Photosynthesis Photosynthesis in nature • • Autotrophs: biotic producers; 1. Photoautotrophs: transform sunlight energy into chemical energy by combining it with CO2 and H2O . 2. Chemoautotrophs; obtain inorganic nutrients and combine it with CO2 to make organic food. Heterotrophs: biotic consumers; obtains organic food by eating other organisms or their by-products (includes decomposers). Photosynthesis • Occurs in plants, algae, certain other protists, and some prokaryotes These organisms use light energy to drive the synthesis of organic molecules from carbon dioxide and (in most cases) water. They feed not only themselves, but the entire living world. (a) On land, plants are the predominant producers of food. In aquatic environments, photosynthetic organisms include (b) multicellular algae, such as this kelp; (c) some unicellular protists, such as Euglena; (d) the prokaryotes called cyanobacteria; and (e) other photosynthetic prokaryotes, such as these purple sulfur bacteria, which produce sulfur (spherical globules) (c, d, e: LMs). (a) Plants (c) Unicellular protist 10 m (e) Pruple sulfur bacteria Figure 10.2 (b) Multicellular algae (d) Cyanobacteria 40 m 1.5 m The chloroplast Sites of photosynthesis • Organ: leaves (major site) – Gas exchange: stomata • Plant cell: mesophyll • Organelle: chloroplast – Double membrane – Thylakoids, grana, stroma – Pigment: chlorophyll Summary of Photosynthesis Light energy 6 CO2 + 12 H2O C6H12O6 + 6 O2 + 6 H2O Redox reactions • Photosynthesis is a redox process - water is oxidized, e- (along w/ H+) are transferred to CO2, reducing it to sugar Reactants: Products: Figure 10.4 12 H2O 6 CO2 C6H12O6 6 H2O 6 O2 Photosynthesis: an overview 2 major steps: 1. light reactions (“photo”) – NADP+ (electron acceptor) to NADPH – Photophosphorylation: ADP ---> ATP 2. Calvin cycle (“synthesis”) – Carbon fixation: carbon into organics The Nature of Sunlight • Light – Is a form of electromagnetic energy, which travels in waves • Wavelength – Is the distance between the crests of waves – Determines the type of electromagnetic energy The Nature of Sunlight • The electromagnetic spectrum – Is the entire range of electromagnetic energy, or radiation • The visible light spectrum – Includes the colors of light we can see – Includes the wavelengths that drive photosynthesis The Nature of Sunlight The Electromagnetic Spectrum 10–5 nm 10–3 nm Gamma rays X-rays UV 1m 106 nm 106 nm 103 nm 1 nm Infrared Microwaves 103 m Radio waves Visible light 380 450 500 Shorter wavelength Figure 10.6 Higher energy 550 600 650 700 Longer wavelength Lower energy 750 nm The Nature of Sunlight • Pigments Light – Are substances that absorb visible light – The also reflect light, which include the colors we see Figure 10.7 Reflected Light Chloroplast Absorbed light Granum Transmitted light Which Wavelengths of Light are Important for Photosynthesis? Three different experiments helped reveal which wavelengths of light are photosynthetically important. The results are shown below. EXPERIMENT RESULTS Absorption of light by chloroplast pigments Chlorophyll a Chlorophyll b Carotenoids Wavelength of light (nm) Chlorophyll b and carotenoids are called accessory pigments; they can absorb colors that chlorophyll a can not. They allow plants to capture more of the energy in light. Rate of photosynthesis (measured by O2 release) Which Wavelengths of Light are Important for Photosynthesis? (b) Action spectrum. This graph plots the rate of photosynthesis versus wavelength. The resulting action spectrum resembles the absorption spectrum for chlorophyll a but does not match exactly (see part a). This is partly due to the absorption of light by accessory pigments such as chlorophyll b and carotenoids. Which Wavelengths of Light are Important for Photosynthesis? Aerobic bacteria Filament of alga 500 600 700 400 (c) Engelmann‘s experiment. In 1883, Theodor W. Engelmann illuminated a filamentous alga with light that had been passed through a prism, exposing different segments of the alga to different wavelengths. He used aerobic bacteria, which concentrate near an oxygen source, to determine which segments of the alga were releasing the most O2 and thus photosynthesizing most. Bacteria congregated in greatest numbers around the parts of the alga illuminated with violet-blue or red light. Notice the close match of the bacterial