Photosynthesis Chapter 10 A.P. Biology Liberty Senior High School Mr. Knowles Photosynthesis • Conversion of unusable light energy into usable chemical energy. C6H12O6 The Players Prokaryotic Photosynthesizers • Purple Sulfur Bacteria • Cyanobacteria Eukaryotic Photosynthesizers • Protists (Algae) • Plants (single-celled and multicellular) Photosynthesis Two Step Process: • Light-Dependent Reactions (Light Reactions)-produce ATP and NADPH. • Light-Independent Reactions (Calvin Cycle)-fix CO2 into sugars. • Light Reactions –Occur in the grana. –Split water, release oxygen, produce ATP, and form NADPH • The Calvin Cycle –Occurs in the stroma –Forms sugar from carbon dioxide, using ATP for energy and NADPH for reducing power • An overview of photosynthesis H2 O CO2 Light NADP ADP + P LIGHT REACTIONS CALVIN CYCLE ATP NADPH Chloroplast Figure 10.5 O2 [CH2O] (sugar) Light Dependent Reactions • Pigments absorb the light energy. • Different pigments are used in different organisms. • Each with different absorption spectra. • Why? • A photosystem – Is composed of a reaction center surrounded by a number of light-harvesting complexes Thylakoid Photosystem Photon STROMA Thylakoid membrane Light-harvesting complexes Primary election acceptor e– Transfer of energy Figure 10.12 Reaction center Special chlorophyll a molecules Pigment molecules THYLAKOID SPACE (INTERIOR OF THYLAKOID) • Light-harvesting complexes –Consist of pigment molecules bound to particular proteins. –Funnel the energy of photons of light to the reaction center. Light-Dependent Reactions • Photon captured by pigments arranged in a network with proteins embedded in a membrane- Photosystem (I and II). • In bacteria, this membrane is the cell membrane. • In protists (algae) and plants, this membrane is the thylakoid of the chloroplasts. Photosystems • In plants, the reaction center molecule is a type of chlorophyll a called P700 for photosystem l. • In the sulfur bacteria, the reaction center molecule is P870 for photosystem l. The First Photosynthesizers • The purple sulfur bacteria more than 3 billion years ago. • Used P870 in photosystem l. • The excited electron is ejected from the pigment and travels in a circular path. Used to power a proton pump to make ATP- chemiosmosis. • This circular movement is called - Cyclic Photophosphorylation. A Comparison of Chemiosmosis in Chloroplasts and Mitochondria • Chloroplasts and mitochondria – Generate ATP by the same basic mechanism: chemiosmosis – But use different sources of energy to accomplish this • The spatial organization of chemiosmosis – Differs in chloroplasts and mitochondria Key Higher [H+] Lower [H+] Chloroplast Mitochondrion CHLOROPLAST STRUCTURE MITOCHONDRION STRUCTURE H+ Diffusion Thylakoid space Intermembrance space Membrance Electron transport chain ATP Synthase Matrix Figure 10.16 ADP+ Stroma P H+ ATP • In both organelles – Redox reactions of electron transport chains generate a H+ gradient across a membrane • ATP synthase – Uses this proton-motive force to make ATP Cyclic Photophosphorylation. • Produces ATP from light energy. • Major limitation: no biosynthesis (no carbohydrates made from CO2. Therefore, no long term storage of energy. • These bacteria must find other sources of hydrogen to reduce CO2. • Inefficient. How do you make a better photosynthesizer? The Evolutionary Process Continues! Make another Photosystem-The Advent of Noncyclic Photosysthesis Noncyclic Electron Flow • Noncyclic electron flow –Is the primary pathway of energy transformation in the light reactions • Produces NADPH, ATP, and oxygen H2O CO2 Light NADP+ ADP CALVIN CYCLE LIGHT REACTIONS ATP NADPH O2 [CH2O] (sugar) Primary acceptor Primary acceptor H2O e 2 2 H+ e 3 NADPH 5 Light + H+ P700 P680 Light 6 ATP Figure 10.13 NADP+ + 2 H+ PC e– e– 1 e– 8 NADP+ reductase Cytochrome complex + O2 7 Fd 4 Pq Photosystem II (PS II) Photosystem-I (PS I) • A mechanical analogy for the light reactions e– ATP e– e– NADPH e– e– e– Mill makes ATP e– Figure 10.14 Photosystem II Photosystem I Enter the Competition! Cyanobacteria • Another photosystem was created. • Photosystem II uses a slightly different form of chlorophyll a called P680 (absorbs shorter wavelengths of light, more energy). • Excited electron enters the electron transport chain and powers proton pumps. • This leads to chemiosmosis and the synthesis of ATP. Photosystem II • The excited electron is eventually passed to the P700 molecule in Photosystem I. • Next, Photosystem I can absorb another photon and excites an electron. • This electron provides the reducing power in the form of NADPH. Photosystems II and I • Together, these are called Noncyclic Photophosphorylation. • Why does II come before I? • Why does noncyclic photo. Produce NADPH instead of NADH? • How is the excited electron replaced in the P680 of Photosystem II? Making Oxygen Gas! • The P680 is a strong oxidizer and it removes an e- from the Z protein. • The Z protein obtains an e from H2O. Z enzyme + • H2 O H + OH • OH- reassembled into O2 and H2O. + • H remain in the thylakoid space. In summary… • The Light Reactions: Photosystem II makes ATP from light energy. Photosystem I makes NADPH from light energy. Both occur in the thylakoid spaces of chloroplasts. Cyclic Electron Flow • Under certain conditions -Photoexcited electrons take an alternative path. -Calvin cycle requires more ATP and NADPH. -This path makes up the difference. • In cyclic electron flow – Only photosystem I is used – Only ATP is produced Primary acceptor Primary acceptor Fd Fd NADP+ Pq NADP+ reductase Cytochrome complex NADPH Pc Figure 10.15 Photosystem II ATP Photosystem I • The light reactions and chemiosmosis: the organization of the thylakoid membrane H2O CO2 LIGHT NADP+ ADP CALVIN CYCLE LIGHT REACTOR ATP NADPH STROMA (Low H+ concentration) O2 [CH2O] (sugar) Cytochrome complex Photosystem II Photosystem I NADP+ reductase Light 2 H+ 3 NADP+ + 2H+ Fd NADPH + H+ Pq Pc 2 H2O THYLAKOID SPACE (High H+ concentration) 1⁄ 2 1 O2 +2 H+ 2 H+ To Calvin cycle STROMA (Low H+ concentration) Thylakoid membrane ATP synthase ADP ATP P Figure 10.17 H+ World’s Largest Organism! The Dark Reactions • Use the ATP and the NADPH made in the light reactions and build sugar molecules from the CO2 in the atmosphere. • Needs a source of energy (from ATP) and the H to reduce the CO2 (from NADPH). • The Calvin Cycle, Fig. 10.12 and 10.13. • Concept 10.3: The Calvin cycle uses ATP and NADPH to convert CO2 to sugar • The Calvin cycle – Is similar to the citric acid cycle – Occurs in the stroma • The Calvin Cycle has three phases – Carbon fixation – Reduction – Regeneration of the CO2 acceptor The Calvin Cycle H2O CO2 Input (Entering one 3 at a time) CO2 Light NADP+ ADP CALVIN CYCLE LIGHT REACTION ATP Phase 1: Carbon fixation NADPH Rubisco O2 [CH2O] (sugar) 3 P P Short-lived intermediate 3 P P Ribulose bisphosphate (RuBP) P 6 3-Phosphoglycerate 6 ATP 6 ADP CALVIN CYCLE 3 ADP 3 6 P ATP P 1,3-Bisphoglycerate 6 NADPH Phase 3: Regeneration of the CO2 acceptor (RuBP) 6 NADPH+ 6 P P 5 (G3P) 6 P Glyceraldehyde-3-phosphate (G3P) P 1 Figure 10.18 G3P (a sugar) Output Glucose and other organic compounds Phase 2: Reduction The Calvin Cycle • Also called the C3 cycle because of a 3-carbon molecule (PGA). • It is the reverse of glycolysis. • Taking C atoms from atmospheric CO2 and making sugars-Carbon Fixation. • Occurs in the stroma. The Calvin Cycle • First step is dependent on RuBP carboxylase enzyme (RUBISCO) which adds C from CO2. • When the temperature >28° C or the concentration of CO2 falls, RUBISCO will also oxidize instead of adding C’s- Photorespiration- the opposite of the Calvin Cycle. RUBISCO Activity >28° C Photorespiration • Loses 1/4 to 1/2 of all fixed C that enters the Calvin Cycle. • The C3 plants