Psychology 110: Biological Psychology

Lecture 13: Motor control

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 Outline

• we will look at the basic components of the motor system: muscles,

motoneurons and proprioceptive receptors.

• we will then look at the hierarchy of processing that exists in the motor
system, beginning with spinal reflexes, and continuing to look at the
primary motor cortex, and motor association cortices.

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 Muscles

Muscles can exert force only by contraction. The movement produced by

this contraction is determined by the muscle’s points of attachment
(tendons).

E.g. because of their attachments,

contraction of the biceps flexes the
arm, while contraction of the triceps
extends the arm (we call this
muscular synergism).

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 Muscle fibers

Muscle fibers consist of many individual fibers bound together with

connective tissue. 4
 Mechanism of muscle contraction

Muscle fibers consist of two types of interlocking protein – actin and myosin.

During contraction the myosin heads bind to actin, and then bend to slide
filaments past one another, shortening the muscle.
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 Types of muscle fiber

i). Fast muscle:

• reacts quickly, fatigues rapidly

• used for activities in which muscle tension changes frequently e.g.
locomotion
• ‘white meat’ in poultry (used to beat wings)

ii). Slow muscle:

• react slowly but resist fatigue

• used to maintain posture
• ‘dark meat’ in poultry (used for standing)

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 Innervation of muscle fiber

Motoneurons leave the ventral root of the spinal cord and branch multiple
times before synapsing with individual muscle fibers at the neuromuscular
junction.

ACh is released and elicits an

action potential in the muscle
fiber itself.

The influx of Ca2+ and Na+

into the fiber triggers the
molecular changes in action
and myosin that produce
contraction.

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 Innervation ratio

The innervation ratio is the ratio of motor axons to muscle fibers.

Lower innervation ratios permit more fine motor control.

For example, the muscles involved in eye movements have about one
motoneuron for every three muscle fibers (1:3 innervation ratio), while
those for the leg have one motoneuron controlling several hundred fibers
(~1:300). 8
 Motoneuron recruitment

Motoneurons differ in size and electrophysiological properties.

Large motoneurons have wide diameter axons to conduct impulses fast, but
are difficult to initially excite.

Small motoneurons have small axons and conduct impulses slowly, but are
easy to initially excite.

SIZE PRINCIPLE:

Muscle tension is increased by recruitment of

increasing numbers of motoneurons in fixed
order according to their size. Weak stimulation
activates only the small neurons, and stronger
stimulation recruits additional larger neurons.
Henneman The principle is similar to range fractionation.

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 Sensory feedback (proprioception)

Proprioception refers to the detection of the position and movement of our

body.

There are two proprioceptive sensory structures that are associated with
muscles:

i). Muscle spindles

ii). Golgi tendon organs

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 Muscle spindles

A muscle spindle consists of a nerve fiber wrapped around an intrafusal

muscle fiber (extrafusal muscle fibers are responsible for the main
contraction of the muscle).

If the muscle fiber is stretched this stretches the nerve fiber and causes
the nerve to fire action potentials.

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 Spinal reflexes: the stretch reflex

1. Weight is placed in
subject’s hand.

2. MA is stretched.

3. SNA is excited.

4. SNA connects
monosynaptically with
MNA exciting the
stretched muscle.

5. Via an interneuron,
SNA also inhibits MNB
ensuring that the
antagonistic muscle
(MB) is relaxed when
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MA is contracted.
 Muscle spindles: efferent nerves

Muscle spindles also receive input from efferent nerves called gamma
motoneurons (alpha motoneurons go to main muscle fibers).

Activity in gamma fibers causes a contraction in the length of the spindle,

which modifies its sensitivity.

The importance of this system is reflected in the fact that 30% of all
motoneurons are gamma.

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 Golgi tendon organ

Golgi tendon organs detect stretch in the tendon, not the muscle, and are
sensitive to contraction.

The Golgi tendon organ has feedback inhibition on the motoneurons

stimulating the muscle. If they detect a muscle overload (too much
contraction) they fire, and inhibit the neuron that is causing the muscle
contraction.

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 Outline

• we will begin by looking at the basic components of the motor system,

including muscles, motoneurons and proprioceptive receptors.

• we will then look at the hierarchy of processing that exists in the motor
system, beginning with spinal reflexes, and continuing to look at the
primary motor cortex, and motor association cortices (premotor cortex
and the supplementary motor area.

