The Calvin cycle is organized into three basic stages: fixation, reduction, and regeneration. In plants, carbon dioxide (CO2) enters the leaves through stomata, where it diffuses over short distances through intercellular spaces until it reaches the mesophyll cells. Once in the mesophyll cells, CO2 diffuses into the stroma of the chloroplast, the site of light-independent reactions of photosynthesis. These reactions actually have several names associated with them. Other names for light-independent reactions include the Calvin cycle, the Calvin-Benson cycle, and dark reactions. The most outdated name is dark reactions, which can be misleading because it implies incorrectly that the reaction only occurs at night or is independent of light, which is why most scientists and instructors no longer use it. The light-independent reactions of the Calvin cycle can be organized into three basic stages: fixation, reduction, and regeneration.
In the stroma, in addition to CO2,two other components are present to initiate the light-independent reactions: an enzyme called ribulose bisphosphate carboxylase (RuBisCO) and three molecules of ribulose bisphosphate (RuBP). RuBP has five atoms of carbon, flanked by two phosphates. RuBisCO catalyzes a reaction between CO2 and RuBP. For each CO2 molecule that reacts with one RuBP, two molecules of 3-phosphoglyceric acid (3-PGA) form. 3-PGA has three carbons and one phosphate. Each turn of the cycle involves only one RuBP and one carbon dioxide and forms two molecules of 3-PGA. The number of carbon atoms remains the same, as the atoms move to form new bonds during the reactions (3 atoms from 3CO2 + 15 atoms from 3RuBP = 18 atoms in 3 atoms of 3-PGA). This process is called carbon fixation because CO2 is “fixed” from an inorganic form into organic molecules.
ATP and NADPH are used to convert the six molecules of 3-PGA into six molecules of a chemical called glyceraldehyde 3-phosphate (G3P). This is a reduction reaction because it involves the gain of electrons by 3-PGA. Recall that a reduction is the gain of an electron by an atom or molecule. Six molecules of both ATP and NADPH are used. For ATP, energy is released with the loss of the terminal phosphate atom, converting it to ADP; for NADPH, both energy and a hydrogen atom are lost, converting it into NADP+. Both of these molecules return to the nearby light-dependent reactions to be reused and reenergized.
At this point, only one of the G3P molecules leaves the Calvin cycle and is sent to the cytoplasm to contribute to the formation of other compounds needed by the plant. Because the G3P exported from the chloroplast has three carbon atoms, it takes three “turns” of the Calvin cycle to fix enough net carbon to export one G3P. But each turn makes two G3Ps, thus three turns make six G3Ps. One is exported while the remaining five G3P molecules remain in the cycle and are used to regenerate RuBP, which enables the system to prepare for more CO2 to be fixed. Three more molecules of ATP are used in these regeneration reactions. The process of photosynthesis is often described as turning sunlight into sugars, and while that's broadly true, there are two distinct biochemical reactions taking place. The first uses the sunlight to create energy inside the cell and the second takes carbon dioxide and uses it to make sugars. The second is the Calvin cycle although the name is a little unfashionable nowadays. It's politer to refer to it as the Calvin–Benson-Bassham cycle or the reductive pentose phosphate cycle, but with all due apologies to Misters Benson and Bassham, the Calvin Cycle is quicker to write. Turning carbon dioxide into sugar may sound fairly magical, but it becomes a more conceivable when you consider that both carbon dioxide (CO2) and glucose (C6H12O6) contain roughly the same sort of elements. The Calvin cycle just adds on all the extra elements required. Having said that, the 'just' is still a fairly major task, requiring different enzymes all working in the correct order. The carbon dioxide molecules diffuse into the cells through small holes in the underside of the leaf. The first enzyme that picks them up is called Rubisco. Despite sounding like a small corporate venture, Rubisco is actually one of the most important enzymes in the world. Without Rubisco, plants would not be able to make sugars, which means that animals would not be able to survive on plants. Rubisco catalysis the connection of the small molecule ribulose-1.5-bisphosphate phosphate (RuBP) to carbon dioxide - therefore fixing the inorganic CO2 as an organic molecule. RuBP contains 5 carbons as well as oxygen, hydrogen and phosphate and it bonds to the CO2 to create a 6 carbon molecule. This promptly splits into two small 3 carbon molecules as shown in the reaction scheme below: These two 3 carbon molecules then go through a series of reduction stages, during which they react with the ATP (energy molecule) and NADPH (reducing molecule) that were produced during the light reactions of photosynthesis. Even though the Calvin cycle doesn't require any light itself, it is completely reliant on molecules created by the light-reactions. This stage creates two molecules of the 3-carbon "glyceraldehyde 3-phosphate" - which can be turned into useful plant sugars by further reactions. In order to continue running the Calvin cycle, and the reason that it is a cycle rather than just a process, the Rubisco must be recycled in order to go and pick up new carbon dioxide molecules. To do this also requires molecules of glyceraldehyde-3-phosphate - which are modified and then joined together to re-form the RuBP. The final result of all this is that for every 3 rounds of the cycle three molecules of RuBP go in, 3 RuBP come out, and one new glyceraldehyde 3-phosphate is made. When put like that it might seem like a lot of effort for very little, in reality it's a very stable and important cycle. As the components of the cycle are all recycled, the Rubisco can just keep picking up carbon dioxide and shooting out sugars, turning an inorganic gas into an energy molecule useful for life. --- Credit link for image 1 Credit link for image 2 Featured Image by Jon Sullivan. More of his awesome photos can be found here. The first draft of this was incorrectly posted too early and contained some of my place-holder notation and Unchecked Mathematics. Apologies to all who followed a broken link, and thanks to the commentators for alerting me to the issue (it slipped my mind that I'd set it to publish).
