Glycolysis and Oxidative Pentose Phosphate Pathway

Lesson Prepared Under MHRD project “National

Mission on Education Through ICT”

Discipline: Botany

Paper: Plant Metabolism

National Coordinator: Prof. S.C. Bhatla

Lesson: Glycolysis and Oxidative Pentose Phosphate

Pathway

Lesson Developer: Dr Manju A. Lal

Department/College: Kirori Mal College

Lesson Reviewer: Dr Girish Mishra

Department/College: Botany, University of Delhi

Language Editor: Vinee Khanna

Department/College: Department of Genetics, University of

Delhi South Campus

Lesson Editor: Dr Rama Sisodia, Fellow in Botany ILLL

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 Glycolysis and Oxidative Pentose Phosphate Pathway

Learning Outcomes

After reading the lesson the reader should be able to elucidate the:

 Important contributions of scientists, who have helped in elucidating the

glycolytic pathway.
 The biochemical reactions that occur during glycolysis.
 Regulation of glycolytic pathway.
 Fermentation.
 Oxidative Pentose Phosphate pathway (Hexose Monophosphate Shunt).

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 Glycolysis and Oxidative Pentose Phosphate Pathway

Table of Contents

Chapter: Glycolysis and Oxidative Pentose Phosphate

Pathway

 Introduction
 Overview of respiration

 Glycolysis
 Important contributions of the scientists which led to
elucidation of glycolytic pathway
 Glycolytic pathway
 Preparatory phase
 Payoff phase
 Regulation of glycolysis
 Why are glycolytic intermediates phosphorylated
 Unique features of plant glycolysis
 Reoxidation of cytosolic NADH
 Fate of pyruvate
 Fermentation
 Pyruvate metabolism in roots growing under anorexia
conditions
 Pyruvate metabolism in muscles
 Significance of fermentation

 Oxidative pentose phosphate pathway

 Pathway
 Significance of the pathway
 Regulation of oxidative pentose phosphate pathway

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Introduction

Any living organism has the capacity to generate energy from the complex
organic compounds. This process is called respiration. The metabolic energy,
which is released during the process, is conserved in the form of ATP. ATP
molecules are energy currency of the cell and it is in this form, the energy is
utilized for various biosynthetic reactions of the cell, and also for various cellular
activities. This includes cell division, cell elongation, ion transport across cell
membrane, cytoplasmic movements and cytoplasmic transport. In other words
most of the activities, which occur in the cell, require ATP. So, it becomes
essential to study the biochemistry of respiration so as to understand how is this
ATP generated during the processes?

Besides ATP generation, intermediates of the respiratory pathway serve as the

precursors for the biosynthesis of various bio molecules in the cell.

Respiration requires uptake of O2 (in aerobic respiration) for the breakdown of

carbon compounds with the simultaneous release of CO2. The most common form
of carbon compounds translocated in the plants is sucrose. It is translocated in
the sieve tubes of the phloem tissues from source to sink. Source is the site
where sucrose is being generated while sinks are the sites where sucrose is being
catabolized. The reaction for aerobic degradation of sucrose can be written as:

C12H22O11 + 12 O2 --------------- 12 CO2 + 11 H2O

The reaction is a coupled redox reaction where sucrose is completely oxidized to

CO2and O2 is reduced to water.

The reaction is exergonic reaction. Standard free energy change during the
reaction is, Δ Go’ = -5760 kJ mole-1 of sucrose.

In a green plant, both the processes i.e., photosynthesis and respiration are
occurring simultaneously in presence of sunlight. Photosynthetic process is
limited only to green parts of the plant, which are exposed to sunlight, while
respiration occurs in all parts of the plant and also is independent of the presence
of sunlight. It has been reported that in some herbaceous plants, 30 – 60 % of
the photosynthetic products are lost due to respiration.

Respiration can be i) maintenance respiration or ii) growth respiration.

Maintenance respiration is the respiration rate, which is required to maintain the
cellular activities, which the organism requires for its sustenance, but growth will

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 Glycolysis and Oxidative Pentose Phosphate Pathway

not take place. However, new tissues will be formed at the expense of energy
produced during growth respiration. In growth respiration energy released during
respiration is much more than actually required for only maintenance of the
organism.

Respiration rate can be measured at the organism level, which means the
exchange of gases in between the organism and the environment. It is called
organismal respiration. The biochemical activity, which occurs at the cellular level
and makes it possible for organismal respiration to happen, is called cellular
respiration. We will be studying about cellular respiration.

Overview of respiration

Respiration is an energy releasing multistep process. If sufficient energy is

released during a reaction, it can be conserved in the form of ATP. Respiration
may occur in presence or in absence of O 2. Generally the plants are growing in
presence of atmospheric O2 (21%). This amount of O2 is sufficient to carry out
the aerobic respiration without showing any limiting effect of O 2 on the rate of
respiration. So, plants normally do not grow in deficiency of O 2. However, in the
water logged soil conditions or the roots growing deep in the soil, oxygen supply
may become limiting, creating anorexia (the absence of O2) or hypoxia
(abnormally low O2) conditions for the plant roots. In such case either the plant
develops aerenchyma, a plant cell death controlled development in roots, to
ensure O2 supply, or anaerobic respiration may occur in the plant roots.

In order to understand the respiration process we will be studying both aerobic

and anaerobic respiration in plants.

To understand the respiration process, we will be studying the process under the
following heads:

1. In the first lesson we will be studying glucose metabolism, which occurs in

cytoplasm. This includes the study of glycolytic reactions (conversion of
glucose to pyruvate), fermentation (conversion of pyruvate to either lactic
acid or alcohol) and oxidative Hexose Monophosphate Shunt.
2. The second lesson will include pyruvate metabolism in presence of O 2,
which will include, oxidative decarboxylation of pyruvate and Tri Carboxylic
Acid cycle, which occurs inside the mitochondrial matrix.
3. The third lesson will include oxidation of the reduced coenzymes (e.g.,
NADH, FADH2), which are produced during TCA, by the electron transport
chain localized in the inner mitochondrial membrane. It is during this phase

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O2 is consumed. Besides this, you will also be studying cyanide resistant

respiration, which is characteristic of the plant cells.
4. In the fourth lesson you will be studying about ATP synthesis.

