Citric acid cycle carbon and carrier balance 2026-10-18
The carbon sequence is oxaloacetate plus an acetyl group to citrate , then isocitrate , alpha-ketoglutarate , succinyl-CoA , succinate , fumarate , malate and back to oxaloacetate . The numbers count the cycling carbon skeleton, excluding coenzyme A. Two carbon dioxide releases accompany the changes .[ref]
Biology 2e by Mary Ann Clark and others. §7.3, Steps in the Citric Acid Cycle, steps 1–8 and Figure 7.11
Per acetyl group: three NADH, one FADH2 equivalent, one GTP or ATP by substrate-level phosphorylation, and two carbon dioxide. Succinate dehydrogenase retains FADH2 and transfers its Electrons to ubiquinone, forming ubiquinol. The FADH2 equivalent and ubiquinol count the same Electron pair.[ref]
Biology 2e by Mary Ann Clark and others. §7.3, Citric Acid Cycle and Steps in the Citric Acid Cycle, steps 3–8, especially step 6, enzyme-bound reduced carrier; §7.4, Q and Complex II: ubiquinone reduction to QH2
A useful check adds the stepwise changes. The carbon count is ; the NADH count is . Each NADH carries two Electrons, and the succinate dehydrogenase route supplies another pair, so one turn exports Electrons. This arithmetic is an accounting identity, not an ATP yield calculation. Converting it to ATP requires the separate energy coupling and transport assumptions discussed under oxidative phosphorylation.
For a chosen ideal conversion of of acetyl groups, multiplication gives of NADH, of ubiquinol, of GTP and of carbon dioxide. The Electron output is . If all these Electrons subsequently reach oxygen with no competing sinks, four Electrons per oxygen molecule require of oxygen. That is consumption by the connected electron transport chain; it is not an oxygen term in the cycle's own substrate ledger.
No cycle step directly consumes molecular oxygen. In ordinary aerobic mitochondrial operation, however, the mitochondrial respiratory chain must reoxidize NADH and ubiquinol to restore the oxidized electron carriers. Without that outlet, a finite oxidized-carrier pool cannot sustain continued turnover. Other electron acceptors and other forms of metabolism require their own analysis.[ref]
How Cells Obtain Energy from Food. Molecular Biology of the Cell, 4th edition by Bruce Alberts and others; Chapter 2, How Cells Obtain Energy from Food, The Citric Acid Cycle Generates NADH by Oxidizing Acetyl Groups to CO2: lack of direct O2 consumption and need to regenerate NAD+
The pyruvate dehydrogenase complex acts upstream: one three-carbon pyruvate gives one two-carbon acetyl group, one carbon dioxide and one NADH. These are additional to the cycle ledger.[ref] Consequently, processing one pyruvate through both stages gives three carbon dioxide and four NADH, while the succinate dehydrogenase contribution and the substrate-level phosphorylation contribution each remain one. Counting the upstream step inside the cycle would obscure where the extra carbon dioxide and NADH arise.
Electron carrier 2026-10-18
An electron carrier cycles between oxidation and reduction. Ubiquinone carries two Electrons, while cytochrome c carries one at a time.[ref]
Biology 2e by Mary Ann Clark and others. §7.4, Q and Complex II; Complex III: mobile carriers and electron capacities
For example, transferring the two Electrons supplied by one NADH requires two one-Electron deliveries by cytochrome c. It does not require two permanently consumed cytochrome c molecules: an available electron carrier can unload, return and reload. Electron carriers recycle material while the net redox conversion supplies free energy.
Mitochondrial respiratory chain 2026-10-18
NADH supplies two Electrons to respiratory complex I, then ubiquinone, respiratory complex III, cytochrome c and cytochrome c oxidase; terminal oxygen reduction forms water. Succinate dehydrogenase, or respiratory complex II, supplies another entry to ubiquinone. Respiratory complex I, respiratory complex III and cytochrome c oxidase move protons outward; respiratory complex II does not.[ref]
For an original Electron balance, supply exactly of NADH to an ideal preparation with no other electron donor or competing electron acceptor. The donor supplies of Electrons. Complete reduction of one oxygen molecule needs four Electrons, so the corresponding oxygen consumption is . Adding the two formal half-reactions makes the result explicit:The cancelling Electrons determine the donor-to-oxygen ratio. These chemical equations do not count transmembrane protons or determine an ATP yield. Chemical protons consumed in water formation and transported protons have different roles in the ledger.

