Photosynthesis Chapter 10 Objectives Compare the overall reaction of photosynthesis with the overall reaction for respiration Describe where the processes of photosynthesis occur Describe the basic processes involved in photosynthesis: water splitting to obtain electrons, redox reactions of the electron transport chains, electron and energy shuttling by means of ATP and NADPH, and the coupling of the light-dependent reactions and the Calvin cycle Explain how pigments capture light and excite electrons Describe the structural and functional differences between photosystem I and photosystem II, and cyclic and noncyclic photophosphorylation Outline the steps in the cyclic fixation of carbon in the Calvin cycle and where these occur Contrast the C4 and CAM photosynthetic systems with the simpler C3 system, and how they are adaptions to hot, dry climates Introduction Overall equation is reversal of cellular respiration 6CO2+12H2O+energy--->C6H12O6+6O2+6H2O Increasingly probing studies provided knowledge about how photosynthesis works van Helmont-developed early ideas about where plants obtain materials for growth showed that soil not sufficient concluded that water important that plants restore “bad” air Ingenhousz-plants only restore air when exposed to light Priestly-showed Autotrophs are Producers Autotroph-means self-feeding applies to any organism that makes own food without eating, decomposing or absorbing other organisms or organic molecules Photosynthetic autotrophs include plants, algae and photosynthetic bacteria Site of Photosynthesis Photosynthesis occurs in chloroplasts in all photosynthetic organisms except monerans Leaves (specifically, mesophyll cells) are primary site of photosynthesis Light-absorbing pigment is chlorophyll located in protein complexes in internal membranes of chloroplasts Sugars assembled in stroma Underlying Processes Oxygen produced by splitting water demonstrated using 18O-labeled reactants given C18O2 does not release 18O2 plant given H218O does give off 18O2 plant Photosynthesis is redox process oxidized--->1/2O2+2H++2e CO2 reduced to glucose by addition of e-’s and H+’s compare with respiration where glucose oxidized and O2 reduced H2O In photosynthesis, electrons travel “uphill” from water to glucose, adding light energy captured by chlorophyll In respiration, electrons travel “downhill” from glucose to water, releasing energy to ATP Overview Photosynthesis is a two-stage process light-dependant reactions convert light energy to chemical energy, releases O2 as waste product occurs in thylakoid membranes and produces energy shuttles ATP and NADPH Calvin cycle cyclic series of steps that assemble organic molecules from CO2 occur in stroma and use energy and electrons from ATP and NADPH in carbon fixation light not required but usually run during day as require shuttles from light-dependant reactions The Light Reactions Driven by visible light light is electromagnetic radiation only small fraction of em radiation perceived by organisms different wavelengths=different colors leaf absorbs some wavelengths (red-orange and blue-violet) and reflects others (green) in plants light absorbed by chlorophyll a, chlorophyll b and carotenoids only chlorophyll a directly involved in light reactions; other pigments act as “antenna” molecules to broaden range of energy absorbed The Photosystems Light behaves like particles-photons When pigment absorbs photon, energy level of one electron is raised to excited, unstable state if pigment is isolated from molecular environment, excited electron loses energy as heat or light and returns to normal level chlorophyll fluoresces red In chloroplasts, 200-300 chlorophyll molecules grouped with proteins to form antenna assembly around two chlorophyll a molecules-reaction center chlorophylls excited electrons passed from antenna chlorophylls to reaction center chlorophylls then to primary electron acceptor series of redox reactions • final is oxidation of reaction center chlorophyll and reduction of primary electron acceptor Two photosystems (antenna assembly+primary electron acceptor) identified absorb at different wavelengths photosystem I-absorbs maximally at 700nm (P700) photosystem II-absorbs maximally at 680nm (P680) function together to carryout non-cyclic electron transport also known as non-cyclic photophosphorylation photosystem I can also carryout cyclic electron transport (cyclic photophosphorylation) thought to be the earliest form of photosynthesis • present in many primitive photosynthetic bacteria synthesizes only ATP Chemical Energy Generation Electron transport chains generate ATP, NADPH and O2 kinetic energy of light absorbed and excites electrons excited electrons passed along electron transport chain-series of redox reactions released energy used to generate ATP, NADPH and O2 production supplied of NADPH requires 2 electrons to PS I by PS II replaced in PS II by splitting water H2O ---> 1/2O2 + 2H+ + 2e- Chemiosmosis Powers ATP synthesis H+ ions from splitting water and those pumped across thylakoid membrane by electron transport chain form gradient across thylakoid membrane (inside to outside) ATP synthase provides port for H+ to diffuse back into stroma releases energy and phosphorylates ADP to ATP similar process to ATP generation in mitochondria known as photophosphorylation Carbon Fixation ATP and NADPH from light-dependant reactions power Calvin cycle net result of Calvin cycle is 3C molecules from CO2 using energy and electrons in ATP and NADPH from light-dependant reactions CO2 added to 5C intermediate ribulose-1,5bisphosphate (RuBP) catalyzed by RuBP carboxylase/oxygenase (rubisco) Number of rearrangements occur in many steps, using energy in ATP and oxidation of NADPH last step in cycle regenerates RuBP all steps occur simultaneously but ultimately regenerate starting reactants, hence cycle Three RuBP enter cycle for each 3C molecule released from chloroplast Calvin cycle occurs in chloroplast stroma 3C molecules exported to cytoplasm used to synthesize glucose and other organic molecules Plants that use only Calvin cycle to fix carbon called C3 plants first identifiable product of carbon fixation is 3C molecule Carbon-fixing Variations C3 plants conserve water by closing stomata allows buildup of O2 in leaves Rubisco fixes O2 rather than CO2 called photorespiration uses ATP and NADPH but makes no sugars C4 plants adapted to conserve water and prevent photorespiration CO2 incorporated into 4C molecule in mesophyll cells diffuses into bundle sheath cells and released enters Calvin cycle in bundle sheath chloroplasts CAM (crassulacean acid metabolism) plants incorporate carbon during night stomata open at night, closed during day CO2 incorporated in 4C molecule and stored in vacuole at night during day, 4C molecules exported into cytoplasm and CO2 released CO2 enters Calvin cycle C4 separate carbon incorporation and fixation spatially CAM plants separate carbon incorporation and carbon fixation temporally