Figure: Overview of the pathways involved in respiration

Source: Author

Glycolysis and Hexose Monophosphate Shunt

Some of the important contributions of the scientists, which have led to the
elucidation of glycolytic pathway:

 In 1860, Louis Pasteur (French chemist and microbiologist) observed that

the vital force of the living cells (yeast cells), was required to carry out
fermentation process.
 In 1897, Hans Buchner and Eduard Buchner (German biologist brothers)
prepared cell free extract of yeast for therapeutic purposes. In order to
preserve the extract they added sucrose, which was a common preservative
used at that time. They observed that alcoholic fermentation occurred, even
without any intact living yeast cell being present. So, it was an accidental
discovery by which it could be demonstrated that fermentation took place
even without the presence of any living cell. This observation contradicted

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the postulate of Louis Pasteur that fermentation required the vital force of
the cell.
 In 1905, Arthur Harden & W.J.Young demonstrated and estimated
quantitatively the CO2, which was produced when yeast extract was added
to glucose solution. They observed that CO2 production declined significantly
after some time of fermentation, however, it resumed after inorganic
phosphate was added to the medium. They observed that the inorganic
phosphate disappeared from the medium because it was incorporated as
the sugar phosphate, as fructose 1,6 di-phosphate (presently known as
fructose 1,6 biphosphate). Harden and young observed that if fructose 1,6
diphosphate was added to the medium, ethanol and carbon-di-oxide were
produced quickly. They concluded that fructose 1,6 di-phospate was the
possible intermediate in the reaction.
 At about the same time, Robinson observed the presence of second sugar,
glucose-6-phosphate in the medium, which was present along with fructose
1, 6 diphosphate in the ratio of 3:1.
 Harden & Young further observed that the dialyzed yeast extract was not
able to carry out fermentation process. When yeast extract was heated to
50◦C or more than that, it lost its capacity to ferment. However, this activity
was restored when the inactive dialyzed fraction of the yeast extract was
mixed with the heated fraction. So, they concluded that the activity of the
yeast extract was depending upon the presence of two kinds of factors
present in the extract, i.e., one fraction was heat labile and non dialyzable
(this fraction was called zymase), while the other fraction, which was heat
stable but was lost due to dialysis (this fraction was called cozymase).
Cozymase was consisting of metal ions, ATP, ADP and other coenzymes
such as NAD+.
 Further interesting observations were made using inhibitors:
Embden used iodoacetate, a known inhibitor of enzymes containing
sulfahydryl groups. Addition of iodoacetate to yeast extract led to
accumulation of fructose 1,6 biphosphate. Embden postulated the cleavage
of fructose 1,6 biphosphate. The enzyme aldolase was later on isolated by
another biochemist Otto Meyerhoff. Embden also demonstrated that by
adding another inhibitor fluoride, 3-phosphoglycerate and 2-
phosphoglycerate accumulated. By these observations it was clear that
conversion of glucose to alcohol required number of steps, each one being
catalyzed by specific enzymes.

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So, you can comprehend that by the use of following approaches, it was possible
to elucidate the complete pathway for glucose breakdown to alcohol:

1. By substituting glucose with other possible intermediates.

2. By demonstrating the requirement of cofactors by removing them by dialysis
3. By using inhibitors of various enzymes of possible metabolic pathway, which
led to accumulation of the substrates of those enzymes.

The pathway was named as EMP pathway after the names of the three
scientists, Gustav Embden, Otto Meyerhoff (Germany), and Jacob Parnus,
(Poland) whose work had helped in elucidating the pathway. Most of the
research occurred during the period of 1890-1930.

Glycolysis

The term Glycolysis is derived from Greek word, glykos-“sweet” or “sugar” and
lysis- means “splitting”. This is the process in which series of chemical reactions
takes place to convert a molecule of glucose to 2 molecules of pyruvate.

C6H12O6 + 2 NAD+ + 2 ADP + 2 Pi ------ 2 CH3COCOOH + 2 NADH + 2 ATP

This pathway operates in the cytosol of the cell however, in plants it operates in
the plastids also. Glycolysis is the sole source of energy in the cells respiring in
absence of oxygen e.g., erythrocytes in the mammals, in some of the bacteria,
some plant cells, which are modified to store starch and derive most of their
energy from glycolysis.

Glucose is broken down in ten steps and ten enzymes catalyze the reactions.

The glycolytic pathway consists of two phases:

1. Preparatory phase: The first five reactions of the glycolysis constitute

preparatory phase. During this phase two phosphorylation reactions take
place, one isomerization reaction and one cleavage of the six carbon sugar
to two molecules of three carbon sugars occurs. During preparatory phase
two ATP molecules are consumed coupled to phosphorylation reactions.
2. Payoff phase: During this phase oxidation reaction will result in reduction
of 2 molecules of NAD+ to NADH. Two molecules of pyruvate and four
molecules of ATP are generated.

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1) Preparatory phase:
The starting compound is glucose. The glucose may be derived from
sucrose by the action of invertase or sucrose synthase, or from starch by
the action of the enzyme starch synthase. Sucrose may have been
translocated from the plastids facilitated by the translocators present in
the plastid membrane. Glucose is metabolized by the following reactions:

Figure: The reactions of the preparatory phase: two phosphorylation, (1 & 3) one
isomerization (2) and one cleavage of 6 carbon compound to 2 molecules of 3-
carbon sugars. (4)
Source: Author, ILLL in-house

Step 1: Phosphorylation of glucose: glucose is phosphorylated and

Glucose-6-phosphate is produced. One molecule of ATP is used during the
reaction. The reaction is catalyzed by the enzyme hexokinase.

Hexokinase, Mg2+
Glucose + ATP-------------------- glucose-6-phosphate + ADP
ΔG0’ = - 16.7 kJ mole-1

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Phosphorylation of glucose is endergonic reaction

Glucose + Pi ------------- Glucose-6-phosphate + H2O ΔGo’ = 13.8 kJmol-
1
(A)

ATP hydrolysis is endergonic reaction

ATP + H2O ---------- ADP + Pi Δ Go’ = -30.5 kJ mol-1 (B)

Reaction A and B are coupled together so that net change in energy during the
reaction will be:
Δ Go’ = 13.8 + (-30.5) kJ mol-1 = - 16.7 kJ mol-1
Hexokinase is a kinase enzyme, which transfers phosphate group from terminal
end of ATP to a hexose sugar including that of glucose. The true substrate of a
kinase enzyme is not the ATP rather Mg ATP complex, because Mg2+ shields the
two phosphate groups of the ATP. The terminal phosphate group is acted upon
by the enzyme and is transferred to hexose sugar.

Another enzyme, which phosphorylates glucose to Glucose-6-phosphate is

Glucokinase. Glucokinase differs from hexokinase in its affinity for glucose.
Glucokinase has less affinity for glucose than hexokinase. Km value of hexokinase
for glucose is less than that of Glucokinase. Role of Glucokinase is important in
utilizing glucose for storage purposes rather than mobilizing for respiration.

