Primer · Nutritional Biochemistry

Energy Metabolism

How glucose, fat, and protein all become ATP — and how the body runs the same machinery in reverse to build fat, build glucose, and make ketones — mapped against the B vitamins that make every step run.

Abstract

Three fuels — glucose, fat, and, when needed, protein — funnel down to one molecule, acetyl‑CoA, and one hub, the Krebs cycle. This paper is built around flow more than mechanism: the diagrams carry the argument, and the text is there to name what the arrows mean. A small, fixed set of B‑vitamin‑derived coenzymes runs the whole system — TPP (B1), FAD (B2), NAD+ (B3), CoA (B5), PLP (B6), and biotin (B7) — and the same handful of molecules (pyruvate, acetyl-CoA, citrate) keep reappearing as the junctions where the body decides which way to run the traffic: toward ATP, or toward building something.

A note on sourcing

The pathway facts here are standard biochemistry (Lehninger, Harper's Illustrated Biochemistry) and not in dispute. Organizing the pathways around which B vitamin does which specific job follows the teaching style of Chris Masterjohn.

The path in brief

Three fuels in, four ways the traffic can reverse, six parts

  1. Part IGlycolysis splits glucose into two pyruvate in the cytosol on niacin (NAD+) alone — then pyruvate dehydrogenase, the very next step, pulls in thiamine (B1) along with three more cofactors to make acetyl-CoA.
  2. Part IIβ-oxidation spirals a fatty acid down two carbons per turn, needing riboflavin, niacin, and pantothenic acid every single turn.
  3. Part IIIBoth routes hand off to the Krebs cycle as acetyl-CoA — whose two decarboxylation steps are the densest cofactor dependencies in the whole system.
  4. Part IVProtein feeds in too: transamination, a PLP (B6)-dependent swap, strips the amine off any of twenty amino acids and hands the carbon skeleton into one of six points on the pathways above.
  5. Part VRun in reverse or sideways, the same junctions build fat (lipogenesis), build glucose (gluconeogenesis), make NADPH and nucleotide precursors (pentose phosphate pathway), and make ketone bodies — four branches off the pathways above.
  6. Part VIAll of it mapped to eight B vitamins, each gating a specific, nameable step — with links out to the full nutrient page for each.

Follow the arrows first. The names are secondary.

THE BIG PICTURE

Glucose and fat both converge on acetyl-CoA, which the Krebs cycle feeds to the electron transport chain Glucose flows through glycolysis to pyruvate, then through pyruvate dehydrogenase to acetyl-CoA. Fat flows through beta-oxidation, repeating, to the same acetyl-CoA. Amino acids feed into the Krebs cycle at several points via transamination. Acetyl-CoA enters the Krebs cycle, which produces NADH and FADH2 that the electron transport chain converts to ATP. Glucose Part I Pyruvate Fatty Acid Part II Acyl-CoA PDH repeats Acetyl-CoA Krebs cycle NADH FADH2 ETC → ATP Amino Acids Part IV
Part I

Glycolysis

Ten steps in the cytosol, no oxygen needed: five that spend ATP priming glucose for cleavage, then five — run twice, once per 3-carbon half — that repay it with a profit. Only one step is a redox reaction; it's the only place niacin is used directly in these ten steps.

I.1Investment, then payoff, twice over

Glycolysis from glucose to pyruvate Glucose is phosphorylated twice, spending two ATP, then split into two three-carbon units. Each unit is oxidized at GAPDH, reducing NAD+ to NADH, then converted to pyruvate while regenerating two ATP per unit. Glucose Hexokinase ATP ADP Glucose-6-P PFK-1 ATP ADP Fructose-1,6-BP Aldolase splits into two 3-carbon units G3P ×2 GAPDH NAD+ NADH 1,3-BPG PGK → Enolase → PK 2 ADP 2 ATP Pyruvate ×2

Net per glucose: 2 ATP, 2 NADH, and 2 pyruvate. The investment phase (hexokinase, PFK-1) spends 2 ATP; the payoff phase runs twice — once per triose — repaying that loan and netting 2 ATP plus 2 NADH.

I.2The bridge to the cycle: pyruvate dehydrogenase

Pyruvate isn't a Krebs cycle intermediate. One more step, pyruvate dehydrogenase (PDH), decarboxylates it to acetyl-CoA first — and that single enzyme pulls in five cofactors at once, more than any other step in this paper.

