The Electron Transport Chain is responsible for the synthesis of most of the ATP in our body. In order to understand how the electron transport chain works, it is critical that you have a good understanding of what the mitochondria are and how it is organized.
The Mitochondria have an inner and an outer membrane. The inner membrane folds in and out on itself and these folds are called Cristae. Cristae increase the total surface area of the inner membrane. The center of the mitochondrion is called the matrix and is analogous to the cytoplasm of a cell. The Electron Transport Chain reactions take place on the inner membrane.
The term, electron transport refers to the proteins on the inner membrane of the mitochondria that will take hydrogen atoms and electrons from NADH and FADH2 and then ultimately use the energy in the electrons to make ATP. Recall that NAD+ and FAD picked up high energy electrons and hydrogens from C-H bonds in glycolysis (from the cytoplasm) and the citric acid cycle (in the matrix of the mitochondria).
In the inner membrane of the mitochondrion is a series of protein complexes that will receive the electrons and pass them from one complex to another. NADH passes 2 high energy electrons onto a protein complex (Complex I) in the inner membrane of the mitochondria. This complex is called NADH dehydrogenase. NADH dehydrogenase does two things. First, it accepts a pair of high energy electrons from NADH. Second, it uses some of the energy from these electrons to undergo a conformational change. This conformational change is associated with the movement of 4H+ ions from the mitochondrial matrix to the intermembranous space (the space between the inner and outer membranes of the mitochondria). Next, these two new electrons on Complex I are moved to Coenzyme Q (CoQ). Coenzyme Q is also called ubiquinone.
FADH2 also passes a pair of high energy electrons to a protein complex (Complex II), also called Succinate dehydrogenase. Complex II accepts the electrons but does not go through any conformational change that is associated with the movement of H+ ions. However, Complex II does pass the electrons to CoQ just like Complex I did. CoQ is a mobile shuttle that moves easily through the membrane and is able to relocate and react with Complex III. Complex III has a long name (Coenzyme Q-Cytochrome c Oxidoreductase). Complex III also goes by the name Cytochrome bc1 Complex. Complex III will undergo a conformational change that is associated with the movement of 4H+ ions from the mitochondrial matrix to the intermembranous space. The two electrons are then moved from Complex III to Cytochrome C (Cyt c). Cyt c another mobile shuttle that is a soluble protein in the intermembranous space that moves easily along the membrane and reacts with Complex IV. Complex IV, also called Cytochrome c Oxidase, uses some of the electron energy to undergo a conformational change that is associated with the movement of 2 H+ ions from the mitochondrial matrix to the intermembranous space. Oxygen receives the 2 electrons from Complex IV and reacts with H+ available in the surrounding fluid to make H2O or water.
A review of figure 11 below reveals that one NADH results in the movement of 10 H+ ions from the mitochondrial matrix to the intermembranous space. One FADH2 results in the active transport of 6 H+ ions. The important message in all of this is that electron energy is used to transport H+ ions to the intermembranous space and this sets up an electrochemical gradient that favors the movement of H+ ions back into the matrix. This is allowed to happen through another protein complex called ATP synthase. The diffusion of H+ ions through ATP synthase is called "chemiosmosis."
ATP synthase is made up of two main components. The component found underneath or in the mitochondrial matrix is capable of rotating and also binds ADP, Pi and ATP. ATP synthase also contains channels that allow the diffusion of H+ ions. As H+ ions diffuse back into the matrix, ATP synthase is physically rotated and enabled to react with ADP and Pi. There is a resulting phosphorylation of ADP and this yields ATP. This process is called Oxidative Phosphorylation.
For each pair of electrons that move from Complex I to Complex IV, about 2.5 ATP can be produced. For each pair of electrons that move from Complex II to Complex IV, about 1.5 ATP can be produced. Therefore, if we round up, it is often stated that each NADH yields 3 ATP while each FADH2 will yield 2 ATP.
Electron Transport Chain. Image created by JS at BYU Idaho F2013.
The image above illustrates the Electron Transport Chain. The protein complexes on the inner mitochondrial membrane use high energy electrons from NADH and FAD2 to move H+ ions to the intermembranous space. The H+ concentration gradient is then used to make ATP through the enzyme complex called ATP Synthase. Oxygen is the final electron acceptor and becomes water.
A quick recap of what has happened so far might go like this: Electrons and hydrogen ions were harvested from the C-H bonds of glucose. These high energy electrons with hydrogen are carried from the reactions of glycolysis and the citric acid cycle to the electron transport chain on the inner membrane of the mitochondria. The electron transport chain takes these high energy electrons and gradually "uses" the energy to pump hydrogen ions into the intermembranous space. As the energy in the electrons is used, the electrons don't have enough energy to form a C-H bond anymore, but they can form an O-H bond. Thus, oxygen comes along and accepts the electrons and hydrogen to form water. The cycle is complete and water can once again be used by a plant somewhere to participate in the photosynthetic reactions that will excite O-H bond electrons again. The hydrogen ions that have been pumped into the intermembranous space are allowed to flow down their electrochemical gradient through ATP synthase. ATP is generated as a result and ATP is used to run the many molecular processes in our cells that keep us healthy and alive. As energy is released in these many reactions of metabolism a little bit is lost as heat. Indeed, metabolism is responsible for a portion of our body heat.
