Photosynthesis

Wednesday, March 06, 2013 11:00 PM

Photosynthesis

Photosynthesis - the process of converting energy in sunlight to energy in chemical bonds, especially glucose 6CO2 + 6H2O + light -> C6H12O6 + 6O2 Begins with light-absorbing pigments in plant cells o A pigment molecule is able to absorb energy from light only within a narrow range of wavelengths o In order to absorb as much of the entire bandwidth from sunlight as possible, different pigments, capable of absorbing different wavelengths, act together to optimize energy absorption o These pigments include the green chlorophyll a and chlorophyll b and the carotenoids, which are red, orange, or yellow o When the light is absorbed into one of these pigments, the energy from the light is incorporated into electrons within the atoms that make up the molecule o These energized electrons ("excited" electrons) are unstable and almost immediately reemit the absorbed energy o The process ends when the energy is absorbed by one of two special chlorophyll a molecules, P680 and P700; these two chlorophyll molecules, named with numbers that represent the wavelengths at which they absorb their maximum amounts of light (680 and 700 nanometers), are different from other chlorophyll molecules because of their association with various nearby pigments o Together with these other pigments, chlorophyll P700 forms a pigment cluster called photosystem I (PS I); chlorophyll P680 forms photosystem II (PS II)

Noncyclic Photophosphorylation
Photophosphorylation - phosphorylation using energy derived from light; noncyclic photophosphorylation begins with PS II and follows these steps: 1. Photosystem II - electrons trapped by P680 in PS II are energized by light 2. Primary electron acceptor - two energized electrons are passed to a molecule called the primary electron acceptor; this electron acceptor is called "primary" because it is the first in a chain of electron acceptors 3. Electron transport chain - electrons pass through an electron transport chain, which consists of proteins that pass electrons from one carrier protein to the next; some carrier proteins, like ferredoxin and cytochrome, include nonprotein parts containing iron; the electron transport chains in photosynthesis are analogous to those in oxidative phosphorylation 4. Phophorylation - as the two electrons move "down" the electron transport chain, they lose energy; the energy lost by the electrons as they pass along the electron transport chain is used to phosphorylate, on average, about 1.5 ATP molecules 5. Photosystem I - the electron transport chain terminates PS I (with P700); here the electrons are again energized by sunlight and passed to a primary electron acceptor (different from the one associated with PS II)

6. NADPH - the two electrons pass through a short electron transport chain; at the end of the chain, the two electrons combine with NADP+ and H+ to form NADPH; NADPH is a coenzyme; like NADH in respiration, NADPH is an energy-rich molecule 7. Splitting of water - the two electrons that originated in PS II are now incorporated into NADPH; the loss of these two electrons from PS II is replaced when water is split into two electrons; a manganese-containing protein complex catalyzes the reaction; the two electrons from water replace the lose electrons from PS II, one of the hydrogen ions provides the H in NADPH, and the oxygen contributes to the oxygen gas that is released Photophosphorylation takes the energy in light and the electrons in water to make ATP and NADPH; because these reactions require light, they are often called the light-dependent reactions or simply light reactions + + H2O + ADP + Pi + NADP + light -> ATP + NADPH + O2 + H

Cyclic Photophosphorylation

Occurs when the electrons energized in PS I are "recycled" Energized electrons from PS I join with protein carriers and generate ATP as they pass along the electron transport chain Electrons in cyclic photophosphorylation return to PS I; here they can be energized again to participate in cyclic or noncyclic photophosphorylation Cyclic photophosphorylation occurs simultaneously with noncyclic photophosphorylation to generate additional ATP Two electrons passing through cyclic photophosphorylation generate, on average, about 1 ATP