distribution to the action spectrum in part b. CONCLUSION photosynthesis. Light in the violet-blue and red portions of the spectrum are most effective in driving Chlorophyll • Chlorophyll a in chlorophyll a in chlorophyll b CH2 – Is the main photosynthetic pigment • Chlorophyll b, Xanthophyll (yellow) and Carotene (orange), – Are the major accessory pigments • Phycoerythrins (red) and Fucoxanthins (brown) – Are minor accessory pigments CH3 CHO Figure 10.10 CH C H3C C C C H C C N C N C Mg N C C C C H C N C H3C CH3 H CH2 H H C C C O C H C CH3 CH3 Porphyrin ring: Light-absorbing “head” of molecule note magnesium atom at center C O O CH2 C C CH2 C O O CH3 CH2 Hydrocarbon tail: interacts with hydrophobic regions of proteins inside thylakoid membranes of chloroplasts: H atoms not shown Pigments • • What you should have seen from Activity A of the lab. Rf = pigment travel distance/solvent travel distance. 1. 2. 3. 4. Carotene (greatest/farthest) Xanthophyll Chlorophyll a Chlorophyll b (least/shortest) Photosystems • Light harvesting units of the thylakoid membrane • Composed mainly of protein and pigment antenna complexes • Antenna pigment molecules are struck by photons. Their main role is to harvest photons and transfer light energy to the reaction center. • Energy is passed to reaction centers (redox location) • Excited e- from chlorophyll is trapped by a primary eacceptor Noncyclic electron flow • Photosystem II (P680): – photons excite chlorophyll e- to an acceptor – e- are replaced by splitting of H2O (release of O2) – e-’s travel to Photosystem I down an electron transport chain (Pq~cytochromes~Pc) – as e- fall, ADP ---> ATP (noncyclic photophosphorylation) • Photosystem I (P700): – “fallen” e- replace excited e- to primary e- acceptor – 2nd ETC ( Fd~NADP+ reductase) transfers e- to NADP+ ---> NADPH (...to Calvin cycle…) • These photosystems produce equal amounts of ATP and NADPH The light reactions and chemiosmosis: the organisation of the thylakoid membrane H2O CO2 LIGHT NADP+ ADP LIGHT REACTOR Chemiosmosis: The process of making ATP that STROMA (Low H concentration) occurs from energy created by making a proton concentration HO (H+) gradient THYLAKOID SPACE across (High H concentration) membranes. O2 [CH2O] (sugar) Cytochrome Photosystem II complex Photosystem I NADP+ reductase Light 2 H+ Fd NADP+ + 2H+ Pc 2 1⁄ 2 1 O2 +2 H+ 2 H+ To Calvin cycle Thylakoid membrane ATP synthase ADP ATP P Figure 10.17 3 NADPH + H+ Pq 2 STROMA (Low H+ concentration) ATP NADPH + + CALVIN CYCLE H+ The Calvin cycle • • • Takes place in Stroma 3 molecules of CO2 are ‘fixed’ into glyceraldehyde 3phosphate (G3P) Phases: 1. Carbon fixation - each CO2 is attached to RuBP (rubisco enzyme) 2. Reduction - electrons from NADPH reduces to G3P; ATP used up 1. Regeneration - G3P rearranged to RuBP; ATP used; cycle continues Calvin Cycle, net synthesis • For each G3P (and for 3 CO2)……. Consumption of 9 ATP’s & 6 NADPH (light reactions regenerate these molecules) • G3P can then be used by the plant to make glucose and other organic compounds Cyclic electron flow • Alternative cycle when ATP is deficient • Photosystem I used but not II; produces ATP but no NADPH • Why? The Calvin cycle consumes more ATP than NADPH……. • Cyclic photophosphorylation Alternative carbon fixation methods, I • Photorespiration: hot/dry days; stomata close; CO2 decrease, O2 increase in leaves; O2 added to rubisco; no ATP or food generated • Two Solutions….. • 1- C4 plants: two photosynthetic cells, bundlesheath & mesophyll; PEP carboxylase (instead of rubisco) fixes CO2 in mesophyll; new 4C molecule releases CO2 (grasses, corn, sugar cane). Alternative carbon fixation methods, II • 2- CAM plants: open stomata during night, close during day (crassulacean acid metabolism); cacti, pineapples, etc. A review of photosynthesis 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 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 • 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 A Comparison of Chemiosmosis in Chloroplasts and Mitochondria • The spatial organisation of chemiosmosis Key Higher [H+] Lower [H+] Chloroplast Mitochondrion – Differs in chloroplasts and MITOCHONDRION STRUCTURE mitochondria Intermembrance space Membrance Matrix Figure 10.16 CHLOROPLAST STRUCTURE H+ Diffusion Electron transport chain ATP Synthase ADP+ Thylakoid space Stroma P H+ ATP