BIOLOGY. Photosynthesis CAMPBELL. Reece Urry Cain Wasserman Minorsky Jackson. Lecture Presentation by Nicole Tunbridge and Kathleen Fitzpatrick

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1 CAMPBELL BIOLOGY TENTH EDITION Reece Urry Cain Wasserman Minorsky Jackson 10 Photosynthesis Lecture Presentation by Nicole Tunbridge and Kathleen Fitzpatrick

2 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 CO 2 and other inorganic molecules Almost all plants are photoautotrophs, using the energy of sunlight to make organic molecules

4 Figure 10.1

5 Figure 10.1a Other organisms also benefit from photosynthesis.

6 Photosynthesis occurs in plants, algae, certain other unicellular eukaryotes, and some prokaryotes These organisms feed not only themselves but also most of the living world

7 Figure μm 1 μm (a) Plants (d) Cyanobacteria 40 μm (b) Multicellular alga (e) Purple sulfur bacteria (c) Unicellular eukaryotes

8 Figure 10.2a (a) Plants

9 Figure 10.2b (b) Multicellular alga

10 Figure 10.2c 10 μm (c) Unicellular eukaryotes

11 Figure 10.2d (d) Cyanobacteria 40 μm

12 Figure 10.2e 1 μm (e) Purple sulfur bacteria

13 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 O 2

14 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

15 Figure 10.3

16 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 organelles allows for the chemical reactions of photosynthesis

17 Chloroplasts: The Sites of Photosynthesis in Plants Leaves are the major locations of photosynthesis Chloroplasts are found mainly in cells of the mesophyll, the interior tissue of the leaf Each mesophyll cell contains chloroplasts CO 2 enters and O 2 exits the leaf through microscopic pores called stomata

18 A chloroplast has an envelope of two membranes surrounding a dense fluid called the stroma Thylakoids are connected sacs in the chloroplast which compose a third membrane system Thylakoids may be stacked in columns called grana Chlorophyll, the pigment which gives leaves their green colour, resides in the thylakoid membranes

19 Figure 10.4 Leaf cross section Chloroplasts Vein Mesophyll Stomata CO 2 O 2 Chloroplast Mesophyll cell Thylakoid Thylakoid Stroma Granum space Outer membrane Intermembrane space Inner membrane 20 μm 1 μm

20 Figure 10.4a Leaf cross section Chloroplasts Vein Mesophyll Stomata Chloroplast CO 2 O 2 Mesophyll cell 20 μm

21 Figure 10.4b Chloroplast Thylakoid Thylakoid Stroma Granum space Outer membrane Intermembrane space Inner membrane 1 μm

22 Figure 10.4c Stroma Granum 1 μm

23 Figure 10.4d Mesophyll cell 20 μm

24 Tracking Atoms Through Photosynthesis: Scientific Inquiry Photosynthesis is a complex series of reactions that can be summarized as the following equation: 6 CO H 2 O + Light energy C 6 H 12 O O H 2 O The overall chemical change during photosynthesis is the reverse of the one that occurs during cellular respiration

25 The Splitting of Water Chloroplasts split H 2 O into hydrogen and oxygen, incorporating the electrons of hydrogen into sugar molecules and releasing oxygen as a by-product

26 Figure 10.5 Reactants: 6 CO 2 12 H 2 O Products: C 6 H 12 O 6 6 H 2 O 6 O 2

27 Photosynthesis as a Redox Process Photosynthesis reverses the direction of electron flow compared to respiration Photosynthesis is a redox process in which H 2 O is oxidized and CO 2 is reduced Photosynthesis is an endergonic process; the energy boost is provided by light

28 Figure 10.UN01 becomes reduced becomes oxidized

29 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 the electron acceptor NADP + to NADPH Generate ATP from ADP by photophosphorylation

30 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

31 Figure Light H 2 O NADP + LIGHT REACTIONS ADP + P i Thylakoid Stroma Chloroplast

32 Figure Light H 2 O NADP + Thylakoid LIGHT REACTIONS ADP + P i ATP NADPH Stroma Chloroplast O 2

33 Figure Light H 2 O CO 2 NADP + Thylakoid LIGHT REACTIONS ADP + P i ATP NADPH CALVIN CYCLE Stroma Chloroplast O 2

34 Figure Light H 2 O CO 2 NADP + Thylakoid LIGHT REACTIONS ADP + P i ATP NADPH CALVIN CYCLE Stroma Chloroplast O 2 [CH 2 O] (sugar)

