Teaching alveolar ventilation with simple, inexpensive

models
Stephen E. DiCarlo
Advan in Physiol Edu 32:185-191, 2008. doi:10.1152/advan.90156.2008

You might find this additional info useful...

This article cites 22 articles, 12 of which can be accessed free at:

/content/32/3/185.full.html#ref-list-1

This article has been cited by 13 other HighWire hosted articles, the first 5 are:
Using a Physics Experiment in a Lecture Setting to Engage Biology Students with the
Concepts of Poiseuille's Law
Jennifer L. Breckler, Tina Christensen and Wendy Sun
CBE Life Sci Educ 2013; 12 (2): 262-273.
[Abstract] [Full Text] [PDF]

Combination of didactic lectures and review sessions in endocrinology leads to

improvement in student performance as measured by assessments
Ayisha Qureshi, Cassy Cozine and Farwa Rizvi
Advan in Physiol Edu, March , 2013; 37 (1): 89-92.
[Abstract] [Full Text] [PDF]

Learning by doing: construction and manipulation of a skeletal muscle model during

lecture

Downloaded from on April 7, 2015

David W. Rodenbaugh, Heidi L. Lujan and Stephen E. DiCarlo
Advan in Physiol Edu, December , 2012; 36 (4): 302-306.
[Abstract] [Full Text] [PDF]

A simple model to demonstrate the balance of forces at functional residual capacity

Praghalathan Kanthakumar and Vinay Oommen
Advan in Physiol Edu, June , 2012; 36 (2): 170-171.
[Full Text] [PDF]

Does sex (female versus male) influence the impact of class attendance on examination
performance?
Ronald N. Cortright, Heidi L. Lujan, Julie H. Cox and Stephen E. DiCarlo
Advan in Physiol Edu, December , 2011; 35 (4): 416-420.
[Abstract] [Full Text] [PDF]

Updated information and services including high resolution figures, can be found at:
/content/32/3/185.full.html

Additional material and information about Advances in Physiology Education can be found at:
http://www.the-aps.org/publications/advan

This information is current as of April 7, 2015.

