Study

guides and introductions to

tutorials
Our course management software does not permit a return to an assignment once you have
submitted it. You can see the study questions and feedback, but the introductions with reading
assignments, learning objectives, etc. are no longer accessible. This document reproduces all of
that information in the order in which the assignments are due.

Electricity 1
To understand how neurons generate, conduct, and transmit signals you will need a good
command of basic electricity. This survey of ten questions addresses common misconceptions
about electricity, electric current, charge carriers, conductors, etc.
No grade will be recorded for this exercise. When you complete Electricity 1 you will
immediately have access to feedback. You may repeat this assessment as many times as you
wish.
Note bene Some of these questions would give a physics major a hard time. They did so for
me. All the same, when you read the feedback the correct answers should make sense.
Introduction to the study guides
Your instructor has prepared a series of study guides, one for each class, to assist you in learning
the fundamentals of cellular and molecular neuroscience. Study guides 1-3 will become available
two weeks before the start of class. Each subsequent guide will come out a week in advance of
the corresponding class. Each study guide includes lists of learning objectives, recommended
textbook readings, recommended resources to consult before and after class, and one or more
study questions.
Study questions are for learning they will not be graded, although for full credit you do need to
submit answers by the due date. Feedback on the study questions (a.k.a., the answer key) will
become available no later than one week after each corresponding class.

Introduction to the study guides

Learning the fundamentals of neuroscience
Prior to each class you are urged to:
complete any recommended followup work from the previous class
read the recommended sections of the textbook
review the PowerPoint notes pages posted in Resources
review any additional recommended resources
Following each class you are urged to:
obtain help with any concepts with which you are having difficulty
try to answer the study questions on your own, then work with your group if necessary
To develop complete confidence in your understanding of the material and in your ability to
apply the related skills, you are strongly encouraged to work in groups and come in for office
hours if and when you need help with any of the material.

Before the first class: recommended resources and assessment

A solid understanding of how neurons and nervous systems function requires reasonable
familiarity with basic electricity, including the properties of charged particles and electric
current, and Ohm's Law. You may want to review one or more of the tutorials listed below,
especially if you have a weak background in the physical sciences. Even students with a strong
background in the sciences may benefit from completing the assessment "Electricity 1."
basic_electricity.pdf (Resources section, tutorials folder)
Ohms_Law.pdf (Resources section, tutorials folder)
capacitance.pdf (Resources section, tutorials folder)
Electricity 1 (Tests & Quizzes)

Studyguide 1 22 Aug 16
Mon Aug 22: A biological basis for behavior
Class notes: 01_Intro_F16.pdf
Recommended textbook reading: None assigned, but if you have time, then the following
readings will get you off to a very good start.

Purves Ch 1 "Studying the nervous system" introduces model organisms, neurons and
glial cells, neural circuits, neural systems, and much of the terminology to be used in the
course.

Nicholls Ch 1 "Principles of signaling and organization" introduces important topics that

will be introduced in the first few weeks of the course.

Kandel Chs 1 & 2 are a useful introduction to the entire course, touching on several of the
topics to be covered later in more detail.

Accessing journal articles: As a member of Rice University you have full library privileges,
including online access to journal articles. You should have direct access but if not then you may
need to use the Library's OneSearch tool. If you do not know how to access online materials
from Fondren Library then you will want to open and follow the tutorial "Retrieving_
journal_articles.pdf" in the Tutorials folder in the Resources section.
Follow-up article (online): Glickstein, M. Golgi and Cajal: The neuron doctrine and the 100th
anniversary of the 1906 Nobel prize. Current Biology 16(5): R147-R151, 2006. This article
nicely follows (or precedes) the discussion in class of Cajal, Golgi, reticular theory, and the
neuron doctrine. Some of the information, such as the discussion of dendritic spines, may prove
valuable later.
What you should be able to do following this class
You should be able to articulate the contradictory views of the role of the brain in
behavior throughout the 19th century and half of the 20th and cite evidence that was used
to support the opposing views. Be able to describe, in your own words, the models for the
roles of the brain and neurons that were finally accepted, and why they were accepted.
What is meant by the neuron doctrine? Be prepared to cite evidence that clearly
established the role of neurons as the basic functional units of the nervous system.
Be able to access primary literature online.

