Photosyntheis; Summary of what you need to know. Photosynthesis is simply the process by which organisms convert solar energy to chemical energy

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1 Photosyntheis; Summary of what you need to know. Understand the various forms of potential energy utilized in the photosynthetic and respiration processes; Describe the similarities and differences between the electron transport chains utilized in photosynthesis and aerobic respiration and how the oxidation and reduction of molecules and the flow of electrons through these electron transport chains is use to generate stable forms of energy in plants and animals (after the respiration lectures; Explain the general flow of electrons from water to NADP + through a series of mobile and membrane protein bound electron carriers in the Z-scheme and what ultimately powers this process; Discuss why photorespiration occurs and how plants have evolved to reduce this process Photosynthesis is simply the process by which organisms convert solar energy to chemical energy 12H20 + 6CO2 C6H12O6 + 6O2 + 6H2O or, in a more balanced form: 6H20 + 6CO2 C6H12O6 + 6O2 This is an energy requiring reaction - the energy source is sunlight Plants produce sugars as a source of food. However, they produce way more than they need to survive. This is good because all other life on earth must survive on the food energy obtained by this excess All photosynthesis occurs in the chloroplast, so let's review the anatomy of a chloroplast Electron Micrograph of a Chloroplast

2 The innermost membrane of the chloroplast is called the thylakoid membrane. The thylakoid membrane is folded upon itself forming many disks called grana (singular = granum). The "cytoplasm" of the chloroplast is called the stroma The photosynthesis reactions can be broken down into two components: The light-dependent reactions (the "light" reactions) - occur on the thylakoid membranes The light-independent reactions (the "dark" reactions) - occur in the stroma o Keep in mind that these are descriptive terms, but you should notice that the lightindependent reactions use materials produced in the light-dependent reactions, so in a sense, they are dependent on light... just don't fixate on this, OK? The light-dependent reactions Goal: To trap sunlight energy and store it as chemical energy to use in all life functions Why: ATP is a good source of energy, but it does not store well o This is like money, if you had 1 million dollars, would you rather carry around singles or gold? Where do these reactions occur?: on the membranes of the thylakoids How is it done? o Pigments - light harvesting molecules form an antenna complex on the thylakoid membranes o Each pigment absorbs a different type of light

3 Why is this good? It enables the plant to utilize a much wider range of light o In green plants, the primary photosynthetic pigments are Chlorophylls a & b. Wavelengths of light absorbed? - many wavelengths in the visible spectrum

4 How do the light-dependent reactions proceed? 1. A photon is absorbed by photosystem II (P680) 2. An electron is raised from a low energy state to a high energy state. The electron then falls down to the low energy state, releasing its energy. However, this energy is not lost, it is picked up by an adjacent pigment molecule where it is used to raise an electron to a higher energy state, etc. etc., until this energy reaches the photosystem (like a bucket brigade or "the wave" at a football game) 3. At the photosystem, the electron is raised, but instead of falling back down, it is stolen by another, electron deficient molecule in the electron transport chain 4. Meanwhile the photosystem's stolen electron is replenished by photolysis, or the splitting of H2O to form H +, electrons, and O2 (note: the H + is kept in side the thylakoid membrane). The O2 resulting is the source of all oxygen in our atmosphere 5. The electron travels down the electron transport system (ETS). Along the way, more H + is pumped into the thylakoid compartment. 6. The electron eventually reaches photosystem I (P700), where it waits until the electron is excited by another photon 7. The electron is stolen by another electron acceptor from a second ETS 8. The final fate of the electron is in converting NADP + to NADPH a. This captures the high-energy electron and puts it into a mobile molecule, NADPH (an "electron taxi"). These high-energy electrons will be used in the Calvin Cycle to reduce CO2 to make sugars 9. The H + is pumped into the thylakoid membrane to generate an H + gradient (i.e. source of potential energy) use to later synthesize ATP Light Reaction animation NADPH, NADH - Electron Carrier Molecules NADP+ and NAD+ are related molecules that serve as electron carriers NADP+/NADPH o Enzymes react NADP+ with either free electrons and H+ to form NADPH o NADPH floats off in the stroma and can be used to reduce a carbon (ultimately converting CO2 to Sugars) NAD+/NADH o Eznymes react NAD+ with food molecules containing C-H bonds to form NADH and H+ o The electrons are carried off in the matrix (or transported from the cytoplasm into the matrix) and then given to the electron transport chain o This reduction powers the pumping of H+ into the intermembrane space (this is explained in the respiration lectures

5 Non-Cyclic Photophosphorylation - A More Detailed Look The form of photosynthesis with which we are most familiar is non-cyclic photophosphorylation. It consists of two sets of pigments to excite. They are called PS1, or photosystem 1, and PS2, or photosystem 2. PS1 is better excited by light at about 700 nm, and is thus sometimes called P PS2 cannot use photons of wavelength longer than 680 nm, and is thus sometimes called P- 680.

