31

and several students were openly proud of their final

Version even before I graded it. A larger number of
students performed the exercise in a way that they
believed would satisfy me, so self-assessment may be
necessary for maximum development of students in this
context. I read many thoughtful and constructive criticisms of the first versions of papers, and I believe that the
students learned a little about each other and about the
literature on amino acid metabolism during this exercise.
Moreover, the librarians at our college were pleasantly
surprised at the interest shown by students in the library
services and were impressed with their ability to learn to
use indexes and to locate information efficiently on their
own. Development of identity, the capacity for intimacy,
and the ability to sustain productivity were all facilitated
by this simple activity, and many other such projects could
be incorporated into our courses and curricula without
jeopardizing the quality of medical education.
Ultimately, whether or not an educator should design
courses and curricula to foster ego development in
students depends on the educator's values. I wonder why
the present system of medical education, which seems to
stifle intellectual development and creativity in students,
is valued relatively highly by most basic scientists who also
value intellectual advancement and creative research in
themselves and in their peers. Perhaps if we could
understand our apparent ambivalence towards ego development and creativity, we could use our own ingenuity
more effectively to increase our productivity in all of our
endeavors.

References
1Erikson, E H (1980) Identity and the Life Cycle; W W Norton and
Company, New York
2Wilber, K, Engler, J and Brown, D P (1986) 'Transformations of
Consciousness', New Science Library, Shambhala Publications Inc,
Boston
3Levinson, D J (1978) 'The Seasons of a Man's Life', Alfred A Knopf,
New York
4Gilligan, C (1982) 'In a Different Voice', Harvard University Press,
Cambridge, MA
5Keller, E F (1985) 'Reflections on Gender and Science', Yale
University Press, New Haven
6Gallagher, J M and Reid, D K (1981) 'The Learning Theory of Piaget
and Inhelder', Brooks/Cole Publishing Co, Monterey, CA
7pfeiffer, R J (1983) Early-Adult Development in the Medical Student,
Mayo Clin Proc 58, 127-134
8Mann, R D (1967) 'Interpersonal Styles and Group Development',
John Wiley and Sons, New York
9Bloom, B S (1956) 'Taxonomy of Educational Objectives Handbook I:
Cognitive Domain', Longmans Green, New York
lNeufeld, V R and Barrows, H S (1974) 'The McMaster Philosophy: An
Approach to Medical Education', J Med Educ 49, 1040-1050
11Morley, C G D and Biumberg, P (1987) 'Learning Medical School
Biochemistry Through Self-Directed Case-Oriented Study', Biochem
Educ 15, 184-188
12Bion, W R (1961) 'Experiences in Groups', Tavistock Publications,
London
13Anzieu, D (1984) 'The Group and the Unconscious', (translated by B
Kilborne), Routledge and Kegan Paul, London
14janis, I L (1972) 'Victims of Groupthink', Houghton Mifflin Co,
Boston

BIOCHEMICAL EDUCATION 17(1) 1989

15Van Winkle, L J (1980) A Framework for Teaching Amino Acid

Metabolism, Biochem Educ 8, 72-75
16Van Winkle, L J (1985) A Summary of Amino Acid Metabolism Based
on Amino Acid Structure, Biochem Educ 13, 25-26

Metabolism: Where to Begin?

RONALD W BROSEMER

Program in Biochemistry~Biophysics
Washington State University
Pullman, WA 99164-4660, USA
Introduction
Metabolism is an area covered in all introductory biochemistry lecture courses, whether the course be designed
as a summary for undergraduates in allied biological
disciplines or as an exhaustive (and exhausting) coverage
for biochemists and others who use biochemical principles
extensively. Yet regardless of the level of presentation,
metabolism presents challenges to the lecturer that are
usually greater than those presented by other broad
biochemical topics, such as structure of biomolecules and
gene function.
One of the special hurdles in teaching metabolism is the
perception that it is not a 'hot' field. Much of the material
covered in introductory courses first appeared in the preElvis Presley era, and most of the rest before Watergate.
Many students would love to learn all the latest details on
how to manipulate and move a gene without taking time
to comprehend what in fact the enzyme product of that
gene is doing in a cell. Medical students complain that too
much effort is spent learning pathways and not enough
time on practical applications - - applications that require
a firm understanding of metabolic facts and principles.
Metabolism is certainly an active field of research albeit
one not as well funded as other areas, but these
contemporary topics are generally difficult to incorporate
into introductory courses. So much time must be devoted
to covering basic metabolic pathways, there is little room
left for the recent advances in control and integration.
Comparison of the latest editions of introductory biochemistry texts with prior editions usually shows that the
chapters on metabolism contain the fewest changes. How
can we understand the implications of "To be or not be
be" without first spending considerable time and effort
learning that "A" is for apple and "B" is for ball?
Goals
Before undertaking the task of preparing a series of
lectures, an instructor should have a clear idea of the goals
of the presentations. I strongly believe that these goals
should be made available in writing to students during the
first class. Since the subject matter of metabolism in
introductory courses presents special difficulties, it is
critical that the instructor's expectations for the majority
of students be clearly defined. In order to stimulate each