Equilibrium potentials
All living cells maintain an electrical potential difference (Em) across their plasma membranes.
The first model that attempted to explain why cells have membrane potentials suggested,
incorrectly, that Em is the equilibrium potential for potassium. An equilibrium potential is the
electrical potential difference that results from an asymmetric distribution of a single type of ion
across a semipermeable membrane. It turns out that for most cell types Em is a diffusion potential
resulting from asymmetric distributions of two or more ions, and not an equilibrium potential.
Nevertheless, the concept of an equilibrium potential is absolutely critical toward understanding
how nerve and muscle cells generate and conduct action potentials, how neurons communicate
via synapses, and how various stimuli activate (or in some cases deactivate) sensory receptor
cells.
In excitable cells including neurons, muscle cells, and receptor cells, external or internal stimuli
cause ion channels to open and/or close, producing a temporary change in Em that serves as a
signaling mechanism in the form of a graded potential or an action potential. To predict the
influence of any intervention on Em we must be able to predict the direction in which an ion
moves and the magnitude of the electric current that its movement produces. To do so we must
learn to work with equilibrium potentials.

Studyguide 2 24 Aug 16
Weds Aug 24: Animal electricity
Class notes: 02_AnimElec_F16.pdf
Recommended textbook reading:
Purves Ch 2, "Electrical signals of nerve cells" p. 25-36 concisely introduces the
electrical properties of neurons. We will cover this material in much more detail, taking a
historical, experimental approach.
Nicholls Ch 1 "Principles of signaling and organization." In particular, pages 10-15
provide an overview of material to be covered today; Ch 6 "Ionic Basis of the Resting
Potential" presents theory behind the resting membrane potential along with some
experimental evidence.
What you should be able to do following this class
Be prepared to cite the evidence that neural activity is electrical in nature and why for so
long there was much reluctance to accept a role for electricity in nerve signaling.
Be able to articulate the rationale behind the model of the resting potential as an
equilibrium potential and Bernstein's explanation for action potentials. Why was it
considered unlikely that a nerve axon passively conducts electricity all the way from the
input side to the output side? How did Hermann's model get around this problem?
Follow-up article (online): Piccolino, Marco. Animal electricity and the birth of
electrophysiology: The legacy of Luigi Galvani. Brain Res Bull 46(5): 381-407, 1998. For
students interested in the history of neuroscience, this article is a nice review of the progress of
science from the discovery that animal tissue has electrical properties up to the Hodgkin-Huxley
experiments.

Related tutorials:
It will probably be of benefit to spread the tutorials out a bit. Complete the first one, make sure
that you have all of the principles well in hand, and wait a bit before moving on to the next one.
The due dates are spread out to facilitate the learning value.
[Tests & Quizzes] "Equilibrium potentials" (two parts) uses a simple model to present the
basic equilibrium potential concept.
[Tests & Quizzes] "Ion movement pt 1" (two parts). What happens to permeant ions
when a system is not in equilibrium? Part 2 provides a study question.
[Tests & Quizzes] "Ion movement pt 2." When ion channels open and ions cross a cell
membrane, what happens to the ion concentrations on both sides? Knowing how to
answer this question is critical to developing an intuitive understanding of resting, action,
generator, and receptor potentials in neurons. This tutorial is also available in Resources
as a pdf (ion_movement.pdf). You might want to work with it as a group.

Ion movement pt 1
Movement of ions that are not in equilibrium
Our two compartment model represents an artificial situation, of course. One difference between
the example in Equilibrium potentials pt 1 and a real cell is that the resting potential difference
Em across a cell membrane is usually NOT an equilibrium potential. To fully understand how
ions support signaling within neurons, we need to know how ions move when Em Eion, that is,
when the membrane potential is not the same as the equilibrium potential of the ion.

Studyguide 3 26 Aug 16
Fri Aug 26: Testing the Bernstein hypothesis
Class notes: 03_F16_Testing.pdf
Recommended articles:

[in Resources] Adrian1917_chIV.pdf. The Adrian article, now in the public domain, is
Chapter IV from the 1917 book by Keith Lucas (published posthumously). It describes
the rationale behind the vapor cooling experiment using the straightforward language of
the day.

[access through Fondren] Keynes, Richard, J Exp Physiol 208: 179-180, 2005. This
article discusses the scientific impact of JZ Young's 1938 publication on the function of
squid giant nerve fibers.

