BIOLOGY. Photosynthesis. The Process That Feeds the Biosphere. Autotrophs sustain themselves without eating anything derived from other organisms

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1 0 Photosynthesis CAMPBELL BIOLOGY TENTH EDITION Reece Urry Cain Wasserman Minorsky Jackson energy ECOSYSTEM CO H O Photosynthesis in chloroplasts Cellular respiration in mitochondria Organic O powers most cellular work Lecture Presentation by Nicole Tunbridge and Kathleen Fitzpatrick Heat energy 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 Autotrophs sustain themselves without eating anything derived from other organisms Autotrophs are the producers of the biosphere, producing organic from CO and other inorganic Almost all plants are photoautotrophs, using the energy of sunlight to make organic 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

2 0 μm μm Figure 0. (a) Plants (d) Cyanobacteria 40 μm 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 (b) Multicellular alga (e) Purple sulfur bacteria (c) Unicellular eukaryotes 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 Researchers are exploring methods of capitalizing on the photosynthetic process to provide alternative fuels. Concept 0.: Photosynthesis converts light energy to the chemical energy of food Chloroplasts are structurally similar to and likely evolved from photosynthetic bacteria ( endosymbiont theory) The structural organization of these organelles allows for the chemical reactions of photosynthesis

3 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 enters and O exits the leaf through Stroma Granum space microscopic pores called stomata Mesophyll Leaf cross section Chloroplasts Vein Stomata CO O Chloroplast Mesophyll cell Outer membrane Intermembrane space Inner membrane 0 μm A chloroplast has an envelope of two membranes surrounding a dense fluid called the stroma s are connected sacs in the chloroplast which compose a third membrane system s may be stacked in columns called grana Chlorophyll, the pigment which gives leaves their green colour, resides in the thylakoid membranes Stroma Granum space μm Chloroplast Outer membrane Intermembrane space Inner membrane μm Tracking Atoms Through Photosynthesis: Scientific Inquiry Photosynthesis is a complex series of reactions that can be summarized as the following equation: CO H O energy C H O O H O The overall chemical change during photosynthesis is the reverse of the one that occurs during cellular respiration The Splitting of Water Chloroplasts split H O into hydrogen and oxygen, incorporating the electrons of hydrogen into sugar and releasing oxygen as a by-product Reactants: CO H O Products: C H O H O O 3

4 Photosynthesis as a Redox Process Photosynthesis reverses the direction of electron flow compared to respiration Photosynthesis is a redox process in which H O is oxidized and CO is reduced The Two Stages of Photosynthesis: A Preview Photosynthesis consists of the light reactions (the photo part) and Calvin cycle (the synthesis part) H O CO Photosynthesis is an endergonic process; the energy boost is provided by light LIGHT REACTIONS ADP P i CALVIN CYCLE becomes reduced Stroma becomes oxidized Chloroplast O [CH O] (sugar) The light reactions (in the thylakoids) Split H O Release O Reduce the electron to Generate from ADP by photophosphorylation Concept 0.: The light reactions convert solar energy to the chemical energy of and Chloroplasts are solar-powered chemical factories Their thylakoids transform light energy into the chemical energy of and The Calvin cycle (in the stroma) Forms sugar from CO, using and Begins with carbon fixation, incorporating CO into organic 4

5 The Nature of Sunlight 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 m 0 5 nm 0 3 nm nm 0 3 nm 0 nm (0 9 nm) 0 3 m Gamma rays Shorter The wavelength electromagnetic spectrum is the entire range Higher of energy electromagnetic energy, Lower or radiation energy X-rays UV Infrared Visible light Microwaves Radio waves nm Longer wavelength 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 Photons are not tangible objects, but they act like objects in that each of them has a fixed quantity of energy Photosynthetic Pigments: The 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 Chloroplast Absorbed light Transmitted light Reflected light Granum 5

6 release) Absorption of light by chloroplast pigments A spectrophotometer measures a pigment s ability to absorb various wavelengths Figure 0.9 White light Refracting prism Chlorophyll solution 3 Photoelectric tube 4 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 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. Figure 0.0a Chlorophyll a Chlorophyll b 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 Carotenoids Wavelength of light (nm) (a) Absorption spectra Rate of photosynthesis (measured by O (b) Action spectrum

