Problem-Based Learning

in the Biology Curriculum

B A R R Y S. K E N D L E R P AT R I C I A A. G R O V E

B ackground
Problem-based learning (PBL) is a pedagogical
approach to learning that involves presentation of a cur-
riculum-related problem or situation whose solution
students who discuss a problem that has been analyzed
previously. In each case, a faculty member (tutor)
guides the session to promote critical analysis and
explore different perspectives.
There are generally four broad objectives in PBL.
requires students to practice skills of analysis, integra-
These are structuring of knowledge for application to
tion, and application. The instructors role is to serve
real-world situations, development of reasoning, devel-
only as a facilitator, providing guidance, but not to pro-
opment of effective learning skills, and increase in moti-
vide the solution.
vation for learning (Barrows, 1986).
The procedure involves selection by the instructor
Now often used in the curricula of medical, dental,
of a problem or case that is turned over to students, usu-
and nursing schools, PBL originated at McMaster
ally in small groups, who collectively work on its reso-
University in the mid-1970s (Barrows & Tamblyn,
lution. Typically, students brainstorm the problem,
1980). During the past quarter century, it has been
form initial hypotheses, and evaluate possible solutions
adopted by health professional schools worldwide and
under the guidance of the instructor (who serves as a
is recognized by the World Federation for Medical
resource and will answer specific questions without
Education and the World Health Organization as a valu-
solving the problem for them). Thus, PBL is student-
able educational technique (Walton & Matthews,
centered and encourages students to actively discuss
1989). However, PBL may also be adapted for use in
and attempt to solve issues that are raised by the case
biology courses at all scholastic levels.
presented to them.
Several formats lend themselves to PBL. These
range from small-group discussions to large groups of Experiences with PBL
PBL has been incorporated into several of the lower
and upper division courses in the biology curriculum of
BARRY S. KENDLER, Ph.D., is Professor of Biology at Manhattan
College, Bronx, NY 10471; e-mail: barry.kendler@mountsaint
our department with notable success. We here describe
vincent.edu. PATRICIA A. GROVE, Ph.D., is Professor of Biology at PBL in two different settings. One is in Introductory
College of Mount Saint Vincent, Riverdale, NY 10471-1093. Biology laboratory for non-majors, and the other

348 THE AMERICAN BIOLOGY TEACHER, VOLUME 66, NO. 5, MAY 2004
involves a colloquium for senior biology majors. In
each, the class size was approximately 20 students. The
former involved a problem called The Riddle of the
Dying Kudu, based on information found in Watson
(1987) and the latter is a case called Habitat
Preservation: The Case of Insufficient Funds.
The Riddle of the Dying Kudu
The following typed statement of the problem was
distributed to undergraduates during a laboratory ses-
sion near the end of an Introductory Biology course for
non-majors.
The Riddle of the Dying Kudu*
Background: Severe drought in the early 1980s
in the northwestern Transvaal of South Africa led
to a large increase in mortality of the greater kudu
(Tragelaphus strepsiceros). Despite their weight
of up to 1000 pounds, this antelope is considered
to be highly drought resistant. It can normally
thrive even on the edge of deserts. However, its nor-
mal range of about one antelope per 100 acres had
been reduced by fencing constructed by farmers to
prevent crop damage. Post-mortem examinations
indicated emaciation and death due to malnutri-
tion, despite the presence of leaves in the gut of this
ruminant. However, no disease was found, and the
kudus internal parasite burden was not excessive.
It was also determined that the protein content of
the leaves was adequate, but analysis of the feces
indicated that the protein was not being digested.
Objective: Explain what caused the kudu to die of
malnutrition, despite the presence of adequate food.
*Case written by Barry Kendler
The students are given sufficient time to read the
problem and are then asked to report any terms whose
meaning is unclear. Frequently, post-mortem, emaci-
ation, and ruminant, are troublesome terms. Having
adequately defined these terms, the instructor/tutor
then informs the class that they may ask questions
regarding anything about the problem but they, them-
selves, would have to explain what caused the animals
to die. Following are excerpts from an audiotape of a
session with typical questions asked by students and
the respective answers by the instructor/tutor:
Student: What is meant by their internal parasite
burden was not excessive?