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 Spinal reflexes

An animal with damage to its spinal cord is still capable of many complex
behaviors. For example, the animal can often stand, and if stimulated, walk
as well.

Spinal cord contains a simple oscillator circuit, called a central pattern

generator, that can produce simple locomotion.

At first cell A transmits the

same number of spikes as
cell B

Negative feedback reaches B

out of phase with the input
because of synaptic delays,
causing rhythmic output.

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 Corticospinal pathway

Consists of motoneurons that travel from the motor cortex, through the
brainstem, and down the spinal cord.

Cross over in the medulla so that the left motor cortex controls the right side
of the body and vice versa.
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 Primary motor cortex (M1)

The hands, lips and tongue are over represented in M1.

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 Spot the difference?

SENSORY HOMUNCULUS MOTOR HOMUNCULUS

In general, areas with many motor outputs also receive a lot of

somatosensory input. There are some subtle differences, e.g. human ears
receive little motor output, and our eyes have few somatosensory inputs.
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 Primary motor cortex (M1)

• lesions of M1 cause paralysis

• electrical stimulation elicits twitches and jerks in limbs

• M1 neurons show direction selectivity for movements

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 Muscles or movements?

Monkey must move joystick from central location to one of eight peripheral
locations (indicate by yellow dot), but by altering the monkey’s hand
orientation, one can alter which muscles are used to make the movement.

One third of M1 neurons encode which muscles are used, one half encode the
movement direction irrespective of the muscles used. 21
 Premotor cortices

Supplementary motor area (SMA) a.k.a. medial premotor cortex (MPC)

Premotor cortex a.k.a. lateral premotor cortex (LPC)

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 LPC: involvement in conditional learning

Conditional tasks involve learning an arbitrary relationship (‘If X then Y’).

TASK 1: Color instructs either pull or turn

a lever.

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 LPC: involvement in conditional learning

Conditional tasks involve learning an arbitrary relationship (‘If X then Y’).

TASK 1: Color instructs either pull or turn

a lever.

TASK 2: Color instructs which shape to

choose.

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 LPC: involvement in conditional learning

Lesions of LPC impair conditional tasks, but

only when the stimulus instructs a specific
motor act.

(UC = unoperated control)

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 Medial premotor cortex (MPC)

Monkeys were trained to raise their

arms for a peanut. Monkeys with MPC
lesions were poor at doing this, failing to
raise their arms as many times per
minute as unoperated controls.

NO CUE CUED

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 Medial premotor cortex (MPC)

Monkeys were trained to raise their

arms for a peanut. Monkeys with MPC
lesions were poor at doing this, failing to
raise their arms as many times per
minute as unoperated controls.

If the monkeys heard a cue tone

however, which told them to raise their
arms for a peanut, the monkeys with
MPC lesions could now perform the
task.

Thus, MPC monkeys seem to have a

problem generating self-initiated
NO CUE CUED
actions, but have no problem when
instructed by an external cue. 27
 Neuroimaging results

• simple movements activate M1

• increasing the motor complexity of the task extends the area of

increased blood flow to include MPC

• when the subject mentally rehearses the complex movement, the area of
activation is restricted to MPC

• when the sequence is indicated by external cues, lateral premotor cortex

is activated instead

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Motor system organization: functional hierarchy

PREFRONTAL

Rules, strategies

MEDIAL PREMOTOR
LATERAL PREMOTOR
Internally generated
Externally cued actions
actions

PRIMARY MOTOR

Movements, muscles

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Brain-machine interface

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Brain-machine interface (BMI)

Matt Nagel was stabbed in the neck

while breaking up a fight at a fireworks
display. Paralyzed from the neck down,
he was implanted with an electrode
array in his motor cortex (similar to the
monkeys).

With this array he was able to control a

cursor on a computer screen (here he
uses it to draw a circle) and type. He
was also able to check email, surf the
internet and play some video games.

Future hopes are that such devices will

be able to control prosthetic and robotic
arms and control wheelchair. A further
aim is to make the implants wireless.
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 Summary

• we’ve seen how muscles are controlled to move joints in a co-

ordinated manner.

• we’ve looked at the sensory feedback mechanisms that are

available to help control the movement.

• we’ve looked at the evidence that the motor system is

organised hierarchically.

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