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Photosynthesis: Life   from   Light  and   Air 2007-2008
Energy needs of life ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],consumers producers
How are they connected? Heterotrophs Autotrophs making energy & organic molecules from light energy making energy & organic molecules from ingesting organic molecules Where ’ s  the ATP? oxidation = exergonic reduction = endergonic glucose  +  oxygen    carbon  +  water  +  energy dioxide C 6 H 12 O 6 6O 2 6CO 2 6H 2 O ATP  + + + +  water  +  energy    glucose  +  oxygen carbon dioxide 6CO 2 6H 2 O C 6 H 12 O 6 6O 2 light energy  + + +
What does it mean to be a  plant ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],ATP N P K … H 2 O glucose CO 2
Plant structure  ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object]
stomate ,[object Object],[object Object]
Chloroplasts chloroplasts in plant cell cross section of leaf leaves chloroplast absorb sunlight & CO 2 make energy & sugar chloroplasts contain chlorophyll CO 2
[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],Plant structure  thylakoid chloroplast ATP H + H + H + H + H + H + H + H + H + H + H + outer membrane inner membrane thylakoid granum stroma
Photosynthesis ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],It ’ s  not  the Dark Reactions !
[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],Light reactions ATP thylakoid chloroplast H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H +
[object Object],[object Object],ETC of Respiration generates H 2 O
ETC of Photosynthesis ,[object Object],[object Object],generates O 2
The ATP that “Jack” built ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],sunlight breakdown of C 6 H 12 O 6 respiration photosynthesis ATP H + ADP + P i H + H + H + H + H + H + H + H +
Pigments of photosynthesis  ,[object Object],[object Object],[object Object],[object Object],[object Object],How does this molecular structure fit its function?
A Look at Light ,[object Object],R O Y G B I V
Light: absorption spectra ,[object Object],[object Object],[object Object],[object Object],[object Object],Why are plants green?
Photosystems of photosynthesis  ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],reaction center antenna pigments
ETC of Photosynthesis chlorophyll  a chlorophyll  b Photosystem II Photosystem I
1 ETC of Photosynthesis sun Photosystem II P680 chlorophyll a e e
1 2 ETC of Photosynthesis Photosystem II P680  chlorophyll a Inhale, baby !   e e ATP thylakoid chloroplast O O Plants SPLIT water! fill the e –  vacancy O H H H H e e e - e - H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + + H
1 2 3 4 ATP energy to build carbohydrates ATP thylakoid chloroplast Photosystem II P680  chlorophyll a ETC of Photosynthesis ATP H + H + H + ADP + P i H + H + H + H + H + H + H + H + e e e e to Calvin Cycle H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H +
sun 5 Photosystem II P680  chlorophyll a Photosystem I P700  chlorophyll b e e ETC of Photosynthesis fill the e –  vacancy e e e e e e
6 electron carrier 5 sun Photosystem II P680  chlorophyll a Photosystem I P700  chlorophyll b $$ in the bank… reducing power ! ETC of Photosynthesis e e e e NADPH  to Calvin Cycle
split H 2 O ETC of Photosynthesis O ATP to Calvin Cycle sun sun H + H + H + H + H + H + H + H + H + H + H + e e e e
ETC of Photosynthesis ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object]
Experimental evidence ,[object Object],[object Object],Proved O 2  came from H 2 O  not  CO 2  = plants split H 2 O ! 6CO 2 6H 2 O C 6 H 12 O 6 6O 2 light energy  + + + 6CO 2 6H 2 O C 6 H 12 O 6 6 O 2 light energy  + + + Experiment 1 6C O 2 6H 2 O C 6 H 12 O 6 6O 2 light energy  + + + Experiment 2
Noncyclic Photophosphorylation ,[object Object],[object Object],[object Object],ATP
Cyclic photophosphorylation ,[object Object],[object Object],[object Object], 18  ATP  + 12  NADPH 1  C 6 H 12 O 6  ATP
Photophosphorylation  NONcyclic photophosphorylation cyclic photophosphorylation ATP NADP
Photosynthesis summary  ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],… stay tuned for the Calvin cycle
Stomates
Photosynthesis:   The Calvin Cycle   Life   from   Air 2007-2008
The Calvin Cycle Whoops! Wrong Calvin… 1950s  |  1961
Remember what it means to be a  plant… ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],+  water  +  energy    glucose  +  oxygen carbon dioxide 6CO 2 6H 2 O C 6 H 12 O 6 6O 2 light energy  + + +
Light reactions ,[object Object],[object Object],[object Object],[object Object],   energy    reducing power       build stuff  !! photo synthesis   ATP
How is that helpful? ,[object Object],[object Object],[object Object],CO 2 NADPH NADP reduces CO 2 carbon fixation C 6 H 12 O 6 NADP
From CO 2     C 6 H 12 O 6 ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object]
From Light reactions to Calvin cycle ,[object Object],[object Object],[object Object],[object Object],[object Object],ATP stroma thylakoid
starch, sucrose, cellulose & more Calvin cycle RuBP RuBisCo 1. Carbon fixation 2. Reduction 3. Regeneration of RuBP ribulose bisphosphate ribulose  bisphosphate  carboxylase used to make glucose   G3P glyceraldehyde-3-P PGA phosphoglycerate C C C 1C CO 2 5C 3C 6 NADP 6 NADPH 6 ADP 6 ATP 3 ADP 3 ATP 3C 3C C C C C C C C C C C C C C C C 6C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C = = | H | H | H | H | H | H C C C – – 5C
glycolysis glucose C-C-C-C-C-C fructose-1,6bP P- C-C-C-C-C-C -P DHAP P- C-C-C G3P C-C-C -P pyruvate C-C-C Remember  G3P? glyceraldehyde 3-phosphate Photosynthesis ATP 2 ADP 2 ATP 4 ADP 4 NAD + 2 2
To G3P and Beyond ! ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],To G3P and beyond !