Recommended textbook reading:

Purves Ch 2 "Electrical signals of nerve cells."

Nicholls Ch 8 "Electrical signaling in neurons."

What you should be able to do following this class

You should be able to make predictions that follow from the Bernstein model for
electrical activity in cells, cite experiments that tested the predictions, and explain the
findings and their significance. Be able to explain what predictions could or could not be
tested in the early 1900s and why.
4

Be able to distinguish a cathodal from an anodal stimulus and threshold from

subthreshold stimulus, and their respective influences on nerve activity. For example,
why can a cathodal but not anodal stimulus generate an action potential?

Be prepared to employ one or more analogies to explain the respective roles of passive
and active forces in regenerating an action potential. Be able to cite experiments
involving extracellular recordings that led up to the Hodgkin and Huxley voltage clamp
experiments and what their results revealed. Be prepared to explain the importance of the
re-discovery of the squid giant axon to neuroscience.

Related tutorial: [Tests & Quizzes] "Ion currents" works through principles governing
movements of ions through channels, the currents that they carry, and the effects of ionic
currents on Em.

Ion movement pt 2
When ions move across a cell membrane, what is the impact on the intracellular concentration?
This tutorial can be a real eye-opener! It is designed to dispel a common misconception about
membrane potentials that is shared by nearly all beginning (and many advanced) students of
neural science. Completing this tutorial will be a formative (learning) experience. It is not
designed to test you for prior knowledge and no grade will be recorded.

Studyguide 4 29 Aug 16
Mon Aug 29: Discovering the diffusion potential
Class notes: 04_F16_Discover.pdf
Recommended article: [access through Fondren] Hodgkin, AL and AF Huxley, "Action
potentials recorded from inside a nerve fibre," Nature 144: 710-711, 1939. If you carefully
peruse this brief communication by Hodgkin and Huxley reporting the first recorded intracellular
action potential you should find the answer to one of the study questions.
Recommended textbook reading:
Purves Ch 2 "Electrical signals of nerve cells" p. 36-38; Box 2C "Action potential form
and nomenclature" p. 39
Nicholls Ch 6 "Ionic basis of the resting potential"
Kandel Ch 6, "Membrane potential and the passive electrical properties of the neuron"
sums up modern theory regarding membrane and action potentials and provides a good
background for the coming class on conduction of the nerve impulse.

What you should be able to do following this class

Be able to distinguish the features and significance of extracellular versus intracellular

recordings including polarity and relative magnitudes. You should be able to look at a
recording and know whether it is extracellular or intracellular based upon information
provided.
Be able to explain the significance of the features of the first intracellular recording of a
nerve action potential. Why did the appearance of these features lead to a revision of the
Bernstein model for resting and action potentials? Be prepared to cite experiments that
supported the sodium theory and the diffusion potential model and explain the rationale.
Be able to explain how a diffusion potential differs from an equilibrium potential. Be
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prepared to determine an equilibrium potential or diffusion potential, knowing the

relevant ion concentrations. Given sufficient information, be able to use the Nernst and/or
Goldman equations to determine intracellular concentration of an ion, whether or not an
ion is in equilibrium, and/or permeability coefficient for an ion. Several study questions
will address these objectives.
Related tutorial: [Tests & Quizzes] "Ion conductance" (two parts) introduces another way of
measuring ion and current flow through membrane channels. Part 2 presents a multiple part
study question. You are encouraged to go through the tutorial before the next class on the ionic
basis of the action potential.
Working with the Nernst & GHK equations: [Tests & Quizzes} "Nernst and GHK" includes
five additional study questions in which you work with the two equations to solve quantitative
problems. Learning these strategies will help you with quantitative problems that may appear on
a homework set or exam.
Assigned journal article
In class today you will receive copies of a public release from EurekAlert! Science News and
selected pages from the corresponding journal article. The complete article, cited below, can be
accessed by anyone with a campus internet connection or off-campus VPN connection.
Copyright laws prohibit posting of the article on Owl-Space.
Pirta Hotulainen et al., MIM-Induced Membrane Bending Promotes Dendritic Spine Initiation.
Developmental Cell, June 2015 DOI: 10.1016/j.devcel.2015.04.014.
For complete instructions please see the reading guide for this article, posted in Resources in the
folder (you guessed it!) Reading guides.

