OpenStax-CNX module: m56017 1 Bis2A 06.3 Photophosphorylation: The light reactions in photosynthesis ∗ Mitch Singer Based on The Light-Dependent Reactions of Photosynthesis† by OpenStax College This work is produced by OpenStax-CNX and licensed under the Creative Commons Attribution License 4.0‡ Abstract By the end of this section, you will be able to: • Explain how plants absorb energy from sunlight • Describe short and long wavelengths of light • Describe how and where photosynthesis takes place within a plant How can light be used to make food? When a person turns on a lamp, electrical energy becomes light energy. Like all other forms of kinetic energy, light can travel, change form, and be harnessed to do work. In the case of photosynthesis, light energy is converted into chemical energy, which photoautotrophs use to build carbohydrate molecules (Figure 1). ∗ Version 1.2: Jun 19, 2015 11:25 am -0500 † http://cnx.org/content/m44448/1.12/ ‡ http://creativecommons.org/licenses/by/4.0/ http://cnx.org/content/m56017/1.2/ OpenStax-CNX module: m56017 2 Figure 1: Photoautotrophs can capture light energy from the sun, converting it into the chemical energy used to build food molecules. (credit: Gerry Atwell) 1 What Is Light Energy? The sun emits an enormous amount of electromagnetic radiation (solar energy). Humans can see only a fraction of this energy, which portion is therefore referred to as visible light. The manner in which solar energy travels is described as waves. Scientists can determine the amount of energy of a wave by measuring its wavelength, the distance between consecutive points of a wave. A single wave is measured from two consecutive points, such as from crest to crest or from trough to trough (Figure 2). http://cnx.org/content/m56017/1.2/ OpenStax-CNX module: m56017 3 Figure 2: The wavelength of a single wave is the distance between two consecutive points of similar position (two crests or two troughs) along the wave. Visible light constitutes only one of many types of electromagnetic radiation emitted from the sun and other stars. Scientists dierentiate the various types of radiant energy from the sun within the electro- magnetic spectrum. The electromagnetic spectrum is the range of all possible frequencies of radiation (Figure 3). The dierence between wavelengths relates to the amount of energy carried by them. http://cnx.org/content/m56017/1.2/ OpenStax-CNX module: m56017 4 Figure 3: The sun emits energy in the form of electromagnetic radiation. This radiation exists at dierent wavelengths, each of which has its own characteristic energy. All electromagnetic radiation, including visible light, is characterized by its wavelength. Each type of electromagnetic radiation travels at a particular wavelength. The longer the wavelength (or the more stretched out it appears in the diagram), the less energy is carried. Short, tight waves carry the most energy. This may seem illogical, but think of it in terms of a piece of moving a heavy rope. It takes little eort by a person to move a rope in long, wide waves. To make a rope move in short, tight waves, a person would need to apply signicantly more energy. The electromagnetic spectrum (Figure 3) shows several types of electromagnetic radiation originating from the sun, including X-rays and ultraviolet (UV) rays. The higher-energy waves can penetrate tissues and damage cells and DNA, explaining why both X-rays and UV rays can be harmful to living organisms. 2 Absorption of Light Light energy initiates the process of photosynthesis when pigments absorb the light. Organic pigments, whether in the human retina or the chloroplast thylakoid, have a narrow range of energy levels that they can absorb. Energy levels lower than those represented by red light are insucient to raise an orbital electron to a populatable, excited (quantum) state. Energy levels higher than those in blue light will physically tear the molecules apart, called bleaching. So retinal pigments can only see (absorb) 700 nm to 400 nm light, which is therefore called visible light. For the same reasons, plants pigment molecules absorb only light in the wavelength range of 700 nm to 400 nm; plant physiologists refer to this range for plants as photosynthetically active radiation. The visible light seen by humans as white light actually exists in a rainbow of colors. Certain objects, such as a prism or a drop of water, disperse white light to reveal the colors to the human eye. The visible light portion of the electromagnetic spectrum shows the rainbow of colors, with violet and blue having shorter wavelengths, and therefore higher energy. At the other end of the spectrum toward red, the wavelengths are longer and have lower energy (Figure 4). http://cnx.org/content/m56017/1.2/ OpenStax-CNX module: m56017 5 Figure 4: The colors of visible light do not carry the same amount of