51-65 PAGE N.

2. Nerve cells and behavior

• Can memory and learning be studied at the level of
nerve cells and their interconnections?

• A beginning is made by relating changes In the

activity of nerve cells to the modification of simple
behaviors
Nerve cells and behavior
 Module Objectives

• The Cellular Components of the Nervous System

• Structures of the neuron

• Role of neurotransmitters on behaviour

The Neuron

• The neuron is the basic building block of

the nervous system

• They are often grouped in bundles

called nerves.
 The 4 parts of Neuron

1. 1. Dendrites are specialized to receive signals

from neighboring neurons and carry them back
to the cell body

• Thin, bushy-like structures that receive

information from outside the neuron

• Relays the information into the cell body

 The 4 parts of Neuron

1. 1. Dendrites are specialized to receive signals

from neighboring neurons and carry them back
to the cell body

• Thin, bushy-like structures that receive

information from outside the neuron

• Relays the information into the cell body

 The Neuron
2. The Cell body
contains the cell
nucleus

• The cell body relays

the information
down to the axon
 The Neuron

3. 3. Axon: A thin, long structure that transmits

signals from the cell body to the axon terminal.

4. 4. Axon Terminal is the last step for the relay of

information inside the neuron.
Let’s Review!
The cell body is
covered with
Axon Terminals
 The Neuron

Once the information hits the

terminal, it is transmitted outside
the cell by neurotransmitters, which
reside in the axon terminal.
Neuron Internal Structure
 CNS Support Cells

• Neuroglia (“glue”) provide physical support, control

nutrient flow, and are involved in phagocytosis

• Astrocytes: Provide physical support, remove debris

(phagocytosis), and transport nutrients to neurons
• Microglia: Involved in phagocytosis and brain immune
function
• Oligodendrocyte: Provide physical support and form the
myelin sheath around axons in the brain

• Schwann Cells form myelin for PNS axons

The Neurons communication
 The Neurons communication

1. Electrical Communication

2. Chemical Communication
 Electrical communication

• Action potential is an electrical current sent

down the axon.

• The activity within the neurons is electrical. This

current causes the neuron to “fire”

• This is an “all-or-none” process

Electrical communication

Action potentials travel down the axon like

a wave of energy
The cell membrane of a resting
neuron is more negative on the
inside of the cell than on the
outside. When the neuron is
stimulated, the permeability of
the membrane changes, allowing
Na+ to rush into the cell. This
causes the inside of the cell to
become more positive. This local
change starts a similar change in
the adjoining segment of the
neuron’s membrane. In this
manner, the electrical impulse
moves along the neuron. From:
Molecular Cell Biology, by Lodish
et al. 1986, 1990 by Scientific
American Books.
 Nerve Impulse Propagation

• The impulse continues to

move toward the cell
body
• Impulses travel faster
when fibers have a myelin
sheath
 Resting Membrane Potential
• Resting membrane potential (RMP) is the difference in
voltage between the inside and outside of the axon
membrane
• NA+ ions are in high concentration outside the cell,
while K+ ions are in high concentration inside the cell
• At rest, sodium-potassium transporters (pumps) push three
NA+ ions out for every two K+ ions they push in, causing the
exterior of the nerve cell membrane to be slightly positive
relative to the inside of the axon

2.23
 The Action Potential

• AP is a stereotyped change in
membrane potential
• If RMP moves past threshold,
membrane potential quickly moves to
+40 mV and then returns to resting
• Ionic basis of the AP:
• NA+ in: upswing of spike
• Diffusion, electrostatic pressure
• K+ out: downswing of spike

2.24
 Properties of the Action Potential
• The action potential:
• Is an “all or none” event: RMP either passes threshold or
doesn’t
• Is propagated down the axon membrane
• Notion of successive patches of membrane
• Has a fixed amplitude: AP’s don’t change in height to
signal information
• Has a conduction velocity (meters/sec)
• Has a refractory period in which stimulation will not
produce an AP (limits the firing rate)

2.25
 Synaptic transmission
• The Synapse is the space between neurons
• The synaptic gap or cleft

• Information must be transmitted across the synapse to other

neurons via the neurotransmitters.