7 release) Absorption of light by chloroplast pigments Figure 0.0 Chlorophyll a Chlorophyll b 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 Accessory pigments called carotenoids absorb excessive light that would damage chlorophyll ( photoprotection ) Carotenoids Wavelength of light (nm) (a) Absorption spectra Rate of photosynthesis (measured by O (b) Action spectrum Aerobic bacteria Filament of alga (c) Engelmann s experiment The action spectrum of photosynthesis was first demonstrated in 883 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 Figure 0. CH 3 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 7

8 membrane membrane Energy of electron Excitation of Chlorophyll by Figure 0. 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 Photon Chlorophyll molecule Excited state Heat Photon (fluorescence) Ground state (a) Excitation of isolated chlorophyll molecule (b) Fluorescence A Photosystem: A Reaction-Center Complex Associated with -Harvesting Complexes A photosystem consists of a reactioncenter complex (a type of protein complex) surrounded by lightharvesting complexes The light-harvesting complexes (pigment bound to proteins) transfer the energy of photons to the reaction center Photon harvesting complexes Photosystem Reaction- Center complex Transfer Special pair of of energy chlorophyll a THYLAKOID SPACE (INTERIOR OF THYLAKOID) (a) How a photosystem harvests light STROMA electron Pigment 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 Photon harvesting complexes Photosystem Reaction- Center complex Transfer Special pair of of energy chlorophyll a THYLAKOID SPACE (INTERIOR OF THYLAKOID) (a) How a photosystem harvests light STROMA electron Pigment 8

9 There are two types of photosystems in the thylakoid membrane I (PS II) functions first (the numbers reflect order of discovery) and is best at absorbing a wavelength of 80 nm The reaction-center chlorophyll a of PS II is called P80 Linear 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 using light energy (PS I) is best at absorbing a wavelength of 700 nm The reaction-center chlorophyll a of PS I is called P700 Figure H H O O 3 ½ P80 4 Pq Cytochrome complex 5 Pc P reductase H Figure H H O O 3 ½ P80 4 Pq Cytochrome complex 5 Pc P reductase H I (PS II) Pigment (PS I) There are 8 steps in linear electron flow:. A photon hits a pigment and its energy is passed among pigment until it excites P80. An excited electron from P80 is transferred to the primary electron (we now call it P80 ) I (PS II) Pigment (PS I) 3. H O is split by enzymes, and the electrons are transferred from the hydrogen atoms to P80, thus reducing it to P80 P80 is the strongest known biological oxidizing agent O is released as a by-product of this reaction 9

10 Figure H H O O 3 ½ P80 4 Pq Cytochrome complex 5 Pc P reductase H Figure H H O O 3 ½ P80 4 Pq Cytochrome complex 5 Pc P reductase H I (PS II) Pigment (PS I) 4. Each electron falls down an electron from the primary electron 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 synthesis I (PS II) Pigment (PS I). In PS I (like PS II), transferred light energy excites P700, which loses an electron to an electron P700 (P700 that is missing an electron) accepts an electron passed down from PS II via the electron H H O O 3 ½ P80 4 Pq Cytochrome complex 5 Pc P reductase H The energy changes of electrons during linear flow through the light reactions shown in a mechanical analogy Mill makes Pigment (PS I) I (PS II) 7. Each electron falls down an electron from the primary electron of PS I to the protein ferredoxin () 8. The electrons are then transferred to and reduce it to The electrons of are available for the reactions of the Calvin cycle I This process also removes an H from the stroma 0

11 Cyclic Flow In cyclic electron flow, electrons cycle back from to the PS I reaction center Cyclic electron flow uses only photosystem I and produces, but not ; No oxygen is released Pq Cytochrome complex reductase H 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 Pc I A Comparison of Chemiosmosis in Chloroplasts and Mitochondria Figure 0.7 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 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 Key Inner membrane Matrix Higher [H ] Lower [H ] H synthase ADP P i H Diffusion CHLOROPLAST STRUCTURE space membrane Stroma