RuBisCo  ,[object Object],[object Object],[object Object],[object Object],[object Object],I’m green  with envy !   It’s not easy  being green !
Accounting  ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object]
Photosynthesis summary ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],NADP ADP
Light Reactions O 2 H 2 O Energy Building Reactions ATP ,[object Object],[object Object],[object Object],NADPH sunlight H 2 O ATP O 2 light energy  + + + NADPH
Calvin Cycle sugars CO 2 Sugar Building Reactions ADP ,[object Object],[object Object],[object Object],[object Object],ATP NADPH NADP CO 2 C 6 H 12 O 6  + + + NADP ATP + NADPH ADP
Putting it all together Sugar Building Reactions Energy  Building Reactions ,[object Object],[object Object],[object Object],[object Object],O 2 H 2 O sugars CO 2 ADP ATP NADPH NADP CO 2 H 2 O C 6 H 12 O 6 O 2 light energy  + + + sunlight
Energy cycle Photosynthesis Cellular Respiration O 2 CO 2 plants animals ,  plants ATP The Great Circle of Life,Mufasa ! even though this equation is a bit of a lie … it makes a  better story H 2 O sun glucose CO 2 H 2 O C 6 H 12 O 6 O 2 light energy  + + + CO 2 H 2 O C 6 H 12 O 6 O 2 ATP energy  + + +
Summary of photosynthesis ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],6CO 2 6H 2 O C 6 H 12 O 6 6O 2 light energy  + + +
Supporting a biosphere ,[object Object],[object Object],[object Object],[object Object],[object Object]
The poetic perspective… ,[object Object],[object Object],Then all the plants, cats,   dogs, elephants & people … are really particles of air woven together by strands of sunlight! sun air
Photosynthesis: Variations on the Theme 2007-2008
Remember what  plants  need… ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],What structures have plants evolved to supply these needs?    sun    ground    air O O C
Leaf Structure H 2 O CO 2 O 2 H 2 O O 2 CO 2 Transpiration vascular bundle Gas exchange phloem (sugar) xylem (water) stomate guard cell palisades layer spongy layer cuticle epidermis
Controlling water loss from leaves ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object]
When stomates close…  ,[object Object],[object Object],[object Object],[object Object],xylem (water) phloem (sugars) The best laid schemes of mice and men… and plants !     H 2 O O 2 CO 2 CO 2 O 2
Inefficiency of RuBisCo: CO 2  vs O 2 ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],photo synthesis photo respiration
Calvin cycle when CO 2  is abundant G3P to make  glucose RuBisCo C3 plants 6C unstable intermediate 1C CO 2 5C RuBP 3C PGA ADP ATP 3C NADP NADPH ADP ATP 3C G3P 5C
Calvin cycle when O 2  is high  to  mitochondria ––––––– lost as CO 2  without   making ATP photorespiration Hey Dude, are you high on oxygen ! RuBisCo It’s so   sad to see a good enzyme, go  BAD! 5C RuBP 3C 2C O 2
Impact of Photorespiration  ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object]
Reducing photorespiration  ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object]
C4 plants  ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],sugar cane corn
C4 leaf anatomy PEP (3C) + CO 2     oxaloacetate (4C) CO 2 light reactions C4 anatomy C3 anatomy ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],bundle sheath cell RuBisCo PEP carboxylase stomate CO 2 O 2
Comparative anatomy C3 C4 Location, location,location !   PHYSICALLY separate C fixation from Calvin cycle
CAM ( Crassulacean Acid Metabolism ) plants ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],It’s all in the timing !
