A problem-solving approach to teaching electrochemical driving force to undergraduates

W. F. Nolan
Advan in Physiol Edu 259:S1-S3, 1990. You might find this additional info useful... Updated information and services including high resolution figures, can be found at: http://advan.physiology.org/content/259/6/S1.citation Additional material and information about Advances in Physiology Education can be found at: http://www.the-aps.org/publications/advan

This infomation is current as of September 2, 2011.

Downloaded from advan.physiology.org on September 2, 2011

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 1990 by the American Physiological Society. ISSN: 1043-4046, ESSN: 1522-1229. Visit our website at http://www.the-aps.org/.

A problem-solving approach to teaching driving force to undergraduates

WILLIAM Department F. NOLAN of Biology, University of Hartford,

electrochemical

-- -West ---- Hartford, 1 Connecticut Ml - 17 * -

NOLAN, WILLIAM F. A problem-solving approach to teaching electrochemical driving force to undergraduates. Am. J. Physiol. 259 (Adv. Physiol. Educ. 4): Sl-S3, 1990.-The diversity of students in the typical undergraduate general physiology course demands consideration of several important educational objectives: the instructor seeks to provide students with a sound understanding of basic physiological concepts, sharpen their analytical skills, and demonstrate how universal physicochemical laws determine function within living systems. These objectives converge in a problem-solving approach to electrochemical driving force and ion flux across biological membranes. undergraduate physiology; analytical skills

problem-solving approach to ECDF and have found this to be an effective tool in developing students reasoning ability while guiding them to a more complete understanding of the forces that determine the movement of ions. My objective here is to describe this problemsolving method of teaching ECDF. It is not my intention to present an exhaustive treatment of membrane potential or bioelectrical signal generation; yet I will begin with a brief description of these phenomena to provide the context within which ECDF is developed in the undergraduate general physiology course. More exhaustive treatments of these topics are available (l-5).
BACKGROUNDMATERIAL

Downloaded from advan.physiology.org on September 2, 2011

of teaching physiology to undergraduates resides in the diversity of the abilities and career objectives of the students. It is likely that some students in each class will pursue careers in physiology or medicine but most will not. Many will pursue careers unrelated to science. Because effective teaching demands a clear perception of educational objectives and because educational objectives must address student needs, the heterogeneity of the typical undergraduate physiology class in the typical liberal arts institution challenges the instructor with several important teaching goals. These include providing the students with a firm understanding of the basic physiological concepts on which higher order life processes are based, enhancing their awareness of unifying concepts, e.g., the dependence of living systems on fundamental physicochemical laws, and improving their problem-solving skills. When teaching electrochemical driving force (ECDF), the instructor has the opportunity to accomplish all three of these educational objectives: ECDF is a basic but important physiological concept because of its central role in bioelectrical signal generation. It also illustrates how essential processes within organisms, e.g., action potential generation and intercellular communication, rely on the same laws that rule the nonliving universe. Finally, if the instructor takes a problem-solving approach to teaching ECDF, the students analytical skills can be enhanced. Although a quantitative treatment of ECDF is often seen in neurobiology textbooks designed for graduate students (5), it is only occasionally found in introductory physiology texts (1). Thus supplemental material may be needed if the instructor wishes to include some quantitative aspects of the subject in an undergraduate general physiology course. I have been using as an instructional aid a simple
PART OF THE CHALLENGE
1043-4046/90 $1.50