Advances in Physiology Education is dedicated to the improvement of teaching and learning physiology, both in specialized
courses and in the broader context of general biology education. It is published four times a year in March, June, September and
December by the American Physiological Society, 9650 Rockville Pike, Bethesda MD 20814-3991. Copyright © 2008 by the
American Physiological Society. ISSN: 1043-4046, ESSN: 1522-1229. Visit our website at http://www.the-aps.org/.
Adv Physiol Educ 32: 185–191, 2008;
doi:10.1152/advan.90156.2008.
Teaching alveolar ventilation with simple, inexpensive models
Stephen E. DiCarlo
Department of Physiology, Wayne State University School of Medicine, Detroit, Michigan
Submitted 23 June 2008; accepted in final form 7 August 2008
DiCarlo SE. Teaching alveolar ventilation with simple, inexpen-
sive models. Adv Physiol Educ 32: 185–191, 2008; doi:10.1152/
advan.90156.2008.—When teaching and learning about alveolar ven-
tilation with our class of 300 first-year medical students, we use four
simple, inexpensive “models.” The models, which encourage re-
search-oriented learning and help our students to understand complex
ideas, are distributed to the students before class. The students
anticipate something new every day, and the models provide elements
of surprise and physical examples and are designed to help students to
understand 1) cohesive forces of the intrapleural space, 2) chest wall
and lung dynamics, 3) alveolar volumes, and 4) regional differences in
ventilation. Students are drawn into discussion by the power of
learning that is associated with manipulating and thinking about
objects. Specifically, the models encourage thinking about complex
interactions, and the students appreciate manipulating objects and
actually understanding how they work. Using models also allows us to
show students how we think as well as what we know. Finally,
students enjoy taking the models home to demonstrate to friends and
family “how the body works” as well as use the models as future study
aids.
pulmonary compliance; intrapleural cohesive forces; relaxation curves
HOW WE TEACH is much more important than what we teach
because the traditional “lecture then test” format and accom-
panying “cookbook” laboratories discourage a deep under-
standing of the subject as well as the development of lifelong
skills such as critical thinking, problem solving, communica-
tion, and interpersonal skills (5). This is important because in
many schools, the classes are teacher centered rather than
dynamic student-centered experiences that engage students in
research-oriented learning. In this setting, the teacher perpetu-
ates the process by transferring knowledge to students through
lectures and note taking rather than active involvement and
personal investment in the process (5). Teachers can encourage
research-oriented learning by carefully considering the type
and organization of information as well as the instructional
strategies we use (5). For example, we all realize that reading
the recipe is not as useful as preparing the meal when learning
how to cook. How, then, do we change our approach from
reading to “doing” (5)?
Simple, inexpensive models encourage research-oriented
learning and are often used to explain complex ideas because
models promote logic, reasoning, and creativity (1, 2, 6 –9, 15,
17–19). Physical models relate the unknown to the familiar and
provide a new perspective on information gathering. Models
also encourage a see-touch interaction to supplement new
information processing.
Active participation with physical models can reach all types
of learners via the visual, auditory, kinesthetic, and tactile
Address for reprint requests and other correspondence: S. E. DiCarlo,
Wayne State Univ. School of Medicine, 540 E. Canfield Ave., Detroit, MI
48201 (e-mail: sdicarlo@med.wayne.edu).
1043-4046/08 $8.00 Copyright © 2008 The American Physiological Society
Refresher Course
schemes of learning (12, 21, 26), and manipulation of models
is supported by constructivism (24). Constructivist learning
theory is based on the idea that knowledge is actively con-
structed by the learner. Social constructivism suggests that
knowledge creation is a shared experience rather than an
individual experience (14). In this context, the important part is
not that students manipulate things physically but that they do
it for a purpose and engage in discussion about it (24). This is
important because learning is a social process as well as an
individual process (4).
When teaching and learning about alveolar ventilation with
our class of 300 first-year medical students, we use four simple,
inexpensive models. The models encourage research-oriented
learning and help our students understand complex ideas. The
models are distributed to the students before class and are
designed to help students to understand 1) cohesive forces of
Downloaded from on April 7, 2015
the intrapleural space (Figs. 1 and 2), 2) chest wall and lung
dynamics (Figs. 3 and 4), 3) alveolar volumes (Fig. 5), and
4) regional differences in ventilation (Fig. 6). Students are
drawn into discussion by the power of learning that is associ-
ated with these experiences. The models encourage thinking
about the complex interactions, and students appreciate manip-
ulating objects and actually understanding how they work.
Using models also allows us to show students how we think as
well as what we know. Finally, students enjoy taking the
models home to demonstrate to friends and family “how the
body works” as well as use the models as future study aids.
We begin teaching about alveolar ventilation by asking
“How does air get into the lungs?” Students are told that air