CAM plants succulents cacti pine apple
C4 vs CAM Summary 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 solves CO 2  / O 2  gas exchange   vs. H 2 O loss   challenge
Why the C3 problem? ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],We’ve all got baggage !
2007-2008 It’s not so easy as it looks… Any Questions??
Ghosts of Lectures Past (storage) 2007-2008
A second look inside a leaf… ,[object Object],[object Object],[object Object],[object Object],photosynthesis gas exchange water loss xylem (water) for Calvin cycle waste from light reactions for light reactions phloem (sugars) O 2 CO 2 H 2 O O 2 CO 2
C4 photosynthesis ,[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],PHYSICALLY separated C fixation from Calvin cycle CO 2 O 2 CO 2 O 2

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Chapter 10(3)

  • 1. Photosynthesis: Life from Light and Air 2007-2008
  • 2.
  • 3. How are they connected? Heterotrophs Autotrophs making energy & organic molecules from light energy making energy & organic molecules from ingesting organic molecules Where ’ s the ATP? oxidation = exergonic reduction = endergonic glucose + oxygen  carbon + water + energy dioxide C 6 H 12 O 6 6O 2 6CO 2 6H 2 O ATP  + + + + water + energy  glucose + oxygen carbon dioxide 6CO 2 6H 2 O C 6 H 12 O 6 6O 2 light energy  + + +
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  • 7. Chloroplasts chloroplasts in plant cell cross section of leaf leaves chloroplast absorb sunlight & CO 2 make energy & sugar chloroplasts contain chlorophyll CO 2
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  • 18. ETC of Photosynthesis chlorophyll a chlorophyll b Photosystem II Photosystem I
  • 19. 1 ETC of Photosynthesis sun Photosystem II P680 chlorophyll a e e
  • 20. 1 2 ETC of Photosynthesis Photosystem II P680 chlorophyll a Inhale, baby ! e e ATP thylakoid chloroplast O O Plants SPLIT water! fill the e – vacancy O H H H H e e e - e - H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + + H
  • 21. 1 2 3 4 ATP energy to build carbohydrates ATP thylakoid chloroplast Photosystem II P680 chlorophyll a ETC of Photosynthesis ATP H + H + H + ADP + P i H + H + H + H + H + H + H + H + e e e e to Calvin Cycle H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H + H +
  • 22. sun 5 Photosystem II P680 chlorophyll a Photosystem I P700 chlorophyll b e e ETC of Photosynthesis fill the e – vacancy e e e e e e
  • 23. 6 electron carrier 5 sun Photosystem II P680 chlorophyll a Photosystem I P700 chlorophyll b $$ in the bank… reducing power ! ETC of Photosynthesis e e e e NADPH to Calvin Cycle
  • 24. split H 2 O ETC of Photosynthesis O ATP to Calvin Cycle sun sun H + H + H + H + H + H + H + H + H + H + H + e e e e
  • 25.
  • 26.
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  • 28.
  • 29. Photophosphorylation NONcyclic photophosphorylation cyclic photophosphorylation ATP NADP
  • 30.
  • 32. Photosynthesis: The Calvin Cycle Life from Air 2007-2008
  • 33. The Calvin Cycle Whoops! Wrong Calvin… 1950s | 1961
  • 34.
  • 35.
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  • 39. starch, sucrose, cellulose & more Calvin cycle RuBP RuBisCo 1. Carbon fixation 2. Reduction 3. Regeneration of RuBP ribulose bisphosphate ribulose bisphosphate carboxylase used to make glucose G3P glyceraldehyde-3-P PGA phosphoglycerate C C C 1C CO 2 5C 3C 6 NADP 6 NADPH 6 ADP 6 ATP 3 ADP 3 ATP 3C 3C C C C C C C C C C C C C C C C 6C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C = = | H | H | H | H | H | H C C C – – 5C
  • 40. glycolysis glucose C-C-C-C-C-C fructose-1,6bP P- C-C-C-C-C-C -P DHAP P- C-C-C G3P C-C-C -P pyruvate C-C-C Remember G3P? glyceraldehyde 3-phosphate Photosynthesis ATP 2 ADP 2 ATP 4 ADP 4 NAD + 2 2
  • 41.
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  • 48. Energy cycle Photosynthesis Cellular Respiration O 2 CO 2 plants animals , plants ATP The Great Circle of Life,Mufasa ! even though this equation is a bit of a lie … it makes a better story H 2 O sun glucose CO 2 H 2 O C 6 H 12 O 6 O 2 light energy  + + + CO 2 H 2 O C 6 H 12 O 6 O 2 ATP energy  + + +
  • 49.