Calvin Cycle
The Calvin cycle "fixes" CO2; it takes chemically unreactive, inorganic carbon dioxide and incorporates it into an organic molecule that can be used in biological systems; the Calvin cycle repeats six times; steps include: 1. Carboxylation: 6CO2 combine with 6RUBP to produce 12PGA - the enzyme ribulose bisphosphate carboxylase / oxygenase, or rubisco, catalyzes the merging of CO2 and RuBP (ribulose bisphosphate); the Calcin cycle is referred to as C3 photosynthesis because the first product formed, PGA (phosphoglycerate), contains three carbon atoms; other names include the CalvinBenson cycle and the carbon reduction cycle 2. Reduction: 12 ATP and 12 NADPH are used to convert 12PGA to 12G3P - the energy in the ATP and NADPH molecules is incorporated into G3P (glyceraldehyde 3-phosphate of PGAL), thus making G3P a very energy-rich molecule; ADP, Pi, and NADP+ are released and then re-energized in noncyclic photophosphorylation 3. Regeneration: 6ATP are used to convert 10G3P to 6RuBP - regenerating the 6RuBP originally used to combine with 6CO2 allows the cycle to repeat 4. Carbohydrate synthesis - the two remaining G3_ are used to build glucose, a common energystoring molecule; other monosaccharides like fructose and maltose can be formed; in addition, glucose molecules can be combined to form disaccharides like sucrose and polysaccharides like starch and cellulose No light is directly used in the Calvin cycle; these reactions are called the light-independent reactions or even the dark reactions; still, the process cannot occur in the absence of light because it is dependent upon the energy from ATP and NADPH, and these two energy-rich molecules can be created only during photophosphorylation, which can occur only in light + + Calvin cycle = 6CO2 + 18ATP + 12NADPH + H -> 18ADP + 18Pi + 12NADP + 1 glucose

Chloroplasts
Chloroplasts - the sites where both the light-dependent and light-independent reactions of photosynthesis occur; have the following areas: 1. Outer membrane - consists of a double layer of phospholipids 2. Intermembrane space - the narrow area between the inner and outer membranes 3. Inner membrane - also a double phospholipid bilayer 4. Stroma - the fluid material that fills the area inside the inner membrane; the Calvin cycle occurs here, fixing carbon from CO2 to generate carbohydrate precursors (G3P) 5. Thylakoids - pancake-like membranes suspended in the stroma; individual membrane layers are thylakoids; an entire stack of thylakoids is a granum (plural, grana); the membranes of the thylakoids contain the protein complexes (including PS I and PS II), cytochromes, and other electron carriers of the light-dependent reactions 6. Thylakoid lumen - the inside of the thylakoid; hydrogen ions (protons) accumulate here

Chemiosmosis in Chloroplasts
Chemiosmosis - the mechanism of ATP generation that occurs when energy is stored in the form of a proton concentration gradient across a membrane; the process in chloroplasts is analogous to ATP generation in mitochondria; these are the steps of photophosphorylation: 1. Hydrogen ions (protons) accumulate inside the thylakoids - H+ are released into the lumen of the thylakoid when water is split by PSII; also, H+ are carried from the stroma into the lumen by a cytochrome in the electron transport chain (ETC) between PS II and PS I 2. A pH and electrical gradient across the thylakoid membrane is created - as H+ accumulate inside the thylakoid, the pH decreases; since some of these H+ come from outside the thylakoids (from the stroma), the H+ concentration decreases in the stroma and its pH increases; this creates a pH gradient consisting of differences in the concentration of H+ across the thylakoid membrane from a stroma pH 8 to a thylakoid pH 5 (a factor of 1000); since H+ are positively charged, their accumulation on the inside of the thylakoid creates an electric gradient (or voltage) as well 3. ATP synthases generate ATP - the pH and electrical gradient represent potential energy like water behind a dam; similar to a dam, channel proteins, called ATP synthases, allow the H+ to flow through the thylakoid membrane and out to the stroma; the energy generated by the passage of H+ (like the water through turbines in a dam) provides the energy for the ATP synthases to phosphorylate ADP to ATP; the passage of about three H+ is required to generate one ATP 4. The Calvin cycle produces G3P using NADPH and CO2 and ATP - at the end of the ETC following PS I, electrons combine with NADP+ and H+ to produce NADPH; with NADPH, ATP, and CO2, two G3P are generated and subsequently used to make glucose or other carbohydrates