Membrane potential ( Vm) is the separation of charge across a cell membrane and is established by the selective permeability of the membrane and the active transport of ions across it. The result is a differential distribution of ions and an excess of negative charge on the membranes inner surface. For any ion, X, present in unequal concentrations across the membrane, there are two forces acting on it. First, there is a chemical driving force (CDF) resulting from the concentration gradient. Second, there is an electrical driving force (EDF) resulting from the interaction between the charge of the ion and Vm. If these forces are equal in magnitude but opposite in direction, there will be no net, or electrochemical, driving force acting on that ion (Fig. 1A). Under these conditions, ion X will be in electrochemical equilibrium and will exhibit no net flux in either direction across the membrane. The membrane potential that exactly balances the CDF, thus establishing electrochemical equilibrium, is obtained from the Nernst equation E x = (RT/zF)ln(X,/XJ where E, is the equilibrium potential for ion X, R is the gas constant, T is the absolute temperature, z is the valance of ion X, F is the Faraday constant, Xe the extracellular concentration of X, and Xi the intracellular concentration of X. Converting to log base 10 for a monovalent ion at 18OC,the Nernst equation reduces to c E x = 58 lOg(XJXi) If Vm does not equal E,, then X will be out of electrochemical equilibrium and there will exist an ECDF equal in magnitude to the difference between Vm and E, (ECDF), = Vm - E,
Physiological Society
Sl

Copyright

0 1990 the American

s2

TEACHING

ELECTROCHEMICAL

DRIVING

FORCE

1\ I=)

EDF CDF -

NET DF=CDF-EDF

This will result in a net passive flux of X across the membrane in the same direction as the ECDF (Fig. lI3). Thus X will carry current, Ix, across the membrane in accordance with a modified version of Ohms law I x = gxwn - Ex)

r\

FIG. 1. A: example of a neuron where electrical and chemical driving forces (EDF and CDF, respectively) acting on cation X are equal and in opposite directions, resulting in no net, or electrochemical, driving force (DF) for X, i.e., electrochemical equilibrium with respect to X. B: example of a neuron where outward CDF for cation X exceeds inward EDF, resulting in outward net driving force acting on X. Vm, membrane potential; [x+]i, [x+1,, intraand extracellular concentrations of cation X, respectively.

whether the membrane current it carries is depolarizing (excitatory) or hyperpolarizing (inhibitory).

GRAPHIC AND NUMERICAL OF ELECTROCHEMICAL ANALYSIS DRIVING FORCE

Downloaded from advan.physiology.org on September 2, 2011

where g, is the membrane conductance for ion X. In the steady state, when the cell is not signaling (i.e., at resting Vm), the concentration gradients of ions that are not in electrochemical equilibrium and that, therefore, exhibit net passive flux across the membrane, are maintained by active transport. Thus the passive flux of an ion in one direction will be offset by active transport in the opposite direction. The end result for all ions is no net movement of charge across the membrane and a stable, resting Vm. The deflection of Vm from rest is the basis of bioelectrical signal generation. This is accomplished by changing membrane conductance to an ion (or ions) that is (are) out of electrochemical equilibrium, resulting in a net membrane current and, therefore, a change in Vm. The membrane conductance for specific ions is altered by opening or closing specific gated ion channels. A change in membrane conductance for any ion X, which is not in electrochemical equilibrium, will cause a change in Ix Ar, = 4Lu4n - Ex)

Thus the balance between active and passive fluxes will be transiently disrupted, a net movement of charge across the membrane will occur, Vm will deflect from its resting value, and a bioelectrical signal will be generated. For any given change in g,, the direction of the resulting change in 1x and its magnitude, i.e., the nature of the signal, will be determined by the direction and magnitude of the electrochemical driving force acting on X. I have found that most students easily grasp the concept and mechanism of changes in membrane conductance. Computing the numerical value of electrochemical driving force is also simple. However, understanding the mechanistic basis of ECDF and, in particular, determining its direction for a particular ion under a given set of conditions are often considerably more difficult. Note that this last determination is critical because for any ion the direction of its electrochemical driving force will determine the direction of its passive flux and, therefore,