energy. Violet has the shortest wavelength and therefore carries the most energy, whereas red has the longest wavelength and carries the least amount of energy. (credit: modication of work by NASA) 2.1 Understanding Pigments Dierent kinds of pigments exist, and each has evolved to absorb only certain wavelengths (colors) of visible light. Pigments reect or transmit the wavelengths they cannot absorb, making them appear in the corresponding color. Chlorophylls and carotenoids are the two major classes of photosynthetic pigments found in plants and algae; each class has multiple types of pigment molecules. There are ve major chlorophylls: and a related molecule found in bacteria called a, b, c and d bacteriochlorophyll. Chlorophyll a and chlorophyll b are found in higher plant chloroplasts and will be the focus of the following discussion. With dozens of dierent forms, carotenoids are a much larger group of pigments. The carotenoids found in fruitsuch as the red of tomato (lycopene), the yellow of corn seeds (zeaxanthin), or the orange of an orange peel (β -carotene)are used as advertisements to attract seed dispersers. In photosynthesis, carotenoids function as photosynthetic pigments that are very ecient molecules for the disposal of excess energy. When a leaf is exposed to full sun, the light-dependent reactions are required to process an enormous amount of energy; if that energy is not handled properly, it can do signicant damage. Therefore, many carotenoids reside in the thylakoid membrane, absorb excess energy, and safely dissipate that energy as heat. Each type of pigment can be identied by the specic pattern of wavelengths it absorbs from visible light, which is the a, absorption spectrum. chlorophyll b, The graph in Figure 5 shows the absorption spectra for chlorophyll and a type of carotenoid pigment called β -carotene (which absorbs blue and green light). Notice how each pigment has a distinct set of peaks and troughs, revealing a highly specic pattern of absorption. Chlorophyll a absorbs wavelengths from either end of the visible spectrum (blue and red), but not green. Because green is reected or transmitted, chlorophyll appears green. Carotenoids absorb in the short-wavelength blue region, and reect the longer yellow, red, and orange wavelengths. http://cnx.org/content/m56017/1.2/ OpenStax-CNX module: m56017 6 Figure 5: (a) Chlorophyll a, (b) chlorophyll b, and (c) β -carotene are hydrophobic organic pigments found in the thylakoid membrane. Chlorophyll a and b, which are identical except for the part indicated in the red box, are responsible for the green color of leaves. β -carotene is responsible for the orange color in carrots. Each pigment has (d) a unique absorbance spectrum. Many photosynthetic organisms have a mixture of pigments; using them, the organism can absorb energy from a wider range of wavelengths. Not all photosynthetic organisms have full access to sunlight. Some organisms grow underwater where light intensity and quality decrease and change with depth. Other organisms grow in competition for light. Plants on the rainforest oor must be able to absorb any bit of light that comes through, because the taller trees absorb most of the sunlight and scatter the remaining solar radiation (Figure 6). http://cnx.org/content/m56017/1.2/ OpenStax-CNX module: m56017 7 Figure 6: Plants that commonly grow in the shade have adapted to low levels of light by changing the relative concentrations of their chlorophyll pigments. (credit: Jason Hollinger) When studying a photosynthetic organism, scientists can determine the types of pigments present by generating absorption spectra. An instrument called a spectrophotometer can dierentiate which wave- lengths of light a substance can absorb. Spectrophotometers measure transmitted light and compute from it the absorption. By extracting pigments from leaves and placing these samples into a spectrophotometer, scientists can identify which wavelengths of light an organism can absorb. Additional methods for the identication of plant pigments include various types of chromatography that separate the pigments by their relative anities to solid and mobile phases. 3 What happens when a compound absorbs a photon of light? When a compound absorbs a photon of light, the compound becomes "excited", in the sense that it has this extra energy. This is illustrated in gure 7 schematically. The compound has some ground state, it absorbs a photon and now contains this excess absorbed energy and is considered "excited". The question because what does the compound do with this excess absorbed energy and how does it get back to its "ground" state. http://cnx.org/content/m56017/1.2/ OpenStax-CNX module: m56017 8 Figure 7: Energy diagram (energy story) of what happens to a molecule that absorbs a photon of light. What does the excited atom or molecule do to return to its ground state. The energy in this molecule can then be dissipated in a variety of ways as the excited electron decays back to its ground state. There are four possible outcomes, which are schematically diagrammed in Figure 8 below. These energy options are: 1. The energy can be dissipated as heat. - 2. The energy can be dissipated as uorescence as the e returns to a lower orbital. 