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Step 2: Glucose-6-phosphate is converted to fructose-6-phosphate: The

reaction is reversible and is catalyzed by the enzyme
phosphohexoisomerase (phosphoglucoisomerase). Glucose is an aldose
sugar while fructose is a ketose. C-1 of the aldehydic group of the glucose
is reduced to alcoholic group while the C2 is oxidized to ketonic form. So,
there is no net oxidation or reduction during the reaction. There is no net
change in free energy and the reaction can go in either direction.

Step 3: Fructose-6-phosphate is further phosphorylated to fructose1, 6-

bisphosphate via enzymatic reaction carried by phosphofructokinase. This
reaction requires another molecule of ATP. Since the reaction is
endergonic, it is coupled to another exergonic reaction of hydrolysis of
ATP. This reaction is a committed pathway for the glycolysis. Once the
sugars are converted to fructose 1,6 bisphosphate it is ensured that the
sugars will enter the glycolytic pathway. Prior to this reaction glucose-6-
phosphate or fructose-6-phosphate may have other roles to play in the
cell, but once fructose 1,6-biphosphate has been produced it is sure to
enter the glycolytic pathway. It is one of the regulatory steps of the
glycolysis, which we will study under the regulation of glycolysis.

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Phosphofructokinase is allosterically regulated enzyme. Allosteric modulators are

ATP, ADP, AMP, and Phosphoenolpyruvate. ATP and PEP being the negative
modulator while ADP, AMP are the positive modulators. ATP binds to two places
of the enzyme molecule. One at the active site along with another substrate,
i.e., fructose-6-Phosphate. Another site is the regulatory site. Binding of ATP at
the regulatory site as a negative modulator results in change of the
conformation of the enzyme molecule. As a result there is lowering of affinity of
the enzyme for the substrate. Citrate also inhibits the activity.

Figure: Structure of phosphofructokinase

Source: http://biomoocnews.blogspot.in/2012/09/daily-newsletter-
september-27-2012.html
Compound in which two phosphate groups are attached to two different groups
of the molecule is called bi-phosphate rather calling them di-phosphate.
Compounds in which two phosphate groups are attached to the same group are
called di-phosphates.

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Step 4. Cleavage of fructose 1, 6 biphosphate: fructose 1,6 biphosphate is

cleaved to two molecules of triose phosphates, i.e., glyceraldehyde-3-
phosphate and dihydroxyacetone phosphate. This is a condensation
reaction and is catalyzed by enzyme fructose 1,6 bi phosphate aldolase,
which is also simply known as aldolase. The free energy change is small
and the reaction is reversible. The basic side chain of the lysine residue,
which is present at the active site of the enzyme, plays a key role in the
condensation reaction.

Step 5. Out of the two molecules of triose phosphates, which are

produced in the above reaction, glyceraldehyde-3-phosphate is on the
main pathway for further glycolytic reactions. Dihydroxyacetone
phosphate is converted to another molecule of glyceraldehyde-3-
phosphate after it has been metabolized. As a result of this, 2 molecules of
glyceraldehyde-3-phosphate will be produced from one molecule of
fructose1,6 biphosphate. The enzyme which catalyzes this interconversion
of the two triose phosphates is called triose phosphate isomers.

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2) Payoff phase of glycolysis

The second phase of glycolysis is also called payoff phase, since ATP
molecules are produced during this phase. In the earlier phase, which you
have studied, 2 ATP molecules were used for conversion of one molecule
of glucose to one molecule of fructose 1,6 bisphosphate. During this payoff
phase, the two molecules of triose phosphates, which have been
generated from one molecule of glucose, are converted to two molecules
of pyruvate in a series of reactions. In the following steps you are going to
study the conversion of one triose phosphate to one molecule of pyruvate.
However, similar reactions will result in conversion of triose phosphate to
another molecule of pyruvate.

Figure: The pay off phase of glycolysis, during which the two molecules of triose
phosphates are converted to two molecules of pyruvate.
Source: Author

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Step 6. Oxidative step: During this step glyceraldehyde-3-phosphate is

converted to 1,3-biphosphoglycerate.

This reaction involves two kinds of reactions,

i. An oxidative reaction in which electron transfer take place from

glyceraldehyde-3-phosphate to NAD+

Glyceraldehyde-3-phospate + H2O + NAD+ -- 3-phosphoglycerate + NADH + H+

RCHO + H2O + NAD+ --------- RCOOH + NADH + 2H+

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During reduction NAD+ can accept two electrons but only one proton is
covalently bound to NAD+. Other proton is present free in the solution as
hydrogen ion. So, NAD+ reduction is written as:

NAD+ + 2e + 2H+ --------- NADH + H+

Source: Author, ILLL in-house

ii. Phosphorylation reaction, in which one inorganic phosphate is
added to 3-phosphoglycerate.

3-phosphoglycerate + Pi ---- 1,3-biphosphoglycerate + H2O

Glyceraldehyde-3-phosphate dehydrogenase catalyzes this reaction.

Change in standard free energy is as follows:

ΔGo’ overall = ΔGo’ oxidation + ΔGo’ phosphorylation

= (-43.1 kJ mol-1) + ( 49.3 kJ mol-1)
= 6.2 kJ mol-1 ( 1.5 kcal mol-1

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This change in standard free energy is calculated for one mole of

glyceraldehyde-3-phosphate. Since one molecule of glucose produces 2
molecules of glyraldehyde-3-phosphate, standard free energy change for
one mole of glucose will be 12.4 kJ mole-1. The ΔGo’ value under cellular
conditions is slightly negative (-1.29 kJ mole-1).

The compound 1, 3-biphosphoglycerate is a high energy compound, since

it has a high phosphate group transfer potential. The carboxylic group is
not free but is anhydride with phosphoric acid. This type of anhydride is
called acyl phosphate which has a very high standard free energy of
hydrolysis, i.e., G0’ = -49.3kJ mole-1. The reaction occurs at the cysteine
residue present at the active site of the enzyme Glyceraldehyde-3-
phosphate dehydrogenase. Since heavy metals such as Hg2 + forms
irreversible covalent bond with the –SH group of cysteine, the enzyme
activity is inhibited.

Step 7. This is the reaction of glycolysis in which ATP is produced. One

phosphate group (the one which is attached to C-1 of the molecule) from
1,3-biphosphoglycerate is transferred to ADP molecule resulting in the
formation of ATP and 3-phosphoglycerate. The reaction is catalyzed by
phosphoglycerate kinase. ATP synthesis is an endergonic reaction and
requires 30.5 kJmole-1, while hydrolysis of 1,3-biphosphoglycerate is an
exergonic reaction in which energy released is -49.3kJ mole-1. The energy
released during the exergonic reaction is more than the one required for
the synthesis of ATP molecule. The two reactions are coupled together and
o ’
the net change in standard free energy is Δ G -18.8 kJ mole-1. ATP
formation, in such a way which occurs without the electron transfer, is
called substrate level phosphorylation.

