Warm-up Objective: Describe how the chemical products of the light-trapping reactions couple to the synthesis of carbohydrates. Warm-up: What is the advantage of the light reaction producing H and ATP on the stroma side of the thylakoid membrane? Stomates Remember what it means to be a plant Photosynthesis: The Calvin Cycle Life from Air 2006-2007 Need to produce all organic molecules necessary for growth carbohydrates, lipids, proteins, nucleic acids Need to store chemical energy (ATP) produced from light reactions in stable form can be moved around plant saved for a rainy day Autotrophs Making energy & organic molecules from light energy photosynthesis carbon + water + energy + oxygen dioxide 6CO + 2 6H 2 O + light C 6 H 12 O 6 6O energy + 2 Light reactions Convert solar energy to chemical energy ATP energy H reducing power What can we do now? build stuff!! 1
How is that helpful? Want to make C 6 H 12 O 6 synthesis How? From what? What raw materials are available? From C 6 H 12 O 6 has very little chemical energy fully oxidized C 6 H 12 O 6 contains a lot of chemical energy reduced endergonic carbon fixation C 6 H 12 O 6 H reduce Reduction of C 6 H 12 O 6 proceeds in many small uphill steps each catalyzed by specific enzyme using energy stored in ATP & H From Light reactions to Calvin cycle Calvin cycle chloroplast stroma Need products of light reactions to drive synthesis reactions ATP H starch, sucrose, cellulose & more Calvin cycle 3. Regeneration of C C ribulose bisphosphate 3 ATP 3 ADP used to make C C C C 1C 1. Carbon fixation ribulose bisphosphate carboxylase 6C glyceraldehyde-3-p PGA phosphoglycerate C = C = C 2. Reduction 6 ATP H H H C C C 6 H 6 ADP 6 H H H C C C Remember? glycolysis C-C-C-C-C-C fructose-6p P-C-C-C-C-C-C-P DHAP P-C-C-C C-C-C-P pyruvate C-C-C 2 ATP 2 ADP 2 NAD + 2 NADH 4 ADP 4 ATP Calvin Cycle Ribulose 1,5-bisphosphate () () 3 ADP 3 ATP 2P i Reforming Glyceraldehyde 3-phosphate () Carbon dioxide () Carbon fixation Glyceraldehyde 3-phosphate () () Glucose and other sugars Stroma of chloroplast 3-phosphoglycerate () (PGA) 6 ATP 6 ADP 1,3-bisphosphoglycerate () 6 H Reverse of glycolysis 6 + 6P i Glyceraldehyde 3-phosphate () () 2
To G-3-P and Beyond! Glyceraldehyde-3-P end product of Calvin cycle energy rich 3 carbon sugar C3 photosynthesis G-3-P = important intermediate G-3-P carbohydrates lipids amino acids nucleic acids Enzyme which fixes carbon from air ribulose bisphosphate carboxylase the most important enzyme in the world! it makes life out of air! definitely the most abundant enzyme Accounting The accounting is complicated 3 turns of Calvin cycle = 1 3 1 () 6 turns of Calvin cycle = 1 C 6 H 12 O 6 (6C) 6 1 C 6 H 12 O 6 (6C) 18 ATP + 12 H 1 C 6 H 12 O 6 any ATP left over from light reactions will be used elsewhere by the cell Photosynthesis summary Light reactions produced ATP produced H consumed H 2 O produced O 2 as byproduct Calvin cycle consumed produced (sugar) regenerated ADP regenerated ADP Summary of photosynthesis Where did the come from? Where did the go? Where did the H 2 O come from? Where did the H 2 O go? Where did the energy come from? What s the energy used for? What will the C 6 H 12 O 6 be used for? Where did the O 2 come from? Where will the O 2 go? What else is involved not listed in this equation? 6CO + 2 6H 2 O + light C 6 H 12 O 6 6O energy + 2 Supporting a biosphere On global scale, photosynthesis is the most important process for the continuation of life on Earth each year photosynthesis synthesizes 160 billion tons of carbohydrate heterotrophs are dependent on plants as food source for fuel & raw materials 3