Ion currents
Directions and magnitudes of ion and current flow and their effects on
Em
It is important to be able to predict the direction in which specific ions will move, the direction
of electric current carried by the ions, and the impact on Em when an abrupt change in membrane
permeability to one or more ions perturbs the steady state. Is the ion movement polarizing
(making Em more negative) or depolarizing (making Em more positive)? We may also need to
estimate magnitude of current flow using Ohms Law.
Ohms Law, V = iR is both one of the most important principles in electricity and (thankfully)
one of the simplest. Ohms Law applies not only to electrical circuits but also to the movement
of ions through membrane channels. Ohms Law can be written as E = iR, where E (volts) is the
electromotive force, i is current, and R is the resistance to current flow. When we are working
with ion flow across membranes we use the expression (Em - Eion) in place of V or E.

Studyguide 5 31 Aug 16
Weds Aug 31: Ionic basis of the action potential
Class notes: 05_F16_IonicBasis.pdf

Recommended textbook reading:

Purves Box 2B "The remarkable giant nerve cells of squid" p. 36; Box 3A "The voltage
clamp method" page 42; Box 3B "Threshold" p. 49; Ch 3 "Voltage-dependent membrane
permeability" p. 41-49
Nicholls Ch 7 "Ionic basis of an action potential"
Kandel Ch 7 "Propagated signaling: the action potential" up to p. 158
Recommended for review:
Two potential conceptual stumbling blocks for this class are how an ion moves with respect to its
equilibrium potential and the meaning of ion conductance. If you aren't sure of your grasp of
these concepts then please go back through the related tutorials.
Ion currents
Ion conductance
What you should be able to do following this class
Be able to explain the principles of voltage clamping and its use in studying very
rapid membrane-mediated events.
Be able to explain the principle behind use of a Ag/AgCl electrode for measuring Em
and injecting or withdrawing current.
Be able to explain "capacitive current"
Be prepared to predict current direction across a membrane at varying clamp voltages
and with/without blocking agents, to identify the ion(s) responsible, and to predict
whether or not the circuit will have to inject or withdraw current to maintain a constant
membrane potential.
You should be able to explain how the timing of Na and K voltage-gated channels
results in a fast, predictable, all-or-none action potential.
Be prepared to explain the rationale behind the models of the resting and action
potentials, present and past.
Study problems: [Tests & Quizzes] "Nernst and GHK" has you work with these two important
equations to calculate unknown variables such as Em, Eion, or intracellular concentration of an
ion.
The resources on basic electricity and Ohms Law (Resources, folder electricity) on OwlSpace should help with the concepts introduced in this tutorial.

Ion conductance
NOTE FOR FALL 2016: There is an error in the feedback to question 2 regarding the physical
units that make up 1 siemens. CORRECT FEEDBACK: From Ohm's Law: 1 volt = 1 ampere 1
siemens. Rearranging, one siemens = one ampere-volt-1 (i.e., 1 ampere/1volt).
Movements of ions across cell membranes are responsible for the electrical currents that produce
the resting, graded, and action potentials that are so important for cell maintenance, for
intercellular signaling, for sensory reception, and for motor control. To study these currents we
need to introduce a new electrical term, namely ion conductance (gion). Ion conductance refers to
the ease with which an ion passes through a single channel or through many channels in a cell
membrane. Here are some reasons for learning to work with ion conductance.
Changes to gNa and gK are responsible for the action potentials in excitable cells

Hodgkin, Huxley, and Goldman used ion conductances and Ohms Law to derive their
equation for the diffusion potential
The goals of two of the most important techniques in neuroscience, voltage clamping and
patch clamping, are to measure ion conductance of individual channels or areas of
membranes
Investigators use ion conductance per unit area of membrane as a measure of ion channel
density
Conductance of mutant ion channels was used to learn how many subunits make up an
ion channel

The difference between sodium conductance and potassium conductance explains the reversal
potentials of cation channels