By the end of this section, you will be able to:
After the energy from the sun is converted and packaged into ATP and NADPH, the cell has the fuel needed to build food in the form of carbohydrate molecules. The carbohydrate molecules made will have a backbone of carbon atoms. Where does the carbon come from? The carbon atoms used to build carbohydrate molecules comes from carbon dioxide, the gas that animals exhale with each breath. The Calvin cycle is the term used for the reactions of photosynthesis that use the energy stored by the light-dependent reactions to form glucose and other carbohydrate molecules. The Interworkings of the Calvin Cycle Figure 1. Light-dependent reactions harness energy from the sun to produce ATP and NADPH. These energy-carrying molecules travel into the stroma where the Calvin cycle reactions take place. In plants, carbon dioxide (CO2) enters the chloroplast through the stomata and diffuses into the stroma of the chloroplast—the site of the Calvin cycle reactions where sugar is synthesized. The reactions are named after the scientist who discovered them, and reference the fact that the reactions function as a cycle. Others call it the Calvin-Benson cycle to include the name of another scientist involved in its discovery (Figure 1). The Calvin cycle reactions (Figure 2) can be organized into three basic stages: fixation, reduction, and regeneration. In the stroma, in addition to CO2, two other chemicals are present to initiate the Calvin cycle: an enzyme abbreviated RuBisCO, and the molecule ribulose bisphosphate (RuBP). RuBP has five atoms of carbon and a phosphate group on each end. RuBisCO catalyzes a reaction between CO2 and RuBP, which forms a six-carbon compound that is immediately converted into two three-carbon compounds. This process is called carbon fixation, because CO2 is “fixed” from its inorganic form into organic molecules. ATP and NADPH use their stored energy to convert the three-carbon compound, 3-PGA, into another three-carbon compound called G3P. This type of reaction is called a reduction reaction, because it involves the gain of electrons. A reduction is the gain of an electron by an atom or molecule. The molecules of ADP and NAD+, resulting from the reduction reaction, return to the light-dependent reactions to be re-energized. One of the G3P molecules leaves the Calvin cycle to contribute to the formation of the carbohydrate molecule, which is commonly glucose (C6H12O6). Because the carbohydrate molecule has six carbon atoms, it takes six turns of the Calvin cycle to make one carbohydrate molecule (one for each carbon dioxide molecule fixed). The remaining G3P molecules regenerate RuBP, which enables the system to prepare for the carbon-fixation step. ATP is also used in the regeneration of RuBP. Figure 2. The Calvin cycle has three stages. In stage 1, the enzyme RuBisCO incorporates carbon dioxide into an organic molecule. In stage 2, the organic molecule is reduced. In stage 3, RuBP, the molecule that starts the cycle, is regenerated so that the cycle can continue. In summary, it takes six turns of the Calvin cycle to fix six carbon atoms from CO2. These six turns require energy input from 12 ATP molecules and 12 NADPH molecules in the reduction step and 6 ATP molecules in the regeneration step.
Check out this animation of the Calvin cycle. Click Stage 1, Stage 2, and then Stage 3 to see G3P and ATP regenerate to form RuBP.
