10/2/2011. Outline. The Process That Feeds the Biosphere. Autotrophs. Photosynthetic Organisms

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1 Chapter 10 hotosynthesis Outline 1. hotosynthesis overview. 3. igments 4. -dependent rxn 5. -independent rxn () 6. C3, C4 and CAM The rocess That Feeds the Biosphere hotosynthesis is the process that converts solar energy into chemical energy Directly or indirectly, photosynthesis nourishes almost the entire living world Autotrophs Autotrophs sustain themselves without eating anything derived from other organisms Autotrophs are the producers of the biosphere, producing organic molecules from CO and other inorganic molecules Almost all plants are photoautotrophs, using the energy of sunlight to make organic molecules 3 4 hotosynthetic Organisms Figure 10. hotosynthesis occurs in plants, algae, certain other protists, and some prokaryotes These organisms feed not only themselves but also most of the living world (a) lants (b) Multicellular alga (c) Unicellular protists 10 m (d) Cyanobacteria 40 m 5 (e) urple sulfur bacteria 1 m 6 1

2 Heterotrophs 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 BioFlix: hotosynthesis 7 8 hotosynthesis 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 Chloroplasts: The Sites of hotosynthesis in lants 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 9 10 Chloroplast Structure Figure 10.4a Leaf cross section Chloroplasts Vein CO enters and O 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 Mesophyll Chloroplast Stomata CO O Mesophyll cell m

3 Figure 10.4b Chloroplast Stroma Thylakoid Granum Thylakoid space Outer membrane Intermembrane space Inner membrane 1 m 14 What is the stomata Chlorophyll a is found in the: 1. The watery matrix of the chloroplast. ores that let CO in 3. A stack of thylakoids 33% 33% 33% 1. Thylakoid membrane. Thylakoid lumen 3. Stroma 33% 33% 33% The watery matrix of... ores that let CO in A stack of thylakoids Thylakoid membrane Thylakoid lumen Stroma Tracking Atoms Through hotosynthesis The Splitting of Water hotosynthesis is a complex series of reactions that can be summarized as the following equation: Chloroplasts split H O into hydrogen and oxygen, incorporating the electrons of hydrogen into sugar molecules and releasing oxygen as a by-product 6 CO + 1 H O + energy C 6 H 1 O O + 6 H O

4 Figure 10.5 hotosynthesis as a Redox rocess Reactants: roducts: 6 CO 1 H O C 6 H 1 O 6 6 H O 6 O hotosynthesis reverses the direction of electron flow compared to respiration hotosynthesis is a redox process in which H O is oxidized and CO is reduced hotosynthesis is an endergonic process; the energy boost is provided by light 19 0 Figure 10.UN01 hotosynthesis two stages becomes reduced Energy 6 CO 6 H O C 6 H 1 O 6 6 O becomes oxidized 1. dependent stage: Happens in the thylakoids energy is converted to chemical energy. independent reactions ( ): Happens in the stroma Carbon is fixed from CO to glucose 1 The Two Stages of hotosynthesis: An Overview The Two Stages of hotosynthesis: An Overview hotosynthesis consists of the light reactions (the photo part) and cycle (the synthesis part) The light reactions (in the thylakoids) Split H O Release O Reduce NAD + to NADH Generate from AD by photophosphorylation The cycle (in the stroma) forms sugar from CO, using and NADH The cycle begins with carbon fixation, incorporating CO into organic molecules 3 4 4

5 Figure H O Figure H O AD + i AD + i Reactions Reactions NADH Chloroplast Chloroplast 5 O 6 Figure Figure H O CO H O CO Reactions AD + i Reactions AD + i NADH NADH Chloroplast Chloroplast O 7 O [CH O] (sugar) 8 The light reactions convert solar energy to the chemical energy of and NADH Figure 10.1 Chloroplasts are solar-powered chemical factories Their thylakoids transform light energy into the chemical energy of and NADH

