Action

Chapter 8:
• Page 327 – 366 ‘Learning and performing new skills’, except:
• page 339 ‘Where is it?’ Box
• page 350 - 351 ‘Patting your head while…’ Box
• page 360 - 361 ‘Contributions of the Basal Ganglia…’ Box

Victor A.F. Lamme

Brain and Cognition
Dept Psychology, University of Amsterdam
[email protected] 1
Mike, the headless chicken
1945 - 1947

Walked
Was fed with fluids and corn
Lived for 18 months
Earned $4500 per month (now 50K)
2
Hierarchical organization of motor pathways: from ‘concept’ to muscle
The final common pathway of all motor output:
the ventral horn Alpha motor neuron and the stretch reflex

Dorsolateral part
Distal muscles
Fine movements

Ventromedial part
Proximal muscles
Posture

4
Diseases that affect Alpha Motor Neurons

Lou Gherig’s disease or ALS: Poliomyelitis: viral infection

Amyotrophic Lateral Sclerosis that selectively attacks AMN’s

5
The Stretch Reflex: keeping posture
- Muscle Spindle senses stretching
- Activates Alpha Motor Neuron > muscle contracts
- Gamma motor neuron contracts muscle spindle during voluntary
movement so that they stay ‘short enough’ to sense stretching
when muscles are short. Input from Pons

6
Reciprocal
inhibition of
antagonistic
muscles

When extensor
(quadriceps)
contracts, flexor
relaxes

7
Crossed extensor reflex: as one limb flexes, the other extends!

This starts to look like walking…

8
Indeed the spinal cord can do walking ‘all on its own’
Decerebrated Cat (high level spinal cord cut) walks on treadmill, in different tempos
Central Pattern Generator generates locomotion

The Central Pattern Generator ‘creates’ locomotion movements

all on its own. The only thing that is required is a high level ‘gait’
command (or it can be set in motion by proprioceptive feedback)

9
Other fully automatic and highly
coordinated movement patterns
- Swallowing, Breathing, Orienting

10
Golgi Tendon Reflex: protecting from overload

11
Withdrawal Reflex: protecting from damage

12
Lower motor control
• Stretch reflex
• Golgi Tendon reflex
• Withdrawal reflex
• Antagonistic relaxation
• Crossed extension reflex
• Central Pattern Generators
• Many more…

• These do all the ‘work’

• Only thing that is needed is a
central ‘command’
• And some checks and balances…
13
Mike, the headless chicken
1945 - 1947

14
Lower motor control
• Stretch reflex
• Golgi Tendon reflex
• Withdrawal reflex
• Antagonistic relaxation
• Crossed extension reflex
• Central Pattern Generators
• Many more…

• These do all the ‘work’

• Only thing that is needed is a
central ‘command’
• And some checks and balances…
15
Pyramidal and Extrapyramidal systems project to spinal cord

Dorsolateral Ventromedial
Distal, fine Proximal, coarse
movements movements
Deliberate Posture
movement
Orienting
Tonus
Balance
Tonus
16
 Extrapyramidal systems
Rubrospinal tract
• upper motor neurons in red nuclei
• control muscle tone and distal limb muscles that
perform more precise movements
Tectospinal tract
• upper motor neurons in superior and inferior colliculi
• Receive visual (superior) and auditory (inferior) info
• Reflex-like Orienting Response: head, neck, upper limbs
move towards visual and auditory stimuli
Vestibulospinal tract
• Info from vestibulococlear nerve (VIII) (inner ear)
• Monitor position and movement of the head to
maintain Posture and Balance
Reticulospinal tract
• Reticular Formation, input from many pathways
• Controls many reflexes (excitability)
• State of arousal

17
Lower motor control
• Stretch reflex
• Golgi Tendon reflex
• Withdrawal reflex
• Antagonistic relaxation
• Crossed extension reflex
• Central Pattern Generators
• Extrapyramidal systems

• These do all the ‘work’

