296

184 Neuron Structure and Function

Key Idea: Neurons are electrically excitable cells that signals from receptors (in the central nervous system)
are specialized to process and transmit information via to effectors. Neurons consist of a cell body (soma) and
electrical and chemical signals. Increased axon diameter and long processes (dendrites and axons). Conduction speed
myelination both increase conduction speed along a neuron. increases with axon diameter and with myelination. Faster
Neurons transmit information in the form of electrochemical conduction speeds enable more rapid responses to stimuli.

Motor (efferent) neuron Axon: A long extension of the cell transmits

the nerve impulse to another neuron or to
Transmits impulses from the CNS an effector, e.g. muscle. Motor axons may
to effectors (muscles or glands). be very long and, in the peripheral nervous
system, many are myelinated.

Dendrites are thin processes from

the cell body that receive stimuli.

lse
pu tion
Im ec Axon branches of motor
ir
Cell body or soma Myelin d neurons have synaptic
containing the organelles sheath knobs at each end. These
to keep the neuron alive release neurotransmitters,
and functioning. Node of Ranvier, chemicals which transmit
a gap in the the impulse between
myelin sheath neurons or between a
neuron and a muscle cell.

Axon hillock region

(generation of action potential)

Where conduction speed is important, Myelinated Neurons

the axons of neurons are sheathed
Diameter: 1-25 µm
within a lipid and protein rich substance
Conduction speed: 6-120 ms-1
called myelin. Myelin is produced by
oligodendrocytes in the central nervous
system (CNS) and by Schwann cells in
the peripheral nervous system (PNS). At
intervals along the axons of myelinated
neurons, there are gaps between Myelin
neighbouring Schwann cells and their
sheaths. These are called nodes of Node of Ranvier
Ranvier. Myelin acts as an insulator,

Roadnottaken
increasing the speed at which nerve Axon
impulses travel because it prevents ion
Myelin layers wrapped
flow across the neuron membrane and
Schwann cell wraps around axon TEM cross section through
forces the current to 'jump' along the
only one axon and a myelinated axon
axon from node to node. produces myelin

1. What is the function of a neuron? transmit information in the form of electrochemical Motor (efferent) neuron
signals from receptors (in the central nervous system) to effectors.

2. What factors increase the speed of conduction along a neuron?

3. How does myelination increase the speed of nerve impulse conduction?

4. What is the advantage of faster conduction of nerve impulses?

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 297

185 The Nerve Impulse

Key Idea: The nerve impulse is an electrical signal that the membrane quickly depolarizes by allowing positively
propagates along the axon of a neuron. charged sodium ions to flood into the cell via voltage gated
Nerve cells (neurons) maintain a polarized membrane when ion channels. To restore and maintain the polarity of the cell
not transmitting a signal. When induced to transmit a signal sodium-potassium pumps move sodium ions out of the cell.

Membrane potential

The resting neuron The nerve impulse

+ +
Na Na
Direction of impulse
+ + + + + + + + + + + + + + + + + + + + + + + + – – – – – + + + + +
– – – – – – – – – – – – – – – – – – – – – – – – – + + + + + – – – – –
– – – – Axon – – – – – – – – – – – – – – – – – – – – – + + + + + – – – – –
+ + + + + + + + + + + + + + + + + + + + + + + + – – – – – + + + + +

Area of impulse
Next area to be stimulated Area returning to resting state

When a neuron is not transmitting an impulse, the inside When a neuron is stimulated, the distribution of charges on
of the cell is negatively charged relative to the outside and each side of the membrane briefly reverses. This process
the cell is said to be electrically polarized. The potential of depolarization causes a burst of electrical activity to
difference (voltage) across the membrane is called the pass along the axon of the neuron as an action potential.
resting potential. For most nerve cells, this is about -70 As the charge reversal reaches one region, local currents
mV. Nerve transmission is possible because this membrane depolarize the next region and the impulse spreads along
potential exists. the axon.

