Photosynthesis: Variations on the Theme. AP Biology

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1 Photosynthesis: Variations on the Theme

2 Remember what plants need Photosynthesis u light reactions light H 2 O sun ground u Calvin cycle CO 2 air O C O What structures have plants evolved to supply these needs?

3 Alternative mechanisms of carbon fixation have evolved in hot, arid climates: Dehydration is a problem for plants, sometimes requiring trade-offs with photosynthesis. On hot, dry days, plants close stomata, which conserves H2O but also limits photosynthesis. The closing of stomata reduces access to CO2 and causes O2 to build up. These conditions favor an apparently wasteful process called photorespiration

4 Photorespiration: An Evolutionary Relic? In most plants (C3 plants), initial fixation of CO2, via rubisco, forms a three-carbon compound (3-phosphoglycerate, aka. PGA). In photorespiration, rubisco adds O2 instead of CO2 in the Calvin cycle, producing a two-carbon compound. Photorespiration consumes O2 and organic fuel and releases CO2 without producing ATP or sugar.

5 O 2 and organic fuel Photorespiration Calvin Cycle (CO 2 and NO ATP or Sugar)

6 Leaf Structure: cuticle epidermis palisades layer vascular bundle xylem (water) phloem (sugar) spongy layer O 2 H 2 O CO 2 Transpiration Gas exchange stomate O 2 H 2 O CO 2 guard cell

7 Controlling water loss (Transpiration): Hot or dry days u stomates close to conserve water u guard cells gain H 2 O = stomates open lose H 2 O = stomates close u adaptation to living on land, but creates PROBLEMS!

8 When stomates close Closed stomates lead to u O 2 build up from light reactions u CO 2 is depleted in Calvin cycle causes problems in Calvin Cycle O 2 xylem (water) CO phloem (sugars) O 2 CO 2 H 2 O

9 Inefficiency of RuBisCo: CO 2 vs O 2 RuBisCo in Calvin cycle u carbon fixation enzyme normally bonds C to RuBP CO 2 is the optimal substrate reduction of RuBP building sugars u when O 2 concentration is high RuBisCo bonds O 2 to RuBP O 2 is a competitive substrate oxidation of RuBP breakdown sugars photosynthesis photorespiration

10 Calvin cycle when CO 2 is abundant: 1C CO 2 G3P to make glucose ATP ADP RuBP 5C 5C RuBisCo 6C unstable intermediate G3P 3C C3 plants 3C PGA NADPH ATP NADP 3C ADP

11 Calvin cycle when O 2 is high: Hey Dude, are you high on oxygen! RuBP 5C O 2 RuBisCo 2C 3C to mitochondria lost as CO 2 without making ATP It s so sad to see a good enzyme, go BAD! photorespiration

12 Impact of Photorespiration: Oxidation of RuBP u short circuit of Calvin cycle u loss of carbons to CO 2 can lose 50% of carbons fixed by Calvin cycle u reduces production of photosynthesis no ATP (energy) produced no C 6 H 12 O 6 (food) produced u if photorespiration could be reduced, plants would become 50% more efficient strong selection pressure to evolve alternative carbon fixation systems

13 Reducing photorespiration: Separate carbon fixation from Calvin cycle u C4 plants PHYSICALLY separate carbon fixation from Calvin cycle w different cells to fix carbon vs. where Calvin cycle occurs w store carbon in 4C compounds different enzyme to capture CO 2 (fix carbon) w PEP carboxylase different leaf structure u CAM plants separate carbon fixation from Calvin cycle by TIME OF DAY fix carbon during night w store carbon in 4C compounds perform Calvin cycle during day

14 C4 plants: A better way to capture CO 2 u 1st step before Calvin cycle, fix carbon with enzyme PEP carboxylase store as 4C compound u adaptation to hot, dry climates have to close stomates a lot different leaf anatomy u sugar cane, corn, other grasses sugar cane corn

15 PEP (3C) + CO 2 oxaloacetate (4C) organic acid C4 leaf anatomy: light reactions O 2 PEP carboxylase CO 2 C3 anatomy stomate PEP carboxylase enzyme bundle sheath cell CO 2 RuBisCo u higher attraction for CO 2 than O 2 better than RuBisCo u fixes CO 2 in 4C compounds u regenerates CO 2 in inner cells for RuBisCo keeping O 2 away from RuBisCo C4 anatomy

16 Comparative anatomy: C3 Location, location,location! C4 PHYSICALLY separate C fixation from Calvin cycle

17 CAM (Crassulacean Acid Metabolism) plants: Adaptation to hot, dry climates u separate carbon fixation from Calvin cycle by TIME close stomates during day open stomates during night u at night: open stomates & fix carbon in 4C storage compounds u fix carbon with enzyme PEP carboxylase too! u in day: release CO 2 from 4C acids to Calvin cycle increases concentration of CO 2 in cells It s all in the timing! u succulents, some cacti, pineapple

18 PEP (3C) + CO 2 malic acid (4C) organic acid CAM plants: cacti succulents pineapple

19 C4 v. CAM Summary: Solves CO 2 / 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 = 2 different times night vs. day

20 Why the C3 problem? Possibly evolutionary baggage We ve all got baggage! u Rubisco evolved in high CO 2 atmosphere there wasn t strong selection against active site of Rubisco accepting both CO 2 & O 2 Today it makes a difference u 21% O 2 vs. 0.03% CO 2 u photorespiration can drain away 50% of carbon fixed by Calvin cycle on a hot, dry day u strong selection pressure to evolve better way to fix carbon & minimize photorespiration

21 It s not so easy as it looks Any Questions??

22 A second look inside a leaf Gas exchange & water flow u CO 2 in u O 2 out u H 2 O out for Calvin cycle waste from light reactions for light reactions photosynthesis O 2 CO 2 xylem (water) phloem (sugars) gas exchange water loss H 2 O O 2 CO 2

23 C4 photosynthesis: CO 2 PHYSICALLY separated C fixation from Calvin cycle O 2 O 2 CO 2 Outer cells u light reaction & carbon fixation u pumps CO 2 to inner cells u keeps O 2 away from inner cells away from RuBisCo Inner cells u Calvin cycle u glucose to veins

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