1,3-biphosphoglycerate + ADP ------ 3-phosphoglycerate + ATP

ΔGo’ = - 18.8 kJ mole-1

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Since one glucose molecule produces two molecules of 1.3-

biphosphoglycerate, 2 ATP molecules will be generated during this
reaction. In the initial preparatory phase 2 ATP molecules were consumed
for the production of fructose 1,6 biphosphate, so the net difference in
number of ATP molecules required or spent till this reaction becomes zero.

Step 8. In this reaction, 3-phosphoglycerate is converted to 2-

phosphoglycerate, by the change in the position of a phosphate group
from C -3 to C-2 of the molecule. The reaction is catalyzed by the enzyme
phosphoglycerate mutase.

Amino acid Histidine is present on the active site of the enzyme

phosphoglycerate mutase. This histidine is initially phosphorylated by 2,3-
biphosphoglycerate (2,3-biphosphoglycerate is an activator of the enzyme).
During the reaction catalyzed by the enzyme, in the first step phosphate group
from the phosphorylated histidine is transferred to –OH group attached to C-2 of
3-phosphoglycerate resulting in the formation of intermediate 2,3-
phosphoglycerate. In the subsequent step phosphoryl group from C-3 of the
molecule is transferred to histidine residue of the enzyme regenerating the
phosphorylated enzyme.

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Step 9. Dehydration of 2-phosphoglycerate: 2-phosphoglycerate has a

low phosphoryl group transfer potential (ΔGo’ value of hydrolysis of this
compound is -17.6 kJ/mole-1). Dehydration of this compound generates
phosphoenolpyruvate, which has a high phosphoryl group transfer
potential, with a standard free energy change of -61.9 kJ/mole-1. Enolase
catalyzes this reaction. The enzyme requires Mg2+. Mg2+complexes with
the carboxyl group of 2-phosphoglycerate facilitating reaction with the
enzyme enolase.

Step 10. Synthesis of pyruvate from phosphoenolpyruvte: This is the

second reaction in the glycolytic pathway, in which ATP is synthesized,
when the phosphoryl group is transferred from PEP to ADP. The reaction is
catalyzed by the enzyme pyruvate kinase. This is another example of
substrate level phosphorylation. Hydrolysis of PEP is exergonic reaction
with value of standard free energy change -61.9 kJ/mole-1. This release of
energy is coupled to ATP synthesis from ADP, which requires 30.5 kJ/mole -
1
.

Pyruvate kinase requires K + and Mg+ for the activity. Like PFK, this
enzyme is also an allosteric enzyme. ATP is the negative modulator of the
enzyme while ADP being the positive modulator. In case ATP accumulates
in the cell and the cell does not need further production of ATP, activity of
pyruvate kinase is inhibited. Reverse happens when ADP accumulates and
the cell needs to produce ATP, pyruvate kinase activity is promoted.

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Summary of glycolysis
During glycolysis one molecule of glucose will produce two molecules of pyruvate. In
the preparatory phase 2 molecules of ATP are consumed and during the payoff phase
2 molecules of NAD+ are reduced to 2 molecules of NADH and 4 molecules of ATP are
produced. So, there will be net gain of 2 ATP molecules and 2 NADH. The net reaction
of glycolysis can be written as,

Glucose + 2 NAD+ + 2 ADP + 2 Pi ------ 2 Pyruvate + 2NADH +2H+ + 2 ATP

Figure: Figure showing the complete pathway of glycolysis, i.e., conversion of glucose
to pyruvate
Source: Author

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Deficiency of glycolytic enzymes leads to hemolytic anemia. In hemolytic anemia

there is premature RBC destruction. A mature RBC is 100% dependent on
anaerobic glycolysis for making their ATP. There will be no ATP production due to
deficiency of glycolytic enzymes. As a result of this Na+/K+ pump activity will be
reduced which is required for the shape and function of RBC resulting in swelling
and breaking of RBC.

Regulation of glycolysis
Any metabolic pathway needs to be regulated, i.e., it should operate when the
cell needs the products of the pathway and should be switched off when these
compounds are not required by the cell, otherwise the energy required for the
operation of the pathway will be a wasteful expenditure. In the multistep pathway
some of the steps are the regulatory steps, which are catalyzed by the regulatory
enzymes. Some of the allosteric regulators of these enzymes are ATP, ADP,
NADH, and NAD+.

Three reactions of the glycolysis are the regulatory steps.

1) Conversion of glucose to glucose-6-phosphate catalyzed by hexokinase.

2) Fructose to fructose1, 6-bisphosphate catalyzed by phosphofructokinase
3) Formation of pyruvate from PEP catalyzed by pyruvate kinase.

In animals primary control point is the activity of PFK. PFK is allosterically

regulated enzyme. When ADP accumulates in the cell, it is an indication that the
cell needs to produce ATP. ADP is the positive modulator of PFK and its
accumulation will result in the production of fructose 1,6 bisphosphate, which will
activate the activity of the enzyme pyruvate kinase. This kind of control
mechanism is called feed forward effect.

In plants the enzyme pyruvate kinase is insensitive to fructose 1, 6 bisphosphate.

It the PFK which is sensitive to accumulation of PEP, i.e., when PEP accumulates,
activity of PFK is inhibited. This is called bottom up regulation. In plants PEP is
metabolized by PEP carboxylase also, in addition to PEP being metabolized by
pyruvate kinase. PEP carboxylation will result in generation of malate, which can
be further be metabolized in the mitochondria. When PEP accumulates and is not
being metabolized, it will inhibit the activity of PFK. Inorganic phosphate
however, will decrease the effect of PEP on PFK. So, in plants it is the ratio of
cytosolic PEP/Pi, which regulates the activity of PFK and in turn regulates
glycolysis.

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Figure: Regulation of glycoysis

Source: Author

Why are the glycolytic intermediates phosphorylated?

Out of the ten intermediates of the cycle nine are phosphorylated. What is the
significance of this? The phosphate group present in the compound may be
serving the following functions:

1. Most of the phosphorylated sugars cannot be transported out of the cell,

because of the fact that membrane lacks their transporters. So, the energy
is not required to retain them in the cell inspite of the large difference in
their concentrations inside and outside the cell.
2. Energy of the high-energy phosphate group of ATP is partially retained in
the form of high-energy phosphorylated compounds, which ultimately is
conserved in the form of ATP in the payoff phase of glycolytic cycle.
3. Presence of the charged phosphate groups in the substrates makes it easier
for them to bind to the active sites of the enzyme and it also increases the
specificity of the reaction.