32
of us who teaches metabolism to grapple consciously and
openly with goal definition and to invite comment and
refutation, I present some thoughts on setting these goals.
I limit my comments to introductory courses for students
who will not be entering biochemistry, molecular biology,
or closely allied fields. The type of course I am addressing
tends to be a 15- to 20-week sequence that is the only
formal biochemistry course taken, with about one-third of
the lectures devoted to metabolism. I include first-year
medical courses, fully aware that there is a wide gap
between what medical students should know about biochemistry and what they will in fact condescend to learn.

Memorization
More than with most other students, we hear the lament:
"I don't have to learn all those pathways, because I can
look them up in a book." Any self-respecting instructor
rejects that argument out of hand, b e c a u s e . . . At this
point each instructor can insert his or her own favorite
recipe, such as: I had to do it; it is good for you; you
should be ready to accept hard work and challenges; if I
cover it in lecture, it is important enough to memorize.
Experience has convinced me that for the large majority
of students in these introductory terminal courses,
memorization of more than just a bare minimum of
structures and reactions results in a decreased understanding of the underlying concepts in metabolism. This caveat
does not apply to advanced courses (including introductory courses for biochemistry students and fellow
molecular biologists), since we cannot truly understand
basic principles in depth without knowing the underlying
facts. As biochemists and instructors, we tend to equate
the biochemistry major (who is basically a reflection of
our own interests and abilities) with students in the
terminal elementary courses. What is good for the major
goose is not necessarily good for the minor gander. Since
we live in a 24-hour world with 10 or 15 week periods for
turning a naive human brain into a storehouse of
erudition, we must make unsatisfying compromises. As
much as we would like to see all students know as much as
we do, there is simply not enough time both to memorize
structures and reactions and to acquire a working knowledge of how they fit together in a living cell. For most
students, memorization becomes the end in itself; their
satisfaction or relief at achieving this immediate goal
leaves little incentive for the deeper, and more important,
appreciation of metabolism.

Thermodynamics and chemistry

Among my more unorthodox views is the opinion that a
qualitative approach should be taken for coverage of
thermodynamics in these survey courses. If we do some
searing soul searching, I feel that most of us would
reluctantly admit that these students do not in fact have to
know about R and T and natural logarithms in order to
have a good grasp of the principles of free energy changes.
They should know that -13,000 calories/mole is a much
better driving force than -5,000, but this can be clearly

BIOCHEMICAL EDUCATION 17(1) 1989

presented without 'RT In'. How many students have we

lost by discussing a gas constant in a system where there
are so few gases? The introduction of standard free energy
is a great disservice in these courses. We expend so much
effort and time on the intricacies of standard free energy,
while somewhere along the line the fact is mumbled that
most metabolites are not present in 1 M concentrations in
real cells. Free energy change can be effectively presented
by simply explaining that it depends upon the actual
concentrations of substrates and products and upon how
far these lie from equilibrium. The concepts of equilibrium constant, of LeChatelier's Principle (both of
which were covered in introductory chemistry in some
detail) and of coupling reactions via common intermediates suffice to produce a working, effective and longlasting understanding of thermodynamics for unaccredited
biochemists. My major goal in the segment on thermodynamics is to encourage students to think in terms of
energy-producing or energy-requiring reactions and how
one is used to drive the other.
Although organic chemistry is a prerequisite for the
type of biochemistry course I am addressing, we instructors all too often fail to appreciate the fact that a large
portion of our clientele do not grasp the essence of a
molecular structure written on paper or a blackboard. Let
me illustrate with the structural formula for glyceraldehyde 3-phosphate. When we biochemists see this formula,
we immediately perceive the aldehyde group (which can
be oxidized or reduced), a free hydroxyl, and a phosphate
ester. For many students, this structural formula is just a
random accumulation of the letters C, H, O, and P. Yes,
they should have learned better in organic chemistry; but
the fact is many did not. By the way, I have met several
students who received high grades in a basic organic
chemistry course but who have no comprehension of
structural formulae. As biochemists we should not be
expected to teach these elementary chemical concepts,
but we should be well aware that many students in the
audience see all the structures we present as just an
alphabetic jumble.

