Photosynthesis Chapter 8 Photosynthesis • Energy for the majority of life on Earth ultimately comes from the sun through photosynthesis • Photosynthesis is the process that converts energy of sunlight the into chemical energy, especially carbohydrates • Carbon Fixation: Photosynthesis collects the energy of sunlight and captures it into a chemical form • Photosynthesis uses CO2 to build carbohydrates (C6H12O6) 2 Photosynthesis • Photoautotrophs: photosynthetic organisms that can produce their own ‘food’ – these organisms fix the energy of sunlight into carbohydrate form – they then use the carbohydrates they produce for cellular respiration • Heterotrophs: organisms that require an alternative food source – these organisms consume photoautotrophs along with the carbohydrates they produce – they then use the carbohydrates they consume for cellular respiration 33 Photosynthesis • Oxygenic photosynthesis: produces O2 - carried out by cyanobacteria, 7 groups of algae, all land plants • Photosynthetic organisms serve as a food and energy source for the majority of life on on earth • Photosynthetic organisms directly or indirectly provide energy for the majority of the life on earth - most living things depend on the sun for energy - a few exceptions include chemoautotrophs: autotrophic organisms that utilize energy from other sources, live independently from the sun 44 Oxygenic Photosynthesis • Energy Fixation • Maintains O2 in the atmosphere • vital for aerobic respiration • Removes CO2 from the atmosphere • green house gases • H2O production 55 Photosynthesis 6CO2 + 12H2O carbon dioxide water C6H12O6 + 6H2O + 6O2 glucose water oxygen • Reactants: 6CO2 + 12H2O • Products: C6H12O6 + 6H2O + 6O2 • Photosynthesis vs. Cellular Respiration: 6CO2 + 12H2O C6H12O6 + 6O2 C6H12O6 + 6H2O + 6O2 6CO2 + 12H2O 66 Photosynthesis vs. Cellular Respiration • Photoautotrophs • Chloroplasts – grana and • ETC • Heterotrophs • Mitochondria thylakoids - mitochondrial membrane, cisternae – NADP reduced to NADPH • Krebs Cycle - Carbohydrates – ATP synthase, ATP CO2 production • Calvin Cycle – CO2 reduced to carbohydrates oxidized to • ETC - NAD+ reduced to NADH - ATP synthase, ATP 7 production Photosynthesis Photosynthesis is divided into two reactions: 1. Light-dependent reactions • capture energy from sunlight • reduce NADP+ to NADPH • produce ATP • occur in chloroplasts 2. Carbon fixation reactions • use ATP and NADPH to synthesize organic molecules from CO2 • Occurs in stroma 8 9 Photosynthesis Overview • Photosynthesis takes place in chloroplasts • Thylakoid membrane – internal membrane arranged in flattened sacs • contain chlorophyll and other pigments • Grana – stacks of thylakoid membranes • Stroma – semiliquid substance surrounding thylakoid membranes 10 Fig. 8.1-1 Fig. 8.1-2 Discovery of Photosynthesis • The work of many scientists led to the discovery of how photosynthesis works: - Jan Baptista van Helmont (1580-1644) - Joseph Priestly (1733-1804) - Jan Ingen-Housz (1730-1799) - F. F. Blackman (1866-1947) 13 Discovery of Photosynthesis • C. B. van Niel, 1930‘s - proposed a general formula: CO2+H2A + light energy CH2O + H2O + 2A - H2A is the electron donor • van Niel identified water as the source of the O2 released from photosynthesis • Robin Hill confirmed van Niel’s proposal that energy from the light reactions fuels carbon fixation 14 Energy from Light Reactions Fuels Carbon Fixation • Photon: a particle of light - acts as a discrete bundle of energy - energy content of a photon is inversely proportional to the wavelength of the light • Photoelectric effect: removal of an electron from a molecule by light - occurs when photons transfer energy to electrons • Light energy is absorbed by pigments: molecules that absorb visible light 15 Energy from Light Reactions Fuels Carbon Fixation 16 Pigments • Pigments: molecules that absorb visible light - chlorophyll a - chlorophyll b - carotenoids • Each pigment has a characteristic absorption spectrum: the range and efficiency of photons it is capable of absorbing. 17 18 Pigments • Chlorophyll a – primary pigment in plants and cyanobacteria - absorbs violet-blue and red light • Chlorophyll b – secondary pigment absorbing light wavelengths that chlorophyll a does not absorb 19 Pigments • Structure of pigments: • Porphyrin ring: complex ring structure with alternating double and single bonds - magnesium ion at the center of the ring - photons excite electrons in the ring - electrons are shuttled away from the ring 20 21 22 Pigments • Accessory pigments: secondary pigments absorbing light wavelengths other than those absorbed by chlorophyll a - increase the range of light wavelengths that can be used in photosynthesis - include: chlorophyll b, carotenoids, phycobiloproteins - carotenoids also act as antioxidants 23 Photosynthesis: Photosystems • Photosystem: Enzyme complexes for photosynthesis – enzymes use light to oxidize H2O and reduce CO2 to carbohydrates • Located in the thylakoid membrane of plants, algae and cyanobacteria or the cytoplasmic membrane of photosynthetic bacteria 24 24 Photosystem Organization • A photosystem consists of: 1. an antenna complex of hundreds of accessory pigment molecules 2. a reaction center of one or more chlorophyll a molecules • Energy of electrons is transferred through the antenna complex to the reaction