Tutor: All animals in the wild are parasitized to
varying degrees by worms and/or protozoa. In
this case, the parasite levels were within normal
limits, and thus, would not have been the cause
of malnutrition.
S: Were pesticides used by farmers on their crops
responsible for the deaths?
T: No evidence of pesticide involvement was found.
S: Did the antelope die because they were fenced in,
which caused problems with their behavior?
T: Although their range was more limited than pre-
viously, they had sufficient room to move around
fairly large areas. It was not as if they were caged,
as in a zoo.
S: Did the kudu die because they didnt have
enough water to drink?
T: No. They can obtain sufficient water from the
leaves of plants that they consume. A source of
drinking water is not needed by these animals.
S: Did the drought destroy the plants that they nor-
mally ate?
T: Some plants were killed by the drought but oth-
ers survived. Some plant species, especially those
living in arid regions, have roots that penetrate
deep into the earth and can access water even
during drought conditions. In other words, they
are drought-tolerant or drought-resistant.
S: Were the plants that were killed by the drought
those that the kudu normally ate?
T: The kudu feed on leaves of trees that were most-
ly unaffected by the drought.
S: Did the drought affect the digestive system of the
antelopes? Were their digestive enzymes not able
to digest the leaves that were eaten?
T: No. Remember that the walls of plant cells are
made of cellulose and that no mammals possess
the digestive enzymes, called cellulases, needed
to digest plant material.
S: What then is the source of the enzymes?
T: We studied five kingdoms of living organisms.
What are they?
S: Animals, Plants, Protists, Fungi, Monera.
T: OK. Which of these could be the source of
enzymes?
S: Maybe bacteria or protozoa?
T: OK, lets just refer to these organisms collectively
as microbes, which is short for microorganisms.
The digestive tract of ruminants is specialized so
that microbes can live there, and it is these organ-
isms that are responsible for the actual digestion
of food from plant sources.
S: Where did these microbes originally come from?
How did they get into the digestive tract?
PROBLEM-BASED LEARNING 349
 T: They entered the kudus body with milk con-
sumed after birth.
S: Did the microbes die from the drought?
T: Microbes live inside the antelopes gut and for
them there is never any drought. However, the
microbes were affected by something.
S: Was it due to too much heat?
T: No. Remember that all mammals, and birds also,
are homeotherms; that is, they have a relatively
constant body temperature, independent of the
temperature of the outside environment.
S: Did some sort of antibiotic kill the microbes?
T: No, antibiotics were not the culprit, but that is a
good guess.
S: Did pesticides kill the microbes?
T: Pesticides did not kill the microbes but that, too,
is good thinking.
S: Did some other chemical kill them?
T: Yes.
S: Did the chemical come from the soil?
T: No.
S: Did it come from the leaves that were eaten?
T: Yes.
S: Did the trees make the chemical?
T: Yes, but why should trees make the chemical? It
takes a lot of metabolic energy to make chemicals
and natural selection would work against those
plants that wasted their energy making unneces-
sary chemicals. Hence, during the course of evolu-
tion, they would have been eliminated in favor of
species that were less wasteful of their resources.
S: Was the chemical used to protect the trees
because they were being eaten?
T: Yes, but plants are always being eaten. Why
should they make the chemical at this point in
time?
S: Is it because of the drought?
T: No. Drought, by itself, does not cause the tree to
make such chemicals, although other chemicals
that promote survival are made.
S: Was this chemical produced by trees that were
on the other side of the fence?
T: Yes, but in much lower amounts.
S: Was it made by the trees because they were
attacked more often since the kudu had less
space?
350 THE AMERICAN BIOLOGY TEACHER, VOLUME 66, NO. 5, MAY 2004
T: Yes. Exactly. The same trees were attacked more
frequently by the kudu because the animals
range was limited.