General principles
Glycolysis, the Krebs cycle, respiratory chain, oxidative
phosphorylation, gluconeogenesis - - certainly the
enzymic bases for these key pathways and processes must
be covered in detail. But how many of us take these
concepts beyond the memorization stage? What general
principles are illustrated by these paths, principles that
might remain long after the monotonous individual
reactions have been forgotten? For example, how is an
alcohol usually oxidized; how is a ketone incorporated
into an aliphatic group; why are there so many phosphorylated metabolites; how are phosphate groups added
to or removed from compounds; why are sulfur, but not
oxygen, esters used extensively in intermediate metabolism? Even students who elegantly and successfully resist
learning metabolic pathways often can appreciate some of
these basic concepts.

33
Medical students present a unique problem in that they
want a specific disease attached to each fact they are
expected to retain. Since there are relatively few metabolic disorders associated with the basic pathways mentioned above, we must emphasize to them that it is
sometimes helpful to learn about the healthy condition,
even though insurance companies will not cover expenses
associated with the hale and hardy. If this fails to elicit
their interest, one can point out that they will probably
make loads of money from diabetic patients. Ears tend to
perk up.
At this point in the course, several choices must be
made. For example, how many additional metabolic
pathways should be covered? Most instructors surely
include the pentose phosphate pathway, B-oxidation, and
fatty acid synthesis. I choose not to cover any of these
three key paths, except in broad outline form. The reason
is an attempt to strike a balance between covering many
individual reactions and introducing underlying themes of
how metabolism as a whole has evolved. With only a
couple of exceptions, the students have already seen every
class of enzyme-catalyzed reaction involved in the above
mentioned pathways; few new concepts are introduced
that cannot be covered by simply pointing out what the
starting and end metabolites of these pathways are. In
fact many of these reactions can be introduced on
examinations to test the students' comprehension of
metabolic patterns with questions about reactions that
they have not seen but that are analogous to reactions
encountered in lecture.

Coverage
There is a widespread but unfortunate philosophy among
students that: "If it is not covered in lecture, I will not
learn it. I expect you as an instructor to discuss every
important topic in lecture, or I am not getting my money's
worth". Or, as medical students would phrase it: "If it
might be on the Board exams, lecture on it". Yet is it not
our major responsibility as instructors to nurture students
to the point where they can function perfectly well without
us? How can we get this critical point across so long as we
bow to demands to cover "everything" in lecture? This is
perhaps the overriding reason for concentrating on only a
couple of tissues or organisms in metabolism. A true
understanding of the basic principles in just one system
should give the learner the intellectual tools necessary to
explore new areas via textbooks, personal interactions
with experts, or further coursework.
These personal views of presenting introductory
metabolism to a general undergraduate or medical student
audience have developed over many years and many
attempts to find an effective approach. The good old days
of "memorize or perish" have gone the way of quarterpound textbooks. We must find ways to develop in
students a sense of the majesty and unity of myriad
metabolic paths and interactions, but for most students we
are graciously granted about 15 hours of impersonal
contact to accomplish this grand task. I wish I knew the
magic formula; my comments here are designed to
encourage the quest for a collective Holy Grail.

Integration
There are many topics that lend themselves to a scaffold
around which integration of metabolism can be constructed; control of glycogen metabolism and starvationdiabetes are two good examples. Whatever the chosen
topics for integration, I believe emphasis should be placed
on delving into the details of only a few examples toward
the end of the metabolism segment rather than discussing
many examples piecemeal as they might arise during
introduction of the various enzymic steps.

Photosynthesis?
Attempts to cover several different biological systems
should be curtailed. I suggest it is pedagogically preferable
to cover one or two tissues in some depth than to survey
the animal, plant and microbial kingdoms superficially.
The most obvious topic example is photosynthesis. If the
framework for introducing metabolism is plants, photosynthesis is a suitable subject. If animal (ie mammalian)
systems are emphasized, photosynthesis should not be
covered. This aggrieves students in the plant sciences,
while delighting those in nutrition. I prefer to cover in
some detail the metabolism of only two human tissues,
liver and muscle, while occasionally mentioning other
examples such as alcoholic fermentation in yeast. Even in
the case of the latter topic, alcohol dehydrogenase leads
into a discussion of liver function.

BIOCHEMICAL EDUCATION 17(1) 1989

B.A.S.C.
Biochemistry forms the basis of modern biology and a
considerable part of what is taught in Biology Courses at
the highschool and junior college level is biochemistry.
Students find biochemistry at once fascinating and
'difficult', and so too do some of their teachers. In
recognition of this The Biochemical Society in London has
formed a working Group called B.A.S.C. (Biochemistry
Across the School Curriculum) to provide resource
materials to assist in the teaching of biochemistry within
the Biology syllabus. Further information about the
activities of this Group and on the availability of the
resource material may be obtained from the Editorial
Office of Biochemical Education or directly from the
Research and Information Officer, The Biochemical
Society, 7 Warwick Court, Holborn, London WC1R 5DP,
UK.