CHAPTER 10

1. CHAPTER 10 PHOTOSYNTHESIS

2. PHOTOSYNTHESIS • WHAT HAPPENS DURING THIS PROCESS??? • PLANTS TAKE THE ENERGY FROM THE SUN AND CARBON DIOXIDE AND CONVERT IT INTO SUGAR AND OXYGEN

3. THE ENERGY CYCLE

4. TRACKING ATOMS THROUGH PHOTOSYN.

5. SITE OF PHOTOSYNTHESIS

6. OVERVIEW OF PHOTOSYNTHESIS

7. WAVELENGTH • LIGHT IS A FORM OF ELECTROMAGNETIC ENERGY THAT TRAVELS IN RHYTHMIC WAVES WHICH ARE DISTURBANCES OF ELECTRIC AND MAGNETIC FIELDS • WAVELENGTH- THS DISTANCE BETWEEN THE CRESTS OF ELECTROMAGNETIC WAVES • THE ELECTROMAGNETIC SPECTRUM RANGES FROM WAVELENGTHS THAT ARE LESS THAN A NANOMETER (GAMMA RAYS) TO MORE THAN A KILOMETER (RADIO WAVES) • VISIBLE LIGHT- IS DETECTABLE BY THE HUMAN EYE, IS ONLY A SMALL PORTION OF THE ELECTROMAGNETIC SPECTRUM AND RANGES FROM ABOUT 380 TO 750 nm. THE WAVELENGTHS MOST IMPORTANT FOR PHOTOSYNTHESIS ARE WITHIN THIS RANGE

8. THE LIGHT SPECTRUM

9. PROPERTIES OF LIGHT • LIGHT ALSO BEHAVES AS IF IT CONSISTS OF DISCRETE PARTICLES OR QUANTA CALLED PHOTONS (PACKETS OF ENERGY) • EACH PHOTON HAS A FIXED QUANTITY OF ENERGY WHICH IS INVERSELY PROPORTIONAL TO THE WAVELENGTH OF LIGHT • EX: A PHOTON OF VIOLET LIGHT HAS NEARLY TWICE AS MUCH ENERGY AS A PHOTON OF RED LIGHT • THE SUN RADIATES THE FULL SPECTRUM OF ELECTROMAGNETIC ENERGY • THE ATMOSPHERE ACTS AS A SELECTIVE WINDOW THAT ALLOWS VISIBLE LIGHT TO PASS THROUGH WHILE SCREENING OUT A SUBSTANTIAL FRACTION OF OTHER RADIATION • THE VISIBLE RANGE OF LIGHT IS THE RADIATION THAT DRIVES PHOTOSYNTHESIS • BLUE AND RED, THE 2 WAVELENGTHS MOST EFFECTIVELY ABSORBED BY CHLOROPHYLL, ARE THE COLORS MOST USEFUL AS ENERGY FOR THE LIGHT REACTIONS

10. PHOTOSYNTHETIC PIGMENTS: THE LIGHT RECEPTORS • LIGHT MAY BE REFLECTED, TRANSMITTED, OR ABSORBED WHEN IT CONTACTS MATTER • PIGMENTS = SUBSTANCES THAT ABSORB VISIBLE LIGHT • DIFFERENT PIGMENTS ABSORB DIFFERENT WAVELENGTHS OF LIGHT • WAVELENTHS THAT ARE ABSORBED DISAPPEAR, SO A PIGMENT THAT ABSORBS ALL WAVELENGTHS APPEARS BLACK • WHEN WHITE LIGHT, WHICH CONTAINS ALL THE WAVELENGTHS OF VISIBLE LIGHT, ILLUMINATES A PIGMENT, THE COLOR YOU SEE IS THE COLOR MOST REFLECTED OR TRANSMITTED BY THE PIGMENT • EXAMPLE: A LEAF APPEARS GREEN BECAUSE CHLOROPHYLL ABSORBS RED AND BLUE LIGHT BUT TRANSMITS AND REFLECTS GREEN LIGHT • EACH PIGMENT HAS A CHARACTERISTIC ABSORPTION SPECTRUM OR PATTERN OF WAVELENGTHS THAT IT ABSORBS. IT IS EXPRESSED AS A GRAPH OF ABSORPTION VERSUS WAVELENGTH