6 The Nature of Sunlight roperties of 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 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 also behaves as though it consists of discrete particles, called photons 31 3 Figure m 10 5 nm 10 3 nm 1 nm 10 3 nm 10 6 nm (10 9 nm) 10 3 m hotosynthetic igments: The Receptors Gamma rays X-rays UV Infrared Microwaves Radio waves igments are substances that absorb visible light Different pigments absorb different wavelengths Wavelengths that are not absorbed are reflected or transmitted Visible light Leaves appear green because chlorophyll reflects and transmits green light nm Shorter wavelength Longer wavelength Higher energy Lower energy Figure 10.8 Chloroplast Reflected light Absorbed light Granum Animation: and igments Right-click slide / select lay 35 Transmitted light 36 6

7 Spectrophotometers A spectrophotometer measures a pigment s ability to absorb various wavelengths Figure 10.9 TECHNIQUE White light Refracting prism Chlorophyll solution hotoelectric tube Galvanometer This machine sends light through pigments and measures the fraction of light transmitted at each wavelength Slit moves to pass light of selected wavelength. Green light High transmittance (low absorption): Chlorophyll absorbs very little green light. 37 Blue light Low transmittance (high absorption): Chlorophyll absorbs most blue light. 38 Absorption spectrum 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 Figure RESULTS (a) Absorption spectra (b) Action spectrum Absorption of light by chloroplast pigments Rate of photosynthesis (measured by O release) Chlorophyll a Chlorophyll b Carotenoids Wavelength of light (nm) Aerobic bacteria Filament of alga 39 (c) Engelmann s experiment Engelmann s experiments 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 He used the growth of aerobic bacteria clustered along the alga as a measure of O production 41 hotosynthetic igments 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 4 7

8 Figure CH 3 CH 3 in chlorophyll a CHO in chlorophyll b Excitation of Chlorophyll by orphyrin ring 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 Hydrocarbon tail (H atoms not shown) Figure 10.1 A hotosystem: A Reaction-Center Complex Associated with -Harvesting Complexes Energy of electron hoton Chlorophyll molecule Excited state Heat hoton (fluorescence) Ground state 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 (a) Excitation of isolated chlorophyll molecule (b) Fluorescence Figure hoton STROMA electron Figure 10.13a hoton hotosystem harvestincenter Reaction- complexes complex harvesting complexes hotosystem Reactioncenter complex STROMA electron Thylakoid membrane Transfer Special pair of igment of energy chlorophyll a molecules molecules THYLAKOID SACE (INTERIOR OF THYLAKOID) (a) How a photosystem harvests light Thylakoid membrane Chlorophyll rotein subunits (b) Structure of photosystem II STROMA THYLAKOID SACE Thylakoid membrane Transfer of energy Special pair of chlorophyll a molecules igment molecules THYLAKOID SACE (INTERIOR OF THYLAKOID) 47 (a) How a photosystem harvests light 48 8

9 Figure 10.13b Thylakoid membrane rotein subunits Chlorophyll STROMA THYLAKOID SACE A primary electron 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 is the first step of the light reactions (b) Structure of photosystem II hotosystems hotosystems There are two types of photosystems in the thylakoid membrane hotosystem II (S II) functions first (the numbers reflect order of discovery) and is best at absorbing a wavelength of 680 nm hotosystem I (S I) is best at absorbing a wavelength of 700 nm The reaction-center chlorophyll a of S I is called 700 The reaction-center chlorophyll a of S II is called Linear Electron Flow 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 and NADH using light energy 1. A photon hits a pigment and its energy is passed among pigment molecules until it excites 680. An excited electron from 680 is transferred to the primary electron (we now call it )

10 Figure Linear Electron Flow is a very strong oxidizing agent 3. H O is split by enzymes, and the electrons are transferred from the hydrogen atoms to 680 +, thus reducing it to 680 igment molecules O is released as a by-product of this reaction hotosystem II (S II) Figure Linear Electron Flow H e H O + 1 / O Each electron falls down an electron transport chain from the primary electron of S II to S I 5. Energy released by the fall drives the creation of a proton gradient across the thylakoid membrane hotosystem II (S II) igment molecules Diffusion of H + (protons) across the membrane drives synthesis Figure Linear Electron Flow H + 1 / O 3 H O q 4 6. In S I (like S II), transferred light energy excites 700, which loses an electron to an electron Cytochrome complex c (700 that is missing an electron) accepts an electron passed down from S II via the electron transport chain igment molecules hotosystem II (S II)