flows from a region of higher pressure to a region of lower
pressure by bulk flow. Accordingly, when alveolar pressure
(PA) is the same as the atmospheric pressure [barometric
pressure (PB)], no air flow occurs, because no pressure gradi-
ent exists.
Thus, to initiate air flow into gas exchange sites of the lung,
PA must decrease below PB, or PB must increase above PA, as
occurs during mechanical ventilation. To expel gases from
alveoli, PA must exceed PB. With normal breathing, PA must
change because PB does not fluctuate on a minute-to-minute
basis.
How does PA decrease below PB? When the inspiratory
muscles contract, the thoracic cavity and lungs enlarge, which
decreases PA. Air then moves into the alveoli by bulk flow until
PA and PB equalize at end inspiration.
Specifically, when the diaphragm contracts, its insertion is
pulled toward its origin, flattening the dome, increasing the
vertical dimension of the thoracic cavity (the thoracic cavity
and lungs enlarge). Increasing the volume of the thoracic
cavity reduces PA below PB. This concept is explained by
Boyle’s law [Boyle’s law (P1V1 ⫽ P2V2)], which describes the
inverse relationship between pressure (P) and volume (V; gas
at a constant temperature). Increasing thoracic volume causes
fewer collisions between gas molecules and between gas mol-
185
Refresher Course
186 STAYING CURRENT IN RESPIRATORY PHYSIOLOGY
Fig. 1. The respiratory system can be modeled as an air-filled balloon sur-
rounded by a larger fluid-filled balloon. The outside of the outer fluid filled
balloon represent the atmosphere. The small liquid-filled compartment be-
tween the two balloons represents the pleural space and intra-pleural pressure.
The air-filled compartment inside the inner air-filled balloon represents the
alveoli and conducting airways of the lungs and intra-alveolar pressure.
ecules and the thorax. The lower number of collisions de-
creases the pressure exerted by the gas, which reduces PA
below PB so air flows into the lungs until PA again equals PB
at end inspiration.
Once air enters the lungs, how does air get out of the lungs?
The lung is stretched during inspiration, so when the inspira-
tory muscles relax, the lung recoils to compress the alveolar
gas volume. This elevates PA above PB so air is expelled
(expired) until PA again equals PB at end expiration. Specifi-
cally, when the diaphragm relaxes, its elasticity causes it to
return to its dome shape, decreasing the vertical dimension of
the thoracic cavity. Again, the lung is stretched during inspi-
ration, so when the inspiratory muscles relax, the lung recoils
to compress the alveolar gas volume. This elevates PA above
PB so air is expelled (expired) until PA again equals PB at end
expiration.
Importantly, this explanation, although correct, requires fur-
ther elaboration (and this is when we introduce the model).
Specifically, how does increasing the vertical dimension of the
thoracic cavity expand the lungs?
To answer this question, students are told that the respiratory
system can be modeled as an air-filled balloon surrounded by
a larger water-filled balloon (10). The outside of the outer
Fig. 2. The model used to facilitate the understanding of the cohesive forces
of the intrapleural space consists of two microscope slides with a few droplets
of water (joined oval structures) placed between them. The slides move easily
over one another horizontally; however, it is very difficult to pull them apart.
Advances in Physiology Education • VOL
Downloaded from on April 7, 2015
Fig. 3. The bony, moderately flexible ribcage can be modeled by cutting the
corners of a wire coat hanger (top, the dashed lines indicate the location for
cutting the hanger into two chest cage models; note that two models are
obtained from each hanger). The model used to simulate the lung is a rubber
band (bottom, shaded circle). The lungs inside the chest cavity, with the
respective pleural surfaces held together by cohesive forces, are simulated by
placing the rubber band around the arms of the coat hanger (1). The volume of the
lung is higher than its equilibrium volume as simulated when the rubber band is
stretched open (2). Chest cavity volume is less than its equilibrium volume as
simulated when the ends of the hanger are compressed (3). The equilibrium
volume of the combined lung-chest cage represents a point [functional residual
capacity (FRC)] where the tendency of the lung to deflate (rubber band to collapse,
inward pointing arrows) is balanced by the tendency of the chest cage to expand
(hanger to spring open, outward pointing arrow) (4). These equal but opposite
recoil forces create an intrapleural pressure that is below atmospheric pressure.
Furthermore, as a person inhales to expand the chest cage and lung, intrapleural
pressure becomes increasingly subatmospheric as a consequence of the force
tending to separate the lung and chest wall.
water-filled balloon represent the atmosphere. The small liq-
uid-filled compartment between the two balloons represents the
pleural space, and the air-filled compartment inside the inner
air-filled balloon represents the alveoli and conducting airways
of the lungs (Fig. 1) (10, 20). The pressures in these three
compartments are referenced to each other by making the
atmospheric pressure (outside the chest wall) equal to 0
mmHg. Before the start of inspiration, the pressure inside the
alveoli (intra-alveolar pressure) is also 0 mmHg, and the
intrapleural pressure is ⫺4 mmHg.
What creates the negative intrapleural pressure? The fluid-
filled balloon surrounding the air-filled balloon forms a double
membrane similar to the pleural sac. Similarly, the outer
surface of the lung is lined with a membrane called the visceral
32 • SEPTEMBER 2008