6 Energy enters the system when PS2 becomes excited by light. Electrons are shed by the excited PS2 (oxidation), which grabs electrons from water, producing a molecule of oxygen gas for every two waters split. PS2 thus returns it to its unexcited state (reduction). The electrons are passed through a chain of oxidation-reduction reactions. Each arrow in the diagram above actually represents a reaction like this one: Each element in the pathway is reduced by the electrons, and turns right around to reduce its neighbor in the pathway by giving it the electrons, thus becoming reoxidized and ready for the next electrons to pass through the photosystem H + Pumping. Electrons leaving photosystem II are transferred transferred through a series of molecules and ultimately end up transferred to to a molecule called plastoquinone (PQ). The PQ reacts with two hydrogen ions from the stroma and the two electrons from photosystem II to form PQH2. 2H + stroma + 2e - + PQ PQH2 The PQH2 diffuses across the thylakoid membrane, passes the two electrons to the next electron carrier and releases the two hydrogen ions into the lumen. PQH2 PQ + 2H + lumen + 2e - PQ can then diffuse back across the membrane to repeat the process. The net result of the Q cycle is to move two hydrogen ions from the stroma to the lumen.

7 Electron Recovery Part of Photosystem II has the ability to split water and release oxygen. PSII is the only known biological molecule capable of oxidizing water. The electrons produced by the oxidation of water supply a steady source of electrons for Photosystem II. 2H2O 4H + + 2e - + O2 Cyclic Photophosphorylation Sometimes an organism has all the reductive power (NADPH) that it needs to synthesize new carbon skeletons, but still needs ATP to power other activities in the chloroplast. Many bacteria can shut off PS2, allowing the production of ATP in the absence of glucose. A proton gradient is generated across the membrane using the mechanisms of photosynthesis. This type of energy generation is called cyclic photophosphorylation. This may seem counter-intuitive. It appeared from noncyclic phtotphosphorylation that PS1 was responsible for NADPH production, while in cyclic photophosphorylation it is important for ATP production. This apparent dichotomy can be resolved when we understand what makes PS1 both a good candidate for noncyclic photophosphorylation and for NADPH production. PS1 is very good at transferring an electron, whether it be to NADP or to ferredoxin (fd). It is a powerful reductant. PS2, on the other hand, is better at grabbing electrons from water to transfer them to quinone (Q). It is a good oxidant. As you can see, the electron transferred is not derived from water, but from PS1 itself. It therefore must be recycled to PS1.

8 The light-independent reactions Goal: to take the recently created NADPH and ATP and store their energy by constructing sugars from CO2 Where: in the stroma of the chloroplast Where does the CO2 come from? o The atmosphere - the leaf opens up its stomates and lets CO2 in o When this happens, H2O is inadvertently released o The plant must always balance its carbon intake with water loss How is CO2 converted into sugar? o The energy is stored by converting CO2 into sugars in the Calvin-Benson Cycle

9 The Calvin-Benson Cycle A molecule of CO2 is taken in by the cell and is combined with RuBP (a five carbon sugar, abbreviated as 5C) to form a 6C intermediate sugar via an enzyme called RuBisCo. The 6C then breaks down to form 2 PGA's (3-phosphoglycerate) - each a 3C The PGA's undergoes a cyclic pathway, the Calvin-Benson cycle, which will eventually spit out 2 PGAL's (phosphoglyceraldehyde G3P in your book). Two PGAL's can form a sugar phosphate, which can then form a sugar Calvin Cycle animation Photorespiration One of the biggest faux pas (that's French for big "mistakes") of evolution RuBisCo is not only attracted to CO2, but it can also use O2 in the Calvin-Benson Cycle When O2 is used in the Calvin-Benson Cycle, no energy is stored - in fact, energy is lost! The reaction is as follows: O2 + RUBP 1 PGA + 1 Phosphogylcolate o There is very little use for phosphoglycolate in the plant, so the plant must spend energy to convert the phosphoglycolate back to a useful molecule and reclaim the two carbons o The conversion of phosphyglycolate occurs in the peroxysome Why does photorespiration occur?

