To be successful today 1 Pull out Photosynthesis

To be successful today… 1) Pull out Photosynthesis Lab 2) Turn in Photosynthesis Part II Study Guide 3) TEST: Photosynthesis (parts I & II) TOM!! BYOD Color of the day is Phones are in put away in backpacks

Photosynthesis Part II: The Calvin Cycle, Environmental Conditions, & Preventing Photorespiration

Photosynthesis: An Overview • The net overall equation for photosynthesis is: 6 CO 2 + 6 H 2 O light C 6 H 12 O 6 + 6 O 2 • Photosynthesis occurs in 2 “stages”: 1. The Light Reactions (or Light-Dependent Reactions) 2. The Calvin Cycle (or Calvin-Benson Cycle or Dark Reactions or Light-Independent Reactions) 3

Photosynthesis: An Overview • To follow the energy in photosynthesis, Light Reactions light thylakoids light ATP NADPH Calvin Cycle stroma Organic compounds (carbs) 4

Phase 2: The Calvin Cycle • In the Calvin Cycle, chemical energy (from the light reactions) and CO 2 (from the atmosphere) are used to produce organic compounds (like glucose). • The Calvin Cycle occurs in the stroma of chloroplasts. 5

Three Steps of the Calvin Cycle 6

Phase 2: The Calvin Cycle • The Calvin Cycle first involves the process of carbon fixation. • This is the process of assimilating carbon from a non-organic compound (ie. CO 2) and incorporating it into an organic compound (ie. carbohydrates). CARBON FIXATION 7

Phase 2: The Calvin Cycle Step 1: Carbon Fixation • 3 molecules of CO 2 (from the atmosphere) are joined to 3 molecules of Ru. BP (a 5 -carbon sugar) by Rubisco (an enzyme also known as Ru. BP carboxylase) This forms 3 C C Rubisco C 3 carbon dioxide molecules C C C C molecules which each have 6 carbons (for a total of 18 carbons!) 3 Ru. BP molecules 8

Three Steps of the Calvin Cycle 9

Phase 2: The Calvin Cycle Where did the NADPH and ATP come from to do this? Step 2: Reduction • The three 6 -carbon molecules (very unstable) split in half, forming six 3 -carbon molecules. • These molecules are then reduced by gaining electrons from NADPH. • ATP is required for this molecular rearranging C C C C C C C C C ATP ADP P NADPH NADP+ 10

Phase 2: The Calvin Cycle Where did these 3 extra carbons come from? • There are now six 3 -carbon molecules, which are known as G 3 P or PGAL. • Since the Calvin Cycle started with 15 carbons (three 5 -carbon molecules) and there are now 18 carbons, we have a net gain of 3 carbons. • One of these “extra” 3 carbon G 3 P/PGAL molecules will exit the cycle and be used to form ½ a glucose molecule. C C C C C 11

Phase 2: The Calvin Cycle • Once the Calvin Cycle “turns”twice, those 2 molecules of G 3 P (a 3 -carbon carbohydrate) will combine to form 1 molecule of glucose (a 6 carbon carbohydrate molecule) OR another organic compound. C C C G 3 P (from 1 turn of the Calvin Cycle) C C C glucose 12 C

Phase 2: The Calvin Cycle • We say that the Calvin Cycle turns twice to make one molecule of glucose (6 -carbons), but really it turns 6 times; each entering carbon dioxide molecule represents one “turn” of the cycle, and 6 carbon dioxide molecules must be incorporated into organic compounds in order for one 6 -carbon glucose molecule to be produced. 13

Calvin Cycle 14

Three Steps of the Calvin Cycle 15

Phase 2: The Calvin Cycle Where does the ATP Step 3: Regeneration of Ru. BP come from to do this? • Since this is the Calvin Cycle, we must end up back at the beginning. • The remaining 5 G 3 P molecules (3 -carbons each!) get rearranged (using ATP) to form 3 Ru. BP molecules (5 -carbons each). C C C C 5 G 3 P molecules Total: 15 carbons C C ATP ADP P 3 Ru. BP molecules Total: 15 carbons 16

Phase 2: The Calvin Cycle CO 2 o sc bi Ru Ru. BP NADPH ATP NADP+ ADP P ORGANIC COMPOUND

Talk to a Partner: The Calvin Cycle 18

Talk to a Partner: The Calvin Cycle • Glucose is what we usually think of as being the major product of photosynthesis; however, G 3 P (also known as PGAL) is the real product, and though it is often used to make glucose it can also be used as a carbon skeleton to form other organic molecules. 19

