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Photosynthesis Chapter 10 Overview: The Process That Feeds the Biosphere Photosynthesis is the process that converts light energy into chemical energy Directly or indirectly, photosynthesis nourishes almost the entire living world Photosynthesis Autotrophs: Organisms that produce their own organic substances from inorganic substances using energy from the atmosphere Plants, algae, and some bacteria are photoautotrophs Photoautotrophs The first photoautotrophs were the cyanobacteria These organisms were responsible for oxygenating the Earth’s atmosphere Utilize sunlight for energy (sunlight is converted to ATP to fuel the process of making sugars) Photoautotrophs Pigments, such as chlorophyll, absorb light energy Photon = A measurable unit of light Photosynthetic pigments include chlorophylls (a, b, c), carotenoids (orange and red) and xanthophylls (gold and yellow) Chloroplasts Chloroplasts are the sites of photosynthesis in plants and most algae Chloroplasts are found in the mesophyll (ground) tissue of leaves A typical mesophyll cell has 30-40 chloroplasts Chloroplast Structure Chloroplasts have a double membrane (inner and outer) A space filled with fluid lies between the two membranes Inside of the chloroplasts are thylakoids, arranged in stacks called grana, which are surrounded by a clear, thick fluid called stroma Leaf Structure The petiole is the attachment point of the leaf to the plant The blade is the “leafy” portion Leaves, like all parts of the plant body, consist of three major tissue types: epidermis, ground (middle) tissue and vascular (transport) tissue Leaf Structure Openings on the underside of the leaf, called stomata, allow for the passage of gasses into and out of the leaf Carbon dioxide enters the leaf Oxygen, produced by cells in the leaf, exits Where does the oxygen come from? Carbon dioxide is absorbed by plants, but the oxygen that is released comes from the splitting of water 6 CO2 + 12 H2O + Light Energy C6H12O6 + 6 H2O + 6 O2 A few other things to know... Photosynthesis has two major phases: the Light Reactions (the photo part) and the Calvin Cycle (the synthesis part) The light reactions (in the thylakoids) split water, release O2, produce ATP, and form NADPH The Two Stages of Photosynthesis: A Preview The Calvin cycle (in the stroma) forms sugar from CO2, using ATP and NADPH Instead of NAD+, a substance called NADP+ accepts electrons in photosynthesis (hey, think of it as P for Photosynthesis!) A few other things to know... Photophosphorylation: Adding a phosphate group to a molecule using energy from sunlight. Carbon fixation: Incorporation of carbon from CO2 in the air into organic molecules such as sugar. Sunlight Light can be measured in packets called photons Photosynthetic pigments, such as chlorophyll, absorb parts of the visible light spectrum more readily than others. Visible Light Energy decreases from violet to red, while wavelength increases along the same path This means that energy is inversely proportional to wavelength Reflected Light The portion of the visible light spectrum that is reflected is the color that the object appears Chlorophyll is green since it absorbs other colors and reflects green The Light Reactions Begin with the absorption of light energy by thylakoids, where light energy is transformed into the chemical energy of ATP and NADPH This is called the excitation of chlorophyll by light energy CH3 CHO in chlorophyll a in chlorophyll b Porphyrin ring: light-absorbing “head” of molecule; note magnesium atom at center Hydrocarbon tail: interacts with hydrophobic regions of proteins inside thylakoid membranes of chloroplasts; H atoms not shown H2O Light LIGHT