Remember what plants need! Photosynthesis. Photosynthesis: Variations on the Theme " Leaf Structure. Controlling water loss from leaves

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1 Remember what plants need! Photosynthesis O light reactions C O! light! sun! H2O! ground Photosynthesis: Variations on the Theme Calvin cycle!! air vascular bundle Leaf Structure phloem (transports What structures have plants evolved to supply these needs? Controlling water loss from leaves xylem (transports water) sugar) Guard cells experience opposite forces! With sun, lots of glucose made, osmosis inward (stomates full & turgid remain open) = Light reactions produce O2 as a waste produce Calvin cycle takes up gain H2O! With LOTS of sun, evaporation of water, osmosis outward (stomates shrink & become flaccid and thus close) = lose H2O O2 H2 O On very hot or dry days: Stomates close to conserve water! This is an adaptation to Transpiration stomate - loss of H2O Gas exchange AP Biology O2 H2O living on land, but guard cell it also creates PROBLEMS!

2 When stomates close! Closing stomata does conserve water BUT it lead to O 2 build up from light reactions CO 2 is depletion by Calvin cycle! causes problems in Calvin Cycle!! O 2 CO 2 H 2 O How So? Inefficiency of : CO 2 vs O 2 Affinity in Calvin cycle carbon fixation enzyme! normally bonds CO 2 to RuBP! CO 2 is the optimal substrate When the enzyme binds to CO 2 it fixes the carbon and starts the process of producing sugar = BUT when O 2 concentration is high! bonds O 2 to RuBP! O 2 is a competitive substrate When O 2 is added, the product splits into a two carbon compound that is changed into CO 2 by peroxisomes and mitochondria. Photosynthesis Photorespiration = occurs in the light and consumes O 2 while producing CO 2 G3P to make glucose Calvin cycle when CO 2 is abundant ATP ADP G3P RuBP 1C CO 2 C3 plants 6C unstable intermediate PGA Calvin cycle when O 2 is high Hey Dude, are you high on oxygen?! It s so sad to see a good enzyme, go BAD! RuBP O 2 photorespiration 2C Lost as CO 2 without producing Sugar or ATP (This O 2 also does not get used in cellular respiration which would produce ATP) NADPH ATP NADP ADP

3 Impact of Photorespiration Reducing photorespiration! Oxidation of RuBP! In C3 plants carbon fixation takes place as part of the normal Calvin Cycle and happen in mesophyll cells short circuit of Calvin cycle loss of carbons to! Adaptation for HEAT: Separate carbon fixation from Calvin cycle! can lose 50% of carbons fixed by Calvin cycle! PHYSICALLY separate carbon fixation from Calvin cycle reduces production of photosynthesis # different cells fix carbon vs. run the Calvin cycle # Initial cells store carbon in a 4C compounds as a first product! different enzyme to capture (fix carbon) # PEP carboxylase! different leaf structure! no ATP (energy) produced (O2 consumed by photorespiration and not used in cellular respiration)! no C6H12O6 (food) produced if photorespiration could be reduced, plant would become 50% more efficient CAM plants! separate carbon fixation from Calvin cycle by TIME OF DAY! fix carbon during night! strong selection pressure to evolve alternative carbon # store carbon in 4C compounds fixation systems! perform Calvin cycle during day! Both fixation and Calvin cycle do occur in same cell though PEP () + oxaloacetate (4C) C4 leaf anatomy light reactions! A better way to capture O2 Mesophyll Cell:! 1st step before Calvin cycle, PEP carboxylase fix carbon with enzyme PEP carboxylase # store as 4C compound oxaloacetate that is converted to malate! Only then is 4C compound exported to corn Bundle Sheath cells where is released and incorporated by the Calvin Cycle. adaptation to hot, dry climates bundle sheath cell higher attraction for than O2! better than which has a high affinity for O2. in high O2 environment of the Mesophyll cell where light reactions occur! different leaf anatomy sugar cane, corn, other grasses stomate! PEP carboxylase enzyme:! have to close stomates a lot! Rubisco NOT responsible for fixing C3 anatomy sugar cane fixes in 4C compounds regenerates in inner cells for! keeping O2 away from 1 ATP is used up to convert pyruvate back to PEP, the acceptor C4 anatomy

4 Comparative anatomy C3 Location, location,location! CAM (Crassulacean Acid Metabolism) plants! Adaptation to hot, dry climates C4 separate carbon fixation from Calvin cycle by TIME (temporal)! close stomates during day to converse water! open stomates during night when it is less hot At night: open stomates & fix carbon in 4C storage compounds = an organic acid During the day: close stomates & release from 4C acids to Calvin cycle It s all in the timing!! increases concentration of in mesophyll cells so Rubisco fixes more than O2 PHYSICALLY separate C fixation from Calvin cycle succulents, some cacti, pineapple C4 vs CAM Summary CAM plants ( crassus - Latin for thick) solves / O2 gas exchange vs. H2O loss challenge cacti succulents pineapple CAM plants separate 2 steps of C fixation anatomically in 2 different cells separate 2 steps of C fixation temporally = 2 different times night vs. day Mesophyll cell

5 Why the C3 problem?! Possibly evolutionary baggage 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 21% O 2 vs. 0.03% CO 2 We ve all got baggage! 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 resulted in evolution of C4 and CAM plants Review: C3, C4, CAM C3, C4, and CAM truly refer to the alternative method of carbon fixation -- grabbing carbon out of the air -- and not the Calvin Cycle itself. $ They *all* use the Calvin Cycle for sugar generation, but they differ in how they turn carbon from thin air into organic matter.! In C4, CO 2 is fixed into 4-carbon storage compounds like oxaloacetate & malate (hence C4)! In CAM, CO 2 is fixed into organic acids like malic acid & isocitric acid (hence Crassulacean Acid Metabolism)! In C3, while CO 2 is initially fixed into a 6-carbon molecule, it is unstable & quickly breaks down to 3-carbon glyceraldehyde-3-phosphate (G3P) (hence C3) C4 & CAM should be seen as variations on *carbon fixation*, because plants had to evolve alternative systems given the limitations of their enzymes and their need to conserve water. It s not so easy as it looks Any Questions??

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