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In: Anatomy and Physiology

What are the steps of the electron transport chain transport protons to the intermembrane space of...

What are the steps of the electron transport chain transport protons to the intermembrane space of the mitochondria?

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Expert Solution

The electron transport chain is a series of proteins and organic molecules found in the inner membrane of the mitochondria. Electrons are passed from one member of the transport chain to another in a series of redox reactions. Energy released in these reactions is captured as a proton gradient, which is then used to make ATP in a process called chemiosmosis. Together, the electron transport chain and chemiosmosis make up oxidative phosphorylation. The key steps of this process, include:

  • Delivery of electrons by NADH and FADH2. Reduced electron carriers (NADH and FADH2 from other steps of cellular respiration transfer their electrons to molecules near the beginning of the transport chain. In the process, they turn back into NAD+and FAD, which can be reused in other steps of cellular respiration.

  • All of the electrons that enter the transport chain come from NADH and FADH2_22​start subscript, 2, end subscript molecules produced during earlier stages of cellular respiration: glycolysis, pyruvate oxidation, and the citric acid cycle.

  • NADH is very good at donating electrons in redox reactions (that is, its electrons are at a high energy level), so it can transfer its electrons directly to complex I, turning back into NAD+^++start superscript, plus, end superscript. As electrons move through complex I in a series of redox reactions, energy is released, and the complex uses this energy to pump protons from the matrix into the intermembrane space.

  • FADH2 is not as good at donating electrons as NADH (that is, its electrons are at a lower energy level), so it cannot transfer its electrons to complex I. Instead, it feeds them into the transport chain through complex II, which does not pump protons across the membrane.

  • Because of this "bypass," each FADH2 molecule causes fewer protons to be pumped (and contributes less to the proton gradient) than an NADH.

  • Beyond the first two complexes, electrons from NADH and FADH2travel exactly the same route. Both complex I and complex II pass their electrons to a small, mobile electron carrier called ubiquinone (Q), which is reduced to form QH2_ and travels through the membrane, delivering the electrons to complex III. As electrons move through complex III, more H+ ions are pumped across the membrane, and the electrons are ultimately delivered to another mobile carrier called cytochrome C (cyt C). Cyt C carries the electrons to complex IV, where a final batch of H+ ions is pumped across the membrane. Complex IV passes the electrons to O2 which splits into two oxygen atoms and accepts protons from the matrix to form water. Four electrons are required to reduce each molecule of O2, and two water molecules are formed in the process.

  • Electron transfer and proton pumping. As electrons are passed down the chain, they move from a higher to a lower energy level, releasing energy. Some of the energy is used to pump H+ ions, moving them out of the matrix and into the intermembrane space. This pumping establishes an electrochemical gradient.

  • Splitting of oxygen to form water. At the end of the electron transport chain, electrons are transferred to molecular oxygen, which splits in half and takes up H+ to form water.

  • Gradient-driven synthesis of ATP. As H+ ions flow down their gradient and back into the matrix, they pass through an enzyme called ATP synthase, which harnesses the flow of protons to synthesize ATP.

Chemiosmosis

Complexes I, III, and IV of the electron transport chain are proton pumps. As electrons move energetically downhill, the complexes capture the released energy and use it to pump H+t ions from the matrix to the intermembrane space. This pumping forms an electrochemical gradient across the inner mitochondrial membrane. The gradient is sometimes called the proton-motive force, and you can think of it as a form of stored energy, kind of like a battery.

Like many other ions, protons can't pass directly through the phospholipid bilayer of the membrane because its core is too hydrophobic. Instead, H+ions can move down their concentration gradient only with the help of channel proteins that form hydrophilic tunnels across the membrane.

In the inner mitochondrial membrane, H+ions have just one channel available: a membrane-spanning protein known as ATP synthase. Conceptually, ATP synthase is a lot like a turbine in a hydroelectric power plant. Instead of being turned by water, it’s turned by the flow of H+ ions moving down their electrochemical gradient. As ATP synthase turns, it catalyzes the addition of a phosphate to ADP, capturing energy from the proton gradient as ATP.


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