Maintaining membrane potential

▶ Membrane potential is produced by the use of a sodium/ Outside cell Inside cell
potassium pump. Sodium from inside the cell is pumped 1
K+
out of the cell and potassium from outside the cell is
pumped into the cell. However, because three sodium Na+
ions are pumped out of the cell for every two potassium
ions that are pumped in, a positive charge builds up on
the outside of the cell relative to the inside of the cell. 2

▶ This produces a potential difference (voltage) across

the cell membrane, called the resting potential. Because
membrane potential is defined by the interior of the cell
relative to the exterior of the cell, the membrane potential
will be negative.

▶ Note that the sodium-potassium pump produces a 3

sodium ion concentration gradient that would drive
sodium into the cell, while also producing a potassium ion
concentration that would drive potassium from inside the
cell to outside the cell.

1. (a) Define the term 'resting potential':

(b) Define the term 'action potential':

2. How is the resting potential produced and maintained?

3. How is an action potential propagated along a nerve fibre (axon)?

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298

186 The Speed of Nerve Impulses

Key Idea: The diameter and myelination of neurons affects the speed of transmission in the nerve fibres of a frog was
the speed of the nerve impulse. around 30 ms-1. Classic experiments carried out in the 1930s
Nerve impulses move along a nerve fibre at a variety of by J.B. Hursh on the mylenated nerve fibres of cats and R.
speeds, from less than 1 ms-1 to nearly 200 ms-1 in the shrimp J. Pumphrey et al on the unmylenated nerve fibres of squid
giant nerve fibre. The first attempts to measure these speeds showed the effect of mylenation and the diameter of the
were in 1849 by Hermann von Helmholtz, who showed that nerve fibre on impulse speed.

Nerve fibre diameter vs speed of transmission

Adapted from J.B. Hursh and R. J. Pumphrey et al

Cat
Diameter (µm) 3.0 3.2 3.4 4.0 4.4 5.0 6.2 6.2 6.4 7.6 8.0 8.8 9.0 10.0

Conduction
17.0 21.9 24.3 36.4 30.4 26.7 42.5 46.1 43.7 60.7 46.2 55.4 54.5 80.0
velocity (ms-1)

Squid
Diameter (µm) 30 40 43 50 108 130 130 148 280 294 357 413 415 567

Conduction
2.2 4.7 6.4 5.6 5.9 7.8 5.9 8.2 15.0 11.0 13.8 17.4 18.6 20.0
velocity (ms-1)

80

70

60
Nerve fibre conduction speed (ms-1)

50

40

30

20

10

0
0 1 2 3 4 5 6 7 8 9 10 11 12
Diameter cat nerve fibre (µm)

0 100 200 300 400 500 600

Diameter squid nerve fibre (µm)

1. Plot the diameter and speed of transmission of both cat (mylenated) and squid (nonmylenated) nerves. Include a key:

2. Calculate the coefficient of determination (R2) for each set of data (see Activity 134 for details):

(a) Cat

(b) Squid:

3. Which appears more important for speed of transmission: mylenation, or nerve fibre diameter?

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188 Electrical Impulses in the Nerve

Key Idea: A nerve impulse occurs in response to a stimulus is this property that enables neurons to transmit electrical
and involves the transmission of a membrane depolarization impulses. When a nerve is stimulated, a brief increase in
along the axon of a neuron. membrane permeability to Na+ temporarily reverses the
The plasma membrane of cells, including neurons, contain membrane polarity (a depolarization). After the nerve
sodium-potassium ion pumps that actively pump sodium ions impulse passes, the sodium-potassium pump restores the
(Na+) out of the cell and potassium ions (K+) into the cell. The resting potential. In neurons that are unmyelinated, the nerve
action of these ion pumps in neurons creates a separation impulse is propagated along the entire length of the nerve.
of charge (a potential difference or voltage) either side of In myelinated neurons, the nerve impulse jumps between the
the membrane and makes the cells electrically excitable. It nodes of Ranvier, speeding up conduction (below).