Photorespiration

Rubisco is the most common protein on earth; however, it is not a particularly efficient molecule; it can fix CO2 and oxygen Photorespiration - fixation of oxygen Photorespiration leads to two problems 1. The carbon dioxide-fixing efficiency is reduced because, instead of fixing only carbon dioxide, rubisco fixes some oxygen as well 2. The products formed when oxygen is combined with RuBP do not lead to the production of useful, energy-rich molecules like glucose; instead, peroxisomes are found near chloroplasts where they function to break down photorespiration products; thus, considerable effort is

made by plants to rid the cell of the products of photorespiration; since the early atmosphere in which primitive plants originated contained very little oxygen, it is hypothesized that the early evolution of rubisco was not influenced by its oxygen-fixing handicap

C4 Photosynthesis

Improving upon photosynthetic efficiency, some plants have evolved a special "add-on" feature to C3 photosynthesis When CO2 enters the leaf, it is absorbed by the usual photosynthesizing cells, the mesophyll cells Instead of being fixed by rubisco into PGA, the carbon dioxide combines with PEP (phosphoenolpyruvate) to form OAA (oxaloacetate or oxaloacetic acid) The fixing enzyme is PEP carboxylase OAA, the first product of this pathway, has 4 carbon atoms, thus the name C4 photosynthesis OAA is then converted to malate and the malate is shuttled through plasmodesmata to specialized cells within the leaf, the bundle sheath cells Here malate is converted to pyruvate and carbon dioxide The pyruvate is then shuttled back to the mesophyll cells where one ATP (broken down to AMP, instead of ADP) is required to convert the pyruvate back to PEP Then the process repeats; the overall effect is to move carbon dioxide from mesophyll cells to the bundle sheath cells; thus carbon dioxide is spatially segregated The purpose for moving carbon dioxide to bundle sheath cells is to increase the efficiency of photosynthesis The bundle sheath cells surround the leaf veins and are themselves surrounded by densely packed mesophyll cells Since bundle sheath cells rarely make contact with an intercellular space, very little oxygen reaches them When malate delivers carbon dioxide to a bundle sheath cell, rubisco begins the Calvin cycle (C3 photosynthesis) Because little oxygen is present, rubisco can fix carbon dioxide without competition from oxygen; thus, little photrespiration takes place, and photosynthesis is more efficient In order for photosynthesis to occur, leaf pores, called stomata, just be open to allow carbon dioxide to enter; however, when the stomata are open, water can escape The higher rate of photosynthesis among C4 plants allows them to reduce the time that the stomata are open, thereby, reducing water loss; thus, C4 plants are found in hot, dry climates, where they possess an adaptive advantage over C3 plants; this advantage apparently compensates for the additional energy requirement (1 ATP to AMP) for C4 photosynthesis In short, C4 photosynthesis provides two advantages: minimizes photorespiration and reduces water loss

CAM Photosynthesis
Crassulacean acid metabolism (CAM) - an "add-on" feature to C3 photosynthesis; almost identical to C4 photosynthesis, with the following changes: 1. PEP carboxylase still fixes carbon dioxide to OAA, as in C4; instead of malate, however, OAA is converted to malic acid (a minor difference, since malate is merely the ionized form of malic acid) 2. Malic acid is shuttled to the vacuole of the cell (not moved out of the cell to bundle sheath cells as in regular C4)

3. At night, stomata are open, PEP carboxylase is active, and malic acid accumulates in the cell's vacuole 4. During the day, stomata are closed (the reverse of other plants); at this time, malic acid is shuttled out of the vacuole and converted back to OAA (requiring 1 ATP to ADP), releasing carbon dioxide; the carbon dioxide is now fixed by rubisco, and the Calvin cycle proceeds. In CAM, carbon dioxide is temporally segregated; the advantage of CAM is that photosynthesis can proceed during the day while the stomata are closed, greatly reducing water loss; CAM plants can grow in hot, dry environments with cool nights