectrum Aerobic bacteria Filament of alga (c) Engelmann s experiment 400 500 600 700 21

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 22 Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

CH 3 in chlorophyll a CHO in chlorophyll b Porphyrin ring: light-absorbing head of molecule; note magnesium atom at center Hydrocarbon tail: interacts with hydrophobic regions of proteins inside thylakoid membranes of chloroplasts; H atoms not shown 23

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) funnel the energy of photons to the reaction center Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 24

A primary electron acceptor in the reaction center accepts an excited electron from chlorophyll a Solar-powered transfer of an electron from a chlorophyll a molecule to the primary electron acceptor is the first step of the light reactions 25 Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Thylakoid membrane Fig. 10-12 Photon Light-harvesting complexes Photosystem Reaction-center complex STROMA Primary electron acceptor Transfer of energy Special pair of chlorophyll a molecules Pigment molecules THYLAKOID SPACE (INTERIOR OF THYLAKOID) 26

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 Photosystem I (PS I) is best at absorbing a wavelength of 700 nm The reaction-center chlorophyll a of PS I is called P700 Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 27

Linear Electron Flow During the light reactions, there are two possible routes for electron flow: cyclic and linear Cyclic electron flow uses only photosystem I and produces ATP, but not NADPH Linear electron flow, the primary pathway, involves both photosystems and produces ATP and NADPH using light energy Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 28

A mechanical analogy for the light reactions ATP NADPH Mill makes ATP Photosystem II Photosystem I 29

Primary acceptor 2 A photon hits a pigment and its energy is passed among pigment molecules until it excites P680 1 Light P680 An excited electron from P680 is transferred to the primary electron acceptor Photosystem II (PS II) Pigment molecules How linear electron flow during the light reactions generates ATP and NADPH 30

Fig. 10-13-2 How linear electron flow during the light reactions generates ATP and NADPH 1 / 2 1 2 H + H 2 O + 3 O 2 Light Primary acceptor 2 P680 Photosystem II (PS II) Pigment molecules P680 + (P680 that is missing an electron) is a very strong oxidizing agent H 2 O is split by enzymes, and the electrons are transferred from the hydrogen atoms to P680 +, thus reducing it to P680 O 2 is released as a byproduct of this reaction 31

Fig. 10-13-3 How linear electron flow during the light reactions generates ATP and NADPH 1 / 2 1 2 H + H 2 O + 3 O 2 Light Primary acceptor 2 P680 Photosystem II (PS II) Pq Pigment molecules 4 Cytochrome complex 5 ATP Pc 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 32

Fig. 10-13-4 How linear electron flow during the light reactions generates ATP and NADPH 1 / 2 2 H + H 2 O + 3 O 2 Primary acceptor 2 P680 Pq 4 Cytochrome complex 5 Pc Primary acceptor P700 In PS I (like PS II), transferred light energy excites P700, which loses an electron to an electron acceptor Light 1 Light 6 Photosystem II (PS II) Pigment molecules ATP Photosystem I (PS I) P700 + (P700 that is missing an electron) accepts an electron passed down from PS II via the electron 33 transport chain

Each electron falls down an electron transport chain from the primary electron acceptor of PS I to NADP + and reduce it to NADPH 2 H + H 2 O + 3 1 / 2 O 2 Primary acceptor 2 P680 Pq 4 Cytochrome complex 5 Pc Primary acceptor P700 Fd e 7 Light 8 NADP + reductase NADP + + H + NADPH 1 Light 6 ATP Photosystem II (PS II) Pigment molecules Photosystem I (PS I) The electrons of NADPH are available for the reactions of the Calvin cycle 34

Photon Stroma 1 Photosystem II Electron transport chain Provides energy for synthesis of ATP by chemiosmosis Photon Photosystem I NADP + H + 6 NADPH Thylakoid membrane Primary acceptor 2 4 Primary acceptor 5 P680 P700 Thylakoid space H 2 O 3 1 2 O 2 + 2 H + 35

A mechanical analogy for the light reactions ATP NADPH Mill makes ATP Photosystem II Photosystem I 36

A Comparison of Chemiosmosis in Chloroplasts and Mitochondria Chloroplasts and mitochondria generate ATP by chemiosmosis, but use different sources of energy Mitochondria transfer chemical energy from food to ATP; chloroplasts transform light energy into the chemical energy of ATP Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 37

Chloroplast The production of ATP by chemiosmosis in photosynthesis Stroma (low H + concentration) Light H + Light NADP + + H + NADPH H + H + ADP + P ATP H + Thylakoid membrane H 2 O 1 2 O 2 + 2 H + H + H + Photosystem II Electron transport Thylakoid space chain (high H + concentration) H + H+ H+ Photosystem I H + H + H + H + H + That s PHOTOSYNTESIS!!!!! ATP synthase 38

Oxidative phosphorylation, using electron transport and chemiosmosis in the mitochondrion. H + H + H + H + H + Intermembrane space Protein complex of electron carriers H + Electron carrier H + H + H + ATP synthase Inner mitochondrial membrane Mitochondrial matrix Electron flow NADH FADH 2 FAD NAD + H + H + H + 1 2 H 2 O O 2 + 2 H + ADP + P H + ATP Electron Transport Chain Chemiosmosis OXIDATIVE PHOSPHORYLATION That s CELLULAR RESPIRATION!! 39

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 H 2 O to NADPH Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 40

The Calvin cycle uses ATP and NADPH to convert CO 2 to sugar The Calvin cycle, like the citric acid 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 Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 41

Input 3 (Entering one at a time) CO 2 Phase 1: Carbon fixation Rubisco 3 P Short-lived intermediate 3 P P 6 P Ribulose bisphosphate 3-Phosphoglycerate (RuBP) P 6 6 ADP ATP 3 ATP 3 ADP Calvin Cycle 6 P P 1,3-Bisphosphoglycerate Phase 3: Regeneration of the CO 2 acceptor (RuBP) 5 P G3P 3 phases of Calvin cycle: Carbon fixation (catalyzed by rubisco) Reduction Regeneration of the CO 2 acceptor (RuBP) Output 1 P G3P (a sugar) 6 P Glyceraldehyde-3-phosphate (G3P) Glucose and other organic compounds 6 NADPH 6 NADP + 6 P i Phase 2: Reduction 42

Alternative mechanisms of carbon fixation have evolved in hot, arid climates On hot, dry days, plants close stomata, which conserves H 2 O but also limits photosynthesis The closing of stomata reduces access to CO 2 and causes O 2 to build up These conditions favor a seemingly wasteful process called photorespiration consumes O 2 Photorespiration and organic fuel and releases CO 2 without producing ATP or sugar Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 43

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 O 2 in our atmosphere 44 Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Figure 10.21 Sugarcane Pineapple C 4 CO 2 CAM CO 2 Mesophyll cell Organic acid 1 CO 2 incorporated (carbon fixation) Organic acid Night CO 2 CO 2 Bundlesheath cell Calvin Cycle 2 CO 2 released to the Calvin cycle Calvin Cycle Day Sugar (a) Spatial separation of steps Sugar (b) Temporal separation of steps 45

H 2 O CO 2 Primary acceptor Primary acceptor Fd O 2 H 2 O Pq Cytochrome complex NADP + reductase NADP + + H + NADPH Pc Photosystem II ATP Photosystem I O 2 46

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