Axon myelination is a feature of vertebrate Depolarized region

nervous systems and it enables them (node of Ranvier)
to achieve very rapid speeds of nerve Saltatory conduction in
– myelinated axons
conduction. ––
+ ––
In a myelinated neuron, action potentials + –
are generated only at the nodes, which ++
is where the voltage-gated channels +++ Axon
occur. The axon is insulated so the action ––– + ++
potential at one node is sufficient to trigger – Schwann cell
––
an action potential in the next node and
the impulse 'jumps' along the axon (called
saltatory conduction). This contrasts with
a non-myelinated neuron in which voltage-
gated channels occur along the entire
The charge will passively depolarize
length of the axon. the adjacent node of Ranvier to
As well as increasing the speed of threshold, triggering an action
potential in this region and
conduction, the myelin sheath reduces
subsequently depolarizing the next
energy expenditure because the area over node, and so on. Action potential is
which depolarization occurs is less. The generated in the
number of sodium and potassium ions axon hillock region.
that need to be pumped to restore resting
potential is, therefore, also less. Myelinated axons have gated
channels only at their nodes.
The electrical impuse
The depolarization in an axon can be shown as a
+50 mV change in membrane potential (in millivolts). A stimulus
must be strong enough to reach the threshold potential
Membrane potential

before an action potential is generated. This is the

voltage at which the depolarization of the membrane
becomes unstoppable.
0 ` When at rest, voltage-gated Na+ channels are
closed (1).
` Voltage-gated Na+ channels open and the
membrane depolarizes as Na+ floods into the
-50 mV Threshold potential cell (2).
` Voltage gated Na+ channels close and voltage-
Resting potential
-70 mV gated K+ channels open, allowing K+ to move out of
the cell (3).
` A delay in closing voltage gated K+ causes
hyperpolarization. Na+/K+ pumps restore the
0 1 2 3 4 5 membrane potential (4). During this refractory
Elapsed time in milliseconds period, the nerve cannot respond, so nerve
impulses are discrete.

1. What happens to a nervous signal if the threshold potential is not reached?

2. Contrast the placement of voltage-gated ion channels in myelinated and nonmyelinated neurons:

3. Based on the oscilloscope trace above, how many times can a nerve cell respond per second?

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 299

187 Chemical Synapses

Key Idea: Synapses are junctions between neurons, or the membrane of the post-synaptic (receiving) cell. Arrival
between neurons and receptor or effector cells. Nerve of an action potential at the axon terminal causes release
impulses are transmitted across synapses. of the neurotransmitter, which diffuses across the cleft and
Action potentials are transmitted across junctions called produces an electrical response in the post-synaptic cell (an
synapses. Almost all synapses in vertebrates are chemical example of signal transduction). Cholinergic synapses are
synapses, which involve the diffusion of a signal molecule or named for the neurotransmitter they release, acetylcholine
neurotransmitter from one cell to another. Chemical synapses (ACh). In the example pictured below, ACh results in
can occur between two neurons, between a receptor cell depolarization (excitation) of the post-synaptic neuron.
and a neuron, or between a neuron and an effector, e.g. Unlike electrical synapses, in which transmission can occur
muscle fibre or gland cell. The synapse consists of the axon in either direction, transmission at chemical synapses is
terminal (synaptic knob), a gap called the synaptic cleft, and always in one direction (unidirectional).

The structure of a cholinergic synapse

Step 2: ACh diffuses across the
synaptic cleft to receptors on the
Step 1: The arrival of an action potential at receiving membrane. Diffusion
the axon terminal causes an influx of calcium across the cleft delays the impulse
ions and induces the vesicles to release the transmission by about 0.5 ms.
neurotransmitter acetylcholine (ACh) into the
synaptic cleft. A high frequency of impulses Synaptic
results in release of more neurotransmitter. knob

Step 3: ACh binds to

receptors on the receiving
Di Ca2+ (post-synaptic) membrane.
rec
tio Ca2+
no
f im
pu
lse

Axon of the
neuron.

