Ageing and Endings A

Neurophysiology and Skeletal Muscle Physiology

Membrane Transport

Simple
diffusion
Facilitated
diffusion

Description

Energy
required

Passive

Passive

Potassium
channels

ATP

SodiumpotassiumATPase
Sodiumglucose
symporter
Sodiumcalcium
exchanger

Primary
active
transport
Secondary
active
transport

Lipid-soluble substances can pass directly

through phospholipid bilayer
Obeys Ficks Law of diffusion rate or flux
Carrier (no continuous pore) or channel
(continuous pore) proteins in membrane
Molecules move down electrochemical
gradient
Compared to simple diffusion, has higher flux
rates, saturability, specificity and selectivity
Movement of molecules against
electrochemical gradient
Coupled to utilisation of chemical energy
Use of an existing ion gradient (generally
sodium) to move another molecule against its
electrochemical gradient
Co-transport/symport or countertransport/antiport

Four main types of membrane transport

Types

Ion
gradient

Examples
Water
Urea

of ion channel gating:

Extracellular ligand.
Intracellular ligand.
Voltage.
Stretch.
Background or leak channels.

Action Potential Propagation

Mechanism of regenerative propagation:
Triggering event opens sodium channels, causing depolarisation.
Depolarisation opens adjacent voltage-gated sodium channels.
Cycle of depolarisation and channel opening continues.
Transient deactivation of sodium channels allows unidirectional propagation.
No loss of signal, unlike passive spread of depolarisation (all or nothing).
Conduction velocity is determined by how easily and quickly local polarisation can
cause the adjacent section of axon to reach threshold and active voltage-determined
sodium channels, and proportional to:
Myelination (markedly increases membrane resistance, but does not affect
internal resistance).
Axon diameter.
Temperature.

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Ageing and Endings A

Neurophysiology and Skeletal Muscle Physiology

Action potential propagation

Action Potential Phases
SODIUM
CHANNELS

Closed

Open

Able to open

Closed

Closed

Unable
to open

Able to open

Resting State

Hyperpolarisation
(Refractory Period)

Repolarisation

Depolarisation

Depolarising Stimulation

Resting State

MEMBRANE POTENTIAL
(Mv)

65 (ENa+)

Threshold
-70
-90 (EK+)
POTASSIUM
CHANNELS

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Closed

Open

Closed

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Ageing and Endings A

Neurophysiology and Skeletal Muscle Physiology

Stages of Synaptic Transmission
Depolarisation reaches presynaptic terminal.
Opening of voltage-gated calcium channels and increased calcium concentration in
presynaptic terminal.
Calcium triggers fusion of synaptic vesicles to presynaptic membrane (via v-SNAREs on
vesicle and t-SNAREs on membrane).
Neurotransmitter release and diffusion across synaptic cleft.
Binding of neurotransmitter with ligand-gated postsynaptic channels.
Neuromuscular Junctions and Motor Units
Neuromuscular junction (NMJ) synapse between a motor axon and a muscle fibre:
One per muscle fibre, but multiple per axon.
Involves spatially-opposed specialisation of both the axon and the muscle fibre.
Neurotransmitter is acetylcholine (ACh)

Receptor type

Neuromuscular Central nervous system

junction

Autonomic nervous
system

Excitatory
ionotropic
~1,000
40-50 mV

Excitatory or inhibitory
metabotropic
~20
Profound

Excitatory or inhibitory
ionotropic and metabotropic
~2
0.2-0.5 mV

Active zones
Potential caused
by single axon
Comparison of NMJ to other synapse types

Typical end plate potential (EPP) causes vesicles to be released from ~100 (out of
~1,000) active zones, causing a potential of 40-50 mV.
EPP terminated by:
Degradation of ACh by acetylcholinesterase.
Diffusion of ACh out of cleft.