Phase 2: The Calvin Cycle Quick recap: • In the Calvin Cycle, energy and electrons from the Light Reactions (in the form of ATP and NADPH) and carbon dioxide from the atmosphere are used to produce organic compounds. • The Calvin Cycle occurs in the stroma inside the chloroplasts (inside the cells…). • Carbon dioxide, ATP, and NADPH are required (reactants). • Organic compounds (G 3 P) are produced (products). 20

Photosynthesis: A Recap • So, as a broad overview of photosynthesis, • The Light Reactions (Phase 1) capture the energy in sunlight and convert it to chemical energy in the form of ATP and NADPH through the use of photosystems, electron transport chains, and chemiosmosis. • The Calvin Cycle (Phase 2) uses the energy transformed by the light reactions along with carbon dioxide to produce organic compounds. 21

Photosynthesis: A Recap • Photosynthesis Animation 22

Talk to a Partner 23

Photosynthesis: A Recap The photosynthetic equation: Provides the carbon to produce organic compounds during the Calvin Cycle Based on this equation, how could the rate of photosynthesis be measured? The organic compound ultimately produced during the Calvin Cycle light 6 H 2 O Split during the light reactions to replace electrons lost from Photosystem II 6 CO 2 6 O 2 Excites electrons during the light reactions C 6 H 12 O 6 Produced as a byproduct of the splitting of water during the light reactions 24

Environmental Factors & Photosynthesis • The rate (or speed) of photosynthesis can vary, based on environmental conditions. • Light intensity • Temperature • Oxygen concentration 25

Environmental Factors & Photosynthesis • Light intensity • As light intensity increases, so too does the rate of photosynthesis. • This occurs due to increased excitation of electrons in the photosystems. light saturation point • However, the photosystems will eventually become saturated. • Above this limiting level, no further increase in photosynthetic rate will occur. 26

Environmental Factors & Photosynthesis • Temperature • The effect of temperature on the rate of photosynthesis is linked to the action of enzymes. • As the temperature increases up to a certain point, the rate of photosynthesis increases. • Molecules are moving faster & colliding with enzymes more frequently, facilitating chemical reactions. • However, at temperatures higher or lower than this point, the rate of photosynthesis decreases. • Enzymes are denatured. 27

Environmental Factors & Photosynthesis • Oxygen concentration • As the concentration of oxygen increases, the rate of photosynthesis decreases. • This occurs due to the phenomenon of photorespiration 28

Photorespiration • Photorespiration occurs when Rubisco (Ru. BP carboxylase) joins oxygen to Ru. BP in the first step of the Calvin Cycle rather than carbon dioxide. • Whichever compound (O 2 or CO 2) is present in higher concentration will be joined by Rubisco to Ru. BP. • Photorespiration prevents the synthesis of glucose AND utilizes the plant’s ATP. • Photorespiration is a negative process for photosynthetic organisms Photosynthesis More CO 2 More O 2 Rubisco joins CO 2 to Ru. BP occurs; glucose is produced Rubisco joins O 2 to Ru. BP Photorespiration occurs; glucose is 29 NOT produced

Photorespiration • As the concentration of O 2 increases and the concentration of CO 2 decreases (due to the closure of the stomata to prevent excessive water loss), photorespiration is favored over photosynthesis. • Again, photorespiration is unfavorable for photosynthetic organisms. • It consumes ATP and does not produce glucose 30

Photorespiration • Photorespiration is primarily a problem for plants under water stress due to drought. • When plants are under water stress, their stomata close to prevent water loss through transpiration. • However, this also limits gas exchange. • O 2 is still being produced (through the light reactions). • Thus, the concentration of O 2 is increasing. • CO 2 is not entering the leaf since the stomata are closed. • Thus, as the CO 2 is being used up (in the Calvin Cycle) and not replenished, the concentration of CO 2 is decreasing. 31

Photorespiration • Strong selective pressure against photorespiration has favored the proliferation of adaptations that increase the evolutionary fitness of those organisms who possess these adaptations • Some plant species that live in hot, dry climates (where photorespiration is an especially big problem) have developed mechanisms through natural selection to prevent photorespiration. • C 4 plants • CAM plants 32