STAYING CURRENT IN RESPIRATORY PHYSIOLOGY
Fig. 4. Top: diagram of the lung and chest cage. Arrows show the movement
of the chest cage and lung. Bottom: Separate relaxation curves for the lung
(right) and chest cage (left) along with the combined lung-chest cage relaxation
curve (middle). The combined lung-chest cage curve is the algebraic sum of the
separate lung and chest cage curves. The slope of each relaxation curve
corresponds to the compliance for the structure(s). At end expiration (point A),
recoil or relaxation pressure for the lung and chest cage alone are equal but
opposite. At this point, lung volume corresponds to FRC. As additional air
volume is inhaled into the lung, the lung is stretched further and exhibits a
greater recoil pressure. At the same time, the chest cage is less compressed, so its
negative recoil pressure diminishes as it approaches its equilibrium volume. When
a slightly larger air volume is inhaled, the chest cage reaches its equilibrium
volume (0-mmHg relaxation pressure, point B), and the lung and lung-chest
relaxation curves intersect (point C). Thereby, at this lung volume, all measured
relaxation for the lung-chest cage system is from the lung because the chest cage
is at its equilibrium volume (point B), or the volume it would assume if the lung
were not present. If an even greater air volume is inhaled (point D), both the
lung and chest cage are stretched beyond their equilibrium volumes. Note that
the compliance curve for the combined lung-chest cage becomes more flat-
tened (less compliant) at this point because the lung and chest cage are both
tending to recoil toward smaller equilibrium volumes. If the total lung-chest
cage system is returned to resting end expiration (point A) and air is expelled,
a negative relaxation pressure results for both the chest cage and combined
lung-chest cage (point E). At this point, the chest cage is compressed as more
and more air is expelled, with the negative recoil pressure resulting from the
tendency of the chest to expand toward its equilibrium volume (point B). At
the same time, the lung contributes little positive relaxation pressure
because it is close to its equilibrium volume (i.e., 0-ml volume) because it
is stretched very little. Students are encouraged to manipulate their models
as they discuss thes relationships with the instructor. [From the Integrated
Medical Curriculum (6a)].
Advances in Physiology Education • VOL
Refresher Course
187
Downloaded from on April 7, 2015
Fig. 5. Alveolar ventilation is the volume of fresh air introduced into the
gas-exchanging regions of the lungs per minute. Top: the model consists of a
paper cutout of the anatomic dead space with the alveoli as well as two
columns of paper, each with four segments, with each segment representing
150 ml of air. One column has three dark blue segments and one light blue
segment, and the second column has three light blue segments and one dark
blue segment. Dark blue segments represent atmospheric (fresh air), and light
blue segments represent alveolar air (air that has previously equilibrated with
pulmonary artery blood). The anatomic dead space and columns of air are sized
such that one of the 150-ml segments fits perfectly into the anatomic dead
space while the remaining three segments fit perfectly into the alveoli (bottom).
pleural, and the inside of the thorax is lined with a membrane
called the parietal pleura. These pleural membranes juxtapose
to form a pleural sac around the lung. The space within the
pleural sac contains a few milliliters of fluid. The intrapleural
fluid creates cohesive forces that hold the visceral and parietal
pleura together. The cohesive forces opposes the tendency of
the ribcage to expand (increase in size) and the tendency of the
lungs to collapse (decrease in size) (10, 20). Opposing these
directionally opposite forces creates the negative intrapleural
pressure.
Intrapleural fluid also creates a slippery surface allowing the
lungs to slide within the chest against the thoracic wall and,
when the chest expands during inspiration, the lungs are
compelled to follow so that the lungs and chest expand as a
single unit.
This concept and physiological significance of the cohesive
forces of the intrapleural space are often difficult for students
to grasp. To help students understand this concept, we and
many others use a simple model (19). The model used to
facilitate the understanding of this concept consists of two
32 • SEPTEMBER 2008
Refresher Course
188 STAYING CURRENT IN RESPIRATORY PHYSIOLOGY
Fig. 6. A Slinky hanging from the ceiling can be used to model the effects of
gravity on regional differences in ventilation. The coils of the Slinky closest to
the ceiling are wide apart, stretched, and therefore on the end of the compliance
curve (it is difficult to stretch further). In sharp contrast, the coils at the bottom
are very close together, compressed, and therefore on the steep part of the
compliance curve (it is easy to stretch). If a subject starts inspiration from FRC,
apical alveoli will start off at a larger volume than the basal ones; thus, apical
alveoli will be less able to accommodate any increase in volume (i.e., they are
less compliant than basal alveoli). Thus, if inspiration starts from FRC, most
of the incoming air goes preferentially to basal alveoli, which started off less
distended and thus more compliant. A Slinky is a deformable spring distorting