• Only thing that is needed is a
central ‘command’
• And some checks and balances…
18
Pyramidal system
• Voluntary (conscious) control of skeletal muscles:
• begins at upper motor neurons of primary motor
cortex and other cortical areas (SMA, PMC)
• axons descend into brain stem and spinal cord to
synapse on lower (alpha) motor neurons

Corticobulbar tract
• Towards cranial nerve nuclei that move eye, jaw,
face, and some muscles of neck and pharynx
(throat)

Corticospinal tract
• Corticospinal tracts visible along ventral surface
of medulla oblongata as pair of thick bands, the
pyramids
• Control of all non-facial somatic muscles
• Lateral CS tract crosses at pyramidal decussation
at high level
• Anterior CS tract cross to opposite side of spinal
cord at lower level in anterior white commissure
19
Figure 15–9
Damage to corticospinal tract
• Paralysis / paresis
• Spasticity / flaccidity:
• A pattern of weakness in the extensors (upper limbs) or flexors (lower limbs), known as 'pyramidal weakness'

• Changed reflexes: Babinski sign

20
‘Pyramidal Weakness’ of the right upper limb

Notice the weakness of the

upper limb (arm) extensors, so
that the wrist and elbow is in a
forced flexion position

Note: better therapy is Constraint Induced

Motor Therapy: forcing the weak parts to
work by tying up the strong half

https://www.youtube.com/watch?v=ddTiNFm-jvo&list=PLk73KWQbHBt0knk9do1p91oNZDTBj3qiM 21
Lower motor control
• Stretch reflex
• Golgi Tendon reflex
• Withdrawal reflex
• Antagonistic relaxation
• Crossed extension reflex
• Central Pattern Generators
• Extrapyramidal systems

• These do all the ‘work’

• Only thing that is needed is a
central ‘command’ (pyramidal)
• And some checks and balances…
22
 23
Introducing Spot, Boston Dynamics
(Checks) and Balances
- Perturbations
- Outside forces
- Uneven terrain

- Slopes
- Require different balance setting
between agonist and antagonist

24
25
Similar problems in humans and animals
when they lack a functional cerebellum

26
Cerebellum provides a
subcortical-cortical loop

-Fine tuning of movements

(lesions or alcohol result in ataxia)
-Timing of automated movement
sequences, motor memory
-Maybe also timing in general

27
 28
SEE: https://www.youtube.com/watch?v=xf1okjCwdOg
• Spinocerebellum
• Balance, Walking, Affected by alcohol use
• Neocerebellum
• Control of fine movements, Finger to nose test, Speech
• Vestibulocerebellum
• Coordination of eye movements with body movements, VOR (vestibulo-ocular-reflex)

29
Cerebellar Ataxia

Endpoint tremor, slurred speech

30
Lower motor control
• Stretch reflex
• Golgi Tendon reflex
• Withdrawal reflex
• Antagonistic relaxation
• Crossed extension reflex
• Central Pattern Generators
• Extrapyramidal systems

• These do all the ‘work’

• Only thing that is needed is a
central ‘command’ (pyramidal)
• And some checks and balances…
31
Basal Ganglia, Anatomy
• Striatum
• Caudate nucleus
• Putamen
• Globus Pallidum
• Externa D2
• Interna
• Substantia Nigra (Dopamine) D1
• Subthalamic nucleus

BG, function
• Direct pathway
• Indirect pathway
• Dopamine release by SNc
• D1, D2 receptors in striatum
 D2
D1

Indirect pathway inhibits GPe (which inhibits

GPi/SNr), so that GPi/ SNr activity increases > more
inhibition of thalamus, cortex > less movements

Direct pathway inhibits GPi / SNr so that

their inhibitory effect is diminished > more
activation of cortex > more movements

GPi / SNr act as inhibitor of thalamus, so

that in the end cortex and output to spinal
cord (i.e. movements) are suppressed
D2
D1
Huntington’s disease, Chorea, Hyperkinesia, Hereditary, Rare (1:10.000)

https://www.youtube.com/watch?v=OveGZdZ_sVs https://www.youtube.com/watch?v=dOufTOc3zKA&list=PLYPhva9ifXMzLEjIPkOhZuPJQOSw2pZwJ
35
Parkinson’s disease