REACTIONS Chloroplast H2O Light LIGHT REACTIONS ATP NADPH Chloroplast O2 H2O CO2 Light NADP+ ADP + Pi LIGHT REACTIONS CALVIN CYCLE ATP NADPH Chloroplast O2 [CH2O] (sugar) Photosystems A photosystem consists of a reaction center surrounded by light-harvesting complexes The light-harvesting complexes (pigment molecules bound to proteins) funnel the energy of photons to the reaction center Photosystems • A primary electron acceptor in the reaction center accepts an excited electron from chlorophyll a Solar-powered transfer of an electron from a chlorophyll a molecule to the primary electron acceptor is the first step of the light reactions Thylakoid Photosystem Photon Thylakoid membrane Light-harvesting complexes Reaction center STROMA Primary electron acceptor e– Transfer of energy Special chlorophyll a molecules Pigment molecules THYLAKOID SPACE (INTERIOR OF THYLAKOID) Photosystems • Photosystem II functions first and is best at absorbing a wavelength of 680 nm • Photosystem I is best at absorbing a wavelength of 700 nm • The two photosystems work together to use light energy to generate ATP and NADPH Noncyclic Electron Flow During the light reactions, there are two possible routes for electron flow: cyclic and noncyclic Noncyclic electron flow, the primary pathway, involves both photosystems and produces ATP and NADPH H2O CO2 Light NADP+ ADP CALVIN CYCLE LIGHT REACTIONS ATP NADPH O2 [CH2O] (sugar) Primary acceptor Energy of electrons e– Light P680 Photosystem II (PS II) H2O CO2 Light NADP+ ADP CALVIN CYCLE LIGHT REACTIONS ATP NADPH O2 [CH2O] (sugar) Energy of electrons Primary acceptor 2 H+ 1/ 2 + O2 Light H2O e– e– e– P680 Photosystem II (PS II) H2O CO2 Light NADP+ ADP CALVIN CYCLE LIGHT REACTIONS ATP NADPH O2 [CH2O] (sugar) Primary acceptor Energy of electrons Pq 2 H+ + 1/ 2 O 2 Light H2O e– Cytochrome complex Pc e– e– P680 ATP Photosystem II (PS II) H2O CO2 Light NADP+ ADP CALVIN CYCLE LIGHT REACTIONS ATP NADPH O2 [CH2O] (sugar) Primary acceptor Primary acceptor e– Energy of electrons Pq 2 H+ 1/ 2 + O2 Light H2O e– Cytochrome complex Pc e– e– P700 P680 Light ATP Photosystem II (PS II) Photosystem I (PS I) H2 O CO2 Light NADP+ ADP CALVIN CYCLE LIGHT REACTIONS ATP NADPH O2 [CH2O] (sugar) Primary acceptor Primary acceptor e– Pq Energy of electrons 2 H+ e– H2O Cytochrome complex + 1/2 O2 Light Fd e– e– NADP+ reductase Pc e– e– NADPH + H+ P700 P680 Light ATP Photosystem II (PS II) NADP+ + 2 H+ Photosystem I (PS I) e– ATP e– e– NADPH e– e– e– Mill makes ATP e– Photosystem II Photosystem I Cyclic Electron Flow • Cyclic electron flow uses only photosystem I and produces only ATP Cyclic electron flow generates surplus ATP, satisfying the demand in the Calvin cycle Primary acceptor Primary acceptor Fd Fd NADP+ Pq NADP+ reductase Cytochrome complex NADPH Pc Photosystem I Photosystem II ATP The Calvin Cycle The Calvin cycle, like the citric acid cycle, regenerates its starting material after molecules enter and leave the cycle The cycle builds sugar from smaller molecules by using ATP and NADPH The Calvin Cycle Carbon enters the cycle as CO2 and leaves as a sugar named glyceraldehyde-3-phosphate (G3P) For net synthesis of one G3P, the cycle must take place three times, fixing three molecules of CO2 The Calvin Cycle The Calvin cycle has three phases: Carbon fixation (catalyzed by Rubisco) Reduction Regeneration of the CO2 acceptor (RuBP) H2 O CO2 Input Light (Entering one CO2 at a time) 3 NADP+ ADP CALVIN CYCLE LIGHT REACTIONS ATP Phase 1: Carbon fixation NADPH Rubisco O2 [CH2O] (sugar) 3 P Short-lived intermediate P P 6 3-Phosphoglycerate 3 P P Ribulose bisphosphate (RuBP) 6 6 ADP CALVIN CYCLE ATP H2O CO2 Input Light (Entering one CO2 at a time) 3 NADP+ ADP CALVIN CYCLE LIGHT