Synaptic vesicles Step 4: Ion channels in the

containing ACh. membrane open, causing an
influx of Na+. This response
may or may not reach the
threshold for an action
potential. The strength of the
The response of a receiving cell to the arrival response is related to how
of a neurotransmitter depends on the nature much ACh is released.
of the cell itself, its location in the nervous
system, and the particular type of Mitochondria
neurotransmitter involved. Cholinergic provide energy Step 5: The Ach is quickly deactivated by
synapses are found in the autonomic nervous for active transport. the enzyme acetylcholinesterase located on
system (which controls unconscious bodily the membrane. Components of the
functions) and the neuromuscular junction neurotransmitter are actively reabsorbed
(between motor neurons and muscle cells). back into the synaptic knob, recycled, and
repackaged back into vesicles.

1. (a) What is a synapse?

(b) What defines a cholinergic synapse?

2. What causes the release of neurotransmitter into the synaptic cleft?

3. Why is there a brief delay in impulse transmission across the synapse?

4. What determines the strength of the response in the receiving cell?

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 301

189 Chemicals at Synapses

Key Idea: Many exogenous chemicals may increase or the reuptake of neurotransmitters by acting on transporter
decrease the effect of neurotransmitters at synapses. proteins that remove a neurotransmitter from the synaptic
Exogenous chemicals, e.g. drugs, may act at synapses, either cleft. Many recreational and therapeutic drugs work through
mimicking or blocking the usual effect of a neurotransmitter, their action at synapses, controlling the response of the
whether it be excitatory or inhibitory. They may also block receiving cell to incoming action potentials.

Blocking transmission Blocking reuptake

Neonicotinoids are a group of neuro-active insecticides. They Once signal transmission is complete, the neurotransmitter
mimic the action of acetylcholine on cholinergic receptors, is reabsorbed by the presynaptic cell, or broken down
but unlike acetylcholine they are not broken down by the by enzymes. Certain drugs can block the reuptake of the
enzyme acetylcholinesterase. This means they can bind to the neurotransmitter, allowing it to continue to stimulate the
cholinergic receptors permanently and block the transmission receptor. The drug cocaine is an example of this.
of nerve signals.
Cocaine blocks the reuptake of the neurotranmitter dopamine
This has two effects. Initially, it leads to over-stimulation as by binding to the dopamine transporter. This allows dopamine
the nerve continues to fire until ion exchange ceases, and to accumulate in the synaptic cleft, stimulating the dopamine
eventually to paralysis as the nerve can not 'reset'. receptors on the postsynaptic cell.

Dopamine
Cocaine blocks
transporter
dopamine transporter
Neonicotinoids mimic stopping the reuptake
ACh action, binding of dopamine
to ACh receptors and
opening Na+ gates
ACh
Dopamine
Synaptic cleft
Cocaine
Membrane of
post-synaptic Synaptic cleft
neurone

Post-synaptic Membrane of
Na+ Sodium-gated Dopamine
neurone post-synaptic neurone
channel receptor

Result: Depolarisation and impulse generation

Post-synaptic
neurone
Result: Continued dopamine stimulus

Neonicotinoids are commonly used as insecticides because Dopamine is part of the reward-motivation pathways in the
of their far greater specificity to insects than other animals. brain, so cocaine use produces a euphoric effect and repeated
However, they are a broad spectrum insecticide and so can kill use leads to addiction. However, the body quickly builds a
both target and non target insects. The use of neonicotinoids tolerance to cocaine so for each use more is required for the
is now restricted in many countries due to their (unintended) same effect.
effects on honey bees.