• This is an electrochemical process

 The “synapse” is the physical gap between
pre- and post-synaptic membranes (~20-30
nMeters)
 Presynaptic membrane is typically an axon
 The axon terminal contains
 Mitochondria that provide energy for axon functions
 Vesicles (round objects) that contain neurotransmitter
 Cisternae that are a part of the Golgi apparatus:
recycle vesicles
 Postsynaptic membrane can be
 A dendrite (axodendritic synapse)
 A cell body (axosomatic synapse)
 Another axon (axoaxonic synapse)
 Postsynaptic density (thickening) lies under the
axon terminal and contains receptors for
transmitters
 100 trillion synapses
 Neurotransmitters

• chemical substances that reside in the axon

terminals

• They communicate to other neurons by binding

to receptors on neighboring neurons
 Chemical Communication
• The communication between neurons is chemical

• Neurotransmitter are
either neutralized by an
enzyme or taken back
up by the neuron that
released it in reuptake.

• At least 50 different
types of
neurotransmitters have
been identified
 Synaptic Transmission
• The neurotransmitters are released from the
vesicles and then attach to receptors located on
the postsynaptic neuron.

• These neurotransmitters are in contact with the

dendrite of the postsynaptic neuron only briefly.

• The chemical is almost immediately destroyed or

reabsorbed
 Neurotransmitters

• At least 50 different types of neurotransmitters

have been identified

• Acetylcholine
• GABA
• Serotonin
• Dopamine
• Norepinephrine
• Endorphins
Electrical Potentials across
Nerve Cell Membranes
• Neurons employ several different types of electrical signal
to encode and transfer information.
• The best way to observe these signals is to use an
intracellular microelectrode to measure the electrical
potential across the neuronal plasma membrane.
• A typical microelectrode is a piece of glass tubing pulled to a
very fine point (with an opening of less than 1 µm diameter)
and filled with a good electrical conductor, such as a
concentrated salt solution.
• This conductive core can then be connected to a voltmeter,
such as an oscilloscope, to record the transmembrane
voltage of the nerve cell.
What is microelectrode technique?
• in physiology, the technology used for measuring
the electric, concentration, and oxidation potentials
of cells and their parts and for influencing them
with electric current and various substances
(strictly localized).
First type of electrical Phenomenon
• The first type of electrical phenomenon can be
observed as soon as a microelectrode is inserted
through the membrane of the neuron.
• Resting membrane potential
Upon entering the cell, the microelectrode reports a
negative potential, indicating that neurons have a
means of generating a constant voltage across their
membranes when at rest. This voltage, called the
resting membrane potential, depends on the type of
neuron being examined, but it is always a fraction of a
volt (typically –40 to –90 mV).
Receptor potentials
• The electrical signals produced by neurons are caused by
responses to stimuli, which then change the resting
membrane potential.
• Receptor potentials are due to the activation of sensory
neurons by external stimuli, such as light, sound, or heat.
• For example, touching the skin activates Pacinian
corpuscles, receptor neurons that sense mechanical
disturbances of the skin.
• These neurons respond to touch with a receptor
potential that changes the resting potential for a fraction
of a second (Figure 2.1A).
Types of neuronal electrical signals.
In all cases, microelectrodes are used to measure changes in the resting membrane potential
during the indicated signals.

(A) A brief touch causes a receptor potential in a Pacinian corpuscle in the skin.

Pacinian corpuscle an encapsulated ending of a sensory nerve that acts as a receptor for
pressure and vibration.
 Pacinian corpuscle
• Pacinian corpuscle or lamellar corpuscle or Vater-Pacini corpuscle; is one of
the four major types of mechanoreceptors (specialized nerve ending with
adventitious tissue for mechanical sensation) found in mammalian skin.
• This type of mechanoreceptor is found in both glabrous (hairless) and
hirsute (hairy) skins, viscera, joints and attached to periosteum of bone,
primarily responsible for sensitivity to vibration.
• Few of them are also sensitive to quasi-static or low frequency pressure
stimulus.
• Most of them respond only to sudden disturbances and are especially
sensitive to vibration of few hundreds of Hz.
• The vibrational role may be used for detecting surface texture, e.g., rough
vs. smooth.
• Most of the Pacinian corpuscles act as rapidly adapting mechanoreceptors.
• Groups of corpuscles respond to pressure changes, e.g. on grasping or
releasing an object.
• Changes in potential are the first step in generating
the sensation of vibrations (or “tickles”) of the skin
in the somatic sensory system.