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Unique features of plant glycolysis

1. In animals, it is mainly glucose, which is catabolized, while in plants,

sucrose is the form in which carbohydrates are mobilized. So it is the
sucrose, which enters the glycolytic pathway.
2. In plants glycolysis occurs also in plastids, in addition to the cytosol.
3. PEP may also be metabolized to malate in addition to its conversion to
pyruvate. Alternate metabolic pathway gives more flexibility to plants in
order to adjust to changing environmental conditions.
4. Regulation of glycolysis is different in plants from that of occurring in
animals. In plants it is the bottom-up regulation while in animals it is feed
forward regulation.

Reoxidation of cytosolic NADH:

NADH produced during glycolysis needs to be re-oxidized so that the glycolysis

continues operating. Glycolysis will stop if there is no oxidized NAD + available.
This re-oxidation of NADH is possible by means of the following pathways:

1. NADH is re-oxidized during fermentation process under anaerobic

conditions.
2. In plants, reoxidation of NADH or NAD(P)H is possible by the external
dehydrogenase of the electron transport chain, which are localized on the
outside of the inner mitochondrial membrane. These dehydrogenases
transfer electrons from NADH/NAD(P)H directly to Ubiquinone and
ultimately the electrons are transferred to O2.
3. The cytosolic NADH needs to be transported to mitochondria where these
can be reoxidized by the electron transport chain, which is localized in the
inner mitochondrial membrane. The inner mitochondrial membrane is
impermeable to NADH, so there are mechanisms by which the reducing
equivalents can be transported through the mitochondrial membrane. This
movement of NADH across inner mitochondrial membrane is called NADH
shuttle system. The direction of the NADH transfer is determined by the
ratio of NADH/NAD in between cytosol and mitochondrial matrix across the
mitochondrial membrane. If the ratio is low in mitochondria, NADH
equivalents are translocated from cytosol to mitochondria, while in case the
ratio is high, the NADH is shuttled fro mitochondria to cytosol. We are going
to study the NADH shuttle system in the following text. In plants, NADH can
be shuttled by following mechanisms.

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 Glycolysis and Oxidative Pentose Phosphate Pathway

a) Glycerol phosphate shuttle: Cytosolic dihydroxyacetone

phosphate is reduced to glycerol phosphate, which is coupled with
the oxidation of NADH. DHAP can diffuse through the outer
mitochondrial membrane. The membrane is freely permeable to
glycerol phosphate, which reaches the outside of the inner
mitochondrial membrane. Glycerol phosphate donates electrons
directly to Ubiquinone of the electron transport chain which is
localized in the inner mitochondrial membrane and 2 ATP molecules
will be produced as a result of electron transfer through the chain
and ultimately to O2, which is the terminal electron acceptor in the
electron transport chain.

Figure: Diagram showing NADH shuttle system: the above figure showing malate-
oxaloacetate shuttle, while the lower figure showing glycerol phosphate shuttle
OAA- Oxaloacetae, Tr-transaminase, DHAP-dihydroxyacetone phosphate, Gl-3P-
glyceraldehyde-3-phosphate, cG1-3PD-cytosolic glyceraldehyde-3-phosphate
dehydrogenase, mG1-3PD-mitochondrial 3 glyceraldehyde-phosphate
dehydrogenase.

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 Glycolysis and Oxidative Pentose Phosphate Pathway

b) Malate/oxaloacetate shuttle: In this kind of shuttle system NADH

is shuttled through inner mitochondrial membrane in the form of
malate, which is synthesized as a result of reduction of oxaloacetate
by the dehydrogenase using cytosolic NADH. Malate when reaches
the mitochondrial matrix, is reoxidized to oxaloacetate, which is
translocated back to cytosol.
c) Malate- aspartate shuttle: Exchange of the carbon compounds
takes place in the form of malate and aspartate, instead of malate
and oxaloacetate, which you have studied above. In the
mitochondria matrix, oxaloacetate is aminated to form amino acid
aspartate by transamination reaction (4). Aspartate is shuttled to
cytosol (5). The aspartate and α–keto glutarate, produced as a result
of transamination reaction in the mitochondrial matrix, are
exchanged with malate and glutamate (5 & 2) from the cytosol. The
oxaloacetate is reduced to malate coupled with the oxidation of
NADH to NAD+ and is recycled to mitochondria where it is further
oxidized to oxaloacetate reducing NAD+ to NADH. In this way
reducing equivalents are transferred from cytosol to mitochondria
and NADH is oxidized in mitochondria.

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 Glycolysis and Oxidative Pentose Phosphate Pathway

Figure: Details of malate aspartate NADH shuttle, numbers indicate the

sequence of reactions.

Figure: Diagram showing utilization of glycolysis intermediated and products in

the cell.
Source: https://adapaproject.org/bbk/tiki-

index.php?page=Leaf%3A+Why+is+glycolysis+called+the+basic+metabolic+pat
hway+of+life%3F

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 Glycolysis and Oxidative Pentose Phosphate Pathway

Video:Glycolysis Overview
Source: http://vcell.ndsu.nodak.edu/animations/glycolysis_overview/movie-
flash.htm

Video: Glycolysis reactions

Source:http://vcell.ndsu.nodak.edu/animations/glycolysis_reactions/movie-
flash.htm

Fate of Pyruvate
Fermentation
Fermentation is a process involved in the conversion of sugars or starch to either
acids or ethanol in absence of oxygen. Glycolysis can continue even in absence of
oxygen if NADH is oxidized to NAD+ and plants can continue to obtain some
amount of energy produced in glycoysis. The rate of glycoysis increases in
absence of oxygen. This response of glycolysis to absence of oxygen is known as
Pasteur effect. Pasteur effect was named after the discoverer of this observation,
a French microbiologist Louis Pasteur. Increased glycolytic pathway is coupled
with up regulation of the genes responsible for the production of glycolytic
enzymes and also for the enzymes responsible for formation of fermentation
enzymes, which you will be studying in the following text.

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 Glycolysis and Oxidative Pentose Phosphate Pathway

Pyruvate metabolism in roots of the plants growing in water logged

conditions
The end product of glycolysis is pyruvate. Its fate is determined by the presence
or absence of oxygen. If the oxygen is available in the cell, pyruvate enters
mitochondria and is metabolized there, which we are going to study in the next
lesson. However, in absence of oxygen, pyruvate is metabolized in cytosol by
fermentation.