T: Now can anyone give the entire solution for the
riddle of the dying kudu?
S: The chemical produced by the plant was respon-
sible for protecting the plants against attack by
the kudu. The microbes were killed and so the
kudu were unable to digest the leaves and they
died.
T: Yes. Now can you identify the chemical? Hint: It
is the same chemical that imparts a bitter taste to
acorns. We mentioned this earlier in the semes-
ter in a different context.
S: Tannin?
T: Yes. Tannins belong to a group of phytochemi-
cals, that is chemicals produced by plants,
known as secondary metabolites. They are specif-
ically formed to repel potential herbivores.
S: But dont squirrels eat acorns?
T: Good observation! The answer is yes, they cer-
tainly do. Apparently they can tolerate the bitter-
ness. However, acorns differ in their tannin con-
tent. Those from white oaks have less tannin
than those from red oaks. Perhaps squirrels pre-
fer eating acorns from white oaks or maybe they
eat such acorns exclusively. How would you
determine which of these possibilities is true?
S: Look it up somewhere?
T: Where would you look it up?
S: The Internet?
T: Perhaps, but what if it wasnt there?
S: A book on squirrels?
T: Same answer as before.
S: Design an experiment?
T: Do it!
S: I would feed squirrels the different kinds of
acorns and see which was eaten and which was
not.
T: OK, that is one way of doing it. Is there another,
more objective way?
S: I would make a platform and I would place a cer-
tain number of acorns from white and red oaks
on the platforms. Then I would observe the
squirrels eating habits to find out if they liked
white or red oak acorns.
T: Very good!
 Following the question and answer session, stu-
dents are given a second printed handout that provides
the solution in greater detail (Figure 1). They are given
time to read it and asked if there are any questions. If so,
the questions are answered.
The entire PBL session can take anywhere from
about one to two hours, depending on the sophistica-
tion and biology background of the class. In all the
many sessions where this problem has been used, stu-
dents eventually solve it on their own. However, in some
cases, hints must be given to prevent the class from
reaching a dead end. Examples of hints in the form of
questions include:
What is necessary for digestion of foods from
plant sources?
How do plant cells differ from animal cells?
What are sources of digestive enzymes?
If you are a tree, how do you protect yourself
from being eaten?
Which substances can affect growth or survival
of microorganisms?
Habitat Preservation: The Case of
Insufficient Funds
As mentioned previously, this case was one of two
used in our senior-level Colloquium for biology majors,
which meets weekly for one hour. Cases written for this
capstone course were designed to integrate information
from core and elective courses in the biology curricu-
lum. During Week 1 of these two-week units, students
analyzed the problem, identified the pertinent informa-
tion they already knew, as well as what they needed to
find out, and allocated tasks within the group. Resource
materials (Figure 2) were placed on reserve in the
library; many students went beyond these to find addi-
tional information. Week 2 was a student-led discussion
of the issues leading to a consensus recommendation
reached by the end of the class session.
Habitat Preservation:
The Case of Insufficient Funds**
Background: The town of Menhaden Bay,
Connecticut lies on the coast of Long Island Sound.
This rural community wants to expand its tax base
to create more jobs and raise money to comply with
a court order to improve its city services. To do this,
the town is negotiating to sell some surrounding
wilderness land. There are three main regions of
approximately equal size.
The first is a salt marsh estuary along the coast.
This area is home to many species of birds, fishes,
Figure 1. Solution to the Riddle of the Dying Kudu
Leaves are digested by microbial enzymes in the rumen. Chemical
analysis of the leaves revealed that they had an excessive amount of
tannins. High tannin levels interfered with normal microbial metabo-
lism, thus culminating in starvation of the kudu.
The elevated tannin levels are attributed to plant defenses against
excessive browsing by the kudu. Under normal conditions, browsing is
limited to less than two minutes per tree. Browsing also occurs infre-
quently since there is about one kudu per 100 acres. Fencing,
designed to limit access of kudu into populated areas and farms
where they were at risk of being shot, had serious consequences dur-
ing the drought, because they were forced to feed more often from
the same trees.