10 When this evolved, the concentration of O2 was low - this was not a problem Plants have since evolved ways to reduce the damage caused by O2 in the Calvin-Benson Cycle Plants must spend up to 40% of their energy stored in sugars to deal with the damage created by RuBisCo fixing O2 in the Calvin-Benson Cycle Major efforts have been made to modify the properties of Rubisco to eliminate the oxygenation reaction, especially using molecular genetics all the results so far indicate that the two reactions cannot be separated. modifications in Rubisco that reduce the oxygenase activity also reduce the carboxylase activity Nature, however, has worked out a system to avoid photorespiration, it is called the C4 photosynthetic pathway. We will discuss this next lecture - can'tcha wait? Leaf Anatomy - The leaf is the primary photosynthetic organ of the plant It is has a large surface area to maximize light harvesting They are thin so that light will penetrate through to the bottom cells Composed of the lamina (blade) and the petiole (stalk) o Leaf shape is highly variable - there are literally 1000's of different shapes of leaves Simple leaf - developmentally it has one lamina per leaf o Look where the petiole meets the stem - you should see a bud Compound leaf - developmentally it has many distinct lamina per leaf o Look where each leaflet meets the rachis, there is no bud. o Look where the rachis meets the stem, there is a bud o Therefore, the entire structure is a leaf Leaf Epidermal Anatomy The outer epidermis is covered with a waxy cuticle to prevent water loss o This also prevents gas exchange :( Stomata (sing = stoma) are pores in the surface of leaves which allow for gas exchange o Pore surrounded by two guard cells o Guard cells open and close to allow gas exchange o The density of stomates is dependent upon ecological conditions like humidity and CO2 concentration The underside of leaves is usually covered with hairs (trichomes). o Many functions: catch water, reduce airflow, produce wax, etc. Leaf Internal Anatomy A "typical" leaf cross section

11 Upper and Lower Epidermis - protective function o Lower epidermis generally contains more stomates than upper epidermis (in dicots) o Epidermal cells lack chloroplasts Palisade Mesophyll - tightly packed cells on the upper surface o Contain three to five times as many chloroplasts as those of the spongy parenchyma. o Chloroplasts remain usually near the cell wall, since this adjustment guarantees optimal use of light Spongy Mesophyll - loosely arranged cells o Creates air spaces to facilitate gas exchange Leaves can have other uses besides photosynthesis

12 C 3 Photosynthesis The photosynthetic pathway we discussed in the previous lecture is known as the C3 pathway The first stable molecule formed after CO2 fixation is a 3-Carbon molecule Most (>90%) of all angiosperms are C3 plants Possess "typical" mesophyll arrangement Rubisco is exposed to O2 and the plant loses energy due to photorespiration C4 Photosynthesis - a Mechanism to Reduce the Effects of Photorespiration

13 Some plants have been observed to fix CO2 and initially form a 4-Carbon molecule. What is up with that? These same plants have an odd cross-sectional anatomy, called kranz anatomy (kranz is German for "wreath") Cross Section of Zea mays displaying Kranz anatomy The vascular bundles are surrounded by a special type of mesophyll cell which are collectively called the bundle sheath o The mesophyll cells do not have Rubisco o The bundle sheath cells have Rubisco and fix CO2 just like in C3 plants o But where do they get the CO2? The mesophyll cells have another CO2-fixing enzyme, PEP carboxylase o CO2 + PEP (phosphoenol pyruvate) >>> OAA (Oxaloacetate), a 4-Carbon compound o PEP Carboxylase has NO affinity for O2 o OAA >>> Malate o Malate is shuttled into the bundle sheath cell o The CO2 is removed, forming Pyruvate, a 3-Carbon compound o Pyruvate is shuttled back to the mesophyll cell where it is converted to PEP (requires ATP) o CO2 enters the Calvin-Benson cycle (exactly the same as in a C3 plant)

14 C4 Photosynthesis C4 Photosynthesis is found in many plants, mostly in drier climates C4 photosynthesis has evolved independently many times All of the enzymes involved in C4 photosynthesis were already present in the plant, so nothing new needed to evolve, just the sequence of operation Corn and sugar cane, two of the 10 most important crops worldwide, C4 plants C4 plants are more efficient than C3 plants in hot, dry environments, but in moist or cold environments, C3 plants can be more efficient PEP carboxylase has a much greater affinity for CO2 at high temps than does Rubisco, so it can assimilate carbon much more efficiently However, the CO2 shuttling costs energy, so this efficiency is lost in cooler temperatures

15 Also, the CO2 shuttling becomes saturated at a much lower CO2 concentration than does Rubisco, so when CO2 levels are high (such as when the stomates are wide open in moist tropical plants) C3 is more efficient C4 Photosynthesis and CAM animation Carbon dioxide yield of C4 and C3 plants of open grasslands in different parts of the world CAM Photosynthesis As if C4 wasn't enough, there is yet another addition/modification to the typical C3 photosynthetic pathway, called Crassulacean Acid Metabolism (CAM) Found almost exclusively in plants in xeric (dry) environments o mainly in succlents, cacti, etc. Plants open stomates during the night - they are kept closed during the day to conserve water The light-dependent reactions occur during the day, creating ATP and NADPH as expected During the night, the stomates of a CAM plant open, CO2 is taken up into the plant and incorporated into a variety of organic acids During the day, the light-dependent reactions proceed, making more ATP and NADPH - this promotes the release of CO2 from the organic acids and Rubisco operates as normal (but in a greatly CO2-enriched environment)

16 This is more of an adaptation to conserve water than to reduce the effects of photorespiration Food For Thought: What type of plant, C3 or C4, would you expect to show the greatest improvement if it were grown in an artificial environment with no O2? Global CO2 levels are on the rise - will this help or hinder plant growth on the earth? What about agricultural plants?

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