Under hypoxia (3 kPa oxygen) or anoxia (0 kPa oxygen) conditions, pyruvate will
not be oxidized further in mitochondria. NADH produced during oxidative
decarboxylation of pyruvate in mitochondria is not oxidized by electron transport
chain because of absence of oxygen. O2 is the terminal electron acceptor in the
electron transport chain. As a result NADH, which is produced as a result of
reduction of NAD+ during glycolysis, will accumulate. The accumulated NADH will
inhibit oxidative decarboxylation of pyruvate. In that case pyruvate needs to be
metabolized in cytosol by fermentation, which is coupled with oxidation of NADH
to NAD+. NAD+ is required for the continued operation of glycolysis.

Generally plants are exposed to ample supply of oxygen and they do not face
hypoxia or anoxia conditions. In water logged conditions or flooded soil, initially
roots are exposed to hypoxia followed by anoxia conditions of the soil and the
pyruvate needs to be metabolized by the fermentation process. The principal
products of glycolysis in either hypoxia or anoxia are lactate and ethanol.
Sometimes alanine, succinate and γ–aminobutyrate (GABA) may also be formed.
The types of end products in the oxygen deprived condition are determined by
the plant species, genotype of the plants, duration and the severity of the oxygen
deprivation.

Initially cytosolic lactic acid dehydrogenase (LDH) reduces pyruvate to lactic acid
reoxidizing NADH to NAD+. Accumulation of lactic acid lowers the pH of the
cytosol (acidosis), which inhibits the activity of LDH, and activity the enzyme
pyruvate decarboxylase is stimulated, the enzyme, which has lower pH optima.
Pyruvate decarboxylase (PDC) catalyzes decarboxylation of pyruvate to
+
acetaldehyde. The enzyme required Mg and thiamine pyrophosphate as the
cofactor (Thiamine itself is vitamin B1). PDC activity results in the formation of
acetaldehyde. The acetaldehyde is reduced to ethanol by the enzyme alcohol
dehydrogenase (ADH), reoxidizing NADH to NAD+. Ethanol is nonpolar and can
diffuse across plasma membrane. As a result of this cytosolic pH is stabilized at

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 Glycolysis and Oxidative Pentose Phosphate Pathway

slightly lower pH. Both of the enzymes, lactic acid dehydrogenase (LDH) and
alcohol dehydrogenase (ADH) are similar in many ways. One of the similarities is
that both the enzymes are NAD+ linked dehydrogenases. Cell membranes of the
root cells are permeable to the product of fermentation i.e., ethyl alcohol, which
will leach out in the soil.
Flood tolerant plants are able to switch to ethanol production to avoid cytoplasmic
acidosis.

Figure: Diagram showing fate of pyruvate in by the yeast extract and the muscle
extract, x 2 indicates that each glucose molecule produces two molecules of
pyruvate
Source: Author
Pyruvate metabolism in muscles

Pyruvate is metabolized in the muscles by its reduction to lactate by the enzyme

lactate dehydrogenase (LDH). LDH reoxidizes NADH to NAD +. During period of
extensive exercise, lactate is formed in the muscles due to oxygen deprivation.
Lactate accumulation in the muscles causes fatigue.

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 Glycolysis and Oxidative Pentose Phosphate Pathway

Lactate is the dead end of the anaerobic glycolysis, but it can be recycled in the
muscles to form pyruvate and further to glucose by means of gluconeogenesis,
which will be discussed elsewhere.

Figure: Diagram showing reoxidation of cytosolic NADH by the lactate

dehydrogenase and alcohol dehydrogenase.
Source: Author
Dental plaque and fermentation: bacteria in the mouth grow using sucrose and
form the sticky colonies on the tooth surface. Bacteria grow below the surface of
the plaque under anaerobic conditions, forming lactic acids. The lactic acid
destroys the enamel of the tooth causing the decay.

Significance of fermentation
During glycolysis there is net gain of 2 ATP molecules and 2 molecules of reduced
NADH, when one molecule is converted to 2 molecules of pyruvate. NADH needs
to be oxidized back to NAD+ so that glycolysis can continue. Oxidized NAD+ is
required for glycolysis to continue. In the presence of oxygen, NADH+ is oxidized
by the electron transport chain, which is present in the inner mitochondrial
membrane. However, in absence of oxygen, NADH+ is oxidized during
fermentation process.

So, in order for the glycolysis to continue in absence of oxygen, oxidation of the
NADH+ by fermentation is required, otherwise NADH will accumulate and
glycolysis will stop in absence of oxidized NAD+ and 2 molecules of ATP, which
were being generated during glycolysis will also not be produced.

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 Glycolysis and Oxidative Pentose Phosphate Pathway

Figure: Significance of fermentation

Source: Author
During fermentation there is net gain of 2 ATP molecules per glucose molecule,
which can be utilized for the growth of the organisms. Contrary to this number
of ATP molecules produced during aerobic respiration varies from 30-32. The
famous French microbiologist, Louis Pasteur observed, when yeast switched
from aerobic conditions to anaerobic, there was increased glycolytic rate and
there was increased rate of carbohydrate breakdown. This effect was called
Pasteur effect. Under the anaerobic conditions, there was increased production
of glycolytic intermediates and also increased expression of the genes encoding
glycolytic and fermentation enzymes.
Fermentation has been commercially exploited. Recently with the growing
awareness about use of ethanol as an alternative source of fuel, there has been
growing interest in this process. Scientists are interested in using cellulose as
the source for generation of biofuels.

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 Glycolysis and Oxidative Pentose Phosphate Pathway

Oxidative Pentose Phosphate Pathway:

Pathway
Additional pathway of glucose metabolism is Oxidative Pentose Phosphate
Pathway. This pathway is not an alternative to glycolysis, rather occurs in
addition to glycolytic pathway. Almost 10-25 % of the glucose is metabolized
through this pathway. In plants, this pathway occurs both in cytosol and the
plastids. Especially in the non green parts of the plants or also in the chloroplasts
in dark, since one of the function of this pathway is to generate NADPH, which is
required for biosynthetic reactions such as in lipid biosynthesis. In green plants,
NADPH is primarily synthesized during the light reactions of the photosynthesis.

We are going to study the metabolic reactions of the pathway. The pathway
consists of two phases:

i) Oxidative phase: Glucose-6-phosphate dehydrogenase catalyzes the

conversion of Glucose-6-phosphate to 6-phosphogluconate. Reduction of
NADP to NADPH is coupled to this oxidative reaction. The pathway is also
known as Phosphogluconate pathway, since the first compound formed is
Phosphogluconate. In the next step, 6-phosphogluconate is decarboxylated
to keto-pentose (ribulose-5-phosphate), which is again coupled to reduction
of NADP+ to NADPH. The reaction is catalyzed by the enzyme 6-
phosphgluconate dehydrogenase. In the oxidative phase one molecule of
glucose-6-phosphate will produce one molecule of ribulose-5-phosphate,
one molecule of CO2 and 2 molecules of reduced NADPH.