11 Figure Linear Electron Flow H + 1 / O H e O q 4 Cytochrome complex 5 c Each electron falls down an electron transport chain from the primary electron of S I to the protein ferredoxin (Fd) 8. The electrons are then transferred to NAD + and reduce it to NADH 1 6 The electrons of NADH are available for the reactions of the cycle hotosystem II (S II) igment molecules hotosystem I (S I) This process also removes an H + from the stroma 61 6 Figure Figure H + 1 / O H e O 3 q 4 Cytochrome complex c 7 Fd 8 reductase + H NADH Mill makes NADH hotosystem II (S II) igment molecules hotosystem I (S I) 63 hotosystem II hotosystem I 64 Cyclic Electron Flow Figure Cyclic electron flow uses only photosystem I and produces, but not NADH No oxygen is released Cyclic electron flow generates surplus, satisfying the higher demand in the cycle q Fd Cytochrome complex c Fd reductase + H NADH Occurs when there is a build up of NADH hotosystem II hotosystem I

12 Cyclic Electron Flow Some organisms such as purple sulfur bacteria have S I but not S II Cyclic electron flow is thought to have evolved before linear electron flow Cyclic electron flow may protect cells from light-induced damage A Comparison of Chemiosmosis in Chloroplasts and Mitochondria Chloroplasts and mitochondria generate by chemiosmosis, but use different sources of energy Mitochondria transfer chemical energy from food to ; chloroplasts transform light energy into the chemical energy of Spatial organization of chemiosmosis differs between chloroplasts and mitochondria but also shows similarities A Comparison of Chemiosmosis in Chloroplasts and Mitochondria Figure Mitochondrion Chloroplast In mitochondria, protons are pumped to the intermembrane space and drive synthesis as they diffuse back into the mitochondrial matrix In chloroplasts, protons are pumped into the thylakoid space and drive synthesis as they diffuse back into the stroma MITOCHONDRION STRUCTURE Intermembrane space Inner membrane Matrix Electron transport chain H synthase Diffusion CHLOROLAST STRUCTURE Thylakoid space Thylakoid membrane Stroma 69 Key Higher [H ] Lower [H ] AD i H 70 Figure and NADH are produced on the side facing the stroma, where the cycle takes place In summary, light reactions generate and increase the potential energy of electrons by moving them from H O to NADH STROMA (low H concentration) HO 1 1 / O THYLAKOID SACE + H + (high H concentration) STROMA (low H concentration) hotosystem II 4 H + Thylakoid membrane Cytochrome complex hotosystem I q 4 H + c synthase AD + i H + Fd reductase 3 NADH + H To

13 Figure 10.UN08 The wavelength of light that the chlorophyll a pigment found at the reaction center in photosystem I absorbs is: nm. 700 nm nm nm 5% 5% 5% 5% nm 700 nm 680 nm 800 nm The electron donor for photosystem II is: The final electron from photosystem I is: 1. NADH. Water 3. NADH 4. Oxygen 5% 5% 5% 5% 1. NAD+. Water 3. NAD+ 4. Oxygen 5% 5% 5% 5% NADH Water NADH Oxygen NAD+ Water NAD+ Oxygen The cycle uses the chemical energy of and NADH to reduce CO to sugar The cycle, like the citric acid cycle, regenerates its starting material after molecules enter and leave the cycle Takes place in the stroma of the chloroplasts Needs: 18, 1 NADH, 6 Carbon dioxide (CO ) to make 1 glucose The cycle builds sugar from smaller molecules by using and the reducing power of electrons carried by NADH 77 13