Kudu are vigorous feeders, stripping leaves and bark from branches
and breaking off twigs.Trees respond to this attack within minutes by
increasing their tannin production.This is detected by kudu and they
normally move on to another tree.The combination of fencing and
drought forced the animals to return to their feeding sites at shorter
and shorter intervals. Under these circumstances, tannin concentra-
tions remained high, too high to allow normal microbial function.
Some kudu were impaled on the fences as they unsuccessfully
attempted to reach areas with more palatable plants. Further investi-
gation indicated that undisturbed trees, growing near ones damaged
by browsing, also exhibited an increase in tannin levels, in contrast to
trees that were over 10 feet away. It was hypothesized that a plant
pheromone was released into the air that, when detected by adjacent
trees, stimulated their own tannin production.Tannin production
requires the expenditure of energy by the plant, which explains why
high concentrations are not present at all times. (Watson, 1988).
and arthropods. It is an important nursery for
young scallops, though the local commercial scal-
lop industry is in decline. It also serves as a major
stopping place for migrating waterfowl including
several species of ducks and brant. Amateur
wildlife watchers have seen harbor seals just off the
coast every summer. A local population of sedge
wrens can be found in a grassy marsh that abuts
the meadow. Summer censuses have documented a
small population whose numbers have ranged
from 2-5 nesting pairs during the last three breed-
ing seasons. There are no other documented breed-
ing populations of sedge wrens in Connecticut,
although the species has been reported in New
York and Massachusetts.
The second is a range of hills that is heavily forest-
ed with sugar maples and other hardwood trees. It
contains the nesting sites for several species of rap-
tors, some that hunt in the salt marsh, and others
that hunt in the nearby meadow. Coyotes and a
PROBLEM-BASED LEARNING 351

Figure 2. List of Resources
(On reserve in Library when case was used in 1994)
Blackwell,W.H. 1990. Poisonous and Medicinal Plants.
Cooper, S.R. and G.S. Brush. 1991. Long-term history of Chesapeake
Bay anoxia. Science, 254, 992996.
Foin,T.C. 1976. Ecological Systems and the Environment.
Grossman, J. 1993. How green are these fairways? Audubon, 95(5),
9099.
Leopold, A. 1949. A Sand County Almanac.
McDowell,W.H. 1986. Power Plant Operation on the Hudson River. In:
K. Limburgh, M.A. Moran, and W.H. McDowell (Eds.), The Hudson
River Ecosystem.
McHarg, I.L. 1969. Design With Nature.
National Science Foundation. 1972. Patterns and Perspectives in
Environmental Science.
Newell, R.I.E. 1988. Ecological changes in Chesapeake Bay: Are they
the result of overharvesting the American Oyster, Crassostrea vir-
ginica? Chesapeake Research Consortium Publication, 129, 536-
546.
Odum, H.T. 1983. Systems Ecology: An Introduction.
Pickett, S.A. and P.S.White. 1985. The Ecology of Natural Disturbance
and Patch Dynamics.
Ranwell, D.S. 1972. Ecology of Salt Marshes and Sand Dunes.
Roughgarden, J., R.M. May, and S.A. Levin (Eds.) 1989. Perspectives in
Ecological Theory. (See especially Chaps. 19 21.)
Smith, R.L. 1980. Ecology and Field Biology (3rd ed.).
Strong, D.R. et al. (eds.). 1984. Ecological Communities: Conceptual
Issues and Evidence.
Van Dyne, G.M. (ed.). 1969. The Ecosystem Concept in Natural Resource
Management.
Wilson, E.O. (ed.). 1988. Biodiversity.
small herd of ungulates coexist less than peaceful-
ly. The hills also contain several patches of a beau-
tiful but very rare orchid, the pink ladys-slipper,
Cypripedium acaule. This plant was widely used
in the 19th century as a sedative, and in treatment
of nervous headaches, hysteria, insomnia, and
depression. Heavy collecting of this plant because
of its remarkable therapeutic properties has great-
ly diminished its prevalence in the Northeast.