under its own weight. (Slinky is a registered trademark of Poof-Slinky.)
microscope slides with a few droplets of water placed between
them (Fig. 2). The slides move easily over one another hori-
zontally; however, it is very difficult to pull them apart.
Similarly, intrapleural fluid creates a slippery surface allowing
the lungs to slide within the chest against the thoracic wall. The
intrapleural fluid also creates cohesive forces that hold the
visceral and parietal pleura together. Thus, when the chest
expands during inspiration, the lungs are compelled to follow
so that the lungs and chest expand as a single unit.
Lung and Chest Wall Dynamics Model
The lung and chest wall dynamics model is based on that
described in Refs. 22 and 23. Once this concept (i.e., lung and
chest wall are held together by cohesive forces) is fully
understood and demonstrated with the model, it becomes
necessary to discuss the complex interactions between the lung
and chest wall.
To do this, students are shown that the bony, moderately
flexible chest cage is analogous to a punctured tennis ball (at
this point tennis balls, previously cut in half, are tossed into the
audience). Because the flexible ribcage is compressible but
does not collapse, the chest cavity has considerable air volume
when inside and outside pressure are equal, or transmural
pressure is 0 mmHg. The model used to simulate the bony,
moderately flexible ribcage consists of the corners of a wire
coat hanger (Fig. 3, top; note that two models are obtained
from each hanger). When no forces are acting on the coat
hanger, its equilibrium volume can be represented by the area
of the triangle. Of course, the volume can be increased by
pulling out on the arms of the corners of the coat hanger and
Advances in Physiology Education • VOL
decreased by pushing in on the arms of the corners of the coat
hanger (Fig. 3, bottom).
In contrast, students are told that if the lung is removed from
the thoracic cage, it closely resembles a collapsed balloon
(balloons are distributed to the students). When the pressure
inside the lung equals outside pressure, or transmural pressure
is 0 mmHg, lung volume is close to 0. The model used to
simulate the lung is a rubber band (Fig. 3, bottom). When no
forces are acting on the rubber band, its equilibrium volume
(represented by the area of the rubber band) is close to 0.
However, the lung, like the rubber band, can be stretched, and
the volume (area of the circle) increased.
When the rubber band is placed around the two arms of the
corners of the coat hanger, the triangular area between the two
arms of the corner of the coat hanger and the rubber band can
be used to represent the volume of air in the lungs at a given
point in time. Furthermore, the inwardly directed recoil force of
the rubber band opposes the outwardly directed recoil force of the
coat hanger (Fig. 3, bottom). When there are no forces acting on
this model, the inwardly and outwardly directed recoil forces
reach a balance, such that the inwardly directed force of the rubber
band is equal and opposite to the outwardly directed force of the
coat hanger. This point is analogous to the end of a quiet
exhalation. The volume of air left in the lungs at the balance point
Downloaded from on April 7, 2015
is equal to the functional residual capacity (FRC).
At this point, contraction of the muscles of inspiration
causes the chest wall to expand and the volume of the lungs
increases. Similarly, pulling out on the arms of the hanger
stretches the rubber band (increasing the volume) as well as
increasing the inwardly directed recoil force of the rubber band
while simultaneously decreasing the outwardly directed recoil
force of the hanger (Fig. 3). Relaxation of the muscles of
inspiration allows the increased recoil force of the lungs to
drive quiet (passive) exhalation. Similarly, releasing the force
on the arms of the hanger allows the model to return to its
equilibrium position.
Thus, with the lungs placed inside the chest cavity and their
respective pleural surfaces held together by cohesive forces
(the rubber band placed around the arms of the coat hanger;
Figs. 3 and 4), the volume of the lung is higher than its
equilibrium volume (the rubber band is stretched open),
whereas chest cavity volume is less than its equilibrium vol-
ume (the ends of the hanger are compressed). Again, the
equilibrium volume of the combined lung-chest cage repre-
sents a point at which the tendency of the lung to deflate
(rubber band to collapse) is balanced by the tendency of the
chest cage to expand (hanger to spring open). Specifically, at
the equilibrium volume of the combined lung-chest cage, the
lung (rubber band) is expanded above and the chest cage
(hanger) is compressed below their respective equilibrium
volumes (Fig. 3, bottom). The equilibrium volume of the
combined lung-chest cage corresponds to resting end expira-
tion, or the position (volume) the combined lung-chest cage
would assume when the respiratory muscles are relaxed. At
resting end expiration, the recoil pressure of the lung tending to
deflate (rubber band collapse) is opposed to an equal but
opposite recoil pressure of the chest wall tending to expand
(hanger springing out; Fig. 3). These equal but opposite recoil
forces create an intrapleural pressure that is below atmo-
spheric pressure. Specifically, a “tug of war” between the lung
and chest wall increases the intrapleural space and decreases
32 • SEPTEMBER 2008