would not efficiently photosynthesize in tropical climates. • Enter the competition! • Concept 10.4: Alternative mechanisms of carbon fixation have evolved in hot, arid climates • On hot, dry days, plants close their stomata – Conserving water but limiting access to CO2 – Causing O2 to build up Photorespiration: An Evolutionary Relic? • In photorespiration –O2 substitutes for CO2 in the active site of the enzyme rubisco. –The photosynthetic rate is reduced. C3 Leaf Structure C4 Leaf Structure C4 Plants • C4 plants minimize the cost of photorespiration –By incorporating CO2 into four carbon compounds in mesophyll cells –Are exported to bundle sheath cells, where they release CO2 used in the Calvin cycle C4 leaf anatomy and the C4 pathway Mesophyll cell Mesophyll cell Photosynthetic cells of C4 plant leaf CO CO 2 2 PEP carboxylase Bundlesheath cell PEP (3 C) Oxaloacetate (4 C) ADP Vein (vascular tissue) Malate (4 C) ATP C4 leaf anatomy BundleSheath cell Pyruate (3 C) CO2 Stoma CALVIN CYCLE Sugar Vascular tissue Figure 10.19 Comparsion of C3 and C4 Plants C3 Plants • Are temperate • plants. • Perform light and • dark reactions in the same cellmesophyll cells. C4 Plants Are tropical plants. (Corn and sugar cane). Perform light reactions in mesophyll cells but the Calvin cycle in bundle sheath cells. The Cost to C4 Plants • It costs 2 ATP to transport each CO2 into a bundle sheath cell. • The energetic cost of C4 Photosynthesis is twice that of C3 Photosynthesis. • Photosynthesis is advantegous in hot climates. • C4 plants outcompete C3 plants in tropical climates. C3 Leaf Structure C4 Leaf Structure What about plants in extremely hot climates? Must conserve water! Transpiration • Water loss from the leaf tissue through the stomata. (Analogy: Sweating.) CAM Plants • CAM plants –Open their stomata at night, incorporating CO2 into organic acids • During the day, the stomata close –And the CO2 is released from the organic acids for use in the Calvin cycle CAM Plants • Succulent plants must conserve water loss from stomata. Cacti and pineapple use another strategy. • Crassulacean Acid Metabolism (CAM) Plants. • The stomata are closed during the day to prevent water loss and reduce photorespiration by preventing CO2 from leaving the leaf. Open the stomata at night. CAM Plants • Perform C3 and C4 pathways in the same cell (mesophyll) but at different times. • The C4 pathway is used at night when the stomata are open. Prevent CO2 losses • The C3 pathway is used during the day when the stomata are closed and there is a need to reduce water loss. The CO2 for making sugars during the day come from organic molecules made during the previous night, none from atmosphere. The CAM pathway is similar to the C4 pathway Pineapple Sugarcane C4 Mesophyll Cell Organic acid Bundlesheath cell (a) Spatial separation of steps. In C4 plants, carbon fixation and the Calvin cycle occur in different types of cells. Figure 10.20 CAM CO2 CO2 CALVIN CYCLE Sugar 1 CO2 incorporated Organic acid into four-carbon organic acids (carbon fixation) 2 Organic acids release CO2 to Calvin cycle CALVIN CYCLE Sugar Night Day (b) Temporal separation of steps. In CAM plants, carbon fixation and the Calvin cycle occur in the same cells at different times. The Importance of Photosynthesis: A Review Light reaction Calvin cycle H2O CO2 Light NADP+ ADP +P1 RuBP 3-Phosphoglycerate Photosystem II Electron transport chain Photosystem I ATP NADPH G3P Starch (storage) Amino acids Fatty acids Chloroplast Figure 10.21 O2 Light reactions: • Are carried out by molecules in the thylakoid membranes • Convert light energy to the chemical energy of ATP and NADPH • Split H2O and release O2 to the atmosphere Sucrose (export) Calvin cycle reactions: • Take place in the stroma • Use ATP and NADPH to convert CO2 to the sugar G3P • Return ADP, inorganic phosphate, and NADP+ to the light reactions