11. ABSORPTION SPECTRUM FOR CHLOROPHYLL

12. ABSORPTION SPECTRUM • THE ABSORPTION SPECTRUM FOR A PIGMENT IN SOLUTION CAN BE DETERMINED BY USING A SPECTROPHOTOMETER, AN INSTRUMENT USED TO MEASURE WHAT PROPORTION OF A SPECIFIC WAVELENGTH IS ABSORBED OR TRANSMITTED BY THE PIGMENT • SINCE CHLOROPHYLL a IS THE LIGHT-ABSORBING PIGMENT THAT PARTICIPATES DIRECTLY IN THE LIGHT REACTIONS, THE ABSORPTION SPECTRUM OF CHLOROPHYLL a PROVIDES CLUES AS TO WHICH WAVELENGTHS OF VISIBLE LIGHT ARE MOST EFFECTIFVE FOR PHOTOSYNTHESIS

13. THE ACTION SPECTRUM • THE ACTION SPECTRUM FOR PHOTOSYNTHESIS DOES NOT EXACTLY MATCH THE ABSORPTION SPECTRUM FOR CHLOROPHYLL a • SINCE CHLOROPHYLL a IS NOT THE ONLY PIGMENT IN CHLOROPLASTS THAT ABSORB LIGHT, THE ABSORPTION SPECTRUM FOR CHLOROPHYLL a UNDERESTIMATES THE EFFECTIVENESS OF SOME WAVELENGTHS • EVEN THOUGH ONLY SPECIAL CHLOROPHYLL a MOLECULES CAN PARTICIPATE DIRECTLY IN THE LIGHT REACTIONS, OTHER PIGMENTS, CALLED ACCESSORY PIGMENTS, CAN ABSORB LIGHT AND TRANSFER THE ENERGY TO CHLOROPHYLL a

14. PHOTOEXCITATION OF CHLOROPHYLL • COLORS OF ABSORBED WAVELENGTHS DISAPPEAR FROM THE SPECTRUM OF TRANSMITTED AND REFLECTED LIGHT • THE ABSORBED PHOTON BOOSTS ONE OF THE PIGMENT MOLECULE’S ELECTRONS IN ITS LOWEST-ENERGY STATE (GROUND STATE) TO AN ORBITAL OF HIGHER POTENTIAL ENERGY (EXCITED STATE)

15. THE EXCITED STATE IS UNSTABLE, SO EXCITED ELECTRONS QUICKLY FALL BACK TO THE GROUND STATE ORBITAL, RELEASING EXCESS ENERGY IN THE PROCESS • THIS RELEASED ENERGY MAY BE: • DISSIPATED AS HEAT • RERADIATED AS A PHOTON OF LOWER ENERGY AND LONGER WAVELENGTH THAN THE ORIGINAL LIGHT THAT EXCITED THE PIGMENT. THIS AFTERGLOW IS CALLED FLUORESCENCE

16. PIGMENT MOLECULES DO NOT FLOURESCE WHEN IN THE THYLAKOID MEMBRANES, BECAUSE NEARBY PRIMARY ELECTRON ACCEPTOR MOLECULES TRAP EXCITED STATE ELECTRONS THAT HAVE ABSORBED PHOTONS • IN THIS REDOX REACTION, CHLOROPHLL IS PHOTO-OXIDIZED BY THE ABSORPTION OF LIGHT ENERGY AND THE ELECTRON ACCEPTOR IS REDUCED • BECAUSE NO PRIMARY ELECTRON ACCEPTOR IS PRESENT, ISOLATED CHLOROPHYLL FLOURESCES IN THE RED PART OF THE SPECTRUM AND DISSIPATES HEAT

17. STRUCTURE OF CHLOROPHYLL

18. REFLECTED LIGHT IN CHLOROPLASTS

19. CHLOROPHYLL STRUCTURE

20. PHOTOSYSTEMS • ANTENNAE COMPLEX • CHOROPHYLL a, CHLOROPHYLL b, AND CAROTENOID MOLECULES ARE LIGHT-GATHERING ANTENNAE THAT ABSORB PHOTONS AND PASS THE ENERGY FROM MOLECULE TO MOLECULE. THIS PROCESS OF RESONANCE ENERGY TRANSFER IS CALLED INDUCTIVE RESONANCE • DIFFERENT PIGMENTS WITHIN THE ANTENNAL COMPLEX HAVE SLIGHTLY DIFFERENT ABSORPTION SPECTRA, SO COLLECTIVELY THEY CAN ABSORB PHOTONS FROM A WIDER RANGE OF THE LIGHT SPECTRUM THAN WOULD BE POSSIBLE WITH ONLY ONE TYPE OF PIGMENT MOLECULE