The third is an expansive but irregularly shaped
meadow region with an abundant population of
voles and other rodents, foxes, and spiders. It is fre-
quented by migrating passerines. It also contains
352 THE AMERICAN BIOLOGY TEACHER, VOLUME 66, NO. 5, MAY 2004
some pitcher plants, and the largest patch in New
England of an endangered species of wildflower
called the Scanlan collar. Several streams run from
the hills through the meadow and into the salt
marsh. Alewives migrate up these streams annual-
ly to spawn.
Objective: You are a member of the Nature
Conservancy, and wish to buy this land to preserve
it in its natural state. Menhaden Bay has agreed to
sell all or part of the land to you, but you only have
enough funds to buy half of it.
There are currently several development proposals
under consideration by the town as well. Assuming
that you can preserve half the land, the remainder
likely will be developed in one of four ways. You
must recommend to the board of directors of the
Nature Conservancy which half of the land to buy,
and thus which development schemes will proceed.
Any other novel but reasonable solutions can be
advocated as well. Be sure to consider both envi-
ronmental preservation and the social impact of
each proposal.
You may assume that the economic value of each
development scheme is essentially the same, and
that the land price per acre is the same in each of
the three habitats.
The students are presented the following proposals:
Proposal 1. Buy the salt marsh and part of the
meadow and hill regions. The remaining portion
of the meadow will become a golf course, while
the hill areas will be converted to a series of sur-
vival game arenas where people play soldier and
shoot paint pellets at each other.
Proposal 2. Buy the hills and half the salt marsh.
The meadow and half the salt marsh will be
cleared for eight-story condominiums to provide
affordable housing for middle class families and
senior citizens.
Proposal 3. Buy the hills and half the meadow. The
salt marsh will be converted to quahog grow-out
pens, and half the meadow will house a new
poultry farm.
Proposal 4. Buy the salt marsh and half the mead-
ow. Connecticut Power will be constructing a
water pump storage station in the hills This sta-
tion is expected to provide competitively-priced
hydroelectric power for coastal Connecticut.
Some pipes and power lines would have to be
run through the remainder of the meadow.
**Case written by Patricia Grove and Michael Judge
 The following objectives for the sessions were estab-
lished:
1. Identify the important species to conserve.
2. Identify habitat(s) important for those species.
3. Determine priorities in environmental conserva-
tion.
4. Balance environmental and social concerns.
5. Appreciate the interaction among species.
6. Appreciate the differences in importance of vari-
ous species for maintaining an ecosystem.
7. Appreciate the role of habitat size and/or degree
of fragmentation in whether or not that habitat
will support a species.
8. Appreciate the impact that development will
have on adjacent areas.
9. Develop negotiating and persuasion skills.
Students were surveyed at the end of the sessions
to allow them to self-assess how well the objectives
were met.
Below are some objectives for the sessions of prob-
lem-based learning that you have just completed.
Indicate for each how well your group satisfied the
objective. Use the scale below for your answers:
4 This objective was extremely well satisfied by
my group.
3 This objective was somewhat satisfied by my
group.
2 This objective was poorly satisfied by my group.
1 This objective wasnt satisfied by my group at
all.
Survey data (meanSD, N=40) from the 1994 collo-
quium cohort show that PBL was an effective way to
meet the learning objectives.
3.5 (0.64) The identification of important species
to conserve.
3.4 (0.56) The identification of habitat(s) impor-
tant for those species that one wants to
conserve.
3.6 (0.49) The determination of priorities in envi-
ronmental conservation.
3.3 (0.54) The balancing of environmental and
social concerns.
3.0 (0.65) An appreciation of the interactions
among species.
3.0 (0.65) An appreciation of the differences in
importance of various species for main-
taining an ecosystem.
2.9 (0.73) An appreciation of the role of habitat
size and/or degree of fragmentation in
whether or not that habitat will support
a species.
3.4 (0.69) An appreciation of the impact that
development will have on adjacent
areas.
