Determine how the plant utilizes the energy from the light dependent reactions to build sugars from CO2. Identify some common adaptations to this metabolic pathway which allow plants to thrive in various environments. https://www.youtube.com/watch?v=hj_WKgn L6MI The following reactions all take place in the Stroma of the chloroplast. ATP and NADPH produced from the light dependent reactions are used to provide energy for synthesis of sugar from molecules of CO2. This is termed carbon fixation. This occurs in a cycle, whereby the primary reactant, a 5carbon molecule termed ribulose bisphosphate (RuBP), is regenerated constantly. For every three CO2 molecules that are fixed, one 3-carbon sugar is produced that can leave the cycle. Rubisco catalyzes the addition of CO2 to RuBP, and a 6-Carbon intermediate breaks into two three carbon molecules called 3-Phosphoglycerate (3PG). Think of these molecules as being low energy; they need to be energized to be useful to the plant. RuBisCO (Ribulose Bisphosphate Carboxylase/Oxygenase) is an enzyme that catalyzes the addition of CO2 to RuBP, the primary reactant of the Calvin Benson Cycle. Each of the 3PG molecules is phosphorylated by ATP, and then reduced by NADPH into a sugar called Glyceraldehyde 3-phosphate (G3P). If RuBisCO catalyzes three carboxylations, how many G3P molecules would we have? fun fact: RuBisCO is slow, and can only catalyze about 3-10 molecules of CO2 per second per enzyme. If the cell used all 6 molecules of G3P produced from 3 CO2 molecules, it would run out of RuBP very quickly. Rather, 5 of the G3P molecules are used to regenerate the 3 molecules of RuBP necessary for the reaction. This means, for every three CO2 molecules fixed, only one G3P molecule is available to be used for other purposes. RuBisCO not only catalyzes the addition of CO2 to RuBP, it can also catalyze the addition of oxygen. No useable sugar results from this reaction, though energy is still used. This is termed photorespiration. In this way, CO2 and O2 are competing for the active sites of RuBisCO. What conditions do you think could affect the rate of photorespiration? The main factor that affects the rate at which photorespiration occurs is O2 concentration vs CO2 concentration. O2 wins out in high temperatures, or in periods of water stress for the plant. Plants that undergo too much photorespiration are not productive, and will die if they can’t synthesize sugars. The “normal” form of photosynthesis we just learned is called C3 photosynthesis (the first step produces a 3 Carbon molecule). C2 is another term for the photorespiratory pathway (The first step produces a 2 Carbon molecule). Some plants have evolved other adaptations in order to reduce photorespiration. Two commonly studied adaptations are: C4 Photosynthesis CAM Photosynthesis some plants are constantly exposed to high temperatures, and minimal H2O. They require adaptations to survive. C4 Plants have a different leaf anatomy, with no spongy mesophyll. In C3 plants, Mesophyll cells are responsible for complete photosynthesis, whereas in C4 plants, Mesophyll cells sequester Carbon in a 4 Carbon molecule via a different metabolic pathway. This Carbon is then released into Bundle Sheath Cells where O2 concentrations are low and the normal Calvin-Benson Cycle can occur without much photorespiration. EP carboxylase is more efficient at carboxylation in high temperatures Amazingly, C4 Photosynthesis has evolved on up to 40 independent occasions, acting as a prime example of convergent evolution. Over 7500 species of plants utilize C4 Photosynthesis (~3% of terrestrial plant species). Corn is an example, as are many grasses and other plants exposed to large amounts of sunlight and higher temperatures. The C4 Rice Project Crassulacean Acid Metabolism (CAM) is a mechanism of storing Carbon Dioxide in a four carbon acid in Vacuoles overnight, so the plant can have its stomata remain shut during the day, and only open it at night. Plants that have evolved this mechanism exist in arid environments, and can avoid rapid evaporation of H2O from stomata. Cactus are examples of CAM Plant https://vimeo.com/7316737