C 3 Plants • C 3 plants, which are “normal” plants, perform the light reactions and the Calvin Cycle in the mesophyll cells of the leaves. • The bundle sheath cells of C 3 plants do not contain chloroplasts palisade mesophyll spongy mesophyll bundle sheath cells 33

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C 4 and CAM Plants • C 4 plants and CAM plants modify the process of C 3 photosynthesis to prevent photorespiration. • C 4 plants perform the Calvin Cycle in a different location within the leaf than C 3 plants. • CAM plants obtain CO 2 at a different time than C 3 plants. 35

C 4 and CAM Plants • Both C 4 and CAM plants separate the initial fixing of CO 2 (carbon fixation) from the using of CO 2 in the Calvin Cycle. (by either space or time) • Both C 4 and CAM plants first use the enzyme PEP carboxylase instead of Rubisco so they are able to fix CO 2 in spite of the relatively high concentrations of O 2. Then they use that CO 2 separately in a normal Calvin cycle. 36

C 4 Plants: Preventing Photorespiration • Plants that use C 4 photosynthesis include corn, sugar cane, and sorghum. • In this process, CO 2 is transferred from the mesophyll cells into the bundle-sheath cells, which are impermeable to CO 2. • This increases the concentration of CO 2. • Thus, the Calvin Cycle is favored over photorespiration. • The bundle-sheath cells of C 4 plants do contain chloroplasts. 37

C 4 Plants: Preventing Photorespiration • Remember, Rubisco will join whichever compound is present in highest concentration (O 2 or CO 2) to Ru. BP; by shuttling CO 2 into the bundle-sheath cells from which CO 2 cannot escape, the concentration of CO 2 is increased, which leads to the joining of CO 2 to Ru. BP and the resultant production of organic compounds through the Calvin Cycle. 38

C 4 Plant Examples 39

C 4 Plants: Preventing Photorespiration • C 4 plants use the Hatch. Slack pathway prior to the Calvin Cycle: • PEP carboxylase adds carbon dioxide to PEP, a 3 -carbon compound, in the mesophyll cells. • This produces a 4 -carbon compound (which is why it’s known as C 4 photosynthesis). 40

C 4 Plants: Preventing Photorespiration • PEP carboxylase adds CO 2 to phosphoenolpyruvate (PEP), forming the four-carbon product oxaloacetate. • PEP carboxylase has a much higher affinity for CO 2 than does rubisco and no affinity for O 2. • Therefore, PEP carboxylase can fix carbon efficiently when rubisco cannot—that is, when it is hot and dry and stomata are partially closed, causing CO 2 concentration in the leaf to fall and O 2 concentration to rise. 41

C 4 Plants: Preventing Photorespiration • This 4 -carbon molecule then moves into the bundle-sheath cells via plasmodesmata (gaps between cell walls). • Once inside the bundle–sheath cells, CO 2 is released and the Calvin Cycle begins. 42

C 4 Plants: Preventing Photorespiration Talk to a Partner!! 43

C 4 Plants: Preventing Photorespiration How did the Hatch. Slack pathway come into existence? ? If the Hatch-Slack pathway helps to prevent photorespiration, why wouldn’t ALL plants have this adaptation? 44

CAM Plants: Preventing Photorespiration • Plants that use CAM photosynthesis include succulent plants (like cacti) and pineapples. • In CAM (crassulacean acid metabolism) photosynthesis, plants open their stomata at night to obtain CO 2 and release O 2. • This prevents them from drying out by keeping their stomata closed during the hottest & driest part of the day. 45

CAM PLANT EXAMPLES 46

CAM Plants: Preventing Photorespiration • When the stomata are opened at night, the CO 2 is converted to an organic acid (via the C 4 pathway) and stored overnight in the gaps of the mesophyll. 47

CAM Plants: Preventing Photorespiration • During the day – when light is present to for the Light Reactions to power the Calvin Cycle – carbon dioxide is released from the organic acid and used in the Calvin Cycle to produce organic compounds. • Remember: • Even though the CO 2 is taken in at night, the Calvin Cycle cannot occur because the Light Reactions can’t occur in the dark! 48

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Avoiding Photorespiration • Both C 4 and CAM plants – which are primarily found in hot, dry climates – have evolutionary adaptations which help prevent photorespiration. • C 4 plants perform the Calvin Cycle in the bundlesheath cells. • CAM plants open their stomata at night and store the CO 2 until morning. 50

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Created by: Cheryl Boggs Richmond, VA 2015