NADPH is used as a reducing power for biosynthetic reactions, while

ribulose-5-phosphate is converted to ribose-5-phosphate by the action of
the enzyme phosphopentoisomerase. Ribose-5-phosphate is used as a
precursor for nucleotide biosynthesis. The first two reactions of the
oxidative phase are irreversible reactions and are regulated by the ratio of
NADPH/NADP+ in the cell. NADPH being the negative regulator of the
enzyme and the NADP+ is the positive regulator. The pathway will operate
in the cells, which have the requirements for both NADPH and ribose-5-
phosphate, such as the dividing cells. However, if the cell needs NADPH, but
does not require ribose-5-phosphate, then the non oxidative phase of the
pathway will convert ribulose-5-phosphate back to glucose-6-phosphate,
which further can be utilized for NADPH generation by means of the

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 Glycolysis and Oxidative Pentose Phosphate Pathway

oxidative pathway. In the following section we are going to study the

nonoxidative phase of the pathway.

Glucose-6-phosphate + 2 NADP+----- ribulose-5-phosphate +

CO2+ 2 NADPH + 2H+

Figure: Oxidative stage of Pentose Phosphate Pathway.

Source: Author

ii) Non-oxidative phase: many of the reactions of this phase are quite
similar to those of the Calvin cycle, which is also called reductive pentose
phosphate pathway. There are two most important enzymes, which catalyze
the inter conversion of the various sugars. These are transaldolase and
transketolase. Transaldolase and transketolase together interconvert
various sugars depending upon the need of the cell.

Transaldolase detach C3 unit from a ketose sugar and transfer it to other aldose
sugar, while transketolase detach C2 unit for a ketose sugar to an aldose sugar.
A number of sugar interconversions are possible which are part of the non-
oxidative phase of the pathway and have been shown in the given figure.

Ribulose-5-phosphate is interconverted to ribose-5-phosphate and

xylulose-5-phosphate by the enzymes phosphopentoisomerase and

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 Glycolysis and Oxidative Pentose Phosphate Pathway

phosphopentoepimerase respectively. Various transformations occur in

between the sugars as given in the figure.

Figure: Non-Oxidative stage of Pentose Phosphate Pathway.

Source: Author

Significance of oxidative pentose phosphate pathway

1. In the plastids of non-green tissues (such as the root cells) and in
chloroplasts in the dark, this pathway is the sole source of NADPH, which
otherwise is produced during the light reaction of photosynthesis. NADPH is
required for lipid biosynthesis and nitrogen assimilation. In animal tissues
this pathway is the sole source of NADPH. NADPH is required for reduction
of glutathione, which is critical for maintaining the plasma membrane.
2. In plants cytosolic NADPH can also be oxidized by the external NADPH
dehydrogenase present on the cytosolic side of the inner mitochondrial
membrane, which transfers the electrons to electron transport chain directly
via Ubiquinone, and is rotenone insensitive. So, this can contribute to ATP
generation.

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 Glycolysis and Oxidative Pentose Phosphate Pathway

3. The pathway supplies ribose-5-phosphate, which is required for nucleotide

biosynthesis.
4. Another intermediate of this pathway is erythrose-4-phosphate, which is the
precursor of plant phenolic compounds, aromatic amino acids and also of
lignin when it combines with phosphoenolpyruvate, which is a glycolytic
intermediate. There are reports that the enzymes catalyzing the pathway
are induced in plants under stress conditions.

Figure: Diagram showing the utilization of the intermediates and products of the
pentose pathway in the cell.
Source:https://adapaproject.org/bbk/tiki-
index.php?page=Leaf%3A+Why+is+glycolysis+called+the+basic+metabolic+pat
hway+of+life%3F

Regulation of oxidative pentose phosphate pathway:

Oxidative and the non-oxidative phase of the pathway operate depending upon
the requirements of the cell. The oxidative phase is regulated by the ratio of
NADPH/NADP+. If there is need of NADPH but ribose-5-phosphate is not required,
ribose-5-phosphate is converted back to glucose-6-phspate by the non-oxidative
reactions.
6 X ribose-5-phosphate----- 5 X glucose-6-phosphate

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 Glycolysis and Oxidative Pentose Phosphate Pathway

Since hexose phosphate is re-generated, the pathway is also called Hexose

Monophosphate Shunt (HMS).

In case the cell needs ribose-5-phosphate for nucleotide biosynthesis, but does
not require NADPH, following reaction will meet the requirement:
5 glucose-6-phosphate +ATP-- 6 ribose-6-phosphate + ADP + H+

People deficient in the enzyme glucose-6-phosphate dehydrogenase will exhibit

symptoms of hemolytic anemia, on being given anti malarial drug pamaquine.
Pamaquine increases the reactive oxygen species and NADPH is required for
protective mechanism, and in the deficiency of the above said enzyme, the RBC
are not able to produce NADPH which is also required to maintain hemoglobin in
its reduced (ferrous) state.

Video: https://www.youtube.com/watch?v=-jBAZCxzcBY (CC)

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 Glycolysis and Oxidative Pentose Phosphate Pathway

Summary

 Respiration is a multistep energy releasing process. Some of the released

energy is conserved in the form of ATP.
 Respiration may occur in presence or in absence of O2. Initial steps of
glucose breakdown are similar both in the presence or absence of O2.
 Glucose breakdown pathway is known as glycolysis, which was elucidated
by a number of scientists including Embden, Meyerhoff and Parnus.
 Net gain of glycolysis is 2 ATP, 2 NADH and 2 molecules of pyruvate.
 In the absence of O2, pyruvate is metabolized in cytosol by fermentation.
Fermentation can be either lactic acid or alcohol.
 In absence of O2, fermentation is necessary for reoxidation of NADH, so that
anaerobic respiration continues.
 In presence of O2 NADH produced during glycolysis needs to be transported
to mitochondria by NADH shuttle system, so that these are oxidized by
electron transport chain.
 In presence of O2, pyruvate is transported to mitochondria where it is
metabolized further.
 Glycolysis is regulated by the regulatory enzymes catalyzing regulatory
steps, especially the ones which are catalyzed by phosphofructokinase and
pyruvate kinase. In plants regulation of glycolysis is bottom up regulation
while in animals it is feed forward regulation.
 The other pathway of glucose metabolism in cytosol is by means of
oxidative Pentose Phosphate Pathway. The pathway includes the first two
oxidative reactions in which 2 molecules of NADPH are produced along with
loss of one carbon as CO2 and one molecule of ribulose 5 phosphate is
produced. The non-oxidative part of the pathway includes interconversion of
sugars finally glucose-6-phosphate is generated. So, the pathway is also
called Hexose Monophosphate Shunt. The pathway is significant because
the reducing power is generated in the form of NADPH, besides production
of ribose-5-phsphate and other sugars which are required for biosynthesis
of some of the biomolecules of the cell.

