1. Photosynthesis Chapter 8

2. Photosynthesis Overview Energy for all life on Earth ultimately comes from photosynthesis 6CO 2 + 12H 2 O C 6 H 12 O 6 + 6H 2 O + 6O 2 Oxygenic photosynthesis is carried out by ◦ Cyanobacteria ◦ 7 groups of algae ◦ All land plants – chloroplasts 2

3. Chloroplast 3 Thylakoid membrane – internal membrane ◦ Contains chlorophyll and other photosynthetic pigments ◦ Pigments clustered into photosystems Grana – stacks of flattened sacs of thylakoid membrane Stroma lamella – connect grana Stroma – semiliquid surrounding thylakoid membranes

4. Stages 4 Light-dependent reactions ◦ Require light 1.Capture energy from sunlight 2.Make ATP and reduce NADP + to NADPH Carbon fixation reactions or light-independent reactions ◦ Does not require light 3.Use ATP and NADPH to synthesize organic molecules from CO 2 Animation

5. Discovery of Photosynthesis Jan Baptista van Helmont (1580–1644) ◦ Demonstrated that the substance of the plant was not produced only from the soil Joseph Priestly (1733–1804) ◦ Living vegetation adds something to the air Jan Ingen-Housz (1730–1799) ◦ Proposed plants carry out a process that uses sunlight to split carbon dioxide into carbon and oxygen (O 2 gas) 5

6. F.F. Blackman (1866–1947) ◦ Came to the startling conclusion that photosynthesis is in fact a multistage process, only one portion of which uses light directly ◦ Light versus dark reactions ◦ Enzymes involved 6

7. C. B. van Niel (1897–1985) ◦ Found purple sulfur bacteria do not release O 2 but accumulate sulfur ◦ Proposed general formula for photosynthesis  CO 2 + 2 H 2 A + light energy → (CH 2 O) + H 2 O + 2 A ◦ Later researchers found O 2 produced comes from water Robin Hill (1899–1991) ◦ Demonstrated Niel was right that light energy could be harvested and used in a reduction reaction 7

8. Pigments Molecules that absorb light energy in the visible range Light is a form of energy Photon – particle of light ◦ Acts as a discrete bundle of energy ◦ Energy content of a photon is inversely proportional to the wavelength of the light Photoelectric effect – removal of an electron from a molecule by light 8

9. Absorption spectrum When a photon strikes a molecule, its energy is either ◦ Lost as heat ◦ Absorbed by the electrons of the molecule  Boosts electrons into higher energy level Absorption spectrum – range and efficiency of photons molecule is capable of absorbing 9

10. Organisms have evolved a variety of different pigments Only two general types are used in green plant photosynthesis ◦ Chlorophylls ◦ Carotenoids In some organisms, other molecules also absorb light energy 10 Many pigments

11. Chlorophylls Chlorophyll a ◦ Main pigment in plants and cyanobacteria ◦ Only pigment that can act directly to convert light energy to chemical energy ◦ Absorbs violet-blue and red light Chlorophyll b ◦ Accessory pigment or secondary pigment absorbing light wavelengths that chlorophyll a does not absorb 11

12. 12 Structure of chlorophyll Porphyrin ring ◦ Complex ring structure with alternating double and single bonds ◦ Magnesium ion at the center of the ring Photons excite electrons in the ring Electrons are shuttled away from the ring

13. Relative effectiveness of different wavelengths of light in promoting photosynthesis Corresponds to the absorption spectrum for chlorophylls 13 Action spectrum

14. 14 Other Pigments Carotenoids ◦ Carbon rings linked to chains with alternating single and double bonds ◦ Can absorb photons with a wide range of energies ◦ Also scavenge free radicals – antioxidant  Protective role Phycobiloproteins ◦ Important in low-light ocean areas

15. Photosystem Organization Antenna complex ◦ Hundreds of accessory pigment molecules ◦ Gather photons and feed the captured light energy to the reaction center Reaction center ◦ 1 or more chlorophyll a molecules ◦ Passes excited electrons out of the photosystem 15

16. Antenna complex Also called light-harvesting complex Captures photons from sunlight and channels them to the reaction center chlorophylls In chloroplasts, light-harvesting complexes consist of a web of chlorophyll molecules linked together and held tightly in the thylakoid membrane by a matrix of proteins 16

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18. Reaction center Transmembrane protein– pigment complex When a chlorophyll in the reaction center absorbs a photon of light, an electron is excited to a higher energy level Light-energized electron can be transferred to the primary electron acceptor, reducing it Oxidized chlorophyll then fills its electron “hole” by oxidizing a donor molecule 18

19. Light-Dependent Reactions 1. Primary photoevent ◦ Photon of light is captured by a pigment molecule 2. Charge separation ◦ Energy is transferred to the reaction center; an excited electron is transferred to an acceptor molecule 3. Electron transport ◦ Electrons move through carriers to reduce NADP + 4. Chemiosmosis ◦ Produces ATP 19 Capture of light energy

20. Cyclic photophosphorylation In sulfur bacteria, only one photosystem is used Generates ATP via electron transport Anoxygenic photosynthesis Excited electron passed to electron transport chain Generates a proton gradient for ATP synthesis 20

21. Chloroplasts have two connected photosystems Oxygenic photosynthesis Photosystem I (P 700 ) ◦ Functions like sulfur bacteria Photosystem II (P 680 ) ◦ Can generate an oxidation potential high enough to oxidize water Working together, the two photosystems carry out a noncyclic transfer of electrons that is used to generate both ATP and NADPH 21