35 BioFlix: The Carbon Cycle

36 BioFlix: Photosynthesis

37 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

38 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

39 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

40 Figure m 10 5 nm 10 3 nm 1 nm 10 3 nm 10 6 nm (109 nm) 10 3 m Gamma rays X-rays UV Infrared Microwaves Radio waves Visible light nm Shorter wavelength Higher energy Longer wavelength Lower energy

41 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

42 Figure 10.8 Light Reflected light Chloroplast Absorbed light Granum Transmitted light

43 Animation: Light and Pigments

44 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

45 Figure 10.9 White light Refracting prism Chlorophyll solution 2 3 Photoelectric tube Galvanometer 1 4 Slit moves to pass light of selected wavelength. Green light The high transmittance (low absorption) reading indicates that chlorophyll absorbs very little green light. Blue light The low transmittance (high absorption) reading indicates that chlorophyll absorbs most blue light.

46 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

47 Absorption of light by chloroplast pigments release) Figure Chlorophyll a Chlorophyll b Carotenoids Wavelength of light (nm) (a) Absorption spectra Rate of photosynthesis (measured by O 2 (b) Action spectrum Aerobic bacteria Filament of alga (c) Engelmann s experiment

48 Figure 10.10a Absorption of light by chloroplast pigments Chlorophyll a Chlorophyll b Carotenoids Wavelength of light (nm) (a) Absorption spectra

49 Figure 10.10b release) Rate of photosynthesis (measured by O 2 (b) Action spectrum

50 Figure 10.10c Aerobic bacteria Filament of alga (c) Engelmann s experiment

51 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 O 2 He used the growth of aerobic bacteria clustered along the alga as a measure of O 2 production

52 Chlorophyll a is the main photosynthetic pigment Accessory pigments, such as chlorophyll b, broaden the spectrum used for photosynthesis The difference in the absorption spectrum between chlorophyll a and b is due to a slight structural difference between the pigment molecules Accessory pigments called carotenoids absorb excessive light that would damage chlorophyll

53 Figure CH 3 CH3 CHO in chlorophyll a 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

54 Video: Space-Filling Model of Chlorophyll a

55 Accessory pigments called carotenoids function in photoprotection; they absorb excessive light that would damage chlorophyll

56 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

57 Figure Energy of electron e Excited state Heat Photon Chlorophyll molecule Photon (fluorescence) Ground state (a) Excitation of isolated chlorophyll molecule (b) Fluorescence

58 Figure 10.12a (b) Fluorescence

59 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

60 Figure Thylakoid membrane Thylakoid membrane Photosystem Photon Lightharvesting complexes Reactioncenter complex STROMA Primary electron acceptor e Chlorophyll STROMA Transfer of energy Special pair of chlorophyll a molecules THYLAKOID SPACE (INTERIOR OF THYLAKOID) (a) How a photosystem harvests light Pigment molecules Protein subunits (b) Structure of a photosystem THYLA- KOID SPACE

61 Thylakoid membrane Figure 10.13a Photosystem Photon Lightharvesting complexes Reactioncenter complex STROMA Primary electron acceptor e Transfer of energy Special pair of chlorophyll a molecules THYLAKOID SPACE (INTERIOR OF THYLAKOID) (a) How a photosystem harvests light Pigment molecules

62 Figure 10.13b Thylakoid membrane Chlorophyll STROMA Protein subunits (b) Structure of a photosystem THYLA- KOID SPACE

63 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

64 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

65 Photosystem I (PS I) is best at absorbing a wavelength of 700 nm The reaction-center chlorophyll a of PS I is called P700

66 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

67 There are 8 steps in linear electron flow: 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 + )

68 Figure 10.UN02 H 2 O CO 2 Light LIGHT REACTIONS NADP + ADP ATP CALVIN CYCLE NADPH O 2 [CH 2 O] (sugar)

69 Figure Primary acceptor e 2 1 Light P680 Photosystem II (PS II) Pigment molecules

70 Figure Primary acceptor 1 2 H + ½ + O 2 Light 3 H 2 O e e e 2 P680 Photosystem II (PS II) Pigment molecules

71 Figure H + ½ + O 2 3 H 2 O e e Primary acceptor e 2 4 Electron transport chain Pq Cytochrome complex Pc 1 Light P680 5 ATP Photosystem II (PS II) Pigment molecules

72 Figure H + ½ + O 2 Light 3 H 2 O e e Primary acceptor e 2 P680 4 Electron transport chain Pq Cytochrome complex 5 Pc Primary acceptor e P700 6 Light ATP Photosystem II (PS II) Pigment molecules Photosystem I (PS I)

