Panel 13–1 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.

CH2OH CH2O P
Step 1
Glucose is
O hexokinase O
phosphorylated by ATP to
form a sugar phosphate.
The negative charge of the
OH
+ ATP
OH
+ ADP + H+
phosphate prevents HO OH HO OH
passage of the sugar
phosphate through the
plasma membrane, OH OH
trapping glucose inside glucose glucose 6-phosphate
the cell.

Step 2 O H
A readily
reversible
6 CH2O P
C1 1CH2OH
rearrangement of
the chemical 5 O H C OH C O
2 phosphoglucose 2
structure isomerase P OH2C CH2OH
(isomerization) 6 O 1
4 1 HO C H HO C H
moves the OH 3 3
5 HO 2
carbonyl oxygen HO 3 2 OH H C OH H C OH
from carbon 1 to 4 4 4 3 OH
carbon 2, forming
OH H C OH H C OH OH
a ketose from an 5 5
aldose sugar. (ring form)
(See Panel 2–3, (ring form)
6
CH2O P 6
CH2O P
pp. 70–71.) (open-chain form) (open-chain form)

glucose 6-phosphate fructose 6-phosphate

Step 3
The new hydroxyl
group on carbon 1 is P OH2C O CH2OH P OH2C O CH2O P
phosphofructokinase
phosphorylated by ATP, in
preparation for the formation HO + ATP HO + ADP + H+
of two three-carbon sugar OH OH
phosphates. The entry of
sugars into glycolysis is OH OH
controlled at this step, through fructose 6-phosphate fructose 1,6-bisphosphate
regulation of the enzyme
phosphofructokinase.

Step 4 CH2O P CH2O P

The six-
carbon sugar is
cleaved to produce C O C O
two three-carbon P OH2C O CH2O P
molecules. Only the HO C H aldolase HO C H H O
glyceraldehyde HO + C
3-phosphate can H C OH H
proceed immediately OH
H C OH
through glycolysis. OH H C OH
(ring form) CH2O P
CH2O P
(open-chain form)
dihydroxyacetone glyceraldehyde
fructose 1,6-bisphosphate phosphate 3-phosphate

Step 5
The other H O
product of step 4, CH2OH
triose phosphate isomerase C
dihydroxyacetone
phosphate, is C O
isomerized to form H C OH
glyceraldehyde CH2O P
3-phosphate. CH2O P
dihydroxyacetone glyceraldehyde
phosphate 3-phosphate
Step 6
The two molecules glyceraldehyde 3-phosphate
of glyceraldehyde 3-phosphate O H dehydrogenase O O P
are oxidized. The energy- C C
generation phase of glycolysis + NAD+ + Pi + NADH + H+
begins, as NADH and a new H C OH H C OH
high-energy anhydride linkage
to phosphate are formed (see CH2O P CH2O P
Figure 13–5).
glyceraldehyde 1,3-bisphosphoglycerate
3-phosphate

Step 7 O O P phosphoglycerate kinase O O–

The transfer
to ADP of the high- C C
energy phosphate + ADP + ATP
group that was H C OH H C OH
generated in step 6
forms ATP. CH2O P CH2O P
1,3-bisphosphoglycerate 3-phosphoglycerate

Step 8
The remaining O O– O O–
phosphate ester linkage in C phosphoglycerate mutase C
3-phosphoglycerate, 1
which has a relatively low
H C OH H C O P
free energy of hydrolysis, 2
is moved from carbon 3
to carbon 2 to form 2- 3
CH2O P CH2OH
phosphoglycerate. 3-phosphoglycerate 2-phosphoglycerate

Step 9
The removal of O O– O O–
water from 2-phosphoglycerate C enolase C
creates a high-energy enol
phosphate linkage.
H C O P C O P + H2O

CH2OH CH2
2-phosphoglycerate phosphoenolpyruvate

Step 10
The transfer to O O– O O–
ADP of the high-energy C pyruvate kinase C
phosphate group that was +
generated in step 9 forms + ADP + H + ATP
C O P C O
ATP, completing
glycolysis.
CH2 CH3
phosphoenolpyruvate pyruvate

NET RESULT OF GLYCOLYSIS O O–

C
CH2OH
O C O

NADH ATP ATP CH3

OH
HO OH
O O–
OH
C
ATP ATP NADH ATP ATP
C O

CH3
In addition to the pyruvate, the net products are two molecules
glucose two molecules of ATP and two molecules of NADH. of pyruvate