22. Photosystem I transfers electrons ultimately to NADP +, producing NADPH Electrons lost from photosystem I are replaced by electrons from photosystem II Photosystem II oxidizes water to replace the electrons transferred to photosystem I 2 photosystems connected by cytochrome/ b 6 - f complex 22 The connection

23. Noncyclic photophosphorylation Plants use photosystems II and I in series to produce both ATP and NADPH Path of electrons not a circle Photosystems replenished with electrons obtained by splitting water Z diagram 23

24. Photosystem II Resembles the reaction center of purple bacteria Core of 10 transmembrane protein subunits with electron transfer components and two P 680 chlorophyll molecules Reaction center differs from purple bacteria in that it also contains four manganese atoms ◦ Essential for the oxidation of water b 6 -f complex ◦ Proton pump embedded in thylakoid membrane 24

25. Photosystem I 25 Reaction center consists of a core transmembrane complex consisting of 12 to 14 protein subunits with two bound P 700 chlorophyll molecules Photosystem I accepts an electron from plastocyanin into the “hole” created by the exit of a light-energized electron Passes electrons to NADP + to form NADPH

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29. Chemiosmosis Electrochemical gradient can be used to synthesize ATP Chloroplast has ATP synthase enzymes in the thylakoid membrane ◦ Allows protons back into stroma Stroma also contains enzymes that catalyze the reactions of carbon fixation – the Calvin cycle reactions 29

30. Production of additional ATP Noncyclic photophosphorylation generates ◦ NADPH ◦ ATP Building organic molecules takes more energy than that alone Cyclic photophosphorylation used to produce additional ATP ◦ Short-circuit photosystem I to make a larger proton gradient to make more ATP 30

31. Carbon Fixation – Calvin Cycle To build carbohydrates cells use Energy ◦ ATP from light-dependent reactions ◦ Cyclic and noncyclic photophosphorylation ◦ Drives endergonic reaction Reduction potential ◦ NADPH from photosystem I ◦ Source of protons and energetic electrons 31

32. Calvin cycle Named after Melvin Calvin (1911–1997) Also called C 3 photosynthesis Key step is attachment of CO 2 to RuBP to form PGA carboxylase /oxygenase Uses enzyme ribulose bisphosphate carboxylase /oxygenase or rubisco 32

33. 3 phases 1. Carbon fixation ◦ RuBP + CO 2 → PGA 2. Reduction ◦ PGA is reduced to G3P 3. Regeneration of RuBP ◦ PGA is used to regenerate RuBP 3 turns incorporate enough carbon to produce a new G3P 6 turns incorporate enough carbon for 1 glucose 33

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35. Output of Calvin cycle Glucose is not a direct product of the Calvin cycle G3P is a 3 carbon sugar ◦ Used to form sucrose  Major transport sugar in plants  Disaccharide made of fructose and glucose ◦ Used to make starch  Insoluble glucose polymer  Stored for later use 35

36. Energy cycle Photosynthesis uses the products of respiration as starting substrates Respiration uses the products of photosynthesis as starting substrates Production of glucose from G3P even uses part of the ancient glycolytic pathway, run in reverse Principal proteins involved in electron transport and ATP production in plants are evolutionarily related to those in mitochondria 36

37. Photorespiration Rubisco has 2 enzymatic activities ◦ Carboxylation  Addition of CO 2 to RuBP  Favored under normal conditions ◦ Photorespiration  Oxidation of RuBP by the addition of O 2  Favored when stoma are closed in hot conditions  Creates low-CO 2 and high-O 2 CO 2 and O 2 compete for the active site on RuBP 37

38. Types of photosynthesis C 3 ◦ Plants that fix carbon using only C 3 photosynthesis (the Calvin cycle) C 4 and CAM ◦ Add CO 2 to PEP to form 4 carbon molecule ◦ Use PEP carboxylase ◦ Greater affinity for CO 2, no oxidase activity ◦ C 4 – spatial solution ◦ CAM – temporal solution 38

39. C 4 plants Corn, sugarcane, sorghum, and a number of other grasses Initially fix carbon using PEP carboxylase in mesophyll cells Produces oxaloacetate, converted to malate, transported to bundle-sheath cells Within the bundle-sheath cells, malate is decarboxylated to produce pyruvate and CO 2 Carbon fixation then by rubisco and the Calvin cycle 39

40. C 4 pathway, although it overcomes the problems of photorespiration, does have a cost To produce a single glucose requires 12 additional ATP compared with the Calvin cycle alone C 4 photosynthesis is advantageous in hot dry climates where photorespiration would remove more than half of the carbon fixed by the usual C 3 pathway alone 40 The cost of doing business this way

41. CAM plants Many succulent (water-storing) plants, such as cacti, pineapples, and some members of about two dozen other plant groups Stomata open during the night and close during the day ◦ Reverse of that in most plants Fix CO 2 using PEP carboxylase during the night and store in vacuole 41

42. When stomata closed during the day, organic acids are decarboxylated to yield high levels of CO 2 High levels of CO 2 drive the Calvin cycle and minimize photorespiration 42 CAM plants in action

43. Compare C 4 and CAM Both use both C 3 and C 4 pathways C 4 – two pathways occur in different cells CAM – C 4 pathway at night and the C 3 pathway during the day 43