3. The energy can be transferred by resonance to a neighboring molecule as the e orbital. - 4. The energy can change the reduction potential such that it can become an e - excited e - donor to a proper e - returns to a lower donor. Linking this acceptor can lead to the transduction of chemical energy this will be discussed in quite some detail in a later video and in lecture. For now lets just leave this as the ability of the excited molecule be converted into chemical work. In other words the excited state can be used in red/ox reactions. http://cnx.org/content/m56017/1.2/ OpenStax-CNX module: m56017 9 Figure 8 As the excited electron decays back to its original orbit, the energy can be released in a variety of ways. While many of the antenna or auxiliary pigments absorb light energy and transfer it to the reaction center (as depicted in option III in gure 8) it is what happens at the reaction center that we are most concerned with (option IV in gure 8). Here a chlorophyll or bacteriochlorophyll molecule absorbs the energy and an electron is excited. This energy transfer is sucient to allow the reaction center to donate the electron in a red/ox reaction to a second molecule. This initiates the photophosphorylation electron transport reactions. The result is an oxidized reaction center that must now be reduced in order to start the process again. How this happens is the basis of electron ow in photophosphorylation and will be described in detail below. 4 Photophosphorylation a brief synopsis Photophosphorylation is the process of converting light into chemical energy, ATP and NADPH. Photosynthesis is the integration of these light-reactions driving the reduction of CO2 to sugars, specically Glyceraldehyde-3-Phosphate, a triose. In this module we will focus on the rst part of photosynthesis, the light-reactions or the generation of ATP and NADPH from light. First and foremost it is important to realize that photophosphorylation and photosynthesis are very ancient sets of reactions. When we think of photosynthesis we mainly think of green plants; taking up http://cnx.org/content/m56017/1.2/ OpenStax-CNX module: m56017 CO 2 10 2 and giving o O . But this is a special, and from an evolutionary perspective, relatively new form of photophosphorylation. While extremely ecient and complicated, form of photophosphorylation that produces O photophosphorylation. 2 oxygenic photophosphorylation, the as a product, is only part of the picture of the evolution of Photophosphorylation has its roots in the anaerobic world, between 3 billion and 1.5 billion years ago, when life was abundant in the absence of molecular oxygen. atively shortly after electron transport chains and Photophosphorylation probably evolved rel- anaerobic respiration began to provide metabolic diversity. Think of photophosphorylation this way: it is simply a form of an electron transport chain. The major dierence is that instead of electrons being donated by a very strong reducing compound, such as NADH, light energy is used to "energize" an electron into a "high energy state". This "energized" electron can be donated to an electron transport chain, and as it decays, that is, as it passes from one electron carrier to another via red/ox reactions protons are pumped across a membrane. The pumping of these protons across a membrane leads to the generation of a PMF, which in turn results in the production of ATP. If enough light energy or photons can be absorbed and transferred to electrons, and if those electrons can have a lower (that is a more negative) reduction potential than NADP/NADPH, then they can be used reduce NADP to form NADPH. Therefore, photophosphorylation requires a compound that can absorb light energy or photons, use that energy to excite an electron and then donate that excited electron to NADPH. That compound is chlorophyll or bacteriochlorophyll. The nal piece of the photophosporylation story is nding something to reduce the oxidized bacteriochlorophyll, under anaerobic conditions, reduced sulfur compounds such as SH 2 0 and even elemental S are excellent electron donors (look at the redox tower provided in gure 9 to see more potential electron donors). anoxygenic photophosphorylation pathways could either make NADPH, in a noncyclic photophosphorylation or ATP, in a processes called cyclic photophosphoryla- These early, simple process called tion per donated electron. At some point, about 1.5 billion years ago, a chlorophyll molecule evolved that when oxidized (when a photon of light was absorbed and transferred to the electron which is ejected) had 2 a higher (more positive) reduction potential than O . Which meant that the oxidized form of chlorophyll could be reduced by water and generate molecular oxygen. for ever. That event, the great That event changed the shape of the planet oxygen event now begins to accumulate molecular oxygen, a toxic, highly corrosive and reactive compound, into the environment and life on this planet would change forever. 