3.3 (0.54) The development of negotiating and
persuasion skills.
Students were also surveyed about what they did
and did not like about the PBL sessions. A representa-
tive sample of their comments for this case (1994
cohort) is included below.
What did you like about these sessions of
problem-based learning?
They were challenging and involved us to inquire
further to get answers.
It gave us a chance to actively participate in dis-
cussion which I feel is important.
Gave everyone an opportunity to speak not like
an oral report one person to audience. Better
than listening to speakers. Good teamwork.
The topics were very interesting. The topic con-
cerning conservation was a good reminder of the
threat to rare species and land.
The fact that we were able to encompass all of our
knowledge from various courses to solve these
problems.
It was different/interesting.
Gave student an opportunity to think about and
discuss problems in an open forum environment.
Improved problem-solving skills and group deci-
sion making.
Hearing different viewpoints and arguments that
one necessarily hadnt thought of. Interesting and
informative topics.
Learned how to constructively think out a prob-
lem, weigh both sides, devise a game plan.
I like the fact that individual work effort is evident
even when a group is working on a case.
It was great. It got students to learn more infor-
mation, and feel free to disagree, etc. in a comfort-
able environment. All ideas were taken well. No
one ever felt outcast because of opinions, ideas,
agreements, disagreements. Was well thought-out!
Awesome sessions.
PROBLEM-BASED LEARNING 353
 I felt more intellectual because I wasnt memoriz-
ing the material. It allowed me to use my mind, to
come up with the best possible solutions.
What didnt you like about these sessions of
problem-based learning?
Nothing.
There was not enough preliminary information
given to us.
I didnt like the fact that we didnt find out what
the real outcome was for the conservation topic.
Some things arent as easy to prove as you might
hope or think.
Too short! Not enough time to research and discuss
ideas and proposals.
Case Writing
Questions that may be asked to serve as a guide in
case-writing have been suggested by Hafler (1989). The
case should be written so as to serve one purpose.
Accurate data from a real situation can be woven into a
narrative that outlines a puzzle and thus holds student
interest. Hafler stipulates that cases should have a list of
354 THE AMERICAN BIOLOGY TEACHER, VOLUME 66, NO. 5, MAY 2004
references and/or resources, and objectives or guiding
questions. A note with each case providing additional
detail and background information about the case should
be given to the instructor/tutor, but not to the student.
Prospects
A resource on PBL (Savin-Baden, 2000) and an in-
depth review of that book (Macdonald 2001) have been
published recently. The National Center for Case
Studies Teaching at the University of Buffalo, State
University of New York, is developing a comprehensive
collection of case studies (http://ublib.buffalo.
edu/libraries/projects/cases/ubcase.htm).
Heliker (1994) stated that the role of the educator
today is to probe student thinking toward conceptual
understand, to increase depth of student knowledge, to
provide student learning opportunities, and, by encour-
aging the articulation and contrasting of different points
of view, to allow the emergence of common under-
standings. ... Introduction of PBL into the curriculum
can provide a mechanism to achieve these learning
objectives.
References
Barrows, H. (1986). A taxonomy of prob-
lem-based learning methods.
Medical Education, 20, 481-486.
Barrows, H. S. & Tamblyn, R. Problem-
Based Learning. NY: Springer.
Hafler, J.P. (1989). Case-writing in medical
education: a study at Harvard Medical
School. Ed.D. Thesis, Graduate
School of Education, Harvard
University.
Heliker, D. (1994). Meeting the chal-
lenge of the curriculum revolution:
problem-based learning in nursing
education. Journal of Nursing
Education, 33, 45-47.
Macdonald, R. (2001). Book review.
Studies in Higher Education, 26, 385-
386.
Savin-Baden, M. (2000). Problem-based
learning in higher education: untold
stories. Buckingham, UK: Society for
Research into High Education and
Open University Press.
Walton, H. & Matthews, M (1989).
Essentials of problem-based learn-
ing. Medical Education, 23, 542-558.
Watson, L. (1987). Beyond supernature,
46-50. NY: Bantam Books.