The reasoning that is applied in analyzing ECDF by this method may be summarized as follows. 1) The electrical and chemical driving forces acting on an ion may be in the same or in opposite directions. If they are in the same direction (e.g., from inside to outside the cell), then the direction of the ECDF acting on that ion will be in that direction as well, and its magnitude will equal the sum of EDF and CDF. If EDF and CDF are in opposite directions, then the direction of the greater of these forces will determine the direction of the ECDF. The magnitude of the ECDF in this situation will equal the difference between the EDF and the CDF (Fig. 1B) 2) In cases where the directions of the EDF and CDF are opposite, their magnitudes must be compared to determine the direction of the ECDF. The strength of the EDF acting on an ion is equal to the membrane potential. Because the equilibrium potential for any ion is the membrane potential required to balance its CDF, the Nernst equation can be used to calculate the millivolt equivalent of its CDF. Thus the equilibrium (i.e., Nernst) potential can be compared with the given Vm (also in mV), and the direction of the ECDF for any ion can be determined. Sample problem 1. Consider a situation where K ion has an intracellular fluid (ICF) concentration of 350 mM and an extracellular fluid (ECF) concentration of 50 mM. What will be the direction and magnitude of the ECDF acting on K at Vm = -60 mV? Solution: the magnitude of the EDF is 60 mV (the given membrane potential) and is directed toward the cell interior because of electrostatic attraction. The magnitude of the CDF, computed from the Nernst equation, is 49 mV. Its direction is from the region to higher to lower concentration or, in this case, toward the ECF. The magnitude of the ECDF is equal to 60 - 49, or 11 mV. Its direction is the same as that of the EDF. Solving this problem is facilitated by the graphic representation of the given conditions (Fig. 2). Once the direction and relative magnitude (indicated by arrow length) of the EDF and CDF are drawn, the direction and relative

TEACHING

ELECTROCHEMICAL

DRIVING

FORCE

s3

EDF (6OmW \

CDF (49mV) Net DF (1lmV)

to carry their analysis one step further and determine whether a given change in conductance will result in a depolarization or a hyperpolarization of the cell membrane. Thus the importance of ECDF in signal generation is emphasized.
CONCLUSION

[K 1. =35O mM I

[K le =50 mM

FIG. 2. Example chemical equilibrium. tions.

of a case where potassium ion is out of electroSee SampZe probZem I and Fig. 1 for abbrevia-

magnitude of the ECDF becomes apparent. Although classroom discussion should certainly emphasize actual neuron structure and membrane differentiation with respect to ion channels, the use of a simple circle diagram, as in Fig. 2, provides a quick way of representing intraand extracellular fluid compartments for directional analysis of driving force. Sample problem 2. If ICF K ion concentration is 350 mM, what ECF K concentration is required for the condition of no net passive flux of K across the membrane at Vm = -60 mV? Solution: because K must be in electrochemical equilibrium for it to exhibit no net passive transmembrane flux, the problem reduces to one of determining the ECF concentration of K needed to yield a K Nernst potential of -60 mV given that ICF K concentration is 350 mM. Solving the Nernst potential for ECF K after making appropriate substitutions yields a value of 29.2 mM. I generally assign several of these problems dealing with both anions and cations under various conditions of Vm and concentration gradient. I also ask my students

This approach to teaching electrochemical driving force helps students comprehend the forces that govern ion flux across biological membranes and sets the stage for more elaborate analysis of bioelectrical signals. It also challenges their reasoning ability and sharpens their problem-solving skills. For the students who will not pursue a career in science, perhaps the major benefit of a thorough understanding of ECDF and ion flux is its contribution to the development of an expensive worldview that places living systems within the domain of universal laws.
Downloaded from advan.physiology.org on September 2, 2011
Received 2 November 1989; accepted in final form 14 June 1990.

REFERENCES ECKERT, R., D. RANDALL, AND G. AUGUSTINE. AnimaZ Physiology: Mechanisms and Adaptations (3rd ed.). New York: Freeman, 1988. HODGKIN, A. L. The ionic basis of electrical activity in nerve and 2. muscle. BioZ. Reu. 26: 339-409, 1951. 3 HODGKIN, A. L., AND A. F. HUXLEY. Currents carried by sodium . and potassium ions through the membrane of the giant axon of LoZigo. J. PhysioZ. Lond. 116: 449-472, 1952. 4. HODGKIN, A. L., AND A. F. HUXLEY. The components of the membrane conductance in the giant axon of Loligo. J. Physiol. Lond. 116: 473-496,1952. 5. KANDEL, E. R., AND J. H. SCHWARTZ. PrincipZes of Neural Science. New York: Elsevier, 1983.

1.