5 Simple Anoxygenic Photophosphorylation Systems Introduction For the early photophosphorylation systems no oxygen was generated. These reactions evolved in anaerobic environments, there was very little molecular oxygen available. Two sets of reactions evolved under these conditions, both directly from anaerobic respiratory chains. These are known as the light reactions because they require the activation of an electron (an excited electron)from the absorption of light energy by bacteriochlorophyll. The light reactions are categorized either as cyclic or as noncyclic photophosphorylation. To help you better understand the similarities of photophosphorylation to respiration, gure 9 below is an electron tower that will be useful in our discussion of photosphosphorylation. http://cnx.org/content/m56017/1.2/ OpenStax-CNX module: m56017 11 Figure 9: Electron tower that has a variety of common photophosphorylation components. PSI and PSII refer to Photosystems I and II of the oxygenic photophosphorylation pathways. For the examples in Figure 8 and Figure 9 P840 is similar in reduction potential as is PSI. Cyclic Photophosphorylation In cyclic photophosphorylation the bacteriochlorophyll and eject an electron forming bacteriochlorophyll ox . red molecule absorbs enough light energy to energize The electron reduces a carrier molecule in the reac- tion center which in turn reduces a series of carriers via red/ox reactions. carriers found in respiration. + suciently large, protons, H ox duce bacteriochlorophyll These carriers are the same If the change in reduction potential from the various red/ox reactions are are translocated across the membrane. Eventually the electron is used to re- and the whole process can start again. This is called cyclic photophosphorylation because the electrons make a complete circuit: bacteriochlorophyll is the source of electrons and is the nal electron acceptor. ATP is produced via the how cyclic photophosphorylation works. http://cnx.org/content/m56017/1.2/ F1 F0 ATPase. The schematic in gure 10 below demonstrates OpenStax-CNX module: m56017 12 Figure 10: Cyclic Photophosphorylation. The reaction center P840 absorbs light energy and becomes excited, denoted with an *. The excited electron is ejected and used to reduce an FeS protein leaving an oxidized reaction center. The electron its transferred to a quinone, then to a series of cytochromes which in term is reduces the P840 reaction center. The process is cyclical. Note the gray array coming from the FeS protein going to a ferridoxin (Fd), also in gray. This represents an alternative pathway the electron can take and will be discussed below in non-cyclic photophosphorylation. NOTE the same electron that leaves the P480 reaction center is not necessarily the same electron that eventually nds its way back to reduce the oxidized P840. Non-cyclic photophosphorylation In cyclic photophosphorylation electrons cycle from bacteriochlorophy (or chlorophyll) to a series of electron carriers and eventually back to bacteriochlorophyll (or chlorophyll): there is no loss of electrons, they stay in the system. In non-cyclic photophosphorylation the electrons are removed from the system, they eventually end up on NADPH. That means there needs to be a source of electrons, a source that has a higher reduction ox potential than bacteriochlorophyll (or chlorophyll) that can donate electrons to bacteriochlorophyll reduce it. to An electron tower is proved below so you can see what compounds can be used to reduce the oxidized form of bacteriochlorophyll. The second requirement, is that when bacteriochlorophyll becomes oxidized and the electron is ejected it must reduce a carrier that has a lower (more negative) reduction potential than NADP/NADPH (see the electron tower). http://cnx.org/content/m56017/1.2/ In this case, electrons can ow from energized OpenStax-CNX module: m56017 13 bacteriochlorophyll to NADP forming NADPH and oxidized bacteriochlorophyll. ox the system and end up on NADPH, to complete the circuit bacteriochlorophyll 0 2 Electrons are lost from is reduced by an external electron donor, such as H S or elemental S . This is diagrammed in gure 11 below. Figure 11: Non-cyclic photophosphorylation. In this example, the P840 reaction center absorbs light energy and becomes energized, the emitted electron reduced a FeS protein and