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  • 52. Photosynthesis: Variations on the Theme 2007-2008
  • 53.
  • 54. Leaf Structure H 2 O CO 2 O 2 H 2 O O 2 CO 2 Transpiration vascular bundle Gas exchange phloem (sugar) xylem (water) stomate guard cell palisades layer spongy layer cuticle epidermis
  • 55.
  • 56.
  • 57.
  • 58. Calvin cycle when CO 2 is abundant G3P to make glucose RuBisCo C3 plants 6C unstable intermediate 1C CO 2 5C RuBP 3C PGA ADP ATP 3C NADP NADPH ADP ATP 3C G3P 5C
  • 59. Calvin cycle when O 2 is high to mitochondria ––––––– lost as CO 2 without making ATP photorespiration Hey Dude, are you high on oxygen ! RuBisCo It’s so sad to see a good enzyme, go BAD! 5C RuBP 3C 2C O 2
  • 60.
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  • 64. Comparative anatomy C3 C4 Location, location,location ! PHYSICALLY separate C fixation from Calvin cycle
  • 65.
  • 66. CAM plants succulents cacti pine apple
  • 67. C4 vs CAM Summary 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 solves CO 2 / O 2 gas exchange vs. H 2 O loss challenge
  • 68.
  • 69. 2007-2008 It’s not so easy as it looks… Any Questions??
  • 70. Ghosts of Lectures Past (storage) 2007-2008
  • 71.
  • 72.

Notas del editor

  1. So, in effect, photosynthesis is respiration run backwards powered by light. Cellular Respiration oxidize C 6 H 12 O 6  CO 2 & produce H 2 O fall of electrons downhill to O 2 exergonic Photosynthesis reduce CO 2  C 6 H 12 O 6 & produce O 2 boost electrons uphill by splitting H 2 O endergonic
  2. A typical mesophyll cell has 30-40 chloroplasts, each about 2-4 microns by 4-7 microns long. Each chloroplast has two membranes around a central aqueous space, the stroma. In the stroma are membranous sacs, the thylakoids. These have an internal aqueous space, the thylakoid lumen or thylakoid space. Thylakoids may be stacked into columns called grana.
  3. Not accidental that these 2 systems are similar, because both derived from the same primitive ancestor.
  4. NADH is an input ATP is an output O 2 combines with H + ’s to form water Electron is being handed off from one electron carrier to another -- proteins, cytochrome proteins & quinone lipids So what if it runs in reverse??
  5. Two places where light comes in. Remember photosynthesis is endergonic -- the electron transport chain is driven by light energy. Need to look at that in more detail on next slide
  6. Photons are absorbed by clusters of pigment molecules ( antenna molecules) in the thylakoid membrane. When any antenna molecule absorbs a photon, it is transmitted from molecule to molecule until it reaches a particular chlorophyll a molecule = the reaction center . At the reaction center is a primary electron acceptor which removes an excited electron from the reaction center chlorophyll a. This starts the light reactions. Don’t compete with each other, work synergistically using different wavelengths.
  7. Two places where light comes in. Remember photosynthesis is endergonic -- the electron transport chain is driven by light energy. Need to look at that in more detail on next slide
  8. PS II absorbs light Excited electron passes from chlorophyll to the primary electron acceptor Need to replace electron in chlorophyll An enzyme extracts electrons from H 2 O & supplies them to the chlorophyll This reaction splits H 2 O into 2 H + & O - which combines with another O - to form O 2 O 2 released to atmosphere Chlorophyll absorbs light energy (photon) and this moves an electron to a higher energy state Electron is handed off down chain from electron acceptor to electron acceptor In process has collected H + ions from H 2 O & also pumped by Plastoquinone within thylakoid sac. Flow back through ATP synthase to generate ATP.
  9. Need a 2nd photon -- shot of light energy to excite electron back up to high energy state. 2nd ETC drives reduction of NADP to NADPH. Light comes in at 2 points. Produce ATP & NADPH
  10. Need a 2nd photon -- shot of light energy to excite electron back up to high energy state. 2nd ETC drives reduction of NADP to NADPH. Light comes in at 2 points. Produce ATP & NADPH
  11. Two places where light comes in. Remember photosynthesis is endergonic -- the electron transport chain is driven by light energy. Need to look at that in more detail on next slide
  12. 1 photosystem is not enough. Have to lift electron in 2 stages to a higher energy level. Does work as it falls. First, produce ATP -- but producing ATP is not enough. Second, need to produce organic molecules for other uses & also need to produce a stable storage molecule for a rainy day (sugars). This is done in Calvin Cycle!