1. Describe three ways in which exogenous chemicals can affect signalling at synapses:

2. (a) Explain how neonicotinoids work:

(b) Explain why neonicotinoid based pesticides have been regulated in many countries:

3. Explain how cocaine produces its effects:

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302

190 Summation at Synapses

Key Idea: The totality of signals from presynaptic cells (decrease membrane potential) by opening Na+ channels.
determines if an action potential is reached in the post Inhibitory neurotransmitters e.g. GABA, increase the
synaptic cell. polarization of the membrane (hyperpolarization, increase
The nature of synaptic transmission in the nervous system membrane potential) by opening Cl- channels. It is the
allows the integration (interpretation and coordination) of sum of all excitatory and inhibitory inputs that leads to the
inputs from many sources. These inputs can be excitatory final response in a post-synaptic cell. Synaptic integration
(causing depolarization) or inhibitory (making an action is behind all the various responses we have to stimuli. It is
potential less likely). Excitatory neurotransmitters, e.g. also the most probable mechanism by which learning and
acetylcholine, reduce the polarization of the membrane memory are achieved.

Summation at synapses
Graded postsynaptic responses may sum to produce an action
potential. Impulse transmission across chemical synapses has several Pre-synaptic neuron
advantages, despite the delay caused by neurotransmitter diffusion.
Action
Chemical synapses transmit impulses in one direction to a precise potential
location and, because they rely on a limited supply of neurotransmitter, Direction Threshold
they are subject to fatigue (inability to respond to repeated stimulation). of impulse
This protects the system against overstimulation.

Synapses allow inputs from many sources to be integrated. The

response of a post-synaptic cell is often not strong enough on its own
to generate an action potential. However, because the strength of
the response is related to the amount of neurotransmitter released, Multiple synapses with
subthreshold responses can sum together to produce a response in the Soma of post- the post-synaptic cell
post-synaptic cell. This additive effect is called summation. Summation synaptic neuron
can be temporal or spatial (below).

EPSP -IPSP cancellation Spatial summation Temporal summation

Action Action
potential potential

Threshold Threshold Threshold

Hyperpolarization
No action potential

If an inhibitory signal reaches a synapse at Impulses from spatially separated axon Several impulses may arrive at the synapse
the same time as an excitatory signal, the terminals may arrive simultaneously at in quick succession from a single axon. The
changes to membrane potential cancel out, different regions of the same post-synaptic individual responses sum to reach threshold
producing no response. neuron. The responses from the different and produce an action potential in the post-
places sum to produce an action potential. synaptic neuron.

1. (a) Explain what is meant by summation:

(b) In simple terms, distinguish between temporal and spatial summation:

2. How does hyperpolarization reduce the chance of an action potential occurring?

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 303

191 The Perception of Pain

Key Idea: Nociceptors are free nerve endings in the skin that responded to by free nerve endings called nociceptors.
send pain signals to the brain. These are found in the skin, muscles, joint, and anywhere
Pain is an important stimulus. It tells us when damage has where harmful stimuli are likely to be felt. Nociceptors
occurred and causes us to either deal with or move away respond to a wide range of stimuli including mechanical,
from the object causing pain. Stimuli that cause pain are thermal, and chemical stimuli.

Receptors in the skin

Nociceptor stimuli
Meissner's corpuscles
(light touch) Chemical stress Mechanical stress Thermal stress
Nociceptor (pain)

James Heilman CC 3.0

Kronoman CC 3.0
E.g lactic acid in Exogenous chemical E.g. pressure from E.g. skin burn
muscles. e.g. bee sting cut/ puncture

Damaged
cells
Acid sensitive Mast cells release Mechanically Heat sensitive
ion channels histamines and gated ion ion channels
(ASICs) serotonin channels

Pacinian corpuscle
Pressure or
(pressure)
Acid build up Chemical stretch Heat
(H+) molecule

Enzyme

Acid sensing Na+ or K+ Pressure- Enzyme-

ion channel channel sensitive Na+ controlled ion
channel channel

Nociceptors are free nerve endings found Depolarization Depolarization Depolarization Depolarization
in the dermis. They respond to damaging
stimuli by sending pain signals to the brain. Damaged cells release
A graze (above) or burn is painful because substances that
they trigger many of these receptors. depolarize nociceptors.