21. REACTION CENTER CHLOROPHYLL • ONLY ONE OF THE MANY CHLOROPHYLL a molecules in each complex can actually transfer an excited electron to initiate the light reactions. This specialized chlorophyll a is located in the reaction center • PRIMARY ELECTRON ACCEPTOR: Located near the reaction center, a primary electron acceptor molecule traps excited state electrons released from the reaction center chlorophyll • The transfer of excited state electrons from chlorophyll to primary electron acceptor molecules is the first step of the light reactions. The energy stored in the trapped electrons powers the synthesis of ATP and NADPH.

22. HOW A PHOTOSYSTEM HARVEST LIGHT

23. PHOTOSYSTEMS • TWO TYPES OF PHOTOSYSTEMS ARE LOCATED IN THE THYLAKOID MEMBRANES, PHOTOSYSTEM I AND PHOTOSYSTEM II • THE REACTION CENTER OF PHOTOSYSTEM I HAS A SPECIALIZED CHLOROPHYLL a MOLECULE KNOWN AS P700 (THE FAR RED PORTION OF THE SPECTRUM) • THE REACTION CENTER OF PHOTOSYSTEM II HAS A SPECIALIZED CHOROPHYLL a MOLECULE KNOWN AS P680, WHICH ABSORBS BEST AT A WAVELENGTH OF 680 nm • P700 AND P680 ARE IDENTICAL CHLOROPHYLL a MOLECULES, BUT EACH IS ASSOCIATED WITH A DIFFERENT PROTEIN. THIS AFFECTS THEIR ELECTRON DISTRIBUTION AND RESULTS IN SLIGHTLY DIFFERENT ABSORPTION SPECTRA.

24. NONCYCLIC ELECTRON FLOW DURING LIGHT REACTIONS

25. MECHANICAL ANALOG FOR LIGHT REACTIONS

26. CYCLIC ELECTRON FLOW

27. MODEL OF THYLAKOID MEMBRANE

28. LIGHT REACTIONS VIDEO

29. THE CALVIN CYCLE

30. PHOTORESPIRATION • PHOTORESPIRATION= A METABOLIC PATHWAY THAT CONSUMES OXYGEN INSTEAD OF CO2, • IT OCCURS BECAUSE THE ACTIVE SITE OF RUBISCO CAN ACCEPT O2 AS WELL AS CO2 • PRODUCES NO ATP MOLECULES • DECREASES PHOTOSYNTHETIC OUTPUT BY REDUCING ORGANIC MOLECULES USED IN THE CALVIN CYCLE • WHEN THE [02] IN THE LEAF’S AIR SPACES IS HIGHER THAN [CO2], RUBISCO ACCEPTS O2 AND TRANSFER IT TO RuBP. (THE “PHOTO” IN PHOTORESPIRATION REFERS TO THE FACT THAT THIS PATHWAY USUALLY OCCURS IN LIGHT WHEN PHOTOSYNTHESIS REDUCES CO2 AND RAISES O2 IN THE LEAF SPACES

31. RUBISCO TRANSFERS O2 TO RuBP RESULTING 5-C MOLECULE SPLITS INTO 2-C MOLECULE 3-C MOLECULE (GLYCOLATE) (PGA) LEAVES CHLOROPLAST & GOES TO PEROXISOME STAYS IN CALVIN CYCLE GLYCOLATE IS BROKEN DOWN INTO CO2 IN THE MITOCHONDRION

32. SOME SCIENTISTS BELIEVE THAT PHOTOREPSIRATION IS A METABOLIC RELIC FROM EARLIER TIMES WHEN THE ATMOSPEHRE CONTAINED LESS O2 AND MORE CO2 • SOME CROP PLANTS (E.G. SOYBEANS) LOSE AS MUCH AS 50% OF THE CARBON FIXED BY THE CALVIN CYCLE TO PHOTORESPIRATION • PHOTORERESPIRATION IS FOSTERED BY HOT, DRY, DAYS WHEN PLANTS CLOSE THEIR STOMATA TO PREVENT DEHYDRATION BY REDUCING WATER LOSS FROM THE LEAF • CERTAIN SPECIES OF PLANTS, WHICH LIVE IN HOT ARID CLIMATES, HAVE EVOLVED ALTERNATE MODES OF CARBON FIXATION THAT MINIMIZES PHOTORESPIRATION.