Age of onset > 50

Common: 1:100
Drugs, Genes

Tremor of hands,
Rigidity, Bradykinesia
(slow and small
movements)

Medication:
L-dopa, stemcells, DBS
https://www.youtube.com/watch?v=Be2Enu65ZE8 36
Higher motor control
• M1: Primary Motor Cortex
• SMA: Supplementary Motor Area
• PMC: PreMotor Cortex
• PPC: Posterior Parietal Cortex

• Action planning
• Action selection
• Affordances
• Mirror Neurons
• Reference frames
37
 • M1: direct motor control
• PMC (with PPC): externally guided, stimulus
driven action (e.g. catching ball)
• SMA (with PFC): internally guided action
(e.g. selecting which object to pick up)

Cortical areas with

cortico-spinal fibers
M1, SMA, PMC, PPC, others
38
HEMIPLEGIA
• Half sided paralysis due to lesions of
upper motor neurons coming from M1

APRAXIA
• Loss of motor skill, not due to muscular,
upper (M1) or lower motor (Spinal Cord)
neuron deficit
• Lesions to SMA, PMC, PPC
• Ideomotor Apraxia: rough idea of
movement can be executed (SMA, PMC)
• Ideational Apraxia: No idea what to do,
uses wrong tools (PPC)
Ideomotor Apraxia

Notice how selective the

deficit is: many movements
can be roughly executed,
some totally not

40
https://www.youtube.com/watch?v=vTFdNk7JIoo
Higher motor control
• M1: Primary Motor Cortex
• SMA: Supplementary Motor Area
• PMC: PreMotor Cortex
• PPC: Posterior Parietal Cortex

• Action planning
• Action selection
• Affordances
• Mirror Neurons
• Reference frames
41
In M1 (and PMC)
neurons encode
movement direction

Individual motor
neurons encode vector
of movement, i.e. are
tuned for the direction
of limb movement

But tuning is fairly

broad…

42
Individual M1 neurons are broadly
tuned. Actual movement is encoded by
vector sum of population of M1 cells

43
Individual M1 neurons are broadly
tuned. Actual movement is encoded by
vector sum of population of M1 cells

44
The Affordance Competition Hypothesis

Sensory inputs create many

potential motor responses
(affordances). Depending on
needs and potential payoffs,
one of these has to be selected

45
Premotor Cortex:
encodes population
vectors of multiple
potential actions (go
to red, go to blue),
until color cue is
given to perform
one action and not
the other:

Note surge in
activity for ‘go to
red’, suppression of
activity for ‘go to
blue’ after cue
46
BUT: PMC also has many neurons for specific types of movements,
regardless of where and where to

Precision Grip
vs
Power Grip

Similar for
- Reach
- Hold
- Tear
- …
47
Another feature of PMC: Mirror Neurons: Neurons that encode an action,
yet also are activated by seeing (or hearing) the same action performed by others

48
 Doing Seeing

Mirror
Neurons

Filimon et al., Neuroimage 2007

Mirror neurons are widespread in motor cortex and parietal cortex.

Overlapping fMRI activation for acted and perceived movements
Contagious
Behavior
Posterior Parietal Cortex (PPC):
translating movement from
retinal (eye-centered) to hand-,
head-, or body-centered
reference frames

51
Higher motor control
• M1
Mind Reading?
• SMA
• PMC
• PPC

• Action planning
• Action selection
• Affordances
• Mirror Neurons
• Reference frames
52
Individual M1 neurons are broadly tuned. Actual movement is encoded
by vector sum of population of M1 cells

This is used to ‘read’ the planned movement from

the motor cortex, and use this in prosthetics

53
Training a computer algorithm (classifier) to ‘recognize’ specific patterns of neural
activity (combinations of activity increases and decreases of specific intentions to move)

54
Training the subject to turn his motor cortex on and off (biofeedback)

55
Training the classifier to recognize specific commands

56
Testing the interface with online movement

57
Decoding intentions from the brain: Prosthetics & Cyborgs

58
https://www.youtube.com/watch?v=zVMDHCnT-d4