A CLOSER LOOK 3.5
A NEW LOOK AT THE ATP BALANCE SHEET
Historically, it was believed that aer. obic metabolism of one molecule of glucose resulted in the production of 38 ATP. However, more recent evidence indicates that this number overestimates the total ATP produc tion and that only 32 molecules of ATP actually reach the cytoplasm The explanation for this conclusion is that new evidence indicates that the energy provided by NADH and FADH is required not only for ATP produc- tion, but also to transport ATP across the mitochondrial membrane.Thisa added energy cost of ATP metabolism reduces the estimates of the total ATP yield from glucose. Specific details of this process follow For many years, it was believed that for every three H pumped into the intermembrane space, one mol ecule of ATP was produced and could be used for cellular energy. Although it is true that approximately three H must pass through the H+ channels (i.e., respiratory assemblies) to pro- duce one ATP, it is now known that another H is required to move the ATP molecule across the mitochon- drial membrane into the cytoplasm. The ATP and H+ are transported into the cytoplasm in exchange for ADP and Pi, which are transported into the mitochondria to resynthesize ATP. Therefore, while the theoretical yield of ATP from glucose is 38 mol- ecules, the actual ATP yield, allowing for the energy cost of transport, is only 32 molecules of ATP per glucose. For details of how these numbers are obtained, see the section "Aerobic ATPTally."
IN SUMMARY
Oxidative phosphorylation or aerobic ATP production occurs in the mitochondria as a re- sult of a complex interaction between the Krebs cycle and the electron transport chain. The primary role of the Krebs cycle is to complete the oxidation of substrates and form NADH and FADH to enter the electron transport chain. The end result of the electron transport chain is the formation of ATP and water. Water is formed by oxygen accepting electrons hence, the reason we breathe oxygen is to use it as the final accep tor of electrons in aerobic metabolism. AEROBIC ATP TALLY It is now possible to compute the overall ATP pro- duction as a result of the aerobic breakdown of glu- ose or glycogen. Let's begin bycountingthetotal
energy yield of glycolysis. Recall that the net ATP production of glycolysis was two ATP per glucose molecule. Furthermore, when O2 is present in the mitochondria, two NADH produced by glycolysis can then be shuttled into the mitochondria, with the energy used to synthesize an additional five ATP (Table 3.2). Thus, glycolysis can produce two ATP directly via substrate-level phosphorylation and an additional five ATP by the energy contained in the two molecules of NADH How many ATP are produced as a result of the oxidation-reduction activities of the Krebs cycle? Table 3.2 shows that two NADH are formed when pyruvate is converted to acetyl-CoA, which results in the formation of five ATP. Note that two GTP (sim- ilar to ATP) are produced via substrate-level phos phorylation. A total of six NADH and two FADH are produced in the Krebs cycle from one glucose mol ecule. Hence, the six NADH formed via the Krebs l cycle results in the production of a total of 15 ATP