The poetic perspective All the solid material of every plant was built by sunlight out of thin air All the solid material of every animal was built from plant material Any Questions?? air sun Then all the cats, dogs, rats, people & elephants are really strands of air woven together by sunlight! 2006-2007 Photosynthesis: Variations on the Theme Remember what plants need Photosynthesis light reactions light sun H 2 O ground Calvin cycle air 2006-2007 What structures have plants evolved to supply these needs? vascular bundle Leaf Structure xylem (water) cuticle phloem (sugar) epidermis palisades layer spongy layer Transpiration O 2 H2 O stomate O 2 H 2 O guard cell A second inside a leaf Gas exchange & water flow in for Calvin cycle O 2 out waste from light reactions H 2 O out for light reactions photosynthesis O 2 CO2 gas exchange water loss H 2 O O 2 xylem (water) phloem (sugars) 4
Controlling water loss from leaves Hot or dry days stomates close to conserve water guard cells gain H 2 O = stomates open lose H 2 O = stomates close adaptation to living on land, but creates PROBLEMS! Closed stomates Closed stomates lead to O 2 builds up from light reactions is depleted in Calvin cycle causes problems in Calvin Cycle Inefficiency of : vs O 2 in Calvin cycle carbon fixation enzyme normally bonds C to reduction of building sugars photosynthesis when O 2 concentration is high bonds O to O 2 is alternative substrate photorespiration oxidation of breakdown sugars to make Calvin cycle review ATP ADP H 1C 6C unstable intermediate PGA ATP ADP to make Calvin cycle review 1C C3 plants PGA Calvin cycle with O 2 A good enzyme, gone BAD! O 2 photorespiration 2C to mitochondria lost as without making ATP 5
Impact of Photorespiration Oxidation of short circuit of Calvin cycle loss of carbons to can lose 50% of carbons fixed by Calvin cycle reduces production of photosynthesis no ATP (energy) produced no C 6 H 12 O 6 (food) produced if photorespiration could be reduced, plant would become 50% more efficient strong selection pressure to evolve alternative systems Reducing photorespiration Separate carbon fixation from Calvin cycle C4 plants physically separate carbon fixation from actual Calvin cycle different enzyme to capture PEP carboxylase stores carbon in 4C compounds different leaf structure CAM plants separate carbon fixation from actual Calvin cycle by time of day fix carbon (capture ) during night store carbon in organic acids perform Calvin cycle during day C4 plants A better way to capture 1st step before Calvin cycle, fix carbon with enzyme PEP carboxylase store as 4C compound adaptation to hot, dry climates have to close stomates a lot different leaf anatomy sugar cane, corn, other grasses C4 Plants corn sugar cane PEP carboxylase light reactions O 2 Comparative anatomy Separate reactions in different cells light reactions carbon fixation Calvin cycle C3 C4 Location, location, location! PEP carboxylase enzyme higher affinity for than O 2 (better than ) fixes in 4C compounds regenerates in inner cells for phosphoenolpyruvate () + oxaloacetate (4C) 6
C4 photosynthesis Physically separated C fixation from Calvin cycle Outer cells light reaction & carbon fixation pumps to inner cells keeps O 2 away from inner cells away from Inner cells O 2 O 2 CO Calvin cycle 2 to veins CAM (Crassulacean Acid Metabolism) plants Different adaptation to hot, dry climates separate carbon fixation from Calvin cycle by time close stomates during day open stomates during night at night, open stomates & fix carbon in storage compounds organic acids: malic acid, isocitric acid in day, close stomates & release from storage compounds to Calvin cycle increases concentration of in cells succulents, some cacti, pineapple CAM plants C4 vs CAM Summary solves / O 2 gas exchange vs. H 2 O loss challenge C4 plants separate 2 steps of C fixation anatomically in 2 different cells CAM plants separate 2 steps of C fixation temporally at 2 different times Why the C3 problem? Possibly evolutionary baggage evolved in high atmosphere there wasn t strong selection against active site of accepting both & O 2 Today it makes a difference Any Questions?? 21% O 2 vs. 0.03% photorespiration can drain away 50% of carbon fixed by Calvin cycle on a hot, dry day strong selection pressure to evolve better way to fix carbon & minimize photorespiration 2006-2007 7