Sodium pump
Electrogenic contribution of the Na/K pump
When we have a diffusion potential, sodium ions constantly diffuse in, potassium ions constantly
diffuse out, and the sodium and potassium currents are equal and opposite. Passive
Na+/K+ exchange is electrically neutral, therefore if there were no active transport of these ions
the Na+ and K+ gradients would decay. The gradients do not decay because an ATP-dependent
sodium pump maintains them by pumping out Na+ in exchange for K+.
Unfortunately (from the viewpoint of students who are trying to understand these relationships)
active transport of Na+ and K+ is NOT electrically neutral. With each cycle, three Na+ are
pumped out for every two K+ pumped in. By pumping out an excess positive ion for each ATP
hydrolyzed, the pump makes a small direct contribution to the Em. We refer to this contribution
as the electrogenic effect. To understand the significance of the electrogenic effect it is critical
that students completely understand the rationale behind the following two statements.
1. The electrogenic effect is not responsible for the membrane potential; it does make
Em somewhat more negative than the diffusion potential.
2. A cell at rest is in a steady state in which the pumping of excess sodium ions out of the cell is
offset by sodium ions leaking into a cell at a slightly higher rate than the rate at which potassium
ions leak out.

Threshold currents
Potassium and sodium currents on approach to threshold
When an excitable cell is depolarized quickly enough with a strong enough voltage, Em reaches
what we call a threshold value. At threshold, sufficient voltage-gated sodium channels open to
support an increased sodium current, which depolarizes the membrane even more, opening even
more sodium channels, further increasing sodium current, etc. This positive feedback cycle is
also called the Hodgkin cycle, after the physiologist who first proposed the mechanism.
What is going on with sodium and potassium currents on approach to threshold? Does it get
progressively easier to trigger an action potential, or does it become more difficult? These
questions are important when we consider that neurons may receive dozens, hundreds, or even
thousands of excitatory and inhibitory inputs at once. How fine tuned are these neurons? If a
neuron receives a thousand inputs how much difference can one single synapse make?

Studyguide 6 2&7 Sep 16

Fri Sep 2 & Mon Sep 7: Conduction of the nerve impulse
Class notes: 06_F16_Conduction.pdf
Recommended for review:
You may want to review "capacitance.pdf" (Resources: tutorials), which presents the
concept of electrical capacitance in circuits and membranes.
"Heat_analogy.pdf" (Resources: tutorials), is adapted from an explanation on p. 115 of
Nicholls' 4th edition of "From neuron to brain." This document compares conducted
current to conducted heat.
Recommended textbook reading:
Purves Box 2A " Passive membrane properties" p. 30-31; Box 3C " Multiple sclerosis "
p. 54
Nicholls Ch 8 "Electrical signaling in neurons"
Kandel Ch 6 "Membrane potential and the passive electrical properties of the neuron"
starting on p. 135 "The functional properties..." deals with the same subject matter that
we will discuss in class.
What you should be able to do following this class

Be prepared to relate the events of the Hodgkin cycle to the phases of action potential
conduction - activation, generation of the action potential, refractory period. You must be
able to relate the phases to ion channel activation, timing of opening, closing, and
inactivation, and the effects on ion conductance.

The role of passively conducted current in action potential regeneration should be clear now.

How do the properties of graded and action potentials differ, and why?
Why does passively conducted current decay with distance down an axon?
Why is there a time delay before a membrane downstream from a stimulus or action
potential achieves the full change in membrane potential?
Be prepared to predict qualitative differences in conduction velocity based upon axon
diameter and whether or not an axon has a myelin sheath; you will not be asked to
reproduce the equations that describe Vx or Vt, however you may be asked to use them.

Tutorials:

"Threshold_currents" (Tests & Quizzes) helps with understanding how neurons generate
and propagate action potentials

"Sodium pump" (Tests & Quizzes) provides valuable background information for the workshop
next Monday Sep 12

Studyguide 7 9 Sep 16
Fri Sep 9: Ion channel function
Class notes: 07_F16_ChanFunc.pdf

Recommended textbook reading:

Purves Ch 4 "Ion channels and transporters" p. 57-69

Nicholls Ch 4 "Ion channels and signaling" (all)

Kandel Ch 5 "Ion channels" p. 100-110

What you should be able to do following this class

You will be presented with experimental situations in which you are asked to predict results such
as the magnitude and/or direction of ion movement and/or current. You might be asked to design
an experiment or explain why an experiment is or isn't practical. Of course you should know
what factors influence channel conductance and you should be able to differentiate between
microscopic and macroscopic currents. In what ways do channels select for specific ions? For
example, what mechanisms can make a channel specific for potassium, sodium, or chloride ions?
You should be able to interpret current-voltage plots and to prepare current-voltage plots based
upon information that you are given.