73 Figure H + ½ + O 2 Light 3 H 2 O e e Primary acceptor e 2 P680 4 Electron transport chain Pq Cytochrome complex 5 Pc Primary acceptor e P700 Fd 7 e e 6 Electron transport chain 8 NADP + reductase Light NADP + + H + NADPH ATP Photosystem II (PS II) Pigment molecules Photosystem I (PS I)

74 3. H 2 O is split by enzymes, and the electrons are transferred from the hydrogen atoms to P680 +, thus reducing it to P680 P680 + is the strongest known biological oxidizing agent O 2 is released as a by-product of this reaction

75 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

76 6. 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

77 7. Each electron falls down an electron transport chain from the primary electron acceptor of PS I to the protein ferredoxin (Fd) 8. 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

78 The energy changes of electrons during linear flow through the light reactions can be shown in a mechanical analogy

79 Figure e e e Mill makes ATP e e e NADPH e ATP Photosystem II Photosystem I

80 Cyclic Electron Flow In cyclic electron flow, electrons cycle back from Fd to the PS I reaction center Cyclic electron flow uses only photosystem I and produces ATP, but not NADPH No oxygen is released

81 Figure Primary acceptor Pq Fd Cytochrome complex Primary acceptor Fd NADP + reductase NADP + + H + NADPH Pc Photosystem II ATP Photosystem I

82 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

83 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

84 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

85 Figure MITOCHONDRION STRUCTURE CHLOROPLAST STRUCTURE Intermembrane space H + Diffusion Thylakoid space Inner membrane Matrix Electron transport chain ATP synthase Thylakoid membrane Stroma Key Higher [H + ] ADP + P i H + ATP Lower [H + ]

86 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

87 Figure Photosystem II Light 4 H + Cytochrome complex Light Photosystem I Fd NADP + reductase 3 NADP + + H + H 2 O e e Pq 1 O 2 THYLAKOID SPACE ½ +2 H + 4 H + (high H + concentration) 2 Pc NADPH To Calvin Cycle STROMA (low H + concentration) Thylakoid membrane ATP synthase ADP + P i H + ATP

88 Figure 10.18a Photosystem II Cytochrome complex Light Photosystem I Light 4 H + Fd Pq H 2 O 1 THYLAKOID SPACE (high H + concentration) e e 2 ½ O 2 +2 H + 4 H + Pc STROMA (low H + concentration) Thylakoid membrane ATP synthase ADP + P i H + ATP

89 Figure 10.18b Cytochrome complex Light Photosystem I Fd NADP + reductase 3 NADP + + H + Pq 2 4 H + Pc THYLAKOID SPACE (high H + concentration) NADPH To Calvin Cycle ATP synthase ADP + P i H + ATP STROMA (low H + concentration)

90 Concept 10.3: The Calvin cycle uses the chemical energy of ATP and NADPH to reduce 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

91 Carbon enters the cycle as CO 2 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 CO 2 The Calvin cycle has three phases 1. Carbon fixation (catalyzed by rubisco) 2. Reduction 3. Regeneration of the CO 2 acceptor (RuBP)

92 Figure 10.UN03 H 2 O CO 2 Light LIGHT REACTIONS NADP + ADP ATP CALVIN CYCLE NADPH O 2 [CH 2 O] (sugar)

93 Figure Input 3 CO 2, entering one per cycle Rubisco Phase 1: Carbon fixation 3 P P 3 P RuBP P 6 3-Phosphoglycerate P Calvin Cycle

94 Figure Input 3 CO 2, entering one per cycle Rubisco Phase 1: Carbon fixation 3 P P 3 P RuBP P 6 3-Phosphoglycerate P 6 ATP 6 ADP Calvin Cycle 6 P P 1,3-Bisphosphoglycerate 6 NADPH 6 NADP + 6 P i 6 G3P P Phase 2: Reduction 1 G3P Output P Glucose and other organic compounds

95 Figure Input 3 CO 2, entering one per cycle Rubisco Phase 1: Carbon fixation 3 P P 3 P RuBP P 6 3-Phosphoglycerate P 6 ATP 6 ADP 3 ATP 3 ADP Phase 3: Regeneration of RuBP 5 P G3P Calvin Cycle 6 1,3-Bisphosphoglycerate G3P 6 P P P 6 6 NADP + 6 P i NADPH Phase 2: Reduction 1 G3P Output P Glucose and other organic compounds

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