8 Photosynthesis CAMPBELL BIOLOGY IN FOCUS. Urry Cain Wasserman Minorsky Jackson Reece
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1 CAMPBELL BIOLOGY IN FOCUS Urry Cain Wasserman Minorsky Jackson Reece 8 Photosynthesis Lecture Presentations by Kathleen Fitzpatrick and Nicole Tunbridge
2 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
3 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
4 Figure 8.1
5 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
6 Photosynthesis occurs in plants, algae, certain other protists, and some prokaryotes These organisms feed not only themselves but also most of the living world
7 40 µm Figure 8.2 (d) Cyanobacteria 10 µm (b) Multicellular alga 1 µm (a) Plants (e) Purple sulfur bacteria (c) Unicellular eukaryotes
8 Figure 8.2a (a) Plants
9 Figure 8.2b (b) Multicellular alga
10 10 µm Figure 8.2c (c) Unicellular eukaryotes
11 40 µm Figure 8.2d (d) Cyanobacteria
12 1 µm Figure 8.2e (e) Purple sulfur bacteria
13 Concept 8.1: Photosynthesis converts light energy to the chemical energy of food The structural organization of photosynthetic cells includes enzymes and other molecules grouped together in a membrane This organization allows for the chemical reactions of photosynthesis to proceed efficiently Chloroplasts are structurally similar to and likely evolved from photosynthetic bacteria
14 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 chloroplasts
15 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
16 Figure 8.3 Leaf cross section Chloroplasts Vein Mesophyll Stomata CO2 O2 Chloroplast Mesophyll cell Thylakoid Granum Thylakoid Stroma space Outer membrane Intermembrane space Inner membrane 1 µm 20 µm
17 Figure 8.3a Leaf cross section Chloroplasts Vein Mesophyll Stomata CO2 O2
18 Figure 8.3b Chloroplast Mesophyll cell Thylakoid Granum Thylakoid Stroma space Outer membrane Intermembrane space Inner membrane 1 µm 20 µm
19 Figure 8.3c Mesophyll cell 20 µm
20 Figure 8.3d Granum Stroma 1 µm
21 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
22 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
23 Figure 8.4 Reactants: Products: 6 CO2 C6H12O6 12 H2O 6 H2O 6 O2
24 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
25 Figure 8.UN01 becomes reduced becomes oxidized
26 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 the electron acceptor, NADP+, to NADPH Generate ATP from ADP by adding a phosphate group, photophosphorylation
27 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 Animation: Photosynthesis
28 Figure 8.5 CO2 H2O Light NADP+ ADP + Pi Light Reactions Calvin Cycle ATP NADPH Chloroplast O2 [CH2O] (sugar)
29 Figure H2O Light NADP+ ADP + Pi Light Reactions Chloroplast
30 Figure H2O Light NADP+ ADP + Pi Light Reactions ATP NADPH Chloroplast O2
31 Figure H2O CO2 Light NADP+ ADP + Pi Light Reactions ATP NADPH Chloroplast O2 Calvin Cycle
32 Figure H2O CO2 Light NADP+ ADP + Pi Light Reactions Calvin Cycle ATP NADPH Chloroplast O2 [CH2O] (sugar)
33 Concept 8.2: 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
34 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
35 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
36 Figure nm 10 3 nm Gamma rays 103 nm 1 nm X-rays UV 1m (109 nm) 106 nm Infrared Microwaves 103 m Radio waves Visible light Shorter wavelength Higher energy nm Longer wavelength Lower energy
37 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
38 Figure 8.7 Light Reflected light Chloroplast Absorbed light Granum Transmitted light
39 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
40 Figure 8.8 Technique White light Refracting prism Chlorophyll solution 2 1 Slit moves to pass light of selected wavelength. Galvanometer 3 4 Green light Blue light Photoelectric tube The high transmittance (low absorption) reading indicates that chlorophyll absorbs very little green light. The low transmittance (high absorption) reading indicates that chlorophyll absorbs most blue light.