Pyruvate dehydrogenase converts pyruvate to acetyl-CoA using five cofactors at once Pyruvate plus thiamine pyrophosphate, lipoic acid, FAD, NAD+, and coenzyme A yields acetyl-CoA, CO2, and NADH. 5 COFACTORS AT ONCE TPP (B1) · Lipoic acid · FAD (B2) · NAD+ (B3) · CoA (B5) Pyruvate Pyruvate Dehydrogenase releases CO2, reduces NAD+ → NADH Acetyl-CoA → Krebs cycle, Part III

Same five cofactors, same reaction template, repeat almost exactly at α-ketoglutarate dehydrogenase inside the Krebs cycle (III.2) — thiamine status can throttle carbon flow into the cycle from two separate directions.

Part II

β-Oxidation

Fat is dismantled in the mitochondrial matrix by a four-step loop that repeats, not a line that runs once. A C16 chain like palmitate spins through it seven times, releasing eight acetyl-CoA and reducing seven FAD and seven NAD+ along the way.

II.1Four steps, repeated until nothing is left but acetyl-CoA

One turn of the beta-oxidation spiral Acyl-CoA dehydrogenase reduces FAD, enoyl-CoA hydratase adds water, 3-hydroxyacyl-CoA dehydrogenase reduces NAD+, and thiolase cleaves off acetyl-CoA using coenzyme A, shortening the chain by two carbons. The shortened chain re-enters at the top for the next turn. Fatty acid + CoA + ATP (activation) Acyl-CoA (Cₙ) trans-Δ²-Enoyl-CoA 3-Hydroxyacyl-CoA 3-Ketoacyl-CoA Acyl-CoA Dehydrogenase FAD → FADH2 Enoyl-CoA Hydratase + H2O 3-OH-Acyl-CoA DH NAD+ → NADH Thiolase + CoA-SH Acetyl-CoA → Krebs cycle repeats with Acyl-CoA (Cₙ₋₂)

Per turn: 1 FADH2, 1 NADH, 1 acetyl-CoA. A C16 chain (palmitate) spins through seven turns, releasing eight acetyl-CoA for the Krebs cycle below.

Part III

The Krebs Cycle

Every acetyl-CoA — from glucose or from fat — enters the same eight-step wheel. Oxaloacetate regenerates at the end of each turn, so the cycle never runs out of a place to put the next acetyl group.

III.1Eight steps, one wheel

The Krebs cycle Citrate, isocitrate, alpha-ketoglutarate, succinyl-CoA, succinate, fumarate, malate, and oxaloacetate arranged in a circle, with NAD+, FAD, and GTP cofactor changes labeled at each step. Acetyl-CoA Citrate Synthase Aconitase Isocitrate DH NAD+→NADH, CO2 α-KG Dehydrogenase NAD++CoA→NADH, CO2 5-cofactor complex, like PDH Succinyl-CoA Synthetase GDP+Pi→GTP Succinate DH FAD→FADH2 = ETC Complex II Fumarase + H2O Malate DH NAD+→NADH Citrate Isocitrate α-Ketoglutarate Succinyl-CoA Succinate Fumarate Malate Oxaloacetate

Per turn: 3 NADH, 1 FADH2, 1 GTP, 2 CO2. The two decarboxylation steps and the succinate dehydrogenase step double as a direct entry point into the electron transport chain.

III.2Two steps worth noticing

Where the cycle repeats itself — and where it plugs straight into the ETC

α-Ketoglutarate dehydrogenase is structurally almost identical to pyruvate dehydrogenase (I.2) and needs the same five cofactors — B1, lipoic acid, B2, B3, B5 — a second checkpoint where thiamine status alone can throttle the cycle. Succinate dehydrogenase is the only Krebs cycle enzyme embedded in the inner mitochondrial membrane: it's also Complex II of the electron transport chain, so this one step belongs to both pathways at once.

Part IV

Protein & Amino Acid Metabolism

Amino acids — from dietary protein, or from muscle during fasting or very-low-carb intake — can be stripped of their nitrogen and fed into the same pathways as carbon skeletons. Every one of the twenty shares the same entry mechanism.

IV.1Transamination: one cofactor swaps the amine for a keto group

An aminotransferase (ALT and AST are the two on a standard liver panel) transfers the amino acid's –NH₂ group onto α-ketoglutarate. The amino acid becomes a keto acid; α-ketoglutarate becomes glutamate. Same swap, all twenty times, always on the same cofactor.