1. What is the purpose of nociceptors?

2. What kind of stimuli do nociceptors respond to?

3. For each of the following, identify which type of receptor nociceptors responds:

(a) Placing finger into too hot water:

(b) Banged shin:

(c) Standing on a sharp stone in bare feet:

4. Explain why nociceptors are also called polymodal receptors:

5. Capsaicin, found in chilli peppers, activates heat activated ions channels in nociceptors. What is the result of this?

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304

192 What is Consciousness?

Key Idea: Consciousness is an emergent property of billions Studies tend to show that consciousness is not limited to any
of neurons working together. one part of the brain, although some parts of the brain appear
What exactly consciousness is and what causes it is a matter to be more important in this regard than others. How the
of intense scientific research and debate. Consciousness different parts of the brain communicate is also important.
is an emergent property of billions of neurons in the brain In a conscious person, signals are passed around the brain
working together to produce the state of being aware. At but in an unconscious person, signals tend to remain in the
its simplest, consciousness is being aware of internal and same place as where they were experienced. For example,
external sensations e.g. your internal thoughts and feelings, an unconscious person's brain may receive pain signals but
or your effects on the environment. However, discovering because those signals are not passed around the brain, the
what makes us or any other organism conscious is difficult. person does not experience the sensation of pain.

Consciousness as an emergent property.

▶ Emergence is the property of small building blocks together
forming bigger units that have different properties than just
the sum of their parts. It is complexity arising from simplicity.
In the case of consciousness, this is a property that emerges
from many millions or billions of neurons working together.
None of the neurons is conscious but put together with
billions of interconnections, a conscious organism emerges.
Levels of consciousness
▶ A simple way of thinking about an organism's consciousness
is asking the question, 'do organisms have experiences?'.
At which level of complexity do organisms stop having
experiences and become unconscious of their surroundings?

Georgia Pinaud PD
▶ Consciousness probably exists at various levels, rather than
simply being conscious or not conscious.
▶ The more sensory inputs an organism has, the more aware
of its surroundings it will be. However, that may only be one
Dogs normally fail the mirror test because they don't use sight as
level of consciousness. The better an organism can process
their primary recognition sense. Tests using scent show that dogs
the information it receives, the more conscious it is likely to may in fact be self-aware.
be. For example, instead of simply reacting to a stimulus in
a repetitive way, an organism might first decide what that 1. What is emergence and how does it relate to
stimulus means and then decide on a course of action. consciousness?
▶ Being aware of others and how they relate to you is another
level of consciousness. This could include anticipating
another's actions or the effect of your actions on them. This
requires an awareness of time and self, or communicating
with others to carry out a task, which requires being able to
empathise with others.
Consciousness in nonhuman animals
▶ It is difficult enough to study what exactly consciousness
is and where it comes from in humans, let alone other
mammals, but what about non mammalian animals,
2. (a) Why is it difficult to determine where consciousness
or invertebrates?
comes from in an organism?
▶ We know that many birds display consciousness on similar
levels as some mammals, e.g. squirrels and scrub jays
show similar behaviours when storing or burying food. They
will re-hide food they buried if they think they were being
watched, showing that they are conscious of others' possible
intentions. Birds such as the Eurasian magpie have also
passed the 'mirror test', showing that they are
self-aware.
▶ We also know that an octopus can plan ahead. Octopuses
have been observed carrying coconut shells or other objects
for hiding in at a later time; they can remember people, and (b) Why is it difficult to determine that consciousness
can solve puzzles. A study in 2022 seemed to show that they even exists in other animals?
experience pain on an emotional level. However, invertebrate
brains are not like mammalian brains. Indeed, although an
octopus has a central brain, it is unlike that of a vertebrate's.
Each of its arms also contains a dense set of neurons that
acts as a 'mini-brain.' These neurons allow the arms to act
independently, although the octopus can control them
individually if it wants to. This makes the question of where
consciousness resides even more difficult.

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