41 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 Accessory pigments include chlorophyll b and a group of pigments called carotenoids An action spectrum profiles the relative effectiveness of different wavelengths of radiation in driving a process
42 Results Absorption of light by chloroplast pigments Figure 8.9 Chlorophyll a Chlorophyll b Carotenoids Wavelength of light (nm) Rate of photosynthesis (measured by O2 release) (a) Absorption spectra 400 (b) Action spectrum Aerobic bacteria Filament of alga (c) Engelmann s experiment
43 Absorption of light by chloroplast pigments Figure 8.9a Chlorophyll a Chlorophyll b Carotenoids Wavelength of light (nm) (a) Absorption spectra
44 Rate of photosynthesis (measured by O2 release) Figure 8.9b 400 (b) Action spectrum
45 Figure 8.9c Aerobic bacteria Filament of alga (c) Engelmann s experiment 700
46 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
47 Chlorophyll a is the main photosynthetic pigment Accessory pigments, such as chlorophyll b, broaden the spectrum used for photosynthesis A slight structural difference between chlorophyll a and chlorophyll b causes them to absorb slightly different wavelengths Accessory pigments called carotenoids absorb excessive light that would damage chlorophyll Video: Chlorophyll Model
48 Figure 8.10 CH3 CH3 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
49 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
50 Figure 8.11 Energy of electron e Excited state Heat Photon (fluorescence) Photon Chlorophyll molecule Ground state (a) Excitation of isolated chlorophyll molecule (b) Fluorescence
51 Figure 8.11a (b) Fluorescence
52 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
53 Photosystem STROMA LightReactionharvesting center complexes complex Primary electron acceptor Thylakoid membrane Photon e Transfer of energy Special pair of chlorophyll a molecules Pigment molecules THYLAKOID SPACE (INTERIOR OF THYLAKOID) (a) How a photosystem harvests light Thylakoid membrane Figure 8.12 Chlorophyll Protein subunits (b) Structure of a photosystem STROMA THYLAKOID SPACE
54 Figure 8.12a Thylakoid membrane Photon Photosystem STROMA ReactionLightharvesting center complexes complex Primary electron acceptor e Transfer of energy Special pair of chlorophyll a molecules Pigment molecules THYLAKOID SPACE (INTERIOR OF THYLAKOID) (a) How a photosystem harvests light
55 Thylakoid membrane Figure 8.12b Chlorophyll Protein subunits (b) Structure of a photosystem STROMA THYLAKOID SPACE
56 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
57 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
58 Photosystem I (PS I) is best at absorbing a wavelength of 700 nm The reaction-center chlorophyll a of PS I is called P700
59 Linear Electron Flow Linear electron flow involves the flow of electrons through both photosystems to produce ATP and NADPH using light energy
60 Linear electron flow can be broken down into a series of steps 1. A photon hits a pigment and its energy is passed among pigment molecules until it excites P An excited electron from P680 is transferred to the primary electron acceptor (we now call it P680+) 3. 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
61 Figure 8.UN02 H2O CO2 Light NADP+ ADP Calvin Cycle Light Reactions ATP NADPH O2 [CH2O] (sugar)
62 Figure Primary acceptor e 2 P680 1 Light Pigment molecules Photosystem II (PS II)
63 Figure Primary acceptor 2 H+ H O + 3 1/ O 2 e e e P680 Light Pigment molecules Photosystem II (PS II)
64 Figure Primary acceptor 2 H+ H O + 3 1/ O 2 e Electron transport chain Pq Cytochrome complex Pc e e P680 5 Light ATP Pigment molecules Photosystem II (PS II)