Transamination moves an amine group from an amino acid to alpha-ketoglutarate An amino acid with an R group and an amine group loses the amine to become a keto acid. The amine group, carried by PLP, attaches to alpha-ketoglutarate, converting it to glutamate. Amino Acid R – CH(NH₂) – COOH α-Ketoglutarate –NH₂ carried by PLP (B6) Keto Acid R – CO – COOH Glutamate R = a different side chain for each of the 20 amino acids

Glutamate's amine is then released as free ammonia (glutamate dehydrogenase, NAD+/NADP+) and disposed of via the liver's urea cycle — an ATP-costly system separate from the ATP-generating pathways above.

IV.2Six doors, twenty amino acids

The keto acid lands at one of six points already drawn above: pyruvate, acetyl-CoA, or a Krebs cycle intermediate. Land at pyruvate or a Krebs intermediate and the carbon can run back to glucose (glucogenic, via gluconeogenesis). Leucine and lysine land only at acetyl-CoA, which can't run backward — purely ketogenic. A few amino acids split across two doors and count as both.

Amino acid entry points into glycolysis and the Krebs cycle Six docking points -- pyruvate, alpha-ketoglutarate, oxaloacetate, succinyl-CoA, fumarate, and acetyl-CoA -- each receiving carbon skeletons from a specific group of amino acids after transamination. Acetyl-CoA is marked as ketogenic only, with no path back to glucose; the other five are glucogenic. SIX DOORS INTO THE PATHWAYS ABOVE Ala · Ser · Gly · Cys Pyruvate Glu · Gln · His Pro · Arg α-Ketoglutarate Asp · Asn Oxaloacetate Ile · Met · Val · Thr Succinyl-CoA Phe · Tyr Fumarate Leu · Lys +Ile,Phe,Tyr,Trp (partly) Acetyl-CoA ketogenic only — no path back to glucose

Every arrow above is PLP-dependent (B6). Alanine, serine, glycine, and cysteine feed pyruvate; glutamate, glutamine, histidine, proline, and arginine feed α-ketoglutarate; aspartate and asparagine feed oxaloacetate; isoleucine, methionine, valine, and threonine feed succinyl-CoA; phenylalanine and tyrosine feed fumarate. Leucine and lysine are the only two amino acids that are purely ketogenic — every other amino acid can, at least partly, run back to glucose.

Part V

Where the Pathways Branch

Everything above runs toward ATP. The same junction molecules — citrate, pyruvate, glucose-6-phosphate, acetyl-CoA — also sit at the start of four branches that run the traffic somewhere else: building fat, building glucose, building nucleotide precursors, or building an alternative fuel for when glucose is scarce.

V.1Lipogenesis: citrate leaves the cycle to build fat

When acetyl-CoA and ATP are both abundant, citrate backs up and is exported out of the mitochondria instead of continuing around the Krebs wheel.

Lipogenesis: citrate exported to the cytosol is rebuilt into fatty acids Citrate crosses from the mitochondria to the cytosol, where ATP-citrate lyase splits it back into acetyl-CoA. Acetyl-CoA carboxylase, using biotin, converts it to malonyl-CoA, and fatty acid synthase, using NADPH, builds fatty acids and triglycerides. MITOCHONDRIA CYTOSOL Citrate shuttle Citrate ATP-Citrate Lyase Acetyl-CoA Acetyl-CoA Carboxylase Biotin (B7) Malonyl-CoA Fatty Acid Synthase NADPH (from Part V.3) Fatty Acids → Triglycerides

The rate-limiting step, acetyl-CoA carboxylase, needs biotin (B7) — the same vitamin that runs pyruvate carboxylase in gluconeogenesis below.

V.2Gluconeogenesis: running glycolysis backward, with three detours

Liver and kidney rebuild glucose from lactate, alanine, and glycerol. Seven of the ten glycolysis steps just run in reverse; three are one-way, so the body uses different enzymes to get around them.

Gluconeogenesis rebuilds glucose from pyruvate using three bypass enzymes Lactate, alanine, and glycerol feed into pyruvate, which pyruvate carboxylase and PEPCK convert to PEP, bypassing pyruvate kinase. PEP runs the reverse of glycolysis to fructose-1,6-bisphosphate, which fructose-1,6-bisphosphatase converts to fructose-6-phosphate, bypassing PFK-1. Glucose-6-phosphatase, liver and kidney only, produces free glucose, bypassing hexokinase. Lactate · Alanine Glycerol Pyruvate Pyruvate Carboxylase + PEPCK Biotin (B7) · bypass 1 PEP reverse of Part I steps Fructose-1,6-BP bypass 2 Fructose-6-P → Glucose (bypass 3, liver/kidney)

Sky-colored arrows are the same enzymes as Part I, run backward. Gold arrows are gluconeogenesis-only detours around the three irreversible glycolysis steps — the first needs biotin.