• Similar sorts of receptor potentials are observed in

all other sensory neurons during transduction of
sensory signals.
synaptic potentials
• Another type of electrical signal is associated with
communication between neurons at synaptic
contacts.
• Activation of these synapses generates synaptic
potentials, which allow transmission of information
from one neuron to another.
• An example of such a signal is shown in Figure 2.1B.
•
Types of neuronal electrical signals.

(B) Activation of a synaptic contact onto a hippocampal pyramidal

neuron elicits a synaptic potential.

In this case, activation of a synaptic terminal innervating a hippocampal pyramidal neuron

causes a very brief change in the resting membrane potential in the pyramidal neuron.
Synaptic potentials serve as the means of exchanging information in complex neural circuits
in both the central and peripheral nervous system.
Pyramidal neurons are also common in
subcortical structures such as the hippocampus
and the amygdala. They are named for their
shape: typically they have a soma (cell body)
that is shaped like a teardrop or rounded
pyramid.
Use of electrical signals—as
• The use of electrical signals—as in sending electricity
over wires to provide power or information—presents
a series of problems in electrical engineering.
• A fundamental problem for neurons is that their
axons, which can be quite long (remember that a
spinal motor neuron can extend for a meter or more),
are not good electrical conductors.
• Although neurons and wires are both capable of
passively conducting electricity, the electrical
properties of neurons compare poorly to an ordinary
wire.
• To compensate for this deficiency, neurons have
evolved a “booster system” that allows them to
conduct electrical signals over great distances
despite their intrinsically poor electrical
characteristics.
• The electrical signals produced by this booster
system are called action potentials (which are also
referred to as “spikes” or “impulses”).

• An example of an action potential recorded from

the axon of a spinal motor neuron is shown in
Figure 2.1C
Types of neuronal electrical signals.

(C) Stimulation of a spinal reflex produces an action potential in a spinal motor neuron.

An example of an action potential recorded from the axon of a spinal motor neuron is
shown in Figure 2.1C
 Spinal motor neuron
• Motor neurons of the
spinal cord are part of the
central nervous system
(CNS) and connect to
muscles, glands and
organs throughout the
body. These neurons
transmit impulses from
the spinal cord to skeletal
and smooth muscles
(such as those in your
stomach), and so directly
control all of our muscle
movements.26-Mar-2018
Recording passive and active electrical signals in a nerve cell.

(A) Two microelectrodes are

inserted into a neuron;
one of these measures
membrane potential while the
other injects current into the
neuron.
• If the current delivered in this way makes the
membrane potential more negative
(hyperpolarization), nothing very dramatic
happens.
• The membrane potential simply changes in
proportion to the magnitude of the injected
current.
• Such hyperpolarizing responses do not require any
unique property of neurons and are therefore
called passive electrical responses.
• A much more interesting phenomenon is seen if
current of the opposite polarity is delivered, so that
the membrane potential of the nerve cell becomes
more positive than the resting potential
(depolarization).
• In this case, at a certain level of membrane
potential, called the threshold potential, an action
potential occurs.
 Recording passive and active electrical signals in a nerve cell.