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 Glycolysis and Oxidative Pentose Phosphate Pathway

Exercises

Q.1 Match the followings:

Reaction Enzyme
i) Fructose-6-phosphate--F1,6,BiP Hexokinase
ii) Glucose+ATP-- G-6-P pyruvate kinase
iii) 3-PGAldehyde+NAD++iP--1,3BiPGA + NADH mutase
iv) PEP + ADP----- Pyruvate+ ATP PFK
v) 3-Pglycerate---- 2-Pglycerate phosphhexoisomerase
vi) G-6-P ----- F-6-P G3P dehydrogenase

Q.2 Name the following:

i) Primary regulatory enzyme of the glycolytic pathway in plants.
ii) The oxidative reaction of the glycolysis.
iii) Mechanism of ATP synthesis during glycolysis.
iv) The enzyme, which catalyze condensation reaction in glycolysis.
v) The reducing power generated during oxidative pentose phosphate
pathway.
vi) Cellular site/sites in plants where glycolysis occur.
vii) The scientist/scientists who for the first time observed that fermentation
can be carried out by cell free extract of yeast.
viii) The mechanism of regulation of the activity of PFK.

Q.3 Fill in the blank:

i) PEP + ADP --------- pyruvate + ATP
Aldolase
ii) ____________ --------- G-3-P + DHAP
G-3-P dehydrogenase
iii) G-6-P + NADP -------- ___________+ NADPH
enolase, Mg2+
iv) 2-PGA -------_______ + H2O
v) Ribulose-5-P ------- xylulose-5-P
transaldolase
vi) Sedoheptulose-7-P + G-3-P------________+ F-6-P

Q.4 How was glycolytic pathway discovered? Give in brief the important
contributions of the scientists?

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 Glycolysis and Oxidative Pentose Phosphate Pathway

Q.5 Write down significance of the following pathways in the cell:

i) Fermentation
ii) Oxidative pentose phosphate pathway

Q.6 What is the significance of glycolytic intermediates being

phosphorylated?

Q.7 Write down the Hexose Monophosphate Pathway diagrammatically.

Q.8 Find out the difference in structure and function of NADPH & NADH.

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 Glycolysis and Oxidative Pentose Phosphate Pathway

Abbreviations used in the text:

PEP Phosphoenolpyruvate
G-3-P Glyceraldehyde-3-phosphate
DHAP Dihydroxyacetone phosphate
S7P Sedoheptulose-7-phosphate
FAD Flavin Adenine Dinucleotide
NAD+ Nicotinamide Adenine dinucleotide
NADP+ Nicotinamide Adenine Dinucleotide Phosphate
EMP Embden Meyerhoff Parnus
ATP Adenosine Triphosphate
ADP Adenosine diphosphate
PDC Pyruvate decarboxylase
ADH Alcohol dehydrogenase
GABA γ-amino butyric acid

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 Glycolysis and Oxidative Pentose Phosphate Pathway

Glossary

Acidosis: acidification of cytosol due to formation of lactic acid or any other

acids.
Aerenchyma: Anatomical feature of roots found in hypoxic conditions, showing
large, gas-filled intercellular spaces in the root cortex.
Anhydride: Product of condensation of two carboxyl or phosphate groups in
which water is eliminated.
Anoxic: Environmental condition, which refers to absence of O2.
Cell-free extract: cell membranes are broken and the filtered homogenate is
called cell free extract.
Coenzyme: An organic cofactor required for the action of certain enzymes.
Dialysis: removal of small molecules from a solution of macromolecules by their
diffusion through semipermeable membrane.
Exergonic: energy releasing
Endergonic: energy absorbing
Free energy: it is the energy available to do work isothermally. It is
thermodynamic quantity.
Hypoxia: Reduced O2 content in air or in a body of water.
Phophorylation: formation of phosphate derivative of a biomolecule usually by
the transfer of a phosphoryl group from ATP.
Redox: it is an oxidation reduction reaction in which electrons are transferred
from a donor to an acceptor molecule.
Standard free energy change: free energy change for a reaction occurring
under a set of standard conditions: temperature 298 K; pressure 1 atm and all
solutes at! M concentration. G 0’ denotes the standard free energy change at pH
7.0 in 55.5 water.
Sulfahydryl groups: -SH groups present in the side chain of an amino acid.
Translocator: An enzyme that catalyze membrane transport.
Transamination: Enzymatic transfer of an amino group from an α–amino acid to
an α–keto acid.

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 Glycolysis and Oxidative Pentose Phosphate Pathway

Key to the answers:

Q.1 i0 PFK ii) Hexokinase iii) G3P dehydrogenase iv) Pyruvate kinase v) mutase
vi) phosphohexoisomerase
Q.2 i) Pyruvate kinase ii) conversion of G3P to 1,3 phosphoglycerate catalyzed by
enzyme G3P dehydrogenase iii) substrate level phosphorylation iv) Aldolase v)
NADPH vi) cytosol and plastids vii) Hans Buchner & Eduard Buchner viii) allosteric
regulation
Q.3 i) Pyruvate kinase ii) Fructose 1, 6 biphosphate iii) 6-phosphogluconate iv)
PEP v) epimerase vi) Erythrose 4 phosphate

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 Glycolysis and Oxidative Pentose Phosphate Pathway

References
Buchanan Bob B., Wilhelm Gruissem and Russell L.Jones, Biochemistry and
Molecular Biology of plants, 2000, The American Society of Plant Biologists

Campbell Mary K., Shawn O.Farrel, Biochemistry 7 th edition 2012, Brooks/Cole,

cengage Learning.

Nelson David L., Michael M. Cox, Lehninger Principles of Biochemistry, 2013,

Macmillan Higher Education

Taiz Lincoln, Eduardo Zeigger, Plant Physiology 5 th edition, 2010, Sinauer

Associates Inc., Publishers Sunderland, Massachusetts, U.S.A.

Armstrong, Frank B., Biochemistry 2nd edition, 1984, Oxford University Press
Inc.

Jones Russel, Helen Ougham, Howard Thomas and Susan Waaland, The
Molecular Life of Plants, 2013, American Society of Plant Biologists.

Further reading

https://adapaproject.org/bbk/tiki-
index.php?page=Leaf%3A+Why+is+glycolysis+called+the+basic+metabolic+pat
hway+of+life%3F
http://www.uic.edu/classes/bios/bios100/lecturesf04am/lect12.htm
http://www.ncbi.nlm.nih.gov/pubmed/12753973

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