STAYING CURRENT IN RESPIRATORY PHYSIOLOGY
the intrapleural pressure (13). Furthermore, as a person inhales
to expand the chest cage and lung, intrapleural pressure be-
comes increasingly subatmospheric as a result of the force
tending to separate the lung and chest wall.
The hanger and rubber band model can be used to under-
stand changes that occur with obstructive and restrictive pul-
monary diseases. For example, many restrictive pulmonary
diseases result in a decreased compliance of the lung and/or
chest. To model this condition, simply add a second or thicker
rubber band to the hanger (22, 23). This situation reduces
equilibrium volume, makes inspiration more difficult, and
requires more muscular effort to move the chest wall (spread
the arms of the hanger) and inflate the lungs (stretch the rubber
bands).
Some obstructive pulmonary diseases result in an increased
compliance of the lungs. This can be demonstrated by using a
thinner rubber band than was used for the normal state (22, 23).
With the thinner rubber band, inspiration is easier because the
lungs are more compliant. In addition, the inwardly directed
recoil force of the lungs, used to drive exhalation, is reduced,
resulting in a prolonged expiration. Finally, a “new” equilib-
rium point is found between the rubber band (lungs) and the
hanger.
We have found that using this model during discussion of
the separate relaxation curves for the lung and chest cage, as
well as the combined lung-chest cage relaxation curve (Fig. 4),
greatly enhances the student’s understanding (parenthetically,
how many of our students really reach an understanding of
these curves?). Students are encouraged to manipulate the
model as the instructor describes the specific components
shown in Fig. 4.
Alveolar Volume Model
Once the dynamics of the lung and chest wall are under-
stood, we ask our students, “What is alveolar ventilation?” As
you know, alveolar ventilation is the volume of fresh air
introduced into the gas-exchanging regions of the lungs per
minute. This concept, seemingly straightforward, is con-
founded and confused by two important features: 1) anatomic
dead space and 2) “fresh” air.
To help unconfound and unconfuse this concept, students
are told that at the end of a normal expiration (just before the
next inspiration) the conducting airways (anatomic dead space)
are filled with alveolar gas. Thus, as a tidal inspiration begins,
the alveoli must first receive the gas that was in the anatomic
dead space from the last exhalation. This gas does not raise
alveolar PO2 or lower alveolar PCO2 very much because it has
the same composition as the alveolar gas. After the dead space
volume is inspired, the alveoli receive fresh air until the tidal
volume is completed. The last portion of the fresh air, of
course, remains in the conducting airways.
The model used to demonstrate this is shown in Fig. 5,
which shows a paper cutout of the anatomic dead space with
the alveoli as well as two columns of paper with four segments,
with each segment representing 150 ml of air (top). As shown
in Fig. 5, top, one column has three 150-ml segments (dark
blue boxes) and one 150-ml segment (light blue box), and the
second column has three 150-ml segments (light blue boxes)
and one 150-ml segment (dark blue box). The dark blue
segments represent atmospheric (fresh air) and light blue seg-
ment represent alveolar air (air that has previously equilibrated
Advances in Physiology Education • VOL
Refresher Course
189
with pulmonary artery blood) (Fig. 5, top). The anatomic dead
space and columns of air are sized such that one of the 150-ml
segments fits perfectly into the anatomic dead space while the
remaining three segments fit perfectly into the alveoli (Fig. 5,
bottom).
To use the model, we start at the end of a normal expiration
(just before the next inspiration) when the conducting airways
are filled with alveolar gas. At this point, the students place the
column of air with one light blue segment into the anatomic
dead space and the three dark blue segments projected into the
atmosphere. Thus, as a tidal inspiration begins (and the stu-
dents slide the column of air into the alveoli), the alveoli must
first receive the gas that was in the anatomic dead space from
the last exhalation (alveolar air, light blue segment). After the
dead space volume is inspired, the alveoli receive fresh air
(atmospheric air, dark blue segments) until the tidal volume is
completed. The last portion of the fresh air, of course, remains
in the anatomic dead space (dark blue segment). Thus, three
segments of fresh air (450 ml) entered the anatomic dead
space, but only two segments of fresh air entered the alveoli.
Since alveolar volume is only the volume of fresh air entering
the alveoli, alveolar volume is 300 ml. That is, during inspi-
ration, tidal volume is 450 ml. Since the dead space is 150 ml,
this means that the alveolar volume (the volume of fresh air
Downloaded from on April 7, 2015
reaching the airspaces) is 300 ml.
At the end of a normal inspiration (just before the next
expiration), the conducting airways are filled with fresh atmo-
spheric air. At this point, the students place the column of air
with one dark blue segment (fresh atmospheric air) into the
anatomic dead space and the three light blue segments into the
alveolus (old alveolar air; Fig. 5, bottom). Thus, as a tidal
expiration begins (and the students slide the column of air into
the atmosphere), the atmosphere must first receive the gas that
was in the anatomic dead space from the last inspiration (fresh
atmospheric air, dark blue segment). After the dead space
volume is expired, the atmosphere receives alveolar air (air that
had previously exchanged, light blue segments) until the tidal
volume is completed. The last portion of the alveolar air, of
course, remains in the anatomic dead space (light blue seg-
ment). Thus, three segments of alveolar air (450 ml) entered
the anatomic dead space, but only two segments of alveolar air
entered the atmosphere. Although this concept is confounded
by the anatomic dead space and confused (not necessarily
complex) because of the “fresh” air concept, the model signif-
icantly helps our students grasp the concepts.
Distribution of Alveolar Ventilation Model
Up to this point, we have air entering and exiting the lungs;
however, where does it go? To answer this question, we
discuss the distribution of alveolar ventilation and the effect of
gravity. The effect of gravity on the distribution of ventilation
within the lung is very important since this has implications for
the matching of ventilation to perfusion in both normal and
pathological lungs as well as during one-lung ventilation for
thoracotomy procedures.
Therefore, we emphasize that, because of gravity, there are
unique compliance characteristics for different alveoli ranging
from the base to the apex of the upright lung. The alveoli at the
top of the lung are ventilated less than those at the bottom. The
reason is that the alveoli at the top are on a less compliant part
of the compliance curve, e.g., they are already “stretched,” and,
32 • SEPTEMBER 2008
Refresher Course
190 STAYING CURRENT IN RESPIRATORY PHYSIOLOGY
for any given pressure change, they will expand less than their
counterparts at the base.