in turn reduces ferridoxin. Reduced ferridoxin (Fdred ) can now reduce NADP to form NADPH. The electrons are now removed from the system, nding their way to NADPH. The electrons need to be replaced on P840, which requires an external electron donor. In this case, H2 S serves as the electron donor. Thought questions. It should be noted that per electron donated to the system, either NADPH or ATP is made, but not both. This puts a limit on the versatility of the system. But what would happen if both systems coexisted simultaneously? That is, if both ATP and NADPH could be formed from one electron? Additionally, these systems require compounds such as reduced sulfur to act as electron donors, not necessarily widely found compounds. What would happen if a chlorophyll 2 2 ox molecule would have a reduction potential higher than that of molecular the O /H O reaction? Answer, a planetary game changer. http://cnx.org/content/m56017/1.2/ OpenStax-CNX module: m56017 14 6 Oxygenic Photophosphorylation Generation of NADPH and ATP The overall function of light-dependent reactions is to convert solar energy into chemical energy in the form of NADPH and ATP. This chemical energy supports the light-independent reactions and fuels the assembly of sugar molecules. The light-dependent reactions are depicted in Figure 12. Protein complexes and pigment molecules work together to produce NADPH and ATP. Figure 12: A photosystem consists of a light-harvesting complex and a reaction center. Pigments in the light-harvesting complex pass light energy to two special chlorophyll a molecules in the reaction center. The light excites an electron from the chlorophyll a pair, which passes to the primary electron acceptor. The excited electron must then be replaced. In (a) photosystem II, the electron comes from the splitting of water, which releases oxygen as a waste product. In (b) photosystem I, the electron comes from the chloroplast electron transport chain discussed below. The actual step that converts light energy into chemical energy takes place in a multiprotein complex photosystem, two types of which are found embedded in the thylakoid membrane, photosystem II (PSII) and photosystem I (PSI) (Figure 13). The two complexes dier on the basis of what they oxidize called a (that is, the source of the low-energy electron supply) and what they reduce (the place to which they deliver their energized electrons). antenna proteins to which the chlororeaction center where the photochemistry takes place. Each photosystem is serviced by the light-harvesting complex, which passes energy from sunlight to the reacBoth photosystems have the same basic structure; a number of phyll molecules are bound surround the tion center; it consists of multiple antenna proteins that contain a mixture of 300400 chlorophyll molecules as well as other pigments like carotenoids. The absorption of a single a and b photon or distinct quantity or packet of light by any of the chlorophylls pushes that molecule into an excited state. In short, the light http://cnx.org/content/m56017/1.2/ OpenStax-CNX module: m56017 15 energy has now been captured by biological molecules but is not stored in any useful form yet. The energy is transferred from chlorophyll to chlorophyll until eventually (after about a millionth of a second), it is delivered to the reaction center. Up to this point, only energy has been transferred between molecules, not electrons. : Figure 13: In the photosystem II (PSII) reaction center, energy from sunlight is used to extract electrons from water. The electrons travel through the chloroplast electron transport chain to photosystem I (PSI), which reduces NADP+ to NADPH. The electron transport chain moves protons across the thylakoid membrane into the lumen. At the same time, splitting of water adds protons to the lumen, and reduction of NADPH removes protons from the stroma. The net result is a low pH in the thylakoid lumen, and a high pH in the stroma. ATP synthase uses this electrochemical gradient to make ATP. What is the initial source of electrons for the chloroplast electron transport chain? a.water b.oxygen c.carbon dioxide d.NADPH http://cnx.org/content/m56017/1.2/ OpenStax-CNX module: m56017 16 The reaction center contains a pair of chlorophyll a molecules with a special property. Those two chlorophylls can undergo oxidation upon excitation; they can actually give up an electron in a process called a photoact. It is at this step in the reaction center, this step in photosynthesis, that light energy is converted into an excited electron. All of the subsequent steps involve getting that electron onto the energy carrier NADPH for delivery to the Calvin cycle where the electron is deposited onto carbon for long-term storage in the form of a carbohydrate.PSII and PSI are two major components of the photosynthetic chain, which also includes the