Studyguide 8 12 Sep 16
Mon Sep 12: Discovering the sodium pump
***BRING A CALCULATOR (OR CELL PHONE WITH CALCULATOR APP) TO CLASS
TODAY***
In today's class we will go through three of the classic experiments leading to the discovery that
an ATP-dependent Na/K exchanger maintains the sodium and potassium gradients. Reviewing
the class notes and the recommended tutorial on the sodium pump experiments should be
adequate preparation for class.
Class notes: 08_F16_Na_pump.pdf
Tutorial: Don't forget to go through the "Sodium_pump" tutorial in Tests & Quizzes before
class.
What you should be able to do following this class
By the end of the class you should have a solid understanding of the role of sodium-potassium
ATPase (the sodium pump) in cells. In particular, you should be able to:

Explain the rationale and interpret results for the three experiments discussed in class
Explain why cells have resting membrane potentials
Distinguish the two effects of sodium pump activity, namely (1) maintenance of the
sodium and potassium gradients and (2) the electrogenic effect, and their significance in
terms of resting membrane potentials
Explain why the inward sodium current exceeds the outward potassium current in a cell
at rest

Studyguide 9 14-16 Sep 16

Wed & Fri Sep 14 & 16: Studying channel structure
Class notes: 09_F16_ChanStruc.pdf

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Recommended video clip:

Membrane protein insertion video (might be a bit elementary, but it is very clear):
http://www.youtube.com/watch?v=PUy_Em5dXmc
Recommended textbook reading:
Purves Ch 4 "Ion channels and transporters"
Purves Ch 6 "Neurotransmitters and their receptors" (use for reference - reading is rather
tedious)
Nicholls Ch 5 "Structure of ion channels" (all)
Kandel Ch 6 "Ion channels" p. 116-123
What you should be able to do following this class
Be prepared to predict the orientation of a membrane protein and number of
transmembrane segments from a hydropathicity map and/or use a diagram of a channel
subunit to predict the pattern of signal sequences in an mRNA transcript coding for a
channel subunit.
Be prepared to apply one or more of the experimental strategies that we examined in
class for learning about channel structure and subunit structure
Be prepared to suggest ways in which expression of channel proteins in Xenopus oocytes might
be used to reveal channel structure.

Studyguide 10 19 Sep 16
Class notes: 10_F16_ChanDiver.pdf
Recommended videos:
Kinesin
I picked these two because they are relatively short. A Google search will turn up some longer
animations that you may find useful.
www.youtube.com/watch?v=4TGDPotbJV4
www.youtube.com/watch?v=YAva4g3Pk6k
Dynein

https://www.youtube.com/watch?v=-7AQVbrmzFw

Recommended textbook reading:

Purves Ch 4 "Ion channels and transporters" (all)
Nicholls Ch 5 "Structure of ion channels" (all)
Kandel Ch 6 "Ion channels" p. 116-123
What you should be able to do following this class
Be prepared to explain what sorts of modifications of the same basic protein "architecture" have
produced an incredible diversity of channel types, including channels that serve very different
functions, such as conducting anions instead of cations.

Studyguide 11 21 Sep 16
Weds Sep 21: Introduction to synaptic transmission
Class notes: 11_F15_Syntran.pdf

11

Recommended textbook reading:

Purves Ch 5 "Synaptic transmission" p. 77-81
Nicholls Ch 11 "Mechanisms of direct synaptic transmission" p. 185-187; 207-210
Kandel Ch 10 "Overview of synaptic transmission" (all)
What you should be able to do following this class
Be prepared to identify a general type of synapse based upon properties that are described
for it.
Be prepared to cite observations that can be made in order to determine whether a
receptor is ionotropic, metabotropic, or electrical
Be able to describe the mechanisms by which materials are transported from soma to
axon terminal, and some of the challenges facing such transport.

First exam
The exam, scheduled for Sep 30, will include questions on material from the beginning of the
course through "Introduction to synaptic transmission." The second exam period begins with
"Presynaptic mechanisms role of calcium."

Nernst and GHK

Here are a few more study questions that focus on quantitative work. You will need to be
comfortable using the Nernst and/or Goldman-Hodgkin-Katz equations to calculate an
equilibrium potential, membrane potential, or intracellular concentration of an ion.

12