CHAPTER 10 PHOTOSYNTHESIS INTRODUCTION PHOTOSYNTHESIS- conversion of solar energy from the sun to chemical energy stored in sugar and other organic molecules - an organism obtains the organic compounds it needs for energy and carbon skeletons by one of two ways: Autotrophic nutrition/ Heterotrophic nutrition AUTOTROPHS Autotrophs can sustain themselves without eating other organisms or substances that come from other organisms - produce organic molecules from CO2 and other inorganic raw materials that come from the environment - they are the main sources of organic compounds for all other organisms called PRODUCERS of the biosphere - plants are autotrophs PHOTOAUTOTROPHS- organisms that use light as a source of energy to synthesize organic compounds - photosynthesis also occurs in algae and some prokaryotes HETEROTROPHS Heterotrophs are unable to make their own food, so they live on compounds produced by other organisms - they are the biosphere’s CONSUMERS - most common form is when an animal eats plants or other animals - some consume the remains of dead organisms DECOMPOSERS - almost all heterotrophs are completely dependent on photoautotrophs for food and oxygen CHLOROPLASTS All green parts of a plant have chloroplasts, but the main sites of photosynthesis are leaves - Color of the leaf is from CHLOROPHYLLgreen pigment found inside the chloroplast - light energy absorbed by chlorophyll drives the synthesis of food molecules in the chloroplast SOME RELATED TERMS - Chloroplasts are found mainly in the cells of the MESOPHYLL- interior tissue in leaf - CO2 enters the leaf and oxygen exits by way of microscopic pores called STOMATA - 2 membranes enclose the STROMA- the fluid inside the chloroplast - inside the inner membrane is another membrane system called THYLAKOIDS - in some places the thylakoid sacs are stacked in columns called GRANA - chlorophyll is found within the thylakoid membranes SUMMARY EQUATION 6 CO2 + 6 H2O + Light energy C6H12O6 + 6 O2 The overall chemical change is the reverse of the one that occurs in cellular respiration - but plants do not make food by simply reversing respiration SPLITTING OF WATER It was originally thought that the oxygen given off by plants came from CO2 and not water - chloroplasts split water into hydrogen and oxygen - original idea challenged by C.B. van Niel in the 1930s - studying photosynthesis in bacteria - 20 years later van Niel’s hypothesis was confirmed when scientists used an isotope to trace the fate of oxygen atoms during photosynthesis REDOX REACTIONS In respiration, energy is released from sugar when electrons are transported by carriers to oxygen - electrons lose potential energy as they are pulled down the chain, and mitochondria use that energy to make ATP Photosynthesis REVERSES the flow of electrons - water is split and electrons (along with H+) are transferred from water to CO2, reducing it to sugar - electrons increase in potential energy as they move from water to sugar OVERVIEW OF LIGHT REACTIONS AND CALVIN CYCLE Photosynthesis actually has 2 stages- the LIGHT REACTIONS and the CALVIN CYCLE LIGHT REACTIONS - steps of photosynthesis that convert solar energy to chemical energy - light energy that is absorbed by chlorophyll drives a transfer of electrons and hydrogen from water to an acceptor called NADP+, which stores the electrons - water is split, and O2 is given off - NADP+ is related to NAD+ (in respiration) - light reactions use solar power to reduce NADP+ to NADPH by adding a pair of electrons and an H+ - the light reactions generate ATP by the addition of a phosphate to ADP PHOSPHORYLATION - the light reactions produce no sugar CALVIN CYCLE - named for Melvin Calvin - incorporates CO2 from the air into organic molecules present in the chloroplast > CARBON FIXATION - then reduces fixed carbon to carbohydrate by the addition of electrons - reducing is provided by NADPH - Calvin cycle makes sugar, but can only do so with the NADPH and ATP made in the light reactions - sometimes called the dark reactions, or the light-independent reactions because steps do NOT require light directly The light reactions take place in the THYLAKOIDS The dark reactions take place in the STROMA LIGHT Light travels in waves of energy WATER WAVELENGTH- distance between crests of electromagnetic waves - range from very small (gamma rays) to very large (radio waves) ELECTROMAGNETIC SPECTRUM- entire range of radiation VISIBLE