cytochrome complex. electron transport The cytochrome complex, an enzyme composed of two protein complexes, transfers the electrons from the carrier molecule plastoquinone (Pq) to the protein plastocyanin (Pc), thus enabling both the transfer of protons across the thylakoid membrane and the transfer of electrons from PSII to PSI. The reaction center of PSII (called primary electron acceptor, P680) delivers its high-energy electrons, one at the time, to the and through the electron transport chain (Pq to cytochrome complex to plastocyanine) to PSI. P680's missing electron is replaced by extracting a low-energy electron from water; 2 thus, water is split and PSII is re-reduced after every photoact. Splitting one H O molecule releases two electrons, two hydrogen atoms, and one atom of oxygen. Splitting two molecules is required to form one molecule of diatomic O 2 gas. About 10 percent of the oxygen is used by mitochondria in the leaf to support oxidative phosphorylation. The remainder escapes to the atmosphere where it is used by aerobic organisms to support respiration. As electrons move through the proteins that reside between PSII and PSI, they lose energy. That energy is used to move hydrogen atoms from the stromal side of the membrane to the thylakoid lumen. Those hydrogen atoms, plus the ones produced by splitting water, accumulate in the thylakoid lumen and will be used synthesize ATP in a later step. Because the electrons have lost energy prior to their arrival at PSI, they must be re-energized by PSI, hence, another photon is absorbed by the PSI antenna. That energy is relayed to the PSI reaction center (called P700). + P700 is oxidized and sends a high-energy electron to NADP to form NADPH. Thus, PSII captures the energy to create proton gradients to make ATP, and PSI captures the energy to reduce NADP + into NADPH. The two photosystems work in concert, in part, to guarantee that the production of NADPH will roughly equal the production of ATP. Other mechanisms exist to ne tune that ratio to exactly match the chloroplast's constantly changing energy needs. 6.1 Generating an Energy Carrier: ATP As in the intermembrane space of the mitochondria during cellular respiration, the buildup of hydrogen ions inside the thylakoid lumen creates a concentration gradient. The passive diusion of hydrogen ions from high concentration (in the thylakoid lumen) to low concentration (in the stroma) is harnessed to create ATP, just as in the electron transport chain of cellular respiration. The ions build up energy because of diusion and because they all have the same electrical charge, repelling each other. To release this energy, hydrogen ions will rush through any opening, similar to water jetting through a hole in a dam. In the thylakoid, that opening is a passage through a specialized protein channel called the ATP synthase. The energy released by the hydrogen ion stream allows ATP synthase to attach a third phosphate group to ADP, which forms a molecule of ATP (Figure 13). The ow of hydrogen ions through ATP synthase is called chemiosmosis because the ions move from an area of high to an area of low concentration through a semi-permeable structure. http://cnx.org/content/m56017/1.2/ OpenStax-CNX module: m56017 17 1 Visit this site : and click through the animation to view the process of photosynthesis within a leaf. Some interesting and informative videos • wiley presents photosynthesis2 • YouTube photosynthesis3 • YouTube photosynthesis and electron tower usage by Dr. Easlon 4 7 Section Summary The pigments of the rst part of photosynthesis, the light-dependent reactions, absorb energy from sunlight. A photon strikes the antenna pigments of photosystem II to initiate photosynthesis. The energy travels to the reaction center that contains chlorophyll a to the electron transport chain, which pumps hydrogen ions into the thylakoid interior. This action builds up a high concentration of ions. The ions ow through ATP synthase via chemiosmosis to form molecules of ATP, which are used for the formation of sugar molecules in the second stage of photosynthesis. Photosystem I absorbs a second photon, which results in the formation of an NADPH molecule, another energy and reducing power carrier for the light-independent reactions. 