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Transcript
INNOVATIONS
TEACHING
THE
A
NERVE
CONDUCTION
“TRAVELING
FLAME”
N
D
I
D
E
A
S
TO UNDERGRADUATES:
ANALOGY
REVISITED
Sabyasachi S. Sircar and Om P. Tandon
Department
of Physiology,
University
College
of Medical
FLAME
A popular textbook of physiology (1) summarizes the
analogy as follows: “A cigarette is a cylindrical structure filled with material at chemical potential which
VOLUME
15 : NUMBER
1 -ADVANCES
110095,
India
In our introductory classeson nerve-muscle physiology, we present to our students the core concepts
outlined above after elaborating them as follows.
When one end of a rod is held over the flame, there are
two ways in which the heat can get transferred to the
other end. If the rod is a good conductor of heat, then
in a very short time, the other end of the rod gets
heated up due to transfer of heat through conduction.
If, on the other hand, the rod is a poor conductor of
heat but nevertheless combustible, e.g., wood, then it
ANALOGY
1043 - 4046 / 96 - $5.00 - COPYRIGHT
Delhi
o 1996 THE AMERICAN
IN PHYSIOLOGY
S78
PHYSIOLOGICAL
EDUCATION
-JUNE
SOCIETY
1996
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TRAVELING
Sbabdara,
may be released asheat. It is excited by applying heat
to it. This at first charges the cigarette with heat in a
quite passive fashion, but when a critical temperature
is reached, something new happens. The chemical
energy of the cylinder is liberated as heat which
spreads out from the active region and raises the
temperature of the neighboring region. When the
temperature here reaches a critical value, this region
will also become active and so propagation will
proceed. If this paragraph is now reread substituting
the words nerve for cigarette, electricity for heat, a fair
description of the nerve conduction will be obtained.”
A knowledge of the factors affecting conduction of
bioelectric signalsis important not only in disciplines
related to the nervous system but also in cardiology
for understanding the pathophysiology of arrhythmias
and the mode of action of antiarrhythmic drugs.
Although the cable properties of nerve are crucial to
the understanding of the mechanism of nerve conduction, some popular textbooks (2, 3) make no mention
of it at all. Authors who avoid discussing cable
properties probably believe that most medical students will not appreciate the implications of the
electric equivalent circuit diagram (Fig. 1). Such a
surmiseunderlines the necessity of developing alternative strategies for explaining the cable properties, like
the ‘ ‘traveling flame’ ’ analogy mentioned here. The
analogy is not new; the name is. Also new is the
proposal for extrapolating the analogy for deducing
the factors affecting nerve conduction.
THE
Sciences,
INNOVATIONS
A
ri
N
D
I
D
E
of an axon.
resistance
r,, extracel(radial);
c,,
It would be incorrect to consider the two modes of
heat transfer as mutually exclusive. In fact, the traveling of the flame is critically dependent on heat
conduction.
The flame must conduct heat to its
adjacent area and raise its temperature to the ignition
point. Hence, factors affecting heat conduction also
affect the traveling flame. It would be of distinct
advantage to harness the speed of conduction with
the fidelity of the traveling flame. This can be achieved
through a model system (Fig. 2) in which small
combustible blocks are interposed between long conducting segments. These blocks, although boosting
the transmission
fidelity, also act as brakes on the
speed of transmission.
Therefore, longer lengths of
conductor with fewer blocks would favor a higher
transmission
speed. However, the advantage is selflimiting. If the heat losses along the metal take the
temperature below the ignition temperature
of the
combustible
material, then the transmission
would
El heat
q heat
conductor
insulator
combustible
Esl
15
FIG. 2.
flame
equivalent
of nerve
to one end of the rod.
: NUMBER
1 - ADVANCES
conduction.
IN PHYSIOLOGY
s79
Flame
EDUCATION
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1996
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burns itself out, conveying the flame to the other end
in the process. There are several differences between
the two modes of heat transfer. 1) In the combustible
rod, the transfer of heat is effected by the traveling
flame, whereas in the conducting rod, only the heat
travels; the flame remains at its own place. 2) A
conductor can be used several times over to transfer
heat; a combustible rod has to be replaced after every
episode of heat transfer. 3) The heat transfer is much
faster in conductors. 4) The heat transfer by combustion can be viewed as a high-fidelity transmission (the
temperature
of the flame remains constant as it
travels), whereas the conduction of heat is invariably
associated with decremental temperatures.