LIGHT- radiation that is most important to life (380 to 750 nm) - detected as colors Sometimes light behaves like it is made up of particles - these are called PHOTONS - act like objects in that each has a fixed amount of energy - this energy is inversely related to the wavelength of the light (shorter wavelength = greater energy of photon) - visible light drives photosynthesis PIGMENTS PIGMENT- a substance that absorbs visible light - different pigments absorb different wavelengths of light - wavelengths that are absorbed disappear - wavelengths that are transmitted are the ones we see - if a pigment absorbs all colors, it appears black Wavelength can be measured with a SPECTROPHOTOMETER - directs beams of light through a solution of a pigment and measures the fraction of the light TRANSMITTED at each wavelength ABSORPTION SPECTRUM- a graph plotting a pigment’s light absorption http://www.biology.lsu.edu/introbio/tutorial/Spec/spectrophotometer.JPG - blue and red light work best for photosynthesis - green is least effective ACCESSORY PIGMENTS Chlorophyll a is not the only important pigment in chloroplasts - only chlorophyll a can participate directly in the light reactions - other pigments capture light energy and transfer it to chlorophyll a - CHLOROPHYLL b- slightly different structure gives it a different color: Chlorophyll a = blue-green Chlorophyll b = yellow-green Other accessory pigments such as CAROTENOIDS are various shades of yellow and orange - broaden the spectrum of colors used for photosynthesis - function in photoprotection- absorb excess light energy that would damage chlorophyll “EXCITING” CHLOROPHYLL When a pigment molecule absorbs a photon, one of the molecule’s electrons is elevated to a higher energy level - the pigment molecule is then said to be “excited” - only photons of specific wavelengths are absorbed - the photons are absorbed by clusters of pigment molecules in the thylakoid membrane The electron cannot remain in this excited state for long - it is unstable - usually, when pigments absorb light, their excited electrons drop to a lower energy level, releasing energy as heat * what makes the top of an automobile so hot on a sunny day PHOTOSYSTEMS Inside the thylakoid membrane, chlorophyll is organized along with proteins and other organic molecules into PHOTOSYSTEMS - has a light-gathering complex made up of chlorophyll a, chlorophyll b, and carotenoids - enables the photosystem to harvest more light over more of the spectrum When any pigment molecule absorbs a photon: - energy is transferred from pigment to pigment until it reaches a particular chlorophyll a - only this chlorophyll a molecule is located in the REACTION CENTER- where the first light-driven reaction of photosynthesis takes place Along with the chlorophyll a molecule in the reaction center is the PRIMARY ELECTRON ACCEPTOR - In a Redox reaction, chlorophyll a in the reaction center loses one of its electrons to the primary electron acceptor - this happens when light excites the electron to a higher energy level - the electron acceptor “catches” the excited electron before it can return to its ground state Each photosystem functions in the chloroplast as a light-harvesting unit - the transfer of electrons from chlorophyll to the primary electron center is the first step of the light reactions 2 PHOTOSYSTEMS There are 2 types of photosystems found in the thylakoid membrane: PHOTOSYSTEM I & PHOTOSYSTEM II - each has a unique reaction center - the reaction center of photosystem I is called P700 because it most effectively absorbs light of wavelength 700 nm - the reaction center of photosystem II is called P680 - the 2 pigments are identical chlorophyll a molecules, but they are associated with different proteins- affects light-absorbing properties NEXT: HOW THE 2 PHOTOSYSTEMS WORK TOGETHER NONCYCLIC ELECTRON FLOW The key to making NADPH and ATP (needed for dark reactions) is a flow of electrons through the photosystems and other molecules in the thylakoid membrane There are 2 possible routes for electron flow: Cyclic and Noncyclic STEPS OF NONCYCLIC ELECTRON FLOW - most common route 1. Photosystem II absorbs light and an electron moves to a higher energy level, becoming excited - this electron is captured by the primary electron acceptor - chlorophyll is oxidized; electron “hole” must be filled 2. An enzyme extracts electrons from water and supplies them to photosytem