12 and are produced on the side facing the stroma, where the Calvin cycle takes place In summary, light reactions generate and increase the potential energy of electrons by moving them from H O to I 4 H Cytochrome complex Pq e e H O THYLAKOID SPACE ½ O H 4 H (high H concentration) Pc reductase 3 H To Calvin Cycle Concept 0.3: The Calvin cycle uses the chemical energy of and to reduce CO to sugar The Calvin cycle, like the citric acid cycle, regenerates its starting material after enter and leave the cycle The cycle builds sugar from smaller by using and the reducing power of electrons carried by STROMA (low H concentration) membrane synthase ADP P i H Figure 0.UN03 H O CO LIGHT REACTIONS O ADP CALVIN CYCLE [CH O] (sugar) Carbon enters the cycle as CO and leaves as a sugar named glyceraldehyde 3-phospate (G3P) For net synthesis of G3P, the cycle must take place three times, fixing 3 of CO The Calvin cycle has three phases. Carbon fixation (catalyzed by rubisco). Reduction 3. Regeneration of the CO (RuBP)

13 Figure ADP Phase 3: Regeneration of RuBP 3 P RuBP 5 P G3P Rubisco P Input 3 CO, entering one per cycle Calvin Cycle Phase : Carbon fixation 3 P P P 3-Phosphoglycerate P P,3-Bisphosphoglycerate G3P P ADP P i Phase : Reduction Concept 0.4: 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 H O but also limits photosynthesis The closing of stomata reduces access to CO and causes O to build up P G3P Output Glucose and other organic compounds These conditions favor an apparently wasteful process called photorespiration Photorespiration: 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 Calvin cycle, producing a two-carbon compound Photorespiration consumes O and organic fuel and releases CO without producing or sugar Photorespiration may be an evolutionary relic because rubisco first evolved at a time when the atmosphere had far less O and more CO 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 3

14 C 4 Plants C 4 plants minimize the cost of photorespiration by incorporating CO into fourcarbon compounds There are two distinct types of cells in the leaves of C 4 plants: Bundle-sheath cells are arranged in tightly packed sheaths around the veins of the leaf Mesophyll cells are loosely packed between the bundle sheath and the leaf surface Photosynthetic cells of C 4 plant leaf Mesophyll cell Bundlesheath cell Vein (vascular tissue) C 4 leaf anatomy Stoma Sugar production in C 4 plants occurs in a three-step process:. The production of the four carbon precursors is catalyzed by the enzyme PEP carboxylase in the mesophyll cells PEP 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 3. Within the bundle-sheath cells, they release CO that is then used in the Calvin cycle The C 4 pathway Mesophyll cell PEP carboxylase Oxaloacetate (4C) Malate (4C) Bundlesheath cell Pyruvate CO (3C) PEP (3C) ADP Calvin Cycle Sugar Vascular tissue CO CAM Plants Since the Industrial Revolution in the 800s, 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 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 Calvin cycle 4

15 Figure 0. The Importance of Photosynthesis: A Review Sugarcane Pineapple C 4 CO CO CAM Mesophyll cell Bundlesheath cell Organic acid CO Calvin Cycle Sugar (a) Spatial separation of steps Organic acid CO Calvin Cycle Sugar Night Day (b) Temporal separation of steps 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 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 in our atmosphere Figure 0.a Figure 0.3 O CO MAKE CONNECTIONS The Working Cell Flow of Genetic Information in the Cell: DNA RNA Protein (Chapters 5 7) Movement Across Cell Membranes (Chapter 7) Energy Transformations in the Cell: Photosynthesis and Cellular Respiration (Chapters 8 0) H O Sucrose (export) Mesophyll cell Nucleus mrna Nuclear pore DNA H O Chloroplast CO Protein 3 Ribosome mrna Rough endoplasmic Protein reticulum (ER) in vesicle Vacuole LIGHT REACTIONS: I ADP P i RuBP 3-Phosphoglycerate CALVIN CYCLE G3P Starch (storage) 4 Vesicle Golgi forming apparatus Protein Plasma membrane 5 Cell wall 9 7 H O Photosynthesis in chloroplast 0 Organic O CO O CO H O 8 Cellular respiration in mitochondrion Transport pump HO O Sucrose (export) 5

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