At this point, it is important to remind students that the
directionally opposite recoil forces of the lung and chest wall
(the tug of war) create an intrapleural pressure that is below
atmospheric pressure [the tug of war between the lung and
chest wall increases the intrapleural space and decreases the
intrapleural pressure (13)]. Furthermore, as a person inhales to
expand the chest cage and lung, intrapleural pressure becomes
increasingly subatmospheric as a result of the force tending to
separate the lung and chest wall. Thus, because the alveoli at
the top of the lung are stretched (greater force tending to
separate the lung and chest wall), the intrapleural pressure is
more negative at apical alveoli and transmural pressure (intra-
alveolar minus intrapleural pressure) is greater due to an
expanded apical alveoli.
To model this concept, the students are given a small Slinky
(Fig. 6) and are told that the lungs are like a Slinky, hanging
from the ceiling (11). The coils closest to the ceiling are wide
apart, already stretched, and therefore on the end of the
compliance curve. In contrast, the coils at the bottom are very
close together, compressed, and therefore on the steep part of
the compliance curve. The students are encouraged to consider
how this effects regional distribution of air flow during inspi-
ration.
Specifically, because of the way in which the lungs are
suspended in the chest cavity and are subjected to gravity, a
gradient in pleural pressure exists from the apex to the base of
⬃7.5 cmH2O. Intrapleural pressure surrounding apical alveoli
is more negative than intrapleural pressure surrounding basal
alveoli. Thus, transpulmonary pressure is greater for apical
alveoli because these alveoli are “stretched” open. If a subject
starts inspiration from FRC, apical alveoli will start off at a
larger volume than basal alveoli due to the more negative
intrapleural pressure; thus, apical alveoli will be less able to
accommodate any increase in volume (i.e., they are less com-
pliant than basal alveoli). Thus, if inspiration starts from FRC,
most of the incoming air goes preferentially to basally located
alveoli, which started off less distended and thus more com-
pliant.
The Slinky perfectly illustrates this concept because the coils
at the top are already stretched, and, for any given pressure
change, these coils will expand less than their counterparts at
the base, which are not stretched. In addition, using this model,
students are able to see the changes in FRC associated with the
supine position.
Discussion
When teaching and learning about alveolar ventilation with
our class of 300 first-year medical students, we use four simple,
inexpensive models. We (my graduate students and I) construct
the models and place the models into plastic bags. I believe that
this is an important lesson for graduate students. Our graduate
students must understand that teaching is no less difficult than
research (25) and classes must be planned as carefully and
thoroughly as experiments. This includes reading the educa-
tional research, designing and testing equipment (materials),
and planning activities. The models are distributed to the
students before class, and these activities are conducted in a
very large lecture room. The students expect something new
every day, which generates energy, excitement, and motiva-
Advances in Physiology Education • VOL
tion. The models encourage thinking about the complex inter-
actions, and students appreciate manipulating the objects and
actually understanding how they work. Using models also
allows us to show students how we think as well as what we
know. Finally, students enjoy taking the models home to
demonstrate to friends and family “how the body works” as
well as use the models as future study aids.
We believe that it is important to plan an activity for students
because activities engage the audience, build curiosity and
create a challenge. That is, learning is not a spectator sport, and
students are not vessels passively waiting to be filled with a
predetermined body of knowledge. Students do not learn by
simply sitting in a classroom listening to the teacher, memo-
rizing prepackaged assignments, and spitting out answers (3).
Students learn when they are actively involved in learning (16).
Students must do more than just listen: they must read, write,
discuss, and be engaged in solving problems (3). Remember,
most of the learning occurs outside the classroom when stu-
dents read, reflect, write, or experience the information. Mod-
els support these activities as well as provide elements of
surprise and concrete examples.
REFERENCES
Downloaded from on April 7, 2015
1. Chan V, Pisegna J, Rosian R, DiCarlo SE. Model demonstrating
respiratory mechanics for high school students. Adv Physiol Educ 15:
1–18, 1996. [Erratum. Adv Physiol Educ 17: 58 –58, 1997.]
2. Chan V, Pisegna JM, Rosian RL, DiCarlo SE. Construction of a model
demonstrating neural pathways and reflex arcs. Adv Physiol Educ 16:
14 – 42, 1996.
3. Chickering AW, Gamson ZF. Seven Principles for Good Practice in
Undergraduate Education (online). http://www.csuhayward.edu/wasc/
pdfs/End%20Note.pdf [11 August 2008].
4. Committee on Developments in the Science of Learning, Committee
on Learning Research, and Educational Practice and the National
Research Council. How People Learn: Brain, Mind, Experience and
School. Washington, DC: National Academy, 2000.
5. DiCarlo SE. Cell biology should be taught as science is practised. Nat Rev
Mol Cell Biol 7: 290 –296, 2006.
6. DiCarlo SE, Sipe E, Layshock JP, Varyani S. Experiment demon-
strating skeletal muscle biomechanics. Adv Physiol Educ 20: 59 –71,
1998.
6a.DxR Development Group. Integrated Medical Curriculum Homepage
(online). http://imc.meded.com/ [11 August 2008].
7. Giuliodori MJ, DiCarlo SE. Myelinated vs. unmyelinated nerve conduc-
tion: a novel way of understanding the mechanisms. Adv Physiol Educ 28:
80 – 81, 2004.
8. Giuliodori MJ, DiCarlo SE. Simple, inexpensive model spirometer for
understanding ventilation volumes. Adv Physiol Educ 28: 33, 2004.
9. Giuliodori MJ, DiCarlo SE. An improved model for simulating obstruc-
tive lung disease. Adv Physiol Educ 32: 167, 2008.
10. Goodman BE. Pulmonary and renal pressure-flow relationships: what
should be taught? Adv Physiol Educ 25: 15–28, 2001.
11. Hopkins SR, Henderson AC, Levin DL, Yamada K, Arai T, Buxton
RB, Prisk GK. Vertical gradients in regional lung density and perfusion
in the supine human lung: the Slinky effect. J Appl Physiol 103: 240 –248,
2007.
12. Lujan HL, DiCarlo SE. First-year medical students prefer multiple
learning styles. Adv Physiol Educ 30: 13–16, 2006.
13. Maron MB. Relating basic concepts of pulmonary mechanics to clinical
situations. Adv Physiol Educ 273: 85–95, 1997.
14. Prawat RS, Floden RE. Philosophical perspectives on constructivist
views of learning. Educ Psychol 29: 37– 48, 1994.
15. Raghavan K, Glaser R. Model-based analysis and reasoning in science:
the MARS curriculum. Sci Educ 79: 37– 61, 1995.
16. Rao SP, DiCarlo SE. Active learning of respiratory physiology improves
performance on respiratory physiology examinations. Adv Physiol Educ
25: 55– 61, 2001.
32 • SEPTEMBER 2008
 Refresher Course
STAYING CURRENT IN RESPIRATORY PHYSIOLOGY 191