II, replacing the lost electrons - this reaction splits water into 2 H atoms and an O atom - the O immediately combines with another O to form O2 (released) 3. Each excited electron passes from the primary electron acceptor of photosystem II to photosystem I by an electron transport chain - very similar to the one in cellular respiration 4. As electrons “fall” down the chain the energy that they release is harnessed by the thylakoid membrane to make ATP - called photophosphorylation (specifically noncyclic photophosphorylation) - this ATP will provide chemical energy to make sugar in the dark reactions 5. When electrons reach the bottom of the electron transport chain, they fill a “hole” in the reaction center of photosystem I - this hole is created when light energy drives an electron to the primary acceptor of photosystem I 6. The primary electron acceptor of photosystem I gives the electrons to a second electron transport chain - this chain transmits electrons to ferredoxin (Fd), an iron-containing protein - an enzyme called NADP+ reductase transfers electrons from Fd to NADP+ - this becomes NADPH, the other molecule needed in the dark reactions CYCLIC ELECTRON FLOW In CYCLIC ELECTRON FLOW, photosystem I is used, but not photosystem II - the electrons cycle back from ferredoxin (Fd) to the cytochrome complex, and then to the reaction center of photosystem I - NADPH is not made, and oxygen is not released - ATP is still made! - this is called CYCLIC PHOTOPHOSPHORYLATION Why is cyclic electron flow needed? - Noncyclic electron flow makes equal amounts of NADPH and ATP - the Calvin cycle (dark reactions) use more ATP than NADPH - cyclic flow makes up the difference by only producing ATP CHEMIOSMOSIS Chloroplasts and mitochondria both generate ATP by chemiosmosis - electron transport chain in a membrane pumps protons across the membrane as electrons fall down the chain - potential energy is stored in the form of an H+ gradient across a membrane - an ATP synthase complex is also built into the membrane - ATP synthase couples the diffusion of H+ down their gradient to the phosphorylation of ADP - some of the electron carriers are very similar in mitochondria and chloroplasts (cytochromes) - ATP synthase complexes are very similar DIFFERENCES IN CHEMIOSMOSIS - In mitochondria, electrons are extracted from food molecules - the photosystems of chloroplasts use light energy to drive electrons to the top of the chain (do not need food) - inner membrane of mitochondrion pumps protons from the mitochondrial matrix out to the intermembrane space - it is the thylakoid membrane of the chloroplast that pumps protons from the stroma into the thylakoid space - thylakoid membrane makes ATP as the H+ diffuse from the thylakoid space back to the stroma - ATP is made in the stroma - NADPH is produced on the side of the membrane facing the stroma CALVIN CYCLE The Calvin cycle is similar to the Krebs cycle in that a starting material is regenerated after molecules enter and leave the cycle - Carbon enters the Calvin cycle in the form of CO2 and leaves in the form of sugar - cycle uses ATP and NADPH - the carbohydrate produced in the Calvin cycle is not glucose, but a 3-carbon sugar named GLYCERALDEHYDE-3PHOSPHATE (G3P) - for one molecule of G3P to be made, the Calvin cycle must take place 3 times, fixing 3 molecules of CO2 - the Calvin cycle can be divided into 3 phases: Phase 1: Carbon fixation - the Calvin cycle attaches each CO2 molecule to a five-carbon sugar named RIBULOSE BISPHOSPHATE (RuBP) - the enzyme that catalyzes this reaction is RUBISCO - the product of this reaction immediately splits to form 2 molecules of 3PHOSPHOGLYCERATE (for each CO2) Phase 2: Reduction - each phosphoglycerate receives a phosphate group from ATP and becomes 1,3-bisphosphoglycerate - next a pair of electrons from NADPH reduces the 1,3-bisphosphoglycerate to G3P (the sugar) - not all of the G3P made is given off; some must be recycled to regenerate RuBP Phase 3: Regeneration of RuBP - the carbon skeletons of 5 molecules of G3P are rearranged into 3 molecules of RuBP - 3 ATP are needed for this step SUMMARY OF CALVIN CYCLE TOTALS (for making 1 G3P): - 9 ATP are consumed - 6 NADPH are consumed - the light reactions regenerate ATP and NADPH - the G3P made is the starting material for metabolic pathways that make other organic compounds - neither the light reactions nor the Calvin cycle can make sugar alone