Figure 3. Living in the harsh conditions of the desert has led plants like this cactus to evolve variations in reactions outside the Calvin cycle. These variations increase efficiency and help conserve water and energy. (credit: Piotr Wojtkowski) The shared evolutionary history of all photosynthetic organisms is conspicuous, as the basic process has changed little over eras of time. Even between the giant tropical leaves in the rainforest and tiny cyanobacteria, the process and components of photosynthesis that use water as an electron donor remain largely the same. Photosystems function to absorb light and use electron transport chains to convert energy. The Calvin cycle reactions assemble carbohydrate molecules with this energy. However, as with all biochemical pathways, a variety of conditions leads to varied adaptations that affect the basic pattern. Photosynthesis in dry-climate plants (Figure 3) has evolved with adaptations that conserve water. In the harsh dry heat, every drop of water and precious energy must be used to survive. Two adaptations have evolved in such plants. In one form, a more efficient use of CO2 allows plants to photosynthesize even when CO2 is in short supply, as when the stomata are closed on hot days. The other adaptation performs preliminary reactions of the Calvin cycle at night, because opening the stomata at this time conserves water due to cooler temperatures. In addition, this adaptation has allowed plants to carry out low levels of photosynthesis without opening stomata at all, an extreme mechanism to face extremely dry periods. Photosynthesis in ProkaryotesThe two parts of photosynthesis—the light-dependent reactions and the Calvin cycle—have been described, as they take place in chloroplasts. However, prokaryotes, such as cyanobacteria, lack membrane-bound organelles. Prokaryotic photosynthetic autotrophic organisms have infoldings of the plasma membrane for chlorophyll attachment and photosynthesis (Figure 4). It is here that organisms like cyanobacteria can carry out photosynthesis. Figure 4. A photosynthetic prokaryote has infolded regions of the plasma membrane that function like thylakoids. Although these are not contained in an organelle, such as a chloroplast, all of the necessary components are present to carry out photosynthesis. (credit: scale-bar data from Matt Russell) The Energy CycleLiving things access energy by breaking down carbohydrate molecules. However, if plants make carbohydrate molecules, why would they need to break them down? Carbohydrates are storage molecules for energy in all living things. Although energy can be stored in molecules like ATP, carbohydrates are much more stable and efficient reservoirs for chemical energy. Photosynthetic organisms also carry out the reactions of respiration to harvest the energy that they have stored in carbohydrates, for example, plants have mitochondria in addition to chloroplasts. 6CO2+6H2O→C6H12O6+6O2 is the reverse of the overall reaction for cellular respiration: 6O2+C6H12O6→6CO2+6H2O Photosynthesis produces oxygen as a byproduct, and respiration produces carbon dioxide as a byproduct. In nature, there is no such thing as waste. Every single atom of matter is conserved, recycling indefinitely. Substances change form or move from one type of molecule to another, but never disappear (Figure 5). Figure 5. In the carbon cycle, the reactions of photosynthesis and cellular respiration share reciprocal reactants and products. (credit: modification of work by Stuart Bassil) CO2 is no more a form of waste produced by respiration than oxygen is a waste product of photosynthesis. Both are byproducts of reactions that move on to other reactions. Photosynthesis absorbs energy to build carbohydrates in chloroplasts, and aerobic cellular respiration releases energy by using oxygen to break down carbohydrates in mitochondria. Both organelles use electron transport chains to generate the energy necessary to drive other reactions. Photosynthesis and cellular respiration function in a biological cycle, allowing organisms to access life-sustaining energy that originates millions of miles away in a star. Section SummaryUsing the energy carriers formed in the first stage of photosynthesis, the Calvin cycle reactions fix CO2 from the environment to build carbohydrate molecules. An enzyme, RuBisCO, catalyzes the fixation reaction, by combining CO2 with RuBP. The resulting six-carbon compound is broken down into two three-carbon compounds, and the energy in ATP and NADPH is used to convert these molecules into G3P. One of the three-carbon molecules of G3P leaves the cycle to become a part of a carbohydrate molecule. The remaining G3P molecules stay in the cycle to be formed back into RuBP, which is ready to react with more CO2. Photosynthesis forms a balanced energy cycle with the process of cellular respiration. Plants are capable of both photosynthesis and cellular respiration, since they contain both chloroplasts and mitochondria.
1.Which part of the Calvin cycle would be affected if a cell could not produce the enzyme RuBisCO? 2. Explain the reciprocal nature of the net chemical reactions for photosynthesis and respiration.
1. None of the cycle could take place, because RuBisCO is essential in fixing carbon dioxide. Specifically, RuBisCO catalyzes the reaction between carbon dioxide and RuBP at the start of the cycle. 2. Photosynthesis takes the energy of sunlight and combines water and carbon dioxide to produce sugar and oxygen as a waste product. The reactions of respiration take sugar and consume oxygen to break it down into carbon dioxide and water, releasing energy. Thus, the reactants of photosynthesis are the products of respiration, and vice versa. |
Why is it important that carbon dioxide molecules from the atmosphere enter the Calvin cycle
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