Map · Energy metabolism · Glycolysis

Glycolysis glucose → pyruvate · 10 enzymes + PFKFB3

Cytosol. Real crystal coordinates (RCSB PDB). PFKFB3 sets F2,6BP, the allosteric activator of PFK-1. Pyruvate then branches to LDH (fermentation) or PDH (acetyl-CoA).

★ Master Switch of Glycolysis — available now

REGULATOR

PFKFB3 (iPFK-2 / FBPase-2)

F6P + ATP → F2,6BP + ADP + H⁺  |  F2,6BP + H₂O → F6P + Pᵢ

Bifunctional kinase + phosphatase that sets the level of F2,6BP, the most potent allosteric activator of PFK-1. Two active sites, two separate mechanisms.

EC 2.7.1.105 / EC 3.1.3.46 · PDB 2AXN·2DWP·2I1V·3QPV·3QPW

▶ OPEN SIMULATOR

Why it matters: PFKFB3 controls glycolytic flux by making and breaking fructose-2,6-bisphosphate. Overexpressed in cancer cells (Warburg effect); a major drug target.

The 10 Glycolytic Steps

GLUCOSE
①
STEP 1

Hexokinase I HK1 · not glucokinase (GCK)

Glucose + ATP → G6P + ADP

Induced-fit domain closure toward glucose (drawn — bound alone, glucose doesn't seal the C-terminal cleft); Mg²⁺-assisted in-line phosphoryl transfer; conserved Asp657 proposed as catalytic base.

EC 2.7.1.1 · human HK1

▶ OPEN SIMULATOR
→
G6P
STEP 2

Phosphoglucose isomerase (PGI)

G6P ⇌ F6P

Aldose-ketose isomerization: ring opening, base-catalyzed proton abstraction via cis-enediol(ate) intermediate, ring closure.

EC 5.3.1.9

▶ OPEN SIMULATOR
→
F6P
STEP 3

Phosphofructokinase-1 (PFK-1)

F6P + ATP → F1,6BP + ADP

Rate-limiting committed step. Allosterically activated by AMP & F2,6BP, inhibited by ATP/citrate. In-line phosphoryl transfer onto the 1-OH.

EC 2.7.1.11

▶ OPEN SIMULATOR
→
F1,6BP
STEP 4

Aldolase (class I)

F1,6BP ⇌ DHAP + G3P

Retro-aldol cleavage through a Schiff-base (protonated imine) between Lys and C2 of the sugar; stabilized carbanion intermediate.

EC 4.1.2.13

▶ OPEN SIMULATOR
→
DHAP + G3P
STEP 5

Triosephosphate isomerase (TPI)

DHAP ⇌ G3P

The perfect enzyme: kcat/KM near the diffusion limit in GAP→DHAP. Glu165 abstracts the pro-R proton from the CH₂OH end of DHAP to a cis-enediolate stabilized by His95/Lys12, then reprotonates C2 to give D-G3P; the loop-6 "lid" seals the pocket.

EC 5.3.1.1

▶ OPEN SIMULATOR
STEP 6

GAPDH (oxidation → acyl phosphate)

G3P + Pᵢ + NAD⁺ → 1,3-BPG + NADH

Cys152 thiolate attacks the aldehyde → thiohemiacetal; hydride transfer to NAD⁺; phosphorolysis of the thioester → acyl phosphate.

EC 1.2.1.12

▶ OPEN SIMULATOR
→
1,3-BPG
STEP 7

Phosphoglycerate kinase (PGK)

1,3-BPG + ADP → 3PG + ATP

First ATP-producing step (substrate-level phosphorylation). Two domains hinge closed (arc ≈32° [schematic]; frame 2XE7 is semi-closed); the TS is neutralized by a basic patch (Arg38 · Lys215 · Lys219) and Mg²⁺, while Arg122/Arg65/Arg170 + His62 clamp the 3-phosphate.

EC 2.7.2.3

▶ OPEN SIMULATOR
→
3PG
STEP 8

Phosphoglycerate mutase (dimeric PGAM)

3PG ⇌ 2PG

Phosphohistidine (His11) shuttle: enzyme phosphorylates the substrate, then reclaims the phosphate at the adjacent position — a Ping-Pong swap.

EC 5.4.2.11

▶ OPEN SIMULATOR
→
2PG
↓
PEP
←Lactate
↗pyruvate
→pyruvate
→Acetyl-CoA
⤷ PYRUVATE branches → PDH → Acetyl-CoA → TCA cycle (aerobic)  |  LDH → Lactate (fermentation · Cori cycle)  |  PC → gluconeogenesis (next)