Axoplasmic Transport

Anterograde
Mechanism
Substances
transported
Speed of
transport

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Retrograde

Microtubule system of tracks, using kinesin/dynein motors that are

energised by ATP
Synthesised proteins (packed by budding off Material absorbed at
in membrane-enclosed vesicles from Golgi
terminals (e.g. trophic
apparatus
factors
~250 mm day
Fast - ~400 mm/day
Slow (cytoskeletal and soluble proteins)
~2 mm/day

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Ageing and Endings A

Neurophysiology and Skeletal Muscle Physiology

Skeletal Muscle Structure and Proteins
Skeletal muscle fibre single elongated, multinucleated (~1,000 nuclei per
centimetre), contractile cell.
Fibril/Myofibril sub-division of a skeletal muscle fibre composed of 100-400
sarcomeres.
Sarcomere repeating unit of skeletal muscle, being the smallest unit of contraction
(defined as one Z line to the next).
Striated appearance of skeletal muscle fibres due to filamentous proteins:
Myosin (thick and dark).
Actin (thin and light).
Z line anchoring point of actin filaments.
M line consists of proteins that act to stabilise myosin filaments.

Sarcomere structure
Group

Protein

Band/Line

Role

Contractile

Myosin
Actin
Troponin
Tropomyosin
M proteins

A band
I band
I band
I band
M line

Muscle contraction

C proteins
Alpha-actinin
Titin

A band
Z line
Z line to M line (A
band to I band)

Regulatory
Structural

Nebulin

Major skeletal muscle proteins

Prevention of constant contraction (rigor)

Securing of myosin in centre of
sarcomere
Stabilisation of A band
Stabilisation of Z line
Stabilisation of sarcomere and prevention
of overextension
Stabilisation of actin

Sarcolemma cell membrane of skeletal muscle fibre.

Connective tissue sheaths:
Endomysium surrounding individual fibres.
Perimysoium surrounding muscle fascicles/bundles.
Epimysium surrounding entire muscle.
Structural protein framework allows transmission of force of contraction laterally to
sarcolemma, basal lamina and connective tissue sheaths, from which transmission can
occurs along sheaths to tendons.

Matt Schiller

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Ageing and Endings A

Neurophysiology and Skeletal Muscle Physiology

Skeletal Muscle Operation
Sliding filament theory:
Muscle operates by the relative sliding of actin and myosin filaments.
Sliding is caused by cross-bridges that project from myosin filaments interacting
with specific sites on actin filaments.
Cross-bridges have a limited range of movement (~10 nm).
Cross-bridges act cyclically, splitting a molecule of ATP per cycle.
Maximum force is produced when the maximum number of sites of interaction are
aligned (of which there are a finite number).
Regulation of contraction by tropomyosin and troponin:
Troponin covers binding sites on actin filaments in resting state.
Troponin has a calcium binding site for calcium released by nervous stimulation.
When calcium binds to troponin, a conformation change occurs that exposes
binding sites on actin and allows skeletal muscle contraction.
Stages of muscle contraction and relaxation:
Action potential in surface membrane.
Action potential conducted down transverse tubules (T tubules) to sarcoplasmic
reticulum.
Excitation-contraction coupling opening of channels in sarcoplasmic reticulum.
Release of calcium from sarcoplasmic reticulum.
Operation of cross-bridge cycle between contractile proteins (contraction).
Calcium pump in sarcoplasmic reticulum causes re-compartmentalisation of
calcium.
Cross-bridge detachment occurs (relaxation).
Types of contraction:
Isotonic contraction where constant tension produces a change in muscle
length.
Isometric contraction where tension change is recorded while muscle length
remains constant.
Twitch and fused tetanus:
Twitch response of skeletal muscle to a single action potential.
Fused tetanus response of skeletal muscle to a high frequency of action
potentials (about four times that of a twitch).
Motor unit a single motor neuron and the muscle fibres that it innervates (vary in
size from several to over 1000 muscle fibres).
Size principle smaller motor units are recruited before larger ones, permitting the
application of fine forces in small increments.
Muscle force can be increased by increasing:
Frequency of motor nerve firing.
Motor unit recruitment.

Speed of contraction
Force
and relaxation

Size

Fatigue
resistance

Fast fibres Fast

High
Large Low
Slow fibres Slow
Low
Small High
Comparison of fast and slow skeletal muscle fibres
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