14 6 molecules of CO 6 molecules of ribulose bisphosphate (RuB) 1 CO uptake 6 AD phase 6 molecules of ribulose CALVIN phosphate (R) CYCLE 3 RuB Carbon regeneration reduction phase phase 10 molecules of G3 Glucose and other carbohydrate synthesis 1 molecules of glyceraldehyde-3- phosphate (G3) Through a series of molecules reactions G3 is of glyceraldehyde-3- rearranged into new phosphate (G3) RuB molecules or another sugar CO molecules are captured by RuB, resulting in an unstable intermediate that is immediately broken apart into GA 1 molecules of phosphoglycerate (GA) 1 1 AD 1 NADH 1 NAD + GA is phosphorylated by and reduced by NADH. Removal of a phosphate results in formation of G3. The Carbon enters the cycle as CO and leaves as a sugar named glyceraldehyde 3-phospate (G3) For net synthesis of 1 G3, the cycle must take place three times, fixing 3 molecules of CO The cycle has three phases Carbon fixation (catalyzed by rubisco) Reduction Regeneration of the CO (RuB) 80 Figure Input Figure Input 3 (Entering one CO at a time) 3 (Entering one CO at a time) hase 1: Carbon fixation hase 1: Carbon fixation Rubisco Rubisco 3 Short-lived intermediate 3 Short-lived intermediate 3 Ribulose bisphosphate (RuB) 6 3-hosphoglycerate 3 Ribulose bisphosphate (RuB) 6 3-hosphoglycerate 6 6 AD 6 1,3-Bisphosphoglycerate 6 NADH 6 6 i 6 Glyceraldehyde 3-phosphate (G3) hase : Reduction 81 1 G3 (a sugar) Output Glucose and other organic compounds 8 Figure Input 3 (Entering one at a time) CO 1 st step: Carbon uptake = carbon fixation 3 Ribulose bisphosphate (RuB) 3 AD 3 hase 3: Regeneration of the CO (RuB) 5 G3 hase 1: Carbon fixation Rubisco 3 Short-lived intermediate 6 3-hosphoglycerate 6 6 AD 6 1,3-Bisphosphoglycerate 6 NADH 6 6 i 6 Glyceraldehyde 3-phosphate hase : (G3) Reduction CO and a phosphorylated five carbon molecule, ribulose bisphosphate (RuB) combines with the help of an enzyme, rubisco, to form a six carbon molecule that spontaneously splits into two 3 carbon molecules, phosphoglycerate (GA) CO + RuB rubisco GA 1 G3 (a sugar) Output Glucose and other organic compounds 83 14

15 cd Step: Carbon Reduction hosphoglycerate (GA) is converted to glyceraldehyde-3-phosphate (G3). is used to phosphorylate GA and NADH is used to reduce GA 3 CO molecules, 9 and 6 NADH are needed for the cycle to produce 1 three carbon sugar (G3) This needs to happen times to make a glucose molecule So 6 CO molecules, 18 and 1 NADH are needed for the cycle to produce 1 glucose During the light dependent reactions Carbon is fixed. HO is split to produce oxygen 3. NADH and are produced 4. and 3 5% 5% 5% 5% ZdFTY OD1dQ&feature=related Carbon is fixed HO is split to produ.. NADH and are... and 3 15

16 lants have mitochondria lants carry out cellular respiration 1. True. False 50% 50% 1. True. False 50% 50% True False True False 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 H O but also limits photosynthesis The closing of stomata reduces access to CO and causes O to build up These conditions favor an apparently wasteful process called photorespiration hotorespiration: An Evolutionary Relic? In most plants (C 3 plants), initial fixation of CO, via rubisco, forms a three-carbon compound (3- phosphoglycerate) In photorespiration, rubisco adds O instead of CO in the cycle, producing a two-carbon compound hotorespiration consumes O and organic fuel and releases CO without producing or sugar hotorespiration C 4 lants hotorespiration may be an evolutionary relic because rubisco first evolved at a time when the atmosphere had far less O and more CO hotorespiration limits damaging products of light reactions that build up in the absence of the 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 cycle C 4 plants minimize the cost of photorespiration by incorporating CO into four-carbon compounds in mesophyll cells This step requires the enzyme E carboxylase E carboxylase has a higher affinity for CO than rubisco does; it can fix CO even when CO concentrations are low These four-carbon compounds are exported to bundle-sheath cells, where they release CO that is then used in the cycle