  13. Where did the energy come from? The Sun Where did the electrons come from? Chlorophyll Where did the H 2 O come from? The ground through the roots/xylem Where did the O 2 come from? The splitting of water Where did the O 2 go? Out stomates to air Where did the H + come from? The slitting of water Where did the ATP come from? Photosystem 2: Chemiosmosis (H+ gradient) What will the ATP be used for? The work of plant life! Building sugars Where did the NADPH come from? Reduction of NADP (Photosystem 1) What will the NADPH be used for? Calvin cycle / Carbon fixation
  14. CO 2 contains little energy because it is fully oxidized Reduce CO 2 in a series of steps to synthesize a stable energy storage molecule
  15. 1. A five-carbon sugar molecule called ribulose bisphosphate, or RuBP, is the acceptor that binds CO2 dissolved in the stroma. This process, called CO2 fixation, is catalyzed by the enzyme RuBP carboxylase, forming an unstable six-carbon molecule. This molecule quickly breaks down to give two molecules of the three-carbon 3-phosphoglycerate (3PG), also called phosphoglyceric acid (PGA). 2. The two 3PG molecules are converted into glyceraldehyde 3-phosphate (G3P, a.k.a. phosphoglyceraldehyde, PGAL) molecules, a three-carbon sugar phosphate, by adding a high-energy phosphate group from ATP, then breaking the phosphate bond and adding hydrogen from NADH + H+. 3. Three turns of the cycle, using three molecules of CO2, produces six molecules of G3P. However, only one of the six molecules exits the cycle as an output, while the remaining five enter a complex process that regenerates more RuBP to continue the cycle. Two molecules of G3P, produced by a total of six turns of the cycle, combine to form one molecule of glucose.
  16. Gluconeogenesis == making new glucose
  17. Where did the CO 2 come from? air Where did the CO 2 go? carbohydrate (G3P) Where did the H 2 O come from? soil (xylem) Where did the H 2 O go? split to donate e- & H+ Where did the energy come from? sun What’s the energy used for? to make ATP in light reactions What will the C 6 H 12 O 6 be used for? the work of plant life Where did the O 2 come from? splitting of H 2 O Where will the O 2 go? air or respiration What else is involved…not listed in this equation? ATP, ADP, NADP, NADPH
  18. On sunny hot days, balancing 2 processes/ 2 forces: Lots of sun so guard cells producing glucose, therefore increasing sugar concentration, therefore increase osmosis into guard cells = turgid & open Lots of sun so plant starts to wilt, therefore guard cells shrink = flaccid & closed
  19. The key point is how carbon dioxide is grabbed out of the air -- carbon fixation -- and then handed off to the Calvin cycle. C4 plants separate the 2 steps of carbon fixation anatomically. They use 2 different cells to complete the process. CAM plants separate the 2 steps of carbon fixation temporally. They do them at 2 different times. The key problem they are trying to overcome is that Rubisco is a very inefficient enzyme in the presence of high O2. In high O2, Rubisco bonds oxygen to RuBP rather than carbon, so the plants have to keep O2 away from Rubsico. 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.
  20. PEP = phosphoenolpyruvate C4 plants solve the photorespiration problem by fixing carbon in outer ring of mesophyll cells using a different enzyme -- PEP carboxylase -- which has a higher affinity for CO2 than O2 (better than Rubisco), so it fixes CO2 in 4C "storage" compounds (oxaloacetate, malate). It then passes the carbon on by regenerating CO2 in the inner bundle sheath cells for Rubisco to use in the Calvin cycle.
  21. Crassulacean Acid Metabolism ( CAM ) Succulents are in the family Crassulaceae the name Crassula is derived from the Latin "crassus" meaning thick and refers to the leaves of these succulent plants CAM plants solve the photorespiration problem by fixing carbon at night (when stomates are open), and put it in "storage" compounds (organic acids like malic acid, isocitric acid) and then in the day (when stomates are closed), they release the CO2 from the "storage" compounds to the Calvin cycle (thereby increasing CO2 in the cells, improving Rubisco's efficiency)
  22. 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 solid stuff. In C4, CO2 is fixed into 4-carbon "storage" compounds like oxaloacetate & malate (hence C4) In CAM, CO2 is fixed into organic acids like malic acid & isocitric acid (hence Crassulacean Acid Metabolism) In C3, while CO2 is initially fixed into a 6-carbon molecule, it is unstable & quickly breaks down to 3-carbon phosphoglycerate (PGA) (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.