5) The
high-fidelity transmission
of the combustible
rod is
achieved at the expense of considerable energy expenditure. Transfer of heat by conduction is thus a far
more efficient mode of heat transfer, the heat losses
notwithstanding.
VOLUME
S
‘i
FIG. 1.
Electric
equivalent
circuit
of the cable properties
lular
resistance
(longitudinal);
r,, membrane
membrane
capacitance;
ri, axoplasmic
resistance.
Traveling
applied
A
INNOVATIONS
A
extinguish itself at the first block. Thus, for a given
temperature of the flame, there will be an optimal
length of the conductor at which it must be interrupted with a combustible thermal booster.
NERVE CONDUCTION
CORRELATES
VELOCITY:
ANALOGICAL
USING
E
A
S
THE
ANALOGY
A student once pointed out a paradox that a thicker
axon conducts faster but a thicker stick burns slowly.
The reason was explained to him as follows: the rate
of the rise of temperature in a thin stick is faster
because there is lessmaterial to be heated up (comparable to a lesser charge on the membrane). Nevertheless,the paradox does highlight the futility of stretching the analogy beyond a point.
Address for reprint requests: S. Sircar, Dept.
of Texas, Austin, TX 787 12-l 064.
11 January
1995; accepted
of Zoology,
in final form
4 January
University
1996.
References
1. Davson,
H., and M. G. Eggleton.
Principles
of Human
Physiology.
London: Churchill,
1968, p. 875.
2. Ganong,
W. F. Review of Medical
Physiology.
East Norwalk,
CT: Appleton-Lang,
1991, p. 45-52.
3. Guyton,
A. C. Textbook
of Medical
PbysioZogy.
Philadelphia,
PA: Saunders, 1991, p. 59-60.
IN PHYSIOLOGY
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1996
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1 - ADVANCES
D
A CAVEAT
Nerve conduction is speeded up if 1) the threshold for
the action potential (comparable to ignition temperature) is low; 2) the peak positivity of the action
potential (comparable to flame temperature) is high;
3) the axon has a larger diameter and, therefore, a
higher axoplasmic conductance (comparable to the
casewhere the combustible material is a good conduc-
: NUMBER
I
We begin our lectures on nerve-muscle physiology
with an overview of nerve conduction as indicated
above, emphasizing the need of a potential/chemical
energy stored in the conducting medium for transmitting signalswithout damping. It is in that context that
the lectures on membrane potential are begun. Later,
when we discuss the cable properties using the
electrical equivalent circuit, the students enthusiastically associate the description with its analogy discussedearlier. They also retain the concepts better, as
adjudged through periodic classtests.
Received
15
D
tor of heat aswell, e.g., a magnesium ribbon); 4) large
lengths of the axon are myelinated so that they
conduct electrotonically with action potentials occurring only at the nodes of Ranvier. Myelination minimizes the short circuiting of potentials across the
membrane (comparable to heat lossesin the absence
of insulation) and reduces the membrane capacitance
(comparable to a lower specific heat).
The speed of heat transfer in the new model system
would be contingent upon the variables relating to
combustion and to conduction. The ignition temperature and the temperature of the flame are in the
former category. The flame will travel fast if the
ignition temperature of the combustible material is
low (which would minimize the delay in ignition) and
flame temperature is high (which enables a speedier
heating of the area adjacent to the flame, since the rate
of heat transfer is proportional to the temperature
difference). The role of ignition temperature explains
why the flame travels faster on an inflammable material. The role of flame temperature explains why a
blazing flame travels faster than a smoldering fire.
Factors that would affect the speed of conduction are
the specific heat of the conductor and its insulation. A
low specific heat would allow quick rise of the
temperature of the matter adjacent to the heat source.
Better insulation prevents heat losses and thereby
channels the heat along the conductor, resulting in
speedier heat transfer. The corresponding analogues
in nerve conduction are easily identifiable. Conduction in an unmyelinated nerve can be likened to the
heat transfer through combustion, where the flame
represents the action potential. Indeed, the action
potential, as it is conducted on a membrane, has been
called the “traveling cathode. ” Saltatory conduction is
analogous to the model system in Fig. 2.
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N