Glucose is not the only molecule with C-H bond energy to use in metabolic reactions. Lipids and Proteins are also metabolized by our cells.
We will continue on with our summary of the metabolic process using the Electron Transport Chain process magnified from the “Metabolism Summary” image from section 8.1. (Green numbers from summary correlate with green numbers on the image below.)
Electron Transport Chain, from the “Big Picture” of Metabolism:Glycolysis, Citric Acid (Krebs) Cycle, Electron Transport Chain, Beta Oxidation and Lipolysis.Image created at BYU-Idaho by JS 2010
11 NADH is carrying a proton and 2 high energy electrons that need to be “dropped off”. FADH2 is also carrying high energy electrons and a couple of protons. These electron “carriers” are able to donate these electrons to an enzyme complex found in the inner mitochondrial membrane. Think of the “electron transport chain” as being like a bucket brigade. A series of proteins pass 2 electrons from one to another. Sometimes when the electrons are passed, a little bit of the energy from the electrons is used to induce a conformational change in some of the protein structures. This conformational change results in the transport of protons from the inside of the mitochondria to the intermembranous space (the space between the inner and outer mitochondrial membranes). Also, some of the energy released as the electrons move through the electron transport chain is given off as heat. NADH donates to the electrons to the electron transport chain at complex I and FADH2 donates electrons at complex II. There are more protons pumped from electrons moving down the transport chain from complex I than from complex II. For this reason, NADH yields more ATP ultimately than FADH2. Whether from NADH or FADH2, any donated electrons will move down the transport chain to the last protein acceptor and cannot go back to previous components of the chain.
Oxygen accepts the electrons from the last protein complex (complex IV) of the chain. As oxygen accepts the electrons, the oxygen becomes reactive and capable of forming a covalent bond with two protons and water is formed (H2O). If you have ever wondered what oxygen does and why it is so important to breathe into our bodies, now you know. Oxygen is the final electron acceptor.
Notice that NADH becomes NAD+ at the beginning of the electron transport chain. Also, FADH2 becomes FAD. This recycles these electron carriers such that they can be used again in earlier metabolic reactions. This has been mentioned, but it is worth mentioning again. Without NAD+, reactions that use NAD+ cannot occur.
Think about this question: What if there were no oxygen available? Predict what would happen to the metabolic pathways discussed…
Answer: If there were no oxygen, then the high energy electrons would sit on the protein complexes in the electron transport chain. They can’t go back and they can’t move forward because there is no FINAL electron acceptor. Complex I would have electrons that couldn’t move as well, so NADH could not “drop off” any more electrons. Since NADH could not become NAD+, it would not be very long before a severe shortage of NAD+ stopped the metabolic reactions from occurring. This would basically grind the Citric Acid Cycle to a stop. Also, pyruvate could not become Acetyl CoA. Basically, none of the mitochondrial metabolic processes could occur. Aerobic Metabolism would not be possible. However, Glycolysis would still be possible. You might recall the possibility to regenerate NAD+ in the cytoplasm if Lactic acid is made (Anaerobic Metabolism).
Hopefully, now you can see why oxygen is so important.
12 In step 11, we learned that as high energy electrons passed down the chain of protein acceptors, energy was used to move H+ ions into the intermembranous space. This generates a proton gradient. This means that there will be a higher concentration of protons in the intermembranous space than there is inside the mitochondrial matrix. This proton gradient represents “potential energy” because the protons will try to flow down their gradient if a passageway opens up and allows such movement.
Step 12 represents the idea sometimes referred to as chemiosmosis. This is a term that refers to the fact that protons tend to flow down their gradient through a selective protein channel. This protein channel, called ATP-Synthase is a very intricate and specialized molecular machine. This protein literally turns as the protons come through it and this kinetic energy is used to bring ADP and inorganic phosphate together so that ATP is created. The synthesis of ATP through chemiosmosis is referred to as Oxidative Phosphorylation.
While glycolysis gives us 2 ATP per glucose molecule, the electron transport chain gives us approximately 34 ATP per glucose molecule. We say “approximately” because it is difficult to say exactly how many ATP we get. This is because some ATP is used to shuttle molecules in and out of the mitochondria and there is likely some “leaking” that occurs when protons from the intermembranous space accidentally escape by some other way than through the ATP synthase enzyme complex. However, it is generally accepted that aerobic metabolism yield between 18 and 19 times more ATP than anaerobic metabolism. It is a good thing that our cells have mitochondria!
Although aerobic metabolism gives us much more ATP, it takes longer to do it. You might think of aerobic metabolism being useful for endurance activities but less useful for activities that require both high speed and high-intensity work.
Conversion Rate to ATP
ATP Yield through Electron Transport
2 NADH from Glycolysis
2 NADH from Pyruvate Oxidation
6 NADH from Citric Acid Cycle
3 ATP / NADH
2 FADH from Citric Acid Cycle
2 ATP /FADH
Net Total from Electron Transport
Net Total from Glycolysis + Citric Acid Cycle (Remember 2 ATP used to start glycolysis)
4 - 2 + 2 = 4 ATP
Total ATP from the complete aerobic metabolism of one glucose molecule
~38 ATP (minus some loss)
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