Panel 131 Details of the 10 steps of glycolysis

For each step, the part of the molecule that undergoes a change is shadowed in blue, and the name of the enzyme that catalyzes the reaction is in a yellow box.

Step 1

Glucose is phosphorylated by ATP to form a sugar phosphate. The negative charge of the phosphate prevents passage of the sugar phosphate through the plasma membrane, trapping glucose inside the cell.

CH2OH O OH OH
glucose hexokinase

CH2O P O OH OH
glucose 6-phosphate

+
OH

ATP

+
OH

ADP

H+

HO

HO

A readily reversible 6 CH2O P rearrangement of 5 the chemical O structure (isomerization) 4 1 moves the OH carbonyl oxygen 2 HO OH 3 from carbon 1 to carbon 2, forming OH a ketose from an aldose sugar. (ring form) (See Panel 23, pp. 7071.)

Step 2

O C1 H HO H H C C C C
2

H
1CH2OH

OH H OH OH

phosphoglucose isomerase

C C C C

O H OH
4 P OH2C 5 6

HO H H

O HO
3

CH2OH
2

3 4

4 5

OH

OH

OH
(ring form)

CH2O P 6 (open-chain form)

CH2O P 6 (open-chain form)

fructose 6-phosphate

glucose 6-phosphate

Step 3

The new hydroxyl group on carbon 1 is phosphorylated by ATP, in preparation for the formation of two three-carbon sugar phosphates. The entry of sugars into glycolysis is controlled at this step, through regulation of the enzyme phosphofructokinase.

P OH2C

O HO

CH2OH

phosphofructokinase

P OH2C

O HO

CH2O P

+
OH

ATP

+
OH

ADP

H+

OH
fructose 6-phosphate

OH
fructose 1,6-bisphosphate

Step 4

The sixcarbon sugar is cleaved to produce two three-carbon molecules. Only the glyceraldehyde 3-phosphate can proceed immediately through glycolysis.

CH2O P C
P OH2C

CH2O P C
aldolase

O H OH OH HO

O H H O C H C OH

O HO

CH2O P OH

HO H H

C C C

C H

OH
(ring form)

CH2O P
(open-chain form) dihydroxyacetone phosphate

CH2O P
glyceraldehyde 3-phosphate

fructose 1,6-bisphosphate

Step 5

The other product of step 4, dihydroxyacetone phosphate, is isomerized to form glyceraldehyde 3-phosphate.

CH2OH C O

H
triose phosphate isomerase

O C

OH

CH2O P
dihydroxyacetone phosphate

CH2O P
glyceraldehyde 3-phosphate

Step 6

The two molecules of glyceraldehyde 3-phosphate are oxidized. The energygeneration phase of glycolysis begins, as NADH and a new high-energy anhydride linkage to phosphate are formed (see Figure 135).

O C H C

glyceraldehyde 3-phosphate dehydrogenase

O C H C

O P

+
OH

NAD+

Pi

+
OH

NADH

H+

CH2O P
glyceraldehyde 3-phosphate

CH2O P
1,3-bisphosphoglycerate

Step 7

The transfer to ADP of the highenergy phosphate group that was generated in step 6 forms ATP.

O C H C

O P

phosphoglycerate kinase

O C H C

+
OH

ADP

+
OH

ATP

CH2O P
1,3-bisphosphoglycerate

CH2O P
3-phosphoglycerate

Step 8

The remaining phosphate ester linkage in 3-phosphoglycerate, which has a relatively low free energy of hydrolysis, is moved from carbon 3 to carbon 2 to form 2phosphoglycerate.

O
1

O C C OH
phosphoglycerate mutase

O C H C

2 3

O P

CH2O P

CH2OH
2-phosphoglycerate

3-phosphoglycerate

The removal of water from 2-phosphoglycerate creates a high-energy enol phosphate linkage.

Step 9

O C H C

O
enolase

O C C

O P

O P

H2O

CH2OH
2-phosphoglycerate

CH2
phosphoenolpyruvate

Step 10 The transfer to ADP of the high-energy phosphate group that was generated in step 9 forms ATP, completing glycolysis.

O C C

O
pyruvate kinase

O C C

O P

ADP

ATP

CH2
phosphoenolpyruvate

CH3
pyruvate

NET RESULT OF GLYCOLYSIS

O C

CH2OH O HO OH OH
ATP ATP NADH ATP ATP NADH ATP ATP

C CH3 O C C CH3

OH

glucose

In addition to the pyruvate, the net products are two molecules of ATP and two molecules of NADH.

two molecules of pyruvate