8 Art Connections Exercise 1 (Solution on p. 19.) Figure 13 What is the source of electrons for the chloroplast electron transport chain? a. Water b. Oxygen c. Carbon dioxide d. NADPH 9 Review Questions Exercise 2 Which of the following structures is not (Solution on p. 19.) a component of a photosystem? a. ATP synthase b. antenna molecule c. reaction center 1 http://openstaxcollege.org/l/light_reactions 2 http://www.wiley.com/legacy/college/boyer/0470003790/animations/photosynthesis/photosynthesis.htm 3 https://www.youtube.com/watch?v=joZ1EsA5_NY 4 https://www.youtube.com/watch?v=joZ1EsA5_NY http://cnx.org/content/m56017/1.2/ OpenStax-CNX module: m56017 18 d. primary electron acceptor Exercise 3 + How many photons does it take to fully reduce one molecule of NADP (Solution on p. 19.) to NADPH? a. 1 b. 2 c. 4 d. 8 Exercise 4 Which complex is not (Solution on p. 19.) involved in the establishment of conditions for ATP synthesis? a. photosystem I b. ATP synthase c. photosystem II d. cytochrome complex Exercise 5 (Solution on p. 19.) From which component of the light-dependent reactions does NADPH form most directly? a. photosystem II b. photosystem I c. cytochrome complex d. ATP synthase 10 Free Response Exercise 6 (Solution on p. 19.) Describe the pathway of electron transfer from photosystem II to photosystem I in light-dependent reactions. Exercise 7 What are the roles of ATP and NADPH in photosynthesis? http://cnx.org/content/m56017/1.2/ (Solution on p. 19.) OpenStax-CNX module: m56017 19 Solutions to Exercises in this Module to Exercise (p. 17) Figure 12 A. to Exercise (p. 17) A to Exercise (p. 18) B to Exercise (p. 18) C to Exercise (p. 18) B to Exercise (p. 18) A photon of light hits an antenna molecule in photosystem II, and the energy released by it travels through other antenna molecules to the reaction center. chlorophyll a The energy causes an electron to leave a molecule of to a primary electron acceptor protein. The electron travels through the electron transport chain and is accepted by a pigment molecule in photosystem I. to Exercise (p. 18) Both of these molecules carry energy; in the case of NADPH, it has reducing power that is used to fuel the process of making carbohydrate molecules in light-independent reactions. Glossary Denition 1: absorption spectrum range of wavelengths of electromagnetic radiation absorbed by a given substance Denition 2: antenna protein pigment molecule that directly absorbs light and transfers the energy absorbed to other pigment molecules Denition 3: carotenoid photosynthetic pigment that functions to dispose of excess energy Denition 4: chlorophyll a form of chlorophyll that absorbs violet-blue and red light and consequently has a bluish-green color; the only pigment molecule that performs the photochemistry by getting excited and losing an electron to the electron transport chain Denition 5: chlorophyll b accessory pigment that absorbs blue and red-orange light and consequently has a yellowish-green tint Denition 6: cytochrome complex group of reversibly oxidizable and reducible proteins that forms part of the electron transport chain between photosystem II and photosystem I Denition 7: electromagnetic spectrum range of all possible frequencies of radiation Denition 8: electron transport chain group of proteins between PSII and PSI that pass energized electrons and use the energy released by the electrons to move hydrogen ions against their concentration gradient into the thylakoid lumen http://cnx.org/content/m56017/1.2/ OpenStax-CNX module: m56017 20 Denition 9: light harvesting complex complex that passes energy from sunlight to the reaction center in each photosystem; it consists of multiple antenna proteins that contain a mixture of 300400 chlorophyll a and b molecules as well as other pigments like carotenoids Denition 10: P680 reaction center of photosystem II Denition 11: P700 reaction center of photosystem I Denition 12: photoact ejection of an electron from a reaction center using the energy of an absorbed photon Denition 13: photon distinct quantity or packet of light energy Denition 14: photosystem group of proteins, chlorophyll, and other pigments that are used in the light-dependent reactions of photosynthesis to absorb light energy and convert it into chemical energy Denition 15: photosystem I integral pigment and protein complex in thylakoid membranes that uses light energy to transport + electrons from plastocyanin to NADP (which becomes reduced to NADPH in the process) Denition 16: photosystem II integral protein and pigment complex in thylakoid membranes that transports electrons from water to the electron transport chain; oxygen is a product of PSII Denition 17: primary electron acceptor pigment or other organic molecule in the reaction center that accepts an energized electron from the reaction center Denition 18: reaction center complex of chlorophyll molecules and other organic molecules that is assembled around a special pair of chlorophyll molecules and a primary electron acceptor; capable of undergoing oxidation and reduction Denition 19: spectrophotometer instrument that can measure transmitted light and compute the absorption Denition 20: wavelength distance between consecutive points of equal position (two crests or two troughs) of a wave in a graphic representation; inversely proportional to the energy of the radiation http://cnx.org/content/m56017/1.2/
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