17. Rodenbaugh DW, Collins HL, Chen CY, DiCarlo SE. Construction of
a model demonstrating cardiovascular principles. Adv Physiol Educ 22:
67– 83, 1999.
18. Rodenbaugh DW, Collins HL, DiCarlo SE. Spirometry: simulations of ob-
structive and restrictive lung diseases. Adv Physiol Educ 26: 222–223, 2002.
19. Rodenbaugh DW, Collins HL, DiCarlo SE. A simple model for under-
standing cohesive forces of the intrapleural space. Adv Physiol Educ 27:
42– 43, 2003.
20. Sherwood L. Human Physiology: From Cells to Systems. Belmont, CA:
Wadsworth, 1997.
21. Slater JA, Lujan HL, DiCarlo SE. Does gender influence learning style pref-
erences of first-year medical students? Adv Physiol Educ 31: 336–342, 2007.
22. Stockert B. Pulmonary ventilation teaching aid. Adv Physiol Educ 27:
41– 42, 2003.
23. Stockert B. Pulmonary ventilation teaching aid: part 2. Adv Physiol Educ
27: 86 – 87, 2003.
24. Tobin K. The Practice of Constructivism in Science Education. Washing-
ton, DC: AAAS, 1993.
25. Vander AJ. The Claude Bernard Distinguished Lecture. The excitement
and challenge of teaching physiology: shaping ourselves and the future.
Adv Physiol Educ 12: 3–16, 1994.
26. Wehrwein EA, Lujan HL, DiCarlo SE. Gender differences in learning
style preferences among undergraduate physiology students. Adv Physiol
Educ 31: 153–157, 2007.

Downloaded from on April 7, 2015

Advances in Physiology Education • VOL 32 • SEPTEMBER 2008