REACTIONS ATP Phase 1: Carbon fixation NADPH Rubisco O2 [CH2O] (sugar) 3 P P Short-lived intermediate 3 P P 6 P 3-Phosphoglycerate Ribulose bisphosphate (RuBP) 6 ATP 6 ADP CALVIN CYCLE 6 P P 1,3-Bisphosphoglycerate 6 NADPH 6 NADP+ 6 Pi 6 P Glyceraldehyde-3-phosphate (G3P) 1 P G3P (a sugar) Output Glucose and other organic compounds Phase 2: Reduction H2O CO2 Input Light (Entering one CO2 at a time) 3 NADP+ ADP CALVIN CYCLE LIGHT REACTIONS ATP Phase 1: Carbon fixation NADPH Rubisco O2 [CH2O] (sugar) 3 P P Short-lived intermediate 3 P P 6 P 3-Phosphoglycerate Ribulose bisphosphate (RuBP) 6 ATP 6 ADP 3 ADP 3 CALVIN CYCLE 6 P ATP P 1,3-Bisphosphoglycerate 6 NADPH Phase 3: Regeneration of the CO2 acceptor (RuBP) 6 NADP+ 6 Pi P 5 G3P 6 P Glyceraldehyde-3-phosphate (G3P) 1 P G3P (a sugar) Output Glucose and other organic compounds Phase 2: Reduction Calvin cycle animation: http://www.science.smith.edu/departments/ Biology/Bio231/calvin.html Alternative mechanisms of carbon fixation Dehydration is a problem for plants, sometimes requiring tradeoffs with other metabolic processes, especially photosynthesis On hot, dry days, plants close stomata, which conserves water but also limits photosynthesis Alternative mechanisms of carbon fixation The closing of stomata reduces access to CO2 and causes O2 to build up These conditions favor a seemingly wasteful process called photorespiration Photorespiration: An Evolutionary Relic? • In most plants (C3 plants), initial fixation of CO2, via the enzyme rubisco, forms a three-carbon compound • In photorespiration, rubisco adds O2 to the Calvin cycle instead of CO2 Photorespiration: An Evolutionary Relic? Photorespiration consumes O2 and organic fuel and releases CO2 without producing ATP or sugar • Photorespiration may be an evolutionary relic because rubisco first evolved at a time when the atmosphere had far less O2 and more CO2 Photorespiration: An Evolutionary Relic? In many plants, photorespiration is a problem because on a hot, dry day it can drain as much as 50% of the carbon fixed by the Calvin cycle C4 Plants C4 plants minimize the cost of photorespiration by incorporating CO2 into four-carbon compounds in mesophyll cells These four-carbon compounds are exported to bundle-sheath cells, where they release CO2 that is then used in the Calvin cycle LE 10-19 Photosynthetic cells of C4 plant leaf Mesophyll cell PEP carboxylase Mesophyll cell CO2 Bundlesheath cell The C4 pathway Oxaloacetate (4 C) PEP (3 C) Vein (vascular tissue) ADP Malate (4 C) ATP C4 leaf anatomy Stoma Bundlesheath cell Pyruvate (3 C) CO2 CALVIN CYCLE Sugar Vascular tissue CAM Plants CAM plants open their stomata at night, incorporating CO2 into organic acids Stomata close during the day, and CO2 is released from organic acids and used in the Calvin cycle Sugarcane Pineapple CAM C4 CO2 Mesophyll cell Organic acid Bundlesheath cell CO2 CO2 incorporated into four-carbon Organic acid organic acids (carbon fixation) CO2 CALVIN CYCLE Sugar Spatial separation of steps CO2 Organic acids release CO2 to Calvin cycle Night Day CALVIN CYCLE Sugar Temporal separation of steps The Importance of Photosynthesis: A Review The energy entering chloroplasts as sunlight gets stored as chemical energy in organic compounds Sugar made in the chloroplasts supplies chemical energy and carbon skeletons to synthesize the organic molecules of cells The Importance of Photosynthesis: A Review In addition to food production, photosynthesis produces the oxygen in our atmosphere Light reactions Calvin cycle H2O CO2 Light NADP+ ADP + Pi RuBP Photosystem II Electron transport chain Photosystem I ATP NADPH 3-Phosphoglycerate G3P Starch (storage) Amino acids Fatty acids Chloroplast O2 Sucrose (export)