center 25 26 Photosystem Organization • At the reaction center, the energy from the antenna complex is transferred to chlorophyll a • This energy causes an electron from chlorophyll to become excited - chlorophyll absorbs a photon, looses an electron • The excited electron is transferred from chlorophyll a to an electron acceptor, quinone • Water donates an electron to chlorophyll a to replace the excited electron 27 28 Light-Dependent Reactions Light-dependent reactions occur in 4 stages: 1. Primary Photoevent – a photon of light is captured by a pigment molecule 2. Charge Separation – energy is transferred to the reaction center; an excited electron is transferred to an acceptor molecule 3. Electron Transport – electrons move through carriers to reduce NADP+ 4. Chemiosmosis – produces ATP (ATP synthase) 29 30 Light-Dependent Reactions • In sulfur bacteria, only one photosystem is used for cyclic photophosphorylation 1. Hydrogen sulfide oxidized - electrons joins a proton to produce hydrogen - elemental S and protons produced as products 2. An electron is recycled to chlorophyll - this process drives the chemiosmotic synthesis of ATP • This system only produces energy 31 Light-Dependent Reactions • In chloroplasts, two linked photosystems are used in noncyclic photophosphorylation 1. Photosystem I - reaction center pigment (P700) with a peak absorption at 700nm 2. Photosystem II - reaction center pigment (P680) has a peak absorption at 680nm • This system produces energy, O2, and 32 NADPH for biosynthesis of carbohydrates Light-Dependent Reactions • Photosystem II acts first: - accessory pigments shuttle energy to the P680 reaction center - excited electrons from P680 are transferred to the cytochrome/b6-f complex - connects photosystems II and I - electron lost from P680 is replaced by an electron released from hydrolysis: the splitting of water 33 Light-Dependent Reactions • The b6-f complex is a series of electron carriers - an electron transport chain - electron carrier molecules are embedded in the thylakoid membrane - protons are pumped into the thylakoid space to form a proton gradient 34 Light-Dependent Reactions • Photosystem I: - receives energy from an antenna complex - energy is shuttled to P700 reaction center - excited electron is transferred to a membranebound electron carrier - electrons are used to reduce NADP+ to NADPH - electrons lost from P700 are replaced from the b6-f complex 35 Fig. 8.13 Z Diagram of Photosystems I and II Fig. 8.13-1 Fig. 8.13-2 Fig. 8.13-3 Light-Dependent Reactions • ATP is produced via chemiosmosis: - ATP synthase is embedded in the thylakoid membrane - protons have accumulated in the thylakoid space - protons move into the stroma only through ATP synthase - ATP is produced from ADP + Pi 40 41 Carbon Fixation Reactions Calvin cycle • biochemical pathway that allows for carbon fixation • incorporates CO2 into organic molecules • occurs in the stroma • uses ATP and NADPH as energy sources 42 Carbon Fixation Reactions To build carbohydrates, cells need: 1. Energy - provided by ATP from light-dependent reactions 2. Reduction Potential: hydrogen atoms - provided by NADPH from photosystem I 43 Carbon Fixation Reactions The Calvin cycle has 3 phases: 1. Carbon Fixation - RuBP + CO2 2 molecules PGA 3 phosphoglycerate 2. Reduction - PGA is reduced to G3P glyceraldehyde-3-phosphate 3. Regeneration of RuBP - G3P is used to regenerate RuBP 44 Carbon Fixation Reactions • Carbon fixation – the incorporation of CO2 into organic molecules - occurs in the first step of the Calvin cycle: ribulose-bis-phosphate + CO2 5 carbon molecule 2(PGA) 1 carbon 3 carbons • Reaction is catalyzed by rubisco 45 46 Fig. 8.16-1 Fig. 8.16-2 Fig. 8.16-3 Carbon Fixation Reactions • Glucose is not a direct product of the Calvin cycle - 2 molecules of G3P leave the cycle - each G3P contains 3 carbons - 2 G3P are used to produce 1 glucose in reactions in the cytoplasm 50 Carbon Fixation Reactions • During the Calvin cycle, energy is needed. The energy is supplied from: - 18 ATP molecules - 12 NADPH molecules 51 Carbon Fixation Reactions • The energy cycle: - photosynthesis uses the products of respiration as starting substrates - respiration uses the products of photosynthesis as starting substrates 52 53 Photorespiration • Rubisco has 2 enzymatic activities: 1. Carboxylation – the addition of CO2 to RuBP - favored under normal conditions 2. Photorespiration – the oxidation of RuBP by the addition of O2 - favored in hot conditions - CO2 and O2 compete for the active site on RuBP. 54 Fig. 8.19-1 Fig. 8.19-2 Photorespiration • Some plants can avoid photorespiration by using an enzyme other than rubisco. - PEP carboxylase - adds CO2 to phosphoenolpyruvate (PEP) - a 4 carbon compound is produced - CO2 is later released from this 4-carbon compound and used by rubisco in the Calvin cycle 57 58 Photorespiration • C4 plants - use PEP carboxylase to capture CO2 - CO2 is added to PEP in mesophyll cell - the resulting 4-carbon compound is moved into a bundle sheath cell where the CO2 is released and used in the Calvin cycle 59 60 61 Photorespiration • CAM plants - CO2 is captured at night when stomata are open - PEP carboxylase adds CO2 to PEP to produce a 4 carbon compound - this compound releases CO2 during the day - CO2 is then used by rubisco in the Calvin cycle 62 63