II. BACTERIAL GROWTH AND MICROBIAL METABOLISM

D. Cellular Respiration

1. Aerobic Respiration

d. The Electron Transport Chain and Chemiosmosis

Fundamental statements for this learning object:

1. Aerobic respiration involves four stages: glycolysis, a transition reaction that forms acetyl coenzyme A, the citric acid (Krebs) cycle, and an electron transport chain and chemiosmosis.
2. During various steps in glycolysis and the citric acid cycle, the oxidation of certain intermediate precursor molecules causes the reduction of NAD⁺ to NADH + H⁺ and FAD to FADH₂. NADH and FADH₂ then transfer protons and electrons to the electron transport chain to produce additional ATPs by oxidative phosphorylation.
3. The electron transport chain consists of a series of electron carriers that eventually transfer electrons from NADH and FADH₂ to oxygen.
4. The chemiosmotic theory states that the transfer of electrons down an electron transport system through a series of oxidation-reduction reactions releases energy. This energy allows certain carriers in the chain to transport hydrogen ions (H⁺ or protons) across a membrane.
5. As the hydrogen ions accumulate on one side of a membrane, the concentration of hydrogen ions creates an electrochemical gradient or potential difference (voltage) across the membrane called proton motive force.
6. This proton motive force provides the energy necessary for enzymes called ATP synthases, also located in the membranes mentioned above, to catalyze the synthesis of ATP from ADP and phosphate.
7. During aerobic respiration, the last electron carrier in the membrane transfers 2 electrons to half an oxygen molecule (an oxygen atom) that simultaneously combines with 2 protons from the surrounding medium to produce water as an end product.

Learning Objectives for this Section

The Electron Transport Chain and Chemiosmosis

During various steps in glycolysis and the citric acid cycle, the oxidation of certain intermediate precursor molecules causes the reduction of NAD⁺ to NADH + H⁺ and FAD to FADH₂. NADH and FADH2 then transfer protons and electrons to the electron transport chain to produce additional ATPs by oxidative phosphorylation (def).

As mentioned in the previous section on energy, during the process of aerobic respiration, coupled oxidation-reduction reactions and electron carriers are often part of what is called an electron transport chain (def), a series of electron carriers that eventually transfers electrons from NADH and FADH₂ to oxygen. The diffusible electron carriers NADH and FADH₂ carry hydrogen atoms (protons and electrons) from substrates in exergonic catabolic pathways such as glycolysis and the citric acid cycle to other electron carriers that are embedded in membranes. These membrane-associated electron carriers include flavoproteins, iron-sulfur proteins, quinones, and cytochromes. The last electron carrier in the electron transport chain transfers the electrons to the terminal electron acceptor, oxygen.

The chemiosmotic theory (def) explains the functioning of electron transport chains. According to this theory, the transfer of electrons down an electron transport system through a series of oxidation-reduction reactions (def) releases energy (see Fig. 1). This energy allows certain carriers in the chain to transport hydrogen ions (H⁺ or protons) across a membrane.

Depending on the type of cell, the electron transport chain may be found in the cytoplasmic membrane or the inner membrane of mitochondria.

• In prokaryotic cells, the protons are transported from the cytoplasm of the bacterium across the cytoplasmic membrane to the periplasmic space located between the cytoplasmic membrane and the cell wall .
• In eukaryotic cells, protons are transported from the matrix of the mitochondria across the inner mitochondrial membrane to the intermembrane space located between the inner and outer mitochondrial membranes (see Fig. 4).

As the hydrogen ions accumulate on one side of a membrane, the concentration of hydrogen ions creates an electrochemical gradient or potential difference (voltage) across the membrane. (The fluid on the side of the membrane where the protons accumulate acquires a positive charge; the fluid on the opposite side of the membrane is left with a negative charge.) The energized state of the membrane as a result of this charge separation is called proton motive force (def) or PMF.

This proton motive force provides the energy necessary for enzymes called ATP synthases (see Fig. 5), also located in the membranes mentioned above, to catalyze the synthesis of ATP from ADP and phosphate. This generation of ATP occurs as the protons cross the membrane through the ATP synthase complexes and re-enter either the bacterial cytoplasm (see Fig. 2) or the matrix of the mitochondria. As the protons move down the concentration gradient through the ATP synthase, the energy released causes the rotor and rod of the ATP synthase to rotate. The mechanical energy from this rotation is converted into chemical energy as phosphate is added to ADP to form ATP.

Proton motive force is also used to transport substances across membranes during active transport and to rotate bacterial flagella.

At the end of the electron transport chain involved in aerobic respiration, the last electron carrier in the membrane transfers 2 electrons to half an oxygen molecule (an oxygen atom) that simultaneously combines with 2 protons from the surrounding medium to produce water as an end product (see Fig. 3).

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