65 Figure Primary acceptor 2 H+ H O + 3 1/ O 2 e Electron transport chain Pq e Cytochrome complex Pc e e Primary acceptor P680 P700 5 Light Light 6 ATP Pigment molecules Photosystem II (PS II) Photosystem I (PS I)
66 Figure Primary acceptor + 2H HO + 3 1/ O e Electron transport chain Pq e Cytochrome complex Pc P680 Fd e e e Primary acceptor 7 Electron transport chain 8 e NADP+ reductase P700 5 Light Light 6 ATP Pigment molecules Photosystem II (PS II) Photosystem I (PS I) NADP+ + H+ NADPH
67 4. Each electron falls down an electron transport chain from the primary electron acceptor of PS II to PS I 5. 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
68 6. In PS I (like PS II), transferred light energy excites P700, causing it to lose an electron to an electron acceptor (we now call it P700+) P700+ accepts an electron passed down from PS II via the electron transport chain
69 7. Excited electrons fall down an electron transport chain from the primary electron acceptor of PS I to the protein ferredoxin (Fd) 8. The electrons are transferred to NADP+, reducing it to NADPH, and become available for the reactions of the Calvin cycle This process also removes an H+ from the stroma
70 The energy changes of electrons during linear flow can be represented in a mechanical analogy
71 Figure 8.14 Mill makes ATP Photo n n Photo NADPH Photosystem II Photosystem I
72 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 Spatial organization of chemiosmosis differs between chloroplasts and mitochondria but also shows similarities
73 In mitochondria, protons are pumped to the intermembrane space and drive ATP synthesis as they diffuse back into the mitochondrial matrix In chloroplasts, protons are pumped into the thylakoid space and drive ATP synthesis as they diffuse back into the stroma
74 Figure 8.15 CHLOROPLAST STRUCTURE MITOCHONDRION STRUCTURE Intermembrane space Inner membrane Matrix Key H+ Diffusion Electron transport chain Thylakoid membrane ATP synthase Stroma ADP + P i + Higher [H ] Lower [H+ ] Thylakoid space H + ATP
75 Figure 8.15a CHLOROPLAST STRUCTURE MITOCHONDRION STRUCTURE Intermembrane space Inner membrane Matrix Key H+ Electron transport chain Higher [H ] Lower [H+ ] Thylakoid space Thylakoid membrane ATP synthase ADP + P + Diffusion Stroma i + H ATP
76 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
77 Figure 8.UN02 H2O CO2 Light NADP+ ADP Calvin Cycle Light Reactions ATP NADPH O2 [CH2O] (sugar)
78 Figure 8.16 Photosystem II 4 H+ Light Cytochrome complex Light NADP+ reductase Photosystem I 3 Fd Pq H2O e 1 THYLAKOID SPACE (high H+ concentration) e NADPH Pc 2 / O2 +2 H+ 12 NADP+ + H+ 4 H+ To Calvin Cycle STROMA (low H+ concentration) Thylakoid membrane ATP synthase ADP + P H+ i ATP
79 Figure 8.16a Photosystem II 4 H+ Light Cytochrome complex Light Photosystem I Fd Pq H2O e 1 THYLAKOID SPACE (high H+ concentration) STROMA (low H+ concentration) e Pc 2 /2 O2 +2 H+ 1 Thylakoid membrane 4 H+ ATP synthase ADP + P H+ i ATP
80 Figure 8.16b Cytochrome complex Light NADP+ reductase Photosystem I 3 Fd NADP+ + H+ NADPH Pc 2 4 H+ THYLAKOID SPACE (high H+ concentration) To Calvin Cycle ATP synthase ADP + P H+ i ATP STROMA (low H concentration) +
81 Concept 8.3: The Calvin cycle uses the chemical energy of ATP and NADPH to reduce CO2 to sugar The Calvin cycle, like the citric acid cycle, regenerates its starting material after molecules enter and leave the cycle Unlike the citric acid cycle, the Calvin cycle is anabolic It builds sugar from smaller molecules by using ATP and the reducing power of electrons carried by NADPH