33. C4 PLANTS • MANY PLANTS SPECIES PREFACE THE CALVIN CYCLE WITH REACTIONS THAT INCORPORATE CARBON DIOXIDE INTO 4-CARBON COMPOUNDS • THESE PLANTS ARE CALLED C4 PLANTS • THE C4 PATHWAY IS USED BY SEVERAL THOUSAND SPECIES IN AT LEAST 19 FAMILIES INCLUDING CORN AND SUGARCANE • THIS PATHWAY IS ADPATIVE, BECAUSE IT ENHANCES CARBON FIXATION UNDER CONDITIONS THAT FAVOR PHOTORESPIRATION, SUCH AS HOT, ARID ENVIRONMENTS

34. LEAF ANATOMY OF C4 PLANTS • LEAF ANATOMY OF C4 PLANTS SPATIALLYL SEGREGATES THE CALVIN CYCLE FROM THE INITIAL INCORPORATION OF CO2 INTO ORGANIC COMPOUNDS. THERE ARE 2 DISTINCT TYPES OF PHOTOSYNTHETIC CELLS: • 1) BUNDLE-SHEATH CELLS-AROUND VEINS OF THE LEAF • 2) MESOPHYLL CELLS-MORE LOOSELY ARRANGED IN THE AREA BTW. THE BUNDLE SHEATH AND LEAF SURFACE

35. C4 ANATOMY AND PATHWAY MESOPHYLL CELLS PUMP CO2 INTO BUNDLE-SHEATH CELLS, MINIMIZING PHOTORESPIRATION AND ENHANCING SUGAR PRODUCTION BY MAINTAINING [CO2]

36. CAM PLANTS • A SECOND PHOTOSYNTHETIC ADAPTATION EXISTS IN SUCCULENT PLANTS ADAPTED TO VERY ARID CONDITIONS • THESE PLANTS OPEN THEIR STOMATA AT NIGHT AND CLOSE THEM DURING THE DAY • THIS CONSEVES WATER DURING THE DAY, BUT PREVENTS CO2 FROM ENTERING LEAVES

37. CAM PLANTS • WHEN STOMATA ARE OPEN AT NIGHT, CO2 IS TAKEN UP AND INCORPORATED INTO A VARIETY OF ORGANIC ACIDS. • THIS MODE OF CARBON FIXATION IS CALLED CRASSULACEAN ACID METABOLISM (CAM) • THE ORGANIC ACIDS MADE AT NIGHT ARE STORED IN VACUOLES OF MESOPHYLL CELLS UNTIL MORNING, WHEN STOMATA CLOSE • DURING DAYTIME, LIGHT REACTIONS SUPPLY ATP AND NADPH FOR THE CALVIN CYCLE. AT THIS TIME, CO2 IS RELEASED FROM THE ORGANIC ACIDS MADE THE PREVIOUS NIGHT AND IS INCORPORATED INTO SUGAR IN THE CHLOROPLASTS

38. THE CAM AND C4 PATHWAYS: • ARE SIMILAR IN THAT CO2 IS FIRST INCORPORATED INTO ORGANIC INTERMEDIATES BEFORE IT ENTERS THE CALVIN CYCLE • DIFFER IN THAT THE INITIAL STEPS OF CARBON FIXATION IN C4 PLANTS ARE STRUCTURALLY SEPARATE FROM THE CALVIN CYCLE; IN CAM PLANTS, THE 2 STEPS OCCUR AT SEPARATE TIMES • REGARDLESS OF WHETHER THE PLANT USES A C3, C4, OR CAM PATHWAY, ALL PLANTS USE THE CALVIN CYCLE TO PRODUCE SUGAR FROM CO2

39. CALVIN CYCLE VIDEO

40. PHOTOSYNTHESIS REVIEW