17 Figure 10.0 Figure 10.0a hotosynthetic cells of C 4 plant leaf Mesophyll cell Bundlesheath cell Vein (vascular tissue) C 4 leaf anatomy Stoma The C 4 pathway Mesophyll cell E carboxylase Oxaloacetate (4C) Bundlesheath cell Malate (4C) CO E (3C) AD yruvate (3C) CO hotosynthetic cells of C 4 plant leaf Mesophyll cell Bundlesheath cell Vein (vascular tissue) C 4 leaf anatomy Sugar Vascular tissue Stoma Figure 10.0b The C 4 pathway Mesophyll cell E carboxylase Oxaloacetate (4C) Bundlesheath cell Malate (4C) CO E (3C) AD yruvate (3C) Sugar CO In the last 150 years since the Industrial Revolution, CO levels have risen greatly Increasing levels of CO may affect C 3 and C 4 plants differently, perhaps changing the relative abundance of these species The effects of such changes are unpredictable and a cause for concern Vascular tissue CAM lants Figure 10.1 Some plants, including succulents, use crassulacean acid metabolism (CAM) to fix carbon CAM plants open their stomata at night, incorporating CO into organic acids Stomata close during the day, and CO is released from organic acids and used in the cycle Sugarcane Mesophyll cell Bundlesheath cell C 4 Organic acid CO CO 1 ineapple CO incorporated (carbon fixation) CO released to the cycle CAM Organic acid CO CO Night Day 101 Sugar (a) Spatial separation of steps Sugar 10 (b) Temporal separation of steps 17

18 Figure 10.1a Figure 10.1b Sugarcane ineapple The Importance of hotosynthesis: A Review Figure 10. H O CO 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 lants store excess sugar as starch in structures such as roots, tubers, seeds, and fruits In addition to food production, photosynthesis produces the O in our atmosphere Reactions: hotosystem II Electron transport chain hotosystem I Electron transport chain Chloroplast AD + i RuB NADH 3-hosphoglycerate G3 Starch (storage) 105 O Sucrose (export) 106 Figure 10.UN0 Figure 10.UN03 3 CO O H O q Cytochrome complex c Fd reductase + H NADH Regeneration of CO Carbon fixation 3 5C 6 3C 5 3C hotosystem II hotosystem I Reduction G3 (3C)

19 Important Concepts Know the vocabulary in this lecture Know the overall reaction of photosynthesis, what is being reduced and oxidized Understand the light spectrum, what wavelengths and colors are absorbed by chlorohyll, know the pigments involved in photosynthesis, the general structure of chlorophyll, know what wavelength is absorbed by the different types of chlorophyll a Important Concepts Know the leaf structure and the functions of the components including the Stomata, veins, Chloroplasts, mesophyll layer Know the chloroplast structure Where do the two stages of photosynthesis (light dependent and light independent) take place Know what happens during the light dependent stage, including what is needed, what is produced, what molecules are donating and accepting electrons, who do they give them to or get them from? Important Concepts Understand the role of the reaction centers, photosystem I, photosystem II, the electron transport chains in the light dependent reaction, know the differences between photosystem I and II Understand how is produced during the light dependent stage, and how NADH is produced Understand the cyclic electron flow, when does this happen and what are the benefits? Know that you need 18, 1 NADH, and 6CO to make one glucose molecule in the calvin cycle Important Concepts Know what happens during the light independent () stage, know what is needed and what is produced, including: glyceraldehyde-3- phosphate (G3) which are made into one glucose. Know the three stages of the cycle and what goes into and out of each stage and what is the purpose of the stage Know the role of the enzymes rubisco and E carboxylase Know what C3 plants do, and know the C4 and CAM adaptations, and what photorespiration is 19

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