82 Carbon enters the cycle as CO2 and leaves as a sugar named glyceraldehyde 3-phospate (G3P) For net synthesis of one G3P, the cycle must take place three times, fixing three molecules of CO 2 The Calvin cycle has three phases Carbon fixation Reduction Regeneration of the CO2 acceptor
83 Phase 1, carbon fixation, involves the incorporation of the CO2 molecules into ribulose bisphosphate (RuBP) using the enzyme rubisco
84 Figure 8.UN03 H2O CO2 Light NADP+ ADP Calvin Cycle Light Reactions ATP NADPH O2 [CH2O] (sugar)
85 Figure Input 3 as 3 CO2 Phase 1: Carbon fixation Rubisco 3 P 3 P P RuBP 6 P 3-Phosphoglycerate Calvin Cycle P
86 Figure Input 3 as 3 CO2 Phase 1: Carbon fixation Rubisco 3 P 3 P P P 6 P 3-Phosphoglycerate RuBP 6 ATP 6 ADP Calvin Cycle 6 P P 1,3-Bisphosphoglycerate 6 NADPH 6 NADP+ 6 Pi 6 P G3P 1 P G3P Output Phase 2: Reduction Glucose and other organic compounds
87 Figure Input 3 as 3 CO2 Phase 1: Carbon fixation Rubisco 3 P 3 P P P 6 P 3-Phosphoglycerate RuBP 6 ATP 6 ADP Calvin Cycle 3 ADP 3 ATP Phase 3: Regeneration of RuBP P 1,3-Bisphosphoglycerate 6 NADPH 6 NADP+ 6 Pi P 5 G3P 6 P G3P 1 P G3P Output 6 P Phase 2: Reduction Glucose and other organic compounds
88 Phase 2, reduction, involves the reduction and phosphorylation of 3-phosphoglycerate to G3P
89 Phase 3, regeneration, involves the rearrangement of G3P to regenerate the initial CO2 receptor, RuBP
90 Evolution of Alternative Mechanisms of Carbon Fixation in Hot, Arid Climates Adaptation to dehydration is a problem for land 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
91 In most plants (C3 plants), initial fixation of CO2, via rubisco, forms a three-carbon compound (3phosphoglycerate) In photorespiration, rubisco adds O2 instead of CO2 in the Calvin cycle, producing a two-carbon compound Photorespiration decreases photosynthetic output by consuming ATP, O2, and organic fuel and releasing CO2 without producing any ATP or sugar
92 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
93 C4 Plants C4 plants minimize the cost of photorespiration by incorporating CO2 into a four-carbon compound An enzyme in the mesophyll cells has a high affinity for CO2 and can fix carbon 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
94 Figure 8.18 Sugarcane 1 Pineapple CO2 CO2 C4 Mesophyll Organic cell acid Bundlesheath cell CO2 2 CAM Night CO2 2 Calvin Cycle Calvin Cycle Sugar Sugar (a) Spatial separation of steps Organic acid 1 Day (b) Temporal separation of steps
95 Figure 8.18a Sugarcane
96 Figure 8.18b Pineapple
97 Figure 8.18c CO2 C4 Mesophyll cell Bundlesheath cell 1 Organic acid CO2 2 CO2 Calvin Cycle Sugar Sugar CAM Night Organic acid Calvin Cycle (a) Spatial separation of steps CO2 1 2 Day (b) Temporal separation of steps
98 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
99 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 the chloroplasts and in structures such as roots, tubers, seeds, and fruits In addition to food production, photosynthesis produces the O2 in our atmosphere
100 Figure 8.19 H2O CO2 Light NADP+ ADP + P Light Reactions: Photosystem II Electron transport chain Photosystem I Electron transport chain i RuBP 3-Phosphpglycerate Calvin Cycle ATP NADPH G3P Starch (storage) Chloroplast O2 Sucrose (export)
101 Figure 8.UN04
102 Figure 8.UN05 El e O2 t or sp an tr in on a tr ch H2O ec El Primary acceptor Primary acceptor Pq NADP+ reductase Cytochrome complex Pc Photosystem II ATP ct ro n ch tr ai an n sp Fd or Photosystem I t NADP+ + H+ NADPH
103 Figure 8.UN06 3 CO2 Carbon fixation 3 5C 6 3C Calvin Cycle Regeneration of CO2 acceptor 5 3C Reduction 1 G3P (3C)
104 Figure 8.UN07 ph 4 ph 7 ph 4 ph 8
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