• (B) Inserting the voltage-

measuring microelectrode
into the neuron reveals a
negative potential, the resting
membrane potential.
• Injecting current through the
current-passing
microelectrode alters the
neuronal membrane
potential.
• Hyperpolarizing current
pulses produce only passive
changes in the membrane
potential. While small
depolarizing currents also
elict only passive responses,
• depolarizations that cause the
membrane potential to meet Action potentials are active responses in the sense
or exceed threshold that they are generated by changes in the
additionally evoke action permeability of the neuronal membrane.
potentials.
• The action potential, which is an active response
generated by the neuron,is a brief (about 1 ms)
change from negative to positive in the
transmembrane potential.
• Importantly, the amplitude of the action potential is
independent of the magnitude of the current used
to evoke it; that is, larger currents do not elicit
larger action potentials.
• The action potentials of a given neuron
are therefore said to be all-or-none,
because they occur fully or not at all.
• If the amplitude or duration of the
stimulus current is increased sufficiently,
multiple action potentials occur, as can
be seen in the responses to the three
different current intensities shown in
Figure 2.2B (right side).
• It follows, therefore, that the intensity
of a stimulus is encoded in the
frequency of action potentials rather
than in their amplitude.
• This arrangement differs dramatically
from receptor potentials, whose
amplitudes are graded in proportion to
the magnitude of the sensory stimulus,
or synaptic potentials, whose amplitude
varies according to the number of
synapses activated and the previous
amount of synaptic activity
• Because electrical signals are the basis of
information transfer in the nervous system, it is
essential to understand how these signals arise.
Remarkably, all of the neuronal electrical signals
described above are produced by similar
mechanisms that rely upon the movement of ions
across the neuronal membrane.
How Ionic Movements Produce
Electrical Signals
• Electrical potentials are generated across the
membranes of neurons—and, indeed, all cells—
because
(1) there are differences in the concentrations of specific
ions across nerve cell membranes, and
(2) the membranes are selectively permeable to some of
these ions.

These two facts depend in turn on two different kinds of

proteins in the cell membrane (Figure 2.3).
• Ion transporters and ion channels are responsible for ionic movements across neuronal
membranes.
• Transporters create ion concentration differences by actively transporting ions against
their chemical gradients.
• Channels take advantage of these concentration gradients, allowing selected ions to
move, via diffusion, down their chemical gradients.
How Ionic Movements Produce
Electrical Signals
• The ion concentration gradients are established by proteins
known as active transporters, which, as their name
suggests, actively move ions into or out of cells against
their concentration gradients. The selective permeability of
membranes is due largely to ion channels, proteins that
allow only certain kinds of ions to cross the membrane in
the direction of their concentration gradients.
• Thus, channels and transporters basically work against
each other, and in so doing they generate the resting
membrane potential, action potentials, and the synaptic
potentials and receptor potentials that trigger action
potentials.
take the case of a membrane that is permeable
only to potassium ions (K+). If the
concentration of K+ on each side of this
membrane is equal, then no electrical
potential will be measured across it (Figure
2.4A).
Increasing the KCl
concentration in
compartment 1 to 10
mM initially causes a
small movement of K+
into compartment
2 (initial conditions) until
the electromotive force
acting on K+ balances the
concentration gradient,
and the net
movement of K+
becomes zero (at
equilibrium).
However, if the concentration of K+ is not the same on the two sides, then an
electrical potential will be generated.

For instance, if the concentration of K+ on one side of the membrane (compartment

1) is 10 times higher than the K+ concentration on the other side (compartment 2),
then the electrical potential of compartment 1 will be negative relative to
compartment 2 (Figure 2.4B).
This difference in electrical potential is generated because the potassium ions flow
down their concentration gradient and take their electrical charge (one positive
charge per ion) with them as they go.

Because neuronal membranes contain pumps that accumulate K+ in the cell

cytoplasm, and because potassium-permeable channels in the plasma membrane
allow a transmembrane flow of K+, an analogous situation exists in living nerve cells.

A continual resting efflux of K+ is therefore responsible for the resting membrane

potential.
SUMMARY
Nerve cells generate electrical signals to convey information over substantial
distances and to transmit it to other cells by means of synaptic connections.
These signals ultimately depend on changes in the resting electrical potential
across the neuronal membrane. A resting potential occurs because nerve cell
membranes are permeable to one or more ion species subject to an
electrochemical gradient. More specifically, a negative membrane potential at
rest results from a net efflux of K+ across neuronal membranes that are
predominantly permeable to K+. In contrast, an action potential occurs when a
transient rise in Na+ permeability allows a net flow of Na+ in the opposite
direction across the membrane that is now predominantly permeable to Na+.
The brief rise in membrane Na+ permeability is followed by a secondary,
transient rise in membrane K+ permeability that repolarizes the neuronal
membrane and produces a brief undershoot of the action potential. As a result
of these processes, the membrane is depolarized in an all-or-none fashion
during an action potential.
When these active permeability changes subside, the membrane potential
returns to its resting level because of the high resting membrane permeability
to K+.
QUIZ 1
• Describe “knee-jerk” reflex briefly.