Photosynthesis Chapter 8 Adapted by G. Cornwall, Ph.D. From Raven’s Biology, McGraw Hill Publishing Photosynthesis Overview • Energy for all life on Earth ultimately comes from photosynthesis 6CO2 + 12H2O C6H12O6 + 6H2O + 6O2 • Oxygenic photosynthesis is carried out by – Cyanobacteria – 7 groups of algae – All land plants – chloroplasts 2 Chloroplast • Thylakoid membrane – internal membrane – Contains chlorophyll and other photosynthetic pigments – Pigments clustered into photosystems • Grana – stacks of flattened sacs of thylakoid membrane • Stroma lamella – connect grana • Stroma – semiliquid surrounding thylakoid membranes 3 Stages • Light-dependent reactions – Require light 1.Capture energy from sunlight 2.Make ATP and reduce NADP+ to NADPH • Carbon fixation reactions or lightindependent reactions – Does not require light 3.Use ATP and NADPH to synthesize organic molecules from CO2 4 5 Pigments • Molecules that absorb light energy in the visible range • Light is a form of energy • Photon – 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 6 Absorption spectrum • When a photon strikes a molecule, its energy is either – Lost as heat – Absorbed by the electrons of the molecule • Boosts electrons into higher energy level • Absorption spectrum – range and efficiency of photons molecule is capable of absorbing 7 • Organisms have evolved a variety of different pigments • Only two general types are used in green plant photosynthesis – Chlorophylls – Carotenoids • In some organisms, other molecules also absorb light energy 8 Chlorophylls • Chlorophyll a – Main pigment in plants and cyanobacteria – Only pigment that can act directly to convert light energy to chemical energy – Absorbs violet-blue and red light • Chlorophyll b – Accessory pigment or secondary pigment absorbing light wavelengths that chlorophyll a does not absorb 9 • Structure of chlorophyll • 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 10 • Action spectrum – Relative effectiveness of different wavelengths of light in promoting photosynthesis – Corresponds to the absorption spectrum for chlorophylls 11 • Carotenoids – Carbon rings linked to chains with alternating single and double bonds – Can absorb photons with a wide range of energies – Also scavenge free radicals – antioxidant • Protective role • Phycobiloproteins – Important in low-light ocean areas 12 Photosystem Organization • Antenna complex – Hundreds of accessory pigment molecules – Gather photons and feed the captured light energy to the reaction center • Reaction center – 1 or more chlorophyll a molecules – Passes excited electrons out of the photosystem 13 Antenna complex • Also called light-harvesting complex • Captures photons from sunlight and channels them to the reaction center chlorophylls • In chloroplasts, light-harvesting complexes consist of a web of chlorophyll molecules linked together and held tightly in the thylakoid membrane by a matrix of proteins 14 Reaction center • Transmembrane protein– pigment complex • When a chlorophyll in the reaction center absorbs a photon of light, an electron is excited to a higher energy level • Light-energized electron can be transferred to the primary electron acceptor, reducing it • Oxidized chlorophyll then fills its electron “hole” by oxidizing a donor molecule 15 Light-Dependent Reactions 1. Primary photoevent Capture of light energy – 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 16 Cyclic photophosphorylation • In sulfur bacteria, only one photosystem is used • Generates ATP via electron transport • Anoxygenic photosynthesis • Excited electron passed to electron transport chain • Generates a proton gradient for ATP synthesis 17 Chloroplasts have two connected photosystems • Oxygenic photosynthesis • Photosystem I (P700) – Functions like sulfur bacteria • Photosystem II (P680) – Can generate an oxidation potential high enough to oxidize water • Working together, the two photosystems carry out a noncyclic transfer of electrons that is used to generate both ATP and NADPH 18 19 • The Two Photosystems in Plants Work Together – Photosystem II drives ATP production • Electron is passed to the reaction center of PS I (Replaces electron lost through excitation) • the energy released is used to synthesize ATP – Photosystem I drives NADPH production. • Electron is passed to NADP+ NADPH 20 Noncyclic photophosphorylation • Plants use photosystems II and I in series to produce both ATP and NADPH • Path of electrons not a circle • Photosystems replenished with electrons obtained by splitting water • Z diagram 21 Chemiosmosis • Electrochemical gradient used to synthesize ATP • Chloroplast has ATP synthase enzymes in the thylakoid membrane – Allows protons back into stroma • Stroma also contains enzymes that catalyze the reactions of carbon fixation – the Calvin cycle reactions 22 • Making more ATP – It takes more energy