at The End of This Lecture, Student Will Be Able To

Lecture No.
9
Glycolysis
• At the end of this lecture, student will be able to
–Explain the energetics of glycolysis

–Describe the role of different enzymes in glycolysis
–Describe the importance of by products in glycolysis
1
Faculty of Pharmacy ©M. S. Ramaiah University of Applied Sciences
2
Bioenergetics
glucose + 2 NAD+ + 2 ADP + 2 Pi 

2 pyruvate + 2 NADH + 2 ATP
•Hexokinase ------------------------ ATP
•Phosphofructokinase ----------- ATP
•G-3-P Dehydrogenase ---------- NADH ----- 3 ATP
• Phosphoglycerate Kinase----------------------- ATP
•Pyruvate Kinase ----------------------------------- ATP
3
Balance sheet for ~P bonds of ATP:
 2 ATP expended
 4 ATP produced (2 from each of two 3C fragments from
glucose)
 Net production of 2 ~P bonds of ATP per glucose.
Glycolysis - total pathway, omitting H+:
In aerobic organisms:
 pyruvate produced in Glycolysis is oxidized to CO2 via Krebs
Cycle
 NADH produced in Glycolysis & Krebs Cycle is reoxidized via
the respiratory chain, with production of much additional
ATP.
4
G ly c e ra ld e h y d e -3 -p h o s p h a te
D e h y d ro g e n a se
2
H O + H+ O O PO 3
+
1 C NAD NADH 1 C
Fermentation: + P i
H C OH H C OH
2 2
2 2
Anaerobic 3 CH 2O PO 3 3 CH 2O PO 3
organisms lack a g ly c e ra ld e h y d e - 1 ,3 -b is p h o s p h o -
respiratory chain. 3 -p h o s p h a te g ly c e ra te
They must reoxidize NADH produced in Glycolysis

through some other reaction, because NAD+ is needed for
the Glyceraldehyde-3-phosphate Dehydrogenase reaction.
Usually NADH is reoxidized as pyruvate is converted to
a more reduced compound.
The complete pathway, including Glycolysis and the
reoxidation of NADH, is called fermentation. 5
Faculty
Faculty of Pharmacy
of Pharmacy ©M. S.©M. S. Ramaiah
Ramaiah University
University of Applied
of Applied Sciences
Sciences
L a c ta te D e h y d ro g e n a s e
O O O O
C NADH + H+ NAD+ C
C O HC OH
CH 3 CH 3
p y ru v a te la c ta te
E.g., Lactate Dehydrogenase catalyzes reduction of

the keto in pyruvate to a hydroxyl, yielding lactate, as
NADH is oxidized to NAD+.
Lactate, in addition to being an end-product of
fermentation, serves as a mobile form of nutrient energy,
Faculty of Pharmacy
6
Faculty of Pharmacy ©M. S.©M. S. Ramaiah
Ramaiah University
of Applied Sciences
Sciences
L a c ta te D e h y d ro g e n a s e
 O 
O O O
C NADH + H+ NAD+ C
C O HC OH
CH 3 CH 3
p y ru v a te la c ta te
Skeletal muscles ferment glucose to lactate during

exercise, when the exertion is brief and intense.
Lactate released to the blood may be taken up by other
tissues, or by skeletal muscle after exercise,
Faculty of Pharmacy
7
Faculty of Pharmacy ©M. S.©M. S. Ramaiah
Ramaiah University
of Applied Sciences
Sciences
Lactic Acidosis
• Normal value- 4-15 mg/ dl

• Increased in exercise, Shock, respiratory disease,
cancer, low pyruvate dehydrogenase activity, Von
Gierke`s Disease
• Oxygen debt
8
P y ru v a te A lc o h o l
D e c a rb o x y la s e D e h y d ro g e n a se
O 
O
CO H NADH + H+ NAD +
H
C 2 O
C O C H C OH
CH CH 3 CH 3
3
p y ru v a te a c e ta ld e h y d e e th a n o l
Some anaerobic organisms metabolize pyruvate to

ethanol, which is excreted as a waste product.
NADH is converted to NAD+ in the reaction
catalyzed by Alcohol Dehydrogenase.
9
Glycolysis, omitting H+:
Fermentation, from glucose to lactate:
glucose + 2 ADP + 2 Pi  2 lactate + 2 ATP
Anaerobic catabolism of glucose yields only 2 “high

energy” bonds of ATP.
10
Go' G
Glycolysis Enzyme/Reaction kJ/mol kJ/mol
Hexokinase -20.9 -27.2
Phosphoglucose Isomerase +2.2 -1.4
Phosphofructokinase -17.2 -25.9
Aldolase +22.8 -5.9
Triosephosphate Isomerase +7.9 negative
Glyceraldehyde-3-P Dehydrogenase -16.7 -1.1
& Phosphoglycerate Kinase
Phosphoglycerate Mutase +4.7 -0.6
Enolase -3.2 -2.4
Pyruvate Kinase -23.0 -13.9
*Values in this table from D. Voet & J. G. Voet (2004) Biochemistry, 3 rd Edition, John
Wiley & Sons, New York, p. 613.
Faculty of Pharmacy
11
©M. S. Ramaiah University of Applied Sciences
Flux through the Glycolysis pathway is regulated by control
of 3 enzymes that catalyze spontaneous reactions:
Hexokinase, Phosphofructokinase & Pyruvate Kinase.
Hexokinase ----- inhibited by G6P
PFK---------------- inhibited by ATP, Citrate, H+
12
Pasteur effect
• In 1861 Louis Pasteur figured out, that in the

absence of oxygen, yeast consumes more glucose
than in the presence of oxygen. A phenomenon
that has become known as Pasteur effect in the
literature.
• Inhibition of glycolysis by oxygen
• Because of inhibition of PFK
13
Crabtree effect
• Inhibition of oxygen consumption by adding glucose

• Increasing concentrations of glucose accelerates
glycolysis
• Production of appreciable amounts of ATP through
substrate-level phosphorylation.
• This reduces the need of oxidative phosphorylation
done by the TCA cycle via the electron transport chain
• Thus decreases oxygen consumption
14
Summary
 Pyruvate produced in Glycolysis is oxidized to CO2 via Krebs
Cycle
 NADH produced in Glycolysis & Krebs Cycle is reoxidized via
the respiratory chain, with production of much additional
ATP
 Skeletal muscles ferment glucose to lactate during
exercise, when the exertion is brief and intense
 Inhibition of the Glycolysis enzyme
Phosphofructokinase when [ATP] is high prevents
breakdown of glucose in a pathway whose main role is
to make ATP.
15
Conversion of Pyruvate to Acetyl CoA
• Pyruvate dehydrogenase complex

• Require 5 cofactor –
1. TPP
2. Lipoamide
3. FAD
4. CoA
5. NAD+
16
17
Regulation
18

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Lecture No.

–Explain the energetics of glycolysis

glucose + 2 NAD+ + 2 ADP + 2 Pi 

They must reoxidize NADH produced in Glycolysis

E.g., Lactate Dehydrogenase catalyzes reduction of

Skeletal muscles ferment glucose to lactate during

• Normal value- 4-15 mg/ dl

Some anaerobic organisms metabolize pyruvate to

Anaerobic catabolism of glucose yields only 2 “high

• In 1861 Louis Pasteur figured out, that in the

• Inhibition of oxygen consumption by adding glucose

• Pyruvate dehydrogenase complex

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