V.3The pentose phosphate pathway: G6P branches for NADPH

Before glucose-6-phosphate (G6P) commits to glycolysis, some of it can branch off to make NADPH — the electron donor lipogenesis and antioxidant defenses run on — and ribose-5-phosphate for nucleotide synthesis.

The pentose phosphate pathway branches off glucose-6-phosphate to make NADPH and ribose-5-phosphate Glucose-6-phosphate is oxidized to 6-phosphogluconate, reducing NADP+ to NADPH, then to ribulose-5-phosphate, reducing a second NADP+ and releasing CO2. Ribulose-5-phosphate converts to ribose-5-phosphate, used for nucleotide synthesis. Glucose-6-P G6PD NADP+ → NADPH 6-Phosphogluconate 6PGD NADP+ → NADPH, CO2 Ribulose-5-P Ribose-5-P → nucleotides (DNA / RNA)

2 NADPH per G6P, spent above in lipogenesis (V.1) and on glutathione regeneration. The pathway's non-oxidative half (not shown) can run in reverse and needs TPP (B1) — the same cofactor as pyruvate dehydrogenase.

V.4Ketogenesis: when acetyl-CoA outpaces oxaloacetate

Fasting or very-low-carb intake drops oxaloacetate (it's pulled toward gluconeogenesis), so the liver's mitochondria can't condense all the acetyl-CoA β-oxidation is producing. The surplus becomes ketone bodies instead.

Ketogenesis converts surplus acetyl-CoA into beta-hydroxybutyrate and acetone Two acetyl-CoA combine to acetoacetyl-CoA, then a third acetyl-CoA forms HMG-CoA, which splits into acetoacetate. Acetoacetate becomes either beta-hydroxybutyrate, using NADH, or acetone, spontaneously. 2 × Acetyl-CoA Thiolase Acetoacetyl-CoA +CoA HMG-CoA Acetoacetate +NADH β-Hydroxybutyrate spontaneous Acetone

β-Hydroxybutyrate travels to other tissues — brain included — as an alternative fuel to glucose. Acetone is mostly exhaled, which is where the fruity breath in ketosis or diabetic ketoacidosis comes from.

Part VI

The B-Vitamin Cofactors

Each B vitamin converts into an active coenzyme before it can do anything. A deficiency doesn't cause a generic drop in “energy” — it stalls a specific, nameable step. The individual nutrient pages carry deficiency signs, intake targets, and food sources for each.

VitaminCoenzymeActs inRole
B1 · Thiamine TPP I, III, V Pyruvate dehydrogenase (I.2) and α-ketoglutarate dehydrogenase (III.2) — both decarboxylation steps — plus the reversible half of the pentose phosphate pathway (V.3).
B2 · Riboflavin FAD / FMN II, III Electron acceptor for acyl-CoA dehydrogenase in every turn of β-oxidation and for succinate dehydrogenase in the Krebs cycle, which feeds those electrons directly into Complex II.
B3 · Niacin NAD+ / NADP+ I–V NAD+ runs GAPDH, β-oxidation, and three Krebs steps. Its cousin NADP+ is what the pentose phosphate pathway (V.3) reduces to NADPH — the electron donor lipogenesis (V.1) spends.
B5 · Pantothenic acid Coenzyme A I–V CoA is the handle every acyl group is carried by — acetyl-CoA, malonyl-CoA, HMG-CoA — through β-oxidation, the Krebs cycle, lipogenesis, and ketogenesis alike.
B6 · Pyridoxine PLP IV PLP runs every transamination in Part IV — swapping an amine for a keto group and handing twenty carbon skeletons into six points on the pathways above.
B7 · Biotin Carboxylase prosthetic group III, V Pyruvate carboxylase refills the Krebs cycle (III) and starts gluconeogenesis (V.2); acetyl-CoA carboxylase is the rate-limiting step of lipogenesis (V.1). Same vitamin, opposite directions.
Reading the map

TPP, CoA, FAD, and NAD+/NADP+ do almost all the direct chemistry. B6 and biotin sit at the branch points instead — B6 decides what protein becomes, biotin decides whether pyruvate builds glucose or acetyl-CoA builds fat. See Metabolic Health for how this fits into mitochondrial capacity and insulin sensitivity.

Glycolysisβ-oxidationKrebs cycleTCA cyclePyruvate dehydrogenaseTransaminationGlucogenic amino acidsKetogenic amino acidsLipogenesisGluconeogenesisPentose phosphate pathwayKetogenesisNADPHThiamineRiboflavinNiacinPantothenic acidPyridoxineBiotinMitochondrial capacityATP