to fix Carbon (1 ½ ATP / NADPH) – PS I can short circuit to make extra ATP via Cyclic phosphorylation. Card Quiz A What high energy molecule is the final product of photosynthesis? Oxygen ATP NADPH Glucose Card Quiz A Where is the chlorophyll located in a plant? Cristae Outer membrane of the chloroplast Thylakoid membrane Stroma Card Quiz A A particle of light is a ____. Wave Photon Proton Newton Card Quiz A What happens at the reaction center of a photosystem? Light is absorbed An electron is energized NADP+ is reduced ATP is formed Card Quiz A Which of the following is made during the light reactions. ADP and NADP+ Glucose and ATP ATP and NADPH NADPH and Carbon Dioxide Card Quiz Answers Green Red Blue Green Green Carbon Fixation – Calvin Cycle • To build carbohydrates cells use • Energy – ATP from light-dependent reactions – Cyclic and noncyclic photophosphorylation – Drives endergonic reaction • Reduction potential – NADPH from photosystem I – Source of protons and energetic electrons 30 Calvin cycle • Named after Melvin Calvin (1911–1997) • Also called C3 photosynthesis • Key step is attachment of CO2 to RuBP to form PGA • Uses enzyme ribulose bisphosphate carboxylase/oxygenase or rubisco 31 3 phases 1. Carbon fixation – RuBP + CO2 → PGA 2. Reduction – PGA is reduced to G3P 3. Regeneration of RuBP – PGA is used to regenerate RuBP • • 3 turns incorporate enough carbon to produce a new G3P 6 turns incorporate enough carbon for 1 glucose 32 33 Output of Calvin cycle • Glucose is not a direct product of the Calvin cycle • G3P is a 3 carbon sugar – Used to form sucrose • Major transport sugar in plants • Disaccharide made of fructose and glucose – Used to make starch • Insoluble glucose polymer • Stored for later use 34 Energy cycle • Photosynthesis uses the products of respiration as starting substrates • Respiration uses the products of photosynthesis as starting substrates • Production of glucose from G3P even uses part of the ancient glycolytic pathway, run in reverse • Principal proteins involved in electron transport and ATP production in plants are evolutionarily related to those in mitochondria 35 36 Photorespiration • Rubisco has 2 enzymatic activities – Carboxylation • Addition of CO2 to RuBP • Favored under normal conditions – Photorespiration • Oxidation of RuBP by the addition of O2 • Favored when stoma are closed in hot conditions • Creates low-CO2 and high-O2 • CO2 and O2 compete for the active site on RuBP 37 Types of photosynthesis • C3 – Plants that fix carbon using only C3 photosynthesis (the Calvin cycle) • C4 and CAM – – – – – Add CO2 to PEP to form 4 carbon molecule Use PEP carboxylase Greater affinity for CO2, no oxidase activity C4 – spatial solution CAM – temporal solution 38 C4 plants • Corn, sugarcane, sorghum, and a number of other grasses • Initially fix carbon using PEP carboxylase in mesophyll cells • Produces oxaloacetate, converted to malate, transported to bundle-sheath cells • Within the bundle-sheath cells, malate is decarboxylated to produce pyruvate and CO2 • Carbon fixation then by rubisco and the Calvin cycle 39 • C4 pathway, although it overcomes the problems of photorespiration, does have a cost • To produce a single glucose requires 12 additional ATP compared with the Calvin cycle alone • C4 photosynthesis is advantageous in hot dry climates where photorespiration would remove more than half of the carbon fixed by the usual C3 pathway alone 40 CAM plants • Many succulent (water-storing) plants, such as cacti, pineapples, and some members of about two dozen other plant groups • Stomata open during the night and close during the day – Reverse of that in most plants • Fix CO2 using PEP carboxylase during the night and store in vacuole 41 • When stomata closed during the day, organic acids are decarboxylated to yield high levels of CO2 • High levels of CO2 drive the Calvin cycle and minimize photorespiration 42 Compare C4 and CAM • Both use both C3 and C4 pathways • C4 – two pathways occur in different cells • CAM – C4 pathway at night and the C3 pathway during the day 43 Card Quiz B Which structure is responsible for gas exchange ? Matrix Stroma Stoma Thylakoid Question 4 Making glucose via photosynthesis costs ____ATPs and ___NADPHs. However, 1 molecule of glucose is enough to make ___ ATPs. 18, 12, 36 12, 18, 36 12, 36, 18 36, 18, 12 Question 9 What is the function of Rubisco? Absorption of photon energy Reduction of NADP+ Chemiosmosis Carbon fixation Card Quiz B Where could a botanist expect to find C4 plants? Canada Costa Rica Tundra Mount Everest Card Quiz B The end product of photosynthesis is the starting material of cellular respiration. a. This is true b. This is false Card Quiz B Which of the following would prevent the formation of phosphoglycerate? Denaturation of the Rubisco enzyme Non-functional b6-f complex Anaerobic conditions All of the above