Escolar Documentos
Profissional Documentos
Cultura Documentos
1
2
3
4
Energyyield (%)
23.9
47.8
71.7
95.6
AGnet (kJ/mol)
156
107
58
9
q (eqn 2)
37.3
51.2
41.6
8.6
References
1 Lupianez J.A., Garcia-Salguero L., Torres N.V., Peragon J. and
Melendez-Hevia E. (1996) Comp. Biochem. Physiol. l13B, 439-443
PII: S0307-4412(97)00118-0
A Guided Discovery Approach for Learning
Glycolysis
EMERIC SCHULTZ
Department of Chemistry
Bloomsburg University
Bloomsburg, PA 17815
USA
Introduction
Metabolic pathways are, at the most basic level, a set of
successive chemical transformations. For those in the
know however, each pathway is pregnant with possibilities
for concept development. From the simple beauty of
'naked reactions', connections can be made to many
unifying principles including: (1) the energetics of each
transformation and of the overall process; (2) the balance
between entropy and enthalpy effects; (3) the multi-level
control aspects of the process; (4) the overall strategy of
the process in relationship to other pathways.
Expected student learning of metabolic pathways runs
the gamut from the very superficial to the very inclusive.
For some nursing students, as well as other students in
allied health sciences, often only the names of a small
number of the key compounds in the pathway (most
usually glycolysis and the citric acid cycle), and perhaps
the names of a few enzymes are required. The learning
here is essentially rote. For students in a junior/senior
level biochemistry course or sequence, especially one
taught from a chemical perspective, a thorough understanding of the actual transformations as well as the
energy and control connections is routinely expected.
Unfortunately, in this case also, most of the learning is
rote.
The clientele for our biochemistry sequence consists of
primarily biology majors (greater than 90%). For this
206
audience the nuances of structural changes are often lost,
if indeed they have ever been comprehended. These
students often default to a wholly inadequate rote
memorization of names, enzymes and structures. The
structures of intermediates presented on examinations by
these students often range from the inept, to the bizarre,
to the comical. Students handicapped in this way seldom
see the forest of integrated concepts from the trees of
individual reactions.
A recent editorial in this journal I celebrated the 25th
anniversary of the Metabolic Pathways Charts and
commented on student strategies for studying metabolic
pathways. In the same issue, Nicholson, the leading
'inventor' of the Charts, in addition to describing the
history behind the development of the modern Charts,
presented the pentose phosphate pathway in a manner
that separated the two overlapping 'missions' of this
pathway. 2 The argument was made that the pathway
needed to be assimilated by the student in a way that was
non arbitrary. The suggested way of learning this pathway
was highlighted by considering the different possibilities
that would arise given different cellular requirements.
Another recent paper described three different styles by
which the essentials of the urea cycle could be learned by
students? One of these styles involved the use of a
nmenonic in order to help students remember the
different components of the cycle.
Ryder and Leach describe a unique tutorial approach
that 'requires students to move away from a view of glycolysis and gluconeogenesis as sequences of metabolites and
enzymes, towards an appreciation of the links between
metabolic pathways.. ?.4 1 agree. The challenge for me in
teaching a metabolic pathway is to impart both the
chemical sense in the specific transformations (you can
not walk away from bonding), the energetic sense in the
thermodynamics of each step and the overall process, and
the biological sense in terms of economy of effort dictated
by evolutionary perfection. However, this cannot occur
unless the essentials of the pathway have been assimilated
by the student. In my opinion, more attention needs to be
paid to teaching students how to learn the rudiments of
specific pathways. We need to have a greater assurance
that students understand these nuts and bolts aspects
before attending to the more difficult concepts detailed
above.
This communication describes a unique way of learning
the glycolytic pathway in step-wise fashion. The pedagogy
involves clear rote components that are connected to a set
of generalizations that develop and deepen important
biochemical concepts.
207
(a) Beginning = setting up substrates from which
energy can be extracted by phosphate transfer;
this will be a top row of four reactions = upper
tier.
(b) M i d d l e = a unique event (like the top of a
mountain from which we can see both sides) that
connects one mission to another; this reaction
connects the top row of four reactions to the
bottom row of four reactions.
(c) E n d = u s i n g the high energy substrate compounds to generate metabolic energy in the form
of ATP; this will be a bottom row of four reactions = lower tier.
(3) Number each of the reactions in the pathway.
The result of this discussion is shown in Figure 1.
Step 2 Identify the reactions in which ATP, ADP and
phosphate are involved either as substrates or as products
of reactions:
[]
[]
[]
[]
[]
7-V
[]
ATP
[]
--
ADP
[]
?W"
ATP
ADP
[]
[]
[]
[]
[]
[]
[]
[]
ATP
[]
ADP
ATP
ADP
208
(a) Glycolysis yields not only biological energy in the
form of ATP, but also biological energy in the
form of reducing power, specifically NADH.
(b) Since this reaction is the 'special' one in the
middle of the pathway, its logical to infer that this
is where the N A D H is made.
(c) N A D H is made from NAD + (a reduction); therefore the glycolytic intermediate has to be
oxidized.
(d) Since there are 2 molecules of glyceraldehdye3-phosphate that are going to be oxidized, 2
molecules of NAD + will be reduced to NADH;
the reaction is updated to reflect this fact.
(e) Given our guiding principles, we expect the oxidation to occur at the end of the molecule where
there is no phosphate; we also expect the
phosphate to be added at this end; (the type of
linkage that results, a mixed anhydride, is similar
to the high energy phosphate anhydrides that the
students have already had experience with in
ATP); the resulting compound is 1,3-bisphosphoglycerate.
(f) The logic of this compound and its high energy
status as the first compound in the lower tier in
the pathway is emphasized.
[]
ATP
[]
ADP
[]
ATP
[]
ADP
2 Pi-~.
[]
